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Caleb L. Loughran

Publications and source records attributed to Caleb L. Loughran.

8 recordsLinked to original sources
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↗
USGS2015

The effect of research activities and winter precipitation on voiding behaviour of Agassiz’s desert tortoises ( Gopherus agassizii )

Context: There is little information available on how research activities might cause stress responses in wildlife, especially responses of threatened species such as the desert tortoise ( Gopherus agassizii ). Aims: The present study aims to detect behavioural effects of researcher handling and winter precipitation on a natural population of desert tortoises in the desert of Southwestern United States, over the period 1997 to 2014, through extensive assessments of capture events during multiple research studies, and capture–mark–recapture survivorship analysis. Methods: Juvenile and adult desert tortoises were repeatedly handled with consistent methodology across 18 years during 10 study seasons. Using a generalised linear mixed-effects model, we assessed the effects of both research manipulation and abiotic conditions on probability of voiding. Additionally, we used a Cormack–Jolly–Seber model to assess the effects of winter precipitation and voiding on long-term apparent survivorship. Key results: Of 1008 total capture events, voiding was recorded on 83 (8.2%) occasions in 42 different individuals. Our top models indicated that increases in handling time led to significantly higher probabilities of voiding for juveniles, females and males. Similarly, increases in precipitation resulted in significantly higher probabilities of voiding for juveniles and females, but not for males. Tortoise capture frequency was negatively correlated with voiding occurrence. Cormack–Jolly–Seber models demonstrated a weak effect of winter precipitation on survivorship, but a negligible effect for both voiding behaviour and sex. Conclusions: Handling-induced voiding by desert tortoises may occur during common research activities and years of above average winter precipitation. Increased likelihood of voiding in individuals with relatively low numbers of recaptures suggested that tortoises may have perceived researchers initially as predators, and therefore voided as a defensive strategy. Voiding does not appear to impact long-term survivorship in desert tortoises at this site. Implications: This study has demonstrated that common handling practices on desert tortoise may cause voiding behaviour. These results suggest that in order to minimise undesirable behavioural responses in studied desert tortoise populations, defined procedures or protocols must be followed by the investigators to reduce contact period to the extent feasible.

Wildlife Research↗
USGS2013

Borehole-explosion and air-gun data acquired in the 2011 Salton Seismic Imaging Project (SSIP), southern California: description of the survey

The Imperial and Coachella Valleys are being formed by active plate-tectonic processes. From the Imperial Valley southward into the Gulf of California, plate motions are rifting the continent apart. In the Coachella Valley, the plates are sliding past one another along the San Andreas and related faults (fig. 1). These processes build the stunning landscapes of the region, but also produce damaging earthquakes. Rupture of the southern section of the San Andreas Fault (SAF), from the Coachella Valley to the Mojave Desert, is believed to be the greatest natural hazard that California will face in the near future. With an estimated magnitude between 7.2 and 8.1, such an event would result in violent shaking, loss of life, and disruption of infrastructure (freeways, aqueducts, power, petroleum, and communication lines) that might bring much of southern California to a standstill. As part of the nation’s efforts to avert a catastrophe of this magnitude, a number of projects have been undertaken to more fully understand and mitigate the effects of such an event. The Salton Seismic Imaging Project (SSIP), funded jointly by the National Science Foundation (NSF) and the U.S. Geological Survey (USGS), seeks to understand, through seismic imaging, the structure of the Earth surrounding the SAF, including the sedimentary basins on which cities are built. The principal investigators (PIs) of this collaborative project represent the USGS, Virginia Polytechnic Institute and State University (Virginia Tech), California Institute of Technology (Caltech), Scripps Institution of Oceanography (Scripps), University of Nevada, Reno (UNR), and Stanford University. SSIP will create images of underground structure and sediments in the Imperial and Coachella Valleys and adjacent mountain ranges to investigate the earthquake hazards posed to cities in this area. Importantly, the images will help determine the underground geometry of the SAF, how deep the sediments are, and how fast earthquake energy can travel through the sediments. All of these factors determine how hard the earth will shake during a major earthquake. If we can improve on our understanding of how and where earthquakes will occur, and how strong their resultant shaking will be, then buildings can be designed or retrofitted accordingly in order to resist damage and collapse, and emergency plans can be adequately prepared. In addition, SSIP will investigate the processes of rifting and magmatism in the Salton Trough in order to better understand this important plate-boundary region. The Salton Trough is a unique rift in that subsidence is accompanied by huge influxes of infilling sediment from the Colorado River. Volcanism that accompanies the subsidence here is muted by these influxes of sediment. The Salton Trough, in the central part of the Imperial Valley, is apparently made up of entirely new crust: young sediment in the upper crust and basaltic intrusive rocks in the mid-to-lower crust (Fuis and others, 1984). Similar to the ultrasound and computed tomography (CT) scans performed by the medical industry, seismic imaging is a collection of techniques that enable scientists to obtain a picture of what is underground. The petroleum industry routinely uses these techniques to search for oil and gas at relatively shallow depths; however, the scope of this project demanded that we image as much as 30 km into the Earth’s crust. This project generated and recorded seismic waves, similar to sound waves, which move downward into the Earth and are bent (refracted) or echoed (reflected) back to the surface. SSIP acquired data in a series of intersecting lines that cover key areas of the Salton Trough. The sources of sound waves were detonations (shots) in deep boreholes, designed to create energy equivalent to magnitude 1–2 earthquakes. The study region routinely experiences earthquakes of these magnitudes, but earthquakes are not located in such a way as to permit us to create the detailed images we need for earthquake hazard assessment. Air gun bursts, generated in the Salton Sea along extensions of our onshore seismic lines, also were utilized as sound-wave sources. Temporary deployments of portable land seismometers, as well as ocean-bottom seismometers (OBSs) on the floor of the Salton Sea, recorded the energy from the land shots and air gun bursts. SSIP is similar to the Los Angeles Regional Seismic Experiments of 1994 and 1999 (LARSE I and II, respectively; Murphy and others, 1996; Fuis and others, 2001). The LARSE surveys demonstrated that the USGS and collaborators can safely and effectively conduct seismic imaging surveys in urban and nonurban areas, on lands owned and/or managed by many different types of agencies and entities. Information was produced that could not have been obtained any other way, and this information was key to changing the leading ideas about earthquake hazards at that time in the Los Angeles region. These surveys produced no significant environmental impact or damage to structures, and they did not trigger earthquakes.

