Horned lizards (Phrynosoma) incapacitate dangerous ant prey with mucus

Horned lizards (Iguanidae, Phrynosomatinae, Phrynosoma) are morphologically specialized reptiles characterized by squat, tank-like bodies, short limbs, blunt snouts, spines and cranial horns, among other traits. They are unusual among lizards in the degree to which they specialize on a diet of ants, but exceptional in the number of pugnacious, highly venomous, stinging ants they consume, especially harvester ants (genus Pogonomyrmex). Like other iguanian lizards, they capture insect prey on the tongue, but unlike other lizards, they neither bite nor chew dangerous prey before swallowing. Instead, they employ a unique kinematic pattern in which prey capture, transport and swallowing are combined into a single feeding stage, apparently leaving the mouth, pharynx, esophagus and stomach vulnerable to bites and stings. Nevertheless, horned lizards consume dozens of harvester ants without harm. We show that their derived feeding kinematics are associated with unique, mucus-secreting pharyngeal papillae that apparently serve to immobilize and incapacitate dangerous ants as they are swallowed by compacting them and binding them in mucus strands. Radially branched esophageal folds provide additional mucus-secreting surfaces the ants pass through as they are swallowed. Ants extracted from fresh-killed horned lizard stomachs are curled ventrally into balls and bound in mucus. We conclude that the pharyngeal papillae, in association with a unique form of hyolingual prey transport and swallowing, are horned lizard adaptations related to a diet of dangerous prey. Harvester ant defensive weapons, along with horned lizard adaptations against such weapons, suggest a long-term, predator-prey, co-evolutionary arms race between Phrynosoma and Pogonomyrmex. J. Exp. Zool. 309A:447-459, 2008. (c) 2008 Wiley-Liss, Inc.     

Seasonal dynamics of entomopathogenic nematodes of the genera Steinernema and Heterorhabditis as a response to abiotic factors and abundance of insect hosts

The seasonal dynamics of entomopathogenic nematodes (EPNs) of the genus Steinernema and Heterorhabditis were studied during one season in meadow and oak wood habitats, in the vicinity of Ceské Budejovice, Czech Republic. The influences of soil temperature, moisture, and abundance of suitable hosts on EPN dynamics were investigated. The host range of these nematodes, in both habitats was also observed. A total of four EPN species were found in both habitats. Steinernema affine was the dominant species both in oak wood and in meadow. Additionally, the oak wood habitat was inhabited by S. kraussei and S. weiseri; the meadow habitat by Heterorhabditis bacteriophora. The mean abundance of total EPN community was 28,000ind./m(2) in oak wood and 11,000ind./m(2) in meadow. The seasonal dynamics of entomopathogenic nematodes in both habitats were characterized by high nematode densities in the beginning of the season, followed by a rapid decrease, and then stabilization. EPN abundances did not show any apparent correlation with soil temperature and moisture, but they were negatively correlated with the abundance of suitable insect hosts. Inter- and intraspecific competition for limited nutrients (hosts) probably played a major role in EPN seasonal dynamics. Broad host range of entomopathogenic nematodes in both habitats was predominantly represented by dipteran and coleopteran larvae. Most common hosts belonged to the families Asilidae, Bibionidae, and Empididae (Diptera), as well as Carabidae and Curculionidae (Coleoptera).     

The Effects of Winter Burning and Grazing on Resources and Survival of Texas Horned Lizards in a Thornscrub Ecosystem

ABSTRACT The ecological effects of land-use practices on reptiles, especially endangered or threatened species, are of conservation and scientific interest. We describe the effects of rotational livestock grazing and prescribed winter burning on resources and survival of the Texas horned lizard (Phrynosoma cornutum) during the summers of 1998 to 2001 in southern Texas, USA. We evaluated survival rates of Texas horned lizards (n = 111) on 6 study sites encompassing 5 different burning and grazing treatments. We also measured indices of cover (i.e., vegetation) and food abundance (i.e., harvester ants [Pogonomyrmex rugosus]). We telemetered and relocated adult lizards daily. We divided the study into 2 seasons, spring (15 Apr–30 Jun) and summer (1 Jul–15 Aug), corresponding to the relative activity of horned lizards. Winter burning provided an increase in food resources and led to increased survival of Texas horned lizards in the second growing season after fire, but grazing-induced changes in vegetation cover reduced survival, likely by increasing lizard vulnerability. Fire and grazing reduced litter and increased bare ground and forb cover but did not affect woody vegetation. Ant activity was greater in burned sites and varied with grazing level, season, and year. Summer survival functions of horned lizards varied by burning treatment, with higher survival observed on burned sites in the second year after burning. Survival rates were ordered from highest in ungrazed sites to lowest in heavily grazed sites. We recognize the limitations of our work resulting from a lack of spatial replication of treatments. However, our mensurative study provides fertile ground for future hypothesis testing regarding the effects of land management on shrubland and grassland reptiles. We propose that future studies focus on the population consequences of variation in burn frequency, burn timing, and grazing intensity.     

