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Title: Florida Entomologist
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Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1986
Copyright Date: 1917
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Subject: Florida Entomological Society
Entomology -- Periodicals
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Insects -- Florida -- Periodicals
Insects -- Periodicals
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FLORIDA ENTOMOLOGIST

(An International Journal for the Americas)

Volume 69, No. 4 December, 1986

TABLE OF CONTENTS

Announcement 70th annual meeting and call for papers ............................ i
MARTINEZ, A. E. AND, T. H. ATKINSON-Annotated Checklist of Bark and
Ambrosia Beetles (Coleoptera: Scolytidae and Platypodidae) Associated
with a Tropical Deciduous Forest at Chamela, Jalisco, Mexico ............. 619
KURCZEWSKI, F. E., AND M. G. SPOFFORD-Observations on the Behaviors of
some Scoliidae and Pompilidae (Hymenoptera) in Florida ................. 636
ALI, A., B. H. STANLEY, AND P. K. CHAUDHURI-Attraction of Some Adult
Midges (Diptera: Chironomidae) of Florida to Artificial Light in the Field 644
PROKOPY, R. J., D. R. PAPAJ, AND T. T. Y. WoNG-Fruit-Foraging Behavior
of Mediterranean Fruit Fly Females on Host and Non-Host Plants ...... 651
DEYRUP, M., AND D. MANLEY-Description of the Male of Pseudomethoca
oculata(Banks) (Hymenoptera: Mutillidae) ...................................... 658
SLATER, J. A.-Aulacoblissus, A New Genus of Micropterous Blissinae from
Venezuela (Hemiptera: Lygaeidae) ............................ ................... 661
PLAGENS, M. J., AND W. H. WHITCOMB-COrn Residue as an Overwintering
Site for Spiders and Predaceous Insects in Florida ............................ 665
LILLIE, T. H., AND D. L. KLINE-Occurrence of Culidoides mississippiensis on
Different Types of Vegetation ........................................................ 672
WALKER, T. J.-Monitoring the Flights of Field Crickets (Gryllus spp.) and a
Tachinid Fly (Euphasiopteryx ochracea) in North Florida ................ 678
ALI, A., AND J. K. NAYAR-Efficacy of Bacillus sphaericus Neide Against Larval
Mosquitoes (Diptera: Culicidae) and Midges (Diptera: Chironomidae) in
the L laboratory .......................................................................... 685
ABOU-SETTA, M. M., R. W. SORRELL, AND C. C. CHILDERS--Life 48: A Basic
Computer Program to Calculate Life Table Parameters bfr an Insect or
M ite Species .......................................................... .................... 690
MENKE, A. S., AND L. A. STANGE-Delta campaniforme rendalli (Bingham) and
Zeta argillaceum (Linnaeus) Established in Southern Florida, and Com-
ments on Generic Discretion in Eumenes s. 1. (Hymenoptera: Vespidae:
Eumeninae) ........ .. .................. ........................................... 697
LUCAS, J. R.-Antlion Pit Construction and Kleptoparasitic Prey ............... 702
GOOI)WIN, J. T.-Immature Stages of Some Western Nearctic and/or Neotropical
Tabanidae (Diptera) ........................................ .......................... 710
PROKOPY, R. J.-Alightment of Apple Maggot Flies on Fruit Mimics in Relation
to Contrast Against Background ........................................ ............ 716
BACKUS, V. L., AND W. H. CADE-Sperm Competition in the Field Cricket
Gryllus integer (Orthoptera: Gryllidae) ........................................ 722



Continued on Back Cover

Published by The Florida Entomological Society














FLORIDA ENTOMOLOGICAL SOCIETY



OFFICERS FOR 1986-87
President .... ................................ ................. D. J. Schuster
President-Elect .................................. ............... J. L. Taylor
Vice-President ...................................... ................ R. S. Patterson
Secretary ................................................... E. R. Mitchell
Treasurer ...................................... ............... A. C. Knapp

M. L. Wright, Jr.
J. E. Eger, Jr.
S*L. S. Osborne
Other Members of the Executive Committee ................. Mathrin
A. Gettman
C. G. Witherington
J. R. McLaughlin


PUBLICATIONS COMMITTEE


E editor ................. ....... ...... ........ .... ........ ....... ........... .. J. R M cLaughlin
Associate Editors ........................................................ A. Ali
C. S. Barfield
J. B. Heppner
M. D. Hubbard
O. Sosa, Jr.
H. V. Weems, Jr.
W. W. Wirth
Business Manager ...................................................... A. C. Knapp

FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30.00 per year in advance, $7.50 per
copy. Membership in the Florida Entomological Society, including subscription to Flor-
ida Entomologist, is $25 per year for regular membership and $10 per year for students.
Inquires regarding membership, subscriptions, and page charges should be addressed
to the Business Manager, P. O. Box 7326, Winter Haven, FL 33883-7326. Florida
Entomologist is entered as second class matter at the Post Office in DeLeon Springs
and Winter Haven, FL.
Authors should consult "Instructions to Authors" on the inside cover of all recent
issues while preparing manuscripts or notes. When submitting a paper or note to the
Editor, please send the original manuscript, original figures and tables, and 3 copies
of the entire paper. Include an abstract and title in Spanish, if possible. Upon receipt,
manuscripts and notes are acknowledged by the Editor and assigned to an appropriate
Associate Editor who will make every effort to recruit peer reviewers not employed by
the same agency or institution as the authors(s). Reviews from individuals working
out-of-state or in nearby countries (e.g. Canada, Mexico, and others) will be obtained
where possible. Page charges are assessed for printed articles.
Manuscripts and other editorial matter should be sent to the Editor, JOHN R.
MCLAUGHLIN, 4628 NW 40th Street, Gainesville, FL 32606.


This issue mailed December 22, 1986















































































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THE 70th ANNUAL MEETING OF THE
FLORIDA ENTOMOLOGICAL SOCIETY
FIRST ANNOUNCEMENT AND CALL FOR PAPERS

The Florida Entomological Society will hold its 70th Annual Meeting on 12-14 Au-
gust, 1987 at the Daytona Beach Hilton, 2637 So. Atlantic Avenue, Daytona Beach, FL
32018; telephone (904)-767-7350. Room rates are $65.00 either single or double (children
included). Pre-registration and registration information will be mailed to members and
will appear in the Newsletter and the March, 1987 Florida Entomologist.
Since many will present papers please copy the form and submit before 1 June, 1987
to:

R. S. Patterson, Chairman
Program Committee, FES
P.O. Box 14565
Gainesville, FL 32604
Phone: (904)-374-5903

Eight minutes will be allotted for presentation of oral papers, with 2 minutes for
discussion. In addition, there will be a separate session for members who may elect to
present a Project (or Poster) Exhibit. The three oral student papers and the three
student Project Exhibits judged to be the best on content and delivery will be awarded
monetary prizes during the meeting. Student participants in the judged sessions must
be Florida Entomological Society Members and must be registered for the meeting.










Martinez & Atkinson: Mexican Bark Beetles 619


ANNOTATED CHECKLIST OF BARK AND
AMBROSIA BEETLES (COLEOPTERA: SCOLYTIDAE AND
PLATYPODIDAE) ASSOCIATED WITH A TROPICAL
DECIDUOUS FOREST AT CHAMELA, JALISCO, MEXICO

ARMANDO EQUIHUA MARTINEZ
Centro de Entomologia y Acarologia
Colegio de Postgraduados
56230 Chapingo, Mexico, Mexico
and
THOMAS H. ATKINSON
Institute de Biologia
Universidad Nacional Autonoma de Mexico
Apdo. 21 San Patricio 48980,
Jalisco, Mexico

ABSTRACT

An annotated checklist is presented of 99 species and 32 genera of bark and ambrosia
beetles (Coleoptera: Scolytidae and Platypodidae) known from the tropical deciduous
forest in the Estaci6n de Biologia Chamela and surrounding areas on the coast of Jalisco,
Mexico. Important components of the fauna include the scolytid tribes Corthylini (26
spp., mostly Araptus, Pityophthorus), Micracini (18 spp., mostly Pseudothysanoes,
Hylocurus) and Cryphalini (16 spp., mostly Hypothenemus). The fauna was poorly
known: 29 species were previously undescribed, 36 are reported from Jalisco for the
first time, and 3 are reported from Mexico for the first time. The area supports a
characteristic fauna; 26 species are known only from the area and another 24 are known
only from the Pacific Slope of Mexico. Most genera show neotropical affinities.

RESUME

Se present un listado comentado de 99 species y 32 g6neros de escarabajos des-
cortezadores y de ambrosia (Coleoptera: Scolytidae y Platypodidae) conocidos de selva
baja caducifolia en la Estaci6n de Biologia Chamela y areas cercanas en la costa de
Jalisco, Mexico. Componentes importantes de la fauna incluyen los escolitidos de las
tribus Corthylini (26 spp., principalmente Araptus, Pityophthorus), Micracini (18 spp.,
principalmente Pseudothysanoes, Hylocurus) y Cryphalini (16 spp., principalmente
Hypothenemus). La fauna era pobremente conocida: 29 species no estaban previamente
descritas, 36 se reportan por primera vez de Jalisco, y 3 se reportan por primera vez
de Mexico. El area mantiene una fauna muy caracteristica; 26 species se conocen
solamente de la zona y otras 24 se conocen solamente del vertiente del pacifico de
Mexico. La mayoria de los generos demuestran afinidades neotropicales.



Bark and ambrosia beetles are important components of temperate forest ecosys-
tems, particularly those with a dominance of conifers. An extensive literature exists on
those species in the northern temperate zones which are able to kill living trees. Even
those species which do not invade healthy trees are among the first insects to invade
moribund or recently dead woody tissue (Wood 1982). Despite the enormous diversity
of neotropical species, information on biology and host associations is scanty, and their











Florida Entomologist 69(4)


importance as a group in tropical forest systems is largely unknown. Bright (1981) and
Wood (1982) recently monographed the Scolytidae of Mexico and Central America, but
the biology and ecology of most of these are still poorly known. Schedl (1972) listed the
world species of Platypodidae but included no biological data; virtually no information
exists on neotropical species. Recent studies in central Mexico have greatly increased
our knowledge of the species in that area (Atkinson and Equihua 1985a, b, c, 1986b,
Atkinson et al., 1986).
We report here the results of an extensive collecting program in tropical deciduous
forest and related communities on the Estacion de Biologia Chamela and surrounding
areas. We discuss the composition of the fauna and its biogeographic affinities and
present a checklist of the species, summarizing distributions, biologies, and host associ-
ations. Preliminary results were summarized earlier (Equihua et al. 1985). There have
been many additions and corrections, and species accounts were not included.

METHODS

The Estacion de Biologia Chamela is a biological reserve and field station of the
Institute de Biologia, Universidad Nacional Aut6noma de Mexico, on the coast of the
state of Jalisco (municipio La Huerta, aprox. 19 30' N, 1050 03' W), Mexico (Figure 1).
The station covers 1600 ha, mostly below 150 m above sea level, and is 2 km from the
coast at its closest. The mean annual temperature is 24.0 C, and the 10 year average
rainfall is 748 mm. Most rain (aprox. 80%) falls in the 4 month period between 2 July
and 4 November; no appreciable rainfall has been detected in the last 10 years between
mid-February and late May (S. H. Bullock, Estaci6n de Biologia Chamela, personal
communication). The vegetation of the station is mostly tropical deciduous forest, with
some subdeciduous forest along the courses of the larger drainages. Estuarine and
riparian communities are found nearby. A recent checklist includes more than 750
species of vascular plants from the station (Lott 1985).
From March, 1982, until February, 1983, a total of 8 collecting trips were made to
the station by one or both of us for periods of 3 days to 2 weeks. Since January, 1985,
Atkinson has resided at the station and has added many host records as well as several
additional species. We collected most beetles from naturally infested host material.
Supplementary collections were made with a blacklight trap and by cutting and leaving
branches or trunks of suspected host plants for possible subsequent infestations. The
type of host tissue consumed (phloem, wood, pith) was noted at the time of collection.
The mating system was determined by direct observation of number and sex of adults
found in gallery systems and gallery architecture when possible.
S. L. Wood (Brigham Young University, Provo, Utah) and D. E. Bright (Biosys-
tematics Research Institute, Agriculture Canada, Ottawa) confirmed identifications and
described new species (Wood 1983, 1984, Bright 1985a, b). Emily J. Lott (Herbario
Nacional, Universidad Nacional Aut6noma de Mexiico) and J. Arturo Solis-Magallanes
(Estaci6n de Biologa Chamela) identified host plants.

FAUNISTIC ANALYSIS

We collected 99 species of bark and ambrosia beetles from the Chamela area (94
Scolytidae and 5 Platypodidae). Most species-(69%) had not been reported from the
area: these were either undescribed (29), unreported from Jalisco (36), or unknown from
Mexico (3). These are summarized by subfamilies and tribes in Table 1. The largest
groups are the Corthylini with 8 genera and 26 species (largest genera Araptus, 9 spp.,
and Pityophthorus, 8 spp.), the Micracini with 18 species in 3 genera (largest genera


December, 1986










Martinez & Atkinson: Mexican Bark Beetles


Fig. 1. Location of the Estaci6n de Biologia Chamela, Chamela, Jalisco, Mexico.











Florida Entomologist 69(4)


TABLE 1. TAXONOMIC SUMMARY OF BARK AND AMBROSIA BEETLES KNOWN FROM
THE CHAMELA REGION, JALISCO, MEXICO.

Family/Subfamily Tribe No. Genera No. Species

Platypodidae
Platypodinae 1 5
Scolytidae
Hylesininae Bothrosternini 1 1
Phloeotribini 1 5
Phloeosinini 3 6
Hypoborini 1 1
Subtotal Hylesininae (6) (13)
Scolytinae Scolytini 3 5
Ctenophorini 3 9
Micracini 3 18
Cactopinini 1 2
Xyleborini 2 5
Cryphalini 5 16
Corthylini 8 26
Subtotal Scolytinae (25) (81)
Subtotal Scolytidae (31) (94)

Total 32 99


Pseudothysanoes, 10 spp., and Hylocurus, 6 spp.), and the Cryphalini with 16 species
in 5 genera (largest genus Hil.,, #II,.,. i .. 11 spp.). The Corthylini is the dominant tribe
in most neotropical areas, and species of the Cryphalini are abundant in most tropical
areas of the world. The relatively minor importance of the Xyleborini is unusual for a
lowland tropical area and is probably related to the dry climate of the area. The diversity
of the Micracini is unusually high compared to any area reported in the literature thus
far.
Biogeographic affinities of the genera are shown in Table 2. Most show neotropical
affinities and these also include the most species. The Mexican element referred to here
includes those genera whose distributions are centered in Mexico, southwestern U.S.,
and northern Central America. The monotypic genus Phloeoterus is known only from
the station and is closely related to Dendroterus (Mexican) and Araptus (neotropical).
The genus Pityophthorus is centered in the nearctic realm, but the species found in
Chamela belong to the scriptor, lautus, and juglandis groups of Bright (1981). These
are basicallt distributed in the eastern and southern U.S., Mexico, and Central America.
The genera Phloeotribus and Scolytus are widely distributed but are well represented
in the neotropics.
The biogeographic affinities of the species are shown in Table 3. Those species
known only from the Chamela area and those from the Pacific slope of Mexico are
approximately of equal importance; species from lowland Mexico and Central America,
and those which are widely distributed in the neotropics follow closely in terms of
number of species. Although at the genus level the Chamela fauna has wider-ranging
affinities, at the species level it shows a very local character. The fauna includes 7
genera which have circumtropical distributions (and include 25 species), but only 8 of
these species are widely distributed. Several species of Hypothenemus, Xyleborus,
Xylosandrus, Platypus, and Hypocryphalus found in the Chamela area are known or
suspected to have been introduced into the new world in historic times. There is little


December, 1986











Martinez & Atkinson: Mexican Bark Beetles


TABLE 2. BIOGEOGRAPHIC AFFINITIES OF GENERA OF BARK AND AMBROSIA
BEETLES FROM THE CHAMELA REGION, JALISCO, MEXICO.

No. No.
Distribution Genera Species Genera

Neotropical 18 51 Araptus, Chramesus, Cnemonyx,
Cnesinus, Corthylus, Dendroterus
Gymnochilus, Hylocurus, Micro-
corthylus, Pseudothysanoes, Pycnar-
thrum, Scolytodes, Scolytopsis,
Stegomerus, Styphlosoma,
Thysanoes, Tricolus
Circumtropical 7 25 Cryptocarenus, Hypocryphalus, Hy-
pothenemus, Platypus, Scolytogenes,
Xyleborus, Xylosandurs
Mexican 4 8 Cactopinus, Chaetophloeus, Dendro-
terus, Phloeoterus
Nearctic 1 8 Pityophthorus*
Other 2 7 Phloeotribus, Scolytus

Total 32 99

*Chamela species belong to the scriptor, lantrus, and juglans groups of Bright (1981), which are basically neotrop-
ical in distribution.


TABLE 3. BIOGEOGRAPHIC AFFINITIES OF SPECIES OF BARK AND AMBROSIA
BEETLES OF THE CHAMELA REGION, JALISCO, MEXICO.

Distribution No. Species

Chamela Region Only 26
Pacific slope of Mexico 24
Lowland Mexico and Central America 21
Widespread Neotropical 17
Widespread World Tropics 8
Other 3

Total 99


reason to believe that the species thus far known only from the vicinity of the station
are really narrow endemics, as most of the Pacific coast of Mexico from Nayarit to
Chiapas has been poorly collected. At the genus level there are broad similarities be-
tween the fauna of Camela and that of southern Morelos, Mexico, another area domi-
nated by tropical deciduous forest and forming part of the Pacific Slope (Atkinson et
al., 1986). We believe that the fauna of Chamela is representative of a distinct fauna of
bark and ambrosia beetles associated with tropical deciduous forest in Mexico. It differs
markedly from that associated with humid lowland forests in southeastern Mexico and
Central America both in generic and tribal level composition and in biogeographic af-
finities (Atkinson and Equihua 1986b). We suspect that this faunal element roughly
coincides with the distribution of tropical deciduous forest given by Rzedowski (1978).










624 Florida Entomologist 69(4) December, 1986


EXPLANATION OF CHECKLIST

This checklist is based entirely on the authors' collections (with the exception of a
few collections made by Felipe Noguera, Estaci6n de Biologia Chamela). There is no
additional material from the region in major Mexican collections nor are there any prior
references in the literature, other than descriptions of new taxa collected by us. The
following information is included for each species: valid name, summary of feeding
habits, degree of host specificity, and reproductive habits (in parentheses), summary
of known distribution, collections, and comments. Classification of feeding habits follows
Wood (1982) (ph=phloeophagy, xm=xylomycetophagy, m=myelophagy, x=xylophagy,
s= spermatophagy). Three degrees of host specificity are recognized: monophagy (=mo,
restricted to hosts of a single genus), oligophagy (=ol, restricted to hosts of a single
family), and polyphagy (=po, found in 2 or more unrelated plant families). Assignment
of degrees of host specificity was based on our observations, a critical review of the
literature, and our admittedly subjective judgment. Terminology for reproductive sys-
tems follows Kirkendall (1983) (mg=monogyny, bg=bigyny, pg=polygyny, ipg=inbred
polygyny). Distributions are summarized from Atkinson and Equihua (1985a,b,c,
1986b), Atkinson et al. (1986), Bright (1981, 1985,a,b), Schedl (1972), and Wood (1982).
An asterisk after the species name indicates a new state record for Jalisco; a double
asterisk indicates a new record for Mexico. Since all collections were made by the
authors in a restricted area with little altitudinal variation only host and collection
numbers are given for the collections. Specimens cited with collection numbers prefixed
by the letter "S" are deposited in the insect collection of the Centro de Entomologia y
Acarologia, Colegio de Postgradusdos, Chapingo, Mexico; collections prefixed by
"FANM" and "THA" are deposited in the Instituto de Biologia, Universidad Nacional
Aut6noma de Mexico, Mexico, D.F. Significant new host associations are indicated in
bold face. Comments on biology and ecology are included when these represent signif-
icant new information. Subfamilies and tribes follow the order given by Wood (1982);
genera and species are ordered alpabetically within tribes.

ANNOTATED CHECKLIST OF THE BARK AND AMBROSIA BEETLES
ASSOCIATED WITH TROPICAL DECIDUOUS FOREST AT
CHAMELA, JALISCO, MEXICO

PLATYPODIDAE

1. Platypus compositus Say, 1824* (xm,mg,po); SE U.S., Mexico, Antilles, Ven-
ezuela, Guyana, Brazil, Argentina. U.V. light (S-755, S-846). No known hosts at
Chamela but polyphagous in the Southeastern U.S. (Blackman 1922, Chamberlin 1939).
2. Platypus excisus Chapuis, 1865* (xm,mg,ol); Mexico, C. America, Cayenne,
Brazil. Lonchocarpous guatemalensis Benth. (NEW HOST) (S-818); Pithecellobium
dulce (Roxb.) Benth. (NEW HOST) (S-478); U.V. light (S-755, S-783, S-821, S-846);
host unknown (S-808). Widely collected in tropical communities in Mexico, always in
leguminous hosts (Atkinson & Equihua 1986b, Atkinson et al. 1986). It is most com-
monly collected in small diameter host material (< 10 cm).
3. Platypus parallels (Fabricius, 1801). (xm,mg,po); tropical America and Africa.
Tabebuia sp. (NEW HOST) (S-428); Guapira sp. (NEW HOST) (S-724); Croton
pseudoniveus Lundell (NEW HOST) (S-817); Ficus sp. (S-819); Spondias purpurea L.
(NEW HOST) (S-825); Brosimum alicastrum Sw. (NEW HOST) (S-883); Thouinidium
decandrum (Humb. & Bonpl.) Radlk. (NEW HOST) (THA-263), U.V. light (S-755,
S-783, S-846); Host unknown (S-857, 868). Most commonly collected species of Platypus











Martinez & Atkinson: Mexican Bark Beetles


on the station; found mostly in large diameter material (> 30 cm) in shaded, more humid
sites.
4. Platypus pulchellus Chapuis, 1865*. (xm, mg, po); Mexico, C. America, Guyana,
Surinam, Brazil. U.V. light (S-755, S-783, S-846). No known hosts at Chamela, assumed
to be polyphagous as are the majority of species in the genus.
5. Platypus segnis Chapuis, 1845*. (xm, mg, po); Mexico, C. America, S. America
to Bolivia. U.V. light (S-755); host unknown (S-351, S-846). No known hosts at Chamela;
polyphagous elsewhere (Atkinson & Equihua 1984b, Atkinson et al. 1986).

SCOLYTIDAE

HYLESININAE

Bothrosternini

6. Cnesinus setulosus Blandford, 1896*. (m,mg,po); Tamaulipas and Jalisco to
Panama. Conocarpus erecta L. (NEW HOST) (S-353); U.V. light (S-755); Acacia sp.
(S-362); Morisonia americana L. (NEW HOST) (THA-372); Bignoniaceae (S-474);
Clytostoma binatum (Thunb.) Sandw. (NEW HOST) (THA-262); host unknown (S-770,
S-860).


Phloeotribini

7. Phloeotribus geminus Wood, 1983. (ph,mg,mo?); known only from Chamela. host
unknown (S-731). Gallery transverse, biramous. Found in small branches, 1.5 cm diame-
ter.
8. Phloeotribus opimus Wood, 1969. (ph,mg,mo); Jalisco, Morelos, to Costa Rica.
Ficus insipida Willd. (THA-383).
9. Phloeotribus setulosus Eichhoff, 1868*. (ph,mg,mo); Veracruz and Jalisco to
Brazil and Peru. Brosimum alicastrum Sw. (S-883). Wood (1982) lists several unrelated
hosts for this species but B. alicastrum appears to be the only host in the Chamela
area. It was only collected once but the characteristic galleries were seen on several
occasions in fallen branches of this tree.
10. Phloeotribus texanus Schaeffer, 1908*. (ph,mg,mo); SE U.S., Nuevo Le6n. Cel-
tis iguaneus (Jacq.) Sarg. (S-807, S-862, S-864). Gallery transverse, biramous; in host
material ranging from 2-10 cm. diameter, frequently associated with C'l( I,... ... sub-
opacus. Teneral adults move about freely and feed on remnants of phloem under bark
before emerging, often nearly erasing the pattern of galleries.
11. Phloeotribus sp. (ph,mg,mo); Undescribed species (S. L. Wood, personal com-
munication), also known from southern Morelos (Atkinson et al., 1986). Phyllostylon
brasilense Capanema (THA-354, THA-369). This Large species of Phloeotribus makes
transverse, biramous galleries in large branches and trunks (> 5 cm diameter) of
weakened or dead host trees. Both the parental and larval galleries heavily score the
sapwood, even in material with thick phloem. Galleries are initiated by females.

Phloeosinini

12. Chramesus exul Wood, 1983. (ph,mg,mo); known only from Chamela. Croton
sp. (S-410, S-411, S-427); C. sp. (S-727); C. pseudoniveus Lundell (NEW HOST) (S-817,
THA-264, FANM-166); host unknown (S-730, S-740, S-758, S-773). Gallery transverse,
biramous.


625











626 Florida Entomologist 69(4) December, 1986

13. Chramesus securus Wood, 1983. (x,mg,mo); known only from Chamela. Lon-
chocarpus sp. (THA-250, FANM-190, FANM-200); Leguminosae (S-365, S-425, S-740,
S-781); host unknown (S-442, S-814). Gallery longitudinal, in phloem and sapwood; lar-
vae mine sapwood.
14. Chramesus subopacus Schaeffer, 1908*. (ph,mg,mo); Florida and Arizona to
Honduras. Sapindaceae (S-778); Celtis iguaneus (Jacq.) Sarg., (S-779, S-807, S-839,
S-862, S-873). The collection from a sapindaceous vine (S-778) is apparently an "acciden-
tal host" since this species is most frequently associated with Celtis spp. The gallery is
transverse and biramous.
15. Chramesus vitiosus Wood, 1969*. (ph,mg,mo); Nayarit and Jalisco. Lonchocar-
pus guatemalensis Benth. (NEW HOST) (S-409); L. constrictus Pitt. (NEW HOST)
(THA-361, THA-362); Leguminosae (S-425, S-728, S-740); host unknown (S-806, S-814,
S-821, S-874). Egg gallery longitudinal, frequently scoring sapwood.
16. Dendrosinus mexicanus Wood, 1983. (x,mg,mo); Jalisco, Morelos. Guapira sp.
(S-372, S-413, S-724). The egg gallery is transverse, biramous, up to 5 cm total length.
The gallery is excavated at a depth of 1.5-3 cm into the wood. Larvae develop entirely
in wood, each individual emerging through a separate hole on emergence. Successful
attacks (with emergence) were observed in host material from 10-40 cm diameter; at-
tacks in smaller diameter material were usually aborted, or larval mortality was virtu-
ally complete. In galleries with active larvae the surrounding wood is normally stained
blue or black, probably by associated fungi.
17. Pseudochramesus sp. ** (ph,mg,mo). Cynometra oaxacana Brandegee (THA-
360). This is the first record of any species of this genus in Central America or Mexico;
all other described species are South American. Although it was only collected once,
galleries have been observed ofi many occasions in its host tree. Galleries are initiated
by females and are transverse, biramous, and almost entirely in the phloem. Successful
attacks have been observed in host material ranging from 3-30 cm diameter.

Hypoborini

18. Chaetophloeus minimus Wood, 1967*. (ph,mg,ol); Jalisco and Colima. Apop-
lanesia paniculata Presl (NEW HOST) (THA-389); Leguminosae (S-360, S-870); host
unknown (S-821). Found in bark of 2 different host species. The gallery is typical of the
genus, consisting of a cave-like nuptial chamber, with 2-3 short egg galleries leading
from it. The egg galleries were longer with respect to the diameter of the nuptial
chamber (3-4 X diam.) than we have observed in other species of the genus (lasius,
sulcatus, mexicanus, confinis, struthanthi, penicillatus) in which the length of these
rarely exceeds the diameter of the chamber. The eggs are not deposited in niches, but
are left tightly packed in the frass left in the egg galleries. Orientation of egg galleries
is irregular, often curved.

SCOLYTINAE

Scolytini

19. Cnemonyx equihuai Wood, 1983, (ph,mg,mo); known only from Chamela area.
Hura polyandra Baill. (S-383, FANM-212, FANM-240). Gallery transverse, biramous,
in branches 2-3 cm diameter.
20. Cnemonyx splendens (Wood, 1961)*. (ph,mg,mo). Jalisco. Hippomane man-
cinella L. (NEW HOST) (THA-320). Transverse galleries in trunks and branches of
host tree. This species appears capable of invading living tissues of its host, as well as











Martinez & Atkinson: Mexican Bark Beetles 627


fallen or killed material. This is the first known locality for this species; it was described
from infested wood intercepted at the U.S.-Mexican border.
21. Scolytopsis puncticollis Blandford, 1896*. (ph,mg,ol); Veracruz and Jalisco to
Argentina, Cuba. Conocarpus erecta L. (NEW HOST) (S-353, THA-258); host unknown
(S-364). This species is also reported from Laguncularia racemosa (L.) Gaertn. and
Terminalia amazonia (Gmel.) Exell in Pulle, both of the Combretaceae (Atkinson and
Equihua 1985b, 1986a). L. racemosa is common in the Chamela area and grows in close
association with C. erecta, also of the Combretaceae.
22. Scolytus cristatus Wood, 1969. (ph,mg,mo); Jalisco to Venezuela. Heteropterys
laurifolia (L.) Adr. Juss. (NEW HOST) (S-469, S-829, S-858). Galleries transverse to
diagonal, uni- or biramous; in phloem of lianas of Heteropterys laurifolia (Malpighiaceae)
ranging in diameter from 3-10 cm. This is the first known host for this species.
23. Scolytus propinquus Blandford, 1896*. (ph,bg,ol); Chihuahua to Costa Rica.
Leguminosae (S-357, S-363); Lonchocarpus guatemalensis Benth. (NEW HOST) (S-
818); L. constrictus Pitt. (NEW HOST) (THA-351); L. eriocarinalis Micheli (NEW
HOST) (THA-370); host unknown (S-780, S-806). Found in host material ranging from
5-30 cm diameter; gallery transverse, biramous.

Ctenophorini

24. Gymnochilus reitteri Eichhoff, 1878*. (ph,mg,mo); Durango to Panama. Ficus
sp. (S-774). Found in broken or fallen branches. Gallery transverse, biramous.
25. Pycnarthrum amersum Wood, 1983 (ph,mg,mo); known only from Chamela
area. Brosimum alicastrum Sw. (S-883). Galleries transverse, biramous, up to 10 cm
total length. Found in host material from 15-50 cm diameter.
26. Pycnarthrumfurnerium Wood, 1971**. (ph,mg,mo); Jalisco to Honduras. Ficus
insipida Willd. (NEW HOST) (THA-351); F. sp. (S-369). First known host for this
species. The gallery is longitudinal and biramous; it was collected in branches ranging
from 1-10 cm diameter.
27. Pycnarthrum hispidum (Ferrari, 1867). (ph,mg,mo); Texas and Florida to
Panama. Ficus sp. (S-378, S-393, S-819); U.V. light (S-755, S-783).
28. Pycnarthrum reticulatum Schedl, 1940*. (ph,mg,mo); Jalisco and Veracruz to
Panama. Ficus sp. (S-393, S-774); U.V. light (S-755, S-821, S-846).
29. Scolytodes amoenus Wood, 1967. (ph,mg,mo); Tamaulipas, Jalisco. Ficus sp.
(S-774).
30. Scolytodes plumeriacolens Wood, 1983. (ph,mg,mo); known only from Chamela
area. Plumeria rubra L. (S-736, S-881). In phloem of small branches, 2-4 cm diameter.
The gallery is basically longitudinal. Like S. plumeriae, it can attack living host plants.
31. Scolytodes plumeriae Wood, 1969*. (ph,mg,mo); Jalisco and Puebla to Costa
Rica. Plumeria rubra L. (S-881). The galleries of this species vary considerably in
orientation but are mostly longitudinal, usually with one but occasionally two branches.
Eggs are placed in widely spaced niches; larval development is entirely in the phloem.
32. Scolytodes tennis (Wood, 1971). (ph,hpg,mo); Jalisco to Honduras. Ficus sp.
(S-404, S-774, S-834), F. insipida Willd. (THA-349). This harem polygynous species
breeds in twigs and small branches (< 2 cm diameter) of native figs. Galleries are
initiated by males, which are joined by up to 5 females, each of which excavates a
separate egg gallery. Attacks are mostly limited to shaded-out branches on living trees.

