Ancestral introns: SGSH

This example illustrates the concept that -- while most human introns are very ancient (very highly conserved, established prior to the emergence of multicellual organisms) -- exceptions exist that confirm that processes creating new introns continue to be operative. It also illustrates that protostomes, notably fly and worm model organisms, are highly unrepresentative in exhibiting much, much greater levels of intron churning than vertebrates.

Although there are 17 human sulfatases, it is straightforward -- because of diagnostic residues and intronation pattern -- to trace back orthologs of individual family members to protostomes and pre-bilaterans. This is done for heparin sulfatase, SGSH, below.

This is rather an interesting situation because 3 of the 7 introns do not date back to early metazoans (eg sponge or anemone) but are post-echinoderm de novo events. Thus intron 2 (human numbering) arose prior to amphioxus, while introns 5 + 7 appear to have arisen after amphioxus but before lamprey.

As new introns are quite rare in chordates, this gene is potentially informative to the phylogenetic tree topology here. Unfortunately the hemichordate gene is not currently available and tunicates do not retain any ancestral information.

The table below shows the phylogenetic origin of human SGSH introns. Fused introns are denoted by ^ showing arthropods have most ancestral introns but also many fusions and insertions.

1 0 1 2 0 1 1   human <--> lamprey 4-5 fused fish, insertions in fugu, ciona fused
1 0 1 2 0^1 1^  amphioxus
1 0^1 2 0^1 1^  sea urchin 
1 0^1 2 0^1 1^  limpet + annelid   
1 0 1 2 0 1 1   anemone
1^0^1^2^0^1^1^  sponge 
1 0^1 2 0^1 1^  silkworm
1 0^1 2 0^1 1^  bee several new distally
1 0^1 2^0^1^1^  beetle several new distally
1^0^1^2^0^1^1^  drosophilids
1 0^1^2^0^1^1^  mosquito ancestral intron 1
1 0 1^2^0^1 1   wasp 2 ancestral
1 0^1 2 0^1 1^  tick
1 0 1 2 0 1 1^  daphnia
1 0^1 2 0^1 1^  ancestral deuterostome note 2 is urchin/amphi, 5 + 7 are amphi/lamprey
1 0^1 2 0^1 1^  ancestral bilateran same or fewer

>SGSH_hsa Homo sapiens human 510 aa 8 exons

0 MSCPVPACCALLLVLGLCRARPRNALLLL 1 
2 ADDGGFESGAYNNSAIATPHLDALARRSLLFRNAFTSVSSCSPSRASLLTGLPQ 0
0 HQNGMYGLHQDVHHFNSFDKVRSLPLLLSQAGVRT 1
2 GIIGKKHVGPETVYPFDFAYTEENGSVLQVGRNITRIKLLVRKFLQTQDDR 2
1 PFFLYVAFHDPHRCGHSQPQYGTFCEKFGNGESGMGRIPDWTPQAYDPLDVL 0
0 VPYFVPNTPAARADLAAQYTTVGRMDQ 1
2 GVGLVLQELRDAGVLNDTLVIFTSDNGIPFPSGRTNLYWPGTAEPLLVSSPEHPKRWGQVSEAYVSLL 1
2 DLTPTILDWFSIPYPSYAIFGSKTIHLTGRSLLPALEAEPLWATVFGSQSHHEVTMSYPMRSVQHRHFRLVHNLNFKMP
FPIDQDFYVSPTFQDLLNRTTAGQPTGWYKDLRHYYYRARWELYDRSRDPHETQNLATDPRFAQLLEMLRDQLAKWQWETHDPWVCAPDGVLEEKLSPQCQPLHNEL* 0

