Annals of the Entomological Society of America Advance Access published November 13, 2015

Annals of the Entomological Society of America, 2015, 1–10 doi: 10.1093/aesa/sav106 Systematics Research article

Redescription of the Soil-Feeding Sinocapritermes mushae (Isoptera: : Termitinae): The First Step of Revision

C.-I. Chiu, Man-Miao Yang, and Hou-Feng Li1

Department of Entomology, National Chung Hsing University, 250 Guoguang Rd., Taichung 40227, Taiwan ([email protected]; [email protected]; [email protected]) and 1Corresponding author, e-mail: [email protected]

Received 10 August 2015; Accepted 14 October 2015

Abstract The soil-feeding termite, Sinocapritermes mushae (Oshima & Maki), was described briefly in Japanese in 1919 and its type locality is Taiwan. S. mushae is the first found species of the genus, and all the other 15 congeners were described from China, and many of them were compared with S. mushae for species differentiation. However, the inconsistent interpretation of the original description of S. mushae soldier morphology occurred in Chinese literature. The winged imago of S. mushae was later described based on a single specimen collected in China, which is insufficient to represent the morphological variation and caused further confusion. To solve these problems, we re-describe and describe the soldier, winged imago, and worker enteric valve of S. mushae based on multiple colony samples collected from the type locality, Taiwan. Partial sequences of mitochondrial 16S gene confirmed that S. mushae is the only Sinocapritermes species found in Taiwan, but Taiwanese popu- lations present high morphological variation. With the intraspecific variation of S. mushae, we demonstrated that most qualitative characters used for species differentiation among Chinese species are not valid. In addi- tion, the morphometric data showed that the five species, Sinocapritermes sinensis Ping & Xu, Sinocapritermes songtaoensis He, Sinocapritermes tianmuensis Gao, Sinocapritermes yuannensis Ping & Xu, and Sinocapritermes xiai Gao & Lam, are not significantly different from S. mushae, and hence they are proposed as junior synonyms. This study indicates that the 15 exclusive Sinocapritermes spp. found in China need to be more thoroughly investigated, and a further revision of Sinocapritermes is necessary.

Key words: termite, , morphology, China, Taiwan

Sinocapritermes Ping and Xu 1986 is a medium genus of soil-feeding between 1963 and 1993. All 16 species occur on mainland China, termite, comprising 16 species from four countries in the Oriental and S. mushae is the only species found outside of China. The inac- Region (Krishna et al. 2013), including China (Krishna et al. 2013), cessible samples and language barrier likely hindered the study of Japan (Takematsu 1994), Taiwan (Oshima and Maki 1919), and Sinocapritermes taxonomy. S. mushae was originally named as Vietnam (Quang et al. 2012). Sinocapritermes belongs to the Procapritermes mushae by Oshima and Maki and reassigned to -Capritermes branch of Termitinae whose soldiers have sym- Sinocapritermes by Ping and Xu in 1986. Because S. mushae was the metric or asymmetrical snapping mandibles (Krishna 1968). This first described species of the genus, many later found species were mandible type allows them to perform a powerful strike on their en- compared with S. mushae, including Sinocapritermes parvulus Yu emies, ants, by a rapid and ballistic movement (Seid et al. 2008). & Ping 1966, Sinocapritermes vicinus Xia et al. 1983, Sinocapritermes and Procapritermes Kemner 1934 are two morpho- Sinocapritermes yuanensis Ping & Xu 1986, Sinocapritermes fujia- logically and phylogenetically similar genera (Ping and Xu 1986, nensis Ping & Xu 1986, Sinocapritermes tianmuensis Gao 1989, Ohkuma et al. 2004, Inward et al. 2007). The main morphological and Sinocapritermes pratensis Ping & Xu 1993. To differentiate difference between these two genera was the number of tibial spurs S. mushae from the other Chinese species, nine qualitative characters of soldier fore legs (Ping and Xu 1986), two for Sinocapritermes and and three quantitative measurements were used (Yu and Ping 1966, three for Procapritermes (Ping and Xu 1986). Ping and Xu 1986, Gao 1989, Ping and Xu 1993). However, con- The taxonomic status of Sinocapritermes is unique from a histor- flicting interpretation of these diagnostic characters of S. mushae oc- ical perspective. S. mushae was collected from Musha (Wushe, curred. For example, the relative length of 2nd and 3rd antennal Nantou County), Taiwan, and first described in Japanese in 1919, segments of soldier that was used for differentiating S. mushae from and the other 15 congeneric species were described in Chinese S. parvulus (S. mushae: 2nd > 3rd vs. S. parvulus: 2nd ¼ 3rd, Yu and

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Fig. 1. Known distribution of S. mushae in Taiwan. Type locality (star), measured (closed circles), and examined (open circles) materials were marked.

