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Morphological discrimination of six species of
the genus Anuraphis (Hemiptera: Aphididae),
including description of a new species
862
Sebastiano Barbagallo,
1
Giuseppe E Cocuzza
Dipartimento di Scienze e Tecnologie Fitosanitarie, University of Catania,
Via S. Sofia 100, Catania, 95123 Italy
The Canadian Entomologist 135: 839 – 862 (2003)
Abstract—The most relevant morphometric characteristics of six species of the
genus Anuraphis Del Guercio were compared. Anuraphis shaposhnikovi sp. nov. is
described and its morphological differences from the closely related species
Anuraphis subterranea are presented. The new species was collected in Sicily and
in the central area of the Italian peninsula on Magydaris pastinacea (Lam.) Paol.
(Apiaceae) and Opopanax chironium (L.) Koch (Apiaceae), which are its secondary
host plants. A key to the viviparous morphs (apterae and alatae) of the seven west
-
ern Palaearctic species living on secondary hosts is provided. Discriminant func
-
tions have been derived to separate both apterae and alatae of A. shaposhnikovi and
A. subterranea.
Barbagallo S, Cocuzza GE. 2003. Discrimination morphologique de six espèces d’Anuraphis
(Hemiptera : Aphididae) et description d’une nouvelle espèce. The Canadian Entomolo-
gist 135 : 839–862.
Résumé—On trouvera ici une comparaison des caractères morphométriques les
plus pertinents de six espèces d’Anuraphis Del Guercio, ainsi que la description
d’Anuraphis shaposhnikovi sp. nov. et des caractéristiques morphologiques qui le
distinguent d’Anuraphis subterranea, une espèce très apparentée. La nouvelle es-
pèce a été récolté en Sicile et dans la région centrale de la péninsule italienne sur
Magydaris pastinacea (Lam.) Paol. (Apiaceae) et sur Opopanax chironium (L.)
Koch (Apiaceae) qui sont ses hôtes secondaires. Un clé permet l’identification des
formes vivipares (aptères et ailées) des sept espèces de la région paléarctique occi
-
dentale qui vivent sur des hôtes secondaires. Des fonctions discrimantes facilitent la
séparation des formes aptères et ailées d’A. shaposhnikovi et d’A. subterranea.
[Traduit par la Rédaction]
Introduction
The aphid genus Anuraphis Del Guercio, 1907 (Hemiptera: Aphididae) is consid
-
ered to be of Palaearctic origin and is among the rather primitive genera of Aphidinae,
Macrosiphini. It includes 10 putative species (Remaudière and Remaudière 1997), of
which only 6 (namely, A. cachryos Barbagallo et Stroyan, 1982; A. catonii Hille Ris
Lambers, 1935; A. farfarae (Koch, 1854); A. ferulae Shaposhnikov, 1995; A. pyrilaseri
Shaposhnikov, 1950; and A. subterranea (Walker, 1852)) have a well-defined taxonomic
status. There are four other nominal species (i.e., A. capparidis Nevsky, 1929;
A. corthusae Nevsky, 1929; A. floris Mozen, 1934; and A. katsurae Shinji, 1952) listed
in the catalogues of either Eastop and Hille Ris Lambers (1976) or Remaudière and
Remaudière (1997), and whose taxonomic status is not yet well known; if they prove to
839
1
Corresponding author (e-mail: sebarbag@unict.it).
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be valid taxa, they may not be representatives of the genus Anuraphis as presently
stated.
The six currently recognized species probably are native to the western
Palaearctic region and perhaps to the Ponto-Mediterranean area where, in particular, the
primary host plant (Pyrus communis L. (Rosaceae)) appears to have originated.
Apiaceae serve as secondary hosts for five of the six species, except for A. farfarae,
which feeds on Asteraceae. Large, myrmecophilous colonies develop on the root collar
parts and the inner sheath area of the basal leaves of the secondary host plant. In suit
-
able ecological conditions, these secondary hosts may be used by some species for con
-
tinuous parthenogenetic reproduction throughtout the year.
Well-known taxa are the widely distributed species A. farfarae, A. subterranea,
and A. catonii; apparently A. pyrilaseri has a less-extensive distribution in the western
Palaearctic, ranging from the Ponto-Mediterranean to southern European areas. All of
these are proven to be holocyclic (Shaposhnikov 1951; Kolesova 1972), at least in some
biotopes and within certain populations. The presence of two life-cycle categories in
A. farfarae, one holocyclic and the other anholocyclic, has prompted Shaposhnikov
(1974) and Shaposhnikov and Sharov (1977) to consider them as two taxa at a
subspecific level (A. farfarae Koch s.s., nominal subspecies, and A. farfarae subsp.
dianae Shaposhnikov). Anuraphis pyrilaseri, a rather variable species in colour, has
both holocyclic and anholocyclic populations; this species usually has continuous par-
thenogenetic reproduction on secondary hosts in the southern Italian regions. Anuraphis
cachryos was originally described from Sicily; it is common in other southern Italian
regions and probably more widespread around the Mediterranean lands and the Middle
East where it was recorded in Iran (Hodjat 1993). Anuraphis cachryos is only known as
an anholocyclic species on secondary hosts. The life cycle of the more recently de-
scribed A. ferulae is not yet known; this aphid is recorded from Ferula spp. (Apiaceae)
in Tadjikistan (Shaposhnikov 1995).
From a morphological point of view, the genus Anuraphis appears to be well de-
fined and characterized, containing a rather homogeneous group of species. The relative
similarity between species, on the other hand, means that the ranges of many
morphometric characters overlap and make it difficult to discriminate species. The ge-
nus Nearctaphis Shaposhnikov, 1950, which includes fourteen species, is the most mor
-
phologically similar genus (Hille Ris Lambers 1970) and is considered the vicariant
Nearctic taxon of the genus Anuraphis.
The true apterous morph of Anuraphis spp. has secondary rhinaria typically
grouped only on the basal part of antennal joint III. A high percentage of “apteriform”
specimens in a colony have typical alatoid features, showing an increase in number and
distribution of secondary rhinaria on antennal joint III (occasionally extended also to
antennal joint IV) and in dorsal body sclerotization, particularly of the abdominal seg
-
ments. These features can occur even in specimens that are without well-differentiated
ocelli and (or) evidence of wing-pad development.
A few years ago, small samples of an Anuraphis sp. with similar morphological
features to A. subterranea, but paler in colour, were collected in Sicily on the basal
parts of Magydaris pastinacea (Lam.) Paol. (Apiaceae) by S Barbagallo. Thereafter, a
few other samples of the same aphid were collected on Opopanax chironium (L.) Koch,
another tall member of the Apiaceae family having a wider Mediterranean distribution.
These additional samples provided an opportunity to find out if the collected aphids be
-
longed to a separate taxon despite their close morphological resemblance to
A. subterranea and, also, to make a broader study of the relationships among other
available populations of Anuraphis species. Several samples of six postulated species of
the genus Anuraphis, most of them collected in Italy, were measured biometrically and
analyzed, and their morphological differences evaluated. Results obtained are illustrated
840 THE CANADIAN ENTOMOLOGIST November/December 2003
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here, whereas short preliminary information on the topic was recently given by
S Barbagallo, I Patti, GE Cocuzza, and MC Mosco (unpublished data) . The Anuraphis
spp. samples collected on Magydaris sp. and Opopanax sp. appear to belong to an
undescribed species, which can be separated both morphologically and by allozyme
analysis (unpublished) from other western Palaearctic species. A new taxon
(A. shaposhnikovi sp. nov.) is described and may be considered a reproductively iso
-
lated offshoot of the strictly linked A. subterranea, showing little genetic distance from
A. cachryos as well. Moreover, a morphometric analysis was computed with the aim to
better clarify the taxonomic status of these aphids and to facilitate the separation of the
new species from the closely related A. subterranea.
Taxonomic treatment
Anuraphis shaposhnikovi sp. nov.
