Tetralicia, Harrison, 1917

Valencia, Luis & Evans, Gregory A., 2024, Review of the genus Tetralicia (Hemiptera: Aleyrodidae) and description of two new species of whiteflies found on avocado trees in Peru, Zootaxa 5403 (2), pp. 197-238 : 209-211

Key to Tetralicia species of the world (puparia)

(see annotated checklist for sources of the illustrations)

1 Body pale, transverse molting suture (tms) reaching the lateral margin; eyes absent; mesothoracic setae (Ts2) and metathoracic setae (Ts3) present; cephalic setae (Cs1) present or absent..................................................... 2

– Body black; other characters variable..................................................................... 4

2 Ts2 absent; Ts3 and Cs1 present, set on raised tubercles; body 1.6x longer than wide; tms extends between the submargin and lateral margin; Chad on Vitex ( Fig. 4E )................................................ tuberculata Bink-Moenen

– Ts2 and Ts3 present, Cs1 if present not set on raised tubercles; body 1.5–1.6x longer than wide; tms extends to the lateral margin; Mexico....................................................................................... 3

3 Cs1 absent; body elongate oval, about 1.6x longer than wide; margin deflexed about 0.3x radius (measured along the tms from its junction with the longitudinal furrow to the lateral margin); submedial region of abdomen with a row of 4 pairs of large round circular areas; Ts2 and Ts3 setae short, each about 20–21 μm; Mexico on Ipomoea ( Fig. 6H )........................................................................................ pallida (Carapia-Ruiz & Sánchez-Flores)

– Cs1 setae present; body elliptical, about 1.5x longer than wide with lateral margin deflexed about 0.4x radius submedial region of abdomen without large round circular areas; Ts2 and Ts3 longer, each about 55 μm; Mexico on Guazuma ( Fig. 6I )..................................................................... palidonigrans (Carapia-Ruiz & Sánchez-Flores)

4 Ts2 absent; Ts3 and Cs1 present or absent.................................................................. 4

– Ts2 and Ts3 present; Cs1 present or absent................................................................ 14

5 Ts3 present; eyes absent................................................................................ 6

– Ts3 absent; eyes present or absent........................................................................ 7

6 Tms extends to the submargin; body 1.8x as long as wide; caudal protuberance not well developed; Chad, on Hyparrhena grass ( Poaceae ) ( Fig. 4C )................................................................ graminicola Bink-Moenen

– Tms extends to lateral margin; body 1.9x as long as wide; vasiform orifice (vo) 1.3x as long as wide, caudal protuberance well developed; USA (CA) on Rhamnus ( Fig. 6F )..................................... oblanceolata (Drews & Sampson)

7 Eighth abdominal segment setae (As8) absent; dorsum with 4 longitudinal rows of circular areas, body elongate slender, 1.8x as long as wide; posterior margin not protruding; Australia on Grevillea ( Proteaceae ) ( Fig. 4A .... debarroi Martin & Carver

– As8 present; dorsum without longitudinal rows of circular areas; body 1.1–2.0x as long as wide; not from Australia nor on Proteaceae ........................................................................................... 8

8 Cephalus with 3 pairs of short setae; eyes absent; body very slender, 2.0x as long as wide; Turkmenistan on Erianthus ( Poaceae ) and Rununculus (Rununculaceae) ( Fig. 4B )...................................................... erianthi Danzig

– Cephalus without setae (except with 1 pair in nevadensis ); eyes present or absent.................................. 9

9 Eyes present; body broadly oval, 1.3x as long as wide; known from Mexico or the USA on Quercus ................... 10

– Eyes absent; body shape variable; known from Europe or the USA, not on Quercus ................................ 12

10 Cephalus with 1 pair of setae; eyes in the shape of a transverse slit; body broadly oval, 1.3x as long as wide, with deflexed margin 0.2x radius; central area of the cephalothorax and abdomen covered with tubercles USA (NV), Mexico, on Quercus ( Fig. 6D )............................................................................. nevadensis (Dooley)

– Cephalus without setae; eyes oval shaped; body broadly oval, 1.3x as long as wide, with deflexed margin 0.3–0.4x radius. 11

11 Lateral margin deflexed 0.4x radius; caudal setae (Cas) extremely long (187), slightly more than 2x As8 (87); medial region of body rugose, submargin pebbly; operculum (op) elongate,1.3x as long as wide ( Fig. 5C )................................................................................................ anahuac (Carapia-Ruiz & Sánchez-Flores)

