The Aphids

SYSTEMATIC TREATMENT OF APHID GENERA

(in alphabetical order)

U

Ucrimyzus

Uhlmannia

Uichancoella

Uroleucon

Utamphorophora

Ucrimyzus Mier Durante & Pérez Hidalgo

Aphidinae: Macrosiphini

A genus for one species found on several genera of Compositae/Asteraceae in Central America, superficially similar to Hyperomyzus but with smaller prothoracic spiracles.

Ucrimyzus villalobosi Mier Durante & Pérez Hidalgo    Apterae are pale dull green, with yellowish antennae, legs and siphunculi; BL 1.6-2.5 mm. Alatae are without dorsal abdominal sclerotisation and have secondary rhinaria distributed ANT III 25-43, IV 5-19, V 0-2. Forming dense colonies on stems of plants in several genera of Compositae/Asteraceae (Bidens, Schkuhria. Senecio, Stevia)in Mexico and Costa Rica (Mier Durante et al. 2013). The life cycle is unknown.

Uhlmannia Börner

Aphidinae: Macrosiphini

A genus for one species found on several genera of Compositae/Asteraceae in Central America, superficially similar to Hyperomyzus but with smaller prothoracic spiracles.

Ucrimyzus villalobosi Mier Durante & Pérez Hidalgo    Apterae are pale dull green, with yellowish antennae, legs and siphunculi; BL 1.6-2.5 mm. Alatae are without dorsal abdominal sclerotisation and have secondary rhinaria distributed ANT III 25-43, IV 5-19, V 0-2. Forming dense colonies on stems of plants in several genera of Compositae/Asteraceae (Bidens, Schkuhria. Senecio, Stevia)in Mexico and Costa Rica (Mier Durante et al. 2013). The life cycle is unknown.

Uichancoella Calilung

Hormaphidinae: Nipponaphidini

One species in the Philippines and Indonesia on Lithocarpus, apparently related to Schizoneuraphis. Only apterae are known.

Uichancoella gabrieli Calilung   Apterae are blackish-brown profusely covered with white wax, leaving only centre of dorsum exposed; BL 1.2-1.6 mm. In a large colony on twigs of Lithocarpus sp. at high altitude in the Philippines (Calilung 1975) and also collected in Sulawesi (BMNH collection, leg. J.H. Martin). Other morphs and life cycle are unknown.

Uroleucon Mordvilko

Aphidinae: Macrosiphini

About 230 species with an often extensive distal band of polygonal reticulation on rather  elongate siphunculi, usually with 5 hairs on the first tarsal segments and often with dorsal abdominal hairs arising from pigmented spots (scleroites). Many species are dark bronzy red or almost black and live on the stems of Compositae/Asteraceae, but some live under the rosette leaves. At least 200 species live on Compositae/Asteraceae, 11 species live on Campanulaceae in the Old World, and 9 species are described from other plant families. The males of many species are green as are the females of some, particularly members of the subgenus Lambersius, South American species included, which live mostly on the tribe Astereae. The other nearctic species live mostly on Heliantheae (16 spp.) or Astereae (13 spp.), while palaearctic species (100 + ) more frequently colonize Cynareae (20 spp.) and Cichorieae (23 spp.). Only 7 species are described from Anthemideae, the hosts of most members of the related genus Macrosiphoniella. From about 1935 to 1975 the genus was commonly called Dactynotus. Accounts are available for western Europe (Hille Ris Lambers 1939a), UK and Ireland (Blackman 2010), Fennoscandia and Denmark (Heie 1995), European Russia (Shaposhnikov 1964), Siberia (Ivanovskaya 1977, Pashchenko 2000a, 2001), palaearctic Inula‑feeders (Holman 1981c), Iran (Rezwani 1991), Kazakhstan (Kadyrbekov 2003a), Japan (Miyazaki 1971), Korea (Lee et al. 2002c, Choi et al. 2012), China (Tao 1963), India (Chakrabarti & Medda 2004), South America (Carvalho et al. 1998; Nieto Nafría et al. 2007, 2019a; Mier Durante et al. 2020), Puerto Rico (Smith et al. 1963), Mexico (Nieto Nafría et al. 2011) and North America (Robinson 1985, 1986). Kanturski & Barjadze (2020) provided a key to Palaearctic Uroleucon species with three hairs on first tarsal segments. Moran et al. (1999) used molecular methods to study the phylogeny and evolution of Uroleucon in North America and Europe. The subgeneric classification needs revision; in particular, it seems likely that there are North American species misplaced in Uromelan, and South American species misplaced in Lambersius. Ortego et al. 2022 established the synonymy of Blanchardaphis with Lambersisus, or alternatively with Uroleucon, depending on the taxonomic criteria used to define the subgenus Lambersius.

Uroleucon achilleae (Koch)    Apterae are red or brownish red with rows of black dorsal spots, black siphunculi and yellow cauda (see influential points/Gallery); BL c.2.4-2.6 mm. On lower leaves of Achillea spp. in Europe, Iran, Kazakhstan and Kyrgyzstan (Kadyrbekov 2012a, 2017a), and introduced to USA (California, Oregon). There is also a trap record from Pakistan (Naumann-Etienne & Remaudière 1995). Monoecious holocyclic, with alate males. Sobhani (1970) studied its biology, life cycle and morphology of U. achilleae in Germany. 2n=12.

Uroleucon (Uromelan) acroptilidis Kadyrbekov, Renxin & Shao    Apterae are brownish; BL 2.9-3.4 mm. On stems of Rhaponticum (= Acroptilon)spp.in south-east Kazakhstan and Xinjiang-Uygur region of western China (Kadyrbekov et al. 2002, Kadyrbekov 2004b, 2005b), and also recorded from Uzbekistan (Kadyrbekov 2013c) and Iran (Barjadze et al. 2017b).

Uroleucon adenocaulonae (Essig)    Apterae are shining reddish, often tinged with olive-green, with dark siphunculi but antennae and legs mainly pale (see aphidtrek.org); BL 2.3-2.9 mm.  On flower stems of Adenocaulon bicolor in western USA (California, Oregon, Washington, Idaho). In Oregon it has also been found on Crepis sp. (Leonard 1974).

Uroleucon (Uromelan) adenophorae (Matsumura)  ( = Uroleucon (Uromelan) adenophoricola Holman)   (Fig.7c)   Apterae are shining dark brown to black; BL 3.7-4.2 mm. On petioles of terminal leaves of Adenophora spp. in Japan, South Korea (Choi et al. 2012 – but note that there are errors in the morphometric data tabulated by these authors), and Mongolia (Holman 1975, as U. adenophoricola).

Uroleucon adesmiae Mier Durante& Ortego     Apterae are dark green to brownish green; BL 2.4-3.2 mm. On stems of Adesmia sp(p). (but not A. boronioides) in Argentina (Mier Durante & Ortego 2008, Nieto Nafría et al. 2019a, Mier Durante et al. 2020).

Uroleucon (Uromelan) aeneum (Hille Ris Lambers)    Apterae are shiny metallic bronze-black (see influentialpoints.com/Gallery); BL 3.0-4.3 mm. On upper parts of stems of thistles (Carduus, Cirsium, Galactites, Onopordon, Silybum), often forming very large colonies. Also recorded from Arctium, Carthamus and Sonchus), but these are likely to be casual occurrences or misidentifications. In Europe, North Africa (Algeria: Peréz Hidalgo et al. 2012b), Turkey, Armenia, Iran, Central Asia, east and west Siberia, and introduced to South America (Argentina, Chile). Monoecious holocyclic, with oviparae and alate males appearing in September (Hille Ris Lambers 1939). Closely related to U. jaceae, and originally described as a subspecies.

Uroleucon alaskense Robinson    Colour of apterae in life is unrecorded, tibiae are probably uniformly dark; BL 2.5-3.1 mm. On Achillea sp. in Alaska.

Uroleucon (Lambersius) altaicum Szelegiewicz   Apterae in life pale green, with only tips of siphunculi dark: BL c.2.6 mm. On Cichorium intybus in Mongolia (Szelegiewicz 1982). Kanturski & Barjadze (2020) provided photomicrographs of this species, distinguished it from U. mulgedii with which it had been previously synonymised, and placed it in the subgenus Lambersius.

Uroleucon (Uromelan) amamianum (Takahashi)    Apterae are bright shiny red to reddish brown, with black antennae, siphunculi,  cauda and distal halves of femora; BL 1.8-2.5 mm. On Solidago and Aster in Japan (Moritsu 1983) and Korea (Lee et al. 2002c), where it was also found on Carpesium abrotanoides, Picris hieracioides and Patrinia scabiosaefolia (the latter probably not a true host). Kanturski & Lee (2020) described and illustrated the ovipara, collected in Korea in October on Solidago virgaurea.

Uroleucon ambiguum Pashchenko    Apterae are shining fuscous brown with brown siphunculi and pale cauda; BL c.3.5 mm. Alatae have only 7-9 secondary rhinaria on ANT III. At stem apices of Artemisia opulenta in east Siberia (Kamchatka; Pashchenko 2000a)

Uroleucon ambrosiae (Thomas)  Plate 28d  Apterae are red-brown to dark brown or dull red, with black siphunculi and a pale cauda; BL 2.5-3.5 mm. Mainly on Ambrosia and Iva in eastern and northern USA, where records from other plants may be misidentifications of other species (Moran 1985). However, populations in south-western USA, and Central and South America are far less specific, colonising numerous species of Compositae/Asteraceae (e.g. Achillea, Ambrosia, Aster, Cichorium, Coreopsis, Eupatorium, Lactuca, Rudbeckia, Senecio, Taraxacum, Xanthium), usually occurring on the flower-stems. This more polyphagous form (regarded in South America as a subspecies, U. ambrosiae ssp. lizerianumBlanchard), has fewer secondary rhinaria on the antennae of both apterae and alatae (Carvalho et al. 1998; Bernays et al. 2000). Bernays & Funk (1999, 2000) found many differences in host selection behaviour between these two forms. Monoecious holocyclic in temperate North America, with alate males; probably mainly or entirely anholocyclic in southern USA, and Central and South America. Records from Tunisia (Ben Halima-Kamel 2012), Algeria (Laamari et al. 2013) and Turkey (Görür et al. 2011b, 2012) require confirmation. U. aaroni Knowlton, described from alatae only, may be a synonym. 2n=12.

Uroleucon  amigoi Nieto Nafría, Ortego & Mier Durante   Apterae are dark brown with dark siphunculi and hyaline antennae, legs and cauda; BL 2.8-3.5 mm. On stems of Adenocaulon chilense, forming relatively compact groups. In southern Chile (Capitán Prat province). Monoecious holocyclic, producing oviparae in early January(Nieto Nafría et al. 2021a).

Uroleucon (Lambersius) anomalae (Hottes & Frison)    Apterae are pale green to yellow-green, siphunculi dark-tipped (see influential points/Gallery); BL c. 1.9-2.0 mm. At apices of flower stems of Symphyotrichum novae-angliae in eastern USA (Moran 1985). 2n=12.

Uroleucon (Uromelan) ariegense Nieto Nafría & Pérez Hidalgo   Colour of aptera in life is unknown, siphunculi and cauda are equally dark; BL c.2.7-3.0 mm (Nieto Nafría & Pérez Hidalgo 2013a). Apterae and alatae were collected in June on an unidentified species of Campanula in southern France (Pyrenean region). The life cycle is unknown.

Uroleucon arnesense Robinson    Apterae are dark green: BL 2.0-2.6 mm. Described from a small species of Solidago in Arnes, Manitoba. Specimens in the BMNH collection from Solidago sp(p) in Ontario (leg. J.P. Sijpkens) and Utah (leg. G.F. Knowlton) also agree closely with paratypes of arnesense. It is possible that the dwarf individuals included in the description of U. pieloui (Richards 1972) were this species.

Uroleucon asteriae Lee, Holman & Havelka    Colour of apterae in life is unrecorded; BL 2.7-3.6 mm. On Aster koraiensis and A. scaber in North Korea (Lee et al. 2002a).

Uroleucon asteromyzon Zhang, Chen, Zhong & Li, in Zhang    Apterae are black; BL c. 2.9 mm. On young shoot terminals of Aster tataricum in Gansu Province, China (Zhang 1999). The type specimens seen had most of the pigment removed in preparation, so certain assumptions had to be made in including this species in the key to aphids on Aster.

Uroleucon asterophagum (Nevsky)    Apterae are yellow-green, with dark antennae and pale siphunculi and cauda; BL 3.0-3.5 mm. On stalks and flowerheads of Aster sp.and Erigeron sp. in Central Asia, and subsequently found on Pseudolinosyris grimmii (Kadyrbekov 2002d). Some specimens collected from Aster himalaicus at Wisley, UK in 1940 (BMNH collection, leg. G. Fox-Wilson) also key to this species.

Uroleucon astronomus (Hille Ris Lambers)    Colour of apterae in life is apparently unrecorded; BL 3.1-4.3 mm. On flower stems of Eurybia macrophylla in eastern USA and Canada, where it is completely host-specific according to Moran (1985), with oviparae and alate males appearing on E. macrophylla in September in Quebec (original description). However, the paratype material includes specimens from Aster (= Symphyotrichum) novae-angliae, and this species has now been recorded from Mexico (Nieto Nafría et al. 2011), where it is apparently much more polyphagous, occurring on plants in several genera of Compositae/Asteraceae, especially those growing at high altitudes.

Uroleucon atripes (Gillette & Palmer)    Apterae are dark reddish brown to blackish red with black legs, antennae and siphunculi; BL 1.8-3.0 mm. On leaves and stems of Aster sp., deforming stems when abundant, and also recorded from Brickellia eupatoroides and Solidago missouriensis.In western USA and British Columbia. Oviparae and alate males occur in October (Palmer 1952).

