A New Detection of the Invasive Mexican Rice Borer (Lepidoptera: Crambidae) From Georgia in the United States Based on Morphological and Molecular ...

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A New Detection of the Invasive Mexican Rice Borer (Lepidoptera: Crambidae) From Georgia in the United States Based on Morphological and Molecular ...
Journal of Integrated Pest Management, (2022) 13(1): 17; 1–5
https://doi.org/10.1093/jipm/pmac014
Brief Communication

A New Detection of the Invasive Mexican Rice Borer
(Lepidoptera: Crambidae) From Georgia in the United
States Based on Morphological and Molecular Data
Ilgoo Kang,1 Blake Wilson,2,4, Blake Carter,3 and Rodrigo Diaz1
1
 Department of Entomology, Louisiana State University Agricultural Center, 404 Life Sciences Building, Baton Rouge, LA, USA,
2
 Louisiana State University AgCenter, Sugar Research Station, Saint Gabriel, LA, USA, 3University of Georgia Extension Service,

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Effingham County, Rincon, GA, USA, and 4Corresponding author, e-mail: bwilson@agcenter.lsu.edu

Subject Editor: Thomas Kuhar

Received 7 February 2022; Editorial decision 12 March 2022

Abstract
The Mexican rice borer, Eoreuma loftini (Dyar), is an invasive herbivore that attacks many gramineous host plants.
The species is an economic pest of several grass crops in North America including sugarcane (Saccharum spp.), rice
(Oryza sativa), corn (Zea mays), and sorghum (Sorghum bicolor).The species was first detected in the United States
in 1980 and has since expanded its range eastward along the Gulf Coast reaching Louisiana in 2008. A disjointed
introduction was detected in Florida in 2012, though range expansion of this population within the state has been
limited. Most recently, a separate introduction was detected along the Atlantic coast of Georgia near the South
Carolina border (32.19884° N, 81.35894° W). Larvae were collected from sugarcane in 2020 and 2021 and identified
as E. loftini.The identification was confirmed with genetic analysis showing >99% similarity with most mitochondrial
cytochrome c oxidase subunit 1 (COI) sequences of known E. loftini. This is the furthest north and east of any
established population in North America. Potential routes of introduction and impacts to crop production along the
U.S. Atlantic Coast are discussed.

Key words: Eoreuma loftini, invasive species, range expansion, Saccharum spp., sugarcane

The Mexican rice borer, Eoreuma loftini (Dyar) (Fig. 1), is an in-                       1984; Fig. 2). Initial detection of E. loftini in the Rio Grande Valley
vasive crop pest that has been expanding its range northward and                         of Texas occurred in 1980 after numerous interceptions of larvae
eastward into the United States. The species is an economic threat to                    in plant material at U.S. ports of entry. While establishment in the
production of grass crops including sugarcane (Saccharum spp.), rice                     region is often attributed to introduction of infested plant material,
(Oryza sativa), corn (Zea mays), and sorghum (Sorghum bicolor).                          evidence suggests the expansion may have occurred unaided. The
Invasion of E. loftini may also have ecological consequences as the                      species was known to be present in the Mexican border states of
stem borer also attacks a broad range of non-crop grasses. The first                     Nuevo Leon and Tamaulipas in the late 1970s, and many of the
record of E. loftini in the U.S. state of Georgia is reported herein.                    border interceptions occurred from plant material that originated
                                                                                         within a few km of the U.S. border (Johnson 1984). Further, the
                                                                                         initial detection in Texas occurred when severe larval infestations
Monitoring and Spread of E. loftini Along the                                            were observed in sugarcane, suggesting that populations were likely
Gulf Coast                                                                               building in the area for some time prior to discovery.
The native range of E. loftini is thought to be western Mexico north                         Following its initial detection in Hidalgo County, Texas, E. loftini
to Arizona and southern California where its presence was noted in                       expanded its range in a northeastern direction at an average rate
association with sugarcane (Dyar 1917, Morill 1925), rice, and nu-                       of 23.0 km/year, reaching the westernmost edge of the Texas rice
merous non-crop grasses (Osborn and Phillips 1946). Although the                         production area in 1989 (Browning 1989, Reay-Jones et al. 2007).
crops that serve as hosts have not been produced in this region for                      Development of pheromone traps capable of detecting male E. loftini
decades, museum specimens reared from larvae collected from non-                         even at low population densities in the early 1990s greatly improved
crop hosts indicate the species remains present in the region (Univ.                     monitoring efforts (Shaver et al. 1990, 1991). Range expansion con-
Cal. Riverside, Entomol. Res. Mus., personal communication). The                         tinued with the leading edge of the invasion moving approximately
species is thought to have spread to eastern Mexico in the 1960s and                     16.5 km/year from 2000 to 2005 (Reay-Jones et al. 2007) reaching
1970s following the expansion of sugarcane to the region (Johnson                        Louisiana in 2008 (Hummel et al. 2010). The eastward spread

