227
Chapter 10
Tallgrass Prairie, Ground Beetles (Coleoptera: Carabidae),
and the Use of Fire as a Biodiversity and
Conservation Management Tool
Robert E. Roughley1, Darren A. Pollock2, and David J. Wade
Department of Entomology, University of Manitoba
Winnipeg, Manitoba, Canada R3T 2N2
Abstract. Fire is a key factor that structures the plant and animal communities of tallgrass prairie. Manipulating
the timing of fires provides a potential method to influence the diversity of these communities. We examined this
potential with a four-year study of ground beetles (Coleoptera: Carabidae) on tallgrass prairie in southern Manitoba.
Using pitfall traps, we compared captures of beetles from 1997 to 2000 in control plots of unburned prairie or in
plots burned in the spring, summer, or fall of 1997. Results showed that species responded individually to different
treatments but that treatment did not alter the composition of the ground beetle community. Further, at least four
years appeared necessary for the beetle community to revert back to its pre-burn status. Thus, for ground beetles, a
mosaic of burn regimes and burn seasons is likely needed to sustain habitat heterogeneity and species diversity.
Résumé. Le feu est un facteur clé de la structure des communautés végétales et animales de la prairie à herbes
hautes. La gestion du calendrier des brûlis peut constituer un moyen d’influer sur la diversité de ces communautés.
Nous examinons cette possibilité dans le cadre d’une étude des carabidés (Coleoptera : Carabidae) de la prairie à
herbes hautes du sud du Manitoba étalée sur quatre ans. Nous comparons les captures de coléoptères effectuées de
1997 à 2000 à l’aide de pièges à fosse dans des parcelles témoins de prairies non brûlées et de parcelles brûlées au
printemps, en été ou en automne, en 1997. Les résultats montrent que chaque espèce réagit à sa manière aux divers
traitements, mais que ces derniers n’influent pas sur la composition de la communauté de carabidés. De plus,
il semble qu’il faille au moins quatre ans à la communauté des coléoptères pour revenir à son état antérieur au
traitement. Ainsi, en ce qui concerne les carabidés, il semble qu’une mosaïque de régimes de brûlis soit nécessaire
pour maintenir l’hétérogénéité de l’habitat et la diversité des espèces.
Introduction
With less than 1% remaining intact, tallgrass prairie is one of most endangered habitats in
Canada (Trottier 1992). Isolated patches are now known only from southeastern Manitoba
and southern Ontario (Trottier 1992). There is thus interest in methods that can be used to
maintain the biodiversity and ecosystem processes of these tallgrass remnants.
Disturbance is one factor that maintains biodiversity (Petraitis et al. 1989) and fire is a
key disturbance in tallgrass ecosystems. The response of fire in tallgrass systems has been
studied for small mammals (Kaufman et al. 1989; Clark and Kaufman 1990), large herbivores
(Vinton et al. 1993; Collins et al. 1998), earthworms (James 1982, 1988), grasshoppers (Evans
1984, 1988), spiders (see Chapter 11), general insect communities (Rice 1932), and selected
1
Deceased.
2
Present address: Eastern New Mexico University, Portales, New Mexico, 88130, USA
Roughley, R. E., D. A. Pollock, and D. J. Wade. 2010. Tallgrass Prairie, Ground Beetles (Coleoptera: Carabidae),
and the Use of Fire as a Biodiversity and Conservation Management Tool. In Arthropods of Canadian Grasslands
(Volume 1): Ecology and Interactions in Grassland Habitats. Edited by J. D. Shorthouse and K. D. Floate.
Biological Survey of Canada. pp. 227-235. © 2010 Biological Survey of Canada. ISBN 978-0-9689321-4-8
doi:10.3752/9780968932148.ch10
228
R. E. Roughley, D. A. Pollock, and D. J. Wade
communities of other organisms (Rice 1932; Collins and Glenn 1997). Many of these studies
also examined the effects of fire on plant taxa (e.g., Hadley and Kieckhefer 1963; Peet et al.
1975; Evans 1984; Pyne 1986; Abrams 1987; Hulbert 1988; Anderson 1990; Collins 1990;
Collins and Gibson 1990; Howe 1999; Sveinson 2001).
