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Cophylogenetic relationships between penguins and their chewing lice
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1 doi: /j x Cophylogenetic relationships between penguins and their chewing lice J. C. BANKS,* R.L.PALMA & A. M. PATERSON* *Bioprotection and Ecology Division, Lincoln University, Canterbury, New Zealand Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand Keywords: chewing lice; coevolution; cophylogeny; cospeciation; failure to speciate; inertia; penguins; Phthiraptera; Sphenisciformes. Abstract It is generally thought that the evolution of obligate parasites should be linked intimately to the evolution of their hosts and that speciation by the hosts should cause speciation of their parasites. The penguins and their chewing lice present a rare opportunity to examine codivergence between a complete host order and its parasitic lice. We estimated a phylogeny for all 15 species of lice parasitising all 17 species of penguins from the third domain of the mitochondrial 12S ribosomal rrna gene, a portion of the mitochondrial cytochrome oxidase subunit 1 gene and 55 morphological characters. We found no evidence of extensive cospeciation between penguins and their chewing lice using TreeMap 2.02b. Despite the paucity of cospeciation, there is support for significant congruence between the louse and penguin phylogenies due to possible failure to speciate events (parasites not speciating in response to their hosts speciating). Introduction It is generally thought that the evolution of obligate parasites, with few means of transferring between different host species, should be tightly linked to the evolution of their own host species (Klassen, 1992). Comparing host and parasite evolution is known as the study of codivergence, coevolution or cophylogenetic descent (Paterson & Banks, 2001). A major aim of coevolutionary studies is to determine the cophylogenetic events that have generated the present distribution of parasite species and to explain how parasites have become associated with their hosts (Brooks & McLennan, 1991). The two major paradigms to explain current hostparasite associations are either that a parasite species has switched to a new host lineage, or that each host species has inherited the association from its ancestor. The two paradigms can be summarized as association by colonization and association by descent (Brooks & Correspondence: Jonathan Banks, Entomology, 320 Morrill Hall, University of Illinois, 505 S. Goodwin Ave, Urbana, IL 61801, USA. Tel.: (+1) (217) ; fax: (+1) (217) ; McLennan, 1991) or as souvenirs and heirlooms (Kliks, 1990). Association by colonization proposes that a parasite s presence on a host is due to parasites switching between hosts (Brooks & McLennan, 1991). Association by colonization suggests that a parasite will speciate following isolation from its ancestral population after colonizing the new host, analogous to speciation following dispersal to new areas by free-living organisms. Association by descent suggests parasites have been inherited from the ancestral host. Association by descent suggests that parasites will speciate when hosts speciate, which is analogous to speciation by free-living organisms following a vicariant event (Brooks & McLennan, 1991), and is known as cospeciation or codivergence (Ronquist, 2003). Chewing lice are a useful group on which to conduct cophylogenetic studies because they are considered to have limited mobility, rely mainly on host-to-host contact for transmission (Hafner & Nadler, 1988), usually appear to have little effect on their host (Rothschild & Clay, 1952; Marshall, 1981) and survive only briefly away from their hosts (Marshall, 1981). For these reasons pocket gophers and their lice, with their remarkably congruent phylogenies (Hafner & Nadler, 156
2 Penguin-louse coevolution ; Hafner et al., 1994), have become the textbook models, e.g. Ridley (1996), of association by descent and of cospeciation between two lineages. Pocket gopher lice have also become the group in which to trial and demonstrate new methods of analysing the extent of cospeciation (for example, Page (1990), Ronquist (1995), Huelsenbeck et al. (1997,2000), Legendre et al. (2002)). However, many other host-parasite systems have considerably less codivergence than the pocket gopher-louse group. It is now recognized that events other than host switching may explain incongruence between host and parasite phylogenies without ruling out a history of association by descent. For example, duplication events (speciation by the parasite without the host speciating), and sorting events, such as missing the boat (the parasite is absent from the host population founding the new host species) or parasite extinctions (Paterson & Banks, 2001), may allow apparently incongruent parasite phylogenies to support a hypothesis of association by descent. Indeed, an analysis of pocket gopher and louse phylogenies allowing these events found even more codivergence between the hosts and their parasites (Page, 1990). A rigorous cophylogenetic study incorporates four stages: first, a robust alpha taxonomy of both hosts and parasites; second, construction of accurate host and parasite phylogenies; third, quantitative comparison of the host and parasite phylogenies; fourth, statistical testing for congruence between the two phylogenies (Clayton et al., 1996). Methods to evaluate the extent of cophylogenetic descent can be divided into two groups. The first group assesses the extent of codivergence by comparing the topology of the independently derived host and parasite phylogenies. Brooks Parsimony Analysis (Brooks et al., 2001), the generalized parsimony method (Ronquist, 1995) implemented in Treefitter, reconciliation analysis (Page, 1994b) as implemented in TreeMap 1 (Page, 1995), and Jungles (Charleston, 1998) as implemented in TreeMap 2.02b (Charleston & Page, 2002), are methods that assume that accurate phylogenies for hosts and parasites are known (Huelsenbeck et al., 2000). The second group of methods does not assume accurate phylogenies are known. Methods such as Data Based Parsimony (Johnson et al., 2001), Parafit (Legendre et al., 2002), and statistical methods based on maximum likelihood (Huelsenbeck et al., 1997) or Bayesian methods (Huelsenbeck et al., 2000); allow the evaluation of codivergence between less than optimal host and parasite phylogenies. While a comparison of the merits of each method is beyond the scope of this paper, TreeMap is the leading method to analyse phylogenetic aspects of host-parasite coevolution (Brooks & McLennan, 2003). The ideal cophylogenetic study should also extensively sample parasites from the host group of interest as it increases the probability of detecting evolutionary changes (Page, 1996). In the past, cophylogenetic studies have tended to choose hosts and parasites that