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Urban buried streams: Abrupt transitions in habitat and biodiversity

Article in The Science of The Total Environment · January 2022

DOI: 10.1016/j.scitotenv.2022.153050

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 Science of the Total Environment 819 (2022) 153050

Contents lists available at ScienceDirect

Science of the Total Environment

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

Urban buried streams: Abrupt transitions in habitat and biodiversity

⁎
Chelsea L. Hintz a, , Michael T. Booth a, Tamara A. Newcomer-Johnson b, Ken M. Fritz b, Ishi Buffam a,c
a
Department of Biological Sciences, University of Cincinnati, Cincinnati, OH 45221, United States of America
b
United States Environmental Protection Agency, Office of Research and Development, Cincinnati, OH 45268, United States of America
c
Department of Landscape Architecture and Management, Swedish University of Agricultural Sciences, Alnarp, Sweden

H I G H L I G H T S G R A P H I C A L A B S T R A C T

• Stream burial is widespread in industrial-

ized and densely populated urban areas.
• Culverts have dramatic impacts on the
structure and function of streams.
• The buried segments of streams had
smaller substrate and deeper water
depths.
• Buried segments also had depleted periph-
yton and macroinvertebrate populations.
• Transitions in many abiotic and biotic pa-
rameters were abrupt rather than gradual.

A R T I C L E I N F O A B S T R A C T

Article history: Stream burial, the rerouting of streams into underground culverts, is common in industrialized and densely populated
Received 11 August 2021 urban areas. While stream burial is common in urban environments, direct characterization of the within-culvert en-
Received in revised form 3 January 2022 vironment is rare and it is unclear if buried reaches reflect neighboring open reaches regarding habitat, biota, and
Accepted 7 January 2022
water chemistry. Additionally, for a buried stream, the entrance and exit of the culvert are abrupt habitat transitions
Available online 14 January 2022
within the stream channel, and it is unknown if these transitions lead to similarly abrupt responses in biotic and abiotic
Editor: Sergi Sabater characteristics or if responses are gradual. Quantifying the within-culvert environment and transitions upon entering/
exiting the culvert has rarely been done but can help inform management practices regarding how these systems are
Keywords: impacted and establish a baseline for evaluating daylighting or stream restoration projects. To understand how cul-
Urban streams verts affect longitudinal biotic and abiotic characteristics of urban streams, we evaluated longitudinal patterns of phys-
Buried streams ical habitat characteristics, stream water physiochemistry, periphyton biomass, and macroinvertebrate density and
Transitions diversity in two urban streams that included long (>100 m) culvert reaches. Abrupt transitions in a suite of abiotic
Macroinvertebrates and biotic variables were observed at the entrances and exits of the culverts whereas some variables showed no re-
Substrate
sponse to the culvert presence. Periphyton biomass and macroinvertebrate density were reduced by 98% and 92%, re-
Culverts
spectively, by culverts in the two streams. Within the culverts, we observed greater water depths (average of 10 cm
outside vs 26 cm within the culvert), finer benthic substrate, and diversity of macroinvertebrates was reduced by
50%. Nutrient concentrations, in contrast, showed no response to the presence of a culvert. Within 60–90 m down-
stream of the culvert exits, most of the measured parameters returned to levels similar to those observed upstream
of the culvert, suggesting that the ecosystem impacts of urban culverts, though dramatic, may be spatially constrained.
Published by Elsevier B.V.

⁎ Corresponding author.
E-mail address: hintzca@mail.uc.edu (C.L. Hintz).

