Ants ofTonga1
James K. Wetterer 2
Abstract: This paper presents combined published, unpublished, and new ant records from 17 islands ofTonga representing all four island groups: Tongatapu (Tongatapu, 'Eua, 'Onevai, Pangaimotu), Ha'apai (Lifuka, Kao, Tofua, 'Uonu kahahake, Nomuka, Nomuka-iki, Mango, Telekitonga), Vava'u (Vava'u, Nua papu, Kapa), and the Niuas (Niuatoputapu, Niuafo'ou). These records increase the list of ants known from Tonga to 53 species. Ten species, including six un described species, are local endemics found only in Tonga or only in Tonga and Samoa: Adelomyrmex sp., Camponotus conicus, Camponotus nig;rifrons, Hypoponera sp., Monomorium sp., Ochetellus sp., Pheidole sp., Pristomyrmex sp., Strumigenys zakharovi, and Vollenhovia samoensis. Another 21 species are broadly distributed Pacific natives: Anochetus g;raeffei, Camponotus chloroticus, Hypoponera confinis, Monomorium liliuokalanii, Monomorium talpa, Odontomachus simillimus, Oligo myrmex atomus, Pheidole oceanica, Pheidole sexspinosa, Pheidole umbonata, Ponera incerta, Ponera tenuis, Pyramica dubia, Rogeria stigmatica, Solenopsis papuana, Stru migenys godeffroyi, Tapinoma minutum, Technomyrmex albipes, Tetramorium in solens, Tetramorium pacificum, and Tetramorium tonganum. Finally, 22 species are not native to the Pacific region, but were brought to the region by human com merce: Anoplolepis g;racilipes, Cardiocondyla emeryi, Cardiocondyla nuda, Hypoponera opaciceps, Hypoponera punetatissima, Monomorium floricola, Monomorium pharaonis, Monomorium sechellense, Paratrechina bourbonica, Paratrechina longicornis, Paratre china vaga, Pheidole fervens, Pheidole megacephala, Plagiolepis alluaudi, Pyramica membranifera, Solenopsis geminata, Strumigenys emmae, Strumigenys rogeri, Tapi noma melanocephalum, Tetramorium bicarinatum, Tetramorium lanuginosum, and Tetramorium simillimum. The number of ant species now known from Tonga is much as would be expected based on the species-area relationship for the neighboring island groups of Fiji, Wallis and Futuna, and Samoa. Differences in ant species richness among these island groups is primarily due to a greater number of local endemics in the island groups with greater land area.
THE KINGDOM OF TONGA consists of about 40 of the Tongan islands are inhabited year 170 islands with a total land area of '"700 round. km2 spread across 350,000 km2 of western Due to the difficulty and expense of travel Polynesia between 15-23° Sand 173-177° to distant and unpopulated islands, most W. Tonga is divided into four major island Tongan ant records come from the three groups (from south to north): Tongatapu, largest and most accessible islands: Tonga Ha'apai, Vava'u, and the Niuas. Only about tapu, 'Eua, and Vava'u. On these islands, hu man activity has heavily impacted most flatter areas (Drake et al. 1996, Steadman and Frei feld 1998, Franklin et al. 1999). Tongatapu, 1 Financial support was provided by Harvard Univer 2 sity, Florida Atlantic University, and Edward O. Wilson. by far the largest island of Tonga (259 km ), Manuscript accepted 10 July 200l. makes up 37% of the total land area of the 2 Honors College, Florida Atlantic University, 5353 kingdom. Tongatapu is very flat, tilting only Parkside Drive, Jupiter, Florida 33458. slightly up to 30-m cliffs along the southern shore (maximum elevation, 82 m). About Pacific Science (2002), vol. 56, no. 2:125-135 two-thirds ofTonga's 100,000 people live on © 2002 by University of Hawai'i Press. Tongatapu. 'Eua, also part of the Tongatapu All rights reserved group, is the third largest island (87 km2) and
125 126 PACIFIC SCIENCE . April 2002
