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The family Polypteridae, commonly known as "bichirs", is a lineage that diverged early in the evolutionary history of Actinopterygii (ray-finned fish), but has been the subject of far less evolutionary study than other members of that clade. Uncovering patterns of morphological change within Polypteridae provides an important opportunity to evaluate if the mechanisms underlying morphological evolution are shared among actinoptyerygians, and in fact, perhaps the entire osteichthyan (bony fish and tetrapods) tree of life. However, the greatest impediment to elucidating these patterns is the lack of a well-resolved, highly-supported phylogenetic tree of Polypteridae. In fact, the interrelationships of polypterid species have never been subject to molecular phylogenetic analysis. Here, we infer the first molecular phylogeny of bichirs, including all 12 recognized species and multiple subspecies using Bayesian analyses of 16S and cyt-b mtDNA. We use this mitochondrial phylogeny, ancestral state reconstruction, and geometric morphometrics to test whether patterns of morphological evolution, including the evolution of body elongation, pelvic fin reduction, and craniofacial morphology, are shared throughout the osteichthyan tree of life.
Our molecular phylogeny reveals 1) a basal divergence between Erpetoichthys and Polypterus, 2) polyphyly of P. endlicheri and P. palmas, and thus 3) the current taxonomy of Polypteridae masks its underlying genetic diversity. Ancestral state reconstructions suggest that pelvic fins were lost independently in Erpetoichthys, and unambiguously estimate multiple independent derivations of body elongation and shortening. Our mitochondrial phylogeny suggested species that have lower jaw protrusion and up-righted orbit are closely related to each other, indicating a single transformation of craniofacial morphology.
The mitochondrial phylogeny of polypterid fish provides a strongly-supported phylogenetic framework for future comparative evolutionary, physiological, ecological, and genetic analyses. Indeed, ancestral reconstruction and geometric morphometric analyses revealed that the patterns of morphological evolution in Polypteridae are similar to those seen in other osteichthyans, thus implying the underlying genetic and developmental mechanisms responsible for those patterns were established early in the evolutionary history of Osteichthyes. We propose developmental and genetic mechanisms to be tested under the light of this new phylogenetic framework.
Osteichthyans (bony fish and tetrapods; Fig. 1) have evolved remarkably diverse body plans since their initial radiation in the Late Silurian ~420 Mya [1,2]. It is therefore not surprising that most major extant lineages have been the subject of extensive evolutionary biology research. As a result, we know much about the evolutionary history and patterns of morphological evolution in osteichthyans, most notably teleost fish and tetrapods (amphibians, reptiles, and mammals). In the age of genomics and advanced molecular techniques, knowledge of these relationships and patterns has proven useful in uncovering the developmental and genetic mechanisms responsible for morphological diversity [e.g., [3-9]].
Figure 1. Consensus view of the phylogeny of extant Osteichthyes, including the position of bichirs (Polypteridae), inferred from phylogenetic analyses of molecular and morphological data.
However, the same cannot be said for one lineage that diverged early in the evolutionary history of Actinopterygii (ray-finned fishes) - the family Polypteridae. Commonly known as "bichirs", this clade includes 12 extant described species (as well as multiple subspecies) that inhabit freshwater rivers and lakes of tropical Africa [10]. Little is known about polypterid evolution primarily because of their long history of phylogenetic and taxonomic confusion, likely owing to their unique collection of "primitive" (e.g., ganoid scales, cartilaginous skeleton, the intestine with a spiral valve) and derived (e.g., highly modified dorsal fins, pectoral fins with lobed base covered with scales, possession of only four gill arches) anatomical features [11-13]. However, recent morphological and molecular analyses have determined that bichirs are a basal lineage of Actinopterygii (ray-finned fishes; Fig. 1) [14-19].
Bichirs provide an attractive system to test whether patterns of morphological evolution are shared throughout the osteichthyan tree of life. If common patterns exist, it would suggest that the underlying genetic and developmental mechanisms responsible for those patterns were established early in the evolutionary history of Osteichthyes. For example, there exist multiple independent derivations of body elongation in modern Actinopterygian fish [e.g., [20-23]], amphibians [e.g., [24]], squamate reptiles [e.g., [25-27]], and numerous extinct tetrapod lineages [e.g., [28,29]]. The loss of pelvic fins and limbs has occurred multiple times over the course of osteichthyan evolution [22,27,30-32]. Furthermore, diverse changes in craniofacial morphology have been the subject of particularly extensive research [5,33-41].
However, the greatest impediment to elucidating the patterns of morphological change in Polypteridae is the lack of a well-resolved, highly-supported phylogenetic tree. In fact, the interrelationships of polypterid species have never been subject to molecular phylogenetic analysis. Those molecular studies that have included polypterid representatives did so in the context of determining "deep" osteichthyan and actinopterygian relationships (Fig. 1) or surveying Hox gene clusters [42].
Extant polypterids comprise two extant genera, Erpetoichthys and Polypterus. Boulenger [43] distinguished these two genera based on the extremely elongate body and absent pelvic fins in Erpetoichthys, and split Polypterus into two taxonomic groups based on the position of the mandible relative to the snout ("lower jaw protrusion" hereafter). Poll [44-46] defined five clusters of species and an ancestral species within extant polypterids based on anatomical features such as relative jaw length, the location and size of the eyes, width of the suboperculum, and proportion of the gular plate. An analysis of 15 anatomical characters suggested, among other things, a sister relationship between Erpetoichthys and P. weeksi, thus rendering Polypterus polyphyletic [47]. However, this study did not incorporate objective rooting or optimality criteria and subsequent parsimony reanalysis revealed that this phylogeny is largely unresolved [12]. These conflicting and inconclusive results warrant additional phylogenetic scrutiny using molecular data.
Here, we infer the first molecular phylogeny of Polypteridae using all 12 recognized species including all but three recognized subspecies to achieve three major objectives. First, we evaluate whether the current species taxonomy underestimates the underlying phylogenetic diversity. Secondly, we examine patterns of evolution in body elongation pelvic fin reduction, and craniofacial morphological evolution in this new phylogenetic context. Finally, we recommend developmental and genetic mechanisms to be tested in the future in conjuction with this new phylogenetic framework.
Phylogenetic analyses
We obtained fresh tissues from commercial dealers of 15 recognized species and subspecies of polypterids including Erpetoichthys calabaricus, Polypterus ansorgii, P. bichir bichir, P. b. lapradei, P. delhezi, P. endlicheri congicus, P. e. endlicheri, P. mokelembembe, P. ornatipinnis. P. palmas buettikoferi, P. p. polli, P. retropinnis, P. senegalus senegalus, P. teugelsi and P. weeksii (Table 1). One to four specimens of each taxon were included in the phylogenetic analyses. Although we included all recognized species of polypterids, we were unable to obtain tissue samples for three subspecies, P. bichir katangae, P. palmas palmas, and P. senegalus meridionalis (Table 1). We obtained DNA sequences of outgroup taxa (Sturgeon [Acipenser transmontanus], Bowfin [Amia calva], Coelacanth [Latimeria chalumnae], Spotted gar [Lepisosteus oculatus], Paddlefish [Polyodon spatula], and Lungfish [Protopterus dolloi]) from GenBank (Additional file 1).
Additional file 1. Information of DNA sequences analyzed in this study.