(Figs. 1–4, Table 1)
Type material
Holotype: apterous viviparous female on Magydaris pastinacea, Buseto Palizzolo
(Trapani, Italy), Scorace woodland, 3.vi.1997 (SB legit). Paratypes: 129 apterous and
56 alatae viviparous females collected on the following host plants, localities in Italy,
and dates — (a) M. pastinacea, Buseto Palizzolo, as above, 9.vi.1999 and 5.vi.2001;
same host plant, Godrano (Palermo), Ficuzza woodland, 4.vi.1999, and Marineo
(Palermo), 2.vi.2000, 17.vii.2000 (ex culture), and 5.vi.2001; (b) Opopanax chironium,
Opi (L’Aquila), 26.vi.1996; Francavilla di Sicilia (Messina), 22.v.2001 and 8.vi.2001.
The holotype is in the collection of the Dipartimento di Scienze e Tecnologie
Fitosanitarie (Entomology section), University of Catania–I, Catania, Italy. Paratypes
are in the same aphidological collection (S Barbagallo) and in the collections of The
Natural History Museum of London, United Kingdom (P Brown); Museum National
d’Histoire Naturelle of Paris, France (G Remaudière); Institute of Entomology – Acad-
emy of Sciences, Ceské Budejovice, Czech Republic (J Holman); Canadian Forest Ser
-
vice, Laurentian Forestry Centre, Sainte-Foy, Quebec, Canada (FW Quednau); Florida
Department of Agriculture and Consumers Services, Gainesville, Florida, United States
of America (S Halbert); Entomology Research Museum, University of California, Riv
-
erside, California, United States of America (D Yanega).
Etymology
The new species is named in honour of the late Georgiy Christoforowich
Shaposhnikov, a research leader in “Anuraphidine” aphids, particularly of the genera
Anuraphis and Dysaphis.
Diagnosis
The general facies of the new taxon, A. shaposhnikovi, is the same as the rather
small and pale specimens of A. subterranea. Compared with the latter, apterae of
A. shaposhnikovi have, on average, less extensive body pigmentation, and both apterae
and alatae show a small reduction in either the number or size of abdominal tubercles.
Nevertheless, the most distinctive feature between the two species appears to be the differ
-
ent length of their ultimate rostral joint; it ranges 0.180–0.240 mm in apterae and
0.174–0.220 mm in alatae of A. shaposhnikovi, whereas it ranges 0.240–0.340 mm in apterae
and 0.214–0.320 mm in alatae of A. subterranea (measured from a random selection of
Volume 135 THE CANADIAN ENTOMOLOGIST 841
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80 apterae and 20 alatae of the latter species from 19 samples from different countries,
collected on Heracleum spp. and Pastinaca spp.). Therefore, in most cases, the ultimate
rostral joint length / length of joint II of hind tarsus (urj/IIht) ratio appears to be
discriminant between the two species. It (i) ranges 1.25–1.60 in apterae of
A. shaposhnikovi (where 15% of the 130 available specimens exceeded 1.50) compared
with 1.50–2.10 in 80 measured apterae of A. subterranea (where only 6% of specimens
have a ratio less than 1.60) and (ii) ranges 1.24–1.54 in alatae of A. shaposhnikovi, with
just less than 4% having a ratio greater than 1.50 (2 alatae out of 56 specimens) com
-
pared with 1.36–1.85 (but only rarely less than 1.50, particularly in summer specimens)
in the corresponding morph of A. subterranea.
842 THE CANADIAN ENTOMOLOGIST November/December 2003
FIGURE 1. Anuraphis shaposhnikovi, apterous viviparous female: antenna (a), distal part of rostrum (b),
distal part of tibia and tarsus of hind leg (c), and siphunculus (d). Scale bar = 0.1 mm.
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The number and distribution of abdominal spinal tubercles also helped to separate
the two Anuraphis species. Apterae and alatae of A. subterranea usually have a com
-
plete set of abdominal spinal tubercles ranging from abdominal segments I to VII, even
in specimens collected in most arid biotopes (Sicily). In A. shaposhnikovi, only 13
apterae specimens (7%) out of 130 apterae and 56 alatae had a complete set of 10 spinal
tubercles on abdominal segments I–V; the other specimens seen lack the presence of
spinal tubercles on some of the ante-siphuncular abdominal segments where they range
1–9 in number (average 4.10 in apterae and 4.40 in alatae). In the same species
post-siphuncular segments usually bear spinal tubercles, at least as single elements on
one side, on abdominal segments VI and VII and occasionally even on abdominal seg
-
ment VIII; their total number on these segments range (0)1–5(6), with an average of
2.70 both for apterae and alatae altogether.
Description
Apterous viviparous female (from secondary hosts)
Body oval in shape and 2.07–3.04 mm long. Frontal profile straight or slightly
sinuose. Dorsal sclerifications well developed on head, prothorax, marginal areas of
meso–metathorax and, as more or less compact bands, from abdominal segments (V) VI
to VII; small sclerifications can be present, in addition, on abdominal tergites anterior
Volume 135 THE CANADIAN ENTOMOLOGIST 843
FIGURE 2. Dorsal view of abdomen (half side) showing the usual distribution of spinal tubercles in
Anuraphis shaposhnikovi (a) and Anuraphis subterranea (b) apterous viviparous female (hairs omitted).
Scale bars = 0.5 mm.
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the siphunculi. Spinal and marginal tubercles surrounded by narrow sclerified bands.
Submarginal muscle sclerites and stigmal plates also well pigmented. Dorsal cuticle re
-
ticulation rather evanescent (unless on sclerified areas of thorax), particularly on the ab
-
dominal central area where it is sometimes hardly visible. Flat tubercles are present
with the following distribution: two pairs on head (one frontal and one occipital pair),
12–34
µ
m wide (there are rarely only three cephalic tubercles); a marginal series from
the prothorax to abdominal segment VII, the latter pair being usually the largest one,
ranging 30–48
µ
m in diameter; paired spinal tubercles set on thoracic terga (where they
are occasionally doubled on one or more segments) and on abdominal segments; here,
there are 4–16 (most frequently 6–9) tubercles altogether, 12–35
µ
m wide, usually pres
-
ent on both abdominal segments VI and VII (occasionally one or two also on abdominal
segment VIII) but not constantly present on abdominal segments anterior the siphunculi
where they range 2–10 in number. Body hairs all short and rather acute apically,
8–12
µ
m long on abdominal segment III up to 14–20 (28)
µ
m long on abdominal seg
-
ment VIII where they are 6–8 in number. Head. Antennae entirely pigmented (except,
sometimes, the paler base of joint III) and rather short, with the flagellum (joints
III–VI) 0.35–0.54 times the body length. Antennal hairs short and inconspicuous,
844 THE CANADIAN ENTOMOLOGIST November/December 2003
FIGURE 3. Anuraphis shaposhnikovi, general aspect of apterous viviparous female (specimen mounted
on a slide). Scale bar = 0.5 mm.
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5–9
µ
m on joint III or 0.17–0.32 times its basal articular diameter. Antennal joint III
bearing 14–64 small, roundish secondary rhinaria, setting ventrally along its entire
length (all specimens seen have alatoid antennal features); joint IV with (0)1–19
rhinaria, mostly placed on its 2/3 apical part; joint VI with processus terminalis
3.10–4.32 times its basal part. Rostrum reaching behind the metacoxae; its apical joint
180–240
µ
m long, with 5–6 short secondary hairs. Urj/IIht ratio ranging 1.25–1.60, and
therefore comparatively shorter, on average, than in other Anuraphis species. Thorax.
Legs rather short, with femora pale at their basal part; tibiae 0.32–0.42 times the body
length and moderately pigmented except at their darker basal and apical parts. Crural
hairs all quite short, 5–10
µ
m long on hind femour (dorsal ones) and from 6–15
µ
m
long (outer side ones) up to 15–25
µ
m long (inner distal ones) on hind tibiae. First tar
-
sal chaetotaxy 3:3:3. Abdomen. Siphunculi truncate-conical and well flanged, as is
usual in the genus, 0.06–0.09 times the body length and 2.00–3.10 times their diameter
at middle length, and shorter (0.70–0.90) than the frontal width (interantennal distance).