Uroleucon australe Nieto Nafría & Mier Durante   Apterae are matte green due to dusting of whitish wax powder; BL 3,2-3.7 mm. On small stems of small plants of Adesmia boronioides in southern Chile (Aysén and Magallanes regions). Holocyclic with abbreviated life cycle; apterous males and oviparae were collected in mid-summer (Mier Durante et al. 2020). Alatae are unknown. Very similar to A. nahuelhuapense, apterae differing in colour in life and shape of siphunculi.

Uroleucon bereticum (Blanchard)    Apterae are shining green to yellowish green, with dusky antennae and dusky/dark siphunculi; BL 2.1-3.1 mm. On various Compositae/Asteraceae (Baccharis, Buva, Conyza, Erigeron, Tanacetum) in South America (Argentina, Brazil, Chile, Peru; Carvalho et al., 1998). Delfino & Starý (2004) redescribed this species, and also described an endemic parasitoid. Nieto Nafría et al. (2007) described an oviparous female and an alate male (collected in March).

Uroleucon bicolor (Nevsky)    Apterae are dark brown with yellowish sides to abdomen and a yellowish spinal stripe between siphunculi; siphunculi are black on basal and brown on distal thirds, with a greenish yellow section in middle; the cauda is yellow; BL 3.5-3.8 mm. In large colonies on stems of Onopordon spp., Senecio spp. and Lactuca viminea in western Siberia and Central Asia. Redescribed from some of Nevsky’s material by Holman (1991a), who suggested that the specimens from Onopordon sp(p). were probably a different species. Alatae are unknown.

Uroleucon (Lambersius) bielawskii (Szelegiewicz)    Colour of apterae in life is unknown; BL 2.9-3.4 mm. On wild Lactuca spp., and (possibly) Hieracium pilosella. In southern and eastern Europe, Turkey and Iran. Kanturski & Barjadze (2020) provided photomicrographs of apterae of this species and placed it in subgenus Lambersius.

Uroleucon bifrontis (Passerini)    Apterae are lustrous olive green with black head and siphunculi; BL 2-3 mm. On leaves and in inflorescences of Inula bifrons and Dittrichia viscosa in Italy and Rumania (Holman 1981a), and now also recorded from Algeria (Laamari et al. 2013).

Uroleucon (Uromelan?) bonitum (Hottes)    Apterae are bright red to brownish red with mainly dark appendages; BL 1.7-2.2 mm. On flower stems of Stephanomeria pauciflora in western USA. Monoecious holocyclic with apterous males in October (original description, as Macrosiphum).

Uroleucon boreale Robinson    Apterae are “dark greenish red, appearing overall as almost black”; BL 2.3-2.8 mm. On unidentified Solidago sp(p). in north-western North America (Yukon, North West Territories).

Uroleucon (Satula) brachychaetum (Olive)    Apterae are dark red to red-brown with blackish antennae and siphunculi, and reddish or yellowish legs; BL 1.4-2.0 mm. On Krigia spp. in North Carolina and Michigan, USA (Moran, 1985).

Uroleucon (Lambersius) bradburyi (Olive)    Apterae are green with mainly dusky-dark appendages, BL c. 2.2-2.6 mm. On Aster sp. in North Carolina, USA.  Alate males were found in October (Olive 1965). 2n=12. [A similar species, but with longer siphunculi and longer R IV+V, has been collected on Aster modestus in Oregon (BMNH collection, leg. D. Hille Ris Lambers.)]  2n=12.

Uroleucon brevirostre Holman    Apterae are blackish brown; BL 3.9-4.2 mm. On flower stems of Mulgedium (= Lagedium) sibiricum (Pashchenko, 2001). Mongolia and Siberia.

Uroleucon (Lambersius) breviscriptum (Palmer)    Apterae are medium green with mainly black appendages (see influential points/Gallery); BL 2.5-3.3 mm. On leaves and stems of  Symphyotrichum laeve (= Aster laevis) in western North America. Oviparae occur in early October (Palmer 1952, as Macrosiphum). U. macgillivrayae in eastern USA and Canada is very similar and a possible synonym.

Uroleucon brevisiphon Carvalho    Colour of apterae in life is unknown, BL 1.8-2.5 mm. Antennae, distal halves of femora, tarsi and siphunculi are dark, and the coxae are also rather dark compared with other South American Uroleucon. On Baccharis spp.in Chile (Carvalho et al. 1998), and there is also a record from a Grindelia sp. (Nieto Nafría et al. 2016b)

Uroleucon (Lambersius) brevitarsus (Robinson)    Apterae are green; BL 2.4-3.2 mm. On terminal parts of  stems of  an unidentified Solidago sp. in Manitoba, Canada. Aphids identified as this species are also recorded from either Solidago or Aster in California (A. Jensen, aphidtrek.org), and from an Erigeron sp. at 2000 m altitude in Mexico (Nieto Nafría et al. 2011).

Uroleucon budhium (H. Banerjee, A.K. Ghosh & Raychaudhuri) (= tenuirostre L.K. Ghosh; = acutirostre Bänziger)    Apterae are (according to original description of budhium)pale brown with dark brown head and siphunculi, antennae and legs mainly pale with dark apices (also dark sensoriated part of ANT III and distal 0.3 of femora), and pale cauda; BL 2.8-4.0 mm. Immatures are creamish white. Described from colonies on undersides of young leaves, flower buds and stems of Echinops cornigerus, and  from “bases and apical shoots” of Anaphalis sp., in Uttar Pradesh, India. Monoecious holocyclic in northern India; the ovipara and alate male were described by Chakrabarti & Medda (2004). Holman (1981c) doubted the association with Echinops, but as well as records from Inula spp. and Blumea sp. (Raychaudhuri, 1980), there is also a large collection from Saussurea albescens (BMNH collection, leg. N.D. Rishi), indicating that it may well colonise Compositae/Asteraceae outside the Inuleae group. However, a record from Ficus sp. (Raychaudhuri, 1980, p.234) should certainly be discounted as vagrants. In northern India (Uttar Pradesh, Meghalaya, West Bengal, Kashmir) and Thailand (Bänziger 1980, as U. acutirostre). [Paratypes of U. acutirostre Bänziger, described from northern Thailand, could not be distinguished from paratypes and other specimens of U. tenuirostre (“=  budhium”) in the BMNH collection, so if the synonymy of tenuirostre with budhium is correct, acutirostre should also be regarded as a synonym (Eastop & Blackman 2005). However, the colour in life of acutirostre is recorded as dark blackish red (Bänziger 1980). The original description of tenuirostre did not record the colour in life.]

Uroleucon bulgaricum Kanturski   Colour of apterae in life unknown: BL 1.9-2.7 mm. On Achillea coarctaca in Bulgaria (Kanturski 2021). Feeding position on plant and life cycle are unknown.

Uroleucon (Lambersius) cadens Moran  (Fig.49c)   Apterae are green, with antennae, legs and apical halves of  siphunculi black; BL 1.5-1.9 mm. On Solidago nemoralis in southern Michigan, falling readily when disturbed.

Uroleucon (Uromelan) calendulae (Nevsky)   The body of the aptera is brownish, appendages somewhat paler; BL c.3.5 mm. Described from flower stems and undersides of upper leaves of Calendula arvensis, an introduced plant, in southern Kazakhstan; the native host is Erigeron aurantiacus (Kadyrbekov 2017a).

Uroleucon (Lambersius) caligatum (Richards)    Apterae are green with mostly black antennae, legs, and siphunculi except at bases, and cauda pale with a dusky tip (see influential points/Gallery); BL 3.2-4.0 mm. On Solidago spp., common in north-eastern USA and Canada. Moran (1981) studied intraspecific variability in performance between three Solidago spp. Cappuccino (1987) compared its population dynamics with that of U. nigrotuberculatum feeding on the same host (Solidago altissima). Closely related to U. luteolum, which has a more southerly distribution in USA, and apparently differs only in the lack of spinal tubercles; further studies are needed, as the development of these tubercles is likely to be environmentally determined.

Uroleucon (Uromelan) campanulae (Kaltenbach)    Apterae are shiny reddish brown to black, with antennae, siphunculi and cauda black and legs bicoloured yellow and black (see influentialpoints.com/Gallery); BL 2.1-3.7 mm. On upper parts of stems and flowers of Campanulaceae (Campanula, Jasione) in Europe, western Siberia and south-west and central Asia. Oviparae and alate males occur in September-October.  Populations from different regions and hosts differ somewhat in morphology (e.g., see Heie 1995), and there is possibly a complex of species. Specimens from Jasione have longer siphunculi than those from Campanula (SIPH 0.23-0.28 × BL and 1.01-1.14 × cauda in apterae from Jasione; SIPH 0.14-0.21 × BL and 0.72-0.98 × cauda in apterae from Campanula), and R IV+V is also longer relative to HT II in Jasione-feeding aphids (Heie, 1995; VFE, unpublished data), suggesting that there may be two distinct species. Hille Ris Lambers (1939) observed that the form from Jasione usually had little success in colonising Campanula. Börner (1950) distinguished populations on Campanula in Austria as a subspecies, U. campanulae ssp. longius.

Uroleucon (Lambersius) canadense (Richards)    Colour of aptera in life was not observed; BL c.1.6-3.2 mm. On Solidago hispida in Canada (Ontario, Quebec). Closely related to and probably synonymous with U. gravicorne;the discriminants given by Richards do not differentiate this species from many specimens of  gravicorne in the BMNH collection.

Uroleucon (Lambersius) capitophoroides (Blanchard)    Apterae are dark pink to dark reddish, frequently with a narrow longitudinal greenish mark on abdominal tergites 2-5. BL 1.9-3,3 mm. Apterae have 5-30 secondary rhinaria on ANT III, more frequently 10-20. Monoecious holocyclic, with alate males, and forming dense colonies on flowering stems of several South American species of Senecio,in Argentina and Chile. Ortego et al 2022, revised the status and taxonomic position of this species previously known as Blanchardaphis capitophoroides and its synonym B. poikila.

Uroleucon carberriense Robinson    Apterae are green, with mainly pale appendages; BL 2.2-3.0 mm. On Chrysopsis (= Heterotheca) villosa in Canada (Manitoba and Alberta).

Uroleucon (Uromelan) carlinae (Börner)    Apterae are reddish brown to blackish brown, BL 2.4-3.0 mm. On Carlina caulescens in southern and central Europe (France, Germany, Poland, Hungary, Italy, Ukraine). Oviparae and alate males appear in early August in Germany (original description).

Uroleucon (Uromelan) carthami (Hille Ris Lambers)    Colour of apterae in life is dark brown to blackish brown (according to Papapanagiotou et al. (2012); BL 2.1-3.3 mm. On Carthamus spp. in southern and central Europe, Algeria (Laamari et al. 2013), Israel, Lebanon, Turkey, and eastward to Pakistan and India (Kashmir). Most records from C. tinctorius in India are probably referable to U. compositae. In the European literature there is also confusion with the very closely related U. jaceae and U. aeneum, which are normally on other Cynareae but can also sometimes occur on Carthamus (Nieto Nafria et al. 1986). Sexual morphs and life cycle unknown. 2n=12 (but 2n=14 is recorded by Khuda-Bukhsh & Kar 1990).

Uroleucon caspicum Rezwani & Lampel    Apterae are shiny dark brown with dark antennae, legs and siphunculi, and pale cauda; BL 3.2-4.7 mm. On Serratula quinquefolia, living on undersides of leaves and on shoots. In Iran and Russia (Caucasus). Monoecious holocyclic, with oviparae and alate males in October (Holman 1991).

Uroleucon (Uromelan) cephalonopli Takahashi    Apterae are medium-sized, broadly spindle-shaped, shiny dark brown-black, with black siphunculi and cauda, antennae and femora mainly black, tibiae mainly yellow; BL c. 3 mm. On upper parts of stems and upper leaves of Cirsium spp. in Japan, Korea, east Siberia and Taiwan (as Macrosiphum ambrosiae; Miyazaki 1971).  There are records of U. cephalonopli from various other genera in North Korea (Lee et al. 2002c), but this species seems to have a strong preference for Cirsium,and there is possible confusion with other very similar species such as U. gobonis. There is a strong possibility that this species is synonymous with U. cameronense (Takahashi), whichwas described from an unknowncomposite plant in Malaya.

Uroleucon chani Robinson    Apterae are brownish green, with antennae (except base of III), distal parts of femora and tibiae, and siphunculi, dark brown/black; BL 2.3-2.9 mm. On Grindelia nana (= hirsutula)in British Columbia, Canada. Oviparae and alate males occur in early October (original description). Aphids identified as this species have now been recorded from Gnaphalium sp. at high altitude (2600m) in Mexico (Nieto Nafría et al. 2011).

Uroleucon chilense (Essig)    Colour in life unrecorded, BL c. 3.5 mm. On Baccharis sp(p). in Coquimbo Province, Chile (Carvalho et al. 1998), and also recorded from Eupatorium candolleanum (Nieto Nafría et al. 2007).

Uroleucon chiliotrichi Nieto Nafría, Ortego & Mier Durante   Apterae are pale green with dark brown antennae and brownish green siphunculi and cauda; BL 2.2-3.0 mm. Other morphs are not known (Nieto Nafría et al. 2021a). On Chiliotrichum diffusum in southern Chile (Coyhaique province). Only known from the type locality.

Uroleucon chondrillae (Nevsky)    Apterae are dark brown with black antennae and siphunculi and yellowish cauda; BL 2.6-3.8 mm. On stems and flower stalks of  Chondrilla juncea in Europe, Russian Volga region and south-west and Central Asia; in Kazakhstan it is recorded also from several other Chondrilla spp. (Kadyrbekov & Aoitzhanova 2005, Kadyrbekov 2017a)). Facultatively holocyclic in the Mediterranean area, where Caresche et al. (1974) studied its biology and host specificity 2n=12.

Uroleucon chrysanthemi (Oestlund)    Apterae are dark brown, with black siphunculi and yellowish cauda; BL 2.2-3 mm. On Bidens spp. in North America. A record from Calendula in Algeria (Laamari et al. 2013) is unlikely and requires confirmation. Monoecious holocyclic, with alate males (Hottes & Frison 1931). It is very similar to U. ambrosiae, and records of U. chrysanthemi from genera other than Bidens may be referable to that species or others in the ambrosiae group.

