Research Article |
Corresponding author: Sopark Jantarit ( fugthong_dajj@yahoo.com ) Academic editor: Ľubomír Kováč
© 2021 Katthaleeya Surakhamhaeng, Louis Deharveng, Sopark Jantarit.
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Citation:
Jantarit S, Deharveng L, Surakhamhaeng K (2021) Three new species of cave Troglopedetes (Collembola, Paronellidae, Troglopedetinae) from Thailand, with a key to the Thai species. Subterranean Biology 40: 129-174. https://doi.org/10.3897/subtbiol.40.73143
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Thailand is today the richest country for the genus Troglopedetes Joseph, 1872, with 17 species described from the country. In this study three troglomorphic new species are described from caves in the western region. They are T. spectabilis sp. nov. and T. rungsimae sp. nov. from Kanchanaburi province and T. takensis sp. nov., from Tak province. The three new species share elongated antennae and have the same number of central mac on Th. III and Abd. IV. However, they differ from one another by the combination of: presence of eyes, antennal length, claw morphology, central head macrochaetotaxy and internal row of dental spines. Troglopedetes spectabilis sp. nov. is remarkably different from its congeners by its extreme long appendages, especially the antennae (ratio antenna: head = 5.5). In the second part of the paper the arrangment of antennal chaetotaxy and the diversity of its phaneres is analyzed in the three new species, homologized and compared with those of two other species described from Thailand (T. meridionalis and T. kae). A total of 22 types of chaetae have now been recognized among the species. A new type of S-chaetae for the genus was discovered in this study. There are 5 types of ordinary chaetae, 15 types of S-chaetae, the subapical organite of Ant. IV and scales. A total of 1,107 to 2,183 antennal chaetae on each side were observed, which includes 308–485 S-chaetae, 687–1,402 ordinary chaetae, 72–295 scales and a subapical organite.
Antennal phaneres, cave, chaetotaxy, Southeast Asia, subterranean habitat, troglomorphy
The genus Troglopedetes Joseph, 1872 is widespread in Thailand in both edaphic and subterranean environments, particularly in the western and northern regions of the country (
Distribution of Troglopedetes in Thailand (green circles) with limestone terrains (red colour) of the country 1 T. maffrei Deharveng & Gers, 1993 2 T. longicornis Deharveng & Gers, 1993 3 T. centralis Deharveng & Gers, 1993 4 T. fredstonei Deharveng, 1988 5 T. leclerci Deharveng, 1990 6 T. microps Deharveng & Gers, 1993 7 T. multispinosus Deharveng & Gers, 1993 8 T. maungonensis Deharveng & Gers, 1993 9 T. calvus Deharveng & Gers, 1993 10 T. dispersus Deharveng & Gers, 1993 11 T. convergens Deharveng & Gers, 1993 12 T. paucisetosus Deharveng & Gers, 1993 13 T. meridionalis Jantarit, Surakhamhaeng & Deharveng, 2020 14 T. kae Jantarit, Surakhamhaeng & Deharveng, 2020 15 T. spectabilis sp. nov. 16 T. takensis sp. nov. 17 T. rungsimae sp. nov. (blue circles) and undescribed species of Troglopedetes found in Thailand (black circles). Regional boundary indicated by grey line; province capitals as small dark dots; scale 1:250,000.
Troglomorphic features in Collembola mostly comprise: large body size, elongated appendages (antennae, legs and furca), elongated and slender claw complex, pointed tenent hair, multiplication of antennal chaetae, blindness and depigmentation (
During our sampling campaigns in limestone caves in Thailand, several Troglopedetes species were collected. Three of them, which were found in caves in the western region (Tak and Kanchanaburi provinces), are described in this study. One of them exhibits strikingly long antennae, not seen in any other species of the genus (ratio of antennae: head > 3 times; antennae: body ≥ 1).
We have already described a high diversity of phaneres on antennae of two species of Troglopedetes (
The specimens of the three new species were collected in the dark zone of caves in Tak and Kanchanaburi Province (Fig.
Ant. antennal segment;
Abd. abdominal segment;
AIIIO apical organ of Ant. III;
Th. thoracic segment;
mac macrochaetae;
mes mesochaetae;
mic microchaeta;
psp pseudopore;
Tita tibiotarsus;
tric trichobothria;
ms S-microchaeta(e);
s or sens S-chaeta(e);
VT ventral tube;
NHM-PSU Princess Maha Chakri Sirindhorn Natural History Museum, Prince of Songkla University, Songkhla, Thailand.
Pseudopore arrangement follows
Subfamily Troglopedetinae Börner, 1913
Thailand, Kanchanaburi province, Sai Yok district, Tham (cave) Morakhot (14°11'02.7"N, 99°01'32.8"E, 426 m a.s.l.).
Holotype : female on slide, Kanchanaburi province: Sai Yok district, Tham (cave) Morakhot, 14°11'02.7"N, 99°01'32.8"E, 426 m a.s.l., 28 February 2019, S. Jantarit, A. Nilsai and K. Jantapaso leg., dark zone of cave, by aspirator (sample # THA_SJ_KRI12). Paratypes: 5 paratypes on slides (all subadults), same locality and date as the holotype.
Holotype and 5 paratypes deposited in NHM-PSU, measurements of holotype in Table
Head | Tergites | Appendages | |||
---|---|---|---|---|---|
Ant. I | 374 | Th. II | 230 | Man | 560 |
Ant. II | 667 | Th. III | 180 | Dens | 736 |
Ant. III | 720 | Abd. I | 100 | Mucro | 53 |
Ant. IVa | 595 | Abd. II | 120 | Furca | 1,349 |
Ant. IVb | 424 | Abd. III | 135 | Claw I | 64 |
Ant. | 2,780 | Abd. IV | 550 | Claw II | 64 |
Head | 500 | Abd. V | 90 | Claw III | 65 |
Abd. VI | 70 | ||||
Body | 1,475 |
Habitus. Highly troglomorphic, slender, with elongate legs, furca and antennae (Figs
Troglopedetes spectabilis sp. nov. A habitus B papilla E of labial palp C ventro-distal complex of labrum D antenna E head chaetotaxy (left = A to G mac nomenclature; right = AMS nomenclature) F Clypeus G outer maxillary lobe H labial basis and ventral chaetotaxy of head, right side I labrum.
