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The
FLORIDA ENTOMOLOGIST
Volume 59, No. 2 June, 1976
CONTENTS
UTMAR. J A., AND W. W. WIRTH-.- RP'ntsion of the N'ew World Specie, 'f For-
cipomn ia. Subgenus Caloforcipomn ia I Diptera: Cerutropgonidae) 109
MNENKE. W. W, AND G. L. GREENF.-Ex.perntelntal Ialhdation ofla Pest llianagec
ment .Model 1:35
HERRING. .J. L.-KeC- to Geinera r.i A nthocoridae of .America North of M eaiia.
with a Descrnphion n'oa ,Nwc Gcnis (Hfemiptera: Heterptera) 14.3
MliCHMORE. W. B..-P.,Winlv.aoirptrns from Florida and the Caribbean Areu. 5.
Americhernes. a Nvu'w Genuc Baseld Upon Chelifer oblongus Sa\ (Cherne.
tihda) 151
BLUTI.FR, G. D. JR.. R. E. STINNER. AND G. L. GREENE .4pplhcatin of Thermon
dt\namnc .Ilo/cl fOr the De)eleopmeint of Lart ie of the Ca'bbagh L,,oper
iLepidoptera. NVectuaite) at Flut tutting Tumiperature., 165
MOCKFORD, E. I... AND H. A. E\ \N.'-l.)esriptlinon. and Record., lofq Soime Philo.
tirsitdue i t fio TI nid(idl. I .l' t I /ndit, (P.scricpleroa. 171
-)OHANY. A. L. ,N[) H. L. CHOM(OY-,Ve Ret'ord. ,s (/ Chigger, (Actanna" Train
biculidae) fi/,m Florihd 18;3
BLSCHMAN. L. I.-Int-.,1on of Florida bl the "Louechig" Plecia nearctica IDip.
tera. Bihbontdev) 191
WOJCIK, B.. \. H. W\HIICOMB. \ND D. H. HA BECK-H.i.s Rang' Tctiong o Tele-
nomus renmus ( H.~mrntlpteir: St chbonie(t.) 195
FISH. D.- In.,we t-Plant RIlation.,hip- f lthe In e tu Irrir u Pitcher, Plant Sarra-
cenia minor 199
CALKINS. C. O.. NM. D HIFTT.EL. *NI[ MK: Ko-.Spoterl/ tind Temporal Dttrl 2
bution of0' t.poiti.n by Plum ('ta, uii.,.-.Conot rachelus nenuphar 205
Dr. Mouric- I't. Prrwo.tl Rev .-it At atd ir.mi The Florkrdo Ent-,nm'lgical So-
etet\ 211
Ahllite., of the .5Tlt .4Annal llehtteng ..q Ti h Flrilta Ent tiorl oIgeal Society............ 213
S( tifcriv / Notes.
KOEHLER, P. G., AND D. E. SHORT-Pasture Mole Cricket Control with an
Applicator-Form ulated Bait .................... ................................................... 181
GLICK, J. I.-Infestation of Native Florida Cattle with the Northern Cattle
Grub, Hypoderma bovis (Diptera: Oestridae) ........................ ........... 190
(Continued on back cover)
Published by The Florida Entomological Society
~OlNGEoi~i
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This issue mailed June 30, 1976
The Florida Entomologist
A REVISION OF THE NEW WORLD SPECIES OF
FORCIPOMYIA, SUBGENUS CALOFORCIPOMYIA
(DIPTERA: CERATOPOGONIDAE)1
JOYCE A. UTMAR AND WILLIS W. WIRTH2
Medical Entomology Project, Smithsonian Institution,
Washington, D. C. 20560, and Systematic Entomology Laboratory,
IIBIII, Agr. Res. Serv., USDA, c/o U. S. National Museum,
Washington, D. C. 20560, respectively
ABSTRACT
The 11 American species of the subgenus Caloforcipomyia of the genus
Forcipomyia are described. Five new species are described and illustrated:
copanensis, hatoensis, hermosa, remigera, and sabalitensis. F. monilis
Goetghebuer, F. diversipes Goetghebuer, and F. splendid Wirth are syn-
onyms of F. glauca Macfie (N. SYNS.).
Saunders (1956) erected the subgenus Caloforcipomyia in the genus For-
cipomyia Meigen for 2 Brazilian species; F. caerulea Saunders was desig-
nated the type-species, and F. varicolor Saunders was also included, both
described in all stages. He suggested that F. auripes Ingram and Macfie and
F. pluvialis Malloch also might belong here but added that the early stages
would be more reliable criteria for determining their placement. F. auripes
is a member of the typical subgenus, and F. pluvialis belongs in the sub-
genus Metaforcipomyia Saunders (Wirth 1965).
Macfie (1934) described F. glauca from Great Britain. Later he described
3 Neotropical species, F. eukosma, F. nigrescens (1939), and F. furcifera
(1940), which belong in the subgenus Caloforcipomyia. Goetghebuer (1936)
described F. diversipes, which is a synonym of glauca (NEW SYNONYMY).
Wirth (1951) described F. splendid, which is also a synonym of F. glauca
(NEW SYNONYMY), and noted its affinities with the Neotropical fauna, es-
pecially with F. nigrescens Macfie. Tokunaga and Murachi (1959) described
F. squamianulipes and assigned it to Caloforcipomyia. Tokunaga (1962)
placed F. takahashii Tokunaga in the subgenus. Dessart (1961), working
with African species, synonymized Goetghebuer's F. rutshuruensis and F.
trimaculata under F. apicalis Goetghebuer. In 1963 he included F. apicalis,
Lasiohelea maculipes Goetghebuer, F. scitula Goetghebuer, F. auripes In-
gram and Macfie, and F. lemuria de Meillon in Caloforcipomyia. Remm
(1961) placed F. monilis Goetghebuer in the subgenus, where this species is a
synonym of F. glauca Macfie (NEW SYNONYMY). Krivosheina (1968) de-
scribed the immature stages of F. glauca (as F. monilis) and provided a key
to the subgenera based on larvae. Other species now in the subgenus Calo-
forcipomyia are F. annulipes Tokunaga (1940), F. takahashii Tokunaga
(1940), and F. swezeyi Edwards (1928).
For general morphological terminology see Wirth (1952) and Chan and
LeRoux (1965). The following special terms are used in the descriptions
and in Tables 1 and 2. Wing length is measured from the basal arculus to
'Portions of this work were supported by Research Contract No. DA-MD-17-74-C-4086 from
the U. S. Army Medical Research and Development Command, Office of the Surgeon General,
Washington, D. C.
2Research Associate, Florida State Collection of Arthropods, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services, Gainesville 32601.
Vol. 59, No. 2
The Florida Enton
TABLE 1. NUMERICAL CHARACTERS
VALUES).
ologist Vol. 59, No. 2
OF FEMALE Caloforcipomyia (MEAN
Wing Spermatheca(e)
Length/Breadth
Species Length Breadth AR TR (microns)
(mm) (mm) 1st 2nd
Caerulea Group
caerulea 0.88 0.31 1.21 1.86 57/38
hermosa 0.83 0.29 1.28 1.69 46/33
Eukosma Group
eukosma 1.65 0.45 0.99 1.54 90/65 90/65
remigera 1.29 0.42 1.27 1.27 54/42 54/42
Glauca Group
glauca 1.40 0.52 1.01 1.63 86/58 95/65
Nigrescens Group
copanensis 1.63 0.54 0.95 1.87 49/35 54/35
hatoensis 1.48 0.55 0.93 1.95 47/32 48/33
nigrescens 1.95 0.60 0.80 1.70 72/48 72/48
sabalitensis 1.53 0.55 0.94 1.50 94/68 98/72
TABLE 2. NUMERICAL CHARACTERS OF MALE Caloforcipomyia (MEAN
VALUES).
Wing
s Length Breadth
Species (mm) (mm) AR TR
(mm) (mm)
Caerulea Group
caerulea 0.89 0.27 1.45 1.85
furcifera 1.11 0.30 1.20 1.90
hermosa 1.04 0.29 1.15 1.56
Eukosma Group
eukosma 1.80 0.50 1.03 1.60
remigera 1.27 0.34 1.24 1.22
Glauca Group
glauca 1.45 0.41 1.20 1.57
Nigrescens Group
copanensis 1.85 0.50 1.29 1.79
hatoensis 1.40 0.39 1.25 1.85
nigrescens 1.95 0.50 1.12 1.60
sabalitensis 1.79 0.50 1.19 1.72
varicolor 1.65 0.55 1.04 1.36
Utmar and Wirth: New World Species of Forcipomyia 111
the wing tip. Costal ratio (CR) is the length of the costa measured from the
basal arculus divided by the wing length. The antennal proportions (AP)
are based on the length of each flagellomere (for convenience hereafter
referred to as segments although the lack of true segmentation is recog-
nized). Antennal ratio (AR) in the female is the combined length of the 5
elongated distal segments divided by the combined length of the shorter
preceding 8 segments; in the male the AR is the combined length of the 4
elongated distal segments divided by that of the preceding 9 segments. The
palpal proportions (PP) are the longitudinal proportions of the segments
of the maxillary palpus. Tarsal ratio (TR) is the length of the basitarsus
divided by the length of the second tarsomere. The female spermatheca is
measured at its greatest length and breadth. Male genitalia terminology is
that of Snodgrass (1957) after Chan and LeRoux (1965).
Measurements and other data were recorded in the manner of Chan and
LeRoux (1965). The data are presented in the following manner: mean
value (minimum value-maximum value, n=number of specimens mea-
sured). Antennal and palpal proportions are given for a typical specimen.
Most of the material studied was from the collection of the U. S. Na-
tional Museum of Natural History, Washington, D. C. Some pinned ma-
terial was examined, but most specimens were observed on microscope
slides, having been cleared and mounted in phenol-balsam after the pro-
cedure of Wirth and Marston (1968).
Particularly useful characters are the shape of the wing, female anal
cone, proximal antennal segments, shape and degree of sclerotization of
the spermathecae, and the female and male genitalia. The shape and other
features of the scales and setae which adorn the bodies of both sexes are also
diagnostic. Color patterns, especially of the wings and legs, were heavily
relied upon by early workers. These are still useful for pinned specimens,
but much of the adornment is lost in preparation of slide mounts.
In this study 11 species of the subgenus Caloforcipomyia are recognized
from the Neotropical Region, of which 5 are described as new. The Neotropi-
cal Region apparently is the site of considerable speciation in this group,
compared with other tropical areas, and many more new species of Calo-
forcipomyia undoubtedly will be discovered as more extensive collections
are made. The only species found in North America is F. glauca, which also
occurs in Europe and extends into the Neotropical Region as far south as
Brazil.
This paper was adapted from a thesis submitted by the senior author to
the University of Maryland in April 1974 for the degree of Master of Science.
She is greatly indebted to Dr. Donald H. Messersmith, Professor of Ento-
mology, for guidance and assistance in the preparation of that thesis. She
also extends thanks to Dr. John A. Davidson of the University of Maryland
for reviewing the manuscript, and to her fellow graduate students, Paul G.
Bystrak, William L. Grogan, Thomas J. Jankowski, Shahin Navai, and Wil-
liam T. Waugh for their help and encouragement.
We gratefully acknowledge the assistance of Richard Lane and authori-
ties of the British Museum (Nat. Hist.) in London and J. Antony Downes
of the Canada Department of Agriculture in Ottawa for their generous loan
of type material.
The Florida Entomologist
Subgenus Caloforcipomyia Saunders
Forcipomyia, subgenus Caloforcipomyia Saunders, 1956: 680. Type-species,
Forcipomyia caerulea Saunders, by original designation.
DIAGNOSIS (after Saunders 1956; Tokunaga and Murachi 1959).-Usu-
ally exhibiting metallic jade green or deep blue pigmentation in the sub-
cutaneous fat body, persisting from the larva through the pupa to the adult.
These pigment granules are soluble in caustic potash (Macfie 1939) but are
retained in phenol-balsam mounts and in specimens in alcohol.
ADULT.-Antenna (Fig. la) elongate; in female proximal segments
barrel shaped to elongate vasiform with short, peglike, hyaline sensilla in
addition to long, slender, hyaline sensory hairs as long as the segment
(Fig. Id); segments 11-15 elongate with long sensory hairs; verticils long.
Palpus usually slender; 3rd segment sometimes swollen or slightly en-
larged, with scattered sensillae or rarely a sensory pore (fig. Ic); 4th and
5th segments completely or partially fused. Wing unmarked or adorned
with dark and pale areas bearing dark or pale scales, which are often stri-
ated and barbed, especially prominent on costal and radial veins; costa
reaching well beyond middle of wing; CR 0.51-0.62; anterior portion of
wing (Fig. 5f) with 2nd radial cell open and elongate; 1st radial cell closed
or slitlike. Fringe of wing composed of fringed hairs (Fig. 3k). Legs with dark
or pale appressed striated scales. Tibial combs present on fore (Fig. 7e) and
hind (Fig. 7f) legs. Claws and empodium well developed in both sexes. Hind
TR 1.22-1.95. One or 2 spermathecae. Male genitalia with posterior margin
of 9th sternum transverse or with shallow caudomedian excavation. Basal
union of the claspettes in form of an inverted V or Y. No backward-directed
claspettes (or "parameres" of some authors) present.
LARVA.-Cylindrical, more elongate than usual for the genus; head
strongly hypognathous with long forward-curving p hairs; prothoracic
pseudopod a simple spinulose cushion; each normal segment bears dorsal
a hairs, dorsolateral hairs arbitrarily homologized with c hairs of sub-
genus Forcipomyia, therefore, b and d hairs considered absent, and with 2
fine lateral hairs, probably e and f. Last abdominal segment bearing a large
dorsal chitinous process extending up and back beyond cauda and anal
gills, with 2 basal and 2 long terminal stout chaetae. Cauda moderately
long and pointed without lateral fringe of short hairs. Anal blood gills 2,
bifid. Anal pseudopod normal.
PUPA.-Relatively spineless, with prominent pair of setigerous proc-
esses, conical at base, on dorsum of thorax just above and behind protho-
racic horns. Abdominal setae few, very small; larval exuviae retained on
last segments; terminal abdominal processes greatly attenuated in both
sexes; male sexual processes dorsal; prothoracic horns slender with a row
of spiracular papillae across tip.
HABITS.-Little is known about the habits and habitats of adults or
immature stages. Larvae taken on rotting wood by Saunders in the forests
of Brazil wriggled actively when disturbed, even "flicking" themselves
several inches after the manner of the cheese skipper (Piophila) (Saunders
1956). One specimen of F. glauca was reared from bamboo.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 113
KEY TO ADULTS OF THE NEW WORLD SPECIES OF Caloforcipomyia3
1. Fem ales ... .......... ..... ..... ........ ..... ... .. .............................. 2
1'. M ales ....... ...................................... 10
2. One spermatheca, usually retort shaped; smaller species,
wing less than 1 mm long caeruleaa group) ........................................ 3
2'. Two spermathecae, not retort shaped; larger species, wing
m ore than 1 m m long .......................... ....... ................................... 4
3. Body with short, broad, dark scales, especially prominent
on radial area of wing; legs dark, knees yellowish; 3rd palpal
segment only moderately enlarged, bearing long scattered
sensilla; proximal antennal segments barrel shaped ...... hermosa n. sp.
3'. Body with longer, narrow, dark scales, not so prominent on
radial area of wing; legs uniformly pale yellow; 3rd palpal
segment stout, swollen at base, bearing a large sensory area;
proximal antennal segments short, vasiform ......... caerulea Saunders
4. Mesonotum reddish to brownish, with numerous golden
scales; legs usually golden yellow; body usually with short,
broad scales; wing with prominent pattern of dark and pale
scales (eukosm a group)...................................... ........... ..................... 5
4'. Mesonotum dull grayish to golden or dark brown, without
golden scales; legs usually stramineous to brownish; body
never with short broad scales; wing without color pattern (ex-
cept faint pale areas distally in glauca) ............................................ 6
5. Wing with macrotrichia mostly yellowish, 3 patches of dark
scales forming a color pattern on anterior margin; no dark
scales on mesonotum; apex of abdomen with only golden
scales; smaller species, wing 1.29 mm long .................... remigera n. sp.
5'. Wing with numerous dark macrotrichia, with pattern of numer-
ous patches of pale macrotrichia; mesonotum with clumps of
broad black scales; larger species, wing 1.80 mm long............
................ .......................................... .......... eukosm a M acfie
6. Wing with an area of pale macrotrichia on anterior margin
past end of costa (glauca group)........................................ glauca Macfie
6'. Wing with uniformly dark macrotrichia, more prominent on
radial area (nigrescens group)....................... ... ............................... 7
7. Body dark brown; legs dark brown with yellowish knees;
wing 1.95 m m long ......................................................... nigrescens M acfie
7'. Body uniformly grayish to brownish; legs stramineous; wing
less than 1.95 m m long .......................... ....... .......................... 8
8. Spermathecae lightly sclerotized, hyaline, ovoid, 0.047 mm
lon g .................................................................................. h a toen sis n sp .
8'. Spermathacae moderately or heavily sclerotized ............................... 9
9. Spermathecae large, 0.096 mm long, broadly oval.................
...................................... .................................... .. sabalitensis n. sp.
3Females of F. furcifera Macfie and F. varicolor Saunders unknown.
The Florida Entomologist
9'. Spermathecae smaller, 0.051 mm long, pyriform with taper-
ing necks ................. ....................................... ............... cop anensis n. sp.
10. Ninth sternum with sclerotized, posteriorly directed, blade-
lik e p processes ...... ........ .............. ........... ........... ....................................... 11
10'. Ninth sternum without such processes .............................................. 12
11. Posterior margin of 9th sternum deeply excavated lateral to
2 medial bladelike processes .......................................... furcifera M acfie
11'. Posterior margin of 9th sternum only slightly excavated lat-
eral to 4 medial bladelike processes .......................... caerulea Saunders
12. Aedeagus with 2 pointed, posteriorly directed, mediolateral
processes ............................................................................... herm osa n. sp.
12'. Aedeagus without mediolateral processes ........................................ 13
13. Aedeagus narrowly triangular with recurved basal arms .................. 14
13'. Aedeagus not narrowly triangular ....................................... ............... 15
14. Telomeres broad, oarlike, with tips expanded; hind TR 1.20....
............................................... .... .... .... ................ rem ig e ra n sp .
14'. Telomeres slender; hind TR 1.60 ................................. eukosma Macfie
15. Aedeagus with slender, posteriorly produced portion, slightly
decurved at apex .................... ................. ........................... glauca M acfie
15'. Aedeagus without slender, posteriorly produced portion with
decurved apex .......... ......... .................................................. ...... .................. 16
16. Aedeagus bearing a distinct nipplelike or pointed distomedian
process borne in a pit or depression in the basal arch, the latter
v a rio u s ............................................................... ..... .. .............. 1 7
16'. Aedeagus with distomedian extremity not set off in a pit or de-
pression, the distal process inconspicuous; basal arch broad
and rounded, half the height of aedeagus .................... sabalitensis n. sp.
17. Aedeagus with basal arms short and stout, the basal arch low;
distomedian process short and broad, bearing a small median
papilla ................................... copanensis n. sp.
17'. A edeagus not as above .................................................... ......... ..... 18
18. Aedeagus with rounded basal arch, the basal arms slender or
stout; distomedian process with slender tip ............................... ..... 19
18'. Aedeagus with anterior margin transverse, the basal arms short
and stout; distomedian process with bluntly pointed tip..........
....... ...... ........... ............. ......................................... h a to en sis n sp .
19. Aedeagus with long, slender, anterior arms of basal arch; TR
1.36; W L 1.95 m m .............................. ........................ varicolor Saunders
19'. Aedeagus with short, stout, anterior arms of basal arch; TR
1.60; W L 1.65 m m ......................................................... nigrescens M acfie
Caerulea Group
Female with 1 retort-shaped spermatheca. Small species, wing less than
1 mm long. Wing uniformly dark, male genitalia with slender, bladelike
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 115
posterior processes on 9th sternum or aedeagus. Included species: caerulea
Saunders, furcifera Macfie, and hermosa n. sp.
1. Forcipomyia (Caloforcipomyia) caerulea Saunders
(Fig. 1)
Forcipomyia (Caloforcipomyia) caerulea Saunders, 1956: 681 (all stages;
Brazil; Fig.)
DIAGNOSIS.-A small species; hind TR 1.65-2.15; 3rd palpal segment
stout, swollen at base; female with proximal antennal segments stout,
irregular, vasiform; 1 pale, retort-shaped spermatheca; anal cone small,
coarsely setose; genital sclerotization broad anteriorly with a transverse
margin, posterior arms narrow, extending as dark oblique areas; male with
4 posteriorly directed, sinuate sternal processes; aedeagus with 2 hyaline,
lateromedial processes with decurved tips.
FEMALE.-Wing Length 0.88 (0.80-0.95, n=6) mm; breadth 0.31 (0.27-
0.35, n= 6) mm.
a
b e
h-i
Fig. 1. Forcipomyia caerulea: a, female antenna; b, male antenna,
last 4 segments; c, female palpus; d, female antenna, segments 10-11; e,
female wing; f, female spermatheca; g, male aedeagus; h, female genitalia;
i, male genitalia, aedeagus removed.
The Florida Entomologist
Head: Brown with long decurved hairs. Mouthparts pale. Antenna (Fig.
la) testaceous, segments 3-10 irregular, vasiform, segments 11-15 elongate;
apical papilla similar to that of F. hermosa (Fig. 2e) bearing a small sub-
terminal bidenticulate process in addition to the hyaline bifid tip. Antennal
sensoria as in Fig. Id. AR 1.21 (1.16-1.33, n=6). AP 8-7-7-8-8-8-8-10-14-14-
15-15-17. Palpus (Fig. Ic) testaceous, stout; 3rd segment swollen at base,
bearing a shallow pit with numerous clavate sensilla; segments 4 and 5
short, completely fused. PP 3-4-12-8.
Thorax: Light brown. Scutellum infused with dark pigment granules.
Pleural area yellow. Halter yellowish. Wing (Fig. le) broad, covered with
brown macrotrichia; costal region covered with narrow, lanceolate, stri-
ated, brown scales. Legs pale yellow with pale, striated scales. Tibial
combs pale. Prothoracic TR 2.35 (2.13-2.53, n=6); mesothoracic TR 2.26
(1.78-2.47, n= 6); metathoracic TR 1.86 (1.65-2.15, n= 6).
Abdomen: Brown with blue-black pigment granules and narrow, brown,
striated scales. Spermatheca (Fig. If) hyaline, retort shaped, dimensions
0.057 x 0.038 (0.040-0.069 x 0.030-0.045, n = 6) mm. Genitalia (Fig lh) with
anal cone small, coarsely setose; genital sclerotization broad anteriorly
with a transverse margin, deeply excavated posteriorly, arms narrow, with
narrow posterior arms extending as darkly sclerotized oblique areas.
MALE.-Wing length 0.89 (0.74-0.99, n = 3) mm; breadth 0.27 (0.24-0.28,
n=3) mm. Antennal plume brown. Antennal segments 12-15 as in Fig. lb.
AR 1.45 (1.28-1.66, n=3). AP 10-7-7-7-6-6-6-8-8-30-22-18-22. Prothoracic TR
2.40 (2.16-2.53, n=4); mesothoracic TR 2.29 (2.05-2.41, n=4); metathoracic
TR 1.85 (1.71-1.95, n=4). Genitalia (Fig. li) with 9th segment tapering an-
teriorly. Posterior margin of sternum slightly indented laterad of trans-
versely produced median portion bearing 2 pairs of bladelike processes,
each darkly sclerotized basally, ending in a sinuate, pointed, hyaline tip.
Posterior margin of tergum transverse, ending at the base of basimeres. Basi-
mere elongate, about 2.2 times longer than broad; telomere slender, yel-
lowish with scattered pigment granules, ending with a subapical hook.
Aedeagus (Fig. Ig) with darkly sclerotized basal arms, posteriorly pro-
duced to a narrowly transverse apex, and bearing 2 tapered, lateromedian,
hyaline processes, each ending in a rounded, decurved tip.
DISTRIBUTION.-Brazil, Colombia, Dominica, Panama.
TYPES.-Holotype, female, allotype, male, paratype, larva, and
pupal exuviae on slide, Rio de Janeiro, Brazil, 8-VII-1923, L. G. Saunders
(Canadian National Collection type no. 6504).
OTHER SPECIMENS EXAMINED.-BRAZIL: Belem, Para, VI-1970,
T. H. G. Aitken, APEG forest, light trap, 1 female. COLOMBIA: Rio Anori,
Antioquia, IX-1970, D. G. Young, tropical rain forest, black light, 1 male;
Rio Raposo, Valle, III-1964, V. H. Lee, light trap, 1 male, 28-VII-1964, 1
male. DOMINICA: Pont Casse, 12-II-1965, W. W. Wirth, rain forest, 1 male,
1 female; VI-1964, O. S. Flint, at light, 1 male, 3 females. PANAMA: Barro
Colorado Island, C. Z., VII-1967, W. W. Wirth, light trap, 2 females.
DISCUSSION.-The males of this species are readily distinguished by
the presence of 2 pairs of distinctive processes extending caudad from the
posterior margin of the 9th sternum. We have examined Saunders' types in
the Canadian National Collection in Ottawa and must point out 2 errors
in Saunders' original description. He stated that the TR was 2.33; in fact this
is the value of the TR of the anterior leg, while on the type the hind TR is
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 117
1.85. The spermatheca of the female holotype is collapsed and poorly vis-
ible, but only 1 spermatheca is present rather than 2. The type series was
reared from larvae on rotten wood in the forest.
2. Forcipomyia (Caloforcipomyia) furcifera Macfie
Forcipomyia furcifera Macfie, 1940:920 (male; Brazil; Fig. genitalia).
FEMALE.-Unknown.
MALE.-Wing length 1.11 mm; breadth 0.30 mm.
Head: Dark brown. Antennal plume pale brown. AR 1.20. AP 20-15-
14-13-12-12-12-12-12-42-35-31-35. Palpus pale brown; 3rd segment some-
what dilated, with small pit; segments 4 and 5 completely fused; terminal
segments in proportion of 20-12.
Thorax: Brown, with a bluish pigment, humeral areas paler. Halter with
white knob. Wing with narrow scales on radial area, rest of wing well
clothed with hairs. Legs pale brown, stramineous, bearing numerous broad
scales. Prothoracic TR 2.00; mesothoracic TR 2.42; metathoracic TR 1.90.
Abdomen: Dark brown with blue-black granules. Genitalia with pos-
terior margin of 9th sternum deeply excavated lateral to posteriorly pro-
duced median portion which bears 2 long bladelike processes. Basimere
dark, reniform; telomere dark, narrow, with tip slightly expanded. Ae-
deagus with curved basal arms and a broad, rounded, apical portion. Basal
union of claspettes V-shaped.
DISTRIBUTION.-Brazil.
TYPE.-Holotype, male, Nova Teutonia, Brazil, 8-V-1939, F. Plau-
mann, deposited in British Museum (Natural History), London.
DISCUSSION.-The small size and possession of sternal processes on
the male allies this species with F. caerulea. The aedeagus, however, bears
no mediolateral processes. Our redescription is based upon examination of
the holotype.
3. Forcipomyia (Caloforcipomyia) hermosa Utmar and Wirth, new species
(Fig. 2)
DIAGNOSIS.-A small brown species adorned with broad dark brown
scales. Hind TR about 1.50-1.90; 3rd palpal segment moderately en-
larged basally, bearing long clavate sensilla; female with proximal an-
tennal segments barrel shaped to subconical; 1 pale retort-shaped sper-
matheca; anal cone finely setose with 2 small apical setae; genital sclerot-
ization forming an inverted Y anteriorly; male with aedeagus bearing 2
posteriorly directed, pointed, mediolateral processes.
FEMALE.-Wing length 0.83 (0.74-1.13, n= 12) mm; breadth 0.29 (0.26-
0.31, n = 12) mm.
Head: Dark brown with blackish pigment granules. Mouthparts yel-
lowish. Antenna (Fig. 2a) pale brown with segments 3-10 barrel shaped to
elongate subconical, segments 11-14 elongate, subequal, segment 15 with
a bifid, apical papilla (Fig. 2e) having a smaller subterminal, bifid pro-
jection. Antennal sensoria as in Fig. 2d. AR 1.28 (1.18-1.38, n=3). AP 10-5-
6-6-7-7-7-10-12-12-13-13-18. Palpus (Fig. 2d) with 3rd segment moderately
enlarged basally, vasiform, bearing long clavate sensilla in shallow de-
pressions. Segments 4 and 5 completely fused. PP 4-4-14-10.
The Florida Entomologist
a
d
1 m
Fig. 2. Forcipomyia hermosa: a, female antenna; b, male antenna, last
4 segments; c, female palpus; d, female antenna, segments 10-11; e, an-
tennal papilla; f, female wing; g, female spermatheca; h, scale from ab-
domen; i, j, scales from wing; k, barbed macrotrichia from wing; 1, male
genitalia; m, female genitalia.
Thorax: Golden brown with blackish pigment granules, especially in
pleural region. Halter pale brown. Wing (Fig. 2f) broad, hyaline, with
costal region covered by dark, striated scales which are broader at wing
base (Fig. 2i,j); numerous brown, lanceolate, striated, barbed macrotrichia
on rest of wing (Fig. 2k). Legs brownish, femorotibial joints yellowish; legs
with appressed, broad, striated scales. Prothoracic TR 2.05 (1.83-2.39, n= 10);
mesothoracic TR 1.88 (1.65-2.13, n=9); metathoracic TR 1.69 (1.47-1.91,
n = 8).
