PROC. ENTOMOL. SOC. WASH. 107(3), 2005, pp. 700-728 A SYSTEMATIC REAPPRAISAL OF THE GENUS DIURAPHIS AIZENBERG (HEMIPTERA: APHIDIDAE) Gary L. Miller, Manya B. Stoetzel, and Ethan C. Kane Systematic Entomology Laboratory, Plant Sciences Institute, Agricultural Research Ser- MD vice, U.S. Department of Agriculture, Bldg. 005, BARC-West, Beltsville, 20705, U.S.A. (email: [email protected]; [email protected]; eckane@sel. barc.usda.gov) — Abstract. Adult female apterae and alates of the genus Diuraphis Aizenberg are de- scribed and illustrated, and keys are provided for identification. Diuraphis elymophila G.- X. Zhang is considered a new synonym of Diuraphis frequens (Walker) and Diuraphis muehlei (Bomer) is considered a new synonym of Diuraphis noxia (Kurdjumov). A phy- logenetic analysis suggests that while the clade that contains Diuraphis noxia + Diuraphis mexicana is monophyletic, the previously recognized subgenus Holcaphis is paraphyletic. Key Words: aphid, Diuraphis, Holcaphis The genus Diuraphis was proposed in are generally recognized (Eastop and Hille 1935 with Brachycolus noxius Mordvilko Ris Lambers 1976, Remaudiere and Re- [= Diuraphis noxia (Kurdjumov)] as its maudiere 1997), their monophyly has not type species (Aizenberg 1935). As currently been tested. defined (Remaudiere and Remaudiere Diuraphis was not well known until the 1997), there are 11 species in the genus re- early 20"^ century when outbreaks of D. ferable to two generally recognized subgen- noxia (the Russian wheat aphid) and D. tri- era. The subgenus Diuraphis sensu stricto tici (the western wheat aphid) in Russia and contains three species: D. mexicana the western United States, respectively, (McVicar Baker), D. muehlei (Borner) and brought attention to the destructiveness of D. noxia (Kurjumov). The subgenus Hol- these aphids on wheat. By the late 1970's, attention was once again focused on Diura- caphis Hille Ris Lambers (1939) comprises phis, especially D. noxia. Substantial range the remaining species and includes: D. extension ofD. noxia was first documented agropyronophaga G.-x. Zhang, D. agrosti- in South Africa in 1978 (Durr 1983) and in dis (Muddathir), D. bromicola (Hille Ris the United States in 1986 (Stoetzel 1987). Lambers), D. calamagrostis (Ossiannils- This aphid spread quickly and now has son), D. elymophila G.-x. Zhang, D. fre- been recorded throughout much of the quens (Walker), D. hold (Hille Ris Lamb- wheat growing regions of the world. In the ers), and D. tritici (Gillette). Historically, United States, its damage to wheat and bar- Diuraphis sensu stricto differs from Hol- ley was extensive and resulted in heavy caphis by the presence of a supracaudal crop losses in some fields (Stoetzel 1987). process on the eighth abdominal tergite and By 1993, D. noxia was a pest in 16 western usually the presence of marginal tubercles states and caused cumulative losses esti- on abdominal segments II-VI (Heie 1992). mated at $500-900 million dollars (Bemal Although the two subgenera of Diuraphis et al. 1993, Morrison and Peairs 1998). VOLUME NUMBER 107, 3 701 Because D. noxia is recognized as an of China, and University of Rostock Insect economically important pest, much of the Collection (URIC), Sagerheide, Germany. literature on Diuraphis has concentrated on Measurements are presented in millimeters this species. However, Zhang et al. (1991) as minimum and maximum ranges of rep- gave a key to Diuraphis and discussed phy- resentative specimens. logenetic relationships. They also included Morphological terms and structures the descriptions of D. agropyronophaga adapted from Stoetzel et al. (1999) are used and D. elymophila. Kovalev et al. (1991) in this work. Those terms are listed below, provided a key to the Diuraphis apterous and equivalent terms that may be found in viviparous females and reviewed the Rus- other literature are listed in parentheses