Urbanization, despite its destructive effects on natural habitats, offers species an opportunity to colonize novel niches. Previous research found that urban Anolis lizards in Puerto Rico had increased adhesive toepad area and more ventral toepad scales, traits that are likely adaptive and genetically based. We further investigated these phenotypic changes using geometric morphometrics to measure differences in toe shape, toepad shape, and lamellar morphology. Our results indicate that the increased toepad area of urban Anolis cristatellus lizards in Puerto Rico is not simply an isometric increase in toe size. Toes of urban populations exhibit multiple disproportional changes compared to forest lizards, with a larger proportion of the toe length covered in adhesive toepad. In addition, the toepads of urban lizards increase more in length than width. Lastly, lizards in urban populations exhibit both increased number of lamellae as well as increased spacing between individual lamellae. We also observed regional variation, with urban specimens having significantly more disparity, suggesting similar processes of urban adaptation are likely happening in parallel across the island, yet with region-specific idiosyncrasies, possibly generating more variation in toepad morphology across urban specimens as compared to forest specimens. Considering the use of geometric morphometrics, we found that specimen preparation, specifically how flat and straight toes are during imaging, to be an important factor affecting our data, more so than specimen size or any other meaningful morphological variation. In addition, we found that landmark and semilandmark data can be used to directly estimate toepad area, offering the opportunity to streamline future studies. In conclusion, our results highlight the value of considering toepad morphology in more detail beyond adhesive pad area or number of lamellae. Geometric morphometrics tools may be employed to elucidate subtle differences in shape to better allow researchers to connect changes in morphology to ecology and adhesive performance.
Evidence suggests that hurricanes can influence the evolution of organisms, with phenotypic traits involved in adhesion, such as the toepads of arboreal lizards, being particularly susceptible to natural selection imposed by hurricanes. To investigate this idea, we quantified trait variation before and after Hurricanes Irma and Maria (2017) in forest and urban populations of the Puerto Rican lizard Anolis cristatellus. We found that the hurricanes affected toe morphology differently between forest and urban sites. In particular, toepads of the forefeet were longer and narrower in forest, but wider in urban populations, compared to pre-hurricane measures. Toepads of the hind feet were larger in area following the hurricanes. Fore and rear toes increased in length following the hurricane. There were no changes in the number of lamellae scales or lamellae spacing, but lamellae 6–11 of the forefeet shifted proximally following the hurricane. We also measured clinging performance and toe shape. We found that toepad area and toe lengths were stronger predictors of adhesive forces than toepad shape. Our results highlight an interaction between urbanization and hurricanes, demonstrating the importance to consider how urban species will respond to extreme weather events. Additionally, our different results for fore and rear feet highlight the importance of evaluating both of these traits when measuring the morphological response to hurricanes in arboreal lizards.
more » « less- PAR ID:
- 10436188
- Publisher / Repository:
- Oxford University Press
- Date Published:
- Journal Name:
- Integrative Organismal Biology
- Volume:
- 5
- Issue:
- 1
- ISSN:
- 2517-4843
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Synopsis -
null (Ed.)Extreme climate events such as droughts, cold snaps, and hurricanes can be powerful agents of natural selection, producing acute selective pressures very different from the everyday pressures acting on organisms. However, it remains unknown whether these infrequent but severe disruptions are quickly erased by quotidian selective forces, or whether they have the potential to durably shape biodiversity patterns across regions and clades. Here, we show that hurricanes have enduring evolutionary impacts on the morphology of anoles, a diverse Neotropical lizard clade. We first demonstrate a transgenerational effect of extreme selection on toepad area for two populations struck by hurricanes in 2017. Given this short-term effect of hurricanes, we then asked whether populations and species that more frequently experienced hurricanes have larger toepads. Using 70 y of historical hurricane data, we demonstrate that, indeed, toepad area positively correlates with hurricane activity for both 12 island populations of Anolis sagrei and 188 Anolis species throughout the Neotropics. Extreme climate events are intensifying due to climate change and may represent overlooked drivers of biogeographic and large-scale biodiversity patterns.more » « less