http://dx.doi.org/10.1016/j.scitotenv.2022.153050
0048-9697/Published by Elsevier B.V.
C.L. Hintz et al.
1. Introduction
The effects of urbanization on stream ecosystems have been coined the
“urban stream syndrome” because urbanization induces fundamental envi-
ronmental modifications which can change stream ecosystem habitat and
function (Walsh et al., 2005). These changes in habitat and function can re-
sult from the increase in impervious cover within the watershed and the as-
sociated increase in runoff volume. Changes to sediment regime, thermal
regime, and pollutant loading can also cause changes to stream ecosystem
habitat and function (Kaushal et al., 2015; Wenger et al., 2009). Urbaniza-
tion can also lead to extreme physical modifications within the stream
channel, and perhaps the most extreme impact of urban development on
streams is their wholesale containment and burial within culverts or pipes
(Elmore and Kaushal, 2008; Julian et al., 2015).
Stream burial is widespread in heavily industrialized and densely popu-
lated urban areas (Elmore and Kaushal, 2008). For example, within the U.S.
cities of Baltimore and Detroit, an estimated 66% and 85% of all streams
have been buried, respectively (Elmore and Kaushal, 2008; Napieralski
and Carvalhaes, 2016). A broader survey of U.S. Census Bureau Urban
Areas in 11 megaregions classified 11,490 km2 of metropolitan areas as
“urban stream deserts,” where stream channels have likely been removed
or buried (Napieralski and Carvalhaes, 2016). In Europe, extensive stream
burial has been documented in London (Napieralski and Carvalhaes,
2016), and country-wide extent of stream burial are 15% and 20% in
Denmark and Sweden, respectively (Wild et al., 2011). Burial happens to
streams of various sizes and flow regimes, and culverts can range in length
from tens of meters to several hundred meters (Beaulieu et al., 2014; Julian
et al., 2015; Olson et al., 2017).
Stream burial abruptly changes key physical drivers of stream ecosys-
tems. These changes include the absence of light, the elimination of subsur-
face interaction, and reduced riparian inputs (Fig. 1). Other physical effects
from stream burial and the presence of a culvert can include increased
water velocities at the culvert exit (Elmore and Kaushal, 2008; Hope
et al., 2014), constricted stream channel width (Beaulieu et al., 2014), lim-
ited sediment depth (Hope et al., 2014), and altered substrate composition
Fig. 1. Conceptual diagram describing hypothesized changes resulting from stream burial based on literature review. Colored boxes (grey and yellow) represent parameters
directly quantified in our study (with the exception of subsurface interactions and velocity). Grey boxes are primary physical responses and could influence changes in the
yellow boxes which represent secondary effects. Blue arrows between boxes represent the directional influence of one variable on another variable. Black arrows within boxes
represent the anticipated effect on a variable in response to burial (“↑” indicates an increase, “↓” indicates a decrease, and “↑↓” indicates inconsistent change within the
literature).
2
Science of the Total Environment 819 (2022) 153050
(Macpherson et al., 2012; Wellman et al., 2000). The impact of culverts on
substrate size is not consistent across streams, however, and depends on a
host of factors including channel slope, type of hydraulic control, substrate
embeddedness, and culvert shape, length, and material (Howley, 2004).
Previous work has demonstrated that culverts can contain larger substrate
(U.S. Department of Agriculture, 2009), smaller substrate (Macpherson
et al., 2012; Wellman et al., 2000), or show no differences in substrate
size (Hope et al., 2014) when compared to adjacent open stream reaches.
Additionally, downstream of culverts there is often bed scouring leading
to pool formation and subsequent deposition of fine substrates (Wellman
et al., 2000). Stream burial simplifies and fragments habitat, decreasing
habitat availability (Elmore and Kaushal, 2008), and modifying ecosystem
processes such as nutrient retention (Beaulieu et al., 2014).
Given the physical changes to the streambed created by burial, changes
in water chemistry might be expected within culverts. However, few stud-
ies have directly observed changes in water chemistry due to stream burial.
In one detailed study of a single urban New Hampshire stream, no differ-
ences were found between buried and open reaches in a variety of solute
concentrations and water chemistry parameters (Hope et al., 2014). Nutri-
ent uptake is slow or undetectable within culverts (Beaulieu et al., 2014,
2015; Pennino et al., 2014), suggesting that culverts primarily transport nu-
trients between open stream reaches with minimal in-stream retention or
removal within the culvert.
Research regarding the impact of burial on stream biota has focused on
how a culvert impacts animal (primarily fish and some macroinvertebrates)
movement (Blakely et al., 2006; Macpherson et al., 2012) and the distribu-
tion of basal resources within streams (Neale and Moffett, 2016). Informa-
tion on algae is limited but biomass (as determined by concentrations of
chlorophyll a) is typically low in culverts and gross primary production is
not supported (Beaulieu et al., 2014; Hope et al., 2014) due to the lack of
light. In contrast, ecosystem respiration is observed in culverts indicating
the presence of heterotrophic organisms and organic matter (Beaulieu
et al., 2014; Hope et al., 2014). Shifts in macroinvertebrate functional feed-
ing groups have been observed in culverts relative to the surrounding
stream, attributed to changes in the distribution of basal resources (Neale
C.L. Hintz et al.
and Moffett, 2016). Downstream drift of aquatic invertebrates can be lower
downstream of culverts and the abundance and diversity of EPT (Ephem-
eroptera-Plecoptera-Trichoptera) invertebrate taxa (taxa indicative of bet-
ter water quality) can be reduced when compared to nearby streams
without culverts (Meyer et al., 2005). Dispersal can be negatively impacted
because culverts act as upstream flight barriers for adult insects, reducing
upstream recruitment (Blakely et al., 2006). The upstream movement of
some fish can be negatively impacted by culverts, with culvert slope (>
3–5%) and hang height (drop to stream) being the most important factors
impacting upstream fish movement (Burford et al., 2009; Favaro et al.,
2014; Macpherson et al., 2012; Poplar-Jeffers et al., 2009).
Streams are naturally heterogenous (Allan, 2004) and it is unclear
whether the properties of buried stream reaches generally reflect adjacent
open reaches (regarding habitat, biota, and water chemistry) or if buried
reaches fundamentally differ in structure and function, leading to habitat
fragmentation (Beaulieu et al., 2014; Blakely et al., 2006; Elmore and
Kaushal, 2008; Macpherson et al., 2012). To date, research has examined
the impacts of culverts by comparing reaches up- and downstream of cul-
verts or by only evaluating a single location within a buried reach. Because
few studies have taken a fine-scale approach in the characterization of bur-
ied reaches, little is known about the spatial variation of physical, chemical,
and biological parameters as channels transition from open to buried to
open reaches.
For a buried stream, the entrance and exit of the culvert are abrupt phys-
ical transitions within the stream channel, and it is unknown if these tran-
sitions lead to abrupt changes in biotic and abiotic characteristics or if
changes observed in the stream are more gradual. Abrupt changes in
land-use can impact biota via edge effects by direct and indirect means
(Murica, 1995), but there can be a spatial lag between riparian change
and in-stream conditions (Feijó-Lima et al., 2018). For example, transitions
in land-use (e.g. forest to pasture) have resulted in responses on different
spatial scales from 10s of m (temperature and sediment size) to 100s of m
(macroinvertebrate diversity and abundance) (Feijó-Lima et al., 2018;
Goss et al., 2014). We expect that when a buried stream enters and exits a
culvert, different ecosystem components will respond in varying ways,
both in the extent of the response and in the spatial lag.
Urbanization is expected to increase (US Census Bureau, 2016) leading
to further stream burial in developing areas (Weitzell et al., 2016), so it will
be pertinent to know whether burial leads to additional downstream effects
(i.e., a spatial lag in community characteristics) as well as the localized eco-
logical impact of burying the stream. Determining the ecological impact of
stream burial – and the spatial extent of this impact – could help inform de-
cisions involving stormwater management, habitat, nutrient processing,
flood control, and the monitoring of restoration projects (Newcomer-
Johnson et al., 2014; Newcomer Johnson et al., 2016).
We aimed to (1) directly characterize the longitudinal spatial variation
in biotic and abiotic ecosystem parameters within buried stream reaches,
(2) determine the magnitude of change for a given parameter upon enter-
ing/exiting a culvert, and (3) evaluate spatial lags in the response of differ-
ent parameters to the abrupt transitions at the entrance and exit of a culvert.
We evaluated two urban streams that included long (>100 m) culvert
reaches by carrying out longitudinal transects of physical habitat character-
istics, stream water physiochemistry, periphyton biomass, and macroinver-
tebrate density and diversity. These parameters are often impacted in urban
streams and can be further altered by stream burial. We hypothesized
(Fig. 1) that buried stream reaches represent altered ecosystems when com-
pared to open upstream and downstream reaches and that we would ob-
serve clear transitions in biotic parameters (such as periphyton biomass
and macroinvertebrates) and some abiotic parameters (such as tempera-
ture) downstream of the entrance and exit of the culvert.
Our specific expectations regarding abiotic parameters were as follows:
At the culvert entrance we hypothesized no change in substrate size or
water depth, given the variable findings of previous research and host of
factors that influence these parameters (Howley, 2004; Macpherson et al.,
2012; Wellman et al., 2000). Within the culverts, we expected that the dis-
tribution of substrate size and water depth would be similar to what was
3
Science of the Total Environment 819 (2022) 153050
observed in open upstream reaches, except at the culvert exit where pool
formation and deposition of fine sediments is often observed (Wellman
et al., 2000). We expected a downstream spatial lag at the culvert exit,
i.e., the presence of a pool and smaller substrate would be present some dis-
tance downstream before the stream reach recovered its upstream qualities
and characteristics. Water temperature was expected to decrease in the cul-
verts due to the lack of solar radiation and gradually recover downstream of
the culvert exit. We expected an increase in inorganic nutrient (N and
P) concentrations as distance into the culvert increased (due to mineraliza-
tion rates > uptake rates), and a decrease in inorganic nutrient concentra-
tions downstream of the culvert as the periphyton community and
associated primary production recover.
Regarding biotic parameters, we expected that periphyton biomass
within the culvert would be minimal due to the elimination of light
(Beaulieu et al., 2014). We expected an abrupt decline in periphyton bio-
mass upon entering the culvert and an abrupt increase upon exiting the cul-
vert to levels observed upstream. We also expected the density and
diversity of macroinvertebrates to be lower within the culverts compared
to upstream reaches because of the elimination of resources (e.g., algae,
leaf litter subsidies; Fuller et al., 2004; Kiffney et al., 2004), and to recover
downstream of the culvert, but with a longer spatial lag than that of periph-
yton.
2. Methods
2.1. Stream description and study design
We monitored two streams whose buried reaches are >100 m in length
– Congress Run and an unnamed tributary (here and throughout referred to
as Princeton) from July 2017 – August 2018. Congress Run and Princeton
are first-order tributaries in the Mill Creek Watershed, an urban watershed
in Cincinnati, OH, USA (29.3% impervious cover). Cincinnati has a temper-
ate climate with no defined dry and wet seasons and receives an average of
1143 mm of precipitation (primarily rainfall) annually (NOAA, https://
www.weather.gov). The Congress Run watershed is 9.9 km2 with 73.6%
developed land (mixed industrial and residential use) and 18.8% impervi-
ous surface cover (US Geological Survey, 2016), but retains a riparian cor-
ridor with an average of 75% canopy cover (summer). The stream is
intermittent within the study reach and has a 183 m long buried reach
