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A Single Endemic and Three Exotic Species of the Termite Genus Coptotermes (Isoptera, Rhinotermitidae) in the New World

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A Single Endemic and Three Exotic Species of the Termite Genus Coptotermes (Isoptera, Rhinotermitidae) in the New World 73 (2): 333 – 348 20.8.2015 © Senckenberg Gesellschaft für Naturforschung, 2015. A single endemic and three exotic species of the termite genus Coptotermes (Isoptera, Rhinotermitidae) in the New World Rudolf H. Scheffrahn *, 1, Tiago F. Carrijo 2, Jan Křeček 1, Nan-Yao Su 1, Allen L. Szalanski 3, James W. Austin 4, James A. Chase 5 & John R. Mangold 5 1 Fort Lauderdale Research and Education Center, 3205 College Avenue, Davie, Florida 33314, USA; Rudolf H. Scheffrahn * [[email protected]]; Jan Křeček [[email protected]]; Nan-Yao Su [[email protected]] — 2 Museu de Zoologia, Universidade de São Paulo, Av. Nazaré 481, 04263-000, São Paulo, SP, Brasil; Tiago F. Carrijo [[email protected]] — 3 Department of Entomology, 319 Agri. Bldg., University of Arkansas, Fayetteville, AR 72701, USA; Allen L. Szalanski [[email protected]] — 4 BASF Corporation, 26 Davis Drive, P.O. Box 13528, Research Triangle Park, NC 27709, USA; James W. Austin [[email protected]] — 5 Terminix International, 860 Ridge Lake Boulevard, Memphis, Tennessee 38120, USA; James A. Chase [[email protected]]; John R. Mangold [[email protected]] — * Correspond ing author Accepted 06.vii.2015. Published online at www.senckenberg.de/arthropod-systematics on 07.viii.2015. Editor in charge: Julia Goldberg. Abstract The termite genus Coptotermes is both large and widespread but, as a whole, lacks robust diagnostic characters for morphological species identification. This has resulted in many taxonomic synonymies leading to the current pool of extant species a few of which are among the world’s most destructive pests of wood. The New World diversity of Coptotermes is far less complicated, but has not been the subject of critical study. Using a large collection representative of endemic Coptotermes from the region, we compared their external morpho­ logy and their 16S genetic marker sequences to identify the three described American species: C. crassus, C. niger, and C. testaceus. We found no consistent differences among populations from Mexico to Bolivia, and therefore, we reassign the former two species as junior synonyms of C. testaceus. We also compared alates and soldiers of the three non­native species of Coptotermes: C. formosanus, C. gestroi, and C. sjostedti (soldiers only) to C. testaceus and provide diagnostic characters to identify all four current New World species. An updated distribution map is provided. Key words Coptotermes crassus, Coptotermes formosanus, Coptotermes gestroi, Coptotermes niger, Coptotermes sjostedti, Coptermes testaceus, synonymy. 1. Introduction The termite genus Coptotermes Wasmann currently con­ ability (EMERSON 1971) and increased body mass with sists of 67 extant species (KRISHNA et al. 2013) distributed colony age (GRACE et al. 1995) compound the difficulty in Asia, Australia, Africa, and the New World. Because of sorting species morphologically. this genus is specious, widespread, pestiferous, and lacks As a result of drastic and provincial proliferation of robust diagnostic interspecific morphology,Coptotermes the Chinese termite fauna (EGGLETON 1999), 40 Cop­ has undergone more synonymies than any other termite totermes species were described from China between genus (KRISHNA et al. 2013). Intraspecific character vari­ 1985 and 1996 (WANG & GRACE 1999; KRISHNA et al. ISSN 1863-7221 (print) | eISSN 1864-8312 (online) 333 Scheffrahn et al.: New World Coptotermes Table 1. Coptotermes testaceus samples used for 16S molecular analysis and measurements. — Key: (*) denotes specimens from near type locality of C. niger; (**) denotes specimens from near type locality of C. crassus; (***) denotes specimens from near type locality of C. testaceus. Species UF code Country Lat. Long. Sample GenBank C. testaceus BZ 5 Belize 17.42 – 88.44 DNA KM588274 C. testaceus BZ 127 Belize 16.229 – 89.094 DNA KM588268 C. testaceus BZ 267 Belize 16.879 – 88.348 DNA KM588293 C. testaceus BO 234 Bolivia – 17.056 – 64.76 DNA KM588280 C. testaceus BO 463 Bolivia – 14.424 – 64.861 DNA KM588281 C. testaceus SA 271 Costa Rica 8.78 – 82.95 DNA KM588285 C. testaceus EC 86 Ecuador – 1.383 – 79.425 DNA KM588269 C. testaceus EC 86a Ecuador – 1.383 – 79.425 DNA KM588269 C. testaceus EC 93 Ecuador – 0.91 – 79.5 DNA KM588279 C. testaceus EC 159 Ecuador – 2.193 – 79.958 DNA KM588289 C. testaceus FG 151 French Guiana 5.038 – 52.956 DNA KM588277 C. testaceus GR 61 Grenada 12.117 – 61.733 DNA KM588278 C. testaceus GUA 658 Guatemala 15.686 – 88.645 DNA KM588292 C. testaceus** CTA 59 Honduras 15.0 – 86 DNA KM588271 C. testaceus** CTA 71 Honduras 15.5 – 88.05 DNA KM588267 C. testaceus MX 8 Mexico 16.478 – 95.157 DNA KM588276 C. testaceus* PN 161 Panama 8.669 – 80.592 DNA KM588266 C. testaceus* PN 453 Panama 9.577 – 79.409 DNA KM588284 C. testaceus* PN 961 Panama 8.22 – 81.861 DNA KM588286 C. testaceus* PN 1091 Panama 8.344 – 82.281 DNA KM588287 C. testaceus* PN 1173 Panama 8.827 – 82.697 DNA KM588290 C. testaceus* PN 1386 Panama 9.32 – 78.999 DNA KM588275 C. testaceus SA 255 Peru – 3.442 – 72.85 DNA KM588288 C. testaceus TT 11 Trinidad/Tobago 10.597 – 61.208 DNA KM588282 C. testaceus TT 603 Trinidad/Tobago 11.285 – 60.601 DNA KM588291 C. testaceus TT 766 Trinidad/Tobago 10.596 – 61.207 DNA KM588272 C. testaceus TT 1705 Trinidad/Tobago 10.159 – 61.005 DNA KM588273 C. testaceus SA 205 Venezuela 3.173 – 65.675 DNA KM588283 C. testaceus SA 204 Venezuela 3.173 – 65.675 DNA KM588270 C. testaceus*** FG 493 French Guiana 5.063 – 53.058 Measure — C. testaceus** HN 61 Honduras 15.755 – 87.455 Measure — C. testaceus** HN 375 Honduras 14.358 – 87.149 Measure — C. testaceus* PN 299 Panama 9.122 – 79.716 Measure — C. testaceus* PN 290 Panama 8.614 – 80.113 Measure — C. testaceus TT 1720 Trinidad/Tobago 10.214 – 61.631 Measure — 2013) of which 18 were reverted into synonymy (HUANG of Coptotermes have been described, all from tropical et al. 2000; LI et al. 2011). Twenty­two species of Cop­ America: C. crassus Snyder, C. niger Snyder, and C. totermes are currently recognized in China of the 44 testaceus (Linnaeus). The two exotic species from Asia, total species in the Oriental Region. Since 1968, a sin­ C. formosanus Shiraki and C. gestroi (Wasmann), have gle new species has been described outside of China in become invasive pests in the subtropical Nearctic region New Guinea (BOURGUIGNON & ROISIN 2010). Three Cop­ and the Neotropics, respectively (CONSTANTINO 2002; totermes species are known from Africa (HARRIS 1966) SCHEFFRAHN & SU 2005). A third enigmatic non­native and recent 16S marker evidence indicates a new species species from Africa, C. sjostedti (Holmgren), is estab­ from Tanzania and Mozambique (JWA, pers. comm.). lished on the West Indian island of Guadeloupe (SCHEFF­ Six Coptotermes species are recorded from Australia; RAHN et al. 2004). however, their identities are still not fully resolved (LO et Coptotermes crassus, C. niger, and C. testaceus have al. 2006). The five Papuan species are well defined mor­ traditionally been identified based on geographical loca­ phologically (BOURGUIGNON & ROISIN 2010). The field of tion with C. crassus spanning Mexico to Panama, C. ni­ molecular genetics is emerging as primary evidence for ger from Guatemala to Colombia, and C. testaceus from identifying cryptic species and synonymies in the mor­ Panama through South America (BECKER 1953; CONSTAN­ phologically challenging rhinotermitid genera of Reticu­ TINO 1998). SCHEFFRAHN et al. (2005) began to doubt the litermes (AUSTIN et al. 2005), Heterotermes (SZALANSKI validity of C. crassus and C. niger based on regional et al. 2004), and Coptotermes (SCHEFFRAHN et al. 2004; morphological comparisons and elected to list the sen­ YEAP et al. 2009). ior moniker, C. testaceus, as the species they found in The history of Coptotermes in the New World is Nicaragua. Over the last 25 years, we have conducted relatively uncomplicated. Only three endemic species many deliberate termite diversity surveys throughout 334 ARTHROPOD SYSTEMATICS & PHYLOGENY — 73 (2) 2015 Fig. 1. New World Coptotermes localities from the University of Florida (UF) collection and type localities from literature. An intercepted shipment and extirpated population of C. formosanus in San Francisco and La Mesa, Calif., respectively, are excluded. All other UF Copto­ termes samples were collected from land­based populations. tropical America yielding over 34,000 colony samples of to a Leica DFC 425 camera. Measurements were taken which Coptotermes compose about 5% of the total. We with a micrometric reticule on the eyepiece of an Olym­ herein use morphology and molecular genetic sequences pus SZX9 stereomicroscope. derived from the surveyed material to reduce, by syn­ We microscopically examined and identified 1,977 onymy, the endemic New World Coptotermes fauna to a samples of the New World­collected Coptotermes. Of single species, C. testaceus. these, we measured specimens selected from colony samples collected at or near the type localities (Table 1): C. crassus (Honduras), C. niger (Panama) and C. testa­ ceus (French Guiana and Trinidad and Tobago). The 2. Materials and methods following morphometric characters were measured, in­ dicating, in parenthesis, the numbered measurement as defined by ROONWAL (1970): soldiers (Table 2) – LH, 2.1. Morphology length of head capsule (9); HH, height of head without postmentum (21); WH, maximum width of head (19); The distribution map (Fig. 1) was created using ArcGIS LM, maximum length of left mandible (37); HF, height desktop ver. 10.1 (ESRI, Redlands, CA) from data ex­ of fontanelle; WF, width of fontanelle; LP, length of pro­ clusive to the University of Florida termite collection, notum (65); WP, width of pronotum (68) and LT, length Fort Lauderdale Research and Education Center, Davie, of hind tibia (85). Alates (Table 3) – LW, length of ante­ Florida. The photographs of the soldiers and alates were rior wing with scale (73); WW, maximum width of an­ taken using a Leica M205C stereomicroscope attached to terior wing; LS, length of anterior wing scale (76); LP, a Leica DFC 425 camera and then the image sequences length of pronotum (65); WP, width of pronotum (68); were combined with Helicon Focus 6 software.
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