Freshwater Biology (2015)
doi:10.1111/fwb.12668
The first to arrive and the last to leave: colonisation and
extinction dynamics of common and rare fishes in
intermittent prairie streams
JAMES E. WHITNEY*,1, KEITH B. GIDO*, ERIKA C. MARTIN* AND KRISTEN J. HASE†
*Division of Biology, Kansas State University, Manhattan, KS U.S.A.
†
Tallgrass Prairie National Preserve, National Park Service, Strong City, KS U.S.A.
SUMMARY
1. The objectives of our research were to examine commonness–rarity patterns in fish communities in
networks of intermittent streams. We quantified species abundance distributions and the importance
of nestedness and turnover to community dissimilarity and then related commonness to colonisation,
extinction and physiological tolerance. Patterns and relationships were evaluated spatially among
sites and temporally within sites during non-drought and drought periods in tallgrass prairie
streams of eastern Kansas, U.S.A.
2. Supra-seasonal drought during 2011–2013 resulted in complete or partial drying of some sites and
provided an opportunity to evaluate whether commonness was predictive of rather than predicted
by colonisation and extinction. Abundance was used to predict re-colonisation in desiccated reaches
and persistence in drying pools.
3. Few species were common, while most were rare regardless of drought, and nestedness drove
community dissimilarity across sites. Common species had higher colonisation and lower extinction
than rarer species, but physiological tolerance was unrelated to commonness.
4. Abundant species were generally the first to re-colonise desiccated reaches, but pre-drought abundance did not predict persistence in partially desiccated reaches.
5. Although common species were the first to colonise and the last to go extinct, we were unable to
determine whether commonness was predictive of rather than predicted by colonisation and extinction. Regardless, our study demonstrates linkages among commonness, colonisation and extinction.
Keywords: drought, metapopulation dynamics, nestedness, species abundance distributions (SADs), turnover
Introduction
A universal pattern in the distribution of species’ abundances across space and time is that a few species are
abundant, while most are rare (Preston, 1948; McGill
et al., 2007; Ulrich, Ollik & Ugland, 2010). Abundant species also tend to have a higher occurrence throughout
space and time (Hanski, Kouki & Halkka, 1993; Gaston,
Blackburn & Lawton, 1997), suggesting that abundance,
occurrence and persistence are linked in an overall phenomenon of commonness (abundant, widespread and
persistent) and rarity (scarce, restricted and infrequent)
(Brown, 1984; Hanski & Gyllenberg, 1993; Magurran &
Henderson, 2003; Magurran, 2007). Identifying mechanisms explaining the commonness–rarity phenomenon
of species abundance distributions (SADs) has long been
a goal of community ecology (Preston, 1948; Williams,
1953) and is still an active area of ecological research
(Matthews, Borges & Whittaker, 2014; Matthews & Whittaker, 2014).
Stream fish communities exhibit the commonness–rarity phenomenon (Sheldon, 1987). Furthermore, in riverine
Correspondence: James E. Whitney, Missouri Cooperative Fish and Wildlife Research Unit, Department of Fisheries and Wildlife Sciences,
University of Missouri, 302 ABNR Building, Columbia, Missouri 65211, U.S.A. E-mail: whitneyj@missouri.edu
1
Present address: Missouri Cooperative Fish and Wildlife Research Unit, Department of Fisheries and Wildlife Sciences, University of
Missouri, Columbia, MO U.S.A.
© 2015 John Wiley & Sons Ltd
1
2
J. E. Whitney et al.
networks composed of intermittent (i.e. rivers that periodically cease to flow) and perennial reaches, common
fishes often occur throughout intermittent or ephemeral
(i.e. streams that dry completely) tributaries and perennial mainstem sections, while rare species are restricted
to perennial reaches, resulting in downstream increases
in species richness (Roberts & Hitt, 2010). This longitudinal pattern of species addition is described as nested,
wherein species-poor headwater communities are a subset of species-rich downstream communities (Patterson
& Atmar, 1986). Turnover (replacement of one species
by another species; Baselga, 2010) is predicted to be a
less common process promoting community dissimilarity in intermittent–perennial stream networks.
Mechanisms explaining nestedness, and by extension
commonness, in intermittent streams can be divided
into two non-mutually exclusive hypotheses. The first
hypothesis predicts that common species possess high
dispersal propensity and can thus colonise a greater
number of habitats leading to a larger overall population size and range (dispersal limitation hypothesis;
Albanese, Angermeier & Peterson, 2009). The second
hypothesis predicts that common species possess high
tolerance of harsh environmental conditions and can
thus persist throughout a greater range of habitats
leading to lower extinction and a larger overall population size and range (environmental tolerance hypothesis; Schlosser, 1990; Townsend & Hildrew, 1994; Taylor
& Warren, 2001). A conceptual model proposed by
Larned et al. (2010) integrated these hypotheses, stating
that longitudinal patterns of nestedness across connected stream habitats were caused by dispersal limitation, whereas temporal patterns of nestedness within
habitats during stream drying were caused by interspecies differences in environmental tolerance. The
degree of stream intermittency creates opportunities to
test these hypotheses, as species that persist in drying
pools would be predicted to have greater tolerance of
high temperatures and low dissolved oxygen than
those that do not (Matthews & Styron, 1981; Smale &
Rabeni, 1995a,b), whereas species that re-colonise fully
desiccated ephemeral reaches would be predicted to
have greater dispersal propensity than those that do
not (Albanese et al., 2009). Periodic or supra-seasonal
drought can enhance stream intermittency and physicochemical stress (Lake, 2003; Magoulick & Kobza, 2003),
thus creating further opportunities for testing these
hypotheses.
The commonness, tolerance and re-colonisation potential of species can be described by three metapopulation
probabilities: probability of occupancy, colonisation and
extinction (Gotelli & Taylor, 1999a,b). A common species
occurs throughout a large number of samples and therefore would have a high probability of occupancy. Species with greater dispersal propensity are expected to
have higher probability of colonisation, and species with
high physiological tolerance are predicted to have lower
extinction probabilities. Therefore, these probabilities can
be used to test the dispersal limitation and environmental tolerance hypotheses. However, colonisation and
extinction may be predicted by commonness, as populations with greater local abundance and regional distribution are less likely to go extinct (Richter-Dyn & Goel,
1972; Lima, Marquet & Jaksic, 1996; Gotelli & Taylor,
1999b; Taylor & Warren, 2001) and are more likely to
provide colonists (Sheldon & Meffe, 1995; Stoll et al.,
2014).