Chaetal types. Four types of chaetae on somites, appendages (except antennae) and mouthparts: scales, present on antennal segment I and II, head, body and ventral side of the furca, absent on legs and ventral tube; ordinary chaetae on all body parts; S-chaetae and trichobothria on tergites; hairs devoid of sockets on outer maxillary lobe. Chaetal types on antennae are much more diverse and described separately further.
Pseudopores
(Figs
Troglopedetes spectabilis sp. nov., continued A distal part of tibiotarsus I and claw complex with clavate tenent hair B distal part of tibiotarsus III and claw complex with clavate tenent hair C ventral side of claw complex D lateral flap and antero-distal face of ventral tube E trochanteral organ F female genital plate G distal part of Manubrium and Mucrodens H mucro.
Mouthparts. Clypeus with 11 mac, symmetrically arranged: 9 ciliated posteriorly and 2 smooth anteriorly (Fig.
Antennae
(Figs
General chaetotaxy
(Figs
Dorsal macrochaetae formula: 4,2/9,4/0,2,4,3 (Figs
Head chaetotaxy. Head with 10–11 peri-antennal mac in line on each side, with 4+4 central mac (chaetae A, B, D, G of
Ventral chaetotaxy of head densely covered with oval scales (65–72 µm long), postlabial chaetae along the linea ventralis as 3 smooth mes anteriorly and 2 smooth chaetae laterally, one mac and an oblique line of 7–8 mes posteriorly on each side (Fig.
Tergite chaetotaxy. Th. II with a collar consisting of a few rows of mac along its anterior and antero-lateral margins, a compact group of 6 central mac on each side (“P3 complex” of
Th. III with 4 mac by side (a group of 3 central and 1 anterior to them), 1 sens at antero-lateral margins, and about 16 mac or long mes at lateral margins (Fig.
Abd. I without central mac, with 1 ms laterally on each side, a row of 3 mics below psp, a6 (sensu
Abd. II with 2 tric on each side and 6 modified mes around them (2 around the internal tric and 3–4 near external tric), 2 mac (1 near internal tric and 1 near external tric), 1 sens near internal tric, 2 mic close to external tric), at least 6 other mes sockets visible at lateral margins (not drawn) (Fig.
Abd. III with 3 tric on each side (1 internal, 2 external) and 9–10 modified mes around tric (3 near internal tric, 6–7 near the two external tric); 4 mac (1 near internal tric and 3 near external tric); 1 sens anterior to internal tric and ms not seen; at least 7 mes at lateral margins (Fig.
Abd. IV with 3 tric on each side (2 antero-lateral, 1 postero-lateral) and about 7–11 modified mes around the two antero-lateral tric; postero-lateral tric without modified mes. Mac distributed as 3 central on each side (1 antero-external to pseudopore, 2 anterior to posterior tergite margin), 1 near postero-lateral tric, and at least 4 external, mixed with at least 13 mes or smaller mac on lateral to posterior margins; 2 sens; at least 9 S-like chaetae sensu
Abd. V with 2 sens detected on each side, and several ordinary chaetae from mes to mac, not counted (Fig.
Legs
(Fig.
Ventral tube
(Fig.
Furca complex
(Fig.
Genital plate
(Fig.
Troglopedetes spectabilis sp. nov. exclusively inhabits a chamber in the dark zone of a cave. Specimens were found in an oligotrophic habitat with a very humid and wet environment on muddy ground. The air temperature in the chamber was 23.5–24.3 °C, soil temperature was 22.9–23.1 °C and the relative humidity was 84%. The cave is undisturbed and rather difficult to access (due to the steep slope up to the entrance). It is located in dry evergreen forest mixed with bamboo forest at moderate altitude (426 m. asl.). Some simple infastructure has been introduced into the cave, including a metal ladder at the entrance.
The species name is derived from the Latin word spectabilis, meaning “remarkable” or “admirable” and referring to its extremely long antennae.
Troglopedetes spectabilis sp. nov. is the most highly troglomorphic species of Troglopedetes. It is clearly distinct from other species of the genus by the extreme length of its appendages, especially the antennae (the ratio of antenna: head is 5.5), by its claw complex and by its furca (Table
Thailand, Tak Province, Tha Song Yang district, Tham (cave) Mae Usu (17°18'15.0"N, 98°09'19.9"E, 172 m a.s.l.).
Holotype : female on slide, Tak Province: Tha Song Yang district, Tham (cave) Mae Usu, 17°18'15.0"N, 98°09'19.9"E, 145 m a.s.l., 7 March 2019, S. Jantarit, A. Nilsai and K. Jantapaso leg., dark zone of cave, by aspirator (sample # THA_SJ_TAK03). Paratypes: 5 paratypes on slides (all subadult), same locality and date as the holotype.
Holotype and 5 paratypes deposited in NHM-PSU, measurements of holotype in Table
Head | Tergites | Appendages | |||
---|---|---|---|---|---|
Ant. I | 192 | Th. II | 200 | Man | 540 |
Ant. II | 390 | Th. III | 212 | Dens | 580 |
Ant. III | 390 | Abd. I | 110 | Mucro | 48 |
Ant. IVa | 322 | Abd. II | 120 | Furca | 1,168 |
Ant. IVb | 230 | Abd. III | 140 | Claw I | 62 |
Ant. | 1,524 | Abd. IV | 655 | Claw II | 62 |
Head | 465 | Abd. V | 80 | Claw III | 65 |
Abd. VI | 60 | ||||
Body | 1,577 |
Habitus. Troglomorphic, slender, with elongate legs, furca and antennae. Length: 1.9–2.0 mm (n = 4) (body 1.4–1.6 mm, head 0.4 mm). Fourth abdominal segment 4–5 times (n = 4) longer than the third one along the dorsal axis. Furca well developed, about 1.4–1.5 (n = 3) times shorter than the body length. Body colour white with spots of orange pigment. Eyes not detected, but with 2+2 pigmented ocular patches.
Chaetal types. Four types of chaetae on somites, appendages (except antennae) and mouthparts: scales, present on antennal segment I and II, head, body and ventral side of the furca, absent on legs and ventral tube; ordinary chaetae on all body parts; S-chaetae and trichobothria on tergites; hairs devoid of sockets on outer maxillary lobe. Chaetal types on antennae are much more diverse and described separately further.
Pseudopores
(Figs
Mouthparts. Clypeus with 13 large mes, symmetrically arranged: 9 ciliated posteriorly and 4 thin smooth anteriorly (Fig.
