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Pentamyzus hille ris lambers, a neotropical genus of the tribe Macrosiphini (Hemiptera: Aphididae: Aphidinae), with the description of a new species PDF

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Preview Pentamyzus hille ris lambers, a neotropical genus of the tribe Macrosiphini (Hemiptera: Aphididae: Aphidinae), with the description of a new species

PROC. ENTOMOL. SOC. WASH. 104(4). 2002. pp. 918-927 PENTAMYZUS HILLE RIS LAMBERS, A NEOTROPICAL GENUS OF THE TRIBE MACROSIPHINI (HEMIPTERA: APHIDIDAE: APHIDINAE), WITH THE DESCRIPTION OF A NEW SPECIES J. M. NiETO Nafria, M. p. Mier Durante, and J. Ortego (JMNN, MPMD) Departamento de Biologia Animal. Universidad de Leon, E-24071 Leon, Spain, (e-mail: [email protected]); (JO) Instituto de Sanidad y Calidad Agrope- cuaria de Mendoza. Boulogne sur Mer, 3050.5500 Mendoza, Argentina. — Abstract. Pentamyzus fueguinus, n. sp., is described and two little-known South American species ofPentamyzus, P. tenuis and P. acaenae, are studied from aphids caught in Tierra del Fuego (Argentina). Considering that Pentamyzus is one of the few genera of Macrosiphini in the Neotropical Region, we present several hypothesis about its evo- lution in the subcontinent. — Resumen. A partir de pulgones capturados en Tierra del Fuego (Argentina) se describe Pentamyzusfueguinus n. sp. y se estudian otras dos especies sudamericanas poco cono- cidas: P. tenuis y P. acaenae. Aprovechando que Pentamyzus es uno de los pocos generos de Macrosiphini con representacion neotropical se analizan varias hipotesis sobre su his- toria evolutiva en el subcontinente. Key Words: Pentamyzus, Aphididae, Macrosiphini, aphid evolution, new species, Tierra del Fuego, Argentina Macrosiphini (Aphidinae) is the richest species). Utamphorophora peruviana (Es- tribe both in species and genera in the fam- sig 1953), Macrosiphum salviae Bartholo- ily Aphididae (Remaudiere and Remaudiere mew 1932, and Wahlgreniella australis 1997, Eastop 1998) and it is very difficult Delfino 1981 may also be native. Penta- to subdivide it into subtribes "because spe- myzus is the only genus of these with spe- ciation in a great scale has taken place dur- cies living on Poaceae or Rosaceae, Vv'hich ing a rather short time rather recently" are very important plant families in the spe- (Heie 1992). The tribe Macrosiphini is ciation of Aphidinae (Heie 1996, von Doh- poorly represented in the Neotropical Re- len and Moran 2000). gion and most ofits species recorded in this Pentamyzus includes fourknown species: region are allochthonous (Smith and Cer- P. acaenae (Schouteden 1904) from Tierra meli 1979, Nieto Nafria et al. 1994). Au- del Fuego (Argentina); the type species P. tochthonous Neotropical species belong to graminis Hille Ris Lambers 1966 from Cal- the genera Microparsus Patch 1908 (several ifornia; and P.falklandicus Hille Ris Lamb- species), Uroleucon Mcrdvilko 1914 (many ers 1974 and P. tenuis Brown 1987 both species), Pentamyzus Hille Ris Lambers from the Falkland Islands. The last three 1966 (3 species), Blanchardaphis Ortego, species live on Poaceae and the first one Nieto Nafria, and Mier Durante 1998 (2 was described on Acaena splendens Hook species) and Nietonafriella Ortego 1998 (1 & Am. (Rosaceae), but this record (Schou- VOLUME 104. NUMBER 4 919 teden 1904) was considered doubtful by Results Hille Ris Lambers (1974) and Brown ( 1987), due to ( 1 ) the known biology of the ideTnhtrifeieeds:pec(ai)esPo.faPcciaitednnaiyez.u(.sSchhaovuetebdeeenn other species ofthe genus, and (2) the pres- ence of grass fragments in the specimen B1r90o4w)nfr1o98m7,saomnpelevsiv1ipaanrdou2s; f(ebm)alP.e,tefnruoims tube of the syntypes collected at San Pi'o sample 3; and (c) a new species from sam- Cape, Ushuaia in 1892. The four species are ples 3 and 4. known only by their respective typical se- ries (Schouteden 1904; Hille Ris Lambers Peutamyzusfiie^uiniis Nieto Nafria, 1966. 