They are adorned with 20–34 (mostly 24–30) rows of spinules, which are not strongly
produced (compared with other congeneric species) and sometimes nearly coalescing
into scales on the basal part of siphunculus. Cauda rounded, 0.50–0.70 times the
siphuncular length and bearing 12–18 curved, long hairs. Genital plate well sclerified,
with 4–10 hairs on the anterior half and 22–32 hairs along its latero-posterior margin,
all short (6–18
µ
m long). Colour. Body yellowish ocherous to brownish (mostly in
specimens kept in laboratory at low temperature (8–10 °C) for 2–3 weeks); nymphs
pale-yellowish to salmon-pinkish.
Volume 135 THE CANADIAN ENTOMOLOGIST 845
FIGURE 4. Anuraphis shaposhnikovi, general aspect of alate viviparous female (specimen mounted on a
slide). Scale bar = 0.5 mm.
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846 THE CANADIAN ENTOMOLOGIST November/December 2003
Length (mm) of antennal joint
No. of secondary
rhinaria
Specimen
No.
Body
length
(mm)
Antennal
flagellum
length (mm) III IV V VI
Urj*
length
(mm)
IIht*
length
(mm)
Siphuncular
length
(mm)
Caudal
length
(mm) III IV
1 2.77 1.18 0.284 0.180 0.120 0.104 + 0.332 0.206 0.160 0.180 0.120 24/30 2/8
2 2.72 1.31 0.340 0.200 0.116 0.108 + 0.368 0.212 0.160 0.180 0.120 43/41 5/9
3 2.43 1.27 0.320 0.180 0.120 0.100 + 0.368 0.200 0.148 0.160 0.100 36/34 5/8
4 2.48 1.35 0.360 0.200 0.136 0.104 + 0.370 0.200 0.160 0.168 0.118 51/53 12/17
5 2.43 1.33 0.320 0.248 0.120 0.102 + 0.360 0.204 0.140 0.180 0.100 31/42 5/5
6 2.17 1.27 0.346 0.184 0.120 0.108 + 0.350 0.200 0.158 0.176 0.108 38/40 5/8
7 2.48 1.18 0.304 0.170 0.116 0.900 + 0.340 0.200 0.152 0.198 0.106 47/42 10/10
8 2.81 1.53 0.488 0.200 0.124 0.112 + 0.428 0.232 0.166 0.200 0.124 59/62 6/12
9 3.04 1.53 0.460 0.220 0.136 0.118 + 0.412 0.240 0.160 0.200 0.120 39/44 4/8
10 2.58 1.49 0.440 0.224 0.128 0.120 + 0.412 0.224 0.152 0.204 0.100 47/43 9/7
11 2.07 1.30 0.392 0.184 0.120 0.104 + 0.360 0.192 0.144 0.160 0.090 68/66 22/19
12 2.07 1.41 0.400 0.220 0.120 0.106 + 0.400 0.200 0.140 0.176 0.100 70/72 22/21
13 1.91 1.38 0.360 0.236 0.124 0.110 + 0.400 0.196 0.140 0.180 0.092 68/71 25/23
14 2.21 1.52 0.480 0.208 0.148 0.116 + 0.406 0.200 0.146 0.184 0.100 73/78 23/24
15 2.13 1.49 0.460 0.200 0.120 0.100 + 0.448 0.200 0.154 0.180 0.090 64/62 12/15
NOTE: Specimen No. 1 is the holotype and specimen Nos. 2–15 are paratypes. Specimen Nos. 1–3 and 11 collected from Magydaris pastinacea, Buseto Palizzolo (Trapani), Sicily, 3.vi.1997; speci-
men No. 4 collected from the same host plant, Godrano (Palermo), Sicily, 4.vi.1999; specimen Nos. 5–7 and 12–14 collected from the same host plant, Marineo (Palermo), Sicily, 2.vi.2000; speci-
men Nos. 8 and 15 collected from Opopanax chironium, Opi (L’Aquila), Italy, 26.vi.1996; and specimen Nos. 9–10 collected from the same host plant, Francavilla di Sicilia (Messina), Sicily,
8.vi.2001.
* Urj is the ultimate rostral joint length and IIht is the length of joint II of the hind tarsus.
TABLE 1. Morphometric data of apterous (1–10) and alatae (11–15) viviparous Anuraphis shaposhnikovi females.
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Alate viviparous female (from secondary hosts)
Body 1.80–2.60 mm long. Head with frontal profile clearly sinuose and usually
bearing four tubercles (occasionally one of the two pairs may be absent, as in apterae).
Abdominal marginal tubercles usually present from abdominal segments I to VII, rarely
lacking from one side on abdominal segment VI or VII; spinal tubercles occurring in a
variable number from 1 to 13 (in most cases 4–8), frequently absent on some of abdom-
inal segments II–V but are present as a rule on abdominal segments VI and VII (at least
on one side). Dorsal abdominal sclerifications dominated, as is usual in the genus, by a
large median patch on abdominal segments IV–VI, followed by single, transverse bands
on abdominal segments VII and VIII; anterior segments of abdomen with more or less
fragmented sclerifications, including submarginal muscle sclerites and marginal patches
on abdominal segments II–IV. Antennae completely pigmented and flagellum 0.46–0.72
times the body length. Processus terminalis 3.20–4.80 times the basal part of joint VI;
joint III bearing 40–94 secondary rhinaria; joint IV with 4–26 rhinaria. Last rostral joint
1.24–1.54 times joint II of hind tarsus. Femora with their basal part pale; tibiae
0.48–0.56 times the body length, usually a little paler in the middle than at their basal
and distal parts. Siphunculi 0.07–0.09 times the body length and bearing 18–28 rows of
small spinules. Other morphological features more or less similar to the corresponding
ones of the apterous morph. Colour. Head and thorax blackish; abdomen yellowish with
olive–brown sclerifications.
Host plants
Eight of the 11 presently available samples of A. shaposhnikovi were collected on
Magydaris pastinacea (Lam.) Paol. in western Sicily, in some woodland areas of
Palermo and Trapani provinces. One sample was collected on Opopanax chironium (L.)
Koch at Opi (L’Aquila), in a central region (Abruzzi) of the Italian peninsula, and two
others in eastern Sicily on the same host plant. Specimens from Opi were previously
identified and cited (Barbagallo and Patti 1998) as A. subterranea but are now identified
as A. shaposhnikovi.
Volume 135 THE CANADIAN ENTOMOLOGIST 847
Character No. Acronym Character
1 bl Body length
2 af Antennal flagellum length (joints III–VI)
3 VIb Basal part of antennal joint VI length
4 VId Distal part of antennal joint VI length
5 fw Frontal width (interantennal distance)
6 urj Ultimate rostral joint length
7 IIht Length of joint II of hind tarsus
8 sl Siphuncular length
9 sw Siphuncular median width
10 cl Caudal length
11 rs Number of spinules rows on siphunculi
12 ot Number of occipital tubercles
13 ft Number of frontal tubercles
14 st1–5 Number of abdominal spinal tubercles on segments 1–5
15 st6–8 Number of abdominal spinal tubercles on segments 6–8
16 mt Number of abdominal marginal tubercles (both side)
TABLE 2. Morphometric characters used for discriminant function analysis in six
species of the genus Anuraphis.
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Distribution
The new aphid species is only known so far from Italy, including Sicily. If the
aphid is definitely proven to have only the two above-named species of Apiaceae
(whose distribution ranges from the western Mediterranean eastwards to the Balkan
peninsula and the Aegean region (Tutin 1964–1980; Pignatti 1982)), as its usual sec
-
ondary hosts, it would be restricted to the same geographical territories.
