Critical thermal maximum of Amur sleeper <Emphasis Type="Italic">Perccottus glenii</Emphasis> in different seasons of the year

Seasonal differences in the critical thermal maximum are revealed in Amur sleeper Perccottus glenii at the age of 1+ and 2 years. Maximum values of the critical thermal maximum determined by two criteria (rotation of the body and immobile gill covers) are registered in the summer, and minimum values of this parameter are observed in the winter. Maximum thermal resistance is registered at a low water heating rate (0.2–0.4°C/h), and minimum resistance is reported for a heating rate 5.0–10.5°C/h. The difference between the values of the critical thermal maximum assessed by two criteria reaches 1–2°C. Features of behavior of Amur sleeper are described at different heating rates. Based on the characteristics of thermal adaptation (critical thermal maximum and optimal temperature for growth), Amur sleeper is similar to warm water eurythermal species (goldfish Carassius auratus and carp Cyprinus carpio).     

Local adaptation of a holoparasitic plant,Cuscuta europaea: variation among populations

Locally adapted parasites have higher infectivity and/or fitness on sympatric than on allopatric hosts. We tested local adaptation of a holoparasitic plant, Cuscuta europaea, to its host plant, Urtica dioica. We infected hosts from five sites with holoparasites from the same five sites and measured local adaptation in terms of infectivity and parasite performance (biomass) in a reciprocal cross‐infection experiment. The virulence of the parasite did not differ between sympatric and allopatric hosts. Overall, parasites had higher infectivity on sympatric hosts but infectivity and parasite performance varied among populations. Parasites from one of the populations showed local adaptation in terms of performance, whereas parasites from one of the populations had higher infectivity on allopatric hosts compared with sympatric hosts. This among‐population variation may be explained by random variation in parasite adaptation to host populations or by time‐lagged co‐evolutionary oscillations that lead to fluctuations in the level of local adaptation.     

Metazoan parasite communities of wild Leporinus friderici (Characiformes: Anostomidae) from Amazon River system in Brazil

This study compared the structure of metazoan parasite communities in two populations of Leporinus friderici from two rivers of the Amazon River system in Brazil. Jainus leporini, Urocleidoides paradoxus, Urocleidoides sp., Tereancistrum parvus, Tereancistrum sp., Clinostomum marginatum, Procamallanus (Spirocamallanus) inopinatus, Contracaecum sp., Octospiniferoides incognita and Ergasilus sp. were found in both hosts. There were differences in species richness of parasites, Shannon index and the evenness between both host populations, because the parasite community showed a similarity of only 33%. Parasites of both host populations had an aggregated dispersion. Host size also contributes to the differences between the populations of hosts investigated, and the feeding habit contributes to the occurrence of endoparasites in L. friderici, an intermediate host for these parasites. The existence of variation in infracommunity and community of parasites for the same host species from different localities indicates the presence of an uneven distribution in terms of species and density of parasites, which are intermediate hosts, and some of them constitute food items for L. friderici in the localities surveyed.     

Community of parasites in Triportheus curtus and Triportheus angulatus (Characidae) from a tributary of the Amazon River system (Brazil)

The present study compared the community of parasites in populations of Triportheus angulatus and T. curtus from a tributary of the Amazon River in northern Brazil. All hosts had one or more parasite species, 862,687 parasites were collected from T. curtus and 302,008 from T. angulatus. Species richness of parasites, Shannon diversity index and evenness index were higher for T. angulatus. The parasite communities of both hosts was similar (99%) and composed by Ichthyophthirius multifiliis, Anacanthorus pitophallus, metacercariae and Procamallanus (Spirocamallanus) inopinatus, with dominance of I. multifiliis. However, low infection level by Piscinodinium pilullare, Dolops sp. and Contracaecum larvae occurred only in T. angulatus, and Spironucleus sp. was found only in T. curtus. There were differences in the prevalence, intensity and mean abundance for some parasites of both hosts. There was aggregated dispersion of parasites in both hosts, but the infection of P. (S.) inopinatus in T. angulatus showed a random dispersion. For T. angulatus and T. curtus, the diversity and abundance of parasites were influenced by the host size. This was the first report of these parasite species for T. curtus, as well as of I. multifiliis, P. pilullare, Dolops sp., P. (S.) inopinatus and Contracecum sp. for T. angulatus.     

