Ferrisia dasylirii (Cockerell), 2012

Kaydan, M. B. & Gullan, P. J., 2012, 3543, Zootaxa 3543, pp. 1-65 : 19-23

Ferrisia dasylirii (Cockerell) stat. rev.

( Figs 2A, 8)

Dactylopius dasylirii Cockerell, 1896: 202 .

Pseudococcus dasylirii ; Fernald, 1903: 101. Change of combination.

Ferrisia virgata (Cockerell) ; incorrect synonymy by Ferris, 1953 a: 362.

Type material (examined). Lectotype: Dactylopius dasylirii Cockerell [designated by Williams (1996)]: adult ♀: on slide with 5 other adult ♀, labels: “ Dactylopius / dasylirii / Dasylirion / wheeleri / Organ, N.M. / Sept. 27 ’96 / (Cockerell) / TYPE” “7279” and “147/11” ( USNM). Paralectotypes: 5 adult ♀, on same slide as lectotype, and second slide with 25 first-instar nymphs, with same labels as lectotype slide ( USNM); 6 adult ♀ (6 slides), mounted from the type material and with different label: “ Pseudococcus / dasylirii (Ckll.) / on Dasylirion wheeleri / Organ, N.M. alt. 5100 ft / Sept. 27,189 6 / Tnsly colr. (7279)” ( USNM) [the label on the latter 6 slides is erroneous in that the collection was not made by Tinsley].

Cockerell (1896) described Dactylopius dasylirii from sotol, Dasylirion wheeleri ( Ruscaceae , or sometimes placed in Nolinaceae ), from a collection that he made in New Mexico near the community of Organ [at 1555 m; 32°26’N 106°36'W]. We have examined a number of collections taken from this host plant in New Mexico and Arizona. The adult females often have rather thick setae associated with the sclerotised area surrounding the dorsal enlarged ducts. Otherwise they resemble adult females from collections on a range of host plants in California that we are treating as the same species and, which like the samples from Dasylirion in Arizona, usually have very few small ventral oral-collar tubular ducts on the margins of the posterior abdominal segments. We have restricted our redescription of F. dasylirii to specimens collected in the southwest USA, but assign many morphologically similar specimens to this species pending further study, especially using freshly collected material and more variable genetic markers. Gullan et al. (2010) sequenced specimens from Aruba, Belize, Jamaica and Florida ( USA) that they considered to be conspecific with populations from Arizona and California ( Fig. 3), although they show some morphological (see below under Variation) and genetic differences from the latter populations. Nur (in lit to J.W. Beardsley) referred to a Californian population on oleander ( Nerium sp. ) from Calexico in Imperial County as species FO and considered it distinct from specimens from Florida ex Coccoloba uvifera that he called species FF. Our specimens from Florida, Aruba, Belize and Jamaica resemble Nur’s voucher specimens of FF in having few or no translucent pores on the hind coxa (many specimens of F. dasylirii from Arizona and California have at least some translucent pores on the hind coxa) and our specimens ex C. uvifera from Aruba ( Fig. 2A) further resemble Nur’s FF in having fewer than 10 (usually ≤7) multilocular disc-pores on abdominal segment VI and few small oral-collar tubular ducts in clusters on the margins of the posterior abdominal segments.

