Conservation challenges in a threatened

hotspot: agriculture and plant biodiversity

losses in Baja California, Mexico

Sula Vanderplank, Exequiel Ezcurra,

Jose Delgadillo, Richard Felger &
Lucinda A. McDade

Biodiversity and Conservation

ISSN 0960-3115
Volume 23
Number 9

Biodivers Conserv (2014) 23:2173-2182

DOI 10.1007/s10531-014-0711-9

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Biodivers Conserv (2014) 23:2173–2182
DOI 10.1007/s10531-014-0711-9

ORIGINAL PAPER

Conservation challenges in a threatened hotspot:

agriculture and plant biodiversity losses in Baja
California, Mexico

Sula Vanderplank • Exequiel Ezcurra • Jose Delgadillo •

Richard Felger • Lucinda A. McDade

Received: 14 September 2013 / Accepted: 21 April 2014 / Published online: 13 May 2014
Ó Springer Science+Business Media Dordrecht 2014

Abstract Modern agricultural practices pose serious threats to biodiversity worldwide.

Species losses from habitat conversion are well documented, but indirect impacts such as
reduced water availability to adjacent ecosystems are less known. San Quintı́n is an
important agricultural valley in the mediterranean climate region of Baja California,
Mexico. The region is also a hotspot of plant species richness and endemism. Plant species
in the region are here analyzed by comparison of the contemporary flora to historical
botanical collections to identify extirpations. Historical collections indicate that habitat
loss to agriculture has been a direct cause of species losses. As importantly, the unsus-
tainable extraction of groundwater has apparently led to salt water intrusion, resulting in
the loss of 22 native plant taxa, including 13 rare plants. Seventy-eight percent of all the
vernal pool taxa have been lost from the flora (including 85 % of the rare taxa) and 11 % of
plants of riparian and pond habitat (including 25 % of the rare taxa) are no longer found in
the region. Unsustainable agricultural practices continue to threaten fragile coastal eco-
systems and are a serious challenge to current and future conservation efforts. Ironically,

Communicated by David Hawksworth.

S. Vanderplank (&)
Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, TX 76107, USA
e-mail: sula.vanderplank@gmail.com

S. Vanderplank
E. Ezcurra
Department of Botany and Plant Sciences, University of California, Riverside, 900 University Avenue,
Riverside, CA 92507, USA

J. Delgadillo
Facultad de Ciencias, Universidad Autónoma de Baja California, Ensenada, BC, Mexico

R. Felger
Herbarium, University of Arizona, P.O. Box 210036, Tucson, AZ 85721, USA

L. A. McDade
School of Botany, Rancho Santa Ana Botanic Garden, Claremont Graduate University, 1500 North
College Ave, Claremont, CA 91711, USA

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these same practices frequently result in abandonment of cultivated areas. Owing to

indirect impacts, conservation of biodiversity and large-scale agricultural operations are
even less compatible on a regional scale than indicated by direct impacts. It is vital that
sustainable agricultural practices be adopted locally and globally to avoid further losses of
biodiversity.