Amur sleeper (Perccottus glenii Dubowski, 1877) spreading in the Danube River Basin

Whereas earlier Amur sleeper (Perccottus glenii) findings in the Danube River were only single specimens, data reported in this study represent the first record on establishment of a local population. An overview of the spread of the Amur sleeper in the Danube River basin is also presented.     

The importance of gobies (Gobiidae,Teleostei) as hosts and transmitters of parasites in the SW Baltic

The parasite fauna of five goby species (Gobiidae, Teleostei) was investigated in the Baltic Sea during the period 1987 to 1990. 13 parasite species were found in samples from the Lübeck Bight:Bothriocephalus scorpii, Schistocephalus sp. (Cestoda);Cryptocotyle concavum, Cryptocotyle lingua, Podocotyle atomon, Derogenes varicus (Digenea);Hysterothylacium sp. (cf.auctum),Contracaecum sp.,Anisakis simplex (Nematoda);Corynosoma sp.,Echinorhynchus gadi, Neoechinorhynchus rutili, Pomphorhynchus laevis (Acanthocephala). The number of parasite species were: 10 in the sand gobyPomatoschistus minutus, 8 in the black gobyGobius niger, 7 in the two-spotted gobyGobiusculus flavescens, 6 in the common gobyPomatoschistus microps, and 5 in the painted gobyPomatoschistus pictus. Neoechinorhynchus rutili occurred only inP. minutus, andCorynosoma sp. only inG. niger. The extent to which the gobies were parasitized clearly depended on the respective ways of life and, moreover, on the kind of prey ingested by the hosts. Additionally, the age of the hosts might be important. The highest rate of parasitism, more than 60%, was reached byHysterothylacium sp. inG. niger and byCryptocotyle concavum inP. microps. Infestation incidence lay mostly below 40% which means a satellite species status (Holmes, 1991). The number of parasite species was highest in summer; the highest intensities of single parasites occurred in spring (Podocotyle atomon) or autumn (Crytocotyle concavum).Bothriocephalus scorpii, Hysterothylacium sp. andPodocotyle infested their juvenile hosts very early, but onlyHysterothylacium was accumulated byG. niger during its whole life span, whereasBothriocephalus persisted also in older gobies in low intensities. The cercariae ofCryptocotyle spp. penetrate actively into their hosts; all the other parasites named were transmitted in larval form by prey organisms which consisted mainly of planktonic and benthic crustaceans. The gobies were final hosts for only 5 parasites; but two species may be transmitted to larger fish, and 6 species to sea birds or mammals. The parasite community of the five gobies may possibly be taken to characterize the ecological quality of the environment of the Lübeck Bight.     

Two new cryptic and sympatric species of the king crab parasite Briarosaccus (Cirripedia: Rhizocephala) in the North Pacific

Rhizocephalan barnacles have been reported to parasitize a wide range of king crab species (Lithodidae). So far all these parasites have been assigned to a single species, Briarosaccus callosus Boschma, 1930, which is assumed to have a global distribution. Here we investigate Briarosaccus specimens from three different king crab hosts from the fjord systems of Southeastern Alaska: Lithodes aequispinus Benedict, 1895, Paralithodes camtschaticus (Tilesius, 1815), and Paralithodes platypus (Brandt, 1850). Using molecular markers and by morphological comparison we show that Briarosaccus specimens from these three commercial exploited king crabs are in fact morphologically distinct from B. callosus, and further represent two separate species which we describe. The two new species, Briarosaccus auratum n. sp. and B. regalis n. sp., are cryptic by morphological means and were identified as distinct species by the use of genetic markers (COI and 16S). They occur sympatrically, yet no overlap in king crab hosts occurs, with B. auratum n. sp. only found on L. aequispinus, and B. regalis n. sp. as parasite of the two Paralithodes hosts. © 2015 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London     

