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Article

Plant Diversity in a Volcanic Crater Interior: Laguna De Apoyo Nature Reserve, Nicaragua

by
Jeffrey K. McCrary
1,
Alain Kheim Meyrat
2,
Ricardo M. Rueda
2 and
Luz Maria Calvo-Irabien
3,*
1
Biodiversity Research Group, Universidad Nacional Autónoma de Nicaragua-Managua, Apdo. Postal #663, Managua 14172, Nicaragua
2
Independent Researcher, León 21000, Nicaragua
3
Centro de Investigación Científica de Yucatán, Unidad de Recursos Naturales, Mérida 97205, Yucatán, Mexico
*
Author to whom correspondence should be addressed.
Conservation 2025, 5(4), 83; https://doi.org/10.3390/conservation5040083
Submission received: 17 October 2025 / Revised: 24 November 2025 / Accepted: 3 December 2025 / Published: 12 December 2025
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Abstract

Volcanic crater interiors in Nicaragua’s Pacific region are a valuable, understudied, and threatened native plant resource. Laguna de Apoyo Nature Reserve encompasses a crater and lake formed following a Quaternary volcanic explosion in Pacific Nicaragua. The flora of the tropical dry forest in the crater’s interior surrounding the lake has not been extensively assessed. We identified 403 native and 72 introduced plant species and their uses through a combination of survey plots, unstructured interviews, expert consultations, and targeted searches for plant species coordinated with key informants in the Reserve. Fabaceae, Euphorbiaceae, and Asteraceae were the most represented native species, whereas the most significant numbers of introduced species were found in Poaceae, Euphorbiaceae, and Fabaceae. Forty-one species have conservation priority status. Documented uses were found for 70% of the native species and 88% of the introduced species. The most significant numbers of plant species with reported use types were ornamentals and fuelwood. This study constitutes the most comprehensive plant species inventory in a protected area of Nicaragua’s tropical dry forest biome. These findings indicate native plant diversity is high, introduced species pose considerable risks, and most species are integrated into local uses. Consequently, management decisions should explicitly promote native diversity, protect threatened species, better control introduced species, and encourage sustainable use.