Antennae
(Figs
General chaetotaxy
(Figs
Dorsal macrochaetae formula: 0,2/8,4/0,2,4,3 (Figs
Head chaetotaxy
(Fig.
Head dorsally densely covered with round to oval scales (33–45 µm long). Ventral chaetotaxy of head densely covered with oval scales (57–67 µm long), postlabial chaetae along the linea ventralis as ciliated mes anteriorly and 3 ciliated chaetae laterally, one mac and an oblique line of 5–6 mes posteriorly on each side (Fig.
Tergite chaetotaxy
(Fig.
Th. III with 4 mac by side (a group of 3 central and 1 anterior to them), 1 sens at antero-lateral margins, and about 12 mac or long mes at lateral margins (Fig.
Abd. I without central mac, with 1 ms laterally on each side, a row of 3 mics below psp, a6 absent, and 5 mes laterally (Fig.
Abd. II with 2 tric on each side and 6 modified mes around them (2 around the internal tric and 4 near external tric), 2 mac (1 near internal tric and 1 near external tric), 1 sens near internal tric, 4 mic (1 close to internal tric and 3 close to external tric), at least 3 other mes at lateral margins (Fig.
Abd. III with 3 tric on each side (1 internal, 2 external) and 8 modified mes around tric (2 near internal tric, 6 near the two external tric); 4 mac (1 near internal tric and 3 near external tric); 1 sens anterior to internal tric and 1 ms; at least 5 mes to small mac at lateral margins (Fig.
Abd. IV with 3 tric on each side (2 antero-lateral, 1 postero-lateral) and about 7 modified mes around the two antero-lateral tric; postero-lateral tric with 1 modified mes. Mac distributed as 3 central on each side (1 antero-external to pseudopore, 2 anterior to posterior tergite margin), 1 near postero-lateral tric, and at least 5 external, mixed with at least 14 mes or smaller mac on lateral to posterior margins; 3 sens; at least 4 S-like chaetae sensu
Abd. V with 2 sens detected on each side, and several ordinary chaetae from mes to mac, not counted (Fig.
Legs
(Fig.
Ventral tube
(Fig.
Furca complex
(Fig.
T. takensis sp. nov. is only known from a large chamber in the dark zone of a cave. Specimens were found as small populations in an oligotrophic habitat, i.e. on the wall and ground surface, with a very humid and wet environment, with small trace of organic matter. Air temperature in the chamber was 22.2–23.0 °C, soil temperature was 21.9 °C and the relative humidity was 85%.
The species name is derived from the name of the province (Tak) where it was found.
Troglopedetes takensis sp. nov. has elongated antennae (the ratio of antenna: head is 3.2), legs and furca (Table
Thailand, Kanchanaburi Province, Sai Yok district, Tham (cave) Khang Khao (14°11'23.8"N, 98°59'37.0"E, 262 m a.s.l).
Holotype : male on slide, Kanchanaburi Province: Sai Yok district, Tham (cave) Khang Khao, 14°11'23.8"N, 98°59'37.0"E, 262 m a.s.l., 28 February 2019, S. Jantarit, A. Nilsai and K. Jantapaso leg., dark zone of cave, by aspirator (sample # THA_SJ_KRI11). Paratypes: 2 subadults on slides, same locality and date as the holotype.
Holotype and 2 paratypes deposited in NHM-PSU, measurements of holotype in Table
Head | Tergites | Appendages | |||
---|---|---|---|---|---|
Ant. I | 95 | Th. II | 128 | Man | 340 |
Ant. II | 240 | Th. III | 120 | Dens | 338 |
Ant. III | 210 | Abd. I | 70 | Mucro | 37 |
Ant. IVa | 200 | Abd. II | 80 | Furca | 715 |
Ant. IVb | 160 | Abd. III | 90 | Claw I | 30 |
Ant. | 905 | Abd. IV | 430 | Claw II | 32 |
Head | 260 | Abd. V | 70 | Claw III | 32 |
Abd. VI | 45 | ||||
Body | 1,033 |
Habitus. Troglomorphic, slender, with elongate legs, furca and antennae. Length: 1.2–1.3 mm (n = 3) (body 0.9–1.0 mm, head 0.2–0.3 mm). Fourth abdominal segment 4–5 times (n = 3, 1 male and 2 subadults) longer than the third one along the dorsal axis. Furca well developed, about 1.4–1.5 (n = 3) times shorter than body length. Body colour white with spots of orange pigment. Eyes absent, no ocular patch.
Chaetal types. Four types of chaetae on somites, appendages (except antennae) and mouthparts: scales, present on antennal segment I and II, head, body and ventral side of the furca, absent on legs and ventral tube; ordinary chaetae on all body parts; S-chaetae and trichobothria on tergites; hairs devoid of sockets on outer maxillary lobe. Chaetal types on antennae are much more diverse and described separately further.
Pseudopores
(Figs
Troglopedetes rungsimae sp. nov. A outer maxillary lobe B papilla E of labial palp C head chaetotaxy (left = A to G mac nomenclature; right = AMS nomenclature) D labial basis and ventral chaetotaxy of head, right side E distal part of tibiotarsus III and claw complex F lateral flap of ventral tube G mucro H male genital plate I distal part of tibiotarsus III and claw complex with clavate tenent hair of an undescribed species co-occurring with T. rungsimae sp. nov.
Mouthparts. Clypeus not visible. Labral formula 4/5,5,4; prelabral chaetae short, bent and ciliated, labral chaetae thinner, longer, smooth and acuminate, those of the distal row slightly larger and longer than those of the median row; the anterior line not cleary seen. Ventro-distal complex of labrum well differentiated, asymmetrical, with 2 distal combs (a larger one with 9–10 teeth on the left side, a smaller one with 10–12 minute teeth on the right side) and an axial pair of sinuous tubules. Distal part of labrum not adorned with spines dorso-distally. Labial palp similar to that described by
Antennae. Antennae (743–905 µm, n = 3), shorter than body+head length (n = 3), 3.1 times (n = 3) longer than the cephalic diagonal. Ant. IV subdivided into two segments, asymmetrically arranged with Ant. IVa longer than IVb (0.57 : 0.43, n = 3), without apical bulb. Length of antennal segments I to IV (IVa+IVb) as 1:2.4:2.2:3.6 (n = 2). Antennal chaetae (scales, 5 types of ordinary chaetae, 13 types of S-chaetae and subapical organit).
General chaetotaxy
(Figs
Head chaetotaxy
(Fig.
