taxonID	type	description	language	source
6A7087EF1378FFDC0AD4FC729C8B0BE8.taxon	type_taxon	Type species. Adelges (Cassiadelges) coccipus Havill & Brunet, by present designation.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFDC0AD4FC729C8B0BE8.taxon	etymology	Etymology. A noun in apposition stemming from the Latin cassida, meaning “ helmet, ” in reference to the prominent cephaloprothoracic shield in adult exules.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFDC0AD4FC729C8B0BE8.taxon	diagnosis	Diagnosis. Adult exulis. Feeds on Larix secondary hosts. Head and pronotum completely covered with plates fused into a cephaloprothoracic shield. Ventral spiracles present on abdominal segments II – VI. Only acuminate setae on abdominal segments VIII – IX.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFDC0AD4FC729C8B0BE8.taxon	discussion	Remarks. Monotypic, including only A. (Ca.) coccipus at present, pending the description of an additional species, A. (Ca.) nr. coccipus, collected from Larix in Sichuan, China (Table 1). This additional species has fully developed legs in the adult stage, so lacking legs is not a diagnostic character of Cassiadelges. Annandina is the only other subgenus in Adelges with a complete cephaloprothoracic shield in the adult exulis stage. Cassiadelges differs from Annandina in feeding on Larix vs. Tsuga secondary hosts and by Annandina having knobbed setae on the distal abdominal segments.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	description	(Fig. 2 A – J)	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	etymology	Etymology. A noun in apposition stemming from the Greek ΚΌΚΚΟς meaning “ berry ” and ΠΟΎς, meaning “ foot, ” in reference to the round vestigial legs. Also, in reference to the developmental similarity to many scale insects (Coccomorpha).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	materials_examined	Type material. HOLOTYPE: 1 adult exulis, BHUTAN, Bumthang, Shingnyer, lat. 27.4595, long. 90.8690, 5 - IV- 2022, on Larix griffithii, Kaka Tshering (USNM). PARATYPES: 1 first instar progrediens, same data (CNC # 5338333 - 15). 1 adult exulis, same data (CNC # 5338333 - 8). 1 first instar progrediens, same data (USNM). 1 adult exulis, same data (YPM # ENT 594747). 1 adult exulis, BHUTAN, Mongar, Sengor, lat. 27.3642, long. 91.0328, 27 - III- 2022, on Larix griffithii, Namgay Sacha, Kaka Tshering (CNC # 5338328 - 4). 1 nymph exulis, same data (NBC). 1 second instar exulis, same data (USNM). 1 first instar sistens, BHUTAN, Mongar, Sengor, lat. 27.3652, long. 91.0346, 27 - III- 2022, on Larix griffithii, Namgay Sacha, Kaka Tshering (CNC # 5338325 - 12). 1 second instar exulis, same data (YPM # ENT 594677). 1 adult exulis, 1 first instar sistens, same data (NBC). 1 first instar progrediens, BHUTAN, Mongar, Sengor, lat. 27.3683, long. 91.0127, 7 - IV- 2022, on Larix griffithii, Kaka Tshering (CNC # 5338330 - 10). 1 first instar progrediens, same data (NBC). 1 first instar sistens, same data (USNM). 1 first instar sistens, same data (YPM # ENT 594732). 1 first instar progrediens, same data (YPM # ENT 594731). 1 adult exulis, BHUTAN, Bumthang, Tangsibi, lat. 27.4891, long. 90.8625, 6 - IV- 2022, on Larix griffithii, Kaka Tshering (CNC # 5338331 - 5). Additional specimens examined. BHUTAN: Bumthang: Shingyner: 1 egg, 4 first instar progredientes, 12 adult exules (CNC), 1 first instar progrediens, 2 adult exules (NBC), 1 first instar progrediens, 2 adult exules (USNM), 2 first instar progredientes, 2 adult exules (YPM), lat. 27.4595, long. 90.8690, 5 - IV- 2022, on Larix griffithii, Kaka Tshering. Mongar: Sengor: 1 late instar nymph exulis, 19 adult exules (CNC), 3 adult exules (NBC), 1 late instar nymph exulis, 5 adult exules (USNM), 3 first instar sistentes, 6 late instar nymph exules, 26 adult exules (YPM), lat. 27.3642, long. 91.0328, 27 - III- 2022, on Larix griffithii, Namgay Sacha, Kaka Tshering. 1 first instar sistens, 2 late instar nymph exules, 24 adult exules (CNC), 1 first instar sistens, 1 late instar nymph exulis (NBC), 4 first instar sistentes, 6 late instar nymph exules, 37 adult exules (USNM), 14 first instar sistentes, 2 first instar sistens exuviae, 11 late instar nymph exules, 6 adult exules (YPM), lat. 27.3652, long. 91.0346, 27 - III- 2022, on Larix griffithii, Namgay Sacha, Kaka Tshering. 3 eggs, 5 first instar progredientes, 1 late instar nymph exulis, 6 adult exules (CNC), 3 first instar progredientes (USNM), 19 first instar progredientes, 9 adult exules (YPM), lat. 27.3683, long. 91.0127, 7 - IV- 2022, on Larix griffithii, Kaka Tshering. Bumthang: Tangsibi: 1 egg, 3 first instar progredientes, 1 late instar nymph exulis, 4 adult exules (CNC), 1 adult exulis (NBC), 3 first instar progredientes, 1 late instar nymph exulis, 3 adult exules (USNM), 1 first instar progrediens, 9 adult exules (YPM), lat. 27.4891, long. 90.8625, 6 - IV- 2022, on Larix griffithii, Kaka Tshering.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	diagnosis	Diagnosis. Adelges (Ca.) coccipus lacks developed legs in all exulis stages after the first instar. The general wax gland structure in the neosistens is similar to Sacchiphantes and Cholodkovskya in that the facets are arranged in a rosette pattern around a central seta, but they are more tightly packed than in the other subgenera. The abdominal spiracles in adult exules are very small and without sclerotization making them difficult to see except at high magnification. The strongly sclerotized cephaloprothoracic shield resembles many Pineus species, but having five pairs of abdominal spiracles and molecular phylogeny places this species definitively within Adelges (Fig. 1).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	description	Description. Neosistens (Fig. 2 A). Body length: n = 7, 0.359 – 0.451 (0.406) mm. Body width: n = 6, 0.202 – 0.293 (0.260) mm. Dorsal wax glands generally on lightly sclerotized plates with 3 – 6 facets, often triangular, sometimes rhomboidal to circular, with a granular appearance, arranged in tightly packed rosettes around a central seta. Head and pronotum with lightly sclerotized dorsal plates, separated from each other and along mid-line by narrow sutures. Each cephalic dorsal plate with five or six wax glands, one apical often folded ventrally in slide-mounted specimens, two anterior arranged transversely, two mesad to ocelli arranged longitudinally, sometimes one postero-spinal. Generally, 2 – 7 facets per cephalic gland. Postero-spinal gland, when present, with fewer facets than the others. The largest wax gland facets on head approximately half the diameter of ocellus. Antennae 3 - segmented, segment III imbricated, with length [n = 8, 0.041 – 0.052 (0.045) mm] 3.1 x its width at midpoint [n = 8, 