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Ecological Connectivity in
a Exurban environment
Using Delaware, Ohio as a case study
Table of Contents
Introduction
and background
Study objectives
Methods and Results Recommendations
and Conclusion
01 02
04
03
01
Introduction and Background
Introduction:
Setting the Scene
(City of Delaware 2020)
(Delaware County 2015)
Setting
the
Scene
(Both maps from Sears 1926b)
Natural vegetation of Ohio, tree canopy
Setting
the
Scene
(ODNR, 1998 and 2005)
Setting
the
Scene
(Everts 1866)
(Perrin & Battle 1880)
(Everts 1875)
Delaware
starts to
urbanize:
Contiguous
habitats get
broken up
into smaller
fragments
Setting
the
Scene
What
Delaware
looks like
today
Aerial shot from Google Earth, shows a
typical landscape configuration in
Delaware. To the bottom is a linear
suburban subdivision, adjacent to a small
habitat fragment with agricultural land to
the north of it and exurban development
everywhere in-between. This is on one of
the fringes of the city. To the right is what
Delaware looks like on its fringes where
there isn’t development consisting mostly
of a mix of fallow and active cropland.
(author photos)
Characterized
by
ongoing
fragmentation
Setting
the
Scene
Aerial shots from Google Earth, shows exurban fragmentation in several
areas of Delaware. (Left) A house sits on a farm lot in a rural residential zone
and (top) more fragmentation including the beginnings of another subdivision.
(author photo).
What
Delaware
looks like
today
Characterized
by
ongoing
fragmentation
Fragmentation
effects
Infrastructure failures
(author photos)
Biodiversity
loss and
extinctions
(author photos)
(Rosenberg et al 2019)
Connectivity defined
By Shannon Hauck, adapted from Firehock and Walker 2015
A large fragment or contiguous area that is broken up into
smaller pieces by fragmentation due to roads or other
human structures, results in species loss unless the
remaining fragments are connected.
Connectivity defined
By Shannon Hauck, adapted from Firehock and Walker 2015
That’s why connectivity
between these fragments,
especially the more isolated
ones, which can be miles
apart, is crucial to population
viability and stability.
Literature Review
Snapshot of Delaware from Google Earth
Literature Review
The main themes were:
Loss of connectivity by habitat fragmentation was the largest issue found in
the literature reviewed.
Several other studies asserted that the importance of wildlife and plant
community connectivity through varying types of movement is a vital
component of landscape structure.
The literature also revealed that the creation of an ecological network,
which is defined as an area with high connectivity with large reserves,
dedicated corridors and high quality stepping stone habitats are essential to
long term viability of Earth's ecosystems.
A complete review with sources can be found in Appendix 5 of my thesis.
02
Thesis Objectives
Thesis Objectives
Study aim: To perform an overlay spatial analysis with
ArcGIS to determine if structural connectivity is feasible for
a wide variety of wildlife, in a environment of rapid land
changes and fluxes, that is, different landscape elements
interacting with each other in dynamic ways across the
rural to urban continuum in a exurban environment using
Delaware, Ohio as a case study.
Thesis Objectives
(1) identify and prioritize ecological connectivity corridors for species, with respect to i) the
importance of identifying and connecting large reserves (intact habitat cores) and habitat fragments
(non-cores), ii) amid the challenge of ongoing fragmentation and iii) investigating the possibility to
utilize or convert existing vacant lots and other open spaces to increase biodiversity.
(2) to create a natural assets dataset with i) the identification of all open spaces regardless of
ownership, ii) a habitat suitability index for all parcels in the city's planning area, to include soil
classifications, geology, slope and landforms, land cover characteristics, tree canopy, vegetation,
disturbance regimes, land use, rivers, tributaries, wetlands and ponds, floodplain characteristics
and percent imperviousness iii) to be shared and used as the basis for the natural assets portion of
the new Parks and Recreation Master Plan, directed to be written by the 2020 Comprehensive Plan
update currently underway (City of Delaware 2020).
(3) this study aims to provide a structural connectivity baseline in which can serve as a jumping off
point for conservation managers and policy makers to perform more detailed functional
connectivity assessments based on focal species to further conservation objectives on the local and
regional scales.
03
Methods and Results
Methods
Remote sensing analysis of GIS overlays utilizing ArcMap 10.7.1, core and
non-core habitat delineation, habitat suitability scoring by geophysical land
analysis, habitat cost surface analysis and ecological corridor delineation
using Least Cost Path (LCP) was utilized in this study to fulfill study
objectives.
Methods
A Map Package for Ohio, derived from Esri’s Green
Infrastructure Initiative was used for habitat suitability and
ecological corridor delineation (connectivity) and included
the following layers: Forested Habitat Fragments, Habitat
Cost Surface and Intact Habitat Cores by Betweenness.
The map gives a statewide view of intact habitat and
conserved areas. The habitat cores shown were derived
using a model built by the Green Infrastructure Center Inc
(Firehock and Walker 2015) and adapted by Esri.
The full set of 20 layers used are detailed
in my thesis.
Methods
The layer was cropped to the Delaware Comprehensive Plan
Growth Boundary (City of Delaware 2020).
The map was tailored and cropped to this boundary and that
boundary was designated as the study area so layers shown
are only of that area with the exception of two habitat
fragments which together were designated as an intact
habitat core (found near the study area but sitting just
outside of it (Deer Haven and Havener Parks) due to its close
proximity to other cores.
Methods
Then the following actions were taken:
1. All open space was identified by county assigned parcel as a unit of measure
(Kaitsa 2020).
2. All semi-natural to natural habitat fragments were classified as either a core
habitat or non-core using the county assigned parcel as a unit of measure (Kaitsa
2020).
3. The resulting cores were tested for size and width requirements (at least 100
acres in size and greater than 200 meters wide) and then converted into unique
polygons (Firehock and Walker 2015; Green Infrastructure Center, 2020). The size
class for each core was assigned following methodology outlined by the Green
Infrastructure Center Inc.
Methods
4. The edge to interior ratio was calculated for all natural to semi-natural cores and
non-cores in acres. Edge/interior ratio represents the relative abundance of core
versus edge habitat.
5. Percentages of streams, wetlands, ponds or other waterbodies, shrub/scrub,
grass/herbaceous, tree canopy and imperviousness were calculated for each parcel
on the NLCDS 2016 tree canopy and land cover layers.
