Tree diversity effects on leaf insect

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Bastien Castagneyrol & Brice Giffard &
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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

288
B. Castagneyrol et al. Forest Ecology and Management 433 (2019) 287–294

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

289
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
290
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
291
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
292
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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these are on printed books ranging between 1478 and 1490. The
fourth specimen is more important as affording a definite clue to a
Cambridge origin, for it is on the second volume of the college
register. A search for and examination of such bindings as remain in
Cambridge might result in the discovery of some inscription or other
evidence which might enable us to identify the binder.
The earliest of the important sixteenth-century binders and stationers
is Garret Godfrey, and his name is first found affixed to a deed in
1503 which itself relates to the position of stationers and
bookbinders in the University. He was a native of the Low Countries,
and from the occurrence of the name Graten on some waste sheets
extracted from some of his bindings, is supposed to have come from
Graten in Limburg. He seems to have occupied a good position in
Cambridge, and was chosen as one of the churchwardens of St
Mary the Great in 1516, and continued to fill offices in connexion with
the church for several years. He died in 1539 and was buried in St
Mary’s, while his will, dated September 12, was proved on October
11.
Godfrey was well supplied with binding tools, and we find him in
possession of at least five rolls. The most important contains figures
of a griffin, a wyvern, and a lion, separated from each other by
branches of foliage. Below the lion are the binder’s initials, the
second having a sign like an arrow-head springing from the top,
representing presumably his merchant’s mark. The workmanship of
this roll very much resembles that of some of the fine London rolls of
the time, those for example of John Reynes or Thomas Symonds,
and it is quite probable that they may all have been engraved by the
same man.
Godfrey’s second roll is divided into five divisions. The first four
contain a turretted gateway with portcullis, a fleur-de-lys, a
pomegranate, and a Tudor rose, each under a canopy and
surmounted by a Royal crown. The fifth contains the binder’s initials
G. G. and between them a shield charged with three horse-shoes.
This introduction of a private heraldic shield into a bookbinder’s
ornament, is, so far as my experience goes, quite without parallel
and certainly requires explanation. Occurring as it does, between the
binder’s initials, it must obviously have some connexion with him,
and the initials are certainly those of Garret Godfrey. The late Sir
Augustus Franks suggested that the initials stood for Guido Gimpus.
The name is not mentioned by Burke in his Armorial, but Papworth
allows a coat “sable, three horse-shoes argent,” to a family, Vytan-
Gimpus, on the authority of Glover’s Ordinary. A third roll containing
the Royal emblem in compartments, in the same order as in the
previous one, also contains his initials, but with no arms. This is a
much narrower roll than the second, and poorer in design. The fourth
and fifth rolls have no mark or initials, one is of diaper work, the other
of interlaced strap work. In addition Godfrey used a few small dies.
Nicholas Speryng, the second stationer, came originally from the
Low Countries, and is found settled in Cambridge, perhaps as early
as 1506, certainly by 1513. It seems likely that he had worked as a
binder before coming to Cambridge, and that he brought some of his
binding tools with him. An examination of his bindings shows that he
often worked in a more old-fashioned style and with older tools than
Godfrey. The ornamental frames on the sides of his bindings are in
many cases built up by a repetition of single stamps, a troublesome
system superseded by the invention of the roll. As he possessed the
stamps he would naturally use them, but he would not have had
them engraved had the use of the roll been then known to him. The
two panel stamps which he used are also probably of foreign make.
These contain representations of the Annunciation on the one side,
and St Nicholas restoring the three murdered children on the other.
The Annunciation was a very favourite subject with bookbinders,
especially those of the Low Countries, while the St Nicholas had
reference to the binder’s Christian name. It is curious to notice that
the surname has been wrongly engraved, and reads Spiernick in
place of Spierinck. Some years ago some fragments of an edition of
Holt’s Grammar, the Lac Puerorum, printed in English at Antwerp by
Adrian van Berghen, were found in the Bodleian, which, from the
indentations still remaining on them, had clearly formed the boards
of one of these bindings, and on one of the fragments the name
Speyrinck was written. The piece of paper had doubtless formed the
wrapper of a parcel addressed to him.
Speryng’s rolls were all engraved after he came to England. The
finest, apparently engraved by the same hand as Godfrey’s, contains
figures of a dragon, a lion, and a wyvern amidst sprays of foliage.
