Aphelenchoides fragariae




Rev 06/13/2022

Strawberry Foliar Nematode 
Strawberry Crimp Nematode
Classification Hosts
Morphology and Anatomy Life Cycle
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                Aphelenchoides fragariae (Ritzema Bos, 1891) Christie, 1942

Various common names include Strawberry crimp nematode, Strawberry spring dwarf nematode, Foliar nematode

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Morphology and Anatomy:

Aphelenchoides fragariae is 0.45-0.8mm long.



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

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Temperate and tropical regions, moist conditions. Widespread in the United States, Europe and Asia as a pest of many plants and seems well adapted to tropical and temperate climates (Scheck, 2021).

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Economic Importance:

C-rated pest in California Nematode Pest Rating System.


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Endoparasite in leaves, but also feeds ectoparasitically on leaf and flower buds in strawberry, for example.  

Nematode enters leaves through stomata or directly.

Aphelenchoides fragariae can swim rapidly; may be negatively geotropic.

Swiss scientist, J. Klingler studied entry of stomata by Aphelenchoides fragariae.  The nematodes aggregated around slits in plastic film even when there were no gases emerging, suggesting recognition of a tactile stimulus.  They had negative or indifferent reaction to oxygen, but strong positive reaction to carbon dioxide emerging from the slits.

That situation would occur at night - no photosynthesis, so CO2 from respiration would emerge from stomata.  Also at night, dew would provide a moisture film and facilitate nematode movement on the leaf surface.   Note, incidentally, that a positive attraction to CO2 is common in soil nematodes.

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More than 250 hosts, in 47 families, including fern, begonia, and strawberry.

The following are listed as hosts in various literature sources:

Aconitum lycoctonum L.  Wolfsbane Ficus benghalensis L.  Banyan Tree Polypodium brasiliense  
Aconitum sp. L.  Aconite Ficus buxifolia   Polypodium percussum Cav. 
Acrostichum flagelliferum Wall.  Swamp Fern Ficus cyathistipula   Polypodium punctatum Swartz.  Clinb Bird's Nest Fe
Acrostichum flagelliferum Wall.  Swamp Fern Ficus elastica Roxb. Decora Rubber-Plant Polypodium repens  
Actaea spicata L.  Baneberry Ficus elastica Roxb. decora variegata Rubber-Plant Polypodium scolopendria Burn. F.  Wart Fern
Adiantum aethiopicum   Maidenhair Fern Ficus elastica Roxb. decora Rubber-Plant Polypodium vulgare L.  European Polypodium
Adiantum capillus-veneris  Fissum Maidenhair Fern Ficus elastica Roxb.  Rubber-Plant Polystichum munitum K. Presl  Giant Holly Fern
Adiantum capillus-veneris   Maidenhair Fern Ficus lyrata Warb.  Fig Potentilla alpina  Hybrid
Adiantum peruvianum   Maidenhair Fern Ficus macrocarpa Hugel  Moreton Bay Fig Primula angustifolia Torr.  Primrose
Adiantum polyphyllum   Maidenhair Fern Ficus pumila L.  Climbing Fig Primula anisiaca   Primrose
Adiantum sp. L.  Maidenhair Fern Ficus religiosa L.  Peepul Primula auricula L.  Auricula
Adiantum tenerum   Maidenhair Fern Ficus rigo   Primula beesiana Forr.  Primrose
Adiantum trapeziforme   Maidenhair Fern Ficus rubiginosa Desf. ex Venten. variegata  Port Jackson Fig Primula denticulata Sm.  Primrose
Ageratum conyzoides L.  Floss Flower Ficus rubiginosa Desf. ex Venten.  Port Jackson Fig Primula denticulata Sm. Cashmiriana Primrose
Anchusa caespitosa Lam.  Bugloss Fragaria chiloensis Duchesne  Strawberry Primula japonica A. Gray  Primrose
Anemia collinia   Flowering Fern Fragaria chiloensis Duchesne  Strawberry Primula luteola Rupr.  Primrose
Anemia phyllitidis Swartz.  Flowering Fern Fragaria chiloensis Duchesne  Strawberry Primula malacoides Franch.  Fairy Primrose
Anemone alpina L.  Windflower Fragaria chiloensis Duchesne Chandler Strawberry Primula obconica Hance.  German Primrose
Anemone angulosa Lam  Windflower Fragaria X ananassa   Strawberry Primula pulverulenta Duthie  Primrose
Anemone coronaria L.  Poppy Anemone Fuchsia sp. L.  Lady's Eardrops Primula rosea Royle  Primrose
Anemone flaccida Friedr. Schmidt.  Poppy Anemone Gardenia jasminoides Ellis.  Gardenia Primula sp. L.  Primrose
Anemone halleri All.  Windflower Geranium cinereum R. Knuth subcaulescens  Cranesbill Primula sp. L. Jewel Primrose
Anemone hupehensis Bowles and Stearn  Japanese Anemone Geranium macrorrhizum L.  Cranesbill Primula wilsoni S.T. Dun  Primrose
Anemone hupehensis Bowles and Stearn japonica  Japanese Anemone Gerbera jamesonii H. Bolus ex Hook. f.  Transvaal Daisy Primula X polyantha Hort.  Polyanthus
Anemone nemorosa L.  European Wood Anemon Glechoma hederacea L.  Ground Ivy Pteris altissima  Repandula ?
Anemone ranunculoides L.  Windflower Gomphrena globosa L.  Globe-Amaranth Pteris altissima  
Anemone sylvestris L.  Windflower Gunnera chilensis Lam.  Pteris biaurita  Allosora
Anemone vitifolia Buch.-Ham.  Windflower Gymnostachyum ceylanicum Arn. & Nees  Pteris biaurita  Repandula
Arisaema amurense Maxim.  Gypsophila cerastioides D. Don  Pteris cretica L. Ouvardii
Arisaema ringens Schott  Helichrysum sp.   Pteris cretica L. Nobilis
Aruncus dioicus Fern.  Goatsbeard Helleborus abchasicus A. Braun.  Hellebore Pteris cretica L. Crispata
Aspidium barteri   Helleborus antiquorum A. Braun.  Hellebore Pteris cretica L. Major
Aspidium capense   Helleborus cyclophyllus Boiss.  Hellebore Pteris cretica L. 
Asplenium adiantum-nigrum   Helleborus foetidus L.  Hellebore Pteris cretica L. Wimsetti
Asplenium bulbiferum G. Forst.  Mother Spleenwort Helleborus intermedius Host  Hellebore Pteris cretica L. Wilsoni
Asplenium decussatum   Helleborus niger L.  Hellebore Pteris cretica L. 
Asplenium dimorphum   Helleborus olympicus Lindl.  Hellebore Pteris cretica L. Albolineata
Asplenium diversifolium   Helleborus orientalis Lam.  Lenten Rose Pteris cretica L. Gaultheri ?