6. Buffers were created around the Olentangy River to reflect the 1000 foot vegetative
buffer outside of urban areas that is required due to its designation as a State Scenic
River (ORC 1547.2). Buffers of 100 feet were created around the river's tributaries for
the same reason.
Methods
7. I split out all parcels identified as open space, habitat fragments (core and non-
core) from the parcel fabric layer into a separate dataset resulting in two datasets,
namely an open space dataset and a built environment dataset.
8. Performed a habitat suitability index by ranking all parcels in both datasets
regardless of ownership with a weighted suitability score of 0 (low suitability) to 5
(high suitability) for habitat determined by current reference conditions on that
parcel.
To determine the suitability for preservation, restoration or conversion of land to a
natural state to improve ecological/habitat connectivity and biodiversity, a
systematic, multiple data point based suitability index-driven methodology was
employed similar to Firehock and Walker 2015; Crossman et al 2012; Parker et al
2008; Pilewski et al 2019 for all 13,021 parcels in the study area with a goal of
identifying and prioritizing ecological corridors and various typologies of open
space for habitat suitability.
Methods
This type of analysis is known as a GIS-based multicriteria decision analysis (GIS-MCDA)
(Malczewski 2006; Pullinger and Johnson 2010). GIS is often recognized as a ‘decision
support system involving the integration of spatially referenced data in a problem solving
environment’ (Cowen 1988).
GIS-MCDA can be thought of as a process that combines geographical data and value
judgments for decision making. As such, with any expert opinion in any discipline, there
lies an inherent scientific reliability or validity error rate or level of uncertainty (Fischhoff
and Davis 2014).
Data layers in ArcMap 10.7.1 were also utilized to assign a habitat suitability score (using
the parcel as a unit of measure) related to the perceived ecological value of each parcel
also following a methodology outlined by the Green Infrastructure Center Inc. Habitat
suitability was measured because greater diversity is frequently associated with better
habitat potential. And areas of higher quality habitat need to be identified to create
stepping stone habitat for increased connectivity.
Methods
I utilized the Cost Distance Tool across a raster Habitat Cost Surface for the study area in
ArcMap to delineate possible ecological corridors. The Cost Distance tool in the Spatial
Analyst toolbox is used to calculate the effective distance, or the cumulative cost of moving
across a distance through varying habitat quality.
Intact Habitat Cores were considered as nodes (source or terminal patches) and the
connections between nodes (riparian ravines, vegetated riparian zones and habitat
fragments (patches) as edges (Urban and Keitt 2001).
This cost surface was used to identify both the permeability of the landscape matrix and then
a Least Cost Path (LCP) network was created using the habitat cores and non-cores in the
study area as well as two larger cores which sit just outside the study area (Delaware and
Alum Creek State Parks) to gauge long distance and close distance permeability and
connectivity.
Corridor/connectivity delineation and selection
Results
The results overwhelmingly showed that terrestrial wildlife, especially
interior specialists are most likely not able to move through the
landscape making connectivity on a large scale not feasible as resistance
and disturbance regimes are high in the surrounding matrix further
complicating any kind of potential terrestrial connectivity.
Larger (over 1,000 acres) intact habitat cores are located well outside of
the study area.
Results
The remaining forested habitat
fragments are widely scattered
among the human dominated
matrix and are mostly small,
isolated and are of high to poor
quality depending on their location,
composition and configuration.
However, connectivity for avian
species is possible through
restoration of these existing
fragments and creation of new
stepping stone habitat within the
urban, suburban, exurban
landscape. (Above) Habitat fragments are
widely dispersed throughout the
study area.
ArcMap 10.7.1
Results
Of the fragments delineated, six
larger natural areas were
identified as habitat cores.
Those cores are Camp
Lazarus/Seymour Woods State
Nature Preserve, Deer
Haven/Havener parks, Stratford
Ecological Center, the future Terra
Alta development (Ryan Homes),
Hickory Woods Park and Gallant
Woods Park.
(Above) Six larger natural areas
were identified as intact habitat
cores.
Camp
Lazarus/
Seymour
Woods
ArcMap 10.7.1
Results
All of them except for Gallant Woods
were found to be concentrated in the
southern portion of the study area
which was also found to be the area
with more geophysical variety and
higher permeability due to lower
levels of disturbance, low density
development and a low number of
intersecting road barriers.
Geophysical variety equals higher
diversity (Anderson et al. 2013). The
area with the highest geophysical
variety is the more permeable area
most of the cores are located.
Camp Lazarus/Seymour Woods
Deer
Haven/
Havener
ArcMap 10.7.1
Results
The habitat suitability analysis revealed:
1. That these cores were classified as small
cores with an average size of 237.5
acres with an average edge/interior
ratio of 58.7 meaning that there is slightly
more interior to edge for all the cores.
(Higher values in the edge/core ratio indicate more
fragmentation in which the patch contains more edge than
interior habitat so those fragments scored lower (Soverel et
al 2010)).
1. Have an average tree canopy of 74
percent, contain an average of 39
percent water (wetlands, ponds, streams,
vernal pools) and contain 11 percent
grassland (savanna ) or prairie and- (Above) Six larger natural
areas were identified as
intact habitat cores.
Camp
Lazarus/
Seymour
Woods
ArcMap 10.7.1
Results
The habitat suitability analysis revealed:
3. The average distance between the
cores is 28,894 feet and average
distance between the cores and the next
largest non-core fragment is 6,312 feet
showing that the smaller non-core
fragments are important to create and
maintain connectivity (see thesis for
more detail on each).
(Above) Six larger natural
areas were identified as intact
habitat cores.
Camp
Lazarus/
Seymour
Woods
ArcMap 10.7.1
Results
The habitat suitability analysis
revealed:
4. Most of the most suitable
areas among the delineated
open spaces is located mostly
in the study area’s fringes with
scattered high quality habitat
throughout the urban,
suburban, exurban and
agricultural zones.
(The darker colors indicate higher
quality).
(Above) Open space delineation and habitat suitability index.
ArcMap 10.7.1
Results
The habitat suitability analysis
revealed:
5. Most of the most suitable
areas among the delineated
built environment is scattered
throughout with the most
suitable parcels being private
HOA parks and institutional
lands to the south and north of
the city’s urban core.
(The darker colors indicate higher
quality).
(Above) Built environment delineation and habitat suitability
index.
