Revision of the aphid subgenus Ramitrichophorus Hille Ris Lambers of genus Macrosiphoniella (Hemiptera, Homoptera: Aphididae, Macrosiphini), with redescriptions of all known species and identification keys Author Stekolshchikov, Andrey V. Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia. Author Buga, Sergey V. 0000-0002-1140-9042 Zoological Department, Belarusian State University, pr. Nezavisimosti 4, Minsk 220030, Republic of Belarus. sergey. buga @ gmail. com; https: // orcid. org / 0000 - 0002 - 1140 - 9042 sergey.buga@gmail.com text Zootaxa 2022 2022-09-12 5183 1 293 342 http://zoobank.org/d6fee369-60f2-483f-8867-01396cda2932 journal article 140860 10.11646/zootaxa.5183.1.23 aef90821-b086-4773-b589-099432880afe 1175-5326 7075519 AD0BED04-C49A-4068-8D3A-22DE8B172744 Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) ( Figs. 3 , 42–63 , Tabl. 4–5 ) Material. Syntype : Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) —2 apterous viviparous female, „ Russia [mistake, right— Ukraine ], Helichrysum arenarium, Smiev , 3.viii.1947 , Leg. Bozhko, Ramitrichophorus medvedevi Bozhko, Det. Bozhko ” (from collections of NHM ); Paratypes : Macrosiphoniella ( Ramitrichophorus ) nasti Szelegiewicz, 1958 —4 apterous viviparous female, „ Polonia , Bydgoszcz, 30.vii.1956 , Helichrysum arenarium , leg. Szelegiewicz, Paratypus , Macrosiphoniella ( Ramitrichophorus ) nasti sp. n. , det. Szelegiewicz, 1957” (from collections of MNHN and NHM ). Additional materials : 2 apterous viviparous females, 03.vii.1947 , Ukraine , Helichrysum arenarium (from collections of MNHN ); 4 apterous viviparous females, 06.viii.1957 , Poland , Kuyavian-Pomeranian Voivodeship , Bydgoszcz, Helichrysum arenarium (from collections of MNHN ); 1 apterous viviparous female, 1 male and 16 oviparous female, 20.ix.1962 , Poland , Kuyavian-Pomeranian Voivodeship , Bydgoszcz, Helichrysum arenarium (from collections of IE BC CAS ); 3 apterous viviparous females, 25.vii.1965 , Poland , Mazovian Voivodeship , Otwock County, Otwock, Helichrysum arenarium (from collections of IE BC CAS ); 37 apterous viviparous females and 16 alate viviparous female, 26.vii.1969 , Russia , Moscow Province , Serpukhovsky District, Luzhki Vill., Helichrysum arenarium (from collections of IE BC CAS ). Also used data from Bozhko (1950 , 1957 , 1976 ), and Szelegiewicz (1958). Etymology. The species was named in honor of the famous Soviet Ukrainian entomologist Sergei Ivanovich Medvedev. FIGURE 42. Apterous viviparous female of Metopeurum medvedevi Bozhko, 1950 (Syntype) . FIGURE 43. Apterous viviparous female of Macrosiphoniella ( Ramitrichophorus ) nasti Szelegiewicz, 1958 (Paratype) . TABLE 4. Biometric data for apterous and alate viviparous females of Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 )

Apterous viviparous females	Alate viviparous females
Number of samples / specimens	7/53 + data of Bozhko (1950 , 1957 , 1976 ) and Szelegiewicz (1958)	1/16 + data of Szelegiewicz (1958)
Length of body	1416–2600	1462–1900
Length of antenna	1291–2150	1780–1978
Length of antenna / length of body	0.70–1.24	1.03–1.24

.....Continued on the next page TABLE 4 (Continued)