Material used for description (in addition to type material): USA: 2 adult ♀ (2 slides, including FBK018), ex Dasylirion wheeleri , Arizona, Santa Catalina Mountains, Mt Lemmon, near Molino Basin , 3.ix.2002, S. Prchal ( BME) ; 8 adult ♀ (8 slides), ex D. wheeleri , Arizona, Santa Catalina Mountains, Mt Lemmon , near “Prison Camp”, 3.ix.2002, S. Prchal ( BME) ; 16 adult ♀ (16 slides, including FBK019), ex Dasylirion sp. , Arizona, Pima Co., Oro Valley, Orange Grove Road , 24.viii.2002, J.F & D.R. Miller ( BME) ; 7 adult ♀ (7 slides, including FBK012), ex D. wheeleri , Arizona, Bisbee, 1429 Franklin St , 31 o 24’34”N, 109 o 55’58”W., 5200 ft [1585 m], 20 viii.2005, A.S. & N. Menke, ( BME) ; 103 adult ♀ (12 slides), ex D. wheeleri , Arizona, Tucson , 10.ix.1943, L.P. Wehrle, plus 5 adult ♀ (5 slides) from same collection ( Wehrle Coll. # 1448) slide-mounted by P.J. Gullan from dry collection in 2002 ( BME) ; 10 adult ♀ (5 slides), ex Dasylirion sp. , Arizona, Pima Co., 1 mile [1.6 km] S of Gardner Canyon Rd & Highway 83, 21.vii.1974, T. Halstead ( CSCA & USNM) ; 4 adult ♀ (4 slides, including FBK026), ex mulberry ( Morus sp. ), California, El Centro 2270, Sandalwood Drive , 17.x.2001, B. Roltsch ( BME) ; 2 adult ♀ (2 slides, including FBK025), ex carob ( Ceratonia siliqua ), California, Calexico, 1018 Sixth St. , 29.x.2001, B. Roltsch ( BME) ; 6 adult (3 slides), ex sotol [ D. wheeleri ], New Mexico, Doña Ana Co., Las Cruces , 13.xi.1980, G.L. Nielsen ( CSCA) .

Material sequenced and/or measured but not used for description: 6 adult ♀ (6 slides, including FBK013), ex Coccoloba uvifera (Polygonaceae) , ARUBA, Baby Beach , 12 o 24’N, 69 o 53’W, 19.vii.2005, P.J. Gullan ( BME) ; 3 adult ♀ (3 slides, including FBK009), ex ornamental shrub, BELIZE, Belize District, Cay Caulker , 4.vi.2000, D.R. Miller ( BME) ; 26 adult ♀ (26 slides, including FBK001), ex Annona squamosa (Annonaceae) , JAMAICA, Kingston, East Street , 7.ix.2003, T. Kondo (3 slides ANIC, 19 BME, 2 USNM, 2 UNCB) ; 1 adult ♀ (1 slide, FBK002), ex Anthurium sp. (Araliaceae) , JAMAICA, Ocho Rios, in garden of Crane Ridge Hotel , 4.ix.2003, T. Kondo & P.S. Cranston ( BME) ; 7 adult ♀ (7 slides), ex Acacia sphaerocephala (Fabaceae) , MEXICO, Oaxaca, Temascal , 1.v.1964, D.H. Janzen ( BME) ; 6 adult ♀ (6 slides, including FBK006), ex Codiaeum variegatum var. pictum (Euphorbiaceae) , USA, Florida, Orange Co., Apopka , 3239 Kelly Park Rd , 8.x.2001, K. Gonzalez ( BME) .