1. Introduction

How, or even whether, protected area policies protect biodiversity in Mesoamerica has been a contentious topic for decades. In Mexico, for instance, many species of conservation importance have known populations only outside of protected areas. Within protected areas, some species with protection needs are in population decline, whereas cosmopolitan species are on the rise [1]. In Costa Rica, where the vast majority of mammal species have been documented within protected areas, a quarter of the mammal species are considered under-protected [2]. In the flagship protected area of northern Nicaragua, Bosawas Biosphere Reserve, the expansion of the agricultural frontier into forests traditionally controlled by indigenous groups has caused substantial habitat loss in recent decades [3].
Poverty and weak governance combine to create complex pressures on the resources and biodiversity present in the Mesoamerican protected areas [4]. Small national size intensifies these pressures. Biodiversity protection is particularly problematic in areas suited to human activities. The tropical dry forest biome is attractive for human inhabitation and farming, yet only 3.3 [5] to 4.5% [6] of its area in Mesoamerica is protected. Four forms of forest habitat loss are in tropical dry forests: urbanization, conversion to cropland and to pastureland, and deforestation [7]. Even without these large-scale events, biodiversity can decline because of smaller-scale forest use that reduces ecosystem function and connectivity or degrades habitats [8]. Loss and degradation in tropical dry forests diminish their ecosystem benefits and resilience to climate change [9].
Nicaragua’s National Protected Areas System (SINAP) manages 76 protected areas, covering over 15% of the national territory [10]. Still, only around 5% of the tropical dry forest lowlands (below 400 masl) is protected. This forest type is segmented into disconnected protected areas spread through the Pacific region [9,11] and is among the most fragmented and rapidly deforested in the western hemisphere [12].
The classical dynamic of the burning-agriculture-cattle progression in the agricultural frontier, eloquently depicted by Belt [13], does not adequately characterize current dynamics in tropical dry forest remnants in the Pacific region of Nicaragua. By the end of the twentieth century, stands of closed-canopy forest in this biome in Nicaragua were remnants from logging operations followed by farming expansion. Many forests in the protected areas, including both lowland tropical dry forest and montane dry forest consist largely of mosaics of farms, woodlots, and forests of varying degrees of degradation [14,15].
Today, Nicaragua’s tropical dry forest is important to both rural and urban people [16]. Forest use can change tree composition in and near protected [15,17]. In general, the biodiversity protection and conservation strategies within SINAP in Nicaragua are limited to a set of laws and regulations prohibiting hunting and extraction of biological and other materials, forest fires, alterations to habitat, and introduction of non-native species, including the principle of “rational and sustainable use of natural ecosystems” [18].
The lowland tropical dry forest region of Nicaragua, covering most of the western half of Nicaragua [9], is blessed with dozens of volcanoes, some of which contain unique closed lakes in the craters. All these volcanic crater lake systems have been declared protected areas within the SINAP network [19]. The catchments of these lakes in the volcanic crater interiors are further protected by special regulations that prohibit the removal of plant material for commercial purposes, all industrial and forestry activities, and the introduction of non-native species [20]. The use of forest resources for non-commercial, household consumption may be permitted under specific concessions, permits, licenses, and quotas [18]. Understanding how well these rules protect strategic biological resources in tropical dry forest requires knowledge of the forest composition as well as of the intimate, dynamic relationships between people and the forests they use.
The fragility of tropical dry forest, sensitive to invasions by introduced species [21], to forest fires, and to overexploitation of botanical resources for fuelwood and other forest products, puts at risk both biodiversity and environmental services such as groundwater supplies [9]. Protected areas offer the potential to preserve populations of threatened plant species and forest integrity [22,23].
One of the protected areas in Nicaragua with a volcanic crater lake is Laguna de Apoyo Nature Reserve (LANR), situated in the departments of Masaya and Granada in the Pacific region of Nicaragua (Figure 1). The entirety of LANR lies in the hot and seasonally dry Pacific lowlands of Nicaragua, in tropical dry forest biome, almost all of it is located below 400 masl [9,11]. The protected area is nestled between urban centers, with over 100,000 people living within 3 km of the crater [24]. The interior of the crater mainly consists of steeply sloped land (average 30% slope), formed by a major pyroclastic explosion approximately 23,000 years ago, in the Pacific region of Nicaragua [25]. The protected area, designated as part of SINAP in 1991, has been managed under a management plan approved in 2010, without any update since then [26].
The principal geographical feature of LANR is its lake, which has attracted biodiversity studies focused on the irreplaceable biodiversity of endemic, threatened fish species [27] and its aquatic malacofauna [28]. In contrast, terrestrial animal biodiversity of LANR has been limited to studies of malacofauna [29], Lepidoptera [30], howler monkeys [31], and birds [32]. Botanical studies have been limited to plant uses [24,33,34].
During the past two decades, the traditional forest uses by households in and around LANR have faced a dramatic competition from tourism [35]. LANR is situated in the midst of traditional population centers, dating back to before Euro-descendant colonization [36]. As populations and land use demands have increased, most of the land in Nicaragua’s Pacific Region has been deforested, placing ever-greater pressure on LANR to provide forest products. As documented previously, the extraction of forest products for domestic use and homeworker enterprises, principally for fuelwood and some artisan products [24,33,34], along with wholesale land clearing for tourism development [37], has increased in recent decades.
This study demonstrates that the LANR crater interior is a vital repository of plant biodiversity within Nicaragua’s tropical dry forest. It highlights the underestimated threat of introduced plant species to native diversity and recognizes opportunities to engage local inhabitants in conservation strategies through regulated plant use.

2. Materials and Methods

Although the volcanic crater interior in LANR is largely forested, it is mainly composed of private properties, which present numerous management challenges due to competing land uses [38]. The entirety of the crater interior is included in the core area of LANR, where strict limitations are applied by the management plan in effect [26]. The buffer zone encompasses farms, woodlots, and villages in the surrounding areas. The obvious fragility of soils due to erosion from steep slopes and the increasing pressure among competing resource uses within the crater [24,37] motivated our interest in the poorly known plant biodiversity.
We conducted observations of terrestrial vascular vegetation in the volcanic crater interior located within LANR, between 2004 and 2017. Plant species were surveyed on twenty-two plots at several locations in the crater interior of LANR (Table S1). These plots ranged from 200 to 1000 m2 and had irregular shapes, reflecting the uneven geography of the LANR crater interior and private property boundaries.
Additional plant species information was documented in 40 daily surveys without plot structure, conducted throughout the crater interior, including mature and young forests, former pastures, riparian and lakeshore areas, agricultural areas, and human settlements. Survey locations were limited to public access areas and those where specific permission was obtained from landowners.
Plant species were identified in the field using dichotomous keys [39,40]; confirmations were made by comparisons of plant samples and photos to voucher material at the University of Central America Herbarium and to the online database for Nicaraguan vegetation managed by the Missouri Botanical Garden [41]. We followed the Missouri Botanical Garden database for plant distribution, nomenclature and systematics, and for assignment of the conservation status of the plant species found in the inventory [41].
Approximate species abundances, estimated from plots and consultations with technicians and key informants from the local communities, were categorized according to the following rubric of individuals per km2: rare, less than 2; limited, 2–10; occasional, 10–20; frequent, 20–200; abundant, more than 200.
Plant uses were documented using pilot group interviews with local informants, incidental observations, expert consultations (including staff of the University of Central America Herbarium, R. Rueda, A.K. Meyrat, and project technicians), and previous studies [24,33,34,41,42,43]. The recorded uses of the plants were classified into the following categories: ornamental; food; fuelwood; living fence; broomstick; medicinal; melliferous; shade trees used for understory cultivation; construction; forage; assorted artisan products.

