[Back to North American Cixiidae]

Family Cixiidae Spinola, 1839

Subfamily Cixiinae Spinola, 1839

Tribe Oecleini Muir, 1922

Genus Haplaxius Fowler, 1904: 97.

Type species: (in original combination): Haplaxius laevis Fowler, 1904: 98.

Synonyms

= Paramyndus Fennah, 1945a (Type species Paramyndus cocois Fennah, 1945a; junior syn. of Myndus crudus Van Duzee 1907); syn. with Myndus by Kramer, 1979: 303-304.
= Myndus Stål, 1862a (auctorum; Type species Flata musiva Germar 1825); syn. by Kramer 1979: 303; removed from syn. by Emeljanov 1989: 62-64; also Holzinger et al. 2002: 128.

Distribution

Widespread in New World.

Recognized species

There are 34 species north of Mexico, plus an additional 30 species in the Neotropics. A related genus (Myxia, see FLOW) was recently  described.

1 Haplaxius balli (Kramer, 1979) – USA: AZ
= Myndus balli Kramer, 1979: 325-326.
= Haplaxius balli (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
2 Haplaxius beameri (Ball, 1933c) – USA: AZ
= Myndus beameri Ball, 1933c: 480.
= Haplaxius beameri (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
3 Haplaxius catalinus (Ball, 1933c) – USA: AZ, NM, TX
= Myndus catalinus Ball, 1933c: 481.
= Haplaxius catalinus (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
4 Haplaxius collinus (Ball, 1933c) – USA: CO, MT; CAN: SK
= Myndus collinus Ball, 1933c: 482.
= Haplaxius collinus Ball, 1933c; comb. by implication Emeljanov 1989: 62.
5 Haplaxius crena (Kramer, 1979) – USA: AZ, CO
= Myndus crena Kramer, 1979: 318-320.
= Haplaxius crena Kramer, 1979; comb. by implication Emeljanov 1989: 62.
6 Haplaxius crudus (Van Duzee, 1907), the “American palm cixiid”  – USA: FL, TX, MS; Belize, Brazil, Cayman Islands (Grand Cayman), Columbia, Costa Rica, Cuba (inc. Isle of Pines), Dominican Republic, Honduras, Jamaica, Mexico (Baja California Sur, Quintana Roo, Yucatán), Panama, Puerto Rico, Venezuela, Trinidad
= Myndus crudus Van Duzee, 1907: 33.
= Paramyndus crudus (Van Duzee, 1907); comb. by implication Fennah 1945b: 142.
= Paramyndus cocois Fennah, 1945a: 424-425.
= Haplaxius crudus (Van Duzee, 1907); comb. by Caldwell 1946: 203; also Emeljanov, 1989: 62.
= Haplaxius cocois (Fennah, 1945a); comb. by Caldwell 1946: 203.
= Haplaxius pallidus Caldwell, 1946: 204; syn. by Kramer 1979: 346-347.
= Haplaxius cocois (Fennah, 1945a); syn. by Kramer 1979: 346; also Emeljanov 1989: 62.
7 Haplaxius dozieri (Kramer, 1979) – USA: MS
= Myndus dozieri Kramer, 1979: 340.
= Haplaxius dozieri (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
8 Haplaxius enotatus (Van Duzee, 1909) – USA: FL, GA, LA, MD, MS, NC, SC, VA; CAN: QC; Cuba
= Myndus enotatus Van Duzee, 1909: 188.
= Haplaxius enotatus (Van Duzee, 1909); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
9 Haplaxius flocki (Kramer, 1979) – USA: AZ, CA; Mexico (Baja California)
= Myndus flocki Kramer, 1979: 342-344.
= Haplaxius flocki (Kramer, 1979); comb. by implication Emeljanov, 1989: 62.
10 Haplaxius fulvus (Osborn, 1903a) – USA: CT, IL, KS, KY, MA, NY, OH, TN; CAN: ON
= Myndus fulvus Osborn, 1903a: 46.
= Haplaxius fulvus (Osborn, 1903a); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
11 Haplaxius gabrielensis Flock, 1951- USA: CA
= Haplaxius gabrielensis Flock, 1951: 169.
= Myndus gabrielensis (Flock, 1951); comb. by Kramer 1979: 328, 330; restored to Haplaxius by Emeljanov 1989: 62.
12 Haplaxius glyphis (Kramer, 1979) – USA: MA, NY, TN; CAN: NB, QC
= Myndus glyphis Kramer, 1979: 311-312.
= Haplaxius glyphis (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
13 Haplaxius impiger (Ball, 1902a) – USA: CO, NM
= Myndus impiger Ball, 1902a: 153.
= Haplaxius impiger (Ball, 1902a); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
14 Haplaxius lophion lophion (Kramer, 1979) – USA: AZ, CO
= Myndus lophion lophion Kramer, 1979: 349-350 (subspecific combination by implication).
= Haplaxius lophion lophion (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
14a Haplaxius lophion alpha (Kramer, 1979) – USA: CO
= Myndus lophion alpha Kramer, 1979: 350.
= Haplaxius lophion alpha (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
15 Haplaxius lunatus (Van Duzee, 1909) – USA: FL
= Myndus lunatus Van Duzee, 1909: 189.
= Haplaxius lunatus (Van Duzee, 1909); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
16 Haplaxius mojavensis (Ball, 1933c) – USA: AZ, CA, NV
= Myndus mojavensis Ball, 1933c: 480.
= Haplaxius mojavensis (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
17 Haplaxius neopusillus (Kramer, 1979) – USA: FL
= Myndus neopusillus Kramer, 1979: 338-339.
= Haplaxius neopusillus (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
18 Haplaxius nevadensis (Kramer, 1979) – USA: AZ, NV
= Myndus nevadensis Kramer, 1979: 325.
= Haplaxius nevadensis (Kramer, 1979); comb. by implication Emeljanov, 1989: 62.
19 Haplaxius nigrifrons (Ball, 1937) – USA: TX
= Myndus nigrifrons Ball, 1937: 179.
= Haplaxius nigrifrons (Ball, 1937); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
20 Haplaxius nolinus (Ball, 1933c) – USA: AZ, NM
= Myndus nolinus Ball, 1933c: 481.
= Haplaxius nolinus (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
21 Haplaxius occidentalis (Van Duzee, 1914) – USA: AZ, CA, ID, OR, UT
= Myndus occidentalis Van Duzee, 1914: 39.
= Haplaxius occidentalis (Van Duzee, 1914); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
22 Haplaxius ovatus (Ball, 1933c) – USA: DE, GA, IA, KS, MA, MD, NE, NJ, OK, VA
= Myndus ovatus Ball, 1933c: 483.
= Haplaxius ovatus (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
23 Haplaxius pictifrons (Stål, 1862a) – USA: CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, LA, MA, MD, MO, MS, NC, NE, NH, NJ, NY, OH, OK, PA, TN, VA; CAN: ON, QC; Cuba
= Myndus pictifrons Stål, 1862a: 307.
= Haplaxius pictifrons (Stål, 1862a); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
= Myndus sordipennis Stål, 1862a: 308.
= Haplaxius sordipennis (Stål, 1862a); comb. by Caldwell 1946: 203; syn. by Kramer 1979: 314, 316.
24 Haplaxius pusillus (Van Duzee, 1909) – USA: CT, FL, GA, MA, NC, NH, NJ, VA
= Myndus pusillus Van Duzee, 1909: 190.
= Haplaxius pusillus (Van Duzee, 1909); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
= Haplaxius perrinei Caldwell, 1951: 34; syn. by Kramer 1979: 336, 338.
25 Haplaxius radicis (Osborn, 1903b) – USA: DC, DE, IL, IN, KS, MD, MS, NC, NY, OH, OK, TN, TX, VA
= Myndus radicis Osborn, 1903b: 42.
= Haplaxius radicis (Osborn, 1903b); comb. by Caldwell 1946: 203.
= Myndus delicatus Van Duzee, 1908: 492.
= Haplaxius delicatus (Van Duzee 1908); comb. by Caldwell 1946: 203; syn. by Kramer 1979: 320-321.
26 Haplaxius rubidus (Ball, 1933c) – USA: TX
= Myndus rubidus Ball, 1933c: 483.
= Haplaxius rubidus (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
27 Haplaxius slossonae (Ball, 1902a) – USA: FL, GA, KS, MO, MS, OK, TX
= Myndus slossoni Ball, 1902a: 154.
= Haplaxius slossoni (Ball, 1902a); comb. by Caldwell 1946: 203.
= Myndus slossonae (Ball, 1902a); emendation to by Kramer 1979: 334, 336.
= Haplaxius slossonae (Ball, 1902a); comb. by implication Emeljanov 1989: 62.
28 Haplaxius texensis (Kramer, 1979) – USA: FL, GA, KS, MO, MS, OK, TX
= Myndus texensis Kramer, 1979: 333.
= Haplaxius texensis (Kramer, 1979); comb. by implication Emeljanov 1989: 62. USA: TX
29 Haplaxius truncatus (Metcalf, 1923) – USA: IA, IL, WI
= Myndus truncatus Metcalf, 1923: 184.
= Haplaxius truncatus (Metcalf, 1923); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
30 Haplaxius viridicatus (Ball, 1933c) – USA: AZ, CA
= Myndus viridicatus Ball, 1933: 483.
= Haplaxius viridicatus (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
31 Haplaxius viridis (Ball, 1902a) – USA: AZ, CA, CO (reported in error: USA: IA, NJ, OH)
= Myndus viridis Ball, 1902a: 153.
= Haplaxius viridis (Ball, 1902a); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.
= Myndus auratus Ball, 1933c: 434; syn. by Caldwell 1946: 203.
= Haplaxius auratus (Ball, 1933c); comb. by Kramer 1979: 340, 342.
32 Haplaxius wheeleri (Wilson, 1996) (in Wheeler & Wilson 1996) – USA: NJ, PA
= Myndus wheeleri Wilson, 1996: 103-104 (in Wheeler & Wilson 1996).
= Haplaxius wheeleri (Wilson, 1996); comb. by Bartlett et al. 2011: 12.
33 Haplaxius xyron (Kramer, 1979) – USA: MI, NY, PA; CAN: ON, QC
= Myndus xyron Kramer, 1979: 312-314.
= Haplaxius xyron (Kramer, 1979); comb. by implication Emeljanov 1989: 62.
34 Haplaxius yuccandus (Ball, 1933c) – USA: AZ
= Myndus yuccandus Ball, 1933c: 482.
= Haplaxius yuccandus (Ball, 1933c); comb. by Caldwell 1946: 203; also Emeljanov 1989: 62.