The objective of this research was to understand how
species’ metapopulation dynamics and tolerance influence commonness in two intermittent tallgrass prairie
stream networks during a drought and non-drought period. Specifically, we first examined commonness–rarity
patterns through space and time across sites positioned
along a stream size gradient. We predicted that sites
would always exhibit the commonness–rarity pattern of
a few abundant and many rare species, but that this pattern would weaken during drought as rare species were
extirpated by harsh environmental conditions. Next, we
sought to determine the contributions of nestedness and
turnover to overall community dissimilarity among and
within sites. We predicted that variation in community
structure would be driven more by nestedness than turnover and that the contribution of nestedness to community dissimilarity would increase during drought as the
community diverged to a subset of tolerant species. Our
next aim was to quantify relationships of commonness
(abundance or occurrence) with metapopulation dynamics and species tolerance to assess the validity of dispersal limitation and environmental tolerance pathways. We
predicted that commonness would be positively related
to colonisation and unrelated to extinction during the
non-drought period, but would be negatively related to
extinction during drought because common species are
predicted to be more tolerant of harsh conditions.
Finally, we used data from desiccated reaches to determine whether catchment abundance was predictive of
re-colonisation and data from drying habitats to test if
local abundance was associated with persistence. We
predicted that abundance would not influence re-colonisation of desiccated reaches or persistence in drying
habitats and that species with higher physiological tolerance would be the last to go extinct from drying habitats.
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
Colonisation, extinction, and commonness of stream fishes
Methods
3
2013, with the exception being that TPNP-2 was not
sampled in August of 2008.
Study area
This study was conducted in stream networks located
on Tallgrass Prairie National Preserve (TPNP) and
Konza Prairie Biological Station (KPBS) in the Flint Hills
region of east-central Kansas, U.S.A. (Fig. 1). Because of
their location in protected reserves, streams in both
catchments were relatively pristine. Tallgrass prairie is
the dominant land cover in both catchments, with gallery forest in riparian areas in lower sections. Grazing
by cattle and/or bison occurs in both catchments, with
some row-crop agriculture also present in lower reaches.
Sites were selected along a gradient of increasing stream
size and permanency (Table 1). Four sites were chosen
on TPNP along an 8.36 river kilometre, 32 m elevational
gradient, and three sites along a 4.16 river kilometre,
56 m elevational gradient, were selected on KPBS
(Fig. 1). Sites were sampled three times (May, August,
November) annually between May 2008 and November
Non-drought and drought study periods
Seasonal drought from a lack of mid- to late summer
(July–September) precipitation is common in both catchments, but these droughts generally do not result in
complete stream drying. Supra-seasonal drought that
occurred in the region during 2011–2013 (Fig. S1)
resulted in complete stream drying, with drying first
occurring in headwater regions and then progressing
downstream (i.e. headwater drying sensu Lake, 2003).
This headwater drying resulted in complete desiccation
(ephemeral) of TPNP-1 and TPNP-2, whereas all other
sites on TPNP and KPBS retained water but ceased to
flow (intermittent). Samples collected during 2008-2010
were assigned to a non-drought period, and samples
taken during 2011–2013 were assigned to a drought period.
Fish sampling
Fishes were sampled in one to four mesohabitats (riffle
and/or pool) at each site using a Smith-Root LR-24 or
LR-20B (Vancouver, WA, U.S.A.) backpack electrofisher
with two dip netters (3.2-mm mesh nets) except in
TPNP-2. In addition to electrofishing, pools in sites 2–4
on the TPNP were seined in a downstream direction
using a 4.57 m 9 1.83 m seine with 3.2-mm mesh. Seining was rarely conducted in KPBS sites because a previous study found electrofishing to be adequate for fish
sampling (Bertrand, Gido & Guy, 2006). All fishes collected were identified to species, measured for total
length (mm) and released. The number caught was used
to describe species abundance, with species abundance
summed across sites during wet and dry periods to
describe among-site spatial patterns, and summed across
sample periods within sites to describe within-site temporal patterns.
Metapopulation probabilities
Fig. 1 Map of study sites on Konza Prairie Biological Station
(KPBS) and Tallgrass Prairie National Preserve (TPNP). Sites were
numbered in order of increasing stream size and permanence.
Light grey shading indicates the Kansas River Basin, whereas dark
grey shading indicates the Neosho River Basin. TPNP sites 1 and 2
are ephemeral (dried completely), whereas TPNP sites 3 and 4 and
all KPBS sites are intermittent (ceased to flow).
Metapopulation probabilities were calculated for each
fish species following the methods of Gotelli & Taylor
(1999a,b) and included probability of occupancy (po:
number of sites occupied/total number of sites), colonisation (pc: number of colonisation events/number of
colonisation opportunities) and extinction (pe: number of
extinction events/number of extinction opportunities).
A colonisation event was defined as a site going from
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
4
J. E. Whitney et al.
Table 1 Physical and hydrological attributes of study sites on Tallgrass Prairie National Preserve (TPNP) and Konza Prairie Biological Station (KPBS). Mean values are averaged across mesohabitats and sample periods with the range in parentheses
Catchment
Site
TPNP
Site
Site
Site
Site
Site
Site
Site
KPBS
1
2
3
4
l
2
3
Mean width
(m)
Mean depth
(m)
Mean site area
(m2)
Catchment
area (km2)
Completely
dry 2011
Completely
dry 2012
1.7
5.3
5.2
5.7
1.8
3.1
4.6
0.16
0.61
0.22
0.36
0.12
0.15
0.19
134
189
522
821
110
282
616
5.7
4.3
15.4
88.4
1.2
3.1
17.5
Yes
No
No
No
No
No
No
Yes
Yes
No
No
No
No
No
(0.0–2.3)
(0.0–5.9)
(3.1–6.7)
(3.6–8.6)
(1.2–3.0)
(1.8–4.3)
(2.9–6.5)
(0.00–0.24)
(0.00–0.81)
(0.05–0.37)
(0.28–0.59)
(0.03–0.16)
(0.03–0.22)
(0.15–0.30)
unoccupied to occupied between two consecutive time
periods, and an extinction event occurred when a site
went from occupied to unoccupied between two consecutive time periods. Extinction defined in this framework
is the disappearance of a species from a locality, and it
does not refer to global extinction. Colonisation and
extinction opportunities were the number of times a site
was unoccupied or occupied, respectively. Species
metapopulations probabilities were calculated across
sites and within sites during the wet and dry periods in
each catchment.