>SGSH_gga chicken 499 aa 8 exons like human
0 MRVWALPLLLGALRLDGAPARNVLLLL 1
2 ADDGGFESGAYNNSAIRTPNLDALARRGLLFQNAFTSVSSCSPSRASVLTGLPQ 0
0 HQNGMYGLHQGVHHFNSFDAVRSLPGLLRQANIRT 1
2 GIIGKKHVGPEAVYPFDFAYTEENSSVLQVGRNITRIKVLVRRFLQSQDVR 2
1 PFFLYVAFHDPHRCGHSQPQYGAFCEKFGNGESGMGWIPDWKPQLYPPKDVQ 0
0 VPHFVPDTPAARADLAAQYTTIGRMDQ 1
2 GIGLVLEELQRAGVLNSTLVIYTSDNGIPFPSGRTNLYWSGTAEPLLLSSPEHPGRWGQVSSAFASLL 1
2 DLTPTILDWFSIPYPSYSIFGTKRVQLTGKSLLPALQSEQPWATAFSSQSHHEATMYYPMRAIQHRQFRLIHNLNYKMPFPIDQDFYVSPTF
QDLLNRTRAGQPTHWNKTLHQYYYRDRWELFDCSQDPTESHNLASDPRYAAIFQMLRAQLLKWQWDTGDPWVCAPNAVLEEKLSPQCQPLHNEL* 0

>SGSH_xtr Xenopus tropicalis frog gappy no fusions
0 MGSRVWYWVIYSLLLLSKAWGRNVLLII 1 
2 GDDAGFESEVYNNTAIHTPNLRDLSKRSLIFKNAFTSVSSCSPSRAAIMTGLPQ 0
0 HQNGMYGLHQDMHHFNSFDDVRSLPLILRQAGIRT 1
2 GIIGKKHIGPESVYPFDFSYTEENSSVLQVGRNITRIKLLVRKFLQSQDQR 2
1 PFLLYVAFHDPHRCGHSQPQYGAFCEKFGNGDPDMGIMPDWSPQYYTPEQVQ 0
0 VPYFIQDTPSARKDIAAQYTTIGRMDQ 1
2 GIGLVLSELYNAGHENDTLVIFSSDNGIPFPNGRTNLYWSGRAEPLLVSSPYHQKRWGQISQSFASLL 1
2 DITPTVLDWFSIPYPNYKIFGKSVQLTGKSLLPALQSEQDWTTVFGSQSHHEVTMYYPMRSVQNLQYLLIHNLNFKMPFPIDQDFYVSPTFQDLLNRTVSGQPTSWFK
TLHNYYYRDRWELYDRSADISEIKNIAEDPAYQDILKSMQNILQKWQSETSDPWMCAPDGVLEEKLEPQCRPLYNEL* 0

>SGSH_fru Fugu reripes fugu 8 exons 499 aa fused exons 4-5 exon 8 split in 3226 bp mus intron Percomorpha not Clupeocephala
0 MFFPLSFVIFSSCIWESDTRNVLLII 1
2 ADDAGFETEVYNNSVVHTPHLRALAQRSLVFNNAFTSVSSCSPSRSAILTGLPQ 0
0 HQNGMYGLHQGVHNFNSFEGVQSLPLLLSKANVHT 1
2 GIIGKKHVGPGSVYPFDFAYTEENGSVLQVGRNITRIKLLVRKFFQAHKEDKVNSQEEER PFFLYVAFHDTHRCGHSQPQYGAFCEKFGNGEKGMGRIPDWKPVYYTPDQVK 0
0 VPPFAPDTPVTRADLAAQYTTVSRLDQ 1
2 GIGLVLQELREAGYENDTLVIYSSDNGIPFPNGRTNLYHSGTAEPMMVSSPEHRKRWGETSQAYVTLL 1
2 DITPTILDWFSVPYPSYSLPGNPRTPVHLTGRSLLTVLSAEPRSWDTVYASQSLHE 0#0 VTMYYPTRSVHQGVYHLLHNLHYRMPFPIDQDLYVSPTFQDLLNRTRRQEPTHWFKSLEQYYYRERWELYDSR 2#1 TDPLETVNLASDPSYSTVLENLRQSLQKWQWETGDPWVCGPDYVLEDKLEPRCRPLYNGL* 0

