Protortonia Townsend, 1898

Williams, D. J. & Gullan, P. J., 2008, A revision of the Neotropical scale insect genus Protortonia Townsend (Hemiptera: Coccoidea: Monophlebidae: Llaveiini), Journal of Natural History 42 (1 - 2), pp. 77-128 : 83-89

Protortonia Townsend

Protortonia Townsend in Townsend and Cockerell, 1898:169.

Type species: Ortonia primitiva Townsend , by monotypy and original designation.

Descriptions of instars of Protortonia

Adult female

Body membranous, broadly oval, posterior end rounded; sometimes attaining a length of over 9.0 mm. Eyes conspicuous, sclerotized, oval and convex. Antennae each normally 11-segmented (occasionally one antenna 9–10-segmented, apparently a developmental abnormality), first and second segments broad, remaining segments gently tapering towards apical segment; each segment bearing flagellate setae, apical segment also with a few fleshy setae. Legs well developed, tibia and tarsus well articulated, each segment with flagellate setae, inner edges of tibia and tarsus with stout, almost lanceolate setae. Trochanters each with three campaniform sensilla on each surface; distal trochanteral seta long and conspicuous. Claws each stout and curved; plantar edge with a series of minute denticles; claw base with a pair of claw digitules, apparently knobbed apically, mostly damaged in specimens examined. Clypeolabral shield and labium degenerate; sclerotized remains of tentorium and outline of labium often faintly discernible. Thoracic spiracles each well developed and heavily sclerotized, with bar present. Abdominal spiracles numbering seven pairs; a pair present on segment VIII lateral to apical cluster of setae; pairs also present on each anterior abdominal segment as far forward as segment II, each spiracle with atrium about as long as diameter of orifice. Anal opening situated well forward from apex of abdomen, surrounded by slender flagellate setae; inner ring of anal tube usually discernible, sclerotized and simple. Vulva often situated almost opposite anal opening. Cicatrices numbering three, situated between vulva and apex of abdomen; middle cicatrix largest, usually elongate and constricted at middle; outer cicatrices smaller and reniform.

Dorsal surface with multilocular disc pores present, each pore usually with an oval or distorted, sunken centre, surrounded by 12–22 outer loculi; fairly numerous, often concentrated near segmental margins or abundant over entire surface; similar pores, but with three or four central loculi, sometimes present near head margin; pores around anal opening sometimes with different arrangement of loculi. Minute duct-like pores sometimes present, shallow, either circular or longer than wide. Abundant short setae, each either flagellate or dagger-shaped, with conspicuous collars, found on all segments. Marginal setae on each segment usually long and stout, present in groups of at least two, usually more; setae on abdominal segments VII and VIII usually longest, with large, conspicuous collars. Other dorsal setae present, thicker than normal dorsal setae, often noticeable in possessing larger collars; sometimes very slender setae present with dome-shaped collars.

Ventral surface with multilocular disc pores, usually similar to those on dorsum, except larger pores sometimes situated around vulva. Minute duct-like pores often present in marginal areas. Setae, often similar to those on dorsum, present in marginal and submarginal areas, usually more slender in medial areas; thicker and longer setae usually present around margins and head; intermediate size setae usually mingled with normal slender setae.

Third-instar nymph

Similar in shape to adult female. Eyes oval. Antennae each with eight or nine segments. Legs well developed, similar to those of adult female but shorter and with fewer setae; trochanters each with three campaniform sensilla on each surface. Claws each stout, usually with three or four minute denticles; claw digitules usually about as long as claw, apically knobbed. Clypeolabral shield triangular, heavily sclerotized. Labium basically 3-segmented, basal segment minute, sometimes difficult to discern. Thoracic spiracles similar to those in adult female. Abdominal spiracles numbering seven pairs, similar to those of adult female but smaller. Anal opening usually conspicuous, surrounded by slender flagellate setae; anal tube with simple inner ring. Cicatrices numbering three, similar in shape to those of adult female but smaller.

Dorsal surface with multilocular disc pores, each often with three, four or five large central loculi, rarely with two, in sunken centre surrounded by ring of outer loculi with thin walls, sometimes outer loculi numerous depending on species; pores scattered over surface, sometimes abundant; pores surrounding anal opening often each with oval centre. Minute duct-like pores usually present, sometimes sparse. Setae numerous, similar to those in adult female but less abundant. Conspicuously long, stout setae usually present on posterior abdominal segments and sometimes present on thorax.

Ventral surface with multilocular disc pores similar to those on dorsum, except commonest locular numbers can differ between on those on venter and dorsum. Minute duct-like pores sparsely present around margins. Setae present in marginal areas similar to those on dorsum, usually more slender in medial areas.

