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Table 1. Summary of data relative to samples analysed and accession numbers for the long-wavelength opsin (LWO), elongation factor la (EF-1a) and the mitochondrial DNA sequences.

Table 1 Summary of data relative to samples analysed and accession numbers for the long-wavelength opsin (LWO), elongation factor la (EF-1a) and the mitochondrial DNA sequences.

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Fig. 1. The basic life cycle in Fordina species without temporary gall. Fd, fundatrix; Fg, fundatrigeniae; Vg, virginoparae (exules); Sxp, sexuparae; 3, male; 2 ov, oviparae; ap, apterae; al, alatae. In brackets a number is given to each distinct morph. Morphs developing inside a gall in the primary host are enclosed by an ellipse. Note that the hatching of the sexual eggs laid by the oviparae in the previous spring may coincide with the new egg laying by sexuals produced in the current season.
Fig. 1. The basic life cycle in Fordina species without temporary gall. Fd, fundatrix; Fg, fundatrigeniae; Vg, virginoparae (exules); Sxp, sexuparae; 3, male; 2 ov, oviparae; ap, apterae; al, alatae. In brackets a number is given to each distinct morph. Morphs developing inside a gall in the primary host are enclosed by an ellipse. Note that the hatching of the sexual eggs laid by the oviparae in the previous spring may coincide with the new egg laying by sexuals produced in the current season.
Two species of this genus have been cited in the Iberian Peninsula: P. cimiciformis von Heyden and P. donisthorpei Theobald. Although P. cimiciformis induces characteristic fusiform-flat green galls on the edge of leaflets of P. terebinthus (Fig. 2), the galls of P. donisthorpei are unknown. Sexuparae of both species have six segmented antennae, but can be distinguished because P. donisthorpei has antennae with the fourth segment longer than the third (shorter in P. cimiciformis) and a striated dorsal cuticle (grid- like in P. cimiciformis) (see descriptions in Pérez Hidalgo & Nieto Nafria, 2003). Wingless exules of P. donisthorpei were described as having the typical Fordina 5-segmented anten- nae, whereas P. cimiciformis was the only species of Fordina whose wingless exules were described as having 6-segmented antennae. Most aphid taxonomists have not questioned the taxonomy of the species Paracletus cimiciformis, as_ its morphs were characterized perfectly, and its life cycle appar- ently is well known (Heie, 1980; Blackman & Eastop, 1994, 2000, 2006; Remaudiére & Remaudiére, 1997; Pérez Hidalgo & Nieto Nafria, 2003). However, Roberti (1983, 1993) ex- pressed his doubts, proposing the existence of two different wingless exule ‘forms’ in this species: ‘form A’, with the usual 5-segmented antennae, eyes composed of just the triommatidium, straight hind tibiae and round-shaped body; and ‘form B’, with 6-segmented antennae, eyes composed of several ommatidia in addition to the triomma- tidium, curved hind tibiae and flat-shaped body. The latter would be the ones von Heyden used to describe the genus Paracletus and the species P. cimiciformis and those cur- rently accepted by most taxonomists as the true P. cimici- formis exules. Moreover, Roberti (1983, 1993) suggested that his proposed ‘form A’ would correspond to what had been known as Forda rotunda Theobald, described as an One taxonomical problem studied in this work has been the possible synonymy between Forda marginata Koch and Forda riccobonii (Stefani). The former produces galls on Pistacia terebinthus, whereas the latter induces them on Pistacia atlantica. Definitive galls clearly differ (Fig. 2), but both are produced on the edge of leaflets. The fundatrigeniae inside the galls, although similar, can be distinguished by a few mor- phological characters (Blackman & Eastop, 1994; Perez Hidalgo & Nieto Nafria, 2003). In contrast, the wingless exules of the two species are indistinguishable (Pérez Hidalgo & Nieto Nafria, 2003). However, the last statement is based on limited data as, contrary to the situation in Forda marginata in which the morphs from the secondary host are well known, the exules of Forda riccobonii, and of other species in the genus, remain poorly known. This led Remaudiére (1999) to create the ‘marginata group’, including four (or six) species of the genus Forda: F. marginata, F. orientalis George (= kaussari Davatchi & Remaudiére), F. pawlowae Mordvilko Fig. 2. Galls of (A) Forda marginata and (C) Paracletus cimiciformis on Pistacia terebinthus and of (B) Forda riccobonii (definitive) on Pistacit atlantica.
