J. Sean Doody

Communal egg-laying in reptiles and amphibians: evolutionary patterns and hypotheses

Communal egg-laying is widespread among animals, occurring in insects, mollusks, fish, amphibians, reptiles, and birds, just to name a few. While some benefits of communal egg-laying may be pervasive (e.g., it saves time and energy and may ensure the survival of mothers and their offspring), the remarkable diversity in the life histories of the animals that exhibit this behavior presents a great challenge to discovering any general explanation. Reptiles and amphibians offer ideal systems for investigating communal egg-laying because they generally lack parental carea simplification that brings nest site choice behavior into sharp focus. We exhaustively reviewed the published literature for data on communal egg-laying in reptiles and amphibians. Our analysis demonstrates that the behavior is much more common than previously recognized (occurring in 481 spp.), especially among lizards (N = 255 spp.), where the behavior has evolved multiple times. Our conceptual review strongly suggests that different forces may be driving the evolution and maintenance of communal egg-laying in different taxa. Using a game theory approach, we demonstrate how a stable equilibrium may occur between solitary and communal layers, thus allowing both strategies to co-exist in some populations, and we discuss factors that may influence these proportions. We conclude by outlining future research directions for determining the proximate and ultimate causes of communal egg-laying.

Invasive species control: understanding conflicts between researchers and the general community

Understanding the reasons for disagreements about conservation issues can facilitate effective engagement between the people involved. Invasive species are often central to such debates, with researchers and members of the public frequently disagreeing about the nature and magnitude of problems posed by the invaders, and the best ways to deal with them. The spread of non-native cane toads (Rhinella marina) throughout Australia has stimulated research on toad impact and control, and has mobilized local communities to reduce cane toad numbers through direct action. Biologists and community groups have disagreed about many toad-related topics, providing an instructive case history about impediments to consensus. Debates about the ecological impacts of cane toads mostly reflect poor communication of available research results (ie scientists have been largely unsuccessful in transmitting their findings to community groups), whereas disagreements about toad control reflect an information vacuum about the effect...

Indirect facilitation of a native mesopredator by an invasive species: are cane toads re-shaping tropical riparian communities?

Top predators can suppress mesopredators both by killing them and by motivating changes in their behavior, and there are numerous examples of mesopredator release caused by declines in top predator populations. Demonstrated cases of invasive species triggering such releases among vertebrate trophic linkages (indirect facilitation), however, are rare. The invasive cane toad, Bufo marinus, has caused severe population-level declines in some Australian predators via lethal toxic ingestion. During a long-term study of the direct impacts of cane toads on predatory monitor lizards in tropical Australia, we documented significant, marked increases in annual counts of a mesopredator, the common tree snake (Dendrelaphis punctulatus). Mean snake counts during surveys of 70-km river transects at two sites increased from <1 individual per survey during 2001–2006, to 8–18 per survey in 2007. These increases occurred approximately 3xa0years following the arrival of cane toads, and 1–3xa0years after 71–96xa0% population declines in three species of predatory monitor lizards (Varanus panoptes, V. mertensi, and V. mitchelli). These data suggest a mesopredator release: the dramatic reduction of predatory monitor lizards caused increases in the tree snake by decreasing predation risk. The increases in tree snake counts were not attributable to either abiotic factors, or a trophic subsidy. The severe declines of predatory monitor lizards, coupled with recent evidence of cascading effects on their prey, suggest that cane toads are re-shaping riparian communities in tropical Australia through both direct negative effects and indirect facilitation.

SPERM STORAGE IN SPERMATHECAE OF THE GREAT LAMPER EEL, AMPHIUMA TRIDACTYLUM (CAUDATA: AMPHIUMIDAE)

The spermathecae of ten female Amphiuma tridactylum were examined by light and electron microscopy during the presumed mating and ovipository seasons (March–August) in Louisiana. Spermathecae were simple tubuloalveolar glands in the dorsal wall of the cloaca. Six of the ten specimens were vitellogenic, and all of these specimens contained sperm in their spermathecae and had secretory activity in the spermathecal epithelium. Two nonvitellogenic females also had sperm in their spermathecae and active epithelial cells, whereas the other nonvitellogenic females lacked stored sperm and secretory activity in the spermathecae. In specimens storing sperm from March–May, the sperm were normal in cytology, and secretory vacuoles were contained within the epithelium. In the August sample, however, evidence of sperm degradation was present, and secretory material had been released into the lumen by an apocrine process. We therefore hypothesize that the spermathecal secretions function in sperm degeneration.

