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Group Title: Bilaterally gynandromorphic Hypodynerus, and a summary of cytologic origins of such mosaic Hymenoptera...(FLMNH Bulletin v.5, no.2)
Title: A bilaterally gynandromorphic Hypodynerus
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Permanent Link: http://ufdc.ufl.edu/UF00001558/00001
 Material Information
Title: A bilaterally gynandromorphic Hypodynerus and a summary of cytologic origins of such mosaic Hymenoptera
Series Title: Bulletin of the Florida State Museum
Physical Description: 26-40 p. : illus. ; 23 cm.
Language: English
Creator: Cooper, Kenneth W ( Kenneth Willard ), 1912-
Publisher: University of Florida
Place of Publication: Gainesville
Publication Date: 1959
Subject: Hypodynerus tuberculiventris   ( lcsh )
Gynandromorphism   ( lcsh )
Hymenoptera   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Bibliography: p. 37-40.
General Note: Cover title.
Statement of Responsibility: Biology of Eumenine wasps, VI.
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Bibliographic ID: UF00001558
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0866
notis - ACK0930
alephbibnum - 000440464
oclc - 05069498
lccn - a 60009050

Table of Contents
    Front Cover
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
    Back Cover
        Page 41
Full Text





Volume 5 Number 2


Kenneth W. Cooper


BIOLOGICAL SCIENCES, are published at irregular intervals. Volumes contain
about 300 pages and are not necessarily completed in any one calendar year.


All communications concerning purchase or exchange of the publication should
be addressed to the Curator of Biological Sciences, Florida State Museum, Seagle
Building, Gainesville, Florida. Manuscripts should be sent to the Editor of the
BULLETIN, Flint Hall, University of Florida, Gainesville, Florida.

Published.11 November 1959

Price for this issue $.35



SYNorsIS: A nearly perfectly divided, left ( )-right ( ) gynandromorph
of the Chilean eumenid wasp Hypodynerus tuberculiventris (Spin.) and a dissec-
tion of its terminal retracted abdominal segments and genitalia are described
and illustrated. A table summarizes the morphology of the seven or eight other
sexual mosaics recorded in the Eumeninae (Vespidae). Although female Hy-
podynerus average considerably larger than males, the approach to dimensional
symmetry of the male and female halves probably reflects nearly equal growth
per food unit (at not-extreme ranges of provisioning) of male and female wasps,
and not a dominant influence of the female half. Indeed, as is usual for such
mosaics although not invariably so, the male and female tissues of this wasp
appear to have differentiated autonomously. The rarity of collection of such
mosaics (frequency < 0.01%) grossly underestimates the frequency of the cytologic
mishaps giving rise to them. The known errors of oocytic meiosis, of suppres-
sion of nuclear activity, and of fertilization leading to gynandromorphy are dis-
cussed; in aggregate their raw frequencies for Hymenoptera probably lie between
0.1% and 1.0% of all eggs.

Although frequent and even mass occurrences of gynandromorphs
and minor sexual mosaics are known in certain exceptional strains
of honeybees (e.g., von Siebold 1864; Rothenbuhler 1958) and colonies
of ants (e.g., Wheeler 1937; Donisthorpe 1946), gynandromorphic
Hlymenoptera are nevertheless extremely rarely encountered by col-
lectors. Thus, to judge from the compilations and observations by
Dalla Torre and Friese (1899), Wheeler (1903), Enderlein (1913),
Mitchell (1929), Krombein (1949), and others, as well as from citations
in Zoological Record, gynandromorphs have been described in fewer
than 200 species of Hymenoptera. If some 280,000 Hymenoptera
have now been described as Bernard (1951) estimates, then less than
a tenth of a percent have gynandromorphic representation, and a
rough estimate of the overall frequency of occurrence of gynandro-
morphs in collections would perhaps be an order of magnitude less
than this, or roughly one-hundredth of a percent. As will be brought
out in the discussion, which reviews what is now known of the origins
of hymenopterous gynandromorphs, the raw frequencies of the events
leading to gynandromorphy (and other mosaicism) are probably con-

Graduate Research Professor of Biology, University of Florida, Gainesville,
Florida. New address: Department of Cytology, Dartmouth Medical School,
Hanover, New Hampshire. Manuscript received 30 August 1959.-ED.


