The diversity of thrips (Insecta: Thysanoptera) on buffel grass (Cenchrus ciliaris) is markedly lower than on native grasses in an urban landscape
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Journal of Urban Ecology, 2020, 1–7
doi: 10.1093/jue/juaa024
Short Notes
The diversity of thrips (Insecta: Thysanoptera) on
buffel grass (Cenchrus ciliaris) is markedly lower
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than on native grasses in an urban landscape
Christopher M. Palmer1,* and Laurence A. Mound2
1
Biodiversity Conservation Division, Northern Territory Department of Natural Resources, Environment, the
Arts and Sport, PO Box 1120, Alice Springs, NT 0871, Australia and 2Australian National Insect Collection,
CSIRO, PO Box 1700, Canberra, ACT 2601, Australia
*Corresponding author. E-mail: cmpalmer2005@gmail.com
Submitted: 27 February 2020; Received (in revised form): 26 July 2020. Accepted: 13 August 2020
Abstract
Buffel grass (Cenchrus ciliaris) is a perennial tussock grass native to parts of Africa and southern Asia but is now distributed
throughout much of mainland Australia. Corresponding with its presence in Australia has been an alteration of fire regimes
and soil nutrients, and a reduction in floral diversity. We investigated whether buffel grass supported a less diverse commu-
nity of thrips (order Thysanoptera) compared to the native grasses Cymbopogon ambiguus and Themeda triandra growing in
Alice Springs, central Australia. The survey data showed that abundance was highest on buffel grass; however, the thrips
community was almost entirely dominated by one species. The thrips communities on the native grasses were similar and
markedly more diverse than that on buffel grass, both in terms of species richness and functional feeding groups. Flower
feeders constituted the greatest proportion of thrips on all three grasses, but the native grasses also supported leaf feeders
and predators. The results indicate that thrips are sensitive to vegetation change, and we suggest that active removal of buf-
fel grass and replacement with local native grasses would help reverse the loss of biodiversity and normal ecological func-
tion in urban areas like Alice Springs. We also suggest that the order Thysanoptera is under-utilised in biodiversity
research.
Key words: arid zone, biodiversity, Australia, exotic weeds, invasive species, Poaceae
Introduction
invade very large areas encompassing a range of ecosystems
Environmental weeds are plants that invade natural communi- and habitats.
ties, and these weeds can have serious deleterious effects on Buffel grass (Cenchrus ciliaris) is a perennial tussock grass na-
ecosystem structure and function. Such effects include: threat- tive to tropical and sub-tropical parts of Africa, the Middle East
ening native biodiversity through competition with native and southern Asia (Lazarides, Cowley, and Hohnen 1997). This
plants for resources and replacement, increasing fuel loads for species was introduced to arid and semi-arid regions of the
fires, and changing water and soil quality and characteristics world (e.g. Australia and Central and South America) as a pas-
(Lonsdale 1999; Williams and West 2000). Some species are ture plant because it is drought tolerant, fire resistant and
much more invasive than others, and the worst weeds can responds quickly to rain. It also has a rapid growth rate, with
C The Author(s) 2020. Published by Oxford University Press.
V
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12 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
prolonged flowering period and prolific seed production (Franks For this study, we surveyed individual plants of the target
2002). With these attributes, it rapidly establishes self-sustain- grass species. This approach differs from other studies (e.g.
ing populations in a wide range of habitats. The species was in- Smyth, Friedel, and O’Malley 2009), that surveyed the space be-
troduced into Australia in the 1870s (Marriott 1955) and was tween target grass species and other species. We took this ap-
sown throughout Queensland and New South Wales from the proach because when it invades, buffel grass often becomes a
1920s (Humphreys 1967), and northern parts of the Northern component of a mixed community of plants, and this has led to
Territory in the 1950s and 1960s (Cameron et al. 1984). Plantings difficulties in establishing a causal relationship between per-
were conducted in central Australia throughout the 1960s and centage cover and altered diversity of mobile organisms (but
1970s for pasture improvement, prevention of soil erosion and see Bonney, Andersen, and Schlesinger 2017).
