Roles of Host Plants in
Boll Weevil Range Expansion
beyond Tropical Mesoamerica
Allan T. Showier
ABSTRACT New findings about the food habits of the boll weevil, Anthonomus Brandis Brandis Boheman (Coleoptera: Curculionidae), refine current theories about range expansion beyond its tropical Mesoamerican origin. The weevil co-evolved with
perennial host plants including cotton, Gossypium hirsutum L.,and patches of host plants were reached by flight, likely instigated
by competition for limited resources. Boll weevils might have sought nectar, pollen, or both for energy to facilitate searching for
patches of host plants. Range expansion was facilitated by shifts to new host plants, and in its dispersion north, the insect reached
the subtropical Lower Rio Grande Valley of Texas, becoming established only when the citrus industry first boomed in the early
1890s. Commonly grown subtropical citrus species bear fruit containing sugars and other nutrients that can sustain active boll
weevils over winter in spite of a 5-mo mandatory fall cotton plowdown. Feeding on citrus and cactus fruit, and possibly on nectars
and pollens, makes nonreproductive host plants (used only for foraging) important for dispersal to and establishment at new
latitudes. Once in south Texas, the pest moved rapidly through the Cotton Belt during warm seasons. To cope with cold temperate
winters, the insect resorted to a poorly adapted dormancy response whereby mortality at study sites is often 100%. Extensive
cotton monocultures support the buildup of large enough summer populations to increase the chance that a few survive and
infest the spring crop. Establishment in temperate fringe areas (central Texas, southern Arizona, and California) and the pest's
range expansion into subtropical northern Argentina are discussed in terms of associations with reproductive (plants that confer
sufficient nutrients for boll weevil reproduction) and nonreproductive, indigenous, and exotic host plants.
he boll weevil, Anthonomus Brandis Brandis Boheman (Coleoptera: Curculionidae), evolved in tropical Mesoamerica
but became established in cotton of the Lower Rio Grande
Valley, TX, in 1892 (Parencia 1978, Haney 2001). From there, it
spread throughout the u.s. Cotton Belt (Burke et a!. 1986, Rummel
and Summy 1997), costing $83.34 billion in losses and $18.67 billion for insecticide-based control between 1893 and 1999 (Haney
2001). Infestations became so injurious that cotton-free winter
periods were instituted by law in some areas, and eradication
programs were launched (Dickerson and Haney 2001, Haney 2001,
Haney et a!. 2001, Johnson and Martin 2001, Texas Department of
Agriculture 2002). The boll weevil also spread south to subtropical
cotton-growing areas of Argentina (Cuadrado 2002, Mas eta!. 2007).
This literature-based research refines and clarifies contemporary
perspectives and presents alternative possibilities on the pest's
relationships with host plants that allowed range expansion well
beyond the Mesoamerican tropics.
T
Mesoamerican Crucible
Tropical Mesoamerica starts at =22° N latitude, near modernday Tampico at the north edge of Veracruz, Mexico (Fig. 1) (Burke
et a!. 1986), and extends into Central America. The oldest boll
234
weevil collected, from -900 C.E., was ensnared in a cotton boll
from tropical Oaxaca (=17° 0' N latitude) (Fig. 1) (Warner and Smith
1968). Mesoamerican cotton grew in warm lowlands, such as in
Veracruz (=19° N latitude) (Fig. 1), where June-October rainfall
is 1,200-2,000 mmjyr (Garcia 1970, Berdan 1987), and cotton is
grown through the cool, dry winter in accordance with its natural
short-daylength flowering habit (Mauney and Phillips 1963, Stark
et a!. 1998).
Boll weevils and Gossypium hirsutum L. evolved together (Burke
et a!. 1986, Brubaker and Wendel 1994 ). The insect also probably
fed and reproduced on G. harknessii Brandg., G. lobatum Gentry,
G. laxum Phil!., CienfueBosia rosei Fryx., Hampea rovirasae Stand!.,
Pseudabutilon lozani (Rose) R. E. Fries, Callirhoe sp., Thespesia
populnea (L.) Soland ex Carrea, as well as other, less preferred plant
species (Fryxell1967, 1969, 1974; Parrott et a!. 1969; Cross et a!.