Ground beetles (Coleoptera: Carabidae) are a particularly useful taxon with which to
study the effects of fire on biodiversity. Taxonomic keys exist for their identification (e.g.,
Lindroth 1960, 1963, 1966, 1968, 1969a, 1969b; Bousquet 1991; Bousquet and Larochelle
1993) and, for many species, biogeographical and ecological information is available (e.g.,
Larochelle 1990; Larochelle and Larivière 2003). Further, ground beetles are easily collected
by passive methods such as pitfall traps and have high species richness; 350 species have
been recorded for Manitoba (Bousquet 1991). Finally, as a taxon, ground beetles inhabit a
variety of ecological niches (herbivores, predators, scavengers, parasitoids) that make their
numbers more responsive to changes in their environment (e.g., Niemelä et al. 1992; Niwa
and Peck 2002).
A remnant patch of tallgrass prairie at the St. Charles Rifle Range (SCRR), near
Winnipeg, Manitoba, has been used to examine the manipulation of fire as a tool to
maintain the biodiversity of plant (Sveinson 2001) and arthropod communities, that is,
ground beetles (this chapter) and spiders (Araneae; see Chapter 11). The 192 ha property
is owned by the Canadian Department of National Defence. This area includes 47.9 ha
of high-quality tallgrass prairie that has remained undisturbed for at least 100 years. An
adjacent piece of the property was last cultivated about 50 years ago (Morgan 1994).
The fire studies at SCRR have relied on the establishment of four replicate blocks of
five plots (50 × 50 m per plot), each in the form of a “+” (Fig. 1). The effect of fire season
was assessed after one plot in each block was burned in early June (spring), early August
(summer), or early September (fall) in 1997. A fourth central plot in each block remained
unburned as a control in arthropod studies. The fifth plot in each block, also unburned, was
used in botanical analyses (Sveinson 2001).
In the study on ground beetles at SCRR, we examined the effects of fire by using pitfall
traps (16 traps per plot) positioned in each of the four plot treatments (spring, summer, fall,
control) per block. Traps were emptied weekly and beetles pooled within plots for each
week. Trap periods varied across years as follows: 28 May to 7 November 1997; 3 April to
10 November 1998; 20 April to 12 November 1999; and 31 March to 10 November 2000.
Fire Effects on Ground Beetle Communities
The four-year study yielded 92 species and 11,364 specimens of ground beetles (Table 1).
A further 11 species were recovered during this period, but elsewhere at SCRR during the
course of an unrelated study (Roughley et al. 2006). Collectively, the 103 species represents
29% of the total number of ground beetles species known from Manitoba (Bousquet and
Larochelle 1993) and attests to the richness of this taxon (Carabidae) at the site.
Regarding the structure of ground beetle communities, the SCRR study suggests
that fire has little apparent effect. In comparisons among treatments within years, no
differences were observed in measures of log abundance or abundance per week, nor
was there a consistent pattern in the abundance levels among treatments from year to
year. Similarly, analysis with a number of measures (log species richness, ShannonWiener, log series alpha indices, Simpson index, and Berger-Parker index) generally
showed no differences in species diversity detected among treatments for any year.
The apparent lack of a burn season effect (spring vs. summer vs. fall) on the ground beetle
Effect of Fire on Ground Beetles in Tallgrass Prairie
229
community at SCRR might reflect the relatively small plot size of 50 × 50 m. For example,
indirect evidence shows that species moved among treatment plots. For collections of beetles
combined across years and treatments, the maximum number of species recovered from an
individual block was 58. However, 92 species were recovered when these collections were
combined across blocks. Regarding this species richness, 45 species of ground beetles were
recovered from a reconstructed tallgrass prairie remnant in Iowa (Larsen and Williams 1999).
Of the most abundant species listed from this site, only Poecilus lucublandus lucublandus
(Say) was common to the results at SCRR. This difference reinforces the observation of
Collins (2000) on the geographical variability of tallgrass communities.
From the qualitative Jaccard similarity index, species composition was most similar in
1997 between fall burn and control treatments. This was expected because the fall burn was
applied near the end of the first field season when beetle activity was expected to be low.