for sian literature. Descriptions, keys to apterae reference: terminal process (= unguis, pro- and alata, and illustrations of Diuraphis of cessus terminalis); secondary sensoria (= Fennoscandia and Denmark were provided secondary rhinaria); cornicle (= siphuncu- by Heie (1992). Halbert et al. (1992) also lus); fundatrix/fundatrices (= stem moth- included keys to North American Diura- er(s)); aptera/apterae (= wingless vivipa- phis. rous female(s)); alata/alatae (= winged vi- Apterae and alatae of the genus Brachy- viparous female(s)); and ovipara/oviparae colus Buckton closely resemble Diuraphis, (= egg-laying female(s)). the main difference is the position of the The information under Specimens Ex- cornicle (Heie 1992). In Brachycolus and amined is organized to conserve space. Ab- most other genera of aphids, the cornicle is breviations for fundatrices, apterae, alatae, on the posterior portion of abdominal ter- oviparae, apterous males, alate males, and gite V, whereas in Diuraphis the cornicle is immatures are listed as: fund.; ap.; al.; ov.; ogintethVeIp(oHseteireio1r99p2o)r.tiHoonwoefvearb,donmeiinthaelr ttehre- alpe.cti;oanl.w<aJ;samnaddiemmat.trheespseactmievelloyc.alIiftya,cboult- relationship between Diuraphis and Bra- on a different date as a previously listed chycolus in a phylogenetic context nor the collection, duplicated information is not re- monophyly of the subgenera of Diuraphis peated. For example, the documentation have been examined. provided for a particular locality may be re- The objectives of this paper are to: (1) corded as: TEXAS: Big Bend. Vll-11- redescribe, illustrate, and present keys for 1978, on Broimis unioloides [= Bromus ca- Dthieuriadpehnitsifiscpaetciioens;ofanadpt(e2r)aetesatntdhealhaytpaoetho-f tharticus], I. M. Miller coll.. (2 al.) USNM; VIII-29-1957. XI-21-1957. lV-14-1978. ecsoimsprtihsatedDiofurtawphoisdisitsincmtonsoupbhgyrloeutpisc (ia.ne.d, on Bromus sp., (15 ap. on 15 si.) USNM. In this hypothetical example, the second Diuraphis (sensu stricto) and Holcaphis). collection was also found at Big Bend, even Materials and Methods though ''Big Bend" was not repeated. Synoptic descriptions are taken from When specimens are mounted on a single original descriptions, types, and identified slide (si.), it is not written as sucli bui is material from the Aphidoidea collection of: assumed. Bracketed (| |) text represents sup- Museum National D'Histoire Naturelle plemental information by the present au- (MNHN), Paris, France; National Museum thors for ckirilication purposes or refers to of Natural History (USNM). Beltsville, the original collection data of laboratory MD, U.S.A.; Canadian National Collection reared specimens. Collection data that are of Insects (CNCl), Ottawa. Canada; The the same except for collection date aic sim- Natural History Museum, London, U.K. ply listed seqLientiall\. Host plants listed in (BMNH); Institute of Zoology Academia the Specimens Examined scctit>ns are sum- Sinica (IZAS), Beijing, People's Republic marized in Table 2. 702 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 1. Data matrix used in the ciadistic analysis. 1111111112 2222222 1234567890 123456789 1245678 B. cerastii 1000110110 0010011 B. ciicubali 0000000210 0010001 B. stellariae 0010111000 0011111 B. asparagi 1010121100 1000010 D. agropyronophagc 0010020310 1011001 D. agrostidis 0001021001 1000011 D. bromicola 0011120011 101111—0 D. calamagrostis 0000121201 10010 D.freqiiens 1001020001 1000100 D. hold 0001010211 1010000 D. me.xicana 1111120301 1110000 D. noxia 1112120101 1111000 D. tritici 1002120300 1010100 — Phylogenetic analysis. Phylogenetic also noted Brachycorynella was similar to analyses of Diuraphis and selected out- Brachycolus. groups were conducted to test the mono- Characters: Thirty morphological char- phyly of Diuraphis and infer relationships acters were examined. Two characters within the genus to test the validity of cur- proved to be autapomorphic and were ex- rently recognized subgenera of Diuraphis cluded from the final data set as parsimony {Diuraphis sensu stricto and Holcaphis). uninformative. Of the remaining 28 char- Nine species {Diuraphis agropyronopha acters, 25 were coded as binary while three G.