Abdomen: Brown with blackish and greenish pigment granules, with
broad, striated, dark brown scales (Fig. 2h). Spermatheca (Fig. 2g) 1, hyaline,
retort-shaped, dimensions 0.046 x 0.033 (0.040-0.055 x 0.030-0.040, n= 9)
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 119
mm. Genitalia (Fig. 2m) with broad, coarsely setose anal cone bearing 2
small setae arising from tubercles near apex. Genital sclerotization of 2
posterior, oblique sclerotizations joined to a narrow Y-shaped mesal por-
tion with an expanded triangular anterior portion.
MALE.-Wing length 1.04 (0.92-1.09, n= 14) mm; breadth 0.29 (0.28-0.32,
n= 14) mm. Antennal plume golden, segments 12-15 as in Fig. 2b. AR 1.15
(1.02-1.27, n=5). AP 12-9-9-9-7-8-7-8-8-31-24-18-25. Prothoracic TR 1.85
(1.64-2.17, n=13); mesothoracic TR 1.65 (1.55-2.17, n=12); metathoracic TR
1.56 (1.48-1.76, n= 11). Genitalia (Fig. 21) with posterior margin of 9th ster-
num with median concavity; tergum with rounded posterior margin. Basi-
mere elongate, ovoid, about 1.9 times longer than broad; telomere slender,
hyaline, ending in a subapical, hooked tip. Ninth segment and basimere with
broad, dark brown, striated scales. Aedeagus complex, with a posterior,
semicircular, sclerotized plate with pigment granules, a broadly triangu-
lar body with recurved, darkly sclerotized basal arms and 2 posteriorly
directed, pointed, mediolateral processes. Basal union almost hyaline,
very slender, Y-shaped.
DISTRIBUTION.-Brazil, Colombia, Costa Rica, Jamaica, Panama.
TYPES.-Holotype, male, allotype, female, Belem, Para, Brazil, VIII,
VI (respectively) 1970, T. H. G. Aitken, APEG Forest, light trap (deposited
in Museum de Zoologia, Universidade de Sdo Paulo, Brazil). Paratypes,
26 males, 11 females, as follows: BRAZIL: same data as types, except dates
II-IX-1970, 10 males, 6 females; Rio Branquinho, Amazonas, VII-1961, E.
J. Fittkau, at light, 1 male, 1 female. COLOMBIA: Rio Anori, Antioquia, IX-
1970, D. G. Young, tropical rain forest, black light, 1 male; Rio Raposo,
Valle, II-VIII-1964, V. H. Lee, light trap, 5 males, 4 females. COSTA RICA:
Navarro, Cartago, VII-1962, F. S. Blanton, light trap, 1 male. JAMAICA:
Hardwar Gap, Hollywell, 16-VI-1970, E. G. Farnworth, light trap, 1 male.
PANAMA: Barro Colorado Island, C. Z., VII-1967, W. W. Wirth, 7 males.
DISCUSSION.-This species is closely related to F. caerulea Saunders.
The small size, relatively broad wing, and enlarged 3rd palpal segments
are characters in common. Mediolateral processes of the aedeagus are pres-
ent on the males of both species but are pointed only in F. hermosa.
ETYMOLOGY.-The specific name is derived from the Spanish: hermoso
(beautiful) and refers to the adornment of broad scales on this species.
Eukosma Group
Female with 2 spermathecae, short slender necks present. Small to
moderate-sized species. Body with numerous appressed, striated, golden
scales, especially numerous on mesonotum and abdomen where their in-
terspersion with patches of blackish scales forms a pattern for species recog-
nition. Wing with yellowish and dark scales forming a prominent pattern.
Dark scales forming a banded pattern on legs, especially on tarsi. Female
genital sclerotization forming a broad plate with rounded anterior margin
and lateral indentation. Male aedeagus narrowly triangular with recurved
basal arms and short, slender, distomedian process. Included species: eu-
kosma Macfie and remigera n. sp.
4. Forcipomyia (Caloforcipomyia) eukosma Macfie
Forcipomyia eukosma Macfie, 1939: 148 (male, female; Brazil; Fig. end
of palpus, radial cells, genitalia, of male).
The Florida Entomologist
FEMALE.-Wing length 1.65 mm; breadth 0.45 mm.
Head: Dark brown. Antenna with segments 4-10 ovoid to elongate; with-
out abrupt change of shape between segments 10 and 11; segments 11-14
elongate, subequal; segment 15 elongate, with a stylet. AR 0.99. AP 20-
16-16-18-20-20-20-20-30-30-30-26-32. Palpus dark brown, slender, without
pit. PP 5-5-15-11.
Thorax: Chestnut brown, with dark greenish granules; mesonotum with
2 sublateral clumps of long black scales on disc; scutellum dark brown.
Wings bearing narrow, dark, striated scales, and with indistinct pattern of
3 large anterior pale spots, and pale areas at periphery in apex of cell Ml,
at apices of veins M2 and M3 + 4, along vein Cul, and near base of anal
cell; wing base and area around r-m crossvein also pale. Legs yellowish,
but broad scales giving legs a banded appearance. Prothoracic TR 1.50;
mesothoracic TR 1.20; metathoracic TR 1.54.
Abdomen: Blackish, with blue-black granules and abundant long
golden scales, and with segmental patches of long black scales. Sper-
mathecae moderately sclerotized, ovoid, tapering to short slender necks;
subequal, each measuring 0.090 by 0.065 mm. Genital sclerotization as in
remigera (Fig. 31).
MALE.-Wing length 1.80 mm; breadth 0.50 mm. Antennal plume yel-
lowish brown. AR 1.03; AP 20-10-10-10-10-10-10-10-9-35-29-18-20. Pro-
thoracic TR 1.84; mesothoracic TR 1.71; metathoracic TR 1.60. Genitalia
with posterior margin of 9th sternum slightly concave mesally. Basimere
large, very dark brown, reniform, bearing almost spinelike bristles on
inner margin near base; telomere yellowish, slender, with end spatulate
and only slightly expanded. Aedeagus narrowly triangular with sclero-
tized recurved basal arms. Basal union of claspettes Y-shaped.
DISTRIBUTION.-Brazil, Colombia, Dominica.
TYPES.-Lectotype, male, allolectotype, female, Nova Teutonia,
Brazil, 19-IX-1937 and 9-VII-1938, F. Plaumann, deposited in British Mu-
seum (Nat. Hist.), London.
SPECIMENS EXAMINED.-BRAZIL: Belem, Para, II, VI-1970, T. H.
G. Aitken, light trap, 2 males. Estrada Rio-Sao Paulo KM 47, 1-II-1944,
P. Wygodzinsky, 1 male (Univ. Sao Paulo); Sao Paulo, H. Florestal, V-
1955, J. Lane, 1 female (Univ. Sao Paulo). COLOMBIA: Rio Raposo, Valle,
II-VIII-1964-65, V. H. Lee, light trap, 6 males, 3 females. DOMINICA:
Clarke Hall, VI-1964, O. S. Flint, light trap, 1 male; I-III-1965, W. W.
Wirth, light trap, 15 males, 6 females. d'Leau Gommier, 16-III-1965, W. W.
Wirth, light trap, 2 males. Fond Figues River, I-III-1965, W. W. Wirth,
12 males, 4 females. Manets Gutter, 10-III-1965, W. W. Wirth, light trap,
3 males, 2 females. South Chiltern Estate, 20-II-1965, W. W. Wirth, 1 male.
DISCUSSION.-Forcipomyia eukosma is a larger species than remigera
and is easily recognized by the dark wing with numerous patches of whitish
scales, the sublateral pair of prominent tufts of long black scales on the
mesonotum, and the alternating areas of long black and golden scales on
the abdomen.
We have examined the syntype male (on slide) which we hereby desig-
nate lectotype, and the pinned syntype female which we designate allolec-
totype.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 121
5. Forcipomyia (Caloforcipomyia) remigera Utmar and Wirth, new species
(Fig. 3)
DIAGNOSIS.-A small, golden-yellow species with pale legs. Hind
TR 1.11-1.37; third palpal segment slender, vasiform; female with proxi-
mal antennal segments subconical; 2 equal, lightly sclerotized, globose
spermathecae; anal cone slender, finely setose; genital sclerotization dis-
tinct, with a semicircular anterior portion, constricted mesally and con-
cave posteriorly, with darkly sclerotized posterior arms; male with tips
of telomeres expanded with a blunt subapical point on inner margin.
a
b
*d
7 f
0Q
Fig. 3. Forcipomyia remigera: a, female antenna; b, male antenna,
last 4 segments; c, female palpus; d, antennal papilla; e, female antenna,
segments 10-11; f, female wing; g, female spermathecae; h, scale from tibia;
i, scale from tarsus; j, scale from abdomen; k, barbed macrotrichia from
wing fringe; 1, female genitalia; m, male genitalia.
(D
The Florida Entomologist
FEMALE.-Wing length 1.29 (1.13-1.40, n= 15) mm; breadth 0.42 (0.36-
0.47, n= 15) mm.
Head: Light brown with many short erect hyaline scales with decurved
tips; antenna and mouthparts paler. Antenna (Fig. 3a) with segments 3-10
subconical, gradually becoming elongate, each slightly paler at base; seg-
ments 11-14 elongate, subequal; segment 15 elongate, ending in a slender,
bifid stylet with a smaller subterminal bifid projection (Fig. 3d). Antennal
sensoria as in Fig. 3e. AR 1.27 (1.03-1.47, n=10). AP 10-9-10-10-11-11-11-12-
20-22-22-20-28. Palpus (Fig. 3c) with 3rd segment elongate, slightly en-
larged at basal 1/3, bearing numerous sensilla in shallow depressions,
segments 4 and 5 completely fused. PP 5-6-18-14.
Thorax: Golden brown with hyaline, striated, barbed scales; pleural
area yellowish, often with bright green pigment granules. Wing (Fig. 3f)
narrow, with 3 patches of pale, striated, barbed scales on radial area, alter-
nating with 3 smaller dark brown patches; numerous dark, lanceolate,
striated, barbed macrotrichia (Fig. 2k) on rest of wing. Legs pale yellowish
with pale appressed striated scales (Fig. 3h). Tarsus with anteromedial
patch of dark brown, striated, appressed scales (Fig. 3i). Prothoracic TR
1.58 (1.38-1.79, n= 15); mesothoracic TR 1.29 (1.11-1.42, n= 15); metathoracic
TR 1.27 (1.11-1.40, n = 15).
Abdomen: Dark brown to greenish brown with erect, golden, striated,
lanceolate and expanded scales as in Fig. 3j. Spermathecae (Fig. 3g) 2,
equal, subglobose to globose, lightly sclerotized, dimensions each 0.054 x
0.042 (0.045-0.074 x 0.035-0.048, n=ll) mm. Genitalia (Fig. 31) with anal
cone small, finely setose. Genital sclerotization hyaline but distinct, semi-
circular anteriorly, constricted mesally and concave posteriorly, form-
ing a darkly sclerotized, recurved portion which extends to bases of the
cerci.
MALE.-Wing length 1.27 (1.01-1.38, n=16) mm; breadth 0.34 (0.27-0.39,
n=16) mm. Antenna with golden plume; segments 12-15 as in Fig. 3b. AR
1.24 (1.13-1.33, n=9). AP 15-9-9-8-8-8-9-9-9-40-30-18-23. Prothoracic TR
1.58 (1.42-1.77, n=15); mesothoracic TR 1.24 (1.12-1.35, n= 15); metathoracic
TR 1.22 (1.09-1.35, n=15). Genitalia as in Fig. 3m. Ninth segment with a
short, narrow, anterior neck; sternum with a shallow, caudomedian exca-
vation; tergum rounded. Basimere reniform with striated, barbed scales and
long setae, about 1.8 times longer than broad; telomere broad, expanded
distally with blunt distomesal point. Aedeagus narrowly triangular with
darkly sclerotized, recurved basal arms. Basal union of claspettes Y-
shaped.
DISTRIBUTION.-Brazil, Colombia, French Guiana, Panama.
TYPES.-Holotype, male, allotype, female, Rio Raposo, Valle, Co-
lombia, VIII-1964, II-1965, respectively, V. H. Lee, light trap (type no.
72212, USNM). Paratypes, 90 males, 17 females, as follows: BRAZIL: Para,
Belem, APEG Forest, T. H. G. Aitken, light trap, III-XI-1970, 28 males, 2
females; sticky trap, VII-1970, 1 female. COLOMBIA: Buenaventura, 7-8-X-
1964, V. H. Lee, light trap, 1 female; Rio Anori, Antioquia, IX-1970, D. G.
Young, tropical rain forest, light trap, 5 males, 1 female; Rio Truando,
Camp Teresita, XI-XII-1967, D. G. Young, 1 male, 1 female; Rio Raposo,
Valle, V. H. Lee, light trap, II-XII-1964-1965, 29 males, 8 females. FRENCH
GUIANA: Cabasson, II-1965, T. H. G. Aitken, light trap, 1 male. PANAMA:
Barro Colorado Island, C. Z., VII-1967, W. W. Wirth, light trap, 26 males,
3 females.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 123
DISCUSSION.-This species closely resembles F. eukosma Macfie but
is a smaller species. The tarsal segments bear only an anteromedial patch
of dark scales and do not appear banded. The aedeagus is similar to that of
eukosma, but the base is not as broad. The telomeres of remigera are broader
and bear a blunt distomesal point, not spatulate as in Macfie's figure of
eukosma.
Externally, F. remigera can be recognized quite readily by the color
pattern of scales on the wing, mesonotum, and abdomen. The wing scales
form a pattern consisting of a pale yellow background with 3 anterior
clumps of larger, dark brown scales. The expanded scales on the mesono-
tum and abdomen are golden yellow only, not dark brown. The male ab-
domen has only golden yellow scales on segments 7, 8, and 9, with a few
very broad, dark brown scales on the bases of the basistyles.
ETYMOLOGY.-The species name is derived from the Latin: remus (oar)
and gerere (to bear) and refers to the expanded telomeres of the male.
Glauca Group
Moderately large species, wing 1.2-1.6 mm long. Body with dark, ex-
panded, striated scales. Femora and tibiae with broad, dark banding. Wing
dark scaled, with one or more large areas of pale scales. Female with 2
spermathecae, duct opening large, neck absent; female anal cone with sev-
eral large setae; genital sclerotization indistinct with sclerotized basal
arms. Male aedeagus with relatively narrow, high basal arch and long,
slender, decurved, distomedian process. One described species: glauca Mac-
fie.
6. Forcipomyia (Caloforcipomyia) glauca Macfie
(Fig. 4)
Forcipomyia glauca Macfie, 1934 (June): 144 (male; England).
Forcipomyia monilis Goetghebuer, 1934 (Aug.): 287 (male; Russia, Fig. ).
NEW SYNONYMY.
Forcipomyia (Caloforcipomyia) monilis Goetghebuer; Remm, 1961: 178
(Estonia; Figs.); Krivosheina, 1968: 58 (USSR; larva, pupa descr., Fig.).
Forcipomyia diversipes Goetghebuer, 1936: 319 (male; Belgium). NEW
SYNONYMY.
Forcipomyia splendid Wirth, 1951: 315 (male, female; Virginia; Fig. male
gen.). NEW SYNONYMY.
Forcipomyia (Caloforcipomyia) splendid Wirth; Wirth, 1965; 125 (status;
distr.).
DIAGNOSIS.-A dull brown species; legs with broad brownish band-
ing. Hind TR 1.37-1.75; palpus slender; female with proximal antennal
segments regular, vasiform; 2 large, pale, unequal, ovoid spermathecae;
anal cone with 2 large, widely separated, medial setae; male aedeagus nar-
rowly produced posteriorly as a blade with a rounded, decurved tip.
FEMALE.-Wing length 1.40 (1.22-1.55, n= 15) mm; breadth 0.52 (0.47-
0.63, n= 15) mm.
Head: Dark brown with long, erect, expanded, hyaline hairs with de-
curved tips. Antenna (Fig. 4a) testaceous, with segments 3-10 regular vasi-
form; segments 11-14 subequal, elongate, segment 15 elongate with apical
papilla (Fig. 4g) bearing 2 tiny teeth and smaller subterminal projection.
The Florida Entomologist
Antennal sensoria as in Fig. 4d. AR 1.01 (0.92-1.08, n = 16). AP 14-13-13-14-
13-13-13-14-19-20-20-21-27. Palpus (Fig. 4c) testaceous with 3rd segment
elongate, slightly enlarged medially, bearing numerous elongate scat-
tered sensilla, segments 4 and 5 partially fused. PP 8-8-22-16.
Thorax: Dull brown. Wing (Fig. 4e) with dark, lanceolate, striated,
barbed scales on radial area, a patch of whitish, striated macrotrichia prox-
imad to the radial patch; brown, narrow, striated macrotrichia covering
rest of wing. Legs yellowish, femora and tibiae broadly brown in midpor-
tion; tarsi with broad bands due to broad brown, striated, barbed scales,
apices pale. Prothoracic TR 1.98 (1.83-2.14, n= 16); mesothoracic TR 2.00
(1.63-2.50, n= 16); metathoracic TR 1.63 (1.45-2.09, n=16).
Abdomen: Brownish green, often with clumps of blue-black pigment
granules; with semi-appressed, lanceolate, striated scales (Fig. 4h). Sper-
a
b
- C~
A3 ~ s
=-e
h
f
Fig. 4. Forcipomyia glauca: a, female antenna; b, male antenna, last
4 segments; c, female palpus; d, female antenna, segments 10-11; e, female
wing; f, female spermathecae; g, antennal papilla; h, scales from abdomen;
i, female genitalia; j, male genitalia.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 125
mathecae (Fig. 4f) 2, large, ovoid, unequal and hyaline, dimensions 0.095 x
0.065 (0.075-0.110 x 0.055-0.080, n=13) mm and 0.086 x 0.058 (0.070-0.095
x 0.040-0.070, n= 13) mm. Genitalia (Fig. 4i) with prominent anal cone bear-
ing 2 widely placed medial setae in addition to some shorter anterior setae.
Posterior margin of 9th sternum doubly sinuate. Genital sclerotization
indistinct, with sclerotized basal arms.
MALE.-Wing length 1.45 (1.21-1.65, n=20) mm; breadth 0.41 (0.39-0.46,
n = 19) mm. Antenna with plume dark brown basally and apical third yel-
low, segments 12-15 as in Fig. 4b. AR 1.20 (1.14-1.35, n=8). AP 15-9-9-9-9-
8-9-10-10-41-27-24-27. Prothoracic TR 2.02 (1.86-2.33, n=19); mesothoracic
TR 1.98 (1.66-2.34, n=20); metathoracic TR 1.57 (1.36-1.75, n=20). Geni-
talia as in Fig. 4j. Posterior margin of 9th sternum with shallow mesal ex-
cavation. Basimere elongate ovoid, about 2 times longer than broad; telo-
mere slender, with a small subapical hook. Aedeagus with heavily sclero-
tized, recurved basal arms and a long, slender, posteriorly produced,
sclerotized distomedian process ending in a rounded, slightly decurved tip.
Basal union of claspettes darkly sclerotized, slender, U- to V-shaped.
DISTRIBUTION.-Northern Europe, North America from Alberta and
Arkansas to Nova Scotia and Florida, West Indies, Mexico and Central
America through South America to Brazil.
TYPES.-Holotype, male, of glauca, from England, Norfolk, Hickling
Broad, VIII-1932, F. W. Edwards, in British Museum (Nat. Hist.), London.
Holotype, male, of rnonilis, Leningrad, USSR (coll. Stackelberg, pre-
sumably in Stackelberg collection in Leningrad). Holotype, male, of
diversipes, Chiny, Belgium, 22-VII-1936, M. Goetghebuer, in Brussels Mus.
(seen by Wirth in 1973, is a female). Holotype, male, of splendid, Mount
Solon, Virginia, 13-VII-1950, W. W. Wirth, light trap (type no. 60969,
USNM).
SPECIMENS EXAMINED.-660 males and females, from the follow-
ing: CANADA: Alberta, Manitoba, Nova Scotia, Ontario, Quebec. U.S.A.:
Alabama, Arkansas, Connecticut, Florida, Georgia, Louisiana, Maine,
Maryland, Massachusetts, Michigan, Mississippi, New Hampshire, New
York, North Carolina, Pennsylvania, Rhode Island, South Carolina,
Virginia, West Virginia, Wisconsin. WEST INDIES: Jamaica, Puerto Rico,
Trinidad. MEXICO: Chiapas, Nayarit, Oaxaca, Veracruz. CENTRAL AMER-
ICA: British Honduras, Costa Rica, El Salvador, Guatemala, Honduras,
Panama. SOUTH AMERICA: Brazil, Colombia.
DISCUSSION.-Wirth has examined the holotypes of F. glauca and F.
diversipes, and we have compared European material, including specimens
determined by Remm as F. monilis, with the holotype of F. splendid. We
believe that only 1 species is involved in Europe and North America, for
which the name glauca has priority, and that this species ranges into the Neo-
tropical Region as far south as southern Brazil.
Nigrescens Group
Moderate to large species, wing 1.48-2.00 mm long. Body with numer-
ous, long, bristlelike setae; broad scales present only on legs, especially
the tarsi. Wing and legs without prominent color pattern. Female with 2
spermathecae; female anal cone with 2 long setae, genital sclerotization
indistinct with 2 small, oblique, sclerotized areas. Male aedeagus with
The Florida Entomologist
low, broad, basal arch, and short, variously shaped, distomedian process.
Five included species: copanensis n. sp., hatoensis n. sp., nigrescens Macfie,
sabalitensis n. sp., and varicolor Saunders.
7. Forcipomyia (Caloforcipomyia) nigrescens Macfie
Forcipomyia nigrescens Macfie, 1939: 156 (male, female; Brazil; Fig.
palpus, antennal segments 10 and 11, spermathecae, radial cells, male
gen.).
FEMALE.-Wing length 1.95 mm; breadth 0.60 mm.
Head: Antenna dark or darkish brown, segments 4-10 long, vasiform,
subequal but narrowing distally; segments 11-14 slightly elongate, sub-
equal; segment 15 elongate, with stylet. AR 0.80. AP 12-12-12-12-12-12-
12-12-16-15-15-15-18. Palpus darkish brown, slender, last 3 segments in pro-
portion of 15-20-40.
Thorax: Dark brown with masses of blue-black granules. Wing with
narrow scales on radial area, well clothed with hairs elsewhere. Legs dark
brown with yellowish knees; bearing dark scales; tarsus dark with color
due to scales. Prothoracic TR 2.00; mesothoracic TR 1.90; metathoracic
TR 1.70.
Abdomen: Blackish with blue-black granules, densely clothed with
scales and bristles. Spermathecae highly sclerotized, ovoid but tapering
toward duct, dimensions of each 0.072 x 0.048 mm, duct sclerotized about
0.004 mm at its commencement. Genitalia as in sabalitensis n. sp.
MALE.-Wing length 1.95 mm, breadth 0.50 mm. Antennal plume dark
brown. AR 1.12. AP 12-10-10-11-11-11-11-11-40-31-17-21. Prothoracic TR
1.57; mesothoracic TR 1.60; metathoracic TR 1.60. Genitalia with margin
of 9th sternum concave; 9th tergum short. Basimere very dark or blackish;
telomere yellow brown with tip only slightly expanded. Aedeagus in form
of a wide arch, with a densely sclerotized, sharp-pointed tip.
DISTRIBUTION.-Brazil, Panama.
TYPES.-The following syntypes in the British Museum (Nat. Hist.),
London: Brazil, Nova Teutonia, 30-VIII-1937, F. Plaumann, 1 male, 1 fe-
male; 11-V-1937, 1 female; 1-IX-1937, 1 female. We have borrowed these
for study and hereby select the male (on slide) as lectotype.
SPECIMENS EXAMINED.-BRAZIL: Nova Teutonia, types as listed
above; X-1962, F. Plaumann, 3 females (pinned, USNM). SAo Paulo, Embu,
V-1955, M. Campos, 1 male. Barueri, 4-VII-1955, K. Lenko, 1 male. Juquia,
VII-1947, J. Lane, 1 male. Rio de Janeiro, Angra, XI-1945, J. Lane, 1 fe-
male. PANAMA: Gamboa, C. Z., Pipeline Road, VII-1967, W. W. Wirth,
malaise trap, 3 females.
DISCUSSION.-This species has the largest wings of the Neotropical
Caloforcipomyia. The aedeagus is also diagnostic. Macfie notes that 1 fe-
male specimen he examined was smaller than the others with a wing length
of 1.70 mm and large spermathecae measuring 0.080 X 0.057 mm. This speci-
men is probably F. sabalitensis n. sp.
8. Forcipomyia (Caloforcipomyia) varicolor Saunders
Forcipomyia (Caloforcipomyia) varicolor Saunders, 1956:683 (all stages;
Brazil; fig.).
FEMALE.-Unknown.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 127
MALE.-Wing length 1.65 mm; breadth 0.55 mm.
Palpus slender with 3rd segment slightly enlarged near base, bearing
circular sensory area with cluster of stalked sensilla; segments 4 and 5
partially fused, but distinct. AR 1.04. AP 15-10-10-10-10-10-10-10-10-40-23-
16-20. Wing narrow, densely hairy on anterior portion, clothed with fine
macrotrichia elsewhere. Legs pale, testaceous. Prothoracic TR 1.80; meso-
thoracic TR 1.52; metathoracic TR 1.36. Genitalia with 9th segment longer
than wide, with short, narrow, anterior neck. Basimere elongate, bearing
squamous setae; telomere slender, slightly thinner medially. Aedeagus
strongly sclerotized, short, broad, V-shaped with posterior slender median
point marked off by hollowed socket. Basal union of claspettes slender,
V-shaped.
DISTRIBUTION.-Brazil.
TYPES.-Holotype, male, and associated pupal exuviae; paratype, 1
larva, Rio de Janeiro, Brazil, 1-VIII-1923, L. G. Saunders, reared on moist
rotting wood, deposited in Canadian National Collection, Ottawa.
DISCUSSION.-Our redescription is from the type, borrowed from
Ottawa. The specimen is in poor condition, wing poorly visible and lack-
ing hind legs. The hind TR measurement was taken from Saunders' original
sketches and notes which are in the files of the USNM. This species closely
resembles F. nigrescens Macfie, differing mainly in its smaller size, lower
TR, and in the shape of the male aedeagus, which shows the heavily sclero-
tized distomedian point of nigrescens, but the basal arms of the arch are
much more slender.
9. Forcipomyia (Caloforcipomyia) sabalitensis Utmar and Wirth,
new species
(Fig. 5)
DIAGNOSIS.-A large, unmarked, yellowish-brown species. Hind TR
1.40-2.13; palpus slender; female with proximal antennal segments elon-
gate, vasiform; 2 large, dark, broadly oval spermathecae; anal cone dark,
bearing 2 long, apical setae; genital sclerotization indistinct, with 2 oblique
sclerotized areas of the posterior arms; male aedeagus with wide arch.
FEMALE.-Wing length 1.53 (1.49-1.58, n=7); breadth 0.55 (0.54-0.57,
n=7) mm.
Head: Vertex dark brown with decurved hyaline hairs. Proboscis long,
testaceous, with dark brown medial band. Antenna (Fig. 5a) with segments
3-10 elongate, vasiform; segments 11-14 elongate, subequal, each paler at
apex; segment 15 having apical papilla similar to that of F. glauca (Fig.
4g), with small bidenticulate tip. Antennal sensoria as in Fig. 5d. AR 0.94
(0.86-1.14, n=8). AP 13-12-13-14-14-13-14-14-20-18-18-18-23. Palpus (Fig.
5c) slender, dark, strongly sclerotized; 3rd palpal segment elongate, bear-
ing scattered clavate sensilla each arising from a shallow depression. Seg-
ments 4 and 5 completely fused. PP 7-7-22-19.
Thorax: Dark brown, scutellum lighter. Halter grayish white. Wing
(Fig. 5e) narrow, hyaline; radial area (Fig. 5f) with dark, lanceolate, stri-
ated scales; brown macrotrichia on rest of wing. Legs uniformly yellowish,
bearing pale, striated scales. Prothoracic TR 1.77 (1.57-2.06, n=9); meso-
thoracic TR 1.85 (1.73-2.08, n=9); metathoracic TR 1.50 (1.40-1.66, n= 8).
Abdomen: Dark brownish with blackish pigment granules; with numer-
The Florida Entomologist
ous long yellowish hairs, without broadened scales; hairs becoming bristle-
like at apex of abdomen. Spermathecae (Fig. 5g) 2, large, dark, broadly
oval; dimensions 0.094 x 0.068 (0.090-0.110 x 0.055-0.080, n=8) mm and
0.098 x 0.072 (0.090-0.110 x 0.060-0.085, n=8) mm. Genitalia as in F. co-
panensis (Fig. 6g) with large anal cone bearing 2 long setae near apex and
several shorter, medial setae, genital sclerotization indistinct, with 2 small
oblique sclerotized areas of the posterior arms.
MALE.-Wing length 1.79 (1.66-1.95, n=9) mm; breadth 0.50 (0.49-0.54,
n = 7) mm. Antenna with dark plume; segments 12-15 as in Fig. 5b. AR 1.19
(1.05-1.29, n =8). AP 17-11-11-10-10-11-11-12-11-52-27-23-23. Prothoracic TR
2.02 (1.77-2.53, n=9); mesothoracic TR 1.93 (1.63-2.36, n=9); metathoracic
TR 1.72 (1.40-2.12, n=9). Male genitalia as in Fig. 5h. Posterior margin of
9th sternum with wide, shallow excavation. Posterior margin of 9th tergum
sinuate. Basimere elongate, darkly sclerotized with many large setae aris-
ing from tubercles, about 2.2 times longer than broad; telomere dark at
base becoming yellowish and slightly expanded medially, and ending with
a hyaline, curved, hooked tip. Aedeagus a low, wide arch half the height of
aedeagus, with darkly sclerotized, recurved basal arms, and very short, in-
conspicuous, decurved distomedian process (shape appears various if not
a
b c
talia.
de
Fig. 5. Forcipomyia sabalitensis: a, female antenna, b, male antenna,
last 4 segments; c, female palpus; d, female antenna, segments 10-11; e,
female wing; f, radial field of wing; g, female spermathecae; h, male geni-
talia.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 129
viewed in perpendicular profile). Basal union of claspettes darkly sclero-
tized, in form of a shallowly notched V.