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Table 1. Number of vertebrae and condition of the lower jaw and pelvic fins in each species and subspecies of Polypteridae.
DNA samples were extracted following the same procedure used in Tokita et al. [48]. A part of the mitochondrial 16S rRNA and cytochrome b (cyt b) genes were amplified by PCR System GeneAmp 2700 (Applied Biosystems, Lincoln, USA) using an Ex Taq polymerase kit (Takara Shuzo Co., Ltd., Otsu, Japan) and the primers provided in Additional file 2. After initial denaturing for 5 min at 95°C, 25-30 cycles were performed with denaturing 1 min at 94°C, annealing 3 min at 50-60°C, and primer extension for 3 min at 72°C, followed by a final elongation of 7 min at 72°C. PCR products were purified by PEG/NaCl precipitation and then sequenced using a Big Dye Terminator Cycle Sequencing Ready Reactions Kit v1.0 and an ABI PRISM 377 and 3130 DNA Sequencer (Applied Biosystems, Lincoln, USA), using the primers provided in Additional file 2.
Additional file 2. List of primers used in this study.
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Because of conserved codon reading frames, the cyt b protein coding sequences could be unambiguously aligned by eye. The 16S rRNA was also aligned by eye, but nucleotide positions that could not be unambiguously aligned were excluded from phylogenetic analysis (data set available from the authors). In some regions of the 16S data, the ability to align the polypterid taxa was improved if we removed sequences for the outgroup taxa (and their corresponding data replaced with "?" in these regions). Because our primary goal is to test the interrelationships of polypterids, and not necessarily their relationship to other osteichthyans, we feel the exclusion of these data is justified. All DNA sequences were deposited in GenBank (Additional file 1). The final data set for subsequent phylogenetic analysis included 925 base pairs (bp) of 16S (256 parsimony informative characters) and 932 bp of cyt b (103, 29, and 292 parsimony informative characters for the first, second, and third codon positions), for a total of 1857 total characters.
We employed partitioned Bayesian analyses to infer the phylogenetic relationships among polypterids. Previous studies have demonstrated that applying separate models of nucleotide evolution to specific subsets of nucleotide data (i.e., "partitioned" or "mixed-model" analyses) improves phylogenetic reconstruction [49-51]. We therefore partitioned the data a priori into four total partitions, one for the 16S rRNA and three for each codon position of cyt b. We determined the appropriate models of sequence evolution from a pool of 16 models (K80, HKY, SYM, GTR with and without I and Γ) for each partition using a JC corrected neighbor-joining tree in PAUP* [52] and the Bayesian Information Criterion [BIC: [53]]. We subsequently conducted Bayesian phylogenetic analyses using the most appropriate models (GTR+I+Γ, GTR+I+Γ, HKY+I+Γ, and GTR+Γ for the 16S and three cyt b codon positions, respectively) in the parallel version of MrBayes v3.1.1 [54,55]. Each Bayesian analysis consisted of 107 generations, using four chains sampled every 1000 generations and default priors (substitution rates, Dirichlet [1, 1, 1, 1, 1, 1]; base frequencies, Dirichlet [1, 1, 1, 1]; gamma shape parameter, uniform [0, 200]; topologies, uniform; branch lengths, unconstrained exponential [λ = 10]).
To determine convergence of the Bayesian analyses, we constructed cumulative posterior probability plots for each clade using the cumulative function in AWTY [56]. Stationarity was assumed when the cumulative posterior probabilities of all clades stabilized. These plots indicated that excluding the first two million generations as burn-in was sufficient. To decrease the chance of reaching apparent stationarity on local optima, four separate analyses were performed. Posterior probability estimates for each clade were then compared between the four analyses using a scatter-plot created by the compare command in AWTY. Posterior probability estimates for clades were similar in all four analyses, and the results of the analyses were combined and the frequency of inferred relationships in these 32,000 trees represented estimated posterior probabilities of clades. Posterior probabilities ≥ 0.95 are considered statistically significant (i.e., "strong") clade support [57].
Morphometric analysis
To investigate shape differences in the head region among Polypterus species, we employed landmark-based geometric morphometrics [58,59]. This approach allows us to describe morphological transformation of certain organismic structures visually and compare morphological differences among taxa quantitatively [59] and has been widely used in contemporary evolutionary biology research [e.g., [34,40,41]]. A total of 13 taxa and 26 specimens caught in wild were included in this analysis. Photographs of both dorsal and lateral views were taken with a digital camera. The photo images of the specimens were imported into the tpsDig2 software [60]. Eight and ten landmarks were placed for dorsal and lateral views, respectively (Additional file 3). The generalized least squares Procrustes superimposition, which removes information unrelated to shape such as position, scale, and orientation, was carried out using Coordgen 6n [61]. Based on the coordinates after the Procrustes superimposition, the partial warp scores were computed and principal component analysis (PCA) was conducted by calculating PCs based on the covariance matrix derived from the partial warp scores using PCAgen6 h [62].
Additional file 3. Landmark definition for morphometric analysis. (A) Polypterus palmas buettikoferi in dorsal view, (B) P. endlicheri congicus in dorsal view, (C) P. p. buettikoferi in lateral view, (D) P. e. congicus in lateral view.
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Ancestral state reconstruction
To evaluate the evolution of body elongation and pelvic fin loss within polypterids, we reconstructed ancestral states using data from extant sampled taxa, the rooted mtDNA phylogram (using mean branch lengths from the posterior distribution) estimated from our Bayesian phylogenetic analyses (above), and Mesquite v2.6 [63]. We removed the non-polypterid outgroups and pruned the data set and trees to include a single exemplar of each species and subspecies. We collected vertebral number for each species from FishBase [64] and preserved specimens. Polypterid species typically exhibit variation in the number of vertebrae (Table 1). Because there is currently no statistically elegant way to incorporate this information into ancestral state reconstruction analyses (e.g., modeling vertebral number in each species as statistical distributions) we used the mean number of vertebrae and coded it as a continuous character [e.g., [65]]. Ancestral vertebrae number for each node was subsequently estimated using weighted squared-change parsimony [66]. Presence of pelvic fin was coded as discrete binary characters and the marginal probabilities of each state at each node were estimated with maximum likelihood [67,68] using the 1-parameter symmetrical Markov model [69]. Ancestral states with a marginal probability ≥ 0.95 were considered resolved with statistical significance.
Results and Discussion
Polypterid phylogeny and hidden diversity
Posterior probabilities (PP) for a vast majority of nodes are statistically significant (≥ 0.95) and the clades with probabilities less than 0.90 are relationships between populations of subspecies (Fig. 2). The analyses support a basal split between Erpetoichthys and Polypterus. In Polypterus, the recently described P. mokelembembe [69] is the sister taxon to all other members of the genus; although support for this relationship is very high (PP = 0.94), it is not significant. The remaining Polypterus are divided into two major clades, one of which (P. bichir + P. endlicheri + P ansorgii) is characterized by lower jaw protrusion (Table 1; see below).
Figure 2. Molecular phylogeny of the extant polypterid species inferred from partitioned Bayesian analyses 16SrRNA and cyt b mitochondrial genes. Branch lengths are means of the posterior distribution. Numbers above or below the node indicate the Bayesian posterior probability that clade is correctly estimated given the model. Posterior probabilities less than 0.50 are not shown. Colors indicate groups defined in Fig. 4.