© The Author(s) 2022. Published by Oxford University Press on behalf of Entomological Society of America.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/),         1
which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.
A New Detection of the Invasive Mexican Rice Borer (Lepidoptera: Crambidae) From Georgia in the United States Based on Morphological and Molecular ...
2                                                                                 Journal of Integrated Pest Management, 2022, Vol. 13, No. 1

                                                                                through southwestern Louisiana from 2008 to 2015 occurred at a
                                                                                rate of 17.9 km/year (Wilson et al. 2015, 2017a). By 2020, the insect
                                                                                had spread to 16 parishes in Louisiana with expansion continuing
                                                                                predictably into key crop-producing regions of the state (Wilson and
                                                                                Huval 2020). Thus, monitoring efforts indicate that range expan-
                                                                                sion in the Unites States occurred at consistent rates, presumably
                                                                                unaided, for 40 years.
                                                                                    The first evidence of disjunct range expansion occurred in 2012
                                                                                when E. loftini was detected in a light trap in Levy County in central
                                                                                Florida (Hayden 2012). The source of this population and mech-
                                                                                anism of spread is unknown, but the establishment in Florida ap-
                                                                                pears to have been the result of a single isolated introduction. Initial
                                                                                distribution and population density in Florida were relatively limited
                                                                                indicating the detection occurred not long after establishment. The
                                                                                population has since begun to expand towards the southeast, though

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                                                                                this expansion has occurred at slower rates than those observed in
                                                                                Texas and Louisiana (VanWeelden et al. 2019). Pheromone trap
                                                                                detections were reported from Marion County (2013), Citrus and
                                                                                Sumter Counties (2014), and Lake County (2019). Expansion in
                                                                                Florida has remained slow. No northward spread has been detected
                                                                                since 2014, and expansion southward was only 14 km from 2015
                                                                                to 2019 (J. Beuzelin, personal communication). Currently, E. loftini
                                                                                is not an economic pest in Florida as the established population re-
                                                                                mains more than 160 km away from areas of commercial production
                                                                                of susceptible grass crops. However, expansion into the Everglades
                                                                                Agricultural Area where sugarcane, rice, and sweet corn are widely
                                                                                cultivated could have large economic impacts (Fig. 3).
                                                                                    No active monitoring programs are currently conducted outside
Fig. 1. Mexican rice borer, Eoreuma loftini (Dyar) (Lepidoptera: Crambidae),    of Louisiana and Florida. Expansion northward and westward in
larva feeding in a sugarcane stalk (top) and adult on a leaf (bottom).
                                                                                Texas may be occurring simultaneously with eastward expansion

Fig. 2. Geographical range expansion of E. loftini in North America. Records compiled from (Johnson 1984, Browning 1989, Reay-Jones et al. 2007, and Wilson
et al. 2015, 2017a).
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Fig. 3. Eoreuma loftini detection site in the U.S. state of Georgia, 2020 and 2021.