2.0.------~ eastern Polynesia (Niue, the Cook Islands, Fiji and French Polynesia combined) to the east (Taylor 1967, Wilson and Taylor 1967, --;; 1.8 Morrison 1996a,b, 1997). In this study, I in .~ ..'" crease the list of ants known from Tonga to Po. '" 53 species. 5 .....• 1.6 o MATERIALS AND METHODS ~ Wallis & Futuna eo In addition to published records of ants from oS 1.4 Tonga, I searched for ant specimens from Niue - Tonga in the collections of several museums: _ Tonga (from Wilson & Taylor 1967) the U.S. National Museum of Natural His 1.2+------,....------,....------l tory (USNM) in Washington, D.C.; the Amer 2.0 3.0 4.0 ican Museum of Natural History (AMNH) log
TABLE 1 Collection Dates and Locales for Ants from Tonga
Collector (Source) Collection Date Island(s) a. Unlisted (Mayr 1870, 1876) <1870 T? b. Cottrell-Dormer (Wilson and Taylor 1967) <1914-<1935 T c. Wilder (Wilson and Taylor 1967) 1923-1925 T, V, "Ha'apai" d. Unknown (USNM) 1930 Niuafo'ou e. Krauss (Wilson and Taylor 1967) 1956 T,E,V f. Rogers & Rogers (NZAC) 1971 Niuatoputapu g. Litsinger (NZAC) 1973-1974 T,E,V h. Krauss (AMNH) 1974-1979 T,E i. Watt (NZAC) 1975-1977 T, E, Niuafo'ou j. Maddison (NZAC) 1975-1977 T,E k. Chernov (Dlussky 1994a) 1976-1977 Niuafo'ou m. Dlussky et al. (Dlussky 1994a) 1980 T, E, V, + many others p. Stechmann et al. (1996) 1986 T r. Carver et al. (1993) 1990-1992 T,E s. Curio (pers. comm.) 1992 Niuafo'ou t. Wetterer (this study) 1995 T, E, V, Lifuka, Kapa
Sources; USNM, u.s. National Museum efNaturai History; AMNH, American Museum of Natural History; NZAC, New Zealand Ar thropod Collection. Islands: T, Tongatapu; E, 'Eua; V, Vava'u. 128 PACIFIC SCIENCE April 2002
TABLE 2 Native Pacific Ants from Different Island Groups of Tonga
Tongatapu Pacific Species Tongatapu 'Eua Ha'apai Vava'u Niuas Range Origin Local endemics Adelomyrmex sp. !!!t !!! 0 Camp(motus conicus ill< em ET Camponotus nigrifrons ill< gj!!! !!! ET Hypoponlffa sp. t 0 Monomorium sp. 0 Ochetellus sp. !!!t !!!t !!! 0 Pheidole sp. !!!t ET Pristomyrmex sp. f S- -- 0 Strumigenys zakharovi !!! ET Vollenhovia samoensis !!!t S- -- ES Wide-ranging Pacific natives Anochetus graeffei !!!t !!! !!!t !!! MWS- CF PO Camponotus chloroticus ill the Niuas. In 1980, G. M. Dlussky, A. A. Za 49. Two species that Dlussky (1994a) listed as kharov, and S. 1. Golovatch collected ants on Pheidole euana Dlussky and Vollenhovia kallisto the islands of Tongatapu, 'Eua, and Vava'u, Dlussky were, in fact, never described (G. as well as an assortment of other Tongan is Dlussky, pers. comm.), so these names are in lands in the Tongatapu group (Onevai, Pan valid. Dlussky (pers. comm.) could not defini gaimotu), Ha'apai group (Nomuka, Nomuka tively distinguish the Vollenhovia specimens iki, Mango, Telekitonga, 'Uonukahahake, from Vollenhovia samoensis Mayr. I there Kao, Tofua), and Vava'u group (Nuapapu). fore list these two species as Pheidole sp. and Together, 46 ant species were collected by the Vollenhovia samoensis. Dlussky (1994a) also two Soviet surveys, which brought the total reported two ant species from Tonga, Adelo number of ant species known from Tonga to myrmex hirsutus Mann and Ochetellus sororis Ants ofTonga Wetterer 129 TABLE 3 Pantropical Exotic Ants from Different Island Groups ofTonga Tongatapu Pacific Species Tongatapu 'Eua Ha'apai Vava'u Niuas Range Origin Anoplolepis gracilipes arnrt t );!!t emt ks MWSNCF PA Cardiocondyla emeryi hInt hInt );!!t t k MWSNCF PA Cardiocondyla nuda aemt );!! );!! );!!t fk MWSNCF PA Hypoponera opaciceps gt @t );!! t M- S- CF PN Hypoponera punctatissima );!!t ! Note: Symbols as in Table 2. PA, pantropical ofAfrican origin; PN, pantropical ofNeotropical origin. " species in Dlussky (I 994a) subsequently synonomized with a more wide-ranging species. (Mann), otherwise known only from Fiji Museum Specimens (Mann 1921). I believe, however, that these specimens are probably undescribed Tongan In the USNM, I found one specimen ofPheidole endemics, which I list as Adelomyrmex sp. and megacephala collected on Niuafo'ou in the Ochetellus sp. (see later in Results for expla Niuas in October 1930. The collector's name nation). was not recorded. This is by far the earliest Swaney (1994), in her travel guide to record of P. megacephala in Tonga. Tonga, described observations of "long In the AMNH, I found ant specimens that legged ants," the common name for Anoplole N. Krauss collected on Tongatapu and 'Eua pis gracilipes (Smith), on Niuafo'ou in the in 1974,1978, and 1979 (Tables 1-3). Niuas attacking newly hatched chicks of the In the MCZ, I found Tongan ant specimens endemic Niuafo'ou incubator bird (Mega collected by W. Cottrell-Darmer (no date) podius pritchardii Gray). Swaney's (1994) ref and by N. Krauss (in 1956) that were pre erence brought to my attention unpublished viously listed by Wilson and Taylor (1967). records of ants collected by E. Curio on In addition, N. Krauss collected Tapinoma Niuafo'ou in 1992 (see in a later section). melanocephalum on Vava'u in February 1956, Stechmann et al. (1996) recorded seven ant a record not included in Wilson and Taylor species tending aphids on Tongatapu in 1986. (1967). Carver et al. (1993) recorded 11 species of In the NZAC, I found ant specimens from ants tending aphids on Tongatapu in 1990 four collections: W. Rogers and G. Rogers 1992. collected ant species on Niuatoputapu in the 130 PACIFIC SCIENCE· April 2002 Niuas between March and September 1971; Categories ofAnt Species J. A. Litsinger collected on Tongatapu, 'Eua, and Vava'u in 1973 and 1974; P. Maddison Ofthe 53 ant species ofTonga, 10 species are collected on Tongatapu and 'Eua from 1975 known only from Tonga or from Tonga and to 1977; J. c. Watt collected on Tongatapu, Samoa, 21 species appear to be broad-ranged 'Eua, and Niuafo'ou in the Niuas in 1975 and Pacific natives, and 22 appear to be exotic to 1977 (Tables 1-3). the Pacific region (Tables 2 and 3). Wilson The previously unpublished records of and Hunt (1967) considered Monomorium Tetramorium insolens (Smith) and Pristo minutum Mayr an exotic tramp. Pacific speci myrmex sp. brought the total number of ant mens previously assigned to this species, how species known from Tonga to 51. This species ever, have been reclassified as Monomorium of Pristomyrmex is currently being described liliuokalanii Forel (see Bolton 1995), which I categorized as a wide-ranging Pacific native. by M. Chang at the MCZ as part of his disser tation research. DISCUSSION Other Unpublished Ant Records Comparison ofthe Tonga Ant Fauna with Those In 1992, E. Curio collected nine ant species ofNeighboring Islands and Island Groups on Niuafo'ou in conjunction with research on the endemic megapodes (E. Curio, pers. The number of ant species now known from Tonga (53) is much as would be expected comm.; Tables 1-3). B. Bolton at the Natural History Museum in London identified Cu based on the species-area relationship of the rio's specimens (E. Curio, pers. comm.). neighboring island groups of Fiji, Wallis and Futuna, and Samoa, to the west and north. Comparing data from these four island groups New Ant Collection on a log-log plot, the number of species (N.) increases linearly with total land area (A) I collected 42 ant species in Tonga (Tables 2 (Figure 1; least squares regression: logN. = and 3), including five that did not match any 2 published descriptions. I tentatively consider 1.177 + 0.1831ogA, R = 0.98). The increase the undescribed Pheidole as the same as Dlus in ant species with land area is primarily due sky's (1994a) Pheidole sp., pending further to an increased number of local endemics study. Two other undescribed species, Adelo in the island groups with greater land area myrmex sp. and Ochetellus sp., are in the same (Table 4). genera as Adelomyrmex hirsutus Mann and Niue, a solitary island directly east of Ochetellus sororis (Mann), reported by Dlussky Tonga, with only 19 known ant species, has (1994a). S. Cover distinguished my specimens fewer than half the number of species ex from types in the MCZ ofthese two Fijian spe pected compared with the numbers known cies. Dlussky (pers. comm.), however, never for Fiji, Wallis and Futuna, Tonga, and Sa compared the Soviet specimens with types. I moa (Figure 1). It seems likely that, due to therefore have tentatively assumed that the incomplete sampling, many inconspicuous Soviet specimens were misidentified and are ant species have been overlooked in Niue. the same undescribed species as those that I found. Finally, I found two undescribed spe Endemic Ant Species in Tonga cies, Hypoponera sp. and Monomorium sp., not reported by any previous collector. My new Eight ant species are known only from Tonga records raise the total number of ant species and two species are known only from Tonga known in Tonga to 53. and Samoa. Mayr described two of these I have donated all ant specimens I col species (Camponotus nigrifrons Mayr in 1870; lected on Tonga to the MCZ, with duplicates Camponotus conicus Mayr in 1876); Dlussky to be sent to appropriate collections in the described one (Strumigenys zakharovi Dlussky Pacific and elsewhere. in Dlussky 1994b). Five Tonga endemic spe- Ants ofTonga . Wetterer 131 TABLE 4 Number ofAnt Species Known from the Islands and Island Groups ofTonga and from the Surrounding Islands and Island Groups Island/Island Land Area Local Pacific Exotic Group (km2) Endemic Native Tramp Total Tongatapu 259 3 15 18 36 'Eua 87 8 20 19 47 Ha'apai group 109 4 17 17 38 Vava'u group 119 1 15 18 34 Niuas group 72 1 11 12 24 Wallis and Futuna 177 0 21 16 37 Niue 259 0 9 10 19 Tonga (total) 699 10 21 22 53 Samoa 3,132 15 27 25 67 Fiji 18,272 56 15 17 88 cies remain undescribed (Adelomyrmex sp., For example, Wilson and Taylor (1967) wrote Hypoponera sp., Monomorium sp., Ochetellus that Pyramica dubia (Brown) is probably a sp., and Pheidole sp.). Vollenhovia samoensis is tramp that originated in New Guinea and known only from 'Eua and from Samoa. spread around the Pacific by human com Pristomyrmex sp. is known only from Niuato merce. In further evidence of this uncertainty, putapu and from Samoa. Dlussky (1994a) placed the origin of P. dubia Only three of the local endemic ant species in Asia. have been found on Tongatapu, and just one Four ant species that are native to the of these has been collected since 1956. In western Pacific region have invaded other contrast, 8 of the 10 have been found on the parts of the world. One such "Pacific-native nearby, smaller island of 'Eua, all of which tramp" is Technomyrmex albipes (Smith), the species have been collected one or more times "white-footed" ant, which is now found as an since 1980. The paucity of endemic species exotic in many areas, including New Zealand, on Tongatapu may relate, in part, to its very Hawai'i, California, Florida, South Africa, flat topography, which offers little habitat India, China, Madagascar, and Saudi Arabia variation and has allowed almost the entire (Wilson and Taylor 1967, Wetterer 1997a, island to come under cultivation. In addition, 1998, McGlynn 1999). In addition, Tetramo the higWy destructive big-headed ant, Phei rium tonganum Mayr is an exotic in Hawai'i dole megacephala, now occurs in high densities and Brazil; Tetramorium pacificum Mayr is in over most of the island (unpub!. data). In Florida, Canada, Central America, and the contrast, the eastern slope of 'Eua has varied Caribbean; and Odontomachus simillimus Smith habitat, is too steep for cultivation, and has is in the Caribbean (McGlynn 1999). not yet been invaded by P. megacephala. Dlussky (1994a) used a different classifica tion scheme than did Wilson and Taylor Native Pacific Ant Species in Tonga (1967) and Wilson and Hunt (1967). Dlussky (1994a) proposed the geographical origin of According to the classifications ofWilson and species, but was not explicit about which spe Hunt (1967), 21 of the ant species found in cies predated human arrival in Polynesia (see Tonga are wide-ranging Pacific natives. Tables 2 and 3). Dlussky (1994a) classified However, it is