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Page 1: Nematode Threats in the NAQS Region

i

Nematode threats in the NAQS region © CSIRO

Nematode Threats in the NAQS Region

(Photo by Tesfa Mekete)

Mike Hodda, Natalie Banks & Sunil Singh

October, 2012

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Nematode threats in the NAQS region © CSIRO

CSIRO advises that the information contained in this publication comprises general

statements based on scientific research up to the date of publication. The reader is

advised and needs to be aware that such information may be incomplete or unable to

be used in any specific situation. No reliance or actions must be made on the

information without seeking prior expert professional, scientific and technical advice.

To the extent permitted by law, CSIRO (including its employees and consultants)

excludes all liability to any person for any consequences, including but not limited to all

losses, damages, costs, expenses and any other compensation, arising directly or

indirectly from using this publication (in part or in whole) and any information or

material contained in it.

If you publish excerpts from the reports, then those excerpts must be an accurate

representation of the test results. You must note on the excerpts that they are from a

larger report held by you and where the report can be inspected.

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Table of contents

Table of contents .................................................................................................ii

List of Figures ..................................................................................................... vi

List of Tables .....................................................................................................vii

Contributors .................................................................................................... viii

Authors ............................................................................................................................... viii

Data and Advice ................................................................................................................. viii

Abbreviations and Acronyms ........................................................................... ix

Summary .............................................................................................................. 1

Key findings ........................................................................................................................... 1

Nematode species identified as threats................................................................................... 2

Introduction ....................................................................................................... 12

Background .......................................................................................................................... 12

Scope .................................................................................................................................... 12

Methodology ...................................................................................................... 14

Data collection...................................................................................................................... 14

Data Filters ........................................................................................................................... 15

Additional criteria for assessing threat posed ...................................................................... 15

Results ................................................................................................................ 17

Nematodes ............................................................................................................................ 17

Geography ............................................................................................................................ 19

Host Plants............................................................................................................................ 20

Pathways............................................................................................................................... 22

Limitations of review ........................................................................................ 26

Assessing threat .................................................................................................................... 26

Verification of material ........................................................................................................ 26

Uncertainty of identifications ............................................................................................... 26

Limited survey data .............................................................................................................. 26

Uncertainty over damage caused.......................................................................................... 27

Climatic suitability for establishment .................................................................................. 27

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Host races or pathotypes ...................................................................................................... 27

Impacts outside the NAQS region ........................................................................................ 28

References .......................................................................................................... 30

Appendix A—List of nematode species evaluated ......................................... 36

Appendix B—List of all host plants considered ............................................. 38

Appendix C—Detailed nematode profiles ...................................................... 40

Ditylenchus angustus............................................................................................................ 41

Ditylenchus destructor ......................................................................................................... 42

Globodera pallida ................................................................................................................ 43

Globodera rostochiensis ...................................................................................................... 44

Heterodera glycines ............................................................................................................. 45

Heterodera zeae ................................................................................................................... 46

Hirschmanniella miticausa .................................................................................................. 47

Hirschmanniella oryzae ....................................................................................................... 48

Longidorus elongatus ........................................................................................................... 49

Meloidogyne enterolobii ...................................................................................................... 50

Meloidogyne exigua ............................................................................................................. 51

Meloidogyne graminicola .................................................................................................... 52

Meloidogyne naasi ............................................................................................................... 53

Meloidogyne thailandica ...................................................................................................... 54

Radopholus arabocoffeae ..................................................................................................... 55

Radopholus bridgei .............................................................................................................. 56

Radopholus citri ................................................................................................................... 57

Radopholus similis ............................................................................................................... 58

Xiphinema americanum s.l. .................................................................................................. 60

Appendix D—Nematodes recorded on host crops considered ..................... 62

Abelmoschus spp. (aibika, okra etc.) ................................................................................ 63

Allium cepa (onion) .......................................................................................................... 63

Ananas comosus (pineapple) ............................................................................................ 63

Annona reticulata (custard apple) .................................................................................... 63

Bambusa spp. (bamboo) ................................................................................................... 63

Bougainvillea spp. ............................................................................................................ 63

Brassica spp. (cabbage, broccoli etc.) ............................................................................. 64

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Calopogonium caeruleum ................................................................................................ 64

Calopogonium mucunoides .............................................................................................. 64

Camellia sinensis (tea) ..................................................................................................... 64

Capsicum spp. (chilli, capsicum etc.) ............................................................................... 64

Carica papaya (pawpaw) ................................................................................................ 65

Celosia argentea (cock's comb) ........................................................................................ 65

Citrullus lanatus (watermelon)......................................................................................... 65

Citrus spp.......................................................................................................................... 65

Cocos nucifera (coconut) ................................................................................................. 66

Coffea sp. (coffee) ............................................................................................................. 66

Colocasia esculenta (taro) ............................................................................................... 66

Coriandrum sativum (coriander) ..................................................................................... 66

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.) .................................................... 66

Dimocarpus longan (longan) ........................................................................................... 67

Dioscorea spp. .................................................................................................................. 67

Dracaena spp.................................................................................................................... 67

Durio spp. (durian) ........................................................................................................... 67

Elettaria cardamomum (cardamom) ............................................................................... 67

Ficus spp. (figs) ................................................................................................................ 67

Gardenia sp. ..................................................................................................................... 67

Glycine max (soybean) ..................................................................................................... 68

Heliconia spp. ................................................................................................................... 68

Hibiscus spp...................................................................................................................... 68

Ipomoea spp...................................................................................................................... 68

Lactuca spp. ...................................................................................................................... 69

Litchi chinensis (lychee) ................................................................................................... 69

Mangifera spp. (mango) ................................................................................................... 69

Manihot esculenta (cassava) ............................................................................................ 69

Manilkara zapota (sapodilla) ........................................................................................... 69

Momordica charantia (bitter melon) ................................................................................ 69

Morus spp. (mulberry) ...................................................................................................... 70

Musa spp. (banana & plantain)........................................................................................ 70

Nephelium lappaceum (rambutan) ................................................................................... 70

Nicotiana tabacum (tobacco) ........................................................................................... 71

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Oryza spp. (rice) ............................................................................................................... 71

Passiflora spp. (passion fruit) .......................................................................................... 72

Persea americana (avocado) ............................................................................................ 72

Petroselinum crispum (parsley) ....................................................................................... 72

Phaseolus vulgaris (haricot bean).................................................................................... 72

Pinus spp. (pine) ............................................................................................................... 73

Piper spp. (pepper) ........................................................................................................... 73

Psidium guajava (guava) .................................................................................................. 73

Psophocarpus tetragonolobus (winged bean) .................................................................. 73

Rhododendron spp. ........................................................................................................... 73

Rosa spp. (rose) ................................................................................................................ 74

Saccharum officinarum (sugarcane) ................................................................................ 75

Saccharum officinarum (sugarcane) ................................................................................ 75

Solanum spp. (potato, aubergine, tomato etc.) ................................................................. 76

Sorghum spp. (sorghum) .................................................................................................. 77

Triticum spp. (wheat)........................................................................................................ 78

Triticum spp. (wheat)........................................................................................................ 78

Vigna spp. (bean).............................................................................................................. 78

Vitis spp. (grape) .............................................................................................................. 79

Xanthosoma spp................................................................................................................ 80

Zea mays (maize) .............................................................................................................. 81

Zea mays (maize) .............................................................................................................. 81

Zingiber officinale (ginger) .............................................................................................. 82

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List of Figures

Figure 1. Process for identification of nematode threats. ...................................................... 16

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List of Tables

Table 1. Greatest nematode threats. ......................................................................................... 3

Table 2. Probable nematode threats ......................................................................................... 5

Table 3. Possible nematode threats in NAQS countries. ....................................................... 10

Table 4. Low nematode threats. ............................................................................................. 11

Table 5. Number of the crops investigated which are hosts for nematode species identified

as threats................................................................................................................................... 18

Table 6. Most frequent crop plant hosts for nematodes in the NAQS region and (number of

nematode species recorded). .................................................................................................... 21

Table 7. Major crop plant hosts outside the NAQS region for nematodes identifed as threats.

.................................................................................................................................................. 21

Table 8. Pathways for greatest nematode threats. Top row: coloured boxes= NAQS

pathways and clear boxes= non-NAQS pathways. .................................................................. 23

Table 9. Pathways for probable nematode threats. Top row: coloured boxes= NAQS

pathways and clear boxes= non-NAQS pathways. .................................................................. 24

Table 10. Pathways for possible nematode threats. Top row: coloured boxes= NAQS

pathways and clear boxes= non-NAQS pathways. .................................................................. 25

Table 11. Genera with unidentified species from NAQS or ASEAN countries. ................... 29

Table 12. List of all nematode species considered as part of this review. ............................. 37

Table 13. List of all host plants considered in this review. .................................................... 39

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Contributors

Authors

Mike Hodda Research Scientist, CSIRO Ecosystem Sciences, Australia

Natalie Banks Technical Officer & Graduate Student, CSIRO Ecosystem

Sciences

Sunil Singh Graduate Student, CSIRO Ecosystem Sciences, Australia

Data and Advice

Richard Davis Plant Pathologist, Northern Australia Quarantine Strategy,

Department of Agriculture, Fisheries and Forestry, Australia

Lynne Jones Plant Pathologist, Northern Australia Quarantine Strategy,

Department of Agriculture, Fisheries and Forestry, Australia

Stephen Mckenna Botanist, Northern Australia Quarantine Strategy, Department

of Agriculture, Fisheries and Forestry, Australia

James Walker Scientific Manager, Northern Australia Quarantine Strategy,

Department of Agriculture, Fisheries and Forestry, Australia

Barbara Waterhouse Senior Botanist, Northern Australia Quarantine Strategy,

Department of Agriculture, Fisheries and Forestry, Australia

Debbie Jennings Technical Officer, Australian National Insect Collection

Takaniko Ruabete Plant Pathologist (Nematodes), Secretariat of the Pacific

Community Suva, Fiji Islands

Dr. Mulawarman Associate Professor, Sriwijaya University, Indonesia

Ariene Garcia-Castillo Scientist, Agri-Food & Veterinary Authority, Singapore

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Abbreviations and Acronyms

ASEAN countries.................. Association of Southeast Asian Nations: Myanmar, Cambodia,

Lao PDR, Vietnam, Thailand, Brunei Darussalam, Malaysia,

Singapore, Philippines, but herein excluding Indonesia because

it is included in the NAQS countries.

NAQS countries .................... Indonesia, Timor Leste, & Papua New Guinea. For the

purposes of this report the Solomon Islands are included

because of data availability and proximity to the NAQs region.

CABI ..................................... Centre for Agricultural Bioscience International

EPPO ..................................... European and Mediterranean Plant Protection Organisation

IPPC ...................................... International Plant Protection Convention

NAQS .................................... Northern Australia Quarantine Strategy

NEMABASE......................... University of California Davis Department of Nematology

Nematode Hosts Database

PNG....................................... Papua New Guinea

SPC ....................................... Secretariat of the Pacific Community

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Summary

Nematode pests are of considerable economic importance but their status in the countries to

the north of Australia is poorly known. In June, 2012, the Northern Australia Quarantine

Strategy group (NAQS) of the Department of Agriculture, Fisheries and Forestry (DAFF)

commissioned this review of available information on the nematodes from the countries

bordering northern Australia to identify any agriculturally important nematodes present in the

region which may represent threats to Australian biosecurity.

This review examines the following factors relevant to the risk of entry into Australia

(especially into Northern Australia) by plant-parasitic nematodes:

Presence in the NAQS region of interest as well as in the ASEAN region more

broadly, together with absence from Australia;

Commonly grown host plants in above regions;

Pathways used to move; and

Impact (defined here as causing documented economic damage).

The review is based on examination of formally- and informally-published, peer-reviewed

and unreviewed literature, public databases from reputable centres of nematology expertise,

opinions and data provided by national experts, as well as unpublished datasets.

Key findings

Records of nematodes from NAQS region countries are sparse, and certainly underrepresent

the nematodes actually present. Records from the ASEAN countries are more extensive, but

still omit species which may be of concern. Data on pathways and hosts in both regions are

also sparse.

Nematode threats can be categorized into 4 groups based on currently available data.

8 species which fulfilled all criteria to be risks fully and thus were considered the

greatest threats.

11 species which fulfilled all criteria except one, and hence were considered probable

threats, because they:

o have not been recorded in the NAQS region countries, but are present in the

ASEAN region immediately to the north; or

o have been recorded from Australia, but have very restricted distributions or

only some host races present within Australia; or

o have not been proven pathogenic to plants.

6 species which could fulfil all criteria if races currently recognised as differing in

host range or pathogenicity are distributed differently in Australia and the NAQS

countries, but where there have been no investigations of whether this is the case.

Hence these species were considered possible threats.

75 species which are known from the ASEAN region countries but fulfilled only some

criteria and hence were considered low threats.

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The crops most likely to host nematodes which were identified as threats were Solanum spp.,

Zea mays, Oryza spp. and Cucurbitaceae spp. (Table 5, p18). Other crops hosting many of

the nematode threats were Capsicum spp., Citrus spp. and Sorghum spp.. Other crops

hosting many of the nematodes identified as threats were Allium cepa, Coffea spp. and

Saccharum officinarum.

Many of the greatest threats had relatively narrow host ranges among the crops investigated,

which may allow targeting for surveillance. Although the host ranges may have been narrow

botanically, most of the hosts were widely distributed and common, meaning that potential

spread and impact are great. Many of the other nematode threats identified had a wide range

of crops as hosts, as well as widely distributed and common hosts.

Significantly, many of the nematodes identified as the greatest or probable threats had host

ranges including Triticum aestivum (wheat) and Glycine max (soy bean). These species are

commercial crops grown outside the NAQS region of Australia, and so are not considered in

detail in this review, but the fact that they are damaged by some of the nematode threats

identified increases the potential negative impact of introduction of the nematodes concerned.

That is, their impact may be greatest in parts of Australia outside the NAQS region.

The pathways most likely to carry damaging nematodes in the region were soil and plants or

plant parts.

Nematode species identified as threats

Greatest proven threats:

Ditylenchus angustus, Globodera pallida, Heterodera glycines, Heterodera zeae,

Hirschmanniella miticausa, Hirshmanniella oryzae, Meloidogyne graminicola, and

Radopholus citri.