3. Results

3.1. Plant Biodiversity in LANR

We documented 497 plant taxa of which 496 were identified to species, among 110 families (Tables S2 and S3; Figure 2). These results encompass 8.6% of the 5796 species and 4.9% of the 225 families of seed plants reported for Nicaragua [41]. Twenty-one taxa were observed only under cultivation and were not included in the following analysis (Table S3).
Only Angiosperms (397 native species, 72 introduced), Ferns (Polypodiopsida: 5 native, none introduced), and Mosses (Lycopodiopsida: 1 native, none introduced) were documented to species level, and Gymnosperms were completely absent in our surveys (Table S2). Ninety-seven Angiosperm families were represented among the native species, with Fabaceae (55 species), Euphorbiaceae (26), Asteraceae (19), and Malvaceae (18) having the most significant number of species. Fifty-eight families were represented with only one or two native species. One hundred seventy-one native species were present at population densities estimated at greater than 20 individuals per km2, in 66 families, of which the most represented families were Fabaceae (28 species), Euphorbiaceae (9), and Asteraceae (8); (Figure 3).
Among the 72 introduced species, there were 35 families. The most prominently represented families were Poaceae (14 species), Euphorbiaceae (8), and Fabaceae (5). Sixteen of the introduced species in 12 families had population densities above the 20 individuals per km2 threshold, of which Fabaceae, Euphorbiaceae, Anacardiaceae, and Poaceae had two species each (Figure 3).
There were no country first reports, nor were there any latitudinal or longitudinal range extensions of any plant species in our study. Nonetheless, there were first reports of 84 species in the Masaya Department-61 native and 21 introduced species-and first reports of two native species in the Granada Department. There were 159 species of trees from 47 families, of which 133 species were native (83.6%), distributed among 43 families. The most highly represented family of native tree species was Fabaceae, with 29 species, as it was also among introduced species, with four of the 26 total introduced species (Table S2).

3.2. Useful Plants in LANR

Thirty per cent of the native species do not have a documented use. Among the 280 species with uses, 44% had only one reported type of use, and only 1.5% had five or more use types. The most abundant use types among the plants were ornamental, fuelwood, food, and medicinal, with 102, 95, 58, and 56 species, respectively. Trees were the growth type most commonly associated with several use types: fuelwood, food, construction materials, furniture, agroforestry, and artisan products. On the other hand, all growth types were found in ornamental, medicinal, melliferous, and forage species (Figure 4a). The tree Spondias mombin had the broadest range of use types (6); Hymenaea courbaril, Gliricidia sepium (Figure 2), Cordia gerascanthus, Samanea saman and Enterolobium cyclocarpum, all trees, had five use types; the palm Bactris guineensis had four. Shrubs with the greatest number of different uses were Acanthocereus tetragonus and Solanum erianthum, each with three use types. Only one herbaceous species, Bromelia pinguin, was found to have as many as three use types (Table S2).
Fourteen per cent of the introduced species had no reported use, and 64% had only one use. Only three species were reported with three use types: Terminalia catappa, Tamarindus indica, and Azadirachta indica (Table S2). As in the case of native species, the most frequent use reports for introduced species were ornamental, food, and medicinal (Figure 4b).
Forty-one of the native species found in the inventory were assigned a conservation status [41] (Table 1). The shrub Euphorbia leucocephala (EN/VU), the epiphyte Hylocereus costaricensis (VU), and the herbs Pseuderanthemum alatum (VU/NT) and Vernonia patens (VU/NT) are the only four native species in the Near Threatened/Vulnerable or higher risk category. The first and third of these are used as ornamentals, while the second has commercial value as a food item.

4. Discussion

This research presents the first botanical inventory focused on a volcanic crater interior in Nicaragua, which is significant because these areas constitute a substantial portion of the dry tropical forest habitat under protection in the country. As was expected, the taxa encountered were indicative of the dry tropical forest habitat of Nicaragua.

