Bactericera cockerelli (tomato/potato psyllid)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Species Vectored
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Risk and Impact Factors
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Bactericera cockerelli (Šulc) 1909
Preferred Common Name
- tomato/potato psyllid
Other Scientific Names
- Paratrioza cockerelli (Šulc) 1909
- Trioza cockerelli (Šulc) 1909
International Common Names
- English: potato, psyllid; tomato psyllid; tomato, psyllid
- Spanish: pulgon saltador de la papa (mexico); pulgon saltador de la tomato (mexico)
- French: psylle de la pomme de terre; psylle de la tomate
Local Common Names
- Germany: Blattsauger, Amerikanischer Kartoffel-; Blattsauger, Tomaten-
- PARZCO (Paratrioza cockerelli)
Summary of InvasivenessTop of page
B. cockerelli is one of the most destructive potato pests in the western hemisphere. It was recognized in the early 1900s that B. cockerelli had the potential to be an invasive and harmful insect, particularly in western United States and Mexico (Šulc, 1909; Crawford, 1914; Compere, 1915; 1916; Essig, 1917). By the 1920s and 1930s, B. cockerelli had become a serious and destructive pest of potatoes in most of the southwestern United States, giving rise to the description of a new disease that became known as ‘psyllid yellows’ (Richards, 1928; 1931; 1933; Binkley, 1929; Richards and Blood, 1933; List and Daniels, 1934; Pletsch, 1947; Wallis, 1955).
In recent years, other solanaceous crops, including tomato, pepper, eggplant, tobacco and tamarillo in a number of geographic areas have suffered extensive economic losses associated with B. cockerelli outbreaks (Trumble, 2008, 2009; Munyaneza et al., 2007a,b; 2008; 2009a,b,c,d; Liefting et al., 2008; 2009; Secor et al., 2009; Espinoza, 2010; Munyaneza, 2010; Crosslin et al., 2010; Rehman et al., 2010; Crosslin et al., 2012a,b; Munyaneza, 2012).
Despite being a native of North America, B. cockerelli is also found in Central America and has recently invaded New Zealand, where it has caused extensive damage to indoor and outdoor solanaceous crops (Teulon et al., 2009; Thomas et al., 2011). B. cockerelli has recently been placed on the list of quarantine pest in EPPO region (EPPO, 2012).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hemiptera
- Suborder: Sternorrhyncha
- Unknown: Psylloidea
- Family: Triozidae
- Genus: Bactericera
- Species: Bactericera cockerelli
Notes on Taxonomy and NomenclatureTop of page
A brief history on the taxonomy and nomenclature of Bactericera cockerelli was recently provided by Butler and Trumble (2012).
B. cockerelli was originally described as Trioza cockerelli by Šulc (1909). In 1910, Crawford erected a new psyllid genus Paratrioza (Crawford, 1910) and Trioza cockerelli was assigned to this genus in 1911 (Crawford, 1911). In 1997, when the genus Paratrioza was reviewed and synonymized with the genus Bactericera, B. cockerelli also changed families from Psyllidae, subfamily Triozinae, to Triozidae (Burckhardt and Lauterer, 1997; Hodkinson, 2009). The subfamily Triozinae was elevated to family status as Triozidae.
Morphological descriptions of B. cockerelli can be found in Crawford (1911, 1914), Essig (1917), Ferris (1925) and Tuthill (1945). Also, a list of the synonyms of B. cokerelli is provided by Tuthill (1945) and Burckhardt and Lauterer (1997). Furthermore, a revised classification of pysllids was recently provided by Burckhardt and Ouvrard (2012).
DescriptionTop of page
B. cockerelli adults are small, measuring about 2.5-2.75 mm long. The adults generally resemble tiny cicadas, largely because they hold their wings angled and roof-like over their body. B. cockerelli adults possess two pairs of clear wings; the front wings bear conspicuous veins and are considerably larger than the hind wings. The antennae are moderately long, extending almost half the length of the body. The overall body colour ranges from pale green at emergence to dark green or brown within 2-3 days, and eventually becomes grey or black thereafter. Prominent white or yellow lines are found on the head and thorax, and dorsal whitish bands are located on the first and terminal abdominal segments. These white markings are spot characteristics of the psyllid, particularly the broad, transverse white band on the first abdominal segment and the inverted ‘V’-shaped white mark on the last abdominal segment (Pletsch, 1947; Wallis, 1955), along with the raised white line around the circumference of the head. Adults are active in contrast to the largely sedentary nymphal stages. These insects are good fliers and readily jump when disturbed.
