1) General zoological data of species
As the name suggests, this species of the five rhinoceroses comes from India and is the largest. Adult Indian rhinoceroses weigh approximately 2,000-4,000 kg. A comprehensive review addresses and illustrates all aspects of Indian rhinoceros questions (Rüedi, 1984). Zoogeographical and evolutionary data are to be found by Groves (1972, 1975).
2) General gestational data
The gestational length of the Indian rhinoceros is 462-491 days (Laurie et al., 1983). Since writing this chapter, a new calf was born to a multiparous female with good breeding dates; her gestation was 491 days long. That placenta weighed 6,650 g.; furthermore, a primiparous female delivered a healthy calf whose placenta weighed 3,550 g and measured 25 cm in greatest diameter. The remarkable aspect of that placenta was the enormous size of the hippomanes. They are depicted below; they weighed 368 g. With the exception of size (female black rhinos weigh about 1,600 kg, the Indian rhino weighs <1,600 kg) that is reflected in the membranes, no significant difference was found in the morphology of placentas from white, black and Indian rhinoceroses (Please see the chapter on white rhinoceros). They implant in both uterine horns, the fetus being located mostly in one horn, and the placenta extending into the other. The weight at birth is 40-80 kg, and females mature at about 3 years. One young is born only.
The placental weight at term, including the membranes but without umbilical cord, varies between 4,200 and 7,600 g in our large experience. The most recently born Indian rhinoceros in San Diego had a placental weight of 6,600 g. It measured 270 cm from side to side, 100 cm in greatest width and was very thin, less that 0.5 cm. In May, 2007, a term stillborn fetus was delivered of a primigravid female at the Wild Animal Park of the San Diego Zoo; in contrast to most other deliveries, this did not occur in a “Boma” but in the enclosure. It weighed 57 kg and had a 4,600 g placenta attaché with very short umbilical cord. The hippomanes attached weighed 180 g, were flattened and measured 18x8.5x2.5 cm.
|Two Greater Indian rhinoceroses at the San Diego Wild Animal Park.|
|Stillborn term calf.|
Early implantational stages were depicted by the sonographic study of Radcliffe et al. (1997). The earliest stage shown occurred 15 days after ovulation, with embryonic definition visible on day 23, and heartbeat on day 26.
The exact time of implantation, however, has not yet been determined. The placenta is diffusely villous, with larger streak-like areas of their absence. This is one reason for the organ to be called "Placenta villosa diffusa incompleta". These bare areas are also referred to as "streets (Strassen)". They show up extremely well when the placenta is trans-illuminated. They generally follow the larger blood vessels.
|Maternal aspect of rhinoceros placenta with white "Strassen" along major vessels.|
|This is a photograph of a portion of the placenta of an Indian rhinoceros, slightly trans-illuminated. It shows broad areas of absent villous (red) regions, so-called "streets" follow the major fetal blood vessels.|
The organ implants in both uterine horns, with the fetus located mostly in one horn, and the placenta extending to both sides.
|Schematic representation of rhinoceros placentation|
4) General characteristics of placenta
The placentas are very large and very thin. One weighing 7,600 g is depicted here. It measured 190 cm in length, 100 cm in greatest width, and 50 cm in smallest diameter. It was 2 mm in thickness. The placenta of a term Indian rhinoceros weighed 5,300 g, with an additional 950 g of amnionic and allantoic membranes. It measured 230 cm in greatest width. Another recently obtained placenta from a term, surviving Indian rhinoceros weighed 5.500 g, had dimensions of 270 x 98 cm and a 5 cm cord but with four blood vessels. In yet another additional specimen from a term, surviving infant, the umbilical cord had three vessels and the weight of the placenta was 5,000 g; yet another weighed 3,550 g, this one from a primipara. These are diffuse placentas without cotyledons, and of an epitheliochorial barrier character. Many placentas have on their maternal surfaces band-like connective tissue areas that are bare of villi. They usually follow the lesser curvature of the placenta and the larger blood vessels; they have been referred to as “Strassen” (streets) in the Suisse literature. They were depicted earlier.