siderably higher than would be judged from the experience of col-
The gynandromorphic Hymenoptera now known are more or
less randomly scattered among sawflies, parasitic Hymenoptera (tere-
brants), ants, wasps and bees, and they probably occur in most species
of Hymenoptera, if not in all. They are conveniently classified to
the four main types recognized by Dalla Torre and Friese (1899):
lateral, the two sides of the insect being of different sex; frontal, hav-
ing a fore-aft division of the sexes; transverse, being dorso-ventrally
divided; and mixed, or combinations of varying complexity of the
first three types. Members of the four classes are not equally fre-
quent, but form a series: lateral > mixed, mixed frontal, and
mixed or frontal > transverse. The male or female parts of gynan-
dromorphs seem to show no tendency to be predominantly left, an-
terior, or dorsal. As might be expected, most gynandromorphs are
only partially so, for they do not have the body equally divided into
male and female sectors. Rather only a portion of the insect (e.g., the
head, the thorax, the head and thorax, etc.) is gynandromorphic in
most cases, and total involvement of the body occurs in less than
10% to 15% of all. The small number of previously known eumenid
gynandromorphs (summarized in table 1) conform with these gen-
eralizations, and the very remarkable new example now to be de-
scribed is the second known instance of a total, lateral, gynandromor-
phic eumenid wasp (figs. 1, 2). Of special interest is the dissection
of the terminal abdominal segments of this gynandromorph (fig. 2),
for the only other eumenid gynandromorph having divided terminalia
was not dissected (see Hoplopus reniformis (GCmnel.) in table 1, and
Leininger 1926).

Hypodynerus tuberculiventris (Spinola)
(Figures 1, 2)
Eumenes tuberculiventris Spinola, 1851, in: Gay, Historia Fis. Pol.
Chile, Zool., VI, p. 267.
Odynerus (Leionotus) tuberculiventris de Saussure, 1853, Et. Fam.
Vesp., I, p. 162
Odynerus (Hypodynerus) tuberculiventris de Saussure, 1855, Et. Fam.
Vesp., III, p. 227
Nortonia tuberculiventris Zavatteri, 1912, Arch. f. Naturgesch.,
LXXVIII, Abt. A, Heft 4, s. 169, Pl. II, fig. 49
Hypodynerus tuberculiventris Bequaert and Ruiz, 1943, Rev. Chilena
Hist. Nat., 54, pp. 30-32, (1941).


LABEL DATA: El Canelo, Prov. Santiago, XI-XII, 1952, Chile. Col-
lected by Louis Pefia (together with some 220 other normal individ-
uals of this species, of which no more than 5% were males).
HABITUS (fig. 1): The left side is entirely of male structure and col-
oration, whereas the right side is conversely entirely female. For the
most part the wasp is well proportioned and nearly symmetrical in
overall appearance, measuring ca. 2.5 mm in head width (between
the outer margins of the compound eyes) and ca. 7.7 mm from frons
to the apex of abdominal tergite-2 (foreshortened in fig. 1). The body
size is accordingly that of an average female, being considerably
larger than an average male. The asymmetry therefore reflects not
the normal sexual dimorphism for body size, but rather the propor-
tional differences between male and female structural parts (e.g., the
compound eyes, mandibles, etc.; see figure 1) normal to wasps of
opposite sex but of corresponding body sizes.
The overall effect of the color pattern, like that of the morphology,
is one of general symmetry. The body of the wasp is predominantly
shining black with bright yellow maculations. Except as noted be-
low, the antennae and legs (from just before the apex of the femur
to the infuscated last tarsal joint) are light reddish brown or rusty.
Without magnification the color pattern is disrupted by only two
marked asymmetries, namely the striking bipartite black and yellow
pattern of the clypeus and the unique yellow spot on the left mesocoxa
(fig. 1). Despite the overall appearance of symmetry, the color pat-
tern of the left side follows faithfully that of the average male and
when magnified it markedly contrasts with that of the female right
side wherever male and female patterns of normal wasps differ.
HEAD (fig. 1): Overall asymmetry slight; nevertheless when
studied in face view or from below (fig. 1), or from above, individual
structures of the left side-being of typical male morphology and
color pattern-present striking contrasts to their counterparts in the
right half which are of normal female constitution and proportions.
For example, on the left side: the ocellar triangle is skewed; the
compound eye is notably larger than that on the right, and of different
shape; the antenna is longer than that on the right, its joints are of
different proportions, and it has 13 segments (12 in Y), the terminal
one being typically hooked as in all males; the frontal carina is more
narrowly separated from the antenna socket; the clypcus is less wide
but proportionately longer; the gena and mandible are markedly
smaller; and so on.


