STUDIES ON THE BIOLOGT, ECOLOGY,
DISTRIBUTION, AND CONTROL OP
EPICAERUS AURIFER BOH. (CURCULIONIDAE),
A NEW PEST OP ALFALFA IN
MEXICO
Dissertation
Presented In Partial Fulfillment of the Requirements
for the Degree Doctor of Philosophy in the
Graduate School of The Ohio State
University
By
George Mallory Boush, B. Sc., M. Sc.
The Ohio State University
1955
Approved by
Adviser
Department of Zool
and Entomology
ACKNOWLEDGEMENTS
The author wishes to express his sincere appreciation
to Dr* D# M# DeLong, Department of Zoology and Entomology,
The Ohio State University, for his inspirational guidance
and counsel in connection with this study and to Drs#
Douglas Barnes and John J# McKelvey of The Rockefeller
Foundation for their advice and encouragement#
Grateful acknowledgement is also made to Mr# Francisco
Gonzolez and Miss Carlota Riess H#, Mexican Agricultural
Program, for their assistance in preparing the drawings;
and to Miss Rose E. Warner, Dr# E# W# Baker, Entomology
Research Branch, U.S#D#A., Dra# Helena Brava and Dra#
Leonila Vasquez G#, University of Mexico, for determining
the species of insects, mites, plants, and mammels res*
pectively as referred to in this dissertation; and to
Mr# E. L# Sleeper who assisted in the preparation of
the section on taxonomy# I should also like to thank
The Rockefeller Foundation for making this study possible#
Finally, I should like to express my heartfelt
gratitude and thanks to my wife, Sara, for her constant
help and encouragement#
ii
Table of Contents
Page
Introduction............
1
Taxonomy
6
..........................
Description of the S t a g e s ........................... 8
Adult
....... 8
....................
Egg ............................................. 12
Larva
Pupa
.......
13
........
15
.....
Life History and Habits
17
Adults .......................................... 17
Qnergence
.......
Sex Ratio
........... .
Mating
18
...............
21
....
22
Preoviposition Period
Ovlposltlon
17
....
23
Number of Eggs and Egg Masses Laid per Female
27
Longevity
28
.....
Description of Damage
....
Behavior Studies
.........
Eggs «..................................
Incubation Period
....
Egg Development and Hatching.............
ill
30
31
32
32
36
Page
Larva ••••••.......
37
Number of Instars
......... .•................
Description of Damage
.♦♦♦3d
.............................. I4I4.
General Habits
Pupa
I4.6
.......................
............ ........ I48
Host and Food Plants
Natural Enemies
Predators
37
...........
53
..................
•53
Parasites........ ........ ..... .... .
Population Dynamics
57
............ •••••••
....«» 57
Distribution Studies.....
»••• 60
General Descriptionof the Area Studied
Method of Determination
......
60
67
.........
Control
....*• 78
Cultural
..............
Chemical
78
........
78
Laboratory ......................
78
F i eld ........................................... 80
Summary and Conclusions
Appendix
....
Literature Cited
Autobiography
........
89
93
.........
.....
100
•
iv
102
Introduction
The present study concerns the biology, ecology,
distribution, and preliminary control studies of the
curculio, Eplcaerus aurlfer Boh*, over a 2-year period
in south-central Mexico* Prom the time of the original
description of this insect by Boheman (IQJ4.O) until the
present, no work, other than Its inclusion in taxonomic
treatises is known. However, three other species of the
genus Eplcaerus; E* lmbrlcatus (Say), E. cognatus Sharp,
and E* formidolosus Boh. are recognized as pests of
agricultural crops. Eplcaerus lmbrlcatus. the imbricated
snout beetle, attacks a wide range of crops In the United
States, especially apples, potatoes, and strawberries;
also E* cognatus is a serious pest of potatoes In the
highlands of Mexico. The genus is apparently indigenous
to the Americas. All of the species recognized by Leng
(1920) and Blackwelder (l9i^7) were originally described
from the southern portion of the United States, Mexico,
and Central America*
Although not yet recorded in the literature, E.
aurlfer, in the larval stage, is a geographically res­
tricted but highly destructive pest of alfalfa* The
adult beetle also attacks the foliage of alfalfa as well
as the foliage of corn, soybeans, and field beans*
However, the principal damage to alfalfa is by the feeding
of the larvae on the primary root of the plant.
1
This tunneling and boring eventually results in the
weakening or destruction of the plant. Attacks by this
pest on a commercial basis result in a progressive
dieback or loss of stand in previously well established
alfalfa fields. Usually, heavily infested fields are
abandoned or plowed under aa a result of these attacks*
Its importance as a potential major pest can be
further realised when it is noted that alfalfa, under
irrigation, is the most important forage crop grown in
Mexico. Alfalfa, as food for cattle, forms the basis
for the dairy Industry. Its productivity is high. When
properly cared for and under favorable conditions it
may remain in production for 5 to 7 years, annually
producing as many as 9 or 10 cuttings. In addition,
certain areas of Mexico are famous for the production
of high quality alfalfa seed*
This problem was undertaken largely because of the
potential economic importance of this Insect and because
of the lack of information concerning its habits, range,
and control*
Inorder to carry out studies on various aspects
of the biology, ecology, distribution, and control of
E. aurlfer. the author established a program of field
and laboratory experiments. Life history studies, pop­
ulation dynamics, movement studies, biological control,
and determination of range and large plot control tests
were done in the field* The laboratory studies included
all physical measurements as well as determination of
the incubation period, longivity, egg-laying habits,
food plant preferences, and preliminary laboratory control
experiments*
As there is no published data concerning the location
and size of commercial alfalfa growing regions of Mexico,
a program was also undertaken in which the author visited
and mapped these areas*
To facilitate the studies on the biology and control
°?
aurlfer it was essential to obtain permission to
utilize an area heavily populated with this insect*
For insectary work on longevity, egg laying studies,
and laboratory insecticide screening, it was necessary
to be able to obtain several thousand individuals of
approximate known age at a given time. Similar land
was needed for field use in biological and insecticidal
control studies as well as population studies*
La Patera, a dairy farm of approximately 200 acres,
located on the northern outskirts of Mexico City, was
found to be heavily infested with E. aurlfer* The owner
of the farm, Mr* Francisco McCann, was greatly concerned
over the possible depredations of this Insect and gener­
ously consented to make available land and labor for
experimental purposes. The farm was completely irrigated
and planted in corn and alfalfa*
k
In late April 1952, on the McCann farm, a lj. acre
field of 3 year old alfalfa of the variety Oaxaca was
found to be particularily heavily infested with E* aurlfer*
This field was immediately designated as a biological
and population study area and efforts were made to keep
it in as undisturbed condition as possible* During the
spring and summer of 1952 and 1953 this field was obser­
ved and examined twice each week. During the remainder
of the year the field was examined at approximately
weekly intervals*
The progress of the various life stages was largely
determined by examination of 10 plants, selected at
random, in the above field at regular weekly intervals*
The larvae and pupae collected at this time, together
with collection data, were placed in vials containing
Peterson* 8 (1914-7) K.A.A.D. solution and later transferred
to 80 per cent ethyl alcohol* These specimens were later
measured and carefully observed for possible parasites*
Samples of plant tissue damaged by this curcullo were
also collected at the same time and placed in 80 per
cent alcohol for more critical observation In the lab­
oratory*
Measurement of adult population fluctuations were
secured by placing five 1 square meter quadrats, at weekly
Intervals, during the rainy season on the alfalfa field
designated for biological studies*
5
A similar alfalfa field, located adjacent to the
biological studies field, was utilized for control exper­
iments* One portion of this field was used to determine
the effects of overgrazing in controlling this pest*
An equal portion was used to determine the effects of
flooding in controlling E. aurlfer* Also at La Patera,
insecticide treatments were applied to the soil of
seedling alfalfa on a field basis as well as foliar
applications of various insecticides to fields of alfalfa
infested with adult curculios*
All of the insectary and laboratory tests on E.
aurlfer were carried out at the Agricultural Experiment
Station at Chapingo located 23 miles southeast of Mexico
City at the National School of Agriculture* Adequate
greenhouse and insectary facilities were available at
Chapingo for these studies* Specimens collected at La
Patera were brought to Chapingo for more critical biological
studies* Cages were also constructed at Chapingo in the
insectary to house various small mammals utilized in
natural predator studies*
6
Taxonomy
Eplcaerus aurlfer Boheman (Original Description)
Boheman, C.H. l81j.O. Schoriherr Genera
et Species, Curculionidium, Vol. VI,
Part 2, p. 278.
Eplcaerus aurlfer Boheman
Sharp, David. 1891. Biologia CentrallAmerlcana Coleoptera, Curculionidae,
Vol. IV, Part 3(1889-1911), P* 119*
Eplcaerus aurlfer Boheman
Dalla Torre, K. W. von and M. P. van
Emden. 1936. Coleopterorum Catalogus
pars ll|.7 , p* lf-2 •
Eplcaerus aurlfer Boheman
Blackwelder, R. E. 19kl* Checklist
of the Coleopterous Insects of Mexico,
Central America, The West Indies, and
South America. U.S. Nat. Mus. Bui. 165,
Part 5, p* 797*
In Sharp (1891) and Blackwelder (19lj.7)f Eplcaerus
ravldus Is synonomized with E. aur1fer.Sharp admits that
he created this synonomy without making an exact comparison
of the types of these two species. He further Indicated
that E. ravldus would prove to be a thickly-scaled and
finely-punctured fcrm of E. aurlfer.