Probable threats:

Ditylenchus destructor, Meloidogyne enterolobii, Meloidogyne naasi, Meloidogyne

thailandica, and Radopholus arabocoffeae (recorded in ASEAN countries but not

NAQS countries);

Globodera rostochiensis, Longidorus elongatus, Meloidogyne exigua, Radopholus

similis, and Xiphinema americanum sensu lato (restricted distribution or limited

number of host races in Australia).

Radopholus bridgei (pathogenicity not demonstrated).

Possible threats:

Meloidogyne arenaria, Meloidogyne hapla, Meloidogyne incognita, Meloidogyne

javanica, Pratylenchus coffeae, and Pratylenchus zeae (recorded from NAQS

countries and Australia, but have races or pathotypes differing in hosts and

pathogenicity which are of biosecurity significance although these have not been

investigated and hence their distributions in the NAQS countries and Australia are

unknown).

Low threats:

73 species as listed in Table 4.

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Table 1. Greatest nematode threats.

Species Country Common crop hosts Pathways

Ditylenchus angustus Indonesia Oryza spp. (rice)

Plants or Plant Parts

Corms, Rhizomes & Tubers

Seedlings & Nursery Stock

Flower & Ornamental Plants

Bulbs

Seeds

Globodera pallida Indonesia Solanum spp. (potato, aubergine, tomato etc.)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Heterodera glycines Indonesia

Beta vulgaris (beetroot)

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.)

Glycine max (soybean)

Vigna spp. (bean)

Zea mays (maize)

Soil

Roots and Rootstock

Seedlings & Nursery Stock

Plants or Plant Parts

Heterodera zeae Indonesia

Bambusa spp. (bamboo)

Capsicum spp. (chilli, capsicum etc.)

Citrus spp.

Oryza spp. (rice)

Saccharum officinarum (sugarcane)

Solanum spp. (potato, aubergine, tomato etc.)

Sorghum spp. (sorghum)

Triticum spp. (wheat)

Zea mays (maize)

Soil

Roots and Rootstock

Seedlings & Nursery Stock

Plants or Plant Parts

Hirschmanniella miticausa PNG, Solomons Colocasia (taro)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

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Hirschmanniella oryzae

Indonesia, Malaysia,

Myanmar, Philippines,

Thailand, Vietnam

Abelmoschus spp. (aibika, okra etc.)

Oryza spp. (rice)

Saccharum officinarum (sugarcane)

Solanum spp. (potato, aubergine, tomato etc.)

Sorghum spp. (sorghum)

Triticum spp. (wheat)

Zea mays (maize)

Plants or Plant Parts

Corms, Rhizomes & Tubers

Seedlings & Nursery Stock

Seeds

Meloidogyne graminicola Indonesia

Allium cepa (onion)

Brassica spp. (cabbage, broccoli etc.)

Glycine max (soybean)

Oryza spp. (rice)

Solanum spp. (potato, aubergine, tomato etc.)

Sorghum spp. (sorghum)

Triticum spp. (wheat)

Zea mays (maize)

Soil

Plants or Plant Parts

Flower & Ornamental Plants

Roots and Rootstock

Radopholus citri Indonesia Citrus spp.

Soil

Roots and Rootstock

Seedlings & Nursery Stock

Plants or Plant Parts

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Table 2. Probable nematode threats.

Species

Basis of

Threat

Rating* Location in Region Common Hosts in Region Pathways

Ditylenchus destructor A Malaysia**

Solanum spp. (potato, aubergine, tomato etc.)

Ipomoea batatus sweet potato

Bulbs

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Bulbs

Seeds

Globodera rostochiensis R D Indonesia Solanum spp. (potato, aubergine, tomato etc.)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Longidorus elongatus D Vietnam

Allium cepa (onion)

Arachis hypogaea (peanut)

Beta vulgaris (beetroot)

Brassica spp. (cabbage, broccoli etc.)

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.)

Ficus spp. (figs)

Lactuca spp.

Phaseolus vulgaris (haricot bean)

Rosa spp. (rose)

Solanum spp. (potato, aubergine, tomato etc.)

Vitis spp. (grape)

Zea mays (maize)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Bulbs

Flower & Ornamental Plants

Meloidogyne enterolobii A Vietnam

Capsicum spp. (chilli, capsicum etc.)

Coffea spp. (coffee)

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.)

Psidium guajava (guava)

Solanum spp. (potato, aubergine, tomato etc.)

Soil

Plants or Plant Parts

Roots and Rootstock

Flower & Ornamental Plants

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Meloidogyne exigua D Thailand

Allium cepa (onion)

Capsicum spp. (chilli, capsicum etc.)

Citrullus lanatus (watermelon)

Citrus spp.

Coffea sp. (coffee)

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.)

Ipomoea spp.

Musa spp. (banana & plantain)

Oryza spp. (rice)

Saccharum officinarum (sugarcane)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Bulbs

Flower & Ornamental Plants

Meloidogyne naasi A Thailand

Allium cepa (onion)

Beta vulgaris (beetroot)

Oryza spp. (rice)

Solanum spp. (potato, aubergine, tomato etc.)

Sorghum spp. (sorghum)

Soil

Plants or Plant Parts

Roots and Rootstock

Flower & Ornamental Plants

Meloidogyne thailandica A Thailand Zingiber officinale (ginger)

Soil

Plants or Plant Parts

Roots and Rootstock

Flower & Ornamental Plants

Radopholus arabocoffeae N Vietnam Coffea spp. (coffee)

Soil

Plants or Plant Parts

Seedlings & Nursery Stock

Radopholus bridgei P Indonesia

Curcuma zedoaria (turmeric)†

Anubias sp., Echinodores sp., Ophiopogon sp (Aquarium

plants)‡

Soil

Plants or Plant parts

Roots and Rootstock

Flower and Ornamental Plants

Aquarium plants

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Radopholus similis R

PNG, Solomons,

Brunei, Indonesia,

Malaysia, Philippines,

Singapore, Thailand

Abelmoschus spp. (aibika, okra etc.)

Ananas comosus (pineapple)

Arachis hypogaea (peanut)

Bambusa spp. (bamboo)

Beta vulgaris (beetroot)

Brassica spp. (cabbage, broccoli etc.)

Camellia sinensis (tea)

Capsicum spp. (chilli, capsicum etc.)

Citrullus lanatus (watermelon)

Citrus spp.

Cocos nucifera (coconut)

Coffea sp. (coffee)

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.)

Dioscorea spp.

Ficus spp. (figs)

Gardenia sp.

Glycine max (soybean)

Heliconia spp.

Hibiscus spp.

Ipomoea spp.

Lactuca spp.

Litchi chinensis (lychee)

Mangifera spp. (mango)

Momordica charantia (bitter melon)

Musa spp. (banana & plantain)

Persea americana (avocado)

Petroselinum crispum (parsley)

Phaseolus vulgaris (haricot bean)

Pinus spp. (pine)

Piper spp. (pepper)

Psidium guajava (guava)

Saccharum officinarum (sugarcane)

Solanum spp. (potato, aubergine, tomato etc.)

Sorghum spp. (sorghum)

Theobroma cacao (cacao)

Triticum spp. (wheat)

Vigna spp. (bean)

Zea mays (maize)

Zingiber officinale (ginger)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Bulbs

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Xiphinema americanum s.l. R Philippines, Thailand

Allium cepa (onion)

Arachis hypogaea (peanut)

Bambusa spp. (bamboo)

Beta vulgaris (beetroot)

Bougainvillea spp.

Brassica spp. (cabbage, broccoli etc.)

Capsicum spp. (chilli, capsicum etc.)

Citrus spp.

Cocos nucifera (coconut)

Coffea sp. (coffee)

Cucurbitaceae spp. (cucumber, pumpkin, squash etc.)

Ficus spp. (figs)

Gardenia sp.

Glycine max (soybean)

Hibiscus spp.

Lactuca spp.

Litchi chinensis (lychee)

Mangifera spp. (mango)

Manilkara zapota (sapodilla)

Morus spp. (mulberry)

Musa spp. (banana & plantain)

Nicotiana tabacum (tobacco)

Oryza spp. (rice)

Passiflora spp. (passion fruit)

Persea americana (avocado)

Phaseolus vulgaris (haricot bean)

Pinus spp. (pine)

Psidium guajava (guava)

Rhododendron spp.

Rosa spp. (rose)

Saccharum officinarum (sugarcane)

Solanum spp. (potato, aubergine, tomato etc.)

Sorghum spp. (sorghum)

Triticum spp. (wheat)

Vigna spp. (bean)

Vitis spp. (grape)

Zea mays (maize)

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Bulbs

Flower & Ornamental Plants

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*—A: present in ASEAN countries only; R: only some host or pathogenicity races present in Australia; D: Distribution very restricted in Australia; P: pathogenicity not

proven

**—unconfirmed report (EPPO)

†— not on main crop host list, but included because native and widespread in Indonesia

‡—not on main crop host list, but included because of unusual pathway

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Table 3. Possible nematode threats in NAQS countries.

Species Country

Percentage of hosts

among crops

investigated*

Pathways

Meloidogyne arenaria Indonesia, PNG,

Solomons 60

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Roots and Rootstock

Flower and Ornamental Plants

Seeds & Nuts

Sea-swept Plant Debris

Meloidogyne hapla Indonesia, PNG 43

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Roots and Rootstock

Flower and Ornamental Plants

Sea-swept Plant Debris

Meloidogyne incognita Indonesia, PNG,

Solomons 78

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Roots and Rootstock

Flower and Ornamental Plants

Sea-swept Plant Debris

Meloidogyne javanica Indonesia, PNG,

Solomons 68

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Roots and Rootstock

Flower and Ornamental Plants

Sea-swept Plant Debris

Pratylenchus coffeae Indonesia, PNG,

Solomons 55

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Roots and Rootstock

Seedlings and Nursery Stock

Flower and Ornamental Plants

Sea-swept Plant Debris

Pratylenchus zeae Indonesia, PNG,

Solomons 27

Soil

Plants or Plant Parts

Corms, Rhizomes & Tubers

Roots and Rootstock

Flower and Ornamental Plants

Seedlings and Nursery Stock

Sea-swept Plant Debris

*—all nematode species identified as possible threats had a wide host range outside the crops investigated

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Table 4. Low nematode threats.

Aphelenchoides besseyi Pratylenchus crenatus

Aphelenchoides fragariae Pratylenchus delattrei

Aphelenchoides ritzemabosi Pratylenchus flakkensis Aphelenchoides subtenuis Pratylenchus goodeyi

Bitylenchus brevilineatus Pratylenchus hexincisus

Bitylenchus dubius Pratylenchus loosi

Bursaphelenchus mucronatus Pratylenchus neglectus

Bursaphelenchus rainulfi Pratylenchus penetrans

Bursaphelenchus singaporensis Pratylenchus pinguicaudatus

Bursaphelenchs thailandiae Pratylenchus pratensis

Cactodera cacti Pratylenchus pseudopratensis

Ditylenchus dipsaci Pratylenchus sudanensis

Helicotylenchus dihystera Pratylenchus teres

Helicotylenchus microcephalus Pratylenchus vulnus

Helicotylenchus multicinctus Quinisulcius acutus

Helicotylenchus pseudorobustus Quinisulcius capitatus

Helicotylenchus varicaudatus Quinisulcius curvus

Heterodera avenae Radopholus duriophilus

Heterodera cruciferae Radopholus nativus

Heterodera fici Rotylenchulus parvus

Heterodera oryzae Scutellonema brachyurum

Heterodera sacchari Scutellonema clathricaudatum

Heterodera schachtii Scutellonema minutum

Heterodera trifolii Trichodora cylindricus

Hemicriconemoides cocophilus Tylenchorhynchus annulatus

Hemicriconemoides mangiferae Tylenchorhynchus clarus

Hemicriconemoides strictathecatus Tylenchorhynchus nudus

Hoplolaimus galeatus Xiphinema basiri

Hoplolaimus indicus Xiphinema brasiliense

Hoplolaimus pararobustus Xiphinema brevicollum

Hoplolaimus seinhorsti Xiphinema index

Meloidogyne fallax Xiphinema insigne

Meloidogyne thamesi Xiphinema italiae

Meloidogyne trifoliophila Xiphinema pachtaicum

Merlinius brevidens Xiphinema rivesi

Pratylenchus alleni Xiphinema vuittenezi

Pratylenchus brachyurus

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Introduction

Background

Nematodes are microscopic round worms which occupy every ecosystem on earth, living

freely or in association with plants or animals. Of the approximately 25 000 known species,

about 15% (3600 species) are plant-parasitic but of these, only a few are considered major

economic pests, causing severe crop losses under certain conditions (Singh, unpublished

data). In developing nations, yield losses due to nematode damage amount to about 15% of

crop production per annum; in developed nations, this figure is 8.8% (Nicol et al., 2011).

Documented damage from plant-parasitic nematodes in Australia is over AUS$600 million

annually (Hodda & Cook, 2009).

Only 15% of species of plant-parasitic nematodes are known from Australia (Hodda & Nobbs

2008, Hodda & Singh 2010, Singh unpublished). Many exotic species have potential to

become invasive and damaging (Singh et al. in press). Because of the considerable cost and

difficulty in managing damaging nematodes once established, quarantine forms a major part

of minimising the impact of plant-parasitic nematodes in Australia.

Considering the potential economic importance of plant-parasitic nematodes, the Northern

Australia Quarantine Strategy group (NAQS) of the Department of Agriculture, Fisheries and

Forestry (DAFF) in June 2012 commissioned CSIRO to review the biosecurity threats posed

by plant-parasitic nematodes in the NAQS region.

Scope

Within Scope

This review examines the following factors relevant to the risk of entry by plant-parasitic

nematodes:

Presence in the NAQS countries as well as the ASEAN countries immediately to the

north;

Commonly grown host plants in above countries and northern Australia;

Pathways used to move (regulated or unregulated); and

Impact (geographic range and level of damage to plant hosts)

Beyond Scope

This review does not consider the following:

crops grown exclusively commercially (many hosts are grown both commercially and

domestically);

native or wild plant hosts;

countries outside the ASEAN region (north of South East Asia and East of the

Solomon Islands);

risk management measures to mitigate the hazard posed by these nematodes;

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detailed climatic or environmental preferences of nematode species (the subject of

current research by the third author);

detailed investigation and analysis of pathways and networks by which the nematodes

might move (the subject of current research by the second author); and

systematic uncertainties regarding the delimitation of nematode species (the subject of

current research by the first author).

