Short-term impacts of prescribed burning on the spider community (Order: Araneae) in a small Ohio grassland.
Rose, Sarah J. ; Goebel, P. Charles
ABSTRACT. Prescribed burning is a management tool that is widely
accepted for prairie management and restoration, yet little is known how
burning may impact the spider community. Although it is generally
thought that prescribed burning may alter the spider community
composition and structure, few studies have examined these shifts in a
controlled manner with both a burned grassland and a nearby unburned
companion grassland. On 25 October 2014 we conducted a prescribed burn
of a grassland at the Gwynne Conservation Area, London, Ohio. Spiders
were sampled using pitfall traps for four weeks pre-burn and six weeks
post-burn in both the treatment grassland and adjacent unburned
grassland. A total of 298 spiders were collected from sixteen families,
over 60 percent of which were in the family Lycosidae. Overall, we found
the prescribed burn did not significantly alter the abundance or
diversity of spiders collected, and interestingly it appears the
community composition of the unburned grassland changed more over the
sample period than the burned grassland. Anecdotal observations also
suggest that some spiders are capable of surviving the fire in situ. As
we continue to study these communities, we will develop a better
understanding of the role that prescribed burning plays in regulating
the structure and composition of the spider communities. Such
information is important to develop process-based restoration and
management practices in grassland ecosystems.
Date of Publication: 1 December 2015
INTRODUCTION
Grassland ecosystems provide many valuable services, including but
not limited to: soil conservation, water quality enhancement, wildlife
habitat, and biodiversity (Risser 1996). Worldwide vast areas of
grasslands have been lost to a variety of human land use (Steinauer and
Collins 1996). Those grasslands that remain are highly fragmented
(Risser 1996) and more susceptible localized extinction events and
invasion by non-native species (Risser 1996), thus leading to the
conclusion that grassland systems should one of the top priorities of
conservation and restoration efforts (Sampson and Knopf 1994).
Before these restoration efforts begin, we need to better
understand the natural disturbance regimes and the influence these
disturbances have on ecosystem structure and composition, as restoration
efforts that emulate natural disturbances and their legacies are more
successful (Long 2009). In grasslands, frequent wildfires, usually in
the fall and ignited by lightning (Risser 1996, Steinauer and Collins
1996), were important natural disturbances. Fire in grasslands helps
reduce the encroachment of woody vegetation (Molles 2008; Hartley 2007),
increases nutrient cycling, and creates warm soil conditions that
promote seed germination (Kozlowski and Ahlgren 1974). Thus, prescribed
burning of grasslands has generally been shown to increase plant
productivity (Kozlowski and Ahlgren 1974), and as a result is considered
an important and inexpensive restoration and management tool (Whelan
1995; Zelhart and Robertson 2009). Yet, even with the known benefits of
burning of grasslands, some are concerned that prescribed burns may
negatively impact small isolated populations of invertebrates (Panzer
2002) or reduce beneficial arthropods such as pollinators and predators
in these ecosystems (Warren et al. 1987).
Despite their diminutive nature, spiders fill an important role in
many ecosystems. As one of the most numerous and higher level predators
of the arthropod world (Warren et al. 1987) they have been shown to be
good biocontrol agents of many pest and invasive species (Wise 1993) and
are important natural enemies of pest insects in many agroecosystems
(Buddie et al 2004). Spiders are a diverse group with multifaceted
methods of prey capture, each that can serve as an indicator to the
habitat in which they reside or utilize. They are also prey for many
animals, including birds, reptiles, amphibians, fish and mammals (Foelix
2011). Spiders are also abundant in most ecosystems (Wise 1993), and are
known to be pioneer colonizers in areas that have been recently altered
or disturbed (Bradley and Ohio Biological Survey 2004; Hodkinson et al.
2001). Spiders are also sensitive and respond quickly to environmental
conditions (Marc et al. 1999), making them a good choice as
bioindicators, especially when considering disturbances and their
effects on ecosystem structure and function.
