The Nymphal Odonate Fauna of Two Watersheds in the
Lower Potomac River Basin, Maryland, with Emphasis on
Rare Taxa
Patrick H. Graves III and Patrick J. Ciccotto
Northeastern Naturalist, Volume 18, Issue 4 (2011): 442–456
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2011 NORTHEASTERN NATURALIST 18(4):445–456
The Nymphal Odonate Fauna of Two Watersheds in the
Lower Potomac River Basin, Maryland, with Emphasis on
Rare Taxa
Patrick H. Graves III1,* and Patrick J. Ciccotto2
Abstract - Sixty percent of Maryland’s odonate species are considered to be in need of
conservation. To prioritize areas for the protection of biodiversity, the Maryland Department
of Natural Resources (MD DNR) has identified 10 watersheds with the highest rates
of occurrence of imperiled and rare stream species, including odonates. We examined
the lotic-breeding odonate fauna of two of these high priority watersheds to determine
the distribution and status of several imperiled odonate species in Maryland. Odonate
nymphs from two Lower Potomac River basin watersheds, Zekiah Swamp Run and
Breton Bay, were collected by volunteers and MD DNR’s Maryland Biological Stream
Survey from 2000–2010 and were identified to species level when possible. Thirty-four
species were collected during this survey, 10 of which are state-listed species. The data
collected in this survey detail the distributions, habitats, and microhabitats of rare odonates
in two priority watersheds in Maryland that can be used to aid in the conservation
of these species and their habitats.
Introduction
Dragonflies and damselflies (Insecta: Odonata) comprise a widespread faunal
group, with approximately 499 species in the United States (NatureServe 2010).
Approximately 18% of the odonates in the US are globally ranked as “at risk”
by The Nature Conservancy and NatureServe (Master et al. 2000, Natureserve
2010). While a number of state checklists and distributional records are available
for the US (e.g., Cruden and Gode 2000, Evans 1988, Johnson and Valley 2005,
Tennessen et al. 1995), few lists are published for purposes of conservation planning
in areas where distributional data, like natal habitat use, are critical for the
protection of biodiversity (Bried 2005, Bried and Ervin 2005, Hunt et al. 2010,
Muise et al. 2007, Orr 2001).
The Atlantic Coastal Plain physiographic province of the southeastern United
States, within which lies over 60% of Maryland, is a center of odonate diversity
and endemism (Kalkman et al. 2008). Well over half (109 species) of the approximately
180 dragonfly and damselfly species that have been documented in
Maryland are considered rare, threatened, or endangered (RTE) in the state (Maryland
Department of Natural Resources 2010). Pollution, sediment disturbance, and
loss and degradation of critical habitats are among the threats to odonates and other
aquatic fauna in Maryland (Kayzak et al. 2005; Southerland et al. 2005a, b). While
the knowledge of adult odonate species distribution is abundant for Maryland
1Monitoring and Non-tidal Assessment Division, Maryland Department of Natural Resources,
580 Taylor Avenue, Tawes Building C-2, Annapolis, MD 21401. 2Department
of Biological Sciences, University of Maryland Baltimore County, 1000 Hilltop Circle,
Baltimore, MD 21250. *Corresponding author - pgraves@dnr.state.md.us.
446 Northeastern Naturalist Vol. 18, No. 4
(Czaplak 1995; Donnelly 1961; Fisher 1940, Moorefield 1941; Orr 1995, 1996a,
1996b, 1996c, 1998a, 1998b, 2000a, 2000b; Root 1923; Solem and Solem 1999),
less is known about the distribution and habitats of odonate nymphs. Detailed distributional
and ecological data for the nymphal stage are needed to better assess the
conservation status of, and potential stressors to, Maryland’s odonate fauna.
The Maryland Biological Stream Survey (MBSS) is a statewide assessment of
the biodiversity of Maryland’s freshwater streams undertaken by the Maryland
Department of Natural Resources (MD DNR). The MBSS has documented the
distributions of, and threats to, a number of aquatic species, notably fishes and other
vertebrates (Kazyak et al. 2005, Kilian et al. 2009, Kline and Morgan 2000, Stranko
et al. 2008). These data have led to the designation of high priority watersheds, i.e.,
areas with high levels of aquatic biodiversity and high numbers of occurrences of
state rare aquatic species. Recent efforts by the MBSS have begun to examine the
distributions of invertebrate species, specifically freshwater mussels and crayfishes
(Ashton 2009, Kilian et al. 2010), to assess their role in the designation of these
conservation watersheds. Similar endeavors are needed for species of other invertebrate
groups, of which little is known about their distributions.
