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Geology topics

Jeffrey E. Lovich

Publications and source records attributed to Jeffrey E. Lovich.

At least 73 records · Page 4Linked to original sources
USGS2017

Unusual population attributes of invasive red-eared slider turtles ( Trachemys scripta elegans ) in Japan: do they have a performance advantage?

The slider turtle ( Trachemys scripta Thunberg in Schoepff, 1792) is native to the USA and Mexico. Due to the popularity of their colorful hatchlings as pets, they have been exported worldwide and are now present on all continents, except Antarctica. Slider turtles are well-established in Japan and occupy aquatic habitats in urban and agricultural areas, to the detriment of native turtles with which they compete. We asked the overall question, do slider turtles in Japan have a performance advantage because they are liberated from the numerous competing turtle species in their native range and released from many of their natural predators? Traits compared included various measures of adult body size (mean, maximum), female size at maturity as measured by size of gravid females, clutch size, population density and biomass, sex ratio, and sexual size dimorphism, the latter two a partial reflection of growth and maturity differences between the sexes. We sampled slider turtle populations in three habitats in Japan and compared population attributes with published data for the species from throughout its native range in the USA. Mean male body sizes were at the lower end of values from the USA suggesting that males in Japan may mature at smaller body sizes. The smallest gravid females in Japan mature at smaller body sizes but have mean clutch sizes larger than some populations in the USA. Compared to most populations in the USA, slider turtles achieve higher densities and biomasses in Japanese wetlands, especially the lotic system we sampled. Sex ratios were female-biased, the opposite of what is reported for many populations in protected areas of the USA. Sexual size dimorphism was enhanced relative to native populations with females as the larger sex. The enhanced dimorphism is likely a result of earlier size of maturity in Japanese males and the large size of mature (gravid) Japanese females. Slider turtles appear to have a performance advantage over native turtles in Japan, possibly as a result of being released from competition with numerous sympatric turtle species in their native range, and the absence of many co-evolved predators and parasites in Japan. This slight competitive edge, coupled with the catholic diet and broad tolerance of varying aquatic habitats of slider turtles, is reflected in their dominance over native and naturalized Japanese turtles in altered aquatic habitats.

Aquatic Invasions↗
USGS2017

Reptiles and amphibians

Summary – We reviewed all the peer-reviewed scientific publications we could find on the known and potential effects of wind farm development, operation, maintenance, and decommissioning on reptiles and amphibians (collectively herpetofauna) worldwide. Both groups are declining globally due to a multitude of threats including energy development. Effect studies were limited to the long-term research by the authors on Agassiz’s Desert Tortoise ecology and behavior at single operational wind farm in California, US and an analysis of the effects of wind farm installation on species richness of vertebrates including reptiles and amphibians in northwestern Portugal. Research on Agassiz’s Desert Tortoise found few demonstrable differences in biological parameters between populations in the wind farm and those in more natural habitats. High reproductive output is due to the regional climate and not to the presence or operation of the wind farm. Site operations have resulted in death and injury to a small number of adult tortoises and over the long-term tortoises now appear to avoid the areas of greatest turbine concentration. Research in Portugal using models and simulations based on empirical data show that vertebrate species richness (including herpetofauna) decreased by almost 20% after the installation of only two large monopole turbines per 250 x 250 m plot. Knowledge of the known responses of herpetofauna to various disturbances allows identification of potential impacts from construction material acquisition in offsite areas, mortality and stress due to impacts of roads and related infrastructure, destruction and modification of habitat, habitat fragmentation and barriers to gene flow, noise, vibration, electromagnetic field generation, heat from buried high voltage transmission lines, alteration of local and regional climate, predator attraction, and increased risk of fire. Research on herpetofauna lags far behind what is needed and, in particular, before-after-control-impact studies are critically needed to identify cause and effect relationships in order to develop effective mitigation strategies for any negative impacts.

