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Annals of the Entomological Society of America Advance Access published May 28, 2015
MORPHOLOGY, HISTOLOGY,
AND
FINE STRUCTURE
Life History and Laboratory Rearing of Bagrada hilaris
(Hemiptera: Heteroptera: Pentatomidae) with
Descriptions of Immature Stages
M. E. TAYLOR,1 C. S. BUNDY,1,2 AND J. E. McPHERSON3
Ann. Entomol. Soc. Am. 1–16 (2015); DOI: 10.1093/aesa/sav048
ABSTRACT Bagrada hilaris (Burmeister) (Hemiptera: Heteroptera: Pentatomidae) is an Old World
pest recently established in North America. Literature on the life history and immature descriptions of
B. hilaris is scattered and variable in content. Therefore, we conducted a study of this insect’s life history
in southern New Mexico from January 2012 to August 2014, reared the bugs in the laboratory, and described the immature stages. Two primary host plants, London rocket (Sisymbrium irio L., a winter annual) and mesa pepperwort (Lepidium alyssoides A. Gray, a summer perennial), were sampled weekly to
record numbers of the various life stages and behavioral activities; broccoli (Brassica oleracea L.) also
was sampled for one season. Adults were found continuously throughout the year. They deposited eggs
individually in the soil near the bases of their host plants from February through October. Nymphs were
found every month of the year. The number of generations per year was difficult to determine due to
such factors as extreme overlap of generations and a lack of reproductive diapause. However, the data
roughly suggest this species is bivoltine, although there is some evidence of a partial third generation.
The bug also was reared from egg to adult under controlled laboratory conditions on fruiting structures
of mesa pepperwort at 25 6 0.01 C under a photoperiod of 14:10 (L:D) h. The incubation period
averaged 7.45 d. The five stadia averaged 3.35, 7.08, 6.39, 7.33, and 10.25 d, respectively. Instars can be
distinguished readily by differences in several morphological features in addition to body size and
coloration.
KEY WORDS Bagrada hilaris, Pentatomidae, life history, immature descriptions
The family Pentatomidae, which occurs worldwide,
contains 900 genera and >4,700 species (Henry
2009). It is represented in America north of Mexico by
60 genera and >200 species (Froescher 1988) including Bagrada hilaris (Burmeister) (Halbert and Eger
2010).
Bagrada is primarily an Old World genus with three
subgenera and 16 species (Rider 2006). Only B. hilaris
occurs in both the Old and New Worlds (Halbert and
Eger 2010).
B. hilaris is considered native to Africa, India, Pakistan, Southeast Asia, and southern Europe, where it is
a pest of cole crops (Brassicaceae) (Arakelian 2008–
2009, Reed et al. 2013, Taylor et al. 2014). It first was
collected in the United States from Los Angeles
County, CA, in 2008 (Garrison 2008–2009) and has
spread north and south in California and east through
Nevada, Utah, Arizona, and New Mexico to Texas
(Reed et al. 2013); it now is established in Mexico
(Sa´nchez-Pen˜a 2014).
1
Department of Entomology, Plant Pathology, and Weed Science,
New Mexico State University, Las Cruces, NM 88003.
2
Corresponding author, e-mail: [email protected].
3
Department of Zoology, Southern Illinois University, Carbondale,
IL 62901.
Although best known as a pest of cole crops, B. hilaris has been reported from at least 76 species in
20 plant families from Poaceae to the Solanaceae
(Table 1). The bugs feed predominantly on leaves and
young seeds. Severe host damage can be caused by
feeding on the apical meristem or cotyledons (Palumbo
and Natwick 2010).
B. hilaris has been reared in the laboratory under
variable conditions. Azim and Shafee (1986) reported
an incubation period of 2–5 d and nymphal stadia of
2–3 d each for the first through fourth instars and 3–4
d for the fifth instar on cabbage (Brassica oleracea L.).
Verma et al. (1993) reported stadia for the first through
fifth instars of 3.33, 4.00, 4.31, 4.54, and 3.82 d, respectively, on Indian mustard [Brassica juncea (L.)], under
controlled conditions. Singh and Malik (1993) reared
the bugs on Indian mustard seeds in the laboratory at
different times of year reporting an incubation period
and nymphal stadia (first–fifth instars combined), respectively, of 6.22 and 14–22 d in April (25.5–34 C),
3.28 and 14–20 d in May (28–30 C), and 6.27 and
29–37 d (28 C; no month given).
A few illustrations and descriptions of the egg and
immature stages have been published for B. hilaris [or
its synonyms: Bagrada cruciferarum Kirkaldy and
Bagrada picta (F.)]. Superficial illustrations were provided by Hutson (1935) (egg, second through fifth instars, and adult) and Narayanan (1958) (egg, first
C The Authors 2015. Published by Oxford University Press on behalf of Entomological Society of America.
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2
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Table 1. Plant associations or feeding records for B. hilaris
Family
Annonaceae
Apiaceae
Asteraceae
Brassicaceae
Caricaceae
Capparaceae
Chenopodiaceae
Compositae
Convolvulaceae
Cucurbitaceae
Euphorbiaceae
Fabaceae
Malvaceae
Poaceae
Species
Asimina triloba (L.)
Daucus carota L.
Bidens pilosa L.
Cynara cardunculus var. scolymus
Dendranthema grandiflorum (Ramat.)
Kitamura
Helianthus annuus L.
Lactuca sativa L.c
Schkuhria pinnata (Lam.) Kuntze ex Thell.
Sonchus arvensis L.
Barbarea sp.
Brassica compestris L.
Brassica juncea (L.) Czern.
Brassica napobrassica (L.) Mill.
Brassica napus L.
Brassica oleracea L. var. italica Plenck.
Brassica oleracae L. var. acephala D.C.
Brassica oleracea L. var. botrytis subvar.
cauliflora D.C.
Brassica oleracea L. var. capitata L
Brassica oleracea var. gongylodes
Brassica oleracea L. var. gemmifera D.C.
Brassica rapa subsp. Pekinensis Rupr.
Brassica rapa L. subsp. oleifera (D.C.) Metzg.
Brassica rapa var. rapa L.
Capsella busa-pastoris (L.) Medikus
Eruca sativa Mill.
Hirschfeldia incana (L.) Lagrese-Fossat
Iberis spp.
Lepidium alyssoides A. Gray
Lepidium latifolium L.
Lepidium capense Thunb.
Lobularia matitima (L.) Desvaux
Mathiola spp.
Raphanus sativus L.
Raphanus raphanistrum L.
