• TABLE OF CONTENTS
HIDE
 Influence of selected sediment...
 Reparing efficient sampling plans...
 Distribution of poneromorph ants...
 Influence of temperature on susceptibility...
 Aganaspis alujai (Hymenoptera:...
 A review of parasitoids (Hymenoptera:...
 Ectoparasitism of damselflies by...
 Effect of adult diet on longevity...
 Progeny fitness of the mealybug...
 Attraction, feeding, and control...
 Revision of the bamboo delphacid...
 Review of the bamboo delphacid...
 The planthopper genus Acanalonia...
 Diversity and distribution of parasitoids...
 Toxicity of commercially available...
 Describing seasonal phenology of...
 Daily timing of mating and age...
 An overview of the red imported...
 Occurrence of clover stem borer,...
 Empicoris subparallelus (Hemiptera:...
 Densities of Heliothis virescens...
 Mating frequency of the male cactus...
 An acrobat ant, Crematogaster obscurata...
 First report of Aulacaspis yasumatsui...
 Exposure to the ripe fruit of tropical...
 Parasitoids of fall armyworm (Lepidoptera:...
 New state record and northeastern...
 Biological notes on Melanagromyza...
 Suitability of different host plants...
 Biology of laboratory-reared Lonomia...
 First record of Phyllophaga sp....
 Wolbachia-associated thelytoky...
 Survey of Tiphia vernalis (Hymenoptera:...
 Endemic parasitoids associated...
 Reduction of feeding by Schistocerca...
 First report of Ectopsocus briggsi...
 Elachista saccharella (Lepidoptera:...
 Cuterebrosis in a Florida horse:...
 Volume 90 author index
 Volume 90 subject index
 Back Matter














Group Title: Florida Entomologist
Title: The Florida entomologist
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Permanent Link: http://ufdc.ufl.edu/UF00098813/00354
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Title: The Florida entomologist
Uniform Title: Florida entomologist (Online)
Abbreviated Title: Fla. entomol. (Online)
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Language: English
Creator: Florida Entomological Society
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Publisher: Florida Entomological Society
Place of Publication: Gainesville Fla
Gainesville, Fla
Publication Date: December 2007
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Subject: Entomology -- Periodicals   ( lcsh )
Insects -- Periodicals -- Florida   ( lcsh )
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periodical   ( marcgt )
Periodicals   ( lcsh )
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Additional Physical Form: Also issued in print.
System Details: Mode of access: World Wide Web.
Language: In English; summaries in Spanish.
Dates or Sequential Designation: Vol. 4, no. 1 (July 1920)-
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 Related Items
Preceded by: Florida buggist (Online)

Table of Contents
    Influence of selected sediment physical parameters on spatial distribution of larval Glyptotendipes paripes (Diptera: Chironomidae) in three central Florida lakes
        Page 593
        Page 594
        Page 595
        Page 596
        Page 597
        Page 598
        Page 599
        Page 600
        Page 601
        Page 602
        Page 603
        Page 604
    Reparing efficient sampling plans based on a spatio-temporal chironomidae (Diptera) larval distribution model
        Page 605
        Page 606
        Page 607
        Page 608
    Distribution of poneromorph ants (Hymenoptera: Formicidae) in the Mexican state of Morelos
        Page 609
        Page 610
        Page 611
        Page 612
        Page 613
        Page 614
        Page 615
    Influence of temperature on susceptibility of Tribolium confusum (Coleoptera: Tenebrionidae) populations to three modified diatomaceous earth formulations
        Page 616
        Page 617
        Page 618
        Page 619
        Page 620
        Page 621
        Page 622
        Page 623
        Page 624
        Page 625
    Aganaspis alujai (Hymenoptera: Figitidae: Eucoilinae), a new species attacking Rhagoletis (Diptera: Tephritidae) in the Neotropical Region
        Page 626
        Page 627
        Page 628
        Page 629
        Page 630
        Page 631
        Page 632
        Page 633
        Page 634
    A review of parasitoids (Hymenoptera: Chalcidoidea) of Trialeurodes floridensis (Hemiptera: Aleyrodidae) with description of a new species from Mexico
        Page 635
        Page 636
        Page 637
        Page 638
        Page 639
        Page 640
        Page 641
        Page 642
    Ectoparasitism of damselflies by water mites in central Florida
        Page 643
        Page 644
        Page 645
        Page 646
        Page 647
        Page 648
        Page 649
    Effect of adult diet on longevity of sterile Mediterranean fruit flies (Diptera: Tephritidae)
        Page 650
        Page 651
        Page 652
        Page 653
        Page 654
        Page 655
    Progeny fitness of the mealybug parasitoid Anagyrus sp. nov. nr. Sinope (Hymenoptera: Encyrtidae) as affected by brood size, sex ratio, and host quality
        Page 656
        Page 657
        Page 658
        Page 659
        Page 660
        Page 661
        Page 662
        Page 663
        Page 664
    Attraction, feeding, and control of Rhagoletis pomonella (Diptera: Tephritidae) with GF-120 and added ammonia in Washington state
        Page 665
        Page 666
        Page 667
        Page 668
        Page 669
        Page 670
        Page 671
        Page 672
        Page 673
    Revision of the bamboo delphacid genus Belocera (Hemiptera: Fulgoroidea: Delphacidae)
        Page 674
        Page 675
        Page 676
        Page 677
        Page 678
        Page 679
        Page 680
        Page 681
        Page 682
    Review of the bamboo delphacid genus Arcofacies (Hemiptera: Fulgoroidea: Delphacidae) from China, with description of one new species
        Page 683
        Page 684
        Page 685
        Page 686
        Page 687
        Page 688
        Page 689
    The planthopper genus Acanalonia in Florida with notes on a recently introduced species, A. excavata (Hemiptera: Fulgoroidae: Acanaloniidae)
        Page 690
        Page 691
        Page 692
    Diversity and distribution of parasitoids of Anthonomus eugenii (Coleoptera: Curculionidae) from Mexico and prospects for biological control
        Page 693
        Page 694
        Page 695
        Page 696
        Page 697
        Page 698
        Page 699
        Page 700
        Page 701
        Page 702
    Toxicity of commercially available household cleaners on cockroaches, Blattella germanica and Periplaneta americana
        Page 703
        Page 704
        Page 705
        Page 706
        Page 707
        Page 708
        Page 709
    Describing seasonal phenology of the leafminer Phyllocnistis citrella (Lepidoptera: Gracillariidae) with pheromone lures: controlling for lure degradation
        Page 710
        Page 711
        Page 712
        Page 713
        Page 714
    Daily timing of mating and age at reproductive maturity in Diaphorina citri (Hemiptera: Psyllidae)
        Page 715
        Page 716
        Page 717
        Page 718
        Page 719
        Page 720
        Page 721
        Page 722
    An overview of the red imported fire ant (Hymenoptera: Formicidae) in mainland China
        Page 723
        Page 724
        Page 725
        Page 726
        Page 727
        Page 728
        Page 729
        Page 730
        Page 731
    Occurrence of clover stem borer, Languria mozardi (Coleoptera: Languriidae), on canola: a new host record
        Page 732
        Page 733
        Page 734
        Page 735
        Page 736
        Page 737
    Empicoris subparallelus (Hemiptera: Heteroptera: Reduviidae), a predatory bug new to the fauna of Florida
        Page 738
        Page 739
        Page 740
        Page 741
    Densities of Heliothis virescens and Helicoverpa zea (Lepidoptera: Noctuidae) in three plant hosts
        Page 742
        Page 743
        Page 744
        Page 745
        Page 746
        Page 747
        Page 748
        Page 749
        Page 750
    Mating frequency of the male cactus moth, Cactoblastis cactorum (Lepidoptera: Pyralidae), under laboratory conditions
        Page 751
        Page 752
    An acrobat ant, Crematogaster obscurata (Hymenoptera: Formicidae), poses an unusual conservation question in the Florida Keys
        Page 753
        Page 754
    First report of Aulacaspis yasumatsui (Hemiptera: Diaspididae) in Africa (Ivory Coast), and update on distribution
        Page 755
        Page 756
    Exposure to the ripe fruit of tropical almond enhances the mating success of male Bactrocera dorsalis (Diptera: Tephritidae)
        Page 757
        Page 758
    Parasitoids of fall armyworm (Lepidoptera: Noctuidae) from a traditional maize crop in the Mexican state of Yucatan
        Page 759
        Page 760
        Page 761
    New state record and northeastern range extension for Caenocholax fenyesi sensu lato (Strepsiptera: Myrmecolacidae)
        Page 762
    Biological notes on Melanagromyza ruelliae (Diptera: Agromyzidae), a seed feeder on the invasive Mexican petunia, Ruellia tweediana (Acanthaceae)
        Page 763
        Page 764
        Page 765
    Suitability of different host plants for nymphs of the sharpshooter Tapajosa rubromarginata (Hemiptera: Cicadellidae: Proconinii)
        Page 766
        Page 767
        Page 768
        Page 769
    Biology of laboratory-reared Lonomia obliqua (Lepidoptera: Saturniidae)
        Page 770
        Page 771
    First record of Phyllophaga sp. aff. capillata (Coleoptera: Melolonthidae) as a soybean pest in the Brazilian "Cerrado"
        Page 772
        Page 773
        Page 774
        Page 775
    Wolbachia-associated thelytoky in Diaphorencyrtus aligarhensis (Hymenoptera: Encyrtidae), a parasitoid of the Asian citrus psyllid
        Page 776
        Page 777
        Page 778
        Page 779
    Survey of Tiphia vernalis (Hymenoptera: Tiphiidae), a parasitoid wasp of Popillia japonica (Coleoptera: Scarabaeidae), in Connecticut
        Page 780
        Page 781
        Page 782
    Endemic parasitoids associated with Anastrepha spp. (Diptera: Tephritidae) infesting guava (Psidium guajava) in southern Bahia, Brazil
        Page 783
        Page 784
        Page 785
    Reduction of feeding by Schistocerca piceifrons piceifrons (Orthoptera: Acrididae), following infection by Metarhizium anisopliae var. acridum
        Page 786
        Page 787
        Page 788
        Page 789
    First report of Ectopsocus briggsi and Trichopsocus dalii (Psocoptera: Psocomorpha: Ectopsocidae and Trichopsocidae) from Iran
        Page 790
        Page 791
    Elachista saccharella (Lepidoptera: Elachistidae), a leafminer infesting sugarcane in Louisiana
        Page 792
        Page 793
        Page 794
    Cuterebrosis in a Florida horse: first equid record for North America
        Page 795
        Page 796
        Page 797
        Page 798
    Volume 90 author index
        Page 799
        Page 800
        Page 801
    Volume 90 subject index
        Page 802
        Page 803
        Page 804
    Back Matter
        Page 805
        Page 806
        Page 807
        Page 808
Full Text



Lobinske et al.: Sediment Influences on Larval Midge Distributions


INFLUENCE OF SELECTED SEDIMENT PHYSICAL PARAMETERS
ON SPATIAL DISTRIBUTION OF LARVAL GLYPTOTENDIPES PARIPES
(DIPTERA: CHIRONOMIDAE) IN THREE CENTRAL FLORIDA LAKES

RICHARD J. LOBINSKE1'2, ARSHAD ALI', ROBERT J. LECKEL, JR.1 AND JAN FROUZ3
'Mid-Florida Research and Education Center and Department of Entomology and Nematology,
Institute of Food and Agricultural Sciences, University of Florida, 2725 Binion Road, Apopka, FL 32703, USA

2Leon County Mosquito Control and Stormwater Maintenance,
501 Appleyard Drive, Suite A, Tallahassee, FL 32304, USA

'Institute of Soil Biology ASCR, Na sadkach 7, CZ-37005, Ceske Bud6jovice, Czech Republic

ABSTRACT

Spatial distribution of larvae of the pestiferous midge, Glyptotendipes paripes Edwards in
relation to selected sediment physical conditions was monitored for 1 year in 3 eutrophic
central Florida lakes. Two of these lakes (Monroe and Wauburg) supported a distribution
pattern of high densities on the firm peripheral sediments of the lakes, while the third lake
(Eustis) displayed the opposite pattern of higher densities in the soft sediments at the lake
center. Linear and multivariate analyses revealed that the presence of larval fecal pellets
and larger sized particles in the Lake Eustis sediments were strongly associated with the
latter distribution pattern. In all 3 lakes, sediment dry weight, and particle size composition
were not significantly associated with larval density. In these lakes, G. paripes distributions
appear to be strongly influenced by the physical structure of the soft, organic sediments at
the lake center; sediments with large particles and considerable accumulation of fecal pel-
lets would support higher densities of the larvae which also exist in relatively firm sedi-
ments close to the lake margin. This provides valuable information to lake managers
attempting population management of this nuisance species.

Key Words: Chironomidae, sediments, larval distribution, ecology

RESUME

Se realize un monitoreo de la distribuci6n espacial de la larva de mosca pestifera, 6C. \p'I i
dipes paripes Edwards con relaci6n a las condiciones fisicas seleccionadas de sedimento por
1 ano en 3 lagos eutr6ficos de la region central de la Florida. Dos de estos lagos (Monroe y
Wauburg) presentaron un patron de distribuci6n de altas densidades sobre los sedimentos
firmes del perif6rico de los lagos, mientras que el tercer lago (Eustis) mostr6 un patron
opuesto de altas densidades en los sedimentos blandos en el centro del lago. Los andlisis de
variables lineares y multiples revelaron que la presencia de las pelotitas de heces de las lar-
vas y particular mas grandes en los sedimentos del Lago Eustis estaban fuertemente aso-
ciados con el patron posterior de distribuci6n. En los 3 lagos, el peso seco de sedimento y la
composici6n del tamano de las particular no fueron significativamente asociados con la den-
sidad de las larvas. En estos lagos, la distribuci6n de G. paripes aparentemente esta influen-
ciada fuertemente por la estructura fisica de los sedimentos organicos blandos en el centro
del lago; los sedimentos con particular grandes y la acumulaci6n considerable de las peloti-
tas de heces soportarian las densidades mas altas de larvas las cuales tambien existen en los
sedimentos relativamente firmes cerca de la orilla del lago. Este trabajo provee una infor-
maci6n valiosa para las personas encargadas del manejo los lagos que tratan a manejar la
poblaci6n de esta fastidiosa especie.


Non-biting midges (Diptera: Chironomidae) 1988). Economic losses associated with nuisance
that emerge in large numbers from natural or midges have been estimated at millions of dollars
man-made (usually eutrophic) lakes are a source annually for many Florida cities (Anonymous
of public nuisance primarily to waterfront resi- 1977). In recent years, research has focused on
dents and businesses in central Florida (Ali 1996). larval control of nuisance midges (Ali 1996; Lobin-
Problems include restriction of outdoor activity, ske & Ali 2006). From both environmental and
soiling of buildings and outdoor equipment, aes- economic viewpoints, accurate implementation of
thetic nuisance, and possible incidences of human control efforts is required to avoid unnecessary
allergy (Gad El Rab et al. 1980; Giacomin & Tassi nontarget impacts and excessive costs. Toward







Florida Entomologist 90(4)


this end, understanding the environmental fac-
tors that influence larval distribution in relatively
large lakes is essential to enable efficient popula-
tion sampling and management. The work of
Lobinske et al. (2002) synthesized some of this
information that helped to produce a computer
model for estimating larval Glyptotendipes paripes
Edwards field population distributions (Lobinske
et al. 2004).
As early as 1959, Provost & Branch (1959) re-
ported that G. paripes larvae most commonly oc-
cur in the peripheral areas of lakes on firm sand
or mud/sand sediments, a pattern categorized as
lake type 1 by Frouz et al. (2004). Similar distri-
bution patterns of G. paripes were reported by
Callahan & Morris (1987) and Cowell & Vodopich
(1981). However, a later investigation of central
Florida lakes (Lobinske et al. 2002) revealed an
opposite distribution pattern of larval G. paripes
predominating on soft, organic sediments in the
deeper, central portions of Lake Yale (Lake
County) that contained substantial amounts of
larval chironomid fecal pellets. This distribution
was categorized lake type 2 by Frouz et al. (2004),
who also reported that fecal pellet accumulations
in the soft, organic sediments of the lake center
influenced the G. paripes larval distribution. Pre-
viously, Bradley & Beard (1969) discussed the
possible importance of chironomid excrements in-
fluencing the particle size of lake bottom sedi-
ments. Since the computer model of Lobinske et
al. (2004) was based on the lake type 1 chironomid
larval distribution pattern, it would not be effec-
tive for a type 2 lake.
This study was conducted to investigate if the
type 2 distribution pattern of G. paripes noted
above would be duplicated in an additional type 2
lake, Lake Eustis, in Lake County. Two known
type 1 lakes, Lake Monroe (Seminole and Volusia
Counties), described by Frouz et al. (2004), and
Lake Wauburg (Alachua County), described by
Lobinske et al. (2002) were selected for compari-
son in this study. Chironomid larval monitoring
programs often require routine collection of large
numbers of samples (Lobinske & Ali 2006) to de-
termine where control measures in a habitat
should be implemented. This is because many
habitats extend over hundreds or thousands of
ha, too large for an economic treatment of the en-
tire habitat. To streamline this process, several
quick and simple sediment parameters were used
in this study to determine if they could produce
reliable information for distinguishing between
lake types and estimating nuisance midge larval
distributions.

MATERIALS AND METHODS

Lakes Eustis and Monroe are each ~4,000 ha
in surface area, and Lake Wauburg ~100 ha. All 3
lakes are shallow and eutrophic. Lake Monroe is


located in the St. Johns River basin, whereas
Lake Eustis is part of the Ocklawaha River basin,
while Lake Wauburg drains into Payne's Prairie.
These lakes are located over a broad section of
central Florida.
Lakes Eustis and Wauburg were sampled for
chironomid larvae and sediments on a monthly
basis from Apr to Sep 2002, and from Jan to Mar
2003. Lake Monroe was sampled during Jun, Jul,
and Sep 2002, and then again in Feb and Mar
2003. For Lake Eustis, 40 sample locations were
randomly determined and distributed over the
entire habitat during each sampling session,
while 20 random locations each were sampled in
Lakes Monroe and Wauburg per session.
A double-hulled pontoon boat was used to
sample Lakes Eustis and Monroe, and a flat-bot-
tom jon boat used to sample Lake Wauburg. At
each sample location in Lakes Eustis and Mon-
roe, water depth was determined with a boat-
mounted depth finder (Model Humminbird Wide
100, Techsonic Industries, Eufaula, IL) while a
weighted sounding line was used to measure
depth in Lake Wauburg. For all lakes, sample lo-
cations were recorded with a Global Positioning
System receiver (Model GPS 12, Garmin Interna-
tional, Olathe, KS). Two 15 x 15-cm Ekman
dredge samples were collected from each loca-
tion. Collected sediments were subjectively clas-
sified as sand, muck, or mixed, based on distinct
visual appearance. Sand samples were composed
almost entirely of sand grains, off-white to pale
gray in color or sometimes tinted green by algae.
Muck samples were dark brown to black, had a
very loose composition, and the constituent parti-
cles were very soft. Mixed samples were com-
posed of sand grains interspersed with various
amounts of muck sediments. One dredge sample
was washed through a 350-pm mesh screen and
the retained material transferred to a 1-L wide-
mouth plastic bottle for transport to the labora-
tory. These benthic samples were examined in a
gridded white pan for midge larvae within 24 h of
collection by standard methods (Ali et al. 1977).
Chironomid larvae were identified with the keys
of Epler (2001) and counted. Nuisance midges (G.
paripes and Chironomus crassicaudatus Mal-
loch) were identified to species and others to con-
venient higher taxonomic level. From the second
dredge sample, 300-400 cm3 of the top 5 cm of sed-
iment was collected and transferred to a labeled
Whirlpak bag, transported to the laboratory, and
stored at -10C until processed and analyzed for
sediment physical parameters. At the time of pro-
cessing, the samples were thawed overnight and
each sample mixed thoroughly. With the method
described in Lobinske et al. (2002), approxi-
mately 1 g of sediment from each sample was
weighed into a tared, labeled beaker and dried at
90C for 24 h to determine percent sediment dry
weight (DW). This gives a fast determination of


December 2007







Lobinske et al.: Sediment Influences on Larval Midge Distributions


relative amount of dry matter to water present in
the sediments. To determine percent particle size
composition, about 1 cm3 of sediment was washed
through a series of 1000, 500, 250, and 125-pm
mesh sieves. The material retained by each sieve
and that passed through all sieves was respec-
tively transferred to tared, labeled beakers and
dried as above. About 1 cm3 volume of sediment
from each sample was transferred to a Petri dish,
flooded with deionized water, and examined with
a stereo dissecting microscope under 4-10x to es-
timate relative content of visibly distinguishable
components. With an eyepiece mounted grid to
estimate volume, the relative composition of fine
particulate organic matter (FPOM), sand, detri-
tus, and fecal pellets were evaluated according to
the following percent volume scale: 0 = 0%, 1 =
<10%, 2 = 10-20%, 3 = 20-50%, 4 = 50-<100%, and
5 = 100%.
Statistical analysis of sediment parameters
between lakes was conducted with Instat V. 3.05
(Graphpad Software, San Diego, CA). Compari-
sons of sediment parameters between lakes for
each basic sediment type (sand, muck, or mixed)
were made with one-way analysis of variance
(ANOVA) with Tukey-Kramer post tests. Ali et al.
(2003) reported that similar sediment parameters
had little or no significant effect on seasonal
abundance of benthic invertebrates in Lake Jesup
(Seminole County), so simple ANOVA was chosen
instead of a time-series analysis. Canonical Cor-
respondence Analysis (CCA) (ter Braak & Smi-
lauer 1998) was used to elucidate comparative in-
fluences of sediment physical parameters on the
chironomid community in each lake with the soft-
ware Canoco for Windows V. 4 (Center for Biome-
try, CPRO-DLO, Wageningen, Netherlands, and
Microcomputer Power, Ithaca, NY). This multi-
variate analysis provides an efficient means to
show how community parameters and organisms
interact with each other. For the current analysis,
only significant (P < 0.05, permutation test) sedi-
ment physical parameters chosen by forward se-
lection and separately for each lake were used in
CCA. Graphical interpretations of larval and sed-
iment distributions were prepared with Slide-
Write V. 6.1 (Advanced Graphics Software, En-
cinitas, CA).

RESULTS AND DISCUSSION

The mean values of selected parameters of
muck, sand, and mixed sediments from Lakes
Eustis, Monroe, and Wauburg are compared in
Table 1. The most consistent series of significant
differences between the lakes can be seen in the
muck sediment parameters. All physical parame-
ters of muck sediment examined from type 2 Lake
Eustis were significantly different from one or
both other lakes. Most noticeable among these
differences were the chironomid fecal pellet val-


ues, lower relative amount of FPOM, and greater
proportion of particles >250 pm in size. As hy-
pothesized for a type 2 lake, the density of larval
G. paripes was significantly greater in Lake Eus-
tis muck sediments than the other lakes. This is
similar to the muck sediments and G. paripes lar-
val distribution in type 2 Lake Yale (Frouz et al.
2004; Lobinske et al. 2002). Possibly related,
McLachlan (1976) reported that G. paripes larvae
showed a preference for larger particles in the
muck sediments of a bog lake. Important signifi-
cant differences also were noted for muck sedi-
ments of type 1 Lake Monroe. Percent particle
sizes from 125 to 1,000 pm were significantly
lower, while proportion of particles <125 pm was
considerably higher, indicating overall finer
grained organic sediments in that lake. Sediment
dry weight was significantly higher for Lake Mon-
roe muck sediments and density ofG. paripes lar-
vae was lowest. For mixed sediments, Lake Eustis
was the only lake with chironomid fecal pellets
present in the sediment. For sand substrates, sed-
iments from Lake Eustis had a significantly
higher dry weight than the other lakes, and fewer
G. paripes larvae. Due to the high sample vari-
ances, this difference was only significant be-
tween Lake Eustis and Lake Wauburg. Other chi-
ronomid midge larvae collected include C. crassi-
caudatus, Cryptochironomus sp., Pseudochiro-
nomus sp., Polypedilum sp., Tanytarsini, and
Tanypodinae; data for these midges are not
shown because of their relatively small numbers
but were included in the community analysis.
Significant correlations were detected for some
muck sediment parameters and G. paripes
log(n+1) larval density (Table 2). Larval density
was negatively correlated with proportion of
muck particles <125 pm, and positively correlated
with proportion of particles retained in 250- and
500-pm pore sieves, and with proportion of fecal
pellet content. This is consistent with the findings
of Frouz et al. (2004), who reported that type 2
sediments which overall contained high concen-
trations of fecal pellets had a higher proportion of
large particles. This allowed for higher levels of
dissolved oxygen concentration within the sedi-
ments conducive for immature G. paripes to build
longer larval tubes than in other sediment types.
Spatial distributions of FPOM, sand, and fecal
pellets are presented in Figs. 1-3. All 3 lakes show
the common distribution of muck (high FPOM
content) sediments at the basin center and more
sand content around the lake perimeter. The high
concentration of fecal pellets in Lake Eustis mid-
dle portion can be clearly seen in Fig. 1. Spatial
distributions of sediment particle size are shown
in Figs. 4-6. Differences between the 3 lakes are
clearly noticeable, including the trend showing a
relatively larger percent composition of particles
>250 pm in Lake Eustis, a more even distribution
of particles <500 pm in Lake Wauburg, and a








Florida Entomologist 90(4)


December 2007


TABLE 1. MONTHLY MEAN SD VALUES OF SELECTED SEDIMENT PHYSICAL PARAMETERS AND GLYPTOTENDIPES
PARIPES LARVAL DENSITIES IN 3 EUTROPHIC CENTRAL FLORIDA LAKES, 2002 TO 2003. ENTRIES IN A ROW
WITH THE SAME LETTER FOLLOWING ARE NOT SIGNIFICANTLY DIFFERENT (P > 0.05) BY ANOVA WITH TUKEY-
KRAMER POST-TESTS.

Parameter Type 2-Lake Eustis Type 1-Lake Monroe Type 1-Lake Wauburg


Depth (ft)
% Dry Weight
FPOM'
Detritus'
Fecal Pellets'
Sand'
% 1000 Jim
% 500 uim
% 250 uim
% 125 uim
% <125 uim
G. paripes larvae/dredge



Depth (ft)
% Dry Weight
FPOM'
Detritus'
Fecal Pellets'
Sand'
% 1000 uim
% 500 uim
% 250 uim
% 125 uim
% <125 uim
G. paripes larvae/dredge



Depth (ft)
% Dry Weight
FPOM'
Detritus'
Fecal Pellets'
Sand'
% 1000 Jim
% 500 pum
% 250 pum
% 125 pum
% <125 pum
G. paripes larvae/dredge


n =211
10.7 2.7 b
7.0 6.0 a
3.5 0.7 a
0.03 0.27 a
2.8 1.0 b
1.1 0.5 a
1.5 3.9 b
19.7 12.6 c
44.7 12.7 c
20.5 8.7 b
14.0 11.6 a
23.3 45.4 b


n=12
8.0 2.2 a
48.6 24.4 a
2.2 0.9 a
0.6 1.0 a
1.3 1.1 b
3.3 1.2 a
2.9 3.5 a
17.2 9.3 b
56.3 12.2 b
21.6 11.8 a
2.0 2.3 a
15.3 33.5 a


n=57
8.1 3.6 b
74.5 3.5 b
0.8 0.6 a
0.2 0.5 a
0.2 0.5 a
4.3 0.4 b
5.2 6.4 b
17.5 7.7 b
59.5 10.0 b
17.3 6.4 a
0.5 0.6 a
1.3 6.6 a


Muck Sediments
n =67
9.0 1.5 a
12.9 4.8 b
4.0 0.4 b
0.07 0.14 ab
0.0 0.0 a
1.5 0.8 b
0.4 0.8 a
5.2 6.8 a
12.5 12.4 a
17.6 8.7 a
64.3 23.7 c
1.2 8.8 a
Mixed Sediments
n=8
6.6 1.8 a
38.0 10.7 a
2.9 0.6 a
1.0 1.4 a
0.0 0.0 a
3.5 0.8 a
2.6 4.0 a
1.9 1.8 a
7.7 7.1 a
40.7 18.0 b
47.1 27.6 b
26.1 42.0 a
Sand Sediments
n=20
5.3 1.8 a
68.5 4.5 a
0.9 0.8 a
0.2 0.4 a
0.0 0.0 a
4.4 0.5 b
0.6 1.9 a
1.1 1.2 a
28.7 22.2 a
57.0 15.6 b
12.7 13.5 b
44.5 128.0 ab


n = 118
9.4 1.7 a
7.9 5.3 a
4.0 0.5 b
0.2 0.6 b
0.01 0.1 a
1.5 0.7 b
1.0 2.1 ab
10.2 7.5 b
34.4 14.6 b
25.0 6.1 c
29.5 21.3 b
8.2 56.9 a


N=19
6.4 2.4 a
30.8 18.4 a
2.7 1.2 a
1.6 1.1 a
0.0 0.0 a
3.1 0.8 a
3.7 5.4 a
14.4 2.6 b
51.1 9.0 b
21.1 5.2 a
9.7 10.0 a
52.4 138.2 a


N =27
5.4 2.6 a
71.3 5.6 a
1.0 0.6 a
1.1 0.9 b
0.0 0.0 a
4.0 0.2 a
4.1 2.8 ab
18.0 6.5 b
58.0 6.3 b
18.1 3.2 a
1.8 1.3 a
119.7 205.4 b


*Relative volume composition values: 0 = 0%, 1 = <10%, 2 = 10-20%, 3 = 20-50%, 4 = 50-<100%, and 5 =100%.

TABLE 2. SIGNIFICANT CORRELATION COEFFICIENTS OF MUCK SEDIMENT PARAMETERS WITH LOG(N+1) GLYPTOTEN-
DIPES PARIPES IMMATURE DENSITY IN 3 EUTROPHIC CENTRAL FLORIDA LAKES, COLLECTED 2002 TO 2003.

Parameter r P N

Particles <125 rim -0.423 <0.0001 391
Particles 250 um 0.448 <0.0001 391
Particles 500 rim 0.273 <0.0001 391
Fecal content 0.379 <0.0001 391






Lobinske et al.: Sediment Influences on Larval Midge Distributions


FPOM


Sand


Fecal Pellet


* 100%

A 50%- <100%

20%- 50%

S10% -20%

* >0% <10%

O 0%


Fig. 1. Spatial distribution and percent relative contents [Fine Particulate Organic Matter (FPOM), Sand and
Larval Fecal Pellets] of all individual sediment samples collected (n = 280) monthly from Lake Eustis (Lake County,
central Florida), Apr to Sep, 2002 and Jan to Mar, 2003.


'c C "e
'^-orf^






Florida Entomologist 90(4)


FPOM


* 100%

A 50 <100%

* 20-50%

V 10-20%

* >0 <10%

0 0%


Fig. 2. Spatial distribution and percent relative contents [Fine Particulate Organic Matter (FPOM) and Sand]
of all individual sediment samples collected (n = 95) monthly from Lake Monroe (Seminole and Volusia Counties,
central Florida), Jun, Jul and Sep in 2002 and Feb and Mar in 2003.


greater relative percent composition of smaller
particles (<250 pm) in Lake Monroe. The study
lakes show a similar pattern in sediment DW dis-
tribution (Figs. 4-6), low sediment DW at the lake
center, and increasing DW around the periphery.
In Lake Monroe, however, sediments in the lake


center had a higher DW (mostly 20-40% com-
pared to 0-20%) than the other two lakes and the
high DW sediments of the perimeter extended
further into the lake (Fig. 5). The higher DW is
probably a product of the smaller particle sizes,
which allowed the sediment to compact more


Sand


December 2007





Lobinske et al.: Sediment Influences on Larval Midge Distributions


FPOM


9 A A A
% AAA A iT 100%
A A Al;
S* A 50 <100%

20- 50%

Sand 10 20%


S* >0 <10%
A o 0%







a) V t 02 A 0
A




V f

Ot T
vs AA







of all individual sediment samples collected (n = 165) monthly from Lake Wauburg (Alachua County, central Flor-
ida), Apr to Sep, 2002 and Jan to Mar, 2003.



ida), Apr to Sep, 2002 and Jan to Mar, 2003.
iA



ida), Apr to Sep, 2002 and Jan to Mar, 2003.






Florida Entomologist 90(4)


125 pm


1~
It
J/
K.


< 125 pm


500 pm


V4
'it
I ^/


250 pm


V.


/


,4 N



E:
jA~ ~ 4r


20-40

0-20


Dry Weight


Fig. 4. Spatial percent composition of sediment particles of different sizes, as retained by specified mesh sizes,
and sediment dry weight in Lake Eustis (Lake County, central Florida), 2002-2003.


densely than in the other 2 lakes. In relation to
G. paripes larval survival, Frouz et al. (2004) re-
ported that the water in small particle dense or-
ganic sediments had lower concentrations of dis-
solved oxygen than larger particle loose sedi-
ments found in type 2 lakes.


Figure 7 shows CCA results of chironomid lar-
vae with selected sediment physical parameters
in the 3 lakes. Larvae of G. paripes, Cryptochi-
ronomus sp., and Pseudochironomus sp. recov-
ered from type 1 Lakes Monroe and Wauburg
were positively correlated with sand content as


1000 pm


~-' yr-
7,


Percent

* 80-100

* 60-80

* 40-60


December 2007


- --






Lobinske et al.: Sediment Influences on Larval Midge Distributions


1000 pm


125 pm


500 pm < 125 pm


250 pm


Percent

* 80-100

* 60-80

* 40-60

m 20-40

FI 0-20


Dry Weight


Fig. 5. Spatial percent composition of sediment particles of different sizes, as retained by specified mesh sizes,
and sediment dry weight in Lake Monroe (Seminole and Volusia Counties, central Florida), 2002-2003.


well as with sediment DW, and negatively corre-
lated with FPOM particles <125 pm in size and
water depth, in agreement with previous reports.
Corresponding with the linear regression analy-
sis, G. paripes larval density in Lake Eustis was
correlated with fecal pellet and FPOM content by
CCA (Fig. 7), while Cryptochironomus sp. and
Pseudochironomus sp. larvae were still influenced


by sediment parameters in the same way as in the
type 1 lakes. Presence of fecal pellets appears to
be the primary factor influencing G. paripes lar-
val distribution in type 2 Lake Eustis, similar to
the fecal pellet/larval distribution association re-
ported by Frouz et al. (2004) for type 2 Lake Yale.
Because fecal pellets and FPOM were closely as-
sociated in Lake Eustis sediments, and no corre-







Florida Entomologist 90(4)


125 pm


< 125 pm


N


Dry Weight


Fig. 6. Spatial percent composition of sediment particles of different sizes, as retained by specified mesh sizes,
and sediment dry weight in Lake Wauburg (Alachua County, central Florida), 2002-2003.


lation of larvae with FPOM in the other lakes was
noted, the correlation of G. paripes larvae with
FPOM was likely a co-linearity artifact.


While the full reasons for the 2 opposite G.
paripes larval distributions are not entirely clear,
an important factor certainly is the differences in


1000 pm


Percent
* 90-100

* 60-80

* 40-60

* 20-40

EI 0-20


500 pm

i>


250 pm


December 2007







Lobinske et al.: Sediment Influences on Larval Midge Distributions


-1.0 41.0

Fig. 7. Canonical Correspondence Analysis of Chironomidae community in Lakes Eustis, Monroe, and Wauburg
(central Florida) with selected physical sediment parameters.


muck sediments found at the lake center. The
presence of fecal pellets and generally larger par-
ticles apparently make the muck sediment more
attractive or suitable for survival ofG. paripes lar-
vae than the fine particle muck sediments encoun-
tered in Lake Monroe or Lake Wauburg, or the
sand sediments of Lake Eustis. Most likely, this is
a result of more oxygen availability in the fecal
pellet dominated sediments, as reported by Frouz
et al. (2004), as well as the ability of larvae to
build longer and deeper protective tubes. The abil-
ity to exploit the lake center habitat also may pro-
vide larvae with some protection from predation
by organisms that typically hunt closer to shore.
For the majority of central Florida lakes, the
lake type 1 distribution of G. paripes larvae is ex-
pected to occur and tools such as the computer
model of Lobinske et al. (2004) can be used to as-
sist managers in targeting control measures. For
lake type 2, this model would be far less effective,


but the use of the simple sediment physical pa-
rameters described in the current study would
provide an effective means to survey and target
larval populations in lake centers. When a lake
manager makes an initial survey of a lake bottom,
the presence/absence of fecal pellets in the lake
center muck sediments will determine if a lake
type 1 or lake type 2 distribution of G. paripes
should be expected. This knowledge will allow
managers to focus surveillance and control mea-
sures on those areas most likely to support nui-
sance larval densities.

LITERATURE CITED

ALI, A. 1996. Pestiferous Chironomidae and their man-
agement, pp. 487-513 In D. Rosen, F. D. Bennett, and
J. L. Capinera (eds.), Pest Management in the Sub-
tropics: Integrated Pest Management-A Florida
Perspective. Intercept, Andover, UK.











ALI, A., R. J. LOBINSKE, J. FROUZ, AND R. J. LECKEL, JR.
2003. Spatial and temporal influences of environ-
mental conditions on benthic macroinvertebrates in
northeast Lake Jesup, central Florida. Florida Sci.
66: 69-83.
ALI, A., M. S. MULLA, B. A. FEDERICI, AND F. W. PELSUE.
1977. Seasonal changes in chironomid fauna and
rainfall reducing chironomids in urban flood control
channels. Environ. Entomol. 6: 619-622.
ANONYMOUS. 1977. Economic Impact Statement. Blind
Mosquito (Midge) Task Force, Sanford Chamber of
Commerce. Seminole County, FL.
BRADLEY, W. H., AND M. E. BEARD. 1969. Mud Lake
Florida, its algae and alkaline brown water. Limnol.
Oceangr. 14: 889-897.
CALLAHAN, J. L., AND C. D. MORRIS. 1987. Survey of 13
Polk County, Florida lakes for mosquito (Diptera:
Culicidae) and midge (Diptera: Chironomidae) pro-
duction. Florida Entomol. 70: 471-478.
COWELL, B. C., AND D. S. VODOPICH. 1981. Distribution
and seasonal abundance of benthic macroinverte-
brates in a subtropical Florida lake. Hydrobiologia
78:97-105.
EPLER, J. H. 2001. Identification Manual for the Larval
Chironomidae (Diptera) of North and South Caro-
lina. A Guide to the Taxonomy of the Midges of the
Southeastern United States, Including Florida. Spe-
cial Publication SJ2001-SP13. North Carolina De-
partment of Environment and Natural Resources,
Raleigh, NC, and St. Johns River Water Manage-
ment District, Palatka, FL.
FROUz, J., R. J. LOBINSKE, AND A. ALI. 2004. Influence
of Chironomidae (Diptera) faecal pellet accumula-
tion on lake sediment quality and larval abundance
of pestiferous midge Glyptotendipes paripes. Hydro-
biologia 518: 169-177.


December 2007


GAD EL RAB, M. O., D. R. THATCHER, AND A. B. KAY.
1980. Widespread IgE-mediated hypersensitivity in
the Sudan to the'green nimitti' midge Cladotanytar-
sus lewisi (Diptera: Chironomidae). II. Identification
of a major allergen. Clinical Exp. Immunol. 41: 389-
396.
GIACOMIN, C., AND G. C. TASSI. 1988. Hypersensitivity
to chironomid Chironomus salinarius (non-biting
midge living in the Lagoon of Venice) in a child with
serious skin and respiratory symptoms. Boll. Inst.
Sieroter. Milan 67: 72-75.
LOBINSKE, R. J., AND A. ALI. 2006. Population monitor-
ing, ecology and control possibilities for nuisance
midges (Diptera: Chironomidae). Tech. Bull. Fla.
Mosq. Control Assoc. 7: 63-66.
LOBINSKE, R. J., A. ALI, AND J. FROUZ. 2002. Ecological
studies of spatial and temporal distributions of Lar-
val Chironomidae (Diptera) with emphasis on Glyp-
totendipes paripes in three central Florida lakes.
Environ. Entomol. 31: 637-647.
LOBINSKE, R. J., J. L. STIMAC, AND A. ALI. 2004. A spa-
tially explicit computer model for immature distri-
butions of Glyptotendipes paripes (Diptera:
Chironomidae) in central Florida lakes. Hydrobiolo-
gia 519: 19-27.
MCLACHLAN, A. J. 1976. Factors restricting the range of
C(. ,'..i.-. -.i-... o paripes Edwards (Diptera: Chirono-
midae) in a bog lake. J. Animal Ecol. 45: 105-113.
PROVOST, M. W., AND N. BRANCH. 1959. Food of chirono-
mid larvae in Polk County lakes. Florida Entomol.
42: 49-62.
TER BRAAK, C. J. F. AND P. SMILAUER 1998. Canoco
Reference Manual and Users Guide to Canoco for
Windows: software for Canonical Community Ordi-
nation (Version 4). Microcomputer Power, Ithaca,
NY.


Florida Entomologist 90(4)







Lobinske & Ali: Computer Model for Larval Midge Sampling


PREPARING EFFICIENT SAMPLING PLANS BASED ON A SPATIO-TEMPORAL
CHIRONOMIDAE (DIPTERA) LARVAL DISTRIBUTION MODEL

RICHARD J. LOBINSKE1'2 AND ARSHAD ALI1
'Mid-Florida Research and Education Center and Department of Entomology and Nematology,
Institute of Food and Agricultural Sciences, University of Florida, 2725 Binion Road, Apopka, FL 32703

2Leon County Mosquito Control and Stormwater Maintenance, 501 Appleyard Drive, Suite A, Tallahassee, FL 32304

ABSTRACT

A spatio-temporal larval distribution model of the nuisance midge Glyptotendipes paripes
Edwards (Diptera: Chironomidae) was used to design resource efficient sampling plans for
the assessment of locations) supporting nuisance population levels of this species in Lake
Monroe (central Florida, USA). The model used bathymetric and sediment maps with lake
water levels and temperatures of the prior month to estimate lake areas supporting rela-
tively low larval populations (<100/m2) that require minimal monitoring effort, and areas
that potentially support nuisance levels of G. paripes larvae, requiring greater sampling re-
sources. The advantage of this system is that the geographic stratification can be altered for
each sampling effort to meet prevailing conditions in the lake and without relying on a ge-
neric stratification that may not match the existing lake situation.

Key Words: Sampling plans, Chironomidae, nuisance, computer model, larvae

RESUME

El modelo de distribuci6n espacio-temporal de larvas de la mosca fastidiosa 6. .... "*.L
paripes Edwards (Diptera: Chironomidae) fue usado para el diseno de un plan de muestreo
para el uso eficiente de los recursos para la evaluaci6n de las localidades que soportan nive-
les de poblaciones fastidiosas de esta especie en el Lago Monroe (en la region central de la
Florida, USA). Este modelo utiliza mapas batimetricos mediaa de la profundidad del agua)
y de sedimento junto con los niveles de aqua y temperatures del lago del mes anterior para
estimar el area del lago que puede soportar poblaciones relativamente bajas de larvas (<100/
m2) y que requieren un esfuerzo minimo para realizar el monitoreo, y las areas que poten-
cialmente pueden soportar niveles fastidiosos de larvas de G. paripes, y que puede requerir
mas recursos para realizar el muestreo. La ventaja de este sistema es que la estratificaci6n
geografica puede ser alterada para cada esfuerzo hecho para el muestreo, tomando en cuenta
las condiciones prevalacientes en el lago y sin defender de una estratificacion gen6rica que
posiblemente no va de acuerdo con la situaci6n existente del lago.


Glyptotendipes paripes Edwards (Diptera: Chi-
ronomidae) is one of the major nuisance midge
species in some parts of central Florida, USA (Ali
1995). Because immature G. paripes are benthic
and distributed in aggregates in lakes that extend
over hundreds or thousands of hectares, popula-
tion sampling for research or management pur-
poses is tedious, time-consuming, and costly. For
highly aggregated populations, an optimized,
stratified sampling plan to examine populations is
often the most efficient method (Cochran 1963). In
this effort, after collecting preliminary samples,
adjustment is made to collect proportionally more
samples from strata with higher variances than
those with low variances. Each sampling stratum
should be internally homogeneous in relation to
the study organism. Based upon this method, G.
paripes larval populations have been successfully
examined in 3 central Florida lakes (Lobinske et
al. 2002). However, water levels in these lakes
may undergo wide fluctuations over the course of a


year or between years. The distributions of the im-
mature G. paripes are associated with water depth
in relation to sediment conditions, so the actual
strata within a lake will vary with the water level.
As part of ongoing research on nuisance chirono-
mids in central Florida, ecological data in relation
to immature G. paripes were used to develop a spa-
tio-temporal model of their distributions (Lobin-
ske 2001; Lobinske et al. 2004). For the present
study, the model was used with selected environ-
mental conditions prevailing in Lake Monroe (Vo-
lusia and Seminole Counties, Florida) to deter-
mine time-specific stratification of the lake for the
enhancement of sampling efficiency. Preliminary
sampling plans for Lake Monroe were tested on 4
occasions between Apr and Jul 2003.