represent various taxonomic levels such as host families or orders and wing or body lice. For example, the gopher-louse study examined 17 louse species that were representative taxa from larger clades containing 122 recognized louse species and the 15 gopher taxa were examples from a group containing 40 species and 450 subspecies (Page, 1996). Similarly an analysis of cophylogeny between seabirds and their lice examined 14 louse species parasitising 11 host species from two relatively large host orders containing over 100 species (Paterson et al., 2000). Choosing taxa to represent higher taxonomic groups has resulted in some aspects of cophylogenetic relationships being neglected. Parasite species with multiple host species (multi-host parasites) are one group that has been especially neglected as they often parasitise closely related hosts. We compared a phylogeny estimated for all 17 species of penguins (Sphenisciformes) (Giannini & Bertelli, 2004) to a phylogeny estimated for all of the 15 species of chewing lice (Phthiraptera: Philopteridae) parasitising penguins. Our study, which is the first to examine all species of chewing lice parasitising an entire host order, did not find evidence for extensive cospeciation. Because we did not exclude any penguin louse species a priori we found several examples where it appears the lice have failed to speciate, i.e. the lice not speciating in response to their hosts speciating (Page, 1994a; Johnson et al., 2003). Failure to speciate has also been called cophylogenetic inertia (Paterson & Banks, 2001) or cophylogeny without cospeciation (Hugot et al., 2001). We suggest that failure to speciate is an additional event that needs to be considered in cophylogenetic studies that examine host parasite groups with multi-host parasites as it can markedly affect the extent of association by descent. Methods Molecular methods Lice were collected from penguins at various southern hemisphere locations (Table 1). Live penguins were restrained, sprayed with pyrethrin insecticide and the plumage searched manually for lice. Penguins found dead were taken back to field camps where they could be searched more thoroughly for lice. Louse specimens were stored in 100% ethanol at room temperature until they could be refrigerated. Frozen penguin carcasses from institutions, such as museums, were also searched and these provided numerous louse specimens from which we could extract DNA suitable for sequencing. Louse specimens were identified from morphological characters (Clay & Moreby, 1967; Banks & Palma, 2003) before the DNA was extracted. Initially, DNA was extracted from lice using the high salt method (White et al., 1990), but later Qiagen DNeasy
3 158 J. C. BANKS ET AL. Table 1 Louse collection sites and hosts. Louse species Host species Location Coordinates latitude Longitude Collector Host status Gene region sequenced A. antarcticus P. adeliae Ross Island, Antarctica S W J. C. Banks Live COI, 12S A. bifasciatus S. humboldti Coquimbo, Chile S E J. C. Banks Dead COI A. bifasciatus S. magellanicus Sea Lion Island, Falklands S W J. C. Banks Live COI, 12S A. brevipes A. patagonicus Kerguelen Is S W M. Gaulthier-Clerc Live COI, 12S E. pachyrhynchus Jackson Bay, New Zealand S E J. C. Banks Live 12S E. robustus Snares Islands, New Zealand S E J. C. Banks Live COI, 12S M. antipodes Otago Peninsula (Genbank S E A. M. Paterson Live 12S accession number Y14910) E. chrysocome New Island, Falklands S E A. van Buren Live COI chrysocome E. chrysocome filholi Snares Islands, New Zealand S E J. C. Banks Live COI, 12S E. robustus Snares Islands, New Zealand S E J. C. Banks Live COI, 12S A. demersus S. demersus Cape Town, South Africa S E J. C. Banks Dead COI, 12S A. gressitti P. papua Sea Lion Island, Falklands S E J. C. Banks Live COI, 12S A. hamiltoni E. schlegeli Macquarie Island, Australia S E K. Edge Live COI, 12S A. keleri E. chrysocome New Island, Falklands S E A. van Buren Live COI, 12S chrysocome A. macquariensis E. chrysocome filholi Macquarie Island, Australia S E K. Edge Live COI, 12S A. mawsoni A. forsteri Ross Island, Antarctica S W P. Ponganis Live COI, 12S A. vanalphenae M. antipodes Otago Museum carcass Unrecorded Unrecorded J. C. Banks Dead COI, 12S A. waterstoni E. minor Coromandel Peninsula S E J. C. Banks Dead COI, 12S Wellington Harbour S E J. C. Banks Live COI, 12S Chatham Island S W J. C. Banks Live COI, 12S Banks Peninsula S E J. C. Banks Live COI, 12S Otago Harbour S E J. C. Banks Dead COI, 12S Phillip Island, Australia S E R. Jessop Dead COI, 12S N. demersus A. patagonicus Macquarie Island, Australia S E K. Edge Live COI, 12S COI, cytochrome oxidase subunit 1. kits were used following the protocol developed by Cruickshank et al. (2001). The head of the louse was separated from the body and then incubated for 48 h at 55 C in a solution containing proteinase K. The DNA was extracted as outlined in the DNeasy protocol. The head and body were retained as voucher specimens. Among the louse species parasitising penguins, we were unable to obtain specimens of A. bicornutus for molecular analysis. Molecular characters for A. bicornutus were coded as missing for the phylogenetic analysis. Portions of the 12S and COI regions were amplified from total genomic extracts using polymerase chain reaction (PCR). All PCR was carried out using a Perkin Elmer 2400 thermal cycler. Reaction profiles for each region were 94 C for 4 min, 40 cycles of 94 C for 20 s, annealing temperature as in Tables 3 and 4 for 30 s, 72 C for 50 s, and finally 72 C for 5 min. See Table 2 for primer sequences. PCR consisted of 2.5 ll of 10 buffer (Roche), 2.5 ll of dntps (1 mm), MgCl 2 (25 mm) as outlined in Tables 3 and 4, 1 ll of each primer (10 lm), 0.25 ll Taq (5 units ll )1, Roche), ll of DNA and water to 25 ll for each reaction. A negative control was incorporated in each amplification round using water rather than DNA. Initially, excess primers and salts were removed from the PCR product by precipitation with isopropanol in the presence of 2.5 M NH 4 Ac followed by a 70% ethanol wash. Later, PCR product was purified using Qiagen Concert rapid PCR purification system kits. Purified PCR fragments were sequenced using BigDye Termination Mix (Perkin-Elmer) and run out on an ABI 373 automated sequencer. Both the sense and antisense strands were sequenced. Sequences were deposited in GenBank (http://www.ncbi.nlm.nih) (Benson et al., Table 2 Primer sequences used in PCR. Primer 1 Sequence Reference 12sai AAACTAGGATTAGATACCCTATTAT Simon et al. (1994) 12sbi AAGAGCGACGGGCGATGTGT Simon et al. (1994) L1091 AAAAAGCTTCAAACTGGGATTA Kocher et al. (1989) GATACCCCACTAT 12sfia CGGGCGATGTGTRCATTMTT This paper C1-J-1718 GGAGGATTTGGAAATTGATTAGTTCC Simon et al. (1994) C1-N-2191 CCCGGTAAAATTAAAATATAAACTTC Simon et al. (1994) COI1 TAAATAAYATRAGDYTTTGDCTKCT This paper COI1R CCYCCNGMNGGRTCAAAAAARGA This paper COI, cytochrome oxidase subunit 1.
