145923291 อนุกรมวิธานของมดสกุล Platythyrea ROGER, 1863 (HYMENOPTERA: FORMICIDAE: PONERINAE) ในประเทศไทย BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 TAXONOMY OF THE ANT GENUS Platythyrea ROGER, 1863 (HYMENOPTERA: FORMICIDAE: PONERINAE) IN THAILAND

NATTHAPORN PHENGSI

BURAPHA UNIVERSITY 2017

59910009_145923291

อนุกรมวิธานของมดสกุล Platythyrea ROGER, 1863 (HYMENOPTERA: FORMICIDAE: PONERINAE) ในประเทศไทย

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

นัฏฐพร เพ็งศรี

วทิ ยานิพนธ์น้ีเป็นส่วนหน่ึงของการศึกษาตามหลกั สูตรวทิ ยาศาสตรมหาบณั ฑิต สาขาวิชาวิทยาศาสตร์ชีวภาพ คณะวิทยาศาสตร์ มหาวิทยาลัยบูรพา 2560 ลิขสิทธ์ิของมห าวทิ ยาลยั บูรพา

TAXONOMY OF THE ANT GENUS Platythyrea ROGER, 1863 (HYMENOPTERA: FORMICIDAE: PONERINAE) IN THAILAND

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

NATTHAPORN PHENGSI

A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE MASTER OF SCIENCE IN BIOLOGICAL SCIENCES FACULTY OF SCIENCE BURAPHA UNIVERSITY 2017 COPYRIGHT OF BURAPHA UNIVERSITY

D

ABST RACT 59910009: MAJOR: BIOLOGICAL SCIENCES; M.Sc. (BIOLOGICAL SCIENCES) KEYWORDS Ant/ Distribution/ Ponerinae/ Taxonomy/ Thailand : NATTHAPORN PHENGSI: TAXONOMY OF THE ANT GENUS Platythyrea ROGER, 1863 (HYMENOPTERA: FORMICIDAE: PONERINAE) IN THAILAND . ADVISORY COMMITTEE: SALINEE KHACHONPISITSAK, WEEYAWAT JAITRONG, 2017

145923291 The Platythyrea Roger, 1863, a ponerine genus, is arboreal ants, nesting in hollow branches or other preformed cavities in live or fallen trees and foraging on tree trunks or other vegetation. Member of the ant genus Platythyrea are rarely collected. The present taxonomic study is the update revision of Platythyrea. It is

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 largely based on the materials deposited in museums of Thailand. A total of seven species of the ant genus Platythyrea were reported from Thailand of which two species are new to science. Based on external morphology of workers, they belong to the two distinct species-groups: P. clypeata group (P. clypeata Forel, 1911, P. grandis sp. nov., and P. janyai Phengsi, Jaitrong, Ruangsittichai et Khachonpisitsak, 2018) and P. paralella group (P. itoi sp. nov., P. paralella (F. Smith, 1859), P. quadridenta Donistrope, 1941, and P. tricuspidata Emery, 1900). The characteristics of external morphology that can be used for separating two species groups are shape of propodeal spiracle, frontal carina condition, shape of propodeal junction, and petiole shape. In terms of distribution, P. paralella is the most widely distributed while P. clypeata is clearly restricted to eastern Thailand. Also, P. itoi sp. nov. is apparently restricted to northern Thailand whereas P. janyai, P. quadridenta, and P. tricuspidata are clearly restricted to southern Thailand. A key to known species and distribution map of the ant genus Platythyrea are presented here.

E

ACKNOWLEDGEMENTS ACKNOWLEDGEMENTS

I would like to express my gratitude to the advisory committee members, Dr. Salinee Khachonpisitsak (advisor), Dr. Weeyawat Jaitrong (co-advisor), Associate Professor Dr. Nantasak Pinkaew (principal examiner), and Assistant Professor Dr. Chanawat Tuntiwaranuruk (member) for their kind advice, comments, suggestions, and discussion throughout this study. 145923291 I also grateful to Assoc. Prof. Decha Wiwatwitaya (Kasetsart University, Thailand) and Prof. Fuminori Ito (Kagawa University, Japan), who kindly allowed to

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 examine the ant specimens for this study in their collections and Yudthana Samung (Mahidol University, Thailand) for imaging assistance in this paper. This work was supported by Grant for Graduate Student 2016–2017 from Faculty of Science, Burapha University, Thailand. Finally, my deep gratitude goes to my family and friends for their presence in my life. They have been encouraging me to get my master's degree and beyond myself.

NATTHAPORN PHENGSI

TABLE OF CONTENTS

Page

ABSTRACT ...... D ACKNOWLEDGEMENTS ...... E

145923291 TABLE OF CONTENTS ...... F LIST OF TABLES ...... G

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 LIST OF FIGURES ...... H CHAPTER 1 ...... 1 INTRODUCTION ...... 1 CHAPTER 2 ...... 3 LITERATURE REVIEWS ...... 3 CHAPTER 3 ...... 18 MATERIALS AND METHODS ...... 18 CHAPTER 4 ...... 27 RESULTS ...... 27 CHAPTER 5 ...... 68 DISCUSSION AND CONCLUSION ...... 68 REFERENCES ...... 71 VITA ...... 77 LIST OF TABLES

Table 2-1. Colony size of the ant genus Platythyrea...... 7 Table 2-2. World distribution of the ant genus Platythyrea based on political regions...... 9 Table 4-1. Distribution of Platythyrea in Thailand. N, Northern; NE, Northeastern; E, Eastern; W, Western; C, Central; and S, Southern...... 67 LIST OF FIGURES

Figure 2-1. External morphology of the ant (Source:https://en.wikipedia.org/wiki/Ant)...... 4 Figure 2-2. External morphology of the ant genus Platythyrea. A, body in profile view; B, head in full-face view; and C, body in dorsal view...... 6 Figure 2-3. World distribution map of the ant genus Platythyrea (Antwiki, 2018)...... 9 Figure 3-1. Map showing collection sites for Platythyrea specimens in Thailand...... 19 Figure 3-2. Term of relative position for Platythyrea specimens: A, head in full-face view; and B, body in dorsal view...... 22 Figure 3-3. Whole body in profile view of Platythyrea...... 23 Figure 3-4. Mesosoma in profile view of Platythyrea...... 23 Figure 3-5. Triangular mandible of Platythyrea clypeata group...... 24 Figure 3-6. Morphometric measurements. A, head in full-face view; B, antenna; C, body in profile view: D, petiole in profile view; and E, petiole in dorsal view...... 26 Figure 4-1. Worker of Platythyrea clypeata. A, body in profile view; B, head in full- face view; and C, body in dorsal view...... 35 Figure 4-2. Queen of Platythyrea clypeata. A, body in profile view; B, head in full- face view; and C, body in dorsal view...... 36 Figure 4-3. Male of Platythyrea clypeata. A, body in profile view; B, head in full-face view; and C, body in dorsal view...... 37 Figure 4-4. SEM images of Platythyrea clypeata (A1–A3) and P. janyai (B1–B3). A1, B1 Sculpture on lateral face of head; A2, B2 Ommatidia of eye; A3, B3 Petiole in profile view...... 38 Figure 4-5. Worker of Platythyrea grandis. A, body in profile view; B, head in full- face view; and C, body in dorsal view...... 42 Figure 4-6. Worker of Platythyrea itoi. A, body in profile view; B, head in full-face view; and C, body in dorsal view...... 46 Figure 4-7. Worker of Platythyrea janyai. A, body in profile view; B, head in full-face view; and C, body in dorsal view...... 50 Figure 4-8. Worker of Platythyrea paralella. A, body in profile view; B, head in full- face view; and C, body in dorsal view...... 56 Figure 4-9. Queen of Platythyrea paralella. A, body in profile view; B, head in full- face view; and C, body in dorsal view...... 57 Figure 4-10. Worker of Platythyrea quadridenta. A, body in profile view; B, head in full-face view; and C, body in dorsal view...... 61 Figure 4-11. Worker of Platythyrea tricuspidata. A, body in profile view; B, head in full-face view; and C, body in dorsal view...... 64 Figure 4-12. Frontal view focusing on the frontal carinae. A, Frontal carinae very widely spaced; and B, Frontal carinae relatively narrowly separated Phengsi et al., 2018)...... 65 Figure 4-13. Dorsal view focusing on the petiole. A, Posterior margin of petiole with two spines, teeth or blunt angles; B. Posterior margin of petiole with three spines, teeth or blunt angles; and C, Posterior margin of petiole without distinct spines, teeth or blunt angles (Phengsi et al., 2018)...... 65 Figure 4-14. Distribution map of the ant genus Platythyrea in Thailand...... 66

CHAPTER 1 INTRODUCTION

1.1 Statements and significance of the problems Platythyrea Roger, 1863 is a ponerine genus of the tribe Platythyreini, with Pachycondyla punctata Bingham, 1903 as the type species (Bolton, 2003; Schmidt & 145923291 Shattuck, 2014). The genus is mainly distributed in pantropical regions, with some species also occurring in subtropical regions of the New World, Africa, Asia, and

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Australia (Bolton, 2003; Schmidt & Shattuck, 2014; Antweb, 2016). Members of the genus have been reported nesting in hollow branches that decompose hollow branches or nesting in other preformed cavities in live or fallen trees, and foraging on tree trunks or other vegetation (Brown, 1975; Ito, 1994; Djiéto-Lordon, Orivel & Dejean, 2001; Yéo, Molet & Peeters, 2006; Molet & Peeters, 2006; Ito, 2016). Some large African species nest under in termitaries of Cubitermes at the base of trees or under rocks that nesting under them increases soil minerals (Brown, 1975; Djieto-Lordon et al., 2001, Schmidt & Shattuck, 2014) whereas other species of this genus are predators. Until now, 38 extant species have been described in the genus Platythyrea. Nine species are recorded from the Neotropical region; 17 in the Ethiopian region; six species in the Australian region and nine species in the Oriental region (Bolton, 1995; Schmidt and Shattuck, 2014; Antwiki, 2016). From there, seven species (Platythyrea bidentata Brown, 1975; P. clypeata Forel, 1911; P. innermis Forel, 1910; P. nicobarensis Forel, 1905; P. parallela F. Smith, 1859; P. quadridenta Donisthorpe, 1941; P. tricuspidata Emery, 1900) have been recorded in Southeast Asia. Specimens of the ant genus Platythyrea are rarely collected because they have a relatively smaller colony size with on average usually a few hundred workers or fewer and Platythyrea workers typically forage individually (Ito, 2016). There are a few studies and publication of the ant genus Platythyrea in the world (Brown, 1975; Kugler, 1976; Lattke, 2003; Yoshimura & Fisher, 2007; Schmidt & Shattuck, 2014). Moreover, the last revision was conducted in 1975 by William L. Brown. Presently, there are no any taxonomic revision of the ant genus Platythyrea. Thus, a taxonomic revision of this genus are necessarily required. 2

In Thailand, Jaitrong and Nabhitabhata (2005) reported only three species: Platythyrea parallela, P. quadridenta, and P. tricuspidata from Thailand. In addition, according to previous reports on ant diversity from Thailand, there were many unnamed species of Platythyrea from Northern, Eastern and Southern Thailand (Prasityousi, 2001; Noon-anant, 2002; Wanichsakulpong, 2007; Sakchoowong Jaitrong & Ogata, 2009) . These lead to uncertain taxonomic status of the ant genus Platythyrea in

145923291 Thailand. So, I plan to examine all specimens of this genus deposited in museums of Thailand in order to describe these species and provide a key to known species and distribution patterns of the ant genus Platythyrea in Thailand. BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 1.2 Objectives of this study To examine the characteristics of external morphology and revise the ant genus Platythyrea in Thailand from worker caste specimens with related diagnosis and produce identification key to species.

1.3 Hypotheses This study investigates ant specimens of the genus Platythyrea to determine whether the characteristics of external morphology that can be used for species classification and identification. Additionally, it is likely to discover ant new species and new records from Thailand.

1.4 Contribution to knowledge To provide a greater insight into the presence of the ant genus Platythyrea in Thailand though a key to species, descriptions of known and new species, and distribution maps of all species.

1.5 Scope of the study To study externally morphological characteristics and distribution patterns of ants in the genus Platythyrea from Thailand. Ant specimens deposited in museums and newly collected from several localities in Thailand were examined. 3

CHAPTER 2 LITERATURE REVIEWS

2.1 Ants Ants belong to a single family, the Formicidae, in the order Hymenoptera (Hölldobler & Wilson, 1990; Bolton, 2003). In the World, a total of 17 extant 145923291 subfamilies, 403 extant genera and subgenera, 15,412 species and subspecies, 3 fossil families, 146 fossil genera, 795 fossil species and subspecies were recorded (Antwiki,

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 2018).

