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363
D.A. Yee (ed.), Ecology, Systematics, and the Natural History of Predaceous
Diving Beetles (Coleoptera: Dytiscidae), DOI 10.1007/978-94-017-9109-0_8,
© Springer Science+Business Media B.V. 2014
Abstract As conspicuous predators throughout ontogeny, dytiscids are central to
freshwater food webs, particularly in lentic systems such as wetlands and ponds.
Adult and larval dytiscids are considered to be generalists, feeding on zooplankton,
aquatic invertebrates, larval amphibians, and fi sh, but some dytiscid species selec-
tively feed on certain prey types relative to others. Selective predation, cannibalism,
intraguild predation, and non-consumptive effects on prey are attributes of dytiscid
feeding that are known to shape food web structure and composition and infl uence
species coexistence. Larval and adult dytiscids are also predators of mosquito larvae
and thus frequently investigated as potential agents for mosquito suppression,
particularly in northern areas and in areas where mosquitoes vector diseases. The
effects of dytiscid predation on food webs and mosquito populations are dependent
on several abiotic and biotic conditions, including vegetation structure, habitat
Chapter 8
Predator-Prey Interactions of Dytiscids
Lauren E. Culler , Shin-ya Ohba , and Patrick Crumrine
L. E. Culler (*)
Dartmouth College , Hanover , NH , USA
e-mail: leculler@gmail.com
S.-y. Ohba
Nagasaki University , Nagasaki , Japan
e-mail: ooba@nagasaki-u.ac.jp
P. Crumrine
Rowan University , Glassboro , NJ , USA
e-mail: crumrine@rowan.edu
With creamy margined, bronze green wing covers, oarlike hind
legs fringed with chestnut-colored hairs, and a pair of
formidable, meat-tong mandibles, what a well-fashioned
submarine predator the diving beetle is.
(Wayne H. McAlsiter
2004 )
364
complexity, and temperature. Dytiscids are also food for other organisms. Odonate
nymphs, fi sh, amphibians, reptiles, birds, and mammals are known predators of
dytiscids, although the extent to which these organisms rely on dytiscids for food
remains unclear. Given the prominent role of dytiscids in freshwater food webs,
future research should be aimed at improving basic knowledge of dytiscid feeding
ecology, using dytiscids to test predator-prey and trophic theory, and examining
how environmental change affects the role of dytiscids as predators of vector and
nuisance species.
Keywords Predation • Trophic ecology • Community structure • Cannibalism
• Biological control • Mosquitoes • Predator-prey interactions • Non-consumptive
effects
8.1 Introduction
Predation is an important component of aquatic systems and plays a critical role in
structuring communities (Batzer and Wissinger 1996 ; Batzer and Sharitz 2006 ) via
consumptive and non-consumptive effects on prey. Knowing the direction and
intensity of predation is vital for understanding the processes that structure
communities (Klecka and Boukal 2012 ) and for cascading predator effects on other
ecosystem characteristics (e.g., secondary production). Cannibalism and intraguild
predation, special cases of predation that are prevalent in aquatic systems, can
further infl uence community structure through density-dependent and size-structure
effects and can help to explain species diversity (Yee 2010 ). Although fi sh and
odonates as predators have received a great deal of attention (e.g., Crowder and
Cooper 1982 ; Gillinsky 1984 ; Mallory et al. 1994 ; Batzer et al. 2000 ; Crumrine
et al. 2008 ), dytiscids are ubiquitous predators in most freshwater habitats (Bay 1974 ).
Dytiscids exert strong top-down impacts on prey assemblages and affect other
ecological attributes of aquatic food webs.
All dytiscid beetles are carnivorous for at least part of their life-cycle. Larvae are
exclusively predaceous, whereas adults may also feed as scavengers (Johnson and
Jakinovich 1970 ; Larson et al. 2000 ; Bofi ll 2014 ). Detection of prey by dytiscids is via
visual (Maksimovic et al. 2011 ), tactile (Friis et al. 2003 ), or chemical (Formanowicz
1987 ) cues and varies between species, life stages, and habitats (Michel and Adams
2009 ). Adults dytiscids are clumsy and inept at capturing active prey (Larson et al.
2000 ) but larvae use a variety of hunting modes, including sit-and-wait and active hunting
(Yee 2010 ), and can be broadly classifi ed as swimmers, fl oaters, and crawlers (Wichard
et al. 2002 ). Larvae of many of the larger dytiscids, such as in the genus Dytiscus ,
are swimmers that pursue their prey by ambushing and trapping it against vegetation
or the water’s surface (Wichard et al. 2002 ). Floating larvae (e.g., Graphoderus ,
Acilius ) are more specialized swimmers that move elegantly through open water
and thus are more active during hunting (Wichard et al.
2002 ). Crawlers, including
larvae in the Hydroporine group, tend to be broad bodied and cling to vegetation and
L.E. Culler et al.