>SGSH_tni Tetraodon nigrovirens fkusion with zfish, 2 novel insertions
0 MLLRISLLIFSCCIWESDTRNVLLIIA 1
2 ADDAGFETEVYNNSVVHTPHLRALAQRGLVFSNAFTSVSSCSPSRSAILTGLPQ 0
0 HQNGMYGLHQGVHNFNSFEGVQSLPLLLRKANIHT 1
2 GIIGKKHVGPGSVYPFDFAYTEENSSVLQVGRNITRIKLLVRKFFQAHKEDRANGQEER 21 PFFLYVAFHDTHRCGHSQPQYGAFCEKFGNGDMGMGRIPDWKPVYYTPEQVK 0
0 VPPFVPDTPASRADLAAQYTTVSRLDQ 1
2 GVGLVLRELRDAGYENDTLVIYSSDNGIPFPNGRTNLYRSGVAEPMIVSSPEHRERWRETSQAYVTLL 1
2 DITPTILDWFSLPYPSYSLPGSPSSPVHLTGRSLLPVLSAEPGNWQTVFASQSLHE 0#0 VTMYYPTRSVHRGAYHLLHNLHYRMAFPVDQDLYVAPTFQDLLNRTRSGEPTHWFKSLGRYYYRERWELYDAR 2#1 ADPLETVNLASDPAYSGVLENLRQSLQKWQWETGDPWVCGPDYVLEDKLEPRCRPLYNGL* 0

>SGSH_dre Danio rerio like human
0 MAFVFAWTLLCLLLCFDVGGCRSRNVLLII 1
2 ADDGGFETDVYNNTVVQTPHLRALSKRSLIFKNAFTSVSSCSPSRSTILTGLPQ 0
0 HQNGMYGLHQGVHHFNSFDGVQSLPLLLKRANIHT 1
2 GIIGKKHVGPGPVYPFDFAYTEETNSVLQVGRNITKIKLLVRKFFQSHKEERSETKEER 21 PFFLYVAFHDPHRCGHSQPQYGVFCEKFGNGESGMGRIPDWEPKYYSPDQVK 0
0 VPYFIPDTPAARADIAAQYTTVSRLDQ 1
2 GIGLVLEELRKAGFENDTLVIYSSDNGIPFPNGRTNLYGSGVKEPMLLSSPEHQQRWGKLSQAYVSLL 1
2 DITPTILDWFSLPYPSYSLSMSQPVELTGRSLLPALISEPSWDTVFSSQSLHEVTMFYPMRSIHKGPYRLLHNLHYRMPFPIDQDFYISPTFQDLLNRTQSGRPTGWFKTL
NEYYYRQRWELFDVRTDPMEKVNLAGDLDYSEVLESLKDLLLKWQWRTEDPWVCEPDAVLEAKLEPECRPLYNGL* 0
 
>SGSH_lra Leucoraja erinacea skate cdna CV547418
CLSLLLGSTGSGALHHNRNVLLIIADDAGFESGVYNNTVVPTPSLGALAAHGLVFTHAFTSVSSCSPSRASLLTGLPQHQNGMYGLHQSVHHFNSFQGVRSLPMLLHQAGIHTGIIGKKH
VGPEEVYPFDYAETEENNSILQVGRNITHIRQLVHTFLTGNSDRPFFLYVAFHDPHRCGHSHPQYGSFCEKFGNGEPGMGRIPDWRPRHYRPEQVTVPPFMPDTPASRSDLAA

>SGSH_ebu Eptatretus burgeri hagfish cdna BJ652802
FFLYIGFHDPHRCGHSHPELGSFCERFGSGEPGTGSIPDWKPQHYSPSEVKVPYFVPDTPAARADLAAQYTTINRLDQGIGLILDELQRAGHEGDTLVIYTSDNGIPFPTGRTNLYQQGLAEPLIVSSPRHRARWGQRTKAMASLLDITPTMLDWFSVPYPEYAIFHGTRPVHLTGRSLLGALTHELSWNVAFGSQSLHEITMFYPMRSIxVGNLHLI

>SGSH_pma Petromyzon_marinus lamprey 8 exons like human
0 1 
2 ADDGGFELGAYNNTAIATPHLDALASLSLRFRHAFTSVSSCSPSRAALLSGLPQ 0
0 HHNGMYGLHQDVHHFNSFEQVRSLPLLLNQSGIRT 1
2 GIIGKKHVGPEAVYPFEFSYTEENNSILQVGRNITHIKQLVRAFLRMDDGR 2
1 PFFLYVAFHDPHRCGHSQPQFGSFCEKFGNGAPGMGTIPDWKPHHYNPDDVK 0
0 VPYFVPDTPTARADLAAQYTTIGRLDQ 1
2 GVGLVLEELRSAGYADDTLVIYSSDNGIPFPGGRTNLYTSGVAEPLLVSSPEDRERWGHSSEAYVSLL 1
2 DITPTMLEWFSVPYPDYAMFKKDEPVVLTGRSLLPALKQEPPWDIAFSSQSHHEVTMYYPMRALRHSHLHLIHNMHFLMPFPVDQDLYVSPTFQDLLNRSMQGQPTHWYKSLREFYYRERW
ELYDTRADPAETCNLAHDAAYADTLRTLQQQLRAWQWATYDPWVCAPDGVLEAQGQYKEHPQCLPLHNLL* 0