1974: Brown 1987). Mier Durante, and Ortego, new species Materials and Methods (Figs. 1, 2B) Apterous viviparous female (n = 413; 67 Supported by project LE20/99 (granted measured) (Table 1; Figs. lA. 2B).—When by the Regional Government of Castilla y alive reddish brown to greenish brown, oval Leon, Spain), four samples of PenTaniyzus in shape, and dorsally little convex and rig- were collected in Tiena del Fuego (Argen- idly sclerotized: tergites of ab.seg. I to IV tina) by J.M. Nieto Nafria and M.P. Mier or V form a continuous plate, tergites of Durante: thoracic segments and ab.seg.VI. VII and 1) On Acaena spletidens (Rosaceae), Us- VII free (sometimes separation metathorax- huaia: Tiena Mayor, 9.1.2000 [49 apterous ab.seg.I and ab.seg.IV-V and abd.seg.V-VI viviparous females, 4 apterous males, and not extending to marginal areas). Mounted nymphs], see P. acaenae; specimens yellowish with distal ^/i-Vs ofan- 2) on Acaena splendens (Rosaceae). Ushua- tenna, u.r.s., and tarsi brown; well-pigment- ia: Glacial Martial. 14.1.2000 [6 apterous ed specimens with other pigmented areas: viviparous females, 2 apterous oviparous parts of antenna, clypeus, third rostral seg- females and nymphs], see P. acaenae; ment, coxae and parts of femora and tibiae, 3) on Alopecunis magellanicus van brac- intersegmental sclerites and marginal spots teolatus (Phil.) M. C. Mariano (Poaceae), on prothorax to ab.seg.IV, plus slightly pig- Rio Grande, 12.1.2000 [288 apterous vivip- mented anal plate and siphunculi. Frontal arous females, 4 apterous oviparous fe- tubercles conspicuous, lateral ones diver- males, 26 apterous males, and nymphs]: see gent and with 1-2 setae; medial one slightly P. tenuis and the new species; and displaced to ventral part ofhead. Dorsal cu- 4) on Hordeum comosiim J. & C. Presl (Po- ticle sclerotized and more or less alveolat- aceae), Rfo Grande: San Pablo Cape, ed. Dorsal setae scarce, stiff, short and 12.1.2000 [192 apterous viviparous females, blunt, as antennal ones; ventral setae point- 4 apterous oviparous females and nymphs]; ed, not too stiff, and longer than dorsal see the new species. ones. Rostrum reaching middle coxae; setae Abbreviations used in the text are as fol- long, pointed and flexible. Legs relatively lows: ab.seg. = abdominal segment(s); short; with setae similar to dorsal ones, ex- ant.Ill, ant.IV = antennal segments III, IV; cept: (1 ) tarsal setae pointed, and (2) ventral ant.Vb, antVpt = base and processus ter- setae on femora and distal setae on tibiae minalis of the antennal segment V: BL = pointed and longer than those, especially a body length; BW = body width; c. = Cau- group of 4 to 6 external and distal setae on da; D = basal diameter of ant.Ill; IAD = tibiae, normally at most 0.5 times tibial inter-antennal distance; h.tb. = hind tibia; width at this point. Proximal half of h.tb. h.t.II = second segment of hind tarsus; s. with 0-4(exceptionally to 9) scent plates = siphunculus; u.r.s. = ultimate rostral seg- placed inside. Abdominal papillae absent. ment. Siphunculius slightly sigmoid and clavate. 920 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Fig. L Peiuamyzusfueguimis. A, Apteorus viviparous female, habitus, dorsal view to right and ventral to left. B, Male, antenna. Scale bar = 1 mm. Illustration by M. Enrique Ortega Lorenzo (Leon. Spain). — — . VOLUME NUMBER 104. 4 921 with a cylindrical part at end and a well- samples 3 and 4, deposited in the authors' developed flange; wrinkled at base, smooth collections and in The Natural History Mu- on swollen part and striate on cylindrical seum (London). Museum National distal neck. Subgenital setae relatively long d'Hisloire Naturelle (Paris), and Zoologisk and pointed. Gonapophyses conspicuous. Museum, K0benhavns Universitet (Copen- Cauda more or less elongated-triangular hagen). — with convex sides, with long, flexible and Etymology. The specific name is from pointed setae. Metric and other meristic Tierra del Fuego, in masculine to agree