Apterous viviparous females	Alate viviparous females
length		437–653	487–558
HFem	length / length of body			0.27–0.41	0.30–0.35
length / W Head		1.01–1.49	1.21–1.34
HTib	length		827–1084	914–1066
	length / length of body			0.45–0.67	0.58–0.67
W Head		382–488	379–419
Depth of frontal sinus / width between antennal bases				0.04–0.12	0.06–0.16
occipital	length	61–106	46–63
	on	length / Ant3 AD	2.71–4.42	2.00–2.67
head	frontal	length	59–106	53–71
length / Ant3 AD		2.67–4.53	2.32–3.00
on Ant1		number	4–8	5–7
		length	41–89	46–66
on Ant3	number	8–18	10–15
	length	26–48	23–33
	length / Ant3 AD		1.13–2.12	0.90–1.44
on AntLB		number	3–4	3–4
		length / AntLB AD	0.77–1.67	1.00–1.50
length of ventral seta on HTr / BD HFem			1.12–1.59	1.00–1.53
on HFem	length of longest	dorsal	41–89	43–63
		ventral	50–81	41–66
		dorso-apical	23–46	25–30
Setae	on longest dorsal HTib longest dorsal / mid-diameter of HTib			44–78	38–51
				1.07–2.08	1.33–1.81
number on 2HT		dorsal	0–3	1–2
		ventral	4–6	3–7
number of spinal and marginal		17–30	16–22
spinal	length	53–111	53–71
	on TerIII	length / Ant3 AD	2.00–4.63	2.10–3.13
marginal		length	61–108	56–78
		length / Ant3 AD	2.67–4.63	2.44–3.38
ventral	length	35–104	58–76
	length / Ant3 AD	1.33–4.47	2.56–3.33
number on TerVI between siphunculi			4–7	4–6
number		5–8	5–7
on TerVIII length			56–111	68–89
length / Ant3 AD		2.47–5.00	2.84–3.67
number on subgenital on anterior half			2–5	2–4
plate	along the hind margin	6–14	7–11
AntLB	99–139	106–156
AntL	PT	438–569	508–602
PT / AntLB		3.71–5.50	3.27–5.64

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Apterous viviparous females	Alate viviparous females
URS	number of accessory setae	4–7	4–7
	length	214–287	228–245
	W Head	0.50–0.63	0.56–0.65
	length / 2HT	1.62–2.00	1.64–1.79
	AntLB	1.80–2.45	1.58–2.14
2HT	length	119–151	129–145
	maximum width length / W Head	3.78–5.18 0.27–0.36	4.43–5.50 0.32–0.37
	AntLB	1.02–1.28	0.89–1.26
SIPH	length	243–400	248–350
	length / length of body	0.10–0.25	0.15–0.18
	length / BW SIPH	2.89–5.67	3.03–4.71
	length / MW SIPH	4.67–7.63	5.69–6.94
	length / Ant3	0.59–0.84	0.50–0.59
CAUD	length	139–216	147–170
	length / BW CAUD	1.09–1.70	1.20–1.56
	number of setae	6–9	6–9
Length of SIPH / length of CAUD		1.37–2.22	1.49–2.06

FIGURE 44. Apterous viviparous female of Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) (26.vii.1969, Russia, Moscow Province, Serpukhovsky District, Luzhki Vill.). FIGURES 45–50. Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) , apterous viviparous female. 45, 3rd antennal segment; 46, head; 47, ultimate rostral segment; 48, hind tarsus; 49, siphunculus; 50, cauda. FIGURE 51. Alate viviparous female of Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) (26.vii.1969, Russia, Moscow Province, Serpukhovsky District, Luzhki Vill.). TABLE 5. Biometric data for oviparous females and males of Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 )

Males	Oviparous females
Number of samples / specimens	1/1	1/16
Length of body	1325	1573–2025
Length of antenna	1338–1361	1533–1791
Length of antenna / length of body	1.01–1.03	0.85–0.99
length	447	497–558
HFem length / length of body	0.34	0.27–0.32
length / W Head	1.09	1.01–1.24
length HTib length / length of body	746–761 0.56–0.57	802–929 0.44–0.52
W Head	410	436–493
Depth of frontal sinus / width between antennal bases	0.02	0.02–0.06

......Continued on the next page TABLE 5. (Continued)

Males	Oviparous females
Se-tae	length occipital on length / Ant3 AD	63	71–91
		2.78–2.94	2.94–4.00
	head length frontal length / Ant3 AD	68	76–101
		3.00–3.18	3.13–4.38
	number on Ant1 length	6	5–7
		43–53	38–66
	number on length Ant3 length / Ant3 AD	13–14	10–16
		30	30–48
		1.33–1.41	1.38–2.00
	number on AntLB length / AntLB AD	3–4	3–5
		1.23–1.38	1.13–1.60
	length of ventral seta on HTr / BD HFem	0.97–1.31	0.91–1.73
	dorsal on length of ventral HFem longest dorso-apical	43–46	49–71
		46–53	43–66
		25–33	29–38
	on longest dorsal HTib longest dorsal / mid-diameter of HTib	41	53–68
		1.07–1.10	1.22–1.59
	dorsal number on 2HT ventral	2–3	2
		5	4–5
	number of spinal and marginal	22	21–30
	length spinal length / Ant3 AD on TerIII length marginal length / Ant3 AD	66	81–99
		2.89–3.06	3.41–4.50
		68	81–91
		3.18	3.08–4.24
	length ventral length / Ant3 AD	56	66–84
		2.44–2.59	2.74–4.00
	number on TerVI between siphunculi	8	7–13
	number	7	9–15
	on TerVIII length	73	84–114
	length / Ant3 AD	3.22–3.41	3.43–5.00
	number on subgenital on anterior half plate along the hind margin	–	13–18
		–	10–19
AntL	AntLB	94–96	114–137
	PT	364–372	430–541
	PT / AntLB	3.87–3.89	3.50–5.00
URS	number of accessory setae	6	6
	length	207	233–252
	W Head	0.51	0.51–0.57
	length / 2HT	1.64–1.71	1.63–1.84
	AntLB	2.16–2.22	1.81–2.18