not possible to determine reli the 21 wide-ranging Pacific natives of Tonga ably which are actually native to Tonga (i.e., as two regional endemics, seven species of predating humans) and which are more re New Guinean origin, two of Asian origin, cent arrivals from other parts of the Pacific. and nine pantropical species of Asian origin 132 PACIFIC SCIENCE· April 2002 (Table 2). One species, Tetramorium insolens, and 3): seven pantropical species of Asian was not found in Dlussky's (1994a) study. origin, 11 pantropical species of African ori gin, two pantropical species of Neotropical origin, and one species of Asian origin. In Exotic Ant Species in Tonga addition, one species, classified by Dlussky According to the classifications ofWilson and (1994a) as Monomorium fossulatum Emery of Hunt (1967), 22 ant species found in Tonga Asian origin, has been synonomized with the are alien tramp species not native to the Pa much broader-ranging tramp species Mono cific, but brought to the region by human morium sechellense. commerce. The scarcity of records from It is striking that only two exotic ant spe Tonga precludes dating the arrivals of exotic cies in Tonga are of N ew World origin: species precisely. However, seven of these Hypoponera opaciceps and Solenopsis geminata. invaders had already arrived in Tonga by This, no doubt, relates to not only geographic the 1860s: Anoplolepis gracilipes, Cardiocondyla proximity and ocean currents, but also tradi nuda (Mayr), Monomorium floricola (Jerdon), tional trade routes. Certainly in the future, Monomorium pharaonis (L.), Paratrechina vaga more arrivals of ants from the New World (Forel), Tapinoma melanocephalum, and Tetra can be expected. morium bicarinatum (Nylander) (Mayr 1870, 1876). Wilson and Taylor (1967) listed six ad Future ofthe Tongan Ant Fauna ditional tramp species in Tonga: Hypoponera punctatissima (Roger), Paratrechina bourbonica Even with the new records published here, (Forel), Paratrechina longicornis (Latreille), the ants of most of the 170 islands of Tonga Pheidole fervens Smith, Solenopsis geminata remain relatively unknown. Only three is (Fabr.), and Tetramorium simillimum (Smith). lands, Tongatapu, 'Eua, and Vava'u, have It is possible that all five have been introduced been well sampled for ants. No doubt many to Tonga within the past 100 yr. undocumented native ant species persist on Nine exotic species now in Tonga were other Tongan islands. However, tramp ants, absent from Wilson and Taylor's (1967) list: particularly Pheidole megacephala, appear to be Cardiocondyla emeryi Forel, Hypoponera opaci rapidly exterminating native ants on Tonga ceps (Mayr), Monomorium sechellense Emery, tapu and 'Eua (unpubl. data). Pheidole megacephala, Plagiolepis alluaudi Forel, Although Pheidole megacephala specimens Pyramica membranifera (Emery), Strumigenys were collected on Tongatapu by Maddison in emmae (Emery), Strumigenys rogeri Emery, 1975, contemporary collections on Tonga and Tetramorium lanuginosum Mayr. Only tapu by Litsinger (in 1973-1974) and Watt one has an unpublished record predating (in 1975-1977) did not include P. mega 1967, Pheidole megacephala. Museum speci cephala, suggesting that P. megacephala pop mens, reported here for the first time, indi ulations in the 1970s were limited. By 1995, cate that P. megacephala invaded the far north however, P. megacephala dominated vast areas of Tonga by 1930. The first P. megacephala of Tongatapu, where it occurred in almost specimens from the main island ofTongatapu every log and under almost every rock (pers. were collected by Maddison in 1975. The obs.). The 1980 Soviet collection documented other eight exotic ant species not on Wilson by Dlussky (1994a) recorded the first P. and Taylor's (1967) list were all collected in megacephala on Vava'u. In 1995, I made the Tonga for the first time in the 1970s or later. first collections of P. megacephala in 'Eua