Classification of ants

Kingdom Animalia

Phylum Arthropoda

Class Insecta

Division Endopterygota

Order Hymenoptera

Family Formicidae

Ants are eusocial insects and play important roles in ecosystems. Some ant genera can spread plant seeds, for example, Rhytidoponera, Pachycondyla, Monomorium, and Iridomyrmex (Buckley, 1982). In addition, the ant genus Oceophylla can be used as a biological control agent in agricultural systems (Offenberg & Wiwatwitaya, 2010; Offenberg, Cuc & Wiwatwitaya, 2013). Ants have distinct external morphology from other insects in having third abdomen fused propodeum and second abdominal segment into a node-like petiole that petiole may be one or two nodes. The second nodes of petiole is called posterior petiole (Figure 2-1). 4

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 2- 1. External morphology of the ant (Source:https://en.wikipedia.org/wiki/Ant).

2.2 External morphology of the ant genus Platythyrea The redescription of external morphology of the ant genus Platythyrea (Figure 2-2) is largely modified from Brown (1975) and Schmidt & Shattuck (2014). Worker. Body slender. Head in full-face view rectangular or triangular, longer than broad, with sides weakly convex or parallel; posterior margin of head concave or straight; occipital corner almost straight or rounded; antenna short or long, antenna scape extending beyond or reaching posterolateral corner of head; clypeus broad or narrow, posterior margin insert between frontal lobes; in profile clypeus convex; lateral portion of clypeus broad; anterior margin convex; mandibles triangular with teeth or without teeth on masticatory margin (depending on species); basal margin of mandible without denticle; with apical tooth (depending on species); eyes relatively moderate to large and convex ( rarely small and flat such as Platythyrea clypeata) , maximum diameter greater than distance from anterior margin of eye to insertion of antenna; located anterior to head midline; ocelli absent; frontal lobes moderately large and 5

widely separated; frontal carinae closely spaced and strongly narrow posteriorly or widely spaced and weakly narrow posteriorly ( Brown, 1975; Schmidt & Shattuck, 2014). Mesosoma elongate, dorsum of mesosoma flat; metanotal groove obsolete; promesonotal suture distinct; propodeum with almost straight or weakly convex dorsal outline; propodeal junction rounded or obtusely angulated (depending on species);

145923291 declivity of propodeum shallowly concave; propodeal spiracle opening circular or elliptical; leg relative long (Brown, 1975; Schmidt & Shattuck, 2014). Petiole cylindrical and sessile; longer than high or almost as long as high, its BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 dorsal outline almost or convex; in dorsal view posterior margin with spine or without spine subpetiolar process well or weakly developed, in dorsal view but flat-topped rather than shallowly domed; petiole elongate with rounded node; node of petiole longer than wide; posterior lobes with one dorsal and two lateral poorly developed; petiole with distinctive flat-topped node; black with reddish appendages; sternopetiolar process well developed into lobe; posterior ventral surface of petiole well developed; third abdominal tergum much longer than postpetiole (Brown, 1975; Schmidt & Shattuck, 2014). Gaster and petiole separated by a distinct impression. Upper plate of the second segment of gaster weakly arched, second pretergite with stridulatory file (Brown, 1975; Schmidt & Shattuck, 2014). Outer surfaces of middle tibiae with hairs thin or lacking hairs; middle and hind tibiae with two comb-like (pectinate) spurs; tarsal claw with internal tooth. Dorsal surface of legs with fine sculpturation, some parts shiny; coarse puncturation small and shallow (Brown, 1975; Schmidt & Shattuck, 2014). Mandibles with fine erect and suberect hairs, body without erect or suberect hairs. Mandibles and dorsal surface of face finely punctate, sides of head with coarse punctures; pronotum finely granulated with coarse punctures on lateropronotum; mesopluron finely striated; propodeum, petiole, and postpetiole finely granulated with wavy striae on propodeum; gaster with wavy transverse striae with poorly defined coarse punctures laterally (Brown, 1975; Bolton, 2003). Queen. Characteristics very similar to worker but queen has winged, thoracic sclerites with corresponding modifications and with ocelli (sometimes absent, which is 6

a unique condition among alate ant queens) (Brown, 1975). Some queen ant species are ergatoid and completely absent in others (Molet & Peeters, 2006). Male. Characteristics very similar to worker but male has winged and with ocelli (Brown, 1975). The ant genus Platythyrea is most similar to the genus Probolomyrmex of the subfamily Proceratinae in having slender body. However, it is easily separated from

145923291 Probolomyrmex by the following characteristics: scapes longer than posterior margin of head in Platythyrea (shorter or just reaching posterior margin of head in Prolobomyrmex); compound eyes present (absent in Probolomyrmex); frontal lobes and BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 frontal carinae present (absent in Probolomyrmex); two metatibia spurs (one spur in probolomyrmex); clypeus uncovered mandibles (clypeus covering mandibles in Probolomyrmex).

Figure 2- 2. External morphology of the ant genus Platythyrea. A, body in profile view; B, head in full-face view; and C, body in dorsal view.

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2.3 Ant biology 2.3.1 Colony and reproduction Members of the genus Platyhtyrea were reported nesting in hollow branches or other preformed cavities in live or fallen trees (Brown, 1975; Djiéto-Lordon et al., 2001; Molet & Peeters, 2006; Yéo et al., 2006). The colony size equals a few hundred workers or fewer such as illustrated below in Table 2-1.

145923291 Platythyrea has reproductive strategies as ponerine genus (Villet, 1992; Molet & Peeters, 2006). Some species are gamergates such as Platythyrea conradti that have ergatoid queens, which interact aggressively with workers to form a dominance BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 hierarchy, but high-ranking workers do not reproduce unless the queen dies (Molet & Peeters, 2006). Some species have alate queens such as Platythyrea quadridenta, Platythyrea tricuspidata and Platythyrea arnoldi (Ito, 1994). Some species have gamergates only, for example, Platythyrea lamellosa, Platythyrea schultzei, and Platythyrea cribrinodis (Villet, 1990, 1991).

Table 2- 1. Colony size of the ant genus Platythyrea.

Scientific name Colony size Platythyrea conradti Emery, 1899 Between 100 to 500 workers ( Lévieux, 1976; Molet & Peeters, 2006; Yéo et al., 2006) Platythyrea lamellose (Roger, 1860) About 115 workers ( Villet et al., 1990) Platythyrea modesta Emery, 1899 Up to 50 workers (Djiéto-Lordon et al., 2001) Platythyrea parallela (F. Smith, 1859) About 50 workers (Wilson, 1959) Platythyrea punctate (F. Smith,1858) Between 23 to 51 workers (Hartmann et al., 2005) Platythyrea quadridenta Donisthorpe, 1941 About 19 workers (Ito, 1995) Platythyrea schultzei Forel, 1910 About 21 workers (Villet, 1991) Platythyrea tricuspidata Emery, 1900 About 21 workers (Ito, 1995)

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Villet (1990, 1991, 1992) examined the division of labor in colonies of Platythyrea cribrinodis, Platythyrea lamellosa and Platythyrea schultzei and found that these species typical age-related polyethism and workers of Platythyrea cribrinodis lay only inviable haploid eggs. Platythyrea punctata, this species has fascinating reproductive behavior. It has various alate queens, parthenogenetic apterous queens, gamergates, and parthenogenetic workers (Schilder, Heinze, Gross and Hölldobler,

145923291 1999; Hartmann, Wantia and Heinze, 2005). The presence of worker policing in such colonies is surprising, since they are virtually clonal and therefore lack any genetic conflict among colony members (Schilder et al., 1999; Hartmann, Wantia, Torres & BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Heinze, 2003). The reason for the look of reproductive conflict is that a reduced number of reproduction leads to increased colony productivity, favoring the maintenance of social control over reproduction (Hartmann et al., 2003).

2.3.2 Behavior Platythyrea workers are very fast runners. They have potent venomous stings, enabling them to rapidly catch prey (Brown, 1975; Djiéto-Lordon et al., 2001). Some species are generalist predators such as Platythyrea conradti, Platythyrea lamellose and Platythyrea modesta (Yéo et al., 2006; Molet & Peeters, 2006; Villet, 1990; Djiéto- Lordon et al., 2001). Research suggests that some species are located near termite mounds (Arnold, 1915; Brown, 1975) but some species prefer adult beetles such as Platythyrea arnoldi whose workers collect nectar on part of their body surface for transportation to the nest areas (Arnold, 1915; Déjean & Suzzoni, 1997). Lévieux (1983) completely confirmed Platythyrea species as a seed feeder. Platythyrea workers typically forage individually (Villet, 1990). Djiéto-Lordon et al. (2001) observed nestmate recruitment in Platythyrea modesta to aid in retrieval of large preys. Interestingly, workers of this species sometimes carry larvae directly to their prey and then bring the prey back to their nest, though this behavior is otherwise unknown within the Ponerinae. P. modesta conducts frequent emigrations to new nest sites with recruitment occurring via use of chemical trails. Ashikin and Hashim (2015) studied daily activity patterns of Platythyrea parallela in Ulu Gombak Forest Reserve, Peninsular Malaysia and found that P. parallela was ‘diurnal’ species after the 24 hours observation performed at two stable colonies. Most individuals of P. parallela were 9

more active during midday hours. Activity patterns were significantly different between daytime and nighttime. These patterns were likely to be related to the temperature regime.

2.4 Distribution The ant genus Platythyrea is pantropical (Figure 2-3). Some species also occur in subtropical regions of the New World, Africa, Asia, and Australia (Brown, 1975;

145923291 Antwiki, 2018). More information is provided in Table 2-2. BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 2- 3. World distribution map of the ant genus Platythyrea (Antwiki, 2018).

Table 2- 2. World distribution of the ant genus Platythyrea based on political regions.

Scientific name Distribution Platythyrea angusta Neotropical Region Brazil, Ecuador, French Guiana, Guyana, Peru, Suriname, Trinidad (type locality) and Tobago, Venezuela. Platythyrea arnoldi Afrotropical Region Democratic Republic of Congo, Mozambique, South Africa, Zimbabwe (type locality). 10

Platythyrea arthuri Afrotropical Region Comoros. Malagasy Region Madagascar (type locality), Mayotte. Platythyrea bicuspis Malagasy Region Madagascar (type locality).

145923291 Platythyrea bidentata Indo-Australian Region Philippines (type locality). Platythyrea brunnipes Australasian Region BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Australia (type locality). Platythyrea brunnipes Australasian Region Australia (type locality). Platythyrea clypeata Oriental Region Sri Lanka (type locality), Thailand and Vietnam. Palaearctic Region China. Platythyrea conradti Afrotropical Region Angola, Benin, Cameroun (type locality), Central African Republic, Democratic Republic of Congo, Equatorial, Guinea, Gabon, Ghana, Guinea, Ivory Coast, Nigeria. Platythyrea cooperi Afrotropical Region South Africa (type locality). Platythyrea cribrinodis Afrotropical Region Botswana, Kenya, Mozambique (type locality), Somalia, Sudan, Uganda, United Republic of Tanzania, Zambia, Zimbabwe. Palaearctic Region Ethiopia. 11

Platythyrea crucheti Afrotropical Region Angola (type locality). Platythyrea dentinodis Australasian Region Australia (type locality). Platythyrea exigua Neotropical Region Brazil (type locality), Paraguay.

145923291 Platythyrea frontalis Afrotropical Region Cameroun (type locality), Democratic Republic of Congo, Ghana, Ivory Coast, BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Uganda. Platythyrea gracillima Afrotropical Region Cameroon, Democratic Republic of Congo (type locality), Ghana, Kenya, Uganda. Platythyrea inermis Indo-Australian Region Philippines (type locality). Platythyrea janyai Oriental Region Thailand (type locality), Malaysia Platythyrea lamellosa Afrotropical Region Angola, Botswana, Democratic Republic of Congo, Namibia, South Africa (type locality), Zimbabwe. Platythyrea lenca Neotropical Region Honduras (type locality). Platythyrea matopoensis Afrotropical Region Zimbabwe (type locality). Platythyrea micans Australasian Region Australia (type locality). Platythyrea mocquerysi Afrotropical Region Comoros. Malagasy Region Madagascar (type locality), Mayotte. 12

Platythyrea modesta Afrotropical Region Cameroun (type locality), Central African Republic, Democratic Republic of Congo, Equatorial Guinea, Gabon, Ghana, Kenya, Mozambique, Nigeria, Saudi Arabia, South Africa, Sudan, Uganda.