ArcMap 10.7.1
Results
Habitat cost surface of the study area
shows high resistance throughout the
study area especially in the urban core
(red) and in the suburban, exurban and
rural areas (yellow).
The Habitat Cost Surface shows that
there is high resistance in the central
core of the city (red), typical of all urban
settlements. The built environment is
very dense in that area and has high
disturbance regimes (high cumulative
human modification over time). The
yellow is mostly suburban, exurban and
agricultural uses and the green equates
to most of the semi-natural habitat
fragments.
ArcMap 10.7.1
(Right) Habitat Cost Surface.
Results
The regional Least Cost Paths shows
low to moderate permeability to the
east and west for terrestrial
movement to Delaware State Park to
the north from the more permeable
area to the south, however, both
potential ecological corridors have
many barriers, shown in red, where
they pass through the periphery of
the urban-exurban matrix.
Groundtruthing (visual survey)
reveals that these corridors are not
feasible for ground movement
especially for interior species.
ArcMap 10.7.1
(Right) Habitat Cost Surface
with LCP Paths.
Results
The results also revealed a higher
permeability to the south mostly due
to its rural nature and large number
of protected areas.
The corridors delineated by the LCP
analysis have high conduciveness for
movement in between Camp
Lazarus/Seymour Woods and Deer
Haven/Havener parks but less so
with the core to the north (Stratford
Ecological Center) because the
riparian zone is not as permeable
due to roads and development (the
red line in the middle of the figure).
ArcMap 10.7.1
(Above) Local LCP paths shown in black between the three
small intact habitat cores in the southern portion of the study
area.
Results
Moderate permeability was found
to exist within the agricultural
zones to the west, east and north
but those are mostly open lands
bisected by several road barriers
with isolated woodlots and
disjointed hedgerows offering low
connectivity in its current state
other than along the riparian
zones of some less urbanized
north-south rivers and streams.
ArcMap 10.7.1
(Above) LCP paths shown in black through the isolated small intact
habitat core (Gallant Woods) and the northern portion of the study
area leading to the closest larger intact habitat core (Delaware
State Park). the park and its adjacent wildlife area is 6,356 acres
and is classified as a medium habitat core using methodology from
the Green Infrastructure Center, Inc.
04
Recommendations and Conclusion
Recommendations
The best solutions revolve around how green spaces of all types and sizes can
be best configured to increase connectivity in a complex exurban environment.
Landscapes should be integrated at multiple scales by connecting semi-
natural areas and developed open spaces (lawn-based parks, the city
cemetery, road verges, road right of ways, utility right of ways, vacant lots,
infill areas, paved greenway buffer zones, tree lawns/street trees, larger
residential yards) for multifunctional connectivity between large intact habitat
cores to increase diversity (especially among more mobile species like birds
and pollinators), making a landscape less sensitive to human induced or
climatic disturbances by building its capacity for resilience.
Ways to increase connectivity
Conservation and restoration of the city’s riparian ravines - The city’s urban
and rural ravines (both protected and unprotected) provide the best
opportunity for connectivity corridors between the identified terminal intact
habitat cores or source patches in the study area if restored. The city of
Toronto, Canada which has similar characteristics as the study area has
created a ravine strategy that Delaware could look to for the management of
its ravines (City of Toronto 2020).
Ways to increase connectivity
Utilizing vacant lots as stepping stone habitat - The creation of habitat in open
spaces and vacant lots for more mobile species - mainly birds, bats and
pollinating insects can be important reservoirs for species richness or
diversity (Burghardt et al 2008; Villasenora et al 2020). For example, semi-
natural vacant lots were found to harbor as many as 60 songbird species in a
recent study in Baltimore (Brodsky 2016).
Ways to increase connectivity
The Northeast Ohio Ecological Consortium (NEOECO) has developed a Vacant
Land Rapid Assessment Procedure (VL-RAP) that provides an efficient way to
screen and evaluate sites using basic ecological and ecosystem principles to
determine their potential suitability as wildlife habitat, stormwater
management, parks, and gardens (Rouse and Bunster-Ossa 2013; Cleveland
Urban Design Collaborative 2008).
Ways to increase connectivity
Utilizing road verges, rights of way and other open spaces to form stepping
stone habitat - Planting pollinator pathways, micro prairies, bioswales, rain
gardens and other green infrastructure can increase plant and animal
biodiversity exponentially depending on which solution is utilized.
Solutions and incentives for cities to improve connectivity through the
adequate use of green infrastructure to address gaps are many. Solutions
range from food forests and micro prairies to wetland parks.
Side swale, Columbus (photos by Kevin Parks, ThisWeekNews)
Examples of Green Infrastructure
(Left) Bioretention planter, Dublin, Ohio. (Right) Bioretention cell near busy four-lane road near the
Scioto River, Upper Arlington, Ohio.
(photos by the author)
Examples of Green Infrastructure
(Left) Pollinator pathway interspersed in
between a large solar array, Newark, Ohio.
(Above) A floodable or sponge park, Powell, Ohio.
(Solar photo by the author; Floodable park by
Jenna Odegard, MAD Scientist Associates)
Examples of Green Infrastructure
Bioretention pond converted from a traditional stormwater retention pond to a wetland bioretention pond, Granville
Land Lab, Granville, Ohio.
(photos by the author)
Examples of Green Infrastructure
Land sharing and land sparing on agricultural lands
More than 100 acres prairie, grass savanna, wetlands, an oak-hickory forest edge and a small
grassy riparian zone on former intensively farmed cropland at the Granville Land Lab, Granville,
Ohio. (photos by the author)
Conclusion
Cities like Delaware can play a fundamental role in the conservation of biodiversity
through strategies that include the introduction of ecosystems and habitats into the
urban fabric, or the preservation of pre-existing ones, as well as the creation of
continuous urban green spaces that guarantee biological connectivity and control
continuing fragmentation indicative of exurban settlements.
It is hoped that the findings generated by this baseline structural connectivity and
suitability analysis, can be used as a jumping off point for further comprehensive
conservation efforts especially in light of continued population growth, rapid
conversion of cropland to development, aging infrastructure concerns and continued
habitat loss through fragmentation among other factors.
Further Research
Further research could be in the form of:
● Some type of functional connectivity assessment or assessments which is the ability of an
area to meet the food and cover requirements for a specific animal species (Juntti and
Rumble, 2006).
