Causes and Consequences of Species Diversity in Forest Ecosystems

Aaron M. Ellison and Frank S. Gilliam
www.mdpi.com/journal/forests
Special Issue Editors
Aaron M. Ellison
Frank S. Gilliam
Special Issue Editors
Aaron M. Ellison
St. Alban-Anlage 66
4052 Basel, Switzerland
This is a reprint of articles from the Special Issue published online in the open access journal Forests
(ISSN 1999-4907) from 2018 to 2019 (available at: https://www.mdpi.com/journal/forests/special
issues/causes consequences diversity)
For citation purposes, cite each article independently as indicated on the article page online and as
indicated below:
LastName, A.A.; LastName, B.B.; LastName, C.C. Article Title. Journal Name Year, Article Number,
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Paramachaerium gruberi Brizicky, an endangered canopy tree of Central American rainforests, shown
here growing on the Osa Peninsula of Costa Rica.
c© 2019 by the authors. Articles in this book are Open Access and distributed under the Creative
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The book as a whole is distributed by MDPI under the terms and conditions of the Creative Commons
license CC BY-NC-ND.
About the Special Issue Editors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Preface to ”Causes and Consequences of Species Diversity in Forest Ecosystems” . . . . . . . ix
Kyle G. Dexter, Ricardo A. Segovia and Andy Griffiths
Exploring the Concept of Lineage Diversity across North American Forests Reprinted from: Forests 2019, 10, 520, doi:10.3390/f10060520 . . . . . . . . . . . . . . . . . . . . . 1
Wenqing Li, Mingming Shi, Yuan Huang, Kaiyun Chen, Hang Sun and Jiahui Chen
Climatic Change Can Influence Species Diversity Patterns and Potential Habitats of Salicaceae Plants in China Reprinted from: Forests 2019, 10, 220, doi:10.3390/f10030220 . . . . . . . . . . . . . . . . . . . . . 19
Ji-Hua Wang, Yan-Fei Cai, Lu Zhang, Chuan-Kun Xu and Shi-Bao Zhang
Species Richness of the Family Ericaceae along an Elevational Gradient in Yunnan, China Reprinted from: Forests 2018, 9, 511, doi:10.3390/f9090511 . . . . . . . . . . . . . . . . . . . . . . . 41
Hong Hai Nguyen, Yousef Erfanifard, Van Dien Pham, Xuan Truong Le, The Doi Bui and Ion
Catalin Petritan
Spatial Association and Diversity of Dominant Tree Species in Tropical Rainforest, Vietnam Reprinted from: Forests 2018, 9, 615, doi:10.3390/f9100615 . . . . . . . . . . . . . . . . . . . . . . . 56
Aaron M. Ellison, Hannah L. Buckley, Bradley S. Case, Dairon Cardenas, Alvaro J. Duque,
James A. Lutz, Jonathan A. Myers, David A. Orwig and Jess K. Zimmerman
Species Diversity Associated with Foundation Species in Temperate and Tropical Forests Reprinted from: Forests 2019, 10, 128, doi:10.3390/f10020128 . . . . . . . . . . . . . . . . . . . . . 69
Aimee Van Tatenhove, Emily Filiberti, T. Scott Sillett, Nicholas Rodenhouse and Michael
Hallworth
Climate-Related Distribution Shifts of Migratory Songbirds and Sciurids in the White Mountain National Forest Reprinted from: Forests 2019, 10, 84, doi:10.3390/f10020084 . . . . . . . . . . . . . . . . . . . . . . 103
Chris J. Peterson
Damage Diversity as a Metric of Structural Complexity after Forest Wind Disturbance Reprinted from: Forests 2019, 10, 85, doi:10.3390/f10020085 . . . . . . . . . . . . . . . . . . . . . . 117
Frank S. Gilliam
Excess Nitrogen in Temperate Forest Ecosystems Decreases Herbaceous Layer Diversity and Shifts Control from Soil to Canopy Structure Reprinted from: Forests 2019, 10, 66, doi:10.3390/f10010066 . . . . . . . . . . . . . . . . . . . . . . 139
R. Travis Belote
Eguale Tadesse, Abdu Abdulkedir, Asia Khamzina, Yowhan Son and Florent Noulekoun