Troglomorphic traits (appendage elongation and body size) in Thai species of the genus Troglopedetes A highly troglomorphic species T. spectabilis sp. nov., where Ant.II and III are fused together B troglomorphic form of an undescribed species C non-troglomorphic form of an undescribed species with eyes and eyes patch.
Tergite chaetotaxy
(Fig.
Th. III with 4 mac by side (a group of 3 central and 1 anterior to them), 1 sens at antero-lateral margins, and about 9 mac or long mes at lateral margins (Fig.
Abd. I without central mac, with 1 ms laterally on each side, a row of 3 mics below psp, a6 absent, and 5 mes laterally (Fig.
Abd. II with 2 tric on each side and 7 modified mes around them (2 around the internal tric and 5 near external tric), 2 mac (1 near internal tric and 1 near external tric), 1 sens near internal tric (Fig.
Abd. III with 3 tric on each side (1 internal, 2 external) and 7 modified mes around tric (2 near internal tric, 5 near the two external tric); 4 mac (1 near internal tric and 3 near external tric); 1 sens anterior to internal tric and 1 ms; at least 6 mic to mes at lateral margins (Fig.
Abd. IV with 3 tric on each side (2 antero-lateral, 1 postero-lateral) and about 7 modified mes around the two antero-lateral tric; postero-lateral tric with 1 modified mes. Mac distributed as 3 central on each side (1 antero-external to pseudopore, 2 anterior to posterior tergite margin), 1 near postero-lateral tric, and at least 5 external, mixed with at least 13 mes or smaller mac on lateral to posterior margins; 3 sens; at least 5 S-like chaetae sensu
Abd. V with 2 sens detected on each side, and several ordinary chaetae from mes to mac, not counted (Fig.
Legs
(Fig.
Ventral tube
(Fig.
Furca complex. Tenaculum with 4 teeth on each ramus, of decreasing size from the basal to the distal one, on a prominent, irregular body, with a postero-basal strong, densely serrated, distally bent chaeta. Manubrium about 1.13 times (n = 3) shorter than mucrodens (mucro+dens). Manubrium dorsally with subequal ciliated mes (none smooth), irregularly arranged in 3–4 rows in two longitudinal stripes separated by a glabrous axial stripe, external row of chaetae distally with at least 10 long ciliated mac, dorso-distal plaque with 4+4 mes and 2+2 pseudopores. Ventrally, with a dense cover of round to oval (15–25 µm long) and thin elongated scales (25–30 µm long). Dens straight, elongate, hairy, slightly and progressively tapering, dorsally with 2 rows of spines, mixed with ciliated mes of various length, thickness and shape. Dorso-external row with 16–18 spines, dorso-internal row with 29 spines (asymmetries between dentes); external spines larger and less sclerotized than the internal ones. Some short ciliated mes interspersed with spines in the external row; dorsally between the two rows of spines a mix of short and long ciliated mes, irregularly arranged in one row distally turning to 3–4 rows proximally; laterally, many short ciliated mes; dorso-distally, 3–(4) stronger ciliated mes; 2+2 psp on dorso-basally between the two rows of spine. Dens ventrally entirely and densely scaled, the scales elongate (25–40 µm long) (oval shape distally), arranged in short lines from 3–5 (distally) to 6–8 scales (proximally). Mucro rather stout, short, 8.9–9.2 (n = 3) times shorter than the dens, with 4 main teeth, the apical one blunt and strong, the subapical one acute and strong, a latero-distal one small and acute, and 1 dorso-basal, minute, acute and strong, with one toothlet basally (Fig.
Genital plate
(Fig.
Troglopedetes rungsimae sp. nov. is only known from a small chamber in the dark zone of a cave. Specimens were found as small populations in an oligotrophic habitat, i.e. on the wall and ground with a very humid and wet environment, without any trace of organic matter. Air temperature in the chamber where specimens were collected was 23.5–24.8 °C, soil temperature was 23.1–23.3 °C and relative humidity was 88–91%.
The species is named in honour of Rungsima Tanthalakha, the Senior Program Director, Research Management and Innovation Management, National Science and Technology Develoment Agency, Thailand, who is interested in karst and cave biodiversity and for her contributions to the study of cave Troglopedetes in Thailand.
Troglopedetes rungsimae sp. nov. has the sixth most elongated antennae of the Troglopedetes species of Thailand (Table
In the same cave, we found another morphotype with a different claw morphology. This type has thin, smooth and clavate tenent hair on all tita (one pointed on Claw I). Claw is long and slender with 2 strong inner teeth, one tooth at 57–73% of inner edge and the other at a 90–91% of inner edge, and a pair of inner basal teeth of unequal size. One small tooth is at 40% on the outer edge. Unguiculus is pointed, narrow, lanceolate and elongate, about 0.65 time shorter than the claw, and its external edge is with 7 toothlets (Fig.
The genus Troglopedetes in Thailand is dominant in the subterranean environment in the northern and western regions and is present in the mountain range bordering Myanmar (unpublished data). It is also spread over a wide area of the central plain region where karst is extremely fragmented (Fig.
Cave Troglopedetes in Thailand exhibit various degrees of adaption to cave life (troglomorphy), from a small set of morphological modifications to a complete set of darkness adaptions, as is also presented in the genus Coecobrya Yosii, 1956 (
Phanere types and their arrangement pattern on each antennal segment that we recognised in the three new species are homologised and compared with the two species currently described from Thailand (T. meridionalis and T. kae, see details in
For the scales (see
Detailed distribution of antennal chaetae in the three new species and in T. meridionalis and T. kae.