From the quantitative Morisita-Horn similarity index, species composition was most similar
in 1997 between the summer and fall burn treatment, and then between the spring burn and
control treatments. A similar result was observed for the spider community (see Chapter 11).
The richness and abundance of spiders and ground beetles peak in the spring, with many
species also having high activity levels in the fall. The timing of the burn treatments may be
important, as certain species will be more vulnerable at specific times during the year.
In summary, however, the variable response to burn treatment confounded the detection
of treatment effects in the community analysis.
Fig. 1. Detail of experimental set-up at the St. Charles Rifle Range. Blocks (A, B, C, D) comprised five plots:
spr = spring burn; sum = summer burn; fall = fall burn; ref = unburned control plot; the fifth plot was used for a
botanical study only. The inset shows the arrangement of pitfall traps with plots.
230
R. E. Roughley, D. A. Pollock, and D. J. Wade
Table 1. Ground beetles species and subspecies (Coleoptera: Carabidae) found at the St. Charles Rifle Range
(SCRR), Winnipeg, Manitoba (1997–2000) (classification follows Bousquet and Larochelle 1993).
Notiophilini
1. Notiophilus aquaticus (Linnaeus, 1758)
2. Notiophilus semistriatus Say, 1823
Cicindelini
3. Cicindela longilabris longilabris Say, 1824
Carabini
4. Calosoma calidum (Fabricius, 1775)
5. Carabus granulatus granulatus Linné, 1758
6. Carabus maeander Fischer von Waldheim, 1820
Elaphrini
7. *Blethisa multipunctata aurata Fischer von
Waldheim, 1828
8. *Elaphrus clairvillei Kirby, 1837
Clivinini
9. Dyschirius globulosus (Say, 1823)
10. Dyschirius integer LeConte, 1852
Bembidiini
11. Bembidion nitidum (Kirby, 1837)
12. Bembidion nudipenne Lindroth, 1963
13. Bembidion graphicum Casey, 1918
14. Bembidion patruele Dejean, 1831
15. Bembidion rapidum (LeConte, 1848)
16. Bembidion mimus Hayward, 1897
17. Bembidion timidum (LeConte, 1848)
18. Bembidion versicolor (LeConte, 1848)
19. Bembidion mutatum Gemminger and Harold, 1868
20. Bembidion quadrimaculatum oppositum Say, 1823
21. Bembidion transparens (Gebler, 1829)
22. Bembidion canadianum Casey, 1924
23. *Bembidion concretum Casey, 1918
24. Bembidion fortestriatum (Motschulsky, 1845)
25. Elaphropus incurvus (Say, 1830)
Patrobini
26. Patrobus lecontei Chaudoir, 1871
27. Patrobus septentrionis Dejean, 1828
28. Patrobus stygicus Chaudoir, 1871
Pterostichini
29. Poecilus corvus (LeConte, 1873)
30. Poecilus lucublandus lucublandus (Say, 1823)
31. Pterostichus commutabilis (Motschulsky, 1866)
32. Pterostichus femoralis (Kirby, 1837)
33. Pterostichus pensylvanicus LeConte, 1873
34. Pterostichus corvinus (Dejean, 1828)
35. †Pterostichus melanarius (Illiger, 1798)
36. Pterostichus caudicalis (Say, 1823)
Zabrini
37. Amara carinata (LeConte, 1848)
38. †Amara apricaria (Paykull, 1790)
39. Amara latior (Kirby, 1837)
40. Amara obesa (Say, 1823)
41. Amara lunicollis Schiødte, 1837
42. Amara cupreolata Putzeys, 1866
43. Amara musculis (Say, 1823)
44. *Amara angustata (Say, 1823)