Book chapter↗
USGS2017

Developing an effective Agassiz's Desert Tortoise monitoring program: Final report to the Coachella Valley Conservation Commission

Agassiz’s desert tortoise (Gopherus agassizii) is a conservation-reliant species with populations north and west of the Colorado River protected as threatened under the Endangered Species Act (Averill-Murray et al. 2012). Since it was listed under this category in 1990, a great deal has been learned about the natural history of the species, and it is now one of the best-studied turtles in the United States (Lovich and Ennen 2013). However, the accumulated body of scientific data available for the species has not yet been translated into recovery or delisting of the species. Successful conservation of any species requires knowledge of their natural history and how vital rates affect their ability to maintain stable populations in the face of natural and anthropogenic stresses. Agassiz’s desert tortoises occur from southwestern Utah to near the Mexican border in California – a distance of over 450 km – but population densities vary greatly across this immense landscape (U.S. Fish and Wildlife Service 2015). Tortoises occur in the Sonoran Desert of California, including the eastern and western ends of the Coachella Valley, where it is one of 27 species covered under the Coachella Valley Multiple Species Habitat Conservation Plan and Natural Community Conservation Plan (CVMSHCP/NCCP). The southern portion of Joshua Tree National Park (JTNP) lies within this 1.1 million acre planning area, and was predicted to be an area of low-density tortoise populations using habitat suitability modeling (Barrows 2011). JTNP is near the southern distributional limit of G. agassizii, yet very little has been published regarding the ecology of tortoises in the Sonoran Desert of California. Reproductive output is an important gross measure of the ability of a population to persist. When integrated with data on fertility and survivorship, this information forms a foundation for assessing population status and formulating effective management strategies (e.g., Congdon et al. 1993, 1994), especially for imperiled species. One aspect of the biology of G. agassizii that has been particularly well-studied is reproductive output. However, most of what we know about this topic comes from research in the Mojave Desert portion of the species’ range (Ernst and Lovich 2009). Comparatively little has been published on the reproductive ecology of populations living in the Sonoran Desert ecosystem of California. Publications by Lovich et al. (1999, 2011, 2012, 2014, 2015) constitute the main body of literature on desert tortoise reproductive ecology in the Sonoran Desert of California, with one study population located at the western end of the CVMSHCP/NCCP area. Collecting data on Agassiz’s desert tortoise ecology in the Sonoran Desert ecosystem is important due to significant differences between the two adjacent desert ecosystems, especially the timing and amounts of annual precipitation, and their potential effects on reproductive output (e.g., Lovich et al. 5 2015). There are also differences in the vulnerability of tortoises to the effects of a warming, drying climate between the two deserts (Barrows 2011; Zylstra et al. 2012). The overall goal of this study was to collect data on demography, reproductive output, and genetic affinities at a study site in the Sonoran Desert portion of JTNP in the eastern end of the CVMSHCP/NCCP area. Specific objectives included: 1) Collect data to establish baselines on tortoise populations and/or their habitat suitability in core habitat within the CVNCCP area, including biotic and abiotic variables affecting persistence of tortoise populations; 2) Compare and contrast with data collected on desert tortoises at USGS/BLM study site near Palm Springs over 16 years; 3) Support long-term modeling efforts needed to determine tortoise population viability; 4) Refine modeled relationships with identified threats such as fire, invasive species and climate change; and 5) Prioritize adaptive management needs for the desert tortoise in and beyond the CVNCCP area. The data from this study will aid in determining baseline estimates of the desert tortoise population size within the planning area as well as establish a marked population of Agassiz’s desert tortoises for future monitoring. Data will be integrated with habitat modeling in order to refine model output. Genetic data will be collected on both the north and south sides of Interstate 10 to determine the potential effects of habitat fragmentation and genetic mixing. Analyses are ongoing and results beyond those presented in this report will be published in peer-reviewed scientific journals following inclusion of additional data collected on the south side of Shavers Valley in 2017-2018.

Report↗
USGS2017

A report on genetic affinities and relatedness of Agassiz’s desert tortoises (Gopherus agassizii) at opposite ends of the Coachella Valley in California

This report summarizes the results for mtDNA and STR genotyping of 41 desert tortoise (Gopherus agassizii) DNA samples from opposite sides of the Coachella Valley: one sample from the west side at the Mesa wind energy facility in the Whitewater Hills and the other from the mouth of Cottonwood Canyon in Joshua Tree National Park, both within the boundaries of the Coachella Valley Multiple Species Habitat Conservation Plan. Additional samples were collected from tortoises on the northern bajadas of the Orocopia Mountains and from the Santa Rosa Mountains and those results will be presented at a later time in a scientific publication. We tested samples for their mtDNA haplotype and 25 STR loci previously used in other studies. We performed assignment testing to determine the genetic affinity of each individual to the geographic region of collection. Despite apparent isolation, both populations appear to be naturally occurring and do exhibit indications of having experienced increased genetic drift (resulting in increased homozygosity, increased inbreeding or a reduction of genetic diversity). The lack of strong evidence for genetic isolation suggests that long-term maintenance of unfragmented landscapes is an important part of tortoise conservation in the region.