Type of chaetae | Distribution on antenna | Location | Position on antennal segment | Number of chaetae | ||||
---|---|---|---|---|---|---|---|---|
T. meridionalis | T. kae | T. spectabilis sp. nov. | T. takensis sp. nov. | T. rungsimae sp. nov. | ||||
Type-1 | Ant. III | dorsal | latero-distal (AIIIO) | 1 | 1 | - | - | 1 |
Ant. III | ventral | latero-diatal (AIIIO) | - | - | 1 | 1 | - | |
Ant.IVa | ventral | proximal | - | - | - | - | 1 | |
Type-2 | Ant. I | ventral | basal | 4 | 4 | - | - | - |
Ant. II | dorsal | basal | 2 | 2 | 2 | 1 | 2 | |
Ant. II | ventral | basal | - | - | 1 | 1 | 1 | |
Type-3 | Ant. I | ventral | proximal | 2 | 2 | 2 | 2 | 2 |
Ant. II | ventral | proximal | 1 | 1 | - | 1 | 1 | |
Ant. II | dorsal | proximal | - | - | 1 | - | - | |
Type-4 | Ant.II | dorsal | AIIIO | - | - | - | - | 2 |
Ant.II | ventral | AIIIO | - | - | - | 5 | - | |
Ant. III | dorsal | AIIIO | 2 | 2 | - | - | 2 | |
Ant. III | ventral | AIIIO | - | - | 2 | 2 | - | |
Ant.IVb | dorsal | lateral | - | - | - | - | 1 | |
Ant.IVb | ventral | lateral | - | - | - | - | 1 | |
Type-5 | Ant. I | ventral | all segment | 7 | 6 | - | 8 | - |
Ant. I | ventral | middle to proximal | - | - | 2 | - | 3 | |
Ant. III | dorsal | AIIIO | 2 | 2 | - | - | 2 | |
Ant. III | ventral | AIIIO | - | - | - | 2 | - | |
Type-6 | Ant. I | dorsal | basal | 3 | 3 | 3 | 2 | 3 |
Ant. I | ventral | basal | - | - | 3 | 3 | 3 | |
Type-7 | Ant. II | ventral | all segment | 3 | 7 | - | - | 6 |
Ant. III | dorsal | lateral | - | 2 | - | - | - | |
Ant. III | ventral | middle to proximal | 3 | 4 | - | - | 5 | |
Ant. IVa | dorsal | middle to proximal | 4 | 4 | - | - | - | |
Ant. I | ventral | all segment | 6 | 10 | 18 | 8 | 13 | |
Type-8 | Ant. II | dorsal | middle to proximal | 9 | 8 | 5 | 8 | 13 |
Ant. II | ventral | all segment | 9 | 9 | 7 | - | 9 | |
Ant. II | ventral | proximal | - | - | - | 8 | - | |
Ant. III | dorsal | all segment | 10 | 15 | - | 14 | 20 | |
Ant. III | ventral | middle to proximal | 11 | 6 | 6 | - | 11 | |
Ant. III | ventral | all segment | - | - | - | 14 | - | |
Ant. IVa | dorsal | all segment | 9 | 10 | - | 7 | 18 | |
Ant. IVa | ventral | all segment | 13 | 5 | 2 | 12 | 20 | |
Ant. IVb | dorsal | all segment | 9 | 9 | 23 | - | 18 | |
Ant. IVb | dorsal | basal | - | - | - | 1 | - | |
Ant. IVb | ventral | all segment | 11 | - | 4 | 8 | 15 | |
Ant. I | ventral | latero-proximal | 2 | 1 | 5 | 1 | 1 | |
Type-9 | Ant. II | ventral | proximal | 2 | 4 | - | - | - |
Ant.IVa | ventral | latero-proximal | - | - | - | - | - | |
Ant. I | ventral | latero-proximal | 14 | 5 | 15 | 12 | 4 | |
Type-10 | Ant. II | ventral | proximal | 2 | 5 | - | - | - |
Ant. II | ventral | lateral | - | - | 24 | - | - | |
Ant. II | ventral | latero-proximal | - | - | - | 24 | 6 | |
Ant. III | dorsal | upper middle | - | 1 | 14 | - | - | |
Ant. III | dorsal | all segment | - | - | - | 9 | - | |
Ant. III | ventral | proximal | 1 | 4 | - | - | 5 | |
Ant. III | ventral | all segment | - | - | 34 | 23 | - | |
Ant. IVa | dorsal | middle to proximal | 1 | 3 | - | - | - | |
Ant. IVa | dorsal | all segment | - | - | 34 | - | - | |
Ant. IVa | ventral | all segment | - | - | 60 | 8 | 12 | |
Ant. IVb | dorsal | middle | 2 | 3 | - | - | 3 | |
Ant. IVb | dorsal | basal | - | - | 3 | - | 7 | |
Ant. IVb | ventral | basal to middle | - | - | 12 | 3 | - | |
Ant. IVa | dorsal | latero-proximal | - | 1 | - | - | - | |
Type-11 | Ant. IVa | dorsal | all segment | - | - | 10 | 10 | 22 |
Ant. IVa | ventral | lateral | - | - | 4 | - | 19 | |
Ant. IVb | dorsal | all segment | 19 | 22 | 42 | 44 | 37 | |
Ant. IVb | ventral | all segment | 35 | 36 | 40 | 18 | 45 | |
Ant. I | ventral | all segment | 8 | 6 | 17 | 8 | 9 | |
Type-12 | Ant. I | ventral | lateral | - | - | - | 3 | - |
Type-13 | Ant. II | ventral | lateral | - | - | - | 5 | 3 |
Ant. III | ventral | lateral | - | - | - | 3 | - | |
Ant. III | ventral | all segment | - | - | 9 | - | - | |
Ant. IVa | dorsal | middle of Ant. IVa | - | 3 | - | - | - | |
Ant. IVa | ventral | lateral | - | - | 1 | - | - | |
Ant. IVa | ventral | middle | - | - | - | 1 | 1 | |
Ant. II | dorsal | proximal | 1 | 1 | 1 | - | - | |
Type-14 | Ant. III | dorsal | all segment | - | - | 17 | - | - |
Type-15 | Ant. III | ventral | middle to proximal | - | - | 3 | - | - |
Ant. III | ventral | all segment | - | - | - | 13 | - | |
Ant. IVa | dorsal | all segment | - | - | 10 | - | - | |
Ant. IVa | ventral | all segment | - | - | 26 | 9 | - | |
Ant. IVb | ventral | all segment | - | - | 22 | 6 | - | |
Total of S-chaetae | 208 | 207 | 485 | 308 | 347 | |||
Ordinary chaetae | Ant. I | dorsal | all segment | 13 | 28 | 58 | 26 | 16 |
Ant. I | ventral | all segment | 27 | 38 | 113 | 58 | ||
Ant. II | dorsal | all segment | 63 | 67 | 157 | 93 | 72 | |
Ant. II | ventral | all segment | 68 | 77 | 161 | 69 | 63 | |
Ant. III | dorsal | all segment | 45 | 38 | 167 | 97 | 64 | |
Ant. III | ventral | all segment | 36 | 45 | 169 | 93 | 62 | |
Ant. IVa | dorsal | all segment | 62 | 59 | 148 | 117 | 64 | |
Ant. IVa | ventral | all segment | 53 | 60 | 142 | 98 | 58 | |
Ant. IVb | dorsal | all segment | 49 | 48 | 150 | 65 | 138 | |
Ant. IVb | ventral | all segment | 67 | 58 | 137 | 75 | 112 | |
Total of ordinary chaetae | 483 | 518 | 1,402 | 790 | 687 | |||
Total S-chaetae and ordinary chaetae | 692 | 726 | 1,887 | 1,098 | 1,034 | |||
Subapical organ | Ant. IVb | dorsal | proximal near the tip | 1 | 1 | 1 | 1 | 1 |
Scales | Ant. I | dorsal | all segment | 19 | 25 | 127 | 58 | 29 |
Ant. II | dorsal | basal to middle | 25 | 13 | 152 | 13 | 30 | |
Ant. II | ventral | basal (to middle) | 4 | 3 | 16 | 57 | 13 | |
Overall antennal chaetae | 741 | 768 | 2,183 | 1,228 | 1,107 |
Regarding ordinary chaetae, the five types mentioned above are present in the new species and their position is the same as described by
The subapical organite of Ant. IV is short, thick, dark, swollen at the tip (4 µm) with protecting chaeta, inserted dorso-internally ca. 38–47 µm from the apex (Figs
With reference to the S-chaetae (sensu
The number of antennal phanneres used for the analysis and comparison in this study was observed from the holotype of each species. We counted a total of 741 to 2,183 chaetae per antenna for the 5 studied species of Thai Troglopedetes, including 207–485 S-chaetae, 483–1,402 ordinary chaetae, 48–295 scales and a single subapical organite (Tables
Number of chaetae of each type along antennal segments in the three new species and in T. meridionalis and T. kae.