The pre-oviposition period is normally about 10 days, with oviposition lasting up to 50 days. Total adult longevity ranges from 20 to 60 days, and females usually live two to three times longer than males (Pletsch, 1947; Abernathy, 1991; Abdullah, 2008; Yang and Liu, 2009). Females lay 300 to 500 eggs over their lifetime (Knowlton and Janes, 1931; Pletsch, 1947; Abdullah, 2008; Yang and Liu, 2009).
The eggs of B. cockerelli are deposited singly, principally on the lower surface of leaves and usually near the leaf edge, but some eggs can be found throughout suitable host plants. Often, females will lay numerous eggs on a single leaf. The eggs are initially light yellow and become dark yellow or orange with time. The eggs measure about 0.32-0.34 mm long, 0.13-0.15 mm wide, and are mounted on a stalk of about 0.48-0.51 mm. Eggs hatch 3-7 days after oviposition (Pletsch, 1947; Wallis, 1955; Capinera, 2001; Abdullah, 2008). Because nymphs prefer sheltered and shaded locations, they are mostly found on the lower surfaces of leaves and usually remain sedentary during their entire development. Nymphs and adults produce large quantities of whitish excrement particles, which may adhere to foliage and fruit.
Nymphs are elliptical when viewed from above, but are very flattened in profile, appearing almost scale-like. Potato psyllid nymphs may be confused with the nymphs of whiteflies, although the former move when disturbed. There are five nymphal instars, with each instar possessing very similar morphological features besides size. The size of the developing wingpads increases with each instar. Nymphal body widths are variable, ranging from 0.23 to 1.60 mm, depending on different instars (Rowe and Knowlton, 1935; Pletsch, 1947; Wallis, 1955). Initially, the nymphs are orange, but they become yellowish-green and then green as they mature. The compound eyes are reddish and quite prominent. During the third instar, the wing pads, light in colour, are evident and become more pronounced with each subsequent molt. A short fringe of wax filaments is present along the lateral margins of the body. Total nymphal development time depends on temperature and host plant and has been reported to have a range of 12 to 24 days (Knowlton and Janes, 1931; Abdullah, 2008; Yang and Liu, 2009).
DistributionTop of page
B. cockerelli is native to North America and occurs mainly in the Great Plains region of the United States, from Colorado, New Mexico, Arizona and Nevada, north to Utah. More recently, its range has expanded to include Wyoming, Idaho, Montana, California, Oregon, Washington, Alberta and Saskatchewan (Pletsch, 1947; Wallis, 1955; Cranshaw, 1993; Ferguson and Shipp, 2002; Ferguson et al., 2003). This insect pest is common in southern and western Texas and has also been documented in Oklahoma, Kansas, Nebraska, South Dakota, North Dakota, Minnesota and as far west as California and British Columbia. Contrary to reports in the older literature (Pletsch 1947; Wallis 1955; Cranshaw 1993, 2001), the potato psyllid does indeed occur in Washington and Oregon, where it appears to overwinter locally (Jensen, 2012) and usually colonizes potato fields in late June and early July (Munyaneza et al., 2009a; Munyaneza 2010; Crosslin et al., 2012a; Munyaneza, 2012). Murphy et al. (2013) reported B. cockerelli overwintering in the Pacific Northwest (north east Oregon, south east Washington state and southwestern Idaho).
Overwintering in areas north of the Texan or Mexican border is a recent development, as is psyllid infestation in southern Idaho and other northern parts of the current range. Before about 2004, potato psyllid was a migratory species, overwintering in northern Mexico and southern Texas and migrating into the Great Plains each summer. Texas, New Mexico, Arizona, Colorado and Nevada saw populations every year. Places farther north were colonized intermittently.
B. cockerelli also occurs in Mexico and Central America, including Guatemala and Honduras (Pletsch, 1947; Wallis, 1955; Rubio-Covarrubias et al., 2006; Trumble, 2008; 2009; Crosslin et al., 2010; Espinoza, 2010; Munyaneza, 2010; Rehman et al., 2010; Rubio-Covarrubias et al., 2011; Aguilar et al., 2013a; Angilar et al., 2013b; Munyaneza et al., 2013b; Munyaneza et al., 2014), and most recently was reported from Nicaragua (Munyaneza, 2012; Bextine et al., 2013b; Munyaneza et al., 2013a). B. cockerelli is also suspected to be present in neighbouring countries, including El Salvador (Bextine et al., 2012; Bextine et al., 2013a). There are no early records of B. cockerelli in Central America, and it is possible that Central America, as well as the northern portions of its current North American range, represents a newly-colonized area.