0.013 – 0.016 (0.015) mm], and approximately 1.4 x longer than segments I [n = 8, 0.015 – 0.026 (0.020) mm] and II [n = 8, 0.016 – 0.024 (0.020) mm] combined. Two primary rhinaria on antennal segment III, one subapical and one approximately ½ the segment length from the apex, each with a small thumb-like projection. A few accessory rhinaria adjacent to the subapical primary rhinarium. Processus terminalis with five setae, the apical seta length [n = 6, 0.046 – 0.056 (0.051) mm] approximately 3 – 4 x longer than the rest. Rostrum reaching past hind coxae, with ultimate segment length [n = 7, 0.033 – 0.040 (0.037) mm] subequal to width of segment III at midpoint [n = 7, 0.036 – 0.045 (0.042) mm]. Stylet length: n = 6, 1.314 – 1.728 (1.579) mm, approximately 3.9 x body length. Coxae, femorotrochanters, and tibiae smooth, second tarsal segments weakly imbricated. Pair of long dorso-subapical setae on foreleg. Pairs of subequal dorso-subapical setae on mid- and hindlegs. Wax glands anterior to coxae lacking. Hind femorotrochanter length: n = 7, 0.045 – 0.057 (0.049) mm. Pronotum with one antero-lateral and sometimes one postero-lateral wax gland on each plate. Mesonotum with one lateral wax gland. Metanotum with one lateral wax gland, usually with fewer facets than on pro- and mesonotum, sometimes lacking. Raised ventral spiracles on meso- and metanotum bearing facets on basal surface. Abdominal wax glands lacking. Abdominal spiracles minute, hardly visible. Acuminate ventral setae in lateral, pleural, and spinal positions on abdominal segments I – VI, in lateral and spinal positions on abdominal segment VII, in a lateral position on abdominal segment VIII, and a pair of long setae on the apex of abdominal segment IX. Abdominal setae not always apparent. Neoprogrediens (Fig. 2 B – C). Body length: n = 12, 0.415 – 0.482 (0.451) mm. Body width 0.244 – 0.295 (0.265) mm. Body lacking wax glands. Head and pronotum with very lightly sclerotized plates, separated from each other and along mid-line by narrow sutures. Each dorsal cephalic plate with six setae, one postero-spinal, one postero-lateral, one antero-spinal, two antero-lateral, one apical. A pair of ventro-spinal setae between antennae. Antennae 3 - segmented, segment III imbricated, length [n = 13, range 0.044 – 0.062 (0.052) mm] approximately 3.1 x its width at midpoint [n = 13, 0.014 – 0.020 (0.017) mm], and subequal to segments I [n = 13, range 0.019 – 0.030 (0.024) mm] and II [n = 13, range 0.022 – 0.031 (0.026) mm] combined. Two primary rhinaria on segment III, one subapical and one approximately ½ the segment length from the apex, each with a small thumb-like projection. A few accessory rhinaria adjacent to the subapical primary rhinarium. Processus terminalis with five setae, the apical seta length [n = 12, 0.038 – 0.051 (0.044) mm] approximately 3 x longer than the rest. Rostrum reaching just anterior to hind coxae. Stylets short [n = 13, 0.411 – 0.680 (0.477) mm], approximately 1.0 x body length. Coxae, femorotrochanters, and tibiae smooth, second tarsal segments imbricated. Pair of dorso-subapical setae on foreleg, one capitate seta approximately 2 x longer than the other acuminate one. Pairs of subequal dorso-subapical setae of mid- and hindlegs. Wax glands anterior to coxae lacking. Hind femorotrochanter length: n = 13, 0.053 – 0.065 (0.058) mm. Prominent, raised ventral spiracles on the meso- and metanotum bearing facets on basal surface (Fig. 2 C). Abdominal spiracles not apparent. Acuminate ventral setae in lateral, pleural, and spinal positions on all thoracic segments and abdominal segments I – VI, in lateral and spinal positions on abdominal segment VII, in a lateral position on abdominal segment VIII, and a pair of long setae on the apex of abdominal segment IX. Adult exulis (Fig. 2 F – J). Body length: n = 21, 0.631 – 1.044 (0.802) mm. Body width: n = 21, 0.579 – 0.877 (0.752) mm. Pronotum width: n = 21, 0.484 – 0.698 (0.600) mm. Body shape transverse oval to nearly round. Dorsal wax glands generally with one to approximately 20 contiguous or nearly contiguous small round pores. Head and prothorax strongly sclerotized on dorsal and ventral surfaces. Dorsal plates of head and pronotum completely fused into cephaloprothoracic shield with distinctly wrinkled surface. Anterior half of cephaloprothoracic shield bearing numerous small wax glands with usually less than 10 thick-walled, round to ovoid, tightly packed facets (Fig. 2 J). Wax glands on both dorsal and ventral surfaces of cephaloprothoracic shield. Wax glands less numerous (often absent), smaller, and with fewer facets on posterior half of cephaloprothoracic shield. The largest wax facets on the head approximately half the diameter of ocellus. Antennae greatly reduced, appearing as a narrow ridge with no apparent segmentation (Fig. 2 H). Rostrum with ultimate segment length [n = 13, 0.031 – 0.042 (0.038) mm] approximately 0.7 x width of segment III at midpoint [n = 13, 0.047 – 0.059 (0.052) mm]. Vestigial legs present as slightly raised cuticular protrusions on pro-, meso-, and metanotum with clusters of approximately 8 – 15 setae (Fig. 2 G). Vestigial legs on meso-, and metanotum unsclerotized. Anterior base of vestigial legs with 0 – 5, usually 1 – 2, small wax glands with one to approximately 12 round facets. Meso- and metanotum, and abdominal segments I – II usually with 1 – 9 small lateral wax glands with approximately 1 – 20 facets each. Mesonotum with pleural wax glands, or an irregular transverse row of 0 – 4 small wax glands consisting of 2 – 8 facets. Prominent, strongly sclerotized and raised ventral spiracles on meso- and metanotum. Rarely, a single small lateral wax gland with a 1 – 14 minute facets on some of abdominal segments III – VII. Minute unsclerotized spiracles on abdominal segments II – VI (Fig. 2 I), often not apparent but sometimes recognizable if internal trachea leading to them are preserved in the specimen. Acuminate setae abundant on abdominal segment IX. Ovipositor short, with length of gonapophyses [n = 10, 0.029 – 0.059 (0.043) mm] approximately 0.9 x the width across apodemes [n = 10, 0.034 – 0.061 (0.044) mm].	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	biology_ecology	Host and Distribution. Larix griffithii in Bhutan (Mongar and Bumthang) (Fig. 3).