Locusts and Grasshoppers: Behavior, Ecology, and Biogeography

Full text

Hindawi Publishing Corporation
Psyche
Volume 2011, Article ID 578327, 4pages
doi:10.1155/2011/578327
Editorial
Locusts and Grasshoppers: Behavior, Ecology, and Biogeography
Alexandre Latchininsky,1Gregory Sword,2, 3 Michael Sergeev,4, 5
Maria Marta Cigliano,6and Michel Lecoq7
1Department of Renewable Resources, University of Wyoming, 1000 E. University Avenue, Laramie, WY 82071, USA
2School of Biological Sciences, University of Sydney, Sydney, NSW 2006, Australia
3Department of Entomology, Faculty of Ecology and Evolutionary Biology, Heep Building, Texas A&M University, College Station,
TX 77842-2475, USA
4Department of General Biology and Ecology, Novosibirsk State University, 2 Pirogova Street, Novosibirsk 630090, Russia
5Laboratory of Insect Ecology, Institute of Systematics and Ecology of Animals, Siberian Branch, Russian Academy of Sciences,
11 Frunze Street, Novosibirsk 630091, Russia
6Division Entomologia, Museo de La Plata, Universidad Nacional de la Plata, Paseo del Bosque S/N,1900 La Plata, Argentina
7CIRAD Bioagresseurs, TA A-106/D, Campus International de Baillarguet, 34398 Montpellier cedex 5, France
Correspondence should be addressed to Alexandre Latchininsky, latchini@uwyo.edu
Received 27 January 2011; Accepted 27 January 2011
Copyright © 2011 Alexandre Latchininsky et al. This is an open access article distributed under the Creative Commons
Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is
properly cited.
Locusts and grasshoppers (L&G) (Orthoptera: Caelifera,
Acridoidea) are an essential component of both, healthy, and
disturbed grassland ecosystems. These insects are abundant
in natural and anthropogenic habitats (rangelands, wetlands,
agricultural fields, lawns, etc.). They stimulate plant growth,
participate in nutrient cycling, and play important role
in food chains [1–5]. Some grasshoppers are proposed as
ecological indicators of ecosystem qualities and efficacy of
ecological networks [6]. On the other hand, when their pop-
ulations grow to catastrophic dimensions, L&G are among
the most devastating enemies of agriculturists. Outbreaks
of locusts such as Schistocerca gregaria (Forsk˚
al, 1775),
Nomadacris septemfasciata (Serville, 1838), Locusta migra-
toria Linnaeus, 1758, Calliptamus italicus (Linnaeus, 1758),
Dociostaurus maroccanus (Thunberg, 1815), Chortoicetes
terminifera (Walker, 1870), and many abundant grasshopper
species continue to occur on all continents except Antarctica
and affect the livelihoods of one in every ten people on Earth.
Such L&G outbreaks are now better controlled and their
frequency and size have been reduced with the application
of preventative strategies [7,8]. However, invasions still
persist. During the outbreak of the Desert locust S. gregaria
in Africa in 2003–2005, over eight million people suffered
from severe 80 to 100% crop losses [9]. To combat the
locust swarms, 13 million hectares in 22 countries on three
continents were treated with broad-spectrum neurotoxins.
Such transcontinental operation, including the food aid for
affected population, cost over half a billion US dollars to the
world community [10].
Losses to L&G are not limited to crop and rangeland
destruction. Besides the economic damage and its subse-
quent negative social impact, L&G outbreaks may seriously
alter ecological processes across landscapes (e.g., carbon and
water cycles). The rapid loss of vegetation cover may result
in soil erosion and increased runoff.L&Gcanalsodestroy
food sources for many animals and thus affect biodiversity;
such effects may be particularly pronounced in isolated
insular ecosystems [11]. Large-scale L&G control programs
can also affect biodiversity, including that of nontarget
grasshoppers [12]. Despite decades of intensive research,
the mechanisms underlying L&G population dynamics (and
for locusts: phase transformation) are not fully elucidated.
Only recently, significant advances were made in our
understanding of L&G behavior and ecology, particularly
individual and group movement, nutritional requirements,
and biochemical mechanisms underlying the transformation
between solitarious and gregarious locust phases [13–15]; see
also review in [16].

2Psyche
Besides the notorious pests, this group of insects includes
many understated rare species which require protection [17–
19]. To complicate the picture, following landscape changes
induced by human agricultural activities, some economic
pests may become exceedingly rare [20]. On the other hand,
many orthopteran species benefit from human-induced
landscape changes and increase their abundance [18,21].
