Puto philo Powell & Miller, 2024

Powell, Erin C. & Miller, Douglass R., 2024, Three new species of Puto giant mealybugs (Hemiptera: Coccomorpha: Putoidae) from Mexico and Guatemala, with keys to New World adult females and World adult males, Zootaxa 5443 (3), pp. 301-336 : 305-317

Puto philo Powell & Miller , sp. n.

Suggested common name: philodendron giant mealybug

Material examined

Holotype adult ♀, second specimen from left of four specimens when examining slide with naked eye. Left label: “ Puto / 521475 / On Monstera / Mexico: at Laredo / Sept. 18, 1952 / Leary, Cary Fouts / Laredo 53226.” Right label gives map of holotype position and states “ Puto philo / Powell & Miller / n.sp.” Holotype slide (USNM) also contains 1 ad. ♀, 1 third-instar ♀, 1 second-instar ♂ (repeated below) .

Paratypes: MEXICO: taken in quarantine at Brownsville , Texas, XI-7-1955, on Philodendron sp. , collector?, 56955 (2 ad. ♀♀ together on 1 slide) NHM ; taken in quarantine at Brownsville , Texas, VII-2-1956, on Philodendron sp. , Heinrich and Allen, 561098 (1 third-instar ♀ on 1 slide) USNM ; taken in quarantine at Brownsville , Texas, VIII-14-1956, on Philodendron cuttings, collector?, 561108 (1 second-instar ♂ on 1 slide) USNM ; taken in quarantine at Brownsville , Texas, IX-19-1956, on Philodendron seedlings, collector?, 561197 (3 third-instar ♀♀, 5 second-instar ♂♂, 1 first-instar nymph together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, V-13-1952, on Philodendron sp. , Cary and Fouts, 52809 (1 second-instar ♂, 2 first-instar nymphs together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, VI-26-1952, on Philodendron sp. , Cary and Fouts, 521001 (1 ad. ♀, 2 second-instar ♀♀, 2 second-instar ♂♂, 2 first-instar nymphs together on 1 slide) FSCA ; taken in quarantine at Laredo , Texas, VII-29-1952, on Philodendron sp. , Lewis and Fouts, 521147 (2 second-instar ♀♀, 2 second-instar ♂♂, 3 first-instar nymphs together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, VIII-15-1952, on Philodendron sp. , Cary and Fouts, 521153 (1 ad. ♀ on 1 slide) UCD ; taken in quarantine at Laredo , Texas , VIII-26-1952, on Philodendron sp. , Cary , Weedmark and Fouts, 521447 (1 ad. ♀, 3 second-instar ♂♂ together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, VIII-30-1952, on Philodendron sp. , Ragsdale and Fouts, 521446 (1 second-instar ♀, 3 second-instar ♂♂, together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, IX-12-1952, on Monstera sp. , Cary and Heinrich, 521445 (2 ad. ♂♂, 5 second-instar ♂♂, 1 second-instar ♀ on 2 slides) USNM ; taken in quarantine at Laredo , Texas , X-7-1952, on Philodendron sp. , Watt, Fouts, Babb, and Ostrem, 521579 (1 ad. ♀, 1 third-instar ♀, 2 second-instar ♂♂ together on 1 slide) FSCA ; taken in quarantine at Laredo , Texas , XI-6-1952, on Philodendron sp. , Leary, Lewis and Fouts, 521648 (5 second-instar ♂♂, 1 first-instar nymph together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, IX-18-1952, on Monstera, Leary, Cary and Fouts , 521475 (1 ad. ♀ holotype, 1 third-instar ♀ and 1 second-instar ♂ on one slide, second slide containing 1 ad. ♀ paratype and 1 second-instar ♂) both USNM ; taken in quarantine at Laredo , Texas, I-12-1953, on “ Monstera canes”, Danos and Fouts, 530415 (1 ad. ♂ on 1 slide) FSCA ; taken in quarantine at Laredo , Texas, I-12-1953, on Monstera canes, Danos and Fouts, 530416 (2 second-instar ♂♂, together on 1 slide) USNM ; taken in quarantine at Laredo , Texas, III-3-1953, on Philodendron sp. , Cary and Fouts, 530612 (2 ad. ♀♀ together on 1 slide) USNM . Vera Cruz , XII-16-1952, on Philodendron sp. , Lewis and Averill, 530437 (1 ad. ♀ on 1 slide) USMN .