Influence of temporal fluctuations in seed abundance on the foraging behaviour of harvester ants (Pogonomyrmex spp.) in the central Monte desert,Argentina

The simultaneous study of the temporal dynamics of foraging behaviour, diet and seed abundance is essential to assess the way in which resources affect the behaviour and ecology of harvester ants. Here, we evaluate how fluctuations in grass seed abundance during three consecutive growing seasons influenced the foraging behaviour and diet of the harvester ants Pogonomyrmex rastratus, P. mendozanus and P. inermis in the central Monte desert, Argentina. Seed abundance of the most consumed grasses varied greatly through ant activity season, and ants altered their foraging behaviour in response to those changes. Foragers spent more time travelling and searching for food, and their foraging trips took longer during the low seed availability season. Foraging distance was very similar among species and, contrary to our expectations, did not vary between seasons. Foraging success of P. rastratus and P. inermis increased during the high availability season. This matched the seasonal pattern of foraging activity, suggesting that colonies may detect seed abundance and regulate their foraging effort with the rate of forager success. Although grass seeds were the main item in the diet of the three species, P. mendozanus, and to a lesser extent P. rastratus, turned more generalist when grass seeds were scarce. In contrast, P. inermis showed a very narrow diet breadth, only harvesting grass seeds in both seasons. Our results indicate the relevance of seed availability on foraging behaviour of harvester ants, which should be taken into account when predicting and evaluating the effect of ants on seed resources as well as numerical responses of harvester ant populations to the temporal and spatial variations in grass seed abundance.     

Phylogeography of the flat-tailed horned lizard (Phrynosoma mcallii) and systematics of the P. mcallii-platyrhinos mtDNA complex

Two species of horned lizards are sympatric along the periphery of the Salton Trough. Phrynosoma mcallii, endemic to the trough, is of conservation concern because its limited habitat has been fragmented by human activities. A more common and widespread species, Phrynosoma platyrhinos, occurs around the periphery of the trough and much further to the North. The two species are syntopic at a few localities, where morphological intermediates have also been found. Here, we used nested clade phylogeographical analysis (NCPA) and analysis of molecular variance (amova), to examine 781 bp of mitochondrial DNA (nad4 and two tRNAs) from 82 individuals of P. mcallii. We tested whether populations of this species were recently connected, or if they were historically isolated prior to human modification of the region. Our NCPA results indicated significant population structure associated with the Colorado River, suggesting limited gene flow and potential isolation across this barrier. Populations west of the Colorado River, currently isolated from one another by human development, show less genetic differentiation. We also collected homologous sequence data from 34 individuals of P. platyrhinos and seven specimens morphologically intermediate between P. mcallii and P. platyrhinos, as a preliminary investigation of hybridization between these two species. From phylogenetic results of these data, we identified a species (Phrynosoma goodei) previously recognized as a subspecies of P. platyrhinos. Six of the morphologically intermediate specimens shared mtDNA haplotypes with P. goodei, while one was nested among P. mcallii haplotypes.     

Helminth parasites in ringed seals (Pusa hispida) from Svalbard, Norway with special emphasis on nematodes: variation with age, sex, diet, and location of host

