Received: 15 February 2020 Accepted: 15 February 2020
DOI: 10.1111/1440-1703.12101
SPECIAL FEATURE : PREFACE
Approaches for general rules of biodiversity patterns in
space and time
Yasuhiro Kubota1,2 | Buntarou Kusumoto1
1
Faculty of Science, University of the Ryukyus, Okinawa, Japan
2
Marine and Terrestrial Field Ecology, Tropical Biosphere Research Center, University of the Ryukyus, Okinawa, Japan
Correspondence
Yasuhiro Kubota, Faculty of Science, University of the Ryukyus, Okinawa, Japan.
Email: kubota.yasuhiro@gmail.com
Understanding the mechanisms for the origin and mainte-
nance of biodiversity is central to biogeography and ecol-
ogy. MacArthur (1972) advocated an exploration to search
for repeated patterns and not simply to accumulate facts.
Using this perspective, emergent macroecological patterns,
such as species–area relationship, species abundance dis-
tribution and decay of similarity with distance (beta diver-
sity) have been reported throughout different spaces, times
and taxa. Nonetheless, attempts in the search for general
(universal) laws generating biodiversity patterns have not
been sufficiently and/or positively argued (Hubbell, 2001;
McGill, 2019; Roughgarden, 2009). Lawton (1999) cast
doubt on the utility of traditional community ecology for
pursuing laws; this was because of the contingency in rela-
tion to the complicated interrelationship between organ-
isms and environments that makes it difficult to find
generalities and instead emphasized the predominance of
population ecology or macroecology. More importantly,
our scientific knowledge of biodiversity remains incom-
plete: indeed, Hortal et al. (2015) define the seven key
knowledge gaps involving species taxonomy (Linnean),
distribution (Wallacean), abundance (Prestonian), evolu-
tionary patterns (Darwinian), species-specific abiotic toler-
ances of (Hutchinsonian), species traits (Raunkiæran) and
biotic interactions (Eltonian) shortfalls. Our ignorance of
biodiversity causes uncertainty in ecological inference
and/or a high degree of contingency involved in target spe-
cies/communities, especially in microecological studies
focusing on particular taxa/sites, which may prevent the
exploration of general rules (even if they exist). In this con-
text, macroecological diversity patterns should always be
revisited in a more honest way using high-quality big data,
but there should not be a pessimistic discourse about the
Ecological Research. 2020;35:289–291. wileyonlinelibrary.com/journal/ere
absence of general theory in biodiversity science
(Simberloff, 2004). Such an approach helps to reduce
the scientific uncertainty related to our ignorance and
may allow us to explore the general law behind repeat-
edly observed diversity patterns. This will contribute to
implementing systematic conservation planning for cap-
turing irreplaceable biodiversity features.
Our research group of biodiversity and conservation bio-
geography (BCB Japan) aims to bridge between local-scale
ecological studies and the concepts/theories of biogeography
and macroecology. We use a variety of datasets (vegetation
plots, forest dynamics plots, species distribution, phylogeny,
functional traits and fossils) and statistical models including
mechanistic null models. For a detailed understanding of
biodiversity patterns, especially those in Asia, we investigate
hierarchical assembly processes through time and space
across local communities, regional-scale assemblages and
global-scale biomes (Kubota, Kusumoto, Shiono, & Tanaka,
2017; Kubota, Kusumoto, Shiono, & Ulrich, 2018a, 2018b;
Kubota, Shiono, & Kusumoto, 2015; Shiono, Kusumoto,
Yasuhara, & Kubota, 2018) and also focus on spatial conser-
vation prioritization to develop protected areas network in
Asia (Kusumoto et al., 2017; Lehtomäki et al., 2018). Our
work motivated us to develop a worldwide research network
(Figure 1) and we run collaborative projects supported by
the Program for Advancing Strategic International Net-
works to Accelerate the Circulation of Talented Researchers,
Japan Society for the Promotion of Science (JSPS) and the
Environment Research and Technology Development Fund
(4–1501 and 4–1802) of the Ministry of the Environment,
Japan. International collaborations promote the exploration
of biodiversity patterns and processes across different spaces,
times and taxa including terrestrial and marine organisms
© 2020 The Ecological Society of Japan 289
290 KUBOTA AND KUSUMOTO

F I G U R E 1 International collaboration network established in the research group for biodiversity and conservation biogeography
(BCB-Japan), University of the Ryukyus (yellow point) by the Program for Advancing Strategic International Networks to Accelerate the
Circulation of Talented Researchers, Japan Society for the Promotion of Science. AU, Aarhus University, Denmark; CSIRO, Commonwealth
Scientific and Industrial Research Organization, Australia; CUNY, City University of New York, USA; FEU, Far Eastern University,
Philippines; HKU, University of Hong Kong; INECOL, Instituto de Ecología A.C., Mexico; JCU, James Cook University, Australia;
NJU, Nanjing University, China; NTHU, National Tsing Hua University, Taiwan; NTU, National Taiwan University, Taiwan; RBG Kew,
Royal Botanic Gardens, Kew, UK; RBG Victoria, Royal Botanic Gardens Victoria, Australia; Smithsonian NMNH, Smithsonian National
Museum of Natural History, USA; UBir, University of Birmingham, UK; UC, University of Canterbury, New Zealand; UH, University of
Helsinki, Finland; UMD, University of Maryland, USA; UMK, Nicolaus Copernicus University in Toru n, Poland; UniNE, University of
Neuchâtel, Switzerland; UOA, University of Auckland, New Zealand; UP, Pedagogical University of Krakow, Poland; UQ, University of
Queensland, Australia; USC, Universidad de Santiago de Compostela, Spain; UvA, University of Amsterdam, the Netherlands

and allow us the opportunity to plan this Special Feature in
Ecological Research.