(Shifflett et al., 2019). Stormwater outfalls and a combined sewer overflow
are present along Congress Run. The Congress Run culvert (approximately
2 m tall and 3 m wide) is constructed of corrugated metal and runs beneath
a golf course (Fig. 2). The study reaches have slopes of 0.89% (upstream),
0.22% (culvert), and 1.16% (downstream). The Princeton watershed
(8.8 km2) is 95.3% developed land and 38.2% impervious surface cover
(US Geological Survey, 2016). Princeton is surrounded by commercial
land use but retains an intact riparian corridor (large trees, brush within
the understory) of 2–3 m on each bank within the study reach (CH, personal
observation). Princeton is a perennial stream with stormwater outfalls pres-
ent and includes a 120 m long buried reach. The culvert is constructed of
corrugated metal with a poured concrete bottom and runs beneath a high-
way interchange (Fig. 2). The Princeton culvert is approximately 4.4 m tall
and 4 m wide. The reaches have slopes of 1.02% (upstream), 0.19% (cul-
vert), and 0.66% (downstream).
Our study design included longitudinal surveys including stream
reaches upstream of the culvert, within the culvert, and downstream of
the culvert to characterize the abiotic and biotic spatial variation within
reaches and across transitions between open and buried reaches (Fig. 2).
Periphyton and physical habitat (such as light, depth, substrate size) were
collected at 13 evenly spaced sites per stream: 3 sites upstream of the cul-
vert (upstream), 7 within-culvert sites (culvert), and 3 sites downstream
of the culvert (downstream). The distance between sites was 30 m in Con-
gress Run and 20 m in Princeton. All measurements described were con-
ducted at baseflow as identified by a local Mill Creek USGS stream gage
(#03259000).
C.L. Hintz et al.
Fig. 2. a) Map of Congress Run. Distance between sites is 30 m. b) Congress Run, downstream opening of culvert. c) Map of Princeton. Distance between sites is 20 m.
d) Princeton, downstream opening of culvert.
2.2. Physical habitat (light, substrate, water depth)
The distribution of substrate size was determined at each site (N =
13 sites per stream) in June 2018. At each site, ≥100 pieces of sub-
strate were randomly selected, and their size (determined by the inter-
mediate axis) was determined using a gravelometer (Wildco, Yulee,
FL). The D18%, D50%, and D84% (the particle size at which 18%,
50%, and 84% of the substrate was smaller and represents the smaller,
median, a coarser fraction of substrate; Olsen et al., 2005) were deter-
mined. For culverted sites, the bottom of the culvert was assigned its
own category and was not included in the calculations of D18%,
D50%, and D84%. Water depth (cm) and an instantaneous measure-
ment of photosynthetically active radiation (PAR, expressed as photo-
synthetic photon flux density, PPFD) were recorded during
periphyton collection (every other month) at each of our 13 sites per
stream. Light readings were collected with a LI-COR LI-192 (LI-COR
Biosciences, Lincoln, NE) and were collected under varying cloud
cover and time of day.
2.3. Water chemistry
Stream water quality parameters were measured every other month at 5
locations within each stream corresponding to 1 upstream site, 3 culvert
sites, and 1 downstream site (Fig. 2). The upstream and downstream sites
were at the beginning and end of each study reach, respectively. The 3
4
Science of the Total Environment 819 (2022) 153050
culvert sites corresponded to the entrance, mid-point, and exit of the cul-
vert. Instantaneous measurements of temperature, dissolved oxygen, pH,
and conductivity were collected with a calibrated YSI ProDSS (Yellow
Springs, OH). The YSI ProDSS was calibrated before each day of sampling
according to the manufacturer's instructions. Water samples (1 L) were col-
lected in a downstream to upstream direction to determine nutrient concen-
trations and were kept on ice until returned to the lab for filtering. Samples
were filtered (0.45 μm) within 24 h of collection (Millipore MFTM mem-
brane filter; Millipore, Billerica, MA) and frozen until analysis. Samples
were analyzed for reactive phosphorus using the ascorbic acid method
(Murphy and Riley, 1962), for NH+ 4 using the phenol-hypochlorite reaction
(Weatherburn, 1967), and for NO− 3 using a spectrophotometric method
(Doane and Horwath, 2003). All methods for determining nutrient concen-
trations were adapted for a microplate reader (Biotek Synergy H1 Hybrid
Microplate reader; Biotek, Winooski, VT). Dissolved organic carbon
(DOC) and total dissolved nitrogen (TN) were determined using a coupled
Shimadzu TOC-VCPH-TNM-1 analyzer. Nutrient concentrations are reported
in units of mg L−1 of the target element, C, N or P. Appropriate calibration
curves (regarding the range of concentrations expected) were used for each
nutrient and an independent QC check was used in each analysis. If samples
fell outside the standard curve, samples were rerun with a new calibration
curve. Additionally, seasonal deployments of calibrated MiniDOT tempera-
ture and oxygen loggers (PME, Vista, CA) were deployed for several days at
5 locations within the streams corresponding to 1 upstream, 3 culvert sites,
and 1 downstream site (Fig. 2). Logging intervals for these continuous de-
ployments was set to 10 min.
C.L. Hintz et al.
2.4. Periphyton biomass (chlorophyll a and ash-free dry mass)
Deployments of artificial substrate (two 10 cm × 10 cm unglazed ce-
ramic tiles attached to a concrete paver, Hauer and Lamberti, 2007) were
used to determine periphyton biomass (N = 13 sites per stream, Fig. 2). Ar-
tificial substrate was used to control for the variability of substrate between
open and culverted reaches (Beaulieu et al., 2014). From July 2017 to Au-
gust 2018, artificial substrate was deployed every two months and collected
after 4-week incubations. Samples were processed for chlorophyll a and
biomass (ash-free dry mass, AFDM). In the field, 2 replicate periphyton
samples were collected from each paver (i.e., 1 replicate per tile). An area
of 9–11 cm2 per tile was scraped and the resultant slurry was transferred
to a 500 mL HDPE bottle. Samples were then diluted with deionized
water to a known volume and a known volume of subsample was filtered
onto two 25-mm diameter, 0.45-μm Whatman GF/F filters (one for chloro-
phyll a and one for AFDM; Wallace et al., 2006). AFDM filters were pre-
combusted and pre-weighed prior to filtering (Wallace et al., 2006). After
filtering, filters (both for chlorophyll and AFDM) were placed into labeled
aluminum foil packets and placed on ice for further processing.
Chlorophyll a samples were extracted in 95% ethanol for 24 h in dark
conditions (Nusch, 1980). After extraction, filters were removed, and the
samples were centrifuged at 1000g for 5 min (Arar, 1997) and 2 mL of liq-
uid was then used for spectrophotometric chlorophyll analysis. Periphyton
samples were read at 750 and 664 nm, then acidified with 0.1 mL of 0.1 N
hydrochloric acid and reread at 750 and 665 nm (APHA, 2005) to account
for pheophytin. Chlorophyll a calculations were made using an ethanol ab-
sorption coefficient (Hansson, 1988). To estimate the amount of AFDM, fil-
ters were dried at 50 °C for 24 h, weighed for dry weight, then combusted
for 2 h at 500 °C to determine final ash weight (Hauer and Lamberti,
2007). For each sampling date, a blank filter was processed following the
same protocol for chlorophyll and AFDM to serve as a QC.
2.5. Macroinvertebrates (density and diversity)
Macroinvertebrates were collected with a D-net (dimensions: 12″ x 10″,
250-μm mesh) at 13 sites per stream once in May 2018. Since Congress Run
is seasonally intermittent, a spring collection date was chosen to ensure we
could collect invertebrates from both streams. The D-net was placed ap-
proximately 5 cm into the sediment and the sediment was disturbed for
0.5 m upstream of the net for 60 s, giving an equal sampling effort at
each site (Resh and Jackson, 1993). Samples were transferred into 1-L bot-
tles with distilled water, placed on ice, and returned to the lab. Within 24 h
upon returning to the lab, samples were washed onto a 250-μm sieve and
material >250 μm in size was preserved with 80% ethanol (Resh and
Hauer, 2017). Samples were subsequently sorted under a dissecting micro-
scope, and macroinvertebrates were identified to family (Cummins et al.,
2008).
2.6. Statistical analyses
To address the spatial extent of responses from stream burial, transition
zones were estimated following the methods described in Feijó-Lima et al.
Table 1
Model used to evaluate the dependence of a given ecosystem parameter on longitudinal distance, to determine transition responses at the entrance or exit of a
culvert. yD represents the stable downstream condition, yU, the stable upstream condition, d, is distance, x0 is the midpoint of the sigmoid in meters, and k is
the slope. The red boxes represent the transition zone for a given variable.
Model Function
yU−yD
Sigmoid logistic f ðdÞ ¼ yU−  −k
d
1þ x0
5
Science of the Total Environment 819 (2022) 153050
(2018). For a given variable, a transition zone is the distance through
which a variable shifts from an upstream stable condition in response to
an abrupt change, and how far the buffering effects of upstream conditions
can be detected (Table 1). Given that our stream reaches included two tran-
sitions (the entrance/presence of the culvert and then the exit/absence of
the culvert), we separated our data into two groups: (1) the upstream-
culvert transition and (2) the culvert-downstream transition. Transition dis-
tances were estimated for each group and variable separately.
To estimate transition distances and potential spatial lags, scatterplots
of all response variables were first created with longitudinal stream dis-
tance as the predictor variable. If a variable showed a monotonic response
to the presence/absence of the culvert, the variable was regressed against
distance using a sigmoidal model (Table 1) using the Levenberg-
Marquardt estimation method in the minpack.lm R package (Elzhov et al.,
2016). If a variable did not show a monotonic response, transition distances
were not estimated. A monotonic response was assessed qualitatively. As
our objectives were to (1) evaluate spatial variation within buried reaches,
(2) determine the magnitude of change upon entering and exiting a culvert,
and (3) to evaluate spatial lags in the response of various stream parameters
to the abrupt transitions of entering and exiting the culvert – a sigmoidal
model allowed us to address these objectives. For variables that were col-
lected bi-monthly (water depth, periphyton and macroinvertebrates), the
mean for each site was used in the regression. Variables were log-
transformed when necessary to meet the assumptions of the model (homo-
geneity of variances and normality). The sigmoidal model was tested
against the null model (y ~ 1) using a log-likelihood test and model as-
sumptions were tested using the nlsTools R package (Baty et al., 2015).
The sigmoidal model allowed us to estimate a stable upstream condition
(yU), stable downstream condition (yD), the location of the midpoint of
the change (x0, in m) and the slope of that change (k; Table 1). The maxi-
mum allowable slope (k) in the model was set to correspond to a complete
(90%) transition within the interval from one site to the next (i.e., over the
course of either 20 m or 30 m depending on the stream). Allowing values of
k above this level did not measurably change the model fit but increased un-
certainty in the k and x0 parameters spuriously. The resulting parameter es-
timates and their associated uncertainty were used as the input in the
inverse estimation methodology (described below) to calculate the length
of transition distances.
Transition distances were estimated separately for the upstream-culvert
and culvert-downstream transition. Transition distances were estimated
using inverse estimation in R with the investR package (Greenwell and
Schubert Kabban, 2014). Inverse estimation infers the possible values of
an explanatory variable (e.g., distance) given values of a response variable
based on a model's confidence interval (Feijó-Lima et al., 2018). For each
transition, the beginning and end of the transition were estimated. yU ±
SE and yD ± SE from the sigmoidal model were used as the starting values
in the inverse estimation to determine the beginning and the end of the
transition, respectively. Bootstrap simulations (1000) were run and the me-
dian value of the bootstrap estimate was used to determine xU (the distance
where the modeled response variable started to deviate from modeled sta-
ble upstream condition, yU) and xD (the distance where the modeled re-
sponse variable started to reach a stable downstream condition, yD).
Rise Decay
C.L. Hintz et al. Science of the Total Environment 819 (2022) 153050