Sisymbrium irio L.
Sisymbrium capense Thunb.
Carica papaya L.
Capparis spinosa L.
Beta vulgaris L.
Chenopodium album L.
Spinacia oleracea L.c
Dahlia sp.
Convolvulus arvensis L.
Citrullus lanatus (Thunb.)c
Cucumis melo L. subsp.
melo var. cantalupo Ser.
Euphorbia hirta L.
Arachis hypogaea L.
Indigofera sp.
Pisum sativum (L.)
Vigna mungo (L.) Hepper
Vigna radiata (L.) R. Wilczek
Abelmoschus esculentus L.
Alcea sp.
Gossypium hirsutum L.
Avena sativa L.
Cynodon dactylon (L.) Pers.
Cyperus rotandus L.
Dactyloctenum aegyptium (L.)
Beauv.
Hordeum vulgare L.
Oryza sativa L.
Pennisetum glaucum (L.) R. Br.
Saccharum officinarum L.
Saccharum spontaneum L.
Sorghum bicolor (L.) Moench
subsp. drummondii (Nees ex
Steud.) de Wet & Harlan
Common name
Locationa
Referenceb
pawpaw
carrot
black jack
artichoke
chrysanthemum
NG
NG
South Africa
NG
NG
Gunn 1918
Gunn 1918
Gunn 1918
Gunn 1918
Gunn 1918
sunflower
lettuce
pinnate false threadleaf
Field sow thistle, corn sow thistle
winter cress
wild rape
Indian mustard
rutabaga
rapeseed, canola
broccoli, broccoflower
kale, collards
cauliflower
USA
NG
South Africa
India
South Africa
South Africa
Sri Lanka
USA (AZ)
India
USA (CA)
USA (CA)
Sri Lanka
Bundy et al. 2012
Gunn 1918
Gunn 1918
Singh and Malik 1993
Gunn 1918
Gunn 1918
Hutson 1935
Palumbo and Natwick 2010
Singh and Malik 1993
Arakelian 2008–2009
Arakelian 2008–2009
Hutson 1935
cabbage
kohlrabi, knoll khol
Brussels sprouts
Chinese cabbage
field mustard
turnip
shepherd’s purse
salad rocket, arugula
shortpod mustard
candytuft
mesa pepperwort
perennial pepperweed
cape pepper-grass
sweet alyssum
stock
radish
jointed charlock
London Rocket
Cape mustard
papaya
caper
sugarbeet
lamb’s quarters
spinach
dahlia
field bindweed
watermelon
cantaloupe
NG
India
USA
USA (AZ)
India
NG
USA (AZ)
South Africa
USA (AZ)
USA
USA (NM)
USA (NM)
South Africa
NG
NG
NG
South Africa
USA (AZ)
South Africa
USA (CA)
Italy
NG
India
USA (AZ)
NG
India
USA (AZ)
USA (AZ)
Gunn 1918
Rakshpal 1949
Bealmear et al. 2012
Palumbo and Natwick 2010
Azim and Shafee 1986
Gunn 1918
Reed et al. 2013
Gunn 1918
Reed et al. 2013
Gunn 1918
Bundy et al. 2012
Bundy et al. 2012
Gunn 1918
Gunn 1918
Gunn 1918
Gunn 1918
Gunn 1918
Reed et al. 2013
Gunn 1918
Arakelian 2008–2009
Ahuja et al. 2008
Gunn 1918
Singh and Malik 1993
Palumbo and Natwick 2010
Gunn 1918
Singh and Malik 1993
Palumbo and Natwick 2010
Palumbo and Natwick 2010
pill-pod spurge
peanut, groundnut
Indigo
pea
black gram
green gram
okra
hollyhock
cotton
oats
Bermuda grass
nut grass
Egyptian crowfoot grass
India
Africa, Asia
India
NG
India
India
India
NG
Africa, India
NG
USA
India
India
Singh and Malik 1993
Hill 1975
Narayanan 1954
Gunn 1918
Gupta and Gupta 1970
Gupta and Gupta 1970
Singh and Joshi 2004
Gunn 1918
Hill 1975
Gunn 1918
Reed et al. 2011
Singh and Malik 1993
Sandhu 1975
barley
rice
pearl millet
sugarcane
Kans grass (kahi)
Sudan grass
NG
India
India
India
India
USA
Gunn 1918
Rakshpal 1949
Gupta and Gupta 1970
Narayanan 1954
Sandhu 1975
Reed et al. 2011
(continued)
2015
TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
3
Table 1. Continued
Family
Polygonaceae
Rhamnaceae
Rubiaceae
Solanaceae
Tropaeolaceae
Zygophylaceae
Species
Sorghum halepense (L.) Pers.
Sorghum vulgare (L.) Pers.
Triticum aestivum L.
Zea mays L.
Polygonum plebeium R.Br.
Ziziphus rotundifolia (Burm. f.)
Wight & Arn.
Cofea spp.
Physalis peruviana L.
Solanum tuberosum L.
Nasturtium integrifolium
(Nutt.) Kuntze
Tribulus terrestris L.
Common name
Locationa
Referenceb
Johnson grass
sorghum
wheat
maize
knotweed
wild jujube
India
India
South Africa
India
India
India
Sandhu 1975
Sandhu 1975
Gunn 1918
Halbert and Eger 2010
Singh and Malik 1993
Singh and Malik 1993
coffee
cape gooseberry
Potato
nasturtium
USA
NG
India
South Africa
Rakshpal 1949
Gunn 1918
Hill 1975
Gunn 1918
puncturevine
India
Sandhu 1975
a
NG: location not given.
b
Only reference of first report of host plant given here.
c
Host records where damage to the plant was not observed.
through fifth instars, and adult). Brief descriptions of
the egg and first through fifth instars were reported by
Rakshpal (1949), Verma et al. (1993), and Azim and
Shafee (1986). Azim and Shafee also included illustrations and a key to the nymphal instars.
Several Old World reports are available on the natural enemies of B. hilaris. Eggs are attacked by scelionid
parasitoids (Hymenoptera) including Gryon karnalense
(Chacko and Katiyar) (as Hadrophanurus karnalensis)
(Chacko and Katiyar 1961), Telenomus samueli (Mani)
(as Liophanurus samueli) (Mani 1941, Mani and
Sharma 1982), Trissolcus spp. (Cheema et al.1973,
Ghosal et al. 2005), and Tiphodytes sp. (Samuel 1949).