– Lateral margin deflexed 0.3x radius; Cas not as long (113–137), slightly less than 2x As8 (65–87) (central region of body fossulate, submargin tiled; cephalothorax and sides of abdomen covered with pebbly sculpturing; abdominal sutures with rows of transverse elliptical markings across each segment; vo elongate, 1.3x as long as wide with op occupying about half the orifice; ( Fig. 6P )................................................... xalapensis (Sánchez-Flores & Carapia-Ruiz)

12 Body round to broadly oval, 1.1–1.3x as wide as long; dorsal surface with rough mosaic sculpturing; deflexed margin 0.2x radius, apparently smooth; Portugal, Spain on Erica ( Fig. 4D )................................. iberiaca Bink-Moenen

– Body elongate slender, 1.8x as wide as long; dorsal surface, more or less smooth, without rough mosaic sculpturing; deflexed margin 0.4x radius with striate sculpturing................................................................ 13

13 Lateral margin crenulate shallowly; vo and operculum (op) longer than wide; As8 and Cas setae short, subequal; tms extending to lateral margin or close to it; widespread in Europe on Erica ( Fig. 3A–E )............................. ericae Harrison

– Lateral margin with urn-shaped, bilobate teeth; vo and op nearly quadrate; As8 short; Cas long; tms extending to submargin USA (CA) on Pluchea ( Fig. 5I )............................................... coachellensis (Drews & Sampson)

14 Vo raised high on a volcano-like projection, posterior margin of seventh abdominal segment with a dark, sclerotized arch; Brazil, on Cecropia ( Figs. 4G–I )........................................................... cecropiae (Bondar)

– Vo not raised on a volcano-like projection, posterior margin of seventh abdominal segment without a dark, sclerotized arch; species not known from Brazil.......................................................................... 15

15 Lateral margin with 2 rows of teeth; body elongate slender, 1.9x as long as wide; eyes absent; op transverse 0.7x as long as wide, filling about 0.7x the vo, the latter 1.2x as long as wide; Mexico on Diospyros ( Fig. 5E )............................................................................................... bidentata (Sánchez-Flores & Carapia-Ruiz)

– Lateral margin with 1 row of teeth; body variable but less than 1.9x as long as wide in most species; eyes present or absent 16

16 Tms extending to lateral margin; eyes absent.............................................................. 17

– Tms not extending to lateral margin; eyes present or absent................................................... 25

17 Cs1present; Ts2 and Ts3 very elongate (except in A. sierrae ); deflexed margin 0.3–0.4x radius....................... 18

– Cs1 absent; Ts2 and Ts3 very short (except in A. popocatepetl ); deflexed margin 0.3x radius........................ 20

18 Vo and op transverse, each 0.7x as long as wide; Ts3 and As8 setae short, body 1.4x as long as wide; margin deflexed 0.4x radius; USA (CA) on a shrub ( Fig. 6O )........................................................ sierrae Sampson

– Vo and op subquadrate (except transverse in A. rotunda ), Ts2, Ts3 and As8 setae long; body 1.3 or 1.6x as long as wide, with margin deflexed 0.3x radius............................................................................ 19

19 Body 1.3x as long as wide; vo and op transverse, 0.7–0.8x as long as wide; submedial region of abdomen speckled with black dots; Cs1, Ts2, Ts3 and Cas setae long and stout; caudal margin rounded; posterior margin of vo floor without a row of cell-like structures; Mexico on unknown host ( Fig. 6M )................................................... rotunda Baker

– Body 1.6x as long as wide; vo 1.1x longer than wide; submedial region of abdomen without black dots; Cs1, Ts2, Ts3 and Cas setae very long, subequal, each about 3x as long as the vo, As8 long, about 2.2x the vo; caudal protuberance not developed; posterior margin of vo floor with a row of 4 roundish,cell-like structures; Peru on Persea ( Figs. 2 , 7 , 8 )........................................................................................... sawyeri Valencia & Evans, n. sp.

