Aphis polii, Barjadze, Shalva, Blackman, Roger L. & Ӧzdemir, Işil, 2015

Barjadze, Shalva, Blackman, Roger L. & Ӧzdemir, Işil, 2015, A new species of Aphis Linnaeus (Hemiptera, Aphididae) living on Teucrium polium L. (Lamiaceae), Zootaxa 4057 (2), pp. 248-256 : 249-255

Aphis polii sp. n.

( Figure 2 , Table 1 )

Material examined. Holotype and paratypes. Although no description has been published, 15 slides of the new species from 3 localities were labelled as “ paratypes by F. Leclant (FL) and G. Remaudière (GR). We have retained this designation, and selected an apt. from one of these samples as holotype. Holotype: apt., coll. no. FL 5092, GR 18130, numbered 1 on slide with 3 other specimens, FRANCE, Source du Lez (Hérault), N 43 0 37 /, E 0 3 0 53/, 36 m a.s.l., 6.xii.1970, on Teucrium polium , leg. F. Leclant. Paratypes: 27 more apt. on 7 slides, same data as holotype; 1 apt. on 1 slide and 5 ovip. on 4 slides, FRANCE, St Baudille, Larzac (Hérault), N 43 0 44 /, E 0 3 0 29/, 829 m a.s.l., 3.xii.1970, on Teucrium polium , leg. F. Leclant (FL 5080, GR 18285); 16 apt. and 1 immature on 4 slides, FRANCE, St Saturnin-les-Apt (Vaucluse), N 43 0 56 /, E 0 5 0 23/, 394 m a.s.l., 7.xii.1972, on Teucrium polium , leg. F. Leclant (FL 5881, GR 18286).

Type specimens are deposited in the Muséum national d’Histoire naturelle, Paris, France (holotype and 32 paratypes on 11 slides), the UMR Centre de Biologie pour la Gestion des Populations, Montpellier, France (8 paratypes on 2 slides) and the Natural History Museum, London, UK (10 paratypes on 3 slides).

Other material examined (not in type series). FRANCE: 5 apt., Éguilles (Bouches du Rhône), N 43 0 35 /, E 0 5 0 18/, 228 m a.s.l., 13.vi.1967, on Teucrium polium , leg. F. Leclant (FL 2038); 1 apt., Éguilles (Bouches du Rhône), N 43 0 35 /, E 0 5 0 18/, 228 m a.s.l., 27.xi.1969, on Teucrium polium , leg. F. Leclant (FL 3854 and GR 18287); 10 apt., Lambesc (Bouches du Rhône), N 43 0 39 /, E 0 5 0 15/, 206 m a.s.l., 24.ii. 1970, on Teucrium polium , leg. F. Leclant and G. Remaudière (FL 4086 and GR 9203); 6 apt., Luberon, (Vaucluse), N 43 0 47 /, E 0 5 0 13/, 700 m a.s.l., 22.iii.1970, on Teucrium polium , leg. G. Remaudière (GR 9275); 10 apt., Luberon (Vaucluse), N 43 0 47 /, E 0 5 0 13/, 700 m a.s.l., 22.iii.1971, on Teucrium polium , leg. F. Leclant and G. Remaudière (FL 4171 and GR 9383); 1 apt., Lourmarin (Vaucluse), N 43 0 45 /, E 0 5 0 21/, 214 m a.s.l., 22.iii.1970, on Teucrium polium , leg. G. Remaudière (GR 9283); 6 apt., Carnoules (Var), N 43 0 18 /, E 0 6 0 11/, 228 m a.s.l., 25.iii.1970, on Teucrium polium , leg. G. Remaudière (GR 9350); 10 apt., La Taillade (Hérault), N 43 0 37 /, E 0 3 0 37/, 221 m a.s.l., 12.iv.1970, on Teucrium polium , leg. F. Leclant and G. Remaudière (FL 4308 and GR 9442); 10 apt., Barcelonne (Drôme), N 44 0 51 /, E 0 5 0 0 2/, 299 m a.s.l., 21.v.1970, on Teucrium polium , leg. G. Remaudière (GR 9470); 8 apt. and 13 al., Rottane ( Corsica), N 42 0 10 /, E 0 9 0 25/, 55 m a.s.l., 4.vi.1970, on Teucrium polium , leg. F. Leclant (FL 4633a and GR 18284); 10 apt., S Castellane (Alpes-de-Haute-Provence), N 43 0 50 /, E 0 6 0 30/, 733 m a.s.l., 1.xi.1989, on Teucrium polium , leg. G. Remaudière (GR 16015); 2 al., Barcelonne (Drôme), N 44 0 51 /, E 0 5 0 0 2/, 299 m a.s.l., 21.v.1970; on Teucrium polium , leg. G. Remaudière (GR 9470); ITALY: 13 apt., Ventimiglia, N 43 0 47 /, E 0 7 0 36/, 9 m a.s.l., 30.iv.1969, on Teucrium polium , leg. D. Hille Ris Lambers (HRL 631 and BM 1984-340), UKRAINE: 17 apt. on 4 slides, Crimea, Alushta, N 44 0 41 /, E 34 0 24 /, 30 m a.s.l., 05.vii, 1960, on Teucrium polium , leg. J. Holman (3707); LEBANON: 7 apt. on 1 slide, Beirut, N 33 0 53 /, E 35 0 30 /, 0 m a.s.l., 3.iv.1966, on Teucrium polium , leg. Moericke (155 and BM 1984-340); IRAN: 6 apt., Rte Chemchak, 20 km E of Teheran, N 36 0 0 0/, E 51 0 29 /, 1800 m a.s.l., 20.v.1955, on Teucrium polium , leg. G. Remaudière (GR i301); 2 apt., Rte Chemchak, 20 km E of Teheran, N 36 0 0 0/, E 51 0 29 /, 1800 m a.s.l., 10.vi.1955, on Teucrium polium , leg. G. Remaudière (GR i490); 2 al. on 1 slide, Rte Tchalus (Chalus), N 36 0 39 /, E 51 0 24 /, 58 m a.s.l., 12.v.1959, on Teucrium sp., leg. G. Remaudière (GR i1612).

