Barbenigma biza Powell & Miller, 2024

Powell, Erin C., Miller, Douglass R. & Keller, Oliver, 2024, Barbenigma Powell & Miller, a bizarre new genus and two new species of scale insects (Hemiptera: Coccomorpha: Ortheziidae) from the United States, Zootaxa 5529 (2), pp. 293-317 : 298-307

publication ID

https://doi.org/10.11646/zootaxa.5529.2.4

publication LSID

lsid:zoobank.org:pub:0EBB38A4-0B20-419B-8DB2-AD43963D6B18

DOI

https://doi.org/10.5281/zenodo.14022389

persistent identifier

https://treatment.plazi.org/id/036887BC-8B28-8F00-FF50-FF1BFCEFE9B7

treatment provided by

Plazi

scientific name

Barbenigma biza Powell & Miller
status

sp. nov.

Barbenigma biza Powell & Miller , sp. nov.

Material examined

Holotype: adult ♀, mounted singly. FLORIDA: Alachua Co., Gainesville, 23 III 1999, on Rumex hastatulus ( Polygonaceae ), coll. R. Esser, E99-879, FSCA_00073114 ( FSCA).

Paratypes: FLORIDA: Alachua Co., Gainesville, Paynes Prairie Preserve , 23 III 1999, on Rumex hastatulus , coll. R. Esser and C. Zamora, E99-879 (1 ad. ♀ and 1 first-instar? nymph (FSCA_00073115) on 2 slides) ( FSCA); Brevard , Co., Merritt Island , 4 VII 1988, on Stenotaphrum secundatum ( Poaceae ), coll. S. Walker (5 ad. ♀♀ (FSCA_00073116– FSCA _00073120), 1 second-instar? nymph ( FSCA _00073121) on 6 slides) ( FSCA, USNM); Palm Beach Co., West Palm Beach, 15 XI 1965, on Stenotaphrum secundatum , coll. R.S. Mullin, N. Bezona, 124942 (2 ad. ♀ (FSCA_00073122–FSCA_00073123 on 2 slides) ( FSCA) .

Other material examined: FLORIDA: Broward Co., Lighthouse Point, 17 VI 1974, on Stenotaphrum secundatum , coll. F. Donaldson (1 second-instar? nymph ( FSCA _00073124) and 1 third-instar? nymph (FSCA_ 00073125) on 2 slides) ( FSCA); Orange Co., Maitland , 5 XI 1969, in soil (lawn), coll. H.M. VanPelt, DPI# 125664 (3 second-instar? nymphs (FSCA_00073126–FSCA_00073128) on 3 slides) ( FSCA); Marion Co., Belleview , 15 IX 1966, in soil, coll. H. Myers (1 second-instar? nymph (FSCA_00073129) on 1 slide) ( FSCA); Miami-Dade Co., Miami, 14 X 1971, on Cocos nucifera ( Arecaceae ), coll. R. Brown, B-7925 (1 second-instar? nymph (FSCA _ 00073130) on 1 slide) ( FSCA).

Etymology: The epithet “ biza ” is an abbreviation of the English word “bizarre” and is to be treated as a noun in apposition.

Adult female ( Figs 1–4 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 )

Description: Slide-mounted holotype 1.25 mm long, 1.1 mm wide; paratypes (n=8) 0.64−1.26 mm long, 0.23−0.95 mm wide; body rotundly oval in mature female ( Figs 1–2 View FIGURE 1 View FIGURE 2 ), narrow elongate in young females ( Fig. 3 View FIGURE 3 ).