Uroleucon chrysopsidicola (Olive)    Apterae are red-brown with black antennae and siphunculi, yellowish legs with black femoral apices and  tarsi, and a pale cauda; BL 1.8-2.5 mm. On Chrysopsis spp. , of which Ch. mariana seems to be the usual host, with single records also from Erigeron annuus and Solidago sp. (Olive 1963), which might be other species. In eastern USA (Florida, Georgia, North Carolina). Monoecious holocyclic, with alate males. 2n=12.

Uroleucon cicerbitae Holman    Apterae are reddish brown with contrastingly whitish cauda and black siphunculi; BL 3.2-4.7 mm. On Cicerbita cacaliifolia (= macrophylla), presumably on upper stems and leaves in spring, and later on inflorescences. In southern Russia (eastern shore of Black Sea). Monoecious holocyclic with oviparae and alate males in October  (Holman 1991).

Uroleucon cichorii (Koch)    Apterae are shining metallic brown with black antennae and siphunculi, black legs except for dusky/dark coxae, pale trochanters and basal halves of femora, and pale yellow cauda (see influentialpoints.com/Gallery); BL 2.7-4.7 mm. On upper parts of stems of Cichorium and related genera of Cichorieae (Crepis, Hieracium, Lactuca, Lapsana, Leontodon, etc.). In Europe, south-west and central Asia, Eritrea, Mongolia, Korea and east Siberia. Monoecious holocyclic, with alate males. A member of a group of closely related species in Europe. Hille Ris Lambers (1939) discussed variation in the group as a whole and erected three subspecies of cichorii based mainly on differences in host colonisation in the field; ssp. cichorii on Cichorium, ssp. leontodontis on Leontodon,and ssp. grossum on Crepis. Here we follow Heie (1995) in treating these as separate species. However, further work is needed on the stability of host associations and taxonomic relationships within this group, and the host specificity of Cichorium-feeding populations in particular is not clearly established, nor is it supported by consistent morphological differences. U. cichorii should therefore be regarded as potentially capable of living on other genera, and the keys have been constructed to take this into account. It is also likely that some records of U. cichorii are misidentified U. picridis and U. hypochoeridis (and vice versa).

Uroleucon ciefi (Olive)    Apterae are dark brown to red-brown with dark antennae, legs and siphunculi, and a pale cauda; BL 1.3-2.3 mm. Alatae have 12-24 rhinaria on ANT III. On Eupatorium spp. in eastern USA (Florida, North Carolina).

Uroleucon cirsicola (Holman)    Apterae are spindle-shaped, dark brassy brown, with mainly black antennae, legs mainly pale brown, siphunculi black on basal half  but brown distally, and cauda yellow; BL 3.3-4.9 mm. On upper parts of stems and undersides of upper leaves of Cirsium spp. A few specimens collected in the type locality on Arctium minus were possibly also this species (Holman 1962), and it is also recorded from Senecio jacobaea and Tragopogon orientalis (Ivanoskaya 1977). In Ukraine (Crimea), Turkey (BMNH collection, leg. N. Tuatay) and west Siberia.

Uroleucon cirsii (L.)    Apterae are bronze brown, with dark head and prothorax, black dorsal abdominal spots, black siphunculi and a yellow cauda (see influentialpoints.com/Gallery); BL 3.2-5.2 mm. On Cirsium spp., forming colonies on upper parts of stems and upper leaves. Ivanoskaya (1977) recorded it from Tragopogon orientalis, although most records from other plant genera are likely to be due to misidentification.Very common throughout Europe and eastward to Iran and west Siberia, and introduced to North America. Monoecious holocyclic, with oviparae and alate males produced in September-October. 2n=10.

Uroleucon (Lambersius) coloradense Robinson    Colour of apterae in life is unknown; BL 2.6-2.8 mm. On Eriogonum flavum (Polygonaceae) in Colorado, USA. The host plant is unusual for the genus and needs to be verified by further collections.

Uroleucon (Uromelan) compositae (Theobald)  (= Dactynotus orientalis Kulkarni; Chakrabarti & Medda 2004)  Plate 28f   Apterae are shining very dark red to almost black, with black siphunculi and cauda (see aphids of Karnataka website); BL 1.9-4.1 mm. On flower-stems, and in low numbers along the mid-ribs of the leaves, of a wide range of Compositae/Asteraceae in tropical and subtropical climates, particularly plants growing in moist or shady situations at the end of the dry season. It is a pest of Carthamus tinctoria (safflower) in India (Blackman & Eastop 2000), and is common on herbaceous Vernonia after the rains in Africa (Eastop 1958). Sometimes it is found on non-composite plants such as Malva and Morus. Widely distributed in Africa and on the Indian subcontinent, and also recorded from Sicily, Turkey, Iran, Réunion, Mauritius, Taiwan and South America (Brazil, Surinam). It is not known to occur in south-east Asia; in Java its role as a general composite feeder seems to be taken over by the closely related U. vernoniae.Apparently it is anholocyclic everywhere, but it is difficult to distinguish from the east Asian species U. gobonis, and it could even possibly be an anholocyclic form of that species. Early African and Indian records of U. jaceae and U. solidaginis on Carthamus should probably all be referred to U. compositae. In southern Africa (Malawi, Tanzania, Zimbabwe) a form that feeds specifically on Vernonia is regarded as a subspecies, U. compositae ssp. evansi (Eastop). 2n=12.

Uroleucon (Lambersius) crepusisiphon (Olive)    Apterae are deep green with blackish antennae, legs and siphunculi, and a dusky-dark cauda; BL 1.7-2.3 mm. On Aster, Eurybia and Symphyotrichum spp. in eastern USA (Michigan, New York, North Carolina). This aphid falls readily from the plant when disturbed (Moran, 1985). Records of U. ambrosiae on these plant generain northern North America are probably this species.

Uroleucon dalmaticum Holman    Apterae are reddish to blackish brown with pale tibiae and whitish or yellowish cauda; BL 1.9-2.9 mm. On undersides of leaves and sometimes in inflorescences of Inula verbascifolia in Croatia(Dalmatia).

Uroleucon debile (Takahashi)    Apterae are dirty green, somewhat pinkish, with black siphunculi and dusky green cauda; BL c. 2.6 mm. On stems of Lactuca (= Ixeris) debilis in Taiwan.

Uroleucon deltense Robinson    Apterae are dark red, with dark brown siphunculi and pale cauda; BL 2.7-3.3 mm. On Lactuca pulchella in Manitoba, Canada. Alatae are unknown.

Uroleucon doellingeriae Pashchenko    Apterae are brown; BL c. 4 mm. On Aster (Doellingeria) scaber in east Siberia (Pashchenko 2000a).

Uroleucon (Uromelan) doronici (Börner)    Apterae are green, shining, with dark apices to segments of  antennae and legs, and black siphunculi and cauda; BL 2.6-5.0 mm. On flower stems and undersides of young leaves of Doronicum spp. in northern and central Europe. Monoecious holocyclic, with oviparae and alate males in September-October in northern Germany (Thieme & Gottschalk 1989).

Uroleucon dubium (Holman)    Apterae are green to brownish green, BL 2.3-3.25 mm. In dense colonies on terminal parts of shoots of  Artemisia sacrorumin Mongolia, and recently recorded from an Artemisia sp. in western Siberia (Altai Republic; Stekolshchikov & Novgorodova 2013).  Presumably monoecious holocyclic; oviparae were collected in July, along with apterous and alate virginoparae, and apterous virginoparae were also found in August (Holman 1975).  Collections at two separate localities suggest that A. sacrorum is the true host, although an unusual one for this group.

Uroleucon (Uromelan) dzhungaricum Kadyrbekov   Apterae are shining brown with distal segments of antennae, joints of legs and entire tarsi, siphunculi and cauda black; BL 4.8-5.8 mm. On Senecio spp. in Kazakhstan (Kadyrbekov 2016a, 2017a). Very similar to and possibly synonymous with the Korean species U. seneciocola.

Uroleucon (Uromelan) echinatum (Kulkarni)    Colour of apterae in life is not recorded; BL 2.3-3.0 mm. On Echinops echinata (recorded as “Echinus echinatus”) in Maharashtra, India (Kulkarni 1980). [The original description did not discriminate this species from U. compositae, butS. Chakrabarti (pers. comm.) has examined type material of this species and found that antennal and dorsal abdominal hairs are shorter than originally reported, and too short for compositae (hairs on ANT III are 26-31 μm, c.0.5 × BD III; hairs on ABD TERG 3 are 47-59 μm, 0.9-1.0 × BD III – note that these measurements were by mistake reversed in the key by Chakrabarti & Medda 2004).]

Uroleucon elbursicum Lampel & Rezwani    Apterae are blackish brown, with pale cauda; BL 2.7-3.3 mm. In inflorescences of Inula thapsoides in Iran. Lampel & Rezwani (original description) tabulated the differences from related species.

Uroleucon elephantopicola Robinson    Apterae are dark, probably brown, but their colour is unrecorded, BL 1.5-1.8 mm. On undersides of leaves, flower stalks and inflorescences of  Elephantopus in USA, described from specimens collected on E. carolianus in Illinois (Robinson 1985), and subsequently found in Florida (Halbert et al. 2000).

Uroleucon (Uromelan) ensifoliae Holman    Apterae are dark reddish brown with black antennae, siphunculi and cauda; BL 3.0-4.6 mm. On undersides of leaves of Inula spp. in eastern Europe (Poland, Slovakia, Czech Republic, Crimea).

Uroleucon (Uromelan?) eoessigi (Knowlton)    Apterae are reddish-brown to maroon, with appendages mainly black (see aphidtrek.org); BL 2.2-3.3 mm. On stems and leaves of  Alcea rosea in western and northern USA, and western Canada, and also found on Iliamna rivularis and anunidentified species of another Nearctic mallow genus Sidalcea (see aphidtrek.org).  Monoecious holocyclic with apterous males.  Knowlton (original description, as Macrosiphum) gave a full account of the life cycle in Utah.

Uroleucon epilobii (Pergande)    Apterae are brown or green with mainly dark appendages (original description); BL c. 2.7 mm. On leaves and stems of Epilobium angustifolium and Epilobium sp. in North America (Kansas, Colorado).  The Colorado aphids (Gillette & Palmer, 1934) differ in colour from original description. The generic position is possibly incorrect.  

Uroleucon (Lambersius) erigeronense (Thomas)  Plate 28g  Apterae are yellowish green with darker spinal stripe or uniformly pale green, siphunculi dark distally with pale bases (see influentialpoints.com/Gallery); BL 2.3-2.8 mm. Typically it occurs on upper parts of stems of  Conyza and Erigeron spp. However, Jensen et al. (2010) have shown that in western USA it can also colonise several other composite genera (with U. escalantii, described from Chrysothamnus nauseosus, as a new synonym), and in South America it has been found on plants in many more composite genera including Aster, Baccharis, Haplopappus, Heterotheca, Grindelia and Lactuca. Of North American origin, introduced to Central and South America, and to Europe, from where it has spread eastward to west Siberia, Iran, Kazakhstan and Pakistan, and south to Algeria (Laamari et al. 2013). It has also been found in Korea on Erigeron spp. and Chrysanthemum lineare (Lee et al. 2002c), and is now reported to be widely distributed in Australia (Brumley & Watson 2017). In the western palaearctic it has only been recorded from Conyza canadensis. Monoecious holocyclic, with oviparae and alate males in September-October in north temperate regions. 2n=12.

Uroleucon essigi Carvalho    Colour of apterae in life is unknown, BL 2.0-2.4 mm. On Baccharis spp. in Coquimbo Province, Chile (Carvalho et al. 1998). It is also recorded from Hysterionica jasionoides (Nieto Nafría et al. 2007).

Uroleucon eumadiae Delfino & Gonzales    Apterae are shining green; BL 2.0-3.1 mm. On flowerheads and stems of Madia spp. in Chile (Delfino & Gonzales 2005).

Uroleucon eupatoricolens (Patch)    Apterae are dull reddish, with black siphunculi and a pale cauda; BL 3.1-4.9 mm. On stems of Eupatorium (and Eutrochium) spp. in north-eastern USA and eastern Canada. Alate males were collected in late August in New Brunswick (MacGillivray 1968).  There are more recent records indicating that this species may have a wider distribution and host range, possibly in relation to montane habitats. Favret et al. (2010) recorded it from Hieracium, Senecio and Verbesina in the Great Smoky Mountains National Park, and an aphid collected on Sonchus sp. at 1800 m altitude in Baja California has also been identified as this species (Nieto Nafría et al. 2011).

Uroleucon (Uromelan) eupatorifoliae (Tissot)    Apterae have head and thorax dark green and abdomen red-brown, antennae and legs pale basally but distally mainly dark brown to black, siphunculi black and cauda dark green (see influential points/Gallery); BL c. 2.2-2.4 mm. Described from  Eupatorium incarnatum (= Fleischmannia incarnata)in Florida, and also recorded from more northerly states in eastern USA. Sexual morphs were collected in December in Florida. [Siphonophora eupatorii Williams, described from Eupatorium ageratoides (= Ageratina altissima) in Nebraska and currently placed in Macrosiphum, could not be distinguished from Uroleucon eupatorifoliae on the basis of the original description, and is possibly an earlier name for Tissot’s species.]

Uroleucon fagopyri Chowdhuri, R.C. Basu, Chakrabarti & Raychaudhuri  (Fig.28b)   Apterae are greenish brown; BL 3.2-3.7 mm. On Fagopyrum cymosum in north-west India. Presumably it is monoecious holocyclic; the ovipara (from an unidentified host plant, but collected with apterous viviparae) was described by Raychaudhuri, L.K. Ghosh & Das (1980).

Uroleucon floricola Robinson    Apterae are dark brown with dark siphunculi and cauda; BL 1.6-2.4 mm. In flowers of Aster sp. in Pennsylvania, USA.

