THESIS

TAXONOMY OF THE ANT GENUS ECHINOPLA SMITH, 1857 (HYMENOPTERA: FORMICIDAE: FORMICINAE) IN THAILAND

SOISUNEE TANANSATHAPORN

GRADUATE SCHOOL, KASETSART UNIVERSITY Academic Year 2018

THESIS APPROVAL GRADUATE SCHOOL, KASETSART UNIVERSITY

DEGREE: Master of Science (Entomology) MAJOR FIELD: Entomology DEPARTMENT: Entomology

TITLE: Taxonomy of the ant genus Echinopla Smith, 1857 (Hymenoptera: Formicidae: Formicinae) in Thailand

NAME: Miss Soisunee Tanansathaporn

THIS THESIS HAS BEEN ACCEPTED BY

THESIS ADVISOR (Miss Monthathip Kongmee, Ph.D.)

THESIS CO-ADVISOR (Associate Professor Nantasak Pinkaew, Ph.D.)

THESIS CO-ADVISOR (Mr. Weeyawat Jaitrong, Ph.D.)

DEPARTMENT HEAD (Assistant Professor Roungthip Masmeatathip, Ph.D.)

APPROVED BY THE GRADUATE SCHOOL ON

DEAN (Associate Professor Somwang Khantayanuwong, Ph.D.)

THESIS

TAXONOMY OF THE ANT GENUS ECHINOPLA SMITH, 1857 (HYMENOPTERA: FORMICIDAE: FORMICINAE) IN THAILAND

SOISUNEE TANANSATHAPORN

A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science (Entomology) Graduate School, Kasetsart University Academic Year 2018

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ABSTRACT Soisunee Tanansathaporn : Taxonomy of the ant genus Echinopla Smith, 1857 (Hymenoptera: Formicidae: Formicinae) in Thailand. Master of Science (Entomology), Major Field: Entomology, Department of Entomology. Thesis Advisor: Miss Monthathip Kongmee, Ph.D. Academic Year 2018

Taxonomy of the ants has been only recently started seriously in Thailand. This study aims to clarify the ant genus Echinopla of Thailand mainly based on the worker caste. Materials used in this study were mainly based on specimens deposited in the collections at the Natural History Museum of the National Science Museum, Pathum Thani Province, Thailand; the Faculty of Science, Prince of Songkla University, Thailand and at the Ant Museum, Faculty of Forestry, Kasetsart University, Thailand. Types or images of type specimens in AntWeb and AntWiki were examined to correct the identification. A total of 11 species are recognized among the Echinopla fauna of Thailand. They belong to the following four species groups: E. melanarctos group (E. cherapunjiensis, E. melanarctos, E. pallipes and E. tritschleri), E. mezgeri group (E. jeenthongi and E. tunkuaduljalilii), E. serrata group (E. fisheri and E. madli) and E. striata group (E. charernsomi, E. lineata and E. striata). Five species (E. cherapunjiensis, E. fisheri, E. tritschleri, E. striata and E. lineata) are records for the first time from Thailand. A dichotomous keys to the species group and Thai species of the genus based on worker caste and quality pictures of all species are provided. Distribution pattern of the species were also discussed.

______/ ____ / ____ Student's signature Thesis Advisor's signature

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ACKNOWLEDGEM ENTS

ACKNOWLEDGEMENTS

I would like to thank and gratitude to my advisor Dr. Monthathip Kongmee (Kasetsart University, Thailand) and co-advisors Associate Professor Nantasak Pinkaew (Kasetsart University, Thailand) and Dr. Weeyawat Jaitrong (National Science Museum, Thailand) for encouragement, instruction and suggestion for completely the thesis. I would like to thank Associate Professor Decha Wiwatwitaya (Kasetsart University, Thailand) and Dr. Nawee Noon-anant (Prince of Songkla University, Thailand), who kindly allowed me to examine ant specimens in their collections. I thank also to Mr. Yudthana Samung (Mahidol University, Thailand), who helped in taking pictures of the Thai Echinopla. I would like to thank the staffs and members in the taxonomic laboratory of the Department of Entomology, Faculty of Agriculture at Kamphaeng Saen, Kasetsart University, Kamphaeng Saen Campus and staffs of the Thailand Natural History Museum, provided some laboratory facilities. This research was partly supported by a Graduate Program Scholarship from the Graduate School, Kasetsart University, and by The Kasetsart Research and Development Institute. Finally, I am especially appreciated to my family and all of my friends for their helped and continuing encouragements throughout this research.

Soisunee Tanansathaporn

TABLE OF CONTENTS

Page

ABSTRACT ...... C

ACKNOWLEDGEMENTS ...... D

TABLE OF CONTENTS ...... E

LIST OF TABLES ...... F

LIST OF FIGURES ...... G

INTRODUCTION ...... 1

OBJECTIVES ...... 3

LITERATURE REVIEW ...... 4

MATERIALS AND METHODS ...... 14

RESULTS ...... 23

DISCUSSION AND CONCLUSION ...... 85

LITERATURE CITED ...... 89

APPENDIX ...... 93 CURRICULUM VITAE ...... 100

LIST OF TABLES

Page

Table 1. Distribution of Echinopla in Thailand. N, Northern; NE, Northeastern; E, Eastern; W, Western; Central and S, Southern...... 84

LIST OF FIGURES

Page

Figure 1 External morphology of the ant...... 5 Figure 2 External morphology of the ant genus Echinopla (workers). Body in profile. A. Echinopla striata; B. E. malanarctos; C. E. madli; D. E. tunkuaduljalilii...... 7 Figure 3 External morphology of the ant genus Echinopla (workers). Head in full-face view. A. Echinopla striata; B. E. malanarctos; C. E. madli; D. E. tunkuaduljalilii. .... 7 Figure 4 External morphology of the ant genus Echinopla (workers). Body from dorsal view. A. Echinopla striata; B. E. malanarctos; C. E. madli; D. E. tunkuaduljalilii...... 8 Figure 5 External morphology of the ant genus Echinopla (queen). Body in profile. A. Echinopla striata; B. E. malanarctos...... 8 Figure 6 External morphology of the ant genus Echinopla (queen). Head in full-face view. A. Echinopla striata; B. E. malanarctos...... 9 Figure 7 External morphology of the ant genus Echinopla (queen). Body from dorsal view. A. Echinopla striata; B. E. malanarctos...... 9

Figure 8 World distribution map of the ant genus Echinopla (AntWiki, 2019)...... 10

Figure 9 Map showing collection sites for Echinopla specimens in Thailand...... 16 Figure 10 Term of relative position for Echinopla specimens: A. head in full-face view and B. body in dorsal view...... 19

Figure 11 Whole body in profile view of Echinopla...... 20

Figure 12 Right mandible of Echinopla showing basal and masticatory margins...... 20

Figure 13 Morphometric measurements. A. head in full-face view; B. antenna; ...... 22 Figure 14 Worker of Echinopla charernsomi. A. body in profile view; B. head in full- face view and C. body in dorsal view...... 33 Figure 15 Worker of Echinopla cherapunjiensis. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 37 Figure 16 Worker of Echinopla fisheri. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 41 Figure 17 Worker of Echinopla jeenthongi. A. body in profile view; B. head in full- face view and C. body in dorsal view...... 47

H

Figure 18 Dealate qyne of Echinopla jeenthongi. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 48 Figure 19 Worker of Echinopla lineata. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 52 Figure 20 Worker of Echinopla madli. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 57 Figure 21 Dealate qyne of Echinopla madli. A. body in profile view; B. head in full- face view and C. body in dorsal view...... 58 Figure 22 Worker of Echinopla melanarctos. A. body in profile view; B. head in full- face view and C. body in dorsal view...... 62 Figure 23 Dealate gyne of Echinopla melanarctos. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 63 Figure 24 Worker of Echinopla pallipes. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 66 Figure 25 Worker of Echinopla striata. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 72 Figure 26 Male of Echinopla striata. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 73 Figure 27 Dealate gyne of Echinopla striata. A. body in profile view; B. head in full- face view and C. body in dorsal view...... 74 Figure 28 Worker of Echinopla tritschleri. A. body in profile view; B. head in full- face view and C. body in dorsal view...... 78 Figure 29 Worker of Echinopla tunkuaduljalilii. A. body in profile view; B. head in full-face view and C. body in dorsal view...... 82 Figure 30 Distribution map of the ant genus Echinopla in Thailand...... 83

INTRODUCTION

Echinopla Smith, 1857 is a formicinae ant genus of the tribe Camponotini, with Echinopla melanarctos Smith, 1857 being the type species (Bolton, 2003; Ward, et al., 2016). Members of the genus are known as arboreal ants (Bharti & Gul, 2012; Zettel & Laciny, 2015) which are most often encountered at the lower vegetation or on rotting wood. The genus is distributed in eastern India, China, various countries in Southeast Asia, New Guinea, and Australia (Bharti & Gul, 2012; Xu & Zhou, 2015; Zettel & Laciny, 2015, 2017). Until now, 36 extant species and three subspecies have been described in the genus Echinopla. Three species are recorded from India; 2 in China; 29 in Southeast Asia; 7 in New Guinea and 2 in Australia (Bharti & Gul, 2012; Jaitrong & Nabhitabhata, 2005; Laciny, et al., 2019; Lui et al., 2015; Tanansathaporn et al., 2018; Xu & Zhou, 2015; Zettel & Laciny, 2015, 2017).

Specimens of the ant genus Echinopla are rarely collected because they have a relatively small colony size and typically nest on canopy and forage individually (Bharti & Gul, 2012; Watanasit, et al., 2005; Zettel & Laciny, 2015). There are a few studies and publications of the genus in the World. Moreover, there are no any taxonomic revision of the genus Echinopla. Thus, a taxonomic revision of the genus are necessarily required.

In Thailand, Jaitrong and Nabhitabhata (2005), Zettel and Laciny (2015), Tanansathaporn et al. (2018), and Laciny et al. (2019) reported six named species: Echinopla charernsomi Tanansathaporn et Jaitrong, 2018; E. jeenthongi Tanansathaporn et Jaitrong, 2018; E. madli Zettel et Laciny, 2015; E. melanarctos Smith, 1857; E. pallipes Smith, 1857 and E. tunkuaduljalilii Laciny, Zettel, Maryati et Noor, 2019. In addition, Watanasit et al. (2005) recorded five unidentified species of the genus Echinopla in their study on “the composition of canopy ants at Ton Nga Chang Wildlife Sanctuary, Songkhla province, southern Thailand”. These lead to uncertain taxonomic status of the genus in Thailand. So, this study plan to examine all specimens of the genus deposited in several museums in Thailand, describe the 2

species (new and named species) and provide an identification key to the known Thai species and the distribution patterns of the ant genus Echinopla in Thailand.

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OBJECTIVES

1. To know and describe the species groups and all species of the Thai Echinopla (new records, known and/or new species) based on worker caste. 2. To provide a key to species group found in Thailand and a key to the Thai species. 3. To know the distribution pattern for each species in Thailand.

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LITERATURE REVIEW

1. Ants

Ants belong to a single family, the Formicidae of the order Hymenoptera (Bolton, 1994, 2003; Hölldobler & Wilson, 1990). In the world, a total of 17 extant subfamilies, 408 extant genera (including subgenera) and 15,519 species (including subspecies) have been listed (AntWiki, 2019).

Classification of ants

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Hymenoptera

Family: Formicidae

Ants are eusocial insects and play important roles in terrestrial ecosystems. Some ant genera can spread plant seeds, for example, Rhytidoponera, Pachycondyla, Monomorium, and Iridomyrmex (Buckley, 1982). In addition, the ant genus Oecophylla can be used as a biological control agent in agricultural systems (Offenberg, et al., 2013).

Ants have distinct external morphology from other insects in having the first abdominal segment fused propodeum and the second abdominal segment into a node- like petiole that petiole may be one or two nodes. The second nodes of petiole is called “postpetiole” (Fig. 1B). 5

A

B

Figure 1 External morphology of the ant. Source: http://www.antwiki.org/wiki/Morphological_Terms.

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2. External morphology of the ant genus Echinopla

The general terminology in the worker caste of the ants follows Hölldobler and Wilson (1990) and Bolton (1994). The important characteristics of worker and queen of the genus Echinopla are as follows (figs. 2–7):

Worker. Body stout. Head in full-face view rectangular to trapezoidal, usually broader than long, with slightly convex side; antenna long, 12-segmented, antennal scape distinctly surpassing posterolateral corner of head; mandible short, subractangular, anterior margin slightly convex, and its masticatory margin armed with five teeth; clypeus broad, anterior margin straight; palpi formular 6, 4; eyes relatively moderately protruding, positioned slightly behind mid-length of head; ocelli absent; frontal lobe chiefly horizontally oriented, usually concealing antennal fossae; frontal carina short, not passing the line connecting mid-points of eyes (Xu & Zhou, 2015).

Mesosoma oblong quadrate, dorsal outline completely arched or interrupted by impressed metanotal groove; promesonotal suture and metanotal groove present or absent; humeral corners of pronotum usually toothed or rarely armed; propodeal spiracles slit-like or elliptical; leg relative long (Xu & Zhou, 2015).

Petiole transverse and stout, subtriangular in profile view, strongly transverse in dorsal aspect; usually spined or toothed laterally and dorsally; rarely petiole node- like and cubic, without spines and tooth (Xu & Zhou, 2015; Zettel & Laciny, 2015).

Gaster globose to ovate, the first segment very large, strongly convex and completely covering the following tergites; hind margin moderately convex (Xu & Zhou, 2015; Zettel & Laciny, 2015).

Body surface usually strong sculptured, longitudinally striate, bluntly spined with background retirugose or densely cribrate with interface retirugose. Rarely body surface shagreened or smooth (Xu & Zhou, 2015). 7

Pilosity and pubescence usually abundant, rarely body dorsum hairless (Xu & Zhou, 2015).

A B

C D

Figure 2 External morphology of theก ant genus Echinopla (workers). Body in profile. A. Echinopla striata; B. E. malanarctosก ; C. E. madli; D. E. tunkuaduljalilii.

ก ก

A B

C D V

Figure 3 External morphology of the ant genus Echinopla (workers). Head in full-face view. A. Echinopla striata; B. E. malanarctos; C. E. madli; D. E. tunkuaduljalilii.

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A B V

C D

V

Figure 4 External morphology of the ant genus Echinopla (workers). Body from dorsal view. A. Echinopla striata; B. E. malanarctos; C. E. madli; D. E. tunkuaduljalilii.