These results reinforce the findings of Boulenger [43] who recognized Erpetoichthys and Polypterus as distinct genera, in addition to two groups within Polypterus based on the condition of the lower jaw (P. mokelembembe was not yet described). The phylogenetic position of Erpetoichthys is not particularly surprising considering numerous biological aspects of this species. The skull of E. calabaricus possesses a variety of unique characteristics that are not found in Polypterus [12]. Furthermore, the ecology of E. calabaricus quite different than Polypterus as it inhabits coastal and estuarine areas unlike Polypterus species that primarily inhabit freshwater [10,70-72].
Our data support Boulenger's conclusion of a close relationship between P. weeksi and P. ornatipinnis [43]. However, the present analyses reveal phylogenetic relationships that are substantially different from other previous studies focusing on the detailed interrelationships of the species. Analyses of 15 morphological and anatomical traits [47] identified close relationships among P. ornatipinnis, P. weeksii, and Erpetoichthys calabaricus, and among P. senagalus, P. retropinnis, P. palmas (although subsequent parsimony reanalysis of these data inferred an almost completely unresolved tree [12]). However, neither of these relationships was inferred by our analyses.
Poll [44-46] recognized close relationships among P. ornatipinnis, P. weeksii, and P. delhezi and that between P. senagalus and P. retropinnis based on morphological characteristics (e.g., relative jaw length, the location and size of the eyes). Our results statistically reject these phylogenetic hypotheses (i.e., alternative hypotheses have a posterior probability > 0.95). Our phylogenetic analyses also statistically reject Poll's hypothesis [45] that P. ansorgii is the sister lineage to all other Polypterus instead, this position is represented by P. mokelembembe.
Our phylogenetic analyses also demonstrate that at least two species, as currently described, are not natural groups - P. endlicheri and P. palmas. Polypterus endlicheri is paraphyletic with respect to both P. bichir and P. ansorgii. Taxonomic changes are clearly needed and we recognize P. e. congicus and P. e. endlicheri as distinct species, P. congicus and P. endlicheri. Hanssens et al. [73] recognized three subspecies in P. palmas: P. p. palmas, P. p. polli, P. p. buettikoferi based on morphometrical inferences. In our phylogeny, the sister relationship between P. p. polli and P. delhezi is strongly supported (PP = 1.0) as well as the relationship between P. p. buettikoferi and P. teugelsi (PP = 1.0). Thus, the morphometric resemblance between P. p. buettikoferi and P. p. polli is clearly the outcome of evolutionary convergence. To understand the evolutionary history of P. palmas complex and make subsequent taxonomic changes, it is necessary to include P. p. palmas in future phylogenetic analyses.
These results imply that morphological characteristics such as body proportion and body coloration used in previous studies are not suitable for inferring phylogeny, possibly because states of these characters tend to be flexibly altered depending on the condition of the species' environment. Furthermore, these phylogenetic results not only have implications for morphological evolution within Polypteridae (below), but also reveal the hidden genetic diversity of this ancient clade of enigmatic fish. That at least two species are polyphyletic (P. endlicheri and P. palmas) is evidence that the current taxonomy of Polypterus underestimates its genetic diversity.
Perhaps more importantly, we find that P. mokelembembe, a species only recently described [74], represents the sister lineage to all other members of the genus. This result suggests that much of the evolutionary diversity of extant Polypteridae has yet to be fully discovered.
As both the 16S and cyt b mitochondrial genes are maternally inherited as a single unit, we acknowledge that our phylogeny specifically tracks the evolutionary history of the mitochondrial genome, and not necessarily the species history. However, we note that there are no obvious instances of recent introgression (e.g., [75]) as all of the named taxa are monophyletic. However, we cannot exclude the possibility that incomplete lineage sorting obscures deeper relationships (see [76] for a review), but we also note that this is unlikely given the rapid coalescence of mitochondrial DNA. Furthermore, we note that, although we have not sampled the entire mitochondrial genome for this analysis, the fact that almost every node in our phylogeny is significantly supported (posterior probability > 0.95) is evidence that we have indeed captured almost as much "phylogenetic signal" as is possible from the mitochondrial genome. Nonetheless, for clarity, we refer to the phylogeny as the "mitochondrial phylogeny" to distinguish it from the true (albeit unknowable) species tree, and acknowledge that a more confident species tree of polypterids awaits future analyses of many more independently evolving loci.
Evolution of body elongation
Our ancestral state reconstructions (Fig. 3) estimate numerous increases and decreases of vertebral number, and thus, changes in body elongation. However, the most striking transformation is seen in Erpetoichthys. Erpetoichthys possesses between 110 and 113 vertebrae (Table 1), but the vertebral number reconstruction for the most recent common ancestor (MRCA) of this genus and Polypterus is ~71. It is not known whether the already relatively high number of vertebrae reconstructed for the MRCA is an artifact of the extremely high number of vertebrae in Erpetoichthys biasing the estimation of the root, or evidence that extinct stem polypterids were already relatively elongate. Regardless, Erpetoichthys is much more elongate than the estimated common ancestor of crown polypterids.
Figure 3. Extant states and result of ancestral state reconstructions of vertebral number using squared-change parsimony.
Anguilliformity (eel-like elongated body form), such as that seen in Erpetoichthys, is seen across diverse lineages of vertebrates including ray-finned fishes, caecilian amphibians, and squamate reptiles [21,23-27,77], and an increase in vertebral number is considered as the common primary factor underlying such body form change [22]. Developmentally, vertebrae emerge from 'somites' that form one after another, with antero-posterior direction in the embryo. They bud off from the anterior end of unsegmented 'presomitic mesoderm' and this budding is regulated by a segmentation clock represented by oscillation of 'clock' genes such as Lunatic fringe [reviewed in [78,79]]. Recently, it was found that the tempo of the segmentation clock is accelerated in snake embryos, "ticking" around four times faster relative to growth rate than in shorter-bodied animals like chick and mouse and consequently generating a large number of somites [80].
Erpetoichthys has around 100 preanal vertebrae, which is almost double the number of other polypterid species [[22], pers. obs.]. Although few embryological studies of Erpetoichthys have been conducted [81,82], the yolk-sac larva and apterolarva of this species appear to be much more elongated and have more somites than those of Polypterus. It is plausible that an increase of vertebrate number in Erpetoichthys was brought about by a similar developmental mechanism to that exploited in snake embryogenesis [80,83], although the precise molecular mechanisms that regulate tempo of segmentation clock in vertebrate embryos have not been identified yet. The present study implies that body elongation observed in Erpetoichthys seems to be brought about in an early phase of polypterid evolution, possibly by accelerating the tempo of the segmentation clock twice as fast as that in other polypterids.
Although the extreme degree of body elongation observed in Erpetoichthys is not found in extant Polypterus, two species: P. bichir and P. teugelsi show a relative increase of vertebrate number (Table 1; Fig. 3). Although the total number of vertebrae is over 60 in both species, the mechanisms by which they achieve this body elongation are quite different. In P. bichir, the number of preanal vertebrae is less than 50 like that in majority of Polypterus species but the number of postanal vertebrae increases into over 14 [22]. On the other hand, P. teugelsi shows relative elongation of the trunk region, possessing at least 54 preanal vertebrae (pers. obs.). Instead, this species has only 10 postanal vertebrae keeping the condition in majority of Polypterus species. Thus, body elongation seems to have been achieved independently in two Polypterus species by employing different developmental mechanism that functions in separate developmental modules [22]. We note that this pattern is surprisingly similar to that seen in squamate reptiles (lizards and snakes) in which body elongation is also achieved by two distinct mechanisms elongating either the trunk or the tail [24-26].