Uroleucon formosanum (Takahashi)    Apterae are shining red-brown with a broad black “saddle”, black siphunculi and a pale yellow cauda (see Kanturski & Lee 2020); BL 2.3-3.4 mm. On stems, and on undersides of leaves along midribs, of Lactuca and related genera in east Asia (Japan, Korea, China, Taiwan, east Siberia), and more recently reported from Youngia japonica in the Mariana Islands (R.H. Miller et al. 2003). Pike et al. (2005) provided a detailed redescription and morphological comparison with the North American Lactuca-feeding species U. sonchellum. Kanturski & Lee (2020) described and illustrated oviparae and alate males collected in October in Korea. Apterae from Y. japonica in India (West Bengal) differ in having fewer secondary rhinaria and fewer caudal hairs and were described as U. formosanum ssp. crepidis A.K. Ghosh, M.R. Ghosh & Raychaudhuri. 2n=12.

Uroleucon fuchuense (Shinji)    Apterae are shining salmon red to reddish brown, with cauda concolorous with body (see Kanturski & Lee 2020, fig.3d), BL  2.5-3.8 mm. On flower stems of Aster spp., especially A. scaber,and also recorded from Pterocypsela raddeana (Moritsu 1958), Cacalia hastata and Saussurea grandifolia (Lee et al. 2002c). In Japan, Korea and east Siberia.  Kanturski & Lee (2020) described and illustrated the ovipara and alate male, collected on A. scaber in Korea in October.

Uroleucon fuscaudatum Chakrabarti & Raychaudhuri    Colour of apterae in life is unrecorded; BL 2.0-2.6 mm. On Inula rubricaulis in northern India (Uttar Pradesh). 2n=12.

Uroleucon garnicai Delfino    Apterae are brown with black siphunculi and a pale/dusky cauda; BL 1.9-2.7 mm. Alatae have 8-14 rhinaria on ANT III. On Eupatorium buniifolium in Argentina (Córdoba, Tucumán), and also recorded from E. patens.

Uroleucon (Uromelan) giganteum (Matsumura)    Apterae are large, shining very dark red, with appendages all black except for bases of femora; BL 4.0-4.3 mm. On Cirsium spp., and also recorded from Dendranthema, Saussurea (= Hemisteptia) lyrata, Serratula wolffii (= coronata) and Synurus spp. In Japan, Korea, east Siberia and Kazakhstan (Kadyrbekov 2017a).

Uroleucon gigantiphagum Moran    Apterae are red-brown to dark brown, antennae and siphunculi black, legs banded with black, cauda pale; BL 2.9-3.4 mm. On Solidago gigantea and S. juncea in Michigan, USA. Paratypes in the BMNH collection are not clearly separable from specimens identified as U. nigrotuberculatum or U. pieloui.

Uroleucon gnaphalii Mier Durante & Nieto Nafría    Colour of apterae in life is unrecorded; BL 2.2-3.9 mm. On Gnaphalium (= Pseudognaphalium) inornatum and Gnaphalium sp. in Mexico (Nieto Nafría et al. 2011). Apterae and alatae were collected in September-October.

Uroleucon (Uromelan) gobonis (Matsumura)    Apterae are shining greenish black to black, with black antennae, siphunculi and cauda, and legs with femora mainly black and tibiae mainly yellow-brown; BL 2.3-3.6 mm. On upper stems and leaves of various Cynareae, especially Arctium, Atractylodes, Carthamus and Saussurea. It can also occur on Cirsium (BMNH collection), but manyrecords of U. gobonis from this host in east Asia are probably referable to U. cephalonopli. In Japan, Korea, Taiwan, China, Mongolia, east and west Siberia (Altai region) and Kazakhstan. Monoecious holocyclic with alate males in Japan, but anholocyclic overwintering also occurs, and reproduction in Taiwan is exclusively parthenogenetic (Takahashi 1923). U. compositae is possibly an anholocyclic form of U. gobonis, as there are no clear morphological discriminants between these two species. 2n=12.

Uroleucon gochnatiae Delfino    Apterae are dark green with black head, antennae, legs, siphunculi and cauda; BL 1.9-2.5 mm. On Gochnatia glutinosa in Argentina (Tucumán), colonising terminal leaves. Nieto Nafría et al. (2007) also recorded it from Hyaloceris cinerea, and described oviparae and alate males found in April.

Uroleucon (Lambersius) gravicorne (Patch)  (Fig.49b)   Apterae are green with dark brown appendages, except for pale cauda and bases of siphunculi; BL 1.6-2.4 mm. On various Compositae/Asteraceae, especially species of Erigeron and Solidago, but also recorded from Aster spp., Chrysanthemum leucanthemum (Olive 1963) and Sonchus sp. (Nieto Nafría et al. 2011). In Michigan, it is common on Erigeron spp. in June-August, but Solidago spp. are the more common hosts in spring and autumn (Moran 1983). Widely distributed in North America, and also now in Central (Panama, Costa Rica; Villalobos Muller et al. 2010) and South America (Carvalho et al. 1998). U. canadense Richards is probably a synonym. Oviparae and alate males occur in October-November. 2n=12.

Uroleucon gredinae Pashchenko    Apterae are black with dark brown appendages, basal parts of antennae and femora paler; BL c. 3.1 mm. On upper parts of Campanula sp. in east Siberia (Pashchenko 2000a). It is not clear how this species differs from U. cichorii.

Uroleucon (Lambersius) grindeliae Barjadze & Kanturski  Colour in life unknown; BL of apterae 1.8-2.6 mm. On Grindelia hirsutula in Oregon, USA (Barjadze & Kanturski 2020). Biology and life cycle unknown.

Uroleucon grossum (Hille Ris Lambers)    Apterae are shining metallic brown with dark antennae and siphunculi, and yellow cauda (see influentialpoints.com/Gallery); BL 2.8-4.9 mm. On upper parts of stems of Crepis spp. in Europe, western Siberia, Central Asia and Mongolia. Monoecious holocyclic, with oviparae and alate males in Europe at end of August (Heie 1995). Further work is needed to confirm its distinction from U. cichorii (q.v.).  2n=12.

Uroleucon hasanicum Pashchenko    Apterae are shining black with pale middle sections of tibiae and pale cauda; BL c. 4.4 mm. Alatae are undescribed.  On shoot apices and flower shoots of Pterocypsela indica (Pashchenko 2000a) in east Siberia (Primorskii).

Uroleucon (Uromelan) helenae (Hille Ris Lambers)    Apterae are shiny black with a bronze tinge, BL 1.7-2.6 mm. In large, rather diffuse colonies on stems of Carlina vulgaris (Stroyan 1957b) in Europe (Croatia, Czech Republic, France, Italy, Switzerland, UK). It is also reported to occur in Kazakhstan, but on Xeranthemum longepapposum (Kadyrbekov 2017a). Oviparae have been reared in culture in England in late September (BMNH collection, leg. H.L.G. Stroyan) but males are unknown.

Uroleucon (Uromelan) helianthicola (Olive)    Apterae are reddish brown with black appendages, including siphunculi and cauda; BL 2.4-3.5 mm. On Helianthus spp., especially wild perennial  species (Rogers et al.1978). Widespread in USA. Sexual morphs are unknown.  2n=12.

Uroleucon (Uromelan) helichrysi Nieto Nafría & Pérez Hidalgo    Colour of apterae in life is unknown, siphunculi and cauda both dark; BL c.3.3-4.1 mm. On Helichrysum sp. in Iran (Nieto Nafría & Pérez Hidalgo 2014). The description includes alatae.

Uroleucon heterothecae Pérez Hidalgo & Nieto Nafría    Colour of apterae in life is unrecorded; BL 2.2-3.2 mm. Apterae and alatae were collected from Heterotheca inuloides in October at high altitude (1750-2300m) in Mexico (Nieto Nafría et al. 2011).

Uroleucon hieracicola (Hille Ris Lambers)    Apterae are bronzy brown, with black siphunculi and pale cauda; BL 2.9-3.5 mm. On flowerheads of Hieracium canadense in Canada (British Columbia, Manitoba, Ontario, Quebec), and also recently found in Maine, USA (A. Jensen, aphidtrek.org). Closely related to, and with little to distinguish it from, U. ambrosiae.

Uroleucon (Uromelan) hieracioides (Bozhko)    Apterae are greenish brown, with black antennae, siphunculi and cauda ; BL c. 2.9 mm. On Hieracium sp. in Ukraine.

Uroleucon hymenocephali Rezwani & Lampel    Apterae are matt greyish brown, with dark brown to blackish tibiae and siphunculi; BL 2.6-3.2 mm. On Hymenocephalus rigidus in Iran.

Uroleucon hypochoeridis (Hille Ris Lambers)    Apterae are brown or reddish brown with black antennae and siphunculi and a pale cauda (see influentialpoints.com/Gallery); BL 2.8-4.4 mm. On upper parts of flowerstalks of Hypochaeris spp., and occasionally also on Crepis, Leontodon and Taraxacum. Widely distributed in Europe, and in Turkey (Remaudière et al. 2006) and Iran (Rezwani 1991), although some of the records may be confused with other similar species, especially U. cichorii. Monoecious holocyclic, with oviparae and alate males in late August-September. Hartbauer (2010) studied its defensive behaviour against natural enemies. 2n=12.

Uroleucon hyssopii Narzikulov & Daniarova    Apterae are brown; BL c. 3.2 mm. On stems and leaves of  Hyssopus seravschanicus (Lamiaceae) in Tajikistan. Monoecious holocyclic with oviparae and alate males in late October (original description).

Uroleucon (Lambersius) idahoensis (Miller)    Apterae are pale green to creamy white, with brown antennae, legs and siphunculi and a green cauda; BL c. 2.3 mm. On leaves and flower stems of Anaphalis spp. (margaritacea, subalpina) in North America. Recorded from Idaho, Colorado (Palmer 1952), New York (Leonard 1963) and New Brunswick (BMNH collection, leg. M.E. MacGillivray). Life cycle and sexual morphs are not recorded.

Uroleucon (Uromelan) illini (Hottes & Frison)    Apterae are lemon yellow, bright green or red, with dark tips to antennae, femora and tibiae, and dark siphunculi and cauda; BL c. 2.7 mm. On both wild and cultivated Helianthus spp. in eastern USA and Canada. Oviparae and alate males are produced in October (Hottes & Frison 1931). 2n=12.

Uroleucon impatiensicolens (Patch)    Apterae are reddish brown to bronze-brown with black siphunculi and pale cauda (see influential points/Gallery); BL 2.5-3.6 mm. On Impatiens spp. in north-eastern North America. Apterae and alatae were re-described by MacGillivray (1968).

Uroleucon (Belochilum) inulae (Ferrari)    Apterae are grass green, with dusky appendages; BL 2.7-3.5 mm. On flower stems and undersides of leaves of Dittrichia (= Inula) viscosa, and possibly related species, in southern Europe and throughout the Mediterranean region, from Portugal to Lebanon, Jordan and Palestine (Hodjat & Ahmadian 1988). The life cycle has apparently not been studied. 2n=12*.

Uroleucon inulicola (Hille Ris Lambers)    Apterae are brown, reddish black or dark bronze, with entirely black antennae, black siphunculi and a rather dusky cauda; BL 2.4-4.7 mm. On Inula spp. in Europe, western Siberia and Central Asia. A record from Senecio in Algeria (Laamari et al. 2013) is unlikely and requires additional confirmation.Holman (1981c) compared several samples of this species and described regional differences, the range of variation encompassing specimens from Sweden that had been described as a subspecies (hirticola Ossiannilsson). Oviparae were described from Sweden (Ossiannilsson 1959, as ssp. hirticola).

Uroleucon iranicum Holman    Apterae are “presumably dark brown or reddish brown”, with blackish siphunculi and a pale cauda; BL 2.3-3.1 mm. On Gundelia tournefortii in Iran, colonising the apical part of the stem. First tarsal segments are unusual for a Uroleucon in having only 3 hairs.

Uroleucon ivae Robinson    Apterae are dark green to brown, appearing almost black; BL 3.0-3.8 mm. On Iva xanthifolia in North America (Manitoba, Colorado, Utah, Idaho, Oregon). Oviparae and alate males occur in September in Manitoba (original description). Some records of U. ambrosiae on Iva may be this species, although U. ambrosiae does also colonise Iva.

Uroleucon (Uromelan) jaceae (L.)    Apterae are dark reddish brown or blackish brown, often shiny, with black antennae, siphunculi and cauda (see influentialpoints.com/Gallery); BL 2.8-4.7 mm. On upper parts of stems of Centaurea spp. in Europe, west Siberia, Middle East, Central Asia and Pakistan, with occasional records from other composite genera, and sometimes also on certain Boraginaceae. Monoecious holocyclic with oviparae and alate males in late September in north-west Europe. Apterae of U. jaceae on Centaurea nigra have secondary rhinaria usually confined to the basal half of ANT III, whereas on other Centaurea spp., and on Carlina, Carthamus and Cynara spp. in southern Europe, secondary rhinaria may extend to 60-70% of the length of the segment, possibly because apterae tend to be more alatiform on less favourable hosts . The name may however being applied to a group of closely-related species with more specific host associations. Börner (1950) described populations on Centaurea scabiosa as a subspecies, U. jaceae ssp. henrichi, but his discriminants do not hold for all populations (e.g. see Heie 1995).Hille Ris Lambers (1939a) observed that specimens from Centaurea rhenana (= C. stoebe) in Italy differed from those from C. jacea in the same locality in having thicker siphunculi with more reticulation, and called this form U. jaceae var. reticulatum. Remaudière & Remaudière (1997) gave this form subspecies status; Kadyrbekov & Aoitzhanova (2005) have subsequently reported it from C. virgata ssp. squarrosa in Kazakhstan, and Kadyrbekov (2014g) from C. scabiosa in Russia (Altai). Most records of U. jaceae from Carduus and Cirsium may be referable to U. aeneum.