● A City Biodiversity Index (CBI). The CBI, pioneered by the city of Singapore (Deslauriers et al
2018), could follow functional connectivity assessments to further identify and prioritize
the city of Delaware’s greenspace for stepping-stone habitat in addition to the prioritized
corridors identified in this study.
● And bioblitzes and other similar citizen science initiatives to document the diversity and
distribution of terrestrial and avian species especially during migration.
These follow-on studies can be performed or hosted by Ohio Wesleyan University students
in the Environmental Science and Geography programs as each student needs to do a capstone
in the local area as part of their graduation requirements.
References
Text References
● Rosenberg K, Dokter A , Blancher P , Sauer J , Smith A , Smith P , Stanton J , Panjabi A , Helft L , Parr
M and Marra P., Decline of the North American Avifauna, Science, 366:6461, pp 120 (2019).
● City of Toronto, Ravine Strategy Implementation Report, 66 pgs (January 2020).
● Burghardt K, Tallamy D and Shriver G., Impact of Native Plants on Bird and Butterfly Biodiversity in
Suburban Landscapes, Conservation Biology, Volume 23, No. 1, 219–224 (2008).
● Villaseñora N, Chianga L, Hernándeza HJ and Escobar M., Vacant lands as refuges for native birds:
An opportunity for biodiversity conservation in cities, Urban Forestry & Urban Greening, 49 (2020).
● Brodskey C., Urban Vacant Lots: Ecological Quality and Social Opportunities in Baltimore,
Maryland, A Dissertation Presented to The Faculty of the Graduate School at the University of
Missouri-Columbia (2016).
● Rouse and Bunster Ossa, Green Infrastructure: A Landscape Approach, 144 pgs (2013). Print.
● Cleveland Urban Design Collaborative, Resilient Cleveland, online report, (2008).
https://www.cudc.kent.edu/resilient-cleveland
● Firehock, K, and Walker RA., Strategic Green Infrastructure Planning: A Multi-Scale Approach
(2015) Print.
● Maps throughout this thesis were created using ArcGIS® software by Esri. ArcGIS® and
ArcMap™ are the intellectual property of Esri and are used herein under license. Copyright © Esri.
All rights reserved. For more information about Esri® software, please visit www.esri.com.
References
Text References
● Anderson MG., Clark M and Sheldon A., Estimating Climate Resilience for Conservation across
Geophysical Settings, Conservation Biology, 28:4, pp 959–970 (2013).
● Taylor, P.D., Fahrig, L. Henein, K. and Merriam,G., Connectivity is a vital element of landscape
structure, Oikos 68:3, pp 571-572 (1993).
● Soverel NO, Coops NC, White JC and Wulder MA, Characterizing the Forest Fragmentation of
Canada’s National Parks, Environmental Monitoring and Assessment, 164:14, pp 481–499 (2010).
● Theobald DM, Landscape Patterns of Exurban Growth in the USA from 1980 to 2020, Ecology and
Society, 10:1, p 32 (2005).
● Lindenmayer D and Hobbs R., eds., Managing and Designing Landscapes for Conservation: Moving
from Perspectives to Principles, Blackwell Publishing, 608 pgs (2007). Print.
● Hansen A, Knight R, Marzluff J, Powell S, Brown K, Gude P and Kingsford J, Effects of exurban
development on biodiversity: Patterns, mechanisms and research themes, Ecological Applications,
15:6, pp 1893–1905 (2005).
● Taylor P, Fahrig L and With K., Landscape Connectivity: A Return to the Basics, Cambridge
University Press, pp 29-43 (2006).
● Tischendorf, L. and Fahrig, L. How should we measure landscape connectivity? Landscape Ecology
15, 633–641 (2000).
● Malczewski J., GIS-based multicriteria decision analysis: a survey of the literature, International
Journal of Geographical Information Science, 20:7, pp 703-726 (2006).
References
Text References
● Pullinger, M.G., Johnson, C.J., Maintaining or restoring connectivity of modified landscapes:
evaluating the least-cost path model with multiple sources of ecological information, Landscape
Ecology, 25, 1547–1560 (2010).
● Fischhoff B and Davis A., Communicating scientific uncertainty, PNAS, p 111 (2014).
● Urban D and Keitt T., Landscape Connectivity: A Graph-Theoretic Perspective, Ecology 82:5, pp
1205-1218 (2001).
● Egan J and Mortensen D., A comparison of land-sharing and land-sparing strategies for plant
richness conserva-tion in agricultural landscapes, Ecological Applications, 22:2, pp 459-471
(2012).
● Balmford B, Green R, Onial M, Phalan B., and Balmford A., How imperfect can land sparing be
before land shar-ing is more favorable for wild species?, Journal of Applied Ecology, 56:1, pp 73-
84 (2019).
● Juntti TM and Rumble M., Arc Habitat Suitability Index computer software, Gen. Tech. Rep. RMRS-
GTR-180WWW. Ft. Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain
Research Station. 31 pages (2006).
● Kaitsa, George, Delaware County Auditor, County agricultural land use statistics. Retrieved 20
August 2020.
● City of Delaware, Comprehensive Plan 2003 – 2008, Delaware Ohio, Department of Planning and
Community Development (2002).
● Ohio Revised Code, Title [15] XV State Scenic River Designation, Chapter 1547.81, online resource
(accessed 17 March 2020) http://codes.ohio.gov/orc/1547.81v1.
References
Graphics and maps
● Eastern Forest and Migratory Bird Decline Graphics, Rosenberg K, Dokter A , Blancher P , Sauer J ,
Smith A , Smith P , Stanton J , Panjabi A , Helft L , Parr M and Marra P., Decline of the North
American Avifauna, Science, 366:6461, pp 120 (2019).
● Side Swale, Columbus, Parks KP, Clintonville Rain Gardens Net Mixed Results, ThisWeek
Community Newspapers (July 2015).
● Floodable park, Powell, Odegard J, MAD Scientists Associates (wetland scientists), Westerville,
Ohio (2020).
● Delaware County Highway Map, Chris Bauserman, Delaware County Engineer (2015).
● Draft City of Delaware Comprehensive Plan Land Use Map (2021).
● Physiographic Regions of Ohio and Glacial Map of Ohio, ODNR (1998 and 2005) respectively.
● Natural Vegetation of Ohio maps, Sears PB, The natural vegetation of Ohio, III. Plant succession,
Ohio Journal of Science, 26, pp 213-31 (1926b).
