John Janovy

Species density distributions as null models for ecologically significant interactions of parasite species in an assemblage

A multiple-kind lottery model is presented for use in determining whether species density distributions in parasite species assemblages reveal regularly occurring species-to-species interactions. The model utilizes a recurrence vector algorithm to rapidly calculate expected frequencies of species per host classes in such assemblages. These calculations have been a computational problem because the probability of a host individual acquiring one species of parasite is not necessarily equal to that of acquiring another species. Thus although the number of possible ways for a host to acquire x parasite species of a possible n is given by the familiar binomial expansion term n![x!(x!(n - x)!], each of these ways can have a different probability. The model is applicable to any system that mimics a multiple-kind lottery in which (1) successes are independent events and (2) it is possible to fail completely to acquire any parasites or their analogs. The algorithm is thus a null model for species density distributions in general. Application of the model is illustrated by host/parasite systems involving snails and trematodes, fish and their protozoan and platyhelminth parasites, and a relatively rich assemblage of parasites in bats.

The Roles of Ecological and Evolutionary Influences in Providing Structure to Parasite Species Assemblages

Parasite species assemblages currently are thought to range from isolationist to interactive, their dynamic properties being related to the number of species and types of hosts involved. The literature contains few experimental tests of this concept, however, and many of the host/parasite systems studied to date are not amenable to experimental manipulation. In this review, the presence of a parasite species, in a sample of host individuals, is considered to be an evolutionary phenomenon, but the parasites population structure is considered to be an ecological one. Studies that allow evaluation of these 2 influences are comparative in nature and include data from a series of homogeneous samples of host populations. A lottery model is presented, in which hosts acquire their assemblages of parasites by Monte Carlo type sampling from multiple kind arrays; the major structuring influence is the relative probability of becoming infected by various parasite species. Claims of parasite species interaction need to be supported by studies showing departures from the predictions of this model. The species density and infraassemblage diversity index distributions are recommended as quantitative tools useful in such work.

A NEW SPECIES OF MYXIDIUM (MYXOSPOREA: MYXIDIIDAE), FROM THE WESTERN CHORUS FROG, PSEUDACRIS TRISERIATA TRISERIATA, AND BLANCHARD'S CRICKET FROG, ACRIS CREPITANS BLANCHARDI (HYLIDAE), FROM EASTERN NEBRASKA: MORPHOLOGY, PHYLOGENY, AND CRITICAL COMMENTS ON AMPHIBIAN MYXIDIUM TAXONOMY

During March 2001–April 2004, 164 adult anurans of 6 species (47 Rana blairi, 35 Rana catesbeiana, 31 Hyla chrysoscelis, 31 Pseudacris triseriata triseriata, 11 Bufo woodhousii, and 9 Acris crepitans blanchardi) from Pawnee Lake, Lancaster County, Nebraska, were surveyed for myxozoan parasites. Of these, 20 of 31 (65%) P. triseriata triseriata and 1 of 9 (11%) A. crepitans blanchardi were infected with a new species of Myxidium. Myxidium melleni n. sp. (Myxosporea) is described from the gallbladder of the western chorus frog, P. triseriata triseriata (Hylidae). This is the second species of Myxidium described from North American amphibians. Mature plasmodia are disc-shaped or elliptical 691 (400–1,375) × 499 (230–1,200) × 23 (16–35) μm, polysporic, producing many disporic pansporoblasts. The mature spores, 12.3 (12.0–13.5) × 7.6 (7.0–9.0) × 6.6 (6.0–8.0) μm, containing a single binucleated sporoplasm, are broadly elliptical, with 2–5 transverse grooves on each valve, and contain 2 equal polar capsules 5.2 (4.8–5.5) × 4.2 (3.8–4.5) μm positioned at opposite ends of the spore. Myxidium melleni n. sp. is morphologically consistent with other members of Myxidium. However, M. melleni n. sp. was phylogenetically distinct from other Myxidium species for which DNA sequences are available. Only with improved morphological analyses, accompanied by molecular data, and the deposit of type specimens, can the ambiguous nature of Myxidium be resolved. Guidelines for descriptions of new species of Myxidium are provided.

