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The impact of habitat quality inside protected areas on distribution of the Dominican Republic’s last endemic non-volant land mammals


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The Hispaniolan solenodon, Solenodon paradoxus, and Hispaniolan hutia, Plagiodontia aedium, are the Dominican Republic’s only surviving endemic non-volant land mammals, and are high priorities for conservation. The country has an extensive protected area (PA) network designed to maintain habitats and benefit biodiversity, but which faces significant anthropogenic threats likely to detrimentally impact both species. We examined how differences in habitats, forest structure, topography, and human activity influence presence of solenodons and hutias across the Dominican Republic. Systematic surveys of seven PAs were undertaken to record indirect signs, with presence-absence data analyzed using a multi-model inference approach incorporating ecological variables from both field and GIS data. Solenodons were detected relatively frequently, whereas detections of hutias were uncommon. Lower elevations, increased surrounding tree cover, canopy closure, and reduced levels of low vegetation are all associated with increased probability of detecting solenodons, whereas agriculture and mangrove represent poor-quality habitat. Increased canopy closure, tree basal area (indicating older-growth forest), and increased rock substrate (providing more den sites) are associated with increased probability of detecting hutias. Our findings indicated that human activities within PAs are likely to negatively affect both species, and conservation activities should focus on preventing encroachment and conversion of forest to agriculture to maintain high-quality forest habitats.
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Journal of Mammalogy, 100(1):45–54, 2019
Published online 30 January 2019
© The Author(s) 2019. Published by Oxford University Press on behalf of American Society of Mammalogists.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (
by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.
The impact of habitat quality inside protected areas on distribution
of the Dominican Republic’s last endemic non-volant land
RJ. K,* MA.C. N, RP. Y, ST. T,
JM. N-M, JL. B,  SJ. B
Durrell Wildlife Conservation Trust, Les Augrès Manor, Trinity, Jersey JE3 5BP, British Channel Islands (RJK, RPY,JMN-M)
Centre for Agri-Environmental Research, University of Reading, Earley Gate, Reading RG6 6AR, United Kingdom (RJK,
Institute of Zoology, Zoological Society of London, Regent’s Park, London NW1 4RY, United Kingdom (MACN,STT)
Sociedad Ornithológica de la Hispaniola, Apto. 401 Residencial Las Galerías, Calle Gustavo Mejia Ricart No. 119 B, Santo
Domingo, Dominican Republic (JLB)
University of East Anglia, Norwich Research Park, Norwich NR4 7TJ, United Kingdom (SJB)
The Hispaniolan solenodon, Solenodon paradoxus, and Hispaniolan hutia, Plagiodontia aedium, are the
Dominican Republic’s only surviving endemic non-volant land mammals, and are high priorities for conservation.
The country has an extensive protected area (PA) network designed to maintain habitats and benet biodiversity,
but which faces signicant anthropogenic threats likely to detrimentally impact both species. We examined how
differences in habitats, forest structure, topography, and human activity inuence presence of solenodons and
hutias across the Dominican Republic. Systematic surveys of seven PAs were undertaken to record indirect
signs, with presence-absence data analyzed using a multi-model inference approach incorporating ecological
variables from both eld and GIS data. Solenodons were detected relatively frequently, whereas detections of
hutias were uncommon. Lower elevations, increased surrounding tree cover, canopy closure, and reduced levels
of low vegetation are all associated with increased probability of detecting solenodons, whereas agriculture and
mangrove represent poor-quality habitat. Increased canopy closure, tree basal area (indicating older-growth
forest), and increased rock substrate (providing more den sites) are associated with increased probability of
detecting hutias. Our ndings indicated that human activities within PAs are likely to negatively affect both
species, and conservation activities should focus on preventing encroachment and conversion of forest to
agriculture to maintain high-quality forest habitats.
El solenodonte de la Hispaniola, Solenodon paradoxus, y la hutia de la Hispaniola, Plagiodontia aedium,
son los únicos mamíferos endémicos terrestres no voladores que sobreviven en la República Dominicana, su
conservación es de alta prioridad. El país tiene una extensa red de áreas protegidas (AP) diseñada para mantener
hábitats y beneciar la biodiversidad, pero se enfrenta a amenazas antropogénicas. Sin embargo, no existen
datos cuantitativos para evaluar las presiones antropogénicas que amenazan a los solenodontes y las hutias.
Examinamos cómo las diferencias en los hábitats, la estructura del bosque, la topografía y la actividad humana
inuyen la presencia de solenodontes y hutias en toda la República Dominicana. Se realizaron encuestas
sistemáticas de siete AP para registrar los signos indirectos de ambas especies, los datos de presencia/ausencia
fueron analizados mediante inferencia multimodelo que incorpora variables ecológicas de los datos de campo y
Sistema de Información Geográca. Los Solenodontes se detectaron relativamente frecuentemente, mientras que
las detecciones de hutias fueron menos comunes. Las elevaciones más bajas, el aumento de la cubierta arbórea
circundante, el cierre del dosel y los niveles reducidos de vegetación baja se asocian con una mayor probabilidad
de detectar solenodones. Mientras que la agricultura y los manglares representan un hábitat de mala calidad
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para el solenodonte. Aumento del cierre del dosel, área basal del árbol (que indica un bosque más antiguo)
y un sustrato con mayor proporcion de roca (que proporciona más sitios para madrigueras) se asocian con una
mayor probabilidad de detectar hutias. Nuestros hallazgos indican que las actividades humanas dentro de las AP
pueden afectar negativamente a ambas especies. Las actividades de conservación deberían enfocarse en mantener
hábitats forestales de alta calidad por medio de prevenir la invasión y la conversión de los bosques a agricultura.
Key words: Caribbean mammals, Hispaniola, hutia, indirect eld signs, solenodon, systematic surveys
Establishment and maintenance of protected areas (PAs)
is a common approach for preserving important regions of
endemism and biodiversity. The role and benets of PAs to
conservation, when properly enforced, are well documented
(Bruner etal. 2001; Rodrigues etal. 2004; Cantú-Salazar and
Gaston 2010; Coetzee etal. 2014). The Caribbean is a glob-
ally important insular biodiversity hotspot, with 74% of 69
mammal species endemic to the region (Mittermeier et al.
2004; Anadón-Irizarry etal. 2012). Key Biodiversity Areas
(KBAs) contain over one-half of all threatened species in the
Caribbean, and 51% overlap partially or completely with PAs
(Anadón-Irizarry etal. 2012).
