A Molecular Phylogeny of Planthoppers (Hemiptera: Fulgoroidea) Inferred from Mitochondrial 16S rDNA Sequences

 

Wen-Bin Yeh1, Chung-Tu Yang1, and Cho-Fat Hui2,*

 

1Department of Entomology, Chung-Hsing University, Taichung, Taiwan 402, R.O.C. Tel: 886-4-22840799-555.  Fax: 886-4-22875024.

2Institute of Cellular and Organismic Biology , Academia Sinica, Taipei, Taiwan 115, R.O.C.  Tel: 886-2-27899512.  Fax: 886-2-27858059.

 

*To whom correspondence and reprint requests should be addressed. 

E-mail: scfhui@yahoo.com

Tel: 886-2-27899512. 

Fax: 886-2-27858059. 

 

Received: 3 September 2003   Accepted: 7 April 2005   Published online: 20 July 2005

 

Abstract          Wen-Bin Yeh, Chung-Tu Yang, and Cho-Fat Hui (2005) A molecular phylogeny of planthoppers (Hemiptera: Fulgoroidea) inferred from mitochondrial 16S rDNA sequences.  Zoological Studies 44(4): xxx-xxx.  Phylogenetic reconstruction of family relationships within the superfamily Fulgoroidea was conducted based on DNA sequences of the mitochondrial 16S rDNA gene.  Sequences of 569 bases of the 3' end of the gene from 59 populations representing 53 species within 15 families were analyzed.  Ranges of the 16S rDNA nucleotide divergences within species were 0%~0.6%, those among species of a given genus were 1.7%~7.8%, and those among genera of the same tribe or subfamily were 8.1%~19.5%.  Scatterplots of total substitutions (Tvs) against transversions (Tv), or transitions (Ts) of the 16S rDNA gene revealed that the mutation rate of Tv was 3 times higher than Ts, and substitutional saturation has not yet been reached.  Phylogenetic reconstruction and bootstrap confidence analysis revealed several basal lineages and 1 advanced group.  The basal lineages included 5 families, where the Cixiidae exhibited close affinity to the Delphacidae, while the phylogenetic positions of the Achilidae, Meenoplidae, and Lophopidae were ambiguous.  Monophyly of the remaining 10 advanced families showed several subdivided family groups with close affinities of the Derbidae to the Tropiduchidae, Dictyopharidae to the Fulgoridae, and Ricaniidae to the Eurybrachidae.  The ancestral lineage of the Tettigometridae which exhibits many ancestral morphological characters was not supported by the 16S rDNA sequence data analysis, and the relationship of the families Flatidae and Nogodinidae was poorly resolved.  In addition, the Issidae was not shown to be monophyletic, although issid members at the subfamily level were well supported.

Key words: Molecular phylogeny, Mitochondrial 16S rDNA, Hemiptera, Fulgoroidea.

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Introduction

Planthoppers constitute a large group of phytophagous insects in the order Hemiptera and family Fulgoroidea including more than 9000 described species with division into 19 families distributed worldwide (O'Brien and Wilson 1985).  These insects occupy extensive ranges of habitats (Denno and Roderick 1990), and some major agricultural pests are included (Wilson and O’Brien 1987).  The Fulgoroidea consists of common herbivores in both agricultural and natural systems, often causing severe damage to their host plants.  Attention has focused on a number of planthopper species because of large damage incurred by the crops of maize, rice, wheat, and forage grasses.  Several species of delphacids exhibit high reproductive potentials and dispersal capabilities which allow them to track changes in favorable resources and therefore predispose them to be agricultural pests (Denno and Roderick 1990).  Nilaparvata lugens (Stål), for example, a delphacid, caused more than US$1.23 billion in losses to rice in South and Southeast Asia annually (Herdt 1987).

The 1st phylogenetic hypothesis concerning the Fulgoroidea was proposed by Muir (1923).  Thereafter, several such hypotheses were proposed based on adult or nymphal morphological characters (Asche 1988, Emeljanov 1991, Chen and Yang 1995).  The Tettigometridae is usually considered the most-ancestral family in the Fulgoroidea since it exhibits many ancestral characters pertaining to the Cicadoidea (Muir 1923).  Other popular relationships of the remaining fulgoroids are those proposed by Asche (1988) and Emeljanov (1991).  Figure 1A shows a cladistic analysis based on 30 adult morphological characters.  It suggests that the Cixiidae is a sister group to the Delphacidae, and both are more ancestral than the others which form several aligned lineages including 1 major group, but the affinity among these lineages is ambiguous (Asche 1988, Wilson et al. 1994).  Asche’s hypothesis was criticized by Emeljanov (1991) based on 50 morphological characters (Fig. 1B), and the Delphacidae has been suggested as being the next-most ancestral and not a sister group of the Cixiidae.  Each of the following lineages in Asche’s hypothesis is further divided in an unequal succession of dichotomies.  Both hypotheses suggest that the Meenoplidae is a sister group to the Kinnaridae, and the Fulgoridae is a sister group to the Dictyopharidae.  These commonly cited fulgoroid phylogenies have frequently been challenged.  The ancestral taxon of the Tettigometridae is questionable, and the primitive characters in tettigometrids are the result of convergence (Fig. 1C) (Bourgoin 1993).  Further molecular evidence using 18S rDNA sequences confirms this controversial point (Bourgoin et al. 1997).  Also, based on the ontogeny of metatarsal characters, Cheng and Yang (1995) suggested that the Delphacidae and Cixiidae constitute the most-advanced lineage of fulgoroids (Fig. 1D).  Thus, the only conclusion generally accepted in fulgoroid systematics is that the Fulgoridae and Dictyopharidae are sister groups.  Obviously, more studies of fulgoroid phylogeny using different characters are needed.

Fig. 1.  Hypotheses of phylogenetic relationships of fulgoroid families.  Phylogenies proposed by Asche (1988) (A), Emelyanov (1991) (B), Bourgoin et al. (1993 1997) (C), and Cheng and Yang (1995) (D).

 

The 3' end sequences of the mitochondrial 16S rDNA gene have been shown to be useful for examining insect relationships from the genus level to the family level (Han and McPheron 1997, Yeh et al. 1998, Marini and Mantovani 2002, Hypša et al. 2002, Whitfield et al. 2002, Ribera et al. 2003).  Mitochondrial 16S rDNA sequences have been used to infer relationships of 6 fulgoroid families of the Tropiduchidae group, and the results indicated that the 3’end sequences of the 16S rDNA are suitable for studying family relationships within the Fulgoroidea since nucleotide divergence increases with increasing taxonomic distance (Yeh et al. 1998).  Nuclear 18S rDNA sequences have been used to evaluate 5 fulgoroid families and the affiliation of the Tettigometridae (Campbell et al. 1995, Bourgoin et al. 1997).  Nucleotide sequences of the 18S rDNA gene may be too conserved for analysis of closely related families since nucleotide divergences between fulgoroid families are in the range of 1.5% to 6.3% (Campbell et al. 1995, Bourgoin et al. 1997).  Also, too few families have been studied so far to obtain a general picture of fulgoroid phylogeny (Campbell et al. 1995, Bourgoin et al. 1997, Yeh et al. 1998).

In this work, sequences of the 16S rDNA gene of 59 populations within 53 species representing 15 families of fulgoroids were analyzed.  The primary aims of this study were to use molecular characters to infer phylogenetic relationships within the Fulgoroidea as well as to address 2 issues that are still being debated: the phylogenetic position of the Tettigometridae, and whether Cixiidae and Delphacidae are ancestral taxa.  Results from fulgoroid 16S rDNA sequences revealed that transversion substitutions are accumulating 3 times faster than transition substitutions.  Phylogenetic analysis showed that members of the same family are grouped together with high bootstrap values, and that the Tettigometridae is not an ancestral taxon.  Finally, the Achilidae, Cixiidae, Delphacidae, Lophopidae, and Meenoplidae were found to constitute the basal lineages, with the remaining 10 families comprising advanced lineages.

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MATERIALS AND METHODS

Taxa examined

In total, 59 populations representing 53 species of 15 families of fulgoroids were included in this study (Table 1).  Specimens of Bruchomorpha oculata, B. jocosa, and Aphelonema histrionica of the Issidae were provided by M. R. Wilson (International Institute of Entomology, London).  Six species selected for outgroup comparisons were Mogannia sp. (Cicadidae, Cica, GenBank accession no.: AF158087), Cosmoscarta kotoensis (Cercopidae, Cerc, accession no.: AF158088), and 4 species of the Cicadellidae: Macrosteles fascifrons (Cic1), Exitianus exitiosus (Cic2), Mocuellus caprillus (Cic3), and Amblysellus grex (Cic4) (Fang et al. 1993).

Figure 1
 

Figure 3
 

Figure 3
 

Table 1.  Taxonomic status of fulgoroid taxa examined in this study with accession nos.

Family

Subfamily or Tribe

Abbrev.

Species name

Accession no.

Achilidae

Plectoderini

Ach1

Betatropis formosana Matsumura

AF158028

 

 

Rhotalini

Ach2

Rhotala formosana

AF158029

 

Cixiidae

Cixiinae

Cix2

Pentastiridius pachyceps (Matsumura)

AF158030

 

 

Cixiinae

Cix3

Cixius circulus Tsaur et Hsu

AF158031

 

 

Cixiinae

Cix4

Cixius inflatus Tsaur et Hsu

AF158032

 

 

Cixiinae

Cix6

Betacixius ocellatus

AF158033

 

Delphacidae

Delphacinae

Del2

Nilaparvata lugens Stål

AF158034

 

 

Delphacinae

Del3

Purohita taiwanensis Muir

AF158035

 

 

Asiracinae

Del4

Ugyops tripunctatus (Kato)

AF158036

 

 

Asiracinae

Del6

Ugyops vittatus (Matsumura) a

AF158037

 

 

Delphacinae

Del7

Nilaparvata lugens Stål

AF158038

 

 

Dlephacinae

Del8

Delphacinae sp.

AF158039

 

Derbidae

Zoraidini

Der1

Zoraida kotoshoensis Matsumura

AF158040

 

 

Zoraidini

Der2

Diostrombus politus Uhler

AF158041

 

 

Otiocerini

Der3

Kamendaka aculeata Yang et Wu

AF158042

 

 

Rhotanini

Der4

Rhotana obaerata Yang et Wu

AF158043

 

 

Otiocerini

Der9

Kamendaka aculeata Yang et Wu

AF158044

 

Dictyopharidae

Dictyopharinae

Dic1

Orthopagus splendens (Germar)

AF158045

 

 

Dictyopharinae

Dic2

Raivuna sp.

AF158046

 

 

Dictyopharinae

Dic3

Saigona gibbosa Matsumura

AF158047

 

 

Dictyopharinae

Dic4

Orthopagus sp. a

AF158048

 

Eurybrachidae

Platybrachinae

Eur1

Platybrachys decemmacula Walker

AF158049

 

Flatidae

Flatini

Fla1

Phylliana alba (Jacobi)

AF158050

 

 

Flatini

Fla2

Mimophantia maritina Matsumura

AF158051

 

 

Nephesini

Fla3

Geisha distinetissima (Walker)

AF158052

 

 

Flatini

Fla4

Mimophantia maritina Matsumura

AF158053

 

 

Flatoidinae

Fla5

Atracis sp.

AF158054

 

Fulgoridae

Aphaeninae

Ful2

Lycorma meliae Kato

AF158055

 

 

Aphaeninae

Ful3

Lycorma olivacea Kato

AF158056

 

 

Fulgorinae

Ful4

Fulgora laternaria Linne'

AF158057

 

 

Aphaeninae

Ful5

Lycorma delicatula White a

AF158058

 

Issidae

Hemisphaerinae

Iss1

Epyhemisphaerius tappanus (Matsumura)

AF158059

 

 

Tonginae

Iss2

Tonga botelensis Kato

AF158060

 

 

Issinae

Iss3

Eusarima astuta Chan et Yang

AF158061

 

 

Caliscelinae

Iss4

Mushya faciata Kato

AF158062

 

 

Hemisphaerinae

Iss5

Hemisphaerius formosus Melichar

AF158063

 

 

Caliscelinae

Iss6

Bruchomorpha oculata Newman

AF158064

 

 

Caliscelinae

Iss7

Bruchomorpha jocosa Stål

AF158065

 

 

Caliscelinae

Iss8

Aphelonema histrionica Stål

AF158066

 

 

Hemisphaerinae

Iss9

Gergithus rotundus Chan et Yang

AF158067

 

 

Hemisphaerinae

Iss10

Gergithus yayeyamensis Hori

AF158068

 

 

Tonginae

Iss11

Tonga westwoodi (Signoret) a

AF158069

 

 

Tonginae

Iss12

Tonga westwoodi (Signoret) a

AF158070

 

 

Issinae

Iss14

Eusarima condensa

AF158071

 

Lophopidae

Lophopinae

Lop1

Lophops carinata (Kirby)

AF158072

 

Meenoplidae

Kermesiinae

Mee2

Nisia serrata Tsaur

AF158073

 

 

Kermesiinae

Mee3

Nisia lansunensis Yang

AF158074

 

 

Kermesiinae

Mee6

Nisia serrata Tsaur

AF158075

 

Nogodinidae

Varciini

Nog1

Mindura subfasciata kotoshonis Matsumura

AF158076

 

 

Pisachini

Nog2

Pisacha naga Distant

AF158077

 

Ricaniidae

 

Ric1

Ricania fumosa (Walker)

AF158078

 

 

 

Ric3

Ricanula pulverosa Stål

AF158079

 

 

 

Ric4

Ricania simulans (Walker)

AF158080

 

 

 

Ric5

Euricania ocella (Walker)

AF158081

 

Tettigometridae

Tettigometrinae

Tet2

Tettigometra sp.