Etymology. The specific name is that suggested by D. Hille Ris lambers, in a letter to F. Leclant dated November 12th, 1969, and is derived from the specific name of its host plant.

Description. Apterous viviparous females (143 specimens /17 samples).

Colour in living specimens: variable from yellow (specimens from Iran, note on slide label by G. Remaudière) to dark green ( Leclant 1978), with dark SIPH and cauda. Pigmentation on slide: ANT I–II brown, ANT III pale (or ANT III and IV both pale when ANT 6-segmented), penultimate antennal segment brown either over whole length or brown on distal part and pale basally, last antennal segment (including PT) brown; head, URS, coxae, trochanters, femora, tarsi (fig. 2 f), SIPH, subgenital and anal plates and cauda brown, tibiae pale with dark apices, dorsum of abdomen in front of SIPH pale, ABD TERG VII–VIII with or without brown cross-bands, presiphuncular and postsiphuncular sclerites absent.

Morphological characters: Body oval (fig. 2 a, b). ANT 5 or 6-segmented (figs. 2 c, d). ANT 0.35–0.70× BL. PT 1.20–2.43× ANT V b or ANT VI b. ANT III without sec. rhin.. Frons convex or with shallow w-shaped outline in dorsal view. Rostrum reaches to or just passes the hind coxae. URS pointed with concave sides and 2 (-3) accessory setae (fig. 2 e), its length 1.28–2.35× ANT V b or ANT VI b and 1.49–2.25× HT II. THR—0.23–1.34. FTC 3:3:2. HT II W 0.21 –0.41× HT II L. Dorsal reticulation distinct. MTu on prothorax 0.010–0.049 mm in diameter. ABD TERG I and VII with small or medium-sized mammariform MTu, respectively 0.006–0.031 mm and 0.007–0.036 mm in diameter. Marginal seta on ABD TERG I is 0.19–2.83× diameter of the MTu on the same tergite. ABD TERG II–VI without MTu. SIPH cylindrical, imbricated, with a small to moderately developed flange (fig. 2 a, b), its length 1.98–5.34× its width in the middle and 0.93–1.98× cauda. ABD TERG VIII with 2 setae. Subgenital plate oval, with spinulose imbrications and 2 setae on the anterior part of the disc and 4–11 setae on the posterior margin (fig. 2 g). Cauda finger-like, without or with a slight constriction in the middle and bearing long and curved 3–7 setae and a lot of strong black spinules (fig. 2 h), its length 0.85–1.95× its basal width. Other morphometric data are given in Table 1 .