365
sediment rather than pursuing prey by swimming (Wichard et al. 2002 ). Once
detected and encountered, larval dytiscids grasp their prey with falcate piercing-
sucking mandibles. They pre-orally inject digestive protease enzymes that liquefy
their prey’s body contents and then proceed to suck the resulting mixture back up
through their mandibles for ingestion (Young 1967 ; Formanowicz 1987 ). This type
of feeding permits tackling prey items that may be quite large, including vertebrates
like fi sh (Fig. 8.1 ) and the tadpoles of bullfrogs and toads (Fig. 8.2 , Smith and
Awan 2009 ). Adults have chewing mouthparts like those of other Coleoptera and
thus are less effi cacious and more gape limited than their larval counterparts.
The diet of any predator is determined by its ability to detect, encounter, attack,
capture, subdue, and digest the various types of prey in its habitat. Any one of these
behavioral interactions may limit a predator’s ability to successfully consume individuals
of a given prey species. For example, prey of a given species may be consumed
because they are more abundant relative to other species in the habitat and therefore
encountered most often. Alternatively, the most abundant prey might be diffi cult for
a predator to successfully capture so alternative prey are pursued. For adult dytiscids,
gut contents can be discerned by dissection of the foregut and inspection of the
contents using a microscope (see Bosi 2001 ; Kehl and Dettner 2003 ). As with any
examination of gut contents, care must be taken in interpreting the results as some
of the material could have been ingested via the guts of other prey organisms (Kehl
and Dettner 2003 ) or could have been accidentally ingested. Visual examination of
gut contents is not possible for dytiscid larvae because the prey are liquefi ed during
ingestion. Polyacrylamide gel electrophoresis has been used to asses gut contents of
Fig. 8.1 Larval Dytiscus
sp. sinking its mandibles
into a small fi sh in a
laboratory aquarium
(Photo courtesy of
Siegfried Kehl 2012)
8 Predator-Prey Interactions of Dytiscids
366
other piercing-sucking predators, such as notonectids (Giller 1984 , 1986 ), and could
be used for studying the diets of larval dytiscids. Laboratory feeding experiments
and preference trials are therefore a common way of assessing what larval and
adult dytiscids consume. Individuals are offered different types of prey in different
proportions and if consumption deviates from the offered proportion the indi-
vidual is considered to exhibit selective predation (e.g., Peckarsky 2006 ; Culler and
Lamp 2009 ).
8.2 What Do Dytiscids Eat?
Dytiscids are considered to be generalists that feed opportunistically on whatever is
available, including conspecifi cs and hetorespecifi cs. Gut content studies, preference
trials, and fi eld observations have shown that zooplankton (Arts et al. 1981 ), insects
(Figs. 8.3 and 8.4 , Johansson and Nilsson 1992 ; Hicks 1994 ), fi sh (Balfour- Browne
1950 ; Dillon and Dillon 1961 ; Le Louarn and Cloarec 1997 ), amphibians (Formanowicz
and Brodie 1982 ; Brodie and Formanowicz 1983 ; Resetarits 1998 ; Rubbo et al.
2006 ; Smith and Awan 2009 ; Inoda et al. 2009 ), reptiles (snakes) (Drummond and
Wolfe 1981 ), and even decaying animal carcasses (Velasco and Millan 1998 ; Barrios
and Wolff 2011 ) are part of their diets. Occasionally, plant material and algae can
also be found in the guts (Deding 1988 ), but plants are considered to be accidentally
ingested (Bosi 2001 ).
Fig. 8.2 A larval Cybister chinensis grasps and consumes a tadpole in the fi eld (Photo by Shin-ya
Ohba 2007)
L.E. Culler et al.
367
Several studies have indicated that both larval and adults stages of certain dytiscid
species selectively feed on certain types of prey relative to others (Koegel 1987 ;
Kehl and Dettner 2003 ; Tate and Hershey 2003 ; Ohba 2009a , b ; Cobbaert et al.
2010 ; Ohba and Inatani 2012 ), sometimes even preferring dead prey to live prey, as
Fig. 8.3 A dragonfl y nymph succumbs to predation by an adult Cybister brevis (Photo courtesy of
Naoto Goto 2003)
Fig. 8.4 Backswimmers (Notonectidae) in a pond serving as prey for a Cybister brevis larva
(Photo by Shin-ya Ohba 2008)
8 Predator-Prey Interactions of Dytiscids
368
is the case with adults of Thermonectus marmoratus (Velasco and Millan 1998 ).
Aditya and Saha ( 2006 ) showed that adult Rhantus sikkimensis preferentially fed on
chironomids versus culicids. Dytiscus circumcinctus larvae preferred mayfl y
nymphs and isopods to caddisfl y larvae whereas the co-occurring D. latissimus had
just the opposite preference (Johansson and Nilsson 1992 ). Yee et al. ( 2013 ) demon-
strated a preference by larval Graphoderus for corixids compared to chironomids
or damselfl ies, but larval Rhantus consumed similar proportions of corixids and
chironomids. In temporary ponds in North Carolina, Dytiscus larvae had a negative
effect on the survival of Pseudacris triseriata tadpoles relative to Bufo americanus
tadpoles (Pearman 1995 ). A few studies have tested the preference of adult and
larval dytiscids feeding on dipterans versus microcrustaceans, with preference noted
for dipterans, including chironomids by adult Potamonectes griseostriatus (Ranta
and Espo 1989 ) and culicids by larval Agabus (Culler and Lamp 2009 ). Some
groups of dytiscids, such as the Hydroporinae, have larvae with elongated nasales
that resemble a pig’s snout (Friis et al. 2003 ) and are presumed adaptations for
capturing microcrustaceans over other types of prey (Galewski 1971 ; de Marzo and
Nilsson 1986 ). In addition to unique morphological adaptations, hunting mode
(Yee 2010 ; Yee et al. 2013 ), hunger level (Hileman et al. 1995 ), visual cues (Nilsson
1986 ), and ontogeny (Friis et al. 2003 ; Ohba 2009b ) are often cited as reasons for
greater consumption of certain prey species relative to others.
