Gavialoid crocodylians are the archetypal longirostrine archosaurs and, as such, understanding their patterns of evolution is fundamental to recognizing cranial rearrangements and reconstructing adaptive pathways associated with elongation of the rostrum (longirostry). The living Indian gharial Gavialis gangeticus is the sole survivor of the group, thus providing unique evidence on the distinctive biology of its fossil kin. Yet phylogenetic relationships and evolutionary ecology spanning ~70 million-years of longirostrine crocodylian diversification remain unclear. Analysis of cranial anatomy of a new proto-Amazonian gavialoid, Gryposuchus pachakamue sp. nov., from the Miocene lakes and swamps of the Pebas Mega-Wetland System reveals that acquisition of both widely separated and protruding eyes (telescoped orbits) and riverine ecology within South American and Indian gavialoids is the result of parallel evolution. Phylogenetic and morphometric analyses show that, in association with longirostry, circumorbital bone configuration can evolve rapidly for coping with trends in environmental conditions and may reflect shifts in feeding strategy. Our results support a long-term radiation of the South American forms, with taxa occupying either extreme of the gavialoid morphospace showing preferences for coastal marine versus fluvial environments. The early biogeographic history of South American gavialoids was strongly linked to the northward drainage system connecting proto-Amazonian wetlands to the Caribbean region.
Many species alter skin color to varying degrees and by different mechanisms. Here, we show that some crocodylians modify skin coloration in response to changing light and environmental conditions. Within the Family, Crocodylidae, all members of the genus Crocodylus lightened substantially when transitioned from dark enclosure to white enclosures, whereas Mecistops and Osteolaemus showed little/no change. The two members of the Family Gavialidae showed an opposite response, lightening under darker conditions, while all member of the Family Alligatoridae showed no changes. Observed color changes were rapid and reversible, occurring within 60-90 minutes. The response is visually-mediated and modulated by serum α-melanocyte-stimulating hormone (α-MSH), resulting in redistribution of melanosomes within melanophores. Injection of crocodiles with α-MSH caused the skin to lighten. These results represent a novel description of color change in crocodylians, and have important phylogenetic implications. The data support the inclusion of the Malayan gharial in the Family Gavialidae, and the shift of the African slender-snouted crocodile from the genus Crocodylus to the monophyletic genus Mecistops.
Conflicts between humans and crocodilians are a widespread conservation challenge and the number of crocodile attacks is increasing worldwide. We identified the factors that most effectively decide whether a victim is injured or killed in a crocodile attack by fitting generalized linear models to a 42-year dataset of 87 attacks (27 fatal and 60 non-fatal) by saltwater crocodiles (Crocodylus porosus) in Australia. The models showed that the most influential factors were the difference in body mass between crocodile and victim, and the position of victim in relation to the water at the time of an attack. In-water position (for diving, swimming, and wading) had a higher risk than on-water (boating) or on-land (fishing, and hunting near the water’s edge) positions. In the in-water position a 75 kg person would have a relatively high probability of survival (0.81) if attacked by a 300 cm crocodile, but the probability becomes much lower (0.17) with a 400 cm crocodile. If attacked by a crocodile larger than 450 cm, the survival probability would be extremely low (<0.05) regardless of the victim's size. These results indicate that the main cause of death during a crocodile attack is drowning and larger crocodiles can drag a victim more easily into deeper water. A higher risk associated with a larger crocodile in relation to victim's size is highlighted by children's vulnerability to fatal attacks. Since the first recently recorded fatal attack involving a child in 2006, six out of nine fatal attacks (66.7%) involved children, and the average body size of crocodiles responsible for these fatal attacks was considerably smaller (384 cm, 223 kg) than that of crocodiles that killed adults (450 cm, 324 kg) during the same period (2006-2014). These results suggest that culling programs targeting larger crocodiles may not be an effective management option to improve safety for children.
