All sequence files are available from the GenBank database (accession number(s) KT998333- KT998452).
Yellow fever continues to be a problem in sub-Saharan Africa with repeated epidemics occurring. The mosquito Aedes bromeliae is a major vector of yellow fever, but it cannot be readily differentiated from its non-vector zoophilic sister species Ae. lilii using morphological characters. Genetic differences have been reported between anthropophilic Ae. bromeliae and zoophilic Ae. lilii and between forest and domestic populations. However, due to the application of different molecular markers and non-overlapping populations employed in previous studies, interpretation of species delimitation is unclear.
DNA sequences were generated from specimens of Ae. simpsoni s.l. from the Republic of Benin, Tanzania and Uganda for two nuclear genes apolipophorin 2 ( apoLp2) and cytochrome p450 ( CYPJ92), the ribosomal internal transcribed spacer region ( ITS) and the mitochondrial cytochrome c oxidase ( COI) barcoding region. Nuclear genes apoLp2 and CYPJ92 were unable to differentiate between species Ae. bromeliae and Ae. lilii due to ancestral lineage sorting, while ITS sequence data provided clear topological separation on a phylogeny. The standard COI barcoding region was shown to be subject to species introgression and unable to clearly distinguish the two taxa. Here we present a reliable direct PCR-based method for differentiation of the vector species Ae. bromeliae from its isomorphic, sympatric and non-biomedically important sister taxon, Ae. lilii, based on the ITS region. Using molecular species verification, we describe novel immature habitats for Ae. lilii and report both sympatric and allopatric populations. Whereas only Ae. lilii is found in the Republic of Benin and only Ae. bromeliae in Tanzania, both species are sympatric in Uganda.
Our accurate identification method will allow informed distribution and detailed ecological studies that will facilitate assessment of arboviral disease risk and development of future targeted vector control.
In Africa, epidemic outbreaks of yellow fever continue despite the availability of an effective vaccine. Effective understanding of disease epidemiology and control requires the ability to reliably identify vectors of yellow fever. The mosquito Ae. bromeliae, a competent vector of yellow fever virus, cannot be reliably morphologically differentiated from its sister species Ae. lilii, which does not bite humans and so does not transmit yellow fever. DNA sequencing of four molecular markers allowed comparisons of how they perform at distinguishing these species. We found that the mitochondrial cytochrome c oxidase ( COI) barcoding region and nuclear apolipophorin 2 ( apoLp2) and cytochrome p450 ( CYPJ92) were unable to reliably distinguish these species. Conversely, genetic variation at the internal transcribed spacer region ( ITS) was able to confirm the vector Ae. bromeliae and non-vector Ae. lilii as distinct species. Based on ITS sequence differences, we developed a robust molecular method to identify the vector Ae. bromeliae from its sister species Ae. lilii. Consequently, we find that these species use the same larval habitats including banana, cocoyam and Dracena spp. in Uganda. Whereas only Ae. lilii appears to be present in Benin and only Ae. bromeliae in Tanzania, we confirm that both species occur in Uganda. Reliable species designation will promote more detailed studies of distribution, ecology and vector status essential for disease risk assessment and mosquito control.
Correctly identifying the vector species involved in mosquito-borne disease transmission is fundamental to predicting disease outbreaks, ascertaining general risk to the human population and targeting control efforts. Despite this necessity, the reliable identification of mosquitoes is problematic in many cases, including the medically important Aedes simpsoni complex. This complex comprises three known species including Ae. simpsoni, Ae. lilii and Ae. bromeliae. Among these three species, Ae. bromeliae is an important vector of yellow fever virus (YFV) and potentially other arboviruses [ 1 ]. Yellow fever has increased in incidence as a result of urbanisation and changes to public health policy [ 2 , 3 ]. The burden of yellow fever in Africa is estimated to be 130,000 cases a year, 85,000 of which result in deaths, despite the availability of a vaccine [ 4 ]. After mass immunisation campaigns at the beginning of the 20 th Century, YFV was successfully reduced in targeted countries. However, YFV outbreaks are causing renewed attention. In West Africa, 13 out of 14 countries known to host YFV now report cases regularly and have experienced epidemics since 2000 [ 5 , 6 ]. Further concern has arisen over identification of a novel YFV genotype implicated in recent outbreaks including the first reported outbreak in Kenya, East Africa (1992–1993) [ 7 ]. Related genotypes have also been reported in Sudan in 2003 and 2005 and more recently in northern Uganda in 2010 [ 8 – 10 ]. Although much attention has been given to understanding yellow fever disease epidemiology [ 11 ], relatively little is known about the mosquito vectors.
Ten species are currently known within the wider Simpsoni Group (Huang 2004). Within this group, Theobald [ 12 – 14 ] originally described three species belonging to the Simpsoni Complex ( Ae. simpsoni, Ae. bromeliae and Ae. lilii). Despite these separate species designations, many entomologists referred to all simpsoni-like mosquitoes within this complex as the nominotypical species, Ae. simpsoni [ 15 – 21 ]. In Uganda, it was found that some populations of Ae. simpsoni s.l. were anthropophilic and attracted to human bait, whereas others were not attracted to man despite local abundance [ 20 ]. Furthermore, mosquitoes collected from zoophilic and anthropophilic populations showed different feeding preferences in the laboratory with a preference for rodents or humans, respectively [ 20 ]. These findings led to a re-examination of mosquito morphology and subsequently Huang [ 22 , 23 ] provided a full description of the component members of the Simpsoni Complex, reviewed their ecology and drew attention to the incorrect use of former nomenclature. Aedes simpsoni, which has only been reported from South Africa and Swaziland, is not implicated in human disease transmission [ 22 , 23 ]. The anthropophilic yellow fever vector Ae. bromeliae is widespread on the African continent [ 23 ]. In contrast, Ae. lilii, only previously reported from Sudan, Ethiopia and Uganda, has never been reported biting man and is thus not considered to be involved in disease transmission [ 22 , 23 ]. Although we know these three species have different distributions, biting behaviour and vectorial abilities, these need to be fully characterised. Understanding ecology and epidemiology requires that species can be distinguished from one another. However, controversy over mosquito taxonomy means that the anthropophilic disease transmitting Ae. bromeliae cannot be reliably distinguished from its zoophilic sister species Ae. lilii [ 24 , 25 ].
Large scale detailed studies of the Simpsoni Complex by Huang [ 22 , 23 , 26 ], reported that Ae. simpsoni s.s. can be distinguished from Ae. bromeliae and Ae. lilii in that it has simple claws on the mid tarsi, as opposed to toothed mid-tarsal claws present in the latter two taxa. Aedes simpsoni can also be easily distinguished from conspecifics by its distinct tarsomere scaling pattern [ 22 , 25 ]. However, Jupp & Kemp [ 24 ] reported variation in tarsal claw morphology of Ae. simpsoni and Ae. bromeliae, and questioned the reliability of this diagnostic character. In this study, we focus on the more widespread taxa of the complex, Ae. bromeliae and Ae. lilii. These sister taxa can sometimes be reliably identified morphologically based on tarsomere banding patterns, but only when this character is exhibited at the extremes of its range as banding patterns overlap between the species [ 25 ]. Lutwama & Mukwaya [ 25 ] questioned the usefulness of tarsomere banding patterns as a diagnostic character for Ae. bromeliae and Ae. lilii. They observed variation in scale ornamentation on an almost continuous scale and found that progeny from the same mother could be identified both as Ae. bromeliae or Ae. lilii based on this morphological character.
The unreliability and practicality of using morphological based methods for routine field-based identification in the Simpsoni Complex has led to attempts to delimit species boundaries through molecular methods. In a study of the Simpsoni Complex, excluding the southerly distributed Ae. simpsoni, Mukwaya et al. [ 27 ] found that anthropophilic populations from Kenya and Uganda form a distinct genetic clade separate from non-sympatric and non-anthropophilic populations from Uganda and Nigeria based on the non-coding internal transcribed spacer ( ITS) region of ribosomal DNA. In that study, species were designated according to both blood feeding preference (because Ae. lilii is zoophilic whereas Ae. bromeliae are human vectors) and on tarsomere banding patterns which were distinctive for some specimens [ 22 , 23 , 27 ]. Recently, Walter et al. [ 28 ] inferred the presence of Ae. bromeliae and Ae. lilii in sympatry in Rabai, Kenya based upon genetic differences between domestic/peri-domestic and forest populations. However, how these putative species relate to those characterised by Mukwaya et al. [ 27 ] remains unclear as they used different molecular markers; two nuclear genes, apolipophorin 2 and cytochrome p450 ( CYPJ92).
A 658 bp region of the cytochrome c oxidase ( COI) gene has been widely adopted as a DNA barcoding standard for species identification because it shows high utility in discriminating between closely related taxa as well as resolving phylogeographic groups within species [ 29 – 31 ]. However, sole use of mitochondrial DNA to delimit species has been questioned because of the potential for pseudogene development and introgression, which may limit the ability of mtDNA markers to resolve closely related species [ 32 , 33 ]. Combined analysis of both mitochondrial and nuclear genes can improve phylogenetic resolution since these markers evolve at different rates and so target different levels of the phylogenetic tree [ 34 ]. Another barcoding candidate is the nuclear internal transcribed spacer ( ITS) regions of the ribosomal gene cistron that comprises the 18S, 5.8S, and 28S genes, an external spacer region and two internal spacer regions ITS1 and ITS2 [ 35 ]. The ITS regions evolve at a rapid rate in the absence of functional constraint [ 36 , 37 ]. Because they are tandemly repeated in the genome, the ITS spacer regions are also subject to concerted evolution whereby paralogues are homogenised by genetic exchange [ 36 , 37 ]. Consequently, paralogues remain genetically similar within species while showing high levels of interspecific divergence [ 37 ]. Although not useful in all taxonomic groups, ITS can differentiate between sister species in a large number of cases and has been widely used to delimit closely related mosquitoes in Anopheles complexes [ 35 , 38 – 42 ].
We seek to expand previous work on genetic differentiation of Ae. bromeliae and Ae. lilii in the Simpsoni Complex by determining how the putative species identified by Mukwaya et al. [ 27 ] and Walter et al. [ 28 ] relate to one another. We achieve this by sequencing the same mosquito samples at previously used molecular markers ( apoLp2, CYPJ92 and ITS). In addition, we were able to evaluate the utility of various molecular markers, including the ITS and COI regions, in determining species bounds. We apply these findings to develop a molecular identification method based on variation in the ITS region to distinguish the disease vector Ae. bromeliae from its non-vector sister species Ae. lilii. Applying this method to mosquitoes from Benin, Uganda and Tanzania enabled us to generate reliable findings of their ecology and distribution.
Mosquitoes of the Simpsoni Complex were collected as larvae from natural breeding sites in Tanzania (n = 36), Uganda (n = 50) and the Republic of Benin (n = 24) from locations detailed in Fig 1 and S1 Table from 2009 to 2014. Immature habitats sampled included the leaf axils of Musa spp. (banana), Colocasia spp. (cocoyam/taro), Dracaena spp. and in tree holes. To avoid biasing the dataset with siblings, each discrete habitat was treated as a separate collection and only one individual per collection was taken for genetic analyses. Where possible, immatures were reared though to adults; otherwise, larvae destined for DNA analysis were stored in 95% ethanol. Adults were desiccated with silica for optimal DNA preservation and either stored in BEEM capsules or pinned. Larvae were preserved in ethanol for later extraction of DNA. All adult mosquitoes were identified as belonging to the Simpsoni Complex using the morphological identification key in Huang [ 26 ], and a subsample of these, and outgroup taxa including Ae. aegypti and Ae. aegypti formosus, were morphologically verified by Dr. Yiau-Min Huang.
Also mapped are the previously described sampling points for Ae. bromeliae (red triangles), Ae. lilii (blue triangles) and Ae. simpsoni s.s (green triangles) based on morphological identification, taken from Huang [ 23 ]. The map was created in QGIS (QGIS Development Team) made with Natural Earth.
DNA was extracted from whole larvae or a single leg of an adult mosquito using the modified phenol-chloroform method in Surendran et al. [ 44 ]. Forty individuals of the Simpsoni Complex were amplified and sequenced for a region of the mitochondrial COI gene with universal primers LCO1490 (5’GGTCAACAAATCATAAAGATATTGG’3) and HCO2198 (5’TAAACTTCAGGGTGACCAAAAAATCA’3) [ 45 ] using a protocol recommended by the Consortium for the Barcode of Life (
Internal transcribed spacer regions 1 and 2 were amplified from six individuals from Uganda and three individuals from Tanzania that were selected to represent the genetic diversity we observed at the COI gene. This was achieved with the 18SFHIN and CP16 primers (5’-GTAAGCTTCCTTTGTACACACCGCCCGT-3’ and 5’-GCGGGTACCATGCTTAAATTTAGGGGGTA-3’, respectively) [ 46 ], as used by Mukwaya et al. [ 27 ]. To generate PCR products, 1 unit of high fidelity MyFi DNA Polymerase (Bioline, UK), 2X MyFi Reaction Buffer, 0.8 μM forward and reverse primer and 1–10 ng of template DNA were used under the following conditions at 30% ramp speed; 95°C for 3 min followed by 30 cycles of 95°C for 30 sec, 58°C for 45 sec and 72°C for 45 sec with no final extension.
Thirteen Individuals, including eight of the same individuals sequenced at the ITS region, were also sequenced for regions of the nuclear genes apolipophorin 2 ( apoLp2) and cytochrome p450 ( CYPJ92) first described by Brown et al. [ 47 ] and used by Walter et al. [ 28 ] in the Simpsoni Complex. This included verified Aedes aegypti formosus collected from a tree hole in Tanzania, for use as an outgroup as was the case in Walter et al. [ 28 ]. Another marker, short-chain dehydrogenase-reductase ( SDR) also used by Walter et al. [ 28 ] in the Simpsoni Complex was not used because it produced multiple nonspecific bands on amplification. PCR products were generated as in Walter et al. [ 28 ]. PCR products were purified with the GenElute PCR clean up kit (Sigma-Aldrich, UK) and Sanger sequenced in forward and reverse directions using the amplification primers. Sequences were generated with BigDye Terminator v3.1 cycle sequencing kit (Applied BioSystems, UK) on an Applied BioSystems 3730 automated sequencer.
ITS products were cloned using the P-GEM cloning kit (Promega, UK) as per instructions. Transformants were blue/white screened and colonies with inserts stored in TE buffer for PCR amplification. Universal M13 primers (5’-TGTAAAACGACGGCCAGT-3’ and 5’-CAGGAAACAGCTATGAC-3’) [ 48 ] were used to amplify cloned ITS products in the following 13μl reaction; 1.25 units of BIOTAQ DNA Polymerase (BioLine, UK), 1 X NH
Cloned ITS sequences were aligned with Mukwaya et al.’s [ 27 ] from several locations in Africa ( S1 Table) and Le Goff et al.’s [ 43 ] sequence data from two Ae. bromeliae originating from the Indian Ocean island of Mayotte available from GenBank (KF135509-10) using the program Geneious v5.4.7 [ 49 ]. Primers were designed based on this alignment to discriminate between species. Several putative species-specific primers were trialled under a wide range of PCR conditions. However, the only primers that worked effectively generated species-specific PCR products that differed by ~30 b.p. Due to this small size difference we recommend that the primer pairs specific for each species are run in separate PCR reactions. Non-specific banding was a feature of all primers tested which we suspect may result from variable indel length among ITS copies within an individual. Although PCR protocols were optimised to reduce non-specific banding, some extra banding can be visible but does not obscure amplification/non-amplification of the species-specific PCR product.
The developed primers are nested within the ITS1-2 region; therefore PCR products encompassing this region were first generated with the 18SFHIN and CP16 primers as described above. These PCR products were purified with the GenElute PCR clean up kit (Sigma, UK) and 0.5 μl was used as template in a 25 μl reaction with 0.6 units of Go Taq Hot Start polymerase (Promega, UK), 1 X NH
Sequence alignment was achieved with Geneious v5.4.7 [ 49 ]. For the apoLp2 and CYPJ92 datasets, files were prepared using seqPHASE [ 50 ] and PHASE v2.1 was then used to infer haplotypes [ 51 ]. The haplotypes of Ae. aegypti formosus outgroups were determined through alignment with the relevant datasets for Ae. aegypti in Brown et al. [ 47 ] and using the program PHASE. Sequences for two Ae. aegypti from Brown et al. [ 47 ] were used as outgroups in conjunction with data generated during the present study. The COI dataset was aligned with two COI sequences of Ae. bromeliae from the Indian Ocean island of Mayotte available from GenBank (KF135496-97) [ 43 ].
Neighbour joining (NJ) trees were constructed in MEGA 6 [ 52 ]as in Mukwaya et al.[ 27 ] and Walter et al. [ 28 ] using the best available substitution model as chosen by JModelTest [ 53 , 54 ]. The Tamura-Nei model with uniform rates among sites was used to construct NJ trees for COI. The Kimura 2 parameter model (K80) with uniform rates among sites was used for ITS sequences and K80 with 0.8 gamma-distributed sites was used to construct trees for apoLp2 and CYPJ92. For all genetic markers, missing data including indels were excluded from analysis. Topological support was determined through 1000 bootstrap replications. The ITS sequence tree was constructed without an outgroup because the high level of divergence between the outgroup and the ingroups presents a challenge for sequence alignment. The two ITS sequences of Ae. bromeliae from Mayotte [ 43 ] were not included in the NJ tree because they did not overlap with the sequences generated here.
A hierarchical AMOVA was performed on the ITS sequence data generated in this study together with Mukwaya et al.’s sequences [ 27 ] (n = 69) in Arlequin v3.5 [ 55 ]. A hierarchical AMOVA was also performed on the apoLp2 (n = 13) and CYPJ92 (n = 13) sequences generated in this study.
Four fixed point substitutions and eight indels of varying length were observed between species in the first 300 bp of the ITS sequence alignment. The neighbour joining tree of ITS sequences revealed two major clades with a bootstrap support of 98% ( Fig 2). As reported previously, the 46 sequences from Mukwaya et al. [ 27 ] clustered into one or other of the two clades depending on their inferred host feeding preference and which they accordingly designated as Ae. bromeliae (anthropophilic) and Ae. lilii (non-anthropophilic). Sequences of all nine Simpsoni Complex individuals from this study also belonged to one or other of these clades, and we therefore identified them as Ae. bromeliae or Ae. lilii with reference to Mukwaya et al.’s [ 27 ] ITS-based species designation.
Red labels represent Ae. bromeliae while blue labels are Ae. lilii. Sequences from Mukwaya et al. [ 27 ] (+) are annotated. Bootstrap support values above 55% are shown.
Mosquitoes collected from Kanyawara, Bundibugyo (Bwamba region) and Najjembe in Uganda represented both ITS clades. In comparison, individuals collected from Tanzania strictly aggregated into the ITS lineage of Ae. bromeliae. AMOVA revealed that almost all sequence differences can be explained by variation between species (92.65%), while there is little variation between populations within groups and within populations (0.25% and 7.19%, respectively).
Sequencing of individuals at both ITS and nuclear genes apoLp2 and CYPJ92 allowed us to compare the ITS-based species designation of Mukwaya et al. [ 27 ] with that of Walter et al. [ 28 ]. Two well-supported lineages were observed in NJ trees for the nuclear genes apoLp2 and CYPJ92 with 100% bootstrap support (Figs 3 and 4). Individuals confirmed as Ae. bromeliae or Ae. lilii using the ITS region with reference to Mukwaya et al. [ 27 ], all fall into a single clade for each of the nuclear genes, apoLp2 and CYPJ92. For both gene trees, the other lineage comprises only the sequences from the forest species in Walter et al. [ 28 ] that they referred to as Ae. lilii. A hierarchical AMOVA revealed that nuclear genes apoLp2 and CYPJ92 could not effectively distinguish between Ae. bromeliae and Ae. lilii with only 0.86% and 18.94% of genetic variance between species groups, respectively. Conversely, there was relatively high genetic variation among populations within species groups (37.04%, 10.28%, respectively) and within populations (62.10%, 70.79%, respectively).
Red labels represent Ae. bromeliae while blue labels are Ae. lilii. Sequences from Walter et al. [ 28 ] (*) and Ae. aegypti outgroups (~) from Brown et al. [ 47 ] are annotated. Bootstrap support values for branches above 55% are shown.
Red labels represent Ae. bromeliae while blue labels are Ae. lilii. Sequences from Walter et al. [ 28 ] (*) and Ae. aegypti outgroups (~) from Brown et al. [ 47 ] are annotated. Bootstrap support values for branches above 55% are shown.
There are two phylogenetic clusters with high bootstrap support (90%, 90%) and two with weak bootstrap support (56% and 43%) in the NJ tree for mitochondrial COI with individuals tending to cluster according to geographic origin ( Fig 5). The sequences from Mayotte cluster with Ae. bromeliae from Tanzania. Individuals designated as Ae. bromeliae or Ae. lilii according to ITS sequence variation tend to group according to species within the same clades, but there are three exceptions. One individual from Uganda, exhibiting the ITS sequence of Ae. bromeliae clusters within a grouping of Ae. lilii while two individuals from Uganda and Benin, identified as Ae. lilii, cluster within a grouping that is otherwise comprised of Ae. bromeliae. A hierarchical AMOVA showed there is more genetic variation within (42.83%) and between populations (55.18%) than between species (1.99%) at the COI gene.
Red labels represent Ae. bromeliae while blue labels are Ae. lilii. Sequences from Le Goff et al. [ 43 ] (^) are annotated. Bootstrap support values for branches above 55% are shown.
Based on fixed differences between species in the ITS sequences, species specific primers were designed to amplify PCR products in Ae. bromeliae or Ae. lilii. Primers designed to amplify in Ae. bromeliae did not amplify a PCR product from Ae. lilii and vice versa ( Fig 6). Application of this method to 110 specimens of Ae. simpsoni s.l. positively identified all individuals as either Ae. bromeliae or Ae. lilii, as detailed in Table 1 . Both species use the same breeding habitats in domestic and peridomestic habitats including the leaf axils of Musa spp., Colocasia spp. and Dracena spp. Despite focussed collection attempts, neither species was found utilising tree holes for immature development. Both species occur in sympatry in Uganda, while only Ae. lilii was collected from the Republic of Benin and Ae. bromeliae was the only species detected in Tanzania ( Fig 1).
PCR products were run with Hyperladder IV (HyIV) and a negative control (NgC).
| Country | Village/town | Ae. bromeliae | Ae. lilli | |
|---|---|---|---|---|
| Tanzania | Mlimba | 5 | 0 | |
| Udagaji village | 19 | 0 | ||
| Chita | 1 | 0 | ||
| Mahenge | 10 | 0 | ||
| Morningside | 1 | 0 | ||
| Total | 36 | 0 | ||
| Benin | Pobe | 0 | 14 | |
| Niaouili Village | 0 | 10 | ||
| Total | 0 | 24 | ||
| Uganda | Kanyawara | 1 | 14 | |
| Bundibugyo | 14 | 6 | ||
| Kapchorwa | 0 | 4 | ||
| Najjembe | 5 | 6 | ||
| Total | 20 | 30 |
Here we used multiple markers to confirm there are two closely related species of the Simpsoni Complex that occur both allopatrically and sympatrically across the sampled range in sub-Saharan Africa. These species correspond to Ae. bromeliae and Ae. lilii of the Simpsoni Complex as characterised by Mukwaya et al. [ 22 , 27 ]. We infer that a third forest taxon originally reported as Ae. lilii in Walter et al. [ 28 ] relates to another species, possibly a another member of the Simpsoni Group. The development of a molecular species identification method for the YFV vector Ae. bromeliae and non-vector Ae. lilii using variation at the ITS region allowed us to make for the first time reliable inferences about mosquito ecology and distribution. Such information is vital for a complete understanding of disease transmission by these species.
All the Ae. simpsoni s.l. mosquitoes tested fell into one or other of two genetically divergent ITS lineages. Based on sequence similarity these correspond directly to the two lineages found by Mukwaya et al. [ 27 ] and we follow their previous designation as Ae. bromeliae and Ae. lilii. We consider their species designation to be reliable as it was based on both morphology and feeding preference. Firstly, even though some individuals could not be identified due to overlap in morphological characters, Mukwaya et al. [ 27 ] were able to identify some individual specimens from the defining characters of leg tarsomere banding pattern and claw morphology described by Huang [ 22 , 23 ]. Secondly, Mukwaya et al. [ 27 ] identified species based upon host feeding behaviour; human landing catches were used to distinguish anthropophilic Ae. bromeliae from non-anthropophilic Ae. lilii which were collected as larvae where human biting members of the Simpsoni Complex were absent. We confirm the existence of Ae. bromeliae and Ae. lilii as two distinct species since, using ITS sequence variation, they remain genetically distinct in sympatry at several locations in Uganda.
Our study enabled us to link the work of Mukwaya et al. [ 27 ] with that of Walter et al. [ 28 ]. Whereas Muwakya et al. [ 27 ] focused on feeding behaviour, Walter et al. [ 28 ] found divergence between forest and peridomestic populations in coastal Kenya at three nuclear genes. Our phylogenetic analysis reveals that the genetic variation in the ITS region does not correspond to the two distinct lineages found in Walter et al. [ 28 ]. Both Ae. bromeliae and Ae. lilii fall into a single phylogenetic clade at nuclear genes apoLp2 (apolipophorin 2) and CYPJ92 (cytochrome p450). This is consistent with these more slowly evolving nuclear genes being unable to resolve recently diverged species due to incomplete lineage sorting [ 56 ]. The forest lineage from Kenya detected by nuclear genes apoLp2 and CYPJ92 is clearly distinct and has no shared ancestral polymorphism with either Ae. bromeliae or Ae. lilii indicating a more distant relationship. Morphological identification of mosquito species is notoriously difficult and there are many species within the wider Simpsoni Group which share morphological characteristics [ 26 ]. We therefore suggest that the forest species reported by Walter et al. [ 28 ] represents a member of the wider Simpsoni Group, rather than the Simpsoni Complex. Possible candidates for this species that are morphologically similar and known from Kenya include Ae. gandaensis, Ae. woodi, Ae. subargenteus and Ae. sampi [ 26 ].
The mitochondrial COI species barcode marker was also unable to distinguish these taxa with most genetic variation occurring between geographic populations rather than between species. Whilst this inability of COI to distinguish Ae. bromeliae and Ae. lilii could be due to incomplete lineage sorting, the phylogeny suggests it is most likely due to mtDNA introgression; there are three cases in which Ae. bromeliae clusters with Ae. lilii or vice versa, two of which occur in sympatry in Uganda. Mitochondrial introgression is fairly common between closely related mosquito taxa [ 33 ]. Given this, and that Aedine mosquitoes may be particularly prone to NUMT’s due to their large genome size [ 57 ] it would seem wise to not rely on mitochondrial markers to distinguish species in this genus without prior confirmation from an additional marker. We found that the ITS region was the only marker able to reliably separate species of the Simpsoni Complex and therefore used it to develop a PCR mediated species identification method. This method is widely applicable across the range of Ae. bromeliae and Ae. lilii except in southern Africa. For use in this region, the method should be modified to accommodate the presence of Ae. simpsoni s.s. Our PCR mediated species diagnostic method removes the difficulties imposed by morphological identification of field specimens and therefore provides a valuable asset to medical entomologists studying arbovirus transmission.
Our identification tool in conjunction with larval sampling from natural habitats has provided further information on the ecology of the Simpsoni Complex. Contrary to the wide range of plant species utilised by Ae. bromeliae as breeding habitats, the immatures of Ae. lilii have only been reported to date from the axils of Sansevieria spp., suggesting a narrower range of immature habitats [ 22 , 23 ]. However, we found that Ae. lilii, like Ae. bromeliae, utilised the plant axils of Musa spp., Colocasia spp. and Dracena spp., suggesting that larval breeding sites are not as restricted as previously reported [ 22 , 23 ]. Walter et al. [ 28 ] hypothesised that selective pressure for use of domestic larval habitats may have driven speciation in the Simpsoni Complex. This is not supported by our findings that showed no obvious differences in larval habitat between species, although our characterisation of larval habitats was not exhaustive. As host choice appears to be an ecological difference between these species, it is possible that divergent selection for anthropophily, and the reliable blood source it provides, could have driven species divergence [ 28 , 58 ].
We have shown here that Ae. bromeliae and Ae. lilii are common in Uganda where they can be found in sympatry in peridomestic habitats. Ae. lilii was the only member of the Simpsoni Complex collected from the Republic of Benin where it is described for the first time, while only Ae. bromeliae was collected in Tanzania. In addition, Mukwaya et al.’s [ 27 ] sequence data shows that mosquitoes from Nigeria are Ae. lilii whereas those from Kenya are Ae. bromeliae. Molecular evidence therefore agrees with the earlier morphological data of Huang [ 22 , 23 ] that Ae. bromeliae is prevalent in East Africa (Tanzania, Kenya, Uganda, Mayotte; Fig 1). This morphological data also indicates a wider distribution across sub-Saharan Africa for Ae. bromeliae ( Fig 1) [ 22 , 23 ], but it would be wise to confirm this using molecular identification. However, given that Ae. bromeliae tends to be readily collected when it is present, the molecular data suggest that only Ae. lilii is present in West Africa ( Fig 1). This difference would be consistent with the early reports that West African populations are predominantly non-anthropophilic [ 59 ]. Based on our preliminary distribution data, Ae. bromeliae may be an important vector of yellow fever in East Africa where YFV has been isolated from the Simpsoni Complex previously and implemented in disease epidemics including the Ethiopian outbreak of 1960–61 [ 59 – 61 ]. In comparison, other mosquito vectors including Ae. aegypti, Ae. luteocephalus, Ae. furcifer and Ae. taylori may be more important for YFV disease transmission in West Africa where yellow fever has been isolated from these vectors and implicated in outbreaks [ 59 , 62 – 65 ].
Emerging/re-emerging arboviruses such as Chikungunya and Zika are causing great concern since they are increasingly responsible for catastrophic epidemics worldwide [ 66 – 72 ]. Members of the Simpsoni Group transmit a range of arboviruses including yellow fever, Babanki and Ngari viruses [ 1 ]. A limited number of arboviral studies have focused on disease transmission in this species group, progress of which is hampered by incomplete taxonomic understanding. There is therefore a great need to resolve the molecular systematics of the wider Simpsoni Group that should in turn be used to develop methods of species identification. In addition to studies of arboviral risk, assessment of mosquito species ranges is required in order to relate these to differences in the distribution of arboviruses and/or arboviral genotypes such as that observed for yellow fever [ 59 ]. Improved identification methods would also facilitate studies on feeding behaviour and genetic introgression which are important for understanding the risks of zoonotic disease emergence. For example, the introgression we observed between Ae. lilii and Ae. bromeliae could increase the propensity of Ae. lilii to feed on humans, resulting in the increased transfer of zoonotic disease into humans. A precedent for this can be seen in Culex pipiens in North America where mixing of the molestus and pipiens genetic forms (predominantly human and bird feeding, respectively) increases transmission of West Nile virus to humans. A wide range of inferences on ecology and epidemiology can now be made with our molecular identification tool that should be used to assess disease risk and provide basic information for vector population control.
(XLSX)
Click here for additional data file.
The authors are grateful to field teams at the Uganda Virus Research Institute and International Institute for Tropical Agriculture for their research assistance.


