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2012 SOUTHEASTERN NATURALIST 11(4):551–566
Nephila clavipes (Araneae: Nephilidae):
A Model Species for Monitoring Climate Change in the
Southeastern United States
Kristin A. Bakkegard1,* and Lawrence J. Davenport1
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 N. 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.
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
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. 1A) are large
(to 25 mm cephalothorax 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
1Department of Biological and Environmental Sciences, Samford University, Birmingham,
AL 35229. *Corresponding author - email@example.com.
552 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 N. 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, N. 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.J. Davenport 553
while temperatures below 0 °C are fatal to adults (Moore 1977). In the summer, 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, N. 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
N. clavipes webs often contain kleptoparasites (Agnarsson 2003).
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.
554 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, N. 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 of the 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).
Materials and Methods
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
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
We accumulated 945 records of Nephila clavipes with county-level locality
data in the continental United States. To date, N. 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
2012 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 of 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.
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 N. 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 list 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
556 Southeastern Naturalist Vol. 11, No. 4
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 the 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, Bugguide.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
the spider pictured. Other websites were less helpful, with the largest problem
being missing locality data. Sites such as Photobucket, Facebook, and YouTube
contained images but no locality data.
Using historical records, documented specimens and current observations, we
have developed the most comprehensive range map of 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 the 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
the 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 introduction of N. clavipes to the east Atlantic and Gulf coasts
may have been promoted by sailing ships. We can also estimate when the spider
was introduced to Alabama. Archer (1940) wrote: “In the last fifteen years, it has
established itself in the urban gardens in Mobile, much to the consternation of the
local citizenry. In the last two years it has been invading gardens in Montgomery.”
Thus, we can date the introduction of N. clavipes to the Mobile area between
1900 and 1925 and its northward spread to Montgomery by 1938.
There are also a few odd records. It is hard to explain the Show Low, AZ record
(MCZ 25983, http://mcz.harvard.edu) collected by F. Matzone in August 1965.
Show Low, a city in east central Arizona, elevation 1950 m, is in the 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
N. clavipes was already present. Another enigmatic area is northern Mississippi.
Dorris and McGaha (1965) included N. 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 Mc-
Gaha (1965) is based indicates that only one specimen of N. clavipes was found,
in Lafayette County. May’s (1933) Thesis and discussions with current Mississippi
arachnologists indicate that N. 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 of Nephila clavipes and the photographer was in Los
Angeles. However, the photograph was that of N. 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 (Orleans 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
Future documentation of the distribution of 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 N. 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
558 Southeastern Naturalist 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.
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 (Auburn 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.
Agnarsson, I. 2003. Spider webs as habitat patches—The distribution of kleptoparasites
(Argyrodes, Theridiidae) among host webs (Nephila, Tetragnathididae). Journal of
Archer, A.F. 1940. The Argiopidae or orb-weaving spiders of Alabama. Alabama Museum
of Natural History 14:1–41.
Banks, N. 1900. Some Arachnida from Alabama. Proceedings of the Academy of Natural
Sciences of Philadelphia 52:529–543.
Banks, N. 1904. The Arachnida of Florida. Proceedings of the Academy of Natural Sciences
of Philadelphia 56:120–147.
Bartram, W. 1943. Travels in Georgia and Florida, 1773–74: A Report to Dr. John
Fothergill. Transactions of the American Philosophical Society 33, part II:121–242.
Reprinted 1996, Penguin Putnam, New York, NY. 701 pp.
2012 K.A. Bakkegard and L.J. Davenport 559
Bishop, S.C., and C.R. Crosby. 1926. Notes on the spiders of the southeastern United
States with descriptions of new species. Journal of the Elisha Mitchell Scientific Society
Brown, K.M. 1974. A preliminary checklist of spiders of Nacogdoches, Texas. Journal
of Arachnology 1:229–240.
Burgess, E. (Ed.). 1875. The spiders of the United States: A collection of the arachnological
writings of Nicholas Marcellus Hentz, M.D. with notes and descriptions by James
H. Emerton. Occasional Papers of the Boston Society of Natural History II. Boston,
MA. 177 pp. with 21 plates.
Carrel, J.E. 1978. Behavioral thermoregulation during winter in an orb-weaving spider.
Symposium of the Zoological Society of London 42:41–50.
Chamberlin, R.V., and W. Ivie. 1944. Spiders of the Georgia region of North America.
Bulletin of the University of Utah 35:1–267.
Coit, J.E. 1915. Citrus Fruits. The MacMillan Company, New York, NY. 503 pp.
