Carpochloroides eugeniae Miller and Stocks, 2022

Miller, Douglass R. & Stocks, Ian C., 2022, New genera and species of felt scales (Hemiptera: Coccomorpha: Eriococcidae), with descriptions of new species and immature instars of described species, Zootaxa 5221 (1), pp. 1-213 : 48-59

Carpochloroides eugeniae Miller and Stocks sp. n.

Type material: Adult female holotype mounted singly on slide, with right label “ Carpochloroides / eugeniae Miller / & Stocks / HOLOTYPE / UCD;” left label “eugeniae / ad ♀ / On Eugenia / acapulcensis / Chivela, Oaxaca / Mexico / Ferris, 1926 / in eriococcid gall.” Holotype is in UCD . Paratypes: MEXICO: Oaxaca: Chivela , II-(?)-1925, on Eugenia acapulcensis, G.F. Ferris (1 ad. ♀ holotype, 45 ad. ♀♀ paratypes, 65 second-instar ♀♀ paratypes, 9 ad. ♁♁ paratypes, 2 second-instar ♁♁ paratypes, 1 second-instar ♁ molting to third-instar prepupa paratype, 50 first-instar nymph paratypes, 47 first-instar nymph shed-skin paratypes, 1 first-instar nymph molting to secondinstar ♁ paratype on 18 slides) UCD (8 slides), USNM (10 slides). This list does not include the slides that have been misplaced. See the “Notes” section of the “First-instar nymph” description for details.

Etymology: The species epithet “ eugeniae ” is based on the scientific name of the host plant of this species.

Field features: The following information is based on observations of the dry galls taken from the original Ferris collection. The galls are formed in small branches at the points where the leaf petioles are attached. Galls vary from 1 to 2 cm in diameter and are nearly spherical in shape. The inner cavity is also spherical and varies from 5‒9 mm in diameter. Two species occur in these galls and although Carpo. eugeniae is the most abundant, adult females of Carpo. mexicanus have been found in the same galls. Each gall contains many specimens, and most specimens within a single gall are synchronized so that the same gall contains individuals primarily in the same life stage. Different galls, though synchronized internally, can be in quite different life stages compared to other galls. The inner cavity of the gall is lined with a white waxy secretion and the cavity is often filled with small balls of felted wax. Under a dissection microscope the insects can be observed scattered over the wall of the cavity with their stylets inserted. Attempts to locate an exit hole from the gall were unsuccessful. The only holes observed are apparently emergence holes formed by parasitoids. If Carpo. eugeniae and Carpo. mexicanus are different species, it is likely that Carpo. eugeniae is the species that induces the gall, not Carpo. mexicanus as suggested by Ferris (1957).

Adult female ( Fig. 23 )

Description: Holotype, slide mounted, 0.8 mm long, 0.6 mm wide (paratypes 0.6–1.2 mm long, 0.4–0.8 mm wide). Body pyriform, without protruding anal lobes. Anal-lobe area dorsally with 2 setae, 1 long and robust, 1 short and thin, and 1 5-locular pore; ventrally with 2 or 3 flagellate setae including elongate anal-lobe seta, and 1 or 2 5-locular pores.

Dorsum with flagellate setae scattered over entire surface, conspicuously shorter than enlarged setae, longest about 13 μm long. Enlarged setae unusually elongate, of 1 size, in transverse rows over surface, forming 3 pairs of longitudinal lines (medial, mediolateral and lateral), mediolateral lines restricted to thorax and anterior abdomen, each line with 1 seta per segment except on thorax and head, where lateral lines each with 2 or 3 setae per segment. Segment IV with 4 enlarged setae (paratypes with 2–4). Largest seta 98 μm long (paratypes 80–120). Enlarged setae straight or slightly curved, with acute or slightly capitate apices, and thin setal base; some setae bifurcate. Macrotubular ducts with internal sclerosis nearly flat, without conspicuous concavity, abundant over surface, except uncommon on posterior 2 or 3 abdominal segments. Microtubular ducts with narrow cup-shaped internal sclerotization similar to sclerotization of macrotubular ducts, about 6 μm long, total sclerotized area approximately 1/8 th length of unsclerotized area; dermal orifice unsclerotized. Microtubular ducts in small numbers over surface, most abundant along body margin. Multilocular pores scattered over entire surface, of 2 kinds: 7-locular pores; 5- locular pores most abundant. Cruciform pores absent. Microtrichia absent.

