Potential changes in the distribution of Carnegiea gigantea under future scenarios

Introduction

Predicting climate change effects on biodiversity is one the most important challenges that researchers face (Parmesan, 2006). Over the last decades several studies have identified the changes induced by anthropogenic emissions of greenhouse gases is affecting the ecology of desert ecosystems (Parmesan & Yohe, 2003; Kimball et al., 2010). Climate change researchers predict that changes to desert ecosystems will alter the nutrient cycles, fire regimes, genetic diversity of populations with implications to evolutionary changes, and cause species range shifts (Loik et al., 2004; Huxman et al., 2004; Bickford et al., 2010; SDNIMP, 2010). In the Southwest United States, the impacts of climate change on plant abundance and distributions have already been reported, including in the Sonoran Desert (Munson et al., 2012).

The saguaro (Carnegiea gigantea) represents one of the most noticeable patterns of plant distribution in the Sonora Desert (Hutto, McAuliffe & Hogan, 1986). The saguaro is a large columnar cactus that grows to a height of 12 m or more. The main stem, which can range up to 75 cm in diameter, has 12 to 25 vertical ribs (Turner, Bowers & Burgess, 1995; Anderson, 2006). This species inhabits rocky and outwash slopes and grows on sandy flats on or near alluvium (Turner, Bowers & Burgess, 1995). The saguaro is very important to the people of the Tohono O’odham Nation, because they rely on this species for food (Beckwith, 2015). The spatial distribution of the saguaro extends through the Sonoran Desert in Arizona, California, and Mexico, but most of its population occurs in Sonora, Mexico (USFR, 2016). Saguaro disease (e.g., bacterial necrosis), air pollution, cattle grazing, wood-cutting, land use changes, urbanization, freezing and drought are significant threats to the saguaro (NPS, 2010; Burquez-Montijo et al., 2013).

However, despite its importance as one of the signature species of the Sonoran Desert, the giant saguaro has been largely ignored by biogeographers. There is limited evidence that the spatial distribution of the saguaro is driven mainly by the climate in the northernmost part of its range (Hutto, McAuliffe & Hogan, 1986; Turner, Bowers & Burgess, 1995; Arundel, 2005). However, the limiting factors affecting the growth of the saguaro in the eastern Sonora State of Mexico have not been identified yet (Turner, Bowers & Burgess, 1995). Hence, there is an urgent need to review our current understanding of the effects of climate on the saguaro distribution.

Species Distribution Modeling (SDM) are correlative models built from the relationships between environmental variables and incomplete presence records (Guisan & Zimmermann, 2000) that have been used to provide understanding on detailed ecological relationships between abiotic predictors and species distributions, and to predict species’ distributions across space and time (e.g., Guisan & Zimmermann, 2000; Araújo & Rahbek, 2006; Elith et al., 2006). The outcome of an SDM is a habitat suitability map, useful for assessing species invasion and proliferation, designing ecogeographic regions, modeling species richness and composition, and supporting conservation planning and spatial prioritization, among others (Ferrier et al., 2002; Franklin, 2010; Benito, Cayuela & Albuquerque, 2013; Guisan et al., 2013).

When forecasting the effect of climate change on species ‘geographical ranges, it is important to consider multiple climate change scenarios (Sala et al., 2000; Parmesan, 2006; Araújo & Rahbek, 2006; Beaumont et al., 2007; Beaumont, Hughes & Pitman, 2008; Bellard et al., 2012) based on four Representative Concentration Pathways (RCPs, IPCC, 2013). RCPs describe scenarios based on assumptions on socio-economic, and greenhouse and air pollutant emissions to provide trajectories for major agents of climate change (Van Vuuren et al., 2011). In this paper we apply SDMs to investigate how habitat suitability for the saguaro can potentially respond to a range of climate change scenarios. Our results provide guidance on the potential impacts of climate change on the saguaro’s geographical range, while increasing our understanding on the impacts of climate change in the ecology of the Sonora Desert.

Materials & Methods

The Study area comprised the Sonora Desert (Fig. 1), which extends from the Southwestern United States into Northern Mexico, including the states of Arizona, California, Baja California, and Sonora. It is rich in both habitat and biodiversity, and encompasses biotic community representing all of the World’s biomes, such as tundra, forest, grassland, chaparral, desert and riparian communities (Arizona-Sonora Desert Museum, 2018). The Sonoran Desert lifeforms include more than 350 birds, 100 reptiles, and more than 2,000 plant species, including the iconic columnar cacti species saguaro (NPS, 2017).

