March 1970] KOGAN AND GOEDEN : STATUS OF Lcma trilineata daturaphila 537

Maheux, G. 1924. Insects of the season in Quebec in Say, T. 1824. Descriptions of coleopterous insects col-
1923. Entomol. Soc. Ontario, 54th Ann. Rep. lected in the late expedition to the Rocky Mountains,
p. 71-73. etc. J. Acad. Natur. Sci. Philadelphia 3(2) : 403-62.
Olivier, A. G. 1808. Entomologie, ou Historie naturelle Schaeffer, C. 1933. Short studies in the Chrysomelidae.
des insectes. Vol. 7. Paris. J. N. Y. Entomol. Soc. 41: 297-325.
Omer-Cooper, G., and P. Miles. 1951. On Lcma trili- Swezey, O. H. 1943. Notes: Lcma trilineata californica
neata—a beetle closely resembling the tobacco slug Schaeffer. Proc. Hawaiian Entomol. Soc. 11(3) :
attacking the Cape gooseberry. S. Afr. T. Sci. 269.
47(12): 330-3.
Symons, T. B. 1905. The common injurious and bene-
Peterson, A. 1951. Larvae of Insects. Columbus, Ohio. ficial insects of Maryland. Md. Agr. Exp. Sta. Bull.
Pt. 2, 416 p. 101: 125-204.
Riley, C. V. 1869. First annual report on the noxious, Tuthill, L. D. 1949. Notes: Occhalia pacifica (Stal).
beneficial and other insects of the State of Missouri. Ibid. 13(3) : 330.
4th Annu. Rep. State Board Agr. 181 -f 7 p.
Van Zwaluwenburg, R. H. 1938. Notes: Lcma nigro-
Sanderson, E. D. 1913. Insect Pests of Farm, Garden vittata Guer. Ibid. 10(1) : 9.
and Orchard. J. Wiley, N. Y. 684 p. 1947. Notes: Lcma enemies in California. Ibid. 13(1):
Sanderson, E. D., and L. M. Peairs. 1931. Insect Pests 23.
of Farm, Garden and Orchard. J. Wiley, N. Y. Walker, C. M. 1899. The sound-producing organs of
568 p. Lcma trilineata. Entomol. News 9(1) : 58-59.

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The Biology of Lema trilineata daturaphila, (Coleoptera: Chrysomelidae) with
Notes on Efficiency of Food Utilization by Larvae 1
MARCOS KOGAN 2 AND RICHARD D. GOEDEN 3
Division of Biological Control, University of California Citrus Research Center and
Agricultural Experiment Station, Riverside 92502

ABSTRACT
The biology of Lcma trilineata daturaphila Kogan & Larvae increased an average of 184 times in weight,
Goeden was studied under controlled laboratory condi- or a total of 60 mg, and consumed an average total of
tions. Adult females oviposited during 3-4.5 months and 2i7.3 mg of food during their development, which yielded
laid a maximum total of 2700 eggs. The mean duration a gross food conversion ratio of 0.276. Slow- and fast-
of the developmental stages (in days) was: egg 4-5, developing larvae were observed to deviate from the mean
larval 8, prcpupal 4-5, pupal 6-7. The insect in all stages in both overall weight gain and gross conversion ratio.
is described and figured, as is the distribution of larval
chemorcceptors.