In grassland habitats, it is expected that the number of
invertebrates (including spiders) would decrease significantly in the
short term following a fire either directly (i.e. mortality) (Reichert
and Reeder 1972) or indirectly (i.e. change in habitat structure and
microclimate) (Hore and Uniyal 2008; Hartley 2007). Although some have
hypothesized that spiders may survive a burn by seeking refuge in the
burrows or non-flammable plant matter (Warren et al. 1987; Jansen 2013),
Bell et al. (2001) suggested that this was unlikely due to the
sensitivity of even the most tolerant spider's physiology to minor
changes in temperature. Rice (1932) found that fire temperatures were
not severe enough to kill animals that were hibernating in the bases of
bunch grass during a spring burn in Illinois, and Brennan et al. (2011)
found that Xanthorrhoea preissii (grass trees) can serve as refugia for
some invertebrates during fire, although significant mortality was
detected. Thus it seems that spiders may be responding to changes in
habitat variables altered by burning in addition to direct mortality as
a result of the fire.
Taking advantage of a scheduled prescribed burn in a restored
grassland in central Ohio planned as a practical experience for students
acquiring red card certification as part of a wildland fire management
course at The Ohio State University, traps were set up to monitor the
spider community in order to determine if there are differences in the
spider community following a prescribed burning. Specifically, our
primary objective was to quantify the changes in spider species
community composition, diversity, and abundance following the prescribed
burn, and compare these changes with an adjacent unburned grassland. We
hypothesized that a large proportion of the spiders in a grassland
treated with prescribed burning will suffer mortality as a result of the
prescribed burn, and we would therefore observe a decrease in diversity
and abundance in spiders in the time period immediately following a
prescribed burn in the burned grassland, but that this decline would not
be observed in the adjacent unburned grassland.
METHODS
Study area
This study utilized two grassland areas at the Gwynne Conservation
area, a 27-hectare (67 acre) demonstration/education area that is part
of The Ohio State University's Molly Caren Agricultural Center
located in London, Ohio (39.95 N, -83.45 W). The administrators of the
wildland firefighter training class (offered through The Ohio State
University) selected the Big Bluestem Prairie (BBS^2 hectares) to be
used for a prescribed burning training, scheduled to occur on 25 October
2014. This prairie was originally established in 1989 and was planted
exclusively as Andropogon gerardii (big bluestem grass), although many
other grass and forb species have naturally established in the site
since establishment. The Prairie Planting (PP 0.8 hectares) was chosen
as a companion site for this study. It is approximately 350-m southeast
of BBS, was established in 1986 as a mixed-species prairie ecosystem,
and was not subjected to any management practices during the spider
sampling period.
Spider Sampling
To characterize the spider community, five pitfall traps were
installed along a transect with a minimum distance between traps of 10
meters, and a minimum distance to the grassland edge of 10 meters. At
each trap location a hole was dug such that a one gallon flower pot fit
snuggly into the hole with the top rim of the pot level with the natural
ground. A 0.9-L deli food container with ~5 cm of propylene glycol/dish
soap solution was placed in the flower pot. Propylene glycol was
selected as it helps to kill and preserve the specimens in the trap and
is less harmful to other wildlife than the alternatives (specifically
ethylene glycol). The dish soap acts to reduce surface tension on the
solution, causing the caught invertebrates to sink into the solution.
The wooden trap, following the design of Bradley and the Ohio Biological
Survey (2004) was then placed securely over the catch container and
flower pot. The roof and base were constructed using V4" plywood.
The base had a 7.6 cm hole cut into the center in which a solo cup with
the bottom removed was inserted to serve as a funnel, guiding the
invertebrates to the catch container. A 0.6-m x 0.6-m piece of chicken
wire was secured of the top of the trap with landscape pins to reduce
the chance of mammalian disturbance to the traps. Traps were installed
on 26 September, 2014, and samples were collected every two weeks
thereafter. The traps in both grasslands were removed on 24 October, and
the prescribed burn occurred on 25 October. Following the prescribed
burn the traps were reinstalled in both areas on 26 October. Samples
were collected weekly for the first two weeks post-burn, while
subsequent samples were collected every two weeks through 7 December,
for a total sample period of four weeks of sampling prior to the burn,
and six weeks of sampling following the burn. Spiders and other
invertebrates separated and stored in 70 percent ethanol until
identification.
Spiders were identified using a Nikon SMZ 1270 stereomicroscope.
Identification to genus was completed following Ubick et al. (2005), and
identification to species utilized resources available from the World
Spider Catalog (2015).
Data Analyses
Prior to analysis all early instar juveniles (i.e., early stage of
development) that were not identifiable past the family level were
excluded from all analyses. We also excluded those families that
represented less than one percent of the total catch over the entire
study period. In addition, Leucauge venusta (family Tetragnathidae) was
also excluded, as only one individual was trapped, and unlike the other
Tetragnathidaes captured, which are ground-dwelling spiders, L. venusta
is an orb-web dwelling species.