Hispaniola, divided politically into Haiti and the Dominican
Republic, is the second largest Caribbean island. The impor-
tance of the PA network in the Dominican Republic is high-
lighted by the fact that 18% (868,314 ha) of the country’s
land area is covered by KBAs, of which 88% is either com-
pletely or partially protected; in comparison, 13% (360,314
ha) of Haiti is covered by KBAs, but only 18% is protected
(Anadón-Irizarry etal. 2012). With 22% of land under strict
protection (IUCN and UNEP-WCMC 2014), the Dominican
Republic has among the highest percentage of PAs of any
country, despite being relatively poor and densely populated
(Holmes 2010). Like many tropical regions, continuing human
population growth places increasing pressure on natural eco-
systems, often leading to unsustainable land-use practices and
damage or loss of forest habitats (Foley etal. 2005; DeFries
etal. 2007). Although intensive exploitation and settlements
that alter the ecosystem of PAs in the Dominican Republic are
not permitted, their boundaries are permeable to encroach-
ment and settlement (Perdomo and Arias 2008). For exam-
ple, creation of infrastructure for development of scientic,
recreational, and tourist activities within PAs is permissible
(Ministerio de Medio Ambiente y Recursos Naturales 2004).
However, a lack of knowledge of regulations and permitted
uses of PAs in local communities, alongside limited enforce-
ment, facilitates ongoing environmental degradation through
resource extraction inside their boundaries, which can drive
declines or extinctions of species that PAs were designated to
protect (Baillie etal. 2004; Caro and Scholte 2007).
The Dominican Republic contains two surviving
native non-volant land mammals, the Hispaniolan hutia
(Capromyidae: Plagiodontia aedium), a large caviomorph
rodent, and the Hispaniolan solenodon (Solenodontidae:
Solenodon paradoxus), a eulipotyphlan insectivore. Both
species always have been considered rare and threatened
(Verrill 1907; Allen 1942; Fisher and Blomberg 2011) and
are currently listed as Endangered by IUCN (2018). They are
both also global conservation priorities based on evolution-
ary distinctiveness (Collen etal. 2011). Habitat loss, invasive
species, persecution, and hunting all are considered poten-
tial threats (IUCN 2018), but the ecology of both species is
poorly understood and available data about status, distribu-
tion, and threats are limited and contradictory. Both species
are considered dependent on native forest and predominantly
use limestone caves as denning sites, although hutias also
reportedly use tree cavities and solenodons reportedly use
fallen logs (Woods 1981; Ottenwalder 1985). Hutias are pri-
marily arboreal and herbivorous, feeding on leaves, fruit,
and bark, whereas solenodons are terrestrial, foraging in soil
and leaf litter for invertebrates (Woods 1981; Sullivan 1983).
Woods (1981) reported that hutias were habitat generalists,
potentially making them more resilient to human pressures,
whereas solenodons were more vulnerable to habitat change.
Ottenwalder (1985) described both species as widely distrib-
uted, but reported that solenodon populations were highly
fragmented and declining in number. Conversely, Sullivan
(1983) reported drastic reductions in hutia populations and
distribution associated with development and deforestation,
and although the species persisted in patches of appropriate
habitat, it was extremelyrare.
We conducted a large-scale eld survey to investigate the
ecology and distribution of Hispaniola’s native land mammals
to increase the conservation evidence base for both species.
Herein, we use the extensive data set generated by this survey
on occurrence of solenodons and hutias in seven PAs across the
Dominican Republic to examine how differences in habitat type,
forest structure, topography, and human activity inuence pres-
ence of these species. We use our ndings to provide recom-
mendations for PA management that can benet conservation
of both species.
M  M
Survey sites and sampling.Between March 2010 and June
2012, data on presence or absence of hutias and solenodons,
together with several measures of ne-scale habitat structure and
composition, were collected from 289 survey points across seven
national parks (NPs) or privately owned PAs in the Dominican
Republic, which together represent 31.2% of the area covered
by the country’s PA network. Selected PAs are widely distrib-
uted, and represent a broad range of habitats, vegetation types,
and topographic or climatic variables (Fig. 1). With only limited
prior knowledge of the distribution of both species, we attempted
to survey ≥ 15 points in each PA to capture variation in species
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distribution. If selected points proved unsafe to access, alterna-
tive randomly allocated points were selected.
Sierra de Bahoruco NP (1,125 km2, 18°10N, 71°31W, 300–
2,367 m elevation; 168 points collected between 5 March 2010
and 20 April 2011)is a mountainous area with diverse habitats
from dry broadleaf forest on lower slopes to wet broadleaf for-
est at higher elevations, which transitions at 1,100 m into pine
forest with shallow soils. Given its high elevational and habitat
variation, we invested extensive survey effort in this PA. In the
absence of high-quality vegetation maps, the PA was stratied
into 400-m elevational bands to ensure we surveyed all vegeta-
tion types, with ca. 20 points per stratum.
Jaragua NP (1,654 km2, 17°49N, 71°32W, 0–331 m eleva-
tion; 22 points collected between 19 July 2010 and 14 January
2011)is a lowland PA containing dry forest, mangroves, and
coastal wetlands. Del Este NP (428 km2, 18°16N, 68°42W,
0–60 m elevation; 16 points collected between 6 July 2010 and
17 June 2011)is another lowland PA containing broadleaf for-
est, karst forest, scrub, savannah, and wetlands. In both PAs,
we were able to stratify survey effort by vegetation type, and
allocated points proportionally to area of each stratum and in
randomly chosen locations (categories in Jaragua: low or no
vegetation cover, dry scrub, dry forest, broadleaf semi-humid
forest, mangrove; categories in Del Este: mangrove, semi-
humid broadleaf forest, broadleaf scrub).
Los Haitises NP (634 km2, 19°01N, 69°37W, 0–287 m ele-
vation; 40 points collected between 13 August 2011 and 23 June
2012) has irregular topography supporting tropical moist for-
est, karst forest, mangroves, wetlands, and coastal forest. Loma
Quita Espuela Scientic Reserve (92 km2, 19°23N, 70°08W,
100–985 m elevation; 19 points collected between 11 August
2011 and 19 December 2011)contains subtropical moist for-
est, cloud forest, rainforest, riparian forest, and wetlands. Loma
Guaconejo Scientic Reserve (23 km2, 19°19N, 69°59W,
0–606 m elevation; 19 points collected between 5 January 2012
and 11 January 2012)contains broadleaf forest, broadleaf scrub,
and pasturelands. Punta Cana Ecological Reserve (11 km2,
18°32N, 68°22W, 0–15 m elevation; ve points collected be-
tween 10 August 2010 and 11 August 2010)is a privately owned
low-elevation PA with coastal scrub and older secondary-growth
dry forest. Selection of points in these PAs was random.
