Search

Trophoblastic invasion and remodeling of the uteroplacental (spiral) arteries in primates are well-documented, but virally nothing is known of the early stages of these phenomena. Therefore, we examined invasion of the maternal vasculature in macaques and baboons at, and immediately following, implantation. Following penetration of the uterine epithelium (day 9), trophoblast spreads along the residual epithelial basal lamina. By day 10, cytoplasmic processes penetrate the epithelial and endothelial basal laminae, and syncytial trophoblast insinuates itself between maternal endothelial cells. As lacunae develop, both syncytial and cytotrophoblast are exposed to maternal blood. Endovascular cytotrophoblast was first observed in subepithelial dilated capillaries and venules. These vessels are lined by increasingly hypertrophied endothelial cells. The spiral arterioles are unmodified at this time. Particularly interesting was the observation that there is rapid extensive endovascular trophoblast invasion of the spiral arterioles immediately beneath the implantation site. By day 14-16 nearly all of the small arterioles directly beneath the site are completely occluded. There is no invasion of the veins in this region. Somewhat later, the deeper arterioles in the principal zone are invaded. Rather than a continuous stream of cells invading the deeper arterioles, these endovascular cells occur in clusters ranging from a few cells to groups of cells that completely plug the lumen. Our results indicate that trophoblastic invasion of maternal vessels occurs very early; and, at least initially, trophoblast can migrate between and along endothelial cells without causing their lysis. The endovascular cells eventually interrupt the endothelial lining of the arterioles and penetrate the walls of the vessels. The occlusion of arterioles underneath the site suggests that circulation through the lacunae at this stage is indirect. Corresponding stages of human development were examined, and no invasion of arterioles could be observed prior to formation of an extensive cytotrophoblastic shell.

The structure of the equine chorionic girdle between days 28 and 42 of gestation was examined. Of particular interest were differentiation of trophoblastic cells within the girdle, adhesion between girdle and endometrium, invasion and displacement of the uterine epithelium, and the nature of the endometrium when girdle cells migrate into it to form endometrial cup cells. The chorionic girdle, identified initially as a band of tall columnar cells, becomes a stratified columnar epithelium indented by clefts and pits. Adhesion to and penetration through the endometrial luminal epithelium are rapid and occur initially in very limited areas. Stromal invasion occurs as strands of contiguous trophoblast cells invade through the basal lamina. Only girdle cells that are adjacent to the basal lamina or have entered the endometrial stroma undergo hypertrophy and differentiate into cup cells. At the initiation of trophoblastic invasion, the luminal epithelium contains numerous, large, intraepithelial, granular lymphocytes; small lymphocytes then accumulate in the stroma, but by day 42 lymphocytes are largely confined to the periphery of the cup. Although adhesion of trophoblast to the endometrial surface is initiated by small groups of girdle cells on restricted areas of the endometrial folds, the area is then increased by new areas of adhesion and by expansion of the initial invasion. Areas of girdle cells that do not attach undergo necrosis, as do superficial portions of areas of invasion. Consequently the girdle cells that form cups may be a minority of the original population. It is suggested that the differentiation of girdle cells is closely programmed and that cells that do not reach the stroma become necrotic at the same time that endometrial cup cells are differentiating.

Rat gestation sites were examined on days 7 through 9 of pregnancy by light microscopy and transmission and scanning electron microscopy to determine the extent of vascular modifications in the vicinity of the mesometrial part of the implantation chamber (mesometrial chamber). At a later time, the mesometrial chamber is, in conjunction with the uterine lumen, the site of chorioallantoic placenta formation. On day 7, in the vicinity of the mesometrial chamber, vessels derived from a subepithelial capillary plexus and venules draining the plexus were dilating. By early day 8, this network of thin-walled dilated vessels (sinusoids) was further enlarged and consisted primarily of hypertrophied endothelial cells with indistinct basal laminas. Sinusoids were frequently close to the mesometrial chamber's luminal surface which was devoid of epithelial cells but was lined by decidual cell processes and extracellular matrix. By late day 8, cytoplasmic projections of endothelial cells extended between healthy-appearing decidual cells and out onto the mesometrial chamber's luminal surface, and endothelial cells were sometimes found on the luminal surface indicating that endothelial cells were migrating. The presence of maternal blood cells in the mesometrial chamber lumen suggested that there was continuity between the chamber and blood-vessel lumens. On day 9, the mesometrial chamber was completely lined with hypertrophied endothelial cells, and sinusoid lumens were clearly continuous with the lumen of the mesometrial chamber. Mesometrial sinusoids and possibly the mesometrial chamber lumen were continuous with vessels in vicinity of the uterine lumen that were fed by mesometrial arterial vessels. Clearing of the mesometrial chamber lumen during perfusion fixation via the maternal vasculature indicated the patency of this luminal space and its confluence with mesometrial arterial vessels and sinusoids. The conceptus occupied an antimesometrial position in the implantation chamber on days 7 through 9, and it was not in direct contact with uterine tissues in the vicinity of the mesometrial chamber. These observations suggest that angiogenesis, not trophoblast invasion or decidual cell death, plays a major role in the opening of maternal vessels into the mesometrial chamber lumen before the formation of the chorioallantoic placenta.