8.3 Selective Predation and Effects on Community
Attributes
When explored within a community context, the consequences of selective predation
include effects on prey abundance and prey taxa richness. In general, and not
surprisingly, dytiscids have high feeding rates and therefore can decrease total
macroinvertebrate abundance or biomass (Arts et al. 1981 ; Arnott et al. 2006 ;
Magnusson and Williams 2009 ; Cobbaert et al. 2010 ), with some macroinvertebrate
groups reduced more than others. In fi shless ponds in north-central Alberta, adults
of Dytiscus alaskanus , via preferential consumption, lowered biomass of several
groups including amphipods, leeches, water bugs, damselfl ies, dipterans, and snails
(Cobbaert et al. 2010 ). Higher zooplankton biomass was also noted, indicating a
possible trophic cascade (Cobbaert et al. 2010 ). Similarly, Tate and Hershey ( 2003 )
used lab experiments and molecular analyses to demonstrate preferential feeding by
larval dytiscids ( Agabetes , Celina , Colymbetes , Derovatellus , Dytiscus , and Rhantus)
on larger prey species, including caddisfl ies, fairy shrimp, water bugs, diptera,
amphipods, and also young-of-year grayling. Neither of these studies reported
changes in taxa richness, but Arnott et al. ( 2006 ) found that Graphoderus liberus
adults reduced zooplankton biomass by 21 % and lowered taxa richness and values
of the Shannon-Wiener diversity index for zooplankton. In general, aquatic invertebrate
predators have been shown to affect community attributes due to selective predation
L.E. Culler et al.
369
(e.g., Murdoch et al. 1984 ; Runck and Blinn 1994 ), although studies directed at
dytiscids are limited (Arnott et al. 2006 ) and the consequences of dytiscid predation
on communities are not yet fully understood.
8.4 Cannibalism and Intraguild Predation
Intraspecifi c predation (cannibalism) is quite common among aquatic organisms
(Fox 1975 ) and has been documented among larval dytiscids (Pajunen 1983 ; Juliano
and Lawton 1990 ; Culler and Lamp 2009 ; Yee 2010 ). There is much less evidence
for cannibalism between adults, most likely due to gape limitation (Johnson and
Jakinovich 1970 ). Cannibalism during the larval stage is probably even more prevalent
than the literature suggests given the generalist foraging patterns and voracity of
many dytiscid species (Fig.
8.5 ). Cannibalism has the potential to function as a
density dependent control on dytiscid populations (Juliano and Lawton 1990 ) and
this effect may be more pronounced when alternative prey is limited in abundance
(Culler and Lamp
2009 ). Under these conditions, cannibalism can be viewed as a
lifeboat strategy that allows individuals to persist under sub-optimal ecological
conditions and even accelerate development in temporary ponds that are prone to
drying (Batzer and Wissinger
1996 ). In some species, such as Potamonectes griseostriatus ,
conspecifi cs make up nearly 10 % of the diet and are among the more common prey
items in the diet of larvae (Pajunen 1983 ).
In general, the factors infl uencing the occurrence and frequency of cannibalism
within Dytiscidae are not unlike those across other orders of aquatic insects. In most
aquatic insects, population size structure plays a key role in determining the frequency
Fig. 8.5 Larval Colymbetes
dolabratus , collected from
a pond near Kangerlussuaq,
Greenland, engaging in
cannibalistic interactions
(Photo courtesy of Gifford
Wong 2010)
8 Predator-Prey Interactions of Dytiscids
370
of cannibalism and larger individuals are almost always the cannibal and smaller
individuals the victim (Wissinger 1992 ; Fagan and Odell 1996 ; Hopper et al. 1996 ;
Wissinger et al. 1996 ; Yee 2010 ). However, the relatively large mandibles possessed by
larval dytiscids confer the ability to subdue large prey items including similar-sized
conspecifi cs (Pajunen 1983 ) and perhaps even larger individuals. Avoidance of canni-
balism may be infl uenced by large differences in size between larvae (Pajunen 1983 )
and the ability to recognize and avoid conspecifi cs (Inoda 2012 ). Given the dearth of
studies on cannibalism among dytiscids, these and other aspects of cannibalism deserve
further inquiry. This is particularly true for dytiscids because they occupy relatively
high trophic positions within fi shless systems and recent modeling studies have
demonstrated the potential for cannibalism to strongly infl uence coexistence among
predators and structure communities (Rudolf 2007 ; Ohlberger et al. 2013 ).
