TAXONOMIC REVISION AND BIOGEOGRAPHY OF OPEN-AIR PROCESSIONAL COLUMN TERMITES (TERMITIDAE, NASUTITERMITINAE) FROM PAPUA

Transcription

1 LAPORAN TAHUNAN KERJASAMA LUAR NEGERI DAN PUBLIKASI INTERNASIONAL (KLN) TAXONOMIC REVISION AND BIOGEOGRAPHY OF OPEN-AIR PROCESSIONAL COLUMN TERMITES (TERMITIDAE, NASUTITERMITINAE) FROM PAPUA Tahun ke 1 dari rencana 3 tahun Dr. Syaukani, S.Si, M.Sc NIDN Dr. Djufri, M.Si NIDN UNIVERSITAS SYIAH KUALA Oktober 2017

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3 CHAPTER I INTRODUCTION Research Background Termites are the most important arthropod decomposers in the lowland rainforest ecosystem (Collins 1983, 1989) and are also directly or indirectly responsible for soil movement and organic matter translocation (Matsumoto and Abe 1979; Abe 1980; Collins 1989; Jones 1996; Gathorne-Hardy, Syaukani and Inward 2006). Indonesian archipelagoes are inhabited by the highest species number of open-air processional column termites, especially in rainforest of Sumatra and Borneo (Tho 1992; Eggleton 2001; Syaukani 2010), and their species distributions have been correlated with past continent history (Gathorne-Hardy et. al.2002). Taxonomy and systematics of those open-air processional column termites are very poorly known, and many complicated problems still remain. In many publications (Haviland 1898; Holmgren 1914; John 1925; Kemner 1934; Snyder 1949; Ahmad 1958; Thapa 1981 and Tho 1992) most species descriptions are very incomplete, with confusing keys. Furthermore no molecular examination has been made so far to test the validity of delicate morphological characters and to analyze intraspecific variation. Hence a revision of the groups with both morphological and molecular data is urgently needed for further taxonomic understanding, which should help biodiversity assessment and conservation management. Our cooperations in termite biology have been continuing since 2001 to present. Our cooperative study plan in termite taxonomy has been implemented through a number of field surveys to Southeast Asian forests, providing service to identifications, finding new taxonomic characters, and managing a number of refference collections. The significance of these joint works have been demonstrated by producing a systematic key for the Nasutitermitinae based on worker characters (Syaukani 2006), identification of wood-feeding Nasutitermitinae (Syaukani 2011), and discovery of three new termite species 1

4 within this group collected from Indonesia (Syaukani 2008, 2010; Syaukani et al. 2011) (as well as many other new species that are in preparation for publication). Track Record in Research Cooperation Joint research with Prof. Yamane has been continued since 2001, when a field survey to northern Sumatra was conducted. I have conducted studies (master's and PhD programs) in Japan supported by a Japanese government scholarship under his supervision. Prof. Yamane often involved me as a counterpart in addition to researchers from LIPI in a number of surveys in Indonesia. He also helped me to obtain research funding from the NEF (Japan) to write a book about the taxonomy of termites (Nasutitermitinae) from the Kerinci Seblat National Park. He was also the editor of the book. He has also helped Indonesian researchers by providing reasearch equipment, help in identification, corrections of manuscripts, and variable recommendation. 2

5 CHAPTER II LITERATURE REVIEW General introduction to termites Termites are eusocial insects belonging to the order Isoptera (Pearce 1997), live in small to large colonies (colonies of some species containing more than a million individuals) (Chhotani 1997). They are commonly known as pests, causing damage to wooden buildings, both dead and living trees in the forest, and also agricultural crops (Roonwal & Chhotani 1989). They have been found to be the most important arthropod decomposers in the tropical forest ecosystem (Collins 1983, 89; Wood & Sands 1978; Gathorne-Hardy et al. 2001). Termites are thought to have originated on the earth around 100 million years ago (Pearce 1997), and fossils of the ancestral Nasutitermitinae are known since 65 million years ago (Grimaldi & Engel 2005). Taxonomically, termites are closely related to the wood-eating cockroach (Pearce 1997). The most primitive termites seem to be closely related to subsocial cockroaches because they share similar microorganisms that digest cellulose. As many as 25 species of protozoan have been found in Mastotermes darwiniensis (a mastotermitid termite) and Cryptocercus punctulatus (a cryptocercid cockroach) (Wilson 1982). A colony of termite consists of workers (or pseudoworkers), soldiers, a king and a queen, and occasionally mymphs and alates (new kings and queens) (Watson & Gay 1991; Pearce 1997). They are morphologically and behaviourally specialized in particular tasks (Lee & Wood 1971; Grimaldi & Engel 2005). Workers and soldiers are sterile castes, in which the development of reproductive organs is suppressed, while kings, queens and alates contribute to reproduction (Lee & Wood 1971; Pearce 1997). Nymphs represent a young stage of reproductives. The developmental pass-ways to these castes are very complicated and often flexible (Miura 2004). Termites are generally known as wood-feeders and their diets are actually quite diverse (Miura & Matsumoto 1998). Higher termites feed on more diverse 7