45. Amara sp.
Chlaeniini
46. Chlaenius sericeus sericeus (Forster, 1771)
47. Chlaenius purpuricollis purpuricollis Randall,
1838
48. Chlaenius pennsylvanica pennsylvanicus Say,
1823
Licini
49. Diplocheila striatopunctata (LeConte, 1844)
50. Badister neopulchellus Lindroth, 1954
51. *Badister obtusus LeConte, 1878
52. *Badister parviceps Ball, 1959
53. Badister transversus Casey, 1920
Harpalini
54. Anisodactylus harrisii LeConte, 1863
55. Anisodactylus rusticus (Say, 1823)
56. Anisodactylus sanctaecrucis (Fabricius, 1798)
57. Stenolophus fuliginosus Dejean, 1829
58. Stenolophus comma (Fabricius, 1775)
59. Stenolophus conjunctus (Say, 1823)
60. Bradycellus lecontei Csiki, 1932
61. Bradycellus congener (LeConte, 1848)
62. Bradycellus neglectus (LeConte, 1848)
63. *Bradycellus nigriceps LeConte, 1868
64. Bradycellus semipubescens Lindroth, 1968
65. *Bradycellus lugubris (LeConte, 1848)
66. Trichocellus cognatus (Gyllenhal, 1827)
67. Acupalpus canadensis Casey, 1924
68. Acupalpus carus (LeConte, 1863)
69. Acupalpus nanellus Casey, 1914
70. Harpalus pensylvanicus (DeGeer, 1774)
71. Harpalus fraternus LeConte, 1852
72. Harpalus indigens Casey, 1924
73. Harpalus ventralis LeConte, 1848
74. Harpalus ochropus Kirby, 1837
75. Harpalus plenalis Casey, 1914
76. Harpalus opacipennis (Haldeman, 1843)
77. *Harpalus amputatus Say, 1830
78. Harpalus herbivagus Say, 1823
79. Harpalus somnulentus Dejean, 1829
80. Selenophorus opalinus (LeConte, 1863)
81. *Selenophorus planipennis LeConte, 1848)
Platynini
82. Synuchus impunctatus (Say, 1823)
83. Agonum gratiosum (Mannerheim, 1853)
84. Agonum lutulentum (LeConte, 1854)
Effect of Fire on Ground Beetles in Tallgrass Prairie
231
Table 1. (continued)
85. *Agonum sordens Kirby, 1837
86. Agonum thoreyi Dejean, 1828
87. Agonum affine Kirby, 1837
88. Agonum corvus (LeConte, 1860)
89. Agonum cupreum Dejean, 1831
90. Agonum cupripenne (Kirby, 1823)
91. Agonum melanarium Dejean, 1828
92. Agonum placidum (Say, 1823)
93. *Agonum propinquum (G. and H., 1868)
94. Platynus decentis (Say, 1823)
Lebiini
95. Cymindis borealis LeConte, 1863
96. Cymindis cribricollis Dejean, 1831
97. Cymindis neglectus Haldeman, 1843
98. Syntomus americanus (Dejean, 1831)
99. Lebia atriventris Say, 1823
100. Lebia divisa LeConte, 1850
101. Lebia fuscata Dejean, 1825
102. Lebia vittata (Fabricius, 1777)
103. Lebia moesta LeConte, 1850
*Species found only in habitats other than tallgrass prairie on SCRR. † Species introduced into North America.
Fire Effects on Individual Species
Effects on individual species of ground beetles were detected. This point is illustrated with
reference to Syntomus americanus (Dejean), Agonum cupreum Dejean, Calosoma calidum
(Fabricius), Poecilus lucublandus lucublandus (Say), Pterostichus melanarius (Illiger),
Bradycellus congener (LeConte), and Lebia divisa LeConte.
Syntomus americanus is a small black native species with widespread distribution
in North America. It is characterized as “a sun-loving species, occurring on sandy rarely
peaty soil with sparse, low vegetation” (Lindroth 1969b: 1056). Larochelle and Larivière
(2003) stated that this species is found on open ground and is active in the sunshine. On
SCRR, S. americanus was one of the more abundant species in most years and in most burn
treatments. Adults of this species responded quickly and effectively to a spring burn, with
high adult abundance that was most apparent immediately after the burn treatment in 1997.
By 2000, the abundance had returned to pre-burn levels. The increased numbers following
the burn treatments were likely more of a response to an opening of the vegetation with
increased sun exposure, rather than a response to the actual burning itself.