Ping 1966), S. vicinus (S. mushae: 2nd > 3rd vs. S. vicinus: Ping and Xu, 1986; S. tianmuensis Gao, 1989; S. vicinus Xia, Gao, 2nd < 3rd, Xia et al. 1983), and S. pratensis (S. mushae: 2nd < 3rd Pan, and Tang, 1983; Sinocapritermes albipennis Tsai & Chen, vs. S. pratensis: 2nd ¼ 3rd, Ping and Xu 1993), comprised three ver- 1963; and S. mushae (Tsai and Chen 1964, Xia et al. 1983, Ping and sions in the literature (2nd < 3rd: Oshima and Maki 1919, Ping and Xu 1986, Gao 1989). S. mushae was first described based on soldier Xu 1993; 2nd ¼ 3rd: Tsai and Chen 1964; 2nd > 3rd: Yu and Ping and worker castes collected by Maki from Musha, Taiwan (Oshima 1966, Xia et al. 1983). Vertex shape of soldier head capsule and Maki 1919). The winged imago of S. mushae was later de- (Oshima and Maki 1919, Tsai and Chen 1964, Ping and Xu 1986, scribed by Tsai and Chen in 1964 based on a single specimen col- 1993) is another contradictory character of S. mushae found in pre- lected in China. vious Chinese literature. Therefore, a need exists to revise the de- For establishing solid Sinocapritermes taxonomy, a clear de- scription of S. mushae. scription of S. mushae is a crucial step. In the current study, we Among the 16 named Sinocapritermes species, the winged imago redescribed S. mushae based on worker, soldier, and winged imago caste was only mentioned in five descriptions as follows: S. sinensis samples collected from multiple colonies from Taiwan. The Annals of the Entomological Society of America, 2015, Vol. 0, No. 0 3

(NMNS), are all winged imagos collected by using Malaise traps, including 15 individuals from four localities of two counties in Taiwan. Morphological observations were conducted by using a LeicaVR M205 C stereomicroscope or LeicaVR DM750 compound micro- scope (Leica Microsystems, Wetzlar, Germany). Measurements and photographs of soldier and imago were taken by LeicaVR MC170 HD digital camera (Leica Microsystems, Wetzlar, Germany) with the software LAS (LeicaVR Application Suite V4.4.0, Leica Microsystems, Wetzlar, Germany). Photograph of worker enteric valve was taken by a single-lens reflex camera (CanonVR EOS 600D; Canon, Tokyo, Japan). Image stacking is carried out by using soft- ware Picolay (free software by H. Cypionka, http://www.picolay. de). Morphological measurements were adopted from Roonwal (1969). Color description followed the Munsell color system (Munsell Color Company 1975). An image of each character was vi- sually compared with Munsell color plate on the computer monitor to describe the hue, value, and chroma. Samples collected from five locations, Dakeng (TW2178), Shuishe Mountain (TW2190), Huisun Forest Station (TW2184), Mingchi (TW2167), and Sileng (TW2211), were subjected to genetic analysis. Total genomic DNA of worker thorax muscles was extracted by using the Puregene DNA Isolation kit (Gentra Systems, Minnesota, USA) following the manufacturer’s protocol. Primers for mitochondrial 16S rRNA are 16Sar (50-CGCCTGTTTATCAAAAACAT-30) and 16Sbr (50-CCGGTCTGAACTCAGATCACGT-30). The 30-ll PCR mixture

