Ecology, 92(3), 2011, pp.

Ó 2011 by the Ecological Society of America

Daily temporal structure in African savanna flower visitation
networks and consequences for network sampling

Institute of Evolutionary Biology, University of Edinburgh, Kings Buildings, West Mains Road, Edinburgh EH9 3JT United Kingdom
School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 3PZ United Kingdom

Abstract. Ecological interaction networks are a valuable approach to understanding
plant–pollinator interactions at the community level. Highly structured daily activity patterns
are a feature of the biology of many flower visitors, particularly provisioning female bees,
which often visit different floral sources at different times. Such temporal structure implies
that presence/absence and relative abundance of specific flower–visitor interactions (links) in
interaction networks may be highly sensitive to the daily timing of data collection. Further,
relative timing of interactions is central to their possible role in competition or facilitation of
seed set among coflowering plants sharing pollinators. To date, however, no study has
examined the network impacts of daily temporal variation in visitor activity at a community
scale. Here we use temporally structured sampling to examine the consequences of daily
activity patterns upon network properties using fully quantified flower–visitor interaction data
for a Kenyan savanna habitat. Interactions were sampled at four sequential three-hour time
intervals between 06:00 and 18:00, across multiple seasonal time points for two sampling sites.
In all data sets the richness and relative abundance of links depended critically on when during
the day visitation was observed. Permutation-based null modeling revealed significant
temporal structure across daily time intervals at three of the four seasonal time points, driven
primarily by patterns in bee activity. This sensitivity of network structure shows the need to
consider daily time in network sampling design, both to maximize the probability of sampling
links relevant to plant reproductive success and to facilitate appropriate interpretation of
interspecific relationships. Our data also suggest that daily structuring at a community level
could reduce indirect competitive interactions when coflowering plants share pollinators, as is
commonly observed during flowering in highly seasonal habitats.
Key words: Africa; competition; ecological networks; facilitation; Kenya; mutualism; pollination;
savanna; temporal structure; visitation webs.

Pollination mutualisms drive one of the most important ecosystem services on Earth and underlie much of
the planet’s terrestrial biodiversity (Kearns et al. 1998,
Bascompte and Jordano 2007). A growing body of work
addresses the community scale of pollination processes
by applying interaction network (web) approaches to
entire plant–pollinator communities. These approaches
allow quantification of both direct and indirect interactions within and between trophic levels, allowing
examination of issues such as the basis of species
coexistence and the consequences of species addition
Manuscript received 3 June 2010; revised 25 August 2010;
accepted 2 September 2010. Corresponding Editor: R. A.
3 Present address: School of Biological Sciences, Woodland Road, Bristol BS8 1UG United Kungdom.
4 Present address: Station Biologique de Roscoff, Centre
National de la Recherche Scientifique, Adaptation et
Diversite´ en Milieu Marin (UMR 7144), Place Georges
Teissier, BP 74, 29682 Roscoff Cedex, France.
5 Corresponding author. E-mail:

or loss (Memmott and Waser 2002, 2004, Traveset and
Richardson 2006, Bascompte and Jordano 2007, Lopezaraiza-Mikel et al. 2007, Aizen et al. 2008).
Generating plant–pollinator interaction networks is
labor intensive and involves the summation of flower
visitation data collected over periods of days, weeks,
months, and occasionally years. In interpreting such
networks, it is important to remember first that not all of
the represented links between plants and pollinators
occur at the same time; and second, that for a given
network, the impact of some plant–pollinator interactions depends critically on their relative timing. For
example, if shared pollinators visit sympatric plant taxa
flowering at the same time (coflowering), plants may
compete for pollinators (Waser and Real 1979, Rathcke
1983, Stone et al. 1996, 1998). However, if visitation by
shared pollinators is partitioned in time, then coflowering plants may facilitate each others’ reproduction by
maintaining a larger pollinator population than any
could alone (Moeller 2004). Thus the interaction
between the two plant species can be negative or
positive, depending on the timing of their interactions
with pollinators.