THE EVOLUTION OF PARASITISM IN THE RED ALGAE: MOLECULAR COMPARISONS OF ADELPHOPARASITES AND THEIR HOSTS1

In several groups of parasites including insect, flowering plant, fungal, and red algal parasites, morphological similarities of the parasites and their specific hosts have led to hypotheses that these parasites evolved from their hosts. But these conclusions have been criticized because the morphological features shared by parasite and host may be the result of convergent evolution. In this study, we examine the hypothesis, originally put forth by Setchell, that adelphoparasitic red algae, that is, parasitic red algae that are morphologically very similar to their hosts, evolved from their specific red algal hosts. Rather than comparing morphological features of parasites and hosts, small-subunit 18S nuclear ribosomal DNA and the internal transcribed spacer regions (ITSs) of the nuclear ribosomal repeat are compared for five parasites, their hosts, and related nonhosts from four red algal orders. These comparisons reveal that each of these adelphoparasites has evolved either directly from the host on which it is currently found, or it evolved from some other taxon that is closely related to the modern host. The parasites Gardneriella tuberifera, Rhodymeniocolax botryoides, and probably Gracilariophila oryzoides evolved from their respective hosts Sarcodiotheca gaudichaudii, Rhodymenia pacifica, and Gracilariopsis lemaneiformis, respectively. The parasite Faucheocolax attenuata evolved from either Fauchea laciniata or Fauchea fryeana and subsequently radiated onto the other host species. Presently this parasite is found on both hosts. Lastly, some parasitic genera such as Plocamiocolax are polyphyletic in their origins. A species of Plocamiocolax from an Antarctic Plocamium cartilagineum appears to have evolved from its host whereas the common Plocamiocolax pulvinata that occurs along the west coast of North America likely evolved from Plocamium violaceum and radiated secondarily onto its present day host, Plocamium cartilagineum.     

Polymorphisms at two enzyme loci (Sod and Odh) in populations of the Amur sleeper,Perccottus glenii (Pisces,Eleotrididae), from its native range and the colonized area: the effect of introduction on genetic variation

In the first half of the 20th century small founder groups of the Amur sleeper, Perccottus glenii, have been transferred from their native range in the Far East to some regions of the European part of Russia. In order to investigate the consequences of the introductions, 15 Far East populations and 11 European populations, that originated from two transfers, were studied for their variability on two enzyme loci (Sod and Odh) using polyacrylamide gel electrophoresis. For both loci, divergence between populations and heterozygosity in the populations were higher in the colonized regions than in the native range. Heterozygote advantage in small founder populations seems to be a factor resisting genetic drift and determining the changes observed in the genetic structure of the introduced populations.     

Life-history traits of Amur sleeper, <Emphasis Type="Italic">Perccottus glenii</Emphasis>, in the invaded Vistula River: early investment in reproduction but reduced growth rate

In recent decades, Amur sleeper is one of the most invasive fish species in Eastern and Central Europe. Generally, it is assumed that success of an invasive species can largely depend on the plasticity of its life-history traits, e.g., indicated by higher investment in reproduction in the initial stage of its invasion. On the other hand, such energy allocation to production of gonads should negatively impact somatic growth rate. The aim of this article was to explore this phenomenon in a non-native population of Amur sleeper inhabiting artificial reservoir on a large lowland river in Central Europe. Through analysis of the population age structure, sex-dependent growth rate and life-history traits we assessed investment in reproduction and its possible relationship with growth pattern as well as compared the results with literature data from native and introduced range. Samples collected monthly from April 2005 to March 2006 were used to estimate sex ratio, GSI, fecundity, eggs diameter frequency over the year, duration of spawning season, and mode of spawning. Age structure was calculated from scales and the same data were also used to back-calculate standard length (SL) at age. The population in the Włocławski Reservoir was represented by eight age groups, and its life span was one of the longest recorded both in its natural range and introduced areas. The weight–length relationship showed that the Amur sleeper grew isometrically and there was no difference between females and males. Although the Amur sleeper growth rate was relatively slow in the Włocławski Reservoir, its increment in the first year of life was comparable to that in other introduced areas and higher than in its natural range. The female age of maturation (1+) found in the reservoir was earlier than observed in its natural range as well as in most introduced areas (2+, 3+). Female length at maturation was similar to that reported elsewhere. The back-calculated standard length (SL) of females and males showed that in the first two age classes (1 and 2) males achieved larger SL than females. Meanwhile, at age 3, 4 and 5, females were larger than males but these differences were insignificant, except for class 5. For both sexes the average annual increment of SL decreased with age but in females the increment was always higher than in males (P < 0.05). The growth rate differences between females and males resulted possibly from their unequal investment in reproduction. Multiple spawning in the Włocławski Reservoir lasted from April to the end of August and was almost 3 months longer than in other regions. Thus, this higher investment in reproduction displayed by, e.g., earlier maturation of females and longer spawning season in the Włocławski Reservoir, may contribute to invasive success of Amur sleeper in newly colonized areas; however, the costs of this strategy result in slower growth in older age classes.     