6 plant materials than lower termites: dead wood, leaf litter, humus, fungi, lichens, dungs and grasses (Wood 1978; Noirot 1992; Miura & Matsumoto 1998). The termites are divided into two major groups based on the associated symbionts in ther hind gut (Bignell and Eggleton 1995). The lower termites have associated with protozoans, while the higher termites have associated with Bacteria (Krishna 1969; Brauman et al. 2001). More than species belonging to 281 genera have been described worldwide in seven families (Mastotermotidae, Kalotermitidae, Termopsidae, Hodotermitidae, Serritermitidae, Rhinotermitidae and Termitidae). The lower termites comprise the first six families, and the higher termites comprise a single family, Termitidae (Kambhampati & Eggleton 2000). As many as 5 families (Kalotermitidae, Termopsidae, Hodotermitidae, Rhinotermitidae and Termitidae) have been found in the Oriental region (Roonwal 1970), and only 3 families (Kalotermitidae, Rhinotermitidae and Termitidae) have been recorded in the Indo- Malayan subregioan (Ahmad 1965; Thapa 198; Tho 1992). Family Termitidae Westwood Family Termitidae is most advanced in social structure and with the highest species number among the seven families. This family consists of approximately 85% of all the known termite species (Krishna 1970; Chhotani 1997; Collins 1989; Kambhampati & Eggleton 2000). Most of the species under the family are confined to ropical areas (Bess1970). Four subfamilies are recognized in this family: Amitermitinae, Apicotermitinae, Macrotermitinae and Nasutitermitinae. Subfamily Nasutitermitinae Hare As the largest subfamily among the higher termites (Family Termitidae), Nasutitermitinae include over 63 genera (Collins 1989) and 550 species (Emerson 1955), with all presently recognized feeding types (Eggleton 2000). This subfamily probably originated in the Neotropical region during the Cretaceous period (Emerson 1955) and features a highly specialised morphology (Collins 1984, 89), 8

7 with the greatest development of defensive nasutes (Deligne et al. 1981; Prestwich 1984). This subfamily was erected by Hare (1937) with an emphasis on the presence of a remarkable specialization, especially the degeneration of mandibles in the soldier accompanied by the development of frontal gland and rostrum (nasus). This is depicted by the series beginning with Syntermes and ending with the highly modified Nasutitermes, Subulitermes and Convexitermes. Nasutitermes is the type genus of Nasutitermitinae (Snyder 1949; Chhotani 1997). A series of publications by Emerson (1949, 1952, 1955) and Ahmad (1950) provided evidence for separate origins of nasute soldier in two major branches of Nasutitermitinae, the Subulitermes-branch (soil-feeding group) and Nasutitermesbranch (non-soil-feeding group), based on imago/worker mandibles. The open-air processional columns termites belong to the Nasutitermes-braanch. Biology of open-air Processional Column Termites The open-air processional column termites consisting of three genera: Hospitalitermes, Lacessititernes and Longipeditermes. Comparing with Lacessititermes and Longipeditermes, the Hospitalitermes has been studied extensively because its unique foraging and feeding behavior (Escherich 1991; Kalshoven 1958; Roonwal 1970; Collins 1979; Jones and Gathorne-Hardy 1995; Miura and Matsumoto 1997; Jones and Brandell 1998). Most termites forage in protected environments (within wood nest, in the soil, or in the galleries), however the members of this group forage above ground or on leaf litter in processional columns (Tho 1992; Jones and Gathorne-Hardy 1995; Miura and Matsumoto 1998). The worker caste of Lacessititermes is thought to be trimorphic (Tho 1992) or polymorphic (Syaukani 2008), while the worker caste of Hospitalitermes is apparently dimorphic (Prasad and Sen-Sarma 1960; Tho 1992) or trimorphic (Miura and Matsumoto 1998). Each morph has a different task. Major worker act as food carrriers in the earlier stage of foraging activity, medium sized workers 9

8 become the dominant food ball carriers in the later stage, and minor workers play the role of gnawers. Like a majority of termites the soldier caste of Hospitalitermes is monomorphic (Snyder 1949), however Syaukani (2009, 2010) found more than one size soldier caste in colonies. The soldier caste of Lacessititermes monomorphic, dimorphic, and trimorphic with small but distinct size difference in most species (Tho 1992; Syaukani 2010). There are two types of soldier caste (dimorphism) are found in Longipeditermes: large and small (Thapa 1981; Tho 1992; Gathorne-Hardy 2001; Miura and Matsumoto 1998). In Hospitalitermes the column of termites leaves their nest in the late afternoon. The nasute soldiers are the first to move out from the nest. The column of mixed workers and soldiers follow closely and in the initial stage there is much vacillation, with termites moving in and out. The column builds up and begins to move forward in a continuous stream, deploying soldiers to guard the flanks (Collins, 1979). Hospitalitermes often march in enormours numbers, with individuals of H. umbrinus involved in foraging excursions (Collins 1979). The distance from nest to feeding area varies horizontally from a few meters to about 40 meters (Abe 1979) which is also the distance Collins (1979) recorded for H. umbrinus. Jander and Daumer (1974) measured a continuous column upto 300 meters long. The termites of Hospitalitermes feed mainly on living lichen and bryophytes, unlike other termites, which generally feed on dead wood or soil. After foraging throughout the night, the columns return to the nest in the following morning with balls of food. Collins (1979) found two types food balls: light and dark coloured. The light balls contained wood constituents, bryophytes, blue-green algae and fungal hyphae while the dark balls consisted mainly of lichen and their spores. Unlike Hospitalitermes, very little has been published on the foraging activity of Lacessititerms and Longipeditermes (Hoare and Jones 1998). Lacessititermes are 10