>SGSH_cin Ciona intestinalis tunicate 2 exons unrelated 507aa total fusion followed by 
0 MFTLKSISFINILFWVYSLSSGSDIRPNVLVLVADDLGFELNAYENNVIKTPNINDLADRGIVYSNAFTTVSSCSPSRSTILTGLPQHQN 1#2 GMYGLHNGYHHFNSFDEVKSLPFLLHENGIRTGIIGKKHVAPEAVYPFDFAETEENNSILQVGRNITRMKELAKQFFSMQLKNESFLLYIGF
HDPHRCGHTHPQYGEFCEKFGNGDYRMGKIPDWKPDYYSPDDVIVPPFVQDTPASRKDISAQYTTISRLDQGVGLIINELKQAGFLESTLILF
TSDNGIPFPNGRTNLYNSGTAGPFILALPVQKHKQAVVDNSYVSLLDITPTVLDWFSITYPQYTLFHRDVKLTGKSLLKDISNSDVAFGSHSL
HEVTMYYPMRSVHKNGLLLIQNLNYLMPFPIDQDFYLSLSFQDLLNRTQTGADLHWSKTLKQYYQRSQLELFNLTSDPLELDNLAYKPEYH
NILTDMQTLLQQWQNTTWDPWRCAPSGVLQDSGAYKLNPTCLPLLNGL* 0

>SGSH_csa Ciona savignyi tunicate 2 exons 77%
0 MVKLDSEYGLIVFFCLASLNSAVLPNALVLVADDLGFELNCYGNNVINTPNIDDLANKGVIYNNAYTTVSSCSPSRSTILTGLPQHQN 1#2  GMYGLHNGYHHFNSFDGVKSLPLLLHANGVRTGIIGKKHVAPEAVYPFDFAETEENNSILQVGRNITKIKELTRQFFLDQPKNESFFLYIGFHDPHRCGHTHPQYGEFCEKFGNRDSG
MGDIPDWKPDIYDPNYVIVPPFVQDTPASRKDISAQYTTISRLDQGVGLVINELKKAGFLDSTLILFTSDNGIPFPNGRTNLYGSGTAEPFFLSNPIQEHKFGEVNQEYVSLLDITPTVL
DWFRISYPEFKLFGREVQLTGKSLLRENTKSNVVFGSQSLHEITMYYPMRSVLQNNLRLIENLNYLMPFSIDQDFYLSLSFQDILNRTQTGTDLPWIKTLKQYYYRPRLELFNLTADPQE
TNNLAYAPEYTNTVSTLRGLLMQWQNVTWDPWRCSPGGVLQDSGAYKFNPTCMSLLNGL* 0s
 
>SGSH_bfl Branchiostoma floridae amphioxus 6 exons 2 fusions shared with urchin but maybe this is ancestral
0 .. 1
2 gDDAGLEMQVYNNTVCKTPHLNSLASRSLTFTQAFTSVSSCSPSRSAILTGqgs 0
0 HQNGMYGLHQGYHHFNSFDTVRSLPLLLNQSGIRT 1
2 GIVGKKHVGPESVYPFEFAHTEENNHIMQVGRNITLIKHLVREFLQQKDDR 2
1 PFFLYIGFHDPHRCGHTNPEYGNFCEKFGNGEPGMGLIPDWTPVHYSPEEVV 00 VPPFVQDTPAARSDLAAQYTTISRLDQ 1 
2 GIGLILQELQAAGHDKDTLILYSSDNGIPFPNGRTNLYNSGMAEPLLLSSPLHTSRWGQTTNSFASLL 12 DVVPTVLDWFGLEYPEYEIFGKNKLVKLTGKSLLPALKEEQSWNTVYASHDLHEITMFYPMRVIRTGDYRLIQNLNFAMPFPID
QDFYLSPAFRGLLNRTRKGQPLHWFNTLKNYYYRPQWELYDLIYNPQETVNLAGNSDYRDVLQELRSQLQAWQKVTYDPWICAPWGVLEDEGPYKDNPVCMSMDNGT* 0