145923291 Palaearctic Region Oman. Platythyrea nicobarensis Oriental Region BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 India (type locality). Platythyrea occidentalis Afrotropical Region Cameroon, Congo, Democratic Republic of Congo, Ghana, Senegal, Sierra Leone (type locality), Uganda. Platythyrea parallela Australasian Region Australia (type locality), New Caledonia. Indo-Australian Region Borneo, Fiji, Guam, Indonesia (type locality), Malaysia, Micronesia (Federated States of), New Guinea, Northern Mariana Islands, Philippines, Samoa, Singapore, Solomon Islands. Malagasy Region Seychelles. Oriental Region Bangladesh, India, Sri Lanka, Thailand, Vietnam. Platythyrea pilosula Neotropical Region Brazil (type locality), Colombia, Cost Rica, Paraguay, Venezuela. Platythyrea prizo Neotropical Region 13

Colombia, Costa Rica (type locality), Guatemala, Honduras, Mexico. Platythyrea punctata Nearctic Region United States (type locality). Neotropical Region Bahamas, Barbados, Brazil, Colombia,

145923291 Costa Rica, Cuba, Dominican Republic, French Guiana, Greater Antilles, Guatemala Guyana, Haiti, Honduras, BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Lesser Antilles, Mexico, Nicaragua, Panama, Puerto Rico, Saint Lucia, Suriname. Platythyrea quadridenta Indo-Australian Region Malaysia, New Guinea (type locality), Philippines. Oriental Region Thailand. Platythyrea sagei Oriental Region India (type locality). Platythyrea schultzei Afrotropical Region Angola, Democratic Republic of Congo, Kenya, Mozambique, Namibia (type locality), South Africa, Uganda, Zambia, Zimbabwe. Platythyrea sinuata Neotropical Region Bolivia, Brazil, Ecuador, French, Guiana, Guyana, Mexico, Peru, Suriname (type locality), Venezuela. Platythyrea strenua Neotropical Region Dominican Republic, Greater Antilles, Haiti (type locality). 14

Platythyrea tenuis Afrotropical Region Cameroun (type locality), Uganda. Platythyrea tricuspidata Indo-Australian Region Borneo, Indonesia (type locality), Malaysia. Oriental Region

145923291 Thailand. Platythyrea turneri Australasian Region Australia (type locality). BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Platythyrea viehmeyeri Afrotropical Region United Republic of Tanzania (type locality). Platythyrea zodion Neotropical Region Ecuador (type locality).

2.5 Historical reviews 2.5.1 Taxonomy Kugler (1976) studied taxonomy of Platythyrea from Costa Rica and describe a new species of Platythyrea based on eleven workers specimens deposited in the Field museum’s insect collections collected by Dr. John Wagner and Dr. John Kethley from berlesates at the La Selva Field Station of the Organization for Tropical Studies in Costa Rica. Lattke (2003) examined the ant genus Platythyrea in Dominican amber and described three new species: P. dentata, P. procera, and P. scalprum. Later, Andrade (2004) investigated Platythyrea from Dominican amber and extant species from Honduras and found two new species of Platythyrea: P. pumilio in Dominican amber and P. lenca from the recent Honduran fauna. Yoshimura and Fisher (2007) examined male ants of the Malagasy region (Madagascar, Mauritius, Reunion, Comores, and Seychelles), made treatment of the genera of Ponerinae and produced a key to subfamilies especially male-based keys to ant subfamilies and to the genera of Ponerinae in the Malagasy region. In conclusion, 15

seven subfamilies known from the Malagasy region in addition to an undetermined taxon are included in the subfamily key. Leyva (2013) studied taxonomy of the genus Platythyrea Roger, 1863 in New world region. Nine species were found: P. angusta, P. exigua, P. lenca, P. pilosula, P. prizo, P. punctata, P. sinuata, P. strenua and P. zodion. A new key, illustrations, diagnoses and distributions of Platythyrea were provided.

145923291 2.5.2 Colony composition Villet, Hart and Crewe (1990) investigated social organization of Platythyrea BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 lamellose and they found that P. lamellosa colonies typically contained 18-276 workers, however, queens were absent. Five to nine ovarioles per ovary were observed in dissected workers. It is worth noting that in each colony examined the only one individual of workers was inseminated and laid diploid reproductive eggs. In addition, adult males were present throughout the year but were very common during summer. Villet (1991) examined social differentiation and division of labour in Platythyrea schultzei. Their results revealed that P. schultzei have obligately queenless colonies with only 10-40 monomorphic workers. A single inseminated worker in each colony plays the important role of the queen. Colony labor of P. schultzei is subdivided into five groups of tasks which include care of eggs, care larvae, care cocoons, and general colony maintenance especially foraging. Schilder et al. (1999) showed that microsatellites discovering clonal structure of populations of Platythyrea punctata. According to their analysis of 314 workers selected from 51 colonies, intraspecific levels of variation at all loci, expressed both in the number of alleles and heterozygosities were detected. A clonal structure, with all individuals from all colonies examined was clearly exhibited in P. punctata population usually sharing the same genotype population. However, an additional genotype was observed in some workers from the only one colony in Puerto Rico. The predominance of thelytoky in P. punctata, together with genetic bottlenecks and founder effects were probably reflected by this low level of genotypic diversity. Brunner, Kellner and Heinze (2009) found that P. punctata colonies usually contain only single reproductive and other workers. If the old reproductive was removed from the colonies, some workers had activated ovaries instead. Later, Kellner 16

and Heinze (2011) confirmed that the facultative parthenogenesis was exhibited in P. punctata. In details, in many field colonies from the Caribbean Islands workers have identical multilocus genotypes and are thus probably clonal, however, the occurrence of males makes an ameiotic mechanism of thelytoky unlikely. Ito (2016) studied nesting and reproductive biology of Platythyrea sp. (parallela-group) in the Bogor Botanic Gardens, West Java, Indonesia. Based on his

145923291 work, this species, always nests in trees and reproduces through both dealated queens and mated workers. In colonies with a single dealated queen, the queen reproduced in only one colony among 13 collected colonies which also contained six mated workers, BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 of which three had active ovaries. In contrast, in the queenless colonies, the proportion of mated workers among the worker population was relatively higher.

2.5.3 Previous reports of Platythyrea in Thailand Prasityousi (2001) studied ant diversity and distribution in Doi Inthanon National Park, Chiang Mai province, northern Thailand. A total of 166 species belonging to 49 genera and 8 subfamilies were recorded. This study reported only one identified species of the ant genus Platythyrea, P. parallela, and also recorded one unidentified species, Platythyrea sp.1. Noon-anant (2002) investigated species diversity and seasonal abundance of ants in Bala Forest at Hala-Bala Wildlife Sanctuary, Narathiwat province, southern Thailand. A total of 255 species in 63 genera and 8 subfamilies were recorded. According to this study, five species of the ant genus Platythyrea including three identified species: P. paralella, P. quadridenta, and P. tricuspidata, and two unidentified species: Platythyrea. sp.1 and Platythyrea. sp.2 were reported. Thenthaworn (2004) investigated the relationship between air quality and ant distribution surrounding Ratchaburi Power Plant, Ratchaburi province, western Thailand. A total of 94 species were reported of which a single of the ant genus Platythyrea, Platythyrea parallela, was recorded from leaf litter and tree. Jaitong and Nabhitabhata (2005) generated the checklist of ant species known in Thailand. Two hundreds and forty-seven species distributed among 55 genera in nine subfamilies were recorded from Thailand. According to this checklist, three species of 17

the ant genus Platythyrea including P. paralella, P. quadridenta, and P. tricuspidata were reported. Watanasit, Tongjerm and Wiwatwitaya (2005) examined the composition of canopy ants at Ton Nga Chang Wildlife Sanctuary, Songkhla province, southern Thailand. A total of 118 morphospecies belonging to 29 genera and 6 subfamilies were identified. Two unidentified species of the ant genus Platythyrea were found.

145923291 Wanichsakulpong (2007) examined the ant species richness in Kasetsart University, Si Racha Campus, Chon Buri province, eastern Thailand. A total of 63 species in 35 genera and 8 subfamilies were reported. From this study, 5 species of the BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 genus Platythyrea, which include Platythyrea parellela, P. quadridenta, P. tricuspidata, Platythyrea sp.1 and Platythyrea sp.2, were found. Sakchoowong et al. (2008) compared the ground-dwelling ant diversity between natural forests and disturbed forests along a natural gas pipeline transect in Thong Pha Phum National Park, Kanchanaburi province, western Thailand. Platythyrea sp.1. was a single unidentified species of the ant genus Platythyrea recorded from the first site, KM5 and plot NF. Plapplueng (2009) compared the ant communities between two study sites at Khao Nan National Park, Nakhon Si Thammarat province, southern Thailand. A total of 228 species in 50 genera and 10 subfamilies were recorded. From this study, P. paralella was the only one species reported from Headquartes station. Jantarit, Wattanasit and Sotthibandhu (2009) studied canopy ants on the deciduous tree (Elateriospermum tapos) in a tropical rainforest at Khao Nan National Park, Nakhon Si Thammarat province, southern Thailand. A total of 123 morphospecies belonging to 31 genera and 5 subfamilies were found of which two unidentified species of the genus Platythyrea were found. Torchote, Sitthicharoenchai and Chaisuekul (2010) investigated species diversity and community composition of ant faunas in three different habitats including mixed deciduous forest, teak plantation, and fruit orchard at Thong Pha Phum district, Kanchanaburi province, western Thailand. There were sixty-two identified species and 67 morphospecies in 49 genera and 9 subfamilies. According to this study, only a single species of the ant genus Platythyrea, P. parallela, was recorded from the mixed deciduous forest. 18

CHAPTER 3 MATERIALS AND METHODS

3.1 Materials 1) 70% Ethyl alcohol 2) Insect pins (No.3)

145923291 3) Card points 4) Stereoscope (ZEISS Discovery.V12 stereoscope) 5) Nikon ECLIPSE E600 microscope

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 3.2 Methods 3.2.1 Study area This taxonomic study is focused in Thailand (Figure 3-1) and areas specific to southern and eastern Thailand where distribution of this genus was found to be most from previous study diversity of ant in Thailand. 3.2.2 Sources of materials This study is mainly based on the materials deposited in the collections at the Natural History Museum of the National Science Museum, Pathum Thani, Thailand (THNHM) and the Ant Museum, Faculty of Forestry, Kasetsart University, Bangkok, Thailand (AMK). Holotypes, paratypes and syntypes of name species were loaned from the following museums listed below. All specimens are dry-mounted on card triangle and deposited in the Natural History Museum of the National Science Museum, Thailand (THNHM).

19

Abbreviation of the type depositories are as follows: AMK Ant Museum, Faculty of Forestry, Kasetsart University, Bangkok, Thailand THNHM Natural History Museum of the National Science Museum, Bangkok, Thailand

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 3- 1. Map showing collection sites for Platythyrea specimens in Thailand.

20

3.2.3 Observations

Morphological observations of worker, male and queen specimens were made with a ZEISS Discovery.V12 stereoscope. Multi-focused montage images were produced using Axio Vision SE64 from a series of source images taken by a Digital AxioCamICc5 camera attached to a ZEISS Discovery.V12 stereoscope.

In addition to morphometric measurements, Scanning Electron Microscope 145923291 images of Platythyrea were made at Microscopic Center, Faculty of Science, Burapha University with a LEO 1450 VP scanning electron microscope on gold coated

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 specimens.

3.2.4 Terminology

Terms of relative position and general terminology of the worker ants follows Hölldobler and Wilson (1990) and Bolton (1995). For the important characters of the worker in the genus Platythyrea used, see Brown (1975), Bolton (2003) and Schmidt and Shattuck (2014). Queen and male characters of the genus, see Brown (1975) and Yoshimura and Fisher (2007).

A complete description of morphological features is given below (see Figures 3-2 to 3-5).

Mandible The appendages with which ants manipulate their environment.

Clypeus Anterior sclerite of the dorsal head, bounded posteriorly by the epistomal suture, which is very commonly referred to as the clypeal suture, or posterior clypeal margin.

Frontal carinae A pair of longitudinal cuticular ridges or flanges on the head, located dorsally behind the clypeus and between the antennal sockets. 21

Frontal lobes The only expression of the frontal carinae in some groups. Sometimes the portion of the torulus closest to the cephalic midline is also raised and expanded into a small, laterally projecting, lobe.

Propodeum The first abdominal segment (AI).

Propodeal declivity The sloping posterior surface of propodeum. 145923291

Mesonotum The tergite of the mesothorax.

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Mesopleuron Pleuron of the mesothorax is the largest pleurite.

Metapleuron Pleuron of the metathorax is located posteriorly on the side of the mesosoma, below the level of the propodeum.

Petiole: The second abdominal segment (AII).