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Methodology

Data collection

Information on the nematodes present in the NAQS and ASEAN countries was compiled

from available national checklists, consultations with national experts and information from

published articles to July 2012.

Nematode distributions were compiled from the published sources listed in the bibliography,

the Australian Faunal Directory, the CABI Crop Protection Compendium, and EPPO

publications.

Host lists were assembled from the CABI Crop Protection Compendium, NEMABASE, the

published sources listed in the bibliography and the comprehensive dataset compiled from all

published sources by the third author as part of his graduate studies (Singh submitted).

Common hosts only were investigated: defined as recorded in more than 3 NAQS or ASEAN

countries (excluding Australia). Lists of common crops were compiled from local experts

and NAQS staff, published lists, unpublished nematode host information from the ASEAN-

NAQS region and the unpublished dataset of the third author. Potential hosts grown

primarily commercially or in other parts of Australia were not investigated in detail, except

for wheat (Triticum aestivum) and soy bean (Glycine max), which, where included in the host

list of the greatest or probable threats, were used to assess the threats posed.

The plant genus Solanum contains over 1500 species, including the potato, tomato and

aubergine as well as many wild relatives (Frodin 2004). Two species of nematodes

considered in this review—Globodera rostochiensis and Globodera pallida—parasitize the

genus Solanum almost exclusively. In this review, the genus was considered as a single

"host". In considering the threat posed by the nematodes and host, the size and wide

distribution of the genus mean that biosecurity threats may be considerably greater than may

appear from the apparently small host list. The plant genera Piper (2000 species), Dioscorea

(800 species) and Ipomoea (600 species) are others considered in the present review which

contain many species but are listed as a single "host".

Pathway categories follow Centre for Agricultural Bioscience International (2012), as

modified by de la Peña et al. (2011) and Singh et al. (in press). Pathways were categorized

as NAQS and non-NAQS pathways based on whether or not they fit within a NAQS pathway

and loosely by what is permitted to be traded under the Torres Strait Treaty

(http://www.dfat.gov.au/geo/torres_strait/index.html). The plant species moving via the

various pathways were not considered in detail because there have been few studies of this

phenomenon. Plant movements are a part of current studies by the second author.

Pathways were determined from the published sources listed in the bibliography, the

unpublished dataset of the third author, and consultation with local experts. Feasibility of

pathways was inferred from distances between hosts as inferred from frequency of

occurrence data, mostly unpublished or informal from local experts or NAQS.

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Data Filters

More than 1200 species of nematodes have been reported as damaging plants (Singh

unpublished). Investigation of all was beyond the scope of this review, and in any case they

are being investigated as part of graduate studies by the third author. Hence a series of filters

was used to select the species most likely to be threats for detailed investigation (see below).

The following filters were applied: only species passing all filters were investigated in detail.

Filter 1: Recorded in NAQS or ASEAN countries.

Filter 2 Common crop hosts present in NAQS or ASEAN countries as well as in

Northern Australia.

Filter 3: Pathway present.

Likewise, many plant species are known as hosts for many of the nematode species

considered. For example, more than 3000 plant species are recorded as hosts for

Meloidogyne incognita and a similar number for M. javanica. Indeed, the hosts for most of

the nematode species investigated in detail include many native and non-crop plants which

may act as reservoirs or dispersal pathways for the nematodes identified as posing threats.

Detailed studies of dispersal are part of separate graduate studies by the second author. The

large numbers of hosts made investigation of all recorded hosts beyond the scope of this

review, so only 60 of the common crop hosts were investigated in detail (Table 13, p39).

Additional criteria for assessing threat posed

The threat posed by the species passing the above filters was assessed using the following

criteria. Whether species met each criterion was assessed using the data sources cited above,

and a level of threat estimated as the number of criteria met. If criterion 3 was not met, then

one of the secondary criteria had to be met.

Criterion 1: Pathogenicity to crops proven.

Criterion 2: Recorded in NAQS countries.

Criterion 3: Absent from Australia. A secondary criterion was that if recorded in

Australia, the species was restricted in distribution or range of races such as to

be a biosecurity threat: the races have been shown to vary in host range and/or

pathogenicity and there is evidence that not all races occur in Australia.

The process is outlined in Figure 1.

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Figure 1. Process for identification of nematode threats.

Plant-parasitic nematodes

List of 99 speciesPresent in ASEAN or

NAQS regions?

Hosts present in NAQS & ASEAN

countries & Australia?

Pathway present?

All criteria met?

Criterion 2 not met: not recorded in

NAQS countries

Criterion 3 not met:

present in Australia

Greatest threats

Yes

YesYes Yes

No

Evaluation criteria1. Damage to plants proven2. Recorded in NAQS countries3. Absent from Australia

Criterion 1 not met:

Damage to plants not

proven

Certain races

absent from

Australia

Host or pathogenicityraces known?

Very restricted

distribution in Australia

Filters

Probable threat

Possible threat

Probable threat

Probable threat

Probable threat

Information not available for Australia

No

Low threat

Low threat

Yes

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Results

Nematodes

A total of 98 nematode species were considered in detail as having passed filters 1-3. They

are listed according to their level of threat in Table 1,Table 2,Table 3 andTable 4 (pages 3, 5,

10 and 11), and consolidated in Table 12 (p37).

Seven species met all the criteria to be considered threats. They are listed in Table 1 (p3),

with full details presented in Appendix C—Detailed nematode profiles (p37).

An additional 11 species met all criteria except one. They are listed in Table 2 (p5), with full

details presented in Appendix C—Detailed nematode profiles (p37).

Five of these species have not been recorded in the NAQS region. Surveying for

nematodes within the NAQS countries have been limited (see Limitations of review, p26),

meaning that these species may be present, but unrecorded. This is more likely if the

species is widespread in the immediately adjacent area.

Five of these species have been recorded from Australia. Two of these have very

restricted distributions within Australia, being restricted to single localities in NSW and

SA, meaning that introduction to new areas is of concern. Three are known to have races

differing in hosts and pathogenicity (which under IPCC means that the distributions of the

races can be considered for biosecurity). The races present in NAQS countries are mostly

not known, however to be included in this group there had to be some evidence that not all

the races were currently present in Australia. All the species in this group are known

major pathogens, so introduction of new races would be a concern.

The pathogenicity of one species has not been demonstrated. Neither has the species been

shown to cause no damage, but few studies are available.

Races differing in host ranges and pathogenicity are recognised for four species of the genus

Meloiodgyne (Root-Knot Nematodes) and two species of the genus Pratylenchus (Root

Lesion Nematodes) found in both the NAQS countries and Australia. All cause large losses

in many parts of the world. None are listed as probable threats because there is no evidence

that only particular races are present in Australia, however there have been very few

investigations, either in Australia or the NAQS countries. Should future studies suggest that

few races are present in Australia, or that the races in the NAQS countries are different, then

the threats posed by these species should be re-assessed.

Of the nematode species identified as greatest or probable threats, most had relatively narrow

host ranges (Table 5, p18). By contrast, many of the probable threats and all of the possible

threats had very wide host ranges. However, the study was limited to crop hosts and many

non-crop hosts are not listed (see Limitations of review, p26). For example, Heterodera zeae

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and Meloidogyne graminicola occur on many graminaceous plants, and the single genus

Solanum, which is the host to Globodera pallida, contains over 1500 species.

Table 5. Number of the crops investigated which are hosts for nematode species identified as threats.

Greatest threats Probable threats Possible threats

Species hosts hosts hosts

Heterodera zeae 8 Radopholus similis 39 Meloidogyne incognita 47

Meloidogyne graminicola 6 Xiphinema americanum 33 Meloidogyne javanica 41

Heterodera glycines 4 Longidorus elongates 12 Meloidogyne arenaria 36

Ditylenchus angustus 1 Meloidogyne exigua 10 Pratylenchus coffeae 33

Globodera pallida 1 Meloidogyne enterolobii 5 Meloidogyne hapla 26

Hirschmanniella miticausa 1 Meloidogyne naasi 5 Pratylenchus zeae 16

Hirschmanniella oryzae 1 Radopholus bridgei 4

Radopholus citri 1 Ditylenchus destructor 3

Globodera rostochiensis 1

Radopholus arabocoffeae 1

Meloidogyne thailandica 1

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Geography

All but one of the species fulfilling all criteria as threats were reported from Indonesia, with

one from PNG and the Solomon Islands. None were from Timor Leste. Only 21 species

have been reported from PNG (SPC 2012): seven of these fulfil most but not all criteria as

threats (Table 1,Table 2 andTable 3, pages 3, 5 and 10). Only twelve species have been

reported from the Solomons (SPC 2012): seven of these fulfil most but not all criteria as

threats (Table 1,Table 2 andTable 3, pages 3, 5 and 10). The species lists from both

countries are almost certainly substantial underestimates resulting from lack of surveys.

There have been no surveys or reports of plant-parasitic nematodes from Timor Leste.

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Host Plants

Detailed lists of nematodes known to parasitize were available for 60 of the 66 common host

plants identified (Table 13, p39). All nematodes known from each of these hosts are listed in

Appendix C—Detailed nematode profiles (p37), with species identified as threats

highlighted.

The crop with the highest numbers of nematode species which were the greatest or potential

threats was Solanum spp. (Table 6, p21). Other crops with many nematode threats recorded

from them were Zea mays, Oryza spp. and Cucurbitaceae spp. (Table 6, p21). Capsicum

spp., Citrus spp., Sorghum spp., Allium cepa, Coffea spp. and Saccharum officinarum hosted

fewer of the nematodes identified as threats, but still considerable numbers (Table 6, p21).

Triticum aestivum (wheat) and Glycine max (soy bean) were both hosts for many of the

greatest or probable threats (Table 7, p21). These species are commercial crops grown

extensively in Australia, but outside the NAQS region. That these species are damaged by

many of the nematode threats identified means that the potential negative impact from the

nematodes includes other parts of Australia and other crops

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Table 6. Most frequent crop plant hosts for nematodes in the NAQS region and (number of nematode species recorded).

Greatest threats (8) Probable threats (11)* Greatest & Probable (19)* All nematodes (98)* Solanum spp. (5) Solanum spp. (6) Solanum spp. (11) Solanum spp. (70)

Oryza spp. (4) Cucurbitaceae spp.(6) Oryza spp. (7) Vitis spp. (61)

Zea mays (4) Capsicum spp. (5) Zea mays (7) Saccharum officinarum (60)

Sorghum spp. (2) Citrus spp. (4) Cucurbitaceae spp.(7) Oryza spp. (58)

Abelmoschus spp. (2) Coffea sp. (4) Sorghum spp. (5) Citrus spp. (58)

Allium cepa (1) Allium cepa (3) Citrus spp. (5) Zea mays (58)

Bambusa spp. (1) Brassica (3) Capsicum spp. (5) Musa spp. (50)

Citrus spp. (1) Ipomoea spp. (3) Allium cepa (4) Sorghum spp. (49)

Cucurbitaceae (1) Lactuca spp. (3) Saccharum officinarum (4) Pinus spp. (44)

Nicotiana tabacum (1) Musa spp. (3) Coffea sp. (4) Brassica (43)

Phaseolus vulgaris (1) Oryza spp. (3)

Saccharum officinarum (1) Phaseolus vulgaris (3)

Vigna spp. (1) Saccharum officinarum (3)

Sorghum spp. (3)

Zea mays (3)

*—partial list only.

Table 7. Major crop plant hosts outside the NAQS region for nematodes identifed as threats.

Greatest threats (8)* Probable threats (11)* Greatest & Probable (19)* All nematodes (98)* Triticum aestivum (4) Triticum aestivum (4) Triticum aestivum (8) Triticum aestivum (55)

Glycine max (2) Glycine max (3) Glycine max (5) Glycine max (48)

*—partial list only.

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Pathways

The pathways through which the greatest number of nematodes can move were plants or

plant parts, and soil (Table 8,Table 9 andTable 10, pages 23, 24 and 25).

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Table 8. Pathways for greatest nematode threats. Top row: coloured boxes= NAQS pathways and clear boxes= non-NAQS pathways.

Nematode species Soil

Plants or

Plant

Parts

Corms,

Rhizomes

& Tubers

Roots and

Rootstock

Seedlings &

Nursery

Stock

Flower &

Ornamental

Plants

Bulbs

Seeds

and

Nuts

Sea-swept

Plant

Debris

Ditylenchus angustus

Globodera pallida

Heterodera glycines

Heterodera zeae

Meloidogyne graminicola

Hirschmanniella miticausa

Hirschmanniella oryzae

Radopholus citri

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Table 9. Pathways for probable nematode threats. Top row: coloured boxes= NAQS pathways and clear boxes= non-NAQS pathways.

Nematode species Soil Plants or

Plant

Parts

Corms,

Rhizomes

& Tubers

Roots and

Rootstock

Seedlings &

Nursery

Stock

Flower &

Ornamental

Plants

Bulbs Seeds

& Nuts

Sea-swept

Plant

Debris

Ditylenchus destructor

Globodera rostochiensis

Longidorus elongatus

Meloidogyne enterolobii

Meloidogyne exigua

Meloidogyne naasi

Meloidogyne thailandica

Radopholus arabocoffeae

Radopholus bridge

Radopholus similis

Xiphinema americanum

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Table 10. Pathways for possible nematode threats. Top row: coloured boxes= NAQS pathways and clear boxes= non-NAQS pathways.

Nematode species

Soil Plants or

Plant

Parts

Corms,

Rhizomes

& Tubers

Roots &

Rootstock

Seedlings

& Nursery

Stock

Flower &

Ornamental

Plants

Bulbs Seeds

& Nuts

Sea-swept

Plant Debris

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus coffeae

Pratylenchus zeae

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Limitations of review

Assessing threat

The international definition of a quarantine pest is ―a pest of potential economic importance

to the area endangered thereby and not yet present there, or present but not widely distributed

and being officially controlled‖ (FAO 2011a). Organisms which meet this definition can be

regulated (FAO 2011b). These criteria are not very specific and results in long lists of

potential threats, investigation of all of which was beyond the resources available for the

present review. There are several more detailed methods for assessing the threats posed by

biological organisms which discriminate threats (Singh et al. in press). Most include

parameters such as climatic suitability, pathways, hosts, and economic impact, but how these

parameters are evaluated differs among methods. A best method has yet to emerge (Singh

2013).