Measurements: apt. (n=143) al. (n=17) ovip. (n=5) Longest seta on ABD TERG III/ANT III BD 0.27–1.35 (0.39) 0.33–1.67 0.40–0.58 SIPH/its basal width 1.36–4.39 (2.41) 2.15–3.87 1.82–2.43 SIPH/its distal width 2.40–6.79 (4.11) 3.45–5.39 3.08–3.73 SIPH/BL 0.07–0.19 (0.11) 0.10–0.14 0.09–0.11 SIPH/ANT V b or ANT VI b 1.06–2.84 (1.98) 1.36–1.85 1.54–1.89 SIPH/URS 0.67–1.76 (0.93) 0.85–1.31 0.83–0.94 SIPH/HT II 1.30–3.16 (1.98) 1.52–2.40 1.59–1.78 SIPH/CAUDA 0.93–1.98 (1.28) 1.18–1.82 1.28–1.37 Longest seta on ABD TERG VIII/ANT III BD 0.38–2.87 (0.50) 0.80–3.27 0.82–1.24

Alate viviparous females (17 specimens /2 samples)

Colour in life unknown. Pigmentation of slide-mounted specimens: ANT rather uniformly brown, except sometimes base of ANT III; head and thorax dark brown; third rostral segment and URS brown; coxae and trochanters brown; femora of fore and mid legs pale brown, of hind legs darker on distal two-thirds; tibiae pale brown except towards apices, tarsi brown; SIPH brown, cauda pale to mid-brown. Dorsal sclerites brown, spiracular plates and marginal sclerites dark brown.

Morphological characters: Body pear-shaped (fig. 2 i). ANT 5- or 6-segmented (5-segmented in all 13 individuals in sample 4633a from Corsica) (fig. 2 k). ANT 0.55–0.68× BL, with PT 1.64–3.16× ANT V b or ANT VI b. ANT III with (4-) 5–8 (-9) secondary rhinaria, varying in size but mostly rather small, distributed in an irregular row over most of its length. Longest setae on ANT III 0.29–0.82× basal diameter of segment, which is more constricted at base than in apterae. Frons convex or with shallow w-shaped outline in dorsal view. Rostrum reaches to the hind coxae. URS pointed with concave sides and 2 accessory setae, its length 1.23–2.00× ANT V b or ANT VI b and 1.51–2.25× HT II. MTu on prothorax medium-sized— 0.015–0.031 mm in diameter. Forewings with media twice-branched, and hind wings with 2 oblique veins. THR 0.39–0.90. FTC: 3:3:2. HT II W 0.21 –0.28× HT II L. ABD TERG I to IV with marginal sclerites. Dorsal reticulation distinct. ABD TERG I and VII with small or medium-sized mammariform MTu, 0.010–0.025 mm of diameter. Marginal seta on ABD TERG I 0.006–0.028 mm, 0.39–2.33× diameter of the MTu on the same tergite. ABD TERG II–VI without MTu. SIPH cylindrical, imbricated, with small flange, its length 2.89–4.44× its width in the middle and 1.18–1.82× cauda. ABD TERG VIII with 2 setae. Postsiphuncular sclerites present, well-developed. Transverse spino-pleural sclerites present on ABD TERG VI, VII and VIII; that on VI variably developed and sometimes fused laterally with postsiphuncular sclerites to form a continuous broad band across the tergite. Subgenital plate with 2 setae on anterior part of disc and 6–9 on posterior margin. Cauda finger-shaped to elongate conical, with 4–6 setae. For other morphometric data see Table 1 .

Oviparous females (5 specimens /1 sample)

Colour in life unknown. Pigmentation of slide-mounted specimens: ANT I, II, IV and V mid-brown, ANT III paler; front part of head, third rostral segment and URS brown; coxae and trochanters brown; fore and mid legs mainly pale brown except tibial apices and tarsi; hind femora, tibiae and tarsi brown; SIPH and cauda dark brown (SIPH sometimes paler basally); anal and subgenital plates dark brown; spiracular plates and marginal tubercles dark brown.