4.1. Native Plant Biodiversity

Very few protected areas in the tropical dry forest in Nicaragua enjoy extensive botanical surveys, and most publications present only tree species diversity within a limited number of fixed plots. Although botanical surveys have been conducted in the tropical dry forest in Río Escalante-Chacocente Wildlife Refuge, located approximately 50 km to the south of LANR, all have focused only on trees [14,44,45]. Tree diversity and conservation have been evaluated in the tropical dry forest of the Miraflor Highlands Protected Landscape [15] and in nearby remnant forests outside protected areas [17]. A list of 303 plant species, including 77 trees, was developed in a private protected area that is not part of the SINAP system [46].
Our results were consistent with those of a recent study of tree diversity, using fixed survey plots, in northern Nicaragua [17]. Twenty-nine of the tree families found in the mentioned study were also found in our study, with notable exceptions Vochysiaceae, Fagaceae, and Asteraceae, the first two of which we do not expect in LANR. The much greater tree species diversity in our study compared to the mentioned study [17], with 159 vs. 84 plant species, may reflect differences in methodology and study site. Our significantly larger species list reflects an advantage of using combined field plant diversity assessment techniques to provide a broader view beyond dominant species in tree diversity.
In another study, tree diversity was evaluated in fixed survey plots in the montane tropical dry forest of the Miraflor Highlands Protected Landscape, located in northern Nicaragua. Of the 72 identified species, 40 are found in LANR, with notable exceptions largely among highland species in the families Clusiaceae, Fagaceae, Moraceae, and Myrtaceae [15].
As in the previously mentioned studies, the number of tree species we documented in the LANR crater interior also exceeded that found in a study using fixed survey plots in seven other tropical dry forests in the Pacific region of Costa Rica and Nicaragua [44]. Among the five most abundant species, at each of the five sites studied in Nicaragua, we documented all except three in the crater interior of LANR. Additionally, all the families identified in their study of the Pacific region of Nicaragua matched those found in LANR. For example, we did not record the presence of Myriocarpa bifurca in LANR, even though it was a prominent species noted in the study conducted at Masaya Volcano National Park, less than 10 km from LANR.
Our inventory largely coincided with that of a study of trees in a tropical dry forest area in succession near LANR [45], with some noteworthy points. That study focused on successional stages formerly used for crops and cattle, which could share characteristics with several areas within the volcanic crater of LANR. However, we did not document five species that the mentioned authors found: Lonchocarpus acuminata, Hippocratea rosea, Bonellia nervosa, Acaciella angustissima, and Eugenia salamensis.
A botanical survey of Reserva Natura, a private property approximately 40 km northwest of LANR in lowland tropical dry forest, which is not part of the SINAP system, yielded similar results to our survey [46]. Of the 303 plant species found in this area, 181 were documented in our survey as well. As in our study, the families with greatest number of taxa present in this location were Fabaceae, Malvaceae, Euphorbiaceae, and Asteraceae, which closely matched our results in LANR.
Our plant species inventory, uniquely extensive for a protected area in Nicaragua’s tropical dry forest, is nonetheless a work in progress that is not exhaustive. Although we did not document any species not previously reported in Nicaragua, we report 61 native species not previously reported within the Masaya Department and two native species not previously reported in the Granada Department [41]. Our results suggest that the Masaya department is considerably understudied as a botanical resource.
As is demonstrated in Table 1, dozens of plant species are regarded as having elevated conservation importance on the national level [41]. However, some species included in this survey warrant additional mention for conservation concerns beyond the criteria of the mentioned Missouri Botanical Garden database.
Dalbergia retusa is listed on the IUCN Red List as Critically Endangered [47], and also on Appendix II of CITES [48]. Swietenia humilis (Figure 2) is listed as Endangered on the IUCN Red List [49] and is included in Appendix II of CITES [48]. Harvest of S. humilis is banned nationally [50]. Cedrela odorata is found on the IUCN Red List as Vulnerable [51], is on Appendix II of CITES [48], and its harvest is also banned within protected areas [50]. Ceiba pentandra is also forbidden for harvesting in Nicaragua, and Pachira fendleri (noted as Pachira quinata) is banned for harvesting within protected areas [50]. All 15 species of Orchidaceae are listed in Appendix II of CITES [48], of which four are listed as Near Threatened nationally (Table 1), but none are in the IUCN Red List. Sideroxylon capiri ssp. capiri and Yucca guatemalensis, both listed as Near Threatened nationally (Table 1), are also listed in the IUCN Red List as Near Threatened [52] and Data Deficient [53], respectively. Carica papaya is listed as Data Deficient in the IUCN Red List, likely because of the difficulty in distinguishing cultivars from wild populations [54].
The extremely low documented population of D. retusa could be attributed to its exceptionally fine wood prized in artisanry, which provides marketing opportunities locally for even small sizes of wood [55]. Only P. fendleri, S. capiri ssp. capiri, C. papaya, and Y. guatemalensis were documented at relatively high abundances; populations of the latter two could be influenced positively by human activity through plantings as food and living fences, respectively.