Subpetiolar process: A ventral cuticular projection of the sternite of the petiole (AII), either below the node or on its anterior peduncle; sometimes absent but when present very variable in shape and size.

Punctate: Fine, impressed points or punctures appearing as pin- pricks.

Pruinose: Fine dust or coarse powder, but which cannot be rubbed off; the brightness of the surface somewhat obscured by the appearance of a bloom Like that of a plum.

Foveae: Depressions or pits.

22

A 145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

B

Figure 3- 2. Term of relative position for Platythyrea specimens: A, head in full-face view; and B, body in dorsal view. 23

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 3- 3. Whole body in profile view of Platythyrea.

Figure 3- 4. Mesosoma in profile view of Platythyrea.

24

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 3- 5. Triangular mandible of Platythyrea clypeata group.

3.2.5 Measurements and indices All worker specimens are measured according to Kugler (1976) and Andrade (2004). Worker measurements for the following parts are made using an ocular micrometer (accurate to 0.01 mm). The abbreviations used for the measurements and indices are as follows and illustrated in Figure 3-6:

TL Total length in profile, roughly measured from the anterior margin of head to the tip of gaster in stretched specimens.

HL Maximum head length in full-face view, measured from the anterior clypeal margin to the midpoint of a line drawn across the posterior margin of head.

HW Maximum head width in full-face view, measured above eyes.

SL Scape length excluding the basal of constriction and condylar bulb.

EL Eye length, maximum length of compound eyes in profile. 25

WL Weber's length, diagonal length of the mesosoma in profile from the anterior margin of pronotum to the posterior basal angle of the metapleuron, excluding neck.

PL Petiole length measured from the anterior margin of the peduncle to the posterior most point of tergite in profile.

PH Petiole height, maximum height of the petiole in profile view. 145923291

PW Petiole width, maximum width of the petiole in dorsal view.

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 CI Cephalic index. HW/HL x 100.

EI Eye index. EL/HW x 100.

SI Scape index. SL/HW x 100.

26

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 3- 6. Morphometric measurements. A, head in full-face view; B, antenna; C, body in profile view: D, petiole in profile view; and E, petiole in dorsal view. 27

CHAPTER 4 RESULTS

TAXONOMY

145923291 Platythyrea Roger, 1863

Platythyrea Roger, 1863: 172. Type-species: Pachycondyla punctata, by subsequent BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 designation of Bingham, 1903: 73. Eubothroponera Clark, 1930: 8. Type-species: Eubothroponera dentinodis, by original designation. Junior synonym of Platythyrea: Brown, 1975: 6.

Diagnosis of the genus (worker). For a more extensive description of the worker caste of the genus, see Schmidt and Shattuck (2014). Some of the important characteristics are reproduced with slight modifications. Workers approximately the same size (monomorphic). Body slender. Head in full-face view rectriangular, longer than broad, with sides weakly convex or parallel, posterior margin almost straight or concave; mandibles triangular, multidentate on masticatory margin (at least for Thai species), basal margin of mandible with or without a denticle; clypeus broad, posterior margin inserted between frontal lobes, anterior clypeal margin convex; frontal lobes widely separated; frontal carina relatively narrowly separated or very widely spaced; eye moderate to large and convex, rarely small and slightly flat (Platythyrea clypeata). Mesosoma in profile almost straight or weakly convex; metanotal groove usually obsolete; propodeal junction obtusely angulated; metapleural gland orifice opening laterally; middle and hind tibia with two pectinated spurs; tarsal claw with a single preapical tooth. Petiole cylindrical, usually clearly longer than broad, its posterodorsal margin in dorsal view convex or concave, without distinct spine or with 2-3 spines at each posterodorsal corner. Helcium usually projects from near midheight on the anterior face of abdorminal segment III. Body without erect and suberect mandible hairs; with erect and suberect hairs. 28

Checklist of the Thai Platythyrea species

Platythyrea clypeata Forel, 1911 = Platythyrea thwaitesi Donisthorpe, 1931 Platythyrea grandis sp. nov. Platythyrea itoi sp. nov. Platythyrea janyai Phengsi, Jaitrong, Ruangsittichai et Khachonpisitsak, 2018

145923291 Platythyrea parallela (F. Smith, 1859) = Platythyrea cephalotes Viehmeyer, 1924 = Platythyrea ceylonensis Donisthorpe, 1941 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 = Platythyrea coxalis Emery, 1893 = Platythyrea coxalis annamita Forel, 1911 = Platythyrea coxalis cylindrica Forel, 1913 = Platythyrea coxalis javana Forel, 1905 = Platythyrea coxalis philippinensis Viehmeyer, 1916 = Platythyrea coxalis tritschleri Forel, 1901 = Platythyrea inconspicua Mayr, 1870 = Platythyrea melancholica aruana Karavaiev, 1925 = Platythyrea parva Crawley, 1915 = Platythyrea pulchella Santschi, 1928 = Platythyrea pusilla Emery, 1893 = Platythyrea pusilla australis Forel, 1915 = Platythyrea pusilla egena Viehmeyer, 1916 = Platythyrea pusilla pacifica Santschi, 1928 = Platythyrea subtilis Emery, 1900 = Platythyrea wroughtoni sechellensis Forel, 1912 = Platythyrea wroughtoni victoriae Forel, 1900 = Platythyrea wroughtonii Forel, 1900 Platythyrea quadridenta Donisthorpe, 1941

Platythyrea tricuspidata Emery, 1900 = Platythyrea tricuspidata penangensis Wheeler, 1929

29

Key to Thai species based on worker caste

1 In dorsal view posterior margin of petiole with 2-3 distinct; spines, teeth or blunt angles (Figs 4-13)...... 2

- In dorsal view posterior margin of petiole round or feebly concave, without distinct spines, teeth or sharp angles (Figs 4-1)...... 4

145923291 2 Antennal scape long, extending beyond posterior margin of head (Fig 4-10); eye relatively smaller (EI  25). ……………………...... 3

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 - Antennal scape relatively short, reaching posterior margin of head (Fig 4-6); eye relatively larger (EI  36)...... P. itoi sp.nov.

3 In dorsal view posterior margin of petiole with 3 distinct spines (Fig.); petiole longer than high...... P. tricuspidata

- In dorsal view posterior margin of petiole clearly concave with distinct lateral blunt angles; petiole almost as long as high...... P. quadridenta

4 Frontal carinae closely spaced or strongly narrowed posteriorly (Fig. 4-12). ………………………………………………..……………..………………. 5

- Frontal carinae widely spaced or weakly narrow posteriorly (Fig. 4-12)...... P. paralella complex

5 Eyes relatively smaller (EL = 0.16 mm; EI  15), with 5-7 ommatidia on longest axis...... 6

- Eyes relatively larger (EL = 0.20 mm; EI  18), with 9 ommatidia on longest axis...... P. janyai

6 Lateral face of head and pronotum finely micropunctate (Fig. 4-5); antennal scape relatively long (SI = 153), by 1/3 of its length extending beyond posterior margin of head. …...……….…………………....…….…..... P. grandis sp. nov.

- Lateral face of head and pronotum punctate with dense foveae (Fig. 4-4); antennal scape relatively short (SI = 125-138), by 1/4 of its length extending beyond posterior margin of head. ………..………………...…...... P. clypeata 30

Platythyrea clypeata Forel, 1911 (Figs. 4-1, 4-2, 4-3, 4-4A1-A3)

Platythyrea clypeata Forel, 1911: 378; Brown, 1975: 50; Bolton, 1995: 336; Xu and Zeng, 2000: 214; Schmidt and Shattuck, 2014: 51. Senior synonym of P. thwaitesi: Brown, 1975: 8.

145923291 Platythyrea thwaitesi Donisthorpe, 1931: 496. Junior synonym of P. clypeata: Brown, 1975: 8.

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Type. The syntype queen from “Pays de Mois" (S.E. Asia)”. The syntype is deposited in MHNG.

Non-type material examined. Nine workers from E. Thailand, Chachoengsao Prov., Tha Takiab Dist., Kaho Ang Reu Nai Wildlife Sanctuary, 27.IX.2002, W. Jaitrong leg., WJT270902-1 (THNHM-I-02423 to THNHM-I-02431, THNHM); 3 workers from same loc., date and collector, WJT270902-1 (THNHM-I-02432 to THNHM-I-02434, THNHM); 6 workers from E. Thailand, Sa Kaeo Prov., Khao Ang Reu Nei Wildlife Sanctuary, 26.VI.2003, W. Jaitrong leg., WJT03-TH-228 (THNHM-I-02435 to THNHM-I-02440, THNHM); 23 workers and 1 male from E. Thailand, Chanthaburi Prov., Soi Dao Dist., 14.V.2008, W. Jaitrong leg., WJT08-E065 (THNHM-I-02441 to THNHM-I-02453, THNHM); 10 workers and 1 delate queen from Laos, Vientiane, Pak Ngum Dist., Ban Phang Dang, 14.VI.2010, W. Jaitrong leg., WJT-LAO-143 (THNHM); 1 delate queen from same loc. and collector, 12.VI.2010 (THNHM).

Redescription of worker. Measurements and indices (n = 37), TL 5.74-6.20, HL 1.29- 1.39, HW 0.86-0.89, SL 1.12-1.18, EL 0.10, WL 1.85-2.05, PL 0.66-0.73, PH 0.46- 0.53, PW 0.40-0.43, CI 61-69, EI 11, SI 125-138. Head in full-face view rectangular, clearly longer than broad, with sides weakly convex or almost parallel, occipital corner roundly convex, and posterior margin feebly concave; antennal scape slender, relatively shorts, lightly extending beyond posterolateral corner of head (by 1/4 of its length); clypeus narrow, in profile with median portion distinctly convex, in full-face view lateral portion relatively broad and 31

anterior margin clearly convex; mandible triangular, its masticatory margin with a large apical tooth, followed by 10 smaller teeth (including basal tooth), large and smaller teeth alternating, but the series as a whole decreasing in size; basal margin of mandible without denticle; eye flat, located laterally anterior to mid-length of head, very small 0.10 mm in maximum diameter, with 5 omatidia along the longest axis, distance between mandibular base and anterior margin of eye about 3 times as long as maximum

145923291 eye length; distance between posterior margin of eye and occipital corner of head in lateral view about 7.3 times as long as distance between mandibular base and anterior margin of eye; frontal lobes close to each other and rounded; frontal carinae strongly BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 narrow posteriorly. Mesosoma elongate, in profile almost straight dorsal outline; promesonotal suture distinct; mesopleuron demarcated from mesonotum and metapleuron by shallow furrow; propodeum with almost straight dorsal outline; propodeal junction obtusely angulated; declivity of propodeum shallowly concave; seen from back propodeal declivity rounded above; legs relatively long; propodeal spiracle opening elliptical. Petiole cylindrical and sessile, clearly longer than high and longer than broad, its dorsal outline almost straight; in profile node of petiole inclined backward; posterodorsal corner formed an acute angle; declivity of petiole deeply concave; in dorsal view node of petiole rectangular, slightly narrower posteriorly, its posterior margin convex and shallow median concave; subpetiolar process developed, subtriangular located anteroventrally, its apex truncate and pointed forward; ventral outline of petiole almost straight. Sculpture: dorsum of head finely punctate; lateral face of head areas behind, above and below eye punctate with dense foveae; dorsa of mesosoma finely micropunctate similar to that of dorsum of head; lateral faces of pronotum, metapleuron and propodeum punctate with sparse shallow foveae; petiole finely micropunctate; gastral gergites I and II finely reticulate; antennal scape with finely micropunctate; coxae microreticulate with smooth and shiny interspaces. Setae: present on tip of gaster; pubescence white, very short and fine. Color: head, mesosoma, petiole and gaster reddish brown to dark brown (tip of gaster yellowish); antenna and leg yellowish brown to reddish brown (funicular paler than scape).

32

Description of dealated queen. Measurements and indices (n = 2), TL 7.49, HL 1.52, HW 1.06, SL 1.32, EL 0.20, WL 2.31, PL 0.79, PH 0.73, PW 0.46, CI 69, EI 18, SI 125. Body size slightly larger than worker. Head in full-face view rectangular, clearly long than broad with convex sides and almost straight posterior margin, occipital corner roundly convex; antenna scape 1/4 of its length extending beyond posterolateral

145923291 corner of head; eye very large and convex, located anterior to mid-length of head, 0.10 mm in maximum diameter with ca. xxx ommatidia on the longest axis; frontal lobe and frontal carinae similar to that of in worker caste; distance between anterior margin of BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 eye and mandibular base almost as long as eye length; ocelli absent. Mesosoma in profile slightly convex dorsal outline; pronotum long and broad; mesoscutum trapezoidal, anterior edge clearly convex in dorsal view, separated from mesoscutellum by a shallow but wide suture and from pronotum by narrow suture; parapsidal lines indistinct, relatively long, straight and running anteriorly to mid-length of mesoscutum; mesoscutellum almost as long as broad; metanotum very short separated from mesoscutellum and propodeum by deep grooves; propodeum relatively long; mesopleuron broad, anepisternum not demarcated from katepisternum; propodeal junction obtusely angulated; declivity of propodeum shallowly concave; seen from back propodeal declivity rounded above. In profile view Petiole relatively short, rhombus, almost as long as broad, its anterior margin weakly convex while posterior margin concave; declivity of petiole shallow concave; subpetiolar process low and subtriangular located anteroventrally, its apex pointed forward; ventral outline of petiole almost straight. Gaster relatively larger than in the worker. Sculpture: setae and color condition similar to that of in the worker caste.