dust control (Keetch 1981; Allan 1997). Buffel grass has now be- There is a tendency to assume that weeds have negative
come dominant in Australia, forming monocultures across the effects on biodiversity only in ‘natural’ areas such as National
rangelands (van Klinken, Panetta, and Coutts 2013), and is esti- Parks, and that the threats to and effects on biodiversity are not
mated to infest 30 million hectares in Queensland alone such an issue in already profoundly altered landscapes such as
(Hannah and Thurgate 2001). Buffel grass has also spread widely towns. However, we also took the approach that potentially
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from the introduction points in central Australia to occur across demonstrating a negative effect of buffel grass on biodiversity
all land tenures (Puckey and Albrecht 2004). in a town could increase public awareness of its impacts, bring-
Concomitant with this nationwide expansion has been an ing the issue closer to the community and increasing support
alteration of fire regimes and soil nutrients, and a reduction in for its long term and widespread management.
biodiversity. In Queensland, converting areas of native mulga to
buffel grass pastures led to declines in soil mineral composition
(Mathers, Harms, and Dalal 2006); and Franks (2002) and Jackson Methods
(2005) demonstrated that native plant species richness
Study sites and focal taxa
was lower in buffel grass-dominated sites. The same result oc-
curred in Alice Springs (Clarke, Latz, and Albrecht 2005). In cen- The area for fieldwork was within Alice Springs (23 420 S, 133
tral Australia, seed viability and germination rate of native 520 E), a small town in the Northern Territory (central Australia)
perennial shrubs is reduced in areas invaded by buffel grass with an area of 328 km2, and a population of 26 000 (id
(Edwards et al. 2019), and longer-lived perennial trees suffered Consulting Pty Ltd 2019). Buffel grass is unmanaged around the
higher mortality due to fires fuelled by buffel grass (Schlesinger, town, and it grows commonly in all disturbed areas. The native
White, and Muldoon 2013). Many ground-dwelling insect taxa grasses Cymbopogon ambiguus (lemon-scented grass) and
are also likely to be negatively affected by such fires (Sands Themeda triandra (kangaroo grass) were surveyed together with
2018); however, there is generally less information on the C. ciliaris, as they are also perennial tussock species that grow
effects of buffel grass on invertebrates compared with its effects albeit less commonly around the town.
on plants. The aim of the sampling was to achieve sufficient coverage
The major aim of this study was to investigate the effect of of the urban environment, and so sampling sites were well-
buffel grass on insect communities, specifically whether insect spread throughout the town. Each grass species was sampled at
communities on buffel grass are different to those occurring on three sites within Alice Springs (Fig. 1), with five or six individ-
native grasses. The insect taxa selected for survey and identifi- ual tussocks of each grass species surveyed at each site.
cation were thrips (order Thysanoptera). Thrips were chosen for Selection and exact size of each site was dependent on the pres-
this study because members of the Poaceae support a particu- ence of appropriate flowering grasses, but each site wasThe diversity of thrips (Insecta: Thysanoptera) on buffel grass | 3
the raw measure as a more intuitive indicator of dominance: a
higher number means reduced diversity and increased
dominance.
To investigate similarity between thrips communities on
each of the three grasses, EstimateS was also used to calculate
the Morisita–Horn index (CMH), one of the few similarity indices
not strongly influenced by species richness and sample size
(Magurran 2004). Values for the index vary from 0 (no similarity)
to 1 (complete similarity). Finally, Statistica (version 13; TIBCO
Software Inc.) was used to perform a cluster analysis on the raw
thrips abundance data, using Euclidean distances to depict the
similarity in thrips communities between the three grass
species.
