Identification of Cotton Fleahopper (Hemiptera: Miridae) Host Plants In

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PLANTÐINSECT INTERACTIONS Identiﬁcation of Cotton Fleahopper (Hemiptera: Miridae) Host Plants in Central Texas and Compendium of Reported Hosts in the United States

1 J. F. ESQUIVEL AND S. V. ESQUIVEL

USDAÐARS, SPARC, Areawide Pest Management Research Unit, 2771 F&B Rd., College Station, TX 77845

Environ. Entomol. 38(3): 766Ð780 (2009) ABSTRACT The cotton ßeahopper, Pseudatomoscelis seriatus (Reuter), is an early-season pest of developing cotton in Central Texas and other regions of the Cotton Belt. Cotton ßeahopper populations develop on spring weed hosts and move to cotton as weed hosts senesce or if other weed hosts are not readily available. To identify weed hosts that were seasonably available for the cotton ßeahopper in Central Texas, blooming weed species were sampled during early-season (17 MarchÐ31 May), mid-season (1 JuneÐ14 August), late-season (15 AugustÐ30 November), and overwintering (1 DecemberÐ16 March) periods. The leading hosts for cotton ßeahopper adults and nymphs were evening primrose (Oenothera speciosa T. Nuttall) and Mexican hat [Ratibida columnifera (T. Nuttall) E. Wooton and P. Standley], respectively, during the early season. During the mid-season, silver-leaf nightshade (Solanum elaeagnifolium A. Cavanilles) was consistently a host for ßeahopper nymphs and adults. Woolly croton (Croton capitatus A. Michaux) was a leading host during the late season. Cotton ßeahoppers were not collected during the overwintering period. Other suitable hosts were available before previously reported leading hosts became available. Eight previously unreported weed species were documented as temporary hosts. A compendium of reported hosts, which includes Ͼ160 plant species representing 35 families, for the cotton ßeahopper is provided for future research addressing insectÐhost plant associations. Leading plant families were Asteraceae, Lamiaceae, and Onagraceae. Results presented here indicate a strong argument for assessing weed species diversity and abundance for the control of the cotton ßeahopper in the Cotton Belt.

KEY WORDS cotton ßeahopper, host plant, cotton pest

The cotton ßeahopper [Pseudatomoscelis seriatus (Re- and pest-speciÞc insecticides for lepidopteran control uter); Hemiptera: Miridae] was Þrst reported in Texas (Armstrong and Camelo 2003, Layton et al. 2003). The cotton (Gossypium hirsutum C. Linnaeus) by Howard aforementioned components have all been incorpo- (1898). The ßeahopper is an early-season pest that rated in the cotton-producing areas of the Southern uses piercing-sucking mouthparts to feed on develop- Blacklands region of Central Texas. Furthermore, the ing cotton fruit (i.e., squares). The affected squares Southern Blacklands region is in the advanced stages are shed by the cotton plant, resulting in yield losses. of boll weevil eradication efforts, and cotton ßeahop- Fleahopper adults typically move from senescing pers could potentially become a post-eradication pest weed hosts to infest cotton (Reinhard 1926a, Almand in Central Texas as well. Host plant data will be in- 1974). Adults have been shown to leave weed hosts strumental in determining sources of cotton ßeahop- and migrate to adjacent cotton (Suh et al. 2004), or in per populations that subsequently infest cotton. some cases, have infested cotton 20 miles from the Historically, plant species in the Croton, Oenothera, original weed host (Gaines and Ewing 1938). Monarda, and Solanum genera have been identiÞed as Sucking insects, including the cotton ßeahopper, leading hosts for the cotton ßeahopper during the have gained importance as pests of cotton in recent overwintering, early-season, summer, and fall seasons, years. Reports attribute this elevated status to the respectively. Previous reports deÞned the leading adoption of transgenic cotton varieties, absence of hosts for the ßeahopper as members of the mentioned traditional early-season insecticide sprays to control genera but the composition of plant species within boll weevils (Anthonomus grandis grandis Boheman), these genera has varied among studies, likely because of differences in plant composition among geograph- ical locations. Furthermore, some of these reports This article reports the results of research only. Mention of a (Gaines and Ewing 1938, Fletcher 1940, Almand 1974, proprietary product does not constitute an endorsement or a recom- mendation by the USDA for its use. Idol and Slosser 2005) focused speciÞcally on selected 1 Corresponding author, e-mail: [email protected]. primary hosts with little regard for other hosts that June 2009 ESQUIVEL AND ESQUIVEL:HOSTS OF THE COTTON FLEAHOPPER 767 may have contributed to the developing ßeahopper ovipositor or male genitalia. Voucher specimens of populations. Reported hosts are further confounded wild host plant species were collected and sent to the by laboratory feeding (Gaylor and Sterling 1976a, b; S.M. Tracy Herbarium at Texas A&M University, Col- Beerwinkle and Marshall 1999) and oviposition stud- lege Station, TX, for identiÞcation and cataloging. ies (Holtzer and Sterling 1980) or all-encompassing To determine the presence of cotton ßeahoppers in statements listing hosts without providing sampling associated temporal host plant species, sampling con- data. For example, Brett (1946) and Hixson (1941) tinued throughout the year beginning on 17 March alluded to the breadth of cotton ßeahopper host plants 2003. Temporal occurrence of plant species was cat- but failed to provide sampling data. Similarly, Eddy egorized into early season (17 MarchÐ31 May), mid- (1927) stated that “other succulent plants” were also season (1 JuneÐ14 August), late season (15 AugustÐ30 hosts. Finally, Fletcher (1930) discussed cyclic November), and overwintering (1 DecemberÐ16 changes in vegetation and their effects on the insect March). These categories coincided with local farm- fauna in Central Texas. Because of this latter point, the ing and production practices. Host plants were pre- prevalence of anecdotal records of host plants, and the sumed to be and deÞned as those species where cotton absence of a current taxonomically accurate database ßeahopper adults and/or nymphs were present. Be- of cotton ßeahopper host plants, the objectives of this cause nymphs do not have wings to move from host to study were to identify temporally available weed spe- host, weed species yielding nymphs were considered cies harboring cotton ßeahoppers in the southern likely reproductive hosts. In total, 76 sites of weed Blackland Prairies of Central Texas and provide a species and six sites of cultivated species (Glycine max deÞnitive and current listing of host plants associated and Medicago sativa) were sampled during the 3-yr with the cotton ßeahopper. study. Although some weed species were only sampled 1 yr because of the variable yearly occurrence or abundance, these data were included to show the Materials and Methods range of plant utilization by cotton ßeahoppers in the Between 17 March 2003 and 17 March 2006, bloom- region. ing herbaceous weeds, alfalfa (Medicago sativa C. Lin- Because of taxonomic revisions of previously re- naeus), and soybeans [Glycine max (C. Linnaeus) E. ported plants and their respective authorities, the Merrill] were sampled for cotton ßeahoppers in Bur- compendium of cotton ßeahopper host plants pre- leson and Robertson counties (Central Texas). As sented here was formulated from a literature review blooming weed species were identiÞed along right- beginning with Howard (1898) through 2008. Refer- of-ways, turn-rows, and ditches, these species were eed journal articles, proceedings, Federal and State subject to sampling. Gaylor (1975) previously con- Agency bulletins, and theses and dissertations with cluded that plant species were more suitable hosts for speciÞc reference to cotton ßeahopper and host plant cotton ßeahoppers when ßowering. Pneumatic air associations were reviewed. Sterling and Dean (1977) samplers and beat sheets were evaluated in prelimi- were consulted for earlier reports of host plant asso- nary studies on row crops, but these tools were not ciations. The botanical nomenclature presented fol- conducive to efÞcient sampling in weed hosts. Be- lows that of Jones et al. (1997). Some previous reports cause of inconsistent plant densities and variable plant only provided common names of host plants. Where architectures in weed species, sweep nets (38.1 cm this occurred, the host plants were assigned to genera diameter) were used for sampling weed hosts. For only (e.g., ragweed ϭ Ambrosia spp.; horsemint ϭ similar reasons, the sweep net was used in previous Monarda spp.) for inclusion in this report. Where host host plant studies (Cleveland 1982, Anderson and species occurred outside of Texas, online searches Schuster 1983, Womack and Schuster 1987, Armstrong were conducted using http://plants.usda.gov/index. and Camelo 2003). Unlike some previous host plant html to locate current taxonomic designations. studies that used variable sample sizes within each The PROC MEANS statement (SAS Institute 2004) study (Cleveland 1982, Snodgrass et al. 1984, Fleischer and the SUM and BY options were used to generate and Gaylor 1987), a uniform protocol of three samples descriptive statistics of the number of cotton ßeahop- (50 sweeps/sample) for each sample site was imple- per adults and nymphs in temporal host plants sam- mented in this study. Because of this sampling proto- pled during this study. Similarly, the mean proportions col, plant species that were in bloom but not in suf- of adults that were female were determined using the Þcient quantity to support 150 sweeps were not MEAN and BY options of the PROC MEANS state- sampled. Plants were sampled twice per week at 3- to ment. 4-d intervals unless precluded by inclement weather. In some instances, multiple sites of the same plant Results and Discussion species were sampled because of the overall abundance of these species. Cotton ßeahopper adults and nymphs were col- Materials sampled by sweep nets were placed in lected from various host plants during each sampling 3.79-liter Ziploc bags, returned to the laboratory, and year. Of the 36 plant species sampled in this study placed in a freezer to facilitate processing of samples. (Table 1), the following eight weed species were pre- Adults and nymphs of cotton ßeahoppers were viously unreported hosts: Amaranthus hybridus C. Lin- counted. Adults were placed under a dissecting scope, naeus; Amaranthus retroﬂexus C. Linnaeus; Bradburia and the ventral side was examined for the presence of pilosa (T. Nuttall); Melilotus indicus (C. Linnaeus) C. 768 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 3

Table 1. Summary of plant species sampled for cotton ﬂeahoppers during 2003–2006 in Central Texas and plant host status