MATERIALS AND METHODS

The model uses the spatial matrix techniques
of Allen et al. (1996, 2001), Brewster & Allen








Florida Entomologist 90(4)


(1997) and Brewster et al. (1997) with the com-
puter software Matlab@ (The Mathworks, Inc.,
Natik, MA) as developed by Lobinske (2001) and
Lobinske et al. (2004) to examine G. paripes pop-
ulations. The model uses spatial maps (in matrix
form) of lake bathymetry modified by lake level
deviation from mean, and sediment conditions to
determine the habitat suitability of each spatial
cell. For this study, the spatial maps were config-
ured as 50 x 50 matrices, with each cell represent-
ing approximately 3.5 ha of surface area. In addi-
tion, inputs for water temperature (as an influ-
ence on development rate), and Secchi disk trans-
parency as an estimator of phytoplankton (midge
larval food) abundance were included in the
model. Within the model, the population is repre-
sented by a three-dimensional matrix consisting
of the "X" and "Y" coordinates that provide spatial
location on the above maps, and the "Z" coordi-
nate that represents number of individuals at
each life stage. A Lefkovitch population growth
matrix (Lefkovitch 1965), detailed by Lobinske et
al. (2004) was applied to the model population at
each spatial grid cell for each time step in the cal-
culation to simulate developmental success and
survival. A dispersal function was applied at the
simulated egg laying stage. Output of projected
log(n+1) transformed population levels in each
spatial cell was in graphic (Fig. 1) and numeric
forms. Lake Monroe was one of the validation
data sets used by Lobinske et al. (2004) to test the
model and Fig. 1 is an example from that valida-
tion dataset. As can be seen clearly, under low wa-
ter levels in the lake, high populations of imma-
tures occurred farther away from the shore,
whereas under high water levels, high popula-
tions occurred close to shore. This scenario was
one of the principal reasons for the present study
because a fixed stratified sampling plan would not
be sensitive to these population shifts and thus,
would not accurately represent the actual strata.
For population management purposes, intensive
sampling is needed in areas anticipated to sup-
port nuisance (>500 immatures/m2) or near-nui-
sance (100 to 500 immatures/m2) population den-
sities. Meanwhile non-nuisance areas anticipated
to support densities below 100/m2 require less
monitoring. When tested for fit to these 2 strata
(<100 and >100 immatures/m2), the model had a
0.88 correct prediction rate for Lake Monroe
(Lobinske et al. 2004). With this success rate, it
was decided to use the model to fit sampling
strata in Lake Monroe based on current lake con-
ditions. Surveys were conducted monthly from
Apr through Jul 2003.
To determine the sampling strata for the lake
for a given month, mean water level at the United
States Geologic Survey data station 02234500
(outlet of Lake Monroe to the St. Johns River) for
the month prior to sampling was used (represent-
ing the approximate development time of imma-


Projected log(n+l) immature
G. paripeslmn
-5


Fig. 1. Example graphic output of a computer model
showing alterations in Glyptotendipes paripes larval
distributions with changing water level; top = historic
mean 1 meter, middle = historic mean, bottom = his-
toric mean + 1 meter, when other input parameters are
kept the same (Secchi disk transparency = 50 cm and
water temperature = 25.0 C).



tures that would be collected during the sample
effort), with water temperature and Secchi disk
transparency data collected during the same
prior month. The model was run for 30 simulated
days and grid cells above and below densities of
100/m2 were assessed. The high density strata
were marked and proportion of the lake area in
each stratum determined. These stratum borders
could then be plotted on lake maps (Fig. 2), and
percentages of areas in the high density stratum
are summarized in Table 1. Based on the previ-
ous month's larval G. paripes data, an optimized
sampling plan based on these 2 strata was calcu-


December 2007







Lobinske & Ali: Computer Model for Larval Midge Sampling


lated by the method of Cochran (1963), according
to the formula:
( NhSEh n
h Y (NhSEh))

where "n," is the calculated number of samples to
be collected the stratum, "N,"is the spatial area of
the stratum, "SE," is the standard error of the
mean for the stratum, and "n" is total number of
samples scheduled to be collected. For the 40 sam-
ples collected in Apr, 14 were collected from the
low density stratum and 26 from the high density
stratum. Either 20 or 40 locations were sampled
per month. A simple random generation computer
program was used to generate random sample lo-
cations in each stratum.
A Global Positioning System receiver was used
to navigate a boat to each sample location. One
Ekman dredge sample was collected at each loca-
tion and the contents processed according to stan-
dard methods (Ali et al. 1977) to enumerate G.
paripes larvae and pupae. Larvae were identified
with the keys of Epler (2001). Immature distribu-
tions in the lake were graphically plotted and
compared to the stratification plan for fit. Strati-


fied mean and standard deviation of immature
densities were calculated for each occasion.

RESULTS AND DISCUSSION

During Apr and May 2003, G. paripes popula-
tions remained below the nuisance threshold of
500 immatures/m2 at all sample locations, even in
the >100/m2 stratum (Fig. 2). During Jun 2003,
five locations were above nuisance threshold level
with mean immature density of 1086/m2 (Table 1).
Seven locations during Jul 2003 exceeded the nui-
sance threshold, with 2 supporting >10,000 imma-
tures/m2; mean immature density was 1,939/m2.
All identified locations with nuisance densities fell
within the predicted high density strata (Fig. 2).
These preliminary data indicate that the com-
puter model was an efficient tool for determining
the geographic boundaries of the sampling strata.
This can be very important in a system like Lake
Monroe, where water levels within the lake can fluc-
tuate by as much as 1.5 m annually. Because G
paripes immature distributions are heavily influ-
enced by water depth, this variation can dramati-
cally alter the locations of high larval densities
within the lake and the geographic boundaries of


Fig. 2. Monthly (Apr to Jul 2003) stratification for immature Glyptotendipes paripes sampling in Lake Monroe,
central Florida, based on projected (by computer model) densities >100/m2 (within shaded area) and <100/m2
(outside shaded area). Dots indicate the sample locations with field densities greater than the nuisance threshold
of 500/m2.







Florida Entomologist 90(4)


December 2007


TABLE 1. PERCENT RELATIVE AREA OF HIGH DENSITY STRATUM, STRATIFIED MEAN DENSITY (NUMBER/M2) AND STAN-
DARD DEVIATION OF IMMATURE GLYPTOTENDIPES PARIPES COLLECTED MONTHLY FROM LAKE MONROE, CEN-
TRAL FLORIDA, USA (APR-JUL 2003).

High density stratum
Sample month (% of lake surface area) Mean SD

Apr 14.3 61 25
May 16.0 390 179
Jun 23.4 1,086 470
Jul 10.4 1,939 1,239


the actual strata with water level changes. Combin-
ing this ability to adjust sampling stratification
with the established method of optimized stratified
sampling, the majority of the sample effort can be
targeted to the strata likely to have the highest vari-
ability in density and most likely the nuisance lev-
els. The remainder of the lake with low variability
will likely support very low populations that require
considerably less monitoring effort. The reduction
in monitoring effort with this system can be highly
advantageous. For example, Lake Monroe is ap-
proximately 4,000 ha in surface area; however, the
projected high density stratum consisted of approx-
imately 416 to 936 ha. Focusing most of the sample
effort on this smaller geographic area would facili-
tate greater sample precision in obviously less time
and labor compared to simple random sampling or
systematic sampling requiring more samples.
Additional work is proposed to compare inde-
pendently collected, systematic samples of imma-
ture G. paripes populations in Lake Monroe to fur-
ther test the efficiency of the model to predict strata
with high population levels within the lake. If con-
firmed, the model should be useful in the planning
of operational control strategies for nuisance
midges in Lake Monroe, and perhaps elsewhere.
For other lakes, detailed bathymetric and sediment
maps for each lake would be needed and rendered
into matrix format. Based on the reports ofFrouz et
al. (2004), G. paripes populations in lakes with or-
ganic sediments containing significant levels ofchi-
ronomid fecal pellets would have a dramatically
different distribution pattern that this model
would not accurately forecast. Therefore, the pre-
liminary sediment mapping should include exami-
nation of these fecal pellets to determine the suit-
ability of this model for use on that particular lake.

ACKNOWLEDGMENTS
The authors gratefully recognize the field and labo-
ratory assistance of Mr. Robert J. Leckel, Jr.

REFERENCES CITED
ALI, A. 1995. Nuisance, economic impact and possibili-
ties for control, pp. 339-364 In P. D. Armitage, P. S.
Cranston, and L. C. V. Pinder (eds.), The Chironomi-
dae: The Biology and Ecology of Nonbiting Midges.
Chapman and Hall, London, UK.


ALI, A., M. S. MULLA, B. A. FEDERICI, AND F. W. PELSUE.
1977. Seasonal changes in chironomid fauna and
rainfall reducing chironomids in urban flood control
channels. Environ. Entomol. 6: 106-111.
ALLEN, J. C., C. C. BREWSTER, J. F. PARIS, D. G. RILEY,
AND C. G. SUMMERS. 1996. Spatiotemporal modeling
of whitefly dynamics in a regional cropping system
using satellite data, pp. 11-124 In D. Gerling and R.
T Mayer (eds.), Bemisia 1995: Taxonomy, Biology,
Damage Control and Management. Intercept, An-
dover, UK.
ALLEN, J. C., C. C. BREWSTER, AND D. H. SLONE. 2001.
Spatially explicit ecology models: A spatial convolu-
tion approach. Chaos Solitons Fractals 12: 333-347.
BREWSTER, C. C., AND J. C. ALLEN. 1997. Spatiotempo-
ral model for studying insect dynamics in large-scale
cropping systems. Environ. Entomol. 26: 473-482.
BREWSTER, C. C., J. C. ALLEN, D. J. SCHUSTER, AND P.
A. STANSLY. 1997. Simulating the dynamics of Bemi-
sia argentifolia (Homoptera: Aleyrodidae) in an or-
ganic cropping system with a spatiotemporal model.
Environ. Entomol. 26: 603-616.
COCHRAN, W. G. 1963. Sampling Techniques. Wiley,
New York.
EPLER, J. H. 2001. Identification Manual for the Larval
Chironomidae (Diptera) of North and South Caro-
lina. A Guide to the Taxonomy of the Midges of the
Southeastern United States, Including Florida. Spe-
cial Publication SJ2001-SP13. North Carolina De-
partment of Environment and Natural Resources,
Raleigh, NC, and St. Johns River Water Manage-
ment District, Palatka, FL.
FROUz, J., R. J. LOBINSKE, AND A. ALI. 2004. Influence
of Chironomidae (Diptera) faecal pellet accumula-
tion on lake sediment quality and larval abundance
of pestiferous midge Glyptotendipes paripes. Hydro-
biologia 518: 169-177.
LEFKOVITCH, L. P. 1965. The study of population growth
in organisms by stages. Biometrics 21: 1-18.
LOBINSKE, R. J. 2001. Ecological Studies of Larval Glyp-
totendipes paripes (Chironomidae: Diptera) in Se-
lected Central Florida Lakes for Creating an
Exploratory Temporal and Spatial Model of Nui-
sance Populations. Ph.D. Dissertation. University of
Florida, Gainesville.
LOBINSKE, R. J., A. ALI, AND J. FROUZ. 2002. Ecological
studies of spatial and temporal distributions of larval
Chironomidae (Diptera), with emphasis on Glyptoten-
dipes paripes (Diptera: Chironomidae) in three cen-
tral Florida lakes. Environ. Entomol. 31: 637-647.
LOBINSKE, R. J., J. L. STIMAC, AND A. ALI. 2004. A spa-
tially explicit computer model for larval distributions
of C. p.i'. -.i *. paripes (Diptera: Chironomidae) in
central Florida lakes. Hydrobiologia 519: 19-27.








Quiroz & Valenzuela: Poneromorph of Morelos, Mexico


DISTRIBUTION OF PONEROMORPH ANTS (HYMENOPTERA: FORMICIDAE)
IN THE MEXICAN STATE OF MORELOS

LUIS N. QUIROZ-ROBLEDO AND JORGE VALENZUELA-GONZALEZ
Institute de Ecologia, A. C. Departamento de Entomologia. Km 2.5 Carretera Antigua a Coatepec No. 361.
Congregaci6n El Haya. C6digo Postal 91070 Xalapa, Veracruz, Mexico
E-mail: quirozl@ecologia.edu.mx; valenjor@ecologia.edu.mx

ABSTRACT

We surveyed poneromorph ants of Morelos state in south-central Mexico with pitfall traps
and hand collecting. In total, we collected 16 poneromorph species representing 9 genera
(Anochetus, Ectatomma, Gnamptogenys, Hypoponera, Leptogenys, Odontomachus, Pachy-
condyla, and Patythyrea). Although all 16 species are previously known from Mexico, our
records increase the number of poneromorphs known from Morelos from 5 to 18. The most
commonly collected species were 0. clarus, G. striatula, E. tuberculum, and H. opaciceps, all
extremely widespread Neotropical species. Some information is provided on distribution,
foraging strati, and nesting.

Key Words: ants, Poneromorphs, nesting, strati

RESUME

Se hizo un inventario de las hormigas Poneromorfas del estado de Morelos, ubicado en la re-
gi6n centro-sur de la Republica Mexicana. Los muestreos se realizaron por medio de colectas
directs y del uso de trampas. En total se encontraron 16 species poneromorfas pertene-
cientes a nueve g6neros (Anochetus, Belonopelta, Ectatomma, Gnamptogenys, Hypoponera,
Leptogenys, Odontomachus, Pachycondyla y Platythyrea), ninguna de estas species son in-
formadas por vez primera para M6xico, pero nuestros registros incrementan el numero de
poneromorfas conocidas para Morelos de 5 a 18. Las species mas comunmente colectadas
fueron 0. clarus, G. striatula, E. tuberculatum y H. opaciceps, todas ellas species neotropi-
cales ampliamente dispersas. Se proporciona alguna informaci6n de su distribuci6n, de sus
estratos de forrajeo y de anidaci6n.


Translation provided by the authors.


In general, poneromorph ants (Bolton 2003),
previously part of the subfamily Ponerinae, have
been considered "primitive" both in terms of their
morphology and habits. These ants have a wide,
mainly Pantropical distribution (Smith 1979;
Hdlldobler & Wilson 1990). Most species live in
small colonies formed by a few dozen to a maxi-
mum of some hundred individuals, with workers
that tend to be monomorphic. They are generally
predators, although some also feed on sugary nec-
tars, fruit, and the secretions of certain homopter-
ans. Most species forage alone, although some ex-
hibit recruiting behavior. Hdlldobler & Wilson
(1990) provide additional information on their bi-
ology and habits.
On the American continent, 25 genera of these
ants are known, of which 17 have been recorded
for Mexico (Bolton 1994; Brandao 1996; Lattke
2003). At present, 76 species of poneromorphs are
recognized for the Mexican Republic, of which
only 5 have been recorded for Morelos (Kempf
1972; Smith 1979; Rodriguez 1986; Brandao
1991, 1996; Cartas 1993; Bolton 1995; Lattke
1995; Quiroz & Valenzuela 1995, 2002; Castaio
1996; Rojas 1996; Longino 1998; Lachaud &


Garcia-Ballinas 2001; Durou et. al. 2002). Baroni-
Urbani (1983), Hdlldobler & Wilson (1990), and
Bolton (1994) provide keys for the identification of
Neotropical genera, and MacKay & MacKay
(1989) for those found in Mexico.
The state of Morelos is situated in south cen-
tral Mexico between 2 important geographic
zones: the Transverse Neovolcanic Belt and the
Rio Balsas Basin. The state's rugged topography
favors several climatic domains and an important
floristic diversity. In addition, Morelos is charac-
terized by the presence of Neartic, Neotropical,
and some autochthonous faunal components
(Anonymous 1981; Aguilar 1990).
The present study provides an inventory of the
poneromorph ant species that occur in the state.
It also includes some information on the distribu-
tion, nesting, and foraging habits of these species.

DESCRIPTION OF THE STUDY AREA

The state of Morelos is located between
1822'06"N and 19007'10"N, and 9803'W and
9830'08"W. Its region (covering a surface area of
4,958 km2) is divided into 33 municipalities, with







Florida Entomologist 90(4)


the state capital located in the city of Cuernavaca.
The 2 highest points in Morelos, Chichinautzin
Mountain and the Popocatepetl Volcano, are located
in the northern region at altitudes of 3,450 m and
5,452 m above sea level, respectively. From these
heights, the state's topography descends southward
to 890 m in the Jojutla Valley and then climbs again
to 1,500 m in the Sierra de San Gabriel in southern
Morelos, near Guerrero (Vidal 1980).
The climate is hot and sub-humid in most
parts of the state. Temperate and sub-humid con-
ditions predominate on mountain slopes, whereas
semi-cold to cold conditions characterize altitudes
above 2,800 m (Anonymous 1981). Tropical decid-
uous forest (TDF) is the most widely distributed
vegetation type in the state, ranging between 890
m and 1,800 m above sea level and covering more
than 50% of the state's surface area. Some dis-
turbed areas of TDF have been transformed into
thorn forests ("huizacheras"), which are generally
characterized by a great number and variety of
Acacia species. Among the ravines and canyons of
the mountainous zone, small patches of temper-
ate cloud forest can be found. Stands of pine, oak,
and fir occur from 1,800 m to 4,000 m. In the high-
est part of the state, bordering the slopes of Popoc-
atepetl, mountain prairie is prevalent. The aver-
age frost line is located at approximately 5,000 m
(Miranda & Hernandez 1963; Rzedowski 1978;
Anonymous 1981).

MATERIALS AND METHODS

We collected ants at 2-4 sites in each of the 33
municipalities of Morelos, in 7 vegetation types at
elevations ranging from 890-3500 m (Fig. 1 and Ta-
ble 1). Ants were collected from pitfall traps and
also captured directly with entomological forceps
and aspirators. Direct collection was conducted by
following a 100 m transect at each sampling site,
with ants captured from soil, under rocks, on fallen
trunks, inside trunks, and on vegetation (grass,
shrubs, and trees) to a height of approximately 2
meters. Two people carried out this procedure,
which lasted 3 h at each of the 77 sites (Table 1).
For trapping, we placed 2 pitfall traps in each
of the selected areas. They consisted of 1-L capac-
ity plastic containers with a diameter of 13 cm.
These were half-filled with commercial anti-
freeze diluted 30% with water as a preservative
and buried so that their openings were at soil
level. Finally, they were covered with a plastic
plate to prevent desiccation and contamination by
garbage and/or water if it rained. Traps were
emptied every 48 h after placement.
We preserved specimens in 70% ethanol. We
mounted vouchers and identified them using
MacKay & MacKay (1989), MacKay & Vinson
1989, Bolton 1994, and Longino's (1998) web
page. We deposited vouchers of all species at the
Institute of Ecology, Xalapa, Veracruz (IEXA).


Fig. 1. Location of the study area. Collection sites (1
to 77) are identified on the map according to numbers,
as indicated in Table 1.


RESULTS AND DISCUSSION

A total of 139 poneromorph ant samples were
obtained, with 56.3% collected directly and 46.3%
from traps. From all samples, 576 specimens were
obtained. Compared with the capture of other ant
subfamilies, relatively few poneromorph speci-
mens were obtained; for example, over 5000
Ecitoninae ants were collected under similar con-
ditions (Quiroz & Valenzuela 2006). Poneromor-
phs tend to form low-population colonies, and in-
dividuals generally forage alone. They can, how-
ever, represent a substantial percentage of ant
biomass in tropical regions due to the abundance
of certain species and to the relatively large size of
some (e.g., Cartas 1993).
Preliminary data show that in the state of Mo-
relos the subfamily Myrmicinae has the greatest
richness in terms of genera (41.5%), followed by
poneromorphs (19.5%), Formicinae (17.1%), Doli-
choderinae (12.2%), Ecitoninae (7.2%), and
Pseudomyrmecinae (2.5%).
Table 2 lists the species collected. The genus
with the greatest species richness is Gnamptoge-
nys (4 species), followed by Hypoponera, Pachy-
condyla, Odontomachus, and Ectatomma (2 spe-
cies each); and Anochetus, Belonopelta, Leptoge-
nys, and Platythyrea (1 each).
A total of 16 species of poneromorph ants were
found in Morelos. Two other species, Hypoponera
foeda and Leptogenys wheeleri, were not collected


December 2007








Quiroz & Valenzuela: Poneromorph of Morelos, Mexico


TABLE 1. COLLECTIONS SITES (CS) FOR PONEROMORPH ANTS IN THE STATE OF MORELOS, MEXICO, LISTED BY MUNIC-
IPALITY. THE COORDINATES AND ALTITUDE OF EACH SITE ARE LISTED.

Municipality CS Altitude Longitude W Latitude N


Amacuzac

Atlatlaucan

Axochiapan

Coatlan del Ri6

Cuautla

Cuernavaca



Emiliano Zapata

Huitzilac


Jantetelco

Jiutepec

Jojutla



Jonacatepec

Mazatepec

Miacatlan

Ocuituco

Puente de Ixtla

Temixco

Temoac

Tepalcingo

Tepoztlan



Tetecala

Tetla del Volcan

Tlalnepantla


Tlaltizapan

Tlaquiltenango


Tlayacapan


1. Amacuzac
2. Huajintlan
3. Atlatlaucan
4. San Juan
5. Axochiapan
6. Quebrantadero
7. Cocoyotla
8. Coatlan
9. Libramiento
10. Calder6n
11. Chamilpa
12. Reforma
13. Flores Mag6n
14. Cuernavaca
15. Tetecalita
16. Calera Chica
17. Coajomulco
18. Cempoala
19. Huitzilac
20. Carr. a Cuautla
21. Carr. a Izucar
22. Tlahuapan
23. Progreso
24. Carr. Jojutla-Ttla
25. Tehuixtla
26. Higuer6n
27. Jicarero
28. Jonacatepec
29. Tlaica
30. Justo Sierra
31. Cuauchichinola
32. Coatetelco
33. Miacatlan
34. Huecahuasco
35. Ocuituco
36. Carr. a Alpuyeca-Xoxocotla
37. Pte. De Ixtla
38. Acatlipa
39. Xochicalco
40. Temoac
41. Amilcingo
42. Agua Fria
43. Tepalcingo
44. Tepoztlan
45. Amatlan
46. Tescal
47. Cartuchos
48. Actopan
49. Tetecala
50. Tetela delV
51. Hueyapan
52. Felipe Neri
53. El Vigia
54. Temilpa
55. Ticuman
56. Acamilpa
57. Huautla
58. Quilamula
59. Tlaquiltenango
60. S. Jos6 de L
61. El bebedero


99 22'10"
99 25'30"
98 53'52"
98 55'40"
98 45'10"
98 47'23"
99 26'57"
99 26'00"
98 53'46"
98 58'42"
99 14'03"
99 13'00"
99 11'05"
99 14'03"
99 10'46"
99 46'05"
99 12'16"
99 21'03"
99 16'02"
98 47'06"
98 47'55"
99 10'04"
99 09'12"
99 16'05"
99 16'15"
99 10'50"
99 13'20"
98 48'09"
98 50'57"
99 21'43"
99 22'27"
99 19'32"
99 21'34"
98 47'08"
98 46'24"
99 14'38"
99 19'11"
99 13'07"
99 17'52"
98 46'32"
98 46'10"
98 58'01"
98 50'37"
99 05'59"
99 02'09"
99 09'02"
99 11'35"
99 24'30"
99 23'55"
98 43'46"
9841'29"
98 59'49"
98 57'34"
99 06'28"
99 10'44"
99 09'25"
99 01'18"
99 01'10"
99 09'37"
99 00'06"
98 59'40"


18 35'55"
18 36'35"
18 56'05"
18 56'09"
18 30'08"
18 31'31"
18 45'15"
18 44'43"
18 47'27"
18 51'41"
18 58'05"
18 56'06"
18 55'05"
18 55'07"
18 46'05"
18 49'53"
19 02'01"
19 06'00"
19 01'42"
18 42'07"
18 43'24"
18 53'07"
18 52'48"
18 34'05"
18 33'34"
18 34'39"
18 36'50"
18 40'58"
18 42'15"
18 43'37"
18 39'49"
18 43'46"
18 46'07"
18 56'13"
18 52'22"
18 41'06"
18 37'00"
18 48'06"
18 47'21"
18 46'23"
18 44'33"
18 35'02"
18 35'47"
18 59'07"
18 58'41"
18 58'02"
18 58'46"
18 43'53"
18 43'45"
18 53'31"
18 53'02"
19 00'24"
19 00'35"
18 42'41"
1841'54"
18 42'59"
18 26'29"
18 30'35"
18 37'44"
18 58'54"
18 55'27"







Florida Entomologist 90(4)


TABLE 1. (CONTINUED) COLLECTIONS SITES (CS) FOR PONEROMORPH ANTS IN THE STATE OF MORELOS, MEXICO,
LISTED BY MUNICIPALITY. THE COORDINATES AND ALTITUDE OF EACH SITE ARE LISTED.

Municipality CS Altitude Longitude W Latitude N

Totolapan 62. Tepetlixpita 1870 98 55'40" 18 58'38"
63. Nepopualco 2040 98 56'34" 18 59'54"
Villa de Ayala 64. Rafael Merino 1133 98 59'04" 18 44'37"
65. Moyotepec 1100 98 59'35" 18 43'12"
66. Chinameca 1050 98 59'46" 18 37'14"
Xochitepec 67. Alpuyeca 1050 99 15'29" 18 44'37"
68. Alcanfores 1060 99 16'13" 18 42'27"
Yautepec 69. Cocoyoc 1320 98 58'55" 18 53'03"
70. Caion de L. 1240 99 06'32" 18 51'27"
71. San Isidro 1105 99 05'35" 18 49'17"
Yecapixtla 72. Yecapixtla 1500 98 51'07" 18 52'52"
73. Xochitlan 1740 98 48'45" 18 53'13"
Zacatepec 74. Chiverias 945 99 13'14" 18 39'14"
75. Galeana 930 99 12'35" 18 38'28"
Zacualpan 76. Zacualpan 1640 98 45'57" 18 47'02"
77. Tlacotepec 1740 98 45'00" 18 48'54"


by us but have been reported for the state
(Brandao 1996). With the new recordings, the
number of poneromorph ant species reaches 18 for
Morelos. None is endemic to the state: 0. clarus is
distributed in the United States and Mexico;
L. mexicana and L. wheeleri have been recorded
only from Mexico; H. foeda is known for Mexico,
Central America, and some Caribbean Islands;
H. opaciceps, P stigma, P villosa, and E. tubercu-


latum are distributed from South America to the
United States; G striatula and G. regulars are
known from Mexico to Argentina; A. mayri, G.
strigata, G. sulcata, 0. laticeps, and E. ruidum are
distributed in Mexico, Central America and
northern South America; B. deletrix is distributed
from Mexico to Costa Rica. One species, Pl. punc-
tata, is known from the United States, Mexico,
Central America, and some Caribbean Islands.


TABLE 2. COLLECTION FREQUENCY (n = 139) OF SPECIES OF PONEROMORPH ANTS COLLECTED IN THE STATE OF MORE-
LOS, MEXICO WITH ALTITUDINAL RANGES AND VEGETATION TYPES WHERE THEY WERE FOUND (A: TROPICAL
DECIDUOUS FOREST; B: THORN FOREST; C: PASTURELAND; D: MANGO ORCHARDS; E: AVOCADO ORCHARDS; F:
ECOTONE BETWEEN TDF AND PINE-OAK FOREST; AND G: URBAN VEGETATION).

Species N Altitude (m) Vegetation

Anochetus mayri 2 945-990 a, c
Belonopelta deletrix 1 1229 a
Ectatomma ruidum 8 950-1510 a, c, d, g
Ectatomma tuberculatum 10 990-1480 a, d
Gnamptogenys regulars 1 1010 a, d
Gnamptogenys striatula 17 1010-1510 a, b, d
Gnamptogenys strigata 7 945-1540 a, g
Gnamptogenys sulcata 4 1010-1160 a
Hypoponera foeda -
Hypoponera punctatissima 1 1130 a
Hypoponera opaciceps 10 1010-1850 a, d, f
Leptogenys mexicana 4 990-1190 a, b
Leptogenys wheeler -
Odontomachus clarus 52 890-1890 a, c, f, b, g, d, e
Odontomachus laticeps 7 1054-1480 a, g, d
Pachycondyla stigma 8 950-1850 a, f
Pachycondyla villosa 1 950 a
Platythyrea punctata 6 950-1850 a, f


December 2007







Quiroz & Valenzuela: Poneromorph of Morelos, Mexico


Finally, H. punctatissima is a pantropical tramp
species, probably of African origin (Wilson & Tay-
lor 1967; Kempf 1972; Brandao 1991, 1996; Bol-
ton 1995; Lattke 1995, 2003; Longino 1998).
Of the 16 species collected during this study,
13 are new records from the state of Morelos:
A. mayri, B. deletrix, E. ruidum, E. tuberculatum,
G. strigata, G. sulcata, G. regulars, H. punctatis-
sima, L. mexicana, 0. laticeps, P stigma, P villosa
and Pl. punctata.
Poneromorphs are widely distributed through-
out Morelos. Only four of the 33 municipalities in
Morelos (Huitzilac, Tlalnepantla, Ocuituco, and
Tetela del Volcan) failed to produce samples of
these ants. Failure to yield samples could occur
because these municipalities are located in the
mountainous northern part of the state.
The most abundant and widely distributed
species were 0. clarus (37.4% of the total number
of samples obtained), G. striatula (12.2%), E. tu-
berculatum and H. opaciceps (7.1%), and E. rui-
dum and P stigma (6%). These 6 species repre-
sent 75.8% of the samples collected (Tables 2 and
3). Next in frequency were G. strigata and 0. lati-
ceps (5%), Pl. punctata (4.3%) G. sulcata and L.
mexicana (3%), and A. mayri (1.4%). The remain-
ing 4 species (B. deletrix, G. regulars, H. punc-
tatissima, and P villosa) were collected only once
each, suggesting that they are uncommon in Mo-
relos or that our sampling methods were ineffec-
tive in detecting them. Due to the methods used,
we could be underestimating the species abun-
dance of tree inhabitants, subterranean habits,
and ants that nest and forage in litter.


90
80so
70
60

% 40
30

to

890-1000 1001-1200 1201-1400 1401-1600 1601-1800 1801-2000
Altitude (m)

Fig. 2. Species richness (dark bars) and collection
frequency (light bars) of poneromorph ants at different
altitudes in the state of Morelos.


The greatest abundance and diversity were
found between 890 and 1600 m; 86.1% of samples
and all species were found in this altitudinal
range. Only 4 species (H. opaciceps, 0. clarus,
Pl. punctata, and P stigma) were obtained above
1600 m. The highest collection altitude recorded
(1,890 m) corresponds to 0. clarus, which was
found in Nepopualco, in the municipality of Toto-
lapan (Fig. 2 and Table 2).
Poneromorphs were encountered in a great di-
versity of vegetational associations (Table 2) in-
cluding tropical deciduous forest (TDF), ecotonal
areas between TDF and pine-oak forests, thorn
forests, pastureland, urban and suburban vegeta-
tion, and a variety of agroecosystems (including
mango, avocado, and guava orchards).


TABLE 3. DISTRIBUTION OF PONEROMORPH SPECIES FOUND IN THE STATE OF MORELOS. TABLE 1 SHOWS THE SITE THAT
CORRESPONDS TO EACH NUMBER.


Species


Locality


Anochetus mayri
Belonopelta deletrix
Ectatomma ruidum
Ectatomma tuberculatum
Gnamptogenys regulars
Gnamptogenys striatula
Gnamptogenys strigata
Gnamptogenys sulcata
Hypoponera foeda
Hypoponera punctatissima
Hypoponera opaciceps
Leptogenys mexicana
Leptogenys wheeler
Odontomachus clarus

Odontomachus laticeps
Pachycondyla stigma
Pachycondyla villosa
Platythyrea punctata


49,56
57
7, 13,32,44, 55,64
7, 21, 32, 43, 44, 48, 65, 68
7
5, 6, 7, 21, 32, 44, 45, 46, 47, 70, 71
12,29,33, 46, 56, 70
7, 8, 20

23
7, 10, 11, 15, 16, 22, 26, 74
49,64, 66

1, 3, 4, 6, 7, 10, 11, 12, 13, 14, 15, 16, 20, 21, 22, 25, 28, 29, 30, 32, 33, 37, 38, 39, 41,
43, 44, 45, 46, 47, 48, 54, 55, 56, 57, 58, 59, 61, 62, 64, 65,66,67,68,69, 70, 71, 72
7, 32,44,65, 70
6, 11,55
58
11,20, 46, 58







Florida Entomologist 90(4)


Most species were found in TDF, the most ex-
tensive vegetational association in Morelos and
one that is especially common in the central and
southern portions of the state. Other landscape
elements, including pastureland, cultivated fields,
and thorn forest (mainly Acacia spp.), are often
interspersed with the more abundant TDF. Pine-
oak forests are encountered in the mountainous,
northern part of the state, and a transitional eco-
tone is a common occurrence between TDF and
the pine-oak forest. This transitional zone ap-
pears to establish the distributional limit for pon-
eromorphs within the state of Morelos.
We found that most species are epigeal forag-
ers (except for G. regulars and P villosa). How-
ever, some can forage underground (L. mexicana,
G. strigata), in litter (G. strigata, G. sulcata, and
B. deletrix), and others on trees (E. tuberculatum,
G. regulars, P stigma, P villosa, and Pl. punc-
tata).
Most species nest in soil (except for A. mayri,
G. sulcata, H. punctatissima, P stigma, and P vil-
losa), in some cases preferentially under rocks
(0. clarus). Others nest in hollow branches
(A. mayri, B. deletrix, G. regulars, H. punctatis-
sima, H. opaciceps, G. strigata, and P stigma), in
litter (B. deletrix), in pre-existing cavities in dead
and living trees (G. sulcata, P villosa), and at the
base of epiphytic plants (P. villosa). Our observa-
tions regarding foraging and nesting sites coin-
cide with those of Lachaud (1990), Lattke (1995),
Longino (1998), and Durou et al. (2002). Data for
B. deletrix and L. wheeleri were taken from Long-
ino (1998), Wilson (1955), and Durou et al. (2002).
Of the 76 species of poneromorphs reported
from Mexico,18 (23.7%) are known for Morelos,
even though this state is relatively small (4,958
km2; 0.25% of the entire area of the country).
Poneromorphs are Pantropical, although in
America they have made some degree of penetra-
tion into the Nearctic region (Brown 1976). Of the
18 species of poneromorphs found in Morelos, 7
are recorded from the United States. The remain-
ing 11 species are distributed only in the Neotro-
pics. This composition of poneromorph ants may
result from the location of the state at the junc-
ture of these 2 biogeographic zones (Brown 1976;
Anonymous 1981; Aguilar 1990).

ACKNOWLEDGMENTS

The assistance of Patricia Galindo, Adriana Trejo
and Teresa Suarez is gratefully acknowledged, espe-
cially for their help in the collection and mounting of
ants. Thanks also to Ingrid Marquez for help with the
translation of this paper.

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Florida Entomologist 90(4)


December 2007


INFLUENCE OF TEMPERATURE ON SUSCEPTIBILITY OF
TRIBOLIUM CONFUSUM (COLEOPTERA: TENEBRIONIDAE) POPULATIONS
TO THREE MODIFIED DIATOMACEOUS EARTH FORMULATIONS


NICKOLAS G. KAVALLIERATOS1, CHRISTOS G. ATHANASSIOU2, BASILEIOS J. VAYIAS2 AND SEVASTI N. MAISTROU2
'Laboratory of Agricultural Entomology, Department of Entomology and Agricultural Zoology,
Benaki Phytopathological Institute, 8 Stefanou Delta str., Kifissia, Attica, Greece
E-mail: nick kaval@hotmail.com

'Laboratory of Agricultural Zoology and Entomology, Agricultural University of Athens,
75 lera Odos str., 11855, Athens, Greece

ABSTRACT

The influence of temperature on the insecticidal effect of 3 commercially available modified
diatomaceous earth (DE) formulations and their mixture against adults and larvae of Tri-
bolium confusum Jacquelin du Val originating from different European geographical loca-
tions was evaluated in laboratory tests. The temperatures tested were 20, 25, and 30 C. The
T confusum populations (strains) originated from Greece, Italy, Portugal, United Kingdom,
Germany, France, and Denmark. Mortality counts were carried out after 7 d of exposure of
T confusum individuals to DE-treated wheat. The 3 modified DEs tested were PyriSec, In-
secto, Protect-It, and a mixture of the three. The dose rates tested were 500 and 1000 ppm.
The strains from Portugal and France were the least susceptible to DEs, while the most sus-
ceptible one was the strain from the United Kingdom. The increase of temperature in many
of the combinations tested increased the efficacy of DE formulations against both adults and
larvae, but there were some cases in which temperature had no effect on DE efficacy. More-
over, adults were more tolerant than larvae to the DEs tested as well as to their mixture, re-
gardless of the strain or temperature. The present study clearly indicates that there are
serious variations in the efficacy of DEs against T confusum individuals originating from
different geographical regions, and several factors affect this efficacy, such as the type of for-
mulation and the temperature level prevailing.

Key Words: Modified diatomaceous earth, populations, temperature, Tribolium confusum,
wheat

RESUME

La influencia de la temperature sobre el efecto de los insecticides en 3 formulaciones comer-
ciales disponibles de suelo de diatomita (SD) modificada y su mezcla en contra de adults y
larvas de Tribolium confusum Jacquelin du Val provenientes de diferentes regions geogra-
ficas de europa fue evaluada en pruebas del laboratorio. Las temperatures probadas fueron
20, 25 y 30 C. Las poblaciones de razas de T confusum provenieron de Grecia, Italia, Portu-
gal, Reino Unido, Alemania, Francia y Dinamarca. Se realizaron los recuentos de mortalidad
despu6s de 7 d en que los individuos de T confusum fueron expuestos a trigo tratado con SD.
Los 3 SD probados fueron PyriSec, Insecto, Protect-It y una mezcla de los tres. Las dosis pro-
badas fueron 500 y 1000 ppm. Las razas de Portugal y de Francia fueron las menos suscep-
tibles a SD, mientras que la mas susceptible fue la raza del Reino Unido. El aumento en la
temperature en muchas de las combinaciones probadas aumento la eficacia de la formacio-
nes de SD contra los adults y larvas, pero hubo unos casos en que la temperature no tuvo
ningun efecto sobre la eficacia de SD. Ademas, los adults fueron mas tolerantes que las lar-
vas hacia los SD probados igual que la mezcla de los 3 SD, sin importar la raza o tempera-
tura. Este studio claramente indica que hay variaciones series en la eficacia de los SD
contra individuos de T confusum que provienen de diferentes regions geograficas, ademas
de various factors que afectan dicha eficacia, como la clase de formulaci6n y el nivel de tem-
peratura prevaleciente.


One of the promising alternatives to tradi- activate" the epicuticular lipids of the insects'
tional insecticides in stored-product protection is cuticle and insects die due to water loss and des-
the application of diatomaceous earths (DEs). DE iccation (Ebeling 1961, 1971; Korunic 1998; Sub-
is composed of fossilized remains of phytoplank- ramanyam & Roesli 2000). DEs have low mam-
tons diatomss) (Korunic 1998). DEs have a physi- malian toxicity and, as inert materials, can pro-
cal mode of insecticidal action, because they "in- vide long-term protection against stored product







Kavallieratos et al.: Diatomaceous Earths against Tribolium confusum


insects (Athanassiou et al. 2005a; Vayias et al.
2006c). Several studies document that the effi-
cacy of DE is affected by several factors such as
grain moisture and relative humidity, tempera-
ture, commodity, target species, and life stage
(Korunic 1998; Arthur 2000b; Subramanyam &
Roesli 2000; Fields & Korunic 2000; Mewis & Ul-
richs 2001; Vayias & Athanassiou 2004; Athanas-
siou et al. 2003, 2005b; Vayias et al. 2006c, Koru-
nic & Fields 2006). In addition, the insect strain is
another factor that can affect significantly the in-
secticidal efficacy of DE and its further assess-
ment as a grain protectant (Fields et al. 2003; Ar-
naud et al. 2005).
The confused flour beetle, Tribolium confusum
Jacquelin du Val, is one of the most serious stored
product insect pests worldwide (Aitken 1975). Al-
though this species develops best in processed
amylaceous commodities, both larvae and adults
can infest, feed, and develop in sound kernels (Ait-
ken 1975). This species has now developed resis-
tance to many chemical insecticides (Zettler 1991;
Arthur & Zettler 1992; Zettler & Arthur 1997).
Many studies have shown that several newer DE
formulations are effective against both T con-
fusum adults and larvae (Arthur 2000b; Vayias &
Athanassiou 2004; Athanassiou et al. 2004, 2005b;
Vayias et al. 2006c). According to these studies, at
the adult stage, T confusum is one of the most tol-
erant insect species to DE, and according to
Arthur (2000b) and Athanassiou et al. (2005b), it
can survive dose rates that can be lethal for other
species (Korunic 1998; Athanassiou et al. 2005b).
On the other hand, its larvae are very susceptible
to DEs (Vayias & Athanassiou 2004). In a recent
work, Vayias et al. (2006b) found considerable
variations in adult mortality when different popu-
lations of T confusum, originating from different
parts of Europe, were exposed to DE-treated
grain. Thus, even if similar experimental condi-
tions are followed, results obtained from studies
from different parts of the world may not be com-
parable, because different populations were used.
Vayias et al. (2006b) used only adults at 1 temper-
ature level; hence, additional experimentation is
needed to examine other life stages at a broader
range of temperatures. In the present study, 3
commercially available modified DE formulations
were evaluated against larvae and adults of T.
confusum populations originating from different
geographical locations of Europe. The impact of
temperature on the effect of DEs on these popula-
tions also was assessed.

MATERIALS AND METHODS

Formulations and Commodity

Three modified and enhanced DE formulations
were used in the tests: (a) PyriSec (Agrinova
Gmbh, Obrigheim/Miihleim, Germany) that con-


tains 1.2% natural pyrethrum (25%), 3.1% piper-
onyl butoxide and 95.7% of SilicoSec, which is a
DE of freshwater origin containing 92% SiO2. The
particle size distribution is between 8 and 12 pm
(Athanassiou et al. 2004), (b) Insecto (Insecto
Natural Products, Inc., Costa Mesa, CA, USA), a
DE of marine origin containing 86.7% SiO2 and
10% food-grade additives. The particle size distri-
bution is between 0.82 and 52.33 pm (Subraman-
yam et al. 1994), (c) Protect-It (Hedley Technolo-
gies Inc., Mississauga, Ontario, Canada) a DE
that contains 83.7% SiO2 with 10% silica aerogel.
The particle size distribution is between 5 and
6 pm (Korunic & Fields 1995), and (d) a mixture
of the 3 above-described DE formulations. The
mixture, which is not commercially available, was
prepared at our laboratory only for experimental
purposes by admixing equal quantities of the
tested DE formulations. The commodity tested
was untreated, clean, hard wheat (var Mexa) har-
vested in 2004. The dockage of the wheat was
minimal (<0.6%) and the moisture content as de-
termined by a Dickey-John moisture meter
(Dickey-John Multigrain CAC II, Dickey-John Co.,
Lawrence, KS, USA) was approx. 11.4%.

Insects and Rearing Cultures

Tribolium confusum adults less than 14 d old
and 3-4th instars were used in the tests. There
were 7 different populations obtained from Dan-
ish Pest Infestation Laboratory (Lyngby, Den-
mark), Central Science Laboratory (York, UK),
Benaki Phytopathological Institute (Kifissia,
Greece), Institute for Stored Product Protection
(Berlin, Germany), University of Molise (Cam-
pobasso, Italy), Laboratoire Denrees StockBes
(Cenon Bordeaux, France), and Tropical Scientific
Research Institute (Lisbon, Portugal) (the above
strains are abbreviated in the text as DK, UK,
GR, GER, IT, FR and POR, respectively). All
strains had been initially collected from the local
fauna of each country. Both adults and larvae of
the populations were reared on wheat flour plus
5% brewers yeast (by weight) at 27 1VC and 65
5% RH. The GR strain was kept at the Benaki
Phytopathological Institute for >10 years, while
the rest were reared in the same laboratory for >7
generations, after introduction from their respec-
tive countries of origin.