Biology
The aphid colonies develop, as in other species of the same genus, on subterra
-
nean parts (root collar area and leaf petioles) and inside the inner sheet surface of basal
leaves; they are usually ant-attended. The aphid life cycle is not known. Colonies on the
two named Apiaceae were usually collected during June in the field and reared in the
laboratory (on excised parts of the host plant) up to about mid-July. Early colonies de
-
tected at the beginning of June in the field were frequently composed of single alatae
with a few nymphs; this alatae may be coming from a primay host. Pyrus communis L.,
and two other wild species (i.e., Pyrus pyraster Burgsd. and Pyrus amygdaliformis
Vill.) are rather common in the area (western and eastern Sicily), but no Anuraphis
spp. pseudogalls have yet been found on their leaves. Vegetative aerial parts of the sec
-
ondary host plants ( M. pastinacea and O. chironium) dry up at the end of June and
848 THE CANADIAN ENTOMOLOGIST November/December 2003
Length (mm) of antennal joint VI
Species and morph
Body
length
(mm)
Antennal
flagellum
length (mm) Basal part Distal part
Urj* length
(mm)
A. cachryos
Apterae (n = 20) 1.86–2.97 0.58–1.16 0.064–0.104 0.186–0.352 0.193–0.244
Alatae (n = 10) 2.01–2.81 1.03–1.37 0.088-.0116 0.290–0.392 0.200–0.240
A. catonii
Apterae (n = 20) 1.63–2.37 0.75–1.17 0.080–0.108 0.272–0.400 0.200–0.240
Alatae (n = 10) 1.50–2.15 0.97–1.35 0.084–0.116 0.320–0.420 0.190–0.220
A. farfarae
Apterae (n = 20) 2.15–3.20 1.04–1.86 0.080–0.152 0.310–0.480 0.268–0.340
Alatae (n = 10) 2.09–2.89 1.36–1.82 0.100–0.136 0.400–0.568 0.264–0.320
A. pyrilaseri
Apterae (n = 20) 2.07–3.30 0.95–1.46 0.072–0.120 0.280–0.440 0.240–0.320
Alatae (n = 10) 1.99–2.99 1.19–1.67 0.080–0.104 0.408–0.560 0.240–0.288
A. shaposhnikovi
Apterae (n = 80) 2.07–2.94 0.97–1.35 0.084–0.120 0.320–0.440 0.180–0.240
Alatae (n = 20) 1.80–2.58 1.16–1.36 0.088–0.120 0360–0.480 0.174–0.216
A. subterranea
Apterae (n = 80) 2.32–3.46 1.01–1.92 0.092–0.148 0.312–0.532 0.240–0.344
Alatae (n = 20) 1.86–3.47 1.24–1.93 0.100–0.144 0.400–0.560 0.214–0.320
NOTE: All specimens were randomly selected from different available samples.
* Urj is the ultimate rostral joint length and IIht is the length of joint II of the hind tarsus.
†
Number of occipital + frontal tubercles on the head.
‡
Number of abdominal spinal tubercles from tergite I to tergite V + from tergite VI to tergite VIII.
§
Number of abdominal marginal tubercles from urite I to urite V + from urite VI to urite VII.
TABLE 3. Range of some relevant morphometric data in six species of the genus Anuraphis.
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during July, so no aphid colonies can survive at this time unless they move down to the
roots (these plants being perennial), but this has not yet been observed.
Comments
It seems that the two known host-plant species (M. pastinacea and O. chironium)
slightly affect some morphological features (i.e., the length of last rostral joint and the
number of abdominal spinal tubercles) in A. shaposhnikovi where specimens living on
Opopanax spp. appear to be morphologically more similar to A. subterranea; the rela
-
tionship seems to be even clearer when specimens of the latter species collected on
Pastinaca spp. are compared.
Other western Palaearctic species of the genus Anuraphis hardly can be confused
with the new taxon here described (see key for comparative differences). Anuraphis
catonii has a smaller body size and a proportionally longer ultimate rostral joint com
-
pared with A. shaposhnikovi; therefore, the urj / body length ratio (Table 4) helps to
separate the two species in most cases, as the ratio is usually 0.10 or greater in
A. catonii (range 0.092–0.126 in apterae and 0.100–0.140 in alatae) and less than 0.10
in A. shaposhnikovi (range 0.070–0.098 in apterae and 0.084–0.105 in alatae). In any
case, A. catonii has a distinctive ecological niche (living it, as far as it is known, only on
Pimpinella spp.) and cannot be confused with the relatively different A. shaposhnikovi.
Anuraphis cachryos usually lack spinal tubercles on abdominal segment VI and VII,
whereas its tibiae are full pigmented (see key for their separation).
Volume 135 THE CANADIAN ENTOMOLOGIST 849
Number of
IIht* length
(mm)
Siphuncular
length
(mm)
Rows of
spinules on
siphunculi
Tubercles
on head
†
Abdominal
spinal
tubercles
‡
Abdominal
marginal
tubercles
§
0.120–0.163 0.120–0.190 18–30 0–2 + 0–2 1–8 + 0–1 10 + 2–4
0.130–0.160 0.138–0.180 20–28 1–2 + 0–2 1–9 + 0–1 10 + 3–4
0.120–0.144 0.120–0.190 18–28 0–2 + 0–2 3–10 + 0–4 10 + 2–4
0.120–0.140 0.120–0.190 12–24 0–2 + 0–2 2–8 + 0–3 10 + 3–4
0.132–0.180 0.210–0.320 34–55 2 + 0 6–9 + 0–2 10 + 0
0.140–0.168 0.185–0.272 30–46 2 + 0 5–10 + 0–1 10 + 0
0.140–0.172 0.200–0.280 23–35 1–2 + 0 2–8 + 0–1 9–10 + 0
0.152–0.180 0.180–0.240 23–34 0–2 + 0 3–7 + 0 10 + 0
0.140–0.170 0.160–0.200 20–34 2–10 + 0–6 10 + 3–4
0.140–0.160 0.152–0.200 18–28 1–2 + 1–2 1–9 + 0–5 10 + 3–4
0.140–0.200 0.180–0.250 27–44 2 + 2 9–10 + 2–5 10 + 3–4
0.144–0.190 0.180–0.240 24–40 2 + 1–2 8–10 + 2–4 10 + 4
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Morphometric analysis
Materials and methods
Apterous and alate viviparous females used for the morphological study are
mounted on permanent slides and belong to species and samples below. They are stored
in the collection of the first author except where otherwise indicated (the number of
specimens for each sample used for multivariate analysis is given in brackets).
Anuraphis cachryos: Cachrys sicula, Caltagirone (Catania), Sicily, 14.iv.1977 [4
apterous], 8.ix.1978 [3 apterous] and 1.v.1978 [2 alatae]; Cachrys libanotis,
Fiumefreddo (Catania), Sicily, 30.iv.1977 [2 apterous]; Echinophora tenuifolia,
Caltagirone (Catania), Sicily, 13.x.1978 [2 apterous and 1 alatae] and 15.ii.1979 [1
apterous]; ibid Palagiano (Taranto), Italy, 14.x.1992 [3 apterous and 4 alatae]; Ferula
communis, S Croce Camerina (Ragusa), Sicily, 21.v.1998 [5 apterous and 3 alatae].
Anuraphis catonii: Pimpinella peregrina, Limina (Messina), Sicily, 7.v.1981 [8
apterous and 4 alatae]; ibid Erice (Trapani), Sicily, 10.vi.1998 [7 apterous and 6 alatae];
Pimpinella saxifraga, Praha (Czech Republic), 2.ix.1969 [4 apterous] (J Holman collec
-
tion); Pimpinella major, Brloh (Czech Republic.), 16.vii.1991 [1 apterous] (J Holman
collection).
Anuraphis farfarae: Petasites albus, Opi (L’Aquila), Italy, 26.vi.1996 [3 apterous
and 1 alatae]; Petasites georgicus Lazarevskaja (ex URSS), 29.x.1974 [2 apterous] (J
Holman collection); Petasites sp., Sambuco (Cuneo) Italy, 12.vi.1998 [2 apterous and 5
alatae]; Tussilago farfara, Sepino (Campobasso), Italy, 18.vi.1993 [2 apterous]; ibid
Rocca S Maria (Teramo) Italy, 28.vi.1996 [8 apterous and 1 alatae]; ibid Lauria
(Potenza), Italy, 23.vi.1998 [3 apterous and 1 alatae]; ibid Praha (Czech Republic),
18.x.1971 [2 alatae] (J Holman collection).