7
If these two species are synonomous, this species
must take the name E* ravldus as this species was des­
cribed on the page preceding E* aurlfer in the same
publication*
Until the types of these two species can be crit­
ically examined by a person familiar with the variabi­
lity in the genus Eplcaerus. it would be less confusing
to consider these two forms as distinct species* It would
seem preferable to retain the name aurlfer In the litera­
ture as It Is the name under which it has been recognized
for over $0 years*
8
Description of the Stages
Adult
The adult beetle Id robust and more or leas cylindrical in form (Figure 1)# The antennae are scaly, strongly
elbowed, inserted at the sides of the beak in deep antennal groves, and 11 -segmented# The last 3 segments of
the antennae form a compact annulated club# The mouthparts are well developed# The paired mandibles are short,
stout, and pincer-llke, possessing a characteristic
conical depression resembling a scar on their outer
surfaces# The beak is short and stout although longer
than the head, not constricted between the eyes, slightly
dilated and strongly notched at the tip# The prothorax
is highly arched with the apex truncate. The elytra
with lij. to 16 rows of indistinct punctures are fused
and completely cover the abdomen when viewed from above.
The inner wings are rudimentary and the metanotum memb­
ranous# The front coxae are contiguous, the middle and
hind coxae are slightly separated# The third segment
of the tarsi is wider than the second and deeply bilobed#
Depending largely upon age, the body may or may not be
densely covered with fine silvery-gray scales# The adults
range from ash-gray to black in color#
The sexes in some cases may be difficult to differ**
entlate without disturbing the genitalia* Usually, how­
ever, in lateral profile the abrupt angle formed at the
apex of the elytra in the female is considerably unlike
the evenly rounded angle formed at the apex of the elytra
in the male* Also, as seen dorsally, the distal portion
of the head of the female is usually greater than half
the width of the thorax, whereas the head of the male
is usually less than half the width of the thorax* In
general, the females are larger than the males.
A summary of the measurements of lengths and widths
of 200 males and 200 females are given in Table I*
Measurements of width was taken across the widest portion
of the elytra, and of length as the distance from the
apex of the snout to the apex of the abdomen*
10
Table I
Summary of measurements of adults;
aurlfer. June 1952,
Chapingo, Mexico*
Sex
Number
Measured
Length (Mm.)
Minimum Maximum Mean
Width (Mm.)
Min* lltax. kean
Male
200
6.9
12.0
10.2 2.5 k>9
Female
200
8.2
llj.,5
11*7 3*0 6.0
3*9
11
Epic r e r u s h ur ifer Bqh
Figure
1
.- Dorsal and lateral aspects of the adult stage
of E* aurlfer*
The eggs are ovoid in shape, slightly tapering
toward the ends (Figure 2-A). When first deposited the
eggs are opaque but later tend to gradually become
translucent as the incubation period increases. Just
prior to hatching the dark head capsule of the embryo
can be clearly seen within the chorion. The chorion
is not superficially marked in any way.
Measurements of 93 ®ggs» within I4.8 hours following
deposition, were made with a dissecting microscope and
an ocular micrometer. The results of these measurements
are summarized in Table II.
Tabid II
Summary of measurements of eggs of EU aurlfer. July 1952,
Chapingo, Mexico
Number
Measured
93
Length (Mm.)
Width (Mm.)
Minimum Maximum Mean Minimum Maximum Mean
1.20
I.I4.6
1 .314. 0.20
O.I4.6
0.32
Color
White
13
Larva
The fully developed fourth instar larva is cres­
cent-shaped or cyphosomatic (figure 2-B). The body is
fleshy, near-white to cream in color, and composed of
13 segments, each showing 2 plicae on the dorsal surface.
All segments with the exception of the posterior 3 or
1^. are equal or nearly equal in size. The larvae are leg­
less; however, the 3 thoracic segments are distinctly
knobbed. The ventral portion of the body bears Ij. to 8
indistinct setae per segment. The abdominal spiracles,
shown in detail in Figure 2-B, are 9 in number and are
found one per segment per side. They are of the annular
biforous type.
The head is distinct, usually exposed and varies
in color from cream in the newly hatched first instar
larva to dark brown in the fully developed fourth instar
larva. The mouthparts are hypognathous, and fitted with
paired heavily sclerotized mandibles. Antennae are present,
but small and indistinct; ocelli are absent.
A summary of the measurements of the larval stages
are shown in Table III. It was also from this table that
the number of larval instars of this insect was calcu­
lated.
Table III
Summary of measurements, in millimeters, of larvae; Eplcaerus aurlfer.
May 1953» Chapingo, Mexico*
Inatar
Number
Wdth. Head £ ap~
Measured Min. Max. Mean
Lth. of Body
Wdth. of Body
Max. Mean Min. Max. Mean
Min*
1*
38
0 *2 8
0 .3 6
0.33
i.ia
1.59
1 .5 2 0 .3 8
0 .5 9
0.1+7
2.
63
0.51+ 0 .8 0
0.65
3 *2 8
5 .2 6
1+.32 0.68
1.1+2
1.02
3.
39
1 .0 3
1.77
1.33
1+.33
3.98
U .6 7 i.ia
1.91
1*67
k.
36
2.10
2 .8 1
2 *14-6
8.9
1+.6
3.1+
11.1+
1 0 .3
2 .2
15
Pupa
As the pupal period approaches, the larva, In the
prepupal stage, shortens and becomes barrel-shaped*
The rather Indistinct segments become deep and well
defined* The head at this time Is deeply retracted
Into the body*
The actual metamorphosis from larva to pupa has
not been observed*
The pupa (Figure 2-C and D) is of the exarfite
type; that is, possessing free appendages* The spiracles
occur as prominent knobs and might possibly serve as
taxonomic characters* Fourteen pupae were measured
and found to have an average length and width of 7*2
Mm* and 2*9 Mm* respectively* The minimum width and
length recorded were 2*6 Mm* and 6*9 Mm*, and the
maximum length and width were 7*6 Mm* and 3*2 Mm*
respectively* The pupa is light brown in color*
16
Ep ie h e r u s
h u r if e r
Bdh
Figure 2.- A, egg stage* B, lateral view of larva*
C, and D, ventral and lateral view of pupa*
17
Life History and Habits
Adults
Emergence
Three alfalfa fields, heavily damaged by the at­
tacks of E*. aurlfer. were observed over a period of
two seasons to determine the time of emergence of
this pest* The fields were located at La Patera dairy
and Chapingo, Mexico, as located previously, and Chaleo,
Mexico* Chaleo is located 30 miles south of Mexico
City* All three fields were in the Mesa Central region
of Mexico and shared approximately the same altitude
and climatic conditions.
Emergence of E* aurlfer in both 1952 and 1953»
at the three areas under observation, always occured
within 2l+ hours following a series of early spring
heavy rains*
The newly emerged individuals, covered with mud,
were usually found within the crown of the alfalfa
plant, head down and half covered with dirt and debris*
The sexes at this time were approximately equal*
Caged males and females refused to copulate or eat
for a period of 2 to 1+ days following emergence.
Also, for two or three days after emergence the insects
frequently defecated a semi-solid substance*
The results of the observations on time of emer­
gence are given in Table TV.
18
Table IV
Dates of emergence of E. aurlfer in 1952 and 1953*
Year
La Patera. P.F.
Chaleo. Mex.
Chapingo. Mex»
1952
May 27
June 8
June 7
1953
June 7
June 11
June 11
Sex Ratio
The sex ratio of E. aurlfer varies with time as
there exists a definite difference between the longe­
vity of the males and females. Observations on sex
ratio would therefore largely depend upon the age of
the sample studied.
In this study, sex ratio was determined by field
collected beetles, In which a
25
square meter quadrat,
replicated ij. times was utilized. Collections were
made at La Patera at I4. periodic times during the season.
As shown in Table V, the sex ratio at the beginning of
the summer was almost 1:1. As the summer progressed,
the ratio of males to females decreased, eventually
reaching a ratio of nearly
1
:3 *
19
Table V
Sex ratios of EU aurlfer at periodic Intervals
during the 1953 season* La Patera, D* P*
Dates
Number
Collected
Per cent
Per cent
Approx*
Male_______ Female______ Sex Ratio
June 15
2,530
lj.6.7
53*3
1:1
July 12
1,809
37*8
62.2
1:1*5
Aug. 9
l,3Ui|-
32.9
67.1
1:2
236
27.0
73*0
1:2.3
Sept. 6
20
Figure 3*- Potted alfalfa and cellulose acetate cages
utilized in the biological studies conducted
in the greenhouse.
■
21
Mating
A series of cages (Figure 3), constructed of cel­
lulose acetate In the form of hollow cylinders 10
inches In diameter and 22 inches high was used to
observe the mating habits of E. aurlfer* Muslin was
sealed over one end of the cages with acetone, and
two 3 inch round windows cut in the sides of the cages
and sealed with fine copper wire were provided for
ventilation. Each cage was placed over a 12 inch flower
pot containing a single 2 year old alfalfa plant of
the variety Oaxaca.
A newly emerged male and female were placed in
each of the ij.2 cages used in these observations.
