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Molecular phylogeny of arthropods and their relatives: polyphyletic origin of arthropodization.

Because the enigmatic arthropod relatives, such as tardigrades, onychophorans, pentastomids, and pycnogonids, have features characteristic of not only both major arthropod groups but also several other animal phyla, analysis of their phylogenetic positions may provide important clues for metazoan phylogeny in general. From analyses of complete or nearly complete 18S rDNA sequences, arthropod monophyly including all the traditional arthropod relatives was well supported. For detailed analyses within arthropods without outgroup effects, we made two reduced alignments. From the analyses of these reduced data sets, arthropods were divided into two clades: one was chelicerates and myriapods, whereas the other included crustaceans plus insects, onychophorans plus pentastomids, and tardigrades. The pycnogonids were grouped closely with the chelicerates. The Cambrian fossil record supports that myriapod-chelicerate and crustacean-insect (including tardigrades, onychophorans, and pentastomids) stems arose separately from a protoarthropod (perhaps nonarthropodized animal, such as a lobopod which has noncalcified cuticles, and nonjointed legs). Thus, the acquisition of truly arthropod characters occurred independently in the two stems. In this case, nonarthropodized characters of tardigrades, onychophorans, and pentastomids are assumed to be primitive.

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Novel rearrangements of arthropod mitochondrial DNA detected with long-PCR: applications to arthropod phylogeny and evolution.

Rearrangements of mitochondrial DNA gene order have been suggested as a tool for defining the pattern of evolutionary divergence in arthropod taxa. We have employed a combination of highly conserved insect-based polymerase chain reaction (PCR) primers with long-PCR to survey 14 noninsect arthropods for mitochondrial gene rearrangements. The size of the amplified fragments was used to order the primer containing genes. Five chelicerates exhibit amplicons that are consistent with the presumptive ancestral arthropod mtDNA gene order. These five species comprise two soft ticks, two prostriate hard ticks, and an opilionid. Six other chelicerates, all metastriate hard ticks, have a different arrangement that was originally discovered by this procedure and has been previously detailed in a complete mtDNA sequence. Three new arthropod mtDNA gene arrangements are described here. They were discovered in a terrestrial crustacean (Isopoda) and two myriapods (Chilopoda, centipede; Diplopoda, millipede). These rearrangements include major realignments of some of the large coding regions and two possible new positions for the tRNA(Met) (M) gene in arthropods. The long-PCR approach affords an opportunity to quickly screen divergent taxa for major rearrangements. Taxa exhibiting rearrangements can be targeted for DNA sequencing of gene boundaries to establish the details of the mtDNA organization.

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Water balance in desert arthropods. Despite their small size, arthropods may be highly adapted for life in xeric conditions.

As judged by the number of species, or of individuals, arthropods are an extremely successful group of desert inhabitants. There is very great structural and physiological diversity within the group, and since adaptations to desert life open to one are not open to all. we should not expect to find the maximum possible development of adaptive features in any arthropod simply because it lives in a desert. Most adult insects fly; their larvae and all other arthropods do not, and their adaptations will differ accordingly. Desert beetles have very impermeable cuticles and tolerate high body temperatures, while desert cockroaches live below the sand. have more permeable cuticles, and absorb water vapor. There is probably no single respect in which all desert arthropods differ from insects of other environments. Perhaps a profitable way of viewing desert animals is to recognize that each is a whole organism with a specific collection of adaptations that must be consistent within themselves and which are associated with a specific mode of life and a specific evolutionary history. The arthropod organization is capable of producing highly efficient desert species. There is, however, a converse way of looking at the situation, Which is often neglected but which may be of general biological interest: does the evolution of adaptations to desert environments necessarily involve loss of viability in more mesic habitats? If so, then what are these disavantages- what, for example, is the disadvantage of a highly impermeable cuticle? In some cases the answer is clear: sandroaches need sand dunes to live in because they are morphologically and behaviorly specialized for this habitat. More often the answer is not obvious.