Mobile phones banned in Punjab educational institutions
by Muhammad IlyasLAHORE: The Punjab government has devised a new strategy to curb drug abuse by imposing a complete ban on use of mobile phones and other sources to connect to social media at all educational institutions across the province.
All private and public sector schools, colleges and universities throughout the province are directed to strictly impose the ban and treat it as ‘most urgent’ to safe guard the youth from drug abuse, according to a notification issued by Punjab’s Director Public Instruction Zulfkar Ali Bhatti on Friday.
All children under the age of 16 are also barred from using social media, it added.
The decision was taken owing to rising use of drugs among students, who connect to drug peddlers through various social media platforms.
Due to drug abuse, incidents of students being found unconscious are on the rise in Lahore. Deaths have also been reported in this connection.
With Pakistan a gateway to the world for around 75 per cent of all drugs produced around the globe in neighbouring Afghanistan, the country is slowing turning into a major consumer for drugs.
Lahore District Inspector General (DIG) Operations Ashfaq Ahmed Khan, while speaking to The Express Tribune, said that police have already launched a crackdown on drug peddlers, operating in the vicinity of educational institutions. “Over 900 suspects have been arrested in the last three months,” he added.
DIG Khan said that efforts are afoot to curb drug supply to the educational institutions and the police have increased surveillance around the schools and colleges in this connection.
The Lahore DIG Operations had earlier stated that the force has been implementing a zero tolerance policy against criminals, particularly drug peddlers targeting the youth.
Lahore police have also arranged awareness campaigns and lectures at different educational institutions across the provincial capital to inform students about the fatal effects of using drugs. He maintained that the basic purpose of delivering awareness lectures is to make students and teachers aware of their basic responsibilities.