Plot methodology.—Each plot was a 20-m-radius circle
(total area: 1,256 m2) around the survey point, within which the
following variables were recorded:
Mammal signs.As both target species have secretive noc-
turnal behaviors (Woods 1981), species presence was based
solely on indirect measures, with no attempts made to survey
using direct observation. All surveys were undertaken by a
team of ve experienced researchers. During daylight hours,
two researchers spent 20min searching for signs of each spe-
cies (Fig. 2). Solenodon presence was determined by presence
of distinctive conical holes (“nose-pokes”) made while foraging
for invertebrates in soil or leaf litter. Hutia presence was deter-
mined by evidence of feeding or gnawing on fruit, bark, and
leaves. Presence also was determined by feces, which is easily
identiable for both species (Mohr 1936–1938; Ottenwalder
1985). For other hutia species, urine marking is sometimes used
to detect presence (e.g., Howe 1974); however, the Hispaniolan
Fig. 1.—a) Map of the Dominican Republic protected area network (stippled gray), showing six surveyed National Parks (solid gray: 1, Del Este;
2, Jaragua; 3, Loma Guaconejo; 4, Loma Quita Espuela; 5, Los Haitises; 6.Sierra de Bahoruco) and one surveyed privately owned reserve (Punta
Cana Ecological Reserve, asterisk). b) Location of the Dominican Republic in the western Caribbean.
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hutia is semiarboreal, and no urine marks were detected during
surveys. Indirect signs of any age were used to conrm presence
of species within survey plots, as the aim of the study was to
understand presence of native mammals in different landscapes
rather than ne-scale temporal habitat or resource use. Evidence
of non-native mammal species was not recorded systematically.
Habitat measures.Dominant habitat was classied as:
broadleaf forest (including dry, semi-humid, and cloud forest),
pine forest, mangrove, agriculture (including plantations, pas-
ture, cultivated areas, and areas where cultivation had ceased
but signs of crop species remained present), and scrub (in-
cluding open grassland not used for pasture, areas of recent dis-
turbance with low vegetation, and dry or wet scrubland). Four
20-m transects were marked out in cardinal directions from the
survey point, and at 2-m intervals along these transects we re-
corded whether the point fell on rock or soil, and number of
vegetation touches by non-grass species in each 50-cm section
of a vertical 250-cm pole; these data were used to determine
percentage rockiness, and vegetation density and heterogeneity
(Willson 1974). Small caves and crevices that native mammals
might use for denning were not recorded as an additional pa-
rameter, because such features would be difcult to measure
or assess in the survey plot. Canopy closure was measured at
10 m along each transect using a canopy-scope (a 25-dot array
on a transparent screen held vertically 20cm from observer—
Brown etal. 2000), and calculated as the mean percentage of
points where sky was not visible (with 100% representing com-
plete canopy closure). Relative biomass was calculated as mean
basal area of 10 trees with > 10cm circumference at breast
height closest to plot center.
Remotely sensed and derived data.ArcMap 10 was used
(ESRI 2015). Point elevation was extracted from a 30-m reso-
lution ASTER Global Digital Elevation Model (DEM—METI
and NASA 2011). Distance to nearest sealed road was calcu-
lated from road data obtained from DIVA-GIS (Hijmans et
al. 2004), accounting for topographical variation rather than
Euclidean distance; detailed data on human settlements across
the Dominican Republic are not available, so this was used
as a proxy measure of likely human disturbance and isolation
(Blake etal. 2007; Hickey 2012). Ametric of surrounding for-
est cover was calculated based on 30-m resolution tree cover
data from 2000 (Hansen etal. 2013), which quanties canopy
closure for all vegetation > 5 m in height; the percentage of
cells around each point with > 75% canopy cover was calcu-
lated for a given species home range, using mean home-range
estimates for each species to calculate cell search radius (hutia:
184 m [n=12]; solenodon: 451 m [n=16]—Kennerley 2014).
Data analysis.The inuence of local and landscape-scale
characteristics on probability of detecting signs of solenodons
or hutias was explored using generalized linear mixed models
(GLMMs) with a binomial error structure (presence=1, ab-
sence=0) and a logit-link, with “PA identity” included as a
random effect, with separate analyses for each species. Points
in Loma Quita Espuela and Loma Guaconejo were excluded
from the hutia model because no signs of this species were
found in any plots or when travelling between points in these
PAs. Individual plots also were excluded from analyses if
data for any explanatory variables were unavailable, result-
ing in 234 and 269 points for hutia and solenodon models,
Fig. 2.—Hispaniolan solenodon (Solenodon paradoxus) eld signs: a) conical-shaped foraging “nose-pokes”; b) feces. Hispaniolan hutia
(Plagiodontia aedium) eld signs: c) gnawed fruit; d) chewed leaf and fecal pellets (photo, Roufs); e) gnawed bark on
tree trunk.
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For each species, a global model including all local and land-
scape-scale variables was tted (Table 1), before a model set of all
possible sub-models, ranked by Akaike’s Information Criterion
corrected for small sample size (AICc), was generated. For the
hutia model, the total number of vegetation touches for the entire
250-cm pole height (“veg250”) was used to describe vegetation
structure, to reect the arboreal nature of this species; for the so-
lenodon model, only the number of vegetation touches in the bot-
tom section of the pole 50cm above ground level (“veg50”) were
considered important for this terrestrial species. Only main effects
with no interaction terms were included, although “rockiness”
also was included as a quadratic term for both species, because
some rockiness might be necessary to provide denning sites, but
extensive rockiness could result in low soil availability and there-
fore insufcient invertebrates for solenodons and fewer trees for
hutias. Correlations between all pairs of potential explanatory
variables were considered, with no evidence of strong collinearity
(r<0.34 in all cases). All input variables were scaled to a mean of
zero and SD of 0.5 to allow direct comparison (Schielzeth 2010).