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The position of arthropods in the animal kingdom: a search for a reliable outgroup for internal arthropod phylogeny.

Morphological evidence for the phylogeny of the animal kingdom has been discussed by numerous authors. DNA sequencing and phylogenetic methods for analyzing these data are alternative approaches to animal phylogeny, but the phenomenon of long branch attraction and poor taxonomic sampling have caused misinterpretations of metazoan relationships. Here we report a cladistic approach to metazoan evolution including 133 18S rDNA sequences of 31 animal phyla. Despite the difficulties associated with analyzing large data sets, our data suggest that the Bilateria and Protostomia are monophyletic. The internal phylogeny of the protostomes is divided into two main clades. One clade includes the classical protostome worms (annelids, sipunculans, echiurans, pogonophorans, and vestimentiferans), mollusks, nemerteans, "lophophorates," platyhelminths, rotiferans, and acanthocephalans, although the internal resolution of the clade is very low. The second clade includes arthropods and other molting animals: tardigrades, onychophorans, nematodes, nematomorphans, kinorhynchs, and priapulans. The arthropods and related phyla lack a ciliated larvae, lack a multiciliate (locomotory) epithelium, and share many features, notably, a reduced coelomic cavity and the presence of a cuticle which molts. The use of these outgroups within the molting clade to root arthropod phylogenies is recommended instead of using annelids or other spiralians. The data are quite conclusive in those phyla with a good taxonomic sampling (i.e., platyhelminths and arthropods).

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Carcass enrichment does not alter decay rates or arthropod community structure: a test of the arthropod saturation hypothesis at the anthropology research facility in Knoxville, Tennessee.

In a test of an arthropod saturation hypothesis, we asked if the 30-yr history of carcass enrichment at the Anthropology Research Facility, Knoxville TN, has altered carcass decay rates or community structure of sarcosaprophagous arthropods, compared with three local nonenriched sites. Over a 12-d period in 1998, using pitfall traps and sweep nets, we sampled a total of 81,000 invertebrates from freshly euthanized pigs (Sus scrofa L.) placed in these sites. From this number, we sorted 69,286 forensically important (sarcosaprophagous) arthropods. The community structure of these organisms, as measured by species and individuals accumulation curves, rarefaction, and nonparametric correlation, was comparable in all four sites in taxonomic similarity, colonization rates, aerial species richness, and ranked abundances of forensically important taxa on a per carcass basis. Measures of carcass decay rate, remaining carcass weight (%) and periodic weight loss, also were similar. In most cases, carcass surface temperatures and maggot mass temperatures were also statistically indistinguishable. Probability-based results and posthoc power analyses of these variables led us to conclude that the sarcosaprophagous arthropod community of the Anthropology Research Facility is representative of surrounding sites.

Animal Feed↗

Does carcass enrichment alter community structure of predaceous and parasitic arthropods? A second test of the arthropod saturation hypothesis at the Anthropology Research Facility in Knoxville, Tennessee.

In a second test of an arthropod saturation hypothesis, we analyzed if the on-campus Anthropology Research Facility (ARF) at the University of Tennessee, Knoxville, with its 20+ yr history of carcass enrichment, is comparable to non-enriched sites in community structure of predatory and parasitic arthropods that prey upon the sarcosaprophagous fauna. Over a 12-day period in June 1998, using pitfall traps and sweep nets, 10,065 predaceous, parasitic, and hematophagous (blood-feeding) arthropods were collected from freshly euthanized pigs (Sus scrofa L.) placed at ARF and at three surrounding sites various distances away (S2-S4). The community structure of these organisms was comparable in most paired-site tests with respect to species composition, colonization rates, and evenness of pitfall-trap abundances on a per carcass basis. Site differences were found in rarefaction tests of both sweep-net and pitfall-trap taxa and in tests of taxonomic evenness and ranked abundances of sweep-net samples. Despite these differences, no evidence was found that the predatory/parasitic fauna at ARF was impoverished with fewer but larger populations as a result of carcass enrichment. Comparison of the sarcosaprophagous and predatory/parasitic faunas revealed a tighter (and more predictable) linkage between carrion feeders (sarcosaprovores) and their carrion than between carrion feeders and their natural enemies (predators and parasitoids), leading us to conclude that ARF is more representative of surrounding sites with respect to the sarcosaprovore component than to the predatory/parasitic component within the larger carrion-arthropod community.