AF158082

 

Tropiduchidae

Tambiniini

Tro1

Kallitaxila sinica (Walker)

AF158083

 

 

Tambiniini

Tro2

Ossoides lineatus Bierman

AF158084

 

 

Catulliini

Tro3

Catullia subtestacea Stål

AF158085

 

 

Tambiniini

Tro4

Ossoides lineatus Bierman

AF158086

 

a Nymph material.

 

DNA extraction, amplification, and sequencing

        Live insects were collected and preserved in 95% ethanol at –20℃.  The entire insect body or the leg of a large specimen was homogenized in a glass homogenizer in 500 μl digestion buffer that contained 100 mM Tris-Cl (pH 8.0), 10 mM EDTA, 100 mM NaCl, 0.5% SDS, 50 mM dithiothreitol, and 0.5 mg/ml proteinase K.  The mixture was incubated at 50℃ overnight, then extracted with phenol-chloroform (modified from Yeh et al. 1998).  Extracted crude DNA was dissolved in 50 μl TE buffer, and an aliquot of 10 μl crude DNA was diluted 10-fold and used as the DNA template in the following amplification reaction.

The polymerase chain reaction (PCR) was employed to amplify a partial sequence of the mitochondrial 16S rDNA gene.  The primers used to amplify the region were 5'-GCCTGTTTATCAAAAACAT-3' and 5'-CCGGTCTGAACTCAGATCA-3' that correspond to nucleotides 13416~13396 and 12866~12884, respectively, of the 16S rDNA gene of Drosophila yakuba (Clary and Wolstenholme 1985).  Amplification was carried out for 39 cycles in a final volume of 100 μl containing 10 mM Tris-Cl (pH 9.0), 50 mM KCl, 1.5 mM MgCl2, 0.01% gelatin, 0.1% Triton-X100, 2 units of SuperTaq polymerase (HT Biotechnology, Taiwan), 0.2 mM of each dNTP, 20 pmoles of each primer, and 2 μl DNA template with the following temperature profile: denaturation for 50 s at 95℃, annealing for 1 min at 50℃, and extension for 1 min at 72℃.  Amplified DNA fragments were separated by agarose gel electrophoresis and extracted from the gel using the Nucleotrap Kit (Macherey-Nagel, Germany).  The resulting DNA product was directly sequenced using the Cycling PCR Sequencing Kit (Perkin Elmer, USA), and 29 cycles were carried out with the following temperature profile: 40 s for denaturation at 95℃, annealing at 50℃, and extension at 72℃ (modified from Yeh et al. 1998).

DNA analysis

        Initial alignment of the mitochondrial sequences was conducted using the Pileup program of the GCG software package (available at http://bioinfo.nhri.org.tw), then manually refined based on the secondary structures of the 16S rRNA sequences (Davis et al. 1994, Fang et al. 1993, Kambhampati et al. 1996).  The proportion of the nucleotide composition of each taxon was calculated using the MEGA version 3 program (Kumar et al. 2004) with a variety of genetic distances.  To correct for AT-richness of the mtDNA sequences and different substitution patterns of transition (Ts) and transversion (Tv), the parameters of nucleotide composition and substitution types were used in the sequence divergence estimation.  The pair-wise distance estimates were based on models that included the proportional, Kimura 2-parameter, and Tamura 3-parameter distance measures.  A statistical analysis system (SAS Institute 2001) was used to test for significant correlations between nucleotide base compositions and substitution patterns of Ts and Tv.

Following the sequence variation estimation, neighbor-joining (NJ) and minimum evolution (ME) implemented in MEGA3 (Kumar et al. 2004) were used for the phylogenetic reconstruction.  Different values of the parameter of α in the gamma distribution were used to determine the effect of heterogeneity in substitution rates among sites.  Sites 40~45, 69~74, 185~186, 279~280, 290~291, 375~377, and 496~497 of the mitochondrial sequence data were excluded from the phylogenetic analysis because they could not be aligned unambiguously.  Bootstrap analyses of 1000 replications were carried out on the trees inferred from the NJ and ME methods.

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RESULTS

Sequence variation

        When gaps were added to the alignment, 569 bases (Appendix I) were analyzed.  Length variations of the partial 16S rDNA sequences among taxa ranged from 534 to 548 bases.  Of the 569 bases examined, 375 bases (66%) were variable.  The average nucleotide composition proportions (± SD) for the fulgoroid sequences were: G, 16.7 (± 1.2); A, 30.6 (± 2.5); T, 43.1 (± 1.7); and C, 9.6 (± 0.5).  A bias towards adenine and thymine is consistent with the base composition of the corresponding 16S rDNA region of other insects (Fang et al. 1993, Dowton and Austin 1994, Kambhampati 1995, Vogler and Pearson 1996, Han and McPheron 1997).  Variable sites were not randomly distributed over the examined 16S rDNA region, and nucleotide divergence patterns in the 3' region of this gene showed that there are 3 highly variable regions (positions 40~74, 279~291, and 372~393; Appendix I).  This result suggests that substitution patterns might be constrained by the function of the gene.  The pattern of substitutions showed the greatest bias with A«T (11.5%), which was larger than changes of A«G (3.2%), T«C (1.8%), and T«G (2.3%), while changes of A«C (0.5%) and G«C (0.1%) were rare.  However, correlation analysis of the base composition and substitution patterns using Kendall’s tau (τ) (Kendall 1938) showed no significant relationship between base composition and substitution patterns (τ = 0.6; τ* = 0.6, α = 0.1).

The uncorrected nucleotide divergences within species were 0%~0.6%, those among species of a given genus were 1.7%~7.8%, and those among genera of the same tribe or subfamily of fulgorids were 8.1%~19.5%.  Regression analysis of total substitutions (Tvs) versus transversions (Tv) and transitions (Ts) revealed that the mutation rate of Tv (with a slope of 0.74; R2 = 0.94) was 3 times higher than that of Ts (with a slope of 0.26; R2 = 0.71), and substitutional saturation due to multiple hits was not yet observed in fulgoroids (Fig. 2).  This information indicates that both Tv and Ts may provide phylogenetic information.

Fig. 2.  Regression analysis of total substitutions (Tvs) versus transitions (Ts, circles) and transversions (Tv, triangles) in the 16S rDNA sequences of fulgoroids.

 

Phylogenetic analysis

Information on the differentiated nucleotide composition and Tv and Ts substitution patterns allowed us to use the 3-parameter estimated distance model (Tamura 1992) in the NJ analysis, and the results are shown in fig. 3A, for which the results of 1000 bootstrap replications are shown in the phylogenetic tree.  Members of the same family were generally grouped together and received significant bootstrap possibilities of from 88% to 99%.  The phylogenetic tree reveals the presence of 2 patterns of lineages.  The basal lineages include 5 families of the Cixiidae, Delphacidae, Meenoplidae, Lophopidae, and Achilidae, and a close relationship is evident in the lineages of the Cixiidae and Delphacidae, although members Del4 and Del6 of the Delphacidae are cohesive to the Cixiidae lineage.  After excluding the 5 basal families, the derived lineages can be divided into many groups and several independently aligned families: (i) Derbidae-Tropiduchidae; (ii) Fulgoridae-Dictyopharidae; (iii) Eurybrachidae-Ricaniidae, and the Flatidae, Nogodinidae, Tettigometridae, and Issidae.  Bootstrap values suggest a robust relationship for each lineage of these groups.  Furthermore, similar phylogenetic topologies were obtained when the data matrix was analyzed under other distance estimation models (Kimura 2-parameter with different values of α in the gamma distribution, Fig. 4a-d).  These trees use family names since members of the same family consistently grouped together (Fig. 3).  The trees support the 2 patterns of lineages but lack resolution among family groups in the advanced lineages.

The minimum evolution result constructed from the 3-parameter estimated distances exhibits the grouping pattern shown in Fig. 3B and is mostly consistent with that from the NJ analysis.  The basal lineages include 5 families, and the relationship between the Delphacidae and Cixiidae is close, but it is necessary to further elucidate the paraphyletic resolution in the Delphacidae.  Relationships among the Meenoplidae, Delphacidae-Cixiidae, Lophopidae, and Achilidae cannot be clearly resolved (Fig. 3B).  Several groups among families in the derived lineages are revealed, including Derbidae-Tropiduchidae, Dictyopharidae-Fulgoridae, and Eurybrachidae-Ricaniidae, but relationships among the remaining families were poorly resolved.  Furthermore, similar phylogenetic topologies were inferred in the minimum evolution analyses when using different values of α in the gamma distribution under the Kimura 2-parameter distance estimate model (Fig. 4e-h).

 

Fig. 3.  Phylogeny of fulgoroids based on the partial mitochondrial 16S rDNA sequences by neighbor-joining (A) and minimum evolution (B) reconstructions based on the 3-parameter Tamura model.  Bootstrap scores exceeding 80% from 1000 replications are given beneath the branches (not shown for branches below the family level).  The black, gray, and hatched rectangular boxes indicate clusters of the outgroups and two defined subdivided groups, respectively.  The black lines indicate families whose phylogenetic relationships are ambiguous according to these analyses.  The dotted-line box indicates advanced lineages.  Taxa of the Issidae (I) are labeled in the subfamily category.

 

Fig. 4.  Phylogenetic tree inferred from the partial mitochondrial 16S rDNA sequences by eighbor-joining (a-d) and minimum evolution (e-h) analyses based on the Kimura-2-parameter distance model.  Correction for rate heterogeneity among sites with different values for the parameter α in the gamma distribution are shown, and bootstrap scores exceeding 80% from 1000 replications are given beneath the branches (not shown for branches below the family level).  The trees use family names since members of the same family grouped together in fig. 3.

 

However, monophyly of the Nogodinidae was not recovered by phylogenetic analysis, with Varciini (Nog1) and Pisachini (Nog2) falling into the Issidae and Flatidae, respectively.  In addition, a non-monophyletic composition was seen in the family Issidae, particularly the subfamily Hemisphaerinae which formed a well-supported cluster with the family Tettigometridae.

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DISCUSSION

The popular view among fulgoroid systematists that the Tettigometridae is the most-ancestral lineage among fulgoroids (Muir 1923, Asche 1988, Emeljanov 1991, Cheng and Yang 1995) was not supported by the 16S rDNA sequence data analyses.  Average nucleotide sequence divergence between the tettigometrid and other fulgoroid families (19.3%) was small when compared to the 2 most-divergent families, i.e., the Meenoplidae (24.5%) and Delphacidae (23.8%).  Also, phylogenetic analyses indicated that the Tettigometridae belongs to a more-advanced lineage of the Fulgoroidea and is grouped with the subfamily Hemisphaerinae of the Issidae.    Based on the 18S rDNA sequence data, Bourgoin et al. (1997) indicated that the Tettigometridae is not a basal family in the Fulgoroidea, and they depicted it as a sister group of the Tropiduchidae.  However, Bourgoin et al. also suggested that the sister taxon of tettigometrids still needed to be rigorously determined partly due to the weak sequence information in the 5’ terminal region of 18S rDNA of the Tropiduchidae.  Obviously, the sister group of Tettigometridae can be further clarified when additional numbers of related sequences are included.

Molecular phylogenies of planthoppers inferred from the 16S rDNA sequences indicate that fulgoroids are monophyletic.  The relationships among families in these analyses are nearly identical: (i) The basal lineages are constituted by the Achilidae, Cixiidae-Delphacidae, Lophopidae, and Meenoplidae; (ii) Robust family groups of the Derbidae-Tropiduchidae, Dictyopharidae-Fulgoridae, and Eurybrachidae-Ricaniidae are aligned with other families in the advanced lineage; and (iii) The Issidae is a non-monophyletic group.   It has been generally accepted that cixiids, delphacids, and meenoplids are relatively ancient families, but the relationships among these 3 families have not been resolved.  For example, it was proposed that cixiids and delphacids are sister groups and are more ancestral than meenoplids based on a cladistic analysis (Asche 1988) with delphacids being more ancestral than cixiids, and cixiids more ancestral than meenoplids (Emeljanov 1991); furthermore, 18S rDNA sequence analysis showed that cixiids and delphacids are sister groups and are more ancestral than the other 5 fulgoroid families (Bourgoin et al. 1997).  However, more-recent relationship reconstruction using cytochrome b (COB) sequences suggested that meenoplids are the most-ancestral family within the fulgoroids (Yeh et al. 1998).  Phylogenetic inferences in these analyses indicated that the Cixiidae has a close affinity to the Delphacidae and constitutes one of the ancestral lineages within the fulgoroids.

Additional evidence from fossil records supports these basal lineages.  The 1st fossil known of an extant member of the Fulgoroidea is of the Cixiidae, which appears at the beginning of the Jurassic period (210 Ma) (Shcherbakov 1996), and the next-oldest fossil is of the Achilidae (135 Ma) (Hamilton 1990).  Most of the other fulgoroid families radiated out in the Cenozoic period (Shcherbakov 1993).  However, sequence analyses revealed an unexpected phylogenetic position of the Lophopidae, which had been considered to be in the advanced lineage based on morphological characters (Asche 1988, Emeljanov 1991, Chen and Yang 1995).  Soulier et al. (1996) pointed out that the Lophopidae is a paraphyletic family.   Molecular results may have been biased from the sample size or the poor alignment of the AT-rich 16S rDNA.  More than 1 taxon of lophopids or an additional sequence, such as 18S rDNA, must necessarily be included to help clarify the evolutionary relationship of the lophopids.

