i , BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 5 Number 2 A BILATERALLY GYNANDROMORPHIC HYPODYNERUS, AND A SUMMARY OF CYTOLOGIC ORIGINS OF SUCH MOSAIC HYMENOPTERA. BIOL- OGY OF EUMENINE WASPS, VI. Kenneth W. Cooper f of . 8 I . -- I * /853 ' UNIVERSITY OF FLORIDA Gainesville 1959 The numbers of THE BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES, are published at irregular intervals. Volumes contain about 800 pages and are not necessarily completed in any one calendar year. OLIVER L. AusTIN, JR., Editor 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. Publi5hed 11 November 1959 Price for this issue $.35 A BILATERALLY GYNANDROMORPHIC HYPODYNERUS, AND A SUMMARY OF CYTOLOGIC ORIGINS OF SUCH MOSAIC HYMENOPTERA. BIOLOGY OF EUMENINE WASPS, VI. KENNETH W. COOPER 1 SYNOPSIS: A nearly perfectly dividad, left (3 )-right (9) gyhandromorph of the Chilean eumenid wasp H!/Podynefus tubercutiventris (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 1987; D6histhorpe 1946), gynandromorphic Hymenoptera are nevertheless extremely rarely encountered by col- lectors. Thus, to judge from the compilations and observations by Dalia Torre and Friese (1899), Wheeler (1908), Enderlein (1918), Mitchell (1929), Ktombein (1949), and others, as well as.from citations - in Zoological Record, gynandromorphs have been described in fewer than 200 species 6f 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 ah order 6f 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 hymenoptdrous gynandromorphs, the raw frequencies of the events leading to gynandromorphy (and other mosaicism) are probably con- ' Graduate Research Professor of Biology, University Rf Florida, Gainesville, Florida. New address: Department of Cytology, Dartmouth Medical School, Hanover, New Hampshire. Manuscript received 86 August 1959.-ED. 26 BULLETIN FLORIDA STATE MUSEUM Vol. 5 siderably higher than would be judged from the experience of col- lectors. The gynandrom6rphic Hymenoptera now known are more or less randomly scattered among sawflies, parasitic Hymenoptera (tere- brants), ants, wasps and bees, and they probabiy occur in most species of Hymenoptera, if not in all. They are conveniently classified to the four main types recognized by Dalia 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; transuerse, 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 g>man- 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 gy-nandromorph having divided terminalia was not dissected (see Hoplopus reniformis (Gmel.) 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, ~t. Fam. Vesp., I, p. 162 Odynerus (Hypodynerus) tuberculiventris de Saussure, 1855, Et. Fam. Vesp., III, p. 227 Nortonia tuberculiuentris Zavatteri, 1912, Arch. f. Naturgesch., LXXVIII, Abt. A, Heft 4, s. 169, Pl. II, fig. 49 Hypodynerus tubercutiventris Bequaert and Ruiz, 1943, Rev. Chilena Hist. Nat., 54, pp. 80:82,(1941). 1959 COOPER: A BILATERALLY GYNANDROMORPH 27 LABEL DATA: El Canelo, Prov. Santiago, XI-XII, 1952, Child. Col- lected by Lodis 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 ahd col- oration. whereas the right side is conversely entirely female. Fof the most part the wasp i5 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- tem 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, j 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 18 segments (12 in 9 ), the terfpinal one being typically hooked as in all males; the frontal carina is more narrowly separated from the antennal socket; the clypeus is lesswide but proportionately longer; the gena and mandible are markedly smaller; and so on. TABLE 1 SUMMARY TABLE OF DESCRIBED ,GYNANDROMORPHIC EUMENID WASPS. TYPE F, F~ONTAL GYNANDROMORPHS; TYPE L, LATERAL GYNANDROMORPHS; TYPE M, MIXED GYNANDROMORP*IS; No TRANSVERSE GYNANDROMORPH HAS SO FAR BEEN REPORTED IN THE EUMENINAE,-SEE TEXT FOR DESCRIPTION OF GYNANDROMORPHIC TYPES. Species Type Head Thorax Abdomen, Reference Eumenes #avicornis Sauss. ?F both antennae S; 8 2 de Saussure 1856 all else 9 Odunerus acoelogaster Perk.1 M' left side S , right 9 left a ; right 9 except for 9 except for an incom- Perkins 1901, 1922 mid and hind legs .