In order to characterize the differences in the spider community
each pitfall trap was treated as an independent replicate and samples
were pooled as either pre- or post-treatment and adjusted to
per-trapping-week. Pitfall traps can be considered independent if there
is sufficient spacing between traps (Woodcock 2005) and several other
studies have also treated individual traps as independent (Moore et al.
2002; Moretti et al. 2002; Obrist and Duelli 1996) with a minimum
distance of 10-m between traps. Furthermore, in order to provide the
most meaningful analysis of these data, even without true replication,
the use of inferential statistics can be used in order to provide the
most meaningful results (Oksanen 2001).
Shannon Diversity Index (Kent and Cocker 1992) was calculated for
each grassland overall, pre- and post-treatment overall, and pre- and
post-treatment by grassland. Comparisons between the grasslands and the
treatments were analyzed using Kruskal-Wallis tests in R (R Core Team
2013).
Species specific responses were analyzed using an indicator species
analysis (Dufrene and Legendre 1997) utilizing the Monte-Carlo
procedures (4999 permutations) with PC-ORD software (McCune and Mefford
1999). Indicator species analysis is a statistical approach that uses
species fidelity (relative frequency of a species within a group) and
exclusivity (relative abundance of a species within a group) to classify
species into groups that reflect environmental conditions represented by
sample units. In addition, the overall differences in the spider
community composition both before and after treatment were determined
using Multi-response Permutation Procedures (MRPP). MRPP is a
nonparametric procedure that is used to test a-priori groups for
differences in composition. (McCune et al 2002). Finally, to further
explore the patterns in spider community, both before and after
treatment, a nonmetric multidimensional scaling (nMDS) ordination plot
was performed with Bray-Curtis distance matrix calculated on per trap
week abundances by trap using the vegan package in R (R Core Team 2013).
Ordination techniques organize sampling entities along gradients to
explain the variability in the data, with nMDS being particularly useful
as it reduces the assumption of linearity (McGarigal et al. 2000).
RESULTS
4here were observable changes in the vegetation structure of both
grasslands during the study as the prescribed burn consumed most of the
vegetation and litter in the BBS, and the PP structure was altered by
snowfall and plant senescence (Fig. 1). It should also be noted that the
first frost [overnight low temperature of 0[degrees] C (32[degrees] F)
or lower] to occur during the sampling period occurred on 30 October
2014 and additionally there was a snowfall event totaling 7.87 cm (3.1
in.) on 17 November 2014. There was a decline in average temperature
highs and lows throughout the duration of the study, consistent with the
change from fall to early winter.
A total of 298 spiders from 14 families and 29 species were
collected. Over 80 percent (244) of these spiders were adults or
juveniles with enough characteristics to identify to species, genus, or
morphospecies, while the remaining nearly 20 percent (54) were only
identifiable to family (SupplementalTable 1). Lycosidae (61.7 percent)
and Linyphiidae (19.5 percent) were the most abundant families.
Comparisons of the Shannon Diversity Index showed a statistically
significant difference between the two grasslands overall (p = 0.02).
When the data for both grasslands was pooled and the pre- and
post-treatment was compared there was no significant difference detected
(p = 0.88). Additionally, comparing each grassland individually for the
pre- and post-treatment there was no significant difference in the
Shannon Diversity Index (p = 0.12 for the both grasslands). Although not
statistically significant, it should be noted that there did appear to
be an increase in the Shannon diversity index for the BBS when comparing
the pre- and the to the post-burn (H' of 1.69 and 1.95
respectively), and the PP showed the opposite trend, with a decrease in
the Shannon diversity index between the two sampling periods (H' of
1.71 and 1.49 respectively) (Fig. 2).
MRPP analysis at both the family and species level did not
demonstrate any statistically significant differences between the two
grasslands {p = 0.08 for both family and species-level analyses), but
comparing the pre-burn to the post-burn overall was significant (p =
0.001) (Table 2). Further analysis, comparing the pre- and post-burn of
each individual grassland showed a significant difference for both
grassland (p = 0.014 family level BBS, p = 0.013 species level BBS, p =
0.009 family level PP, p = 0.005 species level PP). Indicator species
analysis suggests that Varacosa avara (Lycosidae) was associated with
the BBS, and Neoantistea agilis (Hahniidae) was associated with the PP.
There were three indicators of the Pre-burn and one indicator of the
post-burn sampling periods when data from both grasslands were pooled.
