Systematics of Pseudoloxops Kirkaldy (Insecta: Heteroptera: Miridae)

SYSTEMATICS OF PSEUDOLOXOPS KIRKALDY (INSECTA: HETEROPTERA: MIRIDAE)

Ratnawati Hazali BSc. (Hons). (Animal Resource Science and Management)

A thesis submitted in fulfillment of the requirements for the degree of Master of Science

In Systematic Entomology

School of Biological, Earth and Environmental Sciences Faculty of Science University of New South Wales

2013 PLEASETYPE THEUNIVERSIW OF NEWSOUTH WALES Thesis/DissertationSheet

Surnameor Familyname: HAZALI

Firstname: RATNAWATI Othername/s:

Abbreviationfor degreeas givenin the Universitycalendar: M.Sc.

School: Blologlcal, Earth and Environmental Sciences Faculty: Science

Title: Systematlcs of Pseudoloxops Kirkaldy (lnsecta: Heteroptera: Miridae)

Abdllct 450worda maxlmum:(PLEASE TYPE)

The ganus PseuddarcF Kkkaldy (lnsocia:Heteroptera: Miridae: Orthotylinae: O.thotylinl) cunently comprises38 described sp€cissfrom ths Palaearctic,Oriental, Alrotroplcal and Australianregions, as well as islandsin the PacificBasin (Schuh1995, 20Og), and many new species await d€scription. Pseudoloxopslacks a comprehensive and modern systematlc treatnent, All prwious sfudia8 of Psatrtobxops ue rcgiond in scopg, and most authors have not evaluatedthe generic limits of Psetdoloxops acro88its dlstrlbutlonalrange. This proj€ctaddrsssed this taxonomicimpediment and the genericlimits of Psa.rdolo(ops.This syst€maticsudy was basedon an examinationot sp€cim€nsobtained lrom museumcollections, as well as n€w collections.A comparativ€ morphological study based on €xtsrnal and male genitalia characters has been done to identify characters ol dlagno8dcand phylogeneticimportenc€. A phylogeneticanalysis of morphologicaldataset ot 53 taxa and 78 charact€rsis preesnt€d,The r€sults sho$, that the genlA Psaudola

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Date */::frt? ACKNOWLEDGEMENTS

First and foremost, I would like to thank Universiti Malaysia Sarawak (UNIMAS) and the Malaysian Ministry of Higher Education for granting me the scholarship for my graduate study at University of New South Wales. Deepest thanks to my supervisor,

Professor Gerry Cassis, without whose support, guidance and faith, this thesis would not have been completed. I’m definitely proud to be one of his students and grateful to him for accepting me as his student and educating me. I thank Professor Mike Archer for introducing me to Gerry. Without him I would not be in Prof. Cassis’ lab today.

There are many individuals who have been part of my project. Importantly, I would like to thank Celia Symonds, Hannah Finlay and Dr. Nik Tatarnic for their invaluable assistance from the beginning of this project. I appreciate all their time, expertise, guidance and friendship. Special thanks to my lab members particularly Fiona Powell,

Michael Elias, Anna Namyatova, Marina Cheng and Alex Brown. They have been great friends, providing an enjoyable work environment and support throughout this whole process.

I would like to acknowledge those from many institutions around the world who generously provided specimens for this study. I would also like to thank the Evolution

& Ecology Research Centre (E&ERC), School of Biological, Earth and Environmental

Sciences for the administrative and financial assistance given throughout the last semester of my study.

I give a special thank you to Professor Fatimah Hj. Abang for believing in me and for her continual support.

Finally, I would like to acknowledge my parents, Mr. Hazali Hj. Aman and Mrs. Rosiah

Sebli, and my siblings for their love and unwavering support. Special thanks go to

Wahap Marni for his constant encouragement. Last but not least, sincere thanks are given to all the many people who I have not listed here, but who have helped me in one way or another. I am very fortunate to have such wonderful and loving people around me.

ABSTRACT

The genus Pseudoloxops Kirkaldy (Insecta: Heteroptera: Miridae: Orthotylinae:

Orthotylini) currently comprises 38 described species from the Palaearctic, Oriental,

Afrotropical and Australian regions, as well as islands in the Pacific Basin (Schuh 1995,

2008), and many new species await description. Pseudoloxops lacks a comprehensive and modern systematic treatment. All previous studies of Pseudoloxops are regional in scope, and most authors have not evaluated the generic limits of Pseudoloxops across its distributional range. This project addressed this taxonomic impediment and the generic limits of Pseudoloxops. This systematic study was based on an examination of specimens obtained from museum collections, as well as new collections. A comparative morphological study based on external and male genitalia characters has been done to identify characters of diagnostic and phylogenetic importance. A phylogenetic analysis of morphological dataset of 53 taxa and 78 characters is presented. The results show that the genus Pseudoloxops is recognised as a monophyletic group based on synapomorphies of the head. The Southwest Pacific group of 20 species was selected for further systematic revision. A total of 19 new species are described including the one previously described species, Pseudoloxops vitiensis Kirkaldy. Habitus color photographs, illustrations of the male genitalia and distribution maps are provided.

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TABLE OF CONTENTS

ACKNOWLEDGEMENTS ……………………………………………………….. i

ABSTRACT ………………………………………………………………………. iii

LIST OF TABLES ………………………………………………………………… v

LIST OF FIGURES ……………………………………………………………….. vi

CHAPTER I. GENERAL INTRODUCTION …………………………………….. 1

CHAPTER II. PHYLOGENETIC ANALYSIS OF THE GENUS PSEUDOLOXOPS KIRKALDY (INSECTA: HETEROPTERA: MIRIDAE: ORTHOTYLINAE) ……………………………………………………………….. 7 Introduction …………………………………………………………………. 7 Materials and methods ………………………………………………………. 9 Character homology and terminology ………………………………………. 12 Phylogenetic results …………………………………………………………. 17 Discussion …………………………………………………………………… 28 References …………………………………………………………………... 31 Tables and Figures …………………………………………………………... 37

CHAPTER III. A SYSTEMATIC STUDY OF THE GENUS PSEUDOLOXOPS KIRKALDY IN THE SOUTHWEST PACIFIC ………..……………………….. 54 Introduction …………………………………………………………………. 54 Materials and methods ………………………………………………………. 56 Taxonomy …………………………………………………………………… 59 Checklist of Pseudoloxops of the Southwest Pacific (inclusive of Australia) ...... 64 Key to species of Pseudoloxops of the Southwest Pacific (inclusive of Australia) ……………………………………………………………………. 65 References …………………………………………………………………... 156 Tables and Figures …………………………………………………………... 163

CHAPTER 4. GENERAL CONCLUSIONS ……………………………………... 187

LIST OF TABLES

Table 2.1 Pseudoloxops characters and characters states 37

Table 2.2 Pseudoloxops character matrix. (-) inapplicable, (?) unknown 41

Table 3.1 Measurements of Pseudoloxops species of the Southwest Pacific 163

LIST OF FIGURES

Figure 2.1 Colour characters. a. P. baiawaensis. b. P. species #106. c. P. cordiformis. d. P. species #104. e. P. kraussi. f. P. solomonensis. g. P. lenakelensis. h. P. efatensis. i. P. species #83. Scale = 1 mm. 46

Figure 2.2 Pygophore characters, dorsal view: dorsal margin (DM), moderately to strongly concave (a., b. & e.) or weakly concave to straight (c., d. & f.); with left tergal process (ltp) (a.); with medial lobe (ml) (c. & d.); ventral margin (VM) projecting (a., c. - f.) or not projecting (b.); with ventral process (vp) (e. & f.); a. P. species #104, b. P. kimiensis, c. P. species #83, d. P. species #106, e. P. solomonensis, f. P. baiawaensis. Scale = 0.1 mm. 47

Figure 2.3 Left paramere characters, dorsal view: Y-shaped (a. - e., g.); sickle- shaped (f. & h.); apophysis (ap) of Y-shaped form parallel or ca. 45° to sensory lobe (sl) (a. - e.), or right angle to sensory lobe (g.); apophysis slim (a. - e.) or moderately broad to broad (f., g. & h.); shape of distal margin of apophysis bilobed (d.) or rounded (a. - c., e. - h.); with basal process (bp) (a., b., d. & e.); a. P. baiawaensis, b. P. tabwemasana, c. P. species #106, d. P. species #83, e. P. pancheriaphila, f. P. mcilwraithensis, g. P. straatmani, h. P. species #104. Scale = 0.1 mm. 48

Figure 2.4 Right paramere characters, dorsal view: simple C-shaped, without medial process (f.); club-shaped, without medial process (g.); with dorsal medial process only (dmp) (a. - e.); with ventral medial process only (vmp) (h.); apophysis (ap) with teeth (b., c., e., g., h.); medial process with teeth (a. - c., e., h.); a. P. species #104, b. P. species #83, c. P. naraiyawa, d. P. mcilwraithensis, e. P. pancheriaphila, f. P. species #106, g. P. baiawaensis, h. P. tabwemasana. Scale = 0.1 mm. 49

Figure 2.5 Phallotheca characters: a., b., f. & g., right lateral view; c. & d., dorsal view; e., left ventral view; opening, right-dorsolateral, from apical to medial (a., b., d. - g.) or dorsal, from apical to medial (c.); simple, without process (f. & g.); with ventral fold (vf), enlarged (a.) or reduced (b. & e.); with subapical lobe on dorsal surface (dsl) (a. - e.); with basal lobe (bl) on dorsal surface (c. & d.); a. P. species #83, b. P. pancheriaphila, c. P. species #104, d. and e. P. kraussi, f. P. species #106, g. P. tatarnici. Scale = 0.1 mm. 50

Figure 2.6 Aedeagus characters: a., c. & d., right lateral view; b., e. & f., ventral view; with one spicule (a.); with two spicules (b. & c.); with three spicules (d. & e.); with four spicules (f.); simple spicule form (unbranched, margins smooth) (a. - d, f.); complex spicule form (branched, margins serrate) (c. & e.); a. P. species #83, b. P. baiawaensis, c. P. species #104, d. P. species #106, e. P. mcilwraithensis, f. P. straatmani. Scale = 0.1 mm. 51 vi

Figure 2.7 Strict consensus cladogram of ten equally most parsimonious trees. 52

Figure 2.8 Implied weight cladogram of most parsimonious tree. Non- homoplasious synapomorphies are represented by filled circles. Homoplasious synapomorphies are shown as open circles. Bold numbers above nodes are bootstrap values. Numbers below nodes are jackknife values. 53

Figure 3.1 Habitus photographs of Pseudoloxops species of the Southwest Pacific. Plate 1 of 2. 168

Figure 3.2 Habitus photographs of Pseudoloxops species of the Southwest Pacific. Plate 2 of 2. 169

Figure 3.3 Male genitalia of Pseudoloxops baiawaensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; sl = sensory lobe of left paramere; vp = ventral process of pygophore; DM = dorsal margin of pygophore; VM = ventral margin of pygophore; DES = dorsal endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 170

Figure 3.4 Male genitalia of Pseudoloxops efatensis. a. Right paramere. b. Left paramere. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; sl = sensory lobe of left paramere. Scale = 0.1 mm. 171

Figure 3.5 Male genitalia of Pseudoloxops kimiensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; sl = sensory lobe of left paramere; vmp = ventral medial process of right paramere. Scale = 0.1 mm. 172

Figure 3.6 Male genitalia of Pseudoloxops kraussi. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, left ventral view. e. Phallotheca, dorsal view. f. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; bl = basal lobe of phallotheca; bp = basal process of left paramere; dsl = dorsal subapical lobe of phallotheca; dmp = dorsal medial process of right paramere; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 173

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Figure 3.7 Male genitalia of Pseudoloxops lenakelensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Aedeagus, left lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; dsl = dorsal subapical lobe of phallotheca; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 174

Figure 3.8 Male genitalia of Pseudoloxops mcilwraithensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, dorsal view. e. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; dmp = dorsal medial process of right paramere; DES1 = first dorsal endosomal spicule of aedeagus; DES2 = second endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 175

Figure 3.9 Male genitalia of Pseudoloxops naraiyawa. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; al = apical lobe of phallotheca; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; dsl = dorsal subapical lobe of phallotheca; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 176

Figure 3.10 Male genitalia of Pseudoloxops pancheriaphila. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dsl = dorsal subapical lobe of phallotheca; dmp = dorsal medial process of right paramere; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 177

Figure 3.11 Male genitalia of Pseudoloxops sikaiana. a. Pygophore, dorsal view. b. Right paramere, left lateral view. c. Right paramere, ventral view. d. Left paramere. e. Phallotheca, right lateral view. f. Phallotheca, dorsal view. g. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; sl = sensory lobe of left paramere; DES = dorsal endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 178

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Figure 3.12 Male genitalia of Pseudoloxops solomonensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; sl = sensory lobe of left paramere; vp = ventral process of pygophore; DES = dorsal endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 179

Figure 3.13 Male genitalia of Pseudoloxops straatmani. a. Pygophore, ventral view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; sl = sensory lobe of left paramere; vp = ventral process of pygophore; DES1 = first dorsal endosomal spicule; DES2 = second dorsal endosomal spicule; DES3 = third dorsal endosomal spicule; VES = ventral endosomal spicule; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 180

Figure 3.14 Male genitalia of Pseudoloxops tabwemasana. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, left lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dsl = dorsal subapical lobe of phallotheca; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; vmp = ventral medial process of right paramere; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 181

Figure 3.15 Male genitalia of Pseudoloxops tatarnici. a. Pygophore, dorsal view. b. Pygophore, dorsal view, with parameres and pygophore processes in situ. c. Right paramere. d. Left paramere. e. Phallotheca, right lateral view. f. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; dmp = dorsal medial process of right paramere; lp = left paramere; rp = right paramere; sl = sensory lobe of left paramere; DES1 = first dorsal endosomal spicule; DES2 = second dorsal endosomal spicule; DES3 = third dorsal endosomal spicule; VES = ventral endosomal spicule; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 182

Figure 3.16 Male genitalia of Pseudoloxops vitiensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, left lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; dsl = dorsal subapical lobe of phallotheca; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; vp = ventral process of pygophore; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 183 ix

Figure 3.17 Male genitalia of Pseudoloxops woodwardi. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, dorsal view. e. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 184

Figure 3.18 Distribution map of Pseudoloxops species of the Southwest Pacific. Map 1 of 2. 185

Figure 3.19 Distribution map of Pseudoloxops species of the Southwest Pacific. Map 2 of 2. 186

CHAPTER I

INTRODUCTION

The genus Pseudoloxops Kirkaldy (Heteroptera: Miridae: Orthotylinae: Orthotylini) belongs to the nominate tribe Orthotylini, a large group, where generic limits are poorly understood. A number of species rich genera, such as Orthotylus Fieber and

Pseudoloxops are examples where their generic definitions require attention, and there is also a significant taxonomic impediment. Pseudoloxops currently comprises 38 described species from the Palaearctic, Oriental, Australian and Afrotropical regions, and within island archipelagos in the Southwest Pacific (Schuh 1995, 2008).

Pseudoloxops lacks a comprehensive systematic treatment, without a monograph or revision. The morphological diagnosis of Pseudoloxops is based on external characters, and Yasunaga (1997, 1999) defined the genus on the following characters: 1) small and slender body; 2) densely pubescent body; 4) colour pattern of pale yellow, and partly sanguineous or scarlet dorsum often with dark spots; and, 5) anteriorly projected frons.

Linnnavuori (1994) also noted that the second antennal segment is longer than the remaining two segments combined. Yasunaga (1997, 1999) also remarked that some external diagnostic characters appear to be unique to Pseudoloxops, but that it is unlikely to be a monophyletic group because of the great variability in the male genitalia. He recommended a much broader survey of characters to define the genus.

The genus Pseudoloxops was originally described by Kirkaldy (1905), for the European species, P. coccineus (Meyer- Dür, 1843). Carvalho (1956) was the first to provide a key to the known species in his study on Micronesian species, which included four species. The most comprehensive and fundamental taxonomic work on Pseudoloxops was first published by Carvalho (1958), who treated 15 species. Post-Carvalho (1958) the most significant papers describing Pseudoloxops species are by Schuh (1974),

Carvalho (1980), Zou (1987), Liu and Zheng (1994), Linnavouri (1994) and Yasunaga

(1997). Most of these works are regional in scope and make little reference to the generic limits of Pseudoloxops or their possible relationships with other orthotyline genera. Schuh (1974) described a new species of genus Pseudoloxops from South

Africa, P. transvaalensis. Carvalho (1980) described a new species, P. lindrothi from

Sri Lanka. Zou (1987) described two new species from China: P. marginatus and P. guttatus. Liu and Zheng (1994) described an additional two new species from China: P. pilosus and P. punctulatus. Their study noted that P. punctulatus was closely allied to P. guttulatus Kerzhner and Muminov 1974 from Central Asia, differing in the shape of the male pygophore and parameres. Linnavouri (1994) recorded nine species of

Pseudoloxops from sub-Saharan Africa, with the following six described as new to science: P. niobe, P. thetis, P. nike, P. ninos, P. galateia and P. amfitrite. Yasunaga

(1997) revised the Japanese species of Pseudoloxops, and described three new species:

P. imperatorius, P. takaii and P. miyamotoi, and further documented a previously described species, P. miyatakei Miyamoto 1969.

Examination of collections from around the world have shown that a large number of new species require description. Also, many of the described species are poorly defined,

2 and are mostly recognised on colour characters alone. There has been little examination of the male genitalia in this genus.

Little is known of the biology of species of Pseudoloxops. Wheeler and Henry (1992) noted that the majority of Pseudoloxops species appear to be associated with broadleaf host plants. Their feeding habits are almost unknown, although predation was observed for the European species, P. coccineus (Meyer-Dür).

On the basis of current shortfalls in the knowledge of Pseudoloxops, my project had the following objectives:

1. To undertake a comparative morphological study of all described species and

new species provisionally assigned to Pseudoloxops;

2. To determine if the genus Pseudoloxops is a monophyletic group, and if so,

what are sister-group relationships within the genus, based on phylogenetic

analysis of morphological data; and,

3. To undertake a systematic revision of the species of Pseudoloxops from the

Southwest Pacific.

REFERENCES

Carvalho, J.C.M. (1956). Insects of Micronesia: Miridae. Bishop Museum, Honolulu, 7:

1-100.

Carvalho, J.C.M. (1958). A catalogue of the Miridae of the world. Part III. Arquivos do

Museo Nacional Rio de Janeiro, 47: 1-161.

Carvalho, J.C.M. (1980). A beautiful new species of blind bugs from Sri-Lanka:

Pseudoloxops lindrothi (Hemiptera: Miridae). Entomologia Generalis, 6 (2/4): 375-377.

Kerzhner, I.M. and Muminov N.N. (1974). New and little-known capsid bugs

(Heteroptera, Miridae) from Middle Asia. Izvestiya Akademii Nauk Tadzhikskoi SSR,

56: 42-46. (In Russian)

Kirkaldy, G.W. (1905). Neue und wenig bekannte Hemiptera. Wiener Entomologische

Zeitung, 24: 266-268.

Linnavuori, R.E. (1994). Orthotylinae of West, Central and North-east Africa

(Heteroptera, Miridae). Acta Zoologica Fennica, 193: 1-84.

Liu, G.Q. and Zheng, L.Y. (1994). Two new species of genus Pseudoloxops Kirk. from

China (Insecta: Heteroptera: Miridae). Reichenbachia, 30: 119-122.

Meyer-Dür, L.R. (1843). Verzeichnis der in der Schweiz einhimischen Rhynchoten

(Hemiptera Linn.). Erstes Heft. Die Familie der Capsini. Jent und Gassmann, Solothurn.

X + 11–116 + IV pp., 7 pls.

Schuh, R.T. (1974). The Orthotylinae and Phylinae (Hemiptera: Miridae) of South

Africa with a phylogenetic analysis of the ant-mimetic tribes of the two subfamilies for the world. Entomologica Americana, 47: 1-332.

Schuh, R.T. (1995). Plant bugs of the world (Insecta: Heteroptera: Miridae).

Systematic catalog, distributions, host list, and bibliography. New York: New York

Entomological Society, 1329 pp.

Schuh, R.T. (2008). On-line systematic catalog of plant bugs (Insecta: Heteroptera:

Miridae). (http://research.amnh.org/pbi/catalog).

Wheeler, A.G., Jr. and Henry, T.J. (1992). A synthesis of the Holarctic Miridae

(Heteroptera): Distribution, biology and origin, with emphasis on North America.

Lanham, MD: Entomological Society of America, V+282 pp.

Yasunaga, T. (1997). Species of the Orthotylinae genus Pseudoloxops Kirkaldy from

Japan (Heteroptera: Miridae). Bulletin of the Biogeographical Society of Japan, 52: 11-

18.

Yasunaga, T. (1999). The plant bug tribe Orthotylini in Japan (Heteroptera: Miridae:

Orthotylinae). Tijdschrift voor Entomologie, 142: 143-183.

Zou, H.G. (1987). Two new species of Pseudoloxops Kirk. from China (Hemiptera:

Miridae). Acta Zootaxonomica Sinica, 12: 389-392.

CHAPTER II

PHYLOGENETIC ANALYSIS OF THE GENUS PSEUDOLOXOPS KIRKALDY

(INSECTA: HETEROPTERA: MIRIDAE: ORTHOTYLINAE)

INTRODUCTION

The Orthotylinae is a subfamily of the Miridae (Insecta: Heteroptera), the largest family of true bugs comprising about 1,400 genera and over 10,000 described species (Cassis and Gross, 1995; Schuh, 1995; Wheeler, 2001). The subfamily Orthotylinae is divided into three tribes, comprising the cosmopolitan Halticini, the Afrotropical and Palaearctic

Nichomachini, and the largest tribe, Orthotylini. The mirid subfamilies Orthotylinae and

Phylinae comprise about 35% of all described species of plant bugs (Cassis et al.,

2007). The subfamily Orthotylinae is represented by about 220 genera (Schuh and

Slater, 1995). Most of the species in the Orthotylinae and Phylinae are phytophagous and highly host-plant specific (Cassis et al., 2007). The classification of the subfamily is based primarily on comparative studies of male and female genitalic characters (Slater,

1950; Kelton, 1959; Kerzhner and Konstantinov, 1999; Cassis, 2008; Forero, 2008) and the pretarsal structures (Schuh 1974, 1976).

Within the Orthotylini there are significant difficulties in establishing generic limits.

Many of the genera within the tribe are inadequately defined. The genus Pseudoloxops is one such genus, with many new species needing description. In the Orthotylini the male genitalia are critical in distinguishing genera and suprageneric groups. Schuh

(1974) in his study on South African Orthotylinae and Phylinae, proposed four generic

7 groups in the tribe Orthotylini comprising the Falconia, Orthotylus, Sericophanes, and

Zanchius groups. In the Zanchius group the aedeagus is more membranous and has no serrated spicules. In contradistinction the aedeagus of Orthotylus group has less membrane and many species have serrated spicules. However, there remain many taxa in the Orthotylini that do have serrated spicules, and separating the genera is confounded. Thus far, many recent studies have enhanced our understanding of the relationships of the genera within the Orthotylinae based on male genitalia (e.g. Schuh,

1974; Stonedahl and Schwartz 1986, 1988; Cassis 2008, 2010; Forero 2008, 2009a,

2009b; Schaffner and Schwartz, 2008). Cassis’ (2008) work in the Lattinova complex of the tribe Austromirini, proposed a new theory to homologise the endosoma and associated structures of the male genitalia, which is also relevant to studies of the

Orthotylini.

The genus Pseudoloxops consists of 38 species worldwide (Schuh 1995, 2008).

Currently, there is no comprehensive systematic treatment of the genus. All previous studies of Pseudoloxops are regional in scope, and most authors have not evaluated the generic limits of Pseudoloxops across its distributional range (Knight 1935, 1937;

Schuh, 1974; Carvalho, 1980; Zou, 1987; Linnavuori, 1994; Liu and Zheng, 1994;

Yasunaga 1997, 1999). Yasunaga (1997) defined Pseudoloxops based on external morphology and colouration, as follows: 1) small and slender body; 2) densely pubescent body; 3) dorsum pale yellow, and partly sanguineous or scarlet colouration, often with dark spots; and, 4) anteriorly projected frons.

Until now, there is no study that has been done to investigate the monophyly of the genus Pseudoloxops. The aim of this chapter is to conduct the first phylogenetic

8 analysis for the genus. To achieve this analysis I carried out a comparative morphological study of all described and new species provisionally assigned to

Pseudoloxops using a combination of external and male genitalia characters.

MATERIALS AND METHODS

Materials

This study is based on an examination of specimens borrowed from museum collections, as well as new collections. All specimens examined were given a unique specimen identifier (USI) and databased in the online Plant Bug Inventory (PBI) locality database (https://research.amnh.org/pbi/locality/).

In this study only male genitalic characters were treated in the analysis. Female genitalic character was not included in this study because of a lack of specimens. Where

I did not have specimens of described species, character information was coded from previous studies (Knight, 1937; Carvalho, 1956; Linnavuori, 1975, 1986, 1994;

Yasunaga, 1997).

The institutions providing specimens for this study are as follows:

AMNH American Museum of Natural History, New York

ANIC Australian National Insect Collection, CSIRO, Canberra

BMNH Natural History Museum, London

BPBM Bernice P. Bishop Museum, Honolulu

CAS California Academy of Sciences

MEM Mississippi Entomological Museum, Mississippi State University,

Mississippi

PPRI National Collection of Insects, South Africa

QDPI DPI Queensland, Queensland

SAMA South Australian Museum, Adelaide

USNM United States National Museum of Natural History, Smithsonian

Institution, Washington

UNSW University of New South Wales, Sydney

UQIC Insect Collection, University of Queensland, Brisbane

ZISP Zoological Institute, Russian Academy of Sciences

Methods

Comparative Morphology

A study of Pseudoloxops morphological characters followed the comparative methods outlined by Schuh and Brower (2009). Firstly, species were differentiated on the basis of character discontinuities of external and male genitalic characters, and morphometric data. Specimens were examined using a LEICA MZ125 microscope and LEICA

DM5000B compound microscope.

The dissections of the male genitalia were made following the methods described in

Kelton (1959). Terminology presented by Cassis (2008) was followed for male genitalia. Illustrations of the male genitalic structures were made using a camera lucida attached to a LEICA DM5000B compound microscope. These include the characters of the following male genitalic structures: pygophore, parameres, phallotheca and aedeagus.

Habitus photographs were taken of exemplar species of Pseudoloxops using the

Visionary Digital Micro-Imaging photographic system

(http://www.visionarydigital.com/), which uses a Canon 40D SLR attached to a K-2 macroscope. Multiple images were taken of male specimens of selected species and merged using the Helicon Focus software.

Phylogenetic methods

The phylogenetic analysis included 50 ingroup and three outgroup taxa. Of the 38 species currently placed in the genus Pseudoloxops, 13 species were included in the analysis. Twenty-five species were not included in the analysis because male specimens were not available for examination of genitalic characters. Fifteen new Pseudoloxops species out of 19 species described from the Southwest Pacific were included (Chapter

3). The remaining four new species have not been considered in the analysis because only females are known. Twenty-two undescribed species of the Pseudoloxops complex completed the ingroup (new species are either given voucher numbers, e.g. species #81, or the new species names from the Southwest Pacific given in Chapter 3).

Outgroup comparison method was used to root the trees and to determine character polarity (Farris, 1982; Nixon and Carpenter, 1993). Three Orthotylini species were used as outgroups: Orthotylus intratropica Symonds and Cassis, in preparation and O. graciliphilus Symonds and Cassis, in preparation, and one undescribed Orthotylini species. These outgroup were selected based on close relationships between them and

Pseudoloxops, and the availability of that group to this study. Most all of the genera of the tribe Orthotylini are very poorly defined.

A morphological matrix with a total of 78 characters was compiled using Nexus Data

Editor (Page, 2001). These included morphological characters of the head, thorax, colouration and the male genitalia. Fifty-six characters were binary and twenty-two characters were coded as unordered multistate. These morphological characters are coded from examination of specimens and as well as from data obtained from the literature where specimens were not available. Inapplicable characters were coded as

“-”, whereas characters with no information available were coded as “?”. Characters and character states are given in Table 2.1 and the data matrix in Table 2.2.

Parsimony analysis was conducted using TNT’s traditional searching method (Goloboff et al., 2003, 2008). Memory was set to allow a maximum of 10000 trees to be saved.

For Wagner trees, random seed was set from 1, performed at 1000 replications with branch swapping by tree bisection reconnection (TBR), with 10 trees saved per replication and replacement of existing trees. Implied weighting (Goloboff, 1993) using constant of concavity K = 3 was performed. Branch support was calculated in TNT using bootstrap values (Felsenstein, 1985) and jackknife (Lanyon, 1985). Resamplings for both measures were run at 1000 random addition replicates. Characters distributions were examined using MacClade 4.0.8 (Maddison and Maddison, 2005).

CHARACTER HOMOLOGY AND TERMINOLOGY

Colour patterns

A total of 31 characters of colour patterns were used in the phylogenetic analysis. Based on my examination of all the Pseudoloxops species available in collections, I found substantial variation in colour patterns. Examples of colour patterns observed are

12 presented in Figure 2.1. In the majority of the Pseudoloxops species, the ground colouration is pale yellow to yellowish brown with the presence of either red, reddish- brown or orange-red markings (Fig. 2.1) and sometimes with dark brown spots on the dorsal surface (character 12-1) (Fig. 2.1b.). The head colouration is mostly pale yellow to yellowish brown (Figs. 2.1a. - e., g. - i.), sometimes with dark brown (Fig. 2.1f.), and usually with red to reddish-brown markings on the frons, vertex, clypeus, mandibular plate and maxillary plate (13-1; 14-1; 15-2; 16-2).

The pronotum ground colour is either pale yellow to yellowish brown (19-0) (Figs. 2.1a.

- e., g. - i.), greenish yellow, or dark brown (Fig. 2.1f.) with red or reddish-brown markings on the lateral areas (19-1) (Figs. 2.1a. - d.) or on both anterior and lateral areas

(Figs. 2.1e., f.). Some species have an entirely pale yellow to yellowish brown (19-0)

(Fig. 2.1h.), dark brown with red markings (19-1) (Figs. 2.1f., i.) or almost red pronotum (19-2). The proepimeron and proepisternum are generally pale yellow to yellowish brown, and usually with red markings. The mesoscutum is generally pale yellow to yellowish brown (23-0) (Figs. 2.1e., g., h.) or bicoloured with red markings on the anterolateral angles (24-1) (Figs. 2.1a. - d., f., i.), posterolateral angles (25-1)

(Fig. 2.1d.), and medial part (26-1) (Figs. 2.1d., f., i.). Some species have mostly a red or orange-red mesoscutum (23-1). The colour of the scutellum shows considerable variation, being either entirely pale yellow to yellowish brown, with red markings (Figs.

2.1a., d. - f., i.) or mostly to entirely red (Figs. 2.1b., c., g., h.).

The hemelytra are generally pale yellow to yellowish brown or sometimes greenish pale yellow. The colouration of the clavus varies from entirely pale yellow to yellowish brown (28-0), with red markings (29-1), or mostly to entirely red (28-1). For species

13 where the clavus has red markings, most species have the red markings on the basal and apical regions (30-2) (Figs. 2.1b., c., e.). For some species the red patterning is restricted either to the basal (30-0) (Figs. 2.1a., i.) or apical regions (30-1) (Figs. 2.1g., h.). There are a few species with scattered small red spots on the clavus (30-3) (Fig.

2.1d.). The colour of the corium is generally pale yellow brown to yellowish brown (31-

0), either with red (32-1) or without red patterning (32-0). For those with red markings, it is either mostly or entirely red on the exocorium (33-0) (Fig. 2.1a.), mostly on the endocorium, or adjacent to the clavus (33-1) (Fig. 2.1e.), mostly on the exocorium and medial areas adjacent to the clavus (33-2) (Figs. 2.1b., c.), mostly on the basal and posterior areas (33-3) (Fig. 2.1i.), on the basal part only (33- 4) (Fig. 2.1f.) or with minor red markings (33-5) (Figs. 2.1g., h.). The presence of red markings on the apex of the embolium is also defining for species (34-1) (Figs. 2.1d., h.). The cuneus is entirely pale yellow to yellowish brown (35-0), mostly or entirely red to dark red (35-1) (Figs. 1.2a., b., d.) or bicoloured (35-2) (Figs. 2.1c., e. - i.). For species with a bicoloured cuneus, it either has red markings only on the apex (Fig. 2.1c.) or medial basal angle, or sometimes both (Figs. 1.2e. - i.). The legs are generally pale yellow brown to yellowish brown, sometimes with a narrow red to reddish-dark brown band on the apex of the metafemora (42-0) or with broad red to reddish-dark brown markings on the distal half of the metafemora (42-1) (Fig. 1.2d.).

Structure

Forty-seven structural characters were used in the phylogenetic analysis. Ten characters were based on head morphology and a single character of the pronotum was coded. A total of thirty-six characters of the male genitalia were documented.

The head characters are phylogenetically important in defining the genus Pseudoloxops as a monophyletic group. These included the flat or nearly flat head in dorsal view (1-

0), the convex frons as seen in lateral view (3-0), and the greatly tumescent maxillary plate (5-1). The presence of a sulcus on the midline of the vertex was also informative.

The pronotum is variable in structure, with species having either a rectilinear (11-0), excavate (11-1) or weakly bisinuate (11-2) posterior margin.

The male genitalia in Pseudoloxops are highly variable. Six characters of the pygophore were found to be important. These are illustrated in Figure 2.2, and include the shape of the dorsal margin, and as to whether it is moderately to strongly concave (43-1) (Figs.

2.2a., b. & e.) or weakly concave to straight (43-0) (Figs. 2.2c., d. & f.); the presence of the left tergal process on the dorsal margin (45-1) (Fig. 2.2a.); the presence of a medial lobe on the dorsal margin (44-1) (Figs. 2.2c. & d.); the shape of the ventral margin, whether projecting (47-1; 47-2) (Figs. 2.2a., c. - f.) or not projecting (47-0) (Fig. 2.2b.); and, the presence of a right ventral process (48-1) (Figs. 2.2e. & f.).

Twelve important characters were found relating to the structure of the left and right parameres. The left paramere in Pseudoloxops was found to be generally either Y- shaped (49-1) (Figs. 2.3a. - e. & g.) or sickle-shaped (49-0) (Figs. 2.3f. & h.). The variations that were considered to be informative included: whether the apophysis of the

Y-shaped form is either parallel, ca. 45° to the sensory lobe (53-0) (Figs. 2.3a. - e.), or at right angles to the sensory lobe (53-1) (Fig. 2.3g.); the presence of a slim apophysis

(52-0) (Figs. 2.3a. - e.) or moderately broad to broad apophysis (52-1) (Figs. 2.3f., g. & h.); whether the shape of the distal margin of the apophysis is bilobed (51-0) (Fig. 2.3d.)

15 or rounded (51-1) (Figs. 2.3a. - c., e. - h.); and, the presence of a basal process (54-1)

(Figs. 2.3a. - b., d. - e.).

The right parameres in Pseudoloxops show a greater amount of variation in shape compared to the left parameres. Variation in the shape of right paramere is shown in

Figure 2.4. Variants include: a simple C-shaped form, with the absence of a medial process (55-0) (Fig. 2.4f.); a club-shaped form, without a medial process (55-1) (Fig.

2.4g.); a C-shaped form with the presence of a dorsal medial process only (55-2) (Figs.

2.4a. - e.); a C-shaped form with a ventral medial process only (55-3) (Fig. 2.4h.); the presence of teeth on the apophysis (56-1; 57-1; 58-1) (Figs. 2.4b. - c., e., g. - h.); and, the presence of teeth on the medial process (59-1; 60-1) (Figs. 2.4a. - c., e. & h.). The right paramere of species #80, #80A and #81 are different compared to the other species in Pseudoloxops, with the presence of both dorsal and ventral medial processes (55-4).

