SYSTEMATICS Redescription of the Drywood Incisitermes inamurae (Isoptera: ) From Southern Taiwan

1,2 3 1 HOU-FENG LI, NATSUMI KANZAKI, AND NAN-YAO SU

Ann. Entomol. Soc. Am. 102(5): 759Ð765 (2009) ABSTRACT Kalotermes inamurae Oshima 1912 (Isoptera: Kalotermitidae) is reassigned to Incisit- ermes (Krishna, 1961) based on the winged imago that is described herein for the Þrst time. The two size morphs of the soldier caste are redescribed, as is the distinctly different morphology of the soldier of incipient colonies. The distribution of I. inamurae is restricted to the tropical zone of Taiwan. Another tropical species, Incisitermes mcgregori (Light), from Luzon, Philippines, has similar soldier morphology and may be a junior synonym of I. inamurae. The Þve Incisitermes species reported in Japan, China, Taiwan, and the Philippines were compared morphologically.

KEY WORDS Incisitermes, winged imago, soldier, , Taiwan

Kalotermes inamurae Oshima 1912 (Isoptera: Kaloter- termes taylori (Light) are found in the Philippines mitidae) was described in brief based on specimens (Light 1921, 1930). Both (Hagen), collected by Tokie Inamura at Kuraur in Heng-Chun found in the central east coast of China (Huang and Peninsula, Taiwan. In 1913, the original description He 2000) and southern Japan (Indrayani et al. 2004, was translated into English (Oshima 1913), but no Mori 1976), and Incisitermes immigrans (Snyder), additional details were provided. Due to the lack of found at Iwojima Island and South Daito Island, Japan winged imagoes, Oshima did not assign this species to (Ohkuma et al. 2004, Takematsu 1997, Yasuda et al. a subgenus. In 1961, Krishna reorganized the genus 2000), were introduced from the New World. The Þve Kalotermes and assigned most of the species to other Asian species are compared here. genera. Although Krishna (1961) questioned the ge- nus of this species, he did not propose a new combi- Materials and Methods nation, probably due to unavailability of winged ima- goes. In light of a recent collection of K. inamurae from Redescriptions and descriptions are based on ex- southern Taiwan, including the type locality, we re- amination of 53 colony samples collected from 28 describe the large and small soldiers, and we also localities, including the type locality in Heng-Chun describe the incipient soldier and the winged imago Peninsula (Fig. 1). were preserved in 85% for the Þrst time. According to the current Kaloter- ethanol. Morphometric data were obtained by using a mitidae taxonomic system (Krishna 1961), a new com- stereomicroscope (model SZX12, Olympus Optical bination for this species is herein proposed as Incisi- Co., Ltd., Tokyo, Japan) Þtted with a calibrated ocular termes inamurae (Oshima). micrometer. The color pictures of each caste taken Incisitermes species, commonly referred to as dry- with a digital camera (DP70, Olympus Optical Co., wood termites, are mostly found in dead trees or dead Ltd.) can be viewed at http://ßrec.ifas.uß.edu/su/ portions of living trees in their native habitats. Due to hou-feng-li.shtml. The terms “large” and “small” sol- their low moisture requirement, some species are able diers (Oshima 1913, Krishna 1961, Scheffrahn et al. to survive in artiÞcial structures. The highest species 2000) are equal to the terms “long-headed” and “short- diversity of Incisitermes (20 species) is found in the headed” soldiers (Banks and Snyder 1920). Measure- Nearctic and Neotropical Americas (Constantino ments were adopted from Roonwal (1969), and the 1998). A few species are known