Etymology: The species epithet “ philo ” is a noun in apposition based on an abbreviation of the scientific name of the host plant genus, Philodendron .

Adult female

( Fig. 1 )

Description: Slide-mounted holotype 2.49 mm long, 1.56 mm wide; paratypes 2.55–4.83 mm long, 1.78–3.34 mm wide; body elongate oval.

Dorsum with 16 pairs of cerarii; anal-lobe cerarii each containing 11 lanceolate setae (paratypes 10–12) and 23 trilocular pores (paratypes 19–25); frontal cerarii each containing 15 lanceolate setae (paratypes 9–13) and 15 trilocular pores (paratypes 9–12); antefrontal cerarii not distinct, each containing 5–9 loosely clustered setae, without basal sclerotization; cerarii without associated discoidal pores or oral-rim type tubular ducts; and marginal cerarii with basal sclerotization. Anterior and posterior ostioles present. Multilocular disc-pores absent; trilocular pores of 1 size, numerous, scattered over body and present on lips of ostioles; discoidal pores scattered over body. Oral-rim type tubular ducts and oral-collar tubular ducts absent. Dorsal setae lanceolate, without basal sclerotization, forming rows across abdominal segments II–VII, clustered on lips of ostioles and scattered on head and thorax, with many on margin between cerarii especially on head and thorax, absent from medial area of abdominal segment VIII; longest seta on abdomen 21 µm (paratypes 21–25 µm) long, longest seta in abdominal cerarii 31 µm (paratypes 27–33 µm) long.

Anal ring dorsal, bearing 6 anal-ring setae with acute apices; posterior anal-ring setae each about 201 µm long (paratypes 207–217 µm); 1.3 times (paratype 1.5–1.6 times) as long as greatest diameter of anal ring.

Venter with multilocular disc-pores in posterior areas of abdominal segments IV–VIII, in anterior areas of segments V–VIII, sparse on segment IV, those on abdomen present medially and submedially, sparse or absent on margin; also present on medial areas of head near mouthparts; multilocular disc-pores on abdomen predominantly each 9-locular, with some 10-locular, those on head 7-locular. Trilocular pores of 1 size, numerous, scattered over body; discoidal pores scattered over body. Oral-collar tubular ducts of 2 sizes; shorter oral-collar tubular ducts on abdomen, in transverse rows across segments II–VIII, numerous on submarginal areas of segments VI–VIII, rare on head and thorax; longer oral-collar tubular ducts absent from head and thorax of holotype female (paratype females with 0–6 in medial area of head anterior to mouthparts). Long hair-like setae arranged in rows across abdomen, longest seta on medial area of abdomen 62 µm (paratypes 69–76 µm) long, longest seta on medial area of segment V 62 µm (paratypes 60–73 µm) long; head setae broken on holotype (longest seta on head of paratypes 137–163 µm long). Circulus divided into 2 oval parts on segment III, sometimes asymmetrical in size, diameter of each part 117–127 µm (paratypes 66–142 µm). Labium 254 µm long (paratypes 226–257 µm). Antennae each 9-segmented, total length about 802 µm (paratypes 691–794 µm); apical segment about 133 µm long (paratypes 113–133 µm), segment III about 101 µm long (paratypes 88–104 µm); with intersegmental sensilla present between segments III and IV, IV and V, and VI and VII; and with a pair of coeloconic sensilla on segments IV, V and IX. One paratype female with aberrant antennae, having one side with 5 segments and the other side with pseudoarticulations, incompletely forming nine segments. Hind tibiae each with small translucent pores. Femora each about 386 µm long (paratypes 333–392 µm), tibiae each about 464 µm long (paratypes 438–484 µm), tibia/tarsus 2.8 (paratypes 2.7–2.9). Hind trochanters each with 3 or 4 campaniform sensilla on each surface. Tarsal digitules capitate, claw digitules capitate and longer than claw; claw denticle absent; basal spurs present.

Diagnosis: Adult females of P. philo differ from other Puto species by having the following combination of characters: a single, divided circulus; claw denticle absent; and dorsal oral-rim type tubular ducts, including those associated with cerarii, absent.

