The high‐Andean Jivarus Giglio‐Tos (Orthoptera,Acridoidea, Melanoplinae): systematics,phylogenetic and biogeographic considerations

The high‐Andean genus Jivarus Giglio‐Tos from Ecuador, Colombia and Peru is revised. Morphological cladistic analysis indicated that Jivarus montanus and the new species digiticercus sp.n. and rugosus sp.n. must be treated as a separate genus, Maylasacris gen.n. The remaining species included in the analysis are assigned to the genus Jivarus, for which the following six species groups are identified: americanus group, antisanae group, carbonelli group, cohni group, pictifrons group and jagoi group. Twenty‐nine species are recognized for Jivarus, with ten described as new: J. rectus sp.n. , J. megacercus sp.n. , J. spatulus sp . n. , J. auriculus sp.n. , J. riveti sp.n. , J. sphaericus sp.n. , J. discoloris sp . n. , J. profundus sp.n. , J. ronderosi sp.n. and J. guarandaensis sp.n. The following new synonymies are proposed: Jivarus albolineatus Ronderos with J. antisanae (Bolivar) syn.n. , J. cerdai Ronderos and J. osunai Ronderos with J. alienus (Walker) syn.n. , and J. rubriventris Ronderos with J. ecuadorica (Hebard) syn.n. ; the new combinations Jivarus ecuadorica (Ronderos) comb.n. and Maylasacris montanus (Ronderos) comb.n. are proposed. Keys to the species of the genera and a review of the morphological characters defining the taxa are provided. Patterns of distribution of the clades coincide with the geography of the northern Andes of Colombia and Ecuador. Areas of endemism of the Jivarus species groups and Maylasacris are delimited by both the high‐altitude curves, including transverse zones, and the drier climates of the intra‐Andean valleys, clearly indicating recent, post‐glacial palaeogeography, as shown also in vegetation distributions. This paper has been formatted with many embedded links to images of type and paratype specimens, maps based on geo‐referenced specimen data and species keys available on the Orthoptera Species file online ( http://orthoptera.speciesfile.org ).     

A revision of the Haploporinae Nicoll, 1914 (Digenea: Haploporidae) from mullets (Mugilidae): <Emphasis Type="Italic">Haploporus</Emphasis> Looss, 1902 and <Emphasis Type="Italic">Lecithobotrys</Emphasis> Looss, 1902

The status of the nominal species of Haploporus Looss, 1902 and Lecithobotrys Looss, 1902 is re-assessed by means of a comparative morphological study based on newly collected specimens from the western Mediterranean, the re-examination of museum material and a critical evaluation of published data. H. benedeni (Stossich, 1887) (type-species) is described and H. lateralis Looss, 1902 is considered to be its junior synonym. Additional data are given for H. pseudoindicus Rekharani & Madhavi, 1985, H. spinosus Machida, 1996 and H. magnisaccus Machida, 1996. Species parasitising Valamugil spp. from the Indo-West Pacific region, H. indicus Rekharani & Madhavi, 1985, H. spinosus, H. magnisaccus, H. mugilis Liu & Yang, 2002 and H. muscolosaccus Machida, 2003, are considered incertae sedis with respect to their generic affiliation. H. pacificus (Manter, 1963) (syn. Neohaploporus pacificus Manter, 1963), H. pseudoindicus and H. musculosaccus are designated as species inquirendae and H. lossii Al-Bassel, 1990 is considered to be a nomen nudum. Lecithobotrys putrescens Looss, 1902 is described based on newly collected material from Liza spp. Pseudolecithobotrys n. g. is erected to accommodate Lecithobotrys stomachicola Machida, 1996, as P. stomachicola (Machida, 1996) n. comb., from the North Pacific. L. aegyptiacus Hassan, El-Aziz, Khidr & Abu Samak, 1990 is considered to be a synonym of Saccocoelium tensum Looss, 1902, and L. brisbanensis (Martin, 1974) (syn. Paralecithobotrys brisbanensis Martin, 1974), L. vitellosus Sharma & Gupta, 1970 and L. suezcanali Nisreen Ezz El-Dien, Abdel-Rahman, El-Gawady, Imam & Fahmy, 1990 are regarded as species inquirendae. New generic diagnoses are presented for both Haploporus and Lecithobotrys.