Contrasting Species Diversity and Values in Home Gardens and Traditional Parkland Agroforestry Systems in Ethiopian Sub-Humid Lowlands Reprinted from: Forests 2019, 10, 266, doi:10.3390/f10030266 . . . . . . . . . . . . . . . . . . . . . 164
v
Steffi Heinrichs, Christian Ammer, Martina Mund, Steffen Boch, Sabine Budde, Markus
Fischer, Jorg Muller, Ingo Schoning, Ernst-Detlef Schulze, Wolfgang Schmidt, Martin
Weckesser and Peter Schall
Landscape-Scale Mixtures of Tree Species are More Effective than Stand-Scale Mixtures for Biodiversity of Vascular Plants, Bryophytes and Lichens Reprinted from: Forests 2019, 10, 73, doi:10.3390/f10010073 . . . . . . . . . . . . . . . . . . . . . . 186
Yuanyuan Li, Han Y. H. Chen, Qianyun Song, Jiahui Liao, Ziqian Xu, Shide Huang and
Honghua Ruan
Changes in Soil Arthropod Abundance and Community Structure across a Poplar Plantation Chronosequence in Reclaimed Coastal Saline Soil Reprinted from: Forests 2018, 9, 644, doi:10.3390/f9100644 . . . . . . . . . . . . . . . . . . . . . . . 220
Nicholas W. Bolton and Anthony W. D’Amato
Herbaceous Vegetation Responses to Gap Size within Natural Disturbance-Based Silvicultural Systems in Northeastern Minnesota, USA Reprinted from: Forests 2019, 10, 111, doi:10.3390/f10020111 . . . . . . . . . . . . . . . . . . . . . 233
Callie A. Oldfield and Chris J. Peterson
Woody Species Composition, Diversity, and Recovery Six Years after Wind Disturbance and Salvage Logging of a Southern Appalachian Forest Reprinted from: Forests 2019, 10, 129, doi:10.3390/f10020129 . . . . . . . . . . . . . . . . . . . . . 245
vi
About the Special Issue Editors
Aaron M. Ellison is the Deputy Director of, and a senior ecologist at, the Harvard Forest, the
Senior Research Fellow in Ecology at Harvard University in the Department of Organismic and
Evolutionary Biology, and a semi-professional photographer and writer. He received his B.A. in
East Asian philosophy from Yale University in 1982 and his Ph.D. in evolutionary ecology from
Brown University in 1986. After post-doctoral positions at Cornell and with the Organization for
Tropical Studies in Costa Rica, Dr. Ellison taught for a year at Swarthmore College before moving
to Mount Holyoke College in 1990. There, he was the Marjorie Fisher Assistant, Associate, and Full
Professor, founding director of Mount Holyoke’s Center for Environmental Literacy, and Associate
Dean for Science, and he taught biology, environmental studies, and statistics until 2001. Following
a sabbatical year at Harvard in 2001–2002, Dr. Ellison assumed his current position at the Harvard
Forest—Harvard’s 1500-hectare outdoor classroom and laboratory for ecological research. Dr. Ellison
works in wetlands and forests to study the disintegration and reassembly of ecosystems following
natural and anthropogenic disturbances; the evolutionary ecology of carnivorous plants; the
response of plants and ants to global climate change; the application of Bayesian statistical inference
to ecological research and environmental decision-making; and the critical reaction of ecology to
Modernism. He has authored or co-authored over 200 scientific papers, dozens of book reviews and
software reviews, and the books A Primer of Ecological Statistics (2004; 2nd edition 2012), A Field Guide
to the Ants of New England (2012), Stepping in the Same River Twice: Replication in Biological Research
(2017), Carnivorous Plants: Physiology, Ecology, and Evolution (2018), and Vanishing Point (2017)—a
collection of photographs and poetry from the Pacific Northwest. From 2010–2015, Dr. Ellison was
the Editor-in-Chief of Ecological Monographs, in 2012 he was elected a Fellow of the Ecological Society
of America, and he is currently a Senior Editor of Methods in Ecology & Evolution.