This Special Feature comprises eight papers that cover
theoretical and empirical topics: on biodiversity estimation
(Chao et al., 2020), large-scale biodiversity patterns
(Kusumoto et al., 2020; Wei, Chen, Wicksten, & Rowe,
2020), ecosystem classifications (Zhao, Basher, & Costello,
2020), species abundance distribution across multiple taxa
(Ulrich, Matthews, & Kubota, 2020), broad-scale species
co-occurrence patterns (Sánchez-Barradas & Villalobos,
2020; Sato, Kusumoto, Çagan, Kubota, & Murakami, 2020)
and potential impacts of human-induced extinction on
island diversity patterns (Matthews, Leidinger, & Cabral,
2020). All these papers focus on emergent patterns (a top-
down approach) and argue the underpinning processes by
means of various statistical approaches.
Estimation of biodiversity is a fundamental step in
understanding the (in)completeness of our knowledge and
precisely capturing the patterns (such as the latitudinal
diversity gradient or regional diversity anomaly), although it
is much more difficult than empirical ecologists had
thought. To quantify modern/paleo-biodiversity from the
incomplete sample, Chao et al. proposed guidelines for
diversity estimation theory, and demonstrated the examples
using species abundance/incidence datasets. Based on that
framework, Kusumoto et al. revisited coral diversity hot-
spots along latitudinal and longitudinal gradients and
demonstrated a “fairly comparable” coral diversity pattern
at a global-scale. They also emphasized priority areas for
future sampling to improve our limited knowledge (such as
Linnean and Wallacean shortfalls). Wei et al. described the
mid-depth diversity maximum of bivalves in the deep-sea
ecosystem and discussed its potential underpinning pro-
cesses. Their collaborative works on biodiversity estimation
illustrated an approach for testing the generality of diversity
patterns previously reported empirically, for example, the
Coral Triangle marine biodiversity hotspot in the Indo-
Pacific region. Zhao et al. revisited marine geographical
units that had been defined subjectively or empirically and
proposed an alternative classification based on objective
analysis with environmental data. Their analysis may pro-
vide fundamental information for designing a network of
marine protected areas. Ulrich et al. focused on species
abundance data sampled across the world. Their phenome-
nological approach using a statistical model revealed con-
straints on species abundance distribution (consistent
realized parameter space) across multiple taxa including
plants, vertebrates and invertebrates and they argued for
universal mechanisms of community assembly. Their study
was an example of the exploration of general laws using a
current large-scale dataset (such as species abundance). In
the context of biogeography, contributions by Sánchez-
Barradas and Villalobos and by Sato et al. focused on species
range overlaps, and tested hypotheses of the mechanisms of
KUBOTA AND KUSUMOTO
spatial co-occurrence. Sánchez-Barradas and Villalobos
argued that climatic Grinnellian niche overlap is more
important than a trophic Eltonian niche in co-occurrence
among American felids. Sato et al. reported that island con-
ditions and species ecological niches jointly play determinis-
tic roles in shaping species co-occurrence of islands birds at
a global scale. From a viewpoint of biodiversity trends in
the Anthropocene, Matthew et al. simulated the impact of
extinction on species–area relationship, functional richness
and beta diversity on islands. They warned that current bio-
geographical patterns had already been modified by
human-induced extinctions, which may bias our interpreta-
tion. These findings and the related arguments should
prompt us to engage in more comprehensive collaboration
across ecology, biogeography, evolution, paleobiology, geol-
ogy and statistical/mathematical theory, to better interpret
the biodiversity patterns we observed at present.
ACK NO WLE DGE MEN TS
We would like to thank all collaborators who contributed
their papers to this Special Feature; the editors-in-chief of
Ecological Research, Prof. Nakaoka and Prof. Suyama;
and other editors for their cooperation. We are also grate-
ful to editorial coordinators, Dr. Shoko Nakamura and
Yuko Aoshima for their kind help and support.
ORCID
Yasuhiro Kubota https://orcid.org/0000-0002-5723-4962
Buntarou Kusumoto https://orcid.org/0000-0002-5091-
3575
R EF E RE N C E S
Chao, A., Kubota, Y., Zeleny, D., Chiu, C.-H., Li, C.-F., Kusumoto, B.,
… Colwell, R. K. (2020). Quantifying sample completeness and
comparing diversities among assemblages. Ecological Research, 35,
292–314 (in this special feature).