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Fig. 1. Bilateral gynandromorph of Ilypodynerus tuberculiventris (Spin.);
left side of wasp male, right side female.
The body of the gynandromorph is shown in ventral view. The normally
retracted abdominal segments, the genitalia, and the male tergite-VIII and sternite-
IX have been dissected from it (sec fig. 2 of the dissection). Above, head in full
face view, maximum width ca. 2.5 mm.; below, stemite-VII in ventral aspect.
Sec text for full description.

Vol. 5





Fig. 2. Cleared dissection of terminal, retracted abdominal segments and
genitalia of the bilateral gynandromorphic Hypodynerus tuberculiventris (Spin.).
Male parts to left, their labelling in lower case; female parts to right, labelling in
The dissection is in dorsal aspect, and the female parts are displayed in their
normal relations and in dorso-lateral aspect. Male sternites-VIII and -IX have
broken apart, the spiculum is detached, and these elements are viewed from
their internal surfaces. Phallus and gonobase have rotated 180" and are viewed
from their ventral surfaces. The sclerotized apodemes of st-viii, sc ?, T-VIII,
T-IX, VF-1, VF-2 are stippled. Vertical scale to left equals one millimeter.
g, gonobase or basal ring of phallus; gst, gonostipes or basiparamere; la, lamina
aedeaglis, "lateral rod", or "penis valve" of aedeagus; ps, parameral spine; s,
spiculum or median apodeme of sternite-IX (detached); sc ?, scalar remnant of
tergite-IX ?; sp, spiracle of tergite-VIII; st-viii, sternite-VIII broken from ankylo-
sis with sternite-IX; st-ix, sternite-IX; t-viii, tergite-VIII; vol, volsella.
DB, dorsal bridge of hemitergite-VIII; R, ramus of second valvula; SP,
spiracle of hemitergite-VIII; T-VIII, hemitergite-VIII; T-IX, hemitergite-IX; VA-
I, -2, -3, valvulae-1, -2, and -3 respectively; VF-1, -2, valvifers-1 and -2.


The typical and hence contrasting male morphology of the left
side of the head is in turn accentuated by the color pattern differences
between the sexes. Thus, on the left side: the yellow frontal spot is
smaller; the scape and pedicel of the antenna are black above, light
reddish brown behind, and yellow in front and below as in a typical
male (on the right, as normal for the female, the scape is entirely light
reddish brown, and the pedicel is black above and light reddish
brown elsewhere); the clypeus is yellow, contrasting sharply with
the shining black of the right or female side.
THORAX AND APPENDAGES (fig. 1): The left pronotum from above
is very slightly shorter at its outer apical angle, and the left meso-
thorax and left propodeum are slightly less rounded and robust than
on the right (female) side. The wings on the left are slightly shorter
than on the right (foreshortened and not evident in fig. 1). The legs
of the left side are very slightly more slender than on the right, and
the left middle femur is slightly sinuate below.
The yellow ornaments on the two sides of the thorax show few
differences, except that the tegulum on the left is marked basally
with black overlying the rufous medial field and only the posterior
tip is yellow, whereas the posterior third of the tegulum on the right
is yellow and has only a suggestion of black basally. The divided
yellow dot on the left anterior distal half of the left mid femur, the
yellow splash on the outer distal face of the left anterior tibia and
on the anterior distal half of the left mid tibia, and the small yellow
spot on the outer mid sector of the left hind tibia are all male char-
acteristics, and have no counterparts on the right (or female) side.
The orange coloration of the costal field of the right forewing is some-
what brighter than that of the shorter, left (male) wing.
ABDOMEN (segments II-VII; fig. 1)2: As in the cases of the thorax
and propodeum, the left side of the abdomen is slightly less robust
than the right, but this is not obvious without close study. Likewise
the yellow color pattern shows no certain difference on the two sides;
such slight asymmetries as exist are paralleled in normal individuals
of either sex. However, the left side of the undissected specimen
had seven segments as in a normal male, whereas but six occurred