We report the results of a comprehensive survey of nymphal odonates in two
high priority watersheds in the Lower Potomac River basin, Zekiah Swamp Run
and Breton Bay. These two watersheds, along with eight others throughout the
state, have been designated by MD DNR as areas that provide the best opportunities
to maintain populations of aquatic RTE species. The Zekiah Swamp Run and
Breton Bay watersheds are located in an area where large human population growth
is projected over the next 30 years (Maryland Department of Planning 2008). The
first step in rare species conservation is to identify those places where the species
occurs. We examined the nymphal odonates of these two high priority watersheds
to determine the distribution, habitats, and microhabitats of several rare and imperiled
odonate species in the region. We anticipate this information to be useful in the
conservation of Maryland’s odonate diversity and its odonate habitat and in determining
the conservation status of lotic-breeding odonate species in Maryland.
Methods
Study sites
The target watersheds for this study, Zekiah Swamp Run and Breton Bay, are
located southeast of Washington, DC in the Lower Potomac River basin on the
Atlantic Coastal Plain physiographic province (Figs. 1 and 2). Zekiah Swamp Run
and Breton Bay drain approximately 28,300 and 15,600 ha, respectively. In both
watersheds, soils transition from moderately to well-drained silt or loam in the
north to predominately clay soils in the south. Pine-hardwood forests cover over
half of the area in both of these watersheds, with agricultural, urban, wetland, and
minimal proportions of other land-cover types comprising the remaining portions
of the landscape (Boward et al. 1998). The topography is characterized by flat
upland surfaces comprised of alluvial plains and fluvial-estuarine terraces with
steep-sided valleys created by stream incision (Reger and Cleaves 2008). Stream
habitats in Zekiah Swamp Run and Breton Bay range in pH from slightly acidic
2011 P.H. Graves and P.J. Ciccotto 447
to neutral and are influenced strongly by seasonal precipitation, with the larger
tributaries expanding onto the floodplain after significant wet periods and drying
during the summer months. Substrates vary from predominately silt, sand, and
gravel in the smaller tributaries of the watersheds to sand and cobble in the larger
tributaries. Several imperiled aquatic animal species are native to these two watersheds,
including six fish species and two freshwater mussel species, having led
to their designation as conservation priority watersheds.
Figure 1. Rare, threatened, and endangered (RTE) odonate species richness at sites in the
Zekiah Swamp Run watershed, MD.
448 Northeastern Naturalist Vol. 18, No. 4
Odonate data
During March and April 2000–2010, benthic macroinvertebrates were
collected from 327 stream sites using MBSS sampling protocols (Stranko et
al. 2007). Thirty of these sites were sampled by the MBSS staff, and 297 were
sampled by Stream Waders volunteers, a group organized to supplement MBSS
data through volunteer sample collections. Criteria used for selection of MBSS
and Stream Waders sites are outlined in Klauda et al. (2008) and Boward and
Figure 2. Rare, threatened, and endangered (RTE) odonate species richness at sites in the
Breton Bay watershed, MD.
2011 P.H. Graves and P.J. Ciccotto 449
Bruckler (2008). In brief, this sampling protocol consisted of sampling 1.86 m2
(20 ft2) of site-specific characteristic habitats (most productive macroinvertebrate
habitats, as defined in Stranko et al. [2007]) using a 540-micron mesh D-net.
Organisms were dislodged from substrates and organic debris, such as wood
and leaf packs, by disturbing these habitats upstream of the D-net with kicking
and rubbing by hand. Contents of the D-net samples were washed into a sieve
bucket, and large debris was removed, inspected for organisms, and discarded.
The sample was preserved in at least 70% ethyl alcohol.
In the lab, odonate nymphs were sorted from the samples, and late-instar individuals
were identified by DNR taxonomists to species level. Individuals in an
early instar stage or missing a body part were identified to either family or genus
level. Individuals identified to species level were compared to DNR voucher
specimens when that species was present in the voucher collection. Twenty individuals
(eight taxa) were sent to a Maryland odonate expert for verification. The
primary sources used for identification of specimens were Needham et al. (2000)
and Westfall and May (1996). Other sources of information used include Cashatt
and Vogt (2001), Kennedy (1924), and Tennessen (2008). Specimens collected in
this study are housed at MD DNR in Annapolis, MD.
