Two species of linyphiid spiders, Disembolus bairdi Edwards, 1999 and Grammonota pallipes Banks, 1895, were discovered along the southwestern coast of Lake Michigan in Lake County, Illinois representing an Atlantic Coastal Plain disjunct distribution. A brief discussion of known collection sites, habitat preferences, and possible modes of dispersal are given

Draney, Michael L

Steffen, James F

English

Valparaiso University

The Great Lakes Entomologist Volume 47 Numbers 1 & 2 - Spring/Summer 2014 Numbers 1 & 2 - Spring/Summer 2014 Article 3 April 2014 Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders, Disembolus Bairdi and Grammonota Pallipes (Araneae: Linyphiidae) James F. Steffen Chicago Botanic Garden Michael L. Draney University of Wisconsin Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Steffen, James F. and Draney, Michael L. 2014. "Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders, Disembolus Bairdi and Grammonota Pallipes (Araneae: Linyphiidae)," The Great Lakes Entomologist, vol 47 (1) Available at: https://scholar.valpo.edu/tgle/vol47/iss1/3 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 20 THE GREAT LAKES ENTOMOLOGIST Vol. 47, Nos. 1 - 21Plant Science and Conservation, Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe, Illinois, 60022. 2Department of Natural and Applied Sciences, University of Wisconsin-Green Bay, 2420 Nicolet Drive, Green Bay, Wisconsin 54311. *Corresponding author: (e-mail: jsteffen@chicagobotanic.org).Disjunct Lake Michigan Populations of Two Atlantic Coast  Spiders, Disembolus bairdi and Grammonota pallipes (Araneae: Linyphiidae)  James F. Steffen1* and Michael L. Draney2Abstract Two species of linyphiid spiders, Disembolus bairdi Edwards, 1999 and Grammonota pallipes Banks, 1895, were discovered along the southwestern coast of Lake Michigan in Lake County, Illinois representing an Atlantic Coastal Plain disjunct distribution.   A brief discussion of known collection sites, habitat preferences, and possible modes of dispersal are given.____________________Disembolus bairdi Edwards, 1999 is a small erigonine spider species that was described by Edwards (1999) from the coastal marshes of Cape Cod, Mas-sachusetts and Long Island, New York from material collected in 1985 through 1990 in Massachusetts and in 1995 in New York.  The species was found under mats of organic debris, usually Zostera or Spartina, near the highest tide levels in areas protected from direct ocean influence.  To our knowledge, it has not been reported from other locations since the publication of Edwards (1999) (type specimens deposited in United States National Museum, Washington D. C.; Mu-seum of Comparative Zoology, Cambridge, Massachusetts; American Museum of Natural History, New York; and British Museum of Natural History, London). Grammonota pallipes Banks, 1895 is another erigonine spider that has been recorded from Long Island, New York (Banks 1895), Cape Cod, Massachusetts (Edwards 1993), Washington County, Maine (Jennings and Graham 2007), May’s Landing, New Jersey (Bishop and Crosby 1932), Somersworth, New Hampshire (Canadian National Collection of Insects, Arachnids and Nematodes, K. W. Wu, pers. comm.), as well as several locations in Canada (unknown Nova Scotia record reported in Buckle et al. 2001; Quebec records reported in Bélanger and Hutchinson 1992; and Ontario, New Brunswick, and Quebec records in Cana-dian National Collection of Insects, Arachnids and Nematodes, K. W. Wu, pers. comm.).  To our knowledge, these species have not previously been recorded for any of the states bordering Lake Michigan (Sierwald et al. 2005).  In late April of 2005, pitfall traps were established in the dunes and inter-dunal swales (panne) at Waukegan Beach along the west coast of Lake Michigan for the purpose of surveying the spider communities there.  The beach area is located approximately 0.8 km (0.5 miles) north of the harbor in Waukegan, Lake County, Illinois (42° 22’ 21” N, 87° 48’ 47” W).  The interdunal swales are flat, damp depressions located behind and between the foredune and backdunes. The substrate is damp sand with a sparse cover of Salix leaf litter.  The veg-etation, which was patchily distributed, consisted mostly of Salix myricoides, 1Steffen and Draney: Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders,Published by ValpoScholar, 20142014 THE GREAT LAKES ENTOMOLOGIST 21Schizachyrium scoparium, and Juncus balticus.  