DISTRIBUTION.-Brazil, Colombia, Costa Rica, Honduras, Panama.
TYPES.-Holotype, male, allotype, female, Sabalito, Puntarenas,
Costa Rica, VIII-1952, F. S. Blanton (type no. 72215, USNM). Paratypes,
46 males, 43 females, as follows: BRAZIL: Nova Teutonia, Santa Catarina,
VIII-IX-1961, 1963, 1964, 1970, F. Plaumann, 4 males, 1 female; Rio Negro,
Parana, 19-VI-1946, D. Franciscanos, 1 male. COLOMBIA: Rio Truando,
Camp Teresita, Antioquia, 9-VII-1967, D. G. Young, 2 females. COSTA RICA:
Same data as type, 39 males, 35 females; Navarro, Cartago, VII-1962, F. S.
Blanton, 1 male, 1 female; San Vito de Java, Puntarenas, VII-1964, F. S.
Blanton, light trap, 3 females. HONDURAS: Santa Rosa, Copan, II-1964,
F. S. Blanton, 1 female. PANAMA: El Volcan, Chiriqui, 22-VII-1966, A.
Broce, 1 male.
OTHER SPECIMENS EXAMINED.-BRAzIL: Sao Paulo, Batea, V-
1940, and Boracea, VII-1949, J. Lane, 1 male, 2 females; Rio de Janeiro,
D. F., II-1940, 1 female; Km 47, Estrada Rio-Sdo Paulo, Mun. Itaguay, P.
Wygodzinsky coll., 1 female; Salesopolis (Boracea) 4-VIII-1947, Rabello,
Travassos and Lane, 1 male, 1 female.
DISCUSSION.-This large, slender, poorly marked species is closely
related to F. nigrescens Macfie, copanensis n. sp., and hatoensis n. sp. It can
be distinguished by the large, darkly sclerotized, broadly oval female sper-
mathecae. The aedeagus, in the form of a wide arch with short, inconspicuous
distal point, distinguishes it from other species of Caloforcipomyia.
10. Forcipomyia (Caloforcipomyia) copanensis Utmar and Wirth,
new species
(Fig. 6)
DIAGNOSIS.-A large, unicolorous brownish species. Hind TR 1.66-
2.20; palpus slender; female with proximal antennal segments irregular,
vasiform; 2 dark, pyriform spermathecae; anal cone with 2 long, terminal
setae; male with aedeagus broadly wedge-shaped, distally truncate, with a
small, median papilla.
FEMALE.-Wing length 1.63 (1.51-1.76, n=4) mm; breadth 0.54 (0.50-
0.58, n= 4) mm.
Head: Brown, proboscis dark with darker median band. Antenna (Fig.
6a) brown; segments 3-10 elongate, irregular, vasiform; segments 10-14
elongate, subequal; segment 15 elongate, terminating in a papilla similar
to that of F. glauca (Fig. 4g) with 2 tiny apical teeth. Antennal sensoria as
in Fig. 6d. AR. 0.95 (0.91-0.97, n= 4). AP 15-14-15-14-15-14-14-16-22-21-22-22-
26. Palpus (Fig. 6c) slender, darkly sclerotized; 3rd segment slightly en-
larged submedially; segments 4 and 5 partially fused. PP 8-9-25-24.
Thorax: Brown. Halter yellowish. Wing (Fig. 6f) narrow with narrow,
lanceolate scales; fine brown macrotrichia on rest of wing. Legs dark yel-
low. Tibia appears darker distally due to dark scales and tibial combs.
Tarsus testaceous with numerous, striated, barbed scales. Prothoracic TR
2.06 (1.79-2.20, n=4); mesothoracic TR 2.16 (1.82-2.36, n=4); metathoracic
TR 1.87 (1.63-2.20, n=4).
Abdomen: Dark brown, pilose. Spermathecae (Fig. 6e). 2 subequal
darkly sclerotized, pyriform, dimensions 0.049 X 0.035 (0.045-0.050 X 0.035,
130 The Florida Entomologist Vol. 59, No. 2
n=4) mm. and 0.054 X 0.035 (0.045-0.060 X 0.035, n=4) mm. Genitalia (Fig.
6g) with anal cone large, finely pilose, bearing 2 long setae arising from
tubercles near apex, and about 4 small medial setae. Genital sclerotiza-
tion indistinct, hyaline, with only 2 posterior, oblique, dark sclerotized
areas.
MALE.-Wing length 1.85 (1.78-1.93, n=3) mm; breadth 0.50 (0.49-0.51,
n= 3) mm. Antennal plume dark, segments 12-15 of antenna as in Fig. 6b.
AR. 1.29 (n= 1). AP 13-12-12-12-13-13-12-13-13-61-32-24-29. Prothoracic TR
2.07 (1.97-2.17, n=3); mesothoracic TR 2.23 (2.09-2.44, n=3); metathoracic
TR 1.79 (1.66-1.88, n=3). Genitalia (Fig. 6h) with 9th segment slightly con-
stricted near base. Posterior margin of sternum with shallow median ex-
cavation. Ninth tergum with transverse posterior margin. Basimere dark,
elongate ovoid, about 2.2 times as long as broad, with stout setae and very
long ones. Telomere darkly sclerotized basally, slightly expanded api-
cally and ending in a strong subapical hook. Aedeagus arched with large,
darkly sclerotized, recurved basal arms, posterior portion broadly wedge
shaped with distal end truncate, terminating in small, median papilla.
Basal union darkly sclerotized, Y-shaped, wide at base.
a
b
^-rs-
g
Fig. 6. Forcipomyia copanensis: a, female antenna; b, male antenna,
last 4 segments; c, female palpus; d, female antenna, segments 10-11; e, fe-
male spermathecae; f, female wing; g, female genitalia; h, male genitalia.
Utmar and Wirth: New World Species of Forcipomyia 131
DISTRIBUTION.-Costa Rica, Honduras.
TYPES.-Holotype, male, Santa Rosa, Copan, Honduras, II-1964, F. S.
Blanton (type no. 72214, USNM). Allotype, female, Sabalito, Puntarenas,
Costa Rica, VIII-1952, F. S. Blanton, light trap. Paratypes, 2 males, 3 fe-
males, as follows: COSTA RICA: same data as allotype, 1 female; Turri-
alba, Cartago, 15-19-VII-1965, P. J. Spangler, malaise trap, 1 female. HON-
DURAS: same data as holotype, 1 male, 1 female; Siguatepeque, Comayagua,
111-1963, F. S. Blanton, 1 male.
DISCUSSION.-The anal cone of the female of this species is similar to
that of F. sabalitensis and F. hatoensis. The 2 darkly sclerotized, pyriform
spermathecae will distinguish F. copanensis from these species. This species
is smaller than F. nigrescens Macfie, which has dark tapering spermathe-
cae. The structure of the aedeagus is also diagnostic.
11. Forcipomyia (Caloforcipomyia) hatoensis Utmar and Wirth,
new species
(Fig. 7)
DIAGNOSIS.-A small species with golden yellow thorax and legs,
thorax and abdomen without broad scales. Hind TR 1.68-2.00; palpus
slender; female with proximal antennal segments elongate, irregular,
vasiform; 2 pale ovoid spermathecae; anal cone with 2 long apical setae;
genital sclerotization indistinct; male aedeagus rectangular, bearing a
triangular apex arising from a mesal semicircular socket.
FEMALE.-Wing length 1.48 (1.39-1.55, n=3) mm; breadth 0.55 (0.49-
0.58, n= 3) mm.
Head: Brown, with erect hyaline scales with decurved tips. Antenna
(Fig. 7a) with segments 3-10 elongate, irregular, vasiform, segments 11-14
elongate, subequal; segment 15 alongate with a bidenticulate apical pa-
pilla as in F. glauca (Fig. 4g). Antennal sensoria as in Fig. 7d. AR 0.93
(0.88-0.96, n=3). AP 15-12-13-13-14-13-14-15-19-20-19-19-24. Palpus (Fig.
7c) slender, segment 3 without definite sensory pit; segments 4 and 5 par-
tially fused. PP 7-8-19-18.
Thorax: Golden brown with scattered green pigment granules. Wing
similar to that of F. sabalitensis (Fig. 5e). Legs pale yellow, with moder-
ately broad, appressed, striated scales, tarsus appearing darker. Prothoracic
TR 2.13 (1.95-2.20, n=3); mesothoracic TR 2.48 (2.27-2.57, n=3); meta-
thoracic TR 1.95 (1.68-2.13, n =3).
Abdomen: Grayish green with dark blue pigment granules; finely pilose.
Spermathecae (Fig. 7g) pale, subequal, ovoid with short slender necks; di-
mensions 0.048 X 0.033 and 0.047 X 0.032 (0.040-0.050 x 0.030-0.035, n=3)
mm. Genitalia as in F. copanensis (Fig. 6g), anal cone dark with 2 long
apical setae; genital sclerotization indistinct, with 2 dark oblique posterior
areas.
MALE.-Wing length 1.40 (1.26-1.50, n=6) mm; breadth 0.39 (0.36-0.42,
n=6) mm. Antennal plume dark, segments 12-15 of antenna as in Fig. 7b.
AR 1.25 (1.12-1.33, n=4). AP 19-10-10-10-10-9-9-10-12-44-31-23-27. Tibial
combs of fore and hind legs as in Fig. 7e and 7f. Prothoracic TR 2.08 (1.83-
2.23, n=6); mesothoracic TR 2.16 (2.08-2.32, n=6); metathoracic TR 1.85
(1.71-1.95, n= 6). Genitalia (Fig. 7h) with 9th segment slightly indented near
base; 9th sternum with posterior margin transverse. Basimere dark, elon-
The Florida Entomologist
gate, about 2.5 times longer than wide, bearing stout setae in addition to the
usual long setae arising from tubercles; telomere slender, yellowish,
darkly sclerotized basally, with a strong subapical hook. Aedeagus darkly
sclerotized, anterior margin transverse, basal arms short and stout, disto-
median process triangular, bluntly pointed, arising from a mesal semi-
circular socket. Basal union of claspettes broadly Y-shaped.
DISTRIBUTION.-Costa Rica, Colombia, Mexico, Panama.
TYPES.-Holotype, male, allotype, female, El Hato, Chiriqui, Pan-
ama, 16-VI-1964, F. S. Blanton (type no. 72216, USNM). Paratypes, 5 males;
15 females, as follows: COLOMBIA: Rio Anori, Antioquia, IX-1970, D. G.
Young, light trap in tropical rain forest, 4 females. Rio Truando, Antioquia,
XI-1967, D. G. Young, light trap, 1 female. Tres Esquinas, Caqueta, 2-VI-
1968, C. J. Marinkelle, light trap, 1 female. COSTA RICA: Sabalito, Punta-
renas, VIII-1953, F. S. Blanton, light trap, 1 female; Turrialba, Cartago,
15-19-VII-1965, P. J. Spangler, 2 females. MEXICO: Concordia, Sinaloa, 4-
VIII-1964, W. R. Mason, malaise trap, 1 female. PANAMA: Barro Colorado
Island, C. Z., VII-1967, W. W. Wirth, light trap, 2 males; Gamboa, C. Z.,
Pipeline Road, VII-1967, W. W. Wirth, malaise trap, 3 females; El Volcan,
Chiriqui, 22-VII-1966, A. Broce, 2 males, 2 females; Rivira Farm, Chiriqui,
7-VII-1964, F. S. Blanton, 1 male.
DISCUSSION.-This unmarked yellowish species can be recognized by
a
b C
e
Fig. 7. Forcipomyia hatoensis: a, female antenna; b, male antenna,
last 4 segments; c, female palpus; d, female antenna, segments 10-11; e,
tibial comb of anterior leg; f, tibial comb of posterior leg; g, female sper-
mathecae; h, male genitalia.
Vol. 59, No. 2
Utmar and Wirth: New World Species of Forcipomyia 133
the pale, hyaline spermathecae in the female and the stout, rectangulate
male aedeagus with distomedian process arising from a distinct socket.
LITERATURE CITED
CHAN, K. L., AND E. J. LEROUX. 1965. Description of Forcipomyia (Neofor-
cipomyia) saundersi sp. n. and redescription of Forcipomyia (Neo-
forcipomyia) eques (Johannsen) (Diptera: Ceratopogonidae), with an
account of the digestive and reproductive systems. Phytoprotection
46:74-104.
DESSART, P. 1961. Contribution A l'etude des Ceratopogonidae (Diptera)
(II). Revision des Forcipomyia Africains decrits par le Dr. Goetghe-
buer. Bull. Ann. Soc. Roy. Ent. Belg. 97:315-76.
DESSART, P. 1963. Contribution A 1'6tude des Ceratopogonidae (Diptera)
(VII). Tableaux dichotomiques illustr6s pour la determination des
Forcipomyia Africains. Mem. Inst. Roy. Sci. Natur. Belg. 2 ser., fasc.
72, 151 p., 16 pl.
EDWARDS, F. W. 1928. Nematocera:23-102. In Insects of Samoa 4:1-366.
British Museum (Nat. Hist.) London.
GOETGHEBUER, M. 1934. Ceratopogonidae et Chironomidae nouveau ou peu
connus d'Europe (5e note). Bull. Ann. Soc. Roy. Ent. Belg. 74:287-
94.
GOETGHEBUER, M. 1936. Nouvelle contribution A la connaissance des
Ceratopogonides et des Chironomides de Belgique. Bull. Ann. Soc.
Roy. Ent. Belg. 76:319-26.
KRIVOSHEINA, N. P. 1968. A contribution to the biology and morphology of
little studied biting midges of the genus Forcipomyia Meigen (Dip-
tera: Ceratopogonidae). Zool. Zh. 47:578-90 (In Russian).
MACFIE, J. W. S. 1934. A new British Forcipomyia (Diptera, Ceratopogoni-
dae). Ent. Mon. Mag. 70:144.
MACFIE, J. W. S. 1939. A report on a collection of Brazilian Ceratopogo-
nidae (Diptera). Rev. Ent. (Rio de Janeiro) 10:137-219.
MACFIE, J. W. S. 1940. Forcipomyia furcifera sp. n. (Diptera, Ceratopogoni-
dae). Rev. Ent. (Rio de Janeiro) 11:920-2.
REMM, H. J. 1961. A survey of species of the Forcipomyia Meigen (Diptera,
Heleidae) from Estonia. Eesti Loodus Ser. 54:165-95 (In Russian).
SAUNDERS, L. G. 1956. Revision of the genus Forcipomyia based on charac-
ters of all stages (Diptera, Ceratopogonidae). Canad. J. Zool. 34:
657-705.
SNODGRASS, R. E. 1957. A revised interpretation of external reproductive
organs of male insects. Smithson. Misc. Collect. 135:1-60.
TOKUNAGA, M. 1940. Biting midges from Japan and neighboring countries,
including Micronesian Islands, Manchuria, North China, and Mon-
golia (Diptera, Ceratopogonidae). Tenthredo 3:58-165, 1 pl.
TOKUNAGA, M. 1962. Biting midges of Ryukyu Islands (Diptera: Ceratopo-
gonidae). Pac. Insects 4:153-217.
TOKUNAGA, M., AND E. K. MURACHI. 1959. Insects of Micronesia: Diptera,
Ceratopogonidae. Insects Micronesia 12:101-434.
WIRTH, W. W. 1951. New species and records of Virginia Heleidae. Proc.
Ent. Soc. Wash. 53:313-26.
WIRTH, W. W. 1952. The Heleidae of California. Univ. Calif. Publ. Ent.
9:95-266.
WIRTH, W. W. 1965. Family Ceratopogonidae (Heleidae):121-142. In Stone,
A., et al. A Catalog of the Diptera of America North of Mexico.
USDA, Agr. Handb. 276:1-1696.
WIRTH, W. W., AND N. MARSTON. 1968. A method for mounting small in-
sects on microscope slides in Canada balsam. Ann. Ent. Soc. Amer.
61:783-4.
The Florida Entomologist
BOOK REVIEW
ENVIRONMENTAL PHYSIOLOGY. D. Bellamy; G. J. Goldsworthy; K. C.
Goldsworthy; K. C. Hignam; W. Mordue; and J. G. Phillips. Ed. G. J. Phil-
lips. 1975. Halstead Press, A Division of John Wiley and Sons, Inc., New
York. 198 p. This book was written for undergraduate students with the
stated purpose of elucidating the basic physiological processes that have
evolved in various animal groups with the emphasis on the interrelation-
ship between environment and functioning of the organism. The authors
have succeeded in achieving this goal to a much higher degree than I had an-
ticipated they would in such a short book, primarily because of their writing
style and balance of coverage. Their writing is lucid, tightly organized,
and contains a minimum of superfluity. Their explanations are to the point
and have impact. Unlike most books of this ilk, they do not restrict their
choice of examples to vertebrates. They provide numerous examples from
a wide variety of animal groups in demonstrating the variation in physio-
logical solutions to common environmental problems. A surprising
amount of the coverage is devoted to insects thus providing a more com-
plete picture of information on the subject. As such I can recommend this
text to zoologists as well as those interested in the physiological ecology
of arthropods.
I must, however, temper my recommendation by mentioning certain
faults. First, there are a number of errors: for example, spiders are described
as arthropods which utilize trachea for respiratory gas exchange (p. 46) thus
having no requirement for an effective method of transporting their blood.
In actuality most spiders utilize book-lungs and a respiratory pigment,
hemocyanin, for exchange and transport of the respiratory gases. Secondly,
their treatment of certain topics is superficial. Less than a page is devoted
to the subject of respiratory pigments, and the discussion is so incomplete
that the uninformed reader will be left unsatisfied if he tries to understand
the basic concepts. Another major fault is the decidedly qualitative flavor
of the book. In only a few instances are formulae even mentioned. If, as
the authors have stated, this is a book for undergraduates, the lack of a quan-
titative approach seriously diminishes the book's value. A relatively
simple exposition involving quantitative models would present a more
realistic picture of the relationship between physiology and ecology. Fi-
nally, the lack of specific references to much of the source material is dis-
concerting; one cannot easily check for errors nor amplify material he finds
interesting.
These faults notwithstanding, the book can be a valuable supplement
to undergraduate course work in physiology, ecology, and arthropod bi-
ology.
John F. Anderson
The University of Florida
Gainesville, Florida 32611
Vol. 59, No. 2
The Florida Entomologist
EXPERIMENTAL VALIDATION OF A
PEST MANAGEMENT MODEL
W. W. MENKE, AND G. L. GREENE
Department of Industrial Management,
Clemson University, Clemson, S. Car. 29631, and
Agr. Research and Education Center, Quincy, Florida 32351, respectively
ABSTRACT
This paper describes 6 tests for validity which were conducted on a com-
puter-based simulation model designed to describe the population dy-
namics of the velvetbean caterpillar (VBC), Anticarsia gemmatalis Hub-
ner, in a soybean host crop. The tests were increasingly severe, ranging from
the preliminary test, the Turing criterion for reasonableness, to a most
stringent requirement that the model's output for the population dynamics
of a different arthropod (the citrus rust mite, Phyllocoptruta Oleivora
Ashm.) compares favorably with experimental measurements made in the
field. The favorable results reported, each showing close agreement be-
tween the model's output and the observed measurements of the popula-
tion dynamics, are believed to be strong substantiation for the model's va-
lidity. In addition, information regarding the VBC life cycle in Florida,
random sampling methods for measuring insect population densities, and
Geocoris as a predator for the VBC are discussed.
Simulation techniques provide a powerful way to analyze the per-
formance of a system so complex that its mathematical description is diffi-
cult or impossible to formulate. Typically the performance of a system's
simulation model is described by a description of the model, a statistical
analysis of the observed results, theoretical error analysis, a discussion of
the implications of the simulation results, and perhaps plans for future
research or model implementation. It is easy to understand that model
validation is most often restricted to the Turing method, where experts are
asked if the model results are reasonable (Emshoff and Sisson 1970) since
the very reason for developing the model, to avoid risky commitment of
scarce resources, may preclude easy experimental validation.
This paper describes a situation in which, from the very beginning of
model development, the authors planned for validation experiments, be-
lieving that a model is only conditionally acceptable, no matter how
reasonable it may seem, until its performance compares favorably with
the observed performance of the system being modeled. Further, even if
the performance comparisons are not favorable, feedback about differences
between model performance and observations of the real system may be
useful in refining the model to describe more accurately the real system
and to provide additional insight into the system's major interactions. These
principles are basic in theory; in practice they are often slighted because of
the cost involved or for expediency.
MATERIAL AND METHODS
The computer-based simulation model of this analysis has been de-
scribed in previous articles (Menke 1973a, 1973b, 1974). Briefly it simulates
Vol. 59, No. 2
The Florida Entomologist
several generations of the population dynamics of the velvetbean cater-
pillar (VBC), Anticarsia gemmatalis Hubner, in a soybean host crop and the
defoliation damage caused by the insect. The scenario used for model de-
velopment is described below.
The VBC moth overwinters in south Florida and is suspected of invading
north Florida from south Florida and the Caribbean area each year. The
female moth lays its eggs, and about 3 generations of insects develop dur-
ing the year until death in late fall (or out-migration of the adult moth)
reduces the population to essentially zero in north Florida soybean fields.
The eggs, estimated at ca. 800 per female, are laid individually by the
adult over a period of about 8 days. After hatching, the insect passes through
5 to 6 instars as caterpillars and a pupal stage before emerging as an adult
moth to mate, disperse, and lay eggs for the next generation. The length of
time the caterpillar remains in an instar (dwell time) is a stochastic vari-
able influenced by the environmental conditions during the stage. Avail-
able field experimental data indicate the dwell time probability distri-
bution to be approximately normal for all instars. Each, however, has a
different average and standard deviation (Reid 1975). The VBC causes
damage by defoliation of the soybean plant; the amount varies with the
instar.
The soybean plant was considered to have 10 stages of development
from emergence of unifoliate leaves through maturity. The plant's leaf
development between the critical stages in plant maturity, podset to pod-
fill, and the percent defoliation by the pest during this period were of pri-
mary interest in the model since economic crop damage occurs when thresh-
old defoliation levels are exceeded during this time.
Technically, the insect population dynamics are modeled in terms of
stochastic variates characterized by the adult moth invasion, the eggs, the
life cycle for each insect from each egg, and the continuous probability dis-
tributions for the dwell time at each instar.
The crop development portion of the model is a deterministic function
of time with the average value of leaf area per row-ft being the dependent
variable since the number of leaves per acre is so large (even at the seed-
ling stage, total leaf area calculated this way is very large and is approx-
imately 1 x 106 cm2/acre or 2.47 x 106 cm2/hectare). Critical points for the
functional relation are seedling, midbloom, podset, and podfill. All soy-
bean leaf area growth and defoliation by the VBC are expressed in square
centimeters per acre. Insect population counts and leaf damage are calcu-
lated at weekly intervals after the date of similar planting.
The model is presently one-dimensional in that it does not consider in-
migration or out-migration of the VBC after the starting moth invasion, nor
does it consider the spread or diffusion of the insect population within the
field from the point of contact of the initial invading moths. Care was taken
to insure independence between the model's output and observed results by
not using the observed results as feedback or input for the model.
The observed population densities used in these analyses were calcu-
lated from population samples taken in the field by the shake method (Tur-
nipseed 1974). Simple probability considerations reported previously
(Menke 1973b) indicated the need for about 200 row-ft (61 row-m) of plants
in the total sample if small insect populations were to be detected with
high probability. This was achieved by collecting 10 random shake samples,
Vol. 59, No. 2
Menke and Greene: Pest Management Model
each from 20 row-ft of host plants, for each day of sampling for population
density. Random samples were assured by using a computer to generate
random numbers and from these to calculate the coordinates of the start-
ing point in the field where each of the day's 10 random samples should be
collected. Sufficient coordinates were generated and listed in tabular form
to insure that randomness would be preserved for daily sampling for a
period of 4 months if desired. The average of each day's 10 sample popula-
tion counts was used as the best estimate of the insect population on that
day. Since sample averages were available, standard statistical techniques
were used to calculate 95% confidence levels on the sample results.
RESULTS AND DISCUSSION
Model validation is important in assessing the weight to be given to pre-
dictions and inferences obtained from experimenting with the model. It is
additionally important in judging how much time, money, manpower, and
other resources should be used to implement the model's predicted opti-
mum strategies in the real system (Shannon 1975).
The first step (T1) in any validation procedure is to test for reasonable-
ness of results. This, the Turing criterion, was reported in previous papers
(Menke 1973b, 1974) where experimental designs for the model were inter-
preted by using analysis of variance techniques, and it was shown that use-
ful and reasonable biological strategies were identified.
The next step (T2) in validating the model of this paper was accom-
plished during the design phase when care was taken to insure that the pa-
rameters and variable values required as data for the operation of the
model had real physical and technical significance. This, the variable-
parameter validity test (Emshoff and Sisson 1970), was accomplished by
using a combination of data gathering techniques (Menke 1973b, 1974) from
PERT type questions (minimum-usual-maximum values) to the exam-
ination of published literature. These methods of data gathering assured
that only those parameters and variable values that made sense to the prac-
ticing entomologist and could be easily identified and collected in the
field or from the accepted literature would be used.
A more discriminating step (T3) in validation was taken where the gen-
eral character of the model's output of population density versus time was
first compared with the population dynamics observed in the field. The
model's output fell within the 95% confidence region shown in Fig. 1, but
the confidence levels for the experimental field results were unknown.
Nevertheless, while the periodicity of the population cycles and the rela-
tive magnitudes of the population peaks and troughs compared favorably
between the modeled and measured data, specific event timing did not, i.e.,
a time delay was observed between the model output and the results in the
field. The shift in timing as well as the relative size of the population at
the peaks, suggested that the low density populations previously observed
in the field and identified as a first generation population may in fact be a
second generation population. This in turn implied that the occurrence of
very low density first generation population of VBC should be observed
earlier in the year than had ever been previously reported. This was con-
firmed experimentally in the summer of 1974 by Greene's observations.
The verification of the "lost generation" effect (T4) did much to estab-
The Florida Entomologist
7/3 10 17 24 31 8/7 14 21 28 9/11 25
DATE OF COUNT
Fig. 1. VBC measured population dynamics compared to the 95% C. L.
envelope derived from the model.
lish credence in the simulation as a valid normative model, i.e., that the
simulation model was acting as it ought to. Simultaneously it reinforced
speculations that the previously accepted assumptions that a moth invasion
from the Caribbean started a new cycle of VBC population dynamics each
year might not be correct and that VBC life cycle starting conditions should
be further investigated.
Vol. 59, No. 2
Menke and Greene: Pest Management Model
It was felt that a major step towards demonstrating validity with even
higher specificity would be achieved by comparing model output in both
magnitude and timing (at 95% confidence) with real field data (T5) provided
the field data were a random sample of sufficient size to permit confidence
limits on the sample results to be estimated.
A comparison between the measured average VBC population densities
determined from 200 row-ft shake samples taken as previously described
and the model's predictions for the period July through September 1973 is
shown in Fig. 1. The plotted x's identify each field sample average. The
capped vertical line through each average indicates the 95% confidence
range for the measurement. The population density limits on the predic-
tions calculated from the model's simulated output values are shown by
the upper and lower boundaries of the 95% C.L. envelope superimposed
on the experimental results of the figure. The model's simulated output
values are representative of 9 complete and distinct simulation runs over
2 generations of VBC life. The dashed portion of the envelope defines the
highly uncertain population estimates from the model during the start of a
new generation. This uncertainty reflects the sensitivity of the start of a new
generation to the number of ovipositing females surviving from the pre-
ceding generation and to the exact number and development rate of the eggs
they lay. It can be seen that the model predicts that the variance or uncer-
tainty in estimating population densities is reduced as the generation peak
is approached. Additional simulation runs would have to be made in order
to significantly reduce the uncertainty in the model's predictions at the
start of each new generation. Additional runs were not made since the start-
ing period is one where only small numbers of 1st, 2nd, or 3rd instar larvae
exist and, therefore, little defoliation occurs. Hence a higher degree of un-
certainty is allowable than that for periods near the generation peaks where
large numbers of 5th and 6th instar larvae cause significant defoliation.
The close agreement between the measured and predicted data shown in
Fig. 1 was felt to be a rigorous test of the model's performance since it faith-
fully portrayed significant portions of 2 population cycles of the VBC
during times when the population density varied by more than 2 orders of
magnitude from values of several hundred insects per acre (or hectare) to
values of nearly 70 thousand insects per acre (nearly 173 thousand insects
per hectare). Note that the population dynamics plotted for the period from
24 July to 24 August included the intricate overlapping effects on total
population of mating, egg laying, and end of life of 1 generation of the
VBC and the emergence and growth of its progeny.
On 24 August (8/24) the beginning of an infection that was later recog-
nized as a Nomuraea rileyi epizootic was noted. Subsequent agreement be-
tween the 2 sets of data was no longer expected or observed. It is believed
that identification and analysis of causes for the differences between the
population dynamics observed during the epizootic and the model's pre-
dicted values for no infection may hold the key to the efficacy of using the
fungus, Nomuraea rileyi, as a pest management tool for VBC control.