Between the wyvern and dragon, in a wreath, are his mark and
initials. A smaller roll is divided into five compartments, the first four
containing a fleur-de-lys, a turretted gateway, a pomegranate, and a
Tudor rose, each surmounted by a Royal crown and under a canopy,
while the fifth contains his mark and initials. It is an almost exact
copy of one of Godfrey’s rolls, with the badges arranged in a
different order. There are two varieties of this roll. In the earlier the
binder’s initial S is a rather badly engraved letter of a curious script
form, but at a later date it has been re-engraved in the more usual
form. Weale, describing the earlier state in one place, reads the
initials N. G., and in another ascribes the roll to Segar Nicholson, a
later Cambridge stationer.
Speryng had also a roll of diaper work resembling, though clearly
differing from, Godfrey’s, and another roll with a graceful pattern of
twining foliage and flowers. He also, like Godfrey, had several dies.
There is a binding in Westminster Abbey Library on a copy of a
Codex Justiniani printed at Paris in 1515 which both Godfrey and
Speryng appear to have had a hand in with remarkable results. First
the volume was bound by Godfrey, who ornamented the side with a
framework made with his fine broad roll, with two parallel
impressions of the same roll inside the panel. The volume somehow
came later into Speryng’s hands. He filled up the vacant spaces with
his own diaper roll, and then stamped his ornamental roll with his
initials over Godfrey’s, mixing the ornaments in hopeless confusion.
We find the body of Speryng’s wyvern with the head of one of
Godfrey’s animals, and so on.
One roll used by Speryng had belonged to Siberch, and I found it
many years ago on a binding in Westminster Abbey Library.
Commenting on this binding Mr Gray writes: “We have evidence of
Siberch’s grand roll being used by Spierinck on the binding of a
Faber, Paris, 1526; the J is obliterated by the N being stamped over
it, but not completely, for Siberch’s initial can be plainly seen
underneath, and this roll was used with one of Spierinck’s own.” This
explanation is hardly clear. Speryng did not use Siberch’s roll on the
binding and then attempt to stamp an N over the J. What he did do
was to try and cut on the brass tool itself an N which should replace
the J, but the work was clumsily done and portions of the J still show.
The binding of the Faber just mentioned is stamped with Siberch’s
mutilated roll and one of Speryng’s diaper rolls, and this was the only
known example of the use of the mutilated roll. Not long ago,
however, I found in the library of Queen’s College, Oxford, a copy of
the Decretals of Panormitanus printed at Lyons in 1524 in four large
folio volumes in a magnificent Cambridge binding. On it is used the
mutilated roll of Siberch, Speryng’s largest signed roll, and his
smaller floral roll. Speryng died about the end of 1545 and was
buried in St Mary’s Church. His will, dated August 20, 1545, was
proved on January 27 following.
In 1523 an act was passed to prevent the taking of alien apprentices,
and a further act of 1529 prohibited aliens, not householders, from
exercising any handicraft, but in both acts the two Universities of
Oxford and Cambridge were exempted. In the latter year Wolsey
was petitioned by the University to be allowed to have three foreign
licensed booksellers, and when in 1534 the act was passed
restricting the dealings of foreign stationers, special permission was
granted to Cambridge to elect three stationers or printers who might
be aliens or natives. No similar privilege was either asked for by or
granted to Oxford.
The University immediately appointed the three stationers, Nicholas
Speryng, Garret Godfrey, and Segar Nicholson, who appear to have
been themselves instrumental in procuring the grant. Strangely
enough the date of this grant providing for increased activity
apparently exactly coincides with a remarkable cessation of work. No
book is known bound by Speryng printed later than 1533, though he
did not die until the end of 1545, while the latest dated book bound
by Godfrey, who died in 1539, is 1535. After this date we find no
more fine bindings. The beautiful broad rolls and panel stamps went
out of fashion and were superseded by narrow rolls of formal
renaissance ornament. As at Oxford the succession of stationers and
binders continued unbroken; but their work is of little interest.
 LECTURE IV.
TAVISTOCK, ABINGDON, ST ALBAN’S SECOND
PRESS,
IPSWICH, WORCESTER, CANTERBURY, EXETER,
“WINCHESTER,” AND “GREENWICH.”
