Diagnostic Protocol

Rice water weevil

Lissorhoptrus oryzophilus

Information prepared by Rolf Oberprierler & Ben Boyd CSIRO Entomology, Canberra ACT, 2601

December 2004 (reviewed 2008)

The scientific and technical content of this document is current to the date published and all efforts were made to obtain relevant and published information on the pest. New information will be included as it becomes available, or when the document is

reviewed. The material contained in this publication is produced for general information only. It is not intended as professional

advice on any particular matter. No person should act or fail to act on the basis of any material contained in this publication without first obtaining specific, independent professional advice. Plant Health Australia and all persons acting for Plant Health

Australia in preparing this publication, expressly disclaim all and any liability to any persons in respect of anything done by any

such person in reliance, whether in whole or in part, on this publication. The views expressed in this publication are not

necessarily those of Plant Health Australia.

Introduction

(Coleoptera: : Erirhininae)

The name “Rice Water Weevil” applies both to a particular species of weevil, Lissorhoptrus oryzophilus Kuschel, which is one of the most serious pests of cultivated rice (Oryza sativa) in the world, and to a larger group of similar weevils that live on aquatic grasses in the New World and are real or potential rice pests as well. While this protocol deals specifically with L. oryzophilus, the diagnostic key makes provision also for recognising other relevant rice water weevils at group and genus level, so as to facilitate their detection at ports of entry in Australia and the taking of appropriate steps to prevent them from establishing in the country.

Rice Water Weevil (RWW) belongs to the subfamily Erirhininae, a small cosmopolitan group of weevils generally associated with monocotyledonous plants in aquatic and semi-aquatic habitats. The definition, composition and classification of the Erirhininae are poorly understood, and considerable taxonomic expertise is often required to recognise these weevils. The group is not very diverse in Australia but includes a few large (but poorly known) genera, such as Echinocnemus and Aonychus. The of the Australian erirhinine fauna was revised by Zimmerman (1993), whose work should be consulted for details about the various species and their distribution. Echinocnemus occurs

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If you would like more information, or to download a copy of the Industry Biosecurity Plan, visit www.planthealthaustralia.com.au, email [email protected] or phone (02) 6260 4322. PLANT HEALTH AUSTRALIA | Rice Industry Biosecurity Plan 2009 on aquatic grasses in the Old World and includes rice pests in Africa (E. gemellus Marshall, E. obscurus Hustache), India (E. oryzae Marshall) and Japan (E. bipunctatus Roelofs, E. squameus (Billberg)), but the biology and hosts of the Australian species are not recorded. In Africa a similar weevil, Afroryzophilus djibai Lyal, damages rice in Senegal (Lyal 1992). Along with other weevils injurious to rice, Echinocnemus and Afroryzophilus are referred to as “rice weevils”, rather than “rice water weevils”.

In the New World, a particular group of Erirhininae, the tribe Stenopelmini (sometimes incorrectly referred to as Hydronomini, e.g. Zimmerman 1993), is very diverse and important, including both serious agricultural pests and highly successful weed biocontrol agents. These weevils are characterised by having 6 (rather than 7) segments in the antennal funicle (see diagnostic key) and mostly by their body being covered by a more or less continuous, smooth layer of fused (agglutinated) scales. In Australia there are three native genera currently classified in Stenopelmini (Athor, Baeosomus, Niphobolus), and four South American species (Cyrtobagous salviniae Calder & Sands, Neochetina bruchi Hustache, N. eichhorniae Warner, Neohydronomus affinis Hustache) introduced as biocontrol agents of invasive aquatic weeds. None of these species are known to attack rice.

Within the Stenopelmini, a group of eight New World genera (Bagoidellus, Bagoidus, Helodytes, Hydrotimetes, Ilyodytes, Lissorhoptrus, Neobagoidus, Oryzophagus) are collectively known as “rice water weevils”. These genera are classified in a subtribe Lissorhoptrina and characterised by an asymmetrical antennal club, narrow tarsi and long swimming hairs on the middle tibiae (see Hix et al. 2000 for details of swimming behaviour). As far as their hosts are known, most of them feed on aquatic grasses, with Lissorhoptrus and Oryzophagus specifically recorded also from rice. No native Lissorhoptrina occur in Australia, and no exotic species are known to have been introduced.

