Ovaticoccus agavium (Douglas)

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 : 96-106

Ovaticoccus agavium (Douglas)

Coccus agavium Douglas 1888: 150 .

Gymnococcus agavium (Douglas) ; Cockerell 1894b: 1053 (change of combination).

Ripersia agavium (Douglas) ; Newstead 1897: 12-13. (change of combination).

Pseudantonina agaves Chiaromonte 1929: 61-62 (junior synonym).

Ovaticoccus agavium (Douglas) ; Kloet 1944: 86 (change of combination).

Specimens examined: UNITED STATES: California: Los Angeles Co.: Los Angeles, V-28-1940, on Aloe sp. , F.R. Platt (10 ad. ♀♀ on 3 slides) UCD; Pasadena, V-12-1938, on Yucca sp. , Marsh (8 ad. ♀♀ on 4 slides) UCD. Orange Co.: Stanton, VI-7-1966, on Yucca sp. , R. Wyatt (5 ad. ♀♀ on 5 slides) CDA. San Bernardino Co.: Yucca Valley , XI-16-1967, on Agave sp. , Cambin (2 ad. ♀♀, 1 ad. ♁ on 3 slides) UCD; San Diego Co.: Torrey Pines, IV-6-1965, on Agave sp. , J.R. Carlin (1 ad. ♀) UCD. Massachusetts: Suffolk Co.: Cambridge, Harvard Botanical Gardens, VII-15-1921, on Agave sp. , and A. scolymus (= A. potatorum ), H. and E. Morrison (6 ad. ♀♀, 3 second-instar ♀♀, 8 first-instar nymphs on 5 slides) USNM. Missouri: Independent city: Saint Louis, Missouri Botanical Garden, VII-25-1923, on Agave sp. , H.L. Sanford (7 ad. ♀♀, 3 second-instar ♀♀, 3 first-instar nymphs, 6 ad. ♁♁, 1 third-instar ♁, 5 second-instar ♁♁ on 8 slides) USNM. New York: Bronx Co.: Bronx, VIII-31-1944, on Agave sp. , G. Rau (4 ad. ♀♀ on 1 slide) USNM; New York Co.: New York, New York Botanical Gardens, VII-1-1921, on A. neglecta and A. antillarum (?), H.L. Sanford and L.C. Griffith (17 ad. ♀♀, 5 second-instar ♀♀, 6 first-instar nymphs, 1 ad. ♁, 2 fourth-instar ♁♁ on 10 slides) USNM; New York Botanical Garden, III-29-1937, on Agave sp. , G. Rau (2 ad. ♀♀ on 2 slides) UCD, USNM. MEXICO: State (?): XI-1-1952, on Agave sp. , W.B. Wood (1 second-instar ♀, 3 second-instar ♁♁ on 1 slide) USNM; V-4-1960, on cactus plants, J. Hidalgo Jr., (1 ad. ♀ on 1 slide) USNM; XII-22-1967, on Agave sp. , C.H. Spitzer (2 ad. ♀♀ on 1 slide) USNM. Puebla: Zacatepec, VII-16-1967, on Agave sp. , D.R. Miller and J. Villanueva B. (3 ad. ♀♀ on 3 slides) UCD. Vera Cruz: intercepted at New York, VII-21-1927, on A. atrovirens, H.Y. Gouldman (3 ad. ♀♀ on 1 slide) USNM. SPAIN: Intercepted at New York, X-21-1959, on Agave sp. , J. Hidalgo Jr., (1 ad. ♀ on 1 slide) USNM. UNITED KINGDOM: England: Kew Gardens, VI-22-1956, VI-27-1957, and V-26-1964, on Agave sp. , A. decipens , and A. franzosini, S.A. Afifi and K. Boratyński (29 ad. ♀♀, 29 second-instar ♀♀, 40 first-instar nymphs, 8 ad. ♁♁, 4 fourth-instar ♁♁, 5 third-instar ♁♁, 21 second-instar ♁♁ on 16 slides) NHM.