The systematic status of Lcma trilineata datura- technique for rearing larvae and adults used in our
phila Kogan & Goeden, naturally associated with studies on the host selection behavior of this sub-
plants of the genus Datura, especially D. mctcloidcs species (Kogan 1969/ Kogan and Goeden 1969).
de Candolle, in southern California, has been dis-
cussed (Kogan and Goeden 1970). The biology of MATERIALS AND METHODS
the three-lined potato beetle, L. trilineata (Olivier) Rearing Technique.—Bouquets of freshly cut field-
sens, hit., was first described by Harris (1841) and collected or greenhouse-grown D. mctcloides served
Riley (1869). Subsequent authors (Blaisdell 1893; as food for larvae and adults. Field-collected and lab-
Sanderson 1913; Sanderson and Peairs 1931; Britton oratory-reared adults were placed in a 75x45x45-cm,
1932, 1933; Blackaller 1945; Essig 1958) added little glass-topped sleeve cage containing 2 large bouquets
original information to these accounts, with the ex- which served both as food and sites for oviposition.
ception of Omer-Cooper and Miles (1951), who The bouquets were held in water-filled 250-ml poly-
worked with adventive South African populations. ethylene jars and were renewed every 5- to 6-day
Force (1966) provided data on oviposition and adult period. During this process, the used bouquets were
longevity of the California subspecies. placed in the dark compartment of a rectangular,
This study of the biology of L. t. daturaphila was translucent-plastic refrigerator crisper, % of which
undertaken to gain information neded to develop a was blackened with opaque paint. Beetles attracted
from the foliage of the used bouquets to the illumi-
1
These studies were sunported in part by the John Simon nated part of the crisper were collected with an
Guggenheim Memorial Foundation, by Dry-Lands Research Insti- aspirator and returned to the oviposition cage.
tute Grant NM-6E, and by the Brazilian National Research Coun-
cil. aReceived for publication June 4, 1969.
Presently, Associate Entomologist, Illinois Natural History 4
Survey,
3
Section of Economic Entomology, Urbana 61801. M. Kogan. 1969. Host selection studies with Lcma trilineata
Assistant Professor of Biological Control and Assistant daturaphila n. n. (Coleoptera: Chrysomelidae). Ph.D. thesis, Uni-
Entomologist. versity of California, Riverside. 273 p.
538 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 63, no. 2

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FIG. 1.—A, Larval rearing cages (bottom shelf) and 3 pupation cages (top shelf) ; B, constant-level reservoir
for the series of bouquet-filled jars in the larval rearing cage.

Egg clusters were collected from the used bouquets.

Pieces of leaves with the egg clusters attached were
transferred to a 100x20-mm petri dish containing a
moist filter paper. The eggs were incubated in a tem-
perature cabinet at 27±1°C for 3 days. The egg
clusters were transferred to the larval rearing cage
when the heads of the larvae could be distinguished
through the chorions.
The larvae were reared in a 122x45 x60-cm glass-
topped sleeve cage (Fig. 1) containing 3 bouquets in
water-filled polyethylene bottles. These bottles were
interconnected basally by 7 mm-iD neoprene tubing.
The 1st bottle of the series was connected to a con-
stant-level water reservoir (Fig. 1 B) filled to a
height which prevented the water in the bottles from
overflowing but kept the stems of the bouquets con-
stantly submerged. With this system, handling time
was held to a minimum and the bouquets remained
in good condition for ca. 6 days. The egg clusters
were placed on the leaves of the caged bouquets after
incubation and the newly emerged larvae were allowed
to freely disperse. Larvae completed development on
these bouquets in 7-8 days.
The fully grown larvae were transferred from the
larval rearing cages to fresh bouquets in 34x32x35-
cm, glass-topped sleeve cages (Fig. 1 A (top)) fur-
nished with a shallow tray covered with a thin layer
of medium grade vermiculite. Larvae descended from
the bouquets to pupate in the vermiculite. Pupae re-
mained in these cages until adult eclosion. Adults
were held in cages by age groups to await use in ex-
periments.
Life History Studies.—During developmental and
feeding studies, newly hatched larvae were individu-
ally isolated in 100x20-mm petri dishes, the bottoms FIG. 2.—Cage used for isolation of individual females.
of which were covered with a 8-mm-thick layer of (A, 250-ml polyethylene bottles; B, water-refill tube; C,
plaster of paris, which was kept continuously moist. 850-ml plastic container; D, lid; E, wire screen; F,
perforated lid of bottle.)
A fresh disc punched from a D. meteloides leaf with
March 1970] KOC'.AN AND GOEDEN : BIOLOC.Y OF Lcnia trilinccitci daturaphila 539

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FIG. 3.—L. t. daturaphila in its developmental stages on leaves of I), mctcloides. A, Egg cluster; B. 2nd-stadial
larvae; C, early 3rd-stadial larvae feeding gregariously; D, late 3rd-stadial larvae dispersing.