Puto philo is similar to P. lamottei Matile-Ferrero, 1985 in having no oral-rim type tubular ducts in the cerarii, no dorsal oral-rim type tubular ducts in the medial areas, and a divided circulus. Puto philo lacks a claw denticle, whereas P. lamottei has one.

The new species is also similar to P. spath (described below) in that both have a divided circulus, but no dorsal oral-rim type tubular ducts in medial areas, and no oral-rim type tubular ducts in the cerarii. However, P. philo differs from P. spath as follows (character states of P. spath are presented in parentheses): (i) claw denticle absent (claw denticle present); (ii) no multilocular disc-pores in the medial area of the thorax (with multilocular disc-pores in the medial area of the thorax); and (iii) with typical multilocular disc-pores of one size (multilocular disc-pores of two sizes, with large multilocular disc-pores each with a trilocular center).

Puto philo also resembles P. paramoensis Matile-Ferrero, 1985 in having no denticle, no dorsal oral-rim type tubular ducts, and no dorsal multilocular disc-pores. However, it differs from P. paramoensis as follows (character states of P. paramoensis are presented in parentheses): (i) ventral multilocular disc-pores on the thorax absent (with many ventral multilocular disc-pores in the medial and marginal areas of the thorax); (ii) multilocular disc-pores predominantly each with 9 loculi but with some 10-locular disc-pores near the vulva (two distinct sizes of multilocular disc-pores, 10- and 6-locular, with the larger size near the vulva); and (iii) claw digitules capitate (claw digitules simple).

Third-instar female

( Fig. 2 )

Description: Slide-mounted, 1.57–2.08 mm long, 0.96–1.26 mm wide; body elongate oval.

Dorsum with 16 pairs of cerarii; anal-lobe cerarii each containing 7–9 lanceolate setae and 16‒19 trilocular pores; frontal cerarii each containing 6–11 lanceolate setae and 5–8 trilocular pores; antefrontal cerarii not distinct, without sclerotization, each containing loose cluster of 7–11 setae; cerarii without associated discoidal pores or oral-rim type tubular ducts; and marginal cerarii each with basal sclerotization. Anterior and posterior ostioles present. Multilocular disc-pores absent; trilocular pores of 1 size, numerous, scattered over body, with more on dorsum than on venter, also present on lips of ostioles; discoidal pores scattered over body. Oral-rim type tubular ducts and oral-collar tubular ducts absent. Dorsal setae lanceolate, forming rows across abdominal segments II–VII, also clustered on lips of ostioles and scattered on head and thorax, with many on margin between cerarii, but absent from medial area of abdominal segment VIII; longest seta on abdomen 17–23 µm long, longest seta in abdominal cerarii 26–28 µm long.

Anal ring dorsal, bearing 6 anal-ring setae with apices acute; posterior anal-ring setae each 135–157 µm long, 1.3–1.5 times as long as greatest diameter of anal ring.

Venter with multilocular disc-pores present in medial area of thorax near mouthparts, predominantly each 7-locular. Trilocular pores of 1 size, scattered over body; discoidal pores scattered over body. Long oral-collar tubular ducts forming cluster (12–20) on head anterior to mouthparts, absent from thorax and abdomen. Long hair-like setae forming rows across abdomen, longest ventral seta on medial area of abdomen 47–57 µm long, longest seta on medial area of segment V 43–53 µm long, longest seta on head 95–114 long. Circulus divided into 2 oval parts on segment III, sometimes asymmetrical in size, each part 34–62 µm in diameter. Labium 191–206 µm long. Antennae each 8-segmented, total length 579–654 µm; apical segment 114–132 µm long, segment III 80–97 µm long; antennal intersegmental sensilla present between segments V and VI; coeloconic sensilla present on segments IV and VIII. Hind tibia and distal femur with few translucent pores. Femora each 264–285 µm long; tibiae each 316–337 µm long, tibia/tarsus 1.9–2.1. Hind trochanters each with 2 campaniform sensilla on each surface. Tarsal digitules capitate, claw digitules capitate and longer than claw; claw denticle absent but basal spurs present.

Diagnosis: A comparison of the third-instar females of P. philo and P. spath is presented in the “Diagnosis” section of the latter species.

