Association of spiders and
lichen on Robben Island, South Africa: a case report
Aeshita Mukherjee1,
Burkhard Wilske2, Rene A. Navarro3, Ansie
Dippenaar-Schoeman4 & L.G. Underhill3
1,3 Avian Demography Unit, University
of Cape Town, Rondebosch 7701, South Africa.
2 Blaustein
Institutes for Desert Research, Ben-Gurion University of the Negev, Sede Boqer
Campus, 84990 Midreshet Ben-Gurion, Israel 4 Agricultural Research Council, Plant Protection
Research Institute, Pretoria 0001, South Africa
Email: 1 aesh2003@yahoo.com
Date of publication (online): 26 April 2010
Date of publication (print): 26 April 2010
ISSN 0974-7907 (online) | 0974-7893 (print)
Editor: Cleofas R. Cervancia
Manuscript details:
Ms
# o2295
Received
25 August 2009
Final
received 20 January 2010
Finally
accepted 17 February 2010
Citation: Mukherjee, A., B. Wilske, R.A. Navarro, A. Dippenaar-Schoeman
& L.G. Underhill (2010). Association of spiders and lichen on Robben
Island, South Africa: a case report. Journal of Threatened Taxa 2(4): 815-819.
Copyright: © Aeshita Mukherjee, Burkhard Wilske, Rene A. Navarro, Ansie
Dippenaar-Schoeman & L.G. Underhill 2010. Creative Commons Attribution 3.0
Unported License. JoTT allows unrestricted use of this article in any medium
for non-profit purposes, reproduction and distribution by providing adequate
credit to the authors and the source of publication.
Author Details: Dr. Aeshita Mukherjee is an ecologist and was a postdoctoral fellow at the University of
Cape Town, South Africa.
Dr. Burkhard Wilske is an ecophysiologist, botanist having expertise in VOC and
carbon budgeting. He is presently a postdoctoral fellow at the Bio-Geosciences
Institute, University of Calgary, Canada.
Rene A. Navarro is an ecologist having expertise in satellite data analysis and,
presently working on spiders at the University of Cape Town in South Africa.
Prof. Ansie Dippenaar-Schoeman is an arachnologist and a well-known
taxonomist, presently a scientist at ARC, Plant protection, Pretoria, SA.
Prof. L.G. Underhill is a statistician, expertise in employing statistics to ecology.
Presently he is the Director of Avian Demography Unit, at the University of
Cape Town, South Africa.
Author Contribution: The first author designed the study, collected data, compiled and
analyzed the data and wrote this manuscript, while the second author’s input
was interpretation of result, literature search and helping in analysis. The
Third author contributed his time in conducting the field surveys. The fourth
author helped in species identification. The fifth author helped in all the
statistical analysis in the paper and was the project-in-charge.
Acknowledgements: Appreciation is extended to the Robben
Island Authority, SA for letting us conduct field studies and collect
specimens. AM is thankful to the
ADU, University of Cape Town for all the support and financial help during the
course of study. The Authors were also helped by the reviewers’ comments in
making the manuscript scientifically more sound.
Abstract: The
present study is a first record of spider occurrence on Robben Island, South
Africa. Some habitats were rich in lichens. As we know, lichens enhance wildlife habitat in less direct
ways. The objective of the study
was to examine the potential importance of lichens in enriching spider
diversity and abundance. A total
of 260 spiders (170 from lichens and 90 from bush) were collected following the
visual search method over one year. Seasonal trends in overall species richness and abundance indicated that
the relative density of spiders was greater in lichens than in bushes. The result suggests that habitat
structure, such as branch size and epiphytic lichen abundance, can be an
explanation for the greater number of spiders in lichen-rich patches of the
island.
Keywords: Robben
Island, spiders, lichens, visual search
Introduction
With the
exception of caribou, whose dependence on lichen is well documented, references
to lichens in literature on wildlife are few (Richardson & Young 1977;
Lawrey 1986; Blewitt & Cooper-Driver 1990). Lichens are less familiar to most researchers than vascular
plants; in animal studies they are frequently grouped with other fungi or with
mosses. Lichens enhance wildlife
habitat in less direct ways. It is worth noting that the given forage may appear
to be a rather small percentage of an animal’s yearly diet, yet it plays an
important strategic role.