The dunes surrounding these swales were covered by several grass species, including Ammophila breviligu-lata, Calamovilfa longifolia, and Schizachyrium scoparium.  Several species of spiders from a number of families were sampled (see complete list in Table 1).  One male of an unidentified species of Linyphiidae was trapped along with six seemingly conspecific females on 1 May 2005.  The male was tentatively identified as Disembolus bairdi by the second author, and the identities of the male and female specimens were later confirmed by Robert L. Edwards (pers. comm. to first author).  In an attempt to obtain more individuals of the species, litter collections were taken at the site.  Interdunal litter samples from 5–9 May yielded 16 mature females, but no further males.  The spider appeared to be restricted to the interdunal swales, as no individuals were obtained from litter collections on the dunes.  The first author returned to the area on 27–28 December 2005 at which time 6 mature male and 35 mature female D. bairdi were found in the litter, and one female of Grammonota pallipes was found in a back-dune area of the beach.Litter samples also were taken approximately 11 kilometers north of Waukegan Beach at the northern unit of Illinois Beach State Park in Lake County during similar seasons in subsequent years (7 and 30 August and 17 November 2008 and 27 March 2009) to see if D. bairdi was present in similar interdunal habitats.  No individuals of this species were obtained from those samples.  However, six individuals of G. pallipes were collected at this site (one male on 30 August 2008 and five females on 17 November 2008).  A more recent litter sample (19 October 2012) from the southern unit of Illinois Beach State Park near the camp ground (42° 26’ 00.43” N 87° 48’ 15.55” W) produced one male and three females of D. bairdi in a black oak savanna close to the beach on a back-dune, approximately 55 meters from the water’s edge.  This location is approximately 7.2 kilometers north of Waukegan Beach.  One male and one female specimen of D. bairdi have been deposited in the collection of the Field Museum (FMNH) in Chicago, Illinois.  Two males and seven females are in the second author’s collection at the University of Wisconsin Green-Bay, Green Bay, Wisconsin, and two males and 42 females are in the first author’s collection at the Chicago Botanic Garden, Glencoe, IL. Table 1.  Ground-dwelling spiders of Waukegan Beach, Waukegan, Lake Co., IL.  All species were either trapped in pitfall traps or sorted from litter between 1 May 2005 and 28 December 2005.Family SpeciesClubionidae Clubiona abboti L. Koch, 1866Dictynidae Phantyna bicornis (Emerton, 1915)Linyphiidae Agyneta fabra (Keyserling, 1886) Disembolus bairdi Edwards, 1999 Eridantes erigonoides (Emerton, 1882) Grammonota pallipes Banks, 1895 Collinsia plumosus (Emerton, 1884) Idionella formosa (Banks, 1892) Islandiana flaveola (Banks, 1892)Salticidae Marpissa pikei (Peckham & Peckham, 1888) Neon nelli Peckham & Peckham, 18882The Great Lakes Entomologist, Vol. 47, No. 1 [2014], Art. 3https://scholar.valpo.edu/tgle/vol47/iss1/322 THE GREAT LAKES ENTOMOLOGIST Vol. 47, Nos. 1 - 2One male and one female also were sent to Robert L. Edwards at Woods Hole, MA.  Two females of G. pallipes have been deposited at the Field Museum in Chicago while the remainder of the material is held by the authors.  DiscussionFor D. bairdi, the Waukegan Beach site represents a range extension of approximately 1,287 km (800 miles) from the New York and Massachusetts collection localities of Edwards (1999).  The known habitat distribution and collection records suggest that D. bairdi is a shoreline species.  All known re-cords (Edwards 1999 and the present report) are from littoral wrack, and no individuals have been found inland, although litter samples were collected in higher elevation areas at the Waukegan Beach site, approximately 260 meters from the water’s edge.  Although G. pallipes appears to have a broader distribution than D. bairdi, the Illinois records for this species appear to be a westward range extension of approximately 850 km (528 miles) from the previous western collection site of Chatterton, Ontario.  Looking at all the known records, the species still appears to be restricted to the Atlantic Coastal Plain and the Great Lakes/St. Lawrence Seaway region of the United States and Canada.  