Micracini

33. Hylocurus dissimilis Wood, 1984. (x,bg,po); known only from Chamela. Sapin-
daceae (S-727); Sterculiaceae (S-769); Leguminosae (S-822); host unknown (S-426, S-800,











628 Florida Entomologist 69(4) December, 1986


S-877). The gallery is typical of the genus (Wood 1982). This species is capable of
completing several generations in the same piece of host material.
34. Hylocurus elegans Eichhoff, 1872*. (x,bg,po); Nayarit to Colombia. Acacia sp.
(S-362); A. hindsii Benth. (NEW HOST) (S-500, S-820); A. cochliacantha Humb. &
Bonpl. (NEW HOST) (S-475); Lonchocarpus sp. (NEW HOST) (S-373); Leguminosae
(S-425, S-746); Ficus sp. (NEW HOST) (S-404); Cordia dentata Poir. (NEW HOST)
(S-476); Sapindaceae (S-778); Hura polyandra Baill. (NEW HOST) (FANM-212); host
unknown (S-810, S-877). Gallery as described for genus by Wood (1982).
35. Hylocurus inaequalis Wood, 1956*. (x,by,po); Nayarit to Oaxaca. Ziziphus
amole (Sesse & Moc.) M. C. Johnst. (NEW HOST) (S-356); Cordia dentata Poir. (NEW
HOST) (S-476); Acacia sp. (S-362); A. cochliacantha Humb. & Bonpl. (NEW HOST)
(THA-245); Apoplanesia paniculata Presl. (NEW HOST) (THA-350; Lonchocarpus
constrictus Pitt. (NEW HOST) (THA-362); L. sp. (S-485); Leguminosae (S-363, S-406,
S-728, S-766, S-779); Hura polyandra Baill. (NEW HOST) (S-383); host unknown (S-
401, S-496, S-723).
36. Hylocurus micaeceus Wood, 1984. (x,bg,?); known only from Chamela. Host
unknown (S-764). Gallery as described for genus (Wood 1982).
37. Hylocurus scitulus Wood, 1984. (x,bg,mo); known only from Chamela. Acacia
sp. (S-748); A. hindsii Benth. (NEW HOST) (S-809, S-820); A. cochliacantha Humb. &
Bonpl. (NEW HOST) (THA-245); host unknown (S-810, S-875). Gallery as described for
genus by Wood (1982).
38. Hylocurus secus Wood, 1984. (x,bg,?); known only from Chamela. Leguminosae
(S-359). Gallery as described for genus by Wood (1982).
39. Pseudothysanoes acares Wood, 1969*. (ph,bg,mo?); Jalisco and Oaxaca.
Leguminosae (S-836); U.V. light (S-755); host unknown (S-481, S-496).
40. Pseudothysanoes mandibularis Wood, 1984. (ph,bg,mo?); known only from
Chamela. Host unknown (S-856, S-863). Galleries in phloem of unidentified liana 2-5 cm
in diameter; gallery longitudinal, biramous, egg galleries filled with packed frass, eggs
not in niches.
41. Pseudothysanoes mendicus Wood, 1969*. (ph,bg,mo?); Jalisco and Colima.
Leguminosae (S-841). Recent attacks in small branches (3 cm diam.) of a small legumin-
ous tree growing near beach.
42. Pseudothysanoes pumilus Wood, 1969*. (ph,bg,po); Jalisco and Colima. Mor-
isonia americana L. (NEW HOST) (S-476); Lonchocarpus sp. (NEW HOST) (S-355);
Caesalpinia eriostachys Benth. (NEW HOST) (S-402); Leguminosae (S-781). Gallery
longitudinal, biramous.
43. P...... I.,l...,,ii..,, simplex Wood, 1984. (ph,bg,mo?); known only from Chamela.
Host unknown (S-863).
44. Pseudothysanoes spinatulus Wood, 1984*. (ph,bg,mo); Jalisco, Oaxaca.
(Chlorophora tinctoria (L.) Gaud.) (NEW HOST) (THA-133). Galleries are initiated by
males which are later joined by 2 females. Initially each female makes a short longitud-
inal tunnel leading away from the main gallery entrance. At a distance of approximately
1 cm each then makes a transverse, biramous egg gallery, the branches of which are
slightly oblique with respect to the host branch. The overall gallery has an "H" shape,
turned sideways with respect to the axis of the host branch. Eggs are placed individually
in niches and the egg galleries are kept free of frass. Galleries have been observed in
material ranging from 1-5 cm diameter. Old galleries are commonly observed in dead
branches, still attached to the tree.
45. Pseudothysanoes squameus Wood, 1984. (ph,bg,ol); known only from Chamela.
Leguminosae (S-358, S-363, S-766, S-824); Acacia sp. (NEW HOST) (S-362); Lonchocar-
pus sp. (S-485); U.V. light (S-755); host unknown (S-476, S-723). The gallery is longitud-











Martinez & Atkinson: Mexican Bark Beetles


inal, biramous; each branch made by a different female. As the female advances she
packs boring frass in the gallery behind her; eggs are packed in the frass in the galleries,
not in niches.
45. Pseudothysanoes thomasi Wood, 1967*. (ph,bg,mo); Sinaloa and Jalisco. U.V.
light (S-755); Celaenodendron mexicanum Standl. (THA-201). The gallery is essentially
as described for P. spinatulus, however females commonly make one branch of their
egg galleries longer than the other, giving the overall gallery a much more irregular
shape. Eggs are packed individually in niches and galleries are kept free of frass.
Attacks have been observed in material ranging in diameter from 3-30 cm.
46. Pseudothysanoes truncatus Wood, 1984. (x,bg,mo); known only from Chamela.
Ziziphus amole (Sesse & Moc.) M. C. Johnst. (NEW HOST) (S-356, THA-388). The
gallery is similar to that described for P. spinatulus, but is entirely inside the sapwood
of small branches of its host. Wood (1984) in the species description lists Randia sp. as
host; this is due to an error in identification of the host plant.
47. Pseudothysanoes vesculus Wood, 1969*. (x,bg,mo); Jalisco and Colima. Ziziphus
amole (Sesse & Moc.) M. C. Johnst. (NEW HOST) (S-356, THA-388). The gallery is
similar to that of P. truncatus in all respects. Attacks are usually in slightly larger
branches.
48. Thysanoes epicaris Wood, 1969. (x,bg,po); Jalisco. Leguminosae (S-365, S-472);
Pithecellobium mangense (Jacq.) MacBride (NEW HOST) (S-418); Acacia sp. (FANM-
234); Sapindaceae (S-739, S-778).
50. Thysanoes texanus Blackman, 1943. (x,bg,po); Southern Texas to Jalisco.
Leguminosae (S-746); U.V. light (S-755).

Cactopinini

51. Cactopinus atkinsoni Wood, 1983. (ph,mg,mo); known only from Chamela.
Stenocereus cf. chrysocarpus Sanchez-Mejorada (NEW HOST) (S-497). Galleries exca-
vated in ribs of fallen or cut stems of its host, an arborescent cactus, which have dried
without rotting. Apparently several generations are passed in the same piece of host
material. Adult and larval mining was so extensive that the pattern of galleries could
not be fully appreciated. Two other species of arborescent cacti common in the area,
Pachycereus pecten-aboriginum and Cephalocereus purpusii, are not attacked.
52. Cactopinus setosus Wood, 1983. (ph,mg,ol); known only from Chamela area.
Acanthocereus occidentalis Britt. & Rose (NEW HOST) (S-498, S-435, S-801);
Stenocereus standleyi (Gonz. Ort.) Buxbaum (NEW HOST) (S-842); S. fricii Sanches-
Mejorada (NEW HOST) (THA-347). This species attacks wounds on living cacti, fre-
quently at the base of spines. The area mined by the insects seems to be relatively
limited, usually blackened and dry in appearance. It is unclear whether the insect
invades tissues killed by other agents or is capable of invading healthy tissue. It has
not been found in dry stems. All three cacti utilized as hosts are sprawling shrubs; S.
standleyi is particularly common on rocky shores.

Xyleborini

53. Xyleborus affinis Eichhoff, 1868. (xm,ipg,po). Circumtropical. Bursera in-
stabilis McVaugh & Rzed. (FANM-186); host unknown (FANM-176).
54. Xyleborus ferrugineus (Fabricius, 1801). (xm,ipg,po); eastern U.S. to Argen-
tina. U.V. light (S-755, S-846); Heliocarpus pallidus Rose (S-767); Ficus sp. (S-819);
Urera sp. (FANM-174); Sciadodendron excelsum Griseb. (FANM-171); Thouinidium
decandrum (Humb. & Bonpl.) Radlk. (THA-257); host unknown (S-783).


629











Florida Entomologist 69(4)


55. Xyleborus palatus Wood, 1974. (xm,ipg,po); Nayarit and Jalisco. Conocarpus
erecta L. (S-353); Cordia dentata Poir. (S-476); Lonchocarpus guatemalensis Benth.
(S-818); Pithecellobium sp. (S-361); Caesalpinia platyloba S. Wats. (FANM-220);
Leguminosae (S-394, S-870); Croton pseudoniveus Lundell (S-817; host unknown (S-
810). This species excavates large cave-like brood chambers between adjacent growth
rings in a manner similar to Dryocoetoides capucinus (Eichhoff) and Theoborus spp.
(Atkinson and Equihua 1985b).
56. Xyleborus volvulus (Fabricius, 1775). (xm,ipg,po); Florida and northern Mexico
to Argentina. Bursera arborea (Rose) Riley (S-354); B. instabilis McVaugh & Rzed.
(FANM-186); Leguminosae (S-421); Heliocarpus pallidus Rose (S-767); Sterculiaceae
(S-769); Carica papaya L. (S-827); Thouinidum decandrum (Humb. & Bonpl.) Radlk.
(THA-257); Sciadodendron excelsum Griseb. (FANM-171); Hura polyandra Baill.
(FANM-212); Ficus cotinifolia HBK. (FANM-202); U.V. light (S-755, S-783, S-821,
S-846); host unknown (S-770). Common pantropical "weedy" species.
57. Xylosandrus curtulus (Eichhoff, 1869)*. (xm,ipg,po); Nayarit and Veracruz to
Brazil. Ziziphus amole (Sesse & Moc.) M. C. Johnst. (S-356); Sapindaceae (S-798, S-
815); Serjania sp. (S-828); Cupania dentata DC. (S-872); Caesalpinia sp. (FANM-168).
Found in small diameter branches (1-3 cm), often in the pith.

Cryphalini

58. Cryptocarenus heveae (Hagedorn, 1912)*. (m,ipg,po); southern Florida and Col-
ima to Brazil. Annona sp. (S-370); Phoradendron sp. (S-747); Sapindaceae (S-739, S-778,
S-798); Thouinidium decandrum (Humb. & Bonpl.) Radlk. (S-734); Paullinia sessili-
flora Radlk. in Rose (S-743); Spondias purpurea L. (S-738); Caesalpinia sp. (S-823); C.
platyloba S. Wats. (FANM-236); Capparis flexuosa (L.) L. (THA-236); Leguminosae
(S-371, S-841, S-836); U.V. light (S-755, S-783); host unknown (S-838, S-846).
59. Cryptocarenus seriatus Eggers, 1933*. (m,ipg,po); southern Texas and Florida
to Brazil and Argentina. Leguminosae (S-371); Caesalpinia sp. (FANM-168); Sapin-
daceae (S-739, S-778); Sapotaceae (S-798); U.V. light (S-783); host unknown (S-770).
60. Hypocryphalus mangiferae (Stebbing, 1914)*. (ph,mg,mo); circumtropical, in-
troduced into New World. Mangifera indica L. (S-499). The cave-type parental galleries
deeply score the sapwood; larvae tunnel individually away from the parental gallery;
the oviposition pattern was not observed. This species breeds in broken, cut, or shaded
out branches, and appears to be able to kill live branches on occasion. It is potentially
a pest of its host tree.
61. Hypothenemus birmanus (Eichhoff, 1878). (m,ipg,po); Florida to Panama.
Moraceae (S-405, S-429); Ficus sp. (S-404); Trophis racemosa (L.) Urb. (S-484); Anacar-
diaceae (S-416); Leguminosae (S-472); Pithecellobium mangense (Jacq.) MacBride (S-
418); Rubiaceae (S-419); Bignoniaceae (S-474); Sapindaceae (S-701, S-722, S-739); Celtis
iguaneus (Jacq.) Sarg. (S-799, S-839); Laguncularia racemosa (L.) Gaertn. (S-803).
62. Hypothenemus brunneus (Hopkins, 1915). (m,ipg,po); Texas and Florida to
Panama. Acacia sp. (S-363); Leguminosae (S-371, S-421, S-425, S-733, S-740, S-746,
S-758, S-766, S-781, S-792, S-822, S-836, S-841); Caesalpinia eriostachys Benth. (S-402);
Lysiloma microphylla Benth. (S-768); Pithecellobium mangense (Jacq.) MacBride (S-
418); Schrankia sp. (FANM-193); Ziziphus amole (Sesse & Moc.) M. C. Johnst. (S-480);
Moraceae (S-405); Ficus sp. (S-378, S-834, S-774); F. cotinifolia HBK (THA-145);
Chlorophora tinctoria (L.) Gaud. (FANM-185); Celtis iguaneus (Jacq.) Sarg. (S-379,
S-839, S-862, S-807); Ceiba pentandra (L.) Gaertn. (S-403); Anacardiaceae (S-416);
Spondias purpurea L. (S-825); Comocladia engleriana Loes. (THA-155); Convol-
vulaceae (S-417); Serjania brachycarpa Gray (S-487); Sapindaceae (S-739, S-798, S-778);


630


December, 1986











Martinez & Atkinson: Mexican Bark Beetles


Croton pseudonivens Lundell (S-488); Hura polyandra Baill. (F-212); Phoradendron
sp. (S-747); Guazama ulmijfolia Lam. (S-749); Sterculiaceae (S-769); Forchhammeria
pallida Liebm. (S-364, S-737); Poeppigia procera Presl (S-790); Cocos nucifera L. (S-
793); Annona muricata L. (S-837); Cordia sp. (S-476); host unknown (S-351, S-390,
S-42, S-496. S-726, S-488, S-782, S-816, S-833, S-867, S-876, S-843, S-810). Wood (1982)
suggested that this species may be African in origin.
63. Hypothenemus callfornicus Hopkins. 1915*. (ph,ipg,po); California and Florida
to Michoacan and Veracruz. Amaranthaceae (S-495). This species is assumed to be
polyphagous since it is known from a wide variety of hosts (Wood 1982) and is poly-
phagous in southern Morelos (Atkinson et al., 1986). Wood (1982) suggested that this
species may be African in origin.
64. Hypothenemus columbi Hopkins, 1915*. (ph,ipg,po); Texas and Florida to Col-
ombia and Venezula. Alnona sp. (S-730); Ficus sp. (S-378). S-774); Cordia sp. (S-476);
Croton sp. (S-727); C. pseudoniveas Lundell (THA-151); Hura polyandra Baill.
(FANM-212); Matelea sp. (FANM-204); Sarcostemma clausum (Jacq.) Schult. (THA-
186); Ceiba pentandra (L.) Gaertn. (FANM-173); H..' .... Ij., ,r ,. Rose (THA-136);
Schrankia sp. (FANM-193); Spondias purpurea L. (S-738); Acacia sp. (S-748); Jatropha
standleyi -t ,,... i (S-760); U.V. light (S-755); host unknown (S-726, S-741, S-821,
S-846, S-805.
65. Hypothenemus crudiae (Panzer, 1791). (m,ipg,po); Eastern and southern U.S.
to Argentina. circumtropical Jatrophjba ''... i' Steyerm. (S-352. S-760); Jatropha sp.
(S-844); Euphorbia colletioides Benth. (THA-236); Forchhammeria pallida Liebm. (S-
364); Capparis indica (L.) Fawc. & Rendle (S-392); Morisonia americana L. (S-423);
Thouinia paucidentata Radlk. (S-367); Paullinia curtmr L. (S-376); Ficus sp. (S-378,
S-834); Celtis iguaneus (Jacq.) Sarg. (S-379); Anacardiaceae (S-416); Spondias p trpurea
L. (S-825); Comocladia engleriana Loes. (FANM-231); Jacquinia pungens A. Gray
(FANM-241); Ipomoea w/olcottiana Rose (FANM-242); Ceiba pentandra (L.) Gaertn.
(FANM-169); Heliocarpms pallidus Rose (THA-236); Matelea sp. (FANM-204); Sarcos-
tecnma clausum (Jac.) Schult. (THA-186); Rubiaceae (S-419); Manihot chlorosticta
Standl. & Goldman (S-482); Aristolochia taliscana Hook. & Arn. (S-745); Leguminosae
(S-757); Caesalpinia platyloba S. Wats. (FANM-210); Cocos nucifera L. (S-762);
Apocynaceae (S-763); Sterculiaceae (S-769); Carica papaya L. (S-827); Sideroxylon
capiri DC. (S-776); L ., acutangula Roxb. (S-845); host unknown (S-366, S-741, S-843,
S-846.
66. Hypothenemus erectus Leconte, 1876 (m,ipg,po). Southern Texas to Venezuela,
Africa. Acacia hindsii Benth. (S-820); Caesalpinia sp. (S-823); Acacia sp. (S-362);
Cynometra oaxacana Brandegee (S-391); Lonchocarpus sp. (S-373); Pithecellobium
mangense (Jacq.) MacBride (S-418); Leguminosae (S-371, S-472, S-750, S-836); Ziziphus
amole (Sesse & Moc.) M. C. Johnst. (S-480); Bignoniaceae (S-474); Capparisflexuosa
(L.) L. (THA-236); Forchhammeria pallida Liebm. (S-364); Thouinia paucidentata
Radlk. (S-367); Thouinidium. decandrmn (Humb. & Bonpl.) Radlk. (S-374, S-734); Paul-
linia cururu L. (S-376); P. . I r.'..,, Radlk. in Rose (S-743); Serjania brachycarpa
Gray (S-747); Cocos nucifera L. (S-793); Laguncularia racemosa (L.) Gaertn. (S-803);
Annon a mnricata L. (S-837); Annona sp. (S-370); Recchia mexicana Moc. & Sesse
(S-813); Avicennia germinans L. (S-477); Croton pseudoniveus Lundell (S-488, S-493);
C. sp. (S-727); Ficus sp. (S-378, S-404); F. cotinifolia HBK. (THA-151); Trophis
racemosa (L.) Urb. (S-484); Moraceae (S-405, S-429); Celtis iguaneus (Jacq.) Sarg.
(S-379, S-839); Spondias purpurea L. (S-825); Anacardiaceae (S-416); Rubiaceae (S-419);
Cordia sp. (S-424); host unknown (S-390, S-430, S-490, S-726, S-816, S-830, S-835,
S-860, S-810).











Florida Entomologist 69(4)


67. Hypothenemus eruditus Westwood, 1836. (ph-h,ipg,po); circumtropical. Jat-
ropha standleyi Steyerm. (S-759); Jatropha malacophylla Standl. (THA-368); Manihot
chlorosticta Standl. & Goldman (S-482); Lonchocarpus sp. (S-373); Caesalpinia erios-
tachys Benth. (S-402); Pithecellobium mangense (Jacq.) MacBride (S-418); Acacia
hindsii Benth. (S-500, S-809); Leguminosae (S-421, S-489, S-730. S-841); Lonchocarpus
constrictus Pitt. (THA-362); Ziziphus amole (Sesse & Moc.) M. C. Johnst. (S-480);
Capparis indica (L.) Fawc. & Rendle (S-392); Ficus sp. (S-393, S-774); F. cotinifolia
HBK. (THA-145); Trophis racemosa (L.) Urb. (S-484); Moraceae (S-429); Spondias
purpurea L. (S-415, S-825); Anacardiaceae (S-416); Convolvulaceae (S-417); Rubiaceae
(S-419); Cupania dentata DC. (S-872); Sapindaceae (S-701, S-815); Guazuma ulmifolia
Lam. (S-749); Cocos nucifera L. (S-762); Carica papaya L. (S-827); Celtis iguaneus
(Jacq.) Sarg. (S-762, S-807); Cordia sp. (S-476); Avicennia germinans L. (S-477); Luffa
acutangula Roxb. (S-845); Ceiba pentandra (L.) Gaertn. (S-403); Bignoniaceae (S-407,
S-474); Sarcostemma clausum (Jacq.) Schult. (THA-186); host unknown (S-366, S-470,
S-496, S-726, S-833, S-871, S-481, S-843, S-810). This species is found in a wide variety
of situations in virtually every plant examined. It has been collected in seeds, fruits,
pith, phloem, wood, and herbaceous plants, generally dead and dry.
68. Hypothenemus interstitialis (Hopkins, 1915)*. (m,ipg,po); Kansas and Florida
to Costa Rica. Leguminosae (S-750, S-792); Acacia sp. (S-362); Schrankia sp. (FANM-
193); Pithecellobium mangense (Jacq.) MacBride (THA-168); Caesalpinia platyloba S.
Wats. (FANM-210); Moraceae (S-405); Ficus sp. (S-834); Forchhammeria pallida
Liebm. (FANM-187); Capparis flexuosa (L.) L. (THA-236); Ceiba pentandra (L.)
Gaertn. (FANM-169); Sida sp. (THA-137); Spondias purpurea L. (S-825); Annona
muricata L. (S-837); Luffa acutangula Roxb. (S-845); Bignoniaceae (S-474); Sapin-
daceae (S-701); Paullinia sessiliflora Radlk. in Rose (S-743); Aristolochia taliscana
Hook. & Arn. (S-745).
69. Hypothenemus seriatus Eichhoff, 1878. (m,ipg,po); circumtropical. Forchham-
meria pallida Liebm. (S-364, S-737); Thouinia paucidentata Radlk. (S-367);
Thouinidium decandrum (Humb. & Bonpl.) Radlk. (S-374); Sapindaceae (S-701, S-739,
S-815); Paullinia sessiliflora Radlk. in Rose (S-743); Ficus sp. (S-369, S-774, S-834);
Trophis racemosa (L.) Urb. (S-484); Moraceae (S-429); Ceiba pentandra (L.) Gaertn.
(S-403); Spondias purpurea L. (S-415, S-791, S-825); Anacardiaceae (S-416); Convol-
vulaceae (S-417); Guazuma ulmifolia Lam. (S-420); Henrya insularis Nees in Benth.
(S-473; Bignoniaceae (S-474); Croton pseudoniveus Lundell (S-488, S-493); Jatropha
standleyi Steyerm. (S-760); Jatropha sp. (S-844); Leguminosae (S-740, S-750, S-758,
S-792, S-836); Caesalpinia sclerocarpa Standl. (THA-167); Lysiloma microphylla
Benth. (S-768); Matelea sp. (S-744); Cocos nucifera L. (S-762); Carica papaya L. (S-
827); Annona muricata L. (S-837); Avicennia germinans L. (S-477); Celtis iguaneus
(Jacq.) Sarg. (S-807); Recchia mexicana Moc. & Sesse (S-813); Luffa acutangula Roxb.
(S-845); host unknown (S-422, S-496, S-723, S-741, S-770. S-821, S-846).
70. Hypothenemus solocis Wood, 1974*. (m,ipg,po); Sinaloa, Nayarit, Jalisco, Col-
ima. Forchhammeria pallida Liebm. (S-364); Capparis flexuosa (L.) L. (THA-236);
Thouinia paucidentata Radlk. (S-367); Thouinidium decandrum (Humb. & Bonpl.)
Radlk. (S-734); Serjania brachycarpa Gray (S-828); Sapindaceae (S-739. S-778);
Leguminosae (S-371, S-733, S-750, S-781, S-836); Lysiloma microphylla Benth. (S-768);
Caesalpinia sp. (S-833); Ziziphus amole (Sesse & Moc.) M. C. Johnst. (S-480); Celtis
iguaneus (Jacq.) Sarg. (S-379); Mangifera indica L. (S-499); Astronium graveolens
Jacq. (FANM-184); Phoradendron sp. (S-747); Cocos nucifera L. (S-793); U.V. light
(S-783); host unknown (S-754. S-833, S-838). Like most species of the genus this is
principally a pith borer of twigs and small stems.


December, 1986











Martinez & Atkinson: Mexican Bark Beetles


71. Hypothenemus squamosus (Hopkins, 1915)*. (m, ipg, po); South Florida to Ver-
acruz and Jalisco. Leguminosae (S-733, S-781); Lysiloma ,iicrophiill Benth. (S-768);
Caesalpinia sp. (S-823); Thouinidium decandrum (Humb. & Bonpl.) Radlk. (S-734);
Serjania brachycarpa Gray (S-828); Sapindaceae (S-739, S-778); Sarcostema clausum
(Jacq.) Schult. (THA-327).
72. Scolytogenes rusticus (Wood, 1974). (ph,mg,mo); Jalisco to Chiapas. U.V. light
(S-755); Ipomoea wolcottiana Rose (NEW HOST) (S-765, THA-154). This species
breeds in branches and trunks of arborescent species of Ipomoea. The "wood" of these
trees consists of loosely bound "rings" of loose fibers. The galleries are basically trans-
verse and biramous and may be found between adjacent rings. It is not known whether
a single gallery may have branches on different levels, or whether each male-female
pair restricts its boring to a given level. It might be classified either as phloeophagous
or xylophagous.
73. Stegomerus pygmaeus Wood, 1967. (ph,mg,mo?); Nayarit to Costa Rica. Sapin-
daceae (S-375); host unknown (S-805, s-859). The galleries are transverse and biramous.
The species breeds in stems of an unknown woody vine, possibly a species of Paullinia.
Although Wood (1982) reported several unrelated hosts for this species, in Chamela it
seems to be monophagous.

Corthylini

74. Araptus sp. 1 (ph,pg,ol). Pithecellobium sp. (S-361); Caesalpinia sclerocarpa
Standl. (THA-371). Gallery radial, in phloem.
75. Araptus sp. 2 (ph,pg,mo?). host unknown (no number).
76. Araptus sp. 3 (m,mg,mo). Euphorbia colletioides Benth. (FANM-164, THA-
263). Excavates pith of small branches of its host, a woody liana. Unlike most species
of the genus, this is monogynous, possibly due to space considerations imposed by its
habits.
77. Araptus sp. 4 (ph,pg,mo?). U.V. light (S-755, S-846).
78. Araptus consobrinus Wood, 1975. (ph,pg,mo); Nayarit and Jalisco. Ficus sp.
(S-369, S-774); F. insipida Willd. (THA-249, THA-349); host unknown (S-407). Gallery
radial and in phloem.
79. Araptus delicatus Wood, 1974. (m,mg,mo). Nayarit, Jalisco. Sarcostemma
clausum (Jacq.) Schult. (NEW HOST) (THA-186, THA-327). Makes axial galleries in
the pith of small diameter stems of its host, a woody liana of the Asclepiadaceae. The
habits appear similar to those of Araptus sp. 3. Individual galleries up to 20 cm long
have been observed with large larvae present. The larvae apparently extend the paren-
tal gallery.
80. Araptusfossifrons Wood, 1975. (s,pg,ol.); Nayarit to Guatemala. Asclepiadaceae
(S-491); Matelea sp. (NEW HOST) (S-744); Marsdenia sp. (FANM-238); Gonolobus sp.
(THA-242). This species breeds in dry fruits of several genera of Asclepiadaceae (vines).
The galleries appear to be radial.
81. Araptus leptus (Bright, 1972)*. (ph,pg,mo?); Jalisco and Veracruz. Mal-
pighiaceae (S-486).
82. Araptus tabogae (Blackman, 1942)*. (ph,pg,m?); Jalisco and Veracruz to
Panama. Paullinia cururu L. (NEW HOST) (S-376). This is the first known host for
this species.
83. Corthylus spinifer Schwarz, 1891. (xm,mg,po). widely distributed in tropical
America. Astronium graveolens Jacq. (FANM-184). Restricted to shaded, moist ar-
royos.
84. Dendroterus fossifrons Wood, 1984. (ph,pg,mo?); known only from Chamela.
Host unknown (S-351). The condition of the host plant, a tree with exfoliating bark, did











634 Florida Entomologist 69(4) December, 1986

not permit identification but it was either a species of Bursera or Jatropha, both of
which host species of this genus.
85. Dendroterus luteolus (Schedl, 1951)*. (ph,pg,ol); Nayarit to Chiapas. U.V. light
(S-755, S-783, S-821); Spondias purpurea L. (NEW HOST) (S-879); Bursera instabilis
McVaugh & Rzed. (THA-346); host unknown (S-421). This species has mostly been
collected in Bursera spp. (Wood 1982, Atkinson et al. 1986). Successful attacks with
large larvae and pupae were observed.
86. Dendroterus mexicanus Blandford, 1904. (ph,pg,mo); Nayarit to Oaxaca. Burs-
era sp. (THA-382).
87. Dendroterus sallaei Blandford, 1904. (ph,pg,mo); Tres Marias Islands, Jalisco
and Tamaulipas to Costa Rica. U.V. light (S-755); Bursera arborea (Rose) Riley (S-354).
Gallery radial and in phloem.
88. Phloeoterus burserae Wood, 1984. (ph,pg,mo); known only from Chamela. Burs-
era instabilis McVaugh & Rzed. (S-794, S-865). Collected on both occasions from re-
cently killed branches (2-4 cm diam.) on live trees. Gallery radial and in phloem.
89. Microcorthylus minimus Schedl, 1950*. (xm,mg,po); Jalisco and Veracruz to
Brazil. Forchhammeria pallida Liebm. (NEW HOST) (S-772); Struthanthus interrup-
tus (HBK.) Blume (NEW HOST) (S-804); Cupania dentata DC. (NEW HOST) (S-872);
Ziziphus amole (Sesse & Moc.) M. C. Johnst. (NEW HOST) (S-356); host unknown
(S-770). The gallery is circular, extended on both side of the entrance, between adjacent
growth rings of its hosts. It is normally found in small stems and branches (1-3 cm
diam.).
90. Pityophthorus costabilis Wood, 1976. (ph,pg,mo); Jalisco, Morelos, Guerrero.
Thevetia ovata (Cav.) DC. (FANM-181). Gallery radial, in phloem.
91. Pityophthorus costatulus Wood, 1976. (ph,pg,mo); Jalisco, Morelos, Guerrero.
Thevetia ovata (Cav.) DC. (FANM-181). Gallery radial, in phloem. The egg galleries of
this species are packed with frass.
92. Pityophthorus diminutivus Bright, 1985. (ph,pg,mo?); known only from
Chamela. Lonchocarpus guatemalensis Benth. (S-761); host unknown (S-771, S-725).
Radial gallery, in phloem.
93. Pityophthorus equihuai Bright, 1985. (ph,pg,mo); known only from Chamela.
U.V. light (S-755); host unknown (S-832); composite shrub (THA-167). Gallery radial,
in phloem of woody vine.
94. Pityophthorus indefessus Bright, 1985. (ph,pg,mo); known only from Chamela.
Astronium graveolens Jacq. (S-416, THA-363, THA-364); host unknown (S-390, S-732).
Radial galleries, in phloem.
95. Pityophthorus indigens Wood, 1976. (ph,pg,mo?); Jalisco. Amphipterygium
adstringens (Schlecht.) Schiede (FANM-227). Galleries radial, in phloem.
96. Pityophthorus nanus Wood, 1964. (ph,pg,mo); Jalisco to Chiapas. Spondias pur-
purea (NEW HOST) (S-738, S-791); Burserafagaroides (HBK.) Engl. (THA-366). This
species has been previously reported only from Bursera spp. (Wood 1982, Atkinson et
al. 1986). Attacks occur in branches and trunks with diameters ranging from 3-30 cm.
97. Pityophthorus trunculus Bright, 1985. (ph,pg,mo?); known only from Chamela.
Host unknown (S-831). Gallery radial, in phloem of woody vine.
98. Styphlosoma granulatum Blandford 1904**. (ph,pg,?). Jalisco, Costa Rica,
Panama. Host unknown (S-351). This is the first report of this genus from Mexico.
99. Tricolus difodinus Bright, 1972*. (xm,mg,po); Nayarit to Guatemala. Sapin-
daceae (S-778). This species is known to be polyphagous in Campeche (Estrada and
Atkinson, unpublished) and is assumed to be so here as well. The gallery is similar to
that of Corthylus spinifer and Microcorthylus minimus.











Martinez & Atkinson: Mexican Bark Beetles 635


REFERENCES CITED

ATKINSON, T. H., AND A. EQUIHUA M. 1985a. Lista commtada de los cole6pteros
Scolytidae y Platypodidae del Valle de Mexico. Folia Entomol. Mex. 65: 63-108.
AND -- 1985b. Notes on biology and distribution of Mexican and Central
American Scolytidae. I. Hylesininae, Scolytinae except Cryphalini and Corth-
ylini. Coleopterists Bull. 39: 227-238.
--- AND 1985c. Notes on biology and distribution of Mexican and Central
American Scolytidae. II. Scolytinae, Cryphalini and Corthylini. coleopterists
Bull. 39: 355-363.
--- AND -- 1986a. Biology of the Scolytidae and Platypodidae (Coleoptera)
in a tropical deciduous forest at Chamela, Jalisco, Mexico. Fla. Entomol. 69:
303-310.
AND -- 1986b. Biology of bark and ambrosia beetles (Coleoptera:
Scolytidae and Platypodidae) of a tropical rain forest in southeastern Mexico with
an annotated checklist of species. Ann. Entomol. Soc. Amer. 79: 414-423.
- E. MARTINEZ F., E. SAUCEDO C., AND A. BURGOS S. 1986. Scolytidae y
Platypodidae (Coleoptera) asociados a selva baja caducifolia y comunidades de-
rivadas en el estado de Morelos, Mexico. Folia Entomol. Mex. 69 (in press).
BLACKMAN, M. W. 1922. Mississippi bark beetles. Miss. Agric. Exp. Sta. Tech. Bull.
11: 1-130.
BRIGHT, D. E. 1981. Taxonomic monograph of the genus Pityophthorus Eichhoff in
North America and Central America. Mem. Ent. Soc. Can. 118: 1-378.
S1985a. New species and records of North American Pityophthorus (Coleopt-
era: Scolytidae), Part IV: the scriptor group. Great Basin Nat. 45: 467-475.
1985b. New species and records of North American Pityophthorus (Coleopt-
era: Scolytidae), Part V: the juglandis group. Great Basin Nat. 45: 476-482.
EQUIHUA M., A., T. H. ATKINSON, AND E. J. LOTT. 1985. Scolytidae y Platypodidae
(Coleoptera) de la Estaci6n de Biologia Chamela, Jalisco. Agroci6ncia 57: 179-93.
KIRKENDALL, L. R. 1983. The evolution of mating systems in bark and ambrosia
beetles (Coleoptera: Scolytidae and Platypodidae). Zool. J. Linnean Soc. 77: 293-
352.
LOTT, E. J. 1985. Listado floristico de la Estaci6n de Biologia Chamela. Institute de
Biologia, Univ. Nacional Aut6noma de M6xico. 47 p.
RZEDOWSKI, J. 1978. Vegetacion de Mexico. Editorial Limusa, Mexico City. 432 p.
SCHEDL, K. E. 1972. Monographie der Platypodidae (Coleoptera). W. Junk, The
Hague. 322 p.
WOOD, S. L. 1982. The bark and ambrosia beetles (Coleoptera: Scolytidae) of North
and Central America, a taxonomic monograph. Great Basin Natur. Mem. 6:
1-1139.
1983. New synonymy and new species of American bark beetles (Coleoptera:
Scolytidae), Part IX. Great Basin Natur. 43: 647-659.
1984. New synonymy and new species of American bark beetles (Coleoptera:
Scolytidae), Part X. Great Basin Natur. 44: 113-119.

ACKNOWLEDGMENTS

We thank Emily J. Lott, Herbario Nacional, and J. Arturo Solis Magallanes, Esta-
cion de Biologia Chamela, Universidad Nacional Aut6noma de M6xico, who identified
host plants and served as guides in the field. Felipe A. Noguera Martinez, Estaci6n de
Biologia Chamela, allowed us to cite some of his collection data. Steven L. Wood,
Department of Zoology, Brigham Young University, and Donald E. Bright, Biosys-
tematics Research Institute, Agriculture Canada, helped identify material and de-
scribed new species. Emily J. Lott and Stephen H. Bullock critically reviewed this
manuscript. Field work was mostly supported by the Centro de Entomologia y
Acarologia, Colegio de Postgraduados.











