Robert L Smith

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• Daly-Engel, T., Smith, R. L., Finn, D. S., Knoderbane, M. E., Phillipsen, I. C., & Lytle, D. A. (2012). 17 novel polymorphic microsatellite markers for the giant water bug, Abedus herberti (Belostomatidae). Conservation Genetics Resources, 4(4), 979-981.
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  Abstract: The giant water bug (Abedus herberti) is a large flightless insect that is a keystone predator in aridland aquatic habitats. Extended droughts, possibly due to climate change and groundwater pumping, are causing once-perennial aquatic habitats to dry, resulting in serious conservation concern for some populations. A. herberti also exhibits exclusive male parental care, which has made it a model organism for studying mating systems evolution. Here we describe 17 novel polymorphic microsatellite loci developed for A. herberti. Number of alleles per locus ranged from 2 to 15, and average observed and expected heterozygosities were 0. 579 and 0. 697, respectively. These loci can successfully resolve both population genetic structure among sites separated by 3-100 km (F ST = 0. 08-0. 21, P < 0. 0001), and divergent mating strategies within local populations, making them highly useful for conservation genetics studies of this vulnerable species. © 2012 Springer Science+Business Media B.V.
• Daly-Engle, T., Smith, R., FinnDS, ., Knoderbane, M., Phillipsen, I., & Lytle, D. (2012). 17 novel polymorphic microsatellite markers for the giant water bug, Abedus herberti (Belostomatidae. Conservation Genetics Resources, 4, 979-981.
• Goforth, C. L., & Smith, R. L. (2012). Subsurface behaviours facilitate respiration by a physical gill in an adult giant water bug, Abedus herberti. Animal Behaviour, 83(3), 747-753.
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  Abstract: Abedus herberti is an aquatic bug that carries a subalar store of atmospheric air for use during submersion. When submersed, adult bugs routinely show two patterns of behaviour we call gaping and dynamic gaping. We studied these behaviours to determine when they are expressed and tested the hypothesis that the behaviours serve a respiratory function. Gaping and dynamic gaping occurred when bugs were positioned sufficiently deep in water that they could not refresh air stores without releasing their hold on the substrate and floating to the surface. Gaping and dynamic gaping occurred more frequently as water depth increased, and bugs performing these behaviours remained submersed longer than bugs that did not express these behaviours. Bugs permitted to express gaping and dynamic gaping remain submersed longer than bugs that were experimentally prevented from performing the behaviours. Bugs also remained submersed longer in high oxygen water than in low oxygen water. We conclude that gaping allows the subalar air store to function as a physical gill and that dynamic gaping is a form of behavioural ventilation. Adult A. herberti are buoyant and must swim or hold onto a substrate to remain submersed. Extending submersion time increases predatory efficiency, reduces the frequency of surfacing, saves energy and reduces predation risk. © 2011 The Association for the Study of Animal Behaviour.
• Goforth, C., & Smith, R. (2012). Subsurface behaviors facilitate oxygen uptake by a physical gill in an adult giant water bug, Abedus herberti (Hemiptera: Belostomatidae). Animal Behavior, 83, 747-753.
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  doi:10.1016/j.anbehav. 2011.12.023
• Goforth, C. L., & Smith, R. L. (2011). Respiratory morphology of the Abedus herberti Hidalgo egg chorion (Hemiptera: Belostomatidae). Journal of Morphology, 272(7), 796-801.
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  PMID: 21472766;Abstract: Although giant water bugs (Hemiptera: Belostomatidae) are large, aquatic insects known for their obligate paternal egg brooding behaviors, little research has focused on the structure of their eggs. The respiratory requirements of developing embryos likely created selection for brooding, so a thorough understanding of the respiratory morphology of belostomatid eggs could help explain how brooding behaviors facilitate embryonic gas exchange. This study used scanning electron microscopy to document the respiratory microstructure of the eggs of Abedus herberti, a back brooding giant water bug. The exochorion is similar to that of other belostomatids in texture and organization except that the respiratory region is confined to the uppermost quarter of the egg. This is the area exposed to the atmosphere by encumbered males. A plastron network made up of densely packed vertical projections demarcates the boundary between the respiratory and nonrespiratory regions of the chorion. The internal chorion is composed of alternate air-filled and denser layers that likely facilitate the movement of oxygen from the aeropyles at the top of the eggs to the developing embryonic tissues. © 2011 Wiley-Liss, Inc.
• Smith, R., Goforth, C. L., & Smith, R. L. (2011). Respiratory morphology of the Abedus herberti Hidalgo egg chorion (Hemiptera: Belostomatidae). Journal of morphology, 272(7).