In addition to the above lists, we identified specimens of F. dasylirii from the following countries: Bahamas, Brazil, Chile, Colombia, Costa Rica, Cuba, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama, USA [Arizona ( AZ) counties: Maricopa and Pima; California ( CA) counties: Alameda, Fresno, Imperial, San Bernardino, San Diego, San Joaquin, San Francisco (under glass) and San Mateo; Florida intercepted in quarantine in CA; Hawai’i: Honolulu; New Mexico: Doña Ana Co.; Texas: Cameron Co.] and Venezuela, on the following plant species: Acacia greggii (Fabaceae) , Acalypha sp. (Euphorbiaceae) , Albizia sp. , A. julibrissin (Fabaceae) , Alpinia purpurata (Zingiberaceae) , Alternanthera sp. (Amaranthaceae) , Ananas comosus (Bromeliaceae) , Andira inermis (Fabaceae) , Annona sp. , A. squamosa (Annonaceae) , Brassica oleracea (Brassicaceae) , Cattleya sp. (Orchidaceae) , Codiaeum sp. (Euphorbiaceae) , Cocos nucifera (Arecaceae) , Cordyline fruticosa (Liliaceae) , Couroupita guianensis (Lecythidaceae) , Croton sp. (Euphorbiaceae) , Cucumis melo (Cucurbitaceae) , Dasylirion sp. (Ruscaceae) , Dieffenbachia sp. (Araceae) , Dracaena marginata , D. massangeana (Ruscaceae) , Elaeis guineensis (Arecaceae) , Eryngium foetidum (Apiaceae) , Fernaldia sp. (Apocynaceae) , Gossypium sp. (Malvaceae) , Hoya sp. (Apocynaceae) , Gossypium sp. (Malvaceae) , Hibiscus sp. (Malvaceae) , Heliconia sp. (Heliconiaceae) , Jatropha berlandii (Euphorbiaceae) , Lantana camara (Verbenaceae) , Mangifera indica (Anacardiaceae) , Morus sp. (Moraceae) , Musa sp. (Musaceae) , Nephelium lappaceum (Sapindaceae) , Nephthytis sp. (Araceae) , orchid sp. ( Orchidaceae ), Philodendron sp. (Araceae) , Pithecellobium dulce (Fabaceae) , Poinsettia pulcherrima (Euphorbiaceae) , Polyscias fruticosa (Araliaceae) , Pothos sp. (Araceae) , Psidium guajava and Psidium sp. (Myrtaceae) , Saintpaulia sp. (Gesneriaceae) , Syngonium sp. (Arecaceae) , Terminalia catappa (Combretaceae) , Theobroma cacao (Malvaceae) , Violeta ( Violaceae ), Yucca sp.(Agavaceae) , Zingiber sp. and Z. officinale (Zingiberaceae) .

ADULT FEMALE. Diagnosis. Ferrisia dasylirii can be diagnosed by the following combination of features: presence of clusters of small oral-collar tubular ducts on ventral margins of last 2 or 3 abdominal segments; ventral oral-collar tubular ducts frequently associated with a minute discoidal pore which almost never touches rim of duct, usually distant from rim by at least half length of duct; dorsal enlarged tubular ducts totalling 54–114 throughout dorsum, rim of each duct often with 1 or 2 oval discoidal pores usually touching outer margin of sclerotised area and sometimes projecting out from margin (on some specimens these discoidal pores are rare); multilocular discpores on venter of abdominal segments VI (1–19) (usually fewer than 15 in one irregular row), VII (13–36), and VIII + IX (13–22); anal lobe cerarii each with 2 conical setae (except some specimens from Dasylirion ( Fig 8b); both pairs of ostioles present; antennae usually ≥600 µm long (apical antennal segment ≥120 µm long); most specimens of F. dasylirii lack translucent pores on the hind coxa, and all examined specimens from the type locality lack these pores. However on adult females from two of the sequenced collections from California (CA 2 and CA 3) and one of the collections from Arizona (AZ 2) translucent pores are present and fairly numerous, and in a second sequenced Arizona collection (AZ 3) 12 of 16 females lack the pores, three females have a few pores on just one coxa and a fourth female has pores on both coxae .