Agonum cupreum is widespread in North America with records predominantly from
western localities, that is, Northwest Territories and British Columbia, south to Arizona,
and east to Michigan and New Brunswick (Bousquet and Larochelle 1993). This species
has the same general habitat description for adults as given for S. americanus (Lindroth
1969b; Frank 1971) and is also a diurnal carabid that is active in sunshine (Larochelle
and Larivière 2003). It is also stated to be “clearly effective as a colonist” and “strongly
favoured by human activities” (Larochelle and Larivière 2003: 21). Agonum cupreum
was collected throughout almost all of the growing season, with peak activity in May.
It was most abundant in the spring burn plot in 2007, being less often collected in other
combinations of treatments and subsequent years.
The natural history and habitat preferences of C. calidum are described by Frank (1971),
Lindroth (1960), and Larochelle and Larivière (2003). With broad distribution in North
America, this species prefers open ground with dry soil, with or without vegetation. Adults
of the species are diurnal. Most individuals at SCRR were collected in June; abundance
dropped precipitously through July and August, with no specimens collected after August.
This pattern indicates that C. calidum is a spring-active species, although Larochelle and
Larivière (2003) note only that the species is active from April to December. Calosoma
calidum was among the 10 most abundant species for most treatments in 1998, but it was
among the top 10 only in spring and summer burn treatments of 2000.
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R. E. Roughley, D. A. Pollock, and D. J. Wade
Introduced and widespread species, such as P. melanarius, are a concern for the
management of tallgrass prairie. This species was introduced on both coasts of North
America and has since been rapidly spreading inland (Lindroth 1966). The earliest known
record for Manitoba is 1956 at Selkirk (40 km NW of SCRR). Individual specimens have
been collected from an amazing variety of habitat types (Larochelle and Larivière 2003) and
are almost ubiquitous in habitats altered by human activities (towns and cities, agricultural
areas, etc.). Indeed, Larochelle and Larivière (2003: 426) stated that P. melanarius is
“strongly favoured by human activities.” For example, it accounted for over 40% of the
total ground beetles collected in an agricultural ecosystem in Alberta (Cárcamo 1995). The
presence of P. melanarius in high numbers on tallgrass prairie may be cause for concern, as
this large species has been shown to be competitively superior to native species of ground
beetles of approximately the same body size (e.g., Currie et al. 1996).
Because of the invasive nature of this species and its preference for disturbed habitats,
we monitored its populations in detail. North American adults of P. melanarius are
dimorphic for wing length. In Europe, where populations are relatively stable, the longwinged form is rare (Larochelle and Larivière 2003). Lindroth (1966) hypothesized that
the prevalence of macropterous individuals in North America was evidence of a recent
introduction and active range expansion.
The population of P. melanarius on SCRR during 1997 was relatively low (about 3%
of the total collection). However, a year after the controlled burning, the numbers increased
to about 8%. This species became one of the 10 most abundant species in the spring and
summer treatments in the year after the burn treatment, but numbers subsequently decreased.
In 1997, about 42% of adult P. melanarius were short winged. In 1998, this number had
increased to 76%. The introduction of non-native or adventive species that are also invasive
may threaten the diversity of native communities. Only 2 of the 103 species of ground
beetles known from SCRR are adventive, P. melanarius and Amara apricaria (Paykull).
This suggests that the tallgrass prairie ecosystem and its elements are apparently able to
exclude many of these weedy species during the successional pathway after burning.
Poecilus lucublandus lucublandus is widespread in North America (Bousquet and
Larochelle 1993) in habitats characterized as “open, moderately dry ground with vegetation
of grasses” (Lindroth 1966: 483). These habitats include gardens, lawns, lakeshores,
gravel pits, and forest edges and clearings (Larochelle and Larivière 2003). On SCRR,
this species had a bimodal pattern of annual abundance of adults. Captures were highest
in spring, lowest in summer, and then high again in autumn. This species was dominant
in all burn treatments and control plots from 1998 to 2000. As Pterostichus lucublandus,
it was unimodal, with a peak in July in an aspen control site within the Interlake region of
Manitoba (Holliday 1991).