Morphological characters: Body broadly oval. ANT 5 -segmented (fig. 2 j). ANT 0.43–0.51× BL. PT 1.39– 1.65× ANT V b. ANT III without sec. rhin. Frons with shallow w-shaped outline in dorsal view. Rostrum reaches beyond the hind coxae. URS pointed with concave sides and 2 accessory setae, its length 1.68–2.02× ANT V b and 1.80–1.93× HT II. MTu on prothorax large-sized— 0.031–0.058 mm of diameter. THR 0.33–0.67. HT II W 0.25 – 0.28× HT II L. Hind tibiae heavily pigmented, swollen over most of length, reaching a maximum width at midlength of about twice the minimum distal width, and bearing 60–73 rather evenly distributed scent plaques (“pseudosensoria”) (fig. 2 l). FTC: 3:3:2. Dorsal reticulation distinct. ABD TERG I and VII with medium-sized mammariform MTu, 0.018–0.028 mm of diameter. Marginal seta on ABD TERG I 0.009–0.012 mm, 0.43–0.61× diameter of the MTu on the same tergite. ABD TERG II–VI without MTu. SIPH cylindrical, imbricated, with normal flange, its length 2.64–3.37× its width in the middle and 1.28–1.37× cauda. ABD TERG VIII with 2 setae. Subgenital plate divided into paired rugose sclerites, with 7–8 setae on posterior edge and 4–5 setae on the central part of each sclerite. Anal plate with setae up to 0.050 mm long. Cauda bluntly conical with 6–7 setae, its length 0.81–1.01× its basal width. For other morphometric data see Table 1 .

Intraspecific variability. A. polii is known from 17 localities in France (including Corsica), Italy, Ukraine, Lebanon and Iran (fig. 1, black circles 1–17). It is unusual for so much material from diverse locations to be available when describing a new species, and this is reflected in the extent of the observed variation. In particular, apterae from Ukraine and Lebanon were longer haired than samples from fifteen localities in three other countries. For example, the longest cephalic frontal setae were 0.018–0.034 mm in specimens from Ukraine and Lebanon, and 0.009–0.023 mm in those from other localities, with only the sample from Iran (0.014–0.023 mm) spanning the two size ranges. No other consistent differences were found between these populations, and the variation in length of setae could be due to differences in macro- or microclimate, or to geographical variation.

Life cycle. This is a monoecious species reported to live in ant-attended colonies on the apical parts of shoots and sometimes on undersides of leaves of Teucrium polium L. (D. Hille Ris Lambers, unpublished field note). Five oviparae were collected in early December at St Baudille, indicating that there is a holocycle in southern France. Leclant (1978, p.79) noted that males are apterous, but unfortunately there are no males on slides in the collections in Paris or Montpellier. Leclant also wrote that he had found the fundatrix, and provided a photograph of a specimen with a relatively short PT and short SIPH and cauda labelled as such. However, when morphometric data for all the apterous specimens collected in early spring were compared with data for apterae collected at other times of year, none of the early spring specimens had the distinctive features that one would expect in a fundatrix morph.

An apterous vivipara was collected along with the oviparae by Leclant in early December at St Baudille, and collections in the same month at St Saturnin-les-Apt and Source du Lez were all of apterous viviparae, indicating that overwintering of parthenogenetic generations (anholocycly) occurs in southern France. The relatively short PT, SIPH and cauda of some specimens may be due to development at low temperature. It seems likely that parthenogenetic overwintering of this species occurs wherever conditions permit.

