Lígia Pizzatto

ESTIMATING SURVIVAL RATES OF UNCATCHABLE ANIMALS: THE MYTH OF HIGH JUVENILE MORTALITY IN REPTILES

Survival rates of juvenile reptiles are critical population parameters but are difficult to obtain through mark-recapture programs because these small, secretive animals are rarely caught. This scarcity has encouraged speculation that survival rates of juveniles are very low, and we test this prediction by estimating juvenile survival rates indirectly. A simple mathematical model calculates the annual juvenile survival rate needed to maintain a stable population size, using published data on adult survival rates, reproductive output, and ages at maturity in 109 reptile populations encompassing 57 species. Counter to prediction, estimated juvenile survival rates were relatively high (on average, only about 13% less than those of conspecific adults) and highly correlated with adult survival rates. Overall, survival rates during both juvenile and adult life were higher in turtles than in snakes, and higher in snakes than in lizards. As predicted from life history theory, rates of juvenile survival were higher in species that produce large offspring, and higher in viviparous squamates than in oviparous species. Our analyses challenge the widely held belief that juvenile reptiles have low rates of annual survival and suggest instead that sampling problems and the elusive biology of juvenile reptiles have misled researchers in this respect.

LIFE-HISTORY ADAPTATIONS TO ARBOREALITY IN SNAKES

If selective forces on locomotor ability and reproductive biology differ among habitats, we expect to see relationships between habitat, morphology, and life-history traits. Comparative (phylogenetically based) analysis of data from 12 pythonid and 12 boid snake species reveals multiple evolutionary shifts in habitat use, notably in the evolution of arboreal habits. Compared to terrestrial and aquatic taxa of the same overall body size, arboreal species have narrower and more laterally compressed bodies and relatively longer tails. Offspring sizes are not affected by arboreality, but presumably reflecting space constraints within their narrow bodies, arboreal species (1) produce smaller clutch sizes relative to maternal body length and (2) have left and right ovaries that overlap little if at all along the length of the body (i.e., the right ovary is positioned anterior to the left ovary) whereas in terrestrial snakes the two ovaries overlap along much of their length. This modification of ovarian morphology in arboreal snakes presumably reduces the degree of bodily distension during vitellogenesis and pregnancy, thus enhancing climbing ability and camouflage among the branches.

Reproductive biology of the false coral snake Oxyrhopus guibei (Colubridae) from southeastern Brazil

The reproductive biology of the false coral snake, Oxyrhopus guibei , was studied through dissection of 496 specimens, combined with observations on captive individuals. Males mature with smaller body size than females, females attain much larger body size, and male-male combat is not expected. Clutch size ranged from 3 to 20, and was correlated with female length. Reproductive cycles in both males and females seem to be continuous, with vitellogenesis and spermatogenesis occurring throughout the year. Reproductive activity in both sexes decreased at the end of the rainy season possibly due to previous intense reproductive activity in more favorable climatic conditions. The smaller number of individuals collected at the end of the rainy season apparently occurs due to the decrease of reproductive activity of this snake.

Infection dynamics of the lungworm Rhabdias pseudosphaerocephala in its natural host, the cane toad (Bufo marinus), and in novel hosts (native Australian frogs).

Host-parasite systems have often evolved over time, such that infection dynamics may become greatly modified from the time of initial contact of the host with the parasite. Biological invasions may be useful to clarify processes in the initial contact of hosts with parasites, and allow us to compare parasite uptake between the ancestral (coevolved) host and novel (noncoevolved) hosts. Cane toads (Bufo marinus) are spreading rapidly through tropical Australia, carrying with them a nematode lungworm (Rhabdias pseudosphaerocephala) congeneric with those found in Australian frogs. We investigated the dynamics of infections of the toad parasite by conducting histologic examinations of cane toads and three native Australian frogs (Litoria dahlii, Litoria nasuta, and Opisthodon ornatus) at 2, 6, and 10 days after experimental exposure to the toad lungworm. More worms were found in toads than in frogs, especially at longer periods postexposure. In toads, the infective larvae entered the skin and muscles within 2 days postexposure, passed into the coelom in 6 days, and reached the lungs at 10 days. In frogs, larvae were found in many organs rather than migrating to consistent target tissues; a few larvae reached the lungs of L. dahlii. Migratory larvae caused increasing inflammation (primarily granulomatous admixed with granulocytes then lymphocytes) through time, especially in frogs. Evolution has resulted in an enhanced ability of the lungworm to locate the target organ (the lungs) of the toad, and an increase in rates of parasite survival within this host.