Asplenium nidus L.  Bird's Nest Fern Hemigraphis alternata T. Anderson  Red Ivy Pteris cretica L. Rivertoniana
Asplenium nidus L.  Bird's Nest Fern Hepatica americana/nobilis Ker-Gawl./Gars.  Liverleaf Pteris dentata Forssk.  Toothed Brake
Asplenium nidus L.  Bird's Nest Fern Hepatica americana/nobilis Ker-Gawl./Gars. Marmorata Liverleaf Pteris denticulata  
Asplenium sp. L.  Spleenwort Heuchera sanguinea Engelm  Coralbells Pteris droogmansiana  
Asplenium sp. L.  Spleenwort Heuchera sp. L.  Alumroot Pteris longifolia  Mariesii
Asplenium trichomanes L.  Maidenhair Spleenwor Horminum pyrenaicum L.  Dragon-Mouth Pteris longifolia  
Aster alpinus L. altaicus  Michaelmas Daisy Huechera sp. L.  Beach Heather Pteris multifida Poir. 
Astrantia biebersteinii Trautv.  Masterwort Hydrangea macrophylla Ser. macrophylla  French Hydrangea Pteris pellucida  Drinkwateri ?
Astrantia carniolica Vul'f.  Masterwort Hydrangea sp.   Hydrangea Pteris podophylla  
Astrantia major L.  Masterwort Hydrophyllum canadense L.  Waterleaf Pteris quadriaurita  
Begonia cucullata Willd. hookeri Vernon Begonia Impatiens balsamina L.  Garden Balsam Pteris sp. L.  Brake
Begonia evansiana X B. socotrana  Schwabenland Red Begonia Iris sp. L.  Flag Pteris tremula  
Begonia foliosa L. B. Sm. & Schub. miniata  Fuchsia Begonia Iris sp. L.  Flag Pteris umbrosa  
Begonia rex Putz.  King Begonia Kalanchoe integra O. Kuntze  Flame Kalanchoe Pulmonaria officinalis L.  Jerusalem Sage
Begonia sp.   Begonia Knautia arvensis T. Coult.  Blue Buttons Pyrus sp.  
Bergenia crassifolia Fritsch  Siberian Tea Knautia macedonica   Ranunculus alpestris  
Bergenia delavayi Engl.  Lantana sp. L.  Shrub Verbena Ranunculus montanus  
Bergenia pacifica Kom.  Ligularia dentata Hara  Ranunculus sp. L.  Buttercup
Blechnum brasiliense Desv.  Ligustrum vulgare L.  Common Privet Rodgersia podophylla A. Gray 
Blechnum cartilagineum   Lilium henryi Bak.  Rodgersia purdomii  
Blechnum fraxineum   Lilium humboldtii Roezl & Leichtl.  Humboldt Lily Rodgersia sambucifolia  
Blechnum gibbum Mett.  Lilium humboldtii X L. pardalinum   Rudbeckia fulgida Perdue speciosa 
Blechnum gibbum Mett.  Lilium longiflorum Thunb. eximium  Trumpet Lily Rumex alpinus  
Blechnum moorei   Lilium longiflorum Thunb. eximium  Trumpet Lily Ruscus hypophyllum L. 
Blechnum sp.   Lilium pardalinum  sunset  Saintpaulia ionantha Wendl.  African-Violet
Bletilla striata Rchb.f.  Lilium pardalinum   Saintpaulia ionantha Wendl.  African-Violet
Bletilla striata Rchb.f.  Lilium philippinense Bak. formosanum grandiflo  Saintpaulia sp. H. Wendl.  African Violet
Bouvardia sp. Salisb.  Lilium pumilum Delile  Coral Lily Salvia farinacea  
Boykinia aconitifolia Nutt.  Lilium regale E. H. Wils.  Regal Lily Salvia sp. L.  Sage
Brassica oleracea Mill. capitata  Cabbage Lilium sargentiae E. H. Wils.  Salvia splendens Sello  Scarlet Sage
Brunnera macrophylla I. M. Johnst.  Siberian Bugloss Lilium sp. L.  Lily Saxifraga ligulata  
Calamintha nepeta Savi  Calamint Lilium speciosum Thunb.  Showy Lily Saxifraga rosacea Moench 
Calceolaria sp. L.  Slipperwort Lilium X hollandicum Bergmans ex Woodcock & Stearn  Candlestick Saxifraga rotundifolia  
Ceterach officinarum Willdenow  Lilium X sulphurgale   Saxifraga stolonifera Meerb.  Strawberryn Geranium
Chrysanthemum hortorum Bailey  Limonium sinuatum Mill.  Scabiosa caucasica  Clive Greaves
Chrysanthemum indicum L.  Lobelia erinus L.  Edging Lobia Scabiosa columbaria  
Chrysanthemum leucanthemum L.  Marguerite Lycoris radiata Herb.  Scabiosa lucida Vill.  Pincushion Flower
Chrysanthemum sp. L.  Chrysanthemum Lygodium circinatum Swartz.  Malay Climbing Fern Scabiosa silenifolia  
Clematis alpina Mill.  Virgin's Bower Lygodium sp. Swartz.  Climbing Fern Scilla sp.  
Clematis heracleifolia DC. davidiana  Virgin's Bower Lygodium volubile Swartz.  Climbing Fern Scrophularia vernalis  