Second-instar female

( Fig. 3 )

Description: Slide-mounted, 1.41–1.74 mm long, 0.79–1.12 mm wide; body elongate oval.

Dorsum with 16 or 17 pairs of cerarii; anal-lobe cerarii each containing 5–7 lanceolate setae and 11–13 trilocular pores; frontal cerarii each with basal sclerotization, containing 5–7 lanceolate setae and 4–6 trilocular pores; antefrontal cerarii not distinct, without sclerotization, each containing 4–6 loosely clustered setae; cerarii each without associated discoidal pores or oral-rim type tubular ducts; and marginal cerarii each with weak basal sclerotization. Ostioles weakly developed. Multilocular disc-pores absent; trilocular pores of 1 size, larger than those on venter, numerous, scattered over body, with more on dorsum than on venter, also present on lips of ostioles; discoidal pores scattered over body. Oral-rim type tubular ducts and oral-collar tubular ducts absent. Dorsal setae lanceolate, few, scattered across body, present on lips of ostioles; longest seta on abdomen 17–19 µm long, longest seta in abdominal cerarii 25–28 µm long.

Anal ring dorsal or bent around body apex, bearing 6 anal-ring setae with apices acute; posterior anal-ring setae each 121–137 µm long, 1.3–1.8 times as long as greatest diameter of anal ring.

Venter with multilocular disc-pores in medial area of pro- and mesothorax, predominantly each 7-locular. Trilocular pores of 1 size, smaller than those on dorsum, scattered over body; discoidal pores scattered over body. Long oral-collar tubular ducts of 1 size forming cluster (6–11) anterior to mouthparts. Long hair-like setae sparse, forming rows across abdomen, longest on head, longest seta on medial area of abdomen 38–56 µm long, longest seta on medial area of segment V 33–49 µm long, longest seta on head 99–111 µm long. Circulus divided into 2 oval parts on segment III, each part 24–38 µm in diameter. Labium 154–167 µm long. Antennae each 7- segmented, total length 506–554 µm, apical segment 115–125 µm long, segment III 96–116 µm long; antennal intersegmental sensilla present between segments IV and V; coeloconic sensilla present on segments III and VII. Hind legs without translucent pores. Femora each 216–233 µm long; tibiae each 233–286 µm long, tibia/tarsus 1.6–1.9. Hind trochanters each with 2 campaniform sensilla on each surface. Tarsal digitules capitate, claw digitules capitate, longer than claw; claw denticle absent; basal spurs present.

Diagnosis: A comparison of the second-instar females of P. philo and P. spath is presented in the “Diagnosis” section of the latter species.

Second-instar male

( Fig. 4 )

Description: Slide-mounted, 1.54–2.56 mm long, 0.88–1.60 mm wide; body elongate oval.

Dorsum with 16 or 17 pairs of cerarii; anal-lobe cerarii each containing 4–8 lanceolate setae and 10–16 trilocular pores; frontal cerarii each with basal sclerotization, containing 5–7 lanceolate setae and 4–11 trilocular pores; antefrontal cerarii not distinct, without sclerotization, each containing 4–7 loosely clustered setae; cerarii without associated discoidal pores or oral-rim type tubular ducts; and marginal cerarii each with weak basal sclerotization. Anterior ostioles absent, posterior ostioles weakly developed. Multilocular disc-pores absent; trilocular pores of 1 size, larger than those on venter, numerous, scattered over body, with more on dorsum than on venter, few present on anterior lip of ostioles; discoidal pores scattered over body. Oral-collar tubular ducts of 1 size, scattered over body. Oral-rim type tubular ducts absent. Dorsal setae lanceolate, few, scattered across body; few present on anterior lip of ostiole; longest seta on abdomen 17–20 µm long, longest seta in abdominal cerarii 24–26 µm.

Anal ring dorsal, bearing 6 anal-ring setae with apices acute; posterior anal-ring setae each 144–163 µm long, 1.5–1.8 times as long as greatest diameter of the anal ring.