Diagnosis. Thirty-five species of the genus Aphis live on the family Lamiaceae worldwide ( Blackman & Eastop 2006). Many of these, including A. polii and the other two species that feed specifically on plants of the genus Teucrium ( A. teucrii and A. alienus ), belong to the frangulae / gossypii group. Members of this group are in general more readily distinguished by biological and molecular rather than morphological differences ( Stroyan 1984; Coeur d’acier et al. 2007; Kim & Lee 2008; Kim et al. 2010; Cocuzza & Cavalieri 2014; Lagos-Kutz et al. 2014). Comparing A. polii with the other Lamiaceae-feeding Aphis species, only A. clinepetae Pashtshenko, 1993 , A. passeriniana (del Guercio, 1900) and A. raji ( Kumar & Burkhardt, 1970) have such a high ratio of URS/HT II; this ratio is 1.49–2.25 in A. polii , 1.50–1.90 in A. clinepetae , 1.54–2.02 in A. passeriniana , and 2.00– 2.20 in A. raji ( Stroyan 1984; Pashtshenko 1993; Blackman & Eastop 2015). Apterous viviparae of A. polii have shorter cephalic frontal setae than those of A. clinepetae ; 0.007–0.032 mm, as opposed to 0.031–0.056 mm in A. clinepetae ( Pashtshenko 1993) . In apterous viviparae of A. passeriniana the rostrum only reaches to about midway between the mid and hind coxae, whereas the rostrum reaches to or just passes the hind coxae in apterae of A. polii , and the alatae of A. passeriniana have conspicuously large and dark postsiphuncular sclerites, much darker than the marginal and posterior abdominal sclerites ( Stroyan 1984), whereas the dorsal abdominal sclerites of alate A. polii are all similarly pigmented. In apterous viviparae of A. raji the URS is distinctly longer (0.190–0.230 mm) than in A. polii (0.091–0.151 mm). A. raji also has setae on ANT III that are about twice as long as the basal diameter of the segment, whereas in A. polii these setae are shorter than the basal diameter ( Kumar & Burkhardt 1970; Blackman & Eastop 2015). These species all have specific associations with different host plants. According to Pashtshenko (1993), A. clinepetae lives on Clinopodium chinense and Nepeta manchuriensis , while according to Stroyan (1984) and García Prieto & Nieto Nafría (2005) A. passeriniana inhabits Salvia spp., and according to Blackman & Eastop (2015) A. raji is recorded from Callicarpa , Colebrookea and Salvia . The morphological feature that they share—the long URS—is probably a convergent adaptation for feeding on the tomentose stems and undersides of leaves of these plants.

The new species seems to be specific to Teucrium polium . Of the other two Aphis species associated specifically with Teucrium , only A. teucrii has been recorded from T. polium ( Holman 2009) . A. alienus is mostly found on stem bases and rhizomes of T. scorodonia ( Stroyan 1984) . A. teucrii seems less specific in its choice of Teucrium spp. and is normally recorded from the aerial parts, apart from one sample in the BMNH collection from roots of T. chamaedrys in Czech Republic. Apterous viviparae of A. alienus and A. teucrii are separated from those of A. polii in a bivariate plot of the length of URS vs. the length of HT II (fig. 3). The same discriminant can be used for alatae. A. alienus and A. teucrii are less easy to separate, and have been suspected to be synonymous ( Blackman & Eastop 2006), but there are consistent differences in the shape of the SIPH of apterous viviparae and in the incidence of marginal tubercles (see key to aphids on Teucrium in Blackman & Eastop 2015).

Five other species of the genus Aphis — A. fabae , A. frangulae , A. gossypii , A. origani and A. salviae have been recorded from Teucrium spp. ( García Prieto & Nieto Nafría 2005; Holman 2009; Blackman & Eastop 2015). The first three of these are polyphagous. A. origani feeds specifically on Origanum and A. salviae on Salvia , so the single records of these species on Teucrium may be based on misidentifications, or sampling of vagrants on unsuitable host plants. The new species can again be distinguished from these four species by its URS/HT II ratio; 1.49–2.25 in A. polii , while it is 0.85–1.50 in the other species.

Figures

FIGURE 2. Aphis polii sp. n. a – h, apterous vivipara; a and b, specimens from Ventimiglia (a) and Éguilles (b) showing variation in shape of siphunculi and cauda; c, 5 - segmented antenna; d, 6 - segmented antenna; e, ultimate rostral segment; f, second segment of hind tarsus; g, subgenital plate; h, cauda; i, alate vivipara; j, ovipara; k, antenna of alate vivipara; l, hind tibia of ovipara.

Tables

TABLE 1. Measurements and ratios of Aphis polii sp. n. (measurements are lengths unless otherwise indicated, and are given in mm). Note: measurements and ratios for holotype are in parentheses.