There is widespread evidence for seasonal partitioning
of visitation by shared pollinators (Stiles 1977, Waser
and Real 1979, Pleasants 1980, Sargent and Ackerly
2008), and interaction network studies have addressed
temporal structure over the same timescale (Lundgren
and Olesen 2005, Basilio et al. 2006, Medan et al. 2006,
Kaiser-Bunbury et al. 2010). The consequences of
summing interactions over a field season were explored
by Jordano et al. (2003) in terms of ‘‘forbidden links,’’
specific plant–pollinator interactions that cannot happen
due to differences in plant and pollinator seasonal
phenology or structural incompatibilities despite their
apparent plausibility in a summed seasonal network.
For this reason Basilio et al. (2006) advise against
representing communities as a single network, but as a
seasonal series. However, with a single exception (Olesen
et al. 2008), network studies have yet to address the finer
scale temporal dynamics of plant–pollinator networks.
In particular, no study has yet examined variation in
network structure over time within a single day.
Whether such structure exists has implications for our
understanding of both the sign and magnitude of direct
and indirect interactions between species both in
pollination networks and more generally. Moreover it
also could inform and influence the collection and
interpretation of interaction data collected over longer
There are good reasons to expect daily temporal
structure in plant–pollinator interactions. Many studies
show that different pollinator taxa are active at different
times of day in a given community (e.g., Herrera 1990,
Willmer and Stone 2004) and that a given pollinator visits
different plants at different times of day (e.g., Armbruster
and Herzig 1984, Stone 1994, Stone et al. 1998). Such
variation is driven by the interaction between ‘‘bottomup’’ influences of floral resource provision (e.g., daily
timing of flower opening and provision of floral rewards;
Armbruster and Herzig 1984, Stone et al. 1999) and ‘‘topdown’’ influences of pollinator biology, including thermal
physiology (Stone 1994, Herrera 1995, Bishop and
Armbruster 1999, Stone et al. 1999), water balance
(Willmer 1988), sexual interactions (Stone 1995, Stone
et al. 1995), predation risk (Munoz and Arroyo 2004),
nesting cycles (Stone 1994, Willmer and Stone 2004), and
phylogenetic inertia (Cunningham 1991). Daily temporal
structuring of activity is particularly well-studied in bees,
whose endothermic flight physiology and need to forage
beyond their individual needs makes them uniquely
sensitive to both bottom-up and top-down structuring
factors (Herrera 1990, Stone 1994, Willmer and Stone
Whether plant–pollinator interactions show daily
temporal structuring at a community level has important consequences for the interpretation of interactions
among coflowering plants that share pollinators. For
pollinators such as bees that regularly remove pollen
from their bodies, daily temporal partitioning of
visitation among coflowering plants can turn a poten-

Ecology, Vol. 92, No. 3

tially negative interaction (competition for pollination)
into a positive interaction (facilitation of fruit set)
(Armbruster and Herzig 1984, Stone et al. 1998, Raine et
al. 2007). Which of these scenarios holds generates
diametrically opposing predictions of the impact of
species gain (by invasion or habitat restoration) or loss
(by extinction). For example, loss of a coflowering
species can result in either competitive release or
facilitation collapse. Sampling schemes that potentially
allow discrimination between these alternatives are
clearly valuable in predicting the pollination services
impact of species loss or gain on a community.
Further, if daily temporal structuring exists, recording
visitation for a limited daily time period risks missing a
subset of plant–pollinator interactions, influencing both
the richness of links recorded and their relative abundance. Although some network studies have recorded
pollinator activity through much of the day (Dupont et
al. 2003, Lundgren and Olesen 2005, Basilio et al. 2006,
Morales and Aizen 2006, Aizen et al. 2008), very few
specify when visitation to each plant taxon was recorded.
Many network studies provide no information on the
daily timing of data collection (e.g., Forup and Memmott
2005, Gibson et al. 2006, Nielsen and Bascompte 2007,
Olesen et al. 2008, Petanidou et al. 2008). Such
information is central to understanding many of the
processes contributing to the structure of such networks,
in asking, for example, whether observation at a specific
plant matched timing of floral reward release.
Here we use time-structured sampling to assess the
sensitivity of inferred flower visitation webs to the daily
timing of data collection and to assess the evidence for
daily temporal structure at a community level. To do
this we constructed networks for four diurnal time
intervals (06:00–09:00, 09:00–12:00, 12:00–15:00, and
15:00–18:00) at four seasonal time points for two
flowering plant communities in a Kenyan savanna
habitat. Our first objective was to quantify variation
between daily time intervals in flower visitation by
specific taxa. We used permutation-based null modeling
to test the hypothesis that activity is randomly
distributed across the four time intervals. Rejection of
this hypothesis implies sensitivity of network contributions of such taxa to daily timing of data collection. Our
second objective was to examine the extents to which
any signature of daily temporal structure, and the taxa
contributing such structure, are consistent through
seasonal time and across sites and years. Our third
objective was to reveal the impact of daily sampling time
on the richness of links sampled and on the proportion
of links missed when sampling is restricted to specific
time intervals.
Study location and sampling dates
Data were collected in 2004 between 3 May and 31
August at Mpala Research Centre (0817 0 N, 37852 0 E,
altitude 1800 m), Laikipia District, central Kenya (Ruiz-