The adult female of this species was described in detail by Boratyński (1958), Miller and McKenzie (1967) and Miller and Miller (1993), and information from those papers is not repeated here. Boratyński also briefly discussed the morphological character states of the first-instar nymph, second-instar female and adult male; he included a drawing and description of a second-instar male (labeled as a third-instar male). Miller and McKenzie (1967) included descriptions of the first-instar nymph, second-instar female (labeled as a second- or third-instar female) and second-instar male (labeled as a third- or fourth-instar male), but did not include illustrations. Afifi (1968) and Hodgson (2020) described and illustrated the adult male but our illustration and description shows slightly different character states, particularly the sensory setae on the tibiae.

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

Second-instar female ( Fig. 48 )

Description: Slide-mounted specimens 0.7–1.0 mm long, 0.3–0.5 mm wide. Body elongate oval, without protruding anal lobes. Anal-lobe areas each dorsally with 2 flagellate setae and 0 or 1 5-locular pore; each ventrally with 3 or 4 flagellate setae including suranal seta and elongate anal-lobe seta, and 0 or 1 5-locular pore.

Dorsum with single longitudinal line of flagellate setae along lateral margin, rarely with 1 or 2 such setae on sublateral areas of posterior abdominal segments. Enlarged setae of 2 sizes: larger size present in 3 pairs of longitudinal lines (medial, mediolateral and lateral); smaller size present in 2 pairs of longitudinal lines, sometimes some or all of small-sized setae replaced by flagellate setae; largest seta 9–12 μm long; enlarged setae variable in shape from thin and conical with straight lateral margins to broad and pear-shaped with curved lateral margins; setal base thin; segment IV with 7–10 setae including 6–10 enlarged setae and 0–4 flagellate setae; segment IV with combined total of 6–10 enlarged setae dorsally and ventrally. Macrotubular ducts absent. Microtubular ducts normally absent, rarely with 1 or 2 on abdomen. Multilocular pores of 1 or 2 kinds: 5-locular pores most numerous, scattered over entire surface, rarely with more pores on head; 3-locular pores rarely present. Cruciform pores absent. Microtrichia absent.

Anal ring ventral, semicircular, incomplete, non-cellular, with 3 setae on each side of ring, each normally shorter than diameter of ring; additional pair of setae often associated with, but not attached to, ring; anal tube weakly sclerotized, dermal orifice sclerotized; without anal flap.

Venter with setae flagellate, longest seta on segment II 15–25 μm long, on segment VII 12–30 μm long; anallobe seta approximately 85μm long. Enlarged setae absent. Macrotubular ducts and microtubular ducts absent. Multilocular pores of 2 kinds: 5-locular pores most abundant, present over entire surface, least abundant on head; 3-locular pores rare, normally on thorax and head. Cruciform pores variable, in clusters of 1–4 pores, on lateral margins of segments V or VI to VII or VIII, also in medial areas from segment II or III to VII, rarely with 1 or 2 pores on head and thorax. Legs with small number of translucent pores on hind coxa; each femur with 3 setae, without proximal seta; each tibia with 4 setae, without middle seta; hind tibia/tarsus 0.7–0.8. Antennae each 6- segmented, 140–150 μm long. Frontal lobes absent. Preantennal pore present. Microtrichia present on prothorax to segment VIII and on ventral surface of hind 2 pairs of coxae.

Notes: The description is based on 41 specimens from five localities. The second-instar female of Ov. agavium is most similar to the second-instar female of Ov. exoticus in lacking dorsal cruciform pores and in having dorsal multilocular pores and similarly shaped anal rings. They differ as follows (character states in brackets are those of Ov. agavium ): each femur with four setae including a proximal seta (with three setae, without a proximal seta); without ventromedial cruciform pores (normally with ventromedial cruciform pores).