There are limitations to this method and transition locations (xU and xD)
cannot be estimated when there are wide SEs surrounding yU and/or yD.
Lastly, a Spearman rank correlation analysis was performed to determine
the correlation between some of our biological parameters. All statistical
analyses were performed in R v. 3.6.2 (R Core Team, 2019). Data from
field datasheets were checked for data entry errors before analysis.
3. Results
3.1. Light
Light (as PAR) was present upstream and downstream of the culvert
while PAR levels dropped to 0 within the culvert (Table 2). Within Congress
Run, instantaneous measurements of light averaged 55 ± 20 μMol
m−2 s−1 (mean ± SE) in the upstream reach and averaged 72 ± 22
μMol m−2 s−1 in the downstream reach across sampling months. In
Princeton, instantaneous measurements of light averaged 113 ± 32 μMol
m−2 s−1 in the upstream reach and averaged 122 ± 36 μMol m−2 s−1 in
the downstream reach across sampling months. Our study design did not
allow us to quantitatively assess transitions in light (sampling sites were
too widely separated, streamside canopy varied, and sampling time of
day was not standardized), but within 20 m (Princeton) or 30 m (Congress
Run) into the culvert PAR levels had dropped to <0.1 μMol m−2 s−1.
3.2. Distribution of substrate size
In both streams, substrate size, as represented by D84%, was smaller
within the respective culverts than outside (Fig. 3a). In Congress Run, sub-
strate was predominantly cobble outside of the culvert and fine gravel
within the culvert (Table 2). The transition zone at the entrance of the cul-
vert was detected near the culvert entrance (Table 3) and the transition in
substrate at the culvert exit was detected within 30 m downstream of the
exit and extended beyond the end of the 90 m downstream study reach
(Fig. 4, Supplementary Table 1). In Princeton, substrate was predominantly
cobble outside of the culvert (both up- and downstream of the culvert) and
coarse gravel within the culvert (Table 2). The first transition zone in
Princeton was detected within the first 20 m of the culvert, and the begin-
ning of the second transition was detected approximately 20 m downstream
of the culvert exit (Fig. 4, Supplementary Table 1).
3.3. Water depth
In both streams, water depth increased as distance into the culverts in-
creased (Fig. 3b). At the culvert exits, deep pools were present that were
not observed anywhere else within our study reaches. In Congress Run,
mean depth upstream of the culvert was shallow (7 ± 2 cm; mean ± SE)
whereas within the culvert, mean depth was much greater (35 ± 2 cm)
(Table 2). Downstream of the culvert, mean depth became shallow again
(15 ± 3 cm). Habitat outside of the culvert (both up- and downstream)
consisted of riffles and runs, transitioning to pool habitat within the culvert.
For the upstream-culvert transition in Congress Run, a transition in depth
was detected within the first 30 m of culvert length and a new stable condi-
tion was detected within 60 m into the culvert (Fig. 4, Supplementary
Table 1). The transition zone at the culvert exit for depth was detected
within 30 m downstream of the culvert exit and a new stable condition
was detected within 30 m of the exit (Fig. 4, Supplementary Table 1).
In Princeton, mean depth upstream of the culvert was shallow (7 ±
1 cm) whereas within the culvert mean depth was greater (17 ± 3 cm)
(Table 2). Downstream of the culvert, mean depth decreased but only
slightly (13 ± 2 cm). Habitat upstream of the culvert consisted of riffles
and runs, transitioning to pool habitat within the culvert. Downstream of
the culvert, habitat consisted of pools, riffles, and runs. Depths within the
culvert were more variable across sampling months in Princeton when com-
pared to Congress Run (Fig. 3b). At the culvert entrance in Princeton, a tran-
sition zone for depth was detected within the first 20 m of the culvert
entrance and a new stable condition near the middle of the culvert
(Fig. 4, Supplementary Table 1). Upon exiting the culvert in Princeton, no
transition in depth could be detected within our 60 m downstream study
reach (p = 0.4253, log-likelihood test between null model and the sigmoi-
dal model; Table 3).
3.4. Water chemistry
Nutrient levels within Congress and Princeton (Table 2, Supplementary
Fig. 1) showed no spatial variation or relationship to culvert presence. In-
stantaneous measurements of temperature, dissolved oxygen, pH, and con-
ductivity did not vary with culvert presence (Table 2) and, therefore, no
transition zones were estimated for any water physiochemistry variables.
In Congress Run, dissolved oxygen ranged from 1.35–11.76 mg L−1, spe-
cific conductance ranged from 343 to 1084 μs cm−1, and stream pH ranged