a no. 14 cork borer was offered to the larvae each day. tracts along the longitudinal axis and slightly with-
These cultures were held at 27±1°C, a relative draws from the poles during incubation. The larval
humidity close to saturation, and a 12/12-hr photo- head capsule can be seen through the translucent
period. chorion toward completion of embryonic development;
The egg production of 10 9 was individually re- at eclosion, it is already sclerotized and pigmented.
corded. Kach of newly emerged females was isolated The eggs are usually laid on the interior surface of a
with 1 or 2 $ in individual cages (Fig. 2) fashioned leaf blade. Their length and diameter at the equa-
from 850-ml plastic food containers (12 cm high, 12 torial line measured 1.22±0.03 and 0.54±0.02 mm,
cm mouth diam, 9.5 cm bottom diam). The neck of a respectively (X±SD of 20 measurements).
250-nil polyethylene bottle projected through a hole B. Larva (Fig. 3-6).—Four larval instars were ob-
in the bottom of each container. The perforated lid served. The morphological features of all instars
of the bottle, once screwed into place, served to fasten were similar. The following description based on the
the container to the bottle. A 5-mm-iD neoprene 4th stadial larva, unless otherwise stated, also de-
tube inserted laterally through a tight-fitting hole in scribes the first 3 instars.
the bottle permitted refilling without opening the cage. The general body coloration is yellowish-white;
The lid of the container was fitted with a fine-mesh the head capsule, pronotal plates, and leg segments,
screen. The stalk of a small shoot of fresh D. mctc- dark brown. The integument of the intersegmental
loides was thus immersed in water, while its foliage folds is smooth; the segmental surfaces are invested
was exposed to the beetles in the upper chamber. with microspines.
The head capsule (Fig. 5 A-C) is round, smooth,
RESULTS AND DISCUSSION and covered with scattered, symmetrically distributed
Morphology of Developmental Stages.—A. Egg hairs. The epicranial suture is deep, its lateral arms
(Fig. 3 A).—The eggs are oblong, glossy yellow, and reaching the base of the antennae. Nine stemmata
are covered with a mucilaginous secretion which are present on each side: 2 close to the hypostomal
serves to attach them to the leaf surface and one ridge, 1 near the antennal socket, and 6 along the
another in clusters of irregular size and shape. The lateral arm of the epicranial suture. The labrum is
mucilage at the cephalic pole is pale brown and dark- triangular and deeply recessed at the middle of the
ens with age. The vitellinic substance gradually con- anterior margin. Each mandible (Fig. 5 E) bears 6
540 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 63, no. 2
ing to surfaces and are used in locomotion. One
thoracic and 8 abdominal spiracles are present. The
pleural areas of the thorax and abdomen bear clusters
of 3 or 4 peglike sensilla atop convex areas situated
immediately below and behind the spiracles. The
abdominal segments each bear rows of sparse, evenly
distributed hairs. The anus is situated dorsally and
is directed cephalad; the excreta thus accumulate
atop the body of the larva.
C. Prepupa and Pupa.—The 4th stadial larva trans-
forms into a prepupa without molting. It rids itself
of the load of excreta on its back (Fig. 6) and be-
comes more rigid and globular in shape. All morpho-
logical features of the larva are maintained in the
prepupa. The prepupal stage begins immediately after
the formation of the pupal case.
The pupa initially is bright yellow but darkens with
age. Its principal proportions are: length 6.2 mm;
width at the base of the pronotum 1.6 mm; width at
the humeri 1.9 mm. In dorsal view, only the protrud-