Florida Entomologist 69(4)


OBSERVATIONS ON THE BEHAVIORS OF SOME SCOLIIDAE
AND POMPILIDAE (HYMENOPTERA) IN FLORIDA

FRANK E. KURCZEWSKI AND MARGERY G. SPOFFORD
Environmental and Forest Biology, State University of New York
College of Environmental Science and Forestry, Syracuse, NY 13210

ABSTRACT

The first report of burrowing behavior in a male nearctic scoliid, Campsomeris
plumipes fossulana, and the first description of the nesting behavior of the pompilid
Anoplius bequaerti are presented. Observations on the nesting behaviors and nests of
the spider wasps Episyron conterminus posters, Anoplius apiculatus pretiosus, and
A. stenotus are also given. New prey families for Auplopus mellipes mellipes and
Agenioideus birkmanni are included. An emendation is made to an earlier paper
(Kurczewski 1981) on nesting behavior of Florida Pompilidae.

RESUME

Se present el primer report del comportamiento del barrenador de un macho
scoliid neartico, Campsomeris plumipesfossulana, y la primer descripci6n del compor-
tamiento de anidar del pompilid Anoplius bequaerti. Tambien se reportan observaciones
sobre el comportamiento de anidar y de nidos de la avispa-arafa Episyron conterminus,
Anoplius apiculatus pretiosus, y de A. stenotus. Se incluyen nuevas families de presa
de Auplopus mellipes mellipes y de Agenioideus birkmanni. Se hace una enmienda de
una publicaci6n previa (Kurczewski 1981) sobre el comportamiento de anidar de Pom-
pilidae de la Florida.



Little information exists on female behavior in the nearctic Scoliidae, and nothing
is known about the behavior of the males (Krombein 1979). Much of the behavioral
information reported for species of nearctic Pompilidae is fragmentary and slight be-
cause of the solitary nature of the females and their often cryptic nesting situations (see
Evans and Yoshimoto 1962). Therefore, any information obtained on the behaviors of
species in these 2 families of aculeate Hymenoptera should be reported to add to the
knowledge about these incompletely studied groups.
With this goal in mind we present information on the burrowing behavior and resting
"cell" of a male of the scoliid Campsomeris plumipes fossulana (Fabricius) and on the
nesting behaviors and nests of the pompilids Priocnemis cornica (Say), Episyron con-
terminus posters (Fox), Anoplius apiculatus pretiosus (Banks), A. bequaerti (Dreis-
bach), and A. stenotus (Banks). The observations were made on the sandy firetrails of
the Archbold Biological Station (ABS), Lake Placid, Highlands County, and on the
sand-flats beside the Peace River at Arcadia (Ar), DeSoto County, Florida. The species
are treated in phylogenetic arrangement following Krombein (1979). The prey and wasp
specimens have been deposited in the invertebrate and insect collections, respectively,
of the S.U.N.Y. College of Environmental Science and Forestry. New prey records for
the pompilids Auplopus mellipes mellipes (Say) and Agenioideus birkmanni (Banks)
are included. These specimens are part of the insect collection of the Archbold Biological
Station.


December, 1986











Kurczewski & Spofford: Burrowing Wasps


Family SCOLIIDAE

Campsomeris plumipes fossulana (Fabricius)

ABS; 9 March 1986; 1430 (EST). One male flew from the vegetation bordering a
firetrail, made 3 low circular flights, each ca. 2.5 m in diameter, and landed on sand
near dried leaves and pine needles. He searched in a slight depression beneath this
debris for 40 sec, and then began to burrow into the sand using the mandibles and
forelegs. After 2 min his head and forelegs were buried from view. As the male dug
deeper, he twisted his body back and forth, forming an arc of ca. 1800. When his head
and thorax were below the sand surface, he began to push sand upward with the
extended abdomen, and, as the anterior half of the abdomen disappeared from sight,
his hindlegs and abdomen were both used to push the sand to the surface. After 9.5
min the wasp had disappeared from view, but every several sec his hindtarsi broke
through the accumulation pushing damp sand upward. Thereafter, the only evidence of
burrowing by the male was the occasional enlargement of the pile. No sand was pushed
upward after 20 min. He never backed out to the surface to clear away the sand accumu-
lation. This area was excavated 90 min later and a quiescent male in a position horizontal
to the sand surface was located at a depth of 4.5 cm.

Family POMPILIDAE

Priocnemis cornica (Say)

In 1981 one of us (FEK) described some aspects of nesting behavior and the nest of
Priocnemis (Priocnemis) sp. in Florida (ABS). At that time it was believed that the
observations pertained to an undescribed species of Priocnemis. However, M. S. Was-
bauer (pers. comm.) has informed us that the specimen in question is probably a P.
cornica with a red metasoma. He has examined other females similar to the Florida
specimen with red metasoma and varying amounts of red on the posterior femora and
tibiae from Turrialba, Costa Rica and Puerto Vallarta, Jalisco, Mexico and believes they
are nothing more than color variants of the usually all-black P. cornica. Conspecificity
is further suggested by the similarity in nesting behavior between the female observed
in Florida and several all-black females of this species (see Evans and Yoshimoto 1962).

Auplopus mellipes mellipes (Say)

Collection data: ABS; 31 March 1983; A. Schreffler. "Flying across the Plaza."
Prey: ?Aysha velox (Becker), immature (Anyphaenidae), with all legs amputated at
the coxal-trochanter joints.

Agenioideus birkmanni (Banks)

Collection data: ABS; 20 April 1983; M. Deyrup. "On bark of Pinus elliottii."
Prey: Platycryptus undatus (DeGeer), ? (Salticidae).

Episyron conterminus posters (Fox)

Ar; 30 March 1972; 1126 (EST); ABS; 11, 16 March 1986; 1315, 1450 (EST). A wasp
was observed digging a burrow with a 100 angle in a bare, white, sandy slope. She
stopped, turned 180 in front of the entrance, cleaned, and flew to her cachement site,











638 Florida Entomologist 69(4) December, 1986


a plant 2 m away. She apparently had lost or misplaced the spider because she searched
on the plant and the adjacent hillside for 16 min. The female then returned to the
burrow without the spider and began digging again. She dug for 5 additional min,
backed out, turned 180', cleaned, flew to the cachement site, and began searching on
the plants in the area. She finally abandoned the 5.2 cm-long burrow and began digging
2 m to the east. She also abandoned her second burrow, only 1 cm long, flew back to
the cachement site, searched on plants in the area, and then flew away. It is highly
unusual for a pompilid to begin a 2nd excavation after abandoning an initial nearly
completed one without first capturing, paralyzing, and caching another spider to replace
the one that was stolen or misplaced.
Another female had cached her paralyzed spider on the leaf of a plant 5 cm above
the sand surface and 60 cm from her nest entrance. She grasped the prey by a leg, flew
with it, and released it on the sand, 30 cm to the opposite side of the entrance. She then
pulled it backwards on the ground, grasping it with the mandibles by a leg, released it
5 cm from the entrance, walked to the opening, turned around, walked back to the prey,
pulled it backwards by a leg, and released it beside the entrance. She grasped the prey
by its spinnerets with the mandibles, and pulled it into the burrow.
The wasp appeared headfirst in her entrance 4 min after entering and began flinging
sand alternately into the burrow with the forelegs and hammering this sand into place
with the end of the abdomen. She came onto the sand surface to distances of 1-6 cm
from the entrance to obtain loose sand for the fill, even though the tumulus was only
2.5 cm long. After filling the burrow and entrance, the female spent 2 min placing small
twigs atop the fill. She eventually removed one of the several twigs placed on the filled
entrance but left the others in place. Her final closure took 12 min to completion. A
second wasp was collected as she closed her nest. She did not place debris atop the filled
entrance.
The tumuli in front of the 0.7 and 0.6 cm-wide entrances were 2.0 cm wide and 2.5
cm long and 4.5 cm wide and 6.0 cm long, respectively. The straight burrows, 3.8 and
6.4 cm long, terminated in cells 3.3 and 3.9 cm beneath the sand surface, respectively,
(Figs. la,b). The cells, 0.5 cm high and wide and 0.6 and 0.9 cm long, respectively,
contained a thoroughly paralyzed Araneus bonsallae (McCook), Y, placed on its left
side and a quiescent Eriophora ravilla (C. L. Koch), immature 9, on its right side (both
Araneidae), with the cephalothoraces placed toward the entrances. The eggs, 0.04 and
0.06 cm wide and 0.15 and 0.22 cm long, were placed obliquely on the right side of the
spider's abdomen, nearly halfway from the base (Fig. 2a) and almost longitudinally on
the left side of the spider's abdomen, nearly midway from its base (Fig. 2b). The wasps
weighed 13 and 24 mg and the spiders, 18 and 54 mg.

Anoplius apiculatus pretiosus (Banks)

Ar; 29 March 1972; 12 March 1986; 1130 (EST). A female was observed constructing
a burrow 16 cm from her prey, which lay dorsum upward on the sand. She pulled the
spider on the ground into the opening, grasping it by its spinnerets, but it became
wedged just inside. The wasp then spent 23 min digging around the spider, pulled it
farther downward into the opening, and covered it nearly completely with loose sand,
using the forelegs alternately. The resulting depression from which she took the sand
measured 3.0 x 4.5 cm. The wasp then walked across the sand and began another
burrow, 80 cm away. She periodically interrupted digging in her 2nd burrow, walked
to the 1st excavation, examined the exposed parts of the spider with her antennae,
walked back to the 2nd site, and resumed digging.
After digging in the 2nd excavation for 12 min, the wasp walked to the 1st site,








Kurczewski & Spofford: Burrowing Wasps


1


b


5 cm


L


Fig. 1. Burrows and cells of species of Pompilidae, as viewed from the side: a, b,
Episyron conterminus posters; c, Anoplius apiculatus pretiosus; d, A. bequaerti.


2


d


Fig. 2. Abdomens of prey spiders of species of Pompilidae, as seen in side view,
showing placements of wasps' eggs: a, b, Episyron conterminus posters; c, Anoplius
apiculatus pretiosus; d, A. bequaerti.











640 Florida Entomologist 69(4) December, 1986

grasped the spider by a leg with the mandibles, pulled the prey across the sand, released
it, dorsal side upward, and started a 3rd excavation, 1.1 m from the prey. She stopped
digging, walked to the spider, pulled it backwards, grasping it as before, and released
it 1 cm from the opening. She then dug alternately with the forelegs, backed out period-
ically while removing sand, and occasionally walked to and examined the spider with
her antennae. After 7 min of digging, the wasp grasped the spider by a hindleg with
the mandibles, placed its abdomen inside the hole, pulled it out of the hole, put it dorsum
up beside the entrance, and examined it for several sec. She then resumed digging and
eventually covered the spider almost completely with sand except for the ends of its
legs. After digging for 21 min, the wasp grasped the spider by a hindleg and, once
again, placed its abdomen inside the entrance. She then dug beneath it, but had consid-
erable difficulty attempting to pull the spider into the burrow by its spinnerets, using
the mandibles. The prey became wedged in the opening and the wasp spent another 5
min alternating between digging around the spider with the forelegs and pulling it down
the burrow by its spinnerets with the mandibles.
Two min after pulling the prey inside the female appeared headfirst in the entrance,
closing the burrow. She came out, flung sand backward rapidly, using the forelegs
alternately, and backed inside and hammered the sand into place with the end of the
abdomen. After 9 min of closing, the wasp flew away but was captured. This closure
was essentially identical to one observed 14 yr earlier at the same locality.
The entrances to the nests from 1972 and 1986, 0.9 and 1.0 cm wide, had not been
filled flush. The rather straight burrows, 8.5 and 9.5 cm long, led to cells, 7.0 and 7.5
cm beneath the surface (Fig. Ic). The cells, 0.6 and 0.7 cm high and wide, respectively,
and 0.8 and 1.0 cm long, contained rather lightly paralyzed Arctosa sp., immature and
penultimate 9 (Lycosidae) with the cephalothoraces positioned outward and dorsum
upward. The eggs, 0.04 and 0.06 cm wide and 0.15 and 0.18 cm long, were both laid
obliquely on the left sides of the spiders' abdomens near their bases (Fig. 2c). The wasps
weighed 12 and 21 mg and the spiders, 26 and 88 mg.

Anoplius bequaerti (Dreisbach)

ABS; 11 March 1986; 1030-1210 (EST). Three females were observed walking on the
sand, flicking their wings, and antennating the sand surface. Periodically, these wasps
paused and dug briefly in the sand with the mandibles and forelegs, presumably search-
ing for prey.
One wasp had captured a spider. After constructing her burrow, she dragged the
prey on the sand by a hindleg with the mandibles and pulled it into her nest by its
spinnerets, using the mandibles. The female appeared headfirst in her entrance, 23.5
min after entry with prey, and began shoving sand backward with the mid- and hindlegs
while packing this sand into place with the end of the abdomen. She then came onto the
surface and, bending the forelegs medially, raked sand backwards into the burrow,
using the forelegs alternately. As she backed in and hammered the sand into the burrow
with the end of the abdomen, her mid- and hindlegs continued to shove sand backward.
During this behavior, the antennae were curled at the ends, the forelegs were held
laterally, and the wings were held flat on the dorsum. After 27.5 min of closing, the
female flew off but was captured.
The entrance was 0.5 cm in diameter. The tumulus was evenly dispersed around the
entrance. The burrow entered the sand at a 400 angle to the horizon for 3 cm, plunged
vertically for 2.3 cm, and then curved back on itself for an additional 6 cm (Fig. Id).
Its total length was thus 11.3 cm and it terminated in a cell, 7.4 cm beneath the surface.
The cell, 0.7 cm high and wide and 0.95 cm long, contained a rather thoroughly paralyzed











Kurczewski & Spofford: Burrowing Wasps


Lycosa sp., immature (Lycosidae) placed cephalothorax outward and ventral side up-
ward. The wasp's egg, 0.07 cm wide and 0.25 cm long, was affixed obliquely to the right
side of the spider's abdomen near the base (Fig. 2d). The wasp weighed 46 mg and the
spider, 52 mg.

Anoplius stenotus (Banks)

Ar; 13 March 1986; 1159 (EST). A female was seen digging at the base of a steer
hoofprint, periodically backing out 3-4 cm from the opening, and removing sand alter-
nately with the forelegs. Her movements were noticeably slower than those of A.
apiculatus pretiosus (see above). A paralyzed Arctosa sp., immature, had been depo-
sited, dorsum upward, in the shade, 7 cm from the entrance. The wasp occasionally
rested and cleaned her antennae with the forelegs; then she would plug the entrance
from below with damp sand, back out, and remove this accumulation every 4 to 8 (mean,
6.0; N=5) min. After 31 min of burrow construction, the female appeared in the entrance
headfirst, as though finished with the tunnel, but then she continued to dig for an
additional 61 min, possibly because of sand cave-ins within the burrow. Her total burrow
construction time thus exceeded 1.5 h.
Periodic examinations of the spider were made by the wasp at intervals of 1-11
(mean 3.8; N=24) min. During each examination the female extended her antennae
which were curled at the ends toward the spider and held them rigid for 20-30 sec; then,
she turned and walked back into the opening. Six times the wasp repositioned the spider
either dorsum or venter up in the vicinity of (1-4 cm) or within the entrance, each time
grasping it by a hindleg with the mandibles, occasionally holding its body perpendicular
to hers. Before her next to last repositioning of the prey, the wasp enlarged the entrance
from 0.5 cm to 1.1 cm in diameter. She then flung the spider which lay partly in the
entrance out of the opening to a distance of 1 cm from the entrance, while digging with
forelegs, plugged the entrance from inside with damp sand, backed out removing the
accumulation, went inside headfirst, turned around within, and pulled in the prey by
its spinnerets, using the mandibles.


She appeared headfirst 3 min later, began filling the burrow by flinging sand back-
ward, using the forelegs alternately, and hammered this sand into place with the end
of the abdomen. During this behavior her antennae were spread laterally against the
walls and curled at the ends. The wings were held flat on the dorsum.
Her entrance was in the side of the steer hoofprint near the bottom, 5.5 cm beneath
the sand surface. The burrow was traced almost laterally for 3.5 cm but was lost in the
dry, loose, collapsing sand. The wasp weighed 16 mg, and the spider weighed ca. 20-25
mg.

DISCUSSION

Very little is known about the behavior of females of the nearctic Scoliidae (Krom-
bein 1979). Nothing is known about the behavior of the males. The tunneling of the male
of Campsomeris plumipes fossulana as inclement weather approached and its sub-
sequent passivity underground may represent typical Campsomeris behavior despite
the fact that it has never been described. The burrowing behavior of the male evidently
functions in enabling him to achieve an optimal depth for resting during the night and
periods of inclement weather. The female tunnels to locate, sting, and oviposit upon
prey. Her burrowing behavior is more efficient and more pronounced than that of the











642 Florida Entomologist 69(4) December, 1986


male, i.e., she disappears from sight more quickly and her movements are more exag-
gerated (see Kurczewski 1963a). To facilitate this, her body is more robust and her legs
are larger and more spinous than those of the male.
Auplopus mellipes mellipes constructs mud cells in the abandoned nests of certain
Vespidae and Sphecidae, in tunnels in wood, beneath loosened bark, under exposed
roots, and in abandoned bee burrows in the ground (Krombein 1979). The females prey
upon Pisauridae, Gnaphosidae, Thomisidae, and Salticidae (Krombein 1979). As in other
Auplopodini, the spider's legs are amputated prior to its placement in the cell (Evans
& Yoshimoto 1962). Our record of this species preying upon ?Aysha velox (Any-
phaenidae) introduces a new family, genus and species of prey.
The palearctic Agenioideus cinctellus (Spinola) apparently does not excavate its own
nest but, instead, appropriates niches in walls, rotten wood, and abandoned burrows of
other insects (summary in Richards & Hamm 1939). All 3 of the nearctic species of
Agenioideus are attracted to walls, cliffs, and buildings (Evans 1950). The lack of fosso-
rial spines on the forelegs of A. birkmanni hints at its use of pre-existing cavities for
nesting sites. This species has been collected twice previously with prey, in each case a
female of Herpyllus vasifer (Walckenaer) (Gnaphosidae) (Evans 1950, Kurczewski &
Kurczewski 1968). Our record of A. birkmanni preying upon the salticid Platycryptus
undatus introduces a new prey family, genus, and species. Richards & Hamm (1939)
list a variety of Salticidae and Thomisidae as prey of A. cinctellus. Evans (1950) re-
ported A. birkmanni transporting prey in flight but Kurczewski & Kurczewski (1968)
noted this species "pulling" its spider through grass. In the observation presented
herein, the wasp was pulling its prey across the bark of a tree (M. Deyrup, pers.
comm.). Obviously manner of prey transport is dependent upon the size of the spider,
as is presumably the case in the palearctic A. cinctellus (Richards & Hamm 1939).
Episyron conterminus posters nests in sand and preys upon orb-weaving spiders
(Krombein 1979). Rather extensive observations on the nesting behavior and nests of
this species have been published by Krombein (1952, 1955, 1958, 1959, 1964) and by
Kurczewski (1963b, 1981). The only new information we add to the known nesting
behavior of E. conterminus posters is the observation of 1 female placing twigs on its
nest after closing it. In this regard this wasp resembles one female of Poecilopompilus
algidus algidus (Smith) that, likewise, placed debris and pine needles atop its filled
entrance (see Kurczewski 1981). Araneus bonsallae (Araneidae) is a new species of prey
for Episyron conterminus posters.
Anoplius apiculatus (Smith) is another sand-nesting pompilid that has been studied
in some detail (Krombein 1979). Two subspecies, A. apiculatus autumnalis (Banks) and
A. apiculatus pretiosus (Banks), prey almost exclusively on the sand spider, Arctosa lit-
toralis (Lycosidae) (Evans & Yoshimoto 1962). After prey capture and during burrow
construction, the spider is frequently moved close to the entrance and covered with
sand (Krombein 1952, Evans, et al. 1953, present study). Our observations indicate that
A. apiculatus pretiosus, like A. apiculatus autumnalis (see Evans, et al. 1953), may
make several false starts, each time moving the spider to the vicinity of the excavation,
before remaining in 1 place and completing a nest.
Virtually nothing is known about the nesting behavior of Anoplius bequaerti. Krom-
bein (1964) described the prey transport of 1 female. Kurczewski (1981) collected 1 wasp
with an immature Schizocosa sp. (Lycosidae). The record we present herein (Lycosa
sp.) confirms the use of wolf spiders as prey. Our observations on the hunting, closure,
nest structure and dimensions, cell contents, and egg placement provide new informa-
tion for this species of pompilid.
A. stenotus, likewise, has been little studied (Krombein 1979). Krombein & Evans
(1955) described the nest and prey of 1 female. Kurczewski & Kurczewski (1973) and











Kurczewski & Spofford: Burrowing Wasps 643

Kurczewski (1981) noted the prey of 2 other wasps. In all cases, including the present
study, the prey comprised species of Lycosidae. The constant examinations of the spider
and the periodic repositioning of the prey are noteworthy behaviors in this species of
pompilid. The unusually lengthy period of burrow construction and the continual, appar-
ent sand cave-ins that we report are reminiscent of those of A. apiculatus autumnalis
(see Evans, et al. 1953) which also nests in loose sand.

ACKNOWLEDGMENTS

We thank M. Deyrup, Archbold Biological Station, for allowing us to examine 2
pompilids pinned with prey in the ABS collection, and G. B. Edwards, Div. of Plant
Industry, Florida Dept. of Agriculture and Consumer Services, for identifying the prey
spiders. We are grateful to M. S. Wasbauer, Div. of Plant Industry, California Dept.
of Food and Agriculture, for identifying Priocnemis cornica and for providing us with
information about its geographic variation.

REFERENCES CITED

EVANS, H. E. 1950. A taxonomic study of the nearctic spider wasps belonging to the
tribe Pompilini (Hymenoptera: Pompilidae). Part I. Trans. American Ent. Soc.
75: 133-270.
-- C. S. LIN, AND C. M. YOSHIMOTO. 1953. A biological study of Anoplius
apiculatus autumnalis (Banks) and its parasite, Evagetes mohave (Banks)
(Hymenoptera, Pompilidae). J. New York Ent. Soc. 61: 61-78.
-- AND C. M. YOSHIMOTO. 1962. The ecology and nesting behavior of the Pom-
pilidae (Hymenoptera) of the northeastern United States. Misc. Publ. Ent. Soc.
America 3: 65-119.
KROMBEIN, K. V. 1952. Biological and taxonomic observations on the wasps in a
coastal area of North Carolina (Hymenoptera: Aculeata). Wasmann J. Biol. 10:
257-341.
1955. Some notes on the wasps of Kill Devil Hills, North Carolina, 1954
(Hymenoptera, Aculeata). Proc. Ent. Soc. Washington 57: 145-160.
1958. Biological notes on some wasps from Kill Devil Hills, North Carolina,
and additions to the faunal list (Hymenoptera, Aculeata). Proc. Ent. Soc.
Washington 60: 97-110.
-- 1959. Biological notes on some ground-nesting wasps at Kill Devil Hills, North
Carolina, 1958, and additions to the faunal list (Hymenoptera, Aculeata). Proc.
Ent. Soc. Washington 61: 193-199.
1964. Results of the Archbold Expeditions. No. 87. Biological notes on some
Floridian wasps (Hymenoptera, Aculeata). American Mus. Nov. 2201: 1-27.
- Family Pompilidae, pp. 1523-1570. In Krombein, K. V., P. D. Hurd, Jr. D. R.
Smith, and B. D. Burks. 1979. Catalog of Hymenoptera in America north of
Mexico. Vol 2, Apocrita (Aculeata). Smithsonian Instit. Press, Washington, D.C.
AND H. E. EVANS. 1955. An annotated list of wasps collected in Florida,
March 20 to April 3, 1954 (Hymenoptera, Aculeata). Proc. Ent. Soc. Washington
57: 223-235.
KURCZEWSKI, F. E. 1963a. Biological notes on Campsomeris plumipes confluenta
(Say) (Hymenoptera: Scoliidae). Ent. News 74:21-24.
---. 1963b. Some new pompilid prey records from southern Florida (Hymenoptera:
Pompilidae). Florida Ent. 46: 209-213.
1981. Observations on the nesting behaviors of spider-wasps in southern Flor-
ida (Hymenoptera: Pompilidae). Florida Ent. 64: 424-437.
---, AND E. J. KURCZEWSKI. 1968. Host records for some North American Pom-
pilidae (Hymenoptera). First Supplement. J. Kansas Ent. Soc. 41: 367-382.











Florida Entomologist 69(4)


AND -- 1973. Host records for some North American Pompilidae
(Hymenoptera). Third supplement. Tribe Pompilini. J. Kansas Ent. Soc. 46: 65-
81.
RICHARDS, 0. W., AND A. H. HAMM. 1939. The biology of the British Pompilidae.
Trans. Soc. British Ent. 6: 51-114.





ATTRACTION OF SOME ADULT MIDGES
(DIPTERA: CHIRONOMIDAE)
OF FLORIDA TO ARTIFICIAL LIGHT IN THE FIELD

ARSHAD ALI
University of Florida, IFAS, Central Florida
Research and Education Center, P. O. Box 909
Sanford, FL 32771
BRUCE H. STANLEY
E. I. DuPont and Company, Agriclutural Products Department,
Experimental Station, Wilmington, DE 19898
and
PRASANTA K. CHAUDHURI
Department of Zoology, University of Burdwan,
West Bengal, India

ABSTRACT

The attraction of pestiferous species of adult Chironomidae to commercially available
lamps of various colors and wattages was studied by employing New Jersey light traps
along the shoreline of a central Florida lake. Glyptotendipes paripes Edwards, Goel-
dichironomus holoprasinus Goeldi, Chironomus crassicaudatus Malloch, and species
of Tanypodinae were predominant in the collections. A comparison of 100-W incandes-
cent lamps showed that white was preferred over yellow, and both were preferred over
red, orange, green or blue. Analysis of deviance indicated that these differences were
due primarily to differences in intensity (lux), although smaller effects of color and
differences in response among species were detected. No differences were observed in
preference between 60-W white incandescent or fluorescent lamps. These results were
consistent with previous laboratory studies, and indicate that manipulating light inten-
sity, rather than color, may be more appropriate in the overall development of an
integrated control strategy of nuisance chironomid midges.

RESUME

Se studio la atracci6n de plagas de species de Chironomidae por lamparas de various
colors y vatios de tipo New Jersey accesibles comercialmente, a lo largo de la costa de
un lago en la parte central de la Florida. Glypototendipes paripes Edwards, Goel-
dichironomus holoprasinus Goeldi, Chironomus crassicaudatus Malloch, y species de
Tanypodinae predominaron en las colecciones. Una comparaci6n de lamparas incandes-
centes de 100 vatios indic6 que el blanco era preferido sobre el amarillo, y que ambos
eran preferidos sobre el rojo, verde, o azul. Analisis de deviaci6n indic6 que esas diferen-
cias se debian principalmente a diferencias en intensidad (lux), aunque se detect
pequefio efectos por el color y direrencias en reacci6n entire las species. No se observa-


644


December, 1986











Ali et al.: Midge Light Attraction


ron diferencias en preferencia entire lamparas blancas de 60 vatios incandescentes o
fluorescentes. Estos resultados fueron consistentes con studios previous de laboratorio,
e indican que manipulando la intensidad de luz mas que el color, pudiera ser mas aprop-
riado enel desarrollo general de una estrategia de control integrado de plagas de
chironomids.



Many large lakes in the central region of peninsular Florida support heavy popu-
lations of chironomid midge larvae. A number of these lakes are surrounded by homes
and business establishments which are severely affected by the dense swarms of adult
midges which emerge from adjacent lakes. The nuisance and economic problems in
urban areas resulting from the massive accumulations of living and dead adult midges
were reported by Ali (1980); the economic loss (including tourist trade) to just one city
in central Florida may amount to several million dollars (Anonymous 1977).
Control of midges in small bodies of water (up to 200 ha) by the use of larvicides and
insect growth regulators is practiced in some parts of the United States (Ali and Mulla
1977a,b, Polls et al. 1975). The use of these materials, however, is not feasible or
economical in the relatively large size midge problem lakes that exist in Florida, each
covering thousands of hectares. Moreover, many of these habitats are either part of or
feed into river systems. Chemicals added to such systems are subject to displacement
and dilution and may be unacceptable from an environmental standpoint. In such situa-
tions, development of biological and cultural control strategies against the pest insects
are needed.
The manipulation of the attraction behavior of adult midges to artificial light may
facilitate midge reductions in the affected areas and thus prove useful in the overall
development of an integrated control program for these pestiferous insects. Recently,
Ali et al. (1984) investigated the preferences of some Florida pest species of midges to
incandescent light of different colors and intensities (wattage) under laboratory condi-
tions. The present studies were conducted beside a lake in central Florida to elucidate
the attraction of adult midges to various artificial light sources in the field situation.

MATERIALS AND METHODS

These studies were conducted during the summer of 1983 and 1984 near Lake Jessup,
Seminole County, Florida. Six New Jersey light traps equipped with suction mechanism
(Mulhern 1942) were used. Each trap was hung from a wooden pole 3 m above ground
level. Six poles were permanently placed at 20 m intervals along the northwest shore
of the lake parallel to the waterline. The land area behind the traps was lined with thick
and tall vegetation and lacked human dwellings or businesses. Thus, no visible sources
of artificial light were noticed in this area during the study periods at night. A portable
3000-W generator (Dayton, Inc., Dayton, OH) and suitable electrical cords were used
to illuminate bulbs (lamps) fitted in the traps. In these studies, commonly available
incandescent lamps also known as general purpose A-line lamps (Anonymous 1978) were
used to conduct three separate tests. In test 1, 100-W lamps of white (WL), yellow
(YL), orange (OL), blue (BL), green (GL), and red (RL) were used. Test 2 contained
WL, YL, OL, BL, GL, and RL of 25, 40, 40, 100, 100, and 100-W, respectively. In test
3, three 60-W lamps of WL and three 60-W warm white fluorescent (FL) lamps
(Circlite'TM) 60) were used. The WL lamps used in these tests were frosted (inside) while
the inner lining of all colored lamps was enameled.
The qualitative characteristics (wavelengths) of the lamps used are available


645










Florida Entomologist 69(4)


(Anonymous 1978). All of the color lamps emitted broad bands of wavelengths in the
visible spectrum. The wavelengths emitted by BL are between 430 and 490 nm, GL
between 490 and 550 nm, YL between 550 and 590 nm, OL between 590 and 620 nm,
and RL between 620 and 770 nm (Hollingsworth 1961). The radiant energy emitted by
WL and FL was in all visible wavelengths of 390 to 770 nm; the FL light peaks at ca.
180-190 nm, which are shorter than the peak wavelenghts of the WL (Ananymous 1978).
To estimate relative brightness (quantity of light) of the lamps, light intensity of
each lamp was measured with a LI-COR light meter (Li-Cor, Inc., Lincoln, NB) equip-
ped with a photometric sensor (LI-210SB). The intensity was measured in a photo-
graphy dark room. Each lamp was fitted in a NJ trap placed at a fixed location 2 m
above ground level and illuminated. The intensity reading for each lamp was taken at
a 2 m distance from the lamp with the sensor directly facing the light source. Light was
measured at the same location and height for each lamp. The measurements of light
intensity for 100-W WL, YL, OL, BL, GL, and RL were 37.1, 19.7, 11.1, 1.2, 1.4, and
2.1 lux, respectively, while those of 25-W WL, 60-W WL, 40-W YL, and 40-W OL were
4.9, 17.1, 5.8, and 3.1 lux, respectively. The intensity reading for the FL was 15.9 lux.
To attract adult midges emerging from Lake Jessup, the traps in each test were
activated about half an hour before sunset and adult trapping in the field was continued
for a period of 2 hours. Tests 1 and 2 were repeated on six different occasions so that
each lamp in a test occupied a different pole each time. Test 3 was also conducted on
six different occasions and the FL and WL lamps were positioned alternately on each
occasion. The adults trapped during each sampling period were identified and counted
in the laboratory.
The quantitative (number) differences of adult midges taken in different traps in
tests 1 and 2 were analyzed by analysis of variance (ANOVA) and in test 3 by Student's
t-test. The significance of the qualitative factors, species (S), lamp type (L) and color
(C), and the quantitative factor, intensity (I) in attracting the adult midges was eluci-
dated by contingency table analysis using log-linear models with quantitative factors as
described in Ali et al. (1984). Since intensity was measured for a lamp type and not each
lamp, the data for tests 1 and 2 were combined in the analysis to allow the effects of
color to be separated from those of intensity. In this analysis, intensity (I) was fitted
first in the hierarchy of increasingly more complex log-linear models, because it was an
attribute common to all lamps. Next, terms specifying the influence of lamp color, in
addition to its intensity (i.e., C + C x I) were added. Finally, terms defining differences
among the species in their response to intensity and color (i.e., S x I + S x C) were
entered. The three-way S x C x I interaction could not be included because of aliasing
(McCullagh and Nelder 1983) in the experimental design. The analysis for test 3 was
designed to separate the effects of lamp type (L) from differences in species preference
for a lamp type (i.e., S x L). The effects of intensity in test 3 could not be separated
from other aspects of the lamp type. In both tests, the mean residual deviance was
significantly greater (chi-square test) than one implying significant heterogeneity.
Therefore, the F-test using the mean deviances was employed to determine the signifi-
cance of the added terms.