Adult male (macropterous) ( Fig. 49 )

Description: Slide-mounted specimens 1.0– 1.2 mm long, 0.3–0.4 mm wide. Body elongate, segment VIII somewhat produced laterally, giving lobular appearance.

Dorsum with 1 pair of tail-forming pore clusters (tfpc); each cluster with 2 elongate, capitate setae of 2 distinct sizes: longest seta 120–145 μm long; shortest setae 85–115 μm long; with 35–50 multilocular pores. Multilocular pores restricted to tail-forming pore clusters, each with 3, 5, or 7 loculi, 5-locular pores most common. X-type pores (x) normally present near dorsal arm of midcranial ridge (dmcr), rarely absent. Flagellate setae slender, apically acute, approximately same length as those on venter, in clusters of 2 or 3 near each margin of segments II to VIII, also in transverse rows on segments II to VII, with 1 pair of elongate setae in medial area of segment VIII; setae present on metathorax and mesothorax, absent from prothorax; head setae present posterior to postoccipital ridge (por) and along dorsal arm of midcranial ridge. Abdominal sclerotization variable, median tergal plate VIII (mtp) clearly indicated, sometimes present on segments II, V, VI and VII, nearly always present around marginal clusters of setae on segments VI and VII. Metapostnotal ridge (mpr) normally present. Scutellar ridge (sclr) strongly developed. Scutum (sct) sclerotized throughout. Prescutum (prsc) rectangular. Prescutal suture (pscs) weakly developed. Pronotal ridges (prnr) normally strongly developed; pronotal sclerites not observed. Hamulohalteres absent. Mesothoracic wings each with 2 setae near wing base. Postoccipital ridge (por) large, lateral ends normally bifurcate. Dorsal arm of midcranial ridge thin, usually nearly touching postoccipital ridge posteriorly, rarely ending far from postoccipital ridge; connected to lateral and ventral arms anteriorly. Dorsomedial sclerite (dmep) lightly sclerotized. Dorsal eye (dse) approximately 28 μm in diameter. Lateral ocellus (lo) approximately 15 μm in diameter, attached to postocular ridge by small arm. Ocular sclerite (ocs) strongly sclerotized dorsally.

Penial sheath (ps) 130–158 μm long, elongate, width/length 0.63–0.67, style (st) in lateral view straight, apically with several small papillae. Sheath with weak indication of longitudinal division ventrally. Dorsal surface heavily sclerotized; ventral surface with large membranous area as well as ventral slit in medial areas, sclerotized anteriorly and laterally. Basal rod absent.

Venter with flagellate setae present medially, mediolaterally, and laterally on segments II or IV to VIII, with 1 or 2 setae on segment II; thoracic setae, when present, restricted to single pair near anterior legs; head setae present only near anterior portion of ventral arm of midcranial ridge (vmcr). Abdominal sclerotization restricted to posterior 3 or 4 pregenital segments; segment VIII with conspicuous plate in submedial area and with 1 pair of small lateral plates; segments V to VII with light sclerotization in medial or submedial areas. Metathoracic precoxal ridge (pcr 3) weakly developed or absent. Lateropleurites (lpl) on mesothorax continuous. Mesosternum (stn 2) well-developed with large furca (fr). Prosternum (stm 1) varying from short to thin and triangular to rectangular. Mouth tubercle (mt) present. Preoral ridge (pror) heavily sclerotized, not touching postocular ridge. Cranial apophysis (ca) bifurcate. Ocular sclerite (ocs) unsclerotized except immediately surrounding ventral eye (ve). Ventral eye approximately same size as dorsal eye. Prothoracic and mesothoracic legs approximately equal in size, metathoracic legs longest. Trochanters each with 3 campaniform sensilla on each surface. Fleshy setae (fs) on distal portion of tibia, absent from tarsus. Inner setae on apical margin of tibia and inner margin of second tarsal segment enlarged; claw (cla) with small denticle (de). Antennae each 10-segmented, third segment longest, 1.8–2.1 times longer than apical segment, tenth segment apically pointed. Fleshy setae present on segments 3–10; capitate setae (cs) on segments 5 or 6–10.

