2012
SOUTHEASTERN NATURALIST
ll(4):551-566
Nephila clavipes (Araneae: Nephilidae):
A Model Species for Monitoring Climate Change in the
_ Southeastern United States
"
Kristin A. Bakkegard'* and Lawrence J. Davenport'
Abstract - We propose that Nephila clavipes (Golden Silk Orbweaver) be used as a model species to track climate change in the southeastern United States. As an ectotherm, it is
sensitive to changes in local climatic conditions. More importantly, this large, unmistakable spider elicits a strong emotional reaction, especially from those unfamiliar with it.
As its range expands, people will take notice, photograph it, and then post those pictures
in the public domain via blogs and social or scientific websites that can be easily viewed
by scientists. For a starting point, we present the most complete range map to date for this
species. This map shows that A^. clavipes, traditionally restricted to the Gulf and Atlantic
coastal areas of the southeastern United States, has recently (since 2000) expanded its
range out of the Coastal Plain into counties north of the Fall Line.
Introduction
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The past and present distributions of species have often been used in studies
of climate change. Changes in pollen assemblages have facilitated our understanding of north-south plant migrations during the Pleistocene (Delcourt and
Delcourt 1981, Gates 1993), while distribution patterns have documented the
regional and local changes that we are currently experiencing (Malcolm and
Pitelka 2000, Parmesan and Galbraith 2004, Peters 1991). The documentation of
future changes will rely on observations made on key species over many years.
An ideal species for such monitoring should: 1) be sensitive to climate flux, 2) be
large enough to be easily observed, 3) be unmistakably unique in its identifying
characteristics, and 4) have a broadly known (and accessible) baseline distribution record.
We propose that Nephila clavipes L. (Golden Silk Orbweaver, Golden Silk
Spider, or Banana Spider) be used as a model species to track climate change
in the southeastern United States. Our hypothesis is that this species' range will
expand northward and westward out of the Coastal Plain physiographic region,
across the Fall Line and into other, upland physiographic regions (Davenport
2007). This spider is easily recognized and cannot be confused with any other
spider in the United States (Kaston 1978, Levi 1980). Females (Fig. 1 A) are large
(to 25 mm céphalothorax length); have conspicuous tufts of hair on the femora
and tibiae of legs I, II, and IV; make large golden webs (up to 1 m in diameter,
1-3 m above the ground); and are easy to identify from photographs (Howell and
Jenkins 2004). Males are considerably smaller and usually noticed only when in
the web of a female. The only other spider with which adult female N. clavipes
'Department of Biological and Environmental Sciences, Samford University, Birmingham, AL 35229. 'Corresponding author - kbakkega@samford.edu.
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Southeastern Naturalist
Vol. 11, No. 4
could be confused with by an untrained eye is Argiope aurantia Lucas (Blackand-Yellow Argiope or Garden Spider). However, A. aurantia has a rounder, less
angular body and no hairy tufts on its legs (Fig. 1B); in addition, it does not have
a golden-yellow web, and its web often contains a stabilimentum, which inspires
another common name, the Writing Spider (Howell and Jenkins 2004).
Spiders are ectotherms and thus are highly influenced by their microclimate
(especially temperature and humidity), local weather, and large-scale climatic
trends (Pulz 1987). Globally, the genus Nephila is restricted to tropical climes,
with A'^. clavipes as the only species found in the Americas, from Argentina to the
southeastern United States (Kuntner et al. 2008, Levi 1980). Its nothern range
limit in the continent is currently in the southern portion of the southeastern US
(Fig. 2). Therefore, its northward expansion will be limited by winter conditions,
since tropical spiders are more likely to be injured by cold weather. However,
spider eggs in general appear to be highly (to -24 °C) resistant to cold (Kirchner
1987). In winter, female N. clavipes behaviorally thermoregulate by sun-basking.
They can elevate their body temperature during the daytime 7 °C above ambient
air temperature and will change the orientation of their webs to maximize insolation (Carrel 1978). At temperatures below 10 °C, A^. clavipes becomes inactive.