from Australia (Gay color scheme of Sands (1965) was used. Specimens for 1975), India (Roonwal and Chhotani 1989), and Oce- scanning electron micrographs (SEMs) were dehy- ania (Krishna 1961). In East Asia, Þve named Incisit- drated in 95%, 100% ethanol and 1,1,1,3,3,3-hexameth- ermes species including I. inamurae have been yldisilazane for 24 h each consecutively (Nation recorded. Incisitermes mcgregori (Light) and Incisi- 1983). Dehydrated specimens were sputter-coated with platinum, and their images were taken with a 1 Department of Entomology and Nematology, Fort Lauderdale scanning electron microscope (SEM-5510LV, JEOL, Research and Education Center, University of Florida, 3205 College Tokyo, Japan) at 10 kV. Contrast of SEMs was en- Ave., Ft. Lauderdale, FL 33314. hanced by using the software GIMP, version 2.6 (Free 2 Corresponding author, e-mail: houfeng@uß.edu. 3 Forest Pathology Laboratory, Forestry and Forest Product Re- Software Foundation, Inc., Boston, MA), and back- search Institute, 1 Matsunosato, Tsukuba, Ibaraki, 305-8687 Japan. grounds were converted into black. Scale bars were

0013-8746/09/0759Ð0765$04.00/0 ᭧ 2009 Entomological Society of America 760 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5

Fig. 1. (A) Distribution of I. inamurae in Taiwan (gray area). Inset (B), current collection sites (open circle) and the type locality (gray circle). redrawn. Collection localities were mapped by using radius terminating at costal margin at proximal half of the software Google Earth, version 4.3 (Google Inc., wing from suture, radial sector with about eight Mountain View, CA). branches in distal half of wing; media unsclerotized, in some specimens, slightly sclerotized beyond distal one-fourth length of wing; cubitus with Ͼ11 branches Incisitermes inamurae (Oshima) toward posterior margin, several branches close to Calotermes (subg. ?) inamurae Oshima: Oshima 1912: wing scale slightly sclerotized. In ventral view, post- 61Ð63. [female dealate, large soldier, small soldier]. mentum oval, anterior quarter pale yellowish, notice- Kalotermes inamurae: Snyder 1949: 16. ably different from orange yellow posterior part; gena ? Kalotermes inamurae: Krishna 1961: 336. and palps orange yellow. Abdominal sternites and Imagoes (Fig. 2; Table 1). In dorsal view, head femora pale yellow. Tibia, tarsus, and claws orange capsule, antennae, notum, and dorsum orange or or- yellow. Arolia present. ange yellow except for ferruginous compound eyes; Comparison. The noticeably orange head and prono- costal veins including costal margin, subcosta, radius, tum and brownish wing and body of I. minor imago gives and radial sector ferruginous orange; generally giving it a much darker appearance than I. inamurae and I. the winged imago a uniform orange appearance. immigrans in general. The winged imago of I. inamurae Head capsule subcircular. Frons broadly concave resembles that of I. immigrans, but the former is larger. with faint rugosity. Labrum wide linguiform, orange Total length with wings of the I. inamurae alate is Ն13.2 yellow, semitransparent; clypeus narrow, yellow mm compared with Ϸ12.0 mm in I. immigrans. Head white. Compound eye chestnut brown, subcircular, width at eyes in I. inamurae is Ն1.5 mm, whereas in I. protruding and large, contacting antennal socket; immigrans is Ϸ1.1 mm, and pronotum maximum width is ocellus elliptical, yellowish contrasting with orange Ն1.65 mm and Ϸ1.2 mm for I. inamurae and I. immigrans, head capsule, right above compound eye and contact- respectively. Arolia present in