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ing eyes are visible, as the head is mostly concealed
D by the pronotum. The pronotum is produced forward
at 2 points; its posterior margin is bisinuous, with a
shallow, longitudinal groove at its base. The wing
thecae cover most of the 3rd pair of legs. The meta-
notum is produced posteriorly into a point and pre-
sents a keel-shaped, longitudinal, median groove. The
abdomen has 7 apparent tergites and 5 spiracles vis-
ible near their anterolateral angles. The tip of the
FIG. 4.—L. t. daturaphila larval instars. A, 1st; B, abdomen bears 2 inwardly hooked processes.
2nd; C, 3rd; D, 4th. Developmental Life History.—Table 1 summarizes
the duration of the life cycle of L. trilineata datura
teeth, the outermost two show marginal crenulations. phila reared under laboratory conditions on D. mctt
The maxillae bear sensorial hairs on the lacinia (Fig. loides.
5 D). The maxillary palpi are 3-segmented; each seg- Larval Development.—A. Larval Eclosion.—After
ment has 1 large, circular, sensillum placodeum on 4 or 5 days of embryonic development, the larvae
its ventral side and the 3rd segment (Fig. 5 F) is ruptured the chorion with their mandibles and
invested with 11-12 peglike sensilla basiconica at its emerged headfirst through a longitudinal slit along
tip. The prementum bears 2 long sensilla trichodea; the midventral region of the egg. The prothorax and
the postmental plate, 4 sensilla trichodea. The labial legs were freed less than a minute after eclosion be-
palpi are 2-segmented; the 2nd segment (Fig. 5 G) gan. Then, with their abdomens still contained within
bears 1 minute hair arising from a pouchlike socket the egg membranes, the larvae swung the anterior
and 7 peglike sensilla at its tip—3 large and 3 small part of their bodies back and forth until their tarsal
sensilla around a central, larger, slightly acuminate claws grasped the surface of the leaf. Having gained
sensillum. The antennae (Fig. 5 H) are 3-segmented this leverage, the larvae freed the rest of their bodies,
and telescope within a circular socket near the base of aided by continuous waves of peristalsis directed
each mandible. The basal antennal segment bears 3 caudad from the head. The entire act of eclosion
circular sensilla placodea; the 2nd segment, 1 sen- lasted 2-4 min. Eggs in the same cluster usually
sillum placodeum; the membranous apical segment hatched concurrently.
supports at least 3 different kinds of sensorial ele- B. Larval Feeding (Fig. 3 B, C).—Newly hatched
ments : 1 large and thick apically situated sensillum larvae immediately started to feed on the epidermal
basiconicum; 2 smaller sensilla basiconica situated at hairs of the leaves. Larvae hatched from eggs of the
diametrically opposed points near the base of the seg- same cluster fed side by side in close contact, usually
ment ; 1 long sensillum trichodeum arising from a in a circular front (Fig. 3). The larvae initially did
globular protuberance, and 2 sensilla basiconica ex- not perforate the leaves, but rather bored shallow
ternally situated on this same protuberance. cavities in the epidermis and lower parenchyma.
The pronotum bears a dorsal plate divided mid- Symptoms of this early feeding activity were chlorotic
longitudinally, which is reduced in the first 2 instars, spots in the upper epidermis above the areas attacked
but covers almost the entire pronotum of the 3rd and by the larvae. Second-stadial larvae also remained in
4th instars. The legs have complex sclerotized coxal close contact but chewed through the mesophyll and
structures and single-clawed tarsi. The abdomen opposite epidermis and consumed all but the thicker
bears 8 fleshy prolegs, which aid the larva in adher- veins of the leaf blades (Fig. 3 C). Early 4th-, and
March 1970] KOGAN AND GOEDEN : BIOLOGY OF Lcma trilineata daturaphila 541

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FIG. 5.—/-. t. daturaphila larval head and appendages. A, Anterior view; B, lateral view; C, posterior view;
D, maxillae and labium, ventral view; E, mandible, interior lateral view; F, tip of maxillary palpus; G, tip of labial
palpus; H, antenna.

sometimes late 3rd-stadial larvae began to disperse on their backs were carefully removed with a brush
randomly to new leaves when the initial food supply before weighing. Leaf discs were weighed before and
was exhausted (Fig. 3 D ) . Older larvae fed on stems after each 24-hr period of larval feeding. The weight
and flowers in the absence of leaves. The excreta of the excreta was not determined; therefore, no data
that accumulate on the dorsal surfaces of all instars on digestibility are presented. Larval weight in-
apparently is characteristic of larvae belonging to this creased ca. 184 times from the 1st through the 6th
subfamily (Fig. 6). or 7th day, then dropped 30-50% as the prepupal
C. Weight Gain.—Larvae were isolated immedi- stage was reached. This drop in weight was mainly
ately after hatching and were fed discs cut from due to elimination of the exudate used in the con-
freshly excised leaves of D. mctcloidcs. The larvae struction of the pupal case.
were weighed every 24 hr. The accumulated excreta D. Efficiency of Food Utilization.—Efficiency of
542 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 63. no. 2
development. Therefore, the amount of leaf con-
sumed during the 1st days could be only approximated.
These approximations and the determination of the
amount of food consumed during subsequent days em-
ployed a correction factor adapted from Waldbauer
(1962). In calculating the correction factor, 10 leaf
discs of the size and nature used for feeding the lar-
vae were held for 24 hr on a moist filter paper in a
petri dish. The weight of the discs was determined
before and after the 24-hr exposure. The mean
weight gain of 10 leaf discs was then used in the
following calculations:
Fc = [1 - c/2] [I - (L + c L ) ] ,
where: c = correction factor = ^24!/!-
FIG. 6.—L. t. daturaphila 4th-stadial larva in lateral A24I = weight gained by a leaf disc after 24
view showing accumulation of excreta on dorsum. hr exposure in the moist chamber.
I = initial weight of 1 intact disc.
Fc = corrected weight of food consumed.
food conversion has been studied in a few species of L = weight of uneaten food.