4.2. Introduced Species

The range of introduced species in the LANR crater interior (72 introduced species in 35 families), excluding those strictly confined to planted and cultivated areas, is startling and calls for targeted eradication or control measures and ongoing monitoring. Seventeen of these species, from 13 families, were found at densities exceeding 20 stems per km2, showing the widespread penetration of introduced species. The presence of several Poaceae species, such as Johnson grass (Sorghum halepense), in disturbed areas fuels the seasonal fires, which locals promote to clear brush and access fuelwood [26].
Among the 26 introduced tree species, seven were found at high densities, all of which are actively planted as fruit, shade, and decorative trees. Two of them, Azadirachta indica and Anacardium occidentale, were planted within the protected area with technical and financial support from an international reforestation organization [24].
The prohibition on species introductions in LANR was strengthened in 2010 with the implementation of the management plan [26]. Human inhabitation, particularly along the lakeshore within the LANR crater, often involves invasive species introductions [56]. The advent of tourism activity has brought with it another wave of species introductions, with herbs such as Chrysopogon zizanioides, a cultivated species that could become naturalized, and the trees neem, Azadirachta indica and quenepa, Melicoccus bijugatus, both of which have highly invasive potential. These species have already made impacts on forested areas and are commonly planted in tourism areas instead of native species. Most importantly, developed properties typically remove understory plants to create open spaces, which promotes the expansion of introduced and invasive grasses.
The invasive potential of A. indica in tropical forests is now widely accepted, and it appears to be modulated by its allelopathic properties against a wide variety of plants [57], and by its easy germination after dispersal via animal consumption of its fruits [21].
The diverse and abundant introduced herb species, especially those of Poaceae, is concerning because of the promotion of seasonal fires [21], although they are not necessarily an impediment to the establishment of trees in tropical dry forests when these factors are controlled [58]. Invasive grasses may selectively favor the establishment of certain tree taxa such as Fabaceae, however, particularly through seasonal fires [59]. Some species of Poaceae should be considered as a threat to plant conservation in LANR and their control should be included in forest restoration and maintenance schemes.
Although plant species introductions are prohibited in LANR [26], our findings strongly suggest that there is still deficient awareness among users, management professionals, and regulatory staff of the threats posed by introduced species. Our results also provide vital information for developing strategies to protect the biodiversity and environmental benefits of LANR. The impacts of invasive introduced plants in tropical dry forests and their control are a rapidly evolving subject [21], to which the experience of LANR can contribute significantly.

4.3. Useful and Overexploited Plants in LANR

Although the extraction of biological products from LANR flora and fauna is prohibited for most purposes without the emission of permits or concessions [18,26], considerable household forest use, fuelwood extraction and some homeworker enterprises producing brooms and baskets are most evident [33,34].
Our results demonstrate that LANR is a reservoir of numerous species of food and medicinal plants. The traditional use of medicinal plants in the vicinity of LANR is widely known but poorly documented. Vestiges of the roles of forests in the traditional, pre-colonial lifestyles and livelihoods in the region surrounding LANR are evident in the continued presence of traditional healers in the Diriomo municipality just outside the Laguna de Apoyo crater [60].
Although ethno-botanical studies in the Caribbean region, where indigenous and Afro-descendant cultures continue to be prominent, have been numerous and extensive in recent decades [61], relatively scant research has been made in the largely latinized Pacific and Central regions of the country, beyond the notable recompilation of Dávila Bolaños [62] and an archaeobotanical investigation [36], both of which demonstrate a robust societal legacy of plant use in pre-Colombian Pacific Nicaragua.
Protected areas can serve as reservoirs for overexploited plant species through managed use, in circumstances where control over activities occurs [22,63]. Our results suggest some species whose presence is limited or absent which merit consideration as examples of overexploitation.
The greater abundance of the introduced A. indica, compared to the valuable Meliaceae, S. humilis and C. odorata, both of which are common in forests where exploitation is controlled, is a concerning feature of our results. Other highly valuable fine woods, such as Dalbergia retusa and Hymenaea courbaril, are present but not at high densities. Some efforts have been made to repopulate the crater interior with these species by enrichment plantings [64].
The low abundance and number of species of palms (Arecaceae) also suggests possible effects of overexploitation. Although the names of two communities on the edge of the Laguna de Apoyo crater, Pacaya and Pacayita, reflect a legacy of the pacaya palm, Chamaedora tepejilote Liebm., in the general area, we did not note its presence within the LANR crater interior. Its flowers are consumed as food. In addition, for Acrocomia aculeata, which produces fruits culturally associated within the region, a few individuals were encountered. Although thatch roofs, historically built with leaves of several species of Arecaceae, are no longer a hallmark of locally built structures since nearly a century ago in this part of Nicaragua, we wonder if overexploitation of palms, as the growth of towns around LANR in the past couple of centuries, has undoubtedly influenced this plant resource in the area. It is not clear to us whether the generally steep, volcanic slopes of the LANR crater interior are poor habitats for native palms or whether their low numbers are the result of overexploitation.
The low population densities of all Orchidaceae merit consideration. These populations may have suffered from over-harvesting for ornamental purposes. Even local trade in ornamental orchids, when not controlled, can threaten populations [65].