The next place after Cambridge is Tavistock, where a book was
printed in 1525. It was an edition of the De consolatione philosophiæ
of Boethius, translated into verse and divided into eight-line stanzas.
In most manuscripts of the poem the author is given as Johannes
Capellanus, but at the end of this edition are some verses whose
first and last letters give the names of the patroness, Elizabeth
Berkeley, and the translator, Johannes Waltwnem or Walton, said to
have been a canon of Oseney. The colophon runs: “Enprented in the
exempt monastery of Tavestok in Denshyre. By me, Dan Thomas
Rychard, monke of the sayd monastery. To the instant desyre of the
ryght worshypful esquyer Mayster Robert Langdon, anno d.
MDXXV.” It was no doubt produced at the expense of Robert
Langdon of Keverell, a rough woodcut of whose arms occurs at the
end of the book. He was a wealthy Cornish gentleman, who married
an heiress and died in 1548.
The book is printed in a neat and clear black-letter. On the title-page
is a woodcut of the Almighty seated in a diamond-shaped panel, the
outside angles being filled with symbols of the four evangelists.
Copies of this book are not so rare as is generally supposed. There
are two in the Bodleian, and one in Exeter College, Oxford. One is in
the Rylands Library, another in St Andrews University, while Lord
Bute, Mr Christie Miller, and the Duke of Bedford all have examples.
The only other Tavistock book known is the Charter and Statutes of
the Stannary, printed nine years later. The colophon runs: “Here
endyth the statutes of the stannary, Imprented yn Tavystoke ye xx.
day of August, the yere of the reygne off our soveryne Lord Kynge
Henry ye viii. the xxvi. yere.” The book is in quarto, and contains
twenty-six leaves of thirty or thirty-one lines. On the title-page is a
woodcut of the King’s arms supported by angels, which occurs
several times elsewhere in the book. The colophon is on the
penultimate leaf, and on the recto of the last leaf is a woodcut of the
crucifixion of St Andrew, on the verso the same woodcut of the
Almighty as is used in the Boethius. Of this book only one copy is
known, in the library of Exeter College, Oxford. It is a most
beautifully preserved example, entirely uncut, and was bequeathed
to the college with a number of other rare books by the Rev. Joseph
Sandford.
The press at Abingdon is represented by one book only, and of that
only one copy is known, preserved amongst Archbishop Sancroft’s
books in the library of Emmanuel. It is a Breviary printed for the use
of the brethren of St Mary’s Monastery, the black monks of the order
of St Benedict. It is printed in red and black, in double columns, with
thirty-four lines to the page, and there are no headlines, catchwords,
or numbers to the leaves. When complete it should have contained
358 leaves, but unfortunately two leaves of the Kalendar containing
the months from May to August are missing. The Kalendar at
present consists of four leaves, three signed AI, AII, AIII, and one
unsigned leaf; and the missing two leaves should have come
between AII and AIII, which looks as though the printer had originally
omitted them by mistake. But so important an error could not remain
uncorrected. A very curious point about the book is that one part is
set up entirely in quires of sixes, while the rest is set up in a way
common to several early English printers, in quires alternately of
eight and four leaves, thus doing away with the half sheet. The
colophon states that the book was printed by John Scolar in the
monastery of the blessed virgin at Abingdon, in the year of our Lord,
1528, and of Thomas Rowland, the abbot, the seventeenth year.
Another colophon gives the more exact date of September 12.
Throughout the book occur some curious woodcut initials. Two, a
large E and S, are exceedingly bad copies of good work. A large C
contains the figure of a knight in armour, a very close copy of one
used earlier at Cambridge.
The printer, John Scolar, is doubtless the Oxford printer of 1517-18,
but where he had been or what he had been doing in the intervening
ten years we have not the slightest evidence to show. Since he is not
mentioned among the stationers and booksellers in the Oxford Lay
Subsidy Roll of 1524, we may presume he was not in the town.
The last of these monastic presses was at St Alban’s. The earlier
press, spoken of in a previous lecture, was at work from about 1480
to 1486, but we have no reason for believing that it was connected
with the abbey. The town is specifically mentioned in the colophons
with no reference to the abbey, nor is any person connected with the
abbey mentioned as favouring or assisting in the production of the
books. As regards the revived press the case is quite different. The
books were produced at the request of the abbot, Robert Catton, or
his successor, Richard Stevenage, and the latter’s close connexion
with the press is shown by the occurrence of his initials in the device
placed at the end of some of the books.