4 Penguin-louse coevolution 159 Table 3 Reaction conditions and primers used for PCR of the third domain of the mitochondrial 12S ribosomal rrna region (see text for more details). Species Primer 1 Primer 2 Extra MgCl 2 (ll of25mm) A. antarcticus 12sai 12sfia A. bifasciatus L sbi A. brevipes 12sai 12sfia 0 56 L sbi sai 12sfia 0 47 A. demersus L sbi A. gressitti 12sai 12sfia 0 47 A. hamiltoni L sbi A. keleri 12sai 12sfia 0 47 A. macquariensis L sbi A. mawsoni 12sai 12sfia 0 47 A. vanalphenae 12sai 12sfia 0 47 A. waterstoni L sbi N. demersus L sbi Annealing temp ( C) Table 4 Reaction conditions and primers used for PCR of a portion of the COI region. Species Primer 1 Primer 2 Extra MgCl 2 (ll of25mm) A. antarcticus C1-J-1718 C1-N A. bifasciatus C1-J-1718 C1-N A. brevipes C1-J-1718 C1-N C1-J-1718 C1-N COI1 COI1R A. demersus C1-J-1718 C1-N A. gressitti C1-J-1718 C1-N A. hamiltoni C1-J-1718 C1-N A. keleri COI1 COI1R A. macquariensis COI1 COI1R A. mawsoni COI1 COI1R A. vanalphenae C1-J-1718 C1-N A. waterstoni C1-J-1718 C1-N N. demersus C1-J-1718 C1-N COI, cytochrome oxidase subunit 1. Annealing temp ( C) 2002) (GenBank accession numbers AF AF491758, AY AY229936). Our PCR results did not contain ghost bands and had few sequence ambiguities that are both usual signs of the amplification of pseudogenes (Bensasson et al., 2001). For species, such as A. antarcticus, with large insertions, we sequenced 12S twice, using different primer sets and obtained identical sequences. The COI sequences were not punctuated by stop codons when translated into amino acids. Louse taxonomy followed Hopkins & Clay (1953), Clay (1967), and Banks & Palma (2003). We considered the louse species A. strutheus as a nomen dubium following the discussion in Clay (1967). Because there were few genetic differences between individual lice of the same species parasitising different host species (see Table 1 for duplicates sequenced), we used one representative sequence for each gene region from each louse morpho-species in the phylogenetic analysis. COI sequences were aligned using Clustal X (Thompson et al., 1997), then adjusted manually using sequential pairwise comparisons. Alignment of the 368 base pair fragment of COI was straightforward because there were few insertions or deletions (indels). Postulated gaps within COI were adjusted with respect to the amino acid sequence so that codons were not split. Alignment of 12S was somewhat problematic and we aligned 12S manually with respect to the secondary structure of A. waterstoni (Page et al., 2002). Unalignable regions were deleted leaving 293 base pairs of 12S for analysis. Louse phylogeny A mixed model Bayesian analysis using Mr Bayes 3 (Ronquist & Huelsenbeck, 2003) was conducted on 55 louse morphological characters (Banks & Paterson, 2004) and the COI and 12S molecular data. The general time reversible model plus gamma (Rodríguez et al., 1990; Yang et al., 1994) was chosen to analyse the genetic data based on the Akaike information criteria in ModelTest (Posada, 2000). Three independent Bayesian analyses were run for each gene to ensure proper sampling of tree space. Two runs were made of generations, and one of generations, with four chains using flat priors and mixed models, saving trees every 100 generations. All trees prior to stationarity were discarded (1000 or 2000 trees depending on the number of generations). Each run produced 50% majority rule trees of the same topology and converged on similar likelihood values after trees prior to stationarity were discarded. A single run of generations was also conducted with 5000 trees prior to stationarity discarded. Nesiotinus demersus was chosen as the outgroup as, although deeper relationships of ischnoceran lice are not well resolved (Cruickshank et al., 2001), we are confident that N. demersus is not part of Austrogoniodes based on substantial morphological differences. Cophylogenetic analysis Penguin louse associations were collated from several sources (Clay, 1967; Watson, 1967; Pilgrim & Palma, 1982; Palma, 1996,1999; Price et al., 2003) and are shown in Fig. 1. Associations of doubtful validity, for example, lice collected from penguins kept in zoos were not considered. A full list of doubtful records is given in Banks & Paterson (2004). We analysed cophylogenetic relationships between penguins and their chewing lice using a penguin phylogeny estimated from 70 integumentary and breeding characters (Giannini & Bertelli, 2004). However, the
5 160 J. C. BANKS ET AL. A. patagonicus A. forsteri P. antarctica P. papua P. adeliae E. pachyrhynchus E. chrysocome E. chrysolophus E. schlegeli E. robustus E. sclateri M. antipodes S. humboldti S. demersus S. magellanicus S. mendiculus Ea. minor NZ N. demersus A. brevipes A. mawsoni A. gressitti A. antarcticus A. bicornutus A. hamiltoni A. macquariensis A. vanalphenae A. bifasciatus A. demersus A. waterstoni Fig. 1 Tanglegram of penguins and their lice. Lines connecting penguins and lice indicate associations. penguin phylogeny we analysed differed slightly from the published phylogeny as we split Eudyptula minor into two taxa following the discussion in Banks et al. (2002) and combined the two Eudyptes chrysocome subspecies as a single terminal taxon. The cophylogenetic history of the penguins and their lice was reconstructed using TreeMap 1 and TreeMap 2.02b. TreeMap 2.02b requires that multi-host lice (lice parasitising several host species) are subdivided into dummy lineages (Page & Charleston, 2002) to generate hypotheses of cophylogeny correctly. The topology of that portion of the phylogeny containing dummy lineages mirrored that of the host phylogeny. Because TreeMap 2.02b requires the addition of dummy taxa, the significance of the cophylogenetic relationship between the penguins and their lice, without dummy taxa, was assessed using TreeMap 1. Parafit (Legendre et al., 2002), which uses the patristic distances of a host and parasite phylogeny transformed into principle coordinates (Gower, 1966), was used to test the extent of a global hypothesis of coevolution between the lice and their hosts. Treefitter 1.0 (http://www. ebc.uu.se/systzoo/research/treefitter/treefitter.html) was also used to implement the generalized parsimony method of testing for a cophylogenetic relationship between penguins and their chewing lice. Costs used were cospeciation ¼ 0, duplication and sorting ¼ 1, and switching ¼ The cost of fitting the penguin phylogeny to the louse phylogeny was compared to the cost of fitting the host tree to random parasite trees. Both of these methods are able to deal with multi-host lice. Results The 50% majority rule consensus tree of trees estimated from the Bayesian analysis is shown in Fig. 2. The analysis found strong support for two groups within Austrogoniodes. The first was a concii clade of, A. bicornutus, A. hamiltoni, A. keleri and A. macquariensis and the second was a cristati clade of A. vanalphenae,, Fig. 2 Fifty per cent majority rule consensus tree of trees for the penguin chewing lice estimated from a Bayesian analysis of sequences from portions of the third domain of the mitochondrial 12S ribosomal rrna gene (12S), cytochrome oxidase subunit 1 (COI) gene and 55 morphological characters. Clade support values are shown above the lines. A. demersus and A. bifasciatus. Austrogoniodes antarcticus and A. gressitti, as well as A. brevipes and A. mawsoni were also strongly supported as two pairs of sister taxa. Cophylogenetic analysis TreeMap TreeMap 2.02b (Charleston & Page, 2002), with cospeciation events weighted as 0, duplications, lineage losses and host switching events weighted as one and up to two host switches allowed (the maximum number that was feasible with the computer power available), found six scenarios that maximized cospeciation. With zero host switches only three cospeciation events were found