RESULTS AND DISCUSSION

Glyptotendipes paripes Edwards was the predominant species constituting 76-93%
of the total adult midges taken in the three tests. Goeldichironomus holoprasinus
Goeldi, Chironomus crassicaudatus Malloch, and species of Tanypodinae [primarily
Coelotanypus concinnus (Coquillett), Coelotanypus scapularis (Loew), and Procladius
sublettei (Roback)] constituted 1-20%, <1-3%, and <1-2%, respectively, of the total


646


December, 1986











Ali et al.: Midge Light Attraction


adults trapped during these studies. Adults of other midge species, Goeldichironomus
cars (Townes), Chironomus decorus Johannsen, Cryptochironomus fulvus
Johannsen, Parachironomus Sp., Polypedilum halterale (Coquillett), Tribelos sp.,
Rheotanytarsus spp., Tanytarsus spp., and Cricotopus spp. collectively constituted
2-3% of the total collections in the three tests.
The number of adult midges attracted to each lamp (color or type) in the combina-
tions of lamps used in tests 1, 2, and 3, is presented in Fig. 1 as a percent of total adults
of each species (or group) taken in each test. The value of light intensity of each lamp
is also given in the figure. In test 1, WL with the highest intensity (37.1 lux) invariably
attracted significantly higher (1% level) numbers of adults of each species (or group)
than the YL, OL, BL, GL, or RL. In this test, YL (19.7 lux) was the second best
preference of the most predominant midge, G. paripes, while the differences between
the numbers of all midge species (or groups) attracted to the other four lamps, OL (11.1
lux), BL (1.2 lux), GL (1.4 lux), and RL (2.1 lux) were statistically insignificant.
In test 2, there were no significant differences (1% level) between the total number
of adults of each species (or group)attracted to WL (4.9 lux) and YL (5.8 lux). These
two lamps attracted >75% of the total G. paripes and C. crassicaudatus occurring in
all the traps used in the test. After WL and YL, G. paripes was attracted in significantly
higher numbers to the OL (3.1 lux) as compared with the BL (1.2 lux), GL (1.4 lux), or
RL (2.1 lux). The differences between the G. paripes catch totals by the BL, GL, and
RL were statistically insignificant. Chironomus crassicaudatus was apparently at-
tracted more to GL than to OL, BL, or RL, but the differences were not significant at
the 1% level of probability. In contrast to test 1, the attraction behavior of G. holop-
rasinus and species of Tanypodinae differed from that of G. paripes and C. cras-
sicaudatus; G. holoprasinus and Tanypodinae were attracted the most by BL than the
other five lamps in the combination. The statistical comparisons of adult catches of G.
paripes, C. crassicaudatus, G. holoprasinus, species of Tanypodinae, and total midges
by the incandescent WL and the white FL revealed no significant differences except for
the category, "other species" which showed a significant difference at the 5% level of
probability.
The results of the analysis of deviance for the three tests are presented in Table 1.
As shown in previous laboratory study (Ali et al. 1984), the intensity of light was the
most important factor which influenced the midge adult catches in tests 1 and 2 as
indicated by the value of 88.6% total deviance (% TD) attributed to the light intensity.
However, there were significant (95% CL) but relatively small effects of the color (5.7%
TD) and also differences in species preference (1% TD) as shown by the different pat-
terns of color preference of G. paripes from that of the Tanypodinae (Fig. 1). Consider-
ing the magnitude of the % TD associated with the factors studied, it becomes obvious
that, in general, lamp color has little impact upon the adult attraction of the species of
midges encountered in this study and that the light intensity is the primary factor
responsible for their attraction. In test 3, there were no significant differences between
the WL and FL catches (0.1% TD) and the differences in the species response to the
two lamp types were also insignificant (1.1% TD).
Numerous studies on the attraction of adult insects to light have been reported in
the literature with most studies conducted on terrestrial insects of economic importance
as reviewed by Heinton (1974). A variety of night-flying insects have shown specific
attraction responses to a certain range of wavelengths, particularly near the ultraviolet
region (300-390 nm) as shown by the cabbage looper [Trichoplusia ni (Hubner)], tobacco
hornworm [Manduca sexta (Johannsen)], pink bollworm [Pectinophora gossypiella
(Saunders)], corn earworm [Heliothis zea Boddie)], and others (Heinton 1974; Menear
1961; Glick and Hollingsworth 1954; Taylor and Deay 1950). The adult boll weevil (An-



















TEST 1


1001 [40

80- 30
60- I2
2020
40
20 10





100. 40
80-
30
60 20
40-
20 -10


100 40
80
S30
60,
40- 20
20 10

WY 0 B G R


TEST 2

Glyptotendipes paripes
100- 40
80- -30
60-
-20
40-
20- 10


Chironomus crassicaudatus
100- 40

80- 30
60-
-20
40-









20- 10
Goeldichironomus holoprasinu
100- 40
80










30
60





20
40-






20, 10
Tanypodinae
100- 40
80-
-30
60-
*20




40-
20 10







Total
Other species
100- 40
80-
30
60
40- *20


20" 10

Total
100- 40
80o
-30
60.
40. *20
20 n fl =a^10

W Y 0 B G R


December, 1986



TEST 3


100- 40
80- .30
60-
1 20
40-
20. *10


100- -40
80- 30
[30
60-
-20

20- 10




100- -40

80 30
60-
20
40-
20- -10



100- .40
80-
130
60-jJI
-20
40- -- /
20 -10




100- 40
80- 30
60-
20
40-










20- 10


W F


COLORS AND/OR TYPES OF LIGHT
Fig. 1. Attraction of adult chironomid midges in the field to six different color 100-W
incandescent lamps (test 1), to six different color incandescent lamps of different wat-
tage (test 2), and to three 60-W white incandescent and three 60-W white fluorescent
lamps (test 3). The bars represent the percentage of adults attracted to each lamp while
the line graphs indicate the light intensity of each lamp in a test combination. The W,
Y, O, B, G, and R indicate white, yellow, orange, blue, green, and red incandescent
lamps, respectively, while F indicates white fluorescent lamp. In test 2, the lamps were
25, 40, 40, 100, 100, and 100-W, respectively.


648


Florida Entomologist 69(4)












Ali et al.: Midge Light Attraction


TABLE 1. ANALYSIS OF DEVIANCE FOR FIELD STUDIES ON ADULT CHIRONOMID
MIDGEU ATTRACTION TO ARTIFICIAL LIGHT.

Lamp color or type Source of Change in % of total
combination variation df deviance' deviance'

Tests 1 and 2 (70,809 adults caught)
WL, YL, OL, BL, GL, RL I 1 89,946** 88.6
C +CxI 8 5,816** 5.7
SxI+SxC 24 998** 1.0
Residual 262 4,706 4.7
Test 3 (44,734 adults caught)
WL, FL L 1 24 0.1
SxL 4 339 1.1
Residual 145 30,878 98.8

'Mostly '* poriies Chir' i iinhito s crassicaiidtafits, Goeltichironouiii s holoprasiir s, and Tanypodinae.
'Deviance -2 Log,. (L), where L is the likelihood function (multinomial) evaluated at its maximum. The change
in deviance is approximately asymptotically chi-square distributed under the null hypothesis that the factors) does
not affect the catch frequencies. **,significant at the 1V, level based on the F-test.
'Total deviance after replicated and species catch totals were included in the model.


thonomus grandis Boheman) showed maximum attraction to wavelengths in the blue-
green region (470-515 nm) of the visible spectrum (Hollingsworth et al. 1964). Among
adults of aquatic insects, species of Simuliidae (Williams and Davis 1951) and several
other species of Diptera were reported to be attracted to light (Frost 1957). Belton and
Pucat (1967) reported that the adult biting midge [Culicoides sanguisuga (Coquillett)]
showed strong attraction to ultraviolet light. In contrast, the adult mosquito Aedes
aegypti (L.) showed a preference for the near infrared region (Magnum and Callahan
1968). Thus, adults of different insect species or groups can have a preference for a
different band of radiant energy in the electromagnetic specturm.
The attraction of adult chironomid midges to light was previously reported by Frost
(1957) and Belton and Pucat (1967). However, the only qualitative and quantitative data
on the attraction of adult midges to light are those of Ali et al. (1984) showing that
under laboratory conditions, G. paripes, C. crassicaudatus, and Polypedilum halterale
(Coquillett) responded more to the quantity (power or intensity) than to the quality
(color or wavelength) of light. The field observations made in the present study are in
agreement with the laboratory observations of Ali et al. (1984); the most predominant
species, G. paripes showed maximum attraction to the source of highest light intensity.
Other species also showed somewhat similar behavior of attraction although some pref-
erences may have been color or wavelength specific. This study has also revealed that
the adult chironomid response to white incandescent and white fluorescent lights of
equal wattage was similar.
Observations made in the present study are of practical significance for the develop-
ment of an integrated control program against pestiferous chironomid midges. Many
large lakes in the central Florida area support dense populations of G. paripes and C.
crassicaudatus (Ali and Baggs 1982). By the use of proper lights on and/or around the
lakes, adults of these species can be drawn to uninhabited or relatively less urbanized
areas for suitable midge control.


649












Florida Entomologist 69(4)


ACKNOWLEDIGMENTS

This article is Fla. Agric. Exp. Sta. Journal Series No. 7222, and is dedicated to Dr.
John F. Darby, former Director, Central Florida Research and Education Center,
IFAS, University of Florida, upon his retirement.

REFERENCES CITED

ALI, A. 1980. Nuisance chironomids and their control: a review. Bull. Entomol. Soc.
Am. 26: 3-16.
ALI, A., AND R. D. BAGGS. 1982. Seasonal changes of chironomid populations in a
shallow natural lake and in a man-made water cooling reservoir in central Florida.
Mosq. News 42: 76-85.
ALI, A., AND M. S. MULLA. 1977a. Chemical control of nuisance midges in the Santa
Ana River Basin, Southern California. J. Econ. Entomol. 70: 191-195.
ALI, A., AND M. S. MULLA. 1977b. The IGR diflubenzuron and organophosphorus
insecticides against nuisance midges in man-made residential-recreational lakes.
J. Econ. Entomol. 70: 571-577.
ALI, A., S. R. STAFFORD, R. C. FOWLER, AND B. H. STANLEY. 1984. Attraction
of adult Chironomidae (Diptera) to incandescent light under laboratory condi-
tions. Environ. Entomol. 13: 1004-1009.
ANONYMOUS. 1977. Economic impact statement, blind mosquito (midge) task force,
Greater Sanford Chamber of Commerce, Seminole County Florida. 6 pp.
ANONYMOUS. 1978. Light and color. Publ. TP-119, General Electric Co., Cleveland,
Ohio.
BELTON, P., AND A. PUCAT. 1967. A comparison of different lights in traps for
Culicoides (Diptera: Ceratopogonidae). Can. Entomol. 99: 267-272.
FROST, S. W. 1957. The Pennsylvania insect trap. J. Econ. Entomol. 50: 287-292.
GLICK, P. A., AND J. P. HOLLINGSWORTH. 1954. Response of the pink bollworm
moth to certain ultraviolet and visible radiation. J. Econ. Entomol. 47: 81-86.
HEINTON, T. E. 1974. Summary of investigations of electric insect traps. Agric. Res.
Serv. Tech. Bull. 1498. U. S. Dept. of Agriculture, Washington, D. C.
HOLLINGSWORTH, J. P. 1961. Relation of wavelength to insect response. In Response
of insects to induced light. U. S. Dep. Agric., Agric. Res. Serv. 20-10: 9-25.
HOLLINGSWORTH, J. P., R. L. WRIGHT, AND D. A. LINDQUIST. 1964. Spectral
response characteristics of the boll weevil. J. Econ. Entomol. 57: 38-41.
MAGNUM, C. L., AND P. S. CALLAHAN. 1968. Attraction of near-infrared radiation
to Aedes aegypti (Linnaeus). J. Econ. Entomol. 61: 36-37.
MCCULLAGH, P., AND J. A. NELDER. 1983. Generalized linear models. Chapman
and Hall, NY. 261 pp.
MENEAR, J. R. 1961. Response of tobacco and tomato hornworm moths to monoc-
hromatic radiation in the near ultra-violet. M.S. thesis, Virginia Polytechnic In-
stitute & State University, Blacksburg, Va.
MULHERN, T. D. 1942. New Jersey mechanical trap for mosquito surveys. N. J.
Agric. Exp. Stn. Circ. 421.
POLLS, I., B. GREENBERG, AND C. LUE-HING. 1975. Control of nuisance midges
in a channel receiving treated municipal sewage. Mosq. News 35: 533-537.
TAYLOR, J. G., AND H. O. DEAY. 1950. Electric lamps and traps in corn borer
control. Agric. Eng. 31: 503-532.
WILLIAMS, C. B., AND L. DAVIS. 1951. Simuliidae attracted at night to a trap using
ultraviolet light. Nature (London) 179: 924-925.


650


December, 1986











Prokopy et al.: Medfly Fruit-Forr'y;,,g


FRUIT-FORAGING BEHAVIOR OF MEDITERRANEAN FRUIT
FLY FEMALES ON HOST AND NON-HOST PLANTS

RONALD J. PROKOPY AND DANIEL R. PAPAJ
Department of Entomology, University of Massachusetts
Amherst, MA 01003, USA
and
TIM T. Y. WONG
Tropical Fruit and Vegetable Research Laboratory,
Agricultural Research Service, USDA, Honolulu, Hawaii, 96804, USA

ABSTRACT

The hypothesis that female fruit flies (Family Tephritidae) remain longer and dis-
cover fruit more readily on host plants than on non-host plants was examined. Naive,
gravid females of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), were
released on non-fruiting host and non-host plants in a large outdoor cage. Their behavior
within each plant was characterized over 15 min. using a stopwatch and tape recorder.
Although medfly females remained longer on one non-fruiting host (mandarin orange)
than on one non-fruiting non-host (Norfolk pine), they stayed just as long on another
non-fruiting non-host (eldorado) as on mandarin orange. Also, females tended to leave
a second non-fruiting host (tomato) more rapidly than eldorado. In one experiment
where host fruit (kumquats) were artificially placed on the plant branches, females were
just as likely to find such fruit on non-host as on host plants. Thus, medfly females may
forage for fruit on host and certain non-host plants in a like fashion, with vegetative
characteristics of host plants not necessarily promoting greater fruit finding efficiency.

RESUME

Se examine la hip6tesis que las moscas de frutas (Familia Tephritidae) se mantienen
mas tiempo y descubren frutas mas rapidamente en plants hospederas que en plants
no-hospederas. Hembras prefadas de la mosca mediterranea, Ceratitis capitata (Wiede-
man), fueron liberadas en hospederos sin fruto y en plants ho-hospederas en una jaula
al'exterior. Se estudi6 su comportamiento en cada plant durante 15 min. usando un
reloj cron6metro y una grabadora. Aunque hembras de la mosca mediterranea es-
tuvieron mas tiempo en el hospedero sin fruto (naranja mandarina) que en el no-hosped-
ero sin fruto (pino Norfolk), ellas estuvieron tanto tiempo en otro no-hospedero sin fruto
(eldorado) como en naranja mandarina. Las hembras tambi6n tendieron a dejar mas
rapido un segundo hospedero sin fruto (tomate) que a eldorado. En un experiment
donde frutas hospedera (kumquats) se pusieron artificialmente en las ramas de las plan-
tas, las hembras encontraron con la misma probabilidad las frutas en plants no-hos-
pederas que en plants hospederas. De aqui que hembras de la mosca mediterranea
puede forrajear frutas en hospederos y en ciertas plants no-hospederas de la misma
forma, con las caracteristicas vegetativas de la plant hospedera no necesariamente
promoviendo la eficiencia de encontrar las frutas.



Reproductively mature female fruit flies (Family Tephritidae) are often found in
greater numbers on fruiting host plants than on non-host plants (eg. Boller et al. 1971,
Prokopy, unpub. data). This distribution may arise from either or both of two causes:
(a) females arrive on fruiting host plants at a greater rate than on non-host plants; and
(b) females leave fruiting host plants at a lesser rate than non-host plants. The latter











Florida Entomologist 69(4)


mechanism is important in the apple maggot fly, Rhagoletis pomonella (Walsh). Thus,
Diehl et al. (1986) found that apple maggot females spent less total time on certain
non-hosts (such as tomato and pine) than on hosts (apple and hawthorn). In addition,
females discovered fewer artificially-positioned fruit on these particular non-hosts than
on hosts. Interestingly, apple maggot females foraged no differently on other non-hosts
(eg. birch) than on hosts.
No similar information has been available for tephritid flies differing in degree of
specialization from the apple maggot fly. In contrast to R. pomonella, which infests
fruit of fewer than 10 species in a single family, the Mediterranean fruit fly (medfly),
Ceratitis capitata (Weidemann), is known to attack more than 250 species of fruit and
vegetables in numerous families. In this study, we asked whether or not female medflies
remained longer on host plants than on non-host plants, and whether females were more
likely to find artificially-positioned fruit on hosts plants than on non-hosts.

MATERIALS AND METHODS

All medflies tested were wild, having originated from larvae that infested field-col-
lected fruit of unsprayed loquats, Eriobotryia japonica, taken from the Kula area of
the island of Maui in Hawaii. This population of medflies had access to a wide range of
intermixed host and non-host plants (Wong et al. 1983). Upon eclosion, females were
held together with males in cages supplied with food (yeast hydrolysate and sucrose)
and water under laboratory conditions (temperature = ca. 25'C, relative humidity =
ca. 40%, daylength = ca. 13 hours). To test whether a female was in a physiological
state conducive to oviposition site foraging, each female, when 12-17 days old, was
offered a host fruit (kumquat, Fortunella japonica). Only those that attempted ovipos-
ition into kumquat were assayed in the experiments (= ca. 95% of all enclosed females).
For ca. 24 h prior to assay, each female was confined alone in a small lab cage.
All test plants were obtained from local (Hawaiian) nurseries and were rooted in
pots. For Exp. 1, plants included: mandarin orange Citrus reticulata (Rutaceae); el-
dorado, Graptophyllum pictum (Acanthaceae); tomato, Lycopersicon esculentum (Sol-
anaceae); and Norfolk pine, Araucaria heterophylla (Araucariaceae). For Exp. 2, which
was conceived and conducted after Exp. 1 had been completed, plants included: sweet
pepper, Capsicum frutescens var. grossum (Solanaceae); eggplant, Solanum
melongena var. esculentum (Solanaceae) and tomato. Mandarin orange is a major host
plant of medflies (i.e. it supports substantial populations in nature); tomato, pepper,
and eggplant are minor hosts (i.e. each supports low or very low populations in nature);
and eldorado and Norfolk pine are non-hosts (Back and Pemberton 1918). Two speci-
mens of each plant species were employed. Plants within an experiment were approxi-
mately equal in canopy volume. For each plant, we measured the height of the canopy
lowermostt to uppermost leaf), canopy diameter, total number of leaves, and the surface
area of each leaf. In Exp. la and Exp. 2, none of the plants harbored fruit. In Exp. Ib,
we hung by wire 6 evenly-spaced kumquat fruit (20 mm diam) from each plant. Before
being used in tests, the foliage and stems of all plants were rinsed gently but thoroughly
with water to remove any debris.
All tests were conducted in cylindrical 3.5 x 3.5 x 3m clear-nylon-screen field cages
on the grounds of the USDA Tropical Fruit and Vegetable Research Laboratory in
Honolulu. A single plant was positioned 1m above ground at the center of the cage. A
single female was transferred gently (using a small piece of moist filter paper on a
probe) from a lab cage onto a leaf at the center of the plant canopy. To ensure uniformity
of assay procedure among replicates, flies were always released onto the same leaf of
a test plant. Using a stopwatch and tape recorder, we monitored the following behaviors


December, 1986











Prokopy et al.: Medfly Fruit-Foraging 653

f
of each fly: number of times an individual flew or walked to a leaf or stem; total time
spent moving, grooming, or resting; and total time on the plant. We also recorded time
spent feeding (because we observed this in less than 2% of all flies assayed, we excluded
time spent feeding from data analysis). In Exp. Ib, we determined whether or not an
individual visited a fruit. A test was terminated when a fly flew from the plant (none
crawled off), when 15 min elapsed, or when a fly alighted on a fruit (Exp. Ib). Each fly
was tested only once, following which it was immediately offered a kumquat fruit. Those
females that did not attempt oviposition into such fruit after testing on a plant were
considered as not having been in a physiological state conducive to oviposition site
foraging and were excluded from data analysis (= ca. 9% of all individuals tested). To
minimize experimental error owing to variation in temperature (range 23-32C) and
wind speed (range 0-11 km/hr) during test periods (9am-4pm), we rotated the sequence
in which plants were tested such that each treatment was offered once before the same
treatment was offered again.
The proportion of females visiting at least one leaf or stem (excluding the leaf on
which the female was released) was compared among plant species using a G-test at the
0.01 level. To compare treatment medians of time spent moving, grooming, resting, and
total time on plants, we used a median test (Siegel 1956) at the 0.01 level.

RESULTS

In Exp. la, in which no fruit were placed on the test plants, medfly females spent
significantly less time grooming, less time resting, and less total time on Norfolk pine
than on orange, eldorado, or tomato, among which there were no significant differences
(Table 1). Although there were no significant differences in female response to these 4
plant species with respect to time spent moving, movement time was least on Norfolk
pine and greatest on orange. Significantly fewer females flew to Norfolk pine leaves
and stems than to orange or tomato leaves and stems. Otherwise, there were no signif-
icant differences among plants in proportion of females flying or walking to leaves and
stems.
In Exp. Ib, in which 6 kumquat fruit were hung from each plant assessed in Exp.
la, there were no significant differences among plants with regard to proportion of
females alighting on fruit (Table 1). In fact, the smallest percentage of fruit-alighting
females was on orange (a major host plant) and the greatest on eldorado (a non-host
plant). Those females which did alight on a fruit were excluded from further considera-
tion in data analysis (we presume such females were a random sample of the population
of all females available for testing, but we do not know this for certain). Among those
females which did not alight on a fruit during the 15-min test period, significantly less
total time was spent on Norfolk pine than on tomato, and significantly less total time
on tomato than on orange or eldorado. Other parameters of female response were very
similar to those in Exp. la (Table 1).
In Exp. 2, in which no fruit were placed on the test plants and in which the plants
were considerably smaller (Table 3) than those assayed in Exp. 1, medfly females spent
significantly less time grooming, less time resting, and less total time on tomato than
on eggplant or pepper, between which there were no significant differences (Table 2).
There were no significant differences in female response to these 3 plant species in time
spent moving on a plant, or proportion of females visiting leaves and stems by flight or
by walking.






















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Prokopy et al.: Medfly Fruit-Foraging 655


TABLE 2. BEHAVIOR OF MEDFLY FEMALES RELEASED INDIVIDUALLY ON
POTTED HOST PLANTS WITHOUT FRUIT (EXP. 2).

% ? visiting
leaves and stems by Median time (sec) on plant
No. 9
Plant assayed Flight* Walking Moving Grooming Resting Total

Eggplant 24 50a 13a 19a 64a 61a 197a
Pepper 24 67a 25a 14a 79a 73a 161a
Tomato 24 58a 13a 11a Ob 21b 47b

*Values in each column not followed by the same letter are significantly different at the 0.01 level.

TABLE 3. MEASUREMENTS OF PHYSICAL CHARACTERISTICS OF PLANTS ON
WHICH MEDFLY BEHAVIOR WAS ASSESSED (VALUES = AVG. OF 2
PLANTS TESTED OF EACH SPECIES).

Plant canopy (cm)
Species of Total no. Mean leaf
Exp. Plant Height Diameter leaves* area (cm2)*

1 Orange 60 55 145 25
1 Eldorado 67 60 109 35
1 Tomato 71 74 350 16
1 Pine 68 75 512 12

2 Eggplant 21 29 12 71
2 Pepper 18 29 21 35
2 Tomato 27 28 67 14

*For the purposes of this table only, a "leaf" on Norfolk pine was not considered to be an individual needle but
rather the number and dimensions (needle tip to needle tip) of branchlets that bore the needles.

DISCUSSION

The interpretation of our findings is necessarily restricted by the artificial conditions
under which flies were tested. All flies assayed were taken from a small laboratory
cage, placed on a relatively small plant, and permitted only a short time period in which
to forage for egg-laying sites on the plant. Nevertheless, certain patterns emerge from
the data that may pertain to fly behavior in nature.
Our results do not permit us to conclude that gravid medfly females in nature, which
occur in greater numbers on host plants than on non-host plants (Prokopy unpub. data)
do so simply because females tend to remain longer on non-fruiting hosts than on non-
fruiting non-hosts. Indeed, although medfly females did remain longer on one non-fruit-
ing host (orange) than on one non-fruiting non-host (Norfolk pine), females stayed just
as long on another non-fruiting non-host (eldorado) as on orange. Moreover, females
tended to leave a second non-fruiting host (tomato) more rapidly than eldorado.
Medfly females consistently were not inclined to spend very much time on Norfolk
pine plants (a non-host) either moving, grooming, or resting. On Norfolk pine, few
females flew to another "leaf" (see Table 3 for our definition of a Norfolk pine leaf).
Whatever leaf to leaf movement did occur was primarily via walking. Indeed, it ap-
peared to us that medfly females on Norfolk pine were in a continuously agitated state.










Florida Entomologist 69(4)


They had great difficulty grasping a firm hold on the Norfolk pine needles. If this
difficulty arose from chemical properties of Norfolk pine repellent to the flies, we would
have expected to observe a great deal more grooming of the tarsi, antennae or mouth-
parts than we did. Hence, we suspect that the physical properties of Norfolk pine
needles, particularly narrowness, were responsible for eliciting rapid emigration of
medflies from Norfolk pine plants. Similarly, gravid apple maggot females were found
to emigrate comparatively rapidly from white pine plants (a non-host) on which they
were released, again apparently on account of the narrowness of the needles (Diehl et
al. 1986). Interestingly, Lawton (1983) noted that British umbellifers that produce finely
divided leaves have relatively impoverished agromyzid fly faunas compared with umbel-
lifers that produce broader leaves. Possibly leaf geometry has a more profound effect
on herbivore resource utilization than is presently realized.
In every experiment, medfly females spent less total time (though not always sig-
nificantly so) on tomato plants than on any other plants tested except Norfolk pine. This
was due to the comparatively short time medfly females spent grooming and resting,
as well as moving, on tomato. In fact, the median time spent moving on tomato was
remarkably similar in all experiments, as was the median time spent grooming or rest-
ing, even though the tomato plants in Exp. 1 were much larger and bore many more
leaves than those in Exp. 2. Perhaps the relatively small size of tomato leaves (Table
3) and/or the chemical properties of tomato foliage were not conducive to greater dura-
tion of fly residence, even though tomato fruit is a minor host of medflies. Similarly,
gravid apple maggot females were found to leave tomato plants (a non-host) rapidly
(Diehl et al. 1986). In that species, however, the females were observed to be almost
continuously grooming from time of release onto tomato foliage until time of departure,
and frequently exhibited signs of apparent neurological impairment (eg. twitching).
Evidently medfly females are at least somewhat better adapted to foraging on tomato
than are apple maggot flies.
Medfly females behaved on eldorado (a non-host) much as they did on orange (a
major host). Test plants of each of these species had similar numbers and sizes of leaves
(Table 3). Analogously, apple maggot flies behaved in birch trees (a non-host) very
much as they did in apple trees (a major host) in experiments in which the test plants
of each type bore similar numbers and sizes of leaves (Diehl et al. 1986).
While non-fruiting host plants may or may not arrest medflies to a significantly
greater degree than non-fruiting non-host plants, the presence of host fruit on host
plants could increase residence time on hosts over that on non-hosts. Previous studies
indicated that fruit discovery and oviposition extended the time over which a female
apple maggot fly foraged within a host plant before she emigrated (Roitberg et al. 1982).
Since the proportion of medflies finding fruit in host plants was no different than the
proportion finding fruit in non-hosts in our present study, the effect of host fruit on fly
residence time does not necessarily depend on the host status of the plant in which the
fruit is positioned. In other words, a gravid female arriving on either a host plant such
as orange or a non-host plant such as eldorado is equally likely to search for and respond
positively to host fruit. Once having oviposited in a fruit, it is likely a female will remain
longer and continue to forage for fruit on plants of either species.
These inferences are supported by what is known about the stimuli eliciting alight-
ment on a fruit. Medfly females approach and alight on fruit primarily on the basis of
shape, size, and color (Nakagawa et al. 1978, Cytrynowicz et al. 1982), stimuli that are
specific to fruit and bear little or no relation to stimuli emanating from vegetative parts
of a host plant. In fact, medfly females alight frequently on host fruit hung in small
cages devoid of other plant material. Thus, the host plant itself apparently does not
necessarily offer a special context in which a medfly female will forage more intensively
for host fruit.


656


December, 1986











Prokopy et al.: Medfly Fruit-Foraging 657

The efficiency with which flies find fruit, while not necessarily depending on a plant's
host status, probably does depend on the detectability of fruit against the foliar back-
ground of particular plant species. To our eyes, mandarin orange plant foliage hid
kumquat fruit from view more often than did the foliage of any other plant species, host
or non-host. Likewise, apple maggot flies are much less apt to find fruit when fruit is
hidden by adjacent foliage (Drummond et al. 1984).
In sum, the combined findings of this investigation and the study of Diehl et al.(1986)
suggest that medfly and apple maggot fly females (and perhaps females of other tep-
hritid species (eg. Dacus latifrons-Prokopy et al., unpub. data) may at the outset forage
for fruit on host and certain non-host plants in a like fashion. Also, the vegetative
characteristics of host plants do not necessarily promote fruit-finding efficiency in either
the medfly or apple maggot fly. Better quantification of the influence of plant vegetative
structures on tephritid fruit-foraging behavior will require the development of artificial
plants whose physical and chemical characteristics can be manipulated in a more precise
fashion than is possible with living plants.

ACKNOWLEDGMENTS

We thank Jon Nishimoto for his assistance in providing flies and plants and Julia
Connelly for typing the manuscript. This work was supported by the Science and Edu-
cation Administration of the USDA under grant 8200184 from the Competitive Research
Grants Office and by Massachusetts Agricultural Experiment Station Project 599.

REFERENCES CITED

BACK, E. A., AND C. E. PEMBERTON. 1918. The Mediterranean fruit fly in Hawaii.
USDA Bull. 536: 118 p.
BOLLER, E. F., A. HAISCH, AND R. J. PROKOPY. 1971. Sterile-insect release method
against Rhagoletis cerasi: preparatory ecological and behavioral studies, pp. 77-
86 In Sterility Principle for Insect Control or Eradication. Proc. Sympos. Vienna
1970. IAEA, Vienna.
CYTRYNOWICZ, M., J. S. MORGANTE, AND H. M. DE SOUZA. 1984. Visual responses
of South American fruit flies and Mediterranean fruit flies to colored rectangles
and spheres. Env. Ent. 11: 1202-1210.
DIEHL, S. R., R. J. PROKOPY, AND S. HENDERSON. 1986. The role of stimuli as-
sociated with branches and foliage in host selection by Rhagoletis pomonella. pp
191-196 In: R. Cavalloro (ed.). Fruit Flies of Economic Importance 87. Balkema,
Netherlands.
DRUMMOND, F., E. GRODEN, AND R. J. PROKOPY. 1984. Comparative efficacy and
optimal positioning of traps for monitoring apple maggot flies. Env. Ent. 13:
232-235.
LAWTON, J. H. 1983. Plant architecture and the diversity of phytophagous insects.
Ann. Rev. Ent. 28: 23-29.
NAGKAGAWA, S., R. J. PROKOPY, T. T. Y. WONG, J. R. ZIEGLER, S. M. MITCHELL,
T. D. URAGO, AND E. J. HARRIS. 1978. Visual orientation of Ceratitis capitata
flies to fruit models. Ent. Exp. Appl. 24: 193-198.
ROITBERG, B. D., J. C. VAN LENTEREN, J. J. M. VAN ALPHEN, F. GALIS, AND R.
J. PROKOPY. 1982. Foraging behavior of Rhagoletis pomonella, a parasite of
hawthorn. J. Anim. Ecol. 51: 307-325.
SIEGEL, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-Hill,
New York. 312.
WONG, T. T. Y., J. I. NISHIMOTO, AND N. MOCHIZUKI. 1983. Infestation patterns
of the Mediterranean fruit fly and the Oriental fruit fly in the Kula area of Maui,
Hawaii. Env. Ent. 12: 1031-1039.










Florida Entomologist 69(4)


DESCRIPTION OF THE MALE OF
PSEUDOMETHOCA OCULATA (BANKS)
(HYMENOPTERA: MUTILLIDAE)

MARK DEYRUP
Archbold Biological Station
P. O. Box 2057
Lake Placid, FL 33852
and
DONALD MANLEY
Clemson University
Pee Dee Research and Education Center
P. O. Box 5809
Florence, SC 29502

ABSTRACT

The previously unknown male of Pseudomethoca oculata (Banks) is described from
Highlands Co., Florida.

RESUME

Se describe el macho previamente desconocido de Pseudomethoca oculata (Banks)
del condado de Highland en la Florida.



Pseudomethoca oculata (Banks) was described from female specimens in 1921. It
occurs on the Coastal Plain of the southeastern United States from North Carolina to
southern Florida (Mickel 1924). Although females of this species are sometimes rela-
tively abundant, there are no reports of males.
During a study of the Mutillidae of the Archbold Biological Station (Highlands
County, Florida), we obtained from malaise traps a large series of male Pseudomethoca
that differ markedly from any described species. We believe that these are males of P.
oculata because: 1) female P. oculata were abundant at the site of the malaise traps at
the time the males were collected; 2) intensive collecting did not produce specimens of
any undescribed female Pseudomethoca, though we did collect both males and females
of four additional described species of Pseudomethoca; 3) it is much simpler to assume
that these abundant males are associated with the abundant females of P. oculata than
to assume that the males represent an abundant undescribed species whose females
have not yet been observed.
DIAGNOSIS: Distinguished from other North American species by the shape of the
parameres, which are elongate, slender, strongly arcuate, as in Figures 1 and 2, and
by the tegulae (Fig. 3), which are strongly bent downward posteriorly and coarsely
punctate throughout. Otherwise superficially resembles P. simillima (Smith), as dis-
cussed below.
DESCRIPTION OF ALLOTYPE: Black, with dorsal areas of thorax and dorsal area of
propodeum dark red; gastral segments 1 and 2 reddish orange; segments 3-4 dark red.
Long hairs on head, body, appendages black, short hairs on tarsi dark grey, pygidial
hairs light grey.