The second St Alban’s press was at work from 1534 up to the time of
the suppression of the abbey in 1539. One book, however, may be
earlier than this date, a Breviary of St Alban’s use. The unique copy
is in the library of the Marquis of Bute at Cardiff Castle, but I have
never had an opportunity of examining it, nor have any facsimiles
from it been published, so that it is impossible to determine when or
where it was printed. The librarian at Cardiff Castle considers it to
have been printed at the abbey about 1526.
In 1534 was issued the Lyfe and passion of Seint Albon prothomartyr
of Englande, translated by John Lidgate, and printed at the request
of Robert Catton, abbot of St Alban’s. This book, of which two copies
are known, has no printer’s name or place of printing, but is
generally ascribed to Herford’s press. The next book is Gwynneth’s
Confutation of Frith’s Book, dated 1536. Of this there is a copy in the
University Library. It again contains no printer’s name, but has a
device having in the centre the initials R. S. joined by a band,
standing for Richard Stevenage, the last abbot of St Alban’s. In 1537
there is an edition of the Introduction for to lerne to reken with the
pen. A copy was mentioned by Ames as in the possession of W.
Jones, Esq., but this has since disappeared, though a last leaf
containing the device is said to be amongst the Bagford fragments.
There may be some confusion between this and Bourman’s edition,
whose device was very similar to the one used by Herford. The
description given by Herbert makes no mention of printer, place, or
device, merely the date 1537.
Three more books are quoted by Herbert under the year 1538. A
godly disputation between Justus and Peccator, The rule of an
honest life, written by Martin, bishop of Dumience, and An epistle
against the enemies of poor people. Two, if not three, appear to have
had full colophons, stating that they were printed at St Alban’s by
John Herford for Richard Stevenage, but unfortunately no copies of
any of these books are now known, so that we have no book which
contains Herford’s name which was printed at St Alban’s. These
three books were originally mentioned in Maunsell’s catalogue, but
Herbert’s entries are considerably fuller and must have been derived
from some other source. There is no reason to doubt their
genuineness, and we know from other evidence that Herford was at
St Alban’s.
An undated book, printed at St Alban’s by Herford, was in Lord
Spencer’s collection, and is now at Manchester. It is entitled, A very
declaration of the bond and free wyll of man: the obedyence of the
gospell and what the very gospell meaneth. This book, hitherto
unknown to bibliographers, contains the name of the place only, and
has no date or printer’s name.
In 1539 Herford appears to have been indiscreet and to have printed
some very unorthodox book. Among the State papers is a letter from
Stevenage, the abbot, to Thomas Cromwell, in which he writes:
“Sent John Pryntare to London with Harry Pepwell, Bonere [Bonham]
and Tabbe of Powlles churchyard stationers, to order him at your
pleasure. Never heard of the little book of detestable heresies till the
stationers showed it me.” The book spoken of must have been
considered to be of importance, when three of the leading London
stationers were sent down to make inquiries about it. It may be that
this was the undated book last mentioned, for it alone is without the
name or device of the abbot himself, yet if the printer had thought
that he was publishing anything obnoxious to the powers, he would
hardly have added “printed at St Alban’s” to his book. The abbot’s
letter was dated October 12, and two months later the abbey was
handed over to Henry VIII.’s commissioners. Of Herford we hear
nothing more for five years.
On the suppression of the abbey the printing material was moved to
London, where it was used for a short time by a printer, Nicholas
Bourman. Now the name of the last abbot of St Alban’s was Richard
Stevenage or Boreman, and it is quite probable that this Nicholas
was some relative to whom the presses and type were entrusted
during Herford’s absence. The device used by Bourman was almost
identical with that used at St Alban’s by Herford, the initials of
Richard Stevenage in the latter being replaced by N. B., and a book
which had been printed in the abbey, the Introduction to learn to
reckon with the pen, was reprinted by Bourman in 1539. The last
definite date in a St Alban’s book is 1538, and, in the following year,
Herford, the printer, was taken to London and presumably
imprisoned or punished. The abbot was left with the printing material
on his hands, which we may suppose from the occurrence of his
initials in the device to have been at any rate partly his property. He
therefore transferred it to London for the use of his relative until such
time as Herford could resume work. This Herford did in 1544, and
Bourman transferred the material back to him. After this Bourman’s
name is found in no book, though he was a member of the
Stationers’ Company and lived to the year 1560.
