Global Assessment of Reptile Distributions - BloGARD

Long lived reptiles: What causes reptiles to have longer lifespans?

29/10/2018

In a recent publication in Biological Journal of the Linnean Society, we present a comparative analysis exploring patterns and drivers of longevity of 1320 reptile species, spanning all orders.
In recent years, there have been many studies focusing on the effect of different ecological variables and life-history traits on the variation in longevity of specific taxonomic groups, focusing mostly on birds and mammals (with the only large- scale ectothermic study done on squamates by Scharf et al. 2015). In order to expand our knowledge on the effect of environmental and life-history variables on the variation in longevity of animals, we tested the effect of ecological variables (through various hypotheses) related to extrinsic mortality (e.g. predation) on the variation in longevity among and within lizards, snakes, turtles and crocodiles. We found that species living on islands, and in colder and more seasonal environments, live longer. Moreover, sampling more individuals increases the chances of finding older specimens, and should be corrected for when studying maximum longevity.

We hope these analyses will enable us to better understand the drivers of longevity in reptiles (and other taxa). We hope this paper will facilitate more large-scale comparative studies on the causes of the variation in longevity of tetrapods in general.

Species representing long lived reptiles (from each order) I studied in my research, in the top: right, Sphenodon punctatus, left, Aldabrachelys gigantea, in the bottom: right, Gavialis gangeticus, left, Cyclura lewisi.

Author: Gavin Stark

Global lizard trait database

29/8/2018

In a recent publication in Global Ecology and Biogeograpy, I present a vast dataset of over 20 body size, ecological, thermal biology, geographic, phylogenetic and life history traits for global lizards.

Over the past 12 years I have been collecting trait data on lizards to complement GARD’s geographic data and allow asking interesting ecological, evolutionary and biogeographic questions – as well as, hopefully, informing conservation decisions. To this end I've now published geographical, morphological, ecological, physiological and life history data for the 6,657 known species of lizards. In the data paper I present descriptive statistics regarding these traits and point to avenues for future research using the dataset.

Scincus scincus (Photo: Simon Jamison)

I hope these data will facilitate more study into the biology of these most fascinating of creatures, and that the database publication will encourage others to add yet more data and to correct errors I surely have made when assembling them.

Author: Shai Meiri

In Memoriam - Ben Collen

22/5/2018

We are deeply saddened by the untimely loss of our former GARD member Dr. Ben Collen, and extend our deepest condolences to his family and friends. Ben was an outstanding scientist, and conservationist, and will be greatly missed.

Hiding in plain sight: rare lizards are more common than we think

23/11/2017

In a recently published paper in Diversity and Distributions we try to illuminate aspects regarding the biology, and conservation of all narrow ranged lizard species, across the globe.

We defined lizard species with the smallest ranges as those only known from a single locality, with a maximum range extent no larger than 10 km. Surprisingly, more than 900 species, or roughly 1 in seven of all known lizard species, have such small ranges. Furthermore, about 750 of these species have never been seen again after their initial discovery, and more than 200 lizard species are only known to science from a single individual.

Enyailoides altotambo from Ecuador ( Photo: Omar Torres Carvajal)

When exploring different attributes of small ranged species we found that most of them inhabit relatively inaccessible places in tropical climates worldwide. Furthermore, they are mostly small bodied species; many of them are active at night; and live in rocky habitats. Among the different lizard groups geckos and skinks dominate with many rare species.

Riama yumborum from Ecuador (Photo: Omar Torres Carvajal)

Many of these species (such as those inhabiting small islands or caves) may truly have small ranges. However others may actually have larger ranges, and we are simply ignorant of the true extent of their distribution. This is especially true those found in remote, inaccessible places with no obvious barriers to their dispersal. Thus their small ranges are potentially only an artifact of our poor knowledge. Distinguishing between these two possibilities is both illuminating from an ecological and evolutionary perspective and extremely important from a conservation point of view.

This work could help better focus conservation efforts by pointing at the species, and places, that are in the greatest need of protection. Many of the species, especially those which have not been observed for decades, may well be already extinct. However, to-date only six of the species studied have been officially recognized as such. In order to examine the true extent of such extinctions, and try to prevent future ones, the study provides invaluable information for directing future research and conservation efforts.

Lizard species known only from their type localities. Circles: species not observed after 1967. Crosses: species observed after 1967.

Authors: Shai Meiri and Uri Roll

The Hot Eurasian nightlife - How do different environmental forces affect nocturnality in lizards?

10/10/2017

In a recent publication in Global Ecology and Biogeography we explored the prevalence of nocturnality amongst Eurasian lizard species and tried to understand what drives these patterns.

Most animals – at least those that live above ground – are active either during the day or during the night. Being active at either time of day carries with it unique benefits and challenges, and thus particular adaptations. Because of this being nocturnal or diurnal is a trait that is pretty rigid amongst closely related species.