Table 2
Summary table of raw data. Values are mean ± SE. Values are averaged across sampling dates and the entrances and exits of the culverts have been excluded. Temperature
refers to instantaneous point measurements while Diel Temperature Range refers to data from continuous short term deployments. Density and family richness refer to mac-
roinvertebrates.
Congress Run Princeton

Variable Upstream Culvert Downstream Upstream Culvert Downstream

Water Depth (cm) [max] 7 ± 2 [24] 35 ± 2 [60] 15 ± 3 [39] 7± 1 [15] 17 ± 3 [48] 13 ± 2 [30]
Substrate D18% (mm) 14 ± 5 4±1 7±1 12 ±2 8±3 7±2
Substrate D50% (mm) 36 ± 4 8±2 24 ± 3 41 ±4 15 ± 3 24 ± 11
Substrate D84% (mm) 73 ± 9 16 ± 4 85 ± 21 115 ± 13 39 ± 15 56 ± 18
Temperature (°C) (instantaneous) 18 ± 3 17 ± 3 15 ± 2 16 ±4 17 ± 4 15 ± 3
Diel Temperature Range (max - min °C) (continuous)
Fall 1.97 ± 0.182 0.67 ± 0.210 6.18 ± 3.43 4.30 ± 0.31 2.16 ± 0.30 2.59 ± 0.14
Spring 4.06 ± 1.05 – – 5.64 ± 0.65 5.32 ± 0.62 5.08 ± 0.60
Summer 0.79 ± 0.12 0.33 ± 0.13 6.39 ± 0.19 5.81 ± 1.07 4.82 ± 0.48 4.30 ± 0.04
pH 7.9 ± 0.1 8.0 ± 0.1 7.7 ± 0.1 8.2 ± 0.1 8.2 ± 0.1 8.1 ± 0.1
Conductivity (SPC) 653 ± 97 607 ± 108 693 ± 96 1104 ± 95 1079 ± 99 1163 ± 90
PAR (μMol m−2 s−1) 55 ± 20 0.0 ± 0.00 72 ± 22 113 ± 32 0.01 ± 0.01 122 ± 36
−1
NH+4 (mg-N L ) 0.03 ± 0.01 0.03 ± 0.02 0.03 ± 0.01 0.01 ± 0.01 0.01 ± 0.01 0.02 ± 0.01
NO−3 (mg-N L
−1
) 0.33 ± 0.09 0.42 ± 0.06 0.27 ± 0.08 0.20 ± 0.12 0.21 ± 0.11 0.22 ± 0.12
TN (mg L−1) 0.58 ± 0.13 0.69 ± 0.10 0.48 ± 0.12 0.42 ± 0.09 0.43 ± 0.10 0.47 ± 0.10
PO3−
4 (mg-P L−1) 0.07 ± 0.01 0.06 ± 0.01 0.06 ± 0.01 0.03 ± 0.01 0.03 ± 0.01 0.02 ± 0.01
DOC (mg L−1) 2.8 ± 0.4 3.1 ± 0.4 2.4 ± 0.6 3.1 ± 0.2 3.1 ± 0.2 3.1 ± 0.2
Chlorophyll (mg m−2) 9±3 0.4 ± 0.1 6±2 10 ± 3 0.2 ± 0.3 10 ± 3
AFDM (g m−2) 7±2 1 ± 0.2 8±1 7±1 1 ± 0.1 5±1
Density (individual m−2) 136 ± 71 5±3 229 ± 142 260 ± 42 31 ± 10 274 ± 165
Family Richness 1.7 ± 0.6 0.5 ± 0.3 3.2 ± 0.7 3.4 ± 0.6 1.8 ± 0.2 3.9 ± 0.9

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C.L. Hintz et al. Science of the Total Environment 819 (2022) 153050

Fig. 3. Substrate and water depth for Congress Run and Princeton. In both panels, the culverts are represented by the grey boxes, and vertical dashed lines are the culvert
entrance/exit. Flow direction is from left to right. a) Substrate size for D18% (circles), D50% (triangle), and D84% (square) percentiles b) Site water depth for each
stream and sampling date. The black line represents the mean water depth across all sampling dates. Note: the y-axis is reversed in panel b.

from 7.46–9.10 across all sampling months and sites. In Princeton, dis-
solved oxygen ranged from 7.29–12.75 mg L−1, specific conductance
ranged from 975 to 1357 μs cm−1, and pH ranged from 7.77–8.68 across
all sampling months and sites. Culvert sites had a smaller diel temperature
range during fall than the open upstream and downstream sites (Table 2).
3.5. Periphyton biomass – chlorophyll a
In both streams, chlorophyll collected from artificial substrates was
much lower within the culvert than in both the upstream and downstream
reaches (Fig. 5a, Table 2). In Congress Run chlorophyll decreased from 9 ±

Table 3
Parameter estimates for the stable upstream condition (yU), stable downstream condition (yD), the midpoint of the sigmoid (x0), and the slope of the sigmoid (k). P-values
represent the result of a log-likelihood test between the null model and sigmoidal model. If the sigmoidal model did not explain more variation than the null model, then no
sigmoidal model was fit. ND = stable condition not detected within our study reach. * = upper slope (k) limit was reached.
Variable yU ± SE yD ± SE x0 ± SE k ± SE p

Congress Run: Upstream - Culvert

Macroinvertebrate Family Richness – – – – 0.0919
Macroinvertebrate Density (individual m−2) – – – – 0.0502
AFDM (g m−2) 6.1 ± 0.8 1.1 ± 0.5 75 ± 56 21 ± 82.7* < 0.0005
Chlorophyll a (mg m−2) 11.9 ± 0.9 0.4 ± 0.6 57 ± 340 21 ± 3347.5* < 0.0005
Substrate D84% 82.2 ± 7.9 0 ± 16.1 85 ± 20 2.3 ± 1.1 < 0.0005
Water Depth (cm) 6.4 ± 3.4 41.3 ± 4.5 113 ± 11 10.5 ± 7.1 < 0.0005

Congress Run: Culvert - Downstream

Macroinvertebrate Family Richness 0.4 ± 0.27 4.9 ± 1.9 316 ± 26 13.5 ± 9.6 < 0.0005
Macroinvertebrate Density (individual m−2) 2±2 1795 ± 9296 314 ± 57 10.7 ± 11.9 0.0015
AFDM (g m−2) 1.1 ± 0.6 9.9 ± 1.5 285 ± 13 13.4 ± 6.8 < 0.0005
Chlorophyll a (mg m−2) 1.3 ± 0.42 ND ND 4.9 ± 8.3 0.0006
Substrate D84% 12.6 ± 8.7 306 ± 945 377 ± 220 9.6 ± 10.5 0.0001
Water Depth (cm) 34.7 ± 4.4 11.1 ± 8.1 298 ± 34 56 ± 984* 0.0289

Princeton: Upstream - Culvert

Macroinvertebrate Family Richness 3.4 ± 0.4 1.9 ± 0.4 60 ± 8 15.8 ± 155.2 0.0214
Macroinvertebrate Density (individual m−2) 260 ± 30 31 ± 24 58 ± 66 21 ± 676.7* < 0.0005
AFDM (g m−2) 7.4 ± 1.2 0.7 ± 1.2 75 ± 31 21 ± 124.8* 0.0028
Chlorophyll a (mg m−2) 10.4 ± 2.5 0 ± 2.6 64 ± 17 5.5 ± 8.1 0.0111
Substrate D84% 117.6 ± 17.6 22.6 ± 3.0 75 ± 16 21 ± 70.7* < 0.0005
Water Depth (cm) 7.4 ± 0.5 21.5 ± 1.2 102 ± 5 7.2 ± 2.0 < 0.0005

