The Aphids

Two North American oak-feeding species with a short antennal terminal process, indented anal plate and rounded cauda. Apterae have the head and thorax fused and an extensively sclerotised tergum, and alatae have wings strongly resembling those of Patchia, with thickly bordered veins and scale-like imbrication of the wing membrane. Quednau (1999) reviewed and illustrated both species.

Lachnochaitophorus obscurus (Tissot) Apterae are mid- to dark brown or almost black; BL 1.6-2.0 mm. Alatae are dark reddish brown with dusky wings, with dark dorsal abdominal cross-bands and wing veins thickly bordered with fuscous. In small colonies on twigs and leaf petioles of young Quercus (nigra, michauxii), attended by ants, or in small numbers on undersides of leaves, where the alatae look very conspicuous against the green of the leaf. In south-eastern USA (Florida, Georgia, Louisiana). Life cycle unknown; oviparae in the original description (Tissot 1932) were collected in Florida in February. L. bisselli is a synonym. Richards (1965) had apterae of L. querceus (see below).

Lachnochaitophorus querceus Granovsky (= Patchia winforii Miller) Apterae are dark brown to black, “almost shiny”; BL 1.7-1.9 mm. Alatae are dark brown to black, with black abdminal cross-bands and wing veins broadly and heavily bordered with fuscous. In small colonies, mainly on leaf petioles and young bark of tender one-year-old shoots, of Quercus rubra (= borealis) and Q. velutina, and also reported forming dense clusters on twigs and leaf petioles of Q. palustris (Miller 1933b, as Patchia winforii). Closely attended by the ant Crematogaster lineolata, which may construct shelters over the colonies. In north-eastern USA (Pennsylvania, Wisconsin, Massachusetts) and Canada (Ontario). Monoecious holocyclic; oviparae (with swollen hind tibiae and a greatly extended posterior abdomen) and small apterous males occur in September in Wisconsin (Granovsky 1933). Richards (1965) synonymised this species with the more southerly L. obscurus, but Granovsky demonstrated consistent differences between L. querceus and L. obscurus (as bisselli), and these differences also hold for material in the BMNH collection . Richards’ description applies to L. querceus.

Lachnus Burmeister

Lachninae: Lachnini

About 15 palaearctic and one nearctic species of medium to large long-legged aphids, the alatae usually having pigmented wings, mostly associated with Fagaceae and attended by ants. It is a taxonomically difficult genus, as indicated under L. roboris. Accounts are available for central Europe (Heinze 1962, Szelegiewicz 1962a), north-west Europe (Heie 1995), the UK (Blackman et al. 2019b), the Iberian peninsula (Nieto Nafría et al. 2002a), India (A.K. Ghosh 1982b), Middle East (Bodenheimer & Swirski 1957), Korea (Lee et al. 1994) and Japan (Sorin 1980, Binazzi & Remaudière 2007). Binazzi & Remaudière (2010) reviewed, keyed and provided excellent illustrations of the Eurasian species on Fagaceae. Until about 1930 the species were often placed in Pterochlorus, and the name Lachnus was applied to aphids of the genus Cinara. Several species probably do not really belong in this genus, particularly some of those described from Salix, which may be closer to Tuberolachnus.

Lachnus acutihirsutus Kumar & Burkhardt Apterae have black head and thorax and brownish black abdomen, with a smooth membranous conical protrusion in the centre of the dorsal abdomen; BL 2.8-4.4 mm. Alatae have forewings pigmented except for two large clear triangles on basal half and a small clear spot between pterostigma and R_S. On twigs of Quercus incana in northern India (Uttar Pradesh, Himachal Pradesh) and on several Quercus spp. in Pakistan (Naumann-Etienne & Remaudière 1995). Also reported from India on Castanopsis (as Quercus) indica (A.K. Mandal et al. 1986). Kanturski et al. (2017b) provided a redescription of apterae and alatae. Sexuales and life cycle are unknown. 2n=16 (Dutta & Gautam 1993).

Lachnus allegheniensis McCook (= montanus Wilson) Apterae are rusty or yellowish brown, with legs brown to black; BL 3.5-4.5 mm. The rare alate morph was described by Hottes (1954a, as montanus); the forewing veins are heavily bordered with fuscous, which extends between the veins at the apex of the wing. On twigs of Quercus spp. (gambelii, gunnisonii, virginiana), collected in various parts of North America, but of rather sporadic occurrence. Monoecious holocyclic, with apterous males and oviparae in October (Palmer 1952).

Lachnus chosoni Szelegiewicz Apterae are in life very shiny dark brown to black (Kanturski et al. 2018c); BL 3.7-4.1 mm. On bark of older branches of an unidentified Quercus sp. in North Korea (Szelegiewicz 1975a, and see also Binazzi & Remaudière 2010). Kanturski et al. (2014b) re-examined the type material, and Kanturski et al. (2018c) reported this species from South Korea (also from an unidentified Quercus sp.), and described oviparae and apterous males collected in October-November.

Lachnus crassicornis Hille Ris Lambers (fig. 93A) Colour of apterae in life is not recorded; BL 2.2-3.0 mm. On Quercus ithaburensis (BMNH collection, leg. D. Hille Ris Lambers) in Israel. It is also recorded from Q. robur in Romania (Holman 2009). Other morphs and life cycle are unknown. Records from Turkey on Q. aegilops (= Q. macrolepis?) (Canakçioglu 1967, 1975) are referable to L. pseudonudus (see Quercus key D)

Lachnus fici Takahashi Apterae brownish black; BL c. 4.5 mm. Alatae have blackish brown pigment at bases of wings, not extending to Cu_1b in forewing, and wing membrane otherwise only slightly dusky around margin. On branches of an unidentified Ficus sp. in Taiwan, and apparently not recorded since the original description (Takahashi 1933c). The synonymy with Nippolachnus himalayensis proposed by Tao (1958) seems doubtful.