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phytophagous insects, and the results obtained were
summarized by Carne (1966) and Waldbauer (1968). The efficiency of food conversion varied in time as
Some investigations used the fresh weights of the observed by Carne (1966). A larval weight gain
larvae and the leaf material consumed to calculate the plotted against time approximated an S-shaped curve
conversion ratio. Waldbauer (1964) commented on (Fig. 8 ) . Growth approximated a geometric progres-
the advantages of using dry weight instead of fresh sion from eclosion to the 8th day. However, the
weight in calculating food conversion ratios, but such weight of food consumed did not follow such a pro-
a technique has the disadvantage of precluding deter- gression. Recorded amounts of food consumed showed
minations based on the same individuals during the at least 2 phases of slower increase corresponding to
course of their development (Carne 1966), or per- periods following the 2nd and 3rd molts (days 5 and
mitting only an indirect estimate of their dry weight 7, Fig. 8) ; however, this reduction in food intake was
by use of aliquots (Waldbauer 1968). largely compensated by a concomitant increase in
The conversion ratio was calculated by dividing efficiency of conversion.
the body weight gain of a larva by the fresh weight Two tendencies were noted in the growth rates of
of the food it consumed during the same period. The the individuals observed. Data on a typical fast-
leaf discs used in the life-cycle studies were held in growing and a typical slow-growing larva are shown
a petri dish containing moist filter paper. The leaf in Fig. 8. Table 2 shows calculations of the overall
discs gained weight after 24 hr by water absorption, weight gain, total food consumed, and gross conver-
and this weight gain was greater than the amount sion ratio.
consumed by larvae during the first 2-3 days of their Table 2 shows that the slow-growing larva actually

FIG. 7.—L. t. daturaphila pupa. A, Opened pupal case with pupa; B, pupa, dorsal view; C, same, ventral view.
March 1970] KOGAN AND GOEDEN : BIOLOGY OF Lema trilincata daturaphila 543
Table 1.—Summary of the life cycle of L. trilincata consumed more food than the fast-growing larva.
daturaphila reared in the laboratory on D. mctcloidcs.
However, its gross conversion ratio was 0.246, and
Developmental stage Duration in days that of the fast-growing individual was 0.316. Fig. 8
shows that the conversion ratio rose from a low of
gg 4-5 0.037 on the 2nd day to a high of 0.459 on the 7th day
Larval I 1.6 of development and increased nearly constantly day
II 2.1 by day without showing the fluctuations observed by
III 2.1
IV 2.2 Carne (1966). Exception to this steady increase oc-
Prepupal 4-5 curred at the end of the 3rd stadium. This efficiency
Pupal 6-7 in food utilization indicates an excellent adaptation
Preovipositional 3-8 of the beetle to its preferred food plant and meets the
Total from egg to egg 25-33
needs of its accelerated larval development.

80 —
Weight of larva e •
Weight of food consumed

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Slow and f a s t g r o w i n g l a r v a e —

— 60 —

5Of—

3 0 -

20 —

< 1st. instar- -2nd. instar—X—3rd instar- -4th. instor- -••4- -Prepupo

•530 —

•030
10
TIME IN DAYS
FIG. 8.—A, Mean weight increase and weight of food consumed by 10 larvae, and weight increase of 1 SIQW-
and 1 fast-growing larva of L. t. daturaphila; B, mean conversion ratio of 10 larva and conversion ratio of 1 slow
and 1 fast-growing larvae.
544 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 63, no. 2
Table 2.—Summary of calculations of the efficiency of
L. trilineata daturaphila in utilizing D. meteloides leaves
as food during larval development.
Mean of Fast- Slow-
10 developing developing
larvae larva larva
Overall weight
gain (nig) 60.0 81.3 64.9
Total food
consumed (mg) 217.3 257.6 263.3
Gross conversion
ratio* 0.276 0.316 0.246
Mean conversion
ratio±CL 0.187±0.156 0.206+0.178 0.207±0.189
• Gross conversion ratio = overall weight gain divided by total
food consumed.