Neotropical species (from an unpublished list by Lois O’Brien)
1 Haplaxius akko (Kramer 1979:372) – Belize, Costa Rica, Mexico, Nicaragua, Panama
2 Haplaxius brimosis (Kramer 1979:366) – Bolivia
3 Haplaxius caldwelli (Kramer 1979:359) – Guatemala, Honduras, Mexico
4 Haplaxius cabrerensis Ferreira, McKamey, & Sanchez, 2010 – Dominican Republic
   Haplaxius crudus (Van Duzee, 1907: 33) – see above
5 Haplaxius deleter (Kramer 1979: 379) – Panama
Haplaxius delta (Kramer 1979: 376) – Belize, Costa Rica, Guatemala, Panama (to Myxia by Echavarria et al. 2021)
6 Haplaxius dolon (Kramer 1979: 384) – Argentina, Brazil, Uruguay
7 Haplaxius dougwalshi Bahder & Bartlett 2020 – Costa Rica
8 Haplaxius fennahi (Kramer 1979: 382) – Peru
9 Haplaxius frontalis Fowler 1904: 98) – Mexico (Guerrero)
10 Haplaxius gnophos (Kramer 1979: 366) – Brazil, Bolivia, Paraguay, Argentina
11 Haplaxius gomphos (Kramer 1979: 374) – Mexico
12 Haplaxius hochae O’Brien 2006: 313) – Dominica
13 Haplaxius jamaicae (Kramer 1979: 355) – Jamaica, Dominican Republic
14 Haplaxius laevis Fowler 1904: 98 – Mexico (Mexico City)
15 Haplaxius lyssa (Kramer 1979: 364) – Peru, Ecuador, Brazil
16 Haplaxius meadi (Kramer 1979: 388) – Mexico
17 Haplaxius mokos (Kramer 1979: 363) – Peru
18 Haplaxius nimbus (Kramer 1979: 368) – Venezuela
19 Haplaxius pocococo Bahder & Bartlett 2021 (in Barrantes et al. 2021) – Costa Rica
20 Haplaxius phylax (Kramer 1979: 362) – Costa Rica
21 Haplaxius serratus (Caldwell 1946: 204) – Mexico
22 Haplaxius sillos (Kramer 1979: 363) – Peru
23 Haplaxius simplicatus Caldwell, 1946: 204 – Costa Rica, El Salvador, Panama
24 Haplaxius skarphion (Kramer 1979: 386) – Belize, Costa Rica, Guatemala, Honduras, Mexico, Panama
25 Haplaxius spanglerorum (Kramer 1979: 380) – Peru
26 Haplaxius sparagma (Kramer 1979: 374) – Guatemala
27 Haplaxius synavei (Kramer 1979: 378) – Venezuela
28 Haplaxius tekmar (Kramer 1979: 368) – Mexico
29 Haplaxius tekton (Kramer 1979: 382) – Peru
30 Haplaxius thryligma (Kramer 1979: 372) – Brazil
31 Haplaxius vilbastei (Kramer 1979: 379) – Argentina

Economic Importance

Haplaxius crudus (Van Duzee 1907) is usually considered a vector of Lethal Yellowing of coconut palms in Florida (e.g., Howard et al. 1983; Howard 1987, here; although some doubt this because the transmission was not subsequently replicated, but see Dzido et al. 2020), and almost certainly elsewhere. It (and perhaps other species) may be vectors of other diseases in coconut, oil, and ornamental palms.  Haplaxius crudus appears to be the vector of lethal browning (Texas Phoenix palm decline) and possibly foliar decay in palms. Haplaxius crudus has been repeatedly reported in association with palm disease vector surveys.