Stream-reach habitat information was recorded for MBSS sites in these watersheds
from 2000–2010. Microhabitat information (e.g., dominant substrate
[sand, cobble, silt, etc.], habitat components [depth, velocity, rootwad, woody
debris, etc.], channel unit [pool, riffle, etc.]) was recorded, when possible, by the
authors at MBSS Breton Bay watershed sites during 2009 and 2010.
In 2009, 15 sites in the Breton Bay watershed were sampled similarly to the
protocol outlined in Stranko et al. (2007), with the exception that the more productive
odonate habitat was sampled instead. This change was made in an effort
to more accurately capture the odonate fauna at these sites by sampling habitats
utilized by odonates (e.g., sand and silt, undercut banks, rootmats), which would
typically be ignored otherwise.
Results and Discussion
During 2001 and 2008, 341 odonate nymphs were collected from 137 sites in
the Zekiah Swamp Run watershed. This collection is represented by 24 species,
19 genera, and nine families (Table 1). From 2000–2010, 1044 odonate nymphs
were collected from 190 sites in the Breton Bay watershed. The Breton Bay collection
is represented by 25 species, 21 genera, and eight families (Table 1). The
most-collected dragonfly family (Gomphidae), dragonfly species (Boyeria vinosa),
damselfly family (Calopterygidae), damselfly genus (Calopteryx), and damselfly
species (Calopteryx maculata) were similar between the two watersheds. Cordulegaster
and Boyeria were the most collected dragonfly genera in the Zekiah
Swamp Run and Breton Bay watersheds, respectively. Eleven species were present
in both collections. The total number of individuals of each species (as well as the
number of individuals identified to the genus or family level) and the percentage
of sites each species was collected at within each watershed is shown in Table 1.
450 Northeastern Naturalist Vol. 18, No. 4
Table 1. Number of nymphal odonate individuals collected from streams in the Zekiah Swamp Run
and Breton Bay watersheds. The percentage of sites at which a species was collected within the respective
watershed is given in parentheses. An asterisk indicates a species listed as rare, threatened,
or endangered in Maryland (Maryland Department of Natural Resources 2010). Individuals listed
in this table at the family and genus level were not identifiable to species level due to immaturity
or missing body parts.
No. of individuals collected
(% of sites collected at)
Zekiah
Scientific name Common name Swamp Run Breton Bay
Anisoptera (dragonflies)
Aeshnidae 1 -
Aeshna umbrosa Walker Shadow Darner 3 (0.7) -
Basiaeschna janata (Say) Springtime Darner 1 (0.7) 7 (2.1)
Boyeria vinosa (Say) Fawn Darner 30 (16.1) 161 (36.8)
Epiaeschna heros (Fabricius) Swamp Darner 2 (1.5) -
Nasiaeschna pentacantha (Rambur)* Cyrano Darner - 1 (0.5)
Cordulegastridae - -
Cordulegaster bilineata (Carle)* Brown Spiketail 3 (2.2) -
Cordulegaster erronea Hagen in Selys* Tiger Spiketail 17 (6.6) -
Cordulegaster maculata Selys Twin-spotted Spiketail 10 (5.8) 8 (8.4)
Cordulegaster obliqua (Say)* Arrowhead Spiketail 4 (2.2) 3 (1.6)
Cordulegaster sp. A spiketail 45 131
Corduliidae 7 21
Epitheca cynosura (Say) Common Baskettail - 2 (0.5)
Epitheca princeps Hagen Prince Baskettail - 1 (0.5)
Epitheca sp. A baskettail 1 4
Helocordulia selysii (Hagen in Selys)* Selys' Sundragon 4 (2.2) 27 (7.9)
Helocordulia sp. A sundragon 3 6
Neurocordulia sp. A shadowdragon 1 -
Somatochlora linearis (Hagen)* Mocha Emerald 6 (2.9) -
Somatochlora tenebrosa (Say) Clamp-tipped Emerald 1 (0.7) 13 (1.6)
Somatochlora sp. An emerald 3 33
Gomphidae 67 90
Gomphus exilis Selys Lancet Clubtail - 2 (0.5)
Gomphus lividus Selys Ashy Clubtail 1 (0.7) 13 (3.2)
Gomphus rogersi Gloyd* Sable Clubtail 11 (5.8) -
Hagenius brevistylus Selys Dragonhunter 3 (1.5) 9 (4.7)
Progomphus obscurus (Rambur)* Common Sanddragon - 4 (2.1)
Stylogomphus albistylus (Hagen in Selys) Eastern Least Clubtail 16 (8) 29 (11.1)
Stylurus sp. A clubtail - 1 (0.5)
Libellulidae 4 -
Libellula cyanea Fabricius Spangled Skimmer 1 (0.7) -
Libellula vibrans Fabricius Great White Skimmer - 1 (0.5)
Libellula sp. A skimmer 2 4
Pachydiplax longipennis (Burmeister) Blue Dasher - 7 (2.1)
Plathemis lydia (Drury) Common Whitetail - 2 (1.1)
Macromiidae - 6
Didymops transversa (Say) Stream Cruiser - 1 (0.5)
Macromia illinoiensis Walsh Swift River Cruiser - 4 (1.6)
Macromia sp. A river cruiser 1 12
2011 P.H. Graves and P.J. Ciccotto 451
Overall, 38% of the individuals collected were identified to species level, and 47%
and 17% were identified to genus and family level, respectively. Of the dragonflies
collected, 48% were identified to species level. Due to immaturity or the loss of
caudal gills, only 21% of damselflies were identified to species level.