Cryptic and Complex Nesting in the Yellow-Spotted Monitor, Varanus panoptes

Abstract Despite the general importance of nest site choice in reproductive success in taxa with little or no parental care, little is known for reptiles other than turtles. Here we report on the nesting ecology of the Yellow-Spotted Monitor, Varanus panoptes, a large tropical lizard that utilizes warrens (concentrated groups of burrows) in northern Australia. We used radio-telemetry, remote photography, and the complete excavation of a warren to test the hypotheses that 1) warrens are used by multiple individual V. panoptes; and if so, 2) they are used for communal nesting; or alternatively 3) they are used for communal estivation during the dry season. At least six individual V. panoptes utilized the warren system including four females and two males, and burrows were excavated by both sexes. Excavation of the warren revealed no estivating lizards at a time when four radio-telemetered V. panoptes had begun estivation. However, we found two nests in the warren, indicative of either communal nesting or multiple clutches of the same female. Nests were deeper than that recorded for any other reptile and were structurally complex. We discuss the implications of the depth and structure of the nesting burrow for the thermal and hydric environment of the eggs and for hatchling emergence. The warrens usage by multiple individuals raises the possibility that the severe declines in V. panoptes caused by invasive Cane Toads (Bufo marinus) may have important implications for the V. panoptes social structure.

Invasive toads shift predator-prey densities in animal communities by removing top predators.

Although invasive species can have substantial impacts on animal communities, cases of invasive species facilitating native species by removing their predators have rarely been demonstrated across vertebrate trophic linkages. The predictable spread of the invasive cane toad (Rhinella marina), however, offered a unique opportunity to quantify cascading effects. In northern Australia, three species of predatory monitor lizards suffered severe population declines due to toad-induced lethal toxic ingestion (yellow-spotted monitor (Varanus panoptes), Mertens water monitor (V. mertensi), Mitchells water monitor (V. mitchelli). We, thus, predicted subsequent increases in the abundance and recruitment of prey species due to the reduction of those predators. Toad-induced population-level declines in the water monitor species approached 50% over a five-year period spanning the toad invasion, apparently causing fledging success of the Crimson Finch (Neochmia.phaeton) to increase from 55% to 81%. The consensus of our original and published long-term data is that invasive cane toads are causing predators to lose a foothold on top-down regulation of their prey, triggering shifts in the relative densities of predator and prey in the Australian tropical savannah ecosystem.

The ecology and evolution of temperature-dependent reaction norms for sex determination in reptiles: a mechanistic conceptual model

Sex‐determining mechanisms are broadly categorised as being based on either genetic or environmental factors. Vertebrate sex determination exhibits remarkable diversity but displays distinct phylogenetic patterns. While all eutherian mammals possess XY male heterogamety and female heterogamety (ZW) is ubiquitous in birds, poikilothermic vertebrates (fish, amphibians and reptiles) exhibit multiple genetic sex‐determination (GSD) systems as well as environmental sex determination (ESD). Temperature is the factor controlling ESD in reptiles and temperature‐dependent sex determination (TSD) in reptiles has become a focal point in the study of this phenomenon. Current patterns of climate change may cause detrimental skews in the population sex ratios of reptiles exhibiting TSD. Understanding the patterns of variation, both within and among populations and linking such patterns with the selection processes they are associated with, is the central challenge of research aimed at predicting the capacity of populations to adapt to novel conditions. Here we present a conceptual model that innovates by defining an individual reaction norm for sex determination as a range of incubation temperatures. By deconstructing individual reaction norms for TSD and revealing their underlying interacting elements, we offer a conceptual solution that explains how variation among individual reaction norms can be inferred from the pattern of population reaction norms. The model also links environmental variation with the different patterns of TSD and describes the processes from which they may arise. Specific climate scenarios are singled out as eco‐evolutionary traps that may lead to demographic extinction or a transition to either male or female heterogametic GSD. We describe how the conceptual principles can be applied to interpret TSD data and to explain the adaptive capacity of TSD to climate change as well as its limits and the potential applications for conservation and management programs.