Pelvic fin loss
In vertebrates, the body elongation tends to co-occur with a loss or reduction of paired appendages [21,24-26,29,84,85]. Following this evolutionary trend, Erpetoichthys, whose body is extremely elongated, also lacks pelvic fins (Table 1). Maximum likelihood ancestral state reconstructions strongly support the conclusion that pelvic fins were present in the most recent common ancestor of Erpetoichthys and Polypterus (Marginal probability = 0.97; data not shown), and that they were secondarily lost in the Erpetoichthys lineage. Given that the loss of pelvic appendages is frequent in multiple osteichthyan lineages, it is likely that the genetic mechanism responsible for pelvic fin loss in Erpetoichthys is similar to other members of this group. Translocation of Hox gene expression in trunk paraxial and/or lateral plate mesoderm is known to be the potential mechanism underlying limb elimination in vertebrates [[86,87]; but see recent paper by Wolstering et al. [88]]. However, this is a less likely explanation for pelvic fin loss in Erpetoichthys because such alteration of developmental program might also obliterate any vestiges of the pelvic girdle yet Erpetoichthys retains a pair of pelvic girdle elements (pers. obs.). Degeneration of apical ectodermal ridge (AER) and lack of expression of AER-associated genes might account for hindlimb truncation in lizards [89,90] and snakes [86], but this has never been recorded in non-tetrapods. Similarly, failure in establishing the Zone of Polarizing Activity (ZPA) has only been implicated in hindlimb loss in cetaceans [91]. The most promising potential mechanism responsible for pelvic fin loss in Erpetoichthys is the elimination of expression of transcription factor-encoding genes such as Pitx1 and Hoxd9 in the pelvic region - the mechanism that causes pelvic fin loss in some teleost fishes [31,32,92]. Unfortunately, the pattern of pelvic fin reduction in embryogenesis of Erpetoichthys has not been described precisely [82] and thus, discrimination of these various hypotheses is not yet possible. However, future studies of these genetic and developmental mechanisms can do so within the phylogenetic framework constructed here.
Transformation of craniofacial morphology
The first three PCs of the geometric morphometrical analysis of both dorsal and lateral views of the head accounted for over 70% of the total shape variation. In the comparison of the dorsal views, PC1 explains 36% of the total variance and primarily describes positional change of the orbit. PC2 explains 21% of the total variance (Additional file 4). In the comparison of the lateral views, PC1 explains 51% of the total variance and describes positional change of the tip of the mandible, the orbit, and labial end. While, PC2 explaining 14% of the total variance describes positional change of the opercle.
Additional file 4. Thin-plate splines (TPS) of principal component 1 (PC1) and 2 (PC2) for both dorsal and ventral views of the head of Polypterus. Arrows indicate PC value plus positive 0.1 score. (A) PC1 in dorsal view, (B) PC2 in dorsal view, (C) PC1 in lateral view, (D) PC2 in lateral view. Numbers along with each plot indicate landmarks defined in Additional file 3.
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Our morphometric analysis revealed the trend of morphological change in Polypterus species where PC1 positively correlates with the size of the head (Fig. 4a and 4c). Phylogenetically closely-related P. ansorgii, P. bichir and P. endlicheri occupied the most extreme positions on the positive sides of both PC1 and PC2 axes. This suggests that these three species not only tend to have a rostrally-projected lower jaw (a condition already noted by previous authors [43,47]), but also a medially located and up-righted orbit, enlarged mouth, and rostroventral distortion of the opercle. Judging from our mitochondrial phylogeny, the character complex seen in these three species is a derived condition.
Figure 4. Plots of principal component (PC) and centroid size (CS) for morphometric characters of Polypterus. a) CS and PC1 for dorsal view, b) PC1 and PC 2 for dorsal view, c) CS and PC1 for ventral view, d) PC1 and PC2 for ventral view. Grouping corresponds to the clade inferred from molecular phylogenetic analysis (Fig. 2): Blue = Polypterus ansorgei, P. bichir lapradei, P. endlicheri endlicheri, and P. e. congicus; Green = P. mokelembembe; Purple = P. ornatipinnis, P. retropinnis, and P. weeksii; Red = P. delhezi, P. palmas buettikoferi, P. p. polli, P. senegalus senegalus, and P. teugelsi.
Interestingly, P. delhezi displayed a similar tendency of morphological change in PC1 axis of both dorsal and lateral views, indicating that this species actually possesses an intermediate condition where neither the upper or lower jaw show pronounced rostral projection. Because P. delhezi is distantly related to the lower jaw projection clade (P. ansorgii, P. bichir and P. endlicheri), the similar cranial shape seen in two polypterid lineages has apparently evolved independently. In polypterid species whose snout projected more rostrally to the tip of mandible, the range of variation was broader than that in species with lower jaw protrusion (Fig. 4). Furthermore, the range was broadly overlapped between two main clades: the monophyletic group composed of P. ornatipinnis, P. weeksii, and P. retropinnis (purple) and that composed of P. delhezi, P. palmas, P. senegalus, and P. teugelsi (red), showing no lineage-specific pattern of craniofacial transformation.
There are multiple potentially productive avenues of future genetic and developmental research into the mechanisms of craniofacial development in Polypteridae. In vertebrates, morphology of the lower jaw is regulated by cranial neural crest cells, and several transcription factors in BMP and/or FGF signaling pathway appear to be involved in the process [4,38,93-97]. Although we do not know what change in developmental program caused substantial transformation of craniofacial morphology in P. ansorgii, P. bichir and P. endlicheri, the event that occurred in relatively early phase of polypterid evolution might have facilitate the shift into a new ecological niche, affecting their feeding strategy.
The discovery of patterns of morphological evolution among "deep" lineages of organisms permits further exploration of the general underlying genetic and developmental mechanisms that evolved early in evolutionary history. Our new molecular phylogeny of Polypteridae, in conjunction with ancestral state reconstruction and geometric morphometric analyses reveals that many of the patterns of morphological evolution seen in polypterids are shared with other actinopterygian, and indeed, osteichthyan lineages. Thus, this implies the underlying genetic and developmental mechanisms responsible for those general patterns were established early in the evolutionary history of Osteichthyes, or perhaps even older in the craniate tree of life. Moreover, these patterns suggest interesting avenues of future evolution and developmental research that are only possible in a phylogenetic framework.
Authors' contributions
MT designed the study. DS and MT collected data. DS and MT performed the morphometric analyses and MCB performed the phylogenetic analyses. MT wrote the paper with the assistance of DS and MCB. All authors read and approved the final manuscript.