Besides cannibalism, intraguild predation (IGP) is likely to be a common interaction
among dytiscids, particularly among larvae for the reasons noted above. IGP is a
mixed competition-predation interaction that occurs when species that compete for
a common resource also interact as predator and prey (see Figs. 3, 4, and 6 in Polis
et al. 1989 ). Simple mathematical models suggest that IGP should be relatively rare
in nature (Holt and Polis 1997 ), but food web studies indicate that IGP is common
across terrestrial, marine, and aquatic systems (Arim and Marquet 2004 ). More
recent theoretical and empirical work indicates that size-structured interactions
such as cannibalism may promote the coexistence of predators in IGP systems
(Crumrine 2005 ; Rudolf 2007 ). There are few studies that specifi cally examine IGP
among larval dytiscids (e.g., Nilsson and Soderstrom 1988 ; Culler and Lamp 2009 ;
Yee 2010 ). As is the case with cannibalism, IGP is probably more prevalent than the
literature suggests given the generalist foraging patterns of larval dytiscids and high
spatial and temporal overlap among species (Yee 2010 ). Of the studies that have
examined IGP among larval dytiscids, not surprisingly, size differences between
individuals infl uence the outcome of predator-prey interactions between intraguild
predators. In some cases larger larvae consume smaller larvae (Nilsson and Soderstrom
1988 ; Yee 2010 ), but there are also examples of IGP between individuals similar in
size (Culler and Lamp 2009 ; Yee 2010 ). In fact, some genera (e.g., Dytiscus ) do not
appear to consume dytiscid prey smaller than themselves and this may promote
coexistence between relatively large- and small-bodied dytiscids (Yee 2010 ). IGP
among larval dytiscids can be symmetric; that is, both predators consume each other
(Culler and Lamp 2009 ; Yee 2010 ). This appears to be most common among conge-
neric species that are similar in size; although higher levels of aggression may also
lead to greater frequency of IGP among some species (Culler and Lamp 2009 ).
Asymmetric IGP appears to be most common when there is a distinct size difference
between individuals (Nilsson and Soderstrom 1988 ; Yee 2010 ). Large-bodied dytiscids,
such as those in the genera Dytiscus and Cybister , are within the guild of top predators
in fi shless ponds and likely function as intraguild predators of larval dytiscids as well
as other large predatory aquatic insects such as odonate nymphs. Despite this, there
are surprisingly few studies that have examined IGP within this group of insects.
Future studies are warranted because IGP among dytiscids is likely to infl uence
coexistence between competing species and it may help to explain the diversity of
species found in some aquatic systems (Yee
2010 ).
L.E. Culler et al.
371
8.5 Non-consumptive Effects of Dytiscid Predation
In addition to consumptive (i.e., lethal) predator effects that change prey abundance
and taxa richness, non-consumptive (i.e., non-lethal) effects are a major component of
predator-prey interactions (Preisser et al. 2005 ). The presence of a predator can trigger
a cascade of changes in prey foraging behavior, physiology, and the timing of life
history events, with consequences for prey fi tness. These non-consumptive effects can
sometimes outweigh consumptive effects (McPeek and Peckarsky 1998 ) and often
cascade to infl uence ecosystem properties and functions (e.g., Schmitz et al. 2010 ).
Removal of aquatic insect predators, including some dytiscids, resulted in altered
migration strategies and an increase in body size of daphnids in fi shless ponds (Herwig
and Schindler 1996 ). Although specifi c investigations of non- consumptive effects of
dytiscids are uncommon, they do offer insights into how these predators may affect
aquatic prey communities. Ohba et al. ( 2012 ) reported that Culex tritaeniorhynchus
female mosquitoes avoided laying eggs in dytiscid- conditioned water and that smaller
mosquitoes emerged from dytiscid-conditioned water as a result of lowered larval
activity. Smith and Awan ( 2009 ) found that American toad and bullfrog tadpoles altered
activity levels and avoided vegetation when dytiscids were present, presumably to
avoid detection and because dytiscids use vegetation as an ambush perch. Similarly,
wood frog tadpoles avoided areas containing caged dytiscids in experimental
mesocosms (Rubbo et al. 2006 ). Johnson et al. ( 2003 ) found that the presence of
dytiscid larvae and other predators of southern leopard frog eggs shortened the time to
hatching and decreased hatchling size. In these preceding examples, dytiscid-induced
changes in a prey’s behavior and size could be energetically costly and have fi tness
consequences, but dytiscid predators can also increase prey fi tness. For example,
in temporary pools adult dytiscids facilitated dispersal of their prey (Beladjal and
Mertens 2009 ); consumption, mastication, and the passage of fairy shrimp through the
digestive tracts of adult dytiscids ( Ilybius fenestratus and Colymbetes fuscus ) led to
increased fairy shrimp hatching (Beladjal and Mertens 2009 ). Given their numeric
and taxonomic dominance in many aquatic systems, non-consumptive predator
effects, positive or negative, should be further explored in dytiscids, especially because
non-consumptive effects of predation on prey have been shown to result in changes to
population and community dynamics (McPeek and Peckarsky 1998 ) and ecosystem
function (Schmitz et al. 2008 ).