California↗
USGS2017

Climate change and collapsing thermal niches of Mexican endemic reptiles

Recent climate change should result in expansion of species to northern or high elevation range margins, and contraction at southern and low elevation margins due to extinction. Climate models predict dramatic extinctions and distributional shifts in the next century, but there are few ground-truths of these dire forecasts leading to uncertainty in predicting extinctions due to climate change. Previously, we reported on recent extinctions of Mexican Sceloporus lizards by comparing recent surveys to historical distributional records for 48 species at 200 sites. We also ground-truthed extinctions on five continents across 8 lizard families by comparing observed and predicted extinctions from an eco-physiological species distribution model and obtained a high R 2 of 0.72 (1, 2). Here, we derive more detailed predictions for 15 terrestrial reptile families and 142 species for the Mexican and California Biogeographic provinces using all known museum occurrence records, and detailed measures on eco-physiology. We adopt the eco-physiological model of extinction developed earlier but use a species-specific model. We predict massive and rapid extinctions of 22% of the reptile populations in Mexico within the next 50 years. We also predict that 3 of 15 reptile families, all three endemic to the Mexican and Californian biogeographic provinces, will go extinct by 2070, the hallmark of the beginnings of a mass extinction event. However, extinctions may be attenuated by forest cover and by presence of montane environments in contemporary ranges. We describe impacts of altitude on three species (Gopherus morafkai, G. evgoodei, and Gambelia sila) to illustrate regional management strategies (AZ-Mexico, Sinoloa, CA) for reserves in tandem with global strategies of CO2 limits that might limit climate impacts. By carefully selecting new montane preserves adjacent to desert and tropical forest habitats, and by implementing global controls on atmospheric CO2 emissions, extinctions may be reduced to less than 11% of species and only a single reptile family.

Report↗
USGS2016

A morphological review of subspecies of the Asian box turtle, Cuora amboinensis (Testudines, Geomydidae)

The turtle Cuora amboinensis has an extensive distribution covering most of southern mainland Asia, Indonesia, and extending to the Philippine Islands. Unlike many species, C. amboinensis occurs on both sides of Wallace's Line separating Asian and Australian flora and fauna. Four subspecies are currently recognized; Cuora a . kamaroma (southern continental Asia, Java and the northern Philippines [introduced]), C. a . lineata (Kachin Province, Myanmar [Burma] and adjacent Yunnan Province, China), C. a . couro (Sumatra, Java, Sumbawa, and adjacent smaller Indonesian islands); and C. a . amboinensis (Moluccas, Sulawesi, Philippines). Five pattern and 33 morphological characters were examined for variation in 691 individuals from throughout the species' range. Our analyses suggest that only two presently recognized subspecies are valid: amboinensis and kamaroma . Neither couro nor lineata are supported by our analysis. We recommend that C. a . couro should be synonymized with the species C. amboinensis and C. a . lineata with the subspecies C. a . kamaroma .

Proceedings of the Biological Society of Washingto↗
USGS2016

Graptemys pearlensis Ennen, Lovich, Kreiser, Selman, and Qualls 2010 – Pearl River Map Turtle

The Pearl River Map Turtle, Graptemys pearlensis (Family Emydidae), is a moderate-sized aquatic turtle endemic to the Pearl River drainage of Louisiana and Mississippi. This taxon has long been a cryptic species, as it was considered part of G. pulchra before 1992 and part of G. gibbonsi until 2010. Graptemys pearlensis exhibits sexual dimorphism, with adult females being considerably larger (carapace length to 295 mm) than adult males (CL to 121 mm). In the 1960s and 1970s, the species was commonly found in higher abundance than the sympatric G. oculifera, a federally listed species. However, due to habitat degradation and the precipitous decline of native mollusks, the species is now found in lower numbers than G. oculifera throughout much of its range. The current IUCN Red List status is Endangered; however, very little is known about the natural history and ecology of the species, which will make conservation efforts challenging.