Comstock, J.H. 1920. The Spider Book. Doubleday, Page, and Company, Garden City,
NY. 721 pp.
Darwin, C.R. 1845. Journal of Researches into the Natural History and Geology of the
Countries Visited during the Voyage of H.M.S. Beagle Round the World, under the
Command of Capt. Fitz Roy, R.N. 2nd edition. John Murray, London, UK. 519 pp.
Davenport, L.J. 2001. Trapdoor spiders. Alabama Heritage 59:50–52.
Davenport, L.J. 2007. Golden silk orbweavers (and climate change). Alabama Heritage
Decae, A.E. 1987. Dispersal: Ballooning and other mechanisms. Pp. 348–356, In W.
Nentwig (Ed.). Ecophysiology of Spiders. Springer-Verlag, Berlin, Germany. 448 pp.
Delcourt, P.A., and H.R. Delcourt. 1981. Vegetation maps for eastern North America:
40,000 yr B.P. to the present. Pp. 123–165, In R.C. Romans (Ed.). Geobotany II. Plenum
Press, New York, NY. 263 pp.
Dorris, P.R. 1963. Taxonomy, ecology, and distribution of the spiders in five counties of
northern Mississippi. M.Sc. Thesis. University of Mississippi, Oxford, MS. 164 pp.
Dorris, P.R. 1967. The spiders of Mississippi. Ph.D. Dissertation. University of Mississippi,
Oxford, MS. 283 pp.
Dorris, P.R. 1968. A preliminary study of the spiders of Clark County, Arkansas, compared
with a five-year study of Mississippi spiders. Arkansas Academy of Science
Dorris, P.R. 1980. A continuation of spider research in Arkansas: Gulf coastal plains.
Arkansas Academy of Science Proceedings 34:108–112.
Dorris, P.R., and Y.J. McGaha. 1965. A list of spiders collected in northern Mississippi.
Transactions of the American Microscopical Society 84:407–408.
Emerton, J.H. 1902. The Common Spiders of the United States. Ginn and Company,
Boston, MA. 225 pp.
Farr, J.A. 1977. Social behavior of the Golden Silk Spider, Nephila clavipes (Linnaeus)
(Araneae, Araneidae). Journal of Arachnology 4:137–144.
Gaddy, L.L., and J.C. Morse. 1985. Common spiders of South Carolina with an annotated
checklist. South Carolina Agricultural Experiment Station Technical Bulletin 1094.
Gates, D.M. 1993. Climate Change and its Biological Consequences. Sinauer Associates,
Sunderland, MA. 280 pp.
Glick, P.A. 1939. The distribution of insects, spiders, and mites in the air. USDA Technical
Bulletin 673. Washington, DC. 150 pp.
560 Southeastern Naturalist Vol. 11, No. 4
Gosse, P.H. 1859. Letters from Alabama, (US) Chiefly Relating to Natural History.
Morgan and Chase, London, UK. (Reprinted 1993, University of Alabama Press,
Tuscaloosa, AL). 324 pp.
Hardy, L.M. 2010. Additions to the spider fauna of northwestern Louisiana. Southeastern
Naturalist 9 (Monograph):1–40.
Higgins, L. 1992. Developmental plasticity and fecundity in the orb-weaving spider
Nephila clavipes. Journal of Arachnology 20:94–106.
Higgins, L. 2000. The interaction of season length and development time alters size at
maturity. Oecologia 122:51–59.
Higgins, L.E., and E. Ezcurra. 1996. Mathematical simulation of thermoregulatory behavior
in an orb-weaving spider. Functional Ecology 10:322–327.
Higgins, L., and C. Goodnight. 2010. Nephila clavipes females have accelerating dietary
requirements. Journal of Arachnology 38:150–152.
Hill, E.M., and T.E. Christenson. 1981. Effects of prey characteristics and web structure
on feeding and predatory responses of Nephila clavipes spiderlings. Behavioral Ecology
and Sociobiology 8:1–5.
Howell, W.M., and R.L. Jenkins. 2004. Spiders of the Eastern United States: A Photographic
Guide. Pearson Education, Boston, MA. 363 pp.
Jones, J.M. 1859. The Naturalist in Bermuda: A Sketch of the Geology, Zoology, and Botany,
of that Remarkable Group of Islands. Reeves and Turner, London, UK. 200 pp.
Kaston, B.J. 1978. How to Know the Spiders, 3rd Edition. Wm. C. Brown, Dubuque,
IA. 272 pp.
Kirchner, W. 1987. Behavioural and physiological adaptations to cold. Pp. 66–77, In W.