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REPORT OF THE IAU/IAG WORKING GROUP ON
CARTOGRAPHIC COORDINATES AND ROTATIONAL ELEMENTS OF
THE PLANETS AND SATELLITES: 2000
P.K. SEIDELMANN (CHAIR)
U.S. Naval Observatory, Washington, DC, U.S.A.
V.K. ABALAKIN
Institute for Theoretical Astronomy, St. Petersburg, Russia
M. BURSA
Astronomical Institute, Prague, Czech Republic
M.E. DAVIES
RAND, Santa Monica, CA, U.S.A.
C. de BERGH
Observatoire de Paris, Paris, France
J.H. LIESKE
Jet Propulsion Laboratory, Pasadena, CA, U.S.A.
J. OBERST
DLR Berlin Adlershof, Berlin, Germany
J.L. SIMON
Institut de Mecanique Celeste, Paris, France
E.M. STANDISH
Jet Propulsion Laboratory, Pasadena, Ca, USA
P. STOOKE
Univ. of Western Ontario, London, Canada
P.C. THOMAS
Cornell Univ., Ithaca, NY, USA
Abstract.
Every
three years the IAU/IAG Working Group on Cartographic Coordinates
and Rotational Elements of the Planets and Satellites revises tables
giving the directions of the north poles of rotation and the prime
meridians of the planets, satellites, and asteroids. Also presented
are revised tables giving their sizes and shapes. Changes since the
previous report are summarized in the Appendix.
Key words: Cartographic
coordinates, rotation axes, rotation periods, sizes, shapes
1.
Introduction
The
IAU Working Group on Cartographic Coordinates and Rotational
Elements of the Planets and Satellites was established as a
consequence of resolutions adopted by Commissions 4 and 16 at the
IAU General Assembly at Grenoble in 1976. The first report of the
Working Group was presented to the General Assembly at Montreal in
1979 and published in the Trans.
IAU 17B, 72-79, 1980.
The report with appendices was published in Celestial
Mechanics 22, 205-230,
1980. The guiding principles and conventions that were adopted by
the Group and the rationale for their acceptance were presented in
that report and its appendices will not be reviewed here. The second
report of the Working Group was presented to the General Assembly at
Patras in 1982 and published in the Trans.
IAU 18B, 15 1 162, 1983, and
also in Celestial Mechanics 29,
309-321, 1983. The third report on the Working Group was
presented to the General Assembly at New Delhi in 1985 and published
in Celestial Mechanics 39,
103-113, 1986. The fourth report of the Working Group was
presented to the General Assembly at Baltimore in 1988 and was
published in Celestial
Mechanics and Dynamical Astronomy 46,187-204, 1989. The fifth
report of the Working Group was presented to the General Assembly at
Buenos Aires in 1991 and was published in Celestial
Mechanics and Dynamical Astronomy
53, 377-397, 1992.
The sixth report of the Working Group was presented to the General
Assembly at the Hague in 1994 and was published in Celestial
Mechanics and Dynamical Astronomy 63,
127-148, 1996. The seventh report of the Working Group was presented
to the General Assembly at Kyoto, but the changes were sufficiently
minor that the report was not published.
In
1984 the International Association of Geodesy (IAG) and the
Committee on Space Research (COSPAR) expressed interest in the
activities of the Working Group, and after reviewing alternatives,
the Executive Committees of all three organizations decided to
jointly sponsor the Working Group. In 1998 COSPAR informed the
Working Group that, while the reports and expertise of the Working
Group are appreciated, the Working Group does not follow the
scientific structure of COSPAR and they wish to terminate the formal
affiliation.
This
report incorporates revisions to the tables giving the directions of
the north poles of rotation and the prime meridians of the planets
and satellites since the last report. Also, tables giving the sizes
and shapes of the planets, satellites, and asteroids are presented.
2. Definition of
Rotational Elements
Planetary
coordinate systems are defined relative to their mean axis of
rotation and various definitions of longitude depending on the body.
The longitude systems of most of those bodies with observable rigid
surfaces have been defined by references to a surface feature such
as a crater. Approximate expressions for these rotational elements
with respect to the J2000 inertial coordinate system have been
derived. The J2000 coordinate system is defined by the FK5 star
catalog and has the standard epoch of 2000 January 1.5 (JD
2451545.0), TCB. The variable quantities are expressed in units of
days (86400 SI seconds) or Julian centuries of 36525 days.
The
north pole is that pole of rotation that lies on the north side of
the invariable plane of the solar system. The direction of the north
pole is specified by the value of its right ascension a0
and declination d0,
whereas the location of the prime meridian is specified by the angle
that is measured along the planet's equator in an easterly direction with respect to the
planet's north pole from the node Q
(located at right ascension 90°
+ a0)
of the planet's equator on the standard equator to the point B
where the prime meridian crosses the planet's equator. The right
ascension of the point Q
is 90°
+ a0
and the inclination of the planet's equator to the standard equator
is 90° - d0.
Because the prime meridian is assumed to rotate uniformly with the
planet, W accordingly
varies linearly with time. In addition, a0,
d0,
and W may vary with time
due to a precession of the axis of rotation of the planet (or
satellite). If W increases
with time, the planet has a direct
(or prograde) rotation and if W
decreases with time, the rotation is said to be retrograde.
In
the absence of other information, the axis of rotation is assumed to
be normal to the mean orbital plane; Mercury and most of the
satellites are in this category. For many of the satellites, it is
assumed that the rotation rate is equal to the mean orbital period.
The
angle W specifies the
ephemeris position of the prime meridian, and for planets or
satellites without any accurately observable fixed surface features,
the adopted expression for W defines
the prime meridian and is not subject to correction. Where possible,
however, the cartographic position of the prime meridian is defined
by a suitable observable feature, and so the constants in the
expression W = W0 +
Wd, where d is the interval in days from the standard epoch, are chosen so
that the ephemeris position follows the motion of the cartographic
position as closely as possible; in these cases the expression for W may require emendation in the future.
Recommended
values of the constants in the expressions for a0,d0, and
W, in standard equatorial
coordinates with equinox J2000 at epoch J2000, are given for the
planets, satellites, and asteroids in Tables I, II, and III. In
general, these expressions should be accurate to one-tenth of a
degree; however, two decimal places are given to assure consistency
when changing coordinates systems. Zeros are added to rate values (W) for computational consistency and are not an indication of
significant accuracy. Additional decimal places are given in the
expressions for the Moon, Mars, Saturn, and Uranus, reflecting the
greater confidence in their accuracy. Expressions for the Sun and
Earth are given to a similar precision as those of the other bodies
of the solar system and are for comparative purposes only. The
recommended coordinate system for the Moon is the mean Earth/polar
axis system (in contrast to the principal axis system).
3. Definition of
Cartographic Coordinate Systems
In
mathematical and geodetic terminology, the terms 'latitude' and
'longitude' refer to a right-hand spherical coordinate system in
which latitude is defined as the angle between a vector and the
equator, and longitude is the angle between the vector and the plane
of the prime meridian measured in an eastern direction. This
coordinate system, together with Cartesian coordinates, is used in
most planetary computations, and is sometimes called the
planetocentric coordinate system. The origin is the center of mass.
Because of
astronomical tradition, planetographic coordinates (those used on
maps) may or may not be identical with traditional spherical
coordinates. Planetographic coordinates are defined by guiding
principles contained in a resolution passed at the fourteenth
General Assembly of the IAU in 1970. These guiding principles state
that:
(1)
The rotational pole of a planet or satellite which lies on
the north side of the invariable plane will be called north, and
northern latitudes will be designated as positive.
(2)
The planetographic longitude of the central meridian, as
observed from a direction fixed with respect to an inertial system,
will increase with time. The range of longitudes shall extend from 0°
to 360°.
Thus,
west longitudes (i.e., longitudes measured positively to the west)
will be used when the rotation is prograde and east longitudes
(i.e., longitudes measured positively to the east) when the rotation
is retrograde. The origin is the center of mass. Also because of
tradition, the Earth, Sun, and Moon do not conform with this
definition. Their rotations are prograde and longitudes run both
east and west 180°
instead of the usual 360°.
Latitude
is measured north and south of the equator; north latitudes are
designated as positive. The planetographic latitude of a point on
the reference surface is the angle between the equatorial plane and
the normal to the reference surface at the point. In the
planetographic system, the position of a point (P) not on the reference surface is specified by the planetographic
latitude of the point (P')
on the reference surface at which the normal passes through P and by the height (h) of
P above P'.
The
reference surfaces for some planets (such as Earth and Mars) are
ellipsoids of revolution for which the radius at the equator (A)
is larger than the polar semiaxis (C).
Calculations
of the hydrostatic shapes of some of the satellites (Io, Mimas,
Enceladus, and Miranda) indicate that their reference surfaces
should be triaxial ellipsoids. Triaxial ellipsoids would render many
computations more complicated, especially those related to map
projections. Many projections would loose their elegant and popular
properties. For this reason spherical reference surfaces are
frequently used in mapping programs.
Many
small bodies of the solar system (satellites, asteroids, and comet
nuclei) have very irregular shapes. Sometimes spherical reference
surfaces are used for computational convenience, but this approach
does not preserve the area or shape characteristics of common map
projections. Orthographic projections often are adopted for
cartographic portrayal as these preserve the irregular appearance of
the body without artificial distortion.
With
the introduction of large mass storage to computer systems, digital
cartography has become increasingly popular. These databases are
important to irregularly shaped bodies and other bodies where the
surface can be described by a file containing planetographic
longitude, latitude, and radius for each pixel. In this case the
reference sphere has shrunk to a point. Other parameters such as
brightness, gravity, etc., if known, can be associated with each
pixel. With proper programming, pictorial and projected views of the
body can then be displayed by introducing a suitable reference
surface.
Table
IV contains data on the size and shapes of the planets. The first
column gives the mean radius of the body (i.e., the radius of a
sphere of approximately the same volume as the spheroid). The
standard errors of the mean radii are indications of the accuracy of