9 known to feed on leaf litter, twigs, and lichen-bark, and forage mainly at night and in the early morning hours (Tho 1992). The nests of Hospitalitermes are generally on the ground, usually in buttress roots, in the hollow of big stumps, or fallen trees, but they may also be on large tree-trunks (Haviland 1998; Kalshoven 1958). Elwood et. al. (2002) found nest of Hospitalitermes in fern removed from crown of Parashorea tomentalla (Dipterocarpaceae) at heights between 39 and 52 meters in Danum Valley, Sabah (Malaysia). Furthermore, Hospitalitermes make nest on the basal part of the living tree trunks and sometimes in standing dead trees. Syaukani (unpublished data) found a nest of Hospitalitermes made from soil for the first time. The nest was standing alone on the ground, mound-shaped, blackish in color, covering by lichens on the outside surface, and enlarged in size from year to year in Sumatra. Lacessititermes is known to make arboreal nest around small branches of trees, the nest being constructed of a soft, brown carton material (Syaukani 2008). Nests of Longipeditermes found in very decayed stump trees and in hollow of fallen decayed trees in Peninsular of Malaysia (Syaukani, unpublished data). Taxonomic position of the open-air processional column termites The genera of Lacessititermes, Hospitalitermes, and Longipeditermes are belonging to the subfamily Nasutitermitinae. As many as 37 species for Hospitalitermes (Table 1), 18 species for Lacessititermes (Table 2), and three species for Longipeditermes (Table 3) were collected from various regions up to present. Lacessititermes and Hospitalitermes are restricted to the Indo-Malayan subregion and Papuan region, while Longipeditermes is restricted to the Indo- Malayan subregion (Eggleton 2001; Syaukani 2008). Lacessititermes was previously thought only occur in the Indo-Malayan subregion before a species of the genus found in island of Sulawesi in 1985 (Gathorne-Hardy et. al. 2000). The soldiers of Hospitalitermes have a constricted head behind antennal sockets. Because of the shape of the head, this genus is thought to be closely related to Bulbitermes and Longipeditermes (found in the Indo-Malayan 11

10 subregion), Grallotermes and Lacessititermes (found in Indo-Malayan subregion and Papuan region), and to Constrictotermes (restricted to Neotropical region). The soldiers of Hospitalitermes and Lacessititermes can be distinguished from Bulbitermes by having much longer legs and antennal segments. Longipeditermes has distinctive morphology and dimorphic soldiers wich contrast greatly in size. The soldiers of Grallotermes have 13 antennal segments compare to 14 antennal segments in Hospitalitermes and Lacessititermes, and much shorter and more broad-based nasus. So far three species have been described for the Longipeditermes in the Indo-Malayan subregion. There are numerous records for Longipeditermes longipes which is widespread in southern Thailand (Hoare and Jones 1998), Peninsular Malaysia (Tho 1992), Borneo (Collins 1984) and Sumatra (John 1925; Gathorne-Hardy, Syaukani and Eggleton 2001). L. kistneri was collected from a single colony only from Java (Ahmad and Akhtar 1985); L. mandibulatus was collected from Sabah (Thapa 1981), Sumatra (Syaukani 2008), and Peninsular Malaysia (Syaukani, unpublished data). Taxonomic history and the systematic problem of the open-air processional column termites Haviland (1898) and Holmgren (1913; ) were among the first to undertake taxonomic studies of open-air processional column termites. Haviland (1898) recognized 21 groups/sections under the name Termes Linn., with three groups that are today assigned to Hospitalitermes, Lacessititermes, and Longipeditermes. Termes hospitalis (currently Hospitalitermes hospitalis) as type spesies for Hospitalitermes and Termes lacessitus (currently Lacessititermes lacessitus) as type spesies for Lacessititermes. Later Snyder (1949) was assigned Termes longipes (currently Longipeditermes longipes) as type species for Longipeditermes. More than a century before Haviland s (1898) work on termites, there was only one described species of Hospitalitermes by Koenig (1779) for Termes monoceros. Then after Haviland, Wasmann (1902) described a species of Hospitalitermes, Eutermes flaviventris from Malaca (Malay Peninsular). 12

11 Holmgren (1913) transferred Haviland s Lacessitus group under genus Eutermes subgenus Lacessititermes (and added four new species to this group), then transferred Hospitalitermes group under genus Eutermes subgenus Hospitalitermes (and added a new species, E. (H.) buttelli from Sumatra). Later, Oshima (1923) raised subgenus Hospitalitermes to a genus Hospitalitermes, and subgenus Lacessititermes to a genus Lacessititermes by Light (1930). Four keys to the species based on color and size characters for Hospitalitermes published by Holmgren (1913), Ahmad (1958), Prashad and Sen-sarma (1960), Morimoto (1973), and two keys for Lacessititermes published by Holmgren (1913) and Ahmad (1958). However the colour of preserved specimen can fade with time, then it is difficult to use those keys that relay on colour. 13

12 CHAPTER III OBJECTIVE AND BENAFIT Specific Research Objectives 1. To revise taxonomy of open-air processional column termite species on Borneo. Taxonomy of the open-air processional column termite species has been debated for a long time. In many published papers, neither the original authors nor any subsequent reviewers have given a satisfactory diagnosis or description for each taxon so far. 2. To reconstruct the phylogeny of open-air processional column termite species on. Phylogenetic relationships among the open-air processional column termite species are not yet established for the Indonesian fauna. Among the existing problems the status of ambiguous genera, especially of Hospitalitermes to Lacessititermes requires a quick solution. 3. To provide an identification key to the open-air processional column termite species from Borneo based on the combination of morphological and molecular characters would be a first effort in termite taxonomy. This effective key can be used not only by termitologists but also by naturalists in general. An identification guide at species level with full-colour pictures is urgently needed to promote ecosystem surveys. 4. To reveal biodiversity and biogeography of the open-air processional column termite species from Borneo. Information on termite species richness and their distribution is needed to promote ecosystem surveys worldwide. 5. To increase the number of publications by Indonesian researchers in highly reputated international journals, and to trig Indonesian researchers to compete in international research grants. 6. To deposit specimens in appropriate institutions (museum, university, research centre etc) for future research. 7. To strengthen cooperation between Japanese and Indonesian researchers through joit research and cooperation, especially in termite biology. 14