Figure 1. Photographs of similar spiders found in the southeastern United States: A. Female Nephila clavipes. Note the rectangular abdomen and distinctive leg tufts. The web
is also distinctively yellow (not seen in photo). B. Female Argiope aurantia. Note the
rounded abdomen and no leg tufts. Webs usually have a stabilimentum (thicker, zig-zag
portion seen running down center of photo under spider). Both photographs, by W.M.
Howell, were taken in Geneva County, AL, and are used with permission.
^,,.,
2012
K.A. Bakkegard and L..I. Davenport
553
while temperatures below 0 °C are fatal to adults (Moore 1977). In the sumtner, if
exposed to direct sun, the spider thermoregulates by making postural adjustments
in its web (Higgins and Ezcurra 1996, Robinson and Robinson 1974). As temperatures increase, individuals will perform evaporative cooling behaviors at 36.78
°C; death occurs at 41.65 °C (Krakauer 1972). Being a tropical species, A^. clavipes
prefers a relative humidity greater than 80% (Moore 1977).
In temperate zones, adult Nephila clavipes have a life span of one year (Moore
1977). Females make 1 to 5 egg sacs during that year, with the average number
of eggs per clutch ranging from 338.3 ± 117.8 to 585.8 ± 249.0, depending on the
deposition date (Higgins 1992, Moore 1977). Females deposit clutches anywhere
from September through November (Moore 1977, Wilder 1865). This spider
overwinters as eggs and spiderlings, which stay in the egg case 5-7 months until
emerging sometime between mid-spring to June to form a communal web, molt,
and then disperse (Higgins 2000, Hill and Christenson 1981, Moore 1977). Females mature in 3-5 months (8-10 instars), whereas males mature earlier (3-5
instars) and with a significantly smaller body size (Higgins 2000, Higgins and
Goodnight 2010). Juvenile males make their own webs for prey capture until
their final molt. Then within 3 days, they abandon their webs to search for females (Myers and Christenson 1988). Males often co-habitat with females, and
A^. clavipes webs often contain kleptoparasites (Agnarsson 2003).
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1961 — 1999
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Figure 2. Range of Nephila clavipes in the continental United States (one 1965 Arizona
record and one 1935 California record not shown). Dates indicate the earliest record for
the spider. Solid line indicates the Fall Line which delineates the coastal plain from interior physiographic regions. Counties with no records are in white.
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Southeastern Naturalist
Vol. 11, No. 4
Presumably, like many other spiders, Nephila clavipes is capable of
ballooning (Decae 1987). In Simberloff and Wilson's (1969) classic study
of recolonization on mangrove islands, A^. clavipes easily recolonized the emptied islands, apparently by aerial transport. However, it is difficult to determine,
based on the literature, if ballooning is their primary mode of dispersal. Glick
(1939) did not list Nephila in his study ofthe aerial arthropods over Tallulah,
LA (Madison Parish), but over 900 spiders in that study were left unidentified.
Kuntner and Agnarsson (2011) show that as a genus, Nephila is an excellent
colonizer and suggest ballooning as the most likely method of dispersal, although it has not yet been observed to do so. Additionally, N. clavipes may use
an intermediate form of ballooning, where the spiderling makes a silk thread
that is floated in the air; when the thread contacts a structure, the line is made
fast and the spider walks across (Moore 1977).
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Materials and Methods
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In order to fully document the current distribution of Nephila clavipes, we
searched for distributional records in the published literature (including early
natural history narratives going back to 1768), natural history museum collections, and online databases, and solicited observations from experienced field
biologists. We examined specimens from the American Museum of Natural
History, Florida State Collection of Arthropods, and the National Museum of
Natural History. We also performed Google searches to find blogs, looked at social photograph-posting websites such as Flickr, Pbase, Photobucket, Picasa, and
YouTube, and used entomology and hobby-oriented websites such as Bugguide.
net, Dave's Garden, and Spiderzrule. Our last web search was conducted 3 January 2012.
For each distributional record, we recorded its source, date, state, county or
parish, specific locality (if available), photographer or collector (in the case of
specimens), and any additional notes (Bugguide.net proved to be a particularly
rich source of natural history notes). By recording the photographer and examining each photograph and associated data, we were able to determine if the
photographer had posted multiple views of the same spider, and thus avoided
duplicate entries.
