o o V I o MS e Mm us ISSN 0006-9698 Cambridge, Mass. 26 May 2016 Number 548 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) James K. Wetterer,' David Lubertazzi,^ Jignasha D. Rana^ and Edward O. Wilson^ Abstract. The Caribbean region is considered a threatened biodiversity hotspot because of its high levels of biotic endemism and widespread habitat destruction. On Caribbean islands, as elsewhere in the world, ants are a critical component of virtually every terrestrial ecosystem. Nonetheless, the ants of most Caribbean islands remain poorly known. Here, we reviewed historical accounts, compiled published and unpublished ant records, and collected new specimens to document the diversity of ants on the Caribbean island of Barbados, including summary accounts for each species. We expected Barbados to have a relatively depauperate native ant fauna with few endemic species because of high isolation, young age, flat terrain, and high degree of habitat destruction. Furthermore, we expected to find many exotic ants, introduced throughout a long history of international commerce dating back to the 16(X)s. Our investigations increased the list of ant taxa known from Barbados to 69 (46 New World and 23 Old World). Of these, 62 are represented among specimens we collected, and seven are New World species only known from earlier records. We found that Barbados has fewer New World ant species and more Old World ant species than are known for Grenada and St. Vincent, two neighboring Caribbean islands of similar size. Most of the New World ant species in Barbados are continuously distributed from South American and Trinidad, through the Lesser Antilles and are probably native (i.e., predating human arrival). Some New World species in Barbados, however, may be exotics introduced through human commerce, as are all Old World ant species present. Six previously recorded New World species that we did not find may now be extinct in Barbados. These include Attci cephalotes liitea and Crematogaster brevidentata, the only two ant taxa thought to be endemic to Barbados, each collected just once >110 years ago. Key words: ants; biogeography; exotic species; island biogeography; West Indies INTRODUCTION ' Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive, Jupiter, Florida 33458, U.S.A.; e-mail: wetterer@fau.edu. ^Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. The Caribbean region is recognized as a threatened biodiversity hotspot because of its high levels of biotic endemism (e.g., 58% of terrestrial plant species and 51% of terrestrial vertebrate species) and widespread habitat destruction (only 10-15% intact The President and Fellows of Harvard College 2016. 7 B REV I ORA No. 548 Saoiia Mona Si Thi>ma;> Jolin Vieques p'.Tortola ^)f\Anauilla Atlantic Ocean * Si Marlin ^Barbuda Eastern Greater Antilles si^iu^' Nevi^^ t^Anligiia Monlserral^ (luadeloiijK* Marie-dalante Caribbean Sea Lesser Antilles (^IXnniniea 2S W " % Martinique St i.ueia St Vincent A , Barbadc« Aruba ( uravao Leevvarcl Antilles % . ISO Bonaire I^i Blanquilla (? Oi 'I'ortuaa'Marganta (irenada I'obago South America Figure 1. Major islands (>50 km^) of the eastern Caribbean. Unshaded areas are now submerged but were dry land 15 kya when sea levels were ~ 150 m lower. native vegetation remaining) (Solorzano et al, 2005). On Caribbean islands, as elsewhere in the world, ants are critical components of virtually every terrestrial ecosystem (e.g., as scavengers, predators of other arthropods, seed dispersers, etc.) (HoUdobler and Wilson, 1990). Despite their importance, the native ants of most Caribbean islands have remained largely unknown. Now, several destructive exotic ant species are spreading through the region threatening native biodiversity. Here, we have compiled published and un- published ant records and collected new speci- mens to document the diversity of ants from Barbados. Barbados is the fourth largest island of the Lesser Antilles (Fig. 1). Compared with other islands of the Lesser Antilles of similar size, we expected Barbados to have a rela- tively depauperate native ant fauna based on several factors. First, Barbados is the most isolated island of the Lesser Antilles, far to the east of all other islands (Fig. 1). Addi- tionally, Barbados has never been connected to any other land mass, so all terrestrial species that have colonized Barbados had to cross the intervening ocean. Barbados is also one of the youngest islands of the Lesser Antilles, emerging only about 1 million years ago. Finally, compared with other major island in the Lesser Antilles, Barbados is flatter, more developed, and with less intact forest. Soon after European colonization in the 1600s, settlers quickly cleared most of the 2016 ANTS OF BARBADOS (IIYMENOPTERA, FORMICIDAE) 3 forest for agriculture. In contrast, we ex- pected Barbados to be home to many exotic ant species, introduced throughout a long history of international commerce. Barbados has traditionally depended heavily on im- ports from abroad, notably lumber and other building material (Peterson, 1973). Published ant records from Barbados Early reports on the natural history of Barbados mentioned several types of ants (Clark, 1670; Hughes, 1750; Coke, 1808-11; Schomburgk, 1848; see Appendix 1). These accounts all appear to concern well-known pest species: a leaf-cutting ant {Atta sp.), a large carpenter ant [Camponotus atriceps (Smith)], the African big-headed ant [Pheidole megacephala (Fabricius)], the tropical fire ant [Solenopsis geminata (Fabricius)], the ghost ant [Tapinoma melanocephalum (Fabricius)], and the little fire ant [Wasmannia auropunc- tata (Roger)]. Unfortunately, no specimens exist to confirm these early reports, so the species identities in these earliest reports remain uncertain (see Appendix 1). There- fore, despite anecdotal evidence on the importance of these six pest ants in the early history of Barbados, we did not include these early reports in our analyses. In the oldest published ant records with extant voucher specimens. Smith (1877) de- scribed Pseudomyrma pilosula Smith and Pseudomyrma variabUis Smith [both = Pseu- domyrmex gracilis (Fabricius)] from Barbados (date and collector unknown). Additionally, Ward (1989) reported that a specimen of Pseudomyrmex maculatus (Smith) was on the same pin as Smith’s P. variabUis type specimen and presumably came from the same source. Ford (1893) described Atta lutea Ford ( = Atta cephalotes lutea Ford) based on speci- mens that W. G. Jeffreys collected in Barbados. Wheeler (1923) reported addition- al ant specimens that Jeffreys collected in Barbados (presumably before 1893): Trigly- phothrix striatidem Emery (= Tetramorium lanuginosum Mayr) and Camponotus sexgut- tatus grenadensis Ford (= Camponotus sex- guttatus Fabricius). On his way to Colombia in 1896, Ford himself made a short stop in Barbados and collected ants. In several subsequent publica- tions, Ford (1897, 1901, 1902, 1912a, c,d, 1913) reported 13 currently valid ant taxa from Barbados: Brachymyrmex cordemoyi Ford, C. sexguttatus, C. sexguttatus grena- densis (=C. sexguttatus), Cardiocondyla emeryi Ford, Crematogaster brevispinosus brevidentata Ford ( = Crematogaster breviden- tata Ford), Crematogaster steinlieili Ford, Monomorium floricola (Jerdon), Odontoma- chus haematoda insularis Guerin { = Odonto- machus insularis Guerin), Prenolepis vividula antiliana Ford [=Nylanderia vividula antil- lana (Ford)], Pheidole fallax jelskii antillensis Ford {= Pheidole jelskii Mayr), Pheidole subarmata elongatula Ford { = Pheidole sub- armata Mayr), Platythyrea punctata pruinosa Mayr [= Platythyrea punctata (Smith)], S. geminata, and W. auropunctata. Crawley (1921) described Rhizomyrma marshalli Crawley (= Acropyga goeldii Ford) from Barbados. Wheeler (1923) compiled a list of ant taxa known from Barbados based on published records of Ford, plus new material collected by Stoner (in 1918), Wheel- er (in 1920), and Ballou (in 1922). Wheeler’s (1923) list included five previously unreported species: Brachymyrmex heeri obscurior Ford {^Brachymyrmex obscurior Ford), Cremato- gaster carinata Mayr, Crematogaster brevis- pinosa minutior Ford {= Crematogaster cri- nosa Mayr), Monomorium minutum Mayr ( = Monomorium monomorium Bolton), and Pre- nolepis longicornis Latreille [= Paratrechina longicornis (Latreille)]. In total, Wheeler (1923) listed 21 valid taxa from Barbados but omitted published records of Smith ( 1 877; P. gracilis) and Crawley (1921; A. goeldii). 4 B REV I ORA No. 548 Tucker (1953) cataloged all insects known from Barbados, including a very incomplete list of ant taxa as follows: Rhizomyrma sp. { = Acropyga sp.), R. marshalli {=A. goeldii), Brachymyrmex spp., C. sexguttatus, C. sexguttatus grenadensis ( = C. sexguttatus), Crematogaster brevispinosa Mayr, C brevis- pinosa brevidentata ( = C. br evident at a), Monomorium destructor Jerdon {=Tricho- myrmex destructor), Monomorium carbonar- ium ebeninum { = Monomorium ebeninum), P. longicornis, S. geminata, and Solenopsis corticalis Forel. Of these, C. brevispinosa, S. corticalis, and M. destructor were new records, although the first two are most likely misidentifications; we therefore excluded these records. Kempf (1972), in his catalog of Neotropical ants, included records representing 22 valid taxa of ants from Barbados, omitting records in Tucker (1953) and one other previously published record: P. jelskii (Forel, 1912b). After Kempf (1972), other published addi- tions to the Barbados ant species list are Tetramorium bicarinatum (Bolton, 1979), Gnamptogenys cf. striatula (Peck, 1981; probably = Gnamptogenys striatula Mayr), Hypoponera sp. (Peck, 1981), and P. macula- tus (Ward, 1989; see above). Thus, before our current collecting efforts, there were reliable published records from Barbados of 26 currently valid ant taxa, plus two taxa identified only to genus. Some of our current records of tramp ant species in Barbados have been reported earlier, in review papers on these species (Wetterer, 2008, 2009a-c, 2010a^, 201 la^, 2012a-e, 2013a, b, 2014a- c; MacGown and Wetterer, 2013). METHODS We searched for ant specimens from Barbados in the collections of the American Museum of Natural History, Museum of Comparative Zoology, and Smithsonian Institute. We also pinned and identified Barbados ant specimens collected by Jo- hanna P. E. C. Darlington in March 1987. Stefan P. Cover (SPC) and one of us (EOW) collected ants in Barbados 17-20 March 1998. Another author (JKW) collected ants in Barbados 25-29 November 2003, 16-22 June 2006, and 2-14 May 2014 (see Appendix 2 for site details). We conducted much of our sampling in the remaining intact forest, notably in Hackleton’s Cliff and Turner’s Hall Woods, and in small forest patches that remain in deep gullies formed from collapsed caves (e.g., Jack-in-the-Box gully). We collected ants following standard methods used in numerous earlier ant faunal surveys (e.g., Wetterer and Wetterer, 2004; referred to as “direct sampling” by Bestel- meyer et al, 2000). The primary goal was to collect the maximum number of different ant species in the time allotted. To do this, we collected at numerous sites in the widest range of habitats accessible and permitted, spending more time at sites where the new or rare species accumulation rate was greater and adjusting collecting techniques to best sample different habitats (Longino, 2000). We have found that this method proves much more efficient than standardized techniques de- signed for evaluating relative abundances of species and making detailed comparisons among individual sites (such as the Ants of the Leaf Litter [ALL] protocol, which re- quires a minimum of three days sampling per site; Agosti and Alonso, 2000; Fisher et al., 2000). We hand-collected ants, made vegeta- tion beatings, and collected complete colonies from under rocks and logs and inside epi- phytes, twigs, and dead wood. We sifted soil and litter samples using a Davis sifter. When possible, we collected not only workers and sexuals, but also larvae and pupae. One focus was surveying intact forests, where native and endemic species are most commonly found. We also surveyed highly disturbed areas. 2016 ANTS OF BARBADOS (I lYMENOPTERA, FORMICIDAE) 5 particularly around ports, where invasive exotic ants dominate. We have found that botanical gardens, urban parks, and outdoor food markets are particularly rich sources of both native and exotic ant species. John T. Longino confirmed the identifica- tion of our Crematogaster specimens. Jose Pacheco and William Mackay identified a subset of our thief ants {Solenopsis spp.) collected in 2003 and 2006. We did not include unidentified thief ants in our record counts. We classified each species as either New World or Old World and evaluated its known range based on information from a wide range of sources (see Results). Kempf s (1972) catalog of Neotropical ants and Deyrup’s (2003) checklist of Florida ants were partic- ularly helpful sources of range information. We classified Old World records into five biogeographic regions: 1. The Afrotropic or Ethiopian region (22.1 million km2) includes sub-Saharan Africa, the southern and eastern coasts of the Arabian Peninsula, southern Iran, southwestern Pakistan, Madagas- car, the western Indian Ocean islands, Cape Verde Islands, and the southern mid-Atlantic islands. 2. The Palearctic region (54.1 million km^) includes Europe, northern Africa, the northern and central Arabian Peninsula, and Asia north of the Hima- layas, including the main islands of Japan, 3. The Indomalay or Oriental region (7.5 million km-) includes southeastern Pakistan, the Indian subcontinent. Southeast Asia, southern China, the Philippines, Taiwan and Japan’s Ryukyu Islands, and Indonesia west of Wallace’s line. 4. The Australasia region (7.6 million km^) includes Australia, New Guinea, Indonesia east of Wal- lace’s Line, Vanuatu, the Solomon Islands, New Caledonia, and New Zealand. 5. The Oceania region (1.0 million km^) includes the Pacific islands of Eiji, Micronesia, and Polynesia (except New Zealand). RESULTS We documented 69 ant taxa in Barbados (62 are represented among specimens we collected and seven are based solely on earlier records; Tables 1, 2). We found that the one published record of M. monomorium in Barbados was based on misidentified specimens of M. ebeninum. We provisionally considered the published records of B. corde- moyi, O. insularis, N. vividula antillana, and Pheidole fallax in Barbados to be misiden- tifications of B. cf obscurior, Odontoma- chiis ruginodis, Nylcinderia cf. steinheili, and P. jelskii, respectively (see Species Accounts below). We judged 46 of the ant taxa found in Barbados to be of New World ant species and the other 23 to be Old World exotics (Tables 1, 2; see Species Accounts below). Our 1998, 2003, 2006, and 2014 collections yielded a diminishing number of previously unrecorded species, finding 26, 10, 5, and 1, respectively. We found several ant species at just one or two sites, indicating that it is likely we overlooked at least a few species. SPECIES ACCOUNTS C&W = S. P. Cover and E. O. Wilson (1998), JW = J. K. Wetterer (2003, 2006, and 2014). We list vial numbers for specimens we collected (see Appendix 2). * = previously published specimen records from Barbados identified as a different species. BMNH = Natural History Museum London, MCZ = Museum of Comparative Zoology, and SI = Smithsonian Institution. 