B. cockerelli is also widespread in New Zealand (Teulon et al., 2009).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Canada||Restricted distribution||EPPO, 2014; CABI/EPPO, 2015|
|-Alberta||Present, few occurrences||Introduced||Not invasive||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|-British Columbia||Present, few occurrences||Introduced||Not invasive||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|-Manitoba||Present, few occurrences||Introduced||Not invasive||Henne et al., 2010a; CABI/EPPO, 2015|
|-Ontario||Present only under cover/indoors||Introduced||Not invasive||Ferguson and Shipp, 2002; Ferguson et al., 2003; EPPO, 2014; CABI/EPPO, 2015|
|-Quebec||Present only under cover/indoors||Introduced||Not invasive||Ferguson et al., 2003; EPPO, 2014; CABI/EPPO, 2015|
|-Saskatchewan||Present, few occurrences||Introduced||Not invasive||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|Mexico||Present||Native||Invasive||Tuthill, 1945; Trumble, 2008; Rubio-Covarrubias et al., 2011; EPPO, 2014; CABI/EPPO, 2015|
|USA||Present||EPPO, 2014; CABI/EPPO, 2015|
|-Arizona||Widespread||Native||Invasive||Brown et al., 2010; EPPO, 2014; CABI/EPPO, 2015|
|-California||Widespread||Native||Invasive||Trumble, 2008; EPPO, 2014; CABI/EPPO, 2015|
|-Colorado||Widespread||Native||Invasive||Cranshaw, 1993; EPPO, 2014; CABI/EPPO, 2015|
|-Idaho||Widespread||Native||Invasive||Crosslin et al., 2012b; EPPO, 2014; CABI/EPPO, 2015|
|-Iowa||Present||EPPO, 2014; CABI/EPPO, 2015|
|-Kansas||Widespread||Native||Invasive||Goolsby et al., 2012; EPPO, 2014; CABI/EPPO, 2015|
|-Minnesota||Present, few occurrences||Native||Henne et al., 2012; EPPO, 2014; CABI/EPPO, 2015|
|-Montana||Widespread||Native||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|-Nebraska||Widespread||Native||Invasive||Goolsby et al., 2012; EPPO, 2014; CABI/EPPO, 2015|
|-Nevada||Widespread||Native||Invasive||Munyaneza et al., 2007a; EPPO, 2014; CABI/EPPO, 2015|
|-New Mexico||Widespread||Native||Invasive||Henne et al., 2012; EPPO, 2014; CABI/EPPO, 2015|
|-North Dakota||Present, few occurrences||Native||Henne et al., 2012; EPPO, 2014; CABI/EPPO, 2015|
|-Oklahoma||Present, few occurrences||Native||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|-Oregon||Widespread||Native||Invasive||Crosslin et al., 2012a; EPPO, 2014; CABI/EPPO, 2015|
|-South Dakota||Present, few occurrences||Native||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|-Texas||Widespread||Native||Invasive||Munyaneza et al., 2007a; EPPO, 2014; CABI/EPPO, 2015|
|-Utah||Widespread||Native||Invasive||Pletsch, 1947; EPPO, 2014; CABI/EPPO, 2015|
|-Washington||Widespread||Native||Invasive||Munyaneza et al., 2009a; EPPO, 2014; CABI/EPPO, 2015|
|-Wisconsin||Present, few occurrences||Introduced||Henne et al., 2012; CABI/EPPO, 2015|
|-Wyoming||Widespread||Native||Invasive||Wallis, 1955; EPPO, 2014; CABI/EPPO, 2015|
Central America and Caribbean
|El Salvador||Present||Native||Bextine et al., 2012; Bextine et al., 2013; EPPO, 2014; CABI/EPPO, 2015|
|Guatemala||Widespread||Native||Invasive||Munyaneza, 2012; EPPO, 2014; CABI/EPPO, 2015|
|Honduras||Widespread||Native||Invasive||Espinoza, 2010; Aguilar et al., 2013; Aguilar et al., 2013; Munyaneza et al., 2013; EPPO, 2014; Munyaneza et al., 2014; CABI/EPPO, 2015|
|Nicaragua||Widespread||Native||Invasive||Munyaneza, 2012; Bextine et al., 2013; Munyaneza et al., 2013; EPPO, 2014; CABI/EPPO, 2015|
|Australia||Present, few occurrences||Introduced||EPPO, 2017|
|-Western Australia||Present, few occurrences||Introduced||EPPO, 2017|
|New Zealand||Widespread||Introduced||Invasive||Gill, 2006; Teulon et al., 2009; Thomas et al., 2011; EPPO, 2014; CABI/EPPO, 2015|
|Norfolk Island||Present||CABI/EPPO, 2015|
History of Introduction and SpreadTop of page
In North America, driven primarily by wind and hot temperatures in late spring, B. cockerelli annually migrates from its overwintering and breeding areas in southern and western Texas, southern New Mexico, Arizona, California, Idaho and northern Mexico (Pletsch, 1947; Wallis, 1955). The migration occurs especially through the midwestern states and Canadian provinces along the Rocky Mountains (Romney, 1939; Pletsch, 1947; Jensen, 1954; Wallis, 1955). In these regions, damaging outbreaks of potato psyllid in potatoes and tomatoes occurred at regular intervals beginning in the late1800s and extending into the 1940s (List, 1939; Wallis, 1946; Pletsch, 1947). In more recent years, psyllid outbreaks have also occurred in regions outside of the midwestern United States, including southern California, Baja California, Oregon, Washington, Idaho, and Central America (Trumble, 2008; 2009; Munyaneza et al., 2009a; Wen et al., 2009; Crosslin et al., 2010; Munyaneza, 2010; Espinoza, 2010; Butler and Trumble, 2012; Crosslin et al., 2012a,b; Munyaneza, 2012). Information about migration of B. cockerelli within Mexico and Central America is lacking. In the southwestern United States, potato psyllids reappear in overwintering areas between October and November, presumably dispersing southward from northern locations (Capinera, 2001); however, their origin has not been determined.