Anal ring dorsal, situated near body apex, or apical; ring itself represented only by small, lightly sclerotized plate anterior to anal opening, without setae or pores. Anal tube lightly sclerotized at dermal orifice.Anal flap absent.

Venter with flagellate setae noticeably short, longest seta on segment II about 11 μm long, on segment VII 12 μm long; anal-lobe seta 85 μm long (paratypes 55–88 μm), often apically capitate; setae apically acute. Enlarged setae absent. Macrotubular ducts of same size as on dorsum, scattered over entire surface, least abundant on head. Microtubular ducts restricted to lateral areas, most numerous on thorax, lightly scattered over surface. Multilocular pores of 2 kinds: 7-locular pores rare, absent from some paratypes; 5-locular pores abundant over entire surface, least numerous on head, clustered around spiracles. Legs absent. Antennae each 6-segmented (paratypes 5- or 6- segmented), 65 μm long (paratypes 45–62 μm); apical segment with 6 sensory setae (paratypes with 4–6 sensory setae); segment 1 with or without 1 small flagellate seta, remainder of antenna without setae. Mouthparts unusually large, labium undivided. Eyes, frontal lobes, preantennal pores and microtrichia absent.

Notes: The description is based on 45 specimens from one locality. The adult female of Carpo. eugeniae is most similar to the adult female of Carpo. mexicanus but differs as follows (character states in brackets are those of the female of Carpo. eugeniae ): with short dorsal flagellate setae only (with enlarged setae and short flagellate setae); with small macrotubular ducts but without microtubular ducts (with microtubular and macrotubular ducts); with unsegmented antennae (with 5- or 6-segmented antennae); with large apodemes attached to the clypeus (without large apodemes); and without a plate anterior to the anal opening (with a plate).

Second-instar female ( Fig. 24 )

Description: Slide-mounted specimens, 0.4–0.7 mm long, 0.2–0.5 mm wide. Body oval, without protruding anal lobes. Anal-lobe areas each dorsally with 2 setae, 1 longer and slightly more robust, 1 shorter and thin; each ventrally with 2 or 3 flagellate setae including elongate anal-lobe seta.

Dorsum with flagellate setae most abundant near body margin, noticeably smaller than enlarged setae. Enlarged setae arranged in same pattern as on adult female, not as conspicuously long and robust; largest seta 25–38 μm long. Macrotubular ducts absent. Microtubular ducts difficult to discern, apparently in small numbers over surface, similar to those on adult female except smaller. Multilocular pores usually absent, rarely with 4-locular or 5-locular pores on segments II or III. Cruciform pores absent. Without microtrichia.

Anal ring apical, ring variable, sometimes represented only by small sclerotized area, other times forming complete ring, often with 1 pair of short setae on each side of ring associated with, but not on, anal ring sclerotization. Anal tube orifice sclerotized.

Venter with flagellate setae about same length throughout entire surface, longest seta on segment II 6 μm long, on segment VII 5 μm long; anal-lobe seta 40–60 μm long. Enlarged setae absent. Macrotubular ducts small, present in small numbers on head, thorax, and anterior abdominal segments. Microtubular ducts absent. Multilocular pores of 2 kinds: 5-locular pores in small numbers on head, thorax, and anterior abdominal segments, most numerous near spiracles; 4-locular pores rare or absent. Cruciform pores absent. Legs absent, sometimes with wrinkled area where metathoracic legs would be. Antennae each normally 6-segmented, sometimes with 4 or 5 segments; 25–38 μm long. Eyes ventral. Labium with 2 indistinct segments, basal segment not observed. Frontal lobes absent. Preantennal pore absent. Microtrichia absent.

Notes: The description is based on 65 specimens from one locality. The second-instar female of Carpo. eugeniae is unique in having narrow, elongate enlarged setae, legs absent, anal ring reduced, microtubular ducts similar in appearance to small macrotubular ducts, macrotubular ducts present in small numbers on the venter only.

Adult male ( Fig. 25 )

Description: Slide-mounted specimens, 0.8–1.0 mm long, 0.4–0.5 mm wide. Body slightly pear-shaped, without protruding anal lobes.