Adults are attacked by the tachinid parasitoids (Diptera) including Alophora pusilla Meigan (Rakshpal
1949, 1954; Crosskey 1976; Azim and Shafee 1986),
Hyalomya pusilla Meigan (Cheema et al. 1973), and A.
indica (Mesnil) (Cheema et al. 1973, Crosskey 1976).
Rakshpal (1954) suggested that the sex of the bug attacked determined the sex of A. pusilla. Thakar et al.
(1969) reported the predatory mite Bochartia sp. (Acarina: Erythraeidae) feeding on nymphs and adults.
Although numerous studies on the biology of B. hilaris have been published for the Old World, little information is available for North America. In south central
Asia, adults have been collected throughout the year
(Batra 1958, Narayanan 1958, Atwal 1959). They were
most active from October through March feeding on
mature pods of winter crops. During the spring, they
were found beneath debris in harvested fields (Sandhu
1975). In India, Singh and Malik (1993) also reported
year-round survival of adults with peak populations during seedling stages from October to November and another peak during harvest from March to April.
Populations were relatively low from December to January when temperatures were lowest. In the New
World, Garrison (2008–2009) reported adults and
nymphs in June and September to November in California feeding on rape (Brassica napus L.), Alyssum
sp., “daikons,” turnips, and broccoli (B. oleracea). He
also stated it had spread to the Yuma region of Arizona
where it fed on Brassica weeds, seed crops, canola, and
cotton. Bundy et al. (2012) reported populations of
nymphs and adults from several localities in New
Mexico on a variety of host plants, including mesa pepperwort (Lepidium alyssoides A. Gray), from April
through September (2010–2011). Reed et al. (2013) reported two main peaks (life stages not given) in B. hilaris populations in southern California, April to May
and September to October, with adults overwintering
in the soil near a food source. Palumbo (2013) found
adult survival throughout the year in Arizona with
peaks from September to October and March to April
on a variety of host plants including broccoli and
cabbage.
Among the localities mentioned by Bundy et al.
(2012) was Las Cruces. Here, high numbers were observed on mesa pepperwort, suggesting that a detailed
life history study was possible. Presented here is the
life history of B. hilaris in and around Las Cruces
including information on laboratory rearing under controlled conditions and descriptions and illustrations of
the immature stages.
Materials and Methods
Life History. A field survey was conducted from
January 2012 to August 2014 at several sites in and
around Las Cruces, NM. Sampling primarily involved
two host plants: London rocket (Sisymbrium irio L.), a
winter annual species, and mesa pepperwort, a summer
perennial species. For London rocket, collecting was
primarily from December through June and for mesa
pepperwort, May through November; however, a few
individuals were collected outside these periods.
Although London rocket was one of the two primary
hosts, no specimens were collected from this host during the beginning of this project (15 January through
25 May 2012), even though it was sampled extensively
during this time. Responding to a report by a stakeholder of these bugs on broccoli, we began sampling
this plant on 7 November 2012, which continued
through the third week in March when the plants had
senesced. Also added for sampling on 5 December
2012 was a patch of London rocket that was infested
with B. hilaris near the broccoli field, which continued
through 15 May 2013 when plants were destroyed by
the grower.
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ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Counts of all life stages and notes on the bugs’ activities were taken weekly. Samples of mesa pepperwort
consisted of six plants per field site utilizing the beatbucket technique (modified from Bundy and McPherson 2009). Leaves from a single plant were shaken rapidly over a bucket (38.8 by 31.9 by 14.4 cm), and
dislodged insects were counted, determined to
instar(s), if possible, and released. Individuals that
could not be determined to instar in the field were
taken to the laboratory for closer examination. During
the winter months, samples of mesa pepperwort were
expanded to include observations of the bugs on leaf
litter and at the bases of the plants, which remained
green throughout the winter months.
Samples of London rocket and broccoli consisted of
whole plant samples (six plants per species) rather than
the beat-bucket technique. For London rocket, the
beat-bucket technique was ineffective for plants, which
varied in size significantly from winter through spring.
For broccoli, the beat bucket technique was too damaging to the plant.
During weekly sampling of various hosts, bugs often
were observed on the soil near the bases of the plants.
Therefore, small areas of soil within a radius of 20 cm
from the bases were included during sampling to
obtain a better representation of the population.
For all sampling techniques, counts of adults and
each immature stage were recorded, and plant material
was inspected for the presence of eggs. Representative
samples of nymphs were preserved in 80% ethanol
(EtOH) to check field determinations and for use in
immature descriptions (see below). In late February
2013, soil samples from around the perimeter of
plants were collected in hopes of recovering eggs. Soil
(1 liter, total) was collected from the base of each of
six plants using a garden spade to scrape the loose soil
(1.5 cm deep), taken to the laboratory in plastic bags,
and washed through a set of sieves (203.2 mm in diameter, 50.8 mm in depth): number 10 (2.0 mm, mesh
size), number 20 (0.83 mm, mesh size), and number 35
(0.50 mm, mesh size). Debris from the last sieve (number 35) was placed in a large dish (15 cm in diameter
and 2 cm in depth) with water (10 ml), and the contents were examined for eggs.
Laboratory Rearing. Adults (100 females, 100
males) were collected from mesa pepperwort in and
around Las Cruces from early August to mid-October
2012, taken to the laboratory, and placed in an ovipositional cage. The cage consisted of a large terrarium (61
by 32 by 41 cm) lined with paper toweling (to absorb
excess water) and provided with cuttings (30 cm in
length) of mesa pepperwort with fruiting structures.
Cuttings were inserted into a block of floral foam (7 by
7 by 5.5 cm), which previously had been placed in a
small plastic cup (350 ml) filled with distilled water and
covered with paper toweling to avoid excess moisture.
A small petri dish (9 cm in diameter and 1.5 cm in
depth) with distilled water was added to the cage to
provide a water source and filled with cotton balls to
prevent the bugs from drowning. One or two narrow
cheesecloth strips (50.8 by 2 cm) were draped over
the plants and served as ovipositional sites. Adults were
transferred to a new ovipositional cage daily and the
old plant material and toweling examined for eggs. The
eggs were removed with a paint brush or fine-tipped
forceps and placed on moistened filter paper in petri
dishes (10 cm in diameter and 2 cm in depth). The
old cages then were cleaned with soap and water to
ensure no eggs remained.
Nymphs were kept in petri dishes similar to those
for eggs but provided with cuttings (3 cm) of the
fruiting structures of mesa pepperwort for food. Petri
dishes were examined daily, molts recorded, and exuviae removed. Nymphs were grouped by molting dates
to determine stadia. Filter paper was moistened daily,
and food was replaced as needed, approximately
every 2 d.