Metazoan parasites of Neogobius fishes in the Slovak section of the River Danube

Metazoan parasites were investigated in three non‐native fishes (monkey goby Neogobius fluviatilis, bighead goby Neogobius kessleri and round goby Neogobius melanostomus) collected from the former and current main channel of the River Danube and from the River Hron, Slovakia, in November 2003. Thirteen parasite species were identified: Triaenophorus crassus, Diplostomum sp., Tylodelphys clavata, Metorchis xanthosomus, Nicolla skrjabini, Gyrodactylus proterorhini, Pomphorhynchus laevis, Contracoecum sp., Raphidascaris acus, Anguillicola crassus, Unio tumidus, Anodonta anatina and Pseudoanodonta complanata. The maximum parasite diversity was found in N. fluviatilis. Total parasite abundance was significantly higher in N. kessleri, but no significant differences among sampling sites were observed. Pomphorhynchus laevis and glochidia of Anodonta anatina reached 100% prevalence in N. kessleri in the new channel of the Danube and, in general, these species were also the most prevalent parasites in all three goby species. For endoparasites, gobies served mostly as intermediate (digenean, cestodes and nematodes) or paratenic (acantocephalan and nematodes) hosts. All parasite species found are common parasites in the Middle Danube basin. No parasites specific to Neogobius, known from their native populations, were observed.     

Systematics of the Eimerian Parasites from North American Snakes of the Family Colubridae,and their Prevalence in the Colubrids of Iowa*

A survey of 117 Iowa snakes, representing 18 species within 12 genera, revealed the presence of 6 species of Eimeria, 5 of which are described as new and 1 of which (E. zamenis) is redescribed. Those species found, the average length-width dimensions of their oocysts ( in micrometers ), and the respective hosts from which they were isolated were as follows: E. attenuata sp. n., 22.2 × 12.6, from 1 of 25 red-sided garter snakes [Thamnophis sirtalis parietalis (Say)] and 1 of 14 northern water snakes [Natrix sipedon sipedon (Linnaeus)]; E. iowaensis sp. n., 17.8 × 14.5, from 1 of 25 redsided garter snakes; E. hydrophis sp. n., 15.4 × 10.9, from 5 of 14 northern water snakes and 1 of 1 diamond-backed water snake [N. rhombifera rhombifera (Hallowell)]; E. helmisophis sp. n., 13.8 × 10.6, from 1 of 5 western worm snakes [Carphophis amoenus vermis (Kennicott)]; E. collanuli sp. n., 33.1 × 18.3, from 1 of 14 prairie ring-neck snakes (Diadophis punctatus arnyi Kennicott), and E. zamenis, 31.0 × 17.0, from 1 of 6 eastern yellow-bellied racers (Coluber constrictor flaviventris Say) and 1 of 1 eastern milk snake [Lampropeltis triangulum triangulum (Lacépède)]. The overall infection rate for the 117 snakes examined was 9.2%; these data are tabulated. In addition, the possible synonymy of E. lampropeltis with E. zamenis is considered, and the probable status of E. attenuata, E. hydrophis, E. zamenis, and E. annea, as parasites of multiple host species, is reviewed with regard to the phylogenetic relationships of the respective hosts from which they have been reported.     

Inspection and evaluation of host plant by the butterfly Mechanitis lysimnia (Nymph., Ithomiinae) before laying eggs: a mechanism to reduce intraspecific competition

Parasites of all kinds affect the behaviour of their hosts, often making them more susceptible to predators. The associated loss in expected future reproductive success of infected hosts will vary among individuals, with younger ones having more lose than older ones. For this reason, young hosts would benefit more by opposing the effects of parasites than old ones. In a laboratory study, the effects of the trematode Telogaster opisthorchis on the anti-predator responses of the upland bully (Gobiomorphus breviceps) and of the common river galaxias (Galaxias vulgaris) were examined in relation to fish age. In a bully population where parasites were very abundant, the magnitude of the fish's anti-predator responses decreased as the number of parasites per fish increased, and this effect was significantly more pronounced in age 2 + and, to a lesser extent, age 3 + fish than in age 1 + fish. In another bully population where parasites were 10 times less abundant, similar effects were noticeable but not significant, whereas no effects of parasites on the responses of galaxiids to predators were apparent. Differences in the abundance of parasites and in their sites of infection in fish may explain the variability among host populations or species. However, in the bully population with high parasite abundance, parasitism has age-dependent effects on responses to predators, providing some support for the prediction that young fish with high expected future reproductive success invest more energy into opposing the effects of parasites than do older fish.     

