Myrmecological News 21 93-99 Vienna, September 2015
Worldwide spread of Tetramorium caldarium (Hymenoptera: Formicidae)
James K. WETTERER & Francisco HITA GARCIA
Abstract
Tetramorium caldarium (ROGER , 1857) is a tramp ant species originally from Africa that has dispersed around the world through human commerce. From 1862 to 1979, T. caldarium was considered a junior synonym of T. simillimum (SMITH , 1851). To document the worldwide spread of T. caldarium , we compiled > 300 published and unpublished specimen site records. In addition, in order to assess their species boundaries, we examined the type specimens of T. caldarium and T. simillimum . We documented Tetramorium caldarium records from 67 geographic areas (countries, island groups, major Caribbean islands, and US states), including several for which there are no previously published records: Austral Islands, Australia, Benin, Cameroon, Cayman Islands, Congo (Republic), Curaçao, Dubai, El Salvador, Gabon, Guadeloupe, Indonesia, Jamaica, Martinique, Namibia, Panama, Scotland, Senegal, South Africa, Tanzania, and Uganda. Tetramorium caldarium is truly cosmopolitan, with records spread across seven of the world's eight bioregions (all except the Antarctic, which has no ants). Tetramorium caldarium records are particularly common on Atlantic islands and from greenhouses and heated buildings in temperate Europe. Key words: Biogeography, biological invasion, exotic species, invasive species. Myrmecol. News 21: 93-99 (online 14 April 2015) ISSN 1994-4136 (print), ISSN 1997-3500 (online) Received 16 February 2015; revision received 17 March 2015; accepted 17 March 2015 Subject Editor: Birgit C. Schlick-Steiner
James K. Wetterer (contact author) , Wilkes Honors College , Florida Atlantic University , 5353 Parkside Drive , Jupiter , FL 33458 , USA. E-mail: [email protected]
Francisco Hita Garcia , Hessisches Landesmuseum Darmstadt , Friedensplatz 1 , 64283 Darmstadt , Germany. E-mail: [email protected]
Introduction FOREL (1911) listed 15 tramp ant species, spread by human For " N. vividula " and " O. haematodus ", it now appears that commerce, which had achieved or were in the process of none of the constituent species are actually cosmopolitan achieving broad cosmopolitan distributions. Previous re- tramps. What FOREL (1911) considered " T. simillimum " ap- views have examined the worldwide spread of 12 of FO- pears to represent at least two species: T. simillimum and REL 's (1911) cosmopolitan species (WETTERER 2005, 2007, T. caldarium (ROGER , 1857). Even without T. caldarium , 2008, 2009a, b, c, 2010a, b, c, 2011a, 2012d, e, WETTERER T. simillimum is still a highly variable species and might & al. 2009). Of these, eight have become major global pests: actually consist of more than one species (FHG, unpubl.). Anoplolepis gracilipes (SMITH , 1857), Linepithema humile At present, T. caldarium is the only species in this com- (MAYR , 1868), Monomorium destructor (JERDON , 1851), plex whose taxonomic boundaries we can confidently de- Monomorium pharaonis (LINNAEUS , 1758), Paratrechina fine. Here, we examine the worldwide spread of T. cal- longicornis (LATREILLE , 1802), Pheidole megacephala (FA- darium . BRICIUS , 1793), Solenopsis geminata (FABRICIUS , 1804), Taxonomic history and Tapinoma melanocephalum (FABRICIUS , 1793). Four others are widespread, but have not developed into signi- ROGER (1857) described Tetrogmus caldarius (= T. calda- ficant pests: Cardiocondyla emeryi FOREL , 1881, Mono- rium ) from a tropical hothouse growing pineapples in Rau- morium floricola (JERDON , 1851), Tetramorium bicarina- den, Prussia (now in Poland, not Germany as listed by BOL - tum (NYLANDER , 1846), and Tetramorium lanuginosum TON 1979 and others). However, shortly after his original MAYR , 1870. All 12 of these species are relatively simple