Mating did not usually take place until 3 or ij.
days after emergence and occured on the ground as well
as on the stems of the plant. Copulation occurs as fol­
lows: the male approaches the female from the side
or rear, and if not prevented by movements of the
female, mounts from the rear. If she is receptive and
does not flee, he clasps her elytra with first his
front and middle legs and finally with all three pairs
of legs. The aedeagus, which is sheath-like, is then
inserted in the fully extended ovipositor. During the
lengthy copulation, little movement on the part of the
male takes place, although the female may walk about
and frequently feeds.
22
The time for 35 matings ranged from 16 minutes to
31 hours; the average being 3^ hours* Mating usually
occured at night, early morning and on overcast days#
In 9 cages containing 3 marked males and 1 female,
all the males were observed to mate with the female
within 72 hours*
Of the caged pairs, copulation took
place an average of 3 times during the 3 month duration
of the observations.
Although egg production reached its peak during
early July and fell off rapidly thereafter, mating
continued until early fall*
In a separate series of 21 cages utilizing a sin­
gle newly emerged female in each cage, no eggs were
deposited. Apparently mating is necessary for egg pro­
duction#
It Is not known if frequent copulation Is necessary
in the field for maximum egg production. Possibly the
frequency of copulation in the greenhouse was due to
caged conditions or environmental factors different
from those found in the field#
The mating observations in the greenhouse occured
at temperatures ranging from 6l to 88 degrees Fahren­
heit#
Preovlposltlon Period
The preoviposition period of the female varies
from 3 to 7 days# In a series of I4.2 cages, as described
23
under the prior topic, in which 1l 2 newly emerged females
were allowed to mate once, the average preoviposition
period for 31 females which produced eggs was 3.9 days.
Of the 11 unaccounted for Individuals, Ij. died prior to
egg deposition and 7 deposited no eggs.
Observations at La Patera in the biological study
field were in accord with the laboratory results.
Although densely populated with newly emerged adults,
no egg masses were found within a week of emergence
of the first individuals.
Dissection of newly emerged unmated females re­
vealed rudimentary ovaries with no visible eggs, to­
gether with a large mass of fatty tissue. Further
dissection of females which had been allowed to mate
2 days prior to the observations showed a decrease
in the fatty tissue and the partial development of
recognizable eggs.
Oviposltlon
The eggs are deposited in ovoid or linear groups
(Figure 2-A) on the top half of the host plant. There
are two methods of deposition: the eggs are deposited
in the fold of a single leaf or leaflet (Figure i^.);
or they are deposited between two opposing leaves or
leaflets. A sticky, fast-drying substance secreted by
accessory glands located near the ovipositor aids
2k
in anchoring the eggs to the surface of the leaf, or
substrate, as well as sticking the edges of the leaves
together*
Oviposition usually occurs at night or in early
morning. The females are easily disturbed at this time
and feign death readily. Six apparently complete acts
of oviposition were observed. They varied in duration
from approximately 20 to 36 minutes, averaging 25
minutes* Temperatures during the observations ranged
from 6 I4. to 79 degrees Fahrenheit*
A female in the process of egg deposition is
shown in Figure 1|* Prior to oviposition, the female
walks rapidly up and down the stem of the plant, oc­
casionally examining a leaf with her antennae and
front legs* When a suitable site is found, the leaflet
or leaves are pulled or bent into position by the
legs to form a trench-like depression*
The actual
egg laying act occurs with the head of the female
directed away from the plant* The ovipositor is tele­
scopic in action and when fully extended is about
half the length of the elytra. The under surface of
the leaf is usually preferred for oviposition and the
eggs are deposited from the distal part of the leaf
Inward, in a single layer usually in a ovoid mass but
occasionally in a straight line*
25
Of 368 egg masses collected at La Patera June 21,
1952, 59 per cent were formed by the folding over of
a single leaflet and 1(1 per cent by the union of two
leaflets*
Figure I4.#- Adult female beetle ovipositing on alfalfa
27
Numb or of Eggs and Egg Masses Laid per Female
A series of tests were conducted in June and July,
1952 and 195*3 at Chapingo to determine the number of
eggs and egg masses laid per female* A similar series
of Ij.2 caged alfalfa plants, as described previously
under Oviposition. were used during both years in these
observations, A newly emerged male and female curculio
were introduced in each cage. The newly produced egg
masses were tagged daily. Temperatures in the green­
house during the tests ranged from 66 to 89 degrees
Fahrenheit,
Of the 82 caged pairs, 53 females deposited from
1 to 7 egg masses, averaging 2,17 egg masses, each
containing from 9 to 55 eggs with the average being
2lj.,6 eggs per mass. The remaining 29 females died
prior to egg deposition.
Field collections and observations at La Patera
during June 1952 on 202+. egg masses showed a range of
from ij. to lj.9 eggs per mass, with the average being
25*7 eggs per mass.
More than 75 per cent of the egg masses in the
laboratory tests were deposited from June 15 through
July 15, However, egg production remained light and
sporadic into the middle of September,
In the field, the egg masses have been collected
28
from J4. plants, alfalfa, corn, beans, and Amarantus
S£.
In the laboratory, in 6 caged flats containing
alfalfa, corn, beans, wheat, and Irish potatoes, egg
masses were deposited on alfalfa, corn, and wheat.
However, only 1 egg mass containing 7 eggs was found
on wheat and 2 egg masses containing 12 and 18 eggs
respectively was found on corn, whereas, 21 egg masses
were found on the alfalfa plants. Each flat contained
10 pairs of newly emerged beetles*
Longevity
In the laboratory tests conducted at Chaplngo
on preoviposition and oviposition observations as well
as the determination of the number of eggs laid per
female; studies, utilizing the same caged paired adults
were carried out to determine longevity* These obser­
vations were apparently of little value as the longev­
ity of the adults caged in the laboratory wasnearly
one month less than the longevity of similar beetles
caged in the field. The laboratory tests were aban­
doned on August 20, 1952 and on August 17# 1953 because
of the death of the beetles* The longevity of this
species, as an adult population in the field, during
1952 and 1953 Is given in Table VI*
29
Table VI
Presence of adults of E., aurlfer at 3 locations
In Mexico during 1952 and 1953*
Location
Plrst
Obs,
Last Presence
Obs* / (Days)
Chapingo,
Mexico
6/7
10/2
125
6/11
10/6
117
Chaleo,
Mexico
6/8
10/2
126
6/11
10/7
118
La Patera,
D. F.
5/27
126
6/7
10/3
118
9/29
First tast
Obs* Obs*
Presence
(Days)
30
Description of Damage
Alfalfa Is thought to be the only cultivated crop
seriously Injured by E* aurlfer* Injury Is primarily
the result of the feeding of the larvae on the primary
root, however, the adults also damage alfalfa by feeding
upon the foliage (Figures 5 and 6)*
Although probably of little economic significance,
the adult has also been observed, by the author, feeding
occasionally on other cultivated crops* These crops
are Its follows:
1.- Com- Zea mays L*
2.- Beans- Phaseolus vulgaris L*
3*- Soybeans- Sola max L.
ij.*- Wheat- Trltleum sativum L*
The foliar damage to these crops as a result
of the feeding action of the adult beetles may be
described as a general scalloping of the leaf margins
and occasionally the severing of a leaf or leaflet
from the petiole* Feeding may occur over the entire
plant but usually it is restricted to the young tender
growing portions* In alfalfa, this damage is partic­
ularly important as the new shoots in the crown of the
plant, representing the new growth, are frequently
seriously damaged or destroyed.
31
Behavior Studies
Probably the most obvious biological observation
concerning the adults is that they are negatively
phototropic. Eating, mating, oviposition, and movement
from one location to another usually take place at
night or under subdued light conditions. The adults
have been repeatedly stimulated into activity in the
greenhouse during the day by darkening the cages in
which they were located.
Generally from 9 to 10 A.M., until 5 or 6 P.M.,
C.S.T., the adults are inactive and are found under
loose litter, debris or dirt. Often, the head and thorax
are buried in the soil or debris and the entire abdomen
is exposed.
During periods of activity the adults are easily
disturbed and "feign death". At such time the legs
are doubled up under the body and the insect may remain
immobile for 10 or 1$ minutes. They may usually be
revived by placing the beetle in the palm of the hand
and gently blowing upon it.
Eplcaerus aurlfer was rarely collected singly in the
field. Usually, when located, they are in or near alfal­
fa and present in great numbers. They do not appear
to be gregarious but are found locally in large numbers
probably because of adequate food and suitable environ­
ment.
32
Eggs
Incubation Period
A total of 63 egg masses, deposited on alfalfa of
the variety Oaxaca, were tagged within 2i+. hours of
deposition and observed at daily intervals. The tempera­
tures in the greenhouse during the duration of the obser­
vations ranged from 66 to 89 degrees Fahrenheit*
The incubation period varied from 27 to 3^ days*
The results of these observations are shown in
Table VII.
33
Table VII
Incubation period of egg masses of B* aurlfer.
Chapingo, Mexico, 1952*
Incubation
Number
Period
Masses
Per cent
(days)_______________Hatched__________________ Hatched
26
0
0
27
2
3.2
28
6.k
29
k
Ik
22.2
30
20
31.7
31
11
174
32
9
H4-.3
33
2
3.2
3k
1
1.6
35
0
0
Total
63
100
311-
Figure 5*- Alfalfa leaves showing characteristic
feeding damage by the adult stage*
35
Figure 6«« Alfalfa plant showing the results of adult
feeding (Note characteristic crown injury)*
Egg Development and Hatching
When first deposited and during the first week of
incubation, the eggs are opaque and pearly white. During
the second and third week they appear grey; and during
the last 8 to 10 days are translucent and almost color­
less. During this later period, the head capsule of the
embryo can be seen within the chorion.