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The arthropod fauna of Quebec vineyards with particular reference to phytophagous arthropods.

A 3-yr study using different sampling and trapping techniques showed that the arthropod pest fauna in two commercial vineyards in southwestern Quebec was qualitatively and quantitatively different than that of Ontario, Canada, and New York state. We hypothesize that a colder winter climate in addition to the agronomic activity of earthing up around the vines in autumn to protect the roots from freezing in winter contributed to low numbers of pests, such as the grape berry moth, Endopiza viteana Clemens (Lepidoptera, Tortricidae). Once in 3 yr, the density of this pest approached, in one of the vineyards, the action threshold recommended for New York. Therefore, it should be monitored on an annual basis. Another phytophagous arthropod that has the potential to cause sporadic economic damage is the potato leafhopper, Empoasca fabae (Harris). The Asiatic garden beetle, Maladera (= Autoserica) castanea (Arrow), was reported for the first time in Canada. The tarnished plant bug, Lygus lineolaris (Palisot de Beauvois), was also captured by sampling. However, its status as a pest has yet to be clarified.

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Common origin of arthropod tyrosinase, arthropod hemocyanin, insect hexamerin, and dipteran arylphorin receptor.

Dipteran arylphorin receptors, insect hexamerins, cheliceratan and crustacean hemocyanins, and crustacean and insect tyrosinases display significant sequence similarities. We have undertaken a systematic comparison of primary and secondary structures of these proteins. On the basis of multiple sequence alignments the phylogeny of these proteins was investigated. Hexamerin subunits, hemocyanin subunits, and tyrosinases share extensive similarities throughout the entire amino acid sequence. Our studies suggest the origin of arthropod hemocyanins from ancient tyrosinase-like proteins. Insect hexamerins likely evolved from hemocyanins of ancient crustaceans, supporting the proposed sister-group position of these subphyla. Arylphorin receptors, responsible for incorporation of hexamerins into the larval fat body of diptera, are related to hexamerins, hemocyanins, and tyrosinase. The receptor sequences display extensive similarities to the first and third domains of hemocyanins and hexamerins. In the middle region only limited amino acid conservation was observed. Elements important for hexamer formation are deleted in the receptors. Phylogenetic analysis indicated that dipteran arylphorin receptors diverged from ancient hexamerins, probably early in insect evolution.

Amino Acid Sequence↗

[Imported arthropod-borne parasites and parasitic arthropods in dogs. Species spectrum and epidemiologic analysis of the cases diagnosed in 1995/96].

Between January 1995 and December 1996 nonendemic or only regionally occurring arthropodborne parasites including ehrlichiae and parasitic arthropods in Germany were detected in 484 dogs, whereby at least 15 species were involved. Listed in decreasing order, Leishmania infections occurred most frequently, followed by infections/infestations with Babesia canis, Ehrlichia canis, Rhipicephalus sanguineus, Dirofilaria immitis and Dermacentor reticulatus. The other species, namely Babesia gibsoni, Trypanosoma congolense, Hepatozoon canis, Ehrlichia platys, Dirofilaria repens, Dipetalonema reconditum, Dermacentor marginatus, Boophilus microplus and Cordylobia anthropophaga were demonstrated in a few or single dogs. Irrespective of the pathogen-species, most of the dogs had travelled with their owners to Mediterranean countries or to Portugal or had been imported from there. Infections with B. canis were diagnosed very frequently in dogs returning from Hungary. Of the 28 dogs with previous residence in Germany only, twelve and four animals were infected with B. canis and E. canis, respectively, and seven and five dogs were infested with R. sanguineus and D. marginatus, respectively.

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