In countries and regions where there are no significant seasonal changes during the winter, temperature is relatively cool, and where suitable host plants are available (e.g. Mexico, Central America), the potato psyllid is able to reproduce and develop all year round (Espinoza, 2010; Rubio-Covarrubias et al., 2011).
B. cockerelli was accidentally introduced into New Zealand, apparently sometime in the early 2000s (Gill, 2006; Liefting et al., 2009; Teulon et al., 2009; Thomas et al., 2011), and is now established on both North and South Island where it causes extensive damage to potato, tomato, pepper and tamarillo crops (Teulon et al., 2009). It is not clear how B. cockerelli arrived in New Zealand; however, it has been suggested that the psyllid was introduced from the western United States, probably through smuggled primary host plant material (Thomas et al., 2011).
IntroductionsTop of page
Risk of IntroductionTop of page
B. cockerelli is a serious and economically important pest of potatoes, tomatoes and other solanaceous crops in the western United States, southern Canada, Mexico, Central America and New Zealand (Munyaneza, 2012). Suitable host plants are widespread in almost any part the world and, given its current distribution in the Americas and New Zealand, it is thought that B. cockerelli could establish and overwinter outdoors in areas with warm climate and mild winters. It could also establish under protected conditions in many regions.
If introduced into a new area, the migratory behavior of B. cockerelli, which favours quick and long distance dispersal, would put both the site of introduction and surrounding regions at risk.
Habitat ListTop of page
|Terrestrial – Managed||Cultivated / agricultural land||Principal habitat||Harmful (pest or invasive)|
|Cultivated / agricultural land||Principal habitat||Natural|
|Protected agriculture (e.g. glasshouse production)||Principal habitat||Harmful (pest or invasive)|
|Protected agriculture (e.g. glasshouse production)||Principal habitat||Natural|
Hosts/Species AffectedTop of page
B. cockerelli is found primarily on plants within the family Solanaceae. The psyllid attacks, reproduces and develops on a variety of cultivated and weedy plant species (Essig, 1917; Knowlton and Thomas, 1934; Pletsch, 1947; Jensen, 1954; Wallis, 1955), including crop plants such as potato (Solanum tuberosum), tomato (Solanumlycopersicon), pepper (Capsicum annuum), eggplant (Solanum melongena) and tobacco (Nicotiana tabacum) as well as non-crop species such as nightshade (Solanum spp.), groundcherry (Physalis spp.) and matrimony vine (Lycium spp.).
Adults have been collected from plants in numerous families, including Pinaceae, Salicaceae, Polygonaceae, Chenopodiaceae, Brassicaceae, Asteraceae, Fabaceae, Malvaceae, Amaranthaceae, Lamiaceae, Poaceae, Menthaceae and Convolvulaceae, but this is not the complete host range of this psyllid (Pletch, 1947; Wallis, 1955; Cranshaw, 1993). Beside solanaceous species, B. cockerelli has been shown to reproduce and develop on some Convolvulaceae species, including field bindweed (Convolvulus arvensis) and sweet potato (Ipomoea batatas) (Knowlton and Thomas, 1934; List, 1939; Wallis, 1955; Puketapu and Roskruge, 2011; Munyaneza, unpublished data).