Princeton: Culvert - Downstream

Macroinvertebrate Family Richness – – – – 0.0592
Macroinvertebrate Density (individual m−2) 20 ± 9 182 ± 106 161 ± 28 12.6 ± 16.1 0.0076
AFDM (g m−2) 1.0 ± 0.8 4.9 ± 1.1 181 ± 26 56 ± 1102* 0.014
Chlorophyll a (mg m−2) 0.3 ± 1.3 9.8 ± 1.5 179 ± 22 56 ± 965* 0.0009
Substrate D84% 24.4 ± 6.5 67.4 ± 15.8 204 ± 29 56 ± 339* 0.0151
Water Depth (cm) – – – – 0.4253

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C.L. Hintz et al. Science of the Total Environment 819 (2022) 153050

Fig. 4. Transition zones detected for biotic and abiotic parameters in Congress Run and Princeton. In both panels, grey boxes represent the culvert and vertical dashed lines
are the culvert entrance/exit. Flow direction is from left to right. The start and end of the transition zones represent the medians from the inverse estimation method. Green
brackets represent the upstream-culvert transition while dark grey brackets represent the culvert-downstream transition. Horizontal dashed lines represent transitions where
the end of the transition could not be detected.

3 mg m−2 (mean ± SE) upstream of the culvert to 0.4 ± 0.1 mg m−2
within the culvert and recovered to 6 ± 2 mg m−2 downstream of the cul-
vert. In Princeton, chlorophyll decreased from 10 ± 3 mg m−2 upstream of
the culvert to 0.2 ± 0.3 mg m−2 within the culvert and recovered to 10 ±
3 mg m−2 downstream of the culvert. Chlorophyll remained consistently
low throughout the culverts as distance into the culverts increased. Chloro-
phyll upstream and downstream of the culverts showed considerable sea-
sonal variation in Congress Run and Princeton (Fig. 5a) while this
seasonal pattern was absent within the culverts.
Across sampling dates, in Congress Run, a decrease in chlorophyll
and a new stable condition were detected upstream of the culvert en-
trance (approximately 30–60 m upstream of the culvert entrance;
Fig. 4, Supplementary Table 1). This indicates a new stable condition
was detected upstream of the culvert and then was maintained for
most of the culvert length. The cause of the transition occurring up-
stream of the culvert is not known, but it could result from the inter-
mittent flow regime of Congress Run as some of the upstream sites
were dry at the time of sampling or, could represent a limitation of
our study given that we had set distances between sampling points
and it could be by chance that this particular site had lower periphy-
ton biomass. At the culvert exit, chlorophyll began to increase but a
downstream stable condition was not detected downstream within
our 90 m study reach (Table 3).
In Princeton, a chlorophyll transition was detected at the culvert en-
trance and a new stable condition was detected before the next sampling
point 20 m downstream (Fig. 4, Supplementary Table 1). For the culvert-
downstream transition in Princeton, a chlorophyll transition was detected
at the culvert exit (Fig. 4) and a new stable condition was detected by the
next sampling site (20 m downstream of the culvert exit).
3.6. Periphyton biomass - ash-free dry mass
In both streams, periphyton biomass (as AFDM) was present on incu-
bated artificial substrates at lower levels within the culverts than within
the open reaches up- and downstream of the culverts (Fig. 5b, Table 3). In
Congress Run, periphyton biomass decreased from 7 ± 2 g m−2 (mean ±
SE) upstream of the culvert to 1 ± 0.2 g m−2 within the culvert and recov-
ered to 8 ± 1 g m−2 downstream of the culvert. In Princeton, periphyton
biomass decreased from 7 ± 1 g m−2 upstream of the culvert to 1 ±
0.1 g m−2 within the culvert and recovered to 5 ± 1 g m−2 downstream
of the culvert. Periphyton biomass displayed no seasonal variation within
the culvert, whereas upstream and downstream reaches for both culverts
showed seasonal variation (Fig. 5).
In Congress Run, periphyton biomass began to decline 30 m upstream of
the culvert entrance and a new stable condition was detected as the stream
entered the culvert (Fig. 4). At the culvert-downstream transition in

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C.L. Hintz et al. Science of the Total Environment 819 (2022) 153050

Fig. 5. Concentrations of chlorophyll a (mg m−2; panel a) and ash-free dry mass (AFDM g m−2; panel b) on artificial substrate for each sampling date and location within the
study streams. In both panels, the culverts are represented by the grey boxes, the vertical dashed lines are the culvert entrance/exit, and the thick black line represents the
mean across all sampling dates. Flow direction is from left to right.

Congress Run, periphyton biomass began to increase upstream of the cul-
vert exit (within 30 m) and the transition ended 30–60 m downstream of
the culvert exit (Fig. 4, Supplementary Table 1). In Princeton, transitions
in periphyton biomass were detected at the culvert entrance (a decrease
in AFDM) and exit (an increase in AFDM). Both of these transitions were
relatively short and stable conditions were detected before the next down-
stream sampling location for each transition (20 m downstream of the
culvert entrance and 20 m downstream of the exit; Fig. 4, Supplementary
Table 1).
3.7. Macroinvertebrate density
In both streams, macroinvertebrates had lower densities inside the
cuvlert than upstream and downstream of the culverted reach (Fig. 6a;

Taxonomic Order

Diptera

Ephemeroptera

Oligochaetes

Other

Plecoptera

Trichoptera

Fig. 6. a) Mean macroinvertebrate density, b) Mean taxonomic family richness, and c) Proportion of sampled macroinvertebrate taxonomic orders a given site and stream.
Error bars in panels a) and b) represent the standard error. In all panels, the culverts are represented by the grey boxes and vertical dashed lines are the culvert entrance/exit.
Flow direction is from left to right.