Uroleucon jaceicola (Hille Ris Lambers)    Apterae are dark bronze-brown, with black antennae and siphunculi, mainly yellow legs and a yellow cauda (see influentialpoints.com/Gallery); BL 2.9-3.3 mm.  On stems of Centaurea spp., especially nigra, jacea) found mainly low on stem in spring and later on upper parts. In Europe, west Siberia and Central Asia. An Italian population on C. ?paniculata with fewer caudal hairs and fewer secondary rhinaria was described as a subspecies, U. jaceicola ssp. pasqualei Hille Ris Lambers & Stroyan. Another subspecies, U. jaceicola ssp. kirgisica (Narzikulov & Umarov 1969), described from Kirghizia on C. squarrosa, has relatively much shorter siphunculi and longer cauda, and is perhaps a different species. Oviparae and small dark apterous males of U. jaceicola s.str. were found on petioles of radical leaves in October (Hille Ris Lambers 1939a and J.H. Martin, pers. comm). 2n=12.

Uroleucon kamtshaticum Pashchenko    Apterae are shining brown; BL c. 3.9 mm. On upper parts of Aster (two different unidentified spp., possibly the normal hosts?), with single apterae collected from Achillea sp. and Picris kamtshatica (Pashchenko 2000a). In east Siberia (Kamchatka).

Uroleucon kashmiricum (Verma)    Apterae are shiny dark reddish brown to black, with siphunculi black and cauda and middle parts of tibiae yellowish brown; BL 1.5-2.8 mm. On stems and young terminal leaves of Campanula spp. in Tajikistan, Afghanistan and northern India. Records from Compositae/Asteraceae need further confirmation, as does a record from Turkey (Akyürek et al. 2011). Monoecious holocyclic on C. incanescens in Tajikistan (Holman 1974a, as U. narzykulovi), but sexual morphs are undescribed.

Uroleucon katonkae (Hottes)    Apterae are green with dusky to dark brown appendages, cauda and bases of siphunculi dusky yellow; BL c.2.7-2.9 mm. On leaves of  Symphyotrichum laeve (= Aster laevis) and Brickiella grandiflora in western USA. Oviparae and apterous males were collected in October.

Uroleucon kikioense (Shinji)    Apterae are shiny reddish black, with black siphunculi, legs bicoloured yellow and black, and a yellow cauda;  BL c. 2.2 mm. On flower stems and undersides of leaves of Campanulaceae (Adenophora, Campanula, Platycodon) in Japan (Moritsu 1983), Korea (Lee et al. 2002a) and east Siberia. A record from Turkey (Görür et al. 2011b, 2012) requires additional confirmation. Pashchenko (2001) provided a redescription. Monoecious holocyclic on Platycodon grandiflorum in Korea (Lee et al. 2002a).

Uroleucon kumaoni Banerjee, A.K. Ghosh & Raychaudhuri    Apterae are blackish; BL c.4.1 mm. Immatures are greyish white. On undersides of young leaves, apical shoots and basal parts of flower stems of  Cirsium argyracanthum (= Cnicus argyracanthus) and Tricholepis furcata in north-west India). It was apparently visited by a red ant (Banerjee et al. 1969). This species is also recorded from Saussurea albescens and Tricholepis stewartii in Pakistan (Naumann-Etienne & Remaudière 1995). Apart from differences of pigmentation and chaetotaxy of the cauda, the description agrees closely with U. (Uromelan) gobonis.

Uroleucon (Uromelan) lactucicola (Strand)    Apterae are shining very dark reddish brown to black except for pale ANT III and basal part of  IV, pale basal halves of femora and middle parts of tibiae; BL 2.3-3.2 mm. Immatures are red. On stems of Solidago spp, and also recorded from certain other Compositae/Asteraceae (Aster maackii, Cirsium maackii, Ixeridium dentatum, Ixeris chinensis, Lactuca oldhami). In east Asia (China, Korea, Japan, Taiwan, east Siberia). Oviparae and alate males believed to belong to this species were found in November in China (Tseng & Tao 1938).

Uroleucon lebanonense Kanturski & Barjadze   Colour of apterae in life is unknown; BL 2.4-3.0 mm. On Tragopogon coloratus in Lebanon. Apterae and alatae were described by Kanturski & Barjadze (2020), who noted that the host plant species was possibly misidentified, as it has not been previously recorded from Lebanon. Other morphs and life cycle are unknown.

Uroleucon lanceolatum (Patch)    Apterae (usually alatiform) are dark red to reddish brown with black antennae and siphunculi, legs mainly dark except for femoral bases, and a pale cauda; BL 2.2-3.5 mm. Alatae usually have forewings with media only once-branched. On Solidago spp., usually graminifolia (=lanceolata), in north-eastern USA and Canada. An alate male was collected in late August. MacGillivray (1968) redescribed this species.

Uroleucon leonardi (Olive)    Apterae are dark brown to almost black, with black siphunculi and pale yellow cauda: BL 2.4-3.6 mm. On Rudbeckia spp., especially the hairier ones (Moran 1985), in central and north-eastern USA and eastern Canada. The life cycle is not recorded. This species is very similar to U. ambrosiae, and early North American records of that speciesfrom Rudbeckia are probably mostly leonardi.

Uroleucon leontodontis (Hille Ris Lambers)    Apterae are shining brown with black antennae and siphunculi and yellow cauda (see influentialpoints.com/Gallery); BL 3.2-4.3 mm. On Leontodon spp. in Europe. The alata was described and illustrated by Depa & Mróz (2012a). Described as a subspecies of U. cichorii, this aphid has a long narrow R IV+V similar to that of U. picridis, and specimens in the BMNH collection are not clearly distinguishable from the latter species, except by the less frequently developed marginal abdominal tubercles, which could be related to the host on which the aphids develop. Experimental work is needed to confirm its distinctness from U. picridis.

Uroleucon leontopodiicola Lee, Holman & Havelka    Colour of apterae in life is unrecorded; BL 2.9-3.6 mm. On Leontopodium coreanum in Korea (Lee et al. 2002a).

Uroleucon (Lambersius) longirostre (Gillette & Palmer)    Apterae are pale green with tips of antennae, legs and siphunculi dusky, and cauda pale to dusky; BL 2.0-3.0 mm. On Cirsium spp. (engelmannii, undulatum)in western USA (Colorado, Oregon, Washington).

Uroleucon longisetosum Chakrabarti & Verma    Apterae are dark brown; BL 2.7-2.8 mm. On growing shoots of Lactuca, Cicerbita and Prenanthes spp. in north India (Himachal Pradesh, Uttar Pradesh), and also recorded from Lactuca brunoniana in Pakistan (Naumann-Etienne & Remaudière 1995). The alata was described by Chakrabarti & Medda (2004). Monoecious holocyclic, with oviparae and apterous males in October (original description). 2n=10.

Uroleucon (Lambersius) luteolum (Williams)  (= tissoti Boudreaux)    Apterae are shining green or yellow with black antennae and legs, siphunculi black with pale basal fifth, and a pale cauda; BL  2.3-3.5 mm. Common on Solidago spp. in eastern and south-eastern USA, west to Kansas and Nebraska; uncommon further north in Michigan, Ontario and Quebec (Robinson 1986). Also recorded from Aster, Conyza and Erigeron, but some of the records may be misidentified other species (erigeronense, gravicorne, macgillivrayae). Oviparae and alate males occur on Solidago in September-October (Olive 1963). 2n=12.

Uroleucon (Lambersius) macgillivrayae (Olive)    Apterae are shining deep green with black appendages; BL 1.7-3.0 mm. On Aster, Conyza and Erigeron in east Canada (Olive 1967) and north-eastern USA (Illinois, Washington DC; BMNH collection). There is also now a record of this species from Baja California (Nieto Nafría et al. 2011). U macgillivrayae is very similar to, and possibly synonymous with, the western North American species U. breviscriptum (Palmer).  It is also very similar to U. luteolum,but with a slightly shorter R IV+V; at least some of the records of U. luteolum (and its synonym U. tissoti) from Aster, Conyza and Erigeron may be referable to U. macgillivrayae. However, some specimens from Michigan collected from Conyza canadensis andidentified as macgillivrayae (BNHM collection, leg. N. Moran) differ by characters given in the key to aphids on Conyza, and may be a distinct species.

Uroleucon macolai (Blanchard)    Apterae are very variable in colour, usually yellow-brown to red-brown but sometimes green, with a darker spinal stripe, rather shiny; BL 2.1-3.5 mm. Immatures are red or yellow-green.  On stems and young growth of Baccharis spp. in Argentina, Bolivia and Chile. Nieto Nafría et al. (2007) reported that this species also occurred on Hysterionica jasionoides and Proustia cuneifolia, and subsequently recorded it also from Taraxacum officinale (Nieto Nafría et al. 2016b). Records from other plant genera (Carvalho et al. 1998) are possibly in error. Oviparae and alate males from Argentina were described by Remaudière et al. (1991). 2n=12.

Uroleucon (Uromelan) macrosiphon (Hille Ris Lambers)    Apterae are dark reddish to blackish brown; BL 3.4-3.9 mm. On thistles (Carduus, Carlina, Cirsium) in alpine habitats in Austria and northern Italy, and there are also records from Greece (Tsitsipis et al. 2007, as U. jaceae macrosiphum) and Turkey (Görür et al. 2017) that perhaps require further confirmation. Its biology is unknown. A member of the U. jaceae group, and originally described as a subspecies of U. jaceae.

Uroleucon (Lambersius) madia (Swain)    Alatae are dark green, slightly pruinose (colour of apterae unrecorded); BL of apterae (BMNH collection) 2.5-3.4 mm. On flowerheads of Madia spp. in western USA.

Uroleucon malarguense Ortego & Nieto Nafría     Apterae are brown or red-brown, BL 2.0-2.4 mm. Monoecious holocyclic on a Hypochaeris sp. that is “probably a new species” (Nieto Nafría et al. 2007). In Argentina (Mendoza province, at 2615 m). Oviparae and alate males were collected in late March.

Uroleucon (Lambersius) manitobense Robinson    Apterae are green; BL 2.1-2.5 mm. On Aster sp. in Manitoba, Canada

Uroleucon martini (Cockerell)    Apterae are shining dark wine-red, with long black siphunculi; BL 2.6-3.3 mm. Immatures have a bluish waxy bloom. Alatae have numerous secondary rhinaria (at least 40) on ANT III. On flowerheads of Helenium hoopesii in New Mexico, USA (Cockerell 1903). The species has not been recognised since its original description. Apparently the same aphid was collected from numerous other plants in the type locality (Rudbeckia, Frasera, Zigadenus (= Toxicoscordion), Eriogonum, Potentilla, and Ligusticum), but these records may have been partly due to confusion with other species. [A slide of apterae and immatures collected in 1902 from Helenium in the type locality and labelled as “Nectarophora martini” was sent to BMNH by Cockerell; these specimens seem to be Macrosiphum kiowanepum, so Cockerell’s description may include that species.]

Uroleucon maximilianicola Robinson    Apterae are dark wine-coloured to almost black; BL 2.6-3.4 mm. On Helianthus maximiliani in Manitoba, Canada. Aphids identified as this species have now been recorded from Viguiera dentata and Lagascea sp. at an altitude of 2,600m in Mexico (Nieto Nafría et al. 2011).. This species is very closely related to, and perhaps synonymous with, U. ambrosiae.

Uroleucon mendocinum Mier Durante & OrtegoApterae are reddish brown with brown to dark brown antennae and siphunculi and a white cauda; BL 2.7-3.4 mm. Forming dense colonies on branches and stems of Baccharis juncea in Argentina (Nieto Nafría et al. 2007).

Uroleucon mexicanum Nieto Nafría & Mier Durante    Colour of apterae in life is unrecorded: BL 1.8-3.2 mm. Apterae and alatae were collected from plants in several genera of Compositae/Asteraceae in Mexico; Ageratum, Brickellia, Eupatorium, Perymenium, Roldana, Stevia, Viguiera, Zaluzania (Nieto Nafría et al. 2011). Sexual morphs and biology are unknown.

Uroleucon mierae Tizado & Nieto Nafría    Apterae are dark, shiny, reddish brown with black antennae and siphunculi, and legs also mainly black; BL 2.5-3.8 mm. Monoecious holocyclic (with dark green alate males) on Andryala spp., and also recorded from Hispidella hispanica (Tizado & Nieto Nafría 1994) and Rhagadiolus stellatus (BMNH collection, leg. H.L.G. Stroyan).  This species closely resembles U. picridis, but has a lower range of values of R IV+V, and in some published works it will key to U. cichorii; therefore some records of both U. cichorii and U. picridis in the Mediterranean area may apply to this aphid. In Spain, Algeria, France, Corsica, Sicily and Croatia (Korčula).

Uroleucon (Uromelan) minor (Börner)    Apterae are brown with black dorsal spots, legs yellow banded with black, antennae, siphunculi and cauda black; BL 2.6-4.5 mm. Originally found on Tanacetum corymbosum, but the normal hosts seems to be Serratula (and Klasea)spp. Widely distributed in Europe, east to south-west Siberia and Kazakhstan (Kadyrbekov 2017a). Differences from U. jaceae could be host plant-related (Stroyan 1991).

Uroleucon (Uromelan) minosmartelli Barbagallo & Patti    Apterae are shiny dark brown with black antennae, siphunculi and cauda, legs bicoloured ochreous and black; BL 1.3-2.25 mm. On Campanula spp. that live in rocks and walls, colonising leaves (especially on upper sides along basal part of mid-rib) and flower stems. Described from Italy, where it is apparently anholocyclic, at least in the south (original description), and also found in Poland (Osiadicz & Hałaj 2017) and former Yugoslavia (BMNH collection, leg. H.L.G. Stroyan).

Uroleucon (Uromelan) minutum (van der Goot)    Apterae are shiny dark reddish brown to blackish brown with black antennae, distal parts of femora and tibiae, siphunculi and cauda; BL 1.9-2.1 mm. Alatae have only c.8-10 secondary rhinaria on ANT III. On shoot apices and undersides of leaves of  Vernonia cinerea in south India and Sri Lanka. Anholocyclic (David 1956a, as dravidiana). A record of Macrosiphoniella sanborni on V. cinerea (George 1927) should probably be referred to this species.