Pathogens/parasites

Candidatus Phytoplasma palmae [Palm lethal yellowing – 16srIV-A] (Bacteria, Mollicutes, Acholeplasmatales, Acholeplasmataceae) (Howard et al. 1983)

Metarhizium anisopliae (Metchnikoff), Ascomycota: Clavicipitaceae (entomopathogenic fungi) (Guerrero et al. 2020)

Plant associations

• Haplaxius beameri – Agave americana L. (American century plant, Agavaceae), Yucca (Agavaceae)
• Haplaxius catalinus – Yucca
• Haplaxius crudus – (apparently polyphagous on palms) Cocos nucifera L. (coconut palm, Arecaceae), Cynodon dactylon (L.) Pers. (Bermudagrass, Poaceae), Elaeis guineensis Jacq. (Arecaceae), Eragrostis curvula (Schrad.) Nees (Poaceae),  Panicum maximum Jacq. (Poaceae), Paspalum notatum Flügge (bahiagrass, Poaceae), Phoenix dactylifera L. (Canary Island date palm, Arecaceae), Pritchardia pacifica Seem. & H.Wendl (Fiji fan palm, Arecaceae), Sabal palmetto (Walter) Lodd. ex Schult. & Schult. f. (Arecaceae), Stenotaphrum secundatum (Walter) Kuntze (St. Augustine grass, Poaceae), Thrinax radiata Lodd. ex Schult. and Schult. f (Florida thatch palm) (Howard & Wilson 2001, Wilson & Wheeler 2010, Wilson et al. 2014, Hill et al. 2018, Bartlett et al. 2018, Narvaez et al. 2018)
• Haplaxius dougwalshi – Asterogyne martiana (H. Wendl.) H. Wendl. ex Hemsl. (Asteraceae) Bahder et al. (2020: 549)
• Haplaxius enotatus – grass, Juncus (Rush, Juncaceae), ‘marsh grasses’
• Haplaxius gabrielensis – Hesperoyucca whipplei (Torr.) Baker (chaparral yucca, Agavaceae), sedges
• Haplaxius glyphis – Populus (Salicaceae)
• Haplaxius hochae – Euterpe globosa Gaertner (Arecaceae) (O’Brien 2006: 313)
• Haplaxius lophion – Pinus (Pinaceae, Pine)
• Haplaxius mojavensis – Yucca brevifolia Engelm. (Joshua tree, Agavaceae)
• Haplaxius nigrifrons – Nolina Michx. (beargrass, Liliaceae)
• Haplaxius nolinus – Nolina microcarpa S. Watson (sacahuista, Liliaceae), Dasyochloa pulchella (Kunth) Willd. ex Rydb. (low woollygrass, Poaceae, as Triodia pulchella)
• Haplaxius occidentalis – Juncus, Washingtonia filifera (Linden ex André) H. Wendl.(California fan palm, Arecaceae)
• Haplaxius ovatus – Panicum virgatum L. (Poaceae) (Wheeler & Wilson 2014: 359)
• Haplaxius pictifrons – Phalaris arundinacea L. (reed canarygrass, Poaceae), grasses (streamside), Salix sp. (willow, Salicaceae)
• Haplaxius pocococo – coconut palm (Cocos nucifera L.), Arecaceae.
• Haplaxius radicis – Impatiens L. (touch-me-not, Balsaminaceae), grasses, nettles
• Haplaxius rubidus – “Palmetto” (presumably Sabal palmetto (Walter) Lodd. ex Schult. & Schult. f. (cabbage palmetto, Arecaceae))
• Haplaxius simplicatus – Urochloa mutica (Forssk.) T.Q. Nguyen (para grass, as Panicum barbinode)
• Haplaxius skarphion – Cocos nucifera L. (Arecaceae) (Bahder et al. 2020: 549)
• Haplaxius slossonae – Juncus, ‘grasses’
• Haplaxius tekmar – Agave
• Haplaxius texansis – Yucca thompsoniana Trel. (Thompson’s yucca)
• Haplaxius wheeleri – Pinus rigida (Pitch pine)
• Haplaxius yuccandus – Yucca

Hosts from Wilson et al. 1994, Howard & Wilson 2001, O’Brien 2006, Wilson & Wheeler 2010, Wilson et al. 2014, Hill et al. 2018, Bartlett et al. 2018, Narvaez et al. 2018, Bahder et al. 2020; plant names from USDA PLANTS or Tropicos.

Additional hosts listed in FLOW here.

Recognition

Without spines on hind tibiae before apex (a tribal feature), tegulae evident, carina on longitudinal midline (median carina) of frons present; vertex without distinct carina on both longitudinal midline and between anterior portions of eyes; longitudinal midlength of mesonotum about 2x or less longitudinal midlength of vertex.

In 2019, Bahder et al described the genus Myxia Bahder & Bartlett, 2019, which may be diagnosed from Haplaxius by the phallobase subtending aedeagus (not forming a phallotheca); parameres distally bifid in lateral view (with inner lateral tooth near apex); anal tube short and stout, distally projected downward. Myxia appears to be distinctively sexually dimorphic.  A description of a second species Myxia should be published soon, and at least one species currently in Haplaxius appears to belong to Myxia.

Fowler 1904: 97-98 description

Capite modico, subrotundato, vertice latitudine evidenter longiori, ad basim haud emarginato, ante oculos haud vel vix producto, laevi; fronte fere seque longa ac lata, subrotundata, a clypeo sulco sat profundo separata, hoc ad latera carinato; ocello frontali interdum vix distincto; pronoto brevi ; scutello tricarinato, carinis plerumque indistinctis ; tegminibus pellucidis, haud vel vix evidenter vel delicatissime granulatis, vena radiali et vena ulnari interiori paullo pone medium aequidistanter furcatis, areis apicalibus novem vel decern; tibiis simplicibus. [Google translate: Short head, subrotundata, top five clearly extends to the base and not emarginated before eyes or a little long, smooth; the front of the long and wide, about himself, orbicular, separated from the shield to the bottom of a ditch they sat, with a carina on the sides of this have to do; at times scarcely an eye of a distinct of the frontal; slightly short; scutellum very tricarinate, the keels for the most part, indistinct; strong lobe, not even barely feed or clearly granulate radial and ulnar vein set an equal inward slightly forked tip areas, nine or ten; three simple.]