In the present review all the common parameters cited above were included using a simple

dichotomous classification. (Presence of hosts was used a surrogate climatic suitability.)

This produced manageable—but still substantial—lists of nematodes. Detailed

investigations of improved methods may yield some different evaluations (Singh 2013).

Verification of material

Many of the nematode and host records used were unverified. Voucher material is extremely

sparse. Many nematodes are misidentified as major crop pests because pests are the best

known species, and hence have information and identification guides available, or be listed in

widely-available keys. However, the reverse also occurs, most particularly with introduced

pests being misidentified as local species, or major pests not being recognised on the basis of

causing different symptoms in different situations or hosts.

The nematode fauna of Australia's north, the NAQS and ASEAN countries are generally very

poorly characterised. There are undoubtedly many more species present than are currently

recorded. How many of these will be of significance to biosecurity is unknown.

Uncertainty of identifications

Several national lists contained nematodes identified to genus level only (notably PNG and

the Solomon Islands). Altogether 18 genera contain unidentified species (Table 11, p29).

Two genera (Bursaphelenchus and Dolichodorus) were only recorded from unidentified

species. These records were not included in this review although most of the genera contain

recognised major crop pests: this may have lead to some underestimation of the nematode

threats. The unidentified species are included in the lists of nematodes recorded by crop for

reference (Appendix D—Nematodes recorded on host crop, p62).

Limited survey data

No records of plant-parasitic nematodes could be located for several countries (e.g. Myanmar

and Timor Leste). Records from other countries were extremely old and in need of updating

(eg PNG, Thailand).

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Uncertainty over damage caused

Lack of information on damage caused by plant-parasitic nematodes is a major uncertainty in

assessing their biosecurity threats (Singh et al. in press). Many of the species investigated in

detail passed all filters, and are known to parasitise crop plants, but the damage caused has

not been investigated or documented. The threats posed by some of these species may need

reassessing should more information become available.

Climatic suitability for establishment

The suitability of the environment in Northern Australia for establishment of nematodes (eg

climate or soil types) was not included in this review because information on preferences and

limits is limited for many nematode species. Environmental conditions can be important

determinants of nematode abundances and the damage they inflict, as well as their success in

establishing. The suitability of the different parts of Australia for different nematode pests of

plants is the subject of separate studies by the third author. These studies should add

considerably to the present review and should be available early in 2013.

Host races or pathotypes

Under the IPPC, races or pathotypes must differ in hosts or pathogenicity to be considered

significant for biosecurity. Such races have been studied and accepted in many nematode

species. Many have been found in both Australia and the NAQS countries (and the ASEAN

countries, too). Whether the races are a biosecurity threat depends on their distribution in the

countries being considered. Although the races have been known for many years in some

cases (Swanson & Van Gundy, 1984 for Meloidogyne, Kort et al., 1974 for Globodera,

Mizukubo 1995, Mizukubo & Sano 1997 and Kumar 1991 for some species of Pratylenchus),

the particular races present in Australia and the NAQS countries remain inadequately known,

particularly for then nematode genera Meloidogyne and Pratylenchus. Studies in several

countries have found that nematode races often vary substantially between countries, with

significant implications for how nematodes are managed (eg Hodda 2004, Rammah and

Hirschmann 1990, Moritz et al., 2003, Robertson et al., 2009). It is thus entirely likely that

the races of Meloidogyne and Pratylenchus present differ between Australia and the NAQS

or ASEAN countries. However, this has not been demonstrated to date, so the species

involved were only regarded as possible threats in the present review. More reserch is

needed to clarify the situation.

By contrast with Meloidogyne and Pratylenchus, there is evidence that only a very limited

number of races of Globodera rostochiensis and Radopholus similis are present in Australia

(Hodda 2004, Quader et al. 2008, Gafur unpublished). Although the races in the NAQS and

ASEAN countries are yet to be characterised, they are likely to be different, so these species

were regarded as probable threats in the present review. A very limited number of races of

Ditylenchus dipsaci seem present as well, but there have been no reports from the NAQS

countries, so this species is only regarded as a low threat via these countries.

No races are known for Xiphinema americanum sensu lato, but in many ways the situation is

similar to the three species discussed in the preceding paragraph. Xiphinema americanum

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s.l. is a species complex, with many very closely related species extremely difficult to

differentiate by any means, morphological or molecular (Gozel et al. 2006, Hunt 1993).

Hence, nematodes identified as Xiphinema americanum s. l.—particularly older records (see

above)—could be any one of up to about 40 species (Hunt 1993). Although there are records

of Xiphinema americanum s. l. from both Australia and the NAQS countries, the differences

in hosts and locations make it unlikely that they refer to the same species within the complex.

Hence Xiphinema americanum s. l. was regarded as a probable threat.

Impacts outside the NAQS region

The potential of nematodes to damage crop plants other than those common in the NAQS

region of Australia was mostly not considered in this review. Of these crops, only two very

important species were investigated briefly: wheat (Triticum aestivum) and soy bean (Glycine

max). Both crops were hosts to many of the nematode threats identified. The threat posed

by the nematodes to other hosts outside the NAQS region was beyond the scope of this

review, and remains to be investigated.

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Table 11. Genera with unidentified species from NAQS or ASEAN countries.

Genus Only record in genus?

Aphelenchoides

Bursaphelenchus X

Dolichodorus X

Ditylenchus

Globodera

Helicotylenchus

Heterodera

Hemicriconemoides

Hoplolaimus

Longidorus

Meloidogyne

Pratylenchus

Radopholus

Rotylenchulus

Rotylenchus

Scutellonema

Trichodorus

Tylenchorhynchus

Xiphinema

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References

Allen, M.W., Hart, W.H. & Baghott, K.V. (1970). Crop rotation controls the barley root-knot nematode at

Tulelake. California Agriculture 24,4-5.

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Appendix A—List of nematode species evaluated

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Table 12. List of all nematode species considered as part of this review.

Aphelenchoides besseyi Aphelenchoides fragariae

Aphelenchoides ritzemabosi

Aphelenchoides subtenuis

Bitylenchus brevilineatus

Bitylenchus dubius

Bursaphelenchus mucronatus

Bursaphelenchus rainulfi

Bursaphelenchus singaporensis

Bursaphelenchs thailandiae

Cactodera cacti

Ditylenchus angustus

Ditylenchus destructor

Ditylenchus dipsaci

Globodera pallida

Globodera rostochiensis

Helicotylenchus dihystera

Helicotylenchus microcephalus

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Helicotylenchus varicaudatus

Heterodera avenae

Heterodera cruciferae

Heterodera fici

Heterodera glycines

Heterodera oryzae

Heterodera sacchari

Heterodera schachtii

Heterodera trifolii

Heterodera zeae

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Hirschmanniella miticausa

Hirschmanniella oryzae

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Longidorus elongatus

Meloidogyne arenaria

Meloidogyne enterolobii

Meloidogyne exigua

Meloidogyne fallax

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne naasi

Meloidogyne thailandica

Meloidogyne thamesi

Meloidogyne trifoliophila

Merlinius brevidens

Pratylenchus alleni

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus delattrei

Pratylenchus flakkensis

Pratylenchus goodeyi

Pratylenchus hexincisus

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pinguicaudatus

Pratylenchus pratensis

Pratylenchus pseudopratensis

Pratylenchus scribneri

Pratylenchus sudanensis

Pratylenchus teres

Pratylenchus thornei

Pratylenchus vulnus

Pratylenchus zeae

Quinisulcius acutus

Quinisulcius capitatus

Quinisulcius curvus

Radopholus arabocoffeae

Radopholus bridgei

Radopholus citri

Radopholus duriophilus

Radopholus nativus

Radopholus similis

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchus robustus

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema minutum

Trichodora cylindricus

Tylenchorhynchus annulatus

Tylenchorhynchus brassicae

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus mashhoodi

Tylenchorhynchus nudus

Xiphinema americanum

Xiphinema basiri

Xiphinema brasiliense

Xiphinema brevicollum

Xiphinema index

Xiphinema insigne

Xiphinema italiae

Xiphinema pachtaicum

Xiphinema rivesi

Xiphinema vuittenezi

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Appendix B—List of all host plants considered

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Table 13. List of all host plants considered in this review.

Scientific name Common name(s) Scientific name Common name(s)

Abelmoschus spp. aibika, okra Morus spp. mulberry

Allium cepa Onion Musa spp. banana & plantain

Ananas comosus pineapple Nephelium lappaceum rambutan

Annona muricata soursop Oryza spp. rice

Annona reticulata custard apple Passiflora spp. passion fruit

Annona squamosa sugar apple Persea americana avocado

Arachis hypogaea peanut Petroselinum crispum parsley

Bambusa spp. bamboo Phaseolus vulgaris haricot bean

Beta vulgaris beetroot Pinus spp. pine

Bougainvillea spp. bougainvillea Piper spp. pepper

Brassica spp. cabbage, broccoli Psidium guajava guava

Calopogonium mucunoides wild ground nut Psophocarpus

tetragonolobus winged bean

Camellia sinensis tea Rosa spp. rose

Capsicum spp. chilli, capsicum Saccharum officinarum sugarcane

Carica papaya pawpaw Solanum spp. potato, aubergine, tomato

Citrullus lanatus watermelon Sorghum spp. sorghum

Citrus spp. citrus Theobroma cacao cacao

Cocos nucifera coconut Vigna spp. bean

Coffea spp. coffee Vitis spp. grape

Colocasia esculenta taro Xanthosoma spp. new cocoyam

Coriandrum sativum coriander Zea mays maize

Cucurbitaceae spp. cucumber, pumpkin,

squash Zingiber officinale ginger

Dimocarpus longan longan

Dioscorea spp. yam

Dracaena spp. dragon tree

Durio spp. durian

Elettaria cardamomum cardamom

Ficus spp. figs

Gardenia spp. gardenia

Heliconia spp. lobster-claws, wild

plantains

Hibiscus spp. hibiscus

Ipomoea spp. sweet potato

Lactuca spp. lettuce

Litchi chinensis lychee

Mangifera spp. mango

Manihot esculenta cassava

Manilkara zapota sapodilla

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Appendix C—Detailed nematode profiles

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Ditylenchus angustus

Threat: Greatest

Taxonomy

Name: Ditylenchus angustus (Butler, 1913) Filipjev 1936

Synonyms: Tylenchus angustus Butler, 1913

Anguillulina angusta (Butler) Goodey, 1932

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Anguinidae

Common Name: Rice stem nematode

Biological characteristics

The nematode causes ufra disease, a high impact disease of deepwater and lowland rice. Plants parasitized by

this nematode become more susceptable to other plant pathogens, such as Pyricularia oryzae (Bridge et al.,

2005).

A migratory ectoparasite living on the outer surface of plants and feeding on plant shoot tissue. The lifecycle of

the nematode is short, 10-20 days and it completes several generations on one plant. Post-harvest, it survives in

a dessicated state in crop residues (Bridge & Starr, 2007; Whitehead, 1998).

The nematode is widespread in countries in Asia and Africa but is restricted to areas of high humidity (≥75% ).

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Myanmar, Thailand, Vietnam, Malaysia, Philippines, Indonesia

Also Egypt, Sudan, Madagascar, India, Pakistan, Bangladesh

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Oryza spp. (rice)

Filter 3: Pathway present

Proven: Plants or plant parts; seedlings and nursery stock; seeds; soil

Probable: Corms, rhizomes and tubers; flower and ornamental plants; bulbs.

Criterion 1: Recorded in NAQS countries

Yes; Indonesia.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Yield losses range from 4-100% (Bridge et al. 2005, Whitehead 1998).

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Ditylenchus destructor

Threat: Probable

Taxonomy Name: Ditylenchus destructor Thorne 1945

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Anguinidae

Common Name: Potato Tuber Nematode, Potato Rot Nematode

Biological characteristics On potato, plants are weakened and usually die when tubers are heavily infested Such tubers have sunken areas

with cracked, wrinkled and partly detached skin, with dry and mealy flesh, greyish to dark brown or black in

colour. The discoloration may be due to secondary invasion of fungi, bacteria and other nematodes. Iris have

grey to black lesions, often black roots and poorly-developed leaves with yellow tips. Groundnuts have black

discoloration appearing first along longitudinal veins, shrunken kernels, brown to black testae and brown

embryo (Jones & De Waele 1988).

A migratory endoparasite living in the soil, tubers and bulbs. The lifecycle of the nematode is short: 10 days at

28°C (De Waele et al., 1990).

Seems unable to withstand excessive desiccation, so usually restricted to moist soils. Before D. destructor was

described, it was regarded for a long time as a strain or race of D. dipsaci and recorded as such. The nematodes

attacking ground nut may be a separate race (Jones & De Waele 1988).

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Malaysia

Also found in isolated places and countries in Africa, Europe, East and North Asia and The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Solanum spp. (potato, aubergine, tomato etc.), Ipomoea spp., Iris spp.

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts; Corms, Rhizomes & Tubers; Bulbs; Seeds

Criterion 1: Recorded in NAQS countries

No

Criterion 2: Absent from Australia

Recorded from some time ago, but records doubtful and no recent findings, so now regarded as

absent (Hodda 2009a, b, Thistlethwayte 2009, 2010).

Criterion 3: Pathogenicity to crops proven

Yield losses in ground nut (Jones & De Waele 1988) and potato (Winslow 1978).

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Globodera pallida

Threat: Greatest

Taxonomy Name: Globodera pallida (Stone, 1973) Behrens, 1975

Synonyms: Heterodera pallida Stone, 1973

Globodera pallida (Stone) Mulvey & Stone, 1976

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Heteroderidae

Common Name: Potato Cyst Nematode, Pale Nematode

Biological characteristics The cause of severe losses through stunting and plant death in potato. Entry wounds provide a point of entry

for pathogens such as Pseudomonas solanacearum (Jatala et al., 1976) and Verticillium dahliae (Harrison, 1971;

Franco & Bendezu, 1985).

A sedentary endoparasite which feeds on plant roots. Juvenile nematodes hatch from eggs in the soil and enter

the roots where they establish a permanent feeding site. Once mature, males leave the roots to mate with

females which remain within the root with their posterior ends protruding outside. Fertilized females become

enlarged and rounded, eventually dying but retaining up to 500 eggs within the tanned body (Scurrah et al.

2005). Without hosts, cysts may remain viable in soil for up to 30 years (Turner, 1996).