Description of male. Measurements and indices, TL 6.14, HL 1.06, HW 0.92, SL 0.30, EL 0.53, WL 1.88, PL 0.63, PH 0.50, PW 0.40, CI 87, EI 57, SI 32 Body size same as worker. Head in full-face view elliptical, slightly long than broad with convex sides and convex posterior margin; antenna with 13 segments; antennal scape short, shorter than each of III-XIII; II shortest, about 0.3 times as long as antennal scape; eye large, anterior margin of eye convex and located close to mandibular base, ventral and posterior margins convex, while dorsal margin feebly 33

concave; eye diameter 0.53 mm in maximum axis; clypeus broad, its anterior margin sharply convex, posterior margin not inserted between frontal lobe; frontal lobes narrow not cover antennal socket; frontal carinae short reaching mid-length of eye; distance between anterior margin of eye and mandibular base clearly shorter than eye length; three ocelli located on vertex. Mesosoma rather stout; pronotum in dorsal view long and broad, its posterior margin strongly concave; mesoscutum trapezoidal, relatively

145923291 longer than broad, anterior margin clearly convex in dorsal view, separated from mesoscutellum by deep groove and separated from pronotum by transverse suture; parapsidal lines narrow, relatively short, straight and running anteriorly to mid-length BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 of mesoscutum; metanotum very short separated from mesoscutellum and propodeum by shallow groove; propodeum relatively long; mesopleuron large, clearly demarcated by metapleuron by deep and broad suture, anepisternum clearly demarcated from katepisternum by a suture; metapleuron relatively narrow comparing with mesopleuron and demarcated from lateral face of propodeum by shallow suture; propodeal junction rounded; declivity of propodeum flat, seen from back propodeal declivity rounded above. In profile view Petiole rhombus, relatively short, slightly shorter than high, its dorsal outline straight, anterior face sloping down to peduncle, while its posterior margin feebly concave; subpetiolar process low and subtriangular located anteroventrally, its apex pointed forward; ventral outline of petiole feebly concave. Gaster relatively larger than in the worker. Sculpture: setae and color condition similar to that of in the worker caste.

Distribution. China, Laos, Sri Lanka (type locality), Thailand (Chachoengsao, Sa Kaeo and Chantaburi provinces) and Vietnam and also see Figure 4-14 and Table 4-1.

Comparative notes. P. clypeata is similar to P. gracillima Wheeler, 1922, P. janyai and P. prizo Kuger, 1977 in general appearance as they share the following characteristics: body reddish brown; frontal lobe narrow; frontal carinae closely spaced and strongly narrowed posteriorly; mandible triangular, its masticatory margin with a large apical tooth, followed by 9-10 smaller teeth, large and smaller teeth alternating; propodeal spiracle opening elliptical; in dorsal view posterior margin of petiole convex 34

without spines. However, P. clypeata can be easily separated from P. gracillama by the following characteristics: body size smaller (TL = 5.74-6.20 in P. clypeata; TL = 9 mm in P. gracillima); eye smaller and flat (0.1 mm in P. clypeata; 0.3 mm and convex in P. gracillima); clypeus narrow and rather convex (broad and rather flat in P. gracillima); propodeal junction obtusely angulated; declivity of propodeum shallowly concave (propodeum junction rounded, declivity of propodeum flat in P. gracillima);

145923291 seen from back propodeal declivity rounded above (propodeal declivity concave above in P. gracillima); petiole laterally swollen (petiole laterally compressed in P. gracillima); head finely punctate (head rather smooth in P. gracillima). BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 P. clypeata can be distinguished from P. prizo by the following characteristics: head shorter (CI = 61-69 in P. clypeata; CI = 70-72 in P. prizo); eye flat and small (0.1 mm in P. clypeata; eye very large, 0.31 mm P. prizo); eye without erect pubescence (eye covered with extremely fine short erect pubescence in P. prizo) ; propodeum junction obtusely angulated (propodeum junction armed with a pair of short teeth or tubercles in P. prizo); in profile, posterodorsal corner of petiole formed an acute angle (roundly convex in P. prizo). Lastly, this species is separated from P. janyai by the following characteristics: head relatively longer (CI 61-69 in P. clypeata; CI 72-74 in P. janyai); eye clearly smaller (EL 0.10 mm with 5 ommatidia on longest axis in P. clypeata; EL 0.20 mm with 11 ommatidia on longest axis in P. janyai); eye flat (convex in P. janyai); finely punctate with dense shallow foveae (dorsum and lateral face of head finely micropunctate without foveae in P. janyai); in profile petiole slightly longer than high and in dorsal view node of petiole slightly narrower posteriorly (clearly longer than high and in dorsal view node of petiole anteriorly as broad as posteriorly in P. janyai); ventral outline of petiole feebly concave (weakly convex in P. janyai) (see fig. 4-4 for comparison). P. clypeata is distinctly allopatric with P. janyai in distribution. It was recorded from Sri Lanka, Vietnam, Laos and E Thailand (Brown 1975). On the other hand, P. janyai occurs in Malay Peninsula (South Thailand and West Malaysia). 35

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 1. Worker of Platythyrea clypeata. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 36

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 2. Queen of Platythyrea clypeata. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 37

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 3. Male of Platythyrea clypeata. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 38

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 4. SEM images of Platythyrea clypeata (A1–A3) and P. janyai (B1–B3). A1, B1 Sculpture on lateral face of head; A2, B2 Ommatidia of eye; A3, B3 Petiole in profile view.

39

Platythyrea grandis sp. nov. (Fig. 4-5)

Type. Holotype: worker from N. Thailand, Chiang Mai Prov., Muang Dist., Restored Forest, 8.V.2002, S. Sonthichai leg., THNHM-I-02463 (THNHM). Paratype: one worker from W. Thailand, Kanchanaburi Prov., Thong Pha Phum N.P., Natural Forest,

145923291 8.III.2005, W. Sakchooeong leg., THNHM-I02454 (THNHM).

Description of worker (holotype and paratype). Measurements and indices, holotype: BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 TL 8.08 mm, HL 1.66 mm, HW 1.09 mm, SL 1.881 mm, EL 0.16 mm, WL 2.64 mm, PL 0.92 mm, PH 0.66 mm, PW 0.46 mm, CI 66, EI 15, SI 172. Measurements and indices, paratypes: TL 7.59 mm, HL 1.65 mm, HW 1.09 mm, SL 1.88 mm, EL 0.16 mm, WL 2.64 mm, PL 0.92 mm, PH 0.66 mm, PW 0.46 mm, CI 64, EI 15, SI 153. Head in full-face view subrectangular, clearly longer than broad, with sides almost parallel, occipital corner convex and posterior margin almost straight, slightly narrower posteriorly; antennal scape relatively long, extending beyond posterolateral corner of head (1/3 of its length); clypeus broad, posterior margin inserted between frontal lobes; in profile clypeus roundly convex; lateral portion of clypeus slightly broad; anterior margin roundly convex; mandible triangular, its masticatory margin with a large apical tooth, followed by 8 smaller teeth, large and smaller teeth alternating, but the series as a whole decreasing in size toward the basal tooth; basal margin of mandible without denticle; eye flat, located laterally, anteriorly 1/3 of head length, relatively small, 0.17 mm in maximum diameter, with 7 omatidia along the longest axis, distance between mandibular base and anterior margin of eye about 0.4 times as long as maximum eye length; distance between posterior margin of eye and occipital corner of head in profile about 2.6 times as long as distance between mandibular base and anterior margin of eye; frontal lobes close to each other and rounded; frontal carinae strongly narrow posteriorly. Mesosoma elongate, in profile almost straight dorsal outline; promesonotal suture distinct; mesopleuron not demarcated from mesonotum, but demarcated from metapleuron by a broad but shallow furrow; metapleuron not demarcated from lateral face of propodeum; propodeum almost straight dorsal outline; propodeal junction rounded; declivity of propodeum shallowly concave; seen from 40

back propodeal declivity rounded above; legs very long; propodeal spiracle opening elliptical. Petiole cylindrical and sessile, clearly longer than high, its dorsal outline weakly convex; in profile posterodorsal corner of petiole with acute angle overhanging declivity of petiole; declivity of petiole shallowly concave; in dorsal view node of petiole rectangular, its posterior margin convex; subpetiolar process low and subtriangular, located anteroventrally, its apex truncate and pointed forward; ventral

145923291 outline of petiole almost straight. Sculpture: head entirely finely micropunctate; dorsum of mesosoma finely micropunctate, mesopluron, metapleuron and lateral face of propodeum punctate; BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 petiole finely micropunctate; gaster superficially shagreened; coxae and femora superficially reticulate, but smooth and shiny interspaces, tibiae micropunctate. Setae: absent; pubescence white, very short and fine, distributed over whole body and appendages, longer and more oblique on anterior clypeal margin, tip of mandible and hypopygium. Color: body reddish brown; tip of gaster, antenna and legs yellowish brown or paler than body.

Distribution. Thailand (Chiang Mai and Kanchanaburi provinces) and also see Figure 4-14 and Table 4-1.

Etymology. The specific name is a Latin referring to the large body size of the species.

Comparative notes. Platythyrea grandis sp. nov. is similar to P. clypeata, P. gracillima and P. janyai in general appearance as they share the following characteristics: body reddish brown; frontal lobe narrow; frontal carinae closely spaced and strongly narrowed posteriorly; mandible triangular, its masticatory margin with a large apical tooth, followed by 9-10 smaller teeth, large and smaller teeth alternating; propodeal spiracle opening elliptical; in dorsal view posterior margin of petiole convex without spines. However, P. grandis can be easily separated from P. clypeata by the following characteristics: head in full-face view, posterior margin weakly concave (posterior margin almost straight in P. clypeata); antennal scape relatively long, 1/3 of its length extending beyond posterolateral corner of head (antennal scape short, 1 /4 of 41

its length extending beyond posterolateral corner of head in P. clypeata); clypeus broad (clypeus narrow in P. clypeata); eye larger (EI = 15, with 7 ommatidia in P. grandis; EI = 11, with 5 ommatidia in P. clypeata); mesosoma relative longer (WL = 2.64 in P. grandis, WL=1.85-2.05 in P. clypeata); mesopleuron not demarcated from mesonotum (mesopleuron clearly demarcated from mesonotum by shallow furrow in P. clypeata); in profile view, propodeal junction roundly convex (propodeal junction obtusely

145923291 angulated in P. clypeata); in dorsal view, posterior margin of petiole clearly convex (posterior margin of petiole convex with shallow median concave in P. clypeata) ; lateral face of head entirely micropunctate (lateral face of head areas behind, above and BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 below eye punctate with dense foveae in P. clypeata); gaster superficially shagreened finely reticulate in P. clypeata). P. grandis can be distinguished from P. gracillima by the following characteristics: clypeus roundly convex (rather flat in P. gracillima); petiole laterally convex; seen from above longer than high (petiole laterally compressed; seen from above a little more than twice as long as broad in P. gracillima) ; mandible finely micropunctate (mandible finely and densely punctate in P. gracillima); head entirely finely micropunctate (rather smooth in P. gracillima). P. grandis can be separated from P. janyai by the following characteristics: head longer (CI = 66 in P. grandis, CI = 72 in P. janyai) ; antennal scape relatively long, 1/3 of its length extending beyond posterolateral corner of head (clearly extending beyond posterolateral corner of head in P. janyai); eye smaller and flat (EL = 0.17 mm in P. grandis, EL = 0.20 mm and slightly convex in P. janyai) ; seen from back propodeal declivity rounded above ( tapering above in P. janyai) ; dorsal outline of petiole weakly convex ( almost straight in P. janyai). 42

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 5. Worker of Platythyrea grandis. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 43

Platythyrea itoi sp. nov. (Fig. 4-6)