Adults (A) or Plant species Common name nymphs (N) Host status captured Amaranthus hybridus C. Linnaeus Slim amaranth A, N Previously unreported Amaranthus retroflexus C. Linnaeus Redroot amaranth A Previously unreported Ambrosia trifida C. Linnaeus (Sy ϭ A. aptera Giant ragweed A, N Previously reported A.P. de Candolle) Bradburia pilosa (T. Nuttall) Soft goldaster A, N Previously unreported Capsella bursa-pastoris (C. Linnaeus) F. Medikus ShepherdÕs purse Ñ Previously reported; non-host in current study Chaerophyllum tainturieri W. J. Hooker var. Hairyfruit chervil A, N Previously reported tainturieri (Sy ϭ C. ßoridanum (J. M. Coulter & J. Rose) Bush) Chamaecrista fasciculata (A. Michaux) E. Greene Prairie senna A Previously reported (Sy ϭ Cassia fasciculata A. Michaux) Cirsium texanum S. Buckley Southern thistle Ñ Non-host in current study Convolvulus equitans G. Bentham Gray bindweed Ñ Non-host in current study Conyza canadensis (C. Linnaeus) A. Cronquist Horsetail A, N Previously reported var. canadensis (Sy ϭ Erigeron canadensis C. Linnaeus; Sy ϭ Leptilon canadense (C. Linnaeus) N. Britton)) Croton capitatus A. Michaux Woolly croton A, N Previously reported Gaura coccinea T. Nuttall ex F. Pursh Scarlet gaura A Previously reported Glycine max (C. Linnaeus) E. Merrill Soybean A, N Previously reported Helianthus annuus C. Linnaeus Common sunßower Ñ Previously reported; non-host in current study Lamium amplexicaule C. Linnaeus Henbit deadnettle Ñ Previously reported; non-host in current study Medicago polymorpha C. Linnaeus Bur clover A, N Previously reported Medicago sativa C. Linnaeus Alfalfa A, N Previously reported Melilotus indicus (C. Linnaeus) C. Allioni Annual yellow A, N Previously unreported sweetclover Monarda citriodora V. de Cervantes ex M. Lemon beebalm A, N Previously reported Lagasca y Segura Oenothera speciosa T. Nuttall Mexican evening primrose A, N Previously reported Packera glabellus (J. Poiret) Butterweed Ñ Nonhost in this study Parthenium hysterophorus C. Linnaeus Ragweed parthenium A, N Previously reported Phyla nodiflora (C. Linnaeus) E. Greene (Sy ϭ Sawtooth frogfruit A, N Previously reported Lippia nodißora (J. de Lamarck)A. Michaux) Pyrrhopappus carolinianus (T. Walter) A. P. de Carolina false dandelion A, N Previously reported Candolle Rapistrum rugosum (C. Linnaeus) C. Allioni Turnipweed A, N Previously reported Ratibida columnifera (T. Nuttall) E. Wooton & Upright prairie coneßower A, N Previously reported P. Standley (Sy ϭ R. columnaris (J. Sims) D. Don) Rudbeckia amplexicaulis M. H. Vahl Clasping coneßower A, N Previously unreported Ruellia nudiflora (G. Engelmann ex A. Gray) I. Violet ruellia A, N Previously unreported Urban Sisymbrium irio C. Linnaeus Rocketmustard Ñ Nonhost in this study Solanum elaeagnifolium A. Cavanilles Silverleaf nightshade A, N Previously reported Solidago sp. Goldenrod A, N Previously reported Trifolium incarnatum C. Linnaeus Crimson clover A, N Previously reported Trifolium repens C. Linnaeus White clover Ñ Nonhost in this study Verbena neomexicana (A. Gray) J. K. Small var. Hillside vervain A, N Previously unreported hirtella L. Perry Verbesina encelioides (A. Cavanilles) G. Bentham Cowpen daisy A Previously unreported & J. Hooker ex A. Gray Xanthium strumarium C. Linnaeus (Sy ϭ X. Rough cocklebur A Previously reported speciosum T. Kearney)

Common names based on designation by Hatch et al. (1990). Ñ, cotton ßeahoppers were not encountered.

Allioni; Rudbeckia amplexicaulis M. H. Vahl; Ruellia mium amplexicaule C. Linnaeus) yielded cotton ßea- nudiﬂora (G. Engelmann ex A. Gray) I. Urban; Ver- hoppers. In addition to these latter three species, Þve bena neomexicana (A. Gray) J. K. Small variety hirtella more species sampled in our study were nonhosts. L. Perry; and Verbesina encelioides (A. Cavanilles) G. The seasonal occurrence of cotton ßeahopper Bentham and J. Hooker ex A. Gray. Of the 23 previ- adults and nymphs in Central Texas plants are shown ously reported hosts sampled during this study, all but in Tables 2 and 3, respectively. In all years, Oenothera three species (Capsella bursa-pastoris (C. Linnaeus) speciosa T. Nuttall was the leading host for adults F. Medikus; Helianthus annuus C. Linnaeus; and La- during the early season (Table 2). Ratibida columnif- June 2009 ESQUIVEL AND ESQUIVEL:HOSTS OF THE COTTON FLEAHOPPER 769

Table 2. Temporal mean no. of cotton ﬂeahopper adults per 50-sweep sample (per sweep; n) collected in identiﬁed plants of Central Texas, where n represents no. of 50-sweep samples

2003 2004 2005 Season Plant species ͓per 50 sweeps ͓per 50 sweeps ͓per 50 sweeps (per sweep; n)͔ (per sweep; n)͔ (per sweep; n)͔ Early season (17 Mar. to Amaranthus retroflexus C. Linnaeus 18.0 (0.360; 15) Ñ Ñ 31 May) Capsella bursa-pastoris (C. Linnaeus) F. Medikus Ñ 0 (0; 6) 0 (0; 9) Chaerophyllum tainturieri W. J. Hooker 10.6 (0.211; 9) Ñ Ñ Conyza canadensis (C. Linnaeus) A. Cronquist Ñ 0.25 (0.005; 12) Ñ var. canadensis Lamium amplexicaule C. Linnaeus Ñ 0 (0; 3) Ñ Medicago polymorpha C. Linnaeus 23.7 (0.474; 42) 0.17 (0.003; 30) 0.13 (0.003; 30) Medicago sativa C. Linnaeus 17.64 (0.353; 81) 0.23 (0.005; 48) 0.86 (0.172; 36) Melilotus indicus (C. Linnaeus) C. Allioni 14.67 (0.293; 15) Ñ 0.17 (0.003; 18) Oenothera speciosa T. Nuttall 422.78 (8.46; 9) 30.87 (0.617; 30) 10.33 (0.207; 18) Packera glabellus (J. Poiret) 0 (0; 15) Ñ 0 (0; 3) Pyrrhopappus carolinianus (T. Walter) A. P. de 3.63 (0.73; 27) Ñ Ñ Candolle Rapistrum rugosum (C. Linnaeus) C. Allioni 43.22 (0.864; 69) 2.72 (0.054; 54) 0.04 (0.001; 24) Ratibida columniferas (T. Nuttall) E. Wooton 88.13 (1.763; 15) 4.83 (0.097; 12) Ñ and P. Standley (Sy ϭ Ratibida columnaris (J. Sims) D. Don) Rudbeckia amplexicaulis M. H. Vahl 15.8 (0.32; 15) 3.07 (0.061; 15) 4.38 (0.088; 21) Sisymbrium irio C. Linnaeus Ñ 0 (0; 6) Ñ Solanum elaeagnifolium A. Cavanilles 306.67 (6.13; 3) Ñ Ñ Trifolium incarnatum C. Linnaeus 3.50 (0.070; 6) 0.71 (0.014; 21) Ñ Trifolium repens C. Linnaeus Ñ 0 (0; 15) Ñ Xanthium strumarium C. Linnaeus Ñ 0.14 (0.003; 21) Ñ Mid-season (1 Jun. to 14 Amaranthus hybridus C. Linnaeus 0.71 (0.014; 21) 0.33 (0.007; 12) Ñ Aug.) Ambrosia trifida C. Linnaeus Ñ 0 (0; 21) Ñ Bradburia pilosa T. Nuttall 0.08 (0.001; 12) Ñ Ñ Chamaecrista fasciculata (A. Michaux) E. Greene Ñ 0.11 (0.002; 18) Ñ (Sy ϭ Cassia fasciculata A. Michaux) Cirsium texanum S. Buckley 0 (0; 3) Ñ Ñ Convolvulus equitans G. Bentham 0 (0; 6) Ñ Ñ Conyza canadensis 0 (0; 30) 0.26 (0.005; 42) Ñ Croton capitatus A. Michaux Ñ 45.96 (0.919; 24) Ñ Rudbeckia amplexicaulis 0.11 (0.002; 9) Ñ Ñ Gaura coccinea T. Nuttall ex F. Pursh Ñ 0.38 (0.007; 21) Ñ Glycine max (C. Linnaeus) E. Merrill 0.15 (0.002; 48) 0 (0; 42) 0 (0; 6) Helianthus annuus C. Linnaeus Ñ 0 (0; 18) Ñ Medicago sativa 0.51 (0.010; 105) 0.21 (0.004; 48) 0.07 (0.001; 30) Monarda citriodora V. de Cervantes ex M. Ñ Ñ 421.11 (8.422; 9) Lagasca y Segura Phyla nodiflora (C. Linnaeus) E. Greene 0.28 (0.006; 39) Ñ Ñ Ratibida columnifera 0.55 (0.011; 33) 7.83 (0.157; 18) Ñ Ruellia nudiflora (G. Engelmann ex A. Gray) I. 0.22 (0.004; 9) Ñ Ñ Urban Solanum elaeagnifolium 6.44 (0.129; 49) 5.04 (0.100; 27) 4.00 (0.080; 6) Verbena neomexicana (A. Gray) J. K. Small var. 3.26 (0.065; 39) Ñ Ñ hirtella L. Perry Late season (15 Aug. to Amaranthus hybridus Ñ 16.0 (0.320; 15) Ñ 30 Nov.) Ambrosia trifida 13.05 (0.261; 39) 0.92 (0.019; 39) Ñ Conyza canadensis 0.79 (0.016; 33) Ñ Ñ Croton capitatus 207.78 (4.156; 45) 315.03 (6.301; 33) Ñ Lamium amplexicaule Ñ 0 (0; 3) Ñ Medicago sativa 0.05 (0.001; 63) 0.20 (0.004; 45) 0.16 (0.003; 45) Parthenium hysterophorus C. Linnaeus 7.00 (0.140; 36) 0 (0; 18) 16.17 (0.323; 18) Phyla nodiflora 3.23 (0.064; 39) Ñ Ñ Rapistrum rugosum 0.73 (0.015; 15) Ñ Ñ Solanum elaeagnifolium 1.67 (0.033; 6) Ñ 2.21 (0.044; 57) Solidago sp. 32.40 (0.648; 30) 0.20 (0.004; 15) Ñ Verbena neomexicana 12.6 (0.252; 30) Ñ Ñ Verbesina encelioides (A. Cavanilles) G. Bentham 0.20 (0.004; 15) Ñ Ñ and J. Hooker ex A. Gray Overwintering (1 Dec. to Capsella bursa-pastoris 0 (0; 6) Ñ Ñ 16 Mar.) Lamium amplexicaule 0 (0; 66) 0 (0; 30) Ñ Medicago polymorpha Ñ 0 (0; 15) Ñ Medicago sativa 0 (0; 63) 0 (0; 36) 0 (0; 36) Parthenium hysterophorus 0 (0; 6) Ñ Ñ Rapistrum rugosum 0 (0; 18) 0 (0; 12) 0 (0; 15) Sisymbrium irio 0 (0; 24) 0 (0; 15) 0 (0; 21) Trifolium repens 0 (0; 6) Ñ Ñ