Nentwig (Ed.). Ecophysiology of Spiders. Springer-Verlag, Berlin, Germany. 448 pp.
Krakauer, T. 1972. Thermal response of the orb-weaving spider Nephila clavipes. American
Midland Naturalist 88:246–250.
Kuntner, M. 2003. A preliminary specimen database of true nephiline spiders (Tetragnathidae).
Available online at http://www.gwu.edu/~clade/spiders/taxonomyPeet.
htm. Accessed 3 January 2012.
Kuntner, M., and I. Agnarsson. 2011. Phylogeography of a successful aerial disperser:
The golden orb spider Nephila on Indian Ocean islands. BMC Evolutionary Biology
Kuntner, M., J.A. Coddington, and G. Horminga. 2008. Phylogeny of extant nephilid
orb-weaving spiders (Araneae, Nephilidae): Testing morphological and ethological
homologies. Cladistics 24:147–217.
Levi, H.W. 1980. The orb-weaver genus Mecynogea, the subfamily Metinae, and the
genera Pachygnatha, Glenognatha, and Azilia of the subfamily Tetragnathinae north
of Mexico (Araneae: Araneidae). Bulletin of the Museum of Comparative Zoology
Malcolm, J.R., and L.F. Pitelka. 2000. Ecosystems and global climate change: A review
of potential impacts on US terrestrial ecosystems and biodiversity. Pew Center on
Global Climate Change, Arlington, VA. 41 pp.
May, U. 1933. Mississippi spiders. M.Sc. Thesis. Mississippi State College, Starkville,
MS. 53 pp.
McCook, H.C. 1893. American Spiders and their Spinningwork, Volume III. Academy of
Natural Sciences, Philadelphia, PA. 407 pp.
Moore, C.W. 1977. The life cycle, habitat, and variation in selected web parameters in the
spider Nephila clavipes Koch (Araneidae). American Midland Naturalist 98:95–108.
2012 K.A. Bakkegard and L.J. Davenport 561
Myers, L., and T. Christenson. 1988. Transition from predatory juvenile male to matesearching
adult in the orb-weaving spider Nephila clavipes (Araneae, Araneidae).
Journal of Arachnology 16:254–257.
Parmesan, C., and H. Galbraith. 2004. Observed impacts of global climate change in the
US. Pew Center on Global Climate Change, Arlington, VA. 56 pp.
Peters, R.L. 1991. Consequences of global warming for biological diversity. Pp. 99–118,
In R.L. Wyman (Ed.). Global Climate Change and Life on Earth. Routledge, Chapman,
and Hall, New York, NY. 282 pp.
Pulz, R. 1987. Thermal and water relations. Pp. 26–55, In W. Nentwig (Ed.). Ecophysiology
of Spiders. Springer-Verlag, Berlin, Germany. 448 pp.
Riley, C.V., L.O. Howard, E.A. Schwartz, and T. Gill. 1895. The Marx collection of
Arachnida. Canadian Entomologist 27:272.
Robinson, M.H., and B.C. Robinson. 1974. Adaptive complexity: The thermoregulatory
postures of the Golden-web Spider, Nephila clavipes, at low latitudes. American Midland
Simberloff, D.S., and E.O. Wilson. 1969. Experimental zoogeography of islands: The
colonization of empty islands. Ecology 50:278–296.
Tumlison, R., and H.W. Robison. 2010. New records and notes on the natural history of
selected invertebrates from southern Arkansas. Journal of the Arkansas Academy of
Wilder, B.G. 1865. On the Nephila plumipes, or silk spider, of South Carolina. Proceedings
of the Boston Society of Natural History 10:200–210.
Wilder, B.G. 1867. Two hundred thousand spiders. Harper’s New Monthly Magazine
562 Southeastern Naturalist Vol. 11, No. 4
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.
State record Counties (# of records) Reference
AL 1932 Baldwin (14) AMNH
1939 Dale (7), Houston, Mobile (10)1 AMNH
1940 Barbour, Covington, Escambia, Archer (1940)
Montgomery (6), Pike, Tuscaloosa
1978 Lee AUEM
1989 Henry Diamond, pers. comm.6
1990 Geneva Diamond, pers. comm.
1996 Coffee Diamond, pers. comm.
2002 Jefferson Samford University photo voucher
2003 Bullock Diamond, pers. comm.
2004 Conecuh, Russell Diamond, pers. comm.
2006 Clarke, Crenshaw, Monroe, Tallapoosa Diamond, pers. comm. (Clarke,
Crenshaw); Folkerts, pers.
comm.7 (Monroe, Tallapoosa)
2008 Choctaw, Lowndes, Wilcox Ray, pers. comm.8 (Choctaw,
Lowndes); Diamond, pers.