The death of Henry VIII. and the accession of Edward VI. in 1547
gave a fresh impetus to printing, and presses were set up in three
new towns, Ipswich, Worcester, and Canterbury. For some years
previous to this no book had been printed outside London, and with
the religious opinions changing from day to day, even the London
printers themselves hardly knew what they might or might not issue
without fear of prosecution. Numbers of persons who had fled
abroad returned, and with them came an immense number of
foreigners seeking refuge from their own religious persecutions
abroad. As might be expected, the three new presses were mainly
devoted to printing books on the religious questions then occupying
the attention of the public.
At Ipswich, some years before printing was introduced there, we find
a book issued by a publisher. This was an edition of the Historia
Evangelica of Juvencus, and its imprint stated that it was to be sold
at Ipswich in 1534 by Reginald Oliver. A copy occurred in the first
part of Heber’s sale and was bought by Thorpe, the bookseller, but
since then all trace of it has disappeared. The edition was probably
that printed by Joannes Graphaeus at Antwerp with Oliver’s imprint
added to a certain number of copies, and this is rendered more likely
by the fact that the same printer issued in the same year an edition
of the Rudimenta-Grammatices, a school-book which had been
prescribed in 1528 by Cardinal Wolsey for use in the school which
he had just founded at Ipswich. Wolsey prescribed the use of this
book and others, not only for the use of his special school at Ipswich,
but throughout England, and we find a number of editions printed at
Antwerp to be sent over to England for sale. Juvencus was one of
the authors whose book was recommended for use.
Of Reginald Oliver we hear nothing more. He took out letters of
denization in 1535 in which he is described as “of Phrisia,” and is
perhaps the Reynold Oliver who translated a tract about an
earthquake in Mechlin in 1546 printed by Richard Lant.
As a stationer Oliver would be also a bookbinder, and perhaps the
panel stamp may be assigned to him signed with the initials R. O.
and a trade-mark. The panel is divided by a vertical line into two
halves each filled by a large medallion. That on the right contains the
Tudor rose, the left the Royal arms. In the left-hand upper corner is a
shield with the cross of St George, in the right the binder’s mark. If
he had been a London binder, this would have contained the city
arms. There is nothing to connect this binding with Oliver except the
initials, but these are uncommon, and no other stationer with them is
known.
The first person to begin printing at Ipswich was a certain Anthony
Scoloker, whom most writers have considered to be a foreigner, but
who is definitely spoken of in the Subsidy Rolls as an Englishman.
He was an educated man, printing translations of his own from
French and German, and it is not improbable that he had left
England on account of religious persecutions, and had returned
under the milder rule of Edward. His settlement in Ipswich for a few
months may perhaps be traced to the influence of Richard Argentine,
a schoolmaster and physician of that town, who was a vigorous
reformer in the reign of Edward, a violent Catholic in the time of
Mary, and a penitent Protestant again under Elizabeth, dying in 1568
as rector of St Helen’s in Ipswich.
Of the seven books printed by Scoloker at Ipswich, three were
translations by himself, three by Argentine, while the seventh was
translated by Richard Rice, Abbot of Conway. The first book issued
was The just reckoning of the whole number of years to 1547
translated by Scoloker from the German; of this an imperfect copy is
in the University Library. His two other translations were A Godly
disputation between a shoemaker and a parson, of which there is a
copy in a private library, and The Ordinary for all faithful Christians.
Of this there is a copy in the Rylands Library. Argentine’s three
translations are Luther’s Sermon upon John xx. and the true use of
the keys, Ochino’s Sermons, and Ulrich Zwingli’s Certain precepts.
Copies of these three books are not uncommon. The book translated
by Rice was Hermann’s Right Institution of Baptism.
The Just Reckoning was translated by July 6, 1547, and, we may
presume, printed shortly afterwards, but the remaining six books
were all printed in the first five months of 1548. The dates of the
translations of the three books by Argentine are January 28, January
30, and February 13. At the end of Zwingli’s Certain precepts, the
earliest of Argentine’s translations, occurs Scoloker’s device, a hand
reaching from the clouds and holding a coin to a touchstone
inscribed “Verbum Dei.” From the clouds in the left-hand corner a
face, representing the Holy Spirit, blows upon the stone. Below is
printed the text, “Prove the spirits whether they be of God.”