Notes: The description is based on 16 specimens from four localities. Adult males are known for three species of Ovaticoccus : Ov. adoxus , Ov. agavium , and Ov. salviae . The adult male of Ov. adoxus is apterous and quite different from the other two; in fact, Hodgson (2020) suggested that the campaniform sensilla on the trochanter show more similarity to eriococcids in the Gondwanan lineage than those from North America. The adult males of Ov. agavium and Ov. salviae are similar to the adult male of Ov. agavium in lacking hamulohalteres, and in having 10-segmented antennae with fleshy setae and capitate setae, and similar genital capsules. They differ as follows (character states in brackets are of Ov. agavium ): with two small unsclerotized circular areas on scutum next to the scutellum (scutum without unsclerotized areas); wing without hamulohaltere pocket (with pocket); lateral ocellus reduced or absent (present and obvious).

Fourth-instar male (pupa) ( Fig. 50 )

Description: Slide-mounted specimens 1.1–1.2 mm long, 0.4 mm wide. Body elongate, small lobes present on segment VIII.

Dorsum with flagellate setae of approximately same pattern as adult male; often with 1 lateral seta on each of posterior 4 or 5 abdominal segments, enlarged apically. Multilocular pores restricted to lateral margins of abdomen, with 3–13 loculi. Discoidal pores absent. Lobe on lateral margin of segment VIII sclerotized. Hamulohalteres absent. Front wing buds approximately 410 μm long, partially sclerotized. Ocular sclerite lightly sclerotized on dorsal and lateral areas only.

Penial sheath sclerotized, dorsally with 1 small lobe on each side of body, each with 2 elongate setae. Anal opening dorsal, without sclerotized rim.

Venter with multilocular pores of same kinds as on dorsum, present along lateral margins of segments I or II to VII or VIII, in small numbers on medial and mediolateral areas of anterior 4 or 5 segments, also near both pairs of spiracles. Discoidal pores absent. Flagellate setae of same distribution as adult male, slightly longer than dorsal setae, usually acute; longest lateral seta on segment VIII 88–98 μm long. Mouth tubercle present. Legs partially developed, setae indicated only by clear dots. Antennae each 10-segmented, about 200 μm long. Microtrichia on prothorax to segment VIII, and on venter of mid- and hind coxae. Eye represented by sclerotized band.

Notes: The description is based on 6 specimens from two localities. The macropterous pupa of Hy. hyperici described in this work is similar to the macropterous pupa of Ov. agavium in having reduced appendages, reduced or no mouthparts, and the presence of multilocular pores and flagellate setae. They differ as follows (character states in brackets are of Ov. agavium ):in lacking dorsal multilocular pores (pores present on margin).

Third-instar male (prepupa) ( Fig. 51 )

Description: Slide-mounted specimen about 1.0 mm long, 0.3 mm wide, not including wing buds. Small lateral lobes present on segment VIII.

Dorsum with flagellate setae of same pattern as adult male, of 2 types: some setae on medial, mediolateral and lateral areas of abdomen more robust than remaining setae, with apex slightly swollen, other setae apically acute. Multilocular pores either entirely absent or restricted to lateral margins of abdomen, rarely with 1 pore on thorax. Entire dorsum unsclerotized. Front wing buds approximately 19 μm long. Ocular sclerite absent.

Abdominal segment IX small, normally not protruding, unsclerotized except abortive ring near anal opening. Anal opening ventral. Small lobes sometimes visible.

Venter with multilocular pores with 9–13 loculi, scattered over surface except in medial areas of posterior abdominal segments. Flagellate setae apically acute, slightly longer than those on dorsum; longest lateral setae on segment VIII 75–89 μm long. Legs and antennae small and poorly defined, partially sclerotized. Eye absent.