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Results
A total of 6497 thrips specimens were collected, from eight de-
scribed genera and one undescribed genus distributed in three
of the six families of Thysanoptera occurring in Australia
(Table 1). The most abundant genera were Haplothrips (5598 indi-
viduals including 412 larvae), Podothrips (538 adults) and
Anaphothrips (293 adults). The thrips fauna on C. ciliaris was
dominated by adults of Haplothrips froggatti, which constituted
93.8% of the total abundance for this grass (Fig. 2a). Only 0.4% of
the total abundance on C. ciliaris was made up by adults of other
species; a total of 15 individuals. Adults of H. froggatti consti-
tuted 50% of the total abundance of both native grasses, with
much of the remaining diversity made up of adults of several
other genera (Table 1 and Fig. 2a). Arranging the data into func-
tional groups based on feeding preferences, 99.9% of the fauna
on buffel grass was made up of flower feeders; the same group
constituted 65.0% on C. ambiguus, and 78.8% on T. triandra
Figure 1: Map of Alice Springs (central Australia), showing thrips sampling sites (Fig. 2b). Also present in relatively large proportions on the na-
for one introduced grass (C. ciliaris) and two native grasses (C. ambiguus and T. tive grasses were leaf feeders (11.9% on C. ambiguus and 10.9%
triandra). on T. triandra) and predators (23.2% on C. ambiguus and 7.5% on
T. triandra; Fig. 2b).
Mixed ANOVA showed that there were no significant differ-
‘between group’ variable was grass species, and the ‘within ences in the thrips communities inhabiting the three grasses
group’ variable was thrips taxa. The nine samples were included (Table 2), meaning the nine samples were not independent and
as random effects. likely came from one underlying thrips population. There was
Species diversity has two components: species richness (the also no interaction of grass species thrips taxa. The mean
number of species) and evenness (the variability in species number of thrips taxa per site was lowest for C. ciliaris (2.7),
abundances) (Magurran 2004). Information on both components higher for C. ambiguus (5.0), and highest for T. triandra (7.3;
is incorporated into diversity indices, the relative weighting of Table 3). As the nine samples were not independent, data were
each varying with the statistic used. To compare cumulative pooled for the measures of diversity, evenness and dominance
species richness, EstimateS 9.1.0 (Colwell 2013) was used to cal- for each grass shown in Table 3.
culate S(est) for each of the three grass species, which is the Based on the survey data C. ciliaris harboured the greatest
expected number of thrips species on each grass given the sam- abundance but the lowest number of thrips taxa (Tables 1 and
ples obtained in this study. To assess statistical differences of 3). Themeda triandra harboured the lowest abundance but the
S(est) between the three grasses, we compared the 84% confi- greatest number of thrips taxa. Each grass species supported ex-
dence intervals according to MacGregor-Fors and Payton (2013). otic thrips taxa, although the abundance of these taxa on each
To investigate how evenly distributed the sampled thrips grass was very low.
individuals were among thrips species, we also used Excel to Cumulative richness of estimated thrips species (Sest) was
calculate the Simpson Evenness index (ED ¼ D/DMAX) after first statistically lower on C. ciliaris (Sest ¼ 7.33, CI 84% ¼ 5.71–8.95)
calculating the Simpson Diversity index (D) using the method than either C. ambiguus (Sest ¼ 11, CI 84% ¼ 9.15–12.85) or T. tri-
described in Gardener (2017). The Simpson Evenness index andra (Sest ¼ 13, CI 84% ¼ 11.05–14.95) based on calculated S(est)
assumes a value between 0 and 1, where 1 is complete equita- values and 84% CIs (Table 3). Calculated ED values showed that
bility of individuals across the sampled species. the distribution of individuals among thrips species was very
We also calculated the Berger–Parker index (d) (Berger and uneven but similar for both native grasses (0.081 for C. ambiguus,
Parker 1970) as a measure of dominance in the thrips samples. 0.072 for T. triandra; Table 3). The distribution was extremely un-
This expresses the proportional abundance and relative impor- even on C. ciliaris (0.002).