Lizards as a group are thought to be ancestrally diurnal. Most of them remain so to this day. Furthermore, they are ectotherms and are predominantly small bodied tetrapods and could thus be particularly affected by the climatic differences between day and night. For this work we collected distribution range and activity time data for all 1,113 lizard species found throughout mainland Eurasia. We then looked at links between richness patterns of lizards with either temperature or productivity.

Cyrtodactylus trilatofasciatus (Photo: Lee Grismer)

We found that nocturnal lizards have the highest species richness in the tropics and in deserts, and their richness decreases when they get closer to the North Pole. Nocturnal lizards are precluded altogether from the coldest regions inhabited by lizards – in high mountains and the highest latitudes.

Stenodactylus sthenodactylus (Photo Uri Roll)

Ambient temperature has a strong influence on richness patterns of both diurnal and nocturnal lizards, where species numbers increase with an increase in temperature. Productivity was found to be more tightly related to the proportion of nocturnal species – again in a positive relationship.

We think that our results point towards the fact that low temperatures are a limiting factor for lizard activity period. It is possible that the year-round warm nights of tropic regions enabled lizards to move towards nocturnal activity. In hot deserts, perhaps the combination of hot days and aridity make diurnal activity less attractive, whereas nocturnal activity can provide shelter from these extreme conditions

Author: Enav Vidan

Stenodactylus doriae (Photo Uri Roll)

A short history of GARD and how it has been used to highlight gaps in global conservation priorities

9/10/2017

In a paper published in Nature Ecology and Evolution we present the first global maps of all reptiles - and thus complete the global distributions of all tetrapods. We further explore how the new reptile information changes how we think about global conservation priorities. As this is the first place where all of the GARD maps have been used and published, we use this opportunity to share some of the history of GARD itself, as well as the particular work that was carried out for this paper.

The beginnings of GARD
Planning for reptile conservation globally we first needed to map the distribution of all known species. About 8500 of them when we started in 2006, about 10,500 now recognized. This was a time when such global databases were being published for amphibians, birds, and mammals – some of us have been instrumental in assembling those databases, so we felt fairly confident we knew how it should be done.

What we were wondering, however, was whether the fact that reptile distributions were not collated at the time was not because it couldn’t be. A quick survey of available field guides and herpetology books revealed that maps of the sort used to assemble distribution data for birds and mammals were simply unavailable for huge parts of the world, including most of the crucial regions in tropical South America, Africa and Southern Asia.

GARD meeting, Oxford (photo: Uri Roll)

GARDians competing for that 'perfect' picture of a gecko

Thus the Global Assessment of Reptile Distribution working group (GARD) was formed. In the meantime we started recruiting the people who did much of the actual legwork – graduate students who digitized maps from existing sources, as well as the maps that started pouring in from the reptile experts among the GARD members. We had to keep track with constant taxonomic changes, species splits and new reptile species discoveries (many of them by GARD members themselves) – resulting in additional 200 species or reptiles nowadays being added annually.

We finally had at least some data for all the species or reptiles we thought one could map about two year ago. Then we met again to start the immensely important process of reviewing the distribution data to ensure errors were kept to the minimum (a process that is still ongoing).

Early on in compiling the data we got the feeling that lizard ‘hotspots’ were not in the tropics, where virtually all other groups studied so far have the most species. Once the maps were fully compiled this was very evident. The unique thermal requirements of reptiles enable them to thrive in drier habitats, allowing them to evolve and prosper in deserts.

Vipera bornmuelleri (photo: Uri Roll)

Do unique reptilian biologies and ecologies demand particular conservation needs?
Or in other words do the major global conservation priorities designations adequately represent reptiles or do their unique distributions make them less protected. It turns out that many reptiles – predominantly lizards and turtles are left out of global priority regions and protected areas.

Hypsilurus papuensis (photo: Alex Slavenko)

We therefore wanted to explore how the focus of future conservation efforts need to change to properly represent reptiles. To do this we run prioritization optimization procedures which enabled us to highlight various regions of the world predominantly in drylands, savannah, steppe, and also islands that increase in importance when reptile distribution data are added. More broadly this work highlights the need of getting better data for lesser known groups in order to compile truly inclusive conservation planning that encapsulates all of biodiversity.

Ecoregions that increase in importance for conservation, when reptile data are added (dark blue - top decile ecoregions, light blue - top quartile ecoregions)

Authors: Shai Meiri and Uri Roll

Island life only works if you’re easy-going – uncovering predictions of the island syndrome for lizard clutch size variation

18/9/2017

In a recent publication in the Journal of Biogeography we show that Insular lizards with variable clutch sizes follow the predictions of the island syndrome, while lizards with fixed clutches do not.

