Quantitative analyses of squamate dentition demonstrate novel morphological patterns

Squamates are ideal subjects for investigating relationships between diet and dental patterns because they exhibit wide dietary diversity, marked variation in dental shape, and are taxonomically abundant. Despite this, well-established links between diet and dental morphology are primarily qualitative in nature, with specific patterns of squamate dental complexity remaining largely unknown. Here, we use quantitative methods and a broad taxonomic dataset to quantify key patterns in squamate dental morphology, including re-examining the relationship between dentition and diet, testing for differences in complexity between dentigerous elements, and exploring the effect of ontogenetic dietary shifts in dental complexity in two iguanid genera. Our findings support previous research by demonstrating that species consuming more plant material possess more complex teeth. We did not find significant complexity differences between the left and right dentigerous elements nor the upper and lower jaws, with the exception of Amblyrhynchus cristatus, the marine iguana, which possesses significantly more complex dentary teeth than premaxillary and maxillary teeth. We find discordant patterns when testing for dental complexity changes through ontogeny. Amblyrhynchus, which is primarily herbivorous throughout its lifetime, increases dental complexity through ontogeny, whereas Ctenosaura, which is generally insectivorous as juveniles and herbivorous as adults, decreases dental complexity. Although preliminary, this research documents and quantifies novel patterns of squamate dental complexity and exhibits the possibilities for further research on the diversity of squamate dental morphology.

Beautiful on the Inside

Marine iguanas stand in stark contrast to the Galápagos rails. They vary enormously in space by size and color, supporting classification into 11 subspecies (whereas rails are monotypic). And they vary in time, especially males, which change to bright colors in the mating season—some to the point of being bright red and green “Christmas iguanas.” They vary over time in an additional special way: iguanas shrink up to 20% (or 2.7 inches) during the food scarcity of El Niño events. They have evolved the capacity to shut down their normal stress response when the ocean gets warm, thus becoming quiescent and riding out the storm. The larger the iguanas, the more they shrink and the longer they survive. Furthermore, nearly all marine iguana subspecies have increased in size since 1905, while, at the same time, climate change has made El Niño events stronger and longer. Could climate change be driving the evolution of larger marine iguanas? Will iguanas be able to keep up as El Niño worsens?

No impact of a short-term climatic “El Niño” fluctuation on gut microbial diversity in populations of the Galápagos marine iguana (Amblyrhynchus cristatus)

Gut microorganisms are crucial for many biological functions playing a pivotal role in the host’s well-being. We studied gut bacterial community structure of marine iguana populations across the Galápagos archipelago. Marine iguanas depend heavily on their specialized gut microbiome for the digestion of dietary algae, a resource whose growth was strongly reduced by severe “El Niño”-related climatic fluctuations in 2015/2016. As a consequence, marine iguana populations showed signs of starvation as expressed by a poor body condition. Body condition indices (BCI) varied between island populations indicating that food resources (i.e., algae) are affected differently across the archipelago during ‘El Niño’ events. Though this event impacted food availability for marine iguanas, we found that reductions in body condition due to “El Niño”-related starvation did not result in differences in bacterial gut community structure. Species richness of gut microorganisms was instead correlated with levels of neutral genetic diversity in the distinct host populations. Our data suggest that marine iguana populations with a higher level of gene diversity and allelic richness may harbor a more diverse gut microbiome than those populations with lower genetic diversity. Since low values of these diversity parameters usually correlate with small census and effective population sizes, we use our results to propose a novel hypothesis according to which small and genetically less diverse host populations might be characterized by less diverse microbiomes. Whether such genetically depauperate populations may experience additional threats from reduced dietary flexibility due to a limited intestinal microbiome is currently unclear and calls for further investigation.

Three new species of chigger mites (Acariformes: Trombiculidae) from the Galápagos Islands

Three new species of chigger mites are described from two species of iguana endemic to the Galápagos Islands. Odontacarus cruzi sp. nov. and Schoengastia galapa sp. nov. are found on the marine iguana, Amblyrhynchus cristatus Bell, and Eutrombicula pachytrichia sp. nov. is found on the Santa Fe land iguana, Conolophus pallidus Heller. All these mite species were collected, identified, and named by P.H. Vercammen-Grandjean, but remained undescribed until the present.