Venter with multilocular disc-pores in medial area of pro- and mesothorax, predominantly each 7-locular. Trilocular pores of 1 size, smaller than those on dorsum, scattered over body; discoidal pores scattered over body. Oral-collar tubular ducts of 2 sizes, shorter ones scattered over entire body, long oral-collar tubular ducts forming cluster (5–10) anterior to mouthparts. Long hair-like setae sparse, forming rows across abdomen, longest on head, longest seta on medial area of abdomen 46–69 µm long, longest seta on medial area of segment V 43–52 µm long, longest seta on head 103–147 µm long. Circulus divided into 2 oval parts on segment III, in some specimens each part fully separated, in other specimens pieces touching, each part 32–55 µm in diameter. Labium 169–180 µm long. Antennae each 7-segmented, total length 532–584 µm, apical segment 111–116 µm long, segment III 117–129 µm long; antennal intersegmental sensilla present between segments IV and V; coeloconic sensilla present on segments III and VII. Hind legs without translucent pores. Femora each 236–273 µm long; tibiae each 278–327 µm long, tibia/tarsus 1.7–2.2. Hind trochanters each with 2 campaniform sensilla on each surface, one specimen with 3 on one side. Tarsal digitules capitate, claw digitules capitate, longer than claw; claw denticle absent; basal spurs present.

Diagnosis: A comparison of the second-instar males of P. philo and P. spath is presented in the “Diagnosis” section of the latter species.

First-instar nymph (sexes undetermined)

( Fig. 5 )

Description: Slide- mounted, 0.80–1.16 mm long, 0.43–0.70 mm wide; body elongate oval.

Dorsum with 18 or 19 pairs of cerarii, anal-lobe cerarii each containing 2 lanceolate setae and 4–6 trilocular pores; frontal cerarii each containing 2–3 lanceolate setae and 2–4 trilocular pores; antefrontal cerarii containing 1 or 2 setae, with basal sclerotization, situated laterad of frontal cerarius; sometimes with another pair of setae posteriorly without basal sclerotization; cerarii each without associated discoidal pores or oral-rim type tubular ducts; and marginal cerarii with basal sclerotization. Anterior ostioles absent, posterior ostioles weakly developed. Multilocular disc-pores absent; trilocular pores of 1 size, numerous, unevenly scattered across body, those on dorsum larger and more numerous than on venter, absent from lips of ostioles; discoidal pores absent. Oral-rim type tubular ducts and oral-collar tubular ducts absent. Dorsal setae lanceolate, few, forming 2 pairs of longitudinal lines on thorax and abdomen, absent from lips of ostioles; longest seta on abdomen 14–20 µm long, longest seta in abdominal cerarii 24–26 µm long.

Anal ring dorsal, bearing 6 anal-ring setae with apices acute, posterior anal-ring setae each 112–122 µm long, 1.7–1.9 times as long as greatest diameter of the anal ring.

Venter with multilocular disc-pores present near mouthparts, near spiracles, and on thorax, predominantly each with 7-loculi. Trilocular pores of 1 size, scattered over body. Discoidal pores marginal, associated with marginal hair-like setae on abdomen, and in cluster on margin of head. Cluster of 2–4 long oral-collar tubular ducts present anterior to mouthparts. Long hair-like setae longest on head, forming rows across abdomen, longest seta on medial area of abdomen 31–46 µm long, longest seta on medial area of segment V 29–36 µm long, longest seta on head 99–125 µm long. Single undivided circulus present on abdominal segment III, 27–33 µm in diameter. Labium 128– 134 µm long. Antennae each 7-segmented, total length 421–468 µm, apical segment 108–116 µm long, segment III 57–79 µm long; antennal intersegmental sensilla present between segments IV and V; coeloconic sensilla present on segment VII. Hind legs without translucent pores. Femora each 166–237 µm long; tibiae each 176–212 µm long, tibia/tarsus 1.3–1.5. Hind trochanters each with 2 campaniform sensilla on each surface. Tarsal digitules capitate, claw digitules capitate, each longer than claw; claw without denticle; basal spurs present.

Diagnosis: A comparison of the first-instar nymphs of P. philo and P. spath is presented in the “Diagnosis” section of the latter species.

Adult male (macropterous)

( Figs 6 , 14a )

Description: Slide-mounted, 2.85–3.21 mm long, 0.77–0.80 mm wide; body elongate, with segment VIII produced laterally.