Excluding the 5 families (Achilidae, Cixiidae, Delphacidae, Lophopidae, and Meenoplidae), phylogenetic relationships confirm 1 major derived lineage although several phylogenetic relationships within it are poorly resolved.  This polytomous result was possibly caused by the poor alignment of the 16S rDNA AT-rich regions or by the rapid radiation of fulgoroid families in the Cenozoic period (Shcherbakov 1993).  However, some reliable conclusions can be drawn from the phylogenetic analyses.  Molecular data confirm a close relationship between the Dictyopharidae and Fulgoridae, and depict the Derbidae as a sister group of the Tropiduchidae, and the Ricaniidae as a sister group to the Eubrybrachidae.  Characters of female genitalia suggest that the family Achilidae has a close affinity to the Derbidae (Fig. 1C).  However, according to the adult and nymphal morphological characters (Asche 1988, Chen and Yang 1995), the phylogenetic affinity is ambiguous between the Achilidae and Derbidae.  Asche (1988) proposed a polytomous relationship for the following 7 families: Eurybrachidae, Flatidae, Issidae, Lophopidae, Nogodinidae, Ricaniidae, and Tropiduchidae.  Nymphal characters also reveal a polytomous relationship of the Eurybrachidae, Gengidae, Hypothonellidae, Lophopidae, and Ricaniidae (Chen and Yang 1995).  Sequence data in this study further confirm the relationship of the Eurybrachidae and Ricaniidae, although the positions of the remaining Flatidae, Issidae, and Nogodinidae cannot be precisely defined.  The non-monophyletic resolution of the Issidae and Nogodinidae highlights the difficulty in defining their phylogenetic position.  It has been proposed that the Issidae is a nonhomogenous group (Yeh et al. 1998, Emeljanov 1999, Gnezdilov 2003), although many key morphological characters such as the tegmina, wing length, clavus, corium, and aedeagus shape in issids can effectively define the subfamily classification (Chan and Yang 1994).  A revision of the Caliscelidae was described by Emeljanov (1999), whereas efficient molecular sequence is needed to delimitate the category of issid subfamilies.  Furthermore, according to the characters of the tegmina, more than 10 species of issids were revised into the Nogodinidae (Fennah 1984).  Analyses of 16S rDNA sequences elucidate many phylogenetic relationships within the Fulgoroidea, although many questions remain unresolved and more research effort is needed.

Acknowledgments:This work was supported by Academia Sinica and the National Science Council (NSC-85-2321-B005-016 and NSC-86-2313-B005-072) of the R.O.C.

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Appendix I: Alignment of the fulgoroid 16S rDNA sequences.  Secondary structure domains in the last line are based on previous models (see text)                                         81

Ful2  CGCC TGTTT ATCAA AAACA TGT CCTT TTGGAATTTATTT------ AAGG TT T-GGCC T GCTCAATGA- ---T-- -TTAAAT

Ful3  CGCC TGTTT ATCAA AAACA TGT CCTT TTGGAATTTATTT------ AAGG TT TGGGCC T GCTCAATGA- ---T-- -TTAAAT

Ful4  CGCC TGTTT ATCAA AAACA TGT CTTC CAGGGTTTAATTG------ GAGG TT T-GGCC T GCTCAATGCA ---T-- AGTAAAT

Ful5  CGCC TGTTT ATCAA AAACA TGT CCTT TTGTAATTTATTT------ AAGG TT T-GACC T GCTCAATGA- ---T-- -TTAAAT

Dic1  CGCC TGTTT AACAA AAACA TCT TCTT -TAGTGTTTATTA------ AAGA TA T-AACC T GCTCAATGAT ---T-- TTTAAAT

Dic2  CGCC TGTTT ATCAA AAACA TCT CTTT -TTGGTTTTATTT------ ---- -- A-AACC T GCTCAATGAA ---A-- TTTAAAT

Dic3  ???? ????? ????? ????? ??? ???? ??????????????????? ???? ?? ?????? ? ?????????? ?????? ???????

Dic4  CGCC TGTTT ATCAA AAACA TCT TCTT -TAGTGTTTATTA------ AAGA TA T-AACC T GCTCAATGAT ---T-- TTTAAAT

Fla1  TGCC TGTTT AACAA AAACA TGT CTTT -TAGTTTTTATTT------ AAAG TC T-AACC T GCTCAATGAT -TTA-- TTTAAAT

Fla2  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGAATTTGTTA------ AAAG TT G-ATTC T GCTCAATGAT --AA-- TTTAAAT

Fla3  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGGTTTTATTT------ AAAG TC T-AACC T GCTCAATGAA ---G-- TTTAAAT

Fla4  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGAATTTGTTA------ AAAG TT G-ATTC T GCTCAATGAT --AA-- TTTAAAT

Del8  CGCC TGTTT ATCAA AAACA TTT CTTT -TAGTTTTAATCTTA---- AAGG TT G-GGCC T GCTCAATGAA ---T-- ATTAAAT

Nog1  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGTTTATGTTA------ AAAG TC T-ATCC T GCTCAATGAT -TA--- TTTAAAT

Nog2  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGGTTTT-TTA------ AAAG TC T-GGCC T GCTCAATGAA -TAT-- TTTAAAT

Ach1  CGCC TTGTT ATCAA AAACA TGT TTTT -TTGTTTATTTATTT---- AAGA TT G-GCCC T GCTCAATGAT --A--- TTTAAAT

Ach2  CGCC TTGTT ATCAA AAACA TGG TTTT -TAGTTTATATTA------ AAGA TC A-GACC T GCTCAATGAA -TAT-- TTTAAAT

Der1  CGCC TGTTT ATCAA AAACA TGT CTTT -TTGATTTTATTT------ AAGG TT T-TACC T GCTCAATGAT -----T TTTAAAT

Der2  CGCC TGTTT ATCAA AAACA TGT TATT -TTGATTTTATATT----- TAAT AT TTAACC T GCTCAATGAT ---AA- TTTAAAT

Der3  CGCC TGTTT ATCAA AAACA TGT TTTT -TTGTTTTTTTAA------ AAAG AT TAAATC T GCTCAATGAA A-TAAT ATTAAAT

Der4  CGCC TGTTT ATCAA AAACA TGG TTTT -TAGTTTTTAATTT----- AAAA AT T-TGGC T GCCCAATGA- --T--T -CTAAAT

Der9  CGCC TGTTT ATCAA AAACA TGT TTTT -TTGTTTTTTTAA------ AAAG AT TAAATC T GCTCAATGAA ATTAAT ATTAAAT

Tro1  CGCC TGTTT ATCAA AAACA TGT CGTC -TTGATTTTATAT------ GATG TT T-AACC T GCTCAATGAT --TA-- TTTAAAT

Tro2  CGCC TGTTT ATCAA AAACA TGT CTTC -TTGATTTTATTT------ GAAG TC T-GGCC T GCTCAATGAT --TA-- ATTAAAT

Tro3  CGCC TGTTT ATCAA AAACA TCT CTTC -ATGAATTTATTT------ GAAG TC A-GGCC T GCTCAATGAG --T--- TTTAAAT

Tro4  CGCC TGTTT ATCAA AAACA T-T CTTA ---GGTTTTATTT------ GAAG TC T-A-CC T GCTCAATGAT --TA-- ATTAAAT

Ric1  ???? ????? ????? ????? ??? ???? ??????????????????? ???? ?? ?????? ? ?????????? ?????? ???????

Ric3  CGCC TGTTT ATCAA AAACA TGT CTTT -AAGTTTATATTT------ AAAG TC T-GACC T GCTCAATGAT ---TA- TTTAAAT

Ric4  CGCC TGTTT ATCAA AAACA TGT CTTT -AAGTTTTTTTTT------ AAAG TC T-GGCC T GCTCAATGAG -AAT-- TTTAAAT

Ric5  CGCC TGTTT ATCAA AAACA TGT CCTT -GAGAGTTTATTT------ AAGG TC T-GTCC T GCTCAATGAA TTTTTT TTTAAAT

Eur1  CGCC TGTTT ATCAA AAACA TGT CTTT -GAGAGTTTATTT------ AAAG TT T-GTCC T GCTCAATGAA -TTTA- TTTAAAT

Lop1  CACC TGTTT ATCAA AAACA TGT CTTT -AAGATTTATATTT----- AAAG TC T-AACC T GCTCAATGAA ---T-- TTTAAAT

Tet2  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGAGTTTATTA------ AAAG TC A-AACC T GCTCAATGAT -AATT- TTTAAAT

Iss1  CGCC TGTTT ATCAA AAACA TGT CTTT -GAGAGTTTATTT------ AAGG TC T-GACC T GCTCAGTGAT TTT--- ATTAAAT

Iss5  CGCC TGTTT ATCAA AAACA TGT CTTT -AAGAGTTTATTT------ AAAG TC T-GGCC T GCTCAATGAA TT---- TTTAAAT

Iss9  CGCC TGTTT ATCAA AAACA TGT CTTT -GAGGGTTTATTT------ AAAG TC T-GACC T GCTCAGTGAT -TAA-- ATTAAAT

Iss10 CGCC TGTTT ATCAA AAACA TGT CTTT -AAGTTGTTTATTT----- AAAG TC T-GGCC T GCTCAGTGAT TTTA-- ATTAAAT

Iss2  CGCC TGTTT AACAA AAACA TGT CTTG -TAGATTTTTTTA------ TAAG TC T-GTCC T GCTCAGTGAT ---T-- TTTAAAT

Iss11 CGCC TGTTT ATCAA AAACA TGT CTTG -TAGTTTTTTTTA------ TAAG TC T-ATCC T GCTCAGTGAT ---T-- TTTAAAT

Iss12 CGCC TGTTT ATCAA AAACA TGT CTTG -TAGTTTTTTTTA------ TAAG TC T-ATCC T GCTCAGTGAT ---T-- TTTAAAT

Iss3  CGCC TGTTT AACAA AAACA TGT CTTT -TAGTTTTTATTT------ AAGG TC A-AGCC T GCTCAATGAT ATATT- TTTAAAT

Iss14 CGCC TGTTT AACAA AAACA TGT CCTT -TAGTTTTTTTTT------ AAGG TC T-AGCC T GCTCAATGAT GTTAA- TTTAAAT

Iss4  CACC TGTTT ATCAA AAACA TGT CTTA -AAAAAATTTATTT----- TAAG TT T-AACC T GCTCAATGAA TTAT-- TTTAAAT

Iss6  CGCC TGTTT ATCAA AAACA TGT CTTA -ATGTTATTTATTT----- TAAG TC T-AGCC T GCTCAATGAA TTGT-- TTTAAAT

Iss7  CGCC TGTTT ATCAA AAACA TGT CTTA -ATGTTATTTATTT----- TAGG TC T-AGCC T GCTCAATGAA TTAT-- TTTAAAT

Iss8  CGCC TGTTT ATCAA AAACA TGT CTTG -TAGTTTTTTTTA------ TAAG TC T-ATCC T GCTCAATGAA GTGATT TTTAAAT

Cix2  CGCC TGTTT ATCAA AAACA TGT TCTT -TAGATTTATTA------- AAGA TT T-AGCC T GCTCTATGAT -TTAA- ATTAAAT

Cix3  CGCC TGTTT ATCAA AAACA TGT TCTT -TAGATTTTTTA------- AAGG TT T-AACC A GCTCTATGAT --GA-- TTTAAAT

Cix4  CGCC TGTTT ATCAA AAACA TGT TCTT -TAGATTTTTTTA------ AAGG TT T-AACC T GCTCTATGAT --TA-- TTTAAAT

Cix6  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGATTTTTTTA------ AAGG TT T-AACC T GCTCTATGAA --TA-- TTTAAAT

Del2  CGCC TGTTT ATCAA AAACA TTT CTTT -CAGGTATTATATTTATTG AAAG TT T-AACC T GCTCAATGAA -TT-T- TTTAAAT

Del3  TGAC TGTTT ATCAA AAACA TTT TCTT -TTGAT-TTATTTAA---- AAGG TA A-AACC T GCTCACTGAA -TTTT- TTTAAAT

Del4  TGCC TGTTT ATCAA AAACA TGT TCTT -TAGGTTTTATTT------ AAGG TC T-CGCC T GCTCAATGAT -TGAT- TTTAAAT

Del6  TGCC TGTTT ATCAA AAACA TGT TCTT -TAGGTTTTATTT------ AAGG TT T-CGCC T GCTCAATGAT -TAAT- TTTAAAT

Del7  CGCC TGTTT ATCAA AAACA TTT CTTT -CAGGTATTATATTTATTG AAAG TT T-AACC T GCTCAATGAA -TT-T- TTTAAAT

Fla5  CGCC TGTTT ATCAA AAACA TGT CTTT -TTGTTTATATTT------ AAAG TC T-GACC T GCTCAATGAA -TTA-- TTTAAAT

Mee2  TGAA TGTTT ATCAA AAACA TTT CTTC -ATGGGTTTTTTT------ GAAG TA T-TCCC T GCCCTATGAA -TTA-T TTTAAAT

Mee3  TGAA TGTTT ATCAA AAACA TTT CTTC -ATGTTTTTTTTT------ GAAG TA T-TCCC T GCTCTATGAA -TTAGT TTTAAAT

Mee6  TGAA TGTTT ATCAA AAACA TTT CTTC -ATGGGTTTTTTT------ GAAG TA T-TCCC T GCCCTATGAA -TTA-T TTTAAAT

Cerc  CGCC TGTTT ATCAA AAACA TGT TTTT -TAGTTTTTATTA------ AAAA AT TTAATC T GCCCAATGAT -TTT-- TTTTAAT

Cica  CGCC TGTTT ATCAA AAACA TGT CTTT -TAGATTATATTA------ AAAG TC T-AATC T GCCCAATGAT ---T-- TTTTAAT

Cic1  CGCC TGTTT AACAA AAACA TTT CTTT -TTGATTATATAA------ AAAG TA T-TTTC T GCCCAATGAT ATTTA- TTTAAAT

Cic2  CGCC TGTTT AACAA AAACA TTT CTTT -TTGCTTTTATA------- AAAG GT G-AGTC T GCCCTATGGG --AT-- ACTAAAT

Cic3  CGCC TGTTT AACAA AAACA TTT CTTT -TTGTGTGTTTA------- AAAG GT ATCTTC T GCCCTATGAT -TATT- ATTAAAT

Cic4  CGCC TGTTT AACAA AAACA TTT CTTT -TTAGTTTTTTTTA----- AAAG TA C-TTTC T GCCCTATGGT -TTGT- TCTAAAT

           <65a> <---> <65b> <-> <66a><-----------------> <66b><> <67a-> - <---68a--> <----> <--68b-