Life-histories of insular species are hypothesized to slow down, a phenomenon known as the "island syndrome". Insular individuals are thus expected to lay smaller clutches of larger eggs compared with individuals belonging to closely related mainland species. Most lizards have variable clutch sizes and can lay any number between one egg and a species-specific maximum, which can be well over 50 eggs. Many lizards, such as geckos and anoles, however, lay invariant small clutches of one or two eggs, and may thus be unable to manifest some aspects of the island syndrome. We tested whether insular species with either variable or invariant clutch sizes respond to insularity differently by analyzing egg, clutch, hatchling and female sizes and brood frequencies of 2,511 lizard species.

Mediodactylus kotschyi (photo Rachel Schwarz)

Pafilis & Rachel, Kalogria region NW Peloponnes (photo Shai Meiri)

We found that insular species with variable clutch sizes lay smaller clutches of larger eggs, from which larger hatchlings emerge, compared with mainland species, as expected by the island syndrome. Lizards with invariant clutch sizes, however, lay smaller clutches on islands and increase clutch frequency, compared with mainland species, perhaps because of limitations set by the female body cavity and pelvic opening. This may result from lower seasonality of tropical islands, leading to a greater spread of reproductive effort, or as a result from fluctuations in population densities caused by tropical storms. Our results also emphasize the importance of taking differences in life-history traits into account while studying lizard reproductive traits on large phylogenetic scales.

Kampana islet (photo: Rachel Schwarz)

Author: Rachel Schwarz

The latitudinal diversity gradient and interspecific competition: no global relationship between lizard dietary niche breadth and species richness

27/1/2017

In a 2017 publication in Global Ecology and Biogeography, we collated a novel quantitative volumetric dietary dataset for 308 lizard species worldwide from the field and literature. This novel dataset enabled us to test seven competing hypotheses posited to explain dietary niche breadth, focusing on those that are thought to either cause, or be influenced by, the latitudinal diversity gradient.

Notomabuya frenata (photo Alison Gainsbury)

A species’ niche breadth is defined as the suite of environments or resources that the species can inhabit or use. Niche breadth is often invoked to explain the latitudinal diversity gradient. The latitudinal diversity gradient is the increase in species richness or biodiversity that occurs from the poles to the tropics. Despite this pattern having been recognized for over 200 years, the processes that drive and maintain the latitudinal diversity gradient remain unclear. We investigated which processes are important drivers of global lizard dietary niche breadth patterns, focusing on the relationship between niche breadth and species richness.

A major tenant explaining greater species richness in the tropics is interspecific competition. Dietary niche breadth has long been hypothesized to decrease from the poles toward the tropics, as the numbers of competitors increase. Geographical variation in niche breadth is also hypothesized to be linked to high ambient energy levels, water availability, productivity and climate stability – reflecting an increased number of available prey taxa. Range size and body size are also hypothesized to be strongly and positively associated with niche breadth. We sought to determine which of these factors is associated with geographical variation in niche breadth across broad spatial scales and thus potentially drive the latitudinal diversity gradient.

Overall, our findings are consistent with the notion that climate is an important predictor of dietary specialization, with both less rainfall and more stable temperatures associated with narrower dietary niches. Trophic interactions between lizard species and their arthropod prey are sensitive to climate. It is likely that climatic conditions not only affect these interactions but also alter the functional role of other vertebrate predators in terrestrial ecosystems. The sensitivity of dietary niche breadth to climate has important implications for essential ecosystem functions that maintain increased species richness in the tropics, such as food web stability and energy flow.

Norops meridionalis (photo: Alison Gainsbury)

The synergistic effects of a narrow dietary niche and small range size augments the vulnerability of species to habitat loss and climate change. Based on our findings, the ‘competitionist’s paradigm’ seems to be the exception rather than the rule in explaining the latitudinal diversity gradient.

Author: Alison Gainsbury

Large and lonely - lately extinct reptiles were mostly on islands, and usually big

17/8/2016

When exploring the extinction of Late Quaternary reptiles it seems that body-size was an important predictor of extinction rate. Larger reptiles were more likely to go extinct - published now in Global Ecology and Biogeograpy.

As extinction rates in our world increase at an alarming rate, proper conservation actions require detailed knowledge of the factors influencing extinction. These could be both anthropogenic pressured placed on natural environments, as well as particular species traits which make them especially vulnerable to these pressures. Over the last 50,000 years in which humans have spread across the Earth, there have been waves of mass extinctions of birds and mammals wherever humans colonized. These extinctions were particularly pronounced for large-bodied animals.