Book chapter↗
USGS2016

Using climate, energy, and spatial-based hypotheses to interpret macroecological patterns of North America chelonians

Our study investigates how factors, such as latitude, productivity, and several environmental variables, influence contemporary patterns of the species richness in North American turtles. In particular, we test several hypotheses explaining broad-scale species richness patterns on several species richness data sets: ( i ) total turtles, ( ii ) freshwater turtles only, ( iii ) aquatic turtles, ( iv ) terrestrial turtles only, ( v ) Emydidae, and ( vi ) Kinosternidae. In addition to spatial data, we used a combination of 25 abiotic variables in spatial regression models to predict species richness patterns. Our results provide support for multiple hypotheses related to broad-scale patterns of species richness, and in particular, hypotheses related to climate, productivity, water availability, topography, and latitude. In general, species richness patterns were positively associated with temperature, precipitation, diversity of streams, coefficient of variation of elevation, and net primary productivity. We also found that North America turtles follow the general latitudinal diversity gradient pattern (i.e., increasing species richness towards equator) by exhibiting a negative association with latitude. Because of the incongruent results among our six data sets, our study highlights the importance of considering phylogenetic constraints and guilds when interpreting species richness patterns, especially for taxonomic groups that occupy a myriad of habitats.

Canadian Journal of Zoology↗
USGS2016

Aspects of the reproductive ecology of female turtles in New Mexico

Data on reproductive ecology of turtles in New Mexico are limited, and some species living there are among the least studied in the United States. We trapped 4 native species of turtles ( Apalone spinifera, Chrysemys picta, Pseudemys gorzugi, and Trachemys gaigeae gaigeae ) in the Rio Grande and Black River (Pecos River drainage) of New Mexico in June 2012 and 2013 to collect data on female reproductive ecology, including clutch size, egg size, timing of egg production, and percentage of gravid females. During our sampling, we found shelled eggs via X-radiography in only 3 native species: C. picta, P. gorzugi, and T. g. gaigeae. Clutch and egg sizes were within the range of previously reported values, although clutch size for P. gorzugi (10 eggs) is only the second published record for that data-deficient species. Clutch size increased with body size in T. g. gaigeae. We observed few differences between reproductive parameters for turtles in New Mexico and their conspecifics and congeners elsewhere in the United States, other than the observation that female C. picta may mature at smaller body sizes in New Mexico relative to other western populations elsewhere in its vast, primarily eastern North American range.

New Mexico↗
USGS2016

Interdrainage morphological and genetic differentiation in the Escambia Map Turtle, Graptemys ernsti

Graptemys ernsti, the Escambia Map Turtle, inhabits the Escambia/Conecuh River, the adjacent Yellow River, and the Pea River further to the east, all of which have been distinct drainage systems since the Pleistocene. We used continuous and meristic morphological and genetic data to compare populations of G. ernsti and found evidence of differences among the three drainages. Frequency of occurrence of a nasal trident differed among the three drainages. Yellow River specimens possessed unique mitochondrial haplotypes while the Conecuh and the Pea shared haplotypes. Five microsatellite loci identified the drainages as being distinct, with the strongest differentiation between the Yellow River and the other two drainages. While these differences do not appear great enough to warrant taxonomic recognition, they do suggest that each population has a distinct evolutionary and demographic history and that they should therefore be managed separately.

Herpetological Conservation and Biology↗
USGS2016

Energy development

Large areas of the desert southwest are currently developed or being evaluated for construction of utility-scale renewable energy projects. These projects include numerous solar and wind energy facilities some of which will be massive. Unfortunately, peer-reviewed scientific publications are not yet available to evaluate the potential effects of solar-based utility-scale renewable energy development (USRED) on any species of wildlife, including amphibians and reptiles (herpetofauna). Scientific publications on the effects of wind-based USRED and operation (USREDO) are focused almost exclusively on flying wildlife including birds and bats. To the best of our knowledge the only publications on the effects of wind-based USREDO on herpetofauna are three publications on desert tortoise ecology at a wind energy facility near Palm Springs, California. Those studies suggested that not all effects of USREDO were detrimental in the short-term. However, additional research is required to determine if wind energy operation is compatible with conservation of this long-lived species over longer periods of time.