6 Penguin-louse coevolution 161 (Figs 3 and 4). With one host switch there were four cospeciation events (Figs 4 and 5) or three cospeciation events with two host switches. TreeMap 1, with zero host switches, found the same reconstruction as TreeMap 2.02b using similar constraints. When cospeciation events due to dummy taxa were excluded, the number of cospeciation events did not differ significantly from the number obtained when 1000 random louse phylogenies were compared to the host phylogeny (P ¼ 0.89) using TreeMap 1. When dummy Fig. 3 Penguin and louse cophylogenetic history estimated using TreeMap 2.02b with no host switching (the concii lineage is shown in Fig. 4 for clarity). The thick lines represent the penguin phylogeny, thin lines the louse phylogeny. The louse phylogeny is stacked over the penguin phylogeny but offset to the left and down relative to the penguin phylogeny. Filled circles ¼ cospeciation events, open circles ¼ possible failure to speciate events, squares ¼ duplication events. Louse lineages that do not form terminal branches have undergone sorting events. Fig. 4 Megadyptes and Eudyptes penguin and the louse concii clade (A. bicornutus,, A. hamiltoni, A. keleri and A. macquariensis) cophylogenetic history estimated using TreeMap 2.0b with zero or one host switch. The thick lines represent the penguin phylogeny, thin lines the louse phylogeny. The louse phylogeny is stacked over the penguin phylogeny but offset to the left and down relative to the penguin phylogeny. Open circles ¼ possible failure to speciate events, squares ¼ duplication events. Louse lineages that do not form terminal branches have undergone sorting events. From Figs 3 and 5 Penguin Louse E. chrysolophus A. macquariensis A. bicornutus E. schlegeli A. macquariensis A. hamiltoni E. chrysocome A. macquariensis A. keleri A. hamiltoni E. robustus E. sclateri E. pachyrhynchus M. antipodes Pygoscelis lineage
7 162 J. C. BANKS ET AL. Penguin Louse To Fig. 4 E. chrysolophus A.cristati E. schlegeli E. chrysocome E. sclateri E. robustus E. pachyrhynchus M. antipodes A. vanalphenae P. papua A.gressitti P. adeliae A. antarcticus P. antarctica A. gressitti A. patagonicus A. brevipes N. demersus A. forsteri A. mawsoni S. mendiculus A. demersus S. magellanicus A. bifasciatus S. demersus A. demersus S. humboldti A. bifasciatus Ea. minor A. waterstoni Ea. minor A. waterstoni Fig. 5 Penguin and louse cophylogenetic history with one host switch (the concii lineage is shown in Fig. 4 for clarity) estimated using TreeMap 2.02b. The thick lines represent the penguin phylogeny, thin lines the louse phylogeny. The louse phylogeny is stacked over the penguin phylogeny but offset to the left and down relative to the penguin phylogeny. Filled circles ¼ cospeciation events, open circles ¼ possible failure to speciate events, squares ¼ duplication events. The arrow indicates the host switch from the ancestor of Eudyptes to the ancestor of Spheniscus. Louse lineages that do not form terminal branches have undergone sorting events. lineages were included in both TreeMap analyses, there was significantly more cospeciation than if the penguin phylogeny had been compared to 1000 random louse phylogenies (P < 0.01). Host switches TreeMap 2.02b proposed a host switch from the ancestor of the crested penguins to the ancestor of the spheniscid penguins that gave rise to A. bifasciatus and A. demersus (Fig. 5). The scenario with two host switches included the switch to the spheniscid penguins and a switch from the ancestor of Pygoscelis, Megadyptes and Eudyptes to Eudyptula. TreeMap 1 also found the same scenarios. The three other scenarios from TreeMap 2.02b with two host switches suggested the host switches were by multi-host lice expanding their range to new host taxa. For example, one scenario proposed a switch by A. hamiltoni from E. chrysocome to E. schlegeli. Parafit and Treefitter Parafit rejected the null hypothesis that the penguins and their lice have evolved independently (P ¼ 0.001), indicating that there is significant cophylogenetic history within the penguin-louse group. Treefitter also found that the cost of fitting the penguin phylogeny to the louse phylogeny was significantly less than the costs of fitting the penguin phylogeny to random trees (P < 0.003) when the cost of switching was set from 2 to 10. The cost was not significantly different if the cost of switching was set to one. Discussion TreeMap 2.02b found significant shared cophylogenetic history if we included cospeciation events due to dummy taxa, i.e. branches added to the host and parasite trees so that all hosts have only one parasite. If dummy branches are included in a TreeMap 2.02b analysis it is more appropriate to consider cospeciation between dummy branches as host tracking events, i.e. cospeciation and failure to speciate events. If this is done, TreeMap 2.02b then assesses the significance of the maximum extent of association by descent rather than the extent of cospeciation. Parafit and Treefitter do not require the addition of dummy taxa. The Parafit analysis found that the penguin and louse phylogenies were significantly more similar to each other than 999 random phylogenies, suggesting a coevolutionary relationship between the penguins and the lice. Treefitter found the cost of fitting the penguin
8 Penguin-louse coevolution 163 phylogeny to the louse phylogeny was significantly less than cost of fitting it to random phylogenies but only if host switching is made more difficult (i.e. cost >1) than duplication and sorting events. It has been argued that setting a higher cost for host switching is justified if parasites do not have a dispersal phase (Desdevies et al., 2002). As lice rely on host-to-host contact for transmission (Hafner & Nadler, 1988), assigning a higher cost to host switching may be justified. Interpreting the reasons for the presence of multi-host parasites markedly affects the extent of association by descent. It has been suggested that multi-host parasites could be genetically isolated, although morphologically conservative, and could be treated as separate taxa (Page et al., 2004). It has also been suggested that a parasite species could parasitise several closely related hosts if it could maintain genetic contact between populations on divergent hosts and thus have failed to speciate (Paterson & Banks, 2001; Johnson et al., 2003). Alternatively, it could be that only one host has inherited the parasite species and the rest of the associations are due to host switching (Ronquist, 2003). Several penguin species that share the same species of lice also share breeding islands or are occasional visitors, often during moulting, to the breeding sites of other penguin species (del Hoyo et al., 1992; Miskelly et al., 2001). However, there are examples of sympatric penguin species that do not share lice, for example emperor, Aptenodytes forsteri, and Adelie, P. adeliae, penguins mix at several sites in Antarctica (Marchant & Higgins, 1990) but are parasitised by different species of lice (Price et al., 2003). Therefore, host switching cannot be inferred simply by host species living in sympatry. Our analysis of the cophylogenetic relationship between penguin lice, without excluding any parasite taxa a priori, shows that multi-host parasites can contribute either to the extent of association by descent or by association depending on the reason(s) for their multi-host parasitism (Banks & Paterson, 2005). We think it unlikely the multi-host penguin lice are cryptic species. We examined 10 