Pseudo-R-squared values were determined for global GLMMs,
where the marginal R2 represents variance explained by xed fac-
tors (R2
GLMM (m)) and the conditional R2 is interpreted as variance
explained by both xed and random factors (R2
GLMM (c)Nak-
agawa and Schielzeth 2013). To account for model uncertainty,
coefcients were averaged across all models with ∆AICc ≤ 2, in-
cluding zeroes as coefcients when variables were not included
in particular models (Burnham and Anderson 2002). The relative
importance of each predictor was calculated as the summed pos-
terior Akaike weight of models containing that predictor which
were included in the averaged model set (Burnham and Anderson
2002). All statistical analyses were performed in R v3.3.2 (R
Development Core Team 2016), using lme4 (Bates et al. 2012)
and MuMIn (Barton 2013).
Hutia.—Hutia signs were recorded at only 14 points across Del
Este, Jaragua, Sierra de Bahoruco, and Punta Cana. Signs were re-
corded from 15 to 2,019 m, but only in broadleaf and rarely pine
forest (Table 2). Data from Los Haitises were included in analyses
although hutia signs were not found within plots, because local hutia
presence was indicated by signs observed outside plots. The global
model had R2
GLMM (m)=0.61 and R2
GLMM (c)=0.70, indicating good
t, therefore strong likelihood of model-averaging outputs providing
high explanatory power. The total model set comprised 192 models,
with ve models considered highly plausible (ΔAICc ≤ 2; Table3).
All models in this subset included canopy closure, rockiness, and
tree basal area, with the top-ranking model including these three
variables exclusively. The other four explanatory variables (rocki-
ness2, veg250, elevation, distance to road) received weaker support,
with each only appearing in one of the top ve models. Probability
Table 1.—Descriptions of variables used in models to explain occurrence of Hispaniolan hutias (Plagiodontia aedium) and Hispaniolan solen-
odons (Solenodon paradoxus) across protected areas (PAs) in the Dominican Republic.
Description Reason for inclusion
(1) Del Este; (2) Jaragua; (3)
Loma Guaconejo; (4) Loma Quita
Espuela; (5) Los Haitises; (6) Sierra
de Bahoruco; (7) Punta Cana
Included as a random term in all models, because PA was a sampling unit with different survey stratica-
tion in different sites, and with nonindependence of locations. Hutia model excluded PAs (3) and (4)
canopy Amount of canopy (%); 0% com-
pletely open to 100% completely
Solenodons and hutias associated with older undisturbed forest (Woods 1981)
rockiness Measure of rockiness (%) Caves in rocks provide denning sites for both solenodons (Ottenwalder 1985) and hutias (Woods 1981)
tree basal
Mean tree basal area of the 10 clos-
est trees to the survey point with a
circumference > 10cm and within
the plot (m2)
Both species thought to be associated with older-growth forest, represented by larger tree basal areas
(Woods 1981)
elevation Elevation from sea level (m) Conditions are less favorable for solenodons at higher elevations (Ottenwalder 1985); hutias can be
found at most elevations (Woods 1981), but habitats at high elevations might provide poorer-quality diet
(Sullivan 1983)
distance to
Distance from the nearest signi-
cant road or track (m)
Both species are thought to be negatively affected by human presence due to persecution and increased
threat from dogs and cats associated with people (Woods 1981; Sullivan 1983; Ottenwalder 1985; Turvey
etal. 2014)
veg250 Total number of vegetation touches
in all sections of the 250-cm pole
Increased vegetation provides more food for hutias (included in hutia model only)
veg50 Total number of vegetation touches
in the rst 50cm above the ground
Vegetation could affect soil conditions and therefore the invertebrate prey available to solenodons
(included in solenodon model only)
tree cover
Based on the 2000 tree cover data,
percentage of cells within a 184-m
radius (mean diameter of hutia
home range) with > 75% tree cover
Hutias are sensitive to disturbance and degradation or fragmentation of natural habitat (Sullivan 1983)
(included in hutia model only)
tree cover
Based on the 2000 tree cover data,
percentage of cells within a 451-m
radius (mean diameter of solenodon
home range) with > 75% tree cover
Solenodons are associated with older undisturbed forest (Woods 1981) (included in solenodon model only)
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of detecting hutia signs increased with increasing canopy closure,
tree basal area, and amount of rockiness, but decreased with increas-
ing vegetation density, distance from nearest road, and elevation.
The rockiness2 term was negative, indicating that although proba-
bility of detecting hutia signs increased with rockiness, this had a
lessened effect at extreme levels of rockiness.
Solenodon.—Solenodon signs were detected in 89 plots across all
seven PAs, across a wide elevational gradient (13–2,026 m) and in
broadleaf forest, pine forest, and scrub (Table 2). The global model in-
dicated a low level of model t to the data, with R2
GLMM (m)=0.29 and
GLMM (c)=0.61. The total model set was reduced to ve models in
the ∆AICc ≤ 2 subset (Table 4). Canopy closure, elevation, tree cover,
and veg50 appeared in all of the top ve models, with the top-ranking
model containing these parameters plus distance to road and tree basal
area. Probability of detection increased with greater canopy closure and
higher tree cover in the wider landscape, but decreased with increas-
ing elevation and density of low-level vegetation. Of those predictors
receiving weaker support, probability of detection increased with
increasing tree basal area and distance to nearest road but declined with
increasing rockiness; the negative quadratic term indicates a greater
rate of reduction in probability at extreme levels of rockiness.
We used systematic surveys and quantitative analyses to in-
vestigate habitat associations of Hispaniola’s two surviving
endemic non-volant land mammals. Both solenodons and
hutias were more common at lower elevations and sites with
increased canopy closure and larger trees, suggesting they re-
quire older, more pristine forest. Differences in response to
landscape- or site-level features (e.g., surrounding tree cover,
rockiness) are likely to reect ecological differences between
the species in diet and environmental requirements. Hutia signs
were detected at far fewer sites, indicating this species might be
more patchily distributed (lower area of occupancy) or have a
reduced current-day geographic range (lower extent of occur-
rence) than solenodons. Global model t indicated that results
for hutias are robust, but results for solenodons must be inter-
preted with more caution.