Queen. Characteristics very similar to worker but queen has wings. Head with three ocelli; mesosoma with tegulae, massive, each side with a tooth-like or lobe-like process; mesoscutum long and broad; scutellum short and broad; metanotum very short and transverse; dorsum of propodeum short and broad, posterior margin rounded (Xu & Zhou, 2015).

A B

Figure 5 External morphology of the ant genus Echinopla (queen). Body in profile. A. Echinopla striata; B. E. malanarctos.

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A B

Figure 6 External morphology of the ant genus Echinopla (queen). Head in full-face view. A. Echinopla striata; B. E. malanarctos.

A B V V

Figure 7 External morphology of the ant genus Echinopla (queen). Body from dorsal view. A. Echinopla striata; B. E. malanarctos.

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3. Distribution

The ant genus Echinopla is distributed in eastern India, China, various countries in Southeast Asia, New Guinea, and Australia (fig. 8) (Bharti & Gul, 2012; Jaitrong & Nabhitabhata, 2005; Xu & Zhou, 2015; Zettel & Laciny, 2015, 2017). More information is provided in the Appendix.

Figure 8 World distribution map of the ant genus Echinopla (AntWiki, 2019).

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4. Habitats Members of the ant genus Echinopla are arboreal ants. Most species were rarely encountered individually from lower vegetation in evergreen forest (Zettel & Laciny, 2015). Bharti and Gul (2012) found E. cherapunjiensis nesting inside a hollowed twig. No ecological study has been done in the genus to date.

5. Historical reviews

5.1 Taxonomy

The genus Echinopla was firstly described by Smith (1857), without designation of the type species. Later, Wheeler (1911) listed the type species of genera and subgenera of the Formicidae, which he designed Echinopla melanarctos Smith, 1857 as the type species of the genus Echinopla.

Smith (1860) described Mesoxena as a new genus with designed Mesoxena mistura by monotype as the type species. Brown (1973) synonymized the genus with the genus Echinopla.

Since 1857 to 1943, twenty-three species of the genus Echinopla were described (André, 1892; Donisthorpe, 1936, 1941; Emery, 1898; Forel, 1901; Mayr, 1862; Smith, 1857; 1860; Stitz, 1911, 1938). Nearly, 70 years since 1943 (1944– 2011), no taxonomic papers dealing with the genus have been found. During the last ten years, 29 new species were described from Southeast Asia (Bharti & Gul, 2012; Tanansathaporn et al., 2018; Laciny et al., 2019; Zettel & Laciny, 2015, 2017). Up to now, 36 species are listed (Appendix). Among them, six named species are known from Thailand: Echinopla charernsomi, E. jeenthongi, E. madli, E. melanarctos, E. pallipes and E. tunkuaduljalilii. No taxonomic revision has been done in the genus to date.

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5.2 Previous records on the ant genus Echinopla in Southeast Asia

Echinopla is one of the small ant genera of the world and mainly distributed in Asia. Among them, 29 species are known in Southeast Asia (Appendix).

Smith (1857–1863) described nine new species of the ant genus Echinopla from Sundaland (Malaysia, Singapore and Indonesia) and a subspecies from the Philippines. Mayr (1862) and André (1892) added two new species from Java (E. lineata) and Borneo (E. rugosa), Indonesia, respectively. Emery (1898) described a species, E. vermiculata from an area east of the “Wallace Line” in Sulawesi Island.

Jaitrong and Nabhitabhata (2005) generated the checklist of ant species known in Thailand. Two hundred and forty-seven species distributed among 55 genera in nine subfamilies were recorded from Thailand. According to this checklist, two species of the ant genus Echinopla including E. melanarctos and E. pallipes were recorded.

Watanasit et al. (2005) examined the composition of canopy ants at Ton Nga Chang Wildlife Sanctuary, Songkhla province, southern Thailand. A total of 118 morphospecies in 29 genera and 6 subfamilies were identified. Five unidentified species of the genus were found.

Bharti and Gul (2012) described a new Echinopla species, E. cherapunjiensis from India. This species is also distributed in China, Myanmar, Laos and Malaysia (Lui et al., 2015; Xu & Zhou, 2015; Jaitrong et al., 2016; Zettel & Laciny, 2015, 2017).

General and Alpert (2012) generated a synoptic review of the ant genera of the Philippines. A total of 11 subfamilies and 92 genera were found of which specimens of two unidentified species of the genus Echinopla were recorded.

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Xu and Zhou (2015) provided key to the known species of the ant genus Echinopla and reported two species from China including E. striata and E. cherapunjiensis.

Zettel and Laciny (2015) reported 16 species of the ant genus Echinopla from west of Weber’s Line. In the paper, a new species, Echinopla madli was described from Thailand. In 2017, Zettel and Laciny again described a new species and provided description of seven species groups of the genus from Southeast Asia.

Tanansathaporn et al. (2018) described two new species, Echinopla charernsomi and E. jeenthongi from Thailand. Both species were collected individually from lowland dry evergreen forest.

Lastly, Laciny et al. (2019) described a new Echinopla species, E. tunkuabdujalilii from West Malaysia and South Thailand. The species belongs to Echinopla mezegeri species group.

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MATERIALS AND METHODS

1. Materials

1.1 70% Ethyl alcohol + Vials (2–3 ml) 1.2 Insect pins (No. 3) 1.3 Forceps 1.4 Card points 1.5 Umbrella 1.6 Stereoscope (ZEISS Discovery. V12 stereoscope) 1.7 Nikon ECLIPSE E600 microscope

2. Methodology

2.1 Study area

This taxonomic study was focused in Thailand and areas specific to southern western eastern and northeastern Thailand where distribution of the genus was found to be most from previous study diversity of ant in Thailand.

2.2 Sources of materials

This study was mainly based on the materials deposited in the collections at the Natural History Museum of the National Science Museum, Pathum Thani, Thailand (THNHM); the Faculty of Science, Prince of Songkla University, Thailand (PSU) and the Ant Museum, Faculty of Forestry, Kasetsart University, Thailand (AMK). Holotypes, paratypes and syntypes of named species were loaned from the following museums listed below.

Additional specimens were collected from 29 localities in 16 provinces. All specimens were dry-mounted on triangle card and deposited in the Natural History 15

Museum of the National Science Museum, Thailand (THNHM). Sweeping, beating and hand collecting methods were used to collect specimens of the genus from lower vegetation.

Abbreviation of the type depositions are as follows:

AMK Ant Museum, Faculty of Forestry, Kasetsart University, Thailand. MHNG Musee d'Histoire Naturelle Genève, Switzerland. NHMB Naturhistorisches Museum, Switzerland. PSU Faculty of Science, Prince of Songkla University, Thailand. THNHM Natural History Museum of the National Science Museum, Thailand. NHMW Natural History Museum, Vienna, Austria. PUPAC Punjabi University Patiala Ant Collection, India. CASC California Academy of Sciences, U.S.A. OUMNH Oxford University Museum of Natural History, United Kingdom. ITBC BORNEENSIS, Institute for Tropical Biology and Conservation. Malaysia Sabah University, Malaysia.

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s

Figure 9 Map showing collection sites for Echinopla specimens in Thailand.

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2.3 Observations

Most morphological observations were made with a ZEISS Discovery.V12 stereo microscope. Multi-focused montage images were produced using NIS element 3.7 from a series of source images taken by a Nikon MNB42100 digital camera attached to a Nikon ECLIPSE E600 microscope. Workers, dealate or alate queens and males were measured using a micrometer. All measurements were given in millimetres, rounded to the next one hundredth millimetre.

2.4 Terminology

Terms of relative position and general terminology of the worker ants follows Hölldobler and Wilson (1990) and Bolton (1994). For important characters of the worker in the genus Echinopla used, see Zettel and Laciny (2015). Queen characters of the genus, see Zettel and Laciny (2015) and Tanansathaporn et al. (2018).

A complete description of morphological features are given below:

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 clypeus margin.

cribrate Frame with wire netting through which flour.

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

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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.

mesonotum The tergite of the mesothorax.

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).

polyporous Multiple pore, having interstices in the skin or in the substance of the body.

reticulate-punctate Covered with a network of carinae, striate or rugae with punctures in the interspaces.

rugose With multiple wrinkles, especially running approximately in parallel.

striate Marked with striate, or multiple impressed lines.

pubescence Exceptionally short, fine hairs, typically forming a second layer beneath the pilosity.

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A

Posterior Posterolateral

Width Lateral

Length

Anterolateral

Anterior

B

Width Length

Anterior Posterior

Figure 10 Term of relative position for Echinopla specimens: A. head in full-face view and B. body in dorsal view. 20

Mesosoma Petiole Gaster

Pronotum Propodeum

First gastral tergite Head

Lateral face Lateral face of of propodeum pronotum

Figure 11 Whole body in profile view of Echinopla.

Basal margin of mandible

Basal tooth of mandible

T4 Masticatory T3 (apical) margin of mandible T2 Apical tooth of mandible T = tooth number

Figure 12 Right mandible of Echinopla showing basal and masticatory margins.

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2.5 Measurements and indices

The acronyms used for the measurements and indices are as follows:

TL Total length. The added lengths of head (including mandibles), mesosoma, petiole, and gaster. HL Head length. Length of head proper, excluding mandibles, measured along midline from anterior clypeal margin to posterior margin of head. HW Head width. Maximum width of head, in full-face view measured behind eyes (excluding eyes). SL Scape length. Maximum straight line length of antennal scape excluding the basal constriction and condylar bulb. ED Eye diameter. Maximum diameter of eye. PW Pronotal width. Maximum width of pronotum measured in dorsal view including spines. MSL Mesosoma length. Diagonal length of mesosoma in profile, from the point at which pronotum meets cervical shield to posterior margin of metapleuron. PL Petiole length. Length of petiole measured in profile from anterior process to posteriormost point of the tergite, where it surrounds gastral articulation. PH Petiole height. Height of petiole measured in profile from apex of the ventral process vertically to a line intersecting dorsalmost point of node including spines. DPW Dorsal petiole width. Maximum width of petiole in dorsal view including spines. GW Maximum width of gaster measured from dorsal view. CI Cephalic index. HW×100/HL. EI Eye index. ED×100/HW. SI Scape index. SL×100/HW. LPI Lateral petiole index. PH×100/PL. DPI Dorsal petiole index. PW×100/PL.

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A B C

E D

Figure 13 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.

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RESULTS

Taxonomy

A total of 11 species of the ant genus Echinopla are recognized in Thailand based on external morphological characters of the workers caste. They belong to the following four species groups: E. melanarctos group (E. cherapunjiensis, E. melanarctos, E. pallipes and E. tritschleri); E. mezgeri group (E. jeenthongi and E. tunhuabduljalilii); E. serrata group (E. fisheri and E. madli) and E. striata group (E. charernsomi, E. lineata and E. striata). Five species (E. cherapunjiensis, E. tritschleri, E. fisheri, E. lineata and E. striata) are recorded for the first time from Thailand. During the two years of the study, two new species have been added to the ant faunas of Thailand.

Genus Echinopla Smith, 1857

Echinopla Smith, 1857: 79. Type-species: Echinopla melanarctos, by subsequent designation of Wheeler, 1911: 162.

Mesoxena Smith, 1860: 106. Type-species: Mesoxena mistura by monotypy. Junior synonym of Echinopla: Brown, 1973: 182.

Diagnosis of the genus (worker). For a more extensive description of the worker caste of the genus, see Xu and Zhou (2015); Zettel and Laciny (2015). Some of the important characteristics are reproduced with slight modifications. Body stout. Head in full-face view rectangular to trapezoidal, usually broader than long, with slightly convex side. Mesosoma oblong quadrate, dorsal outline completely arched or interrupted by impressed metanotal groove; promesonotal suture and metanotal groove present or absent; humeral corners of pronotum usually toothed or rarely armed; propodeal spiracles slit-like or elliptical; leg relative long (Xu & Zhou, 2015). Petiole transverse and stout, subtriangular in profile view, strongly transverse in dorsal view; usually spined or toothed laterally and dorsally; rarely petiole node-like 24

and cubic, without spines and tooth (Xu & Zhou, 2015; Zettel & Laciny, 2015). Gaster globose to ovate, the first segment very large, strongly convex and completely covering the following tergites; hind margin moderately convex (Xu & Zhou, 2015; Zettel & Laciny, 2015). Body surface usually strong sculptured, longitudinally striate, bluntly spined with background retirugose or densely cribrate with interface retirugose. Rarely body surface shagreened or smooth (Xu & Zhou, 2015). Pilosity and pubescence usually abundant, rarely body dorsum hairless (Xu & Zhou, 2015).

Key to Echinopla species groups based on the worker caste

1 Dorsa of head, mesosoma, first gastral tergite and antennal scape without standing hairs; head clearly longer than broad...... E. mezgeri group

- Dorsa of head, mesosoma, first gastral tergite and antennal scape with dense standing hairs; head shorter than broad, as long as broad or slightly longer than broad...... 2

2 Dorsa of head and mesosoma finely longitudinally striate...... E. striata group - Dorsa of head and mesosoma bluntly spined or densely cribrate or polyporous...... 3 3 Dorsa of head, mesosoma and first gastral tergite with dense blunt spines...... E. melanarctos group - Dorsa of head, mesosoma and first gastral tergite with macropunctures or cribrate or polyporous...... E. serrata group

Echinopla melanarctos group

Xu and Zhou (2015) defined this species group as follows: Eyes located behind mid-length of head. Frontal lobes broad, lateral margin nearly straight or sinuate, completely or partially concealing antennal fossae. Mesosoma dorsum strongly convex, complete or interrupted by impressed metanotal groove. Mesosoma usually constricted at metanotal groove. Petiole transverse, sides and upper margin spined and toothed. Dorsa of head and mesosoma abundantly bluntly spined with background retirugose; gaster (first gastral tergite) abundantly bluntly spined, 25

longitudinally striate or densely cribrate. Body surface with abundant hairs and pubescence.

Currently valid names. Echinopla cherapunjiensis Bharti et Gul, 2012; Echinopla circulus Zettel et Laciny, 2015; Echinopla melanarctos Smith, 1857; Echinopla pallipes Smith, 1857; Echinopla tritschleri Forel, 1901.

Echinopla mezgeri group

Zettel and Laciny (2017) defined this species group as follows: Head clearly longer than broad. Eyes positioned at mid-length of head. Palp formula 5, 3. Antennal socket largely covered by frontal carinae. Mesosoma elongated, with weakly impressed mesometanotal suture; pronotum with tooth-like protrusions. Petiolar node stocky, subtriangular in lateral view, without spines or teeth. First gastral tergite with ventrally curved posterior margin; following gastral segments protruding ventrally. Body surface dull black, with extremely fine sculpture, almost without standing setae.