Ñ, plant species not sampled because of insufÞcient plant stand or completely absent. 770 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 3

Table 3. Temporal mean no. of cotton ﬂeahopper nymphs per 50-sweep sample (per sweep; n) collected in identiﬁed plants of Central Texas, where n represents no. of 50-sweep samples

2003 2004 2005 Season Plant species ͓per 50 sweeps ͓per 50 sweeps ͓per 50 sweeps (per sweep; n)͔ (per sweep; n)͔ (per sweep; n)͔ Early season (17 Mar. to Amaranthus retroflexus C. Linnaeus 0 (0; 15) Ñ Ñ 31 May) Capsella bursa-pastoris (C. Linnaeus) F. Medikus Ñ 0 (0; 6) 0 (0; 9) Chaerophyllum tainturieri W. J. Hooker 0.33 (0.007; 9) Ñ Ñ Conyza canadensis (C. Linnaeus) A. Cronquist Ñ 0.42 (0.008; 12) Ñ var. canadensis Lamium amplexicaule C. Linnaeus Ñ 0 (0; 3) Ñ Medicago polymorpha C. Linnaeus 0.21 (0.004; 42) 0.10 (0.002; 30) 0 (0; 30) Medicago sativa C. Linnaeus 1.22 (0.024; 81) 0 (0; 48) 0.36 (0.007; 36) Melilotus indicus (C. Linnaeus) C. Allioni 0.13 (0.003; 15) Ñ 0 (0; 18) Oenothera speciosa T. Nuttall 27.00 (0.540; 9) 9.07 (0.181; 30) 0.17 (0.003; 18) Packera glabellus (J. Poiret) 0 (0; 15) Ñ 0 (0; 3) Pyrrhopappus carolinianus (T. Walter) A. P. de 1.11 (0.022; 27) Ñ Ñ Candolle Rapistrum rugosum (C. Linnaeus) C. Allioni 0.22 (0.004; 69) 0 (0; 54) 0 (0; 24) Ratibida columnifera (T. Nuttall) E. Wooton and 54.20 (1.084; 15) 13.83 (0.277; 12) Ñ P. Standley (Sy ϭ Ratibida columnaris (J. Sims) D. Don) Rudbeckia amplexicaulis M. H. Vahl 13.07 (0.261; 15) 0 (0; 15) 1.48 (0.030; 21) Sisymbrium irio C. Linnaeus Ñ 0 (0; 6) Ñ Solanum elaeagnifolium A. Cavanilles 30.33 (0.607; 3) Ñ Ñ Trifolium incarnatum C. Linnaeus 0.83 (0.017; 6) 0.05 (0.001; 21) Ñ Trifolium repens C. Linnaeus Ñ 0 (0; 15) Ñ Xanthium strumarium C. Linnaeus Ñ 0 (0; 21) Ñ Mid-season (1 Jun. to 14 Amaranthus hybridus C. Linnaeus 0 (0; 21) 0.08 (0.002; 12) Ñ Aug.) Ambrosia trifida C. Linnaeus Ñ 0 (0; 21) Ñ Bradburia pilosa T. Nuttall 0.17 (0.003; 12) Ñ Ñ Chamaecrista fasciculata (A. Michaux) E. Greene Ñ 0 (0; 18) Ñ (Sy ϭ Cassia fasciculata A. Michaux) Cirsium texanum S. Buckley 0 (0; 3) Ñ Ñ Convolvulus equitans G. Bentham 0 (0; 6) Ñ Ñ Conyza canadensis 0 (0; 30) 0.02 (0.001; 42) Ñ Croton capitatus A. Michaux Ñ 20.50 (0.410; 24) Ñ Gaura coccinea T. Nuttall ex F. Pursh Ñ 0 (0; 21) Ñ Glycine max (C. Linnaeus) E. Merrill 0.04 (0.001; 48) 0 (0; 42) 0 (0; 6) Helianthus annuus C. Linnaeus Ñ 0 (0; 18) Ñ Medicago sativa 0.20 (0.004; 105) 0(0; 48) 0 (0; 30) Monarda citriodora V. de Cervantes ex M. Ñ Ñ 427.33 (8.547; 9) Lagasca y Segura Phyla nodiflora (C. Linnaeus) E. Greene 0.08 (0.002; 39) Ñ Ñ Ratibida columnifera 0.30 (0.006; 33) 3.17 (0.063; 18) Ñ Rudbeckia amplexicaulis 0.11 (0.002; 9) Ñ Ñ Ruellia nudiflora (G. Engelmann ex A. Gray) I. 0.11 (0.002; 9) Ñ Ñ Urban Solanum elaeagnifolium 1.27 (0.254; 49) 1.85 (0.037; 27) 1.00 (0.020; 6) Verbena neomexicana (A. Gray) J. K. Small var. 0.26 (0.005; 39) Ñ Ñ hirtella L. Perry Late season (15 Aug. to Amaranthus hybridus Ñ 0 (0; 15) Ñ 30 Nov.) Ambrosia trifida 0.28 (0.005; 39) 0.03 (0.001; 39) Ñ Conyza canadensis 0 (0; 33) Ñ Ñ Croton capitatus 156.09 (3.122; 45) 126.42 (2.528; 33) Ñ Lamium amplexicaule Ñ 0 (0; 3) Ñ Medicago sativa 0 (0; 63) 0 (0; 45) 0 (0; 45) Parthenium hysterophorus C. Linnaeus 0 (0; 36) 0 (0; 18) 0.17 (0.003; 18) Phyla nodiflora 0.18 (0.004; 39) Ñ Ñ Rapistrum rugosum 0.13 (0.003; 15) Ñ Ñ Solanum elaeagnifolium 0 (0; 6) Ñ 0.05 (0.001; 57) Solidago spp. 2.97 (0.059; 30) 0 (0; 15) Ñ Verbena neomexicana 0.70 (0.014; 30) Ñ Ñ Verbesina encelioides (A. Cavanilles) G. Bentham 0 (0; 15) Ñ Ñ and J. Hooker ex A. Gray Overwintering (1 Dec. to Capsella bursa-pastoris 0 (0; 6) Ñ Ñ 16 Mar.) Lamium amplexicaule 0 (0; 66) 0 (0; 30) Ñ Medicago polymorpha Ñ 0 (0; 15) Ñ Medicago sativa 0 (0; 63) 0 (0; 36) 0 (0; 36) Parthenium hysterophorus 0 (0; 6) Ñ Ñ Rapistrum rugosum 0 (0; 18) 0 (0; 12) 0 (0; 15) Sisymbrium irio 0 (0; 24) 0 (0; 15) 0 (0; 21) Trifolium repens 0 (0; 6) Ñ Ñ