Lichens are
not a popular food item for animals except for a few. However, Common Greenshield, and other lichens do cover or
give shelter to tiny animals, such as water bears, mites, springtails, spiders,
insects, and lizards. Some birds
are known to use lichens in their nests due to their insulating properties
(Richardson & Young 1977) while some amphibians are known to camouflage
perfectly among lichens. Several
species of invertebrates live on and among lichens, using them for concealment,
shelter or food. Over half of the
orders of insects have associations with lichens. There are striking cases of insects mimicking lichens, the
most well known being that of the peppered moth (Biston betularia) on
lichen-covered tree bark (Richardson 1974). Lichens are a part of many food webs that include
invertebrates and their predators. Spiders may prey upon, for example, small
lichen-eating arthropods, such as mites or bark lice. Birds, invertebrates and lichens were all found to be
functionally connected, and all were impacted by forestry practices (Pettersson
et al.1995). Many lichen species contain bitter compounds that may discourage
feeding by invertebrates, however, very little is known about the nutritional
quality of lichens (Lawrey 1986; Blewitt & Cooper-Driver 1990).
Other
literature shows that many small insects eat lichens exclusively (Hale
1972). According to Gerson &
Seaward (1977) and Seaward (1984), bark lice (Psocoptera) are voracious lichen
feeders. Some web spinners
(Embioptera) eat lichens; they live in silk tunnels, which they construct (Ross
1966; Anon 1970). A noctuid moth, Zanclognatha theralis, in
Tennessee, was found to both feed on Usnea
trigosa and to look remarkably like the lichen (Sigal
1984).
Many
orthopterans mimic lichens. Butterflies and moths (Lepidoptera) have the richest association with
lichens (Gerson & Seaward 1977). Many beetles (Coleoptera) live on or in lichens (Richardson 1974). Some examples of flies (Diptera) are
associated with lichens, and a number of studies have found ant (Hymenoptera)
nests built underneath lichen mats; Bailey (1970) observed the dispersal of
lichen propagules by ants. Some
species of termites (Isoptera) in the genus Hospitalitermes feed exclusively on lichens (Kalshoven 1958).
The mites
(Acari: Cryptostigmata) live on lichens in great numbers and in varying degrees
of dependence on them. These mites play important roles in soil energetic and
soil fertility (Seyd & Seaward 1984). At least one species deposits its eggs in lichens; it secretes a
substance which causes the lichen to swell and grow over the eggs, protecting
them. Oribatid mites are also the
largest category of lichen feeders (Lawrey 1987; Seyd & Seaward 1984).
Some
spiders (Araneae) disguise their webs with lichens, and the giant crab spider
of Trinidad constructs nests using lichens of the genus Usnea. Many spiders live in tundra lichens. Except for a few occasional studies no
detailed documentation on spiders using lichens as their habitat has been
reported. While working on Robben
Island, we found it very interesting to note that a few species appeared to be
highly dependent on lichens as their primary habitat. It is also worthy to note that spiders are sensitive to
microhabitat variation (Foelix 1996). This explains the occurrence of certain species of spiders in selected
habitat, governed by particular species of plants (Henschel & Lubin 1997;
Ward & Lubin 1993) and physical characteristics of the habitat like soil
and rocks (Foelix 1996). In
addition, low mobility of certain spider species and an innate preference for
certain habitats increases the effect of patchiness (Brandt & Lubin 1998).
Understanding the aspects outlined above, the present study aimed at
investigating:
(i) seasonal trends in species abundance and
the composition in a lichen spider community;
(ii) comparison of adjacent patches of
vegetation with regard to spider species composition and abundances.