determination of these parameters due to inaccuracies of the
observational data. Because the shape of a rotating body in
hydrostatic equilibrium is approximately a spheroid, this is
frequently a good approximation to the shape of planets, and so the
second and third columns give equatorial and polar radii for
'best-fit' spheroids. The origin of these coordinates is the
center-of-mass with the polar axis coincident with the spin axis.
The fourth column is the root-mean-square (RMS) of the radii
residuals from the spheroid and is an indication of the variations
of the surface from the spheroid due to topography. The last two
columns give the maximum positive and negative residuals to bracket
the spread.
Table
V contains data on the size and shape of the satellites. The first
column gives the mean radius of the body. The standard errors of the
mean radii are indications of the accuracy of determination of these
parameters due to inaccuracies of the observational data. Because
the hydrostatic shape of a body in synchronous rotation about a
larger body is approximately an ellipsoid, that shape has been
selected to describe the shape of the satellites. The next three
columns (2-4) give the axes of the best-fit ellipsoids in the order
equatorial subplanetary, equatorial along orbit, and polar. The
origin of these coordinates is the center-of-mass with the polar
axis coincident with the spin axis. The fifth column is the RMS of
the radii residuals from the ellipsoid and is an indication of the
variations of the surface from the ellipsoid due to topography. The
last two columns give the maximum positive and negative residuals to
bracket the spread.
Table
I. Recommended values for the direction of the north pole of
rotation and the prime meridian of the Sun and planets (2000)
a0,d0
are standard equatorial coordinates with equinox J2000 at epoch J2000.
Approximate
coordinates of the north pole of the invariable plane are a0
= 273 °.85,
d0=
66°.99.
T =
interval in Julian centuries (of 36525 days) from the
standard epoch
d =
interval in days from the standard epoch.
The
standard epoch is 2000 January 1.5, i.e., JD 2451545.0 TCB.
| Sun
|
a0
|
=
|
286°. 13
|
|
|
d0
|
= |
63°.87
|
|
|
W |
=
|
84°.10 + 14°.1844000d
|
|
|
|
| Mercury
|
a0
|
=
|
281.01 - 0.033T
|
|
|
d0 |
=
|
61.45 - 0.005T
|
|
|
W
|
=
|
329.548 + 6.1385025d
|
(a)
|
|
| |
| Venus
|
a0 |
=
|
272.76
|
|
|
d0 |
=
|
67.16
|
|
|
W
|
=
|
160.20 - 1.4813688d
|
(b)
|
|
|
| Earth |
a0
|
=
|
0.00 - 0.641T
|
|
|
d0 |
=
|
90.00 - 0.557T
|
|
|
W
|
=
|
190.16 +<PHONE_NUMBER>d
|
(c)
|
|
|
| Mars |
a0
|
=
|
317.68143 - 0.1061T
|
|
|
d0 |
=
|
52.88650 - 0.0609T
|
|
|
W
|
=
|
176.753 +<PHONE_NUMBER>6d
|
(d)
|
|
|
| Jupiter
|
a0
|
=
|
268.05 - 0.009T |
|
|
d0 |
=
|
64.49 + 0.003T
|
|
|
W
|
=
|
284.95 +<PHONE_NUMBER>d
|
(e)
|
|
|
| Saturn
|
a0
|
=
|
40.589 - 0.036T
|
|
|
d0 |
=
|
83.537 - 0.004T
|
|
|
W
|
=
|
38.90 +<PHONE_NUMBER>d
|
(e)
|
|
|
| Uranus
|
a0
|
=
|
257.311
|
|
|
d0 |
=
|
-15.175
|
|
|
W
|
=
|
203.81
-<PHONE_NUMBER>d
|
(e)
|
|
|
|
|
|
|
|
|
|
|
| Neptune |
a0
|
=
|
299.36 + 0.70 sin N
|
|
|
d0 |
=
|
43.46
- 0.51 cos N
|
|
|
W
|
=
|
253.18
+<PHONE_NUMBER>d-0.48sin
N
|
(e)
|
|
N
|
=
|
357.85 + 52.316T
|
|
|
|
|
|
|
|
|
|
|
|
| Pluto
|
a0
|
=
|
313.02
|
|
|
d0 |
=
|
9.09
|
|
|
W
|
=
|
236.77 - 56.3623195d
|
(f)
|
(a) The 20°
meridian is defined by the crater Hun Kal.
(b) The 0°
meridian is defined by the central peak in the crater Ariadne.
(c)
The expression for W might be in error by as much as 0°.2
because of uncertainty in the length of the UT day and the TT UT on
1 January 2000.
(d)
The 0°
meridian is defined by the crater Airy-0.
(e)
The equations for W for Jupiter, Saturn, Uranus and Neptune refer to the rotation of
their magnetic fields (System III). On Jupiter, System I (WI = 67 °.1
+ 877°.900d)
refers to the mean atmospheric equatorial rotation; System II (WII
= 43°.3
+ 870°.270d) refers to the mean atmospheric rotation north of the south
component of the north equatorial belt, and south of the north
component of the south equatorial belt.
(f) The 0°
meridian is defined as the mean sub-Charon meridian.
Table
II. Recommended values for the direction of the north pole of
rotation and the prime meridian of the satellites (2000)
a0, d0, T,
and d have the same
meanings as in Table I (epoch 2000 January 1.5, i.e., JD 2451545.0
TCB).
Earth:
Moon
a0 = 269°.9949
+ 0°.0031T -
3°.8787sin El
- 0°.1204 sin E2
+ 0.0700 sin E3 - 0.0172 sin E4
+ 0.0072 sin E6
- 0.0052sin El0 + 0.0043sin E13
d0 = 66.5392
+ 0.0130T
+ 1.5419 cos E1 +
0.0239 cos E2
- 0.0278 cos E3 + 0.0068 cos E4
- 0.0029 cos E6
+ 0.0009 cos E7 + 0.0008 cos E10
- 0.0009cos E13
W = 38.3213
+ 13.17635815 d
- 1.4 x 10-12 d2
+ 3.5610 sin E1
+ 0. 1208 sin E2 - 0.0642 sin E3
+ 0.0158 sin E4
+ 0.0252 sin E5 - 0.0066 sin E6
- 0.0047 sin E7
- 0.0046 sin E8 + 0.0028 sin E9
+ 0.0052 sin E 10
+ 0.0040sin E11 + 0.0019 sin E12
- 0.0044 sin E l3
where
El = 125°.045
- 0°.0529921d,
E2 = 250°.089
- 0°.1059842d,
E3 = 260°.008
+ 13°.0120009d,
E4 = 176.625 +
13.3407154d,
E5 = 357.529 +
0.9856003d,
E6 = 311.589 + 26.4057084d,
E7 = 134.963 +
13.0649930d,
E8 = 276.617 +
0.3287146d,
E9 = 34.226+ 1.7484877d,
E10
= 15.134 - 0.1589763d,
E11 = 119.743 +
0.0036096d,
E12 = 239.961 +
0.1643573d,
E13
= 25.053 + 12.9590088d
Mars: I Phobos
a0 = 317.68
- 0.108T
+ 1.79 sin Ml
d0 = 52.90
- 0.061T
- 1.08 cos M1
W = 35.06
+ 1128.8445850d
+ 8.864T 2
-
1.42 sin M1
- 0.78 sin M2
II Deimos
a0 = 316.65
- 0.108T
+ 2.98 sin M3
d0 = 53.52
- 0.061T
- 1.78 cos M3
W = 79.41
+<PHONE_NUMBER>d
- 0.520T2
- 2.58 sin M3 + 0.19
cosM3
where
M1
= 169°.51 - 0°.4357640d,
M2 = 192°.93
+ 1128 °.4096700d
+ 8°.864T2,
M3
= 53°.47 - 0°.0181510d
Jupiter: XVI
Metis
a0 = 268.05
- 0.009T
d0 =
64.49
+ 0.003T
W = 346.09 +
1221.2547301d
XV
Adrastea
a0 = 268.05
- 0.009T
d0 = 64.49
+ 0.003T
W = 33.29
+ 1206.9986602d
V
Amalthea a0 = 268.05
- 0.009T
- 0.84 sin J1
+ 0.01 sin 2J1
d0 = 64.49
+ 0.003T
- 0.36 cos J1
W =
231.67
+<PHONE_NUMBER>d
+ 0.76 sin J1 - 0.01
sin 2J1
XIV Thebe
a0 =
268.05
- 0.009T
- 2.11 sin J2
+ 0.04 sin2J2
d0 = 64.49
+ 0.003T
- 0.91 cos J2
+ 0.01 cos 2J2
W =
8.56
+<PHONE_NUMBER>d
+ 1.91 sin J2 - 0.04
sin 2J2
I
Io
a0
= 268.05
- 0.009T
+ 0.094 sin J3
+ 0.024 sin J4
d0 = 64.50
+ 0.003T
+ 0.040 cos J3
+ 0.011 cos J4
W = 200.39
+<PHONE_NUMBER>d - 0.085 sin J3
- 0. 022 sin J4
II
Europa
a0
= 268.08
- 0.009T
+ 1.086 sin J4
+ 0.060 sin J5
+ 0.015 sin J6
+ 0. 009 sin J7
d0 = 64.51
+ 0.003T
+ 0.468 cos J4
+ 0.026 cos J5
+ 0.007 cos J6
+ 0.002 cos J7
W =
35.67
+<PHONE_NUMBER>d
- 0.980 sin J4 - 0.054 sin J5
- 0.014 sin J6 -
0.008 sin J7
(a)
III
Ganymede a0
= 268.20
- 0.009T
- 0.037 sin J4
+ 0.431 sin J5
+ 0.091 sin J6
d0 =
64.57
+ 0.003T
- 0.016 cos J4
+ 0.186 cos J5
+ 0.039 cos J6
W = 44.04
+ 50.3176081d + 0.033 sin J4
- 0.389 sin J5
(b)
- 0.082 sin J6
IV
Callisto
a0
= 268.72
- 0.009T
- 0.068 sin J5
+ 0.590 sin J6
+ 0.010 sin J8
d0 =
64.83
+ 0.003T
- 0.029 cos J5 + 0.254 cos J6
- 0.004 cos J8
W =
259.73
+ 21.5710715d
+ 0.061 sin J5
- 0.533 sin J6
(c)
-
0.009 sin J8
where
J l = 73°.32
+ 91472°.9T
J 2 = 24°.62 + 45137°.2T,
J 3 = 283°.90 + 4850°.7T,
J 4
= 355.80 + 1191.3T,
J 5 = 119.90 + 262.1T,
J6 = 229.80 + 64.3T,
J 7 = 352.25 +
2382.6T,
J 8 =113.35 +
6070.0T
Saturn: XVIII
Pan
a 0 = 40.6
- 0.036T
d0 = 83.5
- 0.004T
W = 48.8
+<PHONE_NUMBER>d
XV
Atlas
a0 = 40.58
- 0.036T
d0 = 83.53
- 0.004T
W = 137.88
+<PHONE_NUMBER>d
XVI Prometheus
a0 = 40.58
- 0.036T
d0 = 83.53
- 0.004T
W = 296.14 +
587.289000d
XVII Pandora
a0 = 40.58
- 0.036T
d0 = 83.53
- 0.004T
W = 162.92
+<PHONE_NUMBER>d
XI
Epimetheus a0 = 40.58
- 0.036T
- 3.153 sin S1
+ 0.086 sin 2Sl
d0 = 83.52
- 0.004T
- 0.356 cos S1 + 0.005 cos 2SI
W = 293.87 +
518.4907239d
+ 3.133 sin S1 - 0.086
sin 2S1
(j)
X
Janus
a0 = 40.58
- 0.036T
- 1.623 sin S2 + 0.023 sin 2S2
d0 = 83.52
- 0.004T
- 0. 183 cos S2 + 0.001 cos 2S2
W =
58.83
+<PHONE_NUMBER>d
+ 1.613 sin S2 - 0.023 sin 2S2
(j)
I Mimas
a0 = 40.66
- 0.036T
+ 13.56 sin S3
d0 = 83.52
- 0.004T
- 1.53 cos S3
W = 337.46 +
381.9945550d
- 13.48 sin S3
- 44.85 sin S5
(d)
II
Enceladus a0 =
40.66
- 0.036T
d0 = 83.52
- 0.004T
W = 2.82
+<PHONE_NUMBER>d
(e)
III
Tethys
a0 = 40.66
- 0.036T
+ 9.66 sin S4
d0 = 83.52
- 0.004T
- 1.09 cos S4
W = 10.45
+<PHONE_NUMBER>d
- 9.60 sin S4
+ 2.23 sin S5
(f)
XIII Telesto
a0 = 50.51
- 0.036T
d0 = 84.06
- 0.004T
W = 56.88
+<PHONE_NUMBER>d
(j)
XIV Calypso a0 = 36.41
- 0.036T
d0 = 85.04
- 0.004T
W = 153.51 +
190.6742373d
(j)
IV
Dione
a0 = 40.66
- 0.036T
d0 = 83.52
- 0.004T
W = 357.00
+<PHONE_NUMBER>d
(g)
XII Helene
a0 = 40.85
- 0.036T
d0 = 83.34
- 0.004T
W = 245.12 +
131.6174056d
V
Rhea
a0 = 40.38
- 0.036T
+ 3. 10 sin S6
d0 = 83.55
- 0.004T
- 0.35 cos S6
W = 235.16 +
79.6900478d
- 3.08 sin S6
(h)
VI Titan
a0 =
36.41
- 0.036T
+ 2.66 sin S7
d0 = 83.94
- 0.004T
- 0. 30 cos S7
W = 189.64 +
22.5769768d
- 2.64 sin S7
VIII Iapetus
a0 = 318.16
- 3.949T
d0 = 75.03
- 1.143T
W
= 350.20
+ 4.5379572d
IX
Phoebe
a0 = 355.00
d0
= 68.70
W = 304.70
+<PHONE_NUMBER>d
where Sl
= 353°.32 + 75706°.7T, S2
= 28°.72 + 75706°.7T, S3
= 177°.40 - 36505°.5T
S4 = 300.00 -
7225.9T,
S5 =
316.45 + 506.2T,
S6
= 345.20 - 1016.3T,
S7 = 29.80 - 52.1T
Uranus: VI Cordelia
a0 = 257.31
- 0. 15 sin U1
d0 = -15.18
+ 0.14 cos U 1
W = 127.69
- 1074.5205730d - 0.04 sinUl
VII
Ophelia
a0 =
257.31
- 0.09 sin U2
d0 = -
15.18
+ 0.09 cos U2
W = 130.35
-<PHONE_NUMBER>d
- 0.03 sin U2
VIII Bianca
a0
= 257.31
- 0.16 sin U3
d0 = -15.18
+ 0. 16 cos U3
W = 105.46
-<PHONE_NUMBER>d - 0.04 sin U3
IX
Cressida
a0
= 257.31
- 0.04 sin U4
d0 = - 15.18
+ 0. 04 cos U4
W = 59.16
-<PHONE_NUMBER>d - 0.01 sin U4
X
Desdemona a0
=
257.31
- 0. 17 sin U5
d0 = -15.18
+ 0. 16 cos U5
W = 95.08
-<PHONE_NUMBER>d - 0.04 sin U5
XI
Juliet
a0
= 257.31
- 0.06 sin U6
d0 = - 15.18
+ 0.06 cos U6
W = 302.56
-<PHONE_NUMBER>d - 0.02 sin U6
XII
Portia
a0
= 257.31
- 0.09 sin U7
d0 = -
15.18 + 0.09 cos U7
W = 25.03
-<PHONE_NUMBER>d - 0.02 sin U7
XIII Rosalind
a0
= 257.31
- 0.29 sin U8
d0 =
-15.18
+ 0.28 cos U8
W =
314.90
-<PHONE_NUMBER>d -
0.08 sin U8
XIV Belinda
a0
=
257.31
- 0.03 sin U9
d0 = -15.18
+ 0.03 cos U9
W = 297.46
-<PHONE_NUMBER>d -
0.01 sin U9
XV
Puck
a0
= 257.31
- 0.33 sin U10
d0 =
-15.18
+ 0.31 cos U10
W = 91.24
-<PHONE_NUMBER>d -
0.09 sin Ul0
V
Miranda
a0
= 257.43
+ 4.41 sin U11
- 0.04 sin 2U11
d0 =
-15.08
+ 4.25 cos U11
- 0.02 cos 2U11
W = 30.70
-<PHONE_NUMBER>d - 1.27 sin U12
+ 0.15 sin 2Ul2
+
1.15 sin U 11
- 0.09 sin 2U11
I
Ariel
a0
=
257.43
+ 0.29 sin U13
d0 = -15.10
+ 0.28 cos U13
W = 156.22
-<PHONE_NUMBER>d + 0.05 sin U12
+ 0.08 sin U13
II
Umbriel
a0
= 257.43
+ 0.21 sin U14
d0 =
-15.10 + 0.20 cos U14
W = 108.05
- 86.8688923d
- 0.09 sin U12
+ 0.06 sin U14
III
Titania
a0
=
257.43
+ 0.29 sin U15
d0 = -15.10
+
0.28 cos U15
W = 77.74
- 41.3514316d + 0.08 sin U15
IV
Oberon
a0
= 257.43
+ 0.16 sin U16
d0 = -15.10
+ 0.16 cos U16
W = 6.77
- 26.7394932d + 0.04 sin U16
where Ul
= 115°.75 + 54991°.87T, U2
= 141°.69 + 41887°.66T, U3
= 135°.03 + 29927°.35T,
U4 = 61.77 +
25733.59T,
U5
= 249.32 + 24471.46T,
U6 = 43.86 + 22278.41T,
U7 = 77.66 +
20289.42T
U8 = 157.36 +
16652.76T,
U9 = 101.81 +
12872.63T,
U10 = 138.64 +
8061.81T,
U11 = 102.23 - 2024.22T,
U12 = 316.41 + 2863.96T,
U13
= 304.01 - 51.94T,
U14 = 308.71 - 93.17T,
U15 = 340.82 - 75.32T,
U16 =
259.14 - 504.81T
Neptune
III
Naiad
a0
= 299°.36
+ 0°.70
sin N - 6.49 sin N1
+ 0°.25
sin 2Nl
d0 = 43.36 -
0.51 cos N
- 4.75 cos Nl. + 0.09 cos 2Nl
W =
254.06
+ 1222.8441209d - 0.48 sin N
+ 4.40 sin N1 - 0.27 sin 2N1
IV
Thalassa
a0
=
299.36
+ 0.70 sin N
- 0.28 sin N2
d0
=
43.45
- 0.51 cos N
- 0.21 cos N2
W =
102.06
+ 1155.7555612d - 0.48 sin N
+ 0. 19 sin N2
V
Despina
a0
= 299.36
+ 0.70 sin N
- 0.09 sin N3
d0 = 43.45