-x. Zhang, Diuraphis agrostidis (Mud- were treated as non-additive multistate dathir), Diuraphis bromicola (Hille Ris characters. Unknown or indeterminable Lambers), Diuraphis calamagrostis (Os- character states were coded as missing data. siannilsson), Diuraphis frequens (Walker), Character descriptions are outlined below Diuraphis hold (Hille Ris Lambers), Di- and the final data matrix is presented in Ta- uraphis mexicana (McVicar Baker), Diura- ble 1. phis noxia (Kurdjumov), and Diuraphis tri- Apterous viviparous female tici (Gillette)) are included in the present (excluding the fundatrix) analysis. Examination of type material of Diuraphis muehlei (Borner) and Diuraphis Head elymophila Zhang revealed that those spe- 1. Tips of median dorsal head setae: cies are junior synonyms of D. noxia and pointed or tapered (0); blunt (1). D. frequens, respectively. Antennae Brachycolus stellariae (Hardy), Brachy- 2. Base of scape with slight posterior-lat- colus cerastii (Kaltenbach), Brachycolus eral protuberance: absent (0); present cucubali (Passerini), and Brachycorynella (1). asparagi (Mordvilko) are included as out- 3. Antennal tubercle shape: undeveloped groups to provide a context in which to test or flat (0); moderately developed or the proposed monophyly of Diuraphis sen- slightly raised (1). su lato. Diuraphis is suspected to be closely 4. Antennal segment I: entirely pigmented related to Brachycolus and Brachycorynel- (0); pigmented medially (1); pale (2). la. Members of these three currently rec- 5. Antennal segment II venter: usually ognized genera were referable to Brachy- smooth (0); usually with some imbri- colus (Shaposhnikov 1964). Heie (1992) cations (1). ) VOLUME NUMBER 107, 3 703 6. Antennal segment III: entirely pig- usually not raised medially (0): raised mented except for base (0); pigmenta- medially ( 1). tion reduced to base or apex (1); pale 23. Cauda: apically rounded (0); apically (2). pointed (1 ). 7. Antennal segment III: imbricated 24. Mid-ventral caudal spicules: individu- throughout (0); partly imbricated (1). ally separate (0); connected ( I 8. Length of antennal segment IV com- 25. Length of cauda compared to length of pared to length of segment V: segment hind tarsus II: longer (0); shorter or ap- IV usually longer than segment V (0); proximately equal ( 1 ). segment IV usually shorter than seg- ment V (1); length of segment IV sub- Alate viviparous females equal to length of segment V (2); Antennae length of segment IV variable when 26. Antennal segment IV secondary sen- compared to segment V (3). soria: usually present (0); usually ab- 9. Length of antennal segment III com- sent (1). 27. Length of antennal segments IV + V pared to length of antennal segment IV +V: shorter (0); longer (1). compared to antennal segment III: lon- Mouthparts ger (0); shorter or subequal (I ). 10. Ultimate rostral segment accessory se- All phylogenetic analyses were per- tae: present (0); absent (1). formed using PAUP==^ (Swofford 2001). 11. Length of base of antenna! segment VI Maximum parsimony (MP) analyses were compared to ultimate rostral segment: conducted using branch-and-bound search- usually shorter or subequal (0); longer es, and bootstrap analyses involved 1.000 replicates of branch-and-bound searching. (1). MP 12. Number of setae anterolateral to post- In the initial analyses, all characters clypeus: 2-3 (0); 1 (1). were treated as unordered and as having Thorax equal weights. In subsequent analyses, the 13. Lateral prothoracic tubercles: present at successive approximations approach to least sometimes (0); absent (1). character weighting (SACW) was used to 14. Protibiae: not uniformly colored (0); oseglyec(tFatrhreismo1s9t69)c.laFdiosrtitchaellSy AreCliWablaenatloypsoils-, uniformly colored (1). 