1) General Zoological Data
This beautiful African species has elegant, spiraled horns in the grey males; females do not have horns. A relevant web site gives the following information: “The range of the Greater Kudu extends from the east in Tanzania, Eritrea and Kenya into the south where they are found in Zambia, Angola, Namibia, Botswana, Zimbabwe and South Africa. They have also been introduced in small numbers into New Mexico. Their habitat includes thick bushveld, rocky hillsides, dry riverbeds and anywhere with a constant supply of water”. The evolutionary relationships of various bovid taxa were described with cytochrome b mtDNA data by Hassanin & Douzery (1999) and Matthee & Robinson (1999). More recently, Hernández Fernández & Vrba (2005) have summarized the relationships and constructed trees that include the kudu, eland and related species, and Gatesy et al. (1997) aligned the Bovini with Tragelaphinae by mtDNA studies of Artiodactyla.
2) General Gestational Data
Single young are the rule (Hayssen et al., 1993) and gestation is estimated to be 7-9 months with neonates weighing 16 kg (Nowak, 1999). The smaller female fetus shown below was found at necropsy as a singleton that measured 9 cm in crown-to-rump length. It was located in the right uterine horn but the placenta extended into the left horn as well. The normal length of gestation is 212-214 days (Mentis, 1972; Hayssen, V. et al., 1993). The usually single neonate weighs ~15 kg (Hayssen et al. (1993).
Early stages of implantation are not recorded. Implantation occurs on flat caruncles with a typical epitheliochorial, cotyledonary type of placenta evolving. Hradecky (1983) described implantation in right and left uterine horns but has found many more cotyledons (55 and 70) than seen in our immature specimen (21). He described four rows of caruncles and observed cotyledons to develop in the lesser horn as well. Mossman (1987) indicated that the placenta attaches mesometrially and that placentation is superficial. Carter (2008) provided suggestions as to how to access Mossman’s collection.
4) General Characterization of the Placenta
I have had two pregnant uteri available; one came from a female that was euthanized after sustaining severe injuries in fights. In its right uterine horn the dam contained a single female fetus weighing 2,900 g (normally 16,000 g at term!) that had a 37 cm CR length. After removing the fetus, the uterus and placenta together weighed 1,650 g. There were 21 flat cotyledons measuring up to 7 cm in width and 0.8 cm in thickness. The cord had a mesometrial central attachment and measured 13 cm in length. A single corpus luteum was present in the right ovary; several small cysts were present in the left ovary. The placenta differs little from that of the lesser kudu (see that chapter) except for the smaller number of cotyledons that were found at least in this single specimen. Hradecky et al. (1988b) depicted the dense band of connective tissue that separates the villous tissue from the endometrium. In addition to this specimen, Dr. P. Hradecky made available some slides of term placentas which he described in publications here annotated. The histology of these specimens is also here presented.
5) Details of fetal/maternal barrier
This is a typical epithelio-chorial placenta with long strands of maternal connective tissue separating the long, slender villi. The single-layered trophoblastic epithelium has a very large number of binucleate cells. Microvilli extend from the trophoblastic surface. The uterine epithelium in this specimen is single-layered and intact, albeit somewhat autolyzed. No trophoblastic infiltration into the endometrium occurs. Some trophoblastic cells have small yellow granular inclusions, but it is not clear that they are not artifacts of fixation. There is no subchorial “hematophagous region”, as seen in so many other species. The finer histologic features of the kudu placentome were described by Hradecky et al. (1988a). They measured the term villi to be 7 mm long and branched and somewhat different from those of the eland. The septa and epithelia are also described in this contribution. . Hradecky et al. (1988a) described that thee villous surfaces were more ‘corrugated’ than those of the eland. They described that: a) ‘the covering epithelium was 10 to 15 µm thick and was composed of polygonal; cells with visible boundaries and round nuclei’; and further b) ‘the crypt lining was 5 µm thick, had dark spindle-shaped nuclei, and varied between syncytial and cellular types’.
6) Umbilical cord
The cord of this first specimen measured 13 cm in length, had four large blood vessels and a central allantoic duct. In addition to these large chorioallantoic vessels there are numerous small vessels but they are not specifically congregated around the allantoic duct. There were numerous small surface granules composed of squamous metaplasia. The allantoic duct is lined with a flat urothelium. The second specimen’s umbilical cord was mesometrially attached, had four blood vessels and a large allantoic duct. The two arteries were considerably larger than the veins. At this earlier stage, there were no foci of squamous metaplasia covering the surface. Many small blood vessels were present in addition to the four larger ones but at this early stage, these vessels consisted of an endothelial lining only; there was no musculature. The umbilical cord of this specimen was 7 cm long and had no spirals. A term umbilical cord has not yet been measured.