Bioassays

Exposure studies were carried out at 20, 25,
and 30'C and 70 1.5% relative humidity (RH).
Six 1-Kg wheat lots were prepared and placed in
6 cylindrical glass jars (18 cm in diameter, 30 cm
in height). Three of these jars contained wheat
treated with 500 ppm of each DE (one jar per DE),
whereas each of the other 3 jars contained wheat
treated with 1000 ppm of DE. Previous studies







Florida Entomologist 90(4)


have shown that T confusum adults are not sus-
ceptible at doses lower than 500 ppm of some of
these DEs (Arthur 2000a, b; Athanassiou et al.
2005b; Vayias & Athanassiou 2004). In addition, 2
cylindrical jars contained wheat treated with an
equivalent mixture of all of the DEs at the same
dose rates. An additional jar of untreated wheat
was used as control. All jars were shaken manu-
ally for approx. 5 min to achieve distribution of
the dust in the entire grain mass. For each
strain's life stage, 4 samples of 30 g each were
taken from each jar, and each sample was placed
in a cylindrical glass vial (7 cm in diameter, 12 cm
in height). The vials were closed, except for a hole
(3 cm in diameter) in the cap, which was covered
with organdy to allow sufficient aeration. Then,
30 T confusum adults or larvae from each strain
were separately introduced into each vial, and all
vials were placed in incubators set each time at
the aforementioned conditions. Mortality of the
exposed individuals (adults or larvae) was mea-
sured after 7 d of exposure. The entire procedure
was repeated 3 times, by preparing new lots of
wheat each time. During the experimental period
the RH level was maintained with saturated
potassium iodine solution, as recommended by
Greenspan (1977).

Data Analysis and Statistics

Preliminary analysis for both adults and lar-
vae showed homogeneity among the replicates of
each assay according to the Levene test (Levene
1960). The data were initially corrected by Ab-
bott's (1925) formula. The arcsine transformed
data were separately analyzed for adults and lar-
vae. For a given life stage, mortality counts were
analyzed by the GLM Procedure of SAS (1998)
with adult (or larval) mortality as the response
variable and DE formulation, dose rate, strain,
and temperature as main effects. Control mortal-
ity ranged between 2-8% and 3-12% for adults
and larvae, respectively. Means were separated
by Tukey-Kramer (HSD) at P = 0.05 (Sokal &
Rohlf 1995).

RESULTS

Tribolium confusum Adults

All main effects and associated interactions
were significant at P < 0.001 level (formulation:
F = 174.83, df = 3, 2015, dose rate: F = 15313.22,
df = 1, 2015, strain: F = 134.07, df = 6, 2015, tem-
perature: F = 1142.84, df= 2, 2015, formulation x
dose rate: F = 24.75, df = 3, 2015, formulation x
strain: F = 2.15, df = 18, 2015, formulation x tem-
perature: F = 4.54, df = 6, 2015, dose rate x strain:
F = 9.44, df = 6, 2015, dose rate x temperature:
F = 39.20, df = 2, 2015, strain x temperature: F =
2.22, df = 12, 2015).


Significant differences in the mortality of
adults were noted among the T confusum strains
as well as among the temperatures tested. At
20C, mortality of adults at the dose rate of 500
ppm, was less than 20% for all the DEs tested
whereas the respective figures at 1000 ppm
ranged between 37.5% and 83.8%. At this temper-
ature, in most cases, the POR and GR strains
were significantly less susceptible than the other
strains at 500 ppm, while at 1000 ppm the least
susceptible strains were the POR and FR, with
the exception of Protect-It where the least suscep-
tible strains were POR and IT (Tables 1, 2).
The increase of temperature from 20C to 25C
led to a significant increase in mortality of adults
in most of the cases tested. However, in the case of
Insecto, significant differences between 20 and
25C were noted only with the GR and GER
strains. At 25C, as noted for 20C, the most toler-
ant strains were the POR and FR. Although the
POR was more tolerant than the FR at 25C, sig-
nificant differences in adult mortality between
those 2 strains were not noted at either dose lev-
els of Pyrisec and at 1000 ppm of Protect-it. At
500 ppm DEs, significant differences in adult
mortality were not observed between the IT and
GR strain, except for the case of mixture of DEs
(Tables 1, 2).
At 30C, mortality values of T confusum adults
were higher than 90%, regardless of the strain or
formulation. This was noted at 1000 ppm, while
at 500 ppm, adult mortality did not exceed 60%.
At this temperature, strains POR and FR were
the most tolerant strains, whereas the most sus-
ceptible one was the UK strain. Generally, of the
formulations tested, Protect-it and the mixture of
DEs were the most effective, and the least effec-
tive one was Insecto (Tables 1, 2).

Tribolium confusum Larvae

All main effects and associated interactions
were significant at P < 0.001 (formulation: F =
213.85, df = 3, 2015, dose rate:F = 2191.16, df = 1,
2015, strain: F = 157.64, df = 6, 2015, tempera-
ture: F = 747.59, df = 2, 2015, formulation x dose
rate: F = 4.64, df = 3, 2015, formulation x strain:
F = 2.62, df = 18, 2015, formulation x tempera-
ture: F = 29.66, df = 6, 2015, dose rate x strain: F
= 28.95, df = 6, 2015, dose rate x temperature: F =
6,77, df = 2, 2015, strain x temperature: F = 4,
515, df = 12, 2015).
As noted for adults, larval mortality increased
with temperature irrespective of the T confusum
strain, DE formulation, or dose. In most cases, at
temperatures higher than 25C and at the high-
est dose rate tested, larval mortality was 100%.
Regardless of temperature, DE formulation, and
dose rate, significantly more larvae of the UK
strain were dead in comparison with the other 6
strains. On the other hand, the most tolerant


December 2007







Kavallieratos et al.: Diatomaceous Earths against Tribolium confusum


TABLE 1. MEAN MORTALITY ( SE) OF DIFFERENT STRAINS OF T. CONFUSUM ADULTS EXPOSED TO WHEAT TREATED
WITH 500 PPM OF 3 DE FORMULATIONS AND THEIR MIXTURE, AT 3 TEMPERATURE LEVELS.

Temperature (C)

Formulation Strain 20 25 30

Pyrisec GR 7.5 2.6 Ab 20.5 1.3 Bb 38.5 2.2 Cc
IT 20.1 + 3.7 Af 21.1 + 1.6 Ab 27.0 2.7 Bb
DK 13.5 3.4 Ad 28.9 2.6 Bc 40.5 2.8 Cc
POR 2.5 4.5 Aa 12.8 1.5 Ba 15.0 2.6 Ba
UK 18.5 5.4 Aef 35.7 4.8 Bd 54.2 4.2 Cd
FR 10.8 + 1.4 Ac 13.2 1.4 ABa 15.4 2.6 Ba
GER 16.2 3.9 Ae 27.8 2.9 Bc 37.8 4.2 Cc
Insecto GR 5.0 1.5 Ab 12.8 1.5 Bb 15.4 1.3 Bb
IT 10.4+ 2.1 Ac 10.5 3.1Ab 13.5+ 1.1 Bb
DK 2.7 5.4 Aa 5.3 1.7 Aa 37.8 2.7 Bd
POR 10.0 + 5.4 Ac 7.7 1.1Aa 10.0 2.1Aa
UK 11.1 + 2.8 Ac 25.0 + 3.6 Ac 41.7 2.5 Ae
FR 10.8 + 2.9 Ac 7.9 1.7 Aa 10.3 1.3 Aa
GER 1.5 1.2 Aa 22.2 3.3 Bc 32.4 2.7 Cc
Protect-It GR 2.5 1.3 Aa 23.1 1.3 Bb 41.0 1.6 Cc
IT 5.0 1.5 Aa 21.1 1.5 Bb 29.7 2.5 Cb
DK 13.5 2.3 Ac 34.2 2.3 Bd 45.9 1.6 Cd
POR 2.5 1.3 Aa 17.9 1.3 Ba 25.0 3.4 Ca
UK 14.8 + 4.6 Ac 39.3 4.6 Be 54.2 1.5 Ce
FR 10.8 2.7 Ab 21.1 2.7 Bb 25.6 + 4.1 Ca
GER 13.5 2.3 Ac 27.8 2.3 Bc 40.5 7.4 Cc
Mixture GR 5.2 1.3 Aa 25.6 1.2 Bc 43.6 1.6 Cc
IT 20.0 + 1.5 Ac 21.1 + 2.9 Ab 32.4 2.5 Bb
DK 5.4 2.3 Aa 31.6 3.5 Bd 48.6 1.5 Cd
POR 2.5 1.3 Aa 17.9 1.3 Ba 25.0 3.4 Ca
UK 18.5 + 4.6 Ac 35.7 5.9 Be 58.3 1.5 Be
FR 13.5 2.7 Ab 18.4 1.6 Ba 30.8 + 4.1 Cb
GER 18.9 2.3 Ac 30.6 2.3 Bd 45.9 7.8 Ccd

For a given formulation, means in the same column followed by the same lowercase letter are not significantly different, while
means in the same row followed by the same uppercase letter are not significantly different; lowercase letters for strains; uppercase
letters for temperature; temperature df = 2, 36, strain df = 6, 71, Tukey-Kramer test at 5%; for abbreviations of strains see materials
and methods.


strains, in terms of larval mortality, were the POR
and FR, with the former strain being more sus-
ceptible than the latter. The GER strain was more
tolerant than the GR, IT, and DK strains. Signifi-
cantly more larvae of the DK strain were dead in
comparison with the GER, GR, and IT strains at
all formulations and temperatures tested. Signif-
icant differences in larval mortality between the
GR and IT strain were not noted, except for the
case of 500 ppm of Protect-It, in which larvae of
the GR strain were more susceptible than those of
the IT strain (Tables 3, 4).

DISCUSSION

The results of the present study indicate that
DEs could be used with success against T con-
fusum, but their insecticidal efficacy is highly in-
fluenced by several factors such as temperature,
life stage, type of DE formulation, and dose rate.


These findings seem to support those obtained
from recent studies (Rigaux et al. 2001; Arnaud et
al. 2005; Vayias et al. 2006b) indicating that the
efficacy of DEs against a given species depends on
the geographical location of the species origin. For
instance, Arnaud et al. (2005) found significant
differences in susceptibility to DE formulations
among adults of the red flour beetle, Tribolium
castaneum (Herbst) originating from different
geographical locations of the world. In a recent
study, Vayias et al. (2006b) in tests of several DEs
against the same strains of T confusum found sig-
nificant variations, which were not always consis-
tent for all formulations. However, in that study,
the strains were tested only at the adult stage,
and only at 1 temperature level. In light of our
findings, it is clearly evident that the observed
difference in susceptibility among the strains
tested is determined by both formulation and
temperature, indicating the complexity of this








Florida Entomologist 90(4)


TABLE 2. MEAN MORTALITY ( SE) OF DIFFERENT STRAINS OF T. CONFUSUM ADULTS EXPOSED TO WHEAT TREATED
WITH 1000 PPM OF 3 DE FORMULATIONS AND THEIR MIXTURE AT 3 TEMPERATURE LEVELS.

Temperature ( C)

Formulation Strain 20 25 30

Pyrisec GR 65.0 4.5 Ad 76.9 1.3 Bb 92.3 1.5 Ca
IT 55.0 + 1.5 Ac 65.8 4.2 Ba 94.6 2.2 Ca
DK 75.7 3.5 Ae 81.6 1.6 Bc 100 0.0 Cb
POR 42.5 2.5 Aa 66.7 2.7 Ba 92.5 2.3 Ca
UK 74.1+ 4.1 Ae 89.3 4.3 Bd 100 0.0 Cb
FR 48.6 1.6 Ab 65.8 2.2 Ba 94.9 1.4 Ca
GER 73.0 4.4 Ae 80.6 + 2.1 Bc 100 0.0 Cb
Insecto GR 47.5 + 5.6 Ac 69.2 + 3.1 Bc 100 0.0 Ca
IT 45.0 3.4 Abc 60.4 1.4 Bb 100 0.0 Ca
DK 70.3 2.8 Ae 84.2 2.7 Bd 100 0.0 Ca
POR 37.5 1.3 Aa 56.4 3.9 Ba 100 0.0 Ca
UK 74.1 + 1.9 Af 85.7 1.3 Bd 100 0.0 Ca
FR 43.2 2.7 Ab 60.5 4.1 Bb 100 0.0 Ca
GER 59.5 + 1.4 Ad 83.3 2.9 Bd 100 0.0 Ca
Protect-It GR 77.5 3.9 Ac 87.2 2.6 Bb 100 0.0 Ca
IT 72.5 2.5 Ab 81.6 1.4 Ba 100 0.0 Ca
DK 81.1 + 4.2 Ad 92.1 + 1.3 Bc 100 0.0 Ca
POR 65.0 1.5 Aa 79.5 3.1 Ba 100 0.0 Ca
UK 85.2 2.3 Ae 96.4 1.9 Bd 100 0.0 Ca
FR 73.0 3.6 Ab 78.9 2.2 Ba 100 0.0 Ca
GER 83.8 3.8 Ade 91.7 + 1.5 Bc 100 0.0 Ca
Mixture GR 75.0 1.5 Ac 89.7 1.2 Bc 97.4 1.3 Ca
IT 70.0 3.7 Ab 84.1 1.6 Bb 100 0.0 Ca
DK 78.4 1.3 Ad 94.7 1.6 Bd 100 0.0 Ca
POR 62.5 2.5 Aa 76.9 2.6 Ba 100 0.0 Ca
UK 81.5 1.9 Ae 100 0.0 Be 100 0.0 Ba
FR 64.9 4.2 Aa 83.9 1.5 Bb 100 0.0 Ca
GER 73.0 + 1.6 Ac 86.1 4.4 Bb 100 0.0 Ca

For a given formulation, means in the same column followed by the same lowercase letter are not significantly different, while
means in the same row followed by the same uppercase letter are not significantly different; lowercase letters for strains; uppercase
letters for temperature; temperature df = 2, 36, strain df = 6, 71, Tukey-Kramer test at 5%; for abbreviations of strains see Mate-
rials and Methods.


phenomenon. For instance, at 500 ppm, Insecto
was equally effective among temperatures
against 3 of the T confusum strains at the adult
stage. Rigaux et al. (2001) examining the suscep-
tibility of different T castaneum strains to Pro-
tect-It, found that mortality was directly related
to insect mobility, and that the less mobile strains
were the least susceptible. It is generally accepted
that low mobility decreases the contact with DE
particles (Korunic 1998; Subramanyam & Roesli
2000). Apart from mobility, potential variations in
the composition of the epicuticular lipids and cu-
ticle thickness may be responsible for these vari-
ations. Since the use of DEs is currently limited,
probably the strains examined have never been in
contact with DEs. Consequently, the development
of resistance after previous exposure to DEs is
unlikely. Since many DE formulations are now in
the process of registration in several parts of
the world, these variations should be taken into


account. Fields et al. (2003), reporting the results
from an international working group with several
DEs, found considerable variations among strains
originating from different geographical regions.
Although the use of standardized strains is prob-
ably a solution to this variation, especially among
working groups conducting tests with DEs, the
use of a local strain is practically more preferable
since this is adapted to local conditions.
The effect of life stage on the insecticidal effi-
cacy of DE against a given insect species has been
examined in previous studies (Subramanyam &
Roesli 2000; Mewis & Ulrichs 2001; Vayias &
Athanassiou 2004). Vayias & Athanassiou(2004)
found that larvae of T confusum were much more
susceptible than adults when exposed to Silico-
Sec-treated wheat. Our findings support this re-
port not only for Silicosec, but also for the other
DEs tested. As above, these differences among
adults and larvae of a given species could be


December 2007







Kavallieratos et al.: Diatomaceous Earths against Tribolium confusum


TABLE 3. MEAN MORTALITY ( SE) OF DIFFERENT STRAINS OF T. CONFUSUM LARVAE EXPOSED TO WHEAT TREATED
WITH 500 PPM OF 3 DE FORMULATIONS AND THEIR MIXTURE AT 3 TEMPERATURE LEVELS.

Temperature ( C)

Formulation Strain 20 25 30

Pyrisec GR 70.0 + 2.1 Abc 82.5 3.3 Bb 87.5 2.5 Cb
IT 2.5 2.5 Ac 80.0 2.1 Bb 85.0 2.6 Cb
DK 70.4 + 2.1 Abc 97.3 1.3 Bc 97.5 1.3 Bc
POR 67.5 1.3 Ab 80.1 2.1 Bb 85.1 1.5 Cb
UK 77.5 + 3.9 Ad 97.5 1.3 Bc 100 0.0 Bc
FR 67.6 2.5 Ab 70.0 2.1 ABa 72.5 2.5 Ba
GER 62.5 2.5 Aa 72.5 2.5 Ba 87.3 2.4 Cb
Insecto GR 55.0 1.5 Abc 75.0 1.5 Bc 80.0 3.7 Cc
IT 52.5 1.3 Ab 77.5 1.3 Bc 85.0 1.5 Cd
DK 65.1 + 1.5 Ad 82.5 1.4 Bd 87.5 1.3 Cd
POR 55.4 + 2.6 Abc 70.0 2.1 Bb 75.0 1.5 Cb
UK 75.0 3.4 Ae 85.0 1.5 Bd 87.6 2.6 Bd
FR 47.5 2.5 Aa 57.5 3.9 Ba 62.5 3.3 Ca
GER 57.5 + 4.5 Ac 67.5 1.3 Bb 77.5 1.3 Cbc
Protect-It GR 70.0 3.7 Ac 82.6 4.5 Bc 90.0 3.7 Cd
IT 65.0 1.5 Ab 80.0 3.7 Bc 87.5 2.5 Ccd
DK 77.5 2.5 Ad 90.0 3.6 Bd 95.0 2.6 Ce
POR 60.0 + 3.1 Aa 75.0 4.6 Bb 82.5 1.3 Cb
UK 82.5 1.3 Ae 100 0.0 Be 97.5 1.3 Be
FR 57.5 3.9 Aa 65.0 4.5 Ba 72.5 2.5 Ca
GER 67.5 + 2.5 Abc 82.5 2.5 Bc 85.0 1.5 Bbc
Mixture GR 75.0 3.4 Abc 87.5 2.5 Bcd 87.3 3.3 Bb
IT 72.5 4.5 Ab 87.8 2.9 Bcd 87.5 4.5 Bb
DK 85.0 + 2.6 Ad 90.0 1.2 Bd 92.5 1.9 Bc
POR 70.0 4.5 Aab 85.0 1.5 Bbc 87.1 1.6 Bb
UK 85.6 + 1.5 Ad 97.5 1.3 Be 97.5 1.5 Bd
FR 67.5 3.4 Aa 67.5 4.9 Aa 72.5 2.5 Ba
GER 77.5 + 2.5 Ac 82.5 2.5 Bb 85.0 4.5 Bb

For a given formulation, means in the same column followed by the same lowercase letter are not significantly different, while
means in the same row followed by the same uppercase letter are not significantly different; lowercase letters for strains; uppercase
letters for temperature; temperature df = 2, 36, strain df = 6, 71, Tukey-Kramer test at 5%; for abbreviations of strains see Mate-
rials and Methods.


attributed to several factors, such as differences
in morphological traits, epicuticular composition,
cuticle thickness, and agility. One possible expla-
nation is that adults or larvae of the same devel-
opmental stage may have a different thickness of
cuticle or even different composition of epicuticu-
lar lipids. Interestingly, for many of the combina-
tions tested here, the rank of susceptibility is sim-
ilar for adults and larvae. This may suggest that
behavioral characteristics may be among the
dominant reasons for these variations. Further
experimental work is needed to examine the basis
of this hypothesis.
The effect of temperature on the insecticidal
effect of DEs against stored grain insects has been
investigated by several researchers (Aldryhim
1990; Arthur 2000b; Dowdy & Fields 2000; Vayias
& Athanassiou 2004; Athanassiou et al. 2005b).
For the majority of the stored-product beetle spe-
cies examined increase in temperature increases


the DE efficacy (Fields & Korunic 2000; Subra-
manyam & Roesli 2000; Athanassiou et al.
2005b). Increase of temperature enhances inter-
nal water loss through the insects' body surface,
and thus, it is positively related to increased des-
iccation caused by DE particles (Fields & Korunic
2000). Moreover, at high temperatures insects are
more mobile, resulting in increased contact with
the DE particles. However, in the case of T con-
fusum, the effect of temperature is often contra-
dictory. Aldryhim (1990) found that T confusum
adults were more susceptible at 20C than at
30C on wheat treated with the DE Dryacide. In
contrast, Vayias & Athanassiou (2004) found that
both adults and larvae of T confusum strain
noted here as GR were more susceptible to Silico-
Sec-treated wheat and flour at elevated tempera-
tures. Similar results also were reported by
Arthur (2000b) for Protect-It, for both adults of
T confusum and T castaneum. Nevertheless, by







Florida Entomologist 90(4)


TABLE. 4. MEAN MORTALITY ( SE) OF DIFFERENT STRAINS OF T. CONFUSUM LARVAE EXPOSED TO WHEAT TREATED
WITH 1000 PPM OF 3 DE FORMULATIONS AND THEIR MIXTURE AT 3 TEMPERATURE LEVELS.

Temperature ( C)

Formulation Strain 20 25 30

Pyrisec GR 87.5 3.3 Ad 100 0.0 Ba 100 0.0 Ba
IT 85.0 1.5 Acd 100 0.0 Ba 100 0.0 Ba
DK 92.4 1.3 Ae 100 0.0 Ba 100 0.0 Ba
POR 80.0 2.3 Ab 100 0.0 Ba 100 0.0 Ba
UK 95.0 1.5 Ae 100 0.0 Ba 100 0.0 Ba
FR 75.0 2.6 Aa 100 0.0 Ba 100 0.0 Ba
GER 82.5 2.5 Abc 100 0.0 Ba 100 0.0 Ba
Insecto GR 70.0 3.1 Ac 92.5 1.3 Bd 100 0.0 Ca
IT 72.8 2.5 Ac 95.0 1.5 Bde 100 0.0 Ca
DK 80.0 + 0.6 Ad 97.5 1.3 Bef 100 0.0 Ba
POR 67.5 1.3 Ab 80.0 0.9 Bb 100 0.0 Ca
UK 85.0 1.6 Ae 100 0.0 Bf 100 0.0 Ba
FR 62.5 1.3 Aa 72.5 2.5 Ba 100 0.0 Ca
GER 72.5 + 1.4 Ac 87.5 1.3 Bc 100 0.0 Ca
Protect-It GR 97.4 + 1.4 Ad 100 0.0 Aa 100 0.0 Aa
IT 97.3 + 1.3 Ad 100 0.0 Aa 100 0.0 Aa
DK 97.5 + 1.4 Ad 100 0.0 Aa 100 0.0 Aa
POR 85.0 3.4 Ab 100 0.0 Ba 100 0.0 Ba
UK 100 + 0.0 Ad 100 0.0 Aa 100 0.0 Aa
FR 72.5 1.3 Aa 100 0.0 Ba 100 0.0 Ba
GER 92.5 1.4 Ac 100 0.0 Ba 100 0.0 Ba
Mixture GR 100 + 0.0 Ac 100 0.0 Aa 100 0.0 Aa
IT 97.5 + 1.3 Ac 100 0.0 Aa 100 0.0 Aa
DK 100 + 0.0 Ac 100 0.0 Aa 100 0.0 Aa
POR 90.0 3.4 Ab 100 0.0 Ba 100 0.0 Ba
UK 100 + 0.0 Ac 100 0.0 Aa 100 0.0 Aa
FR 75.0 3.5 Aa 100 0.0 Ba 100 0.0 Ba
GER 92.5 1.3 Ab 100 0.0 Ba 100 0.0 Ba

For a given formulation, means in the same column followed by the same lowercase letter are not significantly different, while
means in the same row followed by the same uppercase letter are not significantly different; lowercase letters for strains; uppercase
letters for temperature; temperature df = 2, 36, strain df = 6, 71, Tukey-Kramer test at 5%; for abbreviations of strains see Mate-
rials and Methods.


testing the same strain, Athanassiou et al.
(2005b) found that the influence of temperature is
affected by the exposure interval. Hence, in that
study, T confusum adults were more susceptible
to SilicoSec, at 30C than at 32C at exposures
<48 h, while the reverse occurred at exposures
>7 d. The age/instar of the adults/larvae tested
may be among the crucial factors for these dissim-
ilar results, and this is why previous studies sug-
gest the use of standardized-age individuals of
Tribolium spp. (De Paula et al. 2002; Vayias &
Athanassiou 2004), while for other species, age is
not very important (Athanassiou et al. 2006a). In
an extensive study with various DEs and stored-
product insect species, Fields and Korunic (2000)
found negative or positive correlations with tem-
perature. Although the authors did not test T con-
fusum, their data indicate that from, the species
tested, T castaneum were slightly less susceptible
to DEs at elevated temperatures. Increased tem-


perature is expected to increase feeding and as a
result, metabolic water. Furthermore, the synthe-
sis of cuticular waxes may be faster at higher tem-
perature levels through temperature-mediated
biochemical pathways (Fields & Korunic 2000).
Since, as inert materials, DEs are not affected by
temperature, it seems that other, physiological
factors are responsible for the variations recorded
in the present work. Also, as noted above, the
source of DEs seems to have a certain effect on in-
sect mortality, although Korunic (1997, 1998) re-
ported that there are more important factors than
the diatom species that compose each DE; these
factors are chiefly tapped density, SiO, content, oil
absorbency, particle size, and secondarily pH (Ko-
runic 1997). Nevertheless, the additives that each
DE contains may differentiate the DE efficacy.
Hence, the observed difference in relation to tem-
perature for Insecto could be attributed to the
presence of food additives, which may attract in-


December 2007







Kavallieratos et al.: Diatomaceous Earths against Tribolium confusum


sects regardless of the temperature level; food at-
tractants may lead to DE consumption, causing
internal desiccation. The present findings par-
tially support the aforementioned reports, since,
in the combinations tested here, the strains ex-
amined had either positive relation or no relation
to temperature. In contrast, none of the combina-
tions showed a negative correlation with temper-
ature. Consequently, our findings support the ad-
mission that, like most stored-product insect spe-
cies, T confusum is generally more vulnerable to
DEs at high temperatures.
Vayias et al. (2006a) reported the first results
for the effect of the combination of natural pyre-
thrum with DE against T confusum, but there is
still inadequate information on the effect of this
combination against stored-grain pest species.
Generally, the efficacy of most pyrethroids is neg-
atively related to temperature (Snelson 1987;
Arthur 1996). In the present tests, PyriSec was
more effective at high temperature levels which
may suggest that the DE particles moderated the
effect of temperature on natural pyrethrum. Vay-
ias et al. (2006a) found that natural pyrethrum
was equally effective against T confusum pupae
at 25 and 30C.
The type of the DE formulation is also an im-
portant factor affecting insecticidal effect (Koru-
nic 1998; Fields & Korunic, 2000; Subramanyam
and Roesli, 2000). Our findings are in compliance
with this claim since efficacy of the tested DEs
varied for a given T confusum adult or larvae
strain. As noted above, this fact could be attrib-
uted to the different SiO2 content of the tested
DEs as well as to the different substances that
have been added to the DE formulations in order
to enhance their efficacy. Due to these variations,
Arnaud et al. (2005) proposed as a possible solu-
tion the use of a DE mixture, in which more than
one DE is present. In the same study, the mixture
of the tested DEs was more effective than the ap-
plication of each DE alone. Generally, a mixture
could combine all the positive characteristics of
different DEs, such as the use of low insecticidal
rates and the presence of food additives. Athanas-
siou et al. (2007) found that a mixture of Protect-
It, PyriSec and Insecto were more effective than a
single DE application on both wheat and maize,
against adults of T confusum, the rice weevil, Si-
tophilus oryzae (L.) and the lesser grain borer,
Rhyzopertha dominica (F.).
The increase of dose from 500 to 1000 ppm
moderated or eliminated the differences among
DEs or strains. This is due to the fact that, at 1000
ppm, the mortality level was 100% or close to
100%, which may have concealed any differences.
However, 1000 ppm is a dose rate that is consid-
ered too high, and affects negatively to a great de-
gree the physical properties of the grains, partic-
ularly bulk density (Korunic et al. 1996; Subra-
manyam & Roesli 2000; Athanassiou et al.


2006b). According to Fields and Korunic (2000),
this is the main problem in presenting efficacy
data with DEs: mortality ranges between 0% and
100%, so any increase of dose results in 100%
mortality, covering possible interaction with
other factors that may affect mortality. As a sec-
ondary pest, which cannot infest easily sound
grain seeds directly, the damage on kernels by
T confusum is more gradual in comparison with
other pests, such as the internal feeders S. oryzae
(rice weevil) and R. dominica (lesser grain borer)
(Aitken 1975). Hence, it would take several gener-
ations for this species to cause considerable grain
damage. Although T confusum adults are toler-
ant to DEs, larvae are highly susceptible. Conse-
quently, the residual applications of DEs may
gradually eliminate T confusum populations
through the high immature mortality. As inert
materials, DEs can persist for a long period in
grain without loss in insecticidal efficacy (Atha-
nassiou et al. 2005a; Vayias et al. 2006c).
Our focus was to evaluate the efficacy of DEs
under a wide range of conditions, as well as the
potential interactions of these conditions. DEs are
now in the first steps of acceptance and registra-
tion as grain protectants in many parts of the de-
veloped world. The use of DEs in stored-product
protection meets with several drawbacks as re-
gards some properties of the treated grain such as
reduced bulk density, reduced flowability, and
dusty appearance (Korunic 1997, 1998; Subrman-
yam & Roesli 2000). These drawbacks have led re-
searchers to assess the efficacy of DEs at lower
dose rates by evaluating the combined use of DEs
with other methods, such as heat, botanicals or
entomopathogenic fungi (Dowdy & Fields 2000;
Athanassiou et al. 2006b; Kavallieratos et al.
2006; Michalaki et al. 2006; Vassilakos et al.
2006). This new DE-based approach, which needs
also additional investigation, is likely to provide
the inferences necessary for a more extensive use
of DEs as grain protectants in future.

ACKNOWLEDGMENTS

We thank Cornel Adler, Patrick Ducom, Lise
Stengard Hansen, Anna Paula Pereira, Pasquale Trem-
aterra, and Maureen Wakefield for providing the T con-
fusum strains. This study was partially supported by
the Ministry of Rural Development and Food (Director-
ate of Plant Produce Protection).

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December 2007


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1157-1162.







Florida Entomologist 90(4)


December 2007


AGANASPIS ALUJAI (HYMENOPTERA: FIGITIDAE: EUCOILINAE),
A NEW SPECIES ATTACKING RHAGOLETIS (DIPTERA: TEPHRITIDAE)
IN THE NEOTROPICAL REGION

SERGIO M. OVRUSKI1' ROBERT A. WHARTON2, JUAN RULL3 AND LARISSA GUILLEN3
'PROIMI-Biotecnologia, Divisi6n Control Biol6gico de Plagas, CONICET (T4100MVB),
San Miguel de Tucuman, Tucuman, Argentina

2Department of Entomology, Texas A&M University, College Station, TX 77843-2475, USA

3Instituto de Ecologia, A.C., Unidad de Entomologia Aplicada, Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico

ABSTRACT

A new neotropical species of Aganaspis (Hymenoptera: Figitidae, Eucoilinae) is described
and compared to other Aganaspis species occurring in the Neotropical Region. This thely-
tokous species was reared from pupae of 2 species of fruit flies in the genus Rhagoletis
(Diptera: Tephritidae), both in the suavis species group, and collected from 2 species of na-
tive walnuts in Mexico. Evidence of host specialization through diapause timing and notes
on basic biology are provided.

Key Words: Hymenoptera, Figitidae, Aganaspis, Rhagoletis, Parasitoid

RESUME

Una nueva especie neotropical de Aganaspis (Hymenoptera: Figitidae, Eucoilinae) se des-
cribe y compare con otras species deAganaspis de la region. Esta especie thelyotokica fue
recuperada de pupas de dos species de moscas de la fruta en el g6nero Rhagoletis (Diptera:
Tephritidae), ambas pertenecientes al grupo de species suauis y obtenidas de dos species
nativas de nueces en M6xico. Se provee evidencia de especializaci6n a su hospedero mani-
festada por la regulaci6n del tiempo de la diapausa y algunas notas sobre su biologia basica.


Translation provided by the authors.


The genus Aganaspis Lin (Hymenoptera:
Figitidae, Eucoilinae) comprises only 6 species,
which are distributed in the Neotropical and
Indo-Pacific regions. The American species A. pel-
leranoi (Brethes) andA. nordlanderi Wharton can
be easily differentiated from the Asian species
group (A. daci (Weld),A. contract Lin,A. ocellata
Lin, and A. major Lin) by several morphological
features such as the shape of the scutellar disc
and female antenna, absence of setae on the eyes,
and lack of the median depression in the meta-
pleuron (Wharton et al. 1998). Originally, this ge-
nus was described by Lin (1987) to accommodate
Asian species. Later, following suggestions by G6-
ran Nordlander, Ovruski (1994) placed the neo-
tropical species Ganaspis pelleranoi (Brethes) (=
G. carualhoi Dettmer) in Aganaspis. This place-
ment has been accepted by subsequent authors
(Wharton et al. 1998; Diaz & Gallardo 2000; Fon-
tal-Cazalla et al. 2002; Guimaraes et al. 2003). In
a cladistic analysis of the subfamily Eucoiline
(Fontal-Cazalla et al. 2002), the genus Aganaspis
was included in the "Neotropical grade", an unre-
solved group of Neotropical taxa representing a
morphological transition between the Zaeucoila


group of genera and the 5 genus groups of higher
eucoilines (The Ganaspis Foerster, Chrestosema
Foerster, Trybliographa Foerster, Rhoptromeris
Foerster, and Kleidotoma Westwood groups) rec-
ognized by Nordlander (1982).
Aganaspis daci (originally described as Tryblio-
grapha daci Weld), was first collected in Malaysia
and Borneo, and introduced into Hawaii as a po-
tential biocontrol agent for Bactrocera dorsalis
(Hendel) (Clausen et al. 1965). This species is the
only one of the 4 Asian species for which hosts
have been recorded. Aganaspis daci was intro-
duced into Florida (USA) where it established suc-
cessfully on Anastrepha suspense (Loew), al-
though in low numbers (Baranowski et al. 1993). It
was released in Mexico (Jimenez-Jimenez 1956),
and Costa Rica (Wharton et al. 1981) for biological
control of Anastrepha spp., but its establishment
in both countries is doubtful (Wharton et al. 1998).
Aganaspis pelleranoi is a widespread neotropi-
cal species that occurs from Mexico to Argentina
and is most commonly found attacking Anas-
trepha larvae in several host plant species
(Ovruski et al. 2000). This eucoiline species is a
potential candidate for biological control ofAnas-







Ovruski et al.: A New Aganaspis Species


trepha species of economic importance (Sivinski
et al. 1997; Lopez et al. 1999).Aganaspis nordlan-
deri is only known from Costa Rica attacking lar-
vae of Ceratitis capitata (Wiedemann) and A. stri-
ata Schiner (Wharton et al. 1998) and from Brazil
attacking A. bahiensis Costa Lima, A. striata and
one lonchaeid of the genus Neosilba McAlpine
(Guimaraes et al. 1999; 2003).
Described species ofAganaspis for which biolog-
ical data have been gathered can be considered
specialists in terms of their tephritid fruit fly host
range but as generalists in terms of the number
and diversity of plant species on which they are
able to find their hosts. As far as their biology is
concerned, no thelytokous strains have been found
for any species in the genus and no described spe-
cies ofAganaspis has previously been found to ex-
hibit obligate diapause or infest diapausing species
of temperate tephritids. Nonetheless, existence of
facultative diapause was documented for popula-
tions ofA. pelleranoi living in tropical sub-decidous
and decidous forest in central Veracruz, Mexico
(Aluja et al. 1998), and in subtropical rain-forest in
northwestern Argentina (Ovruski et al. 2004).
In this paper, a new neotropical Aganaspis
species, reared from Rhagoletis ramosae Hernan-
dez-Ortiz on Juglans major var. glabrata (Torr.)
A. Heller, and Rhagoletis zoqui Bush on Juglans
mollis Engeim is described. Evidence of host spe-
cialization and diapause is furnished, and the re-
lationships between the AmericanAganaspis spe-
cies are discussed.

MATERIALS AND METHODS

The description ofA. alujai was based on quan-
titative measurements on 21 female specimens in
Mexico that were reared from Rhagoletis ramosae
and Rhagoletis zoqui recovered from single collec-
tions of Juglans major var. glabrata in Michoacan
and Juglans mollis in Hidalgo, respectively. No
males were reared. Quantitative measurements
were made with an optical micrometer installed
within Zeiss-Stemi SV6 stereo-microscopes. Scan-
ning electron microscope (SEM) images were
taken for detailed observation of characteristic
morphological features. Specimens were prepared
for SEM by taking them out of a 70% alcohol solu-
tion and placing them in 10% KOH for at least 10
minutes to clear structures. Specimens were then
transferred to a Petri dish lined with a paper
towel to allow room temperature evaporation of
KOH. After 1 hour, specimens were cut with dis-
secting tools and placed on top of a carbon conduc-
tive tab affixed to a SEM stub. The specimens
were coated with gold-palladium in a JEOL
Model SINECOAT-JSC-1100. Images were taken
with a JEOL Model JSM-5600LV microscope.
Light microscopy images were taken with a Mac-
roFire camera mounted on a MZ16Apo stereomi-
croscope and post-processed with AutoMontageTM


and PhotoShop. Terminology for the descrip-
tions follows Wharton et al. (1998), with modifica-
tions as used by Fontal-Cazalla et al. (2002).

RESULTS

Species Description

Aganaspis alujai Wharton & Ovruski, n. sp.
(Fig. la-f)

Female. Body length 2.5-3.0 mm; fore wing
length 2.4-2.9 mm. Head and mesosoma black,
metasoma mostly black, tending to castaneous
brown apically and ventrally, with pattern of
lighter coloration slightly variable among speci-
mens, antennae black apically grading to dark
brown over basal half, mandible brown, with apical
teeth black, postgena dark brown, legs yellow
brown with telotarsi, most of mid and hind coxa,
and swollen portion of all femora dark brown. Head
1.9-2.2 times wider than long and 1.03-1.06 times
wider than mesoscutum in dorsal view, 1.11-1.48
times higher than wide in frontal view. Toruli ele-
vated laterally, forming short, shallow, orbital fur-
row between antennal base and internal eye mar-
gin. Vertex and frons bare, polished, unsculptured;
occiput bare, densely and obliquely striate laterally
and dorsal-laterally, smooth and polished mid-dor-
sally. Face shining, polished; clypeal furrows dis-
tinct, widely separated, very weakly converging
dorsally, extending nearly half distance from ven-
tral margin of clypeus to toruli; row of 3-4 long,
weakly decurved setae in each clypeal furrow; 3-4
pairs of decumbent, medially directed setae above
and lateral dorsal margin of clypeal furrow; longi-
tudinal row of 6-7 setae extending along inner eye
margin onto malar sulcus. Eyes weakly bulging,
temples distinctly receding in dorsal view; 1.25-1.5
times higher than wide in lateral view, without vis-
ible setae at 60x. Temple 0.5-0.66 times as long as
eye. Malar space 0.45-0.55 times height of eye; ma-
lar space with deep, complete malar sulcus; gena
weakly striate and setose ventrally. Ocelli ar-
ranged in an isosceles triangle, posterior ocelli
widely separated: distance between them 1.42-1.5
times distance between posterior ocellus and eye.
Antenna (Fig. la) short, 2.4-2.5 times height of
head, without distinct club; scape and pedicel 1.9
and 1.15 times as long as broad, respectively; first 3
flagellomeres slightly broader subapically than
medially, flagellomeres 5-11 broader medially,
nearly moniliform; first flagellomere slightly
(1.06-1.13 times) longer than second; ratio of
length to maximum width, all flagellomeres:
3.1:2.5:2.32-2.4:2.0-2.25:1.6-1.7:1.6-1.7:1.6-1.7:1.5-
1.6:1.5-1.6:1.5-1.6:1.9-2.0. Mesosoma 1.25 times
longer than high; 1.73-1.79 times longer than wide;
1.36-1.45 times higher than wide. Pronotal plate
(Fig. Ib) protruding above anterior margin of meso-
scutum, weakly notched; median bridge 1.42-1.5







Florida Entomologist 90(4)



- lb


December 2007


le


If






-.- '-
rt~


Fig. 1.Aganaspis alujai. a, female antenna; b, pronotal plate; c, scutellar disc and cup in profile; d, scutellar disc
and cup in dorsal view; e, fore, mid, and hind coxae in lateral view; f, complete fore wing, with arrow showing the
area behind Rs&M vein between Sc+R1 and basal section of Cul.


times wider than anterior ocellus, and 0.35-0.40 arm; posterior part 2.16-2.20 times wider than
times as wide as posterior part; anterior part of long, with 10-11 setae near posterior margin.
pronotal plate 5.65-5.70 times wider than long, Pronotum laterally polished, unsculptured, with
with 5-6 setae on posterior margin of each lateral tuft of white setae anteriorly, and 8-10 long, com-







Ovruski et al.: A New Aganaspis Species


pletely decumbent setae posteriorly. Mesoscutum
1.08-1.10 times as wide as long; notauli absent, rep-
resented by a row of 6-7 short setae on each side,
each row converging anteriorly with lateral row of
8-9 short setae; parascutal impression weak,
largely confined to area mesad tegula; mesonotal
keel absent or barely visible as a weak median ele-
vation anteriorly, replaced by weak median depres-
sion on posterior 0.3. Scutellar disc (Fig. Ic, d) dis-
tinctly and finely reticulate; posterior margin of
disc rounded in dorsal view, sinuate in profile;
height of scutellum about 0.9 times length of
scutellar plate in lateral view; fovea behind lateral
keel of scutellum 1.56-1.67 times higher than long.
Scutellar plate (Fig. Id) large, subrounded, 1.2-1.4
times longer than wide, posterior margin strongly
rounded, never truncate, anterior margin slightly
extending through scutellar fovea; width of plate
0.64-0.66 times width of disc; surface nearly flat,
with either 2 or 3 pairs of lateral punctures, each
bearing a short, erect seta. Pit of scutellar plate
(Fig. Id) small, elliptical, abutting posterior rim of
plate, 2.0-2.15 times wider than long, and 0.3-0.4
times width of scutellar plate. Lateral bars (Fig. Ic)
long, 0.92-0.96 times as long as scutellar plate,
smooth to finely striate. Scutellar fovea large, deep,
2.09-2.15 times wider than long. Mesopleuron
smooth, completely without setae; precoxal carina
straight along ventral margin, curving dorsally
along anterior 0.3 and posterior 0.2. Metanotum
densely pubescent laterally. Metapleuron smooth
above, rugulose ventral-posteriorly, with strongly
carinate posterior margin; bare except along poste-
rior margin. Propodeum short, 1.3-1.4 times wider
than long in dorsal view, densely white-pubescent,
the setae almost completely obscuring area laterad
lateral propodeal carina; lateral propodeal carinae
slightly diverging ventrally then abruptly broad-
ened posteriorly delimiting a round, deep fovea
posterior-medially; upper part between lateral car-
inae with long, dense, dorsally-directly, white se-
tae. Anterior face of fore and mid coxae extensively
covered with long setae; hind coxa anterior-medi-
ally with patch of short setae extending more than
half length from base, and with patches of longer
setae ventrally; mid and hind coxae with short,
dense, white pubescence on postero-dorsal margin,
and with some long setae directed backwards (Fig.
le). Fore wing (Fig. If) 2.7-2.9 times longer than
wide, with longest seta on posterior-apical margin
0.08-0.11 times maximum width of wing; marginal
cell 2.5-2.7 times as long as deep, completely closed
along wing margin; Rs 1.55-1.73 times longer than
2r; costal cell with a longitudinal row of setae on
ventral surface extending from basal to apical sec-
tion of cell, otherwise bare; cell immediately distad
basal vein uniformly setose, with vertical row of 3-5
setae parallel to basal vein (Fig. If). Metasoma
1.39-1.45 times longer than high; syntergum 0.85-
0.92 times as long as metasoma, and 1.09-1.12
times length of mesosoma; hairy ring at base of


syntergum absent mid-ventrally, otherwise com-
plete and very well-developed; syntergum emargin-
ate dorsally, maximum length in lateral view 1.15-
1.2 times length along mid-dorsal line; minute
punctures present on posterior third of syntergum
and on visible portions of remaining terga.

Material Examined

The above description is based on 21 females,
all reared from Rhagoletis ramosae from a single
collection of Juglans major var. glabrata. No
males were reared. Female holotype, Mexico, Mi-
choacan, Zirimicuaro, 12-IX-02, ex. Rhagoletis ra-
mosae in Juglans major var. glabrata, Diaz-Fleis-
cher collector (Instituto de Ecologia, A.C.);
paratypes, same data as holotype, 6 females (In-
stituto de Ecologia, A.C.); 12 females (Texas A&M
University); 1 female (US National Museum); 1
female (University of California, Riverside). The
coordinates for the site are 19024'N, 10158'W, at
1,306 m above sea level. The environmental con-
ditions for the site are mean annual temperature
17C and mean annual precipitation 1,350 mm.
Additional examined material includes 78 fe-
males emerging from pupae of Rhagoletis zoqui
collected in 19-IX-03, from Juglans mollis collected
at Lagunita, Hidalgo, Mexico, 2039'N, 9914'W,
2621 m above sea level. Mean annual temperature
and precipitation for the site are 16C and 500 mm.

Etymology

This species is named after Martin Aluja, in
recognition of his numerous contributions to ecol-
ogy and biology of neotropical fruit fly parasitoids.