BBSPre had one indicator species, BBS-post had four indicator species,
PP-Pre had two indicator species, and there were no indicators for the
PP-Post (Table 3).
[FIGURE 1 OMITTED]
The nMDS ordination (Fig. 3) resulted in a two dimension solution
with a final stress of 0.153 and shows some overlap of the spider
communities of the two grasslands prior to the burn, and shifts after 25
October. The resulting location of the post burn plots demonstrates a
greater similarity in the BBS post-burn site to the pre-treatment sites,
whereas the PP shows less similarity to either the pre-burn plots or the
BBS post-burn.
DISCUSSION
Most studies have shown a decrease in spider richness and/or
abundance in the time postburn when compared to pre-burn and/or control
sites (Rice 1932; Dunwiddie 1991; Zelhart and Robertson 2009; Riechert
and Reeder 1972; Pascoe 2003). Although it should be noted that these
studies [except Riechert and Reeder (1972) that used hand collection and
litter sorting] utilized sweep netting sampling exclusively.
Sweep-netting is a technique that is most often utilized to sample
arthropods dwelling on low vegetation (Ozannne 2005; New 1998).
Therefore, in the timeframe immediately following a fire, sweep netting
would yield minimal results, as there would be limited vegetation for
sweeping to occur on. Ground spiders, on the other hand, have been shown
to benefit or have no shortterm impacts from burning in two studies that
utilized pitfall trapping (Hore and Uniyal 2008; Jansen et al. 2013).
Understanding sampling techniques is an important aspect to arthropod
research as there is potential for biases and errors (Leather and Watt
2005). In comparing pitfall traps, sweep nets, and visual searches
Churchill and Arthur (1999) found that 94 percent of families were
captured in pitfalls, 25 percent with sweep nets and 41 percent by
visual sampling. Therefore, when trying to determine short-term effects,
when nearly all the vegetation and litter are presumed to be consumed in
the prescribed burn, pitfall trapping seems to be the most logical
sampling technique to examine the immediate and short term impacts of
prescribed burning on spider communities.
[FIGURE 2 OMITTED]
We did not find evidence to support the hypothesis that there would
be a reduction in diversity and abundance in the time frame immediately
following burning. In fact we observed an increasing trend in the
Shannon Diversity Index in the burned grassland post burning, although
it was not statistically significant. This leads to the conclusion that
spiders are either surviving the fire in situ, or are able to recolonize
the area very quickly. In the mop-up phase of the prescribed burn
numerous spiders were observed on the burned surface (Fig. 4). In
addition, a Varacosa avara (Lycosidae) male was captured in the trapping
period of 23 November 2015 to 07 December 2015 with obvious burn
injuries to his extremities (Fig. 5). As the burn occurred on the 25
October, one explanation is that he suffered the injuries in the
prescribed burn and survived until he was captured in the pitfall trap
several weeks later. Additionally, as there are multiple grassland
habitats in close proximity to the burned grassland at the Gwynne
Conservation Area, and ballooning spiders were observed on sampling days
after the prescribed bum, it is likely that recolonization was also
occurring. Other studies have stated the importance of maintaining
refuge habitat and varying the spatiotemporal variation (i.e. burning on
a rotational basis) among sites in these types of ecosystems (Swengel
2001) in order to provide source populations for recolonization. Thus a
combination of survival and recolonization may be responsible for the
lack of a decline in diversity and abundance.
[FIGURE 3 OMITTED]
[FIGURE 4 OMITTED]
Although shifts in community composition were detected, they seem
more pronounced for the companion unburned grassland then for the
grassland subjected to the prescribed burn. As this study took place in
the fall it is possible that these shifts are just part of the
phenological changes in spider community that occur naturally each year.
As the two grasslands utilized for the study were significantly
different in the pre-burn time frame it is not possible to use the
unburned as a control, therefore we are not able to conclude if any of
the changes were specifically due to the burn. Further studies would
need to be completed to evaluate this in more detail, with greater
sampling size and better replication. It is clear that we still are
lacking in our knowledge of the impact of prescribed burning on the
spider community and further studies are warranted in order for land
managers and restoration ecologists to gain the insights needed for
proper care of these ecosystems.