 

                                                                                                 158

Ful2  AGC CGCAGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTT GA CG

Ful3  AGC CGCAGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTT GA CG

Ful4  AGC CGCAGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GGGGTC TAGGATGAAG GGTTA GA CG

Ful5  AGC CGCAGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTT GA CG

Dic1  AGC CGCAGT AATTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG GA CG

Dic2  AGC CGCGGT AAATTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG AA CG

Dic3  ?GC CGCGGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG AA CG

Dic4  AGC CGCAGT AATTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG GA CG

Fla1  AGC CGCAGT ATATTG ACTGTG CGA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGTATGAAT GGTTG GA CA

Fla2  AGC CGCAGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GATTT GA CG

Fla3  AGC CGCAGT ATATTG ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGTATGAAT GGTTA GA CG

Fla4  AGC CGCAGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GATTT GA CG

Del8  AGC TGCAGT ATTTTA ACTGTA CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTT GA CA

Nog1  AGC CGCGGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG GA CG

Nog2  AGC CGCAGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGTATGAAT GGTTA AA CG

Ach1  AGC CGCGGT AATTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGGTT GA CA

Ach2  AGC CGCGGT AACTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTA GA CA

Der1  AGC CGCAGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTA GA CG

Der2  AGC CGCGGT ATTTTG ACCGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAA GGTTG GA CG

Der3  AGC CGCGGT ATATTG ACCGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TTGAATGAAT GATTT GA CG

Der4  AGC CGCAGG AATTTA ACTGTA CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT TGAGTC TTGTATGAAT GGTTT GA CG

Der9  AGC CGCGGT ATATTG ACCGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TTGAATGAAT GATTT GA CG

Tro1  AGC CGCGGT AATTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG GA CG

Tro2  AGC CGCGGT AATTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TAGAATGAAT GGTTG GA CG

Tro3  AGC CGCGGT AATTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GGGGTC TTGAATGAAT GGTTA GA CG

Tro4  AGC CGCGGT AATTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TAGAATGAAT GGTTG GG CG

Ric1  ??? ?????? ?????? ?????G CAA AGG TAGCATAGTA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTA GA CG

Ric3  AGC CGCAGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGTATGAAT GGTTG GA CG

Ric4  AGC CGCGGT ATTTTG ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTA GA CG

Ric5  AGC CGCGGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGTATGAAT GGTTA GA CG

Eur1  AGC CGCAGT ATTTTG ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAA GGTTA GA CG

Lop1  AGC CGCAGT ATTTTA ACTGTG TAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTG AA TG

Tet2  AGC CGCAGT ATTTTA ACTATA CAA AGG TAGCATAATC  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTA AA CG

Iss1  AGC CGCGGT ATATTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TAGAATGAAT GGTTG AA CG

Iss5  AGC CGCGGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGGATGAAT GGTTG AA CG

Iss9  AGC CGCGGT ATATTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GGGGTC TAGAATGAAT GGTTA AA CG

Iss10 AGC CGCGGT ATATTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGTATGAAT GGTTA AA CG

Iss2  AGC CGCAGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGATT CA CG

Iss11 AGC CGCAGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGATT CA CG

Iss12 AGC CGCAGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGATT CA CG

Iss3  AGC CGCGGT AAATTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGGATGAAT GGTTA GA CG

Iss14 AGC CGCGGT ATTTTG ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTA AA CG

Iss4  AGC CGCAGT AAATTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT AAGGTC TTGAATGAAT GGTTG GA TG

Iss6  AGC CGCAGT AATTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAAGTC TTGAATGAAA GGTTG GA TG

Iss7  AGC CGCAGT AATTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAAGTC TTGAATGAAA GGTTG GA TG

Iss8  AGC CGCAGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT AAAGTC TTGAATGAAT GGTTG GG TG

Cix2  GGC CGCAGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTTTCT TAATT AGGATC TGGAATGAAT GGTTG GA CA

Cix3  GGC CGCGGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTCT TAATT AGGGTC TTGAATGAAT GGTTG GA CG

Cix4  GGC CGCGGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTCT TAATT GGGGTC TTGAATGAAT GGTTG AA CG

Cix6  GGC CGCGGT ATTTTA ACTGTG CTA AGG TAGCATAATA  ATT  A GTCTCT TAATT AGGGTC TTGAATGAAT GGTTG GA CG

Del2  AGC TGCAGT AATTTA ACTGTA CAA AGG TAGCATAGTA  ATT  A GTCTTT TAATT GAGGTC TAGAATGAAT GGTTT AA CA

Del3  AGC CGCAGT ATTTTA ACTGTA CTA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TGGAATGAAT GGTTT AA CA

Del4  GGC CGCGGT AATTTA ACTGTG CTA AGG TAGCGTGATA  ATT  A GTCTTT TAATT GAGGTC TTGTATGAAT GGTTG GA CA

Del6  AGC CGCGGT AATTTA ACTGTG CTA AGG TAGCGTAATA  ATT  A GTCTTT TAATT GGGGTC TTGTATGAAT GGTTG GA CA

Del7  AGC TGCAGT AATTTA ACTGTA CAA AGG TAGCATAGTA  ATT  A GTCTTT TAATT GAGGTC TAGAATGAAT GGTTT AA CA

Fla5  AGC CGCAGT ATATTG ACTGTG CGA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TTGAATGAAT GGTTA AA CG

Mee2  GGC CGCGGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TTGAATGAAT GGGTT AA CA

Mee3  GGC CGCGGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TTGAATGAAT GGGTT AA CA

Mee6  GGC CGCGGT ATTTTA ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAGGTC TTGAATGAAT GGGTT AA CA

Cerc  GGC CGCAGT AATTTG ACTGTG CAA AGG TAGCATAATA  ATT  A GTCTTT TAATT AAAGTC TTGTATGAAT GATTG GA TG

Cica  GGC CGCGGT AAACTG ACCGTG CGA AGG TAGCATAATA  ATT  A GTCTTT TAATT GAAGGC TTGAATGAAT GATTG GA TG

Cic1  GGC CGCAGT ATACTA ACTGTG CAA AGG TAGCAAAATA  ATT  A GTTTCT TAATT AGAAGC TAGAATGAAT GGATA TA TG

Cic2  GGC TGCAGT ATTTTG ACTGTG CAA AGG TAGCATAGTA  ATT  A GTTTCT TAATT AGAAGC TGGTATGAAT GAATT AA TG

Cic3  GGC TGCAGT ATATTG ACTGTG CAA AGG TAGCATAGTA  ATT  A GTTTCT TAATT GGAAGC TTGTATGAAT GGATG TA TG

Cic4  GGC TGCAGT AGCTTG ACTGTG CAA AGG TAGCATAATA  ATT  A GTTTTT TAATT GAAGGC TGGTATGAAT GGAAA TT TG

      --> <69a-> <----> <-69b> <-><70a><--------> <70b> - <71a-> <---> <-71b> <--------> <67b> <> <-

 

 

                                                                                                 242

Ful2  AAAATTAT AC TTTATTTTTTTAATTTAA TTTGAATTTTATTT TTAAG TTAAAAAG CTTAA ATTTA GGAGAGGGACGA TAAGA CC

Ful3  AAAATTAT AC TTTATTTTTTTAATTTAA TTTGAATTTTATTT TTAAG TTAAAAAG CTTAA ATTTA GGAGAGGGACGA TAAGA CC

Ful4  AGAAGTAA AC TTTATTATCTTAATTTTA TTTGAATTTTATTT TTTAG TAAAAAAG CTAAA TTGTT GAAGAGGGACGA TAAGA CC

Ful5  AAAATTAT AC TTTATTTTTGTAATTTAA TTTGAATTTTATTT TTAAG TTAAAAAG CTTGA ATTTT GAAGAGGGACGA TAAGA CC

Dic1  AGAAATAT AC TTTATTTAATTGATTTT- TTTGAATTTTATTT TTTAG TTAAAAAG CTAAA ATTTT GAAGAGGGACGA TAAGA CC

Dic2  AAAAATTT AC TTTATTTTTTTTATTGT- TTTGAATTTTATTT TTTAG TCAAAAAG CTTAA ATATT AAAGAGGGACGA TAAGA CC

Dic3  AGAAATTT AC TTTATTTCATTTATTTT- TTTGAATTTTATAT TTTGG TTAAAAAG CTTAA ATGTT GAAGAGGGACGA AAAGA CC

Dic4  AGAAATAT AC TTTATTTAATTGATTTT- ATTGAATTTTATTT TTTAG TTAAAAAG CTAAA ATTTT GAAGAGGGACGA TAAGA CC

Fla1  AGGAAATT AC TTTATTTTTTTAATTTTT GTTGAATTTTAGAT TTTGG TTAAAATT CCAAA ATTTT TAAAAGGGACGA TAAGA CC

Fla2  AGGAATTA AC TTTATTTAAATTATTTT- TTTTAATTTTTGTT TTTGG TTAAAATA CCAAA ATTTT TATAAGGGACGA TAAGA CC

Fla3  AGGGAATT TC TTTATTTGGGGGAAGTTA ATTTAATTTATGTT TTTGG TTAAAATA CCAGG ATTTT TAAAAGGGACGA TAAGA CC

Fla4  AGGAATTA AC TTTATTTAAATTATTTT- TTTTAATTTTTGTT TTTGG TTAAAATA CCAAA ATTTT TATAAGGGACGA TAAGA CC

Del8  AGAAAATA AC TTTATTAATTAATCTTT- TTTCAATTTTATTT TTTAG TTAAAAAG CTAAA ATTTT TAAATGGGACGA TAAGA CC

Nog1  AGAAATTT TC TTTATTTTTTTTATTTTA TTTGAATTTTTATT TTTGG TTAAAATT CCAAA ATTTT TAAGAGGGACGA TAAGA CC

Nog2  AAAAATTT AC TTTATTTATTTTATTTT- TTTTAACTTTTATT TTTGG TTAAAATA CCAAA GTTTA TAAGAGGGACGA TAAGA CC

Ach1  AGGAAATA AC TTTATTTATTAA?????? ?????????????? ????? ???????? ????? ????? ???????????? ????? ??