Meiolania Platyceps

Varanus priscus from the Melbourne Museum

We set out to examine if a similar pattern could be found for reptiles. In other words, we wanted to know if extinction also favored larger-bodied reptiles. We compiled data on body sizes of all currently known extant species of reptiles, just over 10,000 different species, as well as on 82 species of reptiles known to have gone extinct during the Late Quaternary, following human colonization of their original distribution ranges. By comparing the two groups, extant and extinct, we found that at least for lizards and turtles, extinct species were remarkably large.

Prime examples include the largest lizard to have ever lived, the Megalania monitor Varanus priscus, or the various species of giant tortoises on islands in the Indian and Pacific Oceans. By far the clearest pattern, however, is that extinctions mostly occurred on islands. Almost 90% of all extinct reptile species were endemic to islands!

Hoplodactylus delcourti

The causes for these extinctions are numerous, and include over-harvesting by humans, introduction of invasive carnivores and rats, habitat change by human colonization, and possibly indirect cascade effects caused by the extinction of other, co-existing species. Our study helps us better understand the mechanisms of extinction in reptiles, and therefore might prove useful for pinpointing species which might be vulnerable to anthropogenic pressures in the future, and thus in need of conservation planning.

localities of extinct late Quaternary reptiles

Author: Alex Slavenko

Patterns of species richness, endemism and environmental gradients of African reptiles

28/7/2016

In our recent publication in the Journal of Biogeography, we assembled a comprehensive distribution map of all reptiles in Africa in order to quantify their geographical overlap with the other vertebrate groups, and to assess the environmental correlates underlying these patterns.

The latitudinal gradient of increasing biological diversity towards the equator is one of the best recognized patterns in biogeography, and has been acknowledged for some time. The naturalist, Alexander von Humboldt wrote of his travels over 200 hundred years ago, that as we approach the tropics, "the greater the variety of structure, form, colour, youth and vigor of organic life." A number of well-known hypotheses explaining this pervasive pattern of the increasing number of different species towards the equator have since proliferated. These include elevated ambient energy and precipitation, the number of different habitats or niches, higher plant productivity, and many more.

Until now reptile diversity gradients have remained largely unmapped and the least studied of the terrestrial vertebrates, especially in Africa. This is an important distinction because reptiles are an extremely diverse class of terrestrial vertebrates (over 10,000 species and counting), and as ectotherms, which often thrive in arid regions, their diversity patterns are thought to differ from the classic latitudinal gradient of the other land vertebrates (amphibians, birds, and mammals). In addition, the distinct reptile lineages - amphisbaenians, crocodiles, lizards, snakes, and turtles are likely to respond differently to environmental variables.

Vipera palaestinae (photo: Uri Roll)

To create our geographic distribution map of reptiles in Africa, we obtained data from a variety of field-guides and atlases, museum databases, the primary literature, IUCN assessments, and maps based on expert knowledge of reptile species and the habitats they occupy. A challenging aspect of the project was to ensure that our maps remained current with respect to new species discoveries and taxonomic name changes (which are constantly being revised), and we also had to confirm the validity of type specimen identifications and localities, especially those referenced from obscure sources and archaic museum specimens. We used GIS software to digitize and overlay the maps of each individual African reptile species (1,601 species in total!) one on top of the other, which allowed us to count the number of species present in a given area - which we call “species richness”.

Here is the product of all of that hard work - the first comprehensive richness map of all reptile species in Africa. The colour codes correspond to the number of species from low (blue) to high (red). It shows that the reptile richness map is largely congruent with previously mapped amphibian, bird, and mammal richness showing the classic species latitudinal gradient, including high richness in the arid regions not seen in the other vertebrates. But when you look at the reptile groups distinctly you see that while the overall reptile richness map mostly resembles snakes, lizards in particular are qualitatively very different. Lizard richness hotspots are widely dispersed with high diversity in tropical regions, as well as arid and mountainous areas, where the distribution of the other reptile and non-reptile groups is relatively low.

When we looked at which environmental predictors best explained these species richness maps we found that net primary productivity (the amount of photosynthetic activity by plants) and precipitation explain most of the variation in reptile and other vertebrates. This explains the clear latitudinal pattern seen in their respective maps, which reflects a strong correlation with plant productivity and rainfall as you move closer to the equator. But again, lizards are unique in that none of these environmental correlates explain their distributions. This is because lizards are well adapted to a wide range of habitats including the tropics as well as the harsh conditions of the desert where plant productivity and rainfall are low. We also showed that individual lizard species on average occupy smaller geographic distributions, reflecting their ability to occupy diverse niches.

Our findings that the distribution of lizard species in Africa is unique when compared to the other vertebrate groups now confirms a pattern that has been seen elsewhere in previous studies (i.e. Australia) and most recently by our paper on the global distribution of reptiles. This shows the importance of studying the diverse reptile groups distinctly instead of lumping them all together, and will have bearing on large-scale conservation efforts that do not represent all reptile groups.

Author: Amir Lewin