13 CHAPTER IV RESEARCH METHODS Field Sampling The present survey of open-air processional column termites have cover a large area on Papua, a variety of habitat types and altitudes. Termites collected by finding colonies and manual sampling (Jones and Eggleton 2000; Gathorne-Hardy et al 2000; Syaukani 2008, 09, 10). Termite colonies searched for at any potential site for nesting, and then nest of each colony documented and photographed. Nests opened and termites collected. All the specimens collected kept in 80% ethanol for morpholical examination, and several individuals kept in absolute ethanol for molecular analyses. All the castes and sexes were sampled and were labeled. Other biological data and behavioral information are also documented with manual and digital systems Laboratory works Identification Specimens from the field mainly identified at Syiah Kuala University and Museum Zoologicum Bogoriense (Indonesia). A number of identified termite specimens in the author s reference collection would be much helpful in identifying materials. Final taxonomic confirmation has beenn made at the American Museum of Natural History (New 15

14 York) or the Natural History Museum (London) with the help of termite experts there. 1. Termites sorted into genera and species based on morphological characters of the soldier and worker, nesting and foraging biology, and biogeographical information. 2. Taxonomic fidelity from morphological characters verified using individualspecific mitochondrial DNA (mtdna) cytochrome oxidase II (COII) gene sequence. 3. DNA extraction: Individual worker termites dissected to provide either the head or head and thorax both for molecular analyses. DNA then will be extracted from an individual termite using the E.Z.N.A Mollucs DNA kit (Omega Bio- Tek, Inc., Doraville, GA). 4. Amplification and sequencing: The polymerase chain reaction (PCR) will be used to amplify a ~800-bp fragment of the COII gene from each termite. Primer TL2J3037 and TKN3785 (Liu and Beckenbach 1992, Simon et al. 1994, Jenkins et al. 1999) e used to amplify the fragment and prime 36 forward and reverse sequencing reactions. 5. Data analyses: Individual for electropherograms will be edited using Sequances software (Gene Codes Corp., Ann Arbor, MI) Producing termite digital images Soldier body, head and pronotum, and worker head photographed with a digital microscope, KEYENCE HF VH Worker mandibles and labrum, antennae of soldier and worker will be removed and mounted on slide glasses with Euparal 3C 239 (Waldeck GmbH & Co. KG) (Syaukani 2008). Then photographs will be taken by a Nikon Coolpix 3340 digital camera attached to a Nikon Eclipse E600. Multi-focused montage images produced using Helicon Focus 4.03 Pro. Artifacts/ghosts and unnecessary parts (unfocused appendages, shaking alcohol effects, etc.) surrounding or covering target objects will be erased and cleaned up using the retouching function of Helicon Focus. Finally the background will be 16

15 cleaned up, and the colour balance, contrast and sharpness will be adjusted using Adobe Photoshop CS3-Ext Terminology and Measurement Morphological terminology that used for the soldier and worker follows Ahmad (1950), Roonwal and Chhotani (1989), Thapa (1981), Tho (1992), Sands (1998), Gathorne-Hardy (2001), Syaukani (2008; 2011), and Syaukani et al. (2011). Probably several new terms will be introduced. In the keys and species descriptions the largest soldiers (and major soldiers in dimorphic species) and largest workers will be principally used to keep consistency and partly because the largest individuals have more useful characters than smaller individuals. The number of individuals that will be measured varies among the genera/species. Soldier head (Fig. 1) The shape (mainly pear-shaped, sometimes oval, round, rectangular, or pumpkin-like) and the size of the head greatly vary among genera and species. Coloration and sometimes pilosity are useful, especially for species level sorting. Other characters include: the condition of the posterior margin of the head in dorsal view (round, straight, or indented), the dorsal outline of the head in profile, and the condition of the constriction that divides the head into anterior and posterior parts. Soldier rostrum (nasus) (Fig. 1 & 2). Shape (conical or cylindrical; straight or curved; direction), length, and coloration (uniformly colored or apex/base with different colours) of the rostrum are useful in separating genera and/or species. Pronotum (Fig. 3). Conditions of the anterior and posterior margins of the pronotum in dorsal view (nearly straight or indented in the middle) are useful in soldiers. General shape and coloration are also often useful. Antennae (Fig. 4 & 5). Antenna is moniliform (with bead-like segments), uniformly or differently colored among segments; generally basal segments, i.e., the first and second, are darker than the subsequent segments. The number of segments (generally 11-15) and their relative length and shape are important at genus and species level sorting. The first or basal segment (scape) is long, the second (pedicel) shorter than the first, and the subsequent segments 17

16 (flagellomeres) very variable in length and shape among the species. The separation between neighboring segments is often not clear. Worker mandibles (Fig. 6). The characters pertaining to worker mandibles are extremely important for the phylogenetic study of termites. Generally they are quite stable and useful for genus level taxonomy. Character states are almost same in the imago (alates) and worker of a given species. In both castes the left and right mandibles are asymmetrical, and this supplies more characters than do other insect groups in which the mandibles are symmetrical. Left mandible consists of the apical tooth, 1 st -4 th marginal teeth and molar prominence. Right mandible consists of the apical tooth, fewer marginal teeth (1 st and 2 nd ), molar plate and its inner layer; the position and condition of the inner layer are important; the molar plate has a notch at its proximal end, and the condition of the notch can be useful. 18

17 1 st 5 tibia pronotum head rostrum abdominal tergite femur coxa antenna 1 6 th 5 th 4 th 3 rd Labial palp mandible 2 nd 1 st 4 constriction posterior margin 2 5 th 4 th 6 th Anterior margin 3 rd 2 nd Posterior margin 3 antennae Figs Soldiers and worker of open-air processional column termite. Soldier body in profile (1), head capsule in dorsal view (2), pronotum seen from above (3), soldier (4) and worker (5) antennae. 20