Ach2  AGAAAATA AC TTTATTAATTTAAATTT- TTTGAATTTAATTT TTAAG TAAAAAAG CTTAA GTATT GAAGAGGGACGA TAAGA CC

Der1  AGGAGACA AC TTTATTAATTTTTAT--- ATTTAATTTAATTT TTTAG TTAAAAAG CTAAA ATTTT TAAGAGGGACGA TAAGA CC

Der2  AGAAAATA GC TTTATTTTTTGTAATTT- TTTGAATTTAATTA TCAAG TTAAAAAG CTTGA ATTTA TTAAAGGGACGA TAAGA CC

Der3  AAAAATAA TC TATTTTAATTTTATTTTT TTTGAATTTAAATT TTTTG TTAAAATG CTAAA ATTTA ATAGAGGGACGA TAAGA CC

Der4  AAAAATAA TC TTTATTTTTTTTTT---- TTTTAATTTTAATT TTGTG TTAAAATG CATAA ATTTT TAAGAGGGACGA TAAGA CC

Der9  AAAAATAA TC TATTTTAATTTTATTTTT TTTGAATTTAAATT TTTTG TTAAAATG CTAAA ATTTA ATAGAGGGACGA TAAGA CC

Tro1  AGAAAAAT AC TTTATTTTTTAAAATTT- ATTTAATTTAATTT TATTG TTAAAACG CAATA ATTAT TTGAAGTGACGA TAAGA CC

Tro2  AAAAAAAT GC TTTATTTGATTTATTT-- ATTTAATTTAACTT CATTG TTAAAATG CAATG ATTTT TTAAAGTGACGA TAAGA CC

Tro3  AGAAATAA AC TTTATTTTTTTAAATA-- ATTAAATTTAATTT TTTTG TTAAAAAG CTTAA ATTTA TTGAAGTGACGA TAAGA CC

Tro4  AAAAAAAT GC TTTATTTGATTTATTT-- ATTTAATTTAACTT CATTG TTAAAATG CAATG ATTTT TTAAAGTGACGA TAAGA CC

Ric1  AGAAACAA AC TTTATTGAAATAAATTT- TTTTAATTTTGATT TTGAG TTAGAATA CTTAA ATTTT AAAGGGGGACGA TAAGA CC

Ric3  AAAAACAT AC TTTATTAATTTTTAAT-- TTTTAATTTTTACT TTAAG TTAAAATA CTTAA ATATT AAAGTGGGACGA TAAGA CC

Ric4  AGAAATAA CC TTTATTTATTTTATTAA- ATTTAATTTTTTTT TTAAG TTAAAATA CTTAA ATTTT TAAGAGGGACGA TAAGA CC

Ric5  AGAAATAT AC TTTATTTGAAAAATTTT- TTTGAATTTTAAAT TTAAG TTAAAATT CTTAA ATTTT TAAGAGGGACGA TAAGA CC

Eur1  AAGAATAA AC TTTATTAATTTGATTAA- TTTGAATTTTAATT TCAAG TAAAAAT- CTT?A ATTTT AATAAGGGACGA TAAGA CC

Lop1  AGAAACTT GT TTTATTAAATAAATTTT- TTTGAATTTAAGTT TCTAG TTAAAATT CTAGA ATAAT TAAGAGGGACGA TAAGA CC

Tet2  AGAAATAA AC TTTATTTTTTAAATTAT- TTTGAATTTTTTTT TATAG TAAAAAAA CTATA ATTTG TAAGTGGGACGA TAAGA CC

Iss1  AGAAATTT AC TTTATTGTTTTTATTTTT TTTGAATTTTATTT ATTGG TTAAAAAG CCATT ATGTT TAAGAGGGACGA TAAGA CC

Iss5  AGAAATTT TC TTTATTAATTTAATTCA- ATTGAATTTTATTT TTTTG TTAAAAAG CTTAA ATTTA AAAGAGGGACGA TAAGA CC

Iss9  AGAAATTT AC TTTATTGTTTTTATTTTT TTTGAATTTTATTT ATTGG TTAAAAAG CCATT ATGTT TAAGAGGGACGA TAAGA CC

Iss10 AGAAAATT AC TTTATTTTATTTATTTTT TTTGAACTTTACTT TTTGG TTAAAAGG CCAAT ATTTT AAAGAGGGACGA TAAGA CC

Iss2  AGGAATAT AC TTTATTAATTTAATTTTT TTTGAATTTTATAT TTTTG TTAAAAAG CTTAA GTTAA AAAGAGTGACGA TAAGA CC

Iss11 AGGAATAT AC TTTATTAATTTAATTTTT TTTGAATTTTATAT TTTTG TTAAAAAG CTTAA GTTAT AAAGAGTGACGA TAAGA CC

Iss12 AGGAATAT AC TTTATTAATTTAATTTTT TTTGAATTTTATAT TTTTG TTAAAAAG CTTAA GTTAT AAAGAGTGACGA TAAGA CC

Iss3  AGAAATAA AC TTTATTTAATTTATTTTT TTTGAATTTGTTTT TTTAG TTAAAATT CTAAA ATGTT TTAGAGGGACGA CAAGA CC

Iss14 AGATATGA AC TTTATTTAATTTATTTT- TTTGAATTTGTTTT TTTAG TTAAAATT CTAAA ATGTT TAAGTGGGACGA CAAGA CC

Iss4  AAAAAAAA AC TTTATTAATTTTTTTTGT ATTGAATTTTATTT TTAAT TTAAAAAG ATTAA ATTTT TGAAGGGGACGA TAAGA CC

Iss6  AAAAATCT AC TTTATTAATATTTTTTT- ATTGAATTTTATTT TTAAT TTAAAAAG ATTAA ATTTA TTTAGGGGACGA TAAGA CC

Iss7  AAAAAACT AC TTTATTAATATTTTTTT- ATTGAATTTTATTT TTGAT TTAAAAAG ATTAA ATTTA TTTAGGGGACGA TAAGA CC

Iss8  AAAAAATT TC TTTATTAGTTTTATTAAT TGAAATTTATTTTT AATTT AAAAAGAT TAATT GTTTA AAAGGGGCACGA TAAGA CC

Cix2  GGAGGTAA AC TTTATTATTTTTTTT--- TTTGAATTTAATTT TTTAG TTAAAAAG CTTAA ATTTT TAATTGTGACGA TAAGA CC

Cix3  AGAAAATA AC TTTATTAAAATAAATTTT TCTTAATTTAATAT TTTTG TTAAAAAG CTAAA ATGAT TAAGTGGGACGA TAAGA CC

Cix4  AGAAAATA AC TTTATTAAAATGATTTTT TTTTAATTTAATAT TTTTG TTAAAAAG CTTAA ATAAT TAAGTGGGACGA TAAGA CC

Cix6  AGAAAACA AC TTTATTAATAAAATTTTT TTTTAATTTAATTT TTTAG TTAAAAAG CTTAA ATAAT TAAGTGGGACGA TAAGA CC

Del2  AGAAGTAA GC TTTATTAATTCAATTA-- AGTGAAATTTATTT TTTAG TTAAAAAG CTAAA ATTTA AAAATGGGACGA TAAGA CC

Del3  AGGTGAAA AC TTTCTTATTTAAATTAC- TTTGAATTTAATTT TTTAG TTAAAAAG CTAAA ATTTT TAAATGGGACGA TAAGA CC

Del4  AGAAGATT TC TTTATTTAATTTATTT-- TTTTAATTTAATTT TGTAG TTAAAAAG CTTCA ATTTT TAATTGGGACGA TAAGA CC

Del6  AGAAGATT TC TTTATTTAATTTATTT-- TTTTAATTTAATTT TTTAG TTAAAAAG CTTTA ATTAT TAATTGGGACGA TAAGA CC

Del7  AGAAGTAA GC TTTATTAATTCAATTA-- AGTGAAATTTATTT TTTAG TTAAAAAG CTAAA ATTTA AAAATGGGACGA TAAGA CC

Fla5  AAGAATAA AC TTTATTTAATTAATTTT- TTTTAATTTTTGTT TTTGG TTAAAATA CCAGA ATATT TAAAAGGGACGA TAAGA CC

Mee2  AAAAAATA TC TTTATTAAATTAAATTT- TTTTAAGTTAATTT TAAAG TTAAAATT CTTTA ATAAT TAAGGGGGACGA TAAGA CC

Mee3  AAATAATA GC TTTATTAAATTAAATTT- TTTGAAATTAATTT TAAAG TTAAAATT CTTTA ATTAT TAAGGGGGACGA TAAGA CC

Mee6  AAAAAATA TC TTTATTAAATTAAATTT- TTTTAAGTTAATTT TAAAG TTAAAATT CTTTA ATAAT TAAGGGGGACGA TAAGA CC

Cerc  AGAAATTA TC TTTCTTTATTATTTTA-- ACTGAATTTGATAT TTAGG TGAAAAAG CTTAA ATTTT TCTGTGGGACGA TAAGA CC

Cica  AAAAGCTA TC TTTATTAATATAAAAA-- TTTTAATTTAACAT TTAAG TAAAAAGG CTTAA ATATT TTAGTGGGACGA GAAGA CC

Cic1  TGATATAT AT TTTATTAGGTTTAAAA-- TTAAAATTTTAATT TTGGG TTAAAATG CTCAA TTTCT TTTTAGGGACGA TAAGA CC

Cic2  AGAAATAT AT TTTATTAATTTTAATA-- TTGAAATTTAAATT TTAAG TGAAAATG CTTAA AGGTT TTCTTGGGACGA TAAGA CC

Cic3  AGAAATAT GT TTTATTAATTTTACTA-- TTAAAAATTAAATT TTAAG TCAAAATG CTTAA TTATT TTTCTGGGACGA TCAGA CC

Cic4  AGATACAT GT TTTATTTGATTTAAATT- TATAAATTTAAACT TTAAG TTAAAATG CTTAA GTGAA TTCTTGGGACGA TAAGA CC

      --64b--> <> <------61b-------> <------------> <72a> <------> <72b> <---> <----73a---> <---> <-

 

 