Six RTE dragonfly species were collected in Zekiah Swamp Run and four RTE
dragonfly and two RTE damselfly species were found in Breton Bay (Table 1).
Two of these species, Helocordulia selysii and Cordulegaster obliqua, were collected
in both watersheds. The most collected RTE species was Cordulegaster
erronea (17 individuals) in Zekiah Swamp Run and H. selysii (27 individuals)
in Breton Bay. The least collected RTE species in each watershed was Cordulegaster
bilineata in Zekiah Swamp Run and Nasiaeschna pentacantha in Breton
Bay (one individual each). At least one RTE species was collected at 17% of the
Zekiah Swamp Run sites and 14% of the Breton Bay sites.
In general, RTE odonate species in Breton Bay and Zekiah Swamp Run were
distributed in lotic habitats throughout the watersheds (Figs. 1, 2). Multiple RTE
species were collected at sites in both watersheds. In the Zekiah Swamp Run
watershed, 46% of the sites at which multiple RTE species were collected lie in
the southern region of the watershed, including the most RTE-species-rich site
(3 species) in either watershed. All four multiple RTE species sites in the Breton
Bay watershed lie in the eastern half of the watershed.
All but one of the 17 C. erronea individuals were collected at sites in the southern
region of the Zekiah Swamp Run watershed. These eight sites were small,
shaded tributaries to Zekiah Swamp Run with average depths ranging from 5–30
cm. Combined, seven other odonate species were collected along with C. erronea:
H. selysii, G. rogersi, Stylogomphus albistylus, Calopteryx maculata, B. vinosa,
Cordulegaster maculata, and Cordulegaster obliqua. Two RTE species, H. selysii
and G. rogersi, were collected along with C. erronea at one site in the southwestern
part of this watershed. In addition, C. erronea was found at three of the eight sites
Table 1, continued.
No. of individuals collected
(% of sites collected at)
Zekiah
Scientific name Common name Swamp Run Breton Bay
Zygoptera (damselflies) - -
Calopterygidae - -
Calopteryx maculata (Beauvois) Ebony Jewelwing 17 (11) 63 (18.4)
Calopteryx sp. A jewelwing 44 248
Coenagrionidae 11 29
Argia fumipennis violacea (Hagen) Violet Dancer 1 (0.7) 26 (5.8)
Argia sp. A dancer 3 43
Enallagma divagans Selys* Turquoise Bluet - 1 (0.5)
Enallagma durum (Hagen)* Big Bluet - 5 (1.6)
Enallagma sp. A bluet 1 11
Ischnura posita posita (Hagen) Fragile Forktail - 2 (1.1)
Ischnura sp. A forktail 14 13
Lestidae - -
Lestes sp. A spreadwing 2
452 Northeastern Naturalist Vol. 18, No. 4
at which G. rogersi was collected in the Zekiah Swamp Run watershed. Cordulegaster
erronea was not collected in the Breton Bay watershed. Other than average
wetted width and depth, detailed habitat information was not recorded at these
sites, as they were all sampled by volunteers. Further investigation of these sites
would reveal if the habitat requirements of C. erronea in this watershed are similar
to those reported in New Jersey (Barlow 1995) and Ohio (Glotzhober 2006):
shaded, small, permanent cool- to cold-water seeps/springs with sandy substrate
and relatively free of fish, other odonate species, and organic matter.