Complete gastrointestinal tracts from 257 ringed seals (Pusa hispida) from Svalbard, Norway, were examined for helminth parasites. Three different helminth groups were recorded (acanthocephalans 61.1%; nematodes 38%; cestodes 0.9%). Acanthocephalans (Polymorphidae) and cestodes (Anophryocephalus and Diphyllobothrium sp(p)., as well as unidentified species, were confined to the intestines. The anisakid nematodes Phocascaris phocae, Pseudoterranova sp(p)., Anisakis sp(p)., and Phocascaris/Contracaecum sp(p). were recorded in both stomachs and the anterior part of the small intestines. The abundance of nematodes and acanthocephalans varied significantly with sampling location of the seal hosts. This is likely due to the relative prevalence of Arctic versus Atlantic water in the different fjord systems, which strongly influences the age class and species of fish available as prey for the seals. Adult male ringed seals had significantly higher abundances of nematodes than did adult females or juveniles. Adult males also had significantly higher abundances of acanthocephalans than did adult females, but were not significantly different from juveniles in this regard. Nematode abundance increased significantly with age of male hosts, but this trend was lacking in female seals. Infection parameters appeared to be related to differences in the age of polar cod (Boreogadus saida) exploited by male, female, and juvenile seals.     

Molecular evidence for the origin of workerless social parasites in the ant genus Pogonomyrmex

Abstract.— Speciation of two social parasites from their respective hosts is tested using a molecular phylogeny. Alignment of 711 DNA base pairs of mitochondrial cytochrome b gene was used to assess phylogenetic relationships of inquiline species to their hosts and to other members of the genus. We show that the inquiline social parasites of the North American seed harvester ants are monophyletic, descending from one of the known hosts ( Pogonomyrmex barbatus ) in the recent past and shifting hosts in a pattern similar to that observed in other Hymenopteran social parasites. In addition, the host populations unexpectedly were found to be polyphyletic. Populations of Pogonomyrmex rugosus from an area east of the Chiricahua Mountains in Southern Arizona belong to a mitochondrial clade separate from the more western clade of P. rugosus from the Sonoran and Chihuahuan Deserts. Evidence of mitochondrial DNA introgression between P. rugosus and P. barbatus was also observed. We conclude that Emery's rule does not strictly hold for this system, but that the hosts and parasites are very closely related, supporting a loose definition of Emery's rule.     

Seasonal patterns in the foraging ecology of the harvester ant Pogonomyrmex naegelii (Formicidae, Myrmicinae) in a Neotropical savanna: daily rhythms, shifts in granivory and carnivory, and home range

The temporal dynamics of foraging, diet, and use of space is essential to understand the ecology of harvester ants. Here, we present an account of the foraging ecology of Pogonomyrmex naegelii in Brazilian cerrado savanna. Nests occur on bare ground and contain 166?C580 workers (N?=?3 colonies). Colony activity is unimodal year-round and peaks around the middle of the day. Foragers leave the nest independently and individually search for food in all directions. Ants ventured up to 15?m from nests, with most foraging occurring within 2?m of nests. Colonies tended to have larger home ranges in the dry/cold (April?CSeptember) than in the wet/warm season (October?CMarch). P. naegelii has a generalist and season-dependent diet comprised of many seed species and arthropod prey, and pieces of plant and animal matter. Foragers collected seeds from 34 plant species, predominantly grasses (genera Gymnopogon, Axonopus, Aristida). Over 6,700 seeds can be stored in nest granaries. Ants and termites were the main animal prey retrieved by P. naegelii. The proportion of seeds and arthropods foraged by P. naegelii changes year-round: in the dry/cold season, the diet is predominantly granivorous, whereas in the wet/warm season, seeds and arthropods are retrieved in more balanced proportions. Although food availability was not assessed, year-round diet of P. naegelii matches the pattern of seasonal abundance of grass seeds and arthropod prey in cerrado. Data on harvester ants come mostly from arid habitats; this study is a first assessment of the ecology of a Neotropical Pogonomyrmex from a moderately moist savanna environment.     

下辽河平原农田土壤线虫群落组成的季节变化

下辽河平原农田生态系统在管理过程中频繁的耕作、施肥以及农用化学品施用等引发了一系列问题, 如土壤退化、耕地数量减少以及生产力下降等, 不可避免地对土壤生物健康产生影响。为探究农田土壤人工管理对土壤生物群落动态的影响, 本研究在辽宁沈阳农田生态系统国家野外科学观测研究站开展了农田土壤线虫群落组成的季节变化研究, 对4个季节农田和废弃农田(对照)的土壤线虫群落组成、多度以及多样性等进行了比较分析。研究结果表明, 土壤线虫总多度在废弃农田中显著高于农田, 但季节间差异不显著。季节变化主要显著影响了自由生活线虫的多度, 其在9月达到最高; 季节变化也显著影响了属的数量, 其在非生长季的11月最低。与废弃农田相比, 农田管理显著降低了杂食捕食线虫和食真菌线虫的多度, 土壤食物网结构相对稳定; 而废弃的农田更易受到季节波动的影响, 土壤食物网也受到一定的干扰。     