Type of S-chaetae/antennal segment | Troglopedetes meridionalis | Troglopedetes kae | Troglopedetes spectabilis sp. nov. | Troglopedetes takensis sp. nov. | Troglopedetes rungsimae sp. nov. | |||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Ant. I | Ant. II | Ant. III | Ant. IV | Total | Ant. I | Ant. II | Ant. III | Ant. IV | Total | Ant. I | Ant. II | Ant. III | Ant. IV | Total | Ant. I | Ant. II | Ant. III | Ant. IV | Total | Ant. I | Ant. II | Ant. III | Ant. IV | Total | ||||||
Ant. IVa | Ant. IVb | Ant. IVa | Ant. IVb | Ant. IVa | Ant. IVb | Ant. IVa | Ant. IVb | Ant. IVa | Ant. IVb | |||||||||||||||||||||
Length (µm) | 78 | 150 | 125 | 125 | 125 | 603 | 90 | 215 | 148 | 136 | 138 | 727 | 374 | 667 | 720 | 595 | 424 | 2,780 | 192 | 390 | 390 | 322 | 230 | 1,524 | 95 | 240 | 210 | 200 | 160 | 905 |
Type-1 | - | - | 1 | - | - | 1 | - | - | 1 | - | - | 1 | - | - | 1 | - | - | 1 | - | - | 1 | - | - | 1 | - | - | 1 | 1 | - | 2 |
Type-2 | 4 | 2 | - | - | - | 6 | 4 | 2 | - | - | - | 6 | - | 3 | - | - | - | 3 | - | 2 | - | - | - | 2 | - | 3 | - | - | - | 2 |
Type-3 | 2 | 1 | - | - | - | 3 | 2 | 1 | - | - | - | 3 | 2 | 1 | - | - | - | 3 | 2 | 1 | - | - | - | 3 | 2 | 1 | - | - | - | 3 |
Type-4 | - | - | 2 | - | - | 2 | - | - | 2 | - | - | 2 | - | - | 2 | - | - | 2 | - | 5 | 2 | - | - | 7 | - | 2 | 2 | - | 2 | 6 |
Type-5 | 7 | - | 2 | - | - | 9 | 6 | - | 2 | - | - | 8 | 2 | - | - | - | - | 2 | 8 | - | 2 | - | - | 10 | 3 | - | 2 | - | - | 5 |
Type-6 | 3 | - | - | - | - | 3 | 3 | - | - | - | - | 3 | 6 | - | - | - | - | 6 | 5 | - | - | - | - | 5 | 6 | - | - | - | - | 6 |
Type-7 | - | 3 | 3 | 4 | - | 10 | - | 7 | 6 | 4 | - | 17 | - | - | - | - | - | 0 | - | - | - | - | - | - | - | 6 | 5 | - | - | 12 |
Type-8 | 6 | 18 | 21 | 22 | 20 | 87 | 10 | 17 | 21 | 15 | 9 | 72 | 18 | 12 | 6 | 2 | 27 | 65 | 8 | 16 | 28 | 19 | 9 | 80 | 13 | 22 | 31 | 38 | 33 | 137 |
Type-9 | 2 | 2 | - | - | - | 4 | 1 | 4 | - | - | - | 5 | 5 | - | - | - | - | 5 | 1 | - | - | - | - | 1 | 1 | - | - | - | - | 1 |
Type-10 | 14 | 2 | 1 | 1 | 2 | 20 | 5 | 5 | 5 | 3 | 3 | 21 | 15 | 24 | 48 | 94 | 15 | 196 | 12 | 24 | 32 | 8 | 3 | 79 | 4 | 6 | 5 | 12 | 10 | 36 |
Type-11 | - | - | - | - | 54 | 54 | - | - | - | 1 | 58 | 59 | - | - | - | 14 | 82 | 96 | - | - | - | 10 | 62 | 72 | - | - | - | 41 | 82 | 123 |
Type-12 | 8 | - | - | - | - | 8 | 6 | - | - | - | - | 6 | 17 | - | - | - | - | 17 | 8 | - | - | - | - | 8 | 9 | - | - | - | - | 9 |
Type-13 | - | - | - | - | - | 0 | - | - | - | 3 | - | 3 | - | - | 9 | 1 | - | 10 | 3 | 5 | 3 | 1 | - | 12 | - | 3 | - | 1 | - | 4 |
Type-14 | - | 1 | - | - | - | 1 | - | 1 | - | - | - | 1 | - | 1 | - | - | - | 1 | - | - | - | - | - | 0 | - | - | - | - | - | 0 |
Type-15 | - | - | - | - | - | 0 | - | - | - | - | - | 0 | - | - | 20 | 36 | 22 | 78 | - | - | 13 | 9 | 6 | 28 | - | - | - | - | - | - |
Overall S-chaetae | 46 | 29 | 30 | 27 | 76 | 208 | 37 | 37 | 37 | 26 | 70 | 207 | 65 | 41 | 86 | 147 | 146 | 485 | 48 | 53 | 81 | 47 | 80 | 308 | 38 | 43 | 46 | 93 | 127 | 347 |
Overall ordinary chaetae | 40 | 131 | 81 | 115 | 116 | 483 | 66 | 144 | 83 | 119 | 106 | 518 | 171 | 318 | 336 | 290 | 287 | 1402 | 84 | 162 | 190 | 214 | 140 | 790 | 54 | 135 | 126 | 122 | 250 | 687 |
Total S- and ordinary chaetae | 86 | 160 | 111 | 142 | 193 | 692 | 103 | 181 | 119 | 145 | 177 | 726 | 236 | 359 | 422 | 437 | 433 | 1887 | 132 | 215 | 271 | 261 | 220 | 1099 | 92 | 178 | 172 | 215 | 377 | 1034 |
Subapical organite | - | - | - | - | 1 | 1 | - | - | - | - | 1 | 1 | - | - | - | - | 1 | 1 | - | - | - | - | 1 | 1 | - | - | - | - | 1 | 1 |
Scale | 19 | 29 | - | - | - | 48 | 25 | 16 | - | - | - | 41 | 127 | 168 | - | - | - | 295 | 58 | 70 | - | - | - | 128 | 29 | 43 | - | - | - | 72 |
Overall chaetae | 105 | 189 | 111 | 142 | 194 | 741 | 128 | 197 | 119 | 145 | 178 | 768 | 363 | 527 | 422 | 437 | 434 | 2183 | 189 | 285 | 271 | 262 | 221 | 1228 | 121 | 221 | 172 | 215 | 378 | 1107 |
The ventral side of the antenna is richer in chaetae and S-chaetae than the dorsal side (overall = 1,042 versus 846 in T. spectabilis sp. nov.; 605 versus 494 in T. takensis sp. nov. and 528 versus 507 in T. rungsimae sp. nov.; S-chaetae = 320 versus 166 in T. spectabilis sp. nov.; 212 versus 96 in T. takensis sp. nov. and 195 versus 153 in T. rungsimae sp. nov. (Tables
The distribution of the different types of S-chaetae along with the antennal segments is arranged in more or less clearly defined patterns which are described below, compared with T. meridionalis and T. kae in the following paragraphs, summarised in Tables
