Tree diversity effects on leaf insect

damage on pedunculate oak_ The role

of landscape context and forest stratum
Bastien Castagneyrol & Brice Giffard &
Elena Valdés-Correcher & Arndt Hampe
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 Forest Ecology and Management 433 (2019) 287–294

Contents lists available at ScienceDirect

Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Tree diversity effects on leaf insect damage on pedunculate oak: The role of T
landscape context and forest stratum
⁎,1
Bastien Castagneyrola,1, Brice Giffardb, , Elena Valdés-Correchera, Arndt Hampea
a
BIOGECO, INRA, Univ. Bordeaux, 33610 Cestas, France
b
UMR 1065 Santé et Agroécologie du Vignoble, Bordeaux Sciences Agro, INRA, ISVV, 33175 Gradignan, France

A R T I C LE I N FO A B S T R A C T

Keywords: Tree diversity has long been recognized as a major driver of insect herbivory in forest ecosystems. However,
Biodiversity predicting the strength and direction of tree diversity effects in real-world situations has proven elusive. One
Forest likely reason is that most studies have focused on within-stand dynamics and insufficiently captured other
Leaf traits ecological drivers of insect herbivory that can act at broader (i.e., landscape) and finer (i.e., individual trees)
Leaf insect herbivory
scales. We measured herbivory as leaf area consumed by insect herbivores in pedunculate oaks (Quercus robur)
Forest stratification
Host plant concentration
growing in mixed and pure forest stands in southwestern France. We assessed the effects of oak spatial isolation
Quercus robur within the landscape, tree stand diversity, forest canopy stratification as well as the influence of leaf traits on
insect herbivory. Insect herbivory increased with stand isolation regardless of tree diversity. Diversity effects
were contingent upon the canopy stratum as insect herbivory in mixed stands exceeded that of pure stands only
in the upper stratum. Leaf traits varied between pure and mixed stands and among canopy strata. Insect her-
bivory was negatively correlated with LDMC and positively with SLA. However, the observed effects of tree
diversity, canopy stratum and stand isolation on insect herbivory were only partially driven by variability in oak
leaf traits. Our findings illustrate that, in real-world contexts, insect herbivory can be driven by a complex
interplay of multiple, scale-dependent drivers. They help step forward towards a more profound understanding
of the complex forces drive insect herbivory in managed forest ecosystems.

1. Introduction background insect herbivory under natural, non-outbreak conditions

A long held view in forest ecology is that tree diversity strongly
influences insect herbivory. Extensive research has demonstrated that
trees are generally more prone to suffering damage when grown in
monospecific stands than when associated with other tree species
(Castagneyrol et al., 2014; Vehviläinen et al., 2007), although neutral
(Rosado-Sánchez et al., 2018) or even opposite patterns have also been
reported (Schuldt et al., 2010). The underlying phenomenon, termed
associational resistance, appears to be widespread in forests
(Castagneyrol et al., 2014; but see Haase et al., 2015; Schuldt et al.,
2015). However, to date most empirical evidence on associational ef-
fects in forests stems from case studies of outbreaks of particular pest
species and from highly controlled experiments (Castagneyrol et al.,
2013; Damien et al., 2016; Schuldt et al., 2015; Vehviläinen et al.,
2007). Despite their evident strengths (Grossman et al., 2018; Paquette
et al., 2018; Verheyen et al., 2016), such experiments fail to properly
consider the diversity and complexity of drivers that tend to affect
(Guyot et al., 2016; Moreira et al., 2017; Kozlov and Zvereva 2017).
This lack strongly constrains our understanding of the actual ecological
relevance of associational effects on insect herbivory in real-world si-
tuations.
Patterns of insect herbivory are moulded by a variety of factors
controlling plant accessibility at different scales. At the landscape scale,
herbivore density tends to be highest in those habitat patches where
their resource is most abundant, because the intensity of physical and
chemical cues makes these patches more likely to be found and colo-
nized (Andersson et al., 2013; Hambäck and Englund, 2005; Root,
1973). Within patches, herbivory on individual plants is influenced by
the identity and diversity of their neighbours which alter the focal
plant’s physical and chemical apparency and its colonization
(Castagneyrol et al., 2013; Finch and Collier, 2000; Moreira et al.,
2016). Finally, herbivory is controlled by individual plant traits in-
cluding nutritional quality and anti-herbivore defences (Castagneyrol
et al., 2018b; Finch and Collier, 2000; Schoonhoven, 2005). Insects rely

⁎
Corresponding author.
E-mail address: brice.giffard@gmail.com (B. Giffard).
1
BC and BG share first authorship.

https://doi.org/10.1016/j.foreco.2018.11.014
Received 3 July 2018; Received in revised form 9 November 2018; Accepted 10 November 2018
0378-1127/ © 2018 Elsevier B.V. All rights reserved.
B. Castagneyrol et al. Forest Ecology and Management 433 (2019) 287–294