We are grateful to Nori Satoh, Michio Hori, Naomichi Ogihara, Takushi Kishida, Wataru Yano, and Nobuto Kikukawa for technical support. We also thank Hiroshi Senou (Kanagawa Prefectural Museum of Natural History) and Patrice Pruvost (Muséum d'Histoire naturelle de Paris) for access to museum collections; Kenko Takemura for providing fish specimens; Toshiaki Igarashi for access to literature; Tsutomu Hikida, Thomas Near, Ainsley Seago, Helen Whitaker and two anonymous reviewers for providing valuable comments on the manuscript; Ainsley Seago and Jun Inoue for the illustrations in Fig. 1; Aquarium Shop Water House (Yokohama, Japan) for the pictures in Fig. 2; and Jim McGuire and the Museum of Vertebrate Zoology for providing computing facilities. This study was financially supported by a Grant-in-Aid for the biodiversity research of the 21st Century COE (A14) and a Grant-in-Aid for the biodiversity and evolutionary research of the Global COE (A06) from the Ministry of Education, Culture, Sports and Science and Technology, Japan to Kyoto University.
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Well, we have resided in this lodge on 1st of January 2015. Nice accommodation, cosy rooms, very convenient location – only 30 km to Maasai Mara National Park. The cottages are placed right on the high bank of Mara river, so you can behold big hippo family right from view point near the cottages. Especially I would like to point... More
- Also Known As:
- Mara Wildebeest Resort Kenya/Narok

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Wednesday, November 13, 2013
© Copyright 2015 Gwinnett Daily Post
It was a setup.
Gwinnett County police believe 28-year-old Steven Carden and pal Thomas Smith met their female acquaintance at an Atlanta club that night in July. The girl — six years their junior, pretty, tattooed — invited them back to Norcross' Celadon Apartments to hang out.
By morning, Smith was hit over the head outside Building F and robbed. Carden was shot dead.
Police have now identified the woman and her two alleged male accomplices, charging them all with murder.
Earlier this month, Gwinnett homicide detectives learned that Diara Hood, the 22-year-old woman, and 18-year-old Tyler Estrada were already in custody in DeKalb County on separate armed robbery and aggravated assault charges. On Friday, the GCPD SWAT team served a warrant at 26-year-old Jovian Lanus' home on McCord Livsey Road near Snellville.
Gwinnett County police spokesman Cpl. Jake Smith said all three individuals developed into suspects in the weeks following the July 29 incident but extra legwork was required. Each of the suspects have been charged with murder and two counts of armed robbery, with Lanus facing an extra count of aggravated assault.
All three are being held without bond.
In the days following the incident, the surviving victim reportedly told police he and Carden had been walking through the apartment complex, located off South Norcross-Tucker Road, when they had "a brief conversation with two black males." He said he was then struck from behind by an unknown object.
Drugs are not believed to be a factor in the incident, Smith said.
More like this story
- Mid-trial guilty plea for suspect in fatal Norcross ambush ( June 5, 2014 )
- Norcross murder suspect's confession key issue in trial ( June 4, 2014 )
- Ex-Gwinnett teacher indicted for alleged relationship with student ( February 6, 2014 )
- Brawl at park leads to ecstasy arrest ( June 2, 2009 )
- Two charged in armed robbery of Jimmy John's driver ( November 19, 2014 )


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AWTS has been serving Industrial and Municipal customers, with large scale water treatment plants, both high purity and waste water treatment, Zero Liquid Discharge facilities and hydraulic frackiing processes. We specialize in all types of technologies including pretreatment clairification, filtration, RO, UF, MF membrane processes, degasification systems, chemical ...
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Bio-Reaction, Duall, Flex-Kleen and Met-Pro Systems are now Met-Pro Environmental Air Solutions. For decades, these businesses have provided innovative solutions and a diverse line of products and technologies for the collection of particulates, the purification of air, and the destruction of VOCs. In 2011, a new subsidiary was established in South America, Met-Pro Chile ...
Total Water
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Total Water Treatment Systems Inc. was founded in 1943 as an authorized dealer of Culligan water treatment equipment. Total Water Treatment Systems specializes in high-purity water applications. These applications are quite varied and can range from highly purified water for In Vitro Fertilization, biotech, dialysis and microchip applications to specialized water for ...
Roediger BioEnergie GmbH
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Our business activities include planning, design, installation and commissioning of plants for the fields of lean gas, process gas, landfill gas, digester gas and biogas. In addition, our activities focuse on technical services relating to the operation, servicing and maintenance of environmental facilities.
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Being engaged in researching and manufacturing oil purifier, oil filtration and oil recycling equipment, Chongqing Tongrui Filtration Equipment Manufacturing Co.,Ltd is a very professional and reputable manufacturer with unique technology and rich experience in oil purifier industry, all the products manufactured by us confirm to international standards in quality and ...
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Zena sro produces hollow fiber membranes and modules immersed or housed. Developed membranes / modules can be used in MBR, waste water (industry or municipal), contactor, degassing, lab an biotech applications.
Darco Engineering / Darco Industrial Water Sdn Bhd
based in Singapore, SINGAPORE
Listed on SGX Sesdaq in July 2002, Darco is a provider of integrated engineering and knowledge-based water treatment and environmental engineering solutions. It was established in 1999 to design, fabricate, assemble, install, commission and service engineered water systems for industrial use in Singapore and Malaysia. Darco has developed systems and services based on ...
OSMO Membrane Systems GmbH
based in Korntal-Münchingen, GERMANY
OSMO develops and implements valuable industrial membrane separation plants for a multitude of process applications as well as for water treatment.OSMO enables unique synergies as well as a significantly expanded development budget that lays the perfect foundation for new approaches in plant construction.We are certified in all current regulations and long-term ...
Enervac Corporation
based in Cambridge, ONTARIO (CANADA)
Since 1978, excellence in engineering, manufacturing and customer service has made Enervac Corporation a world leader in the manufacture and development of processing and recovery systems for industrial liquids and gases. Enervac Corporation provides practical and economical solutions to wastewater disposal and recycling. Enervac offers systems for waste water recycling ...
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Midweek Madness - Serious Sam Franchise, 80% Off
Save 80% on the Serious Sam Franchise during this week's Midweek Madness*!
Serious Sam 3: BFE is a glorious throwback to the golden age of first-person shooters where men were men, cover was for amateurs and pulling the trigger made things go boom. Serving as a prequel to the original indie sensation, Serious Sam: The First Encounter, Serious Sam 3 takes place during the EarthÂ?s final struggle against MentalÂ?s invading legions of beasts and mercenaries. Set against the collapsing temples of an ancient civilization and the crumbling cities of 22nd century Egypt, Serious Sam 3 is an exhilarating fusion of classic twitch shooters and modern gameplay features.
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Honduras still appear to have plenty of work left to do in the final days before the 2010 FIFA World Cup South Africa™ as they were held to a goalless draw by 10-man Azerbaijan in Austria today.
Wayward shooting meant Honduras never really tested Azerbaijan goalkeeper Salahat Agayev, even after the 73rd-minute dismissal of Sasa Yunisoglu for a rash tackle.
Ramon Nunez was the only Honduras player who managed to force a save in the entire match, but his left-footed shot from the edge of the area was easy enough for the goalkeeper.
Honduras will face Chile, Spain and Switzerland in South Africa 2010's Group H.


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There was plenty more like this--every epistle dumber than the previous--but you get the general idea.
Old English epistol, from Old French epistle, epistre (Modern French épitre), from Latin epistola "letter," from Greek epistole "message, letter, command, commission," whether verbal or in writing, from epistellein "send to," from epi "to" (see epi-) + stellein in its secondary sense of "to dispatch, send" from PIE *stel-yo-, suffixed form of root *stel- "to put, stand," with derivatives referring to a standing object or place (see stall (n.1)).