[FIGURE 5 OMITTED]
ACKNOWLEDGMENTS
Salaries and research support for this research was provided by
state and federal funds appropriated to the Ohio Agricultural Research
and Development Center (OARDC) at The Ohio State University. We
especially thank the support of the Gwynne Conservation Area for
allowing us access to the site, especially Nick Zachrich and Marne
Titchenell for their assistance in setting up this research project. We
also wish to thank The Ohio State University's Wildland Fire
Management class (ENR 3335) for executing the prescribed burn,
especially Mike Bowden (ODNR) and Dr. Roger Williams (Instructor). We
are very grateful to Dr. Richard A. Bradley for his assistance
throughout this project, including advising on pitfall trap
construction, study design, and tremendous assistance with spider
identification. We would also like to thank Timothy Rose for his
assistance in the field, and John and Elizabeth Rose for their
assistance in constructing the pitfall traps. Finally, we wish to thank
the two anonymous reviewers who provided useful editorial suggestions.
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SARAH J. ROSE (1) and P. CHARLES GOEBEL, School of Environment and
Natural Resources, Ohio Agricultural Research and Development Center,
The Ohio State University, Wooster, OH, USA
(1) Address correspondence to Sarah J. Rose, 2021 Coffey Rd., The
Ohio State University, Columbus, OH 43210. E-mail: rose.891@osu.edu
Table 2
MRPP
Comparison Species/ Delta Delta Delta Delta A
Family 1 2 3 4
BBS vs PP Family 6.675 6.646 na na 0.078
BBS vs PP Species 6.691 6.604 na na 0.075
Pre vs Post Family 3.943 3.878 na na 0.459
Pre vs Post Species 4.001 3.84 na na 0.455
4 categories Family 2.374 2.366 2.26 2.105 0.686
4 categories Species 2.521 2.398 2.308 2.107 0.676
BBS Pre vs Post Family 2.374 2.24 na na 0.654
BBS Pre vs Post Species 2.521 2.308 na na 0.639
PP Pre vs Post Family 2.366 2.105 na na 0.664
PP Pre vs Post Species 2.398 2.107 na na 0.659
Comparison Observed Expected P-Value
Delta Delta
BBS vs PP 6.661 7.225 0.08
BBS vs PP 6.648 7.12 0.08
Pre vs Post 3.91 7.225 <0.01
Pre vs Post 3.921 7.19 <0.01
4 categories 2.271 7.225 <0.01
4 categories 2.334 7.19 <0.01
BBS Pre vs Post 2.307 6.675 0.01
BBS Pre vs Post 2.414 6.691 0.01
PP Pre vs Post 2.236 6.646 0.01
PP Pre vs Post 2.253 6.604 <0.01
Table 3
Indicator Species Analysis by Habitat Type and Sampling
Period Species abbreviation (indicator value, p-value)
Habitat Type
BBS PP Pre Post
Vaav Neag Pasa Agpr
(68.2, 0.01) (50.0, 0.03) (70.0, <0.01) (60.0,0.01)
Pimi
(70.0, <0.01)
Tihe
(81.8, <0.01)
Habitat Type
BBS-Pre BBS-Post PP-Pre PP- Post
Pasa Erau Neag none
(83.3, <0.01) (44.1, 0.04) (48.9, 0.05)
LinUnkl Pimi
(51.8,0.03) (83.3, <0.01)
Vaav
(51.5,0.03)
Pisp
(60.0, 0.03)
Supplemental Table 1
Spiders collected by Grassland and Sampling Period
Family
genus species (abbreviation) Author
Anyphaenidae
Wulfila saltabundus (Wusa) Hentz 1847
Araneidae
Araneidae early instars
Clubionidae
Elaver sp. (Elsp)
Dictynidae
Circurina robusta (Ciro) Simon 1886
Dictynidae unknown I (DiunkI)
Gnaphosidae
Drassyllus sp. (Drsp)
Micaria pulicaria (Mipu) Sundevall 1832
Hahniidae
Neoantistea agilis (Neag) Keyserling 1887
Linyphiidae
Bathyphantes pallidus (Bapa) Banks 1892
Centromerus cornupalpis (Ceco) O. Pickard-Cambridge 1875
Erigone aletris (Eral) Crosby & Bishop 1931
Erigone autumnalis (Erau) Emerton 1882
Grammonota pictilis (Grpi) O. Pickard-Cambridge 1875
Mermessus bryantae (Mebr) Ivie & Barrows 1935
Mermessus maculatus (Mema) Banks 1892
Linyphiidae unknown I (Liunkl)
Linyphiidae unknown II (Liunkll)
Linyphiidae unknown III (Liunkll)
Liocranidae
Agroeca pratensis (Agpr) Emerton 1890
Agroeca sp. (Agsp)
Lycosidae
Paradosa saxatilis (Pasa) Elentz 1844
Pardosa sp. (Pasp)
Pirata/Piratula sp. (Pisp)
Piratula minuta (Pimi) Emerton 1885
Rabidosa punctulata (Rapu) Hentz 1884
Tigrosa helluo (Tihe) Walckenaer 1837
Varacosa avara (Vaav) Keyserling 1877
Lycosidae unknown I (Lyunkl)
Lycosidae unknown II (Lyunkll)
Lycosidae unknown III (Lyunklll)
Lycosidae Early Instars
Mysmenidae
Mysmenidae unknown (Myun)
Oxyopidae
Oxyopes sp. (Oxsp)
Philodromidae
Ebo iviei (Ebiv) Sauer & Platnick 1972
Philodromus sp. (Phsp)
Salticidae
Marpissa lineata (Mali) C.L. Koch 1846
Tetragnathidae
Leucauge venusta (Leve) Walckenaer 1841
Pachygnatha tristriata (Patr) C.L. Koch 1845
Theridiidae
Theridion sp. (Thsp)