Dorsum with robust fimbriate setae ( Figs 1c View FIGURE 1 , 3m View FIGURE 3 ), each with up to five tines, sparse, two present marginally on each side of each abdominal segment, one present submedially on each abdominal segment, roughly in single longitudinal line on thorax and head, holotype with one instance of two setae arising from single socket on head, longest seta on submedial area of abdomen 19 µm long (paratypes 18–23 µm), longest seta on margin 22 µm long (paratypes 16–30 µm), longest seta on head 22 µm long (paratypes 19–24 µm). Dorsal spines each without setal base ( Figs 1o View FIGURE 1 , 3n View FIGURE 3 ), with swollen, rounded apex, sparse, one to three on each side of each abdominal segment, longest spine on submedial area of abdomen 17 µm long (paratypes 14–19 µm), longest spine on thorax and head 14 µm long (paratypes 14–23 µm). Tubular ducts scattered over body ( Figs 1q View FIGURE 1 , 3o View FIGURE 3 ). Multilocular pores absent. Short conical setae similar to simple pores, either dome-shaped or acorn-shaped, with conical projection, definite setal base ( Figs 1b View FIGURE 1 , 3a View FIGURE 3 ), one to several clustered near base of each antenna, 4 µm in height, 4 µm wide (paratypes 3–5 µm by 3–4 µm). Simple pores each with slightly raised, rounded center, scattered over body ( Figs 1e View FIGURE 1 , 3f View FIGURE 3 ). One paratype specimen with structure near each antenna with unequal sides, without base, longest side 7 µm. Anal lobes each bearing enlarged fimbriate seta 48 µm long (paratypes 27–33 µm), enlarged capitate seta broken on holotype (paratypes 48–53 µm), spine with swollen apex, 27 µm long (paratypes 17–21 µm), on each side of anal ring ( Fig. 1n View FIGURE 1 ). Microtrichia absent. Derm smooth.

Anal ring ( Fig. 3l View FIGURE 3 ) wrapping around margin, invaginated in pocket, forming setal basket, with ring of conical pores, bearing six robust anal-ring setae with truncate apices, about equal in size, each about 39 µm long (paratypes 29−33 µm), 0.7 times (paratype 0.7−1.0 times) as long as greatest diameter of anal ring.

Venter with two pairs of thoracic spiracles ( Figs 1d View FIGURE 1 , 3d View FIGURE 3 ), spiracular openings projecting above derm, three pairs of abdominal spiracles ( Fig. 1f View FIGURE 1 ), spiracular openings flush with derm.Atria of thoracic and abdominal spiracles each with ring of pores, without loculi, in single row. Anterior thoracic spiracles with 10 (paratypes 6–9) pores in ring around opening, 12 µm in diameter (paratypes 10–12 µm). Abdominal spiracles with 7–11 (paratypes 7–11) pores in ring around opening, anterior abdominal spiracular opening 13 µm in diameter (paratypes 12–15 µm). Body setae ( Fig. 1k View FIGURE 1 ) with blunt apices, not capitate, those on abdomen longer than those on thorax and head, longest seta on thorax or head 14 µm long (paratypes 10–14 µm), longest seta on abdomen 36 µm long (paratypes 31–39 µm). Robust fimbriate setae and spines with swollen apices marginal. Eye absent. Labium two-segmented, small, about as wide as long, round, bent upward on holotype (paratypes 34–44 µm long by 31–42 µm wide), only one seta arising from basal sockets. Vulva wide, 230 µm wide (paratypes 72–201 µm), with smooth margins, one heavily sclerotized lateral bar on each side of vulva. Margins of abdominal segments with heavily sclerotized areas (probably muscle attachment points) (not illustrated in Figs 1 View FIGURE 1 , 3 View FIGURE 3 , see Fig. 2 View FIGURE 2 ), the posterior two sclerotizations each typically with two associated tubular ducts on each side. Compound duct clusters ( Figs 1i View FIGURE 1 , 3i View FIGURE 3 , 4 View FIGURE 4 Ab and Ba) present marginally, clusters of tubular ducts with quadrilocular centers, spines arising from between quadrilocular tubular ducts, clusters set in sclerotized ring, three pairs on submargins of abdomen, one on each side of each of segments VI –VIII, two medially (one posterior to vulva, one anterior to vulva), eight duct clusters total on abdomen, median clusters 32–34 µm in diameter (paratypes 19–39 µm), 26 ducts per median cluster (paratypes 14–27), marginal clusters 27–28 µm in diameter (paratypes 22–32 µm), 15–21 ducts per marginal cluster (paratypes 10–18). Multilocular pores absent. Dome- and acorn-shaped setae, with raised rounded blunt apex, singly or in cluster on head, about same size as those on dorsum. Simple pores few, scattered on thorax and abdomen, about same size as those on dorsum. Microtrichia absent. Derm smooth. Anal lobes indistinct, each side of lobe area bearing group of two enlarged capitate setae, 52 µm long (paratypes 52–53 µm), one small fimbriate seta (both broken on holotype) (paratypes 10–12 µm), one spine with swollen apex, 21 µm long (paratypes 20–21 µm). Antennae ( Fig. 1a View FIGURE 1 ) each three-segmented, segments short, segments I and III subequal in length, total length about 78 µm (paratypes 53–66 µm), segment I about 30 µm long (paratypes 18–37 µm), segment II about 17 µm long (paratypes 15–18 µm), apical segment about 31 µm long (paratypes 24–35 µm). Apical segments each with one long fleshy apical seta, straight with rounded blunt tip (broken on holotype) (paratypes 34–38 µm), one short fleshy subapical seta, curved with rounded blunt tip, 17 µm long (paratypes 15–20 µm), third fleshy seta at apex, microseta adjacent to apical setae, one capitate spine on apex, unspecialized setae long and straight. Pseudobasal antennal segment absent. Legs ( Figs 1h View FIGURE 1 , 3e View FIGURE 3 ) reduced in size, all about equal, trochanter and femur fused, two campaniform sensilla on each surface of trochanter, tibia and tarsus not fused, tarsus one-segmented, coxa 54 µm long (paratypes 45–51 µm), trochanter + femur 60 µm long (paratypes 54–63 µm), tibia 25 µm long (paratypes 19–22 µm), tarsus 43 µm long (paratypes 34–38 µm), tibia/tarsus 0.6 (paratypes 0.5–0.6), claw 19 µm long (paratypes 14–23 µm). Leg setae sparse, straight, spiniform dorsally and setiform ventrally, enlarged spines between tibia and tarsus absent. Sensory pore between tibia and tarsus present, sensory seta between tibia and tarsus absent. Tarsal digitules absent. Claw digitules each spine-like with acute apex, shorter than claw, claw denticle absent.