9
C.L. Hintz et al.
Table 2). In Congress Run, macroinvertebrate density decreased from
136 ± 71 individuals m−2 (mean ± SE) upstream of the culvert to 5 ±
3 individuals m−2 within the culvert. At some culvert sites in Congress
Run, no macroinvertebrates were collected. Downstream of the culvert,
macroinvertebrate density increased to 229 ± 142 individuals m−2 in Con-
gress Run. The upstream-culvert transition was not modeled because our
hypothesis (i.e. the sigmoidal model) did not explain more variation than
the null model (p = 0.0502, log-likelihood test between null model and
the sigmoidal model; Table 3). Upon exiting the culvert in Congress Run,
a transition in macroinvertebrate density was detected at the culvert exit
and a stable condition was detected before the next downstream sampling
point (30 m downstream).
In Princeton, macroinvertebrate density decreased from 260 ± 42 indi-
viduals m−2 (mean ± SE) upstream of the culvert to 31 ± 10 individuals
m−2 within the culvert. Downstream of the culvert in Princeton, macroin-
vertebrate density increased to 274 ± 165 individuals m−2. In Princeton,
macroinvertebrate density decreased surrounding the culvert entrance
and a stable culvert condition was detected before the next downstream
sampling point (20 m downstream). For the culvert-downstream transition
in Princeton, macroinvertebrate density increased between 20 and 40 m
upstream of the culvert exit and a downstream stable condition was de-
tected near the culvert exit (Fig. 4, Supplementary Table 1). Macroinverte-
brate density in the Princeton culvert recovered to upstream levels before
the culvert exit (Table 2). Lastly, overall, macroinvertebrate densities in
both streams were correlated with periphyton biomass (as AFDM) (via
Spearman rank correlation, p < 0.005, r = 0.54).
3.8. Macroinvertebrate family richness
Macroinvertebrate diversity was low in both streams (Fig. 6b, Table 2)
regardless of stream reach and was primarily composed of Chironomidae
and Oligochaeta (Fig. 6c). The number of families present in the culvert
was lower than in reaches up- and downstream of the culverts (Fig. 6b).
An average of 0.5 and 1.8 families were found within the culvert in Con-
gress Run and Princeton, respectively and at some within-culvert sites in
Congress Run, no macroinvertebrates were found.
In Congress Run, macroinvertebrate family richness was higher up-
stream than within the culvert but our hypothesis (i.e., the sigmoidal
model) did not explain more variation than the null model (p = 0.09) so
the upstream-culvert transition was not modeled (Table 3). For the
culvert-downstream transition in Congress Run family richness began to in-
crease within 30 m downstream of the culvert exit and a stable downstream
level of family richness was detected by 60 m downstream of the culvert
exit (Fig. 4, Supplementary Table 1). In Princeton, transitions in macroin-
vertebrate diversity were detected at the culvert entrance (a decrease in di-
versity). This transition was short, and a stable level of macroinvertebrate
richness was detected before the next downstream sampling location for
each transition (20 m downstream of the culvert entrance) (Fig. 4, Supple-
mentary Table 1). Downstream of the culvert in Princeton, macroinverte-
brate family richness increased but our hypothesis (i.e., the sigmoidal
model) did not explain more variation than the null model (p = 0.42) so
the culvert-downstream transition was not modeled (Table 3).
4. Discussion
We aimed to directly characterize spatial variation in ecosystem param-
eters within buried stream reaches, determine the magnitude of change
upon entering/exiting a culvert, and to evaluate how different parameters
responded spatially to the abrupt transitions of entering and exiting the cul-
vert. Our data show that the within-culvert stream environment was consis-
tently altered across a suite of parameters including water depth, substrate
size, periphyton biomass, and macroinvertebrate density and taxa richness
(Figs. 3, 5, and 6). We also found that there was little spatial variation
within buried reaches, as a stable condition was detected within the cul-
verts for most variables. Culverts have been suggested as agents of fragmen-
tation in lotic systems (Fuller et al., 2015) and here we demonstrate that
10
Science of the Total Environment 819 (2022) 153050
burial indeed serves as an additional stressor on stream ecosystems poten-
tially leading to habitat fragmentation.
4.1. Spatial variation and magnitude of change within buried reaches
Our work highlights that even in degraded urban systems, culverts have
dramatic impacts on the structure and function of streams. Urban streams
are typically characterized by altered channel morphology and reduced bi-
otic richness (Walsh et al., 2005) and stream burial may serve as an addi-
tional stressor on urban ecosystems. We observed that substrate size,
periphyton biomass (chl a and AFDM), and macroinvertebrate abundances
decreased and remained low throughout the culverts in our study.
In-stream habitat can serve as a control on the presence and diversity of
biota and we observed that physical habitat was altered within the culverts.
Specifically, deeper water depths and smaller substrates were observed.
Water depth increased with distance into the culvert and the culverts eval-
uated in this study created pool-like environments at their downstream end.
These pools continued downstream of the culvert exits and pools of this
depth were not observed anywhere else within our studied stream reaches.
Based on our observation that pavers at within-culvert sites washed away
48% of the time during the study period compared with only 9% of pavers
from sites outside of the culverts, culverts had higher velocities within the
culverts during rain events. The physical habitat within our culverts repre-
sented a different habitat type than was observed elsewhere in our study
reaches and this was also reflected in the presence and diversity of biota
within the culverts.
Biotic communities were negatively impacted by the presence of the
culverts – specifically, we observed a loss of periphyton biomass and
lower macroinvertebrate density and taxa richness in culverts than in up-
stream/downstream reaches. The lack of light is an obvious limitation on
periphyton biomass production, and the availability of periphyton can be
a direct control on macroinvertebrates (Kiffney et al., 2004). Macroinverte-
brate communities can also be influenced by substrate size (Cummins et al.,
2008). Smaller-sized substrates provide less stable habitat and are nega-
tively correlated with macroinvertebrate density and diversity (Allan and
Castillo, 2007; Roy et al., 2003) and our study culverts were dominated
by these smaller substrates. Our upstream and downstream reaches had
wider distributions of substrate size and larger predominant substrate
which corresponded to increased macroinvertebrate density and diversity.
Similar relationships have been observed between substrate size and mac-
roinvertebrates in other urban streams (Roy et al., 2003; Wood and
Armitage, 1997).
In both streams, substrate size, periphyton biomass, and macroinverte-
brate abundances generally recovered downstream of the culvert to levels
similar to those observed upstream of the culvert. A new stable downstream
condition could be detected within our study reach (within 90 m for Con-
gress Run and within 60 m for Princeton) for most variables (but see
Fig. 4) suggesting the impacts from the culvert are spatially constrained
for the measured variables and at the spatial scale we monitored. The
downstream recovery of macroinvertebrates that we observed is encourag-
ing and suggests that the impact of culverts may be effectively limited to
within the culvert itself. However, the macroinvertebrate communities
were already impacted upstream of the culvert (i.e. low density and a com-
munity comprised of taxa indictive of poor water quality in these urban
streams (Alberts et al., 2018; Cummins et al., 2008; MBI, 2012) so it re-
mains to be seen if a more diverse macroinvertebrate community in a less
degraded stream would recover to the same extent downstream of the bur-
ied segment.
A suite of measured water chemistry variables (including dissolved N, P,
and DOC) did not change measurably upon entering the culverts or exhibit
any variation with distance into the culverts. This supports previous work
showing that culverts primarily serve as transporters of nutrients rather
than transformers (Beaulieu et al., 2014; Hope et al., 2014; Pennino et al.,
2014). In areas with high levels of urbanization and stream burial, culverts
may represent a missed opportunity for nutrient processing and thus con-
tribute to the downstream export of nutrients (Beaulieu et al., 2015).
C.L. Hintz et al.
4.2. Transition distances
To the best of our knowledge, this is the first detailed spatial character-
ization of within-culvert environments. We aimed to evaluate how various
stream parameters responded to the abrupt transitions of entering and
exiting the culvert and found that individual parameters responded some-
what differently. Estimated transition zone lengths from the inverse estima-
tion methodology ranged from 11 to 106 m for different parameters in
Congress Run and from 6 to 67 m in Princeton (Supplementary Table 1).
Our calculated transition zones were somewhat shorter than those previ-
ously reported in other stream ecosystem studies (described below), but
we are not aware of other studies that have examined transitions in re-
sponse to the entrance/exit of a culvert.
The abiotic parameters that displayed transitions in our systems were
substrate size and water depth. A change in substrate size is often associated
with riparian change in streams and has been previously detected on scales
of hundreds of meters in other regions (Chakona et al., 2009; Feijó-Lima
et al., 2018; Goss et al., 2014). When a riparian corridor transitions from
forest to pasture, a shift to smaller substrates has been detected, attributed
to increased bank erosion (Chakona et al., 2009; Feijó-Lima et al., 2018).
The opposite pattern has been detected when land-use surrounding streams
transitions from agriculture to forest – with larger substrate detected down-
stream of the transition (Goss et al., 2014). In our culverted streams, we ob-