contained 23.5 llofddH2O, 1 ll of dNTP mixture (25 mM), 0.3 llof Taq DNA Polymerase, forward and reverse primers (0.6 ll each, 10 mM), 10 reaction buffer 3 ll, and 1 ll termite DNA template. The thermal cycling for PCR included a pre-cycle denaturation at 95C for 5 min, followed by 35 cycles of denaturation at 95C for 1 min, annealing at 55C for 1 min, extension at 72C for 2 min, and a postcycle extension at 72C for 10 min. Gene sequences were submitted to GenBank (GenBank accession numbers: KT277540–44). For morphological comparison of all named Sinocapritermes species, all characters used for Sinocapritermes species differenti- Fig. 2. Definition of three soldier diagnostic characters, head vertex shape (A), the parallelism of lateral sides of head (B), and the presence/absence of me- ation were analyzed in this study, including nine qualitative char- dian notch on anterior margin of pronotum (C). acters and three quantitative characters. The nine qualitative characters were: 1–8 are soldier characters and 9 is that of available intraspecific variation allows us to reevaluate the reliabil- imago; 1, head vertex shape (Ping and Xu 1986, Gao and Lam ity of previously used diagnostic characters of Sinocapritermes 1990); 2, the presence/absence of median notch on anterior mar- spp. The species with no significantly different morphology from gin of pronotum (Ping and Xu 1986); 3, the parallelism of two S. mushae were proposed as junior synonyms. Partial sequences of lateral sides of head (Ping and Xu 1993); 4, relative lengths of mitochondrial 16S gene of selected Taiwanese samples were also left and right mandible (Ping and Xu 1986); 5, relative length of provided for future molecular identification of the inaccessible 2nd and 3rd antennal segments (Yu and Ping 1966, Xia et al. Chinese samples. 1983, Ping and Xu 1993); 6, the ratio of medial and lateral length of labrum (Yu and Ping 1966); 7, the ratio of head median sutureandheadlengthtomandiblebase(Ping and Xu 1986); 8, the relative values of head length to mandible base and mandible Materials and Methods length (Ping and Xu 1993); and 9, relative values of eye–ocellus Redescriptions are all based on samples collected from Taiwan distance excluding sclerite and ocellus diameter (Xia et al. 1983). (Fig. 1) by two methods, hand-picking and Malaise trapping. For The definitions of qualitative character 1, 2, and 3 are presented hand-picking, termite samples were collected from the interface in Fig. 2. The three quantitative soldier characters were head between soil and wood or in termite’s subterranean gallery sys- width, head length to mandible base at dorsal condyle, and mini- tem (Chiu et al. 2015). Hand-picking samples were immediately mum width of postmentum. preserved in 85% ethanol. In total, 25 colony samples collected from eight localities of five counties in Taiwan, containing worker, soldier, and winged imago castes, were deposited at the Sinocapritermes mushae (Oshima and Maki) National Chung Hsing University Termite Collection, Taichung, Taiwan (NCHU). The other examined samples preserved in the Procapritermes mushae Oshima and Maki 1919: 313–316, figs. 1 National Museum of Natural Science, Taichung, Taiwan and 2 [soldier, worker described and figured]. Tsai and Chen 4 Annals of the Entomological Society of America, 2015, Vol. 0, No. 0

Fig. 3. S. mushae soldier. Semi-lateral view of mandibles (A), and whole body (B). Ventral view of whole body (C). Dorsal view of pronotum, mesonotum, metano- tum, and abdomen (D). Lateral view of head capsule (E).

1964: 86–87, fig. 37 [imago, soldier described and figured]. Sinocapritermes mushae (Oshima and Maki) Ping and Xu 1986:2, Yu and Ping 1966: 130 [soldier compared with S. parvulus]. 7, 12, 16, 18, [soldier compared with S. yunnanensis and Xia et al. 1983: 161–162, [soldier and winged imago com- S. fujianensis]. Gao 1989: 2, 4–5, [key; soldier compared with pared with S. xiai]. S. tianmuensis]. Huang et al. 1989: 496, 505–507, fig. 377 Annals of the Entomological Society of America, 2015, Vol. 0, No. 0 5

Table 1. Measurements of soldier of S. mushae, n ¼ 36 from four Table 2. Measurements of winged imagoes of S. mushae, n ¼ 30 colonies from three colonies