Anuraphis pyrilaseri: Thapsia garganica, Lentini (Siracusa), Sicily, 24.v.1977 [4
apterous and 2 alatae]; ibid Fiumefreddo (Catania), Sicily, 30.v.1998 [2 apterous and 2
alatae]; Ferula communis, Bortigali (Nuoro), Sardinia, 16.iv.1978 [4 apterous and 1
alatae]; ibid Randazzo (Catania), Sicily, 9.xii.1997 [2 apterous and 1 alatae]; Ferulago
campestris, Bronte (Catania), Sicily, 19.iv.1995 [2 apterous]; Laserpitium siler,
Sambuco (Cuneo), Italy, 26.vi.1997 [2 apterous and 2 alatae]; L. hispidum, Lazarevskaja
(ex URSS), 29.x.1974 [1 apterous and 1 alatae) (J Holman collection); Laser trilobum,
Armenia (ex URSS), 22.vi.1965 (1 apterous and 1 alatae) (GCh Shaposhnikov collec
-
tion); Cachrys ferulacea, Roccella Valdemone (Messina), Sicily, 22.v.1999 [2 apterous].
Anuraphis shaposhnikovi: Magydaris pastinacea, Buseto Palizzolo (Trapani),
Sicily, 3.vi.1997 [13 apterous and 1 alatae], 9.vi.1999 [3 apterous] and 5.vi.2001 [11
apterous]; ibid Godrano (Palermo), Sicily, 4.vi.1999 [1 apterous and 1 alatae]; ibid
Marineo (Palermo), Sicily, 2.vi.2000 [20 apterous and 9 alatae] and 17.vii.00 [9
apterous and 5 alatae]; Opopanax chironium, Opi (L’Aquila), 26.vi.1996 [7 apterous
and 6 alatae]; Francavilla di Sicilia (Messina), 22.v.2001 and 8.vi.2001 [16 apterous].
Anuraphis subterranea: Heracleum sphondylium, samples with the following
collection data: Marsworth (Herts.), England, 11.ix.1975 [12 apterous and 3 alatae];
Collalbo (Bolzano), Italy, 2.ix.1992 [8 apterous and 2 alatae]; Vastogirardi (Isernia), It
-
aly, 19.vi.1993 [3 apterous]; S Gregorio Matese (Caserta), Italy, 18.vi.1993 [4 apterous
and 1 alatae]; Udine, Italy, 18.vi.1994 [6 apterous]; Opi (L’Aquila), Italy, 26.vi.1996 [5
apterous]; Sambuco (Cuneo), Italy, 12.vi.1998 [8 apterous and 3 alatae]; Marano Equo
(Rome), Italy, 03.vi.1999 [1 apterous and 1 alatae]; Praha (Czech Republic), 19.vii.1999
[8 apterous and 1 alatae]; Heracleum pyrenaicum, Vigo di Fassa (Trento), Italy,
3.ix.1992 [7 apterous]; H. pyrenaicum subsp. cordatum, Isnello (Palermo), Sicily,
11.vi.1999 [3 alatae]; ibid Novara di Sicilia (Messina) [4 apterous and 2 alatae], and
Roccella Valdemone (Messina), Sicily, 18.vi.1999 [5 apterous]; Floresta (Messina) Italy,
850 THE CANADIAN ENTOMOLOGIST November/December 2003
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Volume 135 THE CANADIAN ENTOMOLOGIST 851
Species and
morph
Antennal
flagellum
length/body length
Urj
length/IIht
length*
Urj
length/siphuncular
length
Urj length/body
length
Siphuncular
length/body
length
Siphuncular
length/siphuncular
diameter
†
Siphuncular
length/IIht length VId/VIb
‡
A. cachryos
Apterae 0.28–0.45 1.25–1.87 1.13–1.67 0.07–0.11 0.06–0.07 1.82–2.40 1.00–1.20 2.41–3.65
Alatae 0.47–0.57 1.32–1.63 1.23–1.68 0.08–0.10 0.06–0.07 1.87–2.67 0.96–1.13 2.46–3.80
A. catonii
Apterae 0.35–0.56 1.43–1.77 1.31–1.72 0.09–0.13 0.06–0.08 1.76–2.71 0.86–1.30 3.00–4.00
Alatae 0.53–0.72 1.52–1.77 1.15–1.67 0.10–0.14 0.07–0.09 1.82–3.25 0.92–1.36 3.33–4.62
A. farfarae
Apterae 0.48–0.69 1.62–2.29 0.97–1.33 0.09–0.15 0.08–0.13 2.80–3.75 1.50–2.00 3.10–4.28
Alatae 0.61–0.70 1.62–2.04 1.00–1.43 0.10–0.13 0.06–0.11 3.08–3.50 1.32–1.62 3.45–4.73
A. pyrilaseri
Apterae 0.38–0.56 1.60–2.12 1.00–1.55 0.08–0.14 0.07–0.10 2.39–3.75 1.30–1.63 2.80–5.00
Alatae 0.53–0.71 1.50–1.84 1.09–1.56 0.08–0.14 0.07–0.10 3.00–3.87 1.14–1.38 4.48–5.65
A. shaposhnikovi
Apterae 0.35–0.54 1.25–1.60 1.00–1.34 0.07–0.10 0.06–0.09 2.00–3.10 1.00–1.40 3.10–4.32
Alatae 0.46–0.72 1.24–1.54 0.98–1.27 0.08–0.11 0.07–0.09 2.25–3.50 1.05–1.34 3.17–4.80
A. subterranea
Apterae 0.38–0.55 1.50–2.11 1.26–1.62 0.09–0.12 0.06–0.09 1.92–3.22 1.05–1.45 2.96–4.87
Alatae 0.55–0.74 1.38–1.85 1.13–1.56 0.08–0.12 0.07–0.10 2.57–3.56 1.00–1.45 3.67–4.63
* Urj is the ultimate rostral joint length and IIht is the length of joint II of the hind tarsus.
†
Siphuncular diameter at their median length.
‡
VId and VIb are the lengths of the distal and basal parts, respectively, of antennal joint VI.
TABLE 4. Variation in range of some relevant morphometric ratio values in six species of the genus Anuraphis (derived from linear values in Table 3).
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18.vi.1999 [1 alatae]; Pastinaca sativa, Barton Mills (Suff.), England, 24.vii.1975 [1
alatae]; Sepino (Campobasso), Italy, 18.vi.1993 [2 apterous]; Scontrone (L’Aquila), It
-
aly, 24.vi.1996 [3 alatae]; Praha, Czech Republic, 23.vii.1986 [5 apterous] (J Holman
collection); Ceský Krumlov, Czech Republic, 8.vi.1984 [1 alatae] (J Holman collec
-
tion); Mount Vitoša, Bulgaria, 4.ix.1988 [2 apterous] (J Holman collection).
852 THE CANADIAN ENTOMOLOGIST November/December 2003
FIGURE 5. Plot of the first and second discriminant functions of apterae (a) and alatae (b) specimens of
Anuraphis pyrilaseri (
), Anuraphis farfarae ( ), Anuraphis cachryos ( ), Anuraphis catonii ( ),
Anuraphis subterranea (
), and Anuraphis shaposhnikovi ( ). Ellipse indicates 95% confidence limits.
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Specimens were measured as illustrated by Ilharco and van Harten (1987) using
an optical compound microscope that was equipped with an eyepiece micrometer.