E. Duration of the Larval Stadia.—Larval develop-

ment usually lasted 8 days, but the duration of each
stadium varied to some degree. Table 1 shows the

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results obtained with 10 larvae reared under con-
trolled conditions.
F. Growth Rate.—The body length and the width
of the head capsules of 10 larvae were measured
daily. Fig. 9 expresses the results graphically. The
average rate of increase of the width of the head
caspule was 1.48.
Post-Larval Development.—A. Prepupal and Pupal
Development.—Certain events marked the onset of FIG. 9.—Growth rate during larval development (mean
the prepupal stage. Fourth stadial larvae ceased feed- of 10 individuals) based on total body length (inter-
rupted line) and width of head capsule (histogram).
ing toward the 8th or 9th day of development and (Vertical lines indicate confidence limits (5% level).)
subsequently lost up to half their maximum weight
before pupating. These larvae also became strongly
positively geotactic and descended to the ground, had a foamy consistency and hardened on contact
whereupon under field conditions they tunneled 2-4 with air. Particles of soil trapped within this sub-
cm into the soil and constructed their pupal cases. stance lent additional rigidity to the cell when it
The prepupal stage ended with pupation after 4-5 hardened. The pupal case was rough externally and
days inside the pupal case. The pupal stage lasted 6-7 generally smooth inside except where a few intrud-
days. ing soil particles caused occasional irregularities. The
B. Formation of the Pupal Case (Fig. 7 C).— cell was oval, 8.5 mm long and 5.3 mm wide exter-
Having reached the proper depth, the larvae pushed nally. In the laboratory, L. trilineata daturaphila was
and compacted the surrounding soil until an oval able to pupate without forming a pupal case. How-
chamber was formed. A whitish liquid expelled ever, the prepupa still produced normal complements
through the mouth was then applied to the walls of of exudate before pupating. The secretion in this case
the chamber by aid of the mouthparts. This exudate was irregularly regurgitated near the pupation site.

FIG. 10.—L. t. daturaphila adults. A, Dorsal view of female (left) and male (right) ; B, copulation.
March 1970] KOGAN AND GOEDEN : BIOLOGY OF Lema trilincata daturaphila 545
Table 3.—Daily, weekly, and total egg production, productive life, and mature egg complement in ovaries after
death of 10 L. t. daturaphila females reared in individual cages on D. mctaloidcs.
Female no.
I II III IV V VI VII VIII IX X x±CL Cumulative
Daily mean 11.5i 13.3 21.5 13.9 26.7 10.3 23..7 15.6 17.7 17.8
Productive life (weeks) 15 15 18 13 6 13 13 18 14 18
Weekly production
Week 1 180 175 226 147 267 14 208 8 194 147 156.6±60.6 156.6
2 109 67 242 68 232 130 275 30 110 205 146.8±61.0 303.4
3 90 0 65 150 248 117 254 73 109 249 135.5±63.1 438.9
4 112 124 177 195 271 140 21 256 248 198 174.2±55.1 613.1
5 130 181 270 174 227 112 178 168 107 111 165.8±38.2 778.9
6 117 144 294 147 77 0 203 41 85 222 133.0±63.5 911.9
7 62 99 207 103 0 313 165 126 189 140.4+65.1 1051.9
8 71 92 140 88 151 171 230 151 139 137.0±34.8 1188.9
9 103 119 196 132 138 211 101 173 187 148.9±28.5 1337.8
10 99 68 167 62 18 309 214 218 150 145.0±66.2 1482.8
11 19 49 15 67 39 119 167 166 96 81.8±41.9 1564.6
12 38 32 143 15 73 143 190 96 156 98.4±44.8 1663.0
13 71 36 156 0 0 40 114 42 103 62.4±38.0 1725.4
14 2 105 114 79 3 73 62.7±35.1 1788.1
15 0 8 118 62 47 47.0 1835.1