1975, Burke and Cate 1979). In Chiapas (=14° 30' to 17° 20' N) (Fig.
1), the boll weevil is a pest of cultivated cotton and infests Hampea
sp., suggesting that it spread from wild-growing hosts to cultivated
cotton (Burke and Cate 1979). Based on Maya and Aztec artifacts
(Berdan 1987), it is believed that cotton cultivation may have begun
as early as 5,800 B.c.E. in Mesoamerica (Roche 1994), and most of
it was G.hirsutum.
American Entomologist.
Winter 2009
UNITED STATES
T.xo~
COlton
JOON
Belt
i
Fig. 1. Locations related to
the boll weevil's origins in
Mesoamerica and range
expansion through Mexico
into Arizona, California,
Texas, and the U.S. Cotton
Belt.
Chi.lp:1S
~.,
o
500
~
Native Mesoamerican reproductive host plants, which supply
nutrients for egg production and sites for oviposition, are mostly
perennial; and flowering is heightened during the cool dry season
(Mauney and Phillips 1963, Wilson and Fryxell1970, Cross et al.
1975, Brubaker and Wendel 1994). Boll weevils evolved with and
adapted to their hosts' patchy distributions and flowering habits,
and did not experience cold winter seasons. Food shortage among
patches of host plants was likely the greatest challenge to survival
(Showier 2009). Greenberg et al. (2003) found that limiting cotton
squares reduced fecundity in boll weevils compared with plentiful
squares; and McGovern et al. (1987) reported that boll weevils discriminate against squares where oviposition has already occurred.
Hence, boll weevils probably flew in search of patches that offered
more pristine resources. Wagner and Villavaso (1999) found that
hypertrophied fat bodies in temperate boll weevils coincide with
dispersal from cotton fields to hibernacula. In the subtropics, high
levels of post-harvest boll weevil movement occur beyond cotton
fields (Summy et al. 1988; Showier 2003, 2006), and experiments
using tethered boll weevils showed that fat fuels flight (Rankin et
al. 1994). In Arizona, the pest oviposits in bolls long after cotton
harvest around 15 October (Henneberry et al. 1990), indicating
that the insects neither shut down reproductively nor search for
hibernacula.
Although boll weevils engage in search behavior, curculionids are
not known to migrate seasonally between specific locations as groups,
or swarms, on prevailing winds (Showier 2006, 2007). Rather, longrange movement of individual boll weevils is opportunistic, relying
largely on wind currents and possibly accidental mechanical transfer
on farm equipment or in loads of harvested cotton. The weevils
Guerra (1988) observed flying upward after feeding on bolls could
have encountered wind currents that directed them to favorable and
unfavorable habitats, akin to wind-dispersed plant seeds. This can
explain, in part, how the occasional boll weevil has been trapped far
from release sites (Johnson eta1.1975, Guerra 1988) or from known
American Entomologist.
Volume 55, Number 4
host plants (Jones et al. 1992). It also explains genetic similarities
among boll weevils in cotton-growing areas <300 km apart (Kim
and Sappington 2004a,b, 2006; Kim et al. 2006, 2008); boll weevils
may have traversed the distances over generations, spreading as
indicated herein. The suggestion that boll weevils from one sampled
location might have moved to other locations where the same natural
genetic marker was found (Kim and Sappington 2004a,b, 2006; Kim
et al. 2006, 2008), however, has not been supported by necessary
data from the vast expanses of unsampled areas; such speculation is
based on assumption, and is by no means definitive as regards boll
weevil origins and dispersal.
Expansion into Tamaulipas and Coahuila
Acceptance of new host plants, or host shifting, was key to the
dispersal of A. gran dis in prehistoric times (Burke and Cate 1979,
Burke et al. 1986, Scataglini et al. 2000). Northward range expansion in Mexico probably occurred gradually between patches of
hosts, such as Cienfuegosia spp. (Jones et al. 1991), but no chronological record exists. The pest's range did not expand explosively
in the late 19th century as claimed by Kim and Sappington (2004a)
because heavy boll weevil infestations of cotton in northern Mexico
(at more northerly latitudes than the Rio Grande Valley of Texas)
during the 1860s are documented (Howard 1897). Jones et al.