Lachnus iliciphilus (del Guercio) Apterae are shining blackish brown; BL 2.6-3.7 mm. Alatae have a pattern of forewing pigmentation very similar to that of L. roboris. On twigs and branches of Quercus spp. (ilex, suber, coccifera) in southern Europe (France, Italy, but apparently not Iberian peninsula). Distribution and biology are uncertain due to confusion in the literature with L. longirostris (= L. pallipes) and L. roboris. Heie’s (1995) account under this name should be referred to L. pallipes. Differences from L. roboris are mainly size-related and require further confirmation by genetic studies.

Lachnus longirostrum David & Ghosh Colour of apterae in life unknown; BL c. 4.4 mm. Alata with hyaline forewings. Described from Salix fragilis in Himachal Pradesh, India, and there are subsequent Indian records from other Salix spp. and from Populus ciliata. A record from Salix sp. in Turkey (Akyürek et al. 2011) needs further confirmation. Biology and life cycle are unknown.

Lachnus margallaensis Kanturski & Zia Apterae in life are dark green; BL 2.4-2.8 mm. Alatae have forewing pigmentation similar to L. acutihirsutus. On branches and shoots of Quercus oblongata in Pakistan (Kanturski et al. 2017b).

Lachnus pallipes (Hartig) (= Lachnus longirostris (Mordvilko); Mróz et al. 2015) Apterae are shining grey-brown to dark reddish or blackish brown, with a clothing of fine pale hairs (see influentialpoints.com/Gallery); BL 2.8-5.0 mm. Alatae have a pattern of forewing pigmentation similar to L. roboris, but the clear patch between the R_sand the media is larger. On older branches and stems of Fagus sylvatica, causing feeding damage by rupture of the cambium, and also on twigs and branches of mostly evergreen species of Quercus, since Mróz et al. (2015) have provided molecular and morphometric data indicating that L. longirostris is a synonym. In Switzerland it is recorded from Castanea sativa (Lampel & Meier 2003, as Schizodryobius longirostris). It may feed in ant shelters on roots of Fagus in summer (Heinze 1962). Monoecious holocyclic; oviparae and small apterous males occur in October. Throughout Europe, and east to the Caucasus. The male genitalia were described and illustrated by Wieczorek et al. (2012), and Mróz et al. (2015) provided redescriptions of both males and oviparae. [It would be advisable to have more proof of the synonymy of L. longirostris with L. pallipes, for example by making host transfers and by analysis of DNA at hypervariable loci.]

Lachnus pseudonudus Kanturski, Wieczorek & Junkiert Appearance in life unknown; BL of aptera 2.6-3.6 mm. On branches and shoots of Quercus macrolepis (= Q. ithaburensis spp. macrolepis) in Turkey (Kanturski et al. 2014b). Other morphs and life cycle are unknown.

Lachnus quercihabitans (Takahashi) Described from some alatae said to be collected (with immatures) on Quercus serrata in Korea, colour in life not noted, BL c. 6 mm. These alatae were described as having hyaline wings, and were placed by Takahashi (1924) in genus Dinolachnus (= Cinara). Some black apterae (BL c. 5.3 mm) from Q. acutissima in the same locality were described under this name by Okamoto & Takahashi (1927), and the much smaller sexual morphs were described by Paik (1972). This species is also recorded from Japan (Tsushima; Takahashi 1933e, as Cinara quercihabitans). However, there is considerable doubt about the identity of the type material, and therefore of the validity of the species, as alatae “agreeing exactly with the type specimens” were collected in the same locality on Abies holophylla (Okamoto & Takahashi 1927). These alatae were described as having long fine hairs, and are probably Cinara longipennis, in which case a new name is required for the Quercus-feeding species, which has apterae (and oviparae) with shorter, thicker hairs than those of C. longipennis. [This species is also reported to occur in China (on Quercus mongolica, Castanea mollissima and Castanopsis sp.; Binazzi & Remaudière 2010). The latter authors provide morphometric data for two apterae from China (BL 4.2-4.4 mm), which seem very similar to L. takahashii, and describe the alate form as having almost wholly black-brown forewings, which is at variance with the original description.]

Lachnus roboris (Linnaeus) Apterae are shining blackish brown (see influentialpoints.com/Gallery); BL 2.5-5.5 mm. Alatae have forewing membrane pigmented except for clear patches on either side of Rs, a clear band running obliquely across base of media and distal part of Cu_1a, and a clear triangle from Cu_1b to the wing-base. On twigs and small branches of Quercus spp. and also sometimes in large numbers on Castanea sativa in southern Europe. Throughout Europe, and also in North Africa, the Middle East and Central Asia. Monoecious holocyclic; apterous oviparae and large alate males occur in September-October. The male genitalia were described and illustrated by Wieczorek et al. (2012). Mróz et al. (2015) provided redescriptions of both sexual morphs. Michel (1942) studied its biology on oaks. Hille Ris Lambers (1956d) synonymised several Lachnus described from various Quercus species under the name roboris, considering variations in length and shape of body hairs and the small-bodied, long-legged forms which occur in summer in southern Europe to be environmentally induced. It nevertheless seems likely that L. roboris is a complex of species with different host plant associations and karyotypes, in which case the name for the southern European species on Castanea would be L. castaneae Hille Ris Lambers. Chromosome numbers of 2n=7, 8, 10, 11, 13, 16 and17 have been recorded, but some of these may apply to L. iliciphilus; a sample from Castanea in Portugal had 2n=10 (Blackman 1990).

Lachnus salicis Chakrabarti & Raha Colour in life is not recorded, probably brownish; BL of aptera 3.5-5.4 mm. Alatae have clear wings. In large dense colonies on stems of Salix babylonica and S. tetrasperma, attended by ants. Sexual morphs are not recorded; apterous viviparae and immatures may overwinter in bark crevices (Chakrabarti & Raha 1988). In Uttar Pradesh, India. Possibly it is a synonym of L. longirostrum, a very similar aphid described from Salix fragilis in Himachal Pradesh (David & Ghosh, in A.K. Ghosh 1982). It is likely that these willow-feeding aphids, with general appearance and biology very like Tuberlachnus salignus but lacking a dorsal tubercle, are wrongly placed in Lachnus.