Also acquired in Old English directly from Latin as pistol. Specific sense of "letter from an apostle forming part of canonical scripture" is c.1200.


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The Ridder Clan Mod is a modification for The Lord of the Rings: The Battle for Middle-Earth II & for The Lord of the Ring: The Battle for Middle-Earth II, The Rise of the Witch-King. This mod changes many things to BFME 2. It will add some new factions, a couple of campaigns, many units & heroes, alot of maps and it will improve the CAH System. This mod changes the gameplay making it work a bit more like BFME 1 and how BFME 2 should have been!
Mordor's Very Special... Orc Pit!
With 2 expansions! The Black Tower and the Tents!


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Creativity involves breaking out of established patterns in order to look at things in a different way. - Edward de Bono, psychologist and writer This post is something new for me. I usually write about our hands-on projects, but I thought it would be fun to take our creative thinking out into the great wide world. In reference to the quote above, not only am I breaking out … [Read more...]
Most Awesome Local Blog!
We recently visited one of our favorite Bay Area kid spots, the Bay Area Discovery Museum in Sausalito, and stayed until the kids could play no more. If you're local, or just might visit San Francisco with young kids, definitely bookmark this as a destination. There are views of the Golden Gate bridge, wonderful nature-based exhibits, and when you walk in you're greeted by this … [Read more...]


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I am happy to post some Holi recipes. Please click the link below each picture to see the recipe. Wishing you all a very Happy Holi! Kaju Katli Besan Laddoo Makmal Poori Mohan Dhal Milk Kheer Rasgulla Shahi Tukra Carrot Halwa Papdi Khakra Chat Pav Bhaji
Tag Archive: Mini Rasgulla
Rasgulla | Easy Rasgulla Recipe
A Royal Treat! I would always think Rasgulla is so difficult to make. My friend always encourages me to try Rasgulla once and see how easy it is. Then I decided to give a try. Once the paneer is ready, you can make the Rasgullas in no time. You just have to pressure cook the …


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Beastly Adventurer
Joined: Nov 2010
Location: here, there, everywhere
Oddometer: 1,858
discussing chinese scooters is always a bit of a minefield, i'm afraid. i always recommend if the person can afford it look elsewhere or for something used, but if for whatever reason a person doesn't want to, i can only try to show them a way to a better seller with a better price, and maybe a chinese make where the chances of getting a better chinese scooter then another may lie
the thing about chinese scooters is, as much as i don't trust the quality, its hard to argue with owners who have 15k, 20k, and 25k miles on theirs, and say they have had little or no problems mechanically. is this the norm? probably not overall, but its out there. and for some, even if they do have problems, it was worth it to them because it comes back down to price. $1000 for a 150cc that went 15K miles so far? maybe it is cheap, and cheaply made. and well, maybe some people just don't care and want a cheap scooter. as i've read in some forums from chinese scoot owners: "so what? you bought one scooter for $4000 that will last three times as long as mine, but i'll just buy 3 scooters over time and still come out paying less then you"
that is the reasoning of many, and if they are riding along fine i really can't tell them how to spend their hard earned money and on what, or put down what they ride. and if they have 10,000 nearly trouble free miles on a china scoot, paid $800 for it, and are happy, happy, happy....even if it breaks tomorrow...i can see why
chinese scooters are a complete crap shoot imo. more knowledgeable scooter people understand and know this. they know what they are buying. an inexpensive gamble. if they lose, they lose. lesson learned. less knowledgeable buyers unfortunately don't know this, and when they lose it hurts. because they expected a good scooter and got trash, or unwittingly bought trash. but not ALL chinese scooters turn out to be complete trash. its just not true, so i can't tell anyone it is
buyer beware, sure. but buy and ride what you want, or can afford. maybe a hard lesson will be learned, or maybe it will turn out ok. it's easy to say to someone "they are all junk. buy Jap or Taiwanese or Italian." but its not my money, so who am i to say anything like that? if someone wants a chinese bike and are intent on getting one i can just point them to some which have better reputations them others, some good pdi advice, a list of parts to replace right off the bat, a decent seller, scootdawg forums, and tell them "good luck"
but telling them to buy a Honda really doesn't do anything for them at all
DudeClone screwed with this post 12-18-2010 at 04:54 PM
Beastly Adventurer
Joined: Sep 2007
Location: In the TARDIS
Oddometer: 3,075
I would like DeBinChe to actually try to help people by posting which Chinese brands and models he has personal experience with and what went wrong with them, in other words be specific. By pointing out the specifics he could save someone a good deal of money and frustration. I suppose the Chinese guy who bought any one of a number of American cars during the 70s and 80s might say that all American cars are crap, not realizing that there may be a huge difference between brands or even models of the same brand.
I am a long time motorcycle mechanic and very experienced with the older Vespa scooters but I've only ridden one Chinese scooter in my entire life. My experience with Lifan and Hensim motorcycles lead me to believe that there may be some decent quality Chinese scooters out there. I'd like to know which are good and which are bad and why.
Xtremely Bad Example
Joined: Oct 2010
Location: Central New England
Oddometer: 313
DudeClone, you forgot one other thing....
If they have a little knowledge of how to turn a wrench, they can usually fix the issues of these cheap scooters without spending too much $$$.
And DaBinChe, if it's coming out of China, It's Not from Japan or Taiwan.
Yes there are companies that contract out to the Chinese manufacturers, the difference between whether the label says Sony or S0ny, is the QC end of things. Sort of like the "Made In USA" back in the 1960's when there were cheap electronics that were "Made In Usa" Usa Japan.
I do love your hypocrisy of buying a chinese Honda Elite. If Chinese products are so bad as you put it, why did you buy one?
Joined: Apr 2009
Location: BryanTexas
Oddometer: 37
A suggestion is to run over to http://scootdawg.proboards.com and browse around. Ya gotta be willing to work on it yourself cause the dealers who'll work on them are few and far between. You'll find a lot of talk about a PDI (post delivery inspection). You basically take the plastic off and check every nut, bolt, put dialectric grease on all electrical connections. Took me 2 days to do this and I haven't had a single problem. It also got me really edumacated about the inner workings of the scoot.
It does 60 without breaking a sweat, 70 if I ask, and gets about 65mpg. So far it's been a lot of fun. Cost $1,800 to my driveway.
Syntroxis screwed with this post 12-18-2010 at 06:20 PM
Beastly Adventurer
Joined: Dec 2007
Location: Santa Cruz Mtns.
Oddometer: 1,255
One of my jobs is fixing chinese scooters...I've pretty much seen it all. Everything from pot metal use to non-fuel line used as fuel line to stripped treads on a brand new bike to incorrect bolts to parts not being the same on the same bike coming from the same factory to lines going where they are not suppose to, you name it and even things you wouldn't believe can occur does occur. I've worked on hundreds if not thousands of these chinese scoots so yeah that makes me more of an expert then most of the folks on here. I can't count how many blown motors I've seen and rebuild, carbs I'd had to fix/replace, electrical issues of every kind. I really can't name a specific thing that goes wrong with chinese scoots cause my experience shows that anything and everything can go wrong with them.
That $1k new scoot that folks think they got for so low will more then catch up to them in repair bills and headaches and down time.