A common problem in studies of species occurrence is the
ability to interpret analyses when uncertainty exists over detec-
tion (Hirzel etal. 2006), which is often exacerbated for rare or
cryptic species (Gibson etal. 2007). We are condent there was
a low probability of positive detection bias for either species,
because indirect signs are distinctive and could not be confused
with other species (Mohr 1936–1938; Ottenwalder 1985). The
potential for false negatives is more likely, and could occur
for two reasons. First, animals might use plots but not leave
signs, for example, if they move through sites but do not use
them for feeding or denning; such false negatives might vary
between species if they defecate at different rates while mov-
ing through sites. This is a particular issue with highly mobile
species (Thompson 2004). However, hutias and solenodons
Table 3.—Results of model selection and model-averaging procedures for explaining occurrence of Hispaniolan hutias (Plagiodontia aedium)
at plots (n=234) across seven protected areas in the Dominican Republic; plots with missing data for any explanatory variable excluded from
analyses. Models ranked in order of increasing AICc differences (∆AICc); K=number of parameters in each model. Model-averaged regression
coefcients (β; ± 95% CI) are averages of βi across all models with ∆AICc ≤ 2, weighted by each model’s Akaike weight wi. Calculations for β
include βi=0 when variables not present in given model. SE=standard error of β. wip=relative variable importance (sum of wi across all models
including that variable).
Model rank Model average
Variable 1 2 3 4 5 βSE wip
canopy 2.91 (0.69, 5.13) 1.13 1.00
rockiness 2.88 (0.46, 5.30) 1.23 1.00
tree basal area 2.10 (0.58, 3.61) 0.77 1.00
(rockiness)2 −0.34 (−2.53, 1.84) 1.11 0.19
elevation −0.22 (−1.72, 1.28) 0.76 0.18
veg250 −0.11 (−1.68, 1.45) 0.79 0.14
distance to road −0.05 (−0.71, 0.61) 0.34 0.14
∆AICc0 1.25 1.40 1.90 1.92
wi0.36 0.19 0.18 0.14 0.14
Table 2.—Summary of plots by habitat classication showing number and percentage of plots where Hispaniolan solenodons (Solenodon
paradoxus) and Hispaniolan hutias (Plagiodontia aedium) were present. Plots in Loma Quita Espuela and Loma Guaconejo excluded for hutia
(see text for details).
Solenodon Hutia
Habitat type Plots Plots with species % Plots Plots with species %
broadleaf 122 57 46.7 98 12 12.2
pine 104 31 29.8 104 2 1.9
mangrove 10 0 0.0 10 0 0.0
agriculture 32 0 0.0 24 0 0.0
scrub 21 4 19.0 15 0 0.0
TOTAL 289 92 31.8 251 14 5.6
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are both thought to be central-place foragers with relatively
static home ranges (Woods 1981; Sullivan 1983; Ottenwalder
1991), reducing the risk of false negatives. Second, animals
could be active in plots, but either signs may not be detected,
or characteristic behaviors are not consistently associated with
production of signs (Gu and Swihart 2004). One method to
reduce such errors is to undertake repeated measures of plots
(MacKenzie and Royle 2005). Unfortunately, this was not fea-
sible in this study because of time constraints, logistical chal-
lenges, and eld conditions. However, basing our analyses
on eld signs rather than direct observations counters these
issues to some extent. Hutias and solenodons live and move
around in close family groups (Woods 1981; Sullivan 1983;
Woods and Ottenwalder 1992), so signs might be expected to
be relatively numerous and more detectable if they are using an
area. Furthermore, signs of both species persist well under all
weather conditions, therefore effectively representing a cumu-
lative record of presence over several weeks: solenodon nose-
pokes last for ca. 2 weeks; hutia and solenodon feces lasts for
> 2 weeks in non-enclosed (i.e., non-cave) environments; and
hutia gnawing and other feeding signs are evident over much
longer periods (Hoy 2011; R.J. Kennerley, pers. obs.). To fur-
ther reduce between-site variation in detectability and min-
imize the risk of false negatives, standardized surveys were
conducted by the same team of skilled eld biologists familiar
with both species, with a relatively small survey area (1,256
m2) searched intensively for 20min. Nonetheless, we recognize
that negative effects of increasing vegetation density (veg50 for
solenodon, veg250 for hutia) on probability of detecting signs
of both species and the negative effect of increasing rockiness
of likelihood of detecting solenodon signs could be at least
partly indicative of reduced detectability of sign in these areas.
Signs recorded for both species were predominantly evidence
of foraging; thus, any identied habitat associations probably
are more closely associated with selection of foraging habitat
rather than den sites. Habitat requirements for these different
activities might differ in both species, and the inuence of the
spatial distribution of foraging and den sites, as well as popu-
lation density, on species’ detection needs further exploration.
The explanatory variables used in this study were chosen
based on hypotheses derived from the limited literature available
on Hispaniolan mammal ecology. Most of these variables were
measurable in the eld, with additional remotely sensed and de-
rived data also used. As data were unavailable regarding human
settlements across the Dominican Republic, distance to nearest
signicant road was used as a proxy for anthropogenic activity, as
presence of a road can make nearby land easier to access and hence
more likely to contain human settlements and resulting habitat
modication (Beever etal. 2003; Benítez-López etal. 2010). The
most recent available tree cover data for the Dominican Republic
are from 2000, making it possible that patterns of landscape-level
forest cover could have changed by the time eldwork was con-
ducted in 2010–2012. We may also not have identied all key
factors affecting native species distribution to include in our anal-
yses; for example, presence of invasive mammals such as black
rats (Rattus rattus), mongooses (Herpestes javanicus), or feral
cats and dogs could represent a competitive or predation threat
strong enough to displace native mammals from human-modi-
ed landscapes and perhaps even areas of good habitat (Sullivan
1983; Turvey etal. 2014, 2017). However, data quantifying such
threats across the Dominican Republic are currently unavailable.
Hutia.—Woods (1981) reported hutias occurred from sea
level to 2,000 m; we recorded hutias across a similar elevational
range. However, our models indicate that likelihood of hutia
presence declines with increasing elevation. Although hutias are
known to feed on a wide variety of plant species (Sullivan 1983;
Woods and Ottenwalder 1992), higher elevations may contain
fewer suitable food plants as vegetation changes from broadleaf
to pine forest. The strong positive relationship between increas-
ing rockiness and presence of hutias also is consistent with pre-
vious suggestions that existence of suitable cavities for den sites
is the most important requirement for good-quality hutia habitat
(Woods 1981; Sullivan 1983). However, our data suggested that
benecial effects may decline at extreme levels of rockiness; if
rock is the dominant substrate, quantity or quality of foraging
habitat may decline to a level that excludes hutias, potentially
due to fewer or less palatable trees being present. The mech-
anism underpinning the weak, counterintuitive observation that
hutia signs are more likely to be recorded closer to roads is un-
clear and requires further investigation.