Several specimens from the Canadian National Collection were collected within 10 km of the Ottawa River in Kingsmere, Quebec and Nepean and Torbolton, Ontario; other specimens were found within 20 km of Lake Ontario in Chatterton, Ontario (K. W. Wu, pers. comm.).  The remainder of the known G. pallipes records are from within 25 km of the Atlantic coast from New Brunswick to New Jersey, with many specimens recorded from coastal Kouchibouguac National Park, New Brunswick. G. pallipes appears to be less confined to beach environments than D. bairdi. In addition to the inland records mentioned above, it has been collected 5 km inland in Maine and 20 km inland in New Jersey and approximately 23 km in-land in Somersworth, New Hampshire (see records cited above).  The available habitat information is consistent with coastal (and similar) microhabitats, with specimens having been collected from sand dunes, sandy beach, dunes, forest in beach heather (Hudsonia tomentosa) litter and under debris, and ponds, which probably means near the shoreline (Bélanger and Hutchinson 1992; K. W. Wu, CNC records, pers. comm.).  Peattie (1922) describes Atlantic Coastal Plain disjunct plant communities in the Great Lakes region, some of which are concentrated in the southern end of Lake Michigan.  Reznicek (1994) gives a detailed discussion of 62 species of Atlantic Coastal plant species known to be disjunct in the Great Lakes region with the highest concentrations occurring in southwest Michigan, northern Indiana, and central and northwestern Wisconsin.  McLaughlin (1932) provides a thor-ough discussion of several possible postglacial avenues of dispersal, including the Mississippi River basin, the Mohawk-Hudson outlet, and the Ottawa connective, which resulted in more Atlantic Coastal Plain plant species being found in the region of the southern end of Lake Michigan than any other inland location.  The prevailing hypothesis is that these plant species gradually migrated along shore-lines of postglacial lakes and drainages.  Reznicek (1994) suggests that, although random long distance dispersal is unlikely, dispersal jumps of varying distance occurred between areas of suitable habitat created along postglacial drainages after which they became established in areas with extensive drying shorelines. While the plant component of these communities has been described, the animals accompanying them are less well known (Thomas 1951).  Thomas (1951) mentions two grasshopper species that presumably followed the sand and gravel outwash plains during post-Wisconsin times, which have survived in sandy areas around the Great Lakes.  Shapiro (1970) details the biogeography of several Atlantic Coastal Plain disjunct butterfly species.  He suggests the post-Wisconsin Hudson-Mohawk outlet of Lake Ontario (12,500–10,500 B.P.), a corridor connecting the 3Steffen and Draney: Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders,Published by ValpoScholar, 20142014 THE GREAT LAKES ENTOMOLOGIST 23Great Lakes and the Coastal Plain, as a means of dispersal for these organisms. He also suggests that the sedge-feeding marsh-living ecology of those insects was important in that dispersal, a pattern that is rare in butterflies with different non-sedge-feeding, non-marsh-living ecologies.  In spiders, ballooning is known as a common means of long distance dispersal (Duffey 1998).  However, Bonte et al. (2003) have shown that ballooning is negatively related to habitat specialization in spiders and also that habitat specialists are characterized by poorly developed dispersal behavior.  Hawes (2007) suggests that high arctic Linyphiidae may have originally colonized those habitats by ballooning, and then ballooning behavior atrophied over evolutionary time.  Therefore, although D. bairdi is currently an apparent habitat specialist, it may have initially dispersed by means of bal-looning during a time of more or less continuous postglacial beach and shoreline habitat availability (McLaughlin 1932) and then later lost ballooning behavior as habitat became more restricted.  The degree of habitat specialization for G. pallipes is less clear, but the species does seem to be present only within a few kilometers of the Atlantic Coast, Great Lakes, or Saint Lawrence/Ottawa River systems, which also suggests fairly specific microhabitat requirements and lack of long-distance ballooning ability.  