Dorsum without lateral tail-forming clusters, multilocular pores, or X-type derm pores. Flagellate setae short, apically acute, about same length as ventral setae, increasing in size posteriorly, some setae with small amount of sclerotization around setal socket, scattered over entire surface, setae on segments VIII and IX unusually abundant. Penial sheath (ps) sclerotized, other abdominal sclerotization variable, normally restricted to lateral areas of segment VIII, sometimes also on segment VII. Thoracic sclerotization absent. Head sclerotization variable, normally with part of ocular sclerite (ocs) visible and with small amount of sclerotization in dorsomedial area. Dorsal eye usually absent, sometimes indicated by small, raised area on ocular sclerite. Rarely dorsal arm of midcranial ridge (dmcr) weakly indicated, dorsal eye (dse) relatively well-developed, ocular sclerite absent.

Penial sheath (ps) 130–180 μm long, width/length 0.8–1.0, style (st) in lateral view slightly curved, apically with several small papillae. Sheath heavily sclerotized on both surfaces, with weak indication of division and with unusually large number of flagellate setae. Basal rod absent.

Venter with flagellate setae present medially on head, thorax, and segments II to VII, also present submedially and laterally on abdomen; 1 or 2 setae on each metepisternum (mtes), rarely with 1 on each mesepisternum (mses). Abdominal sclerotization present on lateral areas of segments VII and VIII. Thoracic sclerotization limited to small areas associated with pro- (pa) and mesothoracic (ma) apophyses and to sclerotization associated with pleural ridges (pr) of each leg. Head sclerotization restricted to ventromedial area near anterior apex of head and to ventral portion of ocular sclerite (ocs). Mouth structure variable, unusually complex for adult male; normally rudiments of clypeus (cl) and labium (lb) present, often possessing setae apically, pharynx (px) present. Cranial apophysis (ca) bifurcate. Ventral and lateral ocelli normally absent, rarely indicated by small, raised areas on ocular sclerite. Proand mesothoracic legs approximately same size, metathoracic legs longest. Fleshy setae absent from legs except slightly enlarged on inner margin of tibia. Tarsal digitules absent, claw digitules not reaching tip of claw, apically acute. Antennae each 5- or 6-segmented, third segment longest, apical segment rounded; fleshy setae present on apical 2 segments, capitate setae absent.

Notes: The description is based on 9 specimens from one locality. The adult male of Carpo. eugeniae is quite variable, particularly in regard to abdominal sclerotization, ocular sclerite development, eye development, mouth structure, setal patterns, and antennal segmentation. It is similar to the adult males of Pseudochermes fraxini (Kaltenbach) and Ovaticoccus adoxus (Ferris) in having little or no head and thoracic sclerotization,unusual antennae,no wings or hamulohalteres, and no tail-forming pore clusters. Pseudochermes fraxini differs as follows (character states of Carpo. eugeniae are in brackets): antennae each 8-segmented (5- or 6-segmented); claw digitules reaching tip of claw (not reaching tip); penial sheath same length as, or shorter than, other abdominal segments (longer). Ovaticoccus adoxus differs from C. eugeniae as follows: antennae each 9-segmented (5- or 6-segmented); multilocular pores present (absent); fleshy setae on distal 6 segments of antenna (on distal 2 segments); fleshy setae on tibiae (absent from tibiae).

Fourth-instar male (pupa) ( Fig. 26 )

Description: Slide-mounted specimen 0.8 mm long, 0.4 mm wide. Body elongate oval, without protruding anal lobes.

Dorsum with flagellate setae of 2 kinds: slightly enlarged setae restricted to lateral margins of segment VIII; thin flagellate setae scattered over surface, basically of same pattern as on second-instar male, increasing in length posteriorly. Without tubular ducts, multilocular pores, or sclerotization.

Penial sheath unsclerotized, without lobular development. Setae present on dorsum only. Anal opening dorsal, with anal-tube orifice weakly sclerotized. Genital opening slightly developed ventrally near sheath apex.

Venter with multilocular pores and tubular ducts absent. Body setae of same distribution as on adult male, noticeably shorter than on dorsum, anal-lobe seta 23 μm long. Mouth opening and cranial apophysis present. Legs and antennae without indication of setae. Antennae each 4- or 5-segmented.