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Abstract Human activity drastically transforms landscapes, generating novel habitats to which species must adaptively respond. Consequently, urbanization is increasingly recognized as a driver of phenotypic change. The structural environment of urban habitats presents a replicated natural experiment to examine trait–environment relationships and phenotypic variation related to locomotion. We use geometric morphometrics to examine claw morphology of five species of Anolis lizards in urban and forest habitats. We find that urban lizards undergo a shift in claw shape in the same direction but varying magnitude across species. Urban claws are overall taller, less curved, less pointed and shorter in length than those of forest lizards. These differences may enable more effective attachment or reduce interference with toepad function on smooth anthropogenic substrates. We also find an increase in shape disparity, a measurement of variation, in urban populations, suggesting relaxed selection or niche expansion rather than directional selection. This study expands our understanding of the relatively understudied trait of claw morphology and adds to a growing number of studies demonstrating phenotypic changes in urban lizards. The consistency in the direction of the shape changes we observed supports the intriguing possibility that urban environments may lead to predictable convergent adaptive change.more » « less
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Abstract Hurricanes cause dramatic changes to forests by opening the canopy and depositing debris onto the forest floor. How invasive rodent populations respond to hurricanes is not well understood, but shifts in rodent abundance and foraging may result from scarce fruit and seed resources that follow hurricanes. We conducted studies in a wet tropical forest in Puerto Rico to better understand how experimental (canopy trimming experiment) and natural (Hurricane Maria) hurricane effects alter populations of invasive rodents (
Rattus rattus [rats] andMus musculus [mice]) and their foraging behaviors. To monitor rodent populations, we used tracking tunnels (inked and baited cards inside tunnels enabling identification of animal visitors' footprints) within experimental hurricane plots (arborist trimmed in 2014) and reference plots (closed canopy forest). To assess shifts in rodent foraging, we compared seed removal of two tree species (Guarea guidonia andPrestoea acuminata ) between vertebrate‐excluded and free‐access treatments in the same experimental and reference plots, and did so 3 months before and 9 months after Hurricane Maria (2017). Trail cameras were used to identify animals responsible for seed removal. Rat incidences generated from tracking tunnel surveys indicated that rat populations were not significantly affected by experimental or natural hurricanes. Before Hurricane Maria there were no mice in the forest interior, yet mice were present in forest plots closest to the road after the hurricane, and their forest invasion coincided with increased grass cover resulting from open forest canopy. Seed removal ofGuarea andPrestoea across all plots was rat dominated (75%–100% rat‐removed) and was significantly less after than before Hurricane Maria. However, following Hurricane Maria, the experimental hurricane treatment plots of 2014 had 3.6 times greater seed removal by invasive rats than did the reference plots, which may have resulted from rats selecting post‐hurricane forest patches with greater understory cover for foraging. Invasive rodents are resistant to hurricane disturbance in this forest. Predictions of increased hurricane frequency from expected climate change should result in forest with more frequent periods of grassy understories and mouse presence, as well as with heightened rat foraging for fruit and seed in preexisting areas of disturbance. -
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Abstract Cyclonic storms, or hurricanes, are expected to intensify as ocean heat energy rises due to climate change. Ecological theory suggests that tropical forest resistance to hurricanes should increase with forest age and wood density. However, most data on hurricane effects on tropical forests come from a limited number of well‐studied long‐term monitoring sites, restricting our capacity to evaluate the resistance of tropical forests to hurricanes across broad environmental gradients.
In this study, we assessed whether forest age and aridity mediate the effects of hurricanes Irma and Maria in Puerto Rico, Vieques and Culebra islands. We leveraged functional trait data for 410 tree species, remotely sensed measurements of canopy height and cover, along with data on forest stand characteristics of 180 of 338 forest monitoring plots, each covering an area of 0.067 ha. The plots represent a broad mean annual precipitation (MAP) gradient from 701 to 4598 mm and a complex mosaic of forest age from 5 to around 85 years since deforestation.
Hurricanes resulted in a 25% increase in basal area mortality rates, a 45% decrease in canopy height and a 21% reduction in canopy cover. These effects intensified with forest age, even after considering proximity to the hurricane path. The links between forest age and hurricane disturbances were likely due the prevalence of tall canopies.
Tall forest canopies were strongly linked with low community‐weighted wood density (WD). These characteristics were on average more common in moist and wet forests (MAP >1250 mm). Conversely, dry forests were dominated by short species with high wood density (WD > 0.6 g cm−3) and did not show significant increases in basal area mortality rates after the hurricanes.
Synthesis . Our findings show that selection towards drought‐tolerant traits across aridity gradients, such as short stature and dense wood, enhances resistance to hurricanes. However, forest age modulated responses to hurricanes, with older forests being less resistant across the islands. This evidence highlights the importance of considering the intricate links between ecological succession and plant function when forecasting tropical forests’ responses to increasingly strong hurricanes.