Disembolus bairdi may be the first example of a true Atlantic Coastal Plain disjunct spider in the Great Lakes region.  Although G. pallipes may also be an Atlantic Coastal Plain disjunct species, its status is less clear because of having a wider distribution, being found at least several kilometers away from littoral environments.  Two wolf spider (Lycosidae) species, Arctosa littoralis (Hentz, 1844) and Pardosa groenlandica (Thorell, 1872) also are found in both geographical locations, but these species seem to be more widespread and less restricted to specific microhabitats.  Although most populations of these spe-cies have been recorded from sandy (in the case of A. littoralis) or cobbly/pebbly (in the case of P. groenlandica) seashore and lakeshore habitats, A. littoralis also has been found in similar sandy habitats on riverbanks and inland, and P. groenlandica is found in alpine scree above timberline across Canada and in the mountain west (Dondale and Redner 1983; Dondale 1999).  Only D. bairdi seems to be truly restricted to littoral microhabitats and may represent an ideal model organism to trace range expansion after the last glaciation.  Only a small segment of Lake Michigan shoreline was examined in this study.  Therefore, it would be important to make a wider search of these littoral areas throughout the Great Lakes region, and in particular, those areas of high concentrations of disjunct plant species, to further document the distribution of D. bairdi and G. pallipes and search for other possible disjunct species restricted to the same microhabitat.  Making these specimens available as identified and digitized museum collections will facilitate researchers in distinguishing habitat specialists and tracing their distributional history and possible mode of dispersal based on known geological events and timelines.  AcknowledgmentsWe are grateful to the late Robert Edwards for kindly agreeing to ex-amine specimens and encouragement in our studies of linyphiids.  We also thank Don Buckle (Saskatoon, Saskatchewan) for helping us find information about Grammonota pallipes and King Wan Wu, Valerie Behan-Pelletier, and Owen Lonsdale of the Canadian National Collection of Insects, Arachnids, and Nematodes (Ottawa, Ontario) for making specimen information available to us. Literature CitedBanks, N.  1895.  A list of spiders of Long Island, N. Y., with descriptions of new species. Journal of the New York Entomological Society 3: 76–93.  4The Great Lakes Entomologist, Vol. 47, No. 1 [2014], Art. 3https://scholar.valpo.edu/tgle/vol47/iss1/324 THE GREAT LAKES ENTOMOLOGIST Vol. 47, Nos. 1 - 2Bélanger, G., and R. Hutchinson.  1992.  Liste annotée des araignées (Araneae) du Quebéc.  Pirata 1: 2–119.  Bishop, S. C., and C. R. Crosby.  1932.  Studies in American spiders: the genus Gram-monota.  Journal of the New York Entomological Society 40: 393–421.  Bonte, G., N. Vandenbroecke, L. Lens, and J. Maelfait.  2003.  Low propensity for aerial dispersal in specialist spiders from fragmented landscapes.   Proceedings Royal Society of London B 270: 1601–1607.  Buckle, D. J., D. Carroll, R. L. Crawford, and V. D. Roth.  2001.  Linyphiidae and Pimoidae of America north of Mexico: Checklist, synonymy, and literature. Part 2, pp. 89–191.  In P. Paquin, and D. J. Buckle (eds.), Contributions à la connaissance des Araignées (Araneae) ďAmérique du Nord.  Fabreries Supplément 10.  Dondale, C. D.  1999.  Revision of the Groenlandica subgroup of the genus Pardosa (Araneae: Lycosidae).  Journal of Arachnology 27: 435–448.  Dondale, C. D., and J. H. Redner.  1983.  Revision of the wolf spiders of the genus Arctosa C. L. Koch in North and Central America (Araneae: Lycosidae).  Journal of Arachnology 11: 1–30.  Duffey, E.  1998.  Aerial dispersal in spiders, pp. 187–191.  In P. A. Selden (ed.), Pro-ceedings of the 17th European Colloquium of Arachnology, Edinburgh 1997.  British Arachnological Society, Burnham Beeches, Bucks.  X + 350 pp.  Edwards, R. L.  1993.  New records of spiders (Araneae) from Cape Cod, Massachusetts, including two possible European immigrants.  Entomological News 104: 79–82.  Edwards, R. L.  1999.  A new Disembolus (Araneae, Linyphiidae) from Cape Cod, Mas-sachusetts and Long Island, New York.  