1. Acropyga goeldii Forel Published Record. No site data; in soil by sugarcane root (1914; J. R. Bovell; Crawley 1921; as Rhizomyrma marshallr, type in BNHM). Specimens Examined. C&W, one site (B68, B74). JW, two sites (2006: 358; 2014: 95). Crawley (1921) described R. marshalli Crawley (= A. goeldii) from specimens that J. R. Bovell collected in Barbados. John Redman Bovell (1855-1928) was the Super- intendent of Agriculture in Barbados, known 6 B REV I ORA No. 548 T.^ble 1. New World ants of Barbados. First = earliest record. New = no. of new site records (1998-2006). First New NWt OW5 Crematogaster crinosa 1920 91 CST bgv Camponotiis sexgiittatiis 1896 80 CST bgv Brachymyrmex cf. obsciirior 1896 74 CST bgv Odontornachiis ruginoclis 1896* 52 CST bgv Wcismannia aiiropiinctata 1896 51 CST bgv Pheidole sculptior 1998 47 CST bgv Cyphomynnex miniitus 1998 46 CST bgv Pheidole jelskii 1896 39 -ST bgv Sulenopsis geminata 1896 39 CST bgv Nylanderia cf. steinheili 1896* 36 CST bgv Brachymyrmex cf. heeri 1998 34 CST bgv Monomorium ebeninum 1918* 33 CST bgv Solenopsis globidaria 1998 31 CST bgv Striimigenys margaritae 1998 29 CST bgv Pheidole exigua 1998 24 CST bg- Anochetus mayri 1998 23 CST bgv Rogeria curvipiibens 1987 23 CST bgv Platythyrea punctata 1896 21 CST bgv Anochetus inermis 1998 16 CST bgv Gnamptogenys striatula 1979* 16 CST b— Pheidole transversostriata 1998 15 -ST b— Camponotus atriceps 1998 14 CST bgv Crematogaster curvispinosa 1998 14 CST bgv Odontornachiis bauri 1998 14 CST bgv Solenopsis pollux 2003 13 CS- bgv Pheidole subarmata 1896 10 CST bgv Rogeria foreli 2003 10 CST bgv Syllophopsis subcoeca 1998 9 — bg- Solenopsis zeteki 2006 8 CS- bg- Solenopsis azteca 2003 8 CS- bgv Nylanderia cf. fulva 1998 7 CST b-v Striimigenys louisianae 2006 7 CST b-v Striimigenys eggersi 2003 5 CST bg- Pheidole susannae 2006 4 CST b— Solenopsis pygmaea 2003 4 C— bgv Leptogenys piibiceps 2003 3 -ST bgv Acropyga goeldii 1914 3 CST b— Striimigenys sclndzi 1998 2 CST b— Pseudoponera stigma 2014 1 CST bgv Hypoponera opaciceps <1960 0 CST b— Crematogaster carinata 1920 0 CST b— Crematogaster brevidentata 1896 0 — b— Crematogaster steinheili 1896 0 — b— Atta cephalotes lutea <1893 0 — b— Pseudomyrmex gracilis <1877 0 CST b-v Pseudomyrmex macula tus <1877 0 -ST b— LG— LGBF LGBF AuOc LGBF LGBF AfPa— AuOc LG— LGBF LGB- LGBF AfPalnAuOc LGB- —Pa LGBF —Pa LGBF LGBF LGBF _G— LGBF —Pa LGB- LGBF L-B- LG— —Pa LG— L LG— LG— LGB- LG— LG— L LG— LG-F LGBF LGBF LG— -G— LGB- LGBF InAuOc LGBF — PalnAuOc LGB LG-F —Pa Oc *First published record uncertain identification. tNew World Range: C = Central America, S = South America. T = T rinidad; b = Barbados, g = Grenada, v = St. Vincent. §01d World Range: L = other Lesser Antilles, G = Greater Antilles, B = Bahamas, F = Florida; Af = Afrotropic, Pa = , Palearctic, In = Indomalaya, Au = Australasia, Oc = Oceania. 2016 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) 7 Table 2. Old World ants of Barbados. Panitrechina longicomis Tapmoma melanocepluiliim Telnmwrium simillinnim Curdiocondyki emery i Pheidole megacepludci Pkigiulepis cdluaudi Monomorium floricola Trichomynnex destructor Tetriimoriiim cakkiriiim Ccirdiocondyla minutior Hypoponera pimctatissima Strumigeriys rogeri Tetramorium kmuginosiim Ccirdiocondyla vemistida Strumigenys emniae Ccirdiocondyla maiiritanica Cardiocondyla obsciirior Nylanderia boiirbonica Pheidole teneriffana S triiniigenys membran if era Syllophopsis sechellensis Leptogenys maxillosa Tetramorium bicarinatum *Symbols as in Table 1. First New NW* OW* 1920 61 CST bgv LGBF AfPalnAuOc 1998 53 CST bgv LGBF AfPalnAuOc 1998 29 CST bgv LGBF AlPalnAuOc 1896 27 CST bgv LGBF AfPalnAuOc 2003 28 CST b-v LGBF AfPalnAuOc 1998 27 — bg- L AfPalnAuOc 1896 17 CST bgv LGBF AfPalnAuOc < 1953 11 CST bgv LGBF AfPalnAuOc 1998 9 CST b— LGBF AfPalnAuOc 1998 6 CST bg- LGBF Af — InAuOc 1998 6 CS- bgv LGBF AfPalnAuOc 1998 6 CST bgv LGBF AfPalnAuOc < 1893 5 C-T b-v LG-F AfPalnAuOc 1998 4 CST b— LGBF Af — In — Oc 2003 4 CST bgv LGBF Af — InAuOc 2006 2 C— bg- G-F AfPalnAu — 1998 2 CS- b— -G-F AfPalnAuOc 1998 2 C-T b-v LGBF AfPalnAuOc 2003 2 -S- bg- LG— AfPa 1998 2 CST b— LGBF AfPalnAuOc 2003 2 — b— L Af— InAuOc 2006 1 CS b-v LG— AfPaIn — Oc 1936 1 CST bgv LGBF AfPalnAuOc for his research with sugarcane. His portrait is now on the two dollar bill of Barbados. LaPolla (2004) determined R. marshalli to be a junior synonym of A. goeldii. Acropyga goeldii is also known from Brazil, Paraguay, Surinam, Guyana, Trinidad, Venezuela, Colombia, and Panama (LaPolla, 2004). Acropyga spp. are obligate mutualists of root-feeding Hemiptera. Acropyga are rarely collected except when researchers are specif- ically seeking them. Acropyga goeldii, often found tending mealy bugs on sugarcane roots, may be fairly common in Barbados, where sugarcane is a major crop. 2. Anochetus inermis Andre Published Records. None. Specimens Examined. C&W, two sites (B13, B21, B51). JW, 14 sites (2003; 87, 97, 99, 100, 119, 121; 2006; 359, 380, 410, 441, 442, 453; 2014; 27, 112). This widespread New World trap-jaw ant is “a nearly uniform tawny to yellowish-red in color” (Brown, 1978) and is the larger and lighter colored of the two Anochetus species present is Barbados. We found this species in both natural and highly disturbed habitats. Anochetus inermis is native to South America and the West Indies, but populations in the Bahamas may be exotic (Deyrup et al., 1998). 3. Anochetus mayri Emery Published Records. None. Specimens Examined. C&W, one site (B19, B30). JW, 22 sites (2003; 104, 107, 110, 130; 2006; 346, 348, 349, 359, 364, 365, 442; 2014; 9, 64, 71, 102, 107, 112, 169, 174, 180, 185, 189). This widespread New World trap-jaw ant is the smaller and darker of the two Anochetus species present is Barbados. Brown (1978, 8 BREVIORA No. 548 p. 617) wrote about the difficulties in defining the taxonomic boundaries of A. mayri as follows (in abbreviated form): ''A. mayri is variable in body size, eye size, antennal scape length, color and sculpture, as well as size and details of form and dentition of the mand- ibles. It is not always easily separated from smaller specimens of the inermis complex on the one hand, or from neglectus on the other, and some of the variation raises the suspicion that mayri may include two or more sibling species.” “The mayri complex is widespread in the West Indies and shows there wide variation in size, color and sculpture.” “After prolonged study of this material, I cannot find any way to separate it into two species, or even into reasonably clearcut geographical forms.” We found this species in both natu- ral and highly disturbed habitats. Anochetus mayri is native to South America, Central America, and the West Indies, but popula- tions in the Bahamas and Florida may be exotic (Deyrup et al., 1998, 2000). 4. Brachymyrmex cf. heeri Published Records. None. Specimens Examined. C&W, two sites (B69, B99, B102). JW, 32 sites (2003: 91, 93, 95, 99, 100, 121, 130; 2006: 337, 343, 348, 349, 354, 363, 365, 380, 442, 451, 453; 2014: 9, 60, 64, 74, 87, 92, 93, 94, 1 19 135, 152, 156, 160, 174). Forel (1874) described Brachymyrmex heeri Forel from greenhouses in Switzerland. Unfortunately, the taxonomy of the genus Brachymyrmex is in disarray, and the species boundaries are not clear. It is likely that there are two or more species identified under the name B. heeri. Some specimens we have identified as B. cf heeri have been identified by other researchers as Brachy- myrmex mimitiis Forel. Small yellow Brachy- myrmex are widespread in the New World. We found this species in both natural and highly disturbed habitats. It is fairly common in Barbados, but it is probably often over- looked because of its very small size. 5. Brachymyrmex cf. obscurior Published Records. No site data (1896; A. Forel; Forel, 1912d, as Brachymyrmex cor- demoyi Forel). Bridgetown (1920; W. M. Wheeler; Wheeler, 1923, as B. obscurior). No site data (Sirjusingh et al, 1992, as B. obscurior). Specimens Examined. No site data (1934; N. A. Weber; MCZ). C&W, one site (B9). JW, 73 sites (2003: 87, 91, 92, 93, 95, 97, 99, 100, 108, 109, 112, 115, 116, 118, 121, 133; 2006: 354, 358, 363, 371, 373, 374, 375, 376, 379, 381, 383, 387, 410, 412, 418, 422, 440441, 442, 451, 456, 457; 2014: 6, 9, 26, 42, 44, 52, 54, 55, 56, 59, 60, 80, 81, 83, 85, 87, 92, 93, 94, 107, 109, 124, 135, 143, 144, 145, 151, 152, 154, 155, 156, 157, 160, 165, 168, 190). Forel (1893) described B. heeri obscurior (= B. obscurior) from St. Vincent, writing that it differed from B. heeri in being brownish in color and slightly hairier and differed from Brachymyrmex patagonicus Mayr in being smaller, slightly hairier, and with slightly longer scapes. Small brown Brachymyrmex are wide- spread and very common in the New World, particularly in highly disturbed environ- ments. Brachymyrmex cf obscurior is variable in size and color, making identification difficult. Some specimens that we list as B. cf. obscurior have been identified by others as B. cordemoyi. It is likely that there may be two or more species identified under this name. Brachymyrmex cf obscurior has been reported from scattered sites outside this region: Hawaii (Wheeler, 1934), New Caledonia (Delsinne et al, 2001), and Samoa (Wetterer and Vargo, 2003). MacGown et al. (2007) consider it an exotic in North America, where it is widespread in southern Florida, with isolated records from Mississippi and Georgia (Ipser et cd., 2005; MacGown et al.. 2016 ANTS OF BARBADOS (IlYMENOPTERA, FORMICIDAE) 9 2007). Brachymynnex cf. obscurior is very common in Bermuda, where it co-exists in close proximity with the two major pest species there: P. megacephaki and Line- pithema biimile (Mayr) (see Wetterer and Wetterer, 2004). 6. Camponotiis atriceps (Smith) Published Records. None. Specimens Examined. C&W, two sites (B78, B97). JW, 12 sites (2003; 89, 123129; 2006; 403409, 415, 417, 423439, 446, 459; 2014: 37, 68, 78, 126). The enormous carpenter ant C. atriceps has been recorded from sites across Central and South America, as well as Trinidad and Tobago (Kempf, 1972). Although Hashmi (1973) synonymized many C. atriceps sub- species, it is likely that C. atriceps is not a single species, but rather a species complex. Camponotiis atriceps nocens Wheeler is known only from Grenada and St. Vincent. In Barbados, C. atriceps nests inside dead branches in wooded areas. Camponotus atriceps has been found in greenhouses in Italy (Jucker et al., 2008) and The Nether- lands (Boer and Vierbergen, 2008). 7. Camponotus sexguttatus (Fabricius) Published Records. No site data (1896; A. Forel; Forel, 1897). Bridgetown (1920; W. M. Wheeler; Wheeler, 1923). Specimens Examined. Bridgetown (1920; W. M. Wheeler; MCZ). Turner’s Hall Woods (1987; JPEC Darlington; MCZ), C&W, four sites (B22, B26, B35, B42, B47, B75, B83, B92, B94, B107). JW, 76 sites (2003: 87, 88, 89, 91, 99, 100-102, 104, 107, 108, 109, 110, 118, 120, 121, 122, 130, 131; 2006: 341-342, 343-345, 346, 348, 350-352, 355-356, 357, 361-362, 364, 366, 368, 370, 380, 381, 388, 390-392, 393-394, 395-398, 399-402, 410-411, 412-414, 420-421, 422, 440-441, 445, 447, 450, 453-454, 457, 460; 2014: 13, 29, 31, 45, 46, 58, 63, 67, 73, 77, 89, 90, 96, 99, 101, 104, 115, 118, 122, 125, 138, 143, 148, 161, 164, 167, 173, 179, 180, 188). This widespread New World carpenter ant (Kempf, 1972) commonly nests inside dead branches. It is extremely common in many habitats. In C. sexguttatus sexguttatus, mi- nor workers are uniformly dark, except for light spots on the gaster, whereas major workers also have a lighter colored head. Camponotus sexguttatus is considered an exotic in Florida and the Bahamas (Deyrup et al. 1998, 2000). 7a. Camponotus sexguttatus grenadensis Forel (= C. sexguttatus) (stat. rev.) Published Records. No site data (^1893; W. G. Jeffreys; Wheeler, 1923). No site data (1896; A. Forel; Forel, 1897). No site data (1918; D. Stoner; Wheeler, 1923). Forel (1897) described C. sexguttatus gre- nadensis from Barbados and Grenada. This form has a much lighter colored head and thorax than the typical form. Emery (1898) considered grenadensis to be a junior syno- nym of Camponotus biguttatus Emery, known only from Bolivia. Wheeler (1923), however, placed biguttatus as a subspecies of sexgutta- tus and separated it from grenadensis and all other subspecies of sexguttatus in the first couplet of his dichotomous key. Wheeler (1923, p. 8) wrote that biguttatus was the sole subspecies with “epinotum in profile convex and arcuate above”; all other subspecies have “epinotum in profile straight or more or less depressed above.” Wheeler (1923, p. 8) also described two additional subspecies of sexgut- tatus from the Lesser Antilles, both of which have lighter head, thorax, and gaster than the typical form: antiguensis Wheeler from Anti- gua (“gaster with cream-colored spots on second segment”) and montserratensis from Montserrat (“gaster immaculate”). Emei7 (1925), in his catalog of known ants, listed biguttatus as a variety of seguttatus and with grenadensis as a junior synonym of biguttatus. Emery (1925), however, did not consider antiguensis and montserratensis, so presum- ably he had not read Wheeler (1923) and did 10 BREVIORA No. 548 not know the characters Wheeler (1923) used to separate biguttatus and grenadensis. Nonetheless, Bolton (1995) listed antiguensis and montserratensh as valid subspecies, but grenadensis as a junior synonym of the Bolivian biguttatus. Following Wheeler (1923), we do not consider grenadensis to be a junior synonym of biguttatus. Although grenadensis specimens are often strikingly lighter in color than the typical form, W. Mackay (personal communication) does not consider this a distinct subspecies. We there- fore have not separated out lighter forms on our list of C sexguttatus records above, and we provisionally consider C. sexguttatus gre- nadensis to be a junior synonym of C. sexguttatus. 8. Cardiocondyla emery i Forel Published Records. No site data (1896; A. Forel; Forel, 1912c). North of Bridgetown (1995; Seifert, 2003). Review (Wetterer, 2012d). Specimens Examined. JW, 27 sites (2003; 86, 89, 95, 114, 116; 2006: 371, 375, 387, 451; 2014: 6, 44, 54, 56, 59, 60, 81, 83, 85, 87, 145, 152, 155, 156, 157, 160, 165, 190). Cardiocondyla emeryi, an African native, is a well-known tramp species distributed worldwide by human commerce (Wetterer, 201 2d). This species is most common in disturbed environments and often co-occurs with dominant exotic ant species (Wetterer, 201 2d). Because of their very small size, all Cardiocondyla species are probably often overlooked. 9. Cardiocondyla mauritanica Forel Published Records. Review (Wetterer, 2012f). Specimens Examined. JW, one site (2006: 356). Cardiocondyla mauritanica is an African native that has become a cosmopolitan tramp species (Wetterer, 20120- In the New World, many older studies recorded this species as its junior synonym, Cardiocondyla ectopia Snelling. This species is most com- mon in disturbed environments. 10. Cardiocondyla minutior Forel Published Records. Review (Wetterer, 2014b). Specimens Examined. C&W, one site (B46). JW, five sites (2003: 112; 2014; 108, 145, 154, 168). Cardiocondyla minutior is an Indo-Malayan native that has become a cosmopolitan tramp species (Wetterer, 2014b). In the New World, many older studies incorrectly recorded this species as Cardiocondyla nuda (Mayr). 