December, 1986











Deyrup & Manley: Male Pseudomethoca oculata


Head clothed with long sparse, erect, black hairs; entire head confluently punctate
except for finely granulate antennal fossae, finely granulate narrow postocular area,
small polished area anterior to median ocellus, small polished area laterad of lateral
ocellus. Apical half of clypeus transversely concave, with a pair of well separated median
teeth. Antennal scape coarsely punctate, carinate ventrally; pedicel and flagellum finely
punctate and hairy as usual in genus.
Thorax covered with sparse long black erect hairs; thorax coarsely, confluently
punctured, except for metapleuron, which is polished, minutely punctate with sparse
silver decumbent hairs on lower half. Tegulae coarsely punctate strongly bent down-
ward posteriorly, and covered with coarse punctures. Mesosternum posteriorly with a
fine brush of hyaline hairs, contrasting strongly with coarse black hairs of adjacent
coxae, and not overlapping into metasternal pit.
Wings completely, evenly infuscated, as usual for genus.
Legs with sparse long black hairs. Femora externally with poorly defined well sepa-
rated small punctures, interspersed with punctures about three times as large. Tibiae
externally with confluent poorly-defined large and small interspersed punctures. Tarsi
densely covered with short dark gray hairs.
Propodeum very coarsely reticulate, with very sparse, long, black, erect hairs.
Gaster generally covered with long black erect hairs. Discal areas terga 1 and 2 with
punctures whose actual pits are separated by more than twice their diameters, each pit
in a shallow vague depression. Tergum 1 with a more densely punctate subapical band;
terga 2-6 densely punctate and hairy apically. Pygidium densely punctate on sides and
on basal half of median area, with hyaline short recumbant hairs; apical median area
smooth and glabrous, not extensively overlapped by basal hairs. Sternum 2 with large
shallow punctures whose pits are generally separated by their own widths. Sterna 3-6
with apical bands of small dense punctures and black hairs; last visible sternum concave,
rugosely punctate, with short, sparse, hyaline hairs.
Genitalia as in Figures 1, 2.
ALLOTYPE: Male, Archbold Biological Station, Lake Placid, Highlands Co., Florida;
2 Oct. 1983; Malaise trap; M. Deyrup. Deposited in Florida State Collection of Ar-
thropods (Galnesville).
PARATYPES: 78 males, Archbold Biological Station, Lake Placid, Highlands Co.,
Florida; Malaise traps; M. Deyrup; 28-IX-83 (3), 30-IX-83 (3), 2-X-83 (2), 4-X-83 (4),
6-X-83 (2), 8-X-83 (3), 16-X-83, 18-X-83, 20-X-83 (4), 22-X-83 (5), 24-X-83, 26-X-83 (4),
30-X-83 (4), 1-XI-83 (2), 15-XI-83 (2), 21-XI-83, 23-XI-83 (4), 27-XI-83 (2), 3-XII-83 (2),
5-XII-83 (2), 9-XII-83, 10-IX-84, 12-IX-84, 24-IX-84, 26-IX-84, 5-X-84, 8-X-84 (2), 10-X-
84, 12-X-84 (2), 15-X-84 (2), 16-X-84, 17-X-84 (2), 18-X-84, 30-X-84 (2), 1-XI-84, 7-XI-84,
9-XI-84, 19-XI-84, 23-IX-85 (3), 7-X-85. The first 10 specimens listed above are depo-
sited in the U.S. National Museum, the next 10 in the Florida State Collection of
Arthropods (Gainesville), the next 20 in the collection of Donald Manley, the remainder
in the collection of the Archbold Biological Station.
VARIATION: The extent of dark red on the thorax is variable. Most specimens have
a black head, dark red dorsal areas on the thorax and propodeum; a few specimens have
a completely black head, thorax and propodeum; one specimen has the head as well as
the thorax dark red. There is no evidence of a seasonal pattern in this variation. Body
length, which depends somewhat on the telescoping of the gastral segments, varies
from about 8.5 mm to 12.5 mm. The length of the forewing varies from 6.3 mm to 8.8
Males of P. oculata strongly resemble those of P. simillima. Shared character states
1) absence of decumbent hairs on the head and body, 2) the shape and sculpture of the
clypeus, 3) the antennal scape carinate, 4) sculpture of the thorax, propodeum, and
gaster, and 5) gastral segments basally red. P. simillima differs in having 1) short


659










Florida Entomologist 69(4)


Fig. 1. Pseudomethoca oculata (Banks), male gentalia, dorsal view.
Fig. 2. Same, lateral view.
Fig. 3. Pseudomethoca oculata (Banks), left tegula.
Fig. 4. Pseudomethoca simillima (Smith), male genitalia, dorsal view.
Fig. 5. Same, lateral view.
Fig. 6. Pseudomethoca simillima (Smith), left tegula.


660


December, 1986











Deyrup & Manley: Male Pseudomethoca oculata


parameres that are not arcuate in dorsal view (Figs. 4, 5), 2) tegulae smooth and shining
on posterior half (Fig. 6), 3) consistently black thorax and propodeum, 4) long erect
hyaline hairs on various parts of the legs and body, and 5) punctures on the occiput
slightly more separated.
Both P. oculata and P. simillima are easily distinguished from males of P. oceola
(Blake) by the presence of very dense, long pale hairs beneath the femora of the latter.

ACKNOWLEDGMENTS

An anonymous reviewer and Dr. William E. Ferguson (San Jose State University)
provided useful suggestions on the manuscript.

REFERENCES CITED

BANKS, N. 1921. New Neartic fossorial Hymenoptera. Ann. Entomol. Soc. Amer. 14:
24-26.
MICKEL, C. E. 1924. A revision of the mutillid wasps of the genera Myrmilloides and
Pseudomethoca occurring in American north of Mexico. Proc. U.S. Nat. Mus.
64(15): 1-51.





AULACOBLISSUS, A NEW GENUS OF
MICROPTEROUS BLISSINAE FROM VENEZUELA
(HEMIPTERA: LYGAEIDAE)

JAMES A. SLATER
Department of Ecology and Evolutionary Biology
University of Connecticut
Storrs, Connecticut 06268

ABSTRACT
Aulacoblissus brailovskyi is described as a new genus and species of Blissinae from
the mountains of Venezuela. It is the most micropterous blissine lygaeid known. The
relationships of the genus are discussed and it is concluded that it is most closely related
to Heteroblissus Barber. Distinguishing characters are given. Illustrations include a
dorsal view of the insect and details of the pygophore, paramere, and sperm reservoir.

RESUME

Se describe a Aulacoblissus brailovski como un nuevo genero y especie de Blissinae
de las montafia de Venezuela. Es el mas conocido micr6ptero blissine lygaeid. Se discu-
ten las relaciones del genero y se concluye que esta muy cercamente relacionado a
Heteroblissus Barber. Se proven caracteristicas distintivas. Ilustraciones incluyen una
vista dorsal del insecto y detalles de la pygophore, paramere, y del dep6sito de esper-
mas.











Florida Entomologist 69(4)


Aulacoblissus, new genus

Body robust, sublinear. Surface completely non-pruinose (except for slight vestiges
of pruinosity mesally on thoracic venter), shining or subshining. Metathoracic scent
gland auricle large, broadly rounded, ear-like. Fore femora moderately incrassate, each
armed below distally with a single small spine. Fore coxal cavities open. Fore tibiae
slightly enlarged distally, not fossorial. Ocelli absent. Eyes set on short broad lateral
head extensions. Body surface coarsely and deeply crenulate with large foveate
punctures.
Paramere (Fig. 5) blade elongate, slender, outer projection prominent and nearly
rectangular without a well differentiated inner projection. Sperm reservoir (Figs. 3, 4),
small, bulb broadened to a subtruncate distal end; wings slender and strap-like not
strongly bent ventrad. Pygophore opening as in Figure 2.
Type species: Aulacoblissus brailovskyi, new species.
This genus will run in my (Slater 1979) key to world blissine genera to the South
African genus Macchiademus to which it is not closely related phylogenetically.
The extreme microptery suggests a possible relationship to such Neotropical genera
as Praetorblissus Slater and Heteroblissus Barber. Both of these genera have open fore
coxal cavities but also have multispinose fore femora. Species of Praetorblissus also
have a derived metathoracic scent gland auricle which curves strongly forward and also
usually have spines on the middle and hind femora.
Although it would be interesting to see nymphs and (if they exist) macropters,
present evidence suggests that Heteroblissus is the sister genus for the following
reasons: 1. The body surface of both taxa lacks pruinosity and is crenulate and coarsely
punctate. 2. The metathoracic scent gland auricle is broadly rounded and ear-like. 3.
The bulb of the sperm reservoir is broadened distally and almost truncate along its
distal margin.
Heteroblissus has a series of elongate striae on the abdominal venter (presumably
stridulatory structures), broadened non-strap-like wings on the sperm reservoir, multi-
spined fore femora, more strongly produced eyes and a series of stout spines along the
shafts of the fore tibiae. All of these appear to be derived features and will readily serve
to distinguish the two genera.
Aulacoblissus shows the greatest degree of wing reduction yet known to occur in
the Blissinae. It is another striking example of the richness and highly specialized
nature of the Neotropical montane lygaeid fauna.

Aulacoblissus brailovskyi, new species
(Fig. 1)

Body above and below nearly uniformly yellowish brown becoming almost black
along posterior margin of scutellum and near extreme distal end of each fourth antennal
segment. Evaporative area dull gray, strongly contrasting with adjacent shining red
brown coloration, extending over inner 2/3 of anterior lobe of metapleuron, onto post-
erior edge of mesopleuron and completely across metasternum to form a continuous
area across venter of body. Punctures on body surface very large and coarse, sometimes
anastomosing with the deeply crenulate and striated areas present over most of body
surface. A few scattered rather elongate hairs present on dorsal surface and on abdom-
inal venter.

Head non-declivent. Tylus reaching middle of first antennal segment. Vertex slightly
convex. Length head 0.70, width 0.94, interocular space 0.62. Lateral margins of pro-


December, 1986










Slater: Micropterous Blissinae 663


-----,
C~------ `;


Fig. 1. Aulacoblissus brailovskyi Dorsal View.


r
I





p1/


f









&e









Florida Entomologist 69(4)


K'
K

127


Fig. 2. Aulacoblissus brailovskyi Pygophore.
Fig. 3. Aulacoblissus brailovskyi Sperm Reservoir Dorsal View.
Fig. 4. Aulacoblissus brailovskyi Sperm Reservoir Lateral View.
Fig. 5. Aulaooblissus brailovskyi Paramere.


notum evenly convex, broadest at level of calli; posterior margin straight. Calli deeply
impressed, appearing almost foveate. Length pronotum 0.76, maximum width 1.30.
Scutellum lacking a median elevation. Scutellar length 0.38, width 0.92. Mesothoracic
wings reduced to a pair of minute tumid yellow lumps at junction of pronotum, scutel-
lum, and metanotum. Abdomen broad but nearly parallel-sided. Connexivum large with
very prominent spiracles placed close to lateral margins of abdomen. Length abdomen
3.12. Labium extending posteriorly to middle of mesosternum, well surpassing fore
coxae. Length labial segments I 0.30, II 0.38, III 0.30, IV 0.34. First antennal segment
broad and rounded, segments II and III terete, segment IV fusiform. Length antennal
segments I 0.30, II 0.60, III 0.70, IV 0.88. Total body length 4.96.


December, 1986







Slater: Micropterous Blissinae


Holotype: male. VENEZUELA: Pmo. de Guaramacal, 3000 m., Bocono, Edo.
Trujillo, Bordon 7-VIII-1981. In Instituto Biologia, UNAM, Mexico, D.F.
It is a pleasure to dedicate this striking new species to Dr. Harry Brailovsky who
is contributing so much to our knowledge of Neotropical Lygaeidae.


ACKNOWLEDGEMENTS

I wish to thank Dr. Harry Brailovsky (Instituto Biologia, UNAM, Mexico) for bring-
ing this interesting insect to my attention: to Ms. Mary Jane Spring and Mrs. Elizabeth
Slater (University of Connecticut) for preparation of the illustrations and aid with the
manuscript respectively.
This work was supported in part by a grant from the National Science Foundation.

REFERENCE CITED

SLATER, J. A. The Systematics, Phylogeny, and Zoogeography of the Blissinae of the
World (Hemiptera, Lygaeidae). 1979. Bull. American Mus. Nat. Hist. 165: 1-180.





CORN RESIDUE AS AN OVERWINTERING SITE
FOR SPIDERS AND PREDACEOUS INSECTS IN FLORIDA

MICHAEL J. PLAGENS' AND WILLARD H. WHITCOMB
Dept. of Entomology & Nematology
University of Florida
Gainesville, FL 32610

ABSTRACT

Corn residue was used as an overwintering site by 24 species of spiders (Araneae)
and 25 predaceous Coleoptera, Hemiptera and Dermaptera in northern Florida. The
two principal microsites were the cavity formed between the leaf sheath and stem, and
between layers of imbricate bracts (husks) of shelled corncobs. Several pest species
were also found in these sites: chinch bugs (Blissus insularis), false chinch bugs
(Pachybrachius vinctus) and rice weevils (Sitophilus orvzae). When both sites were
available, shelled corn cobs were highly preferred by both predators and pests.


RESUME

En un studio hecho en el Norte de Florida, se encontr6 que los residues del cultivo
de maiz son utilizados como lugar de invernaci6n por 24 species de arafas (Araneae)
y por 25 species de depredadores pertenecientes a los ordenes Cole6ptera, Hemiptera
y Dermaptera. Dentro del ecosistema de la plant, los artr6podos se encontraron
habitando principalmente la cavidad localizada entire la uni6n de la hoja y el tallo, asi
como tambi6n entire las diferentes capas bracteas imbricadas de la tusa del maiz. En
estos mismos sitios fueron encontrados varias species de plagas, tales como: chinches
(Blissus insularis). falsos chinches (Pachybrachius vinctus) y gorgojos del arroz


'Present address: 4407 E. Lee St., Tucson, AZ, 85712.


665








666 Florida Entomologist 69(4) December, 1986


(Sitophilus orizae). Cuando ambos sitios estaban disponibles, tanto los depredadores
como los insects plaga se encontraron preferencialmente en la tusa de maiz.



Entomologists have learned to take advantage of the overwintering strategies used
by herbivorous insects such as the boll weevil (Slosser et al. 1984), the southwestern
corn borer (Archer et al. 1983), and the bean leaf beetle (Jeffords et al. 1983). Some
overwintering populations can be limited by tilling, which breaks up the post harvest
trash and exposes the insects, and/or by removing accumulated leaf litter in the borders
where the hibernating stages are found. Data on overwintering of insects is used to
time agricultural management decisions and to construct life tables that span more than
one year.
Likewise, overwintering strategies of predaceous arthropods could be used to manip-
ulate their populations; however, information about stages and sites is limited. Spiders,
due to their long life cycles and slow rates of reproduction, should be particularly
dependent on the adequacy of suitable overwintering sites. The purpose of this study
was to identify the type and location of such sites in the corn field. Other insects using
these same sites were also identified.

METHODS AND MATERIALS

LOCALITY DESCRIPTIONS

Overwintering predaceous insects and spiders were studied in two northern Florida
corn fields. The first was located near Archer, Florida, in southwestern Alachua County
and was studied during the winter of 1982-83. The second field was located on the Tall
Timbers Research Station in northern Leon County and was studied during the winter
of 1983-84. The Archer site was on a deep layer of fine sand that supported xeric
hammock or turkey oak scrub on near by, less disturbed sites. Peanuts, corn, watermel-
ons, and forage grasses were rotated on irrigated fields. Cattle, and to a lesser extent
hogs, were kept nearby, often put in the fields after harvest.
The Tall Timbers corn fields were planted on a sandy clay loam soil and were sur-
rounded by a fire-managed slash pine plantation maintained to encourage wildlife popu-
lations, particularly quail and deer. The post-harvest corn residue was accordingly left
in the field for the birds and other animals to glean.

METHODS AND MATERIALS

SPlant debris left in the fields following harvest included dead weeds, wind-blown
leaves, corn stubble, shattered stalks, and at Tall Timbers only, shelled corn cobs with
many of the bracts still attached. In a preliminary study at each field, these materials
were each gathered and then carefully examined in the field, by breaking apart the
collected materials and sorting them by hand over a white tray. Spiders and insects
were collected and preserved in alcohol. This established that spiders and insects were
abundant in the corn residue but were essentially absent in the other debris.
Further study was directed soley at the corn residue. At Archer the residue con-
sisted of the crown attached to a meter or less of broken stalk; 16 sets of 10 corn stalks
were examined for predators on three visits during December and January. The residue
left by a sheller picker at Tall Timbers also left shelled corn cobs with many of the husks











Plagens & Whitcomb: Overwintering Sites


still attached and covering the cobs. There, 24 sets of 10 corn stalks plus 10 cobs with
husks were examined as described above on three dates in January and February.
Within field sites were not selected at random because the harvesters distribute the
residue unevenly. All collections were made on cool days (<100C), therefore the location
of inactive spiders and insects in the stubble was noted.
Spiders were identified by use of the keys in Kaston (1981), by comparison with
specimens in the Florida State Collection of Arthropods (F.S.C.A.), and verification by
Dr. G. B. Edwards. Hemiptera were identified by Dr. Frank Mead and Coleoptera by
Dr. R. E. Woodruff and Dr. J. Howard Frank. Voucher specimens of all species col-
lected are deposited in the F.S.C.A.

RESULTS

The corn stubble provided two basic hiding places for numerous overwintering spid-
ers and insects. The leaf sheath normally remained attached to the corn stalk, where it
overlayed an indentation along the length of the stalk (Figure 1); this was the only site
provided by the stubble left in the Archer field. The corn stubble that was left at Tall
Timbers also included the shelled cobs with up to 20 enclosing husks. This protected
and insulated site was greatly preferred by the overwintering spiders and insects at
Tall Timbers as shown by their virtual absence from the protected leaf base cavities.
This overwintering site is shown in Fig. 2. At either site, no more than an occasional
specimen was collected from within the corn stalks.
In all, 24 species of spiders and 25 species of predaceous insects were identified. The
spiders collected are listed in Table 1 and the predaceous insects in Table 2.


Leaf Sheath






Corn Stalk










Fig. 1. Cross section of corn stalk and leaf sheath showing the protected cavity
between them.










Florida Entomologist 69(4)


Husks


Corn Cob


















Fig. 2. Cross section of corn cob with enclosing husks showing the protected layers
of air.

DISCUSSION
Overwintering spiders have at least 3 requirements of their quarters. First, the site
must insulate them from the coldest temperatures. Kirchner (1973) studied cold resis-
tance in spiders, finding that many species spending the winter in the open or in hollow
plant stems can survive temperatures as low as -16 to -30C. Kirchner studied species
inhabiting central Europe; the cold tolerance of Florida species has not been investi-
gated.
Second, the spider must avoid desiccation (Edgar and Loenen 1974). Protection from
the sun and wind is provided within plant stems, stalks and bracts. A third requirement
is the need to avoid predation (Danks 1978). Poikilotherms, due to their decreased
metabolic activity at low temperatures, are particularly vulnerable to winter-active
birds and mammals. Gunnarsson (1983) found that overwintering spiders on spruce
branches had much greater survival when foraging birds were excluded. The preference
by overwintering spiders for geometrically complex situations such as birds' nests
(Otzen and Schaefer 1980), Spanish moss (Rosenfeld 1911), and in this case, the numer-
ous enclosing bracts around the corn cob, should be important in providing safety from
homeothermic predators as well as insulation from cold temperatures.
The condition of the stubble left in a corn field is partly determined by the method
used to harvest and by the postharvest treatment of the field. If the corn is chopped
for silage, or if the field is disced and planted to a second crop, virtually no stubble is
left. The combine with a picker head, which was used at the Archer field, leaves just
the corn stalks. On the other hand, the picker sheller, which was used at Tall Timbers,
leaves both the stalks and cobs in the field.


668


December, 1986











Plagens & Whitcomb: Overwintering Sites


TABLE 1. SPIDERS COLLECTED DURING THE
ARCHER (160 PLANTS) AND TALL
FLORIDA, 1982-1984.


WINTER FROM CORN STUBBLE AT
TIMBERS (240 PLANTS) IN NORTH


FAMILY
Genus species Stages' Archer2 Tall Timbers3


DICTYNIDAE
Dictyna sp.
THERIDIIDAE
Achaeranea globosa
Coleosoma acutiventer
Theridion spp.
LINYPHIIDAE
Eperigone near banksi
Ceraticellus similis
MIMETIDAE
Mimetus sp.
OXYOPIDAE
Oxyopes salticus
Oxyopes scalaris
AGELENIDAE
Agelenopsis sp.
HAHNIIDAE
Neoantistea magna
LYCOSIDAE
Pardosa milvina
Pardosa parvula
Pirata sp.
ANYPHAENIDAE
Wulfila saltibunda
CLUBIONIDAE
Castieniara sp.
Chiricanthium inclusum
Clubiona sp.
Trachelus sp.
GNAPHOSIDAE
Cessonia bilineata
Poecilochroa sp.
Zelotes rusticus
THOMISIDAE
Misumenoides formosipes
SALTICIDAE
Metaphidippus galathea


EJ,MJ
MJ,F
LJ,M,F


MJ,LJ
LJ


EJ,MJ,LJ,M,F
EJ,MJ,F
MJ


MJ,LJ


EJ,MJ
MJ,LJ
EG,LJ
MJ,LJ,M,F

MJ


LJ,M


'EJ=Early Juvenile, MJ= Middle Juvenile, LJ Late Juvenile, M=Male, F=Female.
All specimens from Archer were collected from under the leaf sheath.
All specimens from Tall Timbers were collected from between the layered husks around the cobs.


The importance of overwintering sites for predaceous arthropods in agroecosystems
is undeniable. However, the corn residue provides overwintering sites for herbivorous
species as well. We found chinch bugs (Blissus insularis), false chinch bugs (Pachyb-
rachius vinctus), saw tooth grain beetles (Oxyzaephilus surinamensis) and grain
weevils (Sitophilus sp.). In addition, Wright et al. (1983) recorded the southern corn
billbug (Sphenophorus callosus) overwintering in the crowns of corn plants, while in


669











Florida Entomologist 69(4)


TABLE 2. PREDACEOUS INSECTS COLLECTED DURING THE WINTER FROM CORN
STUBBLE AT ARCHER (160 PLANTS) AND TALL TIMBERS (240 PLANTS) IN
NORTHERN FLORIDA.

FAMILY
Genus species Stages' Archer2 Tall Timbers"

FORFICULIDAE
Doru aculeatum M,F 23 35
LABIDURIDAE
Labidura ADULT 1
ANTHOCORIDAE
Cardiastethus assimilis ADULT 1
Lasiochilus sp? LI 3 12
Orius sp. LI 1
LYGAEIDAE
Geocoris uliginosus ADULTS 4
Geocoris punctipes ADULTS 12
MIRIDAE
Spanogonicus sp. LI 1
NABIDAE
Hoplistoscelis deceptivus ADULTS 2 10
REDUVIIDAE
Zelus cervicalis ADULTS 2
CARABIDAE
Calleida decora ADULTS 1
Harpalini spp.
(6 species) ADULTS 14
COCCINELIDAE
Scymnus sp. ADULTS 1 2
STAPHYLINIDAE
Genus spp.
(11 species) ADULTS 80

'LI=Late Instar, M=Males, F-Females
2All specimens from Archer were collected from under the leaf sheath.
'All specimens from Tall Timbers were collected from between the layered husks around the cobs.


more northern latitudes stubble helps to retain a layer of insulating snow that increases
the survival of Lepidoptera pupae (Turnock and Bilodeau 1984).
Two major pests of corn also make use of post harvest corn residue and the remaining
crowns as overwintering sites, namely the European corn borer (Ostrinia nubilalis)
(Brindley and Dicke 1963) and the southwestern cornstalk borer (Diatraea cram-
bidoides) (Metcalf et al. 1962). Burkhardt (1952) also reported fall armyworm (Spodopt-
era fruqiperda) larvae pupating in parts of the corn plant. Thus, the use of corn stubble
in management of predator populations in areas where these insects are important pests
would be unwise.
However, the intensity and diversity of predators in corn residue suggests that
overwintering sites could otherwise be limiting and that providing alternative sites
could increase predator survival. Fye (1985) placed bundles of corrugated fiberboard as
overwintering sites in pear orchards, but did not detect a significant increase in predator
numbers the following spring despite large numbers of predators using the the traps.
This result was attributed to variance in the reservoirs of predators the previous fall
and the need for even higher densities of traps. How overwintering sites might be


670


December, 1986











Plagens & Whitcomb: Overwintering Sites


provided while not benefiting pest species will require intimate knowledge of the over-
wintering biology of both pests and beneficial.

ACKNOWLEDGMENTS

The authors wish to extend sincere thanks to Mr. Jack Simmons who admitted us
to his agricultural fields and to Dr. Edward V. Komerak who granted use of the Tall
Timbers Research Station.
Florida Agricultural Experiment Station Journal Series #7179.

REFERENCES CITED

ARCHER, T. L., E. D. BYNUM, JR., AND A. KNUTSON. 1983. Winter management
of the southwestern cornborer (Lepidoptera: Pyralidae), using several cultural
practices on different dates. J. Econ. Entomol. 76: 872-6.
BRINDLEY, T. A., AND F. F. DICKE. 1963. Significant developments in European
corn borer research. Ann. Rev. Entomol. 8: 155-76.
BURKHARDT, C. C. 1952. Feeding and pupating habits of the fall armyworm in corn.
J. Econ. Entomol. 45: 1035-7.
DANKS, H. V. 1978. Modes of seasonal adaptation in the insects. I. Winter survival.
Canadian Entomol. 110: 1167-205.
EDGAR, W. E., AND M. LOENEN. 1974. Aspects of the overwintering habitat of the
wolf spider Pardosa lugubris. J. Zool. 172: 383-7.
FYE, R. E. (1985) Corrugated fiberboard traps for predators overwintering in pear
orchards. J. Econ. Entomol. 78: 1511-4.
GUNNARSSON, B. 1983. Winter mortality of spruce-living spiders: effect of spider
interactions and bird predation. Oikos 40: 226-33.
JEFFORDS, M. R., G. C. HELM, AND M. KOGAN. 1983. Overwintering behavior and
spring colonization of soybean by the bean leaf beetle (Coleoptera:Chrysomelidae)
in Illinois. Environ. Entomol. 12: 1459-63.
KASTON, B. J. 1981. Spiders of Connecticut. State Geolog. and Natur. Hist. Survey
of Conn. Bull. 70, revised edit. 1020 p.
KIRCHNER, W. 1973. Ecological aspects of cold resistance in spiders, p. 271 to 279
IN Effects of temperature on ectothermic organisms. edit. W. Weiser. New
York 298 p.
METCALF, C. L., W. P. FLINT, AND R. L. METCALFDD. 1962. Destructive and
useful insects, their habits and control. New York 1087 p.
OTZEN, W., AND M. SCHAEFER. 1980. Hibernation of arthropods in bird nests: con-
tribution to winter ecology. Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere 107:
435-48.
ROSENFELD, A. H. 1911. Insects and spiders in Spanish moss. J. Econ. Entomol. 4:
398-409.
SLOSSER, J. E., R. J. FEWIN, J. R. PRICE, L. J. MEINKE, ASND J. R. BRYSON.
1984. Potential of shelter belt management for boll weevil (Coleoptera:Cucur-
lionidae) control in Texas Rolling Plains. J. Econ. Entomol. 77: 377-85.
TURNOCK, W. J., AND R. J. BILODEAU. 1984. Survival of pupae of Mamestra con-
figurata (Lepidoptera:Noctuidae) and 2 of its parasites in untilled and tilled soil.
Canadian Entomol. 116: 257-68.
WRIGHT, R. J., J. W. VANDUYN, AND J. R. BRADLEY, JR. 1983. Seasonal phenology
and biology of the southern corn billbug in eastern North Carolina. J. Georgia
Entomol. Soc. 18: 376-85.











672 Florida Entomologist 69(4) December, 1986


OCCURRENCE OF CULICOIDES MISSISSIPPIENSIS ON
DIFFERENT TYPES OF VEGETATION

THOMAS H. LILLIE', AND DANIEL L. KLINE
Insects Affecting Man and Animals Research Laboratory
U.S.D.A., Agricultural Research Service
Gainesville, Florida 32604

ABSTRACT
An AFS Sweeper was used from 11 February to 8 April 1985 to examine the occurr-
ence of Culicoides mississippiensis Hoffman adults on Spartina aIltert florra Loiseleur,
Ilex vomitoria Aiton, Juniperus silicicola (Small), Juncus roemerianus Scheele, and
an undetermined species of short grass. Spartina alterniflora and I. vomitoria harbored
significantly more specimens than the other plants. The sex ratio was 1:1 on S. alterni-
flora, the dominant plant type in the breeding area. Ilex vomitoria contained a male-
biased sex ratio and significantly more C. mississippiensis adults when in flower than
when not in flower. The association with flowers suggests a response to a visual or
chemical cue.

RESUME

Se us6 un esparavel tipo AFS del 11 de Febrero al 8 de Abril de 1985 para determinar
la frecuencia de adults de Culicoides mississippiensis Hoffman en Spartina alternif-
lora Loiseleur, Ilex vomitoria Aiton, Juniperus silicicola (Small), Juncus roemerianus
Scheele, y una indeterminada especie de hierba corta. Spartina alterniflora y I. vom-
itoria albergaron significativamente mas species que las otras plants. La proporci6n
del sexo fue de 1:1 en S. alterniflora, que es el tipo de plant que predomina en el area
de cruzamiento. Ilex vomitoria contenia una proporci6n de sexo, parcial hacia los machos
y significativamente tenia mas adults de C. mississippiensis cuando estaba florido que
cuando no lo estaba. La asociaci6n con las flores sugiere una reacci6n a un estimulo
visual o quimico.



Some species of bloodsucking insects show a preference for settling at specific sites
between periods of flight. Females may land on a vertebrate host to obtain a bloodmeal,
or rest on man-made structures, plants, or the soil, while they digest a bloodmeal. Also,
both sexes may visit sap flows or flowers to obtain nectar. Knowledge of the association
with a particular host, resting site, or nectar source may be useful when planning a
control program. A repellent can be applied on the host, resting sites can be treated
with a residual insecticide, or extracts from a nectar source can be used as attractants.
Several workers have examined the host range for various species of Culicoides,
but little research has been done regarding resting sites or nectar sources. The latter
are of particular importance because nectar provides energy for sustained flight in
mating swarms (Downes 1969), increases adult longevity (Linley 1966a), and may play
a role in egg maturation (Linley 1966b). The females of some species are not able to
survive to the time of the first oviposition without nectar (Downes 1958).
Neither resting sites nor nectar sources are known for the biting midge, Culicoides
mississippiensis Hoffman (Diptera: Ceratopogonidae), a common pest of man along the

'Present address: Epidemiology Division, U.S. Air Force School of Aerospace Medicine, Brooks Air Force Base,
Texas 78235-5301.











Lillie & Kline: Plant Association of Culicoides 673

Gulf Coast from Texas to southern Florida (Blanton and Wirth 1979). We designed a
study to examine the association between this species and 5 plant species.

MATERIALS AND METHODS

Possible resting sites and nectar sources were determined following preliminary
tests in March and April 1984 near Yankeetown, Levy County, Florida. Adult C. mis-
sissippiensis were found in great numbers, at that time, near flowering shrubs and
breeding sites in the marsh. With this in mind, Juncus roemerianus Scheele, Ilex
vomitoria Aiton, Spartina alterniflora Loiseleur, Juniperus silicicola (Small), and an
undetermined species of short grass were selected. The specific plants ranged in heights
as follows: J. roemerianus, 1-1.5 m; I. vomitoria, 1-2.5 m; S. alterniflora, 0.25-0.75 m;
J. silicicola, 2-3 m; and short grass, 0.1-0.2 m.
We used a portable suction device, an Arbovirus Field Station (AFS) Sweeper
(Meyer et al. 1983), to collect C. mississippiensis adults. Similar devices have been
used in prior studies of biting midges (Bidlingmayer 1961, Tanner and Turner 1975) and
mosquitoes (Nasci 1981). We always began sampling 4-5 hrs after sunrise to avoid the
possible bias of any diel periodicity. The suction device was moved over each type of
plant for 2 min at 3 different sites on 11, 22, and 26 February, 11 and 26 March, and 1
and 8 April 1985.
Since shape and size varied between species of plant selected, the specific pattern
of movement of the sweeper varied also. For the shrubs, J. silicicola and I. vomitoria,
the intake port was moved over the outer canopy and along the branches. For the
grasses, J. roemerianus, S. alterniflora, and the short grass, the device was moved
through the vegetation. For all 5 plant species the sampling device contacted the plant
so as to disturb any resting insects, causing them to take flight and be sucked into the
sweeper. The time of each sample remained constant, but variation in the area sampled
was unavoidable from day to day because the operator determined the pace of the
sweeper around and through the plants. We calculated this variation by measuring the
area the operator was able to cover in 2 min while sampling for C. mississippiensis in
short grass. The average area for 7 samples was 6.7 0.7 m2 (mean standard devia-
tion). Since the sweeper was easy to move through the short grass, the area covered
on other plants was probably less.
One hundred and five samples were obtained in the above manner. We also collected
21 control samples by holding the suction device 45-60 cm away from plants included in
the study. All samples were placed in a styrofoam container with solid CO2 (dry ice) for
transport to the laboratory in Gainesville, where the number of C. mississippiensis
males and females (gravid and non-gravid) was recorded. Data were subjected to a
chi-square/G-statistic with multiple column analysis to assess the significance of differ-
ences we observed.


RESULTS AND DISCUSSION

We collected C. mississippiensis adults on all plants and in the controls. More indi-
viduals were found on S. alterniflora and I. vomitoria than on the other plants (Table
1). The former harbored the most individuals initially, and the sex ratio was approxi-
mately 1:1 for each day we collected specimens (Fig. 1). In contrast, I. vomitoria con-
tained few specimens when we began the study, but the number greatly increased on
the last 2 sampling dates (Fig. 2). Ilex vomitoria had significantly more males (65%)
(p<0.005, X2= 19.15, d.f. = 1) than S. alterniflora (49%) on these 2 dates.
The sex ratio varied significantly (p>0.005, X2=29.08, d.f. = 4) when we combined










Florida Entomologist 69(4)


CULICOIDES MISSISSIPPIENSIS ON
SPARTINA ALTERNIFLORA


I IIFeb 122 Feb 126 Feb II Mar 22 Mar I I Apr 1 8 Apr
SAMPLING DATE (1985)
Fig. 1. Number of Culicoides mississippieCsis adults collected on Spartina alferni-
flora by using a portable suction device.

data for each plant species and compared it statistically. Most ceratopogonid species
have a sex ratio of 1:1 at adult emergence (Kettle 1955). Such a ratio for specimens
collected on S. altermflora should not be unusual (Table 1) because C. mississippiensis
immatures are often found in salt marsh soil where this plant grows (Kline 1986). The
immatures are also associated with soil where J. roemerianus grows; 57% of the speci-
mens found on this plant were female. Samples from J. silicicola, which grows near the
salt marsh in areas not regularly flooded by tides, were composed 57% of females.
The sex ratio was not 1:1 on I. vomitoria (37% female, 63% male). Such a deviation
may occur if there is differential mortality between the sexes (Linley and Mook 1978),
or if the site where specimens are being collected contains an attractant that lures one
sex in greater abundance over the other. Differential mortality may be ruled out in this
case because the ratio remained 1:1 on plants such as S. /'. t '.. t ..., We suspect an
attractant is involved in the relationship between C. mississippiensis and 1. vomitoria.
A sudden increase in the number of specimens (Fig. 2) coincided with the flowering
period for this plant, late March to early May. On 1 April we noticed more midges in
collections obtained on plants in flower than on plants not in flower. Consequently, on
8 April we sampled 5 flowering plants and 3 non-flowering plants to compare the number
of C. mississippiensis associated with each. The data (Table 2) strongly suggest that
the number of specimens is related to the flowering condition. We did not sample


December, 1986


674











Lillie & Kline: Plant Association of Culicoides


CULICOIDES MISSISSIPPIENSIS ON

ILEX VOMITORIA



* Females
SMal32
Males P^


7


0 0 r-^n I1


5

0 A


I Feb 22 Feb 26 Feb| II Mar I 22 Mar I Apr


675


SAMPLING DATE (1985)
Fig. 2. Number of Culicoides mississippiensis adults collected on Ilex comnitoria by
using a portable suction device.

beyond 8 April because I. vomitoria was no longer in flower at the Yankeetown site
soon after that date.
Ilex vomitoria is the most common flowering shrub in the area in late March and
early April. Midges were observed crawling into and out of the flowers and over
branches. Gravid females were found in significantly greater frequency (p<0.005,
X'=44.30, d.f. = 4) on I. vomitoria than on other plants. Since females often do not
survive in the laboratory to the age of their first oviposition without ingesting some


TABLE 1. RESTING SITES OF CULICOIDES MISSISSIPPIENSIS ADULTS AS DETER-
MINED BY PORTABLE SUCTION DEVICE COLLECTIONS.