Currently valid names. Echinopla jeenthongi Tanansathaporn et Jaitrong, 2018; Echinopla mezgeri Zettel et Laciny, 2015; Echinopla tunkuabduljalilii Laciny, Zettel et Noor-Izwan, 2019.

Echinopla serrata group

Xu and Zhou (2015) defined this species group as follows: Eyes located behind mid-length of head. Frontal lobes broad and triangular, laterally bluntly angled, concealing most of antennal fossae. Mesosoma dorsum convex, entire or interrupted by impressed metanotal groove. Mesosoma constricted at metanotal groove or not. Petiole transverse, sides and upper margin spined and toothed. Body surface densely cribrate with interstice coarsely retirugose. Body surface with abundant hairs and pubescence.

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Currently valid names. Echinopla angustata Zettel et Laciny, 2015; Echinopla arfaki Donisthorpe, 1943; Echinopla australis Forel, 1901; Echinopla brevisetosa Zettel et Laciny, 2015; Echinopla crenulata Donisthorpe, 1941; Echinopla dubitata Smith, 1862; Echinopla fisheri Zettel et Laciny, 2015; Echinopla madli Zettel et Laciny, 2015; Echinopla maeandrina Stitz, 1938; Echinopla octodentata Stitz, 1911; Echinopla praetexta Smith, 1860; Echinopla rugose André, 1892; Echinopla serrata Smith, 1859; Echinopla silvestrii Donisthorpe, 1936; Echinopla turneri Forel, 1901; Echinopla wardi Zettel et Laciny, 2015.

Echinopla striata group

Xu and Zhou (2015) defined this species group as follows: Eyes located close to posterior corners of head. Frontal lobes narrow and elevated, lateral margin straight, posterior corner broadly rounded, concealing half of antennal fossae. Mesosoma dorsum convex and interrupted by impressed metanotal groove. Mesosoma constricted at metanotal groove. Petiole transverse, sides and upper margin spined and toothed. Body dorsum longitudinally striate. Body surface with abundant hairs and pubescence.

Currently valid names. Echinopla deceptor Smith, 1863; Echinopla densistriata Stitz, 1938; Echinopla lateropilosa Zettel et Laciny, 2017; Echinopla lineata Mayr, 1862; Echinopla pseudostriata Donisthorpe, 1943; Echinopla senilis Mayr, 1862; Echinopla striata Smith, 1857; Echinopla vermiculata Emery, 1898.

Checklist of the Thai Echinopla species

Echinopla melanarctos species group Echinopla cherapunjiensis Bharti et Gul, 2012 Echinopla melanarctos Smith, 1857 Echinopla pallipes Smith, 1857 Echinopla tritschleri Forel, 1901 27

Echinopla mezgeri species group Echinopla jeenthongi Tanansathaporn et Jaitrong, 2018 Echinopla tunkuaduljalilii Laciny, Zettel, Maryati et Noor, 2019

Echinopla serrata species group Echinopla fisheri Zettel et Laciny, 2015 Echinopla madli Zettel et Laciny, 2015

Echinopla striata species group Echinopla charernsomi Tanansathaporn et Jaitrong, 2018 Echinopla lineata Mayr, 1862 = Echinopla sucki Forel, 1901 Echinopla striata Smith, 1857

Key to the Thai species based on worker caste

1 Dorsa of head, mesosoma, first gastral tergite and antennal scape without standing hairs...... 2

- Dorsa of head, mesosoma, first gastral tergite and antennal scape with dense standing hairs...... 3

2 Dorsa of head, pronotum and first gastral tergite with dense macropunctures; posterior margin of first gastral tergite with numerous small denticles...... E. jeenthongi - Dorsa of head, pronotum and first gastral tergite opaque, with very weak punctures; posterior margin of first gastral tergite without denticles...... E. tunhuabduljalilii 3 Dorsa of head and mesosoma finely longitudinally striate...... 4 - Dorsa of head and mesosoma bluntly spined or densely cribrate...... 6 4. Lateral faces of pronotum, mesopleuron, metapleuron and lateral face of propodeum longitudinally striate...... E. lineata 28

- Lateral face of pronotum, mesopleuron and metapleuron without longitudinal striation...... 5

5 Lateral face of propodeum smooth and shiny; larger species (HW 1.70–1.80 mm); standing hairs on head and mesosoma black; trochanters black to dark brown...... E. striata

- Lateral face of propodeum reticulate-punctate and with ill-defined longitudinal striation; smaller species (HW 1.40–1.60 mm); standing hairs on head and mesosoma yellowish brown; trochanters yellowish brown...... E. charernsomi 6 Dorsa of head, mesosoma and first gastral tergite with dense blunt spines...... 7 - Dorsa of head, mesosoma and first gastral tergite with macropunctures or cribrate...... 10 7 In profile view, dorsum of mesosoma strongly convex and forming a complete arch, not interrupted by impressed or notched metanotal groove; in dorsal view, mesosoma slightly constricted at metanotal groove position; eye relatively small (EI 10–16), distinctly protruded...... 8

- In profile view, dorsum of mesosoma almost straight, not forming a complete arch, interrupted by impressed or notched metanotal groove; in dorsal view, mesosoma strongly constricted at metanotal groove position; eye relatively large (EI >16), moderately convex...... 9 8 Dorsa of head, mesosoma and first gastral tergite well developed long bluntly spines; lateral face of head regularly longitudinally striate; in dorsal view each side of petiole with long spine, longer than half the width of the node; larger species (HW 2.10–2.45 mm); legs and antennal scape black. …...... E. melanarctos - Dorsa of head and mesosoma with weakly developed short blunt spines, first gastral tergite densely cribrate; lateral face of head irregularly rugose; in dorsal view, each side of petiole with short spine, shorter than half width of the node; smaller species (HW 1.55–1.75 mm); legs yellowish brown; antenna reddish brown...... E. pallipes 29

9 Dorsa of head and mesosoma with short blunt spines; lateral faces of head and mesosoma with dense white hairs; dorsal outline of mesosoma indistinctly interrupted by narrowly notched metanotal groove; lateral face of first gastral tergite densely cribrate; body black; mandibles and antennae reddish brown; legs yellowish brown...... E. cherapunjiensis - Dorsa of head and mesosoma with moderately long blunt spines, their apex truncate and bearing a long yellowish hairs; in lateral view, dorsal outline of mesosoma interrupted by distinctly impressed metanotal groove; lateral face of first gastral tergite finely longitudinally striate; body black; mandibles and antennae reddish brown; legs and gastral apex yellowish brown; eyes blackish brown...... E. tritschleri 10 Dorsal outlines of promesonotum and propodeum weakly convex, separating them with shallow metanotal groove; head dorsally and laterally polyporous, with closely set pores; dorsal margin of petiole with small teeth; smaller species (HW 0.90–1.00 mm)...... E. madli - Dorsal outlines of promesonotum and propodeum strongly convex, separating them by deep metanotal groove; head dorsally and laterally polyporous, unevenly large pores and smooth and shiny interspaces; dorsal margin of petiole with large teeth; larger species (HW 1.15–1.25 mm)...... E. fisheri

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Echinopla charernsomi Tanansathaporn et Jaitrong, 2018 (Fig. 14)

Echinopla charernsomi Tanansathaporn et Jaitrong, in Tanansathaporn et al., 2018: 3, figs 1–3.

Type. Holotype. Worker (THNHM-I-01365, examined), NE Thailand, Nakhon Ratchasima province, Wang Nam Khiao district, Sakaerat Environmental Research Station, 14.5097222°N, 101.9319444°E, 17.VIII.2009, W. Jaitrong leg. (THNHM, examined). Paratypes. One worker (THNHM-I-01366), same data as holotype (THNHM, examined); one worker (THNHM-I-01367), Central Thailand, Nakhon Nayok province, Muang Nakhon Nayok, Hin Tang subdistrict, 14.43722222°N, 101.37583333°E, dry evergreen forest, 25.III.2000, W. Jaitrong leg. (THNHM, examined).

Non-type material examined. One worker (THNHM-I-01218), W Thailand, Tak province, Umphang district, Thung Yai W.S., Ka-nga Sod Forest Ranger Station, 20.II.2015, W. Jaitrong leg. (THNHM).

Measurements. Holotype. TL 5.32 mm, HL 1.28 mm, HW 1.60 mm, SL 1.32 mm, ED 0.28 mm, PW 1.52 mm, MSL 1.96 mm, PL 0.52 mm, PH 0.48 mm, DPW 1.40 mm, GW 1.72 mm, CI 125, EI 18, SI 83, LPI 92, DPI 292. Paratype (n = 2). TL 4.72–5.28 mm, HL 1.12–1.16 mm, HW 1.40–1.56 mm, SL1.20–1.28 mm, ED 0.28 mm, PW 1.28–1.52 mm, MSL 1.84–1.96 mm, PL 0.56–0.64 mm, PH 0.52–0.56 mm, DPW 1.24–1.48 mm, GW 1.52–1.80 mm, CI 130–134, EI 18–19, SI 82–84, LPI 88– 91, DPI 226–231.

Diagnosis. Medium-sized, HW 1.40–1.56 mm, head in full-face view broader than long; eye relatively large, moderately convex; dorsum of mesosoma distinctly laterally marginate along its length, armed with numerous teeth; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture present, narrow and shallow; metanotal groove distinct, deeply impressed; in posterior 31

view, dorsal margin of petiole with numerous teeth; lateral margin of petiole with 3–4 large teeth; first gastral tergite globose, its posterior margin without denticles. Dorsa of head, mesosoma and first gastral tergite finely, longitudinally striate; lateral face of pronotum irregularly reticulate-punctate; mesopleuron punctate; lateral face of propodeum irregularly rugose. Dorsa of head, mesosoma, petiole and first gastral tergite with dense black and long hairs mixed with dense pubescence. Body black; trochanters and tip of gaster reddish brown.

Description. Worker (fig. 14). Head in full-face view trapezoidal, narrowed anteriorly and broader than long, lateral margin weakly convex, posterior margin moderately convex, posterolateral corner roundly convex. Eye relatively large, moderately convex, located close to posterior corner of head. Frontal lobe obliquely elevated, in full-face view covering antennal fossa only in part. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight. Frontal carina short, not surpassing level of anterior margin of eye. Antennae 12-segmented; scape moderately long and slender, one third of scape length extending beyond posterior margin of head; funicular segments, each longer than broad; antennal segment II slightly narrower and longer than each of III–XI; terminal segment (XII) almost as long as X and XI combined.

Mesosoma stout, dorsum of mesosoma distinctly marginate at sides along entire length, length roughly 1.3 times pronotum width; propodeum slightly larger than promesonotum. In profile dorsal outline of promesonotum weakly convex, while propodeum strongly convex; both promesonotum and propodeum meeting at deep and broad incision, the metanotal groove; promesonotal suture present, narrow and shallow; mesopleuron flat, not clearly demarcated from metapleuron. In dorsal view mesosoma with waist-like incision in front of propodeum; pronotum trapezoidal or subrectangular, anterior margin convex, posterior margin concave, lateral margin straight; pronotum narrower than head; mesonotum clearly shorter and narrower than pronotum; propodeum more or less subglobose, as broad as pronotum, and longer than mesonotum; anterior and lateral margins of mesosoma armed with numerous 32

teeth. Legs relatively long; femora flat, seen from posterior aspect broad at base then gradually narrowed to apex.

Petiole in profile longer than high, and narrowed dorsally. In dorsal view petiole transverse, about as broad as propodeum, each side with strong posteriorly bent spines; anterior face much narrower than posterior face. In posterior aspect dorsal margin with 12–13 denticles; lateral margin with 3–4 large teeth.

First gastral tergite in dorsal view globose, covering remaining tergites, slightly broader than long, posterior margin roundly convex without denticles.

Dorsum and lateral face of head finely, longitudinally striate; antennal scape superficially reticulate with smooth and shiny interspaces; mandible roughly and irregularly rugose. Dorsa of pronotum, mesonotum and propodeum longitudinally striate; lateral face of pronotum irregularly reticulate-punctate; mesopleuron punctate; lateral face of propodeum irregularly rugose; legs reticulate with smooth and shiny interspaces. First gastral tergite densely, finely, longitudinally striate.

Dorsa of head, mesosoma, petiole, and first gastral tergite with dense erect long hairs mixed with dense pubescence. Scape and legs with sub erect long hairs mixed with sparse pubescence.

Entire body black; apex of mandible, antennal segments X–XII, trochanters, and tip of gaster reddish brown; eye grayish brown.

Remarks. E. charernsomi belongs to the Echinopla striata group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. striata and E. lineata by having a longitudinal striation on the dorsal body surface. However, E. charernsomi can be distinguished from E. striata by much smaller size (HW 1.40–1.60 mm in E. charernsomi, HW 1.72–1.85 mm in E. striata), irregularly reticulate-punctate lateral face of propodeum (smooth and shiny in E. striata), and yellowish brown trochanters (black to dark brown in E. striata). Echinopla charernsomi is separated 33

from E. lineata by a punctate and reticulate-punctate sculpturing on the lateral face of the mesosoma (entirely and finely striate in E. lineata), yellowish brown trochanters (dark brown in E. lineata) and body surface with sparse white hairs (dense white hairs in E. lineata).

Habitat. No biological information is available for E. charernsomi. However, judging from the specimen examined this species inhabits lowland to highland. All specimens of the species in this study were collected from lower vegetation.

Distribution. Thailand (Nakhon Ratchasima, Nakhon Nayok and Tak provinces).

A

B C

Figure 14 Worker of Echinopla charernsomi. A. body in profile view; B. head in full- face view and C. body in dorsal view.

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Echinopla cherapunjiensis Bharti et Gul, 2012 (Fig. 15)

Echinopla cherapunjiensis Bharti & Gul, 2012: e-53, figs. 1-3.

Echinopla cherapunjiensis: Liu et al. 2015: 35; Xu & Zhou, 2015: 32; Zettel & Laciny 2015: 113; Zettel & Laciny 2017: 15.

Type. Holotype worker (PUPAC, Antweb images examined) from Cherapunji, Meghalaya, 1200 m. alt., India, 25.2988°N, 91.7086°E, 2.IV.2009, Irfan Gul leg.