Notes: Barbenigma biza differs from B. boscus (described below) ( B. boscus character states in parentheses) in having: more spines on the dorsal abdomen, spines typically occurring in pairs on submargin of each side of each abdominal segment (spines occurring singly); body setae with blunt, but not capitate, apices (body setae with capitate apices); spines on the dorsum short, 14–23 µm long (spines on the dorsum long, 22–31 µm long); and dorsal fimbriate setae short, 16–30 µm long (dorsal fimbriate setae long, 26–39 µm long).

We illustrated two forms of adult female. The change in body shape between young adult females ( Fig. 3 View FIGURE 3 ) and older females with eggs ( Fig. 1 View FIGURE 1 ) is remarkable, with the young adult females thin and elongate, half the length and only a quarter of the width of mature females. These young adult females are so different in body shape that they were labeled as “male”, perhaps because previous curators thought they were second-instar nymphs or pre-pupae. However, they have both a distinct vulva, with lateral sclerotized bars, and quadrilocular duct clusters that we posit are homologous with an ovisac band ( Figs 3 View FIGURE 3 , 4B View FIGURE 4 ). We found no morphological differences between either form beyond the difference in body shape.

The simple pores on the thorax and abdomen may not be homologous to the ones on the head, and we call these simple pores rather than setae. The ones on the head have a lot of depth, easily visible when the structures are viewed from the side. However, those on the thorax and abdomen (both ventral and dorsal) did not have such obvious depth.

Third-instar? nymph

( Fig. 5 View FIGURE 5 )

Description: Slide-mounted specimen 0.76 mm long, 0.28 mm wide, body elongate oval (n=1).

Dorsum with robust fimbriate setae ( Fig. 5j, n View FIGURE 5 ), often with up to five tines, sparse, in segmental rows on abdomen, thorax, and head, arranged in two pairs of longitudinal lines (submedial and lateral) on abdomen, longest seta on submedial area of abdomen 20 µm long, longest seta on margin 23 µm long, longest seta on head 17 µm long. Dorsal spines without setal base ( Fig. 5m View FIGURE 5 ), with enlarged base, swollen, rounded apices, in segmental rows on abdomen, thorax, and head, arranged in two pairs of longitudinal lines near fimbriate setae, longest spine on submedial area of abdomen 24 µm long, longest spine on thorax and head 24 µm long. Small tubular ducts ( Fig. 5l View FIGURE 5 ) most abundant on abdomen, rare on thorax, present submedially and mediolaterally on abdomen. Multilocular pores absent. Discoidal pores absent. Pair of dome- or acorn-shaped setae ( Fig. 5a View FIGURE 5 ), 4 µm in height, 4 µm width. Anal lobes indistinct, each side of lobe area bearing two enlarged fimbriate setae, 27 µm long, and spine 27 µm long on each side of anal ring. Microtrichia ( Fig. 5k View FIGURE 5 ) uncommon, in small patches on thorax and abdomen. Entire derm alveolar with polygonal and elongate cell-like reticulations ( Fig. 5c, f, k View FIGURE 5 ).