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16 61 94 34
17 112 67 34
18 12 32 50
Total 1203 1299 2715 1172 1122 932 2445 2091 1828 2370
Eggs in ovaries 0 0 0 0 5 0 1 0 2 0

Adult Stage.—A. Emergence.—The mandibles of and usually were destroyed in the process. The beetle
the imago were discernible through the transparent bored its way out of the pupal case through an irregu-
integument of the pupa by the end of the pupal period, lar hole at 1 pole. The cuticle hardened in the field by
the pupal exuviae were shed inside the pupal case the time the beetle reached the surface of the ground;
in the laboratory, the adults remained quiescent after
shedding the pupal exuviae until the hardening proc-
ess was completed.
B. Copulation (Fig. 10 B).—The 1st mating oc-
curred 1-2 days after emergence. Beetles remained
in copula for long periods. A large proportion of this
activity seemed to occur between 10 :00 and 14 :00 hr
in the laboratory; however, this observation was not
experimentally confirmed. The female is synovigenic
and required the permanent presence of the male for
normal egg production. At high population levels in
the field or in crowded laboratory cultures, as many
as 3 $ were observed to mount a single female at 1
time.
C. Preoviposition Period.—Oviposition initially oc-
curred 3-8 days after adult emergence.
D. Number of Eggs per Female.—The longest ovi-
positional period of a female was 18 weeks, during
which time 2715 eggs were deposited. The average
daily egg production of 10 9- was 17.2±5.4 eggs. The
daily mean, weekly, and cumulative egg production of
10 individual 9- as well as the mean weekly and cumu-
lative egg production of all 10 9 are shown in Table
3 and graphically depicted in Fig. 11.
E. Number of Eggs per Cluster.—A total of 100
field- and laboratory-collected egg clusters contained
an average of 30.8 eggs/cluster. The smallest clusters
contained 5-8 eggs; the largest cluster had 74 eggs.
Single isolated eggs were seldom observed. Under
conditions of crowding, lack of food, or unsuitable
FIG. 11.—Mean weekly (boldface numerals on ordi- food, eggs were occasionally laid end-to-end in strands
nate) and cumulative (smaller (light-face numerals) ovi- laterally affixed to the surface.
position by 10 L. t. daturaphila females. (Vertical lines
indicate confidence limits (5% level).) F. Productive Adult Life.—Beetles normally lived
546 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 63, no. 2
for 3-4.5 months. A marked decrease in egg produc- Essig, E. O. 1958. Insects and Mites of Western North
tion was noted among older females. Adults required America. McMillan, N. Y., 1059 p.
a continuous supply of food. However, they fed with Force, D. C. 1966. Reactions of the three-lined potato
beetle, Lcma trilincata (Coleoptera: Chrysomclidae),
greater moderation than larvae and withstood longer to its host and certain nonhost plants. Ann. Entomol.
periods of starvation. The daily feeding activity of Soc. Amer. 59(6) : 1112-9.
the adults was not measured. Harris, T. W. 1841. A report on the Insects of Massa-
chusetts, Injurious to Vegetation. Cambridge. 459 p.
G. Overwintering. — Active adult females were Kogan, M., and R. D. Gocden. 1969. A photometric
found in the Riverside area as early as Jan. 29, 1967, technique for quantitative evaluation of feeding
when their host plants were beginning to show re- preferences of phytophagous insects. Ann. Entomol.
newed growth. Some beetles remained active until Soc. Amer. 62(2) : 319-22.
late November. The only specimens found during the 1970. The systematic status of Lcma trilincata datura-
phila, new name, with notes on the morphology of
winter were a few adults hidden in sheltered loca- chemoreceptors of adults (Coleoptera: Chrysomcli-
tions, mainly the hollow seed pods of D. meteloides dae). Ibid 63(2) : 529-37.
still attached to the plants. A thorough search for Om.r-Coopcr, G., and P. Miles. 1951. On Lcma tri-
overwintering pupae in the soil was unsuccessful. lincata—a beetle closely resembling the tobacco slug
attacking the Cape gooseberry. S. Afr. J. Sci.
Therefore, this species probably overwinters as adults, 47(12) : 330-3.
as is the case with other species of Lcma. Riley, C. V. 1869. First annual report on the noxious,
beneficial and other insects of the State of Missouri.
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Blackaller, A. 1945. El mayate del tomate de cascara, Sand rson, E. D. 1913. Insect Pests of Farm, Garden

