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A fourth contribution to the study of Hawaiian Orthotylus (Heteroptera: Miridae): new species, addenda, revised checklist, and key to the Perrottetia-feeding species.

Abstract--Supplemental data and addenda are provided to complement and correct the work of Polhemus (2011) dealing with Hawaiian Orthotylus species. The following new species are described, accompanied by illustrations of key morphological characters, distribution maps, and color dorsal habitus photographs: O. erythrinae from Oahu. which feeds on Erythrina sandwicensis, and O. broussaisiae from Kauai, which feeds on Broussaisia arguta. Additional distribution records are provided for O. dubautiae D. Polhemus, O. hedyotivorus D. Polhemus, and O. kopikoides D. Polhemus. The name Orthotylus perrottetioides is proposed as a replacement name for Orthotylus perrottetiopsis Polhemus 2011. which due to a lapsus is a primary homonym of Orthotylus perrottetiopsis Polhemus 2003, and addenda to Polhemus (2011) related to this change are detailed. A revised key to Hawaiian species of Orthotylus occurring on Perrottetia is provided, accompanied by color dorsal habitus photographs of the males and females of the four known Perrottetia-feeding Hawaiian species. A revised checklist and updated of Hawaiian Orthotylus species, host plants, and islands of occurrence is also provided.

Key words: Plant bugs, Miridae, Orthotylus, new species, host plants.

INTRODUCTION

The plant bug genus Orthotylus has undergone a remarkable insular radiation within the Hawaiian Islands, with extensive diversification across a wide range of host plants. The current paper describes two new species that feed on two genera and one family not previously documented as plant hosts of Hawaiian Orthotylus. As a result, the genus in the Hawaiian Islands now contains 95 species that feed on 45 plant species in 19 genera representing 16 families. This paper also corrects certain errors that were committed in the course of documenting this complex and speciose insular plant bug radiation, in particular the rectification of a primary homonym. To alleviate the confusion that may have been caused, a revised key is provided to the Hawaiian Orthotylus species feeding on Perrottetia, all of these species are illustrated in color, and a revised checklist of species names, host plants, and islands of occurrence is provided. The latter checklist should be used in preference to that provided in Polhemus (2011).

MATERIAL AND METHODS

All measurements in the descriptions below are provided in millimeters. Morphological terminology for male genitalic structures follows the previous paper of D. Polhemus (2011). CL numbers in the Material Examined sections refer to a collection locality coding system used by the author to cross reference locality data, specimens, and photographs. Holotypes and paratypes are held in the Bishop Museum, Honolulu, Hawaii (BPBM). As with previous papers in this series (D. Polhemus, 2003, 2005, 2011), nomenclature for host plants follows Wagner, et al. (1999).

GENUS ORTHOTYLUS FIEBER, 1858 ADDITIONAL SPECIES FEEDING ON FABACEAE

Two species of endemic Hawaiian Orthotylus have previously been documented to feed representatives of the Fabaceae, both of them occurring on the endemic tree Sophora chysophylla. As reported below, another Hawaiian species is now at hand feeding on the single endemic Hawaiian representative of the widespread genus Erythrina, thus adding another genus to the known assemblage of host plants utilized.

Orthotylus erythrinae, new species Figures 1A, B, 2, 3A C

HOLOTYPE: HAWAIIAN ISLANDS, Oahu, Waianae Mountains, behind Dillingham AFB, 26 April 1991, on Erythrina sandwicensis, A. Asquith, [male] (BPBM).

TYPE LOCALITY: Oahu, Dillingham Air Force Base.