Bacterial and β-proteobacterial diversity in Olea europaea var. mastoidis- and O. europaea var. koroneiki-generated olive mill wastewaters: influence of cultivation and harvesting practice on bacterial community structure

Despite the fact that several approaches have been applied for the bioremediation of olive mill wastewaters, little information is available on bacteria inhabiting these agro-industrial effluents. In the present study, 16S rRNA gene clone libraries were constructed to identify bacterial diversity in olive-oil mill wastewaters generated by two olive varieties, Olea europaea var. mastoidis and O. europaea var. koroneiki. Due to chloroplast excess in wastewater produced from the processing of O. europaea var. koroneiki, a clone library using specific PCR primers for β-Proteobacteria was further constructed. The bacterial diversity in O. europaea var. mastoidis-generated olive mill wastewaters consisted mainly of members of Acetobacteriaceae, Prevotellaceae and Lactobacillaceae, while the majority of β-proteobacteria identified in O. europaea var. koroneiki-generated olive mill wastewaters were placed within the families Comamonadaceae, Oxalobacteraceae, Hydrogenophilaceae and Rhodocyclaceae. At least 17 novel phylogenetic linkages among Bacteria were identified. Olive-oil mill wastewaters microbiota appears to have originated from soil and freshwater environments, while the cultivation and harvesting practice highly influenced the bacterial community structure in olive mill wastewaters. The presence of fecal bacteria in O. europaea var. mastoidis-generated olive mill wastewaters, due to the long harvesting period, should be of concern.     

Interaction of a host plant and its holoparasite: effects of previous selection by the parasite

If parasites decrease the fitness of their hosts one could expect selection for host traits (e.g. resistance and tolerance) that decrease the negative effects of parasitic infection. To study selection caused by parasitism, we used a novel study system: we grew host plants (Urtica dioica) that originated from previously parasitized and unparasitized natural populations (four of each) with or without a holoparasitic plant (Cuscuta europaea). Infectivity of the parasite (i.e. qualitative resistance of the host) did not differ between the two host types. Parasites grown with hosts from parasitized populations had lower performance than parasites grown with hosts from unparasitized populations, indicating host resistance in terms of parasite’s performance (i.e. quantitative resistance). However, our results suggest that the tolerance of parasitic infection was lower in hosts from parasitized populations compared with hosts from unparasitized populations as indicated by the lower above‐ground vegetative biomass of the infected host plants from previously parasitized populations.     

Gastrointestinal parasites of the chimpanzee population introduced onto Rubondo Island National Park,Tanzania

The release of any species into a novel environment can evoke transmission of parasites that do not normally parasitize the host as well as potentially introducing new parasites into the environment. Species introductions potentially incur such risks, yet little is currently known about the parasite fauna of introduced primate species over the long term. We describe the results of long‐term monitoring of the intestinal parasite fauna of an unprovisioned, reproducing population of chimpanzees introduced 40 years earlier (1966–1969) onto Rubondo Island in Lake Victoria, Tanzania, a non‐native habitat for chimpanzees. Two parasitological surveys (March 1997–October 1998 and October 2002–December 2005) identified Entamoeba spp. including E. coli, Iodamoeba buetschlii, Troglodytella abrassarti, Chilomastix mesnili, Trichuris sp., Anatrichosoma sp., Strongyloides spp., Strongylida fam. gen. sp., Enterobius anthropopitheci, Subulura sp., Ascarididae gen. sp., and Protospirura muricola. The parasite fauna of the Rubondo chimpanzees is similar to wild chimpanzees living in their natural habitats, but Rubondo chimpanzees have a lower prevalence of strongylids (9%, 3.8%) and a higher prevalence of E. anthropopitheci (8.6%, 17.9%) than reported elsewhere. Species prevalence was similar between our two surveys, with the exception of Strongyloides spp. being higher in the first survey. None of these species are considered to pose a serious health risk to chimpanzees, but continued monitoring of the population and surveys of the parasitic fauna of the two coinhabitant primate species and other animals, natural reservoir hosts of some of the same parasites, is important to better understand the dynamics of host–parasite ecology and potential long‐term implications for chimpanzees introduced into a new habitat. Am. J. Primatol. 72:307–316, 2010. © 2009 Wiley‐Liss, Inc.     