Eggs, nymphs, and adults were maintained in the
laboratory in an incubator at 25 6 0.01 C under a
photoperiod of 14:10 (L:D) h.
Description of Immature Stages. Descriptions of
the egg and instars are based on 10 individuals. Eggs
and first instars were obtained in the laboratory from
field-collected females; second through fifth instars
were field-collected. All immature stages were preserved in 80% EtOH. Illustrations were made on a
lightbox from digital photographs taken on a Canon
EOS 5D mark II with the Passport system. Digital renderings were created using Adobe Illustrator CS6 and
Photoshop CS6. Measurements of most structures (in
mm) were made using an ocular micrometer; those for
micropylar processes (in mm) were made from scanning
electron micrographs.
Scanning electron micrographs were used in the
description of the egg. Eggs were fixed 2.5% glutaraldehyde, rinsed with 0.1 M imidazole buffer, and run
through a critical point dryer. They were mounted on a
carbon adhesive tab, sputter-coated with gold, and
examined with a scanning electron microscope
(S3400N II, Hitachi High Technologies, Pleasanton,
CA) at an accelerating voltage of 10 kV.
Statistics. Measurements
are
expressed
as
means 6 SE; SEs <0.005 are listed as 0.00. Data on
total developmental time of male and female bugs were
analyzed with the SAS (SAS Institute 2002–2008)
TTEST procedure. Level of significance selected was
0.05.
Voucher Specimens. Selected samples of eggs and
instars have been vouchered in the New Mexico State
Arthropod Museum in Las Cruces, NM.
Results and Discussion
Life History. Adults (n ¼ 8,264) and nymphs
(18,060) of B. hilaris were found continuously throughout the year, including the winter months, on or near
their host plants (Figs. 1 and 2). Mating pairs
(n ¼ 1,687) were found continuously (at least one pair
each week) through the year on mesa pepperwort
(n ¼ 1,294), peaking from mid-June to early October;
and discontinuously on London rocket (n ¼ 393), peaking from early January to mid-May. (Note: percentages
of life stages at low levels are not visible for some dates
on the life history graphs [Figs. 1–6]).
2015
TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
/ / / / /
Jan
Feb
Mar
Apr
May
June
July
Aug
Sept
Oct
5
/ // /
Nov
Dec
50
25
ADULT
0
(N=8264)
25
50
25
EGG 0
(N=642)25
50
1ST 25
INSTAR 0
(N=2715)25
50
2ND 25
INSTAR 0
(N=4249)25
50
3RD 25
INSTAR 0
(N=4290)25
50
4TH 25
INSTAR 0
(N=3061)25
50
5TH 25
INSTAR 0
(N=3745)25
50
Fig. 1. Composite field life cycle of B. hilaris on mesa pepperwort, London rocket, and broccoli in Las Cruces, NM,
2012–2014. Percentage in each sample of total individuals of each stage. Percentages less than 5% may not be visible due to
size of figure. Top horizontal bar represents actively growing periods of each host plant: gray ¼ mesa pepperwort; white
diagonal stripes ¼ mesa pepperwort þ broccoli overlap; black diagonal stripes ¼ broccoli þ London rocket overlap;
black ¼ London rocket þ mesa pepperwort overlap.
/ / / / /
Jan
Feb
Mar
Apr
May
June
July
Aug
Sept
Oct
/ // /
Nov
Dec
50
ADULT
(N=8264)
25
0
25
50
25
EGG 0
(N=642) 25
50
1ST 25
INSTAR 0
(N=2715) 25
50
2ND 25
INSTAR 0
(N=4249)25
50
3RD 25
INSTAR 0
(N=4290)25
50
4TH 25
INSTAR 0
(N=3061) 25
50
5TH 25
INSTAR 0
(N=3745) 25
50
Fig. 2. Composite field life cycle of B. hilaris on mesa pepperwort, London rocket, and broccoli in Las Cruces, NM,
2012–2014. Percentage in each sample of total individuals of the same stage. Percentages less than 5% may not be visible due
to size of figure. Top horizontal bar represents actively growing periods of each host plant: gray ¼ mesa pepperwort; white
diagonal stripes ¼ mesa pepperwort þ broccoli overlap; black diagonal stripes ¼ broccoli þ London rocket overlap;
black ¼ London rocket þ mesa pepperwort overlap.
6
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
/ / / / /
Jan
Feb
Mar
Apr
May
June
July
Aug
Sept
Oct
Nov
/ /
Dec
50
25
ADULT
0
(N=1884)
25
50
25
EGG 0
(N=50) 25
50
1ST 25
INSTAR 0
(N=136) 25
50
2ND 25
INSTAR 0
(N=922) 25
50
3RD 25
INSTAR 0
(N=992) 25
50
4TH 25
INSTAR 0
(N=1051)25
50
5TH 25
INSTAR 0
(N=1409)25
50
Fig. 3. Field life cycle of B. hilaris on London rocket and broccoli in Las Cruces, NM, 2012–2014. Percentage in each
sample of total individuals of each stage. Percentages less than 5% may not be visible due to size of figure. Top horizontal bar
represents actively growing periods of each host plant: gray ¼ broccoli; black diagonal stripes ¼ broccoli þ London rocket
overlap; black ¼ London rocket; white ¼ neither host present.
/ / / / /
Jan
Feb
Mar
Apr
May
June
July
Aug
Sept
Oct
Nov
/ /
Dec
50
25
ADULT
0
(N=1884)
25
50
25
EGG 0
(N=50) 25
50
1ST 25
INSTAR 0
(N=136) 25
50
2ND 25
INSTAR 0
(N=922) 25
50
3RD 25
INSTAR 0
(N=992) 25
50
4TH 25
INSTAR 0
(N=1051)25
50
5TH 25
INSTAR 0
(N=1409)25
50
Fig. 4. Field life cycle of B. hilaris on London rocket and broccoli in Las Cruces, NM, 2012–2014. Percentage in each
sample of total individuals of the same stage. Percentages less than 5% may not be visible due to size of figure. Top horizontal
bar represents actively growing periods of each host plant: gray ¼ broccoli; black diagonal stripes ¼ broccoli þ London rocket
overlap; black ¼ London rocket; white ¼ neither host present.