into Louisiana. Monitoring in adjacent states in the southeastern                     were from S. halepense and P. urvillei, both of which are considered
United States for potential human-aided introductions is not cur-                     invasive pests in the southern United States (Beuzelin et al. 2011).
rently conducted, but may be warranted if spread continues                            Population declines of other crambid stem borers have been reported
unpredictably.                                                                        following establishment of E. loftini in parts of Mexico (Rodriguez-
                                                                                      del-Bosque et al. 2011), although these declines may have resulted
                                                                                      from changes in natural enemy communities rather than competitive
Eoreuma loftini Ecological and                                                        displacement. While ecological impacts following establishment of
Economic Impacts                                                                      E. loftini are thought to be minimal in most regions, more research
Eoreuma loftini is a polyphagous stem borer which attacks nu-                         into this area is needed.
merous crop and non-crop grasses in the family Poaceae (Beuzelin
et al. 2016a, Showler 2019). Crop hosts include corn (Zea mays),
                                                                                      Establishment in Georgia
sugarcane (Saccharum spp.), rice (Oryza sativa), and sorghum
(Sorghum bicolor) in decreasing order of host preference (Showler                     Initial alert of potential detection of E. loftini in Georgia occurred
et al. 2012). Detections have also been reported from wheat                           as the result of a post to a social media group for producers of
(Triticum aestivum) in Texas, though its pest status in the crop is                   sugarcane syrup in the southeastern United States. Photos showed
unknown. Key non-crop grass hosts include johnsongrass (Sorghum                       larvae with great resemblance to E. loftini along with larval injury
halepense), vaseygrass (Paspalum urvillei), brome (Bromus spp.), and                  consistent with that caused by the species to sugarcane in Louisiana
others (Beuzelin et al. 2011, 2013, 2016a). Many other grass species                  and Texas (B. E. Wilson, personal observation). Contact was made
have also been reported as hosts, with early reports suggesting any                   with the private citizen making the initial post who confirmed her lo-
grass species with sufficient stem size to support larvae feeding could               cation in Effingham County, Georgia, USA and agreed to assist with
be attacked (Van Zwaluwenburg 1926, Beuzelin et al. 2016a).                           specimen collection. Six larvae and one pupa were collected from the
    Economic impacts result from damaging infestations that can                       sugarcane patch on 1 December 2020, stored in alcohol, and shipped
occur in crop hosts. Yield loss of up to 15% has been reported from                   to the Louisiana State University Department of Entomology (Baton
unmitigated infestations in rice in Louisiana and Texas (Reay-Jones                   Rouge, LA).
et al. 2007, Wilson et al. 2021). Losses in sugarcane have been re-                        We confirmed the first distribution record of E. loftini from Georgia,
ported to be 15–25% (Wilson et al. 2012, Showler and Reagan                           USA based on both morphological and molecular data. In morphological
2017). Economic impacts to corn and sorghum are less well studied,                    analysis conducted in 2021, each specimen was examined under a micro-
but plant death has been reported to result from extensive internal                   scope. Larvae of E. loftini were distinguished from other crambid larvae
feeding in plants of both crops (Showler et al. 2012).                                by the presence of a single subventral seta on the meso- and metathorax
    Ecological impacts of E. loftini in non-crop habitats have not                    (Beuzelin et al. 2016a). Additional photos from the same field were pro-
been well-studied. The species is known to utilize native and inva-                   vided in September of 2021 and also determined to be E. loftini. To iden-
sive grasses along the U.S. Gulf Coast, particularly during winter                    tify the specimens using molecular method, a 658bp fragment of the
when crop hosts are not available (Beuzelin et al. 2011, 2013). While                 mitochondrial cytochrome c oxidase subunit I gene (COI) of each insect
the species has been found in densities of >3 larvae/m2 in non-crop                   was targeted and amplified using a primer set, LepF1 and LepR1, devel-
habitats in invaded regions, these populations are not likely to be                   oped by. DNA was extracted from larval and pupal specimens using the
sufficient to alter plant communities. Further, the primary grasses                   DNeasy Blood and Tissue Kit (QIAGEN, Hilden, Germany) and amp-
from which E. loftini was recovered in non-crop habitats of Texas                     lified by polymerase chain reactions (PCR). PCR volumes were 25 µl
4                                                                            Journal of Integrated Pest Management, 2022, Vol. 13, No. 1

containing 12.5 µl of DreamTaq Green PCR Master Mix (2X) (Thermo                The mode of introduction of E. loftini into Georgia is unknown,
Scientific), 1–2 µl of template genomic DNA, 9.5 µl of ddH2O, and 1.0 µl    though the apparent disjunct expansion suggests that human-aided
of each primer at 5–10 µM. PCR conditions were 95°C for 3 min; 5            movement is a likely cause. Movement of infested plant material is
initial cycles of 95°C for 30 s, 45°C for 30 s, 72°C for 1 min; 35 cycles   the primary mode of introduction for invasive herbivores (Hulme
of 95°C for 30 s, 51°C for 30 s and 72°C for 1 min; and a final ex-         2009). Although the initial detection occurred in sugarcane, this does
tension at 72°C for 7 min. Crude PCR products were visualized on a          not necessarily implicate movement of sugarcane in the spread to
1.8–2.0% agarose gels to confirm the success of amplification sent to the   Georgia. Infestations of E. loftini in sugarcane in Florida have not
LSU genomics facility. PCR products were initially cleaned by a EtOH        been reported, so transportation of sugarcane from the region would
precipitation clean-up method developed by the facility and sequenced       not likely have contributed to the spread. Infested sugarcane imports
on the 3130xl Genetic Analyzers (Applied Biosystems) using the BigDye       could have come from the much more distant sources in western
Terminator v1.1 chemistry (Applied Biosystems). DNA assembly and            Louisiana or the Rio Grande Valley of Texas, but transportation of
sequence editing were conducted using Geneious Prime 2019.2 (https://       sugarcane more than 1,000 km seems unlikely with closer sources
www.geneious.com). The obtained sequences were compared with COI            available. Further, the species may have been present in Georgia for
sequences of E. loftini deposited in the GenBank database of the National   several years, infesting many of the suitable weedy grass hosts before
Center for Biotechnology Information (NCBI) using the Basic Local           detection in crop plants. Movement of grass materials such as hay or