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1378FFD80AD4FA119E290AF7.taxon	discussion	Remarks. Adelges (Ca.) coccipus feeds at the base of the buds and needle whorls, and on the bark on twigs of Larix griffithii. Similar to many other adelgid species, A. (Ca.) coccipus has morphologically dissimilar sistens and progrediens forms in the first instar, with sistentes having stronger sclerotization and longer rostral stylets (Havill & Foottit 2007). All but one of the first instars collected on March 27 were sistens forms, while all the first instars collected on April 5 – 7 were progrediens forms. The neosistentes on the earlier date were likely the remnants of the overwintering generation that had not developed because all of the specimens in our possession were damaged to some degree. The progredientes collected later were alive and abundant. This could indicate the presence of a longer generation of overwintering sistens individuals with an intervening series of short generations of progredientes individuals. We did not observe a dichotomy among the adults that would indicate morphologically distinct sistens and progrediens, nor did we collect sexupara adults into which a proportion of progrediens nymphs often develop in other species. In addition, their DNA sequences did not match any specimens on Picea, so there is no indication at this point that A. (Ca.) coccipus can complete a holocycle. This is the first known legless species in the Aphidomorpha. The nymph illustrated in Fig. 2 D – E was confirmed as a second instar because it was found emerging from its first instar exuvium. This confirms that all but the first instar lacks legs and have reduced antennae (Fig. 2 E). We are not aware of any other completely legless species within Aphidomorpha, but some species, scattered throughout Aphididae have reduced leg structures. For example, Colopha hispanica Nieto Nafría & Mier Durante, 1987 (Aphididae: Eriosomatinae), some Quadrartus species (Aphididae: Hormaphidinae) (Takahashi 1958; Sorin 2001), Mastopoda pteridis Oestlund, 1886 (Aphidinae: Macrosiphini), and Atarsos grindeliae Gillette, 1911 (Aphidinae: Macrosiphini) all have shortened legs with some segments missing or fused. Being nested well into the Adelgidae, A. (Ca.) coccipus represents an independent evolution of the apodous trait commonly found in scale insects (Hemiptera: Sternorrhyncha: Coccomorpha). In scale insects, legs can be entirely missing, or represented by a variety of reduced morphologies (Gullan & Kosztarab 1997). The morphology of A. (Ca.) coccipus leg vestiges as clusters of setae is similar to some species of flat grass scales (Hemiptera: Acleridae; McConnell 1953) and legless mealybugs (Hemiptera: Pseudococcidae: Pseudococcinae: Serrolecanini; Hendricks & Kosztarab 1999). The phylogeny (Fig. 1) places A. (Ca.) coccipus sister to an undescribed species from China that will be dealt with in a subsequent publication when more specimens are available for a full description. This other species has all of the features of Cassiadelges, but unlike A. (Ca.) coccipus, it has fully developed legs. Comparison between Adelges (Ca.) coccipus and A. (Ca.) nr. coccipus presents an exciting opportunity to examine the evolutionary development of insect limb formation from a new perspective.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	description	(Fig. 4 A – E)	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	etymology	Etymology. From Changaphu, the locality in Bhutan where spruce trees were found to be heavily infested by this species.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	materials_examined	Type material. HOLOTYPE: 1 adult fundatrix, BHUTAN, Thimphu, Changaphu Forest Nursery, lat. 27.4997, long. 89.7176, 2 - IV- 2020, on Picea spinulosa, Kaka Tshering (USNM). PARATYPES: 1 first instar fundatrix, 1 adult fundatrix, same data (NBC). 1 first instar fundatrix, 1 nymph fundatrix, same data (USNM). 1 first instar fundatrix, same data (YPM # ENT 594814). 1 nymph fundatrix, same data (YPM # ENT 594815). 1 adult fundatrix, same data (YPM # ENT 594816). 2 adult gallicolae, BHUTAN, Thimphu, Changaphu, lat. 27.5022, long. 89.7156, 29 - VII- 2021, on Picea spinulosa, Kaka Tshering (CNC # 5338320 - 6, CNC # 5338321 - 9). 1 adult gallicola, same data (NBC). 1 adult gallicola, same data (USNM). 1 adult gallicola, same data (YPM # ENT 594826). Additional material examined. BHUTAN: Bumthang: Khandopang: 2 adult fundatrices, 3 fundatrix exuviae (YPM), 27.4262, 90.9063, 8 - IV- 2022, on Picea spinulosa, Kaka Tshering. Thimphu: Changaphu Forest Nursery: 11 late instar nymph fundatrices (CNC), 1 first instar fundatrix exuvium, 4 late instar nymph fundatrices, 2 adult fundatrices (USNM), 1 first instar fundatrix exuvium, 5 late instar nymph fundatrices, 3 adult fundatrices (YPM), lat. 27.4997, long. 89.7176, 2 - IV- 2020, on Picea spinulosa, Kaka Tshering. 9 fourth instar nymph gallicolae, 17 adult gallicolae (CNC), 1 adult gallicola (NBC), 2 late instar nymph gallicolae, 5 adult gallicolae (USNM), 1 nymph gallicola, 15 adult gallicolae (YPM), lat. 27.4997, long. 89.7176, 29 - VII- 2021, on Picea spinulosa, Kaka Tshering. 16 first instar gallicolae (CNC), 15 galls (YPM), lat. 27.4997, long. 89.7176, 22 - V- 2022 on Picea spinulosa, Kaka Tshering.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	diagnosis	Diagnosis. The neofundatrix of A. (Ch.) changaphuensis (Fig. 4 A) is similar to the neosistens of A. (Ch.) viridanus (Börner 1909; Inouye 1945) in having a wide spinal fissure bisecting the cephalic and prothoracic plates, spinal, pleural, and lateral wax glands on the meso- and metathoracic and first six abdominal segments, and wax glands with facets surrounding a central seta. The A. (Ch.) changaphuensis neofundatrix differs from the A. (Ch.) viridanus neosistens in having separate longitudinal lateral prothoracic plates and differs from all other members of Cholodkovskya in having larger lateral abdominal wax glands arranged around two setae instead of around one seta. The neofundatrix of A. (Ch.) viridanus has not been described to allow direct comparison with the same morph in A. (Ch.) changaphuensis. The general appearance of the wax glands is also similar to neosistentes and neofundatrices in species of Sacchiphantes. The adult fundatrix of A. (Ch.) changaphuensis (Fig. 4 C) is similar to that of A. (Ch.) viridanus (Havill et al. 2023) but A. (Ch.) changaphuensis lacks wax glands antero-lateral to forecoxae. Late instar nymphs (Fig. 4 B) are similar in morphology to the adult but lack an ovipositor. The adult gallicola of A. (Ch.) changaphuensis (Fig. 4 D) differs from A. (Ch.) viridanus because it does not have anterior and posterior wax glands on each side of the head united. Unlike the gallicolae of other Adelges, the medial vein on the hindwings of A. (Ch.) changaphuensis are either faint or lacking entirely. Also note, that the sexuparae of Adelges (Gilletteella) cooleyi (Gillette, 1907) and Adelges (Dreyfusia) nordmannianae (Eckstein, 1890) lack the medial vein on the hindwing, so A. (Ch.) changaphuensis is not the only alate Adelges with this character.