multi-host lice and nine of the 10 within-species comparisons showed no differences in the 12S and COI sequences (data not shown). For example, Australian and New Zealand blue penguins, differ by 4% for COI and 2% for 12S (Banks et al., 2002) and yet the sequences for the louse A. waterstoni collected from these hosts in Australia and New Zealand did not differ at all for the same gene regions. The penguin lice contrast with several studies that have found that there are genetic differences between populations of louse species parasitising different host species. For example, there were differences in the sequences for COI from populations of the louse Physconelloides eurysema parasitising the pigeon hosts Claravis pretiosa, Columbina inca and C. passerina. COI sequences for P. eurysema even varied with the location of the host (Johnson et al., 2002). Failure to speciate is an alternative reason for multihost parasites that supports association by descent. Penguin species with multi-host lice share several characteristics, such as sympatric distributions and morphological similarity that are thought to make failure to speciate possible (Clayton et al., 2004). For example, all six of the morphologically similar Eudyptes species are parasitised by, and all six eudyptids have been reported from the Snares Islands (Miskelly et al., 2001). Straggling by birds, especially during moulting, may provide sufficient opportunities for the lice to maintain genetic contact. The absence of genetic differences found in our comparison of multi-host louse populations meant we could not distinguish failure to speciate from an extremely recent host switch. It may be that the absence of genetic differences between louse populations on different host species is due to a very recent host switch and insufficient time has elapsed since the switch for differences to accumulate between the louse populations. Hugot et al. (2001) suggested that where the same parasite species parasitises closely related hosts it is more parsimonious to propose failure to speciate than host switching. Application of this principle would suggest that the distribution of most of the multi-host penguin lice is due to failure to speciate. Additionally, we can think of no recent changes in the distribution of penguins that would enable so many very recent host switches. However, faster diverging genes are necessary to distinguish recent host switches from failure to speciate events. Mapping the louse phylogeny onto the penguin phylogeny using TreeMap 2.02b (Charleston & Page, 2002) did not find evidence of extensive cospeciation. There were three or four cospeciation events, depending on the number of host switches allowed, which was not significantly more than would be expected if we compared the penguin phylogeny to 1000 random louse phylogenies. Cospeciation events were only 21 29% of the total number of speciation events, which contrasts with the strong evidence of codivergence found from gophers and their chewing lice (Hafner & Nadler, 1988; Hafner et al., 1994), or seabirds and their chewing lice (Paterson et al., 2000). The proportion of cospeciation events in the penguin-chewing louse assemblage was similar to the lowest values found for chewing lice parasitising a range of mammalian taxa (20% for horses and 25% for cats) (Taylor & Purvis, 2003). Why is the extent of cospeciation in the penguin lice so low in comparison to the pocket gopher lice? Hafner et al. (2003) suggested that cospeciation is likely to occur when the hosts have a patchy distribution, the lice have low dispersal abilities and there are constraints that prevent lice establishing on new host taxa. These factors are likely to affect both the penguin and the gopher lice. However, pocket gophers differ from penguins in that gophers have a fossorial lifestyle, are geographically isolated and have small population sizes (Hafner et al., 2003). It seems
9 164 J. C. BANKS ET AL. likely these host specific factors contribute to the extent of cospeciation between gophers and their lice. Often several species of penguins use the same islands for breeding and/or straggle to other species breeding areas (Marchant & Higgins, 1990). Contact at these times may offer lice opportunities to transfer between hosts and thus reduce the extent of cospeciation. Other cophylogenetic studies of lice parasitising bird species sharing nest holes have also found few cospeciation events. For example, toucan lice do not show cospeciation with their hosts (Weckstein, 2004). Although the pocket gopher chewing lice may be a text-book example of cladogenesis by cospeciation, speciation of other parasite groups, such as the penguin chewing lice, is not as tightly linked to the divergence of their hosts. The TreeMap 2.02b analysis suggested duplications were the predominant method by which penguin lice speciated. Duplication events require genetic divergence between louse populations parasitising the same host species. Band-tailed pigeons, Patagioenas fasciata are parasitised by the louse Physcollenoides spenceri in North and South America (Price et al., 2003). Physcollenoides spenceri in North America show mitochondrial divergence levels of 9% from those in South America, which is similar to divergence levels found between different louse species (Johnson & Clayton, 2004). It is possible that these two louse populations are too divergent to interbreed if they were to be re-united in the future. Overlap of the ranges of P. spenceri in the future would give rise to a duplication event. Penguin colonies tend to be geographically distant from each other, especially in the Southern Ocean and geographical isolation followed by re-contact may be a route for duplications to occur. Warham (1975) speculated that glaciation of breeding grounds during colder conditions or shifts in the hydrological convergences may have resulted in penguin populations and species being more isolated than they are now. However, little is known about the effect of climate variation on the historical distribution of penguins and conditions on the subantarctic island breeding grounds of the crested penguins (Warham, 1975). The TreeMap 2.02b scenario with zero host switches postulated that the most recent common ancestor of the penguins was parasitised by six louse species. Six species of lice parasitising a single host taxon seems unlikely. Currently the maximum number of louse species parasitising a single penguin species, the rockhopper penguin, E. chrysocome, is five (Price et al., 2003). However, two of the three subspecies of rockhopper penguins recognized on morphological differences are parasitised by three louse species, while the third subspecies harbours only two louse species (Price et al., 2003; Banks & Paterson, 2004). The scenario with one host switch (Figs 4 and 5) postulated three louse species on the most recent common ancestor of the penguins. This situation seems more likely. However, without information on the probability of the occurrence of host switches, duplications and extinctions, it is difficult to justify the choice of TreeMap scenarios. Genetic data for the penguins would be useful in choosing between scenarios as relative branch lengths could be used to eliminate switches between hosts that were not extant with the lice. The topology of the 50% majority rule consensus tree produced by the Bayesian analysis with A. bicornutus pruned from the phylogeny was identical to the topology of a tree estimated from a maximum likelihood analysis of the genetic data alone (result not shown). Also, A. bicornutus and were sister taxa in both the combined Bayesian analysis in this study and a maximum parsimony (MP) analysis