18 Cutting edge Apical tooth 1 st marginal tooth 3 rd marginal tooth 4 th marginal tooth Molar prominence Left mandible Apical tooth 1 st marginal tooth 2 nd marginal tooth Molar plate Anterior edge Right mandible Posterior edge Inner layer of molar plate Notch Right mandible Fig. 6. Morphology of worker mandibles in the Nasutitermitinae 21

19 anterior part b posterior part a antennae e h Fig. 7. Measured body parts of the open-air processional colomn termite (soldier). Pronotum width (a), and length (b); maximum head width at the anterior part (c), and at the posterior part (d); head capsule length including rostrum (e); rostrum length (f); maximum height of head excluding postmentum (g), and head length measured to base of mandibles (HL) (h). 22

20 CHAPTER V RESULTS As many as two genera and 8 species have been examined during the first survey of the open-air processional columns termites in Kalimantan. The three genera are: Lacessititermes and Hospitalitermes. Genus Hospitalitermes Holmgren Hospitalitermes Holmgren, pp. 48, 60, 62, 66, 72 (as subgenus of genus Eutermes) Genus Hospitalitermes: Oshima, Genus Hospitalitermes: Snyder Genus Hospitalitermes: Ahmad Genus Hospitalitermes: Chhotani Hospitalitermes, as well as Lacessititermes and Longipeditermes, is one of the genera consisting of free-ranging species (Tho 1992), forages above the ground in exposed processional columns (Jones & Gathorne-Hardy 1995), with up to 500,000 individuals in a foraging excursion (Collins 1979). Hospitalitermes, probably paraphyletic (Prasetyo 2007), is slightly similar to Lacessititermes in general morphology (Miura & Matsumoto 1998). Hospitalitermes is characterized by the following features: in the soldier head capsule pale brown to black, with anterior part paler than or darker than the posterior part; lighter part of the rostrum variable in area; antenna much paler than or similar to head capsule in coloration, generally with basal segments darker than the subsequent; pronotum paler than or similar to head capsule in coloration; abdominal tergites generally whitish brown to sepia brown; coxae and femora pale brown to black; tibiae whitish yellow to brown. In dorsal view, head capsule weakly to strongly constricted behind antennal sockets, with the anterior part excluding rostrum smaller than the posterior part in size; its posterior margin nearly straight to roundly convex in dorsal view; dorsal outline (including rostrum) in profile weakly indented to moderately concave; rostrum variable in shape and length; antenna with 14 segments that are variable in length and shape. 23

21 In the worker, antenna whitish yellow to dark sepia brown and 15- segmented; generally apical tooth of left mandible shorter than or equal to first marginal tooth; third weakly to moderately protruding from cutting edge; fourth hidden behind molar prominence; posterior edge of right mandible nearly straight to weakly concave; inner layer of molar plate nearly straight to strongly convex. Tho (1992) mentioned that Hospitalitermes has a dimorphic worker caste, while Miura (2004) found three types of worker (major, medium and small) for H. medioflavus. Lacessititermes is hardly distinguished from Hospitalitermes, except in the worker caste by the absence of notch at molar plate of the right mandible. As many as 36 species of Hospitalitermes have been described from Indo- Malayan and Austro-Malayan subregions (Prasetyo 2007), and eight species have been recorded from Sumatra (Snyder 1949). Among them, two species, H. butteli (Holmgren) and H. nemorosus Ghidini, were transferred to Lacessititermes (Prasetyo 2007). Hospitalitermes umbrinus (Haviland) Termes umbrinus Haviland, p Eutermes umbrinus: Wasman, p Eutermes (Hospitalitermes) umbrinus: Holmgren, p Hospitalitermes umbrinus: Snyder, p Hospitalitermes umbrinus: Collins Hospitalitermes umbrinus: Tho Descriptio Largest soldier Head: head capsule brown to sepia brown, with anterior part slightly paler than the posterior part; rostrum with apical third lighter and basal two-thirds darker in coloration; antenna much paler than head capsule, not uniformly coloured, (except for the first and second segments) yellow to pale brown. Head capsule in dorsal view strongly constricted behind antennal sockets, with anterior part excluding rostrum clearly smaller in size than posterior part; its posterior margin nearly straight; dorsal outline (including rostrum) in profile moderately concave; rostrum 24

22 equal to haft the length of head capsule or more; antenna with 14 segments; third segment longer than fourth and fifth; fourth and fifth almost equal in length; 6 th - 14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above slightly paler than head capsule in coloration, with the paler area larger than darker area on the disc; pronotum with anterior margin nearly straight, and posterior margin very weakly indented in the middle; coxae sepia brown; femora pale brown to brown; tibiae yellow to pale brown. Abdomen: tergites pale brown to brown in coloration. Worker Antenna: pale brown to brown except for the first and second segments; 15- segmented; third segment slightly shorter than or equal to fourth; fourth and fifth almost equal in length; 6 th -15 th gradually becoming elongate toward apex. Left mandible: apical tooth much shorter than first marginal tooth; third marginal strongly protruding from cutting edge; fourth completely hidden behind molar prominence. Right mandible: first marginal tooth with a minute bulge basally in anterior part; posterior edge of second marginal nearly straight; inner layer of molar plate moderately convex Hospitalitermes diurnus Kemner Hospitalitermes diurnus: Snyder Hospitalitermes diurnus Kemner Hospitalitermes diurnus: Roonwal & Maiti, Soldier Head: head capsule pale brown to sepia brown, with anterior part much paler than the posterior part; rostrum darker than anterior part and paler than posterior part of head capsule; antenna dark yellow with first segment darker than the subsequent, paler than the posterior part of head capsule in coloration. Head capsule in dorsal view strongly constricted behind antennal sockets, with anterior part excluding rostrum clearly smaller in size than the posterior part; its posterior margin weakly to moderately indented in the middle; dorsal outline (including rostrum) 25