on a complex system of decision cues for selecting the plants they forage Table 1
on. These can be used hierarchically or sequentially, and their role can Summary of stand characteristics.
vary depending on the animal’s spatial scale of perception or nutritional Stand Coordinates Stand type
status (Andersson et al., 2013; Schoonhoven, 2005). For instance,
herbivores can be attracted by large patches from the distance but their Berganton 44° 45′40.85″N, 0° 49′ 37.58″W Pure
France 44° 44′ 44.10″N, 0° 50′ 50.82″W Pure
final decision to feed on a given plant depends on its traits and its
St Alban 44° 43′ 18.78″N, 0° 45′ 3.25″W Pure
neighbours (Finch and Collier, 2000, 2012; Hambäck et al., 2014). Barlan 44° 44′ 57.00″N, 0° 49′ 53.12″W Pure
While great advances have been made in our understanding of specific Castéra 44° 44′ 0.1″N, 0° 52′ 42.29″W Pure
drivers of insect herbivory, the scale-dependent interplay between dif- Croix d′Hins 44° 43′ 21.69″N, 0° 49′ 32.31″W Pure
ferent drivers remains poorly understood. Hermitage 44° 44′ 50.69″N, 0° 46′ 10.78″W Mixed
Renardière 44° 43′ 42.33″N, 0° 50′ 8.78″W Mixed
Some such interactions have recently been hypothesized. It has for
H5 44° 43′ 8.11″N, 0° 49′ 59.83″W Mixed
instance been suggested that resource isolation at the landscape level H6 44° 43′ 30.63″N, 0° 51′ 10.72″W Mixed
may cause herbivores to dedicate more time to foraging within resource H8 44° 43′ 10.51″N, 0° 50′ 36.85″W Mixed
patches, thus increasing damage in isolated stands as a result of re- H20 44° 42′ 56.23″N, 0° 51′ 48.50″W Mixed
inforced small-scale effects of tree diversity on feeding decisions
(Hambäck et al., 2014; Stutz et al., 2015; Underwood et al., 2014).
However, observational studies demonstrating such an interaction are agricultural lands and woodlands whose most common tree species are
lacking. Within habitat patches, herbivory tends to vary along vertical oaks (Quercus robur, Q. pyrenaica) and birch (Betula pendula). These
gradients in forest canopy as a result of parallel changes in micro- deciduous tree species are also present in variable abundance in the
climate (in particular, higher temperatures in upper strata, Stiegel et al., pine plantations.
2017), leaf traits (Dudt and Shure, 1994; Stiegel et al., 2017; Thomas In early 2009, we selected 12 forest stands for study: six pure stands
et al., 2010), or the diversity and activity of herbivores’ enemies in which Quercus robur was the main species, and six mixed stands
(Aikens et al., 2013). For instance, Stiegel et al. (2017) showed that the consisting of pine-oak mixtures where oaks were abundant (Table 1).
decrease in insect herbivory from lower to upper stratum was ac- For each stand, we mapped the main habitat types in circular buffers of
counted for by the parallel increase of temperatures and decrease in 500 m radius (ca. 78.5 ha). We distinguished pine plantations, decid-
nitrogen content of more sun-exposed leaves. However, the effect of uous forests and open habitats. Open habitats included roads, forest
stratification on herbivory has proven to vary among insect feeding tracks, firebreaks, clearcuts, field and field margins, and young pine
guilds and forest management (Gossner et al., 2014). Tree diversity is plantations (Barbaro et al., 2005). The buffer of 500 m radius provided
known to alter the vertical stratification of forest canopies (Forrester, the largest gradient of habitat variability, avoided spatial overlapping
2017; Vanhellemont et al., 2018). In turn, forest stratification will likely between nearby buffers, and was previously found to be suitable to
alter the effects of tree diversity on herbivores. For instance, both the study plant-herbivore-predator interactions in different landscape
position in the canopy (Stiegel et al., 2017) as well as the density and contexts (Barbaro et al., 2005; Chaplin-Kramer et al., 2011). Habitat
diversity of neighbouring trees can trigger the surface or the C:N ratio mapping was based on satellite images read and analysed with QGIS
of plant leaves, two traits that are commonly related with patterns of version 2.18.13 (QGIS Development Team, 2017).
herbivory (Castagneyrol et al., 2017; Moreira et al., 2017; Loranger
et al., 2013; Pearse, 2011; Schoonhoven, 2005). However, quantifying 2.2. Leaf sampling and measurements
the relative contribution of trait-dependent and trait-independent ef-
fects of plant density and diversity on insect herbivory remains chal- In each stand, we randomly selected individual oak trees and as-
lenging and the few existing studies have yielded conflicting results. signed them to one of three forest strata. All leaves collected below 1 m
Here, we address how tree diversity effects on leaf damage caused were assigned to the lower stratum, which may have included hanging
by defoliating insect herbivores in pedunculate oak (Quercus robur) are low branches of adult trees as well as leaves of younger individuals. The
shaped by ecological drivers independently and interactively acting at canopies were further divided in an intermediate stratum (lower and
the individual (i.e., leaf traits), habitat (i.e., forest stratum) and land- mid-height branches) and an upper stratum (upper branches). We chose
scape (i.e., forest stand isolation) scales. For this purpose, we measured to define these strata in relative instead of absolute terms because tree
leaf herbivory in oaks in the lower, intermediate and upper stratum of height varied among the different stands.
pure and mixed oak forest stands along a gradient of forest isolation at In 2009, we sampled twice a total of 300 oak leaves per stratum in
the landscape level across the season. We hypothesised that (i) insect each stand, in early (July) and late (September) season. The two sam-
herbivory would be lower in mixed oak-pine stands than in pure oak pling dates were considered as temporal replicates. Although leaf her-
stands, and (ii) the difference would be strongest in isolated stands. We bivory is a cumulative process such that early season damage remains
further predicted that (iii) leaf traits and insect herbivory would vary visible in late season, oaks produce three and up to four generations of
among forest strata and that (v) differences in herbivory among forest leaves in the study area. Thus, the first and second campaigns represent
strata would be larger in pure than in mixed stands. By addressing tree independent assessments of leaf insect herbivory.
diversity effects on insect herbivory at different scales, both within and Leaves were collected on a sample of six individuals per stratum and
between stands, our study pursues a better understanding of the hier- per season, that were haphazardly selected each time to ensure statis-
archical mechanisms that drive tree-herbivore interactions in real- tical independence of the temporal replicates. The number of sampled
world landscapes. leaves was held constant between pure and mixed stands and between
temporal replicates, whereas the sampling methodology was adapted to
2. Materials and methods the forest stratum because of technical constraints. Leaves were col-
lected using a pruning shear for the lower stratum, and with a 10 m pole
2.1. Study site and stand selection pruner for the intermediate stratum of all stands and the upper stratum
of mixed stands. Rifle shooting was used to cut branches down in the
The study was carried out in the Landes de Gascogne region (south- upper stratum of pure stands. For the lower stratum, we assembled a
western France), about 40 km southwest of Bordeaux (44°41′N, 300 leaf sample by haphazardly collecting 35 leaves on one hanging
00°51′W). This region harbours the largest plantation forest in Europe branches of six different individuals, plus an additional pool of leaves
with a monoculture of maritime pine (Pinus pinaster) covering ap- from younger individuals. For intermediate and upper strata, we as-
proximately 10 000 km2. The remaining landscape is characterized by sembled the 300 leaf sample by collecting 30–50 leaves on six to ten

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Table 2
Summary of LMM testing the effects of season, forest type, stratum and isolation on herbivory and leaf traits. P-values are indicated within brackets and significant
effects are shown in bold. Marginal (R2m) and conditional (R2c) R2 are reported for the simplified model.
Predictors Total herbivory Leaf surface SLA LDMC
df χ2-value χ2-value χ2-value χ2-value

Season 1 121.06 (< 0.001) 0.66 (0.416) 4.01 (0.045) 122.03 (< 0.001)
Forest type 1 3.2 (0.072) 4.74 (0.029) 11.08 (< 0.001) 13.55 (< 0.001)
Stratum 2 129.79 (< 0.001) 1090.22 (< 0.001) 1240.75 (< 0.001) 102.15 (< 0.001)
Isolation 1 8.27 (0.004) 3.95 (0.047) 1.00 (0.318) 0.05 (0.819)
Forest type × Stratum 2 71.71 (< 0.001) 26.69 (< 0.001) 68.70 (< 0.001) 15.45 (< 0.001)
Forest type × Isolation 1 0.86 (0.352) 0.02 (0.885) 1.59 (0.207) 2.37 (0.123)
Stratum × Isolation 2 1.08 (0.583) 2.89 (0.216) 1.26 (0.533) 2.58 (0.275)
Forest type × Stratum × Isolation 2 0.67 (0.716) 2.21 (0.331) 0.59 (0.744) 0.32 (0.854)
R2m (R2c) 0.36 (0.45) 0.65 (0.66) 0.68 (0.72) 0.36 (0.45)

Fig. 1. Effects of stand isolation, forest type and stratum on leaf herbivory. (A) Effects of stand isolation. Each point represents leaf herbivory averaged ( ± SE) per
stratum and season. (B) Interactive effects of forest type and stratum on leaf herbivory. Each point represents leaf herbivory averaged per season across all forest
stands ( ± SE, n = 6 per forest type). Letters above bars indicate statistical differences between forest types.

branches of 6 different trees (1 or 2 branches per tree). In each sample,
we took every tenth leaf until reaching the scheduled sample size.
Herbivory was estimated by two measurers aware of sample origin.
To reduce variability among observers, we used a grid of 0.25 cm2
(0.5 × 0.5 cm) printed on a transparent plastic sheet and overlaid on
leaves. We calculated the total leaf area removed or affected by insect
herbivores divided by the number of leaves analysed. We initially dis-
entangled damage caused by different feeding guilds (chewers, skele-
tonizers, leaf-rollers, leaf-miners). However, some of these guilds
caused too scant damages to allow separate analyses and we therefore
pooled all types of damages.
We measured three leaf traits: leaf surface, Specific Leaf Area (SLA)
and Leaf Dry Matter Content (LDMC). These traits were measured on 10
leaves per stratum, stand and sampling date following Cornelissen et al.
(2003) after confirming that this sample size satisfyingly captures
variability among strata. We only used undamaged, mature, and fully
expanded leaves. Leaf surface and water-saturated fresh mass were
measured with a planimeter (WinFolia Pro 2007b, Regent Instruments,
Canada Inc.) and a balance (Ohaus EP114 Explorer Pro Analytical
Balance). Leaf surface, SLA and LDMC were first calculated at the level
of individual leaves and then averaged per replicate, stratum, stand and
season.
2.3. Statistical analyses
We calculated the percentage of open area in buffers of 500 m radius
centered on selected stands as a proxy for stand isolation at the land-
scape level. We preferred to use this variable instead of deciduous forest
cover because pedunculate oaks commonly grow below the canopy of
pine plantations without being detectable on satellite images (Gerzabek
et al., 2017). The cover of deciduous forest alone therefore under-
estimates oak abundance in the landscape. We used linear mixed-effect
models (LMM) to analyse the effects of landscape, tree diversity and
canopy stratification on leaf traits and insect herbivory. Fixed effects
were season (early vs. late), forest type (pure vs. mixed stand), stratum
(lower, intermediate and upper) and isolation (percentage of open areas
in the landscape). Season was considered as a temporal replicate. We
tested all two- and three-way interactions between forest type, stratum
and isolation. We declared stand identity as a random factor to account
for the non-independence of samples from the same stand. For each
response variable (herbivory, leaf surface, SLA and LDMC), we first
built the full model and then applied model simplification by sequen-
tially removing non-significant terms, starting with the highest-order
interaction term. We made no attempt to simplify the random factor as
it was imposed by the sampling design. Significant interactions between
stratum and forest type were treated by estimating contrasts among
strata for each forest type separately and contrasts between pure and
mixed stands for each stratum independently. Finally, we estimated