of only morphological characters (Banks & Paterson, 2004). The Bayesian consensus tree presented here and a MP analysis of the morphological data (Banks & Paterson, 2004) found broadly similar groups. For example, the concii clade had the same members in the MP and Bayesian analyses. However, MP and Bayesian analyses found different relationships within the groups. For example, the Bayesian analysis found to be the basal member of the group whereas the morphological data alone had as the most derived member of the group. Generally cophylogenetic studies have concentrated on host specific parasites while multi-host parasites have been ignored. Indeed, several methods of analysing cophylogenetic data at this point in time cannot deal with multi-host parasites (Charleston & Page, 2002) or else multi-host parasites make the methods unwieldy (Johnson et al., 2001). Multi-host parasites have often been (1) treated as an unresolved clade, which more data will resolve (Page, 1994a), (2) eliminated from the analysis (Huelsenbeck et al., 1997,2000) or (3) assumed to be widespread because of recent host switching (Dowling et al., 2003). None of these methods of analysing cophylogenetic history explicitly allow failure to speciate to occur and yet, if closely related host species share a parasite species, it seems possible that the association has been inherited from an ancestor. Additionally, the method chosen to deal with multi-host parasites markedly affects the extent of association by descent. If every association between multi-host lice and their penguin hosts is due to failure to speciate, all of the penguin louse associations can be explained as association by descent. Alternatively if multi-host lice have only recently colonized their present hosts only 51% of the penguin louse associations are due to association by descent. More study is required to distinguish these possibilities. Because we did not exclude any louse species from our coevolutionary analysis, we found that multi-host parasites, usually neglected in other studies, can critically affect the interpretation of the extent of cophylogenetic history in a lineage. Additionally, some of the methods used to analyse cophylogenetic history require that the phylogenies be manipulated to produce the correct
10 Penguin-louse coevolution 165 reconstruction. Identifying the reasons some parasite species appear to be able to maintain genetic contact despite their hosts diverging will be an interesting extension of cophylogenetic studies. Acknowledgments This work was supported logistically by Antarctica New Zealand, collecting was partially funded by Royal Society of New Zealand, Kelly Tarlton s Antarctica, and Claude McCarthy, Lincoln University doctoral and Gordon Williams scholarships. Kevin Johnson provided assistance with the analysis. We are grateful to many people who assisted with the collection of specimens especially Sonja van Alphen, Amy van Buren, Kerri-Anne Edge, Michel Gaulthier-Clerc, Paul Ponganis, Paul Sagar, Cor Vink and Kath Walker. References Banks, J.C., Mitchell, A.D., Waas, J.R. & Paterson, A.M An unexpected pattern of molecular divergence within the blue penguin (Eudyptula minor) complex. 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At the American Music Awards Sunday night, supermodel Gigi Hadid used her opening monologue to obnoxiously mock the incoming First Lady, Melania Trump. With exaggerated pouted lips, squinted eyes and a fake accent, Hadid mockingly spoke lines such as "I love my husband, President Barack Obama and our children Sasha and Malia." Clearly, the intended comic jab was a reference to Melania's Republican National Convention speech that lifted material from a speech given by First Lady Michelle Obama in 2008.
According to Hadid, she has been working on her debut Melania impersonation all week. "I gotta' get her face right," Hadid said before doing the impression. However, Hadid wasn't the only liberal elite mocking the Trumps that night either.
SNL host Jay Pharaoh took shots at President-Elect Donald Trump as well. "I love Bruno Mars. I don't know what color he is, so I can't deport him," Pharoah said while impersonating Trump. "He looks like Mowgli from 'The Jungle Book,' he does. I love 'Uptown Funk,' it was all totally about me. It was originally called 'Uptown Trump.' I'm going to funk this country up bigly."
It wasn't long until people on social media began ridiculing the AMAs for mocking the Trump family. "Started 2 watch the American Music Awards, as soon as they showed how Un-American they R & bashed President Trump, Boycott #NoRespectNoSupport," Twitter user @Yolanda111 tweeted.
Video of the mocking speeches can be viewed below:
Our #AMAs hosts @GiGiHadid and @JayPharoah are holding nothing back tonight. Let's get this party started! pic.twitter.com/w50MYd7nvk
— AMAs (@AMAs) November 21, 2016
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North Dakota Pipeline Protests Escalating, Hundreds Injured

subset: commoncrawl


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|Message view
|« Date » · « Thread »
|« Date » · « Thread »
|"Rich Wagner" <richmwag...@hotmail.com>
|Re: Ant in JDEE (was "Embedding Ant...")
|Fri, 18 Jul 2003 19:45:01 GMT
Hi Erik, I was wondering if you could be more specific about where to look for details about how Ant was integrated into JDEE, e.g. "get JDEE's source and look at x.java", or "look at such-and-such page in the documentation". I'm very interested in seeing what was done (at the code level), but didn't find what I needed when I poked around a bit. So I'd appreciate a little help, if possible, before I chase too many wild geese... ;-) TIA, Rich Wagner Senior Engineer D3 Advantage / Digital Document Delivery Pitney Bowes, Inc. >From: Erik Husby Reply-To: "Ant Users List" To: Ant Users List Subject: Re: >Embedding Ant... Date: Fri, 18 Jul 2003 14:58:33 -0400 > >Check out what was done to integrate Ant into the JDEE, the Java >Development Environment for Emacs. Get JDEE from http://jdee.sunsite.dk/. >The JDEE allows one to run an "Ant Server" process under Emacs thus >eliminating the Ant startup time making builds much faster. > >-- >Erik Husby Team Lead for Software Quality Automation Whitehead >Institute/MIT Center for Genome Research Rm. 2192 320 Charles St Cambridge, >MA 02141-2023 mobile: 781.354.6669 office: 617.258.9227 >email@example.com _________________________________________________________________ MSN 8 with e-mail virus protection service: 2 months FREE* http://join.msn.com/?page=features/virus --------------------------------------------------------------------- To unsubscribe, e-mail: firstname.lastname@example.com For additional commands, e-mail: email@example.com
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Boss Tuners & Metronomes
Results 1 - 9 of 9 matches
Price DropBoss DB-90 Dr. Beat Metronome
Practice in style with the flagship of the Dr. Beat Metronome line. The new DB-90 is loaded with quality...$135.75Compare Compare Now site1prod213017
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9 patterns and 24 beat variations--plus combos and styles and tap tempo. Headphone jack.$39.00Compare Compare Now site1prod213016
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With so many tuners and metronomes to choose from, you might be wondering what's right for you. Honestly, that will depend on your own needs and preferences. For example, if you want a tuner that clips right on to your guitar, go with the TU-10 chromatic tuner. Boasting a stylish design, along with many features derived from other Boss tuners like Accu-Pitch and even a Stream mode, the TU-10 Clip-On tuner will make you sound flawless night after night.
