Taxonomic Identity of the Ghost Ant, Tapinoma Melanocephalum (Fabricius, 1793) (Formicidae: Dolichoderinae)

Home , Aneuretinae, Metasoma, Tapinoma, Tapinoma indicum, Tapinoma melanocephalum, Tapinoma minutum

Zootaxa 4410 (3): 497–510 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2018 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4410.3.4 http://zoobank.org/urn:lsid:zoobank.org:pub:8DB5214E-2CCB-4DC4-BCCE-DB5296E63B8B

Taxonomic identity of the ghost ant, Tapinoma melanocephalum (Fabricius, 1793) (Formicidae: Dolichoderinae)

ROBERTO J. GUERRERO Grupo de Investigación en Insectos Neotropicales, Programa de Biología, Facultad de Ciencias, Universidad del Magdalena, Carrera 32 # 22–08, Santa Marta, Magdalena, Colombia. E-mail: [email protected], [email protected]

Abstract

This paper revises the taxonomy of Tapinoma melanocephalum (Fabricius, 1793) as follows: T. melanocephalum = Tapi- noma luffae (Kuriam, 1955) syn. nov., = Tapinoma melanocephalum coronatum Forel, 1908 syn. nov., = Tapinoma melanocephalum malesianum Forel, 1913 syn. nov. A neotype of Tapinoma melanocephalum (Fabricius, 1793) is designed here. Lectotypes of Tapinoma melanocephalum coronatum Forel, 1908 and T. melanocephalum malesianum Forel, 1913 are designated. Formica wallacei is proposed as a replacement name for Formica familiaris (= T. melanocephalum senior synonym). The worker, queen and male are redescribed and diagnosed. The morphological variability of populations is discussed. All castes are included in full color images.

Key words: Cosmopolitan species, morphological variation, new synonymy, taxonomic stability, tramp species

Introduction

The ghost ant, Tapinoma melanocephalum (Fabricius, 1793) (Formicidae: Dolichoderinae), is one of the earliest described ant species. For more than two centuries, investigators have used body coloration as a diagnostic feature: black head, brown thorax paler below, pale abdomen (e.g. Nickerson & Bloomcamp 2012). However, this "diagnostic" characteristic does not strictly fit all populations throughout the wide distribution of the species. Color variability has led to the description of several varieties and subspecies (Forel 1908; Forel 1913; see Taxonomic synopsis) as well as nomenclatural inconsistencies within the nominal species. Tapinoma melanocephalum is a ubiquitous tramp species, found predominantly in tropical and subtropical areas, dispersed by human commercial activity. Wetterer (2009) mapped the known distribution of T. melanocephalum and concluded that it is one of the most widely distributed ant species worldwide. This species is apparently native to Southeast Asia or the Indo-Pacific region (Wetterer 2009), but populations have been able to live in almost all anthropogenic environments, although very few records come from natural or partially transformed environments within the New World tropics. Tapinoma melanocephalum also can become established within buildings in warm places in temperate latitudes. The ghost ant is considered to be an annoying pest, as it infests homes, offices, and hospitals, where it can act as a mechanical vector of human diseases (for example, in Colombia; Olaya-Másmela 2005). Considering the importance of this synanthropic species, it is essential to provide nomenclatural stability and to propose characteristics for the delimitation of this species of ant. I therefore examined specimens identified as T. melanocephalum and infraspecific categories deposited in museums and biological collections throughout the world. The study of the morphology allowed me to identify a series of taxonomically useful characters for all castes of T. melanocephalum which are discussed here.

Material and methods

I examined 959 specimens (915 workers, 27 queens and 17 males) identified as T. melanocephalum and associated

Accepted by J. Longino: 28 Feb. 2018; published: 18 Apr. 2018 497 names (i.e. subspecies). Of those, 84% of the material was pinned; the rest was preserved in ethanol (80-96%). I also studied type specimens of infra-specific taxa (T. melanocephalum var. coronatum Forel, 1908 and T. melanocephalum var. malesianum Forel, 1913 syntypes) through the loan of Auguste Forel's types from the Museum of Natural History (Geneva, Switzerland) to the laboratory of Dr. Philip S. Ward (University of California at Davis). The following are acronyms for the institutions whose material I examined. Those marked with an asterisk (*) are not registered in Evenhuis (2016).