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	description	Description. Neofundatrix (Fig. 4 A). Body length: n = 4, 0.604 – 0.722 (0.665) mm. Body width: n = 4, 0.374 – 0.437 (0.404) mm. Dorsal wax glands generally on strongly sclerotized plates, and with 3 – 6 round to ovoid facets with a granular appearance, loosely packed and arranged around a central stout seta, sometimes merged with neighboring glands to include up to approximately 20 facets arranged around two or three setae. Head and pronotum with strongly sclerotized spinal and lateral plates, separated from each other and along mid-line by wide fissures. Ocelli borne on small lateral plates. Each spinal cephalic plate with five large wax glands, three apical and two postero-lateral, a sixth slightly raised postero-spinal area with a central seta, usually lacking facets, occasionally with one or several facets. The largest dorsal wax gland facets on the head are slightly smaller than ocellus. Large ventral wax glands on moderately sclerotized sternites posterior to base of antennae with approximately 30 small, round, tightly packed facets. Antennae 3 - segmented, segment III distinctly imbricated, very long and slender [n = 6, 0.125 – 0.132 (0.129) mm], 7.3 x its width at midpoint [n = 6, 0.017 – 0.019 (0.018) mm], approximately 2.3 x longer than segments I [n = 4, 0.019 – 0.034 (0.025) mm] and II [n = 4, 0.020 – 0.033 (0.027) mm] combined. Two primary rhinaria on segment III, one subapical and one approximately 1 / 3 the segment length from the apex, each with a small thumb-like projection. A few accessory rhinaria adjacent to the subapical primary rhinarium. Processus terminalis with five setae, the apical seta length [n = 4, 0.035 – 0.041 (0.038) mm] approximately 2 x the rest. Rostrum reaching past hind coxae, with ultimate segment as long [n = 5, 0.053 – 0.061 (0.057) mm] as width of segment III at midpoint [n = 4, 0.051 – 0.056 (0.054) mm]. Stylets very long [n = 2, 3.353 – 3.467 (3.410) mm], approximately 5.1 x body length. Coxae and femorotrochanters smooth, with tibiae weakly imbricated, and second tarsal segments distinctly imbricated. Pair of dorso-subapical capitate setae of fore- and midlegs with one of the pair approximately 2 x as long as the other, subequal on hindleg. Mid- and hind-coxae with basal wax glands with approximately 8 – 12 moderately packed round to ovoid facets. Hind femorotrochanter length: n = 5, 0.096 – 0.105 (0.101) mm. Pronotum with wax glands on the four corners of each spinal plate, each gland with 1 – 4 facets. Lateral plates of pronotum extended longitudinally with two wax glands, each with approximately 5 – 12 facets. Meso- and metanotum with dorsal spinal, pleural, and lateral plates, each with a single wax gland. Spinal and pleural wax glands round and arranged around one seta, lateral wax glands oval-shaped along longitudinal axis usually arranged around two setae. Abdominal segments I – VI with dorsal spinal, pleural, and lateral plates, segment VII with spinal and lateral plates, and segment VIII with lateral plates. Abdominal segment IX with a transversely extended plate in central position. Each abdominal plate with a single wax gland. Lateral wax glands of abdominal segments I – VIII transverse oval-shaped with 7 – 20 facets usually arranged around two setae. Pleural and spinal wax glands of abdominal segments I – VI and spinal glands of segment VII round to oval with 3 – 7 facets arranged around a single seta. Ventral spiracles on abdominal segments II – VI. All abdominal segments with a ventro-spinal pair of fine acuminate setae, those on segment IX especially long and slender. Adult fundatrix (Fig. 4 C). Body length: n = 6, 1.343 – 1.903 (1.644) mm. Body width: n = 6, 1.042 – 1.226 (1.111) mm. Color in life light green. Dorsal wax glands generally on very lightly sclerotized plates, and with small round, tightly packed facets with borders only slightly darker than the surrounding cuticle. Sclerotized plates on head and pronotum lacking. Head with 8 – 10 small anterior wax glands between ocelli, each gland with approximately 3 – 20 loosely packed facets, the largest of which are approximately half the diameter of the ocellus. Ventral wax glands at base of antennae with approximately 30 – 40 very small tightly packed facets. Antennae 3 - segmented, segment III [n = 6, 0.074 – 0.097 (0.083) mm], approximately 3.6 x its width at midpoint [n = 6, 0.019 – 0.026 (0.023) mm], longer than segments I [n = 6, 0.024 – 0.037 (0.031) mm], and II [n = 6, 0.027 – 0.033 (0.032) mm] combined. Rostrum reaching to hind coxae, with ultimate segment length [(n = 4, 0.068 – 0.091 (0.079) mm] approximately 0.9 x width of segment III at midpoint [n = 4, 0.076 – 0.099 (0.084) mm]. Coxae, femorotrochanters, and tibiae smooth, second tarsal segments weakly imbricated. Mid- and hind-coxae with transversely oval wax glands bearing 10 – 25 facets. Hind femorotrochanter length: n = 6, 0.125 – 0.224 (0.171) mm. All thoracic segments and abdominal segments I – VI with spinal, pleural, and lateral wax glands, abdominal segment VII with lateral and spinal wax glands, segment VIII with lateral wax glands. Spiracles on abdominal segments II – VI strongly sclerotized and conspicuous. Numerous acuminate setae on abdominal segments VIII and IX. Ovipositor with length of gonapophyses [n = 4, 0.117 – 0.157 (0.130) mm] approximately 1.2 x the width across apodemes. Adult gallicola (Fig. 4 D). Body length: n = 16, 1.772 – 2.610 (2.058) mm. Pronotum width: n = 15, 0.587 – 0.812 (0.676) mm. Abdomen width: n = 16, 0.839 – 1.221 (0.960) mm. Color in life light green when newly emerged. Dorsal wax glands generally with round, tightly packed facets interspersed with minute setae. Head with a pair of anterior and a pair of posterior irregular-shaped wax glands with approximately 10 – 30 facets. Posterior glands may be subdivided into small groups of facets around setae. Antenna distinctly five segmented, n = 12, 0.336 – 0.434 (0.381) mm long, segments II – V imbricated. Length of antennal segment III: n = 12, 0.075 – 0.088 (0.080) mm, IV: n = 12, 0.080 – 0.109 (0.093) mm, n = 12, V: 0.101 – 0.137 (0.115) mm. Ratio of length of antennal segments III: IV: V 1.00: 1.16: 1.43. Antennal segment III tapering sharply towards the base, segments IV and V with a gradual taper below rhinarium. Rhinaria on antennal segments III – V oval, subapical, often with sinuate margins, sometimes appearing triangular in profile. Rhinarium occupying approximately 1 / 3 of each segment: III: n = 10, 0.25 – 0.41 (0.34), IV: n = 10, 0.25 – 0.40 (0.33), V: n = 10, 0.30 – 0.40 (0.34). Processus terminalis with four or five acuminate setae, with apical seta length [n = 10, 0.015 – 0.022 (0.017) mm] approximately 1.9 x as long as the rest. Ultimate