Ferrisia dasylirii is most similar to F. virgata as both species have ventral oral-collar tubular ducts of more than one size, clusters of small oral-collar tubular ducts on the ventral margins of the last 2–3 abdominal segments, and a minute discoidal pore usually near each enlarged dorsal tubular duct and most ventral oral-collar tubular ducts but never touching the rim of the duct opening. F. dasylirii can be distinguished readily from F. virgata by the following features: (i) discoidal pores are on the outer margin of the sclerotised area around the rim of the dorsal enlarged tubular ducts on the abdomen ( Fig. 4 Ab) and often each pore and its surrounding sclerotisation projects out from the margin (in F. virgata discoidal pores associated with the sclerotised area around the rim of the dorsal enlarged tubular ducts on the abdomen usually not touching the outer margin of the sclerotised area and almost never projecting out from margin; Fig. 4 Aa); (ii) small oral-collar tubular ducts in clusters on the posterior abdomen with distal end of each duct slightly tapered towards the attachment of inner ductile (in F. virgata the distal end of each duct is rounded); (iii) antennae usually ≥600 µm long with apical segment 120–150 µm long (usually ≤600 µm long with apical segment 105–125 µm long in F. virgata ); (iv) venter of abdominal segment VI usually with ≤15 multilocular disc pores typically forming a single, sometimes irregular, row (in F. virgata ≥15 pores, usually forming at least a partial double row medially); (v) translucent pores usually absent on hind coxa (in F. virgata present on dorsal surface of hind coxa, especially posterolaterally, although often few in number); (vi) each anal lobe with only 2 cerarian setae except some specimens from Dasylirion (in F. virgata sometimes with an extra 1–2 conical cerarian seta(e) that is/are more slender than the other 2 setae). F. dasylirii is also very similar to F. cristinae , F. kondoi and F. williamsi (which have ventral oral-collar tubular ducts of more than one size, and clusters of small oral-collar tubular ducts on the ventral margins of the last 2 or 3 abdominal segments), however, F. dasylirii can be readily distinguished from the other three species by the position of the minute discoidal pores associated with ducts, which in F. dasylirii are always near the enlarged tubular ducts and ventral oral-collar tubular ducts but never touch the rim of the duct opening (discoidal pores always adjacent to duct openings in F. cristinae , F. kondoi and F. williamsi ). F. dasylirii is also close to F. milleri and F. ecuadorensis but can be separated from these two species by the absence of clusters of small oral-collar tubular ducts on the head, thorax and abdominal segments.

Description of slide-mounted specimens (based on measured specimens listed above and excluding those from Aruba, Belize, Jamaica, Mexico and Florida ( USA); Fig. 8). Body elongate oval, 3.14–5.30 mm long, 1.36–2.86 mm wide. Eye marginal, 63–85 µm wide. Antenna 8 segmented, 580–780 µm (generally ≥600 µm) long; apical segment 120–150 µm long, 30–40 µm wide. Clypeolabral shield 190–218 µm long, 170–200 µm wide. Labium 175–215 µm long, 120–188 µm wide. Anterior spiracles 60–90 µm long, 43–50 µm wide across atrium; posterior spiracles 87–110 µm long, 50–75 µm wide across atrium. Circulus quadrate, 150–220 µm wide, divided by an intersegmental line. Legs well developed; hind trochanter + femur 450–600 µm long, hind tibia + tarsus 480–650 µm long, hind claw 35–45 µm long. Ratio of lengths of hind tibia + tarsus to hind trochanter + femur 0.96–1.08, ratio of lengths of hind tibia to tarsus 2.40–3.03, ratio of length of hind trochanter + femur to greatest width of femur 3.64–4.36. Tarsal digitules subequal, each 55–60 µm long. Claw digitules subequal, each 35–45 µm long. Translucent pores present on hind legs, totalling 49–70 on femur and tibia combined, but often absent on coxae (114–150 on coxa, femur and tibia combined with 32–49 on dorsal surface of each coxa on Californian specimens). Ostioles: both pairs present; each anterior ostiole poorly developed, with 22–32 trilocular pores and 4–7 setae; each posterior ostiole with 49–67 trilocular pores and 8–12 setae. Anal ring 120–198 µm wide, with 6 anal ring setae, each seta 170–262 µm long.