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Alignment Search Tool (BLAST).                                              other forages for livestock feed is one possible means of transportation.
     Using BLAST, we confirmed that all the newly obtained COI              Florida is a major producer of hay forage with more than 100,000 ha
sequences were ~99%           (Paspalum spp.) which are the known hosts of E. loftini (Chambliss
matched with seven deposited sequences obtained from specimens              et al. 2006, Beuzelin et al. 2016a). While most of this hay is sold as
collected in Texas (Weslaco, Hidalgo), Louisiana (Baton Rouge),             livestock fodder within Florida, it can generally be transported out of
Florida, northwestern Mexico (Los Mochis, Sinaola), and >~97%               state without restriction (USDA APHIS 2021). Unfortunately, deter-
matched with three deposited sequences obtained from specimens              mination of the mode of introduction with any degree of certainty is
collected in the south-central region of Mexico (Zacatepec, Morelos).       not possible. Future studies could, however, identify the source popu-
The molecular data indicate the E. loftini population introduced to         lation through genetic analyses and comparisons to known popula-
Georgia is closely related with U.S. populations and the population         tions from Texas, Louisiana, Florida, and Mexico.
from northwestern Mexico. Additionally, we confirmed that utilizing             Economic impacts to crop production in Georgia and surrounding
COI barcoding helps quick and accurate identification of E. loftini.        states are possible in coming years. Georgia currently produces ap-
     Because COI barcodes obtained from E. loftini collected in             proximately 190,000 ha of corn and 90,000 ha of wheat with com-
Louisiana and Texas are not available on the GenBank yet, it is not         parable production of both crops in nearby South Carolina (USDA
possible to determine the potential origins of the population intro-        NASS 2021). More than >80% of the corn hectares in Georgia are
duced into Georgia.                                                         planted with insecticidal Bacillus thuringiensis (Bt) varieties (Buntin
     This detection represents a significant range expansion appearing      2021). Transgenic Bt corn varieties that are effective against other corn
to result from a new introduction. The extent of the established range      stalk borers should also minimize losses from E. loftini (Showler et al.
in Georgia remains unknown. Pheromone trap monitoring programs              2013, Beuzelin et al. 2016b). Increased economic losses are likely to
should be established following the methods of Wilson et al. (2015,         occur to non-Bt refuge corn in regions where E. loftini becomes estab-
2016, 2017a, 2017b) to delineate the extent of the invaded range in         lished, however (Showler et al. 2012). The prevalence of irrigation in
Georgia and possibly South Carolina.                                        Georgia corn production may limit E. loftini infestations as the pest is
     The location of the Effingham County detection is approxi-             exacerbated in drought conditions (Reay-Jones et al. 2008, Showler
mately 400 km (250 miles) north of the closest known population             and Castro 2010). Approximately, 90% of field corn is irrigated in
in Sumter County, Florida. Effingham county lies north of the 32nd          the Coastal Plain region surrounding the E. loftini detection site in
parallel, approximately 130 km north of the northern most loca-             Effingham County (Bryant 2021). Sweet corn and sugarcane grown
tion in Louisiana where the species is known to occur (Wilson and           for syrup production will also be at risk of infestation, but these crops
Huval 2020). This indicates that the species may be capable of col-         are not produced on a commercial farm scale in Georgia.
onizing more temperate regions than other sugarcane stem borers
such as the sugarcane borer, Diatraea saccharalis (F.). Indeed, the
cold tolerance of E. loftini populations in Mexico was found to be          Acknowledgments
greater than both D. saccharalis and the neotropical corn stalk borer,
                                                                            We express gratitude to citizen scientists for their willingness to assist in
Diatraea lineolata (Walker) (Rodriguez-del-Bosque et al. 1995).             ­specimen collection. Additional thanks are in order to Randy Richard for pro-
Based on its current known distribution, it is likely the species is cap-   viding the initial alert. This work was funded in part by the USDA NIFA Foun-
able of inhabiting the southern latitudes of Mississippi and Alabama.       dational Grant 1027792, USDA Hatch funds, and the American Sugarcane
Assuming these regions are suitable for E. loftini establishment, their     League. This publication is approved by the Louisiana State University Agri-
colonization in coming years is likely. The rate of expansion during        cultural Center (LSU-MS-2022-270-36697).
biological invasions generally increases with increasing size and
number of source populations (O’Reilly-Nugent et al. 2016). With            References Cited
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