BMNH The Natural History Museum, London, England. CAS California Academy of Sciences, San Francisco, California, USA. CBUM Centro de Colecciones Biológicas de la Universidad del Magdalena-CBUMAG, Santa Marta, Magdalena, Colombia. * CEUA Colección Entomológica de la Universidad de Antioquia, Medellín, Antioquia, Colombia. CPDC Centro de Pesquisas del Cacao, Comissão do Plano de Lavoura, Itabuna, Bahia, Brazil. IAVH Colección de Insectos, Instituto Alexander von Humboldt, Claustro de San Agustín, Villa de Leyva, Colombia. INPA Instituto Nacional de Pesquisas da Amazonia, Colección Sistemática de Entomología, Manaus, Amazonas, Brazil. ICNC Colección de Insectos, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D.C., Colombia. JKWC James K. Wetterer personal collection, Florida Atlantic University, Boca Raton, Jupiter, Florida, USA. * MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA. MEUV Museo de Entomología de la Universidad del Valle - MUSENUV, Cali, Colombia. MHNG Museum of Natural History, Geneva, Switzerland. MIZA Museo del Instituto de Zoología Agrícola, Universidad Central de Venezuela, Maracay, Aragua, Venezuela. MPEG Museu Paraense Emilio Goeldi, Belem, Pará, Brazil. MPUJ Museo Entomológico de la Universidad Javeriana, Bogotá D.C., Colombia. MZSP Museu de Zoología da Universidade de São Paulo, São Paulo, Brazil. PSWC Phillip S. Ward personal collection, University of California at Davis, CA, USA.* UCDC R.M. Bohart Museum of Entomology, Davis, CA, USA. UNAB Colección de insectos de la Facultad de Agronomía, Universidad Nacional de Colombia, Bogotá D.C., Colombia. UNCM Museo de Entomología "Francisco Luis Gallego", Universidad Nacional de Colombia, Medellín, Colombia. USNM National Museum of Natural History-Smithsonian Institution, Washington D.C., USA. WEMC William & Emma Mackay personal collection, University of Texas at El Paso, Texas, USA.*

Tapinoma ants have a very simple morphology (no spines, teeth, nor marked sculpture on the body) that render the separation and identification of the species difficult. To resolve the taxonomy of the ghost ant, I examined the specimens previously identified as T. melanocephalum, including workers, gynes and males. Morphological description and measurements of specimens were performed using a Nikon SMZ 740 binocular stereomicroscope equipped with a micrometer at magnifications of 96X. Morphometric characters were examined in each of the castes (workers, gynes, and males). All measurements were expressed in millimeters with an accuracy of 0.01. The following measurements and indices were used:

Head Length (HL). In full-face view, the maximum length taken from the posterior cephalic margin to the maximum projection of the anterior border of the clypeus. Head Width (HW). In full-face view, the maximum width of the cephalic capsule, recorded posterior to the compound eyes. Scape Length (SL). In full-face view, the maximum length of the first segment of the antenna (scape), excluding

498 · Zootaxa 4410 (3) © 2018 Magnolia Press GUERRERO the neck that occurs just distal of the condylar bulb. Where the scapes were not aligned with the frontal view of the head, it was necessary to place the cephalic capsule obliquely to take a correct measurement of the scape. Weber’s Length (WL). In lateral view of the mesosoma, an oblique line from the most anterior projection of the pronotum to the lower posterior corner of the metapleural sclerite.

Cephalic Index (CI). (HW/HL)*100. Scape Index (SI). (SL/HL)*100.

Digital color images were generated from ant types and additional specimens presented in this paper. Digital images of type material were taken from www.Antweb.org, while those of non-types and the neotype were generated with a Leica digital high resolution camera (Type DFC450), attached to a Leica M205FA automated stereomicroscope. Stacked images were processed with LAS montage module-Leica®. Saturation and brightness were adjusted in Corel PHOTO PAINT X8.

Results

Taxonomic synopsis

Tapinoma melanocephalum (Fabricius, 1793). = Tapinoma australis Sanstchi, 1928a = Tapinoma familiaris (Smith, F., 1860b) = Tapinoma luffae (Kuriam, 1955), New Synonym = Tapinoma melanocephalum coronatum Forel, 1908, New Synonym = Tapinoma melanocephalum malesianum Forel, 1913, New Synonym = Tapinoma nana (Jerdon, 1851) = Tapinoma pellucida (Smith, F., 1857)

Tapinoma melanocephalum (Fabricius, 1793) Figs. 1–28

Neotype worker (Figs 1–3, 10), by present designation. Colombia: Magdalena, Santa Marta, Calabazo, roadside to Pueblito-PNN Tayrona, nest in dead branch on the roadside, secondary dry forest (11.2825°N, -74.0239°W), 22.i.2017, leg. R. Guerrero, nest series, RJG01286 (ICNC-093716). Paraneotype workers (10), dealate queens (2), males (2), same data as neotype (1w, BMNH; 1w, 1dq, 1m CAS; 1w, 1q, ICNC; 2w, MCZ; 1w, MHNG; 2w, 1m, MZSP; 2w, USNM).

