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Article

Diversity and Distribution of Monocot Understory Herbs during Tropical Forest Succession in Northeastern Costa Rica

by
Jennifer W. C. Sun
1,
Robin L. Chazdon
2,3 and
Philip W. Rundel
1,*
1
Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095, USA
2
Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT 06269, USA
3
Forest Research Institute, University of the Sunshine Coast, 90 Sippy Downs Road, Sippy Downs, QLD 4556, Australia
*
Author to whom correspondence should be addressed.
Diversity 2024, 16(8), 439; https://doi.org/10.3390/d16080439
Submission received: 21 May 2024 / Revised: 3 July 2024 / Accepted: 18 July 2024 / Published: 24 July 2024
(This article belongs to the Special Issue Plant Diversity Hotspots in the 2020s)
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Abstract

:
Broad-leaved monocot herbs form one of the most common and diverse growth forms of Neotropical plants. Their significance and frequency of occurrence is particularly notable in the understories of tropical rainforests, where they form a dominant element. We assessed and quantified changes in the cover and diversity of understory herb communities in a chronosequence of 1 ha permanent plots established as part of a multidisciplinary study on tropical forest regeneration in the Atlantic lowlands of northeastern Costa Rica. Sampled were two young stands cleared 12 years ago, two secondary forests with 21 and 39 of years of recovery since clearance, and two stands in old-growth primary forest. Changes in species composition during succession were assessed using Chao’s Jaccard similarity index. Observed species richness ranged from 15 to 26 species in individual plots, with the greatest number of species in the 21-year intermediate-age and fewest in the young 12-year plots. Herb species sampled represented 6 families, 15 genera, and 39 species, with the Araceae contributing the largest number of species. Ten species were sampled in all six stands, while fourteen species were found exclusively in one plot. Herb density (ramets m−2) showed a hump-shade trend, with peak density in the intermediate stands and a lower level in mature and young secondary forests. Mean herb cover in 25 m2 quadrats ranged from 2.0% (young stand) to 22.7% (intermediate-age stand) and differed significantly both among stand types and among sites. Both observed and estimated species richness increased along the chronosequence as a whole, with the highest number of species in primary forest, although only slightly higher than in intermediate-age stands. Over half of the species exhibited some degree of clonal growth, with the extent of clonal spread varying among species and forest stands. Although we did not find a clear pattern between clonality and forest age, we observed a greater number of clonal patches in secondary over primary forest stands.

1. Introduction

Secondary forests are an important feature of tropical landscapes, covering more area than mature forests [1]. In the Neotropics, these forests are increasing in area and value, acting as habitat refugia and biodiversity reservoirs [1,2,3,4], commercial sources of timber and non-timber forest product [5,6,7], and sinks for anthropogenic carbon emissions [8,9]. Tropical secondary forests have until recently been poorly studied, and unifying principles for predicting expected changes along a chronosequence of stand age are lacking. Understanding regeneration patterns within secondary forests has therefore become an important area of research that aligns itself with the need to improve our critical need for sound strategies for tropical forest management and conservation. Although successional dynamics have been well documented in woody tree communities in secondary tropical forests [10,11,12], little is known regarding the regeneration patterns of other functional groups, and we remain far from an integrative view of regeneration [13].
Broad-leaved monocotyledons, the dominant group of understory monocot herbs, represent one of the most abundant and diverse plant forms in tropical forests [14,15,16]. They are distributed across a variety of habitats ranging from dense forest understories to full-sun pastures [17] and display a wide array of ecological traits, including variations in leaf physiology, morphology, architecture, life history, and vegetative reproduction [16,18,19]. Understory herbs are also significant food and habitat sources for diverse communities of frugivores [20] and herbivores [21,22]. Commercially, they are valuable non-timber forest products used in medicine, ornamental horticulture [23,24,25,26,27], and handicraft production in local communities throughout Central and South America [7,28,29].
Despite their diversity, frequency of occurrence, and importance in Neotropical forests, few studies have examined the composition and structure of understory herb communities and enumerated species list for plots of known size [15,30,31], sampled the composition of herb communities in gaps [32,33], and examined herb recovery after anthropogenic disturbances [34,35]. Studies of changes to herb community composition and cover during tropical forest succession are notably lacking.
Here, we assemble a unique dataset on understory herb communities in six 1 ha plots along a chronosequence in northeastern Costa Rica. Specifically, we compared density, percentage coverage, species composition, and species richness between secondary and mature forests spanning three stages of succession. Previous studies along this chronosequence have demonstrated that tree communities in secondary forests are able to recover aboveground biomass, species richness, and species composition at rapid rates [1,12,36,37]. These studies provide evidence for the resilient nature of secondary forests in the study area, though they do not address succession dynamics in herbaceous plants. To our knowledge, this is the first study reporting the behavior of understory herbs during tropical forest succession utilizing this level of taxonomic precision, relatively large plots, and representation of secondary and mature forests.

2. Methods

2.1. Study Area

Research was conducted in January to February 2006 in the Atlantic lowlands of the Sarapiquí region, Heredia Province, northeastern Costa Rica. The natural vegetation is classified as tropical wet forest [38]. Most of the study sites were within La Selva Biological Station, a 1600-hectare reserve of premontane wet forest in the Atlantic lowlands of Costa Rica (10°280 N, 83°590 W). The forest crown varies from 30 to 55 m in height with a closed canopy. The research station has a mean annual rainfall of 4244 mm (1958–2004), with a mean monthly rainfall above 300 mm from May through December. There are peaks of precipitation above 400 mm per month in June–August and November–December and a drier period from January to April. Even in the driest period of February and March, however, rainfall averages are above 150 mm each month. Air temperature is very stable annually, and the daily variation ranges from a maximum monthly mean of 31.7 ± −0.1 °C to a minimum of 20.0 ± −0.2 °C [39] Soils are derived from weathered volcanic deposits and are primarily classified as Ultisols [40]. In the past fifty years, this region has been converted from nearly continuous forest cover to a mosaic of small- and large-scale agriculture fields, cattle pastures, second-growth forests, selectively logged forests, and mature forest fragments [41,42,43].