Identification of microsatellite markers in Plasmodium mexicanum,a lizard malaria parasite that infects nucleated erythrocytes

Reptile and bird hosts of malaria parasites (Plasmodium) have nucleated erythrocytes. Infected blood thus contains a mix of abundant host and scant parasite DNA which has prevented identification of Plasmodium microsatellites. We developed a protocol for isolation of microsatellite markers for Plasmodium mexicanum, a parasite of lizards. The ATT repeat was common in the genome of P. mexicanum, but most (87%) of these repeats were exceptionally long (50–206 + repeats). Seven microsatellite markers with polymerase chain reaction primers are described. The protocol should allow discovery of microsatellites of malaria parasites (with AT‐rich genomes) infecting bird and reptile hosts.     

Parasite transfer from crustacean to fish hosts in the Lübeck Bight,SW Baltic Sea

Four helminth parasites out of 19 species found in the Lübeck Bight, Baltic Sea, were chosen for investigations on the transfer from invertebrate to small-sized fish hosts: larvae of the tapewormsSchistocephalus sp. andBothriocephalus sp. (Cestoda) living in planktonic copepods as primary hosts;Podocotyle atomon (Digenea) andHysterothylacium sp. (Nematoda) were found in benthic crustaceans, especiallyGammarus spp. These hosts were the prey of 3 gobiid fishes,Gobiusculus flavescens (feeding mainly on plankton),Pomatoschistus minutus (preferring benthos), andP. pictus (feeding more on plankton than benthos). Because the fishes selected smaller sizes of crustaceans, they ingested all stages of the copepods but only the smaller-sized groups of gammarids which were often less infested by parasites. In order to evaluate the probability for a fish to be parasitized by a helminth, an infestation potential index (IP) was calculated.Podocotyle atomon andHysterothylacium sp. revealed an IP which was far lower in gobies than expected when the prevalences of the previous hosts were taken into consideration. The IP of tapeworm larvae was mainly influenced by the feeding pressure of the gobiid predators, which might change with developmental stage and season. It is concluded that parasite transfer to the next host decreases when sizes of prey and predator differ only moderately. This mechanism can reduce the numbers of parasites transferred to less suitable or wrong hosts.     

A PHYLOGENETIC STUDY OF PARASITIC GENERA PLACED IN THE FAMILY CHOREOCOLACACEAE (RHODOPHYTA)1

Red algal parasites are common and have a unique type of development in which parasite nuclei are transferred to host cells and “control” host cell development. Previous phylogenetic studies have concentrated on parasites closely related to their hosts, termed adelphoparasites. A second set of parasites, usually classified in a different family or tribe from their host, termed alloparasites, have not been studied phylogenetically. This study concentrates on the wholly parasitic family, the Choreocolacaceae (Gigartinales). Using small subunit rDNA sequence data, we found that all the parasites studied are within the same family as their host. Our data support the placement of Holmsella, species of which parasitize Gracilaria and Gracilariopsis, in the order Gracilariales and suggest that Holmsella is an old parasitic genus. Most other species of the Choreocolacaceae parasitize species of the Rhodomelaceae. The one exception is the hyperparasitism between Harveyella mirabilis (Reinsch) F. Schmitz et Reinke (Rhodomelaceae) and the parasite Gonimophyllum skottsbergii Setchell (Delesseriaceae). The parasites Bostrychiocolax australis Zuccarello et West and Dawsoniocolax bostrychiae (Joly et Yamaguishi‐Tomita) Joly et Yamaguishi‐Tomita are placed within the tribe Bostrychiae as are their hosts. Harveyella mirabilis has a single origin and has switched hosts several times during its passage between the Atlantic and Pacific Oceans. Evidence does not support the continued recognition of the family Choreocolacaceae. Our results also indicate that the distinction between adelphoparasites and alloparasites is unwarranted, with a continuum between newly evolved parasites closely related to their hosts and parasites less closely related to their hosts.