Overview of Reproductive Strategies in Xenodontini (Serpentes: Colubridae: Xenodontinae) with New Data for Xenodon Neuwiedii and Waglerophis Merremii

Abstract Reproductive patterns are highly variable among Neotropical colubrids. Snakes of the Tribe Xenodontini are widely distributed in South America and show continuous reproductive cycles in many areas. We report interspecific and intraspecific variation in reproductive traits among these snakes and also show that seasonal cycles occur mainly in cooler areas. Clutch size relative to body size is similar among species, but Erythrolamprus spp. seem to lay fewer eggs than other species, and Waglerophis merremii from southeastern and southern Brazil lay more eggs than other species. Newborn Erythrolamprus are larger than all other Xenodontini, which may be related to ophiophagy. As in many other snakes, adult females are larger than males, but both sexes attain maturity with a proportionally similar body size. The sexual size dimorphism index is generally lower in smaller bodied species, and combat may be absent in the tribe. Male reproductive cycles are less well studied but seem to be more conservative, being continuous in all species studied (W. merremii, Xenodon neuwiedii, and Liophis miliaris). Phylogeny has an important role in reproductive patterns but climate and life-history traits can also influence tropical and subptropical species.

Reproductive Ecology of Dipsadine Snakes, With Emphasis on South American Species

A relatively large amount of variation occurs in the reproductive ecology of tropical snakes, and this variation is generally regarded as being a consequence of seasonality in climate and prey availability. In some groups, even closely related species may differ in their reproductive ecology; however, in others it seems to be very conservative. Here we explore whether characters related to reproduction are phylogenetically constrained in a monophyletic group of snakes, the subfamily Dipsadinae, which ranges from Mexico to southern South America. We provide original data on reproduction for Leptodeira annulata, Imantodes cenchoa, and three species of Sibynomorphus from southern, southeastern and central Brazil, and data from literature for other species and populations of dipsadines. Follicular cycles were seasonal in Atractus reticulatus, Dipsas albifrons, Hypsiglena torquata, Leptodeira maculata, L. punctata, Sibynomorphus spp. and Sibon sanniola from areas where climate is seasonal. In contrast, extended or continuous follicular cycles were recorded in Dipsas catesbyi, D. neivai, Imantodes cenchoa, Leptodeira annulata, and Ninia maculata from areas with seasonal and aseasonal climates. Testicular cycles also varied from seasonal (in H. torquata) to continuous (in Dipsas spp., Leptodeira annulata, L. maculata, N. maculata and Sibynomorphus spp.). Most dipsadines are small (less than 500 mm SVL), and females attain sexual maturity with similar relative body size than males. Sexual dimorphism occurred in terms of SVL and tail length in most species, and clutch size tended to be small (less than five eggs). Combat behavior occurs in Imantodes cenchoa, which did not show sexual size dimorphism. Reproductive timing, for both females and males, varied among species but in general there were no differences between the tribes of Dipsadinae in most of the reproductive characteristics, such as mean body size, relative size at sexual maturity, sexual size and tail dimorphism, duration of vitellogenesis or egg-carrying in oviducts.