2.2. Data Collection

In 2005, six 1 ha permanent vegetation monitoring plots were established as part of a multidisciplinary study on tropical forest regeneration in northeastern Costa Rica. Among these plots, two were in young secondary forests (EB and JE), two were in “intermediate-age” secondary forests (LSUR and LEPS), and two were in mature forests (LEPP and SV). Both intermediate secondary and one mature forest stand (LEPP) were located within the La Selva Biological Station, approximately 2 km from each other. The remaining three stands were located approximately 5 km northwest of La Selva in the town of Chilamate: two young secondary stands on privately owned farms, and one mature stand (SV) within a moderately sized tract of mature forest owned by the Selva Verde Ecolodge (Figure 1). All secondary forest stands were cleared for pasture in the early to mid-1970s, actively managed for 4–6 years, and abandoned in the late 1970s to early 1980s [42]. Forest age and land-use history were determined using aerial photographs, historic records [44], satellite images, and interviews with local residents (Table 1).
Within each stand, we recorded the abundance of all understory herbs in contiguous 5 × 5 m quadrats along a series of three parallel transects of 5 × 200 m (40 quadrats per transect, total of 3000 m2 per stand). This design samples more species than square plots of equitable area without oversampling clusters of clonally reproducing plants [45]. We defined understory herbs as broad-leaved monocotyledons in the families Araceae, Costaceae, Cyclanthaceae, Heliconiaceae, Marantaceae, and Zingiberaceae. Understory palms (Arecaceae) were not included, as their functional group has been shown to be more appropriately considered as shrubs [19]. With the exception of Cyclanthaceae and addition of Musaceae, these same families form the majority of understory herbs in paleotropical forests as well. Ramets of clonal species were counted individually if no direct connections were observed. To account for size differences among individuals, two experienced field assistants estimated the percentage coverage of herb species within each 5 × 5 m quadrat. The average of the two coverage estimates was used for data analysis. All specimens were identified to species level in the field.

2.3. Statistical Analysis

To assess overall inventory completeness and to compare species richness at stand level, we constructed sample-based rarefaction curves from 100 randomizations of sample order. Patchiness was set to zero so that each plant was sorted at random to a sample within species, and Mao’s tau approximation was used to calculate expected species richness. For stand-level comparisons, we pooled quadrat and transect data at each site with stand as the replication unit. Species richness for each stand age was then compared for the lowest number of individuals found in any site to correct for differences in species density driven by differences in the density of individuals [46,47,48,49]. Additionally, we computed Chao2, a non-parametric incidence-based richness estimator that reduces the bias that undersampling imposes for expected species richness from 100 resamples for all stands [50]. All calculations were made using the program EstimateS [51,52].
Changes in species composition during succession were assessed using Chao’s Jaccard similarity index, an estimator that is more robust to differences in sample size than other similarity metrics [50] (Chao et al. 2005). Pairwise similarities between each pair of sites were calculated using EstimateS 8.0 [51]. To visualize the relationships, we performed a non-metric multidimensional scaling analysis (NMS), an ordination technique based on ranked distances between n entities on k axes that seeks to minimize distortions caused by reductions in dimensionality [53,54]. Ordinations were based on species that occurred in at least two stands, as rare species have the potential to obscure general patterns in any ordination methods [54]. We examined solutions with three axes, but two axes adequately decreased stress in both ordinations. We used Spearman’s rank correlation test to see if stand position was correlated with either dimension in ordination space. Ordinations and tests were conducted in PAST 2.04 [55].
We compared mean density and percentage coverage among stand types and stands using one-way analysis of variance with post hoc comparisons. Data were log-transformed when necessary to meet the assumptions for analysis of variance. We pooled quadrats and transects by stands for analysis and used stand as the replication unit. All statistical tests were conducted in PAST 2.04 [55].

3. Results

3.1. Distribution of Diversity

The species composition for each successional stage is illustrated in Table 2. We recorded 15,304 individuals from 6 families, 15 genera, and 39 species. The Araceae was the most species-rich family with 10 species (25.6% of the total number of species), while the two species of Zingiberaceae present only comprised 5.1% of the species found. Of the total species recorded, seven (18.0%) were recorded only in mature forest, six (15.4%) only in intermediate succession, and three (7%) only in early succession forest stands. Fourteen species were found exclusively in one site and ten in all six sites.
The five most abundant species were Goeppertia micans, Heliconia irrasa subsp. undulata, Spathophilum fulvovirens, G. cleistantha, and Philodendron grandipes. These represent three of the six families and account for 84.3% of all individuals sampled. With the exception of G. cleistantha, these species exhibited strong clonal capabilities and appeared in numerous patches throughout all six sites (Table 2). Conversely, 26 species comprised less than 5% of the total number of individuals sampled. The occurrence of rare species was highest in the mature forest stand located in La Selva Biological Station (LEPP) and lowest in the two early succession stands (EB and JE). Overall, 23 species exhibited some degree of clonal growth, and 23 species occurred in more than one stand type (Table 2).
Although the sample-based rarefaction curve for all stands failed to saturate (Figure 2), this is not unexpected for very species-rich tropical floras, and the difference was small when comparing values for estimated and observed number of species. In the 29-year stand (LEPS), for example, the recorded species richness was 96.0% of what Chao2 estimated true richness to be. For five stands, we were within at least 90.4% of the estimator, which suggests that a substantial portion of the total herb diversity was recorded at each site. In one of the young secondary stands (JE), sampling was within 80.2% of the estimator and the rarefaction curve failed to reach an asymptote. In order to make ecologically and statistically meaningful comparisons, we excluded JE from richness analysis between stands. Species richness was therefore compared using the second-lowest common number of individuals recorded. Observed species richness ranged from 15 to 26 species, with the greatest number of species in the intermediate-age LSUR and the fewest in EB and JE. When we compared stands for a common rarefied number of individuals to correct for differences in species density driven by differences in the density of individuals, both observed and estimated species richness increased along the chronosequence as a whole, with the highest number of species in the mature stand LEPP (Figure 3). Estimated species richness was second-highest in the mature-stand SV, but this number did not differ greatly from those recorded for intermediate secondary forests.

3.2. Density and Coverage

Herb density (ramets m−2) showed a hump-shade trend, with peak density in the intermediate stands and a rapid decline in mature and young secondary forests (Figure 4a). Mean density, ranging from 0.06 to 1.52 ramets m−2, differed significantly both among stand types (one-way ANOVA, F2,717 = 34.53, p < 0.0001) and among sites (one-way ANOVA, F5,714 = 20.04, p < 0.0001). With the exception of the young secondary forests, stands similar in age were not significantly different (Tukey’s post hoc comparisons, p < 0.0001). The stand-type effects reflect higher mean herb density in intermediate succession stands compared to mature and early succession stands, i.e., herbs were roughly twice as abundant in the intermediate stands as in other stands.
Percentage herb cover, which accounts for the variance in size and degree of clonality, also showed a hump-shaped trend (Figure 4b). Mean cover in 25 m2 quadrats ranged from 2.0 to 22.7% and differed significantly both among stand types (one-way ANOVA, F2,717 = 83.72, p < 0.0001) and among sites (one-way ANOVA, F5,714 = 38.51, p < 0.0001). Unlike density patterns, percentage coverage differed significantly among mature forests, but not among young secondary forests (Tukey’s post hoc comparisons, p < 0.001 and p = 0.639, respectively).