Keywords Conservation
Sustainability
Groundwater
Species richness

Vernal pools
Extirpation
Salt-water intrusion

Abbreviations
CFP California Floristic Province
GSQ Greater San Quintı́n

Introduction

Reducing extinction that results from human activities is one of the most pressing chal-
lenges of our time. Identifying the causes of localized extirpations is critical to under-
standing metapopulation dynamics and how to avoid extinctions on a global scale. The link
between habitat loss and species losses is well documented; however, there are additional
complexities that affect vulnerability of species to extirpation in impacted habitats. Risk of
extinction is greatly increased for taxa that are endemic to restricted areas (Ellstrand and
Elam 1993; Myers et al. 2000). This is of particular concern in hotspots of endemism
where rare and/or narrowly endemic species tend to co-occur (Brooks et al. 2002).
Fifty percent of the world’s human population live in coastal areas (Finkl 1994) where
development places considerable pressures on coastal ecosystems. Agricultural operations
adjacent to these populations can be challenging since fresh water availability for crop
cultivation is hampered by salt-water intrusion in many coastal regions (Oude Essink
2001). Often the agricultural practices enhance salt-water intrusion, and cause additional
detrimental effects on soil quality including erosion, desertification, salinization, com-
paction, and pollution (Zalidis et al. 2002). Wetland habitats are particularly susceptible;
these are fragile habitats that often house ecologically restricted species particularly in
fragmented landscapes. Wetlands generally exist because of the interaction of surface
water and ground water (Llamas 1988). The amount of time that soil water is present and
the salinity of ground water affect floristic composition in wetland areas (Baldwin and
Mendelssohn 1998). Case studies in Spain highlight the ecological impacts of changes in
ground water availability, as well as the propensity for these changes to go unnoticed for a
number of years, making the problem a serious challenge by the time it is noticed (Llamas
1988).
The California Floristic Province (CFP) runs along the Pacific coast of North America,
from Oregon, USA, to Baja California, Mexico. The CFP has been designated as a global
biodiversity hotspot, an area of high endemism heavily impacted by human activity, with
2,125 endemic plant species (Myers et al. 2000; Brooks et al. 2002), almost 50 % (1,031)
of which are threatened with extinction or have already become extinct (Brooks et al.
2002). About 10 % of these endemics are restricted to NW Baja California, an area that
constitutes less than 5 % of the CFP (Raven and Axelrod 1978; O’Brien et al. in press).
In 1994 the North American Free Trade Agreement (NAFTA) reduced restrictions on
trans-border commerce, spiking international interest in Baja California for large scale

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export-orientated agricultural and industrial projects, notably those that cannot be easily
developed in the US due to laws protecting the environment (Dedina 2007). The coastal
corridor of CFP in Baja California is facing the greatest development and urbanization
pressures in the state (Riemann and Ezcurra 2007). Despite its high conservation priority,
there is currently no legal protection for any of the coastal lands in the CFP of Baja
California (Riemann and Ezcurra 2005) other than that provided by ZOFEMAT (Zona
Federal Marı́timo-Terrestre, or Mexico’s Federal coast administration) concessions, which
control land use of the tidal zone and immediately adjacent land areas.
The San Quintı́n valley is the largest agricultural valley in Baja California that functions
with autochthonous water sources (Fig. 1) (the Mexicali valley uses allochthonous water
from the Colorado River). This valley presently has 14 large agricultural businesses and
more than 20,000 irrigated hectares under cultivation (Martı́nez-Veloz 2012). Aguirre-
Muñoz et al. (2001) demonstrated that ground-water extraction in the surrounding area far
exceeds sustainable limits and current usage is six times higher than the sustainable level.
In the context of the semi-arid climate of NW Baja California, the environmental impact of
groundwater overdraft is a particularly significant problem for indigenous plants (Zektser
et al. 2005) thus a serious concern for conservation efforts in the region. Many abandoned
agricultural fields can be seen locally (Vanderplank 2010) and in adjacent regions (Cas-
tellanos et al. 2005), likely as a direct result of salt water intrusion. For economic
development in this region, aquaculture (e.g., of oysters, abalone) is a more sustainable
alternative to agriculture (Aguirre-Muñoz et al. 2001).
Despite the threat from human activities, the San Quintı́n region continues to supports
several fragile plant communities that are globally rare. The unusual mix of CFP species
with plants from more xeric regions to the south make the regiona floristically rich area
with 435 taxa documented in recent years in an area of 360 km2, Greater San Quintı́n
(GSQ) (Vanderplank 2011). One hundred and twenty-four plant taxa (35 %) of the native
flora of GSQ are rare and/or locally endemic (67 of these are endemic or near-endemic to
NW Baja California) according to the classification of O’Brien et al. (in press). Clearly,
GSQ merits consideration as a priority region for conservation and the area is currently
under consideration by Mexico’s National Commission for Protected Natural Areas for
declaration as a protected area. The cost of conservation measures continues to increase
even as efforts to secure protection for the area proceed at a deliberate pace (Fuller et al.
2007). Threats to species will continue unabated until these efforts are successful: agri-
cultural practices have already led to the extinction of the narrowly endemic San Quintı́n
Kangaroo Rat (Cartron et al. 2005) and several other organisms are at risk (e.g., locally
endemic reptiles and birds; TNC 2007). This study documents changes in the flora using
historical herbarium specimens to identify plants that have been extirpated.