Throughout
the country various monitoring projects were undertaken as part of the South
African National Survey of Arachnida. At ARC-Plant Protection Research Institute, Pretoria, for the first time
an inventory of the spider families, genera and species known from South Africa
were made. It was found that the
spider fauna of South Africa is remarkably rich when compared with some other
faunas of the world. A total of 67
families of the world’s spider families occur in this region represented by 428
genera and about 2900 species. This represents about 9% of the world’s fauna. However, looking at the literature and other published
reports we found that no documentation on spiders were reported from Robben
Island. Although there exists a
booklet describing the wildlife at Robben Island (Crawford & Dyer 2000) it
does not include any information on spiders.
While
working on birds at the island, we came across different species of spiders and
more interestingly in different microhabitats. We found an appreciable number of jumping spiders along the
shoreline. The present study is an
attempt to record the occurrence of spiders on Robben Island with special
emphasis on those depending on lichens. The study also compared them with the species occurring in the bushes,
which are the closest microhabitat adjacent to the lichens. The paper is a part of the arachnid
survey of the shorelines of South Africa, which is a first of its kind. To the best of our knowledge, this
paper presents the first record from Africa of spiders occurence related with
lichens.
Methods
Study Area
The study
was conducted on Robben Island, (33.80S, 18.360E), a
world heritage site, South Africa. Robben Island is roughly oval in shape; 3.3km long north south, and
1.9km wide, with an area of 5.07km² and is the largest in a string of
islands, which lie along the southwestern coast of Africa. The island is relatively flat, rising
to a height of 30m above sea level, as a result of an ancient erosion
event. It is 7km from the nearest
mainland (Cape Town). Winters are
cool and rainy, whereas summers are hot and dry making a very distinct climate
on the island. The indigenous
vegetation is quite similar to that of the mainland. The island makes a unique habitat, having both indigenous
and exotic species of flora and fauna. For recreation and for game, certain antelopes, mammals and birds were
introduced to the island. The
island harbours over 7000 penguins and several other sea birds and seals. The
government, during the apartheid, introduced ungulates and some other animals
now found on the island, as a resource for entertainment. The island has a distinct habitat of
grassland, and a rocky shore. About 100 human habitations exist on the island, which makes a different
surrounding for the occurrence of spiders. The island was mainly used until 1994 as the political
prison during the apartheid era. As a result the island was never surveyed for any scientific purposes.
Survey
The whole
island has very specific vegetation patches and so we took them separately to
monitor the spider abundance and richness. About 500m from the seashore the lichens grow both on the
rocks and on the semi dry branches. About 100m adjacent to it, bushes of Australian acacia occured void of
lichens. Visual search was
employed to collect the spiders occurring both on the bushes and on the
lichens. Every month collection
was done for 2 hrs (searching 20 random points) at each habitat for three
consecutive days. We also did
pitfall trapping, but the data obtained has not been used in this paper as they
may mislead by mixing of species from other microhabitats.
Data analysis
Spiders
collected were sorted as to species (if adult) or to genus or family if
juvenile. Species identifications
were confirmed by Prof. Dr. Ansie Dippenaar-Schoeman (ARC, Plant protection,
Pretoria) and specimens were deposited in the National Collection for reference.
We analysed
the monthly data from October 2003 to September 2004. Spider data were analysed for seasonal trends in overall
species richness and abundance and in the trends for different species. We
compared the two trapping sites with regard to spider abundance. Similarity indices were calculated to
compare the species between seasons and between the two sites
Results
Over the
year, 260 spiders (170 from lichens and 90 from bushes) were collected. Winter and early spring had lower
numbers of spiders than late spring, summer and autumn. April showed a distinct peak in
numbers, followed by stable numbers throughout summer and a smaller increase in
the autumn as a general trend. In
lichens the numbers were significantly high during autumn while in the bushes
the numbers were high in summer.
The
relative density of spiders was greater in lichens over the 12 monthly
samples. Overall, more spiders
were trapped in lichen (median, range: 41.5, 6-109) than in bushes (22.5, 1-74;
Wilcoxon signed ranks test, z = 2.275, p = 0.023). After square root transformation to normalize the data, we
performed repeated measures analysis on the total number of spiders, with
seasons repeated measures and site as a factor. The effect of season (within subjects) was significant (F3,
12 = 3.578, p = 0.047), but none of the sites (F1, 4 = 1.04, p
= 0.37) nor the interaction between site and season (F3, 12 = 0.643,
p = 0.6) were significant.