(1992) found adult boll weevils on a patch of cotton regrowth 30
km from the nearest cultivated cotton and concluded that they
moved from the south with prevailing winds (Moody et al. 1993,
Westbrook et al. 2000) after cotton harvest. Because wild host
plants were assumed to be absent, they concluded weevils could
only have reached the isolated cotton patch by long-distance dispersal (Jones et al. 1992). A systematic method for determining
the absence of host plants, however, was not indicated, and it is
doubtful that nonreproductive host plants (used only for foraging)
were considered.
Ofthe six published studies that have quantified pollen grains in
235
Table 1. Data on pollen ingestion
Sources of
pollen
Mixedtaxa
Mixedtaxa
Mixedtaxa
Single species
Mixedtaxa
Mixedtaxa
Mixedtaxa
Mixedtaxa
Almond
Citrus
Corn
Melon
Mixed
by boll weevils from six studies
Mean no. pollen
grains/weevil"
n
% weevilswith grains"
856
2,171
144
23
69
56
0.2-149f
4-19f
20-10or
2.7
12.9
18.0
8.1
23.1
5.1
9.6
1.7
13.5
1,204
06
486
5,081
50
40
30
40
50
34.7
82
87
98
31
36
21
100
Regionb
Seasons'
LGC
LRGV
Tamaulipas
All
All
All
Tamaulipas
Tamaulipas
Central Texas
MSDelta
All
Oct-May
Apr-Jul
All
Experimental
Conditionsd
Natural
Natural
Natural
Artificial
Natural
Natural
Natural
Natural
Artificial'
Artificial'
Artificial'
Artificial'
Artificial'
Reference
Benedict et al. 1991
Ibid
Ibid
Ibid
Jones et al. 1992
Jones et al. 1993
Jones and Coppedge 1999
Hardee et al. 1999
Greenberg et al. 2007
Ibid
Ibid
Ibid
Ibid
"-, data not availableinthe reference.
b _, no regionindicatedbecausethe studywasconductedentirelyinthe laboratoryor in cages.LGC,
LowerGulfcoast,Texas;LRGY,
LowerRioGrandeValley,Texas;Central
Texas,Crockett,Munday,and Uvaldecounties;Tamaulipas,a statein northeasternMexico;MSDelta,MississippiDelta,WashingtonCounty.
'-, not applicable.
d Artificial,
no-choicelaboratoryor cagestudies;natural,wild-caughtweevilsdissected.
'Pollen,fromnectar-richbeepellets,waspresentedin dropletsofwater.
fTwenty-twofloweringplantswerecagedseparatelywithbollweevils;dissectionsshowedweevilshad < 10 pollengrains/weevilfrom91% ofthe plantspeciesusedinthe
experiment.
adult boll weevil digestive tracts (Table 1), five involved dissections
of wild-caught weevils. Benedict et al. (1991) identified pollen of 12
plant families from weevils collected in the Lower Rio Grande Valley
(e.g., 17% Asteraceae, 20% Brassicaceae, 24% Poaceae), 9 plant families from the lower GulfCoast (e.g., 6% Asteraceae, 12% Poaceae), and
6 plant families from Tamaulipas, Mexico (e.g., 9% Asteaceae, 15%
Poaceae). In Tamaulipas, Jones et al. (1993) reported ingested pollen
from 1,204 plant species (e.g., 29.9% Compositae, 14.6% Leguminoseae, 5.3% Malvaceae). Weevils from Crockett, Munday, and Uvalde
counties of Central Texas contained pollens from 311 plant taxa (e.g.,
13% Fabaceae, 8% Poaceae, 23% Salicaceae) (Jones and Coppedge
1999). Weevils from Washington County in the Mississippi Delta
had 301 types of pollen (the most commonly encountered families
were Anacardiaceae, Asteraceae, Chenopodiaceae, Amaranthaceae,
Fagaceae, Mavalceae, and Poaceae) (Hardee et al. 1999).