The major histocompatibility complex (MHC) is a dynamic genome region with an essential role in the adaptive immunity of vertebrates, especially antigen presentation. The MHC is generally divided into subregions (classes I, II and III) containing genes of similar function across species, but with different gene number and organisation. Crocodylia (crocodilians) are widely distributed and represent an evolutionary distinct group among higher vertebrates, but the genomic organisation of MHC within this lineage has been largely unexplored. Here, we studied the MHC region of the saltwater crocodile (Crocodylus porosus) and compared it with that of other taxa. We characterised genomic clusters encompassing MHC class I and class II genes in the saltwater crocodile based on sequencing of bacterial artificial chromosomes. Six gene clusters spanning ∼452 kb were identified to contain nine MHC class I genes, six MHC class II genes, three TAP genes, and a TRIM gene. These MHC class I and class II genes were in separate scaffold regions and were greater in length (2-6 times longer) than their counterparts in well-studied fowl B loci, suggesting that the compaction of avian MHC occurred after the crocodilian-avian split. Comparative analyses between the saltwater crocodile MHC and that from the alligator and gharial showed large syntenic areas (>80% identity) with similar gene order. Comparisons with other vertebrates showed that the saltwater crocodile had MHC class I genes located along with TAP, consistent with birds studied. Linkage between MHC class I and TRIM39 observed in the saltwater crocodile resembled MHC in eutherians compared, but absent in avian MHC, suggesting that the saltwater crocodile MHC appears to have gene organisation intermediate between these two lineages. These observations suggest that the structure of the saltwater crocodile MHC, and other crocodilians, can help determine the MHC that was present in the ancestors of archosaurs.
Crocodiles and their kin (Crocodylidae) use asymmetrical (bounding and galloping) gaits when moving rapidly. Despite being morphologically and ecologically similar, it seems alligators and their kin (Alligatoridae) do not. To investigate a possible anatomical basis for this apparent major difference in locomotor capabilities, we measured relative masses and internal architecture (fascicle lengths and physiological cross-sectional areas) of muscles of the pectoral and pelvic limbs of 40 individuals from six representative species of Crocodylidae and Alligatoridae. We found that, relative to body mass, Crocodylidae have significantly longer muscle fascicles (increased working range), particularly in the pectoral limb, and generally smaller muscle physiological cross-sectional areas (decreased force-exerting capability) than Alligatoridae. We therefore hypothesise that the ability of some crocodylians to use asymmetrical gaits may be limited more by the ability to make large, rapid limb motions (especially in the pectoral limb) than the ability to exert large limb forces. Furthermore, analysis of scaling patterns in muscle properties shows that limb anatomy in the two clades becomes more divergent during ontogeny. Limb muscle masses, fascicle lengths and physiological cross-sectional areas scale with significantly larger coefficients in Crocodylidae than Alligatoridae. This combination of factors suggests that inter-clade disparity in maximal limb power is highest in adult animals. Therefore, despite their apparent morphological similarities, both mean values and scaling patterns suggest that considerable diversity exists in the locomotor apparatus of extant Crocodylia.
OBJECTIVES: To investigate the character of immobilization given by alfaxalone in juvenile crocodiles at optimal and at suboptimal temperatures. STUDY DESIGN: Prospective, randomized partial crossover study. ANIMALS: Twenty captive male estuarine (weight 0.6-2.5 kg) and five captive male freshwater crocodiles (weight 0.2-0.6 kg). METHODS: Crocodiles were acclimatized for 24 hours at one of the following environmental temperatures; 32 °C, 27 °C, 22 °C or 17 °C, then received 3 mg kg-1 intravenous (IV) alfaxalone into the dorsal occipital venous sinus. Duration and quality of immobilization was assessed and heart rate (HR) measured. On a separate occasion each crocodile was immobilized at one other environmental temperature. RESULTS: Alfaxalone, 3 mg kg-1 IV, produced immobilization for 55 (range 15-100 minutes in estuarine, and 20 (range 20-25) minutes in freshwater crocodiles at 32 °C. There was no significant difference overall in immobilization times between temperatures, other than that, in estuarine crocodiles, duration was shorter at 32 °C than 22 °C. The character of immobilization was unpredictable, with animals recovering without warning, or having extended recoveries requiring assisted ventilation. Assisted ventilation was necessary mainly at the lower temperatures. Median HR in all temperature treatments decreased within 5 minutes post-injection, but the change in HR over the duration of immobilization was affected by the temperature, with a progressively smaller range of fall as temperature decreased. At 17 °C, two estuarine crocodiles appeared to re-immobilize after initial recovery, became severely bradycardiac and required ventilation and re-warming. CONCLUSIONS AND CLINICAL RELEVANCE: Alfaxalone IV in small captive estuarine and freshwater crocodiles provides adequate induction of immobilization at various temperatures. However, the unpredictable results following induction mean it is unsuitable for field use and should be restricted to environments where intubation and ventilation are available, where animals can be warmed to optimal temperature, and where access to immersion in water can be restricted for 24 hours.