Measurement (mm) Range Mean 6 SD Measurement (mm) Range Mean 6 SD

No. antennal segments 14 14 No. antennal segments 14–15 14.97 6 0.18 Length of left mandible 1.41–1.84 1.64 6 0.11 Head length to mandible base 0.61–0.84 0.74 6 0.06 Length of right mandible 1.52–1.95 1.72 6 0.11 Head length to labrum tip 0.85–1.37 1.12 6 0.14 Head length to mandible tip 2.97–3.69 3.28 6 0.19 Head width, max at eyes 1.07–1.19 1.13 6 0.03 Head length to mandible base 1.52–1.91 1.69 6 0.11 Head width without eyes 0.82–0.97 0.93 6 0.04 Head length to labrum tip 1.91–2.51 2.14 6 0.15 Eye max diam without sclerite 0.24–0.30 0.27 6 0.02 Head width, max 1.06–1.36 1.18 6 0.08 Distance from eye to lower mar- 0.04–0.09 0.07 6 0.01 Head height with postmentum 0.81–1.10 0.95 6 0.08 gin of head, min Postmentum, median length 0.67–1.11 0.85 6 0.08 Ocellus diam, max 0.08–0.13 0.10 6 0.02 Postmentum, max width 0.33–0.45 0.38 6 0.04 Eye-ocellus distance excluding 0.04–0.11 0.07 6 0.02 Postmentum, min width 0.19–0.28 0.23 6 0.02 sclerite Labrum, median length 0.20–0.47 0.35 6 0.07 Forewing length from suture 9.93–12.22 11.11 6 0.57 Labrum, max length 0.30–0.65 0.48 6 0.08 Forewing, max width 2.49–3.20 2.77 6 0.18 Labrum, max width 0.25–0.37 0.29 6 0.03 Pronotum, max length 0.41–0.56 0.49 6 0.04 Pronotum, max length 0.21–0.36 0.30 6 0.03 Pronotum, max width 0.80–0.97 0.89 6 0.05 Pronotum, max width 0.59–0.77 0.66 6 0.05 Total length with wings 11.81–13.81 12.85 6 0.59 Total length 4.17–5.50 4.66 6 0.34 Total length without wings 4.95–6.46 5.84 6 0.36 Hind tibia, max length 0.93–1.22 1.06 6 0.07 Hind tibia, max length 0.83–1.26 1.12 6 0.10