Frank S. Gilliam is a professor of biology at the University of West Florida and a professor emeritus
at Marshall University. He completed his B.S. in biology at Vanderbilt University and received a
Ph.D. in plant ecology at Duke University, studying the fire ecology of southeastern coastal plain pine
forests. Following post-doctoral appointments at Kansas State University to study the fire ecology
of tallgrass prairie and at the University of Virginia to study hardwood forest canopy/atmosphere
interactions, Dr. Gilliam began his 28-year tenure on the faculty of Marshall University in 1990.
His research lies primarily at the conceptual boundary between terrestrial plant communities and
ecosystems, including the movement and cycling of plant nutrients, especially nitrogen (N). These
interests extend to fire ecology and the effects of fire on nutrient cycling, plants, and soils in fire-prone
ecosystems. Additionally, related to his ecosystem approach to ecological research is an interest in
atmospheric deposition and precipitation chemistry, leading to the study of pollutant conditions
(acid deposition, excess N, ozone) in forested areas. Other work includes secondary succession
and the species dynamics of the herbaceous layer of forests, as well as the variety of biotic and
abiotic factors that influence species composition and change within this vegetation stratum. Ongoing
work includes vegetation dynamics in forest ecosystems, N cycling in forest ecosystems, and species
composition and stand structure in longleaf pine forests. Dr. Gilliam currently serves as Associate
Editor for Journal of Ecology and Journal of Plant Ecology. He has authored or co-authored more than 100
peer-reviewed articles, in addition to book chapters and reviews of books, current scientific articles,
and software, and has authored/co-authored the books Terrestrial Plant Ecology, 3rd Edition (1999)
vii
and The Herbaceous Layer in Forests of Eastern North America (2003; 2nd Edition 2014). He is the grateful
husband of Laura P. Gilliam and father of Rachel M. Gilliam, M.Div., and Ian S. Gilliam LTJG USN.
viii
Diversity in Forest Ecosystems”
Forests have the highest plant diversity among terrestrial ecosystems. Among forests, tree
species diversity tends to be highest in the tropics and at low elevations and is positively associated
with increasing precipitation and resource availability. Within forests, trees themselves create
physical structures and habitats for other species. Stratified “layers” consist of species of similar
life forms. The lowest, herbaceous layer is a mixture of resident species (e.g., mosses, liverworts,
ferns, flowering herbs) and transient seedlings of trees and shrubs that eventually grow into higher
strata. Shrub, subcanopy, and canopy layers, in turn, are dominated increasingly by woody shrubs
and trees. Epiphytes, epiphylls, herbaceous vines, and lianas depend on trees for support, while
arthropods, birds, and other animals use them for food and shelter.
Considerable research in recent decades has yielded new insights into the mostly positive
relationships between species diversity and ecosystem processes. There has been a concomitant
escalation of concern that declines in biodiversity of forests caused by increasing human population
size and land-use intensity, together with shifts in biodiversity caused by rapid climatic change
and new disturbance regimes, will compromise the ecosystem “services” forests provide to human
society. Although forests have always been seen as dynamic systems—Henry Chandler Cowles
described ecological succession over a century ago as “a variable approaching a variable, not a
constant”—the rapid increase in atmospheric concentrations of carbon dioxide and other “greenhouse
gases” and frequencies of extreme floods, droughts, fires, and catastrophic cyclonic windstorms
presage extensive changes and rearrangement of forests worldwide. Thus, research on the causes
and consequences of biodiversity in forests now intersects with the anxieties of the Anthropocene.