Observations were made at the time of hatching on
3 egg masses. The larvae utilize their mandibles in
rupturing the membraneous chorion and escape from the end
of the egg. The actual time elapsing from the initial
rupturing of the chorion until the larva clears itself
of the egg ranges from 2 to 6 minutes, averaging approx­
imately 3 minutes.
The majority of the individuals of a given egg mass
hatch at the same time. One egg mass containing 18 eggs
hatched in 28 minutes; another containing 1+3 eggs required
72 minutes for 1+0 of the eggs to hatch. A third egg mass,
containing 12 eggs, required 125 minutes for complete
hatching.
Eighty-four egg masses were collected July 6, 1953
from the biological study field at La Patera and brought
to Chapingo for observation. The egg masses, each in a
single petrl dish, were placed in a constant temperature
cabinet controlled at 70 to 72 degrees P. and 70 to 80
37
percent relative humidity. Only 12 individuals, from a total
of 2187 eggs, hatched. Possibly the living leaf forms an
environmental microclimate, difficult to measure and du­
plicate, which is essential for successful incubation.
This latter possibility might form the basis for some
form of culturally controlling E. aurifer.
Also, the long incubation period required, and the
vulnerability of the egg masses on the upper portion of the
plant might be considered a weak link in the life cycle of
this pest.
Larva
Number of Instars
Since the larvae go into the ground immediately upon
hatching and undergo all of the growth stages there, it was
not feasible to observe a series of individual larvae to
determine the number of instars and the time elapsed in each
stage. Also attempts to collect larval cast skins and head
capsules by screening and flooding the soil were unsuccess­
ful.
To determine the number of instars as well as other
biological data, a regular weekly collecting system was
established. Prom mid-April, 1952, until December, 1953»
a collecting trip was made each Saturday to the biological
study field at La Patera, Ten alfalfa plants were dug each
38
week and examined, along with the associated soil for lar­
vae. The larvae, together with the collection data and the
number of larvae per plant, were placed in K.A.A.D. solution
for examination in the laboratory.
From these field collections it was determined that E,
aurlfer has one complete generation per year. No full grown
larvae or pupae were found in the soil from the middle of
June through August.
Measurements of the width of the head capsule of 176
larvae, as well as length and width of the body, from the
above collections at La Patera are given in table 3* The head
capsule widths separate into 1+ fairly well defined groups.
To determine if an ecdysis had been overlooked, the principle,
as proposed by Dyar (1890), that the widths of the head of
a larva in its successive instars follow a regular geomet­
ric progression in their increase, was applied.
The average widths of the head of successive larval
instars was as follows: 0.33; 0.65; 1*33 and 2.i|6 mm. The
ratio of increase was found to be .5U« Applying this ratio
to the measured widths, the following calculated widths
were obtained: 0.35; 0 .72; 1.33; and 2.1(.6mm. On this basis
it is thought that the larva complete l± instars.
Descrlption of the Damage
Alfalfa is the only plant that the larvae have been
found near or collected from in the field or laboratory. The
39
damage caused by the attacks of the larvae on the roots of
the plant are usually brought to attention by the obvious
progressive dleback and loss of stand in well established
alfalfa fields, (Figures 7 and 8 ). On examination of a
given plant, the primary root is seen to be severely injured
(Figures 9 and 10 ), b^ the excavations and tunnels caused
by the feeding of the larva of E.aurlfer. In general the tun
nels are directed toward the central portion of the root.
The principal damage to the plant appears to be by the de­
cay of the entire central portion of the root, apparently
caused by the entrance of soil fungi and micro-organisms, (F
gure 10).
This decay is rapid and takes place from late summer
through the winter to early spring. Loss of stand, as a
result of the death of the plants, occurs largely in the
winter and early spring.
In a badly infested 3-year-old field of alfalfa of
the variety Kenya at La Patera in April, 1952, a count of
i+OO alfalfa plants, collected by following a ll^-inch plow,
disclosed that 96 per cent of the primary roots were in­
jured. As a result of this damage, the field was taken out
of cultivation and replanted in com. Although a high
population of larvae remained in the soil, the seedling
c o m was not injured.
:
1*0
Figure 7*- Normal three year old alfalfa field (Variety
Oaxaca)*
Ui
Figure 8*- Three year old alfalfa field (Variety Oaxaca)
showing results of attack by the larvae*
Figure 9«~ Alfalfa plants showing damage caused by the
tunneling and feeding In the primary root
by the larval stage*
o
c.
Figure 10#- Cross section of primary root of alfalfa#
A, Injury to central portion of root# B, Healthy root
C , Injured root showing entrance hole#
hk
General Habits
The larva, in all stages, is subterranean. As the
newly emerged forms make their way out of the egg, they
Immediately fall to the ground. By alternately contract­
ing and expanding they rapidly move into the soil at the
base of the plant. Observations have shown that
newly-
emerged larvae perish within an hour or two if the soil is
especially dry and hard.
No observations have been made on the feeding habits
of the first instar forms. Studies after the newly-emerged
larvae enter the soil are difficult because of their minute
size and fragility.
Observations on potted alfalfa inoculated with known
age larvae, in the greenhouse at Chapingo, have indicated
that the larvae pass through the first instar within 10 to
lij. days.
The second instar larvae have been observed feeding
on the primary root. Usually they appear unable to penetrate
the tough central portion of the root and confine their feed­
ing to the soft fleshy epidermal and cortex regions.
The time required for completion of the second instar
in the laboratory varied from 3 to 7 weeks.
The third and fourth instar larvae are found in about
equal numbers throughout the fall and winter. In early
April more than 80 per cent of the larvae are In the fourth
h5
lnstar.
Feeding continues through the fall and winter. The
third and fourth instar forms are responsible for most of
the damage to the primary root. Feeding on the root occurs
from ground surface level to as deep as 53 inches. Most
of the root damage to 3-ysar-old alfalfa occurs from a
depth of 2 or 3 inches to about 16 inches.
In the weekly observations of infested plants at
Chapingo during 1952 and 1953 » f©w larvae were actually
recovered from the root proper. Usually the larvae were
found within 1 to 2 inches of the root. From the almost
haphazard riddling of the root, and the few individuals pre­
sent, it would seem that the larvae move frequently, in the
soil, up and down the shaft of the l*oot, repeatedly feeding
and retiring to the adjacent soil. Damage to the secondary
roots by the larvae may occur, but has not been shown to
take place.
In the greenhouse the larvae appear to be sensitive
to varying degrees of moisture in the soil. When water Is
withheld from the plants preparatory to sifting the soil
for larval recovery, the insects are usually at the very
bottom of the pot where the soil moisture is greatest.
When observations were made following a daily watering
fcoutlne, the larvae were found at all levels in the soil.
In the field, from 1 to 3h individuals have been re-
covered from the soil around a single plant. Although diffi­
cult to form an average, usually 2 or 3 larvae are found
per plant.
Pu£a
The pupa is subterranean. It is found in an ovoid
earthen cavity located at a depth of from ^ to 1). Inches
below the surface of the ground. Usually, the pupal cavity
is within Ij. or 5 inches of the root and is often found with­
in the spreading basal crown of the plant.
The pupa when first transformed is near white but
gradually becomes light brown, and just prior to emergence
takes on a definite grey color.
Although a few (less than 1 per cent) individuals pass
the late fall and winter in the pupal stage, most transform
from prepupae to pupae about the end of April. The pupalstage lasts approximately 1 month.
The life cycle of E. aurlfer is given in diagrammatic
form In Figure 11. The graphs are not quantitatively acc­
urate, but indicate the trend in population size.
■—
& £
(Mi
Ssm.
i.—
&sl.—
V M M
Figure 11
Host and Food Plants
In this discussion, host plant Is limited to those
plants, In the field, on which the female deposits eggs
and on which larval development takes place. As such, E.
aurlfer is known to have but one host plant, Hairy Peruvian
Alfalfa, Medlcago satlva L,
In this study, Food Plant refers to those plants which
serve as a source of food. Eggs may be deposited upon a
food plant and not alter its status if larval development
does not take place. In addition to the cultivated food plants
of this beetle given on page 30, the following wild plants
have been recorded as serving as food for the adults.
1 .- Solanum rostraturn L.
2»- Anoda hastata Cav.
3 .- Eruca satlva Lam.
]+.- Rumex obtuslfollus L.
5.- Malva parvlflora L.
6 .- Paspalum sp.
7.- Abutllon sp.
8 .- Cucumls sp.
9*- Amarantus sp .
The latter plant, Amarantus sp.. of the pigweed group,
serves as an excellent food plant and is usually devoured
when occurring on the margins of a heavily infested alfalfa
field. Egg masses have been collected from this plant but
apparently the larvae cannot develop upon it as they have
never been found in the soil associated with the roots.
Larval development would further be curtailed as this plant
is an annual and dies in late summer, about the time the lar­
vae would be half developed.