Anal ring ( Fig. 5i View FIGURE 5 ) wrapping around margin, invaginated in pocket, forming setal basket, with conical projections mostly obscured, with tubular structure on each side of ring, bearing six short spine-like anal-ring setae with rounded apices, about equal in size, each about 10 µm long, 0.3 times as long as greatest diameter of anal ring.

Venter with two pairs of thoracic spiracles ( Fig. 5c View FIGURE 5 ), spiracular openings each slightly projecting above derm, each atrium with ring of 4–6 pores, without loculi, in single row around spiracle, spiracular opening 10 µm in diameter. Abdomen with three pairs of spiracles ( Fig. 5e View FIGURE 5 ) on anterior segments, each volcano shaped, difficult to distinguish from cell-like reticulations, with unobvious tracheae. Robust fimbriate setae, in segmental rows on abdomen, in two pairs of longitudinal lines on abdomen (submedial and lateral), present marginally on thorax and head, longest seta on abdomen 34 µm long, longest seta on thorax 16 µm long, shorter slightly enlarged setae in cluster on segment VIII, 8 µm long, slightly longer enlarged setae near base of each antenna. Spines with enlarged base, swollen, rounded apices, submarginal on abdomen, thorax, and head, longest spine on head 25 µm long ( Fig. 5f View FIGURE 5 ). Eye absent. Labium two-segmented, small, about as wide as long, round, only one seta arising from all basal sockets. Vulva absent. Multilocular pores absent. Modified quadrilocular ducts absent. Dome- or acorn-shaped setae with raised rounded blunt apex scattered on head, same size as those on dorsum. Small tubular ducts uncommon, arranged in longitudinal line mediolaterally on each side of head, thorax, and anterior abdominal segments. Anal lobes indistinct, each anal lobe area bearing two enlarged fimbriate setae, 28 µm long, one spine (apex obscured by wax) on each side. Entire derm alveolar with polygonal cell-like reticulations. Antennae each three-segmented, segments short, total length 59 µm long, segment I 19 µm long, segment II 16 µm long, apical segment 27 µm long. Apical segment with long fleshy apical seta, straight with rounded blunt tip, 35 µm long, short fleshy subapical seta, curved with rounded blunt tip, 18 µm long, third fleshy seta at apex, 28 µm long, two microsetae adjacent to apical setae, each clubbed apically, unspecialized setae spine-like. Segment I with small, hair-like seta near base. Pseudobasal antennal segment absent. Legs ( Fig. 5b View FIGURE 5 ) reduced in size, all about equal, trochanter and femur fused, with two campaniform sensilla on each surface, tibia and tarsus not fused, tarsus one-segmented, coxa 54 µm long, trochanter + femur 48 µm long, tibia 18 µm long, tarsus 29 µm long, tibia/tarsus 0.6, claw 18 µm long. Leg setae sparse, straight, spiniform dorsally and ventrally, enlarged spines between tibia and tarsus absent. Sensory pore between tibia and tarsus present, sensory setae between tibia and tarsus absent. Tarsal digitules absent. Claw digitules each spine-like with acute apex, shorter than claw, claw denticle absent.

Notes: This specimen was collected with a second-instar nymph (described below) but no adult females. We include it in material examined but not as a paratype for this reason.

Second-instar? nymph

( Figs. 4D View FIGURE 4 , 6 View FIGURE 6 )

Description: Slide-mounted specimen 0.42–0.53 mm long, 0.22–0.35 mm wide, body rotund oval (n=7).