3.3. Floristic Similarity

Species composition showed high similarity among stands close in age. Floristic similarity is calculated as the number of species shared between stands divided by the number in either stand individually. Intermediate stands were most similar (0.993), followed by mature forest stands (0.954). The similarity index was lowest between the intermediate and young secondary stands. SV had relatively high similarity values to LSUR and LEPS. Stands separated along the first axis of the NMS plot (2-d stress = 0.127), with mature and intermediate stands grouping on the left and young stands on the right (Figure 5). Stand position along the first dimension of the ordination was correlated with stand age (Spearman’s rank correlation rs = 0.708, p < 0.001).

4. Discussion

Although the chronosequence approach enables researchers to rapidly assess forest changes across a large time frame, these studies infer rates of change from a single-time investigation and assume that forest stands differ only in age. In a comparison between chronosequence and long-term vegetation dynamics studies [56], certain community-level patterns in woody seedlings were predictable from chronosequence trends, while others were not. More likely, the trajectories of plant communities during succession are shaped by a complex set of interactions among landscape characteristics, local site factors, biotic interactions, and species life histories [57]. In this study, we attempted to minimize the effect of environmental and historical heterogeneity in our interpretation by selecting sites based on verifiable estimates of stand age. Sites were also selected to encompass the same range in land-use history, soil types, and topography. Although this does not replace the need for continued long-term sampling in succession studies, it does offer important insights into the recruitment and recovery process of understory herb communities in tropical forests.
Unlike woody perennial species, understory herbs lack the stiff petioles and support structures needed to attain canopy heights and must therefore employ a different occupation strategy during succession. Over half of the species recorded in this study exhibited some degree of clonal growth, with the extent of clonal spread varying among species and forest stands. Although we did not find a clear pattern between clonality and forest age, we did observe a greater number of clonal patches in secondary forest compared to mature forest stands. Previous studies on the terrestrial herb Aechmea magdalenae (Bromeliacae) have indicated that clonal reproduction is positively correlated with canopy openness [58,59]. Thus, increased clonal activity at high irradiance levels, such as those experienced in the early phases of succession, may be a strategy for rapid lateral growth in pioneer herbaceous species. We also observed that species with strong clonal capabilities tended to be generalists, at least with respect to successional status. These generalists are by far the most abundant species found in our study, which implies a high degree of resilience in the study area.
Changes in herb density reflect interactions between intrinsic life-history traits and density-dependent processes during succession. In our stands, density rapidly increased in the early phases of succession as pioneer species were episodically recruited, forming a strong cohort dominated by fast-growing and light-demanding species (e.g., Heliconia latispatha and Costus laevis). High photosynthetic capacities coupled with clonal capabilities enable many of these pioneer species to proliferate in higher light environments such as open pastures, canopy gaps and forests edges [16,18,19,60]. As the canopy closes and light levels decrease, herb density and coverage eventually decline when pioneer species die off, and density-dependent forces, such as competitive exclusion, regulate the recruitment and survival processes of remaining and new, more shade-tolerant herb species [61,62,63].
The nature and extent of site-specific disturbances can also lead to unpredictable changes in density patterns in the early phases of succession. Mean density of herbs was 16.6-fold higher in EB than JE, despite the plots being in close proximity to each other and sharing the same soils and topography. JE also had the lowest observed species richness among all stands. A likely explanation for our results is the different composition of remnant vegetation in these plots: EB had 22 remnant trees from 15 species that composed a substantial fraction of the plot basal area, whereas JE had only a single remnant palm tree that did not contribute appreciably to plot basal area. Remnant trees have long-lasting positive effects on forest recovery by attracting dispersal agents that deposit or regurgitate seeds [64,65,66,67], and by eliminating competition from thicket-forming colonizers such as ferns and grasses [30,68,69,70,71]. While we did not measure the density of dispersal agents, we did observe thick and pure carpets of fern thickets in JE, which likely inhibited the recruitment of herb individuals and contributed to overall low density and richness of understory herbs at this plot. Similar findings have been found in other systems characterized by weedy grow [64,65,66].
Interestingly, we did not detect a significant difference in percentage coverage between the young secondary forest stands. The observed pattern is largely due to the dominance of the highly clonal Goeppertia micans in the EB plot, which comprised 93.5% of total ramets recorded. The increase in density, however, did not equate to a proportional increase in percentage coverage, as G. micans does not occupy nearly as much space as the other species in this study.
Despite fluctuations in overall density, species richness increased along the chronosequence, whether measured as the Mao’s tau rarefaction or the incidence-based coverage estimator Chao2. The pattern of higher herb density and correspondingly higher species richness revealed density effects in our stands, which we controlled by comparing rarefaction approximation and estimator values at the lowest common number of individuals found across all six stands. With the exception of Anthurium lancifolium Schott and Dracontium gigas (Seem.) Engl., fast-growing pioneer species were also found in older forests, indicating that pioneer species can persist locally long after canopy closure. Demographic studies on Goeppertia inocephala (Kuntze) H. Kenn. and Nicolson have shown that individuals recruited in canopy gaps are able to survive as relict species in mature forests by reducing leaf and inflorescence production [72,73,74]. Surprisingly, we also found that slow-growing mature forest species such as Spathiphyllum fulvovirens Schott were able to colonize early in succession, which has only been shown for tree species [75,76], and underscore the importance of initial floristic composition in the determination of successional pathways. In contrast, less than a quarter of the mature forest species were recruited after canopy development.
Succession changes in species richness and herb density may also be affected by human activities. Among the mature forest stands, species richness and mean density was higher in the stand located within La Selva Biological Station (LEPP) compared to the stand outside the station (SV). Whereas 13% of the total species recorded were found exclusively in LEPP, no species were exclusive to SV. In the past 20–30 years, La Selva Biological Station has been actively protected from hunting by game wardens, whereas forest stands in the surrounding landscape have been exposed to continuous hunting pressure [75]. In response to protection from hunting, populations of terrestrial mammals, such as the agouti (Agouti paca) and collared peccary (Tayassu tajacu), are noticeably higher at La Selva than in surrounding regions [12,76,77]. Previous studies [12] found that the density of canopy tree and palm seedlings and saplings was 4.7- and 2.0-fold higher, respectively, in SV than in LEPP. The most abundant species in both stands, the palm Welfia regia, had a seedling density of 416 seedlings m−2 in SV and 48 seedlings m−2 in LEPP. Because palms are particularly favored by peccaries [77], the reduced populations of seed predators in non-protected areas may have led to the increased abundance of Welfia seedlings and saplings and the competitive exclusion of understory herbs. [78]. Those researchers also demonstrated that close proximity to understory palms can reduce growth and survival in co-occurring plants, attributing most of this impact to increased herbivory, pathogen attacks, and reduced light availability [78].
Although species composition may not recover as rapidly as species richness in tropical forests [3,11,79,80,81,82], floristic reassembly of understory herbs in northeastern Costa Rica is gradually occurring. Our results are contrary to what has been reported in Singapore, where secondary forests failed to recruit species from adjacent mature forests due to a highly fragmented study area and the extinction of seed-dispersing fauna [80]. In comparison, deforestation in the Sarapiquí area has been relatively modest [43], and other studies have demonstrated a high capacity for compositional recovery in woody tree species [37,42,56,83,84], palms [36], and lianas [85].
It is worth noting that stand recovery does not imply that succession always follows a predictable and directional trajectory. More often, species composition in human-influenced landscapes is influenced by a complex set of interacting factors in addition to forest age that lead to idiosyncratic pathways [86,87]. In our study, some of the variability in successional patterns found between sites of similar age may be a result of these complex interactions. Early succession stands were widely separated along the second dimension of our ordination analysis, and the mature stands showed distinct floristic compositions. Such variability was probably responsible for the large difference in pairwise comparisons between young succession stands and the less significant difference in mature forests. Although additional study plots may have been ideal, it was not possible given the intensive nature of our sampling protocol. Instead, we chose to sample a large area in each plot to offset the small number of plots.
Results from this study provide clear insights into how understory herbs respond to changes following disturbance and their potential role in shaping stand-level dynamics during succession. Species richness approaches but does not reach old-growth levels and species composition offers an optimistic forecast of rapid understory herb reassembly in our study area. The data collected here also forms a valuable baseline for assessing the survival and recovery of other understory herb communities following disturbance in comparable tropical forests. Future studies focusing on the interaction of factors that drive change in herb communities may be useful in understanding and predicting successional dynamics in tropical forests.

Author Contributions

Conceptualization, J.W.C.S. and R.L.C.; methodology, J.W.C.S., R.L.C. and P.W.R.; validation J.W.C.S., R.L.C. and P.W.R.; formal analysis, J.W.C.S. investigation J.W.C.S., resources, J.W.C.S., R.L.C. and P.W.R.; data curation; J.W.C.S., R.L.C. and P.W.R.; writing—original draft preparation, J.W.C.S.; writing—review and editing, J.W.C.S., R.L.C. and P.W.R.; visualization, J.W.C.S.; supervision, J.W.C.S., R.L.C. and P.W.R.; project administration; J.W.C.S. and R.L.C.; funding acquisition, P.W.R. All authors have read and agreed to the published version of the manuscripts.

Funding

This research was funded by UCLA Trust Account.

Institutional Review Board Statement

Research permits were granted by the Comisión Institucional de Biodiversidad (Institutional Biodiversity Committee, University of Costa Rica; resolution VI-8315-2014) and authorized by La Selva Biological Station.

Data Availability Statement

Orinal field datais available from the authors on reqest.

Acknowledgments

Much of this research would not have been possible without the field assistance of Mauricio Gaitan and his knowledge of the La Selva flora. Susan Letcher and Arielle Cooley provided thoughtful reviews of the manuscript. We gratefully acknowledge the staff of La Selva Biological Station for their encouragement and support in carrying out this field program.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Chazdon, R.L. Second Growth: The Promise of Regeneration in an Age of Deforestation; University of Chicago Press: Chicago, IL, USA, 2014. [Google Scholar]
  2. Chazdon, R.L. Tropical forests–log ‘em or leave’ em? Science 1998, 281, 1295–1296. [Google Scholar] [CrossRef]
  3. Chazdon, R.L. Tropical forest recovery: Legacies of human impact and natural disturbances. Perspect. Plant Ecol. Evol. Syst. 2003, 6, 51–71. [Google Scholar] [CrossRef]
  4. Wright, S.J.; Muller-Landau, H.C. The future of tropical forest species. Biotropica 2006, 38, 287–301. [Google Scholar] [CrossRef]
  5. Chazdon, R.L.; Coe, F.G. Ethnobotany of woody species in second-growth, old-growth, and selectively logged forests of northeastern Costa Rica. Conserv. Biol. 1999, 13, 1312–1322. [Google Scholar] [CrossRef]
  6. Silver, W.L.; Kueppers, L.M.; Lugo, A.E.; Ostertag, R.; Matzek, V. Carbon sequestration and plant community dynamics following reforestation of tropical pasture. Ecol. Appl. 2004, 14, 1115–1127. [Google Scholar] [CrossRef]
  7. Ticktin, T. The ecological implications of harvesting non-timber forest products. J. Appl. Ecol. 2004, 41, 11–21. [Google Scholar] [CrossRef]
  8. Grace, J. Understanding and managing the global carbon cycle. J. Ecol. 2004, 92, 189–202. [Google Scholar] [CrossRef]
  9. Feldpausch, T.R.; Riha, S.J.; Fernandes, E.C.M.; Wandell, E.V. Development of forest structure and leaf area in secondary forests regenerating on abandoned pastures in Central Amazonia. Earth Interact. 2005, 9, 1–22. [Google Scholar] [CrossRef]
  10. Hubbell, S.P.; Foster, R.B.; O’Brien, S.T.; Harms, K.E.; Condit, R.C.; Wechsler, B.; Wright, S.J.; Loo de Lao, S. Light-gap disturbances, recruitment limitation, and tree diversity in a neotropical rainforest. Science 1999, 283, 554–557. [Google Scholar] [CrossRef]
  11. Capers, R.S.; Chazdon, R.L.; Brenes, A.R. Successional dynamics of woody seedling communities in wet tropical secondary forests. J. Ecol. 2005, 93, 071–1084. [Google Scholar] [CrossRef]
  12. Chazdon, R.L.; Finegan, B.; Capers, R.S.; Salgado-Negret, B.; Casanoves, F.; Boukili, V.; Norden, N. Composition and dynamics of functional groups of trees during tropical forest succession in northeastern Costa Rica. Biotropica 2010, 42, 31–40. [Google Scholar] [CrossRef]
  13. Dewalt, S.J.; Schnitzer, S.A.; Denslow, J.S. Density and diversity of lianas along a chronosequence in a central Panamanian lowland forest. J. Trop. Ecol. 2000, 16, 1–19. [Google Scholar] [CrossRef]
  14. Galeano, G.; Suárez, S.; Balslev, H. Vascular plant species count in a wet forest in the Chocó area on the pacific coast of Colombia. Biodivers. Conserv. 1998, 7, 1563–1575. [Google Scholar] [CrossRef]
  15. Costa, F.R.C. Structure and composition of the ground-herb community in a terra-firme central Amazonian forest. Acta Amaz. 2004, 34, 53–59. [Google Scholar] [CrossRef]
  16. Rundel, P.W.; Cooley, A.; Gerst, K.L.; Riordan, E.C.; Sharifi, M.R.; Sun, J.W.; Tower, J.A. Functional traits of broad-leaved monocot herbs in the understory and forest edges of a Costa Rican rainforest. PeerJ 2020, 8, e9958. [Google Scholar] [CrossRef]
  17. Benítez-Malvido, J. Effect of low vegetation on the recruitment of plants in successional habitat types. Biotropica 2006, 38, 171–182. [Google Scholar] [CrossRef]
  18. Rundel, P.W.; Sharifi, M.R.; Gibson, A.C.; Esler, K.J. Structural and physiological adaptation to light environments in neotropical Heliconia (Heliconiaceae). J. Trop. Ecol. 1998, 14, 789–801. [Google Scholar] [CrossRef]
  19. Cooley, A.M.; Reich, J.A.; Rundel, P.W. Petiole biomechanics of neotropical understory monocots. Am. J. Bot. 2004, 91, 573–581. [Google Scholar] [CrossRef] [PubMed]
  20. Wunderle, J.M.; Willig, M.R.; Henriques, L.M.P. Avian distribution in treefall gaps and understory of terra firme forest in the lowland Amazon. Ibis 2005, 147, 109–129. [Google Scholar] [CrossRef]
  21. Horvitz, C.C.; Schemske, D.W. Effects of plant size, leaf herbivory, local competition and fruit production no survival, growth and future reproduction of a neotropical herb. J. Ecol. 2002, 90, 279–290. [Google Scholar] [CrossRef]
  22. Kay, K.M.; Schemske, D.W. Pollinator assemblages and visitation rates for 11 species of neotropical Costus (Costaceae). Biotropica 2003, 35, 198–207. [Google Scholar]
  23. Berry, F.; Kress, W.J. Heliconia: An Identification Guide; Smithsonian Institution Press: Washington, DC, USA, 1991. [Google Scholar]
  24. Salick, J.; Mejia, A.; Anderson, T. Non-timber forest products integrated with natural forest management, Río San Juan, Nicaragua. Ecol. Appl. 1995, 5, 878–895. [Google Scholar] [CrossRef]
  25. Castro, C.E.F.; Graziano, T.T. Espécies de Heliconia (Heliconiaceae) no Brasil. Rev. Bras. Hortic. Ornam. 1997, 3, 15–28. [Google Scholar]
  26. Lorenzi, H.; de Souza, H.M. Plantas Ornamentais No Brasil; Instituto Plantarum, Nova Odessa: São Paulo, Brazil, 1999. [Google Scholar]
  27. Arruda, R.; Carvalho, V.T.; Andrade, P.C.M.; Pinto, M.G. Heliconias of the Baixo Jurua Extractive Reserve: Economical potential for Amazonian traditional population. Acta Amazon. 2008, 38, 611–616. [Google Scholar] [CrossRef]
  28. Nakazono, E.M.; Bruna, E.M.; Mesquita, R.C.G. Experimental harvesting of the non-timber forest product Ischnosiphon polyphyllus in central Amazonia. For. Ecol. Manag. 2004, 190, 219–225. [Google Scholar] [CrossRef]
  29. Ticktin, T.; Johns, T.; Chapol Xoca, V. Patterns of growth in Aechmea magdalenae (Bromeliaceae) and its potential as a forest crop and conservation strategy. Agric. Ecosyst. Environ. 2003, 94, 123–139. [Google Scholar] [CrossRef]
  30. Royo, A.A.; Carson, W.P. The herb community of a tropical forest in central Panamá: Dynamics and impact of mammalian herbivores. Oecologia 2005, 145, 66–75. [Google Scholar] [CrossRef]
  31. Smith, A.P. Respuestas de hierbas del sotobosque tropical a claros ocasionados por la caída de árboles. Rev. Biol. Trop. 1987, 35, 111–118. [Google Scholar]
  32. Dirzo, R.; Horvitz, C.C.; Quevedo, H.; López, M.A. The effect of gap size and age on understory herb community of a tropical Mexican rain forest. J. Ecol. 1992, 80, 809–822. [Google Scholar] [CrossRef]
  33. Costa, F.R.C.; Magnusson, W. Selective logging effects on abundance, diversity, and composition of tropical understory herbs. Ecol. Appl. 2002, 12, 807–819. [Google Scholar] [CrossRef]
  34. Ribeiro, M.B.N.; Bruna, E.M.; Mantovani, W. Influence of post-clearing treatment on the recovery of herbaceous plant communities in Amazonian secondary forests. Restor. Ecol. 2010, 18, 50–58. [Google Scholar] [CrossRef]
  35. Letcher, S.G.; Chazdon, R.L. Rapid recovery of biomass, species richness, and species composition in a forest chronosequence in northeastern Costa Rica. Biotropica 2009, 41, 608–617. [Google Scholar] [CrossRef]
  36. Norden, N.; Chazdon, R.L.; Chao, A.; Jiang, Y.-H.; Vilchez-Alvarado, B. Resilience of tropical rain forests: Tree community reassembly in secondary forests. Ecol. Lett. 2009, 12, 385–394. [Google Scholar] [CrossRef] [PubMed]
  37. Chazdon, R.L.; Alvarado, B.V.; Letcher, S.G.; Wendt, A.; Sezen, U.U. Effects of human activities on successional pathways: Case studies from lowland wet forests of northeastern Costa Rica. In The Social Lives of Forests: Past, Present, and Future of Woodland Resurgence Get Access Arrow 2014; University of Chicago Press: Chicago, IL, USA, 2014; pp. 129–142. [Google Scholar]
  38. Holdridge, L.R.; Grenk, W.G.; Hatheway, W.H.; Liang, T.; Tosi, J.A. Forest Environments in Tropical Life Zones; Pergamon Press: New York, NY, USA, 1975. [Google Scholar]
  39. Graham, E.A.; Hansen, M.; Kaiser, W.J.; Lam, Y.; Stealey, M.J.; Yuen, E.; Rundel, P.W. Dynamic microclimate boundaries across a sharp tropical rainforest clearing edge. Remote Sens. 2021, 13, 1646. [Google Scholar] [CrossRef]
  40. Sollins, P.; Sancho, F.; Mata, R.; Sanford, R.L. Soils and soil process research. In La Selva: Ecology and Natural History of a Neotropical Rainforest; McDade, L.A., Bawa, K.S., Hespenheide, H., Hartshorn, G.S., Eds.; University of Chicago Press: Chicago, IL, USA, 1994; pp. 35–53. [Google Scholar]
  41. Butterfield, R.P. The regional context: Land colonization and conservation in Sarapiquí. In La Selva: Ecology and Natural History of a Neotropical Rain Forest; McDade, L.A., Bawa, K.S., Hespenheide, H.A., Hartshorn, G.S., Eds.; University of Chicago Press: Chicago, IL, USA, 1994; pp. 299–306. [Google Scholar]
  42. Guariguata, M.R.; Chazdon, R.L.; Denslow, J.S.; Dupuy, J.M.; Anderson, L. Structure and floristics of secondary and old-growth forest stands in lowland Costa Rica. Plant Ecol. 1997, 132, 107–120. [Google Scholar] [CrossRef]
  43. Read, J.M. Land-Cover Change Detection for the Tropics Using Remote Sensing and Geographic Information Systems. Ph.D. Dissertation, Louisiana State University, Baton Rouge, LA, USA, 1999. [Google Scholar]
  44. Pierce, S.M. Environmental history of La Selva biological Station: Colonization and deforestation of Sarapiquí Canton, Costa Rica. In Changing Tropical Forests: Historical Perspectives on Today’s Challenges in Central and South America; Steen, H.K., Rucker, R.P., Eds.; Forest History Society: Durham, NC, USA, 1992; pp. 40–57. [Google Scholar]
  45. Philips, O.; Miller, J.S. Global Patterns of Plant Diversity: Alwyn H. Gentry’s Forest Transect Data Set; Missouri Botanical Garden Press: St. Louis, MO, USA, 2002. [Google Scholar]
  46. Colwell, R.K.; Coddington, J.A. Estimating terrestrial biodiversity through extrapolation. Philos. Trans. R. Soc. B 1994, 345, 101–118. [Google Scholar]
  47. Chazdon, R.L.; Colwell, R.K.; Denslow, J.S.; Guariguata, M.R. Statistical methods for estimating species richness of woody regeneration in primary and secondary rain forests of NE Costa Rica. In Forest Biodiversity Research, Monitoring and Modeling: Conceptual Background and Old World Case Studies; Dallmeier, F., Comiskey, J.A., Eds.; Parthenon Publishing: Paris, France, 1998; pp. 285–309. [Google Scholar]
  48. Gotelli, N.; Colwell, R.K. Quantifying biodiversity: Procedures and pitfalls in the measurement and comparison of species richness. Ecol. Lett. 2001, 4, 379–391. [Google Scholar] [CrossRef]
  49. Longino, J.T.; Coddington, J.; Colwell, R.K. The ant fauna of a tropical rain forest: Estimating species richness three different ways. Ecology 2002, 83, 689–702. [Google Scholar] [CrossRef]
  50. Chao, A.; Chazdon, R.L.; Colwell, R.K.; Shen, T.-J. A new statistical approach for assessing similarity of species composition with incidence and abundance data. Ecol. Lett. 2005, 8, 148–159. [Google Scholar] [CrossRef]
  51. Colwell, R.K. EstimateS: Statistical Estimation of Species Richness and Shared Species from Samples. Version 8.00. 2007. Available online: http://purl.oclc.org/estimates (accessed on 1 January 2010).
  52. Colwell, R.K.; Elsensohn, J.E. EstimateS turns 20: Statistical estimation of species richness and shared species from samples, with non-parametric extrapolation. Ecography 2014, 37, 609–613. [Google Scholar] [CrossRef]
  53. Schiffman, S.S.; Lance, R.M.; Young, F.W. Introduction to Multidimensional Scaling: Theory, Methods, and Applications; Academic Press: New York, NY, USA, 1981. [Google Scholar]
  54. McCune, B.; Grace, J.B. Analysis of Ecological Communities; MjM Software Design: Gleneden Beach, OR, USA, 2002. [Google Scholar]
  55. Hammer, Ø.; Harper, D.A.T.; Ryan, P.D. PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontol. Electron. 2001, 4, 9. Available online: http://palaeo-electronica.org/2001_1/past/issue1_01.htm (accessed on 1 January 2010).
  56. Chazdon, R.L.; Letcher, S.G.; van Breugel, M.; Martínez-Ramos, M.; Bongers, F.; Finegan, B. Rates of change in tree communities of secondary Neotropical forests following major disturbances. Philos. Trans. R. Soc. Lond. Ser. B 2007, 362, 273–289. [Google Scholar] [CrossRef] [PubMed]
  57. Pickett, S.T.A.; Collins, S.L.; Armesto, J.J. Models, mechanisms and pathways of succession. Bot. Rev. 1987, 53, 335–371. [Google Scholar] [CrossRef]
  58. Pfitsch, W.A.; Smith, A.P. Growth and photosynthesis of Aechmea magdalenae, a terrestrial CAM plant in a tropical moist forest, Panama. J. Trop. Ecol. 1988, 4, 199–207. [Google Scholar] [CrossRef]
  59. Villegas, A.C. Spatial and temporal variability in clonal reproduction of Aechmea magdalenae, a tropical understory herb. Biotropica 2001, 33, 48–59. [Google Scholar] [CrossRef]
  60. Dupuy, J.M.; Chazdon, R.L. Effects of vegetation cover on seedling and sapling dynamics in secondary tropical wet forests in Costa Rica. J. Trop. Ecol. 2006, 22, 65–76. [Google Scholar] [CrossRef]
  61. Harms, K.E.; Powers, J.S.; Montgomery, R.A. Variation in small sapling density, understory cover, and resource availability in four neotropical forests. Biotropica 2004, 36, 40–51. [Google Scholar] [CrossRef]
  62. Uriarte, M.; Canham, C.D.; Thompson, J.; Zimmerman, J.K.; Brokaw, N. Seedling recruitment in a hurricane-drive tropical forest: Light limitation, density-dependence and the spatial distribution of parent trees. J. Ecol. 2005, 93, 291–304. [Google Scholar] [CrossRef]
  63. Wills, C.; Harms, K.E.; Condit, R.; King, D.; Thompson, J.; He, F.; Muller-Landau, H.C.; Ashton, P.; Losos, E.; Comita, L.; et al. Nonrandom processes maintain diversity in tropical forests. Science 2006, 27, 527–531. [Google Scholar] [CrossRef]
  64. Guevara, S.; Purata, S.E.; van der Maarl, E. The role of remnant forest trees in tropical secondary succession. Vegetatio 1986, 66, 77–84. [Google Scholar] [CrossRef]
  65. Slocum, M.G.; Horvitz, C.C. Seed arrival under different genera of trees in a neotropical pasture. Plant Ecol. 2000, 149, 51–62. [Google Scholar] [CrossRef]
  66. Guevara, S.; Laborde, J.; Sánchez-Rios, G. Rain forest regeneration beneath the canopy of fig trees isolated in pastures of Los Tuxtlas, Mexico. Biotropica 2004, 36, 99–108. [Google Scholar] [CrossRef]
  67. Schlawin, J.R.; Zahawi, R.A. ‘Nucleating’ succession in recovering neotropical wet forests: The legacy of remnant trees. J. Veg. Sci. 2008, 19, 485–492. [Google Scholar] [CrossRef]
  68. Aide, T.M.; Cavalier, J. Barriers to tropical lowland forest restoration in the Sierra Nevada de Santa Marta, Colombia. Restor. Ecol. 1994, 2, 219–229. [Google Scholar] [CrossRef]
  69. Walker, L.R. Effects of fern thickets on woodland development on landslides in Puerto Rico. J. Veg. Sci. 1995, 5, 425–532. [Google Scholar] [CrossRef]
  70. Suazo Ortuño, I. Aspectos Ecológicos de Pteridium aquilinum (Polypodiaceae) en la Región de Chajul, Chiapas. Master’s Thesis, Facultad de Biología, Universidad Michoacana de San Nicolás de Hidalgo, Morelia, Mexico, 1998. [Google Scholar]
  71. George, L.O.; Bazzaz, F.A. The fern understory as an ecological filter: Emergence and establishment of canopy tree seedlings. Ecology 1999, 80, 833–845. [Google Scholar] [CrossRef]
  72. Matlaga, D.P.; Horvitz, C. Growth and survival across a gap– understory gradient: Contrast in performance of sexually vs. clonally produced offspring. Am. J. Bot. 2009, 96, 439–447. [Google Scholar] [CrossRef] [PubMed]
  73. Matlaga, D.P.; Horvitz, C.C. Large size and high light do not lower the cost of reproduction for the Neotropical herb Goeppertia marantifolia. Am. J. Bot. 2015, 102, 350–357. [Google Scholar] [CrossRef]
  74. Matlaga, D.P.; Snyder, R.K.; Horvitz, C.C. Dispersal of Goeppertia marantifolia clonal offspring increases with greater canopy openness and larger plant size. J. Trop. Ecol. 2017, 33, 107–113. [Google Scholar] [CrossRef]
  75. van Breugel, M.; Bongers, F.; Martínez-Ramos, M. Species dynamics during early secondary forest succession: Recruitment, mortality and species turnover. Biotropica 2007, 35, 610–619. [Google Scholar] [CrossRef]
  76. Hanson, T.; Brunsfeld, S.; Finegan, B. Variation in seedling density and seed predation indicators for the emergent tree Dipteryx panamensis in continuous and fragmented rain forest. Biotropica 2006, 38, 770–774. [Google Scholar] [CrossRef]
  77. Franklin, J.; Rey, S.J. Spatial patterns of tropical forest trees in Western Polynesia suggest recruitment limitations during secondary succession. J. Trop. Ecol. 2007, 23, 1–12. [Google Scholar] [CrossRef]
  78. Beck, H. Seed predation and dispersal by peccaries throughout the Neotropics and its consequences: A review and synthesis. In Seed Fate; Forget, P.M., Lambert, J.E., Hulme, P.E., Wall, S.B.V., Eds.; CABI Publishing: Cambridge, MA, USA, 2005; pp. 77–100. [Google Scholar]
  79. Wang, Y.H.; Augspurger, C. Dwarf palms and cyclanth strongly reduce Neotropical seedling recruitment. Oikos 2004, 107, 619–633. [Google Scholar] [CrossRef]
  80. Corlett, R.T. The ecological transformation of Singapore, 1819–1990. J. Biogeogr. 1992, 19, 411–420. [Google Scholar] [CrossRef]
  81. Clark, D.B. Abolishing virginity. J. Trop. Ecol. 1996, 12, 735–739. [Google Scholar] [CrossRef]
  82. Finegan, B. Pattern and process in neotropical secondary rain forests: The first 100 years of succession. Trends Ecol. Evol. 1996, 11, 119–124. [Google Scholar] [CrossRef]
  83. Dewalt, S.J.; Maliakal, S.K.; Denslow, J.S. Changes in vegetation structure and composition along a tropical forest chronosesquence: Implications for wildlife. For. Ecol. Manag. 2003, 182, 139–151. [Google Scholar] [CrossRef]
  84. Guariguata, M.R.; Ostertag, R. Neotropical secondary succession: Changes in structural and functional characteristics. For. Ecol. Manag. 2001, 148, 185–206. [Google Scholar] [CrossRef]
  85. Letcher, S.G.; Chazdon, R.L. Lianas and self-supporting plants during tropical forest succession. For. Ecol. Manag. 2009, 257, 2150–2156. [Google Scholar] [CrossRef]
  86. Gleason, H.A. The individualistic concept of the plant association. Bull. Torrey Bot. Club 1926, 53, 7–26. [Google Scholar] [CrossRef]
  87. Chazdon, R.L. Chance and determinism in tropical forest succession. In Tropical Forest Community Ecology; Carson, W., Schnitzer, S.A., Eds.; Wiley-Blackwell Publishing: Oxford, UK, 2008; pp. 384–408. [Google Scholar]
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Figure 1. Map of the study area, showing the locations of the six monitoring plots in relation to major rivers (blue line) and roads (black line) in Sarapiquí Province of northeastern Costa Rica. The arrow indicates location of study area within Costa Rica.
Figure 1. Map of the study area, showing the locations of the six monitoring plots in relation to major rivers (blue line) and roads (black line) in Sarapiquí Province of northeastern Costa Rica. The arrow indicates location of study area within Costa Rica.
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Figure 2. Pooled sample-based rarefaction curve (solid line) with 95% confidence intervals (dashed line) for all understory herb species recorded in six forest stands across a chronosequence in northeastern Costa Rica.
Figure 2. Pooled sample-based rarefaction curve (solid line) with 95% confidence intervals (dashed line) for all understory herb species recorded in six forest stands across a chronosequence in northeastern Costa Rica.
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Figure 3. Species richness by site. For rarefaction, Mao’s tau was used for smoothing approximation of observed species richness, and Chao2 is the incidence-based coverage estimator. Error bars show the standard deviation for Mao’s tau and Chao2. Rarefaction and Chao 2 were calculated for each site using the lowest number of individuals observed across all stands (1260 individuals). The JE plot was not included in these comparisons due to the low number of understory herbs sampled.
Figure 3. Species richness by site. For rarefaction, Mao’s tau was used for smoothing approximation of observed species richness, and Chao2 is the incidence-based coverage estimator. Error bars show the standard deviation for Mao’s tau and Chao2. Rarefaction and Chao 2 were calculated for each site using the lowest number of individuals observed across all stands (1260 individuals). The JE plot was not included in these comparisons due to the low number of understory herbs sampled.
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Figure 4. Bar graphs with standard error for (a) mean density (ramet m−2) and (b) mean percentage coverage in 25 m2 quadrats for understory herbs in young secondary (light-gray bars), intermediate secondary (medium-gray bars), and mature (black bars) forest stands.
Figure 4. Bar graphs with standard error for (a) mean density (ramet m−2) and (b) mean percentage coverage in 25 m2 quadrats for understory herbs in young secondary (light-gray bars), intermediate secondary (medium-gray bars), and mature (black bars) forest stands.
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Figure 5. Non-metric multidimensional scaling plot of herbs in young (EB, JE), intermediate (LSUR, LEPS), and mature (LEPP, SV) forest plots.
Figure 5. Non-metric multidimensional scaling plot of herbs in young (EB, JE), intermediate (LSUR, LEPS), and mature (LEPP, SV) forest plots.
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Table 1. Site characteristics for secondary and mature forest monitoring plots in northeastern Costa Rica.
Table 1. Site characteristics for secondary and mature forest monitoring plots in northeastern Costa Rica.
SiteLatitude, LongitudeLocationAge in 2006 (yrs)Prior Land-UseLandscape Matrix
Juan Enriquez (JE)10.46° N, 84.07° WChilamate12 Pasture
(ca. 12 yr)		Pasture, secondary, and mature forests
El Bejuco(EB)10.46° N, 84.67° WChilamate12 Pasture
(ca. 10 yr)		Pasture, secondary, and mature forests
Lindero Sur (LSUR)10.41° N, 84.03° WLa Selva 21 Pasture
(ca. 5 yr)		Mature and secondary forests
Peje Second- growth (LEPS)10.43° N, 84.03° WLa Selva 29 Pasture
(ca. 5 yr)		Mature and secondary forests
Selva Verde (SV)10.44° N, 84.07° WChilamate– Mature forestPasture, secondary, and mature forests
Peje Old-growth (LEPP)10.42° N, 84.04° WLa Selva –Mature ForestMature and secondary forests
Table 2. Mean cover values of broad-leaved understory herb species identified in six 1 ha forest plots across a chronosequence of forest stands in northeastern Costa Rica. Degree of clonality is based on qualitative field observations (SC = strongly clonal; C = clonal; WC = weakly clonal; NC = non-clonal).
Table 2. Mean cover values of broad-leaved understory herb species identified in six 1 ha forest plots across a chronosequence of forest stands in northeastern Costa Rica. Degree of clonality is based on qualitative field observations (SC = strongly clonal; C = clonal; WC = weakly clonal; NC = non-clonal).
YOUNG INTERMEDIATE MATURE CLONALITY
FAMILYSPECIESJEEBLSURLEPSSVLEPP
AraceaeAnthurium lancifolium 0.8 0.1NC
Anthurium ochranthum1.75.811.7105 NC
Dieffenbachia grayumiana17.526.75.80.80.82.5WC
Dieffenbachia hammelii 3.3 NC
Dracontium gigas0.8 NC
Philodendron grandipes0.84.232.541.70.840.8SC
Spathiphyllum friedrichsthalii 0.8SC
Spathiphyllum fulvovirens0.84.212.558.324.266.7NC
Spathiphyllum laeve1.73.317.52033.339.2NC
Spathiphyllum phryniifolium1.78.337.55.828.35WC
CostaceaeCostus bracteatus0.80.813.311.76.76.7C
Costus laevis104.2 2.5 C
Costus malortieanus54.230.08.30.83.3SC
Costus pulverulentus 5.01.7 WC
Costus scaber 8.320.81514.2WC
CyclanthaceaeAsplundia longipetula 0.8 1.7NC
Asplundia sleeperae 1.7NC
Asplundia uncinata 6.75.00.813.3C
Cyclanthus bipartitus11.719.27.529.233.320.8SC
Carludovica sulcata 0.8 SC
Dicranopygium umbrophilum 14.270.8NC
HeliconiaceaeHeliconia hursuita6.7
Heliconia imbricata 0.8 WC
Heliconia irrasa 46.717.518.311.7NC
Heliconia latispatha8.35.81.7 7.5 C
Heliconia mathiasiae 1.714.29.2 2.5C
Heliconia pogonantha 1.7 C
Heliconia umbrophila 1.72.55.0NC
MarantaceaeCalathea lasiostachya 1.715.012.512.52.5NC
Goeppertia cleistantha 65.052.515.07.5C
Goeppertia hammelii 0.1C
Goeppertia inocephala 30.817.50.8 NC
Goeppertia leucostachys 0.8 WC
Goeppertia micans9.287.58078.343.335SC
Goeppertia venusta 0.14.2NC
Ischnosiphon inflatus0.80.80.85.313.311.7WC
Pleiostachya pruinosa 0.80.1 WC
ZingiberaceaeRenealmia cernua 25.83.3 1.7WC
Renealmia pluriplicata 0.89.28.35.82.5WC
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Sun, J.W.C.; Chazdon, R.L.; Rundel, P.W. Diversity and Distribution of Monocot Understory Herbs during Tropical Forest Succession in Northeastern Costa Rica. Diversity 2024, 16, 439. https://doi.org/10.3390/d16080439

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Sun JWC, Chazdon RL, Rundel PW. Diversity and Distribution of Monocot Understory Herbs during Tropical Forest Succession in Northeastern Costa Rica. Diversity. 2024; 16(8):439. https://doi.org/10.3390/d16080439

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Sun, Jennifer W. C., Robin L. Chazdon, and Philip W. Rundel. 2024. "Diversity and Distribution of Monocot Understory Herbs during Tropical Forest Succession in Northeastern Costa Rica" Diversity 16, no. 8: 439. https://doi.org/10.3390/d16080439

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