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  Although giant water bugs (Hemiptera: Belostomatidae) are large, aquatic insects known for their obligate paternal egg brooding behaviors, little research has focused on the structure of their eggs. The respiratory requirements of developing embryos likely created selection for brooding, so a thorough understanding of the respiratory morphology of belostomatid eggs could help explain how brooding behaviors facilitate embryonic gas exchange. This study used scanning electron microscopy to document the respiratory microstructure of the eggs of Abedus herberti, a back brooding giant water bug. The exochorion is similar to that of other belostomatids in texture and organization except that the respiratory region is confined to the uppermost quarter of the egg. This is the area exposed to the atmosphere by encumbered males. A plastron network made up of densely packed vertical projections demarcates the boundary between the respiratory and nonrespiratory regions of the chorion. The internal chorion is composed of alternate air-filled and denser layers that likely facilitate the movement of oxygen from the aeropyles at the top of the eggs to the developing embryonic tissues. J. Morphol., 2011. © 2011 Wiley-Liss, Inc.
• Smith, R., Kern, K. B., Stickney, R. E., Gallison, L., & Smith, R. L. (2010). Metronome improves compression and ventilation rates during CPR on a manikin in a randomized trial. Resuscitation, 81(2).
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  We hypothesized that a unique tock and voice metronome could prevent both suboptimal chest compression rates and hyperventilation.
• Smith, R. L., Schnack, J. A., Schaefer, E. F., & Kehr, A. I. (2008). Ticks, Amblyomma rotundatum (Acari: Ixodidae), on toads, Chaunus schneideri and Chaunus granulosus (Anura: Bufonidae), in Northern Argentina. Journal of Parasitology, 94(2), 560-562.
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  PMID: 18564766;Abstract: This communication provides notes on 2 species of toads, Chaunus schneideri and Chaunus granulosus, infested with ixodid ticks, Amblyomma rotundatum, from the provinces of Corrientes and Formosa in northern Argentina. Chaunus schneideri is a new amphibian host record for A. rotundatum, a species previously reported to parasitize other anurans and also reptiles. We examined 74 ticks on 5 toads. All ticks were A. rotundatum; all adults were females, and all developmental stages were randomly attached to host body parts. Ticks remained attached to one of the toads for from 7 to 17 days after the host was captured. One toad, encumbered with 33 ticks, was moribund when found and died shortly thereafter. © American Society of Parasitologists 2008.
• Smith, G., Trumbo, S. T., Sikes, D. S., Scott, M. P., & Smith, R. L. (2007). Host shift by the burying beetle, Nicrophorus pustulatus, a parasitoid of snake eggs. Journal of Evolutionary Biology, 20(6), 2389-2399.
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  PMID: 17956400;Abstract: Recent work [Ecoscience (2000) vol. 7, 395-397] suggests that the burying beetle Nicrophorus pustulatus may have undergone a remarkable host shift, exploiting snake eggs rather than carrion as resources for breeding. We conducted behavioural and physiological experiments to examine the hypothesis of a host shift and to formulate hypotheses on its origin. Two congeners of N. pustulatus, Nicrophorus orbicollis and Nicrophorus defodiens did not respond to snake eggs with typical breeding behaviour. When N. pustulatus male-female pairs (n = 14) were presented with clutches of snake eggs, the number of offspring but not the mean size of offspring varied with snake egg mass, indicating effective regulation of brood size. When breeding on turtle eggs, N. pustulatus had a more variable response than when exploiting snake eggs, suggesting that turtle eggs are not a primary resource for breeding. Nicrophorus pustulatus presented with both snake eggs and a mouse carcass combined and exploited the two resources within the same nest (10 of 12 trials). Mouse carcasses and snake eggs were treated differently. Carcasses were moved, buried and stripped of hair in a manner characteristic of burying beetles, whereas snake eggs were not moved or buried. Females that discovered a mouse carcass also had a significantly greater juvenile hormone increase than did females discovering snake eggs. Some responses to the two resources, however, were similar. Female N. pustulatus oviposited rapidly in response to either a mouse carcass or snake eggs, and males elevated sex pheromone emission in response to either resource. The efficient use of snake eggs, the ability to regulate brood size and the different responses to snake eggs and carrion suggest that N. pustulatus is well adapted to exploiting snake eggs for breeding. The use of snake eggs by N. pustulatus has potential implications for conservation of oviparous reptiles. © 2007 The Authors.
• Vu, Q. M., Nguyen, H. H., & Smith, R. L. (2007). The termites (Isoptera) of Xuan Son National Park, northern Vietnam. Pan-Pacific Entomologist, 83(2), 85-94.