,:
Results
We accumulated 945 records of Nephila clavipes with county-level locality
data in the continental United States. To date, A'^. clavipes has been recorded in
204 counties of 11 states (Fig. 2, Appendix 1). Of these records, 50.7% were
from website photographs (includes social and science-based), 37.9% were from
natural history museum collections, 5.8% from personal communications with
other scientists, and 5.6% from the literature.
The early writers on the natural history of the southeastern United States
failed to record the presence of Nephila clavipes. However, William Bartram
(1943), in his report to Dr. John Fothergill on his travels in Georgia and Florida
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K.A. Bakkegard and L.J. Davenport
555
during 1773-1774, described a spider that could be N. clavipes. While his large
yellow and black streaked spider sounds much like Argiope aurantia, his description includes "their legs very long and armed with prickles of stiff black hair,"
which is characteristic oí Nephila. He possibly encountered both species but only
mentioned one, later mistakenly fusing their traits together. Philip Henry Gosse
(1859), after spending nine months of 1838-1839 in Dallas County, AL, did not
mention Nephila, even though he described over 120 other terrestrial invertebrates and noted the presence of spiders in mud dauber nests, ¡A
The earliest unambiguous record for Nephila clavipes in the United States is
by Wilder (1865). In 1863, Dr. Burt G. Wilder was a surgeon in the Union Army
stationed on Folly Island, SC, near Charleston. Wilder specifically wrote that he
found no mention of his spider in any publications by earlier workers such as
Nicholas Hentz (Burgess 1875), who was later considered the father of American
arachnology; Hentz spent much of the 1840s in northern and central Alabama
(Davenport 2001). Wilder (1867) was fascinated by the spider's strong yellow
silk and thought it could be used for some practical purpose.
The oldest specimen for which we found a record was collected by the
arachnologist George Marx (1838-1895) in 1885 and is likely to be one of
the specimens referred to by McCook (1893). As an historical aside, Marx's
entire collection, consisting of over 1000 arachnids, was offered up for sale
for $1500 less than a year after his death (Riley et al. 1895). McCook (1893)
discussed Nephila clavipes (and several related species now understood to be
N. clavipes), stating that it had been collected in Louisiana, Texas, and Key
West, FL; he also noted that the spider was probably limited to the Gulf Coast
and did not appear to be abundant. For what he called JV. wilderi, he stated its
range to be "along Southern Atlantic and Gulf Coasts, and southwesterly to
Southern California." A detailed species account, including photographs of its
web, can be found in Comstock (1920). However, Comstock provided few specifics, stating that it was "widely distributed through the southern states."
Equally noteworthy are early twentieth-century studies that do not Mst Nephila
clavipes (or any of its synonyms). These include Banks (1900), who collected
133 species of spiders in the southern half of Alabama, mostly from the Auburn
(Lee County) and Mobile (Mobile County) areas. Many of these specimens were
taken in the autumn, when N. clavipes is most noticeable. Another example is
James Emerton's (1902) book, a guide to the common spiders likely to be found
from Georgia to as far west as the Rocky Mountains. Bishop and Crosby (1926)
did not find the spider while collecting in the Okefinokee Swamp of southern
Georgia during the summer of 1912, although there are several specimens from
the Okefinokee Swamp, collected in 1912 by persons unknown, in the American
Museum of Natural History (AMNH) collection. They also did not list the spider
in their report, which included records from a collection at the Alabama State
Museum (now the University of Alabama Natural History Collection) that covered many southeastern states (Alabama, Florida, Georgia, Kentucky, Louisiana,
Missouri, North Carolina, Tennessee, and Texas; specimens from 1903-1925).
Chamberlin and Ivie (1944) reviewed the spiders of Georgia, also including most
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Southeastern Naturalist
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of South Carolina, parts of north Florida, and a small section of North Carolina.
Their review included field work and a re-examination of drawings by British
naturalist John Abbot (1751-1840), who lived in what is now Screven County,
GA and is believed to have done most of his work on spiders during the late 1700s
(Chamberlin and Ivie 1944). Abbot did not include Nephila in his 567 drawings
of spiders, and Chamberlin and Ivie (1944) provide only two records: Leon
County, FL and Ware County, GA. Both specimens are in fhe AMNH collection.