                                                                                                 328

Ful2  CTATAGATCTT T ATTTTATTTTTGATTTTTTTTTTT-- TAGTTAATT ---TTAAAAATTTTAATTAATAAAT TT TGTTG GGG TGAC

Ful3  CTATAGATCTT T ATTTTATTTTTGATTTTTTTTTTT-- TAGTTTATT --TTTAAAATTTTTATTAATTAAAT TT TGTTG GGG TGAC

Ful4  CTATAGATCTT T ATTTTTTTCCATTTTTTTTTATTT-- TAGTTTATT --TTAGATTTTTTTTGGTTATAAAT TT TGCTG GGG TGGT

Ful5  CTATAGATCTT T ATTTTATTTTTGATTTTTTTTTTT-- TAGTTTATT --TTTAAAATTTTTATGAATTAAAT TT TGTTG GGG TGAC

Dic1  CTATAGATCTT A ATTTAATTTATTTAAGTTTTTTTT-- TTGTTGTCT --TTTTTAATTTATTTAATTTAAAT TT GGTTG GGG CGAC

Dic2  CTATAGATCTT G AATTTATTAAATTAATTTATTTTT-- TTGTTGTTT --TTATTAATTTATTTTAATTAATT TT AGTTG GGG CGAT

Dic3  CTATAGATCTT A AATTTTTTATTATAATTTATTTTT-- TTGTGGTTT --TTTTTAATTTGAAGTAATCAATT TT GGTTG GGG CGAT

Dic4  CTATAGATCTT A ATTTAATTTATTTAAGATAAAAT--- TTGTTGTAT --TTTTTAATTTATTTATTTTAAAT TT GGTTG GGG CGAC

Fla1  CTATAGATCTT T ATAATTCTTATTTTTTTGTTTTTT-- TTGTTGTTT -TAAGATTATTTATTTATGATTTAT TT AGTTG GGG TGAT

Fla2  CTATAGATCTT T AAAAAAAATATTTATTTTTCTTTT-- TTGTTGTAT --TTTAAAATTAAGATATTTTTTTT TT TGTTG GGG TGAC

Fla3  CTATAGATCTT T ATAATATTTTATTATTTAGGTTTT-- TTGTTGTAT --TTTTTTATTTAGTATTTTTTTAT TT AGTTG GGG TGAT

Fla4  CTATAGATCTT T AAAAAAAATATTTATTTTTCTTTT-- TTGTTGTAT --TTTAAAATTAAGATATTTTTTTT TT TGTTG GGG TGAC

Del8  CTATAGATCTT A AAATTTTTTATATTTGAAATAATTT- TATGGGTTA ATTTAAATTTTGTTTGGGAAAATTT TT TGTTG GGG TGAC

Nog1  CTATAGATCTT T AAAATAATTTATTTATTATTTTTTT- TTGTTGTAT --TTATTTTTTATTTATATTTTTTT TT TGTTG GGG TGAT

Nog2  CTATAGATCTT T AAAATTGTTTTTTTTTCTTTTTTT-- TTGATGTGT -TTTTAGGAGAATTATTTCATTTTT TT TGTTG GGG TGAT

Ach1  ??????????? ? ?????????????????????????? ????????? ??????TTTTTTAGGAAGTAAAAAT TT TGTTG GGG TGAC

Ach2  CTATAGATCTT T ATTATTAATTTTTATAAGATGTTTAG TAGATTTTT -----TTTTTTGTGAAAATTATTAA TT TGCTG GGG TGGC

Der1  CTATAGATCTT T ATTATTGATTTTTTATTTTTG----- TTAATTTTT -----TATTTAATTTAATTAATAAT TT TATTG GGG TGAT

Der2  CTATAGATCTT T ATTATTTAGGTTTTATATTATTA--- TTATTGTTT --TTGATTTTTATAATTTAAATAAT TT TATTG GGG TGAT

Der3  CTATAGATCTT T ATTATATTTATAATATTTATTTTTT- TTATTGTTT ---TTTATTATATTATAAATTTAAT TT TATTG GGG TGAT

Der4  CTAAAGATCTT T ATTTAATAATTATTTTAATTTTT--- TTTAA---- --TTTTTTTATTTTAATTTTTAAAT TT GGTTG GGG TGAT

Der9  CTATAGATCTT T ATTATATTTATAATATTTATTTTTT- TTATTGTTT ---TTTATTATATTATAAATTTAAT TT TATTG GGG TGAT

Tro1  CTATAGATCTT T ATTTTATTTTTTTTTTTTTTCTTT-- TTGATGGTT -TTTATTTAAATTTAATTATAAAAT TT TGTTG GGG TGAC

Tro2  CTATAGATCTT T ATTTTTTTTATATTTTTTTTTTTT-- TTGTTGATT -TTATATAATTTTAGTTTAAAAAAT TT TGTTG GGG TGAT

Tro3  CTATAGATCTT A ATTAATTTTTAATATTTTTTTTT--- TTGTTGTAT ----TTTATTGTTATTAAATTTAAT TT TGTTG GGG TGAT

Tro4  CTATAGATCTT T ATTTTTTT-CTATTTTTTTTTTT--- TTGTTGATT -TTATGTATTTTTAGTTTAAAAAAT TT TGTTG GGG TGAT

Ric1  CTATAGATCTT A AAAATATTTACTTTTTATTTTTTTT- TTGTTGTTT --TTTATTTAAATATTATTTATTTT TT AGTTG GGG TGAT

Ric3  CTATAGATCTT A ATAATTTTATTATTTATTTTTTTT-- TTGTTGTTT -TATTAATTTATTTATTAATTTTTT TT AGTTG GGG TGAT

Ric4  CTATAGATCTT G AGAATTTTTTTTTTTTTTTTTTTT-- TTGTTGTTT --TTTATTTGGAATTTTAATTTTCT TT AGTTG GGG TGAT

Ric5  CTATAGATCTT A ATAATTATTTTTTTTTGTTTTTTT-- TTGTTGTGA --TTTAATATATTTAAAATTTTTTT TT TGTTG GGG TGAT

Eur1  CTATAGATCTT - ATAATAATTTTTTTTTTAAAATTT-- TAGTTGTTT -ATAAATTTAAAATAAAAATTTTAA TT TGTTG GGG TGAT

Lop1  CTATAGATCTT A ATAATATAAATAATTAATAAAATTT- TAGATTATA ---TATTGGTAATTTTTTTTTTTAT TT TGTTG GGG TGAT

Tet2  CTATAGATCTT T ATAAATTAAAAGTTTATTTTTTTT-- TTGAATTAT -TTTTATATTAAATTTTTAATTTAT TT TGTTG GGG TGAT

Iss1  CTGTAGATCTT T ATAATTTAAATAATTTTTATTTTTTT GTGTTTGTT --TTTTTTGTTATTATTTGATTTAT TT TGTTG GGG TGAC

Iss5  CTATAGATCTT T ATAATAAAATTTTTTTTTTTTTTTT- TTGTAGTTT ---TTTTAAAAATTAATTTAATTAT TT TGTTG GGG TGAT

Iss9  CTGTAGATCTT T ATAATTTAAATAATTTTTATTTTT-- TTGTTTGTT --TTTTTTGTTATTATTTGATTTAT TT TGTTG GGG TGAC

Iss10 CTGTAGATCTT T ATAATTTAAACTATTTTTATTTTT-- TTGTTTGTC --TTTTTTGTAATTTTTTATTTTAT TT TGTTG GGG TGAT

Iss2  CTATAGATCTT T AAAATTTTTTTTTTTTAATTTTTT-- TTGTTGTTT --ATTGGTTATTTAAATTTATTTTT TT TGTTG GGG AGAT

Iss11 CTATAGATCTT T AAAATTTTTTTTTTTTAATTTTTT-- TTGTTGTTT --ATTGTTTATTTAAATTAATTTTT TT TGTTG GGG AGAT

Iss12 CTATAGATCTT T AAAATTTTTTTTTTTTAATTTTTT-- TTGTTGTTT --ATTGTTTATTTAAATTAATTTTT TT TGTTG GGG AGAT

Iss3  CTATAGATCTA A AGAAGATTTTATTTTATTTTTTTT-- TTGTTGTTT ---TTTATTTAATATAAATTTTTTT TT TGTTG GGG TGAC

Iss14 CTATAGATCTT T AAATAGGTTTATTTTATTTTTTTT-- TTGTTGTTT ---TTTAATTAATATTAATCTTTTT TT TGTTG GGG TGAT

Iss4  CTATAGATCTT A AAAATATTATATTTATATATATTT-- TTGTTGTTT ATATTTATAAAAATTATATTATTTT TT TGTTG GGG TGAC

Iss6  CTATAGATCTT A AAAATATTTTTTTTTTATTAATTT-- TTGTTGTTA AATTTTTTGTAAA-AATATTATTTT TT TGTTG GGG TGAC

Iss7  CTATAGATCTT A AAAATATTTTTTTTTTATTAATTT-- TTGTTGTTA AATTTTTTGAAATTAATATTATTTT TT TGTTG GGG TGAC

Iss8  CTATAGATCTT T AAAATTTTTTATATATTTATTTTT-- TTGTTGTTT -TTAATAAATTTTTAAATAGATTTT TT TATTG GGG TGAT

Cix2  CTATAGATCTT A AAAATATTTTTATATAAATTTTT--- TTGTGGTAT -TCTTTTTTTTGTAGGAATTATTTT TT AGTTG GGG TGAT

Cix3  CTGTAGATCTT T ATAATTAATTAATATAAAATTTT--- TGGTTGTAT --TATTTTTTTTTTTAGTTAATTAT TT TGTTG GGG TGAC

Cix4  CTGTAGATCTT T ATAATTAATTAAAATAATTTTTT--- TGGTTGATA --TTTTTTTTTTTTTAATTAATTTT TT TGTTG GGG TGCC

Cix6  CTGTAGATCTT A AAATATTTAATAAGTTCATTTTT--- TTGTTGTTA TTATGTGTTTCTTATTTAAACTTTT TT TATTG GGG TGAT

Del2  CTTTAGATCTG T AAATTTTAATTGAATAATAATATTA- ATGTGGATT ---TAATTTTTATTAAATAAAATTT TT TGTTG GGG TGAC

Del3  CTATAGATCTT A AATTTTTTTTTATCTTAATTTCT--- TGGTTAATA AAATGTTAAAATTTTAAAAAAAATT TT TGTTG GGG TGAC

Del4  CTGTAGATCTT T AAAATTGGTTTTTTTAATTTTTT--- TTGTTTATT --TAATTTTTATTTTTATTAATTTT TT GGTTG GGG TGAT

Del6  CTGTAGATCTT T AAAATTGATTTTAATAATTTTTT--- TTGTTTATT --TAGTTTTTATTTTTATTAATTTT TT GGTTG GGG TGAT

Del7  CTTTAGATCTT T AAATTTTAATTGAATAATAATATTA- ATGTGGATT ---TAATTTTTATTAAATAAAATTT TT TGTTG GGG TGAC

Fla5  CTATAGATCTT A AAAATATTTTTTTATTTTTTTTTT-- TTGTTGTTT -TTTAAAAAATTTATCTTTTATTTT TT TGTTG GGG TGAT

Mee2  CTATAGATCTT A AAAAAATAAATTTCTTATT------- TGGTTTTTT --TATTTATAAATTAATTTATTTAT TT AGTTG GGG AGAT

Mee3  CTATAGATCTT A AAAAATAAATTTTTATATT------- TAGTTTTTT ---TTATTTTTTAAAATTTATTTTT TT AATTG GGG AGAT

Mee6  CTATAGATCTT A AAAAAATAAATTTCTTATT------- TGGTTTTTT --TATTTATAAATTAATTTATTTAT TT AGTTG GGG AGAT

Cerc  CTATAGATCTT T AAAATTATTTTTTTTTAATATTTTT- AGAATTTAT -ATTTCTTATTAAATTATTAATTTT TT TGTTG GGG TGAT

Cica  CTATAGAATTT G AAATCTATAATTATTAAATTTATT-- TAGATAAAT -ATTAAATTTAATAATTATGAATTT TT TGTTG GGG TGAC

Cic1  CTATAGAACTT T ACTAAATATAATTTAGTTGGTTTT-- TTTTTTATA ----TAAATTTATTAAATTTTATTT TT AGTTG GGG TGAC

Cic2  CTATAGAACTT T ACATC-TAATTTCTAGTTGATTTT-- AACTTTTAT --ACTCTTCTAGGGTTTTTAGATGA TT CGCTG GGG TGGT

Cic3  CTATAGAACTT T ATATTACCTAGTTATAGTTTATTT-- TTGATTATA -ATACTATCTTAATATTACAGTACT TT TGCTG GGG TGGT

Cic4  CTATAGAACTT T ACATTGTTTGTTTTTAATGAATTTT- TACATTAAA --TATTATATTAATTTAAGACTTGT TT TGCTG GGG TGGT

      ---74a----> - <-----------75a----------> <-------> <----------75b----------> <> <80a> <-> <80b

 

 