Discriminant function analysis has been extensively used as a powerful tool to
discriminate and clarify the taxonomic status of related insect species (Blackman and
Peterson 1986; Blackman 1992; Tizado and Nieto Nafria 1994; Sanmartin and Martin
Piera 1999; Williams and Langor 2001). It is applied here to morphometric data of six
species of the genus Anuraphis (namely, A. cachryos, A. catonii, A. farfarae,
A. pyrilaseri, A. shaposknikovi, and A. subterranea) to determine their degree of diver
-
gence. For this purpose, 16 continuous (1–10) and discrete (11–16) variable characters,
both for apterae and alatae viviparous females from secondary hosts, were selected on
the basis of their perceived taxonomical importance (Table 2). Anuraphis ferulae was
not included in the multivariate analysis because it was not available for study; its origi
-
nal description (Shaposhnikov 1995) is based on 8 apterae and 3 alatae specimens alto
-
gether (which apparently are the only specimens as yet known), of which we have seen
an apterous viviparous female collected on Ferula sp., Varzob, Tajikistan (ex URSS),
16.x.1988 (GCh Shaposhnikov collection).
These characters were measured on randomly selected 20 apterae and 10 alatae of
each of these 6 taxa. Wilks’
λ
was applied to evaluate the significance of the analyses
and cluster analysis of Mahalanobis’ distances, calculated on the centroid values in the
discriminant space, was used to construct a dendrogram of affinities among the six spe
-
cies.
To find a discriminant linear function between the phenotypically closely related
species A. shaposhnikovi and A. subterranea, an additional 80 apterae and 22 alatae for
each taxon were measured and submitted separately to the discriminant function
Volume 135 THE CANADIAN ENTOMOLOGIST 853
Group Function Eigenvalue
Variance
explained (%)
Cumulative
variance
Canonical
correlation
Apterae
1 1 40.91 72 72 0.99
2 6.49 12 84 0.93
3 0.22 8 92 0.91
4 3.32 6 98 0.87
5 1.09 2 100 0.72
2 1 13.42 62 62 0.97
2 6.16 29 91 0.93
3 0.05 9 100 0.81
3 1 9.33 100 100 0.95
Alatae
1 1 69.07 76 76 0.99
2 10.48 11 87 0.93
3 5.95 7 94 0.85
4 3.91 4 98 0.81
5 1.90 2 100 0.72
2 1 7.37 58 58 0.94
2 3.13 25 83 0.87
3 2.11 17 100 0.82
3 1 5.57 100 100 0.92
TABLE 5. Canonical discriminant functions of apterae and alatae morphs of six species of
the genus Anuraphis.
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analysis. This analysis generated a linear discriminant function that was used to deter-
mine which species each specimen belonged to.
The statistical analyses were performed on untransformed data using
STATISTICA version 5.1 (StatSoft Inc. 1996).
Results
Morphometric data of most relevant characters for the six nominal western Euro-
pean species of the genus Anuraphis show a high degree of overlap between species
(Tables 3, 4). Nevertheless, the range of their secondary hosts is well known for each
species, and in most cases may strongly contribute to species identification whenever
morphological features lead to reasonable doubts.
The discriminant function analysis on apterae of the six Anuraphis species based
on the selected 16 characters (Table 2) provided five functions (Wilks’
λ
41.939
= 0.0006,
P < 0.00001). The six species were discriminated in multivariate space, and also
showed at the same time the morphological similarity among species (Fig. 5a). The first
discriminant function explains 72% of the total variation (apterae, Table 5). Here,
among the six analyzed species, two main groups of taxa can be recognized, corre
-
sponding to the same results of a genetical analysis (S Barbagallo, I Patti, GE Cocuzza,
and MC Mosco, unpublished data). Anuraphis farfarae and A. pyrilaseri belong to
Group 1, whereas the remaining four species (A. cachryos, A. catonii, A. shaposhnikovi,
A. subterranea) appear to be more closely related to each other (Group 2), even though
they were separated into two subgroups (Fig. 5a). The second, third, fourth, and fifth
functions explain 11%, 8%, 6%, and 2%, respectively, of the total variation and are
helpful in discriminating the Group-2 species.
Group-1 species are characterized, compared with those of Group 2, by longer
siphunculi and a reduced number of body tubercles (i.e., on head and post-siphuncular
abdominal segments).
The dendrogram based on Mahalanobis’ distance data clearly shows the basic di
-
chotomy (Fig. 6). Nevertheless, in Group 2, A. shaposhnikovi shows a closer relationship
854 THE CANADIAN ENTOMOLOGIST November/December 2003
FIGURE 6. Dendrogram of cluster-analysis results based on Mahalanobis’ generalized distances in
apterae of the six species of the genus Anuraphis showing their morphological relationships.
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to A. cachryos, and particularly to A. catonii, than to A. subterranea (but see taxonomic
discussion in the description of A. shaposhnikovi).
A discriminant function analysis of alatae (Wilks’
λ
41.939
= 0.00002, P < 0.00001)
also discriminated the species into two main groups, represented by A. farfarae and
A. pyrilaseri in Group 1 and the four other species in Group 2. Moreover, the first
discriminant function separated, inside the latter group, A. subterranea and A. cachryos
from the overlapping A. shaposhnikovi and A. catonii (alatae, Table 5; Fig. 5b). The
Volume 135 THE CANADIAN ENTOMOLOGIST 855
FIGURE 7. Plot of the first and second discriminant functions of apterae (a) and alatae (b) specimens of
Anuraphis cachryos (
), Anuraphis catonii ( ), Anuraphis subterranea ( ), and Anuraphis
shaposhnikovi (
). Ellipse indicates 95% of the confidence limits.
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other discriminant functions clearly separate the latter two species, and the classifica
-
tion functions correctly placed 100% of specimens into the appropriate group.
When the discriminant function analysis was applied to Group-2 apterae, exclud
-
ing A. pyrilaseri and A. farfarae (Wilks’
λ
34.55
= 0.0033, P < 0.00001), the species were
separated by values of the first discriminant function with a slight overlap between
A. catonii and A. shaposhnikovi (apterae, Table 5; Fig. 7a); nevertheless, the second
discriminant function gave a clear separation between them.
856 THE CANADIAN ENTOMOLOGIST November/December 2003
Coefficient Predicted group
Character*
A.
subterranea
A.
shaposhnikovi Actual group
A.
subterranea
A.
shaposhnikovi
Percentage
correctly
classified
Apterae
Urj 1101.8 677.8 A. subterranea 80 0 100
Cl 1867.2 1638.7 A. shaposhnikovi 0 80 100
VIb –378.6 –132.4 Total 80 80 100
St1–5 –0.23 –1.1
Constant –320.8 –218.7
Alatae
Urj –36.9 –235.4 A. subterranea 20 2 90.91
Cl 0.6 –1.0 A. shaposhnikovi 0 22 100
VIb 211.9 157.2 Total 20 24 95.45
St1–5 1528.7 1726.8
Constant –178.3 –134.1
* Urj is the ultimate rostral joint length, cl is the caudal length, VIb is the length of the basal part of antennal joint VI, and
St1–5 is the number of abdominal spinal tubercles on segments 1–5.
TABLE 6. Fisher’s linear functions and percentages of correctly classified apterae and alatae morphs of
Anuraphis subterranea and Anuraphis shaposhnikovi.
FIGURE 8. General aspect of apterae from secondary host plants (specimen mounted on slides) of
Anuraphis farfarae (a) and Anuraphis pyrilaseri (b). Scale bars = 0.5 mm.
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Similarly, a discriminant function analysis performed on Group-2 alatae (Wilks’
λ
18.12
= 0.00093, P < 0.00001) separated species by values of the second discriminant
function (alatae, Table 5; Fig. 7b).
In the discriminant function analysis of apterae of A. subterranea and
A. shaposhnikovi (Wilks’
λ
176.02
= 0.097, P < 0.00001), values of standardized coeffi
-
cients (derived from 80 specimens for each species) showed that urj had the greatest
weight, followed by VIb, cl, and st1–5 (Table 6). The classification provided by the
discriminant function analysis on the basis of the above-mentioned characters correctly
placed 100% of the specimens into the appropriate group; here, the function based only
Volume 135 THE CANADIAN ENTOMOLOGIST 857
FIGURE 9. General aspect of apterae viviparae from secondary host plants (specimen mounted on
slides) of Anuraphis cachryos (a), Anuraphis catonii (b), Anuraphis ferulae (c), and Anuraphis
subterranea (d). Scale bars = 0.5 mm.