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Entamoeba Clone‐Recognition Experiments: Morphometrics,Aggregative Behavior,and Cell‐Signaling Characterization

Studies on clone‐ and kin‐discrimination in protists have proliferated during the past decade. We report clone‐recognition experiments in seven Entamoeba lineages (E. invadens IP‐1, E. invadens VK‐1:NS, E. terrapinae, E. moshkovskii Laredo, E. moshkovskii Snake, E. histolytica HM‐1:IMSS and E. dispar). First, we characterized morphometrically each clone (length, width, and cell‐surface area) and documented how they differed statistically from one another (as per single‐variable or canonical‐discriminant analyses). Second, we demonstrated that amebas themselves could discriminate self (clone) from different (themselves vs. other clones). In mix‐cell‐line cultures between closely‐related (E. invadens IP‐1 vs. E. invadens VK‐1:NS) or distant‐phylogenetic clones (E. terrapinae vs. E. moshkovskii Laredo), amebas consistently aggregated with same‐clone members. Third, we identified six putative cell‐signals secreted by the amebas (RasGap/Ankyrin, coronin‐WD40, actin, protein kinases, heat shock 70, and ubiquitin) and which known functions in Entamoeba spp. included: cell proliferation, cell adhesion, cell movement, and stress‐induced encystation. To our knowledge, this is the first multi‐clone characterization of Entamoeba spp. morphometrics, aggregative behavior, and cell‐signaling secretion in the context of clone‐recognition. Protists allow us to study cell–cell recognition from ecological and evolutionary perspectives. Modern protistan lineages can be central to studies about the origins and evolution of multicellularity.     

Validation of <Emphasis Type="Italic">Christianella</Emphasis> Guiart, 1931 (Cestoda: Trypanorhyncha) and its taxonomic relationship with <Emphasis Type="Italic">Grillotia</Emphasis> Guiart, 1927

Christianella Guiart, 1931 (Cestoda: Trypanorhyncha) is redefined as a subgenus of Grillotia Guiart, 1927 based on the type-species, G. (C.) minuta (van Beneden, 1849), from the elasmobranch Squatina squatina (Linnaeus). Grillotia smarisgora (Wagener, 1854) is treated as a synonym of G. (C.) minuta, as are G. angeli Dollfus, 1969 and G. bothridiopunctata Dollfus, 1969. Other species included in the subgenus are G. (C.) carvajalregorum Menoret & Ivanov, 2009 (formerly Progrillotia dollfusi Carvajal & Rego, 1983), G. (C.) australis Beveridge & Campbell, 2001, G. (C.) longispinis (Linton, 1890) n. comb. (formerly Rhynchobothrium longispine Linton, 1890) and G. (C.) yuniariae Palm, 2004. The subgenus is similar to Grillotia Guiart, 1927 (sensu stricto), having two bothria and an atypical heteroacanthous armature, but differs in having a single row of intercalary hooks, fewer, elongate segments with testes often in longitudinal columns, a distinctive basal armature, an internal seminal vesicle which extends beyond the cirrus or hermaphroditic sac and no uterine pore. The adults of three species are known, all parasitising members of Squatina Duméril.     

Novel N‐Cycloalkylcarbonyl‐N′‐arylthioureas: Synthesis,Design, Antifungal Activity and Gene Toxicity

A synthesis method of novel N‐cycloalkylcarbonyl‐N′‐arylthioureas was developed. It consists of sequential addition of equimolecular amounts of ammonium isothiocyanate and substituted anilines to cycloalkylcarbonyl chlorides. The identity and purity of products were confirmed by LC/MS spectra, their structure by elemental analysis, IR and ¹H‐NMR spectra. Preliminary antimicrobial screening for standard microorganisms and molecular docking allowed to select several structures for antifungal and genetic toxicity studies. Conducted in vitro screening of 9 compounds for antifungal potential against 11 phytopathogenic fungi and three Phytophthora strains revealed that two N‐(arylcarbamothioyl) cyclopropanecarboxamides at a concentration of 50 μg/ml exhibited activities comparable to the standard antifungal agent ‘Cyproconazole’. Analysis of mutagenicity of novel thioureas using the Salmonella reverse mutagenicity assay (‘Ames Test’) showed a low gene‐toxicity profile.     

