%0 Journal Article %J Fungal Biology %D 2014 %T Aquastella gen. nov.: a New Genus of Saprolegniaceous Oomycete Rotifer Parasites Related to Aphanomyces, With Unique Sporangial Outgrowths %A D. P. Molloy %A Glockling, S. L. %A C. A. Siegfried %A Beakes, G. W. %A James , T. Y. %A Matitsky, S. E. %A Wurdak, E. %A L. Giamberini %A Gaylo, M. J. %A Nemeth, M. J. %K Aquastella acicularis %K Aquastella attenuata %K Convergent evolution %K Phylogenetic analysis %K Saprolegniales %X

The oomycete genus Aquastella is described to accommodate two new species of parasites of rotifers observed in Brooktrout Lake, New York State, USA. Three rotifer species – Keratella taurocephala, Polyarthra vulgaris, and Ploesoma truncatum – were infected, and this is the first report of oomycete infection in these species. Aquastella attenuata was specific to K. taurocephala and Aquastella acicularis was specific to P. vulgaris and P. truncatum. The occurrence of infections correlated with peak host population densities and rotifers were infected in the upper layers of the water column. Sequencing of 18S rRNA and phylogenetic analysis of both species placed them within the order Saprolegniales, in a clade closely related to Aphanomyces. The Aquastella species were morphologically distinct from other rotifer parasites as the developing sporangia penetrated out through the host body following its death to produce unique tapered outgrowths. Aquastella attenuata produced long, narrow, tapering, finger-like outgrowths, whilst A. acicularis produced shorter, spike-like outgrowths. We hypothesize that the outgrowths serve to deter predation and slow descent in the water column. Spore cleavage was intrasporangial with spore release through exit tubes. Aquastella attenuata produced primary zoospores, whereas A. acicularis released spherical primary aplanospores, more typical of other genera in the Aphanomyces clade.

%B Fungal Biology %V 118 %P 544-558 %G eng %U http://dx.doi.org/10.1016/j.funbio.2014.01.007 %R 10.1016/j.funbio.2014.01.007 %0 Journal Article %J Journal of Invertebrate Pathology %D 2013 %T Mode of Action of Pseudomonas fluorescens Strain CL145A, A Lethal Control Agent of Dreissenid Mussels (Bivalvia: Dreissenidae) %A D. P. Molloy %A D. A. Mayer %A L. Giamberini %A Gaylo, M. J. %K Biological control %K Biopesticide %K Dreissena polymorpha %K Hemocyte infiltration %K Pseudomonas fluorescens %K Zebra mussel %X

Pseudomonas fluorescens strain CL145A (Pf-CL145A) has demonstrated promise as an efficacious and selective agent for the control of macrofouling Dreissena spp. mussels. Herein, we report trials to investigate the mode of action of this biocontrol agent against Dreissena polymorpha, the zebra mussel. Exposure to dead Pf-CL145A cells achieved the same temporal pattern and percentage mussel mortality as did live cells, thereby excluding infection as the possible lethal mode of action. Histological analysis revealed pathologies consistent with the cause of death being intoxicating natural products associated with Pf-CL145A cells. Irrespective of whether the mussels were exposed to live or dead Pf-CL145A cells, examination of tissues from histological sections revealed that: (1) at the end of the 24-h treatment period there was massive hemocyte infiltration into the lumina of both the digestive gland and stomach; and (2) mussel deaths occurred following lysis and necrosis of the digestive gland and sloughing of stomach epithelium. These trials provide strong evidence that the lethal mode of action of Pf-CL145A is intoxication.

%B Journal of Invertebrate Pathology %V 113 %P 115-121 %G eng %U http://dx.doi.org/10.1016/j.jip.2012.12.013 %R 10.1016/j.jip.2012.12.013 %0 Journal Article %J Management of Biological Invasions %D 2013 %T Non-target Trials with Pseudomonas fluorescens Strain CL145A, A Lethal Control Agent of Dreissenid Mussels (Bivalvia: Dreissenidae). %A D. P. Molloy %A D. A. Mayer %A Gaylo, M. J. %A Burlakova, L. E. %A Karatayev, A. Y. %A Presti, K. T. %A Sawyko, P. M. %A Morse, J. T. %A Paul, E. A. %K biocontrol %K Dreissena polymorpha %K Dreissena rostriformis bugensis %K quagga mussel %K Zebra mussel %K Zequanox® %X

In an effort to develop an efficacious and environmentally safe method for managing zebra mussels (Dreissena polymorpha) and quagga mussels (Dreissena rostriformis bugensis), we initiated a research project investigating the potential use of bacteria and their natural metabolic products as biocontrol agents. This project resulted in the discovery of an environmental isolate lethal to dreissenid mussels, Pseudomonas fluorescens strain CL145A (Pf-CL145A). In previous published reports we have demonstrated that: 1) Pf-CL145A’s mode of action is intoxication (not infection); 2) natural product within ingested bacterial cells lyse digestive tract epithelial cells leading to dreissenid death; and 3) high dreissenid kill rates (>90%) are achievable following treatment with Pf-CL145A cells, irrespective of whether the bacterial cells are dead or alive. Investigating the environmental safety of Pf-CL145A was also a key element in our research efforts, and herein, we report the results of non-target trials demonstrating Pf-CL145A’s high specificity to dreissenids. These acute toxicity trials were typically single-dose, short-term (24-72 h) exposures to Pf-CL145A cells under aerated conditions at concentrations highly lethal to dreissenids (100 or 200 mg/L). These trials produced no evidence of mortality among the ciliate Colpidium colpoda, the cladoceran Daphnia magna, three fish species (Pimephales promelas, Salmo trutta, and Lepomis macrochirus), and seven bivalve species (Mytilus edulis, Pyganodon grandis, Pyganodon cataracta, Lasmigona compressa, Strophitus undulatus, Lampsilis radiata, and Elliptio complanata). Low mortality (3-27%) was recorded in the amphipod Hyalella azteca, but additional trials suggested that most, if not all, of the mortality could be attributed to some other unidentified factor (e.g., possibly particle load or a water quality issue) rather than Pf-CL145A’s dreissenid-killing natural product. In terms of potential environmental safety, the results of these invertebrate and vertebrate non-target trials are encouraging, but it would be unrealistic to think that dreissenids are the only aquatic organisms sensitive to Pf-CL145A’s dreissenid-killing natural product. Additional testing is needed to better define Pf-CL145A’s margin of safety by identifying the sensitivity of other susceptible organisms. The results of these non-target safety trials – in combination with equally promising mussel control efficacy data – have now led to Pf-CL145A’s commercialization under the product name Zequanox®, with dead cells as the product’s active ingredient. The commercial availability of only dead-cell Zequanox formulations will eliminate the risk of any possible non-target infection by Pf-CL145A, further reducing environmental concerns. During the non-target project reported herein, the limited quantities of Pf-CL145A cells that we were able to culture severely restricted the number and size of our trials. In contrast, the availability of Zequanox will now greatly expand the opportunities for non-target testing. The trials reported herein – exposing non-target organisms under aerated conditions to unformulated, laboratory-cultured cells – clearly point to Pf-CL145A’s potential for high host specificity, but non-target trials with Zequanox – using Pf-CL145A cells cultured, killed, and formulated using industrial-scale protocols – will be even more important as they will define the non-target safety limits of the actual commercial products under a wide range of environmental conditions.