Lachnus shiicola Sorin Apterae are shining brownish black; BL 3.7-4.0 mm. Forewing pigmentation of alata is similar to that of L. tropicalis. On Castanopsis sp. (?cuspidata) in Japan, feeding in scars of the bark of the trunk near its base, and on exposed roots of old trees, covered by ant shelters. Sexuales and life cycle are unknown. Closely related to L. tropicalis, but rather longer-haired and with shorter antennae and HT II (Sorin 1980).

Lachnus sorini Binazzi & Remaudière Apterae are shining blackish, with a large prominent rounded dorsal abdominal tubercle ; BL 3.2-5.0 mm. Alatae have forewing pigmentation similar to that of L. roboris. On Quercus spp. in Japan, and also collected on Castanopsis sp. Oviparae were found on Q. paucidentata (= sessilifolia) in late October-November (Binazzi & Remaudière 2007).

Lachnus swirskii Hille Ris Lambers Apterae are reddish brown, broadly pear-shaped; BL 3.1-4.4 mm. Alatae have forewing pigmentation closely resembling that of L. roboris. On branches of young Quercus spp. (aegilops, infectoria, ithaburensis), recorded from Israel and Turkey (Hille Ris Lambers 1954b; Canakçioglu 1967, 1975). Sexuales and life cycle unknown. Davatchi et al. (1957) described a form with longer hairs on the vertex, collected from Q. persica in Iran, as a subspecies, L. swirskii persicae.

Lachnus takahashii Sorin Apterae are brownish black, not shiny; BL 3.8-5.1 mm. Forewings of alata mainly infuscated, but with several clear patches of irregular shape. On Quercus acutissima in Japan, feeding in scars of the bark of old trees, usually on the basal part of the trunk in ant shelters (Sorin 1980). Sexuales and life cycle are unknown.

Lachnus tatakaensis Takahashi Apterae are brownish black, with many large black circular spots in rows on dorsal abdomen. Antennae black, legs reddish brown and black; BL 5.5-5.7 mm (Takahashi 1937b). Alatae have not been described. On trunk and branches of an unidentified Salix sp. at high altitude in Taiwan. The life cycle is unknown. Possibly this species belongs in either Tuberolachnus or Pterochloroides.

Lachnus tropicalis (van der Goot) (= japonicus Matsumura) Apterae are shining brownish black; BL 3.8-5.3 mm. Alatae have forewing wholly infuscated, except for a clear patch between pterostigma and Rs, and a clear band from base of media to distal part of Cu_1a. On twigs and stems of Quercus spp. Lithocarpus spp. and Castanea spp., in east and south-east Asia (India, east Siberia, China, Japan, Korea, Vietnam, Laos, Java, Malaya). Dense populations of a Lachnus that appears to be this species were also collected on twigs of Casuarina equisetifolia at high altitude in New Guinea (BMNH collection, leg. J. Szent-Ivany). Records from Castanopsis (= Shiia) are possibly all L. shiicola. Monoecious holocyclic in Japan, with apterous oviparae and alate males (Shinji 1927, Moritsu 1983); possibly anholocyclic in warmer regions. L. siniquercus Zhang, described from Q. liaodongensis ( ?liaotungensis Koidzumi) in China (Zhang & Zhong 1982) is closely related, and could be large specimens of tropicalis. The tropicalis/shinjii group perhaps involves a complex of species or races, paralleling that thought to occur in L. roboris. Das & Chakrabarti (1989) recorded a new parasitoid from L. tropicalis, in India. Sakata (1999) studied the effects of ants (Lasius niger) on population dynamics. 2n=12, 13 or 16 (Blackman 1986). [Muramoto (1987) reported chromosome numbers from 2n=14-38, but his results are difficult to interpret and may include polyploid cells and/or preparations of more than one species.]

Lachnus tuatayae Remaudière Apterae are grey with black hind tibiae; BL 2.1-2.8 mm. Alatae have forewing pigmentation similar to that of L. roboris. On Quercus persica in Turkey (Ozdemir et al. 2005; Binazzi & Remaudière 2010). Sexuales and life cycle are unknown.

Lachnus wichmanni Hille Ris Lambers Apterae are dull blackish with greyish waxy markings; Bl 3.3-4.6 mm. Sides of head and sections of the femora and tibiae are orange-brown. Alatae have the forewing membrane infuscated distally, and very thick bands of fuscous along Rs, media and Cu_1b. On trunk and branches of Hippophae rhamnoides, attended by ants. Recorded from Germany and Italy, and recently also from Georgia (Barjadze et al. 2010a). Monoecious holocyclic; apterous oviparae and small apterous males occur in August in southern Germany (Hille Ris Lambers 1956d).

Lachnus yunlongensis Zhang Known only from apterous viviparae (BL c. 4.6 mm) collected on Salix sp. in China (Zhang & Zhong 1985). Biology is unknown. The host recorded in the original description suggests that this species might have greater affinities with Tuberolachnus than with Lachnus. However, R. Chen et al. (2016) have now recorded it as occurring on Cyclobalanopsis (= Quercus) glauca, and their DNA analysis places it within Lachnini.

Laingia Theobald

Chaitophorinae: Siphini

One narrow-bodied species differing from Atheroides by the position of the siphunculi (on abdominal tergite 6, as opposed to abdominal tergite 5 in Atheroides) and by lack of a dorsal sclerotic carapace. Wieczorek (2010) provided a full account with redescriptions of all morphs.

Laingia psammae Theobald Apterae are dirty straw-coloured to greyish green (see influential points.com/Gallery); BL 1.6-2.8 mm. Alatae have dark transverse bars on the dorsal abdomen. Typically found in inflorescences of Ammophila arenaria in sand dunes, but also on Calamagrostis epigeios at more inland locations, and also recorded in Sweden from Elymus, Calamagrositis arundinacea and Deschampsia caespitosa (Heie 1982), and in Iberian peninsula on Carex acutiformis. Sometimes attended by ants. Widely distributed in Europe, and across Asia to east Siberia (Pashchenko 1988a). Monoecious holocyclic with apterous males and oviparae found in Spain in October (Nieto Nafría & Mier Durante 1998); oviparae have also been observed in May and June (Wieczorek 2010). The male genitalia were described and illustrated by Wieczorek et al. (2011). Zhu et al. (2017) found mitochondrial DNA differences between populations in eastern and western regions of northern China.