>SGSH_spu Strongylocentrotus purpuratus (purple urchin) 5 exons 3 fusions
0 MNLMVFHLFLLLILLQNGLTCGKNVLVLV 1
2 ADDGGFEMGVYNNTVIKTPHLDALGKQSLVFKHAYTSVSSCSPSRSVIMTGLPQ 00 HQNGMYGLLNGYHHFNSFDEVRSLPMLLGQAGVRT 1
2 GIIGKKHVGPEAVYPFDYSKTPEDGYPIMQVGRNITLIKQYAREFLQTNDTR 2
1 PFFLYIGFHDAHRCGHTHPEFGQFCEKFGNGQPNMGTITDWTPAKYDPNDVI 00 VPYHVQDTPVARDDISAQYTTVSRLDQ 1
2 GVGLMMKELEAAGYLKDTLIIYTSDNGIPFTSGRTNLYECGSREPFMISSPFHQERWGQTSDAFISLM 12 DITPTVLDFFGIKYPKYKIFKGSVQLTGKSLLPALTSEPSGWNVSLSSHDLHE
ITMFYPQRVIRTPRYRLIHNLNFAMPFPIDQDFFISHTFQDILNRTRNHQPLHWYKTLKDYYYRPEWELFDLIDDPTEVNNLAYVGKYEGLLVDLKEHLVDWQNVTNDPFRCYPWGVLEDAGDYKYSPTCLPLDNGL* 0

>SGSH_lgi Lottia gigantea limpet 5 exons 
0 1 
2 GDDAGLQMSAYGNRDIKSPNFDQLAAKSLVFKHGFTSVSSCSPSR 2#1 SVILTGIPQ 00 HQNGMYGLHHNPHHFNSFDDIRSLPVILGDHGIRT 1
2 GIVGKKHVGPDYVYKFDYEQTEENNSLNQVGRNITFMKLKIQEFLSNNDTR 2
1 PFLMYIGFHDPHRCGHVHPEFGSFCEKFGNGEPGMGVIPDWKPVSYNTDDIL 00 VPYFIQNTEIAKQDIAAQYTTISRLDQ 1
2 GIGLMIKELELAGVLDDTLIIYSSDNGIPFPNGRTNLYDAGMAEPMLISSPSDTHRWGQV 0#0 KSEAMVNLV 12 DIVPTVLEWFGLDYPTYKLNKQ IVKLTGKSLLPILHEEPTSGWNSVYASHDLHEVTMYYPMRVLRKRQYKLIHNINYKMPFPIDQDFYLSPTFQDLLNRTRHK
KNLNWTKSLKSYYYRPQWELYNIINDPQELKNLAYNKQFIDVLRSLKVELNQWQNITNDPWICAPGGVLEASGTYKYTPSCLPLDNDTEEQYDESEYTISVV* 0

>SGSH_aca Aplysia californica (sea hare) Protostomia; Mollusca; Gastropoda
2 RSAILSGLPQHQNGMYGLHHGVHHFNSFDGLRTLPNILSKAGIKTG 2
WIRFHDPHRCGHTHPQYGQFCEKFGNGEPGMGVIPDWTPVSYSAQEVEVPDFVQDTPAAREDIAAQYTTISRMDQG
GIQLILTELEKAGHSNDTLISSLGHGIPFPDGRTNLY