Even though the diversity of pollens inside wild boll weevils
might be noteworthy and nutritious, the average numbers of grains
per weevil were only 2.7-23 (Table 1). Ingested pollen grains are
typically 10-100 11min diameter (Jones and Coppedge 1999); but
when some weevil digestive tracts hold >200 grains, the observed
levels suggestthat despite their capacity to ingest> Il-fold more, they
might be incidentally ingesting pollen grains while drinking nectar, a
possibility first suggested by Benedict et al. (1991). Pollen can drop
into nectar when the weevil, other insects, or wind jostle the anthers;
and some pollen grains are wind-blown from distant sources (Faegri
and van der Pilj 1971) and ingested (Benedict et al. 1991).
Benedict etal. (1991) conducted a set of no-choice assays whereby
adult boll weevils were caged with individual species of flowering
plants. Of the 22 plant species used in the experiment, 20 resulted
in <10 ingested pollen grains, and 18 resulted in <5 grains, but an
overall average of 47 grains/weevil was found for the remaining
two plant species (Table 1). Although the study suggests that boll
weevils seek pollen as food, the weevils were presented with intact
flowers and nectar ingestion was not measured; it is possible that
236
pollen and nectar were sought, or that one was being incidentally
ingested while the weevils actually fed on the other.
Greenberg et al. (2007) similarly found pollen grains inside
laboratory-fed boll weevils (Table 1), but bee pellets were used as
the pollen sources for each assay (Greenberg etal. 2007). In an assay
using Aloe sp. (Jones et al. 1993) and plants used by Greenberg et
al. (2007) (Table 1), adult boll weevils survived on average for up to
-73 d and -55 d, respectively, on bee pellets. Bee pellets, however,
mostly contain substantial amounts of nectar (Iannuzzi 1993, Human
and Nicolson 2006). When Greenberg etal. (2007) mixed the pellets
into a drop of water (Greenberg et al. 2007), the same confounding
condition of pollen grains presented with nectar (or dilute nectar)
likely occurred, but fluid dietary constituents were not considered. Of
the five kinds of bee pellets used, four resulted in an average of <14
ingested grains/weevil (Table 1) (Greenberg et al. 2007). The fifth
bee pellet, made up of almond [Prunus dulcis (Mill.) Webb], pollen
resulted in - 23 ingested grains/weevil (Greenberg et al. 2007), but
almonds are not cultivated in the Lower Rio Grande Valley.Although
Greenberg etal. (2007) identified almond pollen as a positive experimental control, there was no basis indicated for that assumption; and
its presentation in bee pellets makes it inappropriate for determining
whether boll weevils actively feed on nonmalvaceous pollens.
Boll weevils have been assumed to be exclusively pollen feeders
(Burks and Earle 1965, Cate and Skinner 1978) because pollen grains,
and not plant juices, can be observed in dissected specimens and
counted under a microscope (Showier and Abrigo 2007). Although
pollens are sources of amino acids, proteins, fats, carbohydrates,
minerals, and vitamins (Stansleyand Linskens 1974, Dayetal.1990,
Roulston et al. 2000, Villanueva et al. 2002), the nutritive value of
nonmalvaceous pollens to boll weevils has not yet been reliably
established, and published evidence for pollen as a sought-after
food is circumstantial.
Nectars also contain amino acids, carbohydrates, lipids, and
other nutrients (Baker and Baker 1973, 1975; Forcone et al. 1997;
American Entomologist.
Winter 2009
Gardener and Gillman 2001), and adult boll weevil and other insect
longevities can be extended by sugar solutions and honey for as long
as 5 wk (Butler 1968, Benchoter and Leal 1976, Tsiropoulos 1980,
Haynes 1985, Haynes and Smith 1992, Hedin and McCarty 1995).
While concentrations and compositions of nectars can vary (Baker
et al. 1998, Ortiz et al. 2003), it is possible that the boll weevil is a
nectar opportunist, seeking it from a variety of plant species, and
might have survived searches for reproductive hosts by subsisting
on nectar. However, like pollen, the value of nectar to boll weevils
has not yet been reliably determined.