Uroleucon mongolicum Holman    Apterae are reddish brown with dark antennae and siphunculi, pale tibiae and cauda; BL 2.3-3.6 mm. On undersides of leaves of Klasea centaureoides and Klasea sp. in Mongolia. Also recorded from Centaurea monantha (= Stemmacantha uniflora)in Korea, and Pashchenko (2000a) described a population on Carduus sp. in east Siberia as a subspecies, U. mongolicum ssp. cardui. Kanturski & Barjadze (2020) provided photomicrographs of specimens from the type series.

Uroleucon (Uromelan) montanivorum Mosbacher    Apterae are reddish brown to black, with a rather metallic sheen; BL 3.6-4.4 mm. On apical leaves and shoots of Centaurea montanum in southern Germany, France and Switzerland, and recently found in Wales (R. Dransfield, pers. comm.; see influentialpoints.com/Gallery). Oviparae and alate males occur in late August-September.  Mosbacher (1959) gave a full morphological and biological description.

Uroleucon monticola (Takahashi)    Apterae are shiny green, dusky around bases of siphunculi, with distal halves of femora and tibiae brown-black, black siphunculi and a contrastingly pale yellow cauda; BL c. 2.6 mm. On Aster and Erigeron in Japan, Taiwan, China and Korea (all the Japanese specimens from several localities were collected from Aster ageratoides var semiamplexicaulis; Miyazaki 1971). Meguroleucon longqishanense Zhang & Qiao, described from Viola verecunda (= V. arcuata) in Fujian province, China (Qiao & Zhang 1999c) is possibly based on vagrant individuals of U. monticola.

Uroleucon muermosum (Essig)   Apterae are dark reddish to brown or almost black in life; BL c.4.5 mm. Described from a yellow composite plant, and subsequently recorded from Senecio yegua (Nieto Nafría et al. 2016b). Only known from Chile.

Uroleucon (Lambersius) mulgedii (Nevsky)    Apterae are pale yellow-green or reddish brown, with siphunculi dark only at apices; BL 2.6-3.2 mm. On Mulgedium tataricum, feeding on petioles and upper surfaces of leaves, and on stems and in inflorescences, dropping readily when disturbed. Also recorded from Sonchus arvensis (Kadyrbekov & Aoitzhanova 2005), Lactuca serriola (Kadyrbekov 2017a) and Cichorium intybus. In Central Asia, Ukraine (Bozhko 1976) and Bulgaria (Tashev 1967, as Macrosiphum mulgedifolii). This species was placed in subgenus Lambersius by Kanturski & Barjadze (2020).

Uroleucon murale (Buckton)    Apterae are reddish brown with black siphunculi and yellow cauda; BL 2.1-3.2 mm. On upper parts of stems and in inflorescences of Mycelis muralis, and also recorded from Lactuca quercina. Throughout Europe (except Iberian peninsula), eastward to Byelorussia, Ukraine and Iran. North American records are probably erroneous (Robinson 1985). Monoecious holocyclic, with sexuales in July-September in Sweden (Ossiannilsson 1959). Buga & Stekolshchikov (2006) fully described the ovipara. A specimen in the BMNH collection contains an internal cecidomyid parasitoid (?Endaphis sp.).

Uroleucon nahuelhuapense Nieto Nafría & von Dohlen  Apterae are bright emerald green with brownish green siphunculi; BL 2.4-3.5 mm. On small stems of Adesmia boronioides in Neuquén province, Argentina (Nieto Nafría et al. 2019a). Holocyclic on A. boronioides; alatae, oviparae and apterous males were described by Mier Durante et al. (2020).

Uroleucon (Uromelan) neocampanulae (Takahashi)    Apterae are brownish black with black antennae, siphunculi and cauda, legs bicoloured yellowish and black; BL 2.5-3.0 mm. On stems of Campanula spp. and Platycodon grandiflorum in Japan, and east Siberia (Pashchenko 1988a).

Uroleucon (Lambersius) nevadense Robinson    Apterae are shining dark green with antennae, legs and siphunculi mainly black; BL 1.1-1.3 mm. On Brickellia microphylla in western USA (Nevada, Idaho).

Uroleucon  (Uromelan) nigrocampanulae (Theobald)    Apterae are dark brown with black antennae, siphunculi and cauda, and bicoloured yellowish brown/black legs; BL 2.7-4.0 mm. On Campanula spp., feeding on the leaves and causing them to become curled in spring, and later spotted with yellow. Sexual morphs are apparently unrecorded. In Europe and across Asia to east Siberia (Pashchenko 1988a).  This species is very similar to U. rapunculoides.

Uroleucon nigrotibium (Olive)    Apterae are red-brown, with black antennae, legs and siphunculi, and pale cauda (see influential points/Gallery); BL 1.9-2.2 mm. Described from an unidentified Solidago sp. in North Carolina, and recorded as host-specific on Solidago nemoralis in Michigan, where long-lived populations consist mainly of apterae that walk frequently between plants (Moran 1986).

Uroleucon nigrotuberculatum (Olive)    Apterae are bright to dull orange-red with dark antennae, legs and siphunculi, and pale cauda (see influential points/Gallery); BL 2.4-4.3 mm. Monoecious holocyclic on Solidago spp. in North America, and introduced in the early 1990s to Japan, where shortly after introduction it was found on plants in several other composite genera (Callistephus, Chrysanthemum, Helianthus, Rudbeckia), and on Oenothera erythrocephala, but this was apparently only a temporary trait, and it soon became more specific to the (non-native) Solidago altissima (Sugimoto & Matsumoto 2000). At high altitudes (1600-3000m) in Mexico it has been recorded from Eupatorium and Montanoa spp. (Nieto Nafría et al. 2011). Cappuccino (1987) compared its population dynamics with that of U. caligatum feeding on the same host (S. altissima).  Many of the records of U. ambrosiae on Solidago should probably be referred to this species. U. pieloui and U. gigantiphagum are very similar, seeming to differ only in the degree of leg pigmentation and/or sporadic occurrence of marginal abdominal tubercles, and may transpire to be synonyms. 2n=12.

Uroleucon (Lambersius) nodulum (Richards)    Colour of apterae in life is unknown, probably green with black antennae, legs and siphunculi and a pale cauda; BL c. 2.0-2.2 mm. On Aster sp. in Ontario, Canada.

Uroleucon (Lambersius) nuble (Essig)    Colour of apterae in life probably pale green with dark brownish head, antennae, distal parts of femora and tibiae, siphunculi (except at bases) and cauda ; BL c. 3.3 mm. Alatae are unknown. Described from an unidentified composite plant in Chile (Essig 1953), and subsequently recorded from that country on Symphyotrichum (=Aster) squamatum and Senecio smithii (Nieto Nafría et al. 2016b). Only the original description is available, and this does not provide the information needed to include this species in the keys to aphids on Aster and Senecio.

Uroleucon obscuricaudatum (Olive)    Apterae are red-brown to reddish, with dark siphunculi and dusky brown cauda; BL 2.7-3.4 mm. On Helianthus spp.in north-eastern USA and Canada (Robinson 1985). An aphid collected on a species of Eupatorium (?inuloides) at 1600m in Mexico has also been assigned to this species (Nieto Nafría et al. 2011).

Uroleucon obscurum (Koch)    Apterae are reddish brown to bronze with black siphunculi and yellow cauda (see influentialpoints.com/Gallery); BL 2.2-3.7 mm. On upper parts of stems of Hieracium spp. in Europe, south to Spain (Nieto Nafría & Mier Durante 1985), Central Asia and east Siberia. There was early confusion with U. picridis; records of  picridis from Hieracium are probably all or mostly referable to this species. Oviparae and alate males are produced over an extended period from late July-September in north-west Europe (Ossiannilsson 1959).

Uroleucon ochropus (Hille Ris Lambers)    Apterae are probably red-brown; BL c.2.9 mm. Described from Chrysanthemum leucanthemum (= Leucanthemum vulgare); later found in large colonies on Lactuca perennis (Hille Ris Lambers 1966b), and also recorded from Mulgedium plumieri (BMNH collection, leg. R. Stäger). In continental Europe, east to south-west Russia, Ukraine and Iran..

Uroleucon olivei Moran    Apterae are deep red with black antennae and siphunculi, mainly black legs and a pale cauda; BL 2.2-3.8 mm. On stems of Aster spp., forming especially large colonies on A. simplex, and also recorded in mid to late summer from Erigeron annuus and Solidago graminifolia (Moran 1985). In eastern USA and Canada (Manitoba, Ontario).  Many of the records of  U. ambrosiae from Aster in eastern North America should probably be referred to this species.

Uroleucon (Uromelan) omeishanense Tao    Apterae are yellowish green with black tibiae, siphunculi and cauda; BL c.2.8 mm. On Artemisia capillaris in Taiwan.

Uroleucon (Uromelan) orientale (van der Goot)    Apterae are dark reddish brown to almost black, with black siphunculi and brown cauda; BL 2.7-3.7 mm. In large colonies on undersides of leaves of Blumea balsamifera in south-east Asia (Java, Borneo, Sarawak, Philippines). Redescribed by Calilung (1967).

Uroleucon (Uromelan) pachysiphon (Börner)    Colour of apterae in life is unrecorded, possibly reddish brown, with black appendages; BL c.2.9-3.3 mm. On Aster amellus in central Europe (Germany, Czech Republic).

Uroleucon (Uromelan) parvotuberculatum (Olive)    Apterae are green with dark brown appendages including siphunculi and cauda; BL 2.1-2.6 mm. On Helianthus atrorubens in North Carolina, USA, records from some other plant genera (Favret et al. 2010) possibly being vagrants.

Uroleucon patagonicum Nieto Nafria & Seco Fernández    Apterae are shining vermilion red with dark brown/black siphunculi and a white cauda; BL 2.3-2.9 mm. In compact colonies on young shoots of Mutisia spinosa in Argentina (Nieto Nafría et al. 2007) and Chile (Nieto Nafría et al. 2020b).

Uroleucon paucosensoriatum (Hille Ris Lambers)    Apterae are bronzy brown, more reddish brown in region of siphunculi, with black antennae and siphunculi, black legs except pale basal halves of femora and pale cauda with dark tip; BL 1.9-3.6 mm. In eastern USA and Canada it occurs on inflorescences and on stems of  Aster and Symphyotrichum spp., and especially (in Michigan) on A. umbellatus (Moran 1985).  There is also a record from Erigeron annuus (Leonard 1968). Oviparae and immature alate males were found in Maine in September (original description). This species is now recorded to occur in Mexico on a wide spectrum of composite genera (Baccharis, Brickellia, Heterotheca, Stevia, Viguiera)at altitudes of 850-3000 m (Nieto Nafría et al. 2011). 2n=12.

Uroleucon payuniense Ortego & Nieto Nafría    Apterae are green with a dark green pleural abdominal patch; BL 2.5-3.0 mm.  On Grindelia chiloensis in Mendoza province, Argentina (Nieto Nafría et al. 2007).

Uroleucon penae Nieto Nafría & Remaudière    Colour of apterae in life unrecorded; BL 2.5-2.8 mm. On a species of Erigeron in Mexico (Nieto Nafría et al. 2011). The type material includes a single alata.

Uroleucon (Lambersius)  penderum Robinson    Apterae are green with extreme apices of femora, ends of tibia and distal parts of siphunculi dark brown; BL 1.8-2.7 mm. Described from Grindelia integrifolia on Pender Island, British Columbia, Canada, and also found on Grindelia sp. in Oregon, and on a beach-side shrub in California (A. Jensen, aphidtrek.org). An aphid collected on Heterotheca inuloides at an altitude of 1800m in Mexico has also been assigned to this species (Nieto Nafría et al. 2011).  2n=12. [Three other species of Uroleucon subgenus Lambersius have been collected on Grindelia spp. in western N. America; U. richardsi (q.v.), and two undescribed species close to penderum but distinguishable by the characters given in the key.]

Uroleucon pepperi (Olive)    Apterae are dark red-brown to metallic black, BL 2.6-4.0 mm. On Cirsium spp. in western USA, and also recorded from Centaurea dealbata.

Uroleucon (Uromelan) phyteumae (Bozhko)    Apterae are uniformly black, slightly shiny; BL 1.5-3.2 mm. On apical parts of Asyneuma canescens. In eastern Europe (Czech Republic, Ukraine). This aphid has large marginal tubercles on prothorax and abdominal tergites 2-4 (Holman 1969, as asyneumatis).

Uroleucon picridiphagum (Takahashi)    Colour of apterae in life is unrecorded, probably dark brown; BL c.3.2 mm. On Picris hieracioides var. japonica in Japan. Closely related to U. sonchi, but with a slightly longer R IV+V and fewer hairs on cauda.

Uroleucon picridis (Fabricius)    Apterae are dark shiny reddish brown, with black antennae and siphunculi, legs brown black except coxae, trochanters and basal halves of femora yellow, cauda yellow (see influentialpoints.com/Gallery); BL 2.6-3.7 mm. On Picris spp., forming colonies on stems just below flowers. Also recorded from other genera of Cichoriaceae (e.g. Cichorium, Crepis, Hieracium, Ixeridium, Lactuca, Leontodon, Sonchus), although some records are probably misidentifications of other species including, in the Mediterranean area, U. mierae (q.v.). In Europe, Middle East, Kazakhstan (Kadyrbekov 2017a), west and east Siberia, Korea, China, Japan, and there is a 1916 record from Java. U. picridis has now also been introduced to California, USA and seems to be established there (Skvarla et al. 2017). Monoecious holocyclic with oviparae and alate males produced in October.

Uroleucon pieloui (Richards)    Apterae are dull red to dark brown, antennae and siphunculi black, legs yellow-brown banded with black, cauda pale; BL 2.1-4.5 mm. On Solidago, with apparent specialisation on more pubescent species (Moran 1985). In north-eastern USA and Canada. U. gigantiphagum is not clearly separable, and both are also very closely related to, if not synonymous with, U. nigrotuberculatum (q.v.).