Diagnosis

This new species differs fromA. daci and other
Old World species by the absence of obvious setae
on the female compound eye, possession of a
scutellar disc that bulges posteriorly beyond the
apex of the scutellar plate, and absence of a dis-
tinct cleft in the posterior carinate margin of the
metapleuron. The new species shares these and
other features with the 2 previously described
New World species,A. nordlanderi and A. pellera-
noi, but differs from them primarily in details of
the scutellar plate, venation, and setal pattern of
the fore wing. The scutellar plate extends into the
scutellar fovea and is relatively flat in bothA. pel-
leranoi (Fig. 2b, c) and A. alujai (Fig. Ic, d)
whereas in A. nordlanderi (Fig. 3a, b) the plate
does not extend into the scutellar fovea and is
weakly but distinctly recurved in profile. The
plate is distinctly larger (relative to the disc) and
more parallel-sided inA. pelleranoi than inA. alu-
jai. The portion of R1 between 2r and the wing
margin is much longer in A. nordlanderi than in
either of the other 2 species, resulting in a greater








Florida Entomologist 90(4)


December 2007


2a


-m"























































Fig. 2.Aganaspis pelleranoi. a, complete fore wing; b, scutellar disc in dorsal view; c, scutellar d7isc in profile.
Fig. 3.Aganaspis nordlanderi. a, scutellar disc in dorsal view; b, scutellar disc in profile.
Fig. 4.Aganaspis daci. a, scutellar disc in dorsal view; b, scutellar disc in profile.







Ovruski et al.: A New Aganaspis Species


Rs/2r ratio for A. nordlanderi and a marginal cell
in which the greatest height is more anteriorly
displaced (see Figs. If, 2a). The marginal cell is
closed by a distinct vein in bothA. pelleranoi and
A. alujai (usually more distinctly pigmented in
A. alujai), but the marginal cell is open in A. nor-
dlanderi. The costal cell is densely setose in A.
nordlanderi but largely reduced to a single me-
dian line of setae in the other 2 species (the line of
setae extends the full length of the cell, but be-
comes a double row of setae over the apical 0.3-
0.4). The cell immediately distad the basal vein is
more or less uniformly setose in A. alujai and A.
nordlanderi (Fig. If) but is bare adjacent the
basal vein inA. pelleranoi (Fig. 2a). The new spe-
cies can be separated from the previously de-
scribed New World species by its darker colora-
tion. Specimens of A. pelleranoi and A. nordlan-
deri that have been reared from C. capitata and
various species ofAnastrepha Schiner have the fe-
male metasoma generally orange and the legs en-
tirely yellow to yellow-brown whereas the meta-
soma and legs are largely black inA. alujai.

Comments
Based on the characters discussed above, A.
alujai shares more features with A. pelleranoi
than with A. nordlanderi. However, when com-
pared with species such as A. daci, the character
states found in A. nordlanderi appear to be de-
rived relative to those in A. pelleranoi and A. alu-
jai, suggesting that the resemblance between the
latter 2 species may not be a result of shared, de-
rived features. Further assessment of relation-
ships would be premature in the absence of a clear
outgroup for the New World species. Problems as-
sociated with the generic classification were dis-
cussed in Wharton et al. (1998), in which the use
of the name Trybliographa for these species (most
recently by Diaz et al. 2006) was rejected.
Prior to the description of Aganaspis by Lin
(1987), Legner & Goeden (1987) reared an incom-
pletely identified eucoiline figitid from the walnut
husk fly, Rhagoletis complete Cresson, infesting
Juglans microcarpa Berland in western Texas
and southeastern New Mexico. Examination of
voucher material (1 male, 1 female from Davis
Mountains, TX) from the University of California,
Riverside collection revealed that specimens from
Texas are nearly identical to those from central
Mexico and represent either another population


of the same species or a very closely related spe-
cies. A third specimen, collected from walnut
trees in Lincoln Co., New Mexico, is identical to
those from Texas. There are minor differences in
venation (Cul is distinct in the material from
Mexico, but the basal vein is shorter in specimens
from the U.S., and lacks the angular extension of
Cul ventrally) and the scutellum. Thus, we prefer
to restrict the definition of A. alujai to material
from Michoacan and Hidalgo until material from
intermediate areas can be examined.

Biology
The specimens forming the type series were
reared from 1,250 puparia recovered from fruits of
J major var. glabrata in Michoacan. From these
puparia, 351 R ramosae adults with a sex ratio
roughly equivalent to 0.5 and 21 female parasitoid
specimens were recovered. The remaining 878 pu-
paria yielded no flies or parasitoids. Additional col-
lections of Juglans mollis in 2003 in Lagunita,
Hidalgo yielded 1000 pupae of Rhagoletis zoqui,
from which 397 adult R. zoqui at a sex ratio
roughly equivalent to 0.5 and 78 female parasitoid
specimens were recovered. Pupae were collected
from infested fruit on Sep 12, 2002, and on Sep 19,
2003, and kept at ambient temperature and hu-
midity at the INECOL in Xalapa in a pupal rearing
chamber with no temperature control. Xalapa has
a mean annual relative humidity of 66% and mean
annual temperature of 18.6C (CONAGUA, http://
smn.cna.gob.mx/productos/observatorios/histor-
ica/jalapa.pdf). Adult tephritid hosts from Michoa-
can emerged from May 8 through Aug 15, 2003
with an average time from pupal recovery to adult
emergence of 298 days. Parasitoids from Michoa-
can also entered diapause but emerged as adults
from parasitized puparia from Aug 5 through Sep
18, 2003, with an average time from pupal recov-
ery to adult parasitoid emergence of 348 days (Fig.
5A). Adult tephritid hosts from Hidalgo emerged
from Mar 24 through Aug 20, 2004 with an aver-
age time from pupal recovery to adult emergence of
294 d. Parasitoids from Hidalgo also entered dia-
pause and emerged as adults from parasitized pu-
paria from Aug 20 through Oct 24, 2004 with an
average time from pupal recovery to adult parasi-
toid emergence of 387 d (Fig. 5B). In the labora-
tory, the average time from adult emergence from
parasitized pupae until death of A. alujai was
12.76 d at 27C and 75% R.H. (n = 21).

EY


The following key includes all Aganaspis species occurring in the New World.
1. Eye covered with scattered setae (more readily visible in females than males); scutellar plate long, protruding
behind scutellar disc (Fig. 4a, b); posterior carinate margin of the metapleuron interrupted by a median cleft
or depression (Introduced to New World) ........... . ........................ ... A. daci (Weld)
-Eye without setae; scutellar plate short, not extending posteriorly beyond scutellar disc (Figs. Ic, d; 2b, c;
3a, b); posterior carinate margin of the metapleuron continuous, not interrupted by a cleft ............ 2







Florida Entomologist 90(4)


December 2007


"C' 1 21 41 61 81 101 121 141 161 181 201 221 241 261 281 301 321 341 361 381 401
-ol
I-
0 50
E 45


S30
c 40 -- Rhagoletis zoqui


^ -25- Aganaspis alujai .
E 20
Z 15
10


1 21 41 61 81 101 121 141 161 181 201 221 241 261 281 301 321 341 361 381 401



Days elapsed from pupal recovery to adult emergence

Fig. 5. Daily post-diapause emergence patterns of (A) adult Rhagoletis ramosae (black histogram) and adult
Aganaspsis alujai (grey histogram), and (B) adult Rhagoletis zoqui (black histogram) and adult Aganaspsis alujai
(grey histogram).


2. Costal cell densely setose; scutellar cup not extending through scutellar fovea (Fig. 3a), with a long anterior
bridge; posterior margin of scutellar disc slightly concave medially and thus weakly bilobed in dorsal view;
marginal cell completely open. ............... ...................... A. nordlanderi Wharton
-Costal cell with a single longitudinal row of setae on ventral surface, otherwise largely bare; scutellar cup ex-
tending through scutellar fovea, with a short anterior bridge (Figs. Id, 2b); posterior margin of scutellar
disc rounded in dorsal view; marginal cell completely closed ..................................... 3
3. Scutellar plate large (Fig. 2b), nearly parallel-sided, 0.71-0.85 times as wide as scutellar disc; fore wing with area
immediately behind basal vein bare (Fig. 2a); hind leg pale, yellow to yellow-brown .. .A. pelleranoi (Brethes)
-Scutellar plate small (Fig. Id), rounded, 0.60-0.66 times as wide as scutellar disc; fore wing with area behind basal
vein densely setose (Fig. If); hind leg (especially coxa and femur) extensively black ..... A. alujai, new species


DISCUSSION

The observed delay in emergence of adult par-
asitoids with respect to that of adults of their te-
phritid hosts also has been documented by Feder
(1995) for Utetes canaliculatus (Gahan) (reported
as Opius lectus Gahan) attacking Rhagoletis
pomonella Walsh on hawthorn. Such synchroniza-
tion is undoubtedly commonplace among parasi-
toids of univoltine hosts, allowing the hosts to
reach sexual maturity, mate, and oviposit, and al-
low their larvae to reach a suitable developmental
stage for parasitism. (For more detailed discus-
sion of various aspects associated with host-para-


sitoid synchronization, see Godfray 1994.) The
synchronization betweenA. alujai and their hosts
R. ramosae and R. zoqui suggests a strong degree
of host specialization. Although we recognize that
the temperatures at which we took longevity
records for adult parasitoids might exceed those
encountered in their natural habitats, we can con-
clude that the lifespan of adults is relatively short
and therefore synchronized emergence with their
host is a key adaptation ifA. alujai is specialized
to parasite walnut-infesting Rhagoletis flies with
obligate diapause as appears to be the case.
Parasitoids of multivoltine host flies, such as
Anastrepha ludens (Loew),A. obliqua (Macquart),







Ovruski et al.: A New Aganaspis Species


A. fraterculus (Wiedemann),A. striata, andA. ser-
pentina (Wiedemann), also may exhibit faculta-
tive diapause, allowing them to overcome periods
of pronounced host scarcity (Aluja et al. 1998; Siv-
inski et al. 2000). For example, someA. pelleranoi
individuals attacking Anastrepha larvae in Psid-
ium guajaua L. diapaused for up to 11 months in
central Veracruz, Mexico (Aluja et al. 1998).
The sites where Aganaspis were found associ-
ated with walnut-infesting flies in the Rhagoletis
suavis group are separated from each other by
great distance and geographically isolated from
each other by numerous barriers of orographic
and climatic nature. In both sites only females
were recovered from infested tephritid puparia,
strongly suggesting that both strains collected
are fully parthenogenic (i.e., thelytokous).
Whether or not thelytoky arose independently in
both cases or had a common origin could be deter-
mined through genetic analysis of individuals
from both populations or by examining whether
or not both strains possess and share similar
strains of sex ratio distorting agents. The species
ofAganaspis (as currently defined), thus offer ex-
citing possibilities to explore the evolution of the-
lytoky in a group not known to possess this trait
as well as function and evolution of diapause in
temperate vs tropical taxa.

ACKNOWLEDGMENTS

We thank Francisco Diaz-Fleisher and Martin Pale
for collection of infested fruit, University of California,
Riverside (UCR) for the loan of specimens, and Matt
Buffington (UCR and USDA/SEL) for information and
facilitating the loan from UCR. This work was funded
by the Mexican Campana Nacional contra Moscas de la
Fruta (DGSV-SAGARPA-IICA). SO acknowledges fi-
nancial support from the Consejo Nacional de Investi-
gaciones Cientificas y T6cnicas de Argentina
(CONICET, grant PIP No. 5129). RAW was supported in
part by a Mexico-Texas A&M Conacyt grant and in part
by NSF/PEET grant DEB 0328922.

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Myartseva et al.: Parasitoids of T floridensis


A REVIEW OF PARASITOIDS (HYMENOPTERA: CHALCIDOIDEA)
OF TRIALEURODES FLORIDENSIS (HEMIPTERA: ALEYRODIDAE)
WITH DESCRIPTION OF A NEW SPECIES FROM MEXICO

S. N MYARTSEVA, E. RUIZ-CANCINO AND J. M. CORONADO-BLANCO
Division de Estudios de Postgrado e Investigaci6n, UAM Agronomia y Ciencias,
Universidad Aut6noma de Tamaulipas, 87149, Ciudad Victoria, Tamaulipas, M6xico
E-mail: smyartse@uat.edu.mx; eruiz@uat.edu.mx; jmcoronado@uat.edu.mx

ABSTRACT

A new species, Encarsia citricola sp. n., reared from Trialeurodes floridensis (Quaintance)
collected in the State of Guanajuato, Mexico, is described and illustrated. A review and a key
to 9 parasitoid species attacking T floridensis are provided. In addition, 4 species of chalci-
doid wasps are recorded for the first time as parasitoids of T floridensis in the New World.

Key Words: Encarsia citricola sp. n., Guanajuato, Aphelinidae, Encyrtidae, Signiphoridae.

RESUME

Se describe Encarsia citricola sp. n., especie emergida de Trialeurodes floridensis (Quain-
tance) recolectada en el Estado de Guanajuato, M6xico. Se proporciona una revision y clave
de nueve species de parasitoides que atacan esta mosquita blanca. Cuatro species de avis-
pas calcidoideas son nuevos registros como parasitoides de T floridensis en el Nuevo Mundo.


Translation provided by the authors.


The family Aleyrodidae (Hemiptera) is com-
prised of approximately 1450 described species in
the world. Whiteflies are detrimental pests at-
tacking a wide variety of agricultural crops and
ornamental plants, predominantly in the tropics
and subtropics (Martin et al. 2000). About 50
whitefly species are commonly found infesting
economic plants around the world, and 33 species
in 20 genera are considered common and econom-
ically important in the Southeastern United
States (Martin 1987; Hodges & Evans 2005).
Nguyen et al. (1993) listed 73 species in 20 gen-
era, including 7 species in the genus Trialeurodes
(Cockerell) in their catalog of the Aleyrodidae spe-
cies associated with citrus worldwide. Carapia et
al. (2003) reported 13 species in the genus Tri-
aleurodes from Mexico, of which 6 occur on citrus.
Trialeurodes floridensis (Quaintance 1900),
the guava whitefly or the avocado whitefly, is a
polyphagous insect feeding on many plant species
from 20 families, and is widely distributed in the
New World from United States of America (Flor-
ida, Texas, Arizona) to Venezuela (Mound & Hal-
sey 1978). In Mexico, the current known distribu-
tion for T floridensis is from the State of Nuevo
Le6n (Russell 1963).
Five species of chalcidoid wasps in the genus
Encarsia Foerster (Aphelinidae) are known at
present to attack T. floridensis: Encarsia hispida
De Santis, E. meritoria Gahan, E. nigricephala
Dozier, E. pergandiella Howard and E. telema-
chusi Evans (Evans & Serra 2002; Noyes 2006).


This paper presents the list of 9 species of para-
sitoids attacking T. floridensis: 7 species of Encar-
sia, including 1 new species ofEncarsia (Aphelin-
idae), 1 species ofMetaphycus (Encyrtidae), and 1
species of Signiphora (Signiphoridae). A key for
identification of these parasitoid species, informa-
tion on their distribution and alternative hosts,
and new records of specimens collected in Mexico
are provided.

MATERIALS AND METHODS

A large series of whitefly Trialeurodes floriden-
sis pupae were collected on leaves of Citrus sinen-
sis (L.) in San Miguel de Allende, Guanajuato,
Mexico, during Nov 13-16, 2005. In the laboratory,
leaves with colonies of whitefly pupae were main-
tained in plastic containers for parasitoid rearing.
Female and male specimens from some of the
emerged adult wasps were mounted in Canada
balsam for morphological study and identifica-
tion, following the slide-mounting method for
Chalcidoidea described by Noyes (1982). A key for
the identification of these species is provided,
based on characters observed on specimens
mounted on microscopic slides.
The whitefly species from which the parasi-
toids emerged were identified with keys to the
puparium (=pupal cases) by Hamon (2002), Mar-
tin (1987), and Hodges & Evans (2005). Parasi-
toid species were identified with the pictorial
guide to the species of Encarsia parasitic on











whiteflies in North America (Schauff et al. 1996),
the guide to Encarsia parasitoids of Bemisia
tabaci (Polaszek et al. 1992), the key to En-
cyrtidae of Costa Rica (Noyes 2004), and the key
to Metaphycus species parasitizing whiteflies
(Myartseva 2006). Some specimens used in this
study are deposited in the Entomological
Museum of the University of Tamaulipas, UAM
Agronomia y Ciencias, at Ciudad Victoria,
Tamaulipas, Mexico. Morphological terminology
used for the chalcidoid wasps follows that of
Hayat (1998) and Schauff et al. (1996). Taxo-
nomic information, hosts, geographic distribu-
tion, collected material, and references (only


December 2007


information for Mexico) are provided for each
parasitoid species.

RESULTS AND DISCUSSION

Eight parasitoid species of T floridensis were
collected in different Mexican States, including 6
species of the genus Encarsia, 1 species of Meta-
phycus, and 1 species of Signiphora. Moreover,
Encarsia telemachusi Evans is recorded as a par-
asitoid of T floridensis in Haiti. A new species,
Encarsia citricola, is described and illustrated
and a review and key to 9 parasitoid species at-
tacking T floridensis are given.


KEY FOR IDENTIFICATION OF PARASITOIDS ATTACKING TRIALEURODES FLORIDENSIS (FEMALES)

1. Antennal flagellum 4-segmented with three transverse funicular segments and one, greatly elongate club seg-
ment ......................................................... Signiphora aleyrodis Ashmead
-Antennal flagellum 6 or 9-segmented, club 2-3-segmented, not greatly elongate ...................... 2
2. Antennal flagellum 9-segmented, with 6-segmented funicle and 3-segmented club; marginal vein of fore wing
very short, much shorter than stigmal vein. ............................... Metaphycus troas Noyes
-Antennal flagellum 6-segmented with 3-4-segmented funicle; marginal vein of fore wing long, much longer
than stigm al vein......................................................... .......... 3
3. Tarsus of middle leg 4-segm ented ............................................................ 4
Tarsus of middle leg 5-segmented ............................................................ 7
4. Fore wing with an asetose area around the stigmal vein; metasoma completely yellow, midlobe of mesoscutum
bearing with two pairs of setae and with its anterior third to half and head dark brown; first funicular seg-
ment twice as long as wide ........................................ Encarsia nigricephala Dozier
-Fore wing without an asetose area around the stigmal vein ....................................... 5
5. Metasoma yellow with dark brown lateral margins; first funicular segment about 0.5 times as long as second
segment; midlobe of mesoscutum with 4-6 pairs of setae .................. Encarsia variegata Howard
-Metasoma completely yellow; head, mesoscutum, axillae and base of metasoma sometimes infuscate....... 6
6. Second funicular segment intermediate in length between first and third segments and usually without linear
sensilla; sixth segment with pointed apex and elongate, 1.2-1.3 times as long as fifth segment. Male with
fifth and sixth flagellar segments separated ............................ Encarsia hispida De Santis
-Second funicular segment as long as, or slightly shorter than third segment and usually with one linear
sensillum; sixth segment about as long as fifth. Male with fifth and sixth flagellar segments fused
................................................................ Encarsia meritoria Gahan
7. Fore wing with an asetose area around the stigmal vein .......................................... 8
-Fore wing without an asetose area around stigmal vein and infuscate below marginal vein; metasoma com-
pletely dark brown; apical segment of club infuscate ........................ Encarsia citricola sp. n.
8. Body completely yellow. Fore wing hyaline ............................... Encarsia telemachusi Evans
-Body with pronotum, triangular spot on midlobe of mesoscutum and larger area on metasoma slightly infus-
cated. Fore wing slightly infuscated beneath venation ................. Encarsia pergandiella Howard

SYNOPSIS OF PARASITOIDS ATTACKING Coloration. Head yellow; face, cheeks (genae)
TRIALEURODES FLORIDENSIS and occiput below foramen brownish; interocellar
triangle and two triangular spots on frontovertex
Family Aphelinidae behind ocelli infuscated. Antennae yellowish-

1. Description ofEncarsia citricola Myartseva sp. nov. white; scape, pedicel and apical segment of club
slightly infuscated. Pronotum, axillae, tegulae,
Female. Body length: 0.80 mm (without ovipos- mesopleuron and propodeum brown; midlobe of
itor). mesoscutum brownish-yellow, darker along the


Florida Entomologist 90(4)







Myartseva et al.: Parasitoids of T floridensis


middle; scutellum, metanotum and side lobes of
mesoscutum (except apices) light yellow. Fore
wings infuscated below marginal vein, with dark
and strong setae on disc, apical half of disc with
short, thin and light station; marginal vein
slightly infuscated. Legs whitish; middle coxae,
hind coxae and hind femora brownish. Metasoma
brown, ovipositor sheaths yellowish-white.
Morphology. Head about as wide as thorax and
slightly wider than height. Frontovertex 0.5 times
as wide as head width and slightly wider than
long (9:8). Ocelli forming slightly obtuse triangle;
distance between hind ocelli about 0.5 times as
long as distance from hind ocellus to eye and
about as long as distance to occiput margin. Eyes
finely and rarely setose, about 1.4 times as long as
cheeks. Malar sulcus present. Mandible with
three teeth (Fig. 1). Antennae (Fig. 2) inserted im-
mediately at the level of lower margin of eyes.
Distance between toruli about 0.5 times as long as
distance to eye and twice as long as distance to
mouth margin. Scape about 4 times as long as
wide; pedicel 1.5 times as long as wide; first funic-
ular segment shorter than other flagellar seg-
ments and 1.7 times as long as wide, other funic-
ular segments subequal in length and about 1.8
times as long as wide; club 2-segmented, slightly
wider than funicle and about as long as 2 preced-
ing funicular segments combined. First funicular
segment without sensillum, other flagellar seg-
ments with one linear sensillum each, apical seg-
ment with 2 sensilla. Mesosoma with reticulate
sculpture, distinctly visible on midlobe of mesos-
cutum and axillae. Midlobe of mesoscutum (Fig.
3) with 4 pairs of setae, side lobes each with 3 se-
tae, each axilla with 1 seta; midlobe about twice
as wide as long; scutellum shorter and slightly
less than twice as wide as long. Scutellar placoid
sensilla closely placed, separated by distance
about one diameter of sensillum. Distance be-
tween posterior scutellar setae about 1.3 times as
long as that between anterior setae. Fore wing
(Fig. 4) 2.6 times as long as wide, its marginal
fringe about 0.25 times as long as the wing's max-
imum width; disc with bare base, on apical middle
with more short and thin station than below
marginal vein. Submarginal vein with 2 setae,
and 1 long seta on apex; marginal vein with 7 se-
tae along anterior margin and subequal in length
to submarginal vein. Hind wing about 7 times as
long as wide, its marginal fringe longer than max-
imum width of wing (4:3). Tarsal formula 5-5-5.
Midtibial spur (Fig. 5) shorter than basitarsus
(4:5); basitarsus subequal in length to two next
tarsal segments combined. Tergites 1-7 with 2-2-
2-2-2-4-4 setae, respectively. Ovipositor (Fig. 6)
exserted, its base inserted at the level of third
tergite. Ovipositor 1.5 times as long as middle
tibia. Third valvula 0.5 times as long as second
valvifer and 0.3 times as long as ovipositor.
Male. Body length: 0.70-0.80 mm.


Coloration. In coloration similar to female, but
frontovertex without infuscate triangular spots,
antennae without infuscated segments, fore
wings hyaline.
Morphology. Head about 1.2 times wider than
height and 1.8 times wider than frontovertex.
Eyes 1.6 times as long as cheeks. Antennal scape
3.3 times as long as wide (Fig. 7). First funicular
segment 2.5 times as long as wide, funicle seg-
ments 2nd to 4th about 2.2 times as long as wide;
club suffused, 3.7 times as long as wide and about
as long as 2 preceding funicular segments com-
bined. All flagellar segments each with 2 sensilla.
Fore wing 2.5 times as long as wide, its marginal
fringe about 0.3 times as long as wing width. Hind
wing about 6.2 times as long as wide. Genitalia
about 0.6 times as long as middle tibia.
Comments. The female of the new species is
similar to the female of Encarsia quercicola
(Howard), described from Aleuroplatus [=Aley-
rodes]Aleyrodes gelatinosus Cockerell on Quercus
sp. in California, USA. According to the original
description by Howard (1908) and the redescrip-
tion by Viggiani (1986), E. citricola can be distin-
guished by the following characters: in E. querci-
cola-midlobe of mesoscutum with 4+2 setae;
middle coxae pale; club subequal in length to third
and fourth preceding segments combined; ovipos-
itor with base arising on the level of Ti; third val-
vula about one-fourth of the total ovipositor
length; scape about 5 times as long as wide; hind
portion of mesoscutum lemon yellow. In E. citri-
cola-midlobe of mesoscutum with 4+2+2 setae;
middle coxae dark brown; club longer than third
and fourth preceding segments combined; ovipos-
itor with base arising at the level of T3; third val-
vula 0.3 times as long as ovipositor; scape about 4
times as long as wide; midlobe of mesoscutum
brown-yellowish, along middle part fuscous.
Hayat (1998) divided the genus Encarsia spe-
cies into more than 20 species groups. E. citricola
sp. n. can be included in the strenua group.
Material. Holotype female: Mexico, Guana-
juato, San Miguel de Allende, ex Trialeurodes
floridensis on Citrus sinensis, 16-XI-2005 (S.
Myartseva). Paratypes: same data as holotype, 2
females and 2 males. All specimens mounted on
slides in Canada balsam. Holotype and 1
paratype male will be deposited in UCR (Univer-
sity of California, Riverside, USA), other
paratypes in the Entomological Museum of Uni-
versity of Tamaulipas, Cd. Victoria, Mexico.

2. Encarsia hispida De Santis 1948

Synonym: Encarsia brasiliensis (Hempel 1904);
synonymy according to Polaszek et al. 2004.
References: De Santis (1979); Hernandez-Suarez
et al. (2003); Myartseva & Ruiz Cancino (2000);
Myartseva et al. (2000, 2006); Noyes (2002, 2006);
Polaszek et al. (1992, 2004); Schmidt et al. (2001).







Florida Entomologist 90(4)


Figs. 1-7. Encarsia citricola Myartseva, sp. n.: 1-mandible, 2-antenna, female, 3-
and scutellum, 4-basal half of fore wing, 5-ovipositor, 6-antenna, male.


Hosts: Aleuroglandulus malangae [=subtilis
Bondar], Aleurothrixus floccosus, A. porteri, Aleu-
rotrachelus espunae [=rhamnicola (Goux)], A.
rhamnicola, A. socialis, A. trachoides, Aleurotuba
jelinekii, Aleyrodes horridus [=Aleurothrixus floc-


-midlobe of mesoscutum


cosus (Maskell)], A. proletella, A. singularis, A.
spiraeoides, Bemisia tabaci, B. tuberculata, Creni-
dorsum aroidephagus, Dialeurodes sp., Lecanoi-
deus floccissimus, Lipaleyrodes sp., Metaleurodi-
cus minimus, Parabemisia myricae, Siphoninus


December 2007







Myartseva et al.: Parasitoids of T floridensis


phillyreae, Tetraleurodes acaciae, Trialeurodes
abutiloneus, T floridensis, T ricini, T vaporari-
orum, T variabilis.
In Mexico this species was reared from Bemi-
sia tabaci-complex, Tetraleurodes sp., and Tri-
aleurodes variabilis.
World distribution: Nearctic USA, Mexico),
Neotropical (widespread), Palaearctic (Europe),
Afrotropical (South Africa), Oriental, Australa-
sian (Australia).
Mexican Distribution: Guerrero, Sinaloa,
Tabasco, Tamaulipas, Yucatan.
Material: Mexico, Tamaulipas, Ciudad Victo-
ria, ex Bemisia tabaci-complex on Euphorbia sp.,
2 females, 29-VII-1998, 1 female, 20-VII-1998, 1
female, 19-X-1998, sweeping,1 female, 11-XI-
1998; G6mez Farias, El Nacimiento, ex Tetraleu-
rodes sp. on Bauhinia divaricata, 1 female, 3-II-
1999 (S. Myartseva).
Comments. Encarsia hispida was considered
as a junior synonym of E. meritoria Gahan 1927
(Viggiani 1989; Schauff et al. 1996). According to
Polaszek et al. (1992), they are distinct species,
and molecular evidence (Babcock et al. 2001) sup-
ports this view.

3. Encarsia meritoria Gahan 1927

References: Gordh (1979); Heraty & Woolley
(1999); Myartseva & Ruiz Cancino (2000); Noyes
(2002, 2006); Myartseva et al. (2000); Polaszek
et al. (1992, 2004); Schauff et al. (1996); Schmidt
et al. (2001).
Host: Trialeurodes floridensis; all records of
hosts may refer to Encarsia hispida (Polaszek et
al. 2004).
World Distribution: Nearctic (USA, Mexico).
Mexican distribution: Chiapas.
Material: Mexico, Chiapas, Reserva El Triunfo,
sendero Palo Gordo 97/049. Trampa agua, N
15039'22", W 92o48'31", one female, 20-22-VII-
1997 (A. Gonzalez Hdz., J.B. Woolley and L. Mon-
toya), CIB B 97-057; El Encajonado, sendero El
Tuinel, red de golpeo, one female, 28-VII-1997 (A.
Gonzalez Hdz.), CIB 97-070.
Comments. Although Encarsia meritoria
clearly is very closely related to E. hispida, there
appear to be good reasons for maintaining the 2
species as distinct pending DNA sequencing of
populations of true E. meritoria, preferably from
the type locality (Polaszek et al. 2004).

4. Encarsia nigricephala Dozier 1937

References: Alarc6n (1993); De Santis (1979);
Evans (1993); Evans & Polaszek (1997); Evans &
Serra (2002); Heraty & Woolley (1999); Myart-
seva & Ruiz Cancino (2000); Myartseva et al.
(2006); Noyes (2002, 2006); Polaszek et al. (1992);
Schauff et al. (1996); Schmidt et al. (2001);
Schuster et al. (1998).


Hosts: Aleurodicus dispersus, Aleurotrachelus
atratus, A. trachoides, Bemisia tabaci, B. argenti-
folii [=tabaci], B. euphorbiae (probably B. poinset-
tiae Hempel), Crenidorsum sp., Dialeurodes
kirkaldyi, Tetraleurodes acaciae, Trialeurodes
abutiloneus, T. floridensis, T. vaporariorum.
In Mexico this species was reared from the Be-
misia tabaci-complex, Tetraleurodes acaciae and
Trialeurodes vaporariorum.
World distribution: Nearctic (USA, Mexico),
Neotropical (widespread), Oriental (Reunion), Pa-
cific Islands (Micronesia).
Mexican distribution: Sinaloa, Tabasco,
Tamaulipas.
Material: Mexico, Tamaulipas, Ciudad Victo-
ria, ex Trialeurodes vaporariorum, 2 females, 2
males, 25.I-4.II.2006 (S. Myartseva).
Comments. New State record for Tamaulipas,
Mexico.

5. Encarsia pergandiella Howard 1907

Synonyms: Encarsia versicolor Girault, 1908;
Encarsia bemisiae De Santis, 1981; Encarsia
tabacivora Viggiani, 1985.
References: Alarc6n (1993); Arredondo et al.
(1994); Hennessey et al. (1995); De Santis (1981)
(as bemisiae); De Santis & Fidalgo (1994); Evans
(1993); Evans & Serra (2002); Gordh (1979); Heraty
& Woolley (1999); Mound & Halsey (1978); Noyes
(2002, 2006); Myartseva & Ruiz Cancino (2000);
Myartseva et al. (2000, 2006); Polaszek et al.
(1992); Schauff et al. (1996); Schmidt et al. (2001).
Hosts: Aleurocanthus woglumi, Aleurodicus
dispersus, Aleuroglandulus malangae [=subtilis],
Aleuroplatus coronata,A. elemerae,Aleurothrixus
floccosus, Aleurotrachelus rhamnicola, A. socialis,
A. trachoides, Aleyrodes azaleae [=Pealius aza-
leae],A. elevatus,A. lonicerae, A. proletella,A. spi-
raeoides, Asterochiton sonchi [=Trialeurodes uap-
orariorum], Bemisia afer, B. argentifolii [=tabaci],
B. tabaci, Dialeurodes citri, D. kirkaldyi, Pealius
azaleae, Trialeurodes abutiloneus, T floridensis,
T packardi, T vaporariorum, T variabilis.
In Mexico this species was reared from the Be-
misia tabaci-complex, Trialeurodes vaporari-
orum, Tetraleurodes sp. and T. mori.
World distribution: Nearctic (USA, Mexico),
Neotropical (Central and South America), Palae-
arctic (Europe), Australasian (Australia).
Mexican distribution: Colima, Mexico D.F.,
Quintana Roo, Sinaloa, Sonora, Tamaulipas. Ma-
terial. Mexico, Tamaulipas, Ciudad Victoria, ex
Bemisia tabaci-complex on Puansetia sp., 2 fe-
males, 9.IX.1998; ex Trialeurodes vaporariorum,
6 females, 25.I-6.II.2006; Jaumave, ex Trialeu-
rodes vaporariorum on Malva sp., 10 females, 4
males; Tampico, ex Tetraleurodes mori, 1 male,
27.III.2001 (S. Myartseva).
Comments. Encarsia pergandiella is a Nearctic
species. It was introduced into Israel and Europe







Florida Entomologist 90(4)


to control Bemisia tabaci (Evans & Castillo 1998;
Hernandez-Suarez et al. 2003). Information
about introduction of E. pergandiella into Mexico
againstAleurocanthus woglumi Ashby (Alvarado-
Mejia & Gonzalez-Hernandez, 1990, with refer-
ence to Dominguez & Carrillo, 1976) is erroneous.

6. Encarsia telemachusi Evans 2002

E. telemachusi was reared from Trialeurodes
floridensis in Haiti on Bauhinia divaricata
(Evans & Serra 2002).

7. Encarsia uariegata Howard 1908

References: Evans (1993); Evans & Serra
(2002); Heraty & Woolley (1999); Myartseva &
Ruiz Cancino (2000); Myartseva et al. (2006);
Myartseva & Varela Fuentes (in press); Noyes
(2002, 2006); Schauff et al. (1996); Viggiani (1996).
Hosts: Aleurodicus perseae [=Paraleyrodes per-
seae], Aleurothrixus floccosus, Paraleyrodes
naranjae, P perseae, Trialeurodes floridensis.
In Mexicothis species was reared from Trialeu-
rodes floridensis.
World distribution: Nearctic (USA, Mexico), Neo-
tropical (Colombia, Haiti, Honduras, Puerto Rico).
Mexican distribution: Chapas, Guanajuato,
Nuevo Le6n, San Luis Potosi, Tamaulipas. Mate-
rial: Mexico, Nuevo Le6n, 20 km S Monterrey, 2
females, 7.VI.2005 (S. Myartseva & E. Ruiz Can-
cino); San Luis Potosi, Tamuin, 1 female,
30.1.2005 (S. Varela Fuentes); Tamaulipas,
GuBmez, 18 females, 4 males, 10.V.2005 (S.
Myartseva). All specimens were reared from
Paraleyrodes spp. on Citrus spp. Guanajuato, San
Miguel de Allende, ex Trialeurodes floridensis on
Citrus sinensis, 2 females, 2 males, 16.XI.2005;
Chiapas, Tapachula, ex Aleyrodidae on Psidium
guajava, 2 females, 8-9.X.2006 (S. Myartseva).
Comments. The Universal Chalcidoidea Data-
base (Noyes 2006)does not record Encarsia uarie-
gata as occurring in Mexico. Schauff et al. (1996)
indicated that this species is presented in Mexico,
but did not mention any specific localities. We
reared E. uariegata from Paraleyrodes spp. col-
lected in 3 states of Mexico-Nuevo Le6n, San
Luis Potosi and Tamaulipas (Myartseva & Varela
Fuentes in press) and from the states of Guana-
juato and Chiapas. Trialeurodes floridensis repre-
sents a new host for Encarsia uariegata.

Family Encyrtidae

8. Metaphycus troas Noyes 2004

References: Myartseva (2006).
Host: Trialeurodes floridensis.
World distribution: Nearctic (Mexico), Neotro-
pical (Costa Rica).
Mexican distribution: Guanajuato.


Material: Mexico, Guanajuato, San Miguel de
Allende, ex Trialeurodes floridensis on Citrus sin-
ensis, 5 females, 5 males, 16.XI.2005 (S. Myart-
seva).
Comments. Currently, there are 5 described
species of the genus Metaphycus that are known to
parasitize whiteflies, all of which are from the Neo-
tropical region (Noyes 2004; Myartseva 2006;
Noyes& Lozada 2005). Of these, 3 species are
known from Mexico. Trialeurodes floridensis repre-
sents a new host record for the genus Metaphycus.

Family Signiphoridae

9. Signiphora aleyrodis Ashmead, 1900

References: De Santis (1979); Myartseva et al.
(2005); Noyes (2002, 2006); Ruiz et al. (2005).
Hosts: A hyperparasitoid through Aleuropla-
tus coronata, Aleurothrixus floccosus, Aleyrodes
spiraeoides, Bemisia tabaci-complex, Dialeurodes
citri, Tetraleurodes sp., Trialeurodes floridensis, T
uaporariorum. Also known from several species of
armored scales (Diaspididae).
In Mexico, was reared from Aleurothrixus floc-
cosus, Bemisia tabaci-complex, Tetraleurodes
spp., Trialeurodes floridensis and Trialeurodes ua-
porariorum.
World distribution: Nearctic (USA, Mexico),
Neotropical (widespread).
Mexican distribution: Guanajuato, Guerrero,
San Luis Potosi, Tamaulipas.
Material: Mexico, Tamaulipas, Ciudad Victo-
ria, ex Tetraleurodes sp. on Bauhinia diuaricata, 1
female, 7.VII.1998, ex Bemisia tabaci-complex on
Rosa sp., 1 female, 3.IX.1998, ex Aleurothrixus
floccosus on Citrus sinensis, 1 female, 9.V.1999, 1
female, 4.IV.2000, ex Tetraleurodes sp. on Leu-
caena puluerulenta, 2 females, 17.1. 2000, 2 fe-
males, 6.V.2000; Miquihuana, ex Tetraleurodes
sp. on Sophora secundiflora, 1 female, 1 male,
3.X.1998, 3 females, 1.IV.2001, ex Tetraleurodes
sp. on Mahonia trifoliata, 1 female, 13.V.2000;
Jaumave, ex Trialeurodes uaporariorum on Ruta
sp., 1 female, 31.111.2001. San Luis Potosi, ex Te-
traleurodes sp. on Psidium guajaua, 1 female,
12.XI.1999, 1 female, 28.X.1999. Guerrero, Acap-
ulco, ex Aleyrodidae on Pithecellobium sp., 1 fe-
male, 12.VI.2000. Guanajuato, San Miguel de Al-
lende, ex Trialeurodes floridensis on Citrus sinen-
sis, 1 female, 16.XI.2005 (S. Myartseva).
Comments. Trialeurodes floridensis represents
a new host record for Signiphora aleyrodis.

ACKNOWLEDGMENTS

The authors thank Dr. G. Evans (APHIS/PPQ/NIS,
c/o Systematic Entomology Laboratory, ARS, USDA,
BARC-West, Beltsville, Maryland, USA) for the loan of
important publications on Aphelinidae; Dr. J. Noyes
(Department of Entomology, The Natural History Mu-


December 2007








Myartseva et al.: Parasitoids of T floridensis


seum, London, UK) for loan of his excellent books about
Encyrtidae of Costa Rica; and Departamento de Estu-
dios de Postgrado e INvestigaci6n, UAM Agronomia y
Ciencias, Universidad Aut6noma de Tamaulipas,
Ciudad Victoria, M6xico, for financial support of this
work. We thank 2 anonymous reviewers for helpful sug-
gestions and corrections.


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Florida Entomologist 90(4)








Lajeunesse: Ectoparasitism of Damselflies


ECTOPARASITISM OF DAMSELFLIES BY WATER MITES
IN CENTRAL FLORIDA

MARC J. LAJEUNESSE
Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, NY 14853-2701

ABSTRACT

Little is known about the frequency of water mite ectoparasitism (Acari: Hydrachnida)
within and among damselfly species of Central Florida. Here I present a field survey of the
assemblage of damselflies and their water mite parasites at the Archbold Biological Station
(Lake Placid, FL) during late Mar, 2006. During this period, 4 species of damselfly were
abundant: Ischnura hastata (Say) and Nehalennia gracilis Morse captured only at pond
sites; and Argia fumipennis atra Gloyd and Ischnura ramburii (Selys) captured at a lake
site. Only pond damselflies had water mites, and 12.2% and 12.5% of I. hastata and N gra-
cilis were parasitized, respectively. These are 2 novel and unreported odonate-acari associ-
ations for this area. I also examined within-species differences in ectoparasitism by sex,
body size, and wing-cell fluctuating asymmetry. However, these factors did not relate to the
prevalence and intensity of parasitism in the field. My study indicates that brief surveys of
odonates in Central Florida will likely generate novel, unreported associations with para-
sitic water mites-this information is important to address the gap in natural history for
southeastern distributions of North American water mites.

Key Words: Arrenurus, Odonata, Hydrachnida, sex-bias, life-history trade-offs, body size,
asymmetry, field surveys

RESUME

Poco es lo conocido sobre la frecuencia de ectoparasitismo por parte de los acaros acuaticos
(Acari: Hydrachnida) en cada especie y entire species de pequenas libelulas (Odonata:
Zygoptera) en la region central de la Florida. Aqui se present un sondeo de campo del com-
plejo de libelulas (conocidas tambi6n como caballitos del diablo o gallegos) y sus acaros pa-
rasitos en la Estaci6n Biol6gica de Archbold (Lake Placid, Florida) durante la iltima part
de marzo del 2006. Durante de este period, de las 4 species de libelulas mas abundantes,
Ischnura hastata (Say) y Nehalennia gracilis Morse fueron capturadas solamente en sitios
de los lagitos;Argia fumipennis atra Gloyd y Ischnura ramburii (Selys) fueron capturadas
en un sitio del lago. Solamente las libelulas capturadas en los lagitos tenian acaros acuati-
cos, y 12.2% y 12.5% de los I. hastata y N. gracilis estaban parasitizadas, respectivamente.
Estas son 2 asociaciones de odonata-acari claras y que no estaban reportadas para esta area.
Tambien, se examine las diferencias intra-especificas en el ectoparasitismo por el sexo, el ta-
mano de cuerpo, y la asimetria fluctuante de las clulas del ala. Sin embargo, estos factors
no fueron relacionados con la prevalencia e intensidad del parasitismo en el campo. Mi es-
tudio indica que la realizaci6n de sondeos breves de los Odonata en la region central de la
Florida probablemente resultard en el descubrimiento de asociaciones nuevas y no reporta-
das con sus parasites de acaros acuaticos-esta informaci6n es important para dirigirse a
la brecha en la historic natural para la distribuci6n de acaros de aqua en el sureste de Ame-
rica del Norte.


Basic information on the natural history of wa-
ter mite parasitism (Acari: Hydrachnida) within
and among odonate species is scarce (Smith
1988), and is urgently needed to assess the host
range and geographic distribution of these para-
sites (Bush et al. 2001; Rolff 2001). Cook (1976)
broadly surveyed adult water mites of Florida
(mainly Hydrachnida: Arrenurus), and described
many new species, adding 35 to the 14 previously
described. However, nothing is known on the lar-
val ecology of the mites, which are often parasitic
on adult damselflies and dragonflies. Here I re-
port field patterns of larval mite parasitism
among the assemblage of damselflies species


found in the southernmost region of Lake Wales
Ridge, Central Florida.
I also examine whether sex, body size, and fluc-
tuating asymmetry relate to patterns of parasit-
ism in the field. Life-history theory predicts an in-
teraction between the reproductive effort among
sexes and susceptibility to parasitism (Poulin
1996; Johnsen & Zuk 1999). This interaction can
result in the characteristic male-bias of parasit-
ism found in mammals and dioecious plants
(Schalk & Forbes 1997; Moorse & Wilson 2002;
Cornelissen & Stiling 2005). However, the direc-
tion of this bias (either female or male) is depen-
dent on the group of parasites under study (i.e.,








Florida Entomologist 90(4)


for birds there is a female bias in blood parasites,
see McCurdy et al. 1998). Among invertebrate
hosts, odonates are among the very few to show
sex-biases in parasitism (Hecker et al. 2002; La-
jeunesse et al. 2004; cf. Sheridan et al. 2002; Kelly
2005); although studies finding no bias are also
common (Sheridan et al. 2002). Specifically for
damselflies parasitized by water mites, the fac-
tors known to mediate sex-biases in parasitism
include sex differences in immunity, adult behav-
ior and/or phenology (Yourth et al. 2001; Lajeun-
esse et al. 2004; Robb & Forbes 2006). Sex-biases
in parasitism can also be generated as an effect of
body size dimorphism between sexes. Here the
larger sex (or larger individual, more generally)
may harbor a greater intensity of parasitism (i.e.,
number of parasites per host) because a larger
size may represent a greater resource for more in-
dividual parasites (Bush et al. 2001).
In addition to sex and body size, fluctuating
asymmetry has also been linked to parasitism for
many animals including damselflies (c.f. Bonn et
al. 1996). Field variation in ectoparasitism may
be associated with damselfly symmetry, should
the stressful conditions during larval growth (e.g.,
competition, predation, and/or parasitism) that
influence the symmetric development of wings,
also affect the susceptibility to, or recruitment of
parasites (Leung & Forbes 1996). However, wing-
cell asymmetry of adult damselflies is only a
coarse surrogate of physical condition during lar-
val-to-adult emergence and not adult condition
(which is difficult to assess for insects). However
despite this shortcoming, wing cell asymmetry
may explain some variation of field parasitism
given that many species of water mite are first
phoretic on damselfly larvae and then parasitize
hosts shortly after adult emergence (Smith 1988).