1 1 . Cardiocondyla obscurior Wheeler Published Records. None. Specimens Examined. C&W, one site (B3). JW, two sites (2006: 364; 2014: 95). Cardiocondyla obscurior is an Old World tramp species (Seifert, 2003). This species is often misidentified as another tramp, Cardiocondyla nroughtonii (Forel), but may be distinguished from this species based on coloration and discriminate function analysis (Seifert, 2003). These two species also show ecological differences, with C obscurior being more arboreal and C. wrougbtonii more sub- terranean (SPC, personal observation). 12. Cardiocondyla venustula Wheeler Published Records. None. Specimens Examined. C&W, one site (B62). JW, three sites (2006: 373; 2014; 94, 168). Cardiocondyla venustula is an Old World tramp species (Seifert, 2003). This species is most common in disturbed environments. 13. Crematogaster crinosa Mayr Published Records. Bridgetown (1920; W. M. Wheeler; Wheeler, 1923). Specimens Examined. C&W, two sites (B15, B54). JW, 89 sites (2003: 87, 89, 91, 92, 93, 95, 97, 99, 100, 103, 104, 107, 109, 115, 118, 130; 2006: 337, 338, 339, 343, 346, 349, 354, 357, 365, 369, 370, 371, 375, 376, 380, 383, 387, 390, 394, 395, 399, 410, 418, 419, 440, 442, 447, 450, 451, 457; 2014: 9, 21, 2016 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) II 26, 43, 44, 49, 50, 53, 54, 57, 59, 62, 66, 71, 74, 80, 82, 84, 85, 91, 92, 93, 94, 95, 98, 100, 102, 107, 109, 118, 135, 141, 143, 144, 145, 152, 154, 156, 157, 162, 174, 181, 190. Longino (2003) recorded this New World species from the West Indies (Grenada, Nevis, and Saint Vincent) and from many sites in South and Central America, north to Texas; Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Guyana, Mexico, Panama, Para- guay, Peru, Uruguay, Texas, and Venezuela. This was our most commonly collected ant species in Barbados, both in forests and disturbed habitats, often occurring in very high densities. We most often found this species nesting in trees, but it also nests in the ground. 14. Crematogaster curvispinosa Mayr Published Records. None. Specimens Examined. C&W, one site (B60). JW, 13 sites (2003: 87, 100, 122; 2006: 346, 395, 397, 419, 453, 459; 2014: 27, 1 18, 169, 181). Longino (2003) reported this species from the West Indies (Saint Lucia) and from many sites in South and Central America, north to Mexico (Bolivia, Brazil, Colombia, Costa Rica, Guatemala, Guyana, Mexico, Nicaragua, Panama, Peru, and Venezuela). In the West Indies, this species has also been reported from Martinique (Forel, 1893, 1912b) and Saint Vincent (Forel, 1893). Although Longino (2003) found this species was most common in disturbed habitats, we collected it primarily in forest areas. 15. Cyphomyrmex minutus Mayr Published Records. None. Specimens Examined. C&W, three sites (Bll, B16, B37, B50, B81, B88, B106). JW, 43 sites (2003: 86, 89, 93, 97, 99, 100, 104, 107, 112, 131; 2006: 338, 343, 346, 349, 364, 367, 370, 371, 376, 387, 390, 394, 399, 410, 441, 446, 451, 459; 2014: 42, 56, 87, 100, 108, 113, 121, 124, 154, 156, 157, 165, 169, 180, 185). Cyphomyrmex minutus is common across the West Indies, although it has often been misidentified as Cyphomyrmex rimosus. According to Longino (2004): “Snelling and Longino (1992) distinguished three sim- ilar species, hamulafus, minutus, and rimosus, based on differences in size, pilosity, and extent of the median basal groove of the first gastral tergite. Subsequently I have not been able to differentiate these taxa. There is abundant geographic variation. In some localities it appears that there are discrete sympatric forms, but in other areas the distinction is blurred. For example, in Florida there are two discrete forms, a native species that is relatively small and an introduced species that is larger and darker. The key in Snelling and Longino would separate these into minutus and rimosus, respectively. In Costa Rica, specimens that are collected from open areas, usually by finding nests in the soil or foragers on the surface, are relatively larger and with longer scapes than specimens found in wet forest leaf litter, but the size distributions overlap. Until further evidence for discrete species is produced, I prefer to call them all C. rimosus.” In our case, we considered all Cyphomyrmex in Barbados to be C. minutus. This small and inconspicuous fungus- growing ant freezes when disturbed and often may be overlooked. Nonetheless, it is surprising that this common species has not been previously reported from Barbados. Weber (1972) presented a map indicating the range of Cyphomyrmex that includes Barbados but provided no records. Other maps in Weber (1972) seem to include Barbados in the ranges of Trachymyrmex, Mycetophylax, and Mycocepurus, but none of these genera has actually been documen- ted in Barbados. It may be that Weber’s maps were carelessly drawn. 16. Gnamptogenys striatula Mayr Published Records. None, but one proba- ble record. Coles Cave (1979; S. B. Peck; Peck, 1981, as Gnamptogenys sp. striatula complex). 12 BREVIORA No. 548 Specimens Examined. C&W, one site (B80, B82, B86, B89). JW, 15 sites (2003: 87, 89, 105-106, 130, 131-132; 2006: 343, 346, 354, 395, 455, 458, 459; 2014: 8, 76, 100). This species has been reported from sites across the Neotropics, from Mexico to northern Argentina and the West Indies (Lattke et al., 2007). We were unable to find Peck’s specimens, the only Gnamptogenys previously reported from Barbados, but we assume that they are conspecific with our specimens. 17. Hypoponera opaciceps (Mayr) Published Records. None. Specimens Examined. Barbados, soil in cane field; queen (R. W. E. Tucker; SI). R. W. E. Tucker published papers in the 1930s to 1950s. We confirmed that this specimen is correctly identified. Hypoponera opaciceps is a small, subterranean ant that feeds on minute soil arthropods. Originally from the New World, H. opaciceps has spread around the world through human commerce. Although people rarely come in contact with H. opaciceps workers, swarms of queens that can sting make this species an occasional pest. Because of its largely subterranean habits, H. opaciceps is probably often overlooked. 18. Hypoponera punctatissima (Roger) Published Records. None, but one possible record. Coles Cave (1979; S. B. Peck; Peck, 1981, as Hypoponera sp.). We were unable to find Peck’s specimens, the only previously published Hypoponera record from Barbados. Specimens Examined. C&W, three sites (B59, B87, B90, B112). JW, three sites (2006: 370; 2014: 100, 169). This African native is a well-known tramp species widely distributed throughout the tropics and subtropics (Delabie and Blard, 2002). Because of its largely subterranean habits, H. punctatissima is probably often overlooked. 19. Leptogenys maxillosa (Smith) Published Records. None. Specimens Examined. JW, one site (2006: 452). Leptogenys maxillosa is a widespread tramp species originally from Africa (Wilson and Taylor, 1967; Lattke, 2011). In the New World, this species is also known from Brazil, Cuba, Honduras, St. Thomas, and St. Vincent (Ashmead, 1900; Kempf, 1972; Lattke, 2011; P. S. Ward, unpublished record). 20. Leptogenys pubiceps Emery Published Records. None. Specimens Examined. JW, three sites (2003: 94, 100, 111). Leptogenys pubiceps is a species complex with “a distribution range restricted to the Caribbean Basin. It is found on the mainland from Heredia and Limon Provinces of Costa Rica to northern Colombia, the northern slopes of the Venezuelan Andes, the Vene- zuelan Coastal Range and Trinidad. It is found on both the Greater and Lesser Antilles” (Lattke, 2011, p. 218). 21. Monomorium ebeninum Eorel Published Records. No site data (1922; H. A. Ballou; Wheeler, 1923, as M. minutum). Specimens Examined. No site data (1922; H. A. Ballou; MCZ). No site data (1934; N. A. Weber; MCZ). C&W, one site (B44). JW, 32 sites (2003: 87, 89, 93, 97, 118; 2006: 338, 343, 354, 358, 367, 380, 390, 395, 399, 410, 412, 418, 419, 422, 440, 441, 446; 2014: 6, 8, 60, 92, 93, 94, 95, 107, 143, 165). Wheeler (1923) misidentified specimens from Barbados as M. minutum (= M. monomorium). This small black New World arboreal species is widespread and common in the West Indies and Central America (Kempf, 1972). This species has also been collected from several sites in the Elorida Keys, where Deyrup et al. (2000) considered it to be exotic. 22. Monomorium floricola (Jerdon) Published Records. No site data (1896; A. Eorel; Wheeler, 1905; Eorel, 1912c). Review (Wetterer, 2010a). 2016 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) 13 Specimens Examined. JW, 17 sites (2003; 93, 97, 99, 108, 1 12; 2006: 354, 364, 365, 380, 381; 2014: 64, 71, 87, 90, 92, 107, 190). This tiny Asian species is an extremely widespread tramp species but is rarely a major pest (Wetterer, 2010a). Fowler et al. (1993) found that M. Jloricola was the most commonly collected ant at a municipal hospital in Sao Paulo, Brazil, making up 34.5% of the ants collected. 23. Nylanderia boiirbonica (Forel) Published Records. None. Specimens Examined. C&W, two sites (B53, B63). Nylanderia boiirbonica (formerly Paratre- china boiirbonica) is an Old World tramp species spread through human commerce. In the West Indies, this species has been reported from the Bahamas (Smith, 1954; Deyrup et al., 1998), Cuba (Trager, 1984), the U.S. Virgin Islands (Wetterer and Lombard, 2010), and the Florida Keys (Deyrup et al, 1988; Wetterer and O’Hara, 2002). Wetterer (un- published data) has also collected this species on other West Indian islands, including Anguilla, Antigua, Barbuda, and the Cayman Islands. Outdoor populations of N. bourbo- nica are widespread in Florida, and indoor populations are known from several other U.S. states (e.g., see Trager, 1984; MacGown and Forster, 2005). 24. Nylanderia cf. fulva Published Records. None. Specimens Examined. C&W, one site (B66, B73). JW, six sites (2003: 108, 121; 2006; 461; 2014: 94, 95, 165). Nylanderia fulva (Emery) is a New World species that has been reported from numer- ous other sites scattered across the New World tropics and subtropics. Unfortunate- ly, almost all records of N. fulva are of questionable reliability because of problems distinguishing N. fulva from other closely related species, notably Nylanderia pubens (Trager, 1984; Gotzek et al, 2012). Nylan- deria fulva and N. pubens, however, may be distinguished using genetic analyses and male morphology (Gotzek et al, 2012). Nylanderia fulva has undergone population explosions in parts of Colombia, the south- ern United States, and on St. Croix, U.S. Virgin Islands, where locals blamed N. fulva for serious crop damage (e.g., a coconut plantation produced no coconuts because of high densities of plant-feeding Hemiptera tended by the ants; Wetterer and Keularts, 2008). An extreme population boom fol- lowed by a bust may be a common N. fulva pattern and might distinguish this species from the closely related N. pubens (Wetterer et al, 2014). 25. Nylanderia cf. steinheili Published Records. No site data [1896; A. Forel; Forel, 191 2d, as Nylanderia vividula antillana (Forel) = Nylanderia guatemalensis (Forel)]. Specimens Examined. C&W, one site (B27, B29, B40, B48). JW, 35 sites (2003: 89, 100, 101, 121, 130; 2006: 346, 349, 354, 357, 363, 364, 365, 387, 410, 422, 440-441, 446-447, 451; 2014: 9, 27, 75, 87, 90, 94, 95, 102, 110, 112, 118, 124, 169, 174, 180, 185, 189). Nylanderia steinheili (Forel) and N. guate- malensis (Forel) are New World species that are similar in size and appearance. Both species have been reported from many sites in the West Indies, but the identities of most records are uncertain. For example, exten- sive records of N. guatemalensis from Florida may all represent misidentifications of N. steinheili (Kallal and LaPolla, 2012). 26. Odontomachus bauri Emery Published Records. None. Specimens Examined. Turner’s Hall Woods (1987; J. P. E. C. Darlington; MCZ), C&W, two sites (B21, B24, BlOl, B107, B108). JW, 12 sites (2006; 365, 394; 2014: 61, 105, 116, 123, 124, 172, 178, 184, 187, 189). 14 B REV I ORA No. 548 Odontomachus bauri is a widespread New World species, known from South America, Central America, and the West Indies (Brown, 1976). This large black trap-jaw ant is common in intact forest habitats on many West Indian islands. 27. Odontomachus ruginodis Wheeler Published Records. None. Specimens Examined. No site data (<1932; R. Thaxter; MCZ). No site data (1934; N. A. Weber; MCZ). C&W, one site (B7, B14, B17). JW, 51 sites (2003: 87, 89, 93, 99, 103, 112, 114, 116, 117, 118, 119, 121, 130, 131; 2006: 337, 343, 346, 348, 353, 359, 367, 374, 376, 380, 384, 387, 390, 395, 399, 410, 412, 419, 441, 443, 446-447, 450, 453; 2014: 10, 26, 44, 61, 64, 74, 88, 107, 137, 140, 144, 152, 154, 190). Odontomachus ruginodis is a widespread New World species probably native to the West Indies (Deyrup et ah, 1998) but may be exotic in Florida (Deyrup, 1991). Brown (1976) regarded O. ruginodis as synonymous with Odontomachus brunneus. Brown {in Deyrup et cd., 1985), however, changed his mind and again separated them into two distinct species. This species is often misiden- tified as O. insularis (e.g., see below). This species is common in a wide variety of habitats, including highly disturbed urban areas. 27a. Odontomachus insularis Guerin Published Records. No site data (1896; A. Forel; Forel, 1912a). No site data (1918; D. Stoner; Wheeler, 1923). Bridgetown (1920; W. M. Wheeler; Wheeler, 1923). We believe that the several Barbados records of this valid species are actually misidentified O. ruginodis, originally de- scribed as a variety of O. insularis. True O. insularis are known from the Greater Antilles. 28. Paratrechina longicornis (Latrielle) Published Records. Garrison (date unknown; W. Norwcll; Wheeler, 1923). Bridgetown (1920; W. M. Wheeler; Wheeler, 1923). No site data (Sirjusingh et al., 1992). Review (Wetterer, 2008). Unpublished Record. North Point (1969; E. Wartburg; AntWeb, 2016). Specimens Examined. No site data (1934; N. A. Weber; MCZ). Harrison’s Cave garden (2005; J. Endeman). C&W, two sites (Bl, B5). JW, 59 sites (2003: 86, 89, 91, 93, 95, 97, 99, 100, 108, 109, 114, 116, 117, 118, 119, 121, 133; 2006: 354, 357, 363, 367, 371, 373, 374, 376, 378, 380, 382, 383, 387, 395, 412, 418, 422, 444, 448, 456, 457; 2014: 8, 26, 42, 52, 56, 59, 60, 87, 92, 93, 95, 100, 136, 143, 144, 145, 152, 154, 160, 163, 168, 190). The longhorn crazy ant, P. longicornis, is an Old World tramp species spread through- out the tropics and subtropics by human commerce (Wetterer, 2008). Paratrechina longicornis is a ubiquitous agricultural and household pest throughout much of the tropics and subtropics and is a pervasive indoor pest in temperate areas. It thrives even in the most artificial environments, including ships at sea, facilitating its dispersal around the world (Wetterer, 2008). 29. Pheidole exigua Mayr Published Records. None. Specimens Examined. C&W, one site (B96, B109). JW, 23 sites (2003: 99, 107, 114, 119, 130, 131; 2006: 343, 346, 348-349, 358, 363, 376, 441, 450; 2014: 65, 107, 114, 118, 170, 176, 180, 185, 189). Pheidole exigua is a widespread New World species known from South America, Central America, and the West Indies (Wilson, 2003). We collected it in a range of habitats in Barbados. 