I. inamurae and I. immi- ing ocular sclerite. Epicranial suture faint, Y-shaped grans but not in I. minor. The winged imagoes of I. branches ending with ocellus on each side. Antennae mcgregori and I. taylori are unknown. composed of 17Ð19 segments, usually 17 or 18, Þrst Soldier (Figs. 3 and 4; Table 2). Three distinct segment longest, second segment shortest, in general, morphs of soldier caste, large and small forms from ma- the segment length gradually increasing from proxi- ture colonies, and the incipient soldier from young col- mal segments to distal segments, proximal segments onies were found. Material from four mature colonies is orange and grading to yellow at distal segments. Man- represented in our collection, including 10, 11, 28, and 29 dibular dentition as in Fig. 2C. The posterior margin of soldiers, respectively, consisting of 21Ð50% large soldiers the Þrst plus second marginal tooth in left mandible and 50Ð79% small soldiers. In incipient colonies, the Þrst shorter than the anterior margin of the third marginal several soldiers produced were signiÞcantly smaller than tooth, which is the character of the genus Incisitermes soldiers found in mature colonies, a difference also ob- and not of the genus Kalotermes described by Krishna served by the H.-F.L. in I. snyderi and I. schwarzi, and (1961). Pronotum wider than long and wider than reported previously in Neotermes papua (Desneux) head; anterior margin smoothly incised, anterior cor- (Roisin and Pasteels 1991), and Mastotermes darwiniensis ner rounded, posterior margin faintly concave, sides Froggatt (Watson 1974). In general, large soldiers are slightly convex with scattered short and long setae up the most sclerotized and incipient soldiers the least to 0.2-mm-long, posterior corner broadly rounded. sclerotized. Morphometrics of large, small, and incip- Wing membrane transparent and faintly tuberculate; ient soldiers are given in Table 2. costal veins including costal margin, subcosta, radius, Large Soldier (Figs. 3A and B and 4A and B; Table and radial sector sclerotized. Costal margin in forew- 2). Head capsule in dorsal view, rectangular, ferrug- ing with scattered short setae Ϸ0.1 mm, subcosta and inous orange anteriorly grading to yellow posteriorly, September 2009 LIETAL.: REDESCRIPTION OF I. inamurae 761 from 10 ؍ Table 1. Measurements of I. inamurae imagoes, n three colonies including five males and five females

Measurement (mm) Range Mean Ϯ SD No. antennal segments 17Ð19 17.8 Ϯ 0.6 Head length to labrum tip 1.70Ð2.05 1.80 Ϯ 0.12 Head length to medium base 1.30Ð1.60 1.42 Ϯ 0.09 of clypeus Head width, max at eyes 1.50Ð1.65 1.58 Ϯ 0.04 Head ht without postmentum 0.85Ð1.10 1.00 Ϯ 0.08 Labrum width, max 0.58Ð0.70 0.63 Ϯ 0.04 Eye max diam with sclerite 0.40Ð0.47 0.44 Ϯ 0.03 Distance from eye to lower 0.15Ð0.30 0.26 Ϯ 0.04 margin of head, min Ocellus diam, max 0.13Ð0.21 0.17 Ϯ 0.02 Pronotum, max length 1.15Ð1.30 1.26 Ϯ 0.05 Pronotum, median length 1.00Ð1.20 1.08 Ϯ 0.05 Pronotum, max width 1.65Ð1.90 1.78 Ϯ 0.07 Hind tibia, max length 1.18Ð1.35 1.23 Ϯ 0.05 Total length with wings 13.20Ð14.40 13.72 Ϯ 0.38 Total length without wings 7.40Ð8.80 8.12 Ϯ 0.41 Forewing length from suture 10.00Ð10.75 10.35 Ϯ 0.23 Forewing, max width 2.75Ð3.35 2.97 Ϯ 0.17