Non-type material examined. One worker, W Thailand, Tak Prov., Umphang Dist., Thung Yai W.S., Huai Nam Kheao, 23.VI.2015, W. Jaitrong leg. (THNHM); 9 workers, W Thailand, Tak Prov., Umphang Dist., Thung Yai W.S., Ka- nga Sod Forest Ranger Station, 15.II.2015, W. Jaitrong leg., TH15-WJT-383 (THNHM); 1 worker, W Thailand, Tak Prov., Umphang Dist., Thung Yai W.S., Bae Ki Forest Ranger Station 17.II.2015, W. Jaitrong leg. (THNHM); 3 workers from NE Thailand, Loei Prov., Phu Luang W.S., 1,000 m. alt., 9.V.2007, S. Hasin leg., SH07- TH 15 (THNHM); 2 workers, same locality, date and collector, SH07-TH 9 (THNHM); 4 workers, same locality and collector, 15.V.2007, SH07-TH 101 (AMK, THNHM); 1 worker, central Thailand, Phitsanulok Prov., Phu Soi Dao N.P., 8 .VII.2006, D. Wiwatwittaya leg. (AMK); 3 workers, central Thailand, Phetchaburi Prov., Kang Kachan N.P., 28.XI.2006, W. Saguansombat leg., WS280906-1 (THNHM); 2 workers, S Thailand, Nakhon Si Thammarat Prov., Khao Nan N.P., San Yen, Hill Evergreen Forest (HEF), 22.IV.2007, W. Jaitrong leg. (THNHM); 1 worker, same locality, 19.IV.2007, W. Jaitrong leg. (THNHM); 1 worker, S Thailand, Narathiwat Prov., Su-ngai Golok Dist., Toh Dang Peat Swamp Forest, 12.X.2000, S. Hasin leg. (AMK).

35

Measurements. Non-type workers (n = 5): TL 4.45–5.75 mm, HL 1.15–1.30 mm, HW 1.32–1.80 mm, SL 1.15–1.40 mm, ED 0.25–0.30 mm, PW 1.30–1.60 mm, MSL 1.70–2.15 mm, PL 0.50–0.70 mm, PH 0.50–0.60 mm, PW 0.80–0.95 mm, GW 1.80–2.15 mm, CI 110–138, EI 16–23, SI 77–97, LPI 86–100, DPI 229–260.

Diagnosis. Large-sized, HW 1.32–1.80 mm, head in full-face view globose, broader than long; eye relatively large moderately convex; dorsum of mesosoma laterally immarginate; in dorsal view mesosoma with waist-like, incision in front of propodeum; promesonotal surture and metanotal groove weak; in posterior view, dorsal margin of petiole with numerous teeth; first gastral tergite globose, its posterior margin without denticles. Dorsa of head and mesosoma with short blunt spines; lateral faces of head and mesosoma with dense white hairs; lateral face of first gastral tergite densely cribrate. Body black; mandibles and antennae reddish brown; legs yellowish brown.

Description. Worker (fig. 15). Head in full-face view globose, broader than long; lateral and posterior margins of head weakly convex. Eye relatively large, moderately convex, located close to posterior corner of head. Frontal lobe obliquely elevated, in full-face view covering antennal fossa only in part. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus weakly convex, anterior margin almost straight. Frontal carina short, not surpassing level of anterior margin of eye. Antennae 12-segmented; scape moderately long and slender; antennal segment II clearly longer than each of III–XI; terminal segment (XII) almost as long as X and XI combined.

Mesosoma stout, dorsum of mesosoma laterally immarginate at sides in dorsal view; propodeum slightly narrower and longer than pronotum. In profile dorsal outline of mesosoma almost straight, not forming a complete arch, interrupted by impressed or notched metanotal groove, promesonotal surture and metanotal groove weak. In dorsal view mesosoma with waist-like incision in front of propodeum; maximum pronotal width slightly narrower than head excluding eyes. Legs relatively 36

long and slender; femora flat, seen from posterior aspect broad at base then gradually narrowed to apex.

Petiole in profile view subtriangular, longer than high, ventral outline almost straight. In dorsal view petiole transverse, armed with seven teeth and slightly narrower than propodeum width.

First gastral tergite in dorsal view globose, covering remaining tergites, slightly broader than long and its posterior margin convex, without denticles.

Dorsa of head and mesosoma with short blunt spines; lateral faces of head and mesosoma with dense white hairs; lateral face of first gastral tergite densely cribrate.

Entire body black; mandible and antenna reddish brown; legs yellowish brown.

Remarks. Echinopla cherapunjiensis belongs to the E. melanarctos group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. pallipes and E. tritschleri by having dense blunt spines on the dorsa of head and mesosoma. However, E. cherapunjiensis can be distinguished from E. pallipes by dorsum of mesosoma almost straight, not forming a complete arch, interrupted by impressed or notched metanotal groove (strongly convex and forming a complete arch, not interrupted by impressed or notched metanotal groove in E. pallipes); eye relatively large, moderately convex (eye relatively small, distinctly protruded in E. pallipes). This is separated from E. tritschleri by dorsa of head and mesosoma with short blunt spines (moderately long blunt spines in E. tritschleri); lateral face of first gastral tergite densely cribrate mixed with sparse short blunt spines (irregularly longitudinally striate in E. tritschleri).

Habitat. Bharti and Gul (2012) reported that E. cherapunjiensis was found from high altitude range of North-East Himalaya. The species inhabits the areas with 37

wet atmospheric conditions and high annual rainfall and was collected the holotype from a branch of a broad leaved tree. A colony from Thung Yai Wildlife Sanctuary, Tak province, colony no. TH15-WJT-383 was collected from a rotting branch on the forest floor. Other specimens were collected from lower vegetation. This species is distributed in both Indo-China and Sundaland in slightly wet forest (evergreen and hill evergreen forests).

Distribution. India [type locality, (Bharti & Gul, 2012)], China (Liu et al., 2015; Xu & Zhou, 2015; Zettel & Laciny, 2017), Laos, Myanmar (Zettel & Laciny, 2017), Thailand (new record) and Malaysia (Zettel & Laciny, 2015).

A

B C

Figure 15 Worker of Echinopla cherapunjiensis. A. body in profile view; B. head in full-face view and C. body in dorsal view.

38

Echinopla fisheri Zettel et Laciny, 2015 (Fig. 16)

Echinopla fisheri Zettel & Laciny, 2015: 108, figs. 17-20.

Echinopla fisheri: Zettel & Laciny, 2017: 14.

Type. Holotype worker (NHMW, Antweb images examined) from E Malaysia, Borneo, Sabah, Danum Valley, Nature trail, 180 m alt., 4.96072°N; 117.80014°E in rainforest, 24.VII.2010, B.L. Fisher leg.

Non-type material examined. One worker from W Thailand, Kanchanaburi Prov., Thong Pha Phum Dist., Dry Evergreen Forest, 13.IV.2004, C. Bourmas leg. (AMK); 1 worker from E Thailand, Chanthaburi Prov., evergreen forest, 24.XII.2003, W. Jaitrong leg., (THNHM); 1 worker from S Thailand, Trang Prov., Palian Dist., Ton Tae Waterfall, 200-300 m a.s.l., 28.III.2005, W. Jaitrong leg. (THNHM).

Measurements. Non-type worker (n = 3): TL 4.50–4.65 mm, HL 0.85–1.30 mm, HW 1.15–1.25 mm, SL 1.25–1.30 mm, ED 0.20–0.25 mm, PW 1.00–1.15 mm, MSL 1.75–1.95 mm, PL 0.50–0.65 mm, PH 0.40–0.50 mm, PW 0.95–1.05 mm, GW 1.30–1.60 mm, CI 79–83, EI 26, SI 105, LPI 83–122, DPI 158–211.

Diagnosis. Medium-sized, HW 1.15–1.25 mm, head in full-face view oval, almost as long as broad; eye relatively large, moderately convex; dorsum of mesosoma laterally immarginate; pronotum with a pair of sharp tubercles, otherwise tubercles on mesosoma hardly developed; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture indistinct; metanotal groove distinct, deep and broad impression; in profile dorsal outlines of promesonotum and propodeum strongly convex, separating them by deep metanotal groove; in posterior 39

aspect, dorsal margin petiole armed with 4 teeth, while lateral margin with a large tooth; posterior margin of first gastral tergite with numerous ill-defined denticles. Head dorsally and laterally polyporous, unevenly large pores and smooth and shiny interspaces. Dorsal surfaces of head, mesosoma, petiole, and first gastral tergite with long and brown standing hairs. Body black; legs reddish brown; antenna and mandible dark brown.

Description. Worker (fig. 16). Head in full-face view oval, almost as long as broad, lateral margin weakly convex and almost straight posterior margin bearing small medial lobe; eye relatively large, moderately convex, located close to posterior corner of head. Frontal lobe chiefly horizontally oriented, covering antennal fossa only in part. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight. Frontal carina short, not surpassing level of anterior margin of eye. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head; funicular segments, each longer than broad; antennal segment II clearly narrower and longer than each of III–XI.

Mesosoma slender, in profile dorsal outlines of promesonotum and propodeum strongly convex, separating them by deep and broad impression metanotal groove; promesonotal suture only laterally indicated. In dorsal view mesosoma with waist-like incision in front of propodeum; laterally immarginate; pronotum with a pair of sharp tubercles, otherwise tubercles on mesosoma hardly developed; pronotum narrower than head; mesonotum clearly shorter than pronotum; propodeum suboval, narrowed anteriorly, as broad as pronotum and longer than mesonotum, lateral margin armed with numerous teeth. Legs relatively long and slender.

Petiole in profile longer than high, and narrowed dorsally. In dorsal view petiole transverse, about as broad as propodeum. In posterior aspect, dorsal margin armed with 4 teeth, while lateral margin with a large tooth. 40

First gastral tergite in dorsal view globose, covering remaining tergites, slightly longer than broad, along posterior margin with transverse impression; posterior margin with numerous ill-defined denticles.

Dosum and lateral face of head polyporous, unevenly large pores and smooth and shiny interspaces.

Dorsal surfaces of head, mesosoma, petiole, and first gastral tergite with long and brown standing hairs.

Entire body black; legs reddish brown; antenna and mandible dark brown.

Remarks. Echinopla fisheri belongs to the E. serrata group (Xu & Zhou, 2015; Zettel & Laciny, 2017). It is closely related to Echinopla madli by having dense macropunctures or cribrate on the body surface. However, E. fisheri can be distinguished from E. madli by the following characteristics: dorsal outlines of promesonotum and propodeum strongly convex, separating them by deep metanotal groove (weakly convex, separating them with shallow metanotal groove in E. madli); head dorsally and laterally polyporous, unevenly large pores and smooth and shiny interspaces (with closely set pores in E. madli); larger body size (HW 1.15–1.25 mm in E. fisheri; HW 0.90–1.00 mm in E. madli).

Habitat. Little is known about the bionomics of E. fisheri. The holotype was found along a nature trail, 180 m a.s.l. in rainforest (Zettel & Laciny, 2015) while the non-type material examined from West and East Thailand, Kanchanaburi and Chanthaburi provinces was collected from dry evergreen forest. This species seem to be distributed in both Indo-China and Sundaland.

Distribution. Thailand (Kanchanaburi, Chanthaburi and Trang provinces, new record) and Malaysia [type locality, (Zettel & Laciny, 2015)].

41

A

B C

Figure 16 Worker of Echinopla fisheri. A. body in profile view; B. head in full-face view and C. body in dorsal view.

42

Echinopla jeenthongi Tanansathaporn et Jaitrong, 2018 (Figs. 17, 18)

Echinopla jeenthongi Tanansathaporn & Jaitrong, in Tanansathaporn et al., 2018: 6, figs. 4-6.

Types. Holotype worker (THNHM-I-01368, examined), S Thailand, Nakhon Si Thammarat province, Sichon district, Khao Noi subdistrict, near Yod Nam Waterfall, 8.90916667°N, 99.73194444°E, 1.XI.2008, T. Jeenthong leg. (THNHM, examined). Paratypes: One worker (NW310705-1), S Thailand, Surat Thani province, Ban Nasarn district, Julaporn 8, 8.86138889°N, 99.48472222°E, evergreen forest, 400-550 m a.s.l., lower vegetation, 31.VII.2005, N. Noon-anant leg. (PSU, examined); 1 worker (NW150405-2), S Thailand, Nakhon Si Thammarat province, Noppitam district, Noppitum [Noppitam] sub-district, Krung Ching, 8.65722222°N, 99.63638889°E, 100-300 m a.s.l., lower vegetation, 15.IV.2005, N. Noon-anant leg. (PSU, examined); 1 dealate gyne (NW091105-3), S Thailand, Phangnga province, Muang Phangnga, Song Prang sub-district, 8.61111111°N, 98.55055556°E, 200-300 m a.s.l., lower vegetation, 09.XI.2005, N. Noon-anant leg. (PSU, examined).

Measurements. Holotype. TL 3.80 mm, HL 1.12 mm, HW 1.04 mm, SL 1.12 mm, ED 0.24 mm, PW 0.92 mm, MSL 1.62 mm, PL 0.53 mm, PH 0.50 mm, DPW 0.96 mm, GW 1.25 mm, CI 93, EI 23, SI 108, LPI 94, DPI 181. Paratype (n = 3). TL 4.40–4.50 mm, HL 0.89–1.15 mm, HW 0.96–1.00 mm, SL 1.06–1.15 mm, ED 0.23–0.25 mm, PW 0.85–0.86 mm, MSL 1.72–2.35 mm, PL 0.65–0.66 mm, PH 0.53–0.60 mm, DPW 0.73–0.80 mm, GW 1.30–1.58 mm, CI 87–108, EI 24–25, SI 110–115, LPI 80–92, DPI 111–123.

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Diagnosis. Medium-sized, HW 0.96–1.04 mm, head in full-face view elliptical, clearly longer than broad; eye relatively large, weakly convex; mesosoma very elongated and subcylindrical; pronotum with tooth-like angles; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture almost absent; metanotal groove distinct with shallow depression; petiole in profile subtriangular, in dorsal view transverse and armed with two spines; first gastral tergite in dorsal view elliptical, much longer than broad, its posterior margin armed with numerous small denticles. Dorsa and lateral faces of head, mesosoma, petiole and first gastral tergite with dense macropunctures. Dorsa and lateral faces of head, mesosoma petiole, and first gastral tergite without long standing hairs; clypeus, mandible and first gastral sternite with sparse long hairs. Entire body black; antenna dark brown, leg reddish brown.

Description. Worker (fig. 17). Head in full-face view clearly longer than broad, elliptical, with weakly convex lateral margin and convex posterior margin. Eye relatively large, moderately protruding, located at mid-length of head. Frontal lobe horizontal, in full-face view covering antennal fossa only in part. Mandible short, subrectangular, anterior margin slightly convex, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight, while posterior margin strongly convex. Frontal carina short, not surpassing level of anterior margin of eye. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head; antennal segment II clearly narrower and longer than each of III–VI; terminal segment (XII) almost as long as X and XI combined.