Ñ, plant species not sampled because of insufÞcient plant stand or completely absent. June 2009 ESQUIVEL AND ESQUIVEL:HOSTS OF THE COTTON FLEAHOPPER 771 era (T. Nuttall) E. Wooton and P. Standley was the Table 4. Overall mean proportions of cotton fleahopper adults second leading host in the 2 yr that this weed was that were female collected from identified plants in Central Texas from 17 Mar. 2003 through 17 Mar. 2006, where n represents the sampled. The third leading host for adults was Solanum no. of 50-sweep samples yielding female adults elaeagnifolium A. Cavanilles during 2003 but Rud- beckia amplexicaulis M. H.Vahl replaced Solanum elae- Year Plant taxa agnifolium in this position during 2004 and 2005. S. 2003 (n) 2004 (n) 2005 (n) elaeagnifolium did not occur during the early season in Amaranthus hybridus 0.13 (4) 0.49 (10) Ñ these latter years. The leading early-season host for Amaranthus retroflexus 0.32 (11) Ñ Ñ nymphs during 2003 and 2004 was R. columnifera; R. Ambrosia trifida 0.40 (24) 0.37 (9) Ñ amplexicaulis was the leading host during 2005 (Table Bradburia pilosa 1.00 (1) Ñ Ñ 3). Second and third leading hosts varied yearly during Capsella bursa-pastoris 000 Chamaecrista fasciculata Ñ 0.00 (2) Ñ the study but included S. elaeagnifolium, Oenothera Chaerophyllum tainturieri 0.23 (9) Ñ Ñ speciosa, Conyza canadensis (C. Linnaeus) A. Cron- Cirsium texanum 0ÑÑ quist variety canadensis, and Medicago sativa. Convolvulus equitans 0ÑÑ In all years, cotton ßeahopper adults were Þrst col- Conyza canadensis 0.46 (10) 0.57 (6) Ñ Croton capitatus 0.46 (42) 0.43 (57) Ñ lected during late March or early April. The identiÞed Gaura coccinea Ñ 0.69 (4) Ñ leading early-season hosts observed in this study agree Glycine max 0.67 (5) 0 0 with previous reports. However, our early-season ob- Helianthus annuus Ñ0Ñ servations indicated that during 2003 adults were Lamium amplexicaule 00Ñ Medicago polymorpha 0.32 (39) 0.13 (4) 0.83 (2) present in other species before O. speciosa, a previ- Medicago sativa 0.36 (80) 0.30 (17) 0.24 (21) ously reported preferred host. Esquivel (2005) ini- Melilotus indicus 0.26 (10) Ñ 0.00 (2) tially reported the presence of cotton ßeahoppers in Plant taxa 2003 (n) 2004 (n) 2005 (n) R. rugosum (C. Linnaeus) C. Allioni. Fleahopper Monarda citriodora Ñ Ñ 0.48 (9) Oenothera speciosa 0.45 (9) 0.34 (24) 0.27 (15) adults were collected in Chaerophyllum tainturieri Packera glabellus 0Ñ0 W. J. Hooker variety tainturieri, Medicago polymorpha Parthenium hysterophorus 0.36 (23) 0 0.43 (17) C. Linnaeus, Medicago sativa C. Linnaeus, Melilotus Phyla nodiflora 0.34 (31) Ñ Ñ indicus (C. Linnaeus) C. Allioni, Rapistrum rugosum, Pyrrhopappus 0.33 (23) Ñ Ñ carolinianus and Trifolium incarnatum C. Linnaeus on 14 April Rapistrum rugosum 0.38 (24) 0.29 (14) 0.00 (1) during the 2003 early season. Furthermore, the highest Ratibida columnifera 0.41 (23) 0.39 (25) Ñ numbers of adults were collected from R. rugosum on Rudbeckia amplexicaulis 0.47 (15) 0.34 (9) 0.26 (15) this date. Nymphs were not encountered. During Ruellia nudiflora 1.00 (2) Ñ Ñ Sisymbrium irio 000 2004, adults were Þrst collected in O. speciosa on 5 Solanum elaeagnifolium 0.52 (44) 0.36 (26) 0.53 (44) April followed by R. rugosum, T. incarnatum, and Xan- Solidago sp. 0.40 (28) 0.33 (3) Ñ thium strumarium C. Linnaeus on 8 April. Similarly, Trifolium incarnatum 0.35 (5) 0.19 (9) Ñ during 2005, adults were Þrst collected in O. speciosa Trifolium repens 00Ñ Verbena neomexicana 0.28 (49) Ñ Ñ on 7 April followed by R. rugosum and M. polymorpha Verbesina encelioides 1.00 (3) Ñ Ñ on 11 April. Xanthium strumarium Ñ 0.67 (3) Ñ During the mid-season, S. elaeagnifolium was consistently among the weed hosts yielding the highest Ñ, plant species not sampled. average number of cotton ßeahopper adults and nymphs (Tables 2 and 3), indicating that this was also C. Linnaeus harbored adults during the 2004 late- a reproductive host in all sampling years. Other lead- season period. However, only A. trifida harbored cot- ing hosts for adults during this period were Verbena ton ßeahopper nymphs. Parthenium hysterophorus C. neomexicana (A. Gray) J. K. Small variety hirtella, and Linnaeus and S. elaeagnifolium were the two leading Amaranthus hybridus C. Linnaeus during 2003, Croton hosts for adults and nymphs during the 2005 late- capitatus A. Michaux and Ratibida columnifera during season period. Typically, as the year progressed, fewer 2004, and Monarda citriodora V. de Cervantes ex M. and fewer hosts were available for sampling. Indeed, Lagasca y Segura and M. sativa during 2005. These only one additional host (i.e., M. sativa) yielded cotton same species were also leading hosts for nymphs (Ta- ßeahoppers during the late season in 2005. ble 3) during the mid-season; presumably, the avail- The overwintering status of the cotton ßeahopper able adults were reproducing on these hosts. M. cit- has been well documented (Reinhard 1926a, Eddy riodora was again conÞrmed as a preferred host 1927, Gaines 1933, Gaines and Ewing 1938), and it is indicated by the high capture numbers, including accepted that this insect overwinters in the egg stage, nymphs recovered. primarily in C. capitatus. Thus, cotton ßeahoppers Croton capitatus continued to harbor adults and were not collected during the overwintering period in nymphs into the late-season period (Tables 2 and 3) the conduct of this study, despite our sampling efforts during 2003 and 2004. Numerically, this weed species throughout the year. Nonetheless, our insect collec- harbored the most adults and nymphs during 2 sam- tions in C. capitatus during the late-season period pling yr. During 2003, Solidago sp. and V. neomexicana indicate that this weed species is indeed a preferred were the other two leading species harboring nymphs host at this time period. (Ambrosia trifida C. Linnaeus and Solidago harbored Sex ratio data indicated that the proportion of fe- adults). In addition to A. trifida, Amaranthus hybridus males in the identiÞed hosts ranged from 0.13 (A. 772 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 3

Table 5. Compendium of reported host plants for the cotton ﬂeahopper in the United States, 1898–2008

Plant family and species Study location Sampling perioda Reference Acanthaceae Ruellia nudiflora (G. Engelmann ex A. Gray) I. TX Mid-season Current study Urban Amaranthaceae Amaranthus spp. TX No sampling data Reinhard 1926b Amaranthus blitoides S. Watson TX Not provided Reinhard 1926a Amaranthus hybridus C. Linnaeus TX Mid-, late season Current study Amaranthus retroflexus C. Linnaeus TX Early season Current study Amaranthus torreyi (A. Gray) S. Watson TX Overwintering Reinhard 1928 Amaranthus viridis C. Linnaeus TX Not provided Reinhard 1926a AR, LA, MS Late summer Snodgrass et al. 1984 Tidestromia lanuginosa (T. Nuttall) P. Standley TX Oct. Knight 1926 OK Not provided Hixson 1940 TX Apr.ÐJune/fall/overwintering Schuster et al. 1969 AR, LA, MS Spring/early summer Snodgrass et al. 1984 Apiaceae Carum spp. TX No sampling data Reinhard 1926b Chaerophyllum tainturieri W. J. Hooker var. TX Not provided Reinhard 1926ab tainturieri (Sy ϭ C. ßoridanum (J. M. OK Not provided Hixson 1940 Coulter and J. Rose) Bush) TX Early season Current study Limnosciadium pinnatum (A. P. de Candolle) TX Apr./May Reinhard 1927b M. Mathias and L. Constance (Sy ϭ Cynosciadium pinnatum A. P. de Candolle) Torilis arvensis (W. Hudson) J. Link AR, LA, MS June Snodgrass et al. 1984 Torilis nodosa (C. Linnaeus) J. Gaertner TX Apr./May Reinhard 1927 Asclepiadaceae Asclepias viridula Chapman TX Not provided Reinhard 1926a Asteraceae Ambrosia spp. TX No sampling data Reinhard 1926b SC Fall Eddy 1927 TX Not provided Thomas 1936 Ambrosia artemisiifolia C. Linnaeus (Sy ϭ A. TX Overwintering Reinhard 1928 elatior C. Linnaeus) OK JulyÐSept. Hixson 1940b AR, LA, MS June/July Snodgrass et al. 1984 Ambrosia psilostachya A. P. de Candolle TX Not provided Reinhard 1926a TX Overwintering Reinhard 1928 OK Aug.ÐOct. Hixson 1940 Ambrosia trifida C. Linnaeus (Sy ϭ A. aptera TX Overwintering Reinhard 1928b A. P. de Candolle) OK JulyÐSept. Hixson 1940b TX Mid-, late season Current study Amphiachyris spp. TX Not provided Thomas 1936 Amphiachyris dracunculoides (A. P. de TX Not provided Reinhard 1926a Candolle) T. Nuttall TX Overwintering Reinhard 1928 OK Not provided Hixson 1940 Anthemis cotula C. Linnaeus OK Late spring/early summer Hixson 1940, 1941 AR, LA, MS May/June Snodgrass et al. 1984 Bradburia pilosa (T. Nuttall) TX Mid-season Current study Chrysanthemum sp. IL Not provided Knight 1941 Conyza bonariensis (C. Linnaeus) A. Cronquist AR, LA, MS July Snodgrass et al. 1984b (Sy ϭ Erigeron bonariensis C. Linnaeus) Conyza canadensis (C. Linnaeus) A. Cronquist AR, LA, MS July Snodgrass et al. 1984b var. canadensis (Sy ϭ Erigeron canadensis TX Early, mid-, late season Current study C. Linnaeus) Conyza canadensis (C. Linnaeus) A. Cronquist TX Apr./May Reinhard 1927b var. canadensis (Sy ϭ Leptilon canadense TX Overwintering Reinhard 1928b (C. Linnaeus) N. Britton) Coreopsis tinctoria T. Nuttall TX July Snodgrass et al. 1984 Diaperia proflifera (T. Nuttall ex A. P. de TX Apr./May Reinhard 1927b Candolle) T. Nuttall var. prolifera (Sy ϭ Filago prolifera (A. P. de Candolle) N. Britton) Erigeron sp. OK Not provided Hixson 1941 Erigeron annuus (C. Linnaeus) C. Persoon AR, LA, MS MayÐJuly Snodgrass et al. 1984 Erigeron philadelphicus C. Linnaeus AR, LA, MS May Snodgrass et al. 1984 Erigeron strigosus G. H. Muhlenberg ex C. von OK Not provided Hixson 1940b Willdenow var. strigosus (Sy ϭ E. ramosus N. Britton, E. Sterns and J. Poggenburg) Eupatorium compositifolium T. Walter TX Overwintering Reinhard 1928 Eupatorium serotinum A. Michaux OK Aug.ÐOct. Hixson 1940

