Carpochloroides viridis Cockerell

Hodgson, Chris & Miller, Dug, 2010, A Review of the Eriococcid Genera (Hemiptera: Sternorrhyncha: Coccoidea) of South America 2459, Zootaxa 2459 (1), pp. 1-101 : 30-37

Carpochloroides viridis Cockerell

Carpochloroides viridis Cockerell, 1899: 12–13 .

Material examined: No data but labelled Carpochloroides viridis Cockerell, Cornell University No. 981 ( USNM): 2/2adff (1 in fair condition, other poor) + 1/6 first-instar nymphs, many in egg membranes (mostly in poor condition; because of the size of the dorsal setae, many other structures hard to discern with certainty). Slides that were acquired with the Cockerell Collection are labelled as this species, “Co-type, on Eugenia , Brazil, Cockerell Coll.” ( USNM): 1/1 adf (in poor condition) and 1/12 first-instar nymphs (in fair condition) + 1/ 9 first–instar nymphs (good condition). On Eugenia, Ypiranga , Brazil, Hempel ( USNM): 1/1 adf (fair condition, but rather distorted) + 8 first-instar nymphs (in poor condition), + 1/1 adm (in fair to good condition but wings distorted and only 1 antenna complete) + 1/7 first-instar nymphs (fair condition). True type material not seen, but slides from the Cockerell Collection could be from the type series. Type collection data: Brazil, Sao Paulo, Campinas, on Eugenia sp. (Myrtaceae) , -. ix.1898, F. Noack.

Adult female ( Fig. 12 )

Unmounted material. “Adult females, as seen alive, clear green, in form and colour very much like the young fruits of Eugenia .” ( Cockerell, 1899).

Mounted material. Body apparently globular, derm entirely membranous. Length 4–6 mm, width 4–6 mm. No body segmentation visible and without any indications of a margin; apparently without pro- and mesothoracic legs; metathoracic legs and antennae reduced to small prominences.

Dorsum. Derm membranous but with a faint pattern of quite large reticulations probably throughout most of dorsum and on areas laterad to spiracles. Dorsal setae all finely spinose, each 25–35 µm long, with broad basal sockets; very sparse, perhaps most abundant posteriorly on abdomen. Macrotubular ducts absent. Microtubular ducts rather small, each outer ductule 5–7 µm long (rather shorter than those found ventrally), apparently without an inner filament; relatively sparse, with 1 in most dorsal reticulations but perhaps absent posteriorly in areas with loculate pores. Loculate pores with 3–5 (mainly 5) loculi, each 5 µm wide, with 1 per reticulation over a broad area anterior to anal ring. Anal lobes entirely absent; with a group of about 3 setae on each side of anus, but without any clearly defined anterior or posterior suranal setae. Median lobe absent. Anal opening in centre of a sclerotised ring, without setae or pores; outer diameter of sclerotised ring about 25 µm wide; actual opening oval, about 18 µm wide and 5 µm long.

Margin. Undefined, without any setae differentiated from dorsal or ventral setae. However, approximate lateral margin of venter possibly indicated by absence of loculate pores. Eyespots not detected.

Venter. Derm membranous. Setae sparse and similar to those on dorsum; those posteriorly on abdomen possibly in transverse segmental rows but this not clear. Macrotubular ducts absent. Ventral microtubular ducts longer and narrower than those on dorsum; frequent throughout but perhaps less abundant medially posterior to labium. Loculate pores similar in structure to those on dorsum; exact distribution uncertain but apparently in a broad submarginal band which extends mesad to spiracles but absent medially on thoracic and most abdominal segments but with a few pores present just anterior to anus; also with a small group of 7–9 on fleshy margins of spiracular cavity.

Antennae roundish (about 28 µm wide), mildly sclerotised, apparently 1 segmented, with perhaps 3 fleshy setae but structure hard to determine; with a short stout spinose seta laterad to base. Clypeolabral shield large, about 275 µm long, but mouthparts with (barely present on female labelled from Ypiranga on oldest specimens) 2 large internal, sclerotised plate-like apodemes extending anteriorly from anterior margin of the tentorial box, each about 175 µm long and total width about 500 µm; labium probably 2 segmented, with perhaps 5 pairs of setae. Spiracles large, with a wide, strong ventral valve, each in a fairly deep cavity: width of peritremes 90–92 µm. Pro- and mesothoracic legs absent; metathoracic legs represented by a pair of short membranous protuberances, each about 15 µm wide and 8 µm long (not detected on female labelled from Ypiranga); with little sign of any segmentation and no setae or translucent pores; located about mid-way between posterior spiracles and anus. Vulva present a short distance anterior to anus. Also with a pair of small, cone-shaped tubular structures located just anterolaterally to anus, each broadest on derm surface (where duct about 6–8 µm wide), each about 10–12 µm long ending distally in a filamentous inner ductule (not detected on female labelled from Ypiranga).