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B. Castagneyrol et al.
Fig. 2. Interactive effects of stratum and forest type on leaf traits. Dots and
error bars represent means ( ± SE) across seasons and forest types. Letters
above bars indicate statistical differences between strata. Contrasts between
strata are shown for each forest type separately (indicated by different grey
shades).
model coefficients of the simplified model and calculated R2 for fixed
effects (Rm2) and fixed plus random effects (Rc2). Response variables
were log-transformed to improve the distribution of model residuals.
We back-transformed model predictions with an exponential function
to plot them on figures.
We were interested in disentangling the effects of isolation, forest
type, stratum and leaf traits on herbivory. Yet, leaf traits were influ-
enced by both forest type and stratum (see Results). In order to avoid
issues arising from collinearity, we also tested the effect of leaf traits on
herbivory in a separate model where we replaced the factors forest type
and stratum by SLA and LDMC. We then applied the same modelling
procedure as described above. SLA, LDMC and isolation were scaled
and centered to allow comparing coefficient parameter estimates, al-
though we present raw data in the figures.
Finally, we used a structural equation modelling (SEM) to confirm
the indirect trait-mediated effect of tree diversity and forest stratum on
herbivory. We first built a theoretical model in which herbivory was
only explained by SLA and LDMC. SLA and LDMC were endogeneous
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Forest Ecology and Management 433 (2019) 287–294
Fig. 3. Interactive effect of Specific Leaf Area (SLA) and stand isolation on leaf
herbivory. Grey shades and isolines show predictions from mixed-effects
models for early-season data and for an average value of LDMC. Dots show the
original data.
variables. Tree diversity and forest stratum were exogeneous variables
that only influenced SLA and LDMC. We used the piecewiseSEM
package (Lefcheck, 2016) and Shipley’s test of direct separation to
evaluate the probability that none of the paths missing from the hy-
pothesised network contain useful information (in particular direct
paths linking forest stratum and tree diversity to herbivory). This hy-
pothesis was considered rejected if χ2-test of Fisher’s C statistic fell
below the significance level (P < 0.05).
All analyses were done in R version 3.3.1 (R Development Core
Team, 2016) using the packages lmer4, car, multcomp, MuMIn and pie-
cewiseSEM (Bartoń, 2016; Fox and Weisberg 2011; Hothorn et al., 2008;
Kuznetsova et al., 2016, Lefcheck, 2016).
3. Results
3.1. Effects of forest type, stratum, isolation and season on herbivory
Herbivory was on average ( ± SE) 1.42 ± 0.03 cm2 per leaf (cor-
responding to ca. 8% leaf area), which corresponds to background in-
sect herbivory in the study area. Herbivory was 37% higher in the late
season than in the early season (Table 2). It increased with stand iso-
lation (Fig. 1A) and decreased from the lower to the upper stratum
(Fig. 1B). Herbivory did not differ between mixed and pure stands, but
we observed a significant forest type × stratum interaction (Table 2) as
differences between pure and mixed stands were only significant in the
upper stratum (coefficient parameter estimate ± SE: 0.60 ± 0.13,
Fig. 1B). In pure stands, leaf herbivory was significantly lower in the
upper stratum (Fig. 1B) whereas it did not differ between the inter-
mediate and the lower stratum. In mixed stand, leaf herbivory was
higher in the lower stratum than in the intermediate and upper stratum,
whereas the latter two did not differ (Fig. 1B).
3.2. Effects of forest type, stratum and isolation on leaf traits
Leaf traits varied consistently among strata and between forest types
B. Castagneyrol et al.
Upper: -2.73 (***)
Intermediate : -0.24 (***)
Upper: -0.98 (***)
Intermediate : -1.01 (***)
0.11 (*)
Stratum SLA
Upper: 0.18 (***) Mixture :
Intermediate : 0.38 (***) - 0.29 (**)
Tree
LDMC
diversity Mixture: - 0.34 (***)
0.54(**)
(Fig. 2) with a significant forest type × stratum interaction for all traits
(Table 2). Leaf surface was on average ( ± SE) 17.5 ± 0.3 cm2. It was
twice as large in the intermediate and upper stratum than in the lower
stratum, both across forest types and seasons, while there were no
differences between the intermediate and the upper stratum. Differ-
ences between the lower and the intermediate and upper stratum were
larger in pure stands than in mixed stands (Table 2). Oaks had larger
leaves in pure stands than in mixed stands, but only in the upper and
intermediate strata. SLA was on average 15.40 ± 0.15 mm2·mg−1. It
consistently decreased from lower to upper stratum in both forest types
and seasons. All contrasts between strata were significant, but differ-
ences were larger in pure than in mixed stands. SLA tended to be higher
in pure stands than in mixed stands, but this difference was only sig-
nificant in the lower and the intermediate stratum. LDMC was on
average 436.0 ± 1.2 mg·g−1. It consistently increased from lower to
upper stratum in both seasons and forest types. However, differences
among strata were contingent on forest type (Table 2). In pure stands,
LDMC was greater in the upper stratum than in the intermediate and
lower stratum, while the latter two did not differ. In mixed stands, all
contrasts between strata were significant. LDMC was greater in mixed
stands than in pure stands, but this difference was only significant in
the intermediate and the upper stratum.
4. Leaf traits associated with the effects of forest type and stratum
on herbivory
Replacing the factors forest type and stratum in the LMM by the leaf
traits SLA and LDMC, we found that differences in herbivory between
forest types and among strata could be accounted for by LDMC and SLA.
Herbivory decreased with LDMC ([82.4 ± 24.8] × 10−3, df = 1,
χ2 = 11.00, P = 0.001) and increased with SLA ([6.0 ± 37.6] × 10−3,
df = 1, χ2 = 34.90, P < 0.001, Fig. 3). The effect of SLA was however
contingent on stand isolation (significant SLA × isolation interaction:
df = 1, χ2 = 14.46, P < 0.001). The positive coefficient parameter
estimate ( ± SE) for the interaction (0.06 ± 0.02) indicated that the
effect of SLA on herbivory was slightly stronger in more isolated stands
and that the effect of stand isolation was in turn stronger for leaves with
greater SLA. These results are consistent with our observation that SLA
and herbivory both decreased from the lower to the upper stratum.
Model R2 was however lower when the factors forest type and stratum
were replaced by SLA and LDMC (R2m = 0.29 and R2c = 0.42 vs.
R2m = 0.36 and R2c = 0.44, Table 2), suggesting that the measured leaf
traits account largely but not completely for differences in herbivory
arising from differences between forest types and among strata.
In line with these results, the SEM analysis (Fig. 4) confirmed that
there were missing paths in our a priori network (C = 9.69, df = 4,
P = 0.046), indicating that the effects of tree diversity and forest
stratum on insect herbivory were only partially mediated by their effect
on the measured leaf traits. In particular, there was a significant missing
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Forest Ecology and Management 433 (2019) 287–294
Fig. 4. Path diagrams showing the results of the
piecewise SEM. Solid and dashed lines represent
significant direct and indirect relationships
among variables, respectively. Black and grey
lines represent negative and positive correla-
tions, respectively. Standardized coefficients and
significance thresholds are shown along paths.
Herbivory
path between forest stratum and herbivory, indicating an additional,
trait-independent, effect of forest stratum on insect damage.
5. Discussion
The factorial and hierarchical sampling design of this single year-
study enabled us to get a detailed insight into the complex interplay of
environmental drivers that determine patterns of background insect
herbivory across spatial scales (Fig. 5). We did not detect a global dif-
ference in herbivory between mixed and pure stands, yet a finer ana-
lysis revealed that such an effect of tree diversity did exist but only in
certain canopy strata in pure stands. We also observed that both stand
isolation and stratum influenced herbivory. Finally, both stand isolation
and stratification effects were partly explained by the concomitant
variation in the leaf traits SLA and LDMC. This set of interrelationships
clearly illustrates the complex nature of the multiple, scale-dependent
drivers of insect herbivory in real-world contexts. It calls for caution
when interpreting ecological studies that address limited sets of puta-
tive drivers of insect herbivory in simplified environments.
Leaf insect herbivory was not globally reduced in mixed stands
but in certain strata. We found that tree diversity effects on insect
herbivory were not consistent across canopy strata. While mixed stands
experienced lower herbivory than pure stands in the lower and inter-
mediate stratum, the opposite occurred in the upper stratum. Previous
studies on effects of tree diversity on insect herbivory have acknowl-
edged potential stratum effects but, instead of quantifying them, seeked
to reduce them by averaging herbivory at the level of individual trees
(Castagneyrol et al., 2013; Muiruri et al., 2015; Vehviläinen et al.,
2006). Our study pinpoints the pitfalls of such an approach by de-
monstrating that the canopy stratum can exert a significant influence on
the (non-)detection of global effects of tree diversity. The pattern that
we observed might be explained by the fact that insects coming from
other forest stands are likely to arrive in the upper stratum. Reduced
herbivory in the upper stratum of pure stands could then reflect a di-
lution of the recently arrived herbivores among a larger number of host
trees (Bañuelos and Kollmann, 2011; Damien et al., 2016; Otway et al.,
2005).
Leaf insect herbivory increased from the upper to the lower
stratum. In accordance with previous studies (e.g. Reynolds and
Crossley, 1997; Stiegel et al., 2017), we found that insect herbivores
caused most damage in the lower stratum. This trend has several pos-
sible, non-exclusive explanations. First, vertical stratification in her-
bivory can be a direct consequence of stratification in leaf traits. SLA
increased and LDMC decreased towards the lower stratum, suggesting
that its leaves were most palatable and hence most attractive for her-
bivores (Le Corff and Marquis, 1999; Murakami and Wada, 1997;
Stiegel et al., 2017). This assumption was fully confirmed by the dis-
tinct effects that both parameters exerted on levels of herbivory. Al-
though we did not measure microclimate in our stands, it is likely that
B. Castagneyrol et al.
Fig. 5. Summary of herbivory and leaf trait response to forest type and strati-
fication. Grey squares represent herbivory and traits for different forest types
and strata. Differences in square size are proportional to observed changes in
herbivory or traits. The figure therefore illustrates the extent of differences
between forest types and among strata.
light and temperature were the primary drivers of the observed SLA and
LDMC trends (Yamasaki and Kikuzawa, 2003, Stiegel et al., 2017). The
elevated herbivory in the lower stratum might also be triggered to some
extent by reduced levels of chemical defences in the youngest in-
dividuals (Boege and Marquis, 2005; Moreira et al., 2017). Yet this
effect does not explain the observed differences between the inter-
mediate and the upper stratum. Second, stratification of herbivory
could result from stratification of herbivore communities. Several stu-
dies have reported higher abundance or richness of herbivores in lower
canopy strata and linked this trend with greater leaf quality (Le Corff
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Forest Ecology and Management 433 (2019) 287–294
and Marquis, 1999; Murakami et al., 2005) or a vertical transfer of
herbivores from the canopy to the understory (Murakami and Wada,
1997; White and Whitham, 2000). However, relationships between