Data analysis
Statistical analyses were performed in R version 3.1.3 (R
Core Team 2015). Results were considered statistically
significant at a ≤ 0.05 and marginally significant at
a ≤ 0.10. Because of biogeographic differences in fish
communities, the TPNP and KPBS communities were
analysed separately and they represent independent
characterisations of commonness–rarity patterns and
independent tests of the dispersal limitation and environmental tolerance hypotheses in intermittent tallgrass
prairie streams. To verify that our study catchments
exhibited the commonness–rarity pattern, we examined
species abundances distributions using Whittaker plots
(McGill et al., 2007; Ulrich et al., 2010), which predict a
linear decrease in log10 (abundance) with increasing species rank for the commonness–rarity pattern. To assess
the significance of Whittaker plots, we conducted
Shapiro–Wilk’s test of normality on log10(x)-transformed
species abundance data, with larger P-values (i.e. stronger conformance to log-normal distribution; H0 = distribution is log-normal) indicating greater support for the
commonness–rarity pattern. To assess the contribution of
nestedness and turnover to overall community dissimilarity, we used the ‘betapart’ package, with Simpson’s
index chosen to describe turnover, the nestedness component of Sørensen’s index used to describe nestedness and
(0.00–422)
(0.00–228)
(117–711)
(381–1515)
(72–210)
(71–420)
(387–909)
Sørensen’s index chosen to describe overall community
dissimilarity (Baselga & Orme, 2012). We acknowledge
that our limited spatial replication (≤4 sites) may have
hindered our ability to detect nestedness and its contribution to community dissimilarity, but the simple
network architecture of our study catchments meant that
including more sites would be redundant and would not
add more information. Study sites encompassed most
tributary and mainstem habitats within catchments,
and should accurately reflect community dissimilarity
patterns.
We tested the dispersal limitation and environmental
tolerance hypotheses using simple linear regression.
Hypotheses were tested spatially across sites and temporally within sites during non-drought and drought periods. Species served as replicates in all analyses, with
species abundance and occupancy used as response
variables to describe commonness. Using species as
replicates resulted in low statistical power in depauperate sites, so results were interpreted with caution. Predictor variables included probability of colonisation for
the dispersal limitation hypothesis and probability of
extinction for the environmental tolerance hypothesis.
To ensure that significant negative relationships between
extinction and commonness were driven by physiological tolerance, we calculated relationships between commonness and hypoxia and hyperthermia tolerance
values reported in the study by Smale & Rabeni (1995a),
which included values for 21 species captured on TPNP
and KPBS. Although these tolerances were quantified
for fish from Missouri, tolerances should be similar for
Kansas fishes given populations are residing at a similar
latitude (isotherm) and in similar habitats (Smale &
Rabeni, 1995a,b). Because we did not have tolerance
information for all species, degrees of freedom for relationships between tolerance and commonness were
sometimes less than relationships between metapopulation probabilities and commonness, which may compromise our ability to detect an effect of tolerance on
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
Colonisation, extinction, and commonness of stream fishes
commonness. To linearise relationships, all abundance
values were log10(x + 1)-transformed and all metapopulation probabilities were logit-transformed (Warton &
Hui, 2011).
Our analyses assumed that species were independent
replicates, which could bias results if abundance and
occupancy are phylogenetically conserved (Pagel, 1992;
Fisher & Owens, 2004). To assess the phylogenetic
dependence of our response variables, we followed the
semi-quantitative methods of Tedesco et al. (2008) and
Olden & Kennard (2010), which was preferable to other
methods (e.g. independent contrasts; Felsenstein, 1985)
given that a complete phylogeny with branch lengths
was not available. This method assigns phylogenetic
dissimilarities among species according to taxonomy
(different order = 0.75; same order = 0.67; same family = 0.50; same genus = 0.00) and then uses a Mantel
test to assess relationships between phylogenetic dissimilarity and the Euclidean dissimilarity of responses
variables. Because we determined that log10 (abundance)
on TPNP (r = 0.08; P = 0.89) and KPBS (r = 0.01;
P = 0.52) and logit (occupancy) on TPNP (r = 0.07;
P = 0.85) and KPBS (r = 0.06; P = 0.67) were phylogenetically unrelated, we reasoned that species could serve
as independent replicates in our analyses.
Within-site case studies
The supra-seasonal drought that occurred during 2011–
2013 provided an opportunity to further assess relationships among abundance, tolerance and dispersal propensity using within-site case studies. Drought resulted in
complete drying at TPNP-1 in 2011 (drying event 1) and
again in 2012 (drying event 2), and at TPNP-2 in 2012.
Complete drying and re-wetting occurred in September
and November, respectively, in 2011 and August and
November, respectively, in 2012. To ascertain whether
abundance was predictive of colonisation, observed recolonisation at desiccated sites was regressed against
catchment-wide abundance during each re-colonisation
period (i.e. each sample month following re-wetting).
Catchment-wide abundance was averaged across all
sites within a catchment except the focal re-colonisation
site, with averaging performed between the sampling
month corresponding to observed re-colonisation and
the previous sample month. Mean catchment-wide
abundance was weighted by the shortest watercourse
distance of a site to each potential source stream, which
was calculated using Google Earth â. Only those species
that were present at a site before the drought or those
that colonised following the drought were used in
5
analyses. Relationships between re-colonisation and
catchment-wide abundance were assessed using logistic
regression, with significance evaluated using a chisquare test statistic.
To investigate how environmental tolerance and abundance influenced persistence during drought, we compared hyperthermia tolerance, hypoxia tolerance and
pre-drought abundance of species that persisted through
a drying event at TPNP-3 to those that did not. The drying event began in May 2012 (pool area = 482 m2),
peaked in August 2012 (pool area = 117 m2) and subsided in November 2012 (pool area = 490 m2). We used
t-tests to examine whether species that persisted through
this drying event had higher hyperthermia and lower
hypoxia tolerances than those that were present prior to
drought conditions in November 2011 and May 2012 but
were absent in November 2012. Finally, to quantify
whether abundance predicted persistence, we compared
pre-drought abundance averaged between November
2011 and May 2012 between species that persisted versus those that did not using a t-test.
Results
Among-site patterns and relationships
We collected 35 139 individuals of 29 species at TPNP
and 24 608 individuals of 12 species at KPBS, for a grand
total of 59 747 individuals and 34 species (Table 2). As
predicted, both the TPNP and the KPBS exhibited SADs
consistent with the commonness–rarity pattern (i.e.
Shapiro–Wilk’s P > 0.16) during non-drought (wet) and
drought (dry) periods (Fig. 2). However, the strength of
these patterns increased during drought on TPNP and
KPBS, as indicated by increases in log-normal P-values
(Table 3). Nestedness was always more important than
turnover in its contribution to among-site community dissimilarity, with this importance increasing during the
drought period as expected (Table 3). In fact, nestedness
was entirely responsible for among-site community dissimilarity during wet or dry periods on KPBS, as there
was no among-site species turnover.