Types. Holotype: worker (FI130914-1), N. Thailand, Chiang Mai Prov., Chiang Mai University, Main Campus, 3.IX.2014, F. Ito leg., FI30914-1 (THNHM). Paratypes: two workers, N. Thailand, Chiang Mai Prov., Chiang Mai University, Main Campus,

145923291 25.VI.2013, Y. Onishi leg. (specimen numbers 19-T and 20-T, THNHM); one worker, N. Thailand, Chiang Mai Prov., nr Faculty of Agriculture, Chiang Mai University (CMU), 26.XI2012, Y. Onishi, H. Kitori and F. Ito leg., YO261112-1 (THNHM). BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Description of worker (holotype and paratypes). Measurements and indices, holotype: TL 5.68 mm, HL 0.99 mm, HW 0.85 mm, SL 0.66 mm, EL 0.33 mm, WL 1.82 mm, PL 0.59 mm, PH 0.59 mm, PW 0.59 mm, CI 86, EI 38, SI 76. Measurements and indices, paratypes (n = 3): TL 5.01-5.84 mm, HL 0.89-1.05 mm, HW 0.79-0.89 mm, SL 0.62-0.72 mm, EL 0.26-0.33 mm, WL 1.58-1.84 mm, PL 0.50-0.59 mm, PH 0.49- 0.66 mm, PW 0.46-0.53 mm, CI 81-86, EI 30-37, SI 77-84. Head in full-face view rectangular, clearly longer than broad, slightly broader posteriorly, with sides weakly convex, posterior margin concave; antenna scape short; reaching posterolateral corner of head; antennal segment II slightly longer each of III- IX but narrower; clypeus broad; posterior margin ill-defined; anterior clypeal margin strongly convex; in profile median portion of clypeus convex; mandible triangular, with sides weakly concave, its masticatory margin with a large apical tooth, followed by 13- 15 denticle; but the series as a whole decreasing in size toward the basal tooth; basal margin lacking denticle close to basal tooth; eye relatively large and convex, located laterally middle of head, 0.33 mm in maximum diameter, with 27 omatidia along the longest axis, distance between mandibular base and anterior margin of eye about 0.5 times as long as maximum eye length; frontal lobes wide and rounded; area between frontal lobes largely impressed; frontal carinae widely spaced and weakly narrow posteriorly. Mesosoma elongate, in profile almost straight dorsal outline; promesonotal suture distinct; mesonotum separated from propodeal dorsum by shallow impression; mesopleuron narrow demarcated from mesonotum and metapleuron by a shallow groove; metapleuron not demarcated from lateral faces of propodeum; propodeum 44

feebly convex dorsal outline; propodeal junction obtusely angulated, shorter than broad at base; declivity of propodeum shallowly concave; legs relatively short; propodeal spiracle opening circular. Petiole cylindrical and sessile, almost as long as high, its dorsal outline almost straight; in profile posterodorsal corner of petiole blunt angle; declivity of petiole concave; in dorsal view petiole subreticular, its posterior margin concave medially, forming blunt lateral angles; subpetiolar process developed,

145923291 subtriangular, located anteroventrally; its apex pointed forward; ventral outline of petiole concave. Sculpture: dorsa of head, pronotum, mesonotum, propodeum, petiole, and first BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 gastral tergite finely micropunctate and opaque; lateral faces of head, pronotum, mesopleuron, metapleuron, and propodeum, petiole, and first gastral tergite micropunctate with dense foveae; second gastral tergite superficially microreticulate with smooth and shiny interspaces; antennal scape finely micropunctate; coxa reticulate with smooth and shiny interspaces; femur superficially shagreened with smooth and shiny interspaces. Setae: present on tip of gaster; Body covered with very short grey pubescence. Color: head, mesosoma, petiole, and gaster dark to dark brown; antenna scape yellowish brown, while funicular dark brown; madible and legs dark reddish brown.

Distribution. Thailand (Chiang Mai province) and also see Figure 4-14 and Table 4-1.

Etymology. The specific name is dedicated to Professor Dr. Fumonori Ito of the Faculty of Agriculture, Kagawa University, Japan who donated me the holotype.

Comparative notes. Platythyrea itoi sp. nov. is similar to P. paralella in general appearance as they share the following characteristics: body darkness; frontal lobe broad; frontal carinae widely spaced and weakly narrow posteriorly; propodeal spiracle opening circular; in dorsal view, posterior margin of petiole weakly concave or almost straight. However, P. itoi can be easily separated from P. paralella by the following characteristics: eye larger (EI = 0.33 mm, with 27 omatidia along the longest axis in P. itoi; EL = 0.16-0.26 mm, with 12 omatidia along the longest axis in P. paralella); masticatory margin of mandible with a large apical tooth, followed by 13-15 smaller 45

teeth (masticatory margin of mandible with a large apical tooth, followed by 9-11 smaller teeth in P. paralella); propodeum feebly convex dorsal outline (almost straight dorsal outline in P. paralella); propodeal junction obtusely angulated (propodeal junction bluntly angulated in P. paralella); in profile, posterodorsal corner of petiole blunt angle (posterodorsal corner of petiole with very short, obtusely angulated in P. paralella); in dorsal view, petiole subreticular, its posterior margin concave medially,

145923291 forming blunt lateral angles (subrectangular, broadened posteriorly, its posterior margin straight or with shallow median concave in P. paralella); dorsa of head, mesosoma and petiole finely micropunctate and opaquae (dorsa of head, mesosoma and petiole finely BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 punctate with sparse shallow foveae in P. paralella). 46

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 6. Worker of Platythyrea itoi. A, body in profile view; B, head in full-face view; and C, body in dorsal view. 47

Platythyrea janyai Phengsi, Jaitrong, Ruangsittichai et Khachonpisitsak 2018 (Figs.4-4B1-B3, 4-7)

Platythyrea janyai Phengsi, Jaitrong, Ruangsittichai and Khachonpisitsak, 2018: 89, figs. 1, 5B1-B3.

145923291 Type. Holotype: worker, S. Thailand, Phattalung Prov., Sri Banpot Dist., Riang Tong Waterfall, Khao Pu Khao Ya National Park, 28.IX.2007, W. Jaitrong leg., WJT07-TH- 2060 (THNHM-I-02392, THNHM). Paratypes: three workers, same data as holotype BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 (THNHM-I-02393 to THNHM-I-02395, THNHM).

Non-type material examined. Two workers, S. Thailand, Trang Prov., Na Yong Dist., Khao Chong Botanical Garden, 7.XI.2014, W. Jaitrong leg., WJT071114-2 (THNHM- I-02421 to THNHM-I-02422, THNHM).

Description of worker. Measurements and indices, holotype: TL 6.63, HL 1.42, HW 1.06, SL 1.39, EL 0.20, WL 2.21, PL 0.73, PH 0.53, PW 0.40,y CI 74, EI 18, SI 131. Measurements and indices, paratypes (n = 3): TL 6.67-6.96, HL 1.45, HW 1.06, SL 1.42, EL 0.20, WL 2.31, PL 0.79, PH 0.53, PW 0.40, CI 72, EI 18, SI 134. Head in full-face view subrectangular, clearly longer than broad, with sides weakly convex, occipital corner round, and posterior margin almost straight; antenna relatively long; scape slender, clearly extending beyond posterolateral corner of head; antennal segment II narrow, 1.6 times as long as segment III; III longer than each of segments IV–XII; clypeus broad, in profile with median portion distinctly convex, in full-face view lateral portion narrow and anterior margin clearly convex; mandible triangular, masticatory margin with a large apical tooth, followed by 9–10 smaller teeth, larger and smaller teeth alternating, but the series as a whole decreasing in size toward basal tooth; basal margin of mandible without denticle; eye slightly convex, located laterally anterior to mid-length of head, relatively large, 0.20 mm in maximum diameter, with eleven ommatidia on longest axis, distance between mandibular base and anterior margin of eye 1.5 times as long as maximum eye length; with head in profile, distance between posterior margin of eye and occipital corner of head 3.4 times 48

as long as distance between mandibular base and anterior margin of eye; frontal lobes relatively close to each other, with roundly convex lateral margins; antennal socket horizontal, in plane of transverse axis of head, and in dorsal view, half concealed by frontal lobe. Mesosoma elongate, in profile with weakly convex dorsal outline; promesonotal suture distinct; metanotal groove absent; mesopleuron not clearly demarcated from mesonotum, but can be separated from metapleuron by a shallow

145923291 furrow; metapleuron not demarcated from lateral face of propodeum; propodeum with almost straight dorsal outline; propodeal junction rounded; declivity of propodeum shallowly concave; seen from back propodeal declivity tapering above; propodeal BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 spiracle opening elliptical; legs very long. Petiole cylindrical and sessile, clearly longer than high and broad, its dorsal outline almost straight; with petiole in profile posterodorsal corner with acute angles overhanging declivity of petiole; declivity of petiole shallowly concave; in dorsal view petiole rectangular, its posterior margin concave medially; subpetiolar process weakly developed, subtriangular, located anteroventrally; ventral outline of petiole weakly convex. Sculpture: Head (including antennal scape), mesosoma, petiole and gaster finely and densely micropunctate; coxae and femora superficially reticulate but shiny. Setae: pubescence white, very short and fine, distributed over whole body and appendages, longer and more oblique on anterior clypeal margin, tip of mandible and hypopygium. Color: Dorsum of head dark brown, while lateral face of head reddish brown; mesosoma, petiole and gaster dark brown to reddish brown (tip of gaster yellowish); antenna and legs yellowish brown (funicular segments paler than scape).

Distribution. Thailand (Pattalung and Trang provinces) and Malaysia (Ulu Gombak National Park) and also see Figure 4-14 and Table 4-1.

Comparative notes. The new species is similar to P. clypeata, P. grandis and P. gracillima in general appearance as they share the following characteristics: body reddish brown; frontal lobe narrow; frontal carinae closely spaced and strongly narrowed posteriorly; mandible triangular, its masticatory margin with a large apical tooth, followed by 9-10 smaller teeth, large and smaller teeth alternating; propodeal 49

spiracle opening elliptical; in dorsal view posterior margin of petiole convex without spines. However, P. janyai can be easily separated from P. gracillama by the following characteristics: body size smaller ( TL = 6.63 mm in P. janyai, TL = 9 mm in P. graciilima); eye relatively smaller (EL = 0.2 mm in P. janyai, EL = 0.3 mm in P. graciilima); seen from back, propodeal declivity tapering above (propodeal declivity weakly concave in P. gracillima); petiole laterally convex, seen from above longer

145923291 than high (petiole laterally compressed, seen from above a little more than twice as long as broad in P. gracillima) . Differentiate between P. janyai and P. clypeata, see “Comparative notes” under P. clypeata and between P. janyai and P. grandis, see BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 “Comparative notes” under P. grandis.

50

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 7. Worker of Platythyrea janyai. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 51

Platythyrea paralella (F. Smith, 1859) (Figs. 4-8, 4-9)

Ponera parallela F. Smith, 1859: 143. Combination in Platythyrea: Donisthorpe, 1932: 454. Platythyrea paralella: Donisthorpe, 1943: 434; Brown, 1975: 47; Wheeler, G.C. &

145923291 Wheeler, J. 1976: 42; Bolton, 1995: 337; Jaitrong and Nabhitabhata, 37; Schmidt and Shattuck, 2014: 52. Platythyrea inconspicua Mayr, 1870: 961; Emery, 1890: 56. Junior synonym of P. BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 punctata: Forel, 1901: 335; Creighton, 1950: 34; of P. parallela: Brown, 1975: 8. Platythyrea coxalis Emery, 1893e: 189; Viehmeyer, 1916: 111. Junior synonym of P. parallela: Wilson, 1958: 151. Platythyrea pusilla Emery, 1893: 188; Wheeler, W.M. 1919: 50. Junior synonym of P. parallela: Wilson, 1958: 151. Platythyrea subtilis Emery, 1900: 666. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea wroughtoni r. victoriae Forel, 1900: 316; Bingham, 1903: 75. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea wroughtonii Forel, 1900: 315. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea coxalis var. tritschleri Forel, 1901: 338. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea coxalis var. javana Forel, 1905: 5. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea coxalis var. annamita Forel, 1911: 215. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea wroughtoni subsp. Sechellensis Forel, 1912: 159. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea coxalis var. cylindrical Forel, 1913: 5. Junior synonym of P. parallela: Brown, 1975: 8. 52

Platythyrea pusilla var. australis Forel, 1915: 10; Wheeler, G.C. & Wheeler, J. 1971b: 1198. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea parva Crawley, 1915: 133. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea pusilla var. egena Viehmeyer, 1916: 111. Junior synonym of P. parallela: Brown, 1975: 8.