Comment. The function of the small cone-shaped tubular structures anterolateral to the anus is unknown but similar structures have been noted previously on the adult females of Pseudotectococcus anonae Hempel and P. rolliniae Hodgson & Gonçalves ( Hodgson et al., 2004) , which also are gall-inducing eriococcids from South America.

The apodemes associated with the mouthparts probably become more sclerotised with age, as they are barely visible on the youngest specimen. This almost certainly also applies to other species in which these apodemes have been noted, i.e. Aculeococcus morrisoni and Tectococcus ovatus .

For a comparison of the adult female of Carpochloroides with other eriococcid genera, see the “Comment” section after the generic diagnosis of Carpochloroides .

First-instar nymph (gender not determined) ( Fig. 13 )

Unmounted material. Not seen.

Mounted material. Body oval, slightly more pointed posteriorly. Length about 320–410 µm, width 140– 160 µm (but venter perhaps slightly wider).

Dorsum. Derm membranous, without obvious dermal spinules. Dorsal setae strongly spinose; many very long, broadest basally and gradually narrowing to a sharp point, highly variable in length but those medially much longer (longest each 75–100 µm and shortest 15–50 µm), with a narrow basal socket, distributed as follows: mainly in 4 medial lines, with 2 pairs (outer much shorter than inner pair) on abdominal segments I and on pro-, meso- and metathorax and posteriorly on head + 1 pair medially on head; also with submarginal setae as follows: 1 on each side of abdominal segment I, 1 pair on each side of meso- and metathorax but none on prothorax (outer seta of each pair much longer than inner seta); also 1 submarginally on each side near posterior margin of head and 2 submarginally on anterior margin. Other types of setae absent. Discoidal pores (probably modified microtubular ducts) appearing as dark spots, probably representing a sclerotised dermal ring: with single ducts submarginally on abdominal segments III – VI, and submedially on pro-, meso- and metathoracic segments, plus another submedially on head. Loculate pores and macrotubular ducts absent. Anal lobes absent or rounded, membranous; each with a very long, flagellate apical seta about 300+ µm long; marginally with 2 pairs of spinose setae between long apical setae, outer pair 20–30 µm long, inner pair about 8–13 µm long .

Margin. Marginal setae strongly spinose, similar in shape to those on dorsum, rather variable in length, each 25–70 µm long; with 4 between eyespots, 4 or 5 on each side between eyes and point opposite anterior spiracles, 4 or 5 on each side laterally between anterior spiracles and abdomen and with 1 on each side of abdominal segments I–VII. Eyespots rather pronounced, each 13–15 µm wide, situated on margin near base of antennae.

Venter. Derm membranous. With 3 pairs of long setae between antennae plus pairs mesad of meso- and metacoxae, each about 30–35 µm long; no setae present medially on abdominal segments I–VII. Submarginal setae strongly spinose, similar to marginal setae but smaller, each 13–30 µm long; with a pair on each side of abdominal segments I–VII, 1 on each side of thorax and 1 pair anteriorly on head. Discoidal pores apparently similar in structure to those on dorsum, with single pore anterior to each posterior spiracle, posterior to each anterior spiracle and posterior to each scape. Loculate pores probably with mainly 5–7 loculi: with 2 or 3 immediately laterad of each peritreme, in a small cavity, plus 1 postero-laterally to each posterior peritreme near margin. Anus slightly oval, membranous, width perhaps 13–16 µm; without an anal ring or anal ring setae; with 1 small seta on each side, each about 10–12 µm long, plus a much longer seta, 80–100 µm long.