15. Metafemur: not uniformly colored (0); characters were weighted based on the re- scaled consistency index. uniformly colored (1). Abdomen Phylogenetic Results and Discussion 16. Large polygonal dorsal abdominal re- MP analysis generated 16 equally most ticulation: present (0); absent (1). parsimonious topologies all with a tree 17. Intersegmental sclerites: present (0); length of 83, consistency index (Ch of absent (1). 0.37. retention index (Rl) iW" 0.43. and re- 18. Lateral abdominal tubercles: present at scaled consistency index (RC) o^ 0.16. least sometimes (0); absent (1). Three iterations of SACW with characters 19. Apical flange of cornicle: present (0); weighted based on the rescaled consistency absent (1). index resulted in a single most parsimoni- 20. Cornicle with associated basal sclerite: ous topology (Fig. 1 ) with a tree length of present (0); absent (1). 14.28. CI of 0.63. RI of 0.75. and RC o\' 21. Cornicle position: anterior to stigmal 0.47. Bootstrap anal\sis of the unweighted pore VI (0); level or posterior to stig- data set pidduccil rcl;iii\cl\ Un\ support mal pore VI ( 1 ). values, with onl\ two ikkIcs reco\ered in 22. Dorsum of abdominal segment Vlll: greater than 5(Y/c ol" the replicates. 704 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 1 D. calamagrostis L= 14.28 D. hold Cl=0.63 ^/=0.75 D. agrostidis RC= 0.47 D. frequens D. tritici D. bromicola D. mexicana D. noxia B. asparagi D. agropyronophaga B. stellariae B. cucubali B. cerastii Fig. 1. Maximum parsimony topology resulting from three iteration of successive weighting and branch- and-bound analysis. Node labels indicate bootstrap proportions obtained from an analysis of the unweighted matrix. L = Length, CI - Consistency Index, RI = Retention Index, RC = Rescaled Consistency Index. The most parsimonious tree presented in 70%. This clade is united by a slight pos- Fig. 1 conflicts with current taxonomic con- terior-lateral protuberance on the base of cepts of Diuraphis in that the genus is not the scape and a medially raised dorsal sur- recovered as a well-supported monophylet- face of abdominal segment VIII. The ic group. The recovery ofB. asparagi with- grouping ofD. mexicana + D. noxia carries in the Diuraphis clade is problematic, but interesting biogeographic implications since not entirely unprecedented. B. asparagi D. mexicana and D. noxia are of Nearctic shares many similarities with Diuraphis and Palearctic origin, respectively. species and has been grouped with other The remaining Diuraphis species, how- species of Diuraphis is previous studies ever, were not recovered as a monophyletic (e.g., Shaposhnikov 1964). The relative po- sister-group to the D. mexicana + D. noxia sition ofthe cornicle with respect to stigmal clade and thus, it would be inappropriate to pore VI (character 21), although lacking treat this as a subgeneric division ofDiura- strong bootstrap support, is potentially syn- phis sensu lato. Therefore, although there is apomorphic for these taxa. support forDiuraphis sensu stricto, we con- Regarding proposed subgeneric divisions clude that it is not instructive to recognize of Diuraphis, the data examined in this the proposed subgenus Holcaphis (e.g., study provides support for Diuraphis sensu Eastop and Hille Ris Lambers 1976; Re- stricto (D. mexicana + D. noxia) which maudiere and Remaudiere 1997), since it was recovered with bootstrap support of most likely represents a non-monophyletic VOLUME 107, NUMBER 3 705 grouping of the remaining Diuraphis spe- - Ultimate rostral segment without accessory se- tae 6 cies. 5. Lengthofantennal segmentIIIshorterthanan- tennal segments IV + V; ultimate rostral