Subtropical Transition at the Lower Rio Grande Valley
The Lower Rio Grande Valley is a subtropical region along the
southernmost edge ofTexas (=26 9' N at Weslaco, Hidalgo County)
(Fig. 1 and 2). Between it and other major cotton-growing areas
in Texas (e.g., Falfurrias and Kingsville; Fig. 2) and the rest of the
Cotton Belt is =70 km of cotton-free scrub. Several indigenous
reproductive host plants of the boll weevil, including Cienfuegosia drummondi (A. Gray) Lewt., C. sulphurea (Kunth) HassI., and
Thespesia spp., occur in the Lower Rio Grande Valley and along
the south Texas coast (Lukefahr 1956, Lukefahr and Martin 1962,
Parrott et al. 1969, Cross et al. 1975, Burke and Clark 1976, Jones
et al. 1991). Lukefahr and Martin (1962) described C.sulphurea as
being "common" on the coast from Brownsville (25 54' N) (Fig.
2) to Gonzales (29 2' N). Near Brownsville, C. sulphurea blooms
in February; but in Nueces County (27 35' - 27 47' N), it blooms
April-October (Lukefahr and Martin 1962), suggesting that a fringe
0
0
0
0
0
orthern High
..................................
..'Ptains ..·..···
.......................................................
.
"
,
..
,
,
,
TEXAS
................... ...,.~
.
.
"
,
u, ••••••••••••••••.••••
...••••..••
t"J .•.•••.••.•••••.••••.•••••.••.••••••••.••.•••••.••.••••••••.••.•••••.••.•••.•••..••..
W±E
.............. ,
,
S
..............
,
.
60
....
J ~~ ~!~J
MEXICO
Fig. 2. Locations related to the boll weevil in various parts of Texas.
American Entomologist.
.
environment between subtropical and temperate regions exists for
native hosts and boll weevils.
Aside from scattered native reproductive host plants, volunteer
cotton harbors boll weevils during the mild winters (Summy et al.
1988, 1993). On reproductive hosts, only fruiting bodies appear to
have the assemblage of nutrients needed for egg production; omission or reduction of certain nutrients can slow or halt the process
(Vanderzant 1963, 1965; Vanderzantand Richardson 1964; Hilliard
and Keeley 1984).
Esquivel et al. (2004) reported that cotyledon-stage
cotton
supported adult boll weevils for =82 d; seedlings were reported
to sustain them from 19 to 22 d (Fenton and Dunnam 1929, Fye et
al. 1959), and four-leaf stage cotton plants for =62 d (Esquivel et
al. 2004). Regardless of the discrepancy, it is unlikely that seedling
cotton is important as a food source for overwintering boll weevils
(White and Rummel 1978). Fenton and Dunnam (1929) found that
boll weevils survived =8 d on pre fruiting cotton. Rummel and Carroll (1985) determined that 60% of overwintered boll weevils fed
pre fruiting cotton plants died within 10 d, and only 17% survived
until fruiting. Bariola (1984) reported that overwintered
boll
weevils fed cotton terminals (without fruit) lived =23 d. Suh and
Spurgeon (2006) indicated that 75% of overwintered boll weevils
fed prefruiting regrowth cotton died during the initial 3 wk of being
starved; however, 5% lived ~12 wk. Examination of the experimental
materials used by Suh and Spurgeon (2006), however, revealed that
their "vegetative-stage" cotton plants were actively fruiting. Esquivel
et al. (2004) concluded that vegetative-stage cotton is nutritionally
inadequate for extending longevity of
overwintered boll weevils, but Suh and
Spurgeon (2006), perhaps because
their weevils were not actually being
........ ,~
- - .
fed prefruiting cotton plants, suggested
the contrary. Regardless, vegetative.
stage cotton plants after spring planting are probably of minor importance
because regrowth and volunteer cotton
fruit during winter (Summy et al. 1988)
and summer, and other hosts that confer greater longevity are common in
the subtropics (Showier 2006, Showier
and Abrigo 2007). Summy et al. (1993)
estimated that late January regrowth
cotton could harbor substantial numbers of boll weevils in the Lower Rio
Grande Valley.