segment of rostrum length [(n = 14, 0.055 – 0.080 (0.064) mm] 0.9 x the width of segment III at midpoint [n = 14, 0.058 – 0.083 (0.072) mm]. Coxae and femorotrochanters smooth, tibiae very weakly imbricated, and second tarsal segments distinctly imbricated. Hind femur with approximately 15 – 20 long fine acuminate setae along ventral surface. Length of longest setae on hind femur: n = 15, 0.028 – 0.039 (0.034) mm. Hind femorotrochanter length: n = 15, 0.314 – 0.408 (0.350) mm. Each side of pronotum with several small, antero-lateral oval-shaped glands with 2 – 50 facets, a very large postero-lateral band of approximately 120 – 150 facets, and a narrow irregular row, 2 – 5 facets thick, along the posterior edge. Mesonotum with a pair of spinal oval-shaped wax glands bearing approximately 50 – 60 facets between mesothoracic lobes. Metanotum with a pair of wax glands with approximately 30 – 60 facets towards the posterior half of the spinal lobes. Forewing length: n = 16, 2.286 – 3.023 (2.661) mm. Hindwing length: n = 16, 1.377 – 1.892 (1.588) mm. Forewing with medial, cubital, and anal veins straight or only slightly curved back towards the body, leaving the radial sector at acute angles. Hindwing with medial vein faint or lacking. Abdomen with spinal, pleural, and lateral wax glands on segments I – VI, lateral and pleural wax glands on segment VII, only lateral wax glands on VIII, and a post-anal gland on IX. Lateral glands on abdominal segments I – VII transverse, often crescent-shaped. Spinal glands decrease in size posteriorly. Distinct sclerotized spiracles on abdominal segments II – VI. Abdominal segments VII – VIII with numerous ventral fine acuminate setae, some long and slender. Ovipositor with gonapophyses [n = 11, 0.106 – 0.126 (0.114) mm] approximately 1.3 x longer than width across apodemes [n = 11, 0.072 – 0.101 (0.086) mm]. Gall (Fig. 4 E). Light green in color, strobile-shaped. Usually occurring singly per node on terminal and lateral buds of new shoots. Consists of swollen needle bases, fused at the axis of the shoot. Chambers internally unconnected. Gall length [n = 9, 21 – 34 (24.6) mm] approximately 1.9 x the width (n = 9, 10 – 16 (12.7) mm]. Usually without a distal shoot, but occasionally with a distal tuft of shortened needles or short shoot. Distal ends of the needles on gall narrow and sharp, extending approximately 2 – 5 mm from the surface.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	biology_ecology	Hosts and distribution. Picea spinulosa in Bhutan (Bumthang, Thimphu) (Fig. 5).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF137CFFC40AD4FB209C330CBD.taxon	discussion	Remarks. The morphological similarities to A. (Ch.) viridanus (Börner, 1909), and the sister relationship with A. (Ch.) viridanus in the phylogeny (Fig. 1) supports the placement of A. (Ch.) changaphuensis in subgenus Cholodkovskya. The life cycle of A. (Ch.) changaphuensis cannot be determined from our collections. There is no evidence of host alternation because our survey did not yield specimens with matching DNA barcodes from potential secondary hosts. Galls collected on May 22, 2022 were mature and close to opening. This species could be anholocyclic on Picea, or since in the molecular phylogeny (Fig. 1) it is nested well within a clade of species that use Larix species as secondary hosts, it may alternate between Picea and Larix. The other members of Cholodkovskya, A. (Ch.) viridanus, Adelges (Cholodkovskya) viridulus (Cholodkovsky, 1911), and A. (Ch.) oregonensis, all feed on species of Larix. Adelges (Ch.) viridanus has an anholocyclic life cycle in Europe and China with a generation of wingless sistentes that settle on the bark, followed by a series of generations of alates that settle on the needles (Cholodkovsky 1911; Li & Tsai 1973). In Japan, the fundatrix adult and gallicola nymph of A. (Ch.) v iridanus were recently described, indicating that in that region the species can either be holocyclic, alternating between Larix and Picea, or includes recently diverged anholocyclic populations on Picea (Havill et al. 2023). The life cycle of A. (Ch.) v iridulus is anholocyclic on Larix but has only one annual generation of sistens on the bark and no known alate forms. The life cycle of A. (Ch.) oregonensis is unknown, as the species is only known from wingless forms that feed on the needles and the base of needles of Larix in western North America (Annand 1928). Note that A. (Ch.) oregonensis is not sister to A. (Ch.) viridanus or A. (Ch.) changaphuensis in the molecular phylogeny (Fig. 1), so it may not belong in this subgenus. A more detailed morphological analysis of the adelgid species feeding on Larix, and a more robust phylogeny of the family could help to resolve a more informative delimitation of Cholodkovskya.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	description	(Fig. 6 A – C)	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	etymology	Etymology. Genitive derived from its host plant, Abies densa.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	materials_examined	Type material. HOLOTYPE: 1 adult exulis, BHUTAN, Bumthang, Tangsibi, lat. 27.4972, long. 90.8699, 6 - IV- 2022, on Abies densa, Kaka Tshering (USNM). PARATYPES: 1 first instar exulis (CNC # 5338332 - 12), same data. 2 late instar nymph exules, same data (NBC). 1 first instar exulis, 2 late instar nymph exules, same data (USNM). 1 first instar exulis, same data (YPM # ENT 594781). 2 late instar nymph exules, same data (YPM # ENT 594788). Additional material examined. BHUTAN: Bumthang: Tangsibi: 2 first instar exules, 11 late instar nymph exules (CNC), 1 first instar exulis, 2 first instar exulis exuviae, 10 late instar nymph exules (NBC), 4 first instar exules, 6 first instar exulis exuviae, 2 late instar nymph exules (USNM), 8 first instar exules, 17 first instar exulis exuviae, 14 late instar nymph exules, 2 late instar nymph exulis exuviae (YPM), lat. 27.4972, long. 90.8699, 6 - IV- 2022, on Abies densa, Kaka Tshering.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	diagnosis	Diagnosis. The first instar exulis of A. (D.) densae (Fig. 6 A) is similar to Adelges (Aphrastasia) pectinatae (Cholodkovsky, 1888), A. (Ap.) pindrowi Yaseen & Ghani, 1971, A. (D.) nordmannianae, and Adelges (D.) piceae (Ratzeburg, 1844) in having compound wax glands with emarginated clusters of many small facets. It differs from these species in the structure of the wax glands, and that the first instar exulis has wax glands in pleural and lateral positions. Late instar exulis nymphs (Fig. 6 B) are similar to the adult (Fig. 6 C) but lack an ovipositor.