>SGSH_cca Capitella capitata fusion Protostomia Lophotrochozoa Annelida Polychaeta
0 1
2 GDDAGLEAGVYNNSVCKTPNIDRLAARSLLFKYAFTSVSSCSPSR 2#1 aILTGLPQ 00 HQNGMYGLHQGTHHFNSFDQVQSLPAILQKNNIRT 1
2 GIIGKKHVGPSPVYPFDFAATEENNSIMQVGRNITRMRQLARSFLTQKDDR 2
1 PFFLYIGFHDPHRCGHTNPEFGAFCEKFGNGEAGMGRIPDWTPIHYDPDDVE 00 VPYFVPDTPAARMDIANQYTTISRLDQG 1
2 GIGVMMQELEISGHLEDTLVMFTSDNGIPFPLGRTNLGEAGT .. 1
2 SHPLMYTLNIHQVTYTGHSLLPSIEGTTDSPVFSSQSLHEITMYYPMRTIRTKQYRLIKNINYKMPFPIDQDFYLSPTFQNILDRERNHQDQHWIKTLSQYYYRPSYEL
YDLETDPKELKNLVGDAKYSDIFKGLNDQLNDWQNATADPWICSPVGVLEDAGAYKQHPVCMPLLNHL*

>SGSH_nve Nematosetlla vectensis rough 3 exons
0 MAVVRRTIALCRHVAISKIPLALLVLLISSAESRKNVLLII 12 GDDAGFESQVYNNSVCKTPHLNALASRGLVFRxAFTSVSSCSPSRSAIL isilyLYTGLPQHQNGMYGLKQNEHHFHSFDAVKSLPLLLKQHDIRT 12 GIIGKKHVALQ 0 
VHPCDLASTEEHNQINQVGRNITYMKELVKKFLQESIDDPRQFFLYVAFHDPHRCGHTSPQFGPHLTRLRGFSPAPPSRFCEKFGNGDPGMGTIPDWTPVLYKPEDVVVPYFVQDTPAARADIAAQYTTISRLDQGIGIFLEE
LKVAGFDKDTLVIFSSDNGIPFPSGRTNLFDPGMKEPFIVSSPYHTKRWGEVSEAFVSLVDIVPTVLDWFSIEYPSYEIYGYNKVELTGTSVLPILEKEPSSGWDTVYASHNLHEVSMYYPMRVLRTKNYKLIHNLNYKMPFPIDQDFMISPSFQDLLNRTSKGEPTNWYKTLQQYYYRPRWELYDIIKDPHEMNNLATKEKFKVVFQGLKKKLNIWQNSTNDPWICAPGGVPLKSRWYP* 0

>SGSH_reb Reniera sponge no introns
0 MSHYSILLVFFLTCSCFTAHAKRNVLLMVADDEGLETPIYGNNRIKTPNLQRLAQRSLVFNHAFTSVSSCSPSRSCIMTGLPQHQNGMYGLEHAIHHFSSFDGIMSLPRILNKTEKYWTG
IIGKKHVAPESVYPYAYSFTEQDGYNLNQVGRNITLMKELARDFLAQAQKSDLPFFLYIGFFDAHRGCDGFCENFGDGSKGNGVIPDWTPTVYDPDDAEAPYFIPDTPVARKDIANQYKT
ISRLDQGVGLMLDALKDFGFDDDTLILYIPHNGTPFPNAKTNLYESGMVEPMMISNPEDKSRWGKTSDALVSSTNIVPTVLDWFGLKYPDYTVFGPNPTRLETESLLPVXAXEPNKKKAP
VFASHDFHEVTMYYPMRVMRTKDFRLIHNLNFAMPYPLATDLYSSITYLDLLSNVAANKSTHWFKTLHQYYYRDQYELFDIKNDPHELKNLATDPEYASVFEEMKSNLTEWRRITDDPWLCWPSGVLLGSKCNPLYNGL* 0

>SGSH_dme Drosphila melanogaster fly 524aa 1 exon
MQFLQWIFTLWLIAGCSAGPQNVLLLLADDAGFESGAYLNKFCQTPNLDALAKRGLLFNNAFTSVSSCSPSRSQLLTGQAGHSSGMYGLHQGVHNFNVLPDT
GSLPNLIRDQSGGRILSGIIGKKHVGAANNFRFDFEQTEEQHSINQIGRNITRMKEYARQFLKQAKDEKKPFFLMVGFHDPHRCGHITPQFGEFCERWGSGEEGMGSIPDWKPIYY
DWRNLDVPAWLPDTDVVRQELAAQYMTISRLDQGVGLMLKELEAAGVADQTLVIYTSDNGPPFPGGRTNLYEHGIRSPLIISSPNKEDRHHEATAAMVSLLDIYPSVMDALQIPRP
NDTKIVGRSILPVLREEPPIKESDSVFGSHSYHEVTMAYPMRMVRNRRYKLIHNINYWADFPIDQDFYTSPTFQQILNATLRKQTLPWYRSLLQYYQRPEWELYDIKTDPLERFNL
ADKAKYNGTLKQLREQLFDWQVATKDPWRCAPHAVLQEQGVYKDQPVCLTLGHEALQRPKRRILGQYEEYVVFS