Prickly pear cactus, Opuntia spp.,
flowers and fruit sustain boll weevils
for averages of 39.4 d and 82.6 d, and
for maximums of 70 d and 181 d, respectively (Jones et al. 1993, Showier
and Abrigo 2007). Despite the availability of Opuntia spp. and other host
plants in Mexico and southern Texas
(Gaines 1934, Lukefahr 1956, Lukefahr
and Martin 1962, Stoner 1968, Cross et
G LFOF
al. 1975, Vigueras and Portillo 2001),
MEXICO
but cotton in the Lower Rio Grande
Valley remained weevil-free for 30 yr
of commercial production beginning
Volume 55, Number 4
•••••
••••••••••
237
=1860 (Garza and Long 2001); but cotton around Monclova (26°
57' N), Coahuila (Fig. 1), =45' latitude north and 220 km west of the
Lower Rio Grande Valley (Fig. 1), was so heavily infested that the
crop was abandoned in 1862 (Howard 1897).
Grapefruit, Citrus paradisi MacFad., and orange, C.sinensis (L.) Osbeck., endocarps support boll weevils for much longer than Opuntia;
the maximum is only 7 d short of weevils fed large squares (Showier
and Abrigo 2007, Showier 2009). Under orchard conditions, boll weevil food sources are concentrated and appear to support substantial
active populations through winter (Showier 2006); endocarps are
accessible through cracks, holes, or lesions while the fruit is attached
to the plant or on the ground (Showier 2007, Showier and Abrigo
2007). Initial establishment of boll weevils in commercial Lower Rio
Grande Valley cotton during the early 1890s (Parencia 1978, Haney
2001) may have been linked to a simultaneous citrus industry boom
(Waibel 1953).
Sections of sugarcane stalks sustained adult boll weevils for =20
d on average (Showier and Abrigo 2007), and although sugarcane
alone does not support substantial longevity, it might playa role in
the pest's persistence through subtropical winters (Showier 2007).
The limited utility of sugarcane suggests that boll weevils might
survive longer on some sugars than on others (Haynes 1985, Hedin
and McCarty 1995), or that sugarcane lacks nutrients present in
citrus and prickly pear endocarps.
Aside from recorded instances of boll weevil infestations in Monclova (Fig. 1) (Riley 1885, Howard 1897), the history of cotton and
the boll weevil in Mexico has not been well documented. Although
citrus, grapes, and other sugary fruits are cultivated in Coahuila, no
published data are available on seasonal boll weevil populations.
Subtropical overwinter survivorship is relatively high, facilitated
mostly by regrowth cotton (Summy et al. 1988, 1993), citrus (Showier
2006, 2007), and by the boll weevils entering quiescence during
infrequent short cold spells (Guerra et al. 1984). Quiescence is an
immediate response to a decline of limiting environmental factors
that is reversible if those factors revert (Kostal 2006). Guerra et
al. (1984) determined that when the dry season starts in tropical
Mexico, immature boll weevils inside bolls sometimes complete
development to adulthood remaining encapsulated and adults enter
quiescence from January until March. Other researchers reported
boll weevils surviving extended periods in desiccated bolls of Central Texas (Cowan et al. 1963, Walker and Shipp 1963). Boll weevil
activity during Lower Rio Grande Valley winters, however, suggests
that many, or most, are in neither diapause nor quiescence (Summy
et al. 1988; Showier 2006, 2007, 2009).
The Cotton Belt Superhighway
Cotton was grown without boll weevils in the United States
from colonial times until 1892, when the first was collected in the
Lower Rio Grande Valley (Parencia 1978, Haney 2001). Two years
later, several south Texas counties were so infested that the USDA
recommended destroying cotton stalks to suppress overwintering
boll weevils and establishing a "cotton-free" quarantine zone to
impede spread (Haney 2001). The pest was still confined to Texas in
1904, but progressed 800 km from Brownsville (Hunter 1904). Soon
thereafter extensive temperate Cotton Belt mono cultures permitted
range expansion at an average rate of 104 kmjyr (Parencia 1978).
