A systematic reappraisal of the genus Diuraphis Aizenberg (Gemiptera: Aphididae)

2005·14.2 MB·

by  Gary L Miller

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