During the examination of each shake catch for the VBC, the popula-
tions of other insects that were simultaneously collected were classified,
counted, and recorded. Fig. 1 shows the average and the 95% confidence
limits on the observed population density of the insect, Geocoris punctipes
(Say) usually believed to be an aggressive predator of the VBC (W. H.
The Florida Entomologist
Whitcomb, unpublished data). The data clearly show an almost uniformly
high Geocoris population density of about 8,000 insects per acre during the
violent population density fluctuations observed for the VBC. Classical
predator-host theories (Watt 1968) predict gross predator population den-
sity fluctuations similar to the host population's gross variations but lag-
ging them in time phase. Clearly the experimental data collected in north
Florida soybean fields show no such interaction. Evidently the predator
had an ample supply of a more or equally preferred food than that af-
forded by the VBC. In any event, the data suggest that a very large popula-
tion density of Geocoris had little population regulation effect on the
VBC. It is interesting that during 1974, upon replication of these field ex-
periments, the effect remained the same and no other variable was identified
which could have confounded the results. Such conclusions would elimi-
nate any inclination towards using Geocoris as a biological control strat-
egy for management of the VBC. Reevaluation of predator-host relations
between Geocoris and the VBC in a north Florida environment appears to
be indicated.
In order to explore further the capabilities of the simulation model, the
population dynamics of the citrus rust mite, Phyllocoptruta Oleivora Ashm.,
a pest entirely different from the VBC, were simulated for 7 generations
(T6). The parameters (longevity, oviposition rate, mortality factors, male
to female ratio, stadia characteristics, temperature relations, etc.) required
for model operation were not available from data in Florida. However,
Swirski and Amitai's (1958) work in Israel on the citrus rust mite provided
data from which the necessary parameters could be determined for environ-
mental conditions similar to those of south Florida.
A comparison of the initial computer run for the simulated relative
population density versus time (days) for the citrus rust mite with experi-
mental population density measurements made in 2 south Florida orange
groves is shown in Fig. 2. Both the slope of the curve for mite density vs.
time and the densities predicted by the model agree favorably with the ex-
perimentally measured values in the 2 groves except for the regions for
days 165-175 for grove I and days 227-242 in grove II where notches occurred
in the experimental population dynamics curves. Observations in the field
indicated these notches to be coincident with the onset of infection of mites
in the field by the pathogen Hirsutella thompsonii.
In addition to the above agreement, the duration of each generation and
its overlap with at least 2 preceding and 2 succeeding generations as simu-
lated by the model are shown in the upper left portion of Fig. 2. These data
are strikingly similar to the experimental results reported in Swirski and
Amitai's (1958) Fig. 6, "Overlapping of Generations at Rehovot".
The above comparison was the last and most demanding step taken to
test the model's validity. Credence in the model and its predictions was
enhanced by its ability to simulate the population dynamics of 2 such dif-
ferent insects as the VBC and the citrus rust mite.
LITERATURE CITED
EMSHOFF, J. R., AND R. L. SISSON. 1970. p. 204-6. In: Design and uses of com-
puter simulation models, McMillan, N.Y., xvii + 302 p.
Vol. 59, No. 2
Menke and Greene: Pest Management Model 141
1000 K
500 K
Compu ler pred cted
overlap ping of generations
100 K
4-- 7 6
a x
2 '
LLI 5
1K I
0 x
_ ./ simulated
.5 x experimental
X Grove I
6 Grove II
80 100 120 140 160 180 200 220 240 260
K=1000 DAY OF COUNT
Fig. 2. Citrus rust mite measured population dynamics compared to the
model's predictions.
MENKE, W. W. 1973a. Identification of viable biological strategies for pest
management by simulation studies. 1973 Winter Simulation Confer.
Pro. January 1973, p. 32-50.
MENKE, W. W. 1973b. A computer simulation model: the velvetbean cater-
pillar in the soybean agroecosystem. Fla. Ent. 56:92-102.
MENKE, W. W. 1974. Identification of viable biological strategies for pest
management by simulation studies. IEEE Transactions on Systems,
Man, and Cybernetics, Vol. SMC-4, No. 4, July 1974:379-86.
NAYLOR, T. H. (ed.). 1969. p. 232-51. In: The design of computer simulation
experiments, Duke University Press, Durham, N.C. 417 p.
REID, J. C. 1975. Larval development and consumption of soybean foliage
by the velvetbean caterpillar, Anticarsia gemmatalis Hubner (Lepi-
doptera: Noctuidae), in the laboratory. Univ. Fla. Ph.D. Disser.
118 p.
SHANNON, ROBERT E. 1975. Chapter 6. In: Systems simulation the art and
science, Prentice-Hall, Inc., Englewood Cliffs, N. J. xii + 386 p.
The Florida Entomologist
SWIRSKI, E., AND S. AMITAI. 1958. Contributions to the biology of the citrus
rust mite, Pub. of AG. RES. Capital Station, Rehovot, 1958 Series,
No. 233-E.
TURNIPSEED, S. 1974. Sampling soybean insects by various D-vac, sweep,
and ground cloth methods, Fla. Ent. 57:217-23.
WATT, K. E. F. 1968. Chapter 6. In: Ecology and resource management,
McGraw-Hill, N. Y. xii + 450 p.
BOOK REVIEW
INSECT HORMONES. V. J. A. Novak. 1975. Chapman and Hall, London; John
Wiley and Sons, New York, 600 p. I was introduced to Professor Novak's
views of metamorphosis through the first English edition of Insect Hor-
mones published in 1966. The 1975 edition shares the same strong points as
the earlier work, but the weaknesses are now even more striking. Professor
NovAk presents a very personal view of classical insect endocrinology.
In the chapter on "Hormones and Morphogenesis" various theories of
metamorphosis are given equal time such that the currently accepted view
of hormonal control receives the same explicit attention as other concepts
that are only of historical interest. Only NovAk's "gradient factor" theory
of metamorphosis receives a full discussion. In brief, NovAk's theory calls
for the "existence of a tissue-bound factor that determines morphogenesis
by the way it has been distributed with the body." Certainly, developmental
gradients are well known in many animals including insects, and are often
invoked in studies of pattern formation. It is not at all clear, however, how
NovAk's theory would be helpful in solving problems in insect hormone
action. Furthermore, students of insect physiology would get a distorted
view from Insect Hormones of the relative status of concepts of insect meta-
morphosis.
I am disappointed that despite the inclusion of some sections on the ef-
fects of hormones on chromosomes and on various cellular activities, there
is a very limited presentation of the very exciting advances that have been
made during the past 10 years in the study of the mechanisms of action of in-
sect hormones. In a comprehensive book on insect hormones that runs to
600 pages I would expect more than 2 or 3% of the space to be devoted to
how hormones act at the molecular level. More space is actually devoted
to a chapter on pheromones.
The section of methods in insect hormone research contains a useful
discussion of surgical techniques, but a cursory and arbitrary consideration
of other methods. As the author observes "almost all the methods used in
physiological research could be cited . .," so why have a separate section
discussion only a few of them?
The publishers suggest that Insect Hormones ". ..is invaluable to all
post graduate researchers in Insect Endocrinology, Entomology and Pest
Control .. ." Despite the shortcomings of NovAk's book, I would agree that
it is useful as a reference on the insect endocrine system. The sections on
neurohormones and on the early studies with juvenile hormone and ecdy-
sone are useful. There are also many photographs of the endocrine organs
and of insects and tissues treated with hormones. Certainly, the reference
list of well over 100 pages should be helpful. I would caution, however,
that this book should be introduced to students only if they also consider
other sources that more closely reflect the views and activities of insect
endocrinologists today.
Herbert Oberlander
ARS-USDA
Gainesville, Florida 32604
Vol. 59, No. 2
The Florida Entomologist
KEYS TO GENERA OF ANTHOCORIDAE OF AMERICA
NORTH OF MEXICO, WITH DESCRIPTION OF A NEW
GENUS (HEMIPTERA: HETEROPTERA)
JON L. HERRING
Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA'
ABSTRACT
A key to the genera of Anthocoridae known from America north of Mex-
ico is presented. A new genus, Alofa, is proposed for Cardiastethus sodalis
White which is currently placed in the genus Buchananiella Reuter.
All members of the family Anthocoridae are predaceous. Although a
few widely distributed species are encountered in flower heads, most occupy
a wide variety of unexposed niches. They occur beneath loose or dead bark,
in piles of decaying weeds and leaves, in beetle galleries in shelf fungi, in
the nests of birds and woodrats, in bat caves and in epiphytes such as bro-
meliads and orchids. They are often intercepted in interstate and interna-
tional shipments of bulbs, cut flowers, nursery stock and stored grain prod-
ucts.
Many species of anthocorids are of significant economic value as a means
of biological control of agricultural pests. Two North American species
of Orius have been released in Hawaii for aid in controlling certain lepi-
dopterous larvae. Montandoniola has been introduced from the Philippines
into Hawaii and California for the control of the Cuban laurel thrips,
Gynaikothrips ficorum (Marchal), and several Mexican species of Macro-
tracheliella have been released in California for this same pest. The Ca-
nadian workers have been attempting to introduce into Canada species of
Tetraphleps from India and Pakistan for the control of the balsam woolly
aphid, Chermes piceae Ratzeburg.
Existing keys to the genera of Anthocoridae of America north of Mexico
(e.g. Reuter 1884, Poppius 1909) are incomplete and out of date.
One new genus is described in the key: Alofa, proposed for Cardiastethus
sodalis White which is currently placed in the genus Buchananiella Reuter.
KEY TO SUBFAMILIES
1. Antennal segments II, III and IV of equal thickness (if II ab-
normally swollen, then III and IV are differently colored).
Erect hairs on III and IV never longer than 2X diameter of seg-
ment. Metapleural scent gland curved forward or straight
and often prolonged forward by a carina or groove. Hamus
always present and generally attached to the cubitus. Female
with a well-developed ovipositor, male with a single para-
mere ............ ............... ...... Anthocorinae Reuter 1884
1Mail address: c/o U. S. National Museum, Washington, D. C. 20560.
Vol. 59, No. 2
The Florida Entomologist
1'. Antennal segments III and IV always thinner than II and pro-
vided with long erect pubescence which is usually longer
than 2X diameter of the segment. Metapleural scent gland
curved forward or backward. Hamus, if present, attached to
the m-cu. Female with or without a well-developed ovi-
positor. M ale with 1 or 2 param eres........................................... ......... 2
2. Metapleural scent gland curved or directed backward and
never prolonged by a carina or groove. Pubescence abundant
on the margins of the pronotum and costal margins of the
hemelytra. Hamus usually present. Female ovipositor al-
ways present. Male with a single bladelike paramere on the
left side ......................................................... Lasiochilinae Carayon 1972
2'. Metapleural scent gland curved forward or backward and
often prolonged by a carina or groove which attains the an-
terior margin of the metapleura (occasionally (Physopleur-
ella) curved backward without a carina, in which case the
fore femora are enlarged and armed with spines). Pubescence
not abundant along the margins of the pronotum and costal
margins of the hemelytra. Hamus usually absent. Female
ovipositor well developed or vestigial, male with 1 or 2 para-
m eres ........................................... ...................Lyctocorinae R euter 1884
KEY TO THE TRIBES OF ANTHOCORINAE
1. Prothorax generally provided with a distinct visible collar,
more or less large, sometimes transversely rugulose. Tarsi
devoid of pseudorolia between the claws. Male fore tibiae
not spinose on their inner surface. Eighth abdominal segment
of male but slightly asymmetrical, genital segment bearing
a thin falcate paramere on the left side........ Anthocorini Carayon 1958
1'. Prothoracic collar narrow, poorly differentiated, not or only
slightly transversely rugulose. Tarsi with a pair of pseuda-
rolia between the claws. Male fore tibiae spinulose on their
inner surface. Eighth abdominal segment of male asymmetri-
cal, curved to the left, genital segment bearing a spiral para-
mere ........................................ ....................... Oriini Carayon 1958
KEY TO THE GENERA OF ANTHOCORINI
1. Apex of metasternum straight or evenly rounded, the hind
coxae w idely separated .............................................................................. 2
1'. Apex of metasternum triangularly produced, the hind coxae
contiguous or nearly so......................... ..... ................................... 5
2. Prothoracic collar indistinct, enclosed in the antero-lateral
reflexed margins of the pronotum. Metapleural scent gland
very short, straight, its tip elevated (Fig. 1); elytra with seri-
ally arranged, silver, scalelike pubescence..............................
.......... .......................... ................. .. M elanocoris Champion 1900
2'. Thoracic collar distinct, clearly delimited on the anterior
margin of the pronotum, Metapleural scent gland not ele-
vated, often extending forward as a carina. Pubescence normal ........ 3
Vol. 59, No. 2
Herring: Genera of Anthocoridae
2 3
12 3
/i
t '
t~t#'' !'- "' l/
~"Th
'I
/
7
7
/^.
/ /;- /
Z.^ ^->
, ,,
9
10 11 12
Fig. 1-12. Metapleural scent glands: 1) Melanocoris, 2) Elatophilus,
3) Temnostethus, 4) Coccivora, 5) Anthocoris, 6) Tetraphleps, 7) Acompo-
coris, 8) Orius, 9) Paratriphleps, 10) Macrotracheliella, 11) Montandoniola,
12) Lasiochilus.
3. Posterior margin of pronotum twice as wide as head across
eyes, lateral margins narrowly explanate. Scent gland
evenly curved forward and extending anteriorly to anterior
The Florida Entomologist
margin of metapleura as an obtuse carina (Fig. 2). Macropter-
ous forms only .................................................. Elatophilus Reuter 1884
3'. Posterior margin of pronotum not twice as wide as head, lat-
eral margins and scent gland not as above. Micropterous
form s m ost com m on ............................ ..... .................................... 4
4. Head much longer than pronotum, pronotal margins expla-
nate and reflexed anteriorly. Rostrum reaching middle of
mesosternum or beyond. Posterior margin of metasternum
rounded, scent gland transverse, nearly straight (Fig. 3). Apex
of wing in brachypterous form rounded. Macropterous forms
unknown ......................................................... Temnostethus Fieber 1860
4'. Head about as long as pronotum, pronotal margins sinuate
and carinate. Rostrum reaching to anterior coxae. Posterior
margin of metasternum straight, scent gland auriculate with
a prominent opening at its base (Fig. 4). Apex of wing in bra-
chypterous form straight. Macropterous forms very slender,
wings sinuate at middle ..............Coccivora McAtee and Malloch 1925
5. Elytra impunctate or extremely finely punctured, pubescence
usually quite short and rather sparse. Collar completely in
front of anterolateral angles. Head shorter, anteocular por-
tion not as long as ocular-postocular. Rostrum not or barely
surpassing anterior coxae. Scent gland pale margined, straight,
curved slightly forward at apex and joined to a fine carina
which reaches the anterior margin of the metapleura (Fig. 5)......
....................................... .................................... A nthocoris F allen 1814
5'. Elytra densely punctured and heavily clothed with long pu-
bescence, collar partially enclosed in anterior angles. Head
longer, anteocular portion at least equal to ocular-post-
ocular portion. Rostrum at least surpassing fore coxae, scent
gland not pale margined or joined to a carina ..................................... 6
6. Scent gland almost straight, extending toward lateral mar-
gin of metapleura and distinctly elevated at its apex (Fig. 6).
Rostrum variable, at least surpassing fore coxae, and in some
cases, onto abdomen ....................................... Tetraphleps Fieber 1860
6'. Scent gland slender, slightly curved forward, not elevated
at its apex (Fig. 7). Rostrum surpassing hind coxae.................
...............................................................A..... ... A com pocoris R euter 1875
KEY TO GENERA OF ORIINI
1. Apex of metasternum, triangularly produced, posterior coxae
contiguous or nearly so. Head short, hardly produced in front
o f ey es ................................ ............ .............. ..... .. ............... .. ..... ..... . .. 2
1'. Apex of metasternum straight or rounded, middle and hind
coxae widely separated. Head long, well produced in front
of eyes .. ........................ ........................ ................................. 3
2. Body length 2.5 x greatest width. Clavus, at least, distinctly
punctured. Scent gland evenly curved forward to anterior
Vol. 59, No. 2
Herring: Genera of Anthocoridae
margin, evapatorium broad, shining (Fig. 8). Head black or
brown ........... ... .. ................................. .. Orius W olff 1861
2'. Body much more robust, length no more than 2x greatest
width. Pronotal width 3 x median length. Only cuneus dis-
tinctly punctured, occasionally with rows of fine punctures
along the veins. Scent gland usually angulate before curving
forward to anterior margin, evapatorium much narrower,
hardly wider than gland (Fig. 9). Head bright yellow..............
..................................................................... P aratrip hlep s C ham pion 1900
3. Head very long, much longer than median length of pro-
notum, postocular length greater than length of an eye. An-
tennal segment II clavate, all segments of almost equal di-
ameter. Cuneus short, only 1/2 as long as the embolium. Apex
of scent gland opening not tapering, rounded at apex (Fig.
10) ....................................................... Macrotracheliella Champion 1900
3'. Head moderately long, equal to median length of pronotum,
postocular length shorter than length of an eye. Antennal
segment II greatly swollen, its diameter greater than termi-
nal segments, III and IV much paler than I and II. Cuneus
long, 3/4 as long as embolium. Apex of scent gland taper-
ing, acute at apex (Fig. 11)...................... Montandoniola Poppius 1909
KEY TO THE GENERA OF LASIOCHILINAE
1. Antennal segment II 4x as long as I and equal to III + IV.
Pronotum deeply, transversely bisulcate on the disc. Cor-
ium and embolium of almost equal width at apex (not seen)....
................................. .................. ......... Plochiocoris Cham pion 1895
1'. Antennal segment II never 4x as long as I nor equal to
III+ IV. Pronotum with a median sulcus only. Embolium
much narrower than corium at apex (scent gland Fig. 12)......
.................................... ..................................... L asioch ilus R euter 1871
KEY TO THE TRIBES OF LYCOTOCORINAE
1. Scent gland distinctly elbowed at middle, either continued
toward anterior margin or joined at a right angle by a fine
carina which reaches anterior margin. Antennal segments III
and IV very thin and provided with erect pubescence much
longer than the diameter of the segment. Female with ovi-
positor well developed. Fore tibiae of male strongly dilated
from base to apex, their apical width more than 2 x, sometimes
3 x their basal width and provided at their apex with a prom-
in e n t b ru sh ........................ ....................... ....... ....... ..... ............................. 2
1'. Scent gland not elbowed at middle, evenly curved forward
or backward. Antennal III and IV hardly thinner than II and
pubescence usually not more than 2 x the diameter of the seg-
ment. Ovipositor often vestigial. Fore tibia of male less
strongly dilated from base to apex, their apical width no more
than 2 x their basal width, apical brush poorly developed or
absent ................... ................... .................. ... 3
The Florida Entomologist
13 14
17
Fig. 13-22. Metapleural scent glands: 13) Lyctocoris, 14) Xylocoris,
15) Scoloposcelis, 16) Calliodis, 17) Nidicola, 18) Physopleurella, 19) Am-
phiareus, 20) Alofa, 21) Cardiastethus, 22) Dufouriellus.
2. Sides of pronotum distinctly margined, margins reflexed; apex
of abdomen without long hairs. Elytra everywhere thickly
punctate. Scent gland almost straight, joined at a sharp right
Vol. 59, No. 2
,
, ~
r ;;
Herring: Genera of Anthocoridae
angle by a very fine carina which extends to the front margin
of the m etapleura (Fig. 13)...........................................
.............. Lyctocorini Carayon 1972 (one genus, Lyctocoris Hahn 1855)
2'. Sides of pronotum not margined, margins deflexed; apex of
abdomen with several long hairs. Elytra, at most, sparsely
punctate. Scent gland curved forward at middle, almost or
quite reaching front margin of metapleura (Fig. 14)..................
.............. Xylocorini Carayon 1972 (one genus, Xylocoris Dufour 1831)
3. Scent gland short, curved forward but not attaining anterior
margin of metasternum or joined by a carina which attains an-
terior margin. Rostrum reaching midcoxae or beyond. Scu-
tellum without circular depressions....................Scolopini Carayon 1954
3'. Scent gland usually longer, curved forward or backward, ei-
ther attaining the anterior margin of metasternum or joined
by a carina which reaches the anterior margin (Exception: In
Physopleurella, the scent gland is not prolonged as a carina,
but the apex is directed backward). Rostrum short, usually
not surpassing the prosternum. Scutellum usually with a
circular depression on each side of middle.........................
.. ................................... .......... ......... Cardiastethini Carayon 1972
KEY TO THE GENERA OF SCOLOPINI
1. Body depressed, flat dorsally, sides subparallel. Legs robust,
anterior and posterior femora enlarged, provided ventrally
with short, irregular teeth. Clavus and corium impunctate.
Metasternum without a median longitudinal keel (scent
gland Fig. 15) ................................................... Scoloposcelis Fieber 1864
1'. Body oval, not depressed. Legs long and slender, femora not
enlarged nor provided with teeth. Clavus and corium punc-
tate. Metasternum with a median longitudinal keel .......................... 2
2. Collar present, pronotum carinate only. Clavus and corium
with punctures serially arranged. Rostrum reaching midcoxae
(scent gland Fig. 16)........................... ..................................
................................ Calliodis Reuter 1871 (= Asthenidea Reuter 1884)
2'. Collar absent, lateral margins of pronotum and hemelytra
widely, explanate and reflexed. Clavus and corium with
punctures along veins but not serially arranged. Rostrum
reaching between hind coxae (scent gland Fig. 17)................
................................ ................ Nidicola Harris and Drake 1941
KEY TO THE GENERA OF CARDIASTETHINI
1. Metapleural scent gland directed backward and usually pro-
longed by a carina or groove which attains the anterior margin
of the m etapleura (Fig. 18-20) ........................................ ...................... 2
1'. Metapleural scent gland curved evenly forward to anterior
margin of the metapleura (Fig. 21-22) ............................. ............. 4
2. Scent gland not prolonged by a carina to the anterior margin
(Fig. 18). Fore femora enlarged and armed beneath with spines.
The Florida Entomologist
Rostrum shorter than head, second joint awl shaped. Pro-
notum with a median longitudinal groove......................
.............................. .......................... ......... Physopleurella R euter 1884
2'. Scent gland prolonged by a fine carina to the anterior mar-
gin. Fore femora slender, unarmed. Rostrum reaching the an-
terior coxae. Pronotum without a complete longitudinal
g ro o v e ................................................................ .............................. ............... 3
3. Clavus and scutellum punctate. Corium and embolium at
apex of equal width. Metasternum prolonged posteriorly by
a bifid apophysis. Pronotal collar not set off by punctures
bearing long hairs (scent gland Fig. 19) ..........Amphiareus Distant 1904
3'. Clavus and scutellum impunctate. Corium at apex at least
2 x width of embolium. Metasternum without apophysis. Pro-
notal collar set off by punctures bearing long hairs (scent
gland Fig. 20) (Cardiastethus sodalis White, type-species;
gender feminine) ......................................... Alofa Herring, new genus
4. Collar present, prominent long hairs on pronotum and
hemelytra. Basal margin of pronotum very deeply sinuate or
concave, disk of pronotum with a distinct transverse groove
(scent gland Fig. 21) ..................................... Cardiastethus Fieber 1860
4'. Collar absent, pronotum and hemelytra with only a few
prominent long hairs. Basal margin of pronotum more widely,
less deeply sinuate, disk of pronotum without a transverse
groove but with a distinct median longitudinal one (scent
gland Fig. 22) ..................... ................Dufouriellus Van Duzee 1917
LITERATURE CITED
PoPPIUS, B. R. 1909. Beitrage zur Kenntnis der Anthocoriden. Acta.
Soc. Sci. Fennicae 37(9):1-43.
REUTER, O. M. 1884. Monographia Anthocoridarum orbis terrestris. Acta
Soc. Sci. Fennicae 14:555-758, 1885. Separate, Helsingfors, 1884,
p. 1-204.
A disadvantage of being clear is that one must sometimes be clearly
wrong. But it is better to risk that than to resort to obscurity.
S. A. Barnett
Vol. 59, No. 2
151 The Florida Entomologist Vol. 59, No. 2
PSEUDOSCORPIONS FROM FLORIDA AND THE
CARIBBEAN AREA.' 5. AMERICHERNES, A NEW GENUS
BASED UPON CHELIFER OBLONGUS
SAY (CHERNETIDAE)2
WILLIAM B. MUCHMORE3
Department of Biology, University of Rochester,
Rochester, New York 14627
ABSTRACT
The genus Americhernes is defined, with Chelifer oblongus Say, from
eastern United States, as the type species. Also placed in the genus are:
Lamprochernes ellipticus Hoff, from Baja California, Mexico; Lampro-
chernes levipalpus Muchmore, from Utah; Americhernes longimanus, new
species, from Florida and the Gulf Coast; A. reductus, new species, from
the Florida Keys; and A. puertoricensis, new species, from Puerto Rico.
The relationships between this and allied genera are discussed briefly.
As pointed out in an earlier paper (Muchmore 1972), a certain amount
of confusion has resulted from the inclusion in the genus Lamprochernes
TSmosviry of some American forms of pseudoscorpions of basically dif-
ferent morphology. One of the most common of these, Chelifer oblongus
Say, which has long been known as Lamprochernes oblongus (Beier 1932b:
84; Hoff 1949:450) is selected as the type species of the new genus.
Americhernes Muchmore, new genus
TYPE SPECIES: Chelifer oblongus Say, 1821.
DIAGNOSIS: A genus of the family Chernetidae (Muchmore 1974:26).
Generally small pseudoscorpions of elongate form and with relatively
stout appendages and smooth, shiny derm. Usually moderately sclero-
tized and darkly colored; surfaces of carapace and abdominal sclerites
smooth and shiny, those of palps mostly smooth and shiny but with some
areas of small granules; pleural membranes smoothly, longitudinally
striate; vestitural setae weakly denticulate subterminally to acuminate;
setae of genital opercula acuminate, those of spiracular plates and anal
plates finely denticulate; 11th tergite and sternite each with 4-6 long, acu-
minate tactile setae. Carapace with a single transverse furrow at about
middle; 2 eyespots present. Cheliceral hand with 5 setae, of which sb, b and
es are terminally denticulate; flagellum of 3 setae, distalmost larger and
serrate along 1 margin; galea conspicuous, with 5-6 rami. Palps rather ro-
bust; trochanter with prominent dorsal protuberance; femur and tibia dis-
tinctly pedicellate; chelal fingers shorter than hand; venom apparatus
'A continuation of the series originally entitled "Pseudoscorpions of Florida."
'Contribution No. 334, Bureau of Entomology, Division of Plant Industry, Florida De-
partment of Agriculture and Consumer Services, Gainesville, Florida 32602.
'Research Associate, Florida State Collection of Arthropods, Florida Department of Agri-
culture and Consumer Services, Gainesville.
152 The Florida Entomologist Vol. 59, No. 2
well developed in movable finger, apparently absent in fixed finger; on
fixed finger, trichobothrium ist at level of, or proximal to, est; it close to
ist and farther from finger tip than distance between ist and isb; each pedal
tarsus with an elevated slit sensillum near proximal end; tibia of leg IV
(and III) with 2 long, tactile setae on outer margin, 1 near middle and the
other very near distal end; tarsus of leg IV (and III) with long, tactile seta
about 0.25 length of segment from proximal end; male anterior genital
operculum with about 4 long, curved, heavy setae surrounded by 14-18
smaller ones; male posterior operculum with 2-3 small setae on either side
of middle inside of anterior margin and 6-12 larger setae on face; female
anterior genital operculum with a compact group of about 6-10 small
setae near center and 2 sets of 2-3 small setae posterior to this, with 2 con-
spicuous, obliquely oriented lyrifissures just behind the compact group of
setae, and with a longitudinally striated and roughened sclerotized area
lying between the 2 small sets of setae; female posterior operculum with
about 6-10 setae in a row along posterior margin; internal genitalia of male
typical of the family; spermathecae of female in form of 2 separate, curved
tubes, expanded distally into cylindrical sacs.
REMARKS: Americhernes may be distinguished from Lamprochernes
and allied American genera by means of the following key:
1. Tibia of leg IV with a prominent tactile seta very near distal
end of outer margin (Fig. 1); spermatheca of female in form
of single tube divided distally into 2 backward curving cornua
(F ig. 2) ............................................... ................................L am p roch ern es
1'. Tibia of leg IV with a prominent tactile seta near middle of
outer margin in addition to tactile seta near distal end (Fig.
9); spermathecae of female in form of 2 separate tubules with
saclike expansions in distal halves (Fig. 5) .......................................... 2
Fig. 1-2. Lamprochernes chyzeri (TS6msviry): 1) Leg IV; 2) Spermatheca
of female.
Muchmore: New Pseudoscorpion Genus
2. Trichobothrium it of fixed chelal finger clearly farther from
finger tip than the distance between ist and isb ............. Americhernes
2'. Trichobothrium it of fixed chelal finger at least as close to
finger tip as the distance between ist and isb ............Lustrochernes, etc.
Americhernes differs further from Lamprochernes in having a perfectly
smooth carapace, without granules laterally as is the case in Lampro-
chernes, and in the arrangement of setae on the genital operculum of the
female, some of which are tightly grouped in Americhernes but more scat-
tered in Lamprochernes. In neither genus are the tactile setae on leg IV
necessarily acuminate, but are often very finely denticulate subtermi-
nally.
The distinction between Americhernes and the other allied American
genera such as Lustrochernes, Cordylochernes and Mesochernes is less easy
to make, mainly because these other genera are only poorly defined at pres-
ent. However, the distinction made by Beier (1932a:259) with respect to the
placement of trichobothria on the fixed chelal finger seems to hold and is
employed in the key given above. It appears also that Americhernes species
are smaller than those of the other genera, but this may not always be so.