During the 2009–2010 Breton Bay watershed sampling, the microhabitats in
which RTE species were collected were noted when possible. While the sample
size of some of these species is small, we feel the description of the microhabitat
in which these species were found adds to what little is known about the nymphs
of these species and their microhabitat preferences in these watersheds as well as
others throughout Maryland. Cordulegaster obliqua individuals were collected in
both watersheds from small (less than 3 m average wetted width), shallow streams with
sand-dominated substrate. The majority of these streams were temporary, and we
found them dry upon a revisit in the summer. All Progomphus obscurus nymphs, a
burrowing gomphid species, were collected in stable sand substrate with little to no
silt in very shallow waters near the shoreline in three small streams and one large
(6 m average wetted width) stream. Pool/glide habitat dominated the reaches in
which P. obscurus were found. Twenty-four individuals of H. selysii, a species at
or near the northeasternmost extent of its global range in Maryland, were collected
in small, permanent streams with average depths of 20–30 cm, while two were collected
in streams with average depths of 80 cm. Helocordulia selysii collected in
2008–2009 were primarily taken from sand and silt substrates beneath undercut
banks. Nearly all individuals of H. selysii were collected at sites located 3 km or
less from Breton Bay. The majority of individuals of the six species reported from
Breton Bay that typically inhabit lentic systems or slow-moving areas of streams
and rivers were collected from stream sites with dammed pool habitat and reduced
flow resulting from Castor canadensis Kuhl (Beaver) activity or natural blockages
(i.e., fallen tree). The RTE species Nasiaeschna pentacantha and Enallagma durum
were among those collected at these sites.
While the RTE status of Somatochlora tenebrosa was removed with the 2010
state rarity rank update (MD DNR 2010), the microhabitat of this species was observed
and recorded at two different streams in the Breton Bay watershed in 2009
and 2010 and is included here. One of the streams is located in the southeastern
part of the watershed, while the other lies in the northeastern part. In both streams,
S. tenebrosa were found in a cluster of 4–5 late-instar individuals in a small area
beneath undercut banks in stable sand substrate with a thin layer of silt on top.
These individuals accounted for all but two of the S. tenebrosa collected during a
3–4 hour concentrated Somatochlora search at return visits to both stream sites.
The individuals collected during this concentrated search were reared in a lab and
identified as adults to species level.
Stream site characteristics and the microhabitat in which the clusters of S. tenebrosa
were found were similar among the two sites. The streams had a wetted
2011 P.H. Graves and P.J. Ciccotto 453
width of approximately 1.5 m and were shallow (5–10 cm). The individuals were
underneath undercut banks cut less than 20 cm into the bank at bends in the stream.
Immediately upstream of the undercut banks, woody debris created small pool
habitat with reduced flow velocity in the segment containing the undercut bank.
The two S. tenebrosa individuals that were not part of a cluster were collected
separately in run habitat, against a bank containing numerous burrows of Cambarus
diogenes Girard (Devil Crayfish). No observation was made of S. tenebrosa
utilizing the burrows as refugia, but given the large number of burrows in the
stream banks and seepage areas, and the observation by Soluk et al. (1996) of
Somatochlora hineana utilizing C. diogenes burrows, it is worthwhile to investigate
whether Somatochlora species, or any other odonate taxa, are using crayfish
burrows in Breton Bay watershed streams and elsewhere.
A sampling design that targets the most productive odonate habitats and not
the most productive overall benthic macroinvertebrate habitats is encouraged for
future nymphal odonate surveys. Even though only 8% of the sites in the Breton
Bay watershed from 2002–2009 were sampled using the most productive odonate
habitat sampling method, 32% of odonates analyzed in this study were collected
from this watershed using this method. This includes 71% of libellulids, 44% of
gomphids, 50% of coenagrionids, 29% of corduliids, and 52% and 53% of the
RTE-rich genera Cordulegaster and Somatochlora, respectively. By focusing on
odonate-preferred habitats, relatively odonate-poor habitats like riffles would not
be sampled as rigorously as they would when using MBSS (Stranko et al. 2007) or
other benthic macroinvertebrate protocols. This focus on odonate-preferred habitats
allows for more sampling of habitats such as sand, silt, and undercut banks, for
example, that otherwise would not receive the amount of attention they deserve.
Conclusions
The results of these studies highlight the RTE nymphal odonate fauna in two
high priority watersheds of the Potomac River drainage in Maryland. These
studies provide much needed data on the distribution and microhabitats of several
RTE species and reveal odonate richness hotspots in these two watersheds.