Mesosoma very elongated and subcylindrical, promesonotum clearly larger than propodeum in dorsal view. In profile dorsal outline of mesosoma weakly convex; promesonotal suture absent, but laterally indicated by short row of puncture; metanotal groove weakly impressed; propodeal outline strongly convex dorsally; propodeal declivity weakly convex. In dorsal view mesosoma with waist-like incision in front of propodeum; promesonotum clearly longer than broad; pronotum with pair of tooth-like protrusions; maximum pronotal width slightly narrower than head 44

excluding eyes. In dorsal view, propodeum suboval, narrowed anteriorly, clearly longer than broad, with lateral and posterior margins convexs; proprodeal spiracle protruding laterally. Legs relatively long and slender.

Petiole in profile subtriangular, as long as high, ventral outline feebly concave. In dorsal view petiole transverse, about as broad as propodeal width, its anterior face as broad as posterior face. In posterior aspect dorsal margin with two spines; lateral margin without denticles or teeth.

First gastral tergite in dorsal view elliptical, much longer than broad, broader posteriorly, covering remaining tergites, posterior margin convex with numerous small denticles.

Dorsum and lateral face of head finely macropunctate, except frontal lobe punctate; mandible with dense puncture at base, smooth and shiny along masticatory margin and apex; antennal scape finely punctate. Mesosoma entirely macropunctate, except anterior-most portion punctate. Petiole and first gastral tergite macropunctate. Legs reticulate with smooth and shiny interspaces. Dorsum of head, mesosoma, petiole, and first gastral tergite without erect hairs; clypeus, mandible, and apex of scape with sparse erect hairs; gastral sternites with dense hairs.

Entire body black; mandible reddish brown; antenna dark brown, with tip of scape reddish brown and antennal segment XII yellowish brown; coxae, trochanters, and basal half of femora yellowish brown; apical half of femora, tibiae, and tarsi reddish brown.

Dealate gyne (fig. 18). Body size slightly larger than worker. Head in full- face view longer than broad, elliptical, with weakly convex side. Compound eye relatively large, moderately protruding, positioned at mid-length of head. Ocelli present, distance between lateral ocelli longer than distance between anterior and lateral ocellus. Mandible short, subrectangular, outer margin slightly convex, its masticatory margin armed with five teeth. Clypeus broad, its anterior margin almost 45

straight, while posterior margin strongly convex. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head. Mesosoma elongated. In dorsal view, pronotum short, separated from mesoscutum by a suture, its anterolateral corner armed with a short, truncate tooth; mesoscutum large, slightly longer than broad, its anterior margin strongly convex, while posterior margin almost straight; mesoscutellum trapezoidal, slightly broader than long, anterior margin weakly convex, separated from mesoscutum and metanotum by shallow sutures; metanotum very short, separated propodeum by a deep suture; propodeum larger than mesoscutellum, anterior margin feebly concave and posterior margin convex; in profile mesopluron broad, anepisternum separated from katepisternum by a transverse mesoplural groove; metapleuron not demarcated from lateral face of propodeum.

Petiole in profile view subtriangular, as long as high. In dorsal view petiole transverse, slightly narrower than propodeum. In posterior aspect dorsal margin with 2 spines laterally.

First gastral tergite in dorsal view elliptical, much longer than broad, broader posteriorly, covering remaining gastral tergites, posterior margin convex with numerous small denticles.

Sculpture, setae and colour condition similar to those of worker caste. Posterior portion of first gastral tergite with four short erect hairs.

Remarks. Echinopla jeenthongi belongs to the E. mezgeri group (Zettel & Laciny, 2017). It is closely related to E. mezgeri and E. tunkuabduljalilii. All species lack long standing hairs on the body surface. However, E. jeenthongi can be distinguished from E. mezgeri by the following characteristics: entire head, mesosoma, petiole and first gastral tergite finely macropunctate (finely punctate, except mesopleuron and lateral face of propodeum covered by a coarse punctuation in E. mezgeri); posterior margin of first gastral tergite with numerous small denticles 46

(without serration in E. mezgeri); basal half of femora yellowish brown (dark brown in E. mezgeri); sparse short pubescent hairs on dorsum of mesosoma (very dense in E. mezgeri). A third species of the E. mezgeri group from West Malaysia (Laciny et al., 2019) differs from both species by a strongly reduced puncturation of mesosoma and gaster tergite 1, and from E. jeenthongi also by a non-serrate hind margin of gaster tergite 1. This species can be separated from E. tunkuabduljalilii by dorsa of head, pronotum and first gastral tergite with dense macropunctures (very weak punctures in E. tunkuabduljalilii), posterior margin of first gastral tergite with numerous small denticles (without denticles in E. tunkuabduljalilii).

Habitat. Tanansathaporn et al. (2018) reported that E. jeenthongi was collected from lower vegetation in evergreen forest in southern Thailand. This species seem to be restricted to Sunderland.

Distribution. Thailand [Surat Thani, Nakhon Si Thammarat and Phangnga provinces (Tanansathaporn et al., 2018)].

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A

B C

Figure 17 Worker of Echinopla jeenthongi. A. body in profile view; B. head in full- face view and C. body in dorsal view.

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A

B C

Figure 18 Dealate qyne of Echinopla jeenthongi. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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Echinopla lineata Mayr, 1862 (Fig. 19)

Echinopla lineata Mayr, 1862: 689. Senior synonym of E. sucki: Emery, 1925: 211.

Echinopla lineata: Mayr, 1872: 143; Zettel & Laciny 2015: 116.

Echinopla sucki Forel, 1901: 75. Junior synonym of E. lineata: Emery, 1925: 211.

Type. Lectotype and two paralectotype workers (NHMW, not examined) from Indonesia, Java, Jakarta (as Batavia), 1857-1858, Novara Expedition (Mayr 1862).

Non-type material examined. One worker, S Thailand, Pattani Prov., Yaring District, Mangrove Forest, 19.IV.2002, C. Bourmas leg (THNHM); 1 worker, E Malaysia, Borneo, Sabah, Maliau Basin, 270-300 m alt., 8.XI.2011, Sk. Yamane leg. (THNHM); 1 worker, E Malaysia, Borneo, Sabah, Danum Valley, 30.VIII.1995, Sk. Yamane leg. (THNHM); 1 worker, Indonesia, Borneo, E Kalimantan, Kutai N.P., Sangkimah, 12.IX.1993, Sk. Yamane leg. (THNHM); 1 worker, Indonesia, Borneo, 14.VIII.1986, T. Yajima leg. (THNHM).

Measurements. Non-type worker (n = 5): TL 4.85–4.90 mm, HL 1.30–1.40 mm, HW 1.58–1.75 mm, SL 1.15–1.35 mm, ED 0.30–0.35 mm, PW 1.35–1.60 mm, MSL 1.75–2.05 mm, PL 0.35–0.65 mm, PH 0.40–0.65 mm, PW 1.15–1.30 mm, GW 1.70–1.90 mm, CI 111–131, EI 18–23, SI 69–84, LPI 80–114, DPI 246–386.

Diagnosis. Large-sized, HW 1.58–1.75 mm, head in full-face view broader than long; eye relatively large, moderately convex; dorsum of mesosoma distinctly laterally marginate along its length, armed with numerous teeth; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture present, narrow and shallow; metanotal groove distinct, deep impression; in profile view promesonotum and propodeum weakly convex; in posterior aspect, dorsal 50

margin of petiole feebly concave with numerous teeth and with long lateral spine on each side; in the same aspect lateral margin of petiole with 2–3 teeth; first gastral tergite globose, its posterior margin without denticles. Dorsa of head, mesosoma and first gastral tergite finely, longitudinally striate; lateral faces of pronotum and propodeum longitudinally striate. Entire body covered with dense white and long standing hairs mixed with pubescent. Entire body black without metallic lustre; apex of mandible, antennal segments X–XII, trochanters, and tarsi dark brown.

Description. Worker (fig. 19). Head in full-face view broader than long, trapezoidal, narrowed anteriorly, lateral margin and posterior margin weakly convex. Eye relatively large, moderately convex, located close to posterior corner of head. Frontal lobe obliquely elevated, in full-face view covering antennal fossa only in part. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight, while posterior margin strongly convex. Frontal carina short, not surpassing level of anterior margin of eye. Antenna 12-segmented; scape moderately long, slightly extending beyond posterior margin of head; funicular segments, each longer than broad.

Mesosoma stout, dorsum of mesosoma distinctly marginate at sides along its length, anterior and lateral margins of mesosoma armed with numerous denticles; propodeum subglobose, almost as long as promesonotum, but narrower. In profile dorsal outline of promesonotum and propodeum weakly convex; both promesonotum and propodeum meeting at deep and broad incision, the metanotal groove distinct; promesonotal suture present, narrow and shallow; mesopluron flat. In dorsal view mesosoma with waist-like incision in front of propodeum; pronotum trapezoidal, anterior margin slightly convex, posterior margin concave, lateral margin straight; pronotum slightly narrower than head width; mesonotum clearly shorter and narrower than pronotum. Legs relatively long and slender.

Petiole in profile subtriangular, as long as high, ventral outline feebly concave. In dorsal view petiole transverse, about as broad as propodeum, each side with strong 51

posteriorly bent spine; anterior face of petiole much narrower than posterior face of petiole. In posterior aspect dorsal margin with numerous teeth; lateral margin of with 2–3 teeth.

First gastral tergite in dorsal view globose, slightly broader than long, posterior margin roundly convex without denticles.

Dorsa of head, mesosoma and first gastral tergite finely, longitudinally striate; lateral faces of pronotum, mesopleuron, metapleuron and propodeum longitudinally striate; legs densely reticulate; antennal scape punctate.

Entire body covered with dense white and long standing hairs mixed with pubescent.

Entire body black without metallic lustre; apex of mandible, antennal segments X–XII, trochanters, and tarsi dark brown.

Remarks. Echinopla lineata belongs to the E. striata group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. striata and E. charernsomi by having a longitudinal striation on the dorsal body surface. However, E. lineata can be distinguished from E. striata by lateral face of propodeum striate (smooth and shiny in E. striata) and body covered with dense white standing hair and dense pubescence (body covered with blackish standing hairs and sparse pubescence in E. striata). See “Remarks” under E. charernsomi for separating E. lineata from E. charernsomi.

Habitat. The non-type material examined from South Thailand, Pattani Province was collected from mangrove forest, while the non-type specimens from Borneo were collected in rainforest. Thus, this species might inhabit primary forests and is restricted in Sundaland.

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Distribution. India (Bharti & Gul, 2012; Xu & Zhou, 2015), Thailand (Pattani province, new record), Malaysia (Zettel & Laciny, 2015), Singapore (Zettel & Laciny, 2015) and Indonesia [type locality, (Mayr, 1862; Zettel & Laciny, 2015)].

A

B C

Figure 19 Worker of Echinopla lineata. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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Echinopla madli Zettel et Laciny, 2015 (Figs. 20, 21)

Echinopla madli Zettel & Laciny, 2015: 103, figs. 1-4.

Type. Holotype worker (NHMW, Antweb images examined), S Thailand, Satun Province, Thale Ban National Park, 10–16.III.1993, leg. Michael Madl. Paratype worker (CASC, not examined), Borneo, Sabah, Danum Valley, Field Centre, 525 m a.s.l., 4.963056°N; 117.802806°E, 16–26.VIII.2010, rainforest, Ant Course 2010, unknown collector.

Non-type material examined. Two workers, S Thailand, Ranong Prov., Khlong Naka W.S., evergreen forest, 12.VIII.2009, W. Jaitrong leg (THNHM).

Measurements. Non-type worker (n = 2): TL 3.00–3.50 mm, HL 0.70–0.85 mm, HW 0.90–1.00 mm, SL 0.75–1.00 mm, ED 0.15–0.20 mm, PW 0.85–1.05 mm, MSL 1.15–1.25 mm, PL 0.40–0.55 mm, PH 0.35–0.40, PW 0.80–1.00, GW 1.05– 1.15, CI 106–143, EI 17–21, SI 83–100, LPI 73–89, DPI 164–233.

Diagnosis. Small-sized, HW 0.90–1.00 mm, head in full-face view slightly broader than long; eye relatively large; dorsum of mesosoma distinctly laterally marginate along its length, armed with numerous teeth; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture indistinct; metanotal groove distinct, deep impression; in profile mesosoma relatively short, promesonotun and proprodeum weakly convex; in posterior aspect, dorsal margin of petiole armed with 6 small teeth; lateral margin of petiole with 3–4 large teeth; first gastral tergite globose, its posterior margin with denticles. Body surface polyporous, with closely set pores, shiny interspaces. Dorsal surfaces of head, mesosoma, petiole, and first gastral tergite with grey standing hairs. Body black; leg reddish brown to dark brown; antenna dark brown.

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Description. Worker (fig. 20). Head in full-face view slightly broader than long, subtrapezoidal, with slightly convex lateral margin and weakly convex posterior margin. Eye relatively large, moderately protruding; located close to posterior corner of head. Frontal lobe chiefly horizontally oriented, completely covering antennal fossa only in part. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin weakly convex. Frontal carina short. Antennal scape moderately long, weakly s-curved, steadily widened from base to apex; antennal segments IIX–X slightly wider than long; terminal segment (XII) almost as long as X and XI combined.

Mesosoma stout, dorsum of mesosoma distinctly laterally marginate along its length, length roughly 1.2 times pronotum width; propodeum slightly larger than promesonotum. In profile mesosoma relatively short, dorsal outline promesonotun and proprodeum weakly convex; promesonotal suture indistinct; metanotal groove distinct, deep impression. In dorsal view mesosoma with waist-like incision in front of propodeum; pronotum subrectangular, anterior margin convex, lateral margin straight; pronotum about as broad as head excluding eyes; propodeum subglobose, about as broad as pronotum; anterior and lateral margins of mesosoma armed with numerous teeth. Legs moderately long; femora not much widened.

Petiole in profile subtriangular, longer than high, and narrowed dorsally. In dorsal view petiole transverse, about as broad as propodeum. In posterior aspect dorsal margin with 6 small teeth; lateral margin of petiole with 3–4 large teeth.

First gastral tergite in dorsal view globose, about as broad as long, and strongly covering the following tergites, posterior margin moderately convex and with denticles.

Head dorsally and laterally polyporous, with closely set pores. Petiole and first gastral tergite polyporous with smooth and shiny interspaces.

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Dorsal surfaces of head, mesosoma, petiole, and first gastral tergite with dense grey standing hairs.

Body black; antenna dark brown; legs reddish brown to dark brown.