herbivore abundance or diversity and herbivore damage are not
straightforward (Basset et al., 1992; Rhainds and English-Loeb, 2003;
Rossetti et al., 2017); hence the relevance of this explanation is difficult
to gauge in our case. Third, top-down control of herbivores by their
enemies could generate lower herbivory in upper strata, if predation
pressure is higher in this part of the canopy (Sobek et al., 2009; Aikens
et al., 2013). Unfortunately, we possess no empirical data yet on pre-
dator abundance or activity that would enable us to test the validity of
this explanation.
Effects of forest type and stratum on leaf insect herbivory were
partly explained by leaf traits. On the one hand, we detected an in-
teraction between canopy stratum and tree diversity on leaf traits
whereby differences in leaf traits among strata were more pronounced
in pure stands. Recent studies reported that tree diversity may alter
abiotic factors in individual canopies (and in particular light environ-
ment), and hence insect herbivory (Castagneyrol et al., 2018a, 2017;
Muiruri and Koricheva, 2016; Rosado-Sánchez et al., 2017). Given the
different growth form of oaks and pines, it is likely that the vertical
stratification of abiotic factors was far more heterogeneous in mixed
than in pure stands (Forrester, 2017), resulting in neater vertical gra-
dients of leaf traits in pure stands. On the other hand, we found sig-
nificant relationships between leaf traits and herbivory. In particular,
leaf insect herbivory increased with increasing SLA and decreased with
increasing LDMC. However, despite this direct effect of leaf traits on
herbivory, we still detected a direct effect of forest stratum on leaf in-
sect herbivory, while the effect of tree diversity on herbivory seemed to
be primarily driven by an effect of tree diversity on leaf traits. To the
best of our knowledge, to date no study addressing leaf trait-mediated
effects of tree diversity on leaf insect herbivory has systematically
compared leaves from different strata. Yet our results indicate that
controlling for this effect will be critical for future studies of tree-her-
bivore interactions in mixed forests.
Leaf insect herbivory increased with oak isolation at the
landscape scale. A long held view in ecology is that herbivory in-
creases with the density of resources (resource concentration hypoth-
esis, Root, 1973). Yet, we found the opposite. Whereas studies on forest
fragmentation often find that herbivore abundance and richness de-
crease with patch size and isolation (Rossetti et al., 2017), results are
less consistent for herbivory itself (Maguire et al., 2016; Rossetti et al.,
2017; Simonetti et al., 2007). Our results suggest that the variation in
outcomes of previous studies may to a considerable extent be caused by
tree cues at the within-patch and individual tree scales that dilute
among-patch trends in herbivory. For instance, virtually no studies we
are aware of have to date accounted for stratification effects. Yet such
effects could be far more determinant for patterns of herbivory than for
herbivore richness or abundance if they affect the quality of the plant
diet – which was the case in our system. On the other hand, we cannot
exclude that the trend we observed was to some extent also mediated by
differential top-down control of herbivores by predators, as has been
reported by some authors (Gonzalez-Gomez et al., 2006; Maguire et al.,
2015; Rossetti et al., 2014). Likewise, we present here the results of a
single year-study that was conducted in 12 mixed and pure forest stands
in southwestern France. We cannot exclude that our results may reflect
particular abiotic conditions and the management applied to forest
stands in this region.
6. Conclusions
Multiple independent approaches have been used so far to address
the effect of tree diversity on insect herbivory. Studies on the influence
of habitat diversity at the landscape level and tree diversity within
forest patches mainly addressed insect movements and population dy-
namics at large scales. Studies focusing on a much smaller scale
B. Castagneyrol et al.
addressed tree-tree interactions and their effects on herbivores through
changes in leaf traits. Here, we linked knowledge on tree-herbivore
interactions from these different perspectives by integrating landscape-,
forest type- and individual-levels effects on insect herbivores within the
same study. By demonstrating that insect herbivory in mixed stands
exceeded that of pure stands only in the upper stratum, we unravel the
importance of considering small-scale variability of biotic and abiotic
factors when addressing insect herbivory on forest trees. Our findings
therefore help step forward towards a more profound understanding of
the complex forces that drive insect herbivory in forests.
Acknowledgements
We thank Hervé Jactel, Luc Barbaro and Emmanuel Corcket (INRA
UMR 1202 BIOGECO) for their suggestions and advices during the
design of this study. We also thank Aurélie Ornon and Jean-Paul Duvert
for providing experimental sites in the Gigama forest property, and
Jean-Marc Louvet and Rozenn Prud’homme for field assistance. BG was
supported by a grant from the French Ministry of Higher Education and
Research. EVC was founded by the ANR project SPONFOREST (grant
BIODIVERSA 2015-58).
Author contributions
BG conceived the study and acquired the data. BC, BG and EVC
analyzed the data. BC and BG drafted the first version of the manu-
script. All authors wrote the final version of the manuscript.
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York. It was a vast and variegated assemblage, composed of
persons from all sections of the North, and may be said to have
formed a new departure in the history of forces organized to resist
the growing and aggressive demands of slavery and the slave
power. Until this Buffalo convention anti-slavery agencies had been
mainly directed to the work of changing public sentiment by exposing
through the press and on the platform the nature of the slave
system. Anti-slavery thus far had only been sheet lightning; the
Buffalo convention sought to make it a thunderbolt. It is true the
Liberty party, a political organization, had been in existence since
1840, when it cast seven thousand votes for James G. Birney, a
former slaveholder, but who in obedience to an enlightened
conscience, had nobly emancipated his slaves, and was now
devoting his time and talents to the overthrow of slavery. It is true
that this little party of brave men had increased their numbers at one
time to sixty thousand voters. It, however, had now apparently
reached its culminating point, and was no longer able to attract to
itself and combine all the available elements at the North, capable of
being marshaled against the growing and aggressive measures and
aims of the slave power. There were many in the old Whig party
known as Conscience-Whigs, and in the Democratic party known as
Barnburners and Free Democrats, who were anti-slavery in
sentiment and utterly opposed to the extension of the slave system
to territory hitherto uncursed by its presence, but who nevertheless
were not willing to join the Liberty party. It was held to be deficient in
numbers and wanting in prestige. Its fate was the fate of all pioneers.
The work it had been required to perform had exposed it to assaults
from all sides, and it wore on its front the ugly marks of conflict. It
was unpopular for its very fidelity to the cause of liberty and justice.
No wonder that some of its members, such as Gerrit Smith, William
Goodell, Beriah Green, and Julius Lemoyne refused to quit the old
for the new. They felt that the Free-Soil party was a step backward, a
lowering of the standard, that the people should come to them, not
they to the people. The party which had been good enough for them
ought to be good enough for all others. Events, however, over-ruled
this reasoning. The conviction became general that the time had
come for a new organization, which should embrace all who were in
any manner opposed to slavery and the slave power, and this
Buffalo Free-Soil convention was the result of that conviction. It is
easy to say that this or that measure would have been wiser and
better than the one adopted. But any measure is vindicated by its
necessity and its results. It was impossible for the mountain to go to
Mahomet, or for the Free-Soil element to go to the old Liberty party,
so the latter went to the former. “All is well that ends well.” This
Buffalo convention of free-soilers, however low was their standard,
did lay the foundation of a grand superstructure. It was a powerful
link in the chain of events by which the slave system has been
abolished, the slave emancipated, and the country saved from
dismemberment.
It is nothing against the actors in this new movement that they
did not see the end from the beginning; that they did not at first take
the high ground that further on in the conflict their successors felt
themselves called upon to take, or that their free-soil party, like the
old liberty party, was ultimately required to step aside and make
room for the great Republican party. In all this and more it illustrates
the experience of reform in all ages, and conforms to the laws of
human progress—Measures change, principles never.
I was not the only colored man well known to the country who
was present at this convention. Samuel Ringold Ward, Henry
Highland Garnet, Charles L. Remond, and Henry Bibb, were there
and made speeches which were received with surprise and
gratification by the thousands there assembled. As a colored man I
felt greatly encouraged and strengthened for my cause while
listening to these men—in the presence of the ablest men of the
Caucasian race. Mr. Ward especially attracted attention at that
convention. As an orator and thinker he was vastly superior, I
thought, to any of us, and being perfectly black and of unmixed
African descent, the splendors of his intellect went directly to the
glory of his race. In depth of thought, fluency of speech, readiness of
wit, logical exactness, and general intelligence, Samuel R. Ward has
left no successor among the colored men amongst us, and it was a
sad day for our cause when he was laid low in the soil of a foreign
country.
 After the Free Soil party, with “Free Soil,” “Free Labor,” “Free
States,” “Free Speech,” and “Free Men,” on its banner, had defeated
the almost permanently victorious Democratic party under the
leadership of so able and popular a standard-bearer as General
Lewis Cass, Mr. Calhoun and other southern statesmen were more
than ever alarmed at the rapid increase of anti-slavery feeling in the
North, and devoted their energies more and more to the work of
devising means to stay the torrents and tie up the storm. They were
not ignorant of whereunto this sentiment would grow if unsubjected
and unextinguished. Hence they became fierce and furious in
debate, and more extravagant than ever in their demands for
additional safeguards for their system of robbery and murder.
Assuming that the Constitution guaranteed their rights of property in
their fellowmen, they held it to be in open violation of the Constitution
for any American citizen in any part of the United States to speak,
write, or act against this right. But this shallow logic they plainly saw
could do them no good unless they could obtain further safeguards
for slavery. In order to effect this, the idea of so changing the
Constitution was suggested, that there should be two instead of one
President of the United States—one from the North and the other
from the South—and that no measure should become a law without
the assent of both. But this device was so utterly impracticable that it
soon dropped out of sight, and it is mentioned here only to show the
desperation of slaveholders to prop up their system of barbarism
against which the sentiment of the North was being directed with
destructive skill and effect. They clamored for more slave States,
more power in the Senate and House of Representatives, and
insisted upon the suppression of free speech. At the end of two
years, in 1850, when Clay and Calhoun, two of the ablest leaders the
South ever had, were still in the Senate, we had an attempt at a
settlement of differences between the North and South which our