Uroleucon pilosellae (Börner)    Apterae are dark reddish brown with black siphunculi and pale yellow cauda (see influentialpoints.com/Gallery); BL 2.2-2.5 mm. On flower stems of Hieracium spp. throughout Europe, and also in Turkey and Iran (where aphids identified  as this species were found on Scorzonera sp.; Mehrparvar 2017). A record from Leontodon in Algeria (Laamari et al. 2013) requires additional confirmation. The sexual morphs are undescribed; the male is reportedly alate (Hille Ris Lambers 1939a, citing Börner).

Uroleucon pseudambrosiae (Olive)    Apterae are dark brown to red-brown with black antennae, legs and siphunculi, and a pale cauda; BL 2.2-3.4 mm. Recorded from numerous genera and species of Compositae/Asteraceae, but the most common hosts are Lactuca spp. (Olive 1963). Records from Aster prior to 1985 are possibly referable to U. olivei. In eastern USA south to Florida, Mexico, Canada (Manitoba, British Columbia), and introduced to Europe (Poland; Osiadacz & Halaj 2010). The record from Turkey (Akyürek et al. 2010) is based on a misidentification (Juan Nieto Nafría, personal communication). 2n=12.

Uroleucon pseudobscurum Hille Ris Lambers    Apterae are very dark bronze, with black siphunculi and pale yellow cauda; BL 2.5-3.2 mm. In large colonies on stems of Hieracium spp. in Italy, Hungary (BMNH collection, leg. V.F.Eastop), Slovakia, Serbia (Petrović Obradović et al. 2020, Belarus (Stekolshchikov et al. 2010), Georgia (Barjadze et al. 2010b), western Siberia (Stekolshchikov & Novgorodova 2013), Kazakhstan (Kadyrbekov, 2003a) and also found in Japan (Miyazaki 1971, as picridis on Hieracium umbellatum).

Uroleucon pseudomuermosum Carvalho    Colour of apterae in life is unknown; BL 2.3-3.2 mm. On Baccharis sp(p). in Llanquihue Province, Chile (Carvalho et al. 1998).

Uroleucon pseudotanaceti (Verma)    Apterae are brown with black siphunculi; BL 2.0-2.5 mm. Described from undersides of leaves of  Helianthus tuberosus in north-west India, and since found on plants in other composite genera (Chrysanthemum, Cynoglossum). Singh et al. (1980) found oviparae on Chrysanthemum sp. in January. 2n=12.

Uroleucon ptarmicae (Bozhko)    Apterae are brownish red with black siphunculi and pale cauda; BL c.3.1 mm. In dense colonies on flower stems of Achillea cartilaginea (= Ptarmica speciosa) in Ukraine (Bozhko 1976b).

Uroleucon pulicariae (Hille Ris Lambers)    Apterae are dark reddish brown with black siphunculi and pale yellow cauda (see influentialpoints.com/Gallery); BL 2.8-3.1 mm. On Inula spp. and Pulicaria spp. in southern, central and eastern Europe, Middle East, Central Asia, and also recorded from North Korea (Holman 1981c).

Uroleucon pyrethri Holman    Apterae are brown, with siphunculi slightly dusky, darker at base and apex, and a broad pale cauda; BL 3.2-4.0 mm. On Pyrethrum macrophyllum, feeding onundersides of leaves, especially lower ones, which tend to become yellow (original description). In the Caucasus (Georgia). Very similar to U. tussilaginis. 

Uroleucon (Lambersius) queretarense Pérez Hidalgo & Nieto Nafría    Colour of apterae in life is unrecorded; BL 1.7-2.5 mm. Apterae and alatae were collected on Aphanostephus ramosissimus in October at an altitude of 2100m in Mexico (Nieto Nafría et al. 2011).

Uroleucon (Uromelan) rapunculoidis (Börner) (= Dactynotus glomeratae Börner)    Apterae are shiny dark brown with black antennae, siphunculi and cauda; BL 3.0-3.9 mm. On stems and flowers of  Campanula rapunculoides in Europe, and recorded from  C. glomerata and C. sibirica in south-west and central Asia. Oviparae were found in October in the Netherlands (Hille Ris Lambers 1939). It is very similar to U. nigrocampanulae.

Uroleucon (Lambersius) remaudiereorum Nieto Nafría & Mier Durante   Colour of apterae in life is unrecorded; BL 2.3-3.2 mm. Apterae and alatae were collected in October on a species of Conyza at an altitude of 2000m in Mexico (Nieto Nafría et al. 2011).

Uroleucon reynoldense (Olive)    Apterae are dark orange-red; BL 1.6-2.2 mm. On Coreopsis spp. in eastern USA (North Carolina, Michigan). Also recorded from New Mexico, where sexuales were collected in early October (A. Jensen, aphidtrek.org), and found at high altitudes in Mexico on Viguiera spp. and Kuhnia (= Brickellia) sp. (Nieto Nafría et al. 2011). It is also recorded from Jamaica on Solanum torvum (Tannice Hall, pers. comm.), but this is an unlikely host. 2n=12.

Uroleucon (Lambersius) richardsi Robinson    Apterae are green with antennae dark beyond base of ANT III, distal parts of femora and whole of tibiae and tarsi dusky/dark, and siphunculi dark on usually about distal 0.6 of length; BL 1.6-2.0 mm.  On Grindelia spp. in western USA (Oregon, Utah) and Canada (Manitoba), and aphids collected on a species of Viguiera at c.1900m altitude in Mexico have now been identified as this species (Nieto Nafría et al. 2011). Oviparae and apterous males occur in early October in Manitoba (Robinson 1965). 2n=12.

Uroleucon riojanum Nieto Nafría & Mier Durante    Apterae are green with antennae, legs and siphunculi distally dark brown, and a greenish white cauda; BL 1.7-2.2 mm. On Gutierrezia isernii, aligned on both sides of leaves or on stems. In La Rioja province of north-west Argentina, at 2020 m (Nieto Nafría et al. 2007). Similar to U. erigeronense but with a longer R IV+V and thicker siphunculi.

Uroleucon (Uromelan) riparium (Stroyan)    Apterae are dark bronze-brown with black antennae, siphunculi, cauda, tarsi, apices of femora and tibiae; BL 2.9-4.1 mm. Living in small colonies on the flower stems of  Crepis spp. in NW Europe (Scotland, Sweden, Finland), and reported to occur also on Cirsium sp. and Tragopogon pratensis in Kazakhstan (Kadyrbekov 2014e, 2017a); in Finland it was also found on Taraxacum (Heie 1995, citing Heikinheimo). Oviparae and alate males occur from late July (in Finland) to September.

Uroleucon (Lambersius) robinsoni Barjadze & Kanturski   Colour in life unknown; BL of apterae 2.1-2.8 mm. On Grindelia squarrosa in Colorado, USA (Barjadze & Kanturski 2020). Other morphs and life cycle are unknown.

Uroleucon rudbeckiae (Fitch)    Apterae are shiny bright orange-red to brick red with mainly blackish appendages, but with bases of siphunculi  distinctly paler, and a pale/dusky cauda (see influential points/Gallery); BL 2.4-3.2 mm. Often in large colonies on stems of Rudbeckia spp. throughout North America. Records from other genera of Compositae/Asteraceae are likely to be misidentifications. Monoecious holocyclic, with alate males. Lamb & Mackay (2010, 2017) have conducted long-term studies of the population dynamics of this aphid. 2n=12.

Uroleucon (Uromelan) rurale (Hottes & Frison)    Apterae are grass-green with blackish antennae, legs, siphunculi and cauda; BL 2.8-3.4 mm. On Verbesina spp. in eastern USA as far south as South Carolina (further south is it is replaced by the closely-related U. verbesinae). Monoecious holocyclic, with alate males. 2n=10.

Uroleucon russellae (Hille Ris Lambers)    Apterae are bronzy black with black antennae, legs mainly black except for basal halves of femora, black siphunculi and a dusky, sometimes almost black, cauda (see influential points/Gallery); BL 2.5-3.3mm. On stems and leaves, and in inflorescences, of  Anaphalis margaritacea and Antennaria sp., and also collected from three Gnaphalium spp. in California (Hille Ris Lambers 1966a), and from Helichrysum virgineum in British Columbia (Forbes & Chan 1989).  Widespread in western and northern USA and across Canada. Monoecious holocyclic, with alate males (original description). 2n=12.

Uroleucon saussureae (Takahashi)    Apterae are reddish black, with black siphunculi and pale cauda; BL 2.8-3.0 mm. On Saussurea spp. in Japan (Miyazaki 1971).

Uroleucon (Uromelan) scorzonerae Danielsson    Apterae are dark red or blackish red with dark siphunculi and cauda; BL 2.5-3.2 mm. On undersides of leaves of Scorzonera humilis, which become faded and yellow. Only known from Sweden until a recent record from Turkey (Görür et al. 2011b, 2012), which perhaps requires additional confirmation. Apparently it is visited by ants (original description).

Uroleucon (Uromelan) scrophulariae (Bozhko)    Apterae are brown or reddish with black siphunculi and cauda; BL c. 2.7 mm. On Scrophularia vernalis in Ukraine.

Uroleucon (Uromelan) seneciocola (Paik)    Apterae are brown; BL 2.6-2.9 mm. On Senecio pierotii in Korea. This species is similar to U. compositae (or U. gobonis), but with less extensive rhinariation of ANT III, in which respect it resembles U. dzhungaricum.

Uroleucon (Uromelan) siculum Barbagallo & Stroyan    Apterae are dark brown, with appendages mainly black, BL 2.1-3.5 mm. Alatae have 29-58 secondary rhinaria on ANT III.  This species is able to colonise an unusually wide range of hosts within Compositae/Asteraceae (Anthemis, Leucanthemum, Pulicaria), and colonies have also been found more than once on flower stems of Rumex conglomeratus (original description). In Sicily and central Italy (Barbagallo & Patti 1998). A record from Turkey (Görür et al. 2011b, 2012) requires additional confirmation.

Uroleucon sijpkensi Hille Ris Lambers    Apterae are brown, with antennae and legs pale contrastingly banded with black, siphunculi black and cauda pale; BL 2.8-3.4 mm. On Solidago macrophylla in Quebec, Canada. Oviparae were collected in mid-August (original description).

Uroleucon (Uromelan) sileneobium (Narzikulov)    Apterae are shining brown, with antennae, siphunculi and cauda almost black; BL c.3.1 mm. On stems of Galatella sedifolia in Tajikistan, and Kadyrbekov (2017a) reported it from Crepis sp. in Kazakhstan. The host was originally given as Silene scabrifolia, but subsequently corrected (see Remaudière & Remaudière 1997, p.313).

Uroleucon (Uromelan) simile (Hille Ris Lambers)    Apterae are shining reddish brown, with black antennae, siphunculi and cauda, and legs yellowish brown with apices of segments black; BL 2.3-4.1 mm. Alatae have 60-85 secondary rhinaria on ANT III.  Colonies form on flowerstalks of  certain Erigeron spp. (especially E. acris), Conyza canadensis, and it is also recorded from Tanacetum boreale (Pashchenko 1988a). It may be a pest of hybrids of  E. speciosum x macranthus grown as ornamentals (Hille Ris Lambers 1967).  Throughout Europe, Central Asia, India (Kashmir), west and east Siberia. Oviparae and small dark apterous males were found in September in the Netherlands, on the undersides of radical leaves (Hille Ris Lambers 1939a).

Uroleucon simlaense Chakrabarti, A.K. Ghosh & Raychaudhuri    Apterae are grey with blackish antennae and siphunculi; BL c. 3.6-3.9 mm. Alatae have c.62-65 secondary rhinaria on ANT III. On undersides of leavesof  Erigeron sp. in Himachal Pradesh, India. 2n=12.

Uroleucon sinuense Mier Durante & Nieto Nafría     Colour of apterae in life is unrecorded; BL 2.7-4.2 mm. Described from a large sample of apterae and alatae collected in April on a species of Gnaphalium at sea level in Baja California, Mexico (Nieto Nafría et al. 2011).

Uroleucon skurichinae Pashchenko    Apterae are reddish brown or brown, shining; BL c.4.4 mm. Alatae have 46-57 secondary rhinaria on ANT III.  On Saussurea spp., feeding on upper parts of stems and flower shoots, and undersides of leaves (Pashchenko 2000a). E Siberia (Primorskii).

Uroleucon (Uromelan) solidaginis (Fabricius)  (Fig.49d)   Apterae are shining reddish brown with black dorsal spots, antennae and legs mainly yellowish brown with darker bands, siphunculi and cauda black (see influentialpoints.com/Gallery); BL 2.3-4.1 mm. Immatures are bright red. On upper parts of stems of Solidago virgaurea in Europe, Asia, north Africa and North America. Monoecious holocyclic, with oviparae in September-October, and alate males recorded from mid-July to October.

Uroleucon solirostratum (Richards)    Colour of apterae in life is unobserved; BL up to 2 mm. On Solidago flexicaulis in Canada (Manitoba, Ontario) (Robinson 1985).

Uroleucon sonchellum (Monell)    Apterae are brick red to dark red-brown or dull red, with black siphunculi and pale cauda; BL 2.2-3.5 mm. On leaves and stems of Lactuca and Sonchus spp., widely distributed in USA. Monoecious holocyclic, with sexual morphs in August (Colorado) to October-November (North Carolina). Redescribed by Olive (1963), and compared in considerable detail with U. formosanum by Pike et al. (2005). 2n=12.

