Osmoregulation in Crocodilians

Recent crocodilians live primarily in freshwater habitats. Howevertwo species (Crocodylus acutus and C. porosus) are estuarinespecialists; two others (C. niloticus and C. johnstoni) thatare primarily found in fresh water, have estuarine populations.Routes of uptake of water and sodium include drinking, feedingand associated incidental drinking, and integumental and buccaldiffusion. Routes of loss include faeces-cloacal fluid, lingualsalt glands, integumental and buccal diffusion, and respiratoryloss. The least understood route of salt and water exchangeis that of the oral and buccal epithelia, which are much morepermeable to water and sodium than the general integument. Thefreshwater alligator (Alligator mississippiensis) osmoregulatesin a manner typical for an amphibious reptile. Body sodium turnoveris low and the general integument is quite low in permeabilityto sodium. Water turnover is more rapid (in terms of molar exchange)but still relatively low for an aquatic reptile. Most waterexchange occurs across the integument and buccal epithelia.The presence of lingual salt glands in freshwater crocodiliansremains enigmatic, as does the failure of these exocrine glandsin estuarine species to respond to saline loading. Secretiondoes occur after injection of the parasympathetic stimulantmethacholine chloride. The "salt water crocodile" (C. porosus)possesses a suite of osmoregulatory adaptations similar to thosefound in other estuarine reptiles. Water and sodium balanceare maintained primarily by an extremely low general permeabilityto sodium, by economies in water loss, and by excretion of excesssodium by the lingual salt glands. Further work is needed toexamine newly hatched C. porosus, and the possibility of ontogeneticchange in lingual gland function in C. acutus. The importanceof incidental drinking of sea water during feeding (recentlydiscovered in turtles) needs to be evaluated in crocodilians.The use of osmoregulatory data in interpretation of the evolutionaryhistory of the genus Crocodylus needs to be viewed with caution.The hypothesis that all species of Crocodylus originated fromthe transoceanic migration of a saline-tolerant form may notbe the most parsimonious explanation.     

Historical Zoogeography of the Eusuchian Crocodilians: A Physiological Perspective

The historical zoogeography of eusuchian crocodilians has rarelybeen reviewed in any detail and yet is of increasing interestto students of crocodilian biology as large amounts of comparativeinformation on a wide range of species come to hand. Previousinterpretations of crocodilian zoogeography have been basedon one or another of two assumptions–that the major continentalland masses have remained more or less fixed in position, andthat the eusuchians have had only very limited powers of dispersalacross marine barriers. Both of these assumptions are inappropriatein light of our present knowledge of continental drift and crocodilianphysiology. In this paper we attempt a reinterpretation of eusuchian zoogeographybased on new information on their systematic relationships,physiological capacity for marine dispersal, and fossil history.We postulate that anatomical and physiological adaptations toa marine existence have played an important role in eusuchianhistory. We propose that Gavialis and Tomistoma, now restrictedto fresh waters, may have been derived secondarily from ancestorsadapted to salt water. In the case of Tomistoma, similaritiesin lingual gland and buccal cavity anatomy to the true crocodiles(Crocodylus and Osteolaemus) suggest that marine adaptationspredated the divergence of tomistomine and crocodyline stocks.The buccal morphology of Gavialis suggests it also has a marineancestry. Its systematic affinities are uncertain, lying perhapswith Tomistoma or, on other interpretations, with the Mesosuchia.In both cases, the fossil record is not inconsistent with thispossibility. Palaeontological information now available is inadequate toreconstruct the evolutionary history of the Eusuchia in detail.However, saltwater adapted eusuchians are more common in thefossil record than is widely recognized and the likelihood ofdispersal across marine "barriers" by non-alligatorid crocodilianscannot be ignored.     

Behavioral and Physiological Thermoregulation of Crocodilians

Crocodilians, like other reptiles, regulate their body temperaturesby a combination of behavioral and physiological mechanisms.Behaviorally, they seek warm surface water or bask when cooland avoid overheating by the evaporation of water from theirdorsum, evaporation of water by gaping or by retreating to deep,cool water. Physiologically, crocodilians increase cutaneousthermal conductance by increasing blood flow to the skin (andsubdermal musculature) during warming. This hastens the warmingprocess. Cutaneous blood flow is reduced during general coolingand locally if the body temperature exceeds skin temperature.This enables crocodilians to increase body temperature significantlywhile basking in cool shallow water. Large crocodilians appearto be able to alter their rates of heat exchange to a largerextent than small ones and they can do so with less cardiovascularinvolvement. Large crocodilians, with their lower surface/volumeratio, are capable of producing sufficient metabolic heat toelevate their body temperature above water temperature.     

Crocodilians and Their Helminth Parasites: Macroevolutionary Considerations

Crocodilian relationships supported by the phylogenetic relationshipsof digenean and nematode parasites are compared with currentestimates of crocodilian phylogeny. The parasite data support(1) the placement of Gavialis as the sister-group of the alligatoridsand the crocodylids, (2) the monophyly of alligators and caimans,(3) the placement of Caiman (as a monophyletic group) as thesister-group of Melanosuchus plus Paleosuchus, and (4) ancientorigins of Crocodylus consistent with patterns of continentaldrift. The parasite data do not support the monophyly of Crocodylus,but the "misplaced" species (C. palustris and Osteolaemus) havehad few parasites reported from them. There is evidence of widespreadhost-switching, but most of the ambiguity appears to resultfrom uneven representation of parasite groups in host species.This is probably due both to uneven sampling by parasitologistsand to parasite extinctions associated with crocodilian extinctions.     