In December, 2005 a stillborn female calf weighing 77 kg was delivered of an experienced female; it could not be resuscitated. The placenta weighed 6,300 g, was 280 cm long and had the usual characteristics with a 5 cm cord and three vessels. The remarkable aspect of this gestation was the course of the umbilical vein in the fetal abdomen. It was hugely dilated (up to 10 cm) shortly after entering the abdomen, filled with fresh clot and, when dissected further, there was no obstruction or another feature that could explain the aneurysmal distention.
|Fetal surface of rhinoceros placentas with short umbilical cord.|
Exceptionally large hippomanes (368 g).
|Section of hippomanes with crystalloid inclusions.|
|The hippomanes in polarized light microscopy.|
In addition, there were numerous round, yellowish squamous patches projecting on the amnionic surface of one Indian rhinoceros placenta that measured 0.5-1 cm. They were absent in another specimen. Naaktgeboren & Zwillenberg (1961) have discussed these structures at great length, for numerous species, but especially the cow.
|Amnionic surface of Indian rhinoceros placenta with round nodules|
Leaf-like and folded villi were alluded to by Ludwig et al. (1965), representing thinner and others, taller epithelial structures. Histochemical reactions showed only minor differences (Ludwig & Müller, 1965). Occasional binucleated trophoblastic cells have been described, as are also found in the horse and many ungulates. But there is no uterine invasion by trophoblast. The electronmicroscopic study (Ludwig & Villiger, 1965) identified similarities to equine placentas and numerous trophoblastic transport vesicles were seen.
The allantoic sac is anchored to the chorion by thin connective strands (Dolinar et al., 1965). While the amnionic epithelium is very thin and flat, the allantoic sac is lined by cuboidal to columnar epithelium. The allantois is diffusely vascularized.
|Higher magnification of white rhinoceros villi.|
|Higher power of the villous surface of an Indian rhinoceros placenta with "Strassen".|
|Structure of amnion/allantois (left), chorion and villi of a mature Indian rhinoceros placenta at right.|
|Another view of the placental surface (right) with villous branches extending left.|
A complete review of all publications on three species of rhinocerotidae placentas was published by Benirschke & Lowenstine, 1995. This report contained details in tabular form. It is printed at the end of the chapter for the convenience of the reader.
Details of barrier structure
|Umbilical cord of stillborn fetus.|
|Fetal surface of Southern white rhinoceros placenta.|
A remarkably long umbilical cord (57.8 cm) in an Indian rhinoceros was described by Dolinar et al. (1965). The umbilical cord has 4 large umbilical vessels and a very large number of small vessels that provide the circulation for the allantois. It also has a large allantoic duct. A vitelline duct is absent. The surface of umbilical cords has squamous plaques.
|Amnion at left (flat epithelium) allantois right (columnar epithelium).|
9) Trophoblast external to barrier
There is no trophoblast beyond the villous structures.
|Indian rhinoceros chorionic surface with abnormally vacuolated trophoblast at right, overlying debris.|
16) Physiological data
Whatever relevant physiologic data are available have been provided in review form by Silberman & Fulton (1979), and as bibliography by Miller (1983).
17) Other resources
Cell strains of four species are available through CRES at the Zoological Society of San Diego by calling on Dr. Oliver Ryder at email@example.com; none are available of the exceptionally rare Javan rhinoceros.
18) Other aspects of interest:
Additional literature copied for convenience here. (Because this reference is so difficult to obtain in print, it is here reprinted without the figures.)
THE PLACENTA OF THE RHINOCEROTIDAE
Cell strains of four species of rhinoceros are available from the “Frozen zoo” at the Zoological Society of San Diego: www.FrozenZoo@sandiegozoo.org
Amoroso, E.C.: Placentation. In, Marshall's Physiology of Reproduction. A.S. Parkes, ed., 3rd ed., Vol. II. London. Longmans, Green, 1952.
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Benirschke, K. and Calle, P.P.: The placenta of the Beluga whale (Delphinapterus leuca). Verh. Ber. Erkg. Zootiere 36:309-314, 1994.
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