● State of Ohio Historical Maps and City of Delaware Plat Maps, Everts LH, A Illustrated Historical
Atlas of Delaware County, Ohio, LH Everts and Co., Philadelphia (1866).
● Satellite shots taken from Google Earth Pro (2020).
● Original patch dynamic art by Shannon Hauck based on Firehock and Walker 2015).
● Figures created by author out of ArcMap 10.7.1
● Original photos taken by author.
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Ecological Connectivity in an Exurban Environment

  • 1. Ecological Connectivity in a Exurban environment Using Delaware, Ohio as a case study
  • 2. Table of Contents Introduction and background Study objectives Methods and Results Recommendations and Conclusion 01 02 04 03
  • 4. Introduction: Setting the Scene (City of Delaware 2020) (Delaware County 2015)
  • 5. Setting the Scene (Both maps from Sears 1926b) Natural vegetation of Ohio, tree canopy
  • 7. Setting the Scene (Everts 1866) (Perrin & Battle 1880) (Everts 1875) Delaware starts to urbanize: Contiguous habitats get broken up into smaller fragments
  • 8. Setting the Scene What Delaware looks like today Aerial shot from Google Earth, shows a typical landscape configuration in Delaware. To the bottom is a linear suburban subdivision, adjacent to a small habitat fragment with agricultural land to the north of it and exurban development everywhere in-between. This is on one of the fringes of the city. To the right is what Delaware looks like on its fringes where there isn’t development consisting mostly of a mix of fallow and active cropland. (author photos) Characterized by ongoing fragmentation
  • 9. Setting the Scene Aerial shots from Google Earth, shows exurban fragmentation in several areas of Delaware. (Left) A house sits on a farm lot in a rural residential zone and (top) more fragmentation including the beginnings of another subdivision. (author photo). What Delaware looks like today Characterized by ongoing fragmentation
  • 10. Fragmentation effects Infrastructure failures (author photos) Biodiversity loss and extinctions (author photos) (Rosenberg et al 2019)
  • 11. Connectivity defined By Shannon Hauck, adapted from Firehock and Walker 2015 A large fragment or contiguous area that is broken up into smaller pieces by fragmentation due to roads or other human structures, results in species loss unless the remaining fragments are connected.
  • 12. Connectivity defined By Shannon Hauck, adapted from Firehock and Walker 2015 That’s why connectivity between these fragments, especially the more isolated ones, which can be miles apart, is crucial to population viability and stability.
  • 13. Literature Review Snapshot of Delaware from Google Earth
  • 14. Literature Review The main themes were: Loss of connectivity by habitat fragmentation was the largest issue found in the literature reviewed. Several other studies asserted that the importance of wildlife and plant community connectivity through varying types of movement is a vital component of landscape structure. The literature also revealed that the creation of an ecological network, which is defined as an area with high connectivity with large reserves, dedicated corridors and high quality stepping stone habitats are essential to long term viability of Earth's ecosystems. A complete review with sources can be found in Appendix 5 of my thesis.
  • 16. Thesis Objectives Study aim: To perform an overlay spatial analysis with ArcGIS to determine if structural connectivity is feasible for a wide variety of wildlife, in a environment of rapid land changes and fluxes, that is, different landscape elements interacting with each other in dynamic ways across the rural to urban continuum in a exurban environment using Delaware, Ohio as a case study.
  • 17. Thesis Objectives (1) identify and prioritize ecological connectivity corridors for species, with respect to i) the importance of identifying and connecting large reserves (intact habitat cores) and habitat fragments (non-cores), ii) amid the challenge of ongoing fragmentation and iii) investigating the possibility to utilize or convert existing vacant lots and other open spaces to increase biodiversity. (2) to create a natural assets dataset with i) the identification of all open spaces regardless of ownership, ii) a habitat suitability index for all parcels in the city's planning area, to include soil classifications, geology, slope and landforms, land cover characteristics, tree canopy, vegetation, disturbance regimes, land use, rivers, tributaries, wetlands and ponds, floodplain characteristics and percent imperviousness iii) to be shared and used as the basis for the natural assets portion of the new Parks and Recreation Master Plan, directed to be written by the 2020 Comprehensive Plan update currently underway (City of Delaware 2020). (3) this study aims to provide a structural connectivity baseline in which can serve as a jumping off point for conservation managers and policy makers to perform more detailed functional connectivity assessments based on focal species to further conservation objectives on the local and regional scales.
  • 19. Methods Remote sensing analysis of GIS overlays utilizing ArcMap 10.7.1, core and non-core habitat delineation, habitat suitability scoring by geophysical land analysis, habitat cost surface analysis and ecological corridor delineation using Least Cost Path (LCP) was utilized in this study to fulfill study objectives.
  • 20. Methods A Map Package for Ohio, derived from Esri’s Green Infrastructure Initiative was used for habitat suitability and ecological corridor delineation (connectivity) and included the following layers: Forested Habitat Fragments, Habitat Cost Surface and Intact Habitat Cores by Betweenness. The map gives a statewide view of intact habitat and conserved areas. The habitat cores shown were derived using a model built by the Green Infrastructure Center Inc (Firehock and Walker 2015) and adapted by Esri. The full set of 20 layers used are detailed in my thesis.
  • 21. Methods The layer was cropped to the Delaware Comprehensive Plan Growth Boundary (City of Delaware 2020). The map was tailored and cropped to this boundary and that boundary was designated as the study area so layers shown are only of that area with the exception of two habitat fragments which together were designated as an intact habitat core (found near the study area but sitting just outside of it (Deer Haven and Havener Parks) due to its close proximity to other cores.
  • 22. Methods Then the following actions were taken: 1. All open space was identified by county assigned parcel as a unit of measure (Kaitsa 2020). 2. All semi-natural to natural habitat fragments were classified as either a core habitat or non-core using the county assigned parcel as a unit of measure (Kaitsa 2020). 3. The resulting cores were tested for size and width requirements (at least 100 acres in size and greater than 200 meters wide) and then converted into unique polygons (Firehock and Walker 2015; Green Infrastructure Center, 2020). The size class for each core was assigned following methodology outlined by the Green Infrastructure Center Inc.
