Matthew L. Brien

Born to be bad: Agonistic behaviour in hatchling saltwater crocodiles (Crocodylus porosus)

Detailed observations on groups of captive saltwater crocodile (Crocodylus porosus) hatchlings revealed sporadic periods of intense agonistic interactions, with 16 highly distinctive behaviours, in the morning (06:00–08:00 h) and evening (17:00–20:00 h) in shallow water. Ontogenetic changes in agonistic behaviour were quantified by examining 18 different groups of hatchlings, six groups each at 1 week, 13 weeks and 40 weeks after hatching. Agonistic interactions between hatchlings at 1 week of age (mean 7.3 ± 0.65/night) were not well-defined and varied in intensity (low, medium, high), number of individuals that were aggressive, and the outcome, while most interactions involved contact (94.5%). There were also clutch specific differences in the frequency of agonistic interactions. At 13 and 40 weeks, a more hierarchal dominance relationship appeared to be established which primarily involved aggression–submission interactions. Agonistic interactions were more frequent (13 weeks 9.7 ± 0.61/night; 40 weeks 22.2 ± 0.61/night) and intense (medium, high), but shorter in duration, in which the subordinate individual fled in response to an approach by a dominant animal that often gave chase but did not make contact. While the full repertoire of behaviour was displayed by hatchlings at 1 week of age, a smaller subset based on dominance status was displayed among 13- and 40-week-old hatchlings. Agonistic behaviour occurs in C. porosus shortly after hatching and is important in establishing and maintaining dominance hierarchies that are characterised by aggression–submission interactions. This type of interaction appears typical for C. porosus both in the wild and in captivity, and may be important in preventing serious injury in a species equipped with formidable armoury. Dispersal by hatchling C. porosus at around 13 weeks of age appears to be driven by a growing intolerance of conspecifics, while territoriality is apparent at an early age. Consequently, agonistic behaviour and social status may be major contributors to the observed differences in growth rates and survival in captivity.

The good, the bad, and the ugly: agonistic behaviour in juvenile crocodilians.

We examined agonistic behaviour in seven species of hatchling and juvenile crocodilians held in small groups (N = 4) under similar laboratory conditions. Agonistic interactions occurred in all seven species, typically involved two individuals, were short in duration (5–15 seconds), and occurred between 1600–2200 h in open water. The nature and extent of agonistic interactions, the behaviours displayed, and the level of conspecific tolerance varied among species. Discrete postures, non-contact and contact movements are described. Three of these were species-specific: push downs by C. johnstoni; inflated tail sweeping by C. novaeguineae; and, side head striking combined with tail wagging by C. porosus. The two long-snouted species (C. johnstoni and G. gangeticus) avoided contact involving the head and often raised the head up out of the way during agonistic interactions. Several behaviours not associated with aggression are also described, including snout rubbing, raising the head up high while at rest, and the use of vocalizations. The two most aggressive species (C. porosus, C. novaeguineae) appeared to form dominance hierarchies, whereas the less aggressive species did not. Interspecific differences in agonistic behaviour may reflect evolutionary divergence associated with morphology, ecology, general life history and responses to interspecific conflict in areas where multiple species have co-existed. Understanding species-specific traits in agonistic behaviour and social tolerance has implications for the controlled raising of different species of hatchlings for conservation, management or production purposes.

Intra- and interspecific agonistic behaviour in hatchling Australian freshwater crocodiles (Crocodylus johnstoni) and saltwater crocodiles (Crocodylus porosus)