Food habits of Brazilian boid snakes: overview and new data, with special reference to Corallus hortulanus

This study examines the diet of eight boid snakes: Boa c. constrictor, Boa c. amarali, Corallus caninus, C. hortulanus, Epicrates cenchria, E. crassus, E. assisi and Eunectes murinus mainly by analyzing the gut contents of preserved museum specimens, and includes a literature review to present an overview of the diet of Brazilian boids. Mammals constitute the primary prey consumed by the majority of the species. Birds are also frequently consumed by C. hortulanus and Boa contrictor, and are the most important prey for B. c. amarali. Ectotherms (mostly lizards) were only consumed by immature snakes. Such prey is rarely consumed by B. c. amarali and not recorded for Epicrates and Eunectes species in our dissections. C. caninus is likely a mammal specialist and Epicrates prey on birds more opportunistically. The niche overlap index varied from 0.27-0.52 for species occurring in the same bioma and geographic range but it is possible this overlap is lower as most sympatric species explore different macrohabitat. C. hortulanus exhibited a significant relationship between prey size and predator head size; this relationship did not differ among mature and immature snakes. In comparison to immature individuals heavier adult snakes fed on heavier prey items however, the ratio between prey/predator mass decreased with increase in predator mass (or size). Most boids exploit diurnal and nocturnal preys, probably using both sit-and-wait and active tactics. They feed on the ground but boas and C. hortulanus and possibly E. cenchria also exploit arboreal prey.

The effects of experimentally infecting Australian tree frogs with lungworms (Rhabdias pseudosphaerocephala) from invasive cane toads

Invasive species may transmit novel pathogens to native taxa, and lacking a history of coevolutionary interactions with the pathogen, the new hosts may be severely affected. Cane toads (Rhinella marina) were introduced to Australia in 1935, bringing with them a lungworm (Rhabdias pseudosphaerocephala) not found in Australian frogs. Previous studies suggest that most frog species are unaffected by this parasite, but one tree-frog (Litoria caerulea) can harbour high numbers of lungworm. More detailed laboratory studies confirm and extend the earlier results on L. caerulea and show that Rhabdias infection severely depresses the viability of metamorphs of an allied tree-frog species, Litoria splendida. Parasitic larvae infected both of these two closely related tree-frog species, but the two anurans differed in the consequences of infection. Parasitism reduced the survivorship of L. splendida and the stamina of both species. Lungworms did not consistently reduce growth rates or affect heart rates in either tree-frog species. Although L. splendida is potentially vulnerable to the arrival of toad-transported lungworms, rates of host-switching may be reduced by low levels of habitat overlap between the frogs (which are rock-dwelling and arboreal) and the toads (which are terrestrial and most abundant in disturbed habitats).

You Are What You Eat: Parasite Transfer in Cannibalistic Cane Toads

Abstract Pathogen transfer may be an important but poorly understood cost of cannibalism. Does the consumption of smaller conspecifics by Cane Toads (Rhinella marina) result in transfer of viability-reducing parasites such as nematode lungworms (Rhabdias pseudosphaerocephala)? Our experimental trials confirm that cannibalistic toads can indeed become infected, and our results are probably the first evidence of macroparasite transmission via intraspecific predation in amphibians. Our results also show that parasites acquired via cannibalism are viable, develop into fertile adults, and reduce the locomotor performance of the hosts. How cannibalism contributes to nematode transmission and spread in natural populations is not known, but we propose a scenario in which this interaction would be likely to increase the lungworm prevalence, intensity, or persistence.

Insights into the adaptive significance of vertical pupil shape in snakes

Pupil shape in vertebrates ranges from circular to vertical, with multiple phylogenetic shifts in this trait. Our analyses challenge the widely held view that the vertical pupil evolved as an adaptation to enhance night vision. On functional grounds, a variable‐aperture vertical pupil (i) allows a nocturnal species to have a sensitive retina for night vision but avoid dazzle by day by adjusting pupil closure, and (ii) increases visual acuity by day, because a narrow vertical pupil can project a sharper image onto the retina in the horizontal plane. Detection of horizontal movement may be critical for predators that wait in ambush for moving prey, suggesting that foraging mode (ambush predation) as well as polyphasic activity may favour the evolution of vertical pupil shape. Camouflage (disruption of the circular outline of the eye) also may be beneficial for ambush predators. A comparative analysis in snakes reveals significant functional links between pupil shape and foraging mode, as well as between pupil shape and diel timing of activity. Similar associations between ambush predation and vertically slit pupils occur in lizards and mammals also, suggesting that foraging mode has exerted major selective forces on visual systems in vertebrates.