  • 23. Methods 4. The edge to interior ratio was calculated for all natural to semi-natural cores and non-cores in acres. Edge/interior ratio represents the relative abundance of core versus edge habitat. 5. Percentages of streams, wetlands, ponds or other waterbodies, shrub/scrub, grass/herbaceous, tree canopy and imperviousness were calculated for each parcel on the NLCDS 2016 tree canopy and land cover layers. 6. Buffers were created around the Olentangy River to reflect the 1000 foot vegetative buffer outside of urban areas that is required due to its designation as a State Scenic River (ORC 1547.2). Buffers of 100 feet were created around the river's tributaries for the same reason.
  • 24. Methods 7. I split out all parcels identified as open space, habitat fragments (core and non- core) from the parcel fabric layer into a separate dataset resulting in two datasets, namely an open space dataset and a built environment dataset. 8. Performed a habitat suitability index by ranking all parcels in both datasets regardless of ownership with a weighted suitability score of 0 (low suitability) to 5 (high suitability) for habitat determined by current reference conditions on that parcel. To determine the suitability for preservation, restoration or conversion of land to a natural state to improve ecological/habitat connectivity and biodiversity, a systematic, multiple data point based suitability index-driven methodology was employed similar to Firehock and Walker 2015; Crossman et al 2012; Parker et al 2008; Pilewski et al 2019 for all 13,021 parcels in the study area with a goal of identifying and prioritizing ecological corridors and various typologies of open space for habitat suitability.
  • 25. Methods This type of analysis is known as a GIS-based multicriteria decision analysis (GIS-MCDA) (Malczewski 2006; Pullinger and Johnson 2010). GIS is often recognized as a ‘decision support system involving the integration of spatially referenced data in a problem solving environment’ (Cowen 1988). GIS-MCDA can be thought of as a process that combines geographical data and value judgments for decision making. As such, with any expert opinion in any discipline, there lies an inherent scientific reliability or validity error rate or level of uncertainty (Fischhoff and Davis 2014). Data layers in ArcMap 10.7.1 were also utilized to assign a habitat suitability score (using the parcel as a unit of measure) related to the perceived ecological value of each parcel also following a methodology outlined by the Green Infrastructure Center Inc. Habitat suitability was measured because greater diversity is frequently associated with better habitat potential. And areas of higher quality habitat need to be identified to create stepping stone habitat for increased connectivity.
  • 26. Methods I utilized the Cost Distance Tool across a raster Habitat Cost Surface for the study area in ArcMap to delineate possible ecological corridors. The Cost Distance tool in the Spatial Analyst toolbox is used to calculate the effective distance, or the cumulative cost of moving across a distance through varying habitat quality. Intact Habitat Cores were considered as nodes (source or terminal patches) and the connections between nodes (riparian ravines, vegetated riparian zones and habitat fragments (patches) as edges (Urban and Keitt 2001). This cost surface was used to identify both the permeability of the landscape matrix and then a Least Cost Path (LCP) network was created using the habitat cores and non-cores in the study area as well as two larger cores which sit just outside the study area (Delaware and Alum Creek State Parks) to gauge long distance and close distance permeability and connectivity. Corridor/connectivity delineation and selection
  • 27. Results The results overwhelmingly showed that terrestrial wildlife, especially interior specialists are most likely not able to move through the landscape making connectivity on a large scale not feasible as resistance and disturbance regimes are high in the surrounding matrix further complicating any kind of potential terrestrial connectivity. Larger (over 1,000 acres) intact habitat cores are located well outside of the study area.
  • 28. Results The remaining forested habitat fragments are widely scattered among the human dominated matrix and are mostly small, isolated and are of high to poor quality depending on their location, composition and configuration. However, connectivity for avian species is possible through restoration of these existing fragments and creation of new stepping stone habitat within the urban, suburban, exurban landscape. (Above) Habitat fragments are widely dispersed throughout the study area. ArcMap 10.7.1
  • 29. Results Of the fragments delineated, six larger natural areas were identified as habitat cores. Those cores are Camp Lazarus/Seymour Woods State Nature Preserve, Deer Haven/Havener parks, Stratford Ecological Center, the future Terra Alta development (Ryan Homes), Hickory Woods Park and Gallant Woods Park. (Above) Six larger natural areas were identified as intact habitat cores. Camp Lazarus/ Seymour Woods ArcMap 10.7.1
  • 30. Results All of them except for Gallant Woods were found to be concentrated in the southern portion of the study area which was also found to be the area with more geophysical variety and higher permeability due to lower levels of disturbance, low density development and a low number of intersecting road barriers. Geophysical variety equals higher diversity (Anderson et al. 2013). The area with the highest geophysical variety is the more permeable area most of the cores are located. Camp Lazarus/Seymour Woods Deer Haven/ Havener ArcMap 10.7.1
  • 31. Results The habitat suitability analysis revealed: 1. That these cores were classified as small cores with an average size of 237.5 acres with an average edge/interior ratio of 58.7 meaning that there is slightly more interior to edge for all the cores. (Higher values in the edge/core ratio indicate more fragmentation in which the patch contains more edge than interior habitat so those fragments scored lower (Soverel et al 2010)). 1. Have an average tree canopy of 74 percent, contain an average of 39 percent water (wetlands, ponds, streams, vernal pools) and contain 11 percent grassland (savanna ) or prairie and- (Above) Six larger natural areas were identified as intact habitat cores. Camp Lazarus/ Seymour Woods ArcMap 10.7.1
  • 32. Results The habitat suitability analysis revealed: 3. The average distance between the cores is 28,894 feet and average distance between the cores and the next largest non-core fragment is 6,312 feet showing that the smaller non-core fragments are important to create and maintain connectivity (see thesis for more detail on each). (Above) Six larger natural areas were identified as intact habitat cores. Camp Lazarus/ Seymour Woods ArcMap 10.7.1
  • 33. Results The habitat suitability analysis revealed: 4. Most of the most suitable areas among the delineated open spaces is located mostly in the study area’s fringes with scattered high quality habitat throughout the urban, suburban, exurban and agricultural zones. (The darker colors indicate higher quality). (Above) Open space delineation and habitat suitability index. ArcMap 10.7.1
  • 34. Results The habitat suitability analysis revealed: 5. Most of the most suitable areas among the delineated built environment is scattered throughout with the most suitable parcels being private HOA parks and institutional lands to the south and north of the city’s urban core. (The darker colors indicate higher quality). (Above) Built environment delineation and habitat suitability index. ArcMap 10.7.1
  • 35. Results Habitat cost surface of the study area shows high resistance throughout the study area especially in the urban core (red) and in the suburban, exurban and rural areas (yellow). The Habitat Cost Surface shows that there is high resistance in the central core of the city (red), typical of all urban settlements. The built environment is very dense in that area and has high disturbance regimes (high cumulative human modification over time). The yellow is mostly suburban, exurban and agricultural uses and the green equates to most of the semi-natural habitat fragments. ArcMap 10.7.1 (Right) Habitat Cost Surface.