- 0.51 cos N
- 0.07 cos N3
W = 306.51
+ 1075.7341562d - 0.49 sin N
+ 0.06 sin N3
VI
Galatea
a0
=
299.36
+ 0.70 sin N
- 0.07 sin N4
d0 = 43.43
- 0.51 cos N
- 0.05 cos N4
W =
258.09
+<PHONE_NUMBER>d
- 0.48 sin N +
0.05 sin N4
VII
Larissa
a0
= 299.36
+ 0.70 sin N
- 0.27 sin N5
d0 = 43.41
- 0.51 cos N
- 0.20 cos N5
W =
179.41
+<PHONE_NUMBER>d
- 0.48 sin N +
0. 19 sin N5
VIII Proteus
a0 =
299.27
+ 0.70 sin N
- 0.05 sin N6
d0 = 42.91
- 0.51 cos N
- 0.04 cos N6
W =
93.38
+<PHONE_NUMBER>d
- 0.48 sin N +
0.04 sin N6
I
Triton
a0
= 299.36
- 32.35 sin N7
- 6.28 sin 2N7
- 2.08 sin 3N7
- 0.74 sin 4N7 - 0.28 sin 5N7
- 0.11 sin 6N7
- 0.07 sin 7N7 - 0.02 sin 8N7
- 0.01 sin 9N7
d0 = 41.17
+ 22.55 cos N7
+ 2.10 cos 2N7 + 0.55 cos 3N7
+ 0.16 cos 4N7 + 0.05 cos 5N7
+ 0.02 cos 6N7
+
0.01 cos 7N7
W = 296.53
- 61.2572637d + 22.25 sin N7
+ 6.73 sin 2N7
+ 2.05 sin 3N7 + 0.74 sin 4N7
+ 0.28 sin 5N7
+ 0.11 sin 6N7 + 0.05 sin 7N7
+ 0.02 sin 8N7
+ 0.01 sin 9N7
where
N =
357°.85 + 52.316T,
N l = 323°.92+62606°.6T, N2 = 220°.51+55064°.2T,
N3 = 354.27 + 46564.5T,
N4
= 75.31 + 26109.4T,
N5 = 35.36 + 14325.4T,
N6 = 142.61 + 2824.6T,
N7 = 177.85 +
52.316T
Pluto I
Charon
a0 = 313.02
d0 = 9.09
W = 56.77
- 56.3623195d
(a)
The 182°
meridian is defined by the crater Cilix.
(b)
The 128°
meridian is defined by the crater Anat.
(c)
The 326°
meridian is defined by the crater Saga.
(d)
The 162°
meridian is defined by the crater Palomides.
(e)
The 5°
meridian is defined by the crater Salih.
(f)
The 299°
meridian is defined by the crater Arete.
(g)
The 63°
meridian is defined by the crater Palinurus.
(h)
The 340°
meridian is defined by the crater Tore.
(i)
The 276°
meridian is defined by the crater Almeric.
(j)
These equations are correct for the period of the Voyager
encounters. Because of precession they may not be accurate at other
time periods.
Satellites
for which no suitable data are yet available have been omitted from
this table. Nereid is not included in this table because it is not
in synchronous rotation.
Table
III. Recommended rotation values for the direction of the north pole
of rotation and the prime meridian of selected asteroids (2000)
a0, d0, W,
and d have the
same meanings as in the Table I (epoch 2000 January 1.5, i.e., JD
2451545.0 TCB).
243 Ida
a0 = 348°.76
d0 = 87°.12
W =
265°.95 - 1864°.6280070d
(a)
951 Gaspra
a0 = 9°.47
d0
=
26°.70
W = 83°.67 + 1226°.9114850d
(b)
4 Vesta
a0
= 301°
d0
=
41 °
W = 292°
+ 1617°.332776d
433
Eros
a0 = 11°.9
d0
=
20°.8
W
= 324°.08
+ 1639°.3922d
(a)
The 0°
meridian is defined by the crater Afon.
(b)
The 0°
meridian is defined by the crater Charax.
Table
VI contains data on the size and shape of selected asteroids. The
first column gives the mean radius of the body and an estimate of
the accuracy of this measurement. The next three columns give
estimates of the radii measured along the three principal axes. The
fifth column gives the radii of a best-fit ellipsoid. These are
given because an ellipsoid is a common reference shape for
photometric analyses. The last column gives an estimate of the
maximum deviation of the body from the ellipsoid and is an estimate
of the goodness of fit.
The values of the radii and axes in Tables IV, V, and VI are
derived by various methods and do not always refer to common
definitions. Some use star or spacecraft occultation measurements,
some use limb fitting, others use altimetry measurements from
orbiting spacecraft, and some use control network computations. For
the Earth, the spheroid refers to mean sea level, clearly a very
different definition from other bodies in the Solar System.
The uncertainties in the values for the radii and axes in
Tables IV, V, and VI are generally those of the authors, and, as
such, frequently have different meanings. Sometimes they are
standard errors of a particular data set, sometimes simply an
estimate or expression of confidence.
The radii and axes of the large gaseous planets, Jupiter,
Saturn, Uranus, and Neptune in Table IV refer to a one-bar-pressure
surface.
The radii given in the tables are not necessarily the
appropriate values to be used in dynamical studies; the radius
actually used to derive a value of J2 (for example) should always be
used in conjunction with it.
Table
IV Size and shape parameters of the planets
Planet Mean
radius
Equatorial
Polar radius
RMS Maximum
Maximum
(km)
radius
(km)
deviation
elevation
depression
(km)
from
(km)
(km)
spheroid
(km)
Mercury 2439.7 ± 1.0
same
same
1
4.6
2.5
Venus 6051.8
± 1.0
same
same
1
11
2
Earth
6371.00 ± 0.01
6378.14 ± 0.01
6356.75 ± 0.01 3.57
8.85
11.52
Mars
3389.50 ± 0.003
3396.19 ± 0.16
N 3376.20 ± 0.05 3.3
21.183 ± 0.005
7.825 ± 0.005
S 3382.582 ± 0.05
Jupiter* 69911 ± 6
71492 ± 4
66854 ± 10
62.1 31
102
Saturn* 58232 ± 6
60268 ± 4
54364 ± 10
102.9 8
205
Uranus* 25362 ± 7
25559 ± 4
24973 ± 20
16.8 28
0
Neptune*
24622 ± 19
24764 ± 15
24341 ± 30
8
14
0
Pluto
1195 + 5
same
same
*The
radii correspond to a one-bar surface.
Table
V. Size and shape parameters of the satellites
| Planet
|
Satellite
|
Mean
Radius
(km)
|
Subplanetary
equatorial
radius (km)
|
Along Orbit
equatorial
radius (km)
|
Polar
radius
(km)
|
RMS
deviation
from
ellipsoid
(km)
|
Maximum
elevation
(km)
|
Maximum
Depression
(km)
|
Earth
Moon
1737.4 ± 1 same
same
same 2.5
7.5
5.6
Mars I
Phobos
11.1 ± 0.15 13.4
11.2
9.2
0.5
II
Deimos
6.2 ±
0.18 7.5
6.1
5.2 0.2
Jupiter
XVI Metis
21.5 ±
4 30
20
17
XV Adrastea
8.2 ±
4 10
8
7
V
Amalthea
83.5 ± 3
125
73
64
3.2
XIV Thebe
49.3 ± 4
58
49
42
I
Io
1818.1 ±
0.1 1826.5
1815.7
1812.2 1.4
5-10
3
II
Europa
1561
same
same
same
0.5
III
Ganymede
2634
same
same
same
0.6
IV
Callisto
2408
same
same
same
0.6
XIII Leda
5
VI
Himalia
85 ± 10
X
Lysithea
12
VII Elara
40 ± 10
XII Ananke
10
XI
Carme
15
VIII Pasiphae
18
IX
Sinope
14
Saturn
XVIII Pan
10 ± 3
XV Atlas
16 ± 4
18.5
17.2
13.5
XVI Prometheus
50.1 ± 3
74.0
50.0
34.0
4.1
XVII Pandora
41.9 ± 2
55.0
44.0
31.0
1.3
XI
Epimetheus 59.5
± 3
69.0
55.0
55.0
3.1
X Janus
88.8 ± 4
97.0
95.0
77.0
4.2
I
Mimas
198.6 ± 0.6 209.1 ± 0.5
196.2 ± 0.5
191.4 ± 0.5
0.6
II Enceladus
249.4 ± 0.3
256.3 ± 0.3 247.3 ± 0.3
244.6 ± 0.5
0.4
III
Tethys
529.8 ± 1.5
535.6 ± 1.2
528.2 ± 1.2 525.8 ± 1.2
1.7
XIII Telesto
11± 4
15 ± 2.5
12.5 ± 5 7.5 ±
2.5
XIV Calypso
9.5 ± 4
15.0
8.0
8.0
0.6
IV
Dione
560 ± 5
same
same
same
0.5
XII Helene
16
17.5 ± 2.5
0.7
V
Rhea
764 ±4
same
same
same
VI
Titan
2575 ± 2
same
same
same
VII Hyperion
141.5 ± 20
180 ± 20 140
± 20
112.5 ± 20
7.4
VIII Iapetus
718 ± 8
same
same
same
6.1
12
IX Phoebe
110 ± 10
115 ± 10 110
± 10
105 ±10
2.7
Uranus VI
Cordelia
13 ± 2
VII Ophelia
15 ± 2
VIII Bianca
21 ± 3
IX Cressida
31 ± 4
X
Desdemona 27
± 3
XI
Juliet
42 ± 5
XII Portia
54 ± 6
XIII Rosalind
27 ± 4
XIV Belinda 33 ± 4
XV Puck
77 ± 5
1.9
V
Miranda
235.8 ± 0.7
240.4 ± 0.6
234.2 ± 0.9 232.9 ± 1.2
1.6
5
8
I
Ariel
578.9 ± 0.6 581.1 ± 0.9
577.9 ± 0.6
577.7 ± 1.0
0.9 4
4
II
Umbriel
584.7 ± 2.8
same
same
same
2.6
6
III
Titania
788.9 ± 1.8
same
same
same
1.3
4
IV
Oberon
761.4 ± 2.6
same
same
same
1.5
12
2
Neptune
III
Naiad
29 ± 6
IV
Thalassa
40 ± 8
V
Despina
74 ± 10
VI
Galatea
79 ± 12
VII Larissa
96 ± 7
104
89
2.9
6
5
VIII Proteus
208 ± 8
218
208
201
7.9
18
13
I
Triton
1352.6 ± 2.4
II
Nereid
170 ± 25
Pluto
I
Charon
593 ± 13
Table VI. Size and shape parameters of selected asteroids
Asteroid Mean radius
Radii measured along
Radii of best-fit
Maximum
(km)
principal axes
Ellipsoid
deviation
(km) (km)
(km) (km)
from ellipsoid
(km)
243 Ida
15.65 ± 0.6
26.8 12.0
7.6
30.0, 12.6, 9.3 8.4
951 Gaspra
6.1 ± 0.4
9.1
5.2
4.4
9.1, 5.2, 4.7
2.1
216 Kleopatra
108.5
47
40.5
4.
Appendix
This
appendix summarizes the changes that have been made to the tables
since the 1994 report (Celestial Mechanics and Dynamical Astronomy 63, 127-148, 1996).
In Table I, the new value for the W0
of Mercury was the result of a new control network
computation by Robinson et al. (1999). The new values of a0 and
d0
for
Mars are due to Folkner et al (1997). The new value for W
of Mars was the result of a control network computation by Davies
et al (1999). The value for the d term in W
for Jupiter is from Higgins et al. (1996).
The new value for the d
term in W for Jupiter is a
new radio rotation period by Higgins et al. (1996).
In Table II the value of W
for Metis is from Lieske (1997).
In Table III the value for Vesta are from Thomas et al. (1997)
and the values of Eros are from Thomas et al. (2000).
In Table IV the Mars model is that determined by the Mars
Orbiter Laser Altimeter (MOLA) group from Smith et al. 1999.
In
Table V the sizes of the inner satellites of Jupiter are from Thomas
et al. (1998). The sizes of th Galilean satellites are from Davies et
al (1998).
In
Table VI the parameters for 216 Kleopatia are from Ostro et al 2000.
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D. E., Zuber, M. T., Solomon, S. C., Phillips, R. J., Head, J. W.,
Garvin, J. B., Banerdt, W. B., Muhleman, D. O., Pettengill, G. H.,
Neumann, G. A., Lemoine, F. G., Abshire, J. B., Aharonson, O., Brown,
C. D., Hauck, S. A., Ivanov, A. B., McGovern, P. J., Zwally, H. J.,
and Duxbury, T. C.: 1999, 'The global topography of Mars and
implications for surface evolution ', Science,
284, 1495-1503.
Thomas,
P. C., Veverka, J., Simonelli, D., Helfenstein, P., Carcich, B.,
Belton, M. J. S., Davies, M. E., and Chapman, C.: 1994, 'The shape of
Gaspra', Icarus 107, 23-36.
Thomas,
P. C., Veverka, J., Carcich, B., Belton, M. J. S., Sullivan, R., and
Davies, M. E.: 1996, 'The shape of Ida', Icarus.120,
20-32.
Thomas,
P. C., Binzel, R. P., Gaffey, M. J., Zellner, B. H., Storrs, A. D.,
and Wells, E. 1997 Vesta: Spin, pole, size, and shape from HST
images, Icarus 128,
88-94.
Thomas,
P. C., Burns, J. A., Rossier, L., Simonelli, D., Veverka, J. Chapman,
C., Klaasan, K., Johnson, T. V., and Belton, M. J. S.: 1998 The
small inner satellites of Jupiter Icarus
135, 360-371.
Thomas,
P. C., Joseph, J., Carcich, B., Veverka, J., Miller, J. K., Owen, W.,
Williams, B., Robinson, M.: 2000 The shape of Eros from NEAR
imaging data, Icarus,145,
in press.
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|
Lomonosov concentrations of low-mass halos |
|
According to the standard LambdaCDM cosmological framework, the mass spectrum of cold dark matter halos spans over many orders of magnitude, from the largest virialized structures observed in the Universe with masses above $10^{15}$ M$_\odot$, extending down to very small masses, probably as small as the Earth's, i.e. $10^{-6}$ M$_\odot$, or even less. Halos below $10^7$ M$_\odot$ are most probably not able to host stars or gas at all, thus remaining completely dark. Yet, there is a hope to detect them via gravitational lensing, dark matter particle decay or annihilation, or distortions in stellar velocity distribution. Low-mass halos may also have an important impact on future observations of reionization process. The Lomonosov project is aimed at measuring the concentration parameter, which characterizes the density inside dark matter halos, for halos in the mass range $10^7 - 10^{10}$ M$_\odot$, hardly accessible by typical large-scale simulations. The Lomonosov suite consists of four dark matter simulations run at the Lomonosov supercomputer of the Moscow State University computer center (http://parallel.ru). One simulation is a full box and three other are zoom simulations of overdense, typical and underdense regions. The characteristics of each of these simulations are summarized in the table below. |
| Please click here to access the halo catalogs, merger trees and snapshots. |
|
Acknowledgement
If you use these data, please cite Pilipenko et al. 2017. |
last update, mar. 2017