Other more subtle differences are presently being investigated in the at-
tempt to settle the relationships in this entire group.
Americhernes oblongus (Say), new combination
Fig. 3-9
Chelifer oblongus Say, 1821:64.
Chelifer (Lamprochernes) oblongus; Ellingsen, 1909:368.
Chelanops oblongus; Ewing, 1911:79.
Lamprochernes oblongus; Beier, 1932b:84; Hoff, 1945:37; 1949:450; 1958:20;
Hoff and Bolsterli, 1956:165.
The type specimens are lost, and the species is presently based upon a
neotype from Illinois, designated by Hoff (1949). The type specimen (5748-
S-483.2) is in the collection of the Illinois Natural History Survey, Ur-
bana, Illinois.
MATERIAL STUDIED. ILLINOIS: Neotype male from Havana,
Mason County; Massac County, 2 males, 2 females. NEW YORK: Genesee
County, 2 males, 1 female. MASSACHUSETTS: Barnstable County, 5
males, 8 females. NEW JERSEY: Gloucester County, 5 males, 1 female.
MARYLAND: Caroline County, 2 males, 4 females. NORTH CAROLINA:
Carteret County, 4 males, 6 females. SOUTH CAROLINA: Oconee County,
2 males; Pickens County, 1 female. FLORIDA: Alachua County, 2 males;
Lake County, 2 males; Highlands County, 4 males, 1 female. MIS-
SISSIPPI: Harrison County, 2 females; Hinds County, 1 male; Leake
County, 1 female; Choctaw County, 1 male. LOUISIANA: St. Tammany
Parish, 1 female. TENNESSEE: Henderson County, 1 male, 1 female.
DIAGNOSIS: A moderate sized species of the genus, with carapace 0.60-
0.75 mm, with palpal femur 0.45-0.55 mm, and chela 0.70-0.85 mm in length;
with length/depth ratio of chelal hand 1.4-1.75; and with distinct granules
on surfaces of all palpal segments.
DESCRIPTION: Hoff's description (1949:450-452) of the neotype and
several other specimens from Illinois is generally adequate. However, a
The Florida Entomologist
5
Fig. 3-9. Americhernes oblongus (Say): 3) Genital opercula of male;
4) Genital opercula of female; 5) Spermathecae of female; 6) Dorsal view
of left palp; 7) Lateral view of palpal trochanter; 8) Lateral view of right
chela; 9) Leg IV.
few important additions can be made, based upon careful study of the many
individuals mentioned above. Aside from differences of the genitalia, males
and females are quite similar, the latter being on the average a little larger.
Therefore, most of the description applies to both sexes alike.
Surface of carapace entirely without granules, being mostly smooth
Vol. 59, No. 2
Muchmore: New Pseudoscorpion Genus
and shiny, but finely lined laterally; eyespots weak, but always notice-
able; the statement of Hoff to the contrary, only a single transverse fur-
row is visible, lying just at middle of carapace; about 40-50 delicate vesti-
tural setae, mostly terminally or subterminally denticulate, like those
of other parts of body and appendages. Abdominal tergites and sternites
mostly smooth, but with occasional weak sculpturing; pleural membranes
smoothly, longitudinally striate; typical tergal chaetotaxy 10:11:12:12:-
14:15:16:16:16:17:T4T5T4T:2; typical chaetotaxy of 5th-12th sternites-
14:17:18:18:20:20:T3T5T3T:2. Male with anterior genital operculum bear-
ing 4 long, heavy setae surrounded by about 14 smaller ones; posterior
operculum with 2 pairs of small setae just inside anterior margin and about
8 setae on face (Fig. 3). Female anterior genital operculum with a compact
group of about 10 setae near center and 2 sets of 2-3 setae posterior to this;
just behind the compact group of setae are 2 conspicuous, obliquely ori-
ented lyrifissures, and extending posteriorly between the 2 small sets of
setae is a heavily sclerotized area, more or less longitudinally striated
and roughened; posterior operculum with about 8 setae in row along pos-
terior margin (Fig. 4). Anterior spiracular plates with 3-4 and posterior
plates with 1-2 small setae; these setae, as well as those of anal plates,
finely denticulate subterminally. Internal genitalia of male generally
typical of the family, without any particular modifications as far as is
known now. Spermathecae of female in form of 2 separate, curved, saclike
tubes, each with a conspicuous band of gland openings toward the base (Fig.
5).
Chelicera about 0.33 as long as carapace; hand with 5 setae, sb, b, and
es finely denticulate near tips; flagellum of 3 setae, the distalmost larger
and serrate along one edge; galea with 5-6 rami in the distal half, that of
the male only slightly less well developed than that of female.
Palps relatively short and heavy (Fig. 6); femur about 0.75 and chela
about 1.2 as long as carapace; in males (females) femur 1.95-2.3(2.1-2.4),
tibia 1.8-1.9(1.8-2.0) and chela without medical 2.3-2.6(2.4-2.7) times as long
as broad; hand 1.4-1.75(1.5-1.75) times as long as deep; movable finger 0.79-
0.88(0.79-0.89) as long as hand; trochanter with large, conspicuous protu-
berance on dorsal side (Fig. 7). Surfaces of segments generally smooth and
shiny, but with areas of fine granules over most of trochanter, on medial
and lateral sides of femur, on medial side of trochanter, and on medial
and lateral sides of chelal hand at base of fingers. Trichobothria positioned
as shown in Fig. 8; est and ist at about same level on fixed finger; it close to
ist and farther from finger tip than distance between ist and isb. Fixed finger
of chela with 20-25 and movable finger with 25-30 marginal teeth; each
finger with 3-6 external and 1-3 internal accessory teeth.
Legs moderately slender; leg IV (Fig. 9) with femur 2.6-3.0 and tibia
3.1-3.6 times as long as deep. Tibia of leg IV (and III) with 2 long, tactile
setae on outer margin, 1 near middle and other very near distal end; tarsus
of leg IV (and III) with a long tactile seta about 0.25 length of segment
from proximal end. Each tarsus with an elevated slit sensillum on outer
margin close to proximal end; subterminal tarsal setae curved, simple.
MEASUREMENTS (mm): Inclusive of both males and females; males
are usually slightly smaller than females but there is much overlap of the
size ranges for the sexes: Body length 1.7-2.8; carapace length 0.605-0.72.
Chelicera 0.20-0.25 by 0.10-0.12. Palpal femur 0.445-0.54 by 0.20-0.25; tibia
The Florida Entomologist
0.42-0.48 by 0.22-0.27; chela (without pedicel) 0.70-0.835 by 0.29-0.34; hand
(without pedicel) 0.415-0.49 by 0.25-0.32; pedicel about 0.06 long; movable
finger 0.33-0.415 long. Leg IV: entire femur 0.43-0.51 by 0.155-0.185; tibia
0.31-0.37 by 0.095-0.115.
REMARKS: In addition to the states mentioned above, this species has
been reported from Arkansas, District of Columbia, Georgia, Indiana, Ken-
tucky, Michigan, Missouri, Nebraska, Ohio, Pennsylvania, Texas and Vir-
ginia (Hoff 1958) and from Colorado (Hoff 1961). Whether all these records
actually pertain to Americhernes oblongus is not certain at this time.
Americhernes longimanus Muchmore, new species
Fig. 10-13
MATERIAL: Holotype female (WM3493.01001) and paratype male
taken in light traps at the Archbold Biological Station, Highlands County,
Florida, 27-XI-1971 and 16-III-1967, respectively (S. W. Frost); 1 female
paratype from a bolt of Pinus elliottii at Gainesville, Alachua County,
Florida, 2-1-1966 (R. E. Woodruff); 1 female and 1 tritonymph paratype
from Marion County, Florida, 19-1-1948 (H. K. Wallace); 1 female para-
type from Horn Island, Jackson County, Mississippi, 21-IV-1965 (E. A.
Richmond). The types are deposited in the Florida State Collection of
Arthropods, Gainesville.
DIAGNOSIS: A moderate sized species of the genus, a little larger
than A. oblongus, with carapace 0.75-0.82 mm, palpal femur greater than
0.55 mm and chela greater than 0.85 mm in length; and with chelal hand
noticeably long and narrow, the length/depth ratio about 2.0 or greater.
DESCRIPTION OF FEMALE: With the characters of the genus as de-
fined above. Carapace quite smooth and shiny; without any transverse fur-
row; no eyes apparent; with about 70-75 finely denticulate, vestitural setae,
4 at anterior and 8-10 at posterior margin. Abdomen elongate; tergites 3-10
and sternites 4-10 divided; surfaces smooth; pleural membranes smoothly,
longitudinally striate. Tergal chaetotaxy of holotype 14:13:13:20:19:20:-
20:21:24:22:T4TT4TT3T:2, others similar. Sternal chaetotaxy of holotype
15:(3)10(3):(1)8(1):18:21:21:21:20:24:T3TT2TT3T:2, others generally sim-
ilar; anterior operculum with a central, compact group of 9-14 small setae
and 2 sets of 2-4 setae posterior to this, and with 2 slightly oblique lyrifis-
sures and a median longitudinal sclerotized area, as in A. oblongus (Fig. 4);
setae on spiracular and anal plates terminally denticulate. Spermathecae
as shown in Fig. 10; the conspicuous gland openings are confined to the com-
mon base of the 2 tubules.
Chelicera typical; about 0.3 as long as carapace; hand with 5 setae,
sb, b, and es finely denticulate; galea long and with 5-6 small rami; fla-
gellum of 3 setae, the distal 1 serrate along anterior margin; serrula ex-
terior with about 18 blades.
Palps fairly robust (Fig. 11); femur about 0.8 and chela about 1.15 as
long as carapace; femur 2.25-2.45, tibia 1.85-2.1, and chela (without pedicel)
2.55-2.8 times as long as broad; hand (without pedicel) 1.95-2.1 times as
long as deep; movable finger 0.61-0.72 as long as hand. Surfaces of seg-
ments generally smooth but with areas of small granules on lateral side
of trochanter, entire medio-ventral side of femur, medial side of tibia at
distal end, and medial side of chelal hand at base of fingers. Trichobothria
Vol. 59, No. 2
Muchmore: New Pseudoscorpion Genus
13
Fig. 10-13. Americhernes longimanus, new species: 10) Spermathecae of
female; 11) Dorsal view of left palp; 12) Lateral view of right chela;
13) Leg IV.
as shown in Fig. 12; on fixed finger ist at or very slightly proximad of level
of est; it close to ist and farther from finger tip than distance between ist
and isb. Fixed finger of chela with 22-24 and movable finger with 26-28 con-
tiguous, marginal teeth; each finger with 5-7 external and usually 2 in-
ternal accessory teeth.
Legs robust; leg IV (Fig. 13) with entire femur 2.4-2.55 and tibia 2.7-2.9
times as long as deep. Tibia of leg IV with 2 tactile setae on outer margin,
The Florida Entomologist
a longer 1 near middle and a shorter 1 near distal end, both of which are
finely denticulate; tarsus IV with a very long, acuminate, tactile seta about
0.25 length of segment from proximal end. Each tarsus with an elevated
slit sensillum on outer margin close to proximal end; subterminal tarsal
setae curved, simple.
MALE: Very similar to female in all details except the genitalia. An-
terior genital operculum with 6 long, heavy setae surrounded by 17 smaller
ones; posterior operculum with 2 sets of 3 small setae just inside anterior
margin and 13 setae on face. Galea on chelicera with 6 small rami, as well
developed as in female. Palpal femur 2.1, tibia 1.8, and chela 2.5 times as
long as broad; hand 1.9 times as long as deep; movable finger 0.64 as long
as hand.
MEASUREMENTS (mm): Ranges for the 4 females are given first, fol-
lowed in parentheses by figures for the male. Body length 2.2-3.5(3.0). Cara-
pace length 0.755-0.815(0.82). Chelicera 0.23-0.25(0.235) by 0.12-0.13(0.12).
Palpal femur 0.555-0.62(0.63) by 0.245-0.27(0.295); tibia 0.51-0.58(0.57) by
0.265-0.31(0.32); chela (without pedicel) 0.85-0.955(0.93) by 0.325-0.36(0.37);
hand (without pedicel) 0.555-0.61(0.615) by 0.28-0.31(0.325); movable finger
0.35-0.435(0.385) long. Leg IV: entire femur 0.52-0.58(0.57) by 0.215-0.235
(0.26); tibia 0.36-0.41(0.415) by 0.125-0.145(0.15); tarsus 0.21-0.26(0.25) by 0.08-
0.09(0.09).
ETYMOLOGY: The species is named longimanus for the relatively
long hand of the palpal chela.
Americhernes reductus Muchmore, new species
Fig. 14-17
MATERIAL: Holotype male (WM2237.01001) and 6 paratypes (4 fe-
males, 2 tritonymphs) from Rattlesnake Lumps, NE of Sugarloaf Key,
Monroe County, Florida, 7-VIII-1970 (D. Simberloff); 12 male and 4 fe-
male paratypes from several of the Florida Keys, Monroe County, Florida
(R. Silberglied, D. Simberloff). The type specimens are deposited in the
Florida State Collection of Arthropods, Gainesville.
DIAGNOSIS: A small species of the genus, with carapace 0.52-0.64,
palpal femur less than 0.50 mm and chela usually 0.75 mm or less in
length; with trichobothrium it at same level as est on fixed chelal finger;
and with only 4 setae on hand of chelicera.
DESCRIPTION OF MALE (based on 7 mounted specimens, including
holotype): With the characters of the genus as defined above. Carapace
smooth and shiny; with a faint transverse furrow at middle; 2 faint eye-
spots; about 50 finely denticulate vestitural setae, 4 at anterior and 6-10
at posterior margin. Abdomen elongate; tergites 3-10 and sternites 4-10 di-
vided; surfaces smooth; pleural membranes smoothly, longitudinally
striate. Tergal chaetotaxy of holotype 8:10:8:11:12:11:15:15:16:17:-
T3TTTT3T:2 (others similar). Sternal chaetotaxy of holotype 19:(2)2-2
(2):(1)6(1):13:14:15:15:17:14:T2T2T2T:2 (others similar); anterior opercu-
lum with 3 long setae surrounded by 16 shorter ones (other paratypes usu-
ally with 4 long setae); setae on spiracular and anal plates terminally
denticulate. Internal genitalia typical.
Chelicera generally typical; about 0.3 as long as carapace; hand with
only 4 setae (apparently sb wanting), b and es finely denticulate; galea
Vol. 59, No. 2
Muchmore: New Pseudoscorpion Genus
16 \ 17
Fig. 14-17. Americhernes reductus, new species: 14) Dorsal view of right
palp; 15) Lateral view of right chela; 16) Genital opercula of female;
17) Spermathecae of female.
thin, with 5 small rami; flagellum of 3 setae; serrula exterior with 16 or
17 blades.
Palps fairly robust (Fig. 14); femur about 0.8 and chela about 1.2 as long
as carapace; femur 2.3-2.5, tibia 1.95-2.15, and chela (without pedicel) 2.8-
3.1 times as long as broad; hand (without pedicel) 2.0-2.05 times as long as
deep; movable finger 0.74-0.81 times as long as hand. Surfaces of segments
generally smooth, but with small areas of tiny granules on trochanter
and medial parts of femur and tibia. Trichobothria as shown in Fig. 15; on
fixed finger it is at level of est, much farther from finger tip than distance
between ist and isb; ist not far proximad of it. Fixed finger of chela with 25-27
and movable finger with 30-32 contiguous marginal teeth; each finger with
2-3 internal and external accessory teeth.
The Florida Entomologist
Legs robust; leg IV with entire femur 2.65-2.85 and tibia 3.0-3.3 times as
long as deep. Tibia of leg IV with 2 tactile setae on outer margin, both only
moderately elongated and denticulate, 1 near middle, the other near distal
end (the 1 at middle sometimes reduced in size); tarsus IV with a very long,
acuminate, tactile seta about 0.25 length of segment from proximal end.
Each tarsus with an elevated slit sensillum near proximal end.
FEMALE (based on 4 mounted specimens): Very similar to male but
slightly larger. Anterior genital operculum (Fig. 16) with a central group
of 5-7 small setae and 2 sets of 3 setae posterior to this, and with 2 oblique
lyrifissures and a median, longitudinal, sclerotized area, much as in A. ob-
longus; posterior operculum with row of 5-7 setae on posterior margin.
Spermathecae as shown in Fig. 17, rather short tubes with long, cylindrical
expanded ends and no gland openings evident anywhere. Galea a little
better developed than in male.
MEASUREMENTS (mm): Ranges of 7 mounted males given first, fol-
lowed in parentheses by ranges for 4 mounted females: Body length 1.7-
2.25(2.0-2.7). Carapace length 0.52-0.605(0.58-0.64). Chelicera 0.15-0.19(0.185-
0.19) by 0.08-0.10(0.095). Palpal femur 0.435-0.49(0.44-0.50) by 0.18-0.215-
(0.19-0.21); tibia 0.39-0.46(0.41-0.47) by 0.20-0.22(0.20-0.23); chela (without
pedicel) 0.64-0.74(0.70-0.78) by 0.21-0.26(0.24-0.27); hand (without pedicel)
0.37-0.44(0.41-0.48) by 0.19-0.24(0.19-0.24); movable finger 0.295-0.33(0.30-
0.35) long. Leg IV: entire femur 0.37-0.445(0.41-0.46) by 0.13-0.16(0.15-0.17);
tibia 0.26-0.30(0.28-0.32) by 0.08-0.095(0.10); tarsus 0.18-0.215(0.19-0.21) by
0.05-0.065(0.06-0.07).
ETYMOLOGY: The species is named reductus because of the reduced
number of setae on the cheliceral hand.
REMARKS: All of the specimens of A. reductus mentioned above, ex-
cept 1, were taken from under the bark of standing red mangrove trees on
small islands of the Florida Keys; most of the islands possessed no supra-
tidal ground at all, only the trees. The single excepted specimen was taken
from under the bark of a fallen log in the center of Key Largo.
A female specimen which undoubtedly belongs to this species was col-
lected by D. Simberloff from under dead bark of a red mangrove tree on a
small, groundless island at Landivar, Belize City, Belize, 2-II-1970. This
record makes it appear likely that A. reductus is widespread in mangrove
growths throughout the Caribbean area.
Americhernes puertoricensis Muchmore, new species
Fig. 18-20
MATERIAL: Holotype female (WM1828.01001) and 3 female para-
types from Rio Piedras, Puerto Rico, 14-III-1959 (W. M. Nadler). Types
are in the collection of the American Museum of Natural History, New
York.
DIAGNOSIS: A small species of the genus with carapace 0.50-0.55 mm,
palpal femur less than 0.40 mm and chela less than 0.62 mm in length;
with it at same level as est on fixed chelal finger; and with 5 setae on hand
of chelicera.
DESCRIPTION OF FEMALE: With the characters of the genus. Cara-
pace smooth; with a shallow transverse furrow at about middle; 2 faint
eyespots present; about 40 finely denticulate, vestitural setae, 4 at anterior
Vol. 59, No. 2
Muchmore: New Pseudoscorpion Genus
18 "
) 22
21
Fig. 18-22. Americhernes spp. A. puertoricensis, new species: 18) Sper-
mathecae of female; 19) Dorsal view of right palp; 20) Lateral view of left
chela; 21) A. ellipticus (Hoff). Spermatheca of female (other one obscured);
22) A. levipalpus (Muchmore). Spermatheca of female (other one obscured).
margin and 6-8 at posterior margin. Abdomen elongate; tergites 2-10 and
sternites 4-10 divided; surfaces smooth; pleural membranes smoothly,
longitudinally striate. Tergal chaetotaxy of holotype 10:8:9:16:15:14:16:-
17:18:19:T3TTTT4T:2; others similar. Sternal chaetotaxy 13:(3)5(2):(1)6-
(1):15:17:20:18:18:18:T3T2T3T:2; others generally similar; anterior oper-
culum with a central group of 6-8 small setae and 2 sets of 3-4 setae pos-
terior to this, and with 2 oblique lyrifissures and a median longitudinal
sclerotized area, much as in A. oblongus (Fig. 4); posterior operculum with
marginal row of 4-6 small setae. Spermathecae as shown in Fig. 18, each
consisting of a narrow tube and an expanded sac of equal length, with con-
spicuous gland openings at junction of tube and sac.
Chelicera typical; about 0.33 as long as carapace; hand with 5 setae,
sb, b and es finely denticulate; galea long, with 5 or 6 rami; flagellum of
3 setae; serrula exterior with 16-17 blades.
Palps fairly robust (Fig. 19); femur about 0.7 and chela about 1.15 as
long as carapace; femur 2.2-2.3, tibia 1.8-2.0, and chela (without pedicel)
2.7-2.8 times as long as broad; hand (without pedicel) 1.75-1.9 times as long
as deep; movable finger 0.80-0.85 as long as hand. Surfaces of segments
The Florida Entomologist
generally smooth, but with few small granules on trochanter and on
medial sides of femur and tibia. Trichobothria as shown in Fig. 20; on fixed
finger it at level of est, much farther from finger tip than distance between
ist and isb; ist not far proximad of it. Fixed finger of chela with about 24-25
and movable finger with 28-31 contiguous, marginal teeth; each finger with
1-2 internal and 3-5 external accessory teeth.
Legs robust; leg IV with entire femur 2.4-2.55 and tibia 2.8-3.0 times as
long as deep. Tibia of leg IV with 2 denticulate tactile setae on outer mar-
gin, 1 near middle and 1 near distal end, the former longer than the latter;
tarsus IV with a very long, acuminate tactile seta about 0.25 length of seg-
ment from proximal end. Each tarsus with an elevated slit sensillum near
proximal end.
MALE: Unknown.
MEASUREMENTS (mm): Female (ranges for the 4 types): Body
length 1.95-2.0. Carapace length 0.50-0.55. Chelicera 0.17-0.18 by 0.08-0.09.
Palpal femur 0.35-0.38 by 0.155-0.17; tibia 0.33-0.36 by 0.17-0.18; chela (with-
out pedicel) 0.585-0.605 by 0.215-0.22; hand (without pedicel) 0.325-0.34 by
0.18-0.19; movable finger 0.27-0.29. Leg IV: entire femur 0.33-0.36 by 0.13-
0.15; tibia 0.23-0.26 by 0.08-0.09; tarsus 0.18-0.185 by 0.06.
ETYMOLOGY: This species is named for Puerto Rico, where it has been
found.
Americhernes ellipticus (Hoff), new combination
Fig. 21
Lamprochernes ellipticus Hoff, 1944:1; 1956:5; 1958:20.
DIAGNOSIS: A large species of the genus, with carapace longer than
0.95 mm, palpal femur longer than 0.75 mm, and chela longer than
0.25 mm.
The holotype female of this species (from Baja California, Mexico) has
been examined and compared with other species of the genus Americhernes.
Along with other features, the spermathecae (1 of which is illustrated in
Fig. 21) conform well with the general pattern. On the other hand, the sper-
mathecae differ conspicuously from those of the specimens from New Mex-
ico mentioned by Hoff (1956:6), which were described as "hammer-shaped
or, perhaps more appropriately, have an appearance resembling the median
longitudinal section of a mushroom." I have also examined some of these
specimens and quite agree with Hoff's description of their form. Such ham-
mer-shaped spermathecae are commonly found in species of Lustrochernes
or Cordylochernes (unpublished observations). It seems likely, therefore,
that these New Mexican specimens do not really belong to A. ellipticus
even though they resemble that species in many morphological characters.
Further study of related forms in the southwestern United States and in
Mexico is necessary before this problem can be resolved.
Americhernes levipalpus (Muchmore), new combination
Fig. 22
Lamprochernes levipalpus Muchmore, 1971:327.
DIAGNOSIS: Similar in many respects to A. oblongus, but with nearly
smooth palps.
Vol. 59, No. 2
Muchmore: New Pseudoscorpion Genus
The spermathecae of the female paratype was not mentioned in the orig-
inal description. To allow easy comparison with other species, the sper-
matheca is illustrated in Fig. 22. It is similar to that of A. oblongus, but the
band of gland openings is around the base of the bulbous enlargement
rather than on the tubule.
Despite extensive recent collection of pseudoscorpions in Utah by
George F. Knowlton, no further representatives of this species have been
found.
ACKNOWLEDGMENTS
Many thanks are due to all those who have made specimens available
for study. I am indebted to C. H. Alteri for the execution of the illustra-
tions. This work was supported in part by a research grant (GB 37570) from
the National Science Foundation.
LITERATURE CITED
BEIER, M. 1932. Zur Kenntnis der Lamprochernetinae (Pseudoscorp.) Zool.
Anz. 97:258-67.
BEIER, M. 1932b. Pseudoscorpionidea II. Subord. Cheliferinea. Tierreich
58:1-294.
ELLINGSEN, E. 1909. Die Pseudoscorpione des Berliner Museum. Mitt.
Zool. Mus. Berlin 4:355-423.
EWING, H. E. 1911. Notes on pseudoscorpions; a study on the variations
of our common species, Chelifer cancroides, with systematic notes on
other species. J. New York Ent. Soc. 19:65-81.
HOFF, C. C. 1944. New pseudoscorpions of the subfamily Lamprocherne-
tinae. Amer. Mus. Novitates 1271:1-12.
HOFF, C. C. 1945. New pseudoscorpions from Arkansas. Trans. Amer. Micros.
Soc. 64:34-57.
HOFF, C. C. 1949. The pseudoscorpions of Illinois. Bull. Illinois Nat. Hist.
Surv. 24:409-98.
HOFF, C. C. 1956. Pseudoscorpions of the family Chernetidae from New
Mexico. Amer. Mus. Novitates 1800:1-66.
HOFF, C. C. 1958. List of the pseudoscorpions of North America north of
Mexico. Amer. Mus. Novitates 1875:1-50.
HOFF, C. C. 1961. Pseudoscorpions from Colorado. Bull. Amer. Mus. Nat.
Hist. 122:409-64.
HOFF, C. C., AND J. E. BOLSTERLI. 1956. Pseudoscorpions of the Mississippi
River drainage basin area. Trans. Amer. Micros. Soc. 75:155-79.
MUCHMORE, W. B. 1971. A new Lamprochernes from Utah. Ent. News 82:-
327-9.
MUCHMORE, W. B. 1972. Observations on the classification of some Euro-
pean chernetid pseudoscorpions. Bull. British Arach. Soc. 2:112-5.
MUCHMORE, W. B. 1974. Clarification of the genera Hesperochernes and
Dinocheirus (Pseudoscorpionida, Chernetidae). J. Arachnol. 2:25-36.
SAY, T. 1821. An account of the Arachnides of the United States. J. Acad.
Nat. Sci. Philadelphia 2:59-82.
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THE
FIOWING
LINE
P"ll"~e"
1111501ft "Wam I
MVdiG2E:M
The Florida Entomologist
APPLICATION OF A THERMODYNAMIC MODEL FOR THE
DEVELOPMENT OF LARVAE OF THE CABBAGE
LOOPER (LEPIDOPTERA: NOCTUIDAE) AT
FLUCTUATING TEMPERATURES12
G. D. BUTLER, JR.,3 R. E. STINNER,4 AND G. L. GREENE5
ABSTRACT
Rates of development of the larval stage of the cabbage looper, Tri-
choplusia ni (Huibner), at numerous constant temperatures were used in an
algorithm to describe the rate in relation to temperature. Observations at
different fluctuating temperatures confirmed that the algorithm can be
used to predict development at fluctuating temperatures. The technique
can be used in field studies and in computer simulations.
The accurate simulation of insect growth is critical in the develop-
ment of models that can predict changes in populations and produce opti-
mization of pest control. Butler and Watson (1974) reported a technique
for estimating the rate of development of Lygus hesperus Knight at fluc-
tuating temperatures in which one can use developmental rates at constant
temperatures in a computer program (WATBUG). We discuss here the use
of WATBUG when a new temperature-dependent algorithm was used to
predict the development of the larval stage of the cabbage looper, Tri-
choplusia ni (Hiibner), at fluctuating temperatures.
METHODS AND MATERIALS
The duration of the larval stage of the cabbage looper in the labora-
tory in relation to 14 constant and 12 fluctuating temperatures was deter-
mined by Jackson et al. (1969), Fye and McAda (1972), and Toba et al.
(1973). Also, Butler et al. (1975) reared several strains of loopers (field,
laboratory, and a dark mutant) in individual cups of medium (Henneberry
and Kishaba 1966) and obtained 40 sets of observations at 12 constant tem-
peratures. The results obtained by all these workers were used to calculate
the development rates in relation to temperature by using the algorithm
developed by Stinner et al. (1974). The sigmoid function used for this algo-
rithm is:
R,=C/ (l+eA+ *'),
where
R,= rate of development (1/time) at temperature t
C = (maximum development rate) X (eA"R' oPt), i.e., the asymptote
'In cooperation with the Arizona Agricultural Experiment Station.
2Mention of a commercial or proprietary product in this paper does not constitute an en-
dorsement of this product by the USDA.
3Western Cotton Research Laboratory, ARS-USDA, Phoenix, Arizona 85040.
4Department of Entomology, North Carolina State University, Raleigh, North Carolina
27607.
'University of Florida Agricultural Research and Education Center, Quincy, Florida 32351.
Vol. 59, No. 2
The Florida Entomologist
topt = temperature at which the maximum developmental rate occurs
A, B= empirical constants
t'= t, for t 5 topt
t'= 2 tops t, for t > topt
Listings of the program in FORTRAN and for a Wang 7202 computer
are available from the senior author.