Head subrotundate, with the vertex long and smooth, and not emarginate at the base, and not or scarcely produced before the eyes; forehead about as long as broad, roughly subrotundate, separated from the clypeus by a distinct furrow, clypeus carinate at the sides; pronotum short; scutellum with three indistinct carinae; tegmina with the radial and interior ulnar veins forked at about an equal distance, slightly behind the middle, the granulation not or scarcely evident; tibiae simple.

Haplaxius skarphion from Bahder et al 2020

Key to US species from Kramer 1979 (figure references are to Kramer 1979, but I have posted most of them below; this key has had Haplaxius wheeleri inserted in the couplet with  pusillus and neopusillus).

1. Frons with distinct transverse black or brown markings at both base and apex … 2

1.-Frons unmarked or not marked as above …. 9

Haplaxius crudus and H. pictifrons

2. Apex of aedeagus prolonged and acute (figs. 10, 11) … truncatus (USA: IA, IL, WI)

2.- Apex of aedeagus neither prolonged nor acute … 3

3. Left apical appendage of aedeagus subdivided distally into blades (fig. 13) or with acute, posteriorly directed tooth on ventral margin near the base (figs. 16,23) … 4

3.- Left apical appendage of aedeagus not so modified … 6

4. Left apical appendage of aedeagus without posteriorly directed tooth (figs. 13, 14) … glyphis (USA: MA, NY, TN; CAN: NB, QC)

4.- Left apical appendage of aedeagus with acute posteriorly directed tooth on ventral margin near base … 5

5. Left apical appendage of aedeagus distally subdivided into two acute portions (figs. 16, 17) … xyron (USA: MI, NY, PA; CAN: ON, QC)

5.- Left apical appendage of aedeagus not subdivided as above (figs. 22,23) … pictifrons (Widespread E USA & Canada)

Hyplaxius xyron from Kramer 1979

6. Left apical appendage of aedeagus broadly rounded distally (fig. 29) or dorsal margin of aedeagus strongly excavated in the basal half (fig. 27) … 7

6.- Left apical appendage of aedeagus produced and tapered distally (fig. 36); dorsal margin of aedeagus not excavated … 8

7. Left apical appendage of aedeagus produced as blunt finger distally, dorsal aedeagal margin strongly excavated (figs. 27,28) … impiger (USA: CO, NM)

7.- Left apical appendage of aedeagus broadly rounded distally, dorsal aedeagal margin not excavated (figs. 29, 30) … crena (USA: AZ, CO)

Hyplaxius impiger from Kramer 1979

8. Anal flap slender and almost J-shaped (fig. 32); left apical appendage of aedeagus quadrate or subquadrate basally (figs. 32, 35-37); posterior margin of pygofer obtusely angled (fig. 32) … radicis (in part) (Widespread E USA)

8.- Anal flap moderately stout and not at all I-shaped (fig. 38); left apical appendage of aedeagus oval or suboval basally (fig. 38); posterior margin of pygofer broadly rounded (fig. 38) … collinus (USA: CO, MT; CAN: SK)

9. Anal flap long with distal portion broadly downturned (fig. 32); left apical appendage of aedeagus quadrate or subquadrate basally and extended as blunt fingerlike lobe distally (figs. 32, 35-37) … radicis (in part) (Widespread E USA)

9.- Anal flap and left apical appendage not shaped as above … 10

10. Anal flap strongly asymmetrical, left margin with subapical lobe and right margin with a decurved process (fig. 41); ventral margin of aedeagus with two basally directed subtriangular processes (fig. 42) (known only from Florida) … lunatus (USA: FL)

10.- Male genitalia without either of the above features … 11

11. Aedeagus in lateral view tending to roundness, its ventral margin produced as spine directed obliquely ventrocephalad (figs. 44, 66) (exception nigrifrons where spine directed caudad, fig. 61); right side of aedeagus with two curved or partly curved contiguous or nearly contiguous processes originating near the apex (figs. 52, 64, 70) (Southwestern) … 12

11.- Aedeagus in lateral view not tending to roundness, its ventral margin simple or not modified as above; right side of aedeagus with zero to two processes, their features not as above … 21

12. Both pro- and mesonotum unicolorous dark fuscous to black … 13

12.- Both pro- and mesonotum not as above … 15

13. Claval areas of forewings entirely dark … beameri (USA: AZ)

13.- Claval areas of forewings clear hyaline except at extreme bases … 14

14. Forewings with discal embrowning not reaching claval vein; ventral margin of aedeagus strongly convex (fig. 48); left aedeagal process short and simple (fig. 47) … nevadensis (USA: AZ, NV)

14.- Forewing with discal embrowning reaching claval vein at least in distal half; ventral margin of aedeagus not convex (fig. 50); left aedeagal process elongate with surface scabrous (fig. 49) … balli (USA: AZ)

15. Frons and clypeus concolorous honey brown, neither with distinct markings … 16

15.- Frons and clypeus are either not concolorous or frons with darker or lighter linear markings … 17

16. Clavus without dark markings; ventral margin of aedeagus unlobed with spine small (figs. 51, 52) … nolinus (USA: AZ, NM)

16.- Clavus with dark markings; ventral margin of aedeagus bilobed and with spine large (figs. 54, 55) … yuccandus (USA: AZ)

17. Clavus with transverse or round dark markings; the line formed by anterior edge of ventral aedeagal spine and aedeagus not concave (figs. 57, 58) … gabrielensis (USA: CA)

17.- Clavus either unmarked or with only longitudinal embrowning along commissural margin; line formed by anterior edge of ventral aedeagal spine and aedeagus concave … 18

18. Frons and clypeus black or dark fuscous, frons with a pale stripe on each side originating at the lateroventral margin and obliquely extended toward median carina; left ventral margin of anal tube lobed in lateral view (fig. 60), right ventral margin simple; ventral aedeagal spine directed caudad in lateral view (figs. 60, 61) … nigrifrons (USA: TX)

18.- Frons lighter in color than clypeus, frons with a dark stripe on each side originating at the apex of frons and extended ventrally inside of lateral frontal margin; both margins of the anal tube the same, ventral aedeagal spine directed cephalad in lateral view … 19

19. Posterior tip of mesonotum black; forewings usually narrowly embrowned either in part or entirely along commissural margin; aedeagus in left lateral view with well-developed lobe near ventral margin distally (fig. 63); ventral aedeagal spine with base broadly triangular and apex comparatively blunt (fig. 64) … catalinus (USA: AZ, NM, TX)