Dealate qyne (fig. 21). Body size slightly larger than worker. Head in full- face view, subtrapezoidal, slightly broader than long, with weakly convex side. Compound eye relatively large, moderately protruding, located close to posterior corner of head. Ocelli present, distance between lateral ocelli longer than distance between anterior and lateral ocellus. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, its anterior margin weakly convex while posterior margin strongly convex. Antennal scape moderately long.

Mesosoma stout; in dorsal view pronotum short, separated from mesoscutum by a suture; mesoscutum large, slightly longer than broad, its anterior margin strongly convex while posterior margin almost straight; mesoscutellum trapezoidal, slightly broader than long, anterior and posterior margins weakly convex, separated from mesoscutum and metanotum by shallow sutures; metanotum very short, separated from propodeum by a deep suture; propodeum larger than mesoscutellum, its anterior margin almost straight and posterior margin weakly convex; in profile mesopluron large, anepisternum separated from katepisternum by indistinct transverse mesopluron groove.

Petiole in profile subtriangular, longer than high, and narrowed dorsally. In dorsal view petiole transverse, about as broad as propodeum. In posterior aspect, dorsal margin with 6 small teeth, lateral margin of petiole with 2–3 large teeth.

First gastral tergite in dorsal view globose, about as broad as long, and strongly covering the following tergites, posterior margin moderately convex and with denticles.

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Sculpture, setae and colour condition similar to those of worker caste.

Remarks. Echinopla madli belongs to the E. serrata group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. fisheri by having dense macropunctures or cribrate on the body surface. However, E. madli can be distinguished from E. fisheri by smaller body size (HW 0.90–1.00 mm in E. madli; HW 1.15–1.25 mm in E. fisheri), dorsal outlines of promesonotum and propodeum weakly convex, separating them with shallow metanotal groove (strongly convex and separating them by deep metanotal groove in E. fisheri), head dorsally and laterally polyporous, with closely set pores (with unevenly large pores and smooth and shiny interspaces in E. fisheri), and dorsal margin of petiole with small teeth (dorsal margin of petiole with large teeth in E. fisheri).

Habitat. All specimens used in the study were collected individually from lower vegetation in rainforest (evergreen) in Peninsular Malaysia (Sundaland).

Distribution. Thailand (Satun province) [type locality (Zettel & Laciny, 2015)], Ranong province [new locality record for Thailand] and Malaysia (Zettel & Laciny, 2015).

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A

B C

Figure 20 Worker of Echinopla madli. A. body in profile view; B. head in full-face view and C. body in dorsal view.

58

Figure 21 Dealate qyne of Echinopla madli. A. body in profile view; B. head in full- face view and C. body in dorsal view.

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Echinopla melanarctos Smith, 1857 (Figs. 22, 23)

Echinopla melanarctos Smith, 1857: 79, pl. 1, figs 25-29.

Echinopla melanarctos: Bolton, 1995; Jaitrong & Nabhitabhata 2005: 22; Gnatzy & Maschwitz, 2006: 125; Xu & Zhou, 2015: 26; Zettel & Laciny 2015: 110; Zettel & Laciny 2017: 9.

Type. Holotype worker (OUMNH, not examined). Labelled “SINC” and with a Donisthorpe type-label.

Non-type material examined. One worker, S Thailand, Narathiwat Prov., Wang Dist., Bala-Hala W.S., 6.VIII.2000, A. Chaitae leg. (AMK); 1 delate queen, S Thailand, Narathiwat Prov., Wang Dist., Bala-Hala W.S., 22.II.2003, Y. Sittimul leg (AMK).

Measurements. Non-type worker (n = 2): TL 6.25–6.75mm, HL 1.50 mm, HW 2.10–2.45 mm, SL 1.55–1.60 mm, ED 0.25 mm, PW 1.95–2.00 mm, MSL 2.25– 2.40 mm, PL 0.50–0.65 mm, PH 0.5 mm, PW 1.50–1.75 mm, GW 2.40–2.50 mm, CI 140–163, EI 10–12, SI 65–74, LPI 77–100, DPI 269–390.

Diagnosis. Large-sized, HW 2.10–2.45 mm, head in full-face view clearly broader than long; eye relatively small, distinctly protruded; dorsum of mesosoma laterally immarginate; in dorsal view mesosoma not waist-like; promesonotal suture almost absent; metanotal groove weak; in profile mesosoma stout, relatively short, strongly convex dorsal outline; in posterior aspect, dorsal margin of petiole with numerous teeth; first gastral tergite globose, its posterior margin without denticles. Dorsa of head, mesosoma and first gastral tergite with dense blunt spines; lateral face of head regularly longitudinally striate; legs and antennal scape black.

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Description. Worker (fig. 22). Head in full-face view clearly broader than long; lateral margin weakly convex and posterior margin bearing small medial lobe; eye relatively small, distinctly protruded, located close to posterior corner of head. Frontal lobe large, covering antennal socket. Mandible short, subrectangular, its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight. Frontal carina short, not surpassing level of anterior margin of eye. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head; antennal segment II clearly narrower and longer than each of III-XI.

Mesosoma stout. In profile view, dorsal outline of mesosoma strongly convex and forming a complete arch, not interrupted by impressed or notched metanotal groove; in dorsal view, mesosoma slightly constricted at metanotal groove position; promesonotal suture almost absent; metanotal groove weak. In dorsal view mesosoma not waist-like incision in front of propodeum; mesosoma immarginate laterally; pronotum narrower than head; propodeum subglobose, as broad as pronotum. Dorsa of head, mesosoma and first gastral tergite well developed long bluntly spines. Legs relatively long and slender.

Petiole in profile subtriangular, longer than high. In dorsal view petiole transverse, about as broad as propodeum, each side with strong posteriorly bent spine; anterior face of petiole much narrower than posterior face of petiole. In posterior aspect dorsal margin of petiole with numerous teeth and lateral margin without teeth.

First gastral tergite in dorsal view globose, covering remaining tergites, slightly broader than long, its posterior margin convex without denticles.

Dorsa of head, mesosoma and first gastral tergite with dense blunt spines. Lateral face of head regularly longitudinally striate.

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Body black; legs, mandible and antenna dark brown, antennal segment XII and tarsi reddish brown.

Dealate qyne (fig. 23). Similar to that of worker caste but body size being larger. Head in full-face view clearly broader than long; lateral margin weakly convex; eye relatively small, distinctly protruded, located close to posterior corner of head. Ocelli present. Mandible short, subrectangular, its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head.

Mesosoma stout. In dorsal view, pronotum short, separated from mesoscutum by a suture; mesoscutum large, trapezoidal, its anterior margin strongly convex, while posterior margin almost straight; mesoscutellum slightly broader than long, anterior margin weakly convex, separated from mesoscutum and metanotum by shallow sutures; metanotum very short, separated from propodeum by a deep suture; propodeum larger than mesoscutellum, anterior margin straight and posterior margin weakly convex; in profile mesopluron large, anepisternum separated from katepisternum by transverse mesopluron groove.

Petiole in profile subtriangular, longer than high. In dorsal view petiole transverse, about as broad as propodeum. In posterior aspect dorsal margin with numerous teeth; lateral margin without teeth.

First gastral tergite in dorsal view globose, covering remaining tergites, slightly broader than long, its posterior margin convex without denticles.

Sculpture, setae and colour condition similar to those of worker caste; lower portion of katepisternum smoother than in worker.

Remarks. Echinopla melanarctos belongs to the E. melanarctos group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. pallipes by having dense blunt 62

spines on the dorsa of head and mesosoma. However, E. melanarctos can be distinguished from E. pallipes by much larger size (HW 2.10–2.45 mm in E. melanarctos, HW 1.55–1.75 mm in E. pallipes); dorsa of head, mesosoma and first gastral tergite well developed long bluntly spines (weakly developed short blunt spines in E. pallipes); lateral face of head regularly longitudinally striate (irregularly rugose in E. pallipes).

Habitat. No biological information is available for E. melanarctos. However, judging from the specimen examined this species inhabits tropical rain forest. All specimens of the species were collected from lower vegetation.

Distribution. Thailand [Narathiwat province, (Jaitrong & Nabhitabhata, 2005)], Malaysia (Zettel & Laciny, 2015), Singapore [type locality (Smith, 1857)] and Indonesia (Zettel & Laciny, 2015).

A

B C

Figure 22 Worker of Echinopla melanarctos. A. body in profile view; B. head in full- face view and C. body in dorsal view. 63

A

B C

Figure 23 Dealate gyne of Echinopla melanarctos. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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Echinopla pallipes Smith, 1857 (Fig. 24)

Echinopla pallipes Smith, 1857: 80.

Echinopla pallipes: Forel 1913: 133; Bolton 1995: 183; Jaitrong and Nabhitabhata 2005: 22; Gnatzy & Maschwitz 2006: Xu & Zhou, 2015: 26; Zettel & Laciny, 2015: 110.

Type. Holotype worker (not examined). Labelled “SAR. 13” and with a Donisthorpe type-label.

Non-type material examined. One worker, S Thailand, Narathiwat Prov., Tropical Rainforest, Bala-Hala W.S., 24.II.2002, S. Hasin leg (AMK); 4 workers, same locality, 20.X.2003, Y. Sittimul leg (AMK).

Measurements. Non-type workers (n = 5): TL 5.25–5.85 mm, HL 1.10–1.30 mm, HW 1.55–1.75 mm, SL 1.15–1.40 mm, ED 0.20–0.25 mm, PW 1.50–1.65 mm, MSL 1.90–2.00 mm, PL 0.50–0.65 mm, PH 0.35–0.45 mm, PW 0.90–1.05 mm, GW 1.85–1.95 mm, CI 124–141, EI 13–16, SI 74–85, LPI 69–90, DPI 254–330.

Diagnosis. Large-sized, HW 1.55–1.75 mm, head in full-face view broader than long; eye relatively small, distinctly protruded; dorsum of mesosoma immarginate laterally; in dorsal view, mesosoma not waist-like; promesonotal suture and metanotal groove indistinct; in profile dorsum of mesosoma strongly convex and forming a complete arch; first gastral tergite globose, its posterior margin without denticles. Dosa of head and mesosoma with weakly developed short blunt spines, first gastral tergite densely cribrate; lateral face of head irregularly rugose. Body black; legs yellowish brown; antenna reddish brown.

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Description. Worker (fig. 24). Head in full-face view globose, almost as long as broad, lateral margin weakly convex and posterior margin bearing small medial lobe; eye relatively small, distinctly protruded, located close to posterior corner of head. Frontal lobe chiefly horizontally oriented, covering antennal socket. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight. Frontal carina short, not surpassing level of anterior margin of eye. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head; antennal segment II clearly narrower and longer than each of segment III–X.

Mesosoma stout. In profile view, dorsal outline of mesosoma strongly convex and forming a complete arch, not interrupted by impressed or notched metanotal groove; in dorsal view, mesosoma slightly constricted at metanotal groove position. In dorsal view mesosoma not waist-like incision in front of propodeum; mesosoma immarginate laterally; pronotum narrower than head; propodeum subglobose, about as broad as pronotum, longer than promesonotum, and its posterior margin convex. Legs relatively long and slender.

Petiole in profile view, triangular, as broad as high, and narrowed dorsally. In dorsal view, petiole transverse, about as broad as propodeum. In posterior aspect, dorsal margin with numerous teeth, while lateral margin without teeth.

First gastral tergite in dorsal view globose, as long as broad, covering remaining tergites, its posterior margin without denticles.

Dorsa of head and mesosoma with weakly developed short blunt spines, first gastral tergite densely cribrate; lateral face of head irregularly rugose.

Body black; legs yellowish brown; antenna and mandible reddish brown.

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Remarks. Echinopla pallipes belongs to the E. melanarctos group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. melanarctos. See “Remarks” under E. melanarctos.

Habitat. This species inhabits lowland rainforest and might be restricted to Sundaland.

Distribution. Thailand (Jaitrong & Nabhitabhata, 2005), Malaysia [type locality (Smith, 1857; Zettel & Laciny, 2015), Indonesia (Zettel & Laciny, 2015) and Philippines (General & Alpert, 2012).

A

B C

Figure 24 Worker of Echinopla pallipes. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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Echinopla striata Smith, 1857 (Figs. 25, 26, 27)

Echinopla striata Smith, 1857: 80.

Echinopla striata: Mayr 1862: 689; Mayr 1872: 143; Emery 1901: 721; Donisthorpe 194: 709; Bolton 1995: 184; Xu & Zhou 2015: 25.

Type. Syntype workers (OUMNH, Antweb images examined). Labelled “21 MAL.” and with a Donisthorpe type-label.

Non-type material examined. One worker , E Thailand, Chanthaburi Prov., Nam Tok Troknong, Tropical Rainforest, 23.XI.2003, D. Wiwatwitaya leg (AMK); 4 workers and 3 males, S Thailand, Ranong Prov., Khlong Naka N.P., Evergreen Forest, 12.VIII.2009, W. Jaitrong leg., TH09-WJT-60 (THNHM); 1 worker and 3 males, S Thailand, Surat Thani Prov., Ban Na San Dist., Ban Lumphun, Tai Rom Yen N.P., 11.X.2011, W. Jaitrong leg., TH11-WJT-39 (THNHM); 1 worker, S Thailand, Surat Thani Prov., Wipawadee Dist., Khlong Yan, 30.XII.2001, W. Jaitrong leg (THNHM); 1 worker, S Thailand, Nakhon Si Thammarat Prov., Sichon Dist., Khao Nan N.P., Evergreen Forest, 12.XII.2007, W. Jaitrong leg., WJT07-TH1909 (THNHM); 1 worker, S Thailand, Trang Prov., Yantakhao Dist., Thung Khai B.G., 13.XI.2017, W. Jaitrong leg., (THNHM); 1 worker, S Thailand, Song Khla Prov., Ton Nga Chang W.S., 13.IX.2004, N. Noon-anant leg (THNHM); 1 worker, S Thailand, Narathiwat Prov., Bala-Hala W.S., 15.IX.2001, S. Hasin leg (AMK); 1 worker, same locality and collector, 6.VI.2001 (AMK); 1 worker, same locality, 23.II.2003, Y. Sittimul (AMK); 23 workers and 1 dealate gyne, same locality, 30.V.2018, W. Jaitrong leg., WJT300618-2 (THNHM).

Measurements. Non-type worker (n = 10): TL 4.75–4.85 mm, HL 1.30–1.50 mm, HW 1.70–1.80 mm, SL 1.30 –1.40 mm, ED 0.25–0.30 mm, PW 1.60–1.70 mm, 68

MSL 2.00–2.15 mm, PL 0.56–0.70 mm, PH 0.50–0.55 mm, PW 1.30–1.50 mm, GW 1.80–1.92 mm, CI 120–131, EI 14–18, SI 75.00–82.35, LPI 76–93, DPI 243–295.