Population biology of Contracaecum rudolphii sensu lato (Nematoda) in the great cormorant (Phalacrocorax carbo) from northeastern Poland

Parameters related to the occurrence, aggregation, and population structure of the nematode Contracaecum rudolphii Hartwich, 1964 , in the great cormorant ( Phalacrocorax carbo ) from northeastern Poland were analyzed. A total of 491 birds of different ages (adults, immatures, and nestlings) was examined; the cormorants studied were taken from both fresh- and brackish water habitats. Contracaecum rudolphii were found in stomachs of 454 birds (92.5%); the 46,244 nematodes included third- (L3) and fourth-stage larvae (L4), and sub-adult and adult females and males. The distribution of nematodes in the host population were highly aggregated. The occurrence of C. rudolphii was significantly dependent on the host's age and habitat, as well as on season; the proportion of development stages differed significantly depending on birds' age and season (the latter only in adult birds). The infrapopulations of C. rudolphii in the adult cormorants showed distinct density-dependent correlations: that is, as the infrapopulation size increased, the number of adult females C. rudolphii decreased, and the proportion of larvae and sub-adult females increased. A higher proportion of larvae and sub-adult females characterized the component population structure of the nematodes in the cormorant nestlings, compared with adult birds, probably because of immune system deficiency in the immature birds, coupled with the development of the nematode population. Seasonal changes in the C. rudolphii population, observed in the adult cormorants, were not related to seasonality of the L3 occurrence in food; instead, the changes are believed to have resulted from independent processes of elimination of the oldest nematodes and their replacement by larval stages that subsequently matured.     

Diet and foraging of the endemic lizard Cnemidophorus littoralis (Squamata, Teiidae) in the restinga de Jurubatiba, Macaé, RJ.

We investigated the diet and foraging of the endemic teiid lizard Cnemidophorus littoralis in a restinga habitat in Jurubatiba, Macaé - RJ. The stomach contents were removed, analyzed and identified to the Order level. There was no relationship between C. littoralis morphological variables and number, length or volume of preys. Termites (48.7%) and larvae (35.5%) were the most important prey items which occurred in the examined lizards' stomachs. The diet did not differ between males and females. Cnemidophorus littoralis is an active forager and predominantly consumes relatively sedentary prey or prey that is aggregated in the environment. We also found an intact and undigested hatchling of the crepuscular/nocturnal gekkonid lizard Hemidactylus mabouia in the stomach of an adult male of C. littoralis, which indicates that C. littoralis is a potential source of mortality for individuals of H. mabouia in the restinga de Jurubatiba.     

Infection by Contracaecum rudolphii (Nematoda: Anisakidae) in the Neotropic cormorant Phalacrocorax brasilianus,and fishes from the estuary of the Valdivia river,Chile

Seasonal parameters of infection by the nematode Contracaecum rudolphii in the Neotropic cormorant Phalacrocorax brasilianus (Phalacrocoracidae) and their relation with feeding and infection in fishes from River Valdivia, Chile, were determined. The prevalences of infection in birds were similar during seasons, whereas mean intensity and percentages of adult gravid females were higher in spring and summer, respectively. For fishes no seasonal differences were found in infection. Cormorant diet varied seasonally in relation to fish prey consumed and this can be related to seasonal differences in infection parameters of birds. The high infection in birds should be considered as a potential risk for salmoniculture.     

Patterns of spatio-temporal variation in the survival rates of a viviparous lizard: the interacting effects of sex,reproductive trade-offs,aridity, and human-induced disturbance