MATERIALS AND METHODS

I surveyed all damselflies (Odonata: Zygop-
tera) at 3 sites near the Archbold Biological Sta-
tion (ABS; Lake Placid, FL) for 10 days (18-III-
2006 to 30-111-2006). Two of these sites were
ephemeral ponds located near ABS, and a third
was located on the shore of Lake Annie. These
sites have distinct vegetation types (see below),
differ in size, but were primarily fields dominated
by grasses or shrubby hypericum (Hypericum
spp.). The edges of these fields were mostly domi-
nated by palmettos (Serenoa spp.). The vegetation
of the field surrounding my larger seasonal pond
(ca. 1500 m2; hereafter large pond, N 2710'50.7",
W 81' 21'10.8") was mostly hypericum with small
patches of redroot (Lachnanthes caroliniana),
gallery (Ilex glabra), fetterbush (Lyonia lucida)
and several pine snags (Pinus spp.). The edge
habitat surrounding this field had scrubby oak
species (Quercus inopia and Q. germinata). The
second seasonal pond (ca. 200 m2; N 2711'14.1",


W 81o21'27.1") had mostly hypericum plants, and
had an edge habitat composed of palmettos and
maidencane (Panicum hemitomon), but did not
have any pine snags. Finally, the lake site (N
27012'39.3", W 81020'58.7") had mostly southern
ridge saw-hill flora (see description in Abraham-
son et al. 1984).
At the two ponds, I sampled damselflies in the
morning (ca. 10:30 am) and afternoon (ca. 4:00
pm), and rotated these sampling times daily (from
afternoon to morning) to control for differences in
activity between damselfly species. My survey at
Lake Annie occurred daily at 1:00 pm. During
surveys, I collected all possible damselflies with a
sweep-net for 1 h. Rigorous sampling for a narrow
time-frame can provide information on the rela-
tive frequency and abundance of damselfly spe-
cies at a particular site. Captured damselflies
were processed at ABS.
Damselfly species were identified following
Westfall and May (1998) and were verified with
specimens from the ABS invertebrate museum
collected by M.J. Westfall. Individuals were sexed
and divided into two age classes: general (e.g.,
newly emerged with soft, translucent wings) or
adult (e.g., full coloration and rigid, dull wings).
Westfall and May (1996) found an age related
color-polymorphism of adult females inI. hastata,
with young females having an orange coloration
and older females with a darker brown to black
coloration. This distinction was used to separate
age classes for this species.
Symmetry of damselflies was estimated as the
absolute difference in the number of cells (c) be-
tween the nodus and ptrostigma on left (L) and
right (R) wings. The total asymmetry (a) was the
sum of the wing cell asymmetry between fore- and
hind-wings (f and h, respectively). Specifically,
asymmetry was calculated as follows:.

a = c + cI c

When a 0, damselflies were treated as asymmet-
ric in analyses.
Four estimates of body size were measured: (1)
wing-length (the distance between the nodus to
the pterostigma for the right forewing; 0.01 mm
Spi 31-44 caliper), (2) femur- and (3) tarsus-length
(the first right leg of each individual was im-
mersed in oil and had the femur and tarsus mea-
sured with use of a compound microscope at 10x; +
0.01 mm Olympus BX60F5). When the first right
leg was missing, I used the first left leg for mea-
surements. Finally, (4) wet weight was measured
for each damselfly immediately following surveys
( 0.0001 g; Mettler Toledo digital scale, model
AB104-5). These measures of body size were col-
lapsed into a single size estimate with Principle
Component Analyses (PCA; Zar 1984), but were
log10(x + 1) transformed prior to PCA analyses.


December 2007







Lajeunesse: Ectoparasitism ofDamselflies


Water mites are parasitic only during a single
stage in their life cycle, but Smith (1988) provides
more details on this stage or the stages outlined
below. After hatching from eggs, mite larvae are
motile, and in many water mite species (i.e., Ar-
renurus mites) larvae are phoretic on damselfly
larvae. When larval damselflies eclose from the
water, phoretic mites abandon the exuviae and
crawl back onto the newly emerged generall) dam-
selfly. At this stage, phoretic mites locate suitable
areas on the damselfly to attach and insert their
feeding tubes (i.e., ventral side of the thorax or ab-
domen). Several mite larvae can attach to a single
damselfly host, and at this stage they are para-
sitic and feed on the host. Once engorged, mites
drop off their hosts when hosts return to water to
reproduce. Finally, these water mites cycle
through additional larval stages until they reach
adulthood as aquatic predators.
In this study, larval water mites on each dam-
selfly were counted and grouped as either
phoretic (i.e., not attached with legs) or attached
(e.g., with feeding tube inserted, engorged and
without visible legs). Larval mites also were
grouped by their attachment-site on the dam-
selfly, either on the ventral side of the thorax or
abdomen.

RESULTS

Richness of Species Assemblages and Abundances
and Age Classes of Species

The most abundant pond species of damselfly
in late Mar was Ischnura hastata, followed by Ne-
halennia gracilis (Table 1), whereas Argia fumi-
pennis atra Gloyd was more common than
Ischnura ramburii (Selys) at the lake site. How-
ever, abundance estimates were biased against I.
ramburii captures because this species occupied
more offshore areas (i.e., among emergent vegeta-
tion at the lake's edge) than the field area sur-
veyed (ca. 2-20 m from the lake's edge). Of the 2
ponds, the larger had greater richness and abun-
dance of both damselfly species than compared to
the smaller pond-assuming an equal probability
of capture at both ponds (Table 1; I. hastata: x2 =
10.3, df = 1, P = 0.0013; N. gracilis: X2 = 6.7, df =
1, P = 0.0094). However, only a single general ofA.
fumipennis atra and I. ramburii were captured at
the large pond site. These 2 species are more com-
mon to permanent sources of water (Westfall and
May 1998), and it is unknown whether they had
emerged from this site or dispersed from a more
permanent site (e.g., Lake Annie).
Damselfly species captured at pond sites dif-
fered in number of adult captures: 75% of 16 N.
gracilis and 37% of 114 I. hastata were adults
(species contrast: x2 = 8.4, df= [1,128], P = 0.0037).
However, the frequency of adult capture did not
differ among the smaller and larger ponds for


both damselfly species (I. hastata: x2 = 0.1, df =
[1,112], P = 0.76; N. gracilis: X2 = 0.1, df = [1,14],
P = 0.72). For the 2 lake species, 12% of 53A. fumi-
pennis atra and all I. ramburii were adult (n = 7).

Water Mite Groups

In total, 92 larval mites were found (24
phoretic and 68 attached), and those attached
could be divided into 3 groups: orange and spotted
thoracic mites (e.g., common on the ventral side of
the mesepimeron, metepimeron and tergites 1
and 2), dark-red abdominal mites (e.g., generally
on 4t to 7th sternites), and a bright-red mite on the
upper posterior of thoracic mespimeron. By far
the most abundant were the orange thoracic
mites, with 61 attached individuals found on 15
damselflies. There also was over dispersion of tho-
racic mites, where 49% of attached mite larvae
were found on 3 damselfly individuals (Table 1).
The other 2 mite groups were uncommon and
found only on single individuals: 6 abdominal
mites on I. hastata from the small pond and a sin-
gle bright-red mite on N. gracilis from the large
pond.
Based on (a) the coloration of mites, (b) the
body sites to which they attached to hosts, (c) that
all phoretic mites were found on newly emerged
tenerals, and (d) that all mites were found on coe-
nagrionid damselflies, it is likely that these lar-
vae are 2 species of Arrenurus water mite (B.P.
Smith, Biol. Dept., Ithaca College, personal com-
munication 2006). Unfortunately, it is impossible
to assess the species of these mite larvae without
prior knowledge of the adult water mites that oc-
cupy this area (this information is unknown for
ABS), or without rearing larval mites on hosts un-
til they detach and reach adulthood (Botman et
al. 2002). Generally, only adult males are useful
for identifying species.
Finally, the single brightly colored mite found
on N. gracilis had attached to an atypical region
for odonates (see above); suggesting that this mite
is likely more often parasitic on a completely dif-
ferent insect order.

Water Mite Parasitism

The majority of mites were found on L. hastata,
where 12.2% of individuals had 82 mites (19
phoretic and 63 attached; Table 1). However, de-
spite the overall difference in abundance between
I. hastata and N. gracilis, both had similar preva-
lence (x2 = 0.1, df = [1, 128], P = 0.98) and inten-
sity of parasitism (Kruskal-Wallis x2 = 0.05, df =
1, P = 0.81). All phoretic mites were found on
newly emerged tenerals: 2 I. hastata with 2 and
17 phoretic mites, respectively, and 1 Y N. gracilis
with 5 phoretic mites. These 2 damselfly-mite
associations are novel and unreported for Central
Florida.







Florida Entomologist 90(4)


TABLE 1. FREQUENCY OF DAMSELFLY SPECIES AND PARASITISM BY WATER MITES OF 3 SITES NEAR THE ARCHBOLD BI-
OLOGICAL STATION. NUMBER OF DAMSELFLIES CAPTURED (n) AND THEIR RELATIVE FREQUENCY (F) ARE PRE-
SENTED, AS WELL AS THE PREVALENCE OF PARASITISM (P) WITH LOWER (LCI) AND UPPER (UCI) 95%
CONFIDENCE INTERVALS, AND MEDIAN INTENSITY (I) OF PARASITISM WITH INTER-QUARTILE (IQR) AND FULL
RANGE (FR). LCI AND UCI WERE CALCULATED FOLLOWING JAYNES' (1976) BAYESIAN METHOD. THIS
METHOD ESTIMATES UCI WHEN PREVALENCE IS ZERO, AND INDICATES THE LIKELIHOOD OF CAPTURING PAR-
ASITIZED INDIVIDUALS GIVEN THE CAPTURE FREQUENCY (E.G., SURVEY SAMPLE SIZE).

Parasitism estimates

Prevalence Intensity

Damselfly assemblage n F P LCI UCI I IQR FR

All sites
Argia fumipennis atra 53 0.28 0.000 0.000 0.054 -
Ischnura hastata 114 0.60 0.122 0.071 0.191 3.5 1.75-6.25 1-22
Ischnura ramburii 7 0.04 0.000 0.000 0.312 -
Nehalennia gracilis 16 0.08 0.125 0.038 0.364 5.0 1-9 1-9
Lake Annie
Argia fumipennis atra 52 0.90 0.000 0.000 0.055 -
Ischnura ramburii 6 0.10 0.000 0.000 0.348 -
Small pond
Ischnura hastata 40 0.93 0.100 0.041 0.231 4.0 1.75-16.75 1-21
Nehalennia gracilis 3 0.07 0.000 0.000 0.602 -
Large pond
Argia fumipennis atra 1 0.01 0.000 0.000 0.776 -
Ischnura hastata 74 0.83 0.135 0.075 0.232 2.5 1.75-6.25 1-22
Ischnura ramburii 1 0.01 0.000 0.000 0.776 -
Nehalennia gracilis 13 0.15 0.154 0.046 0.428 5.0 1-9 1-9


Damselflies from the large or small pond were
not more likely to have parasites (X2 = 0.56, df = [1,
128], P = 0.45), and there was no site-by-species in-
teraction in likelihood of being parasitized (Wald
X2 = 0.01, df = [1, 128], P = 0.93). Thus, parasitism
frequency was not dependent on the site from
which damselflies emerged in whole, or individu-
ally by species (I. hastata: X2 = 0.3, df = [1,112], P
= 0.58; N. gracilis: 2 = 0.89, df = [1,14], P = 0.34).
I did not find any larval mites onA. fumipennis
atra or I. ramburii at my lake site (Table 1). Thus,
all following analyses are limited to I. hastata and
N. gracilis because these were the only species
parasitized by my water mites. I also combine
data from the adult and general damselflies in all
following analyses, because (a) Arrenurus mites
are only recruited by damselflies during larval
emergence (thus the prevalence and intensity of
parasitism does not accumulate among age
classes), and (b) the frequency of parasitism
among age classes did not differ (I. hastata: X2 =
0.2, df = [1,112], P = 0.62; N. gracilis: X2 = 0.67, df
= [1,14], P = 0.41).

Sex and Parasitism

Males were as common as females in both spe-
cies of damselfly: I. hastata had 59 Y and 55 6
(assuming equal sex ratio: X2 = 0.14, df = 1, P =


0.17) andN. gracilis had 7 Y and 9 6 (X2 = 0.25, df
= 1, P = 0.62). However, no one sex was more
likely to be parasitized (Fig. la; hastata had 7 Y
and 7 6 parasitized: x2 = 0.02, df= 1, P = 0.888;N.
gracilis had 1 Y and 1 6 parasitized: X2 = 0.04, df
= 1, P = 0.85), or have a greater intensity of larval
water mites (I. hastata: Kruskal-Wallis X2 = 1.51,
df = 1, P = 0.22). Too few individuals ofN. gracilis
were parasitized to test whether intensity of wa-
ter mites differed between sexes.
The large and small pond also did not have dif-
ferent sex ratios of both species (I. hastata: X2 =
0.45, df = [1,111], P = 0.5; N. gracilis: X2 = 0.17,
df= [1,14], P = 0.68), and did not contribute to
frequencies of parasitism among sexes (site-by-
sex interaction for I. hastata: Wald X2 = 0.06, df =
1, P = 0.81; and N. gracilis: Wald 2 < 0.01, df = 1,
P = 0.99).

Body Size and Parasitism
Body size measures (i.e., wing-, femur-, and
tarsus-length, and weight) were collapsed into a
single estimate of size (hereafter PC1) with PCA,
and calculated separately for each species. The
PC1 for I. hastata explained 59.3% of the varia-
tion among body size traits (eigenvalue = 2.37),
and 41% for N. gracilis (eigenvalue = 1.64). Both
damselfly species were overall similar in PC1 (t <


December 2007










Lajeunesse: Ectoparasitism ofDamselflies


0.01, df = 94, P = 1.0), however only I. hastata was
sexually dimorphic with females being larger
than males (Fig. Ib; I. hastata: t = 9.01, df = 83, P
< 0.001;N. gracilis: t = 0.25, df = 9, P = 0.81).
Body size did not relate with prevalence of par-
asitism in I. hastata (log-likelihood test, X2 = 0.08,
df= 1, P = 0.77), intensity of infection (r2 = 0.07, F
= 0.79, df = 10, P = 0.39), or relate to size differ-
ences between sexes (e.g., no sex-by-size interac-
tion: Wald x2 = 0.31, df = 1, P = 0.58). Again, too
few parasitized individuals were available to as-
sess whether water mite parasitism related to
body size or sex for N. gracilis.
In general, it does not appear that body size re-
lated to patterns of parasitism by larval mites. Fi-
nally, there were no differences in body sizes
among damselflies emerging from the small or
large pond (all damselflies: t = 0.74, df = 94, P =
0.74; I. hastata: t = -0.45, df = 83, P = 0.63;N. gra-
cilis: t = 0.56, df= 9, P = 0.59).

Symmetry and Parasitism

Symmetry of damselflies did not relate to fre-
quencies of parasitism observed in the field.
Asymmetric damselflies were not more likely to
be parasitized than symmetric individuals for all
damselflies (X2 = 3.09, df= [1,129], P = 0.08) or in-
dividually by species (I. hastata: X2 = 1.83, df =
[1,111], P = 0.17;N. gracilis: X2 = 2.52, df = [1,14],
P = 0.11). Asymmetric damselflies also did not
have a greater intensity of parasitism than sym-
metric individuals (all damselflies: Kruskal-Wal-
lis x2 = 2.89, df = 1, P = 0.09; I. hastata: Kruskal-
Wallis x2 = 0.01, df = 1, P = 1.0; N. gracilis:
Kruskal-Wallis X2 = 1.65, df = 1, P = 0.2), and
there was no site-by-symmetry interaction in pre-
dicting patterns of parasitism (Wald X2 = 0.01, df
= 1, P = 0.94). In addition, there was no variation
in symmetry among the damselflies captured at
the 2 ponds (X2 = 1.83, df = [1,111], P = 0.175). Fi-
nally, the sexes of each damselfly species were
equally symmetric (Fig. lc;L. hastata: X2 = 0.02, df
= [1,111], P = 0.89; N. gracilis: X2 = 0.01, df =
[1,14], P = 0.95), and there was no relationship be-
tween symmetry and body size (I. hastata:
Kruskal-Wallis x2 = 0.21, df = 1, P = 0.65; N. gra-
cilis: Kruskal-Wallis x2 = 1.2, df= 1, P = 0.27).


DISCUSSION


Patterns of field parasitism have been reported
for several odonate-water mite associations in
northeastern North America (e.g., Forbes et al.
2002), but very few studies exist for southern
North America. My survey found 2 novel host as-
sociations for Arrenurus water mites in Central
Florida, where 12.2% of I. hastata and 12.5% of
N. gracilis damselflies had larval mites. Should
my survey have been seasonal, I likely would
have found other novel hosts-given that many


Fig. 1. Contrasts between females (f) and males (m)
of pond species of damselfly in prevalence of water mite
parasitism (a), body size (b) and the frequency of sym-
metry (c) defined as the ratio of symmetric vs. asymmet-
ric individuals in wing cell number. Females and males
are symmetric if 95% CI overlap with 0.5 (dashed bar).
Sample sizes are as follows: Ischnura hastata 59 2/55
and for Nehalennia gracilis 7 /96. Asymmetric 95%
confidence intervals (CI) around proportions (e.g., prev-
alence and frequency of symmetry) were calculated fol-
lowing Jaynes (1976).


1.00-


0.75-


0.50 -


0.25-


0.00 -

1.5-

1.0 -

0.5-

0.0 -

-0.5-

-1.0 -

-1.5-


f m


f m


1.00-


a 0.75-
-H

E
E 0.50-
t


,-
0



uC
000-
0.00-


I-- - -


f m


f m


" I t


m
i. hastata


i I
f m
N. gracilis











additional species of damselfly (and dragonfly)
emerge at different periods throughout the year
(Westfall & May 1996).
Field variation in parasitism by water mites
can occur because of differences in host exposure,
immunity, phenology, and spatial constraints.
However, I could not detect differences in preva-
lence and intensity of parasitism between the two
infected species of damselfly-despite being dis-
tantly related and differing significantly in abun-
dance. Nor could I detect the contribution of sex,
body size and asymmetry of hosts on observed
field patterns in water mite parasitism. At this
time of season, it appears that larval water mites
are distributed randomly among species and
classes (e.g., sex) of damselfly hosts.
These findings corroborate other studies that
find no trend in parasitism among sexes for other
insects (Sheridan et al. 2002), or more generally,
no effect due to body size and asymmetry (M0ller
& Thornhill 1998; Forbes et al. 2004). However,
my ability to detect such patterns, should they
have existed, was poor due to the low prevalence
of parasitism and overdispersion of mites on few
individual hosts-a common problem to field sur-
veys of parasites (Bush et al. 2001). A large num-
ber of parasitized individuals, with high repre-
sentation among various host classes (e.g., sex,
symmetry, etc.), would be required to confidently
assess whether such mediating effects exist. This
is perhaps best achieved through the experimen-
tal manipulation of mite inflections (Lajeunesse
et al. 2004). However in the field, large sample
sizes of parasitized individuals may not be attain-
able for damselfly species that are not abundant
locally, or uncommon at certain times of the sea-
son, as seen for N. gracilis damselflies, where few
were captured, and even fewer were parasitized.
I also did not find any dead larval mites and
was unable to assess whether damselflies had lost
mite larvae prior to surveys. Attached dead mites
can indicate a successful immune response by
damselflies (Yourth et al. 2001). However, a sig-
nificant portion of the surveyed damselflies were
general (63% of L. hastata and 25% of N. gracilis)
and were perhaps too young to yet resist larval
mite parasitism. In addition, there is the possibil-
ity that many of the surveyed damselflies had al-
ready lost mites before sampling. However, no
mite scars were found on any individual (scars
are left when mite larvae detach from hosts;
Smith 1988), nor were there any large, fully en-
gorged mite larvae found on any host. These ob-
servations indicate that the surveyed damselflies
could not have lost mite larvae because mites had
fully matured and detached from hosts to com-
plete their life cycle (Smith 1988). To avoid these
issues, future surveys should examine immunity
to mite larvae feeding tubes (Yourth et al. 2001) or
conduct a seasonal mark-recapture study to de-
termine mite mortality (Lajeunesse et al. 2004).


December 2007


Odonate-water mite associations are a model
system to test key hypotheses on life-history
trade-offs (Mitchell 1967), but the lack of field
data on patterns of host use has hampered
progress on understanding the ecology and evolu-
tion of these mites and their hosts (Bohonak 1999;
Rolf 2002; Forbes et al. 2002; Lajeunesse et al.
2004). Additional field surveys, broadly sampling
a diversity of habitats, can help address these is-
sues and will likely generate novel, unreported
host associations for parasitic water mites.


ACKNOWLEDGMENTS

I thank Jed Sparks, Bruce Smith, Richard Root, Pe-
ter Marks, Michael Stastny, and Stuart Campbell for
helpful discussion on this project, and the Archbold Bi-
ological Station for logistic support. The Department of
Ecology and Evolutionary Biology at Cornell funded
this study as part of a graduate course in Florida field
ecology.

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Florida Entomologist 90(4)


December 2007


EFFECT OF ADULT DIET ON LONGEVITY OF STERILE
MEDITERRANEAN FRUIT FLIES (DIPTERA: TEPHRITIDAE)

JAMES D. BARRY'2, SUSAN B. OPP', JULIA DRAGOLOVICH3 AND JOSEPH G. MORSE1
'Department of Entomology, University of California, Riverside, CA 92521

2Current Address: DuPont Crop Protection, Stine-Haskell Research Center, Newark, DE 19714

'Department of Biological Sciences, California State University, East Bay, CA 94542

ABSTRACT

Fly longevity is critical to sterile release programs for Mediterranean fruit flies (medflies)
because the longer sterile flies are present after a release, the greater the probability of mat-
ing. Current release programs provide sterile, adult medflies with sucrose in an agar matrix
for 2-3 d before release. We used cages to compare the effects of different diets on the longev-
ity of medfly, Ceratitis capitata (Wiedemann). A diet of dry hydrolyzed yeast + sucrose sup-
plied during the pre-release interval did not significantly affect field survivorship of medfly
adult males relative to the standard sucrose diet. Post-release diets, simulating nitrogen
and sugar sources that released medflies may find after release, had significant effects on
medfly survivorship. Hydrolyzed yeast + sucrose resulted in the highest medfly survivor-
ship, followed by sucrose, and then water alone. Finally, diets containing hydrolyzed yeast
were not found to have significant amounts of protein and thus are more likely nitrogen or
amino acid sources for flies, rather than sources of protein.

Key Words: Ceratitis capitata, sterile insect technique, longevity, hydrolyzed yeast, protein

RESUME

La longevidad de las moscas es critical en un program de t6cnicas de insecto est6ril para la
mosca mediterranea de la fruta porque el mas tiempo que las moscas est6riles esten presents
despu6s de ser liberadas, mayor es la probabilidad del apareamiento. Los programs actuales
de liberaci6n de adults est6riles de la mosca mediterranea los proven con sucrosa en una
matriz de agar por 2-3 dias antes de su liberaci6n. Nosotros usamos jaulas para comparar los
efectos de dietas diferentes sobre la longevidad de la mosca mediterranea Ceratitis capitata
(Wiedemann). Una dieta de levadura seca hidrolizada + sucrosa suplida durante el intervalo
pre-liberaci6n no tuvo un efecto significant sobre la sobrevivencia de los machos adults de
la mosca mediterranea en el campo en relaci6n con la dieta estandar de sucrosa. Las dietas de
pos-liberaci6n, que simulan las fuentes de nitr6geno y azucar que la mosca mediterranea li-
berada y que pueden ser encountradas despu6s de ser liberadas, tuvo efectos significantes so-
bre la sobrevivencia de las moscas mediterranea. La levadura seca hidrolizada con sucrosa
result en una sobrevivencia lo mas alta de mosca mediterranea, seguido en orden por la dieta
de sucrosa y despu6s solo el agua. Por iltimo, las dietas que contienen levadura hidrolizada
no tuvieron cantidades significantes de protein y por lo que probablemente son fuentes de ni-
tr6geno o de aminoicidos para las moscas, al contrario de ser fuentes de protein.


The Mediterranean fruit fly (medfly), Ceratitis
capitata (Wiedemann), is a multivoltine, poly-
phagous insect pest that could have a devastating
economic impact if it were to become established
in California or Florida (Robinson et al. 1986; Liq-
uido et al. 1991; Metcalf 1994). Presently, the
sterile insect technique (SIT) is used in these
states as a preventative measure to reduce the po-
tential for Mediterranean fruit fly colonization.
For wild males, reproductive success is the end re-
sult of the following steps: joining a lek, phero-
mone emission, courtship behavior, copulation,
sperm transfer and storage, fertilization of eggs,
and preventing/delaying female re-mating (Yuval
et al. 2002). The goal of SIT is for released sterile


males to mate with any introduced wild females,
resulting in the production of infertile eggs
(Knipling 1955). Sterile male longevity after re-
lease and competitiveness with wild males are
two factors that can impact the effectiveness of a
SIT program.
Protein and carbohydrates are necessary di-
etary components for optimum medfly develop-
ment and fecundity and could impact the effec-
tiveness of sterile males. In the field, tephritid
fruit flies have a larval diet that is often protein-
deficient; whereas the larval diet of laboratory-
reared, sterile flies is protein-rich (Cayol 2000).
Cangussu and Zucoloto (1997) found that larvae
consuming a protein-rich diet followed by a pro-







Barry et al.: Tephritid Adult Diet and Longevity


tein-poor adult diet resulted in females capable of
producing more eggs than females having a pro-
tein-poor larval diet followed by a protein-rich
adult diet. For adults, field sources of carbohy-
drates include injured fruits and honeydew, and
proteins are commonly found in bird feces and
rotting fruit colonized by bacteria (Hendrichs &
Hendrichs 1990; Hendrichs et al. 1991).
Many studies have characterized the effects of
protein-rich and protein-deficient adult diets (i.e.,
diets containing protein + sugar vs. sugar only, re-
spectively) on wild and laboratory medflies (Yuval
et al. 2002; Yuval et al. 2007). Laboratory strains
of medfly fed protein-deficient adult diets some-
times showed an overall reduction in longevity
(Cangussu & Zucoloto 1997; Niyazi et al. 2004), or
had a comparable longevity to protein-fed flies
(Shelly & Kennelly 2002; Shelly & McInnis 2003).
In contrast, Kaspi & Yuval (2000) found 24 h of
starvation resulted in greater mortality for pro-
tein-fed than protein-starved laboratory strain
flies. Protein-fed wild flies and laboratory males
were more likely to emit pheromones in leks than
protein-starved flies (Papadopoulos et al. 1998;
Kaspi et al. 2000; Kaspi and Yuval 2000). Yuval et
al. (1998) found that field-collected lekking wild
males contained higher amounts of protein and
sugar compared to resting males. From these
studies it is clear that there are some benefits of
protein in the adult diet, however the impact
would often depend on field conditions (e.g., like-
lihood of 24 h starvation).
Mating success of medflies can also be im-
pacted by diet. Laboratory strain males provided
with protein were 1.4-2x more likely to mate than
protein-starved males (Taylor & Yuval 1999;
Kaspi et al. 2000; Kaspi & Yuval 2000; Shelly et al.
2002), but in other studies no effects of diet were
found (Shelly & Kennelly 2002; Shelly & McInnis
2003). Niyazi et al. (2004) reported that laboratory
flies provided with a pro-biotic, yeast-enhanced,
sugar diet had a significant mating advantage
over protein-starved flies fed either a pro-biotic or
non-probiotic diet in the laboratory, but there was
no significant effect of diet between these 2 treat-
ments in field cage studies. In one study with wild
flies, the inclusion of protein in the adult diet in-
creased mating success over protein-starved coun-
terparts (Shelly & Kennelly 2002).
In part due to the high mortality of sterile
medflies in the field, releases of sterile medflies
occur twice a week in the Preventative Release
Program in California (Barry et al. 2002). Sterile
adult medflies are provided with sucrose in agar
(1 M sucrose, no protein) 2-3 d before release in
SIT programs such as that in California. It is pos-
sible that sterile males may not find needed nutri-
ents following field release to allow for adequate
longevity and reproductive development so as to
mate with wild females. However, in a study with
field enclosures, Maor et al. (2004) reported that


both protein-starved and protein-fed flies were
able to successfully forage for protein and sugar
when it was available. Improving the pre-release
adult diet is one possible method of increasing
SIT fly longevity in the field.
Kaspi and Yuval (2000) reported that the inclu-
sion of protein hydrolysate in pre-release diets in-
creased mating success while reducing longevity.
In California, the inclusion of hydrolyzed yeast
into the agar matrix of SIT medflies resulted in a
high bacterial population forming on the diet me-
dia, and as a result, the concentration of an anti-
microbial compound (methyl paraben, CHO,)
was increased, which subsequently lowered fly
survivorship (R. V. Dowell, Calif. Dept. of Food &
Agric., pers. comm.).
In this paper we investigated the effects of pre-
and post-release diets on sterile medfly longevity
in small field cages (without plants), and deter-
mined the levels of protein that were provided by
different diets containing hydrolyzed yeast.

MATERIALS AND METHODS

Source of Insects

Sterile medfly puparia (Vienna-7, laboratory
strain) were obtained in Oct of 2002, from the
USDA/CDFA Cooperative Mediterranean fruit fly
Preventative Release Program. The medflies for
our tests were reared at the California Depart-
ment of Food and Agriculture (CDFA) larval rear-
ing center in Waimanalo, Hawaii. Puparia were
dyed with Day-Glo neon red dye (Day-Glo Color
Corp., Cleveland, OH), at 3 g of dye per liter of
medflies, using a mechanical mixer containing 10-
20 L of puparia (D. McInnis, USDA-ARS, pers.
comm.). The puparia were tumbled gently for a
few minutes to dye all of the flies prior to irradia-
tion at 8 d of puparial age. Two days prior to eclo-
sion, puparia were placed in hypoxia for 1-2 h and
irradiated with a dose of 145 Gy from a Cobalt60
pool-type irradiator. Puparia were subsequently
shipped to the David R. Rumsey Emergence and
Release Facility in Los Alamitos, California.

Pre-release and Post-release Diets

Survivorship studies were conducted in River-
side, CA with sterile medflies to simulate treat-
ment of adult male medflies used in the SIT release
program in California, which are held and fed for 2-
3 d before being released into the environment.
Two pre-release diet treatments were offered to
newly closed medflies for 48 h: (1) sucrose in agar,
which is the standard medfly diet used in the CA
preventative release program (94.98% sucrose,
4.95% agar, and 0.07% methyl paraben in 1 L of
distilled, sterile water per 181 g of dry matter)
(Niyazi et al. 2004), or (2) sucrose in agar (as in
diet #1), with the addition of a separate dry mix-







Florida Entomologist 90(4)


ture of hydrolyzed yeast + dry sucrose (1:3, wt:wt).
For the first 48 h, flies were held at the David R.
Rumsey Emergence and Release Facility. After 48
h, groups of 20-30 flies, each in a 0.5-L plastic con-
tainer were transferred in coolers by vehicle to
Riverside, CA (approximately 2.5 h in transit).
One group of flies was randomly selected to be
placed into each of 80 small field cages (without
plants; 30.5 x 30.5 x 30.5 cm3; Bioquip, Rancho
Dominguez, CA), located along a southwest-fac-
ing wall, underneath a cover (6 m x 3 m) to limit
exposure to the wind and sun. The following 4
post-release diet treatments were offered to flies:
(1) 10 g hydrolyzed yeast + dry sucrose (1:3 by
weight) + water, (2) 10 g dry sucrose + water, (3)
water, or (4) nothing (i.e., no food or water pro-
vided). Water was provided in a 20-mL container
with a cotton wick and was changed every 4-5 d.
Dry hydrolyzed yeast and sucrose mixtures were
provided in plastic dishes (and were not mixed
with water). A total of 10 replicate cages with 20-
30 adult males per cage were used for each medfly
diet treatment (2 pre-release x 4 post-release
treatments).
Cages were checked daily for fly mortality, and,
after all flies had died, the average longevity of
flies was determined for each cage. At the cage
site, the average high temperature was 29.7C,
average low temperature was 12.7C, and average
percent relative humidity was 61%.

Statistical Analyses

One-way and two-way Analysis of Variance
(ANOVA) was used to compare square root-trans-
formed (I(x + 0.5)) data of the average medfly lon-
gevity per cage for pre-release and post-release
diets, with means of significant factors separated
by Fisher's Least Significant Difference (LSD)
test (Minitab, Inc. 1998).

Protein Analysis

Protein levels were determined by a modified
Bradford method developed by Bio-Rad (Bio-Rad
Protein Assay, catalog #500-0002) (Bradford 1976).


Hydrolyzed yeast (Fisher Scientific, BP1422-500)
was dissolved in de-ionized water at a concentra-
tion of 1mg yeast per 1 mL water solution. Seven
samples of this solution were prepared, each in
triplicate. Standard protein curves were estab-
lished with 3 concentrations of protein standard.
Bovine serum albumin (1.46 mg/mL) was diluted
with de-ionized water to concentrations of 10 pg/
mL, 5 pg/mL, and 2.5 pg/mL. Three replicates of
all samples and standards were assayed for pro-
tein content. An 800-mL aliquot of sample or
standard was placed in a clean test tube to which
200 mL of reagent was added (Bio-Rad Protein
Dye Reagent catalog #500-0006) and allowed to
sit for 5 minutes before reading in a spectropho-
tometer at 595 nm. A "blank" consisting of 800 pL
de-ionized water plus 200 pL reagent was tested.
The absorbance values for the 3 replicates of each
sample were averaged for the final value. The fi-
nal values were compared to the standard curve
to determine the amount of protein present in the
hydrolyzed yeast. Protein values are presented as
micrograms of protein per milligram of yeast.

RESULTS

Pre-release and Post-release Diets

Adult medfly longevity was significantly af-
fected by post-release diet, but was not affected by
pre-release diet or the interaction of pre- and
post-release diet (Table 1; Pre-release F = 3.52;
df = 1; P = 0.065; Post-release: F = 184.80; df = 3;
P < 0.0001; Interaction: F = 0.99; df = 3;P = 0.401;
Error: df = 72). When data for pre-release diets
were pooled, post-release diet significantly af-
fected adult medfly longevity (Fig. 1; F = 178.92;
df= 3, 76;P < 0.0001). Males that were offered the
post-release hydrolyzed yeast + sucrose + water
diet lived significantly longer than those offered
all 3 other post-release diets, and males provided
the sucrose + water diet were significantly longer
lived than flies offered water and flies offered
nothing. At the end of the trial, there was still ex-
cess hydrolyzed yeast and sucrose mixture left in
those cages where it was provided.


TABLE 1. LONGEVITY OF ADULT MALE MEDFLIES EXPOSED TO DIFFERENT POST-RELEASE DIET REGIMENS.

Longevity after 48 h
Pre-release diet (0-48 h) Post-release diet (>48 h) (mean days SE)

Sucrose+agar No food or water 3.1 + 0.08
Water 3.6 0.05
Sucrose + water 10.9+ 0.70
Sucrose/yeast hydrolysate + water 13.5 1.00
Sucrose + agar + yeast hydrolysate No food or water 3.2 0.05
Water 3.7 0.07
Sucrose + water 12.0 0.90
Sucrose/yeast hydrolysate + water 14.7 0.80


December 2007







Barry et al.: Tephritid Adult Diet and Longevity


1.0

0.9

0.8

' 0.7

R 0.6

3 0.5

S0.4
i 0.3

0.2

0.1

0.0


0 10 20 30 40

Days


Fig. 1. Adult, sterile male medfly survivorship with different post-release diets offered to flies after 48 h. Results
were pooled for both pre-release diets (yeast hydryolsate + sugar and sugar) that were offered to flies for the first
48 h. Day 0 was the day flies were first exposed to post-release diets.


Protein Analysis
Hydrolyzed yeast contained extremely low lev-
els of protein (mean = 3.15 pg protein/mg hydro-
lyzed yeast, SD = 0.28, n = 7). Therefore, the medfly
diet that consisted of hydrolyzed yeast + dry su-
crose (1:3, wt:wt) was approximately 0.08% protein.

DISCUSSION
The inclusion of a hydrolyzed yeast and su-
crose mixture in the post-release adult medfly
diet increased longevity in comparison to flies
provided with only sucrose and water. There was
no increase in the longevity of medflies provided
with hydrolyzed yeast and sucrose mixture prior
to transfer into cages (i.e., as a pre-release diet)
when post-release diets were not provided. This
finding leads us to suggest that incorporating this
mixture of hydrolyzed yeast and sucrose into the
standard pre-release diet of sterile flies would not
be beneficial, unless there were other advantages
(i.e., improved mating success) that would not be
conveyed to these sterile flies if they foraged and


obtained protein in the field. As Maor et al. (2004)
stated, released flies incapable of finding protein
and carbohydrates are likely to die regardless of
the pre-release diet. The length of time a fly is ex-
posed to pre-release conditions (i.e., 2-3 d) may
preclude detectable differences in subsequent fly
longevity in the field.
The impact of pre-release diets on mating suc-
cess may be less important than what food sources
foraging flies are able to find following release.
Kaspi and Yuval (2000) found that mating success
of flies offered a protein or sugar pre-release diet
for 4 d could be altered by 24 h of starvation (with
access to water only) and by offering an apple. For
example, sugar-fed males that were given 24 h ac-
cess to an apple had greater mating success than
protein-fed flies starved for 24 h. Likewise, sugar-
fed and protein-fed flies both provided with access
to apples had comparable mating success. These
findings demonstrate that the fate of pre-release
flies could vary considerably, largely based on
what flies find after they are released.
Mass-reared flies may not die more quickly
when fed a diet containing hydrolyzed yeast be-









Florida Entomologist 90(4)


cause nitrogen does not limit attainment of repro-
ductive maturity (Zucoloto 1992). Thus, because
mass-reared flies are fed a larval diet high in pro-
tein, adult diet is less important for reproductive
maturity than is the case with wild flies. Muller et
al. (1997) found that mortality decreased when
mass-reared adult medflies were fed a diet con-
taining hydrolyzed yeast. We similarly found a
decrease in mortality when cage-confined medfly
males were fed a post-teneral diet containing hy-
drolyzed yeast. Additional studies would be bene-
ficial to continue to tease apart the differences be-
tween mass-reared, sterile medflies, and wild
medflies in relation to diet.
Surprisingly, we found that hydrolyzed yeast is
not a significant source of protein, despite numer-
ous studies that refer to a fly diet containing hydro-
lyzed yeast or yeast hydrolysate as containing pro-
tein or being an important source of protein (Ja-
come et al. 1995; Muller et al. 1997; Shelly et al.
2002; Yuval et al. 2002). It has been found that
yeast hydrolysate contains 19 amino acids and pro-
vides an extrinsic source of nitrogen that is needed
by walnut husk flies to produce eggs (Tsiropoulos
1978). We were initially concerned that handling of
fly diet containing hydrolyzed yeast (e.g., exposure
to heat and/or to feeding by flies for extended peri-
ods of time), as might occur with cage studies or
with storage for use in large-scale SIT programs,
could alter protein content. But we found that the
protein content was so low to begin with that it is
not likely to be affected measurably by adverse
conditions. We note that the tephritid fly literature
is rife with confusing references to "protein diets"
when these diets, for the most part, appear to be
"nitrogen-rich" and have contained hydrolyzed
yeast, not protein. In the future, researchers
should be careful to indicate the identity and rela-
tive ratio of components in diets fed to flies.
Based on fly longevity in small field cages
(without plants), we did not find overwhelming ev-
idence to suggest that the inclusion of hydrolyzed
yeast in SIT pre-release diets is warranted with
medfly. Based on our studies and the findings of
others, we conclude that the ability of sterile flies
to locate and feed on nitrogen and carbohydrate
sources post-release will more significantly affect
sterile fly survivorship than pre-release diet.

ACKNOWLEDGMENTS

We thank the late David Rumsey, Kevin Hoffman,
Robert Dowell, Rufino Santos, Bob Garagliano, and many
other personnel from the USDA/CDFA Cooperative Med-
iterranean fruit fly Preventative Release Program; M.
Durbin for assistance with protein assays; and R. D.
Goeden, K. Hoffman, and T. M. Perring for reviewing an
earlier version of this manuscript. This research was
funded in part by the California Citrus Research Board,
by the Agricultural Research Initiative of the California
State University (grant to S. Opp and Carol Lauzon), and
by the California Department of Food & Agriculture.


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Florida Entomologist 90(4)


December 2007


PROGENY FITNESS OF THE MEALYBUG PARASITOID ANAGYRUS
SP. NOV. NR. SINOPE (HYMENOPTERA: ENCYRTIDAE) AS AFFECTED
BY BROOD SIZE, SEX RATIO, AND HOST QUALITY


JUANG-HORNG CHONG1' AND RONALD D. GETTING'
'Department of Entomology, University of Georgia, College of Agricultural and Environmental Sciences,
Griffin Campus, 1109 Experiment Street, Griffin, GA 30223 USA

'Present address: Clemson University, Pee Dee Research & Education Center,
2200 Pocket Road, Florence, SC 29506 USA

ABSTRACT

Anagyrus sp. nov. nr. sinope Noyes & Menezes (Hymenoptera: Encyrtidae) is a gregarious,
koinobiont parasitoid of the Madeira mealybug, Phenacoccus madeirensis Green (Hemi-
ptera: Pseudococcidae). We investigated effects of larval competition on development, sur-
vival, and progeny body size of the parasitoid over a range of brood sizes and sex ratios. We
recorded the brood size, sex ratio, and fitness parameters of parasitoids developing from sec-
ond instars and adult P. madeirensis. Adult mealybugs as hosts produce larger brood and
progeny body sizes and more female progeny. Second instar P. madeirensis continued to de-
velop after parasitism, and mummies achieved a more advanced developmental stage yield-
ing parasitoid cohorts of larger brood and body sizes, female-biased sex ratio, and longer
developmental time than mummies formed from younger host instars. Overall, the sexual
composition within a brood had little or no effect on the development, survival and body size
of the parasitoid progeny developing from second instar and adult mealybugs. Larval com-
petition among brood-mates developing in adult mealybugs caused a reduction in the prog-
eny tibial length with increasing brood sizes. In contrast, parasitoids developing from
second instar mealybugs had a longer developmental time and little (although positive) or
no response in the progeny tibial length to increasing brood size. We hypothesize that the
continuous development of second instar mealybugs allows the hosts to accumulate addi-
tional resources, thus, reducing competition among parasitoid brood mates for limited re-
sources. The continuous development of hosts of younger developmental stages after
parasitism may have the potential to alter the direction and intensity of larval competition
of the koinobiont gregarious parasitoid.

Key Words: growth potential, host stage preference, larval competition, Phenacoccus madei-
rensis, sexual asymmetry

RESUME

Anagyrus sp. nov. nr. sinope Noyes & Menezes (Hymenoptera: Encyrtidae) es un parasi-
toide koinobionte y gregario de la cochilla de Madeira, Phenacoccus madeirensis Green (He-
miptera: Pseudococcidae). Nosotros investigamos los efectos de la competencia de las larvas
en el desarrollo, sobrevivencia y tamano del cuerpo de la progenie del parasitoide sobre el
rango en el numero de progenie y la raz6n sexual. Nosotros registramos los parametros en
el numero de la progenie de cada grupo, la raz6n sexual, y el estado fisico de los parasitoides
desarrollados sobre el segundo estadio inmaduro y adults de P. madeirensis. Las cochinillas
adults como hospederas produce grupos de progenies mas numerosas, individuos mas
grandes y con una mayor proporci6n de hembras. El segundo estadio inmaduro de P. madei-
rensis continue su desarrollo despu6s de estar parasitados y las momias lograron tener una
etapa de desarrollo mas avanzada, resultando en cohortes de parasitoides con grupos de pro-
genie con mas individuos de cuerpos mas grandes, con una mayor proporci6n de hembras a
machos y con el tiempo de desarrollo mas largo que las momias formadas de los estadios mas
j6venes de hospederos. Sobre todo, la composici6n sexual entire el grupo de progenie tuvo
poco o no efecto sobre el desarrollo, la sobrevivencia, y el tamano de cuerpo de la progenie del
parasitoide que se desarrollaron de cochinillas de segundo estadio y adulto. La competici6n
larval entire las parejas de grupo de progenie desarrolladas en cochinillas adults causo una
reducci6n en la longitud de la tibia de la progenie con un aumento en el numero de progenie
por grupo. Por contrast, los parasitoides que se desarrollaron sobre el segundo estadio de
la cochinilla tuvo un period de desarrollo mas largo y una respuesta ligera (aunque posi-
tiva) o ninguna respuesta en la longitud de la tibia de la progenie en relaci6n al aumento en
el numero de progenie por grupo. Presentamos una hip6tesis de que el desarrollo continue
del segundo estadio de las cochinillas permit que el hospedero acumule recursos adiciona-
les y reduzca la competici6n entire las parejas en el grupo de progenie del parasitoide para







Chong & Oetting: Larval Competition Affected Parasitoid Fitness


los recursos limitados. El desarrollo continue de los hospederos en los estadios menos desa-
rrollados despues de estar parasitadas puede tener el potential para alterar la direcci6n y la
intensidad de la competici6n larval de este parasitoide koinobionte y gregario.