SThe propodeum, or true first abdominal segment, has been dealt with in the
discussion of the thorax; the apparent first abdominal segment, bearing the petiole,
is in fact the second abdominal segment, and the numbering employed here
accordingly counts the propodeum as abdominal tergite-1.


on the right side as is typical of most female aculeates. Although
tergite-VII on the whole appears symmetrical, it has a small notch
apically, slightly left of center at the margin of juncture of male and
female sides. To the right of center, the female tergital half extends
slightly more distally than does that of the male. Sternite-VII, on the
other hand, is very asymmetrical (fig. 1, below), being unequally scal-
loped medially and somewhat prolonged on the left (or male) side.
Sternite-IX (apparent seventh male sternite) projected somewhat to
the right beyond the right apex of the emargination in the undissected
specimen, and was overlain by what in effect is a male hemitergite-
VIII (fig. 2, t VIII). Aside from the anomalous occurrence of an
extra abdominal segment on the left side, before dissection the apex
of the abdomen possessed a deceptively regular contour.3
ABDOMEN (segments VIII-IX and genitalia; fig. 2): Only in the
head of this wasp is the sexual dimorphism and clearly bilateral di-
vision into a left male half and right female half as strikingly and
diagrammatically displayed as in these terminalia. The left eighth
tergite of the male half (t viii) is sclerotized and pigmented over its
exposed apical two-thirds. Proximally, as in a normal male, this ter-
gite is neither pigmented nor strongly sclerotized. But unlike that
of the normal male it is no more than a hemitergite, and it is joined
across the midline by a dorsal bridge (DB), just as a left hemitergite
of a normal female would be, with the wholly retracted, structurally
normal, female hemitergite of the right side (T VIII). Aside from
the dorsal bridge and integumentary membranes connecting the main
exoskeletal plates, there are no other unions between the left (male)
and right (female) halves.
A partially successful attempt was made to keep all the exoskeletal
parts in their normal array and relations during dissection and per-
manent mountings, yet regrettably the left (male) sternite-VIIT (st
viii), which is normally ankylosed to sternite-lX, and the spiculum
(s) were broken from their unions. Nevertheless the arrangement and
structures of sternites-VITI and -IX are clear enough and, except

Surely, in the field, this wasp would have appeared to most collectors quite
normal and a female, just as it did to Pefia. Yet there may be some sufficiently
alert, discerning, and nimble to be certain death to such a diminutive freak.
Consider Morice's (1901) comment, when writing of an equally cryptic gynandro-
morphic bee, Podalirius retusus (L.): "Its curious particolored face caught my
eye, otherwise I should have let it pass as a female, for the other characters were
too inconspicuous to attract attention as it flew past me." (!)


along their medial edges, of normal morphology. The spiculum, on
the other hand, is unduly short and disproportionately broad. It is
not certain whether this is an artifact resulting from damage in dis-
section, or whether the spiculum was in fact so modified in the
gynandromorph. It may well be modified for not only is it a ventral
midline structure, but to a noticeable degree the phallus is also ill-
shapened (as are also the first and second valvulae on the female side),
for it is both broader and somewhat shorter than normal, and has a
shortened, less needle-like, parameral spine (ps). Nevertheless exactly
half-the left half-of a phallus is present, containing each of the
lateral components of the normal eumenid phallus (see figure 2). A
flap of integument (sc ?), the left half of which is elliptically curved
as ordinarily is the case for eumenine scalar remnants of tergite-IX,
and bears a weak apodeme along its left apical margin, completes
the retracted abdominal and genital complements of a normal male.
The female morphology of the right side (fig. 2) is normal insofar
as hemitergites-VIII and -IX, valvifers-1 and -2, and valvula-3 are
concerned. Valvula-1, however, is shorter and somewhat more ro-
bust than usual, and valvula-2 is much shortened and deformed be-
yond its basal third. Nevertheless the female genitalia and retracted
abdominal segments show no suggestion of masculinizationn", and
in every aspect the right side is completely that of a female.
COMMENT: This specimen, then, is a notably clean-cut bilateral
gynandromorph, with the only obvious disturbance of symmetry be-
ing the invasion of the dorsal bridge of right hemitergite-VIII across
the midline onto the male side. The minor proportional defects in
the phallus, perhaps in the spiculum, and in valvulae-1 and -2, can-
not be taken as evidence of "feminization" or masculinizationn", and
perhaps best can be understood as reflecting restraints on the sym-
metrical growth of these very diverse parts brought about by their
apposition in the developing pupa. There is no suggestion that diffus-
ible elements derived from one sex have notably biased the develop-
ment of the other. Rather, as in the case of other gynandromorphs
(but not all, see Leclercq 1953) the evidence indicates that the tis-
sues of each sex seem largely self-determining. Nor is the general
symmetry and correspondence of gross body size of the male and
female halves with that of an average female suggestive of a dom-
inant effect of the female half on the male half, even though the aver-
age female Hypodynerus tuherculiventris (Spin.) is considerably larger
than the average male. Body size in eumenid wasps is, within limits,
a function of food consumed, and the largest males in a population