Examination of the spatial and temporal variation in survival rates provides insight on how the action of natural selection varies among populations of single species. In this study, we used mark-recapture data from seven populations of the viviparous lizard Sceloporus grammicus in Central Mexico and a multi-model inference framework to examine interpopulation variation in the survival of adult males and females. We aimed to analyze the potential effects of aridity, human-induced disturbance, and reproductive costs on the survival rates of these lizards. For females in particular, we also searched for a negative relationship between litter size (adjusted for female size) and female survival. Our results demonstrate seasonal changes in survival for males and females. In three out of our seven study sites female survival decreased during the birthing season. In contrast, male survival did not appear to decrease during the mating season. We found an interaction between site-specific aridity and reproductive season affecting female survival. A decrease in female survival during the birthing season was observed in relatively arid sites. In one of these arid sites we found a negative effect of size-adjusted litter size on female survival: females producing more offspring than those expected for their size were more likely to die. This result represents evidence of a physiological trade-off for gravid females occurring in at least one of the studied populations. Interpopulation variation in the degree of human-induced disturbance could not explain the observed patterns of spatial variation in survival rates. Our results demonstrate wide variation in sex-specific survival patterns of this viviparous lizard and provide evidence that negative associations between reproduction and survival are highly dependent on the local environmental conditions.     

Parasite Prevalence, Abundance, and Diversity in a Semi-free-ranging Colony of Mandrillus sphinx

We examined parasite prevalence, abundance of protozoan cysts and helminth eggs in the feces, and number of parasitic taxa in a population of mandrills (Mandrillus sphinx) in semi-free-ranging conditions in their habitat range, with respect to the annual cycle, sex, age, dominance rank, and female reproductive status. We identified 3 taxa of amebic protozoa (Entamoeba coli, E. histolytica/dispar complex, and Endolimax nana), 1 ciliate protozoa (Balantidium coli), and various nematodes. Prevalence ranged from 1 observation in 874 samples for Trichuris and Mammomonogamus (nematodes) to 100% for Entamoeba coli and Endolimax nana. Daily observation, consistency of fecal samples, and periodic veterinary examination indicated that the mandrills were all healthy, suggesting that the presence of intestinal parasites in the provisioned population is well tolerated. Parasite prevalence, abundance in the feces, and number of taxa varied significantly across the annual cycle. Nematode egg prevalence and abundance were lowest during the dry season. We found no sexual difference and no influence of female dominance rank on parasitic infections. Nematode prevalence increased significantly with age in females, but not in males. There was no influence of age on prevalence of other taxa, abundance in the feces, or number of taxa. Abundance of nematode eggs in the feces was higher in pregnant than in lactating or cycling females. However, births are seasonal in the mandrill colony, and pregnant females were present during the months when nematode egg abundance was also higher in males, suggesting that this may be an influence of climatic seasonality in addition to, or instead of, female status.     

Determinants of helminth infection in a subterranean rodent, the Cape dune mole-rat ( Bathyergus suillus )

The helminth fauna of the largest bathyergid, the Cape mole-rat ( Bathyergus suillus ) was studied throughout an entire calendar year. The species richness encountered was low, with only 3 species of nematodes ( Longistriata bathyergi , Mammalakis macrospiculum, and Trichostrongylus sp.) and 2 species of cestodes ( Taenia sp. and Rodentolepis sp.). At less than 10%, the prevalence for all helminths species was similarly low and may be a result of the solitary lifestyle and the subterranean habitat exploited by this rodent. Clear seasonal patterns were apparent for the most common nematode ( L. bathyergi ), and prevalence and abundance were highest among non-pregnant females compared to males and pregnant females. Dispersal patterns associated with the mating system of the host could explain this pattern. In contrast, the prevalence of the most common cestode ( Taenia sp.) was neither determined by season nor host sex, suggesting that foraging habits may constantly expose B. suillus to this parasite.     

Geographic and host range of the nematode Soboliphyme baturini across Beringia

The nematode Soboliphyme baturini Petrov, 1930, was found to represent a single species with a relatively broad geographic range across Beringia and northwestern North America on the basis of the assessment of molecular sequence data for adult and juvenile parasites. Refuted are hypotheses suggesting that several cryptic species could be partitioned either among an array of mustelid definitive hosts or across the vast region that links North America and Eurasia. Host specificity for this species is examined on the basis of a comprehensive list for definitive hosts, derived from new field surveys and existing literature for S. baturini. Only 5 mustelids (Gulo gulo, Martes americana, M. caurina, M. zibellina, and Neovison vison) appear to have significant roles in the life history, persistence, and transmission of this nematode. Soboliphyme baturini readily switches among M. americana, M. caurina, Mustela erminea, or N. vison at any particular locality throughout its geographic range in North America, although Martes spp. could represent the source for nematodes in a broader array of mustelids. Molecular analyses (243 base pairs of mitochondrial gene nicotinamide dehydrogenase [ND4]) suggest that hypotheses for host specificity across an array of mustelid definitive hosts are not supported. The life cycle of S. baturini is explored through a review of diet literature for 2 marten species, M. americana and M. caurina, and other mustelids across the Holarctic. Shrews (Soricomorpha: Soricidae) comprise >8% of prey for these species of Martes, suggesting their putative role as paratenic hosts. Juvenile nematodes found in the diaphragms of soricids are genetically identical to adult S. baturini found in the stomachs of mustelids at the same locations in both Asia and North America, corroborating a role in transmission for species of Sorex.     