8.6 Dytiscids as Predators of Vector and Nuisance Species
Of Coleopteran predators, dytiscids are the most commonly reported predators of
vector and nuisance species, specifi cally mosquito larvae and pupae (Fig. 8.6 , James
1961 , 1964 , 1965 , 1967 ; Lee 1967 ; Young 1967 ; Borland 1971 ; Notestine 1971 ;
Swamy and Rao 1974 ; Akmetbekova and Childibaev 1986 ; Sugiyama et al. 1996 ;
Mogi 2007 ; Quiroz-Martínez and Rodríguez-Castro 2007 ; Shaalan and Canyon
2009 ). Consumption rates can be as high as 86 mosquito larvae per predator per day
8 Predator-Prey Interactions of Dytiscids
372
(Aditya et al. 2006 ), thus warranting their consideration as agents for natural
mosquito suppression. The most commonly reported predators of mosquitoes in
fi eld studies include the genera Laccophilus , Agabus , and Rhantus (Sailer and Lienk
1954 ; Kuhlhorn 1961 ; James 1964 , 1965 ; Lee 1967 ; Roberts et al. 1967 ; Ohba et al.
2010 , Table 8.1 ). Laboratory observations have confi rmed that adult and larval
dytiscids attack mosquito larvae (Table 8.1 ). For example, Bofi ll ( 2014 ) found that
although adult and larval Laccophullis faciatus rufus consumed early and late instar
Culex quinquefasciatus , adults consumed more later instars, suggesting a potential
synergistic effect of dytiscids on mosquito populations. Mosquito larvae have been
found in the guts of fi eld collected dytiscids (Deding 1988 ; Bosi 2001 ). Radioisotope
studies (James 1965 ) and precipitin tests (Service 1973 ) have confi rmed the
prominent roles of dytiscids as mosquito predators. Moreover, serological methods
(Service 1977 , 1993 ) and DNA analysis (Ohba et al. 2010 ) revealed that some species
consumed malaria vector mosquitoes in their natural wetlands.
The effects of dytiscids on mosquitoes have been studied in diverse habitats (Walton
et al. 1990 ; Campos et al. 2004 ; Carlson et al. 2009 ; Ohba et al. 2013 ) including rice
ecosystems (Mogi and Miyagi 1990 ; Mogi 1993 ; Takagi et al. 1996 ; Mogi et al. 1999 ;
Mwangangi et al. 2008 ; Ohba et al. 2011 ), areas where mosquitoes vector disease
(e.g., Chandra et al. 2008 ; Hassan et al. 2010 ), and in northern areas, where dytiscids
often dominate as top predators and have life cycles that are synchronous with those of
their mosquito prey (e.g., James 1964 ; Nilsson and Svensson 1994 ). In a fi eld study in
India, Chandra et al. ( 2008 ) showed that Acilius sulcatus larvae signifi cantly impacted
mosquito larvae that prevail in cement tanks. A signifi cant decrease in larval density of
different mosquito species was observed 30 days after the introduction of A. sulcatus
larvae, while the removal of A. sulcatus resulted in a signifi cant increase in larval
density. These results highlight the effi cacy of A. sulcatus in suppressing larval mos-
quito populations (Chandra et al. 2008 ). Using artifi cial ponds in Sweden, Lundkvist
et al. ( 2003 ) showed that after colonization by large adult dytiscid predators ( Ilybius ,
Rhantus and Agabus spp.), larval mosquito abundance was signifi cantly reduced.
Interest in dytiscids for their mosquito suppression abilities has spurred research
that examines natural patterns in assemblages of dytiscids and culicids (e.g., Nilsson
and Svensson 1995 ). In northern areas, many species of dytiscids have lifecycles
that coincide with mosquito development. In snowmelt ponds in Greenland, larvae
of Colymbetes dolabratus hatch early in the spring just after pond thaw when mosquito
larvae are abundant and one of the only food sources available (Culler 2013 ).
Fig. 8.6 A larval Dytiscus
sp. eats a mosquito larva
(Photo courtesy of Ary
Farajollahi 2009)
L.E. Culler et al.