Eleven characters of the phallotheca were used in the phylogenetic analysis. The characters included the opening of the phallotheca, whether it is apical only (62-0), right-dorsolateral from apical to medial (62-1) (Figs. 2.5a., b., d. - g.) or dorsal from apical to medial (62-2) (Fig. 2.5c.); the phallotheca without processes (Figs. 2.5f. & g.); with a large ventral fold (65-1) (Fig. 2.5a.) or small ventral fold (65-0) (Figs. 2.5b. & e.); the presence of a subapical lobe on the dorsal surface (67-1) (Figs. 2.5a. - e.); and, the presence of a basal lobe on the dorsal surface (71-1) (Figs. 2.5c. & d.).

The aedeagus in Pseudoloxops is highly diverse, particularly in relation to the number of endosomal spicules, varying from one to four, as shown in Figure 2.6. The marginal surface of species with a single endosomal spicule is smooth whereas for species with

16 multiple endosomal spicules the margin can be smooth or serrate. Seven characters proved to be important in the phylogenetic analysis. These included the number of spicules (cf. one spicule (72-0) - Fig. 2.6a.; two spicules (72-1) - Figs. 2.6b. & c.; three spicules (72-2) - Figs. 2.6d. & e.; and, four spicules (72-3) - Fig. 2.6f.). For species with a single spicule, the following additional characters were informative: the presence or absence of a basally folded aedeagus; whether the spicule is elongate (about five times longer than wide) (74-1) or not elongate (about as long as wide) (74-0); whether the spicule is straight (75-0) or recurved (75-1); and, whether the spicule encloses (76-0) or is distal to the secondary gonopore (76-1). For those species with multiple spicules, the following characters were informative: presence or absence of at least one simple spicule (unbranched and margins smooth) (77-1) (Figs. 2.6b., c., d. & f.) and the presence or absence of at least one complex spicule (branched and margins serrate) (78-

1) (Figs. 2.6c. & e.).

PHYLOGENETIC RESULTS

The analysis of the morphological data matrix (Table 2.2) produced ten equally parsimonious trees with a length of 388 steps, a consistency index of 0.27 and a retention index of 0.59. The strict consensus tree is shown in Figure 2.7. Implied weighting using constant concavity K = 3 resulted in one most parsimonious tree

(Figure 2.8), to which the characters were optimised. Non-homoplasious synapomorphies are shown as filled circles. Homoplasious synapomorphies are shown as open circles. Bold numbers above nodes indicate bootstrap values. Numbers below nodes indicate Jackknife values. Only Bootstrap and Jackknife values greater than 50% are shown on the tree obtained (Fig. 2.8).

Character support for all nodes in Figure 2.8 is discussed below. The characters and character states are given in Table 2.1. The data matrix is given in Table 2.2.

Node 1. The genus Pseudoloxops is defined by two non-homoplasious synapomorphies: head flat or nearly flat (character 1-0) and frons convex in lateral view

(3-0). This node is also supported by five homoplasious characters: the maxillary plate greatly tumescent medially (5-1) (which is not greatly tumescent medially in

Pseudoloxops baiawaensis and species #53), the frons with red markings (13-1) (absent in P. tabwemasana, P. solomonensis, species #09, #14, #94, #97, #98), the bicoloured clypeus (15-1), the apex of scutellum with orange-red to dark red markings (27-0) and, the metafemora apically with red to reddish-brown markings (41-1). This node is supported by a moderate bootstrap value of 55 and a Jackknife value of 67.

Based on this phylogeny, the genus Pseudoloxops is recognised as a monophyletic group. This is most strongly supported by the abovementioned head synapomorphies, which are also diagnostic for the genus, with the frons flat or nearly flat, and convex in lateral view. This head structure is unique in comparison to other genera in the tribe

Orthotylini.

Node 2. This node is supported by five homoplasious characters: vertex without red markings or concolorous with remainder of head (14-0); thoracic sterna unicolorous or without red markings (22-0); the presence of red markings on apex of embolium (34-1); costal fracture with red markings (38-1); and, the abdomen unicolorous or without red markings (39-0).

Node 3. This node is supported by the following characters: weakly visible mandibular plate in dorsal view (4-0); midline of vertex moderately to deeply sulcate (6-1); the bicoloured mesoscutum (23-3); and, the presence of broad red band on apical half of metafemora (42-1).

Node 4. This node is defined by three homoplasious characters: the bicoloured mandibular plate (16-1); cuneus colouration mostly or entirely red to dark red (35-1); and, the presence of Y-shaped left paramere (49-1).

Node 5. This node is supported by a single non-homoplasious synapomorphy, the presence of red markings on anterolateral angles of bicoloured mesoscutum (24-1) and two homoplasious: the moderately to strongly projected frons anteriorly (2-1) and the bicoloured second antennal segment (18-1).

Node 6. There are two major clades at this node. Together they are defined by the presence of Y-shaped left paramere with the apophysis parallel or 45° to sensory lobe

(53-0).

Node 7. This clade, consisting of two subclades (P. amfitirite, species #76, #62, P. amabilis and P. nike, P. sudanicus, P. niobe, #104, #53, P. coccineus), is supported by one non-homoplasious synapomorphy: the absence of teeth on apophysis of the C- shaped right paramere (without medial process) (58-0).

Node 8. This clade is comprised of four species (P. amfitrite, species #76, #62, P. amabilis) and is defined by the bicoloured mesoscutum, with the absence of orange to red markings on the medial part of mesoscutum (26-0).

Node 9. This clade is supported by the presence of a weakly Y-shaped left paramere

(50-0).

Node 10. In this clade, species #62 and P. amabilis are more closely related to each other, with species #76 sister to them. The monophyly of this clade is based on two characters: the absence of red markings on vertex or concolorous with remainder of head (14-0) and the apex of the scutellum pale yellow to yellowish brown (27-1).

Node 11. This clade is comprised of six species (P. nike, P. sudanicus, P. niobe, species

#104, species #53 and P. coccineus). It is supported by the following characters: the presence of dark brown spots on dorsal surface (12-1); the bicoloured mesoscutum with the presence of red markings on posterolateral angle (25-1); and, the aedeagus with two endosomal spicules (72-1).

Node 12. This clade is defined by having orange or red markings on the abdominal venter (39-1).

Node 13. This clade is supported by four homoplasious characters: the posterior margin of pronotum rectilinear (11-0); the yellowish brown or unicolorous second antennal segment (18-0); the presence of orange or red markings on thoracic sterna (22-1); and, the presence of scattered small red spots on the clavus (30-3).

Node 14. The species within this clade share the presence of a sickle-shaped left paramere (49-0).

Node 15. In this clade, species #53 and P. coccineus are more closely related to each other, with species #104 from China the sister-group of the latter two species. The sister-group relationship for species #53 and P. coccineus is based on two homoplasious characters: the absence of dark brown spots on dorsal surface (12-0) and the mostly red orange to red mesoscutum (23-1).

Node 16. This node is supported by the presence of a slim apophysis of the left paramere (52-0).

Node 17. This clade is supported by four homoplasious characters: the absence of red markings on the vertex or concolorous with remainder of head (14-0); the bicoloured cuneus (35-2); the absence of red markings on the costal fracture or concolorous with the remainder of the corium (38-0); and, the right paramere in the C-shaped form, with a dorsal medial process only (55-2).

Node 18. This clade is comprised of four species from the Southwest Pacific (P. solomonensis, P. sikaiana, P. cordiformis and P. baiawaensis). The species in this clade share the following two homoplasious characters: the presence of right ventral process in pygophore (48-1) and the aedeagus with two endosomal spicules (72-1).

Node 19. This clade is defined by the presence of a basal process on left paramere (54-

1).

Node 20. This clade indicates a sister-relationship of P. cordiformis and P. baiawaensis.

Both species have a bicoloured mesoscutum, with the absence of orange to red markings on the medial part (26-0).

Node 22. This node is supported by the presence of a medial lobe on the dorsal margin of the pygophore (44-1).

Node 23. This clade is supported by six homoplasious characters: the mostly or entirely red to reddish-brown mandibular plate (16-2); the pale yellow to yellowish-brown apex of the scutellum (27-1); the left paramere with basal process (54-1); the phallotheca with a complex process (61-1); the presence of a ventral fold on the phallotheca (64-1); and, the aedeagus with a single endosomal spicule.

Node 24. This node is supported by the presence of a subapical lobe on the dorsal surface of the phallotheca (67-1).

Node 25. Six species (P. signatus, P. pancheriaphila, P. kraussi, species #80A, #80 and

#81) form a group, which is supported by a single homoplasy, the mostly or entirely red to orange corium (31-1).

Node 26. Two homoplasious characters define this clade of five species: the bicoloured mesoscutum with the presence of orange to red markings on the medial part (26-1) and the presence of a subtriangular fold on the ventral surface of the phallotheca (66-2).

Node 27. This node is supported by one homoplasious character, the presence of a basal lobe on the dorsal surface of the phallotheca (71-1).

Node 28. Species #80A, species #80 and species #81 form a clade which is supported by a single non-homoplasious synapomorphy, the presence of two medial processes on the C-shaped right paramere (55-4). This node is also supported by six homoplasious characters: the vertex with a moderately carinate posterior margin (7-2); the presence of red markings on the vertex (14-1); the presence of orange or red markings on the abdomen (39-1); the weakly Y-shaped left paramere (50-0); the moderately broad to broad apophysis of the left paramere (52-1); and, the aedeagus with a single spicule, which is distal to the secondary gonopore (76-1).

Node 29. Species #80A and species #80 are sister-species and is supported by one non- homoplasious synapomorphy, the single endosomal spicule with the presence of basal fold (73-1).

Node 30. The following characters support this clade: the eyes of male not reaching the ventral margin of the head in lateral view (9-0) and the absence of reddish markings at the apex of the embolium (34-0).

Node 31. Two homoplasious characters support this clade: the bicoloured mandibular plate (16-1) and the presence of an large fold on the ventral surface of the phallotheca

(65-1).

Node 32. This clade comprises three species from Vanuatu: P. tabwemasana, P. lenakelensis and P. efatensis. It is supported by a single non-homoplasious synapomorphy: the pale to yellowish-brown clavus, with the presence of red markings only on the apical part (30-0). It is also supported by one homoplasy, the pale yellow or yellowish-brown mesoscutum (23-0).

Node 33. P. lenakelensis and P. efatensis are sister-species, which is supported by two homoplasious characters: the eyes in male reaching ventral margin of head in lateral view (9-1) and the presence of medial lobe on dorsal margin of pygopohore (44-0).

Both species are more closely related to each other than with P. tabwemasana.

Node 34. This node, is comprised of two major subclades and supported by a single homoplasious character, the weakly anteromedially projected frons (2-0).

Node 35. This clade of six species (species #99, P. naraiyawa, P. vitiensis, species #07,

P. adamsoni, P. rubroclavus) is supported by one non-homoplasious synapomorphy, the presence of an large single subapical lobe on the dorsal surface of the phallotheca (69-

1).

Node 36. This clade of three species (species #99, P. naraiyawa, P. vitiensis) is defined by three homoplasious characters: the eyes in the male not reaching the dorsal margin of the head in lateral view (8-0); the mostly or entirely red to orange corium (31-1); and, the aedeagus with a somewhat straight, single endosomal spicule (75-0).

Node 37. This clade documents the sister-group relationship of P. naraiyawa and P. vitiensis (with species #99 sister to them). The clade is defined by the following homoplasious characters: the moderately to strongly projected frons (2-1); the yellowish brown or unicolorous second antennal segment (18-0); and, the pale yellow or yellowish-brown mesoscutum (23-0).

Node 38. Species #07, P. adamsoni and P. rubroclavus form a clade, which is supported by two homoplasious characters: the mostly or entirely red clypeus (15-2) and the mostly or entirely red to reddish-brown mandibular plate (16-2).

Node 39. P. adamsoni and P. rubroclavus are sister-species, which is supported by one homoplasy, the eyes in males reaching the ventral margin of the head in lateral view (9-

1).

Node 40. This clade of eleven species is supported by the following homoplasious characters: the presence of red markings on the vertex (14-1); the bicoloured mesoscutum with the presence of orange to red markings on medial part (26-1); and, the presence of a sinuous or bilobed distal margin of the apophysis of the left paramere (51-

0).

Node 41. This clade forms a sister-group relationship between species #83 and P. palauensis. The clade is supported by four homoplasious characters: the eyes in the male reaching the ventral margin of the head in lateral view (9-1); the mostly yellowish- brown to brown corium, with the presence of red markings, mostly on basal and

25 posterior areas (33-3); the presence of reddish markings on the apex of the embolium

(34-1); and, the presence of red markings on the costal fracture (38-1).

Node 42. This clade comprises nine species of Pseudoloxops from French Polynesia. It is supported by one homoplasy, the presence of narrow red band markings on the apical part of the metafemora (42-0).

Node 43. This clade is defined by the presence of two non-homoplasious synapomorphies: the pale yellow to yellowish-brown clavus, without a red marking (29-

0) and the mostly yellowish-brown to brown corium, without a red marking (32-0). This clade is also supported by the following homoplasious characters: the yellowish-brown or unicolorous second antennal segment (18-0); the pale yellow or yellowish-brown mesoscutum (23-0); the bicoloured cuneus, with the absence of red to dark red markings on the medial base angles (36-0).

Node 44. Species #90, #11 and #102 form a clade, which is not supported by any non- homoplastic synapomorphy. Two homoplasious characters support it: the weakly Y- shaped left paramere (50-0) and the presence of an apical lobe on the right dorsal opening of the phallotheca (63-1). The clade has a strong bootstrap support of 77% and

Jackknife value of 82%.

Node 45. Species #11 and species #102 are more closely related to each other than to species #90. Their sister-group relationship is based on one homoplasy, the presence of a single subapical lobe on the dorsal surface of the phallotheca, with a bilobed margin.

This clade is supported by significant bootstrap (53%) and Jackknife values (54%).

Node 46. This clade of five species (species: #09, #14, #94, #98, #97) is defined by the following characters: the eyes in the male not reaching the dorsal margin of the head in lateral view (8-0); the frons without red markings or concolorous with the remainder of the head (13-0); the vertex without red marking or concolorous with the remainder of the head (14-0); the yellowish-brown to brown clypeus (15-0); the yellowish-brown mandibular plate (16-0); the unicolorous or pale yellow to yellowish-brown cuneus (35-

0); the pale or not red forewing membrane veins (40-0); and, the presence of a sinuous or bilobed distal margin of the apophysis of the left paramere (51-0).

Node 47. This clade is supported by three homoplasious characters: the mostly or entirely yellow-brown or dark brown pronotum (19-0); the proepimeron unicolorous or without red marking (20-0); and, the proepisternum unicolorous or without red markings (21-0).

Node 48. This clade is comprised of three species from the Tahiti Islands: species #94,

#98 and #97. Two homoplasious supports this clade: the presence of a ventral fold on the phallotheca, small in size (65-0) and the ventral fold on the phallotheca with a rounded margin (66-0).

Node 49. This clade indicates a sister-group relationship of species #97 and #98, which together are most closely related to species #94. Two homoplasious characters support their sister-group relationship: the eyes in the male reaching the dorsal margin of the head in lateral view (8-0) and the C-shaped right paramere with a ventral medial process only (55-3).

DISCUSSION

In the phylogenetic analysis presented above, the genus Pseudoloxops is a monophyletic group and is strongly supported by two non-homoplasious synapomorphies: the head flat or nearly flat, and with the frons convex in lateral view; and four homoplasious characters: the maxillary plate greatly tumescent medially, the frons with red markings, the apex of scutellum with orange-red to dark red markings, and the metafemora apically with red to reddish-brown markings. In combination, these characters differentiate Pseudoloxops from other described orthotyline genera.

The position of the five most basal Pseudoloxops species (P. miyamotoi, P. mcilwraithensis, P. kimiensis, P. straatmani and species #66) is probably affected by the fact that most of the male genitalia characters were either not coded because of a lack of male specimens available for study or the genitalia were highly autapomorphic

(P. mcilwraithensis).

Eight nodes are supported by at least one non-homoplasious synapomorphy (Fig. 2.8).

However, from the eight nodes, only one clade (Node 1) has greater than 50% bootstrap and Jackknife support. There are two other clades, which have significant bootstrap and

Jackknife values (> 50%) (Nodes 44, 45), with both clades supported by homoplasious synapomorphies only. The monophyly of species: #90, #11 and #102 (Node 44) is the most supported in the analysis, which is defined by the weakly Y-shaped left paramere, and the presence of an apical lobe on the right dorsal opening of the phallotheca, as well as strong resampling support. The sister-group relationship of species #11 and species

#102 (Node 45) is supported by the presence of a single subapical lobe on the dorsal surface of the phallotheca, with a bilobed margin.

Most studies of the genus Pseudoloxops use external characters such as colour patterns and male genitalic characters to define and separate the species (e.g., Knight 1937;

Carvalho 1956, 1980; Carvalho and Wallerstein 1976; Linnavuori 1975, 1994; Liu and

Zheng 1994; Linnavuori 1975, 1994; Yasunaga 1997, 1999). Although most of the characters examined in this study were homoplasious, the external characters, especially the colour characters, were phylogenetically significant and in many cases corroborated by male genitalic characters, particularly those of the parameres. Areekul and Quicke

(2006) suggested that colour characters should be used more in phylogeny based on previous studies done by Gosliner and Johnson (1999).

In terms of distribution, this analysis shows that some of the clades are geographically restricted. For example, 14 species from French Polynesia form a monophyletic group

(Node 34). This clade is supported by the frons being weakly anteromedially projected.

However, two species from Fiji, P. naraiyawa and P. vitiensis Kirkaldy form a group with species #99 from Tahiti. This group is defined by the presence of eyes in the male not reaching the dorsal margin of the head in lateral view, the mostly or entirely red to orange corium and the aedeagus with a somewhat straight, single endosomal spicule.

Species from Asia, Africa and Palearctic form a group which is represented by 10 species in my analysis. It is supported by a single non-homoplasious synapomorphy, the absence of teeth on apophysis of the C-shaped right paramere. However, one species from South Africa, species #106, was placed together with species from islands in the

Pacific Basin and Australia. This analysis also shows that the species of the Southwest

Pacific Islands are more closely related to each other.

The following observational work and materials would test the phylogeny and systematics of the genus Pseudoloxops presented in this study:

1) Broader investigation of orthotyline taxa, which will help to test the current

hypothesis of Pseudoloxops monophyly and relationships with other orthotyline

genera.

2) More collecting of specimens in the island archipelagos of the Pacific Basin to

expand the limited materials than those that were available in my study.

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Table 2.1. Pseudoloxops characters and character states.

1. Head dorsal: (0) flat or nearly flat; (1) swollen/rounded. 2. Frons - dorsal view: (0) weakly projected anteriorly (clypeus visible); (1) moderately to strongly projected anteriorly (clypeus not visible). 3. Frons - lateral view: (0) convex, strongly ventrally oriented; (1) declivent, sloping gently forward. 4. Mandibular plate: (0) barely to slightly visible in dorsal view; (1) greatly visible in dorsal view. 5. Maxillary plate: (0) not swollen medially; (1) tumescent medially. 6. Midline of vertex: (0) weakly sulcate (at most as a weak line without depression); (1) moderately to deeply sulcate (with line depressed). 7. Vertex - posterior margin: (0) rounded; (1) weakly carinate, not sharp; (2) moderately carinate, not sharp. 8. Eyes in lateral view - male: (0) not reaching dorsal margin of head; (1) reaching dorsal margin of head. 9. Eyes in lateral view - male: (0) not reaching ventral margin of head; (1) reaching ventral margin of head. 10. Bucculae: (0) not reaching posterior margin of head; (1) reaching posterior margin of head. 11. Posterior margin of pronotum: (0) rectilinear; (1) excavate; (2) weakly bisinuate. 12. Dorsal colouration - dark brown spots: (0) absent; (1) present. 13. Frons - colouration: (0) without red markings or concolorous with remainder of head; (1) with red markings. 14. Vertex - colouration: (0) no reddish mark or concolorous with remainder of head; (1) with red markings. 15. Clypeus - colouration: (0) yellowish brown to brown; (1) bicoloured; (2) mostly or entirely red. 16. Mandibular plate - colouration: (0) yellowish brown; (1) bicoloured, with minor red highlighting; (2) mostly or entirely red to reddish brown. 17. Antennal segment I - colouration: (0) yellowish brown; (1) bicoloured, with minor red highlighting; (2) mostly or entirely red to reddish brown. 18. Antennal segment II - colouration: (0) yellowish brown or unicolorous; (1) bicoloured, with minor red highlighting; (2) mostly or entirely red. 19. Pronotum - colouration: (0) mostly or entirely yellow brown or dark brown; (1)base colour greenish-yellow brown to dark brown with red to reddish brown markings; (2) mostly or entirely red; (3) greenish-yellow brown. 20. Proepimeron - colouration: (0) unicolorous or without red markings; (1) with red markings at dorsal part or mostly. 21. Proepisternum - colouration: (0) unicolourous or without red markings; (1) with red markings. 22. Thoracic sterna - colouration: (0) unicolourous or without red markings; (1) with orange or red markings; (2) greenish.

Table 2.1. Cont/-.

23. Mesoscutum - colouration: (0) pale yellow/yellowish brown; (1) mostly red/orange to red; (2) mostly greenish-yellow brown; (3) bicoloured. 24. Mesoscutum - anterolateral angle - colouration: (0) without red markings; (1) with red markings. 25. Mesoscutum - posterolateral angle - colouration: (0) without red markings; (1) with red markings. 26. Mesoscutum - medial section - colouration: (0) without orange to red markings; (1) with orange to red markings. 27. Apex of scutellum - colouration: (0) orange-red to dark red; (1) pale yellow to yellowish brown. 28. Clavus - colouration: (0) pale yellow to yellowish brown; (1) mostly or entirely red; (2) mostly or entirely greenish-pale yellow. 29. Clavus - colouration - pale yellow to yellowish brown form: (0) without red marking; (1) with red markings. 30. Clavus - pale yellow to yellowish brown form, with red markings: (0) basal only; (1) apical only; (2) basally and apically; (3) with scattered small red spots. 31. Corium - colouration: (0) yellowish brown to brown; (1) mostly or entirely red to orange; (2) mostly or entirely green. 32. Corium - colouration - for mostly yellowish brown to brown form: (0) without red markings; (1) with red markings. 33. Corium - for mostly yellowish brown to brown form, with red markings: (0) mostly or entirely on exocorium; (1) mostly on endocorium or adjacent to clavus; (2) mostly on exocorium and inner areas adjacent to clavus; (3) mostly on basal and posterior areas; (4) basal only; (5) minor red markings, not as above. 34. Apex of embolium - colouration - reddish: (0) absent; (1) present. 35. Cuneus - colouration: (0) unicolorous or pale yellow to yellowish brown; (1) mostly or entirely red to dark red; (2) bicoloured, with minor red highlighting. 36. Cuneus bicoloured - inner base red to dark red: (0) absent; (1) present. 37. Cuneus bicoloured - outer base red to dark red: (0) absent; (1) present. 38. Costal fracture - colouration: (0) without red markings or concolorous with remainder of corium; (1) with red markings. 39. Abdomen - colouration: (0) unicolourous or without red marking; (1) with orange or red markings; (2) greenish. 40. Forewing membrane veins - colouration: (0) pale or not red; (1) red. 41. Metafemora - colouration - apically red to reddish dark brown: (0) absent; (1) present. 42. Metafemora - colouration - those with red markings: (0) narrow band at apex; (1) broad band, on apical half. 43. Pygophore - dorsal margin shape: (0) weakly concave to straight; (1) moderately to strongly concave.

Table 2.1. Cont/-.

44. Pygophore - medial lobe, on dorsal margin: (0) absent; (1) present. 45. Pygophore - left tergal process on dorsal margin: (0) absent; (1) present. 46. Pygophore - right tergal process on dorsal margin: (0) absent; (1) present. 47. Pygophore - ventral margin shape: (0) not projecting, straight or slightly convex; (1) projecting, convex, sometimes weakly sinuous; (2) projecting, subtriangular, apex acute. 48. Pygophore - right ventral process: (0) absent; (1) present. 49. Left paramere - general shape: (0) sickle-shaped; (1) Y-shaped. 50. Left paramere - Y-shaped: (0) weakly Y-shaped; (1) deeply Y-shaped. 51. Left paramere - shape of distal margin of apophysis: (0) sinuous or bilobed; (1) rounded. 52. Left paramere - apophysis: (0) slim; (1) moderately broad to broad. 53. Left paramere - apophysis of Y-shaped form: (0) parallel to sensory lobe or ca. 45° to sensory lobe; (1) right angle to sensory lobe. 54. Left paramere - basal process: (0) absent; (1) present. 55. Right paramere - general shape: (0) simple C-shaped, without medial process; (1) club-shaped, without medial process; (2) C-shaped, with dorsal medial process only; (3) C-shaped, with ventral medial process only; (4) C-shaped, with two medial processes. 56. Right paramere - apophysis (C-shaped with dorsal medial process only): (0) without teeth or contours; (1) with teeth or contours. 57. Right paramere - apophysis (C-shaped with ventral medial process only): (0) without teeth; (1) with teeth. 58. Right paramere - apophysis (C-shaped without medial process only): (0) without teeth; (1) with teeth. 59. Right paramere - medial process (C-shaped with dorsal medial process only): (0) without teeth; (1) with teeth. 60. Right paramere - medial process (C-shaped with two medial process (ventral) only): (0) without teeth; (1) with teeth. 61. Phallotheca - general shape: (0) simple, without process; (1) complex, with process. 62. Phallotheca - opening: (0) apical only, closed medially; (1) apical to medial - right-dorsolateral orientation; (2) apical to medial - dorsal orientation. 63. Phallotheca - apical lobe on right dorsal margin of opening: (0) absent; (1) present. 64. Phallotheca - subapical fold/lobe/protuberance on ventral surface: (0) absent; (1) present. 65. Phallotheca - size of subapical fold/lobe/protuberance on ventral surface: (0) small (barely raised above surface of phallotheca); (1) large (prominently raised above surface of phallotheca). 66. Phallotheca - margin of subapical fold/lobe/protuberance on ventral surface: (0) rounded; (1) bilobed; (2) subtriangular; (3) subrectangular, with narrowly concave margin; (4) spine-like or finger-like.

Table 2.1. Cont/-.

67. Phallotheca - subapical lobe on dorsal surface (removed from opening): (0) absent; (1) present. 68. Phallotheca - number of subapical lobe on dorsal surface: (0) one; (1) two or more. 69. Phallotheca - size of single subapical lobe on dorsal surface: (0) small (barely raised above surface of phallotheca); (1) large (prominently raised above surface of phallotheca). 70. Phallotheca - margin of single subapical lobe on dorsal surface: (0) rounded; (1) bilobed; (2) finger-like. 71. Phallotheca - basal lobe on dorsal surface: (0) absent; (1) present. 72. Aedeagus - number of endosomal spicules: (0) one; (1) two; (2) three; (3) four or more. 73. Aedeagus - margin of single spicule: (0) simple, without basal fold; (1) with basal fold. 74. Aedeagus - length of single spicule: (0) not elongate (about as long as wide); (1) elongate (about five times longer than wide at base). 75. Aedeagus - subapical shape of single spicule: (0) somewhat straight; (1) recurved. 76. Aedeagus - position of single spicule: (0) close to secondary gonopore; (1) distal to secondary gonopore. 77. Aedeagus - multiple spicule form - with at least one simple spicule (unbranched, tapering to apex, margins smooth): (0) absent; (1) present. 78. Aedeagus - multiple spicules - branched (at least one), complex, with serrate margins: (0) absent; (1) present.

Table 2.2. Pseudoloxops character matrix. (-) innaplicable, (?) unknown

Characters 1 1 1 1 1 1 1 Taxa 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 outgroups Orthotylus graciliphilus 1 0 1 1 0 0 0 1 0 0 1 0 0 0 0 0 Orthotylus intratropica 1 0 1 1 0 0 0 1 0 1 1 0 0 0 0 0 Orthotylini undescribed sp. 1 0 1 1 0 0 0 1 1 1 0 0 0 0 0 2 Pseudoloxops adamsoni 0 0 0 0 1 1 1 1 1 0 2 0 1 0 2 2 amabilis 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 2 amfitrite 0 1 0 0 1 1 1 1 1 1 2 0 1 1 1 2 baiawaensis 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 bifasciatus 0 1 0 0 1 1 1 1 0 1 2 0 1 0 1 2 coccineus 0 1 0 0 1 1 2 1 0 1 0 0 1 1 1 1 cordiformis 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 2 efatensis 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 kimiensis 0 0 0 0 0 1 2 1 1 1 0 0 1 1 1 2 kraussi 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 lenakelensis 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 mcilwraithensis 0 0 0 1 1 0 1 1 1 1 2 0 1 1 1 2 miyamotoi 0 0 0 1 1 0 0 ? ? 1 0 1 1 0 1 2 naraiyawa 0 1 0 0 1 1 1 0 0 1 2 0 1 0 0 2 nike 0 1 0 0 1 1 1 ? ? 1 2 1 1 1 1 1 niobe 0 1 0 0 1 1 1 1 1 1 0 1 1 1 2 2 palauensis 0 0 0 0 1 1 2 1 1 1 2 0 1 1 2 2 pancheriaphila 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 2 rubroclavus 0 0 0 0 1 1 1 1 1 1 2 0 1 0 2 2 sikaiana 0 1 0 0 1 1 1 1 1 1 2 0 1 1 2 1 signatus 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 2 solomonensis 0 1 0 0 1 1 2 1 1 1 2 0 0 0 1 1 straatmani 0 0 0 0 1 1 2 1 1 1 2 0 1 1 1 1 sudanicus 0 1 0 0 1 1 2 1 1 1 2 1 1 1 1 1 tabwemasana 0 1 0 0 1 1 2 1 0 1 2 0 0 0 0 0 tatarnici 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 vitiensis 0 1 0 0 1 1 1 0 0 1 2 0 1 0 1 1 woodwardi 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 2 species #07 0 0 0 0 1 1 1 1 0 1 2 0 1 0 2 2 species #09 0 0 0 1 1 1 1 0 0 1 2 0 0 0 0 0 species #102 0 0 0 0 1 1 1 1 0 1 2 0 1 1 1 1 species #104 0 1 0 0 1 1 1 1 0 1 0 1 1 1 1 1 species #106 0 1 0 0 1 1 1 1 1 1 2 0 1 0 2 1 species #11 0 0 0 0 1 1 1 1 0 1 2 0 1 1 1 1 species #14 0 0 0 0 1 1 1 0 0 1 2 0 0 0 0 0 species #53 0 1 0 0 0 1 0 1 1 1 2 0 1 1 2 2 species #62 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 species #66 0 1 0 0 1 1 1 1 1 1 2 0 1 0 1 1 species #68 0 1 0 0 1 1 2 1 1 1 2 0 1 1 1 2 species #76 0 1 0 0 1 1 1 1 1 1 2 0 1 1 1 1 species #80 0 1 0 0 1 1 2 1 1 1 2 0 1 1 1 2 species #80A 0 1 0 0 1 1 2 1 1 1 2 0 1 1 1 2 species #81 0 1 0 0 1 1 2 1 0 1 2 0 1 1 1 2 species #83 0 0 0 0 1 1 1 1 1 1 2 0 1 1 1 1 species #85 0 0 0 0 1 1 1 1 0 1 2 0 1 1 1 1 species #90 0 0 0 0 1 1 1 1 0 1 2 0 1 1 1 1 species #94 0 0 0 0 1 0 0 0 0 1 2 0 0 0 0 0 species #97 0 0 0 0 1 1 0 1 0 1 2 0 0 0 0 0 species #98 0 0 0 0 1 1 1 1 1 1 2 0 0 0 0 0 species #99 0 0 0 0 1 1 2 0 0 1 2 0 1 0 1 1

41 Table 2.2. Cont/-

Characters 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 Taxa 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 outgroups Orthotylus graciliphilus 0 0 3 0 0 2 2 - - - 1 2 - - 2 - Orthotylus intratropica 0 0 3 0 0 2 1 - - - 1 2 - - 2 - Orthotylini undescribed sp. 2 1 1 1 1 1 1 - - - 1 0 1 0 1 - Pseudoloxops adamsoni 2 1 1 1 1 0 3 1 0 0 1 0 1 2 0 1 amabilis 2 1 1 1 1 0 3 1 0 0 1 0 1 0 1 - amfitrite 2 0 1 1 1 0 3 1 0 0 0 0 1 2 1 - baiawaensis 2 1 1 1 1 0 3 1 0 0 0 0 1 0 0 1 bifasciatus 2 1 1 1 1 0 3 1 0 0 1 0 1 2 0 1 coccineus 2 0 1 1 1 1 1 - - - 0 0 1 3 1 - cordiformis 2 1 1 1 1 0 3 1 0 0 0 0 1 2 1 - efatensis 2 1 1 1 1 0 0 - - - 1 0 1 1 0 1 kimiensis 2 0 1 1 1 0 3 0 0 1 1 0 1 2 1 - kraussi 2 1 1 1 1 0 0 - - - 1 0 1 2 1 - lenakelensis 2 1 1 1 1 0 0 - - - 1 0 1 1 0 1 mcilwraithensis 2 0 1 1 1 0 1 - - - 0 0 1 2 1 - miyamotoi 2 0 1 1 1 1 1 - - - 0 0 1 2 0 1 naraiyawa 1 0 2 1 1 0 0 - - - 1 0 1 2 1 - nike 2 1 1 1 1 0 3 1 1 1 0 0 1 2 1 - niobe 2 0 1 1 1 1 3 1 1 1 0 0 1 3 1 - palauensis 2 2 1 1 1 0 3 1 1 1 1 0 1 2 0 1 pancheriaphila 2 1 1 1 1 0 3 1 0 1 0 0 1 2 1 - rubroclavus 2 0 1 1 1 0 3 1 0 0 1 1 - - 0 1 sikaiana 2 0 1 1 1 0 3 1 0 1 0 0 1 2 1 - signatus 1 0 1 1 1 0 3 1 0 0 1 0 1 2 1 - solomonensis 2 1 1 1 1 0 3 1 0 1 0 1 - - 0 1 straatmani 2 0 1 1 1 0 3 0 0 1 0 0 1 2 0 1 sudanicus 2 1 1 1 1 0 3 1 1 1 0 0 1 2 0 1 tabwemasana 2 1 0 0 0 0 0 - - - 1 0 1 1 0 1 tatarnici 2 1 1 1 1 0 0 - - - 0 2 - - 2 - vitiensis 2 0 1 1 1 0 0 - - - 0 0 1 2 1 - woodwardi 2 1 1 1 1 0 3 1 0 0 1 0 1 2 0 1 species #07 2 1 1 1 1 0 3 1 0 0 1 0 1 2 0 1 species #09 2 0 1 1 1 0 0 - - - 1 0 0 - 0 0 species #102 2 0 1 1 1 0 0 - - - 1 0 0 - 0 0 species #104 2 0 1 1 1 1 3 1 1 1 0 0 1 3 1 - species #106 2 1 1 1 1 0 3 1 0 0 0 0 1 2 0 1 species #11 2 0 1 1 1 0 0 - - - 1 0 0 - 0 0 species #14 1 1 0 0 0 0 0 - - - 1 0 0 - 0 0 species #53 2 0 1 1 1 1 1 - - - 1 0 1 0 0 1 species #62 2 1 1 1 1 0 0 - - - 1 0 1 2 0 1 species #66 2 1 1 1 1 0 3 1 0 0 0 0 1 2 1 - species #68 2 1 1 1 1 0 3 1 0 1 0 0 1 2 1 - species #76 2 1 1 1 1 0 3 1 0 0 0 0 1 2 1 - species #80 2 1 1 1 1 0 3 1 0 0 1 0 1 2 1 - species #80A 2 1 1 1 1 0 3 1 0 1 1 0 1 2 1 - species #81 2 0 1 1 1 1 3 1 0 1 1 0 1 2 1 - species #83 1 1 1 1 1 0 3 1 0 1 1 0 1 0 0 1 species #85 2 1 1 1 1 0 3 1 0 1 0 0 1 2 0 1 species #90 2 0 1 1 1 0 0 - - - 1 0 0 - 0 0 species #94 0 0 0 0 0 0 0 - - - 1 0 0 - 0 0 species #97 0 0 0 0 0 0 0 - - - 1 0 0 - 0 0 species #98 2 0 0 0 0 0 0 - - - 1 0 0 - 0 0 species #99 2 1 1 1 1 0 3 1 0 0 0 0 1 2 1 -