Occurs in many countries, but in highly restricted areas. Similar climatic preferences to its host the potato.

Races are recognised (Kort et al., 1977).

Filter 1: Present in ASEAN-NAQS region

ASEAN region: Malaysia, Indonesia

Australo-Pacific: New Zealand

Also found in isolated places and countries in Africa, Europe, East and North Asia and The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Solanum spp. (potato, aubergine, tomato etc.)

Filter 3: Pathway present

Proven: Soil; corms, rhizomes and tubers

Probable: Plants and plant parts

Criterion 1: Recorded in NAQS countries

Yes; Indonesia.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Yield losses of up to 100% (Brodie & Mai 1989).

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Globodera rostochiensis

Threat: Probable

Taxonomy

Name: Globodera rostochiensis (Wollenweber 1923) Behrens 1975

Synonyms: Heterodera rostochiensis Wollenweber 1923

Common names: Potato Cyst Nematode, Golden Nematode

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Heteroderidae

Biological characteristics

The cause of severe losses through wilting, stunting and plant death in potato. Entry wounds provide a point of

entry for pathogens such as Pseudomonas solanacearum (Jatala et al., 1976) and Verticillium dahliae (Harrison,

1971; Franco & Bendezú, 1985).

A sedentary endoparasite which feeds on plant roots. Juvenile nematodes hatch from eggs in the soil and enter

the roots where they establish a permanent feeding site. Once mature, males leave the roots to mate with

females which remain within the root with their posterior ends protruding outside. Fertilized females become

enlarged and rounded, eventually dying but retaining up to 500 eggs within the tanned body (Scurrah et al.

2005). Without hosts, cysts may remain viable in soil for up to 30 years (Turner, 1996).

Occurs in many countries, but in highly restricted areas. Similar climatic preferences to its host the potato.

Races are recognised (Kort et al., 1977).

Filter 1: Recorded in ASEAN or NAQS countries

Philippines, Indonesia

Also in New Zealand, and found in isolated places and countries in Africa, Europe, East and North

Asia and The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Solanum spp. (potato, aubergine, tomato etc.)

Filter 3: Pathway present

Proven: Soil; corms, rhizomes and tubers

Probable: Plants and plant parts

Criterion 1: Recorded in NAQS countries

Yes; Indonesia.

Criterion 2: Absent from Australia

Presently in a defined area of Victoria only and under regulation. Recently declared eradicated

from Western Australia (Collins et al ., 2010). Only one race found in Victoria (Quader et al.

2008).

Criterion 3: Pathogenicity to crops proven

Yield losses of up to 100% (Brodie & Mai 1989).

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Heterodera glycines

Threat: Greatest

Taxonomy Name: Heterodera glycines Ichinohe 1952

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Heteroderidae

Common Name: Soyabean Cyst Nematode

Biological characteristics A major crop pest preventing nodule formation in plants, causing ―yellow dwarf‖ disease. Also found to

interact with Fusarium solani in ―sudden death‖ syndrome of soybean.

A sedentary endoparasite living inside roots. Juveniles hatch from eggs in soil, enter roots and migrate to

vascular tissue, then become sedentary and establish a feeding site. Adult males leave the roots to fertilize adult

females protruding from the root surface. Fertilised females form lemon-shaped cysts attached to the roots

(Oyekanmi & Fawole, 2010). H. glycines can undergo 3-4 generations per growing season in cool areas and 6-

7 generations in hotter regions (Noel, 1985).

Preferred temperature is 23–28°C: above 34°C development stops (Schmitt & Riggs, 1991; Burrows & Stone,

1985). Without hosts, cysts may remain viable in soil for 6–8 years (Slack et al., 1972). Races are recognised

(Riggs & Schmidt 1988, Niblack et al., 2002).

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Indonesia

Also in isolated places and countries in Africa, Europe, East and North Asia and The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Beta vulgaris (beetroot), Cucurbitaceae spp. (cucumber, pumpkin, squash etc.), Glycine max

(soybean), Phaseolus vulgaris (haricot bean), Solanum spp. (potato, aubergine, tomato etc.), Vigna

spp. (bean), Zea mays (maize)

Filter 3: Pathway present

Proven: Soil

Probable: Roots and Rootstock; Seedlings & Nursery Stock; Plants or Plant Parts

Criterion 1: Present in NAQS region of interest

Yes; Indonesia.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Recorded yield losses range from 10-80% (Jacobsen et al., 1983).

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Heterodera zeae

Threat: Greatest

Taxonomy

Name: Heterodera zeae Koshy, Swarup & Sethi, 1971

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Heteroderidae

Biological characteristics

An important crop pest causing general disease symptoms. Disease symptoms increase in the presence of

Cephalosporium maydis in the field/laborartory testing, indicating that the nematode may increase the plant‘s

susceptability to other plant pathogens.

A sedentary endoparasite living inside roots. Juveniles hatch from eggs in soil, enter roots and migrate to

vascular tissue, then become sedentary and establish a feeding site. Adult males leave the roots to fertilize adult

females protruding from the root surface. Fertilised females form lemon-shaped cysts attached to the roots

The lifecycle is short at 15-18 days, with 6-7 generations in 1 crop season (McDonald & Nicol, 2005). Cysts

remain viable for a long time.

Environmental preferences poorly known.

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Thailand, Indonesia

Also in New Zealand, and found in isolated places and countries in Africa, Europe, East and North

Asia and The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Bambusa spp. (bamboo), Capsicum spp. (chilli, capsicum etc.), Citrus spp., Oryza spp. (rice),

Saccharum officinarum (sugarcane), Solanum spp. (potato, aubergine, tomato etc.), Sorghum spp.

(sorghum), Triticum spp. (wheat), Zea mays (maize)

Filter 3: Pathway present

Proven: Soil

Probable: Roots and Rootstock; Seedlings & Nursery Stock; Plants or Plant Parts

Criterion 1: Recorded in NAQS countries

Yes; Indonesia.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

One of the most economically important nematdoes attacking cereals with reported yield losses of

13-73% (Krusberg et al., 1997).

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Hirschmanniella miticausa

Threat: Greatest

Taxonomy

Name: Hirschmanniella miticausa Bridge, Mortimer & Jackson 1983

Synonyms: None

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Pratylenchidae

Common Name: None

Biological characteristics

The nematode causes ‗miti miti‘ disease or corm rot of taro (Calocasia esculentum). The above ground

symptoms include chlorotic leaves and heavy infestation by this nematode causes plant death due to corm

damage (Bridge et al., 1983). Infested corms, when cut exhibit red streaks radiating from the base of the corm.

A migratory endoparasite living on the outer surface of plants and feeding on corm and root tissue. The

lifecycle of the nematode has not been studied and little is known about its true host range.

The nematode has been reported from Solomon Islands and PNG, from taro grown in dryland soils, rainfed

mountain slopes and in flooded swamp pits.

Filter 1: Recorded in ASEAN or NAQS countries

NAQS region: PNG, Solomon Islands

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Colocasia esculentum (taro)

Filter 3: Pathway present

Proven: Soil; Corms, Rhizomes and Tubers; Plants or Plant Parts; Seedlings and Nursery Stock

Probable: Flower and Ornamental Plants; Bulbs; Sea Swept Plant Debris

Criterion 1: Recorded in NAQS countries

Yes; PNG, Solomon Islands

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

The corms of C. esculenta infested by this nematode are inedible. In the Solomon Islands, the

impacts on production are so severe that cultivation of taro is abandoned in fields with high

populations of this nematode.

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Hirschmanniella oryzae

Threat: Greatest

Taxonomy Name: Hirschmanniella oryzae (van Breda de Hann, 1902) Luc & Goodey 1964

Synonyms: Tylenchus oryzae van Breda de Haan 1902

Anguillulina oryzae (van Breda de Haan) Goodey 1932

Hirschmannia oryzae (van Breda de Haan) Luc & Goodey 1962

Radopholus oryzae (van Breda de Haan) Thorne 1949

Rotylenchus oryzae (van Breda de Haan) Filipjev & Schuurmans Stekhoven 1941

Tylenchus apapillatus Imamura 1931

Anguillulina apapillata (Imamura) Goodey 1932

Rotylenchus apapillatus (Imamura) Filipjev 1936

Hirschmanniella apapillata (Imamura) Siddiqi 1986

Hirschmanniella nana Siddiqi 1966

Hirschmanniella exigua Khan 1972

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Pratylenchidae

Common Name: Rice Root Nematode

Biological characteristics This nematode thrives in rice fields and is able to withstand flooded conditions. The above-ground symptoms

of damage by this nematode are not easily identifiable. The symptoms include slow growth and reduced

tillering, which can be confused with symptoms of nutrient deficiency.

Hirschmanniella oryzae is a migratory endoparasite and feeds on the root tissue of rice plants. The lifecycle of

this nematode under ideal conditions can be completed in 30 days but the number of generations is variable with

1-2 generations per year (Kuwahara & Kisabu, 1970).

The nematode is widespread in rice growing regions in Asia and has restricted distributions in parts of Africa,

Central America and Caribbean, Europe, North America and South America.

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Malaysia, Myanmar, Philippines, Thailand, Vietnam

NAQS region: Indonesia

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Abelmoschus spp. (aibika, okra etc.), Oryzae spp. (rice), Saccharum officinarum (sugarcane),

Solanum spp. (potato, aubergine, tomato etc.), Sorghum spp. (sorghum), Triticum spp. (wheat), Zea

mays (maize)

Filter 3: Pathway present

Proven: Soil, Plants or plant parts; seedlings and nursery stock

Probable: Corms, rhizomes and tubers; flower and ornamental plants; bulbs; sea swept plant debris

Criterion 1: Recorded in NAQS countries

Yes; Indonesia

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Important pest of rice, yield reduced by 25-39% (Babatola & Bridge 1980; Cho et al. 1994)

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Longidorus elongatus

Threat: Probable

Taxonomy

Name: Longidorus elongatus (De Man 1876) Micoletzky 1922

Synonyms: Dorylaimus elongatus De Man 1876

Trichodorus elongatus (De Man 1876) Filipjev 1921

Dorylaimus (Longidorus) elongatus De Man 1876 (Micoletzky 1922)

Dorylaimus tenuis Linstow 1879

Longidorus monohystera Altherr 1953

Longidorus menthasolanus Konicek & Jensen 1961

Taxonomic Position: Nematoda: Dorylaimea: Dorylaimida: Dorylaimina: Longidoridae

Common Name: No common name

Biological characteristics Direct feeding causes severe stunting and galling of root systems in many crops, and is associated with Docking

Disorder of sugarbeet (Whitehead & Hooper 1970). Transmits plant viruses including Rasperry Ringspot Virus

and Tomato Black Ring Virus.

A migratory ectoparasite living on the outer surface of plants and feeding on plant roots. The lifecycle of the

nematode is 9 weeks at 30ºC (Wyss 1970) but probably only one genreation per year occurs in the field

(Thomas 1969).

The nematode is widespread in temperate regions, with more limited occurrence in the tropics.

Filter 1: Recorded in ASEAN-NAQS region

ASEAN region: Myanmar, Thailand, Vietnam, Malaysia, Philippines, Indonesia

Also New Zealand, and found in isolated places and countries in Europe, East and North Asia and The

Americas.

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Allium cepa (onion), Arachis hypogaea (peanut), Beta vulgaris (beetroot), Brassica spp. (cabbage,

broccoli etc.), Cucurbitaceae (cucumber, pumpkin, squash etc.), Ficus spp. (figs), Lactuca spp.,

Phaseolus vulgaris (haricot bean), Rosa spp., Solanum spp., Vitis spp., Zea mays.

Filter 3: Pathway present

Proven: : Soil; Seedlings & Nursery Stock

Probable: Plants or Plant Parts; Corms, Rhizomes & Tubers; Bulbs; Flower & Ornamental Plants

Criterion 1:

Yes; Indonesia.

Criterion 2: Absent from Australia

One record from Lolium sp. in South Australia.

Criterion 3: Pathogenicity to crops proven

Yes (Hunt 1993).

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Meloidogyne enterolobii

Threat: Probable

Taxonomy

Name: Meloidogyne enterolobii Yang & Eisenback, 1983

Synonyms: Meloidogyne mayaguensis Rammah & Hirschmann, 1988

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Meloidogynidae

Common Name: Pacara Earpod Tree Root-Knot Nematode

Biological characteristics

An emerging threat to vegetable production recently included on the quarantine list for EPPO. It causes serious

root galling and plant decline via a combination of a high reproduction rate, large galls and a very wide host

range. It is virulent against several sources of resistance to other Meloidogyne species and can overcome the

important Mi-1 resistance gene (Castagnone-Sereno 2012).

A sedentary endoparasite feeding on underground plant tissues, especially roots. After hatching, juveniles

locate and enter roots, then establish a permanent feeding site. A gall is induced around this feeding site as the

juveniles become sedentary and mature into females. The females remain in the gall for the rest of their

lifecycle and lay eggs in a gelationous matrix outside the root. This species can complete its lifecycle in 24

days (Westerich et al., 2011).

Seems to prefer mild conditions under which many vegetables grow. It is also a particular pest in enclosed

production systems (greenhouses).

Filter 1: Present in ASEAN-NAQS region

ASEAN region: Vietnam

Mainly found in Central America, it is also found in isolated places and countries in Africa, Europe,

East and North Asia

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Capsicum spp. (chilli, capsicum etc.), Coffea sp. (coffee), Cucurbitaceae spp. (cucumber, pumpkin,

squash etc.), Psidium guajava (guava), Solanum spp. (potato, aubergine, tomato etc.), Nicotiana

tabacum (tobacco)

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts; Roots and Rootstock; Flower & Ornamental Plants

Criterion 1: Recorded in NAQS countries

No.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Causes losses of up to 65% in some crops (Cetintas et al. 2007).

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Meloidogyne exigua

Threat: Probable

Taxonomy

Name: Meloidogyne exigua Goeldi, 1892

Synonyms: Heterodera exigua (Goeldi) Marcinowski, 1909

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Meloidogynidae

Common Name: Coffee Root-Knot Nematode

Biological characteristics

Coffee is the main host, however it can parasitise a wide range of other plants as well. Above-ground

symptoms on coffee include yellowing of leaves, leaf fall and tree death in case of heavy infestation.

Adult females are sedentary endoparasites, residing and feeding in specialised cells which form characteristic

galls on the roots. The lifecycle of this nematode can be completed in 15-30 days (Lima & Ferraz 1985).