On days 7 and 8 of pregnancy, mesometrial regions of rat gestation sites were examined by light microscopy and transmission electron microscopy to determine what changes occur before the chorioallantoic placenta forms in that region. By day 7, gestation sites contained a uterine lumen mesometrially and an antimesometrial extension of the uterine lumen, the implantation chamber. The implantation chamber consisted of a mesometrial chamber between the uterine lumen and the conceptus, an antimesometrial chamber that contained the conceptus, and a decidual crypt antimesometrial to the conceptus. Stromal cells that formed the walls of the implantation chamber were closely packed decidual cells, while those that surrounded the uterine lumen were loosely arranged. Late on day 7, a portion of the epithelium lining the mesometrial chamber was degenerating, but this area of initial degeneration was never adjacent to the antimesometrial chamber. By early day 8, most of the epithelial cells lining the mesometrial chamber were degenerating and were being sloughed into the chamber lumen. Although degeneration of these epithelial cells morphologically resembled necrosis, it was precisely controlled, since adjacent epithelial cells lining the uterine lumen remained healthy. The space that separated the denuded luminal surface of the mesometrial chamber from underlying decidual cells became wider and was occupied by an extracellular matrix rich in cross-banded collagen fibrils. Decidual cell processes, that earlier had penetrated the basal lamina beneath healthy epithelial cells, protruded into this matrix and penetrated the basal lamina at the luminal surface. By late day 8, large areas of denuded chamber wall were covered with decidual cell processes, little remained of the basal lamina, and cross-banded collagen fibrils were scarce in the area occupied by decidual cell processes. During the times studied, uterine tissues that formed the walls of the mesometrial chamber were not in direct contact with the conceptus. This study indicates that trophoblast does not play a direct role in epithelial degeneration, basal lamina penetration, or extracellular matrix modifications in the mesometrial region of implantation chambers where part of the chorioallantoic placenta forms, although trophoblast may be required to trigger or modulate some of the changes.

The annular hematoma of the shrew, Blarina brevicauda, is a specialized portion of the yolk-sac wall. In this study, we have examined the fine structure of the different cellular components of the anular hematoma. Small pieces of the gestation sacs from seven pregnant shrews were fixed in glutaraldehyde and osmium tetroxide and processed for transmission electron microscopy. In the area of the trophoblastic curtain, the maternal capillary endothelial cells were hypertrophied and syncytial trophoblast surrounded the capillaries. Cellular trophoblast covered part of the luminal surface of the curtain region, whereas masses of apparently degenerating syncytium were present on other areas of the surface. Maternal erythrocytes, released into the uterine lumen from the curtain region, were phagocytized and degraded by the columnar cells of the trophoblastic annulus. No evidence of iron or pigment accumulation was evident in the parietal endodermal cells underlying the annular trophoblast. Parietal endodermal cells were characterized by cuboidal shape, widely dilated intercellular spaces, and cytoplasm containing granular endoplasmic reticulum. Endodermal cells of the visceral yolk-sac accumulated large numbers of electron-dense granules as well as glycogen in their cytoplasm. Hemopoietic areas and vitelline capillaries were found subjacent to the visceral endoderm. The various portions of the yolk-sac wall of Blarina appear to perform complementary functions which are probably important in maternal-fetal iron transfer.