While museum collections provided "hands-on" documentation, websites
varied greatly in their information content, with Bugguide.net, Flickr.com, and
Spiderzrule.com being the most useful. Hosted by the Iowa State University Department of Entomology, Bugguidc.net encourages a scientific approach to data
collection, with many submitters providing interesting natural history notes on
the density of webs (often noted if high), prey items, and whether or not the spider had been previously noticed in their area. The geotagging function of Flickr
was likewise helpful. Some hobby-oriented websites, such as Dave's Garden,
included locality data, which became an important part of the discussion about
fhe spider pictured. Other websites were less helpful, with the largest problem
being missing locality data. Sites such as Photobuckct, Facebook, and YouTube
contained images but no locality data.
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Discussion
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Using historical records, documented specimens and current observations, we
have developed the most comprehensive range map oí Nephila clavipes to date.
Prior to our study, the most complete map was by Levi (1980), who noted that the
spider was found only in the warmer parts of the southeastern United States. His
most northern record was in Hyde County, NC; his most northwestern record was
from Navajo County, AZ. Our study adds many historical records, most notably,
central Alabama by Archer (1940), and shows fhe northward migration of this
spider out of what had been an exclusively coastal plain distribution.
The earliest records of this spider are centered on major seaports. Jones (1859)
stated that Nephila clavipes (then Epeiria clavipes) was the best known and most
attractive spider on Bermuda, and had been known from that island as early as
fhe seventeenth century. Since many naturalists of that time, including Charles
Darwin (1845), noted the capturing of spiders in ships' sails and riggings, we
suggest that the infroduction of N. clavipes fo fhe east Atlantic and Gulf coasts
may have been promoted by sailing ships. We can also estimate when fhe spider
was infroduced to Alabama. Archer (1940) wrote: "In the last fifteen years, it has
established itself in the urban gardens in Mobile, much fo the consternation of the
local citizenry. In the last two years if has been invading gardens in Montgomery." Thus, we can dafe fhe introduction of A^. clavipes fo fhe Mobile area between
1900 and 1925 and its northward spread fo Monfgomery by 1938.
There are also a few odd records. It is hard to explain fhe Show Low, AZ record
(MCZ 25983, hftp://mcz.harvard.edu) collected by F. Matzone in August 1965.
Show Low, a cify in east central Arizona, elevafion 1950 m, is in fhe Ponderosa
2012
•
K.A. Bakkegard and L.J. Davenport
557
Pine ecosystem and thus experiences cold winter weather. The arachnologist
Donald C. Lowrie collected Nephila clavipes in 1935 in California (AMNH collection). The tag states Fillmore County. While there is no Fillmore County in
California, there is a city of Fillmore in Ventura County in southern California.
Perhaps the spider was introduced via the citrus industry, which brought in plants
from all over the world, including Florida and Brazil (Coit 1915), areas where
A^. clavipes was already present. Another enigmatic area is northern Mississippi.
Dorris and McGaha (1965) included A^. clavipes in their list of spiders from five
counties in north central Mississippi, but do not indicate which county contained
what species. However, Dorris' (1963) Master's Thesis on which Dorris and McGaha (1965) is based indicates that only one specimen of A^. clavipes was found,
in Lafayette County. May's (1933) Thesis and discussions with current Mississippi arachnologists indicate that A^. clavipes is not present, at least in noticeable
numbers, in northern Mississippi. It is also clear that southern Mississippi has
been under-collected historically.
One pitfall that we found in using pictures from the public domain is that the
locality data is not always what it seems to be. We found a geotagged photo on
Flickr with a locality of Los Angeles, CA. In this case, all the data were correct;
it was indeed a photograph oí Nephila clavipes and the photographer was in Los
Angeles. However, the photograph was that of A', clavipes on public display in
the spider pavilion at the Natural History Museum of Los Angeles County. The
second case was of a beautiful photograph posted on the website Dave's Garden
with a locality of Greensboro, NC. Unfortunately, the photographer, who lived at
the time in Greensboro, has since passed away, so we are unable to confirm that
the photograph was actually taken there. Otherwise, that would be an intriguing
record, well outside the expected range of this spider.