145923291 Platythyrea coxalis var. philippinensis Viehmeyer, 1916: 283. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea cephalotes Viehmeyer, 1924: 224. Junior synonym of P. parallela: Brown, BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 1975: 8. Platythyrea melancholica var. aruana Karavaiev, 1925: 75. Junior synonym of P. parallela: Wilson, 1958: 151; Brown, 1975: 8. Platythyrea pusilla var. pacifica Santschi, 1928: 43. Junior synonym of P. parallela: Wilson and Taylor, 1967: 20. Platythyrea pulchella Santschi, 1928: 121. Junior synonym of P. parallela: Brown, 1975: 8. Platythyrea ceylonensis Donisthorpe, 1941: 202. Junior synonym of P. parallela: Brown, 1975: 8.

Type. Holotype worker from Papua New Guinea, Kokoda, 366m (BMNH).

Non-type Material examined. Two queens from N. Thailand, Chiang Mai Prov., Omkoi Dist., Hill evergreen forest, 17.VII.2016, N. Phengsi leg. (THNHM-I-02418 to THNHM-I-02419, THNHM); Four workers from N. Thailand, Lamphun Prov., Li Dist., Mae Li Forest Plantation, Teak Plantation, 18.IX.2013, W. Jaitrong leg., WJT180913-2 (THNHM-I-02403 to THNHM-I-02406, THNHM); one worker from N. Thailand, Chiang Mai Prov., Chiang Dao Dist., 24.VIII.2014, W. Jaitrong leg. (THNHM-I-02402, THNHM); one worker W. Thailand, Tak Prov., Um Phang Dist., Um Phang W.S., Mae Khlong Yai Village, Old field, 950-1000 m alt., 11.IX.2004, W. Jaitrong leg. (THNHM-I-02415, THNHM); one worker from W. Thailand, Kanchanaburi Prov., Thong Pha Phum Dist. 20.XII.2003, W. Jaitrong leg. (THNHM- I-02401, THNHM); one worker from NE. Thailand, Nakhon Ratchasima Prov., 53

Sakaerat Environmental Research Station, Dry evergreen forest, 17.VIII.2009, W. Jaitrong leg. (THNHM-I-02397); two workers from NE. Thailand, Loei Prov., Phu Luang W.S., 18.V.2007, S. Hasin leg. (THNHM-I-02409 to THNHM-I-02410, THNHM); one worker from NE. Thailand, Loei Prov., Phu Luang W.S., Agriculture area, 12.V.2007, S. Hasin leg. (THNHM-I-02408, THNHM); one worker from NE. Thailand, Mukdahan Prov., Cham Cha-E Dist., Cheang Chiang Niam Village,

145923291 8.VI.2007, W. Jaitrong leg. (THNHM-I-02407, THNHM); one worker from NE. Thailand, Phitsanulok Prov., Phu Soi Dao N.P., Hill evergreen forest, 1050 m alt., 8.VII.2008, D. Wiwatwitaya leg. (THNHM-I-02420, THNHM); two workers from E. BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Thailand, Chachoengsao Prov., Tha Kiab Dist., 26.X.2005, W. Jaitrong leg. (THNHM- I-02398 THNHM-I-02399, THNHM); Three workers from S. Thailand, Surat Thani Prov., Ban Nasan Dist., Ban Lamphun, 11.X.2011, W. Jaitrong leg. (THNHM-I-02466 to THNHM-I-02468, THNHM); one worker from S. Thailand, Surat Thani Prov., Ban Ta Khun Dist., Khlong Sang W.S., 15.X.2011, W. Jaitrong leg. (THNHM-I-02464, THNHM).

Description of worker. Measurements and indices, TL 3.86-5.81 mm, HL 0.82-1.12 mm, HW 0.62-0.96 mm, SL 0.59-0.82 mm, EL 0.16-0.26 mm, WL 1.15-1.81 mm, PL 0.49-0.72 mm, PH 0.46-0.60 mm, PW 0.40-0.53 mm, CI 70-88, EI 18-25, SI 75-95. Head in full-face view subrectriangular, longer than broad, with sides weakly convex, occipital corner almost straight, and posterior margin feebly concave; antennal scape relatively short, its apex reaching posterolateral corner of head; clypeus broad; posterior margin insert between frontal lobes; in profile clypeus weakly convex; anterior margin of clypeus roundly convex; mandible triangular, its masticatory margin with a large apical tooth, followed by 9-11 smaller teeth, the series as a whole decreasing in size to basal tooth; basal margin of mandible without a denticle; eye convex, located laterally anterior to mid-length of head, relatively large, 0.16-0.26 mm in maximum diameter, with 12 omatidia along the longest axis, distance between mandibular base and anterior margin of eye about 1.2 times as long as maximum eye length; distance between posterior margin of eye and occipital corner of head in lateral view about 2.4 times as long as distance between mandibular base and anterior margin of eye; frontal lobe wide and rounded; frontal carinae very widely spaced. Mesosoma 54

elongate, in profile with almost straight dorsal outline; promesonotal suture distinct; mesopleuron easily demarcated from mesonotum and metapleuron by a distinct suture; metapleuron not dermarcated from lateral face of propodeum; propodeum almost straight dorsal outline; propodeal junction blunt angle; declivity of propodeum concave; seen from back, propodeal declivity sinuate above; legs relatively short; propodeal spiracle opening circular. Petiole cylindrical, almost as long as high, its dorsal outline

145923291 weakly convex; in profile posterodorsal corner of petiole with very short obtusely angulated; declivity of petiole shallowly concave; in dorsal view petiole subrectangular, broadened posteriorly, its posterior margin straight or with shallow median concave; BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 subpetiolar process low, located anteroventrally; its ventral outline convex; ventral outline of petiole feebly concave. Sculpture: dorsa of head, mesosoma and petiole finely punctate with sparse shallow foveae; lateral face of head finely punctate with dense shallow foveae; lateral face of pronotum, mesopleuron, metapleuron, lateral face of propodeum and lateral face of petiole punctate with sparse large and shallow foveae; first gastral tergite micropunctate with dense foveae; second gastral tergite micropunctate with sparse small and shallow foveae; remaining gastral segments superficially finely micropunctate; coxae and femora superficially reticulate, but smooth and shiny interspaces. Setae: absent on main part of body but series of short setae present on tip of mandible and gaster; pubescence white, very short and fine, distributed over whole body and appendages. Color: entire body black to dark brown; tip of gaster, antenna and leg reddish brown yellowish brown.

Description of dealated queen. Measurements and indices, TL 4.85, HL 0.89, HW 0.63, SL 0.69, EL 0.26, WL 1.58, PL 0.56, PH 0.56, PW 0.40, CI 70, EI 42, SI 110. Head in full-face view subrectangular, clearly long than broad with weakly convex sides and almost straight posterior margin, occipital corner roundly convex; antennal scape relatively short, reaching posterolateral corner of head and size increase gradually to apex; clypeus broad; posterior margin inserted between frontal lobes; anterior margin of clypeus roundly convex; in profile clypeus weakly convex; mandible 55

triangular, its masticatory margin with a large apical tooth, followed by 10-11 smaller teeth, the series as a whole decreasing in size to basal tooth; basal margin of mandible almost straight without denticle; eye large and convex, located anterior to mid-length of head, 0.26 mm in maximum diameter with ca. 17 ommatidia on the longest axis; frontal lobe and frontal carinae similar to that of in worker caste; distance between anterior margin of eye and mandibular base longer than eye length; ocelli present but

145923291 relatively small. Mesosoma rather stout, in profile slightly convex dorsal outline; in dorsal view pronotum relatively long, almost as long as broad; mesoscutum hexagonal, relatively long, slightly longer than broad, anterior margin convex, separated from BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 mesoscutellum by a V-shaped shallow suture and separated from pronotum by transverse suture; parapsidal lines distinct, relatively long, straight and running anteriorly to mid-length of mesoscutum; mesoscutellum small, almost as long as broad; metanotum very short separated from mesoscutellum and propodeum by narrow sutures; propodeum relatively long; mesopleuron relatively long, anepisternum clearly demarcated from katepisternum by a deep suture; metapleuron not dermacated from lateral face of propodeum; propodeal junction obtusely angulated; declivity of propodeum shallowly concave; seen from back propodeal declivity rounded above; propodeal declivity concave. In profile view Petiole relatively short, subrectangular, almost as long as high; declivity of petiole concave; subpetiolar process low and subrectangular located anteroventrally; ventral outline of petiole feebly concave. Gaster larger than in the worker. Sculpture, setae and color condition similar to that of in the worker caste.

Distribution. Australia (type locality), New Caledonia, Borneo, Fiji, Guam, Indonesia (type locality), Malaysia, Micronesia, New Guinea, Northern Mariana Islands, Philippines, Samoa, Singapore, Solomon Islands, Seychelles, Bangladesh, India, Sri Lanka, Thailand (Chiang Mai, Lamphun, Tak, Kanchanaburi, Nakhon, Loei, Mukdahan, Phitsanulok, Chachoengsao and Surat Thani provinces), Vietnam and also see Figure 4-14 and Table 4-1.

Comparative notes. P. paralella is most similar to P. itoi in general appearance. See under P. itoi. 56

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 8. Worker of Platythyrea paralella. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 57

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 9. Queen of Platythyrea paralella. A, body in profile view; B, head in full- face view; and C, body in dorsal view. 58

Platythyrea quadridenta Donistrope, 1941 (Fig. 4-10)

Platythyrea quadridenta Donistrope, 1941: 134. Platythyrea quadridenta: Imai et al., 1984: 67; Ito, 1994: 211; Bolton, 1995: 337; Jaitrong and Nabhitabhata, 2005: 37.

145923291 Type. Holotype worker from Papua New Guinea, Kokoda, 366m (BMNH).

BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Non-type material examined. Eight workers from S. Thailand, Narathiwat Prov., Wang Dist., 6.XI.2002, W. Jaitrong leg., WJT02-TH-312 (THNHM); one worker from W. Malaysia, Ulu Gombak N.P., VI-X.1992, collector unknown.

Description of worker. Measurements and indices (Non-types, n = 9), TL 6.06-6.53 mm, HL 1.29-1.32 mm, HW 0.99 mm, SL 1.32 mm, EL 0.23-0.26 mm, WL 2.05-2.11 mm, PL 0.66-0.73 mm, PH 0.62-0.66 mm, PW 0.50-0.53 mm, CI 75, EI 23-26, SI 133. Head in full-face view rectangular, clearly longer than broad, with sides weakly convex or parallel, and posterior margin almost straight; antennal scape slender, 1/4 of its length extending beyond posterolateral corner of head; antennal segment II narrowest, about 0.5 times as long as segment III; III longer than each of IV-XII; clypeus broad; posterior margin clearly defined, insert between frontal lobes; in profile median portion of clypeus distinctly convex; anterior clypeal margin convex; mandible triangular, its masticatory margin with a large apical tooth, followed by 7 smaller teeth, large and smaller teeth alternating, but the series as a whole decreasing in size toward the basal tooth; basal margin of mandible almost straight without denticle; eye convex, located laterally anterior to mid-length of head, relatively large, 0.17-0.23 mm in maximum diameter, with 12 omatidia along the longest axis, distance between mandibular base and anterior margin of eye about 0.3 times as long as maximum eye length; frontal lobes wide and rounded; area between frontal lobes flat; frontal carinae widely spaced and weakly narrow posteriorly (Fig. 4-11B). Mesosoma elongate, in profile with weakly convex dorsal outline; promesonotal suture distinct; mesopleuron clearly demarcated from mesonotum and metapleuron by a shallow suture with 59

connecting promesonotal suture; metapleuron can be demarcated from lateral face of propodeum by indistinct ridge; propodeum feebly convex dorsal outline; propodeal spines present, obtusely angulated, shorter than broad at base; declivity of propodeum shallowly concave; legs relatively long and slender; propodeal spiracle opening circular. Petiole trapezoid and sessile, its length almost as long as high, dorsal outline convex; in profile posterodorsal corner of petiole with blunt angle overhanging

145923291 declivity of petiole; declivity of petiole concave; in dorsal view petiole subreticular, its posterior margin strongly concave with distinct lateral blunt angles; subpetiolar process weakly developed, subtriangular, located anteroventrally, its apex pointed forward and BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 downward; ventral outline of petiole almost straight or feebly concave. Sculpture: Posterior two thirds of head in full-face view entirely punctate with dense foveae, while anterior 1/3 finely punctate lacking foveae; antennal scape superficially finely micropunctate; dorsum of mesosoma and petiole punctate with dense foveae; mesopleuron and metapleuron finely punctate lacking foveae; gastral segments I and II finely microreticulate; coxa punctate, while femur and tibia superficially reticulate but shiny. Setae: Pubescence white, very short and dense, distributed over whole body and appendages; groups of short standing hairs present on anterior clypeal margin, tip of mandible and tip of gaster (pygidium and hypopygium). Color: Head, mesosoma, petiole, first and second gastral segments black to dark brown; antennal scape and legs reddish brown; antennal funicular, tip of gaster and tarsus yellowish brown.