description, ROGER (1862) designated T. caldarium to be to recognize outside their native ranges. The remaining three a junior synonym of T. simillimum , and this species re- names on FOREL 's (1911) list, Nylanderia vividula (NY- mained in synonymy for more than a century, from 1862 LANDER , 1846), Odontomachus haematodus (LINNAEUS , to 1979. This situation changed with the extensive revi- 1758), and Tetramorium simillimum (SMITH , 1851), have all sions of the genus Tetramorium undertaken by BOLTON posed significant taxonomic problems. These three names, (1976, 1977, 1979, 1980, 1985). Although BOLTON (1977) as used by FOREL (1911), each represented multiple spe- initially continued to treat T. caldarium as a junior syno- cies with different geographic ranges, not single species. nym of T. simillimum , BOLTON (1979) later revived T. caldarium as a valid species, based on distinct morpho- graphy web sites (e.g., earth.google.com, www.tageo.com, logical differences. and www.fallingrain.com). If a site record listed a geo- Junior synonyms of Tetramorium caldarium include the graphic region rather than a "point locale", and we had no African taxa T. pusillum hemisi WHEELER , 1922 from the other record for this region, we used the coordinates of Democratic Republic of Congo (Zaire), T. minutum DONIS - the largest town within the region or, in the case of small THORPE , 1942 from Egypt, and T. pauper transformans SAN - islands and natural areas, the center of the region. We did TSCHI , 1914 from Kenya (BOLTON 1979, 1980, 1985, 1995). not map records of Tetramorium caldarium on boats, found BOLTON (1979) distinguished Tetramorium caldarium in newly imported goods or intercepted in transit by qua- from T. simillimum using four characters that can differ- rantine inspectors, e.g., T. caldarium specimens intercepted entiate the two species. Compared with T. simillimum, T. in 1936 in New York on a shipment from the Dominican caldarium has "frontal carinae less strongly developed", Republic (SI and BMNH). Published records usually in- "antennal scrobes feeble", "cephalic sculpture weak", and cluded collection dates. In a number of cases, publica- "head differently shaped" (i.e., T. simillimum with head tions did not include the collection dates for specimens, but broadening behind eyes and T. caldarium with no such we were able to determine them based on information on broadening). Based on our examination, we concur with the collector's travel dates or limited them by the collector's BOLTON (1979). The four characters listed above distin- date of death. guish both species immediately throughout their whole dis- Results tribution ranges. BOLTON (1979, 1980) placed both T. cal- darium and T. simillimum in the T. simillimum species- We compiled Tetramorium caldarium specimen records group. Most of the other 26 members of this species group from > 300 sites worldwide (Fig. 2), documenting the are known only from Sub-Saharan Africa, with a few spe- earliest known T. caldarium records for 67 geographic areas cies also found in the southwest of the Arabian Peninsula, (Tabs. 1 - 3), including several for which there are no pre- Madagascar, and neighboring islands in the western Indi- viously published records: Austral Islands, Australia, Be- an Ocean. Based on this species richness centered in the nin, Cameroon, Cayman Islands, Congo (Republic), Cura- Afrotropical region, it is likely that T. caldarium and T. si- çao, Dubai, El Salvador, Gabon, Guadeloupe, Indonesia, millimum are both native to this region. This conclusion is Jamaica, Martinique, Namibia, Panama, Scotland, Senegal, also supported by their wide distribution ranges in Africa and South Africa, Tanzania, and Uganda. Our only records of their occurrence in undisturbed and anthropogenic habitats. T. caldarium from France, Germany, and Monaco came Tetramorium