30. Pheidole jelskii Mayr Published Records. No site data (1896; A. Forel; Forel, 1912b). No site data (1918; D. Stoner; Wheeler, 1923). No site data (1998; C&W; Wilson, 2003). Specimens Examined. Bridgetown (1920; W. M. Wheeler; MCZ). No site data (N. A. Weber; MCZ). C&W, two sites (B8, B23, 2016 ANTS OF BARBADOS (IIYMENOPTERA, FORMICIDAE) 15 B49). JW, 37 sites (2003; 89, 91, 92, 93, 95, 97, 99, 100, 103, 104, 110, 112, 121, 133; 2006; 338, 354, 371, 376, 381, 383, 387, 412, 418, 422, 441, 442, 446, 450, 451, 456; 2014; 9, 42, 56, 80, 95, 145, 165). Pheidole jelskii is a widespread New World species known from South America and the West Indies (Wilson, 2003). This large species is common in Barbados, with con- spicuous ground nests in open sunny areas. 30a. Pheidole fallax Mayr Published Records. No site data (Sirjusingh et al, 1992). In a review paper concerning biological control of weevils and whitegrubs, Sirjusingh et al. (1992) recorded P. fallax from Barba- dos. This was most likely a misidentification of P. jelskii, a common species that was first described as P. fallax jelskii. Wilson (2003, p. 290) wrote of P. fallax, “no certain records exist from the Lesser Antilles, where the closely related jelskii abounds.” 31. Pheidole megacephala (Fabricius) Published Records. Review (Wetterer, 2012e). Specimens Examined. JW, 28 sites (2003; 92, 95-96, 103, 109, 113-114, 115; 2006; 346- 347, 375, 376, 379, 389, 410, 451; 2014; 7, 42, 48, 54, 59, 83, 86, 90, 142, 143, 150, 152, 155, 157, 166). Although Wilson (2005) speculated that these are the pest ants reported in Barbados by Clarke (1670), the earliest published re- cords of P. megacephala in the New World come from St. Vincent (Forel, 1893) and the Bahamas (Emery, 1895). There are no published records of P. megacephala from Barbados. The African big-headed ant, P. megace- phala, an African native, is an important pest ant spread throughout the tropics and sub- tropics by human commerce (Wetterer, 2012e). Pheidole megacephala is well known as a household and agricultural pest. Because P. megacephala does not attack humans, this species is often not recognized as a sub- stantial threat. The negative ecological effect of P. megacephala, however, may be greater than any other invasive ant species. In areas where it occurs at high density, few native invertebrates persist. Loss of invertebrate species that serve key functions in the natural community (e.g., important prey species) may have cascading effects, leading to the subsequent loss of additional species. Phei- dole megacephala tends to thrive in open, disturbed habitats with weedy vegetation that can support high densities of plant- feeding Hemiptera, which these ants tend for honey dew. 32. Pheidole sculp tior Forel Published Records. None. Specimens Examined. C&W, four sites (B4, B28, B34, B39, B41, B43, B65, BlOO, B105). JW, 43 sites (2003; 87, 89, 93, 100, 104, 107; 2006; 358, 363, 365, 371, 376, 380, 387, 394, 395, 399, 440, 442, 446-447, 451, 453, 459; 2014; 27, 45, 61, 72, 74, 87, 95, 98, 107, 112, 118, 140, 152, 154, 156, 157, 169, 174, 180, 185, 190). Pheidole sculptior is widespread in the West Indies and northern South America (Wilson, 2003). It is common in a wide range of habitats in Barbados. 33. Pheidole subarmata Mayr Published Records. No site data (1896; A. Forel; Forel, 1912b). Specimens Examined. C&W, one site (B6). JW, nine sites (2003; 89, 99; 2006; 346, 442; 2014; 27, 87, 135, 146, 154). Pheidole subarmata is a widespread New World species known from South America, Central America, and the West Indies, common in disturbed habitats (Wilson, 2003). 34. Pheidole susannae Forel Published Records. None. Specimens Examined. JW, four sites (2006; 453; 2014; 27, 74, 183). 16 B REV I ORA No. 548 Pheidole siisannae is a widespread New World species, known from South America, Central America, and the West Indies, found in a wide variety of habitats (Wilson, 2003). 35. Pheidole teneriffana Forel Published Records. None. Specimens Examined. JW, two sites (2003: 115; 2006: 373). This Old World species was described from the Canary Islands, but Espadaler and Bernal (2003) considered this species an exotic there. It is also known from Sicily, Egypt, the Arabian Peninsula, Israel, Turkey, Malta, Greece, the Iberian Peninsula, California, Cuba, St. Helena, and Ascension Island (Wetterer, 2011c). Wilson (2003) proposed that this species may be native to the Medi- terranean. It does not appear to be very common in Barbados, found only at two highly disturbed sites. 36. Pheidole transversostriata Mayr Published Records. None. Specimens Examined. JW, 15 sites (2006: 343, 346, 349, 365, 367, 370, 390, 394, 397, 419, 442, 450, 453, 457, 459). Pheidole transversostriata is a widespread New World species, known from South America and the West Indies (Wilson, 2003). It is fairly common in Barbados, particularly at forested sites. 37. Plagiolepis alluaudi Emery Published Records. Review (Wetterer, 2014a). Specimens Examined. C&W, one site (B72). JW, 27 sites (2003: 91, 114, 117, 118; 2006: 357, 364, 379, 387, 390, 399; 2014: 6, 9, 44, 48, 56, 60, 61, 72, 74, 83, 85, 87, 107, 140, 153, 154, 157). This small orange ant is an African tramp species that has been spread around the world, particularly in the Pacific, through human commerce (Wetterer, 2014a). Kempf (1972) listed this species in the New World from Bermuda, St. Kitts, and St. Lucia. It also occurs on the Atlantic islands of Bermuda, Ascension, and St. Helena (Wet- terer et al, 2007). 38. Platythyrea punctata (Smith) Published Records. No site data (1896; Eorel, 1901). No site data (1918; D. Stoner; Wheeler, 1923). Turner’s Hall Woods (1995; Schilder et al., 1999). Specimens Examined. JW, 21 sites (2003: 87, 89, 94, 101, 122, 130; 2006: 358, 360, 364, 410, 442, 453; 2014: 28, 60, 71, 74, 118, 169, 177, 180, 186). Platythyrea punctata is a “circum- Caribbean” species found in a wide range of habitats (Brown, 1975). Seal et al. (2011) made genetic analyses of P. punctata speci- mens from a wide range of sites in the West Indies, Central America, Mexico, Texas, and Elorida. Analyses by Seal et al. (2011) in- dicated that populations of P. punctata in Central America are most basal and that populations in Mexico and Texas branched off long before this ant colonized the West Indies and Florida. Seal et al. (2011, p. 11) found that “phylogeographic analysis of P. punctata indicates a recent, one-way coloni- zation from Central America to the West Indies,” and concluded that it is “unlikely that P. punctata is a tramp species aided by human movements and commerce, because island haplotypes were never found on the mainland and only in one case was a mainland haplo- type found in the West Indies.” Seal et al. (201 1 ) estimated that P. punctata spread from Central America and through the West Indies about 150,000 years ago. 39. Pseudoponera stigma (Fabricius) Published Records. None. Specimens Examined. JW, one site (2014: 186). This species has an extensive range in both the New World and Old World tropics. Pseudoponera stigma shows characteristics of a native species throughout its range in both the New World and Old World, with near continuous distributions in intact forest 2016 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) 17 habitats. Wetterer (2012b) concluded that P. stigma most likely originated in tropical South and Central America. 40. Roger ia ciirvipuhens Emery Published Records. None. Specimens Examined. Orange Hill, in rotten wood (1987; J. P. E. C. Darlington; MCZ). C&W, one site (B19, B31). JW, 22 sites (2003: 97, 100, 109, 131; 2006: 343, 346, 349, 399, 410, 419, 447, 453, 457; 2014: 9, 61, 74, 98, 108, 118, 169, 174, 180). Roger ia curvipiibens is a widespread New World species known from northern South America, Central America, and the West Indies (Kugler, 1994; LaPolla and Sosa- Calvo, 2006). We found this species in a wide variety of habitats. 41. Rogeria foreli Emery Published Records. None. Specimens Examined. JW, 10 sites (2003: 97, 104, 109, 121; 2006: 363, 390, 447; 2014: 9, 140, 152). Rogeria foreli is a widespread New World species known from northern South Amer- ica, Central America, the West Indies, and the southwestern United States (Kugler, 1994; LaPolla and Sosa-Calvo, 2006). We found this species in a variety of habitats. 42. Solenopsis azteca Forel Published Records. Review (Pacheco and Mackay, 2013). Specimens Examined. JW, eight sites (2003: 87; 2006: 349, 363, 365, 395, 397, 419, 450). Thief ants are generally overlooked be- cause of their small size and primarily subterranean habits. Thief ants commonly persist at high densities in areas invaded by dominant exotic ants such as P. megacephala and L. humile (Wetterer et al., 2001). Despite the recent pioneering work on the taxonomy of thief ants (Pacheco and Mackay, 2013), much taxonomic work remains to be done on this challenging group. Solenopsis azteca has monomorphic work- ers that range in color from light yellow to light brown (Pacheco and Mackay, 2013). Solenopsis azteca is a New World species with a known range of “Honduras south to Colombia (Cauca) and northern Venezuela, Caribbean Region (Jamaica, Puerto Rico, Antilles)” (Pacheco and Mackay, 2013, p. 112). 43. Solenopsis geminata (Fabricius) Published Records. No site data (1896; A. Forel; Forel, 1912c). No site data (1922; H. A. Ballou; Wheeler, 1923). Bridgetown (1920; W. M. Wheeler; Wheeler, 1923). No site data (Sirjusingh et ai, 1992). Review (Wetterer, 201 la). Specimens Examined. C&W, one site (B93). JW, 38 sites (2003: 86, 89, 90, 91, 93, 97, 98-99, 104, 118, 133; 2006: 337, 338, 340, 343, 346, 357, 358, 363, 371-372, 373, 376- 377, 386, 387, 399, 412, 446-447, 450, 451, 456, 457; 2014: 12, 27, 74, 80, 81, 124, 147, 154). The tropical fire ant, S. geminata, is native to the New World, where in disturbed areas, it can reach very high densities and dominate the invertebrate community (Risch and Car- roll, 1982). It is also an invasive pest in many other parts of the world, including many tropical and subtropical areas where sea turtles nest, such as Australia, islands of the Pacific and Indian Oceans, the Arabian Peninsula, India, South Africa, Greece, and Cyprus (Wetterer, 2011a). 44. Solenopsis globularia (Smith) Published Records. Review (Pacheco and Mackay, 2013). Specimens Examined. C&W, three sites (B2, B56, B64, B70, Bill). JW, 28 sites (2003: 95, 97, 109, 112, 121, 133; 2006: 371, 375, 376, 381, 387, 410, 422, 442, 446-447, 449, 450, 456, 457; 2014: 6, 52, 56, 61, 85, 92, 154, 156, 157). Solenopsis globularia is a widespread New World species that ranges throughout the Caribbean to the Gulf Coast of the United States, through Mexico, to the central and northern part of South America 18 B RE VI ORA No. 548 (Brazil, Ecuador, Guiana) and the Galapagos Islands (Pacheco and Mackay, 2013). It also occurs on the South Atlantic islands of Ascension and St. Helena (Wetterer et al., 2007). 45. Solenopsis pollux Forel Published Records. Review (Pacheco and Mackay, 2013). Specimens Examined. JW, 13 sites (2003: 95, 110, 116, 121, 130; 2006: 337, 338, 346, 358, 371, 390, 451, 459; Pacheco and Mackay, 2013). Solenopsis pollux is a widespread New World species known from South America, Central America, and the West Indies (Pacheco and Mackay, 2013). 46. Solenopsis pygmaea Forel Published Records. Review (Pacheco and Mackay, 2013). Specimens Examined. JW, four sites (2003: 107; 2006: 364, 501, 528; Pacheco and Mackay, 2013). Solenopsis pygmaea is a widespread New World species known from Central America and the West Indies (Pacheco and Mackay, 2013). 47. Solenopsis zeteki Wheeler Published Records. Review (Pacheco and Mackay, 2013). Specimens Examined. JW, eight sites (2006: 380, 381, 395, 397, 399, 410, 440-441, 453). Solenopsis zeteki is a widespread New World species known from northern South America, Central America, and the West Indies (Pacheco and Mackay, 2013). 48. Strumigenys eggersi Emery Published Records. None. Specimens Examined. JW, five sites (2003: 130; 2006: 348-349, 453; 2014: 100, 102). Strumigenys are tiny predatory ants that feed on soil arthropods. All Strumigenys species are probably often overlooked be- cause of their small size and primarily sub- terranean habits. Strumigenys eggersi is a widespread New World species (Bolton, 2000). Brown (1959, p. 46) proposed that this species’ “home range is probably south Brazil and Bolivia, though a lack of collections from central and northern Brazil prevents us from knowing how far north this species extends. Strumigenys eggersi is also known from widely scattered localities in the West Indies, Florida, and southern Mexico . . . and it seems likely that it has been introduced by man at these many points.” Brown (1962, p. 249), however, seemed more unsure of the species’ exotic range, writing that S. eggersi was “wide- spread (possibly by recent introduction) in the West Indies; southern Florida; southern Mexico.” This species appears to be relatively un- common in Barbados. 49. Strumigenys emmae (Emery) Published Records. Review (Wetterer, 2012c). Specimens Examined. JW, four sites (2003: 104, 117; 2006: 379; 2014: 174). Strumigenys emmae is a common Old World tramp found in tropical regions worldwide (Wetterer, 2012c). Strumigenys emmae is one of three Old World dacetine ants, along with Strumigenys membranifera and Strumigenys rogeri, with widespread re- cords in both the Old World and the New World. 50. Strumigenys louisianae Roger Published Records. Review (Wetterer, 2014c) Specimens Examined. JW, seven sites (2006: 337, 450; 2014: 54, 108, 152, 154, 174). Strumigenys louisianae has the broadest geographic distribution of any New World Strumigenys, with an essentially continuous range from Argentina to North Carolina (Wetterer, 2014c). The widespread occurr- ence of S. louisianae in natural habitats across the West Indies suggests that S. louisianae is native throughout most or all of this region (Wetterer, 2014c). However, we found S. louisianae only at two highly 2016 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) 19 disturbed sites, suggesting that it may be exotic to Barbados. 51. Stnnuigenys rnargaritae (Forel) Published Records. Review (MacGown and Wetterer, 2013). Specimens Examined. C&W, two sites (B18, B21, B25). JW, 27 sites (2003; 87, 89, 107, 121, 130; 2006; 337, 343, 364, 367, 370, 380, 387, 395, 419, 446, 450, 453; 2014; 9, 103, 108, 118, 135, 140, 156, 169, 174, 185). Stnimigenys rnargaritae is a widespread New World species (MacGown and Wetterer, 2013). Although rare in many parts of its range, P. rnargaritae appears to have an essentially continuous circum-Caribbean dis- tribution from northern South America to the southeastern US and through the Lesser Antilles from Trinidad to Puerto Rico. If this distribution is truly continuous, it would support the hypothesis that P. rnargaritae, though generally considered exotic to the southeastern US, is actually native through- out its known range. Stnimigenys rnargaritae is commonly found associated with W. auropunctata, an aggressive and often dominant Neotropical species. Stnimigenys rnargaritae may derive protection from predators by associating with W. auropunctata. It is also possible that P. rnargaritae might afford some benefit to W. auropunctata. Strumigenys rnargaritae is common in a wide range of habitats in Barbados. 