Second-instar nymph

Similar to third-instar nymph but setae and pores much less abundant. Antennae each with six segments. Legs well developed; trochanters each with two campaniform sensilla on each surface. Claws each with at least two minute denticles. Clypeolabral shield and labium well developed. Thoracic spiracles more slender than in adult and third-instar nymphs. Abdominal spiracles numbering seven pairs. Anal opening surrounded by slender flagellate setae; anal tube with simple inner sclerotized ring.

First-instar nymph

Body oval. Eyes oval, convex. Antennae each 6-segmented. Legs well developed, slender; trochanters each with two campaniform sensilla on each surface. Claws each slender, with single denticle near tip; claw digitules apically knobbed. Clypeolabral shield and labium well developed. Thoracic spiracles sclerotized. Abdominal spiracles numbering seven pairs, sometimes difficult to locate, each with atrium about same size as a pore. Anal opening simple, not surrounded by rows of flagellate setae as in other instars; anal tube conspicuous. Cicatrix single, circular, medial on last abdominal segment.

Dorsal surface with multilocular disc pores present, not numerous, each with 1–4 central loculi surrounded by a ring of thin-walled outer loculi. Minute duct-like pores sometimes discernible. Distinctive, enlarged spine-like setae present in groups near midline and lateral margins, each seta with a wide but narrow collar and a more or less oblong or oval middle part, either minutely reticulate, nodulose or alveolate, remainder of seta narrower and tapering, bluntly pointed. Conspicuous long, flagellate setae present on margins of abdominal segments VI to VIII or VII and VIII; other long setae present on thorax and sometimes on head. Shorter flagellate setae distributed near enlarged setae.

Ventral surface with multilocular disc pores scattered, similar in structure and distribution to those on dorsum. One or two long setae present marginally on each side of abdominal segment VIII; other setae more slender than those on dorsum.

Adult male

Very few male specimens are available for Protortonia . Schrader (1930) describes the mating behaviour of the adult males of P. near primitiva , but none of his specimens could be located. Foldi (2006) described the adult male of P. ecuadorensis in detail, but the adult males of no other species of Protortonia have been described. However, one adult male of an undescribed Protortonia species from Boyacá, Colombia, is available ( USNM) and it shares the following features, which may be diagnostic of the genus, with the adult male of P. ecuadorensis : antennae each with 10 segments, each flagellar segment elongate, without swellings (each flagellar segment is trinodose in Llaveia oaxacoensis Morrison and Laurencella colombiana ) and with two pairs of long, fleshy caudal extensions on the posterior abdomen [three pairs in L. oaxacoensis ( Morrison 1928) , four pairs in L. colombiana ( Foldi and Watson 2001; Hodgson and Foldi 2006)].

Third-instar male

We have only one definite third-instar (prepupal) male of Protortonia . This specimen was inside a loose wax ‘‘cocoon’’ on a dry cactus cladode with other dry material of P. cacti (ex Opuntia dillenii from Henry Tryon, apparently collected in San Juan, Puerto Rico) and was slide-mounted by PJG in 2006. This specimen is in poor condition but there is a pupa (with wing pads) forming inside. The third-instar male cuticle is female-like in lacking developing wings (except for the cuticle of the pharate pupa) and in having three cicatrices on the posterior ventral abdomen. However, unlike the third-instar female, it has vestigial mouthparts and presumably does not feed. It appears to have 9-segmented antennae and a simple, sclerotized anal ring, wider than that of the associated third-instar female.

Diagnostic features

Adult females of Protortonia can be distinguished from those of other genera of the Llaveiini by the (i) vestigial mouthparts (sometimes remnants of tentorium and outline of labium visible), (ii) three ventral cicatrices posterior to the vulva, and (iii) distinctive multilocular disc pores, each with an oval and sunken centre surrounded by a ring of 12–22 loculi. Adult females of Protortonia look superficially similar to adult females of many species of the Iceryini, especially in the position and number of the cicatrices, but among other features Protortonia always has seven pairs of abdominal spiracles, compared with 2–4 in Iceryini ( Morrison 1928; Unruh and Gullan 2008). The adult females of the different Protortonia species are quite similar to each other morphologically, whereas the female nymphs of each species are usually quite distinct from those of other species. For example, compare the illustrations of the first-instar nymphs of P. azteca ( Figure 2 ), P. cacti ( Figure 5 ) and P. navesi ( Figure 13 ) and note the distinct differences in the dorsal and marginal setae. In contrast, the adult females of these same three species ( Figures 1 , 3 , 10 ) are extremely similar and it takes careful study of well-prepared specimens to see the differences in pore size and setal size and shape among these species. The first-instar nymphs of Protortonia can be distinguished from those of other genera of Llaveiini by the following combination of features (characteristics of other genera in parentheses): (i) a single ventral cicatrix (1–7 cicatrices), (ii) the presence of very long marginal setae on one to three thoracic segments and only the last two or three posterior abdominal segments (on all or almost all body segments), and (iii) distinctive, enlarged dorsal spine-like setae, each with the region above the collar minutely reticulate, nodulose or alveolate, and distinct from the smooth and tapered apical region (spine-like setae usually not or barely textured). The potentially diagnostic features of the adult male of Protortonia are discussed above under ‘‘Adult male’’.