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on urj and st1–5 identified 99.4% of the specimens and can be used as an easy tool for
taxonomic identification:
Discrimination function score = 1102urj – 0.23(st1–5) [1]
where a score greater than 271 classifies the specimen as A. subterranea and a score
less than 271 classifies the specimen as A. shaposhnikovi.
The discriminant function analysis performed on alatae (Wilks’
λ
34.342
= 0.152, P <
0.00001) of the two taxa (comparing 22 specimens for each taxon) also explained 100%
of the variation, and the two species were well discriminated by urj, st1–5, VId, and IIht
(Table 6). The classification function correctly placed 95.45% of the specimens into the
appropriate group:
Discrimination function score = –36.9urj + 0.6(st1–5) + 211.9VId + 1528.7IIht [2]
where a score greater than 329 classifies the specimen as A. subterranea and a score
less than 329 classifies the specimen as A. shaposhnikovi.
Key to Anuraphis spp. viviparous morphs on secondary host
As already mentioned, all currently recognized species of the genus Anuraphis,
including the new species here described, show morphological overlap as can be see
from the morphometric data (except for A. ferulae) (Tables 3, 4).
Shaposhnikov (1964) provided a key to the four species that were known at that
time on both primary and secondary hosts; more recently, Blackman and Eastop (1994)
provided a key to the morphs (fundatrices and alate emigrants) from the primary host
only. Heie (1992) also keyed, described, and illustrated three (A. catonii, A. farfarae,
A. subterranea) species. An illustrated description of A. cachryos with figures on
A. pyrilaseri, including comparative morphological notes with other species, was given
by Barbagallo and Stroyan (1982). Biometric accounts on A. ferulae were given by
Shaposhnikov (1995) who added a key to separate it from the group of remaining spe-
cies of the genus Anuraphis.
Key to Anuraphis species
Apterae viviparae
1. Head as a rule with a single pair of tubercles (occipitals only). Abdominal marginal tubercles confined
to the abdominal segments I–V. Siphunculi 0.20–0.32 mm long, 0.075–0.13 times the body length,
2.40–3.75 times as long as wide at middle length, and bearing 23–55 rows of spinules (these can be
counted easily through the number of spicules along the marginal side of siphunculi). Siphuncular
length / IIiht length ratio 1.30–2.00 ............................2
— Head as a rule with two pairs of tubercles (both occipital and frontal pairs), though they could be occa
-
sionally inconspicuous or even individually lacking from one or both pairs. Abdominal marginal tuber
-
cles on abdominal segments I–V only (A. ferulae) or up to abdominal segment VII (other species).
Siphunculi 0.10–0.25 mm long, 0.046–0.086 times the body length, mostly 1.30–2.80 times (only in
A. subterranea up to 3.20 times) as long as wide at their middle length, and bearing usually 16–44
rows of spinules. Siphuncular length / IIht length ratio 0.82–1.42 ...............3
2. Siphunculi usually dark (as a rule darker than distal part of hind tibiae), 0.21–0.32 mm long, 1.50–2.00
times the second hind tarsomer and densely adorned with 34–55 rows of rather well-distinct and
sclerified spinules. Abdominal segments anterior to siphunculi (unless in small specimens) frequently
adorned with more or less sclerified dorsal bands, even much perforated and broken along the median
line, but particularly expanded and coalescing in alatoid specimens; abdominal spinal tubercles ranging
7–12 in number and frequently extended (though on one side only) up to abdominal segments VI and
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(or) VII–VIII. Aphid dark brown in life, sometimes with a tinge of dark green, living on root collar
parts of Asteraceae of the genera Tussilago, Petasites, and Adenostyles. Widely distributed in western
Palaearctic; also recorded (but probably erroneously) for North America (Fig. 8a).........
.................................A. farfarae (Koch, 1854)
— Siphunculi brown to dark brown (on average, as dark as the distal part of hind tibiae), 0.20–0.28 mm
long, 1.30–1.63 times joint II of hind tarsus, and bearing 23–35 rows of spinules, which basally tend to
coalesce into serrulated embrications. Dorsal abdominal sclerifications variable, from muscle sclerites
alone, on segments anterior to siphunculi, to a more or less compact subtrapezoid macula (mostly pres
-
ent on abdominal segments IV–VI in alatoid specimens). Abdominal spinal tubercles 3–9 in number
and confined, as a rule, up to abdominal segment V (rarely up to abdominal segment VI). Aphid vari
-
able in colour, from pale green to brownish yellow–orange, but not dark brown. Mostly common (in
southern Italy) on the genera Ferula and Thapsia, but living also on several other genera of Apiaceae,
including Laserpitium, Laser, Ferulago, Cachrys, Hippomarathrum, and Opopanax. Distributed from
southwestern Europe to Caucasus and Crimea (Fig. 8b)......A. pyrilaseri Shaposhnikov, 1950
3. Siphunculi short (0.10–0.15 mm), just a little longer than their basal width and 1.30–1.80 times their
diameter at middle length, adorned with 19–28 rows of spinules. Abdominal tubercles, either spinals or
marginals, confined to abdominal segments I–V (i.e., abdominal segments VI and VII lack both spinal
and marginal tubercles). Antennal flagellum (i.e., joints III–VI) short, about 0.30 times the body length.
On Ferula spp., presently known from Tajikistan (Fig. 9c)......A. ferulae Shaposhnikov, 1995
— Siphunculi 0.12–0.25 mm long, corresponding to 1.76–3.20 times their median diameter, and adorned
with 18–44 rows of spinules. Abdominal marginal tubercles present on abdominal segments I–VII, al
-
though occasionally lacking on one or both sides of abdominal segment VI (A. cachryos, A. catonii).
Antennal flagellum 0.28–0.56 times the body length, but only rarely (some specimens of A. cachryos)
less than 0.35 times the body length ...........................4
4. Siphunculi 0.12–0.19 mm long, but if more than 0.15 m the urj length / siphuncular length ratio is
1.30–1.70 and the aphid lives on Pimpinella spp. (A. catonii), or abdominal spinal tubercles range 1–9
in number, lacking usually from abdominal segments VI and VII, and tibiae are fully pigmented
(A. cachryos).....................................5
— Siphunculi 0.16–0.25 mm long, but if less than 0.18 mm the urj length / siphuncular length ratio is
1.00–1.34, but only exceptionally (4% of specimens seen) more than 1.27 and the aphid lives on other
genera of Apiaceae, abdominal spinal tubercles range 4–16 in number, usually present on abdominal
segments VI and VII, and tibiae are paler in the middle than at their basal and distal parts
(A. shaposhnikovi)...................................6
5. Maximum basal diameter of siphunculi ranging 1.92–3.80 (usually 2.5–3.0) times their minimum distal
width (just behind the flange). Abdominal spinal tubercles 1–9 in number (most frequently 4–6), usu
-
ally confined to tergites I–V. Tibiae uniformly well pigmented. Abdominal marginal tubercles compara
-
tively smaller (i.e., that on abdominal segment VII ranges 1.0–2.0 times the maximum diameter of
stigmal pore on that segment). Mostly on Cachrys spp. and Echinophora spp., occasionally also on
Ferula spp. Widely Mediterranean in distribution; recorded in Middle East (Fig. 9a)........