52. Strumigenys membranifera (Emery) Published Records. Review (Wetterer, 2011b). Specimens Examined. C&W, two sites (B67, B87). Strumigenys membranifera is a widespread tramp species of African origin (Wetterer, 2011b). Whereas S. rogeri and S. emmae are almost exclusively tropical, S. membranifera has spread to higher latitudes. Strumigenys membranifera is the only one of the three species with outdoor populations in Europe (e.g., S. membranifera is common in urban parks of southern Spain). In the continental United States, S. membranifera has been found in 12 southern states, whereas S. rogeri and S. emmae have outdoor populations only in peninsular Florida. Strumigenys membranifera occurs in a broad range of habitats, including dense forest, cultivated fields, urban lawns, and gardens. 53. Strumigenys rogeri Emery Published Records. Review (Wetterer, 2012a). Specimens Examined. C&W, two sites (B95, B96). JW, four sites (2003; 89; 2006; 354; 2014; 108, 180). Strumigenys rogeri apparently originated in tropical Africa, where its closest relatives all live, but it has become widespread on tropical islands of the Indo-Pacific and the West Indies and in peninsular Florida (Wetterer, 2012a). Outside of Africa and Florida, there are only a small number of continental records of S. rogeri, including a few from South and Central America and just one from continental Asia, in peninsular Malaysia. It is unclear whether S. rogeri has not yet spread to these continental areas, whether continental ants have competitively excluded S. rogeri, or whether these ants have been simply over- looked in surveys of diverse continental faunae. 54. Strumigenys sclmlzi (Emery) Published Records. None. Specimens Examined. C&W, one site (B36). JW, one site (2006; 363). Strumigenys sclmlzi was originally de- scribed from Brazil and is known from South America, Central America, and Trini- dad (Bolton, 2000). Our records of S. sclmlzi on Barbados are the only records of this species on a noncontinental island. 55. Syllophopsis sechellensis (Emery) Published Records. None. Specimens Examined. JW, two sites (2003; 130; 2014; 343). 20 B REV I ORA No. 548 Syllophopsis sechellensis (formerly Mono- morium sechellense) is native to tropical Asia but has spread by human commerce to many Old World sites. Wheeler (1923) recorded this species from Antigua. The present records are the only others known from the New World. This small yellow species is similar in appearance to SyUophop- sis subcoeca. 56. Syllophopsis subcoeca (Emery) Published Records. None. Specimens Examined. C&W, one site (B19). JW, eight sites (2003; 100, 104, 119; 2006: 354, 387, 443; 2014: 108, 135). We provisionally classify S', subcoeca (for- merly Monomorium subcoecum) as a New World species. All published records for S. subcoeca come from the Neotropics, where it has been considered native (Eernandez, 2007). All other members of Syllophopsis, however, are restricted to the Old World (except S. sechellensis', see above), and it is possible that S. subcoeca is actually an Old World species not yet collected in its native range. Syllophopsis subcoeca is similar in appearance to S. sechellensis', however, the mesopleuron is shiny and unsculptured in subcoeca, whereas it is matte and microretic- ulate in sechellensis. 57. Tapi noma melanocephcdum (Eabricius) Published Records. Review (Wetterer, 2009a). Specimens Examined. C&W, one site (B58). JW, 51 sites (2003: 91, 92, 93, 95, 97, 99, 108, 112, 114, 118, 119, 121; 2006: 337, 338, 354, 358, 364, 373, 376, 380, 383, 387, 390, 394, 410, 412, 418, 440, 441, 442, 447, 450; 2014: 27, 42, 51, 56, 83, 85, 90, 92, 93, 95, 136, 140, 144, 145, 156, 159, 162, 165, 190). The ghost ant, T. melanocephcdum, is a ubiquitous indoor and outdoor pest throughout much of the tropics and sub- tropics and a common indoor pest in temper- ate regions (Wetterer, 2009a). The close resemblance between T. melanocephcdum and a number of Indo-Pacific taxa indicates that T. melanocephcdum probably originated in this region. 58. Tetramorium bicarinatum (Nylander) Published Records. No site data (date unknown; collector unknown; Bolton, 1979). Review (Wetterer, 2009c). Specimens Examined. No site data (1936; E. A. Chapin and R. E. Blackwelder; SI). (1987; J. P. E. C. Darlington; MCZ). JW, one site (2014: 90). Bolton (1979) wrote that he examined T. bicarinatum specimens from Barbados, citing Bolton (1977), but no records from Barbados were listed in the earlier paper. The penny ant, T. bicarinatum, is an Old World tramp species spread throughout the tropics and subtropics by human commerce (Wetterer, 2009c). 59. Tetramorium caldarium (Roger) Published Records. None. Specimens Examined. C&W, one site (BllO). JW, eight sites (2003: 92, 109, 112, 114; 2006: 376, 381; 2014: 83, 165). Tetramorium caldarium is a tramp ant species originally from Africa that has dispersed around the world through human commerce. Erom 1862 to 1979, T. caldarium was considered a junior synonym of Tetra- morium simillimum (Smith), but Bolton (1979) revived T. caldarium as a species. Tetramorium caldarium records are particu- larly common on Atlantic islands and from greenhouses and heated buildings in temper- ate Europe. We found this species only in highly disturbed habitats in Barbados. 60. Tetramorium lanuginosum Mayr Published Records. No site data (^1893; W. G. Jeffreys; Wheeler, 1923). Review (Wetterer, 2010b). Specimens Examined. JW, five sites (2006: 376, 387; 2014: 151, 152, 157). Tetramorium lanuginosum is considered a pantropical tramp (Wetterer, 2010b). Although we found this species at only two 2016 ANTS OF BARBADOS (1 lYMENOPTERA, FORMICIDAE) 21 sites, it was very common in grassy areas ot the University of the West Indies campus. 61. Tetramoriiim similUmuw (Smith) Published Records. None. Specimens Examined. C&W, two sites (BIO, B77). JW, 27 sites (2003; 86, 92, 93, 112, 114, 115, 117, 119, 121, 133; 2006: 337, 376, 381, 383, 385, 387, 418, 422, 441, 442, 456; 2014; 54, 144, 154, 157, 160, 190). This species is a common and widespread Old World tramp species that has spread worldwide through hriman commerce (Bolton, 1977). Concerning T. caldariiim and T. simi- llimum, Bolton (1979, p. 169) wrote: “Of the two there is no doubt that simillimum is the most successful. It has been widely recorded throughout the tropics and subtropics and also occurs fairly frequently in the temperate zones in zoological and botanical gardens and in conservatories and other constantly heated buildings.” We found this species at many highly disturbed sites in Barbados. 62. Trichomyrmex destructor (Jerdon) Published Records. No site data (Tucker, 1953). Review (Wetterer, 2009b). Specimens Examined. JW, 11 sites (2003: 86, 115, 117, 119, 133; 2006: 371, 374, 381, 416, 450; 2014; 42). The destroyer ant, T. destructor (formerly Monomorium destructor) is an Old World tramp species that is a pest in many tropical and subtropical areas (Wetterer, 2009b). It is notorious for living in electrical equipment, chewing through the insulation of electrical wires, and attacking people. 63. Wasmannia auropunctata (Roger) Published Records. No site data (1896; A. Forel; Forel, 1912c). Coles Cave (1979; S. B. Peck; Peck, 1981). Review (Wetterer, 2013a). Specimens Examined. C&W, two sites (B38, B85, B91). JW, 49 sites (2003: 89, 93, 99, 104, 107, 110, 1 14, 122, 130; 2006: 338, 343, 346, 349, 354, 364, 370, 387, 390, 394, 395, 399, 419, 422, 442, 446-447, 450, 457, 459; 20 14:6, 8, 44, 60, 64, 74, 87,95, 98, 100, 102, 107, 111, 113, 121, 135, 145, 158, 169, 185, 189). The little fire ant, W. auropunctata, occurs throughout most of the warmer parts of the New World, from subtropical Argentina to subtropical Mexico and through much of the West Indies, although it is not clear whether this species in native to this entire region. During the past century, exotic populations of W. auropunctata have become established in numerous other places, includ- ing the Galapagos Islands, West Africa (Gabon, Cameroon, and possibly the Re- public of Congo and the Democratic Republic of Congo), Melanesia (New Caledonia, Solo- mon Islands, Vanuatu, and possibly Tuvalu), Polynesia (Wallis and Futuna and Hawaii), the mainland United States (Florida and possibly California), and subtropical Atlantic islands (the Bahamas and Bermuda). In many areas, W. auropunctata can be a significant agricultural pest, not only stinging agricultur- al workers, but also enhancing populations of Hemiptera. Hemiptera cause damage both through sapping plants of nutrients and by increasing the occurrence of diseases, includ- ing viral and fungal infections. Additionally, W. auropunctata has a negative effect on many animals, both invertebrates and vertebrates, although most reports on such effects have been anecdotal. The effects of W. auropunc- tata populations seem to be most severe on tropical islands where it is not native, such as the Galapagos, New Caledonia, and the Solomon Islands. Reports of widespread blindness in both domestic and native mam- mals caused by W. auropunctata stings de- serve serious attention (Wetterer, 2013a). SPECIES WITH NO RECENT RECORDS (POSSIBLY EXTINCT) 64. Atta cephalotes lutea Eorel (stat. rev.) Published Records. No site data (<1893; W. G. Jeffreys; Eorel, 1893; type locale). Specimens Examined. None. 77 BREVIORA No. 548 This species was collected only once. Forel (1893) described A. lutea Forel from Barba- dos but later designated it a subspecies of colombica (Forel, 1908), a classification fol- lowed by Emery (1913). Santschi (1929) revived its status as a species and listed it in Trinidad. Gongalves (1942) listed it as a species and was told by R. W. E. Tucker that it did not occur in Barbados. Borgmeier (1950) repeated its status as a species. Weber (1958, p. 10) designated it a subspecies of cephalotes, writing “Contrary to the alloca- tions of this as a separate species by more recent workers, it appears to be no more than a subspecies of cephalotes. It would seem to be a mutant developed from a sub- species of cephalotes on the pale coral sand of this small island. Perhaps it was intro- duced as typical cephalotes by way of the abundant shipping between Barbados and Trinidad and Demerara several centuries ago. I collected typical cephalotes in all parts of Trinidad and do not believe that lutea occurs there.” Borgmeier (1959, p. 344) wrote (in German), '"Atta lutea Eor. 1893 (Barbados, Jeffreys leg.) was described as a species. The description fits exactly with uncolored specimens of cepha- lotes (seen in the shiny gaster of the smallest workers); Eorel mentions ‘a bunch of hairs a little more abundant of each side of the vertex’, which fits only cephalotes. One hardly understands therefore that Eorel (1913) and Emery (1913, 1923) place lutea as a variety of colombica. Santschi (1929) brought new confusion in that he considered light yellow specimens with shinier forehead of Trinidad as lutea, and Gonsalves (1942) and Borgmeier (1950) repeated this; but Forel says specifi- cally that his animals were dull; only for the 3 mm worker does he give the head as subopaque and the gaster as ‘very shiny’. Of geographic variation, therefore there can be nothing said. The light yellow coloiing alone can establish no subspecies. Such light yellow workers were gathered by Weber himself on Trinidad (1934, 20 XII., no. 33); it appears in the same colony together with normally colored animals and therefore has no nomenclatural value at all.” Weber (1958), however, explicitly stated that he did not believe lutea occurred in Trinidad; thus, “light yellow coloring alone” clearly was not Weber’s (1958) basis for differen- tiating lutea. Weber (1972) resurrected lutea as a sub- species of cephalotes. Bolton (1995) listed lutea as a junior synonym based on Borgme- ier (1959). Borgmeier (1959), however, made this synonymy based on the written description, not through examining type specimens. We therefore chose to follow Weber (1972), the more recent taxonomic designation. The taxonomy of this form deserves further study. This record is the only record for any Atta species from Lesser Antilles (except Trinidad and Tobago), and this may be an exotic to Barbados, as Weber (1958) suggested. 65. Crematogaster brevidentata Eorel Published Records. Bathsheba, 200 m (1896; A. Eorel; Eorel, 1912b; type locale). Specimens Examined. None. Eorel (1912b) described C. brevispinosus brevidentata from Barbados. Longino (2003) examined a syntype worker in the Musee dHistoire Naturelle Geneve and raised bre- videntata to full species, known only from Barbados. Longino (2003, p. 131) wrote: “workers are small, propodeal spines are tiny, there are no erect setae on head or fourth abdominal tergite, erect setae are sparse on mesosoma, and there is a strong anteroventral petiolar tooth.” Several described subspecies of C. brevispi- nosa were later determined to be synonyms of C. crinosa. Of the five New World ant species that we collected, most often in Barbados, C. crinosa is the only one that Eorel did not report among the ants he collected in 1896. It seems likely that either 2016 ANTS OF BARBADOS (IIYMENOPTERA, FORMICIDAE) 23 C hrevidentata is conspecific with C. crinosa or that C hrevidentata has been driven extinct by C crinosa. 66. Crematogaster carinata Mayr Published Records. Bridgetown (1920; W. M. Wheeler; Wheeler, 1923). Specimens Examined. None. Longino (2003) examined specimens of this New World species from many parts of South and Central America. Wheeler’s (1923) specimens, not examined by Longino (2003), are the only record of this species from the West Indies. This arboreal species was collected only in Bridgetown, the capital and main port city of Barbados, suggesting that this species may be exotic to Barbados. Wheeler (1923) may have sampled a tempo- rary population of this species, brought to Barbados on lumber from South America. 67. Crematogaster steinheili Forel Published Records. No site data (1896; A. Forel; Forel, 1912b). Specimens Examined. None. This yellow arboreal ant is a strictly Antil- lean species. It is very common in the Greater Antilles and the Bahamas and has been introduced to Florida. In the Lesser Antilles, it is otherwise known only from a few sites in Antigua (JKW, unpublished data), the near- est site records to the single Barbados site record. Crematogaster steinheili is the only New World ant species in Barbados that is known from Florida or the Bahamas, but not from Trinidad or South America. Thus, it seems likely that C. steinheili is not native to Barbados. Forel (1912b) may have sampled a temporary population of this species, brought to Barbados on lumber from the Greater Antilles. 68. Pseudomyrmex gracilis (Fabricius) Published Records. No site data (2w; <1877; no date; no collector; BMNH; lectotype and paralectotype of P. pilo.sula Smith; Smith 1877; designation by Ward, 1993: 155). No site data (Iw; no date; no collector; BMNH; lectotype of P. variahilis Smith; designation by Ward, 1989: 439). Review (Wetterer, 2010c). Specimens Examined. None. In the earliest ant records from Barbados with extant specimens. Smith ( 1 877 ) described P. pilosula Smith and P. variahilis Smith [both species = P. gracilis (Fabricius)]. It has never been found since. This species is widespread through much of the New World and could be exotic (Wetterer, 2010c), brought to Barbados on lumber from South America. It has been introduced to the southeastern United States and Hawaii (Wetterer, 2010c; MacGown et al., 2013). 