Host Plants and Other Plants AffectedTop of page
|Capsicum annuum (bell pepper)||Solanaceae||Main|
|Convolvulus arvensis (bindweed)||Convolvulaceae||Wild host|
|Ipomoea batatas (sweet potato)||Convolvulaceae||Main|
|Lycium (boxthorn)||Solanaceae||Wild host|
|Medicago sativa (lucerne)||Fabaceae||Other|
|Nicotiana tabacum (tobacco)||Solanaceae||Other|
|Physalis (Groundcherry)||Solanaceae||Wild host|
|Solanum (nightshade)||Solanaceae||Wild host|
|Solanum lycopersicum (tomato)||Solanaceae||Main|
|Solanum melongena (aubergine)||Solanaceae||Other|
|Solanum tuberosum (potato)||Solanaceae||Main|
|Thuja occidentalis (Eastern white cedar)||Cupressaceae||Other|
Growth StagesTop of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page
B. cockerelli has historically been associated with ‘psyllid yellows’ disease of potato and tomato (Richards and Blood, 1933). Psyllid yellows disease is thought to be associated with feeding by psyllid nymphs (List, 1925) and may be caused by a toxin associated with the insect (Carter, 1939), although the actual etiology of the disease is yet to be determined (Sengoda et al., 2010). More recently, this psyllid has been found to be associated with the bacterium ‘Candidatus Liberibacter’ (Hansen et al., 2008; Liefting et al., 2009; Crosslin et al., 2010; Munyaneza, 2010; Munyaneza, 2012; Munyaneza and Henne, 2012) (see ISC datasheet on ‘Candidatus Liberibacter solanacearum’ for details).
The characteristic above-ground plant symptoms of infestation by B. cockerelli in potatoes and tomatoes include retarded growth, erectness of new foliage, chlorosis and purpling of new foliage with basal cupping of leaves, upward rolling of leaves throughout the plant, shortened and thickened terminal internodes resulting in rosetting, enlarged nodes, axillary branches or aerial potato tubers, disruption of fruit set and production of numerous, small, and poor quality fruits (List, 1939; Pletsch, 1947; Daniels, 1954; Wallis, 1955; Munyaneza, 2012; Munyaneza and Henne, 2012).
The below-ground symptoms on potato include the setting of an excessive number of tiny misshapen potato tubers, production of chain tubers and early breaking of dormancy of tubers (List, 1939; Pletsch, 1947; Wallis, 1955). Additional potato tuber symptoms include collapsed stolons, browning of vascular tissue concomitant with necrotic flecking of internal tissues and streaking of the medullary ray tissues, all of which can affect the entire tuber. Upon frying, these symptoms become more pronounced and chips or fries processed from affected tubers show very dark blotches, stripes, or streaks, rendering them commercially unacceptable (Munyaneza et al., 2007a,b; 2008; Secor et al., 2009; Crosslin et al., 2010; Miles et al., 2010; Munyaneza, 2012; Munyaneza and Henne, 2012); see the ISC datasheet on 'Candidatus Liberibacter solanacearum' for details.
List of Symptoms/SignsTop of page
|Fruit / abnormal shape|
|Fruit / reduced size|
|Growing point / dwarfing; stunting|
|Growing point / wilt|
|Leaves / abnormal colours|
|Leaves / abnormal forms|
|Leaves / leaves rolled or folded|
|Leaves / necrotic areas|
|Leaves / wilting|
|Leaves / yellowed or dead|
|Roots / hairy root|
|Stems / fasciation|
|Stems / internal discoloration|
|Stems / stunting or rosetting|
|Stems / wilt|
|Stems / witches broom|
|Vegetative organs / internal rotting or discoloration|
|Vegetative organs / surface cracking|
|Whole plant / discoloration|
|Whole plant / distortion; rosetting|
|Whole plant / dwarfing|
|Whole plant / early senescence|
|Whole plant / plant dead; dieback|
Biology and EcologyTop of page
Recent outbreaks of B. cockerelli in Baja California and coastal California led to the discovery that potato psyllid in those regions is genetically distinct from psyllids that overwinter in southern Texas and eastern Mexico, which suggests there exists two different potato psyllid biotypes, referred to elsewhere as ‘western’ and ‘central’ biotypes (Liu et al., 2006; Jackson et al., 2009). The western biotype differs from southern Texas populations in several life history traits (Liu and Trumble, 2007) and possibly overwinters in different geographic areas to those used by psyllids of the midwestern United States (Trumble, 2008).
Following the 2011 reports of zebra chip in Idaho, Oregon, and Washington (Munyaneza, 2012; Crosslin et al., 2012a,b), a genetic study by Swisher et al. (2012), using high resolution melting analysis of the mitochondrial Cytochrome C Oxidase subunit I-like gene of B. cockerelli, led to the identification of a third biotype (referred to as ‘northwestern haplotype’), so far known only from the Pacific Northwest. This northwestern potato psyllid biotype is genetically different from the central and western biotypes (Swisher et al., 2012). In addition, a recent study by Swisher et al. (2014) identified a fourth biotype of the psyllid, which appears to be distributed in New Mexico and parts of Colorado and referred to as a “southwestern haplotype”. Furthermore, potato psyllids in Mexico, Honduras, El Salvador, and Nicaragua were identified as the central haplotype (Swisher et al., 2013).