19.- Posterior tip of mesonotum pale; forewings neither embrowned nor darkened along commissural margin; aedeagus in left lateral view with a poorly developed lobe or no lobe near ventral margin distally; ventral aedeagal spine with base less or not broadly triangular and apex acute … 20

20. Crossveins at the level of stigma to claval apex entirely pale or white; ventral aedeagal spine with base broad (figs. 66, 67); ventral margin of anal flap straight or slightly concave (fig. 66) … mojavensis (USA: AZ, CA, NV)

20.- Crossveins at the level of stigma to claval apex only partly pale or white; ventral aedeagal spine with base narrow (fig. 70); ventral margin of anal flap convex beyond middle (fig. 69) … texensis  (USA: FL, GA, KS, MO, MS, OK, TX)

21. Aedeagus slender, in lateral view with pair of long processes originating near the apex and extending cephalad; aedeagus in right lateral view with acute process near apex projecting distally; each forewing distinctly paler along costal margin … 22

21.- Aedeagus without three processes as above; each forewing not paler along costal margin … 23

22. Forewings transparent stramineous to pale brown; paired dorsal processes of aedeagus about equal in length and stoutness (figs. 72, 73); anal flap subquadrate in lateral view (fig. 72) … enotatus (USA: FL, GA, LA, MD, MS, NC, SC, VA; CAN: QC; Cuba)

22.- Forewings transparent fuscous at least basally and apically; paired dorsal processes of aedeagus clearly unequal in length and stoutness (figs. 75, 76); anal flap apically decurved in lateral view (fig. 75) … slossonae (USA: FL, GA, KS, MO, MS, OK, TX)

Haplaxius enotatus from Kramer 1979

23. Aedeagus in right lateral view with single upright or slightly curved process originating below dorsal margin, dorsal margin with or without short tooth (figs. 79, 82, 85, 91, 94) … 24

23.- Aedeagus in right lateral view without process or processes as above, dorsal margin with long tooth (fig. 97), basally directed process (fig. 103), or simple … 29

24. Pygofer in lateral view with posterior margin strongly triangular … 25

24.- Pygofer in lateral view with the posterior margin not strongly triangular, usually roundly produced … 26

25. A single process from the right side of aedeagus projecting much beyond the dorsal edge of aedeagus, ventral process on aedeagal shaft lacking lacking (fig. 79, see wheeleri below), a process not originating from flange (fig. 79) … pusillus (USA: CT, FL, GA, MA, NC, NH, NJ, VA)

25.- Single process from the right side of aedeagus not reaching the dorsal edge of aedeagus (fig. 82), a process originating from flange (fig. 82) … neopusillus (USA: FL)

25′.- Process from the right side of aedeagus exceeding the dorsal margin of aedeagus; a strong, acute process present on the ventral aspect of the aedeagal shaft and a serrate region anterior to this process … wheeleri (on pine, USA: NJ, PA)

26. Toothlike projection on the dorsal margin of aedeagus in lateral view either positioned close to the apex or absent … 27

26.- Toothlike projection on the dorsal margin of aedeagus in lateral view near middle … 28

27. Without toothlike projection on the dorsal margin of the aedeagus (figs. 84, 85); posterior lateral angles of pregenital ventral abdominal segments are almost always marked with black (extremely common in Calif. and other western states … occidentalis (USA: AZ, CA, ID, OR, UT)

27.- With toothlike projection on the dorsal margin of aedeagus near the apex (figs. 87, 88); ventral abdominal segments unicolorous (known only from Miss.) … dozieri (USA: MS)

28. A single process on right side of aedeagus upright and slender (fig. 91); triangular projection from venter of the anal flap in the basal half (fig. 90) … viridis (USA: AZ, CA, CO (reported in error: USA: IA, NJ, OH))

28.- Single process on right sides of aedeagus tilting distally and broader (fig. 94); triangular projection from venter of anal flap at or slightly beyond middle (fig. 93) … flocki (USA: AZ, CA; Mexico (Baja California))

29. Dorsal margin of aedeagus with moderately long, toothlike, posteriorly directed projection at or near the apex (figs. 96, 97) … viridicatus (USA: AZ, CA)

29.- Dorsal margin of aedeagus without such a projection …30

30. Anal flap with process originating between ventral margins … 31

30.- Anal flap without process or process originating from ventral margin or margins … 32

31. Longer aedeagal process directed cephalad and parallel or nearly parallel with the long axis of aedeagus (figs. 100, 103); the ventral process of the anal flap irregularly rounded or blunt distally (figs. 99, 102) … ovatus (USA: DE, GA, IA, KS, MA, MD, NE, NJ, OK, VA)

31.- Longer aedeagal process directed ventrocephalad and projecting beyond the ventral margin of aedeagus (figs. 106, 108, 109); the ventral process of theanal flap subtriangular (fig. 105) … crudus (USA: FL, TX, MS; Mesoamerica, Caribbean, northern South America)

32. Aedeagus with two short processes on the right side (figs. Ill, 112); anal flap quadrate (fig. 110) … fulvus (USA: CT, IL, KS, KY, MA, NY, OH, TN; CAN: ON)

32.- Aedeagus without processes on the right side; anal flap not quadrate  … 33

33. Anal flap with a long triangular process at the middle of right ventral margin (fig. 113); apex of aedeagal process footshaped in ventral view (fig. 115) … rubidus (USA: TX)

33.- Anal flap with right margin at most broadly angular (fig. 116) or rounded; apex of aedeagal process tapered to acute tip (fig. 118) … lophion (USA: AZ, CO)

KEY TO MEXICAN AND NEOTROPICAL SPECIES OF HAPLAXIUS (Males Only) (modified from Kramer 1979, figure references from that work)

1. Anal flap in lateral view with single or paired lobe-like projection or processes originating between ventral margins … 2

1.- Anal flap in lateral view either simple or process (or processes) originating directly from one or both ventral margins … 7

2. Aedeagus with three processes … 3

2.- Aedeagus with two processes … 4

3. Projection from anal flap single and almost at the apex of ventral margin (fig. 120); all aedeagal processes with slender bases (figs. 121-122) … jamaicae

3.- Projection from anal flap paired and before the middle of ventral margin (fig. 123); at least dominant aedeagal process with extremely broad base (figs. 123-124) … meadi

4. Both aedeagal processes projecting to or above the dorsal margin of aedeagus in lateral view … 5

4.- Only one or neither aedeagal process projecting to or above the dorsal margin of aedeagus in lateral view … 6

5. Apex of proximal aedeagal process much basad of the apex of the distal aedeagal process (fig. 127); ventral margin of aedeagus smooth distally (fig. 126); frons unicolorous … simplicatus