caldarium workers (Fig. 1) are similar in from the Fauna Europaea website (RADCHENKO 2004) and body size, proportions, and color to another tramp ant spe- lacked any specimen data. BOROWIEC (2014) reported T. cies, Wasmannia auropunctata (ROGER , 1863). Wasmannia caldarium from France, Germany, and Monaco in his cat- auropunctata , however, is slightly smaller, has longer se- alog of European ants based on this same source (L. Boro- tae on the face and dorsum, longer propodeal spines, and wiec, pers. comm.). We obtained a record of T. caldarium a more rectangular petiole in side view. Another impor- from Scotland on the Myrmecology Forum on antfarm.org tant key difference is the presence of a two-segmented (http://antfarm.yuku.com/reply/99651/Queen-ID-request- antennal club in W. auropunctata compared to the three- Small-pictures#reply-99651). segmented club of T. caldarium . The latter also has very In a number of cases, specimens originally identified as weak frontal carinae and reduced antennal scrobes while Tetramorium simillimum were re-identified in later papers both of these characters are strongly developed in W. auro- as T. caldarium , from e.g., Madeira (SAUNDERS 1903, WET - punctata . TERER & al. 2007), the Imatong Mountains, South Sudan (WEBER 1943, BOLTON 1980), and Saint Helena (TAYLOR Methods & WILSON 1961, BOLTON 1976, 1980). BOLTON (1977) Using published and unpublished records, we documented listed three T. simillimum records he later re-identified as the known worldwide range of Tetramorium caldarium. We T. caldarium : from Tres Hermanos and Mayaguez, Puerto obtained unpublished site records from museum speci- Rico (BOLTON 1979) and Pombas, Cape Verde (BOLTON mens in the collections of the Archbold Biological Station 1980). BOLTON (1979) wrote that the only T. caldarium (ABS, identified by M. Deyrup), the California Academy records he knew from the Oriental and Indo-Australian bio- of Science (CAS, identified by FHG), the Natural History geographic regions were three series from India and one Museum London (BNHM, identified by B. Bolton and from New Caledonia (all at the MCZ). FHG), the Museum of Comparative Zoology (MCZ, iden- Problematic records: MEINERT (1861) reported Myr- tified by B. Bolton, S. Cover, X. Espadaler, and FHG), and mica caldaria (= T. caldarium ) from Rosenborg Castle the Smithsonian Institution (SI, identified by B. Bolton). In Gardens in Copenhagen, Denmark, but BOLTON (1995) addition, we used on-line databases with collection infor- listed MEINERT (1861) as having described the gynandro- mation on specimens by Ants of Africa (TAYLOR 2012), morph of T. simillimum and the male of T. caldarium . Bol- AntWeb (www.antweb.org), the Australian National Insect ton (pers. comm.) did not personally examine any Danish Collection (ANIC), the Essig Museum of Entomology (UCB Tetramorium specimens. In the unsorted Tetramorium spe- Essig), Fauna Europaea (RADCHENKO 2004), and the Glo- cimens at the Smithsonian, we found three specimens la- bal Biodiversity Information Facility (www.gbif.org). Fi- beled "Copenhagen 2 / 3 - 1858" and "Copenhagen 17 / 5 nally, JKW collected T. caldarium specimens on numer- - 1858" that are almost certainly specimens collected by ous Atlantic and Caribbean islands, as well as in Florida, Meinert. Stefan Cover identified these as T. caldarium ; we Dubai, and El Salvador. therefore consider all of MEINERT 's (1861) records to be We obtained geographic coordinates for collection sites T. caldarium (and the record of T. simillimum from Den- from published references, specimen labels, maps, or geo- mark to be an error).
94
Fig. 1: Tetramorium caldarium . (a) Head of worker from Hawaii; (b) lateral view of the same worker; (c) dorsal view of the same worker (photos by Eli Sarnat).
Fig. 2: Distribution records of Tetramorium caldarium.