Antennae 6 segmented, 124–128 µm long; setal distribution: scape 3; pedicel 1 + campaniform sensillum; segments III and IV 1 hair-like seta; V 2 setose seta + 1 fleshy seta; VI 3 fleshy setae, 5 short setose setae plus 2 long setae, length of longest about 80–90 µm. Clypeolabral shield about 70–105 µm long; labium 2 segmented; about 50 µm wide, with possibly 5 pairs of short setae on apical segment and 1 pair on proximal segment. Spiracles small, peritremes possibly 5 µm wide, each with a lateral membranous cavity with loculate pores, cavity about 14–17 µm wide. Legs well developed; lengths (metathoracic leg (µm)): coxa 35–45; trochanter + femur 65–80; tibia 38–50; tarsus 33–45; claw 17–23; setae: coxae 2 or 3, trochanter 2, femur 3, tibia 4, tarsus 3; long trochanter seta 65–70 µm long; tarsal campaniform sensillum present; claw with a distinct denticle; both tarsal digitules on all legs capitate and equal in size; claw digitules alike with small capitate apices.

Comment. The first-instar nymphs of C. viridis are immediately recognisable by the extremely long spinose setae on the head, thorax and anterior abdominal segments, almost all of which are longer than the width of a segment. In addition, they have: (i) 6-segmented antennae; (ii) discoidal pores (probably modified microtubular ducts) on both dorsum and venter; (iii) anal lobes not differentiated and unsclerotised; (iv) loculate pores resticted to a cavity laterad to each spiracle (apart from 1 pore just posterior to each posterior spiracle); (v) no cruciform pores, (vi) no median anal plate; and (vii) dorsal setae of just 1 type. The firstinstar nymph of Exallococcus laureliae Miller & González also has rather large dorsal spinose setae but these are much shorter than on C. viridis , and it also differs in having: (i) the anal lobes differentiated and sclerotised; (ii) loculate pores in a fairly wide longitudinal band ventrally; (iii) cruciform pores; and (iv) a median anal plate.

Adult male ( Fig. 14 )

Unmounted material. Not seen.

Mounted material. Total body length 1.3 mm; width across triangular plates 360 µm. Antennae quite long, about 2/3rds total body length; capitate setae only present on apical segment. Body rather setose, almost all fleshy setae (fs); fleshy setae (fs) long and rather similar to hair-like setae (hs). Pores absent. Wings perhaps slight shorter than total body length; without alar setae or alar sensory pores. Hamulohalteres absent.

Head: approximately rather square oval, broadest across genae; length about 230 µm, width across genae about 220 µm. Dorsally, median crest (mc) not demarcated and not reticulated but with a short, indistinct midcranial ridge (dmcr), not reaching postoccipital ridge posteriorly; postoccipital ridge (por) indistinct, with arms extending only laterally; with (on each side) about 10 fs dorsal head setae (dhs); pores absent. Ventral mid-cranial ridge (vmcr) quite short and narrow, extending from lateral arms (lmcr) posteriorly almost to ocular sclerite (ocs); without any reticulation laterally. Genae (g) not reticulated; each with 12–14 fs genal setae (gs). Eyes: with 2 pairs of round, simple eyes; dorsal eyes (dse) placed rather anteriorly, above each scape, each about 35–37 µm wide; ventral simple eyes (vse) placed quite far posteriorly, each about 40–43 µm wide. Ocelli (o) large, situated laterally, each about 16 µm widest, just touching postocular ridge and with an extension up dorsal margin. Ocular sclerite (ocs) only partially sclerotised, particularly around and between simple eyes; lightly reticulated around each vse. Preocular ridge (procr): short both dorsally and ventrally, latter possibly extending to anterior to each vse. Postocular ridge (pocr) strongly developed, extending posteroventrally from near dmcr postero-laterally past each ocellus to neck region. Interocular ridge probably absent. Dorsal ocular setae absent. Ventral mid-cranial ridge setae (vmcrs) 1–3 fs on each side; ventral head setae (vhs): with 6–9 fs + 2–9 hs on each side just anterior to and laterad of each vse, plus about 7 hs medially posterior to vse. Preoral ridge (pror) absent. Cranial apophysis (ca) not detected.

Antennae: 10 segmented and filiform; about 875 µm long (ratio of total body length to antennal length 1:0.67). Fleshy setae several times longer than width of flagellar segments, each about 65–75 µm long. Scape (scp) 50 µm long and 66 µm wide, with 3 fs + 2 hs. Pedicel (pdc) 60 µm long, 54 µm wide; with a few faint concentric ridges, mainly on distal half; with 6 hs + 16 fs; no campaniform pore detected. Segments III–X each about 16–25 µm wide; lengths of segments (µm): III 128; IV 95; V 112; VI 125; VII 103; VIII 75; IX 75 and X 70; approximate number of setae per segment: III–IX 16–20 fs + 0–3 hs; VIII & IX both with 1 antennal bristle, and segment X with 13 fs, 3 hs, 3 or 4 large + 2 smaller antennal bristles + 3 capitate setae (caps); sensilla basiconica not detected.