ners at mandibular articulations. Head capsule when view laterally, covered with dense setae of variable length to 0.2 mm on both dorsal and ventral discs. Frons slopes smoothly from vertex, surface weakly rugose and slightly concave, with Þve to eight long setae at frontal carinae. Clypeus narrow, white, trap- ezoid. Labrum orange yellow, linguiform, pilose in distal half with around Þve long setae up to 0.3 mm. In lateral view, eyespot yellow, elliptical, and situated a short distance behind, and at same height as antennal fossae; a faded yellowish circular spot right above eyespot, Ͻ0.1 mm in diameter, situated close to vertex. Mandibles pitch black distally and grading to chestnut brown at base, robust with a moderate basal hump; left mandible with short distal blade and three marginal teeth, Þrst two prominent, third less prominent; right mandible with distal blade in distal half, and two tri- angular teeth in basal half; dentition as in Fig. 4B; in lateral view, mandibles angling upward Ϸ30Њ from plane of head. Antennae with 13Ð16 segments, rarely 13, the proximal three segments ferruginous orange and grading to yellow at apical segment; Þrst segment longest; third antennal segment subclavate with long setae on distal end; relative length formula usually 2Ͻ3Ͼ4ϭ5. Antennal carinae ferruginous orange, markedly protruding. Postmentum orange-yellow an- teriorly grading to pale yellow posteriorly, bearing setae laterally. Pronotum wide and squarely incised at anterior margin to form an angle Ϸ130Њ; anterior cor- Fig. 2. Scanning electron micrographs of I. inamurae winged imago. Dorsal (A) and oblique (B) views of head and ners sharply rounded; posterior margin with slight pronotum; mandibles (C); right forewing (D). 1 ϩ 2, the concavity; posterior corners broadly rounded. All fused Þrst and second marginal teeth; 3, the third marginal large soldiers with wing pads on meso- and metano- tooth. Scale bars ϭ 500 ␮m. tum. Femora yellow-white, noticeably inßated, slightly longer than tibia. The enlarged third antennal segment, swollen femur, and incised pronotum are the sides parallel with Ͼ50 lateral setae of variable length characters separating the genus Incisitermes from the to 0.2 mm, anterior corners nearly square at mandib- genus Kalotermes. ular articulations, posterior margin evenly rounded, Small Soldier (Figs. 3C and 4C and D; Table 2). median posterior margin slightly concave. Epicranial Head capsule orange yellow in dorsal view, anten- suture faint. Surface of head capsule when viewed nal carinae and frontal carinae protruding and darker, ventrally, pale yellow except for reddish anterior cor- ferruginous orange. Mandibles pitch black; base 762 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5

lighter, dark chestnut brown. Thoracic dorsum or- ange-yellow or yellow; abdominal dorsum lighter, yel- low or pale yellow. Tibiae yellow; femora and sternum yellow-white. Yellow postmentum contrasting with yellow-white genae. In general, small soldier compar- atively less scleritized than large morph. Almost every structure is paler than those of the large soldier by one scale in SandsÕ color scheme (Sands 1965). In dorsal view, head capsule subsquare, shorter than that of large soldier; sides slightly convex with Ͼ50 lateral setae of variable length to 0.2 mm, anterior corners nearly square, posterior corner and posterior margin usually covered by pronotum. Epicranial su- ture faint. Clypeus a narrow, white, trapezoid. Labrum orange yellow, linguiform, pilose in distal half with more than ten long setae over 0.2 mm and less than 5 up to 0.3 mm. Mandibles, much more slender than that of large soldier, with slight basal hump; dentition as in Fig. 4D. Pronotum wide and incised at anterior mar- gin; anterior corners sharply rounded; posterior mar- gin with slight concavity. All small soldiers