2009 Autauga, Calhoun, Macon, Perry, Shelby, BG; Ray, pers. comm.; F, BG,
Washington BG, F
2010 Elmore Godwin, pers. comm.9
AR 1966 Clark2 Dorris (1968)
1979 Union2 Dorris (1980)
2009 Ashley, Union Tumlison and Robison (2010)
AZ 1965 Navajo MCZ
CA 1935 Ventura? AMNH
FL 1893 Monroe (48) McCook (1893)
1903 Miami-Dade (55) AMNH
1904 Charlotte (5), Citrus Banks (1904)
1911 Brevard (13) AMNH
1925 Liberty (9) AMNH
1926 Alachua (65), Putnam (8)3, St. Lucie (9) AMNH
1927 Seminole (9) AMNH
1933 Lee (9), Leon (16), St. Johns (7) AMNH
1934 Manatee4, Orange (15) USNM, FSCA
1940 Calhoun, Sarasota (13) Archer (1940), AMNH
1941 Collier (5) FMNH
1943 Highlands (28), Okaloosa AMNH
1944 Hendry, Santa Rosa (5) Kuntner (2003), AMNH
1945 Escambia AMNH
1946 Osceola AMNH
1950 Levy (13) FSCA
1951 Jackson, Pinellas (13) FSCA
2012 K.A. Bakkegard and L.J. Davenport 563
State record Counties (# of records) Reference
FL 1952 Palm Beach (17) FMNH
1953 Broward (18) FMNH
1954 Volusia (8) FSCA
1963 Columbia, Glades, Marion (17) FSCA
1964 Martin FSCA
1969 Indian River FSCA
1972 Lake (7) CAS
1973 Baker MCZ
1974 Wakulla Farr (1977)
1978 Jefferson5 AMNH
1981 Pasco FSCA
1984 Hillsborough (13) Kuntner (2003)
1986 Holmes Howell, pers. comm.10
1989 DeSoto FSCA
1990 Gilcrist BG
1992 Gadsden NASDB
2004 Duval (7), Flagler JAX, P
2006 Hernando F
2007 Nassau, Polk (7), Sumter, Suwannee KM, BG, OBS, F
2008 Bay, Clay, Union BG, F, F
2010 Okeechobee BG
GA 1912 Charlton (8) AMNH
1916 Thomas AMNH
1932 Lee USNM
1935 Ware AMNH
1946 Screven FMNH
1950 Baker, Ben Hill, Clay FSCA
1960 McIntosh (5) UGA
1967 Long UGA
1972 Liberty AMNH
1973 Dougherty Folkerts, pers. comm.
1976 Bulloch FI
1980 Oconee Smith, pers.comm.11
1984 Glynn (6) USNM
1998 Chatham (11) AMNH
2002 Camden BG
2003 Jasper GB
2005 Bryan, Coffee, Crisp, Lowndes BG, GB, BG, F
2006 Sumter F
2007 Tattnall W
2008 Stewart BG
2009 Jones BG
2010 Laurens BG
Unk Clarke Smith, pers. comm.