Lambersaphis Narzikulov

Chaitophorinae: Chaitophorini

A genus for one species on Populus, differing from Chaitophorus in the very short ANT PT, sparse short needle-like dorsal hairs, crater-like siphunculi without reticulate sculpturing, and semicircular cauda. The alatae have thickly fuscous-bordered wing veins like the North American Chaitophorus populicola (which also has a rounded cauda). Molecular work (T. Liu et al. 2022) now questions the validity of Lambersaphis as a genus separate from Chaitophorus.

Lambersaphis pruinosa (Narzikulov) Apterae are blackish brown (Qiao et al. 2003a); BL c. 1.5-1.7 mm. On young shoots of Populus pruinosa and P. diversifolia (= P. euphratica) in central Asia (Kazakhstan, Tadzhikistan; Aibasov 1971, Narzikulov 1957, 1961b), and now also recorded from Iran (Rezwani 2004), and from an unidentified species of Salix in China (Qiao et al. 2003a). Oviparae and alate males occur in October (BMNH collection, leg. M.N. Narzikulov).

Landisaphis Knowlton & Ma

Aphidinae: Macrosiphini

A genus for one little-known North American species with flattened, mostly spatulate dorsal hairs, a reticulate dorsal cuticle and clavate siphunculi. The sclerites of the spinal hairs on abdominal tergites 6-8 are developed into rugose conical processes (less developed in alatae).

Landisaphis davisi Knowlton & Ma (Fig.37a) Colour of apterae in life is unrecorded, BL 0.9-1.4 mm. Described from apterae collected on undersides of leaves of Chenopodium album, although there are subsequent records from Brassicaceae (Lepidium perfoliatum, Descurainia sophia; BMNH collection), and it has been reared on Capsella bursa-pastoris (A. Jensen, pers. comm). Previously only known from the state of Washington, but it has recently been found on a native Chenopodium sp. in a dry forest/desert habitat in New Mexico (A. Jensen, pers. comm). ..

Latgerina Remaudière

Calaphidinae: Calaphidini

One or two species on Alnus in Mexico, characterised by the apterae with frontal and marginal multidigitate hair-bearing processes. See Remaudière (1981) for all available information on the genus.

Latgerina orizabaensis Remaudière (fig. 27F,G) Apterae are flattened dorsoventrally, pale yellow with black antennae; BL 1.6-2.2 mm. Alatae have black head, thorax and appendages and black dorsal abdominal markings. On undersides of leaves, especially on young branches, of Alnus acuminata ssp. arguta in Mexico (Remaudière 1981). A more pigmented form with paler antennae collected on A. firmifolia was described as a subspecies, L. orizabaensis mexicana; the host plant was given in the original description as A acuminata, but was corrected in Remaudière & Muños Viveros (1985a, p.78). Monoecious holocyclic; oviparae and alate males were produced by both subspecies in September.

Lehrius Gredina

Aphidinae: Macrosiphini

One species in east Siberia with well-developed antennal tubercles and flangeless siphunculi, seemingly close to Jacksonia but with a cauda more typical of a moss-feeding aphid, although it was not found on mosses.

Lehrius papillicaudus Gredina Apterae are yellowish green; BL c.1.2 mm. Alatae are unknown. On Elsholtzia pseudocristata (= E. ciliata) in east Siberia. To judge from its description this species could be Myzus isodonis (Takahashi).

Lepidaphis Kadyrbekov, Renxin & Shao

Aphidinae: Macrosiphini

Two eastern palaearctic species on Lepidium in desert regions, related to Brevicoryne and Brachycolus but with numerous hairs on the front of the head and abdominal tergite 8, and a short antennal terminal process (Kadyrbekov et al. 2002). Life cycles are unknown.

Lepidaphis deformans (Nevsky) Apterae are greenish, with slight wax film; BL 1.7-2.2 mm. The single known alata has 12-14 secondary rhinaria on ANT III. In leaf galls on Lepidium spp. in Uzbekistan, Turkmenistan, southern Kazakhstan and the Xinjiang-Uygur region of western China (Kadyrbekov et al. 2002, 2017a).

Lepidaphis terricola Kadyrbekov, Renxin & Shao Apterae are greenish, with slight grey wax film; BL 1.9-2.1 mm. On roots of Lepidium spp. in south-east Kazakhstan and the Xinjiang-Uygur region of western China (Kadyrbekov et al. 2002).

Linaphis Zhang

Aphidinae: Macrosiphini

One species in China with most of the characters of a Lipaphis, and possibly belonging in that genus, but then one would expect the more usual host to be a crucifer.

Linaphis lini Zhang Apterae are grass-green; BL c. 1.8 mm. On Linum usitatissimum in Yinchuan Province, China. Populations built up to damaging levels in July. The life cycle is unrecorded.

Linosiphon Börner

Aphidinae: Macrosiphini

Three palaearctic and one nearctic species resembling Illinoia in the tendency for siphunculi to be swollen proximal to the subapical, reticulated zone, but the degree of swelling varies considerably within species. Heie (1994) provided an account of two species in Europe.

Linosiphon asperulophagum Holman Apterae are green or pinkish with shiny brown-black dorsum, head and basal antennal joints, and dark siphunculi; BL 1.7-2.3 mm. Alatae are as yet undescribed; a specimen in the BMNH collection (leg. H.L.G. Stroyan) has ANT III with 5-6 small rhinaria, and dorsal abdomen with incomplete dark cross bars and large marginal and postsiphuncular sclerites. On undersides of leaves and upper parts of stems of Asperula odorata in Czech Republic. Records from other localities and from Galium spp. need confirmation as all other specimens in the BMNH collection identified as L. asperulophagum, including those recorded from Hungary on Asperula by Szelegiewicz (1966a), are referable to L. galii (q.v.). Oviparae and alate males occur in October (original description).