module BioCReader
Public Instance Methods
read(path, options)
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 7 def read(path, options) collection = nil File.open(path) do |file| collection = read_from_file_or_string(file, options) end collection end
read_annotation(xml, annotation, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 94 def read_annotation(xml, annotation, options = {}) annotation.id = xml["id"] annotation.text = read_text(xml, "text") read_infon(xml, annotation) read_recursive(xml, annotation, "location") true end
read_collection(xml, collection, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 55 def read_collection(xml, collection, options = {}) collection.source = read_text(xml, "source") collection.date = read_text(xml, "date") collection.key = read_text(xml, "key") read_infon(xml, collection) read_recursive(xml, collection, "document", options) end
read_document(xml, document, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 63 def read_document(xml, document, options = {}) document.id = read_text(xml, "id") if options[:documents].kind_of?(Array) && !options[:documents].include?(document.id) return false end read_infon(xml, document) read_recursive(xml, document, "passage") read_recursive(xml, document, "relation") document.adjust_ref true end
read_from_file_or_string(file_or_string, options)
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 16 def read_from_file_or_string(file_or_string, options) collection = nil xml_doc = Nokogiri::XML(file_or_string) do |config| config.noent.strict.noblanks end xml = xml_doc.at_xpath("//collection") if xml.nil? fail 'Wrong format' end collection = SimpleBioC::Collection.new read_collection(xml, collection, options) collection end
read_infon(xml, obj)
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 42 def read_infon(xml, obj) xml.xpath("infon").each{ |i| obj.infons[i["key"]] = i.content} end
read_int(xml, name)
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 37 def read_int(xml, name) val = read_text(xml, name) val && val.to_i end
read_location(xml, location, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 109 def read_location(xml, location, options = {}) location.offset = xml["offset"] location.length = xml["length"] true end
read_node(xml, node, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 115 def read_node(xml, node, options = {}) node.refid = xml["refid"] node.role = xml["role"] true end
read_passage(xml, passage, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 75 def read_passage(xml, passage, options = {}) passage.text = read_text(xml, "text") passage.offset = read_int(xml, "offset") read_infon(xml, passage) read_recursive(xml, passage, "sentence") read_recursive(xml, passage, "annotation") read_recursive(xml, passage, "relation") true end
read_recursive(xml, obj, name, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 46 def read_recursive(xml, obj, name, options = {}) target_class = SimpleBioC.const_get(name.capitalize) xml.xpath(name).each do |node| instance = target_class.new(obj) ret = send(:"read_#{name}", node, instance, options) obj.instance_variable_get(:"@#{name}s") << instance if ret end end
read_relation(xml, relation, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 102 def read_relation(xml, relation, options = {}) relation.id = xml["id"] read_infon(xml, relation) read_recursive(xml, relation, "node") true end
read_sentence(xml, sentence, options = {})
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 85 def read_sentence(xml, sentence, options = {}) sentence.text = read_text(xml, "text") sentence.offset = read_int(xml, "offset") read_infon(xml, sentence) read_recursive(xml, sentence, "annotation") read_recursive(xml, sentence, "relation") true end
read_text(xml, name)
click to toggle source
# File lib/simple_bioc/bioc_reader.rb, line 32 def read_text(xml, name) node = xml.at_xpath(name) node && node.content end