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  Abstract: Subterranean termite species, their habitat preferences and nest habits at Xuan Son National Park, a lowland and lower mountain evergreen and limestone forest in northern Vietnam, were investigated in 2002 and 2003. A total of 234 collections were obtained from 588 sampling sites, on 12 transects, among four different habitat types. Fifteen species in eight genera and two families were recorded. Termitidae was the dominant family with six genera and 12 species. The genus Odontotermes with five, contained the largest number of species. Five species were new records for northern Vietnam: Odontotermes maesodensis Ahmad, 1965, Nasutitermes ovatus Fan, 1983, Pericaptitermes latignathus (Holmgren, 1913), Pericaptitermes nitobei Shiraki, 1909 and Bulbitermes laticephalus Ahmad, 1965. The inventory included eight fungus-growing species: Macrotermes barneyi Light, 1924, Ma. annandalei (Silvestri, 1914), O. yunnanensis Tsai et Chen, 1963, O. hainanensis Light, 1924, O. formosanus Sharaki, 1909, O. maesodensis Ahmad, 1965, O. graveli (Silvestri, 1914) and Microtermes pakistanicus Ahmad, 1965. Five species: M. barneyi, O. yunnanensis, O. hainanensis, O. formosanus and M. pakistanicus occurred in all habitat types. The scrubland/grassland habitat contained 14 species, forest habitats 12 species, cultivated lands eight species and residential areas contained only six species. In forest habitats 56.2% of sample plots yielded termites, in scrubland/grasslands 54.5%, 32% of plots on cultivated land produced termites and in residential habitats only 18% had termites. Six species identified are considered special pests because their activities weaken earthen structures: M. pakistanicus, M. barneyi, M. annandalei, O. yunnanensis, O. hainanensis and O. formosanus. Nesting patterns of surveyed species are noted with special attention to species that inhabit earthen structures.
• Miettinen, M., Kaitala, A., Smith, R. L., & Ordóñez, R. M. (2006). Do egg carrying and protracted copulation affect mobility in the golden egg bug?. Journal of Insect Behavior, 19(2), 171-178.
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  Abstract: Golden egg bug Phyllomorpha laciniata (Heteroptera, Coreidae) females oviposit on male and female conspecifics that carry ova until they hatch. Embryos benefit from being carried because of diminished risks of predation. Female carriers are never the parents of carried eggs, and males are only rarely the fathers of any carried eggs. Eggs develop and hatch without being carried in the laboratory. Egg carriers may be viewed as victims, exploited by females that encumber them with eggs. The intensity of selection favouring resistance to egg carrying should be proportional to the costs of this behaviour. One possible cost could be a reduction in mobility caused by carried eggs. We compare movement rates among encumbered and unencumbered golden egg bugs of both sexes. Protracted copulations (often exceeding 20 h) typical of this species and mating may also cause reduction in bugs mobility. Therefore, we also evaluate rates of movement of coupled pairs of bugs. Our results indicate that egg loads do not affect the mobility and speed of either males or females. However, copulating pairs are significantly slowed as compared to single bugs. Thus protracted copulations have a clear cost in rates of movement and possibly risks of predation, but there are no apparent mobility costs for egg carrying. © 2006 Springer Science+Business Media, Inc.
• Lytle, D. A., & Smith, R. L. (2004). Exaptation and flash flood escape in the giant water bugs. Journal of Insect Behavior, 17(2), 169-178.
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  Abstract: Although behaviors may remain highly conserved through evolutionary time, the ecological functions they serve can undergo surprising transformations. We used phylogenetic, correlational, and experimental evidence to show how a >150-million year-old behavior, which originally evolved to facilitate migration, has been co-opted for flash flood escape in two distantly related giant water bug species (Hemiptera: Belostomatidae). Using behavioral experiments with simulated rainfall, we showed that species from flash-flooding as well as non-flash-flooding environments are capable of rainfall response behavior (RRB), the ability to use rainfall as a cue to abandon an aquatic habitat. The results suggest that, in addition to allowing individuals to escape flash floods, RRB is the proximate mechanism generating a well-established ecological pattern: The correlation between rainfall and migration to seasonal breeding habitats that has been documented in 13 species throughout the family. Placing RRB in phylogenetic context reveals that for several taxa the behavior is an exaptation (a trait evolved for one function but later co-opted for another) for escaping flash floods. For Lethocerus medius, rainfall response behavior is an addition exaptation because the behavior is used to initiate migration to seasonal rain pools (ancestral function) as well as for flash flood escape (co-opted function). In the distantly related Abedus herberti, rainfall response behavior is a transfer exaptation because it has been co-opted exclusively for flash flood escape and the ancestral function has been lost. These findings emphasize that a phylogenetic framework is needed to fully understand the origins and ecological significance of behaviors.