69. Pseudomyrmex maculatus (Smith) Published Records. “Barbadoes” (Iw; <1877; no collector; BMNH; Ward, 1989). Specimens Examined. None. Ward (1989) found that a specimen of P. maculatus (Smith) was on the same pin as Smith’s P. variabilis specimen that was labeled “Barbadoes.” Ward (1992) also listed this species from Barbados. Smith (1855) described P. maculatus from Brazil. The specimens of P. gracilis and P. macu- latus are of unknown age but must predate Smith (1877). This species could be exotic, brought to Barbados on lumber from South America. DISCUSSION In preliminary biogeographic analyses of West Indian ants, Wilson (1988) reported 18 ant species from Barbados. Here, we docu- mented 69 ant taxa collected in Barbados (46 New World and 23 Old World). Preliminary analyses (Table 3) indicate that there are fewer New World and more Old World ant species recorded from Barbados than from the two Lesser Antilles islands most similar in size to Barbados: St. Vincent and Grenada (Table 3). 24 BREVIORA No. 548 Table 3. Ant species richness of Barbados and NEIGHBORING ISLANDS.* Area (km-) Elev (m) Total NW OW Barbados 462 336 69 46 23 St Vincent^^ 381 1,234 70 58 12 Grenada^ 344 840 74 61 13 *Elev = maximum elevation, NW= New World species, OW= Old World species. ^St Vincent and Grenada data from Wetterer et al. (unpublished data). Only four ant taxa recorded in Barbados are known solely from the West Indies (Table 1 ). Most of the New World ant species we recorded have very broad ranges (Table 1). Of the New World species found in Barbados, 41 also occur in mainland South America, but only 17 are known from Florida. Several of these New World species have been found at a few sites in the Old World (e.g., A. mayri, B. cf. heeri, C. atriceps, N. cf. steinheili, G. striatula in European greenhouses). However, only two species, S. geminata and W. auropimctata, are tramp species that have spread world- wide via human commerce (Table 1). There- fore, it seems likely that most of the New World ants in Barbados are probably native and originated in South America. Some New World species, however, could be exotic to Barbados (e.g., C. steinheili and N. cf. fulva; see Species Accounts), and others could have a mix of both native and exotic populations. For example, one possibility is that light-colored C. sexguttatus grenaden- sis (= C. sexguttatus), known only from Barbados, Grenada, and St. Vincent, is native to Barbados, whereas the widespread typical form of C. sexguttatus is exotic. All 23 Old World ant species found in Barbados have broad worldwide distribu- tions and are certainly exotic to Barbados. These included some major pest species, such as P. megacephala, T. melanocephalum, and T. destructor. Of these 23 species, 18 are also known from South America, 16 from Trini- dad, and 19 from Florida. Possible ant extinctions in Barbados A number of ant species on Barbados may have been driven extinct through human activities, some lost without a trace. By the time of the earliest ant collection in Barbados in the 19th century, little remained of the original vegetation (Forel, 1912b). Six ant species, all of New World origin, are known from Barbados based solely on single collection records >80 years old (Tables 1, 2; see Species Accounts). Four of these ants are widespread arboreal species; C carinata, C. steinheili, P. gracilis, and P. macidatus. These species may have been incidentally extirpated due to widespread deforestation in Barbados. Alternatively, these arboreal ants may have been only temporary populations introduced on im- ported lumber (see Species Accounts). The other two species with no recent records from Barbados are the only apparently endemic taxa: A. cephalotes lutea and C. brevidentata. Jeffreys collected the only known speci- mens of A. cephalotes lutea in Barbados sometime before 1893 (Forel, 1893). Else- where in the New World, colonies of other A. cephalotes subspecies can grow to include several million workers. These ants are highly conspicuous and destructive pests. It seems unlikely that A. cephalotes lutea colonies have survived to the present day undetected in Barbados. This ant may have been purposely exterminated in Barbados as an agricultural pest. Forel (1912b) made the only known collection of the acrobat ant C. breviden- tata at a site 200 m above the coastal town of Bathsheba. The area above Bath- sheba is the site of the largest remaining forest in Barbados, on the slopes of 2016 ANTS OF BARBADOS (HYMENOPTERA, FORMICIDAE) 25 Hackleton’s Cliff. Although we collected extensively in this area, we did not find C. hrevidentata. Future research Building upon the present study on the ants of Barbados, we plan to complete our inventory of the ants of neighboring Grenada and St. Vincent, the southern- most major islands of the Lesser Antilles (Fig. 1). Grenada and St. Vincent differ from Barbados in many ways that can influence biodiversity. Grenada and St. Vincent are much older than Barbados, having first emerged ~ 50 mya, and some researchers have proposed that they were once attached to South America. During low sea levels 15 kya, Grenada was joined with all the Grenadine islands to the north and with an extension to the south, forming an island about ten times Grenada’s current size that was separated from St. Vincent by just a few kilometers and from South America by ~ 70 km (Fig. 1 ). Grenada and St. Vincent are also much more mountainous than Barbados. Our present inventory can serve as a base- line for ant research in Barbados. We collected several ant species at only single locales, and we expect that additional collect- ing in Barbados will find additional species present. We also expect that additional exotic ant species will continue to invade Barbados. For example, the highly destructive red imported fire ant, Solenopsis invicta, original- ly from South America, is now spreading across the West Indies (Wetterer, 2013b) but has not yet been recorded from Barbados. About 30 Old World ant species are known from the West Indies. Of these, major Old World tramp species that have not yet been recorded from Barbados include Mono- moriwn pharaonis and Technomyrmex diffi- cilis. Destructive exotic ants tend to thrive in open, disturbed habitats with weedy vegeta- tion that can support high densities of plant- feeding Hemiptera, which these ants tend for honeydew. This is one reason to encourage the strict protection of what little forest remains in Barbados. ACKNOWLEDGMENTS We thank M. Wetterer and S. Shattuck for comments on this manuscript; S. P. Cover, M. Deyrup, J. T. Longino, L. R. Davis, and R. R. Snelling for assistance in ant identifi- cation; and the National Science Foundation (DEB-0515648) and Elorida Atlantic Uni- versity for financial support. APPENDIX 1. EARLY REPORTS PROM BARBADOS CONCERNING ANTS OP UNCERTAIN SPECIES IDENTITIES Clarke (1670: pp. 70-7 1 ) presented the earliest account of ants in Barbados, writing, “They have ants and pismires of a small size, but of a great industry; they are every where, in hollow ground amongst the root of Trees, upon the Bodie, Branches, Leaves, and Fruit of all Trees; without houses, within their houses, upon their sides, walls, windows, Roofs, Tables, Cupboards, Stools, Beds. Floors, all within and without are covered with them. When they find a dead Cockroch, though he be bigger then a hundred of them, yet they will take hold of him and lift him up, and away they carry him, some going by to assist those that are weary; some (like Officers) lead the way to shew the hole into which he must pass, and if his body do lie a cross that it cannot go into the hole, they give notice to the carriers, that presently turn his body endwise before it come to the hole, and that without any stop, and they never pull contrary wayes. The Planters which are so curious to prevent their coming upon their Tables, Cupboards, and Beds, have little troughs filled with water for the feet of these to stand in; yet all will not prevail, for they will get in the scieling, and so fall down upon them. To keep them from the shelves on which their meat stands, they are forced to hang them to the roof with ropes, and to tar those ropes and the roofs over them. When a Carpet upon a Table is covered over with them, if you kill many, and let them alone but a while, they will carry away all the dead ones. If you set Sugar upon a Table which you have first freed from them, some in the 26 BREVIORA No. 548 room will presently smell it, and make towards it as fast as they can, and having found it, return again without medling with it, and gives notice of this booty, and then they come in thousands and ten thousands, and in an instant fetch it all away so that there is no place safe from these over-busie Creatures.” Wilson (2005, p. 32) speculated on the identity of the ants reported by Clarke (1670), writing, “On Barbados in the mid- 1600s there was an ant that was a serious house pest. Unlike fire ants, its workers lifted and carried large food items, such as cockroaches, in an unusually coordinated fashion. This feature points to the ant genus Pheulole. Among the many species known from the West Indies, only two are candidates: P. jelskii, a native species, and P. megacephalci, which is of African origin. The evidence favours P. megacephalci, a global, invasive ant that has caused similar problems in other tropical countries.” Pheulole, however, have distinctly dimorphic workers, a trait not mentioned by Clarke (1670). An alternative possibility is P. longicomis, whose workers often carry large prey in a coordinated group. Hughes (1750, pp. 93-94) wrote about four species of ants in Barbados: 1. “The Small Red Ant. This is a very small ant: Yet the Part of the Skin it bites continues painful for near Four Hours afterwards. If these are likewise killed, and rubbed upon the Skin, they raise a Blister. The Bodies of these Ants are thickly covered with sharp fine-pointed Bristles, imper- ceptible to the naked Eye.” 2. “The Stinging Ant. This appears to be the same with what is to be seen in England in the Summer Season in most Pasture-Lands.” 3. “The Horse-Ant. This is the largest-sized Ant, and is often to be met with both within and without Doors; and hath nothing peculiar in its Make or Qualities.” 4. “The Sugar-Ant. This is a small whitish Ant, very fond of Sugar, or any sweet and oily Liquids; and consequently very troublesome to the Housewives, it being difficult to keep them from every kind of Victuals.” Judging from Hughes’ ( 1 750) depictions, it seems likely that the “small red ant” is the little fire ant W. aiiropimctata, the “stinging ant” is the tropical fire ant 5. geminala (Wheeler, 1926; Wilson, 2005), the “horse- ant” is the enormous carpenter ant C. atriceps, and the “sugar-ant” is the ghost ant T. melanocephaliim (Wheeler, 1926). Hughes (1750) also mentioned a “Wood-Ant” and a “Great-headed Ant.” Hughes (1750) described the Wood-Ant as “very destructive to Timber” and “very expeditious in building their Nests, which are long hollow T ubes, the Outside being an Incrustation of a gritty clayey Matter.” Hughes ( 1 750) continued that the Great-Headed Ant “no-way differs from the last described [the Wood- Ant], but by the Bigness of its Head.” The description of long tubes of clay indicates that these last two “ants” are actually worker and soldier termites. Castles (1790) wrote extensively about a population explosion of “sugar ants” that devastated Grenada beginning around 1770 but largely receded after a hurricane in 1780. Castles (1790) mentioned that Barbados once experienced the bad effects of these same ants but had been freed of this problem. In a chapter on the history of Martinique, Coke (1808-11) wrote about a highly destructive “sugar-ant” that devastated Martinique between 1763 and 1766 and again around 1770. Coke (1808-1 1 ) also mentioned that “some time before they appeared in Martinico, they ravaged Barbadoes to such a degree, that it was deliberated, whether that island, formerly so flourishing, should not be deserted” (Coke, 1808-11; vol 2: 313). Wheeler (1926) and Wilson (2005) concluded that this 18th century plague ant was probably S. geminala, which may have had a population explosion that was associated with the introduction of an exotic plant- feeding Hemiptera mutualist. In his “History of Barbados,” Schomburgk (1848) devoted four pages to pest ants. Schomburgk (1848) discussed four ant species, which he identified as Formica omnivora, Formica cephalotes, Formica caiistica, and Formica saccharivora. These four species may or may not correspond to 5. geminala, Alla cephaloles, W. auropimclala, and T. melanocephaliim, although it is not certain. Most of the Schomburgk (1848) information on ants came from New World locales other than Barbados, and much of it appears to involve confusion of different ant and termite species. Schomburgk (1848, p. 640) described the ravages of “the Sugar Ant. Formica omnivora, Linn., Myrmica omnivora, Latr.,” drawing heavily on the writings of Castles (1790) concerning Grenada and Coke (1808-11) concerning Martinique. Schomburgk (1848, p. 640) stated these ants “showed themselves about 1760 in Barbados, and caused such devastation that ‘it was deliberated, whether that island, formerly so flourishing, should not be deserted’.” Schomburgk (1848) further wrote, “It has been asserted that this insect had been brought to Barbados from Tobago in some mould imported by Mr. Gidney Clarke, owner of the Bell Plantation. In 1814 they again made their appearance with considerable injury in many districts to the vege- table productions and feathered stock, but they did not continue long,” citing “Dr. Collyns’s MSS” as his source of information. Schomburgk (1848, p. 643) 2016 ANTS OF BARBADOS (I lYMENOPTERA, FORMICIDAE) 27 continued: “They are still to be found in Barbados, but only in small numbers.” As mentioned above, Wheeler (1926) and Wilson (2005) concluded that F. oninivuru was probably S. geminatci. Schomburgk (1848, p. 643) then wrote about the “Cushi or Great-headed ant,” which he identified as F. cephalotes (= A. cepluiloies), a species of the leaf-cutting ant that is a major pest in South and Central America. But Schomburgk’s (1848) own observations on this ant all appear to come from Guyana in South America, and his mention of the presence of this ant in Barbados may be based on Flughes’ (1750) mention of a “Great-headed Ant,” which actually referred to a termite. Schomburgk (1848) also noted a small red ant, citing (and paraphrasing) Flughes’ (1750) account, which appears to refer to W. aiiropimctuta. But then Schomburgk (1848, p. 644) added, “the ant is described by Kollar as Formica caiistica." Later authors have identified F. camtica as either Cephalotes mimitus (Fabricius) (Forel, 1895) or Cephalotes piisillus (Klug) (Kempf, 1951). Neither of these species is small or red or stings, and no Cephalotes is known from Barbados. Finally, Schomburgk (1848, p. 644) wrote, “the Sugar Ant, which will creep through the smallest crevices in order to get to the sugar, is Formica saccharivora.” This second “Sugar Ant” is clearly different from the F. omnivora “sugar ant” he discussed earlier. Although modem taxonomists have been unable to determine the identity of F. saccharivora, Linnaeus ( 1758) described it as an American species the size of Formica caespitum ( = Tetramorium caespitum), the large “pavement ant” of Europe. Such a large ant would not be able to “creep through the smallest crevices.” Instead, the Schomburgk ( 1 848) account of this second sugar ant appears similar to that of Hughes’ (1750) sugar ant; both seem to fit T. melanocephalum (also see Wheeler, 1926). APPENDIX 2. COLLECTION SITE INFORMATION Information given as vial nos.; site; degrees N, degrees W (date). Cover and Wilson, 1998 (six sites) Bl-18, 55-56, 105-106, 110-113: Casuarina Beach Club; 13.07, 59.57 (17, 18, and 21 Mar 1998) B19-54: Turner’s Hall Woods; 13.23, 59.58 (18 Mar 1998) B57-65, 76-77: Graeme Hall Swamp; 13.07, 59.58 (19 Mar 1998) B66-75: Barbados Wildlife Reserve; 13.27, 59.60 (19 Mar 1998) B78-94: Jack-in-the-Box; 13.17, 59.58 (20 Mar 1998) B95-104, 107-109: Hackleton’s Cliff Forest; 13.20, 59.53 (20 Mar 1998) Wetterer, 2003 (27 sites) 86 and 1 17: Black Rock, by apartment; 1 3. 