are larger than the smallest females. This wasp, as a larva, very
probably developed in a female cell with a provender of caterpillar
prey of the bulk normally provided an average zygote (or fertilized
egg, which, in Hymenoptera, ordinarily develops as a female) in
contrast to that stored for an azygote (or unfertilized egg developing
as a male). What is of interest, and which this wasp shows unequivo-
cally, is that male and female eumenid larvae that have consumed
an equal bulk of food (which is neither excessively large nor unusually
scant) will develop to wasps of equal size, or virtually so.

The low frequency with which hymenopterous gynandromorphs
occur in the experience of collectors (< 0.01%) would, were Boveri's
(1888, 1915) hypothesis of their formation valid, permit a rough esti-
mate of the lowest rate at which delayed sperm extraordinarily ferti-
lize one nuclear product of the first cleavage division of haploid eggs.
The result of such an abnormal sequence would be the entrance to
second cleavage of one diploid, or 2N, syngamic nucleus (ordinarily
giving female tissues) and one haploid, or N, matroclinous nucleus
(giving male tissues), the adult product thereby being a sexual mosaic.
The particular pattern of sex mosaicism, and the ultimate percentage
of the individual made up of male rather than female tissues, would
be principally determined by the partition of the diploid and haploid
nuclei of the embryo among the imaginal discs as in any case initially
involving two or more classes of nuclei. Under Bovcri's hypothesis
each gynandromorph commences life as an inseminated egg, and this
much is assuredly true for most gynandromorphs of Habrobracon and
Apis where detailed genetic analysis has been possible. However it
is now very clear, especially from Whiting's research on Habrobracon
as well as from Rothenbuhler's and others' on the honeybee, that the
sequence of events following fertilization that Boveri postulated
occur extremely rarely, if at all, and that gynandromorphs may arise
in a number of different ways.
The commonest modes of origin of gynandromorphs in Hlabrobra-
con and Apis involve events prior to cleavage in the egg. Thus the
commoner origin of gynandromorphs in Apis (but a much rarer one in
Habrobracon) is brought about by effective dispermy; one sperm fer-
tilizes the haploid egg nucleus, the other sperm, instead of remaining
arrested or degenerating, directly enters cleavage to give a clone of
haploid nuclei (Rothenbuhler, Gowen and Park 1952; Rothenbuhler
1958; Whiting 1928, 1931, 1943). In these cases the events are just