The most mileage that I have ever seen on one is 26k miles which is most certainly not the norm. The guy was an older fellow and he rode the thing very conservatively...in the bike lane conservative so that was how he got so many miles on the thing. But boy when I had to do work on that bike everything was literally about to fall apart. But for the most parts chinese scoots have a shelf life of up to 5k miles before the repairs cost more then the value of the bike. It is actually very rare that a bike will go 1k miles without having some type of issue, needing some type of work.
Beastly Adventurer
Joined: Sep 2007
Location: In the TARDIS
Oddometer: 3,075
Your posts are worthless unless you provide make, model, and problem information.
Beastly Adventurer
Joined: Dec 2007
Location: Santa Cruz Mtns.
Oddometer: 1,255
I didn't say coming form Japan or Taiwan, I said Japanese or Taiwanese companies. I bought a honda product and that is the difference. My point is don't get something from a chinese company coming out of china.
What you are saying is like saying that the honda cars coming out of the southern states is not made by honda but instead made by GM...don't be mistaken it is every bit a honda. Maybe you don't understand mfg. Here is an example: when I used to work for Intel in their FAB, they had factories all over the world including undeveloped countries. How they make chips is the same in very location, the processes/procedures/specifications were identical. So it did not matter if your intel chip was from their headquarters in Santa Clara or Isreal or any location it was the same. This is how Honda does things too their mfg plants world wide does things identically to each other, to the standards of Honda specifications.
Beastly Adventurer
Joined: Dec 2007
Location: Santa Cruz Mtns.
Oddometer: 1,255
Are you aware that there isn't that many chinese manufacturers of scooters? You can easily find a half dozen "brands" coming from one manufacturer. The best example of this is the Fly ilbello and it's various versions...just about every American "brand" chinese scooter company has this very same bike from the same manufacturer, they are just badged/"branded" differently here in the US. This is cause the US brands aren't manufacturers only importers. These importers (independent of each other) go to this manufacturer and orders a hundred in X color with Y options and put on their badge/"brand" and brings it over here.
You see the problem is not like normal vehicle problems where a specific vehicle let say toyota prius, with the gas pedal getting stuck from the floor mats, has a specific problem and can be easily address. The problem with chinese scoots is that it is completely unpredictable as to what each individual bike has issues with...like a box of chocolate you never know what you're gonna get. So let me repeat there is no specific issues with any specific model it really is a roll of the dice. Because if there were a specific issue with a specific model this implies that there is some underlying issue like design flaw or manufacturing flaw. In the case of chinese scoots the flaws mainly has to do with the whole manufacturing process, there is no consistency/repeat-ability anywhere so problems will arise from any thing.
Beastly Adventurer
Joined: Jul 2010
Location: Morgan Hill CA
Oddometer: 2,880
There is some truth to this, but this is only true on the low end crappy cheap chinese scoots. But I think that there are some good bikes coming from china, like lifans and the high end companies.
Oh, and hugemoth, do you have any info on Harley building their bikes in china at any point? I have heard about this, but I don't know where. This is the only thing that CANNOT be built in china! Harley motorcycles are the American legend, they are one of few motorcycles being built in America, actually one of very few major American companies other than Ford. Point is, DaBinChe, sure quality may be the same, but it won't be a true Harley. This is to your other post, saying Hondas built in America are still Hondas. This is true. But it is going to be different with Harleys, I mean that is one of the reasons we Harley riders pay $20,000 for a cruiser when a Honda VTX 1800 is like $12,000.
Yes, the Honda doesn't feel quite the same, but this is a factor. I mean what happened to that good old Milwaukee steel? They have been made here for over 107 years, and know they are going to change that?
It is already uncool that my $500 Harley jacket is made there!. But I do have understanding in why this is a possibility, sales. They will reduce costs BIG TIME. But they don't even realize what they are going to start! Oh boy, all those bikers who stay true to good old American Harleys and make fun of rice burners! They are going to loose so many sales it might put the 107+ year old company to an end!
But this is unlikely, hopefully they will come to their senses!
Big Adventurer
Joined: Apr 2010
Location: One of the Burj's
Oddometer: 2,510
My Zinda wasn't too bad. I paid $400 for it in June, and it was less than a year old. It did the job...
Sometimes it didn't do the job...
Yep, "new" crank, new piston and barrel, not sure if they swapped the head too... all for $100.
It's probably worth replacing the wheel bearings if you have one.... I couldn't get over 85kph Vmax with this bearing
They chew out the cush rubbers a bit too...
And they aren't very good for carrying gear when you have to loan half your occy straps to someone else
They aren't bad off the beaten track....
But they get you there....
But best of all.... having mine made me go and buy a KTM 950 se when I got home....
A man should only do the work that is required of him. To do more is a form of greed.
Ride through Oz and Asia
Beastly Adventurer
Joined: Nov 2010
Location: here, there, everywhere
Oddometer: 1,858
^ holy cow
but some people are cheap. or just want a scooter for a year, or do in fact want a mechanical project. many see scooters as disposable items, things to buy and use and throwaway. others ride one or two hundred miles a month. and so they get the cheapest scoots around. cheap plastics, cheap parts? they don't care
the original poster in this thread has a friend who wants a BIG 250cc. even after all the other recommendations for used scooters, smaller brand name bikes at close to the same price, and warnings against.....his friend is still considering a chinese 250cc. some people in this thread (and at scootdawg) have 250cc scooters and have over 12,000 trouble free miles on them over two years. they paid $1700 or so for them. so even if they will have more problems then the bike is worth someday soon....they figure "hey, time for a new scooter!"
some value quality more then others. some want long lasting durability and a high mileage, low maintenance machine. yet others will just say "$5000? no thanks. i paid $1600 and i'll do it again"
and its hard to tell them don't do it if a BIG 250cc is what they want or need, but can't afford otherwise or are too cheap (frugal) except to buy a chinese scoot
plus don't forget, not everyone has a dealer nearby who has brand name scoots. and some people live in rural areas where private party used selections are low, or nothing at all. i live in an area where i can go get a low mileage 2005 Piaggio BV 200 for $2000 tomorrow, or a lower mileage 2006 Kymco People 250 for under $1500. and their is more to choose from. but for many that is not the case. so a new scooter delivered to their door is an attractive offer. especially if they are mechanically inclined
DudeClone screwed with this post 12-19-2010 at 10:46 PM
Beastly Adventurer
Joined: Sep 2007
Location: In the TARDIS
Oddometer: 3,075
I remember Zongshen is the company that will be making HD as well as Piaggio (vespa) but don't know the details. The article implied that the HDs will be for the Asian market, at least at first.
Beastly Adventurer
Joined: Nov 2010
Location: here, there, everywhere
Oddometer: 1,858
well, Zongshen certainly know how to build a lot of scoots. i would think manufacturing for HD and Piaggio would improve the quality of Znen scooters, too. if the company wants it to
how this would happen, idk? i doubt the same quality control that goes into a Piaggio model would be used for the average Znen, but if they can start using some upgraded parts and components sourced from or copied from these bigger players, who knows?
Beastly Adventurer
Joined: Jul 2010
Location: Morgan Hill CA
Oddometer: 2,880
Thanks. I just hope they keep those Harleys in the chinese market. I won't buy a new Harley if they start building them there, only keep my current and buy used. It is ok with me for other bikes like Honda, but as I said, Harley just won't be the same.