with data in Table 1 — the gender of the genus. Alatae viviparous female. Not known Biology and distribution. Pcuhimyzus and probably does not exist. In material col- fuegiiinus is a monoecious species with ap- lected, several thousand nymphs and none terous males. Their biological cycle is short. of them alatoid. Males and oviparous females and many Oviparo—us female (n = 4; 4 measured) male nymphs, which are recognizable by (Table 1 ). Very similar to viviparous fe- their brown or black appendages, were col- male. Proximal half of h.tb. lightly swollen lected in early austral summer. Aphids form and with (exceptionally from 13) 20-44 groups of 4 to 12 specimens on the outface scent plates, placed inside. Subgenital plate of leaves of Alopecurus mageUcmicus var. with (8)13-22 and 13-28 discal and pos- bracteolatus and Hordeum comosum, terior setae, respectively. which grow in moist areas near the Grande Apterous male (—n = 26; 14 measured) River and a little river on the beach at San (Table 1; Fig. IB). In life darker than fe- Pablo Cape, respectively (this moist micro- male, with legs and antenna brown to black habitat is very similar to that reported by when alive. When mounted, heavily pig- Hille Ris Lambers 1966 for Pentamyzus mented: ant.Ill to ant.Vpt, u.r.s.. 1^, distal gramiuis). Grasses of dry neighboring en- part of tibiae and tarsi dark brown, and vironments are not colonized. We looked head, rostrum, legs, intersegmental and for the species without success on different stigmatic sclerites, siphunculi, genitalia and grasses in moist and dry environments of Cauda smoked; several specimens with light the Argentinean TieiTa del Fuego and Pa- pigmented marginal areas on pre-siphun- tagonia. cular segments and one transversal bar on Pentamyzus acaenae (Schouteden 1904) ab.seg.VIII. Frontomedial tubercle ventral, larger than faint cephalic lateral ones, and (Fig. 2A) nodulose as dorsal cuticle ofthorax and ab- Apterous viviparous female (Table 1). domen. Similar in shape to male ofP. acae- When alive light to green and very convex. nae, but with more alveolated and darker Dorsum of body heavily sclerotized. nodu- cuticle, especially on sclerotized areas. lose and unpigmented; tergites of metatho- Ant.III and ant.IV with 41-77 and 13-30 rax to ab.seg.VI form a continuous plate, (exceptionally ant.Vb with 4) small, round tergites of ab.seg. VII and VIII mutually and striated-walle—d secondary sensoria. free. Very few metric differences have been Type material. Holotype: apterous vi- observed with regard to types [Schouteden viparous female (measured specimen num- 1904J, the only known specimens; these ber 2) collected on Alopecurus magellani- differences could be due to different collec- cus var. bracteolatits at Rio Grande (Tierra tion dates, November and January, but dif- del Fuego Province. Argentina), 12.1.2000. ferent volume of samples must also be tak- in collection Universidad de Leon, Depar- en into account. Dorsal setae scarce, stiff, tamento de Biologia Animal. Paratypes: short and blunt; setae of antenna and legs other apterous viviparous females, ovipa- similar in shape and in length, except (1) rous females and males of studied material tarsal setae pointed, and (2) ventral setae of 922 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table Metric and meristic data of Pentamyzusfueguinus and P. acaenae, apterous vivparous females 1. (apt. viv. f.). oviparous females (apt. ov. f.) and males (apt. mal.). In square brackets in the apterous vivparous females column of P. acaenae are data from types. — VOLUME 104. NUMBER 4 923 I [/ n B /•/ Fig. 2. Apterous viviparous female. A. PeiUainyziisacaciuie. B, P.fiici^iiiiuts. Head-prothorax (1). prothorax- mesothorax (2), mesothorax-metathorax (3). metathorax-ab.seg.I (4), ab.seg.IV-ab.seg.V (5). ab.seg.V-ab.seg.VI (6), ab.seg.VI-ab.seg.VlI (7), ab.seg.VII-ab.seg.VIII (8) intersegmental lines. femora and di.stal setae oftibiae pointed and tral summer and winged