with wing pads on meso- and metanotum. In lateral view, head capsule, both dorsal and ven- tral discs covered with setae of variable length to 0.2 mm of same density. Frons sloping smoothly from vertex, surface smooth, faintly concave, with Þve to eight long setae at frontal carinae. Mandible angled upward Ϸ30Њ from plane of head. Antennae with 14Ð17 segments, rarely 17, the proximal three seg- ments darker, ferruginous orange, third segment sub- clavate; antennal formula 2Ͻ3Ͼ4ϭ5. Eyespot yellow, elliptical, and situated a short distance behind anten- nal fossae; a subcircular patch Ͻ0.1 mm with the same color with eyespot situated above eyespot, close to vertex. Femora inßated, as long as tibia. Incipient Soldier (Figs. 3D and 4E and F; Table 2). Head capsule pale yellow or yellow white with orange yellow antennal carinae and frontal carinae. Mandi- bles dark chestnut brown in distal half; base lighter, ferruginous orange. Thoracic dorsum yellow-white and semitransparent; abdominal dorsum white. Tibiae, femora and sternum white. Postmentum yellow- white. In general, incipient soldiers are much smaller than either small or large soldiers in size, and only mandibles and anterior part of head are strongly scle- rotized. In dorsal view, head capsule slightly rounded or subsquare; sides slightly convex with Ͻ50 lateral setae of variable length to 0.2 mm, anterior corners curved inward; posterior corner and posterior margin rounded. Epicranial suture obsolescent. Clypeus a narrow, white, trapezoid. Labrum orange yellow, lin- guiform, pilose in distal half with more than 10 setae Ͼ0.2 mm pointing upward and forward. Mandibles, more slender than that of small soldier, with faintly basal hump, dentition as in Fig. 4F. Pronotum wide and Fig. 3. Scanning electron micrographs of dorsal view of smoothly incised at anterior margin; anterior corners I. inamurae soldiers. Large soldier (A and B), small soldier sharply rounded; posterior margin with slight concav- ϭ (C), and incipient soldier (D). Scale bars 1 mm. ity. Some incipient soldiers with wing pads on meso- and metanotum. In lateral view, head capsule, both dorsal and ven- tral disc, covered with scattered setae. Frons sloping September 2009 LIETAL.: REDESCRIPTION OF I. inamurae 763

Fig. 4. Scanning electron micrographs of I. inamurae soldiers in lateral view and their mandibles in dorsal view. Large soldier (A and B), small soldier (C and D), and incipient soldier (E and F). Right antennae removed for clarity. Scale bars ϭ 500 ␮m. smoothly from vertex, slightly concave, with less than Comparison. The body size and color of I. inamurae Þve long setae up to 0.2 mm at frontal carinae. Man- soldiers are similar to those of I. immigrans, and both dible angled upward Ϸ 30Њ from plane of head. An- are smaller and much lighter than I. minor. The prono- tennae with 11Ð14 segments, usually 11 or 12, the tum of I. inamurae is wider than the head and the proximal three segments darker, orange yellow, third anterior margin is smoothly incised, whereas the segment slightly clavate; antennal formula 2Ͻ3Ͼ4ϭ5. pronotum of I. immigrans and I. minor are as wide as Eyespot pale yellow, elliptic; in lateral view, a subcir- head and the anterior margin is sharply incised. The cular spot with the same color of eyespot situated meso- and metathorax of I. inamurae are also wider above eyespot close to vertex. Femora inßated, as long than the head giving I. inamurae a noticeably wider as tibia. thorax compared with most Incisitermes spp. The third 764 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 102, no. 5 from four colonies; large 16 ؍ from three colonies; small soldiers, n 5 ؍ Table 2. Measurements of I. inamurae incipient soldiers, n from four colonies 16 ؍ soldiers, n