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Table 5. Continued

Plant family and species Study location Sampling perioda Reference Fleischmannia incarnata (T. Walter) R. King AR, LA, MS Oct. Snodgrass et al. 1984b and H. Robinson (Sy ϭ Eupatorium incarnatum T. Walter) Gaillardia pulchella Fougeroux de Bondaroy TX May Reinhard and Owen 1928 TX Laboratory study Beerwinkle and Marshall 1999 Gamochaeta purpurea (C. Linnaeus) A. TX Apr./May Reinhard 1927b Cabrera (Sy ϭ Gnaphalium purpureum C. Linnaeus) Gnaphalium spp. TX Early-season Almand 1974 Almand et al. 1976 TX AprÐJune Holtzer and Sterling 1980 Helenium amarum (C. RaÞnesque-Schmaltz) AR, LA, MS July/Sep./Oct. Snodgrass et al. 1984 H. Rock Helenium amarum (C. RaÞnesque-Schmaltz) TX Not provided Reinhard 1926ab H. Rock var. amarum (Sy ϭ H. tenuifolium TX Overwintering Reinhard 1928b T. Nuttall) OK JulyÐAug. Hixson 1940b Helianthus spp. TX No sampling data Reinhard 1926b Helianthus annuus C. Linnaeus TX Overwintering Reinhard 1928 TX Early season Schuster et al. 1969 TX Laboratory study Beerwinkle and Marshall 1999 Helianthus maximilianii H. A. Schrader TX Not provided Reinhard 1926a TX Overwintering Reinhard 1926b Helianthus petiolaris T. Nuttall OK Not provided Hixson 1940 Helianthus tuberosus C. Linnaeus OK Aug. Hixson 1940 Heterotheca stenophylla (A. Gray) L. Shinners OK Not provided Hixson 1941b var. stenophylla (Sy ϭ Chrysopsis villosa (F. Pursh) T. Nuttall ex A. P. de Candolle var. stenophylla (A. Gray) A. Gray) Heterotheca subaxillaris (J. de Lamarck) N. TX Overwintering Reinhard 1928 Britton and H. Rusby Hymenoxys spp. TX Not provided Thomas 1936 Iva annua C. Linnaeus (Sy ϭ I. ciliata C. von OK JulyÐOct. Hixson 1940b Willdenow) Krigia cespitosa (C. RaÞnesque-Schmaltz) K. TX Apr./May Reinhard 1927b Chambers var. gracilis (A. P. de Candolle) (Sy ϭ Serinia gracilis K.E.O. Kuntze) Lygodesmia juncea (F. Pursh) D. Don ex W. J. OK Not provided Hixson 1940 Hooker Parthenium spp. TX Not provided Thomas 1936 Parthenium hysterophorus C. Linnaeus TX No sampling data Hunter 1926 TX Overwintering Reinhard 1928 TX Early season Schuster et al. 1969 AR, LA, MS July Snodgrass et al 1983 TX Laboratory studies Beerwinkle and Marshall 1999 TX Late season Current study Psilostrophe villosa P. Rydberg OK Not provided Hixson 1940 Pyrrhopappus carolinianus (T. Walter) A. P. de AR/LA/MS July Snodgrass et al. 1984 Candolle TX Early season Current study Ratibida columnifera (T. Nuttall) E. Wooton TX Early season Schuster et al. 1969b and P. Standley (Sy ϭ Ratibida columnaris TX Early, mid-season Current study (J. Sims) D. Don) Rudbeckia amplexicaulis M. H. Vahl TX Early, mid-season Current study Solidago sp. OK Fall Eddy 1927 TX Late season Current study Solidago altissima C. Linnaeus OK Sept. Hixson 1940 Solidago tortifolia S. Elliot OK Aug.ÐSept. Hixson 1940 Symphyotrichum spp. (Sy ϭ Aster spp.) SC Fall Eddy 1927 TX Overwintering Reinhard 1928 Symphyotrichum pilosum (C. von Willdenow) AR, LA, MS Aug. Snodgrass et al. 1984 G. L. Nesom var. pilosum (Sy ϭ Aster pilosus Willdenow) Verbesina encelioides (A. Cavanilles) G. TX Late season Current study Bentham and J. Hooker ex A. Gray Xanthium strumarium C. Linnaeus (Sy ϭ X. TX Overwintering Reinhard 1928b speciosum T. Kearney; Sy ϭ X. americanum OK JulyÐOct. Hixson 1940b T. Walter) AR, LA, MS Sept. Snodgrass et al. 1984 TX Early-season Current study Azioaceae Trianthema portulacastrum C. Linnaeus TX Not provided Reinhard 1926a Berberidaceae Podophyllum peltatum C. Linnaeus TX Not provided Thomas 1936

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Table 5. Continued

Plant family and species Study location Sampling perioda Reference Brassicaceae Capsella bursa-pastoris (C. Linnaeus) F. OK Apr.ÐMay Hixson 1940 Medikus Dimorphocarpa wislizeni (G. Engelmann) R. OK Not provided Hixson 1940b Rollins (Sy ϭ Dithyrea wislizeni G. Engelmann) Lepidium virginicum C. Linnaeus var. medium TX Apr./May Reinhard 1927b (E. Greene) C. L. Hitchcock (Sy ϭ L. medium E. Greene) Lepidium virginicum C. Linnaeus TX Not provided Reinhard 1926a OK MayÐJuly Hixson 1940 AR, LA, MS May Snodgrass et al. 1984 Caprifoliaceae Symphoricarpos orbiculatus C. Moench IL Not provided Knight 1941 Caryophyllaceae Cerastium fontanum J. C. Baumgarten subsp. TX Not provided Reinhard 1926a vulgare (C. J. Hartman) W. Greuter and H. Burdet (Sy ϭ C. vulgatum C. Linnaeus Stellaria media (C. Linnaeus) D. Villars OK Not provided Hixson 1940 AR, LA, MS May Snodgrass et al. 1984 Chenopodiaceae Atriplex spp. TX Not provided Reinhard 1926a TX No sampling data Reinhard 1926b TX Overwintering Reinhard 1928 Atriplex argentea T. Nuttall OK Not provided Hixson 1940 Chenopodium spp. TX No sampling data Reinhard 1926b Chenopodium album C. Linnaeus var. album TX Not provided Reinhard 1926b OK Spring-summer Hixson 1940 Chenopodium incanum (S. Watson) A. A. OK Not provided Hixson 1940 Heller Chenopodium leptophyllum (C. Moquin- OK Not provided Hixson 1940 Tandon) T. nuttall ex S. Watson Dysphania ambrosioides (C. Linnaeus) S. OK Summer Hixson 1940 Mosyakin and SE Clemants (Sy ϭ Chenopodium ambrosioides C. Linnaeus) Monolepis nuttalliana (J. A. Schultes) E. TX Apr./May Reinhard 1927 Greene Clusiaceae Hypericum mutilum C. Linnaeus OK Hixson 1940 Convolvulaceae Ipomoea spp. TX No sampling data Reinhard 1926b Ipomoea cordatotriloba A. Dennstaedt var. TX Not provided Reinhard 1926ab torreyana (A. Gray) D. Austin (Sy ϭ I. triÞda (K. Kunth) G. Don var. torreyana (A. Gray) Shinners) Ipomoea leptophylla J. Torrey OK Not provided Hixson 1940 Cucurbitaceae Citrullus lanatus (C. Thunberg) J. Matsumura TX Not provided Reinhard 1926ab and T. Nakai var. lanatus (Sy ϭ C. citrullus (C. Linnaeus) G. C. W. H. Karsten Euphorbiaceae Chamaesyce maculata (C. Linnaeus) J. K. AR, LA, MS July Snodgrass et al. 1984b Small (Sy ϭ Euphorbia maculata C. Linnaeus) Croton spp. AZ Not provided Knight 1926, 1941 TX No sampling data Reinhard 1926b SC Overwintering Eddy 1928 TX No sampling data Fletcher 1930 TX Fall and overwintering Harned 1933 OK No sampling data Brett 1946 TX Lab study Gaylor 1975 TX No sampling data Beerwinkle and Marshall 1999 Croton capitatus A. Michaux (Sy ϭ C. TX No sampling data Hunter 1926b engelmannii A. M. Ferguson var. albinoides CA, LA, KS, MN, Not provided Knight 1926 A. M. Ferguson) MS, MO TX TX No sampling data Reinhard 1926a AL, AR, GA, MS Overwintering Reinhard 1927, 1928 IL Sept. laboratory specimens Knight 1927 TX TX Not provided Knight 1941

Continued on following page June 2009 ESQUIVEL AND ESQUIVEL:HOSTS OF THE COTTON FLEAHOPPER 775