viviparous females longer than those, especially a group of 4 possibly do not exist (alatoid nymphs have to 8 external and distal on tibiae, normally not been observed). Aphids live on the at least 0.5 times tibial width at this point. leaves or shoots of the rosaceous Acaena Ventral setae pointed and flexible and lon- splendeus, which is very common in the de- ger than dorsal ones. ciduous forest of the mountainous area of Apterous oviparous—female (previously the southern half of Tiena del Fuego; this undescribed) (Table 1). Very similarto vi- plant and other species of Acaena exist in viparous female. With group of 8—16 scent other areas ofArgentinean Tieira del Fuego plates on an internal small swollen area at and Patagonia, but the aphid has not been proximal half of h.tb. Subgenital plate with recorded there. 10-16 discal setae and 16 posterior setae. Pentamyzus tenuis Brown 1987 Apterous male (previously undescribed) — (Table 1). Green, but darker than female, Pentamyzus tenuis is a very peculiar spe- with smoked head and legs when alive; pig- cies (the body is elongated and the front is mentation similar to male of P. fueguinus convex). We collected one female, which is when mounted; dorsal cuticle ofthorax and similar in general to the types (two vivip- abdomen less nodulose as male of P. fue- arous females and two nymphs), but some guinus. Ant.Ill, ant.IV and ant.Vb respec- differences have been observed (Table 2). tively with 61-73, 25-32 and 0-1 second- The host plant in Tierra del Fuego is not ary sensoria, similar to those new species. the typical host plant, Poa alopecurus Biology and distribution. Pentamyzus Kunth. (Brown 1987). The presence of this acaenae is a monoecious species, with ap- aphid in South America, at a locality situ- terous males. Its life cycle is very short. ated approximately eight degrees to the Sexuales occur in the first month ofthe aus- west and two degrees to the south of the 924 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Metric and meristic data ofapterous viviparous females of Pentaiuyzus tenuis from Falkland Islands and Tierra del Fuego. VOLUME 104. NUMBER 4 925 mGP Z 2b mRP mGPj- iGpP) 926 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON to California. The primitive Pentamyzus the last species of a lineage which returned may have been dioecious, as its ancestor. from South America. Later, it diversified into two monoecious The three hypotheses are concomitant subHneages, one rosaceous-feeding and an- with the present theory on the colonization other grass-feeding. The subHneage on Ro- of South America by the Aphidinae (the saceae finally established itself on herba- Holarctic origin of this subfamily and its ceous ones, because there are no other au- subordinated taxa, such as Macrosiphini, is tochthonous rosaceans species in the far generally accepted, Heie 1996). South south (this is not suiprising in monoecious America was invaded in different waves by aphids, for example Chaetosiphon fragae- different lineages of this tribe (so Micro- folii (Cockerell 1901), Amphorophora gei parsus, Pentamyzus and siphunculi-reticu- (Borner 1939), or Acyrthosiphon boreale lated aphids, such as Uroleucon, Nietona- Hille Ris Lambers 1952 [Macrosiphini] liv- friella, and Blanchardaphis, belong to dif- ing on Rosaceae herbaceous species), and ferent generic groups and live on plants of P. acaenae belongs to this sublineage. The different families, the first one on Fabaceae other known Pentamyzus species belong to and last three on Asteraceae). the other sublinage; in this group the most The three hypotheses may be correct, but different species are: (i) P. tenuis, which we prefer 1 or, to a lesser extent, 3. because has a elongated body and is South Ameri- they are the most parsimonious when ex- can, and (ii) P. graminis, which has several plaining the morphological similarities be- derivative characteristics such as small-cla- tween P. fueguinus and P. acaenae and the