Journal of Arachnology 27: 16–18.  Hawes, T. C.  2007.  Ballooning in high Arctic linyphiids: a case of behavioral atrophy? Bulletin of the British Arachnological Society 14: 39–42.  Hentz, N. M.  1844.  Description and figures of the araneides of the United States.  Boston Journal of Natural History 4: 386–396.  Jennings, D. T., and F. Graham, Jr.  2007.  Spiders (Arachnida: Aranae) of Milbridge, Washington County, Maine.  General Technical Report NRS-16 Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station.  204 pp. McLaughlin, W. T.  1932.  Atlantic coastal plain plants in the sand barrens of north-western Wisconsin.  Ecological Monographs 2: 335–383.  Peattie, D. C.  1922.  The Atlantic coastal plain element in the flora of the Great Lakes. Rhodora 24: 57–70, 80–88.  Reznicek, A. A.  1994.  The disjunct coastal plain flora in the Great Lakes region.  Bio-logical Conservation 68: 203–215.  Shapiro, A. M.  1970.  Postglacial biogeography and the distribution of Poanes viator (Hesperiidae) and other marsh butterflies.  Journal of Research on the Lepidoptera 9: 125–155.  Sierwald, P., M. L. Draney, T. Prentice, F. Pascoe, N. Sandin, E. M. Lehman, V. Medland, and J. Louderman.  2005.  The spider species of the Great Lakes States. Proceedings of the Indiana Academy of Science 114: 111–206.  Thomas, E. S.  1951.  Distribution of Ohio animals.  The Ohio Journal of Science 51: 153–167.  Thorell, T.  1872.  Om några Arachniden från Grönland.  Öfversigt af Kongliga Veten-skaps-Akademiens Förhandlingar 29: 147–166. 5Steffen and Draney: Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders,Published by ValpoScholar, 2014

Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders, \u3ci\u3eDisembolus Bairdi\u3c/i\u3e and \u3ci\u3eGrammonota Pallipes\u3c/i\u3e (Araneae: Linyphiidae)

ValpoScholar

The Great Lakes Entomologist 
Volume 47 
Numbers 1 & 2 - Spring/Summer 2014 Numbers 
1 & 2 - Spring/Summer 2014 
Article 3 
April 2014 
Disjunct Lake Michigan Populations of Two Atlantic Coast 
Spiders, Disembolus Bairdi and Grammonota Pallipes (Araneae: 
Linyphiidae) 
James F. Steffen 
Chicago Botanic Garden 
Michael L. Draney 
University of Wisconsin 
Follow this and additional works at: https://scholar.valpo.edu/tgle 
 Part of the Entomology Commons 
Recommended Citation 
Steffen, James F. and Draney, Michael L. 2014. "Disjunct Lake Michigan Populations of Two Atlantic Coast 
Spiders, Disembolus Bairdi and Grammonota Pallipes (Araneae: Linyphiidae)," The Great Lakes 
Entomologist, vol 47 (1) 
Available at: https://scholar.valpo.edu/tgle/vol47/iss1/3 
This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. 
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20 THE GREAT LAKES ENTOMOLOGIST Vol. 47, Nos. 1 - 2
1Plant Science and Conservation, Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe, 
Illinois, 60022. 
2Department of Natural and Applied Sciences, University of Wisconsin-Green Bay, 
2420 Nicolet Drive, Green Bay, Wisconsin 54311. 
*Corresponding author: (e-mail: jsteffen@chicagobotanic.org).
Disjunct Lake Michigan Populations of Two Atlantic Coast  
Spiders, Disembolus bairdi and Grammonota pallipes 
(Araneae: Linyphiidae)  
James F. Steffen1* and Michael L. Draney2
Abstract
 Two species of linyphiid spiders, Disembolus bairdi Edwards, 1999 and 
Grammonota pallipes Banks, 1895, were discovered along the southwestern 
coast of Lake Michigan in Lake County, Illinois representing an Atlantic Coastal 
Plain disjunct distribution.   A brief discussion of known collection sites, habitat 
preferences, and possible modes of dispersal are given.
____________________
Disembolus bairdi Edwards, 1999 is a small erigonine spider species that 
was described by Edwards (1999) from the coastal marshes of Cape Cod, Mas-
sachusetts and Long Island, New York from material collected in 1985 through 
1990 in Massachusetts and in 1995 in New York.  The species was found under 
mats of organic debris, usually Zostera or Spartina, near the highest tide levels 
in areas protected from direct ocean influence.  To our knowledge, it has not 
been reported from other locations since the publication of Edwards (1999) (type 
specimens deposited in United States National Museum, Washington D. C.; Mu-
seum of Comparative Zoology, Cambridge, Massachusetts; American Museum of 
Natural History, New York; and British Museum of Natural History, London). 