Distribution. Thailand (Jaitrong & Nabhitabhata, 2005), Malaysia (Imai et al., 1984; Ito, 1994), Philippines and New Guinea (Donisthorpe, 1941) and also see Figure 4-14 and Table 4-1.

Comparative notes. Platythyrea quadridenta is most similar to P. bidentata Brown, 1975 and P. sagei Forel, 1900 in general appearance as they share the following characteristics: body darkness; frontal lobe broad; frontal carinae widely spaced and weakly narrow posteriorly; propodeal spiracle opening circulars; in dorsal view posterior margin of petiole concave. However, P. quadridenta can be distinguished 60

from P. bidentata by the following characteristics: posterior margin of head straight and parallel sides (posterior margin of head concave and convex sides in P. bidentata); beneath clypeal margin without small teeth hidden (beneath clypeal margin probably one or two small teeth hidden in P.bidentata); in dorsal view petiole clearly longer than broad (almost as longer as broad in P. bidentata); seen from above, pronotum with margins or humeral angles (pronotum without margins or humeral angles in P.

145923291 bidentata) . P. quadridenta can be separated from P. sagei by the following characteristics: head in full-face view rectangular (subtriangular in P. sagei); antennal scape relatively longer (1/4 of its length extending beyond posterolateral corner of head BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 in P. quadridenta; 1/5 of its length extending beyond posterolateral corner of head in P. sagei); declivity of propodeum shallowly concave (almost flat in P. sagei); petiole trapezoid (rectangular in P. sagei); in dorsal view petiole swollen (impressed in P. sagei). 61

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 10. Worker of Platythyrea quadridenta. A, body in profile view; B, head in full-face view; and C, body in dorsal view. 62

Platythyrea tricuspidata Emery, 1900 (Fig. 4-11)

Platythyrea tricuspidata Emery, 1900: 665. Platythyrea tricuspidata: Imai et al., 1984: 67; Bolton, 1995: 337; Ito, 1994: 210; Jaitrong and Nabhitabhata, 2005: 37. Senior synonym of penangensis: Brown,

145923291 1975: 9. Platythyrea tricuspidata subsp. penangensis Wheeler, W.M., 1929: 29. Junior synonym of tricuspidata: Brown, 1975: 9. BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Type. Syntype worker from Indonesia, Sumatra, Si Rambé (MSNG).

Non-type material examined. One worker from S. Thailand, Nakhon Si Thammarat Prov., Khao Nan N.P., Way to San Yen, 500-600 m alt., 17.IV.2007, W. Jaitrong leg., THNHM-02461 (THNHM); One worker from W. Malaysia, Ulu Gombak N.P., Selangor, Malaysia., X.1992, F. Ito leg., FI92MG-603 (THNHM).

Description of worker. Measurements and indices (Non-types, n = 2), TL 8.08 mm, HL 1.55 mm, HW 1.09 mm, SL 1.46 mm, EL 0.30 mm, WL 2.48 mm, PL 0.99 mm, PH 0.80 mm, PW 0.63 mm, CI 70, EI 27, SI 136. Head in full-face view rectangular, clearly longer than broad, with sides convex and posterior margin straight; antenna scape relatively long and slender, one thirds of its length extending beyond posterolateral corner of head; antennal segment II narrowest; antennal segment III almost as long as each of IV-XI; terminal segment (X) slightly longer than III; clypeus broad; in profile median portion of clypeus distinctly convex; posterior margin ill-defined; anterior clypeal margin clearly convex; mandible triangular, its masticatory margin with a large apical tooth, followed by a series of 10- 12 denticles, but the series as a whole decreasing in size toward the basal tooth; basal margin almost straight lacking denticle; eye convex, located laterally anterior to mid- length of head, relatively large, 0.27-0.30 mm in maximum diameter, with 19 omatidia along the longest axis, distance between mandibular base and anterior margin of eye about 0.3 times as long as maximum eye length; frontal lobes wide and rounded; area 63

between frontal lobes shallowly impressed; frontal carinae widely spaced and weakly narrow postriorly (Fig. 4-11B). Mesosoma elongate, in profile pronotum strongly convex dorsal outline; mesonotum and propodeal dorsum straight; promesonotal suture distinct; metanotal groove absent; mesopleuron narrow not clearly demarcated from mesonotum, but clearly separated from metapleuron by a shallow groove; metapleuron not demarcated from dorsal and lateral faces of propodeum; propodeal junction roundly

145923291 convex; propodeal spines present and blunt (but sharper than in P. quadridenta), shorter than broad at base; declivity of propodeum shallowly concave; legs relatively long and slender; propodeal spiracle opening circular. Petiole sessile, clearly longer than high, BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 its dorsal outline weakly convex; in profile posterodorsal corner of petiole with spines overhanging declivity of petiole; declivity of petiole shallowly concave; in dorsal view petiole clearly longer than broad, its posterior margin bearing three blunt and short spines; subpetiolar process developed, subtriangular, located anteroventrally, its apex pointed downward; ventral outline of petiole weakly convex. Sculpture: on head entirely punctate with dense foveae, pronotum, petiole and first gastral tergite finely punctate with dense foveae (but foveae weaker than on head); mesonotum, mesopleuron, propodeal dorsum, upper portion of metapleuron and upper portion of lateral face of propodeum finely punctate with sparse foveae; second gastral tergite finely reticulate but shiny; mandible finely punctate; antenna scape finely micropunctate; coxa macroreticulate but shiny; femur superficially reticulate but shiny; tibia micropunctate. Setae: absent on main part of body but series of short setae present on tip of gaster; pubescence white, very short and fine, distributed over whole body and appendages. Color: Head (excluding antenna, clypeus and mandible), mesosoma, petiole and gastral segments I and II black to dark brown; antenna, clypeus, mandible and legs and tip of gaster dark reddish brown.

Distribution. Thailand (Jaitrong & Nabhitabhata, 2005), Malaysia (Imai et al., 1984) and Indonesia (Emery, 1900) and also see Figure 4-14 and Table 4-1.

64

Comparative notes. Platythyrea tricuspidata is clearly distinguished from another species in the genus by it has three teeth or spines in posterior margin of petiole if seen in dorsal view.

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 11. Worker of Platythyrea tricuspidata. A, body in profile view; B, head in full-face view; and C, body in dorsal view. 65

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 12. Frontal view focusing on the frontal carinae. A, Frontal carinae very widely spaced; and B, Frontal carinae relatively narrowly separated Phengsi et al., 2018).

Figure 4- 13. Dorsal view focusing on the petiole. A, Posterior margin of petiole with two spines, teeth or blunt angles; B. Posterior margin of petiole with three spines, teeth or blunt angles; and C, Posterior margin of petiole without distinct spines, teeth or blunt angles (Phengsi et al., 2018). 66

145923291 BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14

Figure 4- 14. Distribution map of the ant genus Platythyrea in Thailand.

67

Table 4- 1. Distribution of Platythyrea in Thailand. N, Northern; NE, Northeastern; E, Eastern; W, Western; C, Central; and S, Southern.

Species groups and species N NE E W C S Platythyrea clypeata group 1. Platythyrea clypeata  2. Platythyrea grandis sp. nov.  

145923291 3. Platythyrea janyai  Platythyrea paralella group 1. Platythyrea itoi sp. nov.  BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 2. Platythyrea paralella      3. Platythyrea quadridenta  4. Platythyrea tricuspidata 

68

CHAPTER 5 DISCUSSION AND CONCLUSION

5.1 Discussion Until now, 41 species of the ant genus Platythyrea have been known around the world. Among them, seven species are found in Thailand and they belong to two

145923291 species groups (sensu Brown, 1975): P. clypeata group (P. clypeata, P. grandis sp. nov. and P. janyai) and P. paralella group (P. itoi sp. nov., P. paralella, P. quadridenta and P. tricuspidata). BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 Shape of frontal lobe, shape of mandible, mandibular dentition, shape of mesosoma and petiolar node are characters used by Brown (1975) to distinguish the two species groups mentioned above. These morphological characters were confirmed and used by several authors who described new species of the genus after Brown (1975) (Kuger, 1977; Lattke, 2003; De Andrade, 2004; Aria et al., 2011; Phengsi et al., 2018). The present study follows the previous works and also proposed the peculiar shape of propodeal spiracle, frontal carina condition, and shape of propodeal junction as more important characters to separate the two species groups. The P. clypeata species group is characterized by frontal lobe narrow; frontal carina closely spaced and strongly narrowed posteriorly; propodeal spiracle opening elliptical; petiole longer than high, posterior margin concave. The P. paralella species group is separated from the former by frontal lobe broad; frontal carina widely spaced and weakly narrow posteriorly; propodeal spiracle opening circular; petiole as long as high, posterior margin convex or with spines. Sculptures on body, teeth on petiole, size of eye, shape of subpetiolar process and pilosity are used to separate each species. The ant genus Platythyrea is clearly monomorphic in worker caste with a small size variation occurring within species. Thus, body size can be used for separating into large and small species. In this study body size has been used for separating Platythyrea grandis sp. nov. from P. clypeata and P. janyai from P. clypeata. Brown (1975) mentioned that P. paralella has variation in body size and external morphology. The specimens collected from Thailand and identified as P. paralella might contain several species. To solve the problem, DNA information must 69

be applied in future work. The other three species in the P. paralella species group (P. itoi, P. quadridenta and P. tricuspidata) are easily identified using external morphology. Currently, Platythyrea is presently distributed in pantropical and subtropical regions, from New World, Africa, Asia and Australia. The Afrotropical regions harbors the highest number of species (16 species), followed by the Oriental and Neotropical

145923291 regions (8 species in each region) (Antwiki, 2018). A total 41 species of the genus are distributed in the world, of which eight species are distributed in Southeast Asia while seven species distributed in Thailand. Among species of Thailand, P. paralella is the BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 most widely distributed. It can be found from the northern to southern parts of Thailand. In addition, it has been recorded from Laos, Vietnam, Malaysia, Indonesia, Singapore, Philippines, New Guinea, Australia, New Caledonia, Fiji, Guam, Micronesia, Northern Mariana Islands, Samoa, Solomon Islands, Seychelles (Donisthorpe, 1943; Jaitrong & Nabhitabhata, 2005; Jaitrong et al., 2016; Antwiki, 2018). In contrast, five species are specifically distributed: P. itoi sp. nov. is specifically distributed in northern Thailand, P. clypeata is specifically distributed in eastern Thailand, P. janyai, P. quadridenta and P. tricuspidata are specifically distributed in southern Thailand. These points out that Platythyrea species have limits to their geographic ranges whereas P. grandis sp. nov. is distributed in northern and western Thailand. According to available data of habitat types, Thai Platythyrea species occur from the low to high altitude areas. These can be separated into three group based on altitudinal distributional pattern; species of group I, two species (P. grandis and P. itoi), are confined to highland habitats; species of group II, three species ( P. janyai, P. quadridenta and P. tricuspidata), are confined to lowland habitats. These species of both groups might be sensitive to these habitats. Finally, for species of group III, a single species (P. paralella) occurs from the lowland to highland, so, the member of this group can resist against small or weak disturbances to the forest ecosystem. The African and Asian (Oriental and Indo-Australian). Platythyrea faunas have completely different sets of species. The sixteen Platythyrea species dominated in Africa have never been recorded from Asia (Antwiki, 2018). There is no distributed overlapping members between Southeast Asia and Africa. Consequently, P. grandis, 70

P. janyai and P. itoi described in the present study can be confirmed as new species of the genus Platythyrea.

5.2 Conclusion A total of seven species of the ant genus Platythyrea are recognized in Thailand based on external morphological characters of the worker caste. They belong

145923291 to the following two species groups: P. clypeata group (P. clypeata, P. grandis sp. nov. and P. janyai) and P. paralella group (P. itoi sp. nov., P. paralella, P. quadridenta and P. tricuspidata). BUU iThesis 59910009 thesis / recv: 28052561 13:49:13 seq: 14 A key to species based on the worker caste is provided and more detailed description is presented here for each species. The distribution map of the ant genus Platythyrea is presented here. P. paralella is the most widely distributed while P. clypeata is clearly restricted to eastern Thailand. Also, P. itoi sp. nov. is apparently restricted to northern Thailand whereas P. janyai, P. quadridenta and P. tricuspidata are clearly restricted to southern Thailand. REFERENCES

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