March 2011


Guajardo et al. 2010). Sampling was conducted at two
0.5-ha plots (50 3 100 m), named Turkana Boma (TB)
and Junction (JN), located 6 km apart in open Acacia/
Commiphora savanna bushland. These sites were selected to differ in topography and flowering plant species,
allowing us to examine the evidence for daily temporal
structure in communities with differing species compositions. Site JN was on a rocky and dry escarpment with
sparse vegetation, while TB was on flatter, better
watered ground with a thicker vegetation cover (Baldock 2007, Ruiz-Guajardo 2008, Ruiz-Guajardo et al.
2010). Flowering plant species richness was similar at
the two sites across the four sampling months (species
totals ¼ 64 at TB and 48 at JN), although only 40% of
flowering species were common to both sites (see
Appendix A). Data were collected at TB for four
separate monthly flower–visitor networks in May, June,
July, and August and at JN for three networks in June,
July, and August. These sampling periods were selected
to coincide with three distinct phases of the annual
flowering season at Mpala. Sampling in May and June
targeted the major annual flowering period following the
long seasonal rains; sampling in July targeted the main
dry season when flowering plant species richness was
low; and sampling in August targeted a short period of
seasonal rainfall when floral species richness was again
high (Baldock 2007, Ruiz-Guajardo 2008).
Sampling rationale
Our approach was to first examine in detail the
species-rich June networks at TB and JN, assessing the
evidence for daily temporal structure in two sites with
differing flowering plant communities (objective 1). We
constructed fully quantified interaction networks for
these data sets and used a null modeling approach (see
Analysis below) to quantify the extent to which activity
by specific taxa was randomly distributed through daily
time and hence sensitive to time of sampling. We then
summarize the results of methodologically identical null
model analyses for all monthly data sets at each site to
assess the consistency of daily temporal structure across
seasonal time and to identify those taxa showing
significant structure (objective 2). Finally, to illustrate
the effect that choice of observation time interval has on
network structure (objective 3), we summarize variation
in network parameters across time intervals for each
monthly network and time interval network. We identify
the number of unique links per time interval (i.e.,
specific flower–visitor interactions observed in no other
time interval in that network) for all monthly networks
at both sites and quantify the impact, in terms of
numbers of links missed, of restricting sampling to each
single three-hour time interval.
Data collection
Data for each network were collected over two
consecutive weeks, the minimum required to devote
adequate observation effort to each floral source, with at


least two weeks between successive monthly networks.
Floral resource abundance in each plot was quantified at
the start of each week by counting the number of floral
units (defined as an individual flower or collection of
flowers that an insect of ;0.5 cm body length could
walk within or fly between [Saville 1993]) for each plant
species present. Flowering plants were identified to
species using keys and descriptions in Blundell (1992)
and Agnew and Agnew (1994) or to genus and
In each plot and month, visitation was quantified for
all plant species with at least 10 floral units. Insect visits
to flowers of each plant species were recorded during 20min observations in each of four daily sampling intervals
(06:00–09:00, 09:00–12:00, 12:00–15:00, and 15:00–
18:00) encompassing the 12-h period between dawn
and dusk. While high levels of elephant activity
precluded network sampling at night, we checked
nocturnal visitation to the species present in our plots
for individuals near the Mpala Research Centre.
Flowers of most species were closed at night and only
two species, Carissa edulis (Apocynaceae) and Turraea
mombassana (Meliaceae), were visited (by hawkmoths,
shortly after dusk). The night-flowering species are
considered in more detail in the Discussion.
Each plant species was allocated one 20-min observation period in each daily time interval in which its
flowers were open, in each week that it flowered. This
resulted in between 20 and 160 min of observation per
species per monthly network. Only one set of observations could be conducted in August, and each species in
this month received a maximum of 80 min of
observation. To avoid bias in the timing of observations
for each plant species, order of observation periods in
each time interval was randomized across plant species.
Sampling was carried out simultaneously at TB and JN
in June, July, and August with sampling days alternating
between the two plots. Visitation sampling totaled 306 h
across the four months of data collection.
For each plant species and time interval, visitation was
observed for a recorded number of floral units occupying
;1 m2. One visit was recorded every time a visitor made
contact with the sexual parts of a floral unit. Return visits
to the same floral unit were counted as a second visit. All
visitors were insects in the orders Hymenoptera, Diptera,
Coleoptera, and Lepidoptera. Species were identified
visually or captured and identified either to species by
specialist taxonomists or to morphospecies using museum
collections. Butterflies were identified to species using
Larsen (1991). Ant visits were recorded, but were very
rare and are excluded from analyses due to low
taxonomic resolution. Insects that escaped before being
identified were lumped at the taxonomic level for which
identity was certain (family or order).
Quantitative flower–visitor interaction networks.—Fully quantified flower–visitor interaction networks for the



complete (whole-day) and daily time interval June data
sets for each site were generated using software
developed by J. Memmott in Mathematica (Wolfram
Research, Champaign, Illinois, USA). Food web linkage
statistics adapted for use in plant–pollinator networks
were calculated for each monthly and time interval
network to examine the effect of daily time on network
structure. Linkage for plants and visitors represents the
mean number of links for plant species and visitor taxa,
respectively, and is a measure of plant and visitor
generalization. Linkage was calculated for visitor taxa as
lv ¼ L/V and plant species as lp ¼ L/Pv, respectively,
where L is the number of links recorded, V is the number
of visitor taxa recorded in the time interval, and Pv is the
number of plant species visited.
Null model analysis of temporal variation in flower–
visitor links.—Our underlying rationale is to compare
our field data to a null model that specifically precludes
daily temporal structure but is otherwise realistic. If
there is no daily temporal structure in our data, then the
number of different flower–visitor link types observed in
a given time interval should match that observed for an
equal quantity of data drawn at random from the
equivalent whole-day data set (i.e., across all four time
intervals). In contrast, if activity is highly structured in
daily time, specific links will be restricted to specific time
intervals, and the number of different link types
observed in specific time intervals will be lower than
the null expectation. For each time interval we
compared therefore the number of observed links with
that present in randomly generated data sets drawn from
the whole-day data set for the same month at the same
site. One thousand sets of four randomized time interval
networks were generated from each whole-day data set
using software written in Cþþ by D. Roze. In each set of
random networks, each observed visit in the whole-day
data set was randomly assigned to one of the four time
intervals, subject to two constraints: (1) The total
number of visits per time interval matched the observed
value, thus preserving variation in absolute levels of
visitation through the day. (2) Only flower–visitor links
observed in the whole-day data set were permitted, since
good biological reasons may preclude particular links
(forbidden links sensu Jordano et al. 2003).
Null models were developed for two data sets for each
month at each site: one incorporated only visits made by
taxa identified to species, while a second incorporated all
recorded visits. The first approach is conservative in that
it minimizes any potentially artifactual impact of
lumped multispecies groupings, but it also necessitates
exclusion of data in all monthly networks (11–44% of
visits). The second approach included lumped taxa,
reflecting both the difficulty of visitor identification in
biodiverse tropical habitats and the desire to maximize
data content. For each time interval and site, departure
of the observed numbers of links from null expectations
was assessed using a conservative two-tailed approach,

Ecology, Vol. 92, No. 3

i.e., we rejected the null hypothesis if the observed
frequency fell in the bottom 2.5% or above the top
97.5% of the randomly generated values. Results are
presented for all visitor taxa combined and separately
for links involving bees, wasps, flies, beetles, and
butterflies and moths. This subdivision allows us to
ask which taxa contribute to any signature of temporal
structuring observed and reflects the fact that while the
timing of visitation may be highly structured for some
taxa (particularly bees; Willmer and Stone 2004) it may
not be for others.
This null modeling approach involves four tests (one
per time interval) for each data set. Though threshold
significance (alpha) values are routinely adjusted for
multiple tests, the application of such procedures
remains an area of active debate (Perneger 1998, Moran
2003, Nakagawa 2004). Commonly applied corrections
(such as Bonferroni ) can be overly conservative and
increase the risk of making a type II error. We therefore
present the results of null model analyses both with
unadjusted threshold significance levels (i.e., P , 0.05, P
, 0.01) and indicate those that remain significant after
ˇ ´ k adjustment (Quinn and Keough 2002)
the Dunn-Sida
of the P , 0.05 threshold to P , 0.0127. We used the
ˇ ´ k rather than the Bonferroni correction as
this approach slightly improves the power for each
comparison (Quinn and Keough 2002). However, with
four tests, the adjusted significance level is very similar
for the standard Bonferroni correction (0.0125 with
Bonferroni correction). Thus all null model analyses
originally significant at P , 0.01 remain significant after
adjustment (including all those on which our conclusions are based), while analyses significant only at P ,
0.05 are no longer significant. We present unadjusted
ˇ ´k
results, highlighting those for which the Dunn-Sida
adjustment of threshold significance levels has any
Objective 1: flower–visitor interactions are structured
in daily time
The June whole-day and time interval flower–visitor
networks are shown for TB in Fig. 1, and qualitatively
similar patterns are presented for JN in Fig. 2. Summary
statistics for these networks are provided in Table 1. We
recorded 174 links between 40 plant species and 98 insect
taxa across both sites. Bees were dominant flower
visitors (49% of recorded visits at TB and 81% at JN),
contributing more visits than other taxa in time intervals
2, 3, and 4 at both sites and in time interval 1 at JN
(Figs. 1 and 2). Bees were also the most species-rich
visitor taxon (48% of visitor species at TB and 54% at
JN) and were involved in the greatest number of links
(53% at TB and 62% at JN).
Visual comparisons of the time interval networks
show obvious temporal variation in flower–visitor
interactions (Figs. 1 and 2). Visit numbers, visitor
taxon richness, and link richness were all greater in

March 2011


FIG. 1. Quantitative flower visitation networks for the
Turkana Boma site at Mpala Research Centre, central Kenya in
June: (a) whole-day network (all data collected between 06:00
and 18:00), (b) time interval 1 (06:00–09:00), (c) time interval 2
(09:00–12:00), (d) time interval 3 (12:00–15:00), (e) time interval
4 (15:00–18:00). In each network the rectangles represent insect
taxa (top row) and plant species (bottom row), and the
connecting triangles represent links between taxa. The width
of the rectangle for each plant species represents the number of
floral units present in the plot. The width of the rectangle for
each visitor taxon represents the total number of visits to all
plants made, and the widths of the connecting lines represent
the number of visits observed for that link. Visitor taxa are
color-coded as follows: red, bees (Hymenoptera); medium blue,
wasps (Hymenoptera); light blue, flies (Diptera); dark blue,
beetles (Coleoptera); yellow, butterflies and moths (Lepidoptera). All networks are drawn to the same scale.

time intervals 2 (09:00–12:00) and 3 (12:00–15:00) than
in time intervals 1 (06:00–09:00) and 4 (15:00–18:00) at
both sites, with values consistently lowest in time
interval 1 (Table 1). The null model results show that
activity by visitor species is highly structured in daily
time: excluding time interval 1 at TB (and time interval
ˇ ´ k correction is applied), the
1 at JN when Dunn-Sida


FIG. 2. Quantitative flower visitation networks for the
Junction site at Mpala Research Centre, central Kenya in June:
(a) whole-day network (all data collected between 06:00 and
18:00), (b) time interval 1 (06:00–09:00), (c) time interval 2
(09:00–12:00), (d) time interval 3 (12:00–15:00), (e) time interval
4 (15:00–18:00). See legend to Fig. 1 for a full explanation of the
network diagrams.

numbers of links in each time interval were significantly lower than in randomly assembled networks at both
sites (Fig. 3). This reflects the fact that few visitor
species were observed in more than one time interval
(18% and 15% of species at TB and JN, respectively).
The observed temporal structuring is due predominantly to temporal patterns in bee activity: numbers of
links involving bees were consistently below null
expectations (Fig. 3). Of the other visitor groups,
butterflies and moths (TB) and wasps (JN) also showed
significant departures from null expectations (Fig. 3
and see Appendix B for taxa with nonsignificant
results). Very similar patterns and conclusions were
reached in analyses including lumped taxa (see
Appendix C).



Ecology, Vol. 92, No. 3

TABLE 1. Network summary statistics for the whole-day and time interval networks at the (a) Turkana Boma (TB) and (b)
Junction (JN) sites at Mpala Research Centre, central Kenya in June 2004.
Time interval
Network property

1 (06:00–09:00) 2 (09:00–12:00) 3 (12:00–15:00) 4 (15:00–18:00) Whole-day

a) TB June
Visitor taxa
Visitor species 
Plant species observed (P)
Plant species visited (Pv)
Flower visits
Flower–visitor links
Connectance, C (%)
Plant linkage
Visitor linkage
Unique linksà
Percentage of unique links§
Links missed if sampling restricted to interval
Percentage of links missed if sampling restricted
to interval






b) JN June
Visitor taxa
Visitor species 
Plant species observed (P)
Plant species visited (Pv)
Flower visits
Flower–visitor links
Connectance, C (%)
Plant linkage
Visitor linkage
Unique linksà
Percentage of unique links§
Links missed if sampling restricted to interval
Percentage of links missed if sampling restricted
to interval






  Visitor taxa identified to species or morphospecies.
à Defined as a link observed in no other time interval in the same whole-day network.
§ Calculated as a proportion of the total links per time interval.

Objective 2: daily temporal structure persists
through seasonal time
At both TB and JN, plant and visitor taxa in the
networks varied across seasonal time (full details of
flower–visitor links observed in all networks can be
found in Appendix D). Over the four months of
sampling, we recorded 267 links between 46 plant
species and 123 insect taxa at TB and 124 links between
30 plant species and 80 insect taxa at JN. Summary
statistics for all time interval and whole-day networks in
May, July, and August are provided in Appendix E. In
all months, patterns paralleled those seen in the June
networks: bees consistently made more visits than any
other taxon (40–71% of total visits) and were involved in
the greatest number of links (43–61%). Numbers of
visits, visitor taxa richness, and link richness were
highest in either time interval 2 or 3 and lowest in time
interval 1 in all networks (Table 1; Appendix E).
Activity by visitor species was structured in daily time
in all networks except the July network at TB (and the
ˇ ´ k correction is
July network at JN when Dunn-Sida
applied; Appendix F), due primarily to patterns in bee
activity; numbers of links involving bees were more
consistently below null expectations than for other

visitor groups (Fig. 3 and see Appendix G). Again, very
similar patterns were found in analyses including
lumped taxa (data not shown).
Objective 3: impact of observation time interval
on the network
Sampling of flower visitors was highly sensitive to the
timing of observation, with few species observed in
multiple time intervals. The most notable exception was
the honey bee, Apis mellifera, which was recorded in
time intervals 2–4 in three monthly networks (Appendix
H). In contrast, many plants were visited in more than
one time interval (43–78% at TB, 20–63% at JN), and a
notable minority were visited in at least three time
intervals (14–27% at TB, 31% at JN in June; see
Appendix H for examples). Thus, in most networks,
choice of recording time interval had a stronger impact
on the recorded species richness of visiting insects than
of visited plants. These specific patterns are reflected in
linkage summary statistics, which were consistently
higher for the overall network than for any constituent
time interval network in plants and visitors (Table 1;
Appendix E).

March 2011



FIG. 3. Comparisons between the number of flower–visitor links actually observed in each time interval (white) and the mean
(695% CI) observed in 1000 randomized networks (gray) for visitors identified to species in the Turkana Boma (TB) and Junction
(JN) June networks: (a) all visitor species at TB, (b) all visitor species at JN, (c) bees at TB, (d) bees at JN, (e) butterflies and moths
at TB, and (f ) wasps at JN. Significant differences between observed values and null expectations are indicated by asterisks. Results
ˇ ´ k correction of the threshold P , 0.05 for multiple tests. Equivalent
significant at P , 0.01 remain significant with Dunn-Sida
figures for nonsignificant taxa are shown in Appendix B.
* P , 0.05; ** P , 0.01 (non-adjusted significance levels).

All time intervals in each whole-day network contained unique links that were not observed in any other
time interval in the same network (Table 1; Appendix E).
Time intervals 2 and 3 contained the greatest numbers of
unique links in all monthly networks. However, consideration of unique links as a proportion of the total
number of links in each time interval shows them to form
a high proportion of total links in almost all time
intervals in each network (65–100%, see Appendix E).
Exclusion of any time interval during sampling will thus
miss specific links. To investigate the effect of sampling
over a shorter daily time window, we calculated the

proportion of links that would be missed if sampling was
restricted to a single time interval (Table 1; Appendix E).
At least 45% of links from the whole-day network would
be missed if sampling was restricted to any single 3-h
time interval (Appendix E), with the impact minimized if
sampling is restricted to time intervals 2 or 3 as these
contained greater numbers of unique links.
Our results show that daily timing of data collection
has a significant impact on inferred structures of flower
visitation networks for these savanna communities.



With the three-hour interval resolution used here, most
flower–visitor links were restricted to a single time
interval. Visit frequency and link richness were both
highest in the middle of the day, although lower rates of
visitation earlier and later in the day both contributed
novel links to the networks. This temporal structuring
meant that restriction of observations to any single
three-hour time interval resulted in failure to detect a
large proportion of links observed through the day
(Table 1; Appendix E). We first consider the limitations
of our approach, then the causes and consequences of
temporal structure both in pollination networks and
more generally in other networks. We end by addressing
the implications of our results for pollination ecologists
with regard to both sampling networks in the field and
analyzing multiple data sets gathered from the literature.
There are three limitations to our approach. First, we
could not generate high-resolution network data for
nocturnal flower visitation, when nocturnal pollinators
such as moths and bats would have been active. This
decision was driven by safety considerations and does
not weaken our conclusion that timing of data collection
matters in network construction. It is already well
known that plant–pollinator interactions are temporally
structured by night and day (Fenster et al. 2004) and
where nocturnal visitation is substantial there will be an
even greater need to incorporate all 24 hours of the
diurnal cycle into an appropriate sampling strategy if the
causes and consequences of network structure are to be
understood. To our knowledge, only one network study
(Devoto et al., in press) has considered nocturnal plant–
pollinator interactions (but see Clinebell et al. [2004] for
a multispecies approach incorporating 24-h sampling).
Second, we combined laboratory and field identification
of flower visitors. A drawback of laboratory identification of captured specimens is that they cannot return to
make future visits. Other studies have removed this bias
by catching all insects (e.g., Lopezaraiza-Mikel et al.
2007, Forup et al. 2008), but we argue that to do so
would have caused local extinction of some rare flower
visitors in our networks. Finally, although all problematic species were morphotyped by taxonomists, identification to species was not possible in all cases because
the insect fauna of Kenya is still not completely
described. The lack of any contrast between analyses
using specimens identified to species and those including
lumped morphotaxa suggests that uncertainty in species
identities has not significantly influenced our results or
associated inference.
Extent and potential causes of observed daily
temporal structure
Our null modeling approach revealed nonrandom
distribution of specific flower–visitor links through daily
time in almost all of our seasonal networks, a result to
be expected from the large literature on pollinator

Ecology, Vol. 92, No. 3

activity patterns (e.g., Stone et al. 1988, Willmer and
Stone 2004) but rarely explicitly considered in the
network literature. The impact of daily time on network
structure was consistent across two sites differing in
floral community over six of seven seasonal data sets,
suggesting that it is a consistent feature at least of
savanna flower–visitor interaction networks. We do not
suggest that network data collection strategies are the
best way to demonstrate such structure, but it is clear
that it can be demonstrated in such data and should
therefore be acknowledged in sampling for interaction
networks. Bees showed the most consistent evidence of
daily temporal structure among forager taxa, consistent
with the dependence of female reproductive success on
both thermal physiology and floral resource availability
(Herrera 1990, 1995, Stone 1994, Stone et al. 1998,
Willmer and Stone 2004). The strength of the patterns in
bees may also reflect their dominance in our data, as in
many other flower–visitor networks (e.g., Memmott and
Waser 2002, Forup et al. 2008). More detailed analyses
of daily activity patterns in non-bee visitors would assist
our understanding of this observed contrast between
bees and other taxa.
Understanding the causes of temporal structure
requires understanding the relative contributions of
sampling error and the top-down and bottom-up
influences driving the timing of flower–visitor interactions. Revealing temporal structure imposed by topdown aspects of visitor biology requires finer resolution
activity pattern and behavioral data than collected in
this study (e.g., Stone et al. 1995, 1998), but our data do
reveal bottom-up constraints imposed by floral resources. Floral imposition of bottom-up control is sometimes
very obvious: in our networks flowers of the morning
glory Ipomoea kituiensis (Convolvulaceae) opened from
08:00 and were shriveled by noon, while flowers of Sida
ovata and S. schimperiana (Malvaceae) opened only
from 11:00 to 15:00. For these plants, visitation outside
these times is impossible, representing a daily temporal
equivalent of forbidden links sensu Jordano et al. (2003).
In other species, however, the timing of resource
provision is not apparent externally. For example,
pollen release in open flowers of the acacia Senegalia
(Acacia) brevispica (Fabaceae) at Mpala takes place
around noon each day (Baldock 2007), a process that is
clearly reflected in shifts in the visitors with links to this
species. Prior to pollen release, most visitation is by
flower-feeding beetles and nectar feeders, while after
pollen release most vists are by pollen-collecting bees
(Appendix H). As a result, the sets of links involving S.
brevispica before and after pollen release are very
different. (See also Stone et al. [1996, 1998] for similar
patterns for a set of acacia species in Tanzania.) Timing
of pollen release in acacias and other plants is sensitive
to microclimate and can vary at the same site between
days, with knock-on effects on the activity of associated
pollinators (e.g., Stone et al. 1998, 1999, Raine et al.
2007). Which component of the visitor community for