served both of these changes abruptly at the entrance and exit of the
culvert, respectively. Transitions in water depth have been observed in re-
sponse to riparian changes and over long distances (>500 m) (Goss et al.,
2014). As with substrate size, we observed 2 abrupt transitions in water
depth as the stream entered and exited the culverts.
We generally did not observe transitions in water chemistry and water
temperature in response to the presence/absence of a culvert. Transitions
in water chemistry have been observed in response to riparian transitions
in other settings (Chakona et al., 2009; Goss et al., 2014) and were expected
here as well due to the large reduction in periphyton biomass within the
culvert. Given that changes in stream water temperature have been de-
tected in less extreme conditions (e.g., entering long stretches of shade;
Rutherford et al., 2004; Storey and Cowley, 1997), we were surprised
that impacts on water temperature (via instantaneous point measurements)
were relatively minor within the culvert given the complete elimination of
solar radiation and slow water velocity.
Generally, macroinvertebrate density, richness, and periphyton bio-
mass were suppressed within 20–30 m of the stream entering a buried
reach, indicating that burial has an almost immediate effect on the biolog-
ical structure of urban streams. This effect persisted throughout the
length of the buried reach, demonstrating that burial serves as an addi-
tional stressor that can further limit an already depauperate urban stream
biological community. Periphyton biomass displayed abrupt transitions
(within 20–30 m, with the exception of the culvert-downstream transition
in Congress Run) as expected given the elimination of light in the cul-
verts. Previous work has shown that a change in macroinvertebrates can
be detected for hundreds of meters downstream before reaching a stable
condition in response to a riparian change (Chakona et al., 2009; Goss
et al., 2014; Storey and Cowley, 1997; Tanaka et al., 2015), but our mac-
roinvertebrate parameters had very short transition zones at the culvert
entrances and exits (with the exception of the upstream-culvert transition
in Congress Run). Our results do not match our a priori hypothesis that we
would observe changes in chlorophyll and AFDM first and then we would
observe changes in macroinvertebrate parameters with a spatial lag
downstream (consistent with bottom-up control; Goss et al., 2014;
Kiffney et al., 2004). Transitions in these parameters were detected be-
fore our second within-culvert site in each stream (i.e., 20 or 30 m into
the culvert) so we were unable to determine if our a priori hypothesis
was supported at a finer spatial scale. But, overall, macroinvertebrate
densities were correlated with periphyton biomass (as AFDM). The mac-
roinvertebrate communities were dominated by Chironomidae and
Oligochaeta which are primarily collector-gatherers feeding on deposited
fine organic matter (Cummins et al., 2008).
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Science of the Total Environment 819 (2022) 153050
For the upstream-culvert transition, we hypothesized that all transitions
would begin at or downstream of the culvert entrance. Unexpectedly, two
transitions were detected upstream of the culvert entrance (Congress Run,
AFDM and chlorophyll a). The cause of this is not known but it could result
from the intermittent flow regime of Congress Run as some of the upstream
sites were dry at the time of sampling or, could represent a limitation of our
study given that we had set distances between sampling points and it could
be by chance that this particular site had lower periphyton biomass.
4.3. Study limitations
The analysis and interpretation was practically constrained by the lim-
ited number of measurement locations (13 per stream), and the longitudi-
nal spacing of our study sites set the minimum extent over which we
could detect transitions. The spacing was set at 20–30 m as a balance of
practical considerations and our expectation of transitions in the 10s to
100 s of m distance. In fact, many of our measured parameters transitioned
quite abruptly, often in less than the interval between two consecutive sam-
pling locations. While this result is relevant and important and indicates
that in already-urbanized streams the ecosystem effects of culverts may
be relatively transient, we were not able to precisely estimate distances
over which many of the more abrupt transitions occur. Follow-up studies
would ideally use a study design with smaller spacing between sampling
points, as well as extending further up- and down-stream to ensure that
all transitions are captured fully. Finally, there were some potentially im-
portant ecosystem variables that were not directly measured in our study,
for instance stream velocity (Fig. 1), which could vary in response to culvert
presence.
4.4. Conclusions: implications for stream monitoring and restoration
Urbanization is projected to increase globally through 2050 (United
Nations, 2019) and increasing urbanization increases the likelihood of
stream burial (Napieralski and Carvalhaes, 2016). While we have evaluated
stream burial at a reach scale, our results suggest that the impact of burial
for our monitored parameters at a stream network scale could be signifi-
cant. This has also been suggested by previous work which evaluated the
impact of stream burial on nutrient processing (Beaulieu et al., 2015). We
have shown that culverts are altered habitats when compared to upstream
and downstream reaches, but they are not devoid of biota. Biotic parame-
ters (periphyton and macroinvertebrates) were reduced by the presence
of the culvert but recovered downstream of the culvert, suggesting that cul-
verts could represent promising places for restoration projects because the
within-culvert habitat is greatly degraded compared to upstream and
downstream reaches. Restoration projects are often completed on a reach-
size scale and culverts could represent ideal locations for these projects.
The United Nations has deemed 2021–2030 the Decade on Ecosystem Res-
toration (United Nations, 2021) and our work provides a timely and valu-
able starting point for how these types of systems could be impacted and
where potential improvements to habitat and biodiversity could be made.
Restoration projects in aquatic systems occur on a spectrum and restora-
tion in buried streams can range from daylighting (i.e. culvert removal) or
altering culvert design to increase connectivity and passage for aquatic or-
ganisms (e.g. installation of baffles or the implementation of “stream simu-
lation design”). While culvert removal may not be possible in all locations,
improvements to culvert design could be implemented. Past work has
shown that cost-sharing between transportation agencies and conservation
groups can increase both groups' return-on-investment when the replace-
ment of aging, failing, and/or under-sized culverts is paired with surface
road improvements and culvert improvements (Neeson et al., 2018).
While restoration projects would ideally be implemented at a
watershed-scale, large scale projects are often difficult to implement in
urban environments (Bernhardt and Palmer, 2011) because of existing in-
frastructure. Our data suggest that there is potential for measurable im-
provements for a suite of parameters where restoration (via culvert
removal, i.e., daylighting) is feasible. We recognize that restoration is
C.L. Hintz et al.
expensive and can have various goals (Bernhardt et al., 2005) and suggest
that restoration via daylighting should be considered a component of
watershed-scale restorations that can provide broader ecosystem services.
Further, we suggest that culverts should be included in monitoring
plans, especially if restoration is being considered, and should be sampled
where feasible and safe. Given that a new stable condition was observed
for most ecosystem parameters within the culverts, sampling at one or
two central locations could be sufficient to characterize the culvert environ-
ment if appropriate for the study design and research questions. Note that
this may not hold true for shorter culverts where transition zones could
occur over most of a culvert's length. Transitions were apparent in most
of our monitored variables, so managers should be aware of these potential
transition zones when choosing monitoring locations within the culvert.
CRediT authorship contribution statement
Chelsea L. Hintz: Conceptualization, Methodology, Software, Valida-
tion, Formal analysis, Investigation, Resources, Writing – original draft,
Writing – review & editing, Visualization, Funding acquisition, Project ad-
ministration. Michael T. Booth: Conceptualization, Methodology, Valida-
tion, Resources, Writing – review & editing, Visualization. Tamara A.
Newcomer-Johnson: Conceptualization, Methodology, Validation, Re-
sources, Writing – review & editing. Ken M. Fritz: Conceptualization,
Methodology, Validation, Resources, Writing – review & editing. Ishi
Buffam: Conceptualization, Methodology, Validation, Formal analysis, Re-
sources, Writing – review & editing.
Declaration of competing interest
The authors declare that they have no known competing financial inter-
ests or personal relationships that could have appeared to influence the
work reported in this paper.
Acknowledgements
We would like to thank the many students who helped conduct this
work, especially J. Goetz, E. Sanderson, C. Tran, M. Urbanic, and N. Willse
for their sustained help in the field and lab. We would like to thank R. Feijo-
Lima for guidance on the analysis. We would like to thank S. Jacobs from
the US EPA for managing access to Congress Run and for his help in getting
the project started. Finally, we would like to thank the US EPA-University of
Cincinnati Graduate Trainee Program, the University of Cincinnati Sigma
Xi Grant in Aid of Research, and the University of Cincinnati Wieman-
Benedict Award for funding. Although this work was reviewed by the
USEPA and approved for publication, it might not necessarily reflect official
Agency policy. Any use of trade, firm, or product names is for descriptive
purposes only and does not imply endorsement by the U.S. Government.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.scitotenv.2022.153050.
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