BL	0.714–1.377 (1.000)	0.925–1.548	1.060–1.183
BW	0.424–0.976 (0.675)	0.429–0.706	0.655–0.730
ANT	0.349–0.730(0.460)	0.567–1.032	0.500–0.540
ANT I	0.020–0.064 (0.038)	0.036–0.052	0.040–0.048
ANT II	0.032–0.052 (0.040)	0.036–0.052	0.041–0.046
ANT III BD	0.013–0.025 (0.018)	0.009–0.017	0.019–0.022
ANT III (if 6 segmented)	0.056–0.169	0.171–0.270	–
ANT III (if 5 segmented)	0.103–0.222 (0.127)	0.194–0.233	0.143–0.175
Longest seta on ANT III	0.005–0.017 (0.006)	0.005–0.009	0.006–0.009
ANT IV	0.037–0.112 (0.072)	0.079–0.163	0.079–0.095
ANT V (if 6 segmented)	0.057–0.123	0.119–0.136	–
ANT V b or ANT VI b	0.048–0.095 (0.056)	0.063–0.095	0.063–0.077
PT	0.079–0.182 (0.109)	0.142–0.278	0.099–0.111
Longest cephalic frontal seta	0.007–0.032 (0.009)	0.006–0.019	0.010–0.013
Rostrum	0.306–0.524 (0.373)	0.308–0.433	0.406–0.423
URS	0.091–0.151 (0.119)	0.094–0.135	0.126–0.134
Posterior seta on the hind trochanter	0.009–0.037 (0.013)	0.012–0.028	0.015–0.031
HFEM	0.154–0.294 (0.193)	0.198–0.381	0.222–0.226
HTIB	0.270–0.492 (0.334)	0.375–0.677	0.357–0.389
HT II length	0.050–0.082 (0.056)	0.056–0.075	0.067–0.070
HT II width	0.014–0.025 (0.018)	0.013–0.018	0.017–0.019
Marginal seta on ABD TERG I	0.005–0.025 (0.009)	0.006–0.028	0.009–0.012
Longest seta on ABD TERG III	0.005–0.023 (0.007)	0.005–0.025	0.008–0.011
SIPH	0.083–0.231 (0.111)	0.096–0.163	0.111–0.119
Distal width of SIPH	0.022–0.055 (0.027)	0.025–0.036	0.030–0.037
Middle width of SIPH	0.022–0.063 (0.031)	0.025–0.037	0.035–0.042
Basal width of SIPH	0.034–0.089 (0.046)	0.031–0.049	0.049–0.061
Longest seta on ABD TERG VIII	0.006–0.046 (0.009)	0.012–0.049	0.016–0.026
Cauda	0.071–0.135 (0.087)	0.073–0.127	0.086–0.087
Basal width of cauda	0.062–0.117 (0.080)	0.062–0.097	0.086–0.108
Ratios:
PT/ANT III	0.56–1.73 (0.86)	0.63–1.08	0.63–0.75
PT/ANT Vb or ANT VIb	1.28–2.35 (1.95)	1.64–3.16	1.39–1.65
ANT V b or ANT VI b/ANT III	0.31–1.13 (0.44)	0.31–0.52	0.44–0.49
ANT IV/ANT III	0.36–0.90 (0.57)	0.37–0.69	0.54–0.62
ANT III/BL	0.06–0.22 (0.13)	0.16–0.23	0.12–0.17
ANT III/SIPH	0.53–2.05 (1.14)	1.13–2.17	1.20–1.47
Longest seta on ANT III/ANT III BD	0.24–0.94 (0.33)	0.29–0.82	0.30–0.43
Cephalic frontal seta/ANT III BD	0.35–2.13 (0.50)	0.58–1.73	0.53–0.62
Rostrum/ANT III	1.54–6.52 (2.94)	1.45–2.25	2.36–2.96
Rostrum/BL	0.30–0.47 (0.37)	0.31–0.37	0.35–0.39
URS/ANT Vb or ANT VIb	1.28–2.35 (2.13)	1.23–2.00	1.68–2.02
URS/HT II	1.49–2.25 (2.13)	1.51–2.25	1.80–1.93
URS/SIPH	0.57–1.48 (1.07)	0.76–1.18	1.07–1.21
HFEM/BL	0.15–0.26 (0.19)	0.20–0.27	0.19–0.21
HTIB/BL	0.28–0.46 (0.33)	0.38–0.51	0.32–0.36
HT II/ANT V b or ANT VI b	0.72–1.19 (1.00)	0.68–1.00	0.87–1.11