Linosiphon galii Mamontova Plate 26a Apterae are shiny green or brown with variably developed black pigmentation of dorsum, and having dusky brown siphunculi with dark apices; BL 1.7-2.0 mm. Alatae have incomplete dark dorsal abdominal cross bands and large marginal and postsiphuncular sclerites. On undersides of leaves of Galium spp. and Asperula odorata. This species has been confused with L. asperulophagum but has a shorter R IV+V and longer HT II, the ratio between these two providing a reliable discriminant. In Ukraine, Czech Republic, Greece, Hungary, Poland and Germany.

Linosiphon galiophagum (Wimshurst) Apterae are shiny green, their siphunculi pale with dark apices; BL 1.7-2.5 mm. Alatae have dark marginal abdominal and intersegmental pleural sclerites but no dark cross bands. On Galium spp., living on undersides of leaves along veins, and on young shoots. In Europe as far south as Corsica, and eastward to west Siberia. Oviparae occur in October (Heie 1994).

Linosiphon sanguinarium (Hottes & Frison) Apterae have pearly white to yellowish head and posterior part of abdomen including siphunculi and cauda, the thorax and anterior abdomen being bright shining ruby-red (presumably due to sap of host); BL c. 1.6 mm. On undersides of leaves, feeding singly near larger veins, of Sanguinaria canadensis. In north-eastern and north-central USA. Monoecious holocyclic; a fundatrix was collected in early May (original description), but sexual morphs have not been described.

Liosomaphis Walker

Aphidinae: Macrosiphini

Four species resembling Cavariella but without a supracaudal process, and typically associated with Berberis.

Liosomaphis atra Hille Ris Lambers Apterae are dark purplish brown, dirty greenish in centre of dorsal abdomen, with pale legs and antennae, siphunculi dark on distal half; BL c.1.3-1.6 mm. Alatae have a brown dorsal abdominal patch (L.K. Ghosh & Pramanik 1976). On Berberis spp. in central and east Asia (Iran, Kazakhstan, Turkmenistan, Afghanistan, India, Pakistan, China). Mehrparvar (2016) provided a redescription. 2n=17 (male? – Kurl & Chauhan 1988)

Liosomaphis berberidis (Kaltenbach) (= Liosomaphis turanica Narzikulov) Plate 16a (Fig.17e,f) Apterae are yellow to yellow-green, or pinkish to orange-red (two colour forms), slightly wax-powdered (see influentialpoints.com/Gallery); BL 1.1-2.3 mm. Alatae have a dark head, thorax and antennae, but little or no dark dorsal abdominal pigmentation. On undersides of leaves of Berberis and Mahonia. Throughout Europe, eastward to India (BNMH collection), and introduced to Australia, New Zealand, North America, and recently recorded from Argentina (Orlando et al. 2018). A record from Algeria (Laamari et al. 2013) requires additional confirmation because of the anomalous host (Achillea). Oviparae and alate males appear in UK in late September-November. 2n=18.

Liosomaphis himalayensis A.N. Basu Apterae are shiny pale yellowish, with green to yellowish brown markings on thorax and abdomen; BL 1.7-2.5 mm. Alatae have the dorsal abdomen with variable brownish sclerotisation (A.K. Ghosh 1969). On undersides of leaves of Berberis spp. in India, Nepal, China, and also recorded from Java (Noordam 2004). A record from Turkey (Şenol et al. 2017) requires additional confirmation. Alate males were found on snow in December at an altitude of 2490m (L.K. Ghosh & Pramanik 1976). This species could possibly be synonymous with Wahlgreniella viburni (Takahashi). 2n=18.

Liosomaphis ornata Miyazaki Apterae are reddish brown or dark brown, mottled with dull green; BL 1.4-2.1 mm. Alatae have a large dark dorsal abdominal patch. On leaves and young growth of Berberis spp. Japan, China (Zhang et al. 1999, as Rhopalosiphoninus yuzhongensis), east Siberia and India (Himachal Pradesh; Nadda & Joshi 2015). Fundatrices occur in Japan in early May (original description).

Lipamyzodes Heinze

Aphidinae: Macrosiphini

A genus for one species superficially resembling Lipaphis, but perhaps more closely related to Myzus.

Lipamyzodes matthiolae Doncaster Plate 12e Apterae are mid to dark green or blue-green, with the head pale grey and pulverulent; BL 1.9-2.5 mm. Alatae have a solid black dorsal abdominal patch and black siphunculi. Described from Arabis and Matthiola, and subsequently collected on Galium (BMNH collection, leg. J.P. Doncaster). In England and Wales, and alatae have been found on Glaucium grandiflorum in Lebanon (BMNH collection, leg. D. Hille Ris Lambers) and on Cardaria draba in USA (Washington; BMNH collection, leg. L. Fox). There were no records of colonies on host plants from outside Britain until it was found in 2009 in Iran on an Achillea sp. (M. Mehrparvar, pers. comm.). The apparently remarkable host range needs conformation. The life cycle and sexual morphs are unknown.

Lipaphis Mordvilko

Aphidinae: Macrosiphini

About 12 mostly western palaearctic species associated with Brassicaceae, related to Brevicoryne and characterised by weakly developed antennal tubercles, apterae with rather short antennae which are almost always without secondary rhinaria, a sclerotic but often weakly pigmented tergum, and weakly swollen siphunculi. Subgenus Lipaphidiella is distinguished by the presence of a conical, rather scabrous, supracaudal process. Accounts are available by Doncaster (1954a), Heinze (1960), Heie (1992) and Blackman (2010).

Lipaphis alliariae Müller Apterae are dark blue-green to almost black (see influentialpoints.com/Gallery); BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III c. 22, IV 5-6. On Alliaria petiolata in Europe (France, Sweden, Finland, Poland, Germany – and now recorded in England; R. Dransfield, pers. com.). Monoecious holocyclic with oviparae and apterous males in October (Müller 1955a, as erysimi ssp. alliariae).

Lipaphis cochleariae Jacob Apterae are dull olive green with variably-developed brown patches; BL 1.2-2.1 mm. Alatae have secondary rhinaria distributed III 19-30, IV 5-11, V 0-4. On Cochlearia officinalis in the intertidal zone, on rosettes of young plants, or in flower-heads. Only known from UK (England, Scotland, Wales). Monoecious holocyclic, with oviparae and apterous males in October (Stroyan, 1957b).