legislators meant to be final. What those measures were I need not
here enumerate except to say that chief among them was the
Fugitive Slave Bill, framed by James M. Mason of Virginia, and
supported by Daniel Webster of Massachusetts; a bill undoubtedly
more designed to involve the North in complicity with slavery and
deaden its moral sentiment than to procure the return of fugitives to
their so-called owners. For a time this design did not altogether fail.
Letters, speeches, and pamphlets literally rained down upon the
people of the North, reminding them of their constitutional duty to
hunt down and return to bondage runaway slaves. In this the
preachers were not much behind the press and the politicians,
especially that class of preachers known as Doctors of Divinity. A
long list of these came forward with their Bibles to show that neither
Christ nor his holy apostles objected to returning fugitives to slavery.
Now that that evil day is past a sight of those sermons would, I doubt
not, bring the red blush of shame to the cheeks of many.
Living as I then did in Rochester, on the border of Canada, I was
compelled to see the terribly distressing effects of this cruel
enactment. Fugitive slaves, who had lived for many years safely and
securely in Western New York and elsewhere, some of whom had by
industry and economy saved money and bought little homes for
themselves and their children, were suddenly alarmed and
compelled to flee to Canada for safety as from an enemy’s land—a
doomed city—and take up a dismal march to a new abode, empty-
handed, among strangers. My old friend Ward, of whom I have just
now spoken, found it necessary to give up the contest and flee to
Canada, and thousands followed his example. Bishop Daniel A.
Payne, of the African Methodist Episcopal church, came to me about
this time to consult me as to whether it was best to stand our ground
or flee to Canada. When I told him I could not desert my post until I
saw I could not hold it, adding that I did not wish to leave while
Garnet and Ward remained, “Why,” said he, “Ward, Ward, he is
already gone. I saw him crossing from Detroit to Windsor.” I asked
him if he was going to stay, and he answered, “Yes; we are whipped,
we are whipped! and we might as well retreat in order.” This was
indeed a stunning blow. This man had power to do more to defeat
this inhuman enactment than any other colored man in the land, for
no other could bring such brain power to bear against it. I felt like a
besieged city at news that its defenders had fallen at its gates.
The hardships imposed by this atrocious and shameless law
were cruel and shocking, and yet only a few of all the fugitives of the
Northern States were returned to slavery under its infamously wicked
provisions. As a means of recapturing their runaway property in
human flesh the law was an utter failure. Its efficiency was destroyed
by its enormity. Its chief effect was to produce alarm and terror
among the class subject to its operation, and this it did most
effectually and distressingly. Even colored people who had been free
all their lives felt themselves very insecure in their freedom, for under
this law the oaths of any two villains were sufficient to consign a free
man to slavery for life. While the law was a terror to the free, it was a
still greater terror to the escaped bondman. To him there was no
peace. Asleep or awake, at work or at rest, in church or market, he
was liable to surprise and capture. By the law the judge got ten
dollars a head for all he could consign to slavery, and only five
dollars apiece for any which he might adjudge free. Although I was
now myself free, I was not without apprehension. My purchase was
of doubtful validity, having been bought when out of the possession
of my owner and when he must take what was given or take nothing.
It was a question whether my claimant could be estopped by such a
sale from asserting certain or supposable equitable rights in my body
and soul. From rumors that reached me my house was guarded by
my friends several nights, when kidnappers, had they come, would
have got anything but a cool reception, for there would have been
“blows to take as well as blows to give.” Happily this reign of terror
did not continue long. Despite the efforts of Daniel Webster and
Millard Fillmore and our Doctors of Divinity, the law fell rapidly into
disrepute. The rescue of Shadrack resulting in the death of one of
the kidnappers, in Boston, the cases of Simms and Anthony Burns,
in the same place, created the deepest feeling against the law and
its upholders. But the thing which more than all else destroyed the
fugitive slave law was the resistance made to it by the fugitives
themselves. A decided check was given to the execution of the law
at Christiana, Penn., where three colored men, being pursued by Mr.
Gorsuch and his son, slew the father, wounded the son, and drove
away the officers, and made their escape to my house in Rochester.
The work of getting these men safely into Canada was a delicate
one. They were not only fugitives from slavery but charged with
murder, and officers were in pursuit of them. There was no time for
delay. I could not look upon them as murderers. To me, they were
heroic defenders of the just rights of man against manstealers and
murderers. So I fed them, and sheltered them in my house. Had they
been pursued then and there, my home would have been stained
with blood, for these men who had already tasted blood were well
armed and prepared to sell their lives at any expense to the lives and
limbs of their probable assailants. What they had already done at
Christiana and the cool determination which showed very plainly
especially in Parker, (for that was the name of the leader,) left no
doubt on my mind that their courage was genuine and that their
deeds would equal their words. The situation was critical and
dangerous. The telegraph had that day announced their deeds at
Christiana, their escape, and that the mountains of Pennsylvania
were being searched for the murderers. These men had reached me
simultaneously with this news in the New York papers. Immediately
after the occurrence at Christiana, they, instead of going into the
mountains, were placed on a train which brought them to Rochester.
They were thus almost in advance of the lightning, and much in
advance of probable pursuit, unless the telegraph had raised agents
already here. The hours they spent at my house were therefore
hours of anxiety as well as activity. I dispatched my friend Miss Julia
Griffiths to the landing three miles away on the Genesee River to
ascertain if a steamer would leave that night for any port in Canada,
and remained at home myself to guard my tired, dust-covered, and
sleeping guests, for they had been harassed and traveling for two
days and nights, and needed rest. Happily for us the suspense was
not long, for it turned out, that that very night a steamer was to leave
for Toronto, Canada.
This fact, however, did not end my anxiety. There was danger
that between my house and the landing or at the landing itself we
might meet with trouble. Indeed the landing was the place where
trouble was likely to occur if at all. As patiently as I could, I waited for
the shades of night to come on, and then put the men in my
“Democrat carriage,” and started for the landing on the Genesee. It
was an exciting ride, and somewhat speedy withal. We reached the
boat at least fifteen minutes before the time of its departure, and that
without remark or molestation. But those fifteen minutes seemed
much longer than usual. I remained on board till the order to haul in
the gang-way was given; I shook hands with my friends, received
from Parker the revolver that fell from the hand of Gorsuch when he
died, presented now as a token of gratitude and a memento of the
battle for Liberty at Christiana, and I returned to my home with a
sense of relief which I cannot stop here to describe. This affair, at
Christiana, and the Jerry Rescue at Syracuse, inflicted fatal wounds
on the fugitive slave bill. It became thereafter almost a dead letter,
for slaveholders found that not only did it fail to put them in
possession of their slaves, but that the attempt to enforce it brought
odium upon themselves and weakened the slave system.
In the midst of these fugitive slave troubles came the book
known as Uncle Tom’s Cabin, a work of marvelous depth and power.
Nothing could have better suited the moral and humane
requirements of the hour. Its effect was amazing, instantaneous, and
universal. No book on the subject of slavery had so generally and
favorably touched the American heart. It combined all the power and
pathos of preceding publications of the kind, and was hailed by many
as an inspired production. Mrs. Stowe at once became an object of
interest and admiration. She had made fortune and fame at home,
and had awakened a deep interest abroad. Eminent persons in
England roused to anti-slavery enthusiasm by her “Uncle Tom’s
Cabin,” invited her to visit that country, and promised to give her a
testimonial. Mrs. Stowe accepted the invitation and the proffered
testimonial. Before sailing for England, however, she invited me from
Rochester, N. Y., to spend a day at her house in Andover, Mass.
Delighted with an opportunity to become personally acquainted with
the gifted authoress, I lost no time in making my way to Andover. I
was received at her home with genuine cordiality. There was no
contradiction between the author and her book. Mrs. Stowe
appeared in conversation equally as well as she appeared in her
writing. She made to me a nice little speech in announcing her object
in sending for me. “I have invited you here,” she said, “because I
wish to confer with you as to what can be done for the free colored
people of the country. I am going to England and expect to have a
considerable sum of money placed in my hands, and I intend to use
it in some way, for the permanent improvement of the free colored
people, and especially for that class which has become free by their
own exertions. In what way I can do this most successfully is the
subject I wish to talk with you about. In any event I desire to have
some monument rise after Uncle Tom’s Cabin, which shall show that
it produced more than a transient influence.” She said several plans
had been suggested, among others an educational institution pure
and simple, but that she thought favorably of the establishment of an
industrial school; and she desired me to express my views as to
what I thought would be the best plan to help the free colored
people. I was not slow to tell Mrs. Stowe all I knew and had thought
on the subject. As to a purely educational institution, I agreed with
her that it did not meet our necessities. I argued against expending
money in that way. I was also opposed to an ordinary industrial
school where pupils should merely earn the means of obtaining an
education in books. There were such schools, already. What I
thought of as best was rather a series of workshops, where colored
people could learn some of the handicrafts, learn to work in iron,
wood, and leather, and where a plain English education could also
be taught. I argued that the want of money was the root of all evil to
the colored people. They were shut out from all lucrative
employments and compelled to be merely barbers, waiters,
coachmen and the like at wages so low that they could lay up little or
nothing. Their poverty kept them ignorant and their ignorance kept
them degraded. We needed more to learn how to make a good living
than to learn Latin and Greek. After listening to me at considerable
length, she was good enough to tell me that she favored my views,
and would devote the money she expected to receive abroad to
meeting the want I had described as the most important; by
establishing an institution in which colored youth should learn trades
as well as to read, write, and count. When about to leave Andover,
Mrs. Stowe asked me to put my views on the subject in the form of a
letter, so that she could take it to England with her and show it to her
friends there, that they might see to what their contributions were to
be devoted. I acceded to her request and wrote her the following
letter for the purpose named.