( a -like) plete 7th dorsal segment on right ( 0 -like); genita- lia 9 Ancistrocerus callosus (Thoms.7 M left side a , right 9 left 9, except foreleg and outwardly purely 9 ; gen- Renge11917 sterhum (8); right side 6, italia not examined except legs which are 9 Ancistrocerus callosus (Thoms.7 ? left antenna 9 in 9 9 Rengel 1917 structure except for presence of a 13th, terminal, hook-like segment ( 3), right ~ antenna b r.o ken; head otherwise 9 Parancistrocerus pedestris M head an irregular left legs and wings 9 , entirely 9 , including gen- Malloch 1924 (Sauss.) 2 mosaic of a and 9 right a italia characters Hoplopus reniformis.(Gmel.) + L left side 9, right 3 left side. 9, right S left side 9, right 8 ; geni- Leininger 1926 talia similarly divided, but may not be 50-50 for phal- lus, of abnormal shape Pterocheitus phaterata var. L left side 3, right 9 9 9, but apparent sternites- Benoist and Berland cheorierana Sauss. G II, -III, -IV and -V.are 6 1985 on the left side, 9 on the right; sting projecting, so genitalia probably 9 Ancistrocerus parietuni (L.) M left side 9, right 3 left half of scutellum 9 „ externally 9 Kontkanen ,1985 right half a ; all legs 9 ; upper right thorax 3 -like , J. F. Perkins of the British Museum of Natural History kindly informed.me that this specimen is in the museum's collection and is to be referred to Odynerus acbelogaster R. C. L. Perkins. (synonyms: O. relictus R. C. L. Perkins and O. lithophilus R. C, L. Perkins). ' Recorded as Odynerus callosus Thoms. 1 Recorded as Odynerus (Stenodynerus) conformis Sauss. ' Recorded as Odynefus reniformis Gmel, " Recorded as Pterochitus cheurieranus,Sauss. 30 BULLETIN FLORIDA STATE MUSEUM Vol. 5 0" , -'/- 355:~i* 'r' AF. ' 1 W 4 \r /, 7Vit, .4 4 -61/ Fig. 1. Bilateral gynandromorph of Hypod!,nerus tuberculiventris (Spin.); left side of wasb 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 (see fig. 2 of the dissection). Above, head in full face view, maximum width ca. 2.5 mm.; below, sternite-VII in ventral aspect. See text for full description. 1959 COOPER: A BILATERALLY GYNANDROMORPH 31 VA- 2ps gst VA- 1 vol '6: -36 VA-3 9 la V F-2 stix R st viii SC ? T IX V F- 1 SP sp T Vill It vw DB 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 capitals. 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 stippied. Vertical scale to left equals one millimeter. g, gonobase or basal ring of phallus; gst, gonostipes or basiparamere; la, lamina "xedeaglis, "lateral rod", or "penis valve of aedeagus; ps, parameral spine; s, spiculum or median apodeme of sternite-IX (detached); se ?, 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 5econd valvula; SP, spiracle of hemitergite-VIII; T-VIII, hemitergite-VIII; T-IX, hemitergite-IX; VA- 1, -2, -8, valvulae-1, -2, and -8-respectively; VF-1, -2, valvifers-I and -2. 32 BULLETIN FLORIDA STATE MUSEUM Vol. 5 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 pronetum 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 5horter than on the right (foreshortened and not evident in Bg. 1). The legs of the left side are very slightly more slender than (in 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 oveflying 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 fight 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 ' The propodeum, or true Erst 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. 1959 COOPER: A BILATERALLY GYNANDROMORPH 38 on the right side as is typical of most female aculeates. Although tefgite-VII on the whole appears symmetrical, it has h 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 seal - loped medially and somewhat prolonged on the left (of 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 'VIIl). Aside ffom the anomalous occurrence of an extra abdominal segment on the left side, before dissection the apex . of the abdomen possessed a deceptitely regular contour.3 ABDOMEN (segments VIII-IX and ganitalia; Bg. 