seg- Diagnosis of the Genus Diuraphis ment approximately 2 times as long as wide at base D. agropyronophaga Diuraphis is characterized by an elongate - Length of antennal segment III subequal to body, relatively short antennae, antennal tu- longer than antennal segments IV + V; ulti- bercles low or weakly developed, abdomi- mate rostral segment approximately 3 times as nal dorsum usually without pigmented long as wide at base D. tririci sclerites anterior to segment VI but some 6. Abdomen with intersegmental sclerites. al- species have intersegmental abdominal though sclerites may be extremely reduced in some alata; length of hind tarsus II shorter or sclerites, cornicles inconspicuous and usu- subequal to length ofcauda 7 ally without an apical flange, and first tarsal - Abdomen without intersegmental sclerites; segments in adults with 3-3-2 setae. Some length of hind tarsus II longer than length of species have abdominal tubercles, spinal cauda 9 7. Legs stout, e.g., greatest width ofhind tibiae supracaudal process present or absent, and subequal or wider than the length of penulti- dorsal setae occasionally spatulate. Most mate antennal segment; rostrum length sube- species produce wax. Diuraphis species qual to width at base generally are associated with leaves of var- D. calamagrostis (apterae only) (in part) ious cultivated and wild grasses (Poaceae). - Legs more slender, e.g., greatest width ofhind tibiae less than the length of penultimate an- tennal segment; rostrum longer than width at Key to Apterae and Alatae Diuraphis base 8 (alatae of D. calamagrostis not included 8. Antennal segment III usually longer than seg- due to insufficient material) ment IV -I- V, occasionally subequal to seg- ment IV-I-V; cornicles short, approximately '^ 1. Abdominal segment VIII supracaudal process pre- to Vi the length ofthe cauda D. Jiolci sentasawelldevelopedfingerlikeprojection(Fig. - Antennal segment III usually shorter than seg- 9D); cornicle with apical flange; lateralprothorac- ment IV + V, occasionally subequal to seg- ictuberclesandabdominalmarginaltuberclespre- ment IV-I-V; cornicles very short, porelike, ap- sent D. noxia proximately '/,o to 1/5 the length of the cauda - Abdominal segment VIII supracaudal process D. agrostidis either absent, slightly raised, or present as a 9. Cornicle unpigmented. porelike, 'Ao to '/,o the conical or triangular protuberance but not fin- length ofthe cauda; dorsal .scleriteson abdom- gerlike; cornicle without apical flange; lateral inal segment VII absent or reduced to a few prothoracic tubercles and marginal tubercles scattered polygonally sclerotized areas . . . present or absent 2 D. hromicola 2. Prothoracictuberclesormarginaltuberclespre- - Cornicle pigmented, short butelongate. V^ toW sent 3 the length ofthe cauda; dorsal sclerites on ab- - Prothoracic tubercles or marginal tubercles ab- dominal segment VII well developed, extend- sent 4 ing nearly to spiracle D. frequens 3. Length of setae on antennal segment III ap- proximatelyVithe diameterofthebase;apterae Diuraphis ai^ropyroiiophaga G.-x. Zhang with supracaudal process on abdominal seg- (Fig. 2) ment VIII present as a conical or triangular Diuraphis (Holcaphis) agrapyroiiophaga protuberance (Fig. 8D), abdomen without in- tersegmental sclerites, cauda triangular with G.-x. Zhang, 1991:327: Zhang el al.l991: pointed apex D. mexiccmu 123; Remaudiere and Reinaiidierc 1997: - Length of setae on antennal segment III sube- 91 qual to the diameter ofthe base; apterae with- — out supracaudal process on abdominal segment Type material. Aptera hoUnype, No. VIII, abdomen with intersegmental sclerites, 6324-1-1-2, on Agropyron sp.. Nei Mongol Cauda parallel sided with bluntly rounded apex Aulomous Region, Fregzhen County, 19- D. calamagrostis (apterae only) (in part) VM976. G.-x. Zhang and T.