2015
TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
Jan
Feb
Mar
Apr
May
June
July
Aug
Sept
Oct
7
Nov
Dec
50
25
ADULT
0
(N=6380)
25
50
25
EGG 0
(N=592)25
50
1ST 25
INSTAR 0
(N=2579)25
50
2ND 25
INSTAR 0
(N=3327)25
50
3RD 25
INSTAR 0
(N=3298)25
50
4TH 25
INSTAR 0
(N=2010)25
50
5TH 25
INSTAR 0
(N=2336)25
50
Fig. 5. Field life cycle of B. hilaris on mesa pepperwort in Las Cruces, NM, 2012–2014. Percentage in each sample of
total individuals of each stage. Percentages less than 5% may not be visible due to size of figure.
Jan
Feb
Mar
Apr
May
June
July
Aug
Sept
Oct
Nov
Dec
50
ADULT
(N=6380)
25
0
25
50
25
EGG 0
(N=592) 25
50
1ST 25
INSTAR 0
(N=2579) 25
50
2ND 25
INSTAR 0
(N=3327)25
50
3RD 25
INSTAR 0
(N=3298)25
50
4TH 25
INSTAR 0
(N=2010) 25
50
5TH 25
INSTAR 0
(N=2336) 25
50
Fig. 6. Field life cycle of B. hilaris on mesa pepperwort in Las Cruces, NM, 2012–2014. Percentage in each sample of
total individuals of the same stage. Percentages less than 5% may not be visible due to size of figure.
Eggs (n ¼ 642) were found from early February
through early October (Figs. 1 and 2) and were deposited singly in the soil near the bases of sampled plants.
Of these, 592 (92.21%) were collected from the bases
of mesa pepperwort; the remainder was collected
from the bases of London rocket and broccoli.
Eggs never were collected on the host plants (Taylor
et al. 2014).
8
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Table 2. Duration (in days) of immature stages of B. hilaris
Stage
Sex
No. completing stadium
Mean 6 SE
Range
Cumulative mean age
7.45 6 0.06
3.35 6 0.05
7.08 6 0.13
6.39 6 0.13
7.33 6 0.18
10.25 6 0.16
41.85 6 0.36
41.88 6 0.54
41.85 6 0.43
6–9
3–5
5–11
4–10
5–12
8–14
37–50
37–50
38–45
7.45
10.80
17.88
24.27
31.60
41.85
Males þ Females
Males
Females
100
91
84
76
73
68
68
40
28
a
Egg
First instar
Second instar
Third instar
Fourth instar
Fifth instar
Egg through fifth instar
a
Of 2,866 eggs laid in the laboratory, 100 were selected for this study.
The number of generations per year in this species
was difficult to determine because of the multiple host
plants, the extreme overlap of stadia, the variation in
numbers per sample, and the absence of reproductive
diapause (see above). All life stages, with the exception
of the eggs, were present in every month of the year;
eggs were not found in January, November, or December (Figs. 1 and 2). Adults and all nymphal instars overwintered (Figs. 1 and 2). Based on the seasonal
abundance of the various life stages, the data roughly
suggest that overwintering adults reproduced in late
winter and spring and their offspring became adults in
June–August (Figs. 1 and 2). The summer (second)
generation reproduced primarily during July–September, and their offspring reached adults during September–November. These data suggest that this species is
bivoltine. However, there is some evidence that there is
a partial third generation in southern New Mexico,
with overwintering late instars reaching adult in January and February (Figs. 1 and 2).
Dividing the life cycle of B. hilaris into the time
spent on London rocket (the winter annual) and broccoli, and the time spent on mesa pepperwort (the
summer annual), further supports our contention that
this species is bivoltine (Figs. 3–6). Specifically, adults
and first through third instars were first collected on
broccoli in early November, indicating original colonization likely occurred in October, and were found discontinuously into March when the plants senesced
(Figs. 3 and 4). Adults and first through fifth instars
first were observed on London rocket during early
December and were found discontinuously into June
(Figs. 3 and 4). Eggs were found discontinuously from
February into May.
Adults were found continuously throughout the year
on mesa pepperwort, whereas the other life stages
were found discontinuously (Figs. 5 and 6). However,
all stages were most abundant from July through October (85.4% of total individuals collected; Fig. 6). Eggs
were collected discontinuously from late February
through mid-December; first instar nymphs from February into December; and second through fifth instars
from early January discontinuously through December.
Although the majority of the plants had senesced during the winter months, observations at this time showed
that the bugs moved to the bases of the plants where
some green leaves and debris remained and provided
cover.
Laboratory Rearing. Eggs (n ¼ 2,866) were laid in
one of two general patterns: singly (n ¼ 2,760, 96.30%)
or in groups, 2–11 eggs (n ¼ 106); those laid in a group
were either deposited in a mass (n ¼ 38, 35.85%), or in
a row (n ¼ 68, 64.15%). Eggs laid singly were deposited
on the four available substrates: cheesecloth (n ¼ 1,891,
68.51%), paper toweling (n ¼ 653, 23.66%), cage surfaces (n ¼ 205, 7.43%), and plant material (n ¼ 117,
4.24%); eggs laid in groups were deposited on cheesecloth (n ¼ 36, 33.96%), paper toweling (n ¼ 35,
33.02%), cage surfaces (n ¼ 18, 16.98%), and plant
material (n ¼ 17, 16.04%). Adults usually were found
on the plant material.
Egg color ranged from cream-colored to light brown
when laid and changed to a dark pink color by day 5.
Eyespots usually appeared at day 4, and the egg burster
became visible by day 5. The incubation period ranged
from 6–9 d and averaged 7.45 d (Table 2).
The first instar emerged through the pseudopercular
opening at the cephalic end of the egg with the aid of an
egg burster. The first, second, third, fourth, and fifth stadia averaged 3.35, 7.08, 6.39, 7.33, and 10.25 d, respectively. Total development averaged 41.85 d and did not
differ significantly between sexes (males, 41.88 d;
females, 41.85 d; t ¼ 0.55; df ¼ 25; P ¼ 0.5890) (Table 2).
Descriptions of Immature Stages
Egg (Figs. 7–11). Length, 0.93 6 0.03 mm; width,
0.77 6 0.02 mm. Eggs typically laid singly in the soil
(see Taylor et al. 2014) resulting in a layer of soil particles glued to the chorion (Fig. 7). Each egg subcylindrical, cream-colored to light brown; chorion spinose
(Fig. 8); pseudoperculum present at cephalic end, convex and circular, surrounded by 11–18 micropylar processes (Figs. 8 and 9); processes clavate and recurved,
smooth, 47.70 6 0.91 mm in length and 11.15 6 0.20 mm
in diameter, opening subapical, 5.83 6 0.35 mm in diameter (Fig. 10); egg burster roughly T-shaped, heavily
sclerotized, surrounded by membranous region, giving
entire structure a subquadrate appearance (Fig. 11).