%B Management of Biological Invasions %V 4 %P 71-79 %G eng %U http://dx.doi.org/10.3391/mbi.2013.4.1.09 %R 10.3391/mbi.2013.4.1.09 %0 Journal Article %J Journal of Invertebrate Pathology %D 2013 %T Pseudomonas fluorescens strain CL145A A Biopesticide for the Control of Zebra and Quagga Mussels (Bivalvia: Dreissenidae) %A D. P. Molloy %A D. A. Mayer %A Gaylo, M. J. %A Burlakova, L. E. %A Karatayev, A. Y. %A Presti, K. T. %A Sawyko, P. M. %A Morse, J. T. %A Paul, A. T. %K Biological control %K Biopesticide %K quagga mussel %K Zebra mussel %K Zequanox® %X

Zebra mussels (Dreissena polymorpha) and quagga mussels (Dreissena rostriformis bugensis) are the “poster children” of high-impact aquatic invasive species. In an effort to develop an effective and environmentally acceptable method to control their fouling of raw-water conduits, we have investigated the potential use of bacteria and their natural metabolic products as selective biological control agents. An outcome of this effort was the discovery of Pseudomonas fluorescens strain CL145A – an environmental isolate that kills these dreissenid mussels by intoxication (i.e., not infection). In the present paper, we use molecular methods to reconfirm that CL145A is a strain of the species P. fluorescens, and provide a phylogenetic analysis of the strain in relation to other Pseudomonas spp. We also provide evidence that the natural product lethal to dreissenids is associated with the cell wall of P. fluorescens CL145A, is a heat-labile secondary metabolite, and has degradable toxicity within 24 h when applied to water. CL145A appears to be an unusual strain of P. fluorescens since it was the only one among the ten strains tested to cause high mussel mortality. Pipe trials conducted under once-through conditions indicated: (1) P. fluorescens CL145A cells were efficacious against both zebra and quagga mussels, with high mortalities achieved against both species, and (2) as long as the total quantity of bacterial cells applied during the entire treatment period was the same, similar mussel mortality could be achieved in treatments lasting 1.5–12.0 h, with longer treatment durations achieving lower mortalities. The efficacy data presented herein, in combination with prior demonstration of its low risk of non-target impact, indicate that P. fluorescens CL145A cells have significant promise as an effective and environmentally safe control agent against these invasive mussels.

%B Journal of Invertebrate Pathology %V 113 %P 104-114 %G eng %U http://dx.doi.org/10.1016/j.jip.2012.12.012 %R 10.1016/j.jip.2012.12.012 %0 Journal Article %J Parasitology %D 2012 %T Haplosporidium raabei n. sp. (Haplosporidia): A Parasite of Zebra Mussels, Dreissena polymorpha (Pallas, 1771) %A D. P. Molloy %A L. Giamberini %A Stokes, N. A. %A Burreson, E. M. %A M. A. Ovcharenko %K Dreissena polymorpha %K Haplosporidia %K Haplosporidium raabei n. sp. %K phylogeny %K small subunit ribosomal DNA %X

Extensive connective tissue lysis is a common outcome of haplosporidian infection. Although such infections in marine invertebrates are well documented, they are relatively rarely observed in freshwater invertebrates. Herein, we report a field study using a comprehensive series of methodologies (histology, dissection, electron microscopy, gene sequence analysis, and molecular phylogenetics) to investigate the morphology, taxonomy, systematics, geographical distribution, pathogenicity, and seasonal and annual prevalence of a haplosporidian observed in zebra mussels, Dreissena polymorpha. Based on its genetic sequence, morphology, and host, we describe Haplosporidium raabei n. sp. from D. polymorpha – the first haplosporidian species from a freshwater bivalve. Haplosporidium raabei is rare as we observed it in histological sections in only 0·7% of the zebra mussels collected from 43 water bodies across 11 European countries and in none that were collected from 10 water bodies in the United States. In contrast to its low prevalences, disease intensities were quite high with 79·5% of infections advanced to sporogenesis.

%B Parasitology %V 139 %P 463-477 %G eng %U http://dx.doi.org/10.1017/S0031182011002101 %R 10.1017/S0031182011002101 %0 Book Section %B The Zebra Mussel in Europe %D 2010 %T Investigation of the endosymbionts of Dreissena stankovici with Morphological and Molecular Confirmation of Host Species %A D. P. Molloy %A L. Giamberini %A Burlakova, L. E. %A Karatayev, A. Y. %A J. R. Cryan %A Trajanovski, S. L. %A Trajanovska, S. P. %E van der Velde, G. %E Rajagopal, S. %E bij de Vaate, A. %K Dreissena %K endosymbionts %K Lake Ohrid %K mussels %K Republic of Macedonia %X

We investigated the types of endosymbionts present in Dreissena mussels in Lake Ohrid, Republic of Macedonia,
and characterized their intensity, prevalence, location, and host impact. Considering the taxonomic uncertainty
of many Balkan dreissenid populations, accurate host identity was considered a very high priority. Our
efforts using both morphological and molecular approaches confirmed the identity of all mussels examined to
be Dreissena stankovici. Key external shell characteristics of D. stankovici are its possession of a: 1) a carina,
2) a convex posterior ventral surface, and 3) a sharp dorsal longitudinal ridge that is distinctively-peaked in
most specimens and slightly convex to almost straight immediately posterior to its peak. Nucleotide sequence
data were successfully generated for 67 Lake Ohrid Dreissena. For each mussel, sequences were obtained for at
least 1 of the following 3 DNA loci: mitochondrial gene 16S rRNA, nuclear gene 28S rRNA, and mitochondrial
gene cytochrome c oxidase subunit I (COI). Remarkably little intraspecific variation was observed in our data
matrices for these 3 gene regions, and we interpreted this, in combination with BLAST searches of the NCBI
database, as strong evidence that the specimens examined were all conspecific D. stankovici. Specimens of
Dreissena polymorpha and Dreissena rostriformis bugensis were used for comparison purposes in both morphological
and molecular analyses.
This is the first study documenting the endosymbiont community in D. stankovici. This dreissenid population in
Lake Ohrid appears to have a rich endosymbiont assemblage, ranging from commensals to parasites, and is comprised
primarily of ciliates (Conchophthirus, Hypocomagalma, Ophryoglena, and Sphenophrya), trematodes
(Echinostomatidae and Phyllodistomum), and intracytoplasmic prokaryote infections. One of the most interesting
findings of our study was how similar this endosymbiont assemblage was to that reported from European
D. polymorpha populations. This perhaps should not be unexpected, however, since D. stankovici and D. polymorpha
are considered sister taxa with divergence estimated at only several million years ago.