OBJECTIVE: To determine the efficacy of medetomidine for immobilisation of captive juvenile crocodiles over a range of temperatures, and its reversibility with atipamezole. STUDY DESIGN: Prospective experimental study. ANIMALS: Forty male estuarine crocodiles (body weight 2.0 to 4.8 kg). METHODS: Each crocodile was randomly assigned to one of four temperature groups: Group 1:32 °C; Group 2:27 °C; Group 3:22 °C; and Group 4:17 °C (n = 10 for each group). Medetomidine (0.5 mg kg(-1) ) was administered intramuscularly (IM) into the thoracic limb of all crocodiles. After 50 minutes, all animals from each group received 2.5 mg kg(-1) atipamezole IM in the opposite thoracic limb and time to recovery was documented. Heart and respiratory rates and the degree of immobilisation were monitored every 5 minutes until recovery, and behaviour monitored for 7 subsequent days. RESULTS: Onset of immobilisation occurred at 15 ± 10 minutes in Group 1, and at 30 ± 10 minutes in Groups 2 and 3. In Group 4, animals were not immobilised. Recovery following atipamezole was 10 ± 5 minutes at all temperatures. One-way analysis of variance (anova) demonstrated a significant difference in induction times between groups (p < 0.01) but not in recovery times following atipamezole administration (p < 0.25). Heart and respiratory rates decreased markedly following medetomidine administration and increased markedly following atipamezole reversal. CONCLUSIONS AND CLINICAL RELEVANCE: Medetomidine administered in the thoracic limb of juvenile captive estuarine crocodiles provides profound sedation or immobilisation at temperatures of 22 °C and above. Atipamezole administered in the contralateral thoracic limb results in consistent reversal of the effects of medetomidine and a return to normal behaviour within 15-20 minutes regardless of temperature. Even though immobilisation is not induced at 17 °C, profound reversible sedation does occur reliably and repeatably.
Caiman crocodilus apaporiensis has been considered by several authors as an extreme of morphological variation within the Caiman crocodilus complex. Here, we evaluate its position in the Caiman crocodilus complex morphospace using morphological traits from head shape. We examined the holotype and seventeen paratypes of Caiman crocodilus apaporiensis Medem 1955 deposited at the Field Museum of Natural History. We performed multivariate morphometric analyses: principal component analysis (PCA) and discriminant function analysis (DFA), based on 21 cranial traits of of C. c. apaporiensis, C. yacare and the C. crocodilus complex (C. c. chiapasius, C. c. fuscus andC. c. crocodilus). We find a notable separation of C.c. apaporiensis from C. yacare and C. crocodilus complex in the morphospace. We suggest that geographic isolation might have driven this morphological separation from the C. crocodilus complex, but further analysis are necessary to confirm whether these differences are related with genetic differentiation within the complex. In addition, we suggest that environmental heterogeneity might drive the evolution of independent lineages within the C. crocodilus complex.
A number of extant and extinct archosaurs evolved an elongate, narrow rostrum. This longirostrine condition has been associated with a diet comprising a higher proportion of fish and smaller prey items compared to taxa with broader, more robust snouts. The evolution of longirostrine morphology and a bulbous anterior rosette of premaxillary teeth also occurs in the spinosaurid theropod dinosaurs, leading to suggestions that at least some members of this clade also had a diet comprising a notable proportion of fish or other small vertebrates. Here we compare the rostral biomechanics of the spinosaurs Baryonyx walkeri and Spinosaurus c.f. S. aegyptiacus to three extant crocodilians: two longistrine taxa, the African slender-snouted crocodile Mecistops cataphractus and the Indian gharial Gavialis gangeticus; and the American alligator Alligator mississippiensis. Using computed tomography (CT) data, the second moments of area and moments of inertia at successive transverse slices along the rostrum were calculated for each of the species. Size-independent results tested the biomechanical benefits of material distribution within the rostra. The two spinosaur rostra were both digitally reconstructed from CT data and compared against all three crocodilians. Results show that African slender-snouted crocodile skulls are more resistant to bending than an equivalent sized gharial. The alligator has the highest resistances to bending and torsion of the crocodiles for its size and greater than that of the spinosaurs. The spinosaur rostra possess similar resistance to bending and torsion despite their different morphologies. When size is accounted for, B. walkeri performs mechanically differently from the gharial, contradicting previous studies whereas Spinosaurus does not. Biomechanical data support known feeding ecology for both African slender-snouted crocodile and alligator, and suggest that the spinosaurs were not obligate piscivores with diet being determined by individual animal size.
During their evolution towards a complete life cycle on land, stem reptiles developed both an impermeable multi-layered keratinized epidermis and skin appendages (scales) providing mechanical, thermal, and chemical protection. Previous studies have demonstrated that, despite the presence of a particularly armored skin, crocodylians have exquisite mechanosensory abilities thanks to the presence of small integumentary sensory organs (ISOs) distributed on postcranial and/or cranial scales.