Clematis vitalba L.  Virgin's Bower Machaeranthera bigelovii Greene  Senecio nemorensis  
Colchicum sp. L.  Mentha spicata L.  Spearmint Shortia sp.  
Coleus sp. Lour.  Flame Nettle Microlepia platyphylla John Smith  Silene elisabethae Jan 
Coniogramme japonica   Narcissus pseudonarcissus L.  Daffodil Sinningia hybrida  
Convallaria majalis L.  Lily of the Valley Narcissus sp. L.  Narcissus Sinningia sp.  
Crassula sp. L.  Nephrodium polymorphum   Sinningia speciosa Benth. & Hook.  Gloxinia
Crossandra undulaefolia R. A. Salisbury  Firecracker Flower Nephrolepis biserrata   Stachys alopecuros  
Cyclamen persicum Mill.  Florist's Cyclamen Nephrolepis exaltata Schott.  Boston Fern Stachys longifolia  
Cyclamen sp. L.  Cyclamen Nephrolepis exaltata Schott.  Boston Fern Stachys officinalis  
Cyperus alternifolius L.  Umbrella Plant Nerine sp. Herbert  Stachys palustris  
Cypripedium reginae Walt.  Showy Lady's Slipper Nicotiana sp. L.  Stenochlaena tenuifolia T. Moore. 
Cypripedium sp. L.  Lady's Slipper Nicotiana virginica   Stenoglottis longifolia  
Cystopteris bulbifera   Odontoglossum sp. Kunth.  Trop America Strobilanthes dyeranus Mast.  Persian Shield
Cystopteris fragilis Bernhardi  Brittle Fern Omphalodes verna Moench.  Creeping Forgetmenot Symphytum asperum  
Dahlia pinnata Cav.  Dahlia Osmunda regalis L.  Royal Fern Synthyris sp. Benth. 
Dahlia sp.   Osmunda regalis L.  Royal Fern Tellima grandiflora Douglas ex Lindley 
Dahlia sp.   Paeonia lactiflora Pall.  Chinese Peony Teucrium arduini  
Davallia canariensis Sm.  Deer's Foot Fern Paeonia lutea   Teucrium chamaedrys  
Davallia denticulata   Paeonia officinalis L.  Peony Thelypteris decursive-pinnata Ching 
Davallia solida   Paeonia sp. L.  Peony Tiarella cordifolia L.  False Mitterwort
Davallia sp.   Paeonia sp. L.  Peony Tradescantia virginiana  
Digitalis purpurea L.  Common Foxglove Paeonia suffruticosa Andr.  Tree Peony Trifolium repens L.  White Clover
Diplazium australe Wakef.  Paeonia wittmanniana   Trillium grandiflorum Salisb. 
Diplazium expansum   Paphiopedilum sp. Pfitz.  Lady Slipper Triosteum fargesii Franch. 
Diplazium shepherdii   Paphiopedilum sp. Pfitz.  Lady Slipper Ulmus sp. L.  Elm
Diplazium sylvaticum   Pelargonium sp. L'Her.  Geranium Vanda sp. v. X  Miss Joaquim
Doronicum cordatum Schultz-Bip.  Leopard's Bane Peltiphyllum peltatum Engl.  Umbrella Plant Verbena tenera  
Doronicum plantagineum L.  Leopard's Bane Peltiphyllum peltatum Engl.  Umbrella Plant Viola odorata L.  Sweet Violet
Doronicum sp. L.  Leopard's Bane Penstemon barbatus Nutt.  Beard Tongue Viola sp. L.  Violet
Dryopteris austriaca Fern. americana  Wood Fern Penstemon gentianoides   Viola sp. L.  Violet
Dryopteris borrei   Wood Fern Penstemon hirsutus   Viola ugliginosa  
Dryopteris filix-mas Schott  Male Fern Penstemon neomexicanus   Woodwardia paradoxa  
Dryopteris hirtipes O. Kuntze.  Wood Fern Pentas lanceolata Deflers  Star Cluster Woodwardia sp. Sm.  Chain Fern
Dryopteris remota   Wood Fern Peperomia magnoliifolia   Radiator Plant Xanthium canadense  
Dryopteris sp. Adans.  Wood Fern Phlox paniculata   Zebrina pendula Schnizl.  Wandering Jew
Dryopteris X boottii Underw.  Boott's Wood Fern Phyllitis scolopendrium Newm.  Hart's Tongue Fern Zinnia elegans Jacq.  Zinnia
Endymion hispanicus Chouard  Spanish Bluebell Pinus palustris Mill.  Longleaf Pine Zinnia sp. L.  Zinnia
Epipactis palustris Crantz.  Helloborine Pityrogramma calomelanos Link  Gold Fern Zoysia sp.  
Epiphyllum sp. Haw.  Orchid Cactus Pityrogramma calomelanos Link  Gold Fern
Epipremnum aureum Bunt.  Pothos Polypodiaceae sp.   Polypody Family
Episcia lilacina Hanst.f.  Carpet Plant Polypodiaceae sp.   Polypody Family
Eryngium alpinum L.  Eryngo Polypodium aureum L. Mandaianum Rabbit's Foot Fern
Ficus altissima Blume.  Council Tree Polypodium aureum L. Areolatum Rabbit's Foot Fern
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