Large soldier Small soldier Incipient soldier Measurement (mm) Range Mean Ϯ SD Range Mean Ϯ SD Range Mean Ϯ SD No. antennal segments 13Ð16 14.6 Ϯ 1.0 14Ð17 15.2 Ϯ 0.9 11Ð14 12.4 Ϯ 1.3 Third antennal segment length 0.16Ð0.24 0.20 Ϯ 0.02 0.14Ð0.22 0.18 Ϯ 0.02 0.14Ð0.19 0.17 Ϯ 0.02 Third antennal segment width, max 0.15Ð0.18 0.17 Ϯ 0.01 0.13Ð0.16 0.15 Ϯ 0.01 0.12Ð0.15 0.13 Ϯ 0.01 Third antennal segment width, min. 0.08Ð0.10 0.09 Ϯ 0.01 0.07Ð0.09 0.08 Ϯ 0.01 0.07Ð0.08 0.08 Ϯ 0.00 Head length to tip of mandibles 3.85Ð4.90 4.48 Ϯ 0.28 3.15Ð3.95 3.65 Ϯ 0.18 2.70Ð3.35 3.13 Ϯ 0.25 Head length to medium base of clypeus 2.65Ð3.45 3.15 Ϯ 0.22 2.10Ð2.70 2.47 Ϯ 0.15 1.70Ð2.20 2.08 Ϯ 0.21 Head width, max 2.05Ð2.50 2.20 Ϯ 0.12 1.80Ð1.95 1.88 Ϯ 0.04 1.50Ð1.70 1.63 Ϯ 0.08 Head ht, without postmentum 1.50Ð1.75 1.63 Ϯ 0.09 1.20Ð1.40 1.32 Ϯ 0.07 0.95Ð1.25 1.11 Ϯ 0.11 Labrum, max width 0.53Ð0.65 0.58 Ϯ 0.03 0.50Ð0.55 0.52 Ϯ 0.02 0.43Ð0.48 0.45 Ϯ 0.02 Clypeus width, max 0.60Ð0.85 0.74 Ϯ 0.06 0.40Ð0.73 0.64 Ϯ 0.07 0.53Ð0.60 0.55 Ϯ 0.03 Left mandible lengtha 1.85Ð2.10 1.99 Ϯ 0.07 1.65Ð1.88 1.80 Ϯ 0.06 1.55Ð1.70 1.63 Ϯ 0.06 Postmentum, median length 1.83Ð2.50 2.28 Ϯ 0.18 1.53Ð1.88 1.78 Ϯ 0.10 1.15Ð1.55 1.44 Ϯ 0.16 Postmentum, max width 0.75Ð0.90 0.82 Ϯ 0.05 0.70Ð0.80 0.73 Ϯ 0.03 0.55Ð0.65 0.62 Ϯ 0.04 Postmentum, min. width 0.35Ð0.53 0.44 Ϯ 0.05 0.33Ð0.45 0.38 Ϯ 0.04 0.25Ð0.33 0.29 Ϯ 0.03 Pronotum, median length 1.05Ð1.45 1.27 Ϯ 0.10 0.90Ð1.10 1.02 Ϯ 0.07 0.75Ð0.80 0.78 Ϯ 0.03 Pronotum, max width 2.35Ð2.85 2.67 Ϯ 0.13 2.10Ð2.35 2.25 Ϯ 0.09 1.60Ð2.00 1.85 Ϯ 0.16 Pronotum, max length 1.65Ð2.10 1.89 Ϯ 0.12 1.35Ð1.70 1.54 Ϯ 0.10 1.05Ð1.25 1.19 Ϯ 0.08 Hind tibia length 1.28Ð1.53 1.42 Ϯ 0.08 1.10Ð1.28 1.20 Ϯ 0.07 0.95Ð1.08 0.99 Ϯ 0.05 Hind femur length 1.33Ð2.25 1.53 Ϯ 0.21 1.03Ð1.33 1.20 Ϯ 0.07 0.90Ð1.03 1.00 Ϯ 0.05 Hind femur width, max 0.70Ð1.40 0.90 Ϯ 0.16 0.53Ð0.70 0.62 Ϯ 0.04 0.43Ð0.55 0.48 Ϯ 0.05 Total length 9.42Ð12.10 10.68 Ϯ 0.71 7.79Ð10.23 8.85 Ϯ 0.59 6.72Ð7.35 7.15 Ϯ 0.26

a Tip to most distant visible point of ventral condyle. antennal segment of I. inamurae is shorter than the 120.85Њ E; 19-X-2008; 2 incipient soldiers (TW789). fourth and Þfth combined, but in I. immigrans and I. Eluanbi Park: 21.90Њ N, 120.85Њ E; 19-X-2008; 4 large minor, the third antennal segments are longer than soldiers, 1 for SEM, four small soldiers, 2 for SEM fourth and Þfth combined. The third antennal segment (TW794). Longkeng Ecological Reserve Area: 21.90Њ of I. immigrans and I. minor is more clavate than that N, 120.86Њ E; 19-X-2008; 2 incipient soldiers, 1 for SEM of I. inamurae. (TW798). Based on the description by Light (1921), I. mcgre- Material Examined. Incisitermes immigrans, soldiers gori collected at Luzon Island, the Philippines, resem- and alates: USA. Hawaii, Oahu. Kaena Point State Park: bles I. inamurae, especially the mandible dentition, 21.55ЊN, 158.23ЊW, 6-XII-96, J. Chase and J. Mangold pilosity of head, the wing pads on meso- and metano- (HI24 and HI26); Kaneana Cave area: 21.51ЊN, tum, and the uncommonly wide pronotum. Due to 158.23ЊW, 6-XII-96, J. Chase and J. Mangold (HI34). these two species being close in geographic distribu- Incisitermes minor, soldiers and alates: USA. Arizona, tion and having similar soldier morphology, we pro- Hwy 82 riparian: 31.51ЊN, 110.80ЊW, 3-XI-05, J. Chase, pose that I. mcgregori may be a junior synonym