Taxonomy of the ‘Synargis axenus complex’ belonging to the ‘Synargis regulus’ species group,with a phylogenetic reassessment of the genus Synargis Hübner, 1819 (Lepidoptera: Riodinidae: Nymphidiini)

Three new species of Synargis Hübner, 1819, from Paraguay and southern and central Brazil are described: Synargis fandanga sp. nov. from Paraguay (Amambay and Paraguari) and southern Brazil (Paraná and Santa Catarina), Synargis rasqueada sp. nov. from central Brazil (Mato Grosso), and Synargis gorpa sp. nov. from southern Brazil (Paraná, Santa Catarina, and Rio Grande do Sul). Lectotypes are designated for Lemonias axenus Hewitson, 1876, Ematurgina axenus ochrophlegma Sitchel, 1911, Ematurgina acervata Seitz, 1932, and Ematurgina perrupta Seitz, 1932. Ematurgina ochrophlegma f. dissimilis Hayward, 1949, is a new synonym of Synargis bifasciata (Mengel, 1902), and Ematurgina ochrophlegma f. distincta Hayward, 1949, is a new synonym of Synargis axenus (Hewitson, 1876). The revalidation of E. perrupta Seitz, 1932, and the new status Synargis ochrophlegma (Stichel, 1911) are proposed. Ematurgina perrupta ab. roeberi Seitz, 1932, and Ematurgina bifasciata ochrophlegma ab. leucomelaina Breyer, 1930, are considered unavailable names. Based on a previous phylogenetic hypothesis, the phylogeny of the genus Synargis is reassessed, adding these new and revalidated taxa, and nine additional characters. The ‘Synargis regulus’ species group and the ‘Synargis axenus complex’ are recovered as monophyletic, with S. gorpa sp. nov. sister to the remaining species of the ‘S. axenus complex’. Additionally, an up‐to‐date geographical distribution map and a dichotomous key are provided, and the taxonomy of the taxa involved is discussed. © 2013 The Linnean Society of London     

<Emphasis Type="Italic">Legionella</Emphasis>, Protozoa,and Biofilms: Interactions Within Complex Microbial Systems

Currently, the investigation of Legionella ecology falls into two distinct areas of research activity: (1) that Legionella multiply within water sources by parasitizing amoebic or ciliate hosts or (2) that Legionella grows extracellularly within biofilms. Less focus has been given to the overlaps that may occur between these two areas or the likelihood that Legionella employs multiple survival strategies to persist in water sources. It is likely that Legionella interacts with protozoa, bacteria, algae, fungi, etc., and biofilm components in a more complex fashion than multiplication or death due to the presence or absence of single components of these complex microbial systems. This paper addresses gaps that exist in the understanding of Legionella ecology and serves to pinpoint areas of future research. To assume that only one other class of organism is important to Legionella ecology may limit our understanding of how this bacterium proliferates in heated water sources and also limit our strategies for its control in the built environment.     

Two new species of <Emphasis Type="Italic">Anthobothrium</Emphasis> van Beneden, 1850 (Tetraphyllidea: Phyllobothriidae) from carcharhinid sharks,with a redescription of <Emphasis Type="Italic">Anthobothrium laciniatum</Emphasis> Linton, 1890

Anthobothrium laciniatum Linton, 1890 is redescribed based on specimens taken from the dusky shark Carcharhinus obscurus (Lesueur) collected from the Northwestern Atlantic Ocean, and a neotype is designated. A. laciniatum differs from A. cornucopia van Beneden, 1850, A. altavelae Euzet & Ben Hassine, 2002, A. lesteri Williams, Burt & Caira, 2004 and A. spinosum Subhapradha, 1955 in total length. It further differs from A. cornucopia, A. altavelae and A. spinosum in proglottid number, and differs from A. galeorhini Suriano, 2002, A. cornucopia, and A. spinosum in testis number. A. lyndoni n. sp. is described from the sandbar shark C. plumbeus (Nardo). This new species differs from A. laciniatum in ovarian width and from A. cornucopia, A. altavelae, A. galeorhini and A. spinosum in the total number of proglottids. It further differs from A. cornucopia, A. galeorhini, and A. spinosum in total length, and from A. cornucopia and A. galeorhini in the number of testes. A. lyndoni n. sp. differs from A. lesteri in bothridial muscle and ovarian morphology. Anthobothrium caseyi n. sp. is described from Prionace glauca (Linnaeus). This new species differs conspicuously from the other six species of Anthobothrium van Beneden, 1850 (sensu stricto) in the shape of its proglottid laciniations. The taxonomic status of 43 species that have been associated with Anthobothrium is addressed. Taxonomic actions regarding Anthobothrium during the past century have resulted in a polyphyletic taxon.     