%B The Zebra Mussel in Europe %I Backhuys Publishers %C Leiden %P 227-237 %G eng %U https://scholar.google.com/scholar?hl=en&q=Investigation+of+the+endosymbionts+of+Dreissena+stankovici+with+Morphological+and+Molecular+Confirmation+of+Host+Species&btnG=&as_sdt=1%2C33&as_sdtp= %0 Journal Article %J Biological Invasions %D 2007 %T Discovery of Dreissena rostriformis bugensis (Andrusov 1897) in Western Europe %A D. P. Molloy %A bij de Vaate, A. %A Wilke, T. %A L. Giamberini %K biology %B Biological Invasions %V 9 %P 871-874 %G eng %U http://dx.doi.org/10.1007/s10530-006-9078-5 %0 Journal Article %J Diseases of Aquatic Organisms %D 2005 %T Ophryoglena hemophaga n. sp. (Ciliophora: Ophryoglenidae): A Parasite of the Digestive Gland of Zebra Mussels Dreissena polymorpha %A D. P. Molloy %A Lynn, D. H. %A L. Giamberini %K Contractile vacuoles %K Palintomy %K Phototaxis %K Theront %K Tomont %K Trophont %K Zebra mussel %X

Ophryoglena hemophaga n. sp. is described from a freshwater Dreissena polymorpha population in the Rhine delta of the Netherlands. This is the first ophryoglenine species (order Hymenostomatida, suborder Ophryoglenina) recorded as a molluscan parasite. As is typical of ciliates in the suborder Ophryoglenina, O. hemophaga exhibits a polymorphic life history with cystment and reproduction by palintomy. Trophonts were observed within digestive gland lumina, and zebra mussel hemocytes were present in some of their digestive vacuoles. The presence of a single, longitudinal tract of multiple contractile vacuoles represents its most unique feature and distinguishes it from all other described Ophryoglena spp. The number of somatic kineties of O. hemophaga (range 50 to 62) is also a distinguishing feature, since it is the lowest described from any Ophryoglena sp. Other characteristics of this species include: ovoid to elongate trophonts 96 to 288 µm in length, with an elongate macronucleus 41 to 65 µm in length; tomonts 50 to 150 µm in diameter producing a clear mucous cyst envelope, whose thickness is approximately half of the tomont diameter; elongated theronts 96 to 131 µm in length which emerge after 1 to 3 cell divisions taking 36 to 48 h at 20 ± 3°C. Protomonts and theronts are, respectively, negatively and positively phototactic—characteristics that likely aid in maintenance of infection in zebra mussel populations.

%B Diseases of Aquatic Organisms %V 65 %P 237-243 %G eng %U http://dx.doi.org/10.3354/dao065237 %R 10.3354/dao065237 %0 Magazine Article %D 2004 %T Building International Bridges of Understanding and Collaborative Science %A D. P. Molloy %K biology %B Members Update %V 15 %P 6-7 %G eng %0 Report %D 2004 %T A Method for Controlling Dreissena Species %A D. P. Molloy %K biology %C Ottawa %G eng %0 Journal Article %J Diseases of Aquatic Organisms %D 2001 %T Characterization of Intracytoplasmic Prokaryote Infections in Dreissena sp. (Bivalvia: Dreissenidae) %A D. P. Molloy %A L. Giamberini %A Morado, J. F. %A Fokin, S. I. %A Laruelle, F. %K Chlamydiales-like %K Connective tissue %K Epithelium %K Intracytoplasmic prokaryote %K Rickettsiales-like %K Virus-like particles %K Zebra mussels %X

This study characterizes intracytoplasmic infections with prokaryote microorganisms in Dreissena sp. (near Dreissena polymorpha) from northeastern Greece and represents the first report of such infections in freshwater bivalves. Light microscope observations of stained tissues revealed basophilic, cytoplasmic inclusion bodies in 87.5% (28/32) of the mussels sectioned. Inclusions in epithelial cells and connective tissues were noted, respectively, in 34.4 and 71.9% of the sample, with 5 mussels (15.6%) having both tissue types infected. Epithelial cell infections were observed in histological sections only in digestive gland tubules and ducts; within tubules, inclusions were present more often in secretory than digestive cells. Connective tissue infections, however, were systemic; among the 32 mussels sectioned, inclusions were found in the gills (65.6%), foot (12.5%), mantle (9.4%), labial palps (6.3%), digestive gland (6.3%), stomach (6.3%), and gonads (3.1%). Cytoplasmic inclusions (maximum dimension, 138 microm) were prominent enough in the gills to be visible in 17.0% of the 247 mussels dissected. Ultrastructurally, prokaryote cells in gill connective tissues were clearly characteristic of Chlamydiales-like organisms, with each intracytoplasmic inclusion containing a loosely packed mixture of elementary, reticulate, intermediate bodies, and blebs. Prokaryote colonies in digestive gland epithelial cells exclusively contained 1 of 4 morphological cell types and were considered Rickettsiales-like. Hexagonal, virus-like particles were present in the cytoplasm of the largest of these Rickettsiales-like prokaryotes. Although host stress was evident from localized cell necrosis and dense hemocyte infiltration, overall infection was fairly benign, with no major, adverse impact on body condition evident among sectioned or dissected mussels. A possible negative effect was partial constriction of gill water tubes, but at the infection intensity observed (typical range 1 to 7 inclusion bodies per section), significant interference with respiration and other metabolic functions of the gills was highly unlikely.