20 Body slender elliptical, 1.7x as long as wide............................................................... 21

– Body oval to broadly oval, 1.4–1.6x as long as wide…………................................................. 23

21 Mesothorax and metathorax with 3 and 2 pairs of setae, respectively; central region of body with ornate sculpturing; body slender elliptical, vo 1.4x as long as wide, USA (CA) on Eriodictyon ( Fig. 6G )................ ornata (Drews & Sampson)

– Mesothorax and metathorax each with 1 pair of setae; central region of body without ornate sculpturing; vo 1.0–1.1x as long as wide.............................................................................................. 22

22 Ts2 (50) longer than Ts3 (37); Cas 2.5–3.5x as long as As8; Mexico, on a grass, Dichanthelium ( Poaceae ) ( Fig. 6K )....................................................................... popocaepetl (Carapia-Ruiz & Sánchez-Flores)

– Ts2 shorter (37) or subequal to Ts3; Cas more than 4x as long as As8; body 1.7–1.9x as long as wide; lateral margin paler than central region, vo 1.1x as long as wide; USA (CA), Mexico on Arctostaphylos and Salvia ( Fig. 6A ).. laingi (Drews & Sampson)

23 Body 1.4x as long as wide; central region of body smooth; vo 1.1x as long as wide, operculum with many short, spinelike setae along its posterior margin; USA (CA, NV), Mexico on Ceanothus ( Fig. 5G )......................... ceanothi Sampson

– Body 1.5–1.6x as long as wide; central region of body smooth or granulate; vo 1.0–1.1x as long as wide, operculum without many short, spinelike setae along its posterior margin........................................................ 24

24 Body 1.5x as long as wide; dorsal surface granulated; As8 and Cas setae very long; dorsum granulated deflexed submargin with rows of dark spots; Mexico, USA (TX) on Lantana ( Fig. 5L )........................... granulata Sampson and Drews

– Body 1.6x as long as wide, dorsal surface not granulated; As8 setae short; Mexico on Karwinskia . ( Fig. 6N )............................................................................... sampsoni (Sanchez-Flores & Carapia-Ruiz)

25 Body elongate slender, 1.7–1.9x as wide as long; eyes absent; dorsal setae variable................................ 26

– Body oval, 1.3–1.5x as wide as long; eyes present; all dorsal setae short......................................... 27

26 Body 1.7x as long as wide; all dorsal setae including Cas setae short; lateral margin with unilobate teeth; Mexico on Coccoloba and Guazuma ( Fig. 5F )................................................ caudata (Sánchez-Flores & Carapia-Ruiz)

– Body 1.9x as long as wide; dorsal setae longer, Cas setae very long (102); lateral margin with bifurcate teeth; Mexico on Cecropia ( Fig. 6L )............................................ pseudogranulata (Carapia-Ruiz & Sánchez-Flores)

27 Submargin with a row of about 6 conical projections on each side, separated from each by more than the diameter of one projection; Ts2 and Ts3 median length, rod-like with clubbed apices; vo and op about as long as wide; body 1.4x as long as wide, deflexed margin about 0.4x radius; USA (CA), Mexico on Hyptis ( Fig. 6O )................... hyptisemoryi (Gill)

– Submargin without conical projections; Ts2 and Ts3 filiform, not rod-like with clubbed apices; other characters variable.. 28

28 Lateral margin with a round protuberance at the level of the transverse molting suture; Cas and other dorsal setae short, body 1.4x as long as wide; margin deflexed 0.3x radius; USA (CA) on Fouquieria ( Fig. 5K )........................................................................................ fouquieriasplendens (Carapia-Ruiz & Sánchez-Flores)

– Lateral margin without a protuberance at the level of the transverse molting suture; Cas variable..................... 29

29 Cs1 and eyes present................................................................................. 30

– Cs1 absent; eyes present or absent...................................................................... 32

30 Cs1 long (50–85); Cas and As8 setae similar in length; deflexed margin 0.1x body radius with striate sculpturing; abdomen with transverse chain-like sculpture across the sutures; marginal teeth large, with a structure starting anterior to true margin towards the head area divided into inverted “Y” shape; dorsum with rows of elongate tile-like sculpturing; Mexico on Chamaedorea ( Fig. 5H )............................................... chamaedoreaelegans (Sánchez-Flores and Carapia-Ruiz)

– Cs1 short; Cas at least 2x as long as As8 setae; deflexed margin 0.3–0.5x body radius; abdomen smooth; marginal teeth not large, without a structure starting anterior to true margin toward head area divided into inverted “Y” shape............. 31

