Territorial markings

All of the suborder Symphyta and many species in the superfamily Aculeata in the suborder Apocrita are solitary insects. Although not requiring the complex semiochemistry of parasitic or social insects, solitary insects employ pheromones for mating, territorial marking, and host marking. Unfortunately, very few of these have been chemically identified. The pheromones of sawflies and seed wasps were extensively reviewed in 1999 [ 14]. The semiochemicals recently identified in solitary hymenoptera, discussed below, are summarized in Table 2 and Fig. 1. 

Wyatt, T.D. (2003) Pheromones and Animal Behaviour. Cambridge University Press, Cambridge. Yahr, P. (1983) Hormonal influences on territorial marking behavior. In B.B. Svare (Ed.), Hormones and Aggressive Behavior, New York, pp. 145-175. 

Keywords Mammalian semiochemicals Mammalian pheromones Mammalian exocrine secretions Chemical communication Territorial marking... 

The bontebok is a strongly territorial antelope found in the southernmost parts of South Africa, while the blesbok inhabits the arid plains of the central parts of the country. It is practically impossible to translocate these animals even over small distances of a few hundred meters by driving them into unfamiliar areas. In game catching operations, the majority of the driven animals mostly run with their heads down, as though they are more interested in information on the ground than in the source of the threat. This is possibly also the reason why this animal was hunted almost to extinction by the early settlers in South Africa. The territorial behavior of the animal is attributed to territorial mark-... 

The secretion of the red hartebeest is characterized by its high aldehyde content. Of an estimated 100 detectable constituents, 25 are saturated and unsaturated aliphatic aldehydes [138]. Because the aldehydes are highly susceptible to autoxidation, the secretion could therefore only be used for shortterm territorial marking. On the other hand, the conversion of the aldehydes to carboxylic acids could also be transmitting information with a date stamp . In this regard, it is debatable whether fatty acids, which are almost ubiquitous in the animal world, really are such major carriers of semiochemical information in all of the many species in which they are purported to fulfill this role. 

Members of the subfamily Antilopinae, known as dwarf antelope, have well-developed preorbital glands. In both sexes the preorbital gland is a thin-walled pocket anterior to the forward corner of the eye, the secretions of which are used for territorial marking. It would be impossible to present and discuss in detail the enormous volume of chemical information on the preorbital secretions of these animals that has been accumulated over more than 30 years (e.g. [8, 17, 18, 136, 140-143]). The compounds identified in the preorbital secretion of the Cape grysbok, Raphicerus melanotis, are, however, listed in Table 8 [18] to exemplify the compounds that are typically found in these secretions, with one notable exception (see below). 

The overall picture of the secretions of the dwarf antelope seems to suggest that secretions that are produced slowly are more complex. This could be explained in terms of microbiological action, which has more time to contribute to the complexity of a secretion, the slower it is produced. If this is indeed the reason for the complexity of secretions that are produced very slowly, it is possible that, in these animals, with exception of the klipspringer, the long-chain lipid constituents of the secretions could be controlled-release carrier materials rather than semiochemicals. If these heavy compounds were semiochemicals, it could be asked why is it necessary for an animal to spend so much energy to regularly renew its territorial marks. In retrospect, it is possible that up to now too much attention could have been devoted to the heavy constituents of the secretions, while the semiochemically active constituents could have been overlooked because they could be present in such low concentrations that they were not detected by the methods that were employed. 

Salamanders of the ambystomatid family may also use territorial marks. In contrast to red-backed salamanders, which are repelled by conspecific odors or fecal pellets (Jaeger and Gergits, 1979 Jaeger, 1986), spotted salamanders, Amhystoma maculatum, are attracted to paper towels on which conspecifics had lived for 4 days, whether the latter were familiar or unfamiliar (Ducey and Ritsema, 1988). 

Birds do not usually rely on scent for territorial marking. Some species, however, such as burrowing owls, apply odoriferous substances to their nest cavities or burrows. The functions are not well understood. Non-avian predators are thought to be the addressees, placing such marking outside the intraspecific behavior of this chapter. 

Canids a second case study in territorial marking... 

Female mice given a choice between territorial marks of exclusive (one odor) and invaded (mixed odor) territories (in the absence of the owner) chose the first odor. Therefore, females use not only the odor of the territory owner but also the absence or presence of marks by competitors (challengers). The absence of any interfering mark identifies the marking male as one of high quality (Hurst and Rich, 1999). In short, individual A can read the story of interactions between individuals B and C and act accordingly to maximize its own fitness. 

Mykytowycz, R. (1968). Territorial marking by rabbits. Scientific American 218,116-126. 

Thiessen, D.D. and Yahr, P. (1970) Central control of territorial marking in the mongolian gerbil. Physiol Behav 5 275-27. ... 

Wenseleers T., Billen J. and Hefetz A. (2002) Territorial marking in the desert ant Cataglyphis niger does it pay to play Bourgeois J. Insect Behav. 15, 85-93. 

Analysis of hydrocarbons from anal fluids (feces) involved in nest marking by the ant Messor capitatus (Grasso et al., 2005) suggests that territory marking using abdominal secretions issued from the rectal sac may be another labeling function by the way of abdominal secretions. The same authors reported that the hydrocarbons in the feces were similar to those hydrocarbons on the cuticle of workers. 

The complex relationship between certain genera of butterflies belonging to the nymphalid subfamilies Danainae and Ithomiinae, and pyrrolizidine alkaloid containing plants has been discussed. As well as using alkaloids as precursors of pheromones, male butterflies are believed to use pyrrolizidine alkaloids to produce compounds used for territory marking (see above). 

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