The nematode is widespread in coffee growing regions in Central and South America. Incidential reports from

China, Greece, India and Thailand (Srinivasan & D'Souza 1965; Zhang & Xu 1994; Karssen & Van Hoenselaar

1998).

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Thailand

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Allium cepa (onion), Capsicum spp. (chilli, capsicum etc.), Citrullus lanatus (watermelon), Citrus

spp., Coffea sp. (coffee), Cucurbitaceae spp. (cucumber, pumpkin, squash etc.), Ipomoea spp., Musa

spp. (banana & plantain), Oryza spp. (rice), Saccharum officinarum (sugarcane)

Filter 3: Pathway present

Proven: Plants or plant parts; seedlings and nursery stock; seeds; soil

Probable: Corms, rhizomes and tubers; flower and ornamental plants; bulbs.

Criterion 1: Recorded in NAQS countries

No

Criterion 2: Absent from Australia

No, but only recorded from New South Wales.

Criterion 3: Pathogenicity to crops proven

Pathogenic to coffee; caused yield loss of 45% (Barbosa et al., 2004; Barbosa et al., 2010)

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Meloidogyne graminicola

Threat: Greatest

Taxonomy Name: Meloidogyne graminicola Golden & Birchfield, 1965

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Meloidogynidae

Common Name: Rice Root-Knot Nematode

Biological characteristics

Causes serious disease of rice often involving swollen and hooked root tips. Symptoms in dry or shallow water

rice include stunting, reduced tillering, wilting, unfilled spikelets, and chlorosis. In deepwater rice, plants do

not emerge above water (Bridge & Page 1982).

A sedentary endoparasite feeding on underground plant tissues, especially roots. After hatching, juveniles

locate and enter roots, establish a permanent feeding site. A gall is induced around this feeding site as the

juveniles become sedentary and mature into females. The females remain in the gall for the rest of their

lifecycle and lay eggs in a gelationous matrix outside the root.

This species prefers wet conditions but survives in dry conditions as well. M. graminicola can survive in soil

flooded to a depth of 1 m for at least 5 months (Bridge and Page, 1982) and egg masses can remain viable for at

least 14 months in waterlogged soil (Roy, 1982).

Filter 1: Present in ASEAN-NAQS region

ASEAN region: Thailand, Myanmar, Lao PDR, Vietnam, Malaysia, Singapore, Philippines,

Indonesia

Very isolated occurrences outside SE Asia: Europe, East and North Asia and The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Allium cepa (onion), Brassica spp. (cabbage, broccoli etc.), Glycine max (soybean), Oryza spp. (rice),

Solanum spp. (potato, aubergine, tomato etc.), Sorghum spp. (sorghum), Triticum spp. (wheat), Zea

mays (maize)

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts; Roots and Rootstock; Flower & Ornamental Plants

Criterion 1: Recorded in NAQS countries

Yes; Indonesia.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

A major rice pest with recorded yield losses of up to 80% (De Waele & Elsen, 2007).

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Meloidogyne naasi

Threat: Probable

Taxonomy Name: Meloidogyne naasi Franklin, 1965

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Meloidogynidae

Common Name: Barley Root-Knot Nematode

Biological characteristics A serious pest of grasses and other plants, inducing stunting and severe root galling with sometimes more than

100 nematodes per gall. Galls are curved, horseshoe or spiral shaped (Kort, 1972). May interact with

Fusarium spp. to cause corchosis (a disease) in plants (Manzanilla-López & Starr, 2009).

A sedentary endoparasite feeding on underground plant tissues, especially roots. After hatching, juveniles

locate and enter roots, establish a permanent feeding site. A gall is induced around this feeding site as the

juveniles become sedentary and mature into females. The females remain in the gall for the rest of their

lifecycle and lay eggs in a gelationous matrix outside the root. There is usually only 1 generation per growing

season (Rivoal and Cook, 1993).

Prefers temperate climates.

Filter 1: Present in ASEAN-NAQS region

ASEAN region: Thailand

Also in New Zealand, and found in Europe as well as isolated places and countries in North Asia and

The Americas

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Allium cepa (onion), Beta vulgaris (beetroot), Oryza spp. (rice), Solanum spp. (potato, aubergine,

tomato etc.), Sorghum spp. (sorghum), Triticum spp. (wheat)

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts; Roots and Rootstock; Flower & Ornamental Plants

Criterion 1: Recorded in NAQS countries

No.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Reported to cause significant damage with up to 75% yield loss in some crops and 100% losses

under some conditions (Allen et al., 1970; Schneider, 1967).

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Meloidogyne thailandica

Threat: Probable

Taxonomy Name: Meloidogyne thailandica Handoo, Skantar, Carta & Erbe, 2005

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Meloidogynidae

Biological characteristics

Unknown. This species has not been studied in its native locality (Thailand). Species was described based on

specimens obtained from intercepted infected ginger sent to the USA from Thailand (Handoo et al., 2005).

Species within this genus are sedentary endoparasites which enter plant tissues as juveniles and establish a

permanent feeding site inducing gall formation.

Filter 1: Present in ASEAN-NAQS region

Asia: Thailand

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Zingiber officinale (ginger)

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts; Roots and Rootstock; Flower & Ornamental Plants

Criterion 1: Recorded in NAQS countries

No.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

This species has not been proven to damage crops, but very little is known about it and there have

been no tests. All species in the genus cause some degree of damage and at least 10 cause proven

major economic losses, so it is probable that the species is pathogenic under at least some

conditions.

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Radopholus arabocoffeae

Threat: Probable

Taxonomy

Name: Radopholus arabocoffeae Trinh, Nguyen, Waeyenberge, Subbotin, Karssen & Moens, 2004

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchina: Pratylenchidae

Biological characteristics

In controlled laboratory tests, this species caused major damage, but symptoms and effect under field conditions

are untested as yet.

A migratory endoparasite which lives within the host plant, moving through and feeding on internal tissues. The

rest of its life history is unknown.

Appears to be restricted to warm humid conditions.

Filter 1: Present in ASEAN-NAQS region

ASEAN region: Vietnam

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Coffea spp. (coffee)

Filter 3: Pathway present

Proven: Plants or plant parts; seedlings and nursery stock; soil

Criterion 1: Recorded in NAQS countries

No.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Caused the highest level of damage in laboratory testing compared with Pratylenchus coffeae and

Radopholus duriophilus, two other plant-parasitic nematodes of coffee (Trinh et al, 2004). There

is very little information on the impact of this pest in the field.

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Radopholus bridgei

Threat: Probable

Taxonomy

Name: Radopholus bridgei Siddiqi & Hahn, 1995

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchoidea: Pratylenchidae

Common Name: None

Biological characteristics

Limited information is available on host range and symptoms of this species. Other species of the genus to

cause necrosis on roots and allow ingress of other root pathogens, particularly fungi. Tumeric is the type host

but the species has also been intercepted on aquarium plants (Anubias sp., Echinodores sp., Ophiopogon sp.)

imported into Poland from Southeast Asia (Ryss and Karnkowski 2010).

All species from the genus are migratory endoparasites. Information on the lifecycle of this nematode is not

available.

Environmental preferences of this species are poorly known. Thus far it has been reported from Indonesia only.

Filter 1: Recorded in ASEAN or NAQS countries

NAQS region: Indonesia

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Curcuma zedoaria (Tumeric)

Filter 3: Pathway present

Proven: Soil, Plants or Plant Parts (including aquatic plants); Corms, Rhizomes and Tubers; Roots

and Root Stock,

Probable: Flower and Ornamental Plants; Sea Swept Plant Debris

Criterion 1: Recorded in NAQS countries

Yes, Indonesia

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Not known

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Radopholus citri

Threat: Greatest

Taxonomy

Name: Radopholus citri Machon & Bridge, 1996

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchoidea: Pratylenchidae

Common Name: None

Biological characteristics

Causes severely reduced growth in Citrus spp., accompanied by necrosis and root damage. There is limited

information on host range: reported from the field on citrus only, however can be cultured on excised maize

roots and carrots (Machon and Bridge, 1996).

A migratory endoparasite feeding on the roots of Citrus spp.. There is limited information available on the life

cycle of this species.

There is limited information on the environmental preferences of this species, other than its distribution seems

limited.

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN: Indonesia (Type locality is East Java)

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Citrus spp.

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts; Roots and Rootstock

Criterion 1: Recorded in NAQS countries

Yes; Indonesia.

Criterion 2: Absent from Australia

Not recorded from Australia.

Criterion 3: Pathogenicity to crops proven

Pathogenicty demonstrated to Citrus spp. in pot trials at 1000 nematodes per plant (Machon and

Bridge, 1996).

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Radopholus similis

Threat: Probable

Taxonomy Name: Radopholus similis (Cobb, 1893), Thorne 1949

Synonyms: Tylenchus similis Cobb, 1893

Anguillulina similis (Cobb) Goodey, 1932

Rotylenchus similis (Cobb) Filipjev, 1936

Tylenchus granulosus Cobb, 1893 (= senior synonym)

Anguillulina granulose (Cobb) Goodey, 1932;

Tetylenchus granulosus (Cobb) Filipjev 1936;

Radopholus granulosus (Cobb) Siddiqi, 1986

Tetylenchus acutocaudatus Zimmermann, 1898

Anguillulina acutocaudata (Zimmermann) Goodey, 1932

Tylenchorhynchus acutocaudatus (Zimmermann) Filipjev, 1934

Radopholus acutocaudatus (Zimmermann) Siddiqi, 1986

Tylenchus biformis Cobb, 1909

Anguillulina biformis (Cobb) Goodey, 1932

Radopholus biformis (Cobb) Siddiqi, 1986

Radopholus similis smilis Cobb, 1893 (Siddiqi, 1986)

Radopholus citrophilus Huettel, Dickson & Kaplan, 1984

Radopholus similis citrophilus Huettel, Dickson & Kaplan, 1984 (Siddiqi, 1986)

Subspecies: Radopholus similis similis (Cobb, 1893) Thorne, 1949 (Siddiqi, 1986)

Radopholus similis citrophilus Huettel, Dickson & Kaplan, 1984 (Siddiqi, 1986) (= citrus race of

R. similis)

Taxonomic Position: Nematoda: Chromadorea: Panagrolaimida: Tylenchoidea: Pratylenchidae

Common Name: Burrowing nematode

Biological characteristics

The type subspecies is a destructive pathogen of banana and a wide range of other crop and ornamental plants,

but not Citrus spp.. Subspecies citrophilus—also known as the Citrus race—parasitises Citrus spp. , as well as

many other plants but not banana. On banana, heavy infestation of the roots causes root-rot, weakening of the

root system and toppling of the plant, a typical symptom of this nematode. On citrus, the species causes

spreading decline, a slow loss of general health. On pepper, it causes the yellows disease. Damage to many

other crops has been recorded.

A migratory endoparasite residing and feeding in root cortex, corms and tubers. It can complete its lifecycle in

20-25 days under ideal conditions (Gowen et al., 2005).

The burrowing nematode parasitizing banana and other crops is widespread in most tropical and subtropical

areas of the world. The subspecies attacking Citrus spp. is known from Florida, USA, and pssibly a few other

locations in Central and South America. There is considerable variability within the species which is still to be

understood (Moens & Perry 2009) and there may be other races or subspecies (Gafur unpublished).

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Brunei Darussalam, Malaysia, Thailand

NAQS region: Indonesia, PNG, Solomon Islands

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Abelmoschus spp. (aibika, okra etc.), Ananas comosus (pineapple), Arachis hypogaea (peanut),

Bambusa spp. (bamboo), Beta vulgaris (beetroot), Brassica spp. (cabbage, broccoli etc.), Camellia

sinensis (tea), Capsicum spp. (chilli, capsicum etc.), Citrullus lanatus (watermelon), Citrus spp.,

Cocos nucifera (coconut), Coffea sp. (coffee), Cucurbitaceae spp. (cucumber, pumpkin, squash etc.),

Dioscorea spp., Ficus spp. (figs), Gardenia sp., Glycine max (soybean), Heliconia spp., Hibiscus

spp., Ipomoea spp., Lactuca spp., Litchi chinensis (lychee), Mangifera spp. (mango), Momordica

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charantia (bitter melon), Musa spp. (banana & plantain), Persea americana (avocado), Petroselinum

crispum (parsley), Phaseolus vulgaris (haricot bean), Pinus spp. (pine), Piper spp. (pepper), Psidium

guajava (guava), Saccharum officinarum (sugarcane), Solanum spp. (potato, aubergine, tomato etc.),

Sorghum spp. (sorghum), Theobroma cacao (cacao), Triticum spp. (wheat), Vigna spp. (bean), Zea

mays (maize), Zingiber officinale (ginger)

Filter 3: Pathway present

Proven: Soil; Plants or Plant Parts (including aquarium plants); Corms, Rhizomes and Tubers; Roots

and Root Stock; Seedlings & Nursery Stock; Flower and Ornamental Plants

Probable: Sea Swept Plant Debris

Criterion 1: Recorded in NAQS countries

Yes, Indonesia, PNG, Solomon Islands

Criterion 2: Absent from Australia

Present in Australia. There is evidence that a very restricted part of the known worldwide

variation within this species is present (Cobon & Pattison 2003, Kaplan et al. 2000, Stirling &

Pattison 2008, Tan et al. 2010).

Criterion 3: Pathogenicity to crops proven

Damages banana and a wide range of other crops such as ginger (Gowen, Queneherve & Fogain

2005): yields reduced by 40-70% (oranges), 50-80% (grapefruits) and 95% after 10 years (Esser et

al. 1988). Damage to turmeric, cardamoms, ginger and peppers has been reported in various field

locations, along with severe damage in inoculation trials on soyabeans, sorghum, maize and

sugarcane, and moderate damage to roots of aubergines, coffee, tomatoes and potatoes.

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Xiphinema americanum s.l.

Threat: Probable

Taxonomy Name: Xiphinema americanum Cobb 1913 sensu lato

Synonyms: Tylencholaimus americanus (Cobb 1913) Micoletzky 1922

Taxonomic Position: Nematoda: Dorylaimea: Dorylaimida: Dorylaimina: Xiphinematidae

Common Name: American Dagger Nematode

Biological characteristics Direct feeding causes damage to roots in many crops (McElroy, 1992). Transmits plant viruses including

tomato ringspot nepovirus (TomRSV), tobacco ringspot nepovirus (TRSV), cherry rasp leaf nepovirus (CRLV)

and peach rosette mosaic nepovirus (PRLV).

