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40
2003. The Journal of Arachnology 31:40–43
ANYPHAENA (ARANEAE, ANYPHAENIDAE) OVERWINTERING
ON LOWEST LIMBS OF WHITE OAK
David W. Boyd, Jr.: USDA, ARS, Small Fruit Research Station, Poplarville,
MS 39470 USA
Will K. Reeves: Department of Entomology, Clemson University, Clemson,
SC 29634 USA
ABSTRACT. Juvenile Anyphaena sp. were collected from overwintering traps placed on the lowest
limbs of white oak, Quercus alba, in South Carolina. Multiple regression analysis was used to determine
that the number of juvenile Anyphaena sp. found can be predicted by the circumference of the limb, the
distance from the trunk and the distance from the ground. This study helps demonstrate that the limbs of
trees, although often neglected in overwintering studies, can provide a refuge for arthropods.
Keywords: Anyphaena, overwintering, Quercus alba
Many spiders enter a dormant stage during
winter conditions (Schaefer 1977) and those
that overwinter on the trunks of trees are often
surveyed by collecting the spiders with card-
board wrapped around tree trunks (Tamaki &
Halfhill 1968; Tedders 1974; Fye 1985;Mizell
& Schiffhauer 1987; Pekar 1999; Horton et al.
2001). However, the species collected in the
trunk traps are not necessarily the same spe-
cies that are collected during warmer months
from the limbs of the same trees (Peka´r 1999;
Horton et al. 2001) and the limbs are usually
neglected when sampling for overwintering
species. Our research was conducted to deter-
mine if the limbs of white oak trees, Quercus
alba L., were suitable for arthropods to over-
winter, and, if so, where on the limbs they
overwintered.
METHODS
We made traps of gray coroplast (corrugat-
ed plastic, similar to cardboard) by cutting a
sheet of coroplast into sections 15 cm long by
3–3.5 cm wide, providing six longitudinal
cavities in each trap. We placed traps on three
mature white oak trees, Quercus alba L., on
30 October 1998. One tree was located in
Pickens County, South Carolina on the Clem-
son University campus. Two trees were locat-
ed in Greenville County, South Carolina, one
on the Bob Jones University campus and the
other at Reedy River Falls Historical Park.
Trees were selected based on ease of acces-
sibility. Three sets of traps were placed on
limbs greater than or equal to3minlength:
one trap set was proximate to the trunk, one
was in the middle of the limb, and one was
on the terminus of the limb. Two sets of traps
were placed on limbs shorter than 3 m in
length: one trap set was proximate to the trunk
and one on the terminus of the limb. We
placed traps around the limb 2.5 cm apart,
parallel to the limb, and held them in place
with gray duct tape. The diameter of the limb
determined the number of traps around the
limb. Two groups of traps were place around
the limb 3–6 cm apart, one offsetting the other
(Fig. 1).
We used 5 limbs on the oak tree in Pickens
County, each with 3 sets of traps. On the tree
at Reedy River Historical Park in Greenville
County we used three limbs, each with 2 sets
of traps, and on the tree at Bob Jones Uni-
versity we used two limbs, each with 3 sets
of traps. We used a total of 27 traps. The num-
ber of limbs used was based on the number
of limbs reachable at each location with a 3
m ladder. For purposes of regression analysis
the average circumference of the limb at each
trap (circumference at both ends of the trap
set divided by 2), the distance of the trap from
trunk, the distance from the trap to the ground,
and the branching of the limb from the trunk
to the trap were measured. The bark surface
was rated on a scale of 1–3, where 1
5
smooth and 3
5
rough.
41BOYD, Jr. & REEVES—ANYPHAENA OVERWINTERING ON WHITE OAK
Figure 1.—One set of coroplast strip traps.
Traps were removed 24 February 1999 after
three consecutive days of average daily tem-
peratures near freezing (
6
1
8
C) (average tem-
perature for Greenville County was 0.9
8
C for
22–24 Feb. and
2
0.4
8
C for Pickens County).
Traps were placed in plastic bags, taken to the
lab and placed in the freezer. Specimens were
removed from the traps, separated, preserved
in 80% ethanol and identified. Voucher spec-
imens were placed in the Clemson University
Arthropod Collection.
Juvenile Anyphaena sp. were the only ar-
thropods found in numbers large enough to
conduct multiple regression analysis. The total
number of Anyphaena sp. collected was 340.
Multiple regression analysis was conducted
using Minitab. The dependent variable was
the number of Anyphaena, which was stan-
dardized for each trap set by dividing the total
number of Anyphaena by the total number of
traps in each set. The independent variables
were average circumference of the limb, dis-
tance from the trunk, distance from the
ground, number of branches per limb, and
bark surface scale for each trap set.
For multiple regression analysis on Any-
phaena no transformation of the dependent
variable was needed. A tolerance test showed
multicolinearity between polynomials of the
independent variables and the independent
variables. Therefore, only the raw independent
variables were used in the analysis. Stepwise,
forward, and backward model selection tech-
inques all provided the same model. The mod-
el showed no systematic patterns, no outliers,
and no evidence of lack of fit.
RESULTS
Spiders were the most numerous arthropods
collected. All the arthropods collected are list-
ed in Table 1. More Anyphaena sp. were col-
lected near the trunk than the terminus of the
limbs (Table 2).
The multiple regression analysis provide
the following model: Number of Anyphaena
sp.
52
16.1
1
20.3 (circumference of limb)
2
2.74 (distance from the trunk)
1
9.86 (dis-
tance from the ground). This model, with an
R
2
of 70.0%, shows that the number of Any-
phaena overwintering in traps on the bottom
limbs of Q. alba can be predicted by the cir-
cumference of the limb, the distance from the
trunk and the distance from the ground.
DISCUSSION
Schaefer (1977) studied the overwintering
habits of spiders and determined four over-
wintering habit types. Anyphaena sp. is part
42 THE JOURNAL OF ARACHNOLOGY
Table 1.—Arthropods collected from overwintering traps around limbs of white oak.
Class Order Family Species
Arachnida Araneae Agelenidae
Anyphaenidae
Araneidae
Philodromidae
Salticidae
Thomisidae
Coras sp. juv.
Anyphaena sp. juv.
Araneus sp. juv.
Philodromus vulgaris (Hentz)
Philodromus sp. juv.
Eris militaris (Hentz)
Hentzia mitrata (Hentz)
Metacyrba undata (De Geer)
Bassaniana versicolor (Keyserling)
Diplopoda
Insecta Polyxenida
Blattaria
Diptera
Hemiptera
Psocoptera
Polyxenidae
Blattellidae
Syrphidae
Miridae
Ectopsocidae
Polyxenus fasiculatus (Say)
Parcoblatta sp. juv.
Syrphus sp. juv.
Deraeocoris nebulosus (Uhler)
Ectopsocus meridionalis Ribaga
Table 2.—Mean number of Anyphaena sp. juv.
(
6
SE) collected from coroplast traps on white oak,
Quercus alba limbs (South Carolina, 1999). Limb
position is relative to the trunk.
Limb
position Average #
Anyphaena (n)
Proximate
Middle
Terminus
17.4
6
2.0 (10)
12.3
6
2.8 (10)
5.4
6
1.3 (8)
of the majority (45%) of spiders that over-
winter in the juvenile stage (Schaefer 1977).
Tree-dwelling spiders in the genus Anyphaena
are nocturnal wanderers, typically living in fo-
liage from spring through fall, but little of
their ecology or behavior is known (Platnick
1974). They feed on aphids and other prey not
typically active during the day (Marc & Ca-
nard 1997; Marc et al. 1999). Anyphaena spp.
take refuge during the winter but can be active
during warmer days (Turnbull 1960), increas-
ing their ability for survival (Gunnarsson
1985). Other overwintering studies, that in-
cluded Anyphaena spp., sampled only the
trunk or the proximal end of the largest
branch. Bajwa and AliNiazee (2001) found
only four Anyphaena in a four year study.
Horton et al. (2001) found only seven Any-
phaena in a one year study. We demonstrated
that Anyphaena will overwinter on most parts
of the branches with refugia present.
Most refuges available to overwintering
spiders are eliminated when leaves are shed.
Previous studies have shown or suggested that
after leaf-fall spiders move down from the
crown until they find refuge (Duffey 1969;
Horton et al. 2001), which might be the case
with our spiders. The overwintering traps pro-
vided a refuge that otherwise would not have
been available. The diameter of the limbs af-
fected the number of spiders and without ex-
ception, the larger the diameter of the branch
the rougher the bark, which might also pro-
vide refugia.
Horton et al. (2001) collected arthropods
from cardboard bands weekly 23 Aug–07 Dec
1999 in Washington apple and pear orchards.
They found Anyphaena pacifica Banks in
higher numbers (224 total) on a weekly basis
than in overwintering samples (7 total, col-
lected in Jan 2000). They suggested that the
spiders overwinter elsewhere. In Oregon, A.
pacifica was found in low numbers (0.35% of
total catch) during the growing season by
beating the branches over a net (Bajwa &
AliNiazee 2001). In Europe, Marc et al.
(1999) and Marc and Canard (1997) suggested
that A. accentuata (Walckenaer) overwinters
on the tree trunk, but they collected very few
individuals (1% of total catch).
Our study provides information that can be
used in further studies of overwintering ar-
thropods on trees. The branches represent a
large portion of the tree and are often neglect-
ed as a sampling site during the winter
months. Large numbers of spiders on the
limbs could alter decision made in integrated
pest management for landscapes and orchards
such as apple, peach, pear, and pecan.
43BOYD, Jr. & REEVES—ANYPHAENA OVERWINTERING ON WHITE OAK
ACKNOWLEDGMENTS
We thank J. Boyd for technical assistance
and for commenting on an earlier draft of the
manuscript. Thanks also to E. Mockford for
identifying psocids, F.C. Thompson (SEL,
ARS, USDA) for identifying syrphids, F.Coy-
le for identifying salticids, and E. Grissell
(SEL, ARS, USDA) for identifying the chal-
cid wasp. P. Adler, A. Wheeler (both of Clem-
son University), and J. Carroll (USDA, ARS)
made helpful suggestions to improve the man-
uscript, for which we are grateful. This is con-
tribution number 4715 of the South Carolina
Agriculture and Forestry Research System,
Clemson University.
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Manuscript received 1 November 2001, revised 15
October 2002.