Abstract. We examined agonistic behaviour in hatchling Australian freshwater crocodiles (Crocodylus johnstoni) at 2 weeks, 13 weeks, and 50 weeks after hatching, and between C. johnstoni and saltwater crocodiles (Crocodylus porosus) at 40–50 weeks of age. Among C. johnstoni, agonistic interactions (15–23 s duration) were well established by two weeks old and typically involved two and occasionally three individuals, mostly between 17 : 00 and 24 : 00 hours in open-water areas of enclosures. A range of discrete postures, non-contact and contact movements are described. The head is rarely targeted in contact movements with C. johnstoni because they exhibit a unique ‘head raised high’ posture, and engage in ‘push downs’. In contrast with C. porosus of a similar age, agonistic interactions between C. johnstoni were conducted with relatively low intensity and showed limited ontogenetic change; there was also no evidence of a dominance hierarchy among hatchlings by 50 weeks of age, when the frequency of agonistic interactions was lowest. Agonistic interactions between C. johnstoni and C. porosus at 40–50 weeks of age were mostly low level, with no real exclusion or dominance observed. However, smaller individuals of both species moved slowly out of the way when a larger individual of either species approached. When medium- or high-level interspecific interactions did occur, it was between similar-sized individuals, and each displayed species-specific behaviours that appeared difficult for contestants to interpret: there was no clear winner or loser. The nature of agonistic interactions between the two species suggests that dominance may be governed more strongly by size rather than by species-specific aggressiveness.

The relationship between early growth and survival of hatchling saltwater crocodiles (Crocodylus porosus) in captivity

Hatchling fitness in crocodilians is affected by “runtism” or failure to thrive syndrome (FTT) in captivity. In this study, 300 hatchling C. porosus, artificially incubated at 32°C for most of their embryonic development, were raised in semi-controlled conditions, with growth criteria derived for the early detection of FTT (within 24 days). Body mass, four days after hatching (BM4d), was correlated with egg size and was highly clutch specific, while snout-vent length (SVL4d) was much more variable within and between clutches. For the majority of hatchlings growth trajectories within the first 24 days continued to 90 days and could be used to predict FTT affliction up to 300 days, highlighting the importance of early growth. Growth and survival of hatchling C. porosus in captivity was not influenced by initial size (BM4d), with a slight tendency for smaller hatchlings to grow faster in the immediate post-hatching period. Strong clutch effects (12 clutches) on affliction with FTT were apparent, but could not be explained by measured clutch variables or other factors. Among individuals not afflicted by FTT (N = 245), mean growth was highly clutch specific, and the variation could be explained by an interaction between clutch and season. FTT affliction was 2.5 times higher among clutches (N = 7) that hatched later in the year when mean minimum air temperatures were lower, compared with those clutches (N = 5) that hatched early in the year. The results of this study highlight the importance of early growth in hatchling C. porosus, which has implications for the captive management of this species.

Energetic costs of digestion in Australian crocodiles

We measured standard metabolic rate (SMR) and the metabolic response to feeding in the Australian crocodiles, Crocodylus porosus and C. johnsoni. Both species exhibit a response that is characterised by rapidly increasing metabolism that peaks within 24 h of feeding, a postfeeding metabolic peak (peak O2) of 1.4–2.0 times SMR, and a return to baseline metabolism within 3–4 days after feeding. Postfeeding metabolism does not significantly differ between species, and crocodiles fed intact meals have higher total digestive costs (specific dynamic action; SDA) than those fed homogenised meals. Across a more than 100-fold range of body size (0.190 to 25.96 kg body mass), SMR, peak O2, and SDA all scale with body mass to an exponent of 0.85. Hatchling (≤1 year old) C. porosus have unexpectedly high rates of resting metabolism, and this likely reflects the substantial energetic demands that accompany the rapid growth of young crocodilians.

Ontogenetic comparisons of standard metabolism in three species of crocodilians

Due in part to their large size, aggressive temperament, and difficulty in handling, there are few physiological studies of adult crocodilians in the literature. As a result, studies comparing individuals across an ontogenetic series and comparisons among species are also lacking. We addressed this gap in knowledge by measuring standard metabolic rates (SMR) of three species of crocodilians (Crocodylus porosus, C. johnsoni, and Alligator mississippiensis), and included individuals that ranged from 0.22 to 114 kg. Allometric scaling of SMR with body mass was similar among the species, but C. porosus had significantly higher SMR than did C. johnsoni or A. mississippiensis. Differences in SMR among species are potentially related to behavioural differences in levels of aggression; C. porosus are the most aggressive of the crocodilians measured, and have rates of standard metabolism that are approximately 36% higher at the grand mean body size than those measured for C. johnsoni or A. mississippiensis, which are among the least aggressive crocodilians.