Number of individuals

Non-gravid Gravid
Material sampled females females Males TOTAL

Spartina alterniflora 206 34 240 480
Ilex vomitoria 147 62 362 571
Juniperuts silicicola 47 0 35 82
Juncus roemerianus 29 1 23 53
Short grass 12 2 16 30
Control1 32 0 0 32
TOTAL 473 99 676 1248

Specimens in flight near plants rather than resting on plants.


40-



C)
I- 30-
-j



o 20-
cr
w


z 10-


326

183~


2










676 Florida Entomologist 69(4) December, 1986


TABLE 2. PREFERENCE OF CITLICOIDES MISSISSIPPIEWNSIS ADULTS FOR FLOWER-
ING AND NON-FLOWERING ILEX VOMITORIA AS DETERMINED BY
PORTABLE SUCTION DEVICE COLLECTIONS.

Number of individuals
Non-gravid Gravid
Ilexr vomitoria females females Males TOTAL

Flowering
A 117 62 326 505
B 82 43 225 350
C 57 20 83 160
D 10 12 28 50
E 6 4 27 37
CONTROL 14 0 0 14

Non-flowering
F 2 0 2 4
G 3 0 0 3
H 0 0 0 0
CONTROL 0 0 0 0



type of sugar, they were probably on the plants to obtain nectar. Males also ingest
sugar in the laboratory and require similar materials in their natural environment.
Nectar provides energy for metabolic activities, and may play a role in egg maturation.
Other plants evaluated in this study did not produce nectar in flowers like I. vom-
itoria, or they did not grow in areas of high larval density like S. alterniflora. Juniperus
silicicola is a gymnosperm and the others (J. roemerianus and shortgrass) are
monocots. Collections for these 3 species and the controls accounted for only 16% of the
specimens recorded. Of these, only 3 were gravid females. The control samples did not
contain any males or gravid females.
The purpose of the controls was to determine the number of specimens in flight near
a plant as opposed to those resting on the plant. We did not contact the plant while
collecting these samples. It is possible that some midges were disturbed when the AFS
sweeper was 45-60 cm away but this is unlikely because only non-gravid females were
obtained, even when sampling near I. vomitoria that harbored many males and gravid
females (Table 2). The preponderance of non-gravids could have been due to the
operator of the suction device attracting host-seeking females. Such attraction is an
extraneous factor that might lead to false conclusions. For example, the short grass
may be a resting site for females if you consider only data from sweeps through the
grass. However, it is actually a poor resting site if it is assumed that many of the
females were attracted to the operator from other sources.
Further studies are planned to examine the relationship between C. mississippiensis
and flowering plants in the Yankeetown area. Some type of visual or chemical cue is
probably involved. Extracts from I. vomitoria will be evaluated as a midge attractant.
Other efforts will be directed toward the search for alternative sources of nectar. Since
the midges are active in spring, fall, and winter (Kline 1986; Lillie 1985), and I. vom-
itoria flowers for only a short time, other sources of nectar are probably important.
Vaccinium arboreum Marsh is a good candidate plant; it occurs in the area and it is a
source of nectar for bees.











Lillie & Kline: Plant Association of Culicoides


ACKNOWLEDGMENT

We are grateful to Dr. J. R. Linley (Florida Medical Entomology Laboratory, Vero
Beach, Florida) and to Dr. Robert Vander Meer (of this laboratory) for their critical
reviews of the manuscript and for helpful suggestions.

REFERENCES CITED

BIDLINGMAYER, W. L. 1961. Field activity studies of adult Culicoidesfurens. Ann.
Ent. Soc. America 54: 149-56
BLANTON, F. S., AND W. W. WIRTH. 1979. The sand flies (Culicoides) of Florida
(Diptera: Ceratopogonidae). Arthropods Florida Neighboring Land Areas 10/
1-204.
DOWNES, J. A. 1958. The feeding habits of biting flies and their significance in class-
ification. Ann. Rev. Entomol. 3: 249-266.
DOWNES, J. A. 1969. The swarming and mating flight of Diptera. Ann. Rev. Entomol.
14: 271-98.
KETTLE, D. S. 1955. Sex ratios among British Culicoides Proc. Roy. Ent. Soc. Lond.
30: 70-72.
KLINE, D. L. 1986. Seasonal abundance of adult Culicoides spp. (Diptera:
Ceratopogonidae) in a salt marsh in Florida, USA. J. Med. Entomol. 23: 16-22.
LILLIE, T. H. 1985. Diel and seasonal activities of Culicoides spp. near Yankeetown,
Florida. Ph.D. Dissertation, Univ. of Florida, Gainesville. 145 pp.
LINLEY, J. R. 1966a. The ovarian cycle in Culicoides barbosai Wirth and Blanton and
C. furens (Poey) (Diptera: Ceratopogonidae). Bull. Entomol. Res. 57: 1-17.
LINLEY, J. R. 1966b. Effects of supplementary carbohydrate feeding on fecundity
and life-length in Leptoconops becquaerti (Kieff). Bull. Entomol. Res. 57: 19-22.
LINLEY, J. R., AND M. S. MOOK. 1978. Seasonal change in the pupal sex ratio in a
population of Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Proc. Flor-
ida Anti-Mosq. Assoc. 49: 6-11.
MEYER, R. P., W. K. REISEN, B. R. HILL, AND V. M. MARTINEZ. 1983. The AFS
Sweeper, a battery-powered backpack mechanical aspirator for collecting adult
mosquitoes. Mosq. News 43: 346-50.
NASCI, R. S. 1981. A lightweight battery-powered aspirator for collecting resting
mosquitoes in the field. Mosq. News 41: 808-11.
TANNER, G. D., AND E. C. TURNER, JR. 1975. Seasonal abundance of Culicoides
spp. as determined by three trapping methods. J. Med. Ent. 12: 87-91.


677











Florida Entomologist 69(4)


MONITORING THE FLIGHTS OF FIELD CRICKETS
(GRYLLUS SPP.) AND A TACHINID FLY
(EUPHASIOPTERYX OCHRACEA) IN NORTH FLORIDA

THOMAS J. WALKER
Department of Entomology and Nematology
University of Florida
Gainesville, Florida 32611

ABSTRACT

Traps broadcasting synthetic calls sampled flights of Gryllus firmus and G. rubens
for 3 years in a pasture near Gainesville, Florida. Both species flew at all times of year
with a major peak in the fall. Average annual catch of G. firmus for the trap broadcast-
ing firms call was 163 (39% male), with mean monthly catches varying from <1 in
February to 63 in October. The trap broadcasting rubens call captured an annual aver-
age of 8,209 G. rubens (44% male), with mean monthly catches ranging from 29 in
January to 1,956 in September. During July and August, G. rubens flew throughout
the night. Gravid females of Euphasiopteryx ochracea, a parasitoid of Gryllus, were
attracted to the rubens trap but not to the firmus trap. In one year 948 were caught,
with lows of 0 in January, April, and June; and a high of 789 in October.

RESUME

Se muestrearon vuelos de Gryllus firmus y de G. rubens por 3 afos en un past
cerca de Gainesville, Florida, con trampas diseminadoras de llamadas sint6ticas. Ambas
species volaron todo el ano, siendo su apogeo en el otono. El promedio annual de capture
de G. firmus por trampas diseminando llamadas de firmus fue de 163 (39% machos),
con promedios mensuales de capture variando entire menos de 1 en Febrero a 63 en
Octubre. La trampa diseminando llamadas de rubens capture un promedio annual de 8,
209 G. rubens (44% machos), con captures de un promedio mensual entire 29 en Enero
a 1,956 en Septiembre. G. rubens vo16 a trav6s de la noche de Julio a Agosto. Hembras
prefiadas de Euphasiopteryx ochraea que es un parasitoide de Gryllus, fueron atraidos
a la trampa de rubens pero no a la de firmus. Se atraparon 948 en un afo, con nfmeros
de 0 en Enero, Abril, y Junio, y un maximo de 789 en Octubre.



The most common southeastern field crickets, Gryllus firmus and G. rubens, are
dimorphic in length of the metathoracic wings (Walker and Sivinski 1986). Short-winged
morphs are flightless. Long-winged morphs include strong fliers, although some indi-
viduals may not fly. Flying field crickets, like flying mole crickets, often land at broad-
casts of the conspecific calling song (Ulagaraj and Walker 1973, Cade 1979a, Walker
1982, Forrest 1983). To study field cricket flights in north Florida, I operated traps that
emitted the synthesized call of either G. rubens or G. firmus. Larvipositing females of
Euphasiopteryx ochracea, a tachinid parasitoid of crickets, were attracted to the call
of G. rubens.

METHODS

Three traps were installed 30 m apart, in an equilateral triangle, near Gainesville,
Fla., in a 5-ha bermuda- and bahiagrass pasture surrounded by woods of slash pine and


December, 1986











Walker: Monitoring Field Crickets 679


mesic hardwoods (NW corner of sec. 31, tp. R19E, T9S). Each trap consisted of a
1.42-m diameter sheet metal funnel emptying into a screen-bottomed, 2-liter plastic jar
(Fig. 1). Centered above the funnel and protected from rain by a plastic bag was an
electronic sound synthesizer, similar to one described by Walker (1982) but driven by
a programmable microprocessor (Intel 8748). (The synthesizer was made and program-
med at Oldacre Electronics, P.O. Box 12951, Gainesville, FL 32604.) Synthesizers used
in this study broadcast calls of either rubens (4763 Hz carrier frequency, 50.0 pulses
per second, 50% duty cycle; continuous trill) or firmus (3980 Hz, 16.9 p/s, 50% duty
cycle; 4 pulses per chirp, 2 chirps per second) (Fig. 2). Above each synthesizer was a
30 x 43 cm (dia x h) cylindrical electrocution grid of the type used by Mitchell et al.
(1972). The grid was attached to a plywood ring (inner and outer diameters 25 and 60 cm)
that had a 3.5 cm high sheet-metal rim around its outer circumference. The ring served
as a platform to catch fragile, electrocuted insects that might otherwise fall into the
funnel and be destroyed by field crickets trapped in the container below.
One trap was equipped with a rubens synthesizer, one with a firmus synthesizer,
and one (the control) with a silent, imitation synthesizer. Positions for the 3 treatments
were not changed during the 3 years of the study, 1-1-1983 to 31-XII-1985. After 9
months the control was discontinued since it had captured no Gryllus sp. Power supplies
for the electrocution grids failed during the 2nd year and were not replaced.
A time clock turned on the power to the 3 traps each evening shortly before sunset
and off each morning shortly after sunrise. Output of the rubens synthesizer was set at
106 dB, 15 cm above the speaker (through the plastic bag), using a Bruel & Kjaer model
2219 sound level meter. The meter was inadequate to set the level of thefirmus synthe-
sizer because the chirps were too brief. A Tektronix model 214 portable oscilloscope


























Fig. 1. Sound trap, including electrocution grid, plywood ring, sound synthesizer
(protective plastic bag just visible through hole in ring), sheet-metal funnel, and catch-
ing jar. Funnel, ring, and synthesizer are supported by a steel yoke attached at either
end to posts 1.3 m tall.












Florida Entomologist 69(4)


A. Gryllus firms


680


100-
g-
90-

80-

70-
0

L
50-
-o








10'

0



4000


3500


3000


a 2500
0

S2000

E
-o


1500
Z


1983
1984
1 1985


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec


B. Gryllus rubens


--IIU--.-.~


Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec


Fig. 2. Numbers of Gryllus trapped monthly, 1983-85. A. G. firms caught in a trap
broadcasting syntheticfirmus calling song. (Oscillogram is 2 sec of the broadcast signal.)
B. Numbers of G. rubens caught in a trap broadcasting synthetic rubens calling song.
(Oscillogram is 2 sec of the broadcast signal.)


December, 1986


-tA Will -i1-- Ill











Walker: Monitoring Field Crickets


was used to adjust synthesized firmus pulses to agree in amplitude with those of rubens
at 106 dB.
The traps were serviced each morning by counting and removing insects on the
plywood platform and by substituting a fresh jar below. Jars were sometimes placed in
a freezer to kill the contents prior to identification and counting. No trapped insects
were released in the vicinity of the traps.
During much of July and August 1983 the rubens trap was serviced at 2350-2400 h
(EDT) as well as the following morning. On the nights of 5 and 6-VIII-1983 it was
serviced at the end of each quarter of the period between sunset (2020 h) and sunrise-
viz. at 2258, 0136, 0414, and 0652 h.

RESULTS AND DISCUSSION

CONTROL TRAP

During 1-1-1983 to 30-IX-1983 the control trap caught three female mole crickets
(Scapteriscus vicinus) but no other crickets and no Euphasiopteryx.

FIRMS TRAP

Gryllus firmus flew during every month with peak catches in September and Oc-
tober (Fig. 2A). Flights were never large. The greatest 1-night catch was 17 (30 Sep-
tember 1983); catches of 5 or more occurred in July (n= 1), September (3), October (14),
and November (1). Annual catches were 225, 144, and 120. Overall sex ratio was 39%
male, with no evident seasonal trend.
Only 14 crickets other than G. firms were caught in the firmus trap: 3 G. rubens
8 Scapteriscus vicinus, and 3 S. acletus. One of the S. vicinus was a last instar juvenile,
surely flightless and incapable of climbing into the trap. (Perhaps a bird dropped it.)
On 15 occasions a short-winged firmus was captured. These probably made it into the
trap unassisted, since they are capable of climbing wooden posts, and males occasionally
call from perches well above the ground. Short-winged firmus were excluded from the
counts but their presence raises the possibility that some long-winged firmus climbed
rather than flew into the trap. However, all long-winged firmus are included in the
data reported here (e.g. Fig. 2A).

RUBENS TRAP

Gryllus rubens flew during every month, with peak catches in August (1983), Sep-
tember (1984), or November (1985) (Fig. 2B). Flights were often large, the biggest
one-night catches being 572 (30-IX-1983) and 480 (27-XI-1985). Flights resulting in
catches of 100 or more occurred during 6 months: March (n= August (10), September
(16), October (16), November (15), December (3). Annual catches totaled 7984, 4894, and
11,750. Overall sex ratio was 44% male with monthly ratios from November to March
being 49% or more and those from April to September being 43% or less. Flights
seemed to occur whenever weather was favorable, i.e., nearly nightly from mid-April
through early November (Fig. 3). Only during 3 January to 18 February, the coldest
time of year, did G. rubens not fly in this study. The seasonal distribution of catches of
20 or more was bimodal (Fig. 3). The peak of large catches in April-May probably
resulted from maturation of overwintering juveniles and the peak in August from mat-
uration of progeny of the earlier peak.
Nightly timing of rubens flights differed greatly from flights of Scapteriscus vicinus












682 Florida Entomologist 69(4) December, 1986

1.0
Gryllus rubens [--

1-19
8 20 or more

.7

.6

z .5

0


.2
t 2 -








Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Fig. 3. Frequency of nights with catches of 0, 1-19, or 20 or more Gryllus rubens in
the rubens trap.

and S. acletus the only other local crickets known to have frequent mass flights.
Whereas S. vicinus and S. acletus restrict their flights to the first 20 to 90 minutes
after sunset (Forrest 1983), G. rubens apparently flies at all times of night with late-
night numbers equal to or greater than early-night numbers (temperature permitting).
During the 39 nights during July and August 1983 that traps were emptied at midnight
as well as the following morning, 29% of the males (244 of 834) and 23% of the females
(273 of 1186) were captured prior to midnight (Table 1). At that season, sunset to
midnight is 34 to 36% of the period from sunset to sunrise. Chi square tests of the
hypothesis that 35% of captures were prior to midnight and 65% were after midnight
were made separately for males and females and for each of the 3 periods because
preliminary chi square tests showed the data to be heterogeneous (P<0.05). In 3 of the
6 tests the distribution of captures before and after midnight deviated significantly from
35:65 (Table 1), and in every case fewer that 35% were captured prior to midnight.
During the 2 nights that crickets were collected after each quarter between sunset and
sunrise, totals for the quarters were 43, 62, 58, 54. The hypothesis that equal numbers
flew during each quarter was not rejected (chi square; 0.5 >P>0.1).
During the 1st and 3rd, but not the 2nd, of 3 approximately biweekly periods (Table
1), a significantly higher proportion of males than of females was captured prior to
midnight. Why sex ratio changes with season (see above) and sometimes with time of
night is not known.
Cade (1979b) made the only other study of flying times of field crickets. He observed
Gryllus integer (the closest relative of G. rubens) flying to lights at Austin, Texas,
during August 1974. He reported that the greatest numbers flew between 1 and 2 hours
after sunset and that flying had ceased by 4 hours after sunset. On 1 occasion he
continued observations until sunrise and saw no late-flying crickets. Since the species,
season, and latitude were similar in our 2 studies, the contrasting results are unex-
pected. Both species call all night (Cade 1979a, Walker unpublished), and Cade (1979a)











Walker: Monitoring Field Crickets


TABLE 1. G. RIUBENS CAUGHT BEFORE AND AFTER MIDNIGHT DURING 39 NIGHTS.

<2400 h >0000 h % <2400
Date
(1983) M F %M M F %M M F

3-16 July' 75 76 50 160 251 39 328 23b
17-31 July'' 84 96 47 178 197 47 32a 338
2-4, 16-22 Augc 85 101 46 252 465 35 25b 18b
Total 244 273 47 590 913 39 29 23

'Proportion captured before midnight not significantly different from 35'7. (Chi square; P>0.25.)
'Proportion captured before midnight significantly different from 35%. (Chi square; P<0.001.)
'Sex ratio significantly different for periods before and after midnight. (Chi square; P<0.05.)
'Sex ratio not significantly different for periods before and after midnight (Chi square; P>0.90.)

showed that G. integer males and females were attracted to a loudspeaker on the ground
at all hours of night. He also observed their making short flights of a few meters as
they approached the speaker (personal communication). Thus G. integer may fly to an
area only in early evening but may walk or make short flights to a source of calling song
at any time of night. I cannot refute the hypothesis that G. ruben has the behavior that
Cade postulates for G. integer. I observed males and females flying about the trap at
midnight but did not determine whether they were just arriving or had been in the area
since early evening. It is unlikely that late-trapped G. rubens climbed into the trap (as
proposed for short-winged G. firmss. Short-winged G. rubens were locally common
but were never trapped.
Individuals of 11 other sound-signalling insect species were captured in the rubens
trap. The 3 caught in significant numbers were Scapteriscus vicinus (n=787), S. acletus
(34), and Oecanthus celerinictus (14). The calling song of each of these species is a
continuous trill with a lower carrier frequency than rubens. A standard sound trapping
station for S. vicinus and S. acletus (Walker 1982) was operated in the same pasture
30 m from the rubens trap. During the study period this station caught 20,410 vicinus
and 8,765 acletus. Other species caught more than once were Oecanthus niveus (7),
Neonemobius cubensis (2), Tibicen resonans (2), and Diceroprocta olympusa (2). One
(long-winged) Gryllus firms and 1 Gryllus sp. juvenile were caught.
During 1983, gravid females of Euphasiopteryx ochracea were attracted to the ru-
bens trap every month except January, April, and June. However, only 14 flies were
caught during the first 2/3 of the year, and 934 were caught during the final 3rd (Fig.
4). Of the latter, 789 were caught in October. Three or more flies were caught nightly
29 September to 8 November and 10 or more were caught on 29 of those 41 nights.
More than 50 were caught on the nights of 10, 12, 15, 16, and 23 October.
Cade (1975, 1979a, 1981) showed that E. ochracea was an important parasitoid of
Gryllus integer at Austin, Texas, and that larvipositing females were attracted in large
numbers to its host's calling song-which resembles that of rubens but has a faster pulse
rate. He did not report the seasonal distribution of fly activity. Mangold (1978) collected
E. ochracea females attracted to the calling song of Scapteriscus acletus during a 14-
month study at Gainesville, Florida. His maximum catch was fewer than 15, but, as in
this study, most flies were caught September to November. The song of S. acletus has
a carrier frequency of ca. 3 kHz, much lower than that of G. rubens. Although S. acletus
can be a host (Mangold 1978), it is seldom if ever naturally parasitized by E. ochracea.











Florida Entomologist 69(4)


684

300


Euphasiopteryx ochracea


250+


-mE.-..-


Jan Feb Mar Apr May June July


Aug Sep Oct Nov Dec


Fig. 4. Numbers of Euphasiopteryx ochracea caught weekly, 1983, at a trap broad-
casting synthetic rubens calling song. (Oscillogram is 2 sec of the broadcast signal.)


OVERVIEW

Except for continuously cold periods, the 2 species of Gryllus flew at all times of
year, but in much larger numbers in summer and fall. Peak flights appear to correspond
to times when large numbers of adults have recently matured. E. ochracea had large
flights in fall and was nearly absent otherwise. Its natural hosts at Gainesville and its
seasonal life cycle are unknown.

ACKNOWLEDGEMENTS

I am grateful to J. C. Webb for help with the electrocution grids, to Todd Pickard
and Sue Wineriter for help with trap emptying and cricket counting, and to T. G.
Forrest, W. H. Cade, and J. E. Lloyd for criticizing the manuscript. This research was
supported by NSF Grant BNS 81-03554. Florida Agricultural Experiment Station Jour-
nal Series No. 7278.

REFERENCES CITED

CADE, W. 1975. Acoustically orienting parasitoids: fly phonotaxis to cricket song.
Science 190: 1312-1313.
CADE, W. H. 1979a. The evolution of alternative male reproductive strategies in field
crickets. Pages 343-379 in M. S. Blum and N. A. Blum, eds., Sexual selection
and reproductive competition in insects. Academic Press, New York.
CADE, W. H. 1979b. Field cricket dispersal flights measured by crickets landing at
lights. Texas J. Sci. 31: 125-130.
CADE, W. H. 1981. Field cricket spacing, and the phonotaxis of crickets and parasitoid
flies to clumped and isolated cricket songs. Z. Tierpsychol. 55: 365-375.


December, 1986



















. I1 II*11


150+


50+











Walker: Monitoring Field Crickets


MANGOLD, J. R. 1978. Attraction of Euphasiopteryx ochracea, Corethrella sp. and
gryllids to broadcast songs of the southern mole cricket. Florida Entomol. 61:
57-61.
MITCHELL, E. R., J. C. WEBB, A. H. BAUMHOVER, R. W. HINES, J. W. STANLEY,
R. G. ENDRIS, D. A. LINDQUIST, AND S. MASUDA. 1972. Evaluation of cylin-
drical electric grids as pheromone traps for loopers and tobacco hornworms.
Envir. Entomol. 1: 365-368.
ULAGARAJ, S. M., and T. J. WALKER. 1973. Phonotaxis of crickets in flight: attrac-
tion of male and female crickets to male calling songs. Science 182: 1278-1279.
WALKER, T. J. 1982. Sound traps for sampling mole cricket flights (Orthoptera: Gryl-
lotalpidae: Scapteriscus). Florida Entomol. 65: 105-110.
WALKER, T. J., AND J. M. SIVINSKI. 1986. Wing dimorphism in field crickets (Or-
thoptera: Gryllidae: Gryllus). Ann. Entomol. Soc. America 79: 84-90.





EFFICACY OF BACILLUS SPHAERICUS NEIDE AGAINST
LARVAL MOSQUITOES (DIPTERA: CULICIDAE) AND
MIDGES (DIPTERA: CHIRONOMIDAE) IN THE LABORATORY

ARSHAD ALI
University of Florida, IFAS, Central Florida Research and Education Center,
P. O. Box 909, Sanford, FL 32771
and
JAI K. NAYAR
University of Florida, IFAS
Florida Medical Entomology Laboratory, 200 9th Street, S.E.
Vero Beach, FL 32962

ABSTRACT

The primary powders of strains 1593 (IF-119) and 2362 (IF-118 and ABG-6184) of
Bacillus sphaericus Neide were tested as larvicides of laboratory-reared mosquitoes
and field-collected midges, and were compared with the activity of strain 1593-4 (RB-
80), the international standard of B. sphaericus. Two mosquito species, Aedes aegypti
(Linn.) and Ae. taeniorhynchus (Wiedemann), and two midge species, Chironomus
crassocaidatis Malloch and Glyptotendipes paripes Edwards were insensitive to B.
sphaericus (LC5o > 50 ppm). Among the five susceptible mosquito species, Culex quin-
quefasciatus Say, Cx. nigripalpus Theobald, Anopheles albimanus Wiedemann, and
An. quadrimaculatus Say, were most susceptible to ABG-6184 (strain 2362) with LCgo
values of 0.0044, 0.0067, 0.54, and 7.35 ppm, respectively. Wyeomyia mitchellii
(Theobald) was almost equally susceptible to ABG-6184 (LC9o = 0.276 ppm) and strain
1593 (IF-119) (LCg = 0.261 ppm).

RESUME

Los polvos primaries de las razas 1593 (IF-119) y de 2362 (IF-118 y ABG-614) de
Bacillus sphaericus Neide, fueron probados como larvicidas de larvas de mosquitos
criadas en el laboratorio y de larvas de moscas de agua colectadas del campo, y fueron
comparadas con la actividad de la raza 1593-4 (RD-80) que es el patron international de
B. shaericus. Dos species de mosquitos, Aedes aegypti (Linn.) y Ae. taeniorhynchus


685











Florida Entomologist 69(4)


(Wiedemann), y dos moscas de agua, Chironom us crassicaudatus Malloch y Gl/ptltcn-
dipes paripes Edwards, fueron insensitivas a B. sphaericus (LCso 50 ppm). Entre las
cinco species de mosquitos susceptibles, Culex quinquefasciatus Say, Cx. nigripalpus
Theobald, Anopheles albimanus Wiedemann, y An. quadrimaculatus Say, fueron las
mas susceptibles a ABG-6184 (raza 2362) con valores de LCo9 de 0.0044, 0.0067, 0.54,
y 7.35 ppm respectivamente. WyUm'nifii mitchellii (Theobald) fue casi igualmente sus-
ceptible a ABG-6184 (LC9o=0.276 ppm) y a la raza 1593 (IF-119) (LC0o=0.261 ppm).



The microbial agent, Bacillus sphaericus Neide has been known for its mosquito
larvicidal activity for many years (Kellen et al. 1965). However, the isolation of more
potent strains of this mosquito larval pathogen in recent years has drawn the attention
of a large number of researchers, resulting in many articles on the laboratory and field
efficacy of several potent strains of this bacterium (Davidson et al. 1981, Lacey and
Singer 1982, Lacey et al. 1984, Mulla et al. 1984a,b, Mulligan et al. 1980, and others).
Among the different strains of B. sphaericus tested against mosquito larvae, strains
1593 and 2362 have shown one of the highest levels of biological activity against several
species (Anonymous 1985). The origin, bacteriophage type and serotype of B.
sphaericus strains 1593 and 2362, and other mosquito-pathogenic B. sphaericus strains
are summarized in Davidson (1984). At present, there are several experimental primary
powders of strains 1593 and 2362 available from government and university laboratories
and industry for interested mosquito control researchers to conduct efficacy tests.
Reported here is laboratory response of larvae of seven mosquito species and two
chironomid midge species exposed to B. sphaericus strains 1593 and 2362 (primary
powders) supplied by H. T. Dulmage, USDA, Brownsville, Texas. Larvae of these
mosquito and midge species were also simultaneously exposed to the B. sphaericus
international standard (RB-80) received from H. de Barjac, Institut Pasteur, Paris,
France, and to an industrially-produced primary powder (ABG-6184) provided by Ab-
bott Laboratories, N. Chicago, Illinois.


MATERIAL AND METHODS

The mosquito species tested were Aedes aegypti (Linn.), Ae. taeniorhynchus
(Wiedemann), Anopheles albimanus Wiedemann, An. quadrimaculatus Say, Culex
nigripalpus Theobald, Cx. quinquefasciatus Say, and Wyeomyia mitchellii (Theobald).
These species were maintained at the Florida Medical Entomology Laboratory at Vero
Beach, Florida. For midge bioassays, field-collected 3rd and 4th instars of C I.. '"...i,
crassicaudatus Malloch and Glyptotendipes paripes Edwards were used. Larvae of the
former species were collected from Lake Monroe, as described in Ali and Baggs (1982),
while G. paripes was obtained from Lake Jessup located at 5-6 km distance from Lake
Monroe, Seminole County, Florida.
Methods utilized for mosquito bioassays were generally the same as described by
Mulla et al. (1982). Twenty larvae (3rd and 4th instars) of a mosquito or midge species
were placed in 120-ml disposable paper cups containing 100 ml tap water. Distilled
water (pH 6.9 0.2) was used for rearing and testing Wy. mitchellii because of the
possibility of its larval mortality in tap water (Nayar 1982). In the midge studies, 5 g
of sterilized sand per cup was added for the larval acclimatization and to prevent any
cannibalism. For treatments, each strain or preparation of B. sphaericus was suspended
in tap water by using a magnetic stirrer to make its 1% stock suspension (w/v) and, as
needed, serial dilutions were made to obtain the appropriate range of concentrations
for testing.


686


December, 1986











Ali & Nayar: Larvicical Activity of B. sphaericus


Each strain or preparation was tested on at least three different occasions. Each
time, 5-6 concentrations (in suspension) of a strain or preparation were applied to each
of three cups (replicates) receiving a concentration while three cups were left untreated
as controls. The stock suspensions and serial dilutions were freshly prepared on each
occasion. The treated and control cups containing mosquitoes or midges were main-
tained under 14-h photoperiod and 27 t 1C room temperature. After a 48-h exposure
period to the pathogen, larval mortality was assessed. The mortality in the treated cups
in a test was adjusted against any mortality in the controls (Abbott 1925). The corrected
mortality was subjected to log probit regression analysis.

RESULTS AND DISCUSSION

The levels of toxicity of the test strains or preparations of B. sphaericus to five
mosquito species in the laboratory are presented in Table 1. The data concerning the
Aedes and the midge species are not included in the table because their LCo5 values
exceeded 50 ppm, thus showing insensitivity to the test strains of B. sphaericus. Among
the five susceptible species of mosquitoes, Cx. quinquefasciatus was the most suscepti-
ble. The Abbott's (2362) preparation (ABG-6184) was slightly superior in activity against
this species than the Dulmage preparations of strains 2362 (IF-118) and 1593 (IF-119)
as indicated by the 0.0044, 0.0052, and 0.0064 ppm LCgo values, respectively.
All strains and preparations of B. sphaericus showed a high level of biological activ-
ity against Cx. nigripalpus. ABG-6184 (2362) was nearly 2X more active against Cx.
nigripalpus than the international standard (1593-4, RB-80). Strains 1593 (IF-119),
2362 (IF-118), and the international standard showed a similar level of activity against
Cx. nigripalpus (Table 1).
Anopheles albimanus was most susceptible to ABG-6184 (LC9o = 0.54 ppm) followed
by 1593 (IF-119) (LCgo = 0.91 ppm) and 2362 (IF-118) (LCgo = 1.22 ppm). By contrast,
An. quadrimaculatus was several times less susceptible to all the test strains or prep-
arations (Table 1). Against An. quadrimaculatus, strain 2362 (IF-118) was the least
active (LCYo = 27.2 ppm).
Bacillus sphaericus showed good activity against Wy. mitchellii with LC9g values
ranging from 0.261-0.818 ppm. Strains 1593 (IF-119) and Abbott's (2362) preparation
showed superior activity against Wy. mitchellii than the international standard and the
strain 2362 (IF-118). In general, the primary powder, ABG-6184 (strain 2362), proved
the most toxic preparation against the mosquito larvae.
Data on the laboratory activity of several toxic strains and preparations of B.
sphaericus against a number of mosquito species in the genera Aedes, Anopheles,
Culex, Culiseta, Mansonia, and Psorophora are previously available (Anonymous 1985,
Mulla et al. 1984a,b, 1985). The present study adds Wy. mitchellii to the list of suscep-
tible species of mosquitoes to B. sphaericus. This study also reveals that C. cras-
sicaudatus and G. paripes midge larvae were insensitive when exposed for 48 h to the
test strains and preparations of B. sphaericus. The microbial agent was also ineffective
against Ae. aegypti and Ae. taeniorhynchus at exposure rates exceeding 50 ppm. The
insensitivity of Ae. aegypti to the available most toxic strains of B. sphaericus is already
documented (Anonymous 1985, Mulla et al. 1984b).
Among the susceptible species of mosquitoes, Cx. nigripalpus and Cx. quinquefas-
ciatus were highly susceptible; the LCgo values of Cx. quinquefasciatus achieved by
different strains and preparations of B. sphaericus used in the present study were 6-20
times lower than those reported by Mulla et al. (1984b) for B. sphaericus strains 2362
(IF-97 AP) and 1593 (IF-94) against Cx. quinquefasciatus. In contrast, lower LC90
values of An. quadrimaculatus were observed by Mulla et al. (1984b) than those


687











Florida Entomologist 69(4)


TABLE 1. SUSCEPTIBILITY OF LABORATORY REARED LARVAE (3RD AND 4TH IN-
STARS) OF FIVE SPECIES OF MOSQUITOESa TO VARIOUS STRAINS AND
PREPARATIONS OF BACILLUS SPHAERICUS IN THE LABORATORY.