Book chapter↗
USGS2016

A potential predator-prey interaction of an American badger and an Agassiz's desert tortoise with a review of badger predation on turtles

The federally threatened Agassiz’s desert tortoise ( Gopherus agassizii ) was listed under the U.S. Endangered Species Act in 1990, but thus far, recovery efforts have been unsuccessful (U.S. Fish and Wildlife Service [USFWS] 2015). Predation has been identified as a contributing factor to declining G. agassizii populations range-wide (e.g., Esque et al. 2010, Lovich et al. 2014). Understanding and managing for predator-prey dynamics is thus an important part of the recovery and conservation of this threatened species (USFWS 2011). Desert tortoises have a host of predators at all stages of their life cycle. Over 20 species of birds, mammals, and reptiles have been recorded as known or suspected predators (Woodbury and Hardy 1948, Luckenbach 1982, Ernst and Lovich 2009). American badgers ( Taxidea taxus , family: Mustelidae) are confirmed excavators of desert tortoise nests (Turner and Berry 1984). They are also suspected predators of adult desert tortoises, a possibility which has been presented in some studies but without empirical verification (Luckenbach 1982, Turner and Berry 1984). Active mostly at night, badgers are solitary, secretive predators (Lindzey 1978, 1982; Armitage 2004) that are extremely difficult to observe in predatory encounters. Recently, strong circumstantial evidence presented by Emblidge et al. (2015) suggests that badgers do prey on adult Agassiz’s desert tortoises based on observations of more than two dozen dead tortoises in the Western Mojave Desert of California. In this note, we present another case of potential badger predation on a large adult desert tortoise in the Sonoran Desert of California. Collectively, these recent two cases potentially indicate that badger predation may be more common and widespread than previously thought. In addition, we review the worldwide literature of badger predation on turtles in general and summarize reported badger observations in Joshua Tree National Park, where our observation occurred, over a period of 55 years.

California↗
USGS2016

Measuring the impact of invasive species on popular culture: a case study based on toy turtles from Japan

The red-eared slider turtle (Trachemys scripta elegans) is native to portions of the United States of America (USA) and adjacent northeastern Mexico. The bright and colorful hatchlings have long been popular as pets globally but numerous individuals have been released into the wild establishing populations in areas well outside their native range. As a result, slider turtles are now introduced worldwide on all continents, with the exception of Antarctica, and many temperate and tropical islands, including Japan. They are very successful at establishing breeding populations in a variety of habitats, even those in proximity to human development. Once established in large populations, they compete with native turtle species sometimes to the detriment of the latter. Tin toy turtles were popular in Japan for decades, and they were an important export item after World War II. From the 1920s to the 1950s, prior to the widespread establishment of slider populations in Japan, the toys were characterized by muted earth-tone colors representative of native species of Japanese turtles. After the 1950s, toy turtles exhibited brighter combinations of yellow, red and green more typical of slider turtles. This transition may reflect demand for more colorful toys by importing countries like the USA. Alternatively, the change was coincident with the importation of large numbers of colorful slider turtles to Japan via the pet trade and their subsequent establishment and numerical dominance in Japanese wetlands. This switch in toy turtle colors may reflect a cultural transition in awareness of what constitutes the appearance of a typical turtle in Japan. Sliders appear to have been accepted by Japanese consumers as a new cultural norm in the appearance of turtles, a case of art imitating life.

Humans and Nature↗
USGS2015

Relationships of maternal body size and morphology with egg and clutch size in the diamondback terrapin, Malaclemys terrapin (Testudines: Emydidae)

Because resources are finite, female animals face trade-offs between the size and number of offspring they are able to produce during a single reproductive event. Optimal egg size (OES) theory predicts that any increase in resources allocated to reproduction should increase clutch size with minimal effects on egg size. Variations of OES predict that egg size should be optimized, although not necessarily constant across a population, because optimality is contingent on maternal phenotypes, such as body size and morphology, and recent environmental conditions. We examined the relationships among body size variables (pelvic aperture width, caudal gap height, and plastron length), clutch size, and egg width of diamondback terrapins from separate but proximate populations at Kiawah Island and Edisto Island, South Carolina. We found that terrapins do not meet some of the predictions of OES theory. Both populations exhibited greater variation in egg size among clutches than within, suggesting an absence of optimization except as it may relate to phenotype/habitat matching. We found that egg size appeared to be constrained by more than just pelvic aperture width in Kiawah terrapins but not in the Edisto population. Terrapins at Edisto appeared to exhibit osteokinesis in the caudal region of their shells, which may aid in the oviposition of large eggs.