Formica melanocephala Fabricius, 1793:353. Lasius melanocephalus: Fabricius, 1804:417. Myrmica melanocephala: Lepeletier, 1836:185. Formica nana Jerdon, 1851:125. Syntype worker: India: Mysore. Name pre-occupied by Latreille, 1802:263. Synonymy by Emery, 1892:166. Myrmica pellucida F. Smith, 1857:71. Syntype workers: Singapore (BMNH; OXUM, University Museum of Natural History, Oxford, United Kingdom) [not examined]. Two syntype workers imaged on AntWeb: CASENT0903062 (BMNH), CASENT0901925 (OXUM). Synonymy by Mayr, 1886:359. Formica familiaris F. Smith, 1860b:96. Syntype worker: Indonesia: Bacan Island. Synonymy by Forel, 1899:101. Junior homonym of Formica familiaris F. Smith, 1860a:68. Replacement name Formica wallacei, new name. Tapinoma melanocephalum: Mayr, 1862:651; Emery, 1887:249 (male); Forel 1891:102 (description of queen); Forel, 1895:472 (re-description of worker); Kempf, 1972:247; Shattuck, 1994:148; Bolton et al. 2007: digital media (CD); Bolton, 2017: e-catalog (www.antcat.org). Micromyrma melanocephala: Roger 1862:258 (queen, combination in Micromyrma).

TAXONOMY OF THE GHOST ANT Zootaxa 4410 (3) © 2018 Magnolia Press · 499 Tapinoma (Micromyrma) melanocephalum: Wheeler, 1922:923. Tapinoma melanocephalum var. coronatum Forel, 1908: 62. Lectotype worker (Figs 4–6, 11), by present designation: Costa Rica: Puntarenas (middle specimen on the pin, MNHG, AntWeb image and specimen identifier CASENT0909775) [examined]; two paralectotype workers: upper and lower specimen on the same pin as the lectotype. Kempf, 1972:247; Shattuck, 1994:148; Bolton et al. 2007: digital media (CD); Bolton, 2017: e-catalog (www.antcat.org). New Synonym. Tapinoma melanocephalum var. malesianum Forel, 1913:93. Lectotype worker (Figs 7–9, 12), by present designation: Indonesia: Sumatra, Soengei Bamban (MNHG, AntWeb image and specimen identifier CASENT0909776) [examined]; paralectotypes: same data as lectotype (45 workers, 6 queens in MNHG, 1 worker in NHMB). Shattuck, 1994:148; Bolton et al. 2007: digital media (CD); Bolton, 2017: e-catalog (www.antcat.org). New Synonym. Tapinoma (Micromyrma) melanocephalum var. australis Sanstchi, 1928a:53. Syntypes (23 workers), Vanuatu: Santo Island, Hog Harbour (BMNH, Natural History Museum, London, U. K; NHMB, Natural History Museum, Basel, Switzerland) [not examined]. Two syntype workers imaged on AntWeb: CASENT0903063 (BMNH), CASENT0911572 (NHBM). Synonymy by Wilson & Taylor, 1967:80. Bolton et al. 2007: digital media (CD); Bolton, 2017: e-catalog (www.antcat.org). Although it appears Santschi intended australe to be the primary description and australis a redescription, australis was published first and has priority. Tapinoma (Micromyrma) melanocephalum var. australe Santschi, 1928b:475. Redescription and an incorrect spelling of australis. Shattuck, 1994:148; Bolton et al. 2007: digital media (CD); Bolton, 2017: e-catalog. (www.antcat.org). Neoclystopsenella luffae Kurian, 1955:133. Holotype male (described as a Bethylidae): India: Delhi, Kapoor Coll., 2-7-1938, on Luffa, Imperial Entomologist (type material lost). New Synonym. Tapinoma luffae: Brown, 1988:337; Shattuck, 1994:140; Bolton & al 2007: digital media (CD); Bolton, 2017: e- catalog (www.antcat.org).

