AnimaliaHemipteraTriozidaePercyDiana M.Revision of the Hawaiian psyllid genus Swezeyana, with descriptions of seven new species (Hemiptera, Psylloidea, Triozidae)Zookeys155201820187587511310.3897/zookeys.758.23019 Swezeyana Caldwell, 1940 Swezeyana Caldwell, 1940: 389. Type species: Swezeyana elongagena Caldwell, 1940, by original designation.Description.

Adult. General colour variable ranging from pale yellow-brown, to green or yellow-green, to almost black; often with pink or reddish highlights on the fore wing as well as on the body, especially genal processes, legs, and abdomen. Fore wing membrane either with distinct darker patches or clouds of pigmentation, these range from dark brown to red, and in some cases are limited to termination of veins at wing margins and around cross veins between Rs and wing margin, if without distinct patterns of pigmentation, appearing uniformly clear, opaque yellow or fuscous; wing veins pale to red or dark brown, cross veins between Rs and ventral wing margin with or without pigmentation. Adult length including fore wing from 2–5 mm. Fore wing elongate and usually narrow (ratio WL:WW > 2.80, often > 3), acute to bluntly acute apically, either with trifurcation of veins R, M and Cu1, or with vein R branching anterior of bifurcation of M and Cu1; vein Rs long, reaching wing margin distad of M fork, but either with or without complete extension of Rs to wing margin, incomplete termination of Rs usually marked by pigmentation; vein R shorter than Cu1 and terminating at base of Rs, a pseudopterostigma is present between base of Rs and wing margin, and a more or less thickened wing margin (C+Sc) is present from the wing base to the pseudopterostigma, in some cases occupying part or entire area of cell c+sc; with or without one or more partial or complete cross veins traversing cell r1 between vein Rs and ventral wing margin; a single, broadly shaped marginal cluster of radular spines (Figs 3O, 6J) in cells cu1, m1, and m2; surface spinules either present in all cells, dense or sparsely distributed, or few to absent from c+sc, r2 and r1, often relatively sparsely distributed but becoming denser towards wing margin. Hind wing narrow and elongate (> 0.5 length of fore wing), clear or slightly fuscous in basal half. Head moderately deflexed downwards, vertex more or less flat dorsally, with lateral ocelli lying on small tubercles, medial epicranial suture distinct; genal processes extremely long, often upturned at apices, with scattered long setae and usually a single, long subapical seta on each process. Antennae short; antennal segments 10, either entirely dark, or more usually with terminal 3(-7) segments darker, or distal part of segments 3-8 darker; a single rhinarium apically on each of segments 4, 6, 8, 9; 1-2 long setae on each of segments 3-9, terminal segment with two unequal length apical setae. Distal proboscis segment short, darker apically. Thorax somewhat flattened to only moderately arched; vertex and thorax with scattered short to moderately long setae. Legs short, hind legs robust with femur longer than tibia; hind leg with meracanthus reduced to almost absent; metafemur with several stout setae apically; metatibia with or without distinct genual spine basally and typically with 1+2 (occasionally 1+3) sclerotized apical spurs; pro- and mesotarsi subequal in length, metatarsi unequal with extremely long basal tarsus slightly expanded with concave, ridged underside (Fig. 3M). Male terminalia with somewhat elongate subgenital plate; proctiger with pronounced posterior lobes medially, 1-2 long setae usually present on posterior apices of each lobe, length shorter, subequal or longer than paramere; paramere shape variable, generally broad basally and tapering to apex, with two stout setae on the interior apex (sometimes appearing as one from lateral view); distal aedeagus segment apex hooked. Female terminalia with medium to long dorsal and ventral setae; proctiger either truncate and markedly convex apically, with apex broad, blunt, bearing small medial cleft and fringed with stout setae, or dorsal surface more or less straight, apex tapering, lacking medial cleft and distinct fringe of setae; proctiger longer than subgenital plate; anal ring hour-glass shaped (with or without a head compartment at proximal end) and composed of a, usually, uninterrupted, double row of cells, posterior/distal portion of ring margin either smooth or convoluted; subgenital plate ventral surface either convex or more or less straight, apex terminating in a variably shaped beak often bearing a short or more pronounced medial cleft spanned by a short or extended membrane; ovipositor valves small, without serrations (Fig. 9O).

