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45
Journal of Mammalogy, 100(1):45–54, 2019
DOI:10.1093/jmammal/gyz007
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 (http://creativecommons.org/licenses/
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
mammals
RJ. K,* MA.C. N, RP. Y, ST. T,
JM. N-M, JL. B, SJ. 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,
MACN,SJB)
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)
*Correspondent: Rosalind.kennerley@durrell.org
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 benet biodiversity,
but which faces signicant anthropogenic threats likely to detrimentally impact both species. We examined how
differences in habitats, forest structure, topography, and human activity inuence 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 beneciar 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
inuyen 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 Geográca. 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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46 JOURNAL OF MAMMALOGY
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 benets of PAs to
conservation, when properly enforced, are well documented
(Bruner etal. 2001; Rodrigues etal. 2004; Cantú-Salazar and
Gaston 2010; Coetzee etal. 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 etal. 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 etal. 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 etal. 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 etal. 2005; DeFries
etal. 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 scientic,
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 etal. 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 etal. 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 extremelyrare.
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 inuence pres-
ence of these species. We use our ndings to provide recom-
mendations for PA management that can benet 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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KENNERLEY ET AL.—HISPANIOLAN LAND MAMMALS IN PROTECTEDAREAS 47
distribution. If selected points proved unsafe to access, alterna-
tive randomly allocated points were selected.
Sierra de Bahoruco NP (1,125 km2, 18°10′N, 71°31′W, 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 stratied
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°49′N, 71°32′W, 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°16′N, 68°42′W,
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°01′N, 69°37′W, 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 Scientic Reserve (92 km2, 19°23′N, 70°08′W,
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 Scientic Reserve (23 km2, 19°19′N, 69°59′W,
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°32′N, 68°22′W, 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 20min 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
identiable 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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48 JOURNAL OF MAMMALOGY
hutia is semiarboreal, and no urine marks were detected during
surveys. Indirect signs of any age were used to conrm 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 classied 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 difcult 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 20cm from observer—
Brown etal. 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 > 10cm 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 etal. 2007; Hickey 2012). Ametric of surrounding for-
est cover was calculated based on 30-m resolution tree cover
data from 2000 (Hansen etal. 2013), which quanties 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 inuence 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,
respectively.
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, Mongabay.com/Tiffany Roufs); e) gnawed bark on
tree trunk.
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KENNERLEY ET AL.—HISPANIOLAN LAND MAMMALS IN PROTECTEDAREAS 49
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 reect 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 50cm 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 insufcient 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,
coefcients were averaged across all models with ∆AICc ≤ 2, in-
cluding zeroes as coefcients 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).
R
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; Table3).
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.
Explanatory
variables
Description Reason for inclusion
protected
area
(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 stratica-
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
closed
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
area
Mean tree basal area of the 10 clos-
est trees to the survey point with a
circumference > 10cm 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
road
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
etal. 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 50cm above the ground
Vegetation could affect soil conditions and therefore the invertebrate prey available to solenodons
(included in solenodon model only)
tree cover
(hutia)
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
(solenodon)
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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50 JOURNAL OF MAMMALOGY
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
R2
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.
D
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 reect 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 etal. 2006), which is often exacerbated for rare or
cryptic species (Gibson etal. 2007). We are condent 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
coefcients (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
K45555
wi0.36 0.19 0.18 0.14 0.14
Table 2.—Summary of plots by habitat classication 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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KENNERLEY ET AL.—HISPANIOLAN LAND MAMMALS IN PROTECTEDAREAS 51
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 20min. 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 identied 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 inuence 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
signicant 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
modication (Beever etal. 2003; Benítez-López etal. 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 identied 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 etal. 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
benecial 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
K65758
wi0.34 0.22 0.16 0.15 0.13
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52 JOURNAL OF MAMMALOGY
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
concern.
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
condently interpret the cause of this apparent reduction in uti-
lized habitats identied 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
etal. 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 signicantly decrease soil
fertility in the Dominican Republic (Templer etal. 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 reect 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 signicant 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 etal. 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 etal. 2004; Templer etal. 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 sufcient 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 etal. 2008), recent genetic
work has demonstrated that effective population sizes of hutias
are much smaller in the Dominican Republic (Brace etal. 2012;
Turvey etal. 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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KENNERLEY ET AL.—HISPANIOLAN LAND MAMMALS IN PROTECTEDAREAS 53
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 etal. 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 benet 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.
A
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
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