• Kaitala, A., & Smith, R. L. (2002). Do golden egg bugs (Phyllomorpha laciniata: Heteroptera, Coreidae) require conspecifics for oviposition?. Journal of Insect Behavior, 15(2), 171-180.
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  Abstract: Golden egg bug (Phyllomorpha laciniata) females lay eggs on the bodies of conspecifics of both sexes. We investigated to what extent reproduction depended on the availability of conspecifics as oviposition substrate and the acceptability of the host plant as an alternative oviposition substrate in the absence of conspecifics. Mated females were placed in experimental enclosures each containing a sprig of fresh host plant. Each experimental female was subjected to one of three treatments: isolated from conspecifics (solitary), paired with another female, or paired with a male. Solitary females laid a few eggs on the host plant but then stopped laying eggs, and solitary females laid significantly fewer eggs than those enclosed with another female or a male. Females enclosed with a male laid no more eggs than those enclosed with a female. When two previously isolated females were later enclosed together, they soon renewed oviposition. Females in nature contained significantly more oviducal eggs than did females that were enclosed with other females for a short period. Thus the availability of suitable conspecifics as oviposition substrate stimulates the deposition of mature eggs, and reproduction depends on the presence of conspecifics of either sex as oviposition substrate.
• Kaitala, A., & Smith, R. L. (2002). The bug that lays the golden eggs. Natural History, 111(2), 32-37.
• Hoekstra, J. D., & Smith, R. L. (1999). Descriptions of the final instar larvae of Argia sabino Garrison and Argia pima Garrison (Odonata: Coenagrionidae). Proceedings of the Entomological Society of Washington, 101(4), 887-896.
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  Abstract: We illustrate and describe the final instar larvae of Argia sabino Garrison 1994 and Argia pima Garrison 1994 based on preserved exuviae and larvae from Sabino Creek, Arizona, U.S.A. A dichotomous key is provided to integrate A. sabino and A. pima into an existing larval key to North American Argia spp.
• Hoekstra, J. D., & Smith, R. L. (1998). Distribution and habitat of Curicta pronotata (Hemiptera: Nepidae) in southeastern Arizona. Entomological News, 109(5), 366-368.
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  Abstract: Curicta pronotata is known from western Mexico and southeastern Arizona. Previous accounts of the species' Arizona distribution draw from only a few collections and localities. Distributional records clarifying the range and habitat of C. pronotata in Arizona are reported here.
• Smith, R. L., & Horton, C. (1998). Fish predation on giant water bug (Heteroptera: Belostomatidae) eggs in an Arizona stream. Great Basin Naturalist, 58(3), 292-293.
• Lokensgard, J., Smith, R. L., Eisner, T., & Meinwald, J. (1993). Pregnanes from defensive glands of a belostomatid bug. Experientia, 49(2), x12-176.
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  PMID: 8440354;Abstract: The aquatic bug Abedus herberti (Hemiptera: Belostomatidae) secretes a mixture containing four pregnanes (desoxycorticosterone (I), pregnenolone (II), progesterone (III), and 3α-hydroxy-pregn-5-en-20-one (IV)) from its cephalic glands. Pregnanes had previously been characterized from the defensive glands of aquatic beetles (Dytiscidae) and shown to be deterrent to fish. It may be specifically under predation pressure from fish that A. herberti and Dytiscidae evolved their comparable defenses.
• Smith, R. L., & Larsen, E. (1993). Egg attendance and brooding by males of the giant water bug Lethocerus medius (Guerin) in the field (Heteroptera: Belostomatidae). Journal of Insect Behavior, 6(1), 93-106.
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  Abstract: Males of the giant water bug Lethocerus medius (Guerin) typify their monobasic subfamily, the Lethocerinae, in that they do not brood eggs attached to their backs as do males of all members of the subfamily Belostomatinae. Exclusive male parental investment as expressed in the Belostomatinae is extremely rare behavior among animals, and evolution of the trait is obscure. Lethocerus medius males apparently remain with their mates through oviposition and are consistently found in attendance of eggs after the female has departed. This behavior may enhance paternity assurance at no cost in opportunity for polygyny. Two double clutches of eggs were found, from which we infer the potential for polygynous matings and shared parental investment. Male L. medius brood attended egg clutches above the surface of the water, where they may moisten them, shade them, and defend them against predation. Egg attendance/brooding by L. medius and other Lethocerus species may represent a plesiomorphic state from which paternal back- brooding evolved in the Belostomatinae. © 1993 Plenum Publishing Corporation.