Methods

The vascular plant taxa of GSQ were documented between 2005 and 2010 in a rigorous
floristic survey. Thirty-five collecting trips totaling 75 field days were conducted, during
which more than 1,600 plant specimens were collected. As a means to ensure that all areas
with vegetation were included in the survey, the site was gridded into km2 and every grid
square with habitat suitable for plant growth was surveyed on the ground (Vanderplank
2010). These data (excluding Isla San Martı́n) were compared to historical specimen data
to determine which plant species have been extirpated from the flora. The online resources
of BajaFlora.org, SEINET, CONABIO, and the Global Biodiversity Information Facility

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Fig. 1 Large-scale agriculture in the San Quintı́n valley

provided a wealth of verifiable specimen data, and regional herbaria were visited and
searched for collections from GSQ. Taxa documented by herbarium specimens as histor-
ically present in the study area but not located during the floristic study conducted by
Vanderplank (2011) were first examined to exclude possible misidentifications or
nomenclatural changes, and a refined list of putatively extirpated taxa resulted.
Specimens of extirpated taxa were annotated and label data were used to map their
locations with the goal of assessing each plant’s habitat as well as possible causes for its
loss. ESRI ArcMap was used for mapping and relevant data layers were obtained from the
Instituto Nacional de Estadı́stica y Geografı́a, México (INEGI). Extirpated taxa were
evaluated for rarity using the findings of O’Brien et al. (in press); these authors assessed the
rarity of all taxa in the CFP of Baja California, producing three lists of taxa of concern (i.e.,
rare globally, rare in CFP Baja California but more common elsewhere, and ‘‘watch-list’’
taxa). These were together categorized as ‘‘rare’’ for the purposes of the present analysis.
After mapping the specimens, historical information on the collection sites (e.g., aerial
imagery, literature, field notebooks) was used to identify possible causes of extirpation.
For each habitat type (as identified by Vanderplank 2011), species loss was calculated
as fraction of the total recorded taxa (i.e., number documented as occurring plus extir-
pated). Using contingency-table analysis, losses were compared against the total flora
under the null hypothesis that local extirpations should be randomly distributed among
habitats. In habitats where species loss was higher than would be expected by chance, the
departure from randomness was tested using the normal-distribution properties of Pearson
residuals to assess significance.

Results

A total of 37 taxa historically documented from the study area were not relocated during
the study of Vanderplank (2011) see Table 1. Fifteen of these were non-native taxa, often
occurring only in association with irrigated agriculture (Soto 1987), that are not considered
further.
The remaining 22 species, (13 of which are rare) fall into three groups based on the
geographic region or habitat type from which they were collected: (1) the vernal pools in
Ejido El Papalote (seven taxa extirpated, five of which were rare species); (2) the Laguna

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 Table 1 Species not located by Vanderplank (2011) but documented to have occurred in GSQ by voucher specimens in regional herbaria
Family Taxon Collection Location/habitat Status Reason not found

Apiaceae Spermolepis echinata Thorne 60290 (RSA) Santa Maria 2 Transient in the
escarpment flora?
Apiaceae Hydrocotyle ranunculoides Moran 27975 (SD) Riparian 2 Presumed extirpated
Apiaceae Hydrocotyle verticillata Thunb. var. verticillata Moran 26337 (SD) Riparian 2 Presumed extirpated
Apocynaceae Sarcostemma cynanchoides ssp. hartwegii Moran 24762 (SD) Riparian Presumed extirpated
Asteraceae Centromadia parryi ssp. australis Moran 27955 (SD) Vernal pool 1B Presumed extirpated
Asteraceae Psilocarphus tenellus Nutt. var. tenellus Moran 21820 (SD) Vernal pool Presumed extirpated
Asteraceae Pluchea odorata (Cass.) var. odorata Harbison s.n. (SD) Riparian Presumed extirpated
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Asteraceae Stephanomeria pauciflora Moran 19311 (SD) Riparian Presumed extirpated