Lissorhoptrus includes the “rice water weevils” in the strict sense. The genus occurs in both North and South America and currently comprises 19 species, but several undescribed ones are known and a revision of the North American species is in progress by C. W. O‟Brien. At least eight species are thus far recorded as attacking rice: L. oryzophilus Kuschel and L. simplex (Say) in North America, L. erratilis Kuschel, L. isthmicus Kuschel, L. kuscheli O‟Brien and L. persimilis O‟Brien in Central America, and L. bosqi Kuschel and L. tibialis (Hustache) in South America. Species catalogues of Lissorhoptrus are provided by O‟Brien & Wibmer (1982) Wibmer & O‟Brien (1986), O‟Brien & Anderson (1996) and Morrone & O‟Brien (1999).

Lissorhoptrus oryzophilus is the most serious of these rice pests and has spread to other rice-growing areas of the world, in particular Central America and eastern Asia (India, China, Korea, Japan). Its distinction from other Lissorhoptrus species is difficult and relies on subtle differences in the genitalia and metatibial spines of the males. Identification keys to Lissorhoptrus species have been constructed by Kuschel (1952) and O‟Brien (1996), but neither is comprehensive and able to distinguish L. oryzophilus and other rice pests from various undescribed species of the genus that have become known since. Lissorhoptrus oryzophilus had been confused with the very similar L. simplex in the earlier literature, until Kuschel (1952) recognised and described it as a distinct species. References to L. simplex prior to 1952 mostly relate to L. oryzophilus.

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For information about distribution, hosts, biology and symptoms of damage of RWW, see Pest Risk Analysis by Stevens (2004) (Rice Industry Biosecurity Plan - Appendix 1) and http://www.ipm.ucdavis.edu/PMG/r682300511.html.

For technical terms relating to diagnostic characters, see Glossary.

Diagnostic images of RWW

Figures 1-6: adult Lissorhoptrus oryzophilus Kuschel; dorsal, lateral and ventral view, abdominal ventrites, mesotibial swimming hairs, tridentate metatibial mucro of male.

Images of the larva, pupal cocoon and larval feeding scars on rice leaves are available at http://www.ipm.ucdavis.edu/PMG/r682300511.html.

Organisms that occur in Australia with which RWW might be confused

Although RWW is readily distinguishable from all other weevils in Australia by its flattened, curved middle tibiae fringed with long swimming hairs, these hairs are easily abraded, making identification of RWW difficult when specimens are damaged in this or other features, or generally in poor condition.

No identification keys are available to all Australian weevils (ca. 4200 described species), but since RWW lives in aquatic habitats, its distinction is only relevant from a relatively small number of weevils that live in similar environments in Australia. Four groups of such weevils occur in Australia:

Listroderini (Figures 7 -14) This group includes three described native genera, Anorthorhinus, Ethemaia and Steriphus, and two introduced species of Listronotus, L. bonariensis (Kuschel) and L. setosipennis (Hustache), the former of which mines in stems of various grasses and the latter is employed in the biocontrol of the pasture weed Parthenium hysterophorus. All these Australian Listroderini are distinguishable from RWW by having several fine longitudinal carinae (keels) on the rostrum, the scales on the pronotum and elytra separate, not agglutinated, and the ventral side covered with setae only, not scales (at most a few scales laterally). Their larvae feed on the roots, stems and leaves of a variety of plants but are not aquatic (lack spine-like spiracles for tapping air from submerged plant parts, see May 1994).

Bagoini (Figures 15-16) This small weevil group includes only one genus in Australia, Bagous, which was recently taxonomically revised (O‟Brien & Askevold 1992) and currently comprises 27 described species in Australia. Its species are superficially very similar to RWW, with a similar coating of agglutinated scales, but can readily be distinguished by the broad prosternal canal in front of the fore coxae, into

Diagnostic protocol: Rice water weevil | PAGE 3 PLANT HEALTH AUSTRALIA | Rice Industry Biosecurity Plan 2009 which the rostrum is retracted. Other differences include the setose antennal club, a 7-segmented antennal funicle (often seemingly 6-segmented, the apical segment being enlarged and closely fitted to the club), the elytra having a prominent protuberance on the declivity, and the middle and hind tibiae being cylindrical and sharply bent inwards at the apex. Bagous larvae feed on various aquatic plants, such as Azolla, Blyxa, Hydrilla, Maundia, Nymphoides, Utricularia (O‟Brien & Askevold 1992), but lack modified spiracles for breathing under water (May 1994).