Notes: The description is based on six specimens from two localities. No other prepupae of Ovaticoccus species have been described for comparison except those treated in this paper. The macropterous prepupa of Hy. hyperici is similar to the macropterous prepupa of Ov. agavium in having reduced appendages, reduced or no mouthparts, and in the presence of multilocular pores and flagellate setae. They differ as follows (character states in brackets are of Ov. agavium ): dorsal multilocular pores present on posterior abdomen (absent); without a lobe on margin of segment VIII (present).

Second-instar male ( Fig. 52 )

Description: Slide-mounted specimens 0.8–1.0 mm long, 0.4 mm wide. Body elongate oval, without protruding anal lobes. Anal-lobe areas each dorsally with 2 flagellate setae and 0 or 1 5-locular pore; each ventrally with 2 or 3 flagellate setae including suranal seta and elongate anal-lobe seta, 0 or 1 tubular duct and 0 or 1 multilocular pore.

Dorsum with flagellate setae of 2 kinds: largest setae with slightly expanded apices, arranged in 3 pairs of longitudinal lines (medial, mediolateral and lateral), rarely absent; smaller setae present over entire surface, apically acute. Enlarged setae normally absent, rarely with 1 or 2 on abdomen; segment IV with 11 or 12 flagellate setae, without enlarged setae. Macrotubular ducts present over entire surface. Microtubular ducts normally absent, rarely with 1 or 2 on abdomen. Multilocular pores normally restricted to last 3 or 4 abdominal segments near body margin, rarely with few pores in medial areas of anterior abdominal segments and thorax, of 4 kinds: 7-locular pores, 4- locular pores, and 3-locular pores in equal numbers; 5-loculars most abundant. Cruciform pores and microtrichia absent.

Anal ring ventral, semicircular, incomplete, non-cellular, with 3 setae on each side of ring, each normally shorter than diameter of ring; additional pair of setae often associated with, but not attached to, ring; anal tube weakly sclerotized, with dermal orifice sclerotized or unsclerotized; without anal flap.

Venter with flagellate setae on segment II 35–42 μm long, on segment VII 38–43 μm long; anal-lobe seta 95– 115 μm long. Enlarged setae absent. Macrotubular ducts on mediolateral and lateral areas of thorax and abdomen, uncommon on head. Microtubular ducts absent. Multilocular pores scattered over entire surface. Cruciform pores variable, normally restricted to lateral margins of segments V to VII, each segment margin with 1 pore; rarely pores more abundant. Legs with hind coxae without pores; each femur with 3 setae, without proximal seta; each tibia with 4 setae, without middle seta; hind tibia/tarsus 0.8–0.9. Antennae each 7-segmented, 175–183 μm long. Frontal lobes absent. Preantennal pore present. Microtrichia present on ventral surface of hind 2 pairs of coxae.

Notes: The description is based on 29 specimens from three localities. The second-instar male of Ov. agavium is most similar to the second-instar male of Ov. exoticus in having few or no enlarged setae, lacking dorsal cruciform pores and in having similarly shaped anal rings. They differ as follows (character states in brackets are those of Ov. agavium ): each femur with four setae including a proximal seta (with three setae, without a proximal seta); without ventromedial cruciform pores (normally with ventromedial cruciform pores).

First-instar nymph ( Fig. 53 )

Description: Slide-mounted specimens 0.5–0.6 mm long, 0.2–0.3 mm wide. Body elongate, without protruding anal lobes. Anal-lobe areas each dorsally with 2 flagellate setae; each ventrally with 2 flagellate setae including suranal and anal-lobe seta.

Dorsum with flagellate setae in single longitudinal line along body margin. Enlarged setae of 1 size in 3 pairs of longitudinal lines (medial, mediolateral and lateral), in transverse rows from head to segment VII; largest seta 6–9 μm long; enlarged setae pear-shaped with truncate apex; setal base thin; segment IV with 6–8 setae including 6 enlarged setae and 2 flagellate setae; segment IV with combined total of 6–8 enlarged setae on dorsum and venter. Macrotubular ducts, microtubular ducts, multilocular pores, and cruciform pores absent. Microtrichia absent.