A series of peri-implantation stages of the rhesus monkey has been collected; these range from preimplantation blastocysts through initial implantation to early villus formation. The three earliest postimplantation specimens encompass the stages of penetration into and through the uterine luminal epithelium and into endometrial blood vessels. The day of pregnancy was established by radioimmunoassay of estrogen (E) levels to determine the preovulatory E peak, and in each instance the embryo was examined to determine the extent of development. The conceptus collected on day 9.5 of pregnancy was the earliest implantation stage; it ballooned above a depression in the endometrium to which it was firmly attached. A column of syncytial trophoblast penetrated into the uterine epithelium to the basal lamina of the latter. The syncytial trophoblast shared junctional complexes with the uterine epithelial cells to which it was apposed at the margin of the site of epithelial penetration. Basal to the apical junction complexes, processes of syncytium indented uterine epithelial cells. Several epithelial cells had been partially isolated and surrounded by flanges of syncytial trophoblast. In the next specimen, at 10.0 days after ovulation, the uterine epithelium had initiated the epithelial plaque reaction. The trophoblast had extended along the residual basal lamina of the uterine epithelium and into the neck of an adjacent uterine gland. Cytotrophoblast was abundant in the central region of the implantation site, and was intermixed with syncytium which formed the majority of the peripheral trophoblast. In several places clefts had formed in the syncytial trophoblast; these clefts were lined with microvilli, had intermicrovillous caveolae, and consequently more closely resemble the trophoblast that eventually lines the intervillous spaces than the trophoblast involved in initial invasion. In the day-10.5 specimen, in addition to prelacunar clefts, lacunae containing maternal blood were present for the first time. The basal lamina was penetrated in many places, and syncytial trophoblast was interposed between maternal endothelial cells of the underlying vessels. It was concluded that syncytial trophoblast is the first tissue to penetrate the uterine luminal epithelium; that the basal lamina of the uterine luminal epithelium, but not the basal lamina of endothelium, constitutes a temporary barrier to trophoblast penetration; that invasion is accomplished with less destruction of maternal tissue than previously suggested; and that the rapid superficial growth of the placenta is made possible by the early tapping of the endometrial vessels.

Corpora lutea were obtained from ten pregnant rhesus monkeys during implantation, and the ultrastructure of granulosa and theca lutein cells was characterized. Specimens were individually staged with regard to the extent of implantation and the relationship to the rise in circulating progesterone and estrogen which is characteristic of early pregnancy. Structural changes characteristic of granulosa lutein cells occurring during implantation included: change in form of endoplasmic reticulum from predominantly agranular tubules to predominantly granular cisternae; reduction in size and number of lipid droplets; increase in area occupied by the Golgi and increase in length of the cisternae of the Golgi complex; development of numerous microvillus-lined intracellular spaces; increase in numbers of membrane-bound dense bodies including peroxisomelike bodies, multivesicular bodies within lobopodia, and other lysosomelike bodies; and alterations of mitochondrial cristae. These changes were suggestive of the production of a secretory protein, rapid utilization of existing steroid precursor reserves for the production of progesterone, and a reduction in capability for steroid precursor accumulation and processing by granulosa lutein cells. Structural changes characteristic of theca lutein cells occurring during implantation included an increase in size and number of lipid droplets, an increase in agranular endoplasmic reticulum, and an increase in area occupied by the Golgi complex. These changes were suggestive of an increased capability for steroid precursor accumulation and processing, perhaps for estrogen production, by the theca lutein cells.

Implantation sites were obtained from rats at various stages of pregnancy and were studied by light microscopy and scanning electron microscopy. Early in pregnancy the uterine luminal epithelium and the decidual cells in the implantation site formed an implantation chamber containing the conceptus. The epithelial cells lining the chamber and the mouth of the chamber degenerated, and the uterine lumen that was mesometrial to the conceptus was obliterated such that the uterine lumen became discontinuous, and the luminal epithelia of intersite areas were isolated. As the conceptus continued to grow, the decidua-conceptus unit bulged into the intersite areas and was partially covered by an epithelium that eventually became discontinuous and degenerated. Once this had occurred, the luminal epithelium of the intersite areas reestablished contact antimesometrial to the decidua-conceptus unit, and the uterine lumen was again continuous. However, the epithelium lining the lumen was not complete in the mesometrial region because of the vascular connections between the uterine stroma and the placenta. Factors influencing the restructuring of the uterine luminal epithelium were discussed.