tance of the most abundant species in the assemblage, and was The native grasses T. triandra and C ambiguus shared more
calculated using the formula described by Magurran (2004). taxa than did C. ciliaris with either of the native grasses
Although often expressed in its reciprocal form (1/d), we present (Table 4). The Morisita–Horn indices demonstrate that the4 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
Table 1: Thrips species and abundance on three grasses in Alice Springs, central Australia, in 2008–2009a
Grass species
Thrips taxa C. ciliaris C. ambiguus T. triandra Feeding strategy
S.O. Tubulifera
Phlaeothripidae
Phaeothripinae
Haplothrips (Haplothrips) froggatti 3575 1053 253 FF (Mound and Minaei 2007)
H. (Haplothrips) ganglebaueri 0 221 76 FF (Mound 2019)
H. (Haplothrips) sp. 0 3 0 FF (assumed)
H. (Trybomiella) sp. 0 0 5 FF (Moritz and Mound 1999)
Haplothrips larvae 223 116 73 FF (inferred)
Podothrips australis 0 498 40 PR (Mound and Minaei 2007)
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S. O. Terebrantia
Aeolothripidae
Desmothrips sp. 3 0 0 FF/PR (Pereyra and Mound 2010)
Thripidae
Thripinae
Anaphothrips incertus 0 231 57 LF (Mound and Tree 2020)
A. sudanensisb 0 0 1 LF (Mound, Hoddle, and Hastings 2019)
Anaphothrips sp. 4 0 0 LF (Mound and Masumoto 2009)
Arorathrips mexicanusb 6 5 9 FF (Mound, Hoddle, and Hastings 2019)
Frankliniella schultzeib 1 0 3 FF (Mound, Hoddle, and Hastings 2019)
Scolothrips sp. 1 1 0 PR (Mound 2011)
Panchaetothripinae
Phibalothrips longiceps 0 24 0 LF (Mound and Tree 2020)
Thripidae gen. nov. 0 0 15 UNK
Total abundance 3813 2152 532
a
Data are pooled from three samples for each grass.
b
Taxon known to be exotic to Australia; FF, flower feeder; LF, leaf feeder; PR, predator; UNK, Unknown.
assemblages on the native grasses were more similar than communities on the native grasses were less dominated by one
those of either native grass and C. ciliaris (Table 4). or more thrips taxa.
Euclidian distances between thrips communities on each Cenchrus ciliaris had the lowest and T. triandra the highest es-
grass species are shown in Table 4, with a vertical tree plot of timated total thrips species richness of the three grasses
these distances shown in Fig. 3. The lowest distance was be- (Table 3). The Sest 84% confidence intervals for C. ambiguus
tween T. triandra and C. ambiguus, denoting greater similarity (9.15–12.85) and T. triandra (11.05–14.95) overlap, leading to the
between the thrips communities on these grasses. inference that estimated thrips species richness is not statisti-
cally different between the two native grasses, further reflecting
their similarity.