Results

From the 360 models produced with different combinations of target distance and parameters, models with a target distance of 1km had the best performance. Among them, three models had the same tc (4), bf (0. 5), and AUC (0.87). We selected the model with the smallest learning rate (lr = 0.005).

According to this model, the annual precipitation and max temperature of warmest month were found to have the greatest relative influence on saguaro’s habitat suitability, 24.7%, and 22.8%, respectively (Fig. 2). Also, the mean annual temperature showed a significant contribution (15.7%).

According to the partial dependence plots obtained from the best BRT model, the relationship between saguaro’s habitat suitability and the environmental predictors are non-linear. The partial contribution of the individual predictors to the model fit (Fig. 3) indicate a preference of the species for warmer areas with high precipitation in the summer and open landscapes. For max temperature of warmest month, the logit of the probability of presence displayed a constant response to about 35 °C and then showed a steep increase (Fig. 3).

The eastern and southern areas of Arizona, USA, and Sonora, Mexico, showed the largest concentrations of areas with high suitability, with a secondary concentration in central Sonora state (Fig. 4). Cells with high habitat suitability were also concentrated in a northernmost part of the Sonora desert (Fig. 4).

Model forecast under different RCPs indicated significant habitat suitability reductions across the species’ presence range and few opportunities for range expansion (Fig. 5). In all RCPs, models predicted a high contraction of suitable habitat. By 2050, RCPs predict a loss of 6.9%, on average, of the of currently suitable habitat, with values ranging from 5.6% (RCP 2.6) to 8.6% (RCP 4.5). Much of the contiguous loss of suitable habitat is on the western edge of saguaro’s range in Arizona, with a sizeable loss in a central patch of Arizona. This pattern continues into Mexico, with contractions on the western edge of Sonora, jetting into the mainland, and receding north from Sinaloa. The models projected little habitat suitability increases, most notably in Sonora State’s range. By 2070, this pattern of habitat suitability loss is projected to continue and worsen with an additional 1.2% contraction from 2050. The greatest expected change appears to be enlarging inland patches of unsuitable habitat from Arizona to Mexico. Our models suggest a moderate growth in habitat suitability in the northward expansion from saguaro’s actual range. We also observed an expansion in the center of the Sonora State.

Models also suggest that temperature has low impact on defining expansion and contraction areas for RCPs 2.5 and 4.6, where increased rainfall is the main factor explaining the increase of habitat suitability (Fig. 6). For RCPs 6.0 and 8.5 expansion areas seem to be mostly explained by an increment in both temperature and rainfall (Fig. 6).

Discussion

For first time, we used BRT models to describe the relationships of environmental variables and the habitat suitability of the iconic saguaro across the Sonoran Desert, and predicted habitat suitability change under climate warming for four different RCPs.

In general, the predictive performance of boosted regression trees depends on the model parameters, such as lr and tc. BRT models required a large tr value to achieve a minimum predictive error. After testing different combinations of parameters, three models emerged as best candidates: they shared the same tr (4), bf (0.50) and AUC (0.87). We selected the model with smaller lr (0.005), because small values for lr results in a slower learning and requires a higher number of trees to improve the predictive error (De’ath, 2007). Also, small lr values shrinks the contribution of each tree and reliably estimate the response (Elith, Leathwick & Hastie, 2008). BRT models were also expected to be affected by target distance, as it happened. Models with smaller target distance, and therefore larger sample size, produced higher AUC values than models with larger target distances, which also produced lower explained deviance values.

We found that the habitat suitability of the saguaro is strongly related with climate variables, which also agrees with previous studies performed at local extent (Parker, 1993; Drezner & Balling, 2002). According to BRT’s variable importance measures, annual precipitation had the strongest influence on the habitat suitability of the saguaro. Overall, the probability of occurrence increased with the annual precipitation, rising steeply and uniformly up to 300 mm, followed by a steep decrease and a stationary phase. Several other studies have identified precipitation as a key factor for the demography of saguaros. Turner, Bowers & Burgess (1995) observed that the saguaro grows in areas of the Sonoran Desert where summer rainfall is substantial. Drezner (2006) reported that the reproductive success of the saguaro is closely related with global and regional-scale variations and increases in rainfall. Precipitation is also related with patterns in saguaro establishment and survival (Winkler et al., 2018). The lack of water is pointed as a major factor affecting cacti mortality, probably because water limitation can reduce the survivorship of young and juvenile individuals (Pierson & Turner, 1998). Furthermore, desert ecosystems of western United States and Northern Mexico are particularly susceptible to climate variability and specifically to drought (Archer & Predick, 2008).