Impacts of the invasive cane toad on aquatic reptiles in a highly modified ecosystem: the importance of replicating impact studies

AbstractnInvasive species can have dramatic and detrimental effects on native species, and the magnitude of these effects can be mediated by a plethora of factors. One way to identify mediating factors is by comparing attributes of natural systems in species with heterogeneity of responses to the invasive species. This method first requires quantifying impacts in different habitats, ecosystems or geographic locations. We used a long-term, before-and-after study to quantify the impacts of the invasive and toxic cane toad (Rhinella marina) on two predators in a highly modified ecosystem: an irrigation channel in an agricultural landscape. Survey counts spanning 8xa0years indicated a severe population-level decline of 84xa0% in Merten’s Water Monitor (Varanus mertensi) that was coincident with the arrival of cane toads. The impact of cane toads on V. mertensi was similar to that found in other studies in other habitats, suggesting that cane toads severely impact V. mertensi populations, regardless of habitat type or geographic location. In contrast, a decline was not detected in the Freshwater Crocodile (Crocodylus johnstoni). There is now clear evidence that some C. johnstoni populations are vulnerable to cane toads, while others are not. Our results reinforce the need for the replication of impact studies within and among species; predicting impacts based on single studies could lead to overgeneralizations and potential mismanagement.

Rediscovery of the scaly-tailed possum (Wyulda squamicaudata) in the eastern Kimberley

The tropical mammal fauna of Australia is both understudied and, in some cases, imperiled, and the former hinders a complete understanding of the latter. An enigmatic and poorly understood species is the scaly-tailed possum (Wyulda squamicaudata), a species endemic to the Kimberley Region, Western Australia. We describe the rediscovery of the scaly-tailed possum in the east Kimberley, where it has not been recorded since 1917. The discovery: (1) reinforces the hitherto-questioned validity of the east Kimberley record; (2) confirms an extension of the range by 200–300u2009km to the east from populations in the west Kimberley; and thus (3) broadens the climate envelope occupied by the species. Implications of the known distribution for the biology, genetics and conservation of the scaly-tailed possum are briefly discussed.

Chronic effects of an invasive species on an animal community

Invasive species can trigger trophic cascades in animal communities, but published cases involving their removal of top predators are extremely rare. An exception is the invasive cane toad (Rhinella marina) in Australia, which has caused severe population declines in monitor lizards, triggering trophic cascades that facilitated dramatic and sometimes unexpected increases in several prey of the predators, including smaller lizards, snakes, turtles, crocodiles, and birds. Persistence of isolated populations of these predators with a decades-long sympatry with toads suggests the possibility of recovery, but alternative explanations are possible. Confirming predator recovery requires longer-term study of populations with both baseline and immediate post-invasion densities. Previously, we quantified short-term impacts of invasive cane toads on animal communities over seven years at two sites in tropical Australia. Herein, we test the hypothesis that predators have begun to recover by repeating the study 12xa0yr after the initial toad invasion. The three predatory lizards that experienced 71-97% declines in the short-term study showed no sign of recovery, and indeed a worse fate: two of the three species were no longer detectable in 630xa0km of river surveys, suggesting local extirpation. Two mesopredators that had increased markedly in the short term due to these predator losses showed diverse responses in the medium term; a small lizard species increased by ~500%, while populations of a snake species showed little change. Our results indicate a system still in ecological turmoil, having not yet reached a new equilibrium more than a decade after the initial invasion; predator losses due to this toxic invasive species, and thus downstream effects, were not transient. Given that cane toads have proven too prolific to eradicate or control, we suggest that recovery of impacted predators must occur unassisted by evolutionary means: dispersal into extinction sites from surviving populations with alleles for toxin resistance or toad avoidance. Evolution and subsequent dispersal may be the only solution for a number of species or communities affected by invasive species for which control is either prohibitively expensive, or not possible.