Thorax. Prothorax: pronotal ridge (prnr) well-developed but not fused dorsally; pronotal sclerite (prn) represented by a large triangular area dorsolaterally; without lateral pronotal setae (lpns). Medial pronotal setae absent; post-tergite not detected, without post-tergital setae or pores. Sternum (stn 1) lightly sclerotised, with a distinct median and transverse ridge; without prosternal setae (stn 1 s) or pores. Anteprosternal setae (astn 1 s) absent.

Mesothorax: prescutum (prsc) oval, 70 µm long, 130 µm wide; sclerotised but not reticulated, without prescutal setae; prescutal ridge (pscr) well developed anteriorly but quickly narrowing; prescutal suture (pscs) present. Scutum (sct): median area without unsclerotised areas; distance of prescutum from scutellum about 70 µm; scutal setae (scts) few, perhaps 4 hs on each side; lateral margins with faint ridging laterad to prescutum; prealare (pra) and triangular plate (tp) well developed. Scutellum (scl) 170 µm wide and 75 µm long; with a distinct scutellar ridge but no foramen; scutellar setae (scls): 3 or 4 fs on each side; postnotal wing process (pnp) relatively narrow but broadening laterally. Basisternum (stn 2) 255 µm wide and 135 µm long; without a median ridge; bounded anteriorly by a strong marginal ridge (mr), very broad antero-laterally, and with strong precoxal ridges (pcr 2); with about 50 fs basisternal setae (stn 2 s); lateropleurite (lpl) broad, with a distinct extension from marginal ridge along anterior margins; furca (f) well developed, narrow-waisted, arms very divergent and extending almost to marginal ridge anteriorly. Mesopostnotum (pn 2) well developed; postnotal apophysis (pna) well developed. Area bounded anteriorly by scutellum and laterally and posteriorly by mesopostnotum not sclerotised. Mesepisternum (eps 2) not reticulated; subepisternal ridge (ser) long and well developed. Postalare (pa) without postalare setae (pas). Mesothoracic spiracle (sp 2): width of peritreme 30 µm, without associated loculate pores. Postmesospiracular setae (pms) 4–7 fs + 1–3 hs on each side. Tegula (teg) present, with about 3 fs + 2–4 hs tegular setae (tegs).

Metathorax: with perhaps 4 fs metatergal setae (mts) on each side and 2 fs laterally on each side; metapostnotal sclerite (pn 3) not detected. Dorsospiracular setae (dss) about 11 fs on each side. Dorsal part of metapleural ridge (plr 3) absent; ventral part of metapleural ridge well developed; metepisternum (eps 3) sclerotised, each with 3 fs postmetaspiracular setae (eps 3 s) plus 6 fs in a group slightly more medially; precoxal ridge (pcr 3) absent; metasternal apophysis (st 3 a) present. Metepimeron (epm 3) sclerotised, but without setae. Antemetaspiracular setae absent. Metathoracic spiracle (sp 3): width of peritreme 32 µm, without loculate pores. Metasternum (stn 3) membranous, with about 12 fs anterior metasternal setae (amss) and 7 fs + 2 hs posterior metasternal setae (pmss).

Wings: both misshapen; hyaline, maybe 1075 µm long; width unknown; alar lobe (al) probably absent; alar setae absent; circular sensoria absent. Hamulohalteres absent.

Legs: mesothoracic leg shortest. Fleshy setae (fs) present on all segments, each as long as on body; setose but with many setae broken and therefore fs and hs not distinguishable. Coxae (cx): I 120; II 108; III 116 µm long; long setae on each coxa not differentiated. Trochanter (tr) + femur (fm): I 245; II 225; III 245 µm long; each trochanter with 2 campaniform sensoria on each side in a line; long trochanter seta not differentiated. Tibia (tib): I 205; II 205; III 205 µm; some setae spur-like on distal half; with 2 apical spurs (tibs) on all legs; length of longest 20–25 µm. Tarsi (tar) 1 segmented, length: I 80; II 85; III 85 µm long (ratio of length of tibia III to length of tarsus III 1:0.41); tarsal spurs differentiated, each about 25 µm long; tarsal campaniform pore present; tarsal digitules (tdt) capitate, slightly shorter than claw. Claws (c) subequal to or slightly longer than width of tarsus; length III: 30 µm; with a small denticle; claw digitules (cdt) with small capitate apices and slightly longer than claw.