Comparative life cycles and life histories of North American Rhabdias spp. (Nematoda: Rhabdiasidae): lungworms from snakes and anurans.

Abstract The present study used experimental infections to compare the life cycles and life histories of 6 Rhabdias spp. infecting snakes and anurans. Free-living development of anuran lungworms was primarily limited to heterogonic reproduction, and females utilized matricidal endotoky exclusively, whereas snake lungworms primarily reproduced homogonically and, when heterogonic reproduction occurred, females used a combination of releasing eggs and matricidal endotoky. Infective snake lungworms survived for longer periods in fresh water compared to anuran worms. Infective anuran lungworms penetrated into the skin of frogs and toads; few infections resulted from per os infections. In contrast, snake lungworms were unable to penetrate skin; instead, infective juveniles penetrated into snake esophageal tissue during per os infections. Despite separate points of entry, anuran and snake lungworms both migrated and developed in the fascia, eventually penetrating into the body cavity of the host. Worms molted to adulthood inside the body cavity and subsequently penetrated into the hosts lungs, where they fed on blood while becoming gravid. Adult lungworm survival varied among lungworm species, but, in general, snake lungworms were longer lived than anuran worms. Anuran lungworms were poorly suited for transmission via transport hosts, whereas snake lungworms were consistently capable of establishing infections using transport hosts. Overall, these observations suggest that snake and anuran lungworms have discrepant life cycles and life history strategies.

Evolutionary Constraints on Population Structure: The Parasites of Fundulus zebrinus (Pisces: Cyprinodontidae) in the South Platte River of Nebraska

Population and community descriptor values (parasites per host, prevalence per parasite species, variance/mean ratios, species density, and diversity indices) for the 7-species parasite community of 61 relatively homogeneous samples of Fundulus zebrinus (Pisces: Cyprinodontidae) in the South Platte River of Nebraska, U.S.A., taken over a 14-yr period, are reported. South Platte River streamflow fluctuates over 2 orders of magnitude on several time scales-monthly, annually, and over multiple year wet-dry cycles. Relatively homogeneous sampling of a single host species with several parasite species provided a system that allowed assessment of the contribution of evolved parasite life cycles to population structure in an everchanging environment. No significant negative species-to-species associations were observed. Species abundance, order of abundance, and diversity were affected most strongly by streamflow, with high water reducing prevalence and abundance of larval trematode parasites. Each parasite species had its characteristic long- and short-term patterns of variation in population descriptor values, with mostly long-term stability superimposed on sometimes extreme short-term fluctuations of descriptor values. The differences in these characteristic patterns were considered products primarily of the evolved life cycle traits and transmission mechanisms operating in the common fluctuating environment. The parasite community as a whole showed resilience, returning to preperturbation diversity following extended periods of high water.

The life cycle of a horsehair worm, Gordius robustus (Nematomorpha: Gordioidea)

Aspects of the life cycle of the nematomorph Gordius robustus were investigated. Gordius robustus larvae fed to Tenebrio molitor (Coleoptera: Tenebrionidae) readily penetrated and subsequently encysted in the posterior portion of the midgut wall. Parasite development was then arrested in each infected beetle. Upon feeding these cysts to laboratory-reared Gryllus firmus, worm larvae developed to adults in the cricket hemocoel. In an additional experiment, G. robustus larvae fed to G. firmus did not develop to adults. These findings are in contrast to other studies that found direct infection to be an alternative mode of infection. This experiment is the first employing a laboratory-maintained stock of hosts to rear nematomorph worms.