The 14 papers in this book, reprinted from a 2018–2019 Special Issue of the journal Forests,
illustrate these intersections. The first four papers document patterns of diversity at different
temporal and spatial scales. Dexter et al. place North American forest diversity in a phylogenetic
context and highlight that species diversity is not limited to modern forests but has its roots in
evolutionary processes and deep time. W. Li et al. examine diversity in a single family (Salicaceae) in
China, using species distribution models to explore its climate-driven changes in diversity from the
Last Glacial Maximum (22,000 years ago) through the present and into the late 21st century. Wang et
al. zero in on patterns of diversity of woody Ericaceae in China’s Yunnan Province, while Nguyen
et al. focus on spatial-scale-dependent diversity patterns of trees within individual 2-hectare plots in
north-central Vietnam.
The next four papers investigate drivers of diversity in unmanaged forests. Ellison et al. use data
from large (≥15-hectare) forest plots in the Western Hemisphere to look for statistical “fingerprints”
of foundation tree species—those species that control the biodiversity and ecosystem dynamics of the
forests they define and structure. Van Tatenhove et al. document the complex interaction of climatic
factors on changing elevational distributions of forest birds and small mammals in New Hampshire’s
White Mountains. Peterson develops a new index of “damage diversity” and shows how it is related
to climatic drivers (wind disturbance) and in turn, influences structure diversity and complexity in
the forests of eastern North America. Gilliam reviews experimental studies of a quarter-century of
nitrogen addition on an experimental forest in West Virginia. This work has revealed that nitrogen
addition at levels similar to those coming from atmospheric deposition leads to a decline in species
diversity of herbaceous species in the forest understory and a greater sensitivity of the remaining
ix
species to changes in light availability defined by the woody overstory.
The final six papers place forest diversity squarely in the context of human impacts and
management. Belote confirms the expected pattern that water availability and soil fertility positively
affect species diversity and productivity in the United States. He goes on to show—perhaps
unsurprisingly but rarely documented—that people are more likely to manage and modify highly
productive, species rich forests but conserve forests that have fewer species and lower productivity.
In a curious parallel, Tadesse et al. find that home gardens in western Ethiopia have a nearly
threefold higher tree diversity than nearby “natural” parklands. People manage for productivity and
diversity. They cultivate many non-native tree species that provide food, fiber, and lumber, whereas
parklands have primarily native species that provide similar, albeit less-productive, ecosystem
services. Heinrichs et al. illustrate that managing forests in Germany as mixtures rather than
monocultures increases local- and landscape-scale diversity of vascular plants, bryophytes, and
lichens. Y. Li et al. examine diversity of soil arthropods in monoculture plantations of poplar (Populus
deltoides) in eastern China, finding generally higher species richness in older (21-year-old) stands
but also temporal shifts in species composition. Bolton and D’Amato show that managing with
disturbance (harvest gaps) increases diversity of both native and non-native understory plants in
silvicultural systems in Minnesota (USA). Lastly, Oldfield and Peterson draw the useful distinction
between diversity and species composition in their report that salvage logging following wind
disturbance has little effect on diversity (as number of species) but substantial effects on species
composition in forests of north Georgia (USA).
Taken together, the papers in this book cover a broad range of forest types across four continents
and examine a wide range of topics relevant to understanding the causes and consequences of forest
diversity. They also illustrate the central importance on this human-dominated planet of managing
for, and with, species diversity in forests.
Aaron M. Ellison, Frank S. Gilliam
Special Issue Editors
Exploring the Concept of Lineage Diversity across North American Forests
Kyle G. Dexter 1,2,*, Ricardo A. Segovia 1,3 and Andy R. Griffiths 1
1 School of GeoSciences, University of Edinburgh, Edinburgh EH9 3FF, UK; [email protected] (R.A.S.); [email protected] (A.R.G.)