as postulated by Morgan (1905) and, contrary to Boveri's view, the
male tissues are of patroclinous or androgenetic origin. A simple
but uncommon variant is known in the honeybee in which haploid
nuclei derived from two sperm, in addition to the syngamic diploid
nucleus, undergo cleavage without fusion. The resulting gynandro-
morph is then in addition a mosaic of two genetically different, andro-
genetic male tissues (Rothenbuhler 1957, 1958). Turn about, in
Habrobracon the frequent mode of origin of gynandromorphs follows
the normal fertilization of an egg (providing a 2N nucleus ordinarily
developing to female tissues), but thereafter this zygotic nucleus and
one remaining nuclear product of oocytic meiosis (hence N, develop-
ing male tissues) enter cleavage (Whiting 1935, 1943; Whiting and
Wenstrup 1932; Whiting and Whiting 1927). The prime cytologic
abnormality in this case is the production of a binucleate egg, and
it can be shown that the two egg nuclei may be genetically different
and hence cannot, in such a case, be identical products of a first cleav-
age division as Boveri postulated (Whiting 1932; Whiting and Stancati
1931). Gynandromorphs arising in this way have matroclinous or
gynogenetic male parts. Though the commonest type in Habrobra-
con, gynandromorphs of such an origin are found only infrequently
in honeybees (Mackenson 1951; Rothenbuhler 1958).
It might be expected that very rarely a binucleate egg would be
effectively dispermic, with one sperm fertilizing one egg nucleus and
both the other egg nucleus and the other sperm nucleus entering
cleavage independently to give two genetically different clones of
haploid nuclei. The haploid male portions of gynandromorphs aris-
ing in this way would be in part of androgenetic (paternal) origin,
in part of gynogenetic (maternal) origin. Such has in fact been found
in Habrobracon (Whiting 1943). Finally, impaternate gynandro-
morphs arising from wholly unfertilized eggs have been proven to
occur rarely in Apis (Tucker 1958), and evidently are known also in
Habrobracon (see von Borstel 1957). In these cases either the second
meiotic division products of one of the two second meiotic spindles
fail to be segregated, or two of the four nuclear products of meiosis
fuse to form a diploid nucleus heterozygous for the sex alleles. In
Tucker's (1958) opinion, the haploid products of oocytic meiosis may
in certain cases undergo one cleavage division before fusing to give
diploid nuclei. In any case a diploid nucleus results, and this nucleus
and one of the two haploid meiotic products enter cleavage to give
a wholly gynogenetic embryo that is a mosaic for female-determined
(2N) and male-determined (N) nuclei. Thus hymenopterous gynandro-

Vol. 5


morphs may arise in a number of different ways, all of which involve
either oogenetic meiotic mishaps, failures to suppress the activity of
all but one product of these meioses, or failures to arrest the activity
of sperm not participating in syngamy, or combinations of these events.
There are, of course, still other conceivable patterns or combina-
tions, involving no more than these meiotic and physiologic failures,
by which gynandromorphs might arise. What is more, the occurrence
of biparental or other diploid males (and of diploid male tissues in
mosaics), and the fact that haploid-diploid sex determination may not
depend upon simple heterozygosity for sex alleles in all Hymenoptera
(e.g., in forms with very close inbreeding; see Whiting 1947, 1958),
further complicate any basis for estimation of the average frequency
of the raw events underlying gynandromorph production. Certainly,
however, they are far more frequent than the gynandromorphs them-
selves (perhaps occurring in 0.1% to 1.0% of all eggs). On the other
hand the very high rates for some special strains within species, such
as Rothenbuhler's (1958) honeybee stocks which may produce up to
40% gynandromorphs, do not reflect chance breakdowns in oogenetic
meiosis or in mechanisms of physiologic restraint. They have been
proven to be genetically determined oocytic or gametic dysfunctions.
Brief but interesting discussions of the possible nature of certain of
these dysfunctions have been given by von Borstel (1957) and Tucker

Benoit, R. ct L. Berland
1935. Trois cas de gynandromorphisme chez les hym6nopt6res aculkates.
Arch. Mus. ilst. Nat. Paris, ser. 6, t. 12, p. 435-458.
Bernard, F.
1951. Ordre des hymenopt&res. Hymenoptera Linne, 1758. Gen6ralit6s. in:
Trait6 dc Zoologie (6d: P. -P. Grass6), t. 10, premier fasc., p. 771-772,
Paris: Masson et Cie, 975 p.
Borstel, R. C. von
1957. Nucleocytoplasmic relations in early insect development, in: The Be-
ginnings of Embryonic Development (ed.: A. Tyler, R. C. von Borstel,
and C. B. Metz), pp. 175-199. Washington, D. C., 400 pp.
Boveri, Th.
1888. Ueber partielle Befruchtung. Sitzber. Gesell. Morph Physiol., Miin-
chen, Bd. 4, s. 64-72.
1915. Uiber die Entstehung der Eugsterschen Zwitterbienen. Arch. entw.-
mcch. Org., Bd. 41, s. 264-311.


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