California↗
USGS2011

Long-term post-fire effects on spatial ecology and reproductive output of female Agassiz’s desert tortoises ( Gopherus agassizii ) at a wind energy facility near Palm Springs, California, USA

We studied the long-term response of a cohort of eight female Agassiz’s desert tortoises ( Gopherus agassizii ) during the first 15 years following a large fire at a wind energy generation facility near Palm Springs, California, USA. The fire burned a significant portion of the study site in 1995. Tortoise activity areas were mapped using minimum convex polygons for a proximate post-fire interval from 1997 to 2000, and a long-term post-fire interval from 2009 to 2010. In addition, we measured the annual reproductive output of eggs each year and monitored the body condition of tortoises over time. One adult female tortoise was killed by the fire and five tortoises bore exposure scars that were not fatal. Despite predictions that tortoises would make the short-distance movements from burned to nearby unburned habitats, most activity areas and their centroids remained in burned areas for the duration of the study. The percentage of activity area burned did not differ significantly between the two monitoring periods. Annual reproductive output and measures of body condition remained statistically similar throughout the monitoring period. Despite changes in plant composition, conditions at this site appeared to be suitable for survival of tortoises following a major fire. High productivity at the site may have buffered tortoises from the adverse impacts of fire if they were not killed outright. Tortoise populations at less productive desert sites may not have adequate resources to sustain normal activity areas, reproductive output, and body conditions following fire.

California↗
USGS2011

Gopherus agassizii (Desert Tortoise). Non-native seed dispersal

Sahara Mustard (Brassica tournefortii) is a non-native, highly invasive weed species of southwestern U.S. deserts. Sahara Mustard is a hardy species, which flourishes under many conditions including drought and in both disturbed and undisturbed habitats (West and Nabhan 2002. In B. Tellman [ed.], Invasive Plants: Their Occurrence and Possible Impact on the Central Gulf Coast of Sonora and the Midriff Islands in the Sea of Cortes, pp. 91–111. University of Arizona Press, Tucson). Because of this species’ ability to thrive in these habitats, B. tournefortii has been able to propagate throughout the southwestern United States establishing itself in the Mojave and Sonoran Deserts in Arizona, California, Nevada, and Utah. Unfortunately, naturally disturbed areas created by native species, such as the Desert Tortoise (Gopherus agassizii), within these deserts could have facilitated the propagation of B. tournefortii. (Lovich 1998. In R. G. Westbrooks [ed.], Invasive Plants, Changing the Landscape of America: Fact Book, p. 77. Federal Interagency Committee for the Management of Noxious and Exotic Weeds [FICMNEW], Washington, DC). However, Desert Tortoises have never been directly observed dispersing Sahara Mustard seeds. Here we present observations of two Desert Tortoises dispersing Sahara Mustard seeds at the interface between the Mojave and Sonoran deserts in California.

Herpetological Review↗
USGS2011

Gopherus agassizii (desert tortoise). Burrow collapse

In the deserts of the southwestern U.S., burrows are utilized by the Desert Tortoise to escape environmental extremes (reviewed by Ernst and Lovich 2009. Turtles of the United States and Canada. 2nd ed. Johns Hopkins Univ. Press, Baltimore, Maryland. 827 pp.). However, the potential for mortality through burrow collapse and entrapment is poorly documented. Nicholson and Humphreys (1981. Proceedings of the Desert Tortoise Council, pp. 163−194) suggested that collapse due to livestock trampling may cause mortality. In addition, Lovich et al. (2011. Chelon. Cons. Biol. 10[1]:124–129) documented a Desert Tortoise that used a steel culvert as a burrow surrogate. The culvert filled completely with sediment following a significant rain event, entombing the animal and ultimately resulting in its death. We note that this mortality was associated with an anthropogenic structure; because tortoises are prodigious diggers, one might hypothesize that they have the ability to dig out of collapsed natural burrows in most situations. Circumstances described here presented us with an opportunity to test this hypothesis.

Herpetological Review↗