95
Tab. 1: Earliest known records for Tetramorium caldarium Tab. 2: Earliest known records for Tetramorium caldarium from Africa, the Arabian Peninsula, and neighboring is- from Europe, Asia, Oceania, and neighboring islands. UCB lands. Unpublished records include collector, museum Essig = Essig Museum of Entomology. Abbreviations as source, and site. BMNH = Natural History Museum Lon- in Table 1. don. CAS = California Academy of Science. MCZ = Mu- seum of Comparative Zoology. + = no previously pub- Europe Earliest record lished record. * = earliest record unconfirmed. + Poland ≤ 1857 ( ROGER 1857) Earliest record + Denmark ≤ 1858 ( MEINERT 1861) + Madeira ≤ 1847 - 1858 (T.V. Wollaston, BMNH): + England ≤ 1910 ( BOLTON 1979) ≤ multiple sites + Azores ≤ 1930 (A. Méquignon, AntWeb): Fur- + Kenya ≤ 1911 (SANTSCHI 1914 as T. pauper + nas, Sao Miguel ≤ transformans ) + France ≤ 2004 ( RADCHENKO 2004) + Congo (Dem. Rep.) ≤ 1913 (WHEELER 1922 as T. pusillum + Germany ≤ 2004 ( RADCHENKO 2004) ≤ hemisi ) + Monaco ≤ 2004 ( RADCHENKO 2004) + Egypt ≤ 1935 (DONISTHORPE 1942 as T. minu- ≤ tum ) + Spain ≤ 2004 ( REYES & ESPADALER 2005)
+ South Sudan ≤ 1939 ( BOLTON 1980) + Balearic Islands ≤ 2004 ( GOMEZ & ESPADALER 2006) + Mascarene Islands ≤ 1951 ( BOLTON 1980) + Scotland ≤ 2011 (Cantiacus, antfarm.yuku.com): + Glasgow Botanic Gardens + Cape Verde ≤ 1953 ( BOLTON 1980) Asia & Oceania Earliest record + Saint Helena ≤ 1958 (BOLTON 1980) + New Caledonia ≤ 1950 ( BOLTON 1979) + Ivory Coast ≤ 1963 ( BOLTON 1980) + India ≤ 1962 ( BOLTON 1980) + Nigeria ≤ 1980 ( BOLTON 1980) + Indonesia ≤ 1985 (R.H.L. Disney, BMNH): Du- + Cameroon ≤ 1980 (D. Jackson, BMNH): Nkoem von + moga -Bone NP + Saudi Arabia ≤ 1983 ( COLLINGWOOD 1985) + Australia ≤ 1986 (B. Heterick, pers. comm.): Bicton + Oman ≤ 1984 (C OLLINGWOOD & AGOSTI 1996) + Society Islands ≤ 1992 ( MORRISON 1996) + Madagascar ≤ 1991 (A. Pauly, CAS): Ambatondra + Gambier Islands ≤ 1996 ( MORRISON 1997) ≤ zaka + Marquesas Is- ≤ 1996 (MORRISON 1997) + Canary Islands ≤ 1993 ( ESPADALER & BERNAL 2003) + lands + Tanzania ≤ 1993 (A.M. Varela, BMNH): Bukoba + Iran ≤ 2005 ( GHAHARI & COLLINGWOOD 2011) + Yemen ≤ 1993 ( COLLINGWOOD & AGOSTI 1996) + Austral Islands ≤ 2006 (P.D. Krushelnycky, UCB Essig): + Morocco ≤ 1993 ( CAGNIANT & ESPADALER 1993) + Rimatara + Namibia ≤ 1998 (B.L. Fisher, CAS): Waterberg NP + Hawaii ≤ 2007 (R. Chang, AntWeb): Haleiwa + South Africa ≤ 2000 (S. van Noort, CAS): Umtanvuna + Fiji ≤ 2008 (SARNAT 2009) ≤ Nature Reserve
+ Uganda ≤ 2001 (A. Abera, CAS): Kamunyinga Several specimens reported as Tetramorium caldarium + Congo (Rep.) ≤ 2007 (Y. Braet & E. Zassi, TAYLOR on AntWeb from Madagascar and South Africa were mis- ≤ 2012): Brazzaville identifications of local native Tetramorium species (FHG, unpubl.). + Gabon ≤ 2008 (A. Fotso Kuate, TAYLOR 2012): ≤ Awome Discussion