Further studies of this spider should concentrate on these five areas because
of their large number of spider records: Houston (Harris County), TX; Charleston (Charleston County), SC; New Orleans (Orieans Parish), LA; Gainesville
(Alachua County), FL; and Miami (Miami-Dade County), FL. We recommend
monitoring the climate records of each of these areas for future comparisons.
In Nephila clavipes, female fecundity significantly increases with larger body
size, while later-maturing females are less fecund (Higgins 2000). As the spider expands its range northward, it will encounter stronger seasonality. Thus,
we predict a slow spread. However, in the southern parts of its range, eggs and
spiderlings should experience a decrease in winter mortality, providing larger
source populations.
Future documentation of the distribution oí Nephila clavipes will depend on
enlisting the public to provide scientific observations. Such efforts can be made
formally in the manner of the Cornell Laboratory of Ornithology with its citizen
science program. Or they can be made informally by monitoring websites, such
as those devoted to photograph- and video-sharing, social networking, or individual blogs. With range expansion, laypeople unfamiliar with A^. clavipes will
soon encounter it for the first time. Because a large spider elicits a strong emotional reaction—either fascination or revulsion—many people will take a picture
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Vol. 11, No. 4
or video of the spider and post it online, to share it with others, get it identified, or
to determine if it is dangerous. We therefore ask that biologists monitor websites,
or whatever means of scientific and social networking will be available, in order
to track the spread of this spider.
Acknowledgments
Thank you to these individuals (listed alphabetically by last name) and institutions
that checked collections, loaned specimens, provided locality data from specimens under
their curation, or provided locality data from their field notes: J.K. Barnes (University
of Arkansas Arthropod Museum), V.M. Bayless (Louisiana State Arthropod Museum), J.
Beccaloni (Natural History Museum, London), J.H. Boone (The Field Museum of Natural
History), J. A. Coddington (National Museum of Natural History), A. Dean (Texas A & M
University), A.R. Diamond (Troy University), G.B. Edwards (Florida State Collection of
Arthropods), Z. Falin and J. Thomas (University of Kansas Biodiversity Institute), D.R.
Folkerts (Auburn University), J.G. Godwin (Auburn University), H. Guarisco (Sternberg
Museum of Natural History), M. Hodge (Louisiana School for Math, Science, and the
Arts), W.M. Howell (Samford University), J.G. King (Vanderbilt University), J. Knight
(South Carolina State Museum), M.W. LaSalle (Pascagoula River Audubon Center), M.B.
Layton (Mississippi State University), J.V. McHugh and C.L. Smith (Georgia Museum of
Natural History), P. Miller (Northwest Mississippi Junior College), J.C. Morse and A.B.
Harrison (Clemson University Arthropod Collection), G.R. Mullen (Auburn University),
M.F. O'Brien (University of Michigan Museum of Zoology), S. Peyton (Mississippi
Museum of Natural Science), N.I. Platnik and L.N. Sorkin (American Museum of Natural History), T. Pucci (Cleveland Museum of Natural History), R.J. Pupedis (Peabody
Museum of Natural History), J.E. Rawlins (Carnegie Museum of Natural History), C.H.
Ray (Aubum University), J.R. Reddell (Texas Memorial Museum), N. Rios (Tulane University Museum of Natural History), T.L. Schiefer (Mississippi Entomological Museum),
R.M. Shelley (North Carolina State Museum of Natural Sciences), K.B. Simpson (Enns
Entomology Museum), G. Stratton (University of Mississippi), R. Tumlison (Henderson
State University), D. Ubick (California Academy of Sciences), G.M. Ward (University of
Alabama), and J.D. Weintraub (Academy of Natural Sciences of Drexel University). M.E.
Martin (LSU Library, Special Collections) assisted in finding an elusive locality. Special
thanks to L.N. Sorkin who hosted K.A. Bakkegard's visit to the AMNH collection and the
Samford University Interlibrary Loan department. Samford University provided essential
funding for this project.