                                                                                                 408

Ful2  A AAAAAA -AATTTAATC TTTTTT TT TT  TTTTACATTTTTTTAT- GA  --ATATTTGATC CTTA -T- TTTG ATTAAAAGATT

Ful3  A AATAAA -ATTTTAATC TTTATT TT TT  TTTTACATTTTTTTAT- GA  --ATATTTGATC CTTA TT- TTTG ATTAAAAGATT

Ful4  A GATAAA -ATTTTAATC TTTATT TT TT  TTTTACATTTCTTTAT- GT  --GATTTTGATC CTTT TT- TTTG ATTATAAGATT

Ful5  A AATAAA -ATTTTAATC TTTATT TT TT  TTTTACATTTTTTTAT- GT  --CTGTTTGATC CTTT AT- TTTG ATTAAAAGATT

Dic1  A AATAAA -TTTTTAATC TTT-TT TT TT  TTTTACAATTATTTTTG GT  ----TTTTGATC TTTT TT- ATTG ATTATAAGATG

Dic2  G AATAAA -ATTTTAATC TTTTTT TT TT  TTTTACA-TTATATTTG GA  ---TATTTGATC CTTT TTT TTTG ATTAAAAGATA

Dic3  A ATTAAA -CATTTAAAC CTTTTT TT TT  TTTTACATTTATTTTT- GA  --TTTATTGATC TTTA TT- TTTG ATTAAAAGATT

Dic4  A AATAAA -TTTTTAAAC CTTTTT TT TT  TTTTACAATTATTTTTG GT  ----TTTTGATC TTTT TT- ATTG ATTATAAGATG

Fla1  T AATAAA -AATTTTATC TTTATT TA TT  TTTTACATTTTTTTTTG GT  ---TATTTGATC TTAA ATT TTGG ATTATAAGAAA

Fla2  G TTTAAA -ATTTTTGAC TTTAAA TT TT  TTTTACA-TTTTTTTT- GT  -TTAATTTGATC TTGA ATT TTTG ATTATATGATT

Fla3  T AATAAA -ATTTTAAAC TTTATT TC TT  TTTTACAATTATTTTTG GT  ---TTTTTGATC CTTA AAT TTAG ATTATAAGATT

Fla4  G TTTAAA -ATTTTTGAC TTTAAA TT TT  TTTTACA-TTTTTTTT- GT  -TTAATTTGATC TTGA ATT TTTG ATTATATGATT

Del8  A TATAAA TTTTTAAAAC TTTAAA AT TT  TTTTACATTTATAGAT- GT  --ATAAATGATC CAAA CTT TTTG ATTATTAAAAT

Nog1  A GATAAA -AACTTAAAC TTTTTT TT AT  TATTACATTTATTTATG GT  --TTTTTTGATC CTTA ATT TTTG ATTTCAAGATT

Nog2  A TATAAA -ATTTTAAAC TTTATA TT TT  TTTTACATTTATTTTTG GT  --TATATTGATC CTTT TTT TTTG ATTTTAAGATT

Ach1  A TATAAA -ATTTTAAAC TTTATT TT TT  CTTTGCTTTTATTTTTG GT  --TTATTTGATC CTTA AT- TTTG GATTTTAGATT

Ach2  A GGTAAA -ATATTTATC TTTATT TT TT  TTTTACATTTATTTATG TT  --TAGTTTGATC CAGA ATT TTTG ATTGAAAGATT

Der1  A ATTAAA -TTTTTTTAC TTTTTT TT TT  TAGAACAATTTTTTATG GG  ---AATTTGATC TTTT AT- TTTG ATTTTAAGAAA

Der2  A ATTAAA -AATTTAAAC TTTTTT TT TT  ATTAACATTTATTT--- GT  --GAATTAAGTA TTTT TTT TAAG AGAATAAGATA

Der3  A ATTAAA -ATTTTTATC TTTTTT AT TT  TATTACAATAGTTTTTG GT  -TAAATGATCCT AAAT AAT TTAG ATTAATAGATA

Der4  A AATAAA -TATTTAATC TTTTTT TT TT  TTTTACATTTTTTTATG GT  ---TATTTGAGT ---T TTT A--- AATTAAAAGAA

Der9  A ATTAAA -ATTTTTATC TTTTTT AT TT  TATTACAATAGTTTTTG GT  -TAAATGATCCT AAAT AAT TTAG ATTAATAGATA

Tro1  A TTTAAA -ATTTTAATC TTTATT TA TT  TTTTACATTTTTTTTT- GT  --GTTTTTGAAC CATA ATT TTTG TTTATTAGAAA

Tro2  A ATTAAA -AATTTTATC TTTAAT TT TT  TTTTACATTTATTTAT- GT  -TTAGTTTGAAC CTTA ATT TTTG TTTAAAAGAAT

Tro3  T GCTAAA -ATTTTAATC TTTAGT TT TT  TATTACATTTATTTTTG GT  --TAGTTTGAAC TTAT AT- TTTG TTTGTAAGATA

Tro4  A ATTAAA -AATTTTATC TTTAAT TT TT  TTTTACATTTATTTAT- GT  -TTAGTTTGAAC CTTA ATT TTTG TTTAAAAGAAC

Ric1  T ATTAAA -ATTTTAAAC TTTGAT TT AT  TTTTACATTTAATTTTG GC  TTATTTTTGATC CTTC TTT TTTG ATTATAAGATA

Ric3  T ACTAAA -ATTTTAAAC TTTAGT TT AT  TTTTACATTTGTTATT- GT  -TTTTTTTGATC TTTT TT- TTTG ATTATAAGATA

Ric4  T ACTAAA -AATTTAAAC TTTAGT TT AT  TTTTACATTTATTATT- GT  -ATTTATTGATC CTTT TT- TTTG ATTATAAGATA

Ric5  T ACCAAA -AATTTAAAC TTTGGT TT AT  TTTTACATAAAATTTAT GT  -ATTTTTTGATC CTTT TT- TTTG ATTATAAGAAA

Eur1  A ACTAAA -ATTTTAAAC TTTAGT TT TT  TTTTACATAAATAAAT- GT  -ATAAATTGATC TTTA TT- ATTG ATTATAAGATT

Lop1  A GCTAAA -ATTTTAAAC TTTAGT TT TT  TTTAACAATAATTTTT- GT  -GTAATTTGATC TTTT AT- TTTG ATTATAAGATA

Tet2  A AATAAA -ATTTTAAAC TTTTTT TT TT  TTTTACATTTATAAAT- GA  -TTAATTTGATC CTTT AAT ATTG AAAATAAGATC

Iss1  A GTTAAA -ATTTTAAAC TTTAGT TT TT  TTTTACATATATTTATG GT  --TTGTTAGATC TTTA ATT TTTG ATATCAAGATC

Iss5  A GCTAAA -ATATTAAAC TTTAGT TT AT  TTTTACAATTTTTTTTG GT  --TTTTTTGATC CTTT -TT ATTG ATATAAAGAAG

Iss9  A GTTAAA -ATTTTAAAC TTTAGT TT TT  TTTTACATACATTTATG GT  --TTACTAGATC TTTA ATT TTTG ATATTAAGATT

Iss10 A ACTAAA -ATTTTAAAC TTTAGT TT TT  TTTTACATACATAGATG GT  --TTTTTTGATC TTTA ATT TTGG ATATTAAGATT

Iss2  A GATAAA -ATTTTAAAC TTTATT TT TT  TTTTACTTTTTTT-AT- GA  --ATTTTTGATC CTTA ATT TTGG ATTATAAGAAT

Iss11 A GATAAA -AATTTTAAC TTTATT TT TT  TTTTACTTTTTTTG-TG GT  ---TTTTTGATC CTTA ATT TTGG ATTATAAGAAT

Iss12 A GATAAA -AATTTTAAC TTTATT TT TT  TTTTACTTTTTTTG-TG GT  ---TTTTTGATC CTTA ATT TTGG ATTATAAGAAT

Iss3  A ATTAAA -ATTTTAAAC TTTAAT TT TT  TTTCACATTTTTTTAT- GT  --TTATTTGATC CTTC TTT GTTG ATTAAAAGATA

Iss14 A GTTAAA -ATTTTTAAC TTTAAT TT TT  TTTTTCATTTATTTAT- GA  --ATTTTTGATC TTTT TTT GATG ATTAAAAGATA

Iss4  A AATAAA -AATTTAAAC TTTATT TT TT  TTTTACATTTGTTTAT- GA  -AATTTTTGATC CTTT -T- TTGG ATAAAAAGAAA

Iss6  A AATAAA -AATTTAAAC TTTATT TA TT  TTTTACATTTATAGTT- GA  --TTTTTTGATC CTAA GTT TTGG ATAGTAAGATA

Iss7  A AATAAA -AATTTAAAC TTTATT TA TT  TTTTACATTTATAGAT- GA  --TTTTTTGATC CTAG -TT TTGG ATAATAAGATA

Iss8  A ATCAAA -AATTTTAAC TTTAAT TT AT  TTTAACATTAATTTAT- GA  -TTTTTTTGATC CTTA TTT TTGG AGAAAAAGAAC

Cix2  T AAAGAA -ATTTTAAAC TTTTTT TT TT  TTTTACATTGATATAT- GT  --TTTTTTGATC TTTA AT- TTTG ATTAAAAGAGT

Cix3  A AATAAA -TTTTTTAAC TTTTTT TA TT  ATATACATTATTTTTTG GT  ---TTTTTGATC CTTA AT- TTTG ATTAAAAGAAA

Cix4  A GTAAAA -TTTTTTAAC CTTTTT TT TT  TAGGACAATAATTTTTT TT  GGGTTTTTGATC CTTT CAT TTTG ATTAAAAGAAA

Cix6  A AATAAA TTTTTTAAAC TTTTTT TT AT  ATTTACATTTATTTATG AA  --AAATTTGATC CTTT TAT TTTG ATTAAAAGAAA

Del2  A GCTAAA ATCTTAAATC TTTAGT AT TT  TACACCATTTATTTAT- GT  --ATATTTGAAC CTTA ATT TAAG GTTATTAAAAT

Del3  A TTAAAA TTTTTAAATC TTTTAA TC AT  TTTTACATAAATAAAT- GT  --ATATTTGATC TTAA CAT TTAG ATTAAAAAATT

Del4  T TTTAAA TTTTTATTAC TTTATT TT TT  TTTTACATTTATTTAT- GT  --TTAATTGATC TTTA ATT TTTG ATTTTTAGATT

Del6  T TTTAAA TTTTTATTAC TTTATT TT TT  TTTTACATTTATTTAT- GT  --TTAATTGATC TTTA ATT TTTG ATTTTTAGATT

Del7  A GCTAAA ATCTTAAATC TTTAGT AT TT  TACACCATTTATTTAT- GT  --ATATTTGAAC CTTA ATT TAGG GTTATTAAAAT

Fla5  A AATAAA ATTTTTGACC TTTATT TT TT  TTTTACATTTATTTTT- GG  -TTTTATTGATC CTTT ATT TTGG ATTATAAGATA

Mee2  T TTTAAA -AATTTAAAC TTTATT TT TT  TTTTACATCAATTTAT- GT  -TTATATTGATC CAGT ATT TCTG ATTATAAGATA

Mee3  T TATAAA -AAATTAAAC TTTATA TT TT  TTTTACATTAATTGTT- GT  TTTATATTGATC CATT TTT AGTG ATTTAAAGATA

Mee6  T TTTAAA -AATTTAAAC TTTATT TT TT  TTTTACATCAATTTAT- GT  -TTATATTGATC CAGT ATT TCTG ATTATAAGATA

Cerc  A AATTAA -TTAATATAA CTTTTT TA TG  TTTTTTCATTAATTTAT GT  --TTTTATGATC TTTT ATT AATG ATTAATAGATT

Cica  A TTAAAA TATTAATAAC TTTTAA TT AA  ATTTTCATAGATAAAT- GA  --TTATTAGATC CAAT TTT TTTG ATTTTAAGACT

Cic1  A GTTAAA -TTTT-AA-C TTTAAT TT TG  ATAATCATTATTTTAT- GT  -AGTTTTTGATC CATA TA- TATG ATAATAAGATA

Cic2  G GATAAA -TTT-AAA-C TTTATT AT TT  TATATCAATTATTTTT- GT  -TTGTTTAGATC CTTT TGG AATC -----AAGAAT

Cic3  G GATAAA -TTT-AAA-C TTTATA TA AT  AATATCAATTATTAAT- GA  -ATGTTAAGATC TTAC AAC TTAG ATTAAAAGATA

Cic4  G AATAAA -TTT-AAA-C TTTATT AA TA  AAAATCA-TTATAAAT- GT  -TTGTGTTGATC TGTT ATT AAGA ATT--AAGAAT

      > <81a-> <--------> <-81b> <><82a><---------------><82b><----------> <88a><-> <88b><--------->

 

 