These data can also indicate sites for further quantitative sampling, which can be
used to establish a baseline for tracking odonate species and community changes
over time as well as further identify the core range of an RTE species in these
watersheds or a subwatershed therein. This information will help facilitate future
studies to better determine microhabitat requirements and environmental stressors
that may impact these species. Such studies may be particularly important
and timely as Charles County, which contains the majority of the Zekiah Swamp
Run watershed, is one of the fastest growing counties in Maryland (Maryland
Department of Planning 2008). Documenting changes in the distribution and status
of odonates and other rare fauna and flora in this area may provide important
information for planning future development projects in Maryland and in the
conservation of Maryland odonate habitat and diversity.
Our results also contributed to a recently completed status reassessment
of Maryland’s odonate fauna (MD DNR 2010). The resulting status changes
454 Northeastern Naturalist Vol. 18, No. 4
included the downlisting of Gomphus rogersi from state endangered to in need of
conservation and the listing of H. selysii as state threatened. Breton Bay odonate
data, and that from watersheds yet to be sampled, can be used in future status
updates of Maryland odonate species as well as aid in the conservation, management,
and protection of the streams they inhabit.
Acknowledgments
This study was funded in part by State Wildlife Grant funds provided to the State
Wildlife agencies by US Congress, and administered through the Maryland Department
of Natural Resources’ Natural Heritage Program. We thank Stream Waders volunteers for
collecting the great majority of data presented in this report, the MBSS for the collection
and storage of odonates and data, and Richard Orr, Neal Dziepak, and Ellen Friedman for
the identification and verification of certain specimens. This manuscript benefited from
comments by Jason Bried and two anonymous reviewers.
Literature Cited
Ashton, M.J. 2009. Freshwater mussel records collected by the Maryland Department
of Natural Resources’ Monitoring and Non-tidal Assessment Division (1995–2009):
Investigating environmental conditions and potential host fish of select species.
Report no. RAS-MANTA-AIM-10-01. Maryland Department of Natural Resources,
Annapolis, MD. 63 pp. Available online at http://www.dnr.state.md.us/streams/pdfs/
AIM_10_01_mussels.pdf. Accessed 16 December 2010.
Barlow, A.E. 1995. On the status of Cordulegaster erronea Hagen in Selys, 1878 in the
state of New Jersey. Argia 7:6–9.
Boward, D., and R. Bruckler. 2008. Maryland Stream Waders volunteer stream monitoring
manual (Revised edition January 2008). Report no. DNR-12-1242008-273. Monitoring
and non-tidal assessment division, Annapolis, MD. 34 pp. Available online at
http://www.dnr. state.md.us/streams/pdfs/sw2008_man.pdf. Accessed 6 July 2010.
Boward, D., D. Randle, P. Kazyak, H. Dail, and J. Christmas. 1998. Lower Potomac
River Basin environmental assessment of stream conditions. Report no. CBWPMANTA-
EA-96-5. Maryland Department of Natural Resources, Annapolis, MD. 29
pp. Available online at http:/ /www.dnr.state.md.us/irc/docs/00001765.pdf. Accessed
28 July 2010.
Bried, J.T. 2005. Species of adult Odonata from three natural areas in Mississippi. Journal
of the Mississippi Academy of Sciences 50:231–232.
Bried, J.T., and G.N. Ervin. 2005. Distribution of adult Odonata among localized wetlands
in east-central Mississippi. Southeastern Naturalist 4:731–744.
Cashatt, E.D., and T.E. Vogt. 2001. Description of the larva of Somatochlora hineana
with a key to the larvae of the North American species of Somatochlora (Odonata:
Corduliidae). International Journal of Odonatology 4:93–105.
Cruden, R.W., and O.J. Gode. 2000. The Odonata of Iowa. Bulletin of American Odonatology
6:13–48.
Czaplak, D. 1995. Lanthus vernalis and parvulus: New species records for Maryland.
Argia 7:27–28.
Donnelly, T.W. 1961. The Odonata of Washington, DC, and vicinity. Proceedings of the
Entomological Society of Washington 63:1–13.
Evans, M.A. 1988. Checklist of the Odonata of Colorado. Great Basin Naturalist 48:96–101.
Fisher, E.G. 1940. List of Maryland Odonata. Entomological News 51:37–42, 67–72.
Glotzhober, R.C. 2006. Life history of Cordulegaster erronea Hagen (Odonata: Cordulegastridae)
in the laboratory and the field. Bulletin of American Odonatology 10:1–18.
2011 P.H. Graves and P.J. Ciccotto 455
Hunt, P.D., M. Blust, and F. Morrison. 2010. Lotic Odonata of the Connecticut River in
New Hampshire and Vermont. Northeastern Naturalist 17:175–188.