In an effort to identify and develop a sex pheromone and other attractants that can be used to develop improved integrated pest management programs for B. cockerelli, its reproductive biology and the role of chemical signals in sex attraction were studied by Guédot et al. (2010; 2012). It was determined for the first time that the potato psyllid possesses a female-produced pheromone that attracts males (Guédot et al., 2010). Guédot et al. (2012) also showed that adult potato psyllids reach reproductive maturity within 48 hours post-eclosion, with females being mature on the day of eclosion and males at one day post-eclosion. In addition, oviposition generally began two days after mating but was delayed when females mated within two days post-eclosion.
Optimum psyllid development occurs at approximately 27°C, whereas oviposition, hatching, and survival are reduced at 32°C and cease at 35°C (List, 1939; Pletsch, 1947; Wallis, 1955; Cranshaw, 2001; Abdullah, 2008; Yang and Liu, 2009; Yang et al., 2010a; Butler and Trumble, 2012). A single generation may be completed in three to five weeks, depending on temperature. The number of generations varies considerably among regions, usually ranging from three to seven. However, once psyllids colonize an area, prolonged oviposition causes the generations to overlap, making it difficult to distinguish between generations (Pletsch, 1947; Wallis, 1955; Munyaneza et al., 2009a). Both adults and nymphs are very cold tolerant, with nymphs surviving temporary exposure to temperatures of -15°C and 50% of adults surviving exposure to -10°C for over 24 hours (Henne et al., 2010a).
Updated information on the biology and ecology of B. cockerelli was recently provided by Munyaneza (2012), Munyaneza and Henne (2012) and Butler and Trumble (2012). Weather is an important element governing the biology of B. cockerelli and its damage potential; B. cockerelli seems to be adapted for warm, but not hot, temperatures. Cool weather during migrations, or at least the absence of elevated temperatures, has been associated with outbreaks of this insect (Pletsch, 1947; Wallis, 1955; Capinera, 2001; Cranshaw, 2001).
Identification of symbionts associated with B. cockerelli is currently being investigated. ‘Candidatus Carsonella ruddii’, the obligate and primary endosymbiont of psyllids, has been confirmed in the potato psyllid (Nachappa et al., 2011; Hail et al., 2012). Beside Candidatus Liberibacter solanacearum, several other secondary endosymbionts associated with the potato psyllid have recently been reported, including the bacteria Wolbachia, Acinetobacter, Methyllibium, Rhizobium, Gordonia, Mycobacterium and Xanthomonas (Nachappa et al., 2011; Hail et al., 2012; Butler and Trumble, 2012; Arp et al., 2014). Currently, little information is available on the interactions and symbiont relationship between these microorganisms and the potato psyllid.
Natural enemiesTop of page
Notes on Natural EnemiesTop of page
B. cockerelli is attacked by a number of natural enemies, including chrysopid larvae, coccinellids, geocorids, anthocorids, mirids, nabids, syrphid larvae and the parasitoids Tamarixia triozae (Hymenoptera: Eulophidae) and Metaphycus psyllidis (Hymenoptera: Encyrtidae), but little is known about their effects on psyllid populations (Pletsch, 1947; Wallis, 1955; Cranshaw, 1993; Al-Jabar, 1999; Butler et al., 2010; Butler and Trumble, 2012a; Liu et al., 2012). In addition, several entomopathogenic fungi, including Beauveria bassiana, Metarhizium anisopliae and Isaria fumosorosae, have been determined to be effective natural enemies of B. cockerelli, causing psyllid mortality up to 99 and 78% under laboratory and field conditions, respectively (Lacey et al., 2009; 2011).
Means of Movement and DispersalTop of page
Adult potato psyllids are good fliers and can disperse over considerable long distances, especially with the onset of wind and hot temperatures. Adults have been shown to migrate en masse to northern western states of the United States and southern Canadian provinces in the spring from the insect overwintering sites in the southwestern United States and northern Mexico (a distance of several hundred kilometers). Immature stages of B. cockerelli are essentially sedentary and do not actively disperse.
Long distance transport of different life stages of this insect pest is possible, particularly by commercial trade of plant material for propagation and produce in the family Solanaceae, which constitute major hosts for B. cockerelli. Based on the discovery of at least four haplotypes of potato psyllid in North and Central America, seasonal dispersal of this insect into potato crops was recently reviewed and discussed by Nelson et al. (2014).
This insect was introduced into New Zealand, where it was recently found established in tomato glasshouses and several outdoors solanaceous crops (Gill, 2006; Liefting et al., 2009; Teulon et al., 2009; Thomas et al., 2011). It is not clear on how the insect arrived in New Zealand, but it was most likely transported with plant material, possibly as eggs (Crosslin et al., 2010; Thomas et al., 2011).