Scoloker’s residence in Ipswich was a short one, probably because
he did not meet with sufficient encouragement, and by June 1548 he
was settled in London, for a book by John Frith, dated June 30, was
issued by Anthony Scoloker and William Seres, dwelling without
Aldersgate. This was printed by Scoloker and has his device on the
last leaf. At London he continued in business until at least 1550.
The next printer to be noticed is John Overton, if indeed he ever
existed, for it may be stated at once that nothing is known of him but
his name. This is found in the colophon of Bale’s Catalogue of British
Writers from the time of Japhet, son of the most holy Noah, up to the
year 1548, which states explicitly that the book was printed at
Ipswich in England by John Overton on July 31, 1548. On the other
hand we find on the title-page of a number of copies a no less
definite statement that the book was printed at Wesel by
Theodoricus Plateanus on July 31, 1548. One thing is clear from an
examination of the book itself, and that is that it was certainly printed
abroad, and Bale himself, in the introduction which he wrote to John
Leland’s little book, entitled The laborious journey and search of
John Leland, writes, “Since I returned home again from Germany
where as I both collected and emprinted my simple work De
Scriptoribus Britannicis.” Most writers have tried to make the two
statements agree by suggesting that the main body of the book was
printed abroad and brought over in sheets, and that it was completed
after its arrival with two sheets printed by Overton at Ipswich. But
these sheets unfortunately are in exactly the same type as the rest of
the book. The Ipswich colophon occurs in all copies of the book, but
the Wesel imprint only in some. A possible explanation may be found
in the legal restrictions regarding the importation of foreign books. As
regards copies sold abroad it would not matter where they were
printed, but imported copies might meet with difficulties. Hence
perhaps the Ipswich colophon and the suppression of the Wesel
imprint.
The suspicion which gathered round the book and its imprints
naturally spread to the printer himself, Theodoricus Plateanus. Was
he a real person or not? Like John Overton, his name was only
known from its occurrence in this book, and it was often assumed to
be fictitious. A recent fortunate discovery made by Mr Murray of
Trinity has proved that he was a genuine printer. Fragments of
several books in Latin and German were found used in the binding of
an English law-book, and on one of these was a colophon, dated
1548, the same date as Bale’s De Scriptoribus, “Drück tho Wesel by
Dirick van der straten,” and leaves of Bale’s book were among the
fragments. The discovery is particularly fortunate, not only as
supplying the printer’s real name, which may enable us to trace him
further, but also as showing us new founts of type undoubtedly used
by him, and which were used also in several foreign printed English
books which up to now could only be conjecturally assigned to
certain towns or presses. Besides the De Scriptoribus Britannicis
several other of Bale’s works can now be definitely assigned to him.
The last of the three Ipswich printers and the most important was
John Oswen, perhaps from his name and later connexion with
Wales, a Welshman. He printed in the town apparently only during
the latter half of 1548, but in that period issued ten if not more books.
These are all without exception works of the reformers, Calvin,
Oecolampadius, Melanchthon, and others. Calvin is represented by
two books, A brief declaration of the feigned sacrament, and A
treatise on what a faithful man ought to do dwelling among the
papists. Of the latter book there is a copy in the University Library, of
the former no copy is at present known.
Melanchthon’s book Of the true authoritie of the Church, and
Oecolampadius’ Epistle, that there ought to be no respect of
personages of the poor, are both quoted by Herbert though no
copies seem to be known at present.
Of the remaining six books, there are copies of two in the University
Library, bequeathed by Mr Sandars. These are: A new book
containing an exhortation to the sick, of which another copy is in the
British Museum, and a curious little 16mo volume, An invectyve
agaynst dronkennes, of which the Cambridge copy is the only one
known. Several bibliographies speak of it as a fragment, but though
not in good condition it wants only the last leaf which would have
been blank, or contained a device: all the text is complete.
There are copies of two other books in the library of Clare College,
Peter Moone’s Short treatise of certain things abused in the Popish
church long used, and John Ramsey’s Plaister for a galled horse.