Dorsum with hair-like setae ( Fig. 6k View FIGURE 6 ), sparse, in three pairs of longitudinal lines (submedial, mediolateral, submarginal), in segmental rows on abdomen, scattered on thorax and head, longest seta on submedial area of abdomen 6–10 µm long, longest seta on thorax or head 5–9 µm long. Spines absent. Multilocular pores absent. Tubular duct in center of circular area present on head ( Fig. 6b View FIGURE 6 ). Dome-shaped setae, on submedial apex of head, set on raised area, 4–5 µm in height, 4 µm wide, sparse ( Fig. 5l View FIGURE 5 ). Simple pores ( Fig. 6e, j View FIGURE 6 ) scattered on thorax and abdomen, sparse. Anal lobes indistinct, each with single apical seta. Microtrichia absent. Entire derm coriaceous.

Anal ring ( Fig. 6g, h View FIGURE 6 ) usually ventral, sometimes wrapping around margin, invaginated in pocket, forming setal basket, with 6 or 8 pores anterior to anal opening, bearing six short spine-like anal-ring setae each with acute to slightly rounded apex, about equal in size, each about 10 µm long, most specimens with setae coming straight up on slide, difficult to measure, about 0.7 times as long as greatest diameter of anal ring, 15–23 µm in diameter, derm with heavier sclerotization around anal ring.

Venter with two pairs of thoracic spiracles ( Fig. 6c View FIGURE 6 ), spiracular openings projecting above derm, atrium with ring of 4–5 pores, without loculi, in single row, spiracular opening 7–9 µm in diameter.Abdominal spiracles unobvious or absent. Hair-like setae sparse, in two pairs of longitudinal lines (marginal and submedial), longest seta on submedial area of abdomen 4–9 µm long, longest seta on thorax or head 6–9 µm long. Eyes absent. Labium two-segmented, small, about as wide as long, round, 25–31 µm long, 30–34 µm wide, only one seta arising from basal sockets. Vulva absent. Multilocular pores absent. Modified quadrilocular ducts absent. Tubular ducts, each in center of circular area, present on each side of head, sometimes also present between antennal bases, smaller ducts present near base of each leg. Dome- or acorn- shaped setae; 4–5 µm in height, 4 µm wide, sparse. Simple pores sparsely scattered on head, anterior to each coxa, and laterally on anterior abdominal segments, same size as those on dorsum. Cluster of 6 short hair-like setae anterior to anal ring, 3–4 µm long, two elongate hair-like setae on each side of anal ring, 23–27 µm long. Anal lobes indistinct, each lobe area with two hair-like setae, 9–11 µm long, single dome- or acorn-shaped seta set on raised base on margin ( Fig. 6g, h View FIGURE 6 ). Antennae coriaceous, each three-segmented, segments short, total length 27–49 µm long, segment I 8–16 µm long, segment II 9–12 µm long, apical segment 17–22 µm long. Apical segment with long fleshy apical seta, straight with rounded blunt tip, 31 µm long, short fleshy subapical seta, curved with a rounded blunt tip, 9–15 µm long, unspecialized setae hair-like. Pseudobasal antennal segment absent.

Legs coriaceous, reduced in size, all about equal, trochanter and femur fused, with two campaniform sensilla on each surface, tibia and tarsus not fused, tarsus one-segmented, area anterior to each coxa with more sclerotization, coxa 32–39 µm long, trochanter + femur 28–35 µm long, tibia 6–14 µm long, tarsus 9–18 µm long, tibia/tarsus 0.7–0.9, claw 11–15 µm long. Leg setae sparse, straight, spiniform dorsally and setiform ventrally, enlarged spines between tibia and tarsus absent. Sensory pore between tibia and tarsus present, sensory setae between tibia and tarsus absent. Tarsal digitules absent. Claw digitules each spine-like with acute apex, shorter than claw, claw denticle absent.

Notes: Only one of the six nymphs examined was collected in association with an adult female to verify the identification. However, we could find no differences between the specimens and utilized all of them in the description.

First-instar? nymph

( Fig. 7 View FIGURE 7 )

Description: Slide-mounted specimen 0.34 mm long, 0.14 mm wide, body rotund oval (n=1).