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Malcolm, J.R., and L.F. Pitelka. 2000. Ecosystems and global climate change: A review
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,^^
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Southeastern Naturalist
562
Appendix 1. Earliest documented record for Nephila clavipes by state and county (or parish).
Number of records is indicated for those counties when the number of records is >5. References are
listed in the same order as the counties. If only one reference is listed for multiple counties, then
that reference applies to all counties. For pers. comm., the person's initials and affiliation are in the
Acknowledgments. For key to other non-literature reference abbreviations, see Appendix 2 below.
Earliest
record
Counties (# of records)
Reference
AMNH
AMNH
Archer (1940)
1978
1989
1990
1996
2002
Baldwin (14)
Dale (7), Houston, Mobile (10)'
Barbour, Covington, Escambia,
Montgomery (6), Pike, Tuscaloosa
Lee
Henry
Geneva
Coffee
Jefferson
2003
2004
2006
Bullock
Conecuh, Russell
Clarke, Crenshaw, Monroe, Tallapoosa
2008
Choctaw, Lowndes, Wilcox
2009
2010
Autauga, Calhoun, Macon, Perry, Shelby,
Washington
Elmore
AUEM
Diamond, pers. comm.^
Diamond, pers. comm.
Diamond, pers. comm.
Samford University photo voucher
Diamond, pers. comm.
Diamond, pers. comm. "
Diamond, pers. comm. (Clarke,
Crenshaw); Folkerts, pers.
comm.' (Monroe, Tallapoosa)
Ray, pers. comm.* (Choctaw, Lowndes); Diamond, pers.
comm. (Wilcox)
BG; Ray, pers. comm.; F, BG,
BG, F
Godwin, pers. comm.'
AR
1966
1979
2009
Clark'
Union'^
Ashley, Union
Dorris (1968)
Dorris (1980)
Tumlison and Robison (2010)
AZ
1965
Navajo
MCZ
CA
1935
Ventura?
AMNH
FL
1893
1903
1904
1911
1925
1926
1927
1933
1934
1940
1941
1943
1944
1945
1946
1950
1951
Monroe (48)
Miami-Dade (55)
Charlotte (5), Citrus
Brevard(13)
Liberty (9)
Alachua (65), Putnam (8)^ St. Lucie (9)
Seminóle (9)
Lee (9), Leon (16), St. Johns (7)
Manatee"", Orange (15)
Calhoun, Sarasota (13)
Collier (5)
Highlands (28), Okaloosa
Hendry, Santa Rosa (5)
Escambia
Osceola
Levy (13)
Jackson, Pinellas (13)
McCook(1893)
; ,
AMNH
Banks (1904)
AMNH
AMNH
AMNH
AMNH
AMNH
USNM, FSCA
Archer (1940), AMNH
FMNH
AMNH
Kuntner (2003), AMNH
AMNH
AMNH
FSCA
, FSCA ,
State
AL
1932
1939
1940
(;
^ •.
•
:
2012
State
FL
GA
LA
K.A. Bakkegard and L.J. Davenport
563
Earliest
record
Counties (# of records)
Reference
1952
1953
1954
1963
1964
1969
1972
1973
1974
1978
1981
1984
1986
1989
1990
1992
2004
2006
2007
2008
2010
Palm Beach (17)
Broward(18)
Volusia(8)
Columbia, Glades, Marion (17)
Martin
Indian River
Lake(7)
Baker
WakuUa
Jefferson'
':
Pasco
Hillsborough(13)
Holmes
DeSoto
Gilcrist
Gadsden
Duval (7), Flagler
Hernando
Nassau, Polk (7), Sumter, Suwannee
Bay, Clay, Union
Okeechobee
FMNH
FMNH
FSCA
"
FSCA
FSCA
FSCA
CAS
MCZ
Farr(1977)
AMNH
FSCA
Kuntner (2003)
Howell, pers. comm.'"
FSCA
BG
NASDB
JAX, P
1912
1916
1932
1935
1946
1950
1960
1967
1972
1973
1976
1980
1984
1998
2002
2003
2005
2006
2007
2008
2009
2010
Unk
Charlton (8)
Thomas
Ware
Screven
Baker, Ben Hill, Clay
Mclntosh (5)
Long
Liberty
Dougherty
Bulloch
Oconee
Glynn (6)
Chatham (11)
Camden
Jasper
Bryan, Coffee, Crisp, Lowndes
Sumter
Tattnall
Stewart
• Jh.. ^8'.