Erirhinini (Figures 17-20) This groups includes Bagoopsis, Echinocnemus, Tadius and several undescribed genera in Australia and is distinguishable from RWW by a 7-segmented antennal funicle and the third tarsal segment being bilobed. Bagoopsis furthermore has stiff setae on the elytra and a longer, thinner rostrum than RWW, Echinocnemus has non-agglutinated scales and Tadius a prosternal canal into which the rostrum can retract. Echinocnemus larvae feed on roots of semi-aquatic grasses and are not adapted for breathing under water (lack spine-like spiracles, May 1994). The larvae of Tadius tunnel in orchids and similar plants but are undescribed, and the larvae and biology of Bagoopsis are unknown.

Stenopelmini: Stenopelmina (Figures 21-30) As currently classified, the Stenopelmina include three native genera (Athor, Baeosomus, Niphobolus) in Australia and four introduced species of the South American genera Cyrtobagous, Neochetina and Neohydronomus. They are distinguishable from RWW mainly by having a bilobed third tarsal segment (except Cyrtobagous), the basal segment of the antennal club setose (except Cyrtobagous) and the middle tibiae cylindrical and without swimming hairs. Cyrtobagous differs from RWW in having no vestiture (scales or setae) on the dorsal body surface. The larvae of the native genera are unknown, but those of the introduced taxa are properly aquatic, with external, piercing spiracles for tapping air from their hostplants under water (May 1994). The larva of Cyrtobagous (May 1994) is very similar to that of RWW (Lee & Morimoto 1988).

Diagnostic images of Australian weevils likely to be confused with RWW

Figures 7-14: Listroderini, dorsal and lateral views: Anorthorhinus pictipes Blackburn, (Kuschel), Listronotus setosipennis (Hustache), Steriphus caudatus (Pascoe). Figures 15-16: Bagoini, dorsal and lateral views: Bagous australasiae Blackburn. Figures 17-20: Erirhinini, dorsal and lateral views: Echinocnemus sp., Tadius erirhinoides Pascoe. Figures 21-30: Stenopelmini: Stenopelmina, dorsal and lateral views: Baeosomus sp., Cyrtobagous salviniae Calder & Sands, Neochetina eichhorniae Warner, Neohydronomus affinis Hustache, Niphobolus deceptor Blackburn Figures 31-36: Stenopelmini: Lissorhoptrina, dorsal and lateral views: Helodytes litus Kuschel, Neobagoidus carlsoni O‟Brien, Oryzophagus oryzae (Costa-Lima) [these species are not known to occur in Australia, but are very similar to RWW and also rice pests in South America]

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Comparative images of symptoms/damage/trauma caused by target organism

See RWW Pest Risk Analysis by Stevens (2004).

Sampling technique

A. HOW TO COLLECT RWW Hand-pick adults and larvae using fine forceps or similar implement.

B. NUMBER OF SPECIMENS TO BE COLLECTED Up to 10, including both small and large specimens. Smaller ones tend to be males, as are needed for species identification.

C. PREFERRED STAGE TO BE COLLECTED Adults. If no adults are available, larvae can also be collected, although their identification to genus level is less secure and to species currently impossible.

D. HOW TO PRESERVE RWW Preserve both adults and larvae in small vials containing 70-80% ethanol, adding a small paper label with collecting details (location, date, host, etc.) written in fine pencil.

E. HOW TO TRANSPORT RWW Tightly seal vials containing ethanol, place in sealable plastic bag and pack with cushioning material in a strong carton or wooden box for mailing.

F. HOW TO COLLECT PLANT SAMPLE IF REQUIRED Not applicable, as unsuitable for proper identification. If only leaves with damage symptoms are available and no adult weevils or larvae can be located, the leaf parts with damage can be cut off. Entire rice plants including roots may also be collected.

G. HOW TO PRESERVE PLANT SAMPLE Place leaf sample in small vials containing 70-80% ethanol, or keep refrigerated in sealable plastic bag. Place entire rice plants in suitable tight-sealing plastic container and refrigerate.