DESCRIPTION: Male: Overall length 2.90, length from tip of tylus to cuneal fracture 1.90, maximum width (across base of cuneus) 1.00. General coloration pale green, with head darker; appendages predominantly pale green (Fig. 1A). Head: Length 0.28, width across eyes 0.70, olive green; eyes dark red, length 0.20, width 0.20; vertex width 0.30, bearing scattered, semi-erect, simple golden setae only. Antennae: predominantly pale brown, with distal 1/5 of segment II, distal half of segment III, and all of segment IV darker, segment I bearing 3 erect spinose golden setae, all segments thickly covered with short, appressed pale setae, lengths of segments I-IV = 0.28, 0.95, 0.60, 0.35. Rostrum: length 0.70, golden with dark brown apex, reaching to bases of hind coxae. Thorax: Pronotum: Length 0.48, width 0.95, pale green, surface bearing scattered, semi-erect, simple golden setae only. Scutellum: Length 0.40, width 0.50, pale green, scutellar surface bearing scattered, semi-erect, simple golden setae only. Hemelytra: Uniformly pale green, translucent; wing membrane brownish fumate, veins slightly darker; hemelytra bearing scattered, semi-erect, simple golden setae only. Legs: All segments golden yellow, extreme tips of tibiae and terminal tarsi embrowned; all leg segments clothed with very short, pale, recumbent setae, tibiae and apices of femora bearing long, erect, golden brown spines. Ventral surface: Pale green, clothed with short, recumbent pale setae, genitalia: Right paramere: Club-like, apex blunt and rounded (Fig. 3B). Left paramere: Falciform, with a shallow subapical concavity isolating a small apical process (Fig. 3A). Spicules of endosoma as in Fig. 3C, central spicule with a serrate apex.

Female: Overall length 2.90, length from tip of tylus to cuneal fracture 1.95, maximum width (across base of cuneus) 1.10; similar to male in general structure and coloration, with following exceptions: pronotum, scutellum, clavus and inner corium dark yellow, outer corium broadly pale green (Fig. 1B); eyes similar in proportional size to those of male, width 0.20, width of vertex 0.30.

HOST PLANT: Erythrina sandwicensis Degener (Fabaceae).

PARATYPES: same data as holotype, 20 [male][male] 24 [female][female] (BPBM); OTHER SPECIMENS EXAMINED: HAWAIIAN ISLANDS, Oahu, hills behind Dillingham AFB [Air Force Base], 14 December 1968, on Erythrina sandwicensis, W. C. Gagne, 1 [male] 2 [female][female], (BPBM); Kaena Point, April 1955, on Erythrina sandwicensis, Ford 1 [male] 5 [female][female], (BPBM).

ETYMOLOGY: The name "erythrinae" is derived from the host plant genus Erythrina, on which this species Feeds.

DISCUSSION: Orthotylus erythrinae is a small, pale yellow to pale brown colored species feeding on Erythrina sandwicensis on Oahu. This species runs to couplet 76 in Polhemus (2011), where it produces an illogical result because the male left paramere lacks either a hook or a deep incision at the apex. It may instead be recognized by the presence of only simple setae on the dorsum, the slender left paramere which is strongly angled centrally (Fig. 3B), the right paramere that is modestly expanded on its basal section and then slightly bilobate and hooked apically (Fig. 3A), and by the shapes of the endosomal spicules (Fig. 3C).

ECOLOGICAL NOTES: The host plant of this new species, Erythrina sandwicensis, is known to native Hawaiians as the "wiliwili," and was a formerly common lowland dry forest tree that is now reduced to remnant stands at a few sites on Oahu. These remaining wiliwili populations have been further reduced in recent years due to impacts from the introduced gall wasp Quadrastichus erythrinae, which reached Hawaii in 2005. Although the impacts of this inadvertent introduction have been subsequently mitigated to some degree by the introduction of another wasp, Eurytoma erythrinae, whose immatures feed on Quadrastichus, the fact remains that many of the older wiliwili trees on Oahu that formerly served as host plants for Orthotylus erythrinae have now died due to the gall wasp infestation. This, coupled with the fact that wiliwili is a dryland tree which only flushes out with leaves and flowers irregularly in response to rains, means that O. erythrinae is now an uncommonly encountered insect on Oahu.

SPECIES FEEDING ON HYDRANGACEAE

The family Hydrangeaceae consists of 17 genera and 170 species that are widely distributed in the Northern Hemisphere, with a few additional species also reaching Southeast Asia. A single endemic and monotypic genus in this family, Broussaisia, occurs in Hawaii, where its single included endemic species, B. arguta, is a common understory element in wet forests on all the main Hawaiian islands except Niihau and Kahoolawe. Despite extensive collecting on this host plant over the past decade, only one Orthotylus species, described below from Kauai, has so far been found to feed on it.

Orthotylus broussaisiae, new species Figures 1C, D, 3D F, 4

HOLOTYPE: HAWAIIAN ISLANDS, Kauai, Mt. Waialeale, moss forest patch 0.6 mi. W. of summit, 1,490 m, 22[degrees]4'14"N, 159[degrees]30'16"W, 18 May 2005, CL 8467, D. A. Polhemus, on Broussaisia arguta, [male] (BPBM).

DESCRIPTION: Male: Overall length 3.70, length from tip of tylus to cuneal fracture 2.30, maximum width (across base of cuneus) 1.30. General coloration dark brown, with pale markings on scutellum and hemelytra, legs predominantly golden-yellow (Fig. 1C). Head: Length 0.20, width across eyes 0.80, dark brown: eyes dark red, length 0.20, width 0.20; vertex width 0.40, bearing scattered senti-recumbent golden setae only. Antennae: with segment 1 and basal two thirds of segment II pale yellow, remainder of segment II plus entirety of segments III and IV black, segment I bearing 3 erect spinose golden setae, all segments thickly covered with short, appressed pale setae intermixed with scattered short semi-erect golden-brown setae, lengths of segments I-IV-0.40, 1.45, 0.80, 0.40. Rostrum: Length 1.10, reaching to middle coxae, golden-brown, extreme tip darker. Thorax: Pronotum: Length 0.60, width 1.15, uniformly dark brown, surface bearing scattered semi-erect, dark brown, simple setae only. Scutellum length 0.60, width 0.80, medium brown, with posterior apex broadly dark yellow, surface bearing scattered semi-erect, golden, simple setae only. Hemelytra: Clavus dark brown, central section of corium broadly medium brown, anterior angles of corium and basal half of cuneus pale white, anterior half of cuneus pale brown; wing membrane including veins uniformly dark fumate brown: hemelytral surface bearing scattered senti-erect, golden, simple setae only. Legs: Golden-yellow, posterior half of hind lemur dark brown, terminal tarsal segments golden brown; all leg segments clothed with very short, recumbent, gold setae, tibiae and apices of femora bearing long, erect, golden spines. Ventral surface: Pale brown, ostiolar peritreme creamy white, abdominal venter bearing short, recumbent, pale, simple setae, these setae longer and more numerous on lateral margins of genital cavity. Genitalia: Right paramere: very slender basally, greatly expanded distally, distal lobe roughly in the form of an inverted triangle with one lateral apex bearing a short, blunt process (Fig. 3E). Left paramere: greatly expanded basally, margin broadly curving, narrowing abruptly to a short distal arm with a transverse head coming to a sharply acute apex (Fig. 3D). Spicules of endosoma as in Fig. 3F.

Female: Overall length 4.00, length from tip of tylus to cuneal fracture 2.50, maximum width (across base of cuneus) 1.30; similar to male in general structure and coloration, with following exceptions: head dark brown, with lateral margins of vertex bordering inner margins of eyes narrowly dark yellow; pronotum dark blackish brown, with posterior margins of calli broadly dark yellow; hemelytra with clavus dark brown, central corium broadly dark red, anterior angles of corium translucent yellow, basal half of cuneus red, posterior half of cuneus creamy white: wing membrane fumate brown, veins dark brown surrounding inner cell, bright red surrounding outer cell (Fig. 1D); eyes slightly smaller in proportional size to those of male, width 0.20, width of vertex 0.50.

PARATYPES: same data as holotype, on Broussaisia arguta, 1 [female] (BPBM).

Host PLANT: Broussaisia arguta Gaudichaud-Beaupre (Hydrangaceae).

Discussion: Orthotylus broussaisiae is a relatively large, predominantly reddish-brown species from the uplands of Kauai. This species runs to couplet 33 in Polhemus (2011), where it produces an illogical result, because the posterior half of the cuneus is brown, but the species does not feed on any genus of Rubiaceae. If forced past this host plant consideration, the species will run to couplet 50, where it produce an illogical result due to its non-conforming combination of paramere shapes (Fig. 3D, E).

It should be noted that the male holotype is slightly teneral, therefore the darker and more reddish coloration of the female specimen may be more indicative of fully mature coloration in both sexes. If more mature specimens prove to exhibit reddish coloration on the posterior portion of the cuneus, then they will run to R. perrotettiae in the key provided by D. Polhemus (2011). It differs from that species in possessing a transversely expanded head on the distal arm of the left paramere (compare fig. 3D with fig. 21 in Polhemus, 2005), in having the distal section of the right paramere more broadly expanded and with a small process at one of the lateral angles (compare fig. 3E to fig. 23 in Polhemus, 2005), and in the presence of uniformly slender and tapering spinules in the phallotheca (compare fig. 3F to fig. 22 in Polhemus, 2005).

ECOLOGICAL NOTES: The type series was taken by beating a Broussaisia arguta shrub in a small patch of wet forest nestled within a gulch head amid the moorlands in the lee of the Mt. Waialeale summit. This is a one of the wettest places on the planet, averaging over 11,500 mm (452 inches) of rain per year over the past 100 years.

SPECIES FEEDING ON CELASTRACEAE

Four species of Hawaiian Orthotylus are known to feed on Perrottetia, the sole genus of the family Celastraceae occurring in Hawaii. Two of these species were described by D. Polhemus (2005), and two more species were described by D. Polhemus (2011). Due to a lapsus, one of the latter two species was given the same name as one of the former pair, creating a primary hononym. This error is corrected below, and a revised key to species, accompanied by dorsal color habitus photos of all Perrottetia-feeding species, is provided.

Orthotylus perrottetioides, new name Figure 5E, F

Orthotylus perrottetiopsis D. Polhemus, 2011: 59. Primary homonym of O. perrottetiopsis D. Polhemus, 2005: 256.

DISCUSSION: Due to a lapsus, D. Polhemus (2011) reutilized the name Ortholylus perrottetiopsis for a new Orthotylus species from Mt. Kaala, Oahu, despite the fact that it had been previously applied seven years earlier to another Orthotylus species from Maui. The error is rectified herein, and the name Orthotylus perrottetioides proposed as a replacement name for the Oahu species.

REVISED KEY TO SPECIES OF HAWAIIAN ORTHOTYLUS FEEDING ON PERROTTETTIA

The following key separates the four endemic Hawaiian species of Orthotylus that feed on the genus Perrottetia, and is intended to supercede the portions of a larger key including these species that appeared in D. Polhemus (2011). This revised key is accompanied by a set of color figures illustrating the males and females of all four species, which should aid in unambiguous identification.

Males

1. Dorsal surface bearing only simple setae,
lacking recumbent thickened or scale-like setae,
these simple setae may be a mixture of erect
and recumbent, but the recumbent setae not
thickened or scale-like, although they may
form a layer of appressed, gold-colored simple
hairs beneath the more upright setal layer
(Fig. 5E, F); male genitalia as in Fig. 3E-G in
D. Polhemus (2011); endemic to Oahu
                             O. perrottetioides D. Polhemus

--Dorsal surface with numerous, conspicuous
pale, scale-like hairs intermixed with darker
simple hairs, or with thickened, recumbent pale
setae intermixed with darker simple erect setae
(Fig. 5A, B, C, D, G, H)                                  2

2. Hemelytra almost entirely orange or reddish,
with pale markings limited to at most a narrow
transverse band at base of cuneus (Fig. 5G, H);
male genitalia as in fig. 3A D in D. Polhemus
(2011); endemic to Molokai
                              O. perrotteticola D. Polhemus

--Hemelytra bearing a pale area on basal section
of corium (Fig. 5A, B, C, D); Kauai or Maui               3

3. Larger species, body length exceeding 3.1 ram;
dorsal coloration as in Fig. 5A, B; male genitalia
as in figs. 21 23 in D. Polhemus (2005), with fight
paramere greatly expanded and roughly triangular
on distal half; endemic to Kauai
                                O. perrottetiae D. Polhemus

--Smaller species, body length equal to or less than
2.9 mm; dorsal coloration as in Fig. 5C, D; male
genitalia as in figs. 24-25 in D. Polhemus (2005),
with right paramere slender throughout and
bearing a small cap distally; endemic to
Maui                         O. perrottetiopsis D. Polhemus


NEW RECORDS FOR PREVIOUSLY DESCRIBED SPECIES

Continued sorting of undetermined backlog material in the Bishop Museum collections has provided further series of the following species, listed below.

Orthotylus dubautiae D. Polhemus

DISCUSSION: Originally the species was described by D. Polhemus (2011) from a short series of 2 males and 2 females taken on Mohihi Ridge, Kauai, with one additional female from the Alakai Swamp. The new records below indicate this species is widespread on a number of different Dubautia species occurring on the Alakai Plateau. It should be noted that despite the Gagne host plant label on the second series listed, there is no plant taxon named Dubautia "laxifolia." This host record most likely refers to either Dubautia laxa, or Dubautia latifolia, both of which occur on Kauai. It is hypothesized that the labelling represents an inadvertent incorrect spelling of Dubautia latifolia (A. Gray) D. D. Keck, which occurs in the Kokee area.

SPECIMENS EXAMINED: HAWAIIAN ISLANDS, Kauai: Kokee State Park, Kaluapuhi Trail, 1,300 m, 23 June 1980, on Dubautia raillardiodes, W. C. Gagne, 4 [male][male], 2 [female][female] (BPBM); Kokee State Park, 1,100 m, 23 June 1980, W. C. Gagne, on Dubautia laxifolia [sic], 8 [male][male], 12 [female][female] (BPBM).

Orthotylus hedyotivorus D. Polhemus

DISCUSSION: Previously known only from the type series taken at the summit of Aiea Ridge, above Kalauao Valley. The new record below is from a locality further northwest along the Koolau Mountains, but from a similar wet forest zone, and once again feeding on Hedyotis terminalis.

MATERIAL EXAMINED; HAWAIIAN ISLANDS, Oahu: Koolau Mountains, Castle Trail, S. of Kaluanui Stream crossing, 645 m, 21[degrees]34'05"N, 157[degrees]54'59"W, 16 May 2011, on Hedyotis terminalis, CL 8518, D. A. Polhemus, 1 [male], 3 [female][female] (BPBM).

Orthotylus kopikoides D. Polhemus

DISCUSSION: The new record below lies between the two sites from where this species was previously recorded by D. Polhemus (2011).

MATERIAL EXAMINED: HAWAIIAN ISLANDS, Molokai: Kamakou Reserve, Puu Kolekole cabin [21[degrees]06'15"N, 156[degrees]53'47"W], 24 August 1991, on Psychotria sp., A. Asquith, 2 [male][male] 1 [female] (BPBM).

ADDENDA TO PREVIOUSLY PUBLISHED WORKS ON HAWAIIAN ORTHOTYLUS

In addition to the corrected species name noted above, the following corrections should be made to Polhemus (2011):

Abstract--Change O. perrottetiopsis to O. perrottetioides.

Table of Contents--Change O. perrottetiopsis to O. perrottetioides.

Color plate 3--Change O. perrottetiopsis to O. perrottetioides.

Key--Couplet 36--Change O. perrotteticola to O. perrottetiopsis.

Key--Couplet 57--In second half of couplet, the first line should read: "... males with hemelytra yellowish brown, often with a large dark patch centrally ..." Also note that this error was carried over from the version of the key that was presented in D. Polhemus (2005).

Key--Couplet 72 The first line of this couplet should read "Anterior margin of antennal segment II hearing a thick brush of erect pale pilose setae ..." Also note that this error was carried over from the version of the key that was presented in D. Polhemus (2005).

Key--Couplet 94--Change O. perrottetiopsis to O. perrottetioides.

Fig. 3--Change O. perrottetiopsis to O. perrottetioides.

Fig. 4--Change O. perrottetiopsis to O. perrottetioides.

Page 59--Change species header from O. perrottetiopsis to O. perrottetioides.

Page 59--Etymology should read as follows: The name "perrottetioides" is derived from the similar congeneric species Orthotylus perrottetiae, and the Greek eidos, having the form of, and refers to the similarity of this species of O. perrottetiae.

Page 60--Discussion should read as follows: Among the Perrottettia-feeding species known to date, O. perrottetioides can be recognized by the absence of yellow coloration on the dorsum, the extensive bright red coloration on the females, and the distinctive male genitalia. The slender male right paramere is a character state shared with O. perrottetiopsis from Maul.

Page 102--Ecological Notes final line should read as follows: For further notes on the Culvert 59 Ridge type locality, see the previous discussion under Orthotylus perrottetioides.

Page 107--The portion of the checklist dealing with species feeding on Perrotettia should read as follows:

TAXON

Orthotylus perrottetiae

Orthotylus perrottetioides

Orthotylus perrotteticola

Orthotylus perrottetiopsis

HOST PLANT

Perrottetia sandwicensis

Perrottetia sandwicensis

Perrottetia sandwicensis

Perrottetia sandwicensis

ISLAND

Kauai

Oahu

Molokai

Maui

Page 108 Appendix 2, caption for Plate 2, change O. perrottetiopsis to O. perrottetioides.

Caption: Fig. 1. Dorsal color habitus photos of Hawaiian Orthotylus species. A. Male and B. female of Orthotylus erythrinae n. sp. from Dillingham airfield, Oahu. C. Male and D. female of Orthotylus broussaisiae n. sp. from Mt. Waialeale, Kauai.

Caption: Fig. 2. Distribution of O. erythrinae n. sp. on Oahu.

Caption: Fig. 3. Male genitalic structures of Hawaiian Orthotylus species. A. Left paramere, lateral view, B. right paramere, lateral view, and C. endosomal spicules of Orthotylus erythrinae n. sp. from Dillingham airfield, Oahu. D. Left paramere, lateral view, E. right paramere, lateral view, and F. endosomal spicules of Orthotylus broussaisiae n. sp. from Mr. Waialeale, Kauai.

Caption: Fig. 4. Distribution of O. broussaisiae n. sp. on Kauai.

Caption: Fig. 5. Dorsal color habitus photos of Hawaiian Orthotylus species feeding on Perrottetia. A. Male (holotype) and B. female (paratype) of Orthotylus perrottetiae D. Polhemus from Alakai Swamp, Kauai. C. Male (holotype) and D. female (paratype) of Orthotylus perrottetiopsis n. sp. from Makawao Forest Reserve, Maui. E. Male and F. female of Orthotylus perrottetiodes D. Polhemus paratype specimens from Mt. Kaala, Oahu. G. Male and H. female of Orthotylus perrotteticola D. Polhemus paratype specimens from Kahawaiiki Stream, Molokai.

ACKNOWLEDGMENTS

The continuing documentation of Hawaii's Orthotylus biota has been accomplished only with the generous support of many individuals, both in the field and the lab. In regard to the current supplement, special thanks are due to Dr. James K. Liebherr of Cornell University, Ithaca, and Dr. Curtis Ewing of the University of California, Berkeley, for assistance and companionship in the field as we surveyed the wettest place on the planet; to the Hawaii Department of Land and Natural Resources who granted us access to Mt. Waialeale; and to Dr. Neal Evenhuis and Dr. Allen Allison of the Bishop Museum, Honolulu, Hawaii (BPBM) for their continuing logistical and laboratory support.

This paper represents contribution 2013-104 to the Hawaii Biological Survey.

LITERATURE CITED

Polhemus, D. A. 2003. An initial review of Orthotylus in the Hawaiian Islands, with descriptions of twenty-two new species (Heteroptera: Miridae). Journal of the New York Entomological Society 110(3-4): 270-340.

Polhemus, D. A. 2005. Further studies on the genus Orthotylus (Heteroptera: Miridae) in the Hawaiian Islands, with descriptions of thirty-four new species. Journal of the New York Entomological Society 112(4): 22?-333.

Polhemus, D. A. 2011. Continuing studies on the genus Orthotylus in the Hawaiian Islands (Heteroptera: Miridae), with descriptions of thirty-two new species. Entomologica Americana 117(1/2): 37 109.

Wagner, W. L., D. R. Herbst and S. H. Sohmer. 1999. Manual of the Flowering Plants of Hawaii. Revised Edition. Bishop Museum Press, Honolulu, 1919 pp.

DAN A. POLHEMUS (1)

Dept. of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817

(1) E-mail address for correspondence: bugman@bishopmuseum.org

REVISED CHECKLIST OF HAWAIIAN ORTHOTYLUS SPECIES
WITH CONFIRMED BREEDING HOST PLANT ASSOCIATIONS

Table. Taxa are grouped by host plant genus, and by island
endemicity from north to south.

TAXON                          HOST PLANT               ISLAND

Orthotylus elaeocarpi          Elaeocarpus bifidus      Kauai

Orthotylus pisoniacola         Pisonia sandwicensis     Kauai

Orthotylus pisoniae            Pisonia umbellifera,     Oahu
                               Pisonia sandwicensis

Orthotylus pisoniopsis         Pisonia brunoniana       Maui

Orthotylus pisonioides         Pisonia brunoniana,      Hawaii
                               Pisonia sandwicensis

Orthotylus manonophila         Hedyotis terminalis      Kauai

Orthotylus manonoides          Hedyotis terminalis      Oahu

Orthotylus manoniella          Hedyotis                 Oahu
                               centranthoides

Orthotylus hedyotiella         Hedyotis acuminata       Oahu

Orthotylus hedyotiopsis        Hedyotis acuminata       Oahu

Orthotylus hedyoti             Hedyotis terminalis      Oahu

Orthotylus manono              Hedyotis terminalis      Molokai

Orthotylus manonocola          Hedyotis terminalis      Molokai

Orthotylus manonophagus        Hedyotis acuminata       Molokai

Orthotylus manonovorus         Hedyotis terminalis      Lanai

Orthotylus hedyotioides        Hedvotis terminalis      Maui

Orthotylus hedyotiphila        Hedyotis hillebrandii    Maui

Orthotylus hedyoticola         Hedyotis terminalis      Hawaii

Orthotylus ilicis              Ilex anomala             Maui

Orthotylus neoilicis           Ilex anomala             Hawaii

Orthotylus perkinsi            Ilex anomala             Hawaii

Orthotylus kopikocola          Psychotria hexandra      Kauai

Orthotylus kopikovorus         Psychotria mauiensis     Kauai

Orthotylus kara                Psychotria kaduana       Oahu

Orthotylus psychotricola       Psychotria mariniana     Oahu

Orthotylus kopikoides          Psychotria mariniana     Molokai

Orthotylus kopikopsis          Psychotria mariniana     Lanai

Orthotylus                     Psychotria mariniana     Lanai
neopsychotriopsis

Orthotylus neopsychotricus     Psychotria mariniana     Maui

Orthotylus kopikophila         Psychotria mariniana     Maui

Orthotylus kopiko              Psychotria mariniana     Maui

Orthotylus kiko                Psychotria mariniana     Maui

Orthotylus                     Psychotria mariniana     Maui
neospychotriodes

Orthotylus neopsychotriae      Psychotria hawaiiensis   Hawaii

Orthotylus psychotriae         Psychotria hawaiiensis   Hawaii

Orthotylus psychotrioides      Psychotria hawaiiensis   Hawaii

Orthotylus coprosmaphagus      Coprosma waimeae,        Kauai
                               Coprosma kauaiensis

Orthotyus coprosmoides         Coprosma ochracea        Oahu

Orthotylus coprosmicola        Coprosma foliosa         Molokai

Orthotylus pilo                Coprosma foliosa         Molokai

Orthotylus coprosmivorus       Coprosma ochracea        Lanai

Orthotylus coprosmae           Coprosma foliosa,        Maui
                               Coprosma pubens

Orthotylus neocoprosmae        Coprosma foliosa         Maui

Orthotylus coprosmaphila       Coprosma montana         Maui

Orthotylus coprosmopsis        Coprosma menziesii       Hawaii

Orthotylus kanakanus           Coprosma rhynchocarpa    Hawaii

Orthotylus pipturicola         Pipturus albidus         Kauai

Orthotylus tantali             Pipturus albidus         Oahu

Orthotylus pipturiphila        Pipturus albidus         Molokai

Orthotylus pseudotantali       Pipturus albidus         Maui

Orthotylus pipturoides         Pipturus albidus         Maui

Orthotylus pipturi             Pipturus albidus         Hawaii

Orthotylus urerae              Urera kaalae             Oahu

Orthotylus ureraphila          Urera glabra             Maui

Orthotylus urericola           Urera glabra             Hawaii

Orthotylus clermontiopsis      Clermontia faurieri      Kauai

Orthotylus clermontiella       Clermontia kakeana       Oahu

Orthotylus clermontioides      Clermontia               Maui
                               arborescens,
                               Clermontia micrantha,
                               Clermontia kakeana

Orthotylus clermonticola       Clermontia grandiflora   Maui

Orthotylus clermontiae         Clermontia               Hawaii
                               clermontioides

Orthotylus iolani              Clermontia               Hawaii
                               clermontioides

Orthotylus xylosmicola         Xylosma hawaiiense       Kauai

Orthotylus daphne              Xylosma hawaiiense       Oahu

Orthotylus xylosmoides         Xylosma hawaiiense       Maui

Orthotylus xylosmae            Xylosma hawaiiense       Hawaii

Orthotylus diospyropsis        Diospyros hillebrandi    Oahu

Orthotylus diospyroides        Diospyros sandwicensis   Molokai

Orthotylus diospryrivorus      Diospyros sandwicensis   Lanai

Orthotylus diospyrella         Diospyros sandwicensis   Maui

Orthotylus diospyri            Diospyros sandwicensis   Hawaii

Orthotylus perrottetiae        Perrottetia              Kauai
                               sandwicensis

Orthotylus perrottetioides     Perrottetia              Oahu
                               sandwicensis

Orthotylus perrotteticola      Perrottetia              Molokai
                               sandwicensis

Orthotylus perrottetiopsis     Perrottetia              Maui
                               sandwicensis

Orthotylus metrosideropsis     Metrosideros             Oahu
                               polymorpha

Orthotylus metrosideri         Metrosideros             Hawaii
                               polymorpha

Orthotylus sophoroides         Sophora chrysophylla     Maui

Orthotylus sophoricola         Sophora chrysophylla     Hawaii

Orthotylus erythrinae          Erythrina sandwicensis   Oahu

Orthotylus nestegiae           Nestegis sandwicensis    Hawaii

Orthotylus hibisci             Hibiscus arnottianus     Oahu

Orthotylus antidesmoides       Antidesma platyphyllum   Molokai

Orthotylus antidesmae          Antidesma platyphyllum   Hawaii

Orthotylus melicopoides        Melicope oahuensis       Oahu

Orthotylus melicopi            Melicope hawaiiensis     Hawaii

Orthotylus dubautiae           Dubautia raillardiodes   Kauai

Orthotylus dubauticola         Dubautia microcephala    Kauai

Orthtylus olapa                Cheirodendron trigynum   Oahu

Orthotylus broussaisiae        Broussaisia arguta       Kauai

Note: No definitive host plant associations have yet been
established for the following species: O. azalais Kirkaldy
(probably occurs on Metrosideros polymorpha), O. kekele
Kirkaldy, O. kassandra (Kirkaldy), O. kassandroides
Polhemus, O. kassandropsis Polhemus, and O. ulaula
Polhemus. The last four species appear to utilize many
host plants and may possibly be predaceous.


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Article Details
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Author:Polhemus, Dan A.
Publication:Entomologica Americana
Article Type:Report
Geographic Code:1U9HI
Date:Jan 1, 2013
Words:4366
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