Hutias previously were reported from numerous habitats
across Hispaniola, including dry subtropical, humid broadleaf,
Table 4.—Results of model selection and model-averaging procedures for explaining occurrence of Hispaniolan solenodons (Solenodon para-
doxus) at plots (n=269) across seven protected areas in the Dominican Republic; plots with missing data for any explanatory variable excluded
from analyses. Table arrangement and variables as in Table 3.
Model rank Model average
Variable 1 2 3 4 5 βSE wip
canopy 1.04 (0.30, 1.78) 0.38 1.00
elevation −2.84 (−3.98, −1.70) 0.60 1.00
tree cover 0.79 (0.02, 1.56) 0.39 1.00
veg50 −1.15 (−2.18, −0.11) 0.53 1.00
distance to road 0.54 (−0.20, 1.29) 0.38 0.85
tree basal area 0.47 (−0.32, 1.26) 0.40 0.78
rockiness −0.01 (−0.55, 0.55) 0.28 0.29
(rockiness)2 −0.14 (−1.00, 0.73) 0.44 0.13
∆AICc0 0.93 1.59 1.60 1.91
wi0.34 0.22 0.16 0.15 0.13
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pine, swamp, and oodplain forests (Sullivan 1983). Here,
hutias only were recorded in broadleaf and pine forest, with
no signs of presence in mangrove, agriculture, or scrub. Our
ndings show that hutias are particularly associated with old-
growth forest, with increased canopy closure, and tree basal
area associated with higher detection probabilities. It is unclear
whether this contradiction with previously reported habitat
associations represents a genuine contraction in distribution,
but given that our study represents the most extensive and ro-
bust survey for hutias in Hispaniola, any absence from habitats
from which they have previously been reported is cause for
Solenodon.—We detected solenodons across a wide eleva-
tional gradient (13–2,026 m), including in high-elevation pine
forest, which is only present above 1,100 m in the Dominican
Republic. These results are consistent with Ottenwalder (1985),
who reported that solenodons occur mainly at elevations below
1,000 m but can occur up to at least 1,500 m.Higher-elevation
environments containing pine forest might be less favorable
for solenodons because of cooler climate, poorer soils, and
moisture constraints. These characteristics are associated with
lower prey availability, while requiring more energy because of
cooler environments (Ottenwalder 1985).
Wet mangrove is not suitable habitat for a species that for-
ages in soil, but low detection of solenodon signs in agriculture
is more intriguing, particularly as it contradicts previous reports
that farmers in the Dominican Republic regard solenodons as
common (Woods 1981). As with hutias, it is not possible to
condently interpret the cause of this apparent reduction in uti-
lized habitats identied by our survey relative to reports from
the 1980s. It is possible that the range of solenodons has con-
tracted and the species has become largely or completely re-
stricted to forest habitats as a consequence of changes in scale
and intensity of farming in the Dominican Republic (Turvey
etal. 2017). The area of land under agriculture has increased,
and the types of crops grown and associated management
practices have changed during recent decades (Bravo-Ureta
and Pinheiro 1997; Raynolds 2002; González et al. 2009).
Agricultural activities are known to signicantly decrease soil
fertility in the Dominican Republic (Templer etal. 2005), and
absence of solenodons from farmland plots could be a conse-
quence of knock-on effects of reduced soil fertility and chang-
ing landscape and crop structure on abundance and availability
of potential solenodon prey species. The positive relation-
ship between presence of solenodons and increasing distance
from roads also could reect presence of better-quality hab-
itat further from this index of human disturbance, or because
such areas may have reduced levels of other human-associated
threats such as domestic and feral dogs and cats or other po-
tentially harmful invasive species. Previous research suggests
that dogs pose a particularly signicant predation threat to both
hutias and solenodons, and both species are persecuted as per-
ceived crop pests and also occasionally still hunted for food
(Sullivan 1983; Ottenwalder 1991; Turvey etal. 2014).
Secondary regrowth may represent a potentially suitable
habitat, as solenodons were detected reasonably regularly (19%
frequency) in scrub habitat. Abandoned agricultural land in the
Dominican Republic can quickly become reforested with native
vegetation, and forest soil properties and processes become
similar to those of undisturbed old forest sites after only a short
period (Martin etal. 2004; Templer etal. 2005).
Probability of detecting solenodon signs increases with
increasing tree basal area, canopy cover, and percentage tree
cover in the surrounding landscape, supporting previous reports
that their presence is associated with good-quality forest (Woods
1981). Solenodons den predominantly in rock clefts (Ottenwalder
1991), but our data indicated that increasing percentage of rocki-
ness reduced the likelihood of their presence. Their main prey are
invertebrates found in soil or leaf litter (Woods and Ottenwalder
1992; Ottenwalder 1999), and presence of sufcient soil for
foraging is likely to be important in determining occurrence of
solenodons, with increased levels of rockiness reducing prey
availability and foraging opportunity. Reduced probability of re-
cording solenodon signs in plots with denser vegetation could re-
ect reduced sign detectability, but also could arise because thick
vegetation at ground level can reduce prey abundance and hinder
access to soil invertebrates (Ottenwalder 1985).
Conservation implications.—Ottenwalder (1985) considered
that hutias were widespread in the Dominican Republic, and
our survey data indicate that overall this pattern remains true,
with hutias detected in several large PAs with no evidence of
overall geographical contraction in extent of occurrence com-
pared to previous distribution estimates (e.g., Sullivan 1983).
However, the limited frequency of hutia detections in our study,
which covered a substantial area of the Dominican Republic’s
PA network, suggests that this species is rare and localized in
the country. Although this apparent rarity could represent a re-
cent decline in population size (although not in overall range
extent), Sullivan (1983) also noted a lack of evidence of hutia
occurrence in areas of apparently suitable and undisturbed
habitat, and suggested that the species already was rare by the
early 1980s. Any decline in hutia populations might therefore
represent a historical rather than recent event, potentially even
associated with older human-caused disappearances of other
now-extinct Hispaniolan endemic mammals (Turvey 2009).