Species abundance and catchment occurrence were
positively related to colonisation during wet and dry
periods on TPNP and KPBS, providing initial support
for the dispersal limitation hypothesis (Table 4; Figs 3 &
4). Our observations did not support the environmental
tolerance hypothesis because abundance and occupancy
were always negatively related to extinction rather
than only during drought. Moreover, the lower extinction of common species did not result from greater
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
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J. E. Whitney et al.
Table 2 Abundance of fish species encountered at Tallgrass Prairie National Preserve (TPNP) and Konza Prairie Biological Station (KPBS)
during wet (2008–2010) and dry (2011–2013) periods. Species are arranged in order of decreasing total abundance per catchment. Species
codes are the first three letters of the genus and the first three letters of the specific epithet, with the number in parentheses preceding the
code indicating total rank in a catchment
Wet
Dry
Location
Common name
Scientific name
Species code
Site
1
Site
2
Site
3
Site
4
Site
1
Site
2
Site
3
Site
4
Total
TPNP
Central stoneroller
Orangethroat darter
Bluntnose minnow
Cardinal shiner
Redfin shiner
Green sunfish
Orangespotted sunfish
Longear sunfish
Red shiner
Western mosquitofish
Black bullhead
Golden redhorse
Creek chub
Brook silverside
Bluegill
Sand shiner
Topeka shiner
Blackstripe topminnow
Logperch
Yellow bullhead
Largemouth bass
Stonecat
Suckermouth minnow
Spotted bass
Spotted sucker
Channel catfish
Slim minnow
Golden shiner
Bullhead minnow
Southern redbelly dace
Central stoneroller
Orangethroat darter
Creek chub
White sucker
Johnny darter
Green sunfish
Slender madtom
Common shiner
Red shiner
Longear sunfish
Western mosquitofish
Campostoma anomalum
Etheostoma spectabile
Pimephales notatus
Luxilus cardinalis
Lythrurus umbratilis
Lepomis cyanellus
Lepomis humilis
Lepomis megalotis
Cyprinella lutrensis
Gambusia affinis
Ameiurus melas
Moxostoma erythrurum
Semotilus atromaculatus
Labidesthes sicculus
Lepomis macrochirus
Notropis stramineus
Notropis topeka
Fundulus notatus
Percina caprodes
Ameiurus natalis
Micropterus salmoides
Noturus flavus
Phenacobius mirabilis
Micropterus punctulatus
Minytrema melanops
Ictalurus punctatus
Pimephales tenellus
Notemigonus crysoleucas
Pimephales vigilax
Chrosomus erythrogaster
Campostoma anomalum
Etheostoma spectabile
Semotilus atromaculatus
Catostomus commersonii
Etheostoma nigrum
Lepomis cyanellus
Noturus exilis
Luxilus cornutus
Cyprinella lutrensis
Lepomis megalotis
Gambusia affinis
(1)CAMANO
(2)ETHSPE
(3)PIMNOT
(4)LUXCAR
(5)LYTUMB
(6)LEPCYA
(7)LEPHUM
(8)LEPMEG
(9)CYPLUT
(10)GAMAFF
(11)AMEMEL
(12)MOXERY
(13)SEMATR
(14)LABSIC
(15)LEPMAC
(16)NOTSTR
(17)NOTTOP
(18)FUNNOT
(19)PERCAP
(20)AMENAT
(21)MICSAL
(22)NOTFLA
(23)PHEMIR
(24)MICPUN
(25)MINMEL
(26)ICTPUN
(27)PIMTEN
(28)NOTCRY
(29)PIMVIG
(1)CHRERY
(2)CAMANO
(3)ETHSPE
(4)SEMATR
(5)CATCOM
(6)ETHNIG
(7)LEPCYA
(8)NOTEXI
(9)LUXCOR
(10)CYPLUT
(11)LEPMEG
(12)GAMAFF
1,581
369
30
268
1
56
1
12
0
0
0
6
48
0
2
0
0
0
1
3
3
0
0
0
0
0
0
0
0
1,423
130
361
147
0
0
0
0
0
0
0
0
90
52
959
3
825
7
14
72
5
0
2
3
40
0
1
0
46
0
0
0
0
0
0
0
0
0
0
0
0
833
817
405
908
0
0
0
0
0
0
0
0
1,793
837
78
196
61
736
82
119
0
0
3
46
49
0
0
0
0
0
0
16
18
0
0
0
0
0
0
0
0
2,448
1,483
1,196
871
52
26
34
0
1
0
0
0
1,155
1,988
994
1,805
641
235
55
266
250
1
2
25
10
13
44
82
0
11
35
12
22
37
22
6
1
3
0
1
0
–
–
–
–
–
–
–
–
–
–
–
–
1,072
181
0
25
5
18
1
1
4
0
0
3
60
0
2
0
0
0
0
0
1
0
0
0
0
0
0
0
0
861
73
210
20
0
0
0
0
0
0
0
0
524
118
656
36
348
31
139
172
10
0
626
12
44
0
0
0
46
0
0
4
0
0
0
0
2
0
0
0
0
982
271
240
255
0
0
0
0
0
0
0
0
1,625
413
31
137
29
652
83
44
5
0
18
21
21
1
2
0
0
0
0
22
14
0
0
0
0
0
0
0
0
7,196
1,563
1,132
405
101
80
17
27
23
9
6
2
1,236
1,864
1,416
699
1,239
206
1,163
484
892
1,066
4
405
165
257
210
112
0
72
45
12
10
5
12
19
16
14
6
0
1
–
–
–
–
–
–
–
–
–
–
–
–
9,076
5,822
4,164
3,169
3,149
1,941
1,538
1,170
1,166
1,067
655
521
437
271
261
194
92
83
81
69
68
42
34
25
19
17
6
1
1
13,743
4,337
3,544
2,606
153
106
51
27
24
9
6
2
KPBS
physiological tolerance, as hypoxia and hyperthermia
tolerance were unrelated to abundance or occurrence.