Diagnosis. Large-sized, HW 1.70–1.80 mm, head in full-face view broader than long; eye relatively large, moderately convex; dorsum of mesosoma distinctly laterally marginate along its length, armed with numerous teeth; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture narrow; metanotal groove present, with deep impression; in profile view, promesonotum weakly convex, while propodeum strongly convex; in posterior aspect, dorsal margin of petiole with 6–7 denticles, lateral margin with 2–3 denticles; first gastral tergite globose, its posterior margin without denticles. Dorsa of head, mesosoma and first gastral tergite finely, longitudinally striate. Lateral face of propodeum smooth and shiny. Dorsa of head, mesosoma, petiole, and first gastral tergite with dense black and long standing hairs mixed with dense pubescence. Entire body black with metallic lustre; trochanters black to dark brown.

Description. Worker (fig. 25). Large-sized, HW 1.70–1.80 mm, with head in full-face view roughly trapezoidal, narrowed anteriorly and broader than long, posterior corners rounded, lateral margin convex. Eye relatively large, moderately convex, located close to posterior corner of head. Frontal lobe narrow and elevated, lateral margin straight, in full-face view covering antennal socket only in part. Mandible short, subrectangular, and its masticatory margin armed with five teeth. Clypeus broad, anterior margin almost straight, posterior margin convex. Frontal carina short, not passing level of anterior margin of eye. Antenna 12-segmented; scape moderately long and slender.

Mesosoma stout, dorsum of mesosoma distinctly laterally marginate along its length, and armed with numerous teeth. In dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture narrow and shallow; metanotal groove present, with deep impression; pronotum trapezoidal or subrectangular, anterior margin convex, posterior margin concave, lateral margin straight; pronotum narrower than head; mesonotum clearly shorter and narrower than pronotum; 69

propodeum subglobose, as broad as pronotum, and longer than mesonotum; anterior and lateral margins of mesosoma armed with numerous teeth. In profile dorsal outline of promesonotum weakly convex, while propodeum strongly convex; both promesonotum and propodeum meeting at deep and broad incision, the metanotal groove.

Petiole in profile subtriangular, longer than high, and narrowed dorsally. In dorsal view transverse, about as broad as propodeum. In posterior aspect dorsal margin with 6–7 denticles; lateral margin with 2–3 denticles.

First gastral tergite in dorsal view globose, covering remaining tergites, narrowed posteriorly, its posterior margin without denticles.

Dorsa of head, mesosoma and first gastral tergite finely, longitudinally striate; lateral faces of pronotum, mesopleuron and metapleuron without longitudinal striation; lateral face of propodeum smooth and shiny.

Dorsa of head, mesosoma, petiole, and first gastral tergite with dense black and long standing hairs mixed with dense white pubescence.

Entire body black with metallic lustre; mandible, antennal segments X–XII, trochanters black to dark brown.

Male (fig. 26). Body size smaller than that in worker. Head in full-face view slightly broader than long; eye relatively large, moderately convex, three ocelli present, located close to posterior margin when seen in full-face view.

In dorsal view, pronotum narrow, its anterior margin strongly convex while posterior margin straight; mesoscutum trapezoidal, anterior margin strongly convex, posterior margin straight, separated from mesoscutellum by deep groove and separated from pronotum by transverse suture; parapsidal line distinct running to mid- length of mesoscutum; metanotum very short, separated from mesoscutellum by 70

shallow groove and from propodeum by deep but narrow grove; propodeum relatively long, in profile view smoothly convex; mesopluron large, anepisternum demarcated from katepisternum by a suture.

Petiole subtriangular, longer than high, and narrowed dorsally. In dorsal view transverse, clearly narrower than propodeum. In posterior aspect dorsal and lateral margins without denticles.

Gaster in dorsal view suboval, slightly longer than broad, narrowed posteriorly. First gastral tergite not covering other segments.

Dorsa of head, mesosoma and gastral tergites without longitudinal striation; lateral faces of pronotum and mesopleuron with indistinct longitudinal striation.

Dorsa of head, mesosoma, petiole, and gastral tergites with sparse yellowish standing hairs

Dorsal surfaces of head and mesosoma black; propodeum, petiole, gaster, mandible and leg dark brown; antennal segments XI–XII reddish brown.

Dealate qyne (fig. 27). Body size slightly larger than that in worker. Head in full-face view roughly trapezoidal, narrowed anteriorly and broader than long, its posterior corner rounded, and lateral margin convex. Eye relatively large, moderately convex, located close to posterior corner of head. Ocelli present, almost equal in size, located closed to posterior of head, distance between lateral ocelli longer than distance between anterior and lateral ocellus.

Mesosoma stout. In dorsal view, pronotum short, its anterior margin convex while posterior margin strongly concave, and separated from mesoscutum by a suture; mesoscutum large, clearly shorter than broad, its anterior margin strongly convex, white posterior margin almost straight; mesoscutellum elliptical, slightly broader than long, anteriorly margin weakly convex, separated from mesoscutum and metanotum 71

by shallow sutures; metanotum very short, separated propodeum by a deep suture; propodeum broad than mesoscutum, anterior margin almost straight and posterior margin weakly convex; in profile anepisternum separated from katepisternum by a transverse mesoplural groove.

Petiole in profile subtriangular, longer than high, and narrowed dorsally. In dorsal view, transverse, about as broad as propodeum. In posterior aspect dorsal margin with 6–7 small denticles; lateral margin with 2–3 small denticles.

First gastral tergite in dorsal view globose, covering remaining tergites, narrowed posteriorly, its posterior margin without denticles.

Sculpture, setae and colour condition similar to those of worker caste.\

Remarks. Echinopla striata belongs to the E. striata group (sensu Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. charernsomi and E. lineata by having fine longitudinal striation on the dorsal body surface. However, E. striata can be distinguished from E. charernsomi by the following characteristics: much larger body size (HW 1.72–1.85 mm in E. striata; HW 1.40–1.60 mm in E. charernsomi), smooth and shiny lateral face of propodeum (irregularly reticulate-punctate in E. charernsomi), and black to dark brown trochanters (yellowish brown in E. charernsomi). Echinopla striata is separated from E. lineata by lateral faces of pronotum, mesopleuron and metapleuron without longitudinal striation (longitudinally striate in E. lineata); dorsa of head, mesosoma, petiole, and first gastral tergite with dense black and long standing hairs (dense white hairs in E. lineata).

Habitat. Xu and Zhou (2015) recorded a specimen that was collected from the highland semi-evergreen monsoon forest in Yunnan, China. Specimens examined in this study were collected from lowland evergreen forest in eastern and southern Thailand and western Malaysia. Thus, this species can be found in both lowland and highland. A colony from Narathiwat province, South Thailand (WJT300618-2) was 72

found to nest in rotting twig on shrub at about 2 meters above the ground in an evergreen forest.

Distribution. China (Xu & Zhou, 2015), Thailand (Chanthaburi, Ranong, Surat Thani, Nakhon Si Thammarat, Trang, Song Khla, Narathiwat provinces, new record), Malaysia [type locality (Smith, 1857; Xu & Zhou, 2015)], Indonesia (Xu & Zhou, 2015) and Philippines (Forel, 1901).

A

B C

Figure 25 Worker of Echinopla striata. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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A

C B

Figure 26 Male of Echinopla striata. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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A

B C

Figure 27 Dealate gyne of Echinopla striata. A. body in profile view; B. head in full- face view and C. body in dorsal view.

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Echinopla tritschleri Forel, 1901 (Fig. 28)

Echinopla tritschleri Forel, 1901: 74.

Echinopla tritschleri: Forel, 1901: 74; Bolton 1995: 184; Xu & Zhou, 2015: 27; Zettel & Laciny 2015: 113; Zettel & Laciny 2017: 16.

Type. Syntype workers (MHNG and NHMB, Antweb images examined), Indonesia, West [Indrapura].

Non-type material examined. Eleven workers, S Thailand, Nakhon Si Thammarat Prov., Noppitum Dist., Khao Luang N.P., 28.V.2003, W. Jaitrong leg., TH03-WJT-321 (THNHM).

Measurements. TL 5.15–6.00 mm, HL 1.65–1.80 mm, HW 1.55–1.60 mm, SL 1.35–1.45 mm, ED 0.25–0.45 mm, PW 1.35–1.45 mm, MSL 2.00–2.25 mm, PL 0.50–0.60 mm, PH 0.60–0.70 mm, PW 0.75–0.80 mm, GW 1.75–1.85 mm, CI 119– 141, EI 16–29, SI 84–94, LPI 109–140, DPI 225–290.

Diagnosis. Large-sized, HW 1.55–1.60 mm, head in full-face view slightly longer than broad; eye relatively large, moderately convex, located close to mid- length of head; dorsum of mesosoma laterally immarginate; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture weak; metanotal groove distinct with deep impression; in posterior aspect, dorsal margin of petiole with 5-6 distinct teeth; first gastral tergite subglobose, slightly longer than broad, its posterior margin without denticles, anterior portion with few tubercles, sides and disc with punctures that flow together to form longitudinal lines and posterior portion with punctures that flow together into transverse lines. Frons and dorsum of mesosoma with longer and slender tubercles, their apex truncate and bearing a long yellowish hair. Body black; legs yellowish brown; mandible and antenna dark brown. 76

Description. Worker (fig. 28). Head in full-face view rounded, slightly longer than broad. Eye relatively large, moderately convex, located close to mid- length of head. Frontal lobe obliquely elevated. Frontal carina reaching level of anterior margin of eye. Scape moderately long and slender; terminal segment (XII) almost as long as X and XI combined.

Mesosoma stout, dorsum of mesosoma laterally immarginate. In dorsal view, mesosoma with waist-like incision in front of propodeum; pronotum trapezoidal or subrectangular, and narrower than head; promesonotal suture weak; metanotal groove distinct with deep impression; propodeum subglobose, narrowed anteriorly, and as broad as pronotum. In profile view, dorsum of mesosoma almost straight, not forming a complete arch, interrupted by impressed or notched metanotal groove.

Petiole in profile subtriangular, almost as long as high, ventral outline straight. In dorsal view, petiole transverse, slightly narrower than propodeum. In posterior aspect, dorsal margin with 5–6 distinct teeth.

First gastral tergite in dorsal view globose, covering remaining tergites, slightly longer than broad, its posterior margin without denticles, anterior portion with few tubercles, sides and disc with dense punctures that flow together to form longitudinal lines and posterior portion with punctures that flow together into transverse lines.

Dorsa of head and mesosoma with moderately long blunt spines, their apex truncate and bearing a long yellowish hair.

Body black; mandible and antenna dark brown; legs and gastral apex yellowish brown; eye blackish brown.

Remarks. Echinopla tritschleri belongs to the E. melanarctos group (Xu & Zhou, 2015; Zettel & Laciny, 2017) and is similar to E. cherapunjiensis by the 77

following characteristics: dorsa of head and mesosoma with dense blunt spine and in profile view, dorsum of mesosoma almost straight, not forming a complete arch. However, E. tritschleri can be distinguished from E. cherapunjiensis by dorsa of head and mesosoma with moderately long blunt spines (short blunt spines in E. cherapunjiensis); dorsal outline of mesosoma interrupted by distinctly impressed metanotal groove (narrowly notched metanotal groove in E. cherapunjiensis); lateral face of first gastral tergite with dense punctures that flow together to form longitudinal lines (densely cribrate or polyporous in E. cherapunjiensis).

Habitat. Zettel and Laciny (2015) recorded that this species was collected from lowland to highland (180-1700 m a.s.l.) rainforest (evergreen and hill evergreen forests) in Malaysia. The material examined from Nakhon Si Thammarat province, colony no. TH03-WJT-321 was collected from a large dead branch that fell from a high tree. Thus, this species might be a canopy ant.

Distribution. Thailand (Nakhon Si Thammarat Province, new record), Malaysia (Zettel & Laciny, 2015, 2017) and Indonesia (Forel, 1901).

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B C

Figure 28 Worker of Echinopla tritschleri. A. body in profile view; B. head in full- face view and C. body in dorsal view.

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Echinopla tunkuaduljalilii Laciny, Zettel, Maryati et Noor, 2019 (Fig. 29)

Echinopla tunkuaduljalilii Laciny, Zettel, Maryati & Noor 2019: 251, figs 1–5.

Type. Holotype worker (ITBC, not examined), Malaysia, Johor, Segamat Dist., Sg. Batang R.F., N 2°20’49’’, E. 103°09’25’’, 130 m a.s.l., A. Izwan-Noor leg. Paratypes. One worker (examined), S Thailand, Phangnga Prov., Muang Phangnga District, Song Prak Subdistrict, near Ton Periwat Waterfall, N 8°36'40", E 98°33'02", 360–380 m a.s.l., evergreen forest, lower vegetation, 25.V.2005. N. Noon-anant leg. (PSU); 1 worker, same locality and collector, 9.XI.2005, N. Noon-anant leg. (PSU); 1 worker, S Thailand, Nakhon Si Thammarat Prov., Noppitum District, Mount San Yen, 300–450 m a.s.l., evergreen forest, from lower vegetation, 18.VII.2005, N. Noon- anant leg (PSU).

Non-type material examined. One worker, S Thailand, Phangnga Prov., Takuarpa Dist., Sri Phangnga, Evergreen Forest, Lower Vegetation, 25.XII.2002, N. Noon-anant leg (PSU); 1 worker, S Thailand, Yala Prov., Julapon10, Betong 500-600 m, evergreen forest, lower vegetation, 9.II.2006, N. Noon-anant leg. (PSU).

Measurements. TL 4.25–4.80 mm, HL 1.15–1.20 mm, HW 0.95 mm, SL 1 mm, ED 0.25 mm, PW 0.95 mm, MSL 1.80–1.90 mm, PL 0.45–0.60 mm, PH 0.50– 0.55 mm, PW 0.80–1.30 mm, GW 1.30–1.35 mm, CI 79–83, EI 26, SI 105, LPI 83– 122, DPI 158–211.

Diagnosis. Medium-sized, HW 0.95 mm, head in full-face view elliptical, clearly longer than broad; eye relatively large, weakly convex; mesosoma very elongated and subcylindrical; pronotum with tooth-like angles; in dorsal view mesosoma with waist-like incision in front of propodeum; promesonotal suture absent; metanotal groove distinct with narrow and shallow depression; petiole in profile subtriangular, in dorsal view transverse and without teeth, with obtuse lateral corners; first gastral tergite in dorsal view elliptical, much longer than broad, its 80

posterior margin without denticles. Dorsa and lateral faces of head, mesosoma, petiole and first gastral tergite with very weak punctures. Dorsa of head, mesosoma, first gastral tergite and antennal scape without standing hairs; clypeus and mandible with sparse long hairs. Entire body black; antenna dark brown, mandible reddish brown, leg yellowish brown.