Whereas hutias appear to be more numerous than solenodons in
the Massif de la Hotte, Haiti (Turvey etal. 2008), recent genetic
work has demonstrated that effective population sizes of hutias
are much smaller in the Dominican Republic (Brace etal. 2012;
Turvey etal. 2016). Two of the smaller PAs were excluded from
analysis of hutias because no signs of the species were found,
indicating there may be a minimum patch size requirement that
we have not yet explored; thus, scale of habitat fragmentation
might be an important factor in determining presence or per-
sistence of hutias.
Solenodons previously were thought to be more threatened
than hutias, with populations considered to be highly frag-
mented and declining in number, despite reports from farm-
ers that the species can be locally common (Woods 1981;
Ottenwalder 1985). Our results indicate that solenodons are
in fact widespread and reasonably frequently detected (32%
frequency) across the areas that we surveyed. This result may
represent a genuine increase in distribution or abundance, or
alternately that the thorough systematic methodology used in
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our study provided a more accurate assessment of the distribu-
tion of solenodons across the Dominican Republic than in previ-
ous studies.
The Dominican Republic has an extensive NP network
(Holmes 2010), but due to poor enforcement and inadequate
regulation of activities within their boundaries, these PAs are
experiencing anthropogenically driven degradation of biodi-
versity, notably due to deforestation and increasing human
settlement (Perdomo and Arias 2008; Pasachnik etal. 2016).
Monitoring whether PAs continue to provide the necessary
habitat for native species in the face of changing environ-
ments and associated threats is fundamental to biodiversity
conservation, and to justify their continued long-term desig-
nation and management. Our study indicates that hutias may
require more intensive protection measures than solenodons,
due to their apparently more localized distribution and re-
stricted habitat associations. Management actions for both
species should focus on preventing human settlement and
encroachment within PAs and, in particular, improving pro-
tection of core areas of older high-quality forest, a move also
likely to benet many other native species on Hispaniola.
With a considerable proportion of the Dominican Republic
under strict protection, and escalating pressure on land out-
side PAs, these areas are likely to play an increasingly impor-
tant role for securing the future of Hispaniola’s last remaining
native land mammals.
Fieldwork was supported by Darwin Initiative project 17025
(“Building evidence and capacity to conserve Hispaniola’s
endemic land mammals”). We thank P. Martinez, R. and
L.Espinal, Y.and N.Corona, J.R.de la Cruz, and T.Bueno.
RJK was funded through Biotechnology and Biological
Sciences Research Council (BBSRC), the Durrell Wildlife
Conservation Trust, and the Zoological Society of London.
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... In addition, the Biological Reserve Loma Charco Azul is also located within the RBJBE [1]. This Reserve is highly biologically diverse, with species that include two endemic species of mammals of Hispaniola: the selenodont (Solenodon paradoxus) and the Hutia (Plagiodontia aedium), which are both highly threatened by habitat Sustainability 2021, 13,11030 3 of 20 deterioration due to anthropogenic pressure and the expansion, mainly from agricultural encroachment in these protected areas [12]. ...
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This paper assesses the Dominican Republic’s willingness to pay (WTP) for a conservation, restoration, and sustainable development program for the Bahoruco-Jaragua-Enriquillo Biosphere Reserve (RBJBE). Created in 2002, the RBJBE covers approximately 4858 km2, of which approximately 900 km2 correspond to the sea surface. With three core conservation zones, the RBJBE constitutes a complex space with a history of conflicts rooted in exploiting natural resources and social and economic issues. At the same time, it is a biodiversity hotspot of global importance. We present a Contingent Valuation Method to estimate the WTP for a conservation and restoration program called PROBIOSFERA. The non-parametric and parametric estimates of the WTP are DOP 165.00 (≈USD 2.94) and DOP 181.88 (≈USD 3.25), respectively. The socioeconomic variables that positively affect the WTP are income level, educational level, and age. The ecosystem services that are statistically related to WTP are the provisioning and supporting services. Regardless of the monetary valuation scenarios defined, the estimated annual monetized aggregated welfare associated with the RBJBE for Dominican society is DOP 7.2 billion (≈USD 128.1 million).
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Comparative assessment of the relative information content of different independent spatial data types is necessary to evaluate whether they provide congruent biogeographic signals for predicting species ranges. Opportunistic occurrence records and systematically collected survey data are available from the Dominican Republic for Hispaniola’s surviving endemic non‐volant mammals, the Hispaniolan solenodon (Solenodon paradoxus) and Hispaniolan hutia (Plagiodontia aedium); opportunistic records (archaeological, historical and recent) exist from across the entire country, and systematic survey data have been collected from seven protected areas. Species distribution models were developed in maxent for solenodons and hutias using both data types, with species habitat suitability and potential country‐level distribution predicted using seven biotic and abiotic environmental variables. Three different models were produced and compared for each species: (a) opportunistic model, with starting model incorporating abiotic‐only predictors; (b) total survey model, with starting model incorporating biotic and abiotic predictors; and (c) reduced survey model, with starting model incorporating abiotic‐only predictors to allow further comparison with the opportunistic model. All models predict suitable environmental conditions for both solenodons and hutias across a broadly congruent, relatively large area of the Dominican Republic, providing a spatial baseline of conservation‐priority landscapes that might support native mammals. Correlation between total and reduced survey models is high for both species, indicating the substantial explanatory power of abiotic variables for predicting Hispaniolan mammal distributions. However, correlation between survey models and opportunistic models is only moderately positive. Species distribution models derived from different data types can provide different predictions about habitat suitability and conservation‐priority landscapes for threatened species, likely reflecting incompleteness and bias in spatial sampling associated with both data types. Models derived using both opportunistic and systematic data must therefore be applied critically and cautiously. Comparative assessment of the relative information content of opportunistic historical records and survey data was conducted for Hispaniolan solenodon and Hispaniolan hutia, to evaluate whether these different data types provide congruent predictions of species distributions. Correlation between survey models and opportunistic models is only moderately positive, likely reflecting incompleteness and bias in spatial sampling, and indicating that models derived using both data types must be applied critically and cautiously.