of I. J. Mangold, and R.H. Scheffrahn (US605 and US606). inamurae. However, collection of I. mcgregori winged California, Santa Cruz Co., Scotts Valley: 37.04ЊN, imago and further comparisons are necessary to con- 122.02ЊW, 15-X-03, M. Folkins (US552). All I. immi- Þrm their systematic position. grans and I. minor used for comparison are deposited Based on the description by Light (1930), the I. in the University of Florida termite collection, Fort taylori collected at Mindanao Island, the Philippines, Lauderdale Research and Education Center, Florida is much smaller than the four Incisitermes spp. dis- (FLREC). cussed above. The body length and head width of I. Type Material. All known type specimens collected taylori are only Ͻ5.00 mm and Ϸ1.17 mm, respectively, by Tokie Inamura at Kuraru (now, Heng-Chun Re- compared with Ͼ7.0 mm and Ͼ1.67 mm for the others. search Center, Taiwan Forestry Research Institute), Material Measured. TAIWAN, Pingtung Co., Heng- Pingtung Co., Taiwan were initially deposited at In- chung Township: All samples collected by N. Kanzaki, stitute of Science, Government of Formosa (Oshima Y.-C. Lan, H.-F. Li, and J.-F. Tsai. Longluantan: 21.98Њ 1913). The Institute of Science was later divided into N, 120.74Њ E; 16-X-2008; l alate (TW736). Same local- the Agriculture Research Institute, Forestry Research ity: 4 large soldiers, four small soldiers, one for SEM Institute, Industry Research Institute, and Tropical (TW740). Wanlitong: 22.00Њ N, 120.70Њ E; 16-X-2008; 5 Medicine Research Institute in 1939. No type speci- alates, one designated as neotype and one for SEM, men was found by H.-F.L. in 2008 from the four large soldiers, four small soldiers (TW742). Shed- Collection of Taiwan Forestry Research Institute, Tai- ing Nature Park: 21.95Њ N, 120.82Њ E; 17-X-2008; 4 pei (TFRI) and Taiwan Agriculture Research Insti- alates, 2 for SEM (TW754). Same locality: 1 large tute, Wufeng, Taichung, and other four major insec- soldier, 1 for SEM (TW755). Xiangjiaowan Ecological tariums in Taiwan under National Museum of Natural Reserve Area: 21.93Њ N, 120.83Њ E; 19-X-2008; 1 incip- Science, Taichung (NMNS), National Taiwan Univer- ient soldier, 1 for SEM (TW782). Same locality: 4 large sity, Entomology Department, Taipei (NTU), Na- soldiers, 4 small soldiers (D-1). Longzaipu: 21.92Њ N, tional Chung-Hsing University, Entomology Depart- September 2009 LIETAL.: REDESCRIPTION OF I. inamurae 765 ment, Taichung, and National Pingtung University of Light, S. F. 1921. Notes on Philippine termites, II. Philipp. J. Science and Technology, Plant Protection Depart- Sci 19: 23Ð63. ment, Neipu, Pingtung. Hence, we believe the type Light, S. F. 1930. Notes on Philippine termites, IV. Philipp. specimens have been misplaced, mislabeled, or lost J. Sci 42: 13Ð58. and herein designate neotypes. NEOTYPE: male alate Mori, H. 1976. Establishment of the American common dry-wood termite Incisitermes minor (Hagen) in Tokyo. collected at TAIWAN, Pingtung Co., Wanlitong: Њ Њ Shiroari 27: 45Ð47. 22.00 N, 120.70 E; 16-X-2008 (TW742) will be de- Nation, J. A. 1983. A new method using hexamethyldisi- posited in NMNS. NEOPARATYPES: alates, small sol- lazane for the preparation of soft insect tissue for scan- diers and large soldiers collected with neotype from ning electron microscopy. Stain Technol. 55: 347Ð352. the same colony will be deposited in NMNS, NTU, Ohkuma, M., H. Yuzawa, W. Amornsak, Y. Sornnuwat, Y. TFRI, and FLREC. Takematsu, A. Yamada, C. Vongkaluang, O. Sarnthoy, N. Kirtibutr, N. Noparatnaraporn, T. Kudo, and T. Inoue. 