373
Table 8.1 Summary of predaceous diving beetles and species of mosquito prey
Dytiscid species and stage
a Mosquito species Method b Reference
Acilius sulcatrus L Culex quinquefasciatus LE, FE Chandra et al. (
2008 )
Agabus bipustulatus A Not reported FCG Bosi ( 2001 )
Agabus conspicuous A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Agabus disintegratus L Aedes albopictus LE Culler and Lamp ( 2009 )
Agabus erichsoni L A. communis LE, FE Nilsson and Soderstrom
(
1988 )
Agabus japonicus A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Agabus opacus L A. communis LE, FE Nilsson and Soderstrom
(
1988 )
Agabus punctatus L A. albopictus LE Culler and Lamp ( 2009 )
Colymbetes dolabratus L A. nigripes LE, FO Culler pers obs
Colymbetes paykulli A Culex. spp. LE Lundkvist et al. (
2003 )
Cybister brevis A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Cybister brevis L C. mimeticus FO Ohba (2009)
Cybister japonicus A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Dytiscus marginicolis C. incidens LE Lee ( 1967 )
Eretes griseus A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Eretes sticticus A Not reported LE Swamy and Rao ( 1974 )
Graphoderus adamsii A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Hydaticus bowringii A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Hydaticus grammicus A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Hydaticus grammicus A, L C. tritaeniorhynchus LE Sugiyama et al. ( 1996 )
Hydaticus rhantoides A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Hydroglyphus japonicus A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Hydroglyphus japonicus A C. tritaeniorhynchus LE Sugiyama et al. ( 1996 )
Hydroglyphus pusillus A C. pipiens LE Bellini et al. ( 2000 )
Hyphydrus japonicus A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Ilybius ater A Culex. spp. LE Lundkvist et al. (
2003 )
Ilybius fuliginicolis A Culex. spp. LE Lundkvist et al. (
2003 )
Ilybius subaeneus A Not reported FCG Bosi ( 2001 )
Laccophilus diffi cilis A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Laccophilus fasciatus A C. pipiens LE Roberts et al. ( 1967 )
Laccophilus fasciatus
rufus A, L
A. vexans,
C. quinquefasciatus
LE Pitcher and Yee (2014) ,
Bofi ll (
2014 )
Laccophilus maculosus L A. atropalpus FCR James ( 1965 )
Laccophilus maculosus A C. pipiens LE Roberts et al. ( 1967 )
Laccophilus proximus A A. vexans LE Pitcher and Yee (2014)
Laccotorephes
punctipennis L
A. albopictus FE Sulaiman and Jeffery ( 1986 )
Rhantus pulverosus A, L C. tritaeniorhynchus LE Sugiyama et al. ( 1996 )
Rhantus sikkimensis A C. quinquefasciatus LE Aditya et al. ( 2006 )
Rhantus suturalis A C. tritaeniorhynchus LE Ohba and Takagi ( 2010 )
Rhantus suturalis A Not reported FCG Bosi ( 2001 )
Unknown C. annulirostris NR Rae ( 1990 )
Laccophilus spp. A Anopheles gambiae FCP Ohba et al. ( 2010 )
a L larvae, A adults
b LE lab experiment, FE fi eld experiment, FO fi eld observation, FCG fi eld collection and gut
contents, FCP fi eld collection and PCR, FCR fi eld collection and radioisotopes, NR not reported
8 Predator-Prey Interactions of Dytiscids
374
Similar patterns are found elsewhere in northern regions. Agabus erichsoni
completes its life cycle in woodland pools, overwintering as both eggs and adults,
the appearance of the latter coinciding with the winter hatch of mosquitoes
(James 1961 ; James 1967 ). Nilsson and Svensson ( 1994 ) compared assemblages of
dytiscids and mosquitoes in two boreal snowmelt pools that differed in temperature.
Although dytiscid abundance was similar, dytiscid species richness was higher and
mosquito larvae suffered higher mortality in the warmer pools (Nilsson and
Svensson 1994 ). In Canadian rock pools, James ( 1964 ) found Laccophilus maculo-
sus to be the most abundant predator of the mosquito Aedes atropalpus, with a
significant inverse correlation between densities of Ae. atropalpus and larval
L. maculosus . Smaller pools supported high densities of immature mosquitoes but
no larval dytiscids, suggesting that dytiscids may restrict mosquitoes to certain parts
of the potential breeding habitat (James 1964 ). Based on associations between
landscape structure, including forest cover and water permanence, and mosquito
and dytiscids assemblages, Schafer et al. ( 2006 ) suggested that creating permanent
wetlands in an open landscape would favor colonization by diverse dytiscid assemblages
and therefore reduce mosquito colonization. Other studies have also suggested that
construction of aquatic habitats with the goal of attracting a diverse and abundant
predator assemblage may help to reduce pest abundance (Walton 2012 ). Similar to
ideas proposed in conservation biological control (Barbosa 1998 ), these techniques
may be useful because dytiscids are diffi cult to rear and do not lend themselves well
to use as classical biological control agents.
8.7 Environmental Constraints on Predation
Predation is dynamic in response to changing abiotic and biotic environmental
conditions. Factors such as vegetation structure, the presence and abundance of
intraguild predators, and environmental temperature, via behavioral and physiological
modifi cations, can limit or enhance the ability of a predator to consume prey.
Habitat complexity and structure have been shown to shift dytiscid hunting mode
from actively foraging to sit-and-wait (Michel and Adams 2009 ; Yee 2010 ), though
do not necessarily affect overall prey capture rates (Michel and Adams 2009 ).