Abdomen: segments I–VII: tergites and sternites unsclerotised and without obvious oval membranous areas in inter-segmental membranes. Caudal extension (ce) of segment VII absent. Without loculate pores. Dorsal setae (ads) almost all fs, each about 50 µm long, in bands of about 16–20 across segments; pleural setae: dorsopleural setae (dps) all fs in groups of about 10–15 on each side; ventropleural setae (vps) all or mainly fs, in a small group of about 4 setae, but these often joined to dps; ventral setae (avs) sparser than ads, mainly fs, each slightly shorter, about 40 µm long.

Segment VIII: both tergite (at) and sternite (as) sclerotised in transverse bands; tergite with about 30 fs ads, extending full width of segment; sternite with about 20 fs avs across segment; caudal extension (ce) rounded, each with about 14 or 15 fs + 1 hs pleural setae, 1 rather longer (about 60 µm long) and stouter than other setae. Glandular pouches (gp) present, moderately deep, each with loculate pores restricted to within pouch; each with 2 non-capitate setae (gls) about 120 µm long.

Genital segment [note: segmentation in description below speculative!]: apparently divided into 2 sections: (i) a broad anterior part (probably representing segment IX) and a short, triangular posterior section; anterior part about 140 µm wide at base and about 60 µm long; dorsally perhaps not heavily sclerotised with about 30 fs extending across segment and with an oval area medio-posteriorly with about 9 long hs ante-anal setae (aas) and with anal opening present along posterior margin; ventrally, anterior part (segment IX) probably heavily sclerotised, particularly near lateral margins, with about 8 fs on each side and about 9 hs on either side of central indentation. Penial sheath (ps) proper (posterior part) pointed but slightly constricted near mid-length; 70 µm wide at base and 90 µm long; with 7–9 long setae (pss) on each side, apparently restricted to ventral and lateral margins. Basal rod (bra) very short, about 10 µm long. Aedeagus C-shaped, scythe-like, total length perhaps 65 µm. Tip of penial sheath with a few minute pores (psp).

Comment. The adult male of C. viridis can be separated from the other known male eriococcids from the Neotropics in having the following combination of characters: (i) 10 segmented antennae, with fleshy setae clearly longer than width of antennal segments; (ii) very short penial sheath, length subequal to basal width; (iii) hair-like and fleshy setae all long, rather similar and hard to separate; (iv) tarsi all 1 segmented; and (v) dorsal abdominal setae more abundant than ventral abdominal setae (other way round on all other species). In having only 1-segmented tarsi, the male of C. viridis is unusual for Eriococcidae . The only other adult male eriococcid known to the authors with well-developed legs which appears to have a 1 segmented tarsus are Opisthoscelis verrucula Froggatt and Cylindricoccus spiniferus Maskell , both from Australia but these two species are gall-inducing and have a very long thin abdomen. However, O. verrucula also shares with C. viridis : (i) long fleshy setae all over its body; (ii) lacks a hamulohaltere; and (iii) has capitate setae on the antennae restricted to the apical segment. The adult male of C. spiniferus has very few setae but also lacks hamulohalteres and has capitate setae restricted to the apical antennal segment ( Gullan, 1978).

Figures

FIGURE 12. Carpochloroides viridis Cockerell. Adult female, where A = dorsal spinose seta; B = dorsal microtubular duct; C = dorsal loculate pore; D = derm showing reticulate pattern with microtubular ducts; F = view of anal area, K = ventral microtubular duct; L = ventral loculate pore; P = anterior spiracle; S = view of ventral derm showing reticulations, etc; V = antenna, and W = mesothoracic leg.

FIGURE 13. Carpochloroides viridis Cockerell. First-instar nymph, where A = dorsal spinose seta; B = dorsal discoidal pores; K = ventral discoidal pores; L = ventral loculate pores, and W = distal end of metathoracic leg.

FIGURE 14. Carpochloroides viridis Cockerell. Adult male, where C = fleshy seta; D = hair-like seta, and K = distal end of metathotacic leg. For other abbreviations, see under ‘Figure captions’ on p.5.