Myslenkov A.I., Voloshina I.V.
Sikhote-Alin State Biosphere Reserve, 46 Partizanskaya St., Terney. Primorsky Krai, 69210 Russia
Sexual behaviour of the Amur goral from Middle Sikhote-Alin Range (Russia)
is discussed. In the 1970's and 1980's investigations of goral biology
were carried out in both the wild and the enclosure for a total of 5 000
hours (2 000 of them in the wild). About 40 wild gorals were individually
recognized by body colour and horn features.
Five gorals were marked with coloured ear tags. During visual observations the binocular 12x and the telescope 30x - 60x were used. The Amur goral performs mainly all the same behavioural patterns as the Himalayan Goral, but has a different pattern "stroking of the female's head", which is unique among Caprinae. Its repertoire is richer than serow, but poorer than those of Oreamnos and Rupicapra.
Key words: sexual behaviour, pattern, posture, rutting season, mating system, copulation, ungulate, goral,
The Amur Goral (Nemorhaedus caudatus raddeanus) is a rare Asian mammal.
It is included as an endangered species in Category I of the Red Data Book
of USSR (1984) and listed as Vulnerable in the 1996 IUCN Red List of Threatened
Animals (IUCN 1996). Species range occupies China, Korea, and the south
of the Russian Far East.
In Russia, goral range covers the Sikhote-Alin Range in Primorsky Krai and the Little Khingan Range in the southern part of Khabarovsky Krai along the border with China ( Voloshina & Myslenkov 1992).
There are 2 types of goral habitats: 1) rocky forests in mainland river valleys , and 2) rocky areas along the coast. The characteristic feature of the latter is the mix of open (rocks and meadows) and closed (oak forests) habitats. Most of the goral in Russia are concentrated in two Reserves - Lazo and Sikhote-Alin (Voloshina & Nesterov, 1992).
Knowledge of goral biology and behavioural information particularly are insufficient, while this species has spread range in China. The first sexual behaviour description of representative of genus Nemorahaedus was made by us (Myslenkov & Voloshina 1978). Further we studied other aspects of behaviour : feeding, marking, and territorial behaviour (Myslenkov & Voloshina 1989). The rutting behaviour of the Himalayan Goral was studied by Lovari & Apollonio (1994).
Our study were conducted in the Sikhote-Alin State Biosphere Reserve
(45( N, 137( E) which is situated in the Middle Sikhote-Alin Range (Primorsky
Krai, Russia). The most northeastern goral population is here. The numbers
of population are 190 individuals. Goral habitats cover about 2 000 ha
along the coast. The slopes of Abrek Mountain are the main habitat of gorals.
Fig. 1. Home ranges of three adult females and territory of adult male
during the rutting season. Shaded quadrates are centres of activity.
male N 26 ___ females: N 35 _ _ N 29 _ . _ N27 _ . . _
MATERIAL AND METHODS
In the 1970's and 1980's intensive investigations of goral biology in
the Sikhote-Alin Reserve were carried out (Myslenkov & Voloshina 1989).
An experimental enclosure of 6 ha was built on the edge of goral habitat.
Five gorals were captured for the enclosure, and then this group bred to
13 gorals (Voloshina 1981). Our eco-ethological observations were conducted
in both the wild and the enclosure for a total of 5 500 hours (2000 of
them in the wild). One part of the ethological investigation was the study
of sexual behavior in goral. About 40 wild gorals were individually recognized
by body colour and horn features. Five gorals were marked with coloured
Visual observations were made with aid of the binocular 12x and the telescope 30x - 60x. We used the sampling all occurrence (Altmann 1974). Behaviour patterns were classified and named in accordance with the descriptions by Geist (1971), Shaller (1977), and Lovari & Apollonio (1994).
The principal characteristics of goral social organization are territoriality
and small group living. Both sexes have home ranges which they mark year
round. Size of home ranges in males is 22 - 55 ha and in females 5 - 16
ha (Myslenkov 1978; Voloshina 1978) (Fig. 1).
Female's home ranges overlap significantly up to 80 %. Core areas which we define as areas with 50 % of all sighting overlap up to half, and centers of activity that is to say quadrates with maximum being usually don't overlap. Adult males defend their territories from other territorial males during the rutting season. Encounters between neighboring males are rare, but when they do occur, it is usually along territory borders males, where males display threats to each other at a distance. A resident male demonstrates patterns such as "neck-up", "horn-threat","rush-threat", and "chase", stranger - "neck-up" and "flight". After crossing of a territorial border, roles are changed.
Fig. 2. Dynamics of sexual activity.
Index = number of pair contacts per 10 animal/hour
Fig. 3. Seasonal dynamics of courtship posture frequency.
Serious fighting may occur between a territorial male and a subadult
male. Two-year-old males submit entirely to dominant adult male. A 3-year-old
male, that remains in his natal area will be in conflict with the dominant
adult male and fights may lead to physical trauma. This occurs when a territorial
male mates with any female, and 3-year-old male attempts to approach them.