Asteraceae Bidens laevis Hall s.n. (SD) Mormona Presumed extirpated
Amaranthaceae Salicornia depressa Moran 29018, 28989, 29431 Mormona 2 Presumed extirpated
(SD)
Cyperaceae Schoenoplectus californicus Harbison s.n. (SD) Riparian Presumed extirpated
Cyperaceae Eleocharis montevidensis Moran 28998 (SD) Mormona Presumed extirpated
Fabaceae Astragalus didymocarpus Barneby var. didymocarpus Moran 29005 (SD) Mormona 1B Presumed extirpated
Fabaceae Astragalus nuttallianus var. cedrosensis Moran 28981 (SD) Mormona 2 Presumed extirpated
Fabaceae Acmispon maritimus (Nutt.) D.D. Sokoloff var. Moran 28997 (SD) Mormona 4 Presumed extirpated
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brevivexillus
Fabaceae Hoffmannseggia glauca Moran 21780 (SD) El Socorro Dunes Transient in the
flora?
Malvaceae Eremalche exilis Moran 25950 (SD) Vernal pool 2 Presumed extirpated
Plantaginaceae Plantago bigelovii Moran 25944 (SD) Vernal pool 2 Presumed extirpated
Poaceae Deschampsia danthonioides Moran 25949 (SD) Vernal pool 2 Presumed extirpated
Poaceae Leptochloa fusca ssp. uninervia Moran 25358 (SD) Vernal pool Presumed extirpated
Polemoniaceae Navarretia fossalis Moran 25945 (SD) Vernal pool 1B Presumed extirpated
Verbenaceae Verbena scabra Wiggins 4766 (RSA) Mormona 2 Presumed extirpated
2177

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 Table 1 continued
2178

Family Taxon Collection Location/habitat Status Reason not found

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Non-native taxa (not considered in the habitat analysis)
Asteraceae Conyza bonariensis Moran 28988 (SD) Mormona * Did not persist
Asteraceae Cynara cardunculus ssp. cardunculus Moran 21809, Thorne 61028 Non-native ruderal * Did not persist
(SD)
Asteraceae Lactuca serriola Soto 26 (BCMEX) Tomato farm * Agricultural weed
Amaranthaceae Chenopodium cf. hians Standl. Soto 24 (BCMEX) Tomato farm * Agricultural weed
Cucurbitaceae Citrullus lanatus Moran 25080 (SD) Escaping cultivation * Did not persist
Cucurbitaceae Cucurbita pepo Moran 25244 (SD) Escape from cultivation * Did not persist
Plumbaginaceae Limonium sinuatum Thorne 58110 (RSA) Escape from cultivation * Did not persist
Poaceae Bromus ciliatus Soto 5 (BCMEX) Tomato farm * Agricultural weed
Poaceae Echinochloa crus-galli Soto 25 (BCMEX) Tomato farm * Agricultural weed
Poaceae Setaria adhaerens Moran 27956 (SD) Vernal pool * Did not persist
Poaceae Triticum aestivum Moran 10501 (SD) Escape from * Did not persist
cultivation?
Portulacaceae Portulaca oleracea Soto 33 (BCMEX) Tomato farm * Agricultural weed
Solanaceae Nicandra physalodes Moran 26330 (SD) Riparian * Did not persist
Solanum douglasii
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Solanaceae Soto 30 (BCMEX) Tomato farm * Agricultural weed