7) Uteroplacental circulation
No details have been described.
8) Extraplacental membranes
The amnion the first specimen had numerous small foci of squamous metaplasia; the allantoic sac contained yellow urine without hippomanes. The membranes extended into the empty left sac, but there were no implanted cotyledons in that horn, in contrast to the second specimen. The composition of amnionic and allantoic fluid of term kudu was described by Hradecky (1984). It included volume and pH values. There is a large, lobulated allantoic sac but there is no decidua capsularis.
9) Trophoblast external to barrier
There is no infiltration of the basal endometrium.
The cervical canal was filled with dense, yellow mucus and was firmly closed.
No true decidua was found.
11) Various features
There are no other unusual features in this gestation. There is an ovarian bursa (Mossman & Duke, 1973).
The role of hormone production (placental lactogen) by the trophoblastic binucleate cells was explored by Wooding (1982). Atkinson et al. (1993) explored the glycoprotein production in ovine placentas by these cells. No other endocrine information is available. The ovaries of the second specimen measured 24&23 x 14&15 x 10&5 mm.
Greater kudu males have 31 chromosomes, females have 32 chromosomes. This discrepancy is due to a translocation between an autosome and the Y-chromosome (Wallace & Fairall, 1967, 1967; Wurster, 1972; Hsu & Benirschke, 1971). Hybrids between greater kudu and eland have been described (Gray, 1972; Jorge et al., 1976). The latter hybrid reported at least was sterile. Gallagher & Womack (1992) also studied the translocations of many bovid species, including the kudu and suggested that speciation may have been the result of translocation with isolation of resultant karyotypically rearranged individuals. Wallace & Fairall (1967) were the first to study this species’ chromosome complement, to be followed by Herzog et al. (1975). The centromeric heterochromatin was delineated by Dain & Dott (1982).
The lesser kudu has 38 chromosomes and is remarkable because of its fusion chromosome; both X and Y chromosomes have compound fusions, and we believe they are with the same autosome (# 13). Petit et al. (1994/5) also studied the relationships of these unusual animals. Both views are presented at the end in a “pedigree” of possible developments over time. The finding of translocations in both sex chromosomes contrasts with that of many other African ungulates that have only an X/A translocation, but possess a normal Y chromosome. These species thus have different numbers in males and females (Benirschke et al., 1980). Other views of evolutionary relationships, resulting from mtDNA studies, were presented by Matthee & Robinson (1999) and Hassanin & Douzery (1999).
I am not aware of any publications.
15) Pathological features
Griner (1983) saw 20 kudus at autopsy and found the cause that 16 births had occurred at the same time. Trauma killed 6 animals, malnutrition another four and one had myopathy, two had pulmonary abscesses; periodontal disease was another important aspect; a Sertoli cell tumor was found in an animal infected with Trichuris. Macivor & Horak (2003) searched for ixodid ticks in wild kudus and other antelopes in South Africa.
16) Physiologic data
Pospisil et al. (1984) provided data on blood counts in kudu as well as several other antelopes. Hradecky (1982) studied the uteri of a variety of antelopes in zoos and found the kudu to possess a Uterus bicornis similar to that of cattle. In a later publication (1984), he studied the composition of two kudu ‘fetal fluids’ (amnionic and allantoic separately) of specimens 20 and 54 cm long. Interestingly, they differed considerably from one another. The two hemoglobin components of kudu (Hb A and Hb B) were studied by Rodewald et al. (1985) who sequenced the complete amino-acid chains. They found much homology with cattle hemoglobin. The nature of myosin heavy chain isoforms was studied in several African ruminants, including the kudu, by Kohn et al. (2007). The response to fever from pneumonia was the topic of a study by Hetem et al. (2008), while Owen-Smith (1997) studied the energy balance during foraging. Hagey et al. (1997) described the changes in bile acid composition with age a many bovids.
17) Other resources
18) Other remarks – What additional Information is needed?
Early stages of implantation are not on record and more endocrine data are needed.
The animal photographs in this chapter come from the Zoological Society of San Diego. The new fetal specimen comes from Dr. Anneke Moresco at UC Davis. Some slides of a term placenta from a greater kudu were donated by Dr. Petr Hradecky.
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