A method of flushing the oviduct and/or uterus of rhesus monkeys was used to obtain a number of preimplantation stages, of which four cleavage stages and seven blastocysts that were judged to be normal were studied cytologically using transmission electron microscopy. In addition to the usual changes in mitochondria and endoplasmic reticulum that accompany differentiation of the blastomeres, the blastocysts with zonae showed sequestration of areas of cytoplasm. The first indications of junctional complexes were short stretches of parallel membrane with a slightly increased density found in the morula stage. Blastocysts developed typical apical junctional complexes, but in addition showed extensive gap junctions linking trophoblast and inner cell mass, and epiblast and differentiating endoderm. Endodermal differentiation occurred at about the same time that a basal lamina was found under mural trophoblast and epiblast (but not polar trophoblast or endoderm). Enlarged torn zonae were found in association with one blastocyst and unaccompanied by blastocysts, including a case in which the animal subsequently prove to be pregnant. This observation suggests that hatching is a normal feature of zonal escape in this species. The trophoblast of blastocysts without zonae had well-formed apical absorptive areas and, in some instances, long irregular microvilli in the area near the inner cell mass. Cell debris, vacuoles containing debris and isolated cells, although variable, were common features of the preimplantation stage in the rhesus monkey.

Fixed uteri from rats on the afternoon of day 6 of pregnancy were split to expose the implantation chambers, their enclosed blastocysts, and the imprints of the blastocyst on the adjacent epithelium of the chamber. Some of the implantation chambers were prepared for scanning electron microscopy; other chambers were treated with colloidal iron hydroxide, with cationized ferritin, or with the tannic acid method, and subsequently were prepared for transmission electron microscopy. In this manner, the disposition of the surface-coat markers on the surface of the blastocysts, surface of the uterus within the chamber, and the surface of the uterus that had been apposed to a blastocyst were compared. Despite the pronounced morphological differences between the microvilli of the uterine luminal epithelium in the imprint and those in the rest of the chamber, the binding of the markers was remarkably similar. No evidence of removal of surface coat could be found in that area of the uterus in contact with the blastocyst. In addition, in two instances in the cationized ferritin-treated material, and in another instance in tannic acid-stained material, regions of the apparently adhering trophoblastic cell membranes and uterine cell membranes had abundant coat materials and, possibly, even secretory materials interposed. When blastocyst-sized glass beads were introduced into uteri from animals made pseudopregnant or unilaterally pregnant, the beads failed to elicit a decidual response and made an imprint that did not resemble the imprint of a blastocyst in an implantation chamber. It was concluded that, at least in the initial stages of adhesion, the blastocyst does not bring about a physical removal of the demonstrable aspects of the surface coat of the uterus. It was concluded further, that glass beads are not a suitable object for mimicking a blastocyst in the rat uterus.

Rat gestation sites were obtained on days 10 through 16 of normal pregnancy. Light and electron microscopic examination of day-10 sites revealed a consistent complex pattern of stromal cell morphologies. Six distinct regions were identified: an antimesometrial region of epithelioid decidual cells that form the gestation chamber containing the embryo and extraembryonic membranes; an abembryonic antimesometrial decidual region, the decidual crypt, where the cells are separated by large extracellular spaces; a mesometrial region with granule-containing cells and mesometrial decidual cells; a region of spiny cells that are lateral to the antimesometrial decidual cells and continuous with the mesometrial decidual cells; and a region of undifferentiated stromal cells adjacent to the myometrium. Between days 12 and 16, the antimesometrial decidua becomes thinner and is eventually sloughed into the newly formed uterine lumen. The role of the antimesometrial decidual cells is discussed with reference to trophoblast invasiveness, protein synthesis, and especially remodeling of the gestation chamber. Differences between decidua and deciduoma are considered.