First antennal segment: eight types of S-chaetae can be recognised (Figs
Second antennal segment: nine types of S-chaetae can be recognised (Figs
Third antennal segment: eight types of S-chaetae can be recognised (Figs
Fourth antennal segment: nine types of S-chaetae can be recognised (Figs
Fourth antennal segment I (a): seven types of S-chaetae can be recognised: type 1, type 8, type 9, type 10, type 11, type 13, type 15. They are present on the dorsal and ventral side, except two types (1 and 9) that are present on the ventral side only.
Fourth antennal segment II (b): six types of S-chaetae can be recognised: type 4, type 7, type 8, type 10, type 11 and type 15. They are present on the dorsal and ventral side, except one type (7) that is present on the dorsal side only and one type (15) that is present on the ventral side only.
The most frequent antennal S-chaetae are type 8 (from 65 to 137 chaetae) and type 10 (from 36 to196 chaetae), which are present along all antennal segments. The other types are specific to three, two or a single segment(s) (Table
With regard to the abundance of S-chaetae along antennal segments, the result for the new species is similar to that in T. meridionalis and T. kae where type 8 is the most common followed by type 11 and 10 (Tables
The distal subsegment of antenna IV (Ant. IVb) is richer in S-chaetae than the proximal subsegment (Ant. IVa) in T. takensis sp. nov. and T. rungsimae sp. nov., but the number of S-chaetae is similar in T. spectabilis sp. nov.. The ordinary chaetae are more numerous in the proximal subsegment (Ant. IVa) than in the distal subsegment (Ant. IVb) in T. takensis sp. nov. and T. rungsimae sp. nov., but the number is rather similar in T. spectabilis sp. nov. (Table
Antennal chaetotaxy in Thai Troglopedetes is clearly plurichaetotic and extremely complex. Overall 22 types of antennal chaetae have been recognized. They are ordinary chaetae (5 types), S-chaetae (15 types), the subapical organite of Ant. IV and scales. According to
It seems that antennal phanerotaxy in the genus Troglopedetes is richer than in other Entomobryidae described so far. For example, 13 types of chaetae are reported in Alloscopus (
Generally, the length of antennal segments in Thai Troglopedetes ranked as Ant. IV (a+b) > II > III > I (
According to the study of
On Ant. I, the shortest segment, the number of S-chaetae varies depending on species. In T. meridionalis and T. kae, Ant. I has more S-chaetae than Ant. II and III, while in T. spectabilis sp. nov., Ant. I has more S-chaetae than Ant. II only (Table
Ant. II is the second longest segment for the moderate long antennal species (T. rungsimae sp. nov. and also in T. meridionalis and T. kae), but it can be shorter than (T. spectabilis sp. nov.) or subequal to (T. takensis sp. nov.) Ant. III for the species with very long antennae. This segment has less S-chaetae than Ant. III (Table
Ant. III is the second longest segment in species with very long antennae (T. spectabilis sp. nov., and T. takensis sp. nov.) and carries the highest number of S-chaetae after Ant. IV (Table
Meanwhile, the ventral side of the antennal segment possesses a higher number of chaetae than the dorsal one both in overall chaetae and S-chaetae (Table
The diversity of antennal chaetae in Thai Troglopedetes does not differ significantly between the extremely long antennae and the moderately long antennae species. T. spectabilis sp. nov., the longest antennae species so far, has 21 types of chaetae (5 types of ordinary chaetae, 14 type of S-chaetae, a subapical organite and scale), whereas T. takensis sp. nov., T. rungsimae sp. nov. (long antennae) and T. kae (moderate long antennae) all have 20 types of chaetae (5 types of ordinary chaetae, 13 S-chaetal types, subapical organite and scale). Troglopedetes meridionalis (moderate long antennae) has 19 types of chaetae (5 types of ordinary chaetae, 12 S-chaetal types, subapical organite and scale) (Tables
Groups of subcylindrical S-chaetae (type 10) on antennal segments I, II and III deserve to be mentioned here. There are clusters of numerous S-chaetae in three Mediterranean cave species (T. ruffoi, T. absoloni and T. ildumensis) (
Number of subcylindrical type 10 S-chaetae on antennal segments I, II and III ventrally in the three new species from Thailand and three Mediterranean cave species.