5.- Apices of aedeagal processes in alignment (figs. 129-130); ventral margin of aedeagus shagreened distally (figs. 129-130); frons laterally bordered with scarlet … caldwelli

6. Aedeagus smooth with ventral margin simple and longer aedeagal process obliquely directed ventrobasally (figs. 106, 108-109) … crudus

6.- Aedeagus at least in part shagreened with ventral margin produced to form slender, ventrally projecting process and longer aedeagal process not as above (figs. 132-133) … serratus

7. Aedeagal flagellum well developed, nearly as long as aedeagus in lateral view (figs. 136, 142); anal flap in left lateral view with part of right margin exposed distally and ventrally (figs. 138, 141) … 8

7.- Without the above combination of characters … 12

8. With no aedeagal process more than half length of flagellum; frons with narrow dark brown stripe slightly mesad of and following each lateral margin for entire length … 9

8.- With one or more aedeagal processes more than half length of flagellum; frons not marked as above … 10

9. Aedeagal flagellum with slender process near the middle on the right side (fig. 136); mesonotum predominantly yellow or yellowish-brown (Costa Rica) … phylax

9.- Aedeagal flagellum without process on right side (fig. 139); mesonotum predominantly black (Peru) … mokos

10. Aedeagus in right lateral view with ventral-most process originating from the ventral margin (fig. 142) … sillos

10.- Aedeagus in right lateral view with ventral-most process originating above ventral margin… 11

11. Aedeagus in lateral view indented on ventral margin and with a small finlike structure only on the right side (figs. 144-145) … lyssa

11.- Aedeagus in lateral view not indented on ventral margin and with a small finlike structure on both right and left sides (figs. 147-148) … brimosis

12. Forewings or face or both predominantly or entirely darkly fuscus to black … 13

12.- Forewings and face lighter in color and either unmarked or with variable markings … 15

13. Entirely dark species except for a hyaline patch on stigmal area of each forewing; two longer aedeagal processes, one directed dorsocephalad and the other caudad (figs. 150-151) … gnophos

13.- Without both of the above features … 14

14. Forewings dark except for pale costal margin, claval veins, and Cu-1; aedeagus broadly U-shaped with major process originating in basal half and directed posteriorly at or along the dorsal margin (figs. IS3-1S4) … nimbus

14.- Forewings hyaline and only darkly marked in distal third; aedeagus suboval with no processes in the basal half (figs. 156-157) … tekmar 

15. Anal flap with paired, sharply acute processes subapically on ventral margin … 16

15.- Anal flap simple, with blunt fingerlike processes, or otherwise different … 17

16. Processes on ventral margin of anal flap broadly triangular (fig. 159); aedeagus with no processes on or projecting above dorsal margin in lateral view (fig. 162); frons with transverse black markings … laevis 

16.- Processes on ventral margin of anal flap narrow and stiletto-like (fig. 163); aedeagus with three processes on or projecting above dorsal margin in lateral view (figs. 164); frons unicolorous … akko 

17. Aedeagus with well-developed left apical appendage, as wide as and concealing much of associated left process in lateral view (figs. 166, 169, 172) … 18

17.- Aedeagus without well-developed left apical appendage as above … 20

18. Left apical appendage of aedeagus with distal margins entirely serrate (fig. 166); aedeagus in right lateral view with broad, subapically rounded, apically pointed process directed cephalad (fig. 167); frons greenish with midline red or orange-red … thryligma

18.- Left apical appendage of aedeagus with distal margin only partly or not at all serrate; aedeagus in right lateral view without process; frons not colored as above … 19

19. Left apical appendage of aedeagus bluntly oval in left lateral view, with ventral margin of anal flap convex and not exposing right ventral margin (fig. 169) … gomphos

19.- Left apical appendage of aedeagus tapered and produced distally in left lateral view, with left ventral margin of anal flap concave and exposing strongly convex right ventral margin (fig. 172) … sparagma 

20. Aedeagus in lateral view subtriangular, nearly twice broader apically than basally (figs. 175-176); flagellum well developed with two processes at or near apex (fig. 177) … delta (now in Myxia; Echavarria et al. 2021)

20.- Without either of the above features … 21

21. Ventral margin of the anal flap with lobe, fin, or other type of modification before apex in left lateral view … 22

21.- Ventral margin of anal flap simple before apex in left lateral view … 25

22. Anal flap stalk-like in basal half with broad lobe just before apex in left lateral view (fig. 178); aedeagus in lateral view subquadrangular and almost uniform in width (fig. 179) … synavei

22.- Anal flap broad in basal half with ventral modification at or before middle in left lateral view (figs. 181, 184, 187); aedeagus in lateral view not of uniform width … 23

23. Aedeagus with long, broadly based, upright process near middle of left side in lateral view (fig. 181); ventral margin of aedeagus with short, caudally projecting process near apex (fig. 182) … vilbastei

23.- Without either of the above features … 24

24. Anal flap with ventral modification subquadrate in left lateral view (fig. 184); aedeagus in lateral view with two long processes originating in basal half and directed caudad (fig. 185); style with two projections on dorsal margin in distal area (fig. 184) … deleter

24.- Anal flap with ventral modification not subquadrate in left lateral view (fig. 187); aedeagus in lateral view without processes originating in the basal half (fig. 188); style without projections in the distal area (fig. 187) … spanglerorum

25. With free anteriorly projecting process on ventral margin at or near the apex of aedeagus … 26

25.- Without process as above … 28

26. Aedeagal flagellum is well-developed, broad, and half aedeagal length (figs. 191) … fennahi

26.- Aedeagal flagellum poorly developed, narrow, and less than half aedeagal length …. 27

27. Aedeagus in lateral view with large ventral keel and three long processes anteriorly directed on or near dorsal margin (figs. 193-194) … tekton

27.- Aedeagus in lateral view with only ventral process and no long processes on dorsal margin (figs. 196-197) … frontalis 

28. Aedeagus prolonged distally by acute, sharp, ventral projection (figs.199-200); posterodorsal angle of pygofer simple (fig. 199); aedeagus with one anteriorly directed process (fig. 199) … dolon

28.- Aedeagus not prolonged distally (figs. 202-203); posterodorsal angle of pygofer with short, distinct lobe (fig. 202); aedeagus with three anteriorly directed processes (figs. 202-203) … skarphion 

Online resources

3i.
EOL.
Bugguide.
Discover Life.
FLOW.
Hoppers of North Carolina.
Kunzweb Gallery.
American Insects.
BOLD.

Collecting

I’ve had the best luck sweeping grasses and sedges near streams, but mostly I get H. pictifrons (this species is recorded on willow and I have collected from willow). Haplaxius crudus is collected from palm fronds (an ongoing study uses sticky cards). Haplaxius shows up in Lindgren funnel traps (in Pennsylvania) along with other cixiids.