South Carolina↗
USGS2015

Intersexual allometry differences and ontogenetic shifts of coloration patterns in two aquatic turtles, Graptemys oculifera and Graptemys flavimaculata

Coloration can play critical roles in a species' biology. The allometry of color patterns may be useful for elucidating the evolutionary mechanisms responsible for shaping the traits. We measured characteristics relating to eight aspects of color patterns from Graptemys oculifera and G. flavimaculata to investigate the allometric differences among male, female, and unsexed juvenile specimens. Additionally, we investigated ontogenetic shifts by incorporating the unsexed juveniles into the male and female datasets. In general, male color traits were isometric (i.e., color scaled with body size), while females and juvenile color traits were hypoallometric, growing in size more slowly than the increase in body size. When we included unsexed juveniles in our male and female datasets, our linear regression analyses found all relationships to be hypoallometric and our model selection analysis found support for nonlinear models describing the relationship between body size and color patterns, suggestive of an ontogenetic shift in coloration traits for both sexes at maturity. Although color is critical for many species' biology and therefore under strong selective pressure in many other species, our results are likely explained by an epiphenomenon related to the different selection pressures on body size and growth rates between juveniles and adults and less attributable to the evolution of color patterns themselves.

Ecology and Evolution↗
USGS2015

Turbines and terrestrial vertebrates: variation in tortoise survivorship between a wind energy facility and an adjacent undisturbed wildland area in the desert southwest (USA)

With the recent increase in utility-scale wind energy development, researchers have become increasingly concerned how this activity will affect wildlife and their habitat. To understand the potential impacts of wind energy facilities (WEF) post-construction (i.e., operation and maintenance) on wildlife, we compared differences in activity centers and survivorship of Agassiz's desert tortoises ( Gopherus agassizii ) inside or near a WEF to neighboring tortoises living near a wilderness area (NWA) and farther from the WEF. We found that the size of tortoise activity centers varied, but not significantly so, between the WEF (6.25 ± 2.13 ha) and adjacent NWA (4.13 ± 1.23 ha). However, apparent survival did differ significantly between the habitat types: over the 18 year study period apparent annual survival estimates were 0.96 ± 0.01 for WEF tortoises and 0.92 ± 0.02 for tortoises in the NWA. High annual survival suggests that operation and maintenance of the WEF has not caused considerable declines in the adult population over the past two decades. Low traffic volume, enhanced resource availability and decreased predator populations may influence annual survivorship at this WEF. Further research on these proximate mechanisms and population recruitment would be useful for mitigating and managing post-development impacts of utility scale wind energy on long-lived terrestrial vertebrates.

Environmental Management↗
USGS2015

Not putting all their eggs in one basket: bet-hedging despite extraordinary annual reproductive output of desert tortoises

Bet-hedging theory makes the counter-intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short-term reproductive output to minimize the risk of reproductive failure in the long-term. We investigated the long-term reproductive output of an Agassiz's desert tortoise ( Gopherus agassizii ) population and conformance to a bet-hedging strategy of reproduction in an unpredictable but comparatively productive environment. Most females reproduced every year, even during periods of low precipitation and poor germination of food plants, and the mean percentage of reproducing females did not differ significantly on an annual basis. Although mean annual egg production (clutch size × clutch frequency) differed significantly among years, mean clutch size and mean clutch frequency remained relatively constant. During an El Niño year, mean annual egg production and mean annual clutch frequency were the highest ever reported for this species. Annual egg production was positively influenced by maternal body size but clutch size and clutch frequency were not. Our long-term results confirm earlier conclusions based on short-term research that desert tortoises have a bet-hedging strategy of producing small clutches almost every year. The risk of long-term reproductive failure is minimized in unpredictable environments, both through time by annually producing multiple small clutches over a long reproductive lifespan, even in years of low resource availability, and through space by depositing multiple annual clutches in different locations. The extraordinary annual reproductive output of this population appears to be the result of a typically high but unpredictable biomass of annual food plants at the site relative to tortoise habitat in dryer regions. Under the comparatively productive but unpredictable conditions, tortoises conform to predictions of a bet-hedging strategy of reproduction with relatively small but consistent clutch sizes.

Biological Journal of the Linnean Society↗