H. HOW TO TRANSPORT PLANT SAMPLE As in e., or flat between sheets of moist newspaper if rapidly deliverable.

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Diagnostic key to RWW in Australia

Note: Only described weevils similar in external appearance AND living in aquatic and semi-aquatic habitats are included in this key. For illustrations of diagnostic characters, see interactive key as supplied with this protocol.

1. Body ventrally with only setae (at most a few scales laterally), dorsal scales distinct, not fused; rostrum dorsally with 3 or more fine longitudinal carinae (tribe Listroderini) ………………………………………………………………. Anorthorhinus, Ethemaia, Listronotus, Steriphus - Body ventrally and dorsally with scales, which are mostly agglutinated (fused to form a smooth covering); rostrum without any carinae ………………………………………………………………2. 2. Prosternum forming a broad canal in front of procoxae, into which rostrum can be retracted (tribe Bagoini) …………………………………………………………………………………………. Bagous

- Prosternum not forming a canal in front of procoxae, or (in Tadius) canal narrow and extending to behind procoxae (subfamily Erirhininae) ………………………………………………….. 3.

3. Funicle of antenna with 7 segments (tribe Erirhinini)…..Bagoopsis, Echinocnemus, Tadius

- Funicle with 6 segments (tribe Stenopelmini) ……………………………………………………………….. 4.

4. Antennal club with basal segment setose (except Cyrtobagous), symmetrical; tibiae without long swimming hairs; tarsi not compressed, segment 3 bilobed (except Cyrtobagous) (subtribe Stenopelmina)……………………………………………………………………………… ……………………Athor, Baeosomus, Niphobolus, Cyrtobagous, Neochetina, Neohydronomus

- Antennal club with basal segment large, glabrous, asymmetrical in dorsal view; mesotibia with fringe of long swimming hairs on outer and inner edges (fig. 5); tarsi compressed, segment 3 not bilobed (subtribe Lissorhoptrina)………………………………………..5

5. Elytral striae open, ill-defined, punctures large and separate (figs. 31, 32) (Helodytes group) …………………………….... Bagoidellus, Bagoidus, Helodytes, Hydrotimetes, Ilyodytes

- Elytral striae sulciform, punctures small and confluent (figs. 1-2, 33-36) (Lissorhoptrus group) …………………………………………………………………………………………………………………………………6.

6. Scales agglutinated on all abdominal ventrites and mostly on prosternum……………………… ……………………………………………………………………………………………….. Neobagoidus, Oryzophagus

- Scales on abdominal ventrites 3 and 4 and on prosternum loose, not agglutinated (figs. 3-4) (Lissorhoptrus) ………………………………………………………………………………………………………… 7.

7. Metatibial apex in male with large, tridentate mucro (fig. 6) ……………………………………………. ……………………………………………………………Lissorhoptrus oryzophilus, L. kuscheli, L. persimilis

- Metatibial apex in male with single or bidentate mucro………………other described species

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Preparedness for specialist emergency diagnosis a. Proposed methodology for identification procedure for confirmation of RWW Identification of specimens suspected to be RWW should be attempted using the printed or electronic identification key to RWW provided with this protocol. Definitive identification of RWW requires examination and careful comparison of male genitalia and metatibial spines with authentically determined voucher material, and/or illustrations as provided by O‟Brien (1996). Proficiency and experience with the preparation, identification and interpretation of the relevant structures, and with weevil identification in general, is required in this process.

If such expertise is not available, or if specimens suspected to be RWW are too poorly preserved to permit use of the identification key provided with this protocol, the specimens can be sent to the Identification and Advice Service (IAS) of the Australian National Collection (ANIC), CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, for confirmation. Consignments should be marked as such and labelled as for urgent attention by Dr. R. G. Oberprieler. They will be attended to as a matter of high priority. If Dr. Oberprieler is unavailable, Mr Tom Weir and/or Dr. Adam Slipinski will be able to confirm identification at genus level. No charge will be made for a determination of whether a particular specimen is or is not Lissorhoptrus oryzophilus. Further identification of a specimen that is not RWW to genus level will also be provided free of charge, but identification to species of such specimen(s) may attract a standard IAS charge. b. Commitment The Australian National Insect Collection has acquired authentically determined adult, larval and pupal specimens of RWW, as well as adults of other species of Lissorhoptrus and related genera of Lissorhoptrina and other Stenopelmini. These specimens are available for verification and definite identification of RWW in Australia at any time. Copies of all pertinent literature relating to diagnostic features of RWW are also available in the ANIC for emergency diagnosis.