Rochester, March 8, 1853.

 My Dear Mrs. Stowe:
You kindly informed me, when at your house a fortnight ago,
that you designed to do something which should permanently
contribute to the improvement and elevation of the free colored
people in the United States. You especially expressed an
interest in such of this class as had become free by their own
exertions, and desired most of all to be of service to them. In
what manner, and by what means you can assist this class most
successfully, is the subject upon which you have done me the
honor to ask my opinion.... I assert then that poverty, ignorance,
and degradation are the combined evils; or in other words, these
constitute the social disease of the free colored people of the
United States.
To deliver them from this triple malady, is to improve and
elevate them, by which I mean simply to put them on an equal
footing with their white fellow countrymen in the sacred right to
“Life, Liberty, and the pursuit of happiness.” I am for no fancied
or artificial elevation, but only ask fair play. How shall this be
obtained? I answer, first, not by establishing for our use high
schools and colleges. Such institutions are, in my judgment,
beyond our immediate occasions and are not adapted to our
present most pressing wants. High schools and colleges are
excellent institutions, and will in due season be greatly
subservient to our progress; but they are the result, as well as
they are the demand of a point of progress, which we as a
people have not yet attained. Accustomed as we have been, to
the rougher and harder modes of living, and of gaining a
livelihood, we cannot, and we ought not to hope that in a single
leap from our low condition, we can reach that of Ministers,
Lawyers, Doctors, Editors, Merchants, etc. These will doubtless
be attained by us; but this will only be, when we have patiently
and laboriously, and I may add successfully, mastered and
passed through the intermediate gradations of agriculture and
the mechanic arts. Besides, there are (and perhaps this is a
better reason for my view of the case) numerous institutions of
learning in this country, already thrown open to colored youth. To
my thinking, there are quite as many facilities now afforded to
the colored people, as they can spare the time, from the sterner
duties of life, to avail themselves of. In their present condition of
poverty, they cannot spare their sons and daughters two or three
years at boarding-schools or colleges, to say nothing of finding
the means to sustain them while at such institutions. I take it,
therefore, that we are well provided for in this respect; and that it
may be fairly inferred from the fact, that the facilities for our
education, so far as schools and colleges in the Free States are
concerned, will increase quite in proportion with our future
wants. Colleges have been open to colored youth in this country
during the last dozen years. Yet few comparatively, have
acquired a classical education; and even this few have found
themselves educated far above a living condition, there being no
methods by which they could turn their learning to account.
Several of this latter class have entered the ministry; but you
need not be told that an educated people is needed to sustain
an educated ministry. There must be a certain amount of
cultivation among the people, to sustain such a ministry. At
present we have not that cultivation amongst us; and therefore,
we value in the preacher, strong lungs, rather than high learning.
I do not say, that educated ministers are not needed amongst
us, far from it! I wish there were more of them! but to increase
their number, is not the largest benefit you can bestow upon us.
We have two or three colored lawyers in this country; and I
rejoice in the fact; for it affords very gratifying evidence of our
progress. Yet it must be confessed, that in point of success, our
lawyers are as great failures as our ministers. White people will
not employ them to the obvious embarrassment of their causes,
and the blacks, taking their cue from the whites, have not
sufficient confidence in their abilities to employ them. Hence
educated colored men, among the colored people, are at a very
great discount. It would seem that education and emigration go
together with us, for as soon as a man rises amongst us,
capable, by his genius and learning, to do us great service, just
so soon he finds that he can serve himself better by going
elsewhere. In proof of this, I might instance the Russwurms, the
Garnetts, the Wards, the Crummells and others, all men of
superior ability and attainments, and capable of removing
mountains of prejudice against their race, by their simple
presence in the country; but these gentlemen, finding
themselves embarrassed here by the peculiar disadvantages to
which I have referred, disadvantages in part growing out of their
education, being repelled by ignorance on the one hand, and
prejudice on the other, and having no taste to continue a contest
against such odds, they have sought more congenial climes,
where they can live more peaceable and quiet lives. I regret their
election, but I cannot blame them; for with an equal amount of
education and the hard lot which was theirs, I might follow their
example....
There is little reason to hope that any considerable number
of the free colored people will ever be induced to leave this
country, even if such a thing were desirable. The black man
(unlike the Indian) loves civilization. He does not make very
great progress in civilization himself but he likes to be in the
midst of it, and prefers to share its most galling evils, to
encountering barbarism. Then the love of country, the dread of
isolation, the lack of adventurous spirit, and the thought of
seeming to desert their “brethren in bonds,” are a powerful check
upon all schemes of colonization, which look to the removal of
the colored people, without the slaves. The truth is, dear
madam, we are here, and here we are likely to remain.
Individuals emigrate—nations never. We have grown up with this
republic, and I see nothing in her character, or even in the
character of the American people, as yet which compels the
belief that we must leave the United States. If then, we are to
remain here the question for the wise and good is precisely that
you have submitted to me—namely: What can be done to
improve the condition of the free people of color in the United
States? The plan which I humbly submit in answer to this inquiry
(and in the hope that it may find favor with you, and with the
many friends of humanity who honor, love, and coöperate with
you) is the establishment in Rochester, N. Y., or in some other
part of the United States equally favorable to such an enterprise,
of an Industrial College in which shall be taught several
important branches of the mechanic arts. This college to be
open to colored youth. I will pass over the details of such an
institution as I propose.... Never having had a day’s schooling in
all my life I may not be expected to map out the details of a plan
so comprehensive as that involved in the idea of a college. I
repeat, then, I leave the organization and administration to the
superior wisdom of yourself and the friends who second your
noble efforts. The argument in favor of an Industrial College (a
college to be conducted by the best men, and the best workmen
which the mechanic arts can afford; a college where colored
youth can be instructed to use their hands, as well as their
heads; where they can be put in possession of the means of
getting a living whether their lot in after life may be cast among
civilized or uncivilized men; whether they choose to stay here, or
prefer to return to the land of their fathers) is briefly this:
Prejudice against the free colored people in the United States
has shown itself nowhere so invincible as among mechanics.
The farmer and the professional man cherish no feeling so bitter
as that cherished by these. The latter would starve us out of the
country entirely. At this moment I can more easily get my son
into a lawyer’s office to study law than I can into a blacksmith’s
shop to blow the bellows and to wield the sledge-hammer.
Denied the means of learning useful trades we are pressed into
the narrowest limits to obtain a livelihood. In times past we have
been the hewers of wood and drawers of water for American
society, and we once enjoyed a monopoly in menial
employments, but this is so no longer. Even these employments
are rapidly passing away out of our hands. The fact is (every day
begins with the lesson, and ends with the lesson) that colored
men must learn trades; must find new employments; new modes
of usefulness to society, or that they must decay under the
pressing wants to which their condition is rapidly bringing them.
We must become mechanics; we must build as well as live
in houses; we must make as well as use furniture; we must
construct bridges as well as pass over them, before we can
properly live or be respected by our fellow men. We need
mechanics as well as ministers. We need workers in iron, clay,
and leather. We have orators, authors, and other professional
men, but these reach only a certain class, and get respect for
our race in certain select circles. To live here as we ought we
must fasten ourselves to our countrymen through their every day
cardinal wants. We must not only be able to black boots, but to
make them. At present we are unknown in the northern States
as mechanics. We give no proof of genius or skill at the county,
State, or national fairs. We are unknown at any of the great
exhibitions of the industry of our fellow-citizens, and being
unknown we are unconsidered.
The fact that we make no show of our ability is held
conclusive of our inability to make any, hence all the indifference
and contempt with which incapacity is regarded fall upon us, and
that too when we have had no means of disproving the infamous
opinion of our natural inferiority. I have during the last dozen
years denied before the Americans that we are an inferior race;
but this has been done by arguments based upon admitted
principles rather than by the presentation of facts. Now firmly
believing, as I do, that there are skill, invention, power, industry,
and real mechanical genius, among the colored people, which
will bear favorable testimony for them, and which only need the
means to develop them, I am decidedly in favor of the
establishment of such a college as I have mentioned. The
benefits of such an institution would not be confined to the
Northern States, nor to the free colored people. They would
extend over the whole Union. The slave not less than the
freeman would be benefited by such an institution. It must be
confessed that the most powerful argument now used by the
southern slaveholder, and the one most soothing to his
conscience, is that derived from the low condition of the free
colored people of the north. I have long felt that too little
attention has been given by our truest friends in this country to
removing this stumbling block out of the way of the slave’s
liberation.
 The most telling, the most killing refutation of slavery, is the
presentation of an industrious, enterprising, thrifty, and intelligent
free black population. Such a population I believe would rise in
the Northern States under the fostering care of such a college as
that supposed.
To show that we are capable of becoming mechanics I might
adduce any amount of testimony; but, dear madam, I need not
ring the changes on such a proposition. There is no question in
the mind of any unprejudiced person that the Negro is capable of
making a good mechanic. Indeed, even those who cherish the
bitterest feelings towards us have admitted that the
apprehension that negroes might be employed in their stead,
dictated the policy of excluding them from trades altogether. But
I will not dwell upon this point as I fear I have already trespassed
too long upon your precious time, and written more than I ought
to expect you to read. Allow me to say in conclusion, that I
believe every intelligent colored man in America will approve and
rejoice at the establishment of some such institution as that now
suggested. There are many respectable colored men, fathers of
large families, having boys nearly grown up, whose minds are
tossed by day and by night with the anxious inquiry, what shall I
do with my boys? Such an institution would meet the wants of
such persons. Then, too, the establishment of such an institution
would be in character with the eminently practical philanthropy of
your trans-Atlantic friends. America could scarcely object to it as
an attempt to agitate the public mind on the subject of slavery, or
to dissolve the Union. It could not be tortured into a cause for
hard words by the American people, but the noble and good of
all classes would see in the effort an excellent motive, a
benevolent object, temperately, wisely, and practically
manifested.
Wishing you, dear madam, renewed health, a pleasant
passage and safe return to your native land,
I am most truly, your grateful friend,
Frederick Douglass.
 Mrs. H. B. Stowe.