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Nematode feeding causes blotches and necrotic lesions between veins on fern leaves which start as water-soaked spots and then turn brown. 

In strawberries, A. fragariae causes malformed leaves with crinkled edges. (G. Caubel)


A. fragariae symptoms on Begonia (M. Ritter)

Nematode injury to leaves: Drawing by Charles S. Papp, CDFA

Disease symptoms on strawberry are called spring dwarf, spring crimp and red plant.

Endoparasitic activity of A. fragariae causes shoot malformations including twisting and puckering of leaves, discolored areas with a hard and rough surface, undersized leaves with crinkled edges, reddening of petioles, tight aggregation of crowns, shortening of internodes on runners, reduced flower trusses with only one or two flowers or flowers aborted, and death of the crown bud (Maas, 1998; Ploeg and Westerdahl, 2018).

Ectoparasitic feeding on folded crown and runner buds causes small dry, brown areas that are seen on expanded leaves usually near the mid-rib. Occasionally the nematodes are found in strawberry fruit pulp (Tacconi, 1972).

On flowering ornamentals such as violets, anemone and begonia, the feeding areas of foliar nematode appear as irregular, water-soaked patches later turning brown, violet, or black. The affected areas are limited by the leaf veins.

The nematode causes die-back disease of lilies; leaves, flower buds and fruits turn brown and die (Daughtrey et al., 1995). Ferns develop water-soaked, often chevron-like stripes on fronds as movement of the nematodes seems to be delimited by vein. The stripes turn brown in summer when the fern is the driest (Sandeno and Jensen, 1962).

The nematode is easily spread to new hosts when in the anhydrobiotic condition, or by movement of infected but asymptomatic planting stock.

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Roguing plants, burning infested material, starting/replacing with healthy stock, general sanitation. Nematode-free planting stock may be based on inspection by regulatory agencies. For example,   foundation, registered, and certified nursery stock may be inspected and tested for viral, fungal, and bacterial pathogens, nematodes, and varietal purity.


Hot water dips can be used on strawberry plants (15 min. at 47 C.), but cultivars and requirements change - current experiments are being conducted by Westerdahl.  