Pokémon Go Chimchar Community Day guide: best movesets and start times
Follow Polygon online:
by Julia LeeChimchar’s Pokémon Go Community Day will take place on Nov. 16 from 11 a.m. until 2 p.m. in your local time zone.
During this time, Chimchar will spawn in abundance and they’ll have an increased chance to appear Shiny.
How do I get a Shiny Chimchar?
Just loitering around during the Community Day time period should net you at least one Shiny Chimchar.  After the Community Day is over, Chimchar will be added to the Shiny pool and will be able to be caught and hatched, but at a dramatically lower rate.
What’s the Community Day move for it?
Like the other fire-type starter Pokémon, Chimchar learns Blast Burn. Infernape with Blast Burn isn’t super meta-relevant, as it’s not as great as Blaziken, Chandelure, or Moltres, but if you have none of those, it’ll make a fine substitute. Evolve your Monferno up to a Infernape from 11 a.m. until 4 p.m. in your local time in order to get the move.
How do I make the most of this Community Day?
 Like every other Community Day, Lures will also last three hours, so plunk a couple of those down in your local spots to increase the Chimchar spawn.
For the best Infernape, you should make sure it has Fire Spin and Blast Burn as its moves.
If you’re deciding which Chimchar to evolve at the end of the day, try using one of the new search strings. Sort your ‘mons by “recent” using the lower right icon, then type “4*” in the search box at the top. This will let you know if you have any 100% IV Chimchar. Most people won’t be so lucky, so next you’ll want to type “3*” in the search box, which will show you any recent ‘mons that are 80% or above. Appraise these to see which one is best and then evolve your new buddy!