Entry on fruits of host species (e.g. tomato, pepper, eggplant) is possible, especially when they are associated with green parts (e.g. truss tomato). Entry on potato tuber is more unlikely.
Pathway CausesTop of page
Pathway VectorsTop of page
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Flowers/Inflorescences/Cones/Calyx||adults; eggs; nymphs||Yes||Pest or symptoms usually visible to the naked eye|
|Fruits (inc. pods)||eggs; nymphs||Yes||Pest or symptoms usually visible to the naked eye|
|Leaves||adults; eggs; nymphs||Yes||Pest or symptoms usually visible to the naked eye|
|Seedlings/Micropropagated plants||adults; eggs; nymphs||Yes||Pest or symptoms usually visible to the naked eye|
|Stems (above ground)/Shoots/Trunks/Branches||adults; eggs; nymphs||Yes||Pest or symptoms usually visible to the naked eye|
Impact SummaryTop of page
Economic ImpactTop of page
Detailed information on the economic impact of B. cockerelli is provided by Munyaneza (2012). Historically, the extensive damage to solanaceous crops that was observed during the outbreak years of the early 1900s is thought to have been due to B. cockerelli’s association with a physiological disorder in plants referred to as ‘psyllid yellows’ (Richards and Blood, 1933), presumably caused by a toxin that is transmitted during the insect’s feeding activities, especially nymphs (Eyer and Crawford, 1933; Eyer, 1937). However, the nature of this toxin has not yet been identified. ‘Psyllid yellows’ is characterized by yellowing and curling of foliage, stunting or death of plants and a loss in yield (Richards and Blood, 1933; Eyer, 1937). Infected tomato plants produce few or no marketable fruits (List, 1939; Daniels, 1954). In potatoes, psyllid yellows results in yellowing or purpling of foliage, the early death of plants and low yields of marketable tubers (Eyer, 1937; Pletsch, 1947; Daniels, 1954; Wallis, 1955). In areas of outbreaks of psyllid yellows, the disorder was often present in 100% of plants in affected fields, with yield losses exceeding 50% in some areas (Pletsch, 1947).
In recent years, potato, tomato, and pepper growers in a number of geographic areas have suffered extensive economic losses associated with outbreaks of potato psyllid (Trumble, 2008, 2009; Munyaneza et al., 2009b,c,d; Crosslin et al., 2010; Munyaneza, 2010). This increased damage is due to a previously undescribed species of the bacterium liberibacter, tentatively named ‘Candidatus Liberibacter solanacearum’ (syn. Ca. L. psyllaurous) (Hansen et al., 2008; Liefting et al., 2009), now known to be vectored by potato psyllid (Munyaneza et al., 2007a,b; Buchman et al., 2011a,b); see the ISC datasheet on 'Candidatus Liberibacter solanacearum' for details. Potato psyllids acquire and spread the pathogen by feeding on infected plants (Munyaneza et al., 2007a,b). The bacterium is also transmitted transovarially in the psyllid (Hansen et al., 2008), which contributes to the spread of the disease between geographic regions by dispersing psyllids. It also helps maintain the bacterium in geographic regions during the insect’s overwintering period (Crosslin et al., 2010; Munyaneza, 2012).
Symptoms associated with liberibacter in tomatoes and pepper include chlorosis and purpling of leaves, leaf scorching, stunting or death of plants, and production of small, poor-quality fruit (Liefting et al., 2009; McKenzie and Shatters, 2009; Munyaneza et al., 2009c,d; Brown et al., 2010; Crosslin et al., 2010). During the outbreaks of 2001-2003, tomato growers in coastal California and Baja California suffered losses exceeding 50-80% of the crop (Trumble, 2009). In potatoes, foliar symptoms closely resemble those caused by psyllid yellows and purple top diseases (Munyaneza et al., 2007a,b; Sengoda et al., 2009). However, tubers from liberibacter-infected plants develop a defect referred to as ‘zebra chip’, which is not induced by the potential toxin causing psyllid yellows (Munyaneza et al., 2007a,b; 2008; Sengoda et al., 2009). Tubers show a striped pattern of necrosis, which is particularly noticeable when the tuber is processed for chips or fries (Munyaneza et al., 2007a,b; 2008; Miles et al., 2010). Chips or fries from affected plants are not marketable. The defect was of sporadic importance until 2004, when it began to cause millions of dollars in losses to potato growers in the United States, Central America and Mexico (Rubio-Covarrubias et al., 2006; Munyaneza et al., 2007a; 2009b; Crosslin et al., 2010; Munyaneza, 2010). In some regions, entire fields have been abandoned because of zebra chip (Secor and Rivera-Varas, 2004; Munyaneza et al., 2007a; Crosslin et al., 2010; Munyaneza, 2010). The potato industry in Texas estimates that zebra chip could affect over 35% of the potato acreage in Texas, with potential losses annually to growers exceeding 25 million dollars (CNAS, 2006). Finally, quarantine issues have begun to emerge in potato psyllid-affected regions, because some countries now require that shipments of solanaceous crops from certain growing regions be tested for the pathogen before the shipments are allowed entry (Crosslin et al., 2010; Munyaneza, 2012).