+ Senegal ≤ 2010 (F.B. Ndiaye, TAYLOR 2012): BOLTON (1980) wrote that both Tetramorium simillimum ≤ Katané and T. caldarium "have been widely distributed over the earth by human commerce. In the main their outdoor dis- + Benin ≤ 2012 (J.F. Vayssieres, TAYLOR 2012): ≤ Korobourou tribution is more or less restricted to the tropical and sub- tropical zones, but both are found fairly frequently in the + Dubai ≤ 2012 (J.K. Wetterer, MCZ): Palm Deira temperate zones, associated with man and living in hot- houses, zoos, or other constantly heated buildings". Our analyses match BOLTON 's (1980: figs. 9 - 11) general as- sessment of widespread tropical and subtropical outdoor NOVAK & SADIL (1941) listed Tetramorium simillimum populations for Tetramorium caldarium . On Atlantic Is- from Upper Silesia, which almost certainly refers to the type lands (e.g., Bermuda, the Azores, Madeira, the Canary Is- locale of T. caldarium in Silesian Prussia (ROGER 1857), lands, Cape Verde, and St Helena), T. caldarium was found now in Poland. much more commonly than T. simillimum (ESPADALER &
96 Tab. 3: Earliest known records for Tetramorium caldarium islands, JKW frequently collected T. simillimum , but only from the New World. Abbreviations as in Table 1. rarely encountered T. caldarium . Unfortunately, most spe- cimen records published as T. simillimum between 1862 and Earliest record 1979, when T. caldarium was considered a junior syno- + Puerto Rico ≤ 1908 ( BOLTON 1979) nym, have never been re-examined. It would be useful for researchers to confirm the species identity of all museum + Haiti ≤ 1912 - 1913 ( BOLTON 1979) specimens previously identified as T. simillimum . + Mexico ≤ 1929 (collector unknown, MCZ): Coatepec It is unclear whether the differences in the distributions of Tetramorium simillimum and T. caldarium reflect ecolo- + Brazil ≤ 1965 ( BOLTON 1979) gical differences or whether they are largely due to chance + Peru ≤ 1967 ( BOLTON 1979) differences in where each species has been introduced. However, it appears that T. caldarium is a fairly arid- ≤ + Florida 1968 ( BOLTON 1979) adapted species that does not like shade or forests or wood- + Nicaragua ≤ 1986 ( PERFECTO 1990) land in its native range in Africa, whereas T. simillimum is predominantly found in forested habitats (rainforests, dry + Panama ≤ 1987 (D.M. Olson, MCZ): Cerro Pata de ≤ Macha forests, woodland). Although both Tetramorium simillimum and T. calda- + Bahamas ≤ 1989 - 1994 ( MORRISON 1998) rium are certainly widespread in disturbed environments, + Galapagos ≤ 1992 ( PEZZATTI & al. 1998) they remain quite inconspicuous and we found no evidence that either species has significant ecological impacts. This + Barbados ≤ 1998 (E.O. Wilson & S.P. Cover, MCZ): contrasts with the great impacts reported for the very si- ≤ Casuarina Beach Club milar appearing Wasmannia auropunctata (WETTERER & + Bermuda ≤ 2002 ( WETTERER & WETTERER 2004) PORTER 2003). With the addition of Tetramorium caldarium and the + Texas ≤ 2003 ( COOK 2003) removal of Nylanderia vividula and Odontomachus hae- + Curaçao ≤ 2004 (J.K. Wetterer, MCZ): Van Engelen, matodus from FOREL 'S (1911) list, only two of the 14 cos- ≤ zoo mopolitan species are native to the New World: Linepi- thema humile and Solenopsis geminata . In addition to those + Cayman Is- ≤ 2008 (J.K. Wetterer, MCZ): Foster Village + lands on FOREL 's (1911) list, almost 30 more ant species have become cosmopolitan, with broad ranges in both the Old + Martinique ≤ 2008 (J.K. Wetterer, MCZ): Fort -de -France World and New World (e.g., see WETTERER 