Grammonota pallipes Banks, 1895 is another erigonine spider that has been 
recorded from Long Island, New York (Banks 1895), Cape Cod, Massachusetts 
(Edwards 1993), Washington County, Maine (Jennings and Graham 2007), May’s 
Landing, New Jersey (Bishop and Crosby 1932), Somersworth, New Hampshire 
(Canadian National Collection of Insects, Arachnids and Nematodes, K. W. Wu, 
pers. comm.), as well as several locations in Canada (unknown Nova Scotia 
record reported in Buckle et al. 2001; Quebec records reported in Bélanger and 
Hutchinson 1992; and Ontario, New Brunswick, and Quebec records in Cana-
dian National Collection of Insects, Arachnids and Nematodes, K. W. Wu, pers. 
comm.).  To our knowledge, these species have not previously been recorded for 
any of the states bordering Lake Michigan (Sierwald et al. 2005).  
In late April of 2005, pitfall traps were established in the dunes and inter-
dunal swales (panne) at Waukegan Beach along the west coast of Lake Michigan 
for the purpose of surveying the spider communities there.  The beach area is 
located approximately 0.8 km (0.5 miles) north of the harbor in Waukegan, Lake 
County, Illinois (42° 22’ 21” N, 87° 48’ 47” W).  The interdunal swales are flat, 
damp depressions located behind and between the foredune and backdunes. 
The substrate is damp sand with a sparse cover of Salix leaf litter.  The veg-
etation, which was patchily distributed, consisted mostly of Salix myricoides, 
1
Steffen and Draney: Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders,
Published by ValpoScholar, 2014
2014 THE GREAT LAKES ENTOMOLOGIST 21
Schizachyrium scoparium, and Juncus balticus.  The dunes surrounding these 
swales were covered by several grass species, including Ammophila breviligu-
lata, Calamovilfa longifolia, and Schizachyrium scoparium.  Several species 
of spiders from a number of families were sampled (see complete list in Table 
1).  One male of an unidentified species of Linyphiidae was trapped along with 
six seemingly conspecific females on 1 May 2005.  The male was tentatively 
identified as Disembolus bairdi by the second author, and the identities of the 
male and female specimens were later confirmed by Robert L. Edwards (pers. 
comm. to first author).  In an attempt to obtain more individuals of the species, 
litter collections were taken at the site.  Interdunal litter samples from 5–9 
May yielded 16 mature females, but no further males.  The spider appeared 
to be restricted to the interdunal swales, as no individuals were obtained from 
litter collections on the dunes.  The first author returned to the area on 27–28 
December 2005 at which time 6 mature male and 35 mature female D. bairdi 
were found in the litter, and one female of Grammonota pallipes was found in 
a back-dune area of the beach.
Litter samples also were taken approximately 11 kilometers north of 
Waukegan Beach at the northern unit of Illinois Beach State Park in Lake 
County during similar seasons in subsequent years (7 and 30 August and 17 
November 2008 and 27 March 2009) to see if D. bairdi was present in similar 
interdunal habitats.  No individuals of this species were obtained from those 
samples.  However, six individuals of G. pallipes were collected at this site (one 
male on 30 August 2008 and five females on 17 November 2008).  A more recent 
litter sample (19 October 2012) from the southern unit of Illinois Beach State 
Park near the camp ground (42° 26’ 00.43” N 87° 48’ 15.55” W) produced one 
male and three females of D. bairdi in a black oak savanna close to the beach 
on a back-dune, approximately 55 meters from the water’s edge.  This location 
is approximately 7.2 kilometers north of Waukegan Beach.  
One male and one female specimen of D. bairdi have been deposited in 
the collection of the Field Museum (FMNH) in Chicago, Illinois.  Two males 
and seven females are in the second author’s collection at the University of 
Wisconsin Green-Bay, Green Bay, Wisconsin, and two males and 42 females 
are in the first author’s collection at the Chicago Botanic Garden, Glencoe, IL. 
Table 1.  Ground-dwelling spiders of Waukegan Beach, Waukegan, Lake Co., IL.  All 
species were either trapped in pitfall traps or sorted from litter between 1 May 2005 
and 28 December 2005.
Family Species
Clubionidae Clubiona abboti L. Koch, 1866
Dictynidae Phantyna bicornis (Emerton, 1915)
Linyphiidae Agyneta fabra (Keyserling, 1886)
 Disembolus bairdi Edwards, 1999
 Eridantes erigonoides (Emerton, 1882)
 Grammonota pallipes Banks, 1895
 Collinsia plumosus (Emerton, 1884)
 Idionella formosa (Banks, 1892)
 Islandiana flaveola (Banks, 1892)
Salticidae Marpissa pikei (Peckham & Peckham, 1888)
 Neon nelli Peckham & Peckham, 1888
2
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22 THE GREAT LAKES ENTOMOLOGIST Vol. 47, Nos. 1 - 2
One male and one female also were sent to Robert L. Edwards at Woods Hole, 
MA.  Two females of G. pallipes have been deposited at the Field Museum in 
Chicago while the remainder of the material is held by the authors.  
Discussion
For D. bairdi, the Waukegan Beach site represents a range extension of 
approximately 1,287 km (800 miles) from the New York and Massachusetts 
collection localities of Edwards (1999).  The known habitat distribution and 
collection records suggest that D. bairdi is a shoreline species.  All known re-
cords (Edwards 1999 and the present report) are from littoral wrack, and no 
individuals have been found inland, although litter samples were collected in 
higher elevation areas at the Waukegan Beach site, approximately 260 meters 
from the water’s edge.  
Although G. pallipes appears to have a broader distribution than D. bairdi, 
the Illinois records for this species appear to be a westward range extension of 
approximately 850 km (528 miles) from the previous western collection site of 
Chatterton, Ontario.  Looking at all the known records, the species still appears 
to be restricted to the Atlantic Coastal Plain and the Great Lakes/St. Lawrence 
Seaway region of the United States and Canada.  Several specimens from the 
Canadian National Collection were collected within 10 km of the Ottawa River 
in Kingsmere, Quebec and Nepean and Torbolton, Ontario; other specimens 
were found within 20 km of Lake Ontario in Chatterton, Ontario (K. W. Wu, 
pers. comm.).  The remainder of the known G. pallipes records are from within 
25 km of the Atlantic coast from New Brunswick to New Jersey, with many 
specimens recorded from coastal Kouchibouguac National Park, New Brunswick. 
G. pallipes appears to be less confined to beach environments than D. bairdi. 
In addition to the inland records mentioned above, it has been collected 5 km 
inland in Maine and 20 km inland in New Jersey and approximately 23 km in-
land in Somersworth, New Hampshire (see records cited above).  The available 
habitat information is consistent with coastal (and similar) microhabitats, with 
specimens having been collected from sand dunes, sandy beach, dunes, forest in 
beach heather (Hudsonia tomentosa) litter and under debris, and ponds, which 
probably means near the shoreline (Bélanger and Hutchinson 1992; K. W. Wu, 
CNC records, pers. comm.).  
Peattie (1922) describes Atlantic Coastal Plain disjunct plant communities 
in the Great Lakes region, some of which are concentrated in the southern end 
of Lake Michigan.  Reznicek (1994) gives a detailed discussion of 62 species of 
Atlantic Coastal plant species known to be disjunct in the Great Lakes region with 
the highest concentrations occurring in southwest Michigan, northern Indiana, 
and central and northwestern Wisconsin.  McLaughlin (1932) provides a thor-
ough discussion of several possible postglacial avenues of dispersal, including the 
Mississippi River basin, the Mohawk-Hudson outlet, and the Ottawa connective, 
which resulted in more Atlantic Coastal Plain plant species being found in the 
region of the southern end of Lake Michigan than any other inland location.  The 
prevailing hypothesis is that these plant species gradually migrated along shore-
lines of postglacial lakes and drainages.  Reznicek (1994) suggests that, although 
random long distance dispersal is unlikely, dispersal jumps of varying distance 
occurred between areas of suitable habitat created along postglacial drainages 
after which they became established in areas with extensive drying shorelines. 