The Mechanism of Temperature Dependent Sex Determination in Crocodilians: A Hypothesis

Incubation of alligator eggs at 30°C produces 100% females,at 33°C 100% males; temperatures in between produce varyingsex ratios. Wild nests of Alligator mississippiensis show similareffects and the populations are biased towards females. Theincidence and patterns of temperature dependent sex determination(TSD) in other crocodilians are reviewed. Temperature also affectshatchling size and pigmentation patterns, post-hatching growthrates and thermoregulation by juvenile crocodilians. The significanceof temperature sensitive periods defined by temperature shiftexperiments is questioned in relation to a hypothesis to explainthe mechanism of TSD in crocodilians. It is postulated thatthere is an initial sex differentiation mechanism which involvesa quantum period of time and a threshold for a dose of a maledetermining factor. The conditions for induction of males areprecise but exhibit variation between individuals within thepopulation. Females develop by default. The hypothalamus mayhave an important role in a later sex differentiation mechanism.The hypothesis is used to explain the late temperature sensitiveperiods defined by high to low temperature shift experiments,why cooler temperatures are more effective at determining sex,how intermediate temperatures can produce both sexes, the differencesin the pattern between turtles and crocodilians and geographicalsimilarities in the pattern of TSD within crocodilians despitediffering climates. The phylogenetic advantages of TSD in crocodiliansare concerned with the overall reproductive strategy of theanimals. Those crocodilians which are incubated and grow tomaturity under optimal environmental conditions will be bothlarge and male. Larger males are more likely to produce moreoffspring. A review of the effects of the environment on sexdetermination in amphibians and fish suggests that there isa general relationship between size and sex in vertebrates     

Testing Hypotheses of Evolutionary Relationships Among Parasites: The Digeneans of Crocodilians

Parasite systematists may question whether or not all the parasitesof a given host taxon share a common evolutionary history, especiallyone which also includes the host taxon. This study presentsa method for testing hypotheses of such faunal coevolution usingcrocodilians and their digeneans as a model host-parasite system.First, phylogenetic analysis, using numerical methods when largedata bases require such handling, produces estimates of genealogicalrelationships among members of various taxa represented. Theestimates, in the form of branching diagrams (cladograms), caneach be compared with host genealogy and with geographic genealogyfor the areas in which the parasites occur. Congruence of suchpatterns suggests a common evolutionary history and corroboratesan hypothesis of faunal coevolution; incongruence suggests aninvasive or colonizing fauna and falsifies an hypothesis offaunal coevolution. The digeneans of crocodilians apparentlyhave coevolved as a faunal unit with their host group, at leastsince the end of the Mesozoic.     

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.     

湾鳄(Crocodylus porosus)线粒体基因组全序列结构分析与鳄类系统发生

该文测序了湾鳄的线粒体基因组全序列,全长为16,917bp。湾鳄mtDNA结构与其他脊椎动物相似,由22个tRNA,2个rRNA和13个蛋白质编码基因及1个非编码的控制区(D-loop)所组成。除NADH6和tRNAGln、tRNAAla、tRNAAsn、tRNACys、tRNATyr、tRNASer(UCN)、tRNAGlu、tRNAPro在L-链上编码之外,其余基因均在H-链编码。基因排列顺序与已测序的鳄类一致,这显示了鳄类线粒体基因排列顺序上的保守性。但鳄类线粒体基因排列顺序与脊椎动物的典型排列方式相比,有较大的差异,尤其是tRNAPhe基因的重排、tRNASer-tRNAHis-tRNALeu基因族的排列方式等。湾鳄mtDNA和已测序的鳄类一样,缺失轻链复制起始点(OLR)。基于17种鳄mtDNA控制区保守区,采用PAUP4.0最大简约法(Maximumparsimony,MP)构建MP树,邻接法(Neighbor-joiningmethod,NJ)构建NJ树,结果显示:食鱼鳄(Gavialisgangeticus)和假食鱼鳄(Tomistomaschlegelii)聚为一支后再与鳄科(Crocodylidae)的其他物种形成姐妹群,这与基于食鱼鳄和假食鱼鳄的线粒体全序列的分析结果一致,支持将食鱼鳄并入鳄科的观点。结果还支持非洲窄吻鳄(Crocodyluscataphractus)与鳄属(Crocodylus)构成姐妹群,可以单独划分为属的观点。     

几种鳄分子系统发生的探讨

百年来关于扬子鳄的分类学位置存在着很多争议,本测得扬子鳄、暹罗鳄和湾鳄的mtDNA ND4和Cytb基因,并从GenBank中获得密西西比鳄和海龟的DN4基因和Cytb基因相应片段。用Clustal X1.8进行对位排列,以海龟为外群构建分子进化系统树。结果显示,在鳄类动物中,扬子鳄与密西西比鳄的亲缘关系最近,两ND4基因序列碱基差异的20．68％,而Cytb基因序列碱基差异为14．43％,但扬子鳄与密西西相比与鳄的分类问题仍将有待进一步探讨。