Effects of ambient O<Subscript>3</Subscript> on wheat during reproductive development: Gas exchange,photosynthetic pigments,chlorophyll fluorescence,and carbohydrates

The current concentrations of O₃ have been shown to cause significant negative effects on crop yield. The present levels of ozone may not induce visible symptoms in most of plants, but can result in substantial losses in reproductive output. This paper considers the impact of ambient O₃ on gas exchange, photosynthetic pigments, chlorophyll (Chl) fluorescence and carbohydrate levels in the flag leaf of wheat plants during various stages of reproductive development using open-top chambers. Mean O₃ concentration was 45.7 ppb during wheat growth and 50.2 ppb after flag leaf development. Reproductive stage showed higher exceedence of O₃ above 40 ppb compared to the vegetative stage. Diurnal variations in net photosynthetic rate (P _N) and stomatal conductance (g _s), intercellular CO₂ concentration (C _i), Fv/Fm ratio, photosynthetic pigments, soluble sugars, and starch were measured at 10, 30, and 50 days after flag leaf expansion (DAFE). The results showed reductions in P _N, g _s, F_v/F_m ratio, photosynthetic pigments and starch, and increases in C _i, F₀, and soluble sugars in nonfiltered chambers (NFCs) compared to filtered chambers (FCs). Maximum changes in measured parameters were observed at 50 DAFE (i.e. grain filling and setting phase). Diurnal variation in P _N showed double peaked curve in both FCs and NFCs, but delayed peak and early depression in NFCs. Stomatal conductance was significantly lower in NFCs. The study suggests that higher prevalence of ambient O₃ during reproductive development led to significant alteration in physiological vitality of wheat having potential negative influence on yield.     

Listeria monocytogenes exopolysaccharide: origin,structure, biosynthetic machinery and c‐di‐GMP‐dependent regulation

Elevated levels of the second messenger c‐di‐GMP activate biosynthesis of an unknown exopolysaccharide (EPS) in the food‐borne pathogen Listeria monocytogenes. This EPS strongly protects cells against disinfectants and desiccation, indicating its potential significance for listerial persistence in the environment and for food safety. We analyzed the potential phylogenetic origin of this EPS, determined its complete structure, characterized genes involved in its biosynthesis and hydrolysis and identified diguanylate cyclases activating its synthesis. Phylogenetic analysis of EPS biosynthesis proteins suggests that they have evolved within monoderms. Scanning electron microscopy revealed that L. monocytogenes EPS is cell surface‐bound. Secreted carbohydrates represent exclusively cell‐wall debris. Based on carbohydrate composition, linkage and NMR analysis, the structure of the purified EPS is identified as a β‐1,4‐linked N‐acetylmannosamine chain decorated with terminal α‐1,6‐linked galactose. All genes of the pssA‐E operon are required for EPS production and so is a separately located pssZ gene. We show that PssZ has an EPS‐specific glycosylhydrolase activity. Exogenously added PssZ prevents EPS‐mediated cell aggregation and disperses preformed aggregates, whereas an E72Q mutant in the presumed catalytic residue is much less active. The diguanylate cyclases DgcA and DgcB, whose genes are located next to pssZ, are primarily responsible for c‐di‐GMP‐dependent EPS production.     

Tetrathiafulvalene‐[2.2]paracyclophanes: Synthesis,crystal structures,and chiroptical properties

Two racemic tetrathiafulvalene‐[2.2]paracyclophane electron donors EDT‐TTF‐[2.2]paracyclophane 1 and (COOMe)₂‐TTF‐[2.2]paracyclophane 2 have been synthesized via the phosphite mediated cross coupling strategy. Chiral HPLC allowed the optical resolution of the (R_P) and (S_P) enantiomers for both compounds. Solid‐state structures of (R_P)‐ 1 and (rac)‐ 2 have been determined by single crystal X‐ray analysis. Intermolecular π‐π and S???S interactions are disclosed in the packing. Single crystal X‐ray analysis of (R_P)‐ 1 combined with experimental and theoretical circular dichroism spectra allowed the assignment of the absolute configuration of the enantiomers of 1 and 2 .     