Of course, we couldn't talk about Boss tuners without mentioning the legendary TU-3. A new and improved version of the TU-2 (one of the top-selling stage tuners ever), the TU-3 has a smooth 21-segment LED meter with a High-Brightness mode that can be seen on the darkest stages. From 7-string guitars to 6-string basses, the TU-3 will have you in tune quickly and accurately.
Now if it's a metronome that you're after, check out the DB-90 Dr. Beat. Loaded with quality sounds, varied drum patterns, and learning tools, this metronome is ideal for any serious percussionist. In fact, the DB-90 has a note mixing function that lets you create a variety of beats instantly by adjusting the levels of 5 different note values. Combined with an onboard microphone, input for your instrument and MIDI and much more, the DB-90 is just what you need to bring your percussion skills to the next level.
And Boss's tuner and metronome choices don't stop there. You'll even find models that contain both tuner and metronome capabilities, so be sure to explore the entire selection. Whatever you're looking for when it comes to tuners and metronomes, you'll have no problem finding it courtesy of Boss.


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Kegs To Go
Hello Beer Lover and Thanks for Stopping By!
To reserve a keg with us, please follow the simple steps below. Please note keg reservations are not guaranteed unless they have been submitted 5 days in advance.
Step 1: Review available Beers and Keg sizing chart to determine your needs (Beer, Keg Size, Quantity). Keg prices include tax.
|Keg Size
|12 oz
|1/2 Barrel
|1/4 Barrel
|1/6 Barrel
Step 2: Make your Reservation
Please fill out the form below to place your order. While this is our preferred method (wink-wink, nudge-nudge), we also accept reservations via email at email@example.com and via phone at 206-420-2407, option 2.
*Required Information
Step 3: Pick up & Payment
Urban Beer Garden (1050 N 34th St, Seattle, 98103)
- Pick up your reserved keg on the date you indicated in our Urban Beer Garden between 11am-9pm at the bar counter.
- As we are often busy in the evenings, keg pick up is best before 4pm.
Please be sure to bring the following with you:
- Driver's License
- Credit Card
- Form of Payment
Payment & Deposit Policy
At the time of pick up, you will be charged for the price of the keg. We do not charge for a deposit for the kegs or taps, but we do retain your credit card information. All kegs with taps must be returned within 14 days of pickup. All kegs destined for a kegerator must be returned within 60 days of pickup. After the allotted time for the deposit period has elapsed ,your credit card will be charged $150 per keg and $50 per tap. This charge will be refunded when the equipment is returned in working order.
Step 4: Return
Return keg(s) to the bar counter at the Urban Beer Garden between 11am and 9pm. If we have charged your card due to a late return, we will refund you at this time assuming all items are returned in proper working condition.
We'd love to fill it! Bring us your clean, sanitized, and pressurized ball-lock or pin-lock Cornelius keg, and we'll fill it with any beer listed in the chart. Please see our drop off / pick up schedule below.
|Drop Off
|Pick UP
|Monday – Wednesday
|Friday after 3pm
|Friday – Sunday
|Tuesday after 3pm


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On this week's Daily Shoah we take the ISIS bucket challenge and talk about James Foley, ISIS, Social Justice Jihadi rappers and generally removing kebab.
Show Prep for the Wicked:
The story of the terrorist rapper and his poor oppressed mother.
BE SURE TO TUNE IN NEXT MONDAY for our Special Labour Day CommieTacular!


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UK-based consumer electronics retailer Expansys has revealed that it will soon be stocking up on and selling the champagne gold edition of the HTC One for £480.00 or about US$ 795.00 apiece. The handset will also be SIM-free and with no monthly contract.
The information was made available through Expansys's product page for the HTC One (Gold Edition).
This edition, however, is not the same as the limited edition plated with 18-carat gold that was specially made for the 18th MOBO music awards in October last year. HTC, in partnership with Goldgenie, commissioned the production of only 5 real-gold edition HTC One handsets.
The edition arriving in Expansys, instead, will be the one in aluminum casing with a champagne gold hue. The golden shade is beautifully complemented by dark-gray trimming running along its sides.
Despite the golden exterior of the gold HTC One, however, it uses essentially the same hardware as the classic silver edition: quad-core 1.7-GHz Snapdragon 600 processor, 2 gigabytes of RAM, 1080p Full HD display, high-end camera sensor, Android 4.3 Jelly Bean, and HTC Sense 5.5.
HTC is due to release a new flagship Android phone before the first quarter ends. The company is known for manufacturing Android phones with premium look and feel.
In the UK region, Carphone Warehouse and Phones 4u were the first to sell the HTC One Gold Edition but as on-contract items.
Amazon offers the HTC One Gold Edition at a much lower price.
The HTC One is also available in blue, red, silver, and black.


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2004 Jeep Wrangler Top Comparisons
Users ranked 2004 Jeep Wrangler against other cars which they drove/owned. Each ranking was based on 9 categories. Here is the summary of top rankings.
User Reviews
View reviews where users commented on categories.Clear selections
Displaying 21 - 30 of 48 2004 Jeep Wrangler reviews.