Positive identification or confirmation of RWW from Australia will be immediately communicated to appropriate authorities within AFFA, featuring details of the overseas source of sample (if known), reference to this protocol for images of the pest, level of confidence in identity, and date of confirmed identification. Vouchers of all samples of RWW or similar species submitted for identification, or for verification of identification, will be deposited in the ANIC for future reference.

Glossary of terms agglutinated – scales fused into a varnish-like covering bilobed – consisting of two lobes carina (pl. carinae) – fine keel declivity – the posterior declining surface of the elytra elytron (pl. elytra) – front wing of modified into hard shield covering the hind wing

Diagnostic protocol: Rice water weevil | PAGE 7 PLANT HEALTH AUSTRALIA | Rice Industry Biosecurity Plan 2009 funicle (pl. funicles) – middle part of weevil antenna (between basal scape and apical club) consisting of 4-7 segments mesotibia (pl. mesotibiae) – middle tibia metatibia (pl. metatibiae) – hind tibia mucro (pl. mucrones) – spine-like process at apex of tibia procoxa (pl. procoxae) – front coxa (basal leg segment) pronotum (pl. pronota) – dorsal part of prothorax (the distinct segment between head and elytra in beetles) prosternum (pl. prosterna) – ventral part of prothorax protuberance – knob-like swelling rostrum (pl. rostra, rostrums) – the characteristic “snout” of weevils and some other setose – covered with setae seta (pl. setae) – hair in insects spiracle (pl. spiracles) – breathing aperture in insects stria (pl. striae) – row of puntures on elytra sulciform – in the shape of a fine groove tarsus (pl. tarsi) – foot of insects (distal leg segment) taxon (pl. taxa) – unit in taxonomy, i.e. species, genus, tribe, family, etc. tibia (pl. tibiae) – shin segment of the insect leg (between femur and tarsus) tridentate – three-toothed ventrite (pl. ventrites) – ventral segment, mainly of the abdomen vestiture – clothing of setae, scales or other structures covering the body

References

Hix, R. L., Johnson, D. T., & Bernhardt, J. L. 2000. Swimming behavior of an aquatic weevil, Lissorhoptrus oryzophilus (Coleoptera: Curculionidae). Florida Entomologist 83(3): 316-324.

Kuschel, G. 1952. Revisión de Lissorhoptrus Leconte y generos vecinos de America. Revista chilena de entomologia 1: 23-74.

Lee, C.-Y., & Morimoto, K. 1988. Larvae of the weevil family Curculionidae of Japan. Part 2. Hyperinae to Cioninae (Insecta: Coleoptera). Journal of the Faculty of Agriculture, Kyushu University 33(1-2): 131-152.

Lyal, C. H. C. 1990. A new genus and species of rice weevil from the Sahel (Coleoptera: Curculionidae: Erirhininae). Bulletin of Entomological Research 80: 183-189.

May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. pp. 365-728. In: E. C. Zimmerman‟s „Australian Weevils (Coleoptera: Curculionoidea). Volume II – Brentidae, Eurhynchidae, Apionidae and a chapter on Immature Stages by Brenda May‟. CSIRO, Melbourne.

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Morrone, J. J., & O‟Brien, C. W. 1999. The aquatic and semi-aquatic weevils (Coleoptera: Curculionidae, Dryophthoridae and ) of Argentina, with indication of their host plants. Physis (Buenos Aires), Sec. C., 57(132-133): 25-37.

O‟Brien, C. W. 1996. Two new Lissorhoptrus rice pests in northern South America, with a review of the species in Colombia and Venezuela. Transactions of the American entomological Society 122(2-3): 115-134.

O‟Brien, C. W., & Anderson, D. M. 1996. A Catalogue of the Coleoptera of America north of Mexico. Family Curculionidae, subfamily Erirhininae. USDA Agriculture Handbook No. 529-143f.

O‟Brien, C. W., & Askevold, I. S. 1992. Systematics and evolution of weevils of the genus Bagous Germar (Coleoptera: Curculionidae), I. Species of Australia. Transactions of the American Entomological Society 118(4): 331-452.