Trachelidae
Meriola decepta (Mede) Banks 1895
Family 26 Sept.- 26 Oct.-
24 Oct. 7 Dec.
BBS- PP- BBS- PP-
genus species (abbreviation) Pre Pre Post Post
Anyphaenidae 0 0 3 0
Wulfila saltabundus (Wusa) 0 0 3 0
Araneidae 1 0 2 2
Araneidae early instars 1 0 2 2
Clubionidae 0 0 1 0
Elaver sp. (Elsp) 0 0 1 0
Dictynidae 0 0 1 1
Circurina robusta (Ciro) 0 0 1 0
Dictynidae unknown I (DiunkI) 0 0 0 1
Gnaphosidae 3 2 3 0
Drassyllus sp. (Drsp) 3 1 3 0
Micaria pulicaria (Mipu) 0 1 0 0
Hahniidae 0 6 0 2
Neoantistea agilis (Neag) 0 6 0 2
Linyphiidae 11 7 35 5
Bathyphantes pallidus (Bapa) 6 2 6 1
Centromerus cornupalpis (Ceco) 0 0 1 0
Erigone aletris (Eral) 0 0 2 1
Erigone autumnalis (Erau) 0 1 7 1
Grammonota pictilis (Grpi) 0 0 2 0
Mermessus bryantae (Mebr) 2 2 3 1
Mermessus maculatus (Mema) 1 0 1 1
Linyphiidae unknown I (Liunkl) 2 2 11 0
Linyphiidae unknown II (Liunkll) 0 0 1 0
Linyphiidae unknown III (Liunkll) 0 0 1 0
Liocranidae 0 0 6 6
Agroeca pratensis (Agpr) 0 0 4 6
Agroeca sp. (Agsp) 0 0 2 0
Lycosidae 76 55 34 19
Paradosa saxatilis (Pasa) 10 2 0 0
Pardosa sp. (Pasp) 13 12 10 8
Pirata/Piratula sp. (Pisp) 0 0 3 0
Piratula minuta (Pimi) 2 10 0 0
Rabidosa punctulata (Rapu) 4 5 0 2
Tigrosa helluo (Tihe) 10 10 3 0
Varacosa avara (Vaav) 7 1 16 3
Lycosidae unknown I (Lyunkl) 1 0 0 2
Lycosidae unknown II (Lyunkll) 0 1 0 1
Lycosidae unknown III (Lyunklll) 0 0 0 1
Lycosidae Early Instars 29 14 2 2
Mysmenidae 0 0 1 0
Mysmenidae unknown (Myun) 0 0 1 0
Oxyopidae 0 0 1 0
Oxyopes sp. (Oxsp) 0 0 1 0
Philodromidae 0 0 2 0
Ebo iviei (Ebiv) 0 0 1 0
Philodromus sp. (Phsp) 0 0 1 0
Salticidae 0 0 2 0
Marpissa lineata (Mali) 0 0 2 0
Tetragnathidae 3 0 1 0
Leucauge venusta (Leve) 0 0 1 0
Pachygnatha tristriata (Patr) 3 0 0 0
Theridiidae 0 0 2 0
Theridion sp. (Thsp) 0 0 2 0
Trachelidae 1 1 0 3
Meriola decepta (Mede) 1 1 0 3