Proteomics of Galápagos Marine Iguanas Links Function of Femoral Gland Proteins to the Immune System

Communication between individuals via molecules, termed chemosignaling, is widespread among animal and plant species. However, we lack knowledge on the specific functions of the substances involved for most systems. The femoral gland is an organ that secretes a waxy substance involved in chemical communication in lizards. Although the lipids and volatile substances secreted by the femoral glands have been investigated in several biochemical studies, the protein composition and functions of secretions remain completely unknown. Applying a proteomic approach, we provide the first attempt to comprehensively characterize the protein composition of femoral gland secretions from the Galápagos marine iguana. Using samples from several organs, the marine iguana proteome was assembled by next-generation sequencing and MS, resulting in 7513 proteins. Of these, 4305 proteins were present in the femoral gland, including keratins, small serum proteins, and fatty acid-binding proteins. Surprisingly, no proteins with discernible roles in partner recognition or inter-species communication could be identified. However, we did find several proteins with direct associations to the innate immune system, including lysozyme C, antileukoproteinase (ALP), pulmonary surfactant protein (SFTPD), and galectin (LGALS1) suggesting that the femoral glands function as an important barrier to infection. Furthermore, we report several novel anti-microbial peptides from the femoral glands that show similar action against Escherichia coli and Bacillus subtilis such as oncocin, a peptide known for its effectiveness against Gram-negative pathogens. This proteomics data set is a valuable resource for future functional protein analysis and demonstrates that femoral gland secretions also perform functions of the innate immune system.

The Mystery of a Marine Monster: Morphological and Performance Modifications in the World’s only Marine Lizard, the Galápagos Marine Iguana

Reptiles have repeatedly invaded and thrived in aquatic environments throughout history, however fewer than 8% of the 6000 extant species are primarily aquatic. The Galápagos Marine Iguana (Amblyrhynchus cristatus), the world’s only marine lizard, may have had one of the most unique and challenging transitions to aquatic life. Curiously, previous studies have identified relatively few physiological adaptations in Marine Iguanas, however, little is known about the extent of morphological specialisation and performance trade-offs associated with the marine environment. By examining the morphology and locomotory performance of the Marine Iguana in comparison to their closely related mainland ancestors, the Black Spiny-tailed iguana (Ctenosaura similis) and Green Iguana (Iguana iguana), we found variation reflected specialisation to ecological niches. However, variation was more pronounced among subspecies of Marine Iguana, suggesting that little morphological or performance modification is required for iguanids to successfully invade aquatic environments, thus raising the question why there are so few extant aquatic reptilian lineages. We found that specialisation for the marine environment resulted in a trade-off in sprint speed in a terrestrial environment, similar to that seen in extant crocodilians. Reduced performance in a terrestrial environment likely poses little risk to large-bodied apex predators, whereas in iguanids, a performance trade-off would likely incur increased predation. As such, we suggest that this may explain why iguanids and other ancestral lineages have not undergone transitions to aquatic life. Additionally, we found that the magnitude of morphological and performance variation was more pronounced between subspecies of Marine Iguana than between iguanid species.Summary StatementThe Marine Iguana has undergone a unique evolutionary transition to aquatic behaviour, we explore the extent of morphological and performance specialisation required and why there are so few extant marine reptiles.

Trophic niche differentiation mirrors intra-island population structure of Galápagos marine iguanas (Amblyrhynchus cristatus)

Background: Differences in the trophic niche often underlie ecological specialization of individuals and can promote ecological speciation of populations, but studies showing a link between differences in the trophic niche and genetic differentiation of populations are rare. On the island of San Cristóbal (Galapágos archipelago), a strong genetic differentiation between two relatively proximate populations (subspecies; Amblyrhynchus cristatus mertensi and A. c. godzilla) of marine iguanas along the coastline has been observed. Here, we explore whether this genetic differentiation is mirrored in the iguanas’ trophic niche. Results: Although, no significant difference in the number of consumed algal taxa between subspecies were detected, the Schoener index exhibited low diet overlap between A. c. mertensi and A. c. godzilla. The latter was also demonstrated by the PERMANOVA analysis with significantly different diet OTU composition from the fecal samples between subspecies. Stable isotope analysis revealed that subspecies identity was overall more important than site for the iguanas’ resource use. Conclusions: By applying a metabarcoding approach on feces samples in combination with stable isotope analysis of skin sheds, we found that A. c. mertensi and A. c. godzilla differ in their ecological niches. Moreover, stable isotope analysis indicated that marine iguana populations have low spatial foraging distances, which, together with the diet partitioning patterns, might explain, at least partially, the lack of gene flow between these geographically proximate marine iguana populations. Key words: diet analysis, trophic niche, metabarcoding, stable isotopes, marine iguanas, Amblyrhynchus cristatus