>SGSH_bmo Bombyx mori silkworm Protostomia Arthropoda Insecta
0 MRVTGIKLALIFYFFITETVLSDKVRNVLLLL 1
2 ADDGGFEIGAYRNKICQTPNIDELARNGLLFNNAFTSVSSCSP 2#1 RAALLTGSPS 00 HQNGMYGLHHGVHHXNSFDNVTSLPNLLRQNGIMT 1
2 GIIGKKHVGPSSVYQFDYEQTEENNHINQVGRNITHMKLLAREFIASANKENK 2
1 PFFLYVAFHDPHRCGHSDPQYGPFCERFGSGEEGMGTIPDWQPWYYQWDEIQL 00 PYFIQ 0#0 DTEAARRDIAAQYTTMSRLDQG 1
2 GVALILKELESAGHADDTLVIYTSDNGIPFPSGRTNFYDPGLREPLIIRSPSSSARKNEASGAMVSLL 12 DIMPTVLDWFGIEKEMTNDIWDGDTPKSLLPILEKGQTNITLICYSMVSECLRVSHRDIICLLLNASKMFIATLRAGITLRATLGTVKNTTSL
SLHSKKQNRFLCPYVCLSLCMLKSLKLRNGF*

>SGSH_ame Apis mellifera Protostomia Arthropoda Insecta 
0 MPHKNAVLLL 1
2 ADDGGFEMRSYLNKICQTPNLDNLAKESLLFNNAYSSVSSCSP 2#1 SRSSLLTGLPS 00 HQNGMYGLHHGIHHFNSFEKVQSLPKILKKNNIRT 1
2 GIIGKKHIGPSNVYPFDFSQTEENNSILQVGRNITKIKLLVREFFSQNKTK 2
1 PFFLYIGFHDPHRCGHTHPEYGNFCEKFGNGDIGMGTIPDWN 00 PIYYQWEQVKVPYFVQNTEAARRDIAAQYTTISRLDQ 1
2 GVGLILKELEDAGFKDNTLVIYTSDNGIPFPNGRTNLYEP 1#2 GLAEPMMIRSPIPNHRKNSITYSLTSLL 21 DIVPTLLDWFNIPYMDPSPFDTNEISVPFLTGKSLLPLLIQ 1#2 EPIENNTAIFASQTHHEITMYYPMRAIRTKRYKLIHNINYKMPFPIDQDFYVSPTFQ 1#2 DLLNRTKNKQPLPWYKTLENYYERPEWELYDLKYDPEEKNNIASKSSAK 0#0 NIFSDLQERLLKWQKITNDPWLCAPTGVLNDIKIKKPQCMPLQNLI* 0

>SGSH_tca Tribolium castaneum Protostomia Arthropoda Insecta confirmed browser
0 MGKTGLLVLLVIWARVGAENGKQLNVLLIL 1
2 ADDGGFEMGAYRNKICQTPNLDALAKNSLIFNNAYTSVSSCSPS 2#1 RSALLTGMPA 00 HQNGMYGLHQAENHFDSFTNVKSLPNILRENGIRTGIIGKKHVGPKSTYRFDYEQTEENNSILQVGRNITLIKLLAREFLNNSTDK 2
1 PFFLYVAFHDPHRCGHTHPEYGQFCQRFGNGDVGMGLIPDWRPIYYQWDELE 00 LPYYIPDTEAARREVANQYTTISRLDQ 12 GVGLILEELEKSGHADDTLVIYSSDNGTPFPNGRTNLYDSGIAEPMFISSPLHKERHNQVTYSLTSLL 12 DIVPTVLDWFNITEESNEINSKKLTGKSLLPLL 0#0 AIFASHNLHEVTMYYPMRMIRTHRYKLIHNLNYQAPFPIDQDFYLSPTFQ 0#0 DILNRTRNKENLYWFKTLRQYYNRPEWELYDLKHDPVELNNLAGQSDYKDIMHELEKRLSEWQNATSDP* 0