In addition, Butler et al. (1975) reared cabbage loopers during the winter
months of 1972-73 in an outside insectary at Phoenix, Arizona. Previously,
larval development on artificial medium (Henneberry and Kishaba 1966)
was observed by G. L. Greene in temperature cabinets programmed for fluc-
tuating temperatures and in a U. S. Weather Bureau thermograph shelter at
Orlando, Florida during the fall, winter, and spring of 1970-71. During the
same period, C. S. Creighton6 at Charleston, South Carolina, also reared
larvae in a thermograph shelter. The larvae used in Florida and South
Carolina had originated from cabbage plants and had been reared in the
laboratory for more than a year. Each test group began with 10 newly
hatched larvae in 1-oz plastic cups, and 10 cups were started for each group
with 5 larvae 1/2-in. or larger the maximum number held in a single cup.
The results obtained by all these workers were used to test the accuracy of
the predictions of the computer program.
RESULTS AND DISCUSSION
The duration of the larval stage of the cabbage looper can be estimated
from the algorithm of Stinner et al. (1974) by using the following values:
A = 4.9848, B = -0.2247, C = 0.112, TOPT = 33.00C.
The experimental observations used to determine these values and the
rates calculated from the algorithm at 2-hr intervals are shown in Fig. 1.
These rates were used in the WATBUG program with the 2-hr fluctuating
temperatures observed in the laboratory (Fye and McAda 1972, Toba et al.
1973, and GLG), in the insectary at Phoenix (GDB), in the field at Orlando
(GLG), and in the field at Charleston. The observed and calculated dura-
tions of the larval stage at these different fluctuating temperatures were
similar (Fig. 2) on the basis of chi-square tests for each group of observa-
tions. The chi-square for all the 45 observations was 15.2, which is not sig-
nificant (P > 0.99). Thus the values calculated by the algorithm from de-
velopment data obtained at various constant temperatures provided a use-
ful estimate of the duration of the larval stage of the cabbage looper at
fluctuating temperatures. Although the use of the rates calculated by the
algorithm are similar to those obtained from a linear regression equation
for most of the temperature range, the algorithm provides a much better
estimate of development at low temperatures. In comparing the results of
the 2 methods at 12 fluctuating temperatures in the laboratory, the chi-
square estimates provided by the algorithm were 3-times smaller than those
from the linear equation.
A large computer program such as WATBUG is not essential for the
utilization of the developmental rates. Those shown in Fig. 1 show the pro-
'The authors sincerely thank C. S. Creighton of the Vegetable Insects Laboratory, ARS-
USDA, Charleston, S. Car. for providing this information.
Vol. 59, No. 2
166
Butler et al.: Model of Cabbage Looper Larva
.10 -
.09
.08 -
.07- -
n .06-
m .05-
S.04-
.03
.02
.01 -
0-
0 4 8 12 16 20 24 28 32 36
Temperature C
Fig. 1. Observed (*) and predicted (-) rates of development of cabbage
looper larvae at constant temperatures.
portion of development that occurs in 1 day at different temperatures.
Thus, at a constant temperature of 320C, the rate is approximately 0.10;
therefore, 10% of the development takes place each day, that is, the stage
will last 10 days. At fluctuating temperatures where the hourly tempera-
tures are known, the values can be scaled, and the percentage of develop-
ment for each hour summed so on the day and hour that the sum equals or
exceeds 1.0, the stage will have been completed. For applications in which
only the maximum and minimum temperatures are known, the senior author
has a FORTRAN program that contains an algorithm for the calculation
of hourly temperatures by using daily maximum-minimum temperatures,
latitude, longitude, and day of the year.
Obviously when hourly development rates are being summed for ex-
tended periods of cool weather, a simple mini-computer program that uti-
lizes the cited equation or a table of values will greatly simplify the task
and will reduce errors. Furthermore, such a program can easily be ar-
The Florida Entomologist
Laboratory Field
F Fye 0 Orlando
G Graham S Charleston
T Toba 0 Phoenix
100
80
60
40
20
0
0 20 40 60 80
100
Calculated Number of Days
Fig. 2. Observed and calculated duration of the larval stage of the cab-
bage looper at fluctuating temperatures.
ranged to convert the Fahrenheit temperatures of our thermometers as in-
put in Centigrade values generally used in scientific literature. Thus, rear-
ing studies at constant temperatures, particularly in the high and low tem-
perature ranges, provide development rates in relation to temperature
which provide the field entomologist with a comparatively simple method
of following the daily development of a particular stage of an insect. When
this information for several stages of an insect is combined with a book-
keeping method to keep track of numbers, it provides a basis for a computer
simulation of the population such as the WATBUG program.
LITERATURE CITED
BUTLER, G. D., JR., AND F. L. WATSON. 1974. A technique for determining
the rate of development of Lygus hesperus in fluctuating tempera-
tures. Fla. Ent. 57:225-30.
0
Q)
"0
Q)
o
0
Vol. 59, No. 2
Butler et al.: Model of Cabbage Looper Larva
BUTLER, G. D., JR., A. G. HAMILTON, AND A. C. BARTLETT. 1975. Develop-
ment of the dark strain of cabbage looper in relation to temperature.
Environ. Ent. 4:619-20.
FYE, R. E., AND W. C. McADA. 1972. Laboratory studies on the develop-
ment, longevity, and fecundity of six lepidopterous pests of cotton
in Arizona. USDA Tech. Bull. 1454. 73 p.
JACKSON, C. G., G. D. BUTLER, JR., AND D. E. BRYAN. 1969. Time required
for development of Voria ruralis and its host, the cabbage looper,
at different temperatures. J. Econ. Ent. 62:69-70.
HENNEBERRY, T. J., AND A. N. KISHABA. 1966. Pupal size and mortality,
longevity, and reproduction of cabbage loopers reared at several
densities. J. Econ. Ent. 59:1490-3.
STINNER, R. E., A. P. GUTIERREZ, AND G. D. BUTLER, JR. 1974. An algorithm
for temperature-dependent growth rate simulation. Can. Ent. 106:
519-24.
TOBA, H. H., A. N. KISHABA, R. PANGALDEN, AND P. V. VAIL. 1973. Temper-
ature and the development of the cabbage looper. Ann. Ent. Soc.
Am. 66:965-74.
BOOK REVIEW
ECOLOGICAL GENETICS. E. B. Ford. 1975. 4th edition. Chapman and Hall,
London; John Wiley and Sons, Halsted Press, New York. xx + 442 p. $32.00.
The fourth edition of this well known and widely cited book is organ-
ized along the same 15 chapter titles that appeared in the first edition, 1964.
A book treating technical and complex processes and addressed to an audi-
ence presumed to have a wide background in population genetics and evo-
lution must, by any values, be judged successful when it has appeared in 4
editions, the first 3 with a number of printings, in a period of 11 years. The
essence of this success appears to stem from the fact that Ford has largely
by-passed theory and developed his case for evolutionary processes from
studies conducted directly in the field, supported by laboratory breeding
tests, etc. The methods clearly appeal to a wider range of biologists than
the more esoteric treatment of mathematics that generally characterize
population genetics. The first chapter attempts to define ecological genet-
ics and distinguish it from related areas such as evolutionary genetics, pop-
ulation genetics, etc. Although Ford spends about 12 pages drawing this
distinction, one can summarize by saying that it is an analysis of variation
in natural populations seeking to explain its origin and maintenance.
While Ford implies that concepts, generally recognized as those of theo-
retical population genetics are not the object of his discussion, it becomes
clear that he assumes the reader to have some insight on that discipline.
The second, third, and fourth chapters, are largely an expression of Ford's
position on genetic drift versus selection in determining variation. Ford
gives a rather simplified discussion of drift addressed to an individual en-
countering the concept for the first time. On this point, the book is inconsist-
ent because he later discusses a number of equally or more complex proc-
esses with the assumption that the reader has an advanced understanding.
Ford develops a strong case against drift which has apparently won over
many colleagues to his views. While the examples he cites are convincing,
he appears to extrapolate to evolution in general and on this point, many
The Florida Entomologist
students of evolution take exception. Chapter 5, Sympatric Evolution,
summarizes some of the more interesting studies to emerge from the English
School of Ecological Genetics. Ford carries the reader step-by-step through
the investigations that suggest spot patterns evolved separate expressions
in a butterfly population having an essentially sympatric range. Toward
the end of the chapter, the reader is anxiously awaiting the explanation for
the data only to find that Ford can offer no explanation, a situation most
uncharacteristic for this talented author. Chapters 6 through 11 concern
the theory of genetic polymorphism and chromosomal polymorphism. The
principal interest here lies in his explanation for the evolution of hetero-
zygous advantage along the same lines as the evolution of dominance, the
latter which he assumes the student to understand. The development of a
supergene, a number of linked loci controlling related traits, is also de-
scribed in detail. Chapters 12 and 13 concern mimicry and its evolution.
Ford gives a lengthy discussion of the basic theory of mimicry, the support-
ing evidence, and the data for the genetic basis of mimetic patterns. It may
be of interest that Ford appears to feel kin selection is not necessary in ex-
plaining mimicry and that models suffer no significant loss of fitness by
occasional mistakes of the predators. Chapter 14 concerns what Ford terms
transient polymorphism, a process frequently identified as directional se-
lection and his principal examples concern melanism. In Chapter 15, Ford
touches on several topics such as mortality in small populations and speci-
ation; the latter process is given much less space than typical of most
books on evolution.
Ford's writing style often conveys the author's position as clearly as his
written statements and increases the reading interest. Nonetheless a num-
ber of sentences are long and the meaning none too clear. Throughout Ford
notes his reluctance to accept the prediction of mathematical models. How-
ever, he devotes several pages of Chapter 14 to reviewing studies involving
an early model devised by J. B. S. Haldane. Then, he rejects the results be-
cause of an error in Haldane's basic equation. Unfortunately, the citation
used to support the presence of an error is omitted from the References,
though a footnote gives some direction.
The book assumes so much basic knowledge that it is unlikely to be used
as a text and the price seems unduly high. Students of evolution, particu-
larly at the genetic level will nonetheless profit by adding it to their
working libraries. The photographic plates are placed together in the
middle of the book while line-drawn figures appear with the related discus-
sion; the references are largely expanded relative to earlier editions, and
the index is sufficiently complete, allowing the reader to quickly locate
topics of interest. Ford's own research has largely used lepidopterans and
it is an asset to this edition that a wider range of organisms is represented in
the examples. On this point, Ford stresses that he is treating principles and
is not cataloging examples. In conclusion, this book provides an excellent
up-to-date summary of evolutionary studies taking the premise that nat-
ural selection is sufficient for explaining variability. Printing errors in this
book seem to be essentially absent.
Clifford Johnson
University of Florida
Gainesville 32611
Vol. 59, No. 2
The Florida Entomologist
DESCRIPTIONS AND RECORDS OF SOME PHILOTARSIDAE
FROM TRINIDAD, WEST INDIES (PSOCOPTERA)'
EDWARD L. MOCKFORD2 AND HOWARD A. EVANS
Illinois State University, Normal, Illinois,
and University of Leeds, Leeds, England, U. K., respectively
ABSTRACT
Four species of Philotarsidae are recorded from Trinidad, West Indies, 3
of which are new to science. Two new species of Aaroniella are described
and 1 new genus is erected, Broadheadia (type B. caribe new species).
Philotarsus bruchi Williner of Argentina, is transferred to Aaroniella. A
key to the known species of Trinidad Philotarsidae is included.
Despite much collecting and several publications in recent years, the
psocid faunas of South America and the West Indies remain relatively
poorly known. From Trinidad there exist literature records of only 8 spe-
cies of psocids (Mockford 1967, 1971; New 1973; Eertmoed 1973), although
many more species are at hand.
Investigation of the psocids of Trinidad apparently began in February
1959, when the senior author and Mr. Aaron M. Nadler spent 8 days collect-
ing on the island. The senior author returned to Trinidad in August 1961
and spent 3 days collecting psocids there. The Trinidad psocid fauna did
not receive attention again until April 1974, when the junior author began
faunal and ecological investigations there which have lasted for a year.
The family Philotarsidae seems to be poorly represented in South Amer-
ica and the West Indies. A single species has been described from South
America (Williner 1943). One species is known from Hispaniola and Cuba
(Mockford 1974), and 1 is known from Jamaica (Turner 1975).
It is apparent from the field work to date that the Philotarsidae consti-
tute a relatively small, inconspicuous part of the entire psocid fauna of
Trinidad. Four species have been found, represented by only 9 individuals.
Of the 4, 3 are new to science, and 1 represents a new genus. In this paper the
new taxa are described and a key to the species is presented.
Measurements (Table I) are in microns and have an error of 0.231i.
Abbreviations used in the measurements are as follows: Fw=forewing;
Hw= hindwing; f,, f2, f3= first, second, and third flagellomeres; F= posterior
femur + trochanter; T= posterior tibia; t1, t2, t, = posterior first, second and
third tarsomeres; cten.= number of ctenidia on posterior t,; IO/D= least
distance between compound eyes in dorsal view divided by greatest antero-
posterior diameter of compound eye in dorsal view; PO= greatest trans-
verse diameter of compound eye divided by greatest antero-posterior diam-
eter of the eye in dorsal view.
'Contribution No. 339, Bureau of Entomology, Division of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services, Gainesville, Florida 32602.
2Research Associate, Florida State Collection of Arthropods, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services.
Vol. 59, No. 2
The Florida Entomologist
Aaroniella dentata Mockford and Evans, new species
DIAGNOSIS.-With radial pattern of spots in forewing. Except for A.
bruchi (Williner) new combination, differing from other previously de-
scribed species with radial spots by entire border of pterostigma being heav-
ily pigmented. Differing from A. bruchi in absence of spot in M, at apex of
areola postica and longer wing length. Male characterized by absence of
radial spot in R, and toothed posterior margin to clunium.
MALE.-Measurements (Table I).
Head: With epicranial line present; frontal lines absent. Bearing nu-
merous setae with longer ones, each about 3/4 length of f,, on vertex and
anterior margin of clypeus. Antennae bearing setae each about length of
f7. Distal flagellomere terminating in seta longer than the segment. An-
terior ocellus slightly smaller than the others. Lacinial tip (Fig. 4) with
relatively large, concave, median cusp; lateral cusp with a few low,
rounded denticles. Eyes a little larger than those of female.
Thorax: Each pretarsal claw with a basal seta and a pulvillus with a
lightly dilated tip. Pterostigma with a spur vein from its posterior angle.
Abdomen: Clunium (Fig. 11) with 15-20 small denticles, in specimen
examined, either side of median convexity. Epiproct (Fig. 11) with spinous
posterior margin and denticulate lateral lobes. Sense cushion of paraproct
with 21, 18 trichobothria in specimen examined. Hypandrium (Fig. 7) with
gently curved posterior margin dorsally, bearing scattered setae. Phallic
frame (Fig. 8) anteriorly rounded, relatively wide and thick; narrower on
sides. Endophallic sclerotizations complex with 5 denticles along mid-
line. Tip of aedeagal arch tuberculate and ending acuminately.
Color (in alcohol): Head light brown with medium brown mottling on
frons and vertex. Mottling larger and more concentrated around eyes and
bordering very dark brown epicranial line. Ocelli pale brown with very
dark brown interval; entire perimeter of posterior pair very dark brown.
Genae medium brown around antennae, medium grayish brown anteriorly.
Chevrons of clypeus medium brown. Anteclypeus with median, dark gray-
ish brown diffuse patch. Labrum dark grayish brown. Antennae medium
brown with white joints and apical spine. Maxillary palps light brown,
terminal segment medium brown. Eyes black.
Thorax light brown with sclerotized areas medium brown. Anterior half
of mesoscutellum and medio-posterior corners of metanotal lobes very
dark brown. Coxae medium brown, pale brown distally. Femora medium
brown except for pale brown distal tips. Tibiae medium brown with pale
brown proximal tips. Tarsi medium brown. Forewing (Fig. 3) with medium
brown margin and veins except for colorless Cub and Cuia around junc-
tion with Cub. Radial spots pale brown with more distinct, similarly col-
ored spots at vein-margin junctions. Margin and veins of hindwing medium
brown except for pale brown Cu,.
Abdomen covered with scattered dark brown pigment. Dorsally, pig-
ment segmentally concentrated except for very narrow median, unpig-
mented line. Phallic frame medium brown, aedeagus very dark brown; re-
mainder of terminalia dark brown.
FEMALE.-Measurements (Table I).
Head: With epicranial line almost reaching ocellar interval; frontal
lines absent. Bearing 25-30 setae each about length of fs; almost twice this
Vol. 59, No. 2
Mockford and Evans: Trinidad Philotarsidae 173
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The Florida Entomologist
length on posterior margin of vertex and anteriorly on clypeus with 1 on
each side directed forward and about 6-8 along anterior margin directed
downward. Antennae with short setae about 4X width of a flagellomere
in length, distal flagellomere terminating in a seta longer than the seg-
ment. Anterior ocellus of similar size to others. Lacinial tip (Fig. 5) with
relatively large, concave median cusp; lateral cusp with a few low,
rounded denticles.
Thorax: Mesonotal lobes bearing setae of similar size range to those of
head. Each pretarsal claw with basal seta and pulvillus with slightly di-
lated tip. Pterostigma with minute spur vein.
Abdomen: Clunium simple. Epiproct subsemicircular with scattered
setae. Sense cushions of paraprocts with 17, 16 trichobothria in specimen
examined as well as 2 setae without rosettes in each. Subgenital plate (Fig.
9) with arms separated but poorly sclerotized; bearing scattered setae on its
distal third with 2 medial ones about twice as long as others; sculptured
terminal sclerite without setae. Gonapophyses (Fig. 10): relatively short
ventral valve with several minute spinelets on tip; dorsal valve subrec-
tangular with spinous protuberance ventral to apex; lateral valve tri-
angular and shallow with apex a little to ventral half and bearing numer-
ous setae of different lengths. Spermapore plate (Fig. 6) complex with pore
bordered by heavily sclerotized area and a basal area bearing sinuate
bands.
Color (in alcohol): Head light grayish brown with medium brown mark-
ings. Vertex mottled around eyes and dark brown epicranial line. Mottling
much finer on frons with a large median spot. Ocelli pale with very dark
brown inner borders. Clypeus chevroned. Anteclypeus dark gray. Labrum
very dark brown. Genae with dark brown subcuticular (?) pigment. Anten-
nae dark brown, white at joints and tips of terminal spines. Eyes black.
Thorax generally light brown with diffuse dark brown patches on notal
lobes. Medio-posterior corners of latter, anterior half of scutellum, and
posterior margin of meta-postnotum very dark brown. Sclerotized parts of
pleura very dark brown; coxae similar shade with pale brown distal tips.
Femora very dark brown with pale brown tips; tibiae medium brown, tarsi
dark brown. Forewing (Fig. 1) with margin and veins medium brown except
for Culb and Cu,, around junction with Cub. Membrane patches medium
brown. Pterostigma with medium brown pigment bordering a central hya-
line area. Hindwing (Fig. 2) with medium brown margin, veins, and patches
at vein-margin junctions.
Abdomen with diffuse dark brown cuticular and subcuticular pigments,
more concentrated ventrally. Clunium dark brown with a light brown
patch either side of dorsal mid-line. Paraprocts and epiproct dark brown.
Genitalia and subgenital plate very dark brown.
NYMPH.-Essentially same as adult in body shape, ciliation, and color
Fig. 1-10. Aaroniella dentata n. sp. (scales in mm): 1) female, forewing.
2) female, hindwing, scale of Fig. 1. 3) male, forewing, scale of Fig. 1.
4) male, lacinial tip. 5) female, lacinial tip. 6) female, spermapore scler-
ite (distal end up on this and all other terminal abdominal parts except
epiproct and paraproct). Scale of Fig. 9. 7) male, hypandrium, scale of Fig.
9. 8) male, phallosome, scale of Fig. 9. 9) female, subgenital plate. 10) fe-
male, ovipositor valves.
Vol. 59, No. 2
Mockford and Evans: Trinidad Philotarsidae 175
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The Florida Entomologist
pattern, except that head and body paler, legs very light brown, setae of
abdomen grouped in segmental bands.
Holotype male, allotype female, 2 male and 1 female paratypes taken
beating trees, Trinidad (West Indies) Simla, Arima Valley, 24-II-1959, E. L.
Mockford collector. The holotype and allotype will be deposited in the
American Museum of Natural History, New York City. Other records (all
from Trinidad, West Indies, collected by H. A. Evans): 1 female on coffee
(Coffea arabica Linn.), Wells Plantation, Maracas Valley, 19-VI-1974; 1
nymph on cacao (Theobroma cacao Linn.), same location, 25-1-1975; 1
nymph on tonca bean (Dipteryx odorata Willd.), same locality, 27-I-
1975; 1 female on cacao, Sonny Plantation, Ortinola Valley, 25-1-1975;
1 nymph on mango (Mangifera indica Linn.), 2000 ft, Textel Telecommu-
nications Station, Arima-Blanchisseuse Road, 8-XI-1974.
Aaroniella festival Mockford and Evans, new species
DIAGNOSIS.-With radial pattern of spots in forewing (Fig. 16). Ex-
cept for A. bruchi differing from other previously described species with
radial spots by entire border of pterostigma being heavily pigmented. Dif-
fering from A. bruchi in having proximal pigmentation in cell M3 conflu-
ous with that in cells R,, Cu,, and Cu,, forming a large central spot.
FEMALE.-Measurements (Table I).
Head: With epicranial line complete; frontal lines distinct. Anterior
ocellus about half size of posterior pair. Bearing several setae with longer
ones, each about length of f,, on posterior margin of vertex and anterior
margins of clypeus and genae. Anterior setae directed downward except for
1 forward-directed individual on each antero-lateral corner of clypeus. An-
tennae with setae about 3 times as long as width of a segment; terminal seta
longer than remainder of segment. Lacinial tip (Fig. 13) with small,
rounded denticle on median cusp and a few low, rounded denticles on lat-
eral cusp. Eyes slightly triangular in outline.
Thorax: Each pretarsal claw with basal seta and pulvillus with
strongly dilated tip. Pterostigma with no spur vein from its posterior angle.
Hindwing as in Fig. 17.
Abdomen: Clunium simple. Epiproct relatively acutely rounded with 4
long setae (length about half distance across epiproct base) and several
smaller ones. Sense cushions of paraprocts with 15, 17 trichobothria on sin-
gle specimen observed and 2 setae without rosettes in each. Subgenital plate
(Fig. 14) with pigment arms separated but poorly developed; bearing scat-
tered setae except on its distal third; 2 medial setae longer than others.
Gonapophyses (Fig. 18): relatively short ventral valve suddenly tapering
in terminal 1/5 to acuminate tip; dorsal valve with low lobe bearing
minute setae, a large denticle and a few small denticles, near distal end;
Fig. 11-18. Aaroniella n. spp. Fig. 11-12: A. dentata n. sp.: 11) male, epi-
proct and paraproct. 12) female, epiproct and paraproct, scale of Fig. 11.
Fig. 13-18: A. festival n. sp.: 13) female, lacinial tip. 14) female, subgenital
plate. 15) female, spermapore sclerite, scale of Fig. 11. 16) female, fore-
wing. 17) female, hindwing, scale of Fig. 16. 18) female, ovipositor valves,
scale of Fig. 13.
Vol. 59, No. 2
Mockford and Evans: Trinidad Philotarsidae
13
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7
.1
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The Florida Entomologist
lateral valve with dorsal corners irregularly squared-off, bearing numer-
ous setae. Spermapore plate (Fig. 15) highly complex with pore inside scler-
otized plate and loops with crescent-shaped sclerotizations anteriorly.
Color (in alcohol): Head ground color light brown. Vertex with
medium brown mottlings around eyes and similarly colored areas border-
ing dark brown epicranial line. Frons with medium brown triangle with
base adjacent to clypeus. Frontal lines medium brown. Ocellar interval
dark brown with inner borders of otherwise pale ocelli very dark. Genae
light grayish brown. Medium brown chevrons on clypeus. Anteclypeus un-
pigmented. Labrum very dark brown. Maxillary palps light brown with
medium brown terminal segments. Antennae medium brown with white
joints and apical spines. Eyes black.
Thorax dark brown with very dark brown, narrow lateral line just
above coxae. Latter medium brown, pale brown distally. Femora very pale
brown proximally grading to medium shade distally. Tibiae pale brown
with a narrow medium brown band subapically at each of the proximal and
distal tips. Tarsi medium brown. Forewing (Fig. 16) with veins pigmented
medium brown except for Cub, and Cu,, a little before and after junction
with Culb. Radial spots and large patch over junction of M + Cu,, Cu, and
M also medium brown. Pterostigma similarly pigmented except for pale
center. Hindwing (Fig. 17) with medium brown veins and large but faint
patch at junction of M + Cu,, Cu,, and M.
Abdomen unpigmented except for a few small, irregularly scattered
patches of dark brown pigment. Spiracles dark brown. Terminalia dark
brown.
Holotype female, Trinidad (West Indies), Textel Telecommunications
Station Arima-Blanchisseuse Road, 28-IX-1974, H. A. Evans collector.
The type is in the collection of the senior author, Normal, Illinois.
Aaroniella achrysa (Banks)
Graphocaecilius achrysus Banks, 1941:391.
Aaroniella achrysa (Banks), Mockford, 1974:133.
This species was diagnosed and described in detail by Mockford (1974).
It is distinguishable from the other Trinidad species by the accompanying
key. Outside of Trinidad, it is known from Hispaniola, Cuba, and Florida.
Record: 1 female, Belix Plantation, Upper Maracas Valley, on citrus
(Citrus spp.), 25-1-1975, H. A. Evans collector.
Genus Broadheadia Mockford and Evans, new genus
Differing from Aaroniella in (1) microptery, (2) lack of ocelli, (3) both
truncated and pointed setae on body, (4) only 2 tarsal segments, (5) sub-
genital plate with long, stout seta on terminal segment, (6) lacinia with
very large lateral cusp widening toward tip, (7) relatively short antennal
segments.
Type species: Broadheadia caribe new species.
This genus is named for Dr. Edward Broadhead, Reader in Pure and Ap-
plied Zoology at the University of Leeds, England. Dr. Broadhead is well
known for his studies in psocidology.
Vol. 59, No. 2
Mockford and Evans: Trinidad Philotarsidae
Broadheadia caribe Mockford and Evans, new species
FEMALE.-General form as in Fig. 19. Measurements (Table I).
Head: Relatively large with width and length axes more than twice
dimensions of corresponding axes of thorax and 2/3 x those of abdomen.
Epicranial line distinct; frontal lines absent. Setae of head each about
length of f, except 1 pair slightly longer and gently curving about 1/3 the
way along posterior margin of vertex from each eye. Numerous setae di-
rected forward from anterior margins of clypeus and genae. Antennae
(broken) bearing setae about 2-1/2 x width of a flagellomere in length;
t to f, only twice longer than wide. Lacinial tip (Fig. 20) with median cusp
bearing 1 small, median denticle; very large lateral cusp widening toward
tip. Eyes small with black center forming irregular junction with light
brown periphery.
Thorax: Pterothoracic notal lobes poorly developed. Minute fore-
wings only reaching a little over half distance to abdomen. Hindwings ex-
tremely minute. Pointed and truncated setae on terga and forewings. Only
left mesothoracic leg remaining on damaged specimen. Each pretarsal claw
(Fig. 21) with pulvillus slightly dilated at end and a basal seta ventrally.
20
21
rzT^
0.5
0.1
Fig. 19-24. Broadheadia caribe n. gen., n. sp., female: 19) whole speci-
men in dorsal view. 20) lacinial tip. 21) pretarsal claw. 22) epiproct and
paraproct. 23) ovipositor valves. 24) subgenital plate.
The Florida Entomologist
Abdomen: Bearing pointed and truncated setae dorsally and laterally,
very few ventrally. Clunium simple. Epiproct (Fig. 22) semicircular with
about 19 scattered setae, 8 marginal or submarginal including 2 very long
ones posteriorly. Paraproct simple with about 17 scattered setae; nq sense
cushion. Gonapophyses (Fig. 23): ventral valve relatively short with
pointed, curved tip bearing minute spines; dorsal valve narrow with spinous
pointed lobe subapically; lateral valve relatively large; almost equi-
lateral, bearing about 30 scattered setae. Subgenital plate (Fig. 24) with
sclerotized arms separated; well sclerotized posterior sections, without
setae, distinct from poorly-sclerotized, setose anterior sections; terminal
sclerite with relatively long, strong apical seta. Spermapore plate com-
plex but damaged in mounting.
Color (in alcohol): Head light brown with medium brown markings.
Epicranial line dark brown posteriorly. Vertex mottled around eyes and
epicranial line. Frons and anteclypeus light brown, clypeus and labrum
slightly darker. Maxillary palps medium brown with terminal segments
slightly darker in distal 3/4. Antennae medium brown with white joints.
Eyes black.
Thorax light brown with sclerotized parts dark brown. Mesothoracic
femur and coxa light brown; tibia dark brown, paler at tips, and tarsus me-
dium brown. Winglets dark brown.
Abdomen light brown with diffuse, medium brown pigmentation dor-
sally and laterally. Terminalia dark brown.
Holotype female, Trinidad (West Indies) Sonny Plantation, Ortinola
Valley, on cacao, 12-X-1974, H. A. Evans collector. The type is in the col-
lection of the senior author, Normal, Illinois.
DISCUSSION.-Broadheadia is a highly neotenic member of the Philo-
tarsus-Aaroniella-Haplophallus line, as opposed to the Austropsocus-
Zelandopsocus line of Thornton and Smithers (1974). Its closest relative
appears to be Aaroniella, but several features point to a more basic dis-
tinctness than the purely neotenic features. These features are (1) the more
rounded lateral valve (2) acute subapical lobe of dorsal valve, (3) seta of
distal piece of subgenital plate. Several setae are present on the distal piece
of the subgenital plate in A. rawlingsi Smithers (New Zealand) and A. pal-
lida New (Australia), but these 2 species appear to be unique in several fea-
tures and may require a distinct genus or transfer to another established
genus.