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	description	Description. First instar exulis. (Fig. 6 A). Body length: n = 10, 0.431 – 0.588 (0.494) mm. Body width: n = 11, 0.271 – 0.457 (0.335) mm. Dorsal wax glands generally on strongly sclerotized plates. Glands consist of 2 – 6 round, oval, or polygonal delineated areas arranged around a central seta, each area containing approximately 6 – 50 very small round, oval, or polygonal facets, sometimes glands merged, arranged around two or more setae. Head and pronotum with moderately to strongly sclerotized and reticulated spinal and lateral plates, separated from each other and along mid-line by wide fissures. Ocelli borne on separate lateral plates. Each spinal cephalic plate with six large wax glands, two spinal, two lateral, two apical, together filling most of the area of the plate. Ventral wax glands at base of antennae lacking. A pair of spinal flagellate setae on head slightly less than half the distance from apex. Antennae 3 - segmented, segment III imbricated, length [n = 13, 0.076 – 0.089 (0.083) mm], approximately 5.2 x its width at midpoint [n = 13, 0.013 – 0.020 (0.016) mm], and approximately 2.1 x longer than segments I [n = 12, 0.016 – 0.028 (0.021) mm] and II [n = 12, 0.017 – 0.022 (0.019) mm] combined. Two primary rhinaria on segment III, one subapical and one approximately ½ the segment length from the apex, each with a small thumb-like projection. Several accessory rhinaria adjacent to subapical primary rhinarium. Processus terminalis with five setae, the apical seta length [n = 5, 0.033 – 0.037 (0.034) mm] approximately 3.0 x longer than the rest. Rostrum reaching hind coxae, with ultimate segment 0.7 x as long [n = 11, 0.033 – 0.038 (0.036) mm] as width of segment III at midpoint [n = 11, 0.045 – 0.052 (0.050) mm]. Stylet length [n = 4, 1.811 – 1.959 (1.888) mm] approximately 3.8 x body length. Coxae and femorotrochanters smooth, with tibiae and second tarsal segments weakly imbricated. Pair of dorso-subapical capitate setae, subequal in length, on tarsal segment II of fore-, mid- and hindlegs. Wax glands antero-lateral to mid-coxae with approximately 7 – 12 round to oval facets. Hind femorotrochanter length: n = 13, 0.056 – 0.068 (0.063) mm. Pronotum with wax glands on the four corners of each spinal plate, each gland with 4 – 6 facet areas. Lateral prothoracic plates longitudinal oval-shaped, each with 2 wax glands, each gland with 5 – 6 facet areas. Meso- and metathorax with separate lateral, pleural, and spinal plates, lateral plates with wax gland with 5 – 6 facet clusters arranged around 1 – 2 setae, pleural and spinal plates each with a single wax gland with 4 – 5 facet clusters arranged around one seta. Each abdominal plate generally with a single wax gland with 2 – 4 faceted areas surrounding a seta. Abdominal segments I – V with separate lateral, pleural, and spinal wax plates, segment VI – VII with lateral and spinal plates, spinal plates touching or fused at the midline, segment VIII with very small lateral plates lacking wax glands, segment IX with apical plate lacking wax glands. Abdominal segments VII – IX tend to collapse telescopically when slide mounted, making these segments easier to see in exuviae. Lateral abdominal plates ovoid to square, pleural and spinal plates transverse oval. Abdominal segment IX with a pair of flagellate setae at the tip. Adult exulis. (Fig. 6 C). Body length: n = 1, 1.305 mm. Body width: n = 1, 0.770 mm. Moderately to strongly sclerotized dorsal plates generally bearing large, strongly sclerotized hemispheroid protuberant spinal and pleural plates with 1 – 3 stout setae on the surface and bearing no apparent wax glands. Head and pronotum with lightly sclerotized large spinal and small lateral plates, separated from each other and along mid-line by fissures. Ocelli borne on lateral plates. Each spinal cephalic plate with three protuberances: two posterior and one apical. Antennae 3 - segmented, segment III (n = 1, 0.060 mm), approximately 1.3 x longer than segments I (n = 1, 0.028 mm) and II (n = 1, 0.019 mm) combined. Segment III with a subapical primary rhinarium with a few adjacent accessory rhinaria. Processus terminalis with 3 stout setae. Rostrum reaching past mid coxae, with ultimate segment as long (n = 1, 0.062 mm) as 0.7 x the width of segment III at its midpoint (n = 1, 0.080 mm). Coxae and femorotrochanters, and tibiae smooth, distitarsi weakly imbricated. Wax glands with approximately 20 – 40 loosely packed facets antero-lateral to all coxae. Hind femorotrochanter length: n = 1, 0.096 mm. Pronotum with protuberant plates on the four corners of each spinal plate. Meso- and metanotum and abdominal segments I – IV with separate lateral, pleural, and spinal wax plates, each borne on a large protuberance. Abdominal segments V – VIII with only lateral plates with the protuberance becoming less pronounced and less sclerotized posteriorly. Abdominal segment IX with a lightly sclerotized transversely extended post-anal plate with a pair of long acuminate setae. Very lightly sclerotized sternites with wax glands with 4 – 12 loosely packed facets and 3 – 4 setae mesad to the spiracles on abdominal segments IV – VI. Acuminate setae abundant ventrally on segment VIII and dorsally on segment IX, and in a single spinal pair on each of the other abdominal segments. Ovipositor with length of gonapophyses (n = 1, 0.125 mm) 1.2 x width across apodemes (n = 1, 0.108 mm).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	biology_ecology	Hosts and Distribution. Abies densa in Bhutan (Bumthang) (Fig. 7).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1360FFC60AD4FCEE9EFF09A6.taxon	discussion	Remarks. This species was collected from Abies densa in densely congregated colonies on the bark of the main stem and on the underside of lateral branches on twigs. On the twigs, the stylets were inserted mostly into the bark, and occasionally into the base of needles. We tentatively place A. (D.) densae in Dreyfusia because it is sister to A. (D.) knucheli in the molecular phylogeny (Fig. 1). It has some morphological similarities to A. (D.) pindrowi, and both A. (D.). pindrowi and A. (D.) densae have morphological similarities to A. (Ap.) pectinatae, which with its two subspecies, is the only member of Aphrastasia Börner, 1909. Adelges (D.). pindrowi was placed in Dreyfusia by Havill & Foottit (2007) and Favret et al. (2015 b). However, Yaseen & Ghani (1971) remarked that A. (D.) pindrowi closely resembles Adelges (Aphrastasia) pectinatae ishiharai (Inouye, 1936), and Blackman & Eastop (2024) also remarked that A. (D.) pindrowi is “ in the pectinatae group. ” It might be appropriate to place A. (D.). pindrowi and A. (D.) densae in Aphrastasia; however A. (D.) densae is not sister to A. (Ap.) pectinatae in our molecular phylogeny. A more detailed morphological