Dorsum with 1 pair of glandular pouches (gp); each glandular pouch with 2 central setae, each apically acute, setae all approximately same size, 247–302 μm long. Multilocular disc-pores present in glandular pouches, each pouch containing about 263–290 tightly clustered pores, predominantly each with 5 loculi; discoidal pores predominantly situated on margin of pouch; setae forming row across anterior margin of pouch; and with tubular ducts (gpt) forming cluster at base of glandular pouch setae (gls). Multilocular disc-pores forming cluster on prothorax, present in segmental rows from metathorax to abdominal segment VII, predominantly each with 4 or 5 loculi. Discoidal pores present near multilocular disc-pores. Hair-like setae slender, apically acute, approximately same length as those on venter, forming segmental rows, with marginal clusters on each side of each segment; setae present on pro-, meso- and metathorax. Head with setae anterior to postoccipital ridge (por), on ocular sclerite (ocs), and along dorsal arm of midcranial ridge (dmcr). Microtrichia present on all abdominal segments (not illustrated). Posterior ostioles (pos) present, weakly developed, without associated setae or pores. Abdominal tergites (at) with two areas of sclerotization on posterior margin of each of segments II–VII; segment VIII with heavily sclerotized yoke-shaped medial tergal plate (mtp) on anterior margin; metapostnotal sclerite present (pn 3); metapostnotal ridge absent. Scutellum (scl) well developed, with 42–48 setae; scutellar ridge (sclr) well developed. Scutum (sct) sclerotized throughout, except for medial triangular clear area (mta) anterior to scutellum. Prescutum, prescutal suture, and prescutual setae absent. Pronotal ridges (prnr) well developed; pronotal sclerites (prn) present. Hamulohalteres present, each with 3 setae (sometimes only 2) (hams). Wings each 2.62–3.04 mm long, covered in microtrichia, each with alar lobe (al), alar sclerites (alsc) and veins (wv), without setae or sensoria; subcosta (sc) arising from alar sclerites; radius (r) joining subcosta; possible media (m) represented by sclerotized area arising ¼ distance from alar sclerite, cubitus anterior (cua) present, not joined with other veins; cubitus posterior (cup) arising from alar sclerites, with additional band of sclerotization anterior to cubitus anterior; irregular patches of sclerotization present on distal half of each wing. Tegula (teg) each with 12–14 setae. Postoccipital ridge (por) well developed, fused with postocular ridge (pocr). Dorsal arm of midcranial ridge (dmcr) narrow, touching postoccipital ridge posteriorly; with anterior swelling, connected to lateral (lmcr) arms anteriorly; surrounded by median crest (mc) with irregular sclerotization and 13–20 setae on each side, also with membranous area between ocular sclerite and sclerotization surrounding dorsal arm of midcranial ridge; ocular sclerite with setae in 2 clusters, one anterolateral with 8–9 setae and one posteromedial with 7–8 setae on each side. With 3 dorsal simple eyes (de) on each side, those nearest dorsal arm of midcranial ridge (dmcr) largest, each 40–42 μm in diameter, those on lateral edge smallest, each 34–35 μm in diameter. Lateral ocellus (lo) about same size as lateral simple eye, each 42–44 μm in diameter. Anus (an) small, situated on membranous dorsal surface of penial sheath. Penial sheath (ps) 498–525 μm long; elongate, about 237 μm wide; width/length 0.45, apically with several small papillae (pap); anterior portion with 15–17 long setae (each about 48–69 μm long), posterior portion with about 10–13 small setae (each 7–9 μm long). Aedeagus (ae) Ushaped, attached on membranous surface, about 0.95–1.08 mm long, apex bifurcate with 3 small denticles.