A foraging gregarious parasitoid determines
the number and the sex ratio of eggs deposited in
a host, based on the host's age or quality, and the
parasitoid's previous foraging experience and
physiological conditions (Godfray 1994; van Al-
phen & Jervis 1996). The decisions on the brood
size (i.e., the number of progeny emerged from
each mummy) and the sex ratio (proportion of
male progeny) within a single host have signifi-
cant influence on the intensity of competition
among brood mates, and subsequently, on the fit-
ness of the progeny.
The influence of brood size on parasitoids'
growth and development has often been explored
on the basis of competition among brood mates for
limited resources (Smith & Fretwell 1974; God-
fray 1994). There are trade-offs between brood
size and the fitness of parasitoids. Progeny which
emerge from a larger brood suffer from a smaller
body size (Bernal et al. 1999; Mayhew & van Al-
phen 1999; Fidgen et al. 2000), a shortened devel-
opmental time (Stapel et al. 1997; Harvey et al.
1998), and a reduced larval survival rate (Naka-
mura 1995; Allen & Hunt 2001; Milonas 2005).
Adult body size is positively correlated to fitness
of the parasitoids, which is measured by the num-
ber of progeny that can be produced by each fe-
male offspring and the number of females that
can be sired by each male offspring (Ode et al.
1996; Sagarra et al. 2001). Larval competition re-
duces progeny body size, and thus, the fitness of
the parents and offspring. An increase in larval
competition has a greater impact on the fitness of
female offspring, because smaller female parasi-
toids can produce fewer eggs than larger females,
while small males can still produce many sperm
for fertilization (Godfray 1994).
The sexual composition of a brood also had a
significant influence on the fitness of gregarious
parasitoids (Godfray 1986; Ode et al. 1996; West et
al. 2001). Godfray (1986) suggested that the differ-
ential ability of female and male parasitoid larvae
to compete for limited resources leads to sexual
asymmetry in larval competition, which is one of
the causes of variations in progeny sex ratio and
brood size. Larvae of the less competitive sex will
suffer to a greater extent in their fitness from the
competition with their more competitive brood
mates. When the female parasitoid larvae out-
numbered the male larvae, the body size of male
progeny was reduced with an increasing propor-
tion of females (Ode et al. 1996; Fidgen et al. 2000).
In this study, we investigated the developmen-
tal response ofAnagyrus sp. nov. nr. sinope Noyes
& Menezes (Hymenoptera: Encyrtidae) to varia-
tions in brood size, sex ratio, and host quality.


Anagyrus sp. nov. nr. sinope is a gregarious koino-
biont parasitoid of the Madeira mealybug, Phen-
acoccus madeirensis Green (Hemiptera: Pseudo-
coccidae), a serious pest of greenhouse ornamen-
tal production. Anagyrus sp. nov. nr. sinope pre-
ferred third instar immature and pre-
reproductive adult female P. madeirensis for ovi-
position and progeny development (Chong & Oet-
ting 2006a). The parasitoids that developed in
these preferred host stages had a shorter develop-
mental time, a lower mortality rate, a higher pro-
portion of females, and larger brood and body
sizes. First and second instars were considered
the least preferred and of the least suitable for
parasitoid growth and development. However,
younger nymphs that had continued development
to become older nymphs before mummification
produced a similar progeny number and sex ratio
as those mealybugs that were parasitized as
adults. Chong & Oetting (2006a) did not investi-
gate the influence of brood size and sex allocation
patterns on the fitness ofAnagyrus sp. nov. nr. si-
nope. We conducted this study to detect the occur-
rence of larval competition and sexual asymmetry
in larval competition among the brood mates of
Anagyrus sp. nov. nr. sinope, and their conse-
quences to the parasitoid's fitness.

MATERIALS AND METHODS

An Anagyrus sp. nov. nr. sinope colony was es-
tablished in 2002 at the University of Georgia,
Griffin Campus, Griffin, GA, USA, with individu-
als collected from a greenhouse colony ofAnagyrus
loecki Noyes & Menezes (Hymenoptera: En-
cyrtidae), which is another biological control can-
didate agent of P madeirensis. The greenhouse col-
ony ofA. loecki at the Griffin Campus was estab-
lished in 2000 with individuals collected from a
colony maintained at the University of Florida,
Mid-Florida Research and Education Center, Apo-
pka, FL. Anagyrus sp. nov. nr. sinope was not de-
tected in the original colony ofA. loecki in Florida,
leading us to believe that Anagyrus sp. nov. nr.
sinope originated as a local contamination oftheA.
loecki colony established in Griffin, GA (Chong &
Getting 2006b). Phenacoccus madeirensis reared
on sprouted russet potatoes (Solanum tuberosum
L., Solanaceae) were provided as hosts in the labo-
ratory colony. The mummies were collected from
the laboratory colony, isolated individually in gela-
tin capsules, and held at a constant temperature of
25 + 1C in an environmental chamber until adult
eclosion. After eclosion, each female parasitoid
was paired with two males and isolated for 48 h in
a glass vial supplied with a streak of honey solu-







Florida Entomologist 90(4)


tion as a source of moisture and carbohydrates. No
mealybugs were provided in the glass vials; thus,
the parasitoids had no oviposition experience at
the beginning of the experiment.
Phenacoccus madeirensis was reared on
sprouted russet potatoes in an insectary at the
Griffin Campus. Second instars with a body
length of 1.0-1.3 mm and pre-reproductive adult
females of 2 to 3 mm were collected for the exper-
iment. The experiment was conducted as a no-
choice test where mealybugs of a single develop-
mental stage were offered to the parasitoids at a
time. Ten mealybugs of one of the two develop-
mental stages were transferred onto an excised
coleus leaf [Solenostemon scutellarioides (L.)
Codd., Lamiaceae] and were allowed to settle over
a 16-h period. The coleus leaf was kept fresh by
inserting its petiole, through a hole drilled at the
bottom of a petri dish, into a cup of water.
A single parasitoid was released from the vial
and allowed to forage in a petri dish containing a
cohort of 10 mealybugs for 24 h at 25 + 1C, 90 +
2% relative humidity and 14:10 (L:D) h photo-
period. The parasitoid was removed after 24 h and
the mealybug cohort was incubated in the envi-
ronmental chamber. The mealybug cohort was ex-
amined every 5 d and the mummies were col-
lected. The collected mummies were isolated in
individual gelatin capsules and incubated in the
environmental chamber at 25 1C until adult
parasitoid eclosion. The developmental stage of
the mealybugs at the time of mummification, the
developmental duration from egg deposition to
adult eclosion of the parasitoids, and the progeny
number and sex ratio (percentages of males) were
recorded for each mummy. All mummies were dis-
sected at the end of the experiment to verify the
survival of immature parasitoids. All adult para-
sitoids were collected and preserved in 70% etha-
nol within 6 h of eclosion, and their left hind tibial
length was measured with a micrometer under
dissecting microscopes. Hind tibial length was
used as a surrogate for the body length and the
fitness of the parasitoid (Sagarra et al. 2001;
Chong & Oetting 2006a). Thirty seven and 46 rep-
licates were prepared for the adult and second in-
stars, respectively.
We first analyzed the distributions of brood
sizes and sex ratios ofAnagyrus sp. nov. nr. sinope
reared from the 2 host stages by subjecting the
frequency of mummies that yielded a specific
brood size or percentage of males to the analysis
of variance (ANOVA) after an arcsine-transfor-
mation (PROC GLM; SAS Institute 1999). The
mean frequencies of brood size and sex ratio from
each host stage were separated by the Tukey's
honestly significant difference (HSD) test (PROC
GLM; SAS Institute 1999). The influence of host
stages at the time of mummification on the re-
corded fitness parameters also was analyzed with
ANOVA and the means separated with Tukey's


HSD test. The relationship between brood size
and sex ratio, and the respective relationships of
brood size or sex ratio with the developmental
time (female and male combined), survival rate,
and female and male tibial lengths of parasitoids
developed from the 2 host stages were elucidated
with linear regression analyses (PROC REG; SAS
Institute 1999).

RESULTS

Parasitized second instar and pre-reproductive
adult P madeirensis continued to develop after
parasitism. Fifty two percent of the parasitized
pre-reproductive adult mealybugs produced eggs
before mummification, while the rest remained as
pre-reproductive females (Table 1). Adult progeny
ofAnagyrus sp. nov. nr. sinope emerged from these
reproducing and non-reproducing adult mealybug
mummies were of similar brood size (about 3
parasitoids per mummy), sex ratio (30-36%
males), developmental time (about 16 days), sur-
vival rate (97-100%), and tibial length (averaged
0.37 mm for females and 0.30 mm for males).
The majority (46%) of the parasitized second
instar mealybugs mummified at the second in-
star, while 21 and 32% of these mealybugs contin-
ued to develop into third instar and adult mealy-
bugs, respectively, before mummification (Table
1). The hosts that were parasitized as second in-
stars but achieved adulthood before mummifica-
tion yielded significantly more female parasitoid
progeny and larger brood and body sizes than the
mealybugs mummified at second instar. With the
continuous development of second instar hosts af-
ter parasitism, the parasitoid's developmental
time from egg deposition to adult eclosion was sig-
nificantly lengthened from 16 d, when the hosts
were mummified at second instar, to 24 d, when
the hosts were mummified as adults. Parasitoids
which emerged from mealybug mummies that
had achieved adulthood (whether the mummies
were developing from second instars or pre-repro-
ductive adults), had similar sex ratios, brood and
body sizes, and developmental time.
Fifty eight percent of all mummies developing
from the second instar mealybugs produced only
one adult parasitoid from each mummy (Fig. 1A).
This was by far the largest single value (ANOVA:
F7, 3. = 21.32, P < 0.0001) and was observed pre-
dominantly in mummies of which the develop-
ment was arrested in the second instar by the par-
asitoids' development and mummification. The
mummies developing from the pre-reproductive
adult mealybugs yielded a range of brood sizes
from zero to 9 adult parasitoids per mummy, with
the largest brood sizes collected at roughly equal
proportion from mummies that achieved both pre-
reproductive and ovipositing status. The most
common brood sizes developed from adult mealy-
bugs being composed of 2 (24.5%) or 3 (25.8%)


December 2007







Chong & Oetting: Larval Competition Affected Parasitoid Fitness


TABLE 1. MEANS ( SEM) OF SEX RATIO, BROOD SIZE, DEVELOPMENTAL TIME, SURVIVAL RATE, AND FEMALE AND MALE
PROGENY TIBIAL LENGTH OF ANAGYRUS sp. nov. NR. SINOPE FROM DIFFERENT HOST STAGES AT THE TIME
OF PARASITISM AND AT THE TIME OF MUMMIFICATION. THE DEVELOPMENTAL STAGES OF PHENACOCCUS MA-
DEIRENSIS ARE SECOND INSTAR (N2), THIRD INSTAR IMMATURE FEMALE (N3), PRE-REPRODUCTIVE ADULT FE-
MALE (PRE-OVIP) AND OVIPOSITING ADULT FEMALE (OVIP).

Host stage Develop-
Host stage at at mummi- Sex ratio mental Survival Female tibial Male tibial
parasitism fiction n (% males) Brood size time (d) rate (%) length (mm) length (mm)

N2 N2 52 67.3 6.6 a lb 16.1 0.2 c 100 0.30 0.01b 0.26 0.01 b
N3 24 28.3 4.9 b 1.3 0.1 b 18.0 0.3 b 100 0.36 0.01 a 0.29 0.01 a
Pre-Ovip 36 23.6 3.7 b 3.2 0.2 a 23.9 0.6 a 98.6 1.4 0.35 0.01 a 0.29 0.01 a
ANOVA statistics
F value 33.37 85.75 107.30 1.06 22.82 13.04
df 2, 109 2, 109 2, 109 2, 109 2,72 2,60
P value <0.0001 <0.0001 <0.0001 0.3512 <0.0001 <0.0001

Pre-Ovip Pre-Ovip 47 30.3 3.9 3.0 0.2 15.7 0.2 100 0.37 0.01 0.30 0.01
Ovip 51 35.8 3.7 2.9 0.2 16.1 0.3 97.1 1.7 0.37 0.01 0.31 0.01
ANOVA statistics
F value 0.90 0.13 0.93 2.81 0.01 0.79
df 1,96 1,96 1,96 1,96 1,90 1,74
P value 0.3442 0.7210 0.3372 0.0971 0.9342 0.3770


adult parasitoids per mummy (Fig. 1A; ANOVA:
F, 396 = 3.88, P < 0.0001). About 2% of all mummies
from both the host stages failed to produce any
parasitoids.
The mummies developing from the second in-
star mealybugs yielded predominantly single-sex
broods (about 42% all-female and 35% all-male)
(Fig. 1B; ANOVA: F10,30 = 18.71, P < 0.0001). Most
of the all-male broods emerged from mummies
that had achieved only second instar develop-
ment, while the majority of all-female broods
emerged from mummies that had achieved third
instar and adult development. In contrast, the
mummies developing from adult mealybugs
mostly yielded broods containing 30% males
(32.6%), followed by those that had sex ratios of
50% males (23.7%) and all-female (23%) (Fig. 1B;
ANOVA: F10,30 = 16.33, P < 0.0001).
The proportion of male progeny in a single
brood (i.e., the sex ratio) decreased with an in-
crease in the brood size of parasitoids developing
in the second instar mealybugs (% males = 0.53-
0.07xbrood size; F, 165 = 5.91, P = 0.0161). However,
there was only a weak relationship between the
brood size and sex ratio of adult parasitoids devel-
oping from second instar mealybugs (r2 = 0.0346).
By contrast, no linear relationship was detected
between the brood size and sex ratio of adult par-
asitoids developing from pre-reproductive adult
mealybugs (F,,,1 = 0.10, P = 0.7555, r2 = 0.0007).
The developmental time of adult parasitoids
from the second instar mealybugs increased
strongly with the brood size (Fig. 2A; developmen-
tal time = 15.3 + 2.6xbrood size; F1,165 = 125.76,P <


0.0001, r2 = 0.4325). On the other hand, the devel-
opmental time of parasitoids developing from
adult mealybugs was not influenced by brood size
(Fig. 2A; F,133, = 1.82, P = 0.1793, r2 = 0.0135). Sur-
vival rates of parasitoid larvae developing in both
the second instar and adult mealybugs ranged
from 83-100% and were not impacted by the par-
asitoid brood sizes (second instar: F1,165 = 0.30, P =
0.5823, r2 = 0.0018; adult: F13,,, = 1.51, P = 0.1190,
r2 = 0.0466).
Although there was no significant relationship
between brood size and female tibial length of
parasitoids developing from the second instar
mealybugs (Fig. 2B; F,165 = 0.01, P = 0.9170, r2 =
0.0001), the male tibial length increased slightly
with an increase in brood size (Fig. 2C; male tibial
length = 0.26 + 0.Olxbrood size; F, 165 = 15.12, P =
0.0002, r2 = 0.1398). An opposite pattern was ob-
served in the relationship between brood size and
progeny tibial length of parasitoids developing
from adult mealybugs. The tibial length of the fe-
male and male parasitoids developing from adult
mealybugs was significantly reduced with an in-
crease in the number of parasitoids per brood (fe-
male tibial length = 0.41 0.02xbrood size; Fig.
2B; F,133 = 52.38, P < 0.0001, r2 = 0.2970; male tib-
ial length = 0.35 0.Olxbrood size; Fig. 2C; F1,165
0.4841, P < 0.0001, r2 = 0.3218).
The developmental duration of parasitoids de-
veloping from second instar mealybugs was
slightly shortened as the percentage of males was
increased (Fig. 3A; developmental time = 20.7 -
2.7x% males; F1,165 = 11.48, P = 0.0009, r2 = 0.0651).
The development time was not influenced by the







Florida Entomologist 90(4)


(A) Brood size


a

60 -

50 i I
/2

40- B9gI= 3
4
0a aa
30
6

b
20 b 8



10
Sc 9









50 30%



a00%


10 be
bbl 40%


0 60%
1 80%


30 c c 5 100%

0 d c dc dd d dd
Second instar Adult

Host stages

Fig. 1. Distribution of brood sizes (A) and sex ratios (B) of Anagyrus sp. nov. nr. sinope from the second instar
and adult Phenacoccus madeirensis.


sex ratio of parasitoids which emerged from mum-
mies developing from adult mealybugs (Fig. 3A;
F,13 = 0.01, P = 0.9720, r2 = 0.0001). The propor-
tion of male parasitoids in broods developing from
the second instar and adult mealybugs did not in-
fluence the parasitoids' survival rate (second in-
star: F,165 = 1.73, P = 0.1905, r2 = 0.0104; adult:
F,133 = 0.04,P = 0.8359, r2 = 0.0003) and female tib-
ial length (Fig. 3B; second instar: F, 165 = 0.01, P =
0.9620, r2 = 0.0001; adult: F,,,, = 3.09, P = 0.0813,


r2 = 0.0243). Male tibial length again showed dif-
ferent responses to an increase in the proportion
of males in both host stages. The tibial length of
male parasitoids developing from the second in-
star mealybugs was reduced as the percentage of
males in a brood increased (Fig. 3C; male tibial
length = 0.30 0.03x% males; F,165 = 18.15, P <
0.0001, r2 = 0.1633). By contrast, the male tibial
length increased slightly with an increase in the
sex ratio of broods developing from adult mealy-


December 2007








Chong & Oetting: Larval Competition Affected Parasitoid Fitness


Second-instar nymphs


Cgs 35


25 30

a 25

E 20
C
. Is

4)







E 045

040
u



S0.30
2 025
4) JO
Li)5


0 1 2 3 4 5


a A





* ^

:
S ,
S /
U ,


6 7 8 0 1 2 3 4 5 6 7 8 9 10


Brood sizes


Fig. 2. Relationship of the brood size with the developmental time (A), female tibial length (B) and male tibial
length (C) ofAnagyrus sp. nov. nr. sinope developing in the second instar (left hand side) and adult (right hand side)
of Phenacoccus madeirensis. Each point represents the mean measurements of the whole brood. The solid line is the
best fitted linear regression model.



bugs (Fig. 3C; male tibial length = 0.27 + 0.08x% fray 1994). Many optimality models of brood size
males; F,133 = 23.47, P < 0.0001, r2 = 0.1871). and sex ratio have considered the effects of lim-
ited host resources or availability of time and eggs
DISCUSSION (e.g., Charnov & Skinner 1984; Waage & Godfray
1985; Godfray 1986). The primary brood size and
Brood size (or clutch size) and sex ratio are ma- sex ratio (i.e., the brood size and sex ratio at the
jor components of the life history strategy of many time of parasitism) are determined by the female
parasitoid species and have received much atten- parasitoids based on their physiological status
tion both experimentally and theoretically (God- (e.g., energy reserve and egg load) and foraging


Pre-reproductive adults


A



















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Florida Entomologist 90(4)


December 2007


Pre-reproductive adults


u 35

S30

C 25
0
-2
E 20
0


.O
105


0.50

0.45

% 0.40

0.35

0.30

E 0.25

0.20


0.40


E 0.35
-B

0.30


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0.20


A

0



l : -- -
S
*
* S
* S S
* ---- S-- -- S
* S
* S
* 0
* 0


0.00 0.25 0.50 0,75 1.00 0.00 0.25 0.50 0.75 1.00


Sex ratio (% males)

Fig. 3. Relationship of the sex ratio (% males) with the developmental time (A), female tibial length (B) and male
tibial length (C) ofAnagyrus sp. nov. nr. sinope developed in the second instar (left hand side) and adult (right hand
side) ofPhenacoccus madeirensis. Each point represents the mean measurements of the whole brood. The solid line
is the best fitted linear regression model.


experience (e.g., host density and travel time),
and the host's physiological status (e.g., host
stage, size or age). The primary brood size and sex
ratio provide the basis for larval competition
among brood mates, which in turn determine the
fitness of the progeny. We showed in this study
that the fitness consequences of larval competi-
tion for the gregarious parasitoid Anagyrus sp.


nov. nr. sinope were affected in part by the num-
ber of parasitoid larvae sharing limited resources
within a brood, and weakly by the competitive
pressure of one sex exerted on the other.
The model of sexual asymmetry in larval com-
petition suggests that the fitness of the less com-
petitive sex is reduced to a greater extent by an
increase in the proportion of the more competitive


Second-instar nymphs


B


* *
S .a *



**
S


c
C






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.
rr S
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*








0
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6



0







Chong & Oetting: Larval Competition Affected Parasitoid Fitness


sex (Godfray 1986). Overall, the sexual composi-
tion within a brood had little or no effect on the
development, survival, and body size ofAnagyrus
sp. nov. nr. sinope. The body size (using the hind
tibial length as the proxy) of female parasitoids
developing from both second instar and adult
P madeirensis appeared to be constant over a
range of sex ratios (% males). The conservation of
body size is beneficial to the female parasitoids
because their egg loads are reduced with even a
small reduction in the body size (Godfray 1994).
The sensitivity of female parasitoids to the reduc-
tion in body size often makes them more aggres-
sive competitors (Ode et al. 1996; Fidgen et al.
2000) in an effort to secure more resources for
their development to the optimal body size. An in-
crease in the percentage of males within a brood
reduced the intensity of competition from the fe-
male brood mates, which led to a slight increase
in the body size of male Anagyrus sp. nov. nr. si-
nope developing from adult mealybugs. In con-
trast, the body size of male parasitoids developing
from second instar mealybugs was only slightly
reduced by an increase in the percentage of males.
This is the reciprocal effect of the lower percent-
age of males in the mummies achieving adult de-
velopment (more resources to produce larger par-
asitoids) and the higher male ratio (almost all
males) in the mummies that remained as second
instar nymphs (produced smaller parasitoids).
An increase in brood size reduces the amount
of host resources available to individual parasi-
toid larvae, resulting in a reduction in the fitness
of the emerging adult parasitoids (Godfray 1994;
Ode et al. 1996). The negative effect of increased
brood size on progeny fitness has been found in
several studies (Nakamura 1995; Ode et al. 1996;
Zaviezo & Mills 2000; Allen & Hunt 2001; Guin-
nee et al. 2005; Keasar et al. 2006). The body sizes
of male and female Anagyrus sp. nov. nr. sinope
developing from adult P madeirensis were re-
duced with an increase in brood size, which
agreed with the prediction of larval competition
models. The developmental time of parasitoids
developing from the second instar mealybugs was
significantly lengthened because more time was
required to obtain sufficient resources to complete
development in larger broods. The increasing
body size of male Anagyrus sp. nov. nr. sinope de-
veloping from the second instar P madeirensis
contradicted the expectations of larval competi-
tion. We believe the observed contradictions in
the fitness responses ofAnagyrus sp. nov. nr. si-
nope to larval competition when developing in
second instar and adult mealybugs can best be ex-
plained by an alteration in the direction and in-
tensity of larval competition in hosts of different
quality and growth potential.
Larval competition among parasitoid brood
mates occurs when there is a limitation in the
amount of resources (i.e., host tissues) that can be


provided by the hosts for the parasitoid's develop-
ment. Many parasitoids, including Anagyrus sp.
nov. nr. sinope, actively select hosts of a specific
age or size that have sufficient resources to pro-
duce progeny of a higher fitness (Godfray 1994;
Harvey et al. 1998; Zaviezo & Mills 2000; Milonas
2005; Chong & Oetting 2006a). Adult and second
instar P madeirensis represent hosts of two differ-
ent qualities for Anagyrus sp. nov. nr. sinope
(Chong & Oetting 2006a). The adult mealybugs did
not continue to increase their body sizes through
growth after parasitism; effectively providing the
developing parasitoid larvae with a fixed amount
of resources. Therefore, the developing parasitoid
larvae in adult mealybugs were competing against
each other for the limited amount of resources. The
larval competition intensified with an increase in
brood size, which reduced the amount of resources
available to each parasitoid larva. As a result, the
body sizes of female and male parasitoids were re-
duced as the brood size increased. On the other
hand, the developing parasitoid larvae might have
suppressed their development to allow the second
instar mealybugs continued development after
parasitism (Chong & Oetting 2006a). These grow-
ing mealybugs accumulated resources and pro-
vided the developing parasitoid larvae with an in-
creased amount of resources. The increases in de-
velopmental time, brood size, proportion of fe-
males, and body size of Anagyrus sp. nov. nr.
sinope in mummies that achieved more advanced
developmental stage has already been demon-
strated by Chong & Oetting (2006a). Hosts of a
more advanced developmental stage provided suf-
ficient resources to allow the development of a
larger parasitoid brood without the expense on the
body size of the developing parasitoids. This in-
crease in available resources appeared to benefit
the male progeny more than the female progeny
(in terms of the gain in body size), perhaps because
excess resources were available to the male prog-
eny after satisfying the development requirements
of the female progeny. The increase in the male
parasitoid's body size to the increase in brood size
and host quality was, however, weak even when
the parasitoids stimulated the growth of the para-
sitized mealybugs (Vet et al. 1994).
This study showed that some aspects of the fit-
ness of parasitoids are influenced by the number
of parasitoid larvae and, to a lesser extent, the
sexual composition within a brood. The growth
potential of young hosts after parasitism may be
able to alter the direction and intensity of compe-
tition among the larvae ofAnagyrus sp. nov. nr.
sinope, particularly among male progeny. The re-
sults of this study indicated the potential influ-
ence of larval competition on parasitoid fitness
and the importance of considering the conse-
quence of the quality or growth potential of a host
stage on the evolution of brood size and sex ratio
in gregarious koinobiont parasitoids.












ACKNOWLEDGMENTS

We are grateful to John Noyes of the Natural History
Museum, London, UK, for the identification of Anagy-
rus sp. nov. nr. sinope, and Monica Townsend, Sherrie
Stevens, and Randy Harris for technical assistance. We
also thank 2 anonymous reviewers for helpful com-
ments on an earlier version of this manuscript.

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Florida Entomologist 90(4)







Yee: GF-120 Effects on Apple Maggot Fly


ATTRACTION, FEEDING, AND CONTROL OF RHAGOLETIS POMONELLA
(DIPTERA: TEPHRITIDAE) WITH GF-120 AND ADDED AMMONIA
IN WASHINGTON STATE

WEE L. YEE
United States Department of Agriculture, Agricultural Research Service,
Yakima Agricultural Research Laboratory, 5230 Konnowac Pass Road, Wapato, WA 98951

ABSTRACT

Experiments were conducted in 2005 and 2006 in western Washington state to determine ef-
fects of adding ammonium carbonate (AC) and ammonium acetate (AA) to GF-120 NF Nat-
uralyte@ Fruit Fly Bait (Dow AgroSciences, Indianapolis, IN) on attraction, feeding, and
control of the apple maggot fly, Rhagoletis pomonella (Walsh). In the field, sticky yellow
panel traps baited with GF-120 + 10% AC attracted more flies than those baited with GF-
120 + 10% AA or GF-120 alone. In the laboratory, female flies responded more frequently to
sucrose and 20% GF-120 than to water and 40% GF-120 + 10% AA on apples, perhaps be-
cause of the confined conditions during testing. In the field, fly attraction and feeding were
greater for GF-120 + 10% or 2.5% AC or AA than GF-120 alone on apple leaves. In 2 spray
tests with 100 mL of GF-120 alone and GF-120 + 2.5% AC or AA applied on single apple
trees, larval infestations in fruit were reduced up to 99% compared with controls, but there
were no differences among treatments, and none was different from spinosad alone. Results
indicate the attractiveness of GF-120 to R. pomonella can be increased with added ammonia,
but that this does not necessarily result in greater control, perhaps because the added am-
monia volatilizes too quickly. Results suggest that at the spray volumes used, GF-120 alone
or even spinosad alone can greatly reduce local R. pomonella populations in Washington.

Key Words: apple maggot fly, bait sprays, spinosad

RESUME

Se realizaron experiments durante el 2005 y 2006 en el oeste del estado de Washington
para determinar los efectos de anadir carbonate de amonio (CA) y acetate de amonio (AA) al
cebo para moscas de la fruta, GF-120 NF Naturalyte Fruit Fly Bait (Dow AgroSciences, In-
dianapolis, IN) sobre la atracci6n, alimentaci6n y el control del gusano de la manzana,
Rhagoletis pomonella (Walsh). En el campo, trampas de panels pegajosas de color amarillo
con cebo GF-120 + 10% CA atrayeron mas moscas que las que estaban con solo GF-120 + 10%
AA o GF-120. En el laboratorio, las hembras de moscas respondieron mas frecuentemente a
la sucrosa y 20% de GF-120 que con agua y el 40% GF-120 + 10% AA sobre las manzanas,
quizas por la condiciones confinadas del experiment. En el laboratorio, la atracci6n y la ali-
mentaci6n de las moscas fue mayor para el GF-120 + 10% o 2.5% CA o AA que solo para el
GF-120 sobre las hojas de manzana. En 2 de las pruebas de asperci6n con 100 mL de solo GF-
120 y GF-120 + 2.5% CA o AA aplicado sobre un arbol individual de manzana, las infestacio-
nes de las larvas en el fruto fue reducida hasta el 99% comparado con el control, pero no hubo
diferencias entire los tratamientos, y ninguno de ellos fue diferente con respect al trata-
miento con solo el spinosad. Los resultados indican que se puede aumentar la capacidad de
GF-120 para atraer R. pomonella al anadir el amonio, pero este no necesariamente result
en un mayor control, quizas porque el amonio anadido se volatiliza demasiado rapido. Los
resultados sugieren que en el volume de las asperciones usadas, solo con GF-120 o aun solo
el spinosad puede reducir grandamente las poblaciones de R. pomonella en Washington.


Apple maggot fly, Rhagoletis pomonella (Walsh),
is a major quarantine pest of apples, Malus domes-
tica (Borkh.) Borkh., in Washington state and other
parts of the Pacific Northwest of the U.S. Washing-
ton state is the leading producer of apples in the
U.S., with a production value in 2005 of US$1.23 bil-
lion, at an estimated per hectare value of $3,078
over -65,500 harvested ha (USDA 2006). To main-
tain apple production profits, insect pest manage-
ment is vital. Until recently, R pomonella in Wash-
ington was believed to have established only in the


western part of the state, along the lower Columbia
River Gorge in the southern part, and in Spokane in
the eastern part. However, larval infestations of
hawthorns and non-commercial apples were de-
tected in central Washington from 2003 to 2005, re-
sulting in a partial quarantine of Kittitas County in
2004 and Yakima County in 2005 (Washington
State Department of Agriculture 2005), near major
apple-producing areas. There is zero tolerance for
larvae in fresh market apples (Washington State
Department of Agriculture 2001), so any trap cap-







Florida Entomologist 90(4)


tures ofR. pomonella on isolated hawthorn or feral
apple trees in central Washington trigger the spray-
ing of trees to eliminate populations or prevent their
spread (Klaus et al. 2007). The organophosphate in-
secticide phosmet has been effectively used for this
purpose. To date, apple maggot has not been de-
tected in commercial orchards in central Washing-
ton. Despite the effectiveness of phosmet, alterna-
tives are needed because of impending restrictions
on the use of this and other highly toxic organophos-
phates (Food Quality and Protection Act 1996).
Various new insecticides mixed with hydro-
lyzed protein baits may provide an effective alter-
native to conventional organophosphate sprays.
Since around 2000, GF-120 NF Naturalyte
Fruit Fly Bait (Dow AgroSciences, Indianapolis,
IN) (GF-120) has been the most frequently tested
bait against subtropical, tropical, and temperate
fruit flies (e.g., Vargas et al. 2002; Prokopy et al.
2003; Barry & Polavarapu 2004). GF-120 is a
modified, more concentrated and marketed ver-
sion of Solbait originally developed for Anas-
trepha and Ceratitis fly control (Burns et al. 2001;
Moreno & Mangan 2003). The frequent testing of
this bait is the result of its effectiveness against
representative tephritids, ease of use, low spray
coverage needed, and organic labeling. GF-120
contains 0.02% spinosad (wt/vol), an insecticide
derived from fermentation products of the bacte-
rium Saccharopolyspora spinosa Mertz and Yao
that has a high safety profile (Dow AgroSciences
2006), as well as 1% ammonium acetate as an at-
tractant (Thomas & Mangan 2005).
Results with GF-120 against Rhagoletis spe-
cies for control in the field have overall been posi-
tive. However, there are no published studies
showing it can eliminate infestations after only
one season of use. In New York, GF-120 was inef-
fective against R. pomonella (Reissig 2003), but in
Michigan it was effective in one of two years (Pelz
et al. 2005). In California, it was effective against
the walnut husk fly, R. complete Cresson (Van
Steenwyk et al. 2003), and in Washington and
Utah it was effective against the western cherry
fruit fly, R. indifferens Curran (Yee & Chapman
2005; Yee & Alston 2006).
Research has indicated that GF-120 is not or
not highly attractive to Rhagoletis flies (Barry &
Polavarapu 2004; Pelz et al. 2005; Yee & Chap-
man 2005; Yee 2006) and recent research has em-
phasized the need to make GF-120 more attrac-
tive (Pelz-Stelinski et al. 2006). A logical choice of
materials to make the bait more attractive is am-
monia, which is associated with protein-rich foods
and has long been known to attract R. pomonella
(Hodson 1948) and R. indifferens (Frick et al.
1954). Indeed, even though GF-120 has 1% am-
monium acetate, adding more ammonium acetate
to GF-120 enhanced its attractiveness to the east-
ern cherry fruit fly, R. cingulata (Loew) (Pelz-Ste-
linski et al. 2006). However, whether increasing


the attractiveness of GF-120 with additional am-
monia results in improved control of larval infes-
tations in fruit is unclear.
In this study, the objectives were to determine
the attraction, feeding, and control of R.
pomonella with GF-120 and added ammonia in
Washington. Three hypotheses were tested: (1)
adding ammonium carbonate or ammonium ace-
tate to GF-120 in lures increases trap captures
compared with GF-120 alone in lures; (2) adding
ammonia compounds to GF-120 also increases fly
attraction and feeding responses when baits are
sprayed on leaves; and (3) GF-120 containing ad-
ditional ammonia compounds decrease larval in-
festations more than GF-120 alone.

MATERIALS AND METHODS

Study Sites and Experimental Design

Four field and laboratory experiments consist-
ing of various tests were conducted in western
Washington in 2005 and 2006. A summary of the
experimental sites, test years, replicate sizes, ex-
perimental settings, and experimental designs is
shown in Table 1. Study sites were known to be
infested with R. pomonella. Treatments and de-
tails for each of the 4 experiments follow.

Experiment 1: Effects of GF-120 with Ammonia
Compounds on Attraction to Traps

For test 1A, various lures were tested with 14 x
23 cm sticky yellow panel traps (Trece, Adair, OK).
Ammonium carbonate (AC) (Keystone Universal
Corp., Melvindale, MI) and ammonium acetate
(AA) (EMD Brand, Barmstadt, Germany) were
used as sources of additional ammonia. Compari-
sons were: (1) a control, (2) 10 g AC, (3) 17% GF-
120, (4) 40% GF-120, (5) 40% GF-120 + 10% AC,
and (6) 40% GF-120 + 10% AA. Percent GF-120 was
based on vol/vol, but % AC and AA was based on wt/
wt. Blank GF-120, without spinosad, was used. The
10 g AC was placed in a clear plastic vial (Thorton
Plastic Co., No. 55-7, Salt Lake City, UT) with two
1-mm holes on the lid. Ten mL of each of the GF-
120 treatments was placed in a 15-mL polypropy-
lene narrow-mouth Nalgene bottle (Nalge Nunc
International, Rochester, NY) with the cap re-
moved (13 mm diameter opening). A 0.25-g cotton
ball was placed inside each bottle to prevent spill-
age. Each lure was hung 1 to 2.5 cm above the cen-
ter of each trap, which was suspended from a
branch ~2 m above the ground. Lures were not re-
placed during the season. Within each row, 1 trap
was placed in every other tree (cv 'MacIntosh'). One
block consisted of 2 seven- or eight-tree rows. Trees
were 3.0 to 4.6 m wide and spaced 10 m apart.
Traps were rotated each week among trees within a
block. Flies were removed weekly and counted. The
test was conducted 19 Jul to 22 Aug.


December 2007







Yee: GF-120 Effects on Apple Maggot Fly


TABLE 1. SUMMARY OF RHAGOLETIS POMONELLA ATTRACTION, FEEDING, AND CONTROL EXPERIMENTS.

Apple tree
Experiment Test Site Year No. replicates setting Experimental design

1. Attraction, traps 1A V 2005 4 Orchard RBD, blocks: tree rows
1B SCR 2005 4 Scattered trees RBD, blocks: single trees
2. Feeding, containers Lab 2005 10 to 27 single flies CRD
3. Attraction, feeding 3A SCR 2005 3 to 6, on ea. of 8 d Scattered trees RBD, blocks: single trees
3B SCR 2006 4, 5, on ea. of 11 d Scattered trees RBD, blocks: single trees
3C P 2005 4, all in 1 d Orchard Each treatment on same
four trees
3D P 2006 10, 2 on ea. of 5 d Orchard RBD, blocks: single trees
4. Larval infestations 4A P 2005 5 Orchard' CRD, single trees
4B P 2006 5 Orchard' CRD, single trees

V, Vancouver (Clark County) (45 37.45'N, 122 39.78'W); SCR, Saint Cloud Ranch (Skamania County) (45 35.88'N, 122 07.03'W);
P, Puyallup (Pierce County) (47 11.76'N, 122 16.50'W).
RBD, randomized block design; CRD, completely randomized design.
"One or 2 treatments tested on single day (see text for details).
bSingle tree replicates were sprayed.


Test 1B at Saint Cloud Ranch was conducted
with similar methods, with the following differ-
ences. There was no 10-g AC lure. The test was set
up in 4 scattered apple trees (cv'Newtown','Wine-
sap', 'Spitzenberg', and 'Early Transparent') with
a single tree as a block. A replicate trap of the con-
trol and each treatment was hung in each tree.
Positions of traps within trees were rotated
weekly. Trees were 6.1 to 7.6 m wide and 9.1 to
15.2 m apart. The test was conducted 28 Jul to 22
Sep.

Experiment 2: Effects of GF-120 with Ammonia
Compounds on Feeding Responses in the Laboratory

Infested apple and hawthorn fruit were col-
lected in Vancouver and Puyallup in summer/fall
2004 and placed in tubs, where larvae emerged.
The puparia subsequently formed were then col-
lected and placed in moist soil and chilled at ~4C
for 6 months. Puparia were transferred to 20-
27C for adult emergence, after which flies were
immediately placed in 473-mL paper containers
with water and a 5% sucrose solution on a wick,
but no other food. Flies were tested at 3 to 5 d af-
ter emergence. Five 50 pL drops of (1) water, (2)
13% sucrose (wt/wt), (3) 17% GF-120, (4) 20% GF-
120, (5) 40% GF-120, (6) 40% GF-120 + 10% AC, or
(7) 40% GF-120 + 10% AA were placed equidistant
on top of 1 apple (cv 'Fuji') in a 1.9-liter paper con-
tainer with an organdy screen that allowed view-
ing. Blank GF-120 was used. One male or female
fly was then introduced into this test container.
After 1 min, the fly was watched continuously for
60 min, and the numbers of feeding bouts and
feeding durations were recorded. No water was
provided in the test container. The test was con-
ducted in a brightly lit room at 24 to 27C.


Experiment 3: Effects of GF-120 with Ammonia
Compounds on Attraction and Feeding in the Field

Test 3A at Saint Cloud Ranch in 2005 con-
sisted of: (1) 13% sucrose, (2) 17% GF-120, 3) 40%
GF-120, (4) 40% GF-120 + 10% AC, and (5) 40%
GF-120 + 10% AA. Blank GF-120 was used. Ten
mL of each treatment were sprayed on an apple
branch with ~30 leaves with a 32-oz volume spray
bottle (Consolidated Plastics Co., Twinsburg,
OH). The five treatments were applied ~1 m apart
within 1 tree and ~1.5 m above the ground. Trees
were ~6.1 m wide. Numbers of flies <15 cm from
spray drops and feeding on drops were recorded.
Each treatment was observed for a continuous 2-
min period, followed immediately by the next
treatment for a total of 3 periods per treatment
during 30 min. The sex of flies was recorded, but
sexes were pooled for presentation and analyses
because of the low numbers. Sprayed leaves and
branches were removed after the completion of
~30 min. Observations were recorded from 0800-
1300 h. On each day, observations were made on 3
to 6 replicate trees >23 m apart. The test was con-
ducted on 8 dates from 26 Jul to 25 Aug. Positions
of treatments within trees were randomized. For
the entire test, there were 3.7 h of continuous ob-
servations for each of the 5 treatments (18.5 h to-
tal). Test 3B at Saint Cloud Ranch in 2006 was
similar to test 3A, but the 17% GF-120 treatment
was dropped, all GF-120 treatments contained
0.0096% spinosad (wt/vol) (Entrust [80% spi-
nosad], Dow AgroSciences, Indianapolis, IN), and
a spinosad alone treatment (also 0.0096% wt/vol)
was included. (In the rest of the tests in experi-
ments 3 and 4 below, the spinosad concentration
in all GF-120 and spinosad alone treatments was
also 0.0096%.) The data collecting method was







Florida Entomologist 90(4)


the same as in test 3A. The test was conducted on
11 dates from 10 Aug to 7 Sep. For the entire test,
there were 5.4 h of continuous observations for
each of the 5 treatments (27 h total).
Test 3C in Puyallup in 2005 consisted of: (1) a wa-
ter control, (2) 13% sugar, (3) 17% GF-120, (4) 40%
GF-120, (5) 40% GF-120 + 2.5% AC, (6) 40% GF-120
+ 2.5% AA, and (7) spinosad alone. Ten mL of each
were applied on a branch of an apple tree as before.
Trees (3 unidentified varieties: striped, red/late, and
yellow/early) in the orchard were ~5 to 7 m tall and
wide. The design differed from that used in tests 3A
and 3B because the initial thought was that odors
among treatments compared simultaneously within
trees could interfere with one another. Each treat-
ment or control was tested on a single tree by itself,
with 4 replicate trees each day. Observations were
made on different days between 0900 and 1300 h: 8
Aug, 13% sucrose and 17% GF-120; 9 Aug, 40% GF-
120 and spinosad alone; 10 Aug, 40% GF-120 + 2.5%
AC; and 12 Aug, water control and GF-120 + 2.5%
AA. Treated leaves were removed, and the same 4
trees were used for other treatments. Data record-
ing also differed from that used in tests 3A and 3B.
Flies seen every 2 min <15 cm from spray drops and
feeding on drops within 30-min periods were re-
corded, for 16 instantaneous recordings over these
periods. All 4 d were sunny, with high temperatures
of 20.4 to 23.8C, wind speeds of 75 to 140 m/min,
and RH of 50 to 62%. Test 3D in Puyallup in 2006
was similar to test 3C, except that 17% GF-120 was
dropped and all five treatments and the control
were compared simultaneously within single trees,
as in tests 3A and 3B. Data recording was the same
as in test 3C. Observations from two replicate trees
were made on each of five d from 24 Jul to 2 Aug, for
10 total replicates per treatment.