  • 36. Results The regional Least Cost Paths shows low to moderate permeability to the east and west for terrestrial movement to Delaware State Park to the north from the more permeable area to the south, however, both potential ecological corridors have many barriers, shown in red, where they pass through the periphery of the urban-exurban matrix. Groundtruthing (visual survey) reveals that these corridors are not feasible for ground movement especially for interior species. ArcMap 10.7.1 (Right) Habitat Cost Surface with LCP Paths.
  • 37. Results The results also revealed a higher permeability to the south mostly due to its rural nature and large number of protected areas. The corridors delineated by the LCP analysis have high conduciveness for movement in between Camp Lazarus/Seymour Woods and Deer Haven/Havener parks but less so with the core to the north (Stratford Ecological Center) because the riparian zone is not as permeable due to roads and development (the red line in the middle of the figure). ArcMap 10.7.1 (Above) Local LCP paths shown in black between the three small intact habitat cores in the southern portion of the study area.
  • 38. Results Moderate permeability was found to exist within the agricultural zones to the west, east and north but those are mostly open lands bisected by several road barriers with isolated woodlots and disjointed hedgerows offering low connectivity in its current state other than along the riparian zones of some less urbanized north-south rivers and streams. ArcMap 10.7.1 (Above) LCP paths shown in black through the isolated small intact habitat core (Gallant Woods) and the northern portion of the study area leading to the closest larger intact habitat core (Delaware State Park). the park and its adjacent wildlife area is 6,356 acres and is classified as a medium habitat core using methodology from the Green Infrastructure Center, Inc.
  • 40. Recommendations The best solutions revolve around how green spaces of all types and sizes can be best configured to increase connectivity in a complex exurban environment. Landscapes should be integrated at multiple scales by connecting semi- natural areas and developed open spaces (lawn-based parks, the city cemetery, road verges, road right of ways, utility right of ways, vacant lots, infill areas, paved greenway buffer zones, tree lawns/street trees, larger residential yards) for multifunctional connectivity between large intact habitat cores to increase diversity (especially among more mobile species like birds and pollinators), making a landscape less sensitive to human induced or climatic disturbances by building its capacity for resilience.
  • 41. Ways to increase connectivity Conservation and restoration of the city’s riparian ravines - The city’s urban and rural ravines (both protected and unprotected) provide the best opportunity for connectivity corridors between the identified terminal intact habitat cores or source patches in the study area if restored. The city of Toronto, Canada which has similar characteristics as the study area has created a ravine strategy that Delaware could look to for the management of its ravines (City of Toronto 2020).
  • 42. Ways to increase connectivity Utilizing vacant lots as stepping stone habitat - The creation of habitat in open spaces and vacant lots for more mobile species - mainly birds, bats and pollinating insects can be important reservoirs for species richness or diversity (Burghardt et al 2008; Villasenora et al 2020). For example, semi- natural vacant lots were found to harbor as many as 60 songbird species in a recent study in Baltimore (Brodsky 2016).
  • 43. Ways to increase connectivity The Northeast Ohio Ecological Consortium (NEOECO) has developed a Vacant Land Rapid Assessment Procedure (VL-RAP) that provides an efficient way to screen and evaluate sites using basic ecological and ecosystem principles to determine their potential suitability as wildlife habitat, stormwater management, parks, and gardens (Rouse and Bunster-Ossa 2013; Cleveland Urban Design Collaborative 2008).
  • 44. Ways to increase connectivity Utilizing road verges, rights of way and other open spaces to form stepping stone habitat - Planting pollinator pathways, micro prairies, bioswales, rain gardens and other green infrastructure can increase plant and animal biodiversity exponentially depending on which solution is utilized. Solutions and incentives for cities to improve connectivity through the adequate use of green infrastructure to address gaps are many. Solutions range from food forests and micro prairies to wetland parks.
  • 45. Side swale, Columbus (photos by Kevin Parks, ThisWeekNews) Examples of Green Infrastructure
  • 46. (Left) Bioretention planter, Dublin, Ohio. (Right) Bioretention cell near busy four-lane road near the Scioto River, Upper Arlington, Ohio. (photos by the author) Examples of Green Infrastructure
  • 47. (Left) Pollinator pathway interspersed in between a large solar array, Newark, Ohio. (Above) A floodable or sponge park, Powell, Ohio. (Solar photo by the author; Floodable park by Jenna Odegard, MAD Scientist Associates) Examples of Green Infrastructure
  • 48. Bioretention pond converted from a traditional stormwater retention pond to a wetland bioretention pond, Granville Land Lab, Granville, Ohio. (photos by the author) Examples of Green Infrastructure
  • 49. Land sharing and land sparing on agricultural lands More than 100 acres prairie, grass savanna, wetlands, an oak-hickory forest edge and a small grassy riparian zone on former intensively farmed cropland at the Granville Land Lab, Granville, Ohio. (photos by the author)
  • 50. Conclusion Cities like Delaware can play a fundamental role in the conservation of biodiversity through strategies that include the introduction of ecosystems and habitats into the urban fabric, or the preservation of pre-existing ones, as well as the creation of continuous urban green spaces that guarantee biological connectivity and control continuing fragmentation indicative of exurban settlements. It is hoped that the findings generated by this baseline structural connectivity and suitability analysis, can be used as a jumping off point for further comprehensive conservation efforts especially in light of continued population growth, rapid conversion of cropland to development, aging infrastructure concerns and continued habitat loss through fragmentation among other factors.
  • 51. Further Research Further research could be in the form of: ● Some type of functional connectivity assessment or assessments which is the ability of an area to meet the food and cover requirements for a specific animal species (Juntti and Rumble, 2006). ● A City Biodiversity Index (CBI). The CBI, pioneered by the city of Singapore (Deslauriers et al 2018), could follow functional connectivity assessments to further identify and prioritize the city of Delaware’s greenspace for stepping-stone habitat in addition to the prioritized corridors identified in this study. ● And bioblitzes and other similar citizen science initiatives to document the diversity and distribution of terrestrial and avian species especially during migration. These follow-on studies can be performed or hosted by Ohio Wesleyan University students in the Environmental Science and Geography programs as each student needs to do a capstone in the local area as part of their graduation requirements.