23 in profile weakly concave; rostrum more than half as long as head capsule; antenna with 14 segments; third segment longer than fourth and fifth; fourth and fifth almost equal in length; 6 th -14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above paler than posterior part of head capsule, with the pale area larger than darker area on the disc; its anterior and posterior margins weakly indented in the middle; coxae brown; femora pale brown to brown; tibiae yellow to yellowish brown. Abdomen: tergites pale brown to brown in coloration. Largest worker Antenna: yellow to pale brown except for the first segment; 15-segmented; third segment longer than fourth and fifth; fourth and fifth approximately equal in length; 6 th -15 th gradually becoming elongate toward apex. Left mandible: apical tooth much shorter than first marginal tooth; third marginal moderately protruding from cutting edge; fourth almost completely hidden behind molar prominence. Right mandible: posterior edge of first and second marginal teeth weakly concave; first marginal tooth with a minute bulge basally in anterior part; posterior edge of second marginal tooth nearly straight; inner layer of molar plate strongly convex. Hospitalitermes hospitalis (Haviland) Termes hospitalis Haviland, p Eutermes monoceros hospitalis: Wasmann, p Eutermes (Hospitalitermes) hospitalis: Holmgren, p Eutermes (Hospitalitermes) hospitalis: John, pp Hospitalitermes hospitalis: Snyder, p Hospitalitermes hospitalis: Ahmad, p

24 1992. Hospitalitermes hospitalis: Tho, p Description Largest soldier Head: head capsule brown to dark sepia brown with anterior part slightly paler than posterior part; rostrum with apical two-thirds lighter, and basal third darker; antenna (except for the first and second segments) uniformly pale brown to sepia brown, and paler than or similar to the posterior part of head capsule in coloration. Head capsule in dorsal view weakly to moderately constricted behind antennal sockets, with anterior part excluding rostrum clearly smaller in size than posterior part; its posterior margin weakly to moderately indented in the middle; dorsal outline (including rostrum) in profile weakly to moderately concave; rostrum more than half as long as head capsule; antenna with 14 segments; third segment longer than fourth; fourth slightly shorter than or equal to fifth; 6th-14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above paler than or similar to head capsule in coloration, with anterior and posterior margins weakly indented in the middle; coxae brown to sepia brown; femora pale brown to brown; tibiae pale brown. Abdomen: whitish brown to pale brown. Largest worker Antenna: first antennal segment blackish brown to black, and darker than the subsequent segments; consisting of 15 segments; third segment longer than fourth; fourth and fifth almost equal in length; 6 th -15 th elongate toward apex. gradually becoming Left mandible: apical tooth shorter (often clearly shorter) than first marginal tooth; third marginal moderately protruding from cutting edge; fourth completely hidden behind molar prominence. Right mandible: posterior edge of second marginal tooth nearly straight; inner layer of molar plate weakly to moderately concave. 27

25 Hospitalitermes medioflavus (Holmgren) Eutermes (Hospitalitermes) forma medioflavus Holmgren, pp Hospitalitermes medioflavus: Kemner, pp Hospitalitermes medioflavus: Snyder, p Hospitalitermes medioflavus: Ahmad, p Description Largest soldier Head: head capsule brown to dark sepia brown, with anterior and posterior parts similar in coloration; rostrum with apical third lighter, and basal two-thirds darker; antenna (except for the first and second segments) uniformly pale brown to brown, and paler than head capsule. Head capsule in dorsal view weakly constricted behind antennal sockets, with anterior part excluding rostrum extremely smaller in size than posterior part; its posterior margin of head capsule roundly convex; dorsal outline in profile weakly concave; rostrum more than half as long as head capsule length; antenna with 14 segments; third segment longer than fourth and fifth; fourth and fifth approximately equal in length; 6 th -14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above slightly paler than or similar to head capsule in coloration, with the paler area on the disc located close to the posterior part; pronotum with anterior margin nearly straight, and posterior margin weakly indented in the middle; coxae brown to sepia brown; femora pale brown to brown; tibiae dark yellow to pale brown. Abdomen: tergites pale brown to dark sepia brown. Largest worker Antenna: pale brown to brown except for the first and the second segments; consisting of 15 segments; third segment longer than fourth and fifth; fourth and fifth approximately equal in length; 6 th -15 th gradually becoming elongate toward apex. Left mandible: apical tooth shorter than first marginal tooth; third marginal moderately protruding from cutting edge; fourth almost completely hidden behind molar prominence. Right mandible: posterior edge of second marginal tooth straight or weakly concave; inner layer of molar plate weakly convex in anterior part. 28

26 Hospitalitermes sp. A Description Largest soldier Head: head capsule reddish brown to blackish, with anterior part reddish brown and posterior part dark sepia brown to blackish; rostrum with almost entirely lighter than head capsule; antenna similar to anterior part of head capsule. Head capsule in dorsal view strongly constricted behind antennal sockets, with anterior part excluding rostrum much smaller than posterior part in size; posterior margin of head capsule strongly emarginate in the middle; dorsal outline (including rostrum) in profile weakly concave; rostrum more than half as long as head capsule; antenna with 14 segments; third segment clearly longer than fourth; fourth and fifth almost equal in length; 6 th -14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above paler than head capsule in coloration, with anterior and posterior margins weakly indented in the middle; coxae and femora brown; tibiae pale brown. Abdomen: tergites whitish brown. Largest worker Antenna: first and second antennal segments pale brown to brown and darker than the subsequent segments; consisting of 15 segments; third segment slightly longer than fourth; fourth and fifth almost equal in length; 6 th -15 th gradually becoming elongate toward apex. Left mandible: apical tooth clearly shorter than first marginal tooth; third marginal moderately protruding from cutting edge; fourth completely hidden behind molar prominence. Right mandible: posterior edge of second marginal tooth nearly straight; outline of molar plate weakly to moderately convex. 29