Within-site patterns and relationships
As expected, most sites exhibited a log-normal SAD
(Shapiro–Wilk’s P-value > 0.05) characteristic of the
commonness–rarity pattern regardless of non-drought or
drought periods (Table 3). However, TPNP-4 did not
exhibit a log-normal SAD during drought, and KPBS-2
never conformed to the log-normal SAD. Changes in the
strength of commonness–rarity patterns were idiosyncratic across sites, with log-normal SADs strengthening
during drought as expected in TPNP-2, KPBS-1 and
KPBS-3, but weakening at TPNP-1, TPNP-3, KPBS-2 and
TPNP-4. In contrast to among-site patterns, nestedness
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
Colonisation, extinction, and commonness of stream fishes
10 000
TPNP
7
KPBS
Wet
Wet
Count
1000
100
10
1
M
RY
AN
RE
Dry
R
O
XC
LU
IG
HN
ET
YA
PC
LE
M
O
TC
CA
R
AT
M
SE
PE
HS
ET
O
CA
CH
NOTCRY
MINMEL
GAMAFF
ICTPUN
MICPUN
AMEMEL
FUNNOT
LABSIC
PHEMIR
AMENAT
PERCAP
NOTFLA
MICSAL
NOTTOP
LEPMAC
MOXERY
NOTSTR
SEMATR
LEPHUM
CYPLUT
LEPMEG
LEPCYA
LYTUMB
PIMNOT
LUXCAR
ETHSPE
CAMANO
10 000
Dry
Count
1000
100
10
1
CH
F
AF
AM
G
EG
PM
LE
UT
PL
CY
YA
PC
LE
R
O
XC
LU XI
TE
NO G
I
HN
ET OM
TC
CA TR
A
M
SE
PE
HS
ET NO
A
M
CA R Y
RE
PIMVIG
NOTFLA
PIMTEN
PHEMIR
ICTPUN
MINMEL
MICPUN
MICSAL
AMENAT
PERCAP
NOTTOP
FUNNOT
NOTSTR
LEPMAC
LABSIC
SEMATR
MOXERY
AMEMEL
LEPMEG
LUXCAR
LEPCYA
CYPLUT
GAMAFF
LEPHUM
LYTUMB
PIMNOT
ETHSPE
CAMANO
Fig. 2 Rank abundance (Whittaker) plots of species collected from intermittent streams on Tallgrass Prairie National Preserve (TPNP) and
Konza Prairie Biological Station (KPBS) during a wet (2008–2010) and dry (2011–2013) period. Abundance is the number of individuals
caught during each period (see Table 2 for species codes).
was not always the greatest contributor to community
dissimilarity across samples within sites. The contribution of nestedness to community dissimilarity during
wet periods was greater than turnover in two sites, was
lower in two sites and was equivalent in three sites, thus
failing to meet expectations in five of seven sites. The
importance of nestedness to community dissimilarity
increased during drought in four of seven sites and was
more important than the contribution of turnover during
drought in two of seven sites, once again failing to
match our predictions. Community composition was
identical through time in KPBS-2 resulting in no nestedness or turnover, with no turnover or nestedness also
occurring at KPBS-1 during the wet period.
Species that were abundant and persistent within sites
generally had high colonisation probabilities regardless
of drought conditions, further supporting the dispersal
limitation hypothesis (Table 4). However, unlike the
ubiquity of this relationship observed among sites,
colonisation and abundance were unrelated in KPBS-1
and KPBS-2 regardless of drought and were unrelated
during drought at TPNP-2. Furthermore, extinction
probabilities were negatively related to abundance and
persistence irrespective of drought in four sites, but in
TPNP-2, they were unrelated during drought, and in
KPBS-1 and KPBS-2, commonness and extinction were
never related. When commonness was predicted by
extinction, it was not due to physiological tolerance as
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
8
J. E. Whitney et al.
Table 3 Significance of log-normal species abundance distributions (SADs) and the contribution of turnover and nestedness to community
dissimilarity among and within sites during wet and dry periods on Tallgrass Prairie National Preserve (TPNP) and Konza Prairie Biological
Station (KPBS). Shapiro–Wilk’s P-values describes conformance of SADs to the log-normal distribution, with higher values indicating greater
conformance to log-normal SADs. Bold values indicated deviation from the distribution (see Fig. 2 for graphical presentation of among-site
SADs). Sørensen’s index describes overall community dissimilarity among samples
Analysis
Study area
Shapiro–Wilk Pvalue
Wet
Dry
Among-site
TPNP
KPBS
TPNP-1
TPNP-2
TPNP-3
TPNP-4
KPBS-1
KPBS-2
KPBS-3
0.16
0.17
0.18
0.40
0.87
0.25
0.34
0.03
0.21
Within-site
0.22
0.60
0.15
0.51
0.72
0.05
0.99
0.01
0.79
predicted by the environmental tolerance hypothesis, as
hypoxia and hyperthermia tolerance were either unrelated to abundance or occurrence, or were related in a
direction opposite that of our predictions.
Within-site case studies
Species that were more abundant in a catchment were
generally the first to re-colonise desiccated sites, but
later re-colonisation events were not associated with
catchment-wide abundance. Three of the 12 species present at TPNP-1 prior to complete drying in the autumn
of 2011 were able to re-colonise in May 2012 before the
stream dried again in August, but these species were
not necessarily more abundant in the catchment than
those that did not re-colonise (Table 5). Following complete drying at TPNP-1 in August 2012, nine species
were able to re-colonise by November of 2013. The initial two species to re-colonise by May 2013 were more
abundant in the catchment than those that did not
re-colonise by this time (P = 0.05), but subsequent
re-colonisation events were not predicted by catchmentwide abundance. Five species present at TPNP-1 prior to
drought never re-colonised during the continuous wetted period during November 2012–November 2013,
including Ameiurus natalis, Micropterus salmoides, Moxostoma erythrurum, Luxilus cardinalis and Pimephales notatus.
Re-colonisation at TPNP-2 following complete drying in
August 2012 was more rapid than that at TPNP-1, as 8
of the 12 species that eventually re-colonised did so
by May 2013. These species were more abundant in
the catchment than species that did not re-colonise.
Re-colonisation was not predicted by catchment-wide
abundance during August 2013, but Luxilus cardinalis
Turnover
Wet
Dry
Nestedness
Wet
Dry
Sørensen
Wet
Dry
0.15
0.00
0.18
0.08
0.04
0.05
0.00
0.00
0.13
0.11
0.00
0.30
0.21
0.12
0.08
0.00
0.00
0.05
0.25
0.33
0.07
0.10
0.04
0.10
0.00
0.00
0.00
0.29
0.50
0.26
0.03
0.12
0.02
0.13
0.00
0.09
0.40
0.33
0.25
0.19
0.08
0.15
0.00
0.00
0.13
0.40
0.50
0.56
0.24
0.24
0.10
0.13
0.00
0.14
was the most abundant species in the catchment among
those examined when it re-colonised TPNP-2 in November 2013. Minytrema melanops, which was never present
at TPNP-2 prior to drought, colonised this site following
complete drying. Notropis topeka, Ameiurus melas and
Ameiurus natalis did not re-colonise TPNP-2 during our
study.