%B Diseases of Aquatic Organisms %V 44 %P 203-216 %G eng %U http://dx.doi.org/10.3354/dao044203 %R 10.3354/dao044203 %0 Report %D 2001 %T A Method for Controlling Dreissena Species %A D. P. Molloy %K biology %C Washington, D. C. %G eng %0 Report %D 2000 %T Zebra Mussel Information System Compact Disc. 2nd Edition %A D. P. Molloy %A Crosier, D. M. %K biology %C Vicksburg, Mississippi %G eng %0 Journal Article %J Journal of Invertebrate Pathology %D 1999 %T New North American Records of Aquatic Insects as Paratenic Hosts of Pheromermis (Nematoda: Mermithidae) %A D. P. Molloy %A Vinikour, W. S. %A Anderson, R. V. %K Amphinemura delosa %K Clioperla clio %K Diplectrona modesta %K Glossosoma intermedium %K Hesperophylax designatus %K Lepidostoma liba %K Pheromermis pachysoma %K Pheromermis vesparum %K prevalence and intensity of infection %X

Several species of aquatic insects in Trout Park Nature Preserve (Elgin, IL) were observed to have small, black spots (<0.1 mm diameter) visible within their bodies. Microscopic examination revealed these spots to be coiled juveniles of a mermithid (Nematoda: Mermithidae). Based on host habitat (seepage areas and rivulets), host species (aquatic insects), and size (mean diameter of coiled juveniles = 79 μm), it is likely that these mermithids were in the genus Pheromermis. Since adult mermithids were not found, species determination was not feasible, and the possibility of a new species cannot be ruled out. Pheromermis pachysoma and Pheromermis vesparum, however, are two species known to use aquatic insects as paratenic (i.e., transport) hosts in order to reach their definitive hosts, vespid wasps. Wasp larvae are infected by consuming the flesh of adult aquatic insects that contain the coiled juveniles of these Pheromermis spp. Of the 19 macroinvertebrate species examined in this study, Pheromermis juveniles were found in 4 caddisfly species (Hesperophylax designatus, Lepidostoma liba, Glossosoma intermedium, and Diplectrona modesta) and in 2 stonefly species (Clioperla clio and Amphinemura delosa). In addition to all 6 insect species being new host records for Pheromermis infection, this also represents the first report of nematode infection in stoneflies within the Western Hemisphere and of a Pheromermis sp. in Illinois. Among trophic groups, insect detritivores have been frequently recorded infected with coiled Pheromermis juveniles because of their direct consumption of eggs, and we also observed this for detritivores in our investigation (e.g., L. liba and A. delosa). Because C. clio was intensively infected, however, our study also provided evidence that predatory insects can be paratenic hosts. Coiled juveniles were typically present in muscle and fat body and present in almost all body regions. Not every infected paratenic host had external signs of infection; thus, dissections are required for accurate determination of infection prevalence and intensity. Our findings, in conjunction with those made in previous studies, indicate that a wide variety of aquatic insects may be used as paratenic hosts by Pheromermis. Because of their small size, it is highly likely that coiled juveniles are either overlooked or mistaken for melanized integumental wounds during field studies of aquatic insects. A more careful inspection for these coiled juveniles in aquatic insects, especially detritivores and their predators in seepage areas, would probably reveal that Pheromermis is far more common than presently documented.

%B Journal of Invertebrate Pathology %V 72 %P 89-95 %G eng %U http://dx.doi.org/10.1006/jipa.1999.4860 %R 10.1006/jipa.1999.4860 %0 Journal Article %J Journal of Shellfish Research %D 1998 %T The Potential for Using Biological Control Technologies in the Management of Dreissena spp. %A D. P. Molloy %K benthic competitors %K natural enemies %K Parasites %K predators %K quagga mussels %K selectively toxic microorganisms %K Zebra mussels %X

Broadly defined, biological control is the use of one species for the suppression of another. Two groups of organisms have potential as Dreissena control agents: selectively toxic microbes and natural enemies. Selectively toxic microbes are naturally occurring soil and water microorganisms that simply by chance happen to be toxic to Dreissena. Applied at artificially high densities to water, the microbial cells per se or their released metabolites are selectively lethal to Dreissena. In contrast, natural enemies are organisms that, by their evolutionary nature, will debilitate or kill Dreissena, including predators, parasites (both multicellular and

microbial), and benthic competitors (organisms capable of competitively displacing Dreissena from substrates).

Selectively Toxic Microbes: In the short run, these microorganisms represent the most promising candidates as biological control agents, The use of highly-specific, toxic microbes has a clear record of commercial success and environmental safety in the control of invertebrate pests in North America. and strains lethal to Dreissena have been already isolated. Considering how quickly a selectively toxic microbe could move from the laboratory to commercialization, continued research to develop this microbial control strategy is a high priority.

Natural Enemies; In North America, as in Eurasia, there will likely be isolated field reports of major impacts by natural enemies, and, on the whole, we will likely see a cumulative effect of a complex of enemies having a constant, but limited, role in naturally suppressing Dreissena populations. In the majority of cases, Dreissena populations will cause economic and ecological effects at densities well below those that their enemies could naturally maintain. This does not mean, however. that certain natural enemies could not be artificially manipulated (e.g., mass produced and released in an inundative fashion) to cause major Dreissena reductions.

1. Parasites: These natural enemies would likely be the most environmentally-safe biocontrol agents, due primarily to their high host specificity. Comprehensive investigations to identify parasites that are strictly specific to Dreissena and that can be economically mass produced are a key research priority. Parasites from Dreissena’s native Eurasian range should be studied to identify species that would be “nearly risk free” candidates for importation into North America. Research to uncover parasites of Dreissena that are already present in North America should be intensified.

2. Predators: Although organisms, such as fish and birds, have sometimes been documented to consume Dreissena at high rates, this does not necessarily mean that they would be effective candidates for use in control programs. Predators are typically not specific enough in their prey choices. A predator introduced to a waterbody from outside its natural range may highly prefer Dreissena as a prey item, but will also consume other aquatic organisms. The consumption of such nontarget prey could potentially have serious, adverse, ecological impacts.

3. Benthic Competitors; In addition to being micro/macrofouling organisms themselves, the lack of specificity of species that can competitively displace Dreissena from substrates (e.g., sponges) significantly reduces their potential as biocontrol agents.