-s. Zhong coll.. 4. Uslotryimsaetteaerostral segment with a pair of acces- -^ not seen. We ha\c studied a single paratype 706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Table 2. Host plants of Diuraphis. The following Table 2. Continued. host plant information represents a summary of those plants listed in the Specimens Examined sections. Al- Elymus glaucus Buckl. though some of the host data may represent aberrant Diuraphisfrequens (Walker) hosts, they are included for reference purposes. When common names for various hosts were used, they are Elymus sp. recorded as such on the list and the scientific name is Diuraphis tritici (Gillette) added in parentheses for cross-reference. Botanical Elytrigia repens var. repens names listed in the collection data were checked against the Integrated Taxonomic Information System Diuraphisfrequens (Walker) (Anonymous 2004a), The International Plant Names Holcus lanatus L. Index (Anonymous 2004b), and the National Genetic Diuraphis hold (Hille Ris Lambers) Resources Program, Germplasm Resources Informa- tion Network (Anonymous 2004c). Holcus mollis L. Diuraphis hold (Hille Ris Lambers) Agrostis stolonifera Hordeum murinum L. Diuraphis agrostidis Muddathir Diuraphis noxia (Kurdjumov) Agropyron sp. Hordeum vulgare L. Diuraphis agropyronophaga G.-x. Zhang Diuraphis hold (Hille Ris Lambers) Diuraphis noxia (Kurdjumov) Diuraphis noxia (Kurdjumov) Avena sp. Hordeum sp. Diuraphis tritici (Gillette) Diuraphis noxia (Kurdjumov) Barley (see Hordeum sp.) Mountain brome (see Bromus marginatus) Bromus catharticus Vahl Oats (see Avena sp.) Diuraphis mexicana (McVicar Baker) Pascopyrum smithii (Rydb.) A. Love Diuraphis noxia (Kurdjumov) Diuraphis tritici (Gillette) Bromus carinatus Hook. & Arn. Phleumpratense L. Diuraphis mexicana (McVicar Baker) Diuraphis noxia (Kurdjumov) Diuraphis noxia (Kurdjumov) Phleumpratense ssp. nodosum (L.) Arcang. Bromus inermis Leyss. Diuraphisfrequens (Walker) Diuraphis bromicola (Hille Ris Lambers) Diuraphis noxia (Kurdjumov) Bromus marginatus Nees ex Steud. Phleum sp. Diuraphis mexicana (McVicar Baker) Diuraphis noxia (Kurdjumov) Diuraphis tritici (Gillette) Quack grass (see Elytrigia repens var. repens) Bromuspolyanthus Scribn. Triticum aestivum L. Diuraphis mexicana (McVicar Baker) Diuraphis noxia (Kurdjumov) Bromus tectorum L. Diuraphis noxia (Kurdjumov) Triticum sp. Diuraphis noxia (Kurdjumov) Bromus sp. Diuraphis tritici (Gillette) Diuraphis mexicana (McVicar Baker) Wheat (see Triticum sp.) Calamagrostis lanceolata Diuraphis calmagrostis (Ossiannilsson) Calamagrostispurpurea Diuraphis calmagrostis (Ossiannilsson) slide deposited in IZAS with left label, Downy brome (see Bromus tectorum) "9921-1-1 Agropyron 90.VI.1" and right Echinochloa crus-galli (L.) Beauv. label, "PARATYPES, Holcaphis agropy- ZHANG ronophaga 16-VI-1990 Ningxia Diuraphisfrequens (Walker) China." Additional paratypes listed in Elymus dahuricus Turcz. ex Griseb. Diuraphisfrequens (Walker) Zhang et al. (1991—). Field features. Aptera grayish white. — VOLUME NUMBER 107, 3 707 2 Figs. 2-3. 2, Diuraphisaf^ropymnophai^a. A, Right side, aptera dorsum ofhead and antennal segments; left side, aptera venter of head and antennal segments I-II. B, Antenna ofalata. C. Cornicle ofaptera. D. Cauda of aptera. 3, D. agrosticlis. A, Right side, aptera dorsum ofhead and antennal segments; left side, aptera venterof head and antennal segments I-II. B, Antenna ofalata. C. Cornicle of aptera. D, Cauda ofaptera. covered with white powder (Zhang et al. 0.084-0.108 long; terminal process, 0.084- 1991). 0.132 long. Head scleroti/ed, sniotHh. with- Recognition characters. Aptera: Body out spinulation; IcMigest dorsal iicad setae length 2.220-2.232; width through eyes, subequal to width o^ antennal seginenl MI. 0.378. Antenna (Fig. 2A) shorter than body; Rostrum extending lo mosocoxae: tiltimatc segment III 0.150-0.216 long; IV 0.066- segment 0.108-0. 