Of the 12 species present at TPNP-3 prior to drought,
6 persisted through the period of drying to small, isolated pools (Table 6). Species that persisted possessed a
1.1 °C higher hyperthermia tolerance than species that
did not (F1,10 = 5.45; P = 0.04) but did not have greater
hypoxia tolerance (F1,10 = 1.96; P = 0.19) or greater predrought abundance (t = 1.80; P = 0.13).
Discussion
Dispersal limitation and environmental tolerance
Common species possessed higher colonisation and
lower extinction than rarer species in intermittent tallgrass prairie streams, potentially supporting both the
dispersal limitation and environmental tolerance
hypotheses. However, the mechanism behind the environmental tolerance hypothesis was not supported, as
the lower extinction of common species was not related
to hypoxia or hyperthermia tolerance. Mechanistic support behind the dispersal limitation hypothesis was also
equivocal, as species abundant in the catchment tended
to be the first to re-colonise desiccated locations, suggesting colonisation is predicted by rather than predictive of abundance.
Although our findings are likely most representative
of relatively unimpacted tallgrass prairie streams located
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
Colonisation, extinction, and commonness of stream fishes
9
Table 4 Relationships between commonness (abundance and occurrence) and predictor variables describing the dispersal limitation hypothesis (probability of colonisation pc) and environmental tolerance hypothesis (probability of extinction pe; hypoxia and hyperthermia tolerance). Relationships are described by their slope (b), variance explained (r2), statistical significance (P) and degrees of freedom (d.f. = 1 –
number of species) and were calculated among and within sites on Tallgrass Prairie National Preserve (TPNP) and Konza Prairie Biological
Station (KPBS) during a wet (2008–2010) and dry period (2011–2013). Values in bold are significant (P ≤ 0.05), and italicised values are marginally significant (P ≤ 0.10) (see Figs 3 (abundance) and 4 (occurrence) for graphical presentation of among-site relationships)
Abundance
Occurrence
Wet
Location
Predictor
TPNP
Among
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
pc
pe
Hypoxia
Hyperthermia
KPBS
Among
TPNP-1
Within
TPNP-2
Within
TPNP-3
Within
TPNP-4
Within
KPBS-1
Within
KPBS-2
Within
KPBS-3
Within
b
Dry
2
0.80
0.73
0.44
0.24
0.71
0.77
0.77
0.15
0.68
0.35
1.60
0.01
0.66
0.55
2.23
0.72
0.47
0.48
0.37
0.08
0.56
0.62
0.19
0.14
0.00
0.34
4.79
0.29
0.00
0.00
0.00
0.10
0.91
0.71
0.82
0.16
r
P
DF
0.66
0.76
0.02
0.06
0.87
0.68
<0.01
0.01
0.78
0.38
0.06
<0.01
0.71
0.81
0.20
0.41
0.35
0.55
<0.01
<0.01
0.48
0.87
<0.01
0.02
0.05
0.21
0.74
0.17
0.87
0.87
0.02
0.66
0.91
0.75
<0.01
0.02
<0.01
<0.01
0.62
0.37
<0.01
0.01
0.86
0.80
<0.01
0.02
0.46
0.96
<0.01
<0.01
0.17
0.03
0.03
<0.01
0.77
0.80
<0.01
<0.01
0.83
0.60
0.77
0.54
0.14
0.59
0.07
0.07
0.85
0.18
<0.01
<0.01
0.83
0.77
26
26
16
15
7
7
7
7
13
13
10
10
13
13
10
10
12
12
10
10
25
25
16
15
3
3
3
3
3
3
3
3
7
7
7
7
b
Wet
0.98
0.74
0.40
0.10
0.53
0.47
1.79
0.45
0.73
0.45
2.50
0.19
0.42
0.10
2.36
0.27
0.54
0.62
1.04
0.05
0.70
0.60
0.31
0.29
0.27
0.83
4.73
0.02
0.00
0.13
2.72
0.17
0.79
0.46
1.68
0.43
Dry
r
P
DF
b
r
P
DF
0.39
0.45
0.02
0.02
0.52
0.38
0.06
0.19
0.37
0.38
0.17
0.03
0.10
0.02
0.47
0.10
0.56
0.88
0.11
<0.01
0.67
0.83
<0.01
0.09
0.60
0.60
0.35
<0.01
0.07
0.07
0.64
0.15
0.76
0.45
0.05
0.19
<0.01
<0.01
0.59
0.63
<0.01
0.03
0.49
0.19
0.03
0.03
0.24
0.66
0.25
0.65
0.02
0.35
<0.01
<0.01
0.25
0.88
<0.01
<0.01
0.72
0.25
0.23
0.23
0.41
0.98
0.74
0.73
0.20
0.61
<0.01
0.02
0.49
0.18
27
27
16
15
11
11
10
10
11
11
9
9
14
14
10
10
15
15
13
13
26
26
16
15
3
3
3
3
3
3
3
3
11
11
10
10
0.78
0.62
0.76
0.24
0.80
0.75
0.20
0.22
0.80
0.44
0.55
0.31
0.65
0.55
2.23
-0.74
0.77
-0.69
-1.93
0.09
0.64
-0.67
-0.34
-0.10
0.00
-1.12
0.49
0.66
0.00
0.00
0.00
0.00
0.90
-0.65
–0.50
0.12
0.79
0.71
0.06
0.07
0.93
0.56
<0.01
0.02
0.85
0.47
<0.01
0.05
0.73
0.86
0.22
0.49
0.67
0.81
0.22
<0.01
0.55
0.92
<0.01
<0.01
0.04
1.00
<0.01
0.07
1.00
1.00
1.00
1.00
0.95
0.68
<0.01
<0.01
<0.01
<0.01
0.32
0.32
<0.01
0.03
0.97
0.73
<0.01
<0.01
0.78
0.51
<0.01
<0.01
0.14
0.02
<0.01
<0.01
0.15
0.81
<0.01
<0.01
0.72
0.74
0.81
–
0.94
0.38
–
–
–
–
<0.01
0.01
0.89
0.82
26
26
16
15
7
7
7
7
13
13
10
10
13
13
10
10
12
12
10
10
25
25
16
15
3
3
3
3
3
3
3
3
7
7
7
7
2
in the Flint Hills, they may reflect general patterns
across more anthropogenically modified prairie streams
or other systems. They agree with those from a perennial shortgrass prairie stream, where mean occupancy
(Gotelli & Taylor, 1999a) and average population size
(Gotelli & Taylor, 1999b) predicted colonisation and
extinction. Similarly, Taylor & Warren (2001) found that
colonisation and extinction probabilities in an intermittent, Ozark stream were strongly related to mean abundance. Greater propagule supply from larger population
2
b
0.95
0.50
0.43
0.08
0.63
0.46
1.85
0.38
0.50
0.26
0.85
0.08
0.35
0.17
0.88
0.24
0.64
0.66
1.04
2.54
0.80
0.64
0.16
0.04
0.42
1.31
0.57
0.77
0.00
0.90
5.03
0.70
0.95
0.53
1.41
0.50
r2
P
DF
0.72
0.49
0.05
0.02
0.62
0.31
0.05
0.11
0.58
0.44
0.07
0.01
0.22
0.14
0.23
0.27
0.69
0.85
0.09
<0.01
0.74
0.83
<0.01
<0.01
1.00
1.00
<0.01
0.38
0.33
1.00
0.63
0.75
0.94
0.50
0.03
0.22
<0.01
<0.01
0.40
0.59
<0.01
0.06
0.53
0.32
<0.01
0.02
0.47
0.74
0.08
0.17
0.14
0.10
<0.01
<0.01
0.30
0.82
<0.01
<0.01
0.87
0.87
–
–
0.95
0.38
0.42
–
0.21
0.14
<0.01
0.01
0.60
0.15
27
27
16
15
11
11
10
10
11
11
9
9
14
14
10
10
15
15
13
13
26
26
16
15
3
3
3
3
3
3
3
3
11
11
10
10
size (Sheldon & Meffe, 1995; Stoll et al., 2014) and
enhanced extinction risk from smaller population size
(Richter-Dyn & Goel, 1972; Lima et al., 1996) were
invoked as mechanisms by Gotelli & Taylor (1999a,b)
and Taylor & Warren (2001) explaining the positive and
negative relationships of colonisation and extinction,
respectively, with commonness. Colonisation and extinction can also interact to influence commonness, as higher