March 2011


S. brevispica is detected will thus depend on when it is
observed. Examples of bottom-up structuring are also
apparent in many temperate communities, particularly
in bees (Willmer and Stone 2004). Examples include
crepuscular or nocturnal visitation of evening primroses
following late afternoon anthesis (Moody-Weis and
Heywood 2001) and tracking of pollen availability
within and among plant species by Anthophora solitary
bees (Stone 1994, Stone et al. 1999).
Repeatability of patterns across years and seasonal
temporal patterns
Though we generated detailed network data for one
year only, data collected in subsequent years suggest
that daily temporal structure is likely to persist across
years as well as across seasons and sites. We continued
to collect monthly data on floral abundance using the
same protocol for the TB and JN sites throughout 2005
and 2006 and found similar groups of plant species to
coflower in a given month over all three years (2004–
2006; Appendix A). Though equivalent community-level
visitation data are not available for subsequent years,
data collected in TB plot in June 2006 for 12 of the
flowering species sampled in the June 2004 network
support the conclusions based on the 2004 data (217
hours of observation; see Appendix I). Bees dominated
numbers of visits and flower–visitor links as in 2004, and
a high proportion (59%) of flower–visitor links occurred
in only one of the four time intervals. Null model
analysis (Appendix I) also revealed evidence of daily
temporal structure in the 2006 data, with the numbers of
bee and lepidopteran flower–visitor links significantly
lower than null expectations for at least two time
intervals. The repetition of these patterns across years is
consistent with their generation by fundamental aspects
of floral and visitor biology, as we suggest above.
Our seasonal data (Appendix D; to be analyzed in
detail elsewhere) show that in addition to the daily
temporal patterns discussed in detail below there is also
seasonal variation in the species and links involved.
Seasonal variation means that the plants interacting via
shared pollinators, positively or negatively, change over
time. Summing the likely selective impact of such
interactions thus requires incorporation of both seasonal
and daily timescales (Rathcke 1983, Stone et al. 1998).
Consequences of daily temporal structure
for network studies
Where detailed information on the timing of dehiscence and release of floral rewards exists, it is possible to
quantify biologically relevant visitation (i.e., by pollen
vectors) in appropriate limited daily time intervals (e.g.,
Armbruster and Herzig 1984, Stone et al. 1998, 1999,
Willmer and Stone 2004). However, few pollination
network studies sample visitation with such knowledge,
beyond the obvious constraints imposed by floral
opening and closure (as for our Ipomoea and Sida
examples). Sampling of visitation for a limited daily time


window without such information runs the risk of
recording visitation at times when floral rewards are
absent or outside periods when pollen can be harvested
or deposited on receptive stigmas. Visits recorded at
these times may miss pollen vectors altogether. Where
network studies have sampled for only a small portion
of the day in the absence of specific information on the
plants involved, we cannot assume that recorded links
constitute those associated with pollen dispersal and
seed set, with obvious implications for ecosystem service
conclusions made from such networks. In our study
failure to record visitation in any time interval would
omit a subset of link types. Unsurprisingly, the impact is
greatest for the two intervals during the middle of the
day, when visit frequency and visitor richness are
greatest. The numbers of interval-specific links are
lowest for the first and last daily time intervals (Table
1; Appendix E), but ignoring these two intervals would
still result in failure to detect certain links.
Studies that do sample across a broad daily time
window still risk misinterpretation of the relationships
among plants and pollinators if they do not consider the
existence of daily temporal structure. For example, Apis
mellifera visited four plant species at JN in June:
Ipomoea kituiensis (Convolvulaceae) in time interval 2,
Emilia discifolia (Asteraceae) and Hypoestes forskahlii
(Acanthaceae) in time interval 3, and Leucas glabrata
(Lamiaceae) in time interval 4 (Appendix H). The
summation of these data over a daily timescale could
lead to the conclusion that these four plant species are
competing for visits from the same pollinator and are at
risk from interspecific pollen transfer. However, our
data show that A. mellifera visited different plant species
at different times of day, as observed for honey bees in
other studies (e.g., Stone et al. 1996, 1998). Given that
honey bees regularly remove pollen from their bodies,
these plants could instead potentially facilitate each
other’s pollination (Rathcke 1983, Moeller 2004).
Daily temporal structure in species interactions could
have consequences for other types of ecological networks. The extent to which daily temporal variation is
important will depend on the timescale over which
interactions occur and whether species have daily
activity patterns that restrict the times at which they
are available to interact. For example, interactions
between parasitoids and their hosts are unlikely to be
structured in daily time since parasitism takes place over
hours or days. Predator–prey interactions, including
those between mutualistic ant guards and other insects,
are more likely to be structured in daily time since
interactions occur over short time periods and both
interacting species can have specific daily activity
patterns (Willmer and Stone 1997, Raine et al. 2002,
Kronfeld-Schor and Dayan 2003). Hori and Noda’s
(2001) study of intertidal rocky shore food webs
provides an elegant example of the impact of daily
variation in the physical environment on taxa involved
in trophic links, with major shifts between periods