Anal ring ventral, semicircular, incomplete, divided anteriorly and posteriorly, non-cellular, with 3 setae on each side of ring, each normally shorter than diameter of ring; additional pair of setae often associated with, but not attached to, ring; anal tube with dermal orifice sclerotized or unsclerotized; anal orifice unsclerotized; without anal flap.

Venter with flagellate setae, longest seta on segment II approximately 16 μm long, on segment VII 20–27 μm long; longest anal-lobe seta approximately 85 μm long. Enlarged setae absent. Macrotubular ducts and microtubular ducts absent. Multilocular pores fairly consistent; head normally with 5-locular pores, rarely 3-locular pores; spiracles each with 5-locular pores; mesothoracic and metathoracic legs each with 3-locular pores; segments IV and V each with 5-locular pores. Cruciform pores present in sublateral areas of segments V to VIII. Legs without pores; each femur with 3 setae, without proximal seta; each tibia with 4 setae, without middle seta; hind tibia/tarsus 0.7–0.9. Antenna 6-segmented, 105–115 μm long, Frontal lobes absent. Preantennal pore present. Microtrichia on metathorax to segment VIII, not visible on legs.

Notes: The description is based on 57 specimens from four localities. The first-instar nymph of Ov. agavium is most similar to the first-instar nymph of Ov. exoticus in having the dorsomedial longitudinal line of enlarged setae present at least on the abdomen, with similarly shaped anal rings, and usually with ventrolateral cruciform pores on the abdomen. They differ as follows (characters in brackets are those of Ov. agavium ): each femur with four setae including a proximal seta (with three setae, without a proximal seta).

Figures

FIGURE 48. Ovaticoccus agavium (Douglas 1888), second-instar female. Mexico, State (?): November 21, 1952, on Agave sp., W.B. Wood. A=enlarged seta; D=5-locular pore; E=3-locular pore; F=microtubular duct; H=cruciform pore; I=anal ring; L=claw; Q=dorsal surface of coxa.

FIGURE 49. Ovaticoccus agavium (Douglas 1888), adult male, Kew Gardens, England, United Kingdom, June 22, 1956, on Agave decipens, S.A. Afifi and K. Boratyński. See methods section for explanation of abbreviations.

FIGURE 50. Ovaticoccus agavium (Douglas 1888), fourth-instar male (pupa). New York Botanical Garden, New York City, New York Co., New York, USA, July 1, 1921, on Agave neglecta, H.L. Stanford and L.C. Griffith. C=7-locular pore; E=3- locular pore; N=flagellate seta; S=6-locular pore; T=pore with>7 loculi.

FIGURE 51. Ovaticoccus agavium (Douglas 1888), third-instar male (prepupa). Cactus House, Missouri Botanical Garden, St. Louis, Independent City, Missouri, USA, July 25, 1923, on Agave sp., H.L. Stanford. B=slightly enlarged seta; I=anal ring; T=pore with>7 loculi.

FIGURE 52. Ovaticoccus agavium (Douglas 1888), second-instar male. Cactus House, Missouri Botanical Garden, St. Louis, Independent City, Missouri, USA, July 25, 1923, on Agave sp., H.L. Stanford. B=slightly enlarged seta; C=7-locular pore; D=5-locular pore; E=3-locular pore; F=microtubular duct; G=macrotubular duct; H=cruciform pore; I=anal ring; L=claw; N=flagellate seta.

FIGURE 53. Ovaticoccus agavium (Douglas 1888), first-instar nymph. New York Botanical Garden, New York City, New York Co., New York, USA, July 1, 1921, on Agave neglecta, H.L. Stanford and L.C. Griffith. A=enlarged seta; D=5-locular pore; E=3-locular pore; H=cruciform pore; I=anal ring; L=claw; N=flagellate seta.