42 Table 2.2. Cont/-

Characters 3 3 3 3 3 3 3 4 4 4 4 4 4 4 4 4 Taxa 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 outgroups Orthotylus graciliphilus - 0 0 - - 0 2 0 0 - 1 0 1 0 0 0 Orthotylus intratropica - 0 2 0 0 0 2 1 0 - 1 0 1 1 0 0 Orthotylini undescribed sp. - 0 2 0 0 0 1 1 0 - 1 0 1 0 1 0 Pseudoloxops adamsoni 3 0 2 1 0 0 0 1 0 - ? ? ? ? ? ? amabilis - 1 1 - - 1 0 1 0 - ? ? ? ? ? ? amfitrite - 0 1 - - 0 0 1 1 1 ? ? ? ? ? ? baiawaensis 0 1 2 1 1 1 0 1 0 - 0 0 0 0 1 1 bifasciatus 2 0 2 1 0 0 0 1 1 0 1 1 0 0 1 0 coccineus - 1 1 - - 1 1 1 1 1 1 0 1 0 1 0 cordiformis - 0 2 0 0 0 0 1 ? ? 1 0 0 0 1 ? efatensis 5 0 2 1 0 0 0 1 0 - 1 0 0 0 1 0 kimiensis - 1 2 1 0 1 0 1 1 1 1 0 0 0 0 0 kraussi - 1 2 1 0 1 0 1 0 - 1 1 0 0 1 0 lenakelensis 5 0 2 1 0 0 0 1 0 - 1 0 0 0 1 0 mcilwraithensis - 1 2 1 0 1 0 1 1 0 0 0 0 0 1 0 miyamotoi 5 0 1 - - 0 1 1 1 0 ? ? ? ? 1 ? naraiyawa - 0 2 1 0 1 0 1 1 1 1 1 0 0 1 0 nike - 1 1 - - 1 0 1 0 - ? ? ? ? ? ? niobe - 1 1 - - 1 1 1 1 1 ? ? ? ? ? ? palauensis 3 1 2 1 0 1 0 1 1 1 ? ? ? ? ? ? pancheriaphila - 0 1 - - 0 0 1 0 - 1 1 0 0 1 0 rubroclavus 1 0 2 1 0 0 0 1 0 - 0 1 0 0 1 0 sikaiana - 0 2 1 0 0 0 1 0 - 1 0 0 0 1 1 signatus - 1 2 1 0 1 0 1 0 - 1 1 0 0 1 0 solomonensis 4 0 2 1 0 0 0 1 0 - 1 0 0 0 1 1 straatmani 3 0 1 - - 0 0 1 1 1 0 0 0 0 1 1 sudanicus 2 1 2 1 0 1 1 1 1 0 ? ? ? ? ? ? tabwemasana 5 0 0 - - 0 0 1 0 - 1 1 0 0 1 0 tatarnici - 1 2 1 0 0 0 1 0 - 1 0 0 0 0 0 vitiensis - 0 1 - - 0 0 1 0 - 1 0 0 0 1 1 woodwardi 4 1 2 1 0 0 0 1 0 - 1 1 0 0 1 0 species #07 1 0 2 0 0 0 0 1 0 - 0 1 0 0 1 0 species #09 - 0 0 - - 0 0 0 1 0 0 1 0 0 1 0 species #102 - 0 2 0 0 0 0 1 1 0 0 1 0 0 1 0 species #104 - 0 1 - - 1 1 1 1 1 1 0 1 0 1 0 species #106 2 1 1 - - 1 0 1 0 - 0 1 0 0 1 0 species #11 - 0 2 0 0 0 0 1 1 0 0 1 0 0 1 0 species #14 - 0 0 - - 0 0 0 0 - 0 1 0 0 1 0 species #53 0 1 2 1 0 1 1 1 1 1 1 0 1 0 2 0 species #62 0 1 1 - - 1 0 1 0 - 1 0 0 0 1 0 species #66 - 1 1 - - 1 0 1 0 - ? ? ? ? 0 ? species #68 - 0 1 - - 1 0 1 0 - 0 0 0 0 1 0 species #76 - 1 1 - - 1 0 1 1 1 1 0 0 0 1 0 species #80 - 1 2 1 0 0 1 1 0 - 0 1 0 0 1 0 species #80A - 1 2 1 0 0 1 1 0 - 0 1 0 0 1 0 species #81 - 1 2 1 0 1 1 1 1 1 1 1 0 0 1 0 species #83 3 1 2 1 1 1 0 1 0 - 0 1 0 0 1 0 species #85 1 0 2 1 0 0 0 1 1 0 0 1 0 0 1 0 species #90 - 0 2 0 0 0 0 1 0 - 0 1 0 0 1 0 species #94 - 0 0 - - 0 0 0 0 - 0 1 0 0 1 0 species #97 - 0 2 0 0 0 0 0 0 - 0 1 0 0 1 0 species #98 - 0 0 - - 0 0 0 0 - 0 1 0 0 1 0 species #99 - 0 2 1 0 0 0 1 1 1 1 1 0 0 1 0

43 Table 2.2. Cont/-

Characters 4 5 5 5 5 5 5 5 5 5 5 6 6 6 6 6 Taxa 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 outgroups Orthotylus graciliphilus 0 - 1 0 - 0 1 - - 1 - - 1 2 0 1 Orthotylus intratropica 0 - 1 1 - 0 0 - - - - - 0 2 0 0 Orthotylini undescribed sp. 0 - 1 1 - 0 2 0 - - 0 - ? ? ? ? Pseudoloxops adamsoni 1 1 ? 0 0 1 2 ? - - ? - ? ? ? ? amabilis 1 0 ? 1 0 0 1 - - 0 - - ? ? ? ? amfitrite 1 1 ? 1 0 0 1 - - 0 - - ? ? ? ? baiawaensis 1 1 1 0 0 1 1 - - 1 - - 0 1 0 0 bifasciatus 1 1 1 0 0 1 3 - 1 - - - 1 1 0 1 coccineus 0 - 1 1 - 0 2 0 - - 1 ? 1 1 0 1 cordiformis 1 1 ? 0 0 1 2 ? - - 1 - ? ? ? ? efatensis 1 1 1 0 0 1 2 1 - - 1 - 1 1 0 1 kimiensis 0 - 1 1 - 1 3 - 1 - - - 1 ? ? ? kraussi 1 1 1 0 0 1 2 1 - - 0 - 1 1 0 1 lenakelensis 1 1 1 0 0 1 2 1 - - 1 - 1 1 0 1 mcilwraithensis 0 - 1 1 - 0 2 0 - - 0 - 0 1 0 0 miyamotoi 0 - ? 0 - 0 0 - - - - - ? ? ? ? naraiyawa 1 1 1 0 0 1 2 1 - - 1 - 1 1 1 1 nike 1 1 ? 1 0 0 1 - - 0 - - ? ? ? ? niobe 1 0 ? 1 1 0 1 - - 0 - - ? ? ? ? palauensis 1 1 ? 0 0 0 1 - - 1 - - ? ? ? ? pancheriaphila 1 1 1 0 0 1 2 1 - - 1 - 1 1 0 1 rubroclavus 1 1 ? 0 0 ? 2 1 - - 1 - ? ? ? ? sikaiana 1 1 1 0 0 1 2 1 - - 1 - 0 1 0 0 signatus 1 1 1 0 0 1 3 - 1 - - - 1 1 0 1 solomonensis 1 1 1 0 0 0 2 1 - - 1 - 0 1 0 0 straatmani 1 0 1 1 1 0 1 - - 1 - - 0 1 0 0 sudanicus 1 1 ? 1 0 0 2 1 - - 1 - ? ? ? ? tabwemasana 1 1 1 0 0 1 3 - 1 - - - 1 1 0 1 tatarnici 1 1 1 0 0 0 2 1 - - 1 - 0 1 0 0 vitiensis 1 1 1 0 0 1 2 1 - - 1 - 1 1 0 1 woodwardi 1 1 1 0 0 1 2 1 - - 1 - 1 1 0 1 species #07 1 1 1 0 0 1 2 1 - - 1 - 1 1 0 1 species #09 1 1 1 0 0 0 2 1 - - 1 - 1 1 0 1 species #102 1 0 0 0 0 0 2 1 - - 1 - 1 1 1 1 species #104 0 - 1 1 - 0 2 0 - - 1 - 1 2 0 0 species #106 1 1 1 0 0 0 0 - - - - - 0 1 0 0 species #11 1 0 0 0 0 0 2 1 - - 1 - 1 1 1 1 species #14 1 1 1 0 0 0 ? ? ? ? ? ? 1 1 ? 1 species #53 0 - ? 1 - 0 1 - - 0 - - 0 1 0 0 species #62 1 0 1 1 0 0 0 - - - - - ? ? ? ? species #66 1 1 ? 1 1 0 1 - ? ? ? ? ? ? ? ? species #68 1 1 1 0 0 1 1 - - 1 - - 0 0 0 0 species #76 1 0 1 1 0 0 1 - - 0 - - 0 1 0 0 species #80 1 0 1 1 0 1 4 - - - - 1 1 1 0 ? species #80A 1 0 1 1 0 1 4 - - - - 0 1 1 0 1 species #81 1 0 1 1 0 1 4 - - - - 0 1 1 1 1 species #83 1 1 0 0 0 1 2 1 - - 1 - 1 1 0 1 species #85 1 1 0 0 0 0 2 0 - - 1 - 1 1 0 1 species #90 1 0 0 0 0 0 2 1 - - 1 - 1 1 1 1 species #94 1 1 1 0 0 1 2 1 - - 1 - 1 1 1 1 species #97 1 1 1 0 0 0 3 - 1 - - - 1 1 0 1 species #98 1 1 1 0 0 0 3 - 0 - - - 1 0 0 1 species #99 1 1 0 0 0 1 2 1 - - 1 - 1 1 1 1

44 Table 2.2. Cont/-

Characters 6 6 6 6 6 7 7 7 7 7 7 7 7 7 Taxa 5 6 7 8 9 0 1 2 3 4 5 6 7 8 outgroups Orthotylus graciliphilus 0 0 0 - - - 0 2 - - - - 0 1 Orthotylus intratropica - - 0 - - - 0 2 - - - - 0 1 Orthotylini undescribed sp. ? ? ? ? ? ? ? 1 - - - - 1 0 Pseudoloxops adamsoni ? ? ? ? ? ? ? ? ? ? ? ? ? ? amabilis ? ? ? ? ? ? ? 2 - - - - ? 0 amfitrite ? ? ? ? ? ? ? 2 - - - - ? 0 baiawaensis - - 0 - - - 0 1 - - - - 1 0 bifasciatus 0 0 1 0 0 0 0 0 0 0 0 1 - - coccineus 0 4 1 1 - - 1 2 - - - - ? 0 cordiformis ? ? ? ? ? ? ? ? ? ? ? ? ? ? efatensis 1 0 0 - - - 1 ? ? ? ? ? ? ? kimiensis ? ? ? ? ? ? ? ? ? ? ? ? ? ? kraussi 0 3 1 0 0 0 1 0 0 1 1 0 - - lenakelensis 1 3 1 0 0 0 0 0 0 1 0 1 - - mcilwraithensis - - 0 - - - 0 2 - - - - 0 1 miyamotoi ? ? ? ? ? ? ? 0 0 ? ? ? ? - naraiyawa 1 1 1 0 1 0 0 0 0 1 0 0 - - nike ? ? ? ? ? ? ? 1 - ? - ? ? 0 niobe ? ? ? ? ? ? ? 1 - ? - ? ? 0 palauensis ? ? ? ? ? ? ? 0 0 1 0 - ? - pancheriaphila 0 2 1 0 0 0 0 0 0 1 1 0 - - rubroclavus ? ? ? ? ? ? ? 0 ? ? ? ? ? ? sikaiana - - 0 - - - 0 1 - - - - 1 0 signatus 0 0 1 0 0 0 0 1 - - - - 1 0 solomonensis - - 0 - - - 0 1 - - - - 1 0 straatmani - - 0 - - - 0 3 - - - - 1 0 sudanicus ? ? ? ? ? ? ? 1 - - - - ? 1 tabwemasana 1 0 1 1 - - 0 0 0 1 1 0 - - tatarnici - - 0 - - - 0 3 - - - - 1 0 vitiensis 1 3 1 0 1 0 0 0 0 1 0 0 - - woodwardi 0 1 0 - - - 0 0 0 1 1 0 - - species #07 1 2 1 0 1 2 0 0 0 0 1 1 - - species #09 1 2 1 0 0 2 0 0 0 1 1 1 - - species #102 1 1 1 0 0 1 0 0 0 1 1 0 - - species #104 - - 0 - - - 0 1 - - - - 1 1 species #106 - - 0 - - - 0 2 - - - - 1 0 species #11 1 1 1 0 0 1 0 0 0 1 0 0 - - species #14 1 1 1 0 0 0 0 0 0 1 1 0 - - species #53 - - 0 - - - 0 3 - - - - 1 0 species #62 ? ? ? ? ? ? ? ? ? ? ? ? ? ? species #66 ? ? ? ? ? ? ? ? ? ? ? ? ? ? species #68 - - 0 - - - 0 2 - - - - 1 1 species #76 - - 0 - - - 0 1 - - - - 1 0 species #80 ? ? ? ? ? ? ? 0 1 1 1 1 - - species #80A 1 4 1 0 0 2 1 0 0 1 0 1 - - species #81 0 2 1 1 - - 1 0 1 0 1 1 - - species #83 1 1 1 0 0 1 0 0 0 1 1 0 - - species #85 1 1 1 0 0 0 0 0 0 0 1 0 - - species #90 1 1 1 0 0 0 0 0 0 1 1 0 - - species #94 0 0 0 - - - 0 0 0 1 1 1 - - species #97 0 0 1 0 0 0 0 2 - - - - 1 0 species #98 0 0 1 0 0 0 0 1 - - - - 1 0 species #99 0 3 1 0 1 2 0 0 0 1 0 0 - -

Figure 2.1. Colour characters. a. P. baiawaensis. b. P. species #106. c. P. cordiformis. d. P. species #104. e. P. kraussi. f. P. solomonensis. g. P. lenakelensis. h. P. efatensis. i. P. species #83. Scale = 1 mm. 46

Figure 2.2. Pygophore characters, dorsal view: dorsal margin (DM), moderately to strongly concave (a., b. & e.) or weakly concave to straight (c., d. & f.) ; with left tergal process (ltp) (a.); with medial lobe (ml) (c. & d.); ventral margin (VM) projecting (a., c. - f.) or not projecting (b.); with ventral process (vp) (e. & f.); a. P. species #104, b. P. kimiensis, c. P. species #83, d. P. species #106, e. P. solomonensis, f. P. baiawaensis. Scale = 0.1 mm. 47

Figure 2.3. Left paramere characters, dorsal view: Y-shaped (a. - e., g.); sickle-shaped (f. & h.); apophysis (ap) of Y-shaped form parallel or ca. 45° to sensory lobe (sl) (a. - e.), or right angle to sensory lobe (g.); apophysis slim (a. - e.) or moderately broad to broad (f., g. & h.); shape of distal margin of apophysis bilobed (d.) or rounded (a. - c., e. - h.); with basal process (bp) (a., b., d. & e.); a. P. baiawaensis, b. P. tabwemasana, c. P. species #106, d. P. species #83, e. P. pancheriaphila , f. P. mcilwraithensis, g. P. straatmani, h. P. species #104. Scale = 0.1 mm. 48

Figure 2.4. Right paramere characters, dorsal view: simple C-shaped, without medial process (f.); club-shaped, without medial process (g.); with dorsal medial process only (dmp) (a. - e.); with ventral medial process only (vmp) (h.); apophysis (ap) with teeth (b., c., e., g., h.); medial process with teeth (a. - c., e., h.); a. P. species #104, b. P. species #83, c. P. naraiyawa, d. P. mcilwraithensis, e. P. pancheriaphila, f. P. species #106, g. P. baiawaensis, h. P. tabwemasana. Scale = 0.1 mm. 49

Figure 2.5. Phallotheca characters: a., b., f. & g., right lateral view; c. & d., dorsal view; e., left ventral view; opening, right-dorsolateral, from apical to medial (a., b., d. - g.) or dorsal, from apical to medial (c.); simple, without process (f. & g.); with ventral fold (vf), enlarged (a.) or reduced (b. & e.); with subapical lobe on dorsal surface (dsl) (a. - e.); with basal lobe (bl) on dorsal surface (c. & d.); a. P. species #83, b. P. pancheriaphila, c. P. species #104, d. and e. P. kraussi, f. P. species #106, g. P. tatarnici. Scale = 0.1 mm. 50

Figure 2.6. Aedeagus characters: a., c. & d., right lateral view; b., e. & f., ventral view; with one spicule (a.) ; with two spicules (b. & c.); with three spicules (d. & e.); with four spicules (f.); simple spicule form (unbranched, margins smooth) (a. - d, f.); complex spicule form (branched, margins serrate) (c. & e.); a. P. species #83, b. P. baiawaensis, c. P. species #104, d. P. species #106, e. P. mcilwraithensis, f. P. straatmani. Scale = 0.1 mm. 51

CHAPTER III

A SYSTEMATIC STUDY OF THE GENUS PSEUDOLOXOPS KIRKALDY IN

THE SOUTHWEST PACIFIC1

INTRODUCTION

The genus Pseudoloxops Kirkaldy belongs to the plant bug tribe Orthotylini

(Heteroptera: Miridae: Orthotylinae), and currently comprises 38 described species

(Schuh 1995, 2008). It is widely distributed throughout the Palaearctic, Oriental,

Afrotropical and Australian regions, and within island archipelagos in the Pacific Basin

(Schuh 1995, 2008).

The genus Pseudoloxops was first erected by Kirkaldy (1905), for the European species,

P. coccineus (Meyer-Dür, 1843). Despite its undoubted diversity the genus has received scant attention, and previous studies are regional in scope (Knight 1935, 1937;

Carvalho, 1980a; Linnavuori, 1994; Liu and Zheng, 1994; Schuh, 1974; Yasunaga

1997, 1999; Zou, 1987).

1The Australian species in this chapter have been submitted separately as a publication to the journal Zootaxa, with Hazali and Cassis as coauthors. The submitted paper was undertaken by Hazali, including all the observational, descriptive, analytical work, and the preparation of the manuscript. Cassis assisted in project design and species conception, as well as editing of the manuscript. This manuscript remains in review.

Yasunaga (1997) defined Pseudoloxops on the basis of the following characters: 1) small and slender body; 2) densely pubescent body; 3) dorsum pale yellow, and partly sanguineous or scarlet colouration, often with dark spots; and anteriorly projected frons.

This combination of characters is not exclusive to Pseudoloxops, and previous to my study there has been no known nonhomoplasious synapomorphy or unique diagnostic character for the genus. In Chapter 2, I defined Pseudoloxops on the basis of the following characters: the head flat or nearly flat, the frons convex in lateral view and the maxillary plate greatly tumescent medially.

In collections, specimens are often identified as Pseudoloxops because of the red colouration on the lateral margins of the pronotum, often contiguous with red patterning on the hemelytra. However, the diagnostic value of this colouration has not been previously tested in this group of mirids and similar colour patterning is known in unrelated orthotylines, such as the New Zealand species of Zanchius Distant (Eyles,

2005).

An examination of these colour characters and male genitalia of all the species provisionally assigned to the genus Pseudoloxops in collections has been conducted.

This was undertaken to provide a revised definition of the genus across its distributional range. The second aim of this study was to describe the species of Pseudoloxops from the Southwest Pacific, including the Australian species, as well as documenting their host plant associations and distribution.

Based on previous studies, only one species is described from the Southwest Pacific: P. vitiensis Kirkaldy 1908 from Fiji. Carvalho and Wallerstein (1976) redescribed this

species and provided illustrations of male genitalic structures. In my study, a total of 19 new species were recognized and I have described all these species, provided keys, and illustrations of male genitalic characters for most of the species.

MATERIALS AND METHODS

Materials

This study is based on an examination of specimens from museum collections and new field collections. Collecting data such as collecting date, locality, sex, collector’s name, and host plant were recorded in the online Plant Bug Planetary Biodiversity Inventory

Project (PBI) locality database (https://research.amnh.org/pbi/locality/). All specimens examined were databased with a unique specimen identifier (USI), including a matrix cod. These USI numbers comprise eight digits (e.g., USI 00192915).

Acronyms for the institutions are as follows:

AMNH American Museum of Natural History, New York

ANIC Australian National Insect Collection, CSIRO, Canberra

BMNH Natural History Museum, London

BPBM Bernice P. Bishop Museum, Honolulu

MEM Mississippi Entomological Museum, Mississippi State University,

Mississippi

QDPI DPI Queensland, Queensland

SAMA South Australian Museum, Adelaide

USNM United States National Museum of Natural History, Smithsonian

Institution, Washington

UNSW University of New South Wales, Sydney

UQIC Insect Collection, University of Queensland, Brisbane 56

Taxa

For the generic description and diagnosis, I have examined described and undescribed species. For those described species, which I have not examined, I have used information given in the literature, including the original descriptions.

Available specimens of the following described species were examined: P. adamsoni

Knight 1937, P. amabilis Linnavuori 1986, P. amfitrite Linnavuori 1994, P. bakeri

Knight 1937, P. bifasciatus Usinger 1946, P. coccineus (Meyer-Dür 1843), P. iranicus

Kerzhner 1962, P. miyamotoi Yasunaga 1997, P. nigrobasicornis Knight 1937, P. nike

Linnavuori 1994, P. ninos Linnavuori 1994, P. niobe Linnavuori 1994, P. palauensis

Carvalho 1956, P. rubroclavus Knight 1937, P. rubroscutellatus Knight 1935, P. signatus Usinger 1946, P. sudanicus Linnavuori 1975, P. thetis Linnavuori 1994, P. transvaalensis Schuh 1974 and P. vitiensis Kirkaldy 1908.

For the treatment of species from the Southwest Pacific, only Pseudoloxops vitiensis

Kirkaldy 1908 was previously described from this region, and specimens of it were examined for this work. All the other species treated in this Chapter are new to science.

Methods

My taxonomic study was based on comparative morphological methods to discriminate among species, using principles outlined by Schuh and Brower (2009). Species discrimination was based on searching for discontinuities in external and genitalic characters between species, and morphometric data. Specimens were examined using a

LEICA MZ125 stereomicroscope and LEICA DM5000B compound microscope.

The dissections of the male genitalia were made following the methods described in

Kelton (1959). Terminology presented by Cassis (2008) was followed for male genitalia, where possible. The aedeagus is composed of an articulating base (the phallobase), a distal membranous medial tubule (the endosoma) housing endosomal spicule(s), bounding the ductus seminis, all of which are covered by the phallotheca.

Subsequently, the illustrations of the male genitalic structures were made using a camera lucida attached to a LEICA DM500B compound microscope. Characters of the pygophore, parameres, phallotheca and aedeagus of the males were illustrated.

Habitus photographs were taken of exemplar species of Pseudoloxops using the

Visionary Digital Micro-Imaging photographic system

Measurements of the external characters were taken using a digital micrometer attached to a LEICA MZ15 stereomicroscope. All measurements are in millimeters.

Measurements are shown in Table 3.1 and include body length, clypeus to cuneus length, head length and width, pronotum length and width, scutellum length and width, interocular distance, and lengths of first to fourth antennal).

Georeferenced data for each locality were obtained from label data, gazetteers, atlases and other sources. Distribution maps for each species were generated with ArcMap

9.3.1 (ESRI, 2009).

Morphology

For male genitalia, I have used the terminology for endosomal spicules of Cassis

(2008), and Cassis et al. (2010), including the following abbreviations: DES = dorsal endosomal spicule of aedeagus (DES1 and DES 2 = first and second dorsal endosomal spicules where three endosomal spicules are present); VES = ventral endosomal spicule of aedeagus. This terminology was not applied to those species where only a single spicule was present; this was because there was no positional criterion that could be applied.

Additional non-genital abbreviations are used as follows: AI-AIV = first to fourth antennal segments; LI-LIV – first to fourth labial segments; IOD = interocular distance; and, MFL = median flexion line. Further abbreviations are used in the illustrations and are explained in the figure captions.

TAXONOMY

Pseudoloxops Kirkaldy, 1905

Loxops Fieber 1858: 314 (gen. nov.; type species: Capsus coccineus Meyer-Dür 1843

by monotypy; junior homonym of Loxops Cabanis 1847 [Aves]); Carvalho

1958: 127 (catalogue).

Pseudoloxops Kirkaldy 1905: 268 (nomen nudum for Loxops Fieber); Kirkaldy 1906:

127 (catalogue); Oshanin 1910: 849 (catalogue); Reuter 1910: 150 (catalogue);

Stichel 1933: 220 (key); Hedicke 1935: 55 (key); Hsiao 1942: 254 (key); China

1943: 26 (catalogue); Kiritshenko 1951: 125 (key); Wagner 1952: 110, 115

(key, description); Carvalho and Leston 1952: 246 (key); Carvalho 1952: 78

(catalogue); Carvalho 1955a: 76 (key); Carvalho 1956: 61 (key to Micronesian

spp.); Carvalho 1958: 127 (catalogue); Wagner and Weber 1964: 307

(synonymy, description); Wagner 1974: 167 (description); Schuh 1974: 45

(diagnosis); Linnavuori 1994: 23 (syn., description, disc., key to African spp.);

Schuh 1995: 184 (catalogue); Yasunaga 1997: 11 (description, discussion);

Yasunaga 1999: 174 (description, discussion); Kerzhner and Josifov 1999: 273

(catalogue).

Aretas Distant 1909: 450 (gen. nov.; type species: Aretas imperatorius Distant 1909, by

monotypy); Carvalho 1952: 78 (synonymy).

Zonodorellus Poppius 1915: 68 (gen. nov.; type species Zonodorellus lateralis Poppius

1915, by original designation); Carvalho 1980b: 658 (synonymy).

Diagnosis. Pseudoloxops is recognized by the following combination of characters: body elongate-ovate to subparallel; ground colouration pale yellow to yellowish brown, with either red, reddish-brown or orange-red markings; body often densely setose with pale, moderately long to elongate, simple setae; AI usually with pale to dark, long setae; head subquadrangular, dorsum nearly flat to flat; frons anteriorly projected in dorsal view, convex in lateral view; vertex almost flat, midline sulcate, with impressed line posteriorly; maxillary plate greatly tumescent medially; bucculae usually reaching posterior margin of head; eyes contiguous with anterior margin of pronotum; first labial segment exceeding bucculae; AI subbasally enlarged; AII usually about 3x longer than

AI; right paramere usually C-shaped, always with medial processes, usually armed, sometimes barely with medial processes; a Y-shaped left paramere, sometimes sickle- shaped; and, endosoma ranging from single, smooth spicules to multiple, branched, smooth to serrated spicules.

Redescription. Body elongate-ovate to elongate-subparallel. COLOURATION. Body pale yellow to yellowish brown, often with red to reddish-brown markings, sometimes with orange-red markings. Head: mostly pale yellow to yellowish brown, usually with red to reddish-brown markings on frons, vertex, clypeus, mandibular plate and maxillary plate, rarely without red or reddish-brown markings. Thorax: pronotum pale yellow to yellowish brown, often with red to reddish-brown markings on anterior and lateral areas, sometimes entirely yellowish-brown or almost red; proepimeron and proepisternum pale yellow to yellowish brown, usually with red markings; mesoscutum and scutellum pale yellow brown to yellowish brown, often with red markings, sometimes almost orange-red to red; hemelytra pale yellow to yellowish brown, usually with red markings, sometimes with almost orange-red to red; legs mostly pale yellow brown to yellowish-brown, sometimes with red markings on hind femora, membrane veins often with red markings. Abdomen: mostly pale yellow brown to yellowish- brown, sometimes with red or orange-red markings. VESTITURE. Most species with dorsum setose with pale, moderately long to elongate, decumbent to semierect, simple setae, mostly on head, pronotum and hemelytra. Antennae: AI often densely setose with pale, moderately long, decumbent setae, intermixed sparsely with elongate, semierect, bristlelike setae, sometimes with dark setae; AII, AIII and AIV usually densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short, semierect, pale setae. Pleura and sterna: almost bare with pale, long to elongate, simple setae. Abdomen: usually setose with pale, elongate, semierect, simple setae. Legs: femora often densely setose with moderately long to elongate, semierect, simple, pale setae; tibiae usually setose with pale, short, semierect to nearly adpressed setae, intermixed with pale, long, spinelike setae. STRUCTURE. Head: subquadrangular, frons weakly to strongly projected anteriorly in dorsal view, convex in lateral view,

dorsum flat; vertex almost flat to flat, midline moderately to deeply sulcate, posterior margin rounded to carinate; maxillary plate greatly tumescent medially; bucculae usually reaching posterior margin of head. Labium: usually reaching or surpassing apex of metacoxae, sometimes reaching mesocoxae, first labial segment longer than bucculae. Eyes: usually reaching dorsal margin of head in lateral view, sometimes barely reaching dorsal margin, in dorsal view adjacent to anterior margin of pronotum.

Antennae: AI slightly to moderately enlarged subbasally, greater in diameter then AII,

AIII and AIV. Pronotum: trapeziform; anterior part rounded to slightly raised, anterior margin weakly concave medially; posterior margin weakly bisinuate, excavate to rectilinear; lateral margin nearly straight to straight; callar region depressed medially, cali weakly to slightly raised, with weakly to impressed line; disc nearly flat to flat.

Mesoscutum: exposed. Scutellum: disc almost flat to slightly raised. Hemelytra: median flexion line (MFL) weakly convex; cuneus moderately long to elongate. Male genitalia: genital capsule with weakly to strongly concave dorsal margin, sometimes with medial lobe, ventral margin nearly straight to strongly convex, sometimes with ventral process; right paramere ranging from club-shaped to C-shaped, with dorsal or ventral medial process, usually armed; left paramere usually Y-shaped, sometimes sickle-shaped; phallotheca ranging from simple to complex or with processes on dorsal and ventral surfaces; endosoma ranging from single, not serrated spicules to multiple, branched, smooth to serrated spicules.

Host plants. Host plant associations have been recorded for 9 of the 38 described species of Pseudoloxops. They are found from the following genera and families:

Glochidion J.R.&G. Forster (Euphorbiaceae), Castanopsis (D.Don) Spach. (Fagaceae),

Cocculus DC. (Menispermaceae), Ficus L. (Moraceae), Fraxinus L. (Oleaceae),

Pipturus Wedd. (Urticaceae), Pithecellobium Mart. (Fabaceae), Prunus L. (Rosaceae),

Quercus L. (Fagaceae), Styrax L. (Styracaceae), Terminalia L. (Combretaceae) and

Ziziphus Mill. (Rhamnaceae).

Distribution. Palearctic, Oriental, Afrotropical and Australian zoogeographic regions, and island archipelagos of the Pacific Basin.

Remarks. Pseudoloxops is best recognised by the following head characters: subquadrangular head, frons and vertex almost to entirely flat, frons anteriorly projected, weakly to strongly convex, and the maxillary plate tumescent. I have examined the morphology of the type species, P. coccineus and it has the above head morphology, red colour patterning and three endosomal spicules. The generic synonymy of Pseudoloxops as proposed by Carvalho (1952, 1980b) is maintained, with

Aretas Distant and Zonodorellus Poppius as junior synonyms. The type species of the latter two available names were not examined, but on the basis of their original descriptions, there is no basis for removing them from synonymy.

The relationships of Pseudoloxops are poorly understood and it is beyond the scope of this work to ascertain its systematic position within the subfamily Orthotylinae. It clearly belongs to the nominate tribe in that the females possess inter-ramal lobes. The head structure is unique within the Orthotylini, and differs considerably to that found in all the other genus groups recognised by Schuh (1974: Falconia group, Zanchius group,

Sericophanes group and Orthotylus group). The male genitalia are highly variable within Pseudoloxops, and some possess strongly serrated endosomal spicules, as in austromirine Orthotylinae (Cassis, 2008) and other Orthotylini, whereas others have a simple spicule.

Checklist of Pseudoloxops of the Southwest Pacific (inclusive of Australia)

P. baiawaensis n.sp. Papua New Guinea; Malaysia (Sabah)

P. cheesmanae n.sp. Vanuatu (Aneityum Is.)

P. cordiformis n.sp. Australia (Queensland)

P. efatensis n.sp. Vanuatu (Efate Is., Ambrym Is.)

P. gagnei n.sp. Papua New Guinea

P. kimiensis n.sp. Papua New Guinea

P. kraussi n.sp. Vanuatu (Efate Is., Espiritu Santo Is., Maewo

Is., Shepherd Islands, Tanna Is.)