Johnson, J., and S. Valley. 2005. The Odonata of Oregon. Bulletin of American Odonatology
8:100–122.
Kalkman, V.J., V. Clausnitzer, K.B. Dijkstra, A.G. Orr, D.R. Paulson, and J. van Tol.
2008. Global diversity of dragonflies (Odonata) in freshwater. Hydrobiologia
595:351–363.
Kazyak, P.L., J.V. Kilian, S.A. Stranko, M.K. Hurd, D.M. Boward, C.J. Millard, and A.
Schenk. 2005. Report no. CBWP-MANTA-EA-05-6. Maryland Biological Stream
Survey 2004, Volume 9: Stream and riverine biodiversity. Maryland Department of
Natural Resources, Annapolis, MD. 108 pp.
Kennedy, C.H. 1924. Notes and descriptions of naiads belonging to the dragonfly genus
Helocordulia. Proceedings of the United States Museum 64:1–4.
Kilian, J.V., S.A. Stranko, R.L. Raesly, A.J. Becker, and P. Ciccotto. 2009. Enneacanthus
chaetodon (Blackbanded Sunfish): An imperiled element of Maryland’s Coastal Plain
ichthyofauna. Southeastern Naturalist 8:267–276.
Kilian, J.V., A.J. Becker, S.A. Stranko, M.A. Ashton, R.J. Klauda, J. Gerber, and M.
Hurd. 2010. The status and distribution of Maryland crayfishes. Southeastern Naturalist
9:11–32.
Kline, M.J., and R.P. Morgan II. 2000. Current distribution, abundance, and habitat preferences
of the Stonecat (Noturus flavus) in Maryland. Report no. CBWP-MANTAEA-
00-7. Maryland Department of Natural Resources, Annapolis, MD. Available
online at http://www. dnr.state.md.us/streams/pdfs/ea-00-7_ stonecat.pdf. Accessed
15 August 2010.
Klauda, R. P. Kazyak, S. Stranko, M. Southerland, N. Roth, and J. Chaillou. 1998.
Maryland Biological Stream Survey: A state agency program to assess the impact of
anthropogenic stresses on stream habitat quality and biota. Environmental Monitoring
and Assessment 51:299–316.
Maryland Department of Natural Resources (MD DNR). 2010. Rare, threatened, and endangered
animals of Maryland. Maryland Department of Natural Resources, Wildlife
and Heritage Service, Natural Heritage Program, Annapolis, MD. Available online at
http://www.dnr.state.md.us/ wildlife/Plants_Wildlife/rte/pdfs/rte_Animal_List.pdf.
Accessed 17 December 2010.
Maryland Department of Planning. 2008. Historical and projected total population for
Maryland’s jurisdictions. Maryland Department of Planning, Planning Data Services,
Baltimore, MD. Available online at http://www.epa.gov/ttn/naaqs/ozone/areas/pop/
popp_md .pdf. Accessed 4 April 2011.
Master, L.L., B.A. Stein, L.S. Kutner, and G.A. Hammerson. 2000. Vanishing assets:
Conservation status of US species. Pp. 93–108, In B.A. Stein, L.S. Kutner, and J.S.
Adams (Eds.). Precious Heritage: The Status of Biodiversity in the United States.
Oxford University Press, Oxford, UK.
Moorefield, H.H. 1941. Addenda to the Odonata of Maryland. Entomological News
52:133–134.
Muise, C., K.R. Langdon, R.P. Shiftlett, D. Trently, A. Hoff, P. Super, A. Mayor, and B.J.
Nichols. 2007. Checklist of Odonata from Great Smoky Mountains National Park.
Southeastern Naturalist 6 (Special Issue 1):207–214.
NatureServe. 2010. NatureServe Explorer: An online encyclopedia of life [web application].
Version 7.1. NatureServe, Arlington, VA. Available online at http://www.natureserve.
or g/explorer. Accessed 17 December 2010.
Needham, J.G., M.J. Westfall, Jr., and M.L. May (Eds.). 2000. Dragonflies of North
America (Revised Edition). Scientific Publishers, Gainesville, FL. 940 pp.
Orr, R.L. 1995. Great Blue Herons, Dwarf Wedge Mussels, and Gray Petaltails. Argia
7(3):19–21.
456 Northeastern Naturalist Vol. 18, No. 4
Orr, R.L. 1996a. The Odonata of Patuxent Wildlife Research Center and vicinity. Argia
4(2):37–67.