Egg. Known for four species. Pale or light brown, oblong-ovoid with a short, laterally positioned pedicel sub-basally on underside; distinctly hexagonal, honeycomb-like, sculpturing, to semi-hexagonal or rounded indentations dorsally; underside unsculptured, tail apparently lacking.

Immature. Known for four species. 5th instar oblong-ovoid, ventro-dorsally flattened with slightly protruding wing buds and distinct humeral lobes; antennae with 3(-4) segments bearing 3(-4) rhinaria (1 on segments 2-3, and 2 on apical segment) and two long, terminal, simple setae of unequal length; tarsi with broad crescent arolia and extremely small, reduced claws; each terminal tarsus bearing a long capitate seta; anus situated ventrally, circumanal ring broad and composed of a single row of elongate cells; dorsum either with wax producing pores (see Tuthill 1966), or non-wax producing tubercles and tentacles (Fig. 12). Chaetotaxy: 5th instar with either continuous or interrupted coverage of marginal setae; overall setal types, even between closely related species (e.g., S. reticulata and S. tentaculata), highly variable (Fig. 12). Smaller instars only known for S. reticulata and S. tentaculata, in which tubercles are apparent from 2nd instar (Fig. 13).

Biology.

All species for which the biology is known have free-living immatures on the surface of leaves (either lower, or both upper and lower surfaces). Those species with immatures described here with protruding tubercles and tentacles were mostly found on the lower leaf surface among dense indumentum and often close to the mid-rib (Fig. 12N).

Host plant.

All Swezeyana are host specific on a single Hawaiian endemic host plant species, Planchonella sandwicensis (Sapotaceae).

Comments.

Two species groups are recognized, elongagena group and reticulata group, based primarily on the strikingly different forms of female terminalia. The elongagena group has broad, truncate female terminalia with a strongly convex proctiger apex; the proctiger and subgenital plate bear a small to pronounced medial cleft at the apex. In contrast, the reticulata group has tapering terminalia without a medial cleft in the proctiger apex. In both species groups the subgenital plate terminates in a more or less well developed beak with small to pronounced cleft spanned by a membrane. The underlying endoskeleton of the two different forms of female terminalia indicate distinctly different development of the apodemes in the two species groups: broad and short in elongagena group (Fig. 4P), and long and narrow in reticulata group (Fig. 10I). However, not all species treated here are known for both sexes, therefore current assignment to these groups relies on other characteristics and DNA barcode data (see Discussion). Due to the unknown female morphology and inconclusive placement in the molecular phylogeny, S. magnaccai is not placed within a species group. Fore wing characteristics such as wing membrane colouration and pseudoveins/cross veins are found in both groups. The 5th instar immatures may also be diagnostic, with elongagena group having wax producing pores dorsally (illustrated in Tuthill 1966 for S. elongagena) and continuous ring of marginal setae (illustrated in Caldwell 1940 for S. elongagena); while in contrast, the reticulata group are characterized by non-wax producing tubercles and tentacles and lack a contiguous marginal ring of setae (Figs 12–13). However, currently immatures are known for these three species only, so it remains to be tested whether these highly distinct immature morphologies reflect species group assignments.

Note on adult assignment to species group.