P. lenakelensis n.sp. Vanuatu (Tanna Is.)

P. mcilwraithensis n.sp. Australia (Queensland)

P. mourangeensis n.sp. Australia (Queensland)

P. naraiyawa n.sp. Fiji (Viti Levu)

P. pancheriaphila n.sp. New Caledonia

P. sikaiana n.sp. Solomon Islands (Bougainville, Florida Group,

Malaita, New Georgia, San Cristoval, Santa

Ysabel)

P. solomonensis n.sp. Solomon Islands (Guadalcanal, Malaita, San

Cristoval)

P. straatmani n.sp. Solomon Islands (Malaita, Santa Ysabel)

P. tabwemasana n.sp. Vanuatu (Espiritu Santo Is.)

P. tatarnici n.sp. Australia (Northern Territory)

P. vitiensis Kirkaldy 1908 Fiji (Matuku Is., Viti Levu, Vanua Levu)

P. wareo n.sp. Papua New Guinea

P. woodwardi n.sp. Australia (Queensland) 64

Key to species of Pseudoloxops of the Southwest Pacific (inclusive of Australia)

1. Mesoscutum uniformly pale yellow brown or yellowish brown ...... 2

- Mesoscutum not uniformly pale yellow brown or yellowish brown, with orange-red

or red markings ...... 10

2. Antennal segment I mostly red, with pale stripe medially; apex of scutellum with

orange-red or red markings …...... 3

- Antennal segment I uniformly red; apex of scutellum yellowish brown ca. 1/3rd of

length ...... P. cheesmannae

3. Pronotum red on anterior and lateral regions, pronotal disc broadly pale yellow

brown or yellowish brown ...... 4

- Pronotum mostly pale yellow brown, yellowish brown or red entirely ...... 7

4. Head generally brown with yellowish tinge; frons with red midline only; vertex with

red markings on posterior margin medially ...... P. gagnei

- Head dorsum including vertex pale yellow brown; frons mostly red ...... 5

5. Corium mostly red, about 2/3rd with posterior areas pale yellow brown or yellowish

brown ...... 6

- Corium mostly pale yellow brown with broad red horizontal stripe medially with

wings at rest ...... P. lenakelensis

6. Scutellum mostly red anteriorly, anterolateral angles and posteriorly yellowish

brown ………………………………………………………………………P. kraussi

- Scutellum pale yellow brown anteriorly, with medial to posterior areas mostly red

…….…………………………………………………………...P. vitiensis (Kirkaldy)

7. Body mostly greenish-yellow, clavus red on base to along outer areas adjacent to

endocorium, extending to apex; endocorium mostly red ………………...P. tatarnici

- Colouration not as above .…………………………………………………………...8

8. Scutellum uniformly pale yellow brown; cuneus pale yellow brown, lateral margin

tinged olive ……………………………………………...………...... P. tabwemasana

- Scutellum not uniformly pale yellow brown, with red markings; cuneus mostly pale

yellow brown with red markings …………….……………………………………...9

9. Pronotum broadly pale yellow brown with red tinge on callar region laterally;

corium mostly pale yellow brown with minor red markings on lateral areas adjacent

to apex of clavus ………….…………………………………………...... P. efatensis

- Pronotum mostly red with calli pale yellow brown; corium mostly red with

brownish markings, extending to apex of clavus and adjacent to membrane

…………….…………………………………………………………….P. naraiyawa

10. Mesoscutum with only anterolateral angles red ...…………………………………11

- Mesoscutum variously coloured, but anterolateral angles without red markings ....14

11. Hemelytra and scutellum mostly pale yellow brown or medium yellowish-brown

…...... 12

- Hemelytra and scutellum mostly red with pale yellow brown markings

……………………………………….……………………………………………..13

12. Clavus mostly medium yellowish-brown, reddish brown at base; exocorium entirely

reddish-brown; midline of scutellum orange-red; antennal segment III uniformly

pale yellow brown ...………………………………………………….P. baiawaensis

- Clavus mostly pale yellow brown, red on base and apex; base of corium and apex of

embolium red; scutellum uniformly pale yellow brown; apex of antennal segment

III red …………..…………………………………………………..…..P. woodwardi

13. Frons and vertex pale yellow brown with brownish tinge, frons tinged red laterally,

posterolateral angles of vertex reddish; scutellum with heart-shaped red marking;

corium mostly red; distal tip of cuneus red ……………..…………….P. cordiformis

- Frons pale yellow brown with red anteriorly; vertex uniformly pale yellow brown;

scutellum broadly red medially; corium about 2/3rd red, posterior areas pale yellow

brown; apex of cuneus red ca. ½ of length …………...... …………P. pancheriaphila

14. Abdomen generally pale yellow brown or yellowish brown; hind femora yellowish

brown ………………………………………………………………………………15

- Abdomen pale yellow brown with mostly red laterally; antennal segment II pale

yellow brown with red band on medial and apex; hind femora with orange-red band

near apex ...…………………………………………….…………..P. mourangeensis

15. Hind femora uniformly pale yellow brown or yellowish brown .....………………16

- Hind femora not uniformly pale yellow brown or yellowish brown, with red

markings ...... ……………18

16. Head dorsum entirely brownish; scutellum mostly orange-red; clavus broadly

brownish on basal half, tinged red; hemelytra mostly red; cuneus mostly red with

base angles pale yellow brown medially ………..…….……………………P. wareo

- Head dorsum mostly brown with frons and vertex tinged red laterally ...…………17

17. Pronotum, scutellum and clavus uniformly brown; apex of clavus pale yellow

brown; corium mostly red about ½ basal; antennal segment II uniformly yellowish-

brown ...……………………………………...... P. solomonensis

- Pronotum, scutellum and clavus uniformly pale yellow brown; apex of clavus red;

about 2/3rd of corium mostly red; antennal segment II pale yellow brown, base

tinged red .…….....………………………………………………………..P. sikaiana

18. Lateral areas and midline of pronotum with red stripes; head dorsum including frons

and vertex with red midline ………………..………………………………………19

- Pronotum mostly brownish anteriorly and laterally, disc broadly pale yellow brown

with lateral areas red; scutellum broadly red medially, apex brownish; hind femora

mostly red with basal area pale yellow brown ………...………………...P. kimiensis

19. Maxillary plate uniformly red; medial area of mandibular plate with narrow red

band; cuneus mostly red; medial part of hind femora with broad red band

……...…………………………………………………………………..P. straatmani

- Maxillary plate not uniformly red, with yellowish-brown markings posteriorly;

mandibular plate mostly red, posterior margin yellowish brown; cuneus pale yellow

brown with apex and basal margin red; apex of hind femora with narrow red band

..……………………………………………………………P. mcilwraithensis

Pseudoloxops baiawaensis new species

Figures 3.1, 3.3, 3.18

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: MALAYSIA: Sabah: Ranau, 5.9667°N

116.6833°E, 1176 m, 23 Aug 1983, G. F. Hevel and W. E. Steiner, (USI 00191326)

(USNM). PARATYPES: MALAYSIA: Sabah: Ranau, 5.9667°N 116.6833°E, 1176 m,

23 Aug 1983, G. F. Hevel and W. E. Steiner, 11♂ (USI 00191315 - USI 00191325), 8♂

(USI 00191327 - USI 00191334), 5♀ (USI 00191335 - USI 00191339) (USNM).

PAPUA NEW GUINEA: Milne Bay Province: Baiawa, Moi Biri Bay, 9.6°S

149.45°E, 30 m, 08 May 1953 - 11 May 1953, G. M. Tate, ♀ (AMNH_PBI00003985)

(AMNH). Morobe Province: NE Wau, 7.3333°S 146.71667°E, 1200 m, 12 Nov 1976,

W. C. Gagne, ♀ (USI 00042729) (BPBM).

Diagnosis. Pseudoloxops baiawaensis is recognised by the following combination of characters: hemelytra mostly yellowish-brown, with exocorium entirely reddish-brown, extending to cuneus; endocorium with minor red markings at base and lateral areas adjacent to apex of clavus; scutellum with orange-red midline, not reaching anterior margin; membrane brownish posteriorly; genital capsule ventral right margin basally concave, with elongate ventral process, armed with short, spinelike processes; and, endosoma with two spicules, lightly sclerotized, smooth, without serrations.

Description. Male: Body elongate-ovate, total length 3.33-3.60 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: generally yellowish-brown; frons mostly yellowish- brown with frons laterally tinged reddish brown; vertex yellowish brown; clypeus mostly yellowish-brown with red midline, extending to about midpoint of clypeus with narrow latitudinal red stripe, apically yellowish brown, basal part laterally tinged red; mandibular plate broadly red medially, with anterior and posterior regions yellowish brown; maxillary plate entirely reddish-brown; lateral aspect of antennal insertion reddish. Antennae: AI mostly reddish-brown, medial regions paler, with yellowish- brown stripe; AII mostly pale yellow brown, base reddish brown; AIII and AIV pale yellow brown. Labium: mostly yellowish-brown, apex of segment IV dark brown.

Pronotum: lateral areas broadly reddish brown, reaching calli, extending to dorsal margin of propleura, posterolateral angles margin brownish, broadly yellowish-brown medially. Propleuron: proepimeron mostly yellowish-brown, dorsal area with reddish- brown markings horizontally; proepisternum reddish-brown anteriorly, yellowish- brown posteriorly. Mesoscutum: mostly yellowish-brown, anterolateral angles reddish.

Scutellum: mostly yellowish-brown, midline to apical region orange-red, not reaching anterior margin. Hemelytra: clavus mostly yellowish-brown with reddish-brown markings basally, reaching about level of anterior margin of scutellum; exocorium reddish brown entirely; endocorium mostly yellowish-brown, base and medial areas adjacent to apex of clavus reddish brown; costal fracture reddish brown; cuneus mostly reddish-brown with yellowish-brown markings on anterior areas adjacent to membrane and outer base angles; membrane pale with posteriorly brownish, veins mostly pale with reddish markings posteriorly. Legs: yellowish brown. Abdomen: yellowish brown.

VESTITURE. Dorsum densely setose with pale, long to elongate, semierect to decumbent, bristlelike setae. Antennae: AI densely setose with dark, short to moderately

long, semierect to decumbent, bristlelike setae, with a few elongate, semierect, pale and dark setae, intermixed sparsely with short, erect, pale setae; AII and AIII densely setose with pale, short, decumbent to nearly adpressed setae, intermixed sparsely with semierect, pale setae, AII basally with a few moderately long, semierect, pale setae;

AIV densely setose with short, semierect, pale setae, intermixed with a few short, erect, pale setae. Pleura and sterna: almost bare with a few long, simple, pale setae. Legs: femora densely setose with pale, moderately long to elongate, semierect, simple setae, intermixed with a few elongate, semierect, simple, pale setae; tibiae densely setose with short, semierect to nearly adpressed, pale setae, intermixed sparsely with long, semierect, spinelike, pale setae, alternately elongate on hind tibiae, apex with a few short, pale, spinelike setae; hind tibiae with several longitudinal, parallel rows of tiny spinules. Abdomen: densely setose with pale, elongate, semierect, simple setae.

STRUCTURE. Head: subquadrangular; frons convex in dorsal view, moderately projected anteriorly, dorsum flat; vertex flat, midline deeply sulcate, with slightly impressed line posteriorly, margin slightly carinate; frontoclypeal boundary deeply dissected; clypeus weakly visible in dorsal view, enlarged, margin slightly convex basally in lateral view, then straight to apex; mandibular plate moderately sized, barely visible in dorsal view, margin weakly convex; maxillary plate moderately sized, ovoid, greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: reaching apex of mesocoxae; labial formula LII>LI>LIII>LIV; LI longer than bucculae, reaching midpoint of proxyphus. Eyes: moderately sized; not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum.

Antennae: antennifers moderately sized, reaching posterior part of mandibular plate, not reaching maxillary plate; AI long, nearly cylindrical, slightly enlarged subbasally, AI

longer than IOD, 1.4-1.8x longer than IOD, shorter than head width; AII cylindrical,

3.1-3.6x longer than AI, 1.5-1.6x longer than pronotum width; AIII and AIV slender;

AII>AIII>AIV>AI. Pronotum: trapeziform; width 2.3-2.8x length, 1.4-1.5x as wide as head; anterior part slightly raised, rounded, with impressed transverse line, anterior margin weakly concave medially; posterior margin slightly bisinuate; lateral margin nearly straight; callar region deeply depressed medially, calli slightly raised, with impressed line; disc flat. Proepimera: flat. Mesoscutum: exposed, ca. ½ of scutellum length. Scutellum: disc almost flat, width 1.2-1.3x length. Hemelytra: translucent; MFL weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing clavus, reaching anterior region of cuneus. Male genitalia: genital capsule with weakly concave dorsal margin, nearly straight, ventral margin strongly convex, right basal margin concave, with elongate ventral process, armed with short, spinelike processes (Fig. 3.3a); right paramere club-shaped, apophysis unarmed

(Fig. 3.3b); left paramere Y-shaped, sensory lobe thumblike, apex rounded, unarmed, with rounded basal process, apophysis parallel to sensory lobe, ca. 45° to sensory lobe, elongate, subbasally enlarged, moderately hooked subapically, apex rounded (Fig.

3.3c); phallotheca nearly cylindrical, without process, opening apical to dextrodorsal region (Figs. 3.3d, e); endosoma with two spicules, lightly sclerotized, DES elongate, simple, smooth margin, somewhat straight then recurved apically, tapering to apex,

VES short, simple, ventral to DES, position of spicules from left to right-lateral, close to secondary gonopore, wrapped with cuplike membranous process (Fig. 3.3f).

Female: Total length 3.43-4.04 mm. COLOURATION. Mostly as in male (Fig. 3.1).

VESTITURE. As in male. STRUCTURE. Mostly as in male; Antennae: AI longer than

IOD, 1.6-1.9x longer than IOD; AII 2.7-3.0x longer than AI, 1.5-1.7x longer than pronotum width. Pronotum: 2.5-2.6x as wide as long, 1.5-1.7x as wide as head.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Papua New Guinea and Sabah, Malaysia. This species is broadly distributed across the Wallace Line, which is similar with distributions seen in a number of mirids taxa such as within the genus Coridromius as reported by Tatarnic and Cassis (2008).

Remarks. Pseudoloxops baiawaensis is closely allied to the Solomon Island species, P. solomonensis, based on male genitalic structures. Characters shared by both species include: the shape of the phallotheca (narrowed apically in P. solomonensis), the elongate ventral process on the pygophore, the Y-shaped left paramere, the right paramere without a medial process, and, the presence of two endosomal spicules.

However, P. baiawaensis is distinguished from P. solomonensis by having a weakly concave dorsal margin of pygophore, whereas it is moderately concave in P. solomonensis. In addition, the ventral margin of the pygophore is strongly convex in P. baiawaensis. Also, P. baiawaensis has the pronotum broadly yellowish-brown medially with the lateral areas broadly reddish brown, whereas it is mostly brown in P. solomonensis. The clavus in P. baiawaensis is mostly yellowish-brown with red markings basally as compared to mostly brown in P. solomonensis. The corium of this species is mostly yellowish-brown with the exocorium reddish brown entirely, whereas in P. solomonensis it is mostly brown with red markings anteriorly and yellowish- brown posteriorly. P. baiawaensis can also be distinguished from P. solomonensis by having a mostly reddish-brown with yellowish-brown markings cuneus, whereas in P.

solomonensis it is mostly pale yellow brown with red markings at the apex and mediobasal angles.

Pseudoloxops cheesmanae new species

Figures 3.1, 3.19

Etymology. This species is named for the collector, Evelyn Cheesman.

Type material. HOLOTYPE: Male: VANUATU: Tafea: Aneityum Is. Co.: Red Crest,

3 m. N.E. of Anelgauhat, 20.19867°S 169.79803°E, 366 m, Jun 1955, L. E. Cheesman

(USI 00377681) (BMNH).

Diagnosis. Pseudoloxops cheesmanae is recognised by the following combination of characters: frons mostly red anteriorly with yellowish-brown markings anteromedially; vertex uniformly yellowish-brown; AI uniformly red; mesoscutum uniformly yellowish- brown; broad red band on anterior part of scutellum; and, apex of cuneus yellowish brown.

Description. Male: Body elongate, subparallel, total length 4.76 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: mostly yellowish-brown; frons mostly red laterally extending anteriorly, yellowish-brown anteromedially; vertex yellowish brown; clypeus mostly yellowish-brown, with dorsum laterally extending to midline red, posteriorly with red horizontal stripe, ca. level of anterior margin of maxillary plate, extreme tip yellowish brown; mandibular plate mostly red medially, latitudinal, with anterior and posterior area yellow-brown; maxillary plate mostly red, posterior area with yellowish- brown markings medially; lateral aspect of antennal insertion red. Antennae: AI uniformly red; AII yellow-brown, base tinged red; AIII and AIV yellow-brown.

Labium: mostly yellowish-brown, apex of segment IV dark brown. Pronotum: lateral

areas broadly red, reaching calli, extending to dorsal margin of propleura, lateral margin tinged yellowish-brown; anterior areas mostly red; callar region medially yellowish- brown; disc broadly yellowish brown. Propleuron: proepimeron mostly yellowish- brown with narrow red markings dorsally, latitudinal; proepisternum red anteriorly, with yellowish-brown posteriorly. Mesoscutum: uniformly yellowish-brown. Scutellum: broad red band on anterior part of scutellum, with yellowish-brown apically, about slightly more than 1/3rd length. Hemelytra: clavus red basally, reaching anterior region of scutellum, ca. 1/3rd length, remainder broadly yellowish-brown, apical part red about

1/4th of claval commissure; corium mostly red, reaching about apex of clavus, not reaching tip, posterior part mostly yellowish-brown, posterior margin contiguous with cuneus mostly red; costal fracture reddish; embolium to costal margin mostly yellowish-brown, distal end reddish; cuneus mostly yellowish-brown, with red markings mostly at base, with lateral angles yellowish brown, posterior region mostly red with apex yellowish brown; membrane yellowish brown, veins mostly red posteriorly with anterior part adjacent to corium yellowish brown. Abdomen: yellowish brown. Legs: yellowish brown. VESTITURE. Dorsum setose with pale, shiny, short to long, decumbent to semierect, simple setae, intermixed with elongate, simple, pale setae, mostly on head, embolium and cuneus. Antennae: AI densely setose with pale, short to long, decumbent to semierect, simple setae, intermixed sparsely with pale, elongate, semierect bristlelike setae; AII densely setose with pale, short, decumbent to nearly adpressed setae ,intermixed with a few pale, short, semierect setae; AIII and AV densely setose with pale, short, decumbent setae, intermixed with a few short, semierect, pale setae. Legs: femora densely setose with pale, short to moderately long, semierect bristlelike setae, intermixed with a few pale, long, somewhat erect bristlelike setae; tibiae densely setose with pale, short, nearly adpressed, bristlelike setae

,intermixed with a few pale, short, semierect bristlelike setae, remainder sparsely setose with smoky brown, long, suberect, spinelike setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, nearly adpressed bristlelike setae, intermixed with a few pale, long, semierect bristlelike setae. Abdomen: densely setose with pale, short to elongate, semierect, simple setae. STRUCTURE.

Head: subquadrangular; frons convex in dorsal view, somewhat rounded, moderately projected anteromedially, somewhat swollen, remainder almost flat; vertex almost flat in lateral view, midline deeply sulcate, with slightly impressed line posteriorly, margin slightly carinate; clypeus visible in dorsal view, enlarged; mandibular plate moderately sized, visible in dorsal view, margin weakly convex; maxillary plate greatly tumescent, exceeding mandibular plate; bucculae unknown. Labium: not present. Eyes: moderately sized, reaching dorsal margin of head in lateral view, reaching bucculae ventrally, exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized; AI long, slightly enlarged subbasally, AI longer than IOD, 1.5x longer than IOD, shorter than head width; AII cylindrical, 3.2x longer than AI, 1.6x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform, width 2.4x length, 1.4x as wide as head; anterior margin slightly raised, rounded, weakly concave medially; lateral margin somewhat straight, posterior margin slightly excavate; callar region depressed medially, calli slightly raised, with impressed line; disc almost flat. Proepimeron: flat.

Mesoscutum: exposed, about ½ scutellum length. Scutellum: disc almost flat, width

1.3x length. Hemelytra: MFL moderately long, weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed, apex rounded; membrane cells moderately long. Abdomen:

surpassing clavus, about reaching anterior margin of cuneus. Legs: hind femora enlarged, about 2x middle femora; tibiae cylindrical. Male genitalia: Not examined.

Female: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Restricted to Tafea Island, Vanuatu.

Remarks. Pseudoloxops cheesmanae is known from only a single specimen. This species is similar to P. kraussi in colouration, sharing the same red patterning as follows: broadly on anterior and lateral regions of the pronotum, the corium mostly with posterior region yellowish brown and clavus apically. However, P. cheesmanae is distinguished from P. kraussi by having the AI uniformly red, the yellowish-brown apex of the cuneus and the clavus with red only at the base; whereas in P. kraussi, AI is mostly red with a pale stripe medially, the apex of the cuneus is red, with the red markings on the basal regions extending along the medial regions of the clavus, reaching beyond the apex of the scutellum. P. cheesmanae also differs in having a broad red band on the scutellum anteriorly, which is trapeziform in P. kraussi.

Pseudoloxops cordiformis new species

Figures 3.1, 3.19

Etymology. This species is named from the Latin cordiformis, meaning “heart-shaped”, in reference to the heart-shaped red pattern on the scutellum.

Type material. HOLOTYPE: Male: AUSTRALIA: Queensland: North Queensland

Co.: Stephens Is., 9.5°S 143.55°E, 25 Nov 1986 - 27 Nov 1986, Houston & Sadler,

Light Trap (USI 00199279) (QDPI).

Diagnosis. Pseudoloxops cordiformis is recognised by the following combination of characters: frons mostly pale yellow brown with brownish tinge, reddish laterally; heart- shaped red marking on scutellum; M-shaped red pattern on corium with broad red band posteriorly adjacent to membrane; clavus and endocorium with brownish tinge; distal tip of cuneus sanguineous; and, membrane veins pale yellow brown with red spots medially on anterior and posterior margins.

Description. Male: body elongate, subparallel, total length 3.69 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: generally pale yellow brown; frons mostly pale yellow brown, with brownish tinge, reddish laterally adjacent to eyes; vertex pale yellow brown, posterior angles adjacent to eyes reddish; clypeus with narrow red stripe anteriorly, midline red, extending about mid-clypeus with broad latitudinal red stripe, apically pale yellow brown; mandibular plate broad red horizontal marking at midpoint, with anterior and posterior margins pale yellow brown; maxillary plate mostly red, with small, pale yellow brown spot posteriorly; lateral aspect of antennal insertion reddish.

Antennae: AI mostly red, with pale yellow brown stripe; AII pale yellow brown, base red; AIII and AIV uniformly pale yellow brown. Labium: mostly pale yellow brown, apex of segment IV dark brown. Pronotum: lateral regions broadly red, margins tinged pale brown, reaching calli laterally, extending to dorsal margin of propleura, broadly pale yellow brown medially. Propleuron: proepimeron mostly pale yellow brown with pale brown band dorsally, intermixed with narrow red stripe; proepisternum red anteriorly, with pale yellow brown posteriorly. Mesoscutum: mostly pale yellow brown, anterolateral angles red. Scutellum: large heart-shaped red marking, with pale yellow brown markings on anterolateral angles and anteromedial region. Hemelytra: clavus red basally, reaching anterior margin of scutellum, remainder broadly pale yellow brown with brownish tinge, apically red ca. ½ of claval commissure; exocorium mostly red,

reaching about level anterior region of membrane, posteriorly pale yellow brown, endocorium mostly red, reaching ca. ½ of claval commissure, followed by broadly pale yellow brown to about anterior part of membrane, with brownish tinge, remainder with broad red band, posteriorly pale yellow brown; cuneus pale yellow brown, apex sanguineous; membrane pale yellow brown, veins pale yellow brown with red spots on anterior margin and posterior margin medially. Legs: front legs pale yellow brown; missing middle and hind legs. Abdomen: pale yellow brown. VESTITURE. Dorsum densely setose with pale, shiny, moderately to elongate, decumbent, bristlelike setae, intermixed with elongate, semierect to decumbent, bristlelike, pale setae. Antennae: AI densely setose with dark, short to moderately long, semierect to decumbent, bristlelike setae, distally with a few long, semierect pale setae, intermixed sparsely with short, semierect, pale setae; AII and AIII densely setose with pale short, decumbent to adpressed setae, intermixed with a few short, semierect, pale setae; AIV densely setose with pale, short, nearly adpressed setae, intermixed with a few short, semierect, pale setae. Pleura and sterna: almost bare with a few scattered, elongate, pale setae. Legs: forefemora densely setose with pale, short to moderately long, semierect, setae, intermixed with a few elongate, semierect, pale setae. Abdomen: densely setose with pale, elongate, semierect, simple setae; apex of ventral region of pygophore with short setae, blunt tips. STRUCTURE. Head: subquadrangular; frons narrowly convex in dorsal view, somewhat rounded, moderately projected anteromedially, remainder flat; vertex flat, midline deeply sulcate, with slightly impressed line posteriorly, margin slightly carinate; frontoclypeal boundary deeply dissected; clypeus slightly visible in dorsal view, enlarged, margin slightly convex basally in lateral view, then straight to apex, basal part slightly bicompressed laterally; mandibular plate moderately sized, visible in dorsal view, margin weakly convex; maxillary plate moderately sized, ovoid,

greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of eyes. Labium: reaching apex of metacoxae; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching anterior part of proxyphus. Eyes: moderately sized; reaching dorsal margin of head in lateral view, reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum.

Antennae: antennifers moderately sized, reaching posterior margin of mandibular plate;

AI moderately long, slightly enlarged subbasally, AI longer than IOD, 1.9x longer than

IOD, shorter than head width; AII cylindrical, ca. 2.9x longer than AI, 1.7x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform; width 2.3x length, 1.4x as wide as head; anterior part slightly raised, rounded, anterior margin weakly concave medially; posterior margin slightly excavate; lateral margin straight; callar region slightly depressed medially, calli weakly raised, with moderately impressed line; disc flat. Proepimera: flat. Mesoscutum: exposed, ca. ½ scutellum length. Scutellum: disc flat, width 1.2x length. Hemelytra: MFL moderately long, weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing corium, approaching anterior part of cuneus. Male genitalia: genital capsule with moderately concave dorsal margin, ventral margin convex, with apical margin slightly weakly sinuate apically; right paramere with dorsal medial process, short, armed, apophysis elongate, sickle-shaped, apex rounded; left paramere

Y-shaped, sensory lobe thumblike, apex rounded, with rounded basal process, apophysis elongate, slim, parallel to sensory lobe, sinuate subapically, apex hooked, distal tip acute.

Female: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Australia.

Remarks. Pseudoloxops cordiformis is known from only a single specimen. This species can be differentiated from other species by the unique red colour patterning on the hemelytra, the heart-shaped red marking on the scutellum, and the M-shaped red markings on the corium. P. cordiformis is also differentiated from other Southwest

Pacific Pseudoloxops species by only having the distal tip of the cuneus sanguineous, the clavus and endocorium with a brownish tinge, and the membrane veins with red spots on the medial regions of the anterior and posterior margins. This species is allied to P. baiawaensis, P. pancheriaphila, P. tatarnici, P. woodwardi by having the pronotum broadly red only on the lateral regions. In addition, P. cordiformis also shares the presence of red markings on the basal and apical regions of the clavus with P. pancheriaphila and P. tatarnici.

Pseudoloxops efatensis new species

Figures 3.1, 3.4, 3.19

Etymology. This species is named after the island of the type locality.

Type material. HOLOTYPE: Male: VANUATU: Shefa: Efate Is. Co.: Vila, Efate Is,

0-100 m, 17.74097°S 168.32099°E, Feb 1977, N. L. H. Krauss (USI 00042886)

(BPBM). PARATYPES: VANUATU: Malampa: Ambrym Is. Co.: Ranon to Mount

Toyo, 16.15949°S 168.13365°E, 400 m, 02 Sep 1979, W. C. Gagne, 1♂ (USI

00043011) (BPBM). Shefa: Efate Is. Co.: Vila, Efate Is, 0-100 m, 17.74097°S

168.32099°E, Jan 1976, N. L. H. Krauss, 1♀ (USI 00042887) (BPBM).

Diagnosis. Pseudoloxops efatensis is recognised by the following combination of characters: body mostly pale yellow brown; clypeus pale yellow brown with pale red tinge laterally; callar region with red tinge laterally; scutellum broadly red with apex pale yellow brown; apex of clavus red; and, narrow red markings on endocorium adjacent to apex of clavus.

Description. Male: Body elongate, subparallel, total length 4.07–4.26 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: generally yellowish-brown; frons with red markings laterally, extending anteriorly; vertex yellowish brown; clypeus mostly pale yellow brown, basal region with pale red tinge laterally; mandibular plate mostly pale yellow brown anteriorly, with red band medially, posterior areas narrowly pale yellow brown; maxillary plate mostly red anteriorly, posterior areas with pale yellow brown markings medially; antennal insertion tinged red laterally. Antennae: AI mostly red, medially paler, with pale yellow brown stripe; AII mostly pale yellow brown, with red markings on base and apex; AIII generally pale yellow brown with red apex; AIV uniformly pale yellow brown. Labium: mostly pale yellow brown, apical part of segment IV dark brown. Pronotum: mostly pale yellow brown, lateral regions anteriorly with narrow red markings, reaching about middle of pronotum. Propleura: proepimera mostly pale yellow brown with narrow horizontal red markings dorsally; proepisterna red anteriorly, with pale yellow brown posteriorly. Mesoscutum: uniformly pale yellow brown.

Scutellum: mostly red, apex pale yellow brown, ca. 1/3rd length. Hemelytra: clavus mostly pale yellow brown with red markings apically, ca. 1/3rd length of claval commissure; corium mostly pale yellow brown with red markings medially, contiguous with apex of clavus; cuneus mostly pale yellow brown, basal angles red medially, apically ca. 1/3rd length red with yellowish apex; membrane pale yellow, veins mostly pale yellow with red markings posteriorly. Abdomen: uniformly pale yellow brown.

Legs: uniformly pale yellow brown. VESTITURE. Dorsum setose with pale, shiny, moderately long to elongate, semierect to decumbent, bristlelike setae, intermixed with pale, elongate, simple setae. Antennae: AI densely setose with dark, long, semierect to decumbent, simple setae, intermixed sparsely with pale, elongate, erect, simple setae;

AII densely setose with pale, short, decumbent to adpressed setae, intermixed with short to elongate, semierect, pale setae, basally with a few elongate, semierect, pale setae;

AIII and AIV densely setose with short, decumbent to adpressed setae, pale setae, intermixed sparsely with short, semierect setae. Pleura and sterna: almost bare with a few pale, long, simple setae. Legs: femora densely setose with pale, short to elongate, semierect, simple setae, intermixed with a few pale, elongate, semierect, simple setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with pale, elongate, semierect, spinelike setae, apex with a few short, spinelike, pale setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, semierect setae, intermixed with a few elongate, semierect, simple, pale setae. Abdomen: densely setose with pale, elongate, semierect, simple setae. STRUCTURE. Head: subquadrangular; frons narrowly convex in dorsal view, moderately projected anteromedially; vertex flat, midline deeply sulcate, with slightly impressed line posteriorly, posterior margin weakly carinate; frontoclypeal boundary weakly dissected; clypeus weakly visible in dorsal view, enlarged, anterior margin weakly convex in lateral view; mandibular plate moderately sized, exposed in dorsal view, margin weakly convex; maxillary plate moderately sized, somewhat ovoid, moderately tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: reaching apex of metacoxae; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching posterior part of proxyphus. Eyes: moderately sized; reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally;

exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, reaching posterior part of mandibular plate; AI elongate, slightly enlarged subbasally, AI 1.7–2.3x longer than

IOD, shorter than head width across eyes; AII cylindrical, 2.9–3.0x longer than AI, 1.4–

1.7x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform, 2.3–2.6x wider than long, 1.4–1.5x wider than head; anterior margin slightly raised, rounded, weakly concave medially; lateral margins somewhat straight; posterior margin slightly excavate; callar region depressed medially, calli slightly raised, with impressed line; disc flat. Proepimera: weakly inflexed. Mesoscutum: exposed, ca. ½ scutellum length. Scutellum: disc flat, 1.1–1.2x wider than long.

Hemelytra: surpassing apex of abdomen, costal margin weakly sinuate basally; MFL moderately long, weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, ca. 1.5x longer than wide, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing clavus, about reaching anterior margin of cuneus. Male genitalia: genital capsule with convex ventral margin; right paramere with dorsal medial process, perpendicular to apophysis, recurved apically, armed with small teeth, apophysis longer than medial process, apex bulbous, denticulate (Fig. 3.4a); left paramere Y-shaped, sensory lobe thumblike, moderately broad, rounded apically, subbasal left margin with rounded process, apophysis moderately narrow, parallel to sensory lobe, ca. 45° to sensory lobe, slightly longer than sensory lobe, weakly sinuate subapically, apex rounded, crenulated (Fig. 3.4b); phallotheca nearly cylindrical, opening apical to right dorsolateral, ventral surface with left subapical fold, enlarged, rounded, dorsal surface without lobe.

Female: Total length 4.15 mm. COLOURATION. Mostly as male; Head: generally pale yellow brown; with intense red markings on frons and clypeus compared to male.

VESTITURE. As in male. STRUCTURE. Mostly as in male. Eyes: not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI 1.5x longer than IOD; AII 2.9x longer than AI, 1.6x longer than pronotum width. Pronotum:

2.6x wider than long, 1.5x wider than head.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Restricted to Vanuatu.

Remarks. Pseudoloxops efatensis is similar to P. lenakelensis by having a mostly pale yellow brown body, with the scutellum mostly red and the clavus red apically.

Pseudoloxops efatensis also shares with P. lenakelensis the presence of red markings on the medial part of the corium, which is narrow in P. efatensis, and not extending to the exocorium. P. efatensis is distinguished from P. lenakelensis by having only a red tinge on the lateral regions of the pronotum, which is broadly red in P. efatensis, and the right paramere has a shorter apophysis, with a bulbous apex.

Pseudoloxops gagnei new species

Figures 3.1, 3.18

Etymology. This species is named after the collector, the late Dr Wayne Gagne.

Type material. HOLOTYPE: Female: PAPUA NEW GUINEA: Morobe Province:

NE Wau, Morobe District, Mt. Missim, 7.1167°S 146.9167°E, 1600 m, 23 Aug 1978,

W. C. Gagne (USI 00042896) (BPBM).

Diagnosis. Pseudoloxops gagnei is recognised by the following combination of characters: large size; head mostly brown with yellowish tinge; frons with red midline; the posterior margin of vertex red medially; clypeus with red markings basally;

maxillary plate yellowish brown, tinged red anteriorly; pronotum red laterally with yellowish-brown tinge; callar region mostly brownish medially with red midline; disc broadly yellowish-brown medially; diamond-shaped red pattern on scutellum, apex yellowish brown, tinged red; clavus mostly brownish tinged red on half basal part, distal tip of apex red; corium mostly red, posteriorly pale yellow brown; and, hind femora mostly red, paler on apex, yellowish-brown basally.

Description. Female: Body elongate, sub-parallel, total length 5.75 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: mostly brownish with yellowish tinge; frons with red markings on midline; vertex red on posterior medially; clypeus mostly yellowish brown, with red markings basally; mandibular plate broadly red with posterior margin yellowish brown; maxillary plate mostly yellowish-brown, anteriorly tinged red; lateral aspect of antennal insertions reddish. Antennae: AI mostly red, medially yellowish brown with red tinge; AII mostly pale yellow brown, paler at base, tinged red apically;

AIII and AIV pale yellow brown. Labium: mostly yellowish-brown, apex of segment IV dark brown. Pronotum: lateral areas mostly red with yellowish-brown tinge; callar region mostly brownish medially, midline red; disc broadly yellowish-brown medially.

Propleura: proepimeron mostly pale yellow brown, with red horizontal markings at dorsal part; proepisterna red anteriorly, pale yellow brown posteriorly. Mesoscutum: uniformly yellowish-brown. Scutellum: mostly red, diamond-shaped, with anterior part mostly pale yellow brown on anterolateral part, apex yellowish brown, tinged red.

Hemelytra: clavus mostly smoky brown on basal part, reaching beyond apex of scutellum, tinged red at base and apex of scutellum, pale yellow brown towards apex; corium mostly red, reaching level ca. 1/3rd from apex of clavus, broadly pale yellow brown towards posterior margins, lateral areas adjacent to anterior margin of membrane with red spots, posterior part adjacent to medial basal angles of cuneus red; embolium

mostly smoky brown, reaching ca. 1/3rd from apex of clavus, pale yellow brown posteriorly, with red markings basally; cuneus mostly pale yellow brown with orange tinge medially, with red markings on mediobasal angles and apex; costal fracture mostly red; membrane smoky brown, veins mostly red posteriorly, pale anteriorly.

Legs: mostly yellowish-brown; metafemora mostly red, about 2/3rd length, paler on apex, with basal part yellowish brown; metatibiae basally tinged red. Abdomen: yellowish brown. VESTITURE. Dorsum densely setose with pale, elongate, semierect to decumbent, simple setae, intermixed with elongate, semierect, simple, pale setae, mostly on head, pronotum and embolium, with moderately long setae on cuneus.

Antennae: AI densely setose with iridescent, moderately long to elongate, semierect to decumbent, bristlelike setae, intermixed with a few iridescent, elongate, erect, bristlelike setae; AII and AIII densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short to elongate, semierect, simple, pale setae; AIV clothed with short, semierect to nearly adpressed setae, intermixed sparsely with elongate, semierect, simple, pale setae. Pleura and sterna: almost bare with a few pale, long, simple setae. Legs: femora densely setose with pale, moderately long to elongate, semierect, simple setae, intermixed with a few pale, elongate, semierect setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with pale, long, semierect, spinelike setae, elongate on hind tibiae, apex with a few short, smoky brown, spinelike setae; hind tibiae with several longitudinal parallel rows of tiny spinules; tarsi densely setose with pale, short, semierect to adpressed setae, intermixed with a few long, semierect, pale setae. Abdomen: venter densely setose with pale, elongate, semierect, simple setae. STRUCTURE. Head: subquadrangular; frons slightly convex in dorsal view, slightly projected anteriorly; vertex almost flat, midline deeply sulcate, with slightly impressed line posteriorly, posterior margin carinate;

frontoclypeal boundary weakly dissected; clypeus visible in dorsal view, enlarged, margin slightly convex; mandibular plate large, visible in dorsal view, margin rounded; maxillary plate moderately sized, somewhat ovoid, greatly tumescent, exceeding mandibular plate in dorsal view, anteriorly exceeding posterior margin of mandibular plate from anterior view; bucculae about reaching eyes posteriorly. Labium: surpassing mesocoxae, almost reaching trochanter; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching anterior part of proxyphus. Eyes: moderately sized, reaching dorsal margin of head in lateral view, not reaching bucculae ventrally; slightly exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, nearly reaching posterior margin of mandibular plate; AI elongate, slightly enlarged subbasally, AI ca. 2x longer than IOD, shorter than head width; AII cylindrical, 3.2x longer than AI, 1.6x longer than pronotum width; AIII and AIV slender; AII>AIII>AI>AIV. Pronotum: trapeziform; 2.1x wider than long, 1.7x wider than head; anterior part slightly raised medially, anterior margin weakly concave medially; posterior margin rectilinear; lateral margin nearly straight; callar region depressed medially, calli weakly raised, with impressed line; disc almost flat, slightly convex in lateral view. Proepimera: flat. Mesoscutum: exposed, nearly ½ of scutellum length. Scutellum: disc weakly raised, width 1.2x greater than length.