[soldier described and figured]. Gao and Lam 1990: 332, eyes black (2.5Y 2/0); costal margin of wings and radial sector vein [key]. Ping and Xu 1993: 439, [soldier compared with S. pra- brown (5YR 3/2). tensis]. Takematsu 1994: 719–722, [soldier, worker described Mandibular dentition is in Fig. 4A. Wing membrane brown (5YR and figured]. Chung and Chen 1994: 202, [key]. Huang and 3/2), covered with thousands of setae, rounded at the apex; costal Han 1997: 111, fig. 8 [soldier described and figured]. Huang margin sclerotized. Forewing slightly longer and broader than hindw- and Li 1999: 312, fig. 10.55 [soldier described and figured]. ing. Forewing (Fig. 4B) median vein with 50–68 hairs, branching 1–3 Huang et al. 2000: 723–725, fig. 437 [soldier described and times near the apex, cubitus with 7–9 branches. Hindwing (Fig. 4C) figured]. Tsai 2003: 104, 124–128, fig. 32 [soldier, worker cubitus with 6–9 branches. Head hairy, broadly oval (Fig. 4D). described and figured]. Fontanelle barely visible (18–31 lm). Area surrounding the fontanelle concave. Postclypeus slightly swollen, convex at the midline and a peak can be observed at the middle of anterior margin; anteclypeus Soldier (Fig. 3; Table 1) nearly as broad as postclypeus, slightly longer than postclypeus, semi- Mandibles dark (2.5Y 2/10), asymmetrical. Left mandible bent in transparent; labrum slightly longer than its width, broadest in the lower half part (Fig. 3A). Head capsule yellow in dorsal view (2.5Y middle, anterior margin unsclerotized (Fig. 4D). Compound eyes cir- 7/8), subrectangular, posterior margin rounded (Fig. 3B). Median cular and protruding; ocellus oval and white. The distance between suture yellow (10YR 7/8), depressed and not reaching the fontanelle, eye and ocellus is slightly longer or equal to ocellus diameter. the ratio of median suture/head length to mandible base is 0.22– Pronotum hairy on lateral and posterior margin, lateral side broadly 0.51. Fontanelle small, opening toward front. Clypeus trapezoidal. round, anterior margin raised, posterior corner rounded, posterior Labrum yellow (2.5Y 7/8), anterior margin sharply bifurcated. The margin downturned. Mesonotum and metanotum clothed with hairs ratio of medial/lateral length of labrum is 0.59–0.85. Antennae with (20–85 lm), posterior margin strongly bilobed (Fig. 4E). Antennae 14 segments (n ¼ 36), yellow (2.5Y 7/8); first segment longest; forth 14–15 segmented (14 segments: 3.3%; 15 segments: 96.7%; n ¼ 30), segment shortest; relative length formula 3 > 4 < 5 < 6 ¼ 7; segment first segment longest, second segment shorter than first segment, third length gradually decrease after ninth segment. In ventral view, post- segment shortest, fourth segment longer or shorter than fifth segment mentum sclerotized and shaped as in Fig. 3C. In lateral view, head (longer: 66.7%; shorter: 33.3%; n ¼ 30), the segment length gradually vertex shaped as in Fig. 3E. increasing after fifth segment (Fig. 4G). Pronotum, metanotum, mesonotum, and abdomen light yellow In ventral view, postmentum with apical half with a square- (10YR 8/8), with short hairs (10–20 lm) and long hairs (60– shaped, sclerotized plate adjacent to anterior corner, other portions 100 lm) (Fig. 3D). Long hairs generally locate near poster margin. unsclerotized and semitransparent; posterior half sclerotized, haired, Pronotum saddle-shaped. Mesonotum as wide as pronotum. and yellow (7.5YR 5/8; Fig. 4F). Gena and palpi yellow (7.5YR 7/8). Metanotum wider than pronotum and mesonotum. Median notch Abdominal sternites yellow in midline (7.5YR 7/8), brown in two lat- on anterior margin of pronotum not always present. Abdomen with- eral sides (7.5YR 5/4). Coxae, trochanters, femora, tibiae, and tarsi out soil content. Coxae, trochanters, femora, tibiae, and tarsi pale pale yellow (10YR 8/8); claws sclerotized. Tibial spur formula 2:2:2. yellow (10YR 8/8); claws sclerotized. Tibial spur formula 2:2:2.

Worker Enteric Valve (Fig. 5) Winged Imago (Fig. 4; Table 2) Enteric valve (P2; Fig. 5A) completely embed in third segment of In dorsal view, head capsule brown (10R 3/3) at central area and hindgut (P3; Fig. 5A), with 7 cushions armed with sclerotized spines dark brown (10R 1/3) at lateral area; pronotum with frontal half (Fig. 5B). Spines concentrated in middle and tip of each cushion, yellowish brown (5YR 5/8) and posterior half brown (5YR 3/2); an- forming two clusters of spines (Fig. 5B), 2–4 spines on tip, and 4–7 tenna, mesonotum, and metanotum yellow (7.5YR 6/8); compound spines in middle. 6 Annals of the Entomological Society of America, 2015, Vol. 0, No. 0

Fig. 4. S. mushae winged imago. Dorsal view of mandibles (A), right forewing (B), and right hindwing (C). Frontal view of head capsule (D). Dorsal view of whole body (E). Ventral view of postmentum (F). Antenna (G).

Comparison species. In addition, no specific morphological difference was ob- The partial mitochondrial 16S sequences show that the pairwise served from all the examined Taiwanese Sinocapritermes sam- genetic distance among the five selected samples ranged from 0 ples. In this study, both morphological and molecular data to 0.98% (GenBank accession numbers: KT277540–44). The support that S. mushae is the only Sinocapritermes species found limited gene variation supported that these samples are the same in Taiwan. Annals of the Entomological Society of America, 2015, Vol. 0, No. 0 7

Fig. 5. Digestive tube of S. mushae worker. Left view of midgut (M), first segment of hindgut (P1), enteric valve (P2), and third segment of hindgut (P3) (A). Worker enteric valve cushions (left) with line drawing of spine patterns on a cushion (right) (B).