48-h lethal concentration (ppm)
Strain and/or
preparation LCo5 95% CL LC90 95% CL

Anopheles albimanus
1593b 0.31 0.26-0.35 0.91 0.81- 0.98
2362e 0.50 0.44-0.57 1.22 0.99- 1.37
Abbott (2362)"1 0.19 0.16 0.22 0.54 0.48- 0.61
Anopheles quadrimaculatus
1593 2.88 2.50-3.40 9.58 8.16-11.20
2362 8.20 7.14-9.74 27.20 24.20-30.64
Standard (1593-4)e 2.42 2.25-2.60 7.86 7.06- 8.69
Abbott (2362) 2.29 2.09-2.50 7.35 6.86- 7.82
Culex nigripalpus
1593 0.0052 0.0049- 0.0056 0.016 0.012 -0.019
2362 0.0048 0.0046- 0.0050 0.014 0.011 -0.018
Standard (1593-4) 0.0071 0.0066 0.0077 0.012 0.011 0.013
Abbott (2362) 0.0026 0.0021 0.0029 0.0067 0.0061 0.0073
Culex quinquefasciatus
1593 0.0023 0.0020 0.0025 0.0064 0.0057 0.0073
2362 0.0016 0.0014 0.0019 0.0052 0.0042- 0.0064
Abbott (2362) 0.0017 0.0015 0.0019 0.0044 0.0040- 0.0049
Wyeomyia mitchellii
1593 0.052 0.045 0.060 0.261 0.245- 0.278
2362 0.269 0.242-0.294 0.818 0.719-0.920
Standard (1593-4) 0.129 0.116-0.145 0.442 0.437 0.448
Abbott (2362) 0.069 0.062 0.076 0.276 0.266 0.290

aMaintained at the Florida Medical Entomology Laboratory at Vero Beach, FL.
4IF 119, spray dried.
IF 118, spray dried.
'A8G-6184, lot 76-828-BD, spray dried.
'International Standard (RB-80), lyophilized.

achieved for the same species in the present study. Variations in activity levels in
different studies could be attributed to several factors such as larval strain, larval age
instarr) and vigor at the time of testing, and procedures used in bioassays. The type of
formulation, percent of active ingredient in a formulation, and the production of the
bacterium under different conditions would also attribute to the toxicity potential of
different preparations of B. sphaericus.
This study confirms the usefulness of B. sphaericus in the biological control of mos-
quitoes. The host range of available isolates of this microbial mosquito larvicide reported
in several laboratory and field studies (e.g., Anonymous 1985, Mulla et al. 1984a,b,
1985, Ramoska et al. 1978, and others) covers species of Culex, Culiseta, Anopheles,
Mansonia, Psorophora, Wyeomyia (present study), and even some aedine species, such
as Ae. melanimon, Ae. nigromaculis, and Ae. triseriatus. The mosquito host range of
B. sphaericus is considered narrower than that of Bacillus thuringiensis var. israelen-
sis (B.t.i.). However, in general, B. sphaericus is more toxic to many susceptible species
of mosquitoes than B.t.i. and also has the advantage of longer persistence in the treated
habitats (Anonymous 1985, Des Rochers and Garcia 1984) although the persistence
phenomenon of B. sphaericus is not as evident in highly polluted environments (Mulla


688


December, 1986











Ali & Nayar: Larvicical Atiritil of B. sphaericus 689


et al. 1984a). Further research on isolation of more toxic strains, development of new
and more effective formulations, and genetic manipulation to enhance the toxicity of
this microbial insecticide (Lacey 1984, and others) may broaden the host range and
ultimate usefulness of this microbial agent in vector control.

ACKNOWLEDGMENTS

This is Florida Agricultural Experiment Stations Journal Series No. 7224. Gratitude
is expressed to Drs. H. de Barjac (Institut Pasteur, Paris) and H. T. Dulmage (USDA,
Brownsville, Texas), and Abbott Laboratories (N. Chicago, Illinois) for provision of
samples of Bacillus sphaericus. Drs. M. S. Mulla (University of California, Riverside)
and L. A. Lacey (USDA, Gainesville, Florida) kindly reviewed the paper which led to
its improvement.

REFERENCES CITED

ABBOTT, W. S. 1925. A method of computing the effectiveness of an insecticide. J.
Econ. Entomol. 18: 265-267.
ALI, A., AND R. D. BAGGS. 1982. Seasonal changes of chironomid populations in a
shallow natural lake and in a man-made water cooling reservoir in central Florida.
Mosq. News 42: 76-85.
ANONYMOUS. 1985. Informal consultation on the development of Bacillus sphaericus
as a microbial larvicide. WHO/TDR/BCV/SPHAERICUS/85.3, 24 p.
DAVIDSON, E. W. 1984. Microbiology, pathology, and genetics of Bacillus sphaericus:
biological aspects which are important to field use. Mosq. News 44: 147-152.
DAVIDSON, E. W., A. W. SWEENEY, AND R. COOPER. 1981. Comparative field trials
of Bacillus sphaericus strain 1593 and Bacillus thuringiensis var. israelensis
commercial powder. J. Econ. Entomol. 74: 350-354.
DES ROCHERS, B., AND R. GARCIA. 1984. Evidence for persistence and recycling of
Bacillus sphaericus. Mosq. News 44: 160-165.
KELLEN, W. R., T. B. CLARK, J. E. LINDEGREN, B. C. Ho, M. H. ROGOFF, AND
S. SINGER. 1965. Bacillus sphaericus Neide as a pathogen of mosquitoes. J.
Invert. Pathol. 7: 442-448.
LACEY, L. A. 1984. Production and formulation of Bacillus sphaericus. Mosq. News
44: 153-159.
LACEY, L. A., AND S. SINGER. 1982. Larvicidal activity of new isolates of Bacillus
sphaericus and Bacillus thuringiensis (H-14) against anopheline and culicine
mosquitoes. Mosq. News 42: 537-543.
LACEY, L. A., M. J. URBINA, AND C. M. HEITZMAN. 1984. Sustained release formu-
lations of Bacillus sphaericus and Bacillus thuringiensis (H-14) for the control
of container-breeding Culex quinquetasciatus. Mosq. News 44: 26-72.
MULLA M. S., B. A. FEDERICI, AND H. A. DARWAZEH. 1982. Larvicidal efficacy of
Bacillus thuringiensis serotype H-14 against stagnant water mosquitoes and its
effect on nontarget organisms. Environ. Entomol. 11: 788-795.
MULLA, M. S., H. A. DARWAZEH, E. W. DAVIDSON, AND H. T. DULMAGE. 1984a.
Efficacy and persistence of the microbial agent Bacillus sphaericus against mos-
quito larvae in organically enriched habitat. Mosq. News 44: 166-173.
MULLA, M. S., H. A. DARWAZEH, E. W. DAVIDSON, H. T. DUIMAGE, AND S.
SINGER. 1984b. Larvicidal activity and field efficacy of Bacillus sphaericus
strains against mosquito larvae and their safety to nontarget organisms. Mosq.
News 44: 336-342.











690 Florida Entomologist 69(4) December, 1986

MULLA, M. S., H. A. DARWAZEH, L. EDE, B. KENNEDY, AND H. T. DULMAGE.
1985. Efficacy and field evaluation of Bacillus thuringiensis (H-14) and B.
sphaericus against floodwater mosquitoes in California. J. Am. Mosq. Control
Assoc. 1: 310-315.
MULLLIGAN, F. S., C. H. SCHAEFER, AND W. H. WILDER. 1980. Efficacy and
persistence of Bacillus sphaericus and B. thuringiensis H-14 against mosquitoes
under laboratory and field conditions. J. Econ. Entomol. 73: 684-688.
NAYAR, J. K. 1982. Wyeomyia mitchellii: observations on dispersal, survival, nutri-
tion, insemination, and ovarian development in a Florida population. Mosq. News
42: 416-427.
RAMOSKA, W. A., J. BURGESS, AND S. SINGER. 1978. Field applications of a bacte-
rial insecticide. Mosq. News 38: 57-60.





LIFE 48: A BASIC COMPUTER PROGRAM TO CALCULATE
LIFE TABLE PARAMETERS FOR AN INSECT OR
MITE SPECIES

M. M. ABOU-SETTA
Plant Protection Research Institute
Agriculture Research Center
Dokky, Giza 12611, Egypt
and
R. W. SORRELL, AND C. C. CHILDERS
University of Florida, IFAS
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, FL 33850 USA

ABSTRACT

A computer program in BASIC was developed following methods used by Birch
(1948) to calculate life table parameters for an arthropod from experimental data. Actual
data were presented as an example and data previously used by Birch (1948) were
included in this paper for verification. This program was written to run on VAX
minicomputer or IBM microcomputer compatibles.

RESUME

Se desarroll6 un program de computadoras en BASIC siguiendo los metodos usados
por Birch (1948) usando datos experimentales para calcular los parametros de un cuadro
sobre la vida de un artr6podo. Se present datos actuales como ejemplo y se incluy6
datos usados por Birch (1948) como verificaci6n. Este program se escribi6 para ser
usado en microcomputadoras VAX o con microcomputadoras compatibles con IBM.











Abou-Setta et al.: Life Table Computation


Birch (1948) identified the life table parameters to be used in calculating insect
population development by adapting human demography values. The primary popu-
lation parameter was the intrinsic rate of natural increase (rm) which fits the following
approximation for an insect species under a definite condition:

max r, X
1, (L,) (M) 1 (1)


where: e = the base of the natural logarithm, rm = the intrinsic rate of natural
increase, X = the female age, L, = the fraction of females alive at age X, and
M, = the expected number of daughters produced per female alive at age X.

Birch (1948) provided both approximate and precise methods for calculating the
value of rm. The approximate method calculated the mean generation time (T), using
equation (2), then estimated the value of rm using equation (3).

T = I ((NiL 1"M / R,, (2)

rn, In (R,) / T (3)

where: R,, = the net reproductive rate [I ((L,)(MJ))].

Birch (1948) tried different values until the appropriate precise value of rm was
reached.
The technique used by Birch (1948) was used in this program. The program first
calculates X, M,, L\, (M)(L,), and (X)(Lx)(M,) for each interval using the input data.
Applying equations (2) and (3) the approximate r. value is calculated. For each interval,
the product of (M,)(Lx) is then divided by the value of e raised to the power of [(rm)(X)]
to get the value of [e1-(-'"(x (M,)(L,)] as RML for each interval. The sum of RML over
all the intervals is compared to the range of 0.9995 to 1.0005. If the sum of RML is not
in that range, then the program reduces or increases the value of r,,, by 0.0001 and
executes the calculation again in a loop until the sum of RML reaches the indicated
range. The last trial rm value is considered the precise value.
The output of this program will be stored in a file (to be named by the operator).
This file includes the following information for each interval of adult female age: total
progeny per interval (M), number of females alive at age X (L,), mean female age at
each interval mid-point (X), female progeny per female (M,), rate of survival (L), the
product of [(Mx)(Lx)] as (MLx), and the final values of RML to meet the original
formula (equation 1). Finally, the program prints the precise life table parameters of
that study as the sum of RML, the net reproductive rate (R,,), the generation time (T),
the intrinsic rate of natural increase (r,,) and the finite rate of increase (e'").
An example of this program is presented (Fig. 1) using data of Euseius mesembinmfs
(Dean) (Acari: Phytoseitdae) reared on ice plant, Malephora crocea (Jacq.) at 30C
(Abou-Setta and Childers, unpublished data). To evaluate the accuracy of this program,
previously calculated (X), (Mx) and (L,) values from published data by Birch (1948) for
Sitophilus oryzae (L.) (Coleoptera: Curculionidae), were used. The obtained results of
this program gave a calculated rm value of 0.761957 vs. the estimated rm value of 0.76











Florida Entomologist 69(4)


LIFE TABLE DATA SHEET


THE MITE OR INSECT NAME : EUSEIUS
THE TEMPERATURE USED WAS : 30 C


M L

4 20
46 20
34 20
33 20
26 20
27 20
23 20
16 20
15 20
15 19
16 18
7 18
9 16
11 16
9 15
5 15
2 13
2 13


December, 1986


LIFE48.LIS


MESEMBRINUS (DEAN)


X Mx Lx MxLx


4.90
5.90
6.90
7.90
8.90
9.90
10.90
11.90
12.90
13.90
14.90
15.90
16.90
17.90
18.90
19.90
20.90
21.90


0.12
1.40
1.04
1.01
0.79
0.82
0.70
0.49
0.46
0.48
0.54
0.24
0.34
0.42
0.37
0.20
0.09
0.09


1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.95
0.90
0.90
0.80
0.80
0.75
0.75
0.65
0.65


0.122
1.403
1.037
1.007
0.793
0.824
0.701
0.488
0.458
0.458
0.488
0.213
0.275
0.336
0.275
0.153
0.061
0.061


RML

.036599
.329197
.190311
.144473
.089030
.072312
.048180
.026215
.019222
.015035
.012543
.004292
.004316
.004126
.002640
.001147
.000359
.000281


THE OBSERVATION (OBS.) INTERVAL USED HAS
THE DEVELOPMENTAL TIME WAS CONSIDERED AS
THE SEX RATIO WAS (FEMALES/TOTAL): .61
THE FRACTION OF EGGS REACHING MATURITY


THE
IHE
THE
THE
THE


1 DAY
4.4


SUM OF RML
NET REPRODUCTIVE RATE (Ro)
GENERATION TIME (T) IN OBS. INTERVALS
INTRINSIC RATE OF NATURAL INCREASE (rm)
FINITE RATE OF INCREASE


INTERVALS


1

= 1.00028
= 9.15
= 9.00947
= .245714
= 1.27853


COLUMN DEFINITIONS

M Total progeny at each interval for all females
L Number of females alive
X Actual female age (time from egg stage)
Mx Female progeny per female
Lx Proportion surviving at age X
MxLx- Female progeny per female times rate of survival
RML MxLx times (e raised to the power of (-rm times x))
(COLUMNS M & L CONTAIN THE INPUT DATA)

Fig. 1. Life 48 BASIC program output model.


by Birch (1948) (Fig. 2). This program substantially reduces the time and effort required
for calculating precise life table parameters.
This BASIC computer program (Fig. 3) uses either original life history data or
precalculated Mx and L, values. The program accepts the insect or mite name, temper-
ature used, number of intervals, type of interval used, and asks for the data type. If
the operator selects (1), the program will ask for the initial number of females, develop-
mental time of female immatures, percentage that reach maturity from the original
progeny, sex ratio (assuming that it is stable during the progeny time), total fecundity












Abou-Setta et al.: Life Table Computation


LIFE TABLE DATA SHEET


LIFE48.LIS


THE MITE OR INSECT NAME : CALANDRA ORYZAE


THE TEMPERATURE USED WAS : 29 C

X Mx Lx MxLx


4.50
5.50
6.50
7.50
8.50
9.50
10.50
11.50
12.50
13.50
14.50
15.50
16.50
17.50
18.50


20.00
23.00
15.00
12.50
12.50
14.00
12.50
14.50
11.00
9.50
2.50
2.50
2.50
4.00
1.00


0.87
0.83
0.81
0.80
0.79
0.77
0.74
0.66
0.59
0.52
0.45
0.36
0.29
0.25
0.19


17.400
19.090
12.150
10.000
9.875
10.780
9.250
9.570
6.490
4.940
1.125
0.900
0.725
1.000
0.190


RML

.564206
.288922
.085830
.032972
.015197
.007744
.003101
.001498
.000474
.000168
.000018
.000007
.000003
.000002
.000000


THE OBSERVATION (OBS.) INTERVAL USED WAS 1 WEEK
THE DEVELOPMENTAL TIME WAS CONSIDERED AS 4 INTERVALS


THE
THE
THE
THE
THE


SUM OF RML
NET REPRODUCTIVE RATE (Ro)
GENERATION TIME (T) IN OBS. INTERVALS
INTRINSIC RATE OF NATURAL INCREASE (rm)
FINITE RATE OF INCREASE


= 1.00014
= 113.485
= 6.20989
= .761957
= 2.14246


COLUMN DEFINITIONS

X Actual female age (time from egg stage)
Mx Female progeny per female
Lx Proportion surviving at age X
MxLx- Female progeny per female times rate of survival
RML MxLx times (e raised to the power of (-rm times x)
(COLUMNS Mx & Lx CONTAIN THE INPUT DATA)


Fig. 2. Validation of Life 48 program using data presented by Birch (1948).



per interval, and the number of females alive at each interval. If the operator selects
(2), the program will ask for the precalculated M_ and Lx values for each interval instead
of M and L. The program was developed to run on DEC VAX minicomputers or IBM
PC compatible microcomputers. A computer program in Turbo Pascal is planned.

ACKNOWLEDGMENT

Thanks to the AMIDEAST Peace Fellowship Program for supporting the first au-
thor during this study. This publication was in partial fulfillment of his requirements
for the Ph.D. degree.
Florida Agricultural Experiment Stations Journal Series No. 7396.


693











Florida Entomologist 69(4)


December, 1986


100 REM PROGRAM LIFE48 JULY 1985
110 REM M. M. ABOU-SETTA, R. W. SORRELL, AND C. C. CHILDERS
120 REM THIS PROGRAM CAN RUN ON BOTH A DEC VAX MINICOMPUTER
130 REM AND ON THE IBM PC MICROCOMPUTER AND COMPATIBLES IF
140 REM APPROPRIATE CHANGES ARE MADE AS DESCRIBED ON LINES
150 REM 1061 THROUGH 1064
160 REM DEFINE OUTPUT FILE
170 REM INPUT INITIAL PARAMETERS
180 ON ERROR GOTO 1580
190 INPUT ENTER MITE OR INSECT NAME ";TITLES
200 INPUT ENTER THE TEMPERATURE USED (C) ";TEMPS
210 INPUT ENTER THE NUMBER OF OBSERVATIONS ";N
220 PRINT ENTER THE TIME INTERVAL BETWEEN OBSERVATIONS ";
225 INPUT TIMINT$
230 PRINT ENTER THE DEVELOPMENT TIME FROM EGG TO ADULT
240 PRINT FEMALE AS THE NUMBER OF OBSERVATION INTERVALS ";
245 INPUT DT
250 PRINT
260 PRINT CALCULATE LIFE TABLE PARAMETERS USING
270 PRINT (1) LIFE HISTORY DATA ( M and L )"
275 PRINT (2) PRECALCULATED Mx AND Lx DATA"
280 PRINT
290 PRINT PLEASE ENTER YOUR DATA TYPE AS 1 OR 2 ",
300 INPUT DATYPE
305 PRINT
310 IF DATYPE 0 1 AND DATYPE <> 2 THEN 260
320 DIM X(N), M(N), M1(N), MX(N), L(N), LX(N), MXLX(N), XML(N)
325 DIM RML(N)
330 IF DATYPE = 2 THEN 640
340 INPUT ENTER THE INITIAL NUMBER OF FEMALES ";NF
350 PRINT ENTER THE FRACTION OF EGGS LAID REACHING";
355 INPUT MATURITY ";PERM
360 INPUT ENTER THE SEX RATIO AS FEMALES PER TOTAL ";SR
370 PRINT
380 PRINT FOR EACH OF ";N;" INTERVALS"
390 PRINT ENTER THE NUMBER OF EGGS LAID (M) AND ";
391 PRINT "THE NO. OF SURVIVING FEMALES (L)
400 PRINT
410 FOR C = 1 TO N
420 PRINT M FOR INTERVAL ";C;" ";
430 INPUT M(C)
490 PRINT L FOR INTERVAL ";C;" ";
500 INPUT L(C)
510 IF L(C) > 0 AND L(C) <= NF THEN 535
520 PRINT ERROR NUMBER MUST BE GREATER THAN 0 AND LESS";
525 PRINT THAN ";NF
530 GOTO 490
535 PRINT
540 NEXT C
550 PRINT
560 REM CALCULATING THE DAILY AND LIFE TABLE VALUES
570 FOR C = 1 TO N
580 X(C) = C + DT .5
590 M1(C)= M(C) / L(C)
600 LX(C) = L(C) / NF
610 MX(C) = M1(C) SR PERM
620 NEXT C
630 GO TO 740


Fig. 3. Listing of Life 48 program for life table calculations.


694












Abou-Setta et al.: Life Table Computation


640
645
646
647
650
660
670
680
690
700
710
720
730
735
740
750
760
770
780
790
800
810
820
830
840
850
860
865
870
880
890
895
900
910
920
930
940
950
960
965
970
980
990
100(
101(
102'
103'
104(
105'
106C
1061
1062
1063
1064
107C
108(
109(


EACH INTERVAL";


PRINT ENTER THE Mx AND Lx DATA FOR E
PRINT OF ADULT AGE"
PRINT FOR ";N;" INTERVALS"
PRINT
FOR C = 1 TO N
X(C) = C + DT .5
PRINT Mx FOR INTERVAL NO. ";C;"
INPUT MX(C)
PRINT Lx FOR INTERVAL NO. ";C;"
INPUT LX(C)
IF LX(C) / 1 THEN 690
PRINT
NEXT C
PRINT WORKING
FOR C = 1 TO C
MXLX(C)= MX(C) LX(C)
XML(C)= MXLX(C) X(C)
EXML = EXML + XML(C)
RO RO + MXLX(C)
NEXT C
REM CALCULATING THE APPROXIMATE VALUE
RM = LOG( RO ) / (EXML / RO)
FOR C = 1 TO N
RML(C) = MXLX(C) / EXP( X(C) RM )
ERML = ERML + RML(C)
NEXT C
REM FINDING THE ACCURATE (RM) USING THE
REM VALUE AS A GUIDE
IF ERML <= 1.0005 AND ERML =, .9995 THEN
IF ERML <.9995 THEN 960
ERML = 0
PRINT "*";
RM = RM + .0001
FOR C = 1 TO N
RML(C) = MXLX(C) / EXP( X(C) RM )
ERML = ERML + RML(C)
NEXT C
IF ERML > 1.0005 THEN 890 ELSE 1030
ERML = 0
PRINT "*";
RM = RM .0001
FOR C = 1 TO N
RML(C) = MXLX(C) / EXP( RM X(C) )
0 ERML = ERML + RML(C)
0 NEXT C
0 IF ERML < .9995 THEN S60
0 T = LOG( RO ) / RM
0 PRINT ENTER NAME OF FILE IN WHICH RE
0 INPUT "PLACED ";FILES


APPROXIMATE

1030


SULTS WILL BE ";


OPEN FILES FOR OUTPUT AS #1
REM FOR VAX COMPATIBILITY, A COMMA MUST NOT BE PLACED
REM AFTER THE #1 IN LINES 1260,1270,1560 ( PRINT #1 USING )
REM FOR IBM COMPATIBILITY, A COMMA MUST BE PRESENT
REM ( PRINT #1, USING CLOSE #1,
PRINT #1,"
PRINT #1," LIFE TABLE DATA SHEET";
PRINT #1," ";FILE$


695


OF RM


0
0


/












Florida Entomologist 69(4)


December, 1986


1100
1110
1120
1130
1140
1150
1160
1170
1180
1190
1200
1250
1260
1270
1280
1290
1300
1305
1310
1315
1320
1330
1340
1345
1350
1360
1365
1370
1375
1380
1385
1390
1395
1400
1405
1410
1420
1430
1440
1450
1455
1460
1470
1480
1490
1500
1505
1510
1515
1520
1530
1540
1550
1560
1570
1580
1590
1600
1610
1620


PRINT #1,"
PRINT #1," THE MITE OR INSECT NAME : ";TITLES
PRINT #1," THE TEMPERATURE USED WAS : ";TEMP$;" C"
PRINT #1,"
IF DATYPE = 2 THEN 1160
PRINT #1," M L ";
PRINT #1," X Mx Lx MxLx RML"
PRINT #1,
F1$ = ### ###"
F2$ = ##.## ##.## #.## ##.### .######"
FOR C = 1 TO N
IF DATYPE = 2 THEN 1270
PRINT #1, USING Fl$; M(C); L(C);
PRINT #1, USING F2$; X(C); MX(C); LX(C); MXLX(C); RML
NEXT C
PRINT #1,"
PRINT #1," THE OBSERVATION (OBS.) INTERVAL USED WAS "
PRINT #1, TIMINT$
PRINT #1," THE DEVELOPMENTAL TIME WAS CONSIDERED AS "
PRINT #1,DT;" INTERVALS"
IF DATYPE = 2 THEN 1350
PRINT #1," THE SEX RATIO WAS (FEMALES/TOTAL): ";SR
print #1," THE FRACTION OF EGGS REACHING MATURITY
PRINT #1,PERM
PRINT #1," "
PRINT #1," THE SUM OF RML
PRINT #1," = ";ERML
PRINT #1," THE NET REPRODUCTIVE RATE (Ro)
PRINT #1," = ";RO
PRINT #1," THE GENERATION TIME (T) IN OBS. INTERVALS
PRINT #1," = ";T
PRINT #1," THE INTRINSIC RATE OF NATURAL INCREASE (rm
PRINT #1," = ";RM
PRINT #1," THE FINITE RATE OF INCREASE
PRINT #1," = ";EXP(RM)


) I'


PRINT #1," "
PRINT #1," COLUMN DEFINITIONS"
PRINT #1," "
IF DATYPE = 2 THEN 1470
PRINT #1," M Total progeny at each interval for all";
PRINT #1," females"
PRINT #1," L Number of females alive
PRINT #1," X Actual female age (time from egg stage)"
PRINT #1," Mx Female progeny per female"
PRINT #1," Lx Proportion surviving at age X"
PRINT #1," MxLx- Female progeny per female times rate";
PRINT #1," of survival"
PRINT #1," RML MxLx times (e raised to the power of";
PRINT #1," (-rm times x))"
IF DATYPE = 2 THEN 1550
PRINT #1," (COLUMNS M & L CONTAIN THE INPUT DATA)"
GOTO 1560
PRINT #1," (COLUMNS Mx & Lx CONTAIN THE INPUT DATA)"
CLOSE #1,
GOTO 1700
REM ERROR HANDLING ROUTINE
IF ERL > 740 THEN 1640
PRINT DATA ERROR PLEASE CHECK DATA AND RE-ENTER"
IF ERL = 430 THEN RESUME 420
IF ERL = 500 THEN RESUME 490


(C)


;











Abou-Setta et al.: Life Table Computation


1621 IF ERL = 680 THEN RESUME 670
1622 IF ERL = 700 THEN RESUME 690
1640 IF ERL > 1060 THEN 1670
1650 PRINT ERROR OPENING OUTPUT FILE TRY AGAIN"
1660 RESUME 1040
1670 PRINT ERROR IN PROGRAM"
1680 PRINT ERROR #";ERR;" OCCURRED AT LINE ";ERL
1690 RESUME 1700
1700 END


REFERENCE CITED

BIK(H, L. C. 1948. The intrinsic rate of natural increase of an insect population. J.
Anim. Ecol. 17: 15-26.





DELTA CAMPANIFORME RENDALLI (BINGHAM) AND
ZETA ARGILLACEUM (LINNAEUS) ESTABLISHED IN
SOUTHERN FLORIDA, AND COMMENTS ON GENERIC
DISCRETION IN EUMENES s. 1.
(HYMENOPTERA: VESPIDAE: EUMENINAE)

A. S. MENKE
Systematic Entomology Laboratory
Agricultural Research Service, U.S.D.A.
% U. S. National Museum of Natural History
Washington DC 20560
and
L. A. STANGE
Florida Department of Agriculture and Consumer Services
Division of Plant Industry
P. O. Box 1269, Gainesville, Florida 32601

ABSTRACT

Delta campanifbrme rendalli (Bingham), an African wasp, and Zeta argillaceum
(Linnaeus), a South American wasp, are established in southern Florida. These insects
add two more genera to the North American fauna. The existing key to the North
American genera is modified to include Delta and Zeta.

RESUME

La avispa africana Delta campaniforne rendalli, y la avispa sudamericana Zeta
argillaceum (Linnaeus) estan establecidas en el sur de la Florida. Estas afiaden dos
g6neros a mas de insects a la fauna de Norteamerica. Se modific6 la clave de los
generous de Norteamerica para incluir a Delta y a Zeta.











Florida Entomologist 69(4)


Two exotic species of potter wasps are now established in southern Florida. Delta
campanifbrme rendalli is an immigrant from Africa and Zeta argillaceum comes from
South America. Like the related genus Eumenes Latreille, species of Delta Saussure
and Zeta Saussure make nests of mud that are often affixed to substrates easily trans-
ported by man. Thus it is not surprising that D. c. rendalli and Z. argillaceum were
accidently introduced to Florida, where both have become established in the region
around Miami and Fort Lauderdale. Unlike most North American species of the native
genus Eumenes that are black and yellow, the two introduced wasps are largely reddish
brown and black. However, one of the two Floridian Eumenes, smithii Saussure, is
similarly colored. The steplike apical margin of tergum II, a generic feature of Eumenes,
distinguishes E. smithii from D. c. rendalli and Z. argillaceum, both of which have
simple terga. The key to genera presented below will identify the two introduced wasps.
Voucher material of the two species is in the Florida State Collection of Arthropods
in Gainesville, and the National Museum of Natural History, Washington DC. We would
like to thank Jim Carpenter, Museum of Comparative Zoology, Harvard University,
Cambridge, Mass. and Frank Parker, Bee Biology and Systematics Lab., USDA, Utah
State Univ., Logan, Utah for their comments on this paper.

Delta campaniforme rendalli (Bingham)

This wasp was described from Nyasaland, now Malawi, by Bingham (1902), and the
species was later recorded from Zaire and Zimbabwe by Bequaert (1926). Carpenter (in
litt.) says that there is a specimen from Mozambique in the Museum of Comparative
Zoology. D. c. rendalli was apparently first collected in Broward County in December,
1981, a few miles southwest of Fort Lauderdale at Davie (1 female), and also at Fort
Lauderdale (3 males). These, as well as subsequent material, were collected by J. A.
Reinert. The following year three additional males and one female were collected at
Fort Lauderdale (January), and three more males and one female were taken at Davie
(September) visiting flowering Borreria verticillata (L.) G. F. W. Meyer (= Spermacoce
verticillata for a total of 12 specimens.
This material compares well with specimens of Delta campaniforme rendalli in the
USNM from Zaire that were determined by J. Bequaert, and also runs to rendalli in
Bequaert's key (1926) to the African species of Eumenes s. 1. In most of the Florida
males the pronotal dorsum is reddish brown except for black posterolaterally and traces
of yellow along the transverse carina. In one male the reddish brown is replaced by
yellow. The four Zaire males studied (USNM) display both patterns.
Bequaert (1926) treated rendalli as a subspecies of the widely distributed Old World
species campaniforme (Fabricius). Some subspecies of this wasp have been elevated to
species status in recent years, and rendalli may warrant the same treatment. In his
brief report on this wasp, Freeman (1984) treated rendalli (misspelled randalli) as a
species without comment, but as far as we can determine, no one has formally elevated
rendalli to species. Both Giordani Soika (in litt.) and Carpenter (in litt.) have expressed
the opinion that rendalli should be treated as a species, but lacking evidence for this
we have followed a conservative interpretation of the taxon.
Bingham described both sexes of rendalli, but Bequaert (1926) indicated that the
male and female syntypes were not conspecific. He restricted the name to the female
syntype, and later (1928) effectively selected it (according to the provisions of Article
74 (a) of the current Code) as the lectotype.
Interestingly, Delta c. rendalli is also established on the West Indian island of
Jamaica (Freeman, 1984) where it was first captured in 1979. Freeman included a


698


December, 1986











Menke & Stange: New U.S. Genera of Wasps 699


photograph of the mud nest that was attached to the wall of a building. The abandoned
nest was apparently taken over by a species of Pachodynerus.

Zeta argillaceum (Linneaus)

This is a widespread, common neotropical potter wasp (Mexico to Argentina-see
Soika, 1975), but apparently it does not occur in the West Indies except on Trinidad.
We have seen 21 specimens that were collected in Dade Co. over a period of 10 years.
The earliest example was taken in July, 1975 at Miami. By 1980 the wasp had been
collected at Miami Springs and Hialeah, and by 1981 had reached Fort Lauderdale. In
1983 Stange and Woodruff found a single celled nest of argillaceum attached to the
underside of a foam pad on the ground at Hialeah, indicating that the species was
established and reproducing.
Soika (1975), in his review of the genus Zeta, recognized 10 subspecies of argil-
laceum. The Florida population belongs to the typical subspecies which occurs in the
Guianas and Brasil.
The biology of argillaceum was discussed by Taffe (1979) under the name
canaliculatum. He estimated that in Trinidad this wasp had 6 generations per year,
each lasting about 60 days. Taffe stated that argillaceum's biology was similar to the
related Z. abdominale (Drury) whose life history has been discussed in a number of
papers (Freeman and Taffe, 1974, Taffe and Ittyeipe, 1976, and Taffe, 1978, 1979,
1983-the name Eumenes colona was used in the earlier papers). These authors studied
abdominale in Jamaica. The mud nests are built on sunlit substrates sheltered from
rainfall, and are usually 1.5 m from the ground. The nests contained 1 to 22 cells, but
single celled nests predominated. Various lepidopterous caterpillars were provisioned,
especially geometrid loopers. Four to 14 prey were provisioned per cell depending on
caterpillar size. Provisioning of a cell takes 2 to 3 days. The egg is suspended from a
filament attached to the rear of the cell. Development from egg to adult takes 5 to 6
weeks. Zea abdominale and Z. argillaceum are attacked by eulophid wasps of the
genus Melittobia Westwood, and miltogrammine flies of the genus Amobia Robineau-
Desvoidy. Mud nests of abdominale may persist for 5 years and they are utilized by a
variety of inquilines: Pachodynerus nasidens (Latreille), P. jamaicensis Bequaert,
Monobia mochii Soika (all Eumeninae), Megachile concinna Smith (Megachilidae), and
Trypoxylon texense Saussure (Sphecidae).
Zeta argillaceum may be established in Tahiti also. There is a single male in the
National Museum of Natural History collected by Jack Clarke at the Fautaua River,
Oct. 17, 1961, in Tahiti.
These introductions add two more genera to the eumenine fauna of North America:
Delta and Zeta', both names attributable to Saussure (1855). For complete generic
citations and synonyms of these two genera see Carpenter (in press). Both genera run
to the genus Minixi Soika in the recent key to the North American genera of Eumeninae
by Carpenter and Cumming (1985). We have emended their key to include Delta and
Zeta. The characters employed here for Delta may not work for all other species of this
genus.

3. Apical margin of tergum II depressed, set off from rest of tergal sur-
face as a lam ella ................................... ........................... .. ... 3a.
Apical margin of tergum II not depressed, not set off from rest of tergal
surface .............. ............... ...... ....... ...... ... ............. 3b.