Two species of this genus have been cited in the Iberian Peninsula: P. cimiciformis von Heyden and P. donisthorpei Theobald. Although P. cimiciformis induces characteristic fusiform-flat green galls on the edge of leaflets of P. terebinthus (Fig. 2), the galls of P. donisthorpei are unknown. Sexuparae of both species have six segmented antennae, but can be distinguished because P. donisthorpei has antennae with the fourth segment longer than the third (shorter in P. cimiciformis) and a striated dorsal cuticle (grid- like in P. cimiciformis) (see descriptions in Pérez Hidalgo & Nieto Nafria, 2003). Wingless exules of P. donisthorpei were described as having the typical Fordina 5-segmented anten- nae, whereas P. cimiciformis was the only species of Fordina whose wingless exules were described as having 6-segmented antennae. Most aphid taxonomists have not questioned the taxonomy of the species Paracletus cimiciformis, as_ its morphs were characterized perfectly, and its life cycle appar- ently is well known (Heie, 1980; Blackman & Eastop, 1994, 2000, 2006; Remaudiére & Remaudiére, 1997; Pérez Hidalgo & Nieto Nafria, 2003). However, Roberti (1983, 1993) ex- pressed his doubts, proposing the existence of two different wingless exule ‘forms’ in this species: ‘form A’, with the usual 5-segmented antennae, eyes composed of just the triommatidium, straight hind tibiae and round-shaped body; and ‘form B’, with 6-segmented antennae, eyes composed of several ommatidia in addition to the triomma- tidium, curved hind tibiae and flat-shaped body. The latter would be the ones von Heyden used to describe the genus Paracletus and the species P. cimiciformis and those cur- rently accepted by most taxonomists as the true P. cimici- formis exules. Moreover, Roberti (1983, 1993) suggested that his proposed ‘form A’ would correspond to what had been known as Forda rotunda Theobald, described as an One taxonomical problem studied in this work has been the possible synonymy between Forda marginata Koch and Forda riccobonii (Stefani). The former produces galls on Pistacia terebinthus, whereas the latter induces them on Pistacia atlantica. Definitive galls clearly differ (Fig. 2), but both are produced on the edge of leaflets. The fundatrigeniae inside the galls, although similar, can be distinguished by a few mor- phological characters (Blackman & Eastop, 1994; Perez Hidalgo & Nieto Nafria, 2003). In contrast, the wingless exules of the two species are indistinguishable (Pérez Hidalgo & Nieto Nafria, 2003). However, the last statement is based on limited data as, contrary to the situation in Forda marginata in which the morphs from the secondary host are well known, the exules of Forda riccobonii, and of other species in the genus, remain poorly known. This led Remaudiére (1999) to create the ‘marginata group’, including four (or six) species of the genus Forda: F. marginata, F. orientalis George (= kaussari Davatchi & Remaudiére), F. pawlowae Mordvilko Fig. 2. Galls of (A) Forda marginata and (C) Paracletus cimiciformis on Pistacia terebinthus and of (B) Forda riccobonii (definitive) on Pistacit atlantica.
“Species names according to Remaudiére & Remaudiére (1997) and Pérez Hidalgo & Nieto Nafria (2003). ’Code for the various samples analysed for the Forda and Paracletus species. Underlined samples were used for morphological characterization. The species/ samples used in the global phylogenetic analysis of the Iberian Fordini are shaded (name and accession number of gene/s used). “Indicates the original source of aphids from that sample. Poaceae species associated to ant nests were not identified. 4S, Spain; Por, Portugal. °Morph included in the molecular analysis; fg, fundatrigeniae obtained from galls; ex, exules either from transfer experiments * or directly collected from the field. SExules from the transfer experiment of sample P. cimiciformis Le5062 displayed morphs of both 5- and 6-segmented antennae. An individual of each morph was used for molecular analysis, yielding identical sequences both for the LWO gene and for the mitochondrial fragment. /Samples with more than one accession number contained more than one allele. Table 1. Continued.
“Species names according to Remaudiére & Remaudiére (1997) and Pérez Hidalgo & Nieto Nafria (2003). ’Code for the various samples analysed for the Forda and Paracletus species. Underlined samples were used for morphological characterization. The species/ samples used in the global phylogenetic analysis of the Iberian Fordini are shaded (name and accession number of gene/s used). “Indicates the original source of aphids from that sample. Poaceae species associated to ant nests were not identified. 4S, Spain; Por, Portugal. °Morph included in the molecular analysis; fg, fundatrigeniae obtained from galls; ex, exules either from transfer experiments * or directly collected from the field. SExules from the transfer experiment of sample P. cimiciformis Le5062 displayed morphs of both 5- and 6-segmented antennae. An individual of each morph was used for molecular analysis, yielding identical sequences both for the LWO gene and for the mitochondrial fragment. /Samples with more than one accession number contained more than one allele. Table 1. Continued.
Table 2. Primers used for amplification (PCR) and sequencing (seq) of samples analysed. “From Moran et al. (1999).
Table 2. Primers used for amplification (PCR) and sequencing (seq) of samples analysed. “From Moran et al. (1999).