                                                                                                 495

Ful2  AAGATACCTTAGGG ATAACAG CGTTATAAATCTGGAGA GTTC-TTA TTGAT-AGATTTGTTTGCG ACCTCGATG TTGGATTAAAA GT

Ful3  AAGATACCTTAGGG ATAACAG CGTTATAAATCTGGAGA GTTC-TTA TTGAT-AGATTTGTTTGCG ACCTCGATG TTGGATTAAAA GT

Ful4  AAGATACCTTAGGG ATAACAG CGTGATAATTCTGGAGA GTTC-ATA TTGAT-AGAGTTGTTTGCG ACCTCGATG TTGGATTAAAA AT

Ful5  AAGATACCTTAGGG ATAACAG CGTTATAAATCTGGAGA GTTC-TTA TTGAT-AGATTTGTTTGCG ACCTCGATG TTGGATTAAAA GT

Dic1  AAGATACCTTAGGG ATAACAG CGTTATTTAGTTGGATA GTTC-ATA TTGAT-AACTAAGATTGCG ACCTCGATG TTGGATTAAAA TT

Dic2  AAGATACCTTAGGG ATAACAG CGTTATTTAGTTGGAGA GTTC-TTA TTGAT-AATTAAGTTTGCG ACCTCGATG TTGGATTAATA AT

Dic3  TAGATACCTTAGGG ATAACAG CGTTATTTAATTGGAGA GTTC-TAA TCAAT-AATTAAGATTGCG ACCTCGATG TTGGATTAAAA TT

Dic4  AAGATACCTTAGGG ATAACAG CGTTATTTAGTTGGATA GTTC-ATA TTGAT-AGCTAAGATTGCG ACCTCGATG TTGAATTAAAA TT

Fla1  AAGATACCTTAGGG ATAACAG CATAATAATTCTGGAGA GTTC-TAA TCGAT-AGACTTGTTTGTG ACCTCGATG TTGGATTAAAA TT

Fla2  AAGATACCTTAGGG ATAACAG CATAATTTATTTGGAGA GTTC-AAA TCGAT-AATTAAGTTTGTG ACCTCGATG TTGGATTAAAA TT

Fla3  AAGATACCTTAGGG ATAACAG CATAATTTACCTGGAGA GTTC-TTA TTGAT-AGGTTTGTTTGTG ACCTCGATG TTGGATTAAAA TT

Fla4  AAGATACCTTAGGG ATAACAG CATAATTTATTTGGAGA GTTC-AAA TCGAT-AATTAAGTTTGTG ACCTCGATG TTGAATTAAAA TT

Del8  AAGATACCTTAGGG ATAACAG CGTAATAAATTTGTATA GTAC-ACA TAAAT-AAGTTTGTTTACG ACCTCGATG TTGAATTAATA TT

Nog1  AAGATACCTTAGGG ATAACAG CGTTATATATCTGGAGA GTTC-TAA TTGAT-AGATATGTTTGCG ACCTCGATG TTGGATTAATT TA

Nog2  AAGATACCTTAGGG ATAACAG CGTTATATGTTTGGAGA GTTC-TTA TCGAT-AAATATGTTTGCG ACCTCGATG TTGGATTAAAA TT

Ach1  AAGATACCTTAGGG ATAACAG CGTTATATATTTGGAGA GTTC-AAA TTGAT-AAATATGTTTGCG ACCTCGATG TTGGATTAAAT TT

Ach2  AAGATACCTTAGGG ATAACAG CGTAATATATTTGGAGA GTTC-TTA TTGAT-AAATATGTTTGCG ACCTCGATG TTGGATTAAAA TT

Der1  AAGATACCTTAGGG ATAACAG CGTTATTTAATTGGAGA GTTC-ATA TTGAT-AATTAAGATTGCG ACCTCGATG TTGGATTAATA AT

Der2  AAGATACCTTAGGG ATAACAG CGTTATATAACTGGAGA GTTC-TTA TTGAT-AGTTATGTTTGCG ACCTCGATG TTGGATTAAAA TT

Der3  TAGATACCTTAGGG ATAACAG CGTTATATAATTGGAAA GTTC-TTA TTGAT-AATTATGATTGCG ACCTCGATG TTGGATTAAAG TT

Der4  AAGATACCTTAGGG ATAACAG CGTTATAAAATTGGATA GTTC-AAA TTGAT-AATTTTGTTTGCG ACCTCGATG TTGGATTAATA AA

Der9  TAGATACCTTAGGG ATAACAG CGTTATATAATTGGAAA GTTC-TTA TTGAT-AATTATGATTGCG ACCTCGATG TTGGATTAAAG TT

Tro1  AAGATACCTTAGGG ATAACAG CGTAATATATTTGGATA GTTC-TTA TTGAT-AAATATTTTTGCG ACCTCGATG TTGGATTAAAA TT

Tro2  AAGATACCTTAGGG ATAACAG CGTAATTTATTTGGATA GTTC-ATA TTGAT-AAATATATTTGCG ACCTCGATG TTGGATTAATA TT

Tro3  AAGATACCTTAGGG ATAACAG CGTAATTTGTTTGGATA GTTC-AAA TTGAT-AAATAAGTTTGCG ACCTCGATG TTGGATTAAAT TT

Tro4  AAGATACCTTAGGG ATAACAG CGTAATTTATTTGGATA GTTC-ATA TTGAT-AAATATAATTGCG ACCTCGATG TTGGATTAAAA TT

Ric1  AAGATACCTTAGGG ATAACAG CGTTATTCACTTGGAGA GTTC-TAA TCGAT-AAGTGAGTTTGCG ACCTCGATG TTGGATTAAAT TT

Ric3  AAGATACCTTAGGG ATAACAG CGTTATTTGTTTGGAGA GTTC-TTA TTGAT-AAATGAGTTTGCG ACCTCGATG TTGGATTAAAT TT

Ric4  AAGATACCTTAGGG ATAACAG CGTAATTTATTTGGAGA GTTC-TTA TTGAT-AAATAAGTTTGCG ACCTCGATG TTGGATTAAGT TT

Ric5  TAGATACCTTAGGG ATAACAG CGTTATTTAATTGGAGA GTTC-TTA TCTAT-AATTAAGTTTGCG ACCTCGATG TTGGATTAAAT TT

Eur1  AAGATACCTTAGGG ATAACAG CGTTATATATTTGGAGA GTTC-TAA TTGAT-AAATATGATTGCG ACCTCGATG TTGGATTAAAA TT

Lop1  AAGATACCTTAGGG ATAACAG CGTTATTCATTTGAAGA GTCC-ATA TCTGT-AAATGAGTTTGCG ACCTCGATG TTGGATTAAAA TA

Tet2  CAGATACCTTAGGG ATAACAG CGTTATATATTTGGAGA GTTC-AAA TTGAT-AAATATGTTTGCG ACCTCGATG TTGGATTAAAA TT

Iss1  AAGATACCTCAGGG ATAACAG CGTTATAGATTTGGAGA GTTC-TTA TTGAT-AAATTTGTTTGCG ACCTCGATG TTGGATTAAAG TT

Iss5  AAGATACCTTAGGG ATAACAG CGTTATATAGTTGGAGA GTTC-AAA TTGAT-AATTATGTTTGCG ACCTCGATG TTGGATTAAAG TT

Iss9  AAGATACCTCAGGG ATAACAG CGTTATATATTTGGAGA GTTC-TTA TCGAT-AAATTTGTTTGCG ACCTCGATG TTGGATTAAAG TT

Iss10 AAGATACCTCAGGG ATAACAG CGTTATAAATTTGGAGA GTTC-TTA TTGAT-AAATTTGTTTGCG ACCTCGATG TTGGATTAAAA TT

Iss2  AAGATACCTTAGGG ATAACAG CGTTATAAATCTGGAAA GTTC-TAA TTGAT-AGATTTGTTTGCG ACCTCGATG TTGGATTAATT TG

Iss11 AAGATACCTTAGGG ATAACAG CGTTATAAGTCTGGAAA GTTC-AAA TTTAT-AGATTTGTTTGCG ACCTCGATG TTGGATTAACT TA

Iss12 AAGATACCTTAGGG ATAACAG CGTTATAAGTCTGGAAA GTTC-AAA TTTAT-AGATTTGTTTGCG ACCTCGATG TTGAATTAATT TG

Iss3  AAGATACCTTAGGG ATAACAG CGTAATATAATTGGATA GTTC-AAA TTGAT-AATTATGTTTGCG ACCTCGATG TTGGATTAAAC TT

Iss14 AAGATACCTTAGGG ATAACAG CGTAATATAATTGGATA GTTC-AAA TTGAT-AATTATGTTTGCG ACCTCGATG TTGGATTAAAT TT

Iss4  AAGATACCTTAGGG ATAACAG CGTTATATTTCTGGAGA GTTC-TAA TCAAT-AGAATTGTTTGCG ACCTCGATG TTGGATTAACT TA

Iss6  AAGATACCTTAGGG ATAACAG CGTTATAATTCTGGAGA GTTC-TAA TCAAT-AGAATTGTTTGCG ACCTCGATG TTGGATTAATT TT

Iss7  AAGATACCTTAGGG ATAACAG CGTTATAATTCTGGAGA GTTC-TAA TCAAT-AGAATTGTTTGCG ACCTCGATG TTGGATTAATT TA

Iss8  AAGATACCTTAGGG ATAACAG CGTTATAAATTTGGAGA GTTC-TAA TCAAT-AAATTTGTTTGCG ACCTCGATG TTGGATTAACT TA

Cix2  AAGATACCTTAGGG ATAACAG CGTAATATATTTGGAGA GTTC-TTA TTGAT-AAGTATGTTTGCG ACCTCGATG TTGGATTAAGA TT

Cix3  AAGATACCTCAGGG ATAACAG CGTTATACATTTGGAAA GTTC-ATA TTGAT-AAATGTGTTTGCG ACCTCGATG TTGAATTAAGT TT

Cix4  AAGATACCTCAGGG ATAACAG CGTTATACATTTGGAAA GTTCTATA TTGAT-AAATGTGTTTGCG ACCTCGATG TTGAATTAAAG TT

Cix6  AAGATACCTCAGGG ATAACAG CGTTATATATTTGGAGA GTTC-TTA TTAAT-AAATATGTTTGCG ACCTCGATG TTGGATTAAAA TT

Del2  AAGATACCTTAGGG ATAACAG CATAATAAATTTGTAAA GTTC-ATA TATAT-AAATTTGATTGTG ACCTCGATG TTGAATTAATC TA

Del3  AAGATACCTTAGGG ATAACAG CGTAATAAATTTGAATA GTTC-ATA TTGAT-AAATTTGTTTACG ACCTCGATG TTGAATTAATA TA

Del4  AAGATACCTCAGGG ATAACAG CGTTTATTTTTTGGAAA GTTC-TTA TTGAT-AAATTAAATTGCG ACCTCGATG TTGGATTAATA TA

Del6  AAGATACCTCAGGG ATAACAG CGTTTATTTTTTGGAAA GTTC-TTA TTGAT-AAATTAAATTGCG ACCTCGATG TTGGATTAATA TA

Del7  AAGATACCTTAGGG ATAACAG CATAATAAATTTGCAAA GTTC-ATA TATAT-AAATTTGATTGTG ACCTCGATG TTGAATTAATC TA

Fla5  AAGATACCTTAGGG ATAACAG CATAATATAGTTGGAGA GTTC-AAA TCGAT-AATTATGTTTGTG ACCTCGATG TTGGATTAAGA ??

Mee2  AAGATACCTTAGGG ATAACAG CGTAATAAATTTGGGGA GTTC-ATA TTTAT-AAATTTGTTTGCG ACCTCGATG TTGAATTAAAA TT

Mee3  AAGATACCTTAGGG ATAACAG CGTAATAAATTTGGGGA GTTC-ATA TTTAT-AAATTTGTTTGCG ACCACGATG TTGAATTAAAA TT

Mee6  AAGATACCTTAGGG ATAACAG CGTAATAAATTTGGGGA GTTC-ATA TTTAT-AAATTTGTTTGCG ACCTCGATG TTGAATTAAAA TT

Cerc  AAGTTACCTTAGGG ATAACAG CGTGATTTATTTGGAAA GTTC-ATA TTTAT-AAATAAGTTTGCG ACCTCGATG TTGGATTAAGA TT

Cica  AAATTACCTTAGGG ATAACAG CGTTATTATTTTAAAAA GTTC-TTA TTGATAAAATTAGATTGCG ACCTCGATG ??????????? ??

Cic1  AAGTTACCTTAGGG ATAACAG CGTAATTTTAGTGGGGA GTTC-ACA TCTAT-ACTAAATTTTGCG ACCTCGATG TTGAATTAAGA TA

Cic2  TAGTTACCTTAGGG ATAACAG CGTAATTTTATTGGCGA GTTC-ATA TCTAT-A-TTAATTTTGCG ACCTCGATG TTGAATTAAGG AA

Cic3  AAGTTACCTTAGGG ATAACAG CGTAATTATTATGGGAA GGTTATAA TTTAT-A-TAAGTTTTGCG ACCTGAATG TTGAATTAAGG AA

Cic4  AAGTTACCTTAGGG ATAACAG CGTAATTACAATGGAAA GTTC-ATA TTTAT-ATTGATTTTTGCG ACCTCGATG TTGAATTAAGG TA

      <-----74b----> <-----> <------89a------> <------> <-------89b-------> <-------> <---90a---> <-

 

                                                                                        569

Ful2  -- TTCTGTGGG GTAGGT TTTACAATA CT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Ful3  -- TTCTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Ful4  -- TTCTGTGAG GCAGGT TTTACAGTT TA GGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Ful5  -- TTCTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Dic1  -- TTCTGTGGG GTAGGT TTTACAGTT AA GGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Dic2  -- TTCTGTGGG GTAGGT TTCACAGTT TA AGGTC TGTTC GACTT ATAAA TTTTACATGATCTGA GTTCA GACCGG

Dic3  -- TTCTGTGGG GTAGGT TTTACAGTT TA AGGTC TGTTC GACTT ATAAA TTTTACATGATCTGA GTTCA GACCGG

Dic4  -- TTCTGTGGA GTAGG? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Fla1  T- ATTTGTGGG GTAGTT TTTACAATT TT AGGTC TGTTC ????? ????? ??????????????? ????? ??????

Fla2  TA AATTGTGGG GTAGGT TTTACAATT TT AGGTC TGTTC GACC? ????? ??????????????? ????? ??????

Fla3  T- ATTTGTGGG GTAGTT TTTACAATT CT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Fla4  TA AATTGTGGG GTAGGT TTTACAATT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Del8  A- ATTTTTGGA GCAGAA GCTAGAATT TT AGGTC TATAC GACTT TTTAA TATTACATGATCTGA GTTCA GACCGG

Nog1  A- AGCTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Nog2  T- TGCTATGGG GTAGGT TTTATAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Ach1  -- AATTGTGGG GTAGG? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Ach2  T- GTTTGTGGG GTAGGT TTTACAAAT CT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Der1  -- AATTGTAAG GTAGGT TTTACAATG TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Der2  T- ATTTATGGG GTAGGT TTCATAATA TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Der3  -- AATTGTGAG GTAGGT TTCACAATG TT AAGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Der4  -- ATGTATGAG GTAGGT TTCATATTA TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Der9  -- AATTGTGAG GTAGGT TTCACAATG TT AAGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Tro1  T- TTTTGTGGG GTAGGT TTCACAATA TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Tro2  T- TTTTGTGGG GAAGGT TTCTCAATA AT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Tro3  T- GTTTATGGG GTAGGT TTCATAATT CT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Tro4  T- TTTTGTGGG GTAGGT TTCACAATA AT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Ric1  T- GTTTGAGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Ric3  T- GTTTGTGGG GTAGGT TTTACAATT CT AGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Ric4  T- GTTTGTGGG GTAGGT TTCACAATT CT AGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Ric5  T- GTTTGTGGG GTAGGT TTTACAAAA CT AGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Eur1  T- GGTTGTGGG GTAGGT TTTACAATT CT AGGTC TGT?? ????? ????? ??????????????? ????? ??????

Lop1  A- ATTTGTGGG GTAGGT TCTACAATT TT AGGTC TGTTC GACC? ????? ??????????????? ????? ??????

Tet2  T- AATTGTGGG GTAGGT TTTACAAGT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Iss1  T- TACTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Iss5  T- TACTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Iss9  T- TACTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA CTTTACATGATCTGA GTTCA GACCGG

Iss10 T- TACTGTGGG GTAGGT TTTACAGTT TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Iss2  T- TACTGTGGG GTAGGT TTTACAGTT GT GGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Iss11 A- TTTTGTGGG GTAAGT TTTACAATT TT AGGTC TGTTC GACTT TAAAT AGTTACATGATCTGA GTTCA GACCGG

Iss12 T- TACTGTGGG GTAGGT TTTACAGTT GT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Iss3  T- ATTTGTGGG GCAGTT TTTACAATC TT AGGTC TGTTC GACTT TAAAA GTTTACATGATCTGA GTTCA GACCGG

Iss14 T- ATTTGTGGG GCAGTT TTTACAATT TT AGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Iss4  A- ATTTGTGGG GTAGAT TTTACAGTT TT AGGTC TGTTC GAC?? ????? ??????????????? ????? ??????

Iss6  A- ATTTGTGGG GTAGAT TTTACAATT TT AGGTC TGTTC GACTT TAAAA AATTACATGATCTGA GTTCA GACCGG

Iss7  A- ATTTGTGGG GTAGAT TTTACAATT TT AGGTC TGTTC GACTT TAAAA AATTACATGATCTGA GTTCA GACCGG

Iss8  A- TTTTGTGGG GTAGG? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Cix2  A- ATAATTGGG GCAGAA GATTATTTA TT AGGTC TGTTC GACTT TAAAA TCTTACATGATCTGA GTTCA GACCGG

Cix3  T- AATTGTGGA GCAGAA TATACAATA TT AGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Cix4  T- AATTGTGGA GCAGAA TATACAATG TT AGGTC TGTTC GACTT TAAAA ATTTACATGATCTGA GTTCA GACCGG

Cix6  T- AATTGTGGA GCAGAA TATTCAAAG TT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Del2  G- AATTCTGGA GCAAAA AA??????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Del3  A- AATTTTAGA GCAGAA ACTTAAAGT TT AGGTC TGTTC GACTT TAAAT ATTTACATGATCTGA GTTCG AGCCGG

Del4  A- GTTTTTGGT GTAGAT GATAATTAA CT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Del6  A- GTTTTTGGT GTAGAT GATAATTAA CT AGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Del7  G- AATTCTGG- ?????? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Fla5  ?? ????????? ?????? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Mee2  T- TGGGAGGGG GAAGGT TTTTTCTAT AT TGG?? ????? ????? ????? ??????????????? ????? ??????

Mee3  T- TAGGAAGGG GAAGGT TTTTTCTAT AT TGGTC TG??? ????? ????? ??????????????? ????? ??????

Mee6  T- TGGGAGGGG GAAGGT TTTTTCTAT AT TGGTC TGTTC GACTT TAAAA TTTTACATGATCTGA GTTCA GACCGG

Cerc  A- GAATAAGGG GTAGGT TTTTTA??? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Cica  ?? ????????? ?????? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Cic1  A- AAAAAAGAA GCAG?? ????????? ?? ????? ????? ????? ????? ??????????????? ????? ??????

Cic2  A- GATTAAGAA GCAGAA TTCTTAAGT CT AACTT TTAAA TCCT? ????? ??????????????? ????? ??????

Cic3  -- TAGTTTGAG TCTGTT CGACTTTTA AA TTCTT ACTTG ATCT? ????? ??????????????? ????? ??????

Cic4  AA TTTTTAGAG TCTGTT CGACTTTTA AA TCCCT ACTTG ATCT? ????? ??????????????? ????? ??????

      -> <--91a--> <----> <--91b--> <> <92a> <---> <92b> <---> <-----90b-----> <---> <93a->