Hemelytra: MFL weakly convex, reaching beyond apex of scutellum, not reaching 1/3rd of claval commissure; cuneus narrow and long, ca. 2x width; membrane cells long.

Male: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Restricted to Papua New Guinea.

Remarks. Pseudoloxops gagnei is known only from the holotype. This species is similar in appearance to P. kimiensis. Characters shared by both species include: large size, the head mostly brownish with red markings on the frons and posteromedial region of the vertex, the mandibular plate mostly red with posterior margin yellowish brown, the clavus brownish and tinged red on the basal half with the apex red, and the hind femora mostly red, but with the apex yellowish brown in P. gagnei. This species can be differentiated by the maxillary plate being yellowish brown with a red tinge anteriorly, which in contrast is mostly red in P. kimiensis.

Pseudoloxops kimiensis new species

Figures 3.1, 3.5, 3.18

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: PAPUA NEW GUINEA: Eastern Highlands:

No.9, Kimi Creek Camp, NE slopes Mount Michael, 6.41666°S 145.33333°E, 1980 m,

29 Aug 1959, L. J. Brass (USI 00042895) (BPBM).

Diagnosis. Pseudoloxops kimiensis is recognized by the following combination of characters: large size; head mostly brown with yellowish tinge; maxillary plate mostly red with posterior part adjacent to clypeus yellowish brown; pronotum mostly brownish on anterior and lateral regions, broadly pale yellow brown disc with outer areas red; scutellum broadly red medially with apex brownish; corium mostly red with broad roundish pale yellow brown markings posteriorly on endocorium; hind femora mostly red with basal part pale yellow brown; genital capsule with weakly projected ventral margin; right paramere with ventral medial process, perpendicular to apophysis,

serrated margins, apophysis armed; and, sickle-shaped left paramere with projected basal process, apophysis broad.

Description. Male: Body elongate, sub-parallel, total length 5.31 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: generally brownish; frons and vertex mostly brownish with yellow tinge on medial and lateral part, with red markings on midline of frons and posterior margin of vertex medially; clypeus mostly yellowish-brown, with red stripes on basal part and posterior part, not reaching apex; mandibular plate broadly red with posterior margin yellowish brown; maxillary plate mostly red, posterior part adjacent to clypeus yellowish brown; lateral aspect of antennal insertion tinged red.

Antennae: AI mostly red, laterally yellowish brown with red tinge; AII mostly pale yellow brown, paler at base; AIII pale yellow brown; missing AIV. Labium: mostly pale yellow brown, apex of segment IV dark brown. Pronotum: mostly brown on anterior and lateral regions, midline of anterior margin with red spot; disc medially with roundish pale yellow brown markings, with broad bright red markings laterally.

Propleura: proepimeron mostly yellowish-brown, dorsal part with red horizontal band; proepisterna red anteriorly, yellowish-brown posteriorly. Mesoscutum: mostly pale yellow brown with broad pale red markings medially. Scutellum: broadly red medially with anterior part mostly pale yellow brown, apex brownish. Hemelytra: clavus mostly pale yellow brown, basally smoky brown with reddish markings, reaching beyond apex of scutellum, apex red; exocorium entirely red, paler posteriorly, endocorium mostly red, reaching about apex of clavus then broadly with roundish pale yellow brown, posterior margin red, lateral regions adjacent to apex of clavus with red markings, extending to anterior margin of membrane; cuneus mostly pale yellow brown with mediobasal angles red, apical 1/3rd of cuneus red; membrane smoky brown, veins bright red. Legs: mostly pale yellow brown, femora apically and tibiae basally with orange

tinge; metafemora mostly red, ca. 2/3rd length, with basal region pale yellow brown; metatibiae with red band at base. Abdomen: pale yellow brown. VESTITURE. Dorsum densely setose with pale, elongate, semierect to decumbent, simple setae, intermixed with elongate, semierect, simple, pale setae, mostly on head, pronotum and embolium, with moderately long setae on cuneus. Antennae: AI densely setose with iridescent, moderately long to elongate, semierect to decumbent, bristlelike setae, intermixed with a few iridescent, elongate, erect, bristlelike setae; AII and AIII densely setose with pale, short, decumbent to adpressed setae, intermixed with a few short, semierect, pale setae.

Pleura and sterna: almost bare with a few pale, elongate, simple setae. Legs: femora densely setose with pale, moderately long to elongate, semierect, simple setae, intermixed with a few pale, elongate, semierect setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with pale, elongate, semierect, spinelike setae, apex with a few short, spinelike setae; tarsi densely setose with pale, short, semierect to adpressed setae, intermixed with a few long, semierect, pale setae. Abdomen: venter densely setose with pale, short to elongate, semierect, simple setae. STRUCTURE. Head: subquadrangular; frons slightly convex in dorsal view, slightly projected anteromedially, dorsum almost flat; vertex flat, midline deeply sulcate, with slightly impressed line posteriorly, margin carinate; frontoclypeal boundary weakly dissected; clypeus visible in dorsal view, enlarged, margin weakly convex; mandibular plate large, visible in dorsal view, margin rounded; maxillary plate moderately sized, ovoid, moderately tumescent, exceeding mandibular plate in dorsal view, not exceeding mandibular plate anteriorly from anterior view; bucculae about reaching eyes. Labium: surpassing mesocoxae, about reaching trochanter; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching anterior part of proxyphus. Eyes: large, reaching dorsal margin of head in lateral view, reaching

bucculae ventrally; exceeding anterolateral angles of pronotum, ca. ½ eyes width, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, nearly reaching posterior margin of mandibular plate; AI long, slightly enlarged subbasally, AI longer than IOD, 1.9x longer than IOD, shorter than head width; AII cylindrical, 3.6x longer than AI, 1.8x longer than pronotum width; AIII slender.

Pronotum: trapeziform; 2.2x as wide as long, 1.4x as wide as head; anterior part raised medially, rounded, anterior margin weakly concave medially; posterior margin rectilinear; lateral margin nearly straight; callar region depressed medially, calli weakly raised, with impressed line; disc almost flat, slightly convex in lateral view. Proepimera: flat. Mesoscutum: exposed, shorter than ½ of scutellum length. Scutellum: disc slightly raised, width 1.2x length. Hemelytra: MFL weakly convex, reaching beyond apex of scutellum, not reaching 1/3rd of claval commissure; cuneus narrow and long, ca. 2x width; membrane cells long. Male genitalia: genital capsule with moderately concave dorsal margin, ventral margin weakly projected, nearly straight (Fig. 3.5a); right paramere with ventral medial process, moderately broad and long, perpendicular to apophysis, with serrate margins, apex rounded, apophysis shorter than sensory lobe, armed with teeth, apex rounded, armed (Fig. 3.5b); left paramere somewhat sickle- shaped, sensory lobe slightly developed, rounded, with basal process, somewhat rectangular, armed, apex bulbous, apophysis elongate, broad, perpendicular to sensory lobe, apex hooked, distal tip rounded (Fig. 3.5c); phallotheca nearly cylindrical, sinuate margins, apically rounded (Fig. 3.5d).

Female: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Papua New Guinea.

Remarks. Pseudoloxops kimiensis is known only from the holotype. P. kimiensis is similar to P. gagnei, but can be distinguished from it by colour characters. See

‘Remarks’ section of P. gagnei for differential diagnosis.

Pseudoloxops kraussi new species

Figures 3.1, 3.6, 3.19

Etymology. This species is named after the collector, N.L.H. Krauss.

Type material. HOLOTYPE: Male: VANUATU: Shefa: Efate Is. Co.: Vila, Efate Is,

0-100 m, 17.74097°S 168.32099°E, Feb 1970, N. L. H. Krauss, (USI 00042891)

(BPBM). PARATYPES: VANUATU: Penama: Maewo Is. Co.: Sounwari, 15.38056°S

168.12989°E, 18 m, 04 Sep 1979, W. C. Gagne, ♂ (USI 00042889) (BPBM). Sanma:

Espiritu Santo Is. Co.: Luganville, 15.52739°S 167.1639°E, 2 m, 24 Aug 1957,

Unknown, ♂ (USI 00042890) (BPBM). Shefa: Efate Is. Co.: NW: Limestone Plateau,

N of Maat, 17.67393°S 168.2608°E, 100 m, 18 Aug 1957, J. L. Gressitt, ♀ (USI

00042893) (BPBM). Shefa: Shepherds Islands Co.: Emae I.: Mt. Tavani Talimasa, 0-

434 m, 17.08333°S 168.36666°E, 30 Aug 1979, W. C. Gagne, Acacia simplex

(Fabaceae), ♂ (USI 00042881), 3♀ (USI 00042882 - USI 00042884) (BPBM). Tafea:

Tanna Is. Co.: 2 km S Ipeukeul, 275-350 m, 19.51511°S 169.45466°E, 23 Aug 1979,

G. A. Samuelson, ♀ (USI 00042885) (BPBM). Lenakel, 0-200 m, 19.51921°S

169.28217°E, 150 m, Jan 1973, N. L. H. Krauss, ♀ (USI 00042892) (BPBM); Mar

1980, N. L. H. Krauss, ♂ (USI 00042888) (BPBM).

Diagnosis. Pseudoloxops kraussi is recognized by the following combination of characters: head dorsum mostly pale yellow brown; AII yellowish brown, with base and apex tinged red; AIII yellowish brown, apex tinged red; pronotum broadly red on

anterior and lateral regions; mesoscutum uniformly pale yellow brown; scutellum mostly red basally, anterolateral angles and apical part yellowish brown; clavus red from base extending along medial areas adjacent to beyond apex of scutellum, apex red; corium broadly red until apex of clavus, with posterior part pale yellow brown; right paramere with dorsal medial process, short, sinuous margin, apophysis longer than medial process, armed; left paramere Y-shaped, with moderately broad basal process, nearly truncate margin, apophysis with pointed distal tip; phallotheca ventrally with subrectangular fold, sinuous margin, dorsal surface with right apical lobe and subbasal fold, moderately sized; and, endosoma with single spicule, slightly hooked subapically.

Description. Male: Body elongate, subparallel, total length 3.90-4.04 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: dorsum mostly pale yellow brown; frons dorsum mostly red laterally, extending to anterior part with posterior areas adjacent to mandibular plate and clypeus pale yellow brown; vertex pale yellow brown; clypeus mostly yellow-brown with red midline, basal part laterally tinged red, then posterior part with red horizontal stripe about level anterior margin of maxillary plate, extreme tip yellow-brown; mandibular plate broadly red medially, latitudinal, with anterior and posterior areas yellow-brown; maxillary plate mostly red, tinged yellow-brown medially; lateral aspect of antennal insertion with red markings. Antennae: AI mostly red, medial areas paler with yellowish-brown stripe; AII yellowish brown, with base and apex tinged red; AIII yellowish brown, with apex tinged red; AIV uniformly yellowish-brown. Labium: mostly yellowish-brown, apical part of segment IV dark brown. Pronotum: lateral areas broadly red with yellowish-brown tinge, reaching calli mostly, somewhat extending to dorsal margin of propleura, anterior areas mostly red; callar region medially yellowish-brown extending broadly to disc medially. Propleura: proepimera mostly yellowish-brown with narrow red markings dorsally, latitudinal;

proepisterna red anteriorly, with yellowish-brown posteriorly. Mesoscutum: uniformly yellowish-brown. Scutellum: basal part mostly red with anterolateral areas yellowish- brown, apical part mostly yellowish-brown, about ½ length. Hemelytra: clavus red basally, reaching about level anterior margin of scutellum, then extending along medial areas of clavus adjacent to scutellum, reaching beyond apex of scutellum, then broadly yellowish-brown, apical part red ca. 1/3rd of claval commissure; corium mostly red, reaching about apex of clavus, not reaching tip, then posterior part mostly yellowish- brown with posterior margin adjacent to cuneus mostly red, with yellowish-brown markings posteriorly adjacent to membrane; costal fracture red; embolium mostly yellowish-brown, distal end reddish; cuneus mostly yellowish-brown basally, about ½ length, then mostly red to apex, anterior margin red from about anteromedially to mediobasal angles adjacent to membrane; membrane yellowish brown, veins mostly red with anterior part ca. 1/3rd length yellowish brown, anterior margins adjacent to corium tinged red. Abdomen: yellowish brown. Legs: yellowish brown. VESTITURE. Dorsum setose with pale, shiny, short to long, semierect to decumbent, simple setae, intermixed with elongate, semierect, simple, pale setae. Antennae: AI densely setose with pale, moderately long, decumbent setae, intermixed sparsely with elongate, semierect, pale setae; AII densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short, semierect, pale setae, then with a few pale, long, semierect setae;

AIII and AIV densely setose with pale, long, decumbent to adpressed setae, intermixed sparsely with short, semierect, pale setae. Pleura and sterna: almost bare with a few pale, long, simple setae. Legs: femora densely setose with pale, short to long, semierect, simple setae, intermixed sparsely with elongate, semierect, simple, pale setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with pale, long, semierect setae, alternately, longer on hind tibiae; with several

longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, semierect setae, intermixed with a few long, semierect, simple, pale setae. Abdomen: densely setose with pale, elongate, semierect, simple setae. STRUCTURE. Head: subquadrangular; frons widely convex in dorsal view, somewhat rounded, slightly projected in front at middle part, somewhat swollen, dorsum almost flat; vertex flat, midline deeply sulcate, with slightly impressed line posteriorly, posterior margin weakly carinate; frontoclypeal boundary weakly dissected; clypeus slightly visible in dorsal view, enlarged, with margin slightly convex basally in lateral view; mandibular plate moderately sized, barely visible in dorsal view, margin weakly convex; maxillary plate moderately sized, somewhat ovoid, moderately tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: reaching apex of metacoxae; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching posterior part of proxyphus. Eyes: moderately sized, reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, reaching posterior region of mandibular plate, not reaching anterior margin of maxillary plate; AI long, slightly enlarged subbasally, AI 1.4-1.7x longer than

IOD, shorter than head width; AII cylindrical, 3.1-3.4x longer than AI, 1.4-1.6x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform; 2.4-2.6x wider than long, 1.3-1.5x wider than head; anterior margin slightly raised, rounded, weakly concave medially; lateral margin somewhat straight, posterior margin slightly excavate; callar region depressed medially, calli slightly raised, with impressed line; disc flat. Proepimera: weakly inflexed. Mesoscutum: exposed, ca. ½ of scutellum length. Scutellum: disc slightly raised, 1.1-1.3x wider than long. Hemelytra: MFL moderately long, reaching ca. 1/3rd of claval commissure,

weakly convex; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing corium, reaching midpoint of cuneus. Male genitalia: genital capsule with concave dorsal margin, ventral margin strongly convex (Fig. 3.6a); right paramere with dorsal medial process, short, sinuous margin, apophysis longer than medial process, apically armed with robust teeth, apex rounded (Fig. 3.6b); left paramere Y-shaped, sensory lobe thumblike, apex rounded, with basal process, enlarged, with nearly truncate margin, apophysis slim, parallel to sensory lobe, ca. 45° to sensory lobe, slightly longer in length to sensory lobe, moderately hooked subapically, apex rounded, with pointed tip (Fig. 3.6c); phallotheca subcylindrical, opening apical to dextrodorsal region, ventral surface with lobe, subrectangular, with weakly sinuous margin, dorsal surface with right apical lobe, small, rounded margin, with right subbasal fold, moderately sized, thumblike (Figs.

3.6d, e); endosoma with single spicule, moderately long, sclerotized, slender, weakly hooked subapically, tapered apically, position left to right hand side dorsolateral, adjacent to secondary gonopore (Fig. 3.6f).

Female: Total length 3.95-4.42 mm. Similar colour and vestiture as in male (Fig. 3.1).

STRUCTURE. Mostly as in male. Antennae: AI1.4-1.6x longer than IOD; AII 3.0-3.1x longer than AI, 1.3-1.6x longer than pronotum width. Pronotum: 2.5-2.8x wider than long, 1.5-1.7x wider than head.

MEASUREMENTS. Table. 1.

Host plant. Three specimens from Mt. Tavani Talimasa were found on Acacia simplex

(Sparmm.) Pedley (Fabaceae).

Distribution. Restricted to Vanuatu.

Remarks. Pseudoloxops kraussi is mostly closely related to P. cheesmanae. See P. cheesmanae ‘Remarks’ section for differential diagnosis.

Pseudoloxops lenakelensis new species

Figures 3.1, 3.7, 3.19

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: VANUATU: Tafea: Tanna Is. Co.: Lenakel, 0-

200 m, 19.51921°S 169.28217°E, 150 m, Mar 1980, N. L. H. Krauss (USI 00042880)

(BPBM).

Diagnosis. Pseudoloxops lenakelensis is recognised by the following combination of characters: body mostly pale yellow brown, frons mostly red; maxillary plate mostly red with brownish tinge; pronotum red on anterior and lateral regions, posterolateral margins pale yellow brown; scutellum mostly red, apex pale yellow brown; apex of clavus red, greenish tinge along medial region of clavus adjacent to mesoscutum and scutellum; broad red stripe on corium; right paramere with elongate apophysis, sickle- shaped; left paramere apophysis armed with small teeth subapically; phallotheca nearly cylindrical, with large sinistroventral subapical fold, and small armed sinistrodorsal subapical lobe; and, endosoma with single, elongate, fingerlike spicule, membranous basally.

Description. Male: Body elongate, subparallel, total length 4.29 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: frons mostly red; vertex pale yellow brown; clypeus mostly yellowish-brown, with red horizontal stripe, about level anterior margin of maxillary plate, midline tinged red anteriorly; mandibular plate mostly red medially, not reaching clypeus, with anterior and posterior areas yellowish brown; maxillary plate mostly red, with brownish tinge; lateral aspect of antennal insertion red. Antennae: AI reddish-pale yellow; AII and AIII generally pale yellow with red apex; AIV missing.

Labium: mostly yellowish-brown, apex of segment IV dark brown. Pronotum: lateral areas broadly red, reaching calli, extending to dorsal margin of propleura; anterior areas mostly red; midline of callar region pale yellow brown, extending to disc with broadly pale yellow brown colouration. Propleuron: proepimeron mostly yellowish-brown with narrow red markings dorsally, latitudinal; proepisternum red anteriorly, with yellowish- brown posteriorly. Mesoscutum: uniformly pale yellow brown. Scutellum: mostly red, apex pale yellow brown. Hemelytra: clavus mostly yellowish with pale red markings apically, ca. 1/3rd length of claval commissure; corium mostly yellowish with broad horizontal pale red stripe, ca. 1/3rd length of claval commissure; cuneus mostly yellowish with mediobasal angles red, apical part ca. 1/3rd length pale red with yellowish apex; membrane pale yellow, veins mostly pale yellow with red markings posteromedial region. Abdomen: uniformly yellow-brown. Legs: front leg yellow- brown; middle and hind legs missing. VESTITURE. Dorsum setose with pale, shiny, moderately long, decumbent to semierect, bristlelike setae, intermixed with elongate setae, mostly on head, pronotum, embolium and cuneus; clypeus setose with pale, short, simple setae. Antennae: AI densely setose with dark, moderately long, decumbent to semierect bristlelike setae, intermixed with pale, short, erect setae, apically with a few pale, elongate, erect bristlelike setae; AII densely setose with pale, short to moderately, decumbent to adpressed setae, intermixed with a few pale, short, semierect bristlelike setae; AIII densely setose with pale, short, decumbent setae, intermixed with a few pale, short, semierect, pale setae. Pleura and sterna: almost devoid with a few scattered, elongate, simple, pale setae. Legs: missing middle and hind legs; forefemora densely setose with pale, short to moderately long, semierect bristlelike setae, intermixed with a few pale, elongate, suberect bristlelike setae; protibiae densely setose with pale, short, nearly adpressed, bristlelike setae, intermixed sparsely with pale, moderately long,

semierect bristlelike setae, then with a few pale, long, somewhat erect, bristlelike setae; tarsi setose with pale, short, semierect, bristlelike setae. Abdomen: venter densely setose with pale, long to elongate, semierect, simple setae; pygophore ventral apically with short setae, blunt tip. STRUCTURE. Head: subquadrangular; frons convex in dorsal view, somewhat rounded, moderately projected anteromedially, somewhat swollen, dorsum almost flat; vertex almost flat in lateral view, midline deeply sulcate, with slightly impressed line posteriorly, margin slightly carinate; fronclypeal boundary moderately dissected; clypeus visible in dorsal view, enlarged, slightly convex in lateral view; mandibular plate moderately sized, visible in dorsal view, margin weakly convex; maxillary plate moderately sized, somewhat ovoid, greatly tumescent, exceeding mandibular plate; bucculae small, reaching posterior margin of eyes. Labium: reaching apex of metacoxae; labial formula LII>LI>LIV=LIII; LI longer than bucculae, reaching posterior part of proxyphus. Eyes: oval in lateral view, moderately sized, nearly reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, reaching posterior area of mandibular plate, not reaching anterior margin of maxillary plate, adjacent to eyes; AI long, slightly enlarged subbasally, AI 1.8x longer than IOD, shorter than head width;

AII cylindrical, ca. 3x longer than AI, 1.7x longer than pronotal width; AIII slender.

Pronotum: trapeziform, 2.7x wider than length, 1.3x as wide as head; anterior part slightly raised, rounded, with impressed horizontal line, anterior margin weakly concave medially; posterior margin slightly excavate, lateral margin straight; callar region depressed medially, calli slightly raised, with impressed line; disc flat.

Proepimera: weakly inflexed. Mesoscutum: exposed, ca. ½ scutellum length, posterior areas slightly depressed medially, deflexed laterally, weakly convex in lateral view.

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Scutellum: disc almost flat, width 1.2x length. Hemelytra: translucent; MFL moderately long, weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long,

1.5x longer than wide, not greatly narrowed, apex rounded; membrane cells moderately long. Abdomen: surpassing corium, about reaching anterior areas of cuneus. Legs: missing middle and hind legs. Male genitalia: genital capsule subconical, with concave dorsal margin, ventral margin convex (Fig. 3.7a); right paramere with dorsal medial process, armed, apophysis elongate, sickle-shaped, apex rounded, armed with teeth (Fig.

3.7b); left paramere Y-shaped, sensory lobe thumblike, apex rounded, with rounded basal process, apophysis elongate, slim, strongly sinuate subapically, margin armed with small teeth, apex acute (Fig. 3.7c); phallotheca nearly cylindrical, opening dextrodorsally, ventral surface with left subapical fold, enlarged, rounded, margin denticulate, dorsal surface with left subapical lobe, small, convex, armed, right-lateral opening with subapical fold, convex, dextrodorsal margin denticulate subbasally (Fig.

3.7d); endosoma with fingerlike single spicule, apex rounded, smooth margin, membranous basally, positioned left to dextrodorsal, distal to secondary gonopore (Fig.

3.7e).

Female: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Restricted to Vanuatu.

Remarks. Pseudoloxops lenakelensis is known from only the holotype. This species is similar in colouration to P. efatensis by having mostly a pale yellowish-brown body with a mostly red scutellum and the clavus is red apically. Pseudoloxops lenakelensis and P. efatensis also have a red band on the medial part of the corium, which is broader in the former species, extending to the embolium. Pseudoloxops lenakelensis can also

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be distinguished from P. efatensis by having broad red markings on the anterior and lateral regions of the pronotum, whereas in P. efatensis it is mostly pale yellowish- brown, with a reddish tinge on the lateral margins, and the elongate apophysis of the right paramere.

Pseudoloxops mcilwraithensis new species

Figures 3.1, 3.8, 3.19

Etymology. This species is named after the type locality, Mcllwraith Range.

Type material. HOLOTYPE: Male: AUSTRALIA: QUEENSLAND: Mcllwraith Ra.,

N.E. Coen, Cape York Pen., 13.75°S 143.33333°E, 29 Jun 1976 - 05 Jul 1976, J. F.

Donaldson, (USI 00199277) (QDPI). PARATYPES: AUSTRALIA: Queensland:

North Queensland Co.: Mt. Tozer, North Foot 400', 12.75°S 143.21638°E, 122 m, 01

Jul 1948 - 05 Jul 1948, L. J. Brass, 1♂ (USI 00042680) (BPBM). Leo Creek Rd., ca.

500m., Mcllwraith Range, 30 km N.E. of Coen., 13.75°S 143.33333°E, 29 Jun 1976 -

04 Jul 1976, G. B. & S. R. Monteith, 1♂ (USI 00192980), 1♀ (USI 00192981) (UQIC).

Mcllwraith Ra., N.E. Coen, Cape York Pen., 13.75°S 143.33333°E, 29 Jun 1976 - 05

Jul 1976, J. F. Donaldson, 1♀ (USI 00199278) (QDPI).

Diagnosis. Pseudoloxops mcilwraithensis is recognised by the following combination of characters: head mostly yellow-brown with red midline, without orange markings; AI uniformly red; pronotum with red midline, without orange markings; mesoscutum with broad red markings at medial; scutellum with red markings at midline; corium mostly red, with yellow-brown markings; metafemora with minor red markings apically; pygophore without processes; right paramere with unarmed apophysis and sensory lobe;

102

left paramere sickle-shaped, hooked apically; and, endosoma with three spicules, DES1 and DES2 with divided branches.

Description. Male: Body elongate-ovate, total length 4.09-4.53 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: mostly yellow-brown with red stripe at midline, extending from anterior margin of frons to posterior margin of vertex; clypeus mostly yellow-brown with red horizontal markings medially (ca. 1/3rd of length), apex yellow- brown; mandibular plate mostly red with posterior margin yellow-brown; maxillary plate mostly red with posterior margin yellow-brown; outer antennal insertion with red markings. Antennae: AI uniformly red; AII and AIII uniformly yellow-brown; AIV yellow-brown, apex brownish. Pronotum: mostly yellow-brown with red stripes at lateral areas and midline; lateral margin yellow-brown. Propleura: proepimera mostly yellow-brown, dorsal areas with red horizontal markings; proepisterna mostly red with posterior areas yellow-brown. Mesoscutum: yellow-brown with broad red markings medially. Scutellum: mostly yellow-brown with red markings at midline, extending to apex (ca. 1/3rd length). Hemelytra: clavus mostly yellow-brown, basal margin red extending to medial areas of clavus adjacent to scutellum, reaching apical part of scutellum, claval commissure mostly red, apex red (ca. 1/5th of claval commissure); medial areas of corium adjacent to claval suture mostly red, lateral areas adjacent to

R+M vein extending to posterior margin of corium mostly red, medial part extending to areas beyond apex of clavus adjacent to claval suture mostly pale yellow brown; embolium and costal margin mostly pale yellow brown at posterior margin, base red, half part to posterior areas mostly red with pale yellow brown at posterior margin, apex red; cuneus mostly pale yellow brown, apex red (ca. 1/3rd of length), basal margin mostly red with areas adjacent to laterobasal angles pale yellow brown; membrane pale yellow brown, veins red. Abdomen: mostly yellow-brown, lateral part brownish. Legs:

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mostly yellow-brown, metafemora tinged red apically, with narrow red bands at apex.

VESTITURE. Dorsum setose with pale, silky, moderately long to elongate, decumbent to semierect, simple setae. Antennae: AI densely setose with pale, moderately long, decumbent bristlelike setae, intermixed sparsely with pale, elongate, semierect bristlelike setae; AII and AIII densely setose with pale, short, decumbent to adpressed bristlelike setae, intermixed sparsely with pale, short, semierect setae; AIV missing.

Pleura and sterna: almost bare with a few scattered, pale, silky, long, simple setae.

Abdomen: venter densely setose with pale, silky, elongate, semierect, simple setae, denser laterally. Legs: femora densely setose with pale, silky, moderately long, semierect setae, intermixed sparsely with pale, long, semierect setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with smoky brown, elongate, spinelike, semierect setae, alternately; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, decumbent to semierect setae, intermixed with a few long, semierect setae.

STRUCTURE. Head: frons weakly convex in dorsal view, rounded, weakly projected in front anteriorly, dorsum flat; vertex flat in lateral view, midline weakly sulcate, posterior margin slightly carinate; clypeus visible in dorsal view, subrectangular; mandibular plate moderately sized, visible in dorsal view, margin weakly convex, weakly rounded; maxillary plate moderately sized, tumescent, slightly visible in dorsal view, convex; bucculae reaching posterior margin of head. Labium: reaching apex of metacoxae; LI longer than bucculae, reaching proxyphus. Eyes: moderately sized, reaching dorsal margin of head in lateral view, reaching bucculae ventrally; exceeding anterolateral angles of pronotum (ca. 1/3rd of eyes width), posterior margin of eyes contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plates; AI moderately long, AI 1.3-1.4x

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longer than IOD, shorter than head width; AII cylindrical, 3.7-4.4x longer than AI, 1.6-

1.7x longer than pronotum width. Pronotum: trapeziform; ca. 2.3x wider than long, 1.2-

1.3x as wide as head; anterior margin rounded, slightly concave medially; lateral margin nearly straight; posterior margin slightly excavate; calli slightly raised, with impressed line; disc flat. Proepimera: almost flat. Mesoscutum: exposed, ca. ½ scutellum length, weakly convex in lateral view. Scutellum: disc raised, 1.2-1.3x wider than long.

Hemelytra: MFL weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, longer than wide, not greatly narrowed; membrane cells moderately long. Abdomen: reaching posterior margin of corium. Male genitalia: pygophore with somewhat straight dorsal margin, ventral margin convex (Fig. 3.8a); right paramere weakly C-shaped, apophysis bulbous, unarmed, apex somewhat rounded, medial process shorter in length to apophysis, apically rounded (Fig. 3.8b); left paramere sickle-shaped, sensory lobe weakly developed, blunt, apophysis elongate, somewhat slim, right angles to sensory lobe, apical part hooked, base without prominent tumescence (Fig. 3.8c); phallotheca opening apical to dextrodorsal region, dorsal surface with granularlike processes (Fig. 3.8d); endosoma with three spicules, complex arrangement, DES1 elongate, broad basally, reduced at apex, unbranched, strongly sclerotized, with serrate apical margin, DES2 broad basally, with three branches, smooth margins, VES with three branches, two branches comblike, sclerotized, elongate, slender, with apex acute (Fig. 3.8e).

Female: Total length 4.70-4.71 mm. COLOURATION. Mostly as in male (Fig. 3.1).

VESTITURE. As in male. AIV densely setose with pale, short, decumbent to nearly adpressed setae, intermixed sparsely with pale, short, semierect setae. STRUCTURE.

Mostly as in male; Eyes: not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI moderately long, 1.1-1.2x longer than IOD;

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AII 3.7-3.8x longer than AI, AII 1.5-1.6x longer than pronotum width;

AII>AIII>AIV>AI. Pronotum: 2.4-2.6x wider than long, 1.4-1.5x wider than head.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Pseudoloxops mcilwraithensis is known from two localities in

Queensland (Mt. Tozer and Mcllwraith Range).

Remarks. Pseudoloxops mcilwraithensis is similar to P. mourangeensis, but differs by the following characters: the elongate-ovate body; the mandibular plate and maxillary plates mostly red with yellowish-brown posteriorly; AII uniformly pale yellow brown; the proepimera mostly yellowish-brown with red markings dorsally; the proepisterna mostly red with yellowish-brown posteriorly; and, the hind femora yellowish brown, tinged red apically, with a narrow red band on apex.

Pseudoloxops mourangeensis new species

Figure 3.1, 3.19

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: AUSTRALIA: Queensland: 2 miles N. of

Mourangee Stn. Edungalba, 23.71667°S, 149.84999°E, 22 May 1969, T. Campbell and

E. Adams (USI 00033753) (ANIC). PARATYPES: AUSTRALIA: Queensland: 2 miles N. of Mourangee Stn. Edungalba, 23.71667°S, 149.84999°E, 22 May 1969, T.

Campbell and E. Adams, 1♂ (USI 00033754) (ANIC), 1♀ (USI 00033755) (ANIC).

Diagnosis. Pseudoloxops mourangeensis is recognized by the following combination of characters: body elongate-subparallel; head dorsal surface mostly pale yellow brown, frons with orange-red markings anteriorly; vertex with roundish markings; clypeus,

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mandibular plate and maxillary plate red; AII with red bands at midpoint and apex; pronotum with orange-red markings on midline and lateral regions; corium generally pale yellow brown with red stripes; abdomen pale yellow brown with mostly red laterally; and, hind femora mostly pale yellow brown, with orange-red band near apex.

Description. Male: Body elongate, subparallel, total length 5.01-5.05 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: dorsum generally pale yellow brown with orange to red Y-shaped markings, midline red, roundish on posterior margin of vertex, frons anteriorly orange to red; mandibular plate, clypeus and maxillary plate red; antennal insertion red laterally. Antennae: antennifers pale yellow brown, tinged red; AI mostly red; AII mostly pale yellow brown, midpoint and apex red; missing AIII and AIV. Eyes: reddish-brown. Labium: mostly pale yellow brown, LIV mostly dark brown. Pronotum: pale yellow brown with orange-red markings at lateral areas, not reaching margin, reaching lateral part of calli; midline orange-red stripe. Propleura: proepimera mostly red, posterior areas pale yellow; proepisterna entirely red. Mesoscutum: generally pale yellow brown, with broad orange-red markings medially. Scutellum: mostly pale yellow brown, with red midline, apex ca. 1/3rd of length. Hemelytra: clavus mostly pale yellow brown, red basally, not exceeding claval vein, reaching about apex of scutellum, apical part red, claval suture red; endocorium mostly red at medical areas adjacent to clavus, reaching ca. 2/3rd of claval commissure, with red markings on areas adjacent to apex of clavus, medial areas between exocorium and endocorium pale yellow brown, posterior areas adjacent to membrane broadly pale yellow brown, exocorium mostly red, with embolium mostly pale yellow brown, apex red; cuneus mostly pale yellow brown, anterior areas adjacent to membrane broadly red, apical part red ca. 1/3rd of length; membrane smoky brown, veins mostly red posteriorly, pale anteriorly, lateral areas adjacent to medial vein tinged red. Sterna: mostly red. Abdomen: pale yellow brown

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with lateral areas mostly red. Legs: mostly pale yellow brown, hind femora with orange-red band near apex; apex of tarsi black. VESTITURE. Dorsum setose with pale, silky, moderately long to elongate, decumbent to semierect, simple setae, mostly on head and pronotum. Antennae: AI densely setose with pale, silky, long, semierect, simple setae, mostly short at base, intermixed with a few elongate, erect, pale setae; AII densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short, semierect, pale setae. Hemelytra: clavus and corium densely setose with pale, silky, moderately long, decumbent to nearly adpressed, simple setae, intermixed with pale, elongate, semierect setae; cuneus densely setose with pale, silky, moderately long to elongate, decumbent, simple setae. Pleura and sterna: almost bare with a few scattered, pale, simple setae. Abdomen: venter densely setose with pale, long, simple setae, intermixed with pale, elongate, semierect, simple setae, mostly on lateral. Legs: femora densely setose with pale, long, semierect setae, mostly short at apical part, intermixed sparsely with elongate, erect, pale setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with pale, elongate, semierect setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with short, semierect, pale setae, intermixed with a few long, semierect, pale setae. STRUCTURE. Head: frons convex in dorsal view, rounded, weakly projected anteromedially, dorsum slightly declivent; vertex almost flat in lateral view, midline weakly sulcate, posterior margin rounded, with slightly impressed line; clypeus visible in dorsal view, large, margin convex; mandibular plate moderately sized, barely visible in dorsal view, margin nearly straight; maxillary plate large, oval, moderately tumescent mostly anteriorly, barely visible in dorsal view; bucculae small, reaching posterior margin of eyes. Labium: surpassing metacoxae; LI longer than bucculae, reaching anterior part of proxyphus. Eyes: moderately sized; reaching dorsal

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margin of head in lateral view, reaching bucculae ventrally; exceeding anterolateral angles of pronotum in dorsal view, contiguous with anterior margin of pronotum.

Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plate, reaching anterior margin of eyes; AI slightly enlarged subbasally, moderately long, AI longer than IOD, ca. 1.4x longer than IOD, shorter than head width; AII cylindrical, ca. 3x longer than AI. Pronotum: trapeziform; 1.4x wider than head; anterior margin rounded, slightly raised, weakly concave medially; posterior margin slightly bisinuate; lateral margin straight; calli weakly raised, with impressed line; disc flat. Proepimera: almost flat. Mesoscutum: exposed, ca. ½ of scutellum length.

Hemelytra: MFL weakly convex, nearly reaching ½ length of claval commissure; cuneus narrow and long, ca. 2x longer than wide; membrane cells elongate, subrectangular. Abdomen: surpassing clavus, reaching posterior areas of corium. Male genitalia: Not examined.

Female: Total length 4.40 mm. COLOURATION. Mostly as in male but brighter, mostly sanguineous; Antennae: AI mostly sanguineous, medial part paler, with pale yellow brown stripe; AII mostly pale yellow brown, mostly red apically; STRUCTURE.

Mostly as in male. Eyes: reaching dorsal margin of head in lateral view, not reaching bucculae ventrally; AI moderately long, ca. 1.3x longer than IOD, AII 3.3x longer than

AI, 1.6x longer than pronotum width; missing AIV. Pronotum: 2.5x wider than long,

1.3x wider than head. Scutellum: 1.2x wider than long.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Queensland, Australia.

Remarks. Pseudoloxops mourangeensis can be distinguished from P. mcilwraithensis by the following characters: the body elongate-subparallel; clypeus, the mandibular

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plate and maxillary plates red; AII with red bands on the medial part and apex; the proepimera mostly red, pale yellow posteriorly; the proepisterna entirely red; the abdomen pale yellow brown which is mostly red laterally; and, the hind femora mostly pale yellow brown, with a orange-red band near apex.

Pseudoloxops naraiyawa new species

Figures 3.1, 3.9, 3.19

Etymology. This species is named after the type locality.

Type material. HOLOTYPE. Male: FIJI: Viti Levu: Naraiyawa, 17.56°S 178.5°E, 28

Nov 1986 - 30 Nov 1986, R. L. Brown, Light Trap, (USI 00003987) (MEM).

Diagnosis. Pseudoloxops naraiyawa is recognized by the following combination of characters: body elongate-subparallel; frons dorsum with narrow red stripes from midline to lateral margin; clypeus mostly pale yellow brown; pronotum mostly red tinged brown laterally, calli mostly pale yellow brown with red midline; hind femora broadly red from middle to apical part, apex pale yellow brown; pygophore with rounded left dorsal lobe; right paramere with dorsal medial process, armed, apophysis apically rounded, armed; the Y-shaped left paramere, with basal process, sensory lobe apically with a few short, spinelike setae; phallotheca with left subapical fold ventrally, dorsal surface right apical fold, large; and, endosoma with single spicule, elongate, not serrated, membranous basally.

Description. Male: Body elongate, subparallel, total length 4.78 mm (Fig. 3.1).

COLOURATION (Fig. 3.1). Head: frons with a few narrow red stripes from midline to lateral margin, extending to anterior margin, with anterior part medially adjacent to clypeus pale yellow brown, lateral margins red; vertex broadly pale yellow brown;

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clypeus mostly pale yellow brown; mandibular plate and maxillary plate mostly red; outer antennal reddish. Antennae: AI mostly pale yellow brown, with red tinge; AII and

AIII pale yellow brown; AIV missing. Eyes: reddish-brown. Labium: mostly pale yellow brown, LIV mostly dark brown apically. Pronotum: mostly red, lateral margins tinged brown, reaching calli laterally, extending to dorsal margin of propleura, calli mostly pale yellow brown, midline red. Propleura: proepimera mostly pale yellow brown, dorsal areas with red horizontal markings, tinged brown; proepisterna mostly red with brownish tinge anteriorly, mostly pale yellow brown posteriorly. Mesoscutum: mostly pale yellow brown. Scutellum: pale yellow brown with red tinge, apex pale yellow brown. Hemelytra: clavus red basally, reaching about midpoint of scutellum, broadly pale yellow brown at medial part, extending beyond apex of scutellum, reaching ca. 1/3rd of claval commissure, then broadly red, brownish apically; corium mostly red, reaching about apex of clavus, extending to lateral areas of clavus and areas adjacent to membrane, with brownish markings medially between endocorium and exocorium, pale yellow brown posteriorly, margins adjacent to cuneus red, embolium mostly pale yellow brown with red tinge, apex pale yellow brown; cuneus broadly pale yellow brown medially, anterior and lateral margins mostly red, apex red-brown; membrane smoky brown, veins red, lateral areas of large areole posteriorly with red markings. Sterna: pale yellow brown. Abdomen: pale yellow brown. Legs: mostly pale yellow brown; hind femora broadly red from about midpoint of femora to apical part, with apex pale yellow brown; apex of tarsi brownish. VESTITURE. Dorsum setose with pale, moderately long to elongate, semierect to decumbent setae. Antennae: AI densely setose with pale, short to moderately long, semierect to decumbent setae, intermixed sparsely with dark, elongate, semierect setae; AII and AIII densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short,

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semierect, pale setae; missing AIV. Pleura and sterna: almost bare with a few scattered, simple setae. Abdomen: densely setose with pale, elongate, semierect, simple setae, denser laterally, intermixed with short setae on pygophore ventrally. Legs: femora densely setose with pale, short to moderately long, semierect, simple setae, intermixed sparsely with pale, elongate, semierect, simple setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with smoky brown, elongate, semierect, spinelike setae, apex with a few short, smoky brown, spinelike setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, semierect to nearly adpressed setae, intermixed with a few, elongate, semierect, pale setae. STRUCTURE. Head: subquadrangular; frons weakly convex in dorsal view, almost flat, moderately projected anteromedially; vertex almost flat, midline deeply sulcate, posterior margin slightly carinate, with impressed line posteriorly; clypeus visible in dorsal view, large, subrectangular; mandibular plate moderately sized, barely visible in dorsal view, margin weakly convex; maxillary plate moderately sized, ovoid, greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: surpassing apex of metacoxae, reaching trochanter; labial formula LII>LI>LIII>LIV; LI longer than bucculae, reaching posterior margin of proxyphus. Eyes: moderately sized; not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plate, reaching interior margin of eyes; AI long, slightly enlarged subbasally, 1.8x longer than IOD, shorter than head width; AII cylindrical, 3.2x longer than AI, 1.7x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform; 2.3x wider than long, 1.6x as wide as head; anterior margin slightly raised, rounded, weakly

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concave medially; posterior margin slightly bisinuate; lateral margin nearly straight; callar region depressed medially, calli rounded, slightly raised, with impressed line; disc almost flat. Proepimera: flat. Mesoscutum: exposed, ca. ½ of scutellum length.

Scutellum: disc raised, width 1.2x length. Hemelytra: MFL moderately long, weakly convex, reaching ca. 1/3rd of claval commissure; cuneus narrow and long, ca. 2x width; membrane cells long. Abdomen: surpassing clavus, almost reaching anterior part of cuneus. Male genitalia: genital capsule with moderately concave dorsal margin, with medial lobe, with rounded left dorsal lobe, ventral margin projected, apically rounded

(Fig. 3.9a); right paramere with dorsomedial process, armed apically, apophysis longer in length than medial process, margin nearly truncate, apex rounded, apex to margin armed with small, sclerotized teeth (Fig. 3.9b); left paramere Y-shaped, sensory lobe thumblike, apex rounded, unarmed, with rounded basal process, sensory lobe apically with a few short, spinelike setae, apophysis elongate, slim, ca. 45° to sensory lobe, moderately hooked subapically, apex acute (Fig. 3.9c); phallotheca opening apical, dextrodorsal to medial, with fingerlike apical lobe, ventral surface with left subapical fold, large, bilobed, dorsal surface with right apical fold, large, rounded (Fig. 3.9d); endosoma with single spicule, spinelike, elongate, tapering to apex, heavily sclerotized, membranous basally, position left to dextrodorsal, proximal to secondary gonopore

(Fig. 3.9e).

Female: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Fiji.

Remarks. Pseudoloxops naraiyawa is known only from a single specimen. This species is closely allied to the Australian species, P. woodwardi, based on male genitalic

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structures. Characters shared by both species include: the shape of the pygophore with the presence of a medial process, the right paramere with a dorsal medial process, the deeply Y-shaped left paramere with a basal process, and the aedeagus with one endosomal spicule. However, P. naraiyawa is distinguished from P. woodwardi by having elongate apophysis of right paramere with rounded apex, whereas it is much broader in P. woodwardi. Additionally, the apophysis of the left paramere is broader than the latter. The phallotheca in P. naraiyawa is more complex compared to P. woodwardi. P. naraiyawa also has a more elongate and larger body than P. woodwardi.

Both species can be easily differentiated by colouration as P. naraiyawa has mostly red corium with brownish markings extending to apex of clavus and adjacent to membrane, whereas in P. woodwardi, with red markings only on basal part and distal tip of embolium. Apart from that, P. naraiyawa can be distinguished from the others by the mostly red pronotum with the calli pale yellow brown, and the pygophore with a rounded left dorsal lobe.

Pseudoloxops pancheriaphila new species

Figures 3.2, 3.10, 3.19

Etymology. This species is named after the plant genus Pancheria, from which the holotype specimen was collected.

Type material. HOLOTYPE: Male: NEW CALEDONIA: Province Sud: ca. 2 km E of Cap N'dua Lighthouse, 22.36348°S 166.9415°E, 47 m, 21 Apr 2005, Cassis, Wall,

Tatarnic, Monteith, Pancheria sp. (Cunoniaceae), det. NOU Staff NYBG Voucher (USI

00017681) (UNSW). PARATYPES. NEW CALEDONIA: Province Nord: Bai de

Sapins, Pindaii Peninsula, 21.35572°S 164.9732°E, 27 Apr 2005, Cassis, Wall,

Tatarnic, Monteith, Lumnitzera racemosa Willd. (Combretaceae), det. NOU Staff

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NYBG Voucher, 3♂ (USI 00017678 - USI 00017680), 12♀ (USI 00017684 - USI

00017695) (UNSW). Poindimié, 20.93333°S 165.33333°E, 50 m, Jan 1969, N. L. H.

Krauss, 1♂ (USI 00042696) (BPBM). Province Sud: 1.5 km N of Port Boisé,

22.34347°S 166.96388°E, 50 m, 20 Apr 2005, Cassis, Wall, Tatarnic, Monteith,

Gymnostoma sp. (Casuarinaceae), det. NOU Staff NYBG Voucher, 1♂ (USI 00017683)

Alphitonia neocaledonica Guill. (Rhamnaceae), det. NOU Staff NYBG Voucher, 1♀

(USI 00017697) (UNSW). Anse Vata, 22.31666°S 166.43305°E, 08 Nov 1958, C.R.

Joyce, 1♂ (USI 00042698) (BPBM); 16 Nov 1958, C.R. Joyce, Light Trap, 1♂ (USI

00042697) (BPBM); 14 Sep 1962, G.F. Gross, 1♂ (USI 00038835) (SAMA). Cap

N'dua Lighthouse, 22.3909°S 166.9244°E, 200 m, 21 Apr 2005, Cassis, Wall, Tatarnic,

Monteith, Acacia spirorbis Labill. (Fabaceae), det. NOU Staff NYBG Voucher, 1♀

(USI 00017698) (UNSW). Haute Yaté, 22.16805°S 166.90133°E, 411 m, 24 Apr 2005,

Cassis, Wall, Tatarnic, Monteith, Acacia spirorbis Labill. (Fabaceae), det. NOU Staff

NYBG Voucher, 1♂ (USI 00017682) (UNSW). Nouméa, 22.2667°S 166.45°E, 01 Jan

1700, A. M. Lea, 1♀ (USI 00038836) (SAMA); 03 Nov 1940, F. X. Williams, Light

Trap, 1♂ (USI 00042695) (BPBM). ca. 2 km E of Cap N'dua Lighthouse, 22.36348°S

166.9415°E, 47 m, 21 Apr 2005, Cassis, Wall, Tatarnic, Monteith, Pancheria sp.

(Cunoniaceae), det. NOU Staff NYBG Voucher, 1♀ (USI 00017696) (UNSW).

Nouméa, 0-100 m, 22.3067°S 166.45233°E, 50 m, Feb 1959, N. L. H. Krauss, 1♂ (USI

00042905) (BPBM); Feb 1976, N. L. H. Krauss, 1♀ (USI 00042998) (BPBM).

Diagnosis. Pseudoloxops pancheriaphila is recognized by the following combination of characters: head dorsum mostly pale yellow brown; frons with red markings anteriorly; scutellum broadly red medially, margins pale yellow brown; corium mostly red from base to beyond apex of clavus; apex of cuneus half apically red; pygophore subconical, ventral convex apically, with slightly sinuous margin, dorsal margin with lobe; the club-

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shaped right paramere; the Y-shaped left paramere, with basal process, apophysis elongate, longer than sensory lobe; phallotheca apically narrow, basally broad, with apical fold ventrally, dorsal surface with subapical lobe; and, endosoma with single spicule, spinelike, elongate, membranous basally.

Description. Male: Body elongate, subparallel, total length 4.07–4.23 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: dorsal surface mostly pale yellow brown with red markings at antennal insertion, extending to anterior areas of mandibular plate; frons red anterolaterally; clypeus narrowly red at midline to medial areas; mandibular plate medially with narrow red horizontal stripe; maxillary plate mostly red anteriorly.

Antennae: AI mostly red with medial areas medium yellow-brown; AII, AIII and AIV medium yellow-brown. Eyes: reddish-dark brown. Labium: mostly pale yellow brown;

LIV mostly dark brown apically. Pronotum: mostly pale yellow brown with lateral areas red, margin, intermixed with dark brown, extending to calli laterally, posterolateral angles dark. Propelura: proepimera mostly pale yellow brown, dorsal areas with narrow red markings; proepisterna mostly red, with posterior areas pale yellow brown.

Mesoscutum: mostly pale yellow brown, anterolateral angles red. Scutellum: mostly red medially, lateral margin and apex pale yellow brown. Hemelytra: clavus mostly pale yellow brown, with red markings at basal and apical regions; corium mostly pale yellow brown, with basal areas red extending along medial areas adjacent to clavus; cuneus pale yellow brown anteriorly, with red small red spot at mediobasal angle, with red apically (ca. ½ scutellum length). Sterna: pale yellow brown. Abdomen: pale yellow brown. Legs: medium yellow-brown; distal tip apex of tarsi blackish. VESTITURE.

Dorsum densely setose with elongate, decumbent to semierect, simple, shiny, pale setae mostly on head, pronotum and hemelytra. Antennae: AI densely setose with short to long (shorter than AI diameter), dark, semierect bristlelike setae, intermixed with

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elongate, pale, erect bristlelike setae; AII and AIII densely setose with pale, very short, decumbent to adpressed setae, intermixed sparsely with semierect setae, AII mostly with short setae basally intermixed with a few pale, long, erect setae; AIV clothed with pale, short (mostly longer than setae on AII & AIII), decumbent setae, intermixed with a few semierect setae. Pronotum: anterolateral areas with pale, elongate, erect setae.

Pleura and sterna: almost bare with a few scattered, elongate, simple setae. Legs: femora densely setose with pale, moderately long to elongate, semierect setae, intermixed with a few pale, elongate, erect setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with smoky brown, elongate, semierect, spinelike setae, alternately, apex with a few short to moderately long, smoky brown, spinelike setae; tarsi densely setose with pale, short, nearly adpressed setae, intermixed with a few elongate, semierect setae. Abdomen: venter with scattered, pale, elongate, semierect simple setae, denser laterally. STRUCTURE. Head: frons strongly convex in dorsal view, strongly projected anteriorly, dorsum flat; vertex flat in lateral view, midline deeply sulcate, posterior margin slightly carinate, with impressed line; frontoclypeal boundary deeply dissected; clypeus visible in dorsal view, large, convex basally; mandibular plate moderately sized, visible in dorsal view, margin weakly convex; maxillary plate moderately sized, tumescent, moderately, mostly medial, slightly excedding mandibular plate; bucculae reaching posterior margin of head.

Labium: reaching apex of metacoxae; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching posterior margin of proxyphus. Eyes: large; reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally; greatly exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum.

Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plate, reaching interior margin of eyes; AI longer than IOD, 1.5–1.8x longer

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than IOD, shorter than head width; AII cylindrical, 3.1–3.2x longer than AI, 1.6–1.7x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform, width 2.0–2.5x length, 1.3–1.4x wider than head; anterior margin slightly raised, rounded, weakly concave medially; posterior margin weakly excavate, with weakly bisinuate margin; lateral margin straight; callar region deeply depressed medially; calli weakly raised, with impressed line; disc flat. Proepimera: weakly inflexed. Mesoscutum: exposed, ca. ½ of scutellum length. Scutellum: disc almost flat, width 1.1–1.2x length. Hemelytra: MFL weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, reaching about midpoint of membran ca. 2x longer than wide, moderately narrowed; membrane cells moderately long. Abdomen: surpassing clavus, reaching anterior margin of cuneus. Male genitalia: pygophore with moderately concave dorsal margin, with rounded lobe, ventral margin convex, apical margin slightly sinuate (Fig. 3.10a); right paramere club-shaped, apophysis armed with teeth, rounded apically (Fig. 3.10b); left paramere Y-shaped, sensory lobe thumblike, unarmed, apex rounded, subbasal left margin with rounded process, apophysis elongate, slim, parallel to sensory lobe, moderately sinuate subapically, slightly longer in length to sensory lobe, apex slightly hooked, acute (Fig. 3.10c); phallotheca opening apical to dextrodorsal region, dorsal surface with subapical lobe, rounded, ventral surface with apical fold, subtriangular shaped, margin sinuous (Fig. 3.10d, e); endosoma with single spicule, sclerotized, spinelike, elongate, broad basally, tapering to apex, membranous basally; secondary gonopore wrapped with membrane, cuplike (Fig. 3.10f).

Female: Total length 4.11–4.30 mm. COLOURATION. As in male (Fig. 3.2).

VESTITURE. As in male. STRUCTURE. Mostly as in male. Eyes: not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI 1.4–1.7x

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longer than IOD; AII 2.8–2.9x longer than AI, 1.5–1.7x longer than pronotum width.

Pronotum: width 2.1–2.7x length, 1.5–1.6x as wide as head.

MEASUREMENTS. Table 3.1.

Host plant. Acacia spirorbis Labill. (Fabaceae), Alphitonia neocaledonica Guill.

(Rhamnaceae), Gymnostoma sp. (Casuarinaceae), Lumnitzera racemosa Willd.

(Combretaceae) and Pancheria sp. (Cunoniaceae).

Distribution. Found in New Caledonia.

Remarks. Pseudoloxops pancheriaphila is most similar to P. woodwardi on the basis of male genitalia. Characters shared by both species include: the genital capsule with a medial lobe on the dorsal margin; the Y-shaped left paramere with an elongate apophysis, which is slim, and longer than the sensory lobe; the endosoma with a single spicule, which is spinelike, elongate, not serrated, and membranous basally. In contrast, the spicule is nearly straight in P. pancheriaphila compared to subbasally recurved in P. woodwardi. Even though both species are similar in the shape of the left paramere, the sensory lobe is rounded in P. pancheriaphila compared to much broader in P. woodwardi. The right paramere is also different with the presence of a short dorsal medial process, whereas it is elongate in the latter species. Both species also share the following characters: the AI mostly red; the pronotum with red markings only on the lateral regions and the clavus with red markings on the basal and apical regions. P. pancheriaphilla can also be distinguished from P. woodwardi by having a broadly red marking on the scutellum and the endocorium is mostly red.

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Pseudoloxops sikaiana new species

Figures 3.2, 3.11, 3.18

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: SOLOMON ISLANDS: Malaita: Sikaiana, 0-

100 m, 8.43222°S 160.83777°E, Dec 1972, N. L. H. Krauss, (USI 00042918) (BPBM).

PARATYPES. SOLOMON ISLANDS: Bougainville: S. Bougainville, Guaba, 720 m,

19 Jun 1956, E.J. Ford, Jnr, Light Trap, ♀ (USI 00042922) (BPBM). Florida Group:

Vunuha, Small Nggela, 9.1833°S 160.2333°E, 19 Sep 1960, C. W. O'Brien, Light Trap,

♀ (USI 00042919) (BPBM). San Cristoval: Kira Kira, 10.44138°S 161.91666°E, 26

Jul 1960, C. W. O'Brien, Light Trap, ♀ (USI 00042920) (BPBM). Santa Ysabel:

Tatamba, 8.39361°S 159.81°E, 23 Jun 1960, C. W. O'Brien, Light Trap, ♀ (USI

00042921) (BPBM). Western Province: New Georgia Islands Co.: Mouth of Bareki

R., 30 Jul 1965, Unknown, ♂ (USI 00377682) (BMNH).

Diagnosis. Pseudoloxops sikaiana is recognized by the following combination of characters: dorsal surface of the head mostly brown with red markings laterally; maxillary plate reddish brown; anterior and lateral regions of pronotum mostly reddish- brown; mesoscutum with red markings on medial and lateral part; scutellum mostly pale yellow brown with apex red; clavus red on basal and apical part, corium broadly red tinged brownish, posteriorly pale yellow brown; apex and mediobasal of cuneus red; genital capsule with right ventral process, armed; right paramere with dorsal medial process, armed, apophysis broad, apically serrated; the Y-shaped left paramere, sensory lobe moderately broad, apophysis strongly hooked subapically, armed with small teeth; and, endosoma with two spicules, elongate, slender, tapering to apex, subbasally recurved, heavily sclerotized.

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Description. Male: Body elongate, subparallel, total length 3.99-4.10 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: generally brown; frons and vertex mostly brown, with red markings laterally, adjacent to eyes; clypeus mostly yellowish-brown with red midline, then with red horizontal stripe, about level of anterior margin of maxillary plate, basal part tinged red laterally; mandibular plate mostly yellowish-brown, medial part with narrow red horizontal stripe; maxillary plate reddish brown; lateral aspect of antennal insertion reddish. Antennae: AI mostly reddish yellow-brown; AII, AIII and

AIV yellowish brown. Labium: mostly yellowish-brown, apex of segment IV dark brown. Pronotum: mostly reddish-brown anteriorly and laterally, with disc broadly pale yellow brown medially. Propelura: proepimera mostly pale yellow brown, dorsal margin with reddish-brown band; proepisterna mostly reddish-brown anteriorly, pale yellow brown posteriorly. Mesoscutum: mostly pale yellow brown, with red markings at medial and lateral regions. Scutellum: mostly pale yellow brown anteriorly, ca. 2/3rd length, posteriorly red to apex. Hemelytra: clavus mostly red basally, tinged brownish, reaching about midpoint of scutellum, then broadly pale yellow brown to about mid- claval commissure, red towards apex; corium mostly red with brownish tinge, reaching about apex of clavus, then broadly pale yellow brown posteriorly, with red markings on posterior part adjacent to membrane anteriorly; cuneus mostly pale yellow brown, apex and mediobasal angles red; membrane smoky brown, veins mostly red. Abdomen: yellowish brown. Legs: yellowish brown. VESTITURE. Dorsum setose with pale, moderately long to elongate, semierect to decumbent, simple setae. Antennae: AI densely setose with moderately long to elongate, dark, bristlelike, decumbent setae, intermixed sparsely with pale, semierect to decumbent, simple setae, then with a few elongate, dark, semierect, bristlelike setae; AII, AIII and AIV densely setose with short, decumbent, pale setae, intermixed sparsely with short, semierect, pale setae. Pleura and

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sterna: almost bare with a few long, simple, pale setae. Legs: femora densely setose with pale, moderately long to elongate, semierect, simple setae, intermixed sparsely with pale, elongate, semierect, simple setae; tibiae densely setose with short, semierect to nearly adpressed, pale setae, intermixed sparsely with elongate, semierect, spinelike, pale setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with short, nearly adpressed, pale setae, intermixed with a few long, semierect, pale setae. Abdomen: densely setose with elongate, semierect, simple, pale setae. STRUCTURE. Head: subquadrangular; frons convex in dorsal view, strongly projected anteromedially; vertex flat, midline deeply sulcate, with slightly impressed line posteriorly, posterior margin slightly carinate; frontoclypeal boundary deeply dissected; clypeus barely visible in dorsal view, margin slightly convex basally in lateral view, then straight to apex; mandibular plate moderately sized, barely visible in dorsal view, margin weakly convex; maxillary plate, moderately sized, somewhat ovoid, greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: reaching apex of mesocoxae, labial formula LII>LI>LIV>LIII;

LI longer than bucculae, reaching posterior part of proxyphus. Eyes: large, reaching dorsal margin of head in lateral view, reaching bucculae ventrally, exceeding anterolateral angles of pronotum, about nearly ½ eyes width, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, reaching posterior part of mandibular plate; AI moderately long, enlarged subbasally, AI 1.6-1.7x longer than

IOD, shorter than head width; AII cylindrical, ca. 3.7x longer than AI, ca. 1.8x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform; 2.3-2.4x wider than long, 1.4-1.6x wider than head; anterior margin slightly raised, rounded, weakly concave medially; posterior margin weakly bisinuate, lateral margin somewhat straight; callar region depressed medially, calli slightly raised,

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with impressed line; disc flat. Proepimera: flat. Mesoscutum: greatly exposed, ca. ½ scutellum length, posterior areas depressed medially, weakly convex in lateral view.

Scutellum: disc weakly raised, width ca. 1.2x length. Hemelytra: MFL moderately long, weakly convex, reaching beyond apex of scutellum, less than 1/3rd of claval commissure; costal margin slightly sinuate medially, at about level apex of clavus; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing clavus, reaching about anterior areas of cuneus. Male genitalia: genital capsule subconical, with moderately concave dorsal margin, ventral margin strongly convex, apical margin slightly sinuous, with right ventral process, armed (Fig. 3.11a); right paramere with dorsomedial process, armed (Fig. 3.11b, c); left paramere Y-shaped, sensory lobe thumblike, moderately broad, apex rounded, unarmed, with rounded basal process, small, apophysis parallel to sensory lobe, subequal in length to sensory lobe, strongly hooked subapically, margin armed with small teeth, apex rounded with pointed distal tip (Fig. 3.11d); phallotheca opening dextrodorsal, dorsal and ventral surfaces without lobe (Fig. 3.11e, f); endosoma with two spicules, elongate, slender, tapering to apex, recurved, heavily sclerotized,

DES shorter than VES; wrapped with cuplike membrane process (Fig. 3.11g).

Female: Total length 3.99-4.31 mm. COLOURATION. As in male (Fig. 3.2).

VESTITURE. As in male. STRUCTURE. Mostly as in male, body more robust. Eyes: reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Solomon Islands.

Remarks. Pseudoloxops sikaiana is most similar to P. vitiensis Kirkaldy in colouration, but can be distinguished by the following genitalic characters: the right paramere with a

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dorsal medial process and broad apophysis, which is narrowed apically in P. vitiensis; the simple phallotheca, which is without a complex process; and the presence of two, elongate, heavily sclerotized spicules, in comparison to P. vitiensis which has only one spicule.

Pseudoloxops solomonensis new species

Figures 3.2, 3.12, 3.18

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: SOLOMON ISLANDS: Malaita: Kwalo, 600-

750 m, 8.97615°S 160.9832°E, 29 Sep 1957, J. L. Gressitt, Light Trap (USI 00042910)

(BPBM). PARATYPES: SOLOMON ISLANDS: Guadalcanal: Nr. Tatuve, Kolosulu,

19 May 1960, C. W. O'Brien, Light Trap, ♂ (USI 00042909) (BPBM). Suta, 500-1200 m, 9.68333°S 160.13333°E, 27 Jun 1956, J. L. Gressitt, Light Trap, ♀ (USI 00042914)

(BPBM). San Cristoval: Bweinaniawarikiapu, 10.574°S 161.863°E, 12 Aug 1960, C.

W. O'Brien, Light Trap, 3♂ (USI 00042906 - USI 00042908), 3♀(USI 00042911 - USI

00042913) (BPBM).

Diagnosis. Pseudoloxops solomonensis is recognized by the following combination of characters: head mostly brown, frons and vertex tinged red laterally; maxillary plate reddish brown; pronotum mostly brown with reddish-brown laterally; pronotum mostly brown with red markings on lateral and medial part posteriorly; apex of scutellum broadly red with midline red; clavus mostly brown with red basally and apically; corium mostly red basally extending to half of claval commissure, broadly pale yellow brown posteriorly; apex and mediobasal angles of cuneus red; genital capsule with moderately concave dorsal margin, ventral margin widely convex, with elongate ventral

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process, with granular processes; right paramere columnar, without medial process; phallotheca simple, narrowed apically, broad basally; and, endosoma with two spicules, not serrated.

Description. Male: Body elongate-ovate, total length 3.77-3.91 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: generally brown; frons and vertex mostly brown, tinged red laterally; clypeus mostly brown, middle part with red horizontal stripe, about level anterior margin of maxillary plate; mandibular plate mostly yellowish-brown, middle part with narrow red horizontal stripe,; maxillary plate reddish-brown; lateral aspect of antennal insertion tinged red. Antennae: AI mostly reddish-yellow brown; AII mostly pale yellow brown, base tinged red; AIII and AIV uniformly pale yellow brown.

Labium: mostly yellowish-brown, apex of segment IV dark brown. Pronotum: mostly brown with lateral part reddish-brown, reaching calli, extending to dorsal margin of propleura; anterior margin medially to midline of calli brownish; disc brownish.

Propleura: proepimera mostly yellowish-brown, dorsal margin with reddish-brown band; proepisterna mostly reddish-brown, posterior margin yellowish brown.

Mesoscutum: mostly brown, with red markings lateral and medial parts posteriorly.

Scutellum: mostly brown anteriorly, ca. 1/3rd of length, mostly red posteriorly, midline red extending to apex. Hemelytra: clavus mostly brown extending to beyond apex of scutellum, intermixed with red markings basally, reaching about midpoint of scutellum, then mostly red from outer areas adjacent to apex of scutellum, reaching ca. 2/3rd of claval commissure, pale yellow brown towards apex; corium and embolium mostly red basally, reaching ca. ½ of claval commissure, then broadly pale yellowish-brown posteriorly; cuneus mostly pale yellow brown, with posterior part adjacent to membrane orange-red, about ½ of cuneus, apex red, about 1/5th of cuneus, mediobasal angles red; membrane smoky brown, veins mostly red with anterior part adjacent to corium pale

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yellow brown. Abdomen: yellowish brown, genital capsule opening dorsally with red markings. Legs: yellowish brown. VESTITURE. Dorsum densely setose with pale to darkish, iridescent, moderately long to elongate, semierect to decumbent, simple setae, intermixed with elongate, semierect, simple, pale setae, mostly on head, pronotum and hemelytra. Antennae: AI densely setose with iridescent, darkish, moderately long to elongate, semierect to decumbent, bristlelike setae, intermixed sparsely with elongate, semierect, bristlelike setae; AII densely setose with short, decumbent to adpressed, pale setae, intermixed sparsely with short, semierect, pale setae, basal part with moderately long, decumbent, pale setae, intermixed with a few long, erect, pale setae; AIII densely setose with short, decumbent to adpressed, pale setae, intermixed with a few short, semierect, pale setae; AIV densely setose with short, semierect to nearly adpressed, pale setae, intermixed with a few moderately long, semierect, pale setae. Pleura and sterna: almost bare with a few elongate, simple, pale setae. Legs: femora densely setose with short to long, semierect, simple, pale setae, intermixed with a few long, somewhat erect, simple, pale setae; tibiae densely setose with short, decumbent to nearly adpressed, pale setae, intermixed sparsely with elongate, semierect, spinelike, pale setae, alternately; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with short, nearly adpressed, pale setae, intermixed with a few long, semierect, pale setae. Abdomen: setose with short, semierect, simple, pale setae, intermixed with elongate, semierect, simple, pale setae; pygophore ventral apically with short setae, blunt tip. STRUCTURE. Head: subquadrangular, frons convex in dorsal view, narrowly, somewhat rounded, strongly projected anteromedially; vertex flat, midline deeply sulcate, with weakly impressed line posteriorly, posterior margin weakly carinate, somewhat rounded; frontoclypeal boundary deeply dissected; clypeus barely visible in dorsal view, margin slightly convex basally in lateral view, then straight to

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apex; mandibular plate moderately sized, barely visible in dorsal view, margin weakly convex; maxillary plate moderately sized, somewhat ovoid, greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: reaching apex of metacoxae; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching posterior part of proxyphus. Eyes: large, reaching dorsal margin of head in lateral view, reaching bucculae ventrally, exceeding anterolateral angles of pronotum, ca. ½ eyes width, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, reaching posterior part of mandibular plate; AI moderately long, enlarged subbasally, AI 1.5-1.8x longer than IOD, shorter than head width; AII cylindrical, 3.3-3.6x longer than AI, 1.5-1.9x longer than pronotum width; AIII and

AIV slender. Pronotum: trapeziform; 2.2-2.8x wider than long, 1.3-1.4x wider than head; anterior margin slightly raised, rounded, weakly concave medially; posterior margin slightly bisinuate; lateral margin somewhat straight; callar region depressed medially, calli slightly raised, with impressed line; disc almost flat. Proepimera: flat

Mesoscutum: exposed, ca. ½ scutellum length, posterior areas depressed medially, weakly convex in lateral view. Scutellum: disc slightly raised, width 1.1-1.2x length.

Hemelytra: MFL moderately long, weakly convex, reaching beyond apex of scutellum, less than 1/3rd of claval commissure; costal margin weakly sinuate basally; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing corium, reaching about mid-cuneus. Male genitalia: genital capsule with moderately concave dorsal margin, ventral margin projected, widely convex, with elongate ventral process, armed with short, spinelike processes (Fig. 3.12a); right paramere hooked-shaped, apophysis broad, apically rounded, apex to subbasal dorsal margin serrated (Fig. 3.12b); left paramere Y-shaped, sensory lobe thumblike, apex rounded, basal process not prominent, apophysis elongate,

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slim, ca. 45° to sensory lobe, longer in length to sensory lobe, sinuate margin, subapically slightly enlarged, strongly hooked, apex acute (Fig. 3.12c); phallotheca nearly ovoid, apex narrow, opening apical to dextrodorsal region, without process (Fig.

3.12d, e); endosoma with two spicules, DES elongate, heavily sclerotized, simple, smooth margin, somewhat straight then recurved apically, tapering to apex, membranous basally, VES short, slim, lightly sclerotized, simple, ventral to DES, position of spicules dorsal, close to secondary gonopore, wrapped with cuplike membrane process (Fig. 3.12f).

Female: Total length 3.98-4.25 mm. Colouration and vestiture as in male (Fig. 3.2).

STRUCTURE: Mostly as in male, body more robust. Eyes: slightly smaller, not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Solomon Island.

Remarks. Pseudoloxops solomonensis is most closely related to P. baiawaensis from

Papua New Guinea and Sabah, Malaysia, on the basis of male genitalic structures. The distinguishing characters for both species are presented in the ‘Remarks’ section of P. baiawaensis.

Pseudoloxops straatmani new species

Figures 3.2, 3.13, 3.18

Etymology. This species is named in honour of R. Straatman, who is one of the collectors of this species.