'The gender of Delta and Zeta, which are letters of the Greek alphabet, is neuter.











Florida Entomologist 69(4)


3a. Pronotum without pretegular carina .......................... Eumenes Latreille".
Pronotum with pretegular carina (fig. 1) ............................... Minixi Soika.
3b. Pronotum with incomplete humeral carina (fig. 2); petiole spiracle
located at midpoint of segment ........................................... Zeta Saussure.
Pronotum without humeral carina; petiole spiracle located beyond mid-
point of segment ............................................... Delta Saussure.

A few comments on the recognition of Delta and Zeta as genera are warranted
because their history is scattered in the literature. Eumenes, in its original sense as
exemplified by Saussure's (1852, 1855) pioneering work on eumenine wasps, contained
a number of groups that he (1855) called divisions and to which he applied "generic"
names. These division names later were recognized as subgenera (Bequaert, 1926, for
examples), and eventually as separate genera (Soika, 1961, Bluthgen, 1961, for exam-
ple). Bequaert (1926) said "Eumenes ... contains over 200 ... species, many of which
differ greatly in shape." "If the fauna of a limited area be considered, they are rather
easily arranged in natural groups, but whether these are of sufficient value to be re-
garded as valid subgenera is a troublesome question." He recognized subgenera, any-
way. By the 1960's some of these subgenera (Delta for one) had been elevated to genera
(Soika, 1961, Bluthgen, 1961). Soika's paper was a world survey of the groups within
Eumenes in the old sense. He elevated most of the subgenera to genera, and subdivided
the component species into groups. Zeta Saussure, another section of the old Eumenes,
was treated as a genus by Soika (1975) when he reviewed its species. He had recognized
Zeta as a genus in 1972 but did not define its characters. Soika continued his generic
fragmentation of Eumenes s. 1. by publishing a large work in 1978 in which nine new
genera were proposed, mostly for species groups recognized in his 1961 paper under
the genus Omicron. One of these new genera was Minixi.
The total number of "eumenes-like" genera now recognized worldwide is about 25
according to Carpenter (in litt.), but this tally is not easy to elucidate due to the
piecemeal fashion in which they have been recognized and the lack of a definitive global
review of these taxa with descriptions and keys to them. This "evolution" of Eumenes
s. 1. into many genera is paralleled throughout the entire subfamily. In our opinion this















1 Minixi mexicanum 2 Zeta argillaceum

Fig. 1-2, left profile of part of thorax. Fig. 1, Minixi showing pretegular carina
(arrow); Fig. 2, Zeta showing incomplete humeral carina (arrow).


'See Grissell (1974) for identification of the two Florida species.


700


December, 1986











Menke & Stange: New U.S. Genera of Wasps 701

splitting is extreme, and it reflects the fact that specialists in the family fail to appreciate
that genera have an important practical component: a generic name should convey
information to the broad user community. The old cliche "unable to see the forest for
the trees" applies to those who have split Eumenes, a meaningful taxon, into many
small genera with similar facies. Granted, Eumenes in its original sense may not be
monophyletic (Carpenter, in litt. says that it probably was) and thus is open to
taxonomic refinement via some splitting, but we believe this has progressed to an
irrational point. Carpenter and Cumming (1985) quoted Bequaert's 1939 cogent assess-
ment of the trend that he saw toward extreme splitting of eumenine genera: "Such a
procedure, however, not only leaves out the many annectant species, but it fails as a
guide to the study of natural relationships." As Carpenter and Cumming point out, this
statement is still an appropriate criticism of the current state of affairs in this subfamily
(and indeed in the family). Unfortunately, the recent cladistic analysis by Carpenter
and Cumming (1985) only hints at possible future lumping of some eumenine genera.
Because they dealt primarily with the North American fauna, they did not attempt to
make decisions on the validity of many of the "genera" split off from the old Eumenes,
but Carpenter (in litt.) agrees that some are doubtless untenable. In fact, some of the
"eumenes"-like genera are probably paraphyletic.
The many "eumenes"-like genera now recognized are separated for the most part
by characters that may be more appropriately used at the species group level than at
the generic: presence or absence of various thoracic carinae, petiole shape, presence or
absence of a lamella on tergum II and so on. It is unknown how these hold up on a world
basis, but variation seems likely, for as early as 1875 Saussure wrote: "This genus
[Eumenes s. 1.] .... is broken up into peculiar types". "These types are connected by
natural transitions which embarrass one in assigning them very fixed limits." The genus
Ammophila in the Sphecidae contains a number of groups that are as distinctive as
some of the genera derived from Eumenes. To elevate these to genera would destroy
a concept that denotes a substantial amount of information content, a generic concept
that is meaningful, hence practical, to a large audience. The old Eumenes was an easily
recognized taxon. We are not suggesting that everything be put back into Eumenes,
but surely some of the current "genera" are at best subgenera or at most species groups.

ENDNOTE

Contribution No. 635, Bureau of Entomology, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, FL.

REFERENCES CITED

BEQUAERT, J. 1926. The genus Eumenes, Latreille, in South Africa, with a revision
of the Ethiopian species (Hymenoptera). Ann. South African Mus. 23: 483-577.
1928. A study of certain types of diplopterous wasps in the collection of the
British Museum. Ann. Mag. Nat. Hist. (10)2: 138-176.
BINGHAM, C. T. 1902. On the Hymenoptera collected by Mr. W. L. Distant in the
Transvaal, South Africa, with descriptions of supposed new species. Ann. Mag.
Nat. Hist. (7)10: 207-222.
BLUTHGEN, P. 1961. Die Faltenwespen Mitteleuropas (Hymenoptera, Diploptera).
Abhandl. Deutsch. Akad. Wiss. Berlin 1961(2): 1-248.
CARPENTER, J. M. (in press). A synonymic generic checklist of the Eumeninae
(Hymenoptera: Vespidae). Psyche.
CARPENTER, J. M., AND J. M. CUMMING. 1985. A character analysis of the North
American potter wasps (Hymenoptera: Vespidae; Eumeninae). J. Nat. Hist. 19:
877-916.











702 Florida Entomologist 69(4) December, 1986


FREEMAN, Brian. 1984. Delta randalli [sic] (Bingham) established in Jamaica
(Eumenidae). Sphecos (9): 13-14.
FREEMAN, B. AND C. TAFFE. 1974. Population dynamics and nesting behaviour of
Eumenes colona (Hymenoptera) in Jamaica. Oikos 25: 388-394.
GRISSELL, E. Eric. 1974. The Eumenes, or potter wasps, of Florida. Florida Dept.
Agric. Consum. Serv., Ent. Circ. (146): 1-2.
SAUSSURE, HENRI DE. 1852-[1853]. Etudes sur la famille des Vespides 1. Mono-
graphie des guepes solitaires ou de la tribu des Eumeniens, [etc.]. lii + 286 p.
[1854]-1856. Etudes sur la famille des Vespides 3. La monographie des Masa-
riens et un supplement a la monographie des Eumeniens. 352 p.
S1875. Synopsis of American wasps. Smith. Misc. Coll. 254: xxxv + 385 p.
SOIKA, A. GIORDANI. 1961. Les lignees philetiques des Eumenes s.1. du globe (Hym.,
Vesp.). Verhand. XI Internat. Kongr. Ent. Wien 1960 1: 240-245.
1975. Sul genere Zeta (Sauss.). Boll. Mus. Civ. Storia Nat. Venezia. 27: 111-
135.
1978. Revisione degli Eumenidi neotropicali appartenenti ai generi Eumenes
Latr., Omicron (Sauss.), Pararaphidoglossa (sic] Schulth. ed affini. Boll. Mus.
Civ. Stor. Nat. Venezia 29: 1-420.
TAFFE, C. A. 1978. Temporal distribution of mortality in a field population of Zeta
abdominale (Hymenoptera) in Jamaica. Oikos 31: 106-111.
1979. The biology of two West Indian species of mud-wasps (Eumenidae:
Hymenoptera). Biol. J. Linn. Soc. 11: 1-17.
1983. The biology of the mud-wasp Zeta abdominale (Drury) (Hymenoptera:
Eumenidae). Zool. J. Linnean Soc. 77: 385-393.
TAFFE, C., AND K. ITTYEIPE. 1976. Effect of nest substrata on the mortality of
Eumenes colona Saussure (Hymenoptera) and its inquilines. J. Anim. Ecol. 45:
303-311.
VAN DER VECHT, J., AND F. C. J. FISCHER. 1972. Palaearctic Eumenidae. Hymen.
Cat. (n. edit.), Pars 8, v + 199 p.





ANTLION PIT CONSTRUCTION AND
KLEPTOPARASITIC PREY

JEFFREY R. LUCAS
Department of Biology
College of William and Mary
Williamsburg, VA 23185

ABSTRACT

At a site on Archbold Biological Field Station (Lake Placid, Florida), 25% of the diet
of antlion larvae (Myrmeleon spp.) consisted of a single species of an ant (Conomyrma
sp.); however, in about 10% of antlion pits containing prey, Conomyrma successfully
kleptoparasitized, or stole prey from, the antlion. Stolen prey were significantly larger
than other prey items. Features that increase the capture efficiency of the pit (steep
slope and a layer of fine sand on the walls) are disrupted during the attempted escape
of large prey; this decreases the risk of predation by antlions on kleptoparasitic ants.
In the laboratory, antlions preferred fine sand over coarse, and moved longer distances
when placed on coarse sand. The former behavior will increase prey capture success
and reduce the risk of kleptoparasitism. The latter behavior increases the probability
of finding a more suitable habitat.











Lucas: Antlion Pit Construction


RESUME

En un lugar de el Campo de Investigaciones Biol6gicas Archbold (Lake Placid, Flori-
da), 25% de la dieta de la larva de la "hormiga leon" (Myrmeleon spp.) consiste de una
sola especie de hormiga (Conomyrma sp.); sin embargo, alrededor de un 10% de los
pozos de la "hormiga le6n" con presa, Conomyrma cleptoparasit6 o le rob6 la presa a
la "hormiga leon." Las press asi robadas eran significativamente mas grandes que
otras posibles press. Las caracteristicas que aumentan la eficacia capturadora del pozo
(ladera empinada con una capa de arena fina en las paredes se desbaratan cuando una
presa grande intent escapar; esto disminuye el riesgo de depredacion por la "hormiga
leon" sobre la hormiga cleptoparasitica. En el laboratorio, la "hormiga le6n" prefiere la
arena fina en lugar de la gruesa, y se va mas lejos cuando se coloca en arena gruesa.
Su eficacia como predador debe ser mayor cuando se encuentra en la arena fina al mismo
tiempo que reducird el riesgo de cleptoparasitismo. Este ultimo comportamiento au-
menta la probabilidad de encontrar un medio mas propicio.



Various arthropods construct some form of trap to aid in prey capture (Wheeler
1930, Hansell 1984). One of the more unusual traps is constructed by larval antlions
which build conical pitfalls in sand or fine soil (Wheeler 1930, Youthed and Moran 1969).
These pitfall traps serve three general functions: (1) the enhancement of capture effi-
ciency by retarding prey escape (Wilson 1974, Lucas 1982), (2) increasing the distance
from the antlion over which prey can be captured (Griffiths 1980), and (3) avoidance
anti-predator tactics (such as biting or spraying with noxious chemicals) of potentially
harmful prey by retreating under the sand (without the prey) or by pulling them under
the walls of the pit (Lucas and Brockmann 1981, Conner and Eisner 1983).
The first two functions depend on three aspects of pit design: pit diameter, pit-wall
slope, and sand grain size. Large pits generally catch more prey as well as larger range
of prey sizes (Griffiths 1980, Heinrich and Heinrich 1984). An increase in pit slope and
decreasing grain size increases capture efficiency over the entire range of prey sizes
(Lucas 1982). To some extent, all three of these pit-design features can be regulated by
the antlion. Antlions exert some control over pit-wall slope and can compensate for sand
of mixed grain size by discarding large particles and retaining finer particles with which
they line the pit (Lucas 1982). Thus antlions build an efficient trap lined with fine sand
even if they are working in a mixture of coarse and fine sand.
When prey attempt to escape however, the pit structure changes because sand is
knocked off the pit walls. This change in structure is relevant because antlions that are
handling prey often attempt to capture additional prey that enter their pits (Griffiths
1980, Lucas 1985). Capture success under these conditions is significantly lower than
with fresh or undisturbed pits (Lucas 1985). The exact cause of this reduced capture
success has not been studied.
Here I analyze the specific features of prey capture that reduce the capture efficiency
of successive prey. This analysis is used to relate the ability of ants to escape from
antlion pits, and also to enter and steal prey from pits which have been previously
disrupted. Since sand grain size is a critical determinant of the pit efficiency, the capac-
ity of antlions to choose among substrates that differ in grain size is addressed.
METHODS

FIELD

The antlion population was located in a shaded area under a large open shed at
Archbold Field Station, Lake Placid, Florida. On 7-8 July and 23-24 August 1983, 250











Florida Entomologist 69(4)


pits were examined every 4 hr for a 20 hr period (3000 pits total). The diameter of each
pit was measured. Any prey were removed and stored in 75% ethanol after which the
antlion was dug from the pit to determine species and instar (descriptions in Lucas &
Stange 1981). The presence and behavior of ants in the pit was also noted. Only pits
containing prey were dug up during the 4 hr census period. After each 20 hr period,
the diameters of approximately 75 randomly chosen pits were measured. All associated
antlions were then dug up to determine the species of antlions at the site and possible
species differences in pit size. In the laboratory, each prey organism was measured to
the nearest mm and adults were identified to family.

LABORATORY

Two laboratory experiments were conducted using third instar Myrmeleon crudelis
Walker.
(1) Pit structure and distribution of sand particles after prey capture: Ten antlions
were allowed to construct pits in color-coded sand composed of two grain sizes: 125-250
jim diameter (white) and 500-1000 jim diameter (red). The color-coding allowed visual
inspection of the distribution of sand grains in the pit (Lucas 1982). The antlions were
then divided into two groups, and each group was given one of two prey types: vestigial-
winged fruit flies (Drosophila melanogaster Meigen) which were about 2 mm long, and
worker carpenter ants (Camponotus floridanus Buckley) which were about 13 mm long.
(2) Choice of sand for pit construction: Fifty-three M. crudelis were placed in 25 x
15 cm plastic boxes filled with sand to about 6 cm depth. Antlions were given a choice
between fine sand (125-250 jm diameter) placed in one half of the box and one of three
different sizes of coarser sand placed in the other half. Antlions were initially placed in
the center of the fine sand on about 1/2 of the trials, and on the coarse sand for the rest
of the trials. In two tests the antlions were given a single grain size throughout the
box. Total movement before pit construction was estimated by measuring the length of
a furrow left in the sand by the moving antlions. Subsequent placement of the pit was
noted.

RESULTS

FIELD
Three species of antlions occurred in the population. On July 8, 1983, among the 65
M. crudelis dug out, there were 68.8% first instars, 15.6% each of second and third
instars; M. carolinus (Banks) (17 individuals) had 23.5% each of first and second instars
and 53.0% third instars. The 4 M. mobilis Hagen caught were all third instars. On
August 24, 1983, M. crudelis (60 individuals) consisted of 31.7% first instars, 43.3%
second instars and 25.0% third instars; M. carolinus (5 individuals) had 60% second
instars and 40% third instars. The only M. mobilis caught was a second instar.
For all species, later instars built significantly larger pits. Pit size was also signific-
antly different between species; this difference was generally due to the smaller pits of
M. carolinus (Table 1). Of the 3000 pits examined, 65 contained prey. Larger pits
generally contained significantly larger prey, with no residual effect for species or instar
(Table 1). Antlions caught a wide variety of both aerial and terrestrial arthropods, but
the ant Conomyrma (genus being revised, voucher series deposited at the Division of
Plant Industry Gainesville, Florida) (E. Nickerson, pers. comm.) accounted for 25% of
the number of prey caught (Table 2).
Although Conomyrma was an important prey at this site, ants of this species occa-
sionally kleptoparasitized antlions (Table 3) (kleptoparasitism is the intra- or inter-spe-


December, 1986











Lucas: Antlion Pit Construction 705


TABLE 1. MULTIPLE REGRESSION ANALYSIS FOR (A) ANTLION PIT SIZES AND (B)
PREY SIZE AT ARCHBOLD STATION, 1983a

A) Antlion Pit Size

independent variablesb degrees of freedom F value Prob F
antlion species 2 3.02 0.050
carolinus vs crudelis 1 5.22 0.023
carolinus vs mobilis 1 2.88 0.091
crudelis vs mobilis 1 0.24 0.624
antlion instar 2 521.0 0.001
error 211

B) Prey Size

independent variablesb degrees of freedom F value Prob F
antlion pit size 1 4.95 0.007
antlion species 2 1.07 0.351
antlion instar 2 0.12 0.734
error 59

"All regression analysis were run on SAS computer program GLM (Ray 1982).
'No interaction terms were significant (P=0.05), so these were deleted from the regression model.
'Species compared using "contrast" option of GLM.


TABLE 2. PREY OF ANTLIONS AT ARCHBOLD FIELD STATION IN 1983.

length Pit size antliona antlion
Order Family N (SD)(mm) (SD)(mm) species instar


Coleoptera
Chrysomelidae
Chrysomelidae(?)
Coccinellidae
Curculionidae



Scarabaeidae

Scolytidae
Staphylinidae
Tenebrionidae
? (larva)
Lepidoptera
? (microlep.)
? (larva)



Dermaptera
Labiduridae
Labiidae


2(0)
4
2
5
5
5
5
4
4(0)
2
3
6
6

3
13(1)
21
13(4)
17

12
3
3


62(33)
105
20
28b
37
73
80
75
43(1)
43
44
35
35

63
56(1)
80
57(1)
80'

94
18
30


car
cru
cru
cru
cru
cru
mob
car
cru
cru
cru
cru
cru












Florida Entomologist 69(4)


Trichoptera
Leptoceridae
Embioptera
Oligotomidae
Diptera
Anthomyiidae(?)
Chloropidae(?)
Culicidae
Muscidae
Tabanidae
Hemiptera
Pentatomidae
Homoptera
Cicadellidae
Delphacidae
Hymenoptera
Eulophidae
Formicidae
Pheidole



Solenopsis


Camponotus

Conomyrma


1 3

1 8


5
2(0)
15
6(2)
10


1 16 100b


1 5


2
2(0)
2
2
5(0)
3
3
15
16
3(0)
3(0)
3(0)
3(0)
3


58

25

95
52(0)
65 b
78(18)
70b


cru

cru


car
cru
cru
mob
car


cru


88
100

53

56
22(6)
26
68
32(6)
70
60
55b
77
65(23)
35(2)
45(3)
76(9)
78


cru
mob

cru

car
cru
cru
cru
cru
cru
cru
cru
cru
car
cru
cru
cru
mob


"cru = Myrmeleon crudelis; car = M. carolinus; mob = M. mobilis.
denotes pits in which CononWyrWta stole at least part of the prey.

cific stealing of already procured food; Brockmann and Barnard 1979). In three cases,
seven or more ants stole an entire carcass (a carpenter ant, a tabanid fly and a caterpil-
lar). In each of these instances, the ants began a "tug-of-war" with the antlion over the
prey, after which the antlion released the prey and retreated under the sand. Other
instances involved a single ant removing a leg from a culicid, a single ant removing a
leg from a curculionid, and three ants removing the head from a pentatomid bug. In
the latter case, one ant fell back into the pit and was caught by the antlion. This was
the only observed instance of a kleptoparasite being killed. In one other instance, a 25
mm noctuid caterpillar introduced into a pit was stolen by about 30 ants. During the
struggle, the antlion (a third instar M. carolinus) was pulled from the sand and was
also taken by the ants to their nest about 0.5 m away. Thus although Conomyrma is
an important prey species in this habitat, it is also a potential kleptoparasite and pre-
dator, especially when foraging in groups.
All pits in which kleptoparasitism occurred were in poor repair, probably due to
prey capture. Stolen prey were significantly larger than other captured prey (z=3.52,
P<0.001, Mann-Whitney U test). Since large antlion pits were more likely to capture
large prey, kleptoparasitism was also more common with antlions constructing large
pits, (z=2.44, P=0.007, Mann-Whitney U test).


December, 1986











Lucas: Antlion Pit Construction


Laboratory: (1) Effects of Prey on Pit Structure

Pit structure was completely unaltered by the capture of fruit flies. However, in all
cases where carpenter ants were fed, pit slope decreased and the pit tended to fill with
large sand particles (Fig. 1). Thus large prey are more likely to disturb the pit than are
small prey. This disturbance will affect subsequent capture success, and should also
reduce the probability that Conomyrnma would be caught when entering the pit to steal
antlion prey.

Laboratory: (2) Choice of sand grain size

Sand grain size influenced at least two aspects of antlion behavior: total distance
moved and choice of sand for pit construction. Larvae tended to travel farthest before
constructing a pit when given a combination of large grain size in both sides (Table 3).
Nine of 35 antlions placed in the two finer sizes of sand (125-250 and 350-500 rnm)
never moved far enough to test both sides before constructing a pit, whereas all 24
placed in the coarser sand moved at least the length of the box (Table 3; difference
between two groups: p=0.006, Fisher Exact Probability Test). Of the antlions that
potentially had a choice between grain sizes, 94 percent chose the finer sand when
started in the fine sand, and 94 percent chose the finer sand when started in the coarse
sand (Table 3; chi-square=20.1, P<0.001). Thus antlions preferred the finer-grained
sand.


Fig. 1. Distribution of small (white=125-250 jm diameter) and large (black=500-
1000 p.m) sand grains in (A) a newly constructed pit, and (B) a pit after the capture of
a Camponotus floridanus worker. The antlion's head is located at the tip of each arrow
and facing it. Light-colored area is the pit.


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Florida Entomologist 69(4)


December, 1986


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Lucas: Antlion Pit Construction


DISCUSSION

The efficiency of any trap depends upon both the nature of the construction materials
and on the arrangement of those materials (Denny 1976, Lucas 1982). As noted previ-
ously, the antlion exhibits several behaviors that enable it to regulate a number of
aspects of the pit design. When large prey attempt to escape however, the pit tends to
fill with the largest sand particles, disrupting the initial design. The filling of the pit
with large sand particles is caused by the physical properties of sand. Large particles
on a slope have a lower stability than do smaller particles, and therefore are more likely
to fall off the pit walls when they are disturbed (Lucas 1982). This is not a problem
during pit construction, because it helps concentrate the larger particles, allowing for
easier removal. But after prey are caught, the capture success of additional prey is
reduced if large particles are present on the walls and if the walls have a shallow slope.
Since ants can move more easily in pits with shallow walls composed of large sand grains
(Griffiths 1980, Lucas 1982), the risk of kleptoparasitism should increase when the pit
is destroyed by large prey. Thus the ability of antlions to select areas of finer-grained
sand for pit construction aids in prey capture (both initial and successive) and avoidance
of kleptoparasitism.
Antlions have been shown to retreat from potentially harmful prey by drawing back
under the sand (Lucas and Brockmann 1981, Conner and Eisner 1983). While this re-
duces the risk to the antlion, it also allows the prey to escape unharmed. This behavior
was seen several times at Archbold when the antlion was attempting to capture Con-
omyrma ants and when Conomyrma were stealing antlion prey. In this respect, group
foraging may reduce the risk to Conomyrma by increasing the risk to the antlion of
capturing these ants.
Kleptoparasitism is most likely to occur when the host takes large, high quality
prey, because the cost of kleptoparasitism would more likely be offset if the reward was
relatively large (Brockmann and Barnard 1979). Kleptoparasitism by Conomyrma was
also significantly more common for the largest prey caught by antlions. But in this
system pit structure is disrupted by large prey and therefore the risk of predation by
antlions on kleptoparasitic Conmyrma is reduced due to the concomitant drop in the
efficiency of the trap in retarding prey (and kleptoparasite) escape.

ENDNOTE

I thank Greg Capelli, Lynda Peterson and the referees for greatly improving previ-
ous versions of the manuscript. Dr. James Layne provided access to the Archbold Field
Station, Mark Deyrup pointed out the antlion population and identified the prey, and
Everett Nickerson corroborated the identification of Conomyrma. Dr. James Griffin
kindly provided the Spanish Abstract. Authors current address is Dept. of Biology,
University of Redlands, Redlands, CA 92374.

REFERENCES

BROCKMANN, H. J., AND C. J. BARNARD. 1979. Kleptoparasitism in birds. Anim.
Behav. 27: 487-514.
CONNER, J., AND T. EISNER. 1983. Capture of bombardier beetles by antlion larvae.
Psyche 90: 175-178.
DENNY, M. 1976. The physical properties of spider's silk and their role in the design
of orb-webs. J. Exp. Biol. 65: 483-506.
GRIFFITHS, D. 1980. The feeding biology of ant-lion larvae: prey capture, handling,
and utilization. J. Anim. Ecol. 49: 99-125.


709











Florida Entomologist 69(4)


HANSELL, M. H. 1984. Animal Architecture and Building Behaviour. Longman, Lon-
don. 324 p.
HEINRICH, B., AND M. J. E. HEINRICH. 1984. The pit-trapping foraging strategy
of the antlion, Mymeleon immaculatus DeGreer (Neuroptera: Myrmeleontidae).
Behav. Ecol. Sociobiol. 14: 151-160.
LUCAS, J. R. 1982. The biophysics of pit construction by antlion larvae (.ilj.. I. ..
Neuroptera). Anim. Behav. 30: 651-664.
LUCAS, J. R. 1985. Partial prey consumption by antlion larvae. Anim. Behav. 33:
945-958.
LUCAS J. R., AND H. J. BROCKMANN. 1981. Predatory interactions between ants and
antlions (Hymenoptera: Formicidae, and Neuroptera: Myrmeleontidae). J. Kan-
sas Ent. Soc. 54: 228-232.
LUCAS, J. R., AND L. A. STANGE. 1981. Key and descriptions to the Myrmeleon
larvae of Florida (Neuroptera: Myrmeleontidae). Florida Ent. 64: 207-216.
RAY, A. A. (editor). 1982. SAS User's Guide: Statistics. SAS Institute, Cary, N.C.
584 p.
WHEELER, W. M. 1930. Demons of the Dust. W. W. Norton, New York. 378 p.
WILSON, D. S. 1974. Prey capture and competition in the antlion. Biotropica 6: 187-
193.
YOUTHED, G. J., AND V. C. MORAN. 1969. Pit construction by myrmeleontid larvae.
J. Insect Physiol. 15: 867-875.




IMMATURE STAGES OF SOME WESTERN NEARCTIC
AND/OR NEOTROPICAL TABANIDAE (DIPTERA)

JAMES T. GOODWIN
Office of International Agricultural Programs,
Texas A&M University, College Station TX 77843.
Research Associate, Florida State Collection of Arthropods
Florida Dept. Agriculture and Consumer Services
Gainesville, FL 32602

ABSTRACT

Immature stages of Tabanus colombensis Macquart, T. nebulosus De Geer, and T.
tetropsis Bigot are described and illustrated. Separation of the immature stages of these
species from known immatures of American Tabanidae is discussed.

RESUME

Se described e ilustran estados inmaduros de Tabanus colembensis Macquart, T.
nebulosus De Geer, y de T. tetroposis Bigot. Se discute la separaci6n de las etapas
inmaduras de estas species de las conocidas inmaduras americanas de Tabanidae.



Goodwin and Murdoch (1974) summarized the known information on the larvae
and pupae of Neotropical Tabanidae. Information was provided on the larval and/or
pupal stages of 19 species of Tabanus, 9 of which were known only from the Neotropics
and 10 having Neotropical-Nearctic distributions. The larvae and pupae of 4, the larvae
of 1, and the pupae of 3 species were described for the first time. For the remaining


December, 1986











Goodwin: Immature Tabanus


species these authors provided translations of previously published descriptions (Cosca-
ron 1969, Coscaron and Led 1969) or brief comments and citations to existing English
language descriptions (Roberts 1962, Burger 1971). In addition to the above, descrip-
tions of the larvae and pupae of 13 species and 3 species of Tabanus exhibiting western
Nearctic and/or Neotropical distributions have been provided by Burger (1977) and
Lane (1975), respectively.
Below I give descriptions of the larvae and pupae of T. tetropsis Bigot and the larvae
only of T. nebulosus De Geer and T. colombensis Macquart. Pupae of the last 2 species
were described previously (Goodwin and Murdoch 1974). Of these 3 species, the first is
known only from the Nearctic Region, the second only from the Neotropical Region,
and the third from both regions. Their separation from other known Neotropical, Neot-
ropical-Nearctic, or western Nearctic larvae and pupae is discussed. The reader is
referred to Teskey (1969), Goodwin (1972), Tidwell (1973), Goodwin and Murdoch (1974),
Burger (1977), or Lane (1975) for illustrations of the descriptive terminology.

Tabanus colombensis Macquart

Mature larva (Fig. 1): Length 23-27 mm, whitish with contrasting brown pubescent
markings. Head capsule 3.0-3.4 mm long, greatest width 0.8-0.84 mm. Anal segment
2.0-2.2 mm long, ca. 1/5 greater than maximum width. Respiratory siphon 0.74-0.88 mm
long, ca. 1/3 greater than basal diameter; stigmatal spine absent. Striations present only
on non-pubescent areas of anal segment and laterally between pubescent extensions on
thoracic segments; spacings 0.016-0.022 mm on thoracic segments, 0.022-0.032 mm on
anal segment. Anterior pubescence absent from anal segment, elsewhere forming com-
plete annuli which are paler caudally; prothoracic annulus laterally with a single broad
fan-shaped caudal projection; meso- and metathoracic annuli each with 4 caudal projec-
tions laterally, these crossing 2/3 and 1/2 respectively of the non-pubescent areas of the
segments, those of mesothorax, especially middle 2, rather broad. Pseudopodial pubes-
cence forming complete annuli on all pseudopodial segments, united with anterior pubes-
cence ventrolaterally on all, dorsolaterally only on abdominal segments 1-3. Posterior
pubescence absent from pro- and mesothorax, elsewhere forming complete annuli which
are progressively more distinct caudally, preanal annulus expanded laterally and with
4 short anterior projections. Anal segment with an area of pubescence covering anal
lobes and ridges and extending rather broadly dorsally and somewhat anteriorly to near
the dorsal surface, this band with both midlateral and dorsolateral posterior projections,
the latter well separated from posterior annulus, the former approaching (rarely nar-
rowly uniting with) posterior annulus.
Pupae: The pupal stage has been described previously (Goodwin and Murdoch 1974).
The specimens discussed herein agree in general with the previous description.
Collections: Eight larvae, 4 reared, were collected from wet mud and grass roots at
the margins of creeks in Bexar, Dewitt and Goliad counties in Texas. Using size as the
criterion, all larvae were full grown. Collections were made in February and March.
Comments. The larva of T. colombensis would key to T. albocirculus Hine in Good-
win and Murdoch (1974). It can be separated from T. albocirculus as follows: in T.


Fig. 1. Lateral view of the larva of Tabanus colombensis.











Florida Entomologist 69(4)


albocirculus all posterior projections from the anterior mesothoracic annulus are slender
and pointed, and on the anal segment the dorsal pubescent extension from the pubes-
cence of the anal ridge curves posteriorly midlaterally, whereas in T. colombensis the
2 middle posterior projections from the anterior mesothoracic annuli are inflated and
have rounded or blunt apices, and on the anal segment the dorsal pubescent extension
from the pubescence of the anal ridges arches anterodorsally mid-laterally. The combi-
nation of the single broad fan-shaped posterior extension from the prothoracic annulus
and the presence of both a dorsolateral and a lateral posterior projection from the
vertical extension of the pubescence of the anal ridges in T. colombensis separates it
from all species treated by Burger (1977) and Lane (1975) except T. imonoensis Philip.
Separation from T. mnolensis can be made as follows: "Pseudopodial pubescence en-
circling segments 4-10 with very short, posterior extensions dorso- and ventrolaterally
on segments 4 or 5-10" and posterior pubescence "with very short anterior extensions
dorso- and ventrolaterally on segments 6 or 7-10" (Lane 1975: 817-18), whereas in T.
colombensis pseudopodial pubescence lacks any evidence of posterior extensions and
anterior extensions are absent from posterior pubescence on all segments except preanal
(10th). Although Teskey (1969) dealt with the immature stages of predominantly eastern
North American species, some of these have ranges which approach or overlap that of
T. colombensis. The same is true for some species discussed by Goodwin (1973, 1976).
However, the same 2 characters noted above to separate T. colombensis from species
treated by Burger (1977) and Lane (1975) plus the absence of a stigmatal spine in this
species will separate it from all known eastern North American species except T. lineola
Fabricius and T. reinwardtii Wiedemann. Separation of T. colombensis and T. lineola
is dealt with in Goodwin and Murdoch (1974). Tabanus colombensis can be separated
from T. reinwardtii by the fact that there are 1 or more isolated dorsolateral pubescent
spots on the anal segment in the latter species, and no such spots are present in the
former. Finally, T. colombensis can be separated from T. tetropsis (see below) by the
obvious differences in the pubescence pattern of the anal segment.
No attempt is made to provide points of separation for the pupal stages of T. colom-
bensis from those of other similar species. In their key to Tabanus pupae Goodwin and
Murdoch (1974) were unable to separate T. colombensis from T. occidentalis Linn. (as
dorsiger) var. dorsovittatus Macquart. Goodwin and Murdoch (1974) suggest that these
2 species are members of a complex including at least 6 other species with Neotropical
or Neotropical-Nearctic ranges, namely T. triangulum Wiedemann, T. subsimilis sub-
similis Bellardi, T. lineola Fabricius, T. pungens Wiedemann, T. commixtus Walker
(as truquii Bellardi), and T. claripennis (Bigot). To these 8 species can be added T.
similis Macquart based on characters of the immatures as described by Teskey (1969),
Lane (1975) and Burger (1977), and T. tetropsis Bigot described below. Comparisons of
the pupae of this group indicate that the range of variation within each species is often
greater than differences between many of the species, hence an attempt to devise a
scheme for separation of the species on the basis of pupal characters is not practical at
this time. Fairchild (1983) reviewed the status of the adults of the above and several
similar species.

Tabanus nebulosus De Geer

Mature larva (Fig. 2): Length 33-36 mm, whitish with distinct contrasting brown
pubescent pattern. Head capsule 5.27-5.35 mm long, greatest width 1.28-1.34 mm. Anal
segment 3.72-3.78 mm long, only a little longer than greatest width. Respiratory siphon
1.62-1.68 mm long, only slightly longer than broad basally; stigmatal spine absent.
Striations absent. Anterior pubescence absent from anal segment, restricted to mid-


December, 1986




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