 
>SGSH_nvi Nasonia vitripennis Protostomia Arthropoda Insecta
1 PFFLYVAFHDPHRCGHSHPEFGSFCEKFGNGEPGMGHIPDWNPIYYQWEQVK 00 LPYHVQDTEPARRDIAAQYTTMSRLDQ 12 GIGLILKELETAGVKDDTLVIYTSDNGIPYTSGRTNLYDP 1
2 GIIGKKHVGPSHAYPFDFAYTEENESILQVGRNITRIKLLVREFLSS
LIYVGMAEPMIISSPYHTDRHNEATYSMTSLLDITPTLLDWFGLTTDKKIEKHLGSLTGKSLIPLL 
 
>Anopheles gambiae (African malaria mosquito) 1 intron, ancesral
0 MSWHTELLLSLVLLSSVPSEAKNVLLLL 1
2 ADDGGFEMGAYRNRIVQTPFLDALAKESLIFNNAYASVSSCSPSRASILTGMPEHQNGQYGLHNGVHNFNSL
PKVHSISSVLGKAGIRTGLIGKKHVGPDETYKFDYERTEEQYPINQVGRNITQIKLFVREFLHQTKSTSNEPFFLMVSFHDPHRCGHVTPQYGSFCERWG
SGEEGMGLIPDWHPIYYVWDEIDLPYYVPDTQPARYDLAAQYTTISRLDQGVGLVLKELRDAGLEDDTLVVYTSDNGPPMPAARTNLYDPGMAEPMFIRS
PEKGVRRNEVTYSMTSHLDLVPTILEWFNLTHPQPTTLTGRSLLPLLFQEPSNQPDDAVFASQSFHEITMAYPMRAIRTKRYKLIHNLNYQLPFPIDQDF
YVSPTFQDILNRTLANQPVPWYKTLRTYYHRPEWELYDLKMDPTESRNLFGKSSMKDTFQQLSERLQKWLEVTKDPFRCAPDGVLQDTGEYLNIPTCLPL
GH* 0

>SGSH_isc Ixodes scapularis tick dna Protostomia Arthropoda; Chelicerata; Arachnida same forward intron, same fusion; last like lottia
0 1#2 glntetpsa 12 ADDGGFETGVYNNTVCQTPHLVELARRGVVFDRAFTSVSSCSPSRASLLTGLPQ 00 HQNGMYGLHQGVHNFQSFPQVRSLPGILAQHGIRT 1
2 GIVGKKHVGPEIVYPFDFAHTEENNSILQVGRNITRIKHLVRKFLSANESK 2
1 PFFLYVAFHDPHRCGHTHPEYGQFCEKFGDGSIPGMGHMPDWTPQRYEPADVSVPYFVQ 00 DTPAARADIAAQYTTVGRMDQ 1
2 EIGLVLQELEATGFGDDTLVLFSSDNGIPFPSGRTNVYEPGIRDPSIVYDPTRPGSAEKVLSEYSK 0#1 RSDAMVSLL 12 DVTPTVLDWFGIQPPDYDIFGKPVILTGASVLPLVGVDGGGEGASGEERAVFASHSLHEATMYYPMRAVRSRGFKLIHNLGFKMPFPI
DQDFYVSPTFQVPSTLLRSFFKRVVMRQQPSLAYAVLLHCFCRHGTRPDASLYPNMVLCTPSAIFGADLFVNVILSCLC* 0

>SGSH_dpu Daphnia pulex Protostomia Arthropoda Crustacea  4x preliminary assembly not for public release by Joint Genome Institute to the Daphnia Genomics Consortium. 
DIASQYTTISRLDQG
2 GVGLVLDELRKAGKADDTLIIFSSDNGISFPNGRTNMYEPGI
2 GLAVPMFIKSPDDESRRGEMTDIQANLL 12 DIVPTVLDWFDIDYLKYHILKPNQPIRLTGKSLLPLLSGENGMKSDTFYGSHVTHEITMNYPMRTIIQEDRYKLIHNLNAPGTPFPIDQDFYLSPTF