..........................A. cachryos Barbagallo et Stroyan, 1982
— Siphuncular maximum basal diameter ranging 1.20–2.15 (but only exceptionally more than 2.00) times
their minimum distal width. Abdominal spinal tubercles 3–13 in number and usually extended up to ab
-
dominal segment VII. Tibiae usually more or less paler in the middle than at their basal and distal
parts. Abdominal marginal tubercles as a rule comparatively larger (i.e., those on abdominal segment
VII range 1.9–2.8 times the maximum diameter of the near stigmal pore). Living on Pimpinella
spp. (most frequently P. saxifraga, as well as P. major, P. peregrina, and perhaps others). Widely dis
-
tributed in western Palaearctic (Fig. 9b)............A. catonii Hille Ris Lambers, 1935
6. Urj 0.18–0.24 mm and urj/IIht ratio 1.25–1.60 but rarely (up to 15% of specimens) more than 1.50. Ab
-
dominal spinal tubercles frequently lacking on one or both sides from some abdominal segments ante
-
rior the siphunculi (they are usually present on abdominal segments VI and VII, and occasionally to
abdominal segment VIII). Tibiae more or less pale in the middle. Value of the function 1102urj –
0.23(st1–5) less than 271. Collected on Magydaris pastinacea and Opopanax chironium in Sicily and
central Italy (Fig. 3) ........................A. shaposhnikovi sp. nov.
— Urj 0.24–0.34 mm and urj/IIht ratio 1.50–2.10, but rarely (up to 6% of specimens) less than 1.60. Ab
-
dominal spinal tubercles set as a rule in a complete series from abdominal segments I to VII, and occa
-
sionally including abdominal segment VIII. Tibiae usually uniformly pigmented from base to apex.
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860 THE CANADIAN ENTOMOLOGIST November/December 2003
Value of the function 1102urj – 0.23(st1–5) more than 271. Living on the genera Heracleum and
Pastinaca. Widely distributed from western Europe eastwards to central Asia, western Siberia, and the
Middle East. Recorded for North America, but probably erroneously as for A. farfarae (Fig. 9d)
..............................A. subterranea (Walker, 1852)
Alatae viviparae
1. Head with a single pair of tubercles (occipitals). Abdominal marginal tubercles present on abdominal
segments I–V only. Siphunculi 0.18–0.27 mm, adorned with 23–46 rows of spinules .......2
— Head usually with both occipital and frontal tubercles (two pairs), although inconstantly present as a
full set and sometimes being small in specimens with siphunculi up to 0.18 mm (but see couplet 4).
Abdominal marginal tubercles present on abdominal segments I–V only (A. ferulae) or on abdominal
segments I–VII (other species). Siphunculi 0.12–0.24 mm and bearing 12–40 rows of spinules . . . 3
2. Siphunculi 0.19–0.27 mm, 1.32–0.162 times IIht, and bearing 30–46 rows of spinules. Antennal joint
VI with the distal part 3.45–4.75 times its basal one. On Asteraceae of the genera Tussilago, Petasites,
and Adenostyles. Western Palaearctic .................A.farfarae (Koch, 1854)
— Siphunculi 0.18–0.24 mm, 1.14–1.38 times IIht, and bearing 23–34 rows of spinules. Distal part of
antennal joint VI 4.50–5.70 times the basal part of the same joint. On several genera of Apiaceae
(Ferula, Thapsia, Laserpitium, Laser, Ferulago, and others). Southwestern Europe, Caucasus, and
Crimea ..........................A. pyrilaseri Shaposhnikov, 1950
3. Marginal and spinal abdominal tubercles confined to abdominal segments I–V only. Secondary rhinaria
(29–43 in number) present on antennal joint III only. Siphunculi 0.12–0.15 mm long, 1.8–2.4 times as
long as wide in the middle, and bearing 21–29 rows of spinules. On Ferula spp., Tadjikistan
(morphometric data here indicated for A. ferulae follow Shaposhnikov’s (1995) description).....
..............................A. ferulae Shaposhnikov, 1995
— Marginal abdominal tubercles usually present from abdominal segments I to VII. Secondary rhinaria
present on antennal joints III and IV, occasionally also on joint V. Siphunculi 1.8–3.6 as long as wide
in the middle .....................................4
4. Siphunculi 0.12–0.19 mm long and bearing 12–28 rows of spinules. If siphunculi are 0.15 mm or more,
urj/IIht ratio more than 1.50 and antennal joint III has 45–65 secondary rhinaria (A. catonii), or abdom-
inal segments VI and VII usually lack spinal tubercles and tibiae are full pigmented (A. cachryos).
Head tubercles present or not as a full set (i.e., one or both tubercles for each pair may not be present).
Comparatively smaller specimens (body 1.50–2.80 mm long) .................5
— Siphunculi 0.16–0.24 mm long, having 18–40 rows of spinules. If siphunculi are 0.19 mm or less,
urj/IIht ratio usually less than 1.50 and tibiae are rather paler in the middle than apically
(A. shaposhnikovi), or antennal joint III has 58–118 (usually not less than 65) secondary rhinaria and
spinal tubercles on abdominal segments VI and VII are present (A. subterranea). Head tubercles always
present as a complete set (two pairs). Usually more large specimens (body 1.80–3.45 mm long). . . 6
5. Urj/IIht ratio 1.32–1.65. Abdominal spinal tubercles usually confined to abdominal segments I–V and
only exceptionally extended up to abdominal segment VI and (or) VII. Tibiae fully pigmented. Body
2.00–2.80 mm long. Living on Cachrys spp., Echinophora spp., and occasionally on other genera of
Apiaceae but not on Pimpinella spp. From Mediterranean to Middle East in distribution ......
..........................A. cachryos Barbagallo et Stroyan, 1982
— Urj/IIht ratio 1.50–1.75. Abdominal spinal tubercles present as a rule on abdominal segment VI and
(or) VII; however, they may lack on some segments anterior the siphunculi. Tibiae usually rather pale
in the middle with darker basal and apical parts. Body 1.50–2.15 mm long. On Pimpinella spp. Western
Palaearctic ........................A. catonii Hille Ris Lambers, 1935
6. Urj/IIht ratio 1.24–1.54, but rarely exceeding 1.50. Abdominal spinal tubercles 2–13 in number, usually
lacking on some segments anterior to the siphunculi, although as a rule present either on abdominal
segments VI and (or) VII. Secondary rhinaria on antennal joints III (both antennae) 96–185. Value of
the function –36.9urj + 0.6(st1–5) + 211.9VId + 1528.7IIht less than 329. On Magydarys pastinacea
and Opopanax chironium. Presently known from Sicily and central Italy. . . A. shaposhnikovi sp. nov.
— Urj/IIht ratio 1.36–1.85, but only exceptionally less than 1.50. Abdominal spinal tubercles 11–14 in
number, usually present, at least on one side, from abdominal segments I to VII inclusive. Secondary
rhinaria on antennal joints III (both antennae), 120–230. Value of the function –36.9urj + 0.6(st1–5) +
211.9VId + 1528.7IIht greater than 329. On Heracleum spp. and Pastinaca spp. Western Palaearctic
..............................A. subterranea (Walker, 1852)
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Conclusion
Biometric analysis including a multivariate processing system carried out on six
species of the genus Anuraphis shows that this genus is a homogeneous taxonomic
group where each specific component, although reproductively isolated, shares with
others several characteristics, owing to its low degree of morphological differentiation.
Nevertheless, on the basis of their morphology, which includes multivariate analysis,
and of their genetic relationships, two natural groups could be seen for alatae and
apterae.
From the morphological point of view, there is a relative high degree of overlap
-
ping among species for many of their most significant biometric parameters currently
used to discriminate members of the genus. Nevertheless, specimens of apterae and
alatae viviparous morphs can be correctly identified.
Acknowledgments
We thank J Holman (Ceské Budejovice, Czech Republic) for the loan of
Anuraphis (A. farfarae, A. catonii, A. subterranea, A. pyrilaseri) collected in Eastern
European countries; the late GCh Shaposhnikov (St. Petersburgh, Russia) for the loan
of one apterous paratype of A. ferulae; RL Blackman (The Natural History Museum of
London, England) for the revision of the manuscript; FW Quednau (The Laurentian
Forestry Centre, Sainte Foy, Quebec, Canada) for advice and suggestions that improved
the manuscript; and P. Suma (Dipartimento di Scienze e Tecnologie Fitosanitarie,
Università of Catania, Italy) for producing the halftone figures.
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