Molecular resources 

Genbank has data for Haplaxius crudus and Haplaxius sp. (Genbank lists Haplaxius deleter, and H. skarphion, but the generic status of these species requires confirmation); Barcode of Life has data for 7 species, recorded under Myndus (H. balli, H. fulvus, H. glyphis, H. ovatus, H. pictifrons, H. radicis, H. xyron, plus 2 Myndus, but only 1 with data) – Update (May 2017 22 Nov. 2018 28 Jan. 2020); BOLD has 5 species under Haplaxius (but only 1 with data).

The complete mitogenome is now available for Haplaxius crudus (Komondy et al. 2020)

Selected references

Attilio Bianco, P., G. Romanazzi, N. Mori, W. Myrie and A. Bertaccini. 2019. Chapter 11. Integrated Management of Phytoplasma Diseases. Pp. 237-258. In: A. Bertaccini, P. G Weintraub, G. P. Rao and N. Mori (eds). Phytoplasmas: Plant Pathogenic Bacteria – II. Springer Nature Singapore Pte Ltd., Singapore. x + 258 pp.

Bahder, B. W., E. E. Helmick, De-Fen Mou, N. A. Harrison and R. Davis. 2018. Digital PCR technology for detection of palm-infecting phytoplasmas belonging to group 16SrIV that occur in Florida. Plant Disease 102(5): 1008-1014. https://doi.org/10.1094/PDIS-06-17-0787-RE.

Bahder, B. W., E. E. Helmick and N. A. Harrison. 2017. Detecting and differentiating phytoplasmas belonging to subgroups 16SrIV-A and 16SrIV-D associated with lethal declines of palms in Florida using qPCR and high-resolution melt analysis (HRMA). Plant Disease 101(8): 1449–1454. ttps://doi.org/1410.1094/PDIS-1401-1417-0023-RE.

Bahder, B.W., E.A. Barrantes Barrantes, M.A. Zumbado Echavarria, D.F. Mou, E.E. Helmick, and C.R. Bartlett. 2020. A new species of planthopper in the genus Haplaxius (Hemiptera: Auchenorrhyncha: Fulgoroidea: Cixiidae) on palms in Costa Rica and a new country record for Haplaxius skarphion. Zootaxa 4767(4): 543-552. https://doi.org/10.11646/zootaxa.4767.4.4

Bahder, B. W., C. R. Bartlett, E. A. Barrantes Barrantes, M. A. Zumbado Echavarria, A. R. Humphries, E. E. Helmick, E. M. Goss and M. S. Ascunce. 2019. A new genus and species of cixiid planthopper (Hemiptera: Auchenorrhyncha: Fulgoroidea) from the Reserva Privada el Silencio de Los Angeles Cloud Forest in Costa Rica. Zootaxa 4701(1): 65-81. https://doi.org/10.11646/zootaxa.4701.1.5

Barrantes, E.A.B., M.A.Z. Echavarria, C.R. Bartlett, E.E. Helmick, and. B.W. Bahder. 2021. A new species of planthopper in the genus Haplaxius (Hemiptera: Auchenorrhyncha: Fulgoroidea: Cixiidae) from coconut palm (Cocos nucifera) in Costa Rica. Zootaxa 4963(3): 412-422. https://doi.org/10.11646/zootaxa.4963.3.2.

Ball, E. D. 1902. Some new North American Fulgoridae. Canadian Entomologist 34: 147-157.

Ball, E. D. 1933e. The genus Myndus Stål in North America (Homoptera: Fulgoridae). Journal of the Washington Academy of Science 23(10): 478-484.

Bartlett, C. R., L. B. O’Brien and S. W. Wilson. 2014. A review of the planthoppers (Hemiptera: Fulgoroidea) of the United States. Memoirs of the American Entomological Society 50: 1-287.

Bastos, L. F., A.V.F. dos Santos, F.V. Penner, L.M. M. Siqueira, A. G. da Silva, I. C. F. Martins, P.M.P. Lins and T.F.V. Batista. 2019. Spatial analysis and population dynamics of Haplaxius crudus (Hemiptera: Cixiidae) in coconut Amazon. Journal of Agricultural Science 11: 14; 1-12. https://doi.org/10.5539/jas.v11n14p186. (ePub Article #14, 12 pp.)

Beirne, B. P. 1950. The Canadian Cixiidae (Homoptera: Fulgoroidea). The Canadian Entomologist 82: 93-101.

Beltran-Aldana, I.J., A.E. Bustillo-Pardey, and A. Morales-Rodriguez. 2020. Mass rearing of Haplaxius crudus (Hemiptera: Cixiidae), an important insect pest in the palm oil industry of Colombia. American Journal of Entomology 4(2): 39-44. http://doi.org/10.11648/j.aje.20200402.1

Bragard, C., K. Dehnen-Schmutz, F. Di Serio, P. Gonthier, M.-A. Jacques, J.A.J. Miret, A. Fejer Justesen, C.S. Magnusson, P. Milonas, J.A. Navas-Cortes, S. Parnell, R. Potting, P.L. Reignault, H.H. Thulke, W. Van der Werf, A.V. Civera, J. Yuen, L. Zappala, E. Czwienczek, Virag Kertesz, F. Streissl, and A. MacLeod. 2020. Pest categorisation of Haplaxius crudus. EFSA Journal 18(7, article 6224): 1-31.

Caldwell, J. S. 1946. Notes on Haplaxius Fowler with descriptions of new species (Homoptera, Cixiidae). Proceedings of the Entomological Society of Washington 48: 203-206.

Caldwell, J. S. 1951. New Cixiidae from Southern North America with notes on others. (Homoptera: Fulgoroidea). The Ohio Journal of Science. Columbus 51: 34-36.

Cherry, R. H. and F. W. Howard. 1984. Sampling for adults of the planthopper Myndus crudus, a vector of lethal yellowing of palms. Tropical Pest Management 30(1): 22-25.

Dzido, J., R. S. Borges, M. Narvaez, J. Julia, C. Oropeza, and M. Dollet. 2012. Transmission of phytoplasmas associated with Texas Phoenix palm decline (16SrIV-D) to Pritchardia pacifica by the planthopper Haplaxius crudus (Cixiidae) in Yucatan, Mexico. Phytopathology 102(11): S6.2 (abstract).

Dzido, J. L., R. Sánchez, M. Dollet, J. F. Julia, M. Narvaez, S. Fabre, and C. Oropeza. 2020. Haplaxius crudus (Hemiptera: Cixiidae) transmits the lethal yellowing phytoplasmas, 16SrIV, to Pritchardia pacifica Seem. & H. Wendl (Arecaceae) in Yucatan, Mexico. Neotropical Entomology (2020): (PrePrint, 11 pp.). https://doi.org/10.1007/s13744-020-00799-2.