Species/Antennal segment | Ant. I | Ant. II | Ant. III | Country | Source |
---|---|---|---|---|---|
T. absoloni | 3–5 | 15 | 15 | Spain |
|
T. ildumensis | 2–5 | 25–30 | 25–30 | Spain |
|
T. kae | 5 | 5 | 5 | Thailand |
|
T. meridionalis | 14 | 2 | 1 | Thailand |
|
T. ruffoi | 9 | 31 | 34 | Italy |
|
T. rungsimae sp. nov. | 4 | 6 | 5 | Thailand | This study |
T. spectabilis sp. nov. | 15 | 24 | 48 | Thailand | This study |
T. takensis sp. nov. | 12 | 24 | 32 | Thailand | This study |
Although the function of individual chaetal types of Collembola antennae is little known, differences in the morphological structure of chaetae are likely to enable springtails to receive a wide range of different stimuli, responding to light, smell, taste, sound, touch, vibration, stretch, temperature, humidity, a multitude of chemical agents and concentration gradients of oxygen and carbon dioxide (
1 | Central area of head macrochaetae absent | 2 |
– | Central area of head macrochaetae present | 3 |
2 | Eyes absent but each ocular area with 2 black spots; claw without inner teeth; tenent hair pointed; antenna as long as the body | T. takensis sp. nov. |
– | Eyes and ocular patches absent; claw with 2 inner teeth; tenent hair clavate; antenna 0.6 times shorter than the body | T. calvus Deharveng & Gers, 1993 |
3 | Central area of head with 1–2+1–2 macrochaeta(e) (A and E mac) | 4 |
– | Central area of head with 3+3 macrochaetae (A, B or C, and E mac) | 6 |
– | Central area of head with 4–6+4–6 macrochaetae (A, B, E (C, D, F, G) mac) | 8 |
– | Central area of head with 7+7 macrochaetae (A–G mac) | 13 |
4 | Central area of head with 1+1 macrochaetae (A mac); lateral flap of ventral tube with 6 chaetae; tenent hair pointed; internal row of dens with 30–37 spines | T. meridionalis Jantarit, Surakhamhaeng & Deharveng, 2020 |
– | Central area of head with 1–2+1–2 macrochaetae (A and E mac); lateral flap of ventral tube with 7 chaetae; tenent hair clavate; internal row of dens with 18–24 spines | 5 |
5 | Abd. IV with 2+2 central mac; claw with 2 inner teeth | T. paucisetosus Deharveng & Gers, 1993 |
– | Abd. IV with 3+3 central mac; claw with 1 inner tooth | T. convergens Deharveng & Gers, 1993 |
6 | Head macrochaetae with A, B, E mac; body length 1.8–2.2 mm; internal row of dens with 34–41 spines | T. multispinosus Deharveng & Gers, 1993 |
– | Head macrochaetae with A, C, E mac; body length 0.9–1.4 mm; internal row of dens with 25–29 spines | 7 |
7 | Antenna as long as the body; body length 0.9–1.0 mm; lateral flap of ventral tube with 8 chaetae | T. rungsimae sp. nov. |
– | Antenna 0.6 times shorter than the body; body length 1.3–1.4 mm; lateral flap of ventral tube with 7 chaetae | T. dispersus Deharveng & Gers, 1993 |
8 | 1–2+1–2 eyes; outer maxillary lobe with 1 sublobal hair | T. microps Deharveng & Gers, 1993 |
– | Eyes absent; outer maxillary lobe with 2 sublobal hair; Abd. IV with 3+3 central mac | 9 |
9 | Th. II with 9+9 central mac; labial formula: M1m2rel1l2; antenna very long about 1.8 times longer than the body | T. spectabilis sp. nov. |
– | Th. II with 8+8 central mac; inter teeth of claw with 1+1 | 10 |
10 | Head macrochaeta D present; labial formula: M1M2ReL1l2; antenna 0.8 times shorter than the body | T. longicornis Deharveng & Gers, 1993 |
– | Head macrochaeta D absent; labial formula: M1M2REL1l2; antenna 0.4–0.5 times shorter than the body | 11 |
11 | Central area of head with 5+5 macrochaetae; Th. II with 8+8 central mac, Abd.IV with 3+3 central mac; dens 15 times longer than the mucro | T. fredstonei Deharveng, 1988 |
– | Central area of head with 4+4 macrochaetae; Th. II with 8+8 central mac, Abd.IV with 3+3 central mac; dens 8.8–14 times longer than the mucro | 12 |
12 | Lateral flap with 6+6 chaetae; internal row of dens with 26–33 spines; mucro with 5 teeth | T. kae Jantarit, Surakhamhaeng & Deharveng, 2020 |
– | Lateral flap with 7+7 chaetae; internal row of dens with 37–42 spines; mucro with 4 teeth | T. centralis Deharveng & Gers, 1993 |
13 | Eyes absent, outer maxillary lobe with 1 sublobal hair; Th. II with 9+9 central mac; Abd.IV with 3+3 central mac | 14 |
– | 3+3 eyes, outer maxillary lobe with 2 sublobal hairs; Th. II with 8+8 central mac; Abd. IV with 2+2 central mac | T. leclerci Deharveng, 1990 |
14 | Labial formula: M1M2REL1l2; body length 1.1–1.2 mm | T. maungonensis Deharveng & Gers, 1993 |
– | Labial formula: M1M2ReL1l2; body length 1.3–1.75 mm | T. maffrei Deharveng & Gers, 1993 |
We thank Rueangrit Promdam, Areeruk Nilsai, Mattrakan Jitpalo and Kanchana Jantapaso for offering various help in the field. We are most grateful for Martin Ellis for proof-reading the English. We also thank the Division of Biological Science (Biology), Faculty of Science, Prince of Songkla University, and PSU-NHM for providing the facilities and support. This study was supported by the Faculty of Science Research Fund, Prince of Songkla University (contract no. 1-2561-02-001), the National Science and Technology Development Agency (FDA-CO-2563-11031-TH), and the Thailand Research Fund (RSA6280063).