Jones
Laurens
' :.ad
Clarke
AMNH
AMNH
USNM
AMNH
FMNH
FSCA
UGA
UGA
AMNH ^
Folkerts, pers. comm.
FI
Smith, pers.comm."
1885
1923
1924
1933
1940
1959
1966
Orleans (11)
East Baton Rouge (12)
Terrebonne
Cameron
Jefferson (7)
Ascension (5)
Jefferson Davis, St. Landry
Lee
-.iad
F
-í.'-
KM, BG, OBS, F
BG, F, F
BG
USNM
, -•
AMNH
BG
GB
BG, GB, BG, F
P
W
'
.-
•
BG
•
•
BG
BG
Smith, pers. comm.
Kuntner (2003)
FMNH
FMNH
Kuntner (2003)
AMNH
CAS
FSCA, LSAM
' :--' '
•;
Vol. 11, No. 4
Southeastern Naturalist
State
LA
Earliest
record
Counties (# of records)
Reference
1975
1981
1996
2000s
2004
2005
2007
2008
2009
2011
West Feliciana
Plaquemines
Bossier
Bienville, Natchitoches
Livingston
St. James
Tangipahoa
Iberia, St. Tammany, Washington
Ouachita, St. Martin (5)
St. Charles
• -
AMNH
MCZ
Hardy (2010)
Hodge, pers. comm.'"
SR
F
1931
1932
1961
1963
1966
2002
2004
2005
2007
2008
2009
George
Jackson (7)
Yazoo
Lafayette
2011
Hancock, Harrison, Pearl River, Stone
1966
1976
2003
2004
2007
2010
Carteret (8)
New Hanover (13)
Brunswick (6)
Columbus, Onslow
Bladen
1863
1935
1953
1977
1978
1985
2004
2005
2006
2009
2010
2011
Charleston (25)
Hampton
Aiken
Beaufort (8)
Georgetown (10)
Clarendon, Lexington, Orangeburg
Horry
Jasper, Sumter
Berkeley
Colleton
Dorchester, Richland
Barnwell
Wilder (1865)
AMNH
UGCA
Gaddy and Morse (1985)
CUAC
Gaddy and Morse (1985)
SR
CMNH, SR
SR
F
1913
1936
1946
1955
1958
1970
1971
1972
Chambers
Harris (18)
Jefferson
Bee
Liberty
Hardin, Nacogdoches
Brazoria, Matagorda
Galveston (5)
AMNH
AMNH
MCZ
USNM
MCZ
Brown (1974)
NASDB
Moore (1977)
B G
•'& 2
SP, F, BG
•
£
WTB, F
•
F
•
.
••
•'"
.
•..,.
'""I"
MS
NC
sc
TX
•
-
Jackson
:"• • ' •
A d a m s
-
AMNH
YPM
Dorris (1967)
Dorris (1963)
"
•
'
••'
•
-
-•
•
BY
•
':
•"•" '
MEM
•
"'• .•'•'• ' ' ''
Forrest
Walthall
>
Claiborne
Greene, Lamar, Noxubee, Perry
Clarke, Hinds, Lauderdale, Simpson
H y d e
'
..
SR
-
F
..
•''•
<•'
'<»•
-
F
F, B G , F, B G
'••
:i^ii"
'
-¡-¡SJ'ujA
'••
-
1
1.
•"'
Schiefer, Peyton, Schiefer, Layton - all pers. comm.'^
LaSalle, pers. comm.'''
1
Levi (1980)
Levi (1980)
NCSP
BG
NCSP, BG
NCSP
F
F
iJ'
'•!
K.A. Bakkegard and L.J. Davenport
State
TX
565
Earliest
record
Counties (# of records)
Reference
1980
2004
2005
2007
2009
Willacy
Jasper
Fort Bend (15)
Lavaca, Montgomery, Tyler
Brazos
NASDB
W
F
SR, BG, F
TAMUIC
'USNM has an undated specimen that may have been collected by the arachnologist George Marx,
who died 3 Jan 1895. There is a label in the same style and handwriting as another Florida specimen he collected.