We found that intraspecific morphological variations among Head width and head length to mandible base, which represent the examined samples are larger than those described in previous the body size of termite, show that S. guangxiensis, S. xiushanensis, studies (Oshima and Maki 1919, Tsai and Chen 1964, Yu and S. magnus, S. planifrons, and S. albipennis are larger than S. mushae Ping 1966, Xia et al. 1983, Ping and Xu 1986, Gao 1989, Gao and (0–5% overlaps; Fig. 6), and S. parvulus and S. fujianensis are Lam 1990, Ping and Xu 1993, Takematsu 1994). Nine qualitative smaller than S. mushae (no overlap; Fig. 6). Minimum widths of characters used for species identification are not consistently ob- postmentum of S. parvulus, S. pratensis, S. sinicus, and S. vicinus are served among intracolony individuals of S. mushae (Table 3). smaller than that of S. mushae (no overlap; Fig. 6). The ranges of Among the 22 states of the nine qualitative characters, all head width, head length to mandible base, and minimum width were found in S. mushae (Table 3); hence, we could not differenti- of postmentum of S. yuanensis, S. tianmuensis, S. songtaoensis, ate S. mushae from the 14 species, including S. parvulus, S. fujia- S. sinensis, and S. xiai are highly overlapped (50–100%) with that nensis, Sinocapritermes guangxiensis, Sinocapritermes of S. mushae; hence, we are unable to differentiate S. mushae with xiushanensis, Sinocapritermes magnus, Sinocapritermes these five species based on the measurements. Because no qualitative planifrons, Sinocapritermes albipennis, Sinocapritermes yuannen- and quantitative characters could be used to differentiate S. mushae sis, Sinocapritermes pratensis, Sinocapritermes tianmuensis, from S. yuanensis, S. tianmuensis, S. songtaoensis, S. sinensis, and Sinocapritermes songtaoensis, Sinocapritermes sinicus, S. sinensis, S. xiai, we proposed that the five species are junior synonyms of and S. xiai. S. mushae. 8 Annals of the Entomological Society of America, 2015, Vol. 0, No. 0

Table 3. Examination of qualitative characters of S. mushae in four and 10 imagoes (TW2096). Huisun Forest Station: 24.09 N, colonies 121.04 E; 02-II-2012; C.-I. Chiu; 10 soldiers and 10 imagoes (TW2184). Character status Presence of the character/No. examined individual