Notes: The description is based on one specimen from one locality. The pupa of Carpo. eugeniae is unique among the few eriococcid pupae that have been described, in lacking multilocular pores.

Third-instar male (prepupa) ( Fig. 27 )

Description: Slide-mounted specimen 0.6 mm long, 0.4 mm wide. Body elongate oval, without projecting anal lobes.

Dorsum with flagellate setae of 2 kinds: slightly enlarged setae restricted to lateral margins of segment VIII; thin flagellate setae scattered over surface, basically in same pattern as on second-instar male, increasing in length posteriorly. Without tubular ducts, multilocular pores, or sclerotization.

Penial sheath unsclerotized, without lobular development. Setae present on venter only. Anal opening dorsal, with anal tube orifice weakly sclerotized, located near posterior apex. Genital opening slightly developed ventrally near sheath apex.

Venter with multilocular pores and tubular ducts absent. Body setae of same distribution as on adult male, noticeably shorter than on dorsum, anal-lobe seta 20 μm long. Mouth opening, pharynx, and cranial apophysis present. Legs and antennae without indication of setae. Antennae each 4- or 5-segmented.

Notes: The description is based on one specimen from one locality. The prepupa of Carpo. eugeniae is unique among the few eriococcid prepupae that have been described in lacking multilocular pores.

Second-instar male ( Fig. 28 )

Description: Slide-mounted specimens, 0.4–0.5 mm long, 0.2–0.3 mm wide. Body oval, without protruding anal lobes. Anal-lobe areas each dorsally with 2 setae, 1 longer; each ventrally with 3 flagellate setae including elongate anal-lobe seta.

Dorsum with flagellate setae most abundant near body margin, noticeably smaller than enlarged setae. Enlarged setae arranged in same pattern as on adult female, not as conspicuously long and robust; largest seta 20–34 μm long, setae rarely capitate. Macrotubular ducts absent. Microtubular ducts difficult to discern, apparently in small numbers over surface, similar to those on adult female except smaller. Multilocular pores usually absent, rarely with 2 or 3 5-locular pores on thorax. Cruciform pores absent. Without microtrichia.

Anal ring dorsal or apical, ring abortive, normally partially developed around anal tube opening, often with 1 or 2 short setae on each side of ring associated with, but not on, anal ring. Anal tube heavily sclerotized near dorsal surface.

Venter with flagellate setae about same length throughout surface, longest seta on segment II 5 μm long, on segment VII 6 μm long; anal-lobe seta 35–63 μm long. Enlarged setae absent. Macrotubular ducts small, difficult to see, present in small numbers on head, thorax, and anterior abdominal segments. Microtubular ducts absent, or present on posterior part of head and thorax. Multilocular pores of 2 kinds: 5-locular pores in small numbers, always present near spiracles, sometimes present near legs, on lateral margins of 1 or 2 abdominal segments, and near mouthparts; 3-locular pores rare or absent. Cruciform pores absent. Legs present, each femur with 3 setae, all distal; tibiae each with 4 setae; hind tibia/tarsus 0.7–0.9; tarsal and claw digitules apically acute; claws with small denticle near tip. Antennae each usually 6-segmented, 1 specimen with 5 segments; each about 38 μm long; apical segment with 4 sensory setae and 1 or 2 flagellate setae; segment 5 with 1 sensory seta and 1 flagellate seta; segment 4 without setae; segments 2 and 1 each with 1 seta.

Notes: The description is based on six specimens from one locality. The second-instar male of Carpo. eugeniae is unique in having: narrow, elongate enlarged setae; anal ring reduced; microtubular ducts similar in appearance to small macrotubular ducts; macrotubular ducts in small numbers on the venter only.

It is interesting that the second-instar male retains its legs, whereas the second-instar female lacks them; we have discovered a similar situation in the second-instar males of Xerococcus fouquieriae . We have been unable to find any other species of scale insect that shows this kind of sexual dimorphism in the second-instar nymphs.

First-instar nymph ( Fig. 29 )

Description: Slide-mounted specimens 0.2–0.3 mm long, 0.1–0.2 mm wide. Body oval, without protruding anal lobes. Anal-lobe areas dorsally each with 2 short conical setae and 1 longer seta; ventrally each with 2 or 3 elongate setae including anal-lobe seta.