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Males	Oviparous females
2HT	length	121–126	129–149
	maximum width length / W Head	4.17–4.25 0.30–0.31	4.27–5.00 0.28–0.33
	AntLB	1.30–1.32	1.06–1.27
SIPH	length	129–144	200–253
	length / length of body	0.10–0.11	0.12–0.14
	length / BW SIPH	2.91–3.35	2.97–5.06
	length / MW SIPH	3.92–4.07	4.05–5.71
	length / Ant3	0.37–0.41	0.51–0.63
CAUD	length	99	156–190
	length / BW CAUD	0.98	0.92–1.17
	number of setae	6	7–10
Length of SIPH / length of CAUD		1.31–1.46	1.25–1.46

FIGURES 52–54. Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) , alate viviparous female. 52, 3rd antennal segment; 53, siphunculus; 54, cauda. FIGURE 55. Male of Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) (20.ix.1962, Poland, KuyavianPomeranian Voivodeship, Bydgoszcz). FIGURES 56–58. Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) , male. 56, 3rd antennal segment; 57, siphunculus; 58, cauda. FIGURE 59. Oviparous female of Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) (20.ix.1962, Poland, Kuyavian-Pomeranian Voivodeship, Bydgoszcz). FIGURES 60–63. Macrosiphoniella ( Ramitrichophorus ) medvedevi ( Bozhko, 1950 ) , oviparous female. 60, 3rd antennal segment; 61, hind tibia; 62, siphunculus; 63, cauda. Description. Apterous viviparous female. Body elongate elliptical, 1.6–2.0 times as long as wide. The living specimens reddish yellow, with a large black patch on dorsum of abdomen (this patch is absent in macerated specimens), with dark head, antennae, legs (except bases of femora), siphunculi and cauda; with distinct waxy pulverulence, in this case, the aphids are gray with brown-red streaks. Cleared specimens with dark brown front, antennae (except base of 3rd antennal segment), distal half of the rostrum, legs (except base of femora) and siphunculi, with brown head (except front), sclerites at the base of the coxae, peritremes, anal plate and cauda, with light-brown bands and scleroites on thorax and abdomen and subgenital plate. Thorax with more or less wide bands on pro- and mesonotum and row of sclerites on metanotum; abdomen with rows of spinal sclerites, with small marginal sclerites and peritrems on I–VII segments, with antesiphuncular sclerites and with band on VIII tergite; sclerites on the thorax and abdomen are often small, pale and rare; sclerites on VII tergite often and on metanotum sometimes fused to form a short band. Surface of head, thoracic dorsum and abdominal tergites I–V smooth, sometimes weakly wrinkled, on VI–VII abdominal tergites with rows of small pointed spinules, which on VIII abdominal tergite partially fuse to form scales; thoracic venter smooth, ventral abdomen with long rows of small hardly visible spinules, sometimes forming strongly elongate cells. Setae on head, thorax and dorsum long, finely pointed, often bifurcating at the apices, in this case, each of the teeth of the fork is finely pointed. Third antennal segment with 11–30 secondary rhinaria, fourth segment with 0–4, fifth with 0 rhinaria. Rhinaria with internal diameter 6–23 μm, with external diameter 2.7–5.5 times as long as high of rhinaria. Setae on antennae blunt, pointed or rarely very weakly capitate. Rostrum is often very long, reaching abdominal segment III–V. Ultimate rostral segment 5.1–8.2 times as long as its basal width. Setae on legs pointed or finely pointed. Peritremes on abdominal segments I and II separated by a distance equal to or less than diameter of peritreme. Siphunculi with polygonal reticulation on distal 0.33–0.53 of length (largest transverse row in reticulate part of siphunculi consisting of 4–7 cells). Subgenital and anal plates with finely pointed setae. Cauda nearly triangular, with some constriction in the middle, which divides it into broad base and conical and rounded on apex distal part; setae on cauda long and finely pointed. Alate viviparous female. Body elongate elliptical, 2.1–2.5 times as long as wide. The living specimens with dark head and thorax. Cleared specimens with dark brown thorax; abdomen with distinct brown marginal sclerites on segments I–VII, spinal