A migratory ectoparasite living on the outer surface of plants and feeding on plant roots. The lifecycle of the

nematode 1 year (Hunt 1993).

The nematode is widespread, preferring drier sandy soils and moderate temperatures (20-24ºC) but also occurs

in heavy soils and higher or lower temperatures.

Filter 1: Recorded in ASEAN or NAQS countries

ASEAN region: Philippines, Thailand

Also New Zealand, and found in Europe, East and North Asia and The Americas.

Filter 2: Common crop hosts present in ASEAN-NAQS region as well as in Australia

Allium cepa (onion), Arachis hypogaea (peanut), Bambusa spp. (bamboo), Beta vulgaris (beetroot),

Bougainvillea spp., Brassica spp. (cabbage, broccoli etc.), Capsicum spp. (chilli, capsicum etc.),

Citrus spp., Cocos nucifera (coconut), Coffea sp. (coffee), Cucurbitaceae spp. (cucumber, pumpkin,

squash etc.), Ficus spp. (figs), Gardenia sp., Glycine max (soybean), Hibiscus spp., Lactuca spp.,

Litchi chinensis (lychee), Mangifera spp. (mango), Manilkara zapota (sapodilla), Morus spp.

(mulberry), Musa spp. (banana & plantain), Nicotiana tabacum (tobacco), Oryza spp. (rice),

Passiflora spp. (passion fruit), Persea americana (avocado), Phaseolus vulgaris (haricot bean), Pinus

spp. (pine), Psidium guajava (guava), Rhododendron spp., Rosa spp. (rose), Saccharum officinarum

(sugarcane), Solanum spp. (potato, aubergine, tomato etc.), Sorghum spp. (sorghum), Triticum spp.

(wheat), Vigna spp. (bean), Vitis spp. (grape), Zea mays (maize)

Filter 3: Pathway present

Proven: Soil

Probable: Plants or Plant Parts; Corms, Rhizomes & Tubers; Bulbs; Flower & Ornamental Plants

Criterion 1: Recorded in NAQS countries

No.

Criterion 2: Absent from Australia

Widespread in Australia, but a limited number of species within the complex are known (Sturhan

et al., 1997).

Criterion 3: Pathogenicity to crops proven

Yes (Lamberti and Ciancio, 1994).

Notes

Xiphinema americanum sensu lato is a species complex of about 40 species. Xiphinema americanum sensu

stricto is a valid species, but the complex is still to be resolved taxonomically. Differences between species are

very small and intraspecific variability remains unresolved (Hunt 1993, Lamberti and Ciancio, 1994). Hence

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reports may refer to any of the species in the complex, and the complex is dealt with as a whole for quarantine

purposes (eg Lamberti and Ciancio, 1994).

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Appendix D—Nematodes recorded on host crops considered

The following lists include all nematode species that have been found on each host, not just

those recorded in the NAQS countries, ASEAN countries or Australia.

Bold underlined—greatest nematode threats

Bold—probable nematode threats.

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Abelmoschus spp. (aibika, okra etc.) Belonolaimus longicaudatus

Helicotylenchus dihystera

Helicotylenchus multicinctus

Hirschmanniella imamuri

Hirschmanniella oryzae

Hoplolaimus indicus

Meloidogyne arenaria

Meloidogyne incognita

Meloidogyne javanica

Paratrichodorus minor

Paratrichodorus porosus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Radopholus similis

Rotylenchulus reniformis

Scutellonema clathricaudatum

Allium cepa (onion) Aphelenchoides besseyi

Aphelenchoides fragariae

Aphelenchoides subtenuis

Belonolaimus longicaudatus

Ditylenchus destructor

Ditylenchus dipsaci

Ditylenchus sp.

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Longidorus elongatus

Longidorus sp.

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus scribneri

Pratylenchus thornei

Pratylenchus zeae

Rotylenchulus reniformis

Scutellonema clathricaudatum

Scutellonema sp.

Xiphinema americanum

Xiphinema index

Xiphinema pachtaicum

Zygotylenchus sp.

Ananas comosus (pineapple) Helicotylenchus dihystera

Hemicriconemoides mangiferae

Heterodera schachtii

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Longidorus sp.

Macroposthonia onoensis

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus brachyurus

Pratylenchus sp.

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema sp.

Annona reticulata (custard apple) Hoplolaimus pararobustus

Pratylenchus brachyurus

Bambusa spp. (bamboo) Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Heterodera zeae

Macroposthonia xenoplax

Merlinius brevidens

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus neglectus

Pratylenchus penetrans

Radopholus similis

Scutellonema brachyurum

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchulus semipenetrans

Xiphinema americanum

Bougainvillea spp. Helicotylenchus dihystera

Meloidogyne incognita

Meloidogyne javanica

Xiphinema americanum

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Brassica spp. (cabbage, broccoli etc.) Aphelenchoides fragariae

Belonolaimus longicaudatus

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Heterodera cruciferae

Heterodera schachtii

Heterodera trifolii

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus seinhorsti

Longidorus elongatus Longidorus sp.

Meloidogyne arenaria

Meloidogyne graminicola Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus coffeae

Pratylenchus hexincisus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus scribneri

Pratylenchus thornei

Pratylenchus vulnus

Radopholus similis Rotylenchulus parvus

Rotylenchulus reniformis

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Xiphinema americanum Xiphinema index

Xiphinema italiae

Xiphinema pachtaicum

Xiphinema sp.

Calopogonium caeruleum (Not common in study area, see p14) Helicotylenchus dihystera

Calopogonium mucunoides Helicotylenchus dihystera

Meloidogyne arenaria

Meloidogyne incognita

Pratylenchus brachyurus

Pratylenchus coffeae

Camellia sinensis (tea) Ditylenchus destructor

Helicotylenchus dihystera

Hoplolaimus pararobustus

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Meloidogyne thamesi

Paratrichodorus porosus

Pratylenchus brachyurus

Pratylenchus loosi

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus pseudopratensis

Pratylenchus sp.

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema sp.

Xiphinema sp.

Capsicum spp. (chilli, capsicum etc.) Belonolaimus longicaudatus

Ditylenchus destructor

Helicotylenchus dihystera

Hemicycliophora arenaria

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus seinhorsti

Longidorus sp.

Meloidogyne arenaria

Meloidogyne enterolobii

Meloidogyne exigua

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus scribneri

Pratylenchus thornei

Pratylenchus zeae

Radopholus similis

Rotylenchulus reniformis

Scutellonema brachyurum

Tylenchorhynchus clarus

Xiphinema americanum

Xiphinema index

Xiphinema sp.

Zygotylenchus sp.

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Carica papaya (pawpaw) Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Hoplolaimus pararobustus

Meloidogyne arenaria

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus penetrans

Pratylenchus zeae

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchulus sp.

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Celosia argentea (cock's comb) (Not common in study area, see p14) Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus penetrans

Scutellonema sp.

Citrullus lanatus (watermelon) Belonolaimus longicaudatus

Helicotylenchus dihystera

Helicotylenchus multicinctus

Hoplolaimus indicus

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus thornei

Radopholus similis

Rotylenchulus reniformis

Xiphinema pachtaicum

Xiphinema sp.

Citrus spp. Belonolaimus longicaudatus

Ditylenchus destructor

Helicotylenchus dihystera

Helicotylenchus microcephalus

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Hemicriconemoides mangiferae

Hemicriconemoides sp.

Hemicriconemoides strictathecatus

Hemicycliophora arenaria

Hemicycliophora sp.

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus pararobustus

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne exigua

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratrichodorus sp.

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus hexincisus

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus sp.

Pratylenchus thornei

Pratylenchus vulnus

Radopholus citri

Radopholus similis

Radopholus sp.

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchus robustus

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Tylenchulus semipenetrans

Tylenchulus sp.

Xiphinema americanum

Xiphinema basiri

Xiphinema brasiliense

Xiphinema brevicollum

Xiphinema index

Xiphinema insigne

Xiphinema italiae

Xiphinema pachtaicum

Xiphinema sp.

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Cocos nucifera (coconut) Helicotylenchus dihystera

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Meloidogyne incognita

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus zeae

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema sp.

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema basiri

Xiphinema insigne

Xiphinema sp.

Coffea sp. (coffee) Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hoplolaimus pararobustus

Hoplolaimus sp.

Longidorus sp.

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus loosi

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus sp.

Pratylenchus vulnus

Quinisulcius capitatus

Radopholus arabocoffeae

Radopholus duriophilus

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema italiae

Xiphinema sp.

Colocasia esculenta (taro) Aphelenchoides besseyi

Helicotylenchus dihystera

Hirschmanniella miticausa

Hirschmanniella sp.

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus coffeae

Pratylenchus penetrans

Quinisulcius acutus

Rotylenchulus reniformis

Coriandrum sativum (coriander) Meloidogyne arenaria

Meloidogyne incognita

Meloidogyne javanica

Cucurbitaceae spp. (cucumber,

pumpkin, squash etc.) Belonolaimus longicaudatus

Ditylenchus destructor

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Hemicycliophora arenaria

Heterodera glycines

Heterodera trifolii

Hoplolaimus indicus

Hoplolaimus seinhorsti

Longidorus elongatus

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne enterolobii

Meloidogyne exigua

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus vulnus

Radopholus similis

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus clarus

Xiphinema americanum

Xiphinema pachtaicum

Xiphinema rivesi

Zygotylenchus sp.

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Dimocarpus longan (longan) Hemicriconemoides mangiferae

Dioscorea spp. Aphelenchoides besseyi

Helicotylenchus dihystera

Hoplolaimus pararobustus

Longidorus sp.

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Paratrichodorus minor

Paratrichodorus porosus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus loosi

Pratylenchus penetrans

Pratylenchus pratensis

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema clathricaudatum

Scutellonema sp.

Dracaena spp. Helicotylenchus dihystera

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus coffeae

Pratylenchus penetrans

Pratylenchus vulnus

Rotylenchulus reniformis

Scutellonema brachyurum

Durio spp. (durian) Radopholus duriophilus

Radopholus sp.

Elettaria cardamomum (cardamom) Meloidogyne incognita

Meloidogyne javanica

Pratylenchus coffeae

Rotylenchulus reniformis

Ficus spp. (figs) Aphelenchoides fragariae

Helicotylenchus dihystera

Heterodera fici

Heterodera sp.

Longidorus elongatus

Longidorus sp.

Macroposthonia onoensis

Macroposthonia sp.

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus thornei

Pratylenchus vulnus

Radopholus similis

Rotylenchulus reniformis

Rotylenchulus sp.

Scutellonema brachyurum

Scutellonema sp.

Tylenchorhynchus annulatus

Tylenchorhynchus clarus

Xiphinema americanum

Xiphinema index

Xiphinema pachtaicum

Xiphinema sp.

Gardenia sp. Helicotylenchus dihystera

Meloidogyne arenaria

Meloidogyne incognita

Merlinius brevidens

Paratrichodorus porosus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus vulnus

Radopholus similis

Scutellonema brachyurum

Xiphinema americanum

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Glycine max (soybean) (Not common in study area, see p14) Aphelenchoides besseyi

Belonolaimus longicaudatus

Belonolaimus sp.

Ditylenchus destructor

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Helicotylenchus sp.

Heterodera glycines

Hoplolaimus galeatus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Meloidogyne trifoliophila

Paratrichodorus minor

Paratrichodorus porosus

Pratylenchus alleni

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus hexincisus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pseudopratensis

Pratylenchus scribneri

Pratylenchus sp.

Pratylenchus teres

Pratylenchus thornei

Pratylenchus vulnus

Pratylenchus zeae

Quinisulcius acutus

Radopholus similis

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchulus sp.

Scutellonema brachyurum

Scutellonema sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema sp.

Heliconia spp. Meloidogyne incognita

Meloidogyne javanica

Pratylenchus coffeae

Pratylenchus goodeyi

Radopholus similis

Hibiscus spp. Aphelenchoides besseyi

Aphelenchoides fragariae

Aphelenchoides ritzemabosi

Helicotylenchus dihystera

Helicotylenchus multicinctus

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Paratrichodorus minor

Pratylenchus brachyurus

Pratylenchus loosi

Pratylenchus penetrans

Pratylenchus vulnus

Radopholus similis

Rotylenchulus reniformis

Scutellonema sp.

Xiphinema americanum

Ipomoea spp. Aphelenchoides besseyi

Aphelenchoides fragariae

Belonolaimus longicaudatus

Ditylenchus destructor

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus microcephalus

Helicotylenchus multicinctus

Hoplolaimus indicus

Longidorus sp.

Macroposthonia onoensis

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Pratylenchus brachyurus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Radopholus similis

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema sp.

Tylenchorhynchus annulatus

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

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Lactuca spp. Aphelenchoides ritzemabosi

Belonolaimus longicaudatus

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Heterodera cruciferae

Longidorus elongatus

Longidorus sp.

Meloidogyne arenaria

Meloidogyne fallax

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus thornei

Pratylenchus zeae

Radopholus similis

Rotylenchulus parvus

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Trichodora cylindricus

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Xiphinema americanum

Litchi chinensis (lychee) Hemicriconemoides mangiferae

Hemicriconemoides sp.

Hemicriconemoides strictathecatus

Paratrichodorus mirzai

Paratrichodorus sp.

Radopholus similis

Rotylenchulus reniformis

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

Mangifera spp. (mango) Helicotylenchus dihystera

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hemicriconemoides sp.

Hemicriconemoides strictathecatus

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Meloidogyne incognita

Pratylenchus brachyurus

Pratylenchus loosi

Pratylenchus penetrans

Pratylenchus sp.

Pratylenchus teres

Quinisulcius acutus

Radopholus similis

Rotylenchulus reniformis

Tylenchorhynchus annulatus

Xiphinema americanum

Xiphinema basiri

Xiphinema brasiliense

Xiphinema rivesi

Xiphinema sp.

Manihot esculenta (cassava) Helicotylenchus dihystera

Hemicriconemoides mangiferae

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus brachyurus

Pratylenchus coffeae

Rotylenchulus reniformis

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus mashhoodi

Tylenchorhynchus sp.