Because of the possible deleterious effects of this
species on other cultivated crops, an effort was made to
determine the relative food plant preferences of selected
cultivated crops by the adult beetles. Four cages, con­
structed of screen wire and one meter square in size, were
placed in the field at Chapingo June 20, 1952. The ground
surface area in each cage was divided into 6 equal portions,
and into these portions were introduced potted com, wheat,
beans, alfalfa, potatoes and tomatoes. The corn, wheat and
beans were 3 weeks old and the alfalfa was of 2-year-old
transplanted stock. The potatoes and tomatoes were of unknown
age but were approximately 8 to 10 inches high. One hundred
adult beetles collected from the biological study area at
La Patera were introduced into each cage. Observations
wero made 2lj., lj.8 , and 72 hours following introduction of
the beetles and the individual crops visually rated as to
the degree of injury caused by feeding. The results of these
50
observations are given in Table VIII. The adults seemed to
prefer, in descending order of preference; beans, com,
alfalfa, tomatoes, wheat and potatoes.
51
Table VIII
Pood plant preference teat utilizing selected cultlvated plants* Chapingo, Mexico, July 1952*
a
e
.
<!*ge Number
Plant
:
tVariety
A
B
;
c
Corn
:Rocomex 1
3
3
:
3
2
2
:
1
k
3
:
1+
e
Wheat
: -•••
e
Beans
:Gto«10-A-5
2
Alfalfa :Oaxaca
2
Potatoes :Alpha
t
:
t
•
•
D
JRating
•
e
•
a
e
•
3
3
1
1.5
2
3
3.5
2
2
2.5
1
1.5
2
2
e
2
3
2
1
:
3
:
2
:
2
•
Tomatoes :Rutgers
2
#
•
2
<
(
•
e
•
•
•
•
•
•
•
•
Rating Scale
1«- No feeding
3*»- Moderate feeding
2*- Light feeding
lj.*- Heavy feeding
Figure 12*- A general view of the countryside in
the Mesa Central Region of Mexico*
53
Natural Enemies
Predators
Soon after the adult beetles were first seen in the
field in the early summer of 1 9 5 2 , it became apparent that
natural enemies were probably the most important single
factor in the reduction of the adult population. On June
7, in the biological study area at La Patera, an average
of l±2 live beetles were recorded per square meter. Prom this
date through June 2 1 , the number of live beetles per
square meter had been reduced, by natural enemies, to 2 7 *
An accurate index as to the efficiency of these enemies
could be determined by a count of the hard empty elytra
left upon the ground following the apparent eviseration
of the beetles. Peces, of a size and shape similar to those
of a housecat, were frequently found associated with the
remains. When these feces were placed in alcohol and ex­
amined with a dissecting microscope, they were found to be
composed almost entirely of cuticle fragments of E. aurifer.
From this evidence it was thought that the enemies func­
tioned as predators. Similar eviserated adults and asso­
ciated feces were also seen at Chaleo and Chapingo. At
the latter location, an insecticide test against the adults
had to be abandoned on July 1 6 , 1953> because of the reduc-
51+
tion in beetles by this means.
To determine the predator or predators involved, a
bounty was offered to the field laborers at Chapingo for
the capture of any small mammals native to the area.
A total of 12 specimens representing 5 species were
procured in this way. The mammals, with the exception of
a shrew, were placed in wire cages in the insectary and not
fed for 2i| hours following their capture. The feces ex­
creted by each of the animals during this time were examined
and compared with the feces containing the portions of E.
aurlfer collected in the field. The following day, food
(-I cup of dry shelled com), water and 100 live beetles
were introduced into each cage.
In these studies, 3 pocket gophers, Cratogeomys
merriami merriaml; a small spotted skunk, Spllogale sp.;
a large striped skunk, Mephitis sp.; 6 ground squirrels,
Cltellus mexlcanus mexlcanus; and a shrew, Sorex saussurel
saussurel Merriam, were observed.
All of the above species, with the exception of the
pocket gopher, eviserated some of the beetles. Both species
of skunks ate the 100 beetles introduced into their respect­
ive cages within ij. hours and produced feces, as well as
eviserated curculios similar to those found in the field.
Also, 5 of the 6 ground squirrels eviserated and ate from
55
6 to 28 beetles within a i^-hour period following their
introduction into the cages. Afterwards, possibly because
of the presence of the com, the ground squirrels refused
to molest the beetles when they were Introduced into the
cages. The skunks, however, refused to eat the corn and re­
peatedly devoured the beetles soon after they were placed
in the cage.
The pocket gophers were found to be vegetarians and
did not eat any of the beetles. These mammals are frequent­
ly found in alfalfa fields where they are considered a pest
because of their mounds and tunnels; however, they apparent­
ly eat little but the stems and leaves of plants.
The shrew died, probably of starvation, about i^8 hours
after it was captured. Within this time it eviserated ij.8
beetles. The dead beetles were unlike those found in the
field in that the eviseration in this case was accomplished
by partially separating the ventral cuticular surface of
the abdomen from the elytra, rather than the destruction
of the entire curculio except the elytra.
The feces produced by the ground squirrel and shrew
are unlike in size, shape, consistency or color, the feces
associated with the effects of the predators in the field.
Unfortunately, these observations were limited to
laboratory conditions and do not necessarily reflect the
actions of these mammals in nafeure. However, as a result
56
of these observations, showing the close similarity between
the feces and eviserated beetles found in the field and
those produced in the laboratory, the two species of skunks
are thought to be predators of E. aurifer*
57
Parasites
During the summers of 1952 and 1953 at Chapingo, mites,
attached to the ventral membraneous areas of the neck of
E. aurlfer were frequently observed and collected. Speci­
mens sent to Dr. E. W. Baker, Entomology Research Branch,
were identified as belonging to the family Parasitidae.
More exact determination could not be made.
Individual curculios with attached mites were placed
alive in cages with food so that they might be observed.
The beetles did not appear to be inconvenienced by the mites.
Copulation, oviposition and feeding proceeded at an app­
arently noimal rate.
Population Dynamics
To determine properly the dynamics of a population
of a given animal in the field during a known period of
time would involve simultaneous sampling at periodic in­
tervals, in replicated representative areas throughout
the range of the animal. Such a study would require con­
siderable time, money, and transportation facilities. As
these items were at a minimum, It was decided, in the
spring of 1952, to conduct a series of observations in a
single field known to contain an initial high adult pop­
ulation. The biological study area at La Patera was chosen
58
for this study because of its proximity to the office and
the abundance of beetles present.
The results of these observations are summarized in
Figure 13• The beetles were first seen in the field on
May 27, and reached their greatest numbers about 9 days later.
The steady decline in population between June 7 and July
5 was thought to be almost entirely due to the feeding
action of predators. The subsequent decline in population
was probably a result of natural mortality, particularly
of the males, as well as predators. The beetles were last
observed in the field September 27.
A similar study was undertaken at the beginning of the
1953 season, but for reasons unknown to the author, the
population of B. aurlfer had diminished to a level so low
that it was not possible to measure the fluctuations
during the season.
59
'Rcmr/L'E /Yi/mzRs ufzurcuuos f f r FQF
ffitf 1 - S U C F F C l / U F S / l S t f M
rcs
A 221
iH*
£
I**
_i*L
A
!•*
!
*
‘J?
id^
JL
-.=V-
•j^y
--_r?
--------------
V-"_
\ /w
t/l
|-A
/
s'/c^
§ s
\
A
./
A/-?
|§-tf
*<
K
_________ s /
Jog%
/
-._
r?
/A<£Y
— .2. _________ Sj'S-t'X
/
//-?^
/
•s
SH-^ 30
\
/
/A ^
/
lq--« l
.Lw
t8
!
i**X
Figure 13*- Adult population changes of S* aurlfer in a
alfalfa field in Central Mexico in 1952*
d
60
Distribution Studies
General Description of the Area Studied
The biological and control studies of L. aurifer
were carried out in the states of Mexico and Distrito Fed­
eral. (For reference purposes, a map with the states of the
Republic of Mexico, is shown in Figure l[|J. These two states
form a portion of a high, semi-arid plateau region, gen­
erally referred to as the Mesa Central geographic zone
(Wellhausen 1951)• This zone, as shown in Figure 15, varies
in altitude from about 6500 to 8500 feet and has a definite
dry and rainy season. The climate is generally temperate
throughout the year.
Because of the mountainous nature of south-central
Mexico, with the resulting high number of ecological
niches and microclimates, the area is extremely difficult
to classify. Leavenworth (1931) in his classification of
ffuevo Leon points out that
"classification based solely
on precipitation, temperature and evaporation data can never
be satisfactory for ecological purposes in mountainous
areas because (l.) data is often Insufficient or lacking
and (2 .) the relationship between climate and biological
phenomena does not lend itself to mathematical formulae."
Possibly a better index of the climate and soil of a
given region can be gained by an analysis of the vegetation.
61
Leopold (1950) prepared a map of the vegetatlonal zones
of Mexico, giving what he considered the original climax
vegetatlonal types. This map is shown in Figure 1 6 .
A photograph, showing a typical view of the general
terrain in the Mesa Central is given in Figure 12.
At present, the principal cultivated crops in this
region are corn, beans, wheat, vegetable crops and alfalfa.
Generally speaking, it Is probably the most productive
agricultural region In Mexico.
Although probably of limited use in field biological
studies, official meteorological data from the municipality
nearest the various study areas, expressed in the form of
climographs, is presented in the Appendix. The climographs
were thought to be helpful in the comparison of progressive
seasonal precipitation and temperature of one area with that
of another area. The climographs are of the type used and
described by Munns (1922) and Transeau (1935)* The tempera­
ture and precipitation data presented In the climographs
are from a 5-year summary of weather conditions In Mexico
published by Contreras Arias (19U2).