4.4. Plant Biodiversity Conservation

Increased attention to LANR in the past few decades, starting with the development and implementation of the management plan, may have attenuated deforestation and forest degradation in LANR [26,38]. The results of this study provide additional information on the value of this reservoir of diverse and ecologically important plant species, which merit more elaborate conservation strategies in a new generation of protected area management. The current management plan has been in effect for 15 years.
Several obstacles confront policy managers and conservation professionals, reducing institutional capacity to execute and evaluate the effectiveness of plant-based conservation policies in the neotropics. Some of the principal limiting factors are: deficient local technical capacity to identify plant taxa; inadequate management and monitoring systems; limited involvement of local communities in monitoring and conservation activities and insufficient permitting and promotion for sustainable plant use [66]. This botanical inventory provides essential baseline information for the development of improved conservation and habitat restoration efforts, habitat monitoring, identifying and recognizing the threats of invasive species, management of species under conservation threats and incorporating stakeholders into sustainable use strategies [18]. Unlike other botanical inventories for protected areas in tropical dry forests, our results provide considerable insight into the role of different plant life forms, beyond trees, in the biodiversity significance of this protected area.

5. Conclusions

We present the most extensive plant species inventory for any protected area in Nicaragua’s tropical dry forest, including native (403) and introduced species (72). Forty-one species of conservation importance were highlighted, and 280 species were identified for various uses. Our results establish the importance of the crater interior of LANR as a reservoir for plant species with ecological, cultural and conservation significance.
Additionally, the results of this study provide insights into the encroachment of introduced plant species within the protected area. The impacts of plant species introductions in tropical dry forests in this region, and their successful management, are insufficiently understood [21]. Likewise, management of this protected area could incorporate recognition and promotion of sustainable plant uses, which offer opportunities for active participation of local populations in activities that may already be occurring without formal recognition.
LANR can serve as a model system for plant conservation in Nicaragua’s tropical dry forest, and especially for the volcanic crater interiors. We recommend that the management of this protected area consider native plant diversity, including life forms other than trees, as well as the threats posed by introduced species, and the incorporation of plant uses in future management strategies for LANR.
We consider that LANR offers opportunities for plant conservation that can be applied to other volcanic crater lake habitats in the region. The fragility of these sites has been recognized in policy [20]. Well-managed plant life in volcanic craters with crater lakes can serve the added function of protecting the lacustrine environments important for providing environmental services and often inhabited by threatened species [9,27].

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/conservation5040083/s1; Table S1: Size, location and habitat type of the 22 survey plots in Laguna de Apoyo Natural Reserve; Table S2: Native and Introduced Plants in the Crater Interior of LANR; Table S3: Cultivated Plants in the Crater Interior of LANR.

Author Contributions

J.K.M.: conceptualization, methodology, formal analysis, investigation, resources, data curation, writing—original draft preparation, review and editing, visualization, supervision, project administration, funding acquisition. A.K.M.: validation, formal analysis, writing—review, and editing. R.M.R.: investigation, validation, formal analysis, writing—review, and editing. L.M.C.-I.: formal analysis, writing—review and editing; visualization. All authors have read and agreed to the published version of the manuscript.

Funding

Our field work was funded by grants from Organismo Autónomo de Parques Nacionales of Spain, United States Agency for International Development, the Small Grants Fund of the Consulate of Canada in Nicaragua, and Conservation, Food, and Health Foundation.