Design System 42
A little living style guide
This little living style guide is the answer to everything. From which colors to use over what font to choose through the different button styles at your disposal.
This little living style guide is the answer to everything. From which colors to use over what font to choose through the different button styles at your disposal.


This is the app that I worked on during the UC Health Hack 2018. It was designed mainly to locate the optimal location for getting a flu shot based on price, perks, and distance from you. We also wanted to digitize medical proof of getting flu shots by having a camera function to save the record to your phone or to a cloud service like Amazon Web Services. It is still open for further development, so if you are interested in sponsoring it or fleshing it out further, please contact me.
I developed the basic concept of this app, as well as basic proof of concept work with my roomate Ata and three other students. Unfortunately, we weren't quite able to get a technical demo with all of our planned features in place during the hackathon (turns out learning Swift in 24 hours is hard), but we did develop a clear idea of what we wanted the app to do and how to do it.
I am working on finishing the writeup describing the app and its features, and will have it available here for download once it's complete.
Download the Presentation Check it out on GitHub


A BASH framework by In the Loft Studios, especially for PHP Developers.
https://google.github.io/styleguide/shell.xml#Naming_Conventions
psp make bash/lobster alpha
If you are making an app called "Titanium"...
git clone git@github.com:aklump/lobster.git alpha
and hit enter.example_app
as titanium
somewhere on your computer and start building your app. That is the only folder you need from this repository to bundle with your app.titanium/gitignore
to titanium/.gitignore
.example_app.sh
as titanium.sh
; this will be the controller for the app.dist/.lobsterconfig
to titanium/.lobsterconfig
and modify the necessary items.dist/.example_appconfig
to titanium/.titaniumconfig
and use this file for any configuration used by your app.example_app/routes
./docs/public_html
.


Probability of winning anything depending on the number of tickets:
Expectation or mean of how many rounds it will take to win anything, depending on the number of tickets:


We tend to offer more aid when a specific individual is in need than when a group is in need.
Based on doodles-site-example.webflow.io


dotnet publish
Publish a .NET application and its dependencies to a directory for deployment to a hosting system.
More information: home page
- Compile a .NET project in release mode:
dotnet publish --configuration Release path/to/project_file
- Publish the .NET Core runtime with your application for the specified runtime:
dotnet publish --self-contained true --runtime runtime_identifier path/to/project_file
- Package the application into a platform-specific single-file executable:
dotnet publish --runtime runtime_identifier -p:PublishSingleFile=true path/to/project_file
- Trim unused libraries to reduce the deployment size of an application:
dotnet publish --self-contained true --runtime runtime_identifier -p:PublishTrimmed=true path/to/project_file
- Compile a .NET project without restoring dependencies:
dotnet publish --no-restore path/to/project_file
- Specify the output directory:
dotnet publish --output path/to/directory path/to/project_file


(.) factors (.–.) study risk HFRS {border-style cases No Yes control px;} #ffffff;} fever home renal cats hemorrhagic food raising rats regression syndrome Jiangsu Raising rodent disease rooms spare Med controls Provincial cleaning px; {font-family China case-control crops excreta font-size infected logistic onset prevention rodents used Hemorrhagic Infect analyses case data dogs double; effect epidemic exposure matched measures multivariate patients previous virus (Table (p > .) China, Control HFRS. P-value Prev Table analysis border-color border-width factor hantavirus month prior protective reported significant solid; study, study. {font-style {font-weight %CI CI) Confidence Funding HFRS, Medical Odds Presence Regression [], []. age areas bold community conducted courier; defined effects em; health including individual infection interval investigation joint n(%) presence properly pt;} ratio, route taking working


Dancing Drones Command Center
Select a JOCS File
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MoCap:
Ready:
Playing:
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0
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%
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Shane Henry
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Started site development at 05.04.2018
Copyright PrivateGER (2018)
About
I'm PrivateGER, your average hobbyist developer. I'm skilled in a variety of languages and programming related tasks.
Projects
I'm the lead developer and owner of codecaptcha, lead dev of MTU, developer of this site, developer of JSHashcracker and developer of the PrivateGER Music Discord Bot.
Started site development at 05.04.2018
Copyright PrivateGER (2018)


Select your Ward
{{ward.WardName}}
ward
{{ward.OccupiedBeds}}
Patients
{{ward.OccupiedBeds}} Beds Occupied
{{ward.VacantBeds}} Beds Vacant


{% extends 'base.html' %} {# Prevent the helper panel from appearing by overriding it. #} {% block helperPanelContainer %}{% endblock helperPanelContainer %} {% block mainContent %}
Please email us directly<EMAIL_ADDRESS>We welcome negative and positive input. And hellos.
{% endblock mainContent %}


Here’s a list I just received of some interesting items. Most might already be widely known, if not all of them, but they still made me smile.  No purchase necessary. Details insideâ€?.

(The shoplifter special?)

On a bar of Palmolive soap:

-"Directions: Use like regular soap".

(And that would be how???)

On some frozen dinners:

-"Serving suggestion: Defrost".

(But, it’s just a suggestion).

On Nanna’s Tiramisu dessert (printed on bottom):

- "Do not turn upside down".

(Well…duh, a bit late, huh)!

On Marks & Spencer Bread Pudding:

-"Product will be hot after heating".

(And you thought????…)

On packaging for a K-Mart iron:

"Do not iron clothes on body".

(But wouldn’t this save me more time?)

On Boot’s Children Cough Medicine:

"Do not drive a car or operate machinery after taking this medication".

(We could do a lot to reduce the rate of construction accidents if we could just get those 5 year olds with head colds off those forklifts.)

On Nytol Sleep Aid:

"Warning: May cause drowsiness".

(And…I’m taking this because???)

On most brands of Christmas lights:

"For indoor or outdoor use only".

(As opposed to…what?)

On a Japanese food processor:

"Not to be used for the other use".

(Now, somebody out there, help me on this. I’m a bit curious.)

On Nobby’s peanuts:

"Warning: contains nuts".

(Talk about a news flash!)

On an American Airlines packet of nuts:

"Instructions: Open packet, eat nuts".

(Step 3: maybe, uh…fly Delta?)

On a Swedish chainsaw:

"Do not attempt to stop chain with your hands or genitals".

(Was there a lot of this happening somewhere?)

 * I don’t blame the company, I blame the parents for this one:

On a child’s superman costume:

"Wearing of this garment does not enable you to fly".

Now that you’ve smiled at least once, it’s your turn to spread the stupidity and send this to someone you want to bring a smile to (maybe even

a chuckle)…in other words send it to everyone. We all need to smile every once in a while.

I hope it made you smile too :)


You can submit the form now! 1 question is remaining {{numberOfQuestionsRemaining}} questions are remaining Made with Survaider


£2.1m funding boost for Cloudsmith in seed investment round
A BELFAST-based technology firm has closed an equity investment round of £2.1 million - one of the largest seed funding boosts for any start-up in the north.
Cloudsmith, founded in 2013, has developed a potentially world-first artifact repository and caching platform that provides secure digital storage for software assets under development.
The solution helps simplify, secure and accelerate software development and delivery pipelines.
The funding round was led by Frontline Ventures and co-investors MMC Ventures, and locally by Techstart Ventures through its Invest NI fund Techstart.
Cloudsmith’s software-as-a-service (SaaS) offers an advanced logistics platform for software developers, operators and vendors, who need to secure, manage and accelerate their product development and distribution. It distributes millions of artifacts on behalf of its global customer base.
The firm's chief executive Alan Carson said: “We believe that the concept of provenance will play a defining role in the future of software security and quality.
“All companies need end-to-end visibility and control of the assets in their software pipelines. It’s important to have the support of Frontline and MMC as we enter this critical stage in Cloudsmith’s growth.”
Steve Collins, partner at Frontline Ventures, said: “The guys are building what is in effect a foundational element for the next stage in the evolution of software development and distribution. Protecting developers and businesses from the chaos of public software will be a game changer.”
Hal Wilson, partner at Techstart Ventures, added: “This investment is a clear vote of confidence in the Cloudsmith team and their ability to execute.”


---
layout: page
title: "About"
description: "This is what I do."
---
Fernando Sainz
- Full stack web developer since 2007
- Doing mostly Ruby and Rails, Go and Vue.js
- Cofounder of pagebox and wolty
- Working for i22 Digitalagentur.
- Living in Bonn, Germany.


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What does this do?
It's hard being a maintainer/active contributor of an open source project 😭. You have to keep an eye on all the bug reports
and pull requests and provide your review on the same. I have realized that I have been wasting so much of my time
in browsing all the issues/PRs lately. The current flow involves going on an issue page and open all the issues in
new tab in order to see the issue details etc.
To make this process a little bit easier, I have built this small tool. Here you don't need to open the issues in
a new tab. In a single page you can browse all the issues/PRs and comment on the same. This saves a lot of time +
RAM also 😛.
How do I comment on issue?
This will be supported once I implement the GitHub oAuth.
I can't see my repo?
GitHub has an api limit so once you hit it, issues won't be able to load. This will be improved after GitHub oAuth (also you'll be able to browse private repo).
I want to improve this project?
Please do ❤. The project is built on top of Angularjs and hardly contains 100s loc in js.
Who are you BTW?
I'm Akash from INDIA. It's been five years since I fell in love with OSS. I have been contributing to Zulip full time since last two years. You can check out more about me here.