1 26, 59.634 (25 and 28 Nov 2003) 87-88: Hopewell House, gully forest; 13.166, 59.587 (25 Nov 2003) 89-90: Harrison’s Cave, forest; 13.178, 59.579 (25 Nov 2003) 91: Edghill, weeds by field; 13.151, 59.604 (25 Nov 2003) 92; Holetown, beach; 13.183, 59.644 (26 Nov 2003) 93-94: Speightstown, vacant lot; 13.229, 59.648 (26 Nov 2003) 95-96: Crab Hill, sugar; 13.309, 59.646 (26 Nov 2003) 97; Cluffs, scmb; 13.323, 59.629 (26 Nov 2003) 98-99: Farley Hill, sugar and weeds; 13.262, 59.601 (26 Nov 2003) 100-102: Turner’s Hall Woods, secondary forest; 13.225, 59.583 (26 Nov 2003) 103: Proutes, sugar; 13.160, 59.581 (27 Nov 2003) 104-106; Pool, forest patch; 13.174, 59.516 (27 Nov 2003) 107: Hackleton’s Cliff, forest; 13.199, 59.528 (27 Nov 2003) 108: Bathsheba, seagrape; 13.214, 59.528 (27 Nov 2003) 109: Barclay’s Park, seagrape; 13.230, 59.545 (27 Nov 2003) 110-111: Welshman Hall Gully, groomed forest; 13.191, 59.581 (27 Nov 2003) 112: East Point, grass and weeds; 13.161, 59.435 (28 Nov 2003) 113-114: King George V Park, trees and grass; 13.130, 59.466 (28 Nov 2003) 115; Oistins, beach; 13.064, 59.552 (28 Nov 2003) 116: Bridgetown, post office garden; 13.095, 59.626 (28 Nov 2003) 118: The Valley, sugar; 13.118, 59.570 (28 Nov 2003) 119-120: Brighton, base of trees by field; 13.121, 59.533 (28 Nov 2003) 121: Hopewell, flower garden; 13.175, 59.594 (29 Nov 2003) 122-129: Welshman Hall Gully, forest by ball fields; 13.185, 59.585 (29 Nov 2003) 130: Flower Forest, garden; 13.202, 59.572 (29 Nov 2003) 28 B REV I ORA No. 548 131-132: Flower Forest, forest; 13.199, 59.570 (29 Nov 2003) 133: Seawell, airport parking; 13.078, 59.493 (29 Nov 2003) Wetterer, 2006 (41 sites) 337: Black Rock, compost by hotel; 13.126, 59.634 (16 June 2006) 338-342: Edge Hill, by cricket field; 13.152, 59.602 (16 June 2006) 343-345: Hopewell House, gully forest; 13.166, 59.587 (16 June 2006) 346-347: Canefield, forest patch; 13.193, 59.590 (16 June 2006) 348-353: Boarded Hall, forest; 13.209, 59.576 (16 June 2006) 354-356: Breedy’s, scrub forest; 13.251, 59.590 (16 June 2006) 357: Hackleton’s Cliff, base forest; 13.203, 59.536 (16 June 2006) 358: Hackleton’s Cliff, sugarcane above cliff; 13.198, 59.527 (17 June 2006) 359-363: Hackleton’s Cliff, trees at top edge; 13.197, 59.525 (17 June 2006) 364: Hackleton’s Cliff, grass above cliff; 13.197, 59.524 (17 June 2006) 365-366: Hackleton’s Cliff, forest at bottom; 13.202, 59.533 (17 June 2006) 367: Jericho, grass; 13.183, 59.587 (17 June 2006) 370: Welshman Hall, gully forest; 13.163, 59.564 (17 June 2006) 371-372: Fitts, beach picnic area; 13.145, 59.645 (17 June 2006) 373: Bridgetown, Cavans Lane; 13.093, 59.62 (18 June 2006) 374: Bridgetown, Carlisle Wharf; 13.093, 59.623 (18 June 2006) 375: Bridgetown, St. Mary’s Church; 13.096, 59.623 ( 1 8 June 2006) 376-378: Bridgetown, Queen’s Park; 13.096, 59.615 ( 1 8 June 2006) 379: Marine Gardens, by apartments; 13.072, 59.598 ( 1 8 June 2006) 380: Gemswick, trees by grass; 13.086, 59.478 (18 June 2006) 381-382: Foul Bay, beach; 13.095, 59.455 (18 June 2006) 383-386: Cheapside, by cruise terminal; 13.097, 59.634 (18 June 2006) 387-389: Cave Hill, UWI campus; 13.133, 59.636 (19 June 2006) 390-392: Bennett’s, trees by Route 2A; 13.170, 59.617 (19 June 2006) 393-394: Ridgeway, gully forest; 13.186, 59.617 (20 June 2006) 395-398: Plumtree, roadside forest; 13.195, 59.619 (20 June 2006) 399-409, 423-439: Rockless, gully forest; 13.250, 59.630 (20 June 2006) 410-411: Trents, vine-covered trees; 13.298, 59.633 (20 June 2006) 412-417: River Bay, waterfront park; 13.317, 59.600 (20 June 2006) 418: Diamond Corner, by ball field; 13.272, 59.603 (20 June 2006) 419-421: Moore Hill, St. Nicholas Abbey; 13.273, 59.597 (20 June 2006) 422: Lakes Beach, beach; 13.245, 59.556 (20 June 2006) 440-441: Bath, near route H3; 13.182, 59.482(21 June 2006) 442-445: Bath, near beach; 13.185, 59.480 (21 June 2006) 446-449: Codrington College, campus; 13.172, 59.479 (21 June 2006) 450: Sweet Vale, Orchid World; 13.159, 59.555 (21 June 2006) 451-452: Belleplaine, Sport Club; 13.243, 59.564 (22 June 2006) 453-455: Walkers, gully forest; 13.242, 59.576 (22 June 2006) 456: Gregg Farm, store at crossroad; 13.215, 59.592 (22 June 2006) 457-458: Hoytes, secondary forest; 13.230, 59.588 (22 June 2006) 459-461: Jack-in-the-Box, gully forest; 13.168, 59.582 (22 June 2006) Wetterer, 2014 (55 sites) 6-7: Holetown, by apartment; 13.180, 59.635 (2 May 2014) 8-25: Hillcrest, Andromeda Garden; 13.210, 59.518 (3 May 2014) 26: Bathsheba, waterfront park; 13.212, 59.518 (3 May 2014) 27-41: Hackleton’s Cliff, 0.6 km E highway; 13.205, 59.529 (3 May 2014) 42-43: Fitts, waterfront park; 13.146, 59.638 (4 May 2014) 2016 ANTS OF BARBADOS (I lYMENOPTERA, FORMICIDAE) 29 44-47: Eazaretto, park scrub; 13.134, 59.634 (4 May 2014) 48-51: Brandons Beach, park; 13.117, 59.627 (4 May 2014) 52-53; Cheapside, by restaurant; 13.101, 59.624 (4 May 2014) 54: Bridgetown, by restaurant; 13.096, 59.618 (4 May 2014) 55: Bridgetown, Jubilee Gardens; 13.098, 59.618 (4 May 2014) 56-58; Carrington, by ball field; 13.105, 59.605 (4 May 2014) 59: Bridgetown, Independence Square; 13.105, 59.605 (4 May 2014) 60: Trents, scrub; 13.194, 59.629 (5 May 2014) 61-63: Rock Hall, forest; 13.191, 59.610 (5 May 2014) 64-70: Mt. Hillaby, N side forest; 13.212, 59.582 (5 May 2014) 71-73; Mt. Hillaby, S side forest; 13.211, 59.582 (5 May 2014) 74-78; Melvin Hill, forest edge; 13.201, 59.564 (5 May 2014) 80: Ape Hill, golf course; 13.22, 59.60 (6 May 2014) 81-82: Ape Hill, by houses; 13.22, 59.60 (6 May 2014) 83-84; Folkestone, Bellairs Station; 13.192, 59.640 (6 May 2014) 85-86: Holetown, mall; 13.185, 59.638 (6 May 2014) 87-89; Worthing, preserve entrance; 13.071, 59.578 (7 May 2014) 90-91: Worthing, swamp trail; 13.071, 59.577 (7 May 2014) 92: South Point, scrub; 13.047, 59.530 (7 May 2014) 93; Enterprise, scrub; 13.058, 59.538 (7 May 2014) 94: Jackmans, scrub; 13.139, 59.594 (7 May 2014) 95-96: Hackleton, scrub; 13.199, 59.518 (8 May 2014) 98-99: Hackleton’s Cliff, by water tank; 13.205, 59.530 (8 May 2014) 100-101; Hackleton’s Cliff, 750 m E of highway; 13.204, 59.529 (8 May 2014) 102-105: Hackleton’s Cliff, 900 m E of highway; 13.203, 59.527 (8 May 2014) 107-111: Hunte’s Garden, garden; 13.192, 59.551, 8 and 9 May 2014) 112-117 and 126-134: Turner’s Hall Woods, SW forest by ghut; 13.221, 59.584 (9 May 2014) 118-123: Turner’s Hall Woods, forest 1 50 m NE ghut; 13.222, 59.582 (9 May 2014) 124-125: Turner’s Hall Woods, SW edge forest; 13.222, 59.586 (9 May 2014) 135-139: Waterford, scrub; 13.117, 59.600 (10 May 2014) 140-142: Oughterson, by house; 13.143, 59.475 (10 May 2014) 143; Three Houses, park; 13.158, 59.460 (10 May 2014) 144: Church Village, by church; 13.137, 59.489 (10 May 2014) 145-149: Batts Rock Beach, waterfront; 13.134, 59.636 (11 May 2014) 150-151: Bridgetown, parking lot; 13.097, 59.617 (1 1 May 2014) 152-153: Brittons Hill, office garden; 13.094, 59.598 (11 May 2014) 154; Henrys, former campus; 13.093, 59.609 (11 May 2014) 155: Bayville, office plantings; 13.087, 59.609 ( 1 1 May 2014) 156: Rendezvous, tree in yard; 13.079, 59.581 (1 1 May 2014) 157-159: Bayville, museum garden; 13.083, 59.602 (12 May 2014) 160-161: Green Garden, American University of Barbados campus; 13.048, 59.523 (12 May 2014) 162-164: Long Beach, sea grape; 13.061, 59.499 (12 May 2014) 165-167: Wotton, by ball field: 13.074, 59.545 (12 May 2014) 168; Welches Heights, store plants; 13.155, 59.61 1 (12 May 2014) 169-173: Turner’s Hall Woods, forest by old bridge; 13.221, 59.584 (13 May 2014) 174-179: Turner’s Hall Woods, forest 300 m NE of ghut; 13.224, 59.582 (13 May 2014) 180-184: Turner’s Hall Woods, forest 450 m NE of ghut; 13.225, 59.581 (13 May 2014) 185-188: Turner’s Hall Woods, forest 650 m NE of ghut; 13.226, 59.580 (13 May 2014) 189; Turner’s Hall Woods, forest 75 m NE of ghut; 13.222, 59.583 (13 May 2014) 190: Two-Mile Hill, by veterinary office; 13.099, 59.592 (14 May 2014) LITERATURE CITED Agosti, D., and L. E. Alonso. 2000. The ALL protocol; a standard protocol for the collection of ground- 30 B REV I ORA No. 548 dwelling ants, pp. 204^206. In D. Agosti, J. D. Majer, L. E. Alonso, and T. R. Schultz eds. Ants: Standard Methods for Measuring and Monitoring Biodiversity. Washington, D.C., Smithsonian In- stitution Press. Ashmead, W. H. 1900. Report upon the Aculeate Hymenoptera of the islands of St. Vincent and Grenada, with additions to the Parasitic Hymenop- tera and a list of the described Hymenoptera of the West Indies. Transactions of the Entomological Society of London 1900: 207-367. Bestelmeyer, B. T., L. E. Alonso, and R. R. Snelling. 2000. The ants (Hymenoptera: Eormicidae) of Laguna del Tigre National Park, Peten, Guatemala. RAP Bulletin of Biological Assessment 16: 75-83. Boer, P. D., and B. Vierbergen. 2008. Exotic ants in the Netherlands (Hymenoptera: Eormicidae). Entomo- logische Berichten 68: 121-129. Bolton, B. 1977. The ant tribe Tetramoriini (Hyme- noptera: Eormicidae). The genus Tetramorium Mayr in the Oriental and Indo-Australian regions, and in Australia. Bulletin of the British Museum (Natural History) Entomology 36: 67-151. Bolton, B. 1979. The ant tribe Tetramoriini (Hyme- noptera: Eormicidae). The genus Tetramorium Mayr in the Malagasy region and in the New World. Bulletin of the British Museum ( Matured History) Entomology 129-181. Bolton, B. 1995. A New General Catalogue of the Ants of the World. Cambridge, Massachusetts, Harvard University Press. Bolton, B. 2000. The ant tribe Dacetini. Memoirs of the American Entomological Institute 65: 1-1028. Borgmeier, T. 1950. Estudos sobre Atta (Hym. Eor- micidae). Memorias do Instituto Oswaldo Cruz 48: 239-263. Borgmeier, T. 1959. Revision der Gattung Atta Fabricius (Hymenoptera, Eormicidae). Stadia En- tomologica 2: 321-390. Brown, W. L., Jr. 1959. The neotropical species of the ant genus Strumigenys Fr. Smith: group of gundla- chi (Roger). Psyche 66: 37-52. Brown, W. L., Jr. 1962. The Neotropical species of the ant genus Strumigenys Fr. Smith: synopsis and keys to species. Psyche 69: 238-267. Brown, W. L., Jr. 1975. Contributions toward a reclas- sification of the Eormicidae. V. Ponerinae, tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini. Search Agricul- ture 5. Brown, W. L., Jr. 1976. Contributions toward a reclassi- fication of the Eormicidae. Part VI. Ponerinae, tribe Ponerini, subtribe Odontomachiti. Section A. In- troduction, subtribal characters. Genus Odontoma- chus. Studia Entomologica 19: 67-171. Brown, W. L., Jr. 1978. A supplement to the world revision of Odontomachus (Hymenoptera: Formici- dae). Psyche 84: 281-285. Castles, J. 1790. Observations on the sugar ants. In a letter from John Castles, Esq. to Lieut. Gen. Melvill, F. R. S. Philosophical Transactions of the Royal Society of London 80: 688-694. Clarke, S. 1670. A True and Eaithful Account of the Pour Chiefest Plantations of the English in America, To Wit, of Virginia, New-England, Bermudas, Barba- dos: With the Temperature of the Air, the Nature of the Soil, the Rivers, Mountains, Beasts, Powls, Birds, Pishes, Trees. Plants, Eruits, &c.: As Also, of the Natives of Virginia, and New-England, Their Re- ligion, Customs, Pishing, Hunting, &c. London, Clavel, Passenger, Cadman, Whitwood, Sawbridge, and Birch. Coke, T. 1808-11. A History of the West Indies Containing the Natural, Civil, and Ecclesiastical History of Each Island: With an Account of the Missions Instituted in those Islands, from the Commencement of Their Civilisation but More Especially of the Missions which Have Been Estab- lished in That Archipelago by the Society Late in Connection with the Rev. John Wesley. Liverpool, Nuttall, Fisher, and Dixon. Crawley, W. C. 1921. New and little-known species of ants from various localities. Annals and Magazine of Natural History (series 9) 7: 87-97. Delabie, j. H. C., and F. Blard. 2002. The tramp ant Hypoponera punctatissima (Roger) (Hymenoptera: Eormicidae: Ponerinae): new records from the Southern Hemisphere. Neotropical Entomology 31: 149-151. Delsinne, T., H. Jourdan, and J. Chazeau. 2001. Premieres donnees sur la monopolisation de re- ssources par I’envahisseur Wasmannia auropunctata (Roger) au sein d’une myrmecofaune de foret seche neo-caledonienne. Actes des Collogues Insectes Sociaux 14: 1-8. Deyrup, M. 1991. Technomyrmex albipes, a new exotic ant in Florida (Hymenoptera: Eormicidae). Plorida Entomologist 74: 147-148. Deyrup, M. 2003. An updated list of Florida ants (Hymenoptera: Eormicidae). Plorida Entomologist 86: 43-48. Deyrup, M., N. Carlin, J. Trager, and G. Umphrey. 1988. A review of the ants of the Florida Keys. Plorida Entomologist 71: 163-176. Deyrup, M., L. Davis, and S. Buckner. 1998. Composition of the ant fauna of three Bahamian Islands, pp. 23-31. In T. K. Wilson ed. Proceedings 2016 ANTS OF BARBADOS (I lYMENOPTERA, FORMICIDAE) 31 of the Seventh Syniposiiini on the Natural History oj the Bahamas. San Salvador, Bahamian Field Station. Devrup, M., L. Davis, and S. Cover. 2000: Exotic ants in Florida. Transactions of the American Entomo- logical Society 126: 293-326. Deyrup, M., J. Tracer, and N. Carlin. 1985. The genus Odontomachus in the southeastern United States (Flymenoptera: Formicidae). Entomological News 96: 188-195. Emery, C. 1895. Beitriige zur Kenntniss der nordamer- ikanischen Ameisenfauna. (Schluss), Zoologische Jahrbiicher. Abteihing fiir Systematik, Geographic and Biologic der Tiere 8: 257-360. Emery, C. 1898. Aggiunte e correzioni al saggio di un catalogo sistematico dei generi Camponotas, Polyrhachis e alTini. Rendiconti delle Sessioni della Reale Accademia delle Scienze dellTstituto di Bologna 2: 225-231. Emery, C. 1913. Etudes sur les Myrmicinae. [V-VIF] Annales de la Societe Entomologiqae de Belgique 57: 250-262. Emery, C. 1923. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [concl.] Genera Insectorum 174c: 207-397. Emery, C. 1925. Hymenoptera, Fam. Formicidae, subfam. Formicinae. Genera Insectorum 183. Espadaler, X., AND V. Bernal. 2003. Exotic ants in the Canary Islands, Spain (Hymenoptera, Formicidae). Vieraea 31: 1-7. FernAndez, F. 2007. Two new South American species of Monomorium Mayr with taxonomic notes on the genus. Memoirs of the American Entomological Institute 80: 128-145. Fisher, B. L., A. K. F. Malsch, R. Gadagkar, J. H. C. Delabie, H. L. Vasconcelos, and J. D. Majer. 