Venter with multilocular disc-pores (mp), predominantly each with 4 loculi, present near anterior (asp) and posterior spiracles (psp), in marginal longitudinal line clusters on each segment, also present medially between mesothoracic legs, in cluster laterad to proepisternum + cervical sclerite (pepcv), in cluster (of about 15) on unsclerotized area anterior to ocular sclerite and preocular ridge (pocr) together with associated setae and discoidal pores (disc). Hair-like setae, apically acute, approximately same length as those on dorsum (79–94 μm on abdominal segment VIII), present in lateral clusters and segmental rows on abdomen, less abundant anteriorly with a few setae mediolaterally on mesothorax, and medially on metathorax and prothorax. Head setae abundant in cluster (about 21– 31 on each side) on medial area of ocular sclerite (osc) under ventral simple eyes (vse), present (9–12 on each side) near midline of head between antennal base and ventral eye (vse), anterior to preocular ridge. Microtrichia present on abdominal segments IV–VIII. Abdominal sternites with sclerotization weakly developed on medial and mediolateral areas of segments II‒VII, segment VIII with heavier sclerotization mediolaterally. Metapleural ridge (plr 3) well developed, with 2 precoxal ridge extensions, the anterior extension longer, reaching to metasternal apophysis. Mesosternum (stn 2) well developed, with large furca (f); lateropleurites (lpl) on mesothorax triangular, with membranous area between each lateropleurite and mesepisternum (eps 2). Mesopleural ridge (plr 2) with precoxal ridge of mesothorax (pcr 2) and additional posterior ridge extension anterior to mesocoxal articulation. Prosternum (stn 1) with sclerotized prosternal ridge (stnr 1) that ends with prosternal apophysis (stn1a), surrounded by poorly developed triangular sclerite. Proepisternum and cervical sclerite well developed. Preoral ridge (pror) well developed, fused with postocular ridge (pocr). Ventral midcranial ridge (vmcr) well developed anteriorly, extending to lateral arms (lmcr) between antennae, fused with preocular ridge and preoral ridge. Ocular sclerite (ocs) sclerotized throughout, area anterior to preocular ridge unsclerotized. Mouth (m) present, unsclerotized, without tubercle. Cranial apophysis (ca) sclerotized, conical, arising at base of ventral midcranial ridge. With 3 ventral simple eyes (vse) on each side, those nearest dorsal arm of midcranial ridge (dmcr) largest (each 46–48 μm in diameter), those on lateral edge smallest (each 26–30 μm in diameter); with a total of 6 pairs of simple eyes and a pair of ocelli wrapping around head. Metathoracic legs longest, with coxa (cx) 227–268 μm long, trochanter (tr) 143–175 μm long, femur (fm) 527–573 μm long, tibia (ti) 830–899 μm long, tarsus (ta) 269–285 μm long, and claw 65–71 μm long; tibia/tarsus 3.00–3.15. Hind trochanters typically each with 4 campaniform sensilla (camp) on each surface, the left metathoracic trochanter of one specimen with 3. Each claw (cl) without denticle; with pair of basal spurs (bs); digitules on tarsus (tdt) capitate, shorter than claw; digitules on claw (cdt) acute, shorter than claw. Spine-like setae present on tarsus and tibia; also with long hair-like setae on tibia, femur, trochanter, and coxa; many hair-like setae on trochanter, femur, and tibia with satellite setae (sats); short hair-like setae present on all leg segments. Legs without translucent pores.

Antennae 10-segmented, each 2.58–2.85 mm long, with segments III–VII longest, approximately equal in length, third segment about 1.2–1.5 times longer than apical segment, last segment apically rounded or pointed. Antenna with elongate hair-like setae (hs) on segments II–X, many with a very small satellite seta (sats); with short setae (shs) on first 3 segments, and antennal bristles (ab) on apical 3 segments; a pair of antennal intersegmental sensilla present between segments III and IV, IV and V, and VII and VIII (not illustrated); coeloconic sensilla present on segments II and X (not illustrated).

Remarks: The description is based on four specimens collected in association with females and immatures. The illustration utilized two specimens which are in fair condition but with the thorax twisted on both specimens. The genitalia were mounted laterally on three of the four males examined for the description. Male wing venation is interpreted with a maximalist approach, though we recognize that the veins observed may not be true veins and further study is required. For instance, in Matsucoccus pini ( Green, 1925) , only one true vein (the radius) was detected when cross-sections were examined, whereas previously this species was described as having three veins ( Franielczyk-Pietyra et al. 2023). What we have labeled as veins may be thickenings, lines, or folds rather than real veins, particularly what we interpret as the media ( Figs 6 , 13 , and 14). To our knowledge, the wing venation of Putoidae has not been studied in detail.