While the plant component of these communities has been described, the animals 
accompanying them are less well known (Thomas 1951).  Thomas (1951) mentions 
two grasshopper species that presumably followed the sand and gravel outwash 
plains during post-Wisconsin times, which have survived in sandy areas around 
the Great Lakes.  Shapiro (1970) details the biogeography of several Atlantic 
Coastal Plain disjunct butterfly species.  He suggests the post-Wisconsin Hudson-
Mohawk outlet of Lake Ontario (12,500–10,500 B.P.), a corridor connecting the 
3
Steffen and Draney: Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders,
Published by ValpoScholar, 2014
2014 THE GREAT LAKES ENTOMOLOGIST 23
Great Lakes and the Coastal Plain, as a means of dispersal for these organisms. 
He also suggests that the sedge-feeding marsh-living ecology of those insects was 
important in that dispersal, a pattern that is rare in butterflies with different 
non-sedge-feeding, non-marsh-living ecologies.  In spiders, ballooning is known as 
a common means of long distance dispersal (Duffey 1998).  However, Bonte et al. 
(2003) have shown that ballooning is negatively related to habitat specialization 
in spiders and also that habitat specialists are characterized by poorly developed 
dispersal behavior.  Hawes (2007) suggests that high arctic Linyphiidae may have 
originally colonized those habitats by ballooning, and then ballooning behavior 
atrophied over evolutionary time.  Therefore, although D. bairdi is currently 
an apparent habitat specialist, it may have initially dispersed by means of bal-
looning during a time of more or less continuous postglacial beach and shoreline 
habitat availability (McLaughlin 1932) and then later lost ballooning behavior 
as habitat became more restricted.  The degree of habitat specialization for G. 
pallipes is less clear, but the species does seem to be present only within a few 
kilometers of the Atlantic Coast, Great Lakes, or Saint Lawrence/Ottawa River 
systems, which also suggests fairly specific microhabitat requirements and lack 
of long-distance ballooning ability.  
Disembolus bairdi may be the first example of a true Atlantic Coastal 
Plain disjunct spider in the Great Lakes region.  Although G. pallipes may also 
be an Atlantic Coastal Plain disjunct species, its status is less clear because of 
having a wider distribution, being found at least several kilometers away from 
littoral environments.  Two wolf spider (Lycosidae) species, Arctosa littoralis 
(Hentz, 1844) and Pardosa groenlandica (Thorell, 1872) also are found in both 
geographical locations, but these species seem to be more widespread and less 
restricted to specific microhabitats.  Although most populations of these spe-
cies have been recorded from sandy (in the case of A. littoralis) or cobbly/pebbly 
(in the case of P. groenlandica) seashore and lakeshore habitats, A. littoralis 
also has been found in similar sandy habitats on riverbanks and inland, and 
P. groenlandica is found in alpine scree above timberline across Canada and in 
the mountain west (Dondale and Redner 1983; Dondale 1999).  Only D. bairdi 
seems to be truly restricted to littoral microhabitats and may represent an ideal 
model organism to trace range expansion after the last glaciation.  
Only a small segment of Lake Michigan shoreline was examined in this 
study.  Therefore, it would be important to make a wider search of these littoral 
areas throughout the Great Lakes region, and in particular, those areas of high 
concentrations of disjunct plant species, to further document the distribution of 
D. bairdi and G. pallipes and search for other possible disjunct species restricted 
to the same microhabitat.  Making these specimens available as identified and 
digitized museum collections will facilitate researchers in distinguishing habitat 
specialists and tracing their distributional history and possible mode of dispersal 
based on known geological events and timelines.  
Acknowledgments
We are grateful to the late Robert Edwards for kindly agreeing to ex-
amine specimens and encouragement in our studies of linyphiids.  We also 
thank Don Buckle (Saskatoon, Saskatchewan) for helping us find information 
about Grammonota pallipes and King Wan Wu, Valerie Behan-Pelletier, and 
Owen Lonsdale of the Canadian National Collection of Insects, Arachnids, and 
Nematodes (Ottawa, Ontario) for making specimen information available to us. 
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24 THE GREAT LAKES ENTOMOLOGIST Vol. 47, Nos. 1 - 2
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Steffen and Draney: Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders,
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Disjunct Lake Michigan Populations of Two Atlantic Coast Spiders, \u3ci\u3eDisembolus Bairdi\u3c/i\u3e and \u3ci\u3eGrammonota Pallipes\u3c/i\u3e (Araneae: Linyphiidae)

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