LA 1885 Orleans (11) Kuntner (2003)
1923 East Baton Rouge (12) FMNH
1924 Terrebonne FMNH
1933 Cameron Kuntner (2003)
1940 Jefferson (7) AMNH
1959 Ascension (5) CAS
1966 Jefferson Davis, St. Landry FSCA, LSAM
564 Southeastern Naturalist Vol. 11, No. 4
State record Counties (# of records) Reference
LA 1975 West Feliciana AMNH
1981 Plaquemines MCZ
1996 Bossier Hardy (2010)
2000s Bienville, Natchitoches Hodge, pers. comm.12
2004 Livingston SR
2005 St. James F
2007 Tangipahoa BG
2008 Iberia, St. Tammany, Washington SP, F, BG
2009 Ouachita, St. Martin (5) WTB, F
2011 St. Charles F
MS 1931 George AMNH
1932 Jackson (7) YPM
1961 Yazoo Dorris (1967)
1963 Lafayette Dorris (1963)
1966 Jackson MEM
2002 Adams BY
2004 Forrest SR
2005 Walthall F
2007 Claiborne F
2008 Greene, Lamar, Noxubee, Perry F, BG, F, BG
2009 Clarke, Hinds, Lauderdale, Simpson Schiefer, Peyton, Schiefer, Layton
- all pers. comm.13
2011 Hancock, Harrison, Pearl River, Stone LaSalle, pers. comm.14
NC 1966 Carteret (8) Levi (1980)
1976 Hyde Levi (1980)
2003 New Hanover (13) NCSP
2004 Brunswick (6) BG
2007 Columbus, Onslow NCSP, BG
2010 Bladen NCSP
SC 1863 Charleston (25) Wilder (1865)
1935 Hampton AMNH
1953 Aiken UGCA
1977 Beaufort (8) Gaddy and Morse (1985)
1978 Georgetown (10) CUAC
1985 Clarendon, Lexington, Orangeburg Gaddy and Morse (1985)
2004 Horry SR
2005 Jasper, Sumter CMNH, SR
2006 Berkeley SR
2009 Colleton F
2010 Dorchester, Richland F
2011 Barnwell F
TX 1913 Chambers AMNH
1936 Harris (18) AMNH
1946 Jefferson MCZ
1955 Bee USNM
1958 Liberty MCZ
1970 Hardin, Nacogdoches Brown (1974)
1971 Brazoria, Matagorda NASDB
1972 Galveston (5) Moore (1977)
2012 K.A. Bakkegard and L.J. Davenport 565
State record Counties (# of records) Reference
TX 1980 Willacy NASDB
2004 Jasper W
2005 Fort Bend (15) F
2007 Lavaca, Montgomery, Tyler SR, BG, F
2009 Brazos TAMUIC
1USNM 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
2Tumlison & Robison (2010) discusses the validity of these records.
3USNM has an undated specimen (USNM 2055693 -NE 149) collected by G. Marx from this
4USNM 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
5An 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.
6A.R. Diamond, Troy University, Troy, AL
7D.R. Folkerts, Auburn University, Auburn, AL
8C.H. Ray, Auburn University, Auburn, AL
9J.G. Godwin, Auburn University, Auburn, AL
10W.M. Howell, Samford University, Homewood, AL
11C.L. Smith, Georgia Museum of Natural History, Athens, GA
12M. Hodge, Louisiana School of Math, Science, and the Arts, Natchitoches, LA
13T.L. Schiefer, Mississippi Entomological Museum, Starkville, MS; S. Peyton, Mississippi Museum
of Natural Science, Jackson, MS; M.B. Layton, Mississippi State University, MS
14M.W. LaSalle, Pascagoula River Audubon Center, Moss Point, MS
566 Southeastern Naturalist Vol. 11, No. 4
Appendix 2. Key to non-literature references used in Appendix 1.
Abbrev. Full name of reference Location
AMNH American Museum of Natural History New York, NY
AUEM Auburn University Entomology Museum Auburn, AL
BG BugGuide.net http://bugguide.net
BY Backyardnature.net http://backyardnature.net
CAS California Academy of Sciences San Francisco, CA
CMNH Carnegie Museum of Natural History Pittsburgh, PA
CUAC Clemson University Arthropod Collection Clemson, SC
DG Dave’s Garden http://davesgarden.com
F Flickr http://www.flickr.com
FI Forestry Images http://www.forestryimages.org
FMNH Field Museum of Natural History Chicago, IL
FSCA Florida State Collection of Arthropods Gainesville, FL
GB Giff Beaton’s webpage http://www.giffbeaton.com
JAX Jacksonville Shell Club http://www.jaxshells.org
KM Kirk M. Rogers’ webpages http://www.kiroastro.com/writings/
LSAM Louisiana State Arthropod Museum Baton Rouge, LA
MCZ Museum of Comparative Zoology Cambridge, MA
MEM Mississippi Entomological Museum Starkville, MS
NASDB1 Nearctic Spider database http://www.canadianarachnology.
OBS O.B. Sirius blog http://obsirius.blogspot.com
NCSP North Carolina State Parks NRID system http://www.dpr.ncparks.gov/nrid/
P Picasa https://picasaweb.google.com
SP StockphotoPro http://www.stockphotopro.com
SR Spiderzrule http://spiderzrule.com
TAMUIC Texas A&M University Insect Collection College Station, TX
UGCA University of Georgia Collection of Arthropods Athens, GA
USNM National Museum of Natural History Washington, DC
W Webshots http://www.webshots.com
WTB What’s that bug? http://whatsthatbug.com
YPM Yale Peabody Museum of Natural History New Haven, CT
1NASBD has been offline since 17 March 2010. This dataset can be accessed through the Global
Biodiversity Information Facility (http://www.gbif.org/)