Plants must not be damaged, and yield must not be affected negatively.  

Hot water treatment may be used for other crops, also.  

Chemical control can be used; e.g., Parathion as a foliar spray. 

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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Bohmer B, 1981. The harmful effect of Aphelenchoides fragariae and A. ritzemabosi on Fragaria ananassa. Gesunde Pflanzen, 33(5):113-117

CABI Crop Production Compendium 2021. Apehelenchoides fragariae. https://www.cabi.org/cpc/datasheet/6381 Accessed 9/29/21

Chitambar, J. J., Westerdahl, B. B., and Subbotin, S. A. 2018. Plant Parasitic Nematodes in California Agriculture. In Subbotin, S., Chitambar J., (eds) Plant Parasitic Nematodes in Sustainable Agriculture of North America. Sustainability in Plant and Crop Protection. Springer, Cham.

Daughtrey, M. L., Wick, R. L., and Peterson, J. L. 1995. Compendium of flowering potted plant diseases. American Phytopathological society, St. Paul, MN

EPPO Global Database. 2021. Aphelenchoides fragariae https://gd.eppo.int/taxon/APLOFR. Accessed 9/29/2021/21

Goodey, J. B., M. T. Franklin, and D. J. Hooper. 1965. T. Goodey's: The Nematode Parasites of Plants Catalogued Under Their Hosts. Commonwealth Agricultural Bureaux, Farnham Royal, Bucks, England. Third Edition

Ibrahim, S. K., R. N. Perry and D. J. Hooper. 1994. Use of esterase and protein patterns to differentiate two new species of Aphelenchoides on rice from other species of Aphelenchoides and from

Kohl, L. M. 2011. Foliar nematodes: A summary of biology and control with a compilation of host range. Online. Plant Health Progress doi:10.1094/PHP-2011-1129-01-RV

Maas, J.L., Ed. 1998. Compendium of Strawberry diseases 2nd Edition. APS Press. Nemaplex UC Davis Nemabase 2010. Aphelencoides fragariae. Accessed 9/29/21

Maeseneer, J. D. 1964. Leaf-browning of Ficus spp., new host plants of Aphelenchoides fragariae (Ritzema Bos). Nematologica 10:403-408.  

Mor, M., and Y. Spiegel. 1993.  Ruscus hypophyllum: a new host for Aphelenchoides fragariae. Journal of Nematology 25:312-313.  

Ploeg, A. and Westerdahl, B. B. 2018. UC IPM pest Management Guidelines: Strawberry Nematodes. UC ANR Publication 3468

Riedel, R. M. and P. O. Larsen. 1974. Interrelationship of Aphelenchoides fragariae and Xanthomonas begoniae on Rieger begonia. Journal of Nematology 6:215-216.

Sandeno, J.L., and Jensen, H. J., 1962. A foliar nematode disease of western sword-fern, Polystichum munitum. Plant Disease Reporter, 46(10):699-701.

Scheck, H.J. 2021. California Pest Rating Proposal for Aphelenchoides fragariae (Ritzema - Bos, 1891) Christie, 1932 Strawberry crimp nematode, Strawberry spring dwarf nematode, Foliar nematode CDFA, Sacramento.

Siddiqui, I. A., S. A. Sher and A. M. French. 1973. Distribution of Plant Parasitic Nematodes in California. State of California Department of Food and Agriculture, Division of Plant Industry. 324p.

Tacconi R, 1972. Infestations of Aphelenchoides fragariae, Aphelenchoides ritzemabosi and Ditylenchus dipsaci on strawberry in some Italian provinces. Redia, 53:313-319

USDA Phytosanitary Certificate Issuance and Tracking System, Phytosanitary Export Database (PExD) Harmful Organisms Database Report. Aphelenchoides fragariae. Accessed 9/29/2021


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Copyright © 1999 by Howard Ferris.
Revised: June 13, 2022.