analysis of the adelgid species feeding on Abies, and a more robust phylogeny of the family could help to resolve a more informative delimitation of Aphrastasia. It is possible that either A. (D.) densae or A. (G.) glandulae is the same as what Schmutzenhofer (1985) recorded as “ Chermes abietis-piceae ” on Abies densa at Thumsingla (= Phrumsengla). In our survey, both A. (D.) densae and A. (G.) glandulae were collected from Abies densa within 16 km of this locality. Chermes abietispiceae was originally described from northern India, alternating between Picea smithiana and Abies spectabilis (Stebbing 1903 a, b). It was later renamed Chermes himalayensis by Stebbing (1910) and given a more detailed description, but this name is an unavailable synonym of the earlier name. Chermes abietispiceae is currently considered a nomen dubium (Favret 2024) because neither of Stebbing’s descriptions provide enough information about the appearance and arrangement of the wax glands to confirm its identity (Schneider-Orelli & Schneider 1954). Our sample of A. (D.) densae was collected from the stem, not from the needles as described by Schmutzenhofer (1985), who observed their feeding causing a downward twisting of the needles. However, it is possible that, similarly to A. (D.) nordmannianae (Pschorn-Walcher & Zwölfer 1958), A. (D.) densae includes progredientes that feed on the needles that we did not sample. Ghani & Rao (1966) also speculated that A. (D.) knucheli could be the same as Chermes abietispiceae. Additional sampling in the region could resolve this.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	description	(Fig. 8 A – D)	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	etymology	Etymology. Invariant combination of the Dzongkha noun, lep, meaning “ brain ” and Psimon the supervillain from the DC Comics Universe, whose brain is visible through a clear dome on his head. Both are in reference to the large wax glands on the head, that superficially resemble the lobes of a vertebrate brain.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	materials_examined	Type material. HOLOTYPE: 1 adult exulis, BHUTAN, Mongar, Sengor, lat. 27.3394, long. 91.0295, 8 - IV- 2022, on Tsuga dumosa, Kaka Tshering (USNM). PARATYPES: 1 first instar exulis exuvium, same data (CNC # 5338329 - 10). 1 adult exulis, same data (CNC # 5338329 - 7). 1 late instar nymph exulis, 1 adult exulis, same data (NBC). 1 first instar exulis, 2 first instar nymph exulis exuviae, 1 late instar exulis, same data (USNM). 1 late instar exulis, same data (YPM # ENT 594636). 1 adult exulis, same data (YPM # ENT 594641). 1 first instar exulis exuvium, BHUTAN, Mongar, Sengor, lat. 27.3711, long. 91.0234, 7 - IV- 2022, on Tsuga dumosa, Kaka Tshering (NBC). 1 late instar nymph sexupara, same data (USNM). 2 first instar exulis exuviae, same data (YPM # ENT 594634). 1 first instar exulis, BHUTAN, Wangdue Phodrang, Dungdungnyesa, lat. 27.5377, long. 90.1851, 12 - XI- 2021, on Tsuga dumosa, Kaka Tshering (CNC # 5338322 - 9). 1 adult exulis, same data (CNC # 5338322 - 4). 3 first instar exules, 1 first instar exulis exuvium, same data (USNM). Additional material examined. BHUTAN: Mongar: Sengor: 2 late instar nymph exules (USNM), 1 first instar exulis, 6 first instar exulis exuviae, 10 late instar nymph exulis exuviae, 12 late instar nymph exules, 3 late instar nymph sexuparae (YPM), lat. 27.3711, long. 91.0234, 7 - IV- 2022, on Tsuga dumosa, Kaka Tshering. 2 first instar exules exuviae, 1 late instar nymph sexupara, 6 adult exules (CNC), 1 first instar exulis, 9 first instar exulis exuviae, 2 late instar nymph exulis exuviae, 9 late instar nymph exules (USNM), 1 first instar exulis, 16 first instar exulis exuviae, 12 late instar nymph exulis exuviae, 11 late instar nymph exules, 1 late instar nymph sexupara, 12 adult exules (YPM), lat. 27.3394, long. 91.0295, 8 - IV- 2022, on Tsuga dumosa, Kaka Tshering. Wangdue Phodrang: Dungdungnyesa: 1 egg, 3 first instar exules, 3 adult exules (CNC), 2 first instar exules, 1 adult exulis (NBC), 3 first instar exules, 2 first instar exulis exuviae, 1 late instar nymph exulis exuvium, 1 adult exulis (USNM), 4 first instar exules, 2 first instar exulis exuviae, 3 late instar nymph exulis exuviae, 6 adult exules (YPM), lat. 27.5377, long. 90.1851, 12 - XI- 2021, on Tsuga dumosa, Kaka Tshering.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	diagnosis	Diagnosis. Adelges (An.) lepsimon is placed in Annandina by phylogenetic relatedness (Fig. 1), and by the combination of characters outlined in Favret et al. (2015 a): feeding on Tsuga, having the first instar exulis with head and pronotum almost completely covered by plates, with margins lined with wax gland facets, four plates on the mesothorax, and six on the metathorax and first five abdominal segments, the adult with a completely fused cephaloprothoracic shield containing many small facets, and by having capitate setae on the ultimate abdominal segments (Annand 1924; Havill et al. 2022). In A. (An.) lepsimon, the spinal and pleural mesothoracic plates are not always fused, so the diagnosis for Annandina should be amended to specify four or six plates on this segment. The first instar exulis of A. (An.) lepsimon differs from A. (An.) tsugae by having round to oval wax glands proximal to the lateral rows of facets on the head and each thoracic segment, and abdominal segments I – VI with dorso-spinal plates each with a wax gland in the center with 1 – 4 facets. The adult exulis of A. (An.) lepsimon resembles A. (An.) tsugae, which includes a diversity of genetic lineages in different regions of Asia and North America (Havill et al. 2016). A future paper will address morphological differences among these lineages compared to A. (An.) lepsimon.