2011b, c, d, e, + Jamaica ≤ 2010 (J.K. Wetterer, MCZ): Mona 2012a, b, c, f, 2013c, 2014a, b, WETTERER & RADCHENKO 2011, WETTERER & al. 2012). Although most of these new + Guadelo upe ≤ 2011 (J.K. Wetterer, MCZ): Pointe -à-Pitre cosmopolitans are inconspicuous Old World species, two + Guatemala ≤ 2012 ( BRANSTETTER & SAENZ 2012) of the greatest ecological and economic pests are New World natives: Solenopsis invicta BUREN , 1972 and Wasmannia ≤ + El Salvador 2012 (J.K. Wetterer, MCZ): San Salvador auropunctata (WETTERER 2013a, b). With the modern glo- + Dominican ≤ 2012 (LUBERTAZZI & ALPERT 2014) bal economy, one can expect more new cosmopolitans in + Republic the future. Acknowledgments BERNAL 2003, WETTERER & WETTERER 2004, WETTERER We thank M. Wetterer, H.C. Wagner, J. Majer, B. Bol- & al. 2004, 2006). With the exception of one record from ton, and A. Andersen for comments on this manuscript; an urban plaza in Córdoba, Spain (REYES & ESPADALER S. Cover for help, encouragement, and ant identification; 2005), all known outdoor records of T. caldarium at lati- B. Bolton (BMNH), S. Cover (MCZ), M. Deyrup (ABS), tudes higher than 32° come from islands in the Atlantic J. Hogan (ONHM), and T. Schultz (SI) for help with their (Bermuda, the Azores, and Madeira; WETTERER & WET - respective ant collections; J. Woinarski, D. Ward, M. Lush, TERER 2004, WETTERER & al. 2004, 2007) and the Mediter- A. Fotso Kuate, B. Heterick, and M. DaSilva for provid- ranean (the Balearic Islands; GOMEZ & ESPADALER 2006). ing unpublished specimens records; O'Brien for GIS help; In addition, T. caldarium appears to be more common than E. Sarnat for use of his excellent photos; D.P. Wojcik and T. simillimum in greenhouses and heated buildings, prima- S.D. Porter for compiling the FORMIS bibliography; L. rily in temperate areas. Whereas there are many recent in- Lesperance of the FAU library for processing so many in- door records of T. caldarium from temperate areas (Tab. 2), terlibrary loans; Florida Atlantic University, the National we found only one confirmed published record of T. simil- Science Foundation, and the National Geographic Society limum from a temperate site: its type locale in England for financial support. (SMITH 1851). In addition, unconfirmed temperate records of T. simillimum come from England (SAUNDERS 1880, References EMERY 1909), Winnipeg, Canada (AYRE 1977), France AYRE , G.L. 1977: Exotic ants in Winnipeg. – Manitoba Entomo- (BERNARD 1968), and Estonia (RADCHENKO 2004). It is logist 11: 41-44. possible that most or all unconfirmed temperate records BERNARD , F. 1968: Faune de l'Europe & du Bassin Méditerra- of T. simillimum were actually T. caldarium . néen. 3. Les fourmis (Hymenoptera: Formicidae) d'Europe oc- In contrast, in most tropical areas (except Atlantic is- cidentale & septentrionale. – Masson, Paris, 411 pp. lands), Tetramorium simillimum has been collected more BOLTON , B. 1976: The ant tribe Tetramoriini (Hymenoptera: For- often than has T. caldarium. For example, on West Indian micidae). Constituent genera, review of smaller genera and re-