2004. Molecular phylogeny of Asian termites (Isoptera) Acknowledgments of the families Termitidae and Rhinotermitidae based on mitochondrial COII sequences. Mol. Phylogenet. Evol. We thank Rudolf H. Scheffrahn (University of Florida) for 31: 701Ð710. kindly offering specimens of I. immigrans and I. minor for Oshima, M. 1912. The taxonomy and distribution of ter- comparison study and reviewing this article. We also thank mites in Taiwan, pp. 54Ð94. In M. Oshima [ed.], The third Yen-Chiu Lan (Leader University), Jing-Fu Tsai (National ofÞcial report on termites. Taiwan So៮tokufu, Taihoku, Chung-Hsing University), and Chun-Chun Chang (Kenting Japan. National Park Headquater) for assistance in termite collec- Oshima, M. 1913. Notes on the termites of Japan with de- tion and Angelica M. Moncada (University of Florida) for scription of one new species. Philipp. J. Sci. 8: 271Ð280. sorting samples. We thank Paul Skelley (Florida Department Roisin, Y., and J. M. Pasteels. 1991. Polymorphism in the of Agriculture and Consumer Services, Division of Plant giant cocoa termites, Neotermes papua (Desneux). In- Industry) for assistance with scanning electron microscopy. sectes Soc. 38: 263Ð272. This study was supported in part by USDAÐARS grant 58- Roonwal, M. L. 1969. Measurements of termites (Isoptera) 6435-2-276. for taxonomic purposes. J. Zool. Soc. India 21: 9Ð66. Roonwal, M. L., and O. B. Chhotani. 1989. The fauna of India and adjacent countries, Isoptera (termites) vol. 1. References Cited Doon Phototype Printers, Dehradun, India. Sands, W. A. 1965. A revision of the termite subfamily Na- Banks, N., and T. E. Snyder. 1920. A revision of the Nearctic sutitermitinae (Isoptera, Termitidae) from the Ethiopian termites with notes on biology and geographic distribu- Region. Bull. Br. Mus. Nat. Hist. Entomol. Suppl. 4: 1Ð172. tion. Bull. U.S. Nat. Hist. Mus. Wash. 108: 1Ð228. Scheffrahn, R. H., J. Krecek, and N.-Y. Su. 2000. Redescrip- Constantino, R. 1998. Catalog of the living termites of the tions of the dampwood termites Neotermes jouteli and N. New World (Insecta: Isoptera). Arq. Zool. (Sa˜o Paulo) luykxi (Isoptera: Kalotermitidae) from Florida, Cuba, Ba- 35: 135Ð231. hamas, and Turks and Caicos Islands. Ann. Entomol. Soc. Gay, F. J. 1975. An Australian species of Incisitermes Krishna Am. 93: 785Ð794. (Isoptera: Kalotermitidae). J. Aust. Entomol. Soc. 14: 395Ð Snyder, T. E. 1949. Catalog of the termites (Isoptera) of the 398. world. Smithson. Misc. Coll. 112: 1Ð490. Huang, F., and X. He. 2000. Incisitermes minor, pp. 182Ð183. Takematsu, Y. 1997. A new record of Incisitermes immigrans In F. Huang, S. Zhu, Z. Ping, X. He, G. Li, and D. Gao from Japan (Isoptera, Kalotermetidae). Jpn. J. Entomol. [eds.], Fauna Sinica, Insecta vol. 17: Isoptera. Science 65: 634. Press, Beijing, China. Watson, J.A.L. 1974. The development of soldiers in incip- Indrayani, Y., T. Yoshimura, Y. Fujii, Y. Yanase, Y. Okahisa, ient colonies of Mastotermes darwiniensis Froggatt and Y. Imamura. 2004. Survey on the infestation of (Isoptera). Insectes Soc. 21: 181Ð190. houses by Incisitermes minor (Hagen) in Kansai and Yasuda, I., Y. Nakasone, K. Kinjo, and S. Yaga. 2000. Mor- Hokuriku areas. Jpn. J. Environ. Entomol. Zool. 15: 261Ð phology and distribution of termites in Ryuku Islands and 268. North and South Daito Islands. Jpn. J. Entomol. 3: 139Ð Krishna, K. 1961. A generic revision and phylogenetic study 156. of the family Kalotermitidae (Isoptera). Bull. Am. Mus. Nat. Hist. 122: 303Ð408. Received 4 February 2009; accepted 9 June 2009.