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Type material. HOLOTYPE: Male: SOLOMON ISLANDS: Santa Ysabel: Buala,

8.16624°S 159.61898°E, 20 Aug 1964, R. Straatman, Light Trap (USI 00042916)

(BPBM). PARATYPES: SOLOMON ISLANDS: Malaita: Kwalo, 600-750 m,

8.97615°S 160.9832°E, 29 Sep 1957, J. L. Gressitt, Light Trap, ♀ (USI 00042915)

(BPBM). Santa Ysabel: Tatamba, 8.39361°S 159.81°E, 24 Jun 1960, C. W. O'Brien,

Light Trap, ♂ (USI 00042917) (BPBM).

Diagnosis. Pseudoloxops straatmani is recognized by the following combination of characters: frons and vertex generally yellowish-brown with red markings on medial and lateral part; maxillary plate uniformly red; pronotum broadly red laterally with yellowish-brown tinge, medial part mostly yellowish-brown, anterior margin and midline of callar region red, disc with broad red markings medially; mesoscutum with broad red spots laterally; scutellum broadly red medially, midline paler; cuneus mostly red with pale yellow brown basally and apically; genital capsule with ventral process, armed with short spinelike process; right-paramere reverse L-shaped; the Y-shaped left paramere, apophysis perpendicular to sensory lobe, apex hooked; phallotheca simple, nearly cylindrical; and, endosoma with four spicules, not serrated.

Description. Male: Body elongate-ovate, total length 4.90 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: frons yellowish brown with red midline, extending from anterior part to vertex; vertex with red markings on midline and laterally, mostly yellowish-brown posteriorly; clypeus mostly yellowish-brown, with red horizontal stripe, about level anterior margin of maxillary plate, dorsum tinged red; mandibular plate mostly yellowish-brown, with narrow red band medially, latitudinal; maxillary plate uniformly red; lateral aspect of antennal insertion tinged red. Antennae: AI mostly red with yellowish-brown tinge; AII and AIII pale yellow; missing AIV. Labium: mostly yellowish-brown, apex of segment IV dark brown. Pronotum: lateral areas

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broadly red with yellowish-brown tinge, reaching lateral part of calli, extending to dorsal margin of propleura, mostly yellowish-brown medially, anterior margin to midline of callar region red; disc mostly yellowish-brown with broad red spot medially, tinged yellowish-brown on midline. Propleura: proepimera mostly yellowish-brown, dorsal areas with red horizontal markings, margins adjacent to pronotum yellowish- brown; proepisterna mostly red, posterior margin yellowish-brown. Mesoscutum: mostly pale yellow brown, with broad red spots laterally, not reaching lateral margin, sometimes tinged red medially. Scutellum: mostly pale yellow brown, broadly red medially, extending to apex with ca. 1/3rd length red, midline paler. Hemelytra: clavus mostly smoky brown basally, with red markings at medial areas, reaching ca. 2/3rd of scutellum length, then broadly yellowish to pale yellow brown, reaching ca. ½ length of claval commissure, red to apex; endocorium mostly red basally reaching ca. ½ length of claval commissure, mostly yellowish to pale yellow brown posteriorly, with broad red markings adjacent to apex of clavus, contiguous with anterior margin of membrane; exocorium mostly red basally, almost reaching level 2/3rd length of claval commissure, mostly yellowish to pale yellow brown posteriorly; cuneus mostly red, with anterior areas adjacent to embolium and apex yellow to pale yellow brown; membrane pale, tinged pale brown, veins mostly red. Legs: mostly yellowish-brown; hind femora with broad red band on medial part. Abdomen: mostly yellowish-brown, genital capsule opening dorsally with red markings. VESTITURE. Dorsum setose with pale, moderately long to elongate, semierect to decumbent, simple setae, intermixed with elongate, semierect, simple, pale setae. Antennae: AI densely setose with iridescent, moderately long to elongate, semierect to decumbent, bristlelike setae, short on apex, intermixed sparsely with iridescent, elongate, semierect, bristlelike setae; AII and AIII densely setose with short, decumbent to adpressed, pale setae, intermixed with a few

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short, semierect, pale setae; missing AIV. Pleura and sterna: almost bare with a few long, simple, pale setae. Legs: femora densely setose with short to elongate, semierect, simple, pale setae, intermixed sparsely with pale, elongate, somewhat erect, simple setae; tibiae densely setose with short, semierect to nearly adpressed, pale setae, intermixed sparsely with elongate, semierect, spinelike, pale setae, alternately; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with short, nearly adpressed, pale setae, intermixed with a few long, semierect, pale setae.

Abdomen: densely setose with elongate, semierect, simple, pale setae. STRUCTURE.

Head: subquadrangular, frons slightly convex in dorsal view, slightly projected medially, almost flat; vertex almost flat, midline deeply sulcate, with slightly impressed line posteriorly, posterior margin carinate; frontoclypeal boundary weakly dissected; clypeus visible in dorsal view, enlarged, margin slightly convex basally in lateral view, then straight to apex; mandibular plate moderately sized, slightly visible in dorsal view, margin weakly convex; maxillary plate moderately sized, somewhat ovoid, greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head.

Labium: surpassing apex of mesocoxae, nearly reaching metacoxae; labial formula

LII>LI>LIV>LIII; LI longer than bucculae, reaching posterior part of proxyphus. Eyes: large, somewhat rounded, reaching dorsal margin of head in lateral view, reaching bucculae ventrally, exceeding anterolateral angles of pronotum, ca ½ eyes width, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, reaching posterior areas of mandibular plate; AI long, slightly enlarged subbasally, AI

1.8-2.0x longer than IOD, shorter than head width; AII cylindrical, 3.2-3.3x longer than

AI, 1.7-2.0x longer than pronotum width; AIII slender; missing AIV. Pronotum: trapeziform; 2.3-2.5x wider than long, 1.2-1.3x as wide as head; anterior margin slightly raised, rounded, with impressed line, anterior margin slightly concave medially; lateral

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margin slightly sinuate; posterior margin slightly excavate; callar region depressed medially, calli slightly raised, with impressed line; disc almost flat. Proepimera: weakly inflexed. Mesoscutum: exposed, ca. ½ scutellum length, slightly longer, weakly convex in lateral view. Scutellum: disc slightly raised, width 1.2-1.3x length. Hemelytra: costal margin slightly sinuate basally; MFL moderately long, weakly convex, reaching ca.

1/3rd of claval commissure; cuneus moderately long, longer than wide, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing corium, reaching about anterior part of cuneus. Legs: hind femora enlarged, ca. 2x middle femora. Male genitalia: genital capsule with moderately concave dorsal margin, ventral margin convex, moderately sized, with ventral process, armed with short, spinelike processes

(Fig. 3.13a); right paramere reverse L shaped, apophysis elongate, margin denticulate, apex rounded (Fig. 3.13b); left paramere Y-shaped, apophysis thumblike, moderately projected, shorter in length to apophysis, apex rounded, without basal process, apophysis elongate, perpendicular to sensory lobe, broad, subrectangular, subapically strongly hooked, apex acute (Fig. 3.13c); phallotheca nearly cylindrical, without process, apically rounded, opening apical to dextrodorsal region (Figs. 3.13d, e); endosoma with four spicules, moderately sclerotized: VES elongate, leaf-shaped, somewhat broad, tapering to apex, margin smooth; DES1 moderately long, slender, tapering to apex, margin smooth, apex acute; DES2 short, fingerlike, medially membranous then lightly sclerotized to base; DES3 short, spinelike, broader than DES2, membranous basally, VES close to secondary gonopore, DES1, DES2 and DES3 distal to secondary gonopore (Fig. 3.13f).

Female: Total length 4.37 mm. COLOURATION. Mostly as in male but head, pronotum, mesoscutum and scutellum ground colour brownish with red markings, with small, dark brown spots on pronotum, mesoscutum and scutellum. VESTITURE. As in

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male. STRUCTURE: Mostly as in male, body more robust. Eyes: slightly smaller, reaching dorsal margin of head in lateral view, not reaching bucculae ventrally.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Solomon Island.

Remarks. Pseudoloxops straatmani can be differentiated from other Southwest Pacific species by having the reverse L-shaped right paramere and the Y-shaped left paramere, with the apophysis right-angle to the sensory lobe. This species is also distinguished from other species by having the disc of pronotum with broad red spot medially with a yellowish tinge at the midline, and the presence of broad red spots on lateral regions of the mesoscutum, but not reaching the lateral margins. P. straatmani shares the presence of a ventral process on the pygophore with P. baiawaensis, P. sikaiana, P. solomonensis and P. vitiensis. This species is allied to P. tatarnici by having a nearly cylindrical phallotheca, without a process, and the aedeagus has four smooth endosomal spicules.

However, the VES is elongate and unbranched in P. straatmani, whereas in P. tatarnici, the VES is branched. Apart from that, DES2 in P. straatmani is membranous medially and sclerotized at the base, and DES3 is membranous basally.

Pseudoloxops tabwemasana new species

Figures 3.2, 3.14, 3.19

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Male: VANUATU: Sanma: Espiritu Santo Is. Co.:

Tabwemasana Ridge, 15.36809°S 166.74876°E, 1372 m, 02 Sep 1971, G.S. Robinson,

(USI 00038840) (SAMA). PARATYPES: VANUATU: Sanma: Espiritu Santo Is.

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Co.: Tabwemasana Ridge, 15.36809°S 166.74876°E, 1372 m, 02 Sep 1971, G.S.

Robinson, 11♂ (USI 00038841 - USI 00038851), 7♀ (USI 00038852 - USI 00038858),

1 unknown (USI 00038859) (SAMA).

Diagnosis. Pseudoloxops tabwemasana is recognized by the following combination of characters: body elongate; mostly pale yellow brown with red markings on apex of clavus, weakly on medial areas of endocorium adjacent to apex of clavus, with olive markings on embolium to costal margin; genital capsule large, right paramere with ventral medial process, perpendicular to apophysis; left paramere with broad basal process; dorsal surface of phallotheca with two small lobes; and, endosoma with moderately broad single spicule, sclerotized apically and basally, membranous medially.

Description. Male: Body elongate, subparallel, total length 4.28-4.77 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: mostly pale yellow brown, with maxillary plate red anteriorly, pale yellow brown posteriorly; outer antennal insertion tinged red. Antennae:

AI mostly red, medial part paler with pale yellow brown stripe; AII pale yellow brown, reddish at base and apex; AIII pale yellow brown, apex red; AIV pale yellow brown, apex brownish. Labium: mostly pale yellow brown, dark brown apically. Pronotum: uniformly pale yellow brown. Propleura: proepimera mostly pale yellow brown, dorsal margin sometimes tinged red; proepisterna mostly pale yellow brown, anterior part sometimes with red tinge. Mesoscutum: uniformly pale yellow brown. Scutellum: uniformly pale yellow brown. Hemelytra: clavus mostly pale yellow brown with red markings at apical part, ca. 1/3rd of clavus commissure length; corium mostly pale yellow brown, with minor red markings at medial areas adjacent to apical part of clavus; embolium to costal margin with olive markings; cuneus mostly pale yellow brown, lateral margin tinged olive; membrane smoky brown, veins mostly red with

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anterior part pale yellow brown. Abdomen: uniformly pale yellow brown. Legs: uniformly pale yellow brown. VESTITURE. Dorsum densely setose with pale, shiny, elongate, semierect, simple setae, intermixed with pale, shiny, moderately long, decumbent, simple setae. Antennae: AI densely setose with pale, shiny, short to moderately long, decumbent setae, intermixed sparsely with pale, elongate, semierect, bristlelike setae; AII and AIII densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short to moderately long, semierect, pale setae; AIV densely setose with pale, short, nearly adpressed setae, intermixed sparsely with semierect, pale setae. Pleura and sterna: almost bare with a few scattered, pale, elongate, simple setae. Abdomen: venter with scattered, pale, elongate semierect setae, denser laterally. Legs: femora densely setose with pale, short to long, semierect setae, intermixed with pale, elongate, semierect setae; tibiae densely setose with pale, short, nearly adpressed setae, intermixed sparsely with smoky brown, elongate, semierect, spinelike setae, alternately, apex with a few short, smoky brown, spinelike setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, semierect to nearly adpressed setae, intermixed with a few elongate, semierect, pale setae. STRUCTURE. Head: subquadrangular; frons slightly convex in dorsal view, slightly projected in front anteriorly, almost flat; vertex almost flat in lateral view, midline deeply sulcate, with impressed line posteriorly, posterior margin slightly carinate; clypeus visible in dorsal view, large, subrectangular; mandibular plate moderately sized, visible in dorsal view, margin slightly convex; maxillary plate moderately sized, ovoid, greatly tumescent, exceeding mandibular plate; bucculae reaching posterior margin of head. Labium: reaching apex of metacoxae, labial formula

LII>LI>LIII>LIV; LI longer than bucculae, reaching posterior margin of proxyphus.

Eyes: moderately sized, nearly reaching dorsal margin of head in lateral view, nearly

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reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plate, reaching interior margin of eyes; AI long, slightly enlarged subbasally; AI 1.5-1.6x longer than IOD, shorter than head width; AII cylindrical, ca. 3.5-3.8x longer than AI, 1.5-1.6x longer than pronotum; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform; 2.5-2.7x wider than long, 1.5x wider than head; anterior margin slightly raised, rounded, slightly concave medially; posterior margin slightly bisinuate, lateral margin straight; callar region depressed medially, calli slightly raised, with impressed line; disc almost flat. Proepimera: flat.

Mesoscutum: exposed, ca. ½ of scutellum length. Scutellum: disc raised, width 1.2-1.3x length. Hemelytra: MFL weakly convex, reaching ca. 1/3rd of claval commissure; cuneus narrow and long, ca. 2x width; membrane cells long. Abdomen: surpassing clavus, reaching about posterior part of corium. Male genitalia: genital capsule large, with moderately concave dorsal margin, with medial lobe, ventral margin projecting, apically rounded, with sinuous margin (Fig. 3.14a); right paramere with ventral medial process, perpendicular to apophysis, apically armed with small teeth, apex rounded, apophysis bulbous, armed with sclerotized small teeth (Fig. 3.14b); left paramere Y- shaped, sensory lobe moderately broad, apex rounded, unarmed, with basal process, enlarged, rounded, apophysis elongate, slightly hooked apically (Fig. 3.14c); phallotheca opening apically, ventral surface with subapical fold, moderately sized, subtriangular, dorsal surface with two small lobes, convex, with subapical dextrodorsal lobe, moderately sized, sinuous margin (Figs. 3.14d, e); endosoma with single spicule, moderately long, smooth margin, heavily sclerotized, membranous medially, then slightly sclerotized basally, moderately sinuate subapically, apex rounded, position close to secondary gonopore, covered with membrane bag at base (Fig. 3.14f).

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Female. Total length 4.61-4.96 mm. Colouration and vestiture as in male (Fig. 3.2).

STRUCTURE: Mostly as in male. Eyes: slightly smaller, not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI long, 1.4-1.6x longer than IOD; AII 2.9-3.2x longer than AI, 1.4-1.5x longer than pronotum width.

Pronotum: 2.4-2.6x as wide as long, 1.6-1.7x as wide as head. Scutellum: width 1.1-

1.2x length.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Vanuatu.

Remarks. Pseudoloxops tabwemasana is most similar in colouration to P. efatensis and

P. lenakelensis by having a mostly pale yellow brown body and red apex of the clavus.

However, P. tabwemasana differs by the following characters: the large size; the dorsal margin of the genital capsule with a medial lobe, the apex of the ventral margin rounded, margin sinuate; the right paramere with a ventral medial process, armed, perpendicular to apophysis; the left paramere with broad basal process; the phallotheca opening apically, with subtriangular subapical fold ventrally, dorsal surface with two small lobes; and, the endosoma with a single spicule, which is moderately long, with a smooth margin, that is heavily sclerotized, membranous medially, and slightly sclerotized basally and moderately sinuate subapically.

Pseudoloxops tatarnici new species

Fig. 3.2, 3.15, 3.19

Etymology. This species is named in honour of Dr. Nikolai Tatarnic, who is one of the collectors of this species.

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Type material. HOLOTYPE: Male: AUSTRALIA: Northern Territory: 185.6 km N of Barkely Homestead on Tablelands Highway, 18.18795°S 135.73106°E, 239 m, 20

Apr 2009, Cassis, Tatarnic, Monteith, (USI 00017703) (UNSW). PARATYPES:

AUSTRALIA: Northern Territory: 185.6 km N of Barkely Homestead on Tablelands

Highway, 18.18795°S 135.73106°E, 239 m, 20 Apr 2009, Cassis, Tatarnic, Monteith,

1♂ (USI 00017702), 3♀ (USI 00017699 - USI 00017701) (UNSW).

Diagnosis. Pseudoloxops tatarnici is recognized by the following combination of characters: head dorsum mostly yellow, frons with red markings anteriorly; base and apex of AII reddish; apex of AIII reddish; pronotum red laterally, yellowish medially, disc broadly greenish-yellow; mesoscutum uniformly yellow; scutellum mostly greenish-yellow with pale red apex; hemelytra greenish-yellow with red markings on clavus at basal and apical part, endocorium mostly, apex of embolium, apex and mediobasal angles of cuneus, and veins mostly; pygophore with right and left dorsal process, elongate, armed; left paramere with elongate sensory lobe, rounded apically, apophysis slim; phallotheca simple, without process; and, endosoma with four spicules, not serrated.

Description. Male: Body elongate-ovate, body length 3.10-3.57 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: mostly yellow, frons with red markings anteriorly; clypeus mostly yellow with red tinged at dorsum, extending to midline to about 2/3rd length, then transverse red, apical part yellow, lateral areas anteriorly tinged red; mandibular plate mostly yellow with narrow red horizontal markings medially; maxillary plate mostly red; outer antennal insertion red. Antennae: AI mostly red, medial part paler with pale yellow brown stripe; AII pale yellow brown, reddish at base and apex; AIII pale yellow brown, apex reddish; AIV missing. Eyes: reddish brown.

Labium: mostly pale yellow, LIV mostly dark brown. Pronotum: lateral areas broadly

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sanguineous, reaching lateral part of calli, not extending to dorsal margin of propleura, margin yellow; callar region yellow medially; disc broadly greenish-yellow. Propleura: proepimera mostly pale yellow, dorsal areas with red-brown markings; proepisterna anteriorly red, posterior areas pale yellow. Mesoscutum: uniformly yellow. Scutellum: mostly greenish-yellow, apex pale red. Hemelytra: clavus mostly greenish-yellow, sanguineous at basal part, reaching about ½ of mesoscutum length, along outer areas between claval suture and claval vein, extending to apex; corium mostly sanguineous at endocorium; mostly greenish-pale yellow at exocorium and posterior areas; apex of embolium red; cuneus mostly greenish-pale yellow, mediobasal angles and apex red; membrane smoky pale brown, veins red. Sterna: mostly yellow. Abdomen: mostly greenish-yellow. Legs: mostly pale yellow brown; apex of hind femora laterally with dark brown markings; apex of tarsi blackish. VESTITURE. Dorsum densely setose with pale, silky, moderately long to elongate, decumbent to semierect, simple setae, intermixed with long, dark, decumbent, bristlelike setae mostly on corium posteriorly and cuneus. Antennae: AI densely setose with pale, silky, short to moderately long, decumbent, simple setae, intermixed sparsely with pale, silky, elongate, erect setae; AII and AIII densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short, semierect, pale setae. Pleura and sterna: almost bare with a few scattered, pale, silky, simple setae. Abdomen: venter densely setose with pale, silky, short, simple setae, intermixed with long, pale setae, mostly on lateral. Legs: femora densely setose with pale, short to long, semierect setae, intermixed with pale, elongate, semierect setae; tibiae densely setose with pale, short, nearly adpressed setae, intermixed sparsely with dark, long, spinelike, semierect setae, alternately; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, nearly adpressed setae, intermixed with a few elongate, semierect setae.

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STRUCTURE. Head: frons slightly convex in dorsal view, slightly projected anteriorly; vertex flat in lateral view, midline moderately sulcate, posterior margin rounded; posterior margin rectilinear; clypeus visible in dorsal view, large, convex basally; mandibular plate moderately sized, visible in dorsal view, lateral margin weakly convex; maxillary plate moderately sized, tumescent, moderately, mostly medial, slightly visible in dorsal view, slightly exceeding mandibular plate; bucculae small, reaching posterior margin of head. Labium: reaching mesocoxae; labial formula

LII>LI>LIV>LIII; LI longer than bucculae, reaching about proxyphus medially. Eyes: moderately sized; reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally; exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plate, reaching interior margin of eyes; slightly enlarged subbasally, moderately long, AI equal to IOD; AII cylindrical, ca. 3.5x longer than AI;

AIII nearly cylindrical, ca. 1.3x longer than pronotum width; AII>AIII>AI, missing

AIV. Pronotum: trapeziform; ca. 2.3x as wide as long, ca. 1.3x as wide as head; anterior margin rounded, slightly raised, weakly concave medially; posterior margin slightly excavate, with weakly bisinuate margin; lateral margin straight; calli weakly raised, with impressed line; disc flat. Proepimera: slightly inflexed. Mesoscutum: exposed, ca.

1/3rd of scutellum length. Scutellum: disc almost flat; ca. 1.4x as wide as long.

Hemelytra: MFL weakly convex, reaching ca. 1/3rd of claval commissure; cuneus moderately long, reaching about midpoint of membrane, longer than wide (ca. 1.5x width), not greatly narrowed; membrane cells moderately long. Abdomen: surpassing clavus, reaching anterior margin of cuneus. Male genitalia: pygophore with moderately concave dorsal margin; ventral margin projected, moderately convex, margin slightly sinuate, with left and right processes, elongate, armed (Figs. 3.15a, b); right paramere

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weakly C-shaped, with dorsal medial process, apophysis elongate, apex bulbous, armed with a few small teeth, medial process short, armed apically (Fig. 3.15c); left paramere deeply Y-shaped, sensory lobe elongate, fingerlike, unarmed, apex rounded, without basal process, apophysis elongate, slim, parallel to sensory lobe, shorter in length to sensory lobe, subapically hooked (Fig. 3.15d); phallotheca simple, without process (Fig.

3.15e); endosoma with four spicules, DES1, DES2 and DES3 not serrated, margins smooth, VES branched (Fig 3.15f).

Female: Mostly as in male, total length 3.44-3.64 mm (Fig. 3.2). Head: frons broad in dorsal view, rounded; vertex almost flat. Eyes: not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI moderately long, sometimes longer than IOD, 1.0-1.2x longer than IOD; AII cylindrical, 3.1-3.2x longer than AI,

1.2-1.3x longer than pronotum width; AIV slender, pale yellow brown, apex dark brown, densely setose with pale, short, nearly adpressed setae, intermixed sparsely with pale, semierect setae; AII>AIII>AIV>AI. Pronotum: 2.6-2.7x as wide as long, 1.3-1.4x as wide as head. Scutellum: 1.2-1.5x as wide as long.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. P. tatarnici is known only from the type locality, Barkely Homestead on

Tablelands Highway, Northern Territory.

Remarks. P. tatarnici is mostly similar to P. woodwardi, but can be distinguished by the following characters: frons with red markings anteriorly; mandibular plate mostly yellow with narrow red markings on middle part; disc of pronotum broadly greenish- yellow; mesoscutum uniformly yellow; scutellum mostly greenish-yellow, apex pale red; clavus mostly greenish-yellow, with red markings along outer areas between claval suture and vein, extending to apex; corium mostly red on endocorium; pygophore with

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ventral margin projected, margin slightly sinuate, dorsal margin without lobe; with dorsal processes; right paramere with bulbous apex of apophysis; left paramere with rounded sensory lobe, without basal process; endosoma with four spicules, not serrated.

Pseudoloxops vitiensis Kirkaldy 1908

Figures 3.2, 3.16, 3.19

Tichorhinus vitiensis Kirkaldy 1908: 377 (n. sp.).

Pseudoloxops vitiensis: Carvalho 1955b: 224 (new combination); Carvalho 1958: 128

(catalogue); Carvalho and Wallerstein 1976: 690 (descr., disc., DV., MG);

Schuh 1995: 186 (catalogue).

Materials examined. FIJI: Ba: Tavua, Viti Levu, 17.44°S 177.87°E, 07 Nov 1964, G.

F. Gross, 1♀ (USI 00038833) (SAMA). Lau Group: Vatoa, Vatoa, 19.83333°S

178.23333°E, 20 m, 23 Feb 1971, N. L. H. Krauss, 1♀ (USI 00042706) (BPBM).

Moala: Matuku I., 19.1667°S 179.7667°E, 07 Apr 1924, E. H. Bryan, Jr., 3♀ (USI

00042678 – USI 00042679, USI 00042707) (BPBM). Rewa: Rewa [Province, no specific location], 18.08747°S 178.33205°E, 259 m, Mar 1906, Muir, ♀ (USI

00042769) (BPBM), 1♂ (USI 00042770) (BPBM). Vanua Levu: Savusavu,

16.77786°S 179.33976°E, 100 m, Mar 1978, N. L. H. Krauss, 2♂ (USI 00042700, USI

00042701), 2♀ (USI 00042703, USI 00042704) (BPBM). Viti Levu: 70 km E of Nadi,

17.79338°S 178.0781°E, 26 Jul 1967, J. & M. Sedlacek, 1♂ (USI 00042702) (BPBM).

Base of Mt. Evans, nr. Vuda Point, 17.66666°S 177.58333°E, 18 Mar 1978, E. I.

Schlinger, 1♀ (USI 00193437) (CAS). Raki Raki, 17.36666°S 178.15°E, Jan 1955, N.

L. H. Krauss, 1♀ (USI 00042705) (BPBM). Rdg. W of Nandarivatu, 17.61265°S

177.94588°E, 853 m, 11 Sep 1938, E. C. Zimmerman, 1♀ (USI 00042894) (BPBM).

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Diagnosis. Pseudoloxops vitiensis is recognized by the following combination of characters: head generally pale yellow brown, frons with a narrow red stripes extending from midline to lateral margin; maxillary plate entirely sanguineous; AII and AIII pale yellow brown, apex tinged red; anterior and lateral regions broadly red, disc broadly pale yellow brown; mesoscutum uniformly pale yellow brown; half apical part of scutellum red, midline basally red, not reaching margin; clavus red on basal and apical part; corium broadly red extending to apex of clavus; apical part of cuneus broadly red, mediobasal angles red; genital capsule with ventral margin projected, somewhat rounded, with right ventral process, armed with short, spinelike processes; right paramere with dorsal medial process, armed; left paramere Y-shaped, sensory lobe broad, apex rounded, unarmed, with rounded basal process, apophysis parallel to sensory lobe, shorter in height to sensory lobe, phallotheca opening from apical to left lateral subapically, with left subapical fold ventrally, large, dorsal surface with right subapical fold, large, rounded margin; and, endosoma with single spicule, elongate, fingerlike, tapering to apex, heavily sclerotized, membranous basally.

Redescription. Male: Body elongate-ovate, total length 3.80-3.83 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: generally pale yellow brown, broadly on vertex; frons with red midline, with a few narrow red horizontal stripes from midline to lateral margin, extending to anterior margin, with anterior part medially adjacent to clypeus pale yellow brown, lateral margins red; clypeus with red markings on dorsum laterally extending to midline to about 2/3rd length, then transverse red marking, lateral margins red basally, apex pale yellow brown; mandibular plate pale yellow brown with broadly sanguineous horizontal marking; maxillary plate entirely sanguineous; lateral aspect of antennal insertion sanguineous. Antennae: antennifers red; AI mostly red, with medial part yellowish-brown; AII and AIII pale yellow brown, apex tinged red; AIV yellowish-

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brown, apex brownish. Labium: mostly pale yellow brown, apex of segment IV mostly dark brown. Pronotum: lateral part broadly red, margin tinged brown, reaching calli mostly, extending to dorsal margin of propelura, posterolateral margin tinged dark brown; anterior part red, callar region pale yellow brown medially; disc broadly pale yellow brown. Propleura: proepimera mostly pale yellow brown, dorsal areas brownish intermixed with narrow red markings; proepisterna anteriorly red, posterior areas pale yellow brown. Mesoscutum: uniformly pale yellow brown. Scutellum: anterior part mostly pale yellow brown, ca. ½ length, midline red but not reaching anterior margin, red posteriorly extending to apex. Hemelytra: clavus red basally, reaching about midpoint of scutellum, broadly pale yellow brown at medial part, then red apically, ca.

1/3rd of claval commissure; corium mostly red, reaching about apex of clavus, then pale yellow brown posteriorly; embolium mostly pale yellow brown; cuneus mostly pale yellow brown anteriorly, ca. 1/3rd of cuneus, mediobasal angles red, then broadly red to apex; membrane smoky pale brown, veins red. Abdomen: yellowish-brown. Legs: yellowish-brown; apex of tarsi brownish. VESTITURE. Dorsum setose with pale silky, moderately long to elongate, decumbent to semierect, simple setae. Antennae: AI densely setose with pale, short to moderately long, semierect to decumbent, simple setae, intermixed sparsely with dark, elongate, semierect to erect, bristlelike setae; AII,

AIII and AIV densely setose with pale, short, decumbent to adpressed setae, intermixed sparsely with short, semierect, pale setae. Pleura and sterna: almost bare with a few pale, long, silky, simple setae. Abdomen: venter setose with pale, silky, long to elongate, simple setae, denser laterally. Legs: femora densely setose with pale, short to moderately long, semierect, simple setae, intermixed sparsely with pale, elongate, semierect, simple setae; tibiae densely setose with pale, short, semierect to nearly adpressed setae, intermixed sparsely with pale, elongate, semierect, spinelike setae,

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alternately, apex with a few short, spinelike setae; hind tibiae with several longitudinal, parallel rows of tiny spinules; tarsi densely setose with pale, short, nearly adpressed setae, intermixed with a few, elongate, semierect, pale setae. STRUCTURE. Head: subquadrangular; frons strongly convex in dorsal view, moderately projected in front anteriorly, reaching about mid-dorsum of clypeus, dorsum almost flat; vertex almost flat, midline moderately to deeply sulcate, with slightly impressed line posteriorly, posterior margin slightly carinate, rectilinear; clypeus visible in dorsal view, large, subrectangular; mandibular plate moderately sized, barely visible in dorsal view, margin slightly convex; maxillary plate moderately sized, ovoid, greatly tumescent, exceeding mandibular plate; bucculae small, reaching posterior margin of head. Labium: surpassing apex of metacoxae, about reaching trochanter; labial formula

LII>LI>LIV>LIII; LI longer than bucculae, about reaching posterior part of proxyphus.

Eyes: moderately sized, oval shaped; reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally; slightly exceeding anterolateral angles of pronotum, contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plate, reaching interior margin of eyes;

AI moderately long, slightly enlarged subbasally, 1.4-1.6x longer than IOD, shorter than head width; AII cylindrical, ca. 3.1-3.5x longer than AI, 1.5-1.7x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI. Pronotum: trapeziform; 2.3-2.4x wider than long, 1.3-1.5x wider than head, weakly convex in lateral view; anterior margin slightly raised, weakly concave medially; posterior margin slightly bisinuate; lateral margin nearly straight; callar region deeply depressed medially, calli slightly raised, with impressed line; disc flat. Proepimera: flat. Mesoscutum: exposed, ca. ½ of scutellum length. Scutellum: slightly raised, width 1.2-1.4x width. Hemelytra: MFL moderately long, weakly convex, reaching ca. 1/3rd of claval commissure; cuneus

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moderately long, ca. 1.5x width, not greatly narrowed; membrane cells moderately long.

Abdomen: surpassing clavus, reaching about anterior part of cuneus. Male genitalia: genital capsule with moderately concave dorsal margin, ventral margin projected, somewhat rounded, with ventral process, armed with short, spinelike processes (Fig.

3.16a); right paramere with dorsal medial process, short and broad, armed apically, apophysis fingerlike, apex rounded, armed (Fig. 3.16b); left paramere Y-shaped, sensory lobe broad, apex rounded, unarmed, with rounded basal process, apophysis elongate, slim, parallel to sensory lobe, moderately hooked subapically, apex duck head shaped (Fig. 3.16c); phallotheca opening from apical to left lateral subapically, ventral surface with left subapical fold, enlarged, subrectangular with narrowly concave margin, dorsal surface with right subapical fold, enlarged, rounded margin (Figs. 3.16d, e); endosoma with single spicule, elongate, fingerlike, tapering to apex, with rounded apex, slightly recurved subbasally, heavily sclerotized, membranous basally, position left to dextrodorsal region, close to secondary gonopore (Fig. 3.16f).

Female: Total length 3.79-4.34 mm. Colouration and vestiture as in male (Fig. 3.2).

STRUCTURE. Mostly as in male, body robust. Eyes: slightly smaller, not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI moderately long, 1.3-1.5x longer than IOD; AII 3.1-3.3x longer than AI, 1.4-1.6x longer than pronotum width. Pronotum: 2.2-2.9x as wide as long, 1.4-1.6x as wide as head.

Scutellum: 1.3x as wide as long.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Fiji Islands.

Remarks. Pseudoloxops vitiensis Kirkaldy 1908 is most similar to P. sikaiana in colouration, but can be distinguished by the following genitalic characters: the right

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paramere with a narrow dorsal medial process and apophysis; the phallotheca with a left subapical fold ventrally and a right subapical fold dorsally; and, the endosoma with a single, elongate, heavily sclerotized spicule, not serrated, membranous basally.

Pseudoloxops wareo new species

Figures 3.2, 3.18

Etymology. This species is named after the type locality.

Type material. HOLOTYPE: Female: PAPUA NEW GUINEA: Morobe Province:

Wareo, Finschaven, 6.45°S 147.7883°E, 01 Jan 1700, Rev. L. Wagner (USI 00038834)

(SAMA).

Diagnosis. Pseudoloxops wareo is recognized by the following combination of characters: large size; head dorsum entirely brownish, without red markings; clypeus mostly pale yellow brown with reddish tinge; pronotum broadly orange-red with brownish tinge on anterior and lateral regions of pronotum, disc broadly pale yellow brown medially; mesoscutum with orange-red markings on anterolateral angles and posterior margin medially; scutellum mostly orange-red; clavus broadly brownish on basal half, tinged red; and, cuneus mostly red with mediobasal angles pale yellow brown.

Description. Female: Body elongate, subparallel, total length 5.05-5.31 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: frons and vertex entirely brownish; clypeus mostly pale yellow brown, laterally reddish, not reaching base and apex, midline tinged red basally; mandibular plate broadly red with posterior margin adjacent to clypeus paler; maxillary plate with red markings anteriorly and laterally, pale yellow brown posteriorly; lateral aspect of antennal insertion reddish. Antennae: AI mostly red;

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missing others. Labium: mostly pale yellow brown, apex of segment IV dark brown.

Pronotum: anterior and lateral regions broadly orange-red with brownish tinge, paler on anterior margin medially; disc broadly pale yellow brown medially. Propleuron: proepimeron mostly pale yellow brown with orange-red band dorsally, tinged brownish; proepisternum red anteriorly, with pale yellow brown posteriorly. Mesoscutum: mostly pale yellow brown, with orange-red markings on anterolateral angles and posterior margin medially. Scutellum: lateral part mostly red, anterior margin to middle part mostly orange-red, extending to apex, anterolateral angles yellowish brown. Hemelytra: clavus broadly yellowish-brown on basal half, extending beyond apex of scutellum, then mostly pale yellow brown, basal part with red markings reaching about level posterior margin of mesoscutum, with orange-red markings along claval commissure to apex, about 1/4th of commissure, not reaching outer part apex of scutellum; endocorium broadly red, reaching about level ¾ of claval commissure, extending to exocorium, then mostly pale yellow brown posteriorly with orange-red markings adjacent to apex of scutellum and membrane anteriorly; exocorium entirely red, distal margin of embolium pale yellow brown; cuneus almost entirely red with outer base angles adjacent to embolium pale yellow brown; membrane yellowish brown; veins mostly red. Legs: missing. Abdomen: pale yellow brown. VESTITURE. Dorsum setose with pale, moderately long to elongate, decumbent, simple setae. Antennae: AI setose with moderately long, semierect to decumbent, simple, pale setae. Pleura and sterna: almost devoid with a few long, simple, pale setae. Abdomen: densely setose with pale, elongate, semierect, simple setae. STRUCTURE. Head: subquadrangular; frons convex in dorsal view, somewhat rounded, moderately projected anteromedially, somewhat flat; vertex almost flat, midline deeply sulcate, with impressed line posteriorly, posterior margin carinate; frontoclypeal boundary slightly dissected; clypeus visible in dorsal

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view, enlarged, margin slightly convex, bicompressed laterally, about level anterior margin of maxillary plate; mandibular plate large, visible in dorsal view, margin weakly convex; maxillary plate moderately sized, greatly tumescent, exceeding mandibular plate, slightly exceeding posterior margin of mandibular plate from anterior view; bucculae reaching posterior margin of head. Labium: reaching apex of metacoxa; labial formula LII>LI>LIV>LIII; LI longer than bucculae, reaching middle part of proxyphus.