Risk and Impact FactorsTop of page Invasiveness
- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
- Tolerant of shade
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Long lived
- Fast growing
- Has high reproductive potential
- Has high genetic variability
- Host damage
- Negatively impacts agriculture
- Negatively impacts trade/international relations
- Pest and disease transmission
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
Prevention and ControlTop of page
Monitoring B. cockerelli is essential for its effective management. Early season management of this insect is crucial to minimize damage and psyllid reproduction in the field. The adult populations are commonly sampled using sweep nets or vacuum devices, but egg and nymphal sampling requires visual examination of foliage. The adults can also be sampled with yellow water-pan traps. Typically, psyllid populations are highest at field edges initially, but, if not controlled, the insects will eventually spread throughout the crop (Workneh et al., 2012; Butler and Trumble, 2012).
B. cockerelli control is currently dominated by insecticide applications (Goolsby et al., 2007; Gharalari et al., 2009; Berry et al., 2009; Butler et al., 2011; Guenthner et al., 2012), but psyllids have been shown to develop insecticide resistance due to the high fecundity and short generation times (McMullen and Jong, 1971). Therefore, alternative strategies should be considered to limit the impact of the potato psyllid and its associated diseases. Even with conventional insecticides, B. cockerelli tends to be difficult to manage. It has been determined that liberibacter is transmitted to potato very rapidly by the potato psyllid, and that a single psyllid per plant can successfully transmit this bacterium to potato in as little as 6 hours, ultimately causing zebra chip (Buchman et al., 2011a,b). This observed low psyllid density, coupled with a short inoculation access period, represents a substantial challenge for growers in controlling the potato psyllid and preventing zebra chip transmission. Just a few infective psyllids feeding on potato for a short period could result in substantial spread of the disease within a potato field or region (Henne et al., 2010b). Most importantly, conventional pesticides may have limited direct disease control, as they may not kill the potato psyllid quick enough to prevent liberibacter and zebra chip transmission, although they may be useful for reducing the overall population of psyllids.
The most valuable and effective strategies to manage zebra chip would likely be those that discourage vector feeding, such as use of plants that are resistant to psyllid feeding or less preferred by the psyllid. Unfortunately, no potato variety has so far been shown to exhibit sufficient resistance or tolerance to zebra chip or potato psyllid (Munyaneza et al., 2011). However, some conventional and biorational pesticides, including plant and mineral oils and kaolin, have shown some substantial deterrence and repellency to potato psyllid feeding and oviposition (Gharalari et al., 2009; Yang et al., 2010b; Butler et al., 2011; Peng et al., 2011) could be useful tools in integrated pest management programs to manage zebra chip and its psyllid vector.
Information on products used to control B. cockerelli is provided by Munyaneza (2012), Munyaneza and Henne (2012) and Butler and Trumble (2012). Good insecticide coverage or translaminar activity is important because psyllids are commonly found on the underside of the leaves. Also, the different life stages require use of specific insecticides as it has been shown that chemicals controlling adults do not necessarily controls nymphs or eggs. Because several generations often overlap, caution should be taken when selecting and applying insecticides targeted against the potato psyllid in relation to which life stages are present in the crop and timing of insecticide applications. Several predators and parasites of B. cockerelli are known, though there is little documentation of their effectiveness. In some areas such as southern Texas, early planted potato crops are more susceptible to psyllid injury than crops planted mid- to late season (Munyaneza et al., 2012).
Gaps in Knowledge/Research NeedsTop of page
At least four haplotypes of B. cockerelli have so far been identified (Swisher et al., 2012; Swishet et al., 2014) and more may be discovered. Although these psyllid haplotypes have been shown to be different genetically, little is known of their differences in biological traits. This information is essential, especially for pest management purposes. In addition, information on overwintering of B. cockerelli, particularly in the regions with temperate climate (Jensen, 2012; Murphy et al., 2013), is lacking. Furthermore, accurate information on long distance movement and dispersal of B. cockerelli is crucial for predicting temporal and spatial colonization of field crops by this insect. Therefore, it is imperative that studies are conducted to further clarify the biology, ecology, movement and dispersal of B. cockerelli, in order to develop effective management strategies for this insect pest.
ReferencesTop of page
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ContributorsTop of page
04/10/2014 Updated by:
Joseph E Munyaneza, USDA-ARS, USA
19/11/12 Original text by:
Joseph E Munyaneza, USDA-ARS, USA
Distribution MapsTop of page
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