The two species groups (elongagena group and reticulata group) are most easily recognized by the shape of the female terminalia, e.g., extremely convex apex of female proctiger in the elongagena group, versus more or less dorsally straight and tapering in the reticulata group. The elongagena group females have a FP:HW ratio typically < 0.70 (range 0.51–0.69), and FP:RL ratio typically < 2.18 (range 1.41–2.19); whereas in the reticulata group FP:HW ratio is typically > 0.70 (range 0.69–0.95), and FP:RL ratio is typically > 2.18 (range 2.17–2.59). Swezeyana males are less easily assigned to a species group, but elongagena group males have a distal aedeagus segment that is typically shorter than the paramere (PL:AEL ratio range 0.92–1.60), whereas reticulata group males have a distal aedeagus segment that is longer than the paramere (PL:AEL ratio range 0.65–0.88). Notably, the fore wing characters used by Zimmerman (1948) to key out the two species described by Caldwell (S. elongagena and S. reticulata), such as presence/absence of cross veins in cell r1 and presence/absence of distinct patterns of pigmentation are found in both species groups. The key below does not key to species group, rather it employs characters useful in distinguishing species, in particular those co-occurring on the same island.

Note on molecular analyses.

The neighbour-joining analysis of two mitochondrial DNA regions is presented in Fig. 2. Strong support is recovered for the reticulata group, but not for the elongagena group. The topologies recovered in comparative ML and Bayesian analyses differ only at weakly or unsupported nodes. The three nodes that group same island sister taxon pairs (on Kauai, Oahu, and Hawaii) are recovered in all analyses but with variable support (much stronger support in the Bayesian than NJ or ML analyses) (see Fig. 2 and Discussion). Maximum genetic divergence (uncorrected p-distances) among Swezeyana species is 19.9%; maximum intraspecific divergence (3%) was found in S. magnaccai on Oahu.

Key to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Swezeyana">Swezeyana</tp:taxon-name-part></tp:taxon-name> adults
1Fore wings with distinct darker patches or clouds of pigmentation, in some cases only around termination of veins at wing margins and around cross veins between Rs and wing margin in cell r1 (Fig. 1E–I) 2
–Fore wings without distinct darker patches or clouds of pigmentation, appearing uniformly clear or opaque, cross veins between Rs and wing margin in cell r1, if present, unpigmented (Fig. 1A–D) 6
2Antennae short, subequal to head width (ratio AL:HW < 1.1), ratio HW:GP > 1.65, female proctiger strongly convex apically with apex extremely broad and blunt, ratio FP:HW < 0.70, on Hawaii 3
–Antennae longer (AL:HW > 1.1), ratio HW:GP < 1.65, female proctiger more or less straight dorsally with apex bluntly acute, on other islands 4
3Fore wings with extensive patches and clouds of red-brown pigmentation, particularly across the central area of wing, and numerous cross veins (typically more than 5) between Rs and wing margin in cell r1, smaller species with broader wings (ratio WL:WW < 3.1) and wing cell cu1 relatively narrow and high (ratio CUR < 1.33) (Figs 1I, 8A–W) S. rubra sp. n.
–Fore wings with only indistinct fuscous brown on membrane and distinct small darker brown patches around termination of veins at wing margins and around cross veins between Rs and wing margin in cell r1, cross veins fewer (typically less than 5), larger species with narrower wings (ratio WL:WW > 3.1) and wing cell cu1 relatively wide and low (ratio CUR > 1.33) (Figs 1E, 5I–K) S. hawaiiensis sp. n.
4Larger species (WL > 1.9 mm), paramere shape more triangular with a broader base, on Kauai (possibly other islands for S. reticulata) 5
–Smaller species (WL < 1.9 mm), paramere shape more slender with a narrower base, on Oahu (Figs 1F, 11) S. magnaccai sp. n.
5Fore wing with fewer cross veins (typically less than 6) between Rs and wing margin, head with shorter genal processes (ratio HW:GP > 1.36) and shorter antennae (AL < 0.75 mm, ratio AL:HW ≤ 1.40), paramere shorter and broader (ratio PL:HW < 0.20, ratio PL:AEL < 0.80), female terminalia shorter (ratio FP:HW < 0.80) with posterior margin of anal ring less convoluted and incised (Figs 1G, 9A–T; immatures Figs 12A–G, 13A–F) S. reticulata Caldwell, 1940
–Fore wing with more cross veins (typically more than 6) between Rs and wing margin, head with longer genal processes (ratio HW:GP < 1.36) and longer antennae (AL > 0.75 mm, ratio AL:HW ≥ 1.40), paramere longer and narrower (ratio PL:HW > 0.20, ratio PL:AEL > 0.80), female terminalia longer (ratio FP:HW > 0.80) with posterior margin of anal ring more convoluted and incised (Figs 1H, 10A–S; immatures Figs 12H–N, 13G–J) S. tentaculata sp. n.
6Smaller species (WL < 3 mm, AL < 1 mm), wing cell cu1 relatively narrow and high (ratio CUR < 2.1), on Oahu, Molokai, Maui 7
–Larger species (WL > 3 mm, AL > 1 mm), wing cell cu1 relatively wide and low (ratio CUR > 2.1), on Kauai (Figs 1D, 6A–K) S. magna sp. n.
7Genal processes shorter (GP < 0.40 mm, ratio HW:GP > 1.35), paramere shorter (ratio PL:HW < 0.27) and with less extended anteriorly directed apex, on Oahu 8
–Genal processes longer (GP > 0.40 mm, ratio HW:GP < 1.35), paramere longer (ratio PL:HW > 0.27) and with more extended anteriorly directed apex, on Molokai and Maui (Figs 1A, 3A–P) S. elongagena Caldwell, 1940
8Paler species (generally yellow-brown to green), fore wing vein Rs typically longer relative to wing length (ratio WL:Rs 1.57–1.82) resulting in shorter distance between terminations of Rs and M1+2, antennae longer (AL > 0.65 mm), paramere longer (ratios PL:HW > 0.20, PL:AEL > 1.15, PL:SH > 0.90) (Figs 1B, 7A–R) S. oahuensis sp. n.
–Darker species (generally dark brown to black), fore wing vein Rs typically shorter relative to wing length (ratio WL:Rs 1.76–2.03) resulting in longer distance between terminations of Rs and M1+2, antennae shorter (AL ≤ 0.65 mm), paramere shorter (ratios PL:HW < 0.20, PL:AEL < 1.15, PL:SH < 0.90) (Figs 1C, 4A–T) S. atra sp. n.
Note on species descriptions.