during which the shore is either submerged or exposed.
However, while rapid pollen removal means that the
distinction between competition for pollination and
facilitation mediated by shared pollinators can be
influenced by daily time, interactions in predator–prey
networks have longer-term dynamics.
Implications for sampling plant–pollinator networks
We propose that sampling for plant–pollinator
interaction networks should ideally target all daily time
intervals in which flowers of each component species are
open. This approach should guarantee sampling of
pollen vectors among the general visitor assemblage,
although subsequent analyses (for example, of correlations between visitor activity and timing of floral reward
provision and stigma sensitivity) are likely to be
necessary to identify key interactions for the species
involved. While this sets the ideal, compromises do have
to be made in many sampling programs. For example, it
would be difficult to sample multiple distant sites over
entire field seasons using this approach. It is possible
though to always clearly state the timing of data
collection. Moreover, sampling can be rotated through
different time intervals at each field site in order to
reduce bias. For example, Henson et al. (2009) rotated
sampling at 12 field sites through three daily time
intervals over a four-month field season. However, care
still needs to be taken to check that the factors
structuring daily activity patterns (such as ambient
temperature or timing of floral resource provision) are
the same across field sites.
Concluding remarks
This study shows the impact of daily temporal
structure in flower–visitor interactions on network
structure for two communities in an African savanna
habitat. Revealing the extent to which top-down and
bottom-up effects contribute to this structure requires
further detailed study of floral reward provision and
daily activity patterns of visitor taxa. We recommend
that future plant–pollinator network studies employ
explicitly time-structured sampling to maximize the
probability that interactions central to the biology of
the taxa involved are appropriately observed. The
benefits of this approach would be threefold: (1) it
would improve our understanding of the factors
structuring pollination networks; (2) it would allow us
to better appreciate the impact of environmental change
on pollination networks, and (3) it would improve our
ability to restore damaged pollination networks.
We thank John Deeming, Connal Eardley, George Else,
David Greathead, Josef Gusenleitner, Michael Kuhlmann,
Alain Pauly, Adrian Pont, Woijiech Pulawski, Raymond Wahis,
and Andrew Whittington for taxonomic expertise, and the
Natural History Museum, London, and the National Museums
of Kenya for access to collections. We are grateful to Mpala
Research Centre and Wanja Kinuthia at the National Museums
of Kenya for logistical support. We thank Raphael Erengai,

Ecology, Vol. 92, No. 3

Patrick Lenguya, Pat Willmer, Andrew Schnabel, Adriana
Otero Arnaiz, and Anna Watson for additional support in data
collection and field identification. We are grateful to Andrew
Schnabel and three anonymous reviewers for constructive
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NERC quota studentship to the University of Edinburgh
(K. C. R. Baldock) and by U.S. National Science Foundation
grant number DEB-0344519 (G. N. Stone). We thank the
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bees. Advances in the Study of Behavior 34:347–466.

Flowering plant species present at the two sites during network sampling in 2004 and for equivalent months in 2005 and 2006
(Ecological Archives E092-057-A1).

Comparisons of numbers of flower–visitor links per time interval between observed networks and null model results for visitor
species with no significant differences between observed values and null expectations at both study sites in June 2004 (Ecological
Archives E092-057-A2).

Comparisons of numbers of flower–visitor links per time interval between observed networks and null model results for visitor
taxa at both study sites in June 2004 (Ecological Archives E092-057-A3).

Flower–visitor links recorded for each monthly network (Ecological Archives E092-057-A4).

Network summary statistics for each monthly network (Ecological Archives E092-057-A5).

Comparisons of numbers of flower–visitor links per time interval between observed networks and null model results for all
visitor species combined for each monthly network (Ecological Archives E092-057-A6).

Comparisons of numbers of flower–visitor links per time interval between observed networks and null model results for each
visitor taxon with significant results across all monthly networks (Ecological Archives E092-057-A7).

Examples of variation across time intervals in the links involving specific taxa (Ecological Archives E092-057-A8).

Results of null model analyses of flower–visitor data from Turkana Boma site in 2006 (Ecological Archives E092-057-A9).