TW2167 TW2184 TW2096 TW2178 Material Examined 1a 8/10 1/10 3/6 0/10 TAIWAN: Yilan Co., Fushan: 24.76 N, 121.59 E; 6-VI-2005; H.- 1b 2/10 9/10 3/6 10/10 F. Li; 1 soldier (TW502). Same locality: 30-V-2007; 1 soldier 2a 8/10 9/10 1/6 4/10 (TW514). Same locality: 12-VIII-2012; C.-I. Chiu; 15 soldiers 2b 2/10 1/10 5/6 6/10 (TW2189). Same locality: 13-VIII-2012; 25 soldiers (TW2190). 3a 6/10 9/10 6/6 10/10 Mingchi: 24.65 N, 121.47 E; 3-IV-2014; H.-F. Li, C.-I. Chiu and 3b 4/10 1/10 0/6 0/10 W.-R. Liang; 4 workers (TW2167). Taoyuan Co. Sileng: 24.65 N, 4a 1/9 1/10 2/6 0/10 121.43 E; 2-IV-2014; H.-F. Li, C.-I. Chiu and W.-R. Liang; 5 4b 8/9 9/10 4/6 10/10 soldiers (TW2165). Taichung Co., Dakeng: 24.19 N, 120.80 E; 5a 5/10 3/10 3/6 4/10 5b 5/10 2/10 0/6 5/10 27-VII-2011; C.-I. Chiu; 9 soldiers (TW2177). Same locality: 6-VIII- 5c 0/10 5/10 3/6 1/10 2011; 2 workers (TW2178). Same locality: 25-IX-2011; 1 soldier 6a 0/10 0/10 0/6 1/10 (TW2179). Same locality: 10-X-2011; 5 soldiers (TW2180). Same 6b 8/10 1/10 1/6 3/10 locality: 10-X-2011; 1 soldier (TW2181). Same locality: 13-X-2011; 6c 2/10 9/10 5/6 6/10 7 soldiers (TW2182). Same locality: 12-II-2012; 5 soldiers 7a 1/10 0/10 1/6 2/10 (TW2185). Tungmao Mountain: 24.18 N, 120.94 E; 28-I-2013; 7b 4/10 3/10 1/6 5/10 C.-I. Chiu; 3 soldiers (TW2192). Nantou Co., Chunyang: 24.03 N, 7c 5/10 7/10 4/6 3/10 121.16 E; 12-III-200209-IV-2002; C.-S. Lin and W.-T. Yang; 1 8a 7/9 5/10 6/6 9/10 winged imago (NMNS ENT 5237-1025). Same locality: 12-III- 8b 1/9 1/10 0/6 0/10 200209-IV-2002; 1 winged imago (NMNS ENT 5237-1026). 8c 1/9 4/10 0/6 1/10 9a 2/10 10/10 7/10 – Lienhuachih: 23.92 N; 120.89 E; 17-X-200013-XI-2000; C.-S. 9b 8/10 0/10 3/10 – Lin and W.-T. Yang; 1 winged imago (NMNS). Same locality: 11- XII-20007-I-2000; 1 winged imago (NMNS). Same locality: 8- Character definitions: XII-200118-II-2002; 1 winged imago (NMNS ENT 6366-305). 1, head vertex shape (soldier): raised (a), flat (b); Same locality: 6-V-200210-VI-2002; 1 winged imago (NMNS 2, the presence/absence of median notch on anterior margin of pronotum ENT 4686-526). Same locality: 9-XII-20026-I-2003; 1 winged (soldier): presence (a), absence (b); imago (NMNS ENT 6575-469). Same locality: 15-XI-200413- 3, the parallelism of two lateral sides of head (soldier): parallel (a), not par- XII-2004; 1 winged imago (NMNS ENT 6541-998). Same locality: allel (b); 6-VI-20054-VII-2005; 1 winged imago (NMNS ENT 6851-215). 4, the relative lengths of left and right mandible (soldier): left > right (a), left < right (b); Shuishe Mountain: 23.85 N, 120.94 E; 18-III-2011; C.-I. Chiu; 1 5, the relative lengths of 2nd and 3rd antennal segments (soldier): worker (TW2096). Huisun Forest Station: 24.09 N, 121.04 E; 18- 2nd < 3rd (a), 2nd ¼ 3rd (b), 2nd > 3rd (c); XI-2011; C.-I. Chiu; 8 soldiers (TW2183). Same locality: 02-II- 6, medial/lateral length of labrum (soldier): >0.84 (a), 0.76–0.84 (b), 2012; 1 worker (TW2184). Same locality: 3-V-2012; 9 soldiers <0.76 (c); (TW2186). Same locality: 1-VII-2012; 4 soldiers (TW2187). Same 7, length of head median suture/head length to mandible base (soldier): locality: 1-VII-2012; 2 soldiers (TW2188). Same locality: 4-XI- 0.5 (a), 0.4 (b), 0.3 (c); 2012; 1 soldier (TW2191). Same locality: 10-V-2013; 8 soldiers 8, the relative values of head length to mandible base and mandible length (TW2193). Same locality: 1-III-2014; 1 soldier (TW2194). Same lo- (soldier): head length to mandible base < mandible length (a), head length to cality: 22-III-2012; 1 soldier (TW2195). Pingtung Co. Lider: 22.01 mandible base ¼ mandible length (b), head length to mandible base > mandi- N, 120.84 E; M.-L. Jeng and T.-R. Chen; 16-I-201128-I-2011, 2 ble length (c); 9, the relative lengths of eye–ocellus distance excluding sclerite and ocellus winged imagoes (NMNS). Nanren Mountain: 22.08 N, 120.86 E; diameter (winged imago): eye-ocellus distance < ocellus diameter (a), eye- 29-XII-201014-I-2011; M.-L. Jeng and T.-R. Chen; 1 winged ocellus distance ¼ ocellus diameter (b). imago (NMNS). Same locality: 26-I-20118-II-2011, 3 winged imagoes (NMNS). Dahan Mountain: 22.41 N, 120.74 E; 2-IV- Remarks 2013; W.-R. Liang; 10 winged imagoes (TW2219). The winged imagoes collected by using Malaise trap show that its dispersal flight occurred yearlong but mainly in winter and spring. Type Material The winged imagoes collected by hand-picking were found under The depositary institute of S. mushae specimens collected by stone or wood in February (TW2184), March (TW2096), and April Moichiro Maki at Musha, Nantou Co., Taiwan, was not mentioned (TW2167). When stone or wood was removed, the winged imagoes in the original article (Oshima an Maki 1919). The type specimen is were spreading wings and tended to fly, which also indicated that unknown (Krishna et al. 2013). No type specimen was found by C.- their dispersal flight season is in winter and spring. I. Chiu and H.-F. Li from the major insectariums in Taiwan, includ- ing National Taiwan University, Entomology Department, Taipei, Material Measured Collection of Taiwan Forestry Research Institute, Taipei TAIWAN: Yilan Co., Mingchi: 24.65 N, 121.47 E; 3-IV-2014; (TFRI), Taiwan Agriculture Research Institute, Wufeng, Taichung, H.-F. Li, C.-I. Chiu and W.-R. Liang; 10 soldiers and 10 imagoes National Museum of Natural Science, Taichung (NMNS), National (TW2167). Taichung Co., Dakeng: 24.19 N, 120.80 E; 6-VIII- Chung-Hsing University, Entomology Department, Taichung, and 2011; C.-I. Chiu; 10 soldiers (TW2178). Nantou Co., Shuishe National Pingtung University of Science and Technology, Plant Mountain: 23.85 N, 120.94 E; 18-III-2011; C.-I. Chiu; 6 soldiers Protection Department, Neipu, Pingtung. We also contacted the Annals of the Entomological Society of America, 2015, Vol. 0, No. 0 9