27 Genus Lacessititermes Holmgren Lacessititermes Holmgren, pp , 65 (as subgenus of genus Eutermes) Genus Lacessititermes: Light, pp. 16, 17, 19, Genus Lacessititermes: Tho, pp Lacessititermes is characterized by the following features: in the soldier head capsule pale brown to black, generally with anterior part paler than posterior part; lighter part of rostrum variable in area; antenna much paler than or similar to head capsule in coloration, generally basal segments being darker than the subsequent; pronotum paler than or similar to head capsule in coloration; abdominal tergites pale yellow to dark sepia brown; coxae yellowish to dark sepia brown, femora yellow to black; tibiae whitish yellow to pale brown. Head capsule in dorsal view very weakly to moderately constricted behind antennal sockets, with its posterior margin nearly straight to roundly convex; dorsal outline (including rostrum) in profile moderately to strongly concave; rostrum variable in shape, more than half as long as head capsule; antenna with 14 segments that are valiable in length and shape. In the worker antenna yellow to pale brown, with 15 segments; apical tooth of left mandible generally shorter than first marginal tooth; third marginal weakly to moderately protruding from cutting edge; fourth almost completely hidden behind molar prominence; posterior edge of right mandible nearly straight to moderately concave; inner layer of molar plate nearly straight to weakly convex. As many as 16 species have been described in the genus Lacessititermes from the Indo-Malayan subregion of the Oriental region and the Austro-Malayan subregion of the Australian region (Prasetyo 2007). Among them L. laborator (Haviland), L. jacobsoni Kemner and L. artrior (Holmgren) have been collected from Sumatra (Holmgren ; John 1925; Kemner 1930; Snyder 1949; Prasetyo 2007). Recently two Hospitalitermes species, H. butteli and H. nemorosus, were transferred to Lacessititermes (Prasetyo 2007). Lacessititermes albipes (Haviland) Termes albipes Haviland, p Termes (Lacessititermes) albipes: Holmgren, p Lacessititermes albipes: Snyder, p Description 30

28 Largest soldier Head: head capsule entirely brown to sepia brown; rostrum dark brown with a reddish tip; first antennal segment pale brown to brown, darker than the subsequent, similar to head capsule in coloration. Head capsule in dorsal view weakly constricted behind antennal sockets, with posterior margin weakly indented in middle; dorsal outline (including rostrum) in profile moderately concave; antenna with 14 segments; third segment longer than fourth in length; fifth slightly shorter than fourth; 6 th -14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above same as head capsule in coloration; its anterior and posterior margins moderately indented in the middle; coxae and femora brown; tibiae pale brown to brown. Abdomen: tergites dark sepia brown. 31

29 Lacessititermes laborator (Haviland) Termes laborator Haviland, p Eutermes (Lacessititermes) laborator: Holmgren, p Eutermes (Lacessititermes) laborator: Holmgren, p Eutermes (Lacessititermes) laborator: John, pp Eutermes laborator: Snyder, p. 200 Description Largest soldier Head: head capsule pale brown to sepia brown, with anterior and posterior parts similar in coloration; rostrum medially darker, and apical and basal portions lighter; antenna dark yellow to pale brown, paler than head capsule; first segment darker than the subsequent. Head capsule in dorsal view moderately constricted behind antennal sockets, with posterior margin roundly convex; dorsal outline (including rostrum) in profile moderately concave; antenna with 14 segments; third segment longer than second and shorter than fourth; fourth slightly longer than or equal to fifth; 6 th -14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above pale brown to brown; its anterior margin nearly straight, and posterior margin weakly indented in the middle; coxae pale brown; femora dark yellow to pale brown; tibiae whitish yellow to yellow. Abdomen: tergites brown to dark sepia brown. Largest worker Antenna: yellow to pale brown with the first segment slightly darker than the subsequent; consisting of 15 segments; third to fifth segments approximately equal in length; 6 th -15 th gradually becoming elongate toward apex. Left mandible: apical tooth shorter than first marginal tooth; third marginal moderately protruding from cutting edge; fourth completely hidden behind molar prominence. Right mandible: posterior edge of second marginal tooth weakly concave; inner layer of molar plate nearly straight; notch weakly developed. 32

30 1898. Termes sordidus Haviland, p.434. Lacessititermes sordidus (Haviland) Eutermes (Lacessititermes) sordidus: Holmgren, p Eutermes (Lacessititermes) sordidus: John, pp Lacessititermes sordidus: Snyder, p Description Largest soldier Head: head capsule brown to dark sepia brown, with anterior part slightly paler than or similar to posterior part; median third of rostrum dark brown, with apical and basal parts lighter; antenna yellow to pale brown, with first segment darker than the subsequent. Head capsule in dorsal view moderately constricted behind antennal sockets, with posterior margin roundly convex; dorsal outline (including rostrum) in profile strongly concave; antenna with 14 segments; third segment shorter than or equal to fourth; fourth longer than fifth; 6 th -14 th gradually becoming elongate toward apex. Thorax: pronotum seen from above yellow to pale brown, much paler than head capsule; its anterior margin moderately and posterior margin weakly indented in the middle; coxae and femora yellow to dark yellow; tibiae pale yellow to yellow. Abdomen: tergites yellow to pale brown. Largest worker Antenna: yellow to pale brown, with first segment darker than the subsequent; comprising of 15 segments; third and fifth segments clearly shorter than fourth; fifth shorter than sixth; 7 th -15 th gradually becoming elongate toward apex. Left mandible: apical tooth shorter than first marginal tooth; third marginal moderately protruding from cutting edge; fourth completely hidden behind molar prominence. Right mandible: posterior edge of second marginal tooth nearly straight; inner layer of molar plate weakly convex; notch weakly developed. 33