In the two
sites combined, we recorded 15 species, of which four were identified to
species level. In lichens there
were ten species, while in bushes eight species were recorded (Table1). Only three species were common to both
the habitats (Caerostris, Neoscona subfusca and Vidole capensis). Aridna sp. of the family Segestridae was found only on lichen on the whole
island. In lichens the most common
species was Neoscona subfusca (family Araneidae, 85 indv.) while in bushes, Steatoda and Theridion sp. were highest
in number (20 and 15, family Theridiidae). Only three species were common to both the habitats (Caerostris, Neoscona subfusca and Vidole capensis). The overall species diversity was
similar in both the sites (1.85 in lichen and 1.67 in bush).
Plant density
The plant
density at both locations was similar, (site lichen 3.95 and site bush 3.61; Wilcoxon test, z = -0.364,
n1 & n2 = 4, p = 0.715). Site
lichen had four species of shrubs and site bush had three shrub species.
Australian acacia constituted 87% of the shrubs in site lichen and 89% in
bushes. The shrub height and
diameter was similar (Average diameter ranged from 51cm to 59cm, and the height
varied from 26cm to 32cm).
Discussion
The
difference in the site characteristics and its influence on species richness
seemed difficult to explain, as the two sites are apart by approximately
100m. However, a possible
suggestion is vegetation composition, which makes one habitat more suitable
than the other one. Host plant
size is probably a factor, which accounts for the species abundance, as the
larger shrubs generally lead to an increase in species diversity of resource
available (Lawton 1983, 1986). It
is also known that the phytophagous fauna increases with a plant’s structural
complexity (Strong et al. 1984), and a distinct positive association exists
between beetle diversity and plant composition (Southwood et al. 1979). Further it is seen in the study that
the species composition shows a specific trend across the seasons. This could be due to the preference of
the species for a particular host vegetation. Most studies on spiders showed that there is a positive
association between species richness of orb builders and plant canopy complexity
(Stratton et al. 1979; Hatley & MacMohan 1980; Greenstone 1984). A similar pattern was found among the
hunting spiders (Duffey 1962 a,b; Uetz 1991). Although some species show little food discrimination, other
species show marked food and habitat preference (Thornton 1985). The availability of food across the
seasons may be one of the factors influencing the change in the species
composition. On the other hand
many spider species are highly mobile and some move long distances, while some
spiders are not fixed with the same plant structure (Forster & Forster
1999). While it is assumed that increased habitat complexity would provide more
environmental niches and foraging opportunities, the results indicate that this
is not the case for all organisms. Thus, the effects of habitat complexity cannot be generalized. Even within spider communities ground
dwellers may differ in their preference for habitat complexity versus tree
dwelling species suggesting the need to group species by functional groups (Lassau
& Hochuli 2004). There could
be a tremendous effect of season and the age of the trees bearing lichens to
predict spider richness. For
example, the spider richness was correlated and varied greatly with season and
weather in a forested landscape in Sweden (Gunnarsson et al. 2004).
In the
present study, only those specimen were considered that were found in the
habitats using visual search. So
it is more likely that only those were collected that walked/webbed on the
foliage. Therefore the studies do
not represent the whole community of spiders and other invertebrates. However, spiders were sampled from all
possible patches within the habitat, to get an overview in an ongoing study. In an ongoing study we are also
collecting spiders and other invertebrates from other habitats to add to this
preliminary survey. This study in
Africa is the first of its kind representing spider communities on an
island. It is suggested that
habitat structure (branch size and epiphytic lichen abundance) could be an
explanation for the greater number of spiders in lichen-rich patches of the
island. The study also emphasises
the usage of lichen as a very potential habitat, which has never been
documented earlier or studied in detail in South Africa and we hope such studies
will be taken up in the future.
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