Both these books are rhyming attacks on the Catholics. There is
another copy of the first in the British Museum, but the two leaves in
the University Library which were supposed to belong to this edition
are really from the press of William Copland in London. Of Ramsey’s
book the copy at Clare is the only one known, but another edition
was printed at London by Raynalde in the same year. The next book
is a translation of Anthony Marcourt’s work, entitled, A declaration of
the Mass; of this there is a copy in the Bodleian, and finally there is
another issue of Hegendorff’s Domestical or household sermons, of
which there is a copy in the British Museum.
 Title-page of the Exhortation to the Sick, printed at
Ipswich by John Oswen in 1548.

Oswen and Scoloker between them printed within a year at least

eighteen books at Ipswich, all of them expounding the religious
views of the reformers, and examples of fourteen are known. Though
there may have been other inducements which caused these
printers to settle in that town, no doubt one of the chief reasons was
its accessibility to the Continent. Numbers of English people who
had fled abroad during the religious troubles of Henry’s reign were
pouring back into the country, and many would pass through
Ipswich, and would purchase to carry with them for their own use
and for their friends copies of the books which set forth the religious
views for which they had undergone such hardships, but which
seemed at last to be on the winning side. Again Ipswich, while easy
to get to, was equally easy to get away from, so that perhaps
discretion caused the printers to wait and see how their books were
received before they definitely settled in an inland town.
Oswen, though he continued in business a few months longer than
Scoloker, seems also to have found Ipswich an unremunerative
centre for his work, and towards the end of 1548, crossing with his
material from the east to the west of England, settled at Worcester.
On January 6, 2 Edward VI., that is at the beginning of 1549, Oswen
obtained a privilege from the King to print “every kind of book or
books set forth by us concerning the service to be used in churches,
ministration of the sacraments, and instruction of our subjects of the
Principality of Wales and marches thereunto belonging for seven
years, prohibiting all other persons whatsoever from printing the
same.” What this privilege definitely meant, or what use Oswen
made of it, is not clear. He certainly printed nothing concerning the
service of the church, or for the instruction of the subjects, specially
adapted to the Principality of Wales. He printed editions of the
Prayer-book and New Testament, but they were in English and
varied in no respect from the editions put forth by the authorised
printers, Whitchurch and Grafton. The three or four Welsh books
issued in the vernacular during the period in which his privilege was
in force, Salisbury’s Dictionary in English and Welsh and Introduction
to Welsh pronunciation, extracts from the laws of Hoel, and the
Epistles and Gospels in Welsh, books both for the instruction of the
subject and the service of the Church, were printed by other printers.
For these, or some of them, at any rate, privileges had been
obtained from Henry VIII. By the beginning of 1549 Oswen was
settled in Worcester in the High Street, issuing a book as early as
January 30. Besides his printing office in Worcester he appears to
have had another shop absolutely on the Welsh border, for at the
end of some of his colophons are the words: “They be also to sell at
Shrewsbury.”
Oswen’s four editions of the New Testament, from 1548 to 1550, as
given by Herbert, seem to resolve themselves into one quarto edition
dated January 12, 1550. Of the Prayer-book there were certainly
three issues. The first is a quarto dated May 24, 1549. It contains an
injunction which “streightly chargeth and commaundeth that no
maner of person do sell this present boke unbounde, above the price
of II Shillinges and two pence ye piece. And the same bound in
paste or in boordes, not above the pryce of thre shillynges and eyght
pence the piece.” The next edition, a folio, dated July 30, 1549, has
a similar injunction fixing the price at two shillings and sixpence
unbound, and four shillings bound. It will be noticed that His
Highness’s Council in their wisdom decreed that the cost of binding a
quarto and a folio should be exactly the same, namely eighteen
pence. The last issue, the rarest of all, was a folio printed in 1552.
Apart from Prayer-books and Testaments we know of seventeen
books printed by Oswen in the four years between 1549 and 1553.
Seven in 1549, five in 1550, three in 1551, and two in 1553.
The first book issued in 1549 was Henry Hart’s Consultorie for all
Christians. This is dated January 30, but is probably 1549 rather
than 1550, since the printer has printed at the beginning his newly-
granted privilege in full.