2 Royal Botanic Garden Edinburgh, Edinburgh EH3 5LR, UK 3 Instituto de Ecología y Biodiversidad, Santiago 7800003, Chile * Correspondence: [email protected]; Tel.: +44-(0)-131-650-7439
Received: 30 May 2019; Accepted: 16 June 2019; Published: 22 June 2019
Abstract: Lineage diversity can refer to the number of genetic lineages within species or to the number of deeper evolutionary lineages, such as genera or families, within a community or assemblage of species. Here, we study the latter, which we refer to as assemblage lineage diversity (ALD), focusing in particular on its richness dimension. ALD is of interest to ecologists, evolutionary biologists, biogeographers, and those setting conservation priorities, but despite its relevance, it is not clear how to best quantify it. With North American tree assemblages as an example, we explore and compare different metrics that can quantify ALD. We show that both taxonomic measures (e.g., family richness) and Faith’s phylogenetic diversity (PD) are strongly correlated with the number of lineages in recent evolutionary time, but have weaker correlations with the number of lineages deeper in the evolutionary history of an assemblage. We develop a new metric, time integrated lineage diversity (TILD), which serves as a useful complement to PD, by giving equal weight to old and recent lineage diversity. In mapping different ALD metrics across the contiguous United States, both PD and TILD reveal high ALD across large areas of the eastern United States, but TILD gives greater value to the southeast Coastal Plain, southern Rocky Mountains and Pacific Northwest, while PD gives relatively greater value to the southern Appalachians and Midwest. Our results demonstrate the value of using multiple metrics to quantify ALD, in order to highlight areas of both recent and older evolutionary diversity.
Keywords: temperate forests; species richness; assemblage lineage diversity; phylogenetic diversity; evolutionary diversity; United States; trees; TILD
1. Introduction
The evolutionary lineage is a fundamental concept in biology, denoting a group of organisms connected by ancestor-descendent relationships [1]. Evolutionary lineages are hierarchically structured; multiple younger evolutionary lineages can be nested within an overarching older lineage, or clade. Thus, multiple genetically diverged lineages can exist within a single taxonomic species, and multiple species can belong to older evolutionary lineages, such as genera, families or orders. Knowing the number of lineages in different ecological assemblages and biogeographic regions gives insights into evolutionary process, biogeographic history, and conservation priorities. For example, an assemblage or region that houses many lineages can be interpreted as having a richer evolutionary history, and therefore may be a greater priority for conservation than one that houses few. However, the conservation value of lineage diversity has yet to be fully, and persuasively, communicated [2–4]. Providing clear and accurate quantification of lineage diversity may assist its integration into conservation practice.
Forests 2019, 10, 520; doi:10.3390/f10060520 www.mdpi.com/journal/forests1
Forests 2019, 10, 520
In its most basic form, quantifying the number of lineages in assemblages could consist of counting the number of species. However, the term lineage diversity is generally applied when the units are not species, but a shallower or deeper evolutionary level, i.e., within or above the species taxonomic rank (see [5–9] for examples below species rank; see [10–14] for examples above species rank). In this paper, we focus on lineage diversity above the species rank. Employing tree assemblages in the contiguous United States, we explore various metrics by which assemblage lineage diversity (hereafter ALD) might be quantified, using taxonomic and phylogenetic approaches. Given its pertinence to conservation prioritisation, we focus specifically on the richness dimension of ALD.
Taxonomy is a hierarchical system for organising biological diversity. As such, it provides an apparently straightforward means of quantifying ALD at different evolutionary depths, for example by tallying the number of genera, families or orders in assemblages. However, Linnean taxonomic ranks are not ‘natural’ in the sense that they do not directly correlate to any precise evolutionary age. Some clades of a given taxonomic rank may actually be younger than clades of a putatively lower taxonomic rank. For example, the genus Pinus (Pinaceae) may be as old as 100 million years [15], which is older than most angiosperm families [16]. If one were to compare an assemblage of four Pinus species with an assemblage of four angiosperm species belonging to different…