Environmentally-induced shifts in predator behavior can have multiple effects on
ecological communities (Michel and Adams 2009 ), including consumptive (Preisser
et al. 2007 ) and non-consumptive effects, the occurrence of trophic cascades,
and even changes in ecosystem function (Schmitz 2008 ). Habitat structural complexity
has also been suggested to enhance predation due to a reduction in negative
intraspecifi c interactions that are a component of dytiscid interactions (i.e., cannibalism
and intraguild predation, Culler and Lamp 2009 ; Yee 2010 ). Given the increasing
numbers of freshwater habitat restoration projects, such as construction of mitigation
wetlands and restoration of drained wetlands on agricultural fi elds, understanding
effects of habitat structure and complexity on dytiscid predation and effects on
food webs remains a high priority for research. For example, wetland construction
L.E. Culler et al.
375
techniques that include adding coarse woody debris or planting diverse aquatic
vegetation could be useful for projects that have goals of encouraging predator
colonization to reduce pest abundance (e.g., Walton 2012 ). This idea largely parallels
a practice used in agricultural habitats known as conservation biological control,
which is defi ned as the manipulation of habitats to favor the natural enemies of
pests, as to conserve biodiversity and reduce pest problems (Barbosa 1998 ).
Temperature, as a fundamental driver of many biological processes, especially in
poikilothermic animals like insects, can also affect dytiscid predation. Calosi et al.
( 2007 ) showed that temperature can alter the diving behavior of dytiscids, with
frequency of diving increasing at higher temperatures, thus decreasing the amount
of time available for other activities such as foraging. Furthermore, temperature,
due to basic effects on the metabolism of ecothermic animals, can also directly
affect consumption rates, with higher intakes necessary to maintain metabolic activ-
ities at higher temperatures (Brown et al. 2004 ; Rall et al. 2010 ). This is consistent
with results from Nilsson and Svensson ( 1994 ) who showed that prey mortality
from dytiscid predation was higher in warmer pools. In addition to direct effects on
rates of predation by dytiscids, temperature can indirectly affect predation via shifts
in the behavior and phenology of the predators and prey (Culler 2013 ). Understanding
these temperature effects is a research priority, particularly in regions where there is
signifi cant warming occurring (e.g., Arctic and alpine regions) and where dytiscids
occur as top predators and have a strong infl uence on the prey community, particu-
larly mosquito abundance.
8.8 Dytiscids as Prey
The role of dytiscids in the trophic ecology of freshwater food webs is often inves-
tigated from the standpoint of dytiscids as top predators, but dytiscids also make
up parts of the diets of many other organisms, both aquatic and terrestrial.
Odonates are known predators of dytiscids (Fig. 8.7 , Larson 1990 R. Roughley
personal communication), but there are few, if any, published reports of other
aquatic invertebrates feeding on dytiscids. Dytiscids cuticle has been recovered
from dissected fi sh guts (Laufer et al. 2009 ), suggesting that that adult and larval
dytiscids are consumed by fi sh (e.g., Closs 1996 ; Laufer et al. 2009 ). There is
evidence, however, that some fi sh and dytiscids do not typically co-occur
(Schilling et al. 2009 ; de Mendoza et al. 2012 ) and thus the extent to which fi sh
rely on dytiscids for food is not well known. In mountain lakes, the distribution of
Agabus bipustulatus is constrained due to predation by salmonid fi sh, and thus they are
found to only inhabit colder lakes where fi sh are unlikely to occur (de Mendoza
et al. 2012 ). Gerhart et al. ( 1991 ) also showed that dytiscids can secrete defensive
hormones that inhibit feeding by fi sh. Dytiscids are part of the diets of turtles
(Chessman 1984 ; Georges et al. 1986 ; Demuth and Buhlmann 1997 ), bullfrogs
(Korschgen and Moyle
1955 ; Bruggers 1973 ), toads (Whitaker et al. 1977 ), sala-
manders (Whiles et al. 2004 ; Dasgupta 1996 ), and snakes (Peddle and Larson 1999 ).
8 Predator-Prey Interactions of Dytiscids
376
The evidence for snake predation comes from postulation that scratch marks on
the beetle’s cuticle were caused from predator attacks in areas with known snake
populations (Peddle and Larson 1999 ).
Dytiscids transcend the aquatic food web by virtue of bird and mammal diets.
Numerous studies have confi rmed the role of adult and larval dytiscids in the
diets of birds, particularly in birds species that are associated with water (e.g.,
Schubart et al. 1965 ; Abensperg-Traun and Dickman 1989 ; Goutner and Furness
1997 ; Elmberg et al. 2000 ), but also in hawks (Munro 1929 ) and fi nches (Montalti
et al. 2005 ). Pellets collected from colonies of grey herons in northern Poland
consisted of 26–51 % invertebrate remains, mainly the dytiscid beetle Dytiscus
marginalis (Jakubas and Mioduszewska 2005 ). Forty-one percent of regurgitate
material from Glossy Ibises in Spain were dytiscids, primarily Cybister (Macías
et al. 2004 ). In Arkansas, dytiscids make up 19 % of the King Rail’s diet during
the winter months (Meanley 1956 ). Brooks ( 1967 ) presented data on the diets of
various species of shorebirds in Illinois, the majority of which contained adults
of the dytiscid beetles Agabus disintegratus and Hygrotus . Raccoons (Capinera
2010 ) and otters (Brzeziński et al. 1993 ) are also noted predators of dytiscid
beetles. During the warm season, dytiscids are the third most important prey
item in terms of biomass for river otters in eastern Poland (Brzeziński et al.