'Tumlison & Robison (2010) discusses the validity of these records.
'USNM has an undated specimen (USNM 2055693 -NE 149) collected by G. Marx from this
county.
"USNM has an undated specimen (USNM 2055693-2754) collected by R.W. Shufeldt (1850-1934).
It was likely to have been collected well before 1934. The next most recent record for this county
is 2007.
'An undated specimen identified by Levi in 1978. Based on writing style on the tag, it was collected
earlier than 1978. There are currently no other records for this county. -:,/>
.j-: :
•"A.R. Diamond, Troy University, Troy, AL
' ;i,i.'
'D.R. Folkerts, Auburn University, Auburn, AL
•'•• >
*C.H. Ray, Auburn University, Auburn, AL
'J.G. Godwin, Auburn University, Auburn, AL - •
• .. .
••• - '•': ; . - • • ' • • • :
; ;
'-,.'.''
'"W.M. H o w e l l , Samford University, H o m e w o o d , A L
•; -,. , ,-'•
•••,':''
"C.L. Smith, Georgia Museum of Natural History, Athens, GA
"M. Hodge, Louisiana School of Math, Science, and the Arts, Natchitoches, LA
"T.L. Schiefer, Mississippi Entomological Museum, Starkville, MS: S. Peyton, Mississippi Museum of Natural Science, Jackson, MS; M.B. Layton, Mississippi State University, MS
'"M.W. LaSalle, Pascagoula River Audubon Center, Moss Point, MS
• •
'-.x
'it
•i.hl-r.-.
•''':..
r;.-
•"
r..,,.i:
• V
566
--
Vol. 11, No. 4
Southeastern Naturalist
Appendix 2. Key to non-literature references used in Appendix 1.
Abbrev.
Full name of reference
Location
AMNH
AUEM
BG
BY
CAS
CMNH
CUAC
DG
F
FI
FMNH
FSCA
GB
JAX
KM
American Museum of Natural History
Auburn University Entomology Museum
BugGuide.net
Backyardnature.net
California Academy of Sciences
Carnegie Museum of Natural History
Clemson University Arthropod Collection
Dave's Garden
Flickr
Forestry Images
Field Museum of Natural History
Florida State Collection of Arthropods
Giff Beaton's webpage
Jacksonville Shell Club
Kirk M. Rogers' webpages
LSAM
MCZ
MEM
NASDB'
Louisiana State Arthropod Museum
Museum of Comparative Zoology
Mississippi Entomological Museum
Nearctic Spider database
OBS
NCSP
O.B. Sirius blog
North Carolina State Parks NRID system
P
SP
SR
TAMUIC
UGCA
USNM
W
WTB
YPM
Picasa
StockphotoPro
Spiderzrule
Texas A&M University Insect Collection
University of Georgia Collection of Arthropods
National Museum of Natural History
Webshots
What's that bug?
Yale Peabody Museum of Natural History
New York, NY
Auburn, AL
http://bugguide.net
http://backyardnature.net
.;-i' ;
San Francisco, CA
^
^ ;.
Pittsburgh, PA
"'•, ;.., .,
Clemson, SC
http://davesgarden.com
http://www.flickr.com
''"'
http://www.forestryimages.org
''
Chicago, IL
Gainesville, FL
http://www.giffbeaton.com
http://www.jaxshells.org
http://www.kiroastro.com/writings/
georgiaj3hotography.html
Baton Rouge, LA
Cambridge, MA
Starkville, MS
http://www.canadianarachnology.
org/data/canada_spiders
http://obsirius.blogspot.com
http://www.dpr.ncparks.gov/nrid/
gallery.php
<>
https://picasaweb.google.com
http://www.stockphotopro.com
http://spiderzrule.com
College Station, TX
^'"
Athens, G A
'
'Jf'
Washington, DC
http://www.webshots.com
•
http://whatsthatbug.com
New Haven, CT
'NASBD has been offline since 17 March 2010. This dataset can be accessed through the Global
Biodiversity Information Facility (http://www.gbif.org/)
)
•
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