CarGurus https://www.cargurus.com
2004 Jeep Wrangler RubiconReview
Great Way To Get Into Wheeling —
When you buy a Jeep you buy into a lifestyle. The overall onroad performance has much to improve but that isnt the reason why you purchase a Rubicon. The engine has lots of torque and is loud at any speed above zero. The tires are loud at speeds in excess of 80km/h and give off a vibration that you can feel through the jeep. You will not win any races at the stop light with this guy and you will use up the hole hwy on ramp almost every time. The offroad performance is where it counts with this jeep. With low range gearing set at 4:1 and air actuated locking front and rear differentials, you will find it hard to get stuck. This thing crawls over obstacles and powers through the mud. But when you do find yourself in a situation where you are stuck, you will be happy that you signed up to the jeep lifestyle because every jeep owner on the trail would be their to help you out. The appearance is stylish and aggressivie with the rubicon rims and mud terrain tires. As you get into the jeep lifestyle it will be hard not to upgrade you vehicle to improve on its off road abilities. you will find no shortage of parts to help make this vehicle trully unique to you. The cost of this vehicle is relatively high because of the desire of the Rubicon model. But if you are using it for off roading it is worth it. You will get the toys with it that will help you in the sticky situations. The fun factor is really high on this one. If you are just cruising down the road with the top down or rippin up the trails this Jeep fits in everywhere. You will have a blast. Just one thing to remember when you start getting into offroading. Jeep stands for "Just Empty Every Pocket"
Primary Use: Sport/fun (spirited driving, track racing, off-roading, etc.)
Pros: Amazing fun
Cons: If your not going to use it for offroading then dont get a Rubicon
Looking for a Used Wrangler in your area?
CarGurus has 15,877 nationwide Wrangler listings starting at $3,495.
CarGurus https://www.cargurus.com
2004 Jeep Wrangler RubiconReview
Its Just The Best. — The best, go anywhere, do anything, cruising convertible you will ever own. Wanna make the guy in his Escalade Jealous... buy a Jeep. You will not regret it. Its NOT practical, its NOT good on gas, but MAN does it ever make you feel good.
Pros: Top down, doors off, can't beat it.
Cons: Tires are $600 each. You have been warned.
CarGurus https://www.cargurus.com
2004 Jeep Wrangler RubiconReview
My Rubicon — Great gas mileage, solid 4.0L I6, 5spd manual, 4wd with pnuematic lockers for both axles, Unusually low mileage for a wrangler with only 33000 miles on it, lots of customization options and of course summers with the top and doors off.
Primary Use: Commuting to work
Pros: Everything
Cons: nothing
CarGurus https://www.cargurus.com
2004 Jeep Wrangler XReview
It Can Handle Anything You Throw At It — Built it myself and the engine is being worked on for more horsepower currently. The only other thing being done is new wheels and tires and lights. It is a very fun car to drive and I love it to death.
Primary Use: Sport/fun (spirited driving, track racing, off-roading, etc.)
Pros: everything
Cons: gas
CarGurus https://www.cargurus.com
2004 Jeep Wrangler RubiconReview
We Enjoy Having It... — We've had the Rubicon since 2005 and we love it! It isnt the most fuel efficient vehicle because of all of the stuff we've added to it...Winch, high-lift jack, bigger tires, lift kit, light bar....but it's fun to drive and will be super useful this winter in the snow
Primary Use: Sport/fun (spirited driving, track racing, off-roading, etc.)
Pros: Fun, reliable
Cons: Not great on gas
CarGurus https://www.cargurus.com
2004 Jeep Wrangler RubiconReview
Me Likes A Lot, I Can Go Places Other People Have To Walk To Get To — performance, ... none unless you count going up 45% inclines or passing 3 feet of muddy water build quality, ... so/so paint job sucks, rust appearance, ... charm of a brick wall fuel economy , ... no not really fun, .... hell yeah, its a lot of fun
Primary Use: Sport/fun (spirited driving, track racing, off-roading, etc.)
Pros: offroad monster
Cons: no comfort
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2004 Jeep Wrangler XReview
Its A Jeep Thing — 4.0 i6 nothing else better. you can do what ever you want to the engine and it will still run, by far the best engine ever made. So dont buy a new jeep cause there V6s bc chrysler is fucing dumb
Primary Use: Sport/fun (spirited driving, track racing, off-roading, etc.)
Pros: the engine
Cons: nothing
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2004 Jeep Wrangler SEReview
Love My Jeep — A Wranglerr is the best Florida vehicle. I love being to take the top off and ride but when it comes to interstate power, there is none. But hey, who's in a hurry when your riding a Jeep anyway. For a weekend vehicle a Jeep is ideal. As for me, I enjoy riding with the top down on my way to work. Kinda clears my mind.
Primary Use: Commuting to work
Pros: Fun in the sun
Cons: No power
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2004 Jeep Wrangler SportReview
It's Bulletproof — It's a amazing handling vehicle that is built strong. The Jeep look great inside and out. The gas mileage isn't the greatest but if you take it easy, you can get 425 km+ to a tank. The Jeep is so much fun to drive, I never get tired of driving it!!
Primary Use: Commuting to work
Pros: I love the look of them
Cons: hard on gas
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2004 Jeep Wrangler SEReview
Displaying 21 - 30 of 48 2004 Jeep Wrangler reviews.
Reviews From Other Years
2015 Jeep Wrangler Reviews
Wife Loves This Thing
Love to haul the kayak and bikes. take the roof and doors off and hit the town or back woods. Everyone loves it. Wish it came with a back up sensor. The dark saddle trim is perfect with the black c... Read More
2014 Jeep Wrangler Reviews
Great Fun Car
Love the car, glad I had the opportunity to drive one. Just a real feel good car :-) Up high and easy to drive, it can snow, rain, sun shine, the car is ready and so the driver. The top can be down in... Read More
Smooth Ride
Black paint Doors 2 Soft top convertible Passenger seats 4 City economy17 (Est) mpg Highway economy21 (Est) mpg I am amazed at how solid and smooth the ride is- This vehicle is easy to drive, has a t... Read More
2013 Jeep Wrangler Reviews
I love everything about this car. I'm still in the process of building it, but it'll be about another 10 grand before I'm close to the finished look. In the mean time it'll be garaged while I save mon... Read More
Raise it up. Put bigger grippy tires on it. Add a wench and some awesome lights. Outfit it with the images you want. Smile when the guys on the trail single you out for the best looking and functi... Read More
2012 Jeep Wrangler Reviews
2012 Wrangler Sport
One of my favorite vehicles. There are few things like driving with the top and doors off of a quick vehicle with a manual transmission. How driving should be. Sure, it's basically a two-seater and ha... Read More
Fun To Drive
Overall I love driving this car especially in the summer with top and doors off. Handles great in the snow in the winter as well. Only thing for me is I would like to get the 4 door unlimited as carg... Read More
2011 Jeep Wrangler Reviews
Go Anywhere Escape Plan.
I wish there was a few more inches behind the front seats. The overall vehicle has improved greatly over the previous body style but the 3.8 is the worst motor ever. The ride and handling is amazing... Read More
It's A Jeep
Compared to the older jeeps, the 2011 is much smoother and has more power and speed. The 2011 allows you to set your speed at 65 on highways the jeep is able to maintain the speed without driving up t... Read More
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