Manilkara zapota (sapodilla) Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Xiphinema americanum

Momordica charantia (bitter melon) Helicotylenchus dihystera

Meloidogyne incognita

Pratylenchus coffeae

Radopholus similis

Rotylenchulus reniformis

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Morus spp. (mulberry) Helicotylenchus pseudorobustus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Paratrichodorus porosus

Pratylenchus coffeae

Pratylenchus penetrans

Pratylenchus vulnus

Scutellonema brachyurum

Xiphinema americanum

Xiphinema index

Xiphinema pachtaicum

Xiphinema sp.

Musa spp. (banana & plantain) Helicotylenchus dihystera

Helicotylenchus microcephalus

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Helicotylenchus sp.

Helicotylenchus varicaudatus

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Heterodera oryzae

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus coffeae

Pratylenchus goodeyi

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus sp.

Pratylenchus thornei

Pratylenchus vulnus

Pratylenchus zeae

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema minutum

Scutellonema sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus nudus

Tylenchorhynchus sp.

Tylenchulus semipenetrans

Xiphinema americanum

Xiphinema italiae

Xiphinema sp.

Nephelium lappaceum (rambutan) Meloidogyne hapla

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Nicotiana tabacum (tobacco) (Not common in study area, see p14) Aphelenchoides ritzemabosi

Ditylenchus dipsaci

Globodera sp.

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hirschmanniella oryzae

Longidorus sp.

Meloidogyne arenaria

Meloidogyne enterolobii

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Paratrichodorus minor

Paratrichodorus sp.

Paratylenchus sp.

Pratylenchus brachyurus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus sp.

Pratylenchus teres

Pratylenchus vulnus

Pratylenchus zeae

Quinisulcius acutus

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchulus sp.

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Trichodora sp.

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema rivesi

Xiphinema sp.

Oryza spp. (rice) Aphelenchoides besseyi

Ditylenchus angustus

Ditylenchus sp.

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Hemicriconemoides cocophilus

Hemicriconemoides sp.

Heterodera oryzae

Heterodera sacchari

Heterodera schachtii

Heterodera sp.

Heterodera zeae

Hirschmanniella imamuri

Hirschmanniella oryzae

Hirschmanniella sp.

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Macroposthonia onoensis

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne graminicola

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne naasi

Meloidogyne sp.

Merlinius brevidens

Paralongidorus australis

Paralongidorus sp.

Paratrichodorus minor

Paratrichodorus sp.

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus thornei

Pratylenchus zeae

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus annulatus

Tylenchorhynchus brassicae

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus mashhoodi

Tylenchorhynchus nudus

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema italiae

Xiphinema sp.

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Passiflora spp. (passion fruit) Aphelenchoides ritzemabosi

Helicotylenchus dihystera

Meloidogyne arenaria

Meloidogyne incognita

Meloidogyne javanica

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema sp.

Xiphinema americanum

Persea americana (avocado) Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Longidorus sp.

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus thornei

Pratylenchus vulnus

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema brachyurum

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchulus semipenetrans

Xiphinema americanum

Xiphinema brasiliense

Xiphinema italiae

Xiphinema sp.

Petroselinum crispum (parsley) Ditylenchus dipsaci

Heterodera schachtii

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Paratylenchus hamatus

Pratylenchus penetrans

Radopholus similis

Tylenchorhynchus clarus

Tylenchorhynchus sp.

Zygotylenchus sp.

Phaseolus vulgaris (haricot bean) Aphelenchoides ritzemabosi

Belonolaimus longicaudatus

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Heterodera glycines

Heterodera schachtii

Heterodera trifolii

Hoplolaimus galeatus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Longidorus elongatus

Longidorus sp.

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Pratylenchus brachyurus

Pratylenchus goodeyi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus scribneri

Pratylenchus thornei

Pratylenchus vulnus

Quinisulcius acutus

Radopholus similis

Rotylenchulus parvus

Rotylenchulus reniformis

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema index

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Pinus spp. (pine) Belonolaimus longicaudatus

Bursaphelenchus sp.

Bursaphelenchus xylophilus

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus sp.

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne incognita

Merlinius brevidens

Paralongidorus sp.

Paratrichodorus minor

Paratrichodorus porosus

Paratrichodorus renifer

Paratrichodorus sp.

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus crenatus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus sp.

Pratylenchus teres

Pratylenchus thornei

Pratylenchus vulnus

Radopholus similis

Radopholus sp.

Rotylenchulus parvus

Rotylenchus robustus

Rotylenchus sp.

Scutellonema brachyurum

Scutellonema sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema index

Xiphinema italiae

Xiphinema rivesi

Xiphinema sp.

Piper spp. (pepper) Helicotylenchus dihystera

Helicotylenchus varicaudatus

Hemicriconemoides mangiferae

Hoplolaimus seinhorsti

Hoplolaimus sp.

Macroposthonia onoensis

Macroposthonia sp.

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Pratylenchus coffeae

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema clathricaudatum

Scutellonema sp.

Xiphinema sp.

Psidium guajava (guava) Helicotylenchus dihystera

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hoplolaimus indicus

Hoplolaimus pararobustus

Meloidogyne arenaria

Meloidogyne enterolobii

Meloidogyne sp.

Pratylenchus pratensis

Pratylenchus zeae

Radopholus similis

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema italiae

Psophocarpus tetragonolobus (winged

bean) Meloidogyne incognita

Rotylenchulus reniformis

Rhododendron spp. (Not common in study area, see p14) Aphelenchoides fragariae

Aphelenchoides ritzemabosi

Helicotylenchus dihystera

Helicotylenchus multicinctus

Macroposthonia xenoplax

Meloidogyne incognita

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Pratylenchus crenatus

Pratylenchus penetrans

Pratylenchus vulnus

Rotylenchus robustus

Scutellonema brachyurum

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

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Rosa spp. (rose) Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hemicriconemoides mangiferae

Hemicriconemoides strictathecatus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Longidorus elongatus

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus scribneri

Pratylenchus sp.

Pratylenchus thornei

Pratylenchus vulnus

Rotylenchus robustus

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Xiphinema americanum

Xiphinema basiri

Xiphinema index

Xiphinema rivesi

Xiphinema sp.

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Saccharum officinarum (sugarcane) Achlysiella williamsi

Belonolaimus longicaudatus

Cactodera cacti

Helicotylenchus dihystera

Helicotylenchus microcephalus

Helicotylenchus pseudorobustus

Hemicriconemoides cocophilus

Hemicriconemoides mangiferae

Hemicriconemoides sp.

Hemicycliophora arenaria

Hemicycliophora sp.

Heterodera sacchari

Heterodera schachtii

Heterodera sp.

Hirschmanniella imamuri

Hirschmanniella oryzae

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Longidorus sp.

Macroposthonia onoensis

Macroposthonia sp.

Meloidogyne arenaria

Meloidogyne exigua

Meloidogyne incognita

Meloidogyne javanica

Paratrichodorus minor

Paratrichodorus mirzai

Paratrichodorus porosus

Paratrichodorus sp.

Pratylenchus brachyurus

Pratylenchus delattrei

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus scribneri

Pratylenchus sp.

Pratylenchus sudanensis

Pratylenchus teres

Pratylenchus thornei

Pratylenchus zeae

Quinisulcius acutus

Radopholus similis

Radopholus sp.

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchulus sp.

Scutellonema brachyurum

Scutellonema sp.

Tylenchorhynchus annulatus

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus mashhoodi

Tylenchorhynchus nudus

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema italiae

Xiphinema sp.

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Solanum spp. (potato, aubergine, tomato

etc.) Aphelenchoides fragariae

Aphelenchoides ritzemabosi

Belonolaimus longicaudatus

Ditylenchus destructor

Ditylenchus dipsaci

Globodera pallida

Globodera rostochiensis

Globodera sp.

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Hemicycliophora arenaria

Heterodera glycines

Heterodera oryzae

Heterodera schachtii

Heterodera trifolii

Hirschmanniella imamuri

Hirschmanniella oryzae

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus seinhorsti

Longidorus elongatus

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne enterolobii

Meloidogyne fallax

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne naasi

Meloidogyne thamesi

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus renifer

Paratylenchus hamatus

Pratylenchus alleni

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus goodeyi

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus pseudopratensis

Pratylenchus scribneri

Pratylenchus thornei

Pratylenchus vulnus

Pratylenchus zeae

Radopholus similis

Radopholus sp.

Rotylenchulus parvus

Rotylenchulus reniformis

Rotylenchus robustus

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Trichodora cylindricus

Tylenchorhynchus brassicae

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema basiri

Xiphinema index

Xiphinema pachtaicum

Xiphinema sp.

Zygotylenchus sp.

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Sorghum spp. (sorghum) Belonolaimus longicaudatus

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Heterodera avenae

Heterodera zeae

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Longidorus sp.

Macroposthonia onoensis

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne naasi

Meloidogyne sp.

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratrichodorus sp.

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus delattrei

Pratylenchus goodeyi

Pratylenchus hexincisus

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus scribneri

Pratylenchus sudanensis

Pratylenchus thornei

Pratylenchus zeae

Quinisulcius acutus

Radopholus similis

Rotylenchulus parvus

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus annulatus

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus nudus

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema rivesi

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Triticum spp. (wheat) (Not common in study area, see p14) Anguina tritici

Belonolaimus longicaudatus

Bitylenchus dubius

Ditylenchus angustus

Ditylenchus destructor

Ditylenchus dipsaci

Ditylenchus sp.

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Helicotylenchus sp.

Helicotylenchus varicaudatus

Heterodera avenae

Heterodera cruciferae

Heterodera schachtii

Heterodera sp.

Heterodera zeae

Hirschmanniella imamuri

Hirschmanniella oryzae

Hoplolaimus indicus

Hoplolaimus pararobustus

Longidorus sp.

Meloidogyne arenaria

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne naasi

Meloidogyne sp.

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus sp.

Pratylenchus brachyurus

Pratylenchus delattrei

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pinguicaudatus

Pratylenchus pratensis

Pratylenchus scribneri

Pratylenchus sp.

Pratylenchus thornei

Radopholus nativus

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Scutellonema clathricaudatum

Scutellonema sp.

Subanguina radicicola

Trichodora sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus nudus

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema pachtaicum

Xiphinema sp.

Vigna spp. (bean) Belonolaimus longicaudatus

Ditylenchus dipsaci

Helicotylenchus dihystera

Helicotylenchus multicinctus

Helicotylenchus pseudorobustus

Hemicycliophora arenaria

Heterodera glycines

Heterodera schachtii

Hoplolaimus indicus

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus thornei

Pratylenchus vulnus

Quinisulcius acutus

Radopholus similis

Rotylenchulus parvus

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Xiphinema americanum

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Vitis spp. (grape) Belonolaimus longicaudatus

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Helicotylenchus sp.

Hemicriconemoides mangiferae

Hemicriconemoides sp.

Hemicriconemoides strictathecatus

Hemicycliophora arenaria

Heterodera schachtii

Hoplolaimus pararobustus

Hoplolaimus seinhorsti

Longidorus elongatus

Longidorus sp.

Macroposthonia xenoplax

Meloidogyne arenaria

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Meloidogyne thamesi

Merlinius brevidens

Neodolichodorus sp.

Paratrichodorus minor

Paratrichodorus porosus

Paratrichodorus renifer

Paratrichodorus sp.

Paratylenchus hamatus

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus goodeyi

Pratylenchus hexincisus

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus pratensis

Pratylenchus scribneri

Pratylenchus sp.

Pratylenchus thornei

Pratylenchus vulnus

Quinisulcius acutus

Rotylenchulus reniformis

Rotylenchulus sp.

Rotylenchus robustus

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus mashhoodi

Tylenchulus semipenetrans

Xiphinema americanum

Xiphinema brasiliense

Xiphinema brevicollum

Xiphinema index

Xiphinema insigne

Xiphinema italiae

Xiphinema pachtaicum

Xiphinema rivesi

Xiphinema sp.

Xiphinema vuittenezi

Zygotylenchus sp.

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Xanthosoma spp. Helicotylenchus dihystera

Meloidogyne incognita

Meloidogyne javanica

Pratylenchus coffeae

Rotylenchulus reniformis

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Zea mays (maize) Aphelenchoides besseyi

Belonolaimus longicaudatus

Ditylenchus destructor

Ditylenchus dipsaci

Dolichodorus sp.

Helicotylenchus dihystera

Helicotylenchus pseudorobustus

Hemicriconemoides mangiferae

Hemicycliophora sp.

Heterodera avenae

Heterodera glycines

Heterodera oryzae

Heterodera schachtii

Heterodera sp.

Heterodera zeae

Hirschmanniella imamuri

Hirschmanniella oryzae

Hoplolaimus galeatus

Hoplolaimus indicus

Hoplolaimus seinhorsti

Hoplolaimus sp.

Longidorus elongatus

Longidorus sp.

Meloidogyne arenaria

Meloidogyne graminicola

Meloidogyne hapla

Meloidogyne incognita

Meloidogyne javanica

Merlinius brevidens

Paratrichodorus minor

Paratrichodorus porosus

Paratrichodorus sp.

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus crenatus

Pratylenchus delattrei

Pratylenchus goodeyi

Pratylenchus hexincisus

Pratylenchus loosi

Pratylenchus neglectus

Pratylenchus penetrans

Pratylenchus scribneri

Pratylenchus sp.

Pratylenchus thornei

Pratylenchus vulnus

Pratylenchus zeae

Quinisulcius acutus

Quinisulcius capitatus

Quinisulcius curvus

Radopholus similis

Radopholus sp.

Rotylenchulus parvus

Rotylenchulus reniformis

Scutellonema brachyurum

Scutellonema clathricaudatum

Scutellonema sp.

Trichodora sp.

Tylenchorhynchus clarus

Tylenchorhynchus claytoni

Tylenchorhynchus mashhoodi

Tylenchorhynchus sp.

Xiphinema americanum

Xiphinema pachtaicum

Xiphinema sp.

Zygotylenchus sp.

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Zingiber officinale (ginger) Helicotylenchus dihystera

Meloidogyne arenaria

Meloidogyne incognita

Meloidogyne javanica

Meloidogyne sp.

Meloidogyne thailandica

Pratylenchus brachyurus

Pratylenchus coffeae

Pratylenchus flakkensis

Pratylenchus pratensis

Pratylenchus zeae

Radopholus similis

Radopholus sp.

Rotylenchulus reniformis

Tylenchorhynchus mashhoodi

Tylenchorhynchus sp.

Xiphinema sp.