Instars. The first instar is described in detail, but
only significant changes from previous instars are
described for subsequent instars. Length was measured from the apices of the tylus and juga to the
apex of the abdomen (two measurements), width
across the metanotum and abdominal segments 3
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TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
9
Fig. 9. Scanning electron micrograph of a B. hilaris egg
(dorsal view), 130 .
Fig. 7. Scanning electron micrograph of a B. hilaris egg
extracted from soil but covered with adhering soil particles
(lateral view), 100 .
Fig. 8. Scanning electron micrograph of a B. hilaris egg
recovered from cheese cloth (lateral view), 100 .
Fig. 10. Scanning electron micrograph of micropylar
process of B. hilaris, 1,200 .
10
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Fig. 11.
Egg burster of B. hilaris.
and 4 (two measurements). Additional measurements
are given in Table 3.
First Instar (Figs. 12 and 20). Length, 1.19 6
0.03 mm (1.20 6 0.03 mm); width, 0.71 6 0.02 mm
(0.77 6 0.02 mm). Body elliptical; widest at abdominal
segments 3–4; reddish, with head and thorax reddish
brown.
Head reddish brown, often with paired, faint brown
longitudinal stripes adjacent to inner margin of eye,
occasionally second faint pair from near base of tylus to
near posterior margin of head, converging posteriorly;
region posterior to eye and median oval area near posterior margin of head incompletely sclerotized; head
strongly declivent; anterolateral margins generally
straight, weakly arcuate near base of antenniferous
tubercle; compound eyes oval, weakly pedunculate;
tylus distinctly exceeding juga. Antennae foursegmented, often concolorous with head; apical segment longest, fusiform; ratio of antennal segment
lengths 1:1.3:1.4:3. Labium four-segmented, generally
concolorous with head.
Thoracic nota reddish brown. Pro- and mesonota
completely sclerotized, lateral margins weakly arcuate,
posterior margins weakly arcuate medially, substraight
laterally. Metanotum sclerotized in anterior two thirds,
forming plate, this plate narrowed medially, posterior
margins of plate and segment (i.e., membranous area)
straight medially; mediolongitudinal line extending
from anterior margin of pronotum through posterior
margin of metanotal plate. Ratio of pro-, meso-, and
metanotal lengths (sclerotized and membranous portion combined) 1:0.7:0.5. Pleura reddish brown;
sterna reddish. Legs concolorous with body except for
distal tip of tarsomere 2, which is dark brown.
Abdomen, reddish dorsally, with six medial plates
(plate 4 absent in first instar) and nine lateral plates,
last medial (6) and last lateral (9) apparently fused and
adjacent to sclerotized anal peritreme, which completely encircles anus; all plates reddish brown. Medial
plate 1 subrectangular, plate 2 subquadrate, plate 3
suboval, plate 5 suboval, plate 6 arcuate; medial plates
1–3 with paired ostioles. Lateral plates extending
dorsally and ventrally from lateral edge of
abdomen: dorsally, plates 1–7 subtriangular to subquadrate; lateral plate 8 rounded; pseudointersegmental
lines present mesad of lateral plates 1–7. Ventral surface mostly concolorous with corresponding dorsal surface; seven faint medial plates present: plates 1–3
paired, transverse, narrow; plates 4–7 subquadrate,
plate 7 adjacent to anal peritreme; ventral extensions of
lateral plates subquadrate. Abdominal spiracles on segments 2–6 each near lateral margin of corresponding
segment. Single trichobothrium (primary trichobothrium) posteromesad of each spiracle on segments 3–7,
arising from reddish brown sclerite.
Second Instar (Figs. 13 and 21). Length, 1.50 6
0.05 mm (1.51 6 0.05 mm); width, 0.91 6 0.02 mm
(1.14 6 0.04 mm). Head and thorax light brown.
Head, dorsally, more completely sclerotized including region posterior to eye, median oval area near posterior margin of head completely sclerotized;
longitudinal stripes darker; anterolateral margins of
head slightly emarginate; paired incipient ocelli occasionally present posteromesad of compound eyes; eyes
pedunculate; longitudinal groove present adjacent to
base of antenniferous tubercle. Ratio of antennal segment lengths 1:1.8:1.9:3.3.
Thorax, dorsally, with suture (pronotal suture)
extending diagonally from anterolateral margin to
approximately one half the length of pronotum. Proand mesonota with medial area weakly extended posteriorly, substraight laterally. Metanotal plate slightly
more sclerotized medially. Ratio of pro-, meso-, and
metanotal lengths 1:0.6:0.2. Ventral surface light
brown. Legs with tarsomere 2 usually concolorous with
other leg segments.
Abdomen, dorsally, with paired, minute transverse
sclerites along anterior margin of segment 1 midway
between lateral margins and midline; medial and lateral
plates darker; medial plate 4 consisting of paired sclerites; pair of faint white spots on second (and occasionally first) abdominal segment midway between lateral
margins and midline. Ventrally, with medial plates well
developed, plate 7 contiguous with lateral plate 9; a
medial irregular sclerotized spot occasionally present
between paired plate 3 and medial plate 4. Anal peritreme more heavily sclerotized, widest dorsally. Sclerite
surrounding each primary trichobothrium larger. Second smaller trichobothrium (secondary trichobothrium)
present on segments 3–7 adjacent to and slightly laterad of primary trichobothrium, incorporated into lateral abdominal plates. All plates from first instar larger.
Otherwise, like first instar.
Third Instar (Figs. 14–15, 22). Length, 2.30 6
0.04 mm (2.29 6 0.04 mm); width, 1.28 6 0.02 mm
(1.62 6 0.03 mm). Two color forms present, one light
(Figs. 14 and 22) and one dark (Fig. 15). Head, dorsally, with juga and portions of head adjacent to eyes
occasionally cream-colored in light form, usually lacking in dark form. Tylus and juga subequal in length;
paired incipient ocelli present posteromesad of compound eyes. Ratio of antennal segment lengths 1:2:1.9:2.9.