Nematode transmission patterns

The transmission of nematode parasites of vertebrates is reviewed with special reference to the phenomena of monoxeny, heteroxeny, paratenesis, and precocity. Monoxeny is divided into 2 types. Primary monoxeny assumes that there was never an intermediate host in the transmission. Secondary monoxeny assumes the loss of an intermediate host during the course of evolution and its replacement by a tissue phase in the final host. Heteroxeny, or the use of intermediate hosts, is a common feature of many nematode groups. The Spirurida utilize arthropods, the Metastrongyloidea molluscs, and Ascaridida arthropods and vertebrates. Paratenesis, or the use of transport hosts, is a common feature of the transmission of nematode parasites of carnivores. It is postulated that in some instances paratenic hosts have become intermediate hosts and replaced the original intermediate host. Precocity in the development of nematodes in intermediate hosts (including what may have been paratenic hosts) is defined as growth and/or development beyond the expected. Its occurrence among the nematode parasites of vertebrates is reviewed. It is regarded as a transmission strategy which accelerates gamete production in the final host. Precocity could also provide the mechanism for the transfer of a parasite from a predator final host to a prey final host.     

Associations between shorebird abundance and parasites in the sand crab, Emerita analoga, along the California coast

I investigated spatial variation in the prevalence and abundance of 4 species of parasites in the sand crab, Emerita analoga, on 8 sandy beaches along 800 km of the California coast, to assess the importance of bird abundance for the distribution of parasites among sand crab populations. I collected sand crabs and counted shorebirds and gulls at each beach during June and November 1994. Sand crabs served as intermediate hosts for 4 species of parasites, including a trematode, Spelotrema nicolli (Cable and Hunnienen, 1938); an acanthocephalan, Polymorphus kenti (Van Cleave, 1947); a nematode, Proleptus sp., and an unidentified trypanorhynch tapeworm. Among sand crab populations, there was substantial spatial variation in the prevalence and abundance of each parasite species. No latitudinal pattern was apparent for any of the 4 species observed. Temporally, parasite prevalence and abundance was significantly different between dates for all 4 parasites. Specifically, sand crab populations experienced higher trematode, nematode, and trypanorhynch prevalence and abundance in November than in June. In contrast, prevalence and abundance of acanthocephalans were higher in June than in November. There were strong positive associations between bird abundance and prevalence of parasitic infection for trematodes and acanthocephalans for some dates but not for nematodes or trypanorhynchs, which use elasmobranchs as definitive hosts. The spatial variation in prevalence and abundance of trematodes and acanthocephalans observed among sand crab populations may be attributed to the distribution and abundance of shorebirds and gulls that serve as definitive hosts.     

On the life history of the nematode Raphidascaris acus (Bloch, 1779) in the natural environment of the River Bystřice, Czechoslovakia

From April 1967 until July 1968 the life history of the nematode Rughiduscuris acus was followed in the natural environment of the River Bystfice. The incidence, intensity of infec-tion, and also the maturation of the nematode in the definitive host ( Saimo truttu m. fario ) indicate that the life history of this parasite exhibits an obvious annual cycle, when adult egg-producing female nematodes are present only from May until the beginning of July. This seasonal rhythm is largely due to changes in the food composition of the definitive host, although the effects of other factors cannot be excluded. In the River BystHce the main intermediate hosts of R. ucus are the bullhead ( Cottus gobio ) and the Siberian sculpin ( Cottuspoecilopus ) in the trout zone of the river and the stone Ioach ( Noernacheilus barbutulus ) in the lowland reaches. Larvae of the species Prodiamesa olivacca ( Diptera: Chironomidue ) were found to act as reservoir hosts for the second-stage larvae of Raphiduscuris acus.