Experiment 4: Effects of GF-120 with Ammonia
Compound Sprays on Larval Infestations

In all spray tests, single apple trees were
sprayed. In Washington, flies of threat to commer-
cial orchards are generally found in single feral
trees or small patches of trees, and not in the or-
chards themselves, so spot instead of broadcast
sprays were used. The label rates for spot sprays
are 30 to 90 mL of undiluted GF-120 spray solution/
tree (Dow AgroSciences 2006). Treatments in test
4A were delivered at 40 mL of undiluted GF-120 in
100 total mL of spray/tree with RL Flo-Master
pressurized sprayers (Root-Lowell Manufacturing
Co., Lowell, MI). Treatments in test 4A compared:
(1) an unsprayed control, (2) 40% GF-120, (3) 40%
GF-120 + 2.5% AC, (4) 40% GF-120 + 2.5% AA, and
(5) spinosad alone. Trees (mostly cv. 'Jonagold')
within rows in an orchard were used, but because of
the irregular numbers of trees per row, a completely
randomized instead of a randomized block design
was used. Trees were 1.7 to 4.5 m tall and 1.5 to 3.8
m wide, most spaced 4 m from others. Sprays were


initiated <7 d of first fly capture on AC-baited sticky
yellow panel traps. After the first fly capture, AC
lures were removed, leaving 1 unbaited panel on
each tree throughout the test. Fly captures were ei-
ther from within the trapped trees or from sur-
rounding trees. Larval infestations could have orig-
inated from flies from either source. However, adult
flies may have been caught before they able to ovi-
posit, so there may be discordance between fly cap-
tures and larval infestations. Weekly applications
were made 1 Jul to 6 Sep, for 11 total sprays. There
is no maximum number of applications on the GF-
120 label (Dow AgroSciences 2006), and in Califor-
nia, there can be up to 19 aerial or ground applica-
tions per acre per season made for control of exotic
fruit flies (Cheney 2005). Forty nine to 108 apples
were picked from each tree, except from two control
trees, where only 8 and 9 apples were present. Test
4B compared the same treatments, also with 100
mL spray/tree and in a completely randomized de-
sign in an orchard (3 unidentified apple varieties:
striped, red/late, and yellow/early) with 5.0 to 6.7 m
tall and wide trees spaced 4 m apart. Weekly appli-
cations were made 11 Jul to 12 Sep, for 9 or 10 total
sprays (some blocks were harvested before the last
spray). One hundred apples were picked from each
tree. In both tests, apples were placed in tubs and
held for 2 months to allow larvae to emerge, after
which counts of puparia were made.

Statistics

For experiment 1, data were analyzed by ran-
domized block analysis of variance (ANOVA), fol-
lowed by Fisher's LSD test for mean separation.
Two-way ANOVA was also conducted to deter-
mine sex, bait, and sex x bait effects. For experi-
ment 2, Fisher's exact test with R x C tables was
used; differences among proportions also were an-
alyzed with a Tukey-type multiple comparison
test among proportions (Zar 1999). Experiment 3
tests were set up for ANOVA, but there were too
many zero values in some replicates in tests 3A
and 3B for this analysis. Thus, chi-square good-
ness of fit tests were used, with counts pooled from
all replicates and dates and with equal ratios for
all treatments as expected values. Numbers of
flies near bait drops and numbers of flies that fed
were pooled to give higher counts for analyses. In
tests 3C and 3D, chi-square tests were performed
as well, but there also were enough flies to conduct
one-way ANOVA and randomized block ANOVA,
respectively, followed by Fisher's LSD test. In ex-
periment 3, data were not based on separate sam-
ples as in experiment 2, and some treatments had
only 1 or 2 flies, so the Tukey-type multiple com-
parison test among proportions was not per-
formed. For experiment 4, one-way ANOVA were
conducted. Data were subjected to square-root (y)
or square-root (y + 1) transformation (when counts
were low and there were zeros) to stabilize the


December 2007







Yee: GF-120 Effects on Apple Maggot Fly


variance. The Statistical Analysis System (SAS
Institute, Inc. 2004) was used for ANOVAs.

RESULTS

Experiment 1: Effects of GF-120 with Ammonia
Compounds on Attraction to Traps

In test 1A, the 10 g AC lure attracted more flies
than the 40% GF-120 + 10% AC lure, which at-
tracted more than the control and the 17% and
40% GF-120 lures, although not more than the
40% GF-120 + 10% AA lure (Table 2). The control
and treatments attracted more females than
males (two-way ANOVA, F = 8.2; df = 1, 36; P =
0.0070), although the sex response pattern across
treatments was not different (bait: F = 11.1; df = 5,
36; P < 0.0001; sex x bait: F = 2.2; df = 5, 36; P =
0.0736). The percentages of females (n = 86) in the
control, 10 g AC, 17% GF-120, 40% GF-120, 40%
GF-120 + 10% AC, and 40% GF-120 + 10% AA
treatments were 5.8, 54.7, 3.5, 5.8, 20.9, and 9.3%,
respectively, and for males (n = 37), they were 0,
40.5, 18.9, 5.5, 27.0, and 8.1%, respectively.
In test 1B, the 40% GF-120 + 10% AC lure at-
tracted more flies than the control and other GF-
120 lures, including the 40% GF-120 + 10% AA lure
(Table 2). There was no difference in the numbers
of females and males caught (two-way ANOVA, F =
1.8; df = 1, 30; P = 0.1878), and the sex response
pattern was similar across treatments (bait: F =
19.9; df = 4, 30; P < 0.0001; sex x bait: F = 0.4; df =
4, 30; P = 0.8080). The percentages of females (n =
803) in the control, 17% GF-120, 40% GF-120, 40%
GF-120 + 10% AC, and 40% GF-120 + 10% AA
treatments were 3.0, 4.9, 13.7, 65.7, and 12.7%, re-
spectively, and for males (n = 614), they were 2.0,
4.9, 25.0, 57.5, and 10.6%, respectively.

Experiment 2: Effects of GF-120 with Ammonia
Compounds on Feeding Responses in the Laboratory
Because of the low response to the baits, mean
numbers of feeding bouts and feeding durations


were not analyzed (e.g., there were no drinks on
water). Fisher's exact test showed that numbers
that fed were dependent on treatment in females
(P = 0.0054), but not in males (P = 0.2826). Anal-
yses of percentages (Table 3) indicated that fe-
male flies responded less to water and 40% GF-
120 + 10% AA than to 13% sucrose and 20% GF-
120. Unlike females, however, males did not re-
spond differently to any treatment.

Experiment 3: Effects of GF-120 with Ammonia
Compounds on Attraction and Feeding in the Field

In test 3A, fly sightings were infrequent given
that there were 3.7 total h of continuous observa-
tions/treatment. Numbers of sightings of flies
near or feeding on sucrose, 17% GF-120, and 40%
GF-120 were similar (1 or 2), and lower than on
40% GF-120 + 10% AC or 10% AA (Table 4). Un-
like in experiment 1, there were no evident differ-
ences in responses to GF-120 + 10% AC and 10%
AA. In test 3B, fly responses were also low over 5.7
total h of observation/treatment and were similar
to those in test 3A. Responses to sucrose, 40% GF-
120, and spinosad alone were similar and lower
than to GF-120 + 10% AC or 10% AA (Table 4).
In tests 3C and 3D, numbers of fly sightings
were greater than in tests 3A and 3B. The num-
bers near or feeding on water, sucrose, and spi-
nosad alone were similar, lower than on 17% or
40% GF-120, and much lower than on GF-120 +
2.5% AC or 2.5% AA (Table 5). No differences were
seen between GF-120 + 2.5% AC and GF-120 +
2.5% AA treatments.

Experiment 4: Effects of GF-120 with Ammonia
Compound Sprays on Larval Infestations

In test 4A, fewer flies were caught on traps in
all treatments than in the control (Table 6). There
were high levels of larval control with all the
treatments, and GF-120 + 2.5% AC and 2.5% AA
treatments did not perform better than GF-120
alone, and statistically no better than spinosad


TABLE 2. MEAN TOTAL NUMBERS OF RHAGOLETIS POMONELLA + SE CAUGHT OVER THE SEASON PER STICKY YELLOW
PANEL TRAP WITH DIFFERENT LURES ON APPLE TREES AT 2 SITES, WA, 2005.

Test 1A: Vancouver Test 1B: Saint Cloud Ranch
Treatment 19 Jul to 22 Aug 28 Jul to 22 Sep

Control 1.2 + 0.9 c 9.0 + 2.0 b
10 gAC 15.5 2.7 a Not tested
17% GF-120 1.8 1.1 c 17.3 1.8 b
40% GF-120 1.8 1.0 c 66.0 47.9 b
40% GF-120 + 10% AC 7.0 2.3 b 220.3 + 39.1 a
40% GF-120 + 10% AA 2.8 1.8 bc 41.8 4.7 b
Randomized block F = 8.4; df = 5, 15 F = 13.0; df = 4, 12
ANOVA P = 0.0006 P = 0.0003

Blank GF-120 used.
Means within columns followed by the same letter are not significantly different (Fisher's LSD test, P > 0.05).







Florida Entomologist 90(4)


December 2007


TABLE 3. PERCENT OF RHAGOLETIS POMONELLA THAT DRANK OR FED ON WATER OR GF-120 BAITS ON APPLES OVER 1-
H OBSERVATIONS IN THE LABORATORY.

Females Males

Treatment n % Drank or fed n % Drank or fed

Water 22 0.0 b 18 0.0 a
13% Sucrose 27 33.3 a 20 15.0 a
17% GF-120 25 20.0 ab 21 0.0 a
20% GF-120 22 27.3 a 18 11.1 a
40% GF-120 27 7.4 ab 24 16.7 a
40% GF-120 + 10% AC 21 9.5 ab 20 5.0 a
40% GF-120 + 10% AA 12 0.0 b 10 10.0 a

% followed by the same letter within columns are not significantly different (Tukey-type multiple comparison test among pro-
portions, P > 0.05).
Blank GF-120 used.


alone (Table 6). In test 4B, fewer flies were caught
in all treatments than in the control. Within treat-
ments, fewest were caught in the spinosad alone
and most in the GF-120 + 2.5% AC treatment (Ta-
ble 6). Despite different effects on adult captures,
all treatments again resulted in similarly high
levels of larval control. Also, again GF-120 + 2.5%
AC and 2.5% AA treatments did not perform bet-
ter than GF-120 alone or spinosad alone (Table 6).

DISCUSSION

In experiment 1, addition of 10% AC to 40% GF-
120 enhanced the attraction of R. pomonella to
sticky yellow panel traps, supporting the first hy-
pothesis and showing that GF-120 can be modified
to increase fly responses, although it did not at-
tract as many flies as the 10 g AC lure. Addition of


ammonium bicarbonate also substantially in-
creased the attractiveness of the commercial bait
Nu-Lure to R. pomonella inside cages compared
with Nu-Lure alone (Hendrichs et al. 1990). GF-
120 + 10% AC was more attractive than GF-120 +
10% AA, suggesting that when amounts of the
compounds in GF-120 are the same, AC releases
more ammonia than AA. Females were more re-
sponsive to the GF-120 lures on traps than males
in one test, likely because their need for protein is
greater (Webster et al. 1979), but the lures affected
the sexes similarly in terms of relative responses.
In experiment 2, feeding responses by flies to
sucrose and GF-120 with or without added ammo-
nia on apples in containers in the laboratory were
low, but results demonstrated that sucrose and
20% GF-120 attracted females or caused them to
feed more than on water. Female flies appeared


TABLE 4. NUMBERS OF SIGHTINGS OF RHAGOLETIS POMONELLA FEEDING ON OR NEAR SUCROSE, GF-120 BAITS, AND
SPINOSAD ON APPLE LEAVES AT SAINT CLOUD RANCH, WA, 2005 AND 2006.

Treatment No. feeding No. <15 cm from baitb Total fly sightings

Test 3A: 2005
13% Sucrose 1 0 1
17% GF-120 1 1 2
40% GF-120 1 1 2
40% GF-120 + 10% AC 8 6 14
40% GF-120 + 10% AA 4 12 16
Total fly sightings: Chi-Square = 30.9; df = 4; P < 0.0001.
Test 3B: 2006
13% Sucrose 4 0 4
40% GF-120 2 0 2
40% GF-120 + 10% AC 9 1 10
40% GF-120 + 10% AA 7 2 9
Spinosad alone 0 1 1
Total fly sightings: Chi-Square = 12.8; df = 4; P = 0.0121.

2005, blank GF-120 used; 2006, GF-120 baits and spinosad alone both had 0.0096% spinosad (wt/vol).
aExpected cells <5, data not analyzed.
bNot feeding










Yee: GF-120 Effects on Apple Maggot Fly


TABLE 5. NUMBERS OF SIGHTINGS OF RHAGOLETIS POMONELLA DRINKING OR FEEDING ON OR NEAR WATER, SUCROSE,
GF-120 BAITS, AND SPINOSAD ON APPLE LEAVES IN PUYALLUP, WA, 2005 AND 2006.


No. drinking
or feeding


Treatment


Total fly sightings


Test 3C: 2005
Water 0 1
13% Sucrose 0 1
17% GF-120 0 13
40% GF-120 3 41
40% GF-120 + 2.5% AC 6 88
40% GF-120 + 2.5% AA 6 89
Spinosad alone 0 2
Total fly sightings: Chi-Square = 286.7; df= 6; P < 0.0001.
Mean fly sightings SE: One-way ANOVA: F = 36.3; df = 6, 21; P < 0.0001.
Test 3D: 2006
Water 0 1
13% Sucrose 2 5
40% GF-120 1 14
40% GF-120 + 2.5% AC 6 36
40% GF-120 + 2.5% AA 3 26
Spinosad alone 0 0
Total fly sightings: Chi-Square = 51.8; df = 4; P < 0.0001; spinosad alone not included.
Mean fly sightings SE: Randomized Block ANOVA: F = 9.0; df = 4, 45; P < 0.0001.
2005 and 2006, GF-120 baits and spinosad alone had 0.0096% spinosad (wt/vol).


Mean fly sightings SE
(per 30 min)


0.02 0.02 d
0.02 0.02 d
3.20 1.4 c
10.20 1.6 b
22.00 3.4 a
22.02 2.3 a
0.50 0.3 d



0.10 0.1 d
0.50 0.2 cd
1.40 0.5 bc
3.60 0.9 a
2.60 1.0 ab
0.00 0.0 d


"Expected cells <5, data not analyzed with Chi-Square.
Total fly sightings + SE: means followed by the same letter are not significantly different (P > 0.05, Fisher's LSD test).


more responsive to sucrose and baits than males,
based on overall response percentages, consistent
with the attraction results in experiment 1. The
lack of higher responses to GF-120 with added
ammonia than to GF-120 alone was surprising
given the results of experiment 1. In fact, addition
of ammonia compounds numerically lowered re-
sponses of flies to GF-120. Perhaps release rates
inside containers were initially very high and ac-
tually repelled rather than attracted flies. Had ob-
servations lasted longer, rates may have attracted
for a period of time before dissipating to levels
that no longer attracted. The environment inside
containers in the laboratory probably lacked im-
portant cues flies need to respond to ammonia.
In 4 tests in experiment 3, GF-120 alone
seemed not attractive or only slightly attractive
when it was sprayed on apple leaves, but adding
AC and AA to GF-120 clearly made it attractive. A
2.5% concentration of either AC or AA was suffi-
cient to increase responses, but direct compari-
sons are needed to determine if it is as effective as
the 10% concentration. Results support the sec-
ond hypothesis and previous work showing at-
traction of R. cingulata to GF-120 enhanced with
AA (Pelz-Stelinski et al. 2006). In tests 3A and 3B
at Saint Cloud Ranch, GF-120 alone did not ap-
pear to be attractive compared with sucrose con-
trols. The windiness at this site may partially ex-
plain the inability to detect any attractiveness of
GF-120. In contrast, in the different environment


in tests 3C and 3D in Puyallup, GF-120 alone did
appear attractive and this may have been caused
by olfactory cues, visual cues, or both. The higher
fly activity in Puyallup than at Saint Cloud Ranch
could have influenced results and increased the
relative effectiveness of GF-120 alone in Puyal-
lup. Despite the greater attraction to GF-120 +
AC or AA, the low numbers of flies that responded
suggest ammonia release from drops was too low
to elicit strong or immediate responses from a
large percentage of a fly population. High ammo-
nia release from lures in experiment 1 is likely
difficult to duplicate from small spray drops on
leaves. The lack of differences between AC and AA
could also be caused by the low amount of ammo-
nia released from spray drops. In tests 3C and 3D,
the numbers of feeding bouts on all GF-120 treat-
ments were lower than numbers of non-feeding
visits. This is consistent with the idea that GF-
120 causes arrestment of flies, as with the blue-
berry maggot, R. mendax Curran (Pelz et al.
2005). More evidence is needed to confirm this
with R. pomonella.
In experiment 4, adding AC or AA to GF-120
did not reduce fly numbers and larval infestations
compared with GF-120 alone, thus not supporting
the third hypothesis, but all GF-120 treatments
resulted in very high levels of larval control in the
2 tests. In test 4B, more flies were caught on traps
in trees sprayed with GF-120 + AC than in trees
with GF-120 alone, suggesting ammonia from the








Florida Entomologist 90(4)


TABLE 6. MEAN NUMBERS OF RHAGOLETIS POMONELLA FLIES PER TRAP AND LARVAE PER APPLE FRUIT SE IN GF-120
BAIT SPRAY TESTS IN PUYALLUP, WA, 2005 AND 2006.

Treatment No. flies/trap % Decrease No. larvae/fruit % Decrease

Test 4A: 2005
Control 35.2 1.9 a 1.22 0.59 a -
40% GF-120 11.6 3.1 b 67.0 0.07 0.02 b 94.3
40% GF-120 + 2.5% AC 28.2 2.2 b 19.9 0.08 0.04 b 93.4
40% GF-120 + 2.5% AA 17.8 2.6 b 49.4 0.16 0.03 b 86.9
Spinosad alone 14.0 2.8 b 60.2 0.05 0.004 b 95.9
One-way ANOVA F = 12.5 F = 4.4
df = 4, 20 P < 0.0001 P = 0.0098
Test 4B: 2006
Control 119.4 10.1 a 0.99 0.23 a -
40% GF-120 15.0 1.4 c 87.4 0.08 0.02 b 91.9
40% GF-120 + 2.5% AC 24.4 1.6 b 79.6 0.05 0.02 b 94.9
40% GF-120 + 2.5% AA 9.6 1.5 c 92.0 0.13 t 0.05 b 86.9
Spinosad alone 1.8 0.8 d 98.5 0.01 0.01 b 99.0
One-way ANOVA F = 158.7 F = 19.4
df =4, 20 P< 0.0001 P< 0.0001

GF-120 and spinosad alone had 0.0096% spinosad (wt/vol). 100 mL spray/tree.
Means within columns followed by the same letter are not significantly different (Fisher's LSD test, P > 0.05).


AC in GF-120 attracted more flies from surround-
ing trees than ammonia from GF-120 alone. This
apparent influx of adults to the test trees did not
increase larval infestations, however, suggesting
flies were trapped or killed before they oviposited.
With respect to larval infestations, one possible
explanation for the lack of differences between
GF-120 and GF-120 + AC or AA treatments is
that ammonia release rates from enhanced drops
decreased quickly after sprays, so after a few days
or even less time the enhanced GF-120 was the
same as GF-120 alone in attractiveness. Ingredi-
ents that prolong ammonia release may be bene-
ficial for control. Perhaps at 100 mL of spray per
tree, flies were able to find drops through normal
foraging even after the drops lost their ammonia.
It was clear that GF-120 is very effective for fly
control, even if it did not eliminate infestations.
Coverage of all single trees in an area or of entire
orchards with GF-120 may lead to even greater
suppression than that obtained by spraying ran-
domly selected single trees as in the present study
or may even eliminate local fly populations over
time. The success of the 2 tests with GF-120
against R. pomonella in this study contrasts with
the bait's failure in New York (Reissig 2003). The
amount of bait spray used in New York was only
32 mL per tree (compared with 100 mL in the
present study), so perhaps this, climatic, and hab-
itat differences explain the inconsistency.
The high levels of control obtained with GF-
120 in experiment 4 were evident, but the use of
100 mL of spinosad alone per tree resulted in sim-
ilarly high control levels, suggesting bait is not
needed with spinosad for a spray to be effective.


Spinosad seems unattractive compared with 40%
GF-120 alone (experiment 3), so its effectiveness
at this volume probably was unrelated to attrac-
tion. Perhaps no or little olfactory or visual stim-
ulation is needed for flies to find the drops over
time, so flies contacted drops while foraging indis-
criminately on leaves. Spinosad probably re-
mained on leaf surfaces long enough for flies to
find them (before being absorbed or broken down).
Some spray also may have landed directly on the
flies. Spinosad drops were smaller than bait drops
and likely covered larger surface areas. In Michi-
gan, spinosad alone (SpinTor) was also as effec-
tive as GF-120 in reducing larval infestation by
R. pomonella in one of two years (Pelz et al. 2005).
Future studies should determine if baits mixed
with spinosad are more critical when spray vol-
umes are <100 mL per tree than when >100 mL
per tree.
In conclusion, overall results indicate the at-
tractiveness of GF-120 to R. pomonella can be in-
creased with added ammonia, but that this does
not necessarily result in greater control, perhaps
because the added ammonia volatilizes too
quickly to make the enhanced GF-120 different
over time than GF-120 alone. Addition of ammo-
nia in GF-120 to increase attractiveness and con-
trol may be more critical at spray volumes lower
than those used in this study. Results here sug-
gest that at the spray volumes used, GF-120 alone
or even spinosad alone can greatly reduce local
R. pomonella populations in Washington, and
should be useful for local horticultural pest and
disease board treatments in residential areas
where organic materials may be more acceptable


December 2007







Yee: GF-120 Effects on Apple Maggot Fly


than organophosphates. Future studies need to
determine how long ammonia-enhanced baits re-
main attractive and if timed release of ammonia
can improve GF-120 performance in eliminating
larval infestations.

ACKNOWLEDGMENTS

I thank Robert Goughnour, Isiah Goughnour, Mera-
lee Nash, and Peter Chapman for field and laboratory
assistance, Blair Wolfley, Lynell Tanigoshi (Washington
State University, Vancouver), and John Stark (Washing-
ton State University Puyallup Research and Extension
Center) for providing research space, Diane Alston
(Utah State University) and Michael Klaus (Washing-
ton State Department of Agriculture) for reviewing the
manuscript, and the Washington Tree Fruit Research
Commission for funding.

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Florida Entomologist 90(4)


December 2007


REVISION OF THE BAMBOO DELPHACID GENUS BELOCERA
(HEMIPTERA: FULGOROIDEA: DELPHACIDAE)

XIANGSHENG CHEN1'23, LIN YANG12 AND JAMES H. TSAI4
'Guizhou Key Laboratory for Plant Pest Management of Mountainous Region,
Guizhou University, Guiyang, Guizhou Province 550025, P. R. China

'Institute of Entomology, Guizhou University, Guiyang, Guizhou Province 550025, P.R. China

'Institute of Zoology, Chinese Academy of Sciences, Beijing 100080, P.R. China

4Fort Lauderdale Research and Education Center, IFAS, University of Florida,
3205 College Avenue, Fort Lauderdale, FL 33314, USA

ABSTRACT

Planthoppers in the genus Belocera Muir, 1913 (Hemiptera: Fulgoroidea: Delphacidae: Del-
phacinae: Tropidocephalini) feed exclusively on bamboo (Poaceae: Bambusoideae). Three
species, B. sinensis Muir (Macao, Taiwan, Guizhou, Hainan, China), B. nigrinotalis Ding and
Yang (Yunnan, China), B. fuscifrons Chen (Guizhou, China), and 2 new species, B. lanpin-
gensis Chen and Yang sp. nov. (Yunnan, China), B. ampelocalama Chen and Tsai sp. nov.
(Guizhou, China) are described in this manuscript. One new synonymy reported, B. huang-
biana Kuoh, 1980 is a junior synonym of B. sinensis Muir, 1913. The male of B. fuscifrons is
reported and described for the first time. The generic characteristics are redefined. The main
morphological features, male genitalia of 5 species and female genitalia of 3 species are de-
scribed or redescribed and illustrated (excluding B. nigrinotalis). A key for identifying the
species of the genus of Belocera is provided.

Key Words: Hemiptera, Fulgoroidea, Delphacidae, Belocera, new species, bamboo pest

RESUME

Los delfacidos del g6nero Belocera Muir, 1913 (Hemiptera: Fulgoroidea: Delphacidae: Del-
phacinae: Tropidocephalini) se alimentan exclusivamente sobre bambu (Poaceae: Bambus-
oideae). Tres species, B. sinensis Muir (Macao, Taiwan, Guizhou, Hainan, China), B.
nigrinotalis Ding y Yang (Yunnan, China), B. fuscifrons Chen (Guizhou, China) y dos nuevas
species, B. lanpingensis Chen y Yang sp. nov. (Yunnan, China), B. ampelocalama Chen y
Tsai sp. nov. (Guizhou, China) son descritas en este trabajo. Se informa de una nueva sino-
nomia, B. huangbiana Kuoh, 1980 es un sin6nimo "junior" de B. sinensis Muir, 1913. Se re-
porta y describe el macho de B. fuscifrons por la primera vez. Las caracteristicas del g6nero
son redefinadas. Las caracteristicas morfol6gicas principles, la genitalia de los machos de
las 5 species y la genitalia de las hembras de las 3 species estan descritas o redescritas e
ilustradas (excludiendo B. nigrinotalis). Se incluye una clave de identificaci6n para las espe-
cies del g6nero Belocera.


The delphacid genus Belocera (Hemiptera:
Fulgoroidea: Delphacidae) was established by
Muir (1913) based on his type species B. sinensis
from Macao, China. It belongs to the tribe Tropi-
docephalini of the subfamily Delphacinae and is
easily separated from other members of this tribe
by the frons, which is widest in the basal 1/3 (Figs.
2, 13, 24, 32), by the antennae with the scape
forked at its apex, sagittate and slightly flattened;
and by the forewing which bear a fuscous median
longitudinal stripe and have a light yellowish
white costal area (Figs. 3-5, 14-15, 25, 33). This
genus is only known to occur in southern China.
To date, 5 species have been described: B. sinensis
Muir, 1913 (Macao; Taiwan: Taipei; Guizhou: Lu-
odian, Wangmo), B. huangbiana Kuoh, 1980
(Hainan: Nada), B. nigrinotalis Ding & Yang,


1986 (Yunnan: Jinghong, Menghai), B. zhejian-
gensis Zhu, 1988 (Zhejiang: Hangzhou; Anhui:
Langyashan, as transferred to the genus Neobelo-
cera by Ding and Hu in 1991) and B. fuscifrons
Chen, 2002 (Guizhou: Libo).
Species of the genus Belocera feed exclusively on
bamboo (Poaceae: Bambusoideae) (Ding et al. 1986;
Yang & Yang 1986; Yang et al. 1999; Chen 2002;
Chen 2003b). Specimens have been collected on the
leaves of several genera of bamboo including Bam-
busa (Yang & Yang 1986; Chen 2002; this paper),
Dendrocalamus (Chen 2002) Neosinocalamus,
Phyllostachys, andAmpelocalamus (this paper).
In this paper we revise the genus Belocera. One
new synonymy reported is B. huangbiana Kuoh,
1980, a junior synonym of B. sinensis Muir, 1913.
Belocera lanpingensis Chen and Yang, and B. am-






Chen et al.: Revision of the Genus Belocera


pelocalama Chen and Tsai, collected, respectively,
from Lanping County, Yunnan Province, and
Dashahe Nature Reserve, Daozhen County,
Guizhou Province, are described as new to sci-
ence. The male of B. fuscifrons Chen, collected
from Fuqian County of Guizhou Province, is re-
ported and described for the first time. The ge-
neric characteristics are redefined. The salient
morphological features, male genitalia of 5 species
and female genitalia of 3 species are described or
redescribed and illustrated (excluding B. nigrino-
talis). A key for identifying the species is provided.

MATERIALS AND METHODS
The methods and morphological terminology
used in this study follow that of Ding et al. (1986),
Yang & Yang (1986) and Chen (2002). The type
specimens and materials examined are deposited
in the Institute of Entomology, Guizhou Univer-
sity, Guiyang, Guizhou Province, PR. China.

DESCRIPTIVE TAXONOMY
Belocera Muir 1913
(Figs. 1-40)
Belocera Muir, 1913: 239. Type species: B. sin-
ensis Muir, 1913, by original designation.


Belocera Muir: Kuoh et al., 1983, Econ. Ins.
Fauna China 27: 43.
Belocera Muir: Yang & Yang, 1986, Taiwan
Mus. Spec. Publ., 6: 42.
Diagnosis. The genus is readily separated from
other Oriental genera of the tribe Tropidocepha-
lini, subfamily Delphacinae, by the frons widest
at basal 1/3, by the antennae with the scape
forked at its apex, sagittate, by the forewings of-
ten with a fuscous central longitudinal fascia, cos-
tal area light yellow or yellowish white, and by
the aedeagus with complex phallobase, phallus
with long spinous processes at apex.
Description. The distinguishing characteristics
outlined by Muir (1913), Kuoh et al. (1983) and
Yang & Yang (1986), are modified as follow:
Coloration. General coloration yellowish brown
to brown. The scape of antennae with two oblique
bands brown to dark brown. Forewings often with
a fuscous central longitudinal fascia, costal area
light yellowish white (Figs. 3-5, 14-15, 25, 33).
Head and Thorax. Head, including eyes (Figs. 1,
12, 23, 31), wider than pronotum (1.10-1.26:1.00).
Vertex wider at base than long submedially about
2.15-2.75:1.00, apical margin transversely broad-
ened and only slightly produced medially. Y-
shaped carina distinct, submedian carinae unit-
ing at apex, apical margin evenly rounding onto


11


a 4


a5


Figs. 1-11. Belocera sinensis Muir. 1. head and thorax, dorsal view; 2. frons and clypeus; 3. forewing (6 Guizhou:
Libo: Maolan); 4. forewing (6 Guizhou: Luodian: Bamao); 5. forewing (9 Guizhou: Luodian: Bamao); 6. hindwing
(6 Guizhou: Luodian: Bamao); 7. male genitalia, caudal view; 8. male genitalia, lateral view; 9. female genitalia,
ventral view; 10. aedeagus, left side; 11. left genital style, lateral view. Scale bars: = 0.5 mm (Figs. 1-2, 9); 1 mm
(Figs. 3-6); 0.2 mm (Figs. 7-8); 0.1 mm (Figs. 10-11).







Florida Entomologist 90(4)


Figs. 12-22. Belocera fuscifrons Chen. 12. head and thorax, dorsal view; 13. frons and clypeus; 14. forewing (6
Guizhou: Fuquqn: Huangsi); 15. forewing holotypee 9, Guizhou: Libo: Maolan); 16. hindwing (6 Guizhou: Fuquqn:
Huangsi); 17. male genitalia, caudal view; 18. male genitalia, lateral view; 19. female genitalia, ventral view; 20.
aedeagus, left side; 21. left genital style, caudal view; 22. left genital style, lateral view. Scale bars: = 0.5 mm (Figs.
12-13, 19); 1 mm (Figs. 14-16); 0.2 mm (Figs. 17-18); 0.1 mm (Figs. 20-22).


frons. Frons (Figs. 2, 13, 24, 32) in median line
longer than wide at widest part about 1.23-
1.35:1.00, widest above level of ocelli, lateral cari-
nae roundly angulate above level of ocelli, then
converging apically, median carina forked at base.
Eyes strongly emarginate on lower margin. Post-
clypeus wider at base than frons at apex. Rostum
surpassing mesotrochanters. Antennae surpass-
ing frontoclypeal suture with scape sagittate,
slightly flattened, in middle line shorter than
width at apex (0.54-0.81:1.00), pedicel cylindrical,
in middle line longer than scape about 1.92-
2.80:1.00. Pronotum (Figs. 1, 12, 23, 31) shorter
than vertex medially (0.71-0.94:1.00), posterior
margin concave medially, with lateral carinae ex-
tending from near the posterolateral angle of the
vertex to the posterior margin of the ronotum,
running as anterolateral margin, curving inward
and reaching hind margin, lateral discs concave.
Mesonotum longer in middle line than vertex and
pronotum combined (1.94-2.43:1.00), median car-
ina reaching the end of scutellum. Forewings nar-
row and elongate long, longer in middle line than
wide at widest part about 3.09-3.56:1.00, broadly
acute at apex with a series of nearly connected
transverse cross veins present at the posterior 1/3.
Hindwings (Figs. 6, 16, 26, 34) with veins M and
Cu, fused for about half their length, M+Cul, and


Culb with a common stalk. Spinal formula of hind
leg 5-6-4, post-tibial spur with an apical tooth.
Male Genitalia. Anal segment of male (Figs. 7,
17, 27, 35) small, ring-like, ventral margin and
more or less concave, without processes or with a
stout, very short process. Pygofer in lateral view
(Figs. 8, 18, 28, 36) with ventral margin longer
than dorsal margin, posterior margin convex, in
caudal view (Figs. 7, 17, 27, 35) with opening
longer than wide, ventral margin concave or with
medioventral process. Genital styles parallel, flat-
tened, in profile (Figs. 11, 22, 30, 40) nearly "7-
shaped", simple, wider at apex than at base, api-
cal portion angulate cephalad. Aedeagus (Figs.
10, 20, 29, 39) with phallobase, phallobase often
developed and complex, twisted and covered on
basal aspect of phallus. Phallus tubular, moder-
ately long, protruding processes of various
lengths from apex. Supporting plate not recogniz-
able. Diaphragm membranous.
Female Genitalia. First valvifers present, sec-
ond valvifers long and large. Ovipositor not over-
passing the pygofer. Gonangulum (Figs. 9, 19, 38)
distinct, with apex acute, blunt or truncate.
Host Plant. Bambusa multiplex (Lour.) Rae-
schel (Yang & Yang 1986), B. emeiensis Chia and
Fung, Dendrocalamus tsiangii (McClure) (Chen
2002), B. sinospinosa McClure, Neosinocalamus


December 2007






Chen et al.: Revision of the Genus Belocera


30


Figs. 23-30. Belocera lanpingensis Chen and Yang sp. nov. 23. head and thorax, dorsal view; 24. frons and
clypeus; 25. forewing (6); 26. hindwing (6); 27. male genitalia, caudal view; 28. male genitalia, lateral view; 29.
aedeagus, left side; 30. left genital style, lateral view. Scale bars: = 0.5 mm (Figs. 23-24); 1 mm (Figs. 25-26); 0.2 mm
(Figs. 27-30).


affinis (Rendle) Keng f.,Ampelocalamus scandons
and Phyllostachys sp.
Distribution. Oriental Region (southern China).
Discussion. This genus is closely related to
Neobelocera Ding and Yang, 1986 (Chen & Liang
2005), which also feeds on bamboo, but differs in
the following: the latter with the antennae mark-
edly flattened, first segment subsagittate, a longi-


tudinal carina down middle, with the apex un-
equally bifurcate, the inner apical angle much
longer than outer apical angle, the postclypeus in
profile, apical part of median carina bend at
rounded, not angled, the rostrum very short, only
reaching mesotrochanters, and the forewing often
with snatchy blackish brown markings, in dark
portion veins bear white spots.


KEY TO SPECIES OF BELOCERA MUIR

1. Frons with apical part bright yellow ............................................................ 2
-Frons yellowish brown or brown, coloration even from base to apex, or with pairs of small light spots (Figs.
13,24) ......................................................... .................... 3
2. Dorsum of body fuscous or blackish brown (Ding & Yang 1986: Fig. 1); furcation of Cu vein of forewings without
dark spot (Ding & Yang 1986: Fig. 7); male pygofer with ventral margin sharply concave, V-like (Ding &
Yang: Fig. 3); genital styles with apices forked (Ding & Yang 1986: Figs. 4-5); phallobase complex, with de-
veloped process (Ding & Yang 1986: Fig. 6) ........................................ B. nigrinotalis
-Dorsum of body light yellowish brown; furcation of Cu vein of forewings with a dark spot (Figs. 3-5); male py-
gofer with ventral margin evenly concave (Fig. 7); genital styles with apices spoon-shaped (Fig. 11); phal-
lobase simple, semicylindrical covered on dorsal aspect of phallus (Fig. 10) ................. B. sinensis
3. Frons evenly brown, without small light spot (Figs. 32); forewings without blackish brown spot (Fig. 33); male
pygofer with 3 medioventral processes on ventral margin (Figs. 35, 37) ............. B. ampelocalamus
-Frons brown to dark brown, with pairs of small light spots (Figs. 13, 24); forewings with a dark brown spot
on furcation of Cu vein and with a small dark brown spot at the end of each longitudinal vein (Figs. 14, 15,
25); without medioventral processes (Figs. 17, 27) ............................................. 4







Florida Entomologist 90(4)


40


37


.39


n35 36 33

Figs. 31-40. Belocera ampelocalamus Chen and Tsai sp. nov. 31. head and thorax, dorsal view; 32. frons and
clypeus; 33. forewing (6); 34. hindwing (6); 35. male genitalia, caudal view; 36. male genitalia, lateral view; 37. py-
gofer, ventral view; 38. female genitalia, ventral view; 39. aedeagus, left side; 40. left genital style, lateral view.
Scale bars: = 0.5 mm (Figs. 31-32, 38); 1 mm (Figs. 33-34); 0.2 mm (Figs. 35-37, 39-40).


4. Body larger (body length of male including forewing 4.40-4.45mm); male anal segment with ventral margin in-
cised medially, without process (Fig. 17); genital styles relatively longer (Figs. 21, 22); phallus with 3 long,
slender spinous processes at apex (Fig. 20) ............... ...................... .B. fuscifrons
-Body smaller (body length of male including forewing 3.30mm); male anal segment with left ventral margin
concave, with a short, stout process on right side (Fig. 27); genital styles relatively shorter (Fig. 30); phallus
with 1 long, slender spinous process at apex (Fig. 29) .............................. B. lanpingensis


Belocera sinensis Muir, 1913
(Figs. 1-11)

Belocera sinensis Muir, 1913: 240.
Belocera huangbiana Kuoh, 1980:198, syn. nov.
Description. Length of body 2.00-2.10 mm
(male), 2.50-2.55 mm (female); including forewing
3.25-3.50 mm (male), 3.75-3.90 mm (female).
Coloration. General coloration yellowish
brown to brown. Eyes dark brown to blackish
brown, ocelli reddish brown. Antennalscape with
2 dark brown stripes. Frons with apical 3/5 bright
yellow, with basal 2/5 yellowish brown to brown.
Genae bright yellow. Postclypeus, vetex, prono-
tum, mesonotum yellowish brown to brown. Tho-
racic and abdominal pleura bright yellowish
white to bright yellow. Legs light gray. Forewings
transparent, with a light yellowish white band
along anterior margin, behind this with a parallel
dark brown longitudinal marking along the ante-
rior margin then curved into the Rs vein, a dark
spot on furcation of Cu vein, at the end of each


longitudinal vein with a small dark brown spot.
Abdomen with dorsum dark brown, sterna yel-
lowish white, lateral areas of each segment with
several small brown spots.
Head and Thorax. Structural features as in ge-
neric descriptions. Head including eyes wider than
pronotum (1.26:1.00). Vertex wider at base than
long submedially about 2.44:1.00. Frons longer in
middle line than wide at widest part about
1.35:1.00. Antennae with scape longer than wide at
apex about 0.54:1.00, shorter than pedicel about
0.52:1.00. Pronotum shorter than vertex (0.88:1.00).
Mesonotum longer than pronotum and vertex com-
bined (1.94:1.00). Forewings longer in middle line
than wide at widest part about 3.17-3.53:1.00.
Male Genitalia. Male anal segment (Fig. 7) with
ventral margin concave medially, without process.
Pygofer in caudal view (Fig. 7) with opening larger
in length than width, ventral margin concave
evenly, in lateral view (Fig. 8) with dorsal margin
emarginate medially. Genital styles in caudal view
(Fig. 7) parallel, inner angle round, outer angle


December 2007









Chen et al.: Revision of the Genus Belocera


strongly produced, spoon-shaped, in lateral view
(Fig. 11) nearly 7-shaped, outer angle acute at
apex, dorsal margin emarginate. Phallus (Fig. 10)
tubular, slender, constricted subapically, apes com-
plex, with a long and twisted process arising from
left dorsal aspect, curved basad then left, another
short and narrow process arising from base of long
one, directed opposite. Phallobase simple, semicy-
lindrical covered on dorsal aspect of phallus.
Female Genitalia. Female pygofer (Fig. 9) with
first valvifers moderately large. Ovipositor as
long as pygofer. Gonangulum small, acute at
apex, separated from first valvifers and apart
from first valvulae.
Material Examined. 4 males and 2 females,
CHINA: Guizhou Province, Luodian, Bamao
(25025'N, 106044'E), 400 m, 2-VIII-1998, X.-S. Chen;
1 male, Guizhou Province, Libo, Maolan (25024'N,
107052'E), 24-V-1998, X.-S. Chen; 13 males and 15
females, Guangdong Province, Guangzhou
(23008'N, 11314'E), 22-XI-2006, X.-S. Chen.
Host Plant. B. multiplex (Yang & Yang 1986),
B. sinospinosa.
Distribution. South China (Macao; Taiwan:
Taipei; Guizhou: Luodian, Libo; Hainan: Nada).
Discussion. This species resembles B. nigrino-
talis, but differs in the following: the latter with
dorsum of body fuscous or blackish brown; furca-
tion of Cu vein of forewings without dark spot;
male pygofer with ventral margin sharply in-
cised, V-like; genital styles with apex forked;
phallobase complex, with developed process.
Based on the similarity of descriptions and the il-
lustrations, Belocera huangbiana Kuoh, 1980,
should be a junior synonym ofB. sinensis Muir.

Belocera nigrinotalis Ding and Yang, 1986

Belocera nigrinotalis Ding and Yang, 1986:
417.
Description. The descriptions are reproduced
from Ding et al. (1986), and redescriptions of male
genitalia are made based on the illustrations of
Ding et al. (1986). "Length of body 1.90 mm
(male), 2.10 mm (female); including forewing 3.30
mm (male), 3.70 mm (female)."
Coloration. "General coloration dark brown to
blackish brown at dorsal aspect, and bright yel-
low in ventral view. Frons with basal part includ-
ing eyes and antennae except base and middle of
first segment bright yellow, are dark brown. An-
tennae with second segment, legs, pleurites be-
tween fore and median legs dark yellowish brown.
Propleura and mesopleura bright yellow. Tegmina
with a yellow band along anterior margin, behind
this with a parallel longitudinal blackish brown
marking along the anterior margin then curved
into the Rs vein, at the end of each longitudinal
vein with a small blackish brown spot. Female
with coloration slightly paler, pronotum and ovi-
positor brown to dark brown."


Head and Thorax. "Head including eyes wider
than pronotum (1.10:1.00). Vertex wider at base
than long submedially about 2.40:1.00. Frons
longer in middle line than wide at widest part
about 1.30:1.00. Antennae with basal segment
longer than wide at apex about 0.73:1.00, shorter
than second segment about 0.44:1.00. Pronotum
shorter than vertex (0.71:1.00). Mesonotum
longer than pronotum and vertex combined
(2.07:1.00). Tegmina longer in middle line than
wide at widest part about 3.09:1.00."
Male Genitalia. Male anal segment (Ding et al.
1986: Fig. 3) ring-like, without process on ventral
margin. Pygofer in caudal view with opening
larger in length than width, ventral margin in-
cised sharply, V-like. Genital styles in caudal view
(Ding et al. 1986: Figs. 4, 5) parallel, forked at
apex, inner angle same size as outer angle. Phal-
lus (Ding et al. 1986: Fig. 6) tubular, very slender,
long, with two spinous processes subapically, di-
rected basad, of which, left one longer and larger.
Phallobase complex, enwinding base of phallus,
with large process on ventral margin.
Material examined. No specimen has been col-
lected by the authors.
Host Plant. Bamboo (Ding et al. 1986).
Distribution. Southwest China (Yunnan: Jing-
hong, Menghai).
Discussion. This species resembles B. sinensis,
but differs in the following: the latter with dor-
sum of body light yellowish brown; furcation of Cu
vein of forewings with a dark spot; male pygofer
with ventral margin concave evenly; genital
styles with apex spoon-shaped; phallobase sim-
ple, semicylindrical.