62
Figure ill*- Map of the Rebubllc of Mexico showing the
state boundaries*
SONORA
CHIHUAHUA
COAHUILA
SINALOA
N. LEON
DURANGO
ZACATECAS
\ S. LUIS
POTOSi"
AGS.
YUCATAN
GTO.
HIDALGO-
JALISCO
J MEXICO
COL.
MICHOACAN
pf-
;t l a .
CAMPECHE
MOR; puEBLA
TABASCO
VERACRUZ
GUERRERO
OAXACA
CHIAPAS
6k
Figure 15*- Map of the Republic of Mexico showing the
general geographic regions (Wellhausen)•
6$
Figure l6*« Map of the Republic of Mexico showing the
major vegetatlonal regions*
TEMPERATE
BOREAL FOREST
PiNE-OAK FOREST
CHAPARRAL
mesquite-grassland
I
j OESERT
TROPICAL
CLOUD FOREST
RAIN FOREST
TROPICAL EVERGREEN FOREST
C«5l SAVANNAH
TROPICAL DECIDUOUS FOREST
|===1 THORN FOREST
ARID TROPICAL SCRUB
MEXICO
VEGETATION MAP
MUSEUM O f VCfttCWUTC 2QCL0CT
vw <v>e«*rr c v c a u f o r n m
BY A S. LEOPOLO
67
Method of Determination
Two methods were used to determine the present dis­
tribution of E. aurlfer» The first method employed con­
sisted of numerous field collections of the beetles in
a progressively wider circle from the point of known
occurence; and the second consisted of requesting avail­
able collection data for this species from the larger
museums. The following museums cooperated in this respect,
1,- U. S. National Museum, Washington, D. C.
2,- British Museum (Natural History), London, England
3,- American Museum of Natural History, New York, N. Y,
14.,- Zoology Museum, University of Texas, Austin, Texas
5>»- Entomology Museum, Cornell University, Ithaca, N, Y,
6,- Snow Museum, University of Kansas, Lawrence, Kansas
7,- Museum, Illinois Natural History Survey, Urbana, 111.
8,- Entomology Museum, Ohio State University, Columbus, 0.
9,- Museum, La Fundacion Rockefeller, Chapingo, Mexico
10,-Instituto del Biologia, Largo de Chapultepec, Mexico,
D. F.
11,-Museum, Department of Emtomology, University of
California, Berkeley, California
A summary of the collection data of E. aurlfer fur­
nished by the above museums containing identified speci­
mens of this species is given in Table IX. Although the
collection data accompanying many of the specimens listed
68
Is Incomplete, all, with the exception of a specimen
In the British Museum labeled ”Sierre de Durango, Mexico” ,
are from the south-central portion of Mexico.
The author found E. aurlfer to be limited in its
distribution to the valloys of Mexico, Mesquital, Tehuacan,
Morelos and Morelia in south-central Mexico (Table X and
Figure 17). Within these valleys, It appeared to be fur­
ther restricted to local areas where alfalfa was grown.
Collections made on wild plants In the regions bordering
alfalfa growing areas were successful; however, collect­
ions made at random at a distance of a mile or more
from alfalfa were, without exception, unsuccessful.
As E. aurlfer was described originally from Mexico
in 18 )4.0 , prior to the introduction of alfalfa, there
must exist, or have existed, one or more native plants
upon which the larvae are capable of developing.
However, as this species is now thought to develop
and occur regularly on alfalfa, distribution was determined
by examination of representative fields in the major al­
falfa producing areas. These areas, as shown in Figure
1 8 , were located through the assistance of Drs. J. B.
Pitner and Reginald Laird, Agronomists with the joint
office of the Mexican Department of Agriculture and The
Rockefeller Foundation.
About 2,000,000 tons (U. S.) of alfalfa are produced
69
yearly on approximately 100,000 acres in Mexico (Hudson
1950).
To clarify the map showing the commercial alfalfa
producing regions of Mexico (Figure 18), the major areas
are located as follows: (1.) area to the north and east
of Mexico City, situated in the Distrito federal and
state of Mexico; (2.) the southern portion of the state
of Tlaxcala, and throughout the northern and central
portions of the state of Puebla; (3.) immediate vicinity
of the city of Oaxaca, Oaxaca; (Ij..) "Aguas Negras" re­
gion near Actopan, Hidalgo; (5.) "Hi Bajio" region in
the states of Guanajuato and Queretaro; (6.) "La Laguna"
area centered around Torreon, Coahuila; (7») Yaqul valley
area in southern Sonora; and (8.) areas around Zamora
and Moralia, Michoacan.
Table IX
A list of the collection data obtained for E» aurlfer from the U.S. National
Museum, British Museum, and the Museum of La Fundacion Rockefeller*
LOCALITY
DATE
COLLECTOR
Progress©, Mor.
V 27 A 8
A. C. Smith
June
H. H. Smith
Puente de Ixtla, Mor.
Cuernavaca, Mor*
Salle
La Fundacion Rockefeller Museum
Chapingo, Mexico
British Museum (Natural History)
London, England
C. C. Dean Collection, U.S.N.M.
Washington, D.C.
British Museum (Natural History)
Hoege
Wickham Collection, U.S.N.M.
7/7/1900
Cuernavaca, Mor*
COLLECTION
Cuernavaca, Mor.
---
Cuernavaca, Mor.
June
Mexico, Mexico
June, 1922 E. G. Smith
H. H. Smith
British Museum (Natural History)
U. S. N. M.
Esperanza, Mexico
---
Hoege
U. S. N. M.
Esperanza, Mexico
---
Hoege
British Museum (Natural History)
---
U. S. N. M.
Frugus, Mexico
Chapingo, Mexico
9/15A8
Chapingo, Mexico
---
A. C. Smith
La Fundacion Rockefeller Museum
F. Pacheco
La Fundacion Rockefeller Museum
Table IX- Continued
LOCALITY
DATE
COLLECTOR
COLLECTION
Pueblo
U. S. N«, M.
Distrito Federal
J. R. Inda Collection, U.S.N.M.
L« Cenradt
U. S. N.. M.
Wickham
British Museum (Natural History)
H. H. Smith
British Museum (Natural History)
Tepetlapa, Gro«
H. H. Smith
British Museum (Natural History)
Izuear de Matamoros, Pue. — -
Hoege
British Museum (Natural History)
Distrito Federal
mm ~ m a m e»
San Angel, D.F.
Omilteme, Gro*
July
Mexico
-----
Salle
British Museum (Natural History)
Mexico, Puebla
-----
Salle
British Museum (Natural History)
Mexico, Etla
---- -
Salle
British Museum (Natural History)
Salle
British Museum (Natural History)
Salle
British Museum (Natural History)
Salle
British Museum (Natural History)
Mexico, Izuear
Mexico, Orizaba
—
---
Mexico, Oaxaca
Mexico. Sierre de Duran«o — -
British Museum (Natural History)
Table X
A list of the collection data obtained for E* aurlfer by the author
in Central Mexico during 1952 and 1953*
Locality
Approx* Alt*
in Meters
Date of
Collection
Host
(Feeding)
La Patera, D* F*
221+0
May 29, 1952
Alfalfa
Mexico, D. F*
221+0
June 10, 1952
Alfalfa
Puebla, Pue*
2100
July 30, 1953
Alfalfa
Tecamachalco, Pue*
2000
July 30, 1953
Alfalfa
Azucar de Matamoros, Pue*
1550
July 2, 1953
Chaplngo, Max*
2200
July 27, 1953
Soybeans
Texcoco, Mex.
2200
July 29, 1953
Alfalfa
Chaleo, Mex*
2200
June 22, 1953
Alfalfa
Amecameca, Mex*
2250
Sept. 21, 1953
Alfalfa
Toluca, Mex*~ Rancho S* Antonio
2675
June 15, 1953
Alfalfa
Toluca, Mex*« Rancho El Carmen
2675
June 15# 1953
Potatoes
Table X- Continued
A list of the collection data obtained fop B. aurlfer
by the author In Central Mexico during 1952 and 1953*
Locality
Approx, Alt,
(in Meters)
Date of
Collection
Host
(Feeding)
Tlaxcala, Tla.
2100
July 3, 1953
Alfalfa
Actopan, Hgo,
20 I4.O
Sept. 22, 1953
Alfalfa
Ixmilqullpan, Hgo*
2000
Aug. 5, 1953
Alfalfa
Cuernavaca, Mor,
1600
June 13, 1953
Amaranthus sp
Cuernavaca, Mor,- Km, 89
1500
July 9 , 1952
Alfalfa
Zamora, Mich*
1500
Aug. 12, 1952
Corn
Moralia, Mich*
2150
Aug* 25, 1952
Alfalfa
Figure 17*- Map of the Republic of Mexico showing
the distribution of E* aurlfer*
I
vn
EPICAERUS
AURIFER
(BOH-)
. . . -t
• Areas Visited No Insects Found
a
Areas Visited Insects Collected
£ Serious Infestation On Commercial Scale
76
Figure 18♦- Map of the Republic of Mexico showing the
commercial alfalfa producing areas*
EPICAERUS
AURIFER
(BOH.)