%B Journal of Shellfish Research %V 17 %P 177-183 %G eng %0 Journal Article %J Reviews in Fisheries Science %D 1997 %T Natural Enemies of Zebra Mussels: Predators, Parasites, and Ecological Competitors %A D. P. Molloy %A Karatayev, A. Y. %A Burlakova, L. E. %A Kurandina, D. P. %A Laruelle, F. %K Birds %K ciliates %K Dreissena %K Fish %K sponges %K trematodes %X

This paper reviews the international literature on the natural enemies of Dreissena spp. and discusses the biology and ecology of organisms known to be involved in their predation (176 species), parasitism (34 species), and competitive exclusion (10 species). Research on natural enemies, both in Europe and North America, has focused on predators, particularly birds (36 species) and fish (15 and 38 species eating veligers and attached mussels, respectively). Other field‐documented predation includes consumption of pelagic larvae by copepods and coelenterates, and consumption of attached mussels by leeches, crabs, crayfish, and rodents. Cannibalism of veligers by adult zebra mussels has also been reported. Ciliates and trematodes are the most commonly reported obligate parasites, with occasional records of suspected bacterial or ascetosporan infection. Mites, nematodes, leeches, chironomids, and oligochaetes have been observed to be associated symbiotically within the mantle cavity, but with few to no adverse effects. Organisms capable of competitively displacing zebra mussels from hard substrates include sponges, amphipods, algae, bryozoans, hydrozoan coelenterates, and other bivalve species (including interspecific competition among Dreissena spp.).

Although the vast majority of the organisms that are natural enemies in Europe are not present in North America, ecologically similar species do exist on this continent, and zebra mussels represent a novel and abundant organism for these native predators, parasites, and ecological competitors — the new natural enemies of Dreissena. However, the idea that these organisms could eliminate zebra mussel populations, even in limited areas of North America, is far more hopeful than realistic. As in Europe, there will likely be isolated reports of major impacts by natural enemies, and on the whole we will likely see a cumulative effect of a suite of enemies having a constant, but limited, role in suppressing zebra mussel populations.

%B Reviews in Fisheries Science %V 5 %P 27-97 %G eng %U http://dx.doi.org/10.1080/10641269709388593 %R 10.1080/10641269709388593 %0 Journal Article %J Journal of Helminthological Society of Washington %D 1996 %T Survey of the Parasites of Zebra Mussels (Bivalvia: Dreissenidae) in Northwestern Russia, With Comments on Records of Parasitism in Europe and North America %A D. P. Molloy %A Roitman, V. A. %A Shields, J. D. %K biology %B Journal of Helminthological Society of Washington %V 63 %P 251-256 %G eng %0 Journal Article %J Dreissena! (Zebra Mussel Information Clearinghouse Newsletter, Brockport, NY) %D 1994 %T Doing Zebra Mussel Research in Russia %A D. P. Molloy %K biology %B Dreissena! (Zebra Mussel Information Clearinghouse Newsletter, Brockport, NY) %V 5 %P 13-14 %G eng %0 Journal Article %J Journal of Shellfish Research %D 1994 %T Short-term Reduction of Adult Zebra Mussels, Dreissena polymorpha, in the Hudson River Near Catskill, New York: An Effect of Juvenile Blue Crab (Callinectes Sapidus) Predation? %A D. P. Molloy %A Powell, J. %A Ambrose, P. %K Biological control %K blue crabs %K predation %K Zebra mussels %X

During the summer of 1992, a loss of 2- to 3-cm zebra mussels, Dreissena polymorpha, was recorded in the Hudson
River near Catskill, NY. We document this adult population crash and present field and laboratory data suggesting that predation by
blue crabs, Callinectes sapidus, was responsible. Dredge samples on July 8th indicated that the mean density of the largest zebra
mussel size class (length = 2 to 3 cm) in the Hamburg-Catskill-Germantownregion was 664 m-2 (range of 59 to 2,222 m-2, 14rucks
examined). In sharp contrast, extensive dredging on August II th did not recover a single mussel of that size class in that area. The
largest mussel collected on August 11th was only 0.7 em long, with over 100 rocks examined. An abundance of shell fragments and
detached, live mussels were considered evidence of predation, possibly by a large immigrant blue crab population present in the area.
Laboratory trials confirmed that these crabs can aggressively consume zebra mussels, particularly the size class that was virtually
eliminated in the Catskill area, and that their feeding does result in piles of shell fragments and removed, yet unconsumed mussels.
Further supportive evidence of the blue crab predation hypothesis was gained in the summers of 1993 and 1994, when blue crabs did
not migrate into the Catskill area and a massive decline in 2 to 3 cm mussels did not occur.

%B Journal of Shellfish Research %V 13 %P 367-371 %G eng %U http://www.academia.edu/download/34389015/Molloy_et_al._1994.pdf %0 Journal Article %J Proceedings of the IVth International Colloquium on Invertebrate Pathology and Microbial Control %D 1994 %T Technology of Application of Microbials in Aquatic Environments: Use of BTI Against Blackflies in Mountainous Areas %A D. P. Molloy %K biology %B Proceedings of the IVth International Colloquium on Invertebrate Pathology and Microbial Control %V 1 %P 479-480 %G eng %0 Journal Article %J Proceedings of the IVth International Colloquium on Invertebrate Pathology and Microbial Control (Montpellier, France) %D 1994 %T Zebra Mussels (Dreissenidae): Review of Parasitic and Other Symbiotic Organisms Associated With These European Biofouling Bivalves %A D. P. Molloy %K biology %B Proceedings of the IVth International Colloquium on Invertebrate Pathology and Microbial Control (Montpellier, France) %V 1 %P 473-474 %G eng %0 Journal Article %J Journal of Shellfish Research %D 1993 %T Approaches to the Biological Control of Zebra Mussels %A D. P. Molloy %K Biological control %K Zebra mussels %B Journal of Shellfish Research %V 12 %P 147 %G eng %0 Journal Article %J Journal of Shellfish Research %D 1992 %T Biological Control of Zebra Mussels: Screening for Lethal Microorganisms %A D. P. Molloy %A Griffin, B. %K biology %B Journal of Shellfish Research %V 11 %P 234 %G eng %0 Magazine Article %D 1992 %T Do Zebra Mussels Have Parasites? %A D. P. Molloy %K biology %B DpIR (Zebra Mussel Information Clearinghouse Newsletter, Brockport, NY) %V 3 %P 7-8 %G eng %0 Journal Article %J Journal of the American Mosquito Control Association %D 1992 %T Impact of the Black Fly (Diptera: Simuliidae) Control Agent Bacillus thuringiensis var. israelensis on Chironomids (Diptera: Chironomidae) and Other Nontarget Insects: Results of Ten Field Trials %A D. P. Molloy %K biology %X

Except for moderate mortality among fiIter-feeding chironomids, Rheotanytarsus spp., the results of 10 field trials with Bacillus thuringiensis var. israelcnsis (B.t.i.) indicated a wide margin of safety to the chironomid community and other stream nontarget insects. Mayflies, caddisflies and 2 other types of chironomids, i.e., tube-dwelling (Chironominae) and surface-dwelling,c aselessla rvae (mix of Chironominae, Diamesinae, Orthocladiinae, Tanypodinae), did not appear to be affected. The susceptibility of filter-feeding chironomids was considerably less than black flies; for example, 4 months of data collected during an operational black fly control program indicated a mean (+95% Cl) mortality among Rheotanytarsus larvae of 23(15-32)% vs. 98(97-99)% among black flies. Although clearly demonstrating the potential of adverse impact on filter-feeding chironomids in operational black fly programs,
these trials also confirmed the narrow impact of B.t.i. on the overall stream insect community.