I was not only requested to write the foregoing letter for the
purpose indicated, but I was also asked, with admirable foresight, to
see and ascertain, as far as possible, the views of the free colored
people themselves in respect to the proposed measure for their
benefit. This I was able to do in July, 1853, at the largest and most
enlightened colored convention that, up to that time, had ever
assembled in this country. This convention warmly approved the plan
of a manual labor school, as already described, and expressed high
appreciation of the wisdom and benevolence of Mrs. Stowe. This
convention was held in Rochester, N. Y., and will long be
remembered there for the surprise and gratification it caused our
friends in that city. They were not looking for such exhibition of
enlightened zeal and ability as were there displayed in speeches,
addresses, and resolutions; and in the conduct of the business for
which it had assembled. Its proceedings attracted wide-spread
attention at home and abroad.
While Mrs. Stowe was abroad, she was attacked by the pro-
slavery press of our country so persistently and vigorously, for
receiving money for her own private use, that the Rev. Henry Ward
Beecher felt called upon to notice and reply to them in the columns
of the New York Independent, of which he was then the editor. He
denied that Mrs. Stowe was gathering British gold for herself, and
referred her assailants to me, if they would learn what she intended
to do with the money. In answer to her maligners, I denounced their
accusations as groundless, and assured the public through the
columns of my paper, that the testimonial then being raised in
England by Mrs. Stowe, would be sacredly devoted to the
establishment of an industrial school for colored youth. This
announcement was circulated by other journals, and the attacks
ceased. Nobody could well object to such application of money,
received from any source, at home or abroad. After her return to this
country, I called again on Mrs. Stowe, and was much disappointed to
learn from her that she had reconsidered her plan for the industrial
school. I have never been able to see any force in the reasons for
this change. It is enough, however, to say that they were sufficient
for her, and that she no doubt acted conscientiously, though her
change of purpose was a great disappointment, and placed me in an
awkward position before the colored people of this country, as well
as to friends abroad, to whom I had given assurances that the
money would be appropriated in the manner I have described.
 CHAPTER IX.
INCREASING DEMANDS OF THE SLAVE
POWER.