78
Control
Cultural
A general review of the life history and habits
of E. aurlfer indicates several apparent approaches toward
control, by cultural means, of this pest.
The long incubation period of the egg required for
hatching, and the exposed position of the egg masses
would suggest several cultural practices designed to
destroy the eggs, such as heavy grazing or cutting at the
time the egg masses are most abundant. Also, the gregar­
ious habits of the adults, their slow and awkward means
of locomotion, and their inability to fly might also be
used to advantage in restricting the movements of E.
aurlfer.
Chemical
Laboratory
A series of laboratory tests were conducted,
prior to the field tests, to determine the general rel­
ative toxlcitles of selected contact insecticides against
the adult beetles. These tests, following the techniques
used by Morrison (l9l|5) and Stringer (l9ij.9), Tor screening
contact insecticides, consisted of saturating filter papers
with known quantities and concentrations of Insecticides
79
and exposing the insects to the treated papers for a given
period of time.
The laboratory tests were conducted at Chapingo
during June, 1952. Female beetles, collected from the
biological study area at La Patera were used in the tests.
The following materials and concentrations were used in
the tests:
1.- 0.5 per cent parathion
2.- 1.5 p©r cent E.P.N.
3.- 1.5 per cent DDT.
1|..- 2.5 per cent Dl/T.
5.- 0.5 per cent BHC.
6.- 1.5 per cent BHC,
7«- 1.5 per cent toxaphene
8.- 1.5 per cent chlordane
9.- 1.5 per cent dieldrin
10.- 1.5 per cent aldrin
11.- 1*5 per cent methoxychlor
The tests were replicated 8 times and each replicate
consisted of 10 beetles. The treated filter papers (15
centimeters in diameter) were allowed to dry for 15 hours
prior to the test. Ten beetles were introduced under
dish (12.5 centimeters
in diameter) inverted over
a petri
atreat­
ed filter paper. The beetles were allowed to remain in
contact #ith the treated papers for 1 hour, after which
80
they were removed and clean and untreated papers put in
their place. Observations and mortalities were recorded
at intervals of 1, 6, 2ip, and lj.8 hours. An untreated check
was Included in each test.
The tests, as summarized in Table XI, were not
statistically analysed as they were preliminary in nature
and designed only to indicate which materials might
warrant further testing in the field.
Toxaphene, BHC, aldrin, and chlordane appeared prom­
ising, under the conditions of the laboratory tests, and
for that reason, were included in later field tests.
Field
Two general methods of application of the in­
secticides were employed in the field tests. The toxicant
was mixed in the seedbed in which alfalfa was to be sown,
or it was applied to the foliage of the plant when the
adult beetles were present. The soil applications were
designed to control the larvae; the foliage application
was designed to control the adult beetles, thereby prevent­
ing oviposition and a subsequent infestation.
Application of Insecticides to the Foliage
This experiment involved the use of a series of
plots arranged in the form of a randomized block, with each
treatment being replicated
times. The experiment was
81
conducted at La Patera on a 2-year-old field of alfalfa,
of the variety Oaxaca, adjacent to the biological study
area. The roots of the alfalfa In this field had been
previously found to be uninfested; however, many beetles
had migrated into the field from surrounding heavily
infested fields. Each plot was 25 feet by 50 feet in size.
The following materials and concentrations were used
in the test*
1.- 5*0 per cent DDT.
2.- 20.0 per cent toxaphene
3«-
P©r cent BHC
I)..- 2.5 per cent aldrin
5»- 10.0 per cent chlordane
The test materials were applied as dusts with a Hudson
rotary hand duster June 26, 1952. The results of the treat­
ments, expressed as live beetles per square meter, are
summarized in Table XII. These counts were taken iq.8 hours
following the application of the materials.
All of the insecticides resulted in a consistent
high control (90 per cent or better reduction compared
with the check) of the adult beetles.
Applications of Insecticides to the Soil
Small Plot Experiment -- - This experiment was
82
designed in the form of a randomized block, each treatment
being replicated ij. times. The plots were each 10 meters
by 10 meters in size. The field in which this experiment
was conducted was prepared as a seedbed for alfalfa. The
insecticide treatments were applied June 10, 1952, to the
surface of the soil, as dusts, and raked into the top
2 to
3 inches of soil. An effort was made to sprinkle the
materials evenly
over the soil in each of the plots,and
to vary the direction of raking so that the materials
would be more uniformly distributed over the plots. Imme­
diately following the application of the materials, the
field was sown In alfalfa of the Oaxaca variety.
The following materials and concentrations were used
In the test:
1.- Aidrin,
2.0 pounds actual per acre rate
2.- Aidrin,
5*0 pounds actual per acre rate
bieldrin, 2.0 pounds actual per acre rate
Ip.- Dieldrin, 5*0 pounds actual per acre rate
5. BHC, 1.0 pound of the gamma isomer per acre rate
6.- BHC, 3*0 pounds of the gamma isomer per acre rate
7«- Toxaphene, 10.0 pounds actual per acre rate
8.- DDT., 10.0 pounds actual per acre rate
9*- Chlordane, 8.0 pounds actual per acre rate
The effects of the materials In reducing the injury
of the larvae to the primary roots was determined October
83
10, 1953, by examining the roots of 100 plants from each
plot. These figures are summarized in Table XIV. The
degree of infestation in the plots, including the untreated
check, was extremely low 10 to if. per cent injury) and as
a result, no evaluation of the treatments could be made.
Unreplicated Large Plot Experiment
This test was designed to determine the effect­
iveness of three materials: aidrin, dieldrin, and BHC, at
rates of 9 pounds, 9 pounds, and 3 pounds of the gamma
isomer per acre respectively, in controlling the larvae,
when applied on a field basis with standard farm equipment.
The plots were unreplicated and each was slightly
over 1 acre in size. To determine the effectiveness of
the treatments, the infestation counts were taken from
if. locations within each plot.
The materials were applied April 25, 1952, as dusts
on a newly-prepared seedbed, with a 12-foot "Evenflo"
distributor, and mixed into the top 2 to 6 inches of soil
with a tractor-drawn disc harrow. Alfalfa, of the variety
Oaxaca, was immediately sowed upon the treated soil.
The degree of injury, as summarized in Table XIII,
was determined by examination of the primary roots, 100
being taken from if. areas in each plot, October 10, 1953*
As in the small plot soil application test, the level of
31*
infestation was too low (2 to 9 per cent injury) to eval­
uate the materials.
No phytotoxic effects of the insecticides, in either
the foliage application test or the soil application tests,
were observed.
Table XI
Results of preliminary laboratory treatments, expressed as per cent control,
against the adult stage of Eh aurlfer. Chaplngo, Mexico, June 1952.
Treatment and
Concentration
1.
2.
3*
Replicates
U.
5.
0.5# parathion
10
10
0
20
1.5# E.P.N*
0
0
0
1.5# DDT.
10
0
2.5# DDT.
20
0.5# BHC
6.
7.
b'.______ Per cent Control
0
10
0
0
0
0
0
0
0
20
10
10
20
20
10
1 0 .0
10
30
20
10
ko
20
20
21.2
50
60
50
70
50
60
ko
90
5 8 .8
1.5# BHC
14-0
50
i^o
80
70
5o
100
90
6 5 .0
1.5# toxaphene
30
50
ko
20
50
30
30
ko
3 6 .25
1.5# chlordane
10
20
30
10
ij-o
30
10
10
2 0 .0
1.5# dieldrin
0
20
0
20
10
10
0
10
20
fco
10
20
20
30
20
0
2 0 .0
1.5# methoxychlor
0
0
0
0
0
0
0
0
0
Untreated check
0
0
0
0
0
0
0
0
1.5# aidrin
6.25
0
8 .7 5
rnmm**
Table XII
Results of foliar insecticide treatments for control of
E. aurlfer adults, La Patera, D.F., June 26, 1952#
Pounds
Live Beetles Per Square Meter
Treatment and
per
Replicates
Concentration_____ Acre_______ 1*
2.
3.
lu
5.
Per cent
Reduction
Mean____ Over Check
5$ DDT* dust
22
2
1
1
1
1
1*2
91
20$ toxaphene dust
2k
0
2
0
2
0
o.e
3$ BHC dust
20
0
1
0
0
2
0*6
2*5# aidrin dust
23
0
2
2
1
2
i*tv
9k
96
90
10$ chlordane dust
20
0
1
1
2
0
0.8
9k
Untreated check
—
13
10
13
19
13*8
--
Table XIII
Per cent of alfalfa primary roots injured by the larvae of E* aurifer
following soil application of insecticides, ]La Patera, June 10, 1952*
Treatment and
Amount per Acre
1.
2*0 pounds aidrin
3
1
2
5.0 pounds aidrin
2
0
3
1
1.5
2*0 pounds dieldrin
0
2
1
2
1.25
5.0 pounds dieldrin
0
1
1
1
0.75
1«0 pound BHC (g.i.)
1
2
1
0
1.0
3*0 pounds BHC (g.l«) 2
0
0
1
0.75
10 pounds toxaphene
3
2
1
2
2.0
10 pounds DDT*
k
0
3
1
2.0
8.0 pounds chlordane
0
1
0
2
0.75
Untreated check
2
3
3
3
2.75
Replicates
3•
Mean
....