The yolk sac of the little brown bat is unusual in that during the course of gestation both the inner endodermal cells (bordering the yolk sac cavity) and outer mesothelium (facing the exocelom) form simple columnar epithelia which persist throughout gestation. These endodermal cells develop an extensive system of agranular endoplasmic reticulum, numerous lipid droplets and unusual "giant" mitochondria. During development the Golgi apparatus changes position from the apical to the basal side of the nucleus, reversing the polarity of the cells. In general, the endodermal cells have cytological features suggestive of synthetic or secretory activity. The mesothelial cells develop an extensive "absorptive apparatus" in their apices, while large crystalloid-containing granules become numerous in their basal cytoplasm. The mesothelial cells have large deposits of glycogen, especially during mid-gestation, but few mitochondria and little granular endoplasmic reticulum. Endodermal cells do not absorb exogenous protein (peroxidase) even if it is injected directly into the yolk sac cavity. However, placement of peroxidase either in the exocelom or in the maternal vascular system results in the appearance of this protein in the "absorptive apparatus" of mesothelial cells as well as in macrophages in the stroma of the yolk sac. While evidence of absorption was clear, no direct evidence of transport of tracer to fetal blood vascular system was obtained. It is postulated that a major function of the hypertrophied mesothelial cells during gestation is the absorption of proteins and possibly other substances from the exocelomic fluid. The major function of the hypertrophied endodermal cells may be synthesis and secretion of substances into the fetal circulation.

In this continuing series of studies of implantation in the rhesus monkey, eight specimens, ranging in gestation age from 9.5 days to 16.5 days after ovulation, were examined with a focus on localized modifications in the endometrium as a response to implantation. Additionally, evidence of continuing changes in early pregnancy was provided by three specimens at the end of the first month of gestation (days 24, 28, and 35). The responses of the endometrium to pregnancy start with a localized accumulation of stromal eosinophils, which is rapidly followed by epithelial plaque formation in the basal cells of the luminal epithelium and gland necks. Plaque cells hypertrophy, develop marginal dense granules, and accumulate glycogen. They form a pad underlying the margins but not the central zone of the implantation site. However, some degenerating plaque cells are found as early as day 15; and little more than a region of leukocytic infiltration remains of the plaque by day 35. Shortly after the plaque response is initiated there is a striking subepithelial edema surrounding the plaque, and the venular capillaries enlarge by engorgement and by endothelial hyperplasia. The endothelial cells subsequently hypertrophy, resulting in a largely columnar endothelium. There is a localized decidual cell response, consisting of an increase in rough endoplasmic reticulum and in filaments, but only a moderate amount of hypertrophy of these cells. Endometrial granular cells become more conspicuous in the area as they accumulate glycogen. Patches of large pale cells appear in the lumen and walls of arterioles subjacent to the implantation site, but the cytology of these cells provided insufficient clues to their origin (cytotrophoblast?). Although the endometrial responses described are impressive and diverse, their advantages to the organism are not obvious. The hypertrophy of the anastomotic capillary bed that accompanies plaque formation may well provide an extensive vascular network available to the developing trophoblastic lacunae. The role of endometrial granular cells, decidual cells, and even plaque cells may be more related to their largely unexplored secretory activity than to their physical contribution to the formation of the basal plate.

Differentiation of extraembryonic mesoderm in the rhesus monkey was studied from the epithelial penetration stage of implantation (stage 4) through the first week of postimplantation development (to stage 6). It was found that the first cells that appeared between the primitive endoderm (hypoblast) and trophoblast were separated from the latter by a basal lamina but appeared to be either loosely attached to the endoderm or to have been detached from it. Cells in this intermediate position differentiated cytologically into mesenchymal cells, which, by stage 5, had a distinctive intraendoplasmic reticulum marker. This differentiation occurred prior to the time at which the primitive streak could be recognized. By the time the primitive streak was readily discernible (stage 6), the extraembryonic mesoderm had already produced substantial extracellular matrix. The sequence of differentiation was repeated, with a 1- to 2-day lag, in the secondary implantation site. No evidence of a contribution from cytotrophoblast or primitive streak to the extraembryonic mesoderm was found. It is concluded that the origin of the first extraembryonic mesoderm in the rhesus monkey is probably a two-step process, with formation of a reticulum from primitive endoderm followed by differentiation in situ into mesenchymal cells. The first blood vessels formed also differentiated in situ from the extraembryonic mesenchymal cells. Primitive capillaries were identifiable as early as the 13th day of pregnancy.

The transition from the trophoblastic plate stage to the early lacunar stage was examined in a series of implantation sites from the rhesus monkey, timed on the basis of the preovulatory estrogen peak, and prepared for transmission electron microscopy. This transition was compared with specimens from stage 5a, b, and c in the Carnegie collection of human embryos. The transition was marked by the differentiation of a new type of syncytial trophoblast--namely, a unilaminar microvillous polarized syncytium, which developed throughout the trophoblastic plate, forming characteristic intrasyncytial clefts. The rapid development of this type of syncytium created a nonclotting chamber for maternal blood wherever trophoblast intrusion into maternal vessels created confluence. Although the nature of the material in the Carnegie series precluded cytological characterization of the trophoblast, there is evidence that a similar transition occurs in human trophoblast and that in the human also the appearance of lacunae marks a change from an early invasive trophoblast to a situation in which growth is more significant.