Fig. 6. Comparison of the ranges of head width, head length to mandible base, and minimum width of postmentum of the soldier caste among 16 Sinocapritermes species. When the measurement range of S. mushae (gray band) is partially overlapped with that of another species, the overlapping percent- age is labeled. The ranges of S. mushae and other species were obtained from this study and its original descriptions, respectively.

University Museum, The University of Tokyo, Tokyo, Japan, to partially supported by the grant (MOST 102-2313-B-005-037-MY2) from search the type specimens collected by Moichiro Maki, but no type the Ministry of Science and Technology, Taiwan. specimen was found there. Hence, we believe that the type speci- mens have been lost, and herein designate neotypes. NEOTYPE: sol- References Cited dier collected at Taiwan, Nantou Co., Huisun Forest Station: 24.09 N, 121.04 E; 02-II-2012 (TW2184) will be deposited in NMNS. Chiu, C.-I., M.-M. Yang, and H.-F. Li. 2015. Structure and function of subter- NEOPARATYPES: winged imagoes and soldiers collected with neo- ranean gallery systems of soil-feeding Pericapritermes nitobei and type from the same colony will be deposited in NMNS, NCHU, and Sinocapritermes mushae. Insect. Soc. Online Publication. doi: 10.1007/ s00040-015-0416-4. TFRI. Chung, C.-H., and C.-S. Chen. 1994. A review of Taiwanese termites (Insecta, Isoptera) with keys to adults and soldiers. Yushania 11: 193–203. Gao, D.-R. 1989. A new species of Sinocapritermes from Mt. Tinmu, China Acknowledgments (Isoptera: Termitidae). Sci. Technol. Termites 6: 1–5. Gao, D.-R., and P. K. S. Lam. 1990. Notes on the genus Sinocapritermes We are very grateful to Chang-Ti Tang, Yi-Chang Liao, Sheng-Feng Lin, and (Isoptera: Temitidae) from China, with description of a new species. Syst. Bao-Cheng Lai (National Chung Hsing University) for sharing important Entomol. 15: 331–334. comments and review of manuscript, and Chih-Hung Wang (Tokyo Huang, F.-S., and Y.-H. Han. 1997. Isoptera: Rhinotermitidae and University of Agriculture and Technology) for translating Japanese reference. Termitidae, pp. 104–112. In X.-K. Yang (ed.), of the three gorge res- We thank Taiwan Forestry Research Institute and Forestry Bureau, Council ervoir areas of Yangtze river, vol. 1. Chongqing Publishing House, of Agriculture, Taiwan, for issuing the collection permissions. This study was Chongqing, China. 10 Annals of the Entomological Society of America, 2015, Vol. 0, No. 0

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