2.5
Table XIV
Per cent of alfalfa primary roots injured by the larvae of
aurifer
following soil application of insecticides, La Patera, April 25* 1952*
Treatment and
Multiple Determinations
Amount per Acre_________1#
2»
3»
lt+_________ Mean
9*0 pounds aidrin
6
9
1
8
6*0
9*0 pounds dieldrin
7
8
k
1
5.C
3*0 pounds BHC (g*i,)
6
9
8
5
7.0
Untreated check
2
9
7
3
5.25
89
SUMMARY AND CONCLUSIONS
The biology, ecology, distribution, and control of
the weevil, Eplcaerus aurlfer Boheman, was studied in
south-central Mexico over a 2-year period. This insect was
found, for the first time, to be a local but serious pest
of alfalfa. The principal damage to alfalfa was noted to
be a result of the feeding of the larvae on the primary
root. Injury by this means results in the complete destruct­
ion of the central portion of the root, and eventually,
the death of the injured plant. The adults have also been
noted in the field feeding upon the following crops: co m ,
beans, wheat, and soybeans, and upon the following wild
plants: Solanum rostratum L., Anoda hastata Cav., Eruca
satlva Lam., Rumex obtusifollus L., Malva parvlflora L.,
Paspalum sp., Abutilon sp.. Cucumls sp., and Amarantus sp.
Under caged conditions in a food preference test the adults
were found to eat, in descending order of preference, the
foliage of beans, corn, alfalfa, tomatoes, wheat, and pota­
toes.
This insect has one complete generation a year. The
adults appear in late May or early June, and within 2
weeks following emergence, the adult population is at its
peak. At this time, the sex ratio is approximately equal.
90
At the time the adults are last seen in the field, the
latter part of September, there are approximately three
times as many females as males.
Copulation commences 2 to Ij. days after emergence
and eggs are produced 3 to 7 days later. The eggs, de­
posited in the form of a linear or ovoid mass, on the
underside of the leaf, within the trough formed by a folded
leaf or leaflet, or between two leaves, contain an average
of 26 eggs, fertilization is required for the production
of viable eggs and a fertilized female produces an average
of 2.17 egg masses per season. In the field, egg masses
have been collected from alfalfa, corn, beans, and a member
of the pigweed group, Amarantus sp. Of these plants, the
larvae are apparently only capable of developing on alfalfa.
The incubation period of the egg is approximately 30 days.
Upon hatching, the larvae immediately make their way into
the soil.
The number of larval instars was found to be l\.;
the winter being passed in the 3rd or i^th instar, or
occasionally as a newly formed pupa.
The pupal stage, lasting from 3 to ij. weeks, usually
occurs during the month of May. The pupa is found In an
ovoid earthen cavity at a depth of \ to
surface of the ground.
inches below the
91
Natural enemies were found to be Important in reducing
the field population of E. aurifer. Laboratory and field
observations indicate that 2 species of skunks, Spllogale
sp. and Maphltis sp.. are important predators of this pest.
Under caged laboratory conditions, a ground squirrel,
Cltellus mexlcanus mexlcanus and a shrew, Sorex saussurel
saussurel Merriam were also observed as predators. A
mite, identified as belonging to the family Parasitidae,
was observed as an external, apparently non-lethal parasite
of E. aurifer.
The distribution of this weevil, as determined by
field collection and summarization of museum data, was
found to be limited to the south-central portion of Mexico.
From collections on alfalfa, it was found to be present in
the valleys of Mexico, Mesquital, Tehuacan, Morelos, and
Morelia.
Preliminary control studies utilizing organic insect­
icides were conducted in the laboratory and field. Labora­
tory tests against the adult beetles indicated that BHC,
Toxaphene, DDT., chlordane, and aidrin might be worthy of
further field study. Foliar applications of these promising
materials, as dusts on alfalfa, against the adult beetles,
resulted In a uniform, high (90 per cent or better) degree
of reduction over the check.
Both small plot and large plot field tests, employing
92
the mixing of the toxicant, formulated as dusts, in the
seedbed prior to the sowing of alfalfa, were conducted.
Evaluation of these plots was not possible, however, because
of the low level of larval injury to the roots.
93
APPENDIX
Figure 1*- Climograph for Texcoco, Mexico*
Figure 2*- Climographs for Toluca, Mexico and Mexico D.F.
Figure 3*- Climographs for Pueblo and Tehuacan, Puebla*
Figure Ij.*- Climographfl for Cuernavaca and Cautla, Morelos*
Figure 5*- Climographs for Zamora and Morelia, Micho&can.
Figure 6*- Climographs for Actopan and Pachuca, Hidalgo*
9k
CLIMOGRAPH FOR
TfiXCOCO M e i . Mean Annual Precipitation 768 mm.
Mean Annual T em p e ra tu re 16 I*.C____
A L T IT U D E -------------------------- 224 0 Mta
IAN.
< a. a:
J ULY.
Appendix
Figure 1
95
-
I M I J . . H M '. '
Me
M io n
Al T I I
* ■>»
T u l U C fl
M il. —
C L IM O G R A P H
78 ^ IB IB
^ e 'n p p tjtu r* 187 £ --------k v L --------------------------8 8 7 f t U t l
FOR
C d .M « « lc a D F
(T a e u h a y a )
M « o n A n n u o l P r • c i p i f o t t o n 7 B S m m.
M to n Annual T e m p eratu re —1 4 ^ Zfi_____
A L T I T U D E _______________ 2 308 MU
>• n n u a i P r e c i p i t o h o n
j(AN
JAN.
JU L Y .
JU L Y .
Appendix
Figure 2
96
C lim o g r a p h f o r
PutMt PuiM «on A nnuol P r t c ip tlo lio n OQt m m
M to n Annuo I
16.2* C____
A L T I T U D E _______________ EI9Q l i l i i
C LIM O G R AP H FO R
T A h u flg a w
Pum
M ean A n n u a l P re c ip ita tio n 47M mm.
M oan A nn u al T a m p ira tu ra lA i
C
A L T IT U O E
________________ 1 1 2 1 M i l
(AN
|AN
T
JU L Y .
JU LY.
I
Appendix
Figure 3
97
C L IM O i.H A f'H r o R
C u n u tla
Mor.
CLIM O GRAPH FOR
M»on A nnuol P re c ip ita tio n 843 m m.
it n n A • i l t ’ ^nperotufe 21 9 *C
C u ernflW flC fl M n r
M to n A n n u ol P re c ip ita tio n 1Q4Q mm.
M ean Annua I Tem p eratu re 2Q.3*C_____
A L T IT U D E _______________ 1538 MU.
A.■*.!DDE ____
jiAfiL
JAW
JU LY.
JU L Y .
A pp e n d ix
Figure ij.
98
C U W -.G R A PH FOR
Z tfflg fQ
MjCtl-
CLIM O GRAPH FOR
M O f l i f l M ic h .
M acn A nn u ol Proc ip ito tio n 826 y m __
Mean A nnual T em pgrotuf e .. 2.Q-Q f i—
M oon A nn u ol P ro c ip lto tto n 8 3 3 mm,
M oan A nnuol T « m p » n itu n I 7 2 * c
ALTiiuDE
A L T IT U D E
_________________ 1564 M U
lAN
________________ 1823 Mlfc
JAN.
/
JU LY.
JU LY.
Apperxl ix
Figure 5
99
CLIM O G R APH FOR
PflfihllM Hflflt
A e ta n a n H an.
C LIM O G R AP H FOR
M eon A nn u ol P re c ip ita tio n a ib y m —
M oon A nnual T e m p eratu re 152 ..C____
A L T IT U D E
24Q Q MH.
M oon A nn u ol P ro e lp ito lio n
M oon A nn u al Tem per of ure_JU6JLjfi____
altitude _____________
191*
Htl
JAN.
JAN .
JU LY.
J ULY.
Appendix
Figure 6
100
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797•
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Sharp, David* 1891* Biologia Cantrail-Americana. Coleoptera. Curculionldae, Vol* IV, Part 3 (1889"
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tests of some organic compounds against Calandra
granarla L* Ann* Appl. Biol. 36: 213*
Transeau, E* N* 1935. The prairie peninsula* Ecology 16(3):
lj.23-ij.37.
Wellhausen, E* J. et al* 1951* Razas de malz en Mexico,
su origen, caracterlstlcas y distrihuclon.
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102
AUTOBIOGRAPHY
I, George Mallory Boush, was born in Norfolk, Virginia,
June 5, 1926. I received my secondary school education in
the public schools system of Princess Anne County, Virginia.
During World War II, I enlisted in the U. S. Army Air
Corps, and served as Aviation Cadet from June ll+, 1 9^1 to
November 8, 191+5* My undergraduate training was obtained
at Virginia Polytechnic Institute, Blacksburg, Virginia,
from which I received the degree Bachelor of Science in
191+8* From The Ohio State University, I received the degree
Master of Science in 1951* While in residence at The Ohio
State University, I held the position of Research Fellow
with the U. S. Public Health Service from October 1, 1950
to March l5» 1952. From April 1, 19U9 to September 30, 1950
I was employed as Assistant Entomologist at the Tidewater
Field Station, Virginia Agricultural Experiment Station at
Holland, Virginia; and from April 1, 1952 to December 1,
1953 was employed as Assistant Entomologist with the
\
Mexican Agricultural Program of the Rockefeller Foundation
in Mexico City, Mexico, since January 1, 1951+ I have held
the position of Assistant Entomologist at The Kentucky
Agricultural Experiment Station, Lexington, Kentucky.