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Conservation decision-making for threatened species in human-modified landscapes requires detailed knowledge about spatial ecology, but robust data derived from tracking individual animals are often unavailable, with management decisions potentially based on unreliable anecdotal data. Existing data are limited for Hispaniola's two threatened non-volant land mammals, the Hispaniolan hutia (Plagiodontia aedium) and Hispaniolan solenodon (Solenodon paradoxus), with assumptions that hutias are better able to tolerate landscape disturbance. We collected spatial behaviour and habitat use data for Hispaniolan mammals during a multi-year field programme across undisturbed and modified habitats in southwestern Dominican Republic, using GPS units for hutias (11 individuals) and radio-telemetry for solenodons (22 individuals). Although significant differences exist in hutia home range estimates between different GPS error derivation strategies and estimated terrestrial/arboreal behaviour scenarios (95% KDE means = 23,582–28,612 m²), hutias almost exclusively use forest under all estimates (mean observations in forest across all strategies/scenarios = 90.3%, total range = 69.1–100%). Solenodons have larger estimated home ranges (95% KDE mean = 156,700 m²), with differences between wet and dry season estimates, and show much more variation in habitat use than hutias within the same landscape; animals regularly use both forested and modified habitats, being observed most frequently in forest (mean = 74.0%, range = 13.0–99.1%) but also occurring regularly in pasture (mean = 15.9%, range = 0–80.0%) and cropland (mean = 7.7%, range = 0–62.0%), and den in all three habitats. This new baseline on Hispaniolan mammal spatial ecology challenges anecdotal data, and suggests solenodons may be better able to tolerate disturbance and persist in modified landscapes.
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To determine the evolutionary history, relationships and distinctiveness of allopatric populations of Hispaniolan solenodon (Solenodon paradoxus), a highly threatened Caribbean ‘relict’ mammal, to understand spatio-temporal patterns of gene flow and the distribution of diversity across complex large island landscapes and inform spatial conservation prioritization. Historical and modern-day solenodon specimens were analysed from sampling localities across Hispaniola, representing this geotectonically complex island's distinct northern, south-eastern and south-western biogeographic provinces. We successfully amplified mitochondrial cytochrome b and control region sequences from 34 solenodon samples. Bayesian phylogenetic analyses were applied to assess the relationship between these sequence data, and coalescent simulation and approximate Bayesian computation were used to identify which model of solenodon intra-island demographic history best explains the observed patterns of variation. We also conducted morphometric analysis of 110 solenodon specimens to investigate whether allopatric Hispaniolan populations can be differentiated using craniodental characteristics. Unique haplotypes were identified in solenodon samples from each biogeographic region, with no haplotype sharing between regions. Higher marginal posterior probability values were found for a three-population model comprising allopatric northern, south-eastern and south-western Hispaniolan populations, with exceptionally low migration rates inferred between all populations, indicating that they are genetically isolated. Modal estimates of long-term effective female population size are extremely low for south-western and south-eastern populations. Morphometric differentiation is observed between all three populations. Evolutionary differentiation of Hispaniolan solenodons into three distinct populations is congruent with phylogenetic patterns observed in several other Hispaniolan species, with population isolation possibly associated with past marine transgression. We interpret these populations as distinct subspecies, with the two genetically impoverished southern subspecies particularly vulnerable to environmental change. Our improved understanding of Hispaniolan solenodon evolutionary history provides an important baseline for identifying wider patterns of intra-island diversification and prioritizing conservation attention for evolutionarily significant populations.
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The Caribbean Islands Biodiversity Hotspot is exceptionally important for global biodiversity conservation due to high levels of species endemism and threat. A total of 755 Caribbean plant and vertebrate species are considered globally threatened, making it one of the top Biodiversity Hotspots in terms of threat levels. In 2009, Key Biodiversity Areas (KBAs) were identified for the Caribbean Islands through a regional-level analysis of accessible data and literature, followed by extensive national-level stakeholder consultation. By applying the Vulnerability criterion, a total of 284 Key Biodiversity Areas were defined and mapped as holding 409 (54%) of the region’s threatened species. Of these, 144 (or 51%) overlapped partially or completely with protected areas. Cockpit Country, followed by Litchfield Mountain - Matheson’s Run, Blue Mountains (all Jamaica) and Massif de la Hotte (Haiti) were found to support exceptionally high numbers of globally threatened taxa, with more than 40 such species at each site. Key Biodiversity Areas, building from Important Bird Areas, provide a valuable framework against which to review the adequacy of existing national protected-area systems and also to prioritize which species and sites require the most urgent conservation attention.
The insular Caribbean is among the few oceanic-type island systems colonized by non-volant land mammals. This region also has experienced the world’s highest level of historical mammal extinctions, with at least 29 species lost since AD 1500. Representatives of only 2 land-mammal families (Capromyidae and Solenodontidae) now survive, in Cuba, Hispaniola, Jamaica, and the Bahama Archipelago. The conservation status of Caribbean land mammals is surprisingly poorly understood. The most recent IUCN Red List assessment, from 2008, recognized 15 endemic species, of which 13 were assessed as threatened. We reassessed all available baseline data on the current status of the Caribbean land-mammal fauna within the framework of the IUCN Red List, to determine specific conservation requirements for Caribbean land-mammal species using an evidence-based approach. We recognize only 13 surviving species, one of which is not formally described and cannot be assessed using IUCN criteria; 3 further species previously considered valid are interpreted as junior synonyms or subspecies. Of the 12 reassessed species, 5 have undergone a change in threat status since 2008, with 3 species (Capromys pilorides, Geocapromys brownii, Mesocapromys angelcabrerai) increasing in extinction risk by 1 IUCN category, and 2 species (Plagiodontia aedium, Solenodon paradoxus) decreasing in extinction risk by 2 categories. Only 1 change in threat status represents a genuine change; all other changes are mainly associated with new information becoming available. Hunting, habitat loss, and invasive species represent major threats to surviving species, and conservation of the highly threatened Caribbean land-mammal fauna will require a range of targeted management strategies.
The extent to which human activity has influenced species extinctions during the recent prehistoric past remains controversial due to other factors such as climatic fluctuations and a general lack of data. However, the Holocene (the geological interval spanning the last 11,500 years from the end of the last glaciation) has witnessed massive levels of extinctions that have continued into the modern historical era, but in a context of only relatively minor climatic fluctuations. This makes a detailed consideration of these extinctions a useful system for investigating the impacts of human activity over time. This book describes and analyses the range of global extinction events which have occurred during this key time period, as well as their relationship to both earlier and ongoing species losses. By integrating information from fields as diverse as zoology, ecology, palaeontology, archaeology, and geography, and by incorporating data from a broad range of taxonomic groups and ecosystems, this text provides a fascinating insight into human impacts on global extinction rates, both past and present.