• Zeh, D. W., Zeh, J. A., & Smith, R. L. (1989). Ovipositors, amnions and eggshell architecture in the diversification of terrestrial arthropods. Quarterly Review of Biology, 64(2), 147-168.
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  Abstract: Comparison with the Parainsecta (Collembola, Protura) suggests that the ovipositor, amnion and complex chorion are novel features acquired early in the evolution of the insects. Insect diversity is probably at least partially a consequence of this suite of egg-stage characters which reduced constraints on suitable sites for egg deposition, and enabled insect lineages to diversify into previously inaccessible niches. The self-sufficient insect egg, resistant to osmotic rupture, desiccation, and drowning, may explain the low incidence of postzygotic parental investment among insects relative to other terrestrial arthropods. Ovoviviparity, viviparity and parental egg care may have inhibited the evolution of such eggs in other terrestrial arthropod taxa. Cladistic data suggest a historical association between diversification and expansion in type or range of habitats utilized for oviposition and larval development. Insects would probably have been unable to exploit the potential of holometaboly and flight without the capacity to ensure egg survival in the wide range of oviposition substrates provided by terrestrial environments. -from Authors
• Zeh, D. W., & Smith, R. L. (1985). Paternal investment by terrestrial arthropods. Integrative and Comparative Biology, 25(3), 785-805.
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  Abstract: Paternal investment in terrestrial arthropods occurs in three contexts. Prezygotic investment includes indirect contributions to offspring through nutrients provided o the male's mate. Biparental care refers to joint male and female care of offspring. Finally, exclusive paternal care occurs when only the male invests in offspring following oviposition. Examples of exclusive paternal care are known in insects such as assassin bugs (Reduviidae), harvestmen (Opiliones), and millipedes (Diplopoda), although it is far more common in a group of secondarily aquatic insects, namely, the giant water bugs (Belostomatidae). Biparental care is also uncommon and is best developed in burying beetles (Silphidae), dung beetles (Scarabaeidae), and termites (Isoptera). The most pervasive type of paternal investment appears to be prezygotic in the form of spermatophore products and other "nuptial gifts" provided to the female parent.The evolution of paternal investment is a complex process and no single hypothesis nor evolutionary pathway appears adequate to explain the diversity of paternal investment strategies in terrestrial arthropods. As is the case with other animal groups, paternal investment is correlated with certainty of paternity and male territoriality. Ecological factors also appear important, especially in the way these influence the ability of males to enhance the survivorship of offspring and/or the fecundity of their mates. Physically harsh or biotically dangerous habitats and ephemeral, highly prized, productive resources are all associated with high levels of paternal investment. Finally, the indirect sperm transfer strategies common to many terrestrial arthropod species seem to preclude males from contributing materially to their offspring by dissociating parent from progeny. This dissociation may explain in part the relative paucity of high levels of paternal investment by terrestrial arthropods. © 1985 by the American Society of Zoologists.
• Smith, R. L. (1979). Paternity assurance and altered roles in the mating behaviour of a giant water bug, Abedus herberti (heteroptera: Belostomatidae). Animal Behaviour, 27(PART 3), 716-725.
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  Abstract: Natural selection theory predicts that male brooding giant water bugs, Abedus herberti Hidalgo, should possess paternity assurance mechanisms and have altered roles in courtship. Mating behaviour of this species was studied to check these predictions. Females were aggressive in courtship, but male 'display' was an essential element. Copulation always preceded oviposition, and copulation and oviposition were cyclical events under male control. Females engaged in cyclical polyandry in the laboratory, and this threat under natural conditions presumably favoured the male-dominated system of alternating bouts of copulation and oviposition. Male and gravid female bugs seemed equally eager to exchange brooding services for eggs under laboratory conditions. Courtship roles are discussed in relation to relative parental investment and availability of eligible males. © 1979.
• Smith, R. L. (1979). Repeated copulation and sperm precedence: Paternity assurance for a male brooding water bug. Science, 205(4410), 1029-1031.
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  PMID: 17795564;

Presentations

• Goforth, C., & Smith, R. (2012, January). The respiratory behaviors of Abedus herberti (Hemiptera: Belostomatidae). 2006 ESA Annual Meeting. Indianapolis, Indiana.
• Goforth, C., & Smith, R. (2011, January). The respiratory behaviors of Abedus herberti (Hemiptera: Belostomatidae). 2006 ESA Annual Meeting. Indianapolis, Indiana.