By 1917, every cotton-producing county in Georgia was infested
(Fig. 1) (Hunter 1917).
With greater latitudes, winters are generally longer and colder,
238
and incidence of so-called diapause increases (Sterling and Adkisson
1966). Diapause, a form of seasonally mediated dormancy (Tauber et
a1.1986, Taub-Montemayor eta1.1997, Kostal 2006), does not occur
in the Lower Rio Grande Valley (Graham et al. 1978; Summy et al.
1988; Showier 2006, 2007, 2009). In the temperate Cotton Belt, however, cold winters are often characterized by such a dramatic decline
oflivingvegetation that the continued existence of boll weevils relies
on a dormancy response to unfamiliar environmental conditions.
In temperate parts of Texas, survival of adult boll weevils through
winters under field conditions is mostly 0-6%, even in the Rolling
Plains (=32° 20' N) and Central Texas (=30° 40' N) (Davis et al. 1975,
Rummel and Carroll 1983, Slosser eta1.1984, Price et al. 1985), both
of which are <500 km north of the Lower Rio Grande Valley (Fig.
2). This percentage suggests that putative boll weevil diapause is
not particularly effective for individuals. In Central Texas, however,
south of the Rolling Plains (Fig. 2), winter survival in a 19-yr study at
several locations was occasionally high, 100% during 1 yr (Davis et
al. 1975). Similarly, in an 18-yr study, the percentage of overwinter
survival in Mississippi was greater at the south end of the state than
at the north end, and mortality was attributed to low temperatures
(Pfimmer and MerkI1981). Boll weevils in the northern Texas High
Plains (=35° 40' - 36° 30') (Fig. 2) reportedly diapause earlier and
in greater percentages than in Central Texas (Sterling and Adkisson
1966). Bottrell et al. (1972) found that winter mortality in the Rolling
Plains was greatest in January and February, also attributed to low
temperatures. The fact that eradication programs capitalize on the
weevil's weak defense againsttemperate winters by using "diapause
sprays" (Brazzel and Newsom 1959, Brazzel et al. 1961, McKibben
et al. 2001) indicates that the pest's existence in those areas is more
tenuous than in subtropical or tropical regions.
Temperate
Fringes in Arizona and California
In Texas, the southern
fringe of the temperate Cotton Belt is
=28-33° N, estimated by changes in the flowering habit of uncultivated indigenous reproductive host plants (Lukefahr and Martin
1962) and by relatively high boll weevil survivorship through winter
(Davis et al. 1975, Rummel and Carroll 1983, Slosser et al. 1984, Price
et al. 1985). The northernmost part of this fringe extends through the
cotton- and citrus-growing areas of southern Arizona and southern
California (both =33° N) (Fig. 1).
Burke et al. (1986) supposed that the boll weevil's expansion
north through western Mexico occurred before the late Pleistocene
via host shifts until the weevil reached contemporary southern
Arizona. In the 1960s and early 1970s, boll weevils attacked cotton
in southern Arizona and southern California sporadically. The boll
weevil was first found in the Bard Valley of California, near Arizona,
in 1965 (Clark 2001). In Arizona, the pest did not become established
year-round until 1978, when a ban on stub cotton (growing the next
crop from stubble) was lifted (Neal and Antilla 2001, Southwest Boll
Weevil Eradication Program 2007), allowing continuous access to
its primary reproductive, mono cultured host plant.
Cotton is grown on =36,900 ha in southern Arizona (i.e., Cochise,
Pima, Santa Cruz, and Yuma counties; 1981-1987 average). About
2,400 ha of grapefruit, oranges, and tangerines (not including lemons, 1996-1997 data) (USDANational Agricultural Statistics Service
2008) are confined to Yuma county. Imperial County, CA, (Fig. 1)
grows =2,300 ha of cotton with =500 ha of grapefruit and oranges
(California Agricultural Statistics Service 2004, Meiser 2004). In
1983, eradication programs started in southern Arizona and southAmerican Entomologist • Winter 2009
nee
ARGEI TINA
o1__
k_m__
1,000
1
••••
vance, in the general direction of prevailing winds, was =66 kmjyr
(Gomez et al. 1998).