  • 52. References Text References ● Rosenberg K, Dokter A , Blancher P , Sauer J , Smith A , Smith P , Stanton J , Panjabi A , Helft L , Parr M and Marra P., Decline of the North American Avifauna, Science, 366:6461, pp 120 (2019). ● City of Toronto, Ravine Strategy Implementation Report, 66 pgs (January 2020). ● Burghardt K, Tallamy D and Shriver G., Impact of Native Plants on Bird and Butterfly Biodiversity in Suburban Landscapes, Conservation Biology, Volume 23, No. 1, 219–224 (2008). ● Villaseñora N, Chianga L, Hernándeza HJ and Escobar M., Vacant lands as refuges for native birds: An opportunity for biodiversity conservation in cities, Urban Forestry & Urban Greening, 49 (2020). ● Brodskey C., Urban Vacant Lots: Ecological Quality and Social Opportunities in Baltimore, Maryland, A Dissertation Presented to The Faculty of the Graduate School at the University of Missouri-Columbia (2016). ● Rouse and Bunster Ossa, Green Infrastructure: A Landscape Approach, 144 pgs (2013). Print. ● Cleveland Urban Design Collaborative, Resilient Cleveland, online report, (2008). https://www.cudc.kent.edu/resilient-cleveland ● Firehock, K, and Walker RA., Strategic Green Infrastructure Planning: A Multi-Scale Approach (2015) Print. ● Maps throughout this thesis were created using ArcGIS® software by Esri. ArcGIS® and ArcMap™ are the intellectual property of Esri and are used herein under license. Copyright © Esri. All rights reserved. For more information about Esri® software, please visit www.esri.com.
  • 53. References Text References ● Anderson MG., Clark M and Sheldon A., Estimating Climate Resilience for Conservation across Geophysical Settings, Conservation Biology, 28:4, pp 959–970 (2013). ● Taylor, P.D., Fahrig, L. Henein, K. and Merriam,G., Connectivity is a vital element of landscape structure, Oikos 68:3, pp 571-572 (1993). ● Soverel NO, Coops NC, White JC and Wulder MA, Characterizing the Forest Fragmentation of Canada’s National Parks, Environmental Monitoring and Assessment, 164:14, pp 481–499 (2010). ● Theobald DM, Landscape Patterns of Exurban Growth in the USA from 1980 to 2020, Ecology and Society, 10:1, p 32 (2005). ● Lindenmayer D and Hobbs R., eds., Managing and Designing Landscapes for Conservation: Moving from Perspectives to Principles, Blackwell Publishing, 608 pgs (2007). Print. ● Hansen A, Knight R, Marzluff J, Powell S, Brown K, Gude P and Kingsford J, Effects of exurban development on biodiversity: Patterns, mechanisms and research themes, Ecological Applications, 15:6, pp 1893–1905 (2005). ● Taylor P, Fahrig L and With K., Landscape Connectivity: A Return to the Basics, Cambridge University Press, pp 29-43 (2006). ● Tischendorf, L. and Fahrig, L. How should we measure landscape connectivity? Landscape Ecology 15, 633–641 (2000). ● Malczewski J., GIS-based multicriteria decision analysis: a survey of the literature, International Journal of Geographical Information Science, 20:7, pp 703-726 (2006).
  • 54. References Text References ● Pullinger, M.G., Johnson, C.J., Maintaining or restoring connectivity of modified landscapes: evaluating the least-cost path model with multiple sources of ecological information, Landscape Ecology, 25, 1547–1560 (2010). ● Fischhoff B and Davis A., Communicating scientific uncertainty, PNAS, p 111 (2014). ● Urban D and Keitt T., Landscape Connectivity: A Graph-Theoretic Perspective, Ecology 82:5, pp 1205-1218 (2001). ● Egan J and Mortensen D., A comparison of land-sharing and land-sparing strategies for plant richness conserva-tion in agricultural landscapes, Ecological Applications, 22:2, pp 459-471 (2012). ● Balmford B, Green R, Onial M, Phalan B., and Balmford A., How imperfect can land sparing be before land shar-ing is more favorable for wild species?, Journal of Applied Ecology, 56:1, pp 73- 84 (2019). ● Juntti TM and Rumble M., Arc Habitat Suitability Index computer software, Gen. Tech. Rep. RMRS- GTR-180WWW. Ft. Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 31 pages (2006). ● Kaitsa, George, Delaware County Auditor, County agricultural land use statistics. Retrieved 20 August 2020. ● City of Delaware, Comprehensive Plan 2003 – 2008, Delaware Ohio, Department of Planning and Community Development (2002). ● Ohio Revised Code, Title [15] XV State Scenic River Designation, Chapter 1547.81, online resource (accessed 17 March 2020) http://codes.ohio.gov/orc/1547.81v1.
  • 55. References Graphics and maps ● Eastern Forest and Migratory Bird Decline Graphics, Rosenberg K, Dokter A , Blancher P , Sauer J , Smith A , Smith P , Stanton J , Panjabi A , Helft L , Parr M and Marra P., Decline of the North American Avifauna, Science, 366:6461, pp 120 (2019). ● Side Swale, Columbus, Parks KP, Clintonville Rain Gardens Net Mixed Results, ThisWeek Community Newspapers (July 2015). ● Floodable park, Powell, Odegard J, MAD Scientists Associates (wetland scientists), Westerville, Ohio (2020). ● Delaware County Highway Map, Chris Bauserman, Delaware County Engineer (2015). ● Draft City of Delaware Comprehensive Plan Land Use Map (2021). ● Physiographic Regions of Ohio and Glacial Map of Ohio, ODNR (1998 and 2005) respectively. ● Natural Vegetation of Ohio maps, Sears PB, The natural vegetation of Ohio, III. Plant succession, Ohio Journal of Science, 26, pp 213-31 (1926b). ● State of Ohio Historical Maps and City of Delaware Plat Maps, Everts LH, A Illustrated Historical Atlas of Delaware County, Ohio, LH Everts and Co., Philadelphia (1866). ● Satellite shots taken from Google Earth Pro (2020). ● Original patch dynamic art by Shannon Hauck based on Firehock and Walker 2015). ● Figures created by author out of ArcMap 10.7.1 ● Original photos taken by author.