Echavarria, M.A.Z., E.A.B. Barrantes, C.R. Bartlett, E.E. Helmick, and B.W. Bahder. 2021. A new species of planthopper in the genus Myxia (Hemiptera: Auchenorrhyncha: Cixiidae) from the Reserva Privada el Silencio de Los Angeles Cloud Forest in Costa Rica. Zootaxa 4915(3): 351-363. https://doi.org/10.11646/zootaxa.4915.3.4.

Eden-Green, S. J. and M. Schuiling. 1976. “Root acquisition” transmission tests with Haplaxius crudus (Hom: Cixiidae) and Proarna hilaris (Hom: Cicadidae).(Abstract). Principes 20: 66.

Emeljanov, A. F. 1989. To the problem of division of the family Cixiidae (Homoptera, Cicadina). Entomologicheskoe Obozrenie 68(1): 93-106.

Farr, T. H. 1985. The earliest known date of collection of Myndus crudus (Homoptera: Cixiidae) from coconut palms in Jamaica. Florida Entomologist 68(1): 226-227.

Ferreira, M., S. H. McKamey and R. T. Martinez. 2010. New records of Haplaxius (Hemiptera: Cixiidae) in the Dominican Republic, with description of a new species. Zootaxa 2614: 65-68.

Flock, R. A. 1951. A new species of Haplaxius, with a key to California species (Homoptera: Cixiidae). Pan-Pacific Entomologist 27(4): 169-171.

Fowler, W. W. 1904. Order Rhynchota. Suborder Hemiptera-Homoptera. (Continued). Biologia Centrali-Americana; contributions to the knowledge of the fauna and flora of Mexico and Central America 1: 77-84, 85-124.

Flock, R. A. 1951. A new species of Haplaxius, with a key to California species (Homoptera: Cixiidae). Pan-Pacific Entomologist 27(4): 169-171.

Gronenborn, B., J. W. Randles, D. Knierim, Q. Barrière, H. J. Vetten, N. Warthmann, D. Cornu, T. Sileye, S. Winter and T. Timchenko. 2018. Analysis of DNAs associated with coconut foliar decay disease implicates a unique single-stranded DNA virus representing a new taxon. Scientific Reports 8(1): 5698. https://doi.org/10.1038/s41598-018-23739-y

Guerrero, M.R., A.E.B. Pardey, and A.M. Rodríguez. 2020. Efficacy of Metarhizium anisopliae to control adults of Haplaxius crudus (Van Duzee) (Hemiptera: Cixiidae), vector of lethal wilt disease of oil palm in Colombia. International Journal of Tropical Insect Science (2020): 1-7. https://doi.org/10.1007/s42690-020-00234-4.

Gurr, G. M., A. Bertaccini, D. Gopurenko, R. R. Krueger, K. A. Alhudaib, J. A. Liu, and M. J. Fletcher. 2015. Phytoplasmas and Their Insect Vectors: Implications for Date Palm. Pp. 287-314. In: W. Wakil, J. Romeno Faleiro, T.A. Miller. (eds.). Sustainable Pest Management in Date Palm: Current Status and Emerging Challenges. Springer International Publishing, Switzerland.

Halbert, S., S. W. Wilson, B. Bextrine and S. B. Youngblood. 2014. Potential planthopper vectors of palm phytoplasmas in Florida with a description of a new species of the genus Omolicna (Hemiptera: Fulgoroidea). Florida Entomologist 97(1): 90-97.

Hemmati, C., M. Nikooei, and A.M. Al-Sadi. 2020. Four decades of research on phytoplasma diseases of palms: A review. International Journal of Agriculture and Biology https://doi.org/10.17957/IJAB/15.1 (PrePRINT) [Haplaxius crudus]

Hill J. G., Seltzer J. L., Hendon A. & Bartlett C. R. 2018. First report of the American palm cixiid (Hemiptera: Cixiidae) from Mississippi, USA. Transactions of the American Entomological Society 144: 593-597.

Holzinger W. E., A. F. Emeljanov and I. Kammerlander. 2002. The family Cixiidae Spinola, 1839 (Hemiptera: Fulgoromorpha) – a Review. Denisia 4: 113-138.

Howard, F. W. 1980. Population densities of Myndus crudus Van Duzee (Homoptera: Cixiidae) in relation to coconut lethal yellowing distribution in Florida. Principes 24:174-178.

Howard, F. W. 1981. Pigment migration in the eye of Myndus crudus (Homoptera: Cixiidae) and its relation to day and night activity. Insect Science and Its Application 2(3): 129-133.

Howard, F. W. 1981. Attractiveness of date and coconut palms to Myndus crudus and other homopterans. Proceedings of the Florida State Horticultural Society 93:199-201.

Howard, F. W. 1987. Myndus crudus (Homoptera: Cixiidae), a vector of lethal yellowing of palms. Pp. 117-129.  In: M.R. Wilson and L.R. Nault,(eds.). Proceedings of the 2nd International Workshop on Leafhoppers and Planthoppers of Economic Importance : Brigham Young University, Provo, Utah, USA, 28th July-1st August 1986. CAB International Institute of Entomology.

Howard, F. W. 1989. Evaluation of six species of grasses as breeding host of Myndus crudus, a vector of lethal yellowing in palms. Proceedings of the Caribbean Food Crops Society 25: 144-146.

Howard, F. W. 1990. Evaluation of grasses for cultural control of Myndus crudus, a vector of lethal yellowing of palms. Entomologia Experimentalis et Applicata 56: 131-137.

Howard, F. W. 1990. Evaluation of Brachiaria and St. Augustune grasses as breeding hosts of Myndus crudus Van Duzee, a vector of leathal yellowing disease of Palms. Beiträge zur Tropischen Landwirtschaft und Veterinarmediziz (Leipzig) 28: 425-430.

Howard, F. W. 2006. American Palm Cixiid, Myndus crudus Van Duzee (Insecta: Hemiptera: Auchenorrhyncha: Fulgoroidea: Cixiidae). Florida Cooperative Extension Service, Institute of Food and Agricultural Sciences, University of Florida. document EENY-389 (IN704).

Howard, F. W. and C. Oropeza, 1998. Organic mulch as a factor in the nymphal habitat of Myndus crudus (Hemiptera: Auchenorrhyncha: Cixiidae). Florida Entomologist 81(1): 92-97.

Howard, F. W. and D. L. Thomas. 1980. Transmission of palm lethal decline to Veitchia merrillii by a planthopper, Myndus crudus. Journal of Economic Entomology 73: 715-717.

Howard F. W. and S. W. Wilson. 2001.  Hemiptera: Auchenorrhyncha. Pp. 128-161. In: Howard F. W., Moore D., Giblin-Davis R. M. & Abad R. G. 2001. Insects on Palms, CABI Publishing, Oxon (Great Britain).

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