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 scIerotized and pigmeated over its exposed apical tw6-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 n6 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-VIII (st viii), which is normally ankylosed to sternite-IX, and the spiculum (s) were broken from their unions. Nevertheless the arrangement and structures of sternites-VIII 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. Consid6r Morice's (1901) comment, when writing of an equally cryptic gynandro- morphic bee, Podalirius retusus (Ii.): "Its curious particoldred. face caught my eye, otherwise I should have let it pass as a female, ·for the other characters were toDinconspicuous to attract attention as it flew past me." (1) 84 BULLETIN FLORIDA STATE MUSEUM Vol. 5 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 masculinization, and in every aspect the right side is completely that of a female. COMMENT: Thls 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 "masculinization, 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 1958) 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 tuberculiventris (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 1959 COOPER: A BILATERALLY GYNANDROMORPH 85 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- tally, 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 ORIGINS OF HYMENOPTEROUS GYNANDROMORPHS The low frequency with which hymenopterous gynandromorphs occur in-the experience of collectors (< 0.01%) wbuld, 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, matribelinous 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 Boveri'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 Habrobra- con and Apis involve events prior to cleavage in the egg. Thus the commoner origin of gy-nandromorphs 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, 1981, 1948). In these cases the events are just 86 BULLETIN FLORIDA STATE MUSEUM Vol. 5 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 1985,1948; Whiting and Wenstrup 1982; 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 nuciei may be genetically different and hence cannot, in such a case, be identical products of a first cleav- age division as Boveri postulated (Whiting 1982; Whiting and Stancati 1981). 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. ohe 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 1948). Finally, impaternate gynandro- morphs arising from wholly unfertilized eggs have been proven to occur riirely in Apis (Tucker I958), and evidently are known also in Habrobracon ( see von Borstel 1957). In these cases either the second mei6tic division products of one of the two second meiotic spindles fail to be segregated, 6r 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 Pfoducts of oocytic meiosis may in certain cases undergo one cleavage division before fusing to give diploid nuclei. In any ease 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- 1959 COOPER: A BILATERALLY GYNANDROMORPH 87 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 the5e 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 cembina- 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 h6terozygosity 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, h6wever, they are far more frequent than the gynandromorphs them- selves (perhap5 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 dysfunetions have been given by von Borstel (1957) and Tucker (1958). LITERATURE CITED Benoit, R. et L. Berland 1985. Trois cas de gynandromorphisme chez les hymdnoptbres acul6ates. Arch. Mus. Hist. Nat. Paris, ser. 6, t. 12, p. 485-458. Bernard, F. 1951. Ordre des hym6nopt6res. Hymenoptera Linnd, 1758. Gen6ralit65. in: Traita de Zoologie (6d: P. -P. Grass6), t. 10, premier fasc., p. 771-772, Paris: Masson et Cle, 975 8 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 Boyeri, Th. 1888. Ueber partielle Befruchtung. Sitzber. Gesell. Morph Physiol,, Mun- chen, Bd. 4. s. 64-72. 31915, ' Uber die Entstehung der Eugsterschen Zwitterbienen. Ardh. entw.- mech. Org., Bd. 41, s. 264-811. - 38 BULLETIN FLORIDA STATE MUSEUM Vol. 5 Dalia Torie, K. W., und H. Friese 1899. Die hermaphroditen und gynandromorphen Hymenopteren. Ben nat.- · med. Ver. Inbsbrack, Bd, 24, s. -1-96, (Jahrg. 