Diagnosis: The male of P. philo is the only known Puto male with both a bifurcate aedeagus and the claw denticle absent. It is similar to the male of P. yuccae ( Coquillett, 1890) in having the apex of the aedeagus bifurcate, but differs from it as follows ( P. yuccae characters are presented in parentheses): (i) claw without denticle (denticle present); (ii) tarsal digitules clubbed (acute); (iii) 6 pairs of simple eyes (7 or 8 pairs); (iv) with 3, sometimes 2, hamulohaltere setae (with 4); and (v) abdominal segments each with multiple rows of multilocular disc-pores (each with one row).

Figures

FIGURE 1. Puto philo Powell & Miller, sp. n., adult female holotype, Mexico at Laredo, September 18, 1952, Leary, Cary, and Fouts, on Monstera, #521475. A=dorsal seta; B=7-locular pore; C, E=discoidal pores; D, F=trilocular pores; G=oral-collar tubular duct; H=anal-lobe cerarius; I=claw; J=9-locular pore.

FIGURE 2. Puto philo Powell & Miller, sp. n., third-instar female paratype, Mexico at Laredo, September 18, 1952, Leary, Cary, and Fouts, on Monstera, #521475. A= antennal intersegmental sensilla; B, L=trilocular pores; C=long oral-collar tubular duct; D=dorsal seta; E=7-locular pore; F, G=discoidal pores; H=cerarius; I=claw; J=anal ring; K=translucent pores on hind femur and tibia.

FIGURE 3. Puto philo Powell & Miller, sp. n., second-instar female paratype, Mexico at Laredo, September 12, 1952, Cary and Heinrich, on Monstera, #521445. A=trilocular pore; B=discoidal pore; C=long oral-collar tubular duct; D=7-locular pore; E=dorsal seta; F=divided circulus; G=anal-lobe cerarius; H=claw.

FIGURE 4. Puto philo Powell & Miller,sp.n., second-instarmale paratype,Mexico at Laredo,August 26, 1952,Cary,Weedmark, and Fouts, on Philodendron, #521447. A=trilocular pore; B=long oral-collar tubular duct; C=7-locular pore; D=discoidal pore; E=dorsal seta; F, G=oral-collar tubular ducts; H=anal-lobe cerarius; I=claw.

FIGURE 5. Puto philo Powell & Miller, sp. n., first instar (sex undetermined) paratype, Mexico at Laredo, June 26, 1952, Cary and Fouts, on Philodendron, #521001. A=trilocular pore; B= antennal intersegmental sensilla; C=discoidal pore; D=long oral-collar tubular duct; E=dorsal seta; F=7-locular pore; G=anal-lobe cerarius; H=ventral seta with associated trilocular and discoidal pores; I=claw. Note that the illustration utilized two specimens on the same slide.

FIGURE 6. Puto philo Powell & Miller, sp. n., adult male paratypes, Mexico at Laredo, September 12, 1952, Cary and Heinrich, on Monstera, #521445. A=dorsum of head; B=venter of head; C=satellite seta next to long hair-like setae; D=quinquelocular pore; E=quadrilocular pore; F=glandular pouch and tubular ducts; G=lateral view of penial sheath; H=apex of aedeagus; I=claw. Note that the illustration utilized two specimens in fair condition with the thorax twisted on both specimens.

FIGURE 13. Puto brom (?) Powell & Miller, sp. n., adult male, Veracruz, Mexico at Brownsville, July 17, 1975, Burgess, Chase, and Arsego, on bromeliad leaf, #5230. A=dorsum of head; B=venter of head; C=satellite setae next to long hair-like seta; D=first four segments of antenna; E=tegula; F=scutellum; G=sternum; H=antennal intersegmental sensilla; I=wing; J=discoidal pore; K=abdomen, ventral and dorsal views; L=claw; M=quinquelocular pore; N=quadrilocular pore; O=glandular pouch; P=apex of aedeagus; Q=lateral view of penial sheath. Note that the illustrations are based on a single specimen dissected into many pieces.

FIGURE 14. The wings of (A) Puto philo Powell & Miller, sp. n. adult male paratype, Mexico at Laredo, September 15, 1952, Leary, Cary, and Fouts, on Monstera, #521475, and (B) Puto brom (?) n. sp. adult male, Veracruz, Mexico at Brownsville, July 17, 1975, Burgess, Chase, and Arsego, on bromeliad leaf, #5230.