According to our model, the max temperature of warmest month had a strong influence on the habitat suitability for the saguaro. Overall, habitat suitability dramatically increased when the maximum temperature of the warmest month went beyond 36 °C. This relationship may occur because of the saguaro is well adapted to the harsh temperature conditions of the Sonoran Desert (Franco & Nobel, 1989). Temperature has been identified as one of the most important factors for the regeneration and population viability of the saguaro (Turner, Bowers & Burgess, 1995; Drezner, 2006), since it plays a key role in driving the establishment and survival of young saguaros, maintaining its distribution over time (Turner et al., 1966; Nobel, 1982). On the other hand, saguaros are sensitive to extended periods of subfreezing temperatures (Nobel, 1982), and catastrophic freeze events have been reported to increase the mortality of the saguaro (Orum, Ferguson & Mihail, 2016).

Our models show potential impacts of climate change on saguaro’s habitat suitability in the Sonoran Desert, a result that is consistent with previous analyses of climate change in desert ecosystems (SDNIMP, 2010; Munson et al., 2012). All models projected onto different RCPs predict a reduction in habitat suitability for the saguaro. Specifically, results indicate that the eastern and central parts of Mexico, and especially the Sonora State, are more sensitive to changes, and face large habitat suitability decreases.

The impacts of climate change on the distribution of the saguaro have recently been reported at the Saguaro National Park, Arizona (Swann et al., 2018). Climate change seems to affect the saguaro directly through increased drought, the occurrence of extended, extreme freezing events, and indirectly because warmer winters temperatures may enhance the spread of exotic species, such as buffelgrass (Cenchrus ciliaris; Swann et al., 2018). Also, drought directly promotes the decline of saguaro density and growth, and reduces perennial shrub and tree cover (nurse plants), which help to protect saguaro from extreme temperatures (Archer & Predick, 2008). We add on to previous studies, showing for the first time the potential changes in the habitat suitability of C. gigantea, under future climate change scenarios, for the whole Sonoran Desert area. Although much work remains to be done to evaluate the effect of climate change on the distribution of saguaro across the Sonoran Desert, our findings provide a strong reason to engage in that work. Because the saguaro distribution is so poorly documented, conservation planners need reliable assessments to monitor the reduction in suitability at Sonoran Desert.

Conclusion

In this study, we used boosted regression trees to investigate effects of climate change on the saguaro habitat suitability, and to explore the complex relationship between environmental factors and saguaro distribution in the Sonora Desert ecosystem at a regional extent. Based on our results, we reached three conclusions: (1) the performance of BRT algorithms varied with the selection of BRT parameters. Overall, BRT models performed well, which reinforces its use for typical ecological analyses (Elith, Leathwick & Hastie, 2008). As indicated by cross-validation analysis, BRT is a useful algorithm for analyzing and predicting ecological data (De’ath, 2007). (2) BRT models identified precipitation and temperature as the main drivers of the habitat suitability for the saguaro in the Sonoran Desert. (3) Although previous studies have reported impacts of climate change on the saguaro, this study is the first attempt to identify potential impacts of climate change on the saguaro’s habitat suitability across its whole range. Previous studies on the possible effects of climate change on saguaro distribution have mostly focused on local scales (Turner et al., 1966; Pierson & Turner, 1998; Archer & Predick, 2008; Swann et al., 2018), while this study is focused on a regional scale. Regardless of the RCP used, models predict a decrease in the saguaro’s habitat suitability across the study area. Also, our results allow us to conclude that under warming conditions an increase in precipitation is required to ensure a high habitat suitability for saguaros. Because saguaros are much more resistant to extended drought than many other species (Swann et al., 2018), we suggest that elucidating the patterns and drivers of species distribution change under climate warming can provide key ecological knowledge necessary to conserve species at the Sonoran Desert.

Supplemental Information