vate siphunculi, lack of complementary presence of two "special" grass-feeding u.r.s. setae, and lack of marginal papillae species, P. tenuis (because of its body ap- (this last characteristic is also present in pearance) and P. graminis (because its de- other species of the genus) and belongs to rivative characteristics and distribution). In the returned lineage. either case, only molecular analysis could (2) The primitive dioecious Pentamyzus allow us to arrive at a conclusion as to stock was North American, and host di- which of these hypotheses is correct. verging originated in North America. Later, Acknowledgments two sublineages colonized South America via the Andes. Here we found two possi- We thank Roger Blackman and Paul bilities: (2a) The northern species on Ro- Brown (The Natural History Museum, Lon- saceae disappeared. P. acaenae belongs to don) and Ole Heie (Holte, Denmark) for the rosacean sublineage established in the constructive criticism ofthe manuscript. We South, P. graminis belongs to the old north- also thank to E. Mendez, E. Martinez and ern grass sublineage, and P. falklandicus, P. A. Dalmasso, Botanists of the lADIZA fueguinus, and P. tenuis belong to the (Mendoza, Argentina) for identifying the southern grasses sublineage; or (2b) all plant species from Tiena del Fuego. northern species disappeared and P. gra- minis returned from South America, such as Literature Cited in hypothesis 1. (3) Pentamyzus was a North American BrowRni,sPL.am1b98e7r.sK(eHyomotopttehrea,geAnpuhsidPideanet)a.mywziutsh aHidlel-e monoecious and grass-feeding genus. Later scription of a new species from the Falkland Is- it colonized South America, and one sub- lands. Systematic Entomology 12(1): 1-6. lineage (represented by P. acaenae) ac- Eastop. V. E 1998. Why do aphids do that?, pp 37- quired Acaena as a host. As in hypothesis 47. In NietoNafn'a,J. M. andA. E G. Dixon,eds. Aphids in Natural and Managed Ecosystems. 2, there are two possibilities with regard to Universidad de Leon. Leon. P. graminis: (i) it belongs to the non-dis- Footit, R. G. and W. R. Richards 1993. The Insects placed old-lineage (in Fig. 3), or (ii) it is and Arachnids ofCanada. Part 22. The Genera of VOLUME NUMBER 104, 4 927 the Aphids of Canada (Homoptera: Aphidi)idea Nieto Nafn'a, J. M.. M. A. Delfino. y M. R Mier Du- and Phylloxeroidea). Agriculture Canada. Re- rante 1994. Laafidofaunade la Argentina, sucon- search Brancli. publication num. \H^5. Ottawa. ocimiento en 1992. Universidad de Le6n (Secre- 766 pp. tariado de Publicaciones). Leon. 235 pp. Heie. O. E. 1W2. The Aphidoidea (HemipteralofFen- Rcinaudiere. G. and M. Remaudiere 1997. Catalogue no.scandia and Denmark. IV. Family Aphididae: des Aphididae du Monde / Catalogue of the Part I of tribe Macrosiphini of Subfamily Aphi- World's Aphididae. Homoptera Aphidoidea. Insti- dinae. Fauna Entomologica .Scandinavica 25: 1- tut National tic la Recherche Agronomic|ue. Paris. 188. 478 pp. . 1996. The evolutionary history ofaphids and .Schouteden, H. 1904. Aphiden. Hamburger Magal- hypothesis on the eoevolution of aphids and haensische Sammelrei.se 4: 3-6. plants. Bolletino di Zoologia agraria e Bachicol- Smith, C. F. and M. M. Cermeli 1979. An Annotated tura (Serie ID 29(2): 149-155. List of Aphididae (Homoptera) of the Caribbean Hille Ris Lambers D. 1966. Notes on California Islands and South and Central America. North aphids. with descriptions of new genera and new Carolina Agricultural Research Service Technical species (Homoptera: Aphididae). Hilgardia Bulletin 259: 1-131. 37(15): 569-623. von Dohlen, C. D. and N. A. Moran, 2000. Molecular . 1974. On American aphids. with descriptions data support a rapid radiation of aphids on the of a new genus and some new species (Homop- Cretaceous and multiple origins of host alterna- tera: Aphididae). Tijd.schrift voor Entomologie tion. BiologicalJournalofthe Linnean Society71: 1 17(4): 103-155. 689-717.

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