Dorsum without flagellate setae. Enlarged setae not conspicuously enlarged, forming 3 pairs of longitudinal lines on head and thorax (medial, mediolateral and lateral), without mediolateral line on abdomen; enlarged setae short and slender, straight sides, thin setal bases, longest setae about 6 μm long. Tubular ducts intermediate between macro- and microtubular ducts, difficult to discern, normally in small numbers, sometimes absent. Multilocular and cruciform pores absent.

Anal ring bent over abdominal apex, dorsal portion composed of large, sclerotized plate with 2 short setae on each side of ring but not associated with anal opening and small opening for anal tube; ventral portion extension of dorsal plate with 1 pair of conical setae.

Venter with setae flagellate, longest seta on segments II and VII about 7 μm long; anal-lobe seta unusually long, 190–250 μm long. Enlarged setae absent. Tubular ducts of same kind as on dorsum, present on thorax, rarely on head. Multilocular pores with 5 loculi only, with 1 or 2 in atria of each spiracle. Cruciform pores absent. Legs with each femur with 2 or 3 setae, distal only; each tibia with 4 setae; tibia/tarsus 0.9–1.1. Antennae each 6-segmented, about 68 μm long. Frontal lobes, preantennal pores and microtrichia absent.

Notes: The description is based on numerous specimens from one locality. The first-instar nymph of Carpo. eugeniae is similar to the first-instar nymph of C. viridis in having multilocular pores in the atria of the spiracles. The two species differ by the following (character states in brackets are of C. eugeniae ): with unusually elongate and enlarged dorsal setae, longer than width of each abdominal segment (enlarged setae nearly flagellate, not enlarged, shorter than width of each abdominal segment); without a sclerotized plate associated with anal ring (with large, sclerotized plate associated with anal ring).

The above specimens were sent by Gordon Ferris Stanford University, to Harold Morrison, US Department of Agriculture, in 1950 for the latter’s opinion; they were determined as “Eriococcinae” and “ Eriococcidae genus uncertain.” The shipment must have included galls from which specimens were mounted by Morrison, but the slide labels do not look like USDA labels or Stanford University labels. We mounted more specimens from the dry collection and it included 247 specimens of various instars on 51 slides. Unfortunately, most of these specimens have been misplaced (six of the slides have been rediscovered and are in the USNM), but they served as the basis of the extensive descriptions and illustrations presented above. Unfortunately, the prepupa and pupa specimens have not been recovered, but specimens of all other instars are included in the “Type material” section. We decided to include all of the information gleaned from the missing and found specimens because it documents an interesting and unique life history. Hopefully additional collections will be made in the future and the misplaced specimens will be found. When they are found, they are to be considered paratypes since they were used for the original description and are from the same collection as the type specimens listed above.

Figures

FIGURE 23. Carpochloroides eugeniae sp. n., adult female, Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris. A=enlarged seta; C=7-locular pore; D=5-locular pore; F=microtubular duct; G=macrotubular duct; I=anal ring; J=antenna.

FIGURE 24. Carpochloroides eugeniae sp. n., second-instar female, Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris. A=enlarged seta; D=5-locular pore; F=microtubular duct; G=macrotubular duct; I=anal ring; J=antenna.

FIGURE 25. Carpochloroides eugeniae sp. n., adult male (apterous), Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris. See Materials and methods for abbreviations.

FIGURE 26. Carpochloroides eugeniae sp. n., fourth-instar male (pupa), Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris. I=anal ring.

FIGURE 27. Carpochloroides eugeniae sp. n., third-instar male (prepupa), Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris. I=anal ring.

FIGURE 28. Carpochloroides eugeniae sp. n., second-instar male, Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris.A=slightly enlarged seta; D=5-locular pore; E=3-locular pore; F=microtubular duct; G=macrotubular duct; I=anal ring; L=claw.

FIGURE 29. Carpochloroides eugeniae sp. n., first-instar nymph, Chivela, Oaxaca, Mexico, February?, 1925, on Eugenia acapulcensis, G.F. Ferris. A=enlarged seta; D=5-locular pore; F=microtubular duct; L=claw; N=flagellate seta.