Hortal, J., de Bello, F., Diniz-Filho, J. A. F., Lewinsohn, T. M.,
Lobo, J. M., & Ladle, R. J. (2015). Seven shortfalls that beset
large-scale knowledge of biodiversity. Annual Review of Ecology,
Evolution, and Systematics, 46, 523–549.
Hubbell, S. P. (2001). The unified neutral theory of biodiversity and
biogeography. Princeton, NJ: Princeton University Press.
Kubota, Y., Kusumoto, B., Shiono, T., & Tanaka, T. (2017). Phyloge-
netic properties of tertiary relict flora in the east Asian conti-
nental islands: Imprint of climatic niche conservatism and in
situ diversification. Ecography, 40, 436–447.
Kubota, Y., Kusumoto, B., Shiono, T., & Ulrich, W. (2018a). Environ-
mental filters shaping angiosperm tree assembly along climatic and
geographic gradients. Journal of Vegetation Science, 29, 607–618.
Kubota, Y., Kusumoto, B., Shiono, T., & Ulrich, W. (2018b). Multi-
ple filters affect tree species assembly in mid-latitude forest
communities. Oecologia, 187, 245–253.
Kubota, Y., Shiono, T., & Kusumoto, B. (2015). Role of climate and geo-
historical factors in driving plant richness patterns and endemicity
on the east Asian continental islands. Ecography, 38, 639–648.
291
Kusumoto, B., Costello, M., Kubota, Y., Shiono, T., Wei, C.-L.,
Yasuhara, M., & Chao, A. (2020). Global distribution of coral
diversity: Biodiversity knowledge gradients related to spatial res-
olution. Ecological Research, 35, 315–326 (in this special feature).
Kusumoto, B., Shiono, T., Konoshima, M., Yoshimoto, A.,
Tanaka, T., & Kubota, Y. (2017). How well are biodiversity
drivers reflected in protected areas? A representativeness
assessment of geohistorical gradients that shaped endemic flora
in Japan. Ecological Research, 32, 299–311.
Lawton, J. H. (1999). Are there general laws in ecology? Oikos, 84,
177–192.
Lehtomäki, J., Kusumoto, B., Shiono, T., Tanaka, T., Kubota, Y., &
Moilanen, A. (2018). Spatial conservation prioritization for the
east Asian islands: A balanced representation of multitaxon
biogeography in a protected area network. Diversity and Distri-
butions, 25, 414–429.
MacArthur, R. H. (1972). Geographical ecology: Patterns in the distri-
bution of species. Princeton, NJ: Princeton University Press.
Matthews, T. J., Leidinger, L. K. T., & Cabral, J. S. (2020). The effect of
species extinctions on Island biogeographic patterns. Ecological
Research, 35, 372–381 (in this special feature).
McGill, B. J. (2019). The what, how and why of doing
macroecology. Global Ecology and Biogeography, 28, 6–17.
Roughgarden, J. (2009). Is there a general theory of community
ecology? Biology and Philosophy, 24, 521–529.
Sánchez-Barradas, A., & Villalobos, F. (2020). Species geographical
co-occurrence and the effect of Grinnellian and Eltonian niche
partitioning: The case of a Neotropical felid assemblage. Ecolog-
ical Research, 35, 382–393 (in this special feature).
Sato, E., Kusumoto, B., Şekercioglu, Ç. H., Kubota, Y., &
Murakami, M. (2020). The influence of ecological traits and
environmental factors on the co-occurrence patterns of birds
on islands worldwide. Ecological Research, 35, 394–404 (in this
special feature).
Shiono, T., Kusumoto, B., Yasuhara, M., & Kubota, Y. (2018). Roles
of climate niche conservatism and range dynamics in woody
plant diversity patterns through the Cenozoic. Global Ecology
and Biogeography, 27, 865–874.
Simberloff, D. (2004). Community ecology: Is it time to move on?
The American Naturalist, 163, 787–799.
Ulrich, W., Matthews, T. J., & Kubota, Y. (2020). Constraints on the
distribution of species abundances indicate universal mecha-
nisms of community assembly. Ecological Research, 35, 362–371
(in this special feature).
Wei, C.-L., Chen, M., Wicksten, M. K., & Rowe, G. T. (2020).
Macrofauna bivalve diversity from the deep northern Gulf of
Mexico. Ecological Research, 35, 343–361 (in this special feature).
Zhao, Q., Basher, Z., & Costello, M. J. (2020). Mapping near surface
global marine ecosystems through cluster analysis of environmen-
tal data. Ecological Research, 35, 327–342 (in this special feature).
How to cite this article: Kubota Y, Kusumoto B.
Approaches for general rules of biodiversity
patterns in space and time. Ecological Research.
2020;35:289–291. https://doi.org/10.
1111/1440-1703.12101