1993 ). The only other mammals known to ingest dytiscids are humans. Several
species in the genus Cybister are regularly consumed in parts of China (Jäch
2003 ), Thailand (Chen et al. 1998 ), New Guinea (Gressitt and Hornabrook 1977 ),
and Japan (S. Ohba, personal observation).
Fig. 8.7 Dragonfl y nymphs and dytiscids frequently co-occur and engage in intraguild predation.
Here, a large Anax dragonfl y (Odonata: Aeshnidae) nymph consumes a Graphoderus larva
(Photo courtesy of Donald Yee 2007)
L.E. Culler et al.
377
8.9 Future Directions
Dytiscids are an ideal study organisms for basic and applied ecological research
due to their ubiquitous and global distribution and ease of handling in the labora-
tory (Fig. 8.8 ) and fi eld (Fig. 8.9 ). Freshwater food webs in lentic habitats, par-
ticularly in fi shless ponds, are much less studied than those in streams and lakes
(Klecka and Boukal 2012 ). Given the role of dytiscids in these habitats as
top predators, predators of mosquitoes, and as food for other organisms, future
studies are essential for understanding the processes that structure freshwater
lentic communities.
Basic information about dytiscid trophic ecology remains largely unknown.
For example, we do not know the prevalence of generalist versus specialist type
feeding in dytiscids and how this changes with ontogeny, and the role, if any, of
plants in the diets of dytiscids. We also do not know the extent to which fi sh and
other aquatic or terrestrial organisms rely on dytiscids as a part of their diets.
Improved basic knowledge about the position and relative importance of dytiscids
in freshwater food webs will promote the design of studies that address basic and
applied questions.
Fig. 8.8 Dytiscids are ideal for use in laboratory experiments where various factors can be manipulated,
including habitat structure, temperature, and relative abundance of different types of prey. Here,
small plastic cups housing dytiscid larvae are used as microcosms to test the effects of structure
and prey density on antagonistic predator-predator interactions (Photo by Lauren Culler 2008)
8 Predator-Prey Interactions of Dytiscids
378
Dytiscids are easily handled and observed and thus are ideal for testing ecological
theories of predator-prey interactions, species coexistence, and consumptive and non-
consumptive effects of predators on prey. Future studies should address prey choice
and the mechanisms underlying selective predation. These studies are needed for
understanding the relative intensity of intraguild predation and cannibalism among
dytiscids and the role of size structure in shaping these interactions. Behavioral studies
are needed to elucidate the non-consumptive effects of dytiscid predation on the
behavior and life-history traits of prey. The consequences of selective predation, intra-
guild predation, cannibalism, and non-consumptive effects by dytiscids remain
largely unknown but have the potential to strongly infl uence population dynamics
and species coexistence (Yee 2010 ).
Environmental changes that are occurring at local and global scales, including
habitat degradation and restoration and climate change, necessitate further study of
how environmental factors shape dytiscid effects on prey, particularly factors such
as habitat structure and temperature. As predators of vector and nuisance species,
some dytiscids have potential as biological control agents. Measuring their effects
on nuisance prey populations and testing how habitat and environmental factors
infl uence these effects are essential for projects that aim to construct or restore natural
lentic habitats while minimizing increased threats from vectors.
Fig. 8.9 Field experiments are useful for measuring effects of dytiscid predation on prey. Here,
white pans are set up adjacent to a tundra pond and used to measure consumption rates of mosquito
larvae by dytiscid predators (Photo by Lauren Culler 2012)
L.E. Culler et al.
379
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Lauren E. Culler completed her B.S. in Biology in 2005 and M.S. in Entomology
in 2008, both from the University of Maryland, College Park, and her Ph.D. in
Ecology and Evolutionary Biology from Dartmouth College in 2013. Her interests
were shaped by an early fascination with dytiscids and include how environmental
factors affect predator-prey interactions in systems that are linked to humans.
She works with dytiscids in restored agricultural wetlands on the Eastern Shore of
Maryland and tundra ponds in Arctic Greenland.
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Shin-ya Ohba has been at Nagasaki University as associate professor since 2012. He
completed his B.S. in Agriculture in 2002 from Tamagawa University, M.S. in
Agriculture from Ehime University in 2004, and a Ph.D. in Entomology from
Okayama University in 2007. His research interests are basic ecology, the role of
predaceous diving beetles as mosquito predators, and also educational effects of
dytiscid on the students (“eggs” of teacher) in his laboratory.
Patrick Crumrine earned his B.S. in
Biology in 1998 from SUNY Platts-
burgh and his Ph.D. in Biology from
the University of Kentucky in 2003.
The primary focus of his research
is to understand how size-structure
infl uences competition and predation
among aquatic organisms, particu-
larly odonates and dytiscids. Other
research interests include disease
ecology of amphibians and population/
community structure of aquatic turtles.
Most of his work is conducted in small
ponds and wetlands in the mid-Atlantic
region of the United States.
L.E. Culler et al.