Lipaphis erysimi (Kaltenbach) Apterae are yellowish green, dirty green or brownish; BL 1.5-2.3 mm. Alatae have secondary rhinaria distributed III 9-32, IV 2-10, V 0-3. On various Brassicaceae (Arabis, Capsella, Coronopus, Erysimum, Isatis, Lepidium, Matthiola, Sinapis, Sisymbrium, Thlaspi, etc.), but not usually on field Brassica crops. In northern Europe, and probably in western Siberia and Central Asia, but its distribution needs clarification due to past confusion with L. pseudobrassicae. Monoecious holocyclic with apterous males. The name has been commonly applied to the world-wide crucifer pest, L. pseudobrassicae (q.v.), but they differ in karyotype, and Ronquist & Åhman (1990) showed in laboratory trials that L. erysimi performed poorly in comparison with L. pseudobrassicae on Brassica oilseed crop plants. 2n=10.

Lipaphis fritzmuelleri Börner Apterae are dark green; BL 1.3-1.8 mm. Alatae have secondary rhinaria distributed III 11-16, IV 0. On Sisybrium spp., occurring on flowers in spring and on lower leaves in summer. In Sweden, Germany, Austria, west Russia and Iran. Also now recorded from Kazakhstan, where it is reported to occur on Cardamine impatiens as well as S. loeselii (Kadyrbekov 2014e). Monoecious holocyclic with oviparae and small apterous males on Sisymbrium in October. 2n=10.

Lipaphis (Lipaphidiella) jungarica Kadyrbekov, Renxin & Shao Apterae are greenish; BL c. 1.7 mm. Apterae have 3-5 secondary rhinaria on ANT III (based on 1 specimen), alatae have them distributed III 7-12, IV 0-3. On upper sides of leaves of Syrenia siliculosa (= Erysimum siliculosum) in western China (Kadyrbekov et al. 2002). Also recorded from Hypecoum erectum (Papaveraceae), but these could be vagrant alatae (only 1 aptera was found, presumably on Syrenia).

Lipaphis (Lipaphidiella) lepidii (Nevsky) (Fig.37b) Apterae are pale green; BL 1.2-1.6 mm. Alatae have secondary rhinaria distributed III 34-48, IV (0-)1-6, V 0-1. On undersides of leaves, stems and flower-stalks of Lepidium spp. in Greece, Middle East and Central Asia, eastward to Pakistan. In Iran it is also recorded from Sisymbrium officinale (Mokhtari et al. 2012). A Roumanian population with longer siphunculi is regarded as a subspecies, L. lepidii ssp. lepidiicardariae (Knechtel & Manolache 1944, as Myzaphis).

Lipaphis pseudobrassicae (Davis) Mustard Aphid, Turnip Aphid, False Cabbage Aphid Plate 12d Apterae are yellowish green, grey-green or olive-green, with a white wax bloom, which in humid conditions may become a dense mealy coat (see aphids of Karnataka website); BL 1.4-2.4 mm. Alatae have secondary rhinaria distributed III 15-30, IV 3-13, V 0-3. On many genera and species of Brassicaceae, inclunding Barbarea, Brassica, Capsella, Iberis, Raphanus and Rorippa. An important world-wide pest of brassica crops (Blackman & Eastop 2000). Monoecious holocyclic in Japan, with apterous males and oviparae (Kawada & Murai 1979, Sasaki 2021; as L. erysimi), and sexual morphs have also been reported from India, China and New Zealand, but it is predominantly anholocyclic in warm climates. Agarwala et al. (2009) compared its morphology and performance on three different species of Brassicaceae in India. 2n = 8 or 9 (anholocyclic populations in most parts of the world have 2n=9).

Lipaphis rossi Börner Apterae are dark grey green with a slight waxy bloom, with a dark grey brown head, broad dark grey brown bars across the dorsum, and large marginal sclerites; BL 1.2-1.6 mm. Alatae have secondary rhinaria distributed III 27-53, IV 10-26, V 2-12. Monoecious holocyclic on Arabis hirsuta in UK, with oviparae and apterous males in October (Prior 1971). Infested plants have stunted flower stems and deformed inflorescences. Also recorded from Netherlands, Denmark, Sweden and Germany (on Arabidopsis thaliana; Heinze 1960). A form on Coringia orientalis in Ukraine was described as a subspecies, L. rossi ssp. coringiae Bozhko, and populations agreeing with this subspecies have been found in Finland on Galium mollugo (Heikinheimo 1984), and in eastern Kazakhstan on Conringia planisiliqua (Kadyrbekov 2009a). However, there is a possibility that this species will prove to be synonymous with L. turritella, as the only substantive difference seems to be its smaller body size (V.F. Eastop, unpublished data).

Lipaphis (Lipaphidiella) ruderalis Börner ( = Lipaphis berteroaella Mamontova) (Fig.37c) Apterae are greyish green; BL 1.8-2.2 mm. Alatae have secondary rhinaria distributed III 35-40, IV 5-7. On undersides of leaves, stems and flower-stalks of Lepidium spp. in eastern Europe, and also in China (Tao 1999). L. berteroaella Mamontova, described from Berteroa incana and Lepidium ruderale in Ukraine, appears to be a synonym.

Lipaphis sisymbrii Bozhko Apterae are yellow-green; BL c. 1.7-1.8 mm. Alatae have secondary rhinaria distributed III 26-30, IV 8. On Sisymbrium polymorphum in Ukraine, and also reported on the same host as well as on S. loeselii in Kazakhstan (Kadyrbekov 2009a, 2017a).

Lipaphis turritella (Wahlgren) Plate 12c Apterae are greenish yellow to yellowish brown, dusted with white wax; BL 1.5-2.3 mm. Alatae have secondary rhinaria distributed III 40-53, IV 10-24, V 2-12. On Arabis (=Turritis) glabra, causing deformation of inflorescences. Also recorded from Arabis pendula (Kadyrbekov 2017a) and Erysimum cheiranthoides (Ivanoskaya 1977). In Europe, west Siberia and Kazakhstan.