31 With mid-abdominal depressions and tubercles, area near submarginal fold of both subdorsum and submargin with fine transverse lines; vo and op as long as wide, op filling 0.9x the vo; a pair of dark, sclerotized sickle-shaped markings present along the lateral margin of the vo; deflexed margin smooth, 0.2x body radius; eyes prominent; cephalothorax entirely dark, abdomen primarily pale with abdominal depressions and tubercles medially, submargin granulate; venter with small, round, bead-like sculpturing just inside the legs; USA, Mexico (intercepted at U.S. ports of entry from Colombia, Dominican Republic, El Salvador and Peru) on Quercus , Persea , Magnolia ( Fig. 5A )................................ abnormis (Quaintance)

– Without mid-abdominal depressions and with tubercles forming an arch, area close to the submarginal fold of both the subdorsum and the submargin with fine transverse lines; vo 1.1x as long as wide; op 0.8x as long as wide, filling 0.6x the vo; deflexed margin striate, 0.1x body radius; Mexico on Annona ( Fig. 5D )......... annonae (Carapia-Ruiz & Sánchez-Flores)

32 Eyes present; deflexed margin 0.1–0.5x radius............................................................. 33

– Eyes absent, deflexed margin 0.1–0.3x body radius......................................................... 34

33 Deflexed margin 0.5x body radius with pebbly sculpturing; As8 setae set on tubercles; body 1.4x as long as wide; vo and op quadrate; dorsocentral region of abdomen with transverse chain sculpturing along the sutures; Mexico on Quercus ( Fig. 6B ).................................................................. mexicana (Carapia-Ruiz & Sánchez-Flores)

– Deflexed margin 0.1–0.2x body radius with smooth or striate sculpturing; As8 setae not set on tubercles; body 1.3–1.4x as long as wide; Mexico on Ficus ( Fig. 5N )................................... guerrerensis (Carapia-Ruíz & Sánchez-Flores)

34 Posterior margin of body truncated; lateral margin with a narrow translucent band behind the marginal teeth; deflexed margin about 0.3x radius; vo triangular shaped; op heart shaped; Cas very short; USA (CA), Mexico, on Arctostaphylos ( Fig. 5B )............................................................................... acaudata (Drews & Sampson)

– Posterior margin of body pointed, not truncated; lateral margin without a narrow translucent band behind the marginal teeth; Cas medium to long; other characters variable............................................................. 35

35 Vo transverse, 0.8x as long as wide; Cas very long, dorsum smooth with imbricate band on the submargin; margin teeth forked; USA, Mexico, El Salvador, Haiti on various hosts ( Fig. 6E )........................................ nigrans (Bemis)

– Vo quadrate to elongate, 1.1–1.2x as long as wide; Cas medium to very long; other characters variable................. 36

36 Cas and As8 short (8–12 μm) and subequal in length; dorsocentral region darker, cone shaped with yellowish lateral margins; op filling almost filling the vo; body 1.5x as long as wide; dorsum of abdominal segments smooth, without depressions; Mexico on Eriogonum ( Fig. 5J )............................................................. eriogonum (Carapia-Ruiz)

– Cas about 2x as long as As8 setae; dorsocentral region not darker and cone shaped................................ 37

37 Cas short (45–50); Ts2 and Ts3 short (12–14); op filling about 0.7x the vo; body 1.4x as long as wide; marginal teeth truncate; at least lateral margins of body sculptured; Mexico on Guazuma ( Fig. 5M )...... guazumae (Carapia-Ruiz & Sánchez-Flores)

– Cas very long (100–102); Ts2 and Ts3 medium length (18–50); body 1.5–1.6x as long as wide; op filling about 0.9x the vo; marginal teeth variable, central region of the body smooth similar in color to the submarginal and marginal region....... 38

38 Ts2 and Ts3 medium in length (48–50); body 1.5x as long as wide; marginal teeth 2–3 tined; op without 3 small tubercles in a row; Mexico on Hyptis ( Fig. 6C )....................................................... mixteca (Carapia-Ruiz)

– Ts2 and Ts3 shorter (18–20); body 1.6x as long as wide; marginal teeth with truncate apices; op with 3 small tubercles in a row; Guatemala on Acalypha sp ( Fig. 6J )...................................... petenensis Carapia-Ruiz & García-Ochaeta

Figures

FIGURE 2. Tetralicia sawyeri n. sp. puparium. A) drawing with detail of vasiform orifice. Setae: Cs1 (cephalic setae), Ts2 (mesothoracic setae), Ts3 (metathoracic setae), As8 (8th abdominal segment setae), Cas (caudal setae); Structure: lms (longitudinal molting suture), cl (cluster of pores), def (deflexed margin), radius, tms (transverse molting suture), tm (true margin), vo (vasiform orifice), op (operculum), cp (caudal protuberance); B) puparium treated only with hydrogen peroxide; C) puparium treated with potassium hydroxide (10%) and hydrogen peroxide.