Disturbed and new habitats can be important for spreading
and living of some native and alien grasshopper forms
[18,21,22]. At the same time, many of rare grasshopper
species are threatened by anthropogenic influences, such as
overgrazing and ploughing [18]. However, in various areas,
such as temperate Eurasia or in Tropical Madagascar, several
centers of orthopteran diversity and endemism overlap
with areas of frequent L&G outbreaks [23–25]. This means
that problems of plant protection and conservation biology
should be solved on the complex basis of a holistic approach.
However, it is hardly ever the case; pests and rare species are
usually studied separately, and their possible relationships are
not explored.
Although the general patterns of grasshopper distribu-
tion are described for different regions [26–28], the main
factors and processes determining grasshopper diversity
patterns at different scales are still under discussion. Impor-
tance of temperatures and precipitation is evident, but the
distribution of many species, populations, and assemblages
could not be explained by macroclimatic factors only [29].
This means that the role of other factors and processes should
be investigated more thoroughly. At a regional level, it is
possible to establish the general pattern of regional biodiver-
sity and explain how the spatial distribution of populations
permits species with various origins and different ecological
preferences to coexist [30].
An example of this approach is the opening article for
this special issue of Psyche, in which M. G. Sergeev reviews
distribution patterns of over 130 species of grasshoppers
and their kin in the boreal zone. Grasshoppers and their
relatives occupy there almost exclusively open habitats, such
as meadows, mountain steppes and tundras, clearings, open-
ings, bogs, and stony flood plains. The boreal orthopteroid
assemblages exhibit low species diversity and abundance.
Based on the biogeographic analysis, the author concludes
that relationships between the faunas of the Eurasian and
North American parts of the boreal zone are relatively
weak.
Local grasshopper distribution patterns have been dis-
cussed since the beginning of the 20th century. Possible
relationships between grasshopper diversity, plant species
composition, and habitat structure have been discussed for
many decades. The paper of D. H. Branson (second in
this special issue) provides an example of such studies. The
author found these relationships too complicated for simple
explanations. The type, level, strength, and complexity of
these relationships may be determined not only by local but
also by regional patterns. Consequently, to evaluate general
trends in grasshopper diversity one should study all main
regions and ecosystems in the same manner. This idea may
serve as a basis for an ambitious regional study.
The third paper of the special issue is devoted to a
complex terminological issue. Acridologists have used a
variety of terms to describe groups of grasshoppers, includ-
ing assemblage, community, guild, and population. This
terminological diversity has raised the question of whether
one of these descriptors is the correct one. The author, J.
A. Lockwood, argues that a term is correct if it accurately
reflects the conceptual framework of the investigator and
effectively communicates this perspective to others. He
describes the contexts in which the most common terms are
appropriate.
In the next paper, O. Olfert et al. investigate the impact
of climate changes on distribution and relative abundance
of a pest grasshopper of major economic importance in
North America, Melanoplus sanguinipes. Various scenarios
of climatic changes were used to parameterize a bioclimatic
model of this species. Compared to predicted range and
distribution under current climate conditions, model results
indicated that M. sanguinipes would have increased range
and relative abundance in more northern regions of North
America. Conversely, model output predicted that the range
of this crop pest could contract in regions where climate
conditions became limiting. However, some caution has been
expressed by authors. The impact of biotic factors such as
natural enemies should also be considered, and bioclimatic
modeling of grasshopper populations will surely benefit
in the future from a multitrophic approach (host plants-
grasshoppers-natural enemies).
The fifth paper of this special issue by H. Song reviews the
current state-of-the-art regarding locust phase polyphenism
in species other than the two model locusts. Although the
mechanisms of locust phase transformation are relatively
well understood for the Desert locust and the Migratory
locust, they remain largely obscure in nonmodel locust
species. The author found similar density-dependent pheno-
typic plasticity among closely related species. He emphasized
the importance of comparative analyses in understanding the
evolution of locust phase and proposed a phylogeny-based
research framework for future analyses.
In the next paper M. Lecoq et al. present a typology quan-
tifying density-dependent color change in the Red locust
nymphs. This information can contribute to improving
the reliability of the data collected by the National Locust
Centers when surveying this major pest. The authors, in
Madagascar, sampled hoppers from several populations of
different density and measured the color of different body
parts as categorical variables. They found that color change
is positively correlated with population density. This study
is an important contribution to our knowledge of locust
coloration in the field, for which there is currently a weaker
understanding than that for laboratory populations.