Six other books are dated 1549. Of these three, Hegendorff’s
Household Sermons, The Dialogue between the seditious anabaptist
and the true Christian, translated by John Veron, and the Spiritual
Matrimony between Christ and the Church, are given by Herbert, but
he had apparently never seen copies and none are at present
known. Of Certeyne sermons appointed by the King’s majesty to be
read, there are two copies in the British Museum. Herbert quotes A
message from King Edward VI., concerning obedience to religion, as
printed by Oswen at Worcester, August 5, 1549. This is apparently
another edition of the message “to certain of his people assembled
in Devonshire” printed by Grafton in July, but addressed to the
people of Wales.
The last of the year’s books was a Psalter printed in red and black
and issued on September 1, of which there is a copy in the British
Museum.
For the year 1550 there are four books, and of these one St
Ambrose, Of opression, said to be translated by Oswen himself, is
known only from the entry in Herbert, derived from Maunsell’s
catalogue. The other three, Zwingli’s Short pathway to the
Scriptures, The godly sayings of the old, ancient faithful fathers
translated by John Veron, and Gribald’s Notable and marvellous
epistle, are known, the first two from copies in the University Library
and elsewhere, the last from a copy in the Bodleian. To the end of
this year we may ascribe a very rare little octavo, Almanack and
prognostication for the year of our Lord 1551. Practised by Simon
Heringius and Lodowyke Boyard. In April 1551 a new edition was
issued of Bullinger’s Dialogue between the rebel Anabaptist and the
true Christian, which is known from copies in the Bodleian and
Brasenose College Library. Herbert had quoted an earlier edition of
1549, of which no copy is known and which is perhaps due to some
confusion with the present edition. Bullinger’s Most sure and strong
defence of the baptism of children, translated by John Veron, was
also issued this year, and a copy is in the Bodleian. The third and
last book of 1551 was issued in May, Hooper’s Godly and most
necessary annotations in the XIII. chapter to the Romans, of which
there is a copy in the University Library.
From May 1551 until May 1553, exactly two years, no dated book is
known from Oswen’s Worcester press, except the folio Prayer-book
of 1552. Nor is there any undated book to be ascribed to these
years, for Oswen’s books, so far as we know, are all dated. This is
another of those unaccountable gaps that we so often find in the
career of a printer.
The last two Worcester books were issued in 1553, a Homily to be
read in time of pestilence, written by Bishop Hooper and dated May
18, and the Statutes of the seventh year of Edward VI. A copy of the
Homily is in the University Library. It is a handsomely printed thin
quarto, and has on the title-page besides four border-pieces, a
woodcut of the Royal arms with supporters, and another of the
youthful king seated on his throne. The Statutes are described by
Herbert from a copy in his own possession, but where it is at present
is not known.
Oswen printed altogether at Worcester some twenty-one books, so
that it heads the list of provincial presses as regards numbers. The
press is also noticeable for the comparative excellence of the
printing and the variety of good border-pieces and initial letters, very
much superior to the material used by most of the contemporary
printers. From the class of books he issued we can well understand
the cessation of his press as soon as Mary succeeded to the throne.
There remain three books to be noticed which appear to be
connected with Oswen. These are John Sawtry’s Defence of the
marriage of priests, another book with a similar title by Philip
Melanchthon, and lastly a Treatise of baptism and the Lord’s Supper.
The colophon of the first runs: “Printed at Awryk by Jan Troost, 1541
in August,” that of the second, “Printed at Lipse by Ubryght Hoff,”
while the title-page says, “Translated out of latyne into englisshe by
Lewes Beuchame in the yere of the Lorde 1541 in August.” The
colophon of the last runs: “At Grunning 1541 April 27.” The three
books are obviously from the same press; initials, borders, type, all
agree, and copies are in some cases bound together. They are no
less obviously printed in England, but by whom? The only clue so far
is an initial M which appears identical with one used by Oswen, while
on the other hand the borders do not occur in any of Oswen’s books
though he uses a large selection. Again, why should all three books
be dated 1541 which, if they were the work of Oswen, must be a
false date. Though admitting the clue which connects them with
Oswen, it would be safer for the present until more evidence is
forthcoming to class them under “printer unidentified.”
At present, though a good start has been made in the Catalogue of
Early English Books in the University Library, a very great deal
remains to be done, and if done at all to be done very carefully, on
the subject of books with fictitious imprints or with none at all.
Hitherto if there was anything the least strange in the appearance of
a book, or if its subject were of a controversial nature, it was
promptly docketed “printed at a secret press on the Continent.” That