Study on two unrecorded genera,Barycnemis Förster and Phradis Förster (Hymenoptera: Ichneumonidae: Tersilochinae), from South Korea

Six genera of the subfamily Tersilochinae (Barycnemis, Diaparsis, Gelanes, Phradis, Probles and Tersilochus) are recognized in South Korea. Two genera, Diaparsis and Tersilochus, were previously recognized from South Korea, whereas the other four genera are recorded from this country for the first time. All genera found in South Korea, except the almost cosmopolitan Diaparsis, are entirely or predominantly Holarctic. A key to the six genera of Tersilochine occurring in South Korea is provided. Two genera of Korean Tersilochinae (Barycnemis and Phradis) are reviewed here, and a key to the other four species (B. bellator, B. dissimilis, P. kyushuensis and P. nikishenae) is provided in this paper.     

HIBERNATION CHEZ TROIS ESPÈCES DE MÉTOPIINES: HYMENOPTERA,ICHNEUMONIDAE

Résumé Les conditions de l'hibernation sont étudiées chez les trois espèces de Métopiines Chorinaeus funebris Gravenhorst, Triclistus podagricus Gravenhorst et Triclistus pygmaeus Cresson. Ces espèces hibernent à des stades de développement compris entre l'éonymphe et l'adulte non émergé (Tableau I). La durée de l'incubation complémentaire après réactivation par le froid à 2^o varie entre 10 et 30 jours en moyenne (Tableaux II, IV et V). Chez T. podagricus et T. pygmaeus, on a observé également des émergences d'adultes avant l'hiver (Tableau III). I1 semble que T. podagricus ne parvienne au stade d'imago hibernant qu'après une baisse de la température d'élevage. Aucune souche dépourvue de diapause n'a pu être créée, mais pour les trois espèces, deux générations annuelles seraient possibles sous conditions contrôlées. Enfin, l'utilisation éventuelle de T. pygmaeus pour une lutte biologique contre Zeiraphera diniana Guénée est évoquée.

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Summary The conditions of hibernation have been investigated in the three metopiine species Chorinaeus funebris, Triclistus podagricus and Triclistus pygmaeus, all entomophagous on Zeiraphera diniana Guénée (Lepidoptera, Tortricidae) in the Engadine Valley (Switzerland). The hibernating stages are eonymph, pronymph or unemerged adult inside the host pupae (Table I). The length of the complementary incubation after reactivation at 2^o varies between 10–30 days (Tables II, IV and V). A small number of T. podagricus and of T. pygmaeus regularly appear as adults before winter, in rearings of both laboratory and field populations (Table III). It seems that T. podagricus larvae need a reduction in the initial rearing temperature before full development is achieved and the hibernating stage is reached (imago). No strain without diapause could be obtained for any of the three species, but the rearing of two annual generations under controlled conditions is possible in the laboratory. The eventual use of T. pygmaeus as a control agent against Z. diniana is discussed.

     

Length–weight relationship and condition factor of wild,grow‐out and ‘loose‐shell affected’ giant tiger shrimp,Penaeus monodon (Fabricius, 1798) (Decapoda: Penaeidae)

Length–weight relationships and condition factors of wild, cultured, and cultured loose‐shell affected Penaeus monodon (2,609 specimens total) were studied from March to August 2007. The regression equation for healthy cultured shrimps was log W = ?1.811 + 2.721 log L (r² = 0.71); log W = ?1.444 + 2.485 log L (r² = 0.91) for wild shrimps; and log W = ?1.112 + 2.237 log L (r² = 0.92) for loose‐shell affected shrimps. All shrimps showed negative allometric growth, although ancova indicated significant differences (P < 0.05) among them. This study presents the first known reference dealing with LWRs and condition factors of cultured, wild and loose‐shell affected P. monodon.     