Lipaphis unguibrevis Zhang Colour of apterae in life is unrecorded; BL c.1.9 mm. Alatae have secondary rhinaria distributed III 5-8, IV 0-2. On Brassica sp. at 3,800 m in Tibet.

Lithoaphis Takahashi

Hormaphidinae: Nipponaphidini

One little-known east Asian species on Fagaceae. [A second species described in this genus by Takahashi, Lithoaphis shiiae Takahashi has been transferred to Metaniponaphis by Aoki et al. (2021).]

Lithoaphis lithocarpi (Takahashi) Apterae are black, rather densely covered with white secretion, almost circular, and dorsally flattened; BL c. 0.75 mm. On undersides of leaves of Lithocarpus sp. in Taiwan (Takahashi 1929; as Astegopteryx lithocarpi). Other morphs and life cycle are unknown.

Lizerius Blanchard

Lizeriinae

A South American genus with primitive features and affinities to the African genus Paoliella, but with no clear host relationships. Four species are known to be tree-dwelling, two on Combretaceae and three on Lauraceae, but several other species are only known from trapped alatae. Quednau (2010) reviewed the genus and provided keys to apterae and alatae and illustrations of most of the known morphs.

Lizerius acunai (Holman) Apterae are dark brown, with blue-grey wax especially around bases of dorsal processes; BL 1.3-1.8 mm. On undersides of leaves of Nectandra reticularis in Cuba (Holman 1974, as Neolizerius). Other morphs and biology unknown.

Lizerius (Paralizerius) brasiliensis Quednau Apterae are yellow-green, with long marginal processes and eyes reduced to triommatidia; BL 1.2-2.1 mm. Alatae have dark head, pterothorax, antennae and legs. On young shoots and suckers of Terminalia australis in Brazil and Uruguay. Apterous males and oviparae were collected in January (Quednau 1974, and BMNH collection, leg. V.F. Eastop).

Lizerius (Paralizerius) cermelii Quednau Plate 6b Apterae are whitish with green head and darker antennae, legs and posterior abdominal processes; BL 1.8-2.4 mm. Described from a large colony of mainly alatae on Bougainvillea sp. in Brazil. Alatae have been trapped in Argentina and Venezuela, and it is also recorded from Belize, Panama and Costa Rica (Villalobos Muller et al. 2010), although records from Ocotea are possibly in error and require confirmation. Oviparae were found (on Bougainvillea) in Brazil (Curitiba) in early November (Quednau 1974).

Lizerius jorgei Cunha & Sousa-Silva Apterae are bright yellow with small greenish stripes on thorax and anterior abdomen, and eyes reduced to triommatidia; BL 1.2-2.1 mm. Alatae are also bright yellow with a dark brown thorax. On Terminalia brasiliensis in São Paulo, Brazil (Cunha & Sousa-Silva 2019); also found on Persea americana – possibly a less-favoured host. An alate male was collected (on Persea) in January.

Lizerius ocoteae Blanchard Plate 7c, d Apterae are dark olive to brownish black, thickly coated with white wax; BL 1.6-2.0 mm. Alatae are brownish black with a sparse covering of wax meal. In dense colonies on young stems and leaves of Ocotea acutifolia in Argentina (Blanchard 1923), and Uruguay (BMNH collection, leg. V.F. Eastop). Also recorded from Ocotea porphyria (Blanchard 1944). Alatae have been trapped in Brazil. Blanchard (1923) found oviparae and alate males within the colony on Ocotea, but gave no collection date. The genitalia (of an ?apterous male) were described and illustrated by Wieczorek et al. (2011).

Lizerius pichurim Quednau Adult apterae in life unknown; BL of alata 1.3-1.9 mm. On Nectandra picurim and Nectandra sp. (alatae and immatures) in Venezuela, and two alatae have been trapped in Argentina (Quednau 2010). A form collected in the Caribbean (host unknown) was described as a subspecies, L. pichurim carabicus Quednau.

Lizerius pustulatus Quednau Colour in life unknown; BL of aptera c. 1.4-1.5 mm. On Ocotea sp. near glomerata and Nectandra sp. in Venezuela, and alatae have been trapped in Brazil (Quednau 2010).

Lizerius tuberculatus (Blanchard) Apterae are brownish black; BL 1.2-1.5 mm. Alatae are also brownish black, and weakly pruinose. On young growth of Nectandra sp. (Blanchard 1939) in Argentina, and also collected from Ocotea ?glomerata in Venezuela (BMNH collection, leg. M. Cermeli), and from Eugenia sp. in Brazil. Alatae have been trapped in Brazil and Jamaica. Sexuales and life cycle are unknown.

Longicaudinus Hille Ris Lambers

Aphidinae: Macrosiphini

One east Asian species resembling Longicaudus but with differences in antennal morphology and first tarsal chaetotaxy (Hille Ris Lambers 1965).

Longicaudinus corydalisicola (Tao) Apterae are pinkish grey with pale appendages, a pale spinal stripe and glassy white wax secretion on sides and end of abdomen; BL 1.7-2.3 mm. Immatures are pale greenish yellow, and alatae have a dark dorsal abdominal central patch. On Corydalis spp. in Japan and Taiwan, introduced to Great Britain and Ireland (Greenslade et al. 2024).

Longicaudus van der Goot

Aphidinae: Macrosiphini

A rather distinctive genus of about 7 species in Europe and Asia, typically host-alternating between Rosa and Thalictrum. They generally have short siphunculi (completely absent in fundatrices), a long hairy cauda and a very long ANT III which in the alata bears numerous tuberculate rhinaria. The most recent revisions are by Chakrabarti & Banerjee (1991a), Remaudière (1993a), and (for China) Qiao et al. (2006e).

Longicaudus cornutus Chakrabarti & Banerjee Colour of apterae in life is unrecorded; BL c.2.5 mm. On Thalictrum sp. in Uttar Pradesh, India.

Longicaudus dunlopi Hille Ris Lambers Apterae are creamy white; BL 1.7-2.4 mm on Rosa, 2.8-3.0 mm on Thalictrum. Heteroecious holocyclic with Rosa canina as primary host, migrating to Thalictrum flavum in marshy, often flooded situations (Remaudière 1993a). In Europe (France, Netherlands), and apterae collected on Th. chelidonii in India are possibly this species (Chakrabarti & Raychaudhuri 1975), as also may be certain Japanese populations previously described as L. trirhodus ssp. japonicus Hille Ris Lambers (Remaudière 1993a).