%B Journal of the American Mosquito Control Association %V 8 %P 24-31 %G eng %U http://www.biodiversitylibrary.org/content/part/JAMCA/JAMCA_V08_N1_P024-031.pdf %0 Journal Article %J American Entomologist %D 1992 %T Review of: Crosskey, R. W. The Natural History of Blackflies %A D. P. Molloy %K biology %B American Entomologist %V 38 %P 45 %G eng %0 Journal Article %J Journal of Shellfish Research %D 1991 %T Biological Control of Zebra Mussels: Use of Parasites and Toxic Microorganisms %A D. P. Molloy %K biology %B Journal of Shellfish Research %V 10 %P 260 %G eng %0 Book Section %B Bacterial Control of Mosquitoes and Black Flies: Biochemistry, Genetics & Applications of Bacillus thuringiensis israelensis and Bacillus sphaericus. %D 1990 %T Progress in the Biological Control of Black Flies With Bacillus thuringiensis var. israelensis, With Emphasis on Temperate Climates %A D. P. Molloy %E de Barjac, H. %E Sutherland, D. %K biology %B Bacterial Control of Mosquitoes and Black Flies: Biochemistry, Genetics & Applications of Bacillus thuringiensis israelensis and Bacillus sphaericus. %I Rutgers University Press %C New Brunswick, New Jersey %P 161-186 %G eng %0 Journal Article %J Bulletin of the Society of Vector Ecology %D 1989 %T Guide to Recent Literature on Black Flies (Diptera: Simuliidae): Part II %A D. P. Molloy %A Molloy, L. %K biology %B Bulletin of the Society of Vector Ecology %V 14 %P 135-218 %G eng %0 Journal Article %J Bulletin of the Society of Vector Ecology %D 1989 %T Investigation of the Feasibility of Microbial Control of Black Flies (Diptera: Simuliidae) with Bacillus thuringiensis var. israelensis in the Adirondack Mountains of New York %A D. P. Molloy %A Struble, R. H. %K biology %X

A two-year study in the Adirondack Mountains of New York State demonstrated that a groundapplied, larvicide program using Bacillus lhuringiensis var. israelensis (Bli) was a technically feasible method for the area control of black flies. The program required 102 discharge measurements and 480 applications ofcommercial formulations of Bli at dosages equivalent to 10 to 30 ppmfor 1min. Treatments were made to all streams within a 3.2-1an radius of the hamlet of Onchiota between April 1st and July 15th, 1983. The larviciding program was successful, with stream samples(n= 131) indicating a mean larval mortality(± 95% CI) of97.8% (96.9-98.5%). To measure adult activity, 1,008 and 168 one-min, insect net sweeps were made, respectively, in the 32.2 Jan2study area and at a distant control site between May 15th and July 31stboth in 1982 (pretreatment year) and 1983. Analysis of these data (39,766 adults in total) indicated that thelarviciding program resulted in statistically significant seasonal reductions in adul tblack flies: 85.8 percent at the center of the Onchiota community and 58.1 percent at its perimeter, i.e., at 1.6 Ian from the center. Thus, the Bti program achieved an average seasonal reduction of 72.0 percent in the target community. As expected, the largest adult reductions in all three genera, i.e., Prosimulium, Simulium, and Stegopterna, were recorded at the center of the community. These results suggest that in similar habitats adult black fly reductions of ca. 86 percent and 58 percent can be achieved at points 3.2 Ian and 1.6 Ian from untreated areasirrespective of the size of the total larvicided area that a community might decide to establish ..

For communities interested in a Bli - black fly.control program in similar habitats, a first-year base cost of ca. $400/km2 of treated area can be expected. This represents the sum of the preparatory expenses of $70-90/km2•which would be unnecessary in subsequent years, and the actual treatmentexpcnses of$240-380/km2• which would likely decrease after the initial year due to increased efficiency of operation. Equipment puochases. liability insurance. training, etc .•would be in addition to this base cost. Labor would be the primary expense, with the cost of purchasing B Ii formulations less than 5 peocentofthe total budget. Such prices will likely make grourId-appliedcontrol programs with Bli economically feasible in the Adirondack Mountains of New York, since the cost for aerial application of broad spectrum insecticides for adult black fly control is considerably higher.

%B Bulletin of the Society of Vector Ecology %V 14 %P 266-276 %G eng %0 Journal Article %J Bulletin of the Society of Vector Ecology %D 1988 %T Guide to Recent Literature on Black Flies (Diptera: Simuliidae): Part I %A D. P. Molloy %K biology %B Bulletin of the Society of Vector Ecology %V 13 %P 126-220 %G eng %0 Journal Article %J Journal of Freshwater Ecology %D 1988 %T A Simple and Inexpensive Method for Determining Stream Discharge From a Streambank %A D. P. Molloy %A Struble, R. H. %K biology %X

A detailed protocol for the determination of stream discharge is presented which requires neither a current meter nor actual entry into streamwater. The protocol represents a modification of the flotation method of discharge determination and was devised in response to the need for a simple, inexpensive, “bank-side” technique for estimation of stream flow. Discharge is determined through measurements of a stream's cross-sectional area and surface velocity. In tests at high and low flows, mean discharges determined by the proposed flotation method were not significantly different from those calculated using a standardized, current-meter method. This suggested that me proposed flotation method (a 10–20 min procedure) could be used with reasonable accuracy to determine discharge in small to moderate-sized streams.

%B Journal of Freshwater Ecology %V 4 %P 477-481 %G eng %U http://www.tandfonline.com/doi/abs/10.1080/02705060.1988.9665197 %R 10.1080/02705060.1988.9665197 %0 Book Section %B Black Flies: Ecology, Population Management, and Annotated World List %D 1987 %T The Ecology of Black Fly Parasites %A D. P. Molloy %E Kim, K. C. %E Merritt, R. W. %K biology %X

The parasites of Simuliidae and their ecology are discussed in the following framework: (1) Aspects of the host-parasite system (approaches to invading the host; commencement of infection; maintenance of stability of parasitic and black fly populations; host specificity, including a table summarizing ranges of mermithid, fungal, protozoal and viral parasites among simuliid genera); (2) Physiological host-parasite interactions (location of infection within or on simuliid hosts, with a table listing this for protozoal, fungal, viral and mite parasites; effects on the black fly by the parasite, also summarized in a table; effect of the black fly on the parasite); (3) Dispersal of parasites; (4) Alternate hosts; and (5) Conclusions.

