Tree seedling richness, but not neighborhood composition, influences insect herbivory in a temperate deciduous forest community

Abstract Insect herbivores can serve as important regulators of plant dynamics, but their impacts in temperate forest understories have received minimal attention at local scales. Here, we test several related hypotheses about the influence of plant neighborhood composition on insect leaf damage in southwestern Pennsylvania, USA. Using data on seedlings and adult trees sampled at 36 sites over an approximately 900 ha area, we tested for the effects of total plant density, rarefied species richness (i.e., resource concentration and dietary‐mixing hypotheses), conspecific density (i.e., Janzen–Connell hypothesis), and heterospecific density (i.e., herd‐immunity hypothesis), on the proportion of leaf tissue removed from 290 seedlings of 20 species. We also tested for the effects of generic‐ and familial‐level neighborhoods. Our results showed that the proportion of leaf tissue removed ranged from zero to just under 50% across individuals, but was generally quite low (<2%). Using linear mixed models, we found a significant negative relationship between insect damage and rarefied species richness, but no relationship with neighborhood density or composition. In addition, leaf damage had no significant effect on subsequent seedling growth or survival, likely due to the low levels of damage experienced by most individuals. Our results provide some support for the resource concentration hypothesis, but suggest a limited role for insect herbivores in driving local‐scale seedling dynamics in temperate forest understories.

[1]  David L. Erickson,et al.  Trait similarity, shared ancestry and the structure of neighbourhood interactions in a subtropical wet forest: implications for community assembly. , 2010, Ecology letters.

[2]  J. Connell On the role of the natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees , 1971 .

[3]  D. Bates,et al.  Fitting Linear Mixed-Effects Models Using lme4 , 2014, 1406.5823.

[4]  L. Hughes,et al.  Herbivore damage along a latitudinal gradient: relative impacts of different feeding guilds , 2003 .

[5]  B. Bentz,et al.  Mountain pine beetle , 2009 .

[6]  Frederick H. Utech,et al.  Checklist of the vascular plants of Powdermill Nature Reserve, Westmoreland County, Pennsylvania / , 1999 .

[7]  Michael Smithson,et al.  A better lemon squeezer? Maximum-likelihood regression with beta-distributed dependent variables. , 2006, Psychological methods.

[8]  Marlene H. Dortch Washington, DC , 1985, International Society of Hair Restoration Surgery.

[9]  T. Tscharntke,et al.  Sapling herbivory, invertebrate herbivores and predators across a natural tree diversity gradient in Germany’s largest connected deciduous forest , 2009, Oecologia.

[10]  R. B. Root Organization of a Plant-Arthropod Association in Simple and Diverse Habitats: The Fauna of Collards (Brassica Oleracea) , 1973 .

[11]  W. Kurz,et al.  Mountain pine beetle and forest carbon feedback to climate change , 2008, Nature.

[12]  L. Comita,et al.  Species associations structured by environment and land-use history promote beta-diversity in a temperate forest. , 2015, Ecology.

[13]  D. Janzen Herbivores and the Number of Tree Species in Tropical Forests , 1970, The American Naturalist.

[14]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[15]  P. Hambäck,et al.  Effects of plant neighborhoods on plant-herbivore interactions: resource dilution and associational effects. , 2014, Ecology.

[16]  D. Coomes,et al.  Competitive interactions between forest trees are driven by species' trait hierarchy, not phylogenetic or functional similarity: implications for forest community assembly. , 2012, Ecology letters.

[17]  R. Marquis,et al.  Insectivorous Birds Increase Growth of White Oak through Consumption of Leaf‐Chewing Insects , 1994 .

[18]  Philip E. Hulme,et al.  HERBIVORY, PLANT REGENERATION, AND SPECIES COEXISTENCE , 1996 .

[19]  Thorsten M. Buzug,et al.  IR-04-067 Adaptive Speciation : Introduction , 2004 .

[20]  William J. Foley,et al.  Assessing the evidence for latitudinal gradients in plant defence and herbivory , 2011 .

[21]  A. Oaks,et al.  Influence of Light , 1994 .

[22]  C. Tutin,et al.  Composition of the diet of lowland gorillas at Lopé in Gabon , 1990, American journal of primatology.

[23]  D. Burslem,et al.  Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings , 2009, Proceedings of the Royal Society B: Biological Sciences.

[24]  Owen T Lewis,et al.  Guild-specific Patterns of Species Richness and Host Specialization in Plant–herbivore Food Webs from a Tropical Forest , 2022 .

[25]  W. Härdtle,et al.  Tree diversity promotes insect herbivory in subtropical forests of south-east China , 2010, The Journal of ecology.

[26]  D. J. Shure,et al.  The influence of light and nutrients on foliar phenolics and insect herbivory , 1994 .

[27]  W. L. Baker Eastern forest insects , 1972 .

[28]  B. Bachelot,et al.  Rare species advantage? Richness of damage types due to natural enemies increases with species abundance in a wet tropical forest , 2013 .

[29]  E. Bernays,et al.  Dietary Mixing in a Generalist Herbivore: Tests of Two Hypotheses , 1994 .

[30]  Robert K. Colwell,et al.  Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness , 2001 .

[31]  P. Legendre,et al.  vegan : Community Ecology Package. R package version 1.8-5 , 2007 .

[32]  J. Koricheva,et al.  Effects of tree stand species composition on insect herbivory of silver birch in boreal forests , 2006 .

[33]  Russell Dinnage Phylogenetic diversity of plants alters the effect of species richness on invertebrate herbivory , 2013, PeerJ.

[34]  B. Beckage,et al.  Density-dependent mortality and the latitudinal gradient in species diversity , 2002, Nature.

[35]  Wesley T Beaulieu,et al.  Conspecific Negative Density Dependence and Forest Diversity , 2012, Science.

[36]  S. Hubbell,et al.  Conspecific and phylogenetic density‐dependent survival differs across life stages in a tropical forest , 2015 .

[37]  S. Hubbell,et al.  Strong density- and diversity-related effects help to maintain tree species diversity in a neotropical forest. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[38]  A. Agrawal,et al.  Specialist versus generalist insect herbivores and plant defense. , 2012, Trends in plant science.

[39]  Ws. Rasband ImageJ, U.S. National Institutes of Health, Bethesda, Maryland, USA , 2011 .

[40]  D. Pimentel Species Diversity and Insect Population Outbreaks , 1961 .

[41]  J. Downing,et al.  Biodiversity and stability in grasslands , 1996, Nature.

[42]  D. Burslem,et al.  Herbivory of tropical rain forest tree seedlings correlates with future mortality. , 2010, Ecology.

[43]  J. Chave,et al.  Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest. , 2012, Ecology letters.

[44]  Y. Basset Host specificity of arboreal and free‐living insect herbivores in rain forests , 1992 .