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ANYPHAENA (ARANEAE, ANYPHAENIDAE) OVERWINTERING ON LOWEST LIMBS OF WHITE OAK

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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.
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.
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The life cycle of spiders in temperate latitudes (N. Germany) and the adaptations of the species to low winter temperatures were studied. 5 types of life cycle can be distinguished: I. Eurychronous species hibernating in different stages (23% of 277 spiders); II. Stenochronous species reproducing in spring and summer and hibernating as immatures (45 %); III. Stenochronous species reproducing in autumn and hibernating in the egg stage (7 %); IV. Diplochronous species overwintering mainly as adults and reproducing in spring (3 %); V. Winter‐active species (9 %). The developmental cycles are controlled by temperature or photoperiod and can be partly defined by certain mechanisms of dormancy, e. g. retardation of postembryogenesis evocated by short days in spiders of type II (postembryonic diapause), arrest of embryogenesis by high temperatures in type III (egg diapause), and inhibition of the maturation of ovaries induced by short days in type IV (reproductive diapause). Spiders or spider eggs could be supercooled, but were not freezing‐tolerant. They may be adapted to low winter temperatures by their spatial or seasonal distribution, metabolism, or resistance. 84 % of the spiders overwintered in leaf litter or in vegetation near the ground; 7 % remained without shelter in higher vegetation layers. Diapause was characterized by a reduction of oxygen consumption. The supercooling points of some hibernating stages were lowered in response to falling ambient temperatures. Only few spiders were endangered by a high rate of winter mortality. Different strategies of hibernation are realized by safe and endangered species. The advantage of low winter mortality in the former populations can be compensated by high mortality during postembryonic development. Zusammenfassung Zur Winterökologie von Spinnen (Araneida) Es wird eine Übersicht über den Jahreszyklus der Spinnen in den gemäßigten Breiten (Umgebung von Kiel, Schleswig‐Holstein) und die Anpassungen der Arten an die niedrigen Wintertemperaturen gegeben. Nach dem überwinternden Stadium lassen sich 5 Typen des Jahreszyklus unterscheiden: I. Eurychrone Arten, die in verschiedenen Stadien überwintern (23% von 277 Arten); II. Stenochrone Arten mit Fortpflanzungsperiode im Frühjahr und Sommer, die als Jungspinnen überwintern (45 %); III. Stenochrone Arten mit Fortpflanzungsperiode im Herbst, die als Ei überwintern (7%); IV. Diplochrone Arten, die meist als Adulte überwintern und sich im Frühjahr fortpflanzen (3 %); V. Winteraktive (9 %). Diese Entwicklungszyklen werden in unterschiedlicher Weise durch Temperatur oder Photoperiode kontrolliert und können zum Teil mit bestimmten Dormanzmechanismen definiert werden. Besondere Bedeutung hatten eine durch Kurztag bedingte Verlangsamung der Postembryogenese bei Spinnen des Typs II (postembryonale Diapause), Stopp der Embryogenese bei hohen Temperaturen bei Typ III (Eidiapause) und eine durch Kurztag induzierte Hemmung der Ovarienreifung bei Typ IV (reproduktive Diapause). Die Arten oder Entwicklungsstadien hatten eine unterschiedlich große Kälteresistenz (gemessen am Unterkühlungspunkt) und waren nicht gefrierresistent. Am tiefsten konnte das Eistadium unterkühlt werden (bis — 31° C). Die Spinnen sind an die Winterkälte unterschiedlich gut angepaßt. Dies betrifft die räumliche und zeitliche Verteilung, den Stoffwechsel und die Resistenz. 84 % der Arten überwinterten in Laubstreu und bodennaher Vegetation, 7 % frei in der höheren Vegetation. Diapause war mit reduziertem Sauerstoff‐verbrauch verknüpft. Der Unterkühlungspunkt einiger überwinternder Stadien nahm parallel zu sinkenden Umgebungstemperaturen ab. Die Wintermortalität der meisten Arten war im Vergleich zur Generationsmortalität gering. Es lassen sich ‐ durch den Winter ‐ gefährdete und ungefährdete Arten und damit zwei Strategien der Überwinterung unterscheiden. Bei letzterer kann der Vorteil geringer Wintermortalität durch höhere Mortalität während der postembryonalen Entwicklung kompensiert werden.
Article
Results from field experiments indicate that predation occurs among spruce-living spiders during winter in SW Sweden. Field observations of natural activity showed that Philodromus spp and Pityohyphantes phrygianus together make up 80% of the spiders active on spruce in winter. They are therefore potential predators on other overwintering spiders. Laboratory experiments were performed at +4 C to assess the importance of such predation between spiders. Small spiders (length 2.5 mm) which had a mean mortality of 3% only. Among the small spiders the Erigninae spp seemed to be more vulnerable to predation than other taxonomic groups. Predation also occurred when large P. phrygianus were kept together, but such predation caused mortality of less importance to the spider populations than the mortality among small spiders. Differences in spider density and food availability did not change this pattern. Considerable weight increase occurred in subadult P. phrygianus when fed during winter. This suggests that winter foraging specimens increase their fitness. Interspecific predation among spiders is suggested to be an important mortality factor in natural populations at high spider densities in November and December, when the ambient temperature often is above 0 C and when the density of large spiders is not yet substantially reduced by bird predation.
Article
In northern Europe at least, extensive knowledge of the systematics and ecology of spiders leads the authors to consider them as a very suitable group for pest limitation and for biodiagnostic purposes. An examination of both the qualitative and quantitative aspects of perdition by spider populations and communities is discussed as well as the evolution of some human factors occurring in agroecosystems that are likely to induce changes in spider predation such as chemical spraying and cultural practices. Studies addressing the recolonisation of agroecosystems by spiders, taking into account their dispersing abilities and habitat selection are summarised, followed by a discussion of the global efficiency of spiders as predators in such environments, the risks associated with their use and how to maximise their efficiency.The bioindicative value of spiders is presented by referring successively to population level and community level. The growth rate or the reproductive rate observed in natural populations can be correlated with the amount of prey ingested in the field. Thus, these parameters give an indirect estimation of the habitat quality. Two specific field experiments are presented to illustrate this ecological concept. Moreover, the role of spiders as indicators of heavy metal pollution (atmospheric or soil pollution) integrated by organisms living close to sources of pollution is discussed by reference to a set of laboratory and field experiments. Due to the close correspondence between the vegetation architecture and the composition of the associated spider community, it is argued (with a list of examples) that fluctuations in the spider community structure allows the bioevaluation of human disturbances. Based on the composition of the spider communities, methods of ecological classifications of natural habitats in several European countries are presented.