Fig. 3. (A) Maximum-likelihood tree obtained after the analysis of the long-wavelength opsin (LWO) gene for the Iberian species of the tribe Fordini. The same topology was obtained after the analysis of the mitochondrial fragment when third codon positions of the gene encoding subunit 6 of the F-ATPase were excluded, except for the inclusion of G. setulosa (added to the tree with a dashed branch) in this analysis. Bootstrap values (given as a percentage) for the LWO analysis are shown before the slash and for the mitochondrial fragment analysis after the slash, when higher than 50. The primary host used by each species in the Iberian Peninsula is indicated beside the tree. The definitive gall type and capacity according to Inbar et a/. (2004) are also indicated. Four capacity classes have been used depending on the number of aphids inside the galls: low, fewer than 100 aphids per gall; moderate, a few hundred; high, up to 1000; very high, thousands. (B) ML tree obtained for the EF-1a gene for the species indicated. Bootstrap values are indicated on the branches when higher that 50.
Fig. 3. (A) Maximum-likelihood tree obtained after the analysis of the long-wavelength opsin (LWO) gene for the Iberian species of the tribe Fordini. The same topology was obtained after the analysis of the mitochondrial fragment when third codon positions of the gene encoding subunit 6 of the F-ATPase were excluded, except for the inclusion of G. setulosa (added to the tree with a dashed branch) in this analysis. Bootstrap values (given as a percentage) for the LWO analysis are shown before the slash and for the mitochondrial fragment analysis after the slash, when higher than 50. The primary host used by each species in the Iberian Peninsula is indicated beside the tree. The definitive gall type and capacity according to Inbar et a/. (2004) are also indicated. Four capacity classes have been used depending on the number of aphids inside the galls: low, fewer than 100 aphids per gall; moderate, a few hundred; high, up to 1000; very high, thousands. (B) ML tree obtained for the EF-1a gene for the species indicated. Bootstrap values are indicated on the branches when higher that 50.
Fig. 4. Principal component analysis of the morphometric varia- bles significantly different between the wingless exules of F. margin- ata (*) and those of F. riccobonii (A). Factors | and 2, representing 57.01 and 18.12%, respectively, of character variation, are shown.
Fig. 4. Principal component analysis of the morphometric varia- bles significantly different between the wingless exules of F. margin- ata (*) and those of F. riccobonii (A). Factors | and 2, representing 57.01 and 18.12%, respectively, of character variation, are shown.
Fig. 5. (A) Maximum-likelihood (ML) tree from the analysis of the long-wavelength opsin gene sequences belonging to species of the genera Forda and Paracletus. The same topology was obtained in the maximum-parsimony (MP) analysis with the codification of the gaps (see text). For some samples two alleles were sequenced, indicated as al and a2. Bootstrap values (as a percentage) are indicated when higher than 50 (ML/MP). (B) Topology obtained in the ML analysis of the mitochondrial sequences, with bootstrap values shown when higher than 50. Species names are abbreviated with the first letters of the generic and specific names. Morph analysed is indicated as follows: fg, fundatrigeniae; ex, exule; sx, sexupara. The two P. cimiciformis hybrid samples (see text) are highlighted. Identical sequences obtained from exules with 6- and 5-segmented antennae from sample Le5062 are indicated as exS/ex6.
Fig. 5. (A) Maximum-likelihood (ML) tree from the analysis of the long-wavelength opsin gene sequences belonging to species of the genera Forda and Paracletus. The same topology was obtained in the maximum-parsimony (MP) analysis with the codification of the gaps (see text). For some samples two alleles were sequenced, indicated as al and a2. Bootstrap values (as a percentage) are indicated when higher than 50 (ML/MP). (B) Topology obtained in the ML analysis of the mitochondrial sequences, with bootstrap values shown when higher than 50. Species names are abbreviated with the first letters of the generic and specific names. Morph analysed is indicated as follows: fg, fundatrigeniae; ex, exule; sx, sexupara. The two P. cimiciformis hybrid samples (see text) are highlighted. Identical sequences obtained from exules with 6- and 5-segmented antennae from sample Le5062 are indicated as exS/ex6.
Table 3. Average genetic distances“ among Forda (F) and Paracletus (P) species/clades’. “Maximum-likelihood distances using TrN+I+G and HKY+G models for the mitochondrial DNA fragment (upper right) and long-wavelength opsin (LWO) gene (lower-left), respectively; main diagonal, intra-specific/-clade distances for the LWO/mtDNA sequences. °f, formicaria; m, marginata; r, riccobonii; 1 and 2 are the two Paracletus clades (see text). “Average distances between genera Forda and Paracletus for the LWO/mtDNA sequences.
Table 3. Average genetic distances“ among Forda (F) and Paracletus (P) species/clades’. “Maximum-likelihood distances using TrN+I+G and HKY+G models for the mitochondrial DNA fragment (upper right) and long-wavelength opsin (LWO) gene (lower-left), respectively; main diagonal, intra-specific/-clade distances for the LWO/mtDNA sequences. °f, formicaria; m, marginata; r, riccobonii; 1 and 2 are the two Paracletus clades (see text). “Average distances between genera Forda and Paracletus for the LWO/mtDNA sequences.
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