2000. Applying the ALL protocol: selected case studies, pp. 207—214. In D. Agosti, J. D. Majer, L. E. Alonso, and T. R. Schultz eds. Ants: Standard Methods for Measuring and Monitoring Biodiversity. Washington, D.C., Smithsonian Institution Press. Forel, a. 1874. Les fourmis de la Suisse. Systematique, notices anatomiques et physiologiques, architec- ture, distribution geographique, nouvelles experi- ences et observations de moeurs. Neue Denkschrif- ten der Allgemeinen Schweizerischen Gesellschaft fiir die Gesammten Natiirwissenschaften 26. Forel, A. 1893. Note sur les Attini. Annales de la Societe Entomologique de Belgique 37: 586-607. Forel, A. 1895. Nouvelles fourmis de diverses prove- nances, surtout d’Australie. Annales de la Societe Entomologique de Belgique 39: 41 49. Forel, A. 1897. Quelques Formicides de I’Antille de Grenada recoltes par M. H. H. Smith. Transactions of the Entomological Society of London 1897: 297-300. Forel, A. 1901. Nouvelles especes de Ponerinae. (Avec un nouveau sous-genre et une espece nouvelle d’Eciton) Revue Suisse de Zoologie 9: 325-353. Forel, A. 1902. Quatre notices myrmecologiques. Annales de la Societe Entomologique de Belgique 46: 170 182. Forel, A. 1908. Ameisen aus Sao Paulo (Brasilien), Paraguay, etc. Gesammelt von Prof. Herm. v. Ihering, Dr. Lutz, Dr. Fiebrig, etc. Verhandliingen der Kaiser Hell- Kdniglichen Zoologisch-Botanischen Gesellschaft in Wien 58: 340-418. Forel, A. 1912a. Formicides Neotropiques. Part I. Annales de la Societe Entomologique de Belgique 56: 28^9. Forel, A. 1912b. Formicides Neotropiques. Part III. 3me sous-famille Myrmicinae (suite). Genres Cre- mastogaster et Pheidole. Memoires de la Societe Entomologique de Belgique 19: 21 1-237. Forel, A. 1912c. Formicides Neotropiques. Part IV. 3me sous-famille Myrmicinae Lep. (suite). Mem- oires de la Societe Entomologique de Belgique 20: 1-32. Forel, A. 191 2d. Formicides Neotropiques. Part 6. 5me sous-famille Camponotinae Forel. Memoires de la Societe Entomologique de Belgique 20: 59-92. Forel, A. 1913. Fourmis d’ Argentine, du Bresil, du Guatemala and de Cuba. Reques de MM. Bruch, Prof. v. Ihering, Mile Baez, M. Peper et M. Rovereto. Bulletin de la Societe Vaudoise des Sciences Naturelles 49: 203-250. Fowler, H. G., O. C. Bueno, T. Sadatsune, and A. C. Montelli. 1993. Ants as potential vectors of pathogens in hospitals in the state of Sao Paulo, Brazil. Insect Science and Its Application 14: 367-370. Goncalves, C. R. 1942. Contribuiqao para o conheci- mento do genero Atta Fabr., das formigas salivas. Boletim da Sociedade Brasileira de Agronomia, Rio de Janeiro 5: 333-358. Gotzek, D., S. G. Brady, R. J. Kallal, and J. S. Lapolla. 2012. The importance of using multi pie approaches for identifying emerging invasive species: the case of the Rasberry crazy ant in the United States. PLoS ONE 1(9): e45314. doi:10.1371/ journal. pone.O 0045314. Hashmi, a. a. 1973. A revision of the Neotropical ant subgenus Myrmothri.x of genus Camponotas (Hymenoptera: Fomiicidae). Stadia Entomologica 16. Holldobler, B., and E. O. Wilson. 1990. The Ants Cambridge, Massachusetts, Harvard University Press. FIughes, G. 1750. Natural History of Barbados. 1972 reprint edition New York, Arno Press. 32 BREVIORA No. 548 Ipser, R. M., M. a. Brinkman, and W. A. Gardner. 2005. First report of Brachymyrmex obscurior Forel (Hymenoptera: Formicidae) from Georgia. Journal of Entomological Science 40: 250-251. JucKER, C., F. Rigato, and R. Regalin. 2008. Exotic ant records from Italy (Hymenoptera, Formicidae). Bollettino di Zoologia agraria e di Bachicoltura 40: 99-107. Kallal, R. J., and J. S. Lapolla. 2012. Monograph of Nylanderia (Hymenoptera; Formicidae) of the world, part II: Nylanderia in the Nearctic Zootaxa 3508. Kempf, W. W. 1951. a taxonomic study on the ant tribe Cephalotini (Hymenoptera: Formicidae). Revista de Entoniologia ( Rio de Janeiro) 22. Kempf, W. W. 1972. Catalogo abreviado das formigas da regiao neotropical (Hymenoptera: Formicidae). Stadia Entomologica 15. Kugler, C. 1994. A revision of the ant genus Rogeria with description of the sting apparatus (Hymenop- tera: Formicidae). Journal of Hymenoptera Re- search 3: 17-89. Lapolla, J. S. 2004. Acropyga (Hymenoptera; Formici- dae) of the World Contributions of the American Entomological Institute 33(3) Lapolla, J. S., and J. Sosa-Calvo. 2006. Review of the ant genus Rogeria (Hymenoptera: Formicidae) in Guyana. Zootaxa 1330: 59-68. Lattke, j. E. 2011. Revision of the New World species of the genus Leptogenys Roger (Insecta: Hymenop- tera: Formicidae: Ponerinae). Arthropod Systemat- ics & Phylogeny 69: 127-264. Lattke, J. E., F. FernAndez, and E. E. Palacio. 2007. Identification of the species of Gnamptogenys Roger in the Americas. Memoirs of the American Entomo- logical Institute 80: 254—270. Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus 1. Editio decima, reformata Holmiae [= Stockholm], L. Salvii. Longing, J. T. 2000. What to do with the data,, pp. 186-203. In D. Agosti, J. D. Majer, L. E. Alonso, and T. R. Schultz eds. Ants: Standard Methods for Measuring and Monitoring Biodiversity Washing- ton, D.C., Smithsonian Institution Press. Longing, J. T. 2003. The Crematogasler (Hymenoptera, Formicidae, Myrmicinae) of Costa Rica. Zootaxa 151. Longino, J. T. 2004 [Internet]. Olympia, Washington: The Evergreen State College [2004 Feb 22; cited 2016 Mar 19]. Ants of Costa Rica website: Cyphomyrmex rimosus (Spinola 1851). Available from: http://academic.evergreen.edu/projects/ants/ genera/Cyphomyrmex/species/rimosus/rimosus.html. MacGown, j. a., and j. a. Forster. 2005. A pre- liminary list of the ants (Hymenoptera: Formicidae) of Alabama. Entomological News 116: 61-74. MacGown, J. A., J. G. Hill, and M. A. Deyrup. 2007. Brachymyrmex patagonicus (Hymenoptera: Formi- cidae), an emerging pest species introduced into the southeastern United States. Elorida Entomologist 90: 457-464. MacGown, J. A., H. Richter, and R. L. Brown. 2013. Notes and new distributional records of invasive ants in the southeastern United States. Midsouth Entomologist 6: 104—120. MacGown, J. A., and J. K. Wetterer. 2013. Distribu- tion and biological notes of Strumigenys margaritae (Hymenoptera: Formicidae: Dacetini). Terrestrial Arthropod Reviews 6: 247-255. Pacheco, J. A., and W. MacKay. 2013. The Systematics and Biology of the New World Thief Ants of the Genus Solenopsis (Hymenoptera: Formicidae). Lampeter, Wales, Edwin Mellen Press. Peck, S. B. 1981. Community composition and zooge- ography of the invertebrate cave fauna of Barba- dos. Elorida Entomologist 64: 519-527. Peterson, C. E. 1973. Sawdust trail, annals of saw- milling and the lumber trade from Virginia to Hawaii via Maine, Barbados, Sault Ste. Marie, Manchac and Seattle to the Year 1860. Bulletin of the Association for Preservation Technology 5: 8^153. Risen, S. J., and C. R. Carroll. 1982. Effect of a keystone predaceous ant, Solenopsis geminata, on arthropods in a tropical agroecosystem. Ecology 63: 1979-1983. Santschi, F. 1929. Melange myrmecologique. Wiener Entomologische Zeitung 46: 84—93. SCHILDER, K., J. HeINZE, G. RoY, AND B. HOLLDOBLER. 1999. Microsatellites reveal clonal structure of populations of the thelytokous parthenogenetic ant Platythyrea punctata (F. Smith) (Hymenoptera: Formicidae). Molecular Ecology 8: 1497-1507. ScHOMBURGK, R. H. 1848. The History of Barbados; Comprising a Geographical and Statistical Description of the Island; a Sketch of the Historical Events since the Settlement; and an Account of its Geology and Natural Productions. 1998 reprint edition. London, Frank Cass. Seal, J. N., K. Kellner, A. Trindl, and J. FIeinze. 2011. Phylogeography of the parthenogenic ant Platythyrea punctata: highly successful colonization of the West Indies by a poor disperser. Journal of Biogeography 38: 868-882. 2016 ANTS OF BARBADOS (I lYMENOPTERA, FORMICIDAE) 33 Seifert, B. 2003. The ant genus Curdiocondyla (Insecta: Hymenoptera; Eormicidae). A taxonomic revision of the C. elegans, C. bidgarkcy C. hatesii, C. niidci, C. shiickardi, C. sUiinbiiloffii, C. wroiighlonii, C. emeryi, and C. mimttior species groups. Annalen des Naturhistorischen Museums in Wien Serie B Botanik und Zoologie 104B: 203-338. SiRJUSiNGH, C., A. Kermarrec, H. Mauleon, C. Lavis, AND J. Etienne. 1992. Biological control of weevils and whitegrubs on bananas and sugarcane in the Caribbean. Florida Entomologist 75: 548-562. Smith, E. 1 855. Descriptions of some species of Brazilian ants belonging to the genera Pseudomyrma, Eciton and Myrmica (with observations on their economy by Mr. El. W. Bates). Transactions of the Royal Entomological Society of London (2) 3: 156-169. Smith, F. 1877. Descriptions of new species of the genera Pseiidomyrma and Tetraponera, belonging to the family Myrmicidae. Transactions of the Entomolog- ical Society of London 10: 57-72. Smith, M. R. 1954. Ants of the Bimini Island Group, Bahamas, British West Indies (Hymenoptera, For- micidae). American Museum Novitates 1671. Snelling, R. R., and J. T. Longing. 1992. Revisionary notes on the fungus-growing ants of the genus Cyphomyrmex, rimosus group (Hymenoptera; For- micidae: Attini), pp. 479^94. In D. Quintero and A. Aiello eds. Insects of Panama and Mesoamerica: Selected Studies. Oxford, Oxford University Press. SoLORZANO, L. A., S. McKenna, and K. Walker. 2005. The Caribbean Biodiversity Initiative at Conserva- tion International, pp 299-302. In P. Miloslavich and E. Klein eds. Caribbean Marine Biodiversity: The Known and the Unknown. Lancaster, Pennsyl- vania, Destech Publications, Inc. Trager, j. C. 1984. A revision of the genus Paratrechina (Hymenoptera; Formicidae) of the continental United States. Sociobiology 9: 49-162. Tucker, R. W. E. 1953. Insects of Barbados. Journal of the Barbados Museum and Historical Society 20: 155-181. Ward, P. S. 1989. Systematic studies on pseudomyrme- cine ants; revision of the Pseudomyrmex ocidatus and P. subtilissimus species groups, with taxonomic comments on other species. Quaestiones Entomolo- gicae 25: 393^68. Ward, P. S. 1992. Ants of the genus Pseudomyrmex (Hymenoptera: Formicidae) from Dominican am- ber, with a synopsis of the extant Antillean species. Psyche 99: 55-85. Ward, P. S. 1993. Systematic studies on Pseudomyrmex acacia-ants (Hymenoptera: Formicidae: Pseudo- myrmecinae). Journal of Hymenoptera Research 2: 117-168. Weber, N. A. 1958. Nomenclatural notes on Proatta and Atta (Hym.: Formicidae). Entomological News 69: 7-13. Weber, N. A. 1972. Gardening Ants: The Attines. Philadelphia, American Philosophical Society. Wetterer, j. K. 2008. Worldwide spread of the longhorn crazy ant, Paratrechina longicornis (Hymenoptera: Formicidae). Myrmecological News 11: 137-149. Wetterer, J. K. 2009a. Worldwide spread of the ghost ant, Tapinoma melanocephalum (Hymenoptera: Formicidae). Myrmecological News 12: 23-33. Wetterer, J. K. 2009b. Worldwide spread of the destroyer ant, Monomorium destructor (Hymenop- tera: Formicidae). Myrmecological News 12: 97-108. Wetterer, J. K. 2009c. Worldwide spread of the penny ant, Tetramorium bicarinatum (Hymenoptera: For- micidae). Sociobiology 54: 811-830. Wetterer, J. K. 2010a. Worldwide spread of the flower ant, Monomorium floricola (Hymenoptera: Formi- cidae). Myrmecological News 13: 19-27. Wetterer, J. K. 2010b. Worldwide spread of the wooly ant, Tetramorium lanuginosum (Hymenop- tera; Formicidae). Myrmecological News 13: 81-88. Wetterer, J. K. 2010c. Worldwide spread of the graceful twig ant, Pseudomyrmex gracilis (Hyme- noptera: Formicidae). Florida Entomologist 93: 535-540. Wetterer, J. K. 2011a. Worldwide spread of the tropical fire ant, Solenopsis geminata (Hymenop- tera: Formicidae). Myrmecological News 14: 21-35. Wetterer, J. K. 2011b. Worldwide spread of the membraniferous dacetine ant, Strumigenys mem- branifera (Hymenoptera: Formicidae). Myrmeco- logical News 14: 129-135. Wetterer, J. K. 2011c. Worldwide spread of Pheidole teneriffana (Hymenoptera: Formicidae). Florida Entomologist 94: 843-847. Wetterer, J. K. 2012a. Worldwide spread of Roger’s dacetine ant, Strumigenys rogeri (Hymenoptera: Formicidae). Myrmecological News 16: 1-6. Wetterer, J. K. 2012b. Worldwide spread of the stigma ant, Pachycondyla stigma ( Hymenoptera: Eormici- dae). Myrmecological News 16: 39^14. Wetterer, J. K. 2012c. Worldwide spread of Emma’s dacetine ant, Strumigenys emmae (Hymenoptera; Formicidae). Myrmecological News 16: 69-74. Wetterer, J. K. 201 2d. Worldwide spread of Emery’s sneaking ant, Cardiocondyla emeryi (Hymenoptera: Formicidae). Myrmecological News 17: 13-20. Wetterer, J. K. 2012e. Worldwide spread of the African big-headed ant, Pheidole megacephala (Hymenop- tera: Formicidae). Myrmecological News 17: 51-62. 34 BREVIORA No. 548 Wetterer, J. K. 2012f. Worldwide spread of the Moorish sneaking ant, Cardiocondyla mauriUmica (Hymenoptera: Formicidae). Sociobiology 59: 985-997. Wetterer, J. K. 2013a. Worldwide spread of the little fire ant, Wasmannki auropimctata (Hymenoptera: Formicidae). Terrestricd Arthropod Reviews 6: 173-184. Wetterer, J. K. 2013b. Exotic spread of Solenopsis invicta (Hymenoptera; Formicidae) beyond North America. Sociobiology 60: 53-63. Wetterer, J. K. 2014a. Worldwide spread of Alluaud’s little yellow ant, Plagiolepis alluaudi (Hymenoptera: Formicidae). Myrmecological News 19: 53-59. Wetterer, J. K. 2014b. Worldwide spread of the lesser sneaking ant, Cardiocondyla minutior (Hymenop- tera: Formicidae). Florida Entomologist 97: 567-574. Wetterer, J. K. 2014c. Geographic distribution of Strumigenys louisianae (Hymenoptera, Formicidae). Terrestrial Arthropod Reviews 7: 159-170. Wetterer, J. K., O. D.xvis, and J. R. Williamson. 2014. Boom and bust of the tawny crazy ant, Nylanderia fidva, on St. Croix, US Virgin Islands. Florida Entomologist 97: 1099-1103. Wetterer, J. K., X. Espadaler, A. L. Wetterer, D. Aguin-Pombo, and a. M. Franquinho-Aguiar. 2007; Ants (Hymenoptera: Formicidae) of the Madeiran Archipelago. Sociobiology 49: 165-291 . Wetterer, J. K., and J. L. W. Keularts. 2008. Population explosion of the hairy crazy ant, Paratrechina piibens (Hymenoptera: Formicidae), on St. Croix, US Virgin Islands. Florida Entomol- ogist 91: 423-427. Wetterer, J. K., and C. D. Lombard. 2010. Fire ants on an important sea turtle nesting beach in St. Croix, US Virgin Islands. Florida Entomologist 93: 449^50. Wetterer, J. K., and B. C. O’Hara. 2002. Ants of the Dry Tortugas, the outermost Florida Keys. Florida Entomologist 85: 303-307. Wetterer, J. K., and D. L. Vargo. 2003. Ants (Hymenoptera: Formicidae) of Samoa. Pacific Science 57: 4094-19. Wetterer, J. K., and A. L. Wetterer. 2004. Ants (Hymenoptera: Formicidae) of Bermuda. Florida Entomologist 87: 212-221. Wetterer, J. K., A. L. Wetterer, and E. Hebard. 2001. Impact of the Argentine ant, Linepithema hiimile, on the native ants of Santa Cruz Island, California. Sociobiology 38: 709-721. Wheeler, W. M. 1905. The ants of the Bahamas, with a list of the known West Indian species. Bulletin of the American Museum of Natural History 21: 79-135. Wheeler, W. M. 1923. Report on the ants collected by the Barbados-Antigua expedition from the Univ. of Iowa in 1918. Studies in Natural History, Iowa University 10: 3-9. Wheeler, W. M. 1926. The Natural History of Ants. From an Unpublished Manuscript in the Archives of the Academy of Sciences of Paris by Rene Antoine Ferchault de Reaumur. New York, Alfred A. Knopf Wheeler, W. M. 1934. Revised list of Hawaiian ants. Occasional Papers Bernice P. Bishop Museum 10. Wilson, E. O. 1988. The biogeography of the West Indian ants (Hymenoptera: Eormicidae), pp. 214-230. In J. K. Liebherr ed. Zoogeography of Caribbean Insects, Ithaca, New York, Cornell University Press. Wilson, E. O. 2003. Pheidole in the New World: A Dominant, Hyperdiverse Ant Genus. Cambridge, Massachusetts, Harvard University Press. Wilson, E. O. 2005. Early ant plagues in the New World. Nature 433: 32. Wilson, E. O., and R. W. Taylor. 1967. Ants of Polynesia. Pacific Insects Monograph 14. y «k V/ 4- i ii/i