Bradycellus congener is a widespread species (Bousquet and Larochelle 1993) whose
adults are found on “clayish, rather dry soil, usually near water, for example, saline ponds in
the prairie, but not confined to such places” (Lindroth 1968: 888). Most adults are collected
after mid-June, are active at low temperatures, and are often active under snow (Larochelle
and Larivière 2003). On SCRR, adults were present in the late autumn, with the majority
of specimens collected in October and November of each sampling season, often under
significant snow cover.
In contrast to the preceding species, L. divisa has a more restricted distribution and may
be a northern grassland specialist of Great Basin grasslands. Members of the genus Lebia
are ectoparasitic as larvae on pre-pupae or pupae of chrysomelid beetles. The host or hosts
of L. divisa are not known (Madge 1967). Because of the apparent rarity of this species
Effect of Fire on Ground Beetles in Tallgrass Prairie
233
and the spotty distribution of collection records, neither Madge (1967) nor Larochelle and
Larivière (2003) provided a habitat preference. However, the latter stated that specimens
were collected on weeds at the edge of a slough. Lebia divisa may be a grassland specialist
that does well in tallgrass prairie. It was abundant only in the fall burn treatment of 1998.
More specimens of this species were collected in our study than were known from all
previous museum collections combined (Madge 1967; DAP, unpublished data).
Conclusions
Disturbance is required to maintain the native diversity and function of tallgrass prairie
(Trottier 1992; Morgan 1994). Fire is a key form of disturbance, which reduces the
encroachment of shrubs and trees onto grassland habitats. A series of aerial photographs
from 1928 to 1991 document that, in the absence of fire, aspen had increased its coverage
at SCRR by about 41% (Morgan 1994).
As a management tool, fire can be manipulated by the frequency of burns within
and among years, and/or by season (e.g., spring vs. fall). The decision to vary either fire
frequency (Collins 2000) or fire season will depend on various properties of the community,
including its stability and resilience. Higher stability will allow the community to resist
disturbances and maintain its functionality. Higher resilience will allow the community to
return to its former state quickly after a disturbance. If any single kind of disturbance, such
as fire, were to become the predominant management technique, then the communities
would be selected for their resilience to the effects of that disturbance (see Chapter 11). For
ground beetles, results from SCRR suggest that at least four years are needed for the beetle
community to revert back to its pre-burn state.
In the management of tallgrass prairie, adherence to a single burn season (e.g., spring
burn only) is unlikely to maximize biodiversity, as evidenced by the results for ground
beetles. Similarly, Sveinson (2001) showed that, relative to summer or fall burns, spring
burns favoured production of woody plants over forbs and grasses. A mosaic of burn
regimes and burn seasons to sustain habitat heterogeneity with a site likely provides the
best alternative to maintain species diversity.
Acknowledgements
A project of this magnitude would not have been possible without critical funding from the
Department of National Defence, State of Wisconsin Department of Natural Resources, and
a Natural Sciences and Engineering Research Council of Canada scholarship (DJW) and
operating/discovery grant (RER). The patience and understanding of our main contacts at
the SCRR—Drew Craig and Scott Edwards—was very important to this research project.
Summer research assistants, graduate students, and other researchers (Patrice Bouchard,
Jason Diehl, Markus Dyck, Dan O’Brian, Terry Galloway, Ginger Gill, Don Henne,
Wendy Graham, Dave Holder, Jackie LeGal, Steve Mazurat, Stuart McMillan, Steve
Offman, Lydia Stepanovich, Stacie Stiege, Heather White, Carla Wytrykush, and Debra
Wytrykush) provided much of the labour of sampling and sample preparation. The powers
of organization, assistance, and ingenuity of the departmental technicians, Dave Holder
and Lisa Babey, were often called upon to assist with this project. The ideas and concepts
of five former graduate students associated with the Department of Entomology—Jason
Diehl, Don Henne, Brent Elliott, Carla Wytrykush, and Rhéal Lafrenière—helped
crystallize many of our ideas about disturbance, tallgrass prairie, and ecosystems.
234
R. E. Roughley, D. A. Pollock, and D. J. Wade
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