KEY TO THE KNOWN SPECIES OF TRINIDAD PHILOTARSIDAE
1. Adults micropterous and with 2 tarsomeres.........................
................................. ..... ............... Broadheadia caribe n. sp.
1'. Adults macropterous and with 3 tarsomeres ........... Aaroniella........2.
2. Forewing with a large spot in basal half on and surrounding
M -C u, division ...................................................................... A festival n. sp.
2'. Forewing with no spot touching M -Cu, division .................................. 3.
3. Pterostigma continuously and heavily pigmented around its
entire margin ................................................ A. dentata n. sp.
3'. Pigmentation of posterior margin of pterostigma not contin-
uous, forming discrete spots ...................................A. achrysa (Banks).
180
Vol. 59, No. 2
Mockford and Evans: Trinidad Philotarsidae 181
ACKNOWLEDGMENTS
The senior author wishes to acknowledge a travel grant from the Amer-
ican Museum of Natural History, New York City, which covered his ex-
penses on his first trip to Trinidad in 1959. The work of the junior author
was carried out during tenure of a N.A.T.O. overseas studentship adminis-
tered by N.E.R.C.
LITERATURE CITED
BANKS, N. 1941, New neuropteroid insects from the Antilles. Mem. Soc.
Cubana Hist. Nat. 15:385-402.
EERTMOED, G. 1973. Phenetic relationships of the Epipsocetae (Psocoptera):
the higher taxa and the species of two new families. Trans. Amer. Ent.
Soc. 99:373-414.
MOCKFORD, E. L. 1967. The Electrentomoid psocids (Psocoptera). Psyche
74:118-65.
MOCKFORD, E. L. 1971. Peripsocus species of the alboguttatus group (Pso-
coptera: Peripsocidae). J. New York Ent. Soc. 79:89-115.
MOCKFORD, E. L. 1974. Records and descriptions of Cuban Psocoptera. Ent.
Americana 48:103-215.
NEW, T. R. 1973. The Archipsocidae of South America (Psocoptera). Trans.
Royal Ent. Soc. London 125:57-105.
THORNTON, I. W. B., AND C. N. SMITHERS. 1974. The Philotarsidae (Psocop-
tera) of New Caledonia. Pacific Ins. 16:177-243.
TURNER, B. D. 1975. The Psocoptera of Jamaica. Trans. Royal Ent. Soc.
London 126:533-609.
WILLINER, G. J. 1943. Psocopteros de Misiones: generous y species neuvos.
Rev. Soc. Ent. Argentina 12:109-21.
PASTURE MOLE CRICKET CONTROL WITH AN APPLICATOR-
FORMULATED BAIT'-(Note). The southern mole cricket, Scapteriscus acletus
Rehn and Hebard, and the change, S. vicinus Scudder, are major pests of pasture-
grass in the southeastern United States (P. G. Koehler and D. E. Short. 1976. J. Econ.
Ent. In press). P. G. Koehler and D. E. Short (1976. J. Econ. Ent. In press) and D. E.
Short and P. G. Koehler (Fla. Ent. In press) have shown that malathion 2% baits can
provide excellent mole cricket mortality when applied at 1.0 to 2.0 lb AI/acre. How-
ever, a major factor in pasture mole cricket control is economics. More than 75%
of the cost of a bait material is the cost of carriers and attractants. The actual toxi-
cant usually accounts for less than 25% of the manufacturer's cost of the ingredients
in baits.
A major economic savings by the farmer could be realized if baits could be formu-
lated on the farm from materials which are readily available. Finely cracked corn
(grits) or commercial laying mash in a crumble form were chosen for testing as car-
rers, and molasses was selected as a suitable attractant (Fla. Dep. of Agric. 1938.
Fla. Bull. of the Dep. of Agric. New Series No. 3). Malathion 2% baits (B) were formu-
lated from the following ingredients:
100 lbs-finely cracked corn or laying mash (crumble)
5 qts-water
64 fl oz-molasses
51 fl oz-malathion (5 lb AI/gal E.C.)
'Fla. Agricultural Experiment Station Journal Series No. 9033.
182 The Florida Entomologist Vol. 59, No. 2
Malathion baits formulated with cracked corn or mash carriers were compared
to a commercially formulated malathion bait (70% peanut hulls, 20% bran, 10%
wheat shorts, and amyl acetate). The formulated baits were tested with 9 other in-
secticidal formulations to determine their efficacy for mole cricket control.
Experiments were conducted on a golf course damaged by mole crickets near
Gainesville, Fla. Plots measured 10 x 10 ft and were separated by 3 ft borders. A
randomized complete block design was used, with 13 treatments replicated 5 times.
Sprays were applied with the premeasured amount of insecticide in 2 gal of water.
Baits were applied carefully by hand to insure complete coverage. Dead or mori-
bund mole crickets found on the soil surface were counted for the first 3 days fol-
lowing treatment. The total number of dead mole crickets was analyzed by increas-
ing the count by 1 and using a square root transformation. The results are presented
in Table 1.
Malathion baits provided significant mortality at 2.0 lbs AI/acre. The malathion
bait with poultry mash as a carrier gave significantly higher mortalities than the com-
mercially formulated baits. This agrees with the Florida Department of Agricul-
ture's (Fla. Dep. of Agric. 1938. Fla. Bull. of the Dep. of Agric. New Series No. 3)
finding that commercial egg mash is a better carrier in baits than bran. These results
indicate that 2% malathion baits formulated from a 5 lb/gal E.C. will perform as
well or better than the commercially formulated toxic baits which have been shown
to be effective (P. G. Koehler and D. E. Short. 1976. J. Econ. Ent. In press). The mash
formulation also provided significantly higher mortality than any spray formula-
tion included in the experiment.
Agricultural producers utilizing applicator-formulated malathion baits should
realize a major savings in the cost of mole cricket control on pastures. Extension
recommendations for applicator-formulated baits will be available pending label
approval.
We thank Dr. F. G. Martin for the statistical analysis of the mole cricket data.
P. G. Koehler, and D. E. Short. Department of Entomology and Nematology, Uni-
versity of Florida, Gainesville, Florida 32611.
TABLE 1. MEAN NUMBER OF MOLE CRICKETS KILLED IN EXPERIMENTAL PLOTS WITH
VARIOUS INSECTICIDES.
Mean number of
Insecticide* Formulation LB AI/acre dead or
moribund per plot**
Malathion (mash) 2%B 2.0 4.2 a
Malathion (cracked corn) 2%B 2.0 2.8 ab
Primifos-ethyl 2%B 2.0 2.0 b
Chlordane 8 lb/gal EC 5.0 1.8 b
Malathion 2%B 1.0 1.4 bc
0-[5-Chloro-l-(1-methyl- 2 lb/gal EC 2.0 1.4 bc
ethyl(-1H-1,2,4-triazol-
3-yl]o, o-diethyl phos-
phoro-thioate (Ent 29128)
Malathion (commercial) 2%B 2.0 0.6 cd
Diazinon 4 lb/gal EC 4.0 0.2 d
Check 0.2 d
Malathion 57% EC 1.5 0.0 d
Shell SD35651 20% WP 1.0 0.0 d
0-Ethyl 0-[4(methylthio) 6 lb/gal 2.0 0.0 d
phenyl]S-propyl phos-
phoro-dithioate (BayNTN
9306)
Shell WL43775 2.4 lb/gal EC 4.0 0.0 d
at 0.05 level as deter-
*Materials applied 15 September 1975.
**Means followed by the same letter are not significantly different
mined by the Duncans multiple range test.
The Florida Entomologist
NEW RECORDS OF CHIGGERS (ACARINA:
TROMBICULIDAE) FROM FLORIDA1,2
A. L. DOHANY3, AND H. L. CROMROY
Department of Entomology and Nematology,
University of Florida, Gainesville, Florida 32611
ABSTRACT
Chiggers (trombiculid larvae) were collected and identified from the
north, northcentral, and central parts of Florida. Six species are reported
as new to the State and 2 are reported from new hosts.
Chiggers or trombiculid larvae cause trombidiosis in humans in much
of the United States. The effects of chigger attack on humans is normally
characterized by itching and swelling at the site of chigger attachment.
Trombidiosis may also be accompanied by secondary infection and persist-
ent lesions. The primary hosts of chiggers are a variety of birds, mammals
and reptiles. Severe damage to turkeys may be caused by chigger attack
(Everett et al. 1973), and recent reports indicate that chiggers are the cause
of lesions on horses in the Pacific Northwest (Easton and Krantz 1973).
Only a few extensive, area-wide studies of chiggers have been conducted
in the United States (Loomis 1956, Gould 1956, Jenkins 1948, Crossley and
Proctor 1971). Surprisingly little work has been conducted on the chiggers
found in Florida. As a preliminary to systemic acaricide studies for the con-
trol of chiggers (Dohany 1974), a limited study of the distribution of the
chiggers within the general study area was conducted.
METHODS
Chiggers were collected primarily from litter samples and tree holes,
and were obtained by using the Tullgren modification of the Berlese fun-
nel. Some specimens were also collected from black plates, Samples were
taken from Tallahassee (Tall Timbers Research Station) in the northern
part of the state, from Gainesville in the northcentral part of the state,
and a small collection was made from Lakeland in the central part of
the state (Fig. 1).
Chiggers were collected in 80% ethyl alcohol and were mounted di-
rectly into Hoyer's media on microscope slides (Krantz 1970). Cover slips
were applied and the slides were heated over an alcohol burner until
bubbles began to form. After drying, the cover slips were ringed with Glyp-
tal and the cleared chiggers were identified to species. Measurements were
made with an ocular micrometer.
'Florida Agricultural Experiment Station Journal Series No. 6070.
2This study was supported by the Office of the Surgeon General; U. S. Army.
3Present Address: U. S. Army Medical Research Unit, Institute for Medical Research, Kuala
Lumpur, Malaysia.
Vol. 59, No. 2
The Florida Entomologist
0 LAKELAND
Fig. 1. Map of the state of Florida showing chigger collection sites.
RESULTS
Eutrombicula
The most common chiggers that attack humans in the United States are
in the genus Eutrombicula. The 2 most frequently occurring in the south-
eastern United States are E. splendens (Ewing) and E. alfreddugesi (Oude-
mans). Jenkins (1948a) presented the host and habitat information on these
2 species in detail. Numerous specimens of both species were collected from
all locations, primarily by the black plate collection technique and from
live-trapped rodents.
Jenkins (1948), Wharton and Fuller (1952), and Wolfenbarger (1952)
presented an abundant number of hosts from which each of these species
has been collected. The scutal measurements of E. alfreddugesi and E.
splendens are presented in Table 1.
Fonsecia
Two species of the genus Fonsecia were collected: one of the subgenus
Fonsecia and the other of the subgenus Parasecia. The genus Fonsecia is
characterized by having the PL's greater than the AM and the AM greater
than the AL's. The subgenus Parasecia differs from the subgenus Fonsecia
Vol. 59, No. 2
Dohany and Cromroy: New Chigger Records
TABLE 1. SCUTAL MEASUREMENTS OF Eutrombicula alfreddugesi (OUDE-
MANS) AND E. splendens (EWING), AND A COMPARISON OF MEA-
SUREMENTS MADE BY PREVIOUS CHIGGER TAXONOMISTS. ALL
MEASUREMENTS ARE IN MICRONS.
AW PW SB ASB PSB A-P AM AL PL S
Eutrombicula
alfreddugesi
Florida
A
B
C
D
E
F
G
Average
Jenkins (1949)
Wolfenbarger (1952)
Kansas
Eutrombicula
splendens
Florida
A
B
C
D
E
F
G
Average
Jenkins (1949)
Wolfenbarger (1952)
Missouri
Arkansas
Texas
87 40
90 42
94 45
93 45
92 47
93 46
94 44
92 44
90 43
27 31 29 42
25 37 32 47
25 30 36 49
27 36 33 46
26 27 30 44
26 30 31 47
27 40 32 49
26 33 32 46
27 33 33 47
77 88 43 23 26 27 28 29 40
92 43
95 43
95 45
95 47
92 43
100 49
91 44
94 45
91 44
83 94 43
92 104 47
100 112 50
25 --- 34 50
26 38 33 48
28 41 38 50
28 40 35 50
29 40 36 52
29 --- --- 48
27 37 23 48
27 39 35 49
27 37 34 48
27 33 34 47
30 35 33 47
29 36 33 47
by having normal AL scutal setae, while they are peg-like in the subgenus
Fonsecia. Radford (1942) originally proposed the raising of 2 Brazilian
species to the generic level and designating them Fonsecia as the "antero-
lateral setae were represented by tooth-like projections." Brennan and
Loomis (1959) in their review of the genus, placed 7 species in Fonsecia.
Loomis (1956) established the subgenus Parasecia with F. (Parasecia) gur-
neyi (Ewing) being the type species and referred 6 additional species to the
subgenus.
Brennan (1969), in adding 3 new species to the subgenus Parasecia, stated
that; "Parasecia is further distinguished from Fonsecia, whose four species
The Florida Entomologist
are also restricted to the New World, by a sinuous posterior scutal margin
(convex in Fonsecia) and a wide host range (Fonsecia species apparently
confined to reptiles)".
Fonsecia (Fonsecia) palmella Brennan and Loomis was collected from
Berlese samples of 5 treeholes and a sample of Spanish moss (Tillandisis
usneoides) from the Tallahassee area. This species had been previously col-
lected from the southeastern five-lined skink (Eumeces inexpectatus) and
from the five-lined skink (E. fasciatus) from Louisiana (Loomis 1956). This
is the only species of Fonsecia (Fonsecia) that has been reported from the
United States. The scutal measurements of specimens of F. (F.) palmella
are presented in Table 2.
TABLE 2. SCUTAL MEASUREMENTS OF Fonsecia (Fonsecia) palmella
BRENNAN AND JONES. ALL MEASUREMENTS ARE IN MICRONS.
Slide AW PW SB ASB PSB A-P AM AL PL S
A 56 70 26 25 18 15 43 12 37 69
B 55 69 26 27 17 13 41 12 38 64
C 54 70 26 27 16 13 46 12 40 63
D 56 69 26 25 17 12 41 11 35 61
E 55 71 27 28 17 12 42 11 39 62
Average 55 70 26 27 17 13 43 12 38 64
B. & L.* 57 71 27 30 17 14 42 12 37 63
*Brennan and Loomis (1959).
Fonsecia (Parasecia) gurneyi gurneyi (Ewing) was the most abundant
chigger collected from Berlese samples of treeholes. This species was col-
lected from all the collection sites in Gainesville, Tallahassee, and Lake-
land. This species has been reported throughout the southeastern United
States, as far north as Maryland and as far west as Texas. Loomis (1956)
listed a large number of reptiles and mammals that F. (P.) gurneyi gurneyi
was collected from and determined that its principal host in eastern Kansas
seemed to be the five-lined skink (E. fasciatus). The scutal measurements of
10 specimens of F. (P.) gurneyi gurneyi are presented in Table 3.
Euschongastia
Two species of Euschongastia were collected during this study. Farrell
(1956) conducted a detailed study of the Euschongastia of North America.
Euschongastia peromysci (Ewing) was taken from Peromyscus flori-
danus from the Gainesville area. This species has been previously reported
throughout the northeastern United States as far west as Oklahoma. Cross-
ley and Proctor (1971) reported this species from the eastern chipmunk
(Tamias striatus) and the pine vole (Microtus pinetorum) from Georgia.
This is the first record of this species from Florida.
Euschongastia peromysci can be easily distinguished by its unusual
galeal setae, having 1-3 stiff setules that arise near the base, and by its char-
acteristic scutal shape. The scutal measurements of 5 specimens are pre-
sented in Table 4.
Vol. 59, No. 2
Dohany and Cromroy: New Chigger Records
TABLE 3. SCUTAL MEASUREMENTS OF Fonsecia (Parasecia) gurneyi
gurneyi (EWING). ALL MEASUREMENTS ARE IN MICRONS.
Slide AW PW SB ASB PSB A-P AM AL PL S
A 65 76 29 25 15 17 31 26 45 61
B 64 75 32 24 16 18 30 24 42 ---
C 64 74 32 24 17 16 26 24 40 54
D 67 79 32 25 17 17 27 26 41 54
E 66 76 31 21 17 16 31 25 44 54
F 62 74 32 23 16 17 30 24 40 ---
G 64 76 30 31 18 17 31 24 40 55
H 65 78 29 22 17 18 31 24 41 56
I 66 76 31 24 16 18 32 25 43 53
J 65 76 31 24 16 18 30 27 45 59
Average 65 76 31 23 16 17 30 25 42 56
Loomis (55) 61 72 28 22 16 15 28 23 41 48
TABLE 4. SCUTAL MEASUREMENTS OF Euschongastia peromysci
(EWING). ALL MEASUREMENTS ARE IN MICRONS.
Slide AW PW SB ASB PSB A-P AM AL PL S
A 50 59 15 31 13 18 36 53 55 24 x 15
B 48 57 14 39 14 17 37 54 53 21 x 15
C 51 58 14 30 13 17 34 55 57 23 x 15
D 49 58 14 29 15 15 34 50 53 --- 16
E 50 57 --- 30 11 16 34 56 59 25 x 15
Average 50 58 14 30 14 17 35 54 55 23 x 15
Farrell* 50 57 22 27 8 13 27 35 46 27 x ---
*Farrell (1956). Average of 100 specimens.
A single specimen of Euschongastia setosa (Ewing) was collected from
a treehole in Gainesville in December 1971. Additional specimens were not
found, even though extensive samples were made from this same treehole
throughout the year. This collection in the winter corresponds with previ-
ous collections of this species. Farrell (1956) recorded this species occur-
ring from late fall through spring in North Carolina and Pennsylvania.
The specimen of E. setosa (a co-type) that was previously collected from
the closest locality to Gainesville, Okefenokee Swamp, Georgia, was also
collected during December (Ewing 1937).
Euschongastia setosa is characterized by its scutum being wider than the
PL and the sensillae heads being ovoid. The scutal measurements (in pm)
for the 1 specimen collected are:
AW PW SB ASB
PSB A-P AM AL
66 86 28 22 8 19 36 30 63 30 x 15
The Florida Entomologist
Microtrombicula
Microtrombicula crossleyi (Loomis) was collected only from a tree-
hole from the Tallahassee area. This species has previously been reported
from Kansas and Oklahoma (Loomis 1954).
Microtrombicula crossleyi is closely related taxonomically to M. tri-
setica (Loomis and Crossley), both having 3 pairs of sternal setae and mul-
tiple coxal setae. The chief characteristics separating the 2 species are the
reduced scutum of M. trisetica with 3 setae on coxa III while M. crossleyi
has 5 setae on coxa III. The scutal measurements of M. crossleyi are listed
in Table 5.
TABLE 5. SCUTAL MEASUREMENTS OF Microtrombicula crossleyi
(LOOMIS). MEASUREMENTS ARE IN MICRONS.
Slide AW PW SB ASB PSB A-P AM AL PL S
A 34 42 13 22 23 20 19 13 25 32
B 36 42 13 20 21 21 16 16 24 29
C 33 41 11 21 19 19 14 18 22 25
D 32 41 13 21 19 21 18 17 22 28
E 32 37 12 20 21 22 17 16 24 30
Average 33 41 12 21 21 21 17 17 23 29
V-G* 35 46 12 22 20 22 21 18 57 34
*Vercammen-Grandjean (1956).
Walchia
Walchia americana Ewing was collected from a treehole in Gaines-
ville. This species was originally described by Ewing (1942) from a "cotton
mouse" (Peromyscus gossypinus [?] from Tallahassee. It has been collected
from numerous mammal hosts. Loomis (1956) indicated that this species,
like all of the members of the subfamily Walchiinae, seems to occur only
upon mammals. In addition to the type location, this species has been col-
lected from Wisconsin (Farner 1946), Oklahoma, Nebraska, Iowa, Kansas,
Utah (Loomis 1956), California (Gould 1956) and Georgia (Crossley and
Proctor 1971).
The scutum of W. americana, as with the entire genus, has only 4 scutal
setae: 2 AL's, 2 PL's, but lacks an AM. The scutal measurements are pre-
sented in Table 6.
Although relatively few samples were taken during this study, 6 species
were found that were new to the State of Florida, and 2 are recorded from
their hosts for the first time. Thus, it is apparent that little is known of the
chigger distribution of the State of Florida. Florida, with its fairly diverse
habitats and with their relative ease of access would offer a unique oppor-
tunity for distribution and taxonomic studies.
ACKNOWLEDGEMENTS
The authors wish to thank Mr. W. A. Phillis, III for his assistance in the
collection of many of the specimens identified during this study.
Vol. 59, No. 2
188
Dohany and Cromroy: New Chigger Records
TABLE 6. SCUTAL MEASUREMENTS OF Walchia americana EWING. ALL
MEASUREMENTS ARE IN MICRONS.
Slide AW PW SB ASB PSB A-P AM AL PL S
A 47 70 41 20 48 29 --- 25 27 50 X 10
B 45 70 40 19 47 29 --- 28 28 50 X 10
Average 46 70 40 19 47 29 --- 26 27 50 x 10
Loomis* 45 68 38 20 44 30 --- 25 27 51 x ---
*Loomis (1956). Average of 7 specimens.
LITERATURE CITED
BRENNAN, J. M. 1971. Three new species of subgenus Parasecia Loomis
(genus Fonsecia) from northeastern Brazil and a key to the included
species (Acarina: Trombiculidae). J. Parasit. 55:662-6.
BRENNAN, J. M., AND R. B. LOOMIS. 1959. A review of the reptile chiggers,
genus Fonsecia (Acarina: Trombiculidae), with descriptions of two
new American species. J. Parasit. 45:53-64.
CROSSLEY, D. A., JR., AND C. W. PROCTOR, JR. 1971. New records of chigger
species (Acarina: Trombiculidae) in Georgia. J. Georgia Ent. Soc.
6:184-7.
DOHANY, A. L. 1974. Potential use of systemic acaricides in the control of
chigger (Acarina: Trombiculidae) populations on rodents. Disserta-
tion, University of Florida, Gainesville, Florida, 115 p.
EASTON, E. R., AND G. W. KRANTZ. 1973. A Euschoengastia species (Acari:
Trombiculidae) of possible medical and veterinary importance in
Oregon. J. Med. Ent. 10:225-6.
EVERETT, R. E., M. A. PRICE, AND S. E. KUNZ. 1973. Biology of the chigger
Neoschongastia americana (Acarina: Trombiculidae). Ann. Ent.
Soc. Am. 66:429-35.
EWING, H. E. 1937. New species of the subfamily Trombiculinae with a key
to the new world larvae of the akamushi group of the genus Trombic-
ula. Proc. Biol. Sco. Washington 50:167-73.
FARNER, D. S. 1946. A new host and locality record for the trombiculid
mite Walchia americana Ewing, with a note on its morphology.
Proc. Ent. Soc. Wash. 48:185-186.
FARRELL, C. E. 1956. Chiggers of the genus Euschongastia (Acarina: Trom-
biculidae) in North America. Proc. U. S. Nat. Mus. 106:85-235.
GOULD, D. J. 1956. The larval trombiculid mites of California (Acarina:
Trombiculidae). Univ. California Publ. in Ent. 11:1-116.
JENKINS, D. W. 1948. Trombiculid mites affecting man. I. Bionomics with
reference to epidemiology in the United States. Am. J. Hyg. 48:22-
35.
JENKINS, D. W. 1949. Trombiculid mites affecting man. III. Trombicula
(Eutrombicula) splendens Ewing in North America. J. Parasit. 35:
201-3.
KRANTZ, G. W. 1970. A manual of acarology. Oregon State University Book
Stores, Inc. Corvallis, Oregon. 335 p.
LOOMIS, R. B. 1954. A new subgenus and six new species of chigger mites
(Genus Trombicula) from the central United States. Univ. Kansas
Sci. Bull. 36:919-41.
The Florida Entomologist
LOOMIS, R. B. 1956. The chigger mites of Kansas (Acarina, Trombiculidae).
Univ. Kansas Sci. Bull. 37:1195-443.
RADFORD, C. D. 1942. The larval Trombiculinae (Acarina: Trombiculidae)
with descriptions of twelve new species. Parasit. 34:55-81.
VERCAMMEN-GRANDJEAN, P. H. 1956. Revision of the genera: Eltonella
Audy, 1956 and Microtrombicula Ewing, 1950 with descriptions of
fifty new species and transferral of subgenus Chiroptella to genus
Leptotrombidium (Acarina, Trombiculidae). Acarologia 7:34-257.
WHARTON, G. W., AND H. S. FULLER. 1952. A manual of the chiggers. The
biology, classification, distribution, and importance to man of the larvae
of the family Trombiculidae (Acarina). Memoirs Ent. Soc. Washington
4:185 p.
WOLFENBARGER, K. A. 1952. Systemic and biological studies on North
American chiggers of the genus Trombicula, subgenus Eutrombicula.
Ann. Ent. Soc. Am. 45:645-77.
INFESTATION OF NATIVE FLORIDA CATTLE WITH THE
NORTHERN CATTLE GRUB, HYPODERMA BOVIS (DIP-
TERA: OESTRIDAE)1-(Note). Larvae of the common cattle grub, Hy-
poderma lineatum (DeVillers), and the northern cattle grub, Hypoderma
bovis (Linn.), are obligatory parasites that develop in bovine hosts. H.
lineatum has a distribution throughout the United States, except Alaska,
and is a long established pest of cattle in Florida. H. bovis is abundant in
the United States north of Texas, but has not been able to survive in Texas
(Price et al. 1969. Texas A & M Univ. Agric. Ext. Serv. B-1080. p. 17), and is
only occasionally seen in cattle imported into Florida.
One larva of H. bovis was collected from a native cow in Orlando,
Orange Co., Florida on 28 May 1969. Since that time no further records of
the northern cattle grub on native Florida cattle had been made. On 14
November 1975, 1 third larval instar of H. bovis was taken from the back
of a native yearling steer from Cocoa in Brevard County. Third instar H.
bovis were again collected from native yearling steers in Cocoa on 20 and
23 December 1975 and 3 and 6 January 1976. Infestation rates at the times of
collection ranged from 2 to 14 larvae per steer. Most of the larvae col-
lected were mature and near pupation, and there was evidence that other
larvae had previously emerged from the hides of the steers.
These infestations constitute a new state record, and the possibility of
the establishment of this species in central Florida should be considered.
Jayson I. Glick, Department of Entomology, University of Florida,
Gainesville, 32611.
'Florida Agricultural Experiment Station Journal Series No. 9057.
Vol. 59, No. 2
The Florida Entomologist
INVASION OF FLORIDA BY THE "LOVEBUG" PLECIA
NEARCTICA (DIPTERA: BIBIONIDAE)'
LAWRENT L. BUSCHMAN
Department of Entomology and Nematology,
University of Florida, Gainesville, Florida 32611
ABSTRACT
Plecia nearctica Hardy has spread eastward from Mississippi and Louisi-
ana since 1940. Its range expanded approximately 32 km (20 miles) per year
reaching southern Florida and South Carolina by 1975. P. americana
Hardy, a native species, occurs in spring in woodland situations, whereas
P. nearctica occurs throughout the year but most frequently in May and
September and is most common in grassy habitats.
Large populations of the bibionid fly, Plecia nearctica Hardy, the "love-
bug", have attracted much attention in southeastern United States because
they are often encountered while driving on highways and become splat-
tered on windshields and radiators. Hetrick (1970) described the biology of
this insect and reported that similar populations of lovebugs were observed
in southern Louisiana in the mid 1930's. The lovebug did not gain public at-
tention in northcentral Florida until the mid 1960's although it was
thought to be native to the Southeast (Hetrick 1970). When P. americana
Hardy was collected near Gainesville, Florida, this study was initiated to
determine if P. americana was the native lovebug and P. nearctica had re-
cently arrived in Florida.
METHODS
I examined about 500 P. nearctica and 50 P. americana loaned from
insect collections at the following institutions: Philadelphia Academy
of Natural Science; Louisiana State University; University of Georgia;
Florida State Collection of Arthropods, Gainesville; Illinois State Nat-
ural History Survey, Urbana; Field Museum of Natural History, Chi-
cago; Museum of Comparative Zoology, Harvard University.
The curators of insect collections at the following institutions pro-
vided information concerning the Plecia in their collections or reported
that they did not have Plecia from southeastern United States collected
before 1960: Archbold Biological Station; Byrd K. Dozier Collection;
California Insect Survey, University of California; Cornell University;
Auburn University; University of Louisville; Mississippi State Univer-
sity; North Carolina State University; Clemson University; Michigan State
University; California Academy of Science; Indiana University; Penn-
sylvania State University; United State National Museum; American Mu-
'This research supported in part by the state funded research grant to the University of Flor-
ida to study the biology and control of the "lovebug", P. nearctica, L. C. Kuitert project di-
rector. Florida Agricultural Experiment Station Journal Series No. 8029.
Vol. 59, No. 2
The Florida Entomologist
seum of Natural History; Entomology Division, North Carolina Depart-
ment of Agriculture; Purdue University; Museum of Zoology, University
of Michigan.
Specimens were identified using descriptions by Hardy (1940, 1945).
Thompson (1975) has subsequently published additional descriptions. Data
on geographic distribution and biological notes were compiled from per-
sonal observations, data on borrowed specimens, data furnished by curators
when the determinations were by D. E. Hardy, and data published by Hardy
(1940, 1945), who examined about 30,000 bibionids (about 200 Plecia) ob-
tained from 37 insect collections of North and South America.