Belocera fuscifrons Chen, 2002
(Figs. 12-22)

Belocera fuscifrons Chen, 2002: 164.
Description. Length of body 2.50-2.70 mm
(male), 3.00-3.20 mm (female); including forewing
4.20-4.45 mm (male), 5.00-5.30 mm (female).
Coloration. General coloration brown to dark
brown. Eyes dark brown to blackish brown, ocelli
reddish brown. Basal segment of antennae with
two strips blackish brown. Genae with basal part
yellowish brown. Frons, postclypeus, vertex,
pronotum, mesonotum brown to dark brown. Tho-
rax and abdomen with pleura bright yellowish
white to bright yellow. Legs light brown to brown.
Forewings transparent, with a light yellowish
white band along anterior margin, behind this
with a parallel longitudinal dark brown marking
along the anterior margin then curved into the Rs
vein, a dark spot on furcation of Cu vein, at the
end of each longitudinal vein with a small dark
brown spot. Abdomen with dorsum dark brown,
sterna brown medially, lateral areas of each seg-
ment bright yellowish white to bright yellow, with
several dark brown spots. Pygofer yellowish







Florida Entomologist 90(4)


brown to dark brown. Frons of female with 4 pairs
of small light spots beside of median carina.
Head and Thorax. Structural features as in ge-
neric descriptions. Head including eyes wider
than pronotum (1.12:1.00). Vertex wider at base
than long submedially about 2.75:1.00. Frons
longer in middle line than wide at widest part
about 1.35:1.00. Antennae with basal segment
longer than wide at apex about 0.81:1.00, shorter
than second segment about 0.50:1.00. Pronotum
shorter than vertex (0.90:1.00). Mesonotum
longer than pronotum and vertex combined
(2.43:1.00). Tegmina longer in middle line than
wide at widest part about 3.39-3.56:1.00.
Male Genitalia. Male anal segment (Fig. 17)
with ventral margin incised medially, without pro-
cess on ventral margin. Pygofer in caudal view with
opening larger in length than width, ventral mar-
gin concave U-like, in lateral view (Fig. 18) with
dorsal margin slightly emarginated medially, poste-
rior margin near ventral margin produced into a
round process. Genital styles in caudal view (Figs.
17, 21) parallel, inner angle round, outer angle
strongly curved to laterodorsad, in lateral view (Fig.
22) broad at apex, apical margin truncate, outer an-
gle narrowing apically, dorsal margin emarginated.
Phallus (Fig. 20) tubular, slender, constricted sub-
apically, end complex, with 3 spinous processes, pos-
terior one short and thick, left one long and slender.
Phallobase complex, enwinding base of phallus,
ventral margin with a stout process, narrowing api-
cally and curved basally.
Female Genitalia. Female pygofer (Fig. 19)
with first valvifers narrower and smaller. Ovipos-
itor slightly shorter than pygofer. Gonangulum
large, apical margin truncate, connected first
valvifers, approaching first valvulae.
Material Examined. 4 females, CHINA: Guizhou
Province, Libo, Maolan (2524'N, 107'52'E), 550m,
20-V-1998, X.-S. Chen; 2 females holotypee and
paratype), Guizhou Province, Libo, Maolan, 24-25-
V-1998, X.-S. Chen; 6 males, 10 females, Guizhou
Province, Fuquan, Huangsi (2642'N, 107'30'E), 2-
IX-2006, L. Yang and X.-S. Chen.
Host Plant. B. emeiensis, D. tsiangii (Chen
2002).N. affinis.
Distribution. Southwest China (Guizhou: Libo,
Fuquan).
Discussion. This species resembles B. ampelo-
calamus, but differs in the following: the latter
with frons brown evenly, without small light spot;
forewings without any dark brown spot; male py-
gofer with 3 medioventral processes on ventral
margin. It is distinguished from B. sinensis and B.
lanpingensis by larger body size (body length in-
cluding tegmen longer than 4.2 mm).
Belocera lanpingensis Chen and Yang sp. nov.
(Figs. 23-30)
Description. Length of body 2.10 mm (male);
including forewing 3.30 mm (male).


Coloration. General coloration yellowish
brown to brown. Eyes dark brown, ocelli reddish
brown. Basal segment of antennae two blackish
brown stripes. Genae with basal part yellowish
white. Frons and genae with pairs of small light
spots. Frons, postclypeus, vertex, pronotum, and
mesonotum yellowish brown to brown. Thorax
and abdomen with pleura yellowish white. Legs
yellowish brown to brown, with dark brown spots.
Tegmina transparent, with light yellowish white
band along anterior margin, behind this with a
parallel longitudinal dark brown marking along
the anterior margin then curved into the Rs vein,
a dark spot on furcation of Cu vein, at the end of
each longitudinal vein with a small dark brown
spot. Abdomen with dorsum blackish brown,
sterna blackish brown medially, lateral areas of
each segment yellowish white, with several small
dark brown spots. Pygofer yellowish brown.
Head and Thorax. Structural features as in ge-
neric descriptions. Head including eyes wider
than pronotum (1.10:1.00). Vertex wider at base
than long submedially about 2.15:1.00. Frons
longer in middle line than wide at widest part
about 1.23. Antennae with basal segment longer
than wide at apex about 0.63:1.00, shorter than
second segment about 0.36:1.00. Pronotum
shorter than vertex (0.80:1.00). Mesonotum
longer than pronotum and vertex combined
(1.97:1.00). Tegmina longer in middle line than
wide at widest part about 3.47:1.00.
Male Genitalia. Male anal segment (Fig. 27)
with left ventral margin concave, with a stout,
very short process at right side. Pygofer in caudal
view (Fig. 27) with opening larger in length than
width, ventral margin concave U-like, in lateral
view (Fig. 28) with dorsal margin straight, poste-
rior margin near ventral margin produced into an
acute process. Genital styles short, stout and
twisted, in caudal view (Fig. 27) outer margin con-
vex, inner and outer angle round, in lateral view
(Fig. 30) with apical margin truncate, outer angle
narrowing apically, dorsal margin sinuated. Phal-
lus (Fig. 29) tubular, slender, apex acute, with one
thick process subapically on left side, another
spinous process narrow and long, arising from
base of above one, directed base of phallus. Phal-
lobase complex, enwinding base of phallus.
Female. Unknown.
Material Examined. Holotype male, CHINA:
Yunnan Province, Lanping (26'30'N, 99'16'E),
2900 m, 13-VIII-2000, X.-S. Chen.
Host Plant. Phyllostachys sp.
ErL td .'...... The specific name refers to the type
locality, Lanping, Yunnan Province.
Distribution. Southwest China (Yunnan: Lan-
ping).
Discussion. This species resembles B. sinensis,
but differs in the following: the latter with the
apical part of frons bright yellow; male anal seg-
ment without process on ventral margin; male


December 2007







Chen et al.: Revision of the Genus Belocera


pygofer concave evenly; genital styles relatively
long; phallobase simple. It is distinguished from
B. fuscifrons by smaller body size (body length in-
cluding forewing less than 3.5 mm), by male anal
segment with ventral margin incised medially,
and by phallus with 3 spinous processes at apex.

Belocera ampelocalamus Chen and Tsai sp. nov.
(Figs. 31-40)

Description. Length of body 2.4 mm (male), 3.0
mm (female); including forewing 4.40 mm (male),
5.20 mm (female).
Coloration. General coloration brown to dark
brown. Eyes dark brown to blackish brown, ocelli
reddish brown. Basal segment of antennae with 2
dark brown stripes. Genae with basal part yellow-
ish brown. Frons, postclypeus, vertex, pronotum
and mesonotum yellowish brown to brown. Thorax
and abdomen with pleura yellowish white. Legs
yellowish brown to brown. Tegmina transparent,
with a light yellowish white band along anterior
margin, behind this with a parallel longitudinal
dark brown marking along the anterior margin
then curved into the Rs vein, a dark spot on furca-
tion of Cu vein, at the end of each longitudinal
vein with a small dark brown spot. Abdomen with
dorsum dark brown, sterna light brown to brown
medially, lateral areas of each segment yellowish
white, with several small brown spots. Pygofer
light brown. Female with ovipositor dark brown.
Head and Thorax. Structural features as in ge-
neric descriptions. Head including eyes wider
than pronotum (1.17:1.00). Vertex wider at base
than long submedially about 2.56:1.00. Frons
longer in middle line than wide at widest part
about 1.24:1.00. Antennae with basal segment
longer than wide at apex about 0.75:1.00, shorter
than second segment about 0.36:1.00. Pronotum
shorter than vertex (0.94:1.00). Mesonotum
longer than pronotum and vertex combined
(2.24:1.00). Tegmina longer in middle line than
wide at widest part about 3.43:1.00.
Male Genitalia. Anal segment (Fig. 35) with
ventral margin incised medially, without process.
Pygofer in caudal view (Fig. 35), with opening
larger in length than width (2.36:1.00), ventral
margin with 3 medioventral processes (Fig. 37),
pygofer in lateral view (Fig. 36) with dorsal mar-
gin straight, posterior margin convex caudad, sin-
uate. Genital styles moderately long, in caudal
view (Fig. 35) parallel, constricted subapically, in-
ner angle small, outer angle strongly produced,
spoon-shaped, in lateral view (Fig. 40) constricted
at basal 1/3, outer angle acute at apex, dorsal
margin emarginated. Phallus (Fig. 39) tubular,
slender, end complex, with one tear-shaped pro-
cess and three long spinous processes, one long
process arising ventral margin of apex, curved
dorsad, almost reaching base of phallus, another
process arising from right base of long one, curved


ventrad, then left, directed dorsad, the third one
arising from ventral margin of phallus, twisted at
apical 2/3. Phallobase complex, twisted, enwind-
ing base of phallus, a leaf-shaped process arising
from ventral margin, covered middle aspect of
phallus.
Female Genitalia. Female pygofer (Fig. 38)
with first valvifers broad and large. Ovipositor
shorter than pygofer distinctly. Gonangulum
broad and large, apex round and blunt, connected
first valvifers and first valvulae.
Material Examined. Holotype male, CHINA:
Guizhou Province, Daozhen, Dashahe Nature Re-
serve (2853'N, 107'36'E), 640 m, 25-VIII-2004,
X.-S. Chen; 1 female, same data as for holotype.
Host Plant. Ampelocalamus scandons.
ErL td ".'. *. This new species is named after the
generic name of host plant, Ampelocalamus scan-
dons (Bambusoideae).
Distribution. Southwest China (Guizhou:
Daozhen).
Discussion. This species resembles B. fusci-
frons, but differs in the following: the latter with
frons bear several small light spots; forewings
with furcation of Cu vein and the end of each lon-
gitudinal vein with respectively a small dark
brown spot; male pygofer with ventral margin
concave U-like, without medioventral process; fe-
male with first valvifers narrower, apical margin
of gonangulum truncate.

ACKNOWLEDGMENTS

This research was supported by the National Natu-
ral Science Foundation of China (No. 30100015,
30560020), by Program for New Century Excellent Tal-
ents in University, by China Postdoctoral Science Foun-
dation (No. 2005037111), by the Provincial Foundation
for Excellent Youth in Science and Technology Field of
Guizhou (No. 20050520), and by the Nomarch Founda-
tion for Excellent Talents in Science, Technology and
Education Field of Guizhou (No. 2005357). This re-
search was also supported by the Florida Agricultural
Experiment Station.

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Technology Publishing House (in Chinese with Eng-
lish summary).
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cera (Homoptera: Delphacidae) from China. Zootaxa
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CHEN X.-S. 2003b. Key to genera of the tribe Tropido-
cephalini from the People's Republic of China with
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Chen et al.: The Genus Arcofacies from China


REVIEW OF THE BAMBOO DELPHACID GENUS ARCOFACIES
(HEMIPTERA: FULGOROIDEA: DELPHACIDAE) FROM CHINA,
WITH DESCRIPTION OF ONE NEW SPECIES

XIANGSHENG CHEN' 2 LIN YANG'2 AND JAMES H. TSAI4
'Guizhou Key Laboratory for Plant Pest Management of Mountainous Region,
Guizhou University, Guiyang, Guizhou Province 550025, P.R. China

2Institute of Entomology, Guizhou University, Guiyang, Guizhou Province 550025, P.R. China

'Institute of Zoology, Chinese Academy of Sciences, Beijing 100080, P.R. China

4Fort Lauderdale Research and Education Center, IFAS, University of Florida,
3205 College Avenue, Fort Lauderdale, FL 33314, USA

ABSTRACT

Four species in the genus Arcofacies Muir, 1915 in China (Hemiptera: Fulgoroidea: Delpha-
cidae: Delphacinae: Tropidocephalini) feeding exclusively on bamboo (Bambusoideae), are
reviewed. The 4 species are A. fullawayi Muir, 1915 (Fujian: Wuyishan; Taiwan: Taibei,
Gaoxiong, Nantou; Sichuan; Hainan; Guizhou: Luodan, Wangmo), A. maculatipennis Ding,
1987 (Guizhou: Luodian, Guiyang, Xishui, Chishui, Daozhen, Changshui, Fuqian),
A. strigatipennis Ding, 1990 (Fujian: Wuyishan), and A. ampelocalamus Chen sp. nov.
(Guizhou: Daozhen). The main morphological characters and male genitalia of the 4 species
are described or redescribed and illustrated. A key for identifying the species of Arcofacies
from China is provided. The importance of these as a pest on bamboo is discussed briefly, and
biological notes of the new species are given.

Key Words:Arcofacies, bamboo delphacids, Hemiptera, Fulgoroidea, new species, China

RESUME

Se revisa cuatro species del genero Arcofacies Muir, 1915 de la China (Hemiptera: Fulgo-
roidea: Delphacidae: Delphacinae: Tropidocephalini) alimentandose exclusivamente sobre
bambu (Bambusoideae). Las 4 species son: A. fullawayi Muir, 1915 (Fujian: Wuyishan;
Taiwan: Taizhong, Taibei, Gaoxiong, Nantou; Sichuan; Hainan; Guizhou: Luodan, Wangmo),
A. maculatipennis Ding, 1987 (Guizhou: Luodian, Guiyang, Xishui, Chishui, Daozhen,
Changshui, Fuqian), A. strigatipennis Ding, 1990 (Fujian: Wuyishan), y A. ampelocalamus
Chen sp. nov. (Guizhou: Daozhen). Se describe o redecribe e ilustra las caracteristicas mor-
fol6gicas principles de la genitalia de los machos de las 4 species. Se incluye una clave para
la identificaci6n de las species de Arcofacies de la China. Se discute brevemente la impor-
tancia de estas como plagas de bambu y se incluye notas biol6gicas sobre la nueva especie.


The delphacid genus Arcofacies was established
by Muir (1915) based on specimens from Manila,
the Philippines (type species: Arcofacies fullawayi
Muir, 1915). It belongs to the tribe Tropidocephalini
within subfamily Delphacinae (Hemiptera: Fulgor-
oidea: Delphacidae) and is easily separated from
other members in this tribe by the postclypeus at
right angle to frons (Fig. 3), by a white median lon-
gitudinal line extending from the apex of the frons
to end of the mesonotum, along the line bordered
with black or brown stripe (Figs. 1, 10, 18, 24), and
by the forewings often with blackish brown mark-
ings, in dark portion veins bear white spots (Figs. 4,
12, 26). It is known to occur in the Oriental region.
Six species have been recorded worldwide (Muir
1915, 1919; Fennah 1973-1975; Ding 1987, 1990),
occurring in the Philippines (2 species:A. fullawayi


Muir, 1915, A. insignis Muir, 1919), Singapore (1
species:A. fullawayi), Malaysia (2 species:A. fulla-
wayi,A. penangensis Muir, 1919), Indonesia (1 spe-
cies:A. fullawayi), Sri Lanka (1 species:A. truncati-
pennis Fennah, 1973-1975) and China (3 species:A.
fullawayi, A. maculatipennis Ding, 1987, A.
strigatipennis Ding, 1990).
Species of Arcofacies from China feed exclu-
sively on bamboo (Bambusoideae) (Ding 1987,
1990; Yang & Yang 1986; Yang et al. 1999; Chen
2003). Specimens were collected on the leaves of
several genera of bamboo, Bambusa (Yang & Yang
1986), Neosinocalamus and Ampelocalamus (this
paper). Arcofacies fullawayi Muir, A. maculati-
pennis Ding andA. ampelocalamus sp. nov. are of
economic significance due to large populations
feeding on the bamboo in the fields.







Florida Entomologist 90(4)


8


9


Figs. 1-9.Arcofacies fullawayi Muir. 1. head and thorax, dorsal view; 2. frons and clypeus; 3. head and thorax,
lateral view; 4. forewing; 5. hindwing; 6. male genitalia, posterior view; 7. male genitalia, lateral view; 8. aedeagus,
lateral view; 9. right genital style, lateral view. Scale bars = 0.5 mm (Figs. 1-3); 1 mm (Figs. 4-5); 0.2 mm (Figs. 6-
7); 0.1 mm (Figs. 8-9).


In this paper we review the Chinese species of
the genus Arcofacies. Arcofacies ampelocalamus
Chen, collected from Dashahe Nature Reserve,
Daozhen, Guizhou Province, is described as new
to science. The main morphological characters
and male genitalia of 4 species are described and
illustrated in detail. A key to all species found in
China is provided.

MATERIALS AND METHODS

Morphological techniques and terminology fol-
low Yang & Yang (1986), and Ding (1990). Speci-
mens examined are deposited in the Insect Collec-
tion at the Institute of Entomology, Guizhou Uni-
versity, Guiyang, Guizhou Province, China
(IEGU).

DESCRIPTIVE TAXONOMY

Arcofacies Muir

Arcofacies Muir, 1915, Can. Ent., 47: 319. Type
species: Arcofacies fullawayi Muir, 1915, by origi-
nal designation.
Arcofacies Muir: Kuoh et al., 1983, Econ. Ins.
Fauna China, 27: 45.
Arcofacies Muir: Yang & Yang, 1986, Mus.
Spec. Publ. Seri., No. 6: 34.


Arcofacies Muir: Ding, 1990, Journal of Bam-
boo Research, 9: 74.
Arcofacies Muir: Ding et al., 1999, Fauna of In-
sects in Fujian Province of China, 2: 442.
The distinctive characters used by Muir
(1915), Yang & Yang (1986), Ding (1990) and Ding
et al. (1999) are modified as follows:
General color yellowish green to yellowish
brown. A white median longitudinal line extends
from the apex of the frons to the end of mesono-
tum, along the line bordered with dark brown or
black. Lateral parts of pronotum each with ob-
lique white band bordered with brown or dark
brown. Forewings with light brown in basal third,
apical portion hyaline, speckled with dark brown
markings, in dark portion veins bear white spots.
Hindwings hyaline with brown veins.
Head including eyes narrower than pronotum.
Vertex (Figs. 1, 10, 18, 24) trapeziform, with mar-
gins more or less well defined, wider at base than
long submedially (1.70-1.88:1), apical margin dis-
tinctly emarginate at both sides of median point,
lateral carinae concave, submedian carinae trans-
verse. Y-shaped carina without stalk, with very
short arms, connecting submedian carinae which
forms a small cell, in lateral view (Fig. 3) vertex
and frons at right angle. Frons (Figs. 2, 11, 19, 25)
in middle line longer than wide at widest point
(1.75-2.17:1), widest at level of ocelli or at apex,


December 2007


. I







Chen et al.: The Genus Arcofacies from China


16


Figs. 10-17.Arcofacies maculatipennis Ding. 10. head and thorax, dorsal view; 11. frons and clypeus; 12. fore-
wing; 13. hindwing; 14. male genitalia, posterior view; 15. male genitalia, lateral view; 16. aedeagus, lateral view;
17. left genital style, lateral view. Scale bars = 0.5 mm (Figs. 10-11); 1 mm (Figs. 12-13); 0.2 mm (Figs. 14-15); 0.1
mm (Figs. 16-17).


lateral carinae convex at base, nearly straight be-
low level of ocelli, median carina not well devel-
oped throughout, forked at extreme base. Post-
clypeus slightly wider at base than frons at apex,
at right angle to frons (Fig. 3), tricarinate. Ros-
trum almost extending to mesotrochanters. Eyes
in dorsal view with lateral margin emarginated
medially. Lateral ocelli present. Antennae cylin-
drical, scape distinctly longer than wide (1.60-
2.00:1), shorter than pedicel (0.52-0.59:1). Prono-
tum with lateral carinae extending to hind mar-
gin, converging apically, median carina weak.
Forewings tectiform at rest. M and Sc1of wing with
a long common stalk, Cu, arising from end of cross
vein or basad. Spinal formula of hind leg 5-6-4.
Anal segment of male (Figs. 6, 14, 20, 28, 30)
collar-shaped, lateroapical angles produced into
spinous processes or not. Pygofer (Figs. 6, 14, 20,
28) in posterior view with opening longer than


wide (1.29-1.6:1), lateral margins (Figs. 7, 15, 21,
29) strongly produced caudad medially or not,
with a small medioventral process or not. Aedea-
gus tubular or flat, with spinous process or not,
orifice subapical. Diaphragn armature sclerotized
and pigmented, V-shaped. Diaphragm wide,
membraneous. Genital styles (Figs. 9, 17, 23, 32,
33) long, simple, broad at base, narrowing api-
cally, basal angle intumescent, apex twisting out-
ward more or less.
Host Plant. Bambusa multiplex (Lour.) Raeu-
schel, B. oldhamii Munro, B. Multiplex Raeuschel
cv. "Fernleaf" Young (Yang & Yang, 1986), Neosi-
nocalamus affinis (Rendle) Keng f; Ampelocala-
mus scandons (Hsueh and Li) Chen, Wen and
Sheng.
Distribution. Oriental Region (China, the Phil-
ippines, Malaysia, Indonesia, Singapore, Sri
Lanka).


KEY TO SPECIES OF ARCOFACIES MUIR FROM CHINA

1. Lateral carinae of frons, gena and pronotum yellowish brown, without white line (Figs. 1, 2, 10, 11); apical half
of forewing with blackish brown markings (Figs. 4, 12); aedeagus simple, without elongate spinous process
(Figs. 8, 16).................... ..................................... ............. 2
-Lateral carinae of frons, gena and pronotum bordered with white lines (Figs. 18, 19, 24, 25); apical veins of
forewing bordered with brown stripes (Fig. 26); aedeagus with long spinous process at middle or at base
(Figs. 22, 31).......................................................................... 3


W 10







Florida Entomologist 90(4)


December 2007


Figs. 18-23.Arcofacies strigatipennis Ding. 18. head and thorax, dorsal view; 19. frons and clypeus; 20. male gen-
italia, posterior view; 21. male genitalia, lateral view; 22. aedeagus and genital styles, lateral view; 23. left genital
style, posterior view. (All figures are reproduced from Ding, 1990.)


2. Median carina of postclypeus dark brown (Fig. 2); anal segment of male with lateroapical angles truncate, without
spinous process (Figs. 6, 7); aedeagus tubular, acute at apex, dorsal margin concave (Fig. 8); genital styles in
lateral view with base relatively narrow (Fig. 9). ...................................... A. fullawayi
-Median carina of postclypeus white (Fig. 11); anal segment of male with lateroapical angles produced
into stout spinous process (Figs. 14, 15); aedeagus tubular, blunt at apex, bent ventrad at middle, dor-
socaudal margin with several teeth (Fig. 16); genital styles in lateral view with base relatively broad
(Fig. 17) ................................................................. A m aculatipennis
3. Pygofer without medioventral process (Fig. 20); in lateral view caudal margin concave, near ventrocaudal
margin produced into a round protuberance (Fig. 21); aedeagus broad and flat at base, near quadrate,
apex acute, thumb-shaped, two spinous processes arising from dorsal margin and ventral margin (Fig.
22).................................. ... ............... ............ . ...A strigatipennis
-Pygofer with small medioventral process, flake-shaped (Fig. 28); in lateral view caudal margin nearly
straight (Fig. 29); aedeagus with base coniform, apex round and blunt, a long spinous processes arising
from left base (Fig. 31). ..................................................... A. ampelocalamus


Arcofacies fullawayi Muir
(Figs. 1-9)

Arcofacies fullawayi Muir, 1915, Can. Ent., 47:
320.
Arcofacies fullawayi: Muir, 1919, Philip. Jour.
Sci., 15: 526.
Arcofacies fullawayi: Fennah, 1956, Proc. Ca-
lif. Acad. Sci., 28(4): 465.
Arcofacies fullawayi: Kuoh et al., 1983, Econ.
Ins. Fauna China, 27: 45.
Arcofacies fullawayi: Ding, 1990, Journal of
Bamboo Research, 9: 75.


Arcofacies fullawayi: Ding et al., 1999, Fauna
of Insects in Fujian Province of China, 2: 442.
Arcofacies fullawayi: Chen, 2002, Insects from
Maolan Landscape, 158.
Description. Body length (from apex of vertex
to tip of forewing): male 3.40-3.45 mm, female
3.65-3.95 mm.
General color yellowish brown with green. A
white median line from apex of frons to posterior
aspect of mesonotum, bordered with blackish
brown (Figs. 1-2); eyes dark brown to blackish
brown; ocelli dark brown infused with red; anten-
nae with middle and apex of scape, base and near







Chen et al.: The Genus Arcofacies from China


32


33


Figs. 24-33. Arcofacies ampelocalamus Chen, sp. nov. 24. head and thorax, dorsal view; 25. frons and clypeus;
26. forewing; 27. hindwing; 28. male genitalia, posterior view; 29. male genitalia, lateral view; 30. anal segment,
posterior-ventral view; 31. aedeagus, lateral view; 32. left genital style, lateral view; 33. left genital style, posterior
view. Scale bars = 0.5 mm (Figs. 24-25); 1 mm (Figs. 26-27); 0.2 mm (Figs. 28-33).


apex of pedicel ring with dark brown to blackish
brown (Fig. 2); lateral parts of pronotum each
with oblique white band bordered with blackish
brown (Figs. 1, 3); forewings with yellowish brown
over basal third, apical two-thirds hyaline, speck-
led with blackish brown markings as in Fig. 4, in
dark portion veins bear white spots; wings hya-
line with brown veins; legs with fore and median
tibiae pale reddish orange; abdomen with dorsum
of VII-IX segments dark brown, pygofer blackish
brown; coloration of female same as male except
lateral and ventral parts of thorax, abdomen pale
green, ovipositor yellowish brown.
Vertex wider at base than long submedially
about 1.70:1. Frons longer than wide at widest
part (about 1.75:1), widest just above ocelli. An-
tennae surpassing frontoclypeal suture, scape
longer than wide (about 2.00:1), shorter than
pedicel (about 0.59:1).
Male Genitalia. Anal segment of male short,
ring-like, with a big gap at ventral margin, lateroap-
ical angles truncate. Pygofer in posterior view with
opening larger in length than width (about 1.33:1)
(Fig. 6), in lateral view posterior margin strongly
produced caudad near ventral aspect (Fig. 7). Aede-
agus simple, tubular, strongly concave on dorsal
margin, acute at apex, directed dorsad (Fig. 8). Gen-
ital styles long, slender, rounded at base, abruptly
narrowing and slightly twisted near apex (Fig. 9).


Host Plant. Bambusa multiplex (Lour.) Raeus-
chel; B. oldhamii Munro; B. multiplex Raeuschel
cv. "Fernleaf" Young (Yang & Yang, 1986); Neosi-
nocalamus affinis (Rendle) Keng f.
Distribution. China (Fujian, Taiwan, Chong-
qing, Hong Kong, Hainan, Guizhou), the Philip-
pines, Malaysia, Indonesia, Singapore.
Specimens Examined. 2 males, 2 females,
CHINA: Guizhou, Wangmo, Sanglang, 31-VII-
1998 (X.-S. Chen); 1 male, 1 female, Guizhou, Lu-
odian, Bamao, 2-VIII-1998 (X.-S. Chen), 1 male, 2
females, Taiwan, Nantou, Wushe, 850 m, 24-XI-
2002 (X.-S. Chen) (IEGU).

Arcofacies maculatipennis Ding
(Figs. 10-17)

Arcofacies maculatipennis Ding, 1987, Acta
Entomologica Sinica, 30(4): 439.
Arcofacies maculatipennis: Chen and Yang,
2005, Insects from Dashahe Nature Reserve of
Guizhou, 122.
Arcofacies maculatipennis: Chen, 2005, In-
sects from Xishui Landscape, 153.
Arcofacies maculatipennis: Chen, 2006, Insects
from Chishui Spinulose Tree Fern Landscape, 119.
Description. Body length (from apex of vertex
to tip of forewing): male 3.75-4.00 mm, female
4.15-4.40 mm.







Florida Entomologist 90(4)


General color yellowish brown with green. A
white median line from apex of frons to end of me-
sonotum bordered with blackish brown (Figs. 10-
11); eyes dark brown to blackish brown; ocelli red-
dish brown; antennae with middle and apex of
scape, base and near apex of pedicel ring with
dark brown (Figs. 11); lateral parts of pronotum
each with oblique white band bordered with
blackish brown (Fig. 10); Forewings with yellow-
ish brown over basal third, rest area hyaline,
speckled with blackish brown markings as fig-
ured (Fig. 12), in dark portion veins bear white
spots; wings hyaline with brown veins; legs with
fore and median tibiae and tarsi, hind tarsi pale
reddish orange; abdomen with dorsum of VII-IX
segments dark brown to reddish brown, ventral
areas pale reddish orange, pygofer blackish
brown; coloration of female same as male except
lateral and ventral parts of thorax, abdomen
green, ovipositor yellowish brown.
Vertex wider at base than long submedially
about 1.70:1. Frons longer in middle line than
wide at widest part about 2.17:1, widest at level of
ocelli. Antennae surpassing frontoclypeal suture,
scape longer than wide about 1.80:1, shorter than
pedicel about 0.53:1.
Male Genitalia. Anal segment of male short,
ring-like, lateroapical angles each produced into
stout process, acute at apex. Pygofer in posterior
view with opening larger in length than width
about 1.60:1 (Fig. 14), in lateral view ventral an-
gles strongly produced (Fig. 15). Aedeagus simple,
tubular, strongly bent ventrad, dorsal margin con-
vex, with about 19-21 small teeth on dorsoposte-
rior margin, obtuse at apex. Genital styles moder-
ately long, approach ventral margin of anal seg-
ment (Figs. 14-15), broad at base, abruptly narrow-
ing to apex, slightly twisted near apex (Fig. 17).
Host Plant. Neosinocalamus affinis (Rendle)
Keng f.
Distribution. Southwest China (Guizhou).
Specimens Examined. 1 male, 4 females,
Guizhou, Daozhen, 25-VII-1984 (Z.-Z. Li); 1 male, 3
females, CHINA: Guizhou, Xishui, Linjiang, 1-VI-
2000 (X.-S. Chen); 6 males, 1 female, Guizhou, Guiy-
ang, Huanxi Park, 25-VII-1998, 1050 m (X.-S.
Chen); 3 males, 1 female, Guizhou, Guiyang, Hua-
nxi, 28-VIII-1998, 1050 m (X.-S. Chen); 3 males, 6
females, Guizhou, Chishui, 20-IX-2000 (X.-S. Chen);
11 males, 21 females, Guizhou, Changshun, 24-VII-
2006 (X.-S. Chen); 8 males, 13 females, Guizhou,
Fuquan, 11-VIII-2006 (X.-S. Chen and L. Yang).

Arcofacies strigatipennis Ding
(Figs. 18-23)

Arcofacies strigatipennis Ding, 1990, Journal
of Bamboo Research, 9(1): 75.
Arcofacies strigatipennis: Ding, Huang, and
Zhuo, 1999, Fauna of Insects in Fujiang Province
of China, 2: 443.


The description and illustration are repro-
duced from Ding (1990), Ding et al. (1999).
"Body length (from apex of vertex to tip of
forewing): male 3.00 mm, female 3.50 mm."
"General color pale yellowish brown with
somewhat green. A white median line from apex
of frons to end of mesonotum, along lateral cari-
nae of frons, gena, and pronotum with white line;
forewings with pale brown over basal third, rest
area hyaline, along apical veins bordered brown
stripes, in dark portion veins bear white spots;
male with abdomen most blackish brown, female
with abdomen most yellowish brown, ovipositor
brown."
"Anal segment of male in lateral view lateroap-
ical angles each produced into process. Pygofer in
posterior view with opening longer than wide
(Fig. 20), in lateral view posterior margin con-
cave, ventral angles strongly produced (Fig. 21).
Aedeagus broad at base, nearly quadrate, apex
finger-like, middle of dorsal margin and ventral
margin each with a process, the dorsal one slen-
der and curving. Genital styles divergent (Fig.
20), long, attaining ventral margin of anal seg-
ment, rounded at base, narrowing to apex,
twisted near apex (Figs. 22-23)."
Host Plant. Bamboo (Ding 1990).
Distribution. South China (Fujian).
Specimen Examined. No specimen has been
collected by the authors.

Arcofacies ampelocalamus Chen, sp. nov.
(Figs. 24-33)

Description. Body length (from apex of vertex
to tip of forewing): male 2.90-4.00 mm, female
3.30-4.40 mm.
General color pale yellowish brown. Frons,
clypeus, gena, vertex, pronotum and mesonotum
brown to dark brown, a white median line from
apex of frons to end of mesonotum, along lateral
carinae of frons, postclypeus, gena, vertex and
pronotum with white line (Figs. 24-25); eyes dark
brown to blackish brown; ocelli reddish brown;
antennae with dorsal and ventral margins and
apex of scape, base and near apex of pedicel dark
brown to blackish brown (Fig. 25); lateral parts of
pronotum each with oblique white band (Fig. 24);
forewings with pale brown over basal third, rest
area hyaline, along transverse vein and apical
veins bordered brown stripes as figured (Fig. 26),
in dark portion veins bear white spots; wings hy-
aline with pale brown veins; legs with fore and
median tibiae pale reddish orange; abdomen pale
yellowish green, pygofer blackish brown; colora-
tion of female same as male except lateral and
ventral parts of thorax, abdomen including ovi-
positor pale green.
Vertex wider at base than long submedially
about 1.88:1. Frons longer in middle line than
wide at widest part about 1.88:1, widest at apex.


December 2007







Chen et al.: The Genus Arcofacies from China


Antennae surpassing frontoclypeal suture, scape
longer than wide at apex about 1.60:1, shorter
than pedicel about 0.52:1.
Male Genitalia. Anal segment of male short,
ring-like, lateroapical angles each produced into
stout process, relative obtuse at apex. Pygofer in
posterior view with opening larger in length than
width about 1.29:1, in ventral view with a small
medioventral process (Fig. 28), in lateral view
posterior margin nearly straight, ventral angles
slightly produced (Fig. 29). Aedeagus tubular,
with base coniform, apex round and blunt, middle
ventral margin with a small process, a long
spinous processes arising from left base, then
strongly bent ventrad (Fig. 31). Genital styles
moderately long, approach ventral margin of anal
segment, moderately broad at base, narrowing to
apex, slightly twisted near apex (Figs. 32-33).
Host Plant. Ampelocalamus scandons (Hsueh
and Li) Chen, Wen and Sheng.
ErP- *..1. This new species is named after the
generic name of host plant, Ampelocalamus scan-
dons (Poaceae: Bambusoideae).
Distribution. Southwest China (Guizhou).
Specimens Examined. Holotype male, CHINA:
Guizhou, Daozhen, Dashahe, Xiannudong, 600 m,
26-V-2004 (X.-S. Chen). Paratype 16 males, 24 fe-
males, same data as holotype; 2 males, 8 females,
Guizhou, Daozhen, Dashahe, Xiannudong, 660m,
25-VIII-2004 (X.-S. Chen).
Biology. This species maybe has two biological
forms, the smaller form (body length including
tegmen male 2.90 mm, female 3.30-3.50 mm),
feeding on a native bamboo,Ampelocalamus scan-
dons (Hsueh and Li) Chen, Wen and Sheng, with
August as the probable peak periods; the larger
form (body length including tegmen: male 3.75-
4.00 mm, female 4.15-4.40 mm), feeding on a na-
tive bamboo, Neosinocalamus affinis (Rendle)
Keng f, with May as the probable peak periods, so
is its population peak.
Remarks. This species is similar to A. strigati-
pennis Ding, but differs as follows: frons widest at
apex (widest at level of ocelli in the latter); pygofer
in ventral view with short medioventral process (no
medioventral process in the latter), in lateral view
pygofer with posterior margin straight (posterior
margin concave in the latter); two genital styles
closer (two genital styles divergent in the latter);
aedeagus coniform at base, round and blunt at apex
quadratee at base, acute at apex in the latter).

ACKNOWLEDGMENTS

This research was supported by the National Natu-
ral Science Foundation of China (No. 30100015,
30560020), by Program for New Century Excellent Tal-


ents in University, by China Postdoctoral Science Foun-
dation (No. 2005037111), by the Provincial Foundation
for Excellent Youth in Science and Technology Field of
Guizhou (No. 20050520), and by the Nomarch Founda-
tion for Excellent Talents in Science, Technology and
Education Field of Guizhou (No. 2005357). This re-
search was also supported by the Florida Agricultural
Experiment Station.


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Florida Entomologist 90(4)


December 2007


THE PLANTHOPPER GENUS ACANALONIA IN FLORIDA WITH NOTES
ON A RECENTLY INTRODUCED SPECIES, A. EXCAVATA
(HEMIPTERA: FULGOROIDEA: ACANALONIIDAE)

SUSAN HALBERT', LOIS B. O'BRIEN2 AND STEPHEN W. WILSON3
'Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville, FL

'Visiting Scholar, Department of Entomology, University of Arizona, Tucson, AZ

'Department of Biology, University of Central Missouri, Warrensburg, MO

ABSTRACT

Acanalonia excavata Van Duzee, described from Nicaragua, has been found at 4 localities in
Florida. Although 6 species ofAcanalonia have been reported from Florida, 2 species known
from single disjunct records, A. concinnula Fowler andA. uirescens Stal, are either in error,
or finds have not been replicated. A key for the identification of the resulting 5 species known
to occur in Florida is provided. The male and female genitalia ofA. excavata are illustrated,
and the placement of this species in the key to the United States species is indicated.

Key Words: Acanalonia, Fulgoroidea, Acanalonia excavata, new record, Florida

RESUME

Acanalonia excavata Van Duzee, descrita de Nicaragua, fue encontrada en 4 sitios en la Flo-
rida. Aunque 6 species deAcanalonia han sido reportadas en la Florida, 2 de estas species
son conocidas de un solo registro incongruente, A. concinnula Fowler yA. uirescens Stal; es-
tos pueden representar un error o hallazgos que no han sido replicados. Se provee una clave
para la identificaci6n de las 5 species resultantes conocidas que se sabe ocurren en la Flo-
rida. Se ilustra la genitalia del macho y la hembra deA. excavata, y se indica la posici6n de
esta especie en la clave para las species en los Estados Unidos.


Six species of the planthopper genus Acana-
lonia (Hemiptera: Fulgoroidea: Acanaloniidae)
have been recorded from Florida: Acanalonia
bivittata (Say), A. concinnula Fowler, A. conica
(Say), A. pumila Van Duzee, A. seruillei Spinola
1839 (=A. latifrons (Walker 1851)), and A. uire-
scens Stal (Metcalf 1954; Freund & Wilson 1995).
A seventh species, A. excavata Van Duzee, has
been collected in 4 sites in Florida and represents
a new record for the United States. Fruend & Wil-
son (1995) reviewed the species of Acanalonia
known to occur in the United States. The purpose
of this paper is to note the introduction of a spe-
cies new to the United States, to provide a key to
the Florida species that includes A. excavata, and
to evaluate and correct published distribution
records for Florida.
Acanalonia excavata was described from Nica-
ragua by Van Duzee (1933) and has not been re-
ported from any other locality since its original
description. This species recently has been col-
lected on 4 occasions in south Florida. Data for
these specimens are as follows: FLORIDA: Mi-
ami-Dade County, Kendall, 25 IV 1997, coll. J. R.
Martin (1 female); Coral Gables, 6 V 2000, coll. J.
Brambila (1 male); Florida City, 9 VI 2004, coll. E.
T Putland (1 female); Miami, 25 X 2004, coll. E. T


Putland (1 female). Specimens are housed in the
Florida State Collection of Arthropods, Division of
Plant Industry, Florida Department of Agricul-
ture and Consumer Services, Gainesville. Acana-
lonia excavata can be separated from the 18 spe-
cies of United States Acanalonia by the produced
head, pubescent frons, the presence of a strongly
curved spine on the left side of the aedeagus, and
the shape of the posterior margin of the female
terminal abdominal sternite (Figs. 1, 2). This spe-
cies will key to couplet 7 in Freund & Wilson's
(1995) key to the Acanalonia species of the United
States but can be separated from the similar spe-
cies A. conica, A. clypeata, and A. saltonia by the
characters of the head, aedeagus, and female ven-
ter. For comparative purposes, the male and fe-
male genitalia are illustrated in Fig. 2 from spec-
imens with the following collecting data: NICA-
RAGUA: 10-16 km W Managua, 18 X 1970, coll.
E. Moore (male); Managua, 17 VII 1970, coll. L. H.
Rolston (female). Acanalonia excavata is likely to
be a recent introduction into Florida from Central
America. Nothing is known about the biology of
this planthopper; however, the other Acanalonia
species that have been studied are widely poly-
phagous, particularly on woody plants (Wilson &
McPherson 1980, 1981; Freund & Wilson 1995).







Halbert et al.: Planthopper Genus Acanalonia in Florida


plnapaphs by David Zicsk

Fig. 1.Acanalonia excavata. A. Habitus, dorsal view.
B. Habitus, lateral view.


Acanalonia concinnula was described from
Mexico by Fowler (1900) and reported from Ven-
ice, Florida by Ball (1933). The single male speci-
men, upon which the Florida record was based
(housed at the National Museum of Natural His-
tory, Smithsonian Institution, Washington, D. C.),
was misidentified, as determined by comparison
of China's illustrations of the holotype (housed at
the Museum of Natural History, London, UK) pub-
lished by Doering (1932). The Florida specimen is
actually a pale form ofA. bivittata. Thus, there is
no evidence that this Mexican species occurs in
Florida.Acanalonia concinnula is known from the
states of Jalisco, Guerrero, Sinoloa, and Puebla in
Mexico (Metcalf 1954, L.B.O., unpublished data).
Acanalonia concinnula was recorded from Texas
by Melichar (1901) whose specimens are supposed
to be housed in museums in Stockholm, Paris, and
Brussels. We know of no US specimens of this spe-
cies in any collections in the United States.
Acanalonia virescens was described from Mex-
ico by Stal (1864) and reported from Marco, Flor-
ida by Ball (1933). The male specimen upon which
this record is based has not been found; thus, the
presence of this species in Florida cannot be veri-
fied. Doering (1932) recorded this species only
from Texas. There is no evidence thatA. virescens
occurs in Florida.
There has been taxonomic confusion about the
status ofA. latifrons (described from New Orleans
LA, USA) and A servillei (described from Phila-
delphia PA, USA). Fennah (1971) determined
that the type ofA. latifrons corresponded to the
description ofA. servillei. However, it is doubtful
that Fennah was able to examine the type ofA.
servillei because it was housed in Spinola's castle
of Tassarolo until 1979 when it was moved to the
Museo Regionale di Scienze Naturali in Turin,
Italy (Casale 1981). Doering (1932) and Metcalf &


Fig. 2.Acanalonia excavata. A. Head, lateral view. B.
Male, pygofer, anal tube, and aedeagus, left lateral view.
C. Male, genital style, left lateral view. D. Female, ter-
minal segments of abdomen, lateral view. E. Female,
terminal segments of abdomen, ventral view. Arrows in-
dicate diagnostic features (see text). Bar = 1 mm. Draw-
ing by Stephen W. Wilson.


Bruner (1930) used a specimen from Cuba as the
basis for their concept of A. servillei (Fennah
1971). Fennah believed that this specimen was
too large (13-15 mm) to be what Spinola described
as A. servillei, which was 8.5 mm long. According
to Fennah's (1971) description and key, theA. ser-
villei of Metcalf & Bruner (1930) and Doering
(1932) probably was A. ingens (Fennah). Ball
(1933) also synonymized A. servillei and A. lati-
frons stating "[there] is certainly but a single
large blunt-headed species of this genus occur-
ring in the United States." He compared speci-
mens from Philadelphia, the type locality of A.
servillei, with specimens from New Orleans, the
type locality ofA. latifrons, and found them to be
the same species. However, he did not examine
the types. He also synonymizedA. servillei with a
species from "Hayti", which was later found to be
distinct. Metcalf (1954) apparently ignored the
synonymy. It is likely that Fennah (1971) was cor-
rect in his synonymy ofA. latifrons withA. servil-
lei (Freund & Wilson 1995); however, it would be
necessary to compare specimens with the type of
A. servillei to be absolutely certain. For the time
being, we treat the Florida species asA. servillei.
So far, A. excavata is known only from Miami-
Dade County.Acanalonia servillei,A. pumila, and
A. conica appear to be distributed widely in Flor-
ida. FSCA distribution records are disjunct, prob-







Florida Entomologist 90(4)


ably representing localities of collecting activities ern Florida, whereas A. pumila may be a Carib-
rather than actual distribution of the insects. In bean species that also occurs in peninsular Flor-
general, based on FSCA specimens, A. bivittata ida. However, Metcalf (1954) lists North Carolina
may be a northern species that ranges into north- as a location for A. pumila.

KEY TO THE ACANALONIA SPECIES OF FLORIDA

la. Vertex with a prominent median longitudinal carina ....................... Acanalonia servillei Spinola
lb. Vertex without a prominent median longitudinal carina .......................................... 2
2a. Dorsum with a pair of dark longitudinal stripes ............................ Acanalonia bivittata (Say)
2b. Dorsum without a pair of dark longitudinal stripes ............................................... 3
3a. Body less than 7 mm long; forewings hemispherical .................... Acanalonia pumila (Van Duzee)
3b. Body greater than 7 mm long; forewings trapezoidal .............................................. 4
4a. Head extended anteriorly beyond the lateral carina greater than the horizontal length of an eye, in lateral
view; frons with length/width ratio greater than 0.65; head declivent at most 10 ; frons sparsely pubescent
.................................................................. Acanalonia conica (Say)
4b. Head extended anteriorly beyond the lateral carina less than the horizontal length of an eye, in lateral
view; frons with length/width ratio less than 0.65; head declivent about 30 ; frons densely pubescent
........................................................ . Acanalonia excavata (Van Duzee)


ACKNOWLEDGMENTS

We thank Mr. David Ziesk, Division of Plant Indus-
try, Florida Department of Agriculture and Consumer
Services, Gainesville, for providing the automontage
photographs and Dr. Stuart McKamey, National Mu-
seum of Natural History, Washington, D.C., for the loan
of Ball's specimen from Florida. This is Entomology Con-
tribution No. 1068, Florida Department of Agriculture
and Consumer Services, Division of Plant Industry, Bu-
reau of Entomology, Nematology, and Plant Pathology.

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