EVOLUTIONARY AVENUES FOR, AND CONSTRAINTS ON, THE TRANSMISSION OF FROG LUNG FLUKES (HAEMATOLOECHUS SPP.) IN DRAGONFLY SECOND INTERMEDIATE HOSTS

Metacercariae survival patterns and their distribution in second intermediate odonate hosts were examined for 4 species of frog lung flukes. Surveys of aquatic larvae and recently emerged teneral dragonflies and damselflies indicated that prevalence and mean abundance of Haematoloechus spp. metacercariae were significantly lower in teneral dragonflies than larval dragonflies, while there was no significant difference in prevalence or mean abundance of Haematoloechus spp. metacercariae among larval and teneral damselflies. Experimental infections of dragonflies indicated that metacercariae of Haematoloechus coloradensis and Haematoloechus complexus were located in the head, thorax, and branchial basket of dragonflies, whereas metacercariae of Haematoloechus longiplexus and Haematoloechus parviplexus were restricted to the branchial basket of these hosts. Metacercariae of H. coloradensis, H. complexus, and H. longiplexus infected the head, thorax, and abdomen of damselflies, but these insects were resistant to infection with H. parviplexus. Subsequent metamorphosis experiments on experimentally infected dragonflies indicated that most metacercariae of H. longiplexus were lost from the branchial basket during metamorphosis, but most metacercariae of H. coloradensis, H. complexus, and H. parviplexus survived dragonfly metamorphosis. These observations suggest that the observed ecological host specificity of H. longiplexus in semiterrestrial leopard frogs may be due to few metacercariae of H. longiplexus reaching these frogs in a terrestrial environment. Because of the uncertain validity of Haematoloechus varioplexus as a distinct species from its synonym H. parviplexus, their morphological characters were reevaluated. The morphological data on H. varioplexus and H. parviplexus indicate that they differ in their acetabulum length and width, ovary shape, testes length, and egg length and width. Experimental infections of plains leopard frogs, northern leopard frogs, and bullfrogs with worms from bullfrogs indicate that the synonymy of H. parviplexus with H. varioplexus is not warranted, and that these flukes are distinct species, i.e., H. parviplexus in bullfrogs and H. varioplexus in plains leopard frogs and northern leopard frogs.

Ultrastructure of Interlamellar Henneguya exilis in the Channel Catfish

Ultrastructural aspects of interlamellar Henneguya exilis infections in channel catfish are reported. The plasmodium wall of this form differs from that of other species in that it is composed of two outer unit membranes which give rise to a zone of numerous pinocytic canals. Single-membraned canals appeared to be a stable feature of the wall while double-membraned canals are interpreted as those actively carrying out pinocytosis. Evidence suggests that host cellular cytoplasm as well as interstitial material is taken in by plasmodia. Plasmodium wall integrity, aggregation of parasite ectoplasmic components, numbers of pinocytic canals, and number of mitochondria proximal to the wall vary among different plasmodium profiles and may be related to plasmodium maturity. The parasite causes extensive hyperplasia of basal cells, which in turn replaces most other cell types found in noninfected gill filaments. Cytoarchitectural differences between basal cells of noninfected filaments and basal cells adjacent to plasmodia include significantly shorter microfilament bundles in the latter.

HOST STADIUM SPECIFICITY IN THE GREGARINE ASSEMBLAGE PARASITIZING TENEBRIO MOLITOR

Reciprocal cross-stadia experimental infections were used to demonstrate stadium specificity within the gregarine assemblage parasitizing Tenebrio molitor, the yellow mealworm. Gregarina cuneata, Gregarina polymorpha, and Gregarina steini are characteristic parasites of larval T. molitor. Gregarina niphandrodes is a characteristic parasite of adult T. molitor. Experimental infections were produced in all homologous host-parasite combinations. No infection was produced in heterologous or cross-stadia combinations. This study introduces the concept of separate, distinct parasite niches corresponding to separate life cycle stages and established by known, predictable life cycle events within a single host species.

Developmental Niche Structure in the Gregarine Assemblage Parasitizing Tenebrio molitor

The exogenous developmental niche spaces of Gregarina cuneata and Gregarina polymorpha, parasites restricted to larval Tenebrio molitor, and Gregarina niphandrodes, a parasite restricted to adult T. molitor, were determined along temperature and humidity axes and associated with parasite niche space on the host resource axis. Gregarine gametocysts were incubated at controlled humidity (0, 10, and 20 mm Hg vapor pressure deficit) at 20, 25, 30, and 35 C to determine the environmental limits to survival and sporulation