Harley is probably suffering in America due to the economy and such, but that is no excuse to go BUILD bikes in china. There is a big difference between giving Harleys to the chinese market for chinese people to ride, that is great! So they can enjoy Harleys and it will help their profit.
But we all know how many problems this is going to cause with the die hard Harley fans like me. I am not just for Harleys, but I love riding Harleys. I love that big powerful twin feel, that rumble with loud pipes, and that powerful torquey motor that just keeps rumbling forever. That chrome idle and American machine make these things a dream to ride! Just like all the BMW guys love their beamers, I love Harleys.
Don't get me wrong, I also LOVE all other bikes. If it its got 2 wheels and a motor, I'll love it! I'll ride a Honda, Beamer, Kawasaki, Ducati, etc. But Harleys are a part of my life. I love to tour on them.
If Harley really wants to make some more money, they need to get into the SMALL DISPLACEMENT motorcycle industry. Yes, exactly. They need to make a cool 250 cc V twin. Think about it. If a beginner who has never ridden 2 wheels wants a bike, what are the chances he or she will feel confident on a 1600 cc 850 pound monster bike? They will obviously go buy a Honda Rebel or Ninja 250. They should make a 50 cc bike with a real V twin as well.
And for any Harley riders, how cool would a new XR 1200 WITH a VROD ENGINE be?
But we all know what the hard core Harley riders are going to say when their bikes can be considered not only rice burners, but discount ride burners!
btcn screwed with this post 12-20-2010 at 09:37 AM
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I downloaded the Polygon / Stars plug-in but it only let's you draw equilateral triangles. Is there a plug-in or method that let's you create any kind of triangle? I can't just use the line tool because it makes the edges of the shape (the vertices of the shape) round or irregular instead of pointy.
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I recently put in a plug for the N.C. Aquarium at Fort Fisher's albino alligator naming contest, and I'm proud to report the aquarium has opened its exhibit and named its star: Luna.
According to the aquarium, visitors got a sneak peek at the exotic creature in March during "Scales and Tails" and immediately initiated the naming contest.
The aquarium received 1,500 submissions both onsite and online, some from as far away as Germany and Scotland. Entries include Lilly, Allie, Pearl and Opal.
So I've finally grown concerned about all the plastic grocery bags drifting around out there—getting lodged in trees, floating aimlessly along the highway like lost balloons.
I realized if I had a dollar for each homeless bag littering our otherwise pristine environment (except for the beer cans), my 401K and I could reconsider retirement.
Apparently, nobody is offering reimbursement for these little eyesores and wildlife hazards.
In April, The Brunswick Beacon requested access to e-mails generated to and from county government administration department heads.
The county promptly responded to the request, providing the newspaper access per N.C. Public Records Law. Often we have good responses to public records requests from the county.
However, following our most recent request, at a May 20 county government department heads' meeting, extended discussion took place about the state's public records law and the federal Freedom of Information Act.
A lot can happen in 50 years. Just ask any of the original residents of Ocean Isle Beach.
Ocean Isle Beach was a 7-mile stretch of pure beach some 50 years ago. When Odell Williamson and M.C. Gore purchased the island in 1953, they sold only a handful of lots before Hurricane Hazel hit the following year and wiped away most if the island's initial developments.
Over time, the town rebuilt and grew into one of the best restored beaches in the country, according to the American Shore and Beach Preservation Association.
There's something about severe weather that really intrigues me. From swirling clouds and darkening skies, to booms of thunder and flashes of lightning, reporter instincts quickly kick in when rough weather approaches.
Now that June 1 has come and gone, all Brunswick County residents' severe weather intrigue should be piquing as hurricane season is finally here.
To the editor: I appreciate the experience John Heidtke has had and thank him for his service. His column in the May 21 edition was enlightening and reassuring. but the organization he quoted for statistics does not exist.
The Violence Policy Center is a rabid anti-second amendment organization and has nothing to do with the National Rifle Association. The statistics quoted are close but not complete.
To the editor: Just when you thought the Bush/Cheney legacy could sink no lower, new torture pictures surface.
A Sydney paper published a few; Wired Magazine had published others in February 2008. Now, General Taguba confirms U.S. personnel raped men, women and children in their custody and sexually abused them with nightsticks, phosphorescent tubes, etc.
Can Americans handle the truth, or do we prefer to sweep these atrocities under the rug, giving the Bush/Cheney administration a pass?
To the editor: From reading articles in newspapers across the state, particularly Raleigh and Charlotte, it appears media have been misinformed, misunderstand or simply accept statements given them by pro-forced annexation proponents.
Annexation would not come to a screeching halt if those being victimized by forced annexation had a vote. Eighty-six percent of North Carolina's annexations are voluntary; and they have a vote.
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State does surprise inspection of jail
County will get a verbal report on results today
State officials started a surprise annual inspection Thursday of the Nueces County Jail and should finish by today, according to the Texas Commission on Jail Standards executive director.
The last Texas Commission on Jail Standards annual inspection was Aug. 11 and showed no problems. The U.S. Marshals Service pulled federal prisoners from the jail earlier this month after finding shoddy living conditions during an inspection.
The state commission must conduct 20 percent of its annual inspections by surprise visit, said Terry Julian, commission executive director.
"We decided this would be a good opportunity to conduct one of those surprise inspections," Julian said.
The commission also can conduct surprise visits between annual inspections if problems or complaints arise.
County officials should receive a verbal report today after the inspection is finished, Julian said. The three state inspectors then will turn in their information for a full report. The Commission on Jail Standards gives jails a time frame of as much as a year to fix any problems, with the timeframe depending on the violation. Jails that don't remedy problems can face a range of penalties, including closure.
The state findings will have no bearing on whether federal prisoners are returned to the jail, U.S. Marshals spokesman Carlos Trevizo said.
"The state does what they do, and we do what we do," Trevizo said. "They advised us they were coming, but that is about it. It has nothing to do with us returning the prisoners."
Sheriff Rebecca Stutts said Thursday she was not surprised by the unannounced visit.
"The jail inspection normally takes place during the summer months," she said. "This is not out of the ordinary."
State reports from the past five years show inspections in March, April, July and August.
Stutts also said she did not anticipate any problems in the inspections.
"Of course, they are professionals in their field," Stutts said. "They have a list of standards to go by. I look forward to their findings."
Photos taken by marshals June 7 show moldy showers, peeling paint, clogged toilets, exposed wiring and inmates sleeping on floors.
Stutts said cleaning was completed and "continues to be done on a daily basis." Larger maintenance issues, such as plumbing, will take longer to address. Stutts requested a workshop with county commissioners to decide on a plan for the repairs.
County Commissioner Chuck Cazalas, who toured the jail earlier this week, said he saw organizational problems that should be addressed.
"It lacked an organized approach to cleaning and an organized approach to painting," he said. "But in fairness, Sheriff Stutts agreed and said that she is going to get on top of it and put a plan together to have a more systematic, organized way of cleaning and painting."
Cazalas said he could see why marshals made the decision to remove prisoners.
"It's all because they need to get in there and get the job done - there is no excuse for uncleanliness," he said.
Contact Denise Malan at 886-4334 or at HYPERLINK mailto:firstname.lastname@example.com email@example.com. Jaime Powell contributed to this report.
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