Institutional Review Board Statement

This study was conducted in accordance with the Declaration of Helsinki. Protocols for plant analysis and interviews followed indications established by the Ministry of the Environment and Natural Resources (MARENA) in the Terms of Reference for the Development of the Management Plan for Laguna de Apoyo Nature Reserve, emitted June 2005, and in the Research Permit 008-062010, emitted 24 June 2010, which did not require ethical review or approval. The study adhered to ethical research principles by ensuring low-risk involvement of interviewees, voluntary participation, informed consent, and confidentiality of all data collected.

Informed Consent Statement

Verbal informed consent was obtained from all subjects involved in the study. Verbal consent was used in accordance with permits and cultural norms, thereby reducing potential intimidation associated with signing documents which could compromise confidentiality.

Data Availability Statement

Additional data used in these analyses are available in the Supplementary Materials Tables S1 and S2.

Acknowledgments

Assistance in field surveys was provided by numerous interns, volunteers, and staff of Estación Biológica Laguna de Apoyo, as well as by the cooperation and participation of multiple landowners who facilitated access to their properties. We thank the Nicaraguan Foundation for Integral Community Development and the University of Central America (now Universidad Nacional Camilo Sotelo), both of which provided institutional support, as well as the visionary insights and support of A. L. Hammett, Miguel d’Escoto Brockmann, M. M., and Juan Roberto Zarruk Abdallah. We thank the Ministry of the Environment and Natural Resources for research permits. The authors would like to thank Celene Espadas Manrique for providing us with the map in Figure 1. We thank the anonymous reviewers for their comments which have substantially improved this manuscript.

Conflicts of Interest

The authors declare no conflict of interest.

Abbreviations

The following abbreviations are used in this manuscript:
LANRLaguna de Apoyo Nature Reserve
maslmeters above sea level
cmcentimeters
kmkilometers
SINAPNational Protected Area System of Nicaragua
IUCNInternational Union for Conservation of Nature
CITESConvention on International Trade in Endangered Species of Wild Fauna and Flora