Pronotum often with dark ringlike spot adjacent to
pronotal suture. Mesonotum occasionally with paired
2015
TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
11
Table 3. Measurements (means 6 SE, mm) of B. hilaris instarsa
Nymph
b
Body length (juga)
Body length (tylus)c
Head length (juga)d
Head length (tylus)e
Thorax widthf
Abdomen width
Width across eyes
Synthlipsis
Antennal segments
First
Second
Third
Fourth
Beak segments
First
Second
Third
Fourth
Notal lengthsg
Pronotum
Mesonotum
Metanotum
Leg lengths
Protrochanter
Profemur
Protibia
Protarsus
Protarsomeresh
First
Second
Mesotrochanter
Mesofemur
Mesotibia
Mesotarsus
Mesotarsomeresh
First
Second
Metatrochanter
Metafemur
Metatibia
Metatarsus
Metatarsomeresh
First
Second
First instar
Second instar
Third instar
Fourth instar
Fifth instar
1.19 6 0.03
1.20 6 0.03
0.44 6 0.01
0.47 6 0.01
0.71 6 0.02
0.77 6 0.02
0.53 6 0.01
0.39 6 0.01
1.50 6 0.05
1.51 6 0.05
0.47 6 0.01
0.49 6 0.01
0.91 6 0.02
1.14 6 0.04
0.66 6 0.01
0.50 6 0.02
2.30 6 0.04
2.29 6 0.04
0.65 6 0.01
0.66 6 0.01
1.28 6 0.02
1.62 6 0.03
0.86 6 0.03
0.55 6 0.06
3.22 6 0.05
3.16 6 0.05
0.87 6 0.02
0.88 6 0.02
1.92 6 0.06
2.29 6 0.07
1.26 6 0.01
0.85 6 0.01
4.62 6 0.21
4.55 6 0.21
1.04 6 0.04
0.99 6 0.04
2.61 6 0.08
3.04 6 0.13
1.64 6 0.03
1.05 6 0.03
0.09 6 0.01
0.12 6 0.01
0.13 6 0.01
0.27 6 0.01
0.11 6 0.01
0.20 6 0.01
0.21 6 0.01
0.36 6 0.01
0.16 6 0.00
0.32 6 0.00
0.31 6 0.01
0.46 6 0.01
0.24 6 0.01
0.55 6 0.01
0.48 6 0.01
0.58 6 0.01
0.27 6 0.01
0.88 6 0.02
0.68 6 0.01
0.76 6 0.01
0.12 6 0.00
0.15 6 0.00
0.09 6 0.01
0.17 6 0.00
0.17 6 0.00
0.26 6 0.01
0.12 6 0.00
0.20 6 0.00
0.25 6 0.00
0.41 6 0.00
0.17 6 0.01
0.26 6 0.00
0.34 6 0.01
0.58 6 0.01
0.22 6 0.01
0.34 6 0.01
0.46 6 0.02
0.78 6 0.01
0.31 6 0.01
0.44 6 0.01
0.15 6 0.01
0.11 6 0.00
0.08 6 0.01
0.22 6 0.00
0.14 6 0.01
0.04 6 0.00
0.35 6 0.01
0.27 6 0.01
0.05 6 0.02
0.57 6 0.01
0.53 6 0.01
0.14 6 0.00
0.86 6 0.02
1.01 6 0.04
0.08 6 0.01
0.12 6 0.01
0.25 6 0.01
0.28 6 0.01
0.22 6 0.01
0.18 6 0.01
0.33 6 0.01
0.42 6 0.01
0.27 6 0.00
0.25 6 0.01
0.50 6 0.01
0.60 6 0.03
0.37 6 0.01
0.34 6 0.01
0.74 6 0.01
0.91 6 0.01
0.51 6 0.01
0.45 6 0.02
1.12 6 0.03
1.29 6 0.04
0.76 6 0.02
0.07 6 0.00
0.17 6 0.00
0.12 6 0.01
0.27 6 0.01
0.31 6 0.01
0.22 6 0.00
0.10 6 0.00
0.20 6 0.01
0.17 6 0.01
0.40 6 0.01
0.46 6 0.01
0.28 6 0.01
0.16 6 0.01
0.25 6 0.01
0.26 6 0.00
0.63 6 0.02
0.68 6 0.01
0.38 6 0.01
0.23 6 0.01
0.35 6 0.00
0.34 6 0.01
0.92 6 0.02
1.01 6 0.01
0.53 6 0.01
0.39 6 0.01
0.46 6 0.02
0.47 6 0.01
1.32 6 0.03
1.41 6 0.04
0.79 6 0.01
0.08 6 0.01
0.17 6 0.00
0.13 6 0.00
0.30 6 0.01
0.35 6 0.01
0.23 6 0.01
0.11 6 0.00
0.20 6 0.00
0.18 6 0.00
0.45 6 0.01
0.57 6 0.01
0.30 6 0.01
0.17 6 0.01
0.27 6 0.00
0.26 6 0.00
0.71 6 0.02
0.88 6 0.01
0.41 6 0.01
0.24 6 0.01
0.35 6 0.01
0.36 6 0.00
1.05 6 0.02
1.33 6 0.02
0.53 6 0.02
0.41 6 0.01
0.46 6 0.02
0.46 6 0.02
1.56 6 0.05
1.91 6 0.07
0.85 6 0.01
0.08 6 0.00
0.17 6 0.00
0.12 6 0.00
0.21 6 0.00
0.18 6 0.01
0.29 6 0.00
0.26 6 0.02
0.38 6 0.01
0.44 6 0.01
0.49 6 0.02
a
Measurements are based on 10 individuals per instar.
Measured from tip of juga to apex of abdomen with head in normal declivent position.
c
Measured from tip of tylus to apex of abdomen with head in normal declivent position.
d
Measured from tip of juga to base of head in horizontal position.
e
Measured from tip of tylus to base of head in horizontal position.
f
Measured across metanotum.
g
Measured at midline.
h
Total length of segment measurements > overall length because of curvature.
b
dark transverse marks midway between lateral margins
and mediolongitudinal line. Pro- and mesonota usually
cream-colored medially in light form, pronotum creamcolored near lateral margins; pro- and mesonota usually
without cream-colored markings in dark form. Pro- and
mesonota with medial area more extended posteriorly,
posterior margins rounded medially, straight laterally in
pronotum, arcuate laterally in mesonotum; mediolongitudinal line usually extending from anterior margin of
pronotum to posterior margin of mesonotum. Ratio of
pro-, meso-, and metanotal lengths 1:0.8:0.1. Tibiae
occasionally with cream-colored mediolongitudinal
stripe in light form, absent in dark form.