RESULTS
The geographic distribution of P. americana (Fig. la) extends along the
coastal plain of North America from North Carolina into Mexico. This
species has been collected periodically in Florida since 1888.
The geographic distribution of P. nearctica (Fig. Ib) in 1940 (Hardy
1940) extended along the Gulf coast from Mississippi to Costa Rica in Cen-
tral America. This species was first collected in western Florida, Escambia
Co., in 1949. In 1957 it occurred as far east as Leon Co., Florida, and in 1966
it was collected in Putnam Co., Fla. In 1972 it was collected in Highlands
and Sarasota counties, Florida. Currently, P. nearctica is found throughout
Florida except southern Dade and Monroe counties (Fig. Ib). To the north,
P. nearctica was collected in Colleton Co., South Carolina, in 1975.
Habitats recorded for P. americana were live oak hammock, wooded
ravine, and deciduous forest. P. americana has twice been reared from lar-
vae collected in rotten logs (Hardy 1940, and specimens by W. G. Brad-
ley in the Department of Entomology Collection, Louisiana State Univer-
sity).
The largest populations of larvae and adults of P. nearctica are found
in grassy habitats such as Bahia grass (Paspalum sp.) pastures and road-
sides. However, these insects fly extensively so adults can be encountered
in almost any habitat.
Throughout its extensive range, P. americana has been collected only
in April, May, and June; there is no evidence of a fall emergence. P. nearc-
tica has been collected most frequently in May and September, but it has
been collected in every month except November. Most of the collection
dates for P. americana in northcentral Florida are 2 or 3 weeks earlier than
similar dates for P. nearctica.
I observed P. americana flying in several compact swarms 1-2 m high
and 0.3 m wide. Each swarm maintained its position over a glass quart jar in-
verted over a 1 m high emergence trap located under the canopy of several
live oaks. When I kneeled next to one swarm, they began swarming over my
head and followed as I walked away. This behavior was so different from
that of P. nearctica that I was prompted to collect the specimens which
were later identified as P. americana.
DIscuSSIoN
The earlier conclusion that P. nearctica was native to Florida (Hetrick
1970) was probably based on misidentified P. americana specimens in the
Florida State Collection of Arthropods. Reexamination of available spec-
Vol. 59, No. 2
Buschman: Plecia nearctica Invasion
193
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The Florida Entomologist
imens indicates that P. americana is native to southeastern United States
but P. nearctica has recently expanded its range into Florida. P. nearctica
was in Louisiana as early as 1911 (Hardy 1940) but it did not reach Florida
until 1949. The range of P. nearctica has subsequently expanded across
Florida at a rate of about 32 km (20 miles) per year. Although P. nearctica
is capable of extended cross country flight (Hetrick 1970, Sharp et al, 1974),
its dispersal was undoubtedly aided by man, adults are easily transported
accidentally in or on automobiles and the larvae occur in sod which is
often shipped great distances.
The realization that P. nearctica has recently immigrated into Florida
has several consequences. First, immigrant species frequently undergo pop-
ulation explosions in their new habitat because they are no longer subjected
to the natural enemies with which they evolved (Sailer 1974). This suggests
that efforts to find natural enemies in older habitats such as Louisiana and
Central America could yield candidates for possible introduction into
Florida as biological control agents. Second, it would be useful to study
lovebug populations in older habitats to determine if populations remain
at high levels or if they subside in time. Third, P. nearctica is still spread-
ing and it is not clear how far south in Florida or north in the Carolinas it
may eventually extend its range.
The presence of 2 species of Plecia in Florida which have not always been
distinguished raises several points of interest. First, concerning recent pub-
lications on the lovebug, P. nearctica is probably the correct name since
P. americana appears to be a woodland insect that is usually collected in
small numbers. However, habitat records of P. nearctica larvae in hard-
wood leaf litter (Hetrick 1970) need to be reevaluated. Second, until we
can distinguish between larvae of the two species, larval determinations
will be in doubt unless representatives are reared for determination.
Third, a comparative study of the biology and behavior of the 2 species
would be in order.
LITERATURE CITED
HARDY, D. E. 1940. Studies in New World Plecia (Bibionidae-Diptera). J.
Kans. Ent. Soc. 13:15-27.
HARDY, D. E. 1945. Revision of Nearctic Bibionidae including Neotropical
Plecia and Penthetria (Diptera). Univ. Kan. Sci. Bull. 30:367-547.
HETRICK, L. A. 1970. Biology of the "Love-bug", Plecia nearctica (Dip-
tera: Bibionidae). Fla. Ent. 53:23-6.
SAILER, R. I. 1974. Foreign exploration and importation of exotic arthropod
parasites and predators. Pages 97-100 in F. G. Maxwell and F. A.
Harris, eds. Proc. summer inst. biological control of plant insects
and diseases. Univ. Miss. Press, Jackson, Miss.
SHARP, J. L., N. C. LEPPLA, D. R. BENNETT, W. K. TURNER, AND E. W.
HAMILTON. 1974. Flight ability of Plecia nearctica in the laboratory.
Ann. Ent. Soc. Am. 67:735-8.
THOMPSON, F. C. 1975. "Lovebugs", a review of the Nearctic species of Plecia
Wiedemann (Diptera: Bibionidae). Cooperative Econ. Insect Report
25:87-91.
Vol. 59, No. 2
The Florida Entomologist
HOST RANGE TESTING OF TELENOMUS REMUS
(HYMENOPTERA: SCELIONIDAE)1,2
B. WOJCIK, W. H. WHITCOMB, AND D. H. HABECK
Department of Entomology and Nematology,
University of Florida, Gainesville, 32611
ABSTRACT
Eggs of 39 species of Lepidoptera were tested as possible hosts of Tele-
nomus remus Nixon, a scelionid egg parasite. Spodoptera frugiperda (J. E.
Smith) and S. latifascia Walker eggs were used for rearing T. remus and as
checks in the host range testing. Of 7 Arctiidae, 1 Ctenuchidae, 5 Geometri-
dae, 1 Mimallonidae, 21 Noctuidae, 2 Notodontidae, and 2 Pyralidae spe-
cies tested, 11 species of Noctuidae and 1 Pyralidae were parasitized by T.
remus. Eggs of 5 Spodoptera spp., 2 Elaphria spp., Heliothis zea (Boddie),
and Feltia subterranea (F.) had high emergence rates of T. remus. Eggs of
Arzama densa Walker (Noctuidae), and Acigona influsella Hamps (Pyra-
lidae), potential biocontrol agents of aquatic weeds, were not parasitized
by T. remus.
Effective control of the Spodoptera pest complex would be an impor-
tant advancement in worldwide agriculture. Spodoptera spp., which attack
a large range of agricultural crops, cause extensive financial losses due to
crop damage and costs for chemical control. Telenomus remus Nixon3, a
scelionid egg parasite of Lepidoptera, indigenous to Sarawak and New
Guinea, was introduced into Israel for control of the armyworm Spodop-
tera littoralis Boisduval (Gerling 1972). The parasite has been successfully
established in Barbados and Montserrat on S. frugiperda (J. E. Smith), S.
eridania (Cramer), and S. sunia (Guenee); releases have also been made in
Trinidad for control of Spodoptera spp. and Heliothis zea (Boddie) (F. D.
Bennett, personal communication). In the United States, the fall army-
worm (Spodoptera frugiperda) overwinters in Florida and migrates north-
ward. Releases of T. remus in southern Florida would be a feasible initial
step towards the biological control of the Spodoptera pest complex in the
United States.
The biology of T. remus was described by Gerling (1972) and Schwartz
and Gerling (1974). Laboratory observations on host searching and selec-
tion by T. remus on S. littoralis eggs were described by Gerling and Schwartz
'Fla. Agr. Exp. Sta. J. Ser. No. 6049.
2This publication was supported in whole or in part by the National Science Foundation
and the Environmental Protection Agency, through a grant (NSFGB-34718 later known as
BMS 75-04223), to the University of California. The findings, opinions, and recommendations
expressed here are those of the authors and not necessarily those of the University of California,
the National Science Foundation, or the Environmental Protection Agency.
3The use of the name T. remus Nixon is problematical at present. Mr. C. F. W. Muesebeck
(U. S. National Museum) informed us, in litt., that when remus and several other species of
Telenomus associated with Spodoptera eggs are studied critically, they may all prove to be
minutissimus Ashmead, a species originally described from Trinidad, and reared in Puerto Rico
from S. frugiperda.
Vol. 59, No. 2
The Florida Entomologist
(1974). There is no host range information available in the literature. The
unpublished observations of both F. D. Bennett and ourselves have indi-
cated that the host range of T. remus appears to be restricted to Lepidoptera.
Our research was directed toward sufficiently determining the lepidopter-
ous host range of T. remus in order to obtain permission for a release of this
parasite in Florida.
MATERIALS AND METHODS
Cultures of the parasite were obtained from the West Indian Station of
the Commonwealth Institute of Biological Control, Trinidad. They were
processed at the Biological Control Laboratory, Division of Plant Indus-
try, Gainesville, Fla., where they were maintained under quarantine at ca.
250C and 55-70% relative humidity. They were reared in 8-dr, glass shell
vials with cotton-covered, foam plug stoppers which were moistened with
water and honey as needed.
Initially, Spodoptera frugiperda4, S. latifascia Walker4, S. exigua (Hub-
ner)4, and S. eridania5 eggs were tested for parasitism by T. remus. All were
acceptable hosts, with an 80 to 100% parasitism rate. Egg masses containing
ca. 50 eggs each and laid on paper toweling, were exposed in vials with ex-
cess populations of T. remus. Based on Schwartz and Gerling's report
(1974) that single females can successfully parasitize more than 60 eggs,
excess populations of parasites were maintained when equal numbers of
eggs and parasites were present in the test vials. The sex ratio of the para-
sites was ca. 1:1; females oviposited for 5 to 7 days. S. frugiperda and S. lati-
fascia eggs were used for maintaining the T. remus colony and were used as
standards for the host range testing. Lepidopterous eggs less than 2 days old
were obtained from laboratory reared4 or field collected adults and were
exposed for 2 to 3 days in 8-dr, glass shell vials. These vials had an excess
population of less than 6-day old T. remus. As a check, a Spodoptera sp.
egg mass was exposed in each of these vials with the test eggs. When pos-
sible, 3 or more replicates of host eggs obtained from more than 1 female
of each species were tested. Each replicate of test eggs contained ca. 20 eggs
with a minimum of 3 and a maximum of 68 eggs. After exposure, the test eggs
were carefully cleaned of any clinging parasites, removed to individual 1-
dr, glass shell vials, and examined after 10 days for parasite or host larvae
emergence. Exact host larval counts were not always obtained because of
cannibalism. When necessary, the test eggs were dissected and examined for
unemerged T. remus. Unreadable tests (owing to desiccation or molding)
were discarded. In cases where the Spodoptera check egg masses had less
than ca. 80% parasitism, the results of that test were eliminated.
RESULTS AND DISCUSSION
Thirty-nine species of Lepidoptera were tested for parasitism by T. remus
(Table 1). Eleven noctuid species and 1 pyralid species were hosts of T.
remus. Two Lepidoptera which feed on aquatic plants were not parasitized
'Obtained from laboratory colonies maintained at the USDA, ARS, Insect Attractant and
Basic Biology Laboratory, Gainesville, Fla.
'Field collected adults.
Vol. 59, No. 2
Wojcik et al.: Host Range of Telenomus remus
LEPIDOPTERA EGGS THAT WERE TESTED AS POSSIBLE HOSTS OF
Telenomus remus NIXON.* EACH TEST REPRESENTS A SEPA-
RATE EGG MASS AND EXPOSURE TO T. remus. THE TOTAL NUM-
BER OF EGGS PARASITIZED REPRESENTS THE COMBINATION OF
BOTH EMERGED AND UNEMERGED PARASITES WITH NO GREGARI-
OUS PARASITISM OF HOST EGGS. WHERE NO PARASITISM IS
RECORDED, SUCCESSFUL HOST LARVAL DEVELOPMENT OC-
CURRED.
Total No. of Eggs
Total No. of Parasitized/Egg
No. of Tests Eggs Tested Masses Parasitized**
Arctiidae
Apantesis phalerata (Harris) 2
Cisthene plumbea Stretch 2
Estigmene acrea (Drury) 11
Estigmene congrua (Walker) 9
Halisidota longa (Grote) 3
Hyphantria cunea (Drury) 3
Isia isabella (Abbot & Smith) 6
Ctenuchidae
Cisseps fulvicolia (Hubner) 1
Geometridae
Chlorochlamys chloroleucaria
(Guenee) 2
Eupithecia miserulata Grote 1
Metasiopsis sp. 3
Semiothisa sp. 2
Tornos sp. 2
Mimallonidae
Lacosoma chiridota Grote 3
Noctuidae
Amolita fessa Grote 4
Anicla infecta (Oschenheimer) 11
Anticarsia gemmatalis Hubner 3
Argyrogramma verruca (F.) 3
Arzama densa Walker 5
Caenurgia chloropha (Hubner) 6
Doryodes bistrialis (Geyer) 1
Elaphria chalcedonia
(Hubner) 3
Elaphria festivoides (Guenee) 1
Feltia subterranea (F.) 12
Heliothis zea (Boddie) 3
Hypsoropha monilis (F.) 3
Mocis marcida (Guenee) 5
Neoerastria apicosa
(Haworth) 3
Platysenta videns (Guenee) 4
0
14/3
4/2
0
0
0
0
15/2
2/1
93/8
11/2
0
0
14/2
4/2
TABLE 1.
Species
The Florida Entomologist
TABLE 1-Continued.
Total No. of Eggs
Total No. of Parasitized/Egg
Species No. of Tests Eggs Tested Masses Parasitized**
Pseudoplusia includes
(Walker) 7 69 0
Pseudaletia unipuncta
(Haworth) 5 71 2/2
Rachiplusia ou (Guenee) 3 40 0
Spodoptera dolichos (F.) 2 61 36/2
Tarachidia candefacta
(Hubner) 2 23 0
Trichoplusia ni (Hubner) 3 30 2/2
Notodontidae
Undetermined species 2 49 0
Litodonta hydromeli Harvey 3 47 0
Pyralidae
Acigona influsella Hamps 4 63 0
Nomophila noctuella (Denis &
Schiffermueller) 6 149 4/1
*Spodoptera frugiperda (J. E. Smith), S. latifascia Walker, S. exigua (Hubner), and S. eridania
(Cramer) eggs were checked as hosts for T. remus and had a 80-100% parasitism rate.
**The no. of emerged and unemerged parasites/the no. of egg masses that produced any para-
sites.
by T. remus: Arzama densa Walker, a native noctuid, attacking water hya-
cinth and pickerel weed, and Acigona influsella Hamps, an imported py-
ralid, being evaluated as a biocontrol agent for water hyacinth (N. Spen-
cer, personal communication). Based on high emergence rates, all species of
Spodoptera tested were more acceptable hosts than the other species tested.
Heliothis zea, Feltia subterranea (F.) and 2 Elaphria spp. also had appre-
ciable emergence rates of T. remus. Based on the results of these tests, T.
remus should have an adequate host range of economically important pests
in the field to insure its dispersal and survival.
ACKNOWLEDGMENTS
We would like to acknowledge the cooperation received from F. D. Ben-
nett in supplying the parasites and information necessary for initiating this
project. The review of the manuscript by him and D. Gerling is much ap-
preciated. We would like to thank E. Grissell of the Division of Plant In-
dustry, Florida Department of Agriculture and Consumer Services for
helping with the taxonomy of T. remus.
LITERATURE CITED
GERLING, D. 1972. The developmental biology of Telenomus remus Nixon
(Hym., Scelionidae). Bull. Ent. Res. 61:385-8.
GERLING, D., AND A. SCHWARTZ. 1974. Host selection by Telenomus remus,
a parasite of Spodoptera littoralis eggs. Ent. Exp. Appl. 17:391-6.
SCHWARTZ, A., AND D. GERLING. 1974. Adult biology of Telenomus remus
(Hymenoptera: Scelionidae) under laboratory conditions. Ento-
mophaga. 19:482-92.
Vol. 59, No. 2
The Florida Entomologist
INSECT-PLANT RELATIONSHIPS OF THE INSECTIVOROUS
PITCHER PLANT SARRACENIA MINOR'
DURLAND FISH
Dept. of Entomology and Nematology, University of Florida,
Gainesville, 32611
ABSTRACT
Eighty plants of Sarracenia minor Walt from Alachua County, Flor-
ida, were examined for insect prey and living insect associates. Leaf damage
by the noctuid moth Exyra semicrocea (Guen6e) had rendered two-thirds
of the leaves non-functional and young larvae were present in 22.5% of
the remaining functional leaves. Larvae of the sarcophagid Blaesoxipha
jonesi (Aldrich) were present in 64% of the functional leaves and were es-
timated to consume as much as 50% of the insect prey captured by the
leaves. Nests of the sphecid wasp Isodontia mexicana (Saussure) prevented
insect capture in an additional 2.5% of the functional leaves. Over 90% of
the insects captured by the leaves were ants, indicating that S. minor may be
specialized in securing its insect prey.
Pitcher plants seem to illustrate every imaginable relationship be-
tween plants and insects. They capture insects in leaves modified as pit-
fall traps and can absorb amino acids, peptides, and other nutrients from
insect prey (Plummer and Kethley 1964). Several species of aquatic Dip-
tera have adapted to survive in the digestive fluid of pitcher plant leaves
consuming the remains of captured insects and intercepting much of the nu-
trients earned by the plant (Jones 1935). Larvae of noctuid moths known
only from pitcher plants feed directly upon the leaf tissues often inflicting
considerable damage to local plant populations (Jones, 1920). Also,
empty leaves occasionally provide a convenient nesting site for certain
ants (Kannowski 1967) and wasps (Jones 1904).
This report concerns the insect relationships of the Hooded Pitcher
Plant, Sarracenia minor Walt. This species is locally abundant in poorly
drained pinewoods along the Atlantic Coastal Plain from central Florida
to the southern portion of North Carolina (McDaniel 1971). Tall pitchers
with translucent spots on a strongly recurved hood distinguish this pitcher
plant from all other species of Sarracenia (Fig. 1).
MATERIALS AND METHODS
On 18 August 1974, I collected one leaf from 80 plants in a moderately
dense stand of S. minor on the periphery of a small pinewoods pond near
Orange Heights, Alachua County, Florida. The noctuid moth Exyra semi-
crocea (Guen6e) had extensively damaged the plants leaving only one leaf
per plant, on the average, in a functional state. Each leaf was severed at
the base of the petiole, placed upright in a plastic bag, and stored in an in-
sulated box for transportation to the laboratory. Within 24 hours each leaf
'Florida Agricultural Experiment Station Journal Series No. 8019.
Vol. 59, No. 2
The Florida Entomologist
AftL
4
1
Fig. 1. A functional leaf of the Hooded Pitcher Plant, S. minor. (Approx.
1/3 actual size.) Fig. 2. Damage to S. minor leaf by the noctuid moth E.
semicrocea. Fig. 3. Co-occupation of a leaf chamber of S. minor by E. semi-
crocea (above) and B. jonesi (below). Fig. 4. Occupation of a S. minor leaf
chamber by a larva of the sphecid wasp I. mexicana (arrow).
was slit lengthwise with a surgical scalpel and inspected for insect prey
and living insect larvae. Samples of living larvae were reared to maturity
for positive identification.
RESULTS AND DISCUSSION
INSECT ASSOCIATES-The most obvious insect associate of S. minor was the
moth E. semicrocea. Plants were estimated to possess on the average, three
leaves, two of which had been at least partially consumed by E. semi-
crocea (Fig. 2). Leaves damaged by this moth were riddled with holes and
most had toppled over. Leaves in such condition are obviously not func-
tional as insect traps and usually contain no other insect associates. Of the
80 functional leaves 18 (22.5%) also contained moth larvae but did not
Vol. 59, No. 2
200
1
Fish: Insects of Sarracenia minor
show signs of damage. In most of these the larvae had just hatched from
small white eggs secured to the upper inside portion of the leaves.
Such extensive damage by E. semicrocea is surprising since S. minor pop-
ulations tend to be quite isolated and no other host species are known to
occur in Alachua County. Factors which prevent this pest species from
totally destroying its only host plant deserve more careful study.
The sarcophagid Blaesoxipha (Fletcherimyia) jonesi (Aldrich), which in-
habits the leafs digestive fluid, was rather abundant. Single larvae were in
51 (64%) of the leaves, and all 3 larval stages were represented among the
samples. B. jonesi is an obligate pitcher plant inhabitant as are all species
in the subgenus Fletcherimyia (Stone et al. 1965).
The young are deposited as first instar larvae into newly opened leaves
where they complete their larval development consuming freshly cap-
tured insects (Forsyth and Robertson 1975) (Fig. 3). Pupation occurs out-
side the leaf. These fleshy maggots were reported by Hepburn and Jones
(1919) to possess anti-enzymes which protect them from the digestive fluid.
Larvae of a third insect associate, the sphecid wasp Isodontia mexicana
(Saussure), were found in 2 of the samples (2.5%). The bottom and top ex-
tremes of an inhabited leaf were filled with dead plant material forming a
closed chamber. Inside the chamber was a single wasp larva with numer-
ous tree crickets and caterpillars obviously provided by the adult (Fig. 4).
This insect cannot be an obligate pitcher plant inhabitant since its range
exceeds that of the plants (Jones 1935; Bohart and Menke 1963).
All three of the insects inhabiting S. minor are of some disadvantage to
the plant. E. semicrocea obviously renders many of the leaves function-
less as insect traps, although the remaining leaf tissue stays green for some
time and may contribute to the plant by photosynthesis. B. jonesi does not
damage the leaves in any way but it does consume substantial amounts of
insect material captured by the plant. The largest maggots occupy nearly
as much space as the captured insects, which indicates that they may be in-
tercepting as much as 50% of the insect material from the plant.
Occupation of leaves by the wasp larvae also renders the leaves func-
tionless as insect traps since the tops of the pitchers are plugged with dead
plant material. The development time for this species is not known, but if
it is brief the leaves may become functional at a later time.
Considerable interaction occurs among the three insect associates of
S. minor. Leaves attached by E. semicrocea are not suitable as nesting sites
for I. mexicana. However, a leaf once occupied by the wasp may be safe from
moth damage since it seems to attack from the inside of the leaf.
The time of invasion by E. semicrocea is of importance to the survival
of the sarcophagid. B. jonesi can survive only if leaf damage is delayed
until enough insects are captured to support its larval development. Other-
wise, it will surely starve. Occupation of a leaf by B. jonesi does not pre-
vent the moth from attacking and their coexistence was noted in several of
the samples (Fig. 3).
The population of E. semicrocea may have a suppressive effect upon
B. jonesi but its effects on I. mexicana would probably be minimal since
the wasp presumably also nests elsewhere.
INSECT VICTIMS-The insects captured by the pit-fall-trap leaves of S.
minor were mainly ants, with lesser numbers of small Diptera, winged
Hymenoptera, and a few Blattidae. Over 90% of the leaves from which in-
The Florida Entomologist
sect remains were recognizable contained ants, sometimes 100 or more, in
various stages of decomposition. Several leaves contained many newly
captured live ants which boiled out when slit for inspection. The small
Diptera seemed to be associated with the few newly opened leaves as they
had not yet been decomposed and no other insect remains were present.
These observations indicate that S. minor may be more selective in cap-
turing insects than other pitcher plant species, and concur with similar ob-
servations made by Jones (1935). Judd (1959) and Swales (1969, 1972) re-
port 11 orders of insects recovered from the leaf fluid of S. purpurea and
Wray and Brimley (1943) report 12 orders from S. flava.
This apparent prey selectivity may be due to the physical nature of the
leaves as S. minor leaves are much narrower than those of S. purpurea and
S. flava. However, the possibility of a more order-specific chemical attract-
ant should not be overlooked.
CONCLUSIONS
Leaf samples from a north Florida collection site indicate that a pop-
ulation of S. minor can be profoundly affected by its insect associates. E.
semicrocea causes extensive damage to the leaf tissues, decreasing photo-
synthesis and disabling the insect traps. B. jonesi parasitizes the digestive
fluid consuming significant amounts of captured insects. I. mexicana plugs
the openings of leaves preventing the entrance of prospective insect prey.
All of these insect associates of S. minor are detrimental to the host plant.
Insects beneficial to the plant in the form of prey are much less diverse
than those of other pitcher plant species indicating that S. minor may spe-
cialize in capturing ants.
ACKNOWLEDGMENTS
I thank Dr. T. J. Walker for critically reviewing the manuscript and
Drs. E. E. Grissell and R. J. Gagn6 for insect identifications.
LITERATURE CITED
BOHART, R. M., AND A. S. MENKE. 1963. A reclassification of the Sphecinae.
Univ. Calif. Publ. Entomol. 30:91-182.
FORSYTH, A. B., AND R. J. ROBERTSON. 1975. K reproductive strategy and
larval behavior of the pitcher plant sarcophagid fly Blaesoxipha
fletcheri. Can. J. Zool. 53:174-9.
HEPBURN, J. S., AND F. M. JONES. 1919. Occurrence of anti-proteases in the
larvae of the Sarcophaga associates in Sarracenia flava. Contr. Bot.
Lab. Univ. Penn. 4:460-3.
JONES, F. M. 1904. Pitcher plant insects. Ent. News 15:14-7.
JONES, F. M. 1920. Pitcher plants and their moths. Nat. Hist. 21:297-316.
JONES, F. M. 1935. Pitcher plants and their insect associates. In: Walcott,
M. V., 1935. Illustrations of the North American pitcher plants.
Smithsonian Institution, Washington, D.C.
JUDD, W. W. 1959. Studies of the Byron Bog in southwestern Ontario. X.
Inquilines and victims of the pitcher plant Sarracenia purpurea L.
Can. Ent. 91:171-80.
Vol. 59, No. 2
Fish: Insects of Sarracenia minor
KANNOWSKI, P. B. 1967. Colony populations of two species of Dolichoderus
(Hymenoptera: Formicidae). Ann. Ent. Soc. Amer. 60:1246-52.
MCDANIEL, S. 1971. Genus Sarracenia (Sarraceniaceae). Bull. Tall Tim-
bers Res. Sta., No. 9, Tallahassee, Fla.
PLUMMER, G., AND J. B. KETHLEY. 1964. Foliar absorption of amino acids,
peptides, and other nutrients by the pitcher plant, Sarracenia flava.
Botan. Gaz. 125:247-60.
STONE, A., C. W. SABROSKEY, W. W. WIRTH, R. H. FOOTE AND J. R. COUL-
SON. 1965. A Catalog of the Diptera of America north of Mexico.
USDA-ARS Agriculture Handbook, No. 276. Washington, D.C.
SWALES, D. E. 1969. Sarracenia purpurea L. as host and carnivore at Lac
Carre, Terrebonne Co., Quebec. Can. Nat. 96:759-63.
SWALES, D. E. 1972. Sarracenia purpurea L. as host and carnivore at Lac
Carre, Terrebonne Co., Quebec. Part II. Can. Nat. 99:41-7.
WRAY, D. L., AND C. S. BRIMLEY. 1943. The insect inquilines and victims of
pitcher plants in North Carolina. Ann. Ent. Soc. Amer. 36:128-37.
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The Florida Entomologist
SPATIAL AND TEMPORAL DISTRIBUTION OF
OVIPOSITION BY PLUM CURCULIOS,
CONOTRACHELUS NENUPHAR1
C. 0. CALKINS, M. D. HUETTEL AND M. McKOY
Insect Attractants, Behavior, and Basic Biology Research Laboratory,
Agr. Res. Serv., USDA, Gainesville, Florida 32604
ABSTRACT
The spatial and temporal patterns of oviposition of the plum curculio,
Conotrachelus nenuphar (Herbst), were established near Gainesville, Flor-
ida, by placing suitable host fruit at 3 and 6 ft above the ground in wild
plum and oak thickets. Females consistently oviposited more in fruit at the
higher level, and they readily located fruit placed in non-host trees. Ovi-
position occurred from 11 March to 7 October and peaked in early June.
Although the plum curculio is considered a multivoltine species in the
southeastern U. S., the short period when wild host fruit is available and
the relatively long time it takes these insects to complete a generation
means that only one major generation is produced each year in north cen-
tral Florida.
Observations of oviposition by the plum curculio, Conotrachelus nenu-
phar (Herbst), in wild plum thickets suggested that fruit were attacked
more readily in the higher portions of thickets and that fruit at lower levels
often escaped damage. However, there were more fruit near the top of the
trees so it was difficult to determine the reason for the pattern. Also, be-
cause wild plums generally disappear by 1 June in northern and central
Florida, there was a question of whether oviposition would occur through-
out the growing season if host fruit were available or whether oviposition
was generally restricted to the spring months by some intrinsic factor. Ob-
servations in many thickets over a period of 2 years also revealed that most
isolated plum thickets bearing fruit harbored plum curculios even though
those thickets may not have produced fruit the previous year. Apparently
plum curculio females were efficient in locating fruit, especially in the
higher levels of thickets. We conducted this study near Gainesville, Flor-
ida, during 1974, to test the validity of these observations.
MATERIALS AND METHODS
Wild plum thickets are useful for experimenting with the behavior and
population dynamics of the plum curculio because this type of community
is stable and functions as an ecological island. In such habitats, popula-
tion fluctuations are normally not influenced by insecticide applications
or other agricultural practices.
For this study, we selected an isolated plum thicket (Prunus angusti-
folia Marsh.) with a moderately high rate of oviposition by plum curculios.
We also selected a young oak thicket (Quercus sp.) of similar configuration.
The thickets were located ca. 100 m apart and separated by thick woods.
LColeoptera: Curculionidae.
Vol. 59, No. 2
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