section_path: Territorial behaviour
section_name: Territorial behaviour
paragraph_text: Territorial behaviour
  
  Little is known of the territorial behaviour of caecilians, but some frogs and salamanders defend home ranges. These are usually feeding, breeding or sheltering sites. Males normally exhibit such behaviour though in some species, females and even juveniles are also involved. Although in many frog species, females are larger than males, this is not the case in most species where males are actively involved in territorial defence. Some of these have specific adaptations such as enlarged teeth for biting or spines on the chest, arms or thumbs.
  
  In salamanders, defence of a territory involves adopting an aggressive posture and if necessary attacking the intruder. This may involve snapping, chasing and sometimes biting, occasionally causing the loss of a tail. The behaviour of red back salamanders (Plethodon cinereus) has been much studied. 91% of marked individuals that were later recaptured were within a metre (yard) of their original daytime retreat under a log or rock. A similar proportion, when moved experimentally a distance of 30 metres (98 ft), found their way back to their home base. The salamanders left odour marks around their territories which averaged 0.16 to 0.33 square metres (1.7 to 3.6 sq ft) in size and were sometimes inhabited by a male and female pair. These deterred the intrusion of others and delineated the boundaries between neighbouring areas. Much of their behaviour seemed stereotyped and did not involve any actual contact between individuals. An aggressive posture involved raising the body off the ground and glaring at the opponent who often turned away submissively. If the intruder persisted, a biting lunge was usually launched at either the tail region or the naso-labial grooves. Damage to either of these areas can reduce the fitness of the rival, either because of the need to regenerate tissue or because it impairs its ability to detect food.
  
  In frogs, male territorial behaviour is often observed at breeding locations; calling is both an announcement of ownership of part of this resource and an advertisement call to potential mates. In general, a deeper voice represents a heavier and more powerful individual, and this may be sufficient to prevent intrusion by smaller males. Much energy is used in the vocalization and it takes a toll on the territory holder who may be displaced by a fitter rival if he tires. There is a tendency for males to tolerate the holders of neighbouring territories while vigorously attacking unknown intruders. Holders of territories have a "home advantage" and usually come off better in an encounter between two similar-sized frogs. If threats are insufficient, chest to chest tussles may take place. Fighting methods include pushing and shoving, deflating the opponent's vocal sac, seizing him by the head, jumping on his back, biting, chasing, splashing, and ducking him under the water.