Abdomen, dorsally, with paired transverse sclerites
of abdominal segment 1 larger; anterior margin of
medial plate 1 often extended anteromedially, posterior
margin often concave medially; anterior margin of
medial plate 2 often convex medially; medial plate 3
subrectangular; medial plate 4 with paired sclerites
fused, subrectangular. Paired white spots on abdominal
segments 1 and 2 usually more pronounced; additional
pair between medial plates 2 and 3. Cream-colored
spot occasionally present on lateral plates in light form,
never present in dark form. Ventrally, medial plate 7
and lateral plate 9 now fused; medial irregular sclerotized spot occasionally present between paired plates 2
12
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Fig. 12.
Fig. 13.
First instar of B. hilaris (dorsal view).
Fig. 14.
view).
Third instar of B. hilaris, light form (dorsal
Fig. 15.
view).
Third instar of B. hilaris, dark form (dorsal
Second instar of B. hilaris (dorsal view).
and 3; spot between paired plate 3 and medial plate 4
always present; all plates from second instar larger.
Otherwise, like second instar.
Fourth Instar (Figs. 16–17, 23). Length 3.22 6
0.05 mm (3.16 6 0.05 mm); width, 1.92 6 0.06 mm
(2.29 6 0.07 mm). Two color forms present, one light
(Figs. 16 and 23) and one dark (Fig. 17). Head with
juga equal to or slightly exceeding length of tylus;
paired ocelli more evident. Ratio of antennal segment
lengths 1:2.3:2:2.4.
Thorax with cream-colored markings now extending
along lateral and anterior margins of pronotum in light
form, occasionally with faint light spots on pro- and
mesonota, mesad of pronotal suture; markings greatly
reduced or absent in dark form, often present as
narrow stripe parallel to lateral margin (Fig. 17).
Lateral margins of pronotum substraight, those of
mesonotum more strongly arcuate. Mesonotum with
posterior margin rounded medially, strongly arcuate
laterally; wing pads evident, extending to abdominal
segment 2. Metanotal wing pads evident but not as
well defined as anterior pair. Ratio of pro-, meso-, and
metanotal lengths 1:0.9:0.2. Pleura often with creamcolored markings. Femora occasionally with faint
2015
Fig. 16.
view).
Fig. 17.
view).
TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
13
Fourth instar of B. hilaris, light form (dorsal
Fig. 18.
view).
Fifth instar of B. hilaris, light form (dorsal
Fig. 19.
view).
Fifth instar of B. hilaris, dark form (dorsal
Fourth instar of B. hilaris, dark form (dorsal
cream-colored mediolongitudinal stripe in light form,
absent in dark form; tibiae often with cream-colored
mediolongitudinal stripe in light form, absent in dark
form.
Abdomen, dorsally, with paired transverse sclerites
of segment 1 larger; medial abdominal plates 1–4 more
elongate transversely; anterior margin of medial plate 1
often produced laterally as irregular thin extension or
as series of small sclerites; posterior margin of medial
plate 1 slightly concave; anterior margin of medial plate
2 slightly convex; all plates from third instar larger.
Additional pair of white spots present between medial
plates 3 and 4. Cream-colored spot present on lateral
abdominal plates in light form; absent in dark form.
Otherwise, like third instar.
Fifth Instar (Figs. 18–19, 24). Length 4.62 6 0.21
mm (4.55 6 0.21 mm); width, 2.61 6 0.08 mm (3.04 6
0.13 mm). Two color forms present, one light (Figs. 18
and 24) and one dark (Fig. 19). Cream-colored markings (now often tinged with yellow or orange) of head
14
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
Figs. 20–24. Heads of B. hilaris (anterior view). (20) First instar. (21) Second instar. (22) Third instar, light form. (23)
Fourth instar, light form. (24) Fifth instar, light form.
extending to ocellus and, with slight break, to posterior
margin of compound eye in light form; absent in dark
form. Juga exceeding length of tylus. Ratio of antennal
segment lengths 1:3.3:2.5:2.8.
Thorax with cream-colored markings (now often
tinged with yellow or orange) often extending twothirds the length of lateral margins of mesonotum in
light form; light spots more evident on pro- and mesonota; all markings on pro- and mesonota, including
those on lateral margins of mesonotum, greatly reduced
or absent in dark form. Mesonotum with posterior margin more acute medially, more strongly arcuate laterally. Wing pads well developed; mesonotal pads
extending to abdominal segments 3–4, metanotal pads
same length. Ratio of pro-, meso-, and metanotal
lengths 1:1.2:0.1. Femora often with cream-colored
medial longitudinal stripe in light form, absent in dark
form.
Abdomen, dorsally, with transverse sclerites of segment 1 no longer visible; lateral extensions of medial
plate 1 well developed; medial plate 5 and lateral plates
8 fused. All plates from fourth instar larger. Otherwise,
like fourth instar.
Diagnosis. The five instars are readily distinguishable by characters other than differences in body size.
The first instar differs from later instars by the color of
the head and thorax, which is reddish brown; presence
of an incompletely sclerotized medial oval spot on the
head; eye shape, which is weakly pedunculate; absence
of a well-defined groove anterior to the eye; absence of
a pair of thoracic sutures on pronotum; presence of
five dorsal medial abdominal plates; and a single
2015
TAYLOR ET AL.: LIFE HISTORY AND LABORATORY REARING OF B. hilaris
trichobothrium posteromesad of each spiracle on segments 3–7; older instars have a darker head and thorax,
a completely sclerotized medial oval spot on the head,
pedunculate eyes, a well-defined groove anterior to the
eye; a pair of thoracic sutures on pronotum; six dorsal
medial abdominal plates, and two trichobothria posteromesad of each spiracle on abdominals segments 3–7.
The second instar can be distinguished from later
instars by the lack of light and dark forms, the lack of
cream-colored markings on the head and thorax, and
the posterior margin of the mesonotum, which is substraight laterally; older instars have light and dark
forms, light forms with cream colored markings on
head and thorax and dark forms with markings absent
or reduced; and the posterior margin of the mesonotum, which is arcuate laterally. The third instar lacks
wing pads and can be distinguished from the fourth
and fifth instars in which the wing pads are easily discernible. The fourth and fifth instars can be distinguished by the lengths of the wing pads, which reach
abdominal segment 2 in the fourth instar and abdominal segments 3 or 4 in the fifth.
Acknowledgments
We thank the following individuals (all from New Mexico
State University): undergraduate Danielle Lara and research
associate Helen Vessels, for assistance in collection of field
samples; arthropod museum curator, Graeme Davis, for assistance in imaging specimens for illustrations; Peter Cook for
use of the scanning electron microscope; and Tessa Grasswitz
for review of the manuscript.
References Cited
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Received 4 February 2015; accepted 4 May 2015.