An adult male demonstrates a series of threats, and if the subadult does
not retreat, but shows "horn-threat", the fight begins.
The rutting season begins in the end of October and ends in December (Myslenkov & Voloshina 1978). Courtship patterns peak in mid-November, when the greatest number of copulations is also observed (Fig. 1,2). The start of the rut is characterized by active courtship by a male of all females occupying his territory. If a male finds a female which is close to estrus, he spends most of his time near her. Then copulations begin and last for 1-2 days. After that a male begins to look for the next female using intensive olfaction. Thus, mating is divided into 2 stages: courtship and copulations. In the first stage, the key postures displayed by the male are: "low-stretch", "stroking of female's head", and "following". A female reacts to the male by "flight" or "horn-threat". A male approaches for one female, then another female until he finds a female approaching estrus (Fig. 4).
It is necessary to emphasize that "stroking of female's head" is unique pattern among Caprinae. Besides the demonstrative functions, it is probably a male's way of marking a greater number of females occupying or visiting his territory. During the stroking, a male attempts to sniff the posthorneal female gland and mark the female head with secretion of the interdigital gland (Fig. 5). Maybe the secretion of posthorneal gland gives the some information about approaching estrus. It is known that the degree of development and functioning of posthorneal gland correlates positively with the reproductive cycle (Schaffer 1940).
Fig. 4. A male approaches in "low-stretch", a female demonstrates "horn threat".
Thus, male behavior during the courtship stage serves to, detect estrus
females, keep them within his territory, and prepare them for copulation.
A copulation stage begins with the onset of estrus in a female. When a male displays "low-stretch" and the female doesn't threaten him, then he will sniff and lick her anogenital region. Then lip-curl almost always occurs. At the same time a female begins to raise her tail. This reaction (raising of tail) is a signal of the female's readiness for copulation. The copulation period during which the mountings follow lasts from 4 to 78 min (mean 32 min, n = 18). One pair has 2 - 4 such periods daily. The copulations commonly begin from the attempts to mount. Among all mountings, attempts were 38 %, unsuccessful mountings were 18 %, and copulations were 44 %. Duration of one copulation is from 2 to 8 sec (mean 4,1 sec, n = 290) (Fig. 6).
Fig. 5. Key posture of male courtship - "stroking of female's head".
Fig. 6. Mounting in goral.
Analysis of sequence and probability of copulation behavior patterns
shows the clear change in the male's postures (Fig.7).
After standing, one of the mounting type occurs and this is followed by
obligatory licking. Then lip-curl may occur (16 %), but more often the
standing and copulative cycle is repeated. The key postures in the copulation
stage are: in females - "copulative posture" with raised tail; in males
- "licking of anogenital region", and "mounting". A male mates with every
female for 1 - 2 days. The maximum number of copulations was 17 times daily,
and 24 times over for 2 days with one female. The total number of any mountings
was 48 and 59, respectively. Thus, a male mating with several females can
make up to 100 copulations in one breeding season. All male behavior serves
to synchronize the mating process.
Fig. 7. Sequence and probability of copulative postures.
Figure in circle indicates observed number of given posture. Figure on line indicates probability of transition one posture in another. Summarized data on 8 pairs. Single circle - male's postures, double circle - female's posture.
Representatives of the family Bovidae demonstrate different types of
mating systems. The most common mating system is polygyny (Owen-Smith 1977;
Gosling 1986). Among Caprinae we find such variations of polygyneous mating
system as "following", "female defense", and "territorial defense". The
latter is typical for the Amur Goral.
The Amur Goral is a territorial, small-group living animal. He occupies an intermediate place between solitary forest-dwellers (Capricornis crispus) and group open-dwellers ( Rupicapra rupicapra, Oreamnos americanus). The goral habitat is a complex of rocky-meadow and forest environments. The Amur Goral lives under more severe climatic conditions in Primorsky Krai than other subspecies of gorals, but it exhibits territoriality which is typical for species inhabiting areas with greater diversity and productivity of primary production. Stability and limited availability of goral habitats in Primorsky Krai may promote territoriality. Geist (1985) suggested that serow and goral represent the original stage of resource defenders in warm climates. The Japanese serow is typical solitary and territorial mammal inhabiting closed habitats (Kawamichi et al. 1987). The Amur Goral is a territorial species also, but it lives in small groups commonly from 5 to 10 individuals. By contrast Rupicapra and Oreamnos inhabiting such open habitats as alpine meadows may aggregate into herds numbering more than 50 individuals (Schaller 1977).
It is known that species inhabiting forest and open landscapes have the richest repertoire of social signals (Geist 1974), that is, complex habitats lead to complex social behavior. This is supported by comparisons of the behavioural patterns of the Japanese serow, Himalayan goral and Mountain goat argued that (Lovari & Apollonio 1994). The Amur Goral performs all the same behavioural patterns as the Himalayan Goral except for "head down", but also has a different pattern - "stroking of female's head". Its repertoire is richer than serow, but poorer than those of Oreamnos and Rupicapra .
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