Table 5. Continued

Plant family and species Study location Sampling perioda Reference TX Early spring Reinhard 1929 TX Apr.ÐNov. Gaines 1933 OK Fall/overwintering Gaines and Ewing 1938 TX Late summer Fletcher 1940 AR, LA, MS Late summer Hixson 1940, 1941 TX Season-long/overwintering Almand 1974 Almand et al. 1976 TX June/Oct. Snodgrass et al. 1984 Laboratory rearing Breene et al. 1989 Mid-, late season Current study Croton capitatus A. Michaux var. lindheimeri TX Fall/overwintering Schuster et al. 1969b (G. Engelmann and A. Gray) J. Mller of Aargau (Sy ϭ C. capitatus A. Michaux var. albinoides (Ferguson) Shinners) Croton capitatus A. Michaux var. lindheimeri TX No sampling data Hunter 1926b (G. Engelmann and A. Gray) J. Mller of TX Sept. Knight 1926b Aargau (Sy ϭ C. engelmannii A. Ferguson) TX No sampling data Reinhard 1926ab Croton glandulosus C. Linnaeus SC Year-round Eddy 1927 Croton glandulosus C. Linnaeus var. OK Not provided Hixson 1940 septentrionalis J. Mller of Aargau Croton lindheimerianus G. Scheele TX May/Sept. Knight 1926 TX Not provided Reinhard 1926a OK Late summer Hixson 1940, 1941 Croton monanthogynus A. Michaux TX No sampling data Hunter 1926 TX Aug. Knight 1926 TX Not provided Reinhard 1926a OK JulyÐSept. Hixson 1940 Croton pottsii (J. Klotzch) J. Mller of Aargau AZ Aug. Knight 1926b var. pottsii (Sy ϭ C. corymbulosus G. Engelmann) Croton punctatus N. von Jacquin TX Sept. Knight 1926 Croton texensis (J. Klotsch) J. Mller of TX No sampling data Hunter 1926 Aargau TX Not provided Reinhard 1926a OK Late summer Hixson 1940, 1941 IL Not provided Knight 1941 Euphorbia bicolor G. Engelmann and A. Gray OK Not provided Hixson 1940 Euphorbia marginata F. Pursh OK July Hixson 1940 Fabaceae Chamaecrista fasciculata (A. Michaux) E. AR, LA, MS Fall Snodgrass et al. 1984 Greene (Sy ϭ Cassia fasciculata A. TX Mid-season Current study Michaux) Glycine max (C. Linnaeus) E. Merrill AR, LA, MS Snodgrass et al. 1984 TX Mid-season Current study Hoffmannseggia glauca (C. Ortega) I. Eifert OK Not provided Hixson 1940 (Sy ϭ H. falcaria A. Cavanilles) Medicago polymorpha C. Linaneus (Sy ϭ M. OK Not provided Hixson 1940 hispida J. Gaertner) TX Early season Current study Medicago sativa C. Linnaeus OK Not provided Hixson 1940 TX Early season Schuster et al. 1969 TX Early, mid-, late season Current study Melilotus indicus (C. Linnaeus) C. Allioni TX Early-season Current study Melilotus ofﬁcinalis (C. Linnaeus) J. de TX Not provided Reinhard 1926a Lamarck Phaseolus spp. SC Early/mid season Eddy 1927 TX Laboratory study Gaylor 1975 TX Laboratory study Gaylor and Sterling 1975, 1976a Rapistrum rugosum (C. Linnaeus) C. Allioni TX Early season Esquivel 2005, 2008 TX Apr.ÐMay Westbrook et al. 2005 TX Early season Current study Trifolium incarnatum C. Linnaeus AR, LA, MS May Snodgrass et al. 1984 TX Early season Current study Trifolium polymorphum J. Poiret ex J. de TX Apr./May Reinhard 1927 Lamarck and J. Poiret (Sy ϭ T. amphianthum J. Torrey and A. Gray) Vicia sativa C. Linnaeus subsp. nigra (C. OK Not provided Hixson 1940b Linnaeus) J. F. Ehrhart (Sy ϭ V. AR, LA, MS May Snodgrass et al. 1984b angustifolia C. Linnaeus) Vigna unguiculata (C. Linnaeus) W. Walpers TX Not provided Reinhard 1926ab subsp. unguiculata (Sy ϭ V. sinensis (C. Linnaeus) G. Savi) Geraniaceae Erodium cicutarium (C. Linnaeus) LÕHritier TX Apr./May Idol and Slosser 2005 de Brutelle ex W. Aiton

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Table 5. Continued

Plant family and species Study location Sampling perioda Reference Geranium spp. TX No sampling data Reinhard 1926b Geranium carolinianum C. Linnaeus OK Not provided Hixson 1940 TX Not provided Reinhard 1926a Geranium dissectum C. Linnaeus AR, LA, MS May Snodgrass et al. 1984 Hydrophyllaceae Nama hispidum A. Gray TX Early season Schuster et al. 1969 Lamiaceae Calamintha arkansana (T. Nuttall) L. Shinners OK MayÐJuly Hixson 1940b (Sy ϭ C. nuttallii G. Bentham) Lamium spp. TX No sampling data Reinhard 1926b Lamium amplexicaule C. Linnaeus TX Not provided Reinhard 1926a OK Not provided Hixson 1940 Marrubium vulgare C. Linnaeus TX May Reinhard and Owen 1928 OK Not provided Hixson 1940 Monarda spp. TX No sampling data Reinhard 1926b TX Overwintering Reinhard 1928 TX Early spring Harned 1933 TX Not provided Thomas 1936 TX May/June Fletcher 1940 TX Lab study Beerwinkle and Marshall 1999 Monarda citriodora V. de Cervantes ex M. TX Apr./May Knight 1926 Lagasca y Segura TX Mar./Apr./May/June Schuster et al 1969 TX Mid-season Current study Monarda citriodora V. de Cervantes ex M. OK Late spring/early summer Hixson 1940,b 1941b Lagasca y Segura var. citriodora (Sy ϭ M. dispersa J. K. Small; Sy ϭ M. tenuiaristata A. Gray ex J. K. Small) Monarda clinopodioides A. Gray TX No sampling data Hunter 1926 OK Not provided Hixson 1940 Monarda fistulosa C. Linnaeus TX Not provided Reinhard 1926a OK Late spring/early summer Hixson 1940, 1941 Monarda fistulosa C. Linnaeus var. mollis G. OK MayÐJune Hixson 1940b Bentham (Sy ϭ M. mollis C. Linnaeus) Monarda punctata C. Linnaeus TX Not provided Reinhard 1926a TX Late summer Hixson 1940, 1941 IL Not provided Knight 1941 TX Summer Almand 1974 Almand et al. 1976 TX Laboratory study Gaylor 1975 TX Laboratory study Gaylor and Sterling 1976a, b TX May Idol and Slosser 2005 Monarda punctata C. Linnaeus var. lasiodonta OK Not provided Hixson 1940 A. Gray (Sy ϭ M. lasiodonta (A. Gray) J. K. TX Apr./May/June Schuster et al. 1969b Small; Sy ϭ M. punctata C. Linnaeus var. coryi (E. McClintock and C. Epling) V. Cory ex Shinners) Monarda russeliana T. Nuttall ex J. Sims OK MayÐJune Hixson 1940 Prunella vulgaris C. Linnaeus OK Not provided Hixson 1940 Pycnanthemum albescens J. Torrey and A. Gray OK JulyÐOct. Hixson 1940 Salvia reflexa J. Hornemann (Sy ϭ S. lancifolia OK Not provided Hixson 1940b J. Poiret) Stachys cordata J. Riddell TX Not provided Reinhard 1926a Stachys tenuifolia C. von Willdenow OK JulyÐSept. Hixson 1940 Teucrium canadense C. Linnaeus OK JuneÐJuly Hixson 1940 Lythraceae Lythrum alatum F. Pursh var. lanceolatum (S. AR, LA, MS July Snodgrass et al. 1984 Elliot) J. Torrey and A. Gray ex J. Rothrock (Sy ϭ L. lanceolatum S. Elliot) Malvaceae Abelmoschus esculentus (C. Linnaeus) C. TX Not provided Reinhard 1926ab Moench (Sy ϭ Hibiscus esculentus C. Linnaeus) Abutilon malacum S. Watson TX May Reinhard and Owen 1928 Callirho involucrata (T. Nuttall ex J. Torrey TX April/May Reinhard 1927 and A. Gray) A. Gray var. involucrata Gossypium herbaceum C. Linnaeus TX Not provided Reinhard 1926a Gossypium hirsutum C. Linnaeus TX Circumstantial data MayÐ Howard 1898 Aug. TX No sampling data Hunter 1926 MS, NE, TX, IL Not provided Knight 1926, 1941 SC TX Late summer Eddy 1927 LA Overwintering Reinhard 1927, 1928 TX Lab study King and Cook 1932 OK Not provided Thomas 1936

Continued on following page June 2009 ESQUIVEL AND ESQUIVEL:HOSTS OF THE COTTON FLEAHOPPER 777

Table 5. Continued

Plant family and species Study location Sampling perioda Reference TX Not provided Hixson 1940 TX Summer Almand 1974 Almand et al. 1976 TX Lab study Gaylor 1975 AR, LA, MS Lab study Gaylor and Sterling 1976a, b TX JuneÐAug. Snodgrass et al. 1984 Not provided Idol and Slosser 2005 Malvastrum americanum (C. Linnaeus) J. TX No sampling data Hunter 1926 Torrey Malvaviscus drummondii J. Torrey and A. Gray TX Not provided Reinhard 1926a Modiola caroliniana (C. Linnaeus) G. Don TX Apr./May Reinhard 1927 Nyctaginaceae Boerhavia coccinea P. Miller TX Not provided Reinhard 1926a Oleaceae Ligustrum sp. OK Hixson 1940 Onagraceae Calylophus serrulatus (T. Nuttall) P. Raven OK Not provided Hixson 1940b (Sy ϭ Oenothera serrulata T. Nuttall var. TX Early spring Schuster et al. 1969b serrulata) Gaura brachycarpa J. K. Small TX Overwintering Reinhard 1928 TX May Reinhard and Owen 1928 TX Early season Schuster et al. 1969 Gaura coccinea T. Nuttall ex F. Pursh TX June/July Knight 1926 OK Not provided Hixson 1940 TX Mid-season Current study Gaura longiﬂora ƒ. Spach (Sy ϭ G. biennis C. TX May Reinhard and Owen 1928b Linnaeus var. pitcheri J. Torrey and A. Gray) Gaura sinuata T. Nuttall ex N. Sringe TX Apr./May Reinhard 1927 TX Apr./May Idol and Slosser 2005 Gaura tripetala A. Cavanilles TX Apr./May Reinhard 1927 Gaura villosa J. Torrey OK Not provided Hixson 1940 TX Early season Schuster et al. 1969 TX Apr./May Idol and Slosser 2005 Oenothera spp. TX No sampling data Reinhard 1926b SC AprilÐJune Eddy 1928 TX Not provided Thomas 1936 OK No sampling data Brett 1946 Oenothera biennis C. Linnaeus SC Early/mid season Eddy 1927 AR, LA, MS Aug. Snodgrass et al. 1969 Oenothera grandis (N. Britton) B. Smyth TX Apr./May/June Idol and Slosser 2005 Oenothera laciniata J. Hill SC Early-season Eddy 1927 TX Apr./May Reinhard 1927 TX Early spring Fletcher 1940 OK Early spring Hixson 1940, 1941 TX Lab study Gaylor 1975 TX Lab study Gaylor and Sterling 1976a, 1976b AR, LA, MS May/June/July Snodgrass et al. 1984 TX Lab studies Beerwinkle and Marshall 1999 TX April/May/June Idol and Slosser 2005 Oenothera laciniata J. Hill var. laciniata A. TX Early spring Schuster et al. 1969 Griseback TX Early-season Almand 1974 Almand et al. 1976 Oenothera rhombipetala T. Nuttall ex J. Torrey OK Not provided Hixson 1940 and A. Gray Oenothera speciosa T. Nuttall (Sy ϭ O. TX Early spring Fletcher 1940 speciosa T. Nuttall var. speciosa) TX Early spring Schuster et al. 1969b AR, LA, MS May/July Snodgrass et al. 1984 TX Early-season Current study Oenothera speciosa T. Nuttall (Sy ϭ O. TX Early spring Schuster et al. 1969b speciosa T. Nuttall var. childsii (L. H. Bailey) P. Munz) Oenothera villosa C. Thunberg ssp. strigosa (P. OK Not provided Hixson 1940b Rydberg) W. Dietrich and P. H. Raven (Sy ϭ O. strigosa (P. Rydberg) K. Mackenzie and B. Bush) Oxalidaceae Oxalis stricta C. Linnaeus OK Early spring Hixson 1940 POACEAE Bromus sp. OK Not provided Hixson 1940 Sphenopholis obtusata (A. Michaux) F. OK Not provided Hixson 1940 Lamson-Scribner var. major (Sy ϭ S. intermedia (P. Rydberg) P. Rydberg) Urochloa texana (S. Buckley) R. D. Webster TX Not provided Reinhard 1926a (Sy ϭ Panicum texanum S. Buckley) OK Not provided Hixson 1940b