Transformation of chickpea: effect of genotype,explant, <Emphasis Type="Italic">Agrobacterium</Emphasis>-strain and composition of culture medium

Reproducible and high-frequency transgenic plant regeneration from callus and embryo axes of four different genotypes of chickpea (Cicer arietinum) was achieved after Agrobacterium-mediated transformation. Three different strains of Agrobacterium (EHA105, AGL1 and LBA4404) harboring the binary vector pCAMBIA1301 containing β-glucuronidase (GUS) and hygromycin phosphotransferase (hpt) genes under the control of a CaMV35S promoter were used. The highest number of transgenic plants was obtained from cotyledonary node-derived calli of genotype Pusa-256. A highly efficient rooting was achieved on Murashige and Skoog medium supplemented with indole-3-butyric acid. The stable integration of the gene was confirmed by molecular analyses of the transformed plants. Inheritance of GUS and hpt gene was followed through two generations and they showed the expected 3:1 inheritance.     

The complete mitochondrial genome of <Emphasis Type="Italic">Thrinchus schrenkii</Emphasis> (Orthoptera: Caelifera,Acridoidea, Pamphagidae)

Complete nucleotide sequences of mitochondrial genome (mitogenome) of Thrinchus schrenkii (Orthoptera: Acridoidea: Pamphagidae) were determined. It is 15672 bp in length and contains 71.2% A + T. All T. schrenkii protein-coding sequences except for the cytochrome oxidase subunit I (COI) start with a typical ATN codon. Instead, CCG, which is a rare but possible initiation codon, is located at the initiation context of COI. The usual termination codons (TAA and TAG) were found from 12 PCGs. However, the ND5 had incomplete termination codon (T). All tRNA genes could be folded into the typical cloverleaf secondary structure, excluding tRNA ^Ser(AGN) which forms another structure according to the Steinberg–Cedergren tertiary structure. The sizes of the large and small ribosomal RNA genes are 1319 and 848 bp, respectively. The A + T content of the A + T-rich region is 78.7%, which is the lowest among the known mitogenome of Acridoidea.     

Mobile genetic elements: in silico,in vitro,in vivo

Mobile genetic elements (MGEs), also called transposable elements (TEs), represent universal components of most genomes and are intimately involved in nearly all aspects of genome organization, function and evolution. However, there is currently a gap between the fast pace of TE discovery in silico, driven by the exponential growth of comparative genomic studies, and a limited number of experimental models amenable to more traditional in vitro and in vivo studies of structural, mechanistic and regulatory properties of diverse MGEs. Experimental and computational scientists came together to bridge this gap at a recent conference, ‘Mobile Genetic Elements: in silico, in vitro, in vivo’, held at the Marine Biological Laboratory (MBL) in Woods Hole, MA, USA.     

Systematic revision of Hoggicosa Roewer, 1960, the Australian ‘bicolor’ group of wolf spiders (Araneae: Lycosidae)

The Australian wolf spider genus Hoggicosa Roewer, 1960 with the type species Hoggicosa errans (Hogg, 1905) is revised to include ten species: Hoggicosa alfi sp. nov. ; Hoggicosa castanea (Hogg, 1905) comb. nov. (= Lycosa errans Hogg, 1905 syn. nov. ; = Lycosa perinflata Pulleine, 1922 syn. nov. ; = Lycosa skeeti Pulleine, 1922 syn. nov. ); Hoggicosa bicolor (McKay, 1973) comb. nov. ; Hoggicosa brennani sp. nov. ; Hoggicosa duracki (McKay, 1975) comb. nov. ; Hoggicosa forresti (McKay, 1973) comb. nov. ; Hoggicosa natashae sp. nov. ; Hoggicosa snelli (McKay, 1975) comb. nov. ; Hoggicosa storri (McKay, 1973) comb. nov. ; and Hoggicosa wolodymyri sp. nov. The Namibian Hoggicosa exigua Roewer, 1960 is transferred to Hogna, Hogna exigua (Roewer, 1960) comb. nov. A phylogenetic analysis including nine Hoggicosa species, 11 lycosine species from Australia and four from overseas, with Arctosa cinerea Fabricius, 1777 as outgroup, supported the monophyly of Hoggicosa, with a larger distance between the epigynum anterior pockets compared to the width of the posterior transverse part. The analysis found that an unusual sexual dimorphism for wolf spiders (females more colourful than males), evident in four species of Hoggicosa, has evolved multiple times. Hoggicosa are burrowing lycosids, several constructing doors from sand or debris, and are predominantly found in semi‐arid to arid regions of Australia. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 83–123.