%B Black Flies: Ecology, Population Management, and Annotated World List %I Penn. State Univ. Press %C University Park, Pennsylvania %P 315-326 %G eng %0 Journal Article %J Journal of the North American Benthological Society %D 1987 %T Review of: The Black Flies of Pennsylvania (Simuliidae, Diptera): Bionomics, Taxonomy, and Distribution, P. H. Adler and K. C. Kim. %A D. P. Molloy %K biology %B Journal of the North American Benthological Society %V 6 %P 79 %G eng %0 Book Section %B Current and Selected Bibliographies on Benthic Biology, 1984 %D 1985 %T Diptera (Simuliidae) %A D. P. Molloy %A Hannon, B. H. %E Webb , D. W. %K biology %B Current and Selected Bibliographies on Benthic Biology, 1984 %I North American Benthology Society, Illinois Natural History Survey %C Champaign, Illinois %P 33-40 %G eng %0 Magazine Article %D 1984 %T The Black Fly Debate %A D. P. Molloy %K biology %B Naho %V 17 %P 7-10 %G eng %0 Book Section %B Current and Selected Bibliographies on Benthic Biology, 1982 %D 1984 %T Diptera - Simuliidae %A D. P. Molloy %A Hannon, B. H. %E Webb , D. W. %K biology %B Current and Selected Bibliographies on Benthic Biology, 1982 %I North American Benthological Society and Illinois Natural History Survey %C Champaign, Illinois %P 48-57 %G eng %0 Journal Article %J Journal of Agricultural Entomology %D 1984 %T Laboratory Evaluation of Commercial Formulations of Bacillus thuringiensis var. israelensis Against Mosquito and Black Fly Larvae. %A D. P. Molloy %A Wraight, S. P. %A Kaplan, B. %A Gerardi, J. %A Petersen, P. %K biology %X

Three   commercial  black   fly  and  mosquito   larvicidal   formulations   of  Bacillus

thuringiensis Berliner var. israelensis de Barjac were bioassayed to determine their potencies relative to the international standard powder, IPS-78. The potencies of the wettable powder (WP) formulations of Bactimos™ and Vectobac™ and the water dispersible concentrate (WDC) TeknarN were determined to be 4530, 5723, and  336 International Toxic Units (ITU)/mg, respectively, against Simulium spp.; their respective ITU/mg values against Aedes aegypti (Linnaeus) were 3556, 2317, and 1373. Little correlation (r2 = 0.47) was evident between the potency rating (ITU/mg) of the formulations against mosquitoes and black flies; this indicated that the potency rating of a formulation against black flies cannot be simply predicted from the A. aegypti ITU/mg rating on its label. A moderate correlation between formulation potency and mean particle size was observed in the black fly assays (r2 = 0.77), but not against mosquitoes (r2 = 0.20). Aqueou s suspensions of Teknar-WDC had a significantly smaller mean particle size and stayed significantly longer in suspension than other formulations. The relatively long duration of suspension of Teknar-WDC was viewed as important since the toxic crystals wou ld be less likely to settle out in mosquito and black fly habitats. A ratio based on the LC50 values determined from standardized bioassay methods is suggested as a convenient way to express the relative  potency of a formulation against mosquitoes and black flies.

 

%B Journal of Agricultural Entomology %V 1 %P 161-168 %G eng %U http://scentsoc.org/Volumes/JAE/v1/2/00012161.pdf %0 Book %D 1982 %T Biological Control of Black Flies (Diptera: Simuliidae) with Bacillus thuringiensis var. israelensis (Serotype 14): A Review with Recommendations for Laboratory and Field Protocol %A D. P. Molloy %K biology %I Entomological Society of America Miscellaneous Publications 12(4):30 pp %C , %G eng %0 Journal Article %J Journal of Invertebrate Pathology %D 1982 %T Rediscovery of Erynia aquatica in Aedes snowpool Mosquitoes %A D. P. Molloy %A Wraight, S. P. %K biology %B Journal of Invertebrate Pathology %V 40 %P 142-145 %G eng %U http://www.sciencedirect.com/science/article/pii/0022201182900441 %R 10.1016/0022-2011(82)90044-1 %0 Journal Article %J Journal of Economic Entomology %D 1981 %T Factors Influencing the Efficacy of Bacillus thuringiensis var. israelensis As a Biological Control Agent of Black Fly Larvae (Diptera: Simuliidae) %A D. P. Molloy %A Gaugler, R. R. %A H. Jamnback %K biology %X

In laboratory trials with Bacillus thuringiensis Berliner var. israelensis de Barjac against Simulium larvae, the following factors significantly influenced efficacy: black fly instar and species, formulation, temperature, and product manufacturer.

Mortalities were consistently higher among smaller larvae, e.g., after 15 min exposures with Bellon Laboratory primary powder at 11°C, the LC90 value for 2nd–5th instars (111 ppb) was significantly lower than for 7th instars (615 ppb). S. verecundum Stone and Jamnback larvae were significantly more susceptible than S. vittatum Zetterstedt to Bellon primary powder. In trials comparing Bellon preparations by total product weight, significantly higher mortalities were achieved with primary powder than with other preparations. No difference in efficacy, however, was evident between Bellon WP and primary powder when compared solely on the basis of the weight of their AI (Bellon wettable = 50% AI (wt/wt), while Bellon primary = 100% AI). Both the Bellon aqueous suspension and emulsion were less effective than the WP and primary powder. Results with the emulsion were erratic due to adverse effect on feeding behavior. Microscopic observation of test larvae revealed abnormal clumping of their cephalic fan rays and inhibition of normal feeding during exposure to the emulsion. Mortality among S. vittatum larvae treated at 20°C was twice that recorded at 10°C. Tests with Abbott and Bellon primary powders showed the Bellon preparation to be 3 to 4 times more toxic.