The seventh paper of this special issue by S. O. Ely et al.
discusses the diel behavioral activity patterns of solitarious
Desert locust adults. The authors found that the insects
were more attracted to volatiles from potted Heliotropium
ovalifolium in scotophase than in photophase. The attraction
towards the host plant odors, in both photophase and
scotophase, concurs with previous observations on locust
oviposition preferences near these plants.

Psyche 3
Intheeighthpaper,R.B.SrygleyandS.T.Jaronskireport
experiments with Beauveria bassiana (Fungi: Ascomycota),
an entomopathogenic fungus that serves as a biological con-
trol agent of Mormon crickets Anabrus simplex Haldeman
(Orthoptera: Tettigoniidae) and other grasshopper pests.
They demonstrated an immune response of infected Mor-
mon crickets and concluded that circulating phenoloxidase
may be an important enzymatic defense against Beauveria
infection, and that it is associated with attempted clearing
of Beauveria blastospores and hyphae from Mormon cricket
hemolymph.
Alexandre Latchininsky
Gregory Sword
Michael Sergeev
Maria Marta Cigliano
Michel Lecoq
References
[1] I. V. Stebaev, “Periodic changes in the ecological distribution
of grasshoppers in the temperate and the extreme continental
steppe regions, andtheir importance forthe local ecosystems,”
in Proceedings of the International Study Conference on the
Current and Future Problems of Acridology, pp. 207–213,
Centre for Overseas Pest Research, London, UK, 1972.
[2] G. B. Hewitt and J. A. Onsager, “A method for forecasting
potential losses from grasshopper feeding on northern mixed
prairie forages,” Journal of Range Management, vol. 35, pp. 53–
57, 1982.
[3] O.O.OlfertandM.K.Mukerji,“Effects of acute simulated
and acute grasshopper (Orthoptera: Acrididae) damage on
growth rates and yield in spring wheat (Triticum aestivum),”
The Canadian Entomologist, vol. 115, no. 6, pp. 629–639, 1983.
[4] M. G. Sergeev, “Zonal-landscape distribution of Orthoptera
zoomassinMiddleRegionoftheUSSR,”Geographia i
Prirodnyje Resursy, no. 2, pp. 89–92, 1989 (Russian).
[5] G. E. Belovsky, “Do grasshoppers diminish grassland produc-
tivity? A new perspective for control based on conservation,”
in Grasshoppers and Grassland Health. Managing Grasshopper
Outbreaks without Risking Environmental Disaster,J.A.Lock-
wood,A.V.Latchininsky,andM.G.Sergeev,Eds.,pp.7–30,
Kluwer Academic Publishers, Dordrecht, Netherlands, 2000.
[6]C.S.Bazelet,Grasshopper bioindicators of effective large-scale
ecological networks, Ph.D. Dissertation, Department of Con-
servation Ecology and Entomology, Stellenbosch University,
South Africa, 2011.
[7] J.I.Magor,M.Lecoq,andD.M.Hunter,“Preventivecontrol
and Desert Locust plagues,” Crop Protection, vol. 27, no. 12,
pp. 1527–1533, 2008.
[8] G.A.Sword,M.Lecoq,andS.J.Simpson,“Phasepolyphenism
and preventative locust management,” Journal of Insect Physi-
ology, vol. 56, no. 8, pp. 949–957, 2010.
[9] L.Brader,H.Djibo,andF.G.Faye,Towards a more Effective
Response to Desert Locusts and Their Impacts on Food Insecurity,
Livelihoods and Poverty. Independent Multilateral Evaluation
of the 2003–2005 Desert Locust Campaign,FAO,Rome,Italy,
2005.
[10] Y. T. Belayneh, “Acridid pest management in the developing
world: a challenge to the rural population, a dilemma to the
international community,” Journal of Orthoptera Research,vol.
14, no. 2, pp. 187–195, 2005.
[11] A. V. Latchininsky, “Grasshopper outbreak challenges conser-
vation status of a small Hawaiian Island,” Journal of Insect
Conservation, vol. 12, no. 3-4, pp. 343–357, 2008.
[12] M. J. Samways, “Can locust control be compatible with con-
serving biodiversity?” in Grasshoppers and Grassland Health.
Managing Grasshopper Outbreaks without Risking Environ-
mental Disaster,J.A.Lockwood,A.V.Latchininsky,andM.
G. Sergeev, Eds., pp. 173–180, Kluwer Academic Publishers,
Dordrecht, Netherlands, 2000.