1898) Donisthorpe, H. 1946. Fifty gynandromorphic ants taken in a single colony of Myrmica sabuleti Meinert in Ireland. Entomologist, vol. 79, pp. 121-131. Enderlein, G. 1913. Ein hervorragender Zwitter von Xylocopa mendozand aus Argentinien. Mit ein Verzeichnis aller bisher beobachteten gynandromorphen Hy- menopteren. Stett. ent. Zeit.. Bd. 74, s. 124-140. Kontkanen, P. 1985. Eine gynandromorphe Wespe, Ancistrocerus parietuni L. Ann. En- tomol. Fennieae, T. 1, pp. 59-60. Krombein, K. V. 1949. Two new gynandromorphs, with a list 6f previbusly recorded sexual ab- errations in the scolioid wasps. Proc. U. S. Nat. Mus., vol. 100, pp. 55-59. Leclercq, J. 1958. Un cas extraordinaire de gynandromorphisme chez Hatictus sexcinctus, (Hym. Apidae). Bull. Inst. roy. Sci. nat. Belgique, T. 29, no. 46, 4 p. Leininger, H. 1926. Ein_lateraler Zwitter von Odunerus reniformis Ginel. (Hym. Vespidae). Verh. Naturwiss. Ver. Karlsruhe. Bd. 80. a. 162-167. Mackenson, 0. 1951. Viability and sex determination in the honey bee (Apis inellifera L.). Genetics, vol. 36, pA 500-509. Malloch, J. R. 1924. A gynandrom6rph of Odunerus conformis Sauss. (Hymenoptera, Acule- ata). Ent. News, vol. 85, p. 70. Mitchell, T. B. 1929. Sex anomalies in the genus Megachile, with descriptions of new speties. (Hymenoptera: Megachilidae). Trans. Amer. Ent. Soc., Vol. 54, pp 821-888. Morgan, T. H. 1905. An alternative interpretation of the origin of gynandromorphous insects. Science, vol. 21, pp. 682-685. Mo]ice, T. D. 1901. Hermaphrodite specimen of Podalirius retuaus, L. Ent. Mo. Mag., vol. 87, pp. 141-142. 1959 COOPER: A BILATERALLY GYNANDROMORPH 39 Perkins, C. D. F. 1901. Hermaphrodite Hawaiian O(lynerus. Ent, Mo. Mag., vol. 37, pp 139-140. 1922. Some instances of herrnaphr6ditism in bees and wasps. Jour. Torquay Nat. Hist. Soc., vol. 8, pp. 181-186. Rengel, C. 1917. Ein Zwitter von Odi,nerus cattosus Thoms. Sitzber: Gesell. Naturforsch. Fr. Berlin, Jahrg. 1917, s. 99-102. Rothenbuhler, W. D. 1957. · Diploid.male tissue as new evidence on sex determination in honey bees. Jour. Hered., vol. 48, pp. 160-168. 1958. Progress and problems in the analysis of gynandromorphic honey bees. Proc. Xth Int. Congr. Ent. (Montreal), vol. 2 (1956), pp 867-874. Rothenbuhler, W. C., J. W. Gowen, and O. W. Park ~ 1952. Androgenesis with zygogenesis in gynandromorphic honey bees (Apis mellifera L.). Science, vol. 115, pp. 687-638. Saussure, H. de 1856. Etudes sur la famille des vespides. III. La monographie des masariens et un supplement a la monographie des eum6niens. p. 141-142, Paris, 352 p. Sieb6ld, C. Th. von 1864. Ueber Zwitterbienen. Zeitsch. wiss. Zool., Bd. 14, s. 78-80. Tucker, K. W. 1958. Automictic parthenogenesis in the honey bee. Genetics, vol. 43, pp 299-816. Wheeler, W. M. 1908. Some new gynandromorphous ants, with a review of the previously recorded cases. Bull. Amer. Mus. Nat. Hist., vol. 19, pp. 653-688. 1937. Mosaies and other anomdlies among ants. Cambridge: Harvard Uni- versity Press. 95 pp. Whiting, P. W. 1928. Mosaicism and mutation in Habrobracon. Biol. Bull; vol. 54, pp 289-307. 1981. Diploid male parts in gynandromorphs of Habrobracon. Biol. Bull., vol. 61, pp. 478-486. 1982. Modification of traits in mosaics from binucleate eggs of Habrobracon. Biol. Bull., vol. 63, pA 296-309. 1985. Sex determination in bees and wasps. jour. Hered., vol. 26, pp. 268- 278. 1948. Androgenesis in the parasitic wasp Habrobracon. Jour. Hered., vol. 84, pp. 855-866. 40 BULLETIN FLORIDA STATE MUSEUM Vol. 5 1947. Some experiments with Melittobia and other wasps. Jour. Hered., vol. 88, pp. 11-20. 1958. Diploid males and triploid females in Habrobracon and Monnoniella. Bull. Assoc. Southeastern Biol., vol. 5, p. 16. Whiting, P. W., and M. F. Stancati 1981. A gynandromorph of Habrobracon from a post-reduced binucleate egg. Biol. Bull., vol. 61, pp. 481-484. Whiting, P. W., and E. J. Wenstrup 1948. Fertile gynaridromorphs in Habrobracon. Jour. Hered., vol. 28, pp. 81-38. Whiting, P. W., and A. R. Whiting 1927. Gynandrom6rphs and other irregular types in Habrobracon. Biol. Bull., vol. 52, pp. 89-121. Contributions to the BULLETIN OF THE FLORIDA STATE MUSEUM may be in any field of biQ10gy. 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