Longicaudus himalayensis Hille Ris Lambers Described from two alatae collected on ?Quercus, which according to Chakrabarti & Banerjee (1991a) were gynoparae that originated from Thalictrum sp. An aptera from Rosa described as himalayensis (David et al. 1971a) is included in the Rosa key on the basis of David et al.’s description, although it seems to be a different species with shorter siphunculi and higher ratio of R IV+V to HT II (see Remaudière 1993). L. himalayensis is also included in the Thalictrum key on the basis of information in the key by Chakrabarti & Banerjee (1991a), who reared one aptera from Thalictrum. There is however no proper description of the aptera of himalayensis, either from Rosa or Thalictrum.

Longicaudus kumauni Chakrabarti & Banerjee Colour of apterae in life is unrecorded; BL c.1.2 mm. On Thalictrum sp. in Uttar Pradesh, India.

Longicaudus naumanni Remaudière (Fig.11c) Apterae are very pale, presumably whitish green, BL 1.6-2.7 mm. Heteroecious holocyclic between Rosa macrophylla and Aquilegia pubiflora in Pakistan. The fundatrix on Rosa gives rise directly to emigrant alatae, which have 60-90 secondary rhinaria on a very long ANT III (Remaudière 1993a). Sexual morphs are unknown.

Longicaudus netuba (Zhang, Chen, Zhong & Li ) Apterae (fundatrices) are yellowish green; BL (fundatrices) 1.6-1.7 mm (only fundatrices and immatures are described). On young shoots of Rosa omeiensis in Xinjiang, China (Zhang 1999, as Netubusaphis netuba). Possibly this is a synonym of L. naumanni.

Longicaudus trirhodus (Walker) Plate 14h (Fig.11b) Apterae are yellowish green to pale apple-green, with variably expressed slightly darker green cross-bands, lightly wax-dusted (see influentialpoints.com/Gallery); BL 1.6-2.7 mm. Alatae have broad dark transverse sclerites on abdominal tergites III-VI, partly merged to form an irregular-shaped dorsal abdominal patch. In small colonies on leaves and blossom buds of Rosa spp. in spring. Heteroecious holocyclic, migrating for the summer to Aquilegia and Thalictrum spp. In Europe, Asia and North America. A monoecious holocyclic population on Thalictrum majus in Iran, with oviparae differing morphologically from those on Rosa, was described as a subspecies, L. trirhodus ssp. iranicus, by Remaudière (1993a). 2n=12.

Longisiphoniella Chakrabarti, Saha & Mandal

Aphidinae: Macrosiphini

One species in India with a short broad cauda and, considering its subterranean habit, remarkably long siphunculi. [Spinaphis L.K. Ghosh may be an older name for the genus, as the type and only known species Spinaphis multisetosa, described from apterae on an unknown host, is similar in many respects.]

Longisiphoniella subterranea Chakrabarti, Saha & Mandal Appearance of apterae in life is not recorded, BL 2.0-2.5 mm. Collected from roots of various plants (Artemisia sp., Cynoglossum glochidiatum, Conyza bonariensis, Rumex sp.) in west Himalaya. It seems unlikely that all these are true host plants. Alatae have large, protruberant secondary rhinaria distributed III 23-30, IV 0-6, V 0-1.

Longistigma Wilson

Lachninae: Lachnini

Three species have been described in this genus, one in North America and the others in east Asia. Very large, bark-feeding aphids, characterised in the alatae by the elongate pterostigma extending around the tip of the forewing. The differences between the Asian species are not clearly defined.

Longistigma caryae (Harris) Apterae are pale brownish-grey, slightly pruinose, with conspicuous rows of dark dorsal spots, and dark siphunculi (see influentialpoints.com/Gallery); BL 5.1-7.8 mm. Alatae have similarly conspicuous black markings. On the bark of numerous tree species in North America. Bissell (1978) listed 292 host records from trees in 24 genera and 16 families. Monoecious holocyclic, with alate males, in northern USA (Washington D.C.; Wilson 1909), anholocyclic in southern states (Bissell 1978). Marked fluctuations in population size occur, this species being very abundant in some years (Tissot 1944).

Longistigma liquidambarus (Takahashi) Apterae black or brownish black, lightly dusted with wax powder; BL c.7.0-7.7 mm. On Liquidambar formosana in Taiwan and Japan (Kawada & Yamashita 1992), feeding on the bark. Anholocyclic; no sexual morphs are known. Tsumuki et al. (1993) studied its tolerance of low temperature. [Possibly also in India, as A.K. Ghosh (1982) provided a detailed description of apterae and an alata of a Longistigma sp. collected (with immatures) on Berchemia floribunda in Meghalaya. These specimens resembled L. liquidambarus in most respects, but were much smaller (BL of apterae 4.5-5.3 mm). Given the polyphagous tendencies shown by members of this genus, it seems likely that these were indeed L. liquidambarus, and that their small size was due to feeding on a less favourable host.]

Longistigma xizangensis Zhang Apterae of BL 6.5-7.1 mm, colour in life unknown. Recorded from several genera of trees (Salix, Phoebe, Populus, Prunus, Quercus) in Tibet (Zhang & Zhong 1981b).

Loniceraphis Narzikulov

Aphidinae: Macrosiphini

One little-known species overwintering on Lonicera in Central Asia.

Loniceraphis paradoxa Narzikulov Apterae (fundatrices) are whitish green; BL 3.4-3.9 mm. In spring colonies on undersides of leaves and growing points of Lonicera spp., attended by ants. In Central Asia, and recorded also from Turkey (Güçlü 2015). Migrating in the second generation to an unknown secondary host. Gynoparae, oviparae and alate males occur in early October (Mukhamediev & Akhmedov 1982). Trichosiphaphis (Xenomyzus) foliotus (Shaposhnikov in Juchnevitch; nomen nudum) is a synonym (Stekolshchikov & Buga 2006b).