Continued on following page 778 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 3

Table 5. Continued

Plant family and species Study location Sampling perioda Reference Polygonaceae Eriogonum annuum T. Nuttall OK Not provided Hixson 1940 Polygonum spp. SC Fall Eddy 1927 Polygonum amphibium C. Linnaeus var. AR, LA, MS Sept. Snodgrass et al. 1984b emersum A. Michaux (Sy ϭ P. coccineum G. H. Muhlenberg ex C. von Willdenow Polygonum aviculare C. Linnaeus AR, LA, MS Aug. Snodgrass et al. 1984 Polygonum hydropiperoides A. Michaux AR, LA, MS Aug. Snodgrass et al. 1984 Polygonum lapathifolium C. Linnaeus AR, LA, MS Aug. Snodgrass et al. 1984 Polygonum pensylvanicum C. Linnaeus AR, LA, MS JulyÐSept. Snodgrass et al. 1984 Rumex spp. TX Apr./May Reinhard 1927 Rumex acetosella C. Linnaeus OK Early spring Hixson 1940 Rumex hastatulus W. Baldwin OK Not provided Hixson 1940 Portulacaceae Portulaca spp. TX No sampling data Reinhard 1926b Portulaca oleracea C. Linnaeus TX Not provided Reinhard 1926a AL Not provided Romm 1937 Ranunculaceae Ranunculus parviflorus C. Linnaeus OK Early spring Hixson 1940, 1941 Rosaceae Rubus occidentalis C. Linnaeus OK Not provided Hixson 1940 Rubiaceae Galium aparine C. Linnaeus OK Hixson 1940 Scrophulariaceae Castilleja indivisa G. Engelmann TX April/May Reinhard 1927 Solanaceae Petunia axillaris (J. de Lamarck) N. Britton, E. OK Not provided Hixson 1940 Sterns, and J. Poggenburg (Sy ϭ Petunia x hybrida P. L. de Vilmorin) Solanum spp. SC Early/mid-season Eddy 1927 TX No sampling data Reinhard 1926b OK No sampling data Brett 1946 TX Lab study Gaylor 1975 Solanum carolinense C. Linnaeus OK JulyÐOct. Hixson 1940 Solanum dimidiatum C. RaÞnesque-Schmaltz OK Not provided Hixson 1940 (Sy ϭ S. torreyi A. Gray) Solanum elaeagnifolium A. Cavanilles TX Not provided Reinhard 1926a TX Overwintering Reinhard 1927, 1928a OK Late spring/early summer Hixson 1940, 1941 AR, LA, MS June Snodgrass et al. 1984 TX Late summer/early fall Schuster et al. 1969 TX Overwintering Idol and Slosser 2005 TX Early, mid-, late season Current study Solanum rostratum M. Dunal TX Not provided Reinhard 1926a OK JulyÐOct. Hixson 1940, 1941 Solanum tuberosum C. Linnaeus TX Laboratory studies Gaylor and Sterling 1975, 1976a OK MayÐJune Hixson 1940, 1941 Verbenaceae Glandularia bipinnatifida (T. Nuttall) T. TX Early season Schuster et al. 1969 Nuttall var. bipinnatifida Glandularia bipinnatifida (T. Nuttall) T. TX Early season Schuster et al. 1969 Nuttall var. ciliata (G. Bentham) B. L. Turner Phyla nodiflora (C. Linnaeus) E. Greene TX Not provided Reinhard 1926ab (Sy ϭ Lippia nodißora (J. de Lamarck) A. TX Mid-, late season Current study Michaux) Verbena litoralis K. Kunth (Sy ϭ V. brasiliensis AR, LA, MS Sept. Snodgrass et al. 1984 J. Velloso de Miranda) Verbena neomexicana (A. Gray) J. K. Small var. TX Mid-, late season Current study hirtella L. Perry Verbena urticifolia C. Linnaeus OK MayÐJuly Hixson 1940 Zygophyllaceae Kallstroemia hirsutissima A. Vail ex J. K. Small TX Not provided Reinhard 1926a Tribulus terrestris C. Linnaeus TX Not provided Reinhard 1926a OK Not provided Hixson 1940

a Designation of ЉNot providedЉϭhost plants only listed; ЉNo sampling dataЉϭobservations were made but no sampling data provided. b Studies reporting the indicating taxonomical synonym. AR, Arkansas; AZ, Arizona; IL, Illinois; LA, Louisiana; MS, Mississippi; NE, Nebraska; OK, Oklahoma; SC, South Carolina; TX, Texas. hybridus)to1.0[B. pilosa (T. Nuttall)] during 2003 tall ex Pursh). M. polymorpha exhibited the highest (Table 4). During 2004, the range for proportion of proportion of females (0.83) during 2005, whereas females was from 0 [Chamaecrista fasciculata (A. females were not encountered in R. rugosum. For the Michaux) E. Greene] to 0.69 (Gaura coccinea T. Nut- previously reported preferred hosts in the Oenothera, June 2009 ESQUIVEL AND ESQUIVEL:HOSTS OF THE COTTON FLEAHOPPER 779

Monarda, Solanum, and Croton genera, the proportion a tool for early-season control of insect populations of females were relatively high with the exception of (Thomas and Owen 1937; Fleischer and Gaylor 1987). O. speciosa during 2005. Sex ratio and ovipositional The variability of seasonal hosts, identiÞcation of eight preference is affected by the species and maturity of previously unreported weed hosts, and logistics (viz. the host plant (Gaylor 1975, Holtzer and Sterling weed densities, weed distribution, and ongoing boll 1980), and our Þndings concur with Holtzer and Ster- weevil eradication efforts) encountered in our study ling (1980). region suggested that weed suppression as a manage- To clarify the status of plant species sampled during ment tool in Central Texas would require further this study, a review of the available literature from study. 1898 through 2008 is presented in Table 5. Table 5 reßects reports identifying plant species sampled in Acknowledgments Þeld conditions as well as those from laboratory reports evaluating the effects of uncultivated host plants G. D. Jones (Research Biologist, USDAÐARS, Areawide on cotton ßeahopper behavior and biology. Plant spe- Pest Management Research Unit) provided clariÞcations for cies sampled for this study were incorporated in Table the botanical nomenclature, and her efforts are greatly ap- 5, providing a current listing of host plants related to preciated. the cotton ßeahopper. In total, 169 species of plants distributed among 35 families have been identiÞed as References Cited hosts for the cotton ßeahopper. The leading plant families with predominant hosts were the Asteraceae Almand, L. K. 1974. Seasonal abundance, dispersal and con- (39 identiÞed species), Lamiaceae (17 identiÞed spe- trol of the cotton ßeahopper on certain host plants. PhD dissertation, Texas A&M University, College Station, TX. cies), and Onagraceae (14 identiÞed species). Some Almand, L. K., W. L. Sterling, and C. L. Green. 1976. Sea- previous reports only reported common names. sonal abundance and dispersal of the cotton ßeahopper as Where this occurred, this report assigned the common related to host plant phenology. Tex. Agric. Exp. Sta. Bull. names to genera in Table 5, but these assignments are 1170, Texas A&M University, College Station, TX. not included in the number of identiÞed species re- Anderson, R. A., and M. F. Schuster. 1983. Phenology of the ported. Twenty-seven plant taxa were assigned to gen- tarnished plant bug on natural host plants in relation to era only. populations in cotton. Southwest. Entomol. 8: 131Ð136. The composition of plant species sampled in this Armstrong, J. S., and L. D. Camelo. 2003. Lygus species study included the commonly accepted preferred associated with Texas High Plains cotton, alfalfa and weeds. Southwest. Entomol. 28: 197Ð204. hosts (primroses, nightshades, horsemints, and cro- Beerwinkle, K. R., and H. F. Marshall. 1999. Cotton ßea- tons) and other available weed species. Eight previ- hopper (Heteroptera: Miridae) responses to volatiles ously unreported host plants for the cotton ßeahopper from selected host plants. J. Cotton Sci. 3: 153Ð159. were identiÞed in the Southern Blacklands region of Breene, R. G., W. R. Martin, Jr., D. A. Dean, and W. L. Central Texas. Our results further indicated that some Sterling. 1989. Rearing methods for the cotton ßeahop- of these new hosts may harbor cotton ßeahopper per. Southwest. Entomol. 14: 249Ð253. adults before previously reported preferred hosts are Brett, C. H. 1946. The cotton ßea hopper in Oklahoma. available. Furthermore, temporal captures of cotton Okla. Agric. Exp. Sta. Bull. B-303, Oklahoma Agricultural ßeahoppers indicated multiple previously unreported and Mechanical College, Stillwater, OK. Cleveland, T. C. 1982. Hibernation and host plant sequence hosts. In some instances, these hosts were leading studies of tarnished plant bugs, Lygus lineolaris, in the weed species harboring ßeahoppers during respective Mississippi Delta. Environ. Entomol. 11: 1049Ð1052. sampling periods. The identiÞcation of these eight Eddy, C. O. 1927. The cotton ßea hopper. S. C. Agric. Exp. Sta. hosts supplements the compendium of previously Clemson College Bull. 235, Clemson College, Clemson, SC. identiÞed plant species inßuencing the development Eddy, C. O. 1928. Cotton ßea hopper studies of 1927 and of cotton ßeahopper populations (Table 5). 1928. S. C. Agric. Exp. Sta. of Clemson College Bull. 251, Of particular interest for future cotton ßeahopper Clemson College, Clemson, SC. host plant studies is the proposed revision to the cot- Esquivel, J. F. 2005. Occurrence of the cotton ßeahopper in a ton ßeahopper nomenclature (Wheeler 2001). The previously unreported early-season host in Central Texas. Jan. 4Ð7, 2005. 2005 Proceedings of the Beltwide Cotton proposed revision involves changing the speciÞc ep- Conference, National Cotton Council, Memphis, TN. ithet from seriatus to seriata resulting in Pseudatomos- Esquivel, J. F. 2008. Observations and implications of the celis seriata as a more correct identiÞcation for the cotton ßeahopper (Heteroptera: Miridae) in Rapistrum cotton ßeahopper. This proposed change should be rugosum (Capparales: Brassicacea). J. Entomol. Sci. 43: monitored to ensure accurate identiÞcation of insectÐ 434Ð437. host plant associations for the cotton ßeahopper. Fleischer, S. J., and M. J. Gaylor. 1987. Seasonal abundance Findings presented here and the compendium of of Lygus lineolaris (Heteroptera: Miridae) and selected host plants for the cotton ßeahoppers provide a cur- predators in early-season uncultivated hosts: implications rent and taxonomically correct listing of host plants for managing movement into cotton. Environ. Entomol. 16: 379Ð389. associated with the cotton ßeahopper. These associ- Fletcher, R. K. 1930. A study of the insect fauna of Brazos ations may be exploited to provide an alternative for County, Texas, with special reference to the Cicadellidae. managing this insect before the ßeahopper infests cot- Ann. Entomol. Soc. Am. 23: 33Ð56. ton or begins late-season oviposition. Indeed, areaw- Fletcher, R. K. 1940. Certain host plants of the cotton ßea ide management of weed hosts has been proposed as hopper. J. Econ. Entomol. 33: 456Ð459. 780 ENVIRONMENTAL ENTOMOLOGY Vol. 38, no. 3