sclerites on abdominal tergites I–VI very small, light-brown or often absent, antesiphuncular sclerites hardly visible. Third antennal segment with 22–32 secondary rhinaria, fourth segment with 2–11, fifth with 0–1 rhinaria. Rostrum is often very long, reaching abdominal segment II–V. Peritremes on abdominal segments I and II separated by a distance less than diameter of peritreme or fused. Siphunculi with polygonal reticulation on distal 0.48–0.60 of length. Male. Body elongate elliptical, 2.0 times as long as wide. Colour of living specimens unknown. Tibiae of all legs dark brown only on the apices and brown in the middle. Marginal sclerites on I–VII segments distinct and sometimes relatively large. Third antennal segment with 33–37 secondary rhinaria, 4th segment with 9–11 and 5th segment with 4–5 secondary rhinaria. Siphunculi with polygonal reticulation on distal 0.35–0.46 of length. Cauda escuteon-shaped (misshapen specimen?). Hind tibiae with 1–3 round or oval pheromone plates. Oviparous female. Body 1.7–2.1 times as long as wide. Colour of living specimens unknown. Tibiae of all legs dark brown only on the apices and brown or light-brown in the middle. Marginal sclerites on abdomen absent, spinal sclerites on abdominal tergites I–VI often small or also absent. Third antennal segment with 8–23 secondary rhinaria, fourth and fifth segments with 0 rhinaria. Rostrum reaching abdominal segment III–IV. Siphunculi with polygonal reticulation on distal 0.34–0.48 of length (largest transverse row in reticulate part of siphunculi consisting of 4–6 cells). Cauda nearly triangular, sometimes with very weak constriction in the middle. Hind tibiae more or less distinctly swollen on basal half, with 27–55 round or oval pheromone plates, more of them located on basal half. Systematic relationships . The species is very close to Macrosiphoniella ( Ramitrichophorus ) janckei Börner, 1939 . The differences between them are given in the keys. Distribution . Germany ( Mecklenburg-Vorpommern ) ( Müller 1969 ), Poland ( West Pomeranian Voivodeship , Świnoujście and Kamień County, Międzyzdroje; Warmian-Masurian Voivodeship , Olsztyn; Kuyavian-Pomeranian Voivodeship , Bydgoszcz; Greater Poland Voivodeship , Poznań; Mazovian Voivodeship, Otwock County, Otwock; Podlasie Voivodeship, Mońki County, near Dolistowo Stare Vill.; Pomeranian Voivodeship , Nowy Dwór Gdański County, Krynica Morska) ( Szelegiewicz 1958a , 1958b , 1967 , 1968 , 1974 , 1975 , 1978 ; Huculak 1965 ; Achremowicz 1967 , 1975 ; Czylok et al. 1982 ; Nast et al . 1990 ; Węgierek & Wojciechowski 2004 ; Osiadacz & Hałaj 2010 ), Russia ( Moscow Province , Serpukhovsky District, Luzhki Vill.), Ukraine ( Kharkiv Province , Chuhuiv District, near Zmiiv) ( Bozhko 1950 , 1963 , 1976 ), Kazakhstan (?) (Aktobe Region, near Aktobe and in Temir District, near Kalmakkyrgan railway station) ( Smailova 1980 ), Iran ( Rezwani et al. 1994 ). Smailova (1980) indicates that two samples of M. ( R. ) medvedevi were collected from stems and pedicels of Helichrysum arenarium in Kazakhstan (Aktobe Region: near Aktobe and in Temir District, near Kalmakkyrgan railway station). Aphid colonies consisted of apterous viviparous females and immatures of 2–3 instars. However, Kadyrbekov in his summary on the fauna of aphids of Kazakhstan (2017) does not give this species for this territory, indicating that only two species have been found in Kazakhstan — M . ( R .) janckei and M . ( R .) hillerislambersi . Thus, the presence of M. ( R. ) medvedevi in Kazakhstan remains doubtful. Biology. The species lives on stems, inflorescences and the lower surface of leaves of Helichrysum arenarium (L.) Moench, always attended by ants ( Szelegiewicz 1958a , 1958b , 1967 ; Huculak 1965 ; Czylok et al. 1982 ), including Formica cinerea Mayr. ( Szelegiewicz 1958a ) . Apterous male and oviparous females of M . ( R .) medvedevi were found in Poland ( Kuyavian-Pomeranian Voivodeship , Bydgoszcz) by Szelegiewicz on 20.ix.1962 . Previously, another species— Macrosiphoniella nikolajevi Kadyrbekov, 1999 —was included in the subgenus Macrosiphoniella .