Dorsum. Anal lobe cerarii each with 2 conical setae, 35–43 µm long, except some specimens on Dasylirion with an additional 2 or 3 robust setae, 20–36 µm long, slightly thinner than typical conical cerarian setae ( Fig. 8b), with 45–67 trilocular pores and 3–8 auxiliary setae. Dorsal body setae slender, each 15–60 µm long. Trilocular pores each 4–5 µm in diameter. Enlarged tubular ducts totalling 55–114 on dorsum, each duct 30–40 µm long, 5.0–7.5 µm (generally 7.0–7.5 µm) wide at mid-length, rim of duct opening sclerotised, 10–13 µm wide, surrounded by a sclerotised circular area 15–35 µm wide, often enclosing 1 or 2 oval discoidal pores (touching outer margin of sclerotised area and sometimes projecting out from margin) and with 2–7 (generally 2 or 3) setae, each 22–48 µm long, usually either within rim (especially on abdomen) or on edge of rim (especially on head); ducts distributed marginally in clusters of 2–6 on head and thorax, on margins of all abdominal segments in groups of 2–4, but with 5–12 ducts on each side of abdominal segment VII, and also 2–7 medially to submarginally on head and thorax, 5–7 medially to submedially on abdominal segments.

Venter. Body setae slender, each 20–238 µm long, longest setae medially on head; apical setae of anal lobe 260–330 µm long. Multilocular disc pores present on posterior abdominal segments only: 1–19 pores on segment VI, 13–36 on segment VII, 13–22 on segments VIII + IX; each pore 7–10 µm in diameter. Trilocular pores each 3–4 µm in diameter. Minute discoidal pores each 2.5–3.5 µm in diameter, almost always associated with oral-collar tubular ducts, but never touching rim of ducts. Oral-collar tubular ducts on most of venter (excluding margins of posterior abdomen) each 12–15 µm long, 3–4 µm wide, totalling 23–81 (totalling 23– 74 specimens on Dasylirion ), distributed as follows: 12–32 on head and thorax, and on abdominal segments: 6–21 total on segments I–III; 2–7 on IV; 1–8 on V; 2–16 on VI; 5–15 on VII; none on VIII. Small oral-collar tubular ducts each 6.5–13.0 µm long, 2.5–4.0 µm wide, distributed as follows on margin of abdominal segments: 0–1 on each side of VI; 0–9 on each side of VII; 0–8 on each side of VIII ( Fig. 8d).

Variation. Adult females of F. dasylirii display much variation in the presence or absence and abundance of translucent pores on the hind coxae. The type specimens from New Mexico lack translucent pores on the hind coxa, as do adult females from Aruba (ARU), Belize (BEL), Florida (FL) and Jamaica (JAM1 and JAM2). However females from two of the sequenced collections from Imperial County, California (CA2 and CA3), and one of the collections from Dasylirion in Arizona (AZ2) have fairly numerous translucent pores on the hind coxae. In a second sequenced Arizona collection (AZ3; also from Dasylirion ), 12 of 16 females lack translucent pores on the coxae, three females have a few pores on just one coxa and a fourth female has pores on both coxae. The function of translucent pores is not known but it is possible that sex pheromones may be released from the pores ( Williams 1985a).

Specimens from two populations collected from Jamaica show some differences from other specimens of F. dasylirii in having a higher number of small oral-collar tubular ducts marginally on abdominal segments VII and VIII (15–25 on each side of VII; 10–21 on each side of VIII) ( Fig. 8c). Specimens from the Galapagos Islands ( Ecuador), one adult female from Hawaii, and a few others from Florida and Brazil differ from all other females of F. dasylirii in possessing transverse clusters or short rows of small oral-collar tubular ducts submedially on abdominal segments IV–VII at each end of the row of multilocular pores of those segments, with 0–9 ducts on each side of IV; 1–8 on each side of V; 4–10 on each side of VI; 3–6 on each side of VII; and 0–5 on each side body on margin of VIII ( Figure 8e). In addition, the minute discoidal pores on the medial to submedial derm of the ventral abdomen of the latter specimens are 2.5–4.5 µm in diameter and with a distinct rim (the pores in typical specimens of F. dasylirii are <2.5 µm in diameter and with an indistinct rim). The latter specimens may represent a distinct species; additional specimens and DNA data would help to establish the taxonomic status of this morphological form.