The inner cell mass of the blastocyst has differentiated into epiblast and hypoblast (primitive endoderm) prior to implantation. Since endoderm cells extend beyond the epiblast, it can be considered that both parietal and visceral endoderm are present. At implantation, epiblast cells begin to show marked evidence of polarity. They form a spherical aggregate with their basal ends toward the basal lamina and apical ends toward the interior. The potential for an internal space is formed by this change in polarity of the cells. No cytological evidence of separation of those cells that will form amniotic epithelium from the rest of the epiblast is seen until a cavity begins to form. The amniotic epithelium is originally contiguous with overlying cytotrophoblast, and a diverticulum remains in this position during early development. Epiblast forms a pseudostratified columnar epithelium, but dividing cells are situated toward the amniotic cavity rather than basally. The first evidence of a trilaminar disc occurs when a strand of cells contiguous with epiblast is found extending toward visceral endoderm. These presumptive mesoderm cells are undifferentiated, whereas extraembryonic mesoderm cells are already a distinct population forming extracellular materials. After implantation, visceral endoderm cells proliferate forming an irregular layer one to three cells thick. Visceral endoderm cells have smooth apical surfaces, but very irregular basal surfaces, and no basal lamina. At the margins of the disc, visceral endoderm is continuous with parietal endoderm and reflects back over the apices of the marginal visceral endoderm cells. This sacculation by visceral endoderm cells precedes pinching off of the secondary yolk sac from the remaining primary yolk sac.

The structure and development of the female reproductive tract, fetal membranes and placental have not previously been recorded for any member of the family Thyropteridae. Recently implanted embryos were obtained in late January, limb-bud stages in March, and full-term fetuses in late May, suggesting a possible gestation length of approximately five months. It is likely, however, that Thyroptera experiences at least two breeding cycles per year. The uterus was narrowly bicornuate; the corpus uteri was unusually large and lacked the glandular density observed in the cornua. The cervix was long, pleated, and relatively aglandular. The oviducts opened at the apices of the cornua; oviductal papillae were absent. A bursa ovarii surrounded the ovary, but there was a small pore opening to the peritoneal cavity adjacent to the fimbriated end of the oviduct. Never more than a single embryo or fetus was present, and only a single corpus luteum was observed; thus Thyroptera, like most bats, is monovular. Ovoimplantation was interstitial; a decidua capsularis was present early but disappeared by the late limb-bud stage. The decidual reaction involved both glandular epithelium and stromal cells, but most of the decidua was destroyed by term. Amniogenesis was initiated after implantation, by cavitation. Primitive entoderm was formed precociously above, as well as below, the presumptive embryonic disc, and a thin extension of Reichert's membrane passed over the cell mass, separating it from the cytotrophoblast of the chorionic placenta. During the amniogenic period, the yolk-sac entoderm fused to the parietal trophoblast via an intervening Reichert's membrane, forming an extensive bilaminar omphalopleure; this was rapidly converted to a trilaminar structure in early post-implantation stages. An avascular chorio-vitelline relationship involved most of the chorionic wall in early post-implantation stages and persisted to term in the abembryonic hemisphere after the partial inversion of the yolk-sac roof in late presomite embryos. The invaginated yolk-sac roof (splanchnopleure) also persisted to term as a viable paraplacental component. A small sac-like allantois was formed between late presomite and early limb-bud stages but disappeared by the late limb-bud stage. Development of the definitive chorioallantoic placenta resembled that in other bats, but the maternal endothelium disappeared relatively early, and trophoblastic differentiation was precocious. The ultrastructural organization of the interhemal membrane was hemodichorial, and otherwise generally resembled the organization previously described in vespertilionid bats. Similarities and differences in the structure of the uterus, placenta, and paraplacental organs of Thyropteridae, in comparison with other families of bats, are discussed. On the basis of fetal membrane characteristics, the Thyropteridae show closer affinities with the Phyllostomatoidea than with the Vespertilionoidea, to which they are presently assigned.