The results supported the prediction of lower insect diversity
Discussion on the exotic grass; however, the highly abundant though
Each grass was surveyed at three locations around the town, highly skewed fauna on buffel grass was surprising. Haplothrips
meaning nine samples in total. The results of the mixed ANOVA froggatti is also known as the ‘black plague thrips’, as it is well
showed that the nine samples likely came from one underlying known to occur in vast numbers across central Australia
thrips population. The mixed ANOVA also revealed significant (Mound and Minaei 2007). Like the majority of Haplothrips spe-
differences between individual thrips taxa and to what extent cies (Mound and Zapater 2003), H. froggatti and H. ganglebaueri
each occupied the grasses; to be expected with a fauna in Alice breed and feed in flowers. Species of Anaphothrips are also phy-
Springs heavily dominated by four species out of the total of 14 tophagous (on leaves), with A. incertus commonly collected from
taxa that were collected. Australian native and not introduced grasses (Mound and
In this investigation, we asked whether buffel grass sup- Masumoto 2009). Podothrips species are predators of grass-living
ported a similar diversity of thrips species compared to the di- scale insects (Mound and Minaei 2007) that feed on the phloem
versity on native grasses, and if there were any differences of their host plants. Podothrips australis occurs widely through-
whether there were any likely implications regarding ecosystem out inland areas of Australia (Mound and Minaei 2007), and the
function. Data from these surveys showed that buffel grass sup- known collection records are from several native and intro-
ported a community of thrips that was markedly different to duced species of Poaceae (Ritchie 1974). None of the native
that supported by two native grasses growing in the same area. thrips H. ganglebaueri, P. australis or A. incertus was found on
The thrips fauna on the native grasses C. ambiguus and T. trian- C. ciliaris during this study, but all three were abundant on
dra was more similar, and each of the thrips faunas on the na- C. ambiguus and T. triandra, suggesting a distinct preference for
tive grasses was more species-rich and more evenly distributed native grasses. The abundance of the predator P. australis on
than the fauna on C. ciliaris. None of the thrips communities on C. ambiguus and T. triandra indicates that the native grasses also
the three grasses approached equitability, although the thrips support communities of other insect groups that buffel grassThe diversity of thrips (Insecta: Thysanoptera) on buffel grass | 5
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Figure 2: Relative proportions of: (a) thrips taxa and (b) thrips functional groups for each grass, based on feeding preferences. Data for each grass combined from all
sites.
Table 2: Results of ANOVA: two-factor with replicationa
absent. In contrast, the two native grasses supported predators
Source of variation df F p value and leaf feeders in high abundance together with flower
feeders. Although flower feeders constituted the greatest pro-
Between grass species 2 1.079 0.398 portion of thrips on all three grasses, the functional diversity of
Within grass species thrips was higher on the native grasses. These results suggest
Thrips taxa 13 3.023 0.001 that with the expansion of buffel grass there has been a con-
Grass species Thrips taxa 26 1.230 0.240 comitant reduction in resources available to native thrips spe-
a
cies and other insect taxa, with likely lasting reductions in their
Haplothrips larvae are excluded from the analysis.
distribution and abundance and the ecosystem services they
provide.
does not, although other insect taxa were not surveyed during The results of this investigation are congruent with the find-
this study. As buffel grass is more abundant than C. ambiguus ings of previous studies in central Australia that highlighted the
and T. triandra in and around Alice Springs, the native grasses negative impacts of buffel grass on different functional groups
may constitute disjunct patches of more suitable habitat within of ants. Both Smyth, Friedel, and O’Malley (2009) and Bonney,
areas of unsuitable habitat, thereby concentrating populations Andersen, and Schlesinger (2017) reported reduced abundance
of some thrips taxa as well as other insect communities on the of the ‘hot-climate specialist’ group of seed feeding ants in areas
native grasses. infested by buffel grass. Bonney, Andersen, and Schlesinger
The thrips fauna on buffel grass was dominated by flower (2017) also found a strong negative association between buffel
feeders, with other functional feeding groups almost entirely grass and overall ant abundance and species richness,6 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
Table 3: Thrips species richness, diversity and dominance on the three grassesa
Thrips
Grass Mean no. Total No. of No. of Estimated richness Evenness Dominance
of taxa taxon unique known
per site richness taxa exotic S(est) S(est) 84% CI S(est) 84% CI ED d
taxa lower bound upper bound
C. ciliaris 2.7 6 2 2 7.33 5.71 8.95 0.002 1.00
C. ambiguus 5.0 8 2 1 11 9.15 12.85 0.081 0.52
T. triandra 7.3 9 3 3 13 11.05 14.95 0.072 0.55
a
Haplothrips larvae are excluded from all measures.