Comments on species diversity

We believe that there may be at least twice as many species of Protortonia as are dealt with in the following key. Here we discuss other available specimens that are not satisfactory for description but which appear to represent new species. We have seen a species intercepted at New York, USA, from the Dominican Republic on Melicoccus bijugatus (Sapindaceae) that is similar to P. cacti and P. azteca , represented by a single adult female and a second-instar nymph (USNM), but the specimens are not in satisfactory condition to describe and illustrate. We have seen a single adult female of each of two species from Guatemala (AUCC), one on Acacia angustissima (Fabaceae) from Guatemala [city] which is too poor to identify to species and the other on an unknown host from Chicol, Huehuetenango, which has multilocular disc pores with 3–6 central loculi in addition to typical pores with a unilocular centre. From Mexico, there are two adult females (USNM) of a species similar to P. ecuadorensis and P. quernea sp. nov. and collected on Quercus sp. ( Fagaceae ) at Santa Rosa, but we could not see cicatrices on these specimens. Another species from Mexico, intercepted at Hoboken, USA, on Sedum sp. ( Crassulaceae ) is represented by a single poor adult female specimen (BME). Also available are third-instar nymphs and an adult male only (USNM) of a species collected at Boyacá, Colombia, on an unidentified plant, and a third-instar female (USNM) of another species from Jamaica on Elaphoglossum latifolium (Lomariopsidaceae) . In addition, there are unconfirmed records from Colombia of unidentified Protortonia species on Acacia sp., Cupressus lusitanica (Mexican cypress; Cupressaceae ), Pinus sp. ( Pinaceae ), Prunus persica (peach) and Pyrus communis (pear) (both Rosaceae ) ( Ochoa Lázaro 1989), and on Salix humboldtiana (Salicaceae) ( Gallego and Vélez Angel 1992). Foldi (1995) lists P. primitiva on Pinus oocarpa from Puerto Lobo, Mexico, but this record might be the pine-feeding species P. azteca . Furthermore, the species discussed by Schrader (1930, 1931) as P. primitiva (Townsend) from Guatemala differs from the original material of P. primitiva (Townsend) from Mexico, but is represented by a single adult female only (USNM) that we have not been able to describe or illustrate. This specimen is discussed further under ‘‘ P. Primitiva ’’.

Key to adult females of Protortonia species (except P. crotonis )

1. Multilocular disc pores abundant, present in bands across dorsal segments, each band usually at least nine pores deep in submedial areas of each abdominal segment................................................ 2

- Multilocular disc pores, although numerous on dorsum, each band usually only about four pores deep in submedial areas of each abdominal segment. Multilocular disc pores, although numerous on dorsum, each band usually only about four pores deep in submedial areas of each abdominal segment................................................. 3

2. Dorsal setae mostly 50–75 Mm long, about 2 Mm wide at base, collars mostly 10 Mm wide at base, 5–8 Mm long. Multilocular disc pores all with a single oval centre......................................... quernea sp. n.

- Dorsal setae mostly 30–40 Mm long, about 2.5 Mm wide at base, collars about 7.5 Mm wide at base, 5 Mm long. Many multilocular disc pores on head and anterior thorax with a trilocular or quadrilocular centre.. ecuadorensis Foldi

3. Dorsal setae mostly dagger-shaped, most at least 5 Mm wide at base, with collars about 20 Mm wide at base and 10 Mm long.............. navesi Fonseca

- Dorsal setae mostly flagellate, 2.0–5.0 Mm wide at base, with collars at most 10– 15 Mm wide at base and 5.0–7.5 Mm long......................... 4

4. Dorsal setae mostly 45–90 Mm long, many with collars 10–15 Mm wide at base. Multilocular disc pores around vulva about same size as those elsewhere on venter, and with similar number of outer loculi...... primitiva (Townsend)

- Dorsal setae mostly 20–45 Mm long, majority short, with collars only 7.5– 12.5 Mm wide at base. Multilocular disc pores around vulva usually larger than those elsewhere on venter, and with more outer loculi............... 5

5. Dorsal and ventral multilocular disc pores mostly with 12–14 outer loculi; pores around anal opening and vulva usually with 16 outer loculi. Inner tibial setae 40–75 Mm long. Medial cicatrix 240–300 Mm wide.......... azteca (Ferris)

- Dorsal and ventral multilocular disc pores mostly with 16–18 outer loculi, including those around anal opening; pores around vulva with up to 22 outer loculi. Inner tibial setae 30–40 Mm long. Medial cicatrix 175–240 Mm wide................................................... cacti (L.)