1 14 Ioiil:. ;ippn^\iiiiaicl\ 2 0.102 long; V 0.066-0.078 lone; base of VI times as IcMig as u itio al base, siihcciual to — 708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON hind tarsal segment II, with 2 accessory se- IV + V whereas the length ofantennal seg- tae. Pronotum without marginal tubercles. ment III is subequal to longer than antennal Hind tibia 0.468-0.570 long; hind tarsus II segments IV + V in D. tritici. See also 0.114 long. Abdomen smooth with ventral Notes section for D. tritici. surface spiculose, dorsal surface spiculose There has been some confusion in the lit- on segments VI-VIII, with sclerite on seg- erature concerning the publication date of ment VII-VIII; marginal abdominal tuber- D. agropyronophaga (Zhang 1991). The cles and supracaudal process absent. Cor- date was listed by Remaudiere and Remau- nicle (Fig. 2B) pigmented, short, 0.030 diere (1997) as 1990, the date that appears long, apical flange undeveloped; without in the title of Zhang's (1991) publication. associated basal sclerite. Cauda (Fig. 2C) However, this work was not published until 0.138 long, elongate, triangular, with slight May 1991 (as printed on the publication). medial constriction, 6-7 lateral setae. A description of D. agropyronophaga was Alata: Body length 1.416; width through also listed as "sp. nov." in Zhang et al. eyes 0.390-0.342. Antenna (Fig. 2D) short- (1991), but, because the actual time of is- er than body; segmentIII 0.246-0.228 long, suance has not been determined, that work with 6-7 secondary sensoria restricted to would be listed as the last day in 1991. — approximately half circumference of seg- Specimens examined. CHINA: Ni- ment; IV, 0.102 long, with 2-4 secondary ngxia, exAgropyron sp., IZAS: 16-VI-1990 sensoria; V, 0.090-0.096 long, without sec- (2 ap. paratypes, 2 al. paratypes, 2 alatoid ondary sensoria; base of VI, 0.102-0.114 imm. labeled "PARATYPES"). long; terminal process, 0.138 long. Dorsal head setae longer than width of antennal Diuraphis agrostidis (Muddathir) segment III. Rostral length and setae similar (Fig. 3) to apterous female, ultimate segment Holcaphis agrostidis Muddathir 1965:477. 0.102-0.108 long with 2 accessory setae. Diuraphis {Holcaphis) agrostidis: Eastop Pronotum without marginal tubercles. Hind and Hille Ris Lambers 1976:175; Remau- tibia 0.600-0.618 long; hind tarsus II diere and Remaudiere 1997:91. 0.126-0.132 long. Abdominal tergum with- — out patches or bands, surface sculpturing Type material. Holotype, morphotypes, BMNH similar to aptera, abdominal tergite VIII and paratypes deposited in (Mud- with 3-5 setae; abdominal marginal tuber- dathir 1965); not seen. — cles and supracaudal process absent. Cor- Field features. Aptera yellow green nicle short, smooth, 0.024 long, similar to with white powder; head dark, almost aptera. Cauda 0.120-0.120 long, similar to black; antennae, legs, and cauda dark (Heie aptera, with 6 lateral setae. 1992). — Notes. Although Zhang (1991) illus- Recognition characters. Aptera: Body trates D. agropyronophaga with a slightly length 1.470-2.100; width through eyes raised area on abdominal tergite VIII, this 0.318-0.384. Antenna (Fig. 3A) shorter structure was not discemable on the speci- than body; segment III 0.090-0.162 long; mens we examined. IV 0.060-0.084 long; V 0.072-0.084 long; Diuraphis agropyronophaga resembles base of VI 0.066-0.084 long; terminal pro- D. tritici. However, the ultimate rostral seg- cess 0.102-0.120 long. Head sclerotized, ment is shorter than D. tritici (approximate- smooth, without spinulation; longest dorsal ly 2 times as long as wide at the base versus head setae longer than width of antennal approximately 3 times as long as wide at segment III. Rostrum ending before meso- the base for D. tritici). Also, in D. agro- coxae; ultimate segment 0.066-0.084 long, pyronophaga, the length of antennal seg- approximately 1.5 times as long as wide at ment III