97 vision of Triglyphothrix FOREL . – Bulletin of the British Museum MORRISON , L.W. 1997: Polynesian ant (Hymenoptera: Formici- (Natural History) Entomology 34: 281-379. dae) species richness and distribution: a regional survey. – Acta BOLTON , B. 1977: The ant tribe Tetramoriini (Hymenoptera: For- Oecologica 18: 685-695. micidae). The genus Tetramorium MAYR in the Oriental and MORRISON , L.W. 1998: A review of Bahamian ant (Hymeno- Indo-Australian regions, and in Australia. – Bulletin of the ptera: Formicidae) biogeography. – Journal of Biogeography British Museum (Natural History) Entomology 36: 67-151. 25: 561-571. BOLTON , B. 1979: The ant tribe Tetramoriini (Hymenoptera: For- NOVÁK , V. & SADIL , J. 1941: Klí č k ur čování mravenc ů st řední micidae). The genus Tetramorium MAYR in the Malagasy re- Evropy se zvláštním z řetelem k mraven čí zví řen ě Čech a Mor- gion and in the New World. – Bulletin of the British Museum avy. – Entomologicke Listy 4: 65-115. (Natural History) Entomology 38: 129-181. PERFECTO , I. 1990: Indirect and direct effects in a tropical agro- BOLTON , B. 1980: The ant tribe Tetramoriini (Hymenoptera: For- ecosystem: the maize-pest-ant system in Nicaragua. – Ecol- micidae). The genus Tetramorium MAYR in the Ethiopian zoo- ogy 71: 2125-2134. geographical region. – Bulletin of the British Museum (Natu- PEZZATTI , B., IRZAN , T. & CHERIX , D. 1998: Ants (Hymenoptera: ral History) Entomology 40: 193-384. Formicidae) of Floreana: lost paradise? – Noticias de Galapa- BOLTON , B. 1985: The ant genus Triglyphothrix FOREL a synonym gos 59: 11-20. of Tetramorium MAYR (Hymenoptera: Formicidae). – Journal RADCHENKO , A. 2004: Formicidae. – Fauna Europaea version 1.1: of Natural History 19: 243-248.
98 WETTERER , J.K. 2009a: Worldwide spread of the ghost ant, Tapi- WETTERER , J.K. 2012f: Worldwide spread of the Moorish sneak- noma melanocephalum (Hymenoptera: Formicidae). – Myr- ing ant, Cardiocondyla mauritanica (Hymenoptera: Formici- mecological News 12: 23-33. dae). – Sociobiology 59: 985-997. WETTERER , J.K. 2009b: Worldwide spread of the destroyer ant, WETTERER , J.K. 2013a: Exotic spread of Solenopsis invicta (Hy- Monomorium destructor (Hymenoptera: Formicidae). – Myr- menoptera: Formicidae) beyond North America. – Sociobiol- mecological News 12: 97-108. ogy 60: 53-63. WETTERER , J.K. 2009c: Worldwide spread of the penny ant, Te- WETTERER , J.K. 2013b: Worldwide spread of the little fire ant, tramorium bicarinatum (Hymenoptera: Formicidae) . – Socio- Wasmannia auropunctata (Hymenoptera: Formicidae). – Ter- biology 54: 811-830. restrial Arthropod Reviews 6: 173-184. WETTERER , J.K. 2010a: Worldwide spread of the flower ant, Mo- WETTERER , J.K. 2013c: Worldwide spread of the difficult white- nomorium floricola (Hymenoptera: Formicidae). – Myrmeco- footed ant, Technomyrmex difficilis (Hymenoptera: Formici- logical News 13: 19-27. dae). – Myrmecological News 18: 93-97. WETTERER , J.K. 2010b: Worldwide spread of the wooly ant, Te- WETTERER , J.K. 2014a: Worldwide spread of Alluaud's little tramorium lanuginosum (Hymenoptera: Formicidae). – Myr- yellow ant, Plagiolepis alluaudi (Hymenoptera: Formicidae). mecological News 13: 81-88. – Myrmecological News 19: 53-59. WETTERER , J.K. 2010c: Worldwide spread of the pharaoh ant, WETTERER , J.K. 2014b: Worldwide spread of the