Boll weevils presumably entered parts of subtropical northern
Argentina soon thereafter (Scataglini et al. 2000). Cuadrado (2002)
reported that although cotton had not been cultivated for 5 yr in
Misiones Province (=26°-27° 30' S [Fig. 3], which corresponds to
the north latitudes of the Lower Rio Grande Valley [Fig. 2]), active
boll weevils were found year-round. Trapped weevils (n = 1,360)
were dissected, revealing 5,325 pollen grains, only 3.9 jweevil, from
28 plant species (Cuadrado 2002). Just 20 weevils contained> 1%
of the pollen grains (~53 grains) (Cuadrado 2002). Because cotton
had previously been grown in Misiones, regrowth was likely common, perennial, and supported breeding populations. Argentina has
indigenous nectar and pollen sources (Forcone et al. 1997, Chalcoff
et al. 2006), and Opuntia is widespread (Montagnini et al. 1997,
Felker et al. 2005). Perhaps more significantly, Misiones Province
and particularly the locations where the weevils were collected are
major citrus production areas (Agostini et al. 1986, Peres and Timmer 1986, Ares et al. 2006, Segura et al. 2006).
Also, since 1994 subtropical Pilcomayo and Pilaga counties (=26°
S) in Formosa Province (Fig. 3) have been boll weevil eradication
zones (Carmona et al. 2003), but boll weevils have continued arriving from Paraguay, advancing at a rate of =9. 7 5 kmjyr (Mas et al.
2007). Half of the cultivated area in Formosa Province is occupied by
cotton, but citrus is considered to be the regional specialty (Whittle
1998). As in the subtropical Lower Rio Grande Valley, boll weevil
establishment in subtropical parts of Argentina appears linked to
nonreproductive host plants.
SOUTH AMERICA
Acknowledgments
Fig. 3. Locations related to the boll weevil's recent advances into Argentina.
ern California (expanding a few years later to include temperate
central parts of each state), and the pest was eradicated from both
states by 1988 (Beasley and Henneberry 1985, Moore 1985, Clark
2001, Neal and Antilla 2001).
Despite the presence of citrus orchards, the general latitude of
southern Arizona and southern California is greater than that of subtropical areas, and winter attrition may have been comparable to that
of the northernmost edge of the fringe in Central Texas. Combined
with relatively small plantings of cotton on which to concentrate
spray efforts, the small citrus producing area (by comparison, the
Lower Rio Grande Valley typically has =81,000 ha of cotton and
11,000 ha of citrus [Tuxbury 2005, Sauls 2008]), and reinstatement
of the ban on stub cotton and its enforcement during the eradication
program (Arizona Department of Agriculture 2005) made the pest's
existence there untenable.
Northern Argentina
Relatively little is known of the boll weevil's southward range
expansion, but it was reported from Colombia and Venezuela on
Cienfuegosia affinis (Kunth) Hochr., Hampea spp., and Thespesia sp.
in 1952 (Cross et al. 1975, Scataglini et al. 2000). The insect probably moved through South America much like it did in Mexico, using
native reproductive host plants (Szumikowski 1953, Lukefahr and
Martin 1962, Fryxell1969, Scataglini et al. 2000), cultivated cotton,
and nonreproductive hosts. In Paraguay, the boll weevil's rate of adAmerican Entomologist.
Volume 55, Number 4
Thanks to Patrick Moran and Francis Reay-Jones for critical
reviews, and to Veronica Abrigo and Jaime Cavazos for research assistance.
•••••
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Allan T. Showier is a research entomologist with the USDA-ARSat the
Kika de la Garza Subtropical Agricultural Research Center in Weslaco, TX.
He has been researching subtropical boll weevil issues for a decade and is
presently researching pest problems of sugarcane and vegetables. allan.
[email protected].
The Light Weight Townes Trap
· Generalist insect collector, especially effective for
Hymenoptera and Diptera
· Very light and mobile, easy to set up and transport
· Made of sun-resistant polyester and about 2 m in
length
· Complete with tie-down lines and polypropylene
wet-and- dry collection head
American Entomologist
• Winter 2009