FIGURE 3. Tetralicia ericae Harrison, 1917. (A–E) puparia: A) habitus drawing, B) habitus slide specimen, C) abdomen, D) vasiform orifice slide specimen, E) vasiform orifice drawing; (F–K) adult male: F) habitus, G–H) genitalia, I–K) antennae; (L–P) adult female: L) habitus, M–N) ovipositor; O–P) antennae. Sources: A, E, H, I, J, N (Bink-Moenen, 1989); B, C, D, F, G, K, L, M, P (USNM specimen).

FIGURE 4. Puparia of Tetralicia species. A) debarroi Martin & Carver, B) erianthi Danzig, C) graminicola Bink-Moenen, D) iberiaca Bink-Moenen, E) tuberculata Bink-Moenen, F) cecropiae (Carapia-Ruiz, 2020b) G) cecropiae Bondar (type, MZU), H) cecropiae (type, USNM), I) cecropiae (type, Bondar 1923).

FIGURE 5. Puparia of Tetralicia species. A) abnormis (Quaintance), B) acaudata (Drews & Sampson), C) anahuac (Carapia- Ruiz & Sánchez-Flores), D) annonae (Carapia-Ruiz & Sánchez-Flores), E) bidentata (Sánchez-Flores & Carapia-Ruiz), F) caudata (Sánchez-Flores & Carapia-Ruiz), G. ceanothi Sampson, H) chamaedoreaelegans (Sánchez-Flores & Carapia-Ruiz), I) coachellensis (Drews & Sampson), J) eriogonum (Carapia-Ruiz), K) fourqueriasplendens (Carapia-Ruiz & Sánchez-Flores), L) granulata Sampson & Drews, M) guazumae (Carapia-Ruiz & Sánchez-Flores), N) guerrerensis (Carapia-Ruíz & Sánchez- Flores), O) hyptisemoryi (Gill), P) kobei (Sánchez-Flores & García-Ochaeta).

FIGURE 6. Puparia of Tetralicia species. A) laingi (Drews & Sampson), B) mexicana (Carapia-Ruiz & Sánchez-Flores), C) mixteca (Carapia-Ruiz), D) nevadensis (Dooley), E) nigrans (Bemis), F) oblanceolata (Drews & Sampson), G) ornata (Drews & Sampson), H) pallida, I) palidonigrans (Carapia-Ruiz & Sánchez-Flores), J) petensis (Carapia-Ruiz & García-Ochaeta), K) popocatepetl (Carapia-Ruiz & Sánchez-Flores), L) pseudogranulata (Carapia-Ruiz & Sánchez-Flores), M) rotunda Baker, N) sampsoni (Sanchez-Flores & Carapia-Ruiz), O) sierrae Sampson), P) xalapensis (Sánchez-Flores & Carapia-Ruiz).

FIGURE 7. Tetralicia sawyeri n. sp. puparium. A) habitus drawing, B) habitus slide mounted specimen, C) lateral margin (tm=true margin), D) vasiform orifice, E) cephalothorax (sms=submarginal setae; gp=geminate pores, cl=cluster of bright pores, F) abdomen (ad=abdominal depression, gp=geminate pores, bp=bright pores, G–K) vasiform orifice (vo) showing lingula with strongly chitinized, four finger-like projections at its apex and views at different depths of field, showing the muscles responsible for the movement of the operculum (op).

FIGURE 8. Tetralicia sawyeri n. sp. A) adults on avocado leaf; B–H) adult female: B) antenna, C) antenna, F3–F7 =flagellar segments 3–7, rs=round sensoriae, sc=sensorial cone, D) tibia I, E) tibia II, tb=tibial brush, F) tibia III, tc=tibial comb, G) ovipositor, ov=ovipositor, H) cg=cement gland.