lesser sneak- Monomorium pharaonis (Hymenoptera: Formicidae). – Myr- ing ant, Cardiocondyla minutior (Hymenoptera: Formicidae). mecological News 13: 115-129. – The Florida Entomologist 97: 567-574. WETTERER , J.K. 2011a: Worldwide spread of the tropical fire ant, WETTERER , J.K., ESPADALER , X., WETTERER , A.L. & CABRAL , Solenopsis geminata (Hymenoptera: Formicidae). – Myrme- S.G.M. 2004: Native and exotic ants of the Azores (Hymeno- cological News 14: 21-35. ptera: Formicidae). – Sociobiology 44: 1-20. WETTERER , J.K. 2011b: Worldwide spread of the membranifer- WETTERER , J.K., ESPADALER , X., WETTERER , A.L., AGUIN - ous dacetine ant, Strumigenys membranifera (Hymenoptera: POMBO , D. & FRANQUINHO -AGUIAR , A.M. 2006: Long-term Formicidae). – Myrmecological News 14: 129-135. impact of exotic ants on the native ants of Madeira. – Ecolo- WETTERER , J.K. 2011c: Worldwide spread of the yellow-footed gical Entomology 31: 358-368. ant, Nylanderia flavipes (Hymenoptera: Formicidae). – The Flo- WETTERER , J.K., ESPADALER , X., WETTERER , A.L., AGUIN -POMBO , rida Entomologist 94: 582-587. D. & FRANQUINHO -AGUIAR , A.M. 2007: Ants (Hymenoptera: WETTERER , J.K. 2011d: Worldwide spread of Tetramorium lu- Formicidae) of the Madeiran Archipelago. – Sociobiology 49: cayanum (Hymenoptera: Formicidae). – The Florida Entomo- 265-297. logist 94: 827-831. WETTERER , J.K., KRONAUER , D.J.C. & BOROWIEC , M.L. 2012: WETTERER , J.K. 2011e: Worldwide spread of Pheidole tenerif- Worldwide spread of Cerapachys biroi (Hymenoptera: For- fana (Hymenoptera: Formicidae). – The Florida Entomologist micidae: Cerapachyinae). – Myrmecological News 17: 1-4. 94: 843-847. WETTERER , J.K. & PORTER , S.D. 2003: The little fire ant, Was- WETTERER , J.K. 2012a: Worldwide spread of Roger's dacetine mannia auropunctata : distribution, impact, and control. – So- ant, Strumigenys rogeri (Hymenoptera: Formicidae). – Myrme- ciobiology 42: 1-41. cological News 16: 1-6. WETTERER , J.K. & RADCHENKO , A. 2011: Worldwide spread of WETTERER , J.K. 2012b: Worldwide spread of the stigma ant, Pa- the ruby ant, Myrmica rubra (Hymenoptera: Formicidae). – chycondyla stigma (Hymenoptera: Formicidae). – Myrmeco- Myrmecological News 14: 87-96. logical News 16: 39-44. WETTERER , J.K. & WETTERER , A.L. 2004: Ants (Hymenoptera: WETTERER , J.K. 2012c: Worldwide spread of Emma's dacetine Formicidae) of Bermuda. – The Florida Entomologist 87: 212- ant, Strumigenys emmae (Hymenoptera: Formicidae). – Myr- 221. mecological News 16: 69-74. WETTERER , J.K., WILD , A.L., SUAREZ , A.V., ROURA -PASCUAL , WETTERER , J.K. 2012d: Worldwide spread of Emery's sneaking N. & ESPADALER , X. 2009: Worldwide spread of the Argen- ant, Cardiocondyla emeryi (Hymenoptera: Formicidae). – Myr- tine ant, Linepithema humile (Hymenoptera: Formicidae). – mecological News 17: 13-20. Myrmecological News 12: 187-194. WETTERER , J.K. 2012e: Worldwide spread of the African big- WHEELER , W.M. 1922: The ants collected by the American Mu- headed ant, Pheidole megacephala (Hymenoptera: Formici- seum Congo Expedition. – Bulletin of the American Museum dae). – Myrmecological News 17: 51-62. of Natural History 45: 39-269.
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