Uroleucon sonchi (L.)  (= Uroleucon parasonchi Raychaudhuri, Raha & Raychaudhuri; Chakrabarti & Medda 2004)  Plate 28e     Apterae are shiny dark brown, antennae mainly dark, legs mainly pale with black coxae and apices to femora and tibiae, siphunculi black, cauda pale yellow (see influentialpoints.com/Gallery); BL 2.9-4.5 mm. Mainly on Sonchus spp. and other genera in the tribe Cichoriaceae (Lactuca, Cichorium, Hieracium, Ixeridium, Picris, Reichardia),but also sometimes recorded from other Compositae/Asteraceae. It has an almost world-wide distribution. Monoecious holocyclic, with apterous males in cold northern temperate regions, presumably anholocyclic. in milder climates. Subspecies have been described from Cousinia in Afghanistan (afghanica Narzikulov) and Kazakhstan (stepposa Smailova), but neither seem to be reliably differentiated from sonchi s. str. However, there are specimens in BMNH from Pakistan (leg. M.A. Ghani) with short dorsal hairs, and from Nepal (leg. K.C. Sharma) with 0-4 rhinaria on ANT IV of alatae, which could qualify for subspecies status. Carver (1999) gave an account of this aphid in Australia. 2n=12.

Uroleucon (Uromelan) stachydis (Bozhko)    Apterae are shining blackish brown; BL c.3.3 mm. At stem apices of  Stachys spp., recorded from Latvia and Ukraine.

Uroleucon stoetzelae Robinson    Apterae are green with light to dark brown appendages, siphunculi pale at bases and cauda pale; BL 1.3-1.8 mm. Described from Achillea millefolium in Pennsylvania, USA, but also now recorded from Erigeron sp. in Mexico (Nieto Nafría et al. 2011).

Uroleucon (Lambersius) suzannae Robinson    Apterae are very pale green except for tips of appendages; BL 2.0-2.7 mm. Alatae are unknown. On Haplopappus hirtus var. sonchiolius in Oregon, USA.

Uroleucon (Uromelan) syrdariense Kadyrbekov    Apterae are shining blackish brown; BL 3.1-3.4 mm. On stems of Senecio jacobaea in Kazakhstan (Kadyrbekov 2003a).

Uroleucon tanaceti (L.)    Apterae are bright red or reddish brown, with yellowish, black-banded antennae and legs, brown-black siphunculi and yellow cauda (see influentialpoints.com/Gallery); BL 2.2-3.4 mm. On Tanacetum spp., feeding primarily on the lower, often yellowing, leaves. The effects of the aposematic coloration of this aphid in relation to its on-plant distribution was studied by Benedek et al. (2019). In Europe, west Siberia, south-west and Central Asia, eastern Himalayas, and North America. Monoecious holocyclic, with oviparae and alate males in October-November (in southern England). In Finland oviparae of this species have been found on cultivated chrysanthemums (Heie 1995). 2n=12.

Uroleucon (Uromelan) taraxaci (Kaltenbach)    Apterae are shining dark bronze-brown, with black dorsal spots, and black antennae, legs (except femoral bases), siphunculi and cauda (see influentialpoints.com/Gallery); BL 2.5-3.8 mm. On Taraxacum spp., living on undersides and at bases of leaves near ground level. In Europe, south-west and central Asia, North America, and recently recorded from Argentina (Ortego et al. 2018) and Chile (Nieto Nafría et al. 2018). Monoecious holocyclic with oviparae and small apterous males appearing in the northern hemisphere in late September. 2n=12.

Uroleucon tardae Hottes & Frison    Apterae are dark red-brown to blackish red, with black antennae and siphunculi, legs dark except for yellow-brown coxae and basal parts of femora, and cauda concolorous with abdomen; BL 1.7-2.5 mm. On Helenium autumnale,feeding at apices of flower stalks, directly under flowers (original description). In north-eastern USA. Oviparae and alate males occur in October. Redescribed by Olive (1963).

Uroleucon teheranense Nieto Nafría & Pérez Hidalgo  Colour of apterae in life is unknown, siphunculi dark with cauda paler; BL c.2.6-2.8 mm (Nieto Nafría & Pérez Hidalgo 2013a). On Lapsana sp. (presumably communis) in Iran (Tehran province). Alatae and life cycle are unknown.

Uroleucon telekiae (Holman)    Apterae are reddish black to dark bronze, with antennae black except for III and basal part of IV, black siphunculi and pale/dusky cauda; BL 3.0-4.6 mm. On apical parts and causing deformation of young leaves of Telekia speciosa in Europe, and Turkey. Monoecious holocyclic with oviparae and alate males in October (Müller & Steiner 1989).

Uroleucon (Lambersius) tenuitarsum (Gillette & Palmer)    Apterae are apple green with antennae and legs mainly dusky-dark, siphunculi black with a paler base, and a pale green cauda; BL 2.5-3.0 mm. On leaves of  Aster and Symphyotrichum spp. in western USA. Oviparae and alate males occur in early October (Palmer 1952).

Uroleucon tessariae Delfino    Apterae are pale green with dark antennae and siphunculi; BL 2.6-3.1 mm. On flower buds and stalks of Pluchea (= Tessaria) absinthoides, forming dense colonies that disperse quickly when disturbed. In Argentina (Tucumán).

Uroleucon tortuosissimae Rezwani & Lampel    Apterae are matt brown, with dark brown siphunculi and pale cauda; BL 1.8-2.6 mm. On Scorzonera tortuosissima in Iran. 

Uroleucon (Uromelan) tripartitum (Ivanoskaya)    Apterae are greenish brown with dark antennae, legs, siphunculi and cauda; BL c.5.0-5.1 mm. On Bidens tripartita in Siberia. This species seems to closely resemble U. jaceae.

Uroleucon (Uromelan) triphylli Miyazaki  ( = Uroleucon adenophorae Miyazaki 1971 and Holman 1975, nec. Matsumura 1918)   (Fig.7b)  Apterae are reddish brown with mainly black appendages; BL 2.5-3.2 mm. On Adenophora spp. in Japan, Mongolia and Russia (Transbaikalia).

Uroleucon (Uromelan) tschuense Kadyrbekov    Apterae are shining blackish brown; BL 2.7-3.1 mm. On stems of Chamaegeron oligocephalus and Conyza canadensis in Kazakhstan (Kadyrbekov 2003a, 2017a). This species seems close to U. simile.

Uroleucon (Uromelan) tuataiae Olive    Apterae are dark red-brown with blackish antennae, legs, siphunculi and cauda: BL 2.4-3.1 mm.  On Ambrosia spp., living on undersides of leaves and in inflorescences. Recorded from eastern USA (New Jersey, Pennsylvania, North Carolina, Florida) and from Cuba (Holman 1974b). 2n=12.

Uroleucon tucumani (Essig)    Apterae are red-brown, BL c.2 mm. On Baccharis spp.in Argentina and Chile (Carvalho et al. 1998, Nieto Nafría et al. 2007). The latter authors also record its occurrence on Parthenium hysterophorus, Bidens sp. and Conyza sp.  U. littorale(Blanchard), described as dark green in life and collected on B. melastomaefolia (= punctulata)in Argentina, is very similar in morphology according to the original description, and is possibly an earlier name for this species..

Uroleucon tussilaginis (Walker)    Apterae are shining brown, antennae pale basally and dark distally, siphunculi black at base and apex but with yellowish brown middle section, cauda yellow; BL 2.4-4.3 mm. On undersides of leaves of Tussilago and Petasites spp. in Europe, and south-west and central Asia. Monoecious holocyclic, with oviparae and apterous males in late September to November. 2n=8? (Kuznetsova 1974, as Dactynotus basalis Walker?; however the karyotype illustrated resembles that of Acyrthosiphon pisum).

Uroleucon ussuriense Pashchenko    Apterae are blackish, with pale middle sections of tibiae and cauda; BL c.3.2 mm. Alatae have 40-44 secondary rhinaria on ANT III. On Saussurea spp., feeding on upper parts of stems and undersides of apical leaves (Pashchenko 2000a). In east Siberia (Primorskii).

Uroleucon (Uromelan) uyguricum Kadyrbekov, Renxin & Shao    Apterae are brownish; BL 2.1-2.3 mm. On stems of Convolvulus pseudocantabrica in Xinjiang-Uygur region of western China (Kadyrbekov et al. 2002).

Uroleucon vancouverense Robinson    Colour in life of apterae is unrecorded; probably with all appendages dark, including cauda; BL 2.5-4.4 mm. Described from a population on Solidago canadensis var. salebrosa in British Columbia, Canada, and also found in Alberta (BMNH collection, leg. A.M. Harper).

Uroleucon (Lambersius) vera Pashchenko    Apterae are green, BL c. 2.5 mm. On upper parts of stems of  Aster ageratoides in east Siberia (Pashchenko 2001).

Uroleucon (Uromelan) verbesinae (Boudreaux)    Apterae are shining bright red or black, with antennae, legs, siphunculi and cauda all black except for yellow bases of femora (see influential points/Gallery); BL 2.5-3.0 mm. On Verbesina virginica in south-eastern USA (Louisiana, Florida), and on V. myriocephala in Honduras (Evans & Halbert 2007). 2n=10.

Uroleucon (Uromelan) vernoniae (van der Goot)    Apterae are shining black; BL 1.7-3.2 mm. On Vernonia spp. and various other Compositae/Asteraceae (Arctotis, Elephantopus, Emilia, Gynura) in Java (see Noordam 2004).          

Uroleucon vernonicola (Holman)    Apterae are blackish brown, with dark antennae, legs and siphunculi and pale cauda; BL 1.9-2.4 mm. On Vernonia cinerea in Cuba. It appears closely related to U. ambrosiae.

Uroleucon (Lambersius) zacatecense Nieto Nafría & Pérez Hidalgo   Colour of apterae in life is unrecorded; BL 2.2-2.8 mm. On Gymnosperma glutinosum in Mexico (Nieto Nafría et al. 2011). Other morphs including alatae are unknown. This species appears to be very similar to U. erigeronense.

Uroleucon (Lambersius) zayasi (Holman)   Apterae are pale green with dark antennae, legs and siphunculi apices; BL 2.4-2.9 mm. On Solidago sempervirens, colonising upper parts of stems and inflorescences, in Cuba (orig. descr.) and found on an unidentified plant in Costa Rica (BMNH colln, leg. R. Kiester).

Uroleucon (Lambersius) zerogutierrezis (Smith & Knowlton)    Apterae are bright apple green, with antennae dark except basal part of III and siphunculi dark on distal half; BL 1.2-1.8 mm. On Gutierrezia sp. in western USA and on G. sarothrae in Mexico (Nieto Nafría et al. 2011). There are also records from Bidens, Senecio and Chrysothamnus, but there is possible confusion with U. erigeronense, to which it is very similarin every respect except the antennal rhinariation.

Uroleucon zinzalae (Hottes & Frison)    Apterae are green with long dark green-black siph. and yellowish green cauda; BL c. 2.7 mm. Feeding on undersides of leaves along mid-ribs of  Polymnia canadensis in Illinois and Michigan, USA (Moran 1985).

Uroleucon (Lambersius) zymozionense (Knowlton)    Colour of apterae in life is uncertain, probably green with dusky-dark appendages, pale bases of siphunculi and a  pale cauda; BL 1.7-2.3 mm.  On Aster leucanthemifolius in western USA.  Sexuales occur in November (Palmer 1952).

Utamphorophora Knowlton

Aphidinae: Macrosiphini

About 7 species of Hyalomyzus‑like aphids with swollen siphunculi, apterae often having rhinaria on ANT III. Their host associations are mainly with Rosaceae and/or monocots, and perhaps they have greatest affinity with Metopolophium. Taiwanomyzus are also similar but have head dorsally spiculose, and other host associations. Remaudière (1983a) reviewed the genus and provided a key to species, and Cook (1984c) keyed the North American species.

Utamphorophora bossekiae (Gillette & Palmer)     Apterae are pale greenish yellow or pale green, mottled or streaked with darker green: BL c.1.8 mm. On leaves and stems of Rubus deliciosa in western USA (Colorado, Utah).

Utamphorophora bromicola Remaudière    Apterae are pale green with two dark green longitudinal submarginal stripes; BL 2.0-2.8 mm. On Bromus carinatus, feeding onupper sides of leaves close to veins, in Mexico. It is apparently anholocyclic (original description).

Utamphorophora commelinensis (Smith)    Apterae are whitish yellow or yellowish with dark apices to antennal segments. legs and siphunculi; BL 1.3-1.9 mm.  Alatae have a large dark saddle-like dorsal abdominal sclerite joining the bases of the siphunculi. On Commelinaceae (Callista, Commelina) in Puerto Rico, and there are trapped alatae in BMNH collection from Costa Rica, Guatemala, Venezuela and Brazil. There are also Puerto Rican records – probably of vagrants – from Brassica interrifolia and Sonchus oleraceus (Smith et al. 1963).

Utamphorophora crataegi (Monell)   Apterae are lemon to canary yellow, with four widely-spaced grass-green dorsal spots (see Johnson & Lyon 1988 or influential points/Gallery); BL 1.7-2.3 mm. Often forming large colonies on twigs, shoots and leaves of Crataegus spp., causing curling when on leaves. Widely distributed in North America. There is no host alternation; oviparae and alate males occur in September-October (Palmer 1952).

Utamphorophora humboldti (Essig)  Plate 24e   Apterae on grasses are apple green with light brown head and pale siphunculi and cauda; BL 1.9-2.6 mm. Alatae have a green abdomen with variably developed dark intersegmental markings. In aggregations along upper sides of the leaf blades, or hidden in flowerheads, of various Poaceae (Dactylis, Festuca, Lolium, Poa, Polypogon,etc.). Heteroecious holocyclic in North America, with Physocarpus spp. as primary hosts. Introduced into England, where it produces sexual morphs in autumn but is probably mainly anholocyclic on grasses. More recently it was found in the Netherlands on Poa annua, with small colonies also on Juncus bufonius, and in glasshouse conditions it also colonised Avena sativa (Piron 2010b). It is also reported to occur in Greece (Tsitsipis et al. 2007). See Stroyan (1979a) for a detailed account. 2n=20.

Utamphorophora vibei (Hille Ris Lambers)    Colour of apterae in life is unknown; BL 2.6-2.8 mm. Alatae have dark intersegmental markings. On unidentified grasses in Greenland. Monoecious holocyclic with oviparae in early August (original description).