%B Journal of Economic Entomology %V 74 %P 61-64 %G eng %U http://jee.oxfordjournals.org/content/74/1/61.abstract %R 10.1093/jee/74.1.61 %0 Journal Article %J Journal of Economic Entomology %D 1981 %T Field Evaluation of Bacillus thuringiensis var. israelensis as a Black Fly (Diptera: Simuliidae) Biocontrol Agent and Its Effect on Nontarget Stream Insects %A D. P. Molloy %A H. Jamnback %K biology %X

When a small stream was treated at 0.5 ppm/15 min (13°C) with an aqueous suspension of unformulated Bellon primary powder of Bacillus thuringiensis Berliner var. israelensis, 96, 86, 53, and 11% reductions occurred, respectively, in larval populations of Simulium spp. black flies at 20, 180, 350, and 705 m below the treatment point. In contrast, the number of black fly larvae declined by only 5% in the upstream control. Mortality occurred within 2–3 days of treatment, with few dead larvae remaining attached by the 5th day posttreatment. The mortality rates of larvae transported to and reared in the laboratory for 48 h posttreatment closely corresponded to the rates recorded in the stream. Significantly higher mortality rates occurred among smaller larvae. In contrast to the sharp reduction (89%) in black fly larvae in the 20–350 m area below the treatment point, Surber samples indicated increases in mayfly (35%), caddisfly (47%), stonefly (75%), chironomid (19%), and elmid (242%) populations. Thus, no adverse effect on any of these nontarget populations was evident following stream treatment.

%B Journal of Economic Entomology %V 74 %P 314-318 %G eng %U http://doi.org/10.1093/jee/74.3.314 %R 10.1093/jee/74.3.314 %0 Journal Article %J Journal of Nematology %D 1981 %T Mermithid Parasitism of Black Flies (Diptera: Simuliidae) %A D. P. Molloy %K biology %X

Mermithid nematodes are common parasites of black flies and play a significant role in the natural regulation of these medically important insects. Infection levels tend to he moderate and perennial, with epizootics rare and highly localized. Mermithid parasitism almost invariably results in the death of the black fly, and thus considerable attention has focused on the potential of these nematodes as biocontrol agents. Early instar black fly larvae appear most susceptible to infection, and integumental penetration hy mermithid preparasites is the only known mode of entry. Postparasitic nematodes typically emerge before host pupation. However, carryover of parasitism into adult simuliids is an important mechanism for local dispersal and recolonization of upstream areas. Following emergence, the mermithids molt to the adult stage. Copulation ensues, the females then laying eggs which eventually give rise to the next generation of infective preparasites. The number of described species is conservatively estimated at 35-40, with most species within the genera Mesomermis, Gastromermis, and Isomermis. The taxonomy of this group of mermithids is a challenging and little explored area. Host-specificity statements, therefore, must be made cautiously because of these systematic problems and others within the Simuliidae. In most instances, temporal and spatial factors limit the host range of these mermithids among simuliid species. Differential susceptibilities anmng larvae concurrently present within the same microhabitat probably reflect varying degrees of host attractiveness and behavioral-physiological resistance. Effects of parasitism on the host may include prevention of metamorphosis, sterility, intersexual development, and behavior modification. Evaluation of the technical feasibility of mermithid control of black flies has been stymied by the limitations of current inoculum-production technology. Continued advances in in vivo and in vitro culture methods are required to accelerate the research process.

%B Journal of Nematology %V 13 %P 250-256 %G eng %U http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2618114/pdf/250.pdf %0 Journal Article %J Journal of Nematology %D 1980 %T Isolation of Strelkovimermis peterseni, A Mermithid Parasite of Anopheline Mosquitoes %A D. P. Molloy %A Wraight, S. P. %K biology %B Journal of Nematology %V 12 %P 330-332 %G eng %U http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2618027/pdf/330.pdf %0 Journal Article %J Journal of Invertebrate Pathology %D 1980 %T The Pathogenicity of Neoaplectana carpocapsae to Blackfly Larvae %A D. P. Molloy %A Gaugler, R. R. %A H. Jamnback %K biological control of blackflies %K Neoaplectana carpocapsae %K Simulium spp. %X

Laboratory assays indicated that infective-stage juveniles of Neoaplectana carpocapsae are highly pathogenic to Simulium spp. larvae. Instar susceptibility increased with larval size, with early instars being nonsusceptible. High rates of mortality (75 – 100%) were achieved in assays against late instars. These results indicate that N. carpocapsae may have potential value as a blackfly biocontrol agent.

%B Journal of Invertebrate Pathology %V 36 %P 302-306 %G eng %U http://www.sciencedirect.com/science/article/pii/0022201180900397 %R 10.1016/0022-2011(80)90039-7 %0 Journal Article %J Journal of Nematology %D 1979 %T Description and Bionomics of Mesomermis camdenensis n. sp. (Mermithidae), A Parasite of Black Flies (Simuliidae) %A D. P. Molloy %K bionomics %K Mesomermis spp. %K Simulium tuberosum %K taxonomy %X

Mesomermis camdenensis n. sp. is described from larvae of Simulium tuberosum (lundstroem) collected in Camden Valley Creek, Washington County, New York. This species possesses a barrel-shaped vagina, vulval flap. two short separate spicules, terminal mouth, six longitudinal chords, six cephalic papillae, large sexually dimorpbic anaphids, an esophagns of uniform width which extends for less than one-third of the body length, and a cone-shaped tail directed ventrally without appendage. Juveniles also are described and illustrated.

A detailed morphological comparison with the mermithid M. flumenalis Welch is presented. The most pronounced morphological differences between these species are in the shape of the vulva, juvenile tail, and infective stage. Cross-mating trials support the integrity of the new species.

The life cycle of M. camdenensis is closely synchronized with that of its primary host, S. tuberosum larvae. Infected S. tuberosum larvae were first collected in May. Emergence of postparasites from late instars took place from mid-June through mid-October. Sampling data indicate a lower susceptibility to infection among S. venuslum Say larvae.

%B Journal of Nematology %V 11 %P 321-328 %G eng %U http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2617980/ %0 Journal Article %J Mosquito News %D 1977 %T A Larval Black Fly Control Field Trial Using Mermithid Parasites and Its Cost Implications %A D. P. Molloy %A H. Jamnback %K biology %B Mosquito News %V 37 %P 104-108 %G eng %U http://www.biodiversitylibrary.org/content/part/JAMCA/MN_V37_N1_P104-108.pdf %0 Magazine Article %D 1975 %T The Blackfly %A D. P. Molloy %A H. Jamnback %K biology %B The Conservationist %V 29 %P 13 %G eng %0 Journal Article %J Mosquito News %D 1975 %T Laboratory Transmission of Mermithids Parasitic in Black Flies (Simuliidae) %A D. P. Molloy %A H. Jamnback %K biology %B Mosquito News %V 35 %P 337-342 %G eng %U http://www.biodiversitylibrary.org/content/part/JAMCA/MN_V35_N3_P337-342.pdf