colonisation can lead to lower extinction via rescue
effects (Brown & Kodric-Brown, 1977). Finally, neutral
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
J. E. Whitney et al.
10
TPNP
KPBS
4
4
3
3
2
2
Wet
Wet
Dry
Dry
1
0
–2
–1
0
1
1
0
2
–2
–1
0
Logit (pc)
1
2
–2
–1
0
Logit (pe)
1
2
1.0
1.4
Log10 (Abundance+1)
Logit (pc)
4
4
3
3
2
2
1
1
0
0
–2
–1
0
Logit (pe)
1
2
4
4
3
3
2
2
1
1
0
0
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
0.4
Hypoxia tolerance (mg L–1 DO)
4
3
3
2
2
1
1
0
0
36.0
36.5
37.0
37.5
38.0
Hyperthermia tolerance (°C)
0.8
1.2
1.6
1.8
Hypoxia tolerance (mg L–1 DO)
4
35.5
0.6
38.5
35.5
36.0
36.5
37.0
37.5
38.0
Hyperthermia tolerance (°C)
processes (Matthews & Whittaker, 2014) or other traits
(e.g. body size; trophic guild) may be responsible for
commonness, although previous studies have found limited effects of other traits on colonisation or extinction
dynamics of stream fishes (Gotelli & Taylor, 1999b; Taylor & Warren, 2001).
Hyperthermia and hypoxia tolerance did not predict
extinction, although species with greater hyperthermia
tolerance were more likely to persist in a drying pool on
TPNP. The contradiction between the significant effect
of hyperthermia tolerance on persistence compared to its
lack of effect on extinction may stem from differences in
the temporal scope during which persistence (6 months)
and extinction (3 years) were calculated, and in the
38.5
Fig. 3 Relationships testing the dispersal
limitation (probability of colonisation pc)
and the environmental tolerance
hypotheses (probability of extinction pe;
hypoxia tolerance; hyperthermia tolerance) in describing species abundance
during wet (2008–2010) and dry (2011–
2013) periods among sites on Tallgrass
Prairie National Preserve (TPNP) and
Konza Prairie Biological Station (KPBS).
Each observation corresponds to a species in the catchment (see Table 4 for statistical results).
categorical (persistence) versus continuous (extinction
probability) nature of these variables. Non-significant
effects of hypoxia and hyperthermia on commonness
may also be a statistical artefact, as the small number of
species for which tolerance data were available may
have limited our power to detect meaningful relationships (Table 4). The lack of an influence of physiological
tolerance on extinction is surprising given the findings
of Smale & Rabeni (1995b) that showed hypoxia tolerance was an important predictor of community membership in intermittent prairie and Ozark forest streams.
Similarly, Matthews & Styron (1981) found that cyprinids and percids in headwater habitats possessed
greater hypoxia tolerance than their confamilial,
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
Colonisation, extinction, and commonness of stream fishes
TPNP
1
Logit (po)
2
1
0
0
–1
–1
–2
–2
–2
Fig. 4 Relationships testing the dispersal
limitation (probability of colonisation pc)
and the environmental tolerance
hypotheses (probability of extinction pe;
hypoxia tolerance; hyperthermia tolerance) in describing species probability of
occupancy (po) during wet (2008–2010)
and dry (2011–2013) periods among sites
on Tallgrass Prairie National Preserve
(TPNP) and Konza Prairie Biological Station (KPBS). Each observation corresponds to a species in the catchment (see
Table 4 for statistical results).
KPBS
Wet
Wet
Dry
Dry
2
–1
0
Logit (pc)
1
2
2
2
1
1
0
0
–1
–1
–2
–2
–2
–1
0
Logit (pe)
1
2
2
2
1
1
0
0
–1
–1
–2
–2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1
0
0
–1
–1
–2
–2
Species abundance distributions
Although we were unable to unequivocally determine
the causes of commonness, our results clearly indicated
that most fishes in our prairie streams were rare, while
few were common as indicated by consistent confor-
1
2
–2
–1
0
Logit (pe)
1
2
36.0
36.5
37.0
37.5
38.0
Hyperthermia tolerance (°C)
1
downstream counterparts in an upland forest stream.
The limited influence of hypoxia tolerance in the intermittent stream networks we studied suggested hypoxia
may not have occurred in our system even during
drought, but we did not monitor dissolved oxygen. Less
severe environmental conditions would be expected in
our pristine prairie streams than in other prairie streams,
many of which are stressed by nutrient runoff and riparian alteration (Dodds et al., 2004).