O‟Brien, C. W., & Wibmer, G. J. 1982. Annotated checklist of the weevils (Curculionidae sensu lato) of South America (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute 39: 1- 563.

Stevens, M. 2004 (unpubl.) Pest Risk Assessment for Rice Water Weevil, Lissorhoptrus oryzophilus, in Australia.

Wibmer, G. J., & O‟Brien, C. W. 1986. Annotated checklist of the weevils (Curculionidae sensu lato) of North America, Central America, and the West Indies (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute 34: 1-382.

Zimmerman, E. C. 1993. Australian Weevils (Coleoptera: Curculionoidea). Volume III – Nanophyidae, Rhynchophoridae, Erirhinidae, Curculionidae: Amycterinae, Literature consulted. CSIRO, Melbourne.

Websites with information related to identification of RWW http://www.ipm.ucdavis.edu/PMG/r682300511.html http://www.fcla.edu/FlaEnt/fe83p316.htm http://www.agnet.org/library/article/nc122c.html http://www.agnet.org/library/article/eb498.html http://aesrg.tamu.edu/Rice/RiceWeevil.htm http://entowww.tamu.edu/fieldguide/bimg205.html http://www.zo.utexas.edu/faculty/sjasper/beetles/biba-k.html http://www.inhs.uiuc.edu/biod/waterbeetles/bibliography/t_references.html http://cipm.ncsu.edu/cropprofiles/docs/msrice.html http://www.fsca-dpi.org/Coleoptera/Mike/curculio.htm

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Illustrations

All photo credits: Ben Boyd (copyright CSIRO Entomology)

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Figure 1. Lissorhoptrus oryzophilus Kuschel Figure 2. Lissorhoptrus oryzophilus Kuschel (lateral (dorsal view) view)

Figure 3. Lissorhoptrus oryzophilus Kuschel Figure 4. Lissorhoptrus oryzophilus Kuschel (ventral view) (abdominal ventrites showing non-agglutinated scales on ventrites 3 and 4)

Figure 5. Lissorhoptrus oryzophilus Kuschel Figure 6. Lissorhoptrus oryzophilus Kuschel (mesotibia showing swimming hairs) (tridentate metatibial mucro of male)

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Figure 7. Anorthorhinus pictipes Blackburn Figure 8. Anorthorhinus pictipes Blackburn (lateral (dorsal view) view)

Figure 9. Listronotus bonariensis (Kuschel) Figure 10. Listronotus bonariensis (Kuschel) (lateral (dorsal view) view)

Figure 11. Listronotus setosipennis (Hustache) Figure 12. Listronotus setosipennis (Hustache) (dorsal view) (lateral view)

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Figure 13. Steriphus caudatus (Pascoe) (dorsal Figure 14. Steriphus caudatus (Pascoe) (lateral view) view)

Figure 15. Bagous australasiae Blackburn Figure 16. Bagous australasiae Blackburn (lateral (dorsal view) view)

Figure 17. Echinocnemus sp. (dorsal view) Figure 18. Echinocnemus sp. (lateral view)

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Figure 19. Tadius erirhinoides Pascoe (dorsal Figure 20. Tadius erirhinoides Pascoe (lateral view) view)

Figure 21. Baeosomus sp. (dorsal view) Figure 22. Baeosomus sp. (lateral view)

Figure 23. Cyrtobagous salviniae Calder & Figure 24. Cyrtobagous salviniae Calder & Sands Sands (dorsal view) (lateral view)

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Figure 25. Neochetina eichhorniae Warner Figure 26. Neochetina eichhorniae Warner (lateral (dorsal view) view)

Figure 27. Neohydronomus affinis Hustache Figure 28. Neohydronomus affinis Hustache (lateral (dorsal view) view)

Figure 29. Niphobolus deceptor Blackburn Figure 30. Niphobolus deceptor Blackburn (lateral (dorsal view) view)

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Figure 31. Helodytes litus Kuschel (dorsal view)

Figure 32. Helodytes litus Kuschel (lateral view)

Figure 33. Neobagoidus carlsoni O’Brien (dorsal view) Figure 34. Neobagoidus carlsoni O’Brien (lateral view)

Figure 35. Oryzophagus oryzae (Costa-Lima) Figure 36. Oryzophagus oryzae (Costa-Lima) (dorsal view) (lateral view)

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