31 CHAPTER VI FURTHER RESEARCH ACTIVITY (PLAN) No Activity 1 Further specimens identification 2 Worker mandibles examination (morphological characters) 3 Visiting partner research 4 Completing scientific paper 5 Writing text book 6 Completing final report for second year data 34

32 CHAPTER VII CONCLUSION 1. As many as 7 species belonging to three genera of open-air processional column termites are recognized among the material collected from Papua in The two genera are: Lacessititermes and Hospitalitermes 3. Two species are considered to be new to science 35

33 REFERENCES Abe, T Colonization of the Krakatau Islands by termites (Insecta: Isoptera). Physiological Ecology Japan, 21: Ahmad, M The phylogeny of termites based on imago-worker mandible. Bulletin of the American Museum of Natural History, 95: Ahmad, M Key to Indomalayan termites. Biologia, 4: Ahmad, M Termites of Malaysia. I. Nasute genera related to Subulitermes (Isoptera, Termitidae, Nasutitermitinae). Bulletin of the Department of Zoology, University of the Punjab 3: Akhtar, M.S. & Ahmad, M A new nasute termite from Java (Isoptera: Termitidae: Nasutitermitinae). Pakistan Journal Zoology, 17: Chhotani, O. B Fauna of India-Isoptera (Termites) Vol. II. Zoological Survey of India, Calcuta, 800 pp. Collins. N. M Observation of the foraging activity of Hospitalitermes umbrinus (Haviland), (Isoptera, Termitidae) in the Gunong Mulu National Park, Sarawak. Ecological Entomology, 4: Collins. N. M Termite population and their role in litter removal in Malaysian rain forests. In S.L. Sutton, S.L., Whitmore, T.C., Chadwick, A.C. (eds.), Tropical Rain Forest: Ecology and Management. Blackwell Scientific Publication, Oxford, pp Collins. N. M The termites (Isoptera) of the Gunung Mulu National Park, with a key to genera known from Sarawak. Sarawak Museum Journal 30: Collins. N.M Termites. In Lieth, H. and Werger, M.J.A. (eds.). Tropical Rain Forest Ecosystems. Biogeographical and Ecological Studies. Elsevier, Amsterdam, pp Eggleton, P Global patterns of termite diversity. In Abe, T., Bignell, D.E. & Higashi, M. (eds.), Termites: evolution, sociality, symbiosis, ecology. Kluwer Academic Publishers, Dordrecht, the Netherlands, pp Emerson, A.E New genera of termites related to Subulitermes from the Oriental, Malagasy, and Australian regions (Isoptera, Termitidae, Nasutitermitinae). Bulletin of the American Museum of Natural History, 1986: Gathorne-Hardy, F A review of the South-East Asian Nasutitermitinae (Isoptera: Termitidae), with descriptions of one new genus and a new species and including a key to the genera. Journal of Natural History, 35:

34 Gathorne-Hardy FJ, Syaukani, Davies RG, Eggleton P, Jones DT Quaternary rainforest refugia in southeast Asia: using termites (Isoptera) as indicator. Biological Journal of the Linnean Society 75: Gathorne-Hardy, F., Jones, D.T. and Syaukani A regional perspective on the effects oh human disturbance on the termites on Sundaland. Biodiversity and Cpnservation, 11: Gathorne-Hardy FJ, Syaukani, Inward DJG Recovery of termite (Isoptera) assemblage structure from shifting cultivation in Barito Ulu, Kalimantan, Indonesia. Journal of Tropical Ecology 22: Haviland, G.D Observations on termites; with description on new species. Journal of the Linnean Society, Zoology, 26: Holmgren, N Termitenstudien. 3. Systematik der Termiten. Die Familien Mastotermitidae. Kungliga Svenska Vetensskapakademiens Handlingar, 48: Holmgren, N Termitenstudien. 4. Versuch einer systematichen Monographie der Termiten der orientalischen Region. Kungliga Svenska Vetensskapakademiens Handlingar, 50: Holmgren, N Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien, ausgefuhrt im Auftrage der Kgl. Preuss. Akademie der Wissenchaften zu Berlin von H.v. Buttel-Reepen. III. Termiten aus Sumatra, Java, Malacca und Ceylon. Gesammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren Zoologische Jahrbucher Abteilungen Systematik, 36: John, O Termiten von Ceylon, der Malayaischen Halbinsel, Sumatra, Java und den Aru-Inseln. Treubia, 6: Jones, D.T. and Gathorne-Hardy, F Foraging activity of the processional termite Hospitalitermes hospitalis (Termitidae: Nasutitermitinae) in the rain forest of Brunei, north-west Borneo. Insectes Sociaux, 42: Jones, D.T. and Eggleton, P Sampling termites assemblages in tropical forest: testing a rapid biodiversity assessment protocol. Journal of Applied Ecology, 31: Kemner, N.A Fauna Sumatrensis. Brijdrage, 66: Kemner, N. A Systematiche und biologische studien uber die Termiten Javas und Celebes. Kungliga Svenska Vetensskapakademiens Handlingar, 13: Light, S.F Notes on Philippine termites IV. The Philippine Journal of Science, 42: