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Understanding of tropical forests has been revolutionized by monitoring in permanent plots. Data from global plot networks have transformed our knowledge of forests’ diversity, function, contribution to global biogeochemical cycles, and sensitivity to climate change. Monitoring has thus far been concentrated in rain forests. Despite increasing appreciation of their threatened status, biodiversity, and importance to the global carbon cycle, monitoring in tropical dry forests is still in its infancy. We provide a protocol for permanent monitoring plots in tropical dry forests. Expanding monitoring into dry biomes is critical for overcoming the linked challenges of climate change, land use change, and the biodiversity crisis. Understanding of tropical forests has been revolutionised by monitoring in permanent plots. Data from global plot networks have transformed our knowledge of forests' diversity, function, contribution to global biogeochemical cycles, and sensitivity to climate change. Monitoring has thus far been concentrated in rain forests. Despite increasing appreciation of their threatened status, biodiversity, and importance to the global carbon cycle, monitoring in tropical dry forests is still in its infancy. We provide a protocol for permanent monitoring plots in tropical dry forests. Expanding monitoring into dry biomes is critical for overcoming the linked challenges of climate change, land use change, and the biodiversity crisis.
Plants, People, Planet. 2020;00:1–6.
Received: 11 Februar y 2020 
  Revised: 3 April 2020 
  Accepted: 20 April 2020
DOI: 10.1002/ppp3.10112
Expanding tropical forest monitoring into Dry Forests: The
DRYFLOR protocol for permanent plots
Peter W. Moonlight1| Karina Banda-R2,3| Oliver L. Phillips2| Kyle G. Dexter1,4|
R. Toby Pennington1,5| Tim R. Baker2| Haroldo C. de Lima6| Laurie Fajardo7|
Roy González-M.8| Reynaldo Linares-Palomino9,10 | Jon Lloyd11|
Marcelo Nascimento12| Darién Prado13| Catalina Quintana14| Ricarda Riina15| Gina
M. Rodríguez M.3| Dora Maria Villela16| Ana Carla M. M. Aquino17| Luzmila Arroyo18|
Cidney Bezerra19| Alexandre Tadeu Brunello17| Roel J. W. Brienen2|
Domingos Cardoso20| Kuo-Jung Chao21 | Ítalo Antônio Cotta Coutinho22|
John Cunha23| Tomas Domingues17| Mário Marcos do Espírito Santo24| Ted
R. Feldpausch5| Moabe Ferreira Fernandes25| Zoë A. Goodwin1| Eliana
María Jiménez26| Aurora Levesley2| Leonel Lopez-Toledo27| Beatriz Marimon28|
Raquel C. Miatto17| Marcelo Mizushima25| Abel Monteagudo29| Magna Soelma Beserra
de Moura30| Alejandro Murakami18| Danilo Neves31| Renata Nicora Chequín13|
Tony César de Sousa Oliveira17| Edmar Almeida de Oliveira28| Luciano P. de Queiroz25|
Alan Pilon32| Desirée Marques Ramos33| Carlos Reynel9| Priscyla M. S. Rodrigues34|
Rubens Santos35| Tiina Särkinen1| Valdemir Fernando da Silva36| Rodolfo M.
S. Souza36,37| Rodolfo Vasquez29| Elmar Veenendaal38
1Tropical Biodiversity, Royal Bot anic Garden Edinburgh, Edinburgh, UK
2School of Geography, Faculty of Environment, University of Leeds, Leeds, UK
3Fundación Ecosistemas Secos de Colombia, Barranquilla, Colombia
4School of Geosciences, The University of Edinburgh, Edinburgh, UK
5Geography, College of Life and Environmental Sciences, University of Exeter, Exeter, UK
6Instituto de Pesquisas, Jardim Botanico do Rio de Janeiro, Rio de Janeiro, Brazil
7Centro de Ecologia, Instituto Venezolano de Investigaciones Cientificas, Caracas, Venezuela
8Ciencias Básicas de la Biodiversidad, Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia
9Herbario, Departamento Académico de Biología, Universidad Nacional Agraria L a Molina, Lima, Peru
10Center for Conservation and Sustainability, Smithsonian Conser vation Biology Institute, Washington, DC, USA
11Depar tment of Life Sciences, Imperial College London, Ascot, UK
12Laboratório de Ciências Ambientais, Universidade Estadual do Norte Fluminense, Campos Dos Goytacazes, Brazil
13Instituto de Investigaciones en Ciencias Agrarias de Rosario (IICAR), Facultad Ciencias Agrarias, UNR , Universidad Nacional de Rosario, Santa Fe, Argentina
14Escuela de Biología, Facultad de Ciencias Exactas, Pontificia Universidad Católica del Ecuador, Quito, Ecuador
15Real Jardín Botánico, CSIC, Madrid, Spain
16Laboratório de Ciências Ambientas, Universidade Estadual do Norte Fluminense, Campos Dos Goytacazes, Brazil
17Depar tamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de Sao Paulo, Ribeirão Preto, Brazil
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2020 The Authors, Plants, People, Planet © New Phytologist Trust
18Museo de Historia Natural Noel Kempff Mercado, Universidad Autonoma Gabriel Rene Moreno, Santa Cruz de la Sierra, Bolivia
19Unidade Acadêmica de Garanhuns, Universidade Federal Rural de Pernambuco, Recife, Brazil
20Instituto de Biologia, Universidade Federal da Bahia, Salvador, Brazil
21International Master Program of A griculture, National Chung Hsing University, Taichung, Taiwan
22Departamento de Biologia Vegetal, Universidade Federal do Ceará, For taleza, Brazil
23Centro de Tecnologia e Recursos Naturais (CTRN), Universidade Federal de Campina Grande, Campina Grande, Brazil
24Programa de Pós-Graduação em Ciências Biológicas (PPGCB), Centro Ciências Biológicas e da Saúde, Universidade Estadual de Montes Claros, Montes
Claros, Brazil
25Ciencias Biologicas, Universidade Estadual de Feira de Santana, Feira de Santana, Brazil
26Grupo de Ecología y Conservación de Fauna y Flora Silvestre, Universidad Nacional de Colombia Facultad de Ciencias, Bogota, Colombia
27Instituto de Investigaciones sobre los Recursos Naturales, Universidad Michoacana de San Nicolás de Hidalgo, Morelia, Mexico
28Universidade do Estado de Mato Grosso, Campus de Nova Xavantina, Brazil
29Herbario HOXA, Jardín Botánico de Missouri, Oxapampa, Peru
30Embrapa Semiárido, Embrapa, Petrolina, Brazil
31Biologia Vegetal, Universidade Federal de Minas Gerais, Belo Horizonte, Brazil
32Faculdade de Ciências Farmacêuticas de Ribeirão Preto, Universidade de Sao Paulo, Ribeirão Preto, Brazil
33Laboratório de Fenologia, Departamento de Botânica, Instituto de Biociências, Universidade Estadual Paulista, Rio Claro, Brazil
34Departamento de Biologia Vegetal, Universidade Federal de Viçosa, Viçosa, Brazil
35Departamento de Ciências Florestais, Universidade Federal de Lavras, Lavras, Brazil
36Department of Forest Sciences, Universidade Federal Rural de Pernambuco, Recife, Brazil
37Depar tamento de Ciências Atmosféricas, Instituto de Astronomia, Geofífisica e Ciências Atmosféricas, Universidade de São Paulo, Ribeirão Preto, Brazil
38Plant Ecology and Nature Conservation, Wageningen University and Research, Wageningen, The Netherlands
Peter W. Moonlight, Tropical Biodiversity,
Royal Botanic Garden Edinburgh, 20A
Inverleith Row, Edinburgh, EH3 5LR, UK.
Funding information
Newton Fund, Grant/Award Number: NE/
N000587/1, NE/N01247X/1 and NE/
N012550/1; Natural Environment Research
Council, Grant/Award Number: NE/
I027797/1 and NE/I028122/1; Fundação de
Amparo à Pesquisa do Estado de São Paulo,
Grant/Award Number: 2015/50488-5;
CYTED, Grant/Award Number: 418RT0554
Societal Impact Statement
Understanding of tropical forests has been revolutionized by monitoring in perma-
nent plots. Data from global plot networks have transformed our knowledge of for-
ests’ diversity, function, contribution to global biogeochemical cycles, and sensitivity
to climate change. Monitoring has thus far been concentrated in rain forests. Despite
increasing appreciation of their threatened status, biodiversity, and importance to the
global carbon cycle, monitoring in tropical dry forests is still in its infancy. We provide
a protocol for permanent monitoring plots in tropical dry forests. Expanding monitor-
ing into dry biomes is critical for overcoming the linked challenges of climate change,
land use change, and the biodiversity crisis.
floristics, long term plots, tropical dry forests, vegetation dynamics, vegetation structure
Long-term forest plots are sites where all trees above a specified
diameter are numbered, identified, and measured, and where re-
peated censuses record growth, mortality, and recruitment. Such
plots have become widespread in tropical rain forests, exempli-
fied by networks such as RAINFOR (Amazon Forest Inventory
Network; Malhi et al., 2002), AfriTRON (African Tropical Rainforest
Observation Network; Lewis et al., 2009), T-FORCES (Tropical
Forests in the Changing Earth System; Qie et al., 2017), and CTFS-
forestGEO (Center for Tropical Forest Science-Forest Global Earth
Observatory; Anderson-Teixeira et al., 2014). The RAINFOR,
AfriTRON, and T-FORCES networks collectively comprise > 1,000
1 ha plots across the tropics, where every tree with a stem diam-
eter ≥ 10 cm is measured. CTFS-forestGEO employs much larger
(often 50 ha) plots where every stem ≥ 1 cm in diameter is measured,
and this more intensive survey means that there are fewer (<100) of
such plots across the tropics.
These long-term tropical rain forest plots have been extremely
successful in achieving their primary aim of improving our knowl-
edge of tropical forest ecology, including, for example: the rela-
tionships of climate with biomass lvarez-Dávila et al., 2017) and
forest structure (Feldpausch et al., 2012); the role of diversity in car-
bon storage and productivity (Coelho de Silva et al., 2019; Sullivan
et al., 2017); and drivers of monodominance in Amazonia (ter Steege
et al., 2019). In addition, they have helped increase understanding
of community floristic diversity and composition (Baker et al., 2016;
Guevara et al., 2016; Levis et al., 2017), continental scale floristic
patterns (Esquivel-Muelbert et al., 2017; ter Steege et al., 2006; ter
Steege, Pitman, Sabatier, Baraloto, & Salomão, 2013), biome delim-
itation, and mapping (Silva-de-Miranda et al., 2018), and even facil-
itated the discovery of species new to science (reviewed by Baker
et al., 2017). Repeated censuses of these plots have provided insight
into the role of tropical forests in global cycles of carbon, energy,
and water (Pan et al., 2011; Phillips et al., 1998), long-term trends in
forest dynamics (Brienen et al., 2015), and the impacts of extreme
climatic events (Feldpausch et al., 2016; Phillips et al., 2009). As
such, these international standardized networks are a helpful mac-
roecological tool to study humanity's effect on the Earth system and
the vital role that moist tropical forests play in carbon sequestration
and therefore in mitigating the effects of increasing concentration
of atmospheric CO2. Conversely, they have also demonstrated how
tropical forest destruction and degradation account for an estimated
1.3 Pg carbon emissions (Malhi, 2010) and that, following deforesta-
tion, the recovery of forest species composition can take centuries
(Rozendaal et al., 2019). They may also have critical implications at
national levels too - in Peru, for example, long-term permanent plots
have been used to show that the country's intact rain forests have
helped to remove 86% of the country's emissions from the combus-
tion of fossil fuels (Vicuña-Minaño et al., 2018).
Long-term monitoring started in tropical rain forests and has been
concentrated there since. This reflects the importance of such
forests as the largest above-ground terrestrial carbon stock (Pan
et al., 2011) and their unparalleled levels of local (alpha) diversity of
plants and animals (e.g. Bass et al., 2010). However, half of the global
tropics are too seasonally dry to support such forests and instead
are home to tropical dry forests (Figure 1) and savannas (Pennington,
Lehmann, & Rowland, 2018). An estimated one-third of the global
population inhabits the seasonally dry tropics (GLP, 2005), and, as
a consequence, these systems have been commonly and severely
altered (e.g., Fajardo et al., 2005; Janzen, 1988; Linares-Palomino,
Kvist, Aguirre-Mendoza, & Gonzales-Inca, 2010; Portillo-Quintero &
Sánchez-Azofeifa, 2010). Because they can be erroneously viewed
as semi-natural, and because of their smaller stature and lower local
diversity than rain forests, tropical dry forests have been under-ap-
preciated by science and conservation. However, new information
suggests that their floristic diversity at continental scale (gamma
diversity) may approach that of rain forests (Flora do Brasil, 2020;
DRYFLOR, 2016), and that they play an essential role in controlling
the interannual variability in the global carbon cycle (Poulter, Frank,
Ciais, Myneni, & Andela, 2014). It is clear that science and society
cannot continue to largely ignore these tropical dry biomes.
Even thirty years ago tropical dry forests were already considered
the most threatened tropical biome on the planet (Janzen, 1988),
and less than 10% of their original extent remains in many Latin
American countries, which house the largest remaining areas of this
vegetation (Miles et al., 2006; Pennington et al., 2018; Pennington,
Prado, & Pentry, 2000). This high level of loss is not only due to re-
cent conversion but also is a reflection of a long history of defor-
estation and use by early civilizations inhabiting dry forest areas,
especially in Latin America (Murphy & Lugo, 1986).
Landscape modification in tropical dry forest areas has been
exacerbated by their frequently fertile soils, and this also makes
them a continuing focus for agricultural expansion. Although at
local scales plant species richness in tropical dry forests does
FIGURE 1 Dry forest in El Coto de
Caza El Angolo, Piura, Peru in the dry
season showing Ceiba trichistandra (A.
Gray) Bakh. Photograph taken by P.W.
not match that of tropical rain forest, in the Neotropics, at least,
high floristic turnover amongst areas means that at continental
scale their species diversity rivals that of rain forest. For example,
DRYFLOR (Latin American Seasonally Dry Tropical Forest Floristic
Network; 2016) recorded 6,958 woody species from just 1,602
surveys, whereas a current estimate of the number of tree spe-
cies in the moist forests of the Amazon Basin is 6,727 (Cardoso
et al., 2017).
Despite this diversity, tropical dry forests are woefully un-
der-protected. For example, only 1.2% of remaining Brazilian
Caatinga dry forest and 1.4% of Colombian inter-Andean dry
forest are protected (García, Corzo, Isaacs, & Etter, 2014; MMA,
2016), falling massively short of the 17% target set by Aichi bio-
diversity target 11 (CBD, 2011). An integral part of improving the
conservation outlook for tropical dry forests, and of gaining vital
information relevant to their restoration, will come from long-term
ecological monitoring. Such monitoring will be essential to under-
stand how their species grow, reproduce, and recruit, and the
mechanisms behind their mortality, especially in times of climatic
and environmental changes.
The rapid growth of long-term forest monitoring in tropical
rain forests partly reflects internationally agreed, standard proto-
cols for plot establishment. Conversely, the slow adoption of mon-
itorin g in dry biom es is a cons equence, among other factors, of the
lack of agreed protocols. Such lack of consensus in part reflects
the wide physiognomic spectrum of tropical savannas and dry for-
ests. For dry forests, the focus of this paper, this can vary from tall,
closed forest with a 25–30 m canopy, to more open, low, thorny,
and cactus scrub (Pennington et al., 2000). Protocols designed for
1 ha plots in the moist tropics (e.g. Phillips, 2018) fail to capture
th e maj ori t y of gr owt h, mor talit y, or re cruitm ent dy nam ics in these
systems, primarily because mature individuals of many species do
not reach a minimum diameter at breast height (DBH) of 10 cm.
These smaller trees play an important role when describing struc-
ture and functioning of dry forest vegetation (Torello-Raventos
et al., 2013). We urgently need a standard for systematizing the
way with which the large number of researchers now working in
dry forests can measure and monitor these ecosystems. Only with
such a standard protocol in place can we lay the foundations for
generating a rich legacy of scientific and practical advancement in
ecology across the tropics.
In response to this urgent need we here present an approach in
measuring and monitoring tropical ecosystems, specifically adapted
to meet the challenges of long term monitoring in dry forests.
Our protocol, the DRYFLOR Field Manual for Plot Establishment and
Remeasurement (“DRYFLOR Plot Protocol”; please see the Supporting
Information for English, Portuguese and Spanish versions of the
protocol), is based on wide tropical experience and has received rig-
orous field testing in the dry forests, semi-deciduous forests, and
related dr y biomes of Pe ru, sou theas t, an d nor theas t Brazil. The pro-
tocol design is modified and expanded from that used by R AINFOR
(The Amazon Forest Inventory Network; Phillips, Baker, Feldpausch,
& Brienen, 2018) across the Americas and beyond with a particular
emphasis on the Amazon Basin. The new DRYFLOR Plot Protocol cap-
tures most dry forest structure and dynamics and is specifically de-
signed to enable a full and detailed comparison with data captured
by humid forest protocols (Phillips et al., 2018) and by savanna and
dry forest protocols (e.g. by measuring stems ≥ 5 cm diameter and at
130 and 30 cm, rather than ≥10 cm diameter at only 130 cm; in its
provisions for multi stemmed individuals). Physiognomic and dynam-
ics data from the protocol are fully compatible with the ForestPlots
database (Lopez-Gonzalez, Lewis, Burkitt, & Phillips, 2011) and flo-
ristic data with the DRYFLOR database (www.dryfl We be-
lieve it reaches a reasonable compromise between practical field
constraints in terms of time and data captured for the purpose of
estimating species abundances and biomass data, but it also pro-
vides optional modules that can be implemented if a more complete
picture of dry forest dynamics is desired.
The DRYFLOR Plot Protocol is a product of a large, collaborative net-
work of researchers working across Latin American dry forests and
related dry biomes. It is intended to permit the rapid and efficient
collection of inventory data in the dry tropics and facilitate stud-
ies on the structure and function of forests. The development of
this protocol is indebted to both the R AINFOR and the DRYFLOR
networks and three projects funded from 2011 to 2019 by the UK
Research Councils and the Brazilian Research Foundations FAPESP
and FAPERJ. The uptake of the protocol in new geographic areas and
beyond these networks will be a continuing challenge, but provides
the considerable benefit of standardised data capture. This will en-
able further collaborative research at wider spatial scales that is vital
for addressing questions about the current and future ecology of
tropical forests in a rapidly changing world. The societal relevance
of this research will ultimately depend not simply on the application
of a universal dry forest protocol, but also on the development of
lasting, meaningful relationships with local and regional stakehold-
ers and policymakers.
This paper was conceived at two DRYFLOR meetings funded by
CYTED (Iberoamerican Program of Science and Technology net-
work grant #418RT0554). The protocol was designed and tested
across three projects: NERC-Newton-FAPESP Nordeste: New
Science for A Neglected Biome (#NE/N01247X/1; #NE/N012550/1;
#2015/50488-5); NERC-Newton-FAPERJ Dry Forest Biomes in
Brazil: Biodiversity and Ecosystem Services; (#NE/N000587/1); NERC
Niche Evolution of South American Trees and its Consequences (#NE/
I027797/1; #NE/I028122/1). We are grateful for the active involve-
ment of the RAINFOR and DRYFLOR networks; all countries, land-
owners and agencies who have granted us permission and provided
logistical support during the protocol testing; and the support of all
author´s institutions.
T.P. conceived the idea and P.M. led the writing of the manuscript
and plot protocol, with significant input from authors K.B.-R. to
D.M.V. All authors contributed to the design and field testing of the
protocol, and had input in the manuscript. Portuguese translation of
the Supporting Information was done by A.T.B., D.M.V., D.R.M., I.C.,
M.N. T.C.d.S.O, and R.C.M.; Spanish translation was done by C.Q,
K.B.-R., R.L.-P., and R.R.
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Tim R. Baker
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Additional supporting information may be found online in the
Supporting Information section.
How to cite this article: Moonlight PW, Banda-R K, Phillips
OL, et al. Expanding tropical forest monitoring into Dry
Forests: The DRYFLOR protocol for permanent plots. Plants,
People, Planet. 2020;00:1–6.
... The latter consists in promoting the natural resilience of a degraded forest until it reaches a climax state. Such an approach is therefore based on a thorough knowledge of the ecological processes of natural recovery that develop after forest degradation [5]. Indeed, the monitoring of post-cultivation ecological processes is fundamental to the maintaining floristic richness [6] and planning forest recovery [5,7]. ...
... Such an approach is therefore based on a thorough knowledge of the ecological processes of natural recovery that develop after forest degradation [5]. Indeed, the monitoring of post-cultivation ecological processes is fundamental to the maintaining floristic richness [6] and planning forest recovery [5,7]. ...
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Ivorian classified forests have been highly anthropized by cocoa farming. In an attempt to provide guidance to the government on approaches to the restoration of the forest while respecting the aspirations of local populations, permanent plots were set up in the classified forest of Haut-Sassandra, and were monitored and measured for 3 years. This study was intended to analyze the evolution of the vegetation of permanent plots in the classified forest of Haut-Sassandra from 2018 to 2021. The results show that the vegetation evolves with the cessation of some agricultural activities. These plantations are colonized by pioneer species during the first three years of the abandonment of agricultural activities. Mortality rates increased by 477.59% and recruitment rates were reduced by 61.87% in regularly maintained plantations compared to their condition three years ago. However, the plantations with no agricultural activities and those which were not maintained but harvested had the highest recruitment rates of pioneer and heliophilous individuals. In sum, tree species could recolonize the classified forest of Haut-Sassandra if clearing is prohibited in cocoa farms. However, the populations could continue to harvest the pods from the cocoa trees which are already established in the classified forest of Haut-Sassandra.
... To quantify the vegetation structure, measurements of stem diameters and projected canopy areas were made according to the protocols detailed in Torello-Raventos et al. (2013) and Moonlight et al. (2021). Tree height measurements were taken by holding a graduated pole close to the trunk. ...
The architecture of root systems is an important driver of plant fitness, competition and ecosystem processes. However, the methodological difficulty of mapping roots hampers the study of these processes. Existing approaches to match individual plants to belowground samples are low‐throughput and species‐specific. Here, we developed a scalable sequencing‐based method to map the root systems of individual trees across multiple species. We successfully applied it to a tropical dry forest community in the Brazilian Caatinga containing 14 species. We sequenced all 42 individual shrubs and trees in a 14 by 14 m plot using double‐digest restriction‐site associated sequencing (ddRADseq). We identified species‐specific markers and individual‐specific haplotypes from the data. We matched these markers to ddRADseq data from 100 mixed root samples from across the centre (10 by 10 m) of the plot at four different depths, using a newly developed R package. We identified individual root samples for all species and all but one individual. There was a strong significant correlation between below and aboveground size measurements, and we also detected significant species‐level root‐depth preference for two species. The method is more scalable and less labour‐intensive than current techniques, and is broadly applicable to ecology, forestry and agricultural biology.
... The manual is based on: the outcomes from two workshops; analysis of the current version of the SEOSAW dataset; and a review of similar guidance developed for the dry tropics (DRYFLOR et al., 2016;Guerin et al., 2017;Moonlight et al., 2020;White et al., 2012) and wet tropics (Alder & Synnott, 1992;Condit, 1998;Lewis et al., 2009b;Malhi et al., 2002;Phillips et al., 2016;Picard et al., 2008;Poorter et al., 2016). The participants at the workshops, and in subsequent discussions, are listed at the end of the paper. ...
• Here we describe a new network of researchers and long-term, in situ, measurements that will characterize the changing socio-ecology of the woodlands of southern Africa. These woodlands encompass the largest savanna in the world, but are chronically understudied, with few long-term measurements. • A network of permanent sample plots (PSPs) is required to: (a) address management issues, particularly related to sustainable harvesting for energy and timber; (b) understand how the woodlands are responding to a range of global and local drivers, such as climate change, CO2 fertilization, and harvesting; and (c) answer basic questions about biogeography, ecosystem function, and the role humans play in shaping the ecology of the region. • We draw on other successful networks of PSPs and adapt their methods to the specific challenges of working in southern African woodlands. In particular we suggest divergences from established forest monitoring protocols that are needed to (a) adapt to a high level of ecosystem structural diversity (from open savanna to dry forest); (b) quantify the chronic disturbances by people, fire, and herbivores; (c) quantify the diversity and function of the understory of grasses, forbs, and shrubs; (d) understand the life histories of resprouting trees; and (e) conduct work in highly utilized, human-dominated landscapes. • We conclude by discussing how the SEOSAW network will integrate with remote sensing and modeling approaches. Throughout, we highlight the challengesinherent to integrating work by forest and savanna ecologists, and the wide range of skills needed to fully understand the socio-ecology of the southern African woodlands.
... Among tropical forests, seasonally dry forests (TDFs) account for 42% of the world's tropical forests [5], are extensive, but little known in their structure and function compared to tropical rainforests [6,7]. In the Neotropical region, TDFs have been globally reduced by 49% to 66% of their original coverage, occurring in patches, immersed in landscapes dominated by crops and livestock areas [8,9]. ...
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Tropical dry forests (TDFs) represent 42% of all tropical forests; they are extensive, but little is known of their structure and function. The fine litterfall represents the main route of circulation of organic materials and nutrients in these ecosystems. The objective of this study was to compare several remnants of TDFs located in contrasting landscape units—Mountain and Lomerio—and with different precipitation, in terms of the fluxes of organic materials to the soil, derived from the production of fine litterfall from the canopy. The fine litterfall (including woody material up to 2 cm in length) was collected monthly from April 2020 to March 2021, in 29 circular plots of 500 m2 randomly established. High rates of litterfall were recorded in the Lomerio landscape (4.9 Mg ha−1) than in the Mountain landscape (4.5 Mg ha−1). The monthly leaf litter production showed clear seasonal patterns, which were largely driven by the importance of the species in the landscape and the effect of precipitation during the study. Annual fine litter production observed in this study in comparison with other TDFs indicates relevant productivity levels, which contribute to the activation of biogeochemical cycles and improved ecosystem functionality.
... Currently, in addition to existing permanent plots in tropical humid forests, efforts to establish new permanent monitoring plots in areas with limited forest monitoring are underway. Future projects such as SECO ( seek to monitor carbon stocks and sinks in tropical dry forests and woodlands, with adapted protocols for the setup of permanent plots (Moonlight et al., 2021). Funding long-term monitoring projects is challenging due to current financing schemes in academia, which are often linked to short-term outputs. ...
... However, fine-scale early warnings of collapse could potentially be encompassed in ecosystem-specific indicators, and those derived from Red List of Ecosystems assessments (RLIE and EHI), which use ecosystem-specific variables that include ecological complexities, such as connectivity between ecosystem types 133,134 . Data, classifications and maps of ecosystems and integrity would be greatly strengthened by a globally coordinated program to unite remotely sensed data and field observations for training and testing of spatial outputs for ecosystems (comparable to those proposed for plant functional diveristy 125 ) and coordinated experiments and sampling 135,136 . Studies on specific ecosystem types provide examples that unite field-based data collection and remotely sensed data 94,137 , while emerging methods can increasingly capitalize on the wealth of remotely sensed data 97 . ...
Despite substantial conservation efforts, the loss of ecosystems continues globally, along with related declines in species and nature’s contributions to people. An effective ecosystem goal, supported by clear milestones, targets and indicators, is urgently needed for the post-2020 global biodiversity framework and beyond to support biodiversity conservation, the UN Sustainable Development Goals and efforts to abate climate change. Here, we describe the scientific foundations for an ecosystem goal and milestones, founded on a theory of change, and review available indicators to measure progress. An ecosystem goal should include three core components: area, integrity and risk of collapse. Targets—the actions that are necessary for the goals to be met—should address the pathways to ecosystem loss and recovery, including safeguarding remnants of threatened ecosystems, restoring their area and integrity to reduce risk of collapse and retaining intact areas. Multiple indicators are needed to capture the different dimensions of ecosystem area, integrity and risk of collapse across all ecosystem types, and should be selected for their fitness for purpose and relevance to goal components. Science-based goals, supported by well-formulated action targets and fit-for-purpose indicators, will provide the best foundation for reversing biodiversity loss and sustaining human well-being.
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The Seasonally Dry Tropical Forests and Woodlands biome (SDTFW) has its largest nucleus in the Caatinga domain. We characterized the flora and physiognomy of the vegetation in the Pedra da Andorinha Wildlife Refuge (RPA), in Ceará, Brazil. Plant collections were made between March/2015 and May/2021, applying usual botanical methods. All material was deposited in HUVA herbarium. The phytosociological studies covered five 50 m x 50 m plots established (1.25 ha) in which all individuals with diameters at soil level ³ 3 cm were inventoried. We identified 266 vascular plants species distributed among 185 genera and 67 families, including one fern (Marsilea deflexa - Marsileaceae). Fabaceae had the greatest species richness (38 spp.), while Ipomoea was the richest genus (9 spp.). 43.6% of all plant species were herbaceous, with a predominance of therophytes (57.5% of all herbaceous plants). The phytosociological study sampled 1,988 individuals distributed among 24 species of 13 families. The species with the greatest important value were Cordia oncocalyx (Boraginaceae) and Croton blanchetianus (Euphorbiaceae). We classify the local physiognomy as typical caatinga sensu stricto vegetation and rocky vegetation on inselbergs and outcrops. We highlight the richness of herbaceous plants in the local community, which surpass the richness of the woody component.
Soil microbial communities are crucial in ecosystem diversity and are directly related to soil fertility. Lombok is an island in central Indonesia that has low soil fertility and a limited amount of available water. Beneficial microorganisms can be used as a low-cost and environmental friendly tool to increase productivity in dryland agriculture systems. Screening to obtain superior rhizosphere bacteria is one of the options to support the nutrient supply in arid soils. Composites soil samples were taken from five ecosystems in Lombok, West Nusa Tenggara, an arid region in the eastern part of Indonesia to obtain the isolates of nitrogen-fixer rhizobacteria (NFR). Nine Azotobacter and Azopirillium spp were isolated from, rainfed, maize, mixed crop, natural forests, and savanna ecosystems. Abundance of total bacteria and N2-fixers in all ecosystems was relatively high (more than 108 cfu g-1), and the highest total population was recorded in the natural forest. The abundance of N2-fixer rhizobacteria recorded the highest Azotobacter population at 2.64 x 108 cfu g-1 in the maize ecosystem and the highest Azospirillum population at 2.32 x 108 cfu g-1 in the natural forest ecosystem. Additionally, the highest contain of organic C and total nitrogen were obtained in natural forest and savanna ecosystem. Eighteen isolates were obtained and characterized microscopic and macroscopically, consisted of nine Azotobacter sp and nine Azospirillium isolates which are potentially to be used as biogent for improving the growth of upland rice on dry climate zone.
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Vegetation is a key biosphere component to supporting biodiversity on Earth, and its maintenance and proper functioning are essential to guarantee the well-being of humankind. From a broad perspective, a fundamental goal of vegetation ecology is to understand the roles of abiotic and biotic factors that affect vegetation structure, distribution, diversity, and functioning, considering the relevant spatial and temporal scales. In this contribution, we reflect on the difficulties and opportunities to accomplish this grand objective by reviewing recent advances in the main areas of vegetation ecology. We highlight theoretical and methodological challenges and point to alternatives to overcome them. Our hope is that this contribution will motivate the development of future research efforts that will strengthen the field of vegetation ecology. Ultimately, vegetation science will continue to provide a strong knowledge basis and multiple theoretical and technological tools to better face the current global environmental crisis and to address the urgent need to sustainably conserve the vegetation cover of our planet in the Anthropocene.
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The tropical dry forest (TDF) is one of the most threatened ecosystems worldwide due to the historical conversion of its lands to pastures and crops. The TDF of the Cartagena Botanical Garden “Guillermo Piñeres” (JBGP), one of the last TDF relicts near Cartagena, is an isolated fragment of 3 ha located in a humid area caused by the presence of the spring of the Matute stream, one of the primary water sources of Cartagena. We built a 1 ha permanent plot to study the composition and plant structure of this forest. We measured the woody vegetation with a diameter at breast height (DBH) ≥ 2.5 cm. We recorded 1568 individuals and 2023 stems of 62 species distributed in 34 families. Of the individuals registered, 85 % were trees and shrubs, 11 % lianas and 4 % palms. The family with the highest species richness was Sapindaceae, with four, followed by Apocynaceae, Arecaceae, Fabaceae, Meliaceae, Moraceae, Nyctaginaceae, and Urticaceae, each with three species. We conclude that the forest is in a late secondary state of succession. Its location and history of disturbances are essential factors in determining its composition, structure, and diversity.
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Higher levels of taxonomic and evolutionary diversity are expected to maximize ecosystem function, yet their relative importance in driving variation in ecosystem function at large scales in diverse forests is unknown. Using 90 inventory plots across intact, lowland, terra firme, Amazonian forests and a new phylogeny including 526 angiosperm genera, we investigated the association between taxonomic and evolutionary metrics of diversity and two key measures of ecosystem function: aboveground wood productivity and biomass storage. While taxonomic and phylogenetic diversity were not important predictors of variation in biomass, both emerged as independent predictors of wood productivity. Amazon forests that contain greater evolutionary diversity and a higher proportion of rare species have higher productivity. While climatic and edaphic variables are together the strongest predictors of productivity, our results show that the evolutionary diversity of tree species in diverse forest stands also influences productivity. As our models accounted for wood density and tree size, they also suggest that additional, unstudied, evolutionarily correlated traits have significant effects on ecosystem function in tropical forests. Overall, our pan-Amazonian analysis shows that greater phylogenetic diversity translates into higher levels of ecosystem function: tropical forest communities with more distantly related taxa have greater wood productivity. Inventory data from 90 lowland Amazonian forest plots and a phylogeny of 526 angiosperm genera were used to show that taxonomic and phylogenetic diversity are both predictive of wood productivity but not of biomass variation.
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Tropical forests are known for their high diversity. Yet, forest patches do occur in the tropics where a single tree species is dominant. Such “monodominant” forests are known from all of the main tropical regions. For Amazonia, we sampled the occurrence of monodominance in a massive, basin-wide database of forest-inventory plots from the Amazon Tree Diversity Network (ATDN). Utilizing a simple defining metric of at least half of the trees ≥ 10 cm diameter belonging to one species, we found only a few occurrences of monodominance in Amazonia, and the phenomenon was not significantly linked to previously hypothesized life history traits such wood density, seed mass, ectomycorrhizal associations, or Rhizobium nodulation. In our analysis, coppicing (the formation of sprouts at the base of the tree or on roots) was the only trait significantly linked to monodominance. While at specific locales coppicing or ectomycorrhizal associations may confer a considerable advantage to a tree species and lead to its monodominance, very few species have these traits. Mining of the ATDN dataset suggests that monodominance is quite rare in Amazonia, and may be linked primarily to edaphic factors.
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Old-growth tropical forests harbor an immense diversity of tree species but are rapidly being cleared, while secondary forests that regrow on abandoned agricultural lands increase in extent. We assess how tree species richness and composition recover during secondary succession across gradients in environmental conditions and anthropogenic disturbance in an unprecedented multisite analysis for the Neotropics. Secondary forests recover remarkably fast in species richness but slowly in species composition. Secondary forests take a median time of five decades to recover the species richness of old-growth forest (80% recovery after 20 years) based on rarefaction analysis. Full recovery of species composition takes centuries (only 34% recovery after 20 years). A dual strategy that maintains both old-growth forests and species-rich secondary forests is therefore crucial for biodiversity conservation in human-modified tropical landscapes.
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Los bosques primarios intactos de la Amazonía peruana se comportan como sumideros de carbono: un servicio ecosistémico clave a nivel mundial. Este sumidero fue cuantificado en 0.54 Mg C ha-1 año-1 (1990-2017) para los bosques amazónicos intactos de las Áreas Naturales Protegidas (ANPs) de Perú y las zonas de amortiguamiento. En otras palabras, la conservación de bosques intactos en ANPs ayudó a remover 9.6 millones de toneladas de carbono de la atmósfera por año, lo cual equivale aproximadamente al 85% de las emisiones de la quema de combustibles fósiles del país durante el 2012. Este servicio de remoción de CO2 atmosférico es necesario incluir en el inventario nacional de gases de efecto invernadero, y en los compromisos nacionales de reducción de emisiones, por dos razones. Primero, debido a ser un flujo importante, nos ayudaría a tener una aproximación más real del balance de carbono en Perú. Segundo, fortalecería la necesidad de mantener la integridad de estos bosques tanto por el servicio de almacenamiento de carbono (evitar emisiones) como el servicio de sumidero (remoción de emisiones) y la diversidad biológica que albergan. La provisión del servicio de sumidero solo se asegurará con una gestión efectiva y adaptativa de las ANPs. El reporte de este servicio ambiental a nivel nacional debe ser implementado a través del monitoreo a largo plazo de la dinámica del carbono y el impacto del cambio climático a través de la red de parcelas forestales permanentes de RAINFOR (Red Amazónica de Inventarios Forestales) y el proyecto MonANPeru. El establecimiento de este sistema de monitoreo permitirá el desarrollo de los mecanismos financieros para cerrar la brecha y lograr la sostenibilidad de la conservación de los bosques en las ANPs de Perú.
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Less than half of anthropogenic carbon dioxide emissions remain in the atmosphere. While carbon balance models imply large carbon uptake in tropical forests, direct on-the-ground observations are still lacking in Southeast Asia. Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 per year (95% CI 0.14-0.72, mean period 1988-2010) above-ground live biomass. These results closely match those from African and Amazonian plot networks, suggesting that the world's remaining intact tropical forests are now en masse out-of-equilibrium. Although both pan-tropical and long-term, the sink in remaining intact forests appears vulnerable to climate and land use changes. Across Borneo the 1997-1998 El Niño drought temporarily halted the carbon sink by increasing tree mortality, while fragmentation persistently offset the sink and turned many edge-affected forests into a carbon source to the atmosphere.
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Significance Large floristic datasets that purportedly represent the diversity and composition of the Amazon tree flora are being widely used to draw conclusions about the patterns and evolution of Amazon plant diversity, but these datasets are fundamentally flawed in both their methodology and the resulting content. We have assembled a comprehensive dataset of Amazonian seed plant species from published sources that includes falsifiable data based on voucher specimens identified by taxonomic specialists. This growing list should serve as a basis for addressing the long-standing debate on the number of plant species in the Amazon, as well as for downstream ecological and evolutionary analyses aimed at understanding the origin and function of the exceptional biodiversity of the vast Amazonian forests.
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High levels of species diversity hamper current understanding of how tropical forests may respond to environmental change. In the tropics, water availability is a leading driver of the diversity and distribution of tree species, suggesting that many tropical taxa may be physiologically incapable of tolerating dry conditions, and that their distributions along moisture gradients can be used to predict their drought tolerance. While this hypothesis has been explored at local and regional scales, large continental-scale tests are lacking. We investigate whether the relationship between drought-induced mortality and distributions holds continentally by relating experimental and observational data of drought-induced mortality across the Neotropics to the large-scale bioclimatic distributions of 115 tree genera. Across the different experiments, genera affiliated to wetter climatic regimes show higher drought-induced mortality than dry-affiliated ones, even after controlling for phylogenetic relationships. This pattern is stronger for adult trees than for saplings or seedlings, suggesting that the environmental filters exerted by drought impact adult tree survival most strongly. Overall, our analysis of experimental, observational, and bioclimatic data across neotropical forests suggests that increasing moisture-stress is indeed likely to drive significant changes in floristic composition.
Tropical forests are known for their high diversity. Yet, forest patches do occur in the tropics where a single tree species is dominant. Such "monodominant" forests are known from all of the main tropical regions. For Amazonia, we sampled the occurrence of monodominance in a massive, basin-wide database of forest-inventory plots from the Amazon Tree Diversity Network (ATDN). Utilizing a simple defining metric of at least half of the trees >= 10 cm diameter belonging to one species, we found only a few occurrences of monodominance in Amazonia, and the phenomenon was not significantly linked to previously hypothesized life history traits such wood density, seed mass, ectomycorrhizal associations, or Rhizobium nodulation. In our analysis, coppicing (the formation of sprouts at the base of the tree or on roots) was the only trait significantly linked to monodominance. While at specific locales coppicing or ectomycorrhizal associations may confer a considerable advantage to a tree species and lead to its monodominance, very few species have these traits. Mining of the ATDN dataset suggests that monodominance is quite rare in Amazonia, and may be linked primarily to edaphic factors.
In the tropics, research, conservation and public attention focus on rain forests, but this neglects that half of the global tropics have a seasonally dry climate. These regions are home to dry forests and savannas (Figures 1 and 2), and are the focus of this Primer. The attention given to rain forests is understandable. Their high species diversity, sheer stature and luxuriance thrill biologists today as much as they did the first explorers in the Age of Discovery. Although dry forest and savanna may make less of a first impression, they support a fascinating diversity of plant strategies to cope with stress and disturbance including fire, drought and herbivory. Savannas played a fundamental role in human evolution, and across Africa and India they support iconic megafauna.