Swezeyana is a small genus with, in general, considerable morphological homogeneity. The species descriptions below provide details of species specific characteristics not supplied in the generic description above.

Swezeyana elongagena. A head B head (lateral view) C male D head and antenna, inset detail of terminal antennal segments (terminal setae outlined) E female terminalia F detail of apex of female terminalia indicating beak and position of medial clefts in proctiger and subgenital plate (outlined) G male terminalia H paramere and distal aedeagus segment I paramere, inset detail of apex J eggs (distinctly hexagonal sculpturing and pedicel outlined and indicated) K hind leg L base of hind tibia with reduced genual spine indicated M mesotarsi and metatarsi (outlined), concave and ridged underside of basal metatarsus (outlined) N small but distinct meracanthus (indicated) O fore wing with interior edge of ventral margin outlined, inset illustrating broad shape of marginal radular spine cluster P extent of fore wing pseudopterostigma (shaded).

Swezeyana magna sp. n. A head and antenna, inset antenna detail B hind leg C base of hind tibia (reduced genual spine indicated) D small but distinct meracanthus (indicated) E metatarsi, inset comparative size of mesotarsi F dorsum of thorax G proboscis H aedeagus and paramere I extent of pseudopterostigma (shaded) J broad shape of marginal radular spine cluster (outlined) K fore wing, with interior edge of ventral margin outlined.

Swezeyana reticulata. A head B head and antenna (lateral view) C detail of antenna D hind leg, inset highly reduced meracanthus (indicated) and base of hind tibia with genual spine (indicated) E male F female G dorsum of thorax H head and antenna I metatarsi (outlined) J female proctiger (dorsal view) K detail of anal ring (outlined), illustrating circumanal ring pores L female terminalia (subgenital beak indicated, apex outlined) M female subgenital plate (ventral view, apex outlined) N detail of posterior apex of female subgenital plate with beak and membrane (outlined) O ovipositor P female terminalia (dorsal view), lateral valves extending beyond proctiger Q male terminalia R aedeagus and paramere, with details of paramere apex (above interior view, below dorsal view) S fore wing, with interior edge of ventral margin outlined T fore wing detail of pigmented cross veins.

Swezeyana reticulata and Swezeyana tentaculata 5th instar immatures. A–G, M S. reticulata: A detail of dorsal lanceolate setae with inflated and ridged bases anterior of eye B detail of dorsal sub-marginal lanceolate setae with inflated and ridged bases on margin of wing pads C detail of tubercles on margin of abdomen, and marginal narrow, blunt sectasetae D detail of dorsal tubercle bearing small simple setae, and small lanceolate setae with greatly inflated and ridged bases on surrounding surface E lateral view showing arrangement of dorsal tubercles, open arrows indicate position of thoracic and abdominal tubercles typically more darkly pigmented F dorsal view, open arrows indicate position of thoracic and abdominal tubercles typically more darkly pigmented G anal ring H–L, N S. tentaculata: H dorsal view, open arrows indicate position of thoracic and abdominal tentacles typically more darkly pigmented, inset detail of marginal pointed sectasetae I anal ring J, K detail of dorsal sub-marginal long, simple setae with narrowly inflated and ridged bases, K also shows different pigmentation for 1st and 2nd tentacle on abdomen margin L detail of long tentacle with simple setae towards the apex and a pair of small simple setae apically, and longer slightly capitate rod setae towards the base and on surrounding surface M tarsus and antenna (similar for both species) N red-brown S. tentaculata immatures found along the midribs on undersides of leaves among the red-brown leaf trichomes.

Swezeyana reticulata and Swezeyana tentaculata 1st-4th instar immatures. A–F S. reticulata: A 1st instar, inset detail of marginal narrow, blunt sectasetae B 2nd instar with appearance of tubercles C detail of 2nd instar anterior marginal and sub-marginal head setae D detail of 2nd instar marginal tubercles on the thorax bearing simple setae towards apices E 3rd instar, inset detail of marginal narrow, blunt sectasetae F 4th instar, inset detail of dorsal and sub-marginal lanceolate setae with inflated and ridged bases G–J S. tentaculata: G 4th instar, with details of marginal pointed sectasetae, and dorsal and sub-marginal simple setae with narrowly inflated and ridged bases H 3rd instar, with detail of larger dorso-medial tubercles bearing spiral of slightly capitate rod setae; marginal pointed sectasetae, and dorsal and sub-marginal simple setae with narrowly inflated and ridged bases as for 4th instar I detail of 3rd instar pairs of slightly capitate rod setae near the apices of marginal tubercles on the abdomen J detail of 3rd instar marginal pointed sectasetae, and dorsal sub-marginal simple setae.

Swezeyana atra sp. n. A head B head and antenna (lateral view) C antenna D proboscis E dorsum of thorax F male terminalia, inset distal aedeagus segment G paramere and distal aedeagus segment H parameres (dorsal view), inset illustrating paramere apex I paramere (posterior view) J female abdomen K female terminalia L female proctiger (dorsal view) M detail of anal ring N female subgenital plate (ventral view) O detail of posterior apex of female subgenital plate with membrane (outlined) P female terminalia endoskeleton (dorsal view, outlined) Q hind leg, inset reduced meracanthus (indicated) and metatarsi (outlined) R base of hind tibia with highly reduced genual spine (indicated) S extent of fore wing pseudopterostigma (shaded) T fore wing (above) with interior edge of ventral margin outlined, hind wing (below).

Swezeyana tentaculata sp. n. A head B head (lateral view) C head and antenna (lateral view) D dorsum of thorax E antenna F head and antenna G proboscis H hind leg, inset highly reduced meracanthus (indicated) and base of hind tibia with genual spine (indicated) I female terminalia endoskeleton (dorsal view) J female terminalia (subgenital beak indicated, apex outlined) K female proctiger (dorsal view) L female subgenital plate (ventral view, apex outlined) M detail of posterior apex of female subgenital plate with beak and membrane (outlined) N detail of anal ring (dorsal view, outlined) O egg (outlined), inset detail of sculpturing on dorsal service P aedeagus and paramere Q male terminalia R fore wing (above), with interior edge of ventral margin outlined, hind wing (below) S fore wing detail of pigmented cross veins and unpigmented membrane surrounding marginal radular spine clusters.

Neighbour-joining (NJ) analysis (combined COI and cytB data) with 1000 bootstrap replicates in PAUP*. Eight of the nine Swezeyana species are included and 16 other taxa from Triozidae, mostly representing other Hawaiian genera for comparison of divergence; and one species from Carsidaridae as an outgroup (Mesohomotoma hibisci) (Table 1). Outlines of parameres and female terminalia illustrate the two recognized species groups (elongagena group in black, reticulata group in grey, S. magnaccai unplaced), and fore wing images (Fig. 1) show the distribution of patterned/pigmented wings. Regional localities of sample are given, and “imm.” indicates an immature sampled for S. reticulata. NJ bootstrap support values ≥ 60% are indicated for all nodes, and comparative node support from ML and Bayesian analyses are shown for the three same island sister taxa pairs (NJ/ML[bold]/Bayesian[italic]) (see Discussion). Maximum genetic divergence (uncorrected p-distances) is shown for Swezeyana and four other endemic Hawaiian genera; also shown are maximum intraspecific distances (indicated with an asterix) for two widespread non-Hawaiian species with multiple individuals sampled across terrestrial (T. urticae) or oceanic (M. hibisci) landscapes (see Discussion).

Additional taxa sampled for the mitochondrial DNA analysis, with GenBank accession numbers.

SpeciesLocalityGenBank COI/cytB (publication)
Family: Carsidaridae
Mesohomotoma hibisci (Froggatt, 1901)Society Islands (Moorea)KY294174/KY294658 (Percy 2017a)
Society Islands (Raiatea)KY294171/KY294655 (Percy 2017a)
New CaledoniaKY294170/KY294654 (Percy 2017a)
SingaporeKY294176/KY294660 (Percy 2017a)
Family: Triozidae
Anomocephala unica Tuthill, 1942Austral Islands (Rapa)KY293698/KY294177 (Percy 2017a)
Bactericera cockerelli (Šulc, 1909)CaliforniaKY011201/KY011296 (Percy 2017a)
Hemischizocranium aloha (Caldwell, 1940)Hawaiian Islands (Kauai)MG988755/MG989062 (this study)
Hemischizocranium bessi Tuthill, 1956Hawaiian Islands (Hawaii)MG988756/MG989063 (this study)
Hevaheva maculata Caldwell, 1940Hawaiian Islands (Kauai)KY293702/KY294181 (Percy 2017a)
Hevaheva minuta Crawford, 1925Hawaiian Islands (Kauai)KY293703/KY294182 (Percy 2017a)
Hevaheva perkinsi Kirkaldy, 1902Hawaiian Islands (Oahu)KY293704/KY294183 (Percy 2017a)
Hevaheva silvestris Kirkaldy, 1908Hawaiian Islands (Oahu)KY293705/KY294184 (Percy 2017a)
Pariaconus iolani (Kirkaldy, 1902)Hawaiian Islands (Kauai)KY293820/KY294297 (Percy 2017a)
Pariaconus proboscideus Percy, 2017Hawaiian Islands (Hawaii)KY294095/KY294573 (Percy 2017a)
Pariaconus pyramidalis Percy, 2017Hawaiian Islands (Hawaii)KY294124/KY294607 (Percy 2017a)
Pariaconus mauiensis Percy, 2017Hawaiian Islands (Maui)KY293841/KY294316 (Percy 2017a)
Stevekenia aiea Percy, 2017Hawaiian Islands (Kauai)KY971542/KY971544 (Percy 2017b)
Stevekenia nothocestri Percy, 2017Hawaiian Islands (Oahu)KY971541/KY971543 (Percy 2017b)
Trioza remota Foerster, 1848EnglandKY294162/KY294646 (Percy 2017a)
Trioza urticae (Linné, 1758)EnglandKY011191/KY011286 (Wonglersak et al. 2017)
T. urticae GreeceKY011122/KY011219 (Wonglersak et al. 2017)
NorwayKY011175/KY011270 (Wonglersak et al. 2017)
PolandKY011114/KY011212 (Wonglersak et al. 2017)

Fore wings of nine Swezeyana species: A S. elongagena (male) B S. oahuensis (male) C S. atra (male) D S. magna (male) E S. hawaiiensis (female) F S. magnaccai (male) G S. reticulata (male above, female below) H S. tentaculata (male above, female below) I S. rubra (male above, female below). Scale bars: 1 mm.

Swezeyana rubra sp. n. A head B head (lateral view) C female D head and antenna E antenna F proboscis G reduced meracanthus (indicated) H hind leg I metatarsi (outlined), inset comparative size of mesotarsi (outlined) J base of hind tibia K dorsum of thorax L female terminalia (subgenital beak indicated, apex outlined) M female proctiger (dorsal view) N female terminalia (dorsal view), inset apex of proctiger (ventral view, cleft indicated) O detail of anal ring (outlined) P female subgenital plate (ventral view) Q detail of posterior apex of female subgenital plate with beak and membrane (outlined) R aedeagus and paramere S male terminalia, inset aedeagus T paramere (posterior view) U eggs (semi-hexagonal sculpturing, pedicel and unsculptured underside outlined and indicated) V fore wing, with interior edge of ventral margin outlined W fore wing detail of pigmented cross veins.

Swezeyana hawaiiensis sp. n. (female). A head and antenna B antenna C proboscis D proctiger, dorso-lateral view showing anal ring (outlined) E detail of apex of proctiger showing medial cleft and fringe of apical setae (indicated) F female subgenital plate (ventral view) with apical beak and membrane outlined G hind leg H small but distinct meracanthus (indicated) I base of hind tibia (highly reduced genual spine indicated) J detail of atypical 1+3 (indicated) arrangement of sclerotized apical metatibia spurs K fore wing, with interior edge of ventral margin outlined.

Swezeyana magnaccai sp. n. A head B head and antenna (lateral view) C proboscis D dorsum of thorax E hind leg F base of hind tibia G male terminalia H aedeagus and paramere I male terminalia (dorsal view), inset details of paramere apices J fore wing detail of termination of vein R at base of pseudopterostigma K fore wing detail of incomplete termination of vein Rs at wing margin (inset incomplete veins indicated) L fore wing, with interior edge of ventral margin outlined M fore wing detail of unpigmented membrane surrounding marginal radular spine clusters.

Swezeyana oahuensis sp. n. A head B head and antenna (lateral view) C proboscis D antenna E hind leg, inset detail of reduced meracanthus (indicated) and genual spine F dorsum of thorax G female H female terminalia (truncate subgenital plate indicated, apex outlined) I female terminalia (dorsal view) J detail of anal ring (outlined) K female subgenital plate (ventral view) L detail of posterior apex of female subgenital plate with beak and membrane (outlined) M male terminalia (dorsal view) N paramere apex (dorsal view above and outlined, lateral view below) O male terminalia P paramere (posterior view) Q aedeagus and paramere R fore wing, with interior edge of ventral margin outlined.

 TuthillLD (1966) Notes on Swezeyana (Psyllidae: Homoptera). Proceedings of the Hawaiian Entomological Society 19: 309. CaldwellJS (1940) New genera and species of jumping plant-lice from the Hawaiian Islands with descriptions of several immature stages (Homoptera: Psyllidae). Proceedings of the Hawaiian Entomological Society 10: 389–397. ZimmermanEC (1948) Insects of Hawaii: Vol. V. Homoptera: Sternorhyncha: Superfamily Psylloidea. University of Hawaii Press, Honolulu, 11–38.