[13] J. Buhl, D. J. T. Sumpter, I. D. Couzin et al., “From disorder
to order in marching locusts,” Science, vol. 312, no. 5778, pp.
1402–1406, 2006.
[14] M. L. Anstey, S. M. Rogers, S. R. Ott, M. Burrows, and
S. J. Simpson, “Serotonin mediates behavioral gregarization
underlying swarm formation in desert locusts,” Science,vol.
323, no. 5914, pp. 627–630, 2009.
[15] D. A. Cullen, G. A. Sword, T. Dodgson, and S. J. Simpson,
“Behavioural phase change in the Australian plague locust,
Chortoicetes terminifera, is triggered by tactile stimulation of
the antennae,” Journal of Insect Physiology,vol.56,no.8,pp.
937–942, 2010.
[16] M. P. Pener and S. J. Simpson, “Locust phase polyphenism: an
update,” Advances in Insect Physiology, vol. 36, pp. 1–272, 2009.
[17] M. J. Samways and J. A. Lockwood, “Orthoptera conservation:
pests and paradoxes,” Journal of Insect Conservation,vol.2,no.
3-4, pp. 143–149, 1998.
[18] M. G. Sergeev, “Conservation of orthopteran biological diver-
sity relative to landscape change in temperate Eurasia,” Journal
of Insect Conservation, vol. 2, no. 3-4, pp. 247–252, 1998.
[19] A. Foucart and M. Lecoq, “Major threats to a protected
grasshopper, Prionotropis hystrix rhodanica (Orthoptera, Pam-
phagidae, Akicerinae), endemic to southern France,” Journal of
Insect Conservation, vol. 2, no. 3-4, pp. 187–193, 1998.
[20] A. V. Latchininsky, “Moroccan locust Dociostaurus maroccanus
(Thunberg, 1815): a faunistic rarity or an important economic
pest?” Journal of Insect Conservation, vol. 2, no. 3-4, pp. 167–
178, 1998.
[21] M. G. Sergeev, O. V. Denisova, and I. A. Vanjkova, “How
do spatial population structures affect acridid management?”
in Grasshoppers and Grassland Health. Managing Grasshopper
Outbreaks without Risking Environmental Disaster,J.A.Lock-
wood,A.V.Latchininsky,andM.G.Sergeev,Eds.,pp.71–88,
Kluwer Academic Publishers, Dordrecht, Netherlands, 2000.
[22] M. J. Samways and M. G. Sergeev, “Orthoptera and landscape
change,” in The Bionomics of Grasshoppers, Katydids and
Their Kin,S.K.Gangwere,M.C.Muralirangan,andM.
Muralirangan, Eds., pp. 147–162, CAB International, Oxon,
UK, 1997.
[23] M. G. Sergeev, “La s´
echeresse et les sch´
emas de distribution
des criquets en Asie centrale et septentrionale,” S´echeresse,vol.
7, no. 2, pp. 129–132, 1996.
[24] J. A. Lockwood and M. G. Sergeev, “Comparative biogeog-
raphy of grasshoppers (Orthoptera: Acrididae) in North
America and Siberia: applications to the conservation of
biodiversity,” Journal of Insect Conservation,vol.4,no.3,pp.
161–172, 2000.
[25] R. Peveling, “Environmental conservation and locust
control—possible conflicts and solutions,” Journal of Orthop-
tera Research, vol. 10, no. 2, pp. 171–187, 2001.
[26] B. P. Uvarov, Grasshoppers and Locusts,vol.2,Centrefor
Overseas Pest Research, London, UK, 1977.
[27] D. Otte, The North American Grasshoppers—vol. 1. Acrididae:
Gomphocerinae and Acridinae, Harvard University Press,
Cambridge, Miss, USA, 1981.

4Psyche
[28] M. G. Sergeev, Principles of Orthopteroid Insects Distribution in
North Asia, Nauka Publishers, Novosibirsk, Russia, 1986.
[29] K. A. Vandyke, A. V. Latchininsky, and S. P. Schell, ‘Importance
of ecological scale in Montane Grasshopper (Orthoptera:
Acrididae) species structure in similar habitat between differ-
ing soil textures and dominant vegetative canopy coverage,”
Journal of Orthoptera Research, vol. 18, no. 2, pp. 215–223,
2009.
[30] M. G. Sergeev, “Ecogeographical distribution of Orthoptera,”
in The Bionomics of Grasshoppers, Katydids and Their Kin,S.K.
Gangwere, M. C. Muralirangan, and M. Muralirangan, Eds.,
pp. 129–146, CAB International, Oxon, UK, 1997.