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Figure 1. Panoramic view and location of Laguna de Apoyo Nature Reserve in Nicaragua. Credits: photo Adrianus Konings; map Celene Espadas Manrique.
Figure 1. Panoramic view and location of Laguna de Apoyo Nature Reserve in Nicaragua. Credits: photo Adrianus Konings; map Celene Espadas Manrique.
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Figure 2. Images of LANR flora. From top left to right bottom: Acalypha pseudalopecuroides, Chrysothemis pulchella, Dichorisandra hexandra, Metastelma schlechtendalii, Xanthosoma mexicanum, Disocorea floribunda, Adiantum deflectens, Ipomoea setosa, Guarea glabra, Gliricidia sepium, Hymenaea courbaril and Swietenia humilis.
Figure 2. Images of LANR flora. From top left to right bottom: Acalypha pseudalopecuroides, Chrysothemis pulchella, Dichorisandra hexandra, Metastelma schlechtendalii, Xanthosoma mexicanum, Disocorea floribunda, Adiantum deflectens, Ipomoea setosa, Guarea glabra, Gliricidia sepium, Hymenaea courbaril and Swietenia humilis.
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Figure 3. Number of species in the nine most speciose families in the crater interior of LANR. Bar legend: dark blue: native species at population density less than 20 individuals per km2; light blue: native species at densities greater than 20 individuals per km2; dark green: introduced species at less than 20 individuals per km2; light green: introduced species at densities greater than 20 individuals per km2.
Figure 3. Number of species in the nine most speciose families in the crater interior of LANR. Bar legend: dark blue: native species at population density less than 20 individuals per km2; light blue: native species at densities greater than 20 individuals per km2; dark green: introduced species at less than 20 individuals per km2; light green: introduced species at densities greater than 20 individuals per km2.
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Figure 4. Number of species of native (a) and introduced (b) Angiosperm plant species classified by growth form and type of use. Notice differences in vertical scale between the two graphs.
Figure 4. Number of species of native (a) and introduced (b) Angiosperm plant species classified by growth form and type of use. Notice differences in vertical scale between the two graphs.
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Table 1. Conservation priority plant species in Laguna de Apoyo Nature Reserve, based on the Nicaraguan vegetation database, managed by the Missouri Botanical Garden [41]. NT-Near Threatened; VU-Vulnerable; EN-Endangered. Note that some species were classified as mixed categories.
Table 1. Conservation priority plant species in Laguna de Apoyo Nature Reserve, based on the Nicaraguan vegetation database, managed by the Missouri Botanical Garden [41]. NT-Near Threatened; VU-Vulnerable; EN-Endangered. Note that some species were classified as mixed categories.
FamilySpeciesConservation Status
AcanthaceaeJusticia rothschuhii (Lindau) DurkeeNT
AcanthaceaePseuderanthemum alatum (Nees) Radlk. ex LindauVU/NT
AgavaceaeAgave angustifolia Haw.NT
AmaryllidaceaeZephyranthes lindleyana Herb.NT
AnnonaceaeSapranthus palanga R.E. Fr.NT
ApocynaceaeLacmellea panamensis (Woodson) Markgr.NT
ApocynaceaeMetastelma schlechtendalii Decne.NT
AraceaeXanthosoma mexicanum Liebm.NT
ArecaceaeAcrocomia aculeata (Jacq.) Lodd. ex Mart.NT
ArecaceaeAttalea butyracea (Mutis ex L. f.) Wess. BoerNT
AsparagaceaeYucca guatemalensis BakerNT
AsteraceaeVernonia patens KunthVU/NT
CactaceaeHylocereus costaricensis (F.A.C. Weber) Britton & RoseVU
CommelinaceaeDichorisandra hexandra (Aubl.) C.B. ClarkeNT
ConvolvulaceaeIpomoea setosa Ker Gawl.NT
DioscoreaceaeDioscorea floribunda M. Martens & GaleottiNT
EuphorbiaceaeAcalypha pseudalopecuroides Pax & K. Hoffm.NT
EuphorbiaceaeEuphorbia cotinifolia L.NT
EuphorbiaceaeEuphorbia leucocephala LotsyEN/VU
EuphorbiaceaeEuphorbia prostrata AitonNT
FabaceaeMimosa arenosa (Willd.) Poir.NT
FabaceaeMucuna sloanei Fawc. & RendleNT
FabaceaeVachellia hindsii (Benth.) Seigler & EbingerNT
GesneriaceaeChrysothemis pulchella (Donn ex Sims) DecneNT
LecythidaceaeCouroupita nicaraguarensis DC.NT
MeliaceaeGuarea glabra VahlNT
MyrtaceaePsidium friedrichsthalianum (O.Berg.) Nied.NT
NyctaginaceaeBoerhavia diffusa L.NT
OrchidaceaeCatasetum integerrimum Hook.NT
OrchidaceaeCatasetum maculatum Kunth NT
OrchidaceaeEncyclia cordigera (Kunth) DresslerNT
OrchidaceaeTrichocentrum cebolleta (Jacq.) M.W. Chase & N.H. WilliamsNT
PoaceaeAcroceras zizanioides (Kunth) DandyNT
PoaceaeGynerium sagittatum (Aubl.) P. Beauv.NT
PolygonaceaePodopterus mexicanus Bonpl.NT
PteridaceaeAdiantum deflectens Mart.NT
RutaceaeCasimiroa dura A. Pool & CoronadoNT
SapindaceaeCardiospermum halicacabum L.NT
SapindaceaeCupania dentata DC.NT
SapotaceaePouteria sapota (Jacq.) H. E. Moore & StearnNT
SapotaceaeSideroxylon capiri ssp. capiri (Pittier) T.D. Penn.NT
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McCrary, J.K.; Meyrat, A.K.; Rueda, R.M.; Calvo-Irabien, L.M. Plant Diversity in a Volcanic Crater Interior: Laguna De Apoyo Nature Reserve, Nicaragua. Conservation 2025, 5, 83. https://doi.org/10.3390/conservation5040083

AMA Style

McCrary JK, Meyrat AK, Rueda RM, Calvo-Irabien LM. Plant Diversity in a Volcanic Crater Interior: Laguna De Apoyo Nature Reserve, Nicaragua. Conservation. 2025; 5(4):83. https://doi.org/10.3390/conservation5040083

Chicago/Turabian Style

McCrary, Jeffrey K., Alain Kheim Meyrat, Ricardo M. Rueda, and Luz Maria Calvo-Irabien. 2025. "Plant Diversity in a Volcanic Crater Interior: Laguna De Apoyo Nature Reserve, Nicaragua" Conservation 5, no. 4: 83. https://doi.org/10.3390/conservation5040083

APA Style

McCrary, J. K., Meyrat, A. K., Rueda, R. M., & Calvo-Irabien, L. M. (2025). Plant Diversity in a Volcanic Crater Interior: Laguna De Apoyo Nature Reserve, Nicaragua. Conservation, 5(4), 83. https://doi.org/10.3390/conservation5040083

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