and Although these late records could represent Ha'apai. At that time, P. megacephala was recent invasions, it is possible that these in the dominant ant species over large parts of vaders came much earlier, but were simply western 'Eua, but had a limited population on not documented. Lifuka in Ha'apai. The vast population ex Dlussky (1994a) placed the 22 exotic ant pansion on 'Eua appears to have occurred species found in Tonga in four categories, in just 15 yr. The tiny P. megacephala pop according to their presumed origins (Tables 2 ulation on Lifuka is probably even younger. Ants ofTonga . Wetterer 133 Although I found that P. megacephala was lian National Insect Collection in Canberra, relatively uncommon in Vava'u, Kapa, and need to be searched for additional unreported Lifuka, I predict that this ant will spread to ant specimens from Tonga to better under dominate these islands as well. stand the spread or loss of ant species. Three destructive tramp ants known in the Pacific that have not yet been found in Tonga are Linepithema humile (Mayr), Mono ACKNOWLEDGMENTS marium destructor (Jerdon), and Wasmannia auropunctata (Roger). In Hawai'i, Linepithema I thank Edward O. Wilson for first suggesting humile is the primary pest ant at higher that I study ants in Tonga, for advice and elevations (above 900 m) (Reimer 1994, encouragement, and for financing the mount Wetterer et al. 1998). It also has had a great ing of ant specimens; Wilco Liebregts and impact on the native ants of Santa Cruz the personnel of Vaini Research Station for Island in the California Channel Islands acting as my hosts in Tonga; Tony Matthias (Wetterer et al. 2000). Monomorium destructor of Toni's Guest House in Nuku'alofa for is known from many parts of the Pacific, in driving me around Tongatapu; Pila Kami for cluding Samoa, the Cook Islands, and French driving me around the west side of 'Eua; Polynesia (Wilson and Taylor 1967). Mayor Stephan and Daniela at the Hilltop Guest (1922:103) wrote of Monomorium destructor in house on Vava'u for lending me a bicycle and the Florida Keys: "So voracious are these in a kayak; Mingsheng Wang of the MCZ for sects that we are obliged to swing our beds mounting the ants; Stefan Cover of the MCZ from the rafters and to paint the ropes with for identifying the ants; Jo Berry at the New a solution of corrosive sublimate, while all Zealand Arthropod Collection, Mount Albert tables must have tape soaked in corrosive Research Center, for her hospitality and her sublimate wrapped around their legs if ants patience waiting for the return of specimens are to be excluded from them. These pests borrowed from her collection; and Roy have the habit of biting out small pieces of Snelling, Genady Dlussky, Andrea Wetterer, skin, and I have seen them kill within 24 and Margaret Wetterer for comments on the hours rats which were confined in cages." manuscript. Most ofthe geographic data used Perhaps the greatest threat, however, is in this paper carne from the United Nations Wasmannia auropunctata, which has become Environment Programme Islands Web Site established in several Pacific island groups, (UNEP 1998). including the Galapagos (Clark et al. 1982), Wallis and Futuna (Gutierrez 1981 in Jour dan 1997), New Caledonia (Jourdan 1997), Literature Cited the Solomon Islands (Fabres and Brown 1978), and most recently Vanuatu (Rapp Anonymous. 1999. New stinging little fire ant 1999) and Hawai'i (Anonymous 1999). In found on the Big Island and Kauai. areas where it invades, W auropunctata can be NASDA News 7:1. a significant agricultural pest, both through Bolton, B. 1995. A new general catalogue of stinging workers and through enhancing the ants of the world. Harvard University populations ofHomoptera (Spencer 1941). In Press, Cambridge. addition, W auropunctata has direct negative Carver, M., P. J. Hart, and P. W. 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