Description. Worker (fig. 29). Head in full-face view clearly longer than broad, elliptical, with lateral margin weakly convex and posterior margin convex. Eye relatively large, weakly convex, located at mid-length of head. Frontal lobe horizontal, in full-face view covering antennal socket only in part. Frontal carina short, not surpassing level of anterior margin of eye. Antennal scape moderately long, steadily widened from base to apex, slightly extending beyond posterior margin of head; terminal segment (XII) almost as long as X and XI combined.

Mesosoma very elongated and subcylindrical. In profile dorsal outline of mesosoma weakly convex; promesonotal suture absent, but laterally indicated by short row; metanotal groove weakly impressed. In dorsal view mesosoma with waist- like incision in front of propodeum; pronotum with a pair of tooth-like protrusions; maximum pronotal width slightly narrower than head excluding eyes; propodeum suboval, narrowed anteriorly, clearly longer than broad, with lateral and posterior margins convex. Legs relatively long and slender.

Petiole in profile subtriangular, as long as high, ventral outline straight. In dorsal view petiole transverse, about as broad as propodeal width. In posterior aspect dorsal margin without teeth.

First gastral tergite in dorsal view elliptical, much longer than broad, covering remaining tergites, its posterior margin without denticles.

Dorsa and lateral faces of head, mesosoma, petiole and first gastral tergite with very weak punctures; mesopleuron, metapleuron and anterior part of lateral face of propodeum with dense macropunctures. Dorsa of head, mesosoma, first gastral tergite 81

and antennal scape without standing hairs; clypeus and mandible with sparse long hairs.

Entire body black; mandible reddish brown, antenna dark brown, tip of scape reddish brown; coxae, trochanter, yellowish brown; femora, tibiae dark brown.

Remarks. Echinopla tunkuaduljalilii belongs to the E. mezgeri group (Zettel & Laciny, 2017). It is similar to E. jeenthongi by lacking long standing hairs on the body surface. See “Remarks” under E. jeenthongi.

Habitat. No biological information is available for E. tunkuaduljalilii. However, judging from the holotype from western Malaysia and specimen from the southernmost part of Thailand this species inhabits lowland and is probably restricted in Sundaland.

Distribution. Thailand (Phangnga and Yala provinces, new record) and Malaysia [type locality, (Laciny et al., 2019)].

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Figure 29 Worker of Echinopla tunkuaduljalilii. A. body in profile view; B. head in full-face view and C. body in dorsal view.

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Figure 30 Distribution map of the ant genus Echinopla in Thailand.

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

Species groups and species N NE E W C S

Echinopla melanarctos group

1. Echinopla cherapunjiensis / / / /

2. Echinopla melanarctos /

3. Echinopla pallipes /

4. Echinopla tritschleri /

Echinopla mezgeri group

1. Echinopla jeenthongi /

2. Echinopla tunhuabduljalilii /

Echinopla serrata group

1. Echinopla fisheri / / /

2. Echinopla madli /

Echinopla striata group

1. Echinopla charernsomi / /

2. Echinopla lineata /

3. Echinopla striata / /

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DISCUSSION AND CONCLUSION

Discussion

Until now, 36 species and three subspecies of the ant genus Echinopla have been known around the world. Among them, 11 species are found in Thailand and they belong to four species groups (Xu & Zhou, 2015): E. melanarctos group (E. cherapunjiensis, E. melanarctos, E. pallipes and E. tritschleri), E. mezgeri group (E. jeenthongi and E. tunkuabduljalilii), E. serrata group (E. fisheri and E. madli) and E. striata group (E. charernsomi, E. lineata and E. striata). Two new species (E. charernsomi and E. jeenthongi) were described in the present study.

Body sculptures; eye shape and position; shape of frontal lobe and shape of head, mesosoma, petiole and pilosity are characters were used by Xu and Zhou (2015) to distinguish the species groups mentioned above. These morphological characters were confirmed and used by several authors who described new species of the genus after Xu and Zhou (2015) (Laciny et al., 2019; Tanansathaporn et al., 2018; Zettel & Laciny, 2017). The E. melanarctos species group is characterized by dorsa of head, mesosoma and first gastral tergite with dense blunt spines; head clearly broader than long. The E. mezgeri species group is characterized by dorsa of head, mesosoma, and first gastral tergite macropunctate without the standing hairs; head clearly longer than broad. The E. serrata species group is separated from the other group by the following characteristics: dorsa of head, mesosoma and first gastral tergite with polyporous or cribrate; head almost as long as broad. Lastly, the E. striata species group is distinguished from the other groups by dorsa of head and mesosoma finely longitudinally striate; head broader than long. Sculptures on body, teeth or spines on petiole, shape of head, shape of body and pilosity are used to separate each species. The ant genus Echinopla is clearly monomorphic in worker caste with a small size variation occurring within the species. Thus, body size can be used for separating into large and small species. In this study body size has been used for separating Echiopla madli (HW 0.90–1.00 mm) from E. fisheri (HW 1.15–1.25 mm), both are in the same species group. 86

Members of the genus Echinopla are distributed in eastern India, China, various countries in Southeast Asia, New Guinea, and Australia. Twenty nine species are distributed in Southeast Asia. Among them, eleven species are found in Thailand (E. charerensomi, E. cherapunjiensi, E. fisheri, E. jeenthong, E. lineta, E. madli, E. pallipes, E. melanarctos, E. striata, E. trischleri and E. tunkuaduljalilii). Echinopla cherapunjiensis is the widely distributed occurring in western, northeastern, central and southern parts of the country. Seven species: E. jeenthongi, E. lineata, E. madli, E. melanarctos, E. pallipes, E. tritschleri and E. tunkuaduljalilii are restricted to the South (fig. 30). So far, no species of the genus have been recorded from northern Thailand. Xu and Zhou (2015) recorded E. cherapunjiensis from highland semi- evergreen monsoon forest in China, while Jaitrong, et al. (2016) found the species from Laos. Thus, based on the distribution pattern this species might occurs in northern part of Thailand.

According to available data of habitat types, Thai Echinopla species occur from the low to high altitude areas. These can be separated into two groups based on altitudinal distributional pattern: species of group I, eight species (E. charerensomi, E. fisheri, E. jeenthong, E. lineta, E. madli, E. pallipes, E. melanarctos and E. tunkuaduljalilii) are confined to lowland primary forests (evergreen and dry evergreen forests) (Laciny et al., 2019; Tanansathaporn et al., 2018; Zettel & Laciny, 2015, 2017). These species of the groups might be sensitive to these habitats. Species of group II, another three species (E. cherapunjiensi, E. striata and E. trischleri) occurs from the lowland to highland (Bharti & Gul, 2012; Jaitrong et al., 2016; Xu & Zhou, 2015).

Zettel and Laciny (2015) and Tanansathaporn et al. (2018) reported that specimens of the ant genus Echinopla were rarely collected because they have a relatively small colony size, typically nest on canopy and forage individually. Watanasit et al. (2005) examined the composition of canopy ants at Ton Nga Chang Wildlife Sanctuary, Songkhla province, southern Thailand being found five unidentified species of the genus. Seven species in the present study (E. charernsomi, E. fisheri, E. jeenthongi, E. lineata, E. madli, E. melanarctos and E. tunkuaduljalilii) 87

were captured individually from lower vegetation while another four species (E. cherapunjienssis, E. pallipes, E. striata and E. trischleri) were collected from nests in rotting woods or dead twigs, which have been fell from canopies. These point out that the ant genus Echinopla typically inhabits the canopy or lower vegetation.

The species of the genus were found in evergreen forests (dry evergreen, evergreen and hill evergreen) (Bharti & Gul, 2012; Tanansathaporn et al., 2018; Xu & Zhou, 2015; Zettel & Laciny, 2015). As in the present study, all species were collected from the same habitats. This genus has never been found in disturb areas, dry dipterocarp and mixed deciduous forests. So that, member of this genus might be used to be the biological indicators. However, no bionomic information of this genus has been studied in details.

Conclusions

A total of 11 species of the ant genus Echinopla are recognized in Thailand based on external morphological characters of the workers caste. They belong to the following four species groups: E. melanarctos group (E. cherapunjiensis, E. melanarctos, E. pallipes and E. tritschleri); E. mezgeri group (E. jeenthongi and E. tunhuabduljalilii); E. serrata group (E. fisheri and E. madli) and E. striata group (E. charernsomi, E. lineata and E. striata). Five species (E. cherapunjiensis, E. tritschleri, E. fisheri, E. lineata and E. striata) are recorded for the first time from Thailand. During the two years of the study, two new species have been added to the ant faunas of Thailand.

Keys to species groups and the Thai species based on the worker caste are provided. Complete description is presented for each species based on worker, male, dealate gyne if available.

The distribution map of the ant genus Echinopla in Thailand is given. Echinopla cherapunjiensis is the widely distributed occurring in western, 88

northeastern, central and southern parts of the country while E. jeenthongi, E. lineata, E. madli, E. melanarctos, E. pallipes, E. tritschleri and E. tunkuaduljalilii are apparently restricted to southern Thailand. So far, no species of the genus have been recorded from northern part.

The genus confined to evergreen forests (dry evergreen, evergreen and hill evergreen). So far, no species of the genus have been found in dry dipterocarp and mixed deciduous forests or in disturbed areas. Thus, members of the genus might be used as biological indicators.

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APPENDIX World distribution of the ant genus Echinopla based on political regions.

Scientific name Distribution

1. Echinopla angustata Zettel et Laciny, 2015 Indo-Australian Region

Philippines (type locality)

2. Echinopla arfaki Donisthorpe, 1943 Indo-Australian Region

New Guinea (type locality)

3. Echinopla australis Forel, 1901 Australasian Region

Australia (type locality)

Indo-Australian Region

New Guinea

4. Echinopla brevisetosa Zettel et Laciny, 2015 Indo-Australian Region

Philippines (type locality)

5. Echinopla charernsomi Tanansathaporn et Oriental Region Jaitrong, 2018 Thailand (type locality)

6. Echinopla cherapunjiensis Bharti et Gul, 2012 Indo-Australian Region

Malaysia

Oriental Region

India (type locality), Laos, Myanmar

Palaearctic Region

China

94

Appendix (Continued).

Scientific name Distribution

7. Echinopla circulus Zettel et Laciny, 2015 Indo-Australian Region

Borneo, Indonesia, Malaysia (type locality)

8. Echinopla crenulata Donisthorpe, 1941 Indo-Australian Region

New Guinea (type locality)

9. Echinopla deceptor Smith, 1863 Indo-Australian Region

Indonesia (type locality)

10. Echinopla densistriata Stitz, 1938 Indo-Australian Region

Borneo (type locality),

Indonesia, Malaysia

11. Echinopla dubitata Smith, 1862 Indo-Australian Region

Indonesia (type locality), Sulawesi

12. Echinopla fisheri Zettel et Laciny, 2015 Indo-Australian Region

Borneo, Indonesia, Malaysia (type locality)

13. Echinopla jeenthongi Tanansathaporn et Oriental Region Jaitrong, 2018 Thailand (type locality)

14. Echinopla lateropilosa Zettel et Laciny, 2017 Indo-Australian Region

Malaysia (type locality)

95

Appendix (Continued).

Scientific name Distribution

15. Echinopla lineata Mayr, 1862 Indo-Australian Region

Borneo, Indonesia (type locality), Krakatau Islands, Malaysia, Singapore

Oriental Region

India

16. Echinopla madli Zettel et Laciny, 2015 Indo-Australian Region

Borneo, Indonesia, Malaysia

17. Echinopla maeandrina Stitz, 1938 Indo-Australian Region

New Guinea (type locality)

18. Echinopla melanarctos Smith, 1857 Indo-Australian Region

Borneo, Indonesia, Malaysia, Singapore (type locality)

Oriental Region

Thailand

19. Echinopla mezgeri Zettel et Laciny, 2015 Indo-Australian Region

Borneo, Indonesia, Malaysia (type locality)

20. Echinopla mistura Smith, 1860 Indo-Australian Region

Indonesia (type locality)

96

Appendix (Continued).

Scientific name Distribution

21. Echinopla nitida Smith, 1863 Indo-Australian Region

Indonesia (type locality)

22. Echinopla octodentata Stitz, 1911 Indo-Australian Region

New Guinea (type locality)

23. Echinopla pallipes Smith, 1857 Indo-Australian Region

Borneo (type locality), Indonesia, Malaysia

24. Echinopla praetexta Smith, 1860 Indo-Australian Region

Indonesia (type locality)

25. Echinopla pseudostriata Donisthorpe, 1943 Indo-Australian Region

New Guinea (type locality), Philippines

26. Echinopla rugose André, 1892 Indo-Australian Region

Borneo (type locality), Indonesia, Malaysia

27. Echinopla senilis Mayr, 1862 Oriental Region

India, Nicobar Island (type locality)

28. Echinopla serrata Smith, 1859 Indo-Australian Region

Indonesia (type locality)

97

Appendix (Continued).

Scientific name Distribution

29. Echinopla silvestrii Donisthorpe, 1936 Indo-Australian Region

New Guinea (type locality)

30. Echinopla striata Smith, 1857 Indo-Australian Region

Borneo, Indonesia, Malaysia (type locality), Philippines

Palaearctic Region China

31. Echinopla subtilis Zettel et Laciny, 2015 Indo-Australian Region

Borneo, Indonesia, Malaysia (type locality)

32. Echinopla tritschleri Forel, 1901 Indo-Australian Region

Borneo, Indonesia (type locality), Malaysia

33. Echinopla tunkuaduljalilii Laciny, Zettel, Indo-Australian Region Maryati et Noor, 2019 Malaysia (type locality),

Thailand

34. Echinopla turneri Forel, 1901 Australasian Region

Australia (type locality)

35. Echinopla vermiculata Emery, 1898 Indo-Australian Region

Indonesia (type locality), Philippines, Sulawesi

98

Appendix (Continued). Scientific name Distribution

36. Echinopla wardi Zettel et Laciny, 2015 Indo-Australian Region

Malaysia (type locality)

CURRICULUM VITAE

CURRICULUM VITAE

NAME Miss Soisunee Tanansathaporn

DATE OF BIRTH 21 July 1994

BIRTH PLACE Samutsongkhram, Thailand

EDUCATION Kasetsart University

SCHOLARSHIP Scholarship for International Publication from Graduate school, Kasetsart University