0
Logit (pc)
35.5
2
38.5
–1
0.6
0.8
1.0
1.2
1.4
Hypoxia tolerance (mg L–1 DO)
2
36.0
36.5
37.0
37.5
38.0
Hyperthermia tolerance (°C)
–2
0.4
Hypoxia tolerance (mg L–1 DO)
35.5
11
1.6
38.5
mance to the log-normal SAD. Interest in variation of
SADs across spatial environmental gradients, species
richness gradients and temporal disturbance gradients
has gained renewed attention (McGill et al., 2007). We
found that the structure of SADs was fairly constant
across samples that varied in location, drought period
and species richness. However, TPNP-4 shifted away
from the log-normal SAD during drought, as species
that were moderately abundant (total rank 5–15, Table 2)
during wet conditions became much more abundant
during the drought, resulting in greater evenness. This
result contrasted with among-site patterns, wherein lognormal distributions strengthened during drought especially on KPBS, potentially because more rare species
were encountered (Table 2). Furthermore, KPBS-2 never
conformed to the log-normal SAD, as the four species
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
12
J. E. Whitney et al.
Table 5 Sequence of species re-colonisation on Tallgrass Prairie National Preserve (TPNP) sites 1 and 2 following complete drying in
September–November 2011 (drying event 1) and August–November 2012 (drying event 2). Chi-square P-values describes significance of the
relationship between re-colonisation and catchment-wide abundance. Significant relationships are in bold. Species codes are the first three
letters of the genus and first three letters of the specific epithet
Drying event
Site
Re-colonisation period
Colonising species
v2 P-value
1
2
TPNP-1
TPNP-1
2
TPNP-2
May 2012
May 2013
August 2013
November 2013
May 2013
CAMANO, ETHSPE, LEPCYA
CAMANO, ETHSPE
LEPCYA, LYTUMB
CYPLUT, LEPHUM, LEPMAC, LEPMEG, SEMATR
CAMANO, ETHSPE, LEPCYA, LEPHUM, LEPMEG,
LYTUMB, PIMNOT, SEMATR
CYPLUT, MINMEL, MOXERY
LUXCAR
0.33
0.02
0.53
0.89
0.05
August 2013
November 2013
0.19
0.03
Table 6 Comparison of mean (SE) hypoxia tolerance, hyperthermia tolerance and pre-drought abundance of species that persisted versus
those that were extirpated during a drying event on Tallgrass Prairie National Preserve (TPNP) site 3 during May–November 2012. Species
codes are the first three letters of the genus and first three letters of the specific epithet. Values that significantly differed (P < 0.05) between
groups are in bold
Category
Species
Hypoxia
DO mg L
Persisted
Extirpated
AMEMEL, AMENAT, CAMANO, ETHSPE, LEPCYA, LEPHUM
LEPMEG, LUXCAR, LYTUMB, MOXERY, PIMNOT, SEMATR
0.78 (0.10)
0.95 (0.26)
that occurred there always had similar population sizes
(Table 2). Depauperate communities generally exhibit
greater evenness (McGill et al., 2007), but low species
richness cannot explain the high evenness in KPBS-2, as
KPBS-1 also had four species but still exhibited a lognormal SAD. Greater productivity at KPBS-2 than at
KPBS-1 could explain the difference between their SADs,
as productivity increases community evenness by
increasing the abundance of rare species (Whittaker,
1960; Hubbell, 1979; Hurlbert, 2004).
Nestedness, turnover and community dissimilarity
Nestedness was more important than turnover in
describing spatial patterns of community dissimilarity,
with this importance increasing during drought. These
results further confirm the importance of nestedness in
shaping stream fish communities and longitudinal richness gradients in intermittent–perennial stream networks, and despite our limited spatial extent that could
have confounded our nestedness analyses, agree with
findings from other streams in the region. For example,
Taylor (1997) found stream fish communities in pools of
an upland stream were nested according to position and
size in connected and isolated pools, respectively,
suggesting colonisation drove nestedness in connected
pools, whereas extinction drove nestedness in isolated
pools. Spatial nestedness patterns were also found in
1
Hyperthermia
°C
Mean abundance
Log10 (x)
37.3 (0.31)
36.2 (0.40)
80 (40)
7 (2)
five wadeable streams distributed across mesic plains
and upland forest by Roberts & Hitt (2010), with colonisation from nearby large rivers invoked to explain nestedness. Colonisation and extinction also appear to be
strongly associated with nestedness in our intermittent
tallgrass prairie streams. However, it remains uncertain
whether high occurrence is predicted by rather than predictive of colonisation and extinction, confounding our
ability to support or refute the conceptual model of
Larned et al. (2010) that predicted spatial nestedness patterns are driven by differential colonisation.
Turnover and nestedness were of similar importance
in contributing to temporal community dissimilarity patterns within sites across years of varying drought intensity. These results contradict the conceptual model of
Larned et al. (2010), which predicted that nestedness
should increase during drought as the extinction of
intolerant species results in community divergence. Our
inability to support the Larned et al. (2010) conceptual
model is not surprising given that physiological tolerance was unrelated to extinction. Our results also contradict those of Taylor & Warren (2001), who found that
temporal community dissimilarity patterns within pools
of intermittent upland forest streams were explained by
nestedness resulting from the selective extinction of
species. Our results thus suggest species replacement
is as important as species loss in driving community
dissimilarity across years of varying drought intensity in
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668
Colonisation, extinction, and commonness of stream fishes
intermittent tallgrass prairie streams. However, these
patterns could shift in coming decades, as global climate
change (D€
oll & Schmied, 2012; Arnell & Gosling, 2013;
Jaeger, Olden & Pelland, 2014) and greater human water
demand (Gleick, 2003; Meybeck, 2003) increase the frequency, intensity, and duration of drought and intermittency. The selective pressures induced by these
anthropogenic changes could fundamentally alter nestedness, turnover and commonness patterns in intermittent tallgrass prairie streams, but colonisation and
extinction will doubtlessly continue to be important
structuring processes for stream fish communities.
Acknowledgments
Support for this research was provided by the Kansas
State University Division of Biology, the National Park
Service Tallgrass Prairie National Preserve and the
Konza Prairie Long Term Ecological Research programme. We also thank the Nature Conservancy for
access to study locations in both Tallgrass Prairie
National Preserve and Konza Prairie Biological Station.
Josiah Maine, Darin McCullough, Josh Perkin, Tyler
Pilger, Jeff Rogosch, Dustin Shaw and Matt Troia
helped with data collection. Matt Troia also assisted
with creation of the study map. This is contribution
No. 16-055-J from the Kansas Agricultural Experiment
Station.
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Supporting Information
Additional Supporting Information may be found in the
online version of this article:
Figure S1. Palmer Modified Drought Index for National
Oceanic and Atmospheric Administration (NOAA) Kansas Climate Division 6 (NOAA National Climatic Data
Center) and mean daily discharge on Kings Creek (United States Geological Survey gage #06879650) during 01
January 2008 – 01 December 2013. Horizontal bars indicate annual mean daily discharge. Note log10 scale on yaxis for discharge values.
(Manuscript accepted 4 August 2015)
© 2015 John Wiley & Sons Ltd, Freshwater Biology, doi: 10.1111/fwb.12668