Solanaceae Solanum esculentum Moran 29010 (SD) Mormona * Did not persist
The family, taxon name, collection (collector/number/herbarium), abbreviated locality or habitat, and native status are presented, along with possible reasons for not being
relocated. Status is given for non-native species (*) and for rare taxa using rankings that follow those of the California Native Plant Society (O’Brien et al. in press): 1B—rare
and endangered in CFP Baja and elsewhere; 2—rare in CFP Baja but more common elsewhere; 3—more information needed; 4—rare but not currently threatened with
extinction in CFP Baja; *introduced; plants lacking entries are not considered rare
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Table 2 Native species loss by habitat, excluding ruderal species (data from herbarium records and
Vanderplank 2011), riparian and vernal pool habitats have experienced significant species losses
Habitat Native taxa Extirpated taxa Relative loss (%) Residuals Significance (P)

Total Rare Total Rare % Total % Rare Total Rare Total Rare

Saltmarsh 29 9 0 0 0.0 0.0 -1.19 -0.91 ns ns

Dunes 89 35 1 0 1.1 0.0 -1.61 -1.80 ns ns
Riparian 118 27 13 7 11.0 25.9 3.00 2.84 0.003 0.005
Scrub 188 53 1 1 0.5 1.9 -2.71 -1.77 0.007 ns
Middens 16 10 0 0 0.0 0.0 -0.89 -0.96 ns ns
Vernal pools 9 6 7 5 77.8 83.3 9.88 5.95 \0.0001 \0.0001
Total 443 140 24 13 4.9 9.3

Mormona salt pond area (seven taxa extirpated, five of which are rare), and (3) arroyos of
the Santo Domingo and San Simon drainages (six taxa extirpated, including two rare taxa).
The two exceptions to this pattern are Spermolepis echinata from the coastal scrub and
Hoffmannseggia glauca from the dunes.
Table 2 categorizes species loss by habitat and shows that species losses, both overall of
native taxa and of rare taxa, are higher than expected for plants of vernal pool and riparian
habitats but not for other habitats.

Discussion

The only cases of extirpation that we are able to document are species that were previously
vouchered by early collectors. Because the area was not intensively collected historically
(previous studies in this region were not systematic inventories of the flora), it is likely that
various additional species have been lost from the local flora. Vanderplank’s (2011) study
more than doubled the number of vouchered taxa known from this region. As such, it is
likely that other species that were not historically vouchered have been lost from the flora
without evidence.
The analysis presented here clearly shows that extirpation of species from the GSQ area
is closely tied to habitats that are dependent upon water, specifically vernal pools and
riparian edges. Vernal pools are seasonal wetlands that become inundated after winter rains
due to an impervious soil layer; they gradually dry up in late spring or summer. These
pools experience a brief waterlogged stage followed by extreme desiccation (Keeley and
Zedler 1998). As a result of urban sprawl and agriculture, California has lost 93–97 % of
its vernal pools (Baskin 1994, Kangas 2005) and Baja California has lost at least 92 %
overall, with loss being nearly complete in some areas (Guilliams et al. 2012). Vernal pools
were well-documented in GSQ as occurring in Ejido El Papalote before conversion to
agriculture. Moran (1981) earlier reported that two taxa had been extirpated there as a
direct result of agriculture. A total of eight vernal pool taxa previously collected from Ejido
El Papalote have apparently been extirpated by habitat destruction for agriculture; five of
these taxa are considered globally rare. Vernal pool taxa in the region are now found only
in roadside ditches during years of extremely heavy rainfall; these cannot be regarded as
safe from future extirpation as roadsides are constantly reworked.