Dorsum with hair-like setae ( Fig. 7h View FIGURE 7 ) sparse, arranged in rows, with two pairs of longitudinal lines (submedial and submarginal) on abdomen, thorax, and head, about 9 µm long. Spines and fimbriate setae absent. Small tubular ducts ( Fig. 7j View FIGURE 7 ) uncommon, restricted to abdomen. Tubular ducts ( Fig. 7b View FIGURE 7 ) in center of circular area on each side of head, present on alveolar nodules. Multilocular pores absent. Simple pores scattered on abdomen, each with slightly raised center ( Fig. 7e, g View FIGURE 7 ). Pair of acorn- or dome-shaped seta ( Fig. 7a, k View FIGURE 7 ) on submedial apex of head, 4 µm in height, 4 µm in width. Simple pores sparsely scattered on thorax and abdomen. Anal lobes indistinct, each lobe area bearing two hair-like setae, 25 µm long, one acorn-shaped seta on posterior margin ( Fig. 7f View FIGURE 7 ). Microtrichia absent. Entire derm alveolar with polygonal and elongate cell-like reticulations ( Fig. 7b, c, f, i View FIGURE 7 ).

Anal ring ( Fig. 7f View FIGURE 7 ) ventral, near abdominal apex, invaginated in pocket, forming setal basket, with 6 pore-like structures at anterior apex of ring, bearing six short, spine-like, anal-ring setae each with slightly rounded apex, about equal in size, 8 µm long, 0.4 times as long as greatest diameter of anal ring.

Venter with two pairs of thoracic spiracles ( Fig. 7c View FIGURE 7 ), spiracular openings slightly projecting above derm, atrium with ring of 4–5 pores, these without loculi, in single row, spiracular opening 8 µm in diameter.Abdominal spiracles unobvious or absent. Hair-like setae in two pairs of longitudinal lines (submedial and lateral) on each side of body, present on each abdominal segment, 8 µm long. Spines and fimbriate setae absent. Eye absent. Labium not clearly visible. Vulva absent. Multilocular pores absent. Modified quadrilocular ducts absent. Simple pores present near each pair of legs, with associated small tubular duct, also present in submarginal line on each side of some anterior abdominal segments, these about the same size as those on dorsum. Small tubular ducts uncommon, arranged in longitudinal line mediolaterally on each side of head, thorax, and anterior abdominal segments.Anal lobes indistinct, each lobe area bearing enlarged hair-like seta on margin, about 6 µm long, two hair-like ventral setae level with anal ring, each about 25 µm long. Acorn-shaped seta submedially, between antennal bases, 3 µm in height, 3 µm width. Entire derm alveolar with polygonal cell-like reticulations. Cluster of 4 short hair-like setae anterior to anal ring, 3–4 µm long, two elongate hair-like setae on each side of anal ring, 23–27 µm long, acorn-shaped seta on each side of anal ring at posterior apex. Antennae each three-segmented, segments short, total length 39 µm.Apical segment with long fleshy apical seta, straight with rounded blunt tip, 29 µm long, short fleshy subapical seta, curved with rounded blunt tip, 14 µm long, third fleshy seta at apex, 19 µm long, unspecialized setae long and hair-like. Pseudobasal antennal segment absent. Legs reduced in size, all about equal. Trochanter and femur fused, with two campaniform sensilla on each surface. Tibia and tarsus not fused, tarsus one-segmented. Coxa 37 µm long, trochanter + femur 31 µm long, tibia 10 µm long, tarsus 16 µm long, tibia/tarsus 0.6, claw 12 µm long. Leg setae sparse, straight, spiniform dorsally and ventrally, enlarged spines between tibia and tarsus absent. Sensory pore between tibia and tarsus present, sensory setae between tibia and tarsus absent. Tarsal digitules absent. Claw digitules each spine-like with acute apex, shorter than claw, claw denticle absent.

Notes: The illustration and description are based on a single specimen that was mounted laterally and not completely cleared, making it difficult to determine the exact location of some structures. This specimen was collected in association with the holotype adult female.

FSCA

USA, Florida, Gainesville, Division of Plant Industry, Florida State Collection of Arthropods

USNM

USA, Washington D.C., National Museum of Natural History, [formerly, United States National Museum]

FSCA

Florida State Collection of Arthropods, The Museum of Entomology

USNM

Smithsonian Institution, National Museum of Natural History

VI

Mykotektet, National Veterinary Institute

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Hemiptera

Family

Ortheziidae

SubFamily

Carayoneminae

Tribe

Barbenigmini

Genus

Barbenigma

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