is shorter than antennal segments base, shorter than hind tarsal segment II, — — VOLUME NUMBER 107, 3 709 without accessory setae. Pronotum without host of D. agrostidis is Agrostis stolonifera marginal tubercles. Hind tibia .390-0.420 L. (Muddathir 1965, Heie 1992) whereas long; hind tarsus II 0.108-0.120 long. Ab- the host ofD. hold are Holcus spp. See also domen smooth with ventral surface spicu- Notes section for D. bromicola, D. fre- lose, dorsal surface spiculose on segments qiiens, and D. hold. — VI-VIII, with small pleural and interseg- Specimens examined. UNITED KING- mental sclerites, large sclerites on segments DOM: SCOTLAND: Frazerburgh. ex grass, V-VIII; abdominal marginal tubercles and H.L.G.S. coll., BMNH: 13-VIII-1959 (8 supracaudal process absent. Cornicle (Fig. ap., 2 imm.). ENGLAND: Northumberland, 3B) slightly pigmented, short, 0.018-0.024 Newcastle, King's College, ex Agrostis sto- long, apical flange undeveloped; with as- lonifera, K. Muddathir coll., BMNH: 8-?- sociated basal sclerite. Cauda (Fig. 3C) 1963 (2 al.). 0.120-0.144 long, elongate, triangular with rounded apex, 6 lateral setae and usually Diuraphis bromicola (Hille Ris Lambers) one preapical seta. (Fig. 4) Alata: Body length 1.860-1.902; width Holcaphis bromicola Hille Ris Lambers through eyes 0.360-0.372. Antenna (Fig. 1959:281. 3D) shorter than body; segment III 0.192- Diuraphis {Holcaphis) bromicola: Eastop 0.198 long, with 4-5 secondary sensoriare- and Hille Ris Lambers 1976:175; Remau- stricted to approximately half circumfer- diere and Remaudiere 1997:91. ence ofsegment; IV 0.078-0.102 long, with — 0-1 secondary sensoria; V 0.090-0.102 Type material. Two cotype slides seen. long, without secondary sensoria; base of One slide with left label "'Holcaphis brom- VI 0.102-0.108 long; terminal process icola nov. spec cotypes Det. D.H.R.L. " and 0.192-0.210 long. Dorsal head setae sube- right label "N. Germany PI. Bromus iner- qual to width antennal segment III. Rostral mis Loc. Leipzig Date Oct. 1956 Leg. length and setae similar to apterous female, Muhle BM1984-340 "(BMNH). Another ultimate segment 0.060-0.072 long without slide with left label "'Holcaphis bromicola accessory setae. Pronotum without margin- nov. spec cotypes Det. D.H.R.L." and right al tubercles. Hind tibia 0.540-0.636 long; label "N. Germany PI. Bromus inermis hind tarsus II 0.108-0.1 14 long. Abdominal Loc. Leipzig Date 20-V1-1957 Leg. Miihle surface sculpturing and sclerotization simi- BM1984-340 "(BMNH). Cotypes of D. lar to aptera, abdominal tergite VIII with 3- bromicola were originally deposited in the 5 setae; abdominal marginal tubercles and collection of Hill Ris Lambers (1959) but BMNH. supracaudal process absent. Cornicle short, are presently in — smooth, 0.018 long, similar to aptera. Cau- Field features. Oviparous female very da 0.120 long, similar to aptera. light green with grey wax-powder; alata Notes. Diuraphis agrostidis most similar to ovipara but darker with less pov\- closely resembles D. hold. Apterae and ala- der, head and thorax bhiLk (Hille Ris 1.amb- tae of D. agrostidis have intersegmental ers 1959). sclerites that are larger than the adjacent Recognition characters. Aptera: Body spiracle and associated sclerotized area, and length 1.950-2.100; width through eyes antennal segment III is usually shorter than 0.348-0.360. Antenna (Fig. 4A) shorter antennal segments IV + V. Conversely, ap- than body; segment 111 0.204-0.246 long; terae and alatae of D. hold have interseg- IV 0.096-0.108 long; V 0.090-0.102 long: mental sclerites that are subequal or smaller base of VI 0.096-0.108 long; terminal prin- than adjacent spiracle and associated scler- cess, 0.096-0.126 long. Head sclcroti/ed. otized and antennal segment III is usually smooth, vviihoui spiniilatiiMi; longest dorsal longer than antennal segments IV + V The head setae shoriL-r than width nt" antennal