Increased demands of slavery—War in Kansas—John Brown’s raid—His

capture and execution—My escape to England from United States
marshals.

NOTWITHSTANDING the natural tendency of the human mind to

weary of an old story, and to turn away from chronic abuses for
which it sees no remedy, the anti-slavery agitation for thirty long
years—from 1830 to 1860—was sustained with ever increasing
intensity and power. This was not entirely due to the extraordinary
zeal and ability of the anti-slavery agitators themselves; for with all
their admitted ardor and eloquence, they could have done very little
without the aid rendered them, unwittingly, by the aggressive
character of slavery itself. It was in the nature of the system never to
rest in obscurity, although that condition was in a high degree
essential to its security. It was forever forcing itself into prominence.
Unconscious, apparently, of its own deformity, it omitted no occasion
for inviting disgust by seeking approval and admiration. It was
noisiest when it should have been most silent and unobtrusive. One
of its defenders, when asked what would satisfy him as a
slaveholder, said “he never would be satisfied until he could call the
roll of his slaves in the shadow of Bunker Hill monument.” Every
effort made to put down agitation only served to impart to it new
strength and vigor. Of this class was the “gag rule,” attempted and
partially enforced in Congress—the attempted suppression of the
right of petition—the mobocratic demonstrations against the exercise
of free speech—the display of pistols, bludgeons, and plantation
manners in the Congress of the nation—the demand, shamelessly
made by our government upon England, for the return of slaves who
had won their liberty by their valor on the high seas—the bill for the
recapture of runaway slaves—the annexation of Texas for the
avowed purpose of increasing the number of slave States, and thus
increasing the power of slavery in the union—the war with Mexico—
the fillibustering expeditions against Cuba and Central America—the
cold-blooded decision of Chief Justice Taney in the Dred Scott case,
wherein he states, as it were, a historical fact, that “negroes are
deemed to have no rights which white men are bound to respect”—
the perfidious repeal of the Missouri compromise, when all its
advantages to the South had been gained and appropriated, and
when nothing had been gained by the North—the armed and bloody
attempt to force slavery upon the virgin soil of Kansas—the efforts of
both of the great political parties to drive from place and power every
man suspected of ideas and principles hostile to slavery—the rude
attacks made upon Giddings, Hale, Chase, Wilson, Wm. H. Seward,
and Charles Sumner—the effort to degrade these brave men, and
drive them from positions of prominence—the summary manner in
which Virginia hanged John Brown;—in a word, whatever was done
or attempted, with a view to the support and security of slavery, only
served as fuel to the fire, and heated the furnace of agitation to a
higher degree than any before attained. This was true up to the
moment when the nation found it necessary to gird on the sword for
the salvation of the country and the destruction of slavery.
At no time during all the ten years preceding the war, was the
public mind at rest. Mr. Clay’s compromise measures in 1850,
whereby all the troubles of the country about slavery were to be “in
the deep bosom of the ocean buried,” was hardly dry on the pages of
the statute book before the whole land was rocked with rumored
agitation, and for one, I did my best by pen and voice, and by
ceaseless activity to keep it alive and vigorous. Later on, in 1854, we
had the Missouri compromise, which removed the only grand legal
barrier against the spread of slavery over all the territory of the
United States. From this time there was no pause, no repose. Every
body, however dull, could see that this was a phase of the slavery
question which was not to be slighted or ignored. The people of the
North had been accustomed to ask, in a tone of cruel indifference,
“What have we to do with slavery?” and now no labored speech was
required in answer. Slaveholding aggression settled this question for
us. The presence of slavery in a territory would certainly exclude the
sons and daughters of the free States more effectually than statutes
or yellow fever. Those who cared nothing for the slave, and were
willing to tolerate slavery inside the slave States, were nevertheless
not quite prepared to find themselves and their children excluded
from the common inheritance of the nation. It is not surprising
therefore, that the public mind of the North was easily kept intensely
alive on this subject, nor that in 1856 an alarming expression of
feeling on this point was seen in the large vote given for John C.
Fremont and William L. Dayton for President and Vice-President of
the United States. Until this last uprising of the North against the
slave power the anti-slavery movement was largely retained in the
hands of the original abolitionists, whose most prominent leaders
have already been mentioned elsewhere in this volume. After 1856 a
mightier arm and a more numerous host was raised against it, the
agitation becoming broader and deeper. The times at this point
illustrated the principle of tension and compression, action and
reaction. The more open, flagrant, and impudent the slave power,
the more firmly it was confronted by the rising anti-slavery spirit of
the North. No one act did more to rouse the north to a
comprehension of the infernal and barbarous spirit of slavery and its
determination to “rule or ruin,” than the cowardly and brutal assault
made in the American Senate upon Charles Sumner, by Preston S.
Brooks, a member of Congress from South Carolina. Shocking and
scandalous as was this attack, the spirit in which the deed was
received and commended by the community, was still more
disgraceful. Southern ladies even applauded the armed bully for his
murderous assault upon an unarmed northern Senator, because of
words spoken in debate! This more than all else told the thoughtful
people of the North the kind of civilization to which they were linked,
and how plainly it foreshadowed a conflict on a larger scale.
As a measure of agitation, the repeal of the Missouri
Compromise alluded to, was perhaps the most effective. It was that
which brought Abraham Lincoln into prominence, and into conflict
with Stephen A. Douglas (who was the author of that measure) and
compelled the Western States to take a deeper interest than they
ever had done before in the whole question. Pregnant words were
now spoken on the side of freedom, words which went straight to the
heart of the nation. It was Mr. Lincoln who told the American people
at this crisis that the “Union could not long endure half slave and half
free; that they must be all one or the other, and that the public mind
could find no resting place but in the belief in the ultimate extinction
of slavery.” These were not the words of an abolitionist—branded a
fanatic, and carried away by an enthusiastic devotion to the Negro—
but the calm, cool, deliberate utterance of a statesman,
comprehensive enough to take in the welfare of the whole country.
No wonder that the friends of freedom saw in this plain man of Illinois
the proper standard-bearer of all the moral and political forces which
could be united and wielded against the slave power. In a few simple
words he had embodied the thought of the loyal nation, and
indicated the character fit to lead and guide the country amid perils
present and to come.
The South was not far behind the North in recognizing Abraham
Lincoln as the natural leader of the rising political sentiment of the
country against slavery, and it was equally quick in its efforts to
counteract and destroy his influence. Its papers teemed with the
bitterest invectives against the “backwoodsman of Illinois,” the “flat-
boatman,” the “rail-splitter,” the “third-rate lawyer,” and much else
and worse.
Preceding the repeal of the Missouri Compromise I gave, at the
anniversary of the American Anti-Slavery Society in New York, the
following picture of the state of the anti-slavery conflict as it then
existed:

“It is evident that there is in this country a purely slavery

party, a party which exists for no other earthly purpose but to
promote the interest of slavery. It is known by no particular
name, and has assumed no definite shape, but its branches
reach far and wide in church and state. This shapeless and
nameless party is not intangible in other and more important
respects. It has a fixed, definite, and comprehensive policy