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Table 4: Shared thrips species and similaritya of the indigenous flora is one of the key approaches to insect
conservation in urban environments, both in Australia and over-
Grass species No. of shared Similarity Euclidean
seas (e.g. Goddard, Dougill, and Benton 2010; New 2018).
thrips taxa (CMH) distance
In conclusion, we demonstrated that buffel grass altered the
C. ciliaris and C. ambiguus 3 0.766 2591 diversity of Thysanoptera occurring in a semi-arid, urban land-
C. ciliaris and T. triandra 3 0.818 3324 scape, likely leading to diminished ecosystem processes that
T. triandra and C. ambiguus 5 0.956 950 normally occur on locally native plants. Given that buffel grass
is established across arid and semi-arid Australia, it is likely to
a
Haplothrips larvae are excluded from all measures. have had major, irreversible impacts on thrips communities in
habitats throughout these climatic zones. The impacts would
have been greatest on members of the Phlaeothripidae, which
has had several endemic radiations in semi-arid parts of the
country (Mound and Tree 2020).
The results from this investigation indicate that thrips are
sensitive to vegetation change. We are unaware of any similar
studies in Australia using thrips as the focal organism, and sug-
gest that the order Thysanoptera is under-utilised in biodiver-
sity research. This investigation, together with a growing
number of other studies (e.g. Bonney, Andersen, and
Schlesinger 2017) are increasing our understanding of the nega-
tive impacts of buffel grass on invertebrate biodiversity and
ecology in Australia.
Despite the almost ubiquitous presence of buffel grass in
many parts of Australia and its impacts on ecosystems both
here and overseas (e.g. Flanders et al. 2006), it is not listed as a
weed in any Australian state or territory, although it is listed on
the Global Invasive Species Database (2019). This is mainly be-
Figure 3: Vertical tree plot of Euclidian distances between thrips communities
cause it is a pasture plant; but it may also partly be because
on the three grass species. Haplothrips larvae are excluded from the analysis. decisions regarding land management and community support
for such decisions largely occur in towns and cities, and many
indicating that the grass negatively impacts animals occurring of the known negative environmental impacts are on land ten-
in the zone between tussocks, not only on the tussocks ures typically away from built-up areas. We hope that demon-
themselves. strating a negative impact of buffel grass on biodiversity in an
Two aspects of our methodology are worth emphasising. urban setting shows that such impacts are likely systemic, con-
First, we surveyed individual plants of the target grass species, tributing to increased community awareness and support for
as we believe this approach makes it easier to establish links be- the long-term management of this highly invasive species.
tween insect communities and the plants on which they may
depend. This is especially important in floristically heteroge- Acknowledgements
neous landscapes where the cover of the target plant can be
The authors thank two anonymous reviewers for comments
low. Second, we suggest that identifying members of the target
insect group to a high level of taxonomic resolution (genus or on an earlier version of the article. Dianne Bell assisted with
species) provides greater context for conclusions regarding re- fieldwork. Carly Steen (NT Parks and Wildlife) produced the
source use and distributions in space and time, and means the base map in Figure 1.
investigation is more easily repeated.
Our results show that local native grasses provide important
Funding
habitat for local native insect species. We suggest that active re-
moval of buffel grass in Alice Springs and replacement with local This study was undertaken with the financial support of the
native grasses wherever possible would help reverse the loss of Northern Territory Department of Natural Resources,
biodiversity and normal ecological processes. Re-establishment Environment, Arts and Sport.The diversity of thrips (Insecta: Thysanoptera) on buffel grass | 7
Conflict of interest statement. The authors declare no conflict Mathers, N. J., Harms, B., and Dalal, R. C. (2006) ‘Impacts of
of interest. Land-Use Change on Nitrogen Status and Mineralisation in the
Mulga Lands of Southern Queensland’, Austral Ecology, 31: 708–18.
Moritz, G. B., and Mound, L. A. (1999) ‘AQIS Identification Guide –
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