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	description	Description. First instar exulis (Fig. 8 A – B). Body length: n = 10, 0.314 – 0.423 (0.355) mm. Pronotum width: n = 11, 0.171 – 0.211 (0.193) mm. Body width: n = 11, 0.189 – 0.271 (0.220) mm. Dorsal wax glands generally on strongly to moderately sclerotized plates, with facets either arranged as anterior, lateral, or spinal longitudinal lines of individual pores, or as clusters of two to approximately 13 tightly packed, often overlapping pores. Head and pronotum almost completely covered by moderately to strongly sclerotized plates, separated from each other and along mid-line by a narrow fissure. Each cephalic plate lined with a row of single wax gland facets along the lateral, anterior, and spinal margins. Facets along the spinal margin indistinct, often more visible in exuviae (Fig. 8 B) than intact nymphs (Fig. 8 A). Each cephalic plate with a prominent round to oval wax gland with approximately 6 – 13 tightly packed facets, proximal to anterior row of facets. Ventral wax glands at base of antennae with approximately 3 – 4 facets. Antenna 3 - segmented, segment III distinctly imbricated, length [n = 12, 0.055 – 0.062 (0.059) mm], approximately 3.8 x its width at midpoint [n = 12, 0.013 – 0.018 (0.015) mm], and approximately 1.6 x longer than segments I [n = 12, 0.017 – 0.022 (0.020) mm] and II [n = 12, 0.015 – 0.020 (0.018) mm] combined. Two primary rhinaria on segment III, one subapical and one approximately 1 / 2 the segment length from the apex, each with a small thumb-like projection. Several accessory rhinaria adjacent to subapical primary rhinarium. Processus terminalis with 5 setae, the apical seta length [n = 11, 0.028 – 0.040 (0.036) mm] approximately 3 x the rest. Rostrum reaching past hind coxae with ultimate segment 0.8 x as long [n = 9, 0.028 – 0.039 (0.034) mm] as width of segment III at midpoint [n = 9, 0.037 – 0.044 (0.041) mm]. Stylet length [n = 4, 0.630 – 0.773 (0.704) mm] approximately 2.0 x body length. Coxae smooth. Femorotrochanters smooth to weakly imbricated, tibiae and second tarsal segments weakly imbricated. Pair of dorso-subapical setae on foreleg, one capitate seta approximately 2 x longer than the other acuminate one. Pairs of subequal dorso-subapical capitate setae on mid- and hindlegs. Wax glands antero-lateral to mid- and hindcoxae with approximately 1 – 6 round to oval facets. Hind femorotrochanter length: n = 12, 0.046 – 0.054 (0.050) mm. Each thoracic segment lined with a row of single wax gland facets along the lateral, margin, and a round to oval anterior cluster of approximately 2 – 13 wax gland facets, just proximal to lateral row. Meso- and metanotum with spinal and pleural plates, each plate with a single seta and a single row of 1 – 5 indistinct spinal and 0 – 5 lateral wax gland facets. Spinal and pleural plates of the mesothorax often fused. Metathoracic spinal plate sometimes with a central cluster of 1 – 5 wax gland facets. Abdominal segments I – V with separate lateral, pleural, and spinal wax plates, segment VI – VII with lateral and spinal plates, segment VIII with lateral plates, segment IX lacking plates. Abdominal segments I – VI dorso-spinal plates each with wax gland in the center consisting of a cluster of 1 – 4 facets. Abdominal segments I – III with pleural plates usually lacking wax gland facets. Abdominal segments IV – V with pleural plates with 1 – 3 wax facets. Abdominal segments I – VI with lateral plates each with 1 – 3 lateral facets and a small proximal wax gland with 1 – 5 facets. Abdominal spiracles minute, hardly visible. Pairs of small, ventro-spinal acuminate setae on abdominal segments I – VIII. Pair of dorso-lateral setae on abdominal segment VIII. Abdominal segment IX with two short acuminate dorsal setae anterior to anus, two longer acuminate ventral setae, and two long capitate setae at apex. Adult exulis (Fig. 8 D). Body length: n = 13, 0.764 – 1.251 (0.980) mm. Pronotum width: n = 14, 0.402 – 0.626 (0.523) mm. Body width: n = 13, 0.502 – 0.883 (0.726) mm. Dorsal wax glands generally on moderately to strongly sclerotized plates with numerous, up to more than 200, contiguous to nearly contiguous small round granular facets. Head and pronotum with sclerotized plates completely fused into cephaloprothoracic shield with largely smooth surface that extends ventrally to surround the base of antennae. Head with very large anterior wax glands with a lateral area mesad to ocelli curling towards, and sometimes touching posterior cephalic wax glands. Pair of cephalic postero-lateral, oval or irregularly shaped wax glands with approximately 25 – 50 facets. Largest diameter of facets on head subequal to the diameter of the ocellus. Large ventral wax glands on head with approximately 60 – 110 small round facets on surface at base of antennae. Antenna 3 - segmented, segment III [n = 13, 0.029 – 0.056 (0.041) mm] approximately 2.0 x its width at midpoint [n = 13, 0.016 – 0.022 (0.020) mm], approximately 0.9 x the length of segments I [n = 13, 0.016 – 0.033 (0.025) mm] and II [n = 13, 0.014 – 0.029 (0.020) mm] combined. Segment III with subapical primary rhinarium with a few adjacent accessory rhinaria. Processus terminalis with five or six stout setae. Rostrum reaching to anterior margin of hind coxae, with ultimate segment [n = 13, 0.041 – 0.091 (0.064) mm] 0.7 x the width of segment III at midpoint [n = 13, 0.050 – 0.076 (0.063) mm]. Coxae, femorotrochanters, and tibiae smooth, with second tarsal segment weakly imbricated. Large wax glands antero-lateral to coxae with approximately 50 – 100 loosely packed facets. Hind femorotrochanter length: n = 13, 0.064 – 0.144 (0.104) mm. Pronotum with very large lateral wax glands consisting of bands of up to 200 or more facets, and with posteropleural and postero-spinal irregularly shaped wax glands with 20 – 70 facets. Meso- and metathoracic segments and abdominal segments I – V with spinal, pleural, and lateral wax glands, circular or elliptical with numerous tightly packed facets. Plates on meso- and metanotum sometimes fused or touching. Abdominal segments VI – VII with lateral and spinal wax glands, segment VIII with lateral wax glands. Abdominal segment IX with large post-anal wax gland. Abdominal segments II – VI with distinct ventral spiracles. Abdominal segments VIII and IX with numerous capitate setae. Ovipositor with gonapophyses [n = 11, 0.075 – 0.115 (0.094) mm] approximately 1.4 x longer than width across apodemes [n = 11, 0.057 – 0.083 (0.067) mm].	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	biology_ecology	Hosts and Distribution. Tsuga dumosa in Bhutan (Mongar, Wangdue Phodrang) (Fig. 9).	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
6A7087EF1363FFC20AD4FF6F9EBE0E89.taxon	discussion	Remarks. Adelges (An.) lepsimon feeds between the needles on Tsuga dumosa twigs, in a similar fashion to A. (An.) tsugae, which feeds on T. dumosa and other Tsuga species (Havill et al. 2016). The two species were collected together at two sites on different trees, so distinguishing them in the field may be difficult. Adelges (An.) lepsimon was collected more rarely than A. (An.) tsugae, which was also collected in Bumthang, Paro, and Thimphu Districts. In addition to exules of A. (An.) lepsimon, we also collected sexupara nymphs with wing pads (Fig. 8 C), so it is likely that this species migrates to Picea where it completes a holocycle.	en	Havill, Nathan P., Brunet, Bryan, Zembrzuski, Zephyr, Tshering, Kaka (2025): Four new adelgid (Hemiptera: Adelgidae) species from Bhutan, including the first legless species of Aphidomorpha. Zootaxa 5583 (3): 437-461, DOI: 10.11646/zootaxa.5583.3.2, URL: https://doi.org/10.11646/zootaxa.5583.3.2