Neotype measurement: HL: 0.51 mm, HW: 0.45 mm, SL: 0.47 mm; WL: 0.44 mm; CI: 88; SI: 92. Worker measurements (n= 316): HL: 0.46 (0.38–0.60) mm, HW: 0.41 (0.35–0.60) mm, SL: 0.43 (0.35–0.52) mm; WL: 0.39 (0.28–0.60) mm; CI: 89 (80–104); SI: 94 (82–105). Worker diagnosis. Bicolored small ants, with head and mesosoma black to brown, mandibles, antennae, legs and gaster light brown to pale yellow; third and fourth segment of the maxillary palps broad and flattened, spatula- like; basal end of the fifth segment inserted into the ventral face of the fourth segment. Worker description (Figs 13–14): Head. Ovoid, longer than wide with slightly convex lateral margins; posterior margin variable, convex to straight, or sometimes slightly emarginated medially, in all cases, with rounded posterolateral corners. Mandibles subtriangular, straight outer margin over 2/3 length, and distally curved inwards; masticatory margin consisting of strong apical tooth, acute and longer than wide; slender subapical tooth, half length of the apical tooth; third triangular tooth half as long as subapical; fourth tooth slightly longer than third, followed by row of tight denticles decreasing in size approaching curvature leading to undifferentiated basal margin, the latter completely smooth or slightly crenulated. Palpal formula 6:4; maxillary palps with the first segment small and cylindrical, second segment conical and wider distally, sixth segment elongated, digitiform, rounded at tip; segments of labial palps relatively similar in size, 1st and 2nd segments wedge-shaped, 3rd subrectangular, last segment oval, longer than wide, with rounded tip. Clypeus broader than long; anteromedial margin with slight depression, and narrow, barely noticeable; margins lateral to depression relatively straight, those more lateral obliquely straight; posterior margin of clypeus convex, almost reaching base of frontal carinae. Frontal carinae diverging distally. Antennal scapes long, surpassing posterior margin of head by distance similar to length of pedicel, outer margins straight, slightly curved at both base and apex, wider near apical end; pedicel slightly conical with sub-parallel sides, third segment twice as wide as long, fourth-eleventh segments rectangular, much longer than wide, gradually elongating towards apex, last segment twice as long as others, rounded at tip. Eyes relatively large, oval, longer than wide, external margin relatively straight while the internal margin strongly convex, distance from the mandibular insertion approximately 1/2 maximum length of the eye. Mesosoma. In lateral view, promesonotum strongly convex, forming nearly continuous line; metanotal sulcus broad and relatively deep; dorsopropodeum strongly convex, dome-shaped, at same level or slightly below level of promesonotum, declivitous face straight, inclined between 50–60º. Propodeal spiracle nearly circular, slightly longer than wide, opening laterally, well below half-length of lateropropodeum.

500 · Zootaxa 4410 (3) © 2018 Magnolia Press GUERRERO Metasoma. In lateral view, petiole short, flattened dorsal face with sharp apex inclined anteriorly; ventral lobe projecting, with straight anterior face and slightly curved posterior face. First tergum of gaster excavated anteriorly, as in other Tapinoma species. Body surface and pilosity. Cephalic surface opaque, finely covered with punctures and reticulations forming very small alveoli; abundant pubescence very short. Mandibles relatively smooth and slightly opaque, dorsal surface tenuously foveolate, covered with abundant short suberect hairs, and longer suberect hairs towards middle and basal portion; ventral face with long erect hairs. Dorsal face of clypeus covered with pubescence similar to that of cephalic capsule; anterior clypeal margin with two longer and thicker hairs, one on either side of medial depression, two golden relatively shorter setae than previously described hairs, above the clypeal margin are a pair of thick hairs directed anteriorly. Mesosoma completely covered with fine reticulations forming alveoli, relatively larger than those of head; surface rough and opaque, with dense whitish appressed hairs, very short and separated, sometimes imperceptible, giving appearance of glabrous surface, only visible in oblique-dorsal position. Legs with relatively fine granular and reticulated sculpture, relatively opaque, completely covered with dense whitish pubescence.

FIGURES 1–12. Tapinoma melanocephalum multifocus images. 1–3) Full face view of the head, profile, and dorsal view of the T. melanocephalum neotype worker; black arrow shows the enlarged segments of the maxillary palps. 4–6) Full face view of the head and profile of T. melanocephalum coronatum lectotype worker. 7–9) Full face view of the head and profile of T. melanocephalum malesianum lectotype worker. 10–12) Labels of T. melanocephalum neotype, T. melanocephalum coronatum and T. melanocephalum malesianum lectotypes, respectively; lectotypes of T. melanocephalum coronatum and T. melanocephalum malesianum were designated after the AntWeb team imaged those specimens. Photos 1–2 by Tania K. Franco, and photos 3 and 10 by Emira García; photos 4–9, and 11–12 by Will Ericson from www.AntWeb.org.

TAXONOMY OF THE GHOST ANT Zootaxa 4410 (3) © 2018 Magnolia Press · 501 FIGURES 13–18. Tapinoma melanocephalum worker, gyne and male habitus. 13–14) Full face view of the head and profile of the worker [Colombia: Magdalena, Santa