Key to available third-instar females of Protortonia

1. Dorsal setae short, mostly 12–20 Mm long, only about 1.0–2.0 Mm wide at base of shaft (wider across collar). Multilocular disc pores usually with five or six central loculi and 12–16 outer loculi...................... cacti (L.)

- Dorsal setae longer, 20–60 Mm long, at least 5.0 Mm wide at base of shaft (wider across collar). Multilocular disc pores normally with 3–5 central loculi and 10 or fewer outer loculi......................................... 2

2. Most dorsal setae stiff but not spine-like, usually 50–60 Mm long, Ḳ5.0 Mm wide at base of shaft, with rounded or slightly expanded apex.............. 3

- Most dorsal setae spine-like and shorter, 20–50 Mm long, often 5–10 Mm wide at base of shaft, with acute apex................................. 4

3. Most dorsal setae with apex rounded and slightly expanded. Multilocular disc pores abundant, at least nine deep on each dorsal segment near margin. ecuadorensis Foldi

- Most dorsal setae with apex rounded but not expanded. Multilocular disc pores less abundant, only about three or four deep on each dorsal segment near margin................................... primitiva (Townsend)

4. Dorsal setae elongate spine-like, at most 5.0 Mm wide at base of shaft. Longest

marginal dorsal setae on posterior abdominal segments 340 Mm long......

............................................. crotonis Reyne - Dorsal setae almost dagger-shaped, 5–10 Mm wide at base of shaft. Longest

marginal dorsal setae on posterior abdominal segments about 440 Mm long.

navesi Fonseca

Key to available second-instar nymphs of Protortonia

1. Dorsal setae with apex blunt or slightly expanded (capitate). Dorsal multilocular disc pores 7.5–15.0 Mm in diameter, mostly with four or five central loculi.. 2 - Dorsal setae with apex acute. Dorsal multilocular disc pores 6.5–7.5 Mm in diameter, mostly with three or four central loculi................... 3

2. Typical dorsal setae 5–50 (mostly 10–30) Mm long, 1.5–2.5 Mm wide at base, with apex blunt or slightly expanded. Dorsal multilocular disc pores 10–15 Mm in diameter with 1–5 (mostly four or five) central loculi.......... cacti (L.)

- Typical dorsal setae 18–90 (mostly 50–65) Mm long, 2.5–4.0 Mm wide at base, all with apex slightly expanded. Dorsal multilocular disc pores 7.5–10.0 Mm in diameter with 1–6 (mostly four or five) central loculi..... ecuadorensis Foldi

3. Typical dorsal setae slender with acute apex, 25–40 Mm long (longer around anal opening), 2.5–4.0 Mm wide at base.................... crotonis Reyne

- Typical dorsal setae almost dagger-shaped, of 2 sizes: (i) 15–25 Mm long, 5–8 Mm wide at base; (ii) 50–100 Mm long, 6.0–7.5 Mm wide at base... navesi Fonseca

Key to available first-instar nymphs of Protortonia (for P. crotonis based on one exuviae only)

1. Dorsal margin with a series of long flagellate setae on all segments; conspicuously longer seta present on either side of mesothorax and each posterior abdominal segment only.................... navesi Fonseca

- Dorsal margin without a series of long flagellate setae, except for a conspicuously long seta present on each side of one or more segments of thorax and on posterior abdomen; if other flagellate marginal setae present, they are short................................................... 2

2. On thorax, conspicuously long marginal setae confined to one on each side of prothorax only...................................... cacti (L.)

- On thorax, conspicuously long setae present one on each side of prothorax, mesothorax and metathorax, with setae on prothorax shortest.......... 3

3. Conspicuously long marginal seta on prothorax about 55 Mm long. Distinctive enlarged dorsal setae scant (fewer than 20) in dorsal medial and submedial areas of thorax and abdomen........................... crotonis Reyne

- Conspicuously long marginal seta on prothorax 140–160 Mm long. Distinctive enlarged dorsal setae numerous (more than 80) in dorsal medial and submedial areas of thorax and abdomen........................ azteca (Ferris)

Figures

Figure 1. Protortonia azteca (Ferris), adult female.

Figure 2. Protortonia azteca (Ferris), first-instar nymph.

Figure 3. Protortonia cacti (L.), adult female.

Figure 5. Protortonia cacti (L.), first-instar nymph.

Figure 10. Protortonia navesi Fonseca, adult female.

Figure 13. Protortonia navesi Fonseca, first-instar nymph.