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Lcma trilincata (Olivier). Fitofilo 4(6) : 364-70. and Orchard. J. Wiley, N. Y. 684 p.
Blaisdell, F. E. 1893. Notes on the habits of some Sand rson, E. D., and L. M. Peairs. 1931. Insect Pests
species of Coleoptera observed in San Diego County, of Farm, Garden and Orchard. Ibid. 568 p.
Calif. Insect Life 5(1) : 33-36. Waldbau r, G. P. 1962. The growth and reproduction
Britton, W. E. 1932. Insects attacking the potato crop of maxillaectomized tobacco hornworms feeding on
in Connecticut. Conn. Agr. Exp. Sta. Bull 208, normally rejected solanaceous plants. Entomol. Exp.
Entomol. Ser. 26: 101-19. Appl. 5 ( 2 ) : 147-58.
1933. Plant pest handbook for Connecticut. 1. Insects. 1964. The consumption, digestion and utilization of
Ibid. Bull. 344: 68-182. solanaceous and non-solanaceous plants by larvae of
Carne, P. B. 1966. Growth and food consumption dur- the tobacco hornworm, Protoparcc scxta (Johan.)
ing the larval stages of Paropsis atomaria (Coleop- (LepHoptera: Sphingidae). Ibid. 7(3) : 253-69.
tera :Chrysomelidae). Entomol. Exp. Appl. 9 ( 1 ) : 1968. The consumption and utilization of food by in-
105-12. sects. Advan. Insect Physiol. 5: 229-88.

A Taxononiic Revision of the Termitophilous Subtribe Perinthina (Coleoptera:

Staphylinidae). I. The Genera Paraperinthus, Perinthodes, and Physoperinlhus
with a Discussion of Their Integumentary Glands and Their Relationships1
JACQUES M. PASTEELS 2 AND DAVID H. KISTNER
Shinner Institute for the Study of Interrelated Insects, Department of Biology,
Chico State College, Chico, California 95926
ABSTRACT
The genera Paraperinthus and Perinthodcs are rede- The integumentary glands of Parapcrintlms shcasbyi
scribed and illustrated. In addition, one new genus is and Physopcrinthus josensi are described. Their defense
described, PHYSOPERINTHUS. Thirteen new species and post pleural glands are very reduced and both genera
are described: Paraperinthus congoensis (Ccngo Repub- possess a median glandular structure in abdominal seg-
lic), P. dispar (South-West Africa), P. mohalei (South- ment VII, which is now known in all 4 genera of African
West Africa), P. natalensis (Republic of South Africa), Perinthina. Because this structure is also present in the
P. sbeasbyi (Republic of South Africa and South-West Corotocini and not in other Aleocharinae, a close rela-
Africa), Pcrinthodes baucaloxcnus (South-West Africa, tionship between the Perinthina and the Corotocini is
Republic of South Africa), P. coatoni (South-West indicated. Numerous pores belonging to either campani-
Africa), P. em.rsoni (Congo Republic), P. pntoriusi form sensillae or to special glandular cells were found
(South-West Africa), P. rapulum (South-West Africa), in the integument of Physopcrinthus.
P. trinrvoides (Republic of South Africa), Physoperin- The relationships between the genera and the species
thus coatoni (South-West Africa), P. jos.nsi (Ivory are discussed. All the termitophiles studied are host-
Coast). Many new geographic and host data of all specific at the species level.
species are presented.

In recent years many new collections of termi- other groups, these collections have yielded many
tophiles associated with the Nasutitermitinae in new species belonging to the subtribe Perinthina.
Africa have been made. Along with specimens of To do justice to the descriptions of the new genus
and species involved, it is necessary to revise the
1 entire subtribe, probably ultimately for the world, but
This study was financed in part by the National Science
Foundation (Grant no. GB-6284). Received for publication July right now the revision is confined to the species on
30, 1969.
2 the continent of Africa. The majority of these spe-
Permanent address: Universite libre de Bruxelles, Belgium.