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Laguna Mormona, a second site of considerable extirpation, was a large saline pond in
the northern portion of the study area. Although agriculture was clearly encroaching on this
area some decades ago [e.g., Moran (1981) noted that Astragalus didymocarpus was
growing at the edge of a barley field], the semi-saline edges of the pond were not suitable
for cultivation. There has, however, been considerable expansion of irrigated agriculture in
adjacent areas during the last decade. This has likely drawn down the water table thus
increasing salinity of the soil and resulting in large stretches of abandoned farmland with
salt-tolerant species recolonizing the formerly ploughed lands. Laguna Mormona itself is
now a halophytic scrubland that provides some habitat for wildlife but is floristically
depauperate: only a handful of species known to be halophytes were present. These habitat
alterations have apparently resulted in the local extirpation of nine taxa, five considered
globally rare, that were previously found here. Notably, several other species previously
reported from this region were located by Vanderplank (2011) only in riparian habitats,
suggesting that fresh water availability controls their distributions. The time-line of species
loss from Laguna Mormona suggest that the area was considerably less saline as recently
as the 1980s.
Two drainage systems or ‘arroyos,’ Rı́o Santo Domingo and Rı́o San Simon, provide
riparian habitat. Seven taxa that were not relocated occurred in these habitats. Rı́o Santo
Domingo was a historical locality for a small number of taxa that require permanent
wetland conditions (e.g., Hydrocotyle spp.). Arroyos in Baja California are by nature
dynamic systems with regular disturbance events from snow-melt and heavy rainfall, but
they are also heavily dependent on groundwater aquifers. The loss of species associated
with more permanent water (e.g., the two rare taxa of Hydrocotyle) suggests that there has
been a shift in the permanent water status of these aquifers.
Current farming practices are not only a threat to biological conservation, but also to
regional agriculture itself, as witnessed by the growing number of abandoned salinized
lowlands. For large-scale agricultural operations to continue, sustainability of local eco-
system services must be addressed. Soil and water quality in the region can be assessed for
condition and potential using the methodology proposed by Zalidis et al. (2002). The vast
tracts of abandoned farmland in the GSQ region indicate the need for more sustainable
agriculture practices if the valley is to remain productive. Agricultural practices must be
addressed as part of long-term conservation planning for this region, including impacts on
the region’s fresh water aquifers. Sustainable farming practices are essential for the long-
term maintenance of biodiversity in this region. Desalinization plants and waste-water
reuse have been shown to be a sustainable solution in arid lands to the south in Baja
California (Pombo et al. 2008). For desalinization to be successful and sustainable in GSQ,
the precise position of the desalinizers and the local handling of waste-water and discharge
must be carefully planned and implemented, taking into consideration the restricted ranges
of many rare and endemic organisms.

Conclusions

Identifying the causes of species extirpations is critical to our understanding of species

extinctions. In GSQ, destruction of vernal pools resulted in loss of 47 % of the species that
occurred in this habitat (Table 1). Changes in the aquifer affecting riparian areas and ponds
have resulted in loss of 10 % of the riparian flora and 23 % of rare plants known to have
occurred in these habitats in the San Quintı́n region. Agriculture continues to expand into
fragile habitats and agricultural water use in the San Quintı́n region is known to be

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exceeding sustainable levels. The resultant lowered water table and increased salinity in
the San Quintı́n valley have likely been the cause of the local extirpation of at least 24
species from the pond and riparian habitats.
These extirpations indicate that wetland habitats of San Quintı́n have already been
considerably altered by adjacent intensive agriculture. Monitoring of endangered taxa that
remain in the region is strongly recommended so that mitigation efforts can be designed
and implemented with adequate lead time. Further lowering of the water table can be
expected to lead to the extirpation of more species, particularly in vulnerable riparian
habitats. Current agricultural practices are unsustainable, having resulted in soil saliniza-
tion and species loss, and are likely to undermine future conservation efforts as well as
agricultural production if not directly addressed. This methodology is recommended as a
means to evaluate floristic loss in coastal areas worldwide.

Acknowledgments This work represents part of Vanderplank’s master’s research at Claremont Graduate
University and doctoral studies at the University of California, Riverside. Funding was generously provided
by The Jiji Foundation. Jon Rebman (San Diego Natural History Museum) generously assisted with plant
identifications and access to specimen data. Many people helped with field work, Vanderplank is particularly
grateful to Naomi Fraga, Sergio Mata, and Bart O’Brien for their continued support.

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