Gaines, J. C. 1933. A study of the cotton ßea hopper with Layton, M. B., J. L. Long, S. G. Flint, and L. M. Green. 2003. special reference to the spring emergence, dispersal, and Control of tarnished plant bug in Mississippi Delta cotton. population. J. Econ. Entomol. 26: 963Ð971. Proceedings of the 2003 Beltwide Cotton Conferences, Gaines, J. C., and K. P. Ewing. 1938. The relation of wind Nashville, TN, 6Ð10 January 2003. currents, as indicated by balloon drifts, to cotton ßea Reinhard, H. J. 1926a. The cotton ßea hopper. Texas Agri. hopper dispersal. J. Econ. Entomol. 31: 674Ð677. Exp. Sta. 57Ð426-12M Bull. 339, Agricultural and Mechan- Gaylor, M. J. 1975. Effects of temperature, host plants, rainfall, ical College of Texas, College Station, TX. and photoperiod on the population dynamics of the cotton Reinhard, H. J. 1926b. Control of the cotton ßea hopper in ßeahopper, Pseudatomoscelis seriatus (Reuter). PhD disser- Texas. Texas Agri. Exp. Sta. Circ. No. 40, Agricultural and tation, Texas A&M University, College Station, TX. Mechanical College of Texas, College Station, TX. Gaylor, M. J., and W. L. Sterling. 1975. Effects of temper- Reinhard, H. J. 1927. Control and spring emergence of the ature on the development, egg production, and survival cotton ßea hopper. Texas Agri. Exp. Sta. 225Ð227-25M- of the cotton ßeahopper, Pseudatomoscelis seriatus. En- L180 Bull. 356, Agricultural and Mechanical College of viron. Entomol. 4: 487Ð490. Texas, College Station, TX. Gaylor, M. J., and W. L. Sterling. 1976a. Effects of temper- Reinhard, H. J. 1928. Hibernation of the cotton ßea hopper. ature and host plants on population dynamics of the Tex. Agric. Exp. Stn. A3Ð128-12,000-L180 Bull. 377, Agricul- cotton ßeahopper, Pseudatomoscelis seriatus. Tex. Agric. tural and Mechanical College of Texas, College Station, TX. Exp. Sta. Bull. B-1161, Texas A&M University, College Reinhard, H. J. 1929. The value of spring emergence Station, TX. records on the cotton ßea hopper, Psallus seriatus. J. Econ. Gaylor, M. J., and W. L. Sterling. 1976b. Development, sur- Entomol. 22: 765Ð768. vival, and fecundity of the cotton ßeahopper, Pseudato- Reinhard, H. J., and W. L. Owen, Jr. 1928. Investigations on moscelis seriatus (Reuter), on several host plants. control of cotton ßea hopper in 1927. Tex. Agric. Exp. Stn. Environ. Entomol. 5: 55Ð58. A8Ð428-9,000-L180 Bull. 380, Agricultural and Mechan- Harned, R. W. 1933. Cotton ßea hopper, pp. 26Ð27. In Re- ical College of Texas, College Station, TX. port to the Chief of the Bureau of Entomology in the Romm, H. J. 1937. The insect depredators of purslane. Fla. Annual Reports of Department of Agriculture, 1933. U.S. Entomol. 20: 44Ð61. Department of Agriculture, Bureau of Entomology. 47 pp. SAS Institute. 2004. SAS 9.1 for Windows. SAS Institute, Hatch, S. L., K. N. Gandhi, and L. E. Brown. 1990. Checklist Cary, NC. Schuster, M. F., C. A. Richmond, J. C. Boling, and H. M. of the vascular plants of Texas MP-1655, Texas A&M Graham. 1969. Host plants of the cotton ßeahopper in University, College Station, TX. the Rio Grande Valley: phenology and hibernating quar- Hixson, E. 1940. The host relation of the cotton ßea hopper ters. J. Econ. Entomol. 62: 1126Ð1129. (Psallus seriatus Reuter). PhD dissertation, Iowa State Snodgrass, G. L., W. P. Scott, and J. W. Smith. 1984. A survey College, Ames, IA. of the host plants and seasonal distribution of the cotton Hixson, E. 1941. The host relation of the cotton ßea hopper. ßeahopper (Hemiptera: Miridae) in the Delta of Arkan- Iowa State College J. Sci. 16: 66Ð68. sas, Louisiana, and Mississippi. J. Georgia Entomol. Sci. 19: Holtzer, T. O., and W. L. Sterling. 1980. Ovipositional pref- 34Ð41. erence of the cotton ßeahopper, Pseudatomoscelis seria- Sterling, W. L., and D. A. Dean. 1977. A bibliography of the tus, and distribution of eggs among host plant species. cotton ßeahopper, Pseudatomoscelis seriatus (Reuter). Environ. Entomol. 9: 236Ð240. Tex. Agric. Exp. Sta. MP-1342, Texas A&M University, Howard, L. O. 1898. Cotton Þeld insects, pp. 85Ð88. In Some College Station, TX. miscellaneous results of the work of the division of en- Suh, C.P.C., D. W. Spurgeon, and A. E. Knutson. 2004. De- tomology III. U.S. Dep. Agric. Div. Entomol. Bull. 18 (new tecting cotton ßeahopper movement into Þelds with series), USDA, Washington, DC. sticky traps. Jan. 5Ð9, 2004. 2004 Proceedings of the Belt- Hunter, W. D. 1926. The cotton hopper, or so-called “cotton wide Cotton Conference, National Cotton Council, ßea”. U.S. Dep. Agric. Circ. 361, USDA, Washington DC. Memphis, TN. Idol, G. B., and J. E. Slosser. 2005. Spring emergence and Thomas, F. L. 1936. Control of the cotton ßea hopper. Tex. development of the cotton ßeahopper (Heteroptera: Agric. Exp. Stn. Cir. No. 77, Agricultural and Mechanical Miridae) in host plants of the Texas Rolling Plains. South- College of Texas, College Station, TX. west. Entomol. 30: 73Ð84. Thomas, F. L., and W. L. Owen, Jr. 1937. Cotton ßea hopper, Jones, S. D., J. K. Wipff, and P. M. Montgomery. 1997. Vascular an ecological problem. J. Econ. Entomol. 30: 848Ð851. plants of Texas. University of Texas Press, Austin, TX. Westbrook, J. K., J. F. Esquivel, and R. S. Eyster. 2005. Early- King, W. V., and W. S. Cook. 1932. Feeding punctures of season dispersal of cotton ßeahoppers (Pseudatomoscelis mirids and other plant-sucking insects and their effect on seriatus). Jan. 4Ð7, 2005. 2005 Proceedings of the Beltwide cotton. U. S. Dep. Agric. Tech. Bull. No. 296, USDA, Cotton Conference, National Cotton Council, Memphis, TN. Washington DC. Wheeler, A. G. 2001. Format and scope of review, pp. 6Ð10. In Knight, H. H. 1926. On the distribution and host plants of the Biology of the plant bugs (Hemiptera: Miridae): pests, pred- cotton ßea-hopper (Psallus seriatus Reuter) Hemiptera, ators, opportunists. Cornell University Press, Ithaca, NY. Miridae. J. Econ. Entomol. 19: 106Ð108. Womack, C. L., and M. F. Schuster. 1987. Host plants of the Knight, H. H. 1927. Notes on the distribution and host tarnished plant bug (Heteroptera: Miridae) in the North- plants of some North American Miridae (Hemiptera). ern Blackland Prairies of Texas. Environ. Entomol. 16: Can. Entomol. 59: 34Ð44. 1266Ð1272. Knight, H. H. 1941. The plant bugs, or Miridae, of Illinois. Ill. Nat. Hist. Surv. Bull. 22: 1Ð234. Received 28 July 2008; accepted 4 March 2009.