Orr, R.L. 1996b. The Odonata of the Chesapeake and Ohio Canal National Historical
Park. Argia 8(3):6–10.
Orr, R.L. 1996c. Maryland county records. Argia 8(4):18.
Orr, R.L. 1998a. The Odonata of Sideling Hill Creek. Argia 10(1):14–15.
Orr, R.L. 1998b. Gomphid emergence times along the Little Patuxent River, Maryland.
Argia 10(4):8.
Orr, R.L. 2000a. The dragonflies and damselflies of Finzel Swamp (Maryland). Argia
12(1):13–14.
Orr, R.L. 2000b. Preliminary list of the dragonflies and damselflies of Washington, DC.
Argia 13(1):20–22.
Orr, R.L. 2001. The dragonflies and damselflies (Insecta: Odonata) of Cove Point,
Calvert County, Maryland. The Maryland Naturalist 44:5–19.
Reger, J.P., and E.T. Cleaves. 2008. Draft physiographic map of Maryland and explanatory
text for the physiographic map of Maryland. Maryland Department of Natural
Resources, Maryland Geological Survey, Annapolis, MD. Available online at http://
www.mgs.md.gov/c oastal/maps/physio.html. Accessed 20 August 2010.
Root, F.M. 1923. Notes on Zygoptera (Odonata) from Maryland, with a description of
Enallagma pallidum, n. sp. Entomological News 34:200–204.
Solem, R.P., and J.K. Solem. 1999. First Maryland record of Orthemis ferruginea (Odonata:
Libellulidae). Argia 11:6–7.
Soluk, D.A., B.J. Swisher, and D.S. Zercher. 1996. The ecology of Hine’s Emerald Dragonfl
y (Somatochlora hineana): Monitoring populations and determining patterns of
habitat use in the Des Plaines River Valley. Activity summary and report of preliminary
results (January–August 1996). Illinois Natural History Survey, Champaign, IL. 35 pp.
Southerland, M.T., L.A. Erb, G.M. Rogers, and P.F. Kazyak. 2005a. Maryland Biological
Stream Survey 2000–2004, Volume 7: Statewide and basin conditions. Report no.
DNR-12-0305-0109. MD DNR, Annapolis, MD. 46 pp. Available online at http://www.
dnr.state.md.us/streams/pdfs/ea-05-4_statewide.pdf. Accessed 13 August 2010.
Southerland, M.T., L.A. Erb, G.M. Rogers, R. Morgan, K. Eheleman, M. Kline, K. Kline, S.
Stranko, P. Kazyak, J. Kilian, J. Ladell, and J. Thompson. 2005b. Maryland Biological
Stream Survey 2000–2004, Volume 14: Stressors affecting Maryland streams. Report
no. DNR-12-0305-0101. MD DNR, Annapolis, MD. 135 pp. Available online at http://
www.dnr.state.md.us/streams/pdfs/ea-05-11_stressors.pdf. Accessed 13 August 2010.
Stranko, S., D. Boward, J. Kilian, A. Becker, R. Gauza, A. Schenk, A. Roseberry-Lincoln,
M. Ashton, and P. Kayzak. 2007. Maryland Biological Stream Survey, Round Three
Field Sampling Manual (Revised edition January 2009). Report no. CBWP-MANTAEA-
07-01. Monitoring and Non-Tidal Assessment Division, Annapolis, MD. 73 pp.
Available online at http://www.dnr.state.md.us/streams/pdfs a-07-01b_fieldRev2010.
pdf. Accessed 6 July 2010.
Stranko, S.A., R.H. Hilderbrand, R.P. Morgan, M.W. Staley, A.J. Becker, A. Roseberry-
Lincoln, E.S. Perry, and P.T. Jacobson. 2008. Brook Trout declines with land-cover
and temperature changes in Maryland. North American Journal of Fisheries Management
28:1223–1232.
Tennessen, K.J. 2008. Odonata (Chapter 12). Pp. 237–294, In R.W. Merritt, K.W. Cummins,
and M.B. Berg (Eds.), An Introduction to the Aquatic Insects of North America.
Kendall/Hunt, Dubuque, IA.
Tennessen, K.J., J.D. Harper, and S.K. Krotzer. 1995. The distribution of Odonata in
Alabama. Bulletin of American Odonatology 3:49–74.
Westfall, M.J., and M.L. May. 1996. Damselflies of North America. Scientific Publishers,
Gainesville, FL. 649 pp.