Pronotum: trapeziform; ca. 2.3x wider than long, ca. 1.7x wider than head; anterior part weakly raised, rounded, anterior margin slightly concave medially; posterior margin rectilinear; lateral margin straight; callar region deeply depressed medially, calli slightly raised, with impressed line; disc almost flat, weakly concave. Proepimera: flat.

Mesoscutum: not greatly exposed, less than ½ of scutellum length. Scutellum: disc slightly raised, width ca. 1.3x length. Hemelytra: MFL moderately long, weakly convex, reaching beyond scutellum, not reaching 1/3rd of claval commissure; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long.

Male: Unknown.

MEASUREMENTS. Table 3.1.

Host plant. Unknown.

Distribution. Found in Papua New Guinea.

Remarks. Pseudoloxops wareo is similar in appearance to P. kimiensis, but is distinguished from it by the following characters: the pronotum is broadly orange-red with a brownish tinge on the anterior and lateral regions; whereas in P. kimiensis, it is mostly brownish on the anterior and lateral regions, with a broadly pale yellow brown disc and the lateral areas red. Pseudoloxops wareo can also be distinguished from P.

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kimiensis by having a broadly red scutellum with a brownish apex and mostly red cuneus.

Pseudoloxops woodwardi new species

Figures 3.2, 3.17, 3.19

Etymology. This species is named in honour of the late Professor T. E. Woodward, who is one of the collectors of this species.

Type material. HOLOTYPE: Male: AUSTRALIA: Queensland: Brisbane,

27.46785°S 153.02801°E, 22 Mar 1955, T. E. Woodward, Light Trap, (USI 00192915)

(UQIC). PARATYPES: AUSTRALIA: Queensland: Bardon, Brisbane, 27.45°S

152.96666°E, 08 Apr 1947, A.C. Arvier, 1♂ (USI 00192914) (UQIC). Brighton,

27.28333°S 153.05°E, 26 Jan 1955, E.S. Reye & M. B. Wilson, Light Trap, 1♂ (USI

00192920) (UQIC). Brisbane, 27.46785°S 153.02801°E, 03 Jun 1932, Unknown, 1♀

(USI 00192970) (UQIC); 15 Mar 1952, S. Barker, 1♂ (USI 00192978) (UQIC); 29 Mar

1952, J. Letchford, 1♂ (USI 00192913) (UQIC); 31 Mar 1952, S. Barker, 1♂ (USI

00192919) (UQIC); 03 May 1955, R. E. Harrison, 1♂ (USI 00192962) (UQIC); 16 Dec

1955, T. E. Woodward, 1♀ (USI 00192963) (UQIC); Aug 1957, G. Ettershank, 1♀

(USI 00192973) (UQIC); 28 Jan 1958, H.G. Greening, 1♂ (USI 00192918) (UQIC); 01

Mar 1958, H.G. Greening, 1♀ (USI 00192971) (UQIC); 01 Apr 1958, A. Singh, 1♀

(USI 00192974) (UQIC); 22 Feb 1959, P.R. Webb, 1♂ (USI 00192961) (UQIC); 19 Oct

1961, F.J.D. McDonald, 1♂ (USI 00192916) (UQIC); 18 Apr 1963, J. C. Cardale, 1♀

(USI 00192972) (UQIC). Corinda, 27.53333°S 152.98333°E, 01 Sep 1952, J. Davis, 1♀

(USI 00192977) (UQIC). Deception Bay, 27.196°S 153.015°E, 25 Mar 1954, J. D. Beri,

1♂ (USI 00192979) (UQIC). Glen Aplin, 28.73333°S 151.86666°E, 14 Feb 1963, P.

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Kerridge, 1♀ (USI 00192976) (UQIC). Heron Island, 23.45°S 151.9167°E, 10 Nov

1957 - 14 Nov 1957, T. E. Woodward, 3♀ (USI 00192967, USI 00192969, USI

00192975) (UQIC); 11 Sep 1972 - 12 Sep 1972, B. Hocking, 1♂ (USI 00192942)

(UQIC). Maryborough, 25.538°S 152.702°E, 17 Apr 1938, R.V.C. Smythe, 1♂ (USI

00192912) (UQIC). Mt. Tambourine, 27.97253°S 153.20167°E, 16 Feb 1963, R.

Shepherd, 1♂ (USI 00192960) (UQIC). Southport, 27.974°S 153.405°E, 29 Feb 1964,

R. A. McLachlan, 1♀ (USI 00192968) (UQIC). University Grounds, St. Lucia,

Brisbane, 27.49777°S 153.01277°E, 13 Apr 1976, G. Thompson, Acacia maidenii

(Fabaceae), 1♂ (USI 00192917), 2♀ (USI 00192964, USI 00192965) (UQIC).

Diagnosis. Pseudoloxops woodwardi is recognised by the following combination of characters: head mostly pale yellow brown, frons with dark red markings; AI mostly red with pale yellow brown stripe at medial part; pronotum broadly pale yellow brown medially, red laterally; scutellum uniformly pale yellow brown; corium mostly pale yellow brown, sometimes tinged pale green, with red markings on basal part and apex of embolium; metafemora uniformly pale yellow brown; pygophore without tubercle process; apophysis and sensory lobe of right paramere armed with small teeth; left paramere Y-shaped, apophysis slightly sinuate, slim, parallel with sensory lobe, slightly longer in length to sensory lobe, apex slightly hooked; sensory lobe of left paramere broad; and, endosoma with single spicule, elongate, smooth margin, membranous basally.

Description. Male: Body elongate-ovate, total length 3.87-4.44 mm (Fig. 3.2).

COLOURATION (Fig. 3.2). Head: mostly pale yellow brown, frons with dark red markings at dorsum laterally extending to anterior part, anterior medially with minute red markings; clypeus pale yellow brown with minute red markings at dorsum, extending to midline to about 2/3rd length, then horizontal red marking, apical part pale

151

yellow brown, lateral areas basally with minute red markings; mandibular plate pale yellow brown with broad red horizontal markings medially; maxillary plate uniformly red with brown markings at tumescence; outer antennal insertion with red markings.

Antennae: AI mostly red, medial part paler with pale yellow brown stripe; AII pale yellow brown, reddish at base and apex; AIII pale yellow brown, apex red; AIV pale yellow brown, apex brownish. Labium: mostly pale yellow brown, dark brown apically.

Pronotum: lateral areas broadly sanguineous, reaching lateral part of calli, extending to dorsal margin of propleura; medial part broadly pale yellow brown. Propleura: proepimera mostly pale yellow brown, dorsal areas with red markings; proepisterna mostly red, with posterior areas pale yellow brown. Mesoscutum: mostly pale yellow brown, anterolateral angles red. Scutellum: uniformly pale yellow brown. Hemelytra: clavus mostly pale yellow brown with red markings at basal part, reaching about ½ of mesoscutum length then extending to about 2/3rd of scutellum length, apical part red about ¼ of commissure length; corium mostly pale yellow brown, basal part red reaching about level 2/3rd of scutellum length; apex of embolium red; cuneus mostly pale yellow brown with red markings at mediobasal angles, apical part red (about ¼ of length) with dark red at apex; membrane smoky brown, veins red. Abdomen: uniformly pale yellow brown. Legs: uniformly pale yellow brown. VESTITURE. Head: dorsal surface setose with pale, short to long, bristlelike setae. Antennae: AI densely setose with pale, moderately long, semierect bristlelike setae, intermixed sparsely with pale, long, erect bristlelike setae; AII and AIII densely setose with pale, short, decumbent bristlelike setae, intermixed sparsely with pale, short, semierect bristlelike setae, basal part with a few pale, short, erect bristlelike setae; AIV densely setose with pale, short, semierect bristlelike setae, intermixed with a few pale, moderately long, semierect bristlelike setae. Pronotum, mesoscutum and scutellum: setose with pale, long,

152

bristlelike setae, intermixed with pale, long, erect bristlelike setae. Hemelytra: densely setose with pale, long, semierect bristlelike setae, intermixed sparsely with pale, long, erect bristlelike setae. Legs: femora densely setose with pale, short, semierect bristlelike setae, intermixed sparsely with pale, moderately long, erect bristlelike setae; tibiae densely setose with pale, short, semierect bristlelike setae, intermixed sparsely with pale to smoky brown, long tibial spines; hind tibiae with several longitudinal, parallel rows of tiny spicules. STRUCTURE. Head: frons convex in dorsal view, moderately projected anteromedially, somewhat flat; vertex flat in lateral view, midline deeply sulcate, posterior margin carinate; clypeus visible in dorsal view, large, subrectangular; mandibular plate moderately sized, weakly visible in dorsal view, margin slightly convex, somewhat rounded; maxillary plate moderately sized, with tumescence, greatly, slightly exceeding mandibular plate, convex; bucculae reaching posterior margin of head. Labium: reaching apex of mesocoxae; LI longer than bucculae, reaching proxyphus. Eyes: moderately sized; nearly reaching dorsal margin of head in lateral view, nearly reaching bucculae ventrally; exceeding anterolateral angles of pronotum, posterior margin contiguous with anterior margin of pronotum. Antennae: antennifers moderately sized, contiguous with posterior margin of mandibular plates; AI moderately long, 1.2-1.4x longer than IOD; AII cylindrical, 3.2-3.4x longer than AI,

1.3-1.4x longer than pronotum width; AIII and AIV slender; AII>AIII>AIV>AI.

Pronotum: trapeziform; 2.6-2.7x wider than long, 1.4-1.5x wider than head; anterior margin rounded, weakly concave medially; lateral margin weakly concave; posterior margin slightly excavate; calli slightly raised, with impressed line; disc flat.

Proepimera: almost flat. Mesoscutum: exposed, ca. ½ scutellum length; posterior areas depressed medially. Scutellum: disc flat, 1.2-1.4x wider than long. Hemelytra: MFL weakly convex, reaching beyond apex of scutellum, not reaching 1/3rd of claval

153

commissure; cuneus moderately long, longer than wide, ca. 1.5x width, not greatly narrowed; membrane cells moderately long. Abdomen: surpassing clavus, reaching anterior margin of cuneus. Male genitalia: pygophore with weakly concave dorsal margin, with medial lobe, ventral margin convex (Fig. 3.17a); right paramere somewhat

C-shaped, apophysis somewhat bulbous, armed with sclerotized small teeth, with dorsal medial process, somewhat short, apically armed with sclerotized small teeth (Fig.

3.17b); left paramere Y-shaped, sensory lobe broad, unarmed, apically nearly truncate, apophysis elongate, slightly sinuate margins, slim, slightly longer in length to sensory lobe, apex slightly hooked, base with tumescence, rounded (Fig. 3.17c); phallotheca opening dextrodorsally to medial region, ventral surface with left apical fold, reduced, somewhat sinuous margin (Fig. 3.17d); endosoma with single spicule, strongly sclerotized, elongate, somewhat broad basally, tapering to apex, margin sinuate, membranous basally (Fig. 3.17e).

Female: Total length 3.83-4.27 mm. Colouration and vestiture as in male (Fig. 3.2).

STRUCTURE: Mostly as in male; Eyes: not reaching dorsal margin of head in lateral view, not reaching bucculae ventrally. Antennae: AI moderately long, longer than IOD,

1.1-1.4x longer than IOD; AII cylindrical, 3.0-3.3x longer than AI, 1.2-1.4x longer than pronotum width. Pronotum: 2.4-2.9x as wide as long, 1.5-1.6x as wide as head.

Scutellum: 1.1-1.4x as wide as long.

MEASUREMENTS. Table 3.1.

Host plant. The specimens of P. woodwardi found on the university grounds at St.

Lucia, Brisbane, were found on flowers of Acacia maidenii F. Muell (Fabaceae).

Distribution. P. woodwardi is known from several localities in Queensland.

Remarks. Pseudoloxops woodwardi is most similar to P. tatarnici, but differs by the following characters: the frons with dark red markings; the mandibular plate with broad

154

red markings medially; the maxillary plate uniformly red; the anterolateral angles of the mesoscutum red; the scutellum uniformly pale yellow brown; the clavus with red markings only on base and apex; the corium red basally; the pygophore with a rounded ventral margin, the dorsal margin with a lobe and without dorsal processes; the right paramere with a broader medial process and apophysis; the left paramere with a broad sensory lobe, basal process present; and, the endosoma with a single elongate spicule, which is membranous basally.

155

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Table 3.1. Measurements of Pseudoloxops species of the Southwest Pacific.

Length Width Species Total Clyp-Cun Head Pronotum Scutellum Cuneus Head Pronotum Scutellum InterOcDi AI AII AIII AIV baiawaensis Male Mean 3.51 2.52 0.45 0.38 0.43 0.49 0.65 0.95 0.55 0.28 0.44 1.49 0.51 0.46 SD 0.11 0.09 0.02 0.02 0.01 0.02 0.02 0.03 0.02 0.02 0.02 0.05 0.02 0.04 Min 3.33 2.37 0.42 0.34 0.41 0.46 0.63 0.91 0.53 0.26 0.41 1.45 0.48 0.39 Max 3.60 2.59 0.47 0.40 0.45 0.50 0.69 0.99 0.57 0.31 0.48 1.57 0.54 0.49 N = 5 5 5 5 5 5 5 5 5 5 5 5 5 5 Female Mean 3.82 2.71 0.46 0.40 0.45 0.55 0.64 1.01 0.59 0.31 0.54 1.55 0.57 0.52 SD 0.23 0.14 0.02 0.03 0.04 0.05 0.03 0.07 0.06 0.01 0.02 0.11 0.02 0.01 Min 3.43 2.47 0.45 0.35 0.39 0.46 0.60 0.93 0.50 0.29 0.50 1.35 0.54 0.51 Max 4.04 2.80 0.49 0.43 0.48 0.59 0.68 1.10 0.66 0.33 0.56 1.63 0.60 0.54 N = 5 5 5 5 5 5 5 5 5 5 5 5 5 5 cheesmanae Male 4.76 3.44 0.57 0.50 0.52 0.74 0.87 1.20 0.65 0.39 0.59 1.89 0.75 0.73 N = 1 1 1 1 1 1 1 1 1 1 1 1 1 1 cordiformis Male 3.69 2.66 0.48 0.38 0.45 0.55 0.62 0.86 0.54 0.26 0.49 1.43 0.54 0.52 N = 1 1 1 1 1 1 1 1 1 1 1 1 1 1 gagnei Female 5.75 4.09 0.65 0.67 0.64 0.82 0.85 1.42 0.77 0.36 0.72 2.30 0.86 0.65 N = 1 1 1 1 1 1 1 1 1 1 1 1 1 1 kimiensis Male 5.31 3.63 0.57 0.57 0.56 0.91 0.88 1.26 0.66 0.33 0.62 2.23 0.69 2.17 N = 1 1 1 1 1 1 1 1 1 1 1 1 1 1 kraussi Male Mean 3.90 2.78 0.49 0.41 0.47 0.61 0.75 1.04 0.54 0.31 0.49 1.57 0.57 0.56 SD 0.10 0.09 0.02 0.02 0.02 0.03 0.04 0.07 0.01 0.03 0.02 0.07 0.06 0.06 Min 3.77 2.68 0.47 0.39 0.44 0.57 0.69 0.95 0.53 0.28 0.47 1.47 0.51 0.49 Max 4.04 2.90 0.51 0.44 0.49 0.64 0.78 1.16 0.56 0.34 0.52 1.63 0.63 0.63 N = 5 5 5 5 5 5 5 5 5 5 5 4 4 4

163 Table 3.1. Cont/-.

Length Width Species Total Clyp-Cun Head Pronotum Scutellum Cuneus Head Pronotum Scutellum InterOcDi AI AII AIII AIV kraussi Female Mean 4.18 3.00 0.50 0.43 0.49 0.62 0.71 1.13 0.62 0.36 0.53 1.62 0.61 0.54 SD 0.19 0.12 0.01 0.03 0.02 0.05 0.01 0.06 0.02 0.02 0.02 0.07 0.09 0.09 Min 3.95 2.84 0.48 0.39 0.46 0.57 0.70 1.09 0.60 0.34 0.50 1.52 0.51 0.49 Max 4.42 3.15 0.52 0.45 0.51 0.70 0.72 1.22 0.65 0.37 0.54 1.69 0.73 0.68 N = 5 5 5 5 5 5 5 5 5 5 4 4 4 4 lenakelensis Male 4.29 3.02 0.52 0.40 0.48 0.65 0.79 1.06 0.59 0.34 0.60 1.82 0.71 N = 1 1 1 1 1 1 1 1 1 1 1 1 1 mcilwraithensis Male Mean 4.35 3.08 0.54 0.46 0.47 0.70 0.83 1.06 0.57 0.32 0.43 1.73 0.68 SD 0.23 0.16 0.02 0.02 0.01 0.05 0.01 0.05 0.03 0.01 0.01 0.15 Min 4.09 2.90 0.52 0.44 0.47 0.66 0.83 1.02 0.55 0.32 0.43 1.62 Max 4.53 3.17 0.56 0.48 0.48 0.76 0.84 1.11 0.60 0.33 0.44 1.90 N = 3 3 3 3 3 3 3 3 3 3 3 3 1 Female Mean 4.71 3.34 0.56 0.45 0.48 0.78 0.77 1.12 0.59 0.40 0.46 1.73 0.74 0.53 SD 0.00 0.05 0.02 0.01 0.07 0.02 0.02 0.04 0.04 0.01 0.00 0.00 0.04 0.07 Min 4.70 3.30 0.54 0.45 0.43 0.77 0.76 1.09 0.56 0.40 0.46 1.73 0.71 0.48 Max 4.71 3.37 0.57 0.46 0.53 0.80 0.78 1.15 0.62 0.41 0.47 1.73 0.77 0.58 N = 2 2 2 2 2 2 2 2 2 2 2 2 2 2 mourangeensis Male 5.05 3.15 0.49 0.39 0.37 1.01 0.76 1.03 0.55 0.34 0.47 4.71 N = 1 1 1 1 1 1 1 1 1 1 1 1 Female 4.40 3.03 0.53 0.40 0.48 0.83 0.40 0.50 1.64 0.86 N = 1 1 1 1 1 1 1 1 1 1 naraiyawa Male 4.78 3.46 0.55 0.53 0.56 0.74 0.77 1.23 0.66 0.36 0.65 2.08 N = 1 1 1 1 1 1 1 1 1 1 1 1

164 Table 3.1. Cont/-.

Length Width Species Total Clyp-Cun Head Pronotum Scutellum Cuneus Head Pronotum Scutellum InterOcDi AI AII AIII AIV pancheriaphila Male Mean 4.16 2.97 0.51 0.44 0.50 0.63 0.75 1.03 0.59 0.35 0.53 1.68 0.64 0.56 SD 0.08 0.08 0.03 0.02 0.02 0.02 0.02 0.03 0.01 0.01 0.02 0.03 0.02 0.03 Min 4.07 2.89 0.49 0.42 0.47 0.60 0.73 1.00 0.58 0.33 0.52 1.64 0.63 0.52 Max 4.23 3.08 0.55 0.49 0.53 0.65 0.77 1.06 0.60 0.37 0.55 1.72 0.67 0.59 N = 5 5 5 5 5 5 5 5 5 5 5 5 5 5 Female Mean 4.20 2.97 0.50 0.43 0.49 0.62 0.67 1.02 0.61 0.38 0.56 1.62 0.65 0.54 SD 0.06 0.03 0.02 0.03 0.02 0.02 0.04 0.06 0.02 0.03 0.01 0.03 0.02 0.03 Min 4.11 2.93 0.47 0.41 0.47 0.60 0.62 0.97 0.59 0.33 0.55 1.59 0.63 0.50 Max 4.25 3.01 0.53 0.47 0.52 0.64 0.71 1.12 0.63 0.40 0.57 1.66 0.69 0.59 N = 5 5 5 5 5 5 5 5 5 5 5 5 5 5 sikaiana Male Mean 4.05 2.95 0.50 0.46 0.50 0.60 0.74 1.09 0.59 0.31 0.51 1.90 0.74 0.71 SD 0.07 0.09 0.04 0.00 0.00 0.00 0.04 0.02 0.03 0.01 0.02 Min 4.00 2.89 0.47 0.45 0.50 0.60 0.71 1.08 0.57 0.30 0.50 Max 4.10 3.01 0.52 0.46 0.50 0.60 0.77 1.10 0.61 0.31 0.52 N = 2 2 2 2 2 2 2 2 2 2 2 1 1 1 Female Mean 4.20 3.08 0.54 0.46 0.51 0.61 0.76 1.14 0.64 0.36 0.52 1.79 0.67 0.65 SD 0.16 0.13 0.00 0.01 0.03 0.05 0.04 0.06 0.02 0.01 0.03 0.23 0.03 Min 4.08 2.99 0.53 0.45 0.49 0.57 0.73 1.10 0.62 0.35 0.50 1.63 0.65 Max 4.31 3.17 0.54 0.46 0.53 0.64 0.78 1.19 0.65 0.36 0.55 1.95 0.69 N = 2 2 2 2 2 2 2 2 2 2 2 2 2 1 solomonensis Male Mean 3.84 2.78 0.51 0.40 0.48 0.56 0.72 1.00 0.54 0.29 0.48 1.65 0.59 0.61 SD 0.07 0.06 0.01 0.03 0.02 0.01 0.05 0.07 0.03 0.02 0.02 0.10 0.03 Min 3.77 2.72 0.50 0.36 0.46 0.55 0.65 0.93 0.51 0.27 0.46 1.53 0.57 Max 3.91 2.82 0.53 0.42 0.50 0.57 0.75 1.05 0.56 0.31 0.50 1.71 0.61 N = 3 3 3 3 3 3 3 3 3 3 3 3 2 1

165 Table 3.1. Cont/-.

Length Width Species Total Clyp-Cun Head Pronotum Scutellum Cuneus Head Pronotum Scutellum InterOcDi AI AII AIII AIV solomonensis Female Mean 4.10 2.96 0.51 0.41 0.48 0.62 0.70 1.10 0.60 0.35 0.49 1.62 0.67 0.55 SD 0.14 0.02 0.01 0.01 0.03 0.03 0.02 0.03 0.02 0.01 0.00 0.06 0.05 Min 3.98 2.94 0.49 0.41 0.43 0.59 0.67 1.07 0.58 0.34 0.48 1.56 0.64 Max 4.25 2.98 0.53 0.43 0.51 0.65 0.73 1.13 0.62 0.37 0.49 1.67 0.73 N = 4 4 4 4 4 4 4 4 4 4 3 3 3 1 straatmani Male 3.58 2.61 0.46 0.37 0.37 0.50 0.70 0.84 0.48 0.25 0.50 1.66 0.69 N = 1 1 1 1 1 1 1 1 1 1 1 1 1 Female 4.37 3.21 0.53 0.48 0.45 0.67 0.73 1.07 0.58 0.33 0.63 N = 1 1 1 1 1 1 1 1 1 1 1 tabwemasana Male Mean 4.57 3.34 0.53 0.48 0.52 0.73 0.82 1.23 0.64 0.35 0.53 1.90 0.73 0.70 SD 0.20 0.05 0.01 0.01 0.03 0.06 0.02 0.04 0.04 0.02 0.01 0.06 0.04 0.06 Min 4.28 3.28 0.51 0.47 0.47 0.63 0.79 1.18 0.60 0.34 0.52 1.82 0.69 0.62 Max 4.77 3.39 0.54 0.49 0.55 0.76 0.85 1.29 0.70 0.37 0.54 1.97 0.78 0.76 N = 5 5 5 5 5 5 5 5 5 5 5 5 5 5 Female Mean 4.76 3.40 0.53 0.49 0.54 0.73 0.73 1.21 0.64 0.38 0.57 1.75 0.73 0.69 SD 0.15 0.11 0.02 0.02 0.02 0.01 0.02 0.03 0.04 0.01 0.03 0.08 0.05 0.07 Min 4.61 3.26 0.52 0.47 0.52 0.72 0.72 1.18 0.61 0.38 0.54 1.67 0.65 0.61 Max 4.96 3.51 0.55 0.51 0.56 0.75 0.76 1.25 0.69 0.39 0.61 1.86 0.77 0.77 N = 5 5 5 5 5 5 5 5 5 5 5 4 4 4 tatarnici Male 3.57 2.56 0.47 0.41 0.43 0.49 0.74 0.95 0.62 0.36 0.36 1.26 0.51 Count 1 1 1 1 1 1 1 1 1 1 1 1 1 Female Mean 3.53 2.44 0.45 0.35 0.38 0.55 0.68 0.93 0.49 0.34 0.38 1.20 0.54 0.45 SD 0.10 0.06 0.02 0.01 0.05 0.01 0.02 0.01 0.01 0.02 0.02 0.04 0.02 0.03 Min 3.44 2.37 0.44 0.34 0.33 0.55 0.66 0.92 0.48 0.33 0.36 1.15 0.53 0.42 Max 3.64 2.49 0.47 0.36 0.41 0.56 0.70 0.93 0.49 0.36 0.40 1.23 0.56 0.49 N = 3 3 3 3 3 3 3 3 3 3 3 3 3 3

166 Table 3.1. Cont/-.

Length Width Species Total Clyp-Cun Head Pronotum Scutellum Cuneus Head Pronotum Scutellum InterOcDi AI AII AIII AIV vitiensis Male Mean 3.82 2.78 0.51 0.42 0.46 0.55 0.70 0.98 0.53 0.33 0.49 1.59 0.65 0.55 SD 0.02 0.03 0.02 0.00 0.02 0.01 0.03 0.03 0.02 0.01 0.02 0.05 0.02 0.08 Min 3.80 2.76 0.49 0.42 0.44 0.54 0.67 0.95 0.52 0.32 0.47 1.54 0.63 0.50 Max 3.83 2.81 0.53 0.42 0.48 0.56 0.74 1.01 0.56 0.34 0.51 1.65 0.66 0.60 N = 3 3 3 3 3 3 3 3 3 3 3 3 2 2 Female Mean 4.15 3.04 0.51 0.43 0.46 0.60 0.72 1.07 0.60 0.38 0.51 1.62 0.64 0.52 SD 0.22 0.18 0.01 0.03 0.03 0.01 0.03 0.03 0.03 0.02 0.02 0.06 0.08 0.07 Min 3.79 2.74 0.50 0.39 0.41 0.58 0.68 1.04 0.56 0.35 0.50 1.52 0.53 0.44 Max 4.34 3.20 0.53 0.47 0.49 0.61 0.76 1.11 0.63 0.39 0.55 1.68 0.70 0.58 N = 5 5 5 5 5 5 5 5 5 5 5 5 4 3 wareo Male 5.05 3.76 0.58 0.65 0.63 0.74 0.87 1.49 0.81 0.42 N = 1 1 1 1 1 1 1 1 1 1 woodwardi Male Mean 4.05 2.88 0.49 0.41 0.48 0.60 0.70 1.06 0.61 0.34 0.44 1.47 0.53 0.48 SD 0.24 0.15 0.02 0.02 0.03 0.04 0.03 0.04 0.02 0.01 0.02 0.06 0.03 0.03 Min 3.87 2.76 0.47 0.39 0.45 0.56 0.66 1.03 0.58 0.32 0.42 1.38 0.49 0.45 Max 4.44 3.12 0.51 0.44 0.52 0.66 0.73 1.14 0.64 0.36 0.48 1.56 0.57 0.51 N = 5 5 5 5 5 5 5 5 5 5 5 5 5 4 Female Mean 4.01 2.87 0.49 0.39 0.48 0.59 0.67 1.05 0.59 0.38 0.46 1.42 0.53 0.45 SD 0.17 0.10 0.01 0.01 0.03 0.05 0.05 0.08 0.04 0.03 0.03 0.07 0.02 0.04 Min 3.83 2.73 0.47 0.38 0.45 0.54 0.62 0.96 0.55 0.35 0.42 1.35 0.51 0.39 Max 4.27 2.99 0.51 0.41 0.52 0.65 0.74 1.15 0.63 0.41 0.50 1.53 0.54 0.47 N = 5 5 5 5 5 5 5 5 5 5 5 5 4 4

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Figure 3.1. Habitus photographs of Pseudoloxops species of the Southwest Pacific. Plate 1 of 2. Scale = 1 mm.

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Figure 3.2. Habitus photographs of Pseudoloxops species of the southwest Pacific. Plate 2 of 2. Scale = 1 mm.

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Figure 3.3. Male genitalia of Pseudoloxops baiawaensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; sl = sensory lobe of left paramere; vp = ventral process of pygophore; DM = dorsal margin of pygophore; VM = ventral margin of pygophore; DES = dorsal endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 170

Figure 3.4. Male genitalia of Pseudoloxops efatensis. a. Right paramere. b. Left paramere. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; sl = sensory lobe of left paramere. Scale = 0.1 mm.

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Figure 3.5. Male genitalia of Pseudoloxops kimiensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; sl = sensory lobe of left paramere; vmp = ventral medial process of right paramere. Scale = 0.1 mm.

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Figure 3.6. Male genitalia of Pseudoloxops kraussi. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, left ventral view. e. Phallotheca, dorsal view. f. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; bl = basal lobe of phallotheca; bp = basal process of left paramere; dsl = dorsal subapical lobe of phallotheca; dmp = dorsal medial process of right paramere; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 173

Figure 3.7. Male genitalia of Pseudoloxops lenakelensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Aedeagus, left lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; dsl = dorsal subapical lobe of phallotheca; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm.

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Figure 3.8. Male genitalia of Pseudoloxops mcilwraithensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, dorsal view. e. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; dmp = dorsal medial process of right paramere; DES1 = first dorsal endosomal spicule of aedeagus; DES2 = second endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm.

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Figure 3.9. Male genitalia of Pseudoloxops naraiyawa. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; al = apical lobe of phallotheca; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; dsl = dorsal subapical lobe of phallotheca; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 176

Figure 3.10. Male genitalia of Pseudoloxops pancheriaphila. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dsl = dorsal subapical lobe of phallotheca; dmp = dorsal medial process of right paramere; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; SG = secondary gonopore of aedeagus. Scale = 0.1 mm.

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Figure 3.11. Male genitalia of Pseudoloxops sikaiana. a. Pygophore, dorsal view. b. Right paramere, left lateral view. c. Right paramere, ventral view. d. Left paramere. e. Phallotheca, right lateral view. f. Phallotheca, dorsal view. g. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; sl = sensory lobe of left paramere; DES = dorsal endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 178

Figure 3.12. Male genitalia of Pseudoloxops solomonensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, dorsal view. Abbreviations: ap = apophysis of right and left parameres; sl = sensory lobe of left paramere; vp = ventral process of pygophore; DES = dorsal endosomal spicule of aedeagus; VES = ventral endosomal spicule of aedeagus; SG = secondary gonopore of aedeagus. Scale = 0.1 mm.

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Figure 3.13. Male genitalia of Pseudoloxops straatmani. a. Pygophore, ventral view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; sl = sensory lobe of left paramere; vp = ventral process of pygophore; DES1 = first dorsal endosomal spicule; DES2 = second dorsal endosomal spicule; DES3 = third dorsal endosomal spicule; VES = ventral endosomal spicule; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 180

Figure 3.14. Male genitalia of Pseudoloxops tabwemasana. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, left lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dsl = dorsal subapical lobe of phallotheca; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; vmp = ventral medial process of right paramere; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 181

Figure 3.15. Male genitalia of Pseudoloxops tatarnici. a. Pygophore, dorsal view. b. Pygophore, dorsal view, with parameres and pygophore processes in situ. c. Right paramere. d. Left paramere. e. Phallotheca, right lateral view. f. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; dmp = dorsal medial process of right paramere; lp = left paramere; rp = right paramere; sl = sensory lobe of left paramere; DES1 = first dorsal endosomal spicule; DES2 = second dorsal endosomal spicule; DES3 = third dorsal endosomal spicule; VES = ventral endosomal spicule; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 182

Figure 3.16. Male genitalia of Pseudoloxops vitiensis. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, right lateral view. e. Phallotheca, dorsal view. f. Aedeagus, left lateral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; dsl = dorsal subapical lobe of phallotheca; sl = sensory lobe of left paramere; vf = ventral fold of phallotheca; vp = ventral process of pygophore; SG = secondary gonopore of aedeagus. Scale = 0.1 mm. 183

Figure 3.17. Male genitalia of Pseudoloxops woodwardi. a. Pygophore, dorsal view. b. Right paramere. c. Left paramere. d. Phallotheca, dorsal view. e. Aedeagus, ventral view. Abbreviations: ap = apophysis of right and left parameres; bp = basal process of left paramere; dmp = dorsal medial process of right paramere; ml = medial lobe of dorsal margin of pygophore; sl = sensory lobe of left paramere; SG = secondary gonopore of aedeagus. Scale = 0.1 mm.

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Figure 3.18. Distribution map of Pseudoloxops species of the Southwest Pacific. Map 1 of 2. 185

Figure 3.19. Distribution map of Pseudoloxops species of the Southwest Pacific. Map 2 of 2. 186

CHAPTER IV

GENERAL CONCLUSIONS

A comparative morphological study of all described species and new species provisionally assigned to Pseudoloxops has been conducted. The phylogenetic analysis presented in this study is the first undertaken for Pseudoloxops. Based on the results, the genus Pseudoloxops is recognised as a monophyletic group, which is strongly supported by two non-homoplasious synapomorphies: the head flat or nearly flat in dorsal view and the frons convex from lateral view. Pseudoloxops has an Eastern Hemisphere distribution with centres of diversity and endemism in Asia, Africa, Australia and island archipelagos in the Pacific Basin. The clades within Pseudoloxops sometimes correlate with distribution, such as a clade from French Polynesia, although other species from regions, such as those from Australia and the southwest Pacific, are found in different clades of Pseudoloxops.

My systematic study of Pseudoloxops species focused on those from Australia and the southwest Pacific, and is the first systematic review of the genus within this region.

Prior to this study, Pseudoloxops consists of 38 species worldwide. In this study, a total of 20 species from the southwest Pacific were described, with 19 new species and the previously described species from Fiji, Pseudoloxops vitiensis Kirkaldy 1908. This increases the number of described species of Pseudoloxops to 57 species. A key to all the species is provided for the southwest Pacific. Based on a comparative morphological study of external and male genitalic characters, I found that both external and male genitalic characters are useful to separate the species. There is considerable

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variation in colour patterns and the male genitalia, which are useful and often corroborate decisions concerning species discrimination.

Future directions for the study of Pseudoloxops would require increasing the sample of many of the species from all regions of the world. The phylogeny of Pseudoloxops in this study has been limited by the availability of males which has prevented a broader examination of male genitalic characters. Also, there are many species for which there are few female specimens, and an investigation of their genitalia has not been possible, but would be a future test of my classification.

Any future study of Pseudoloxops also requires a broader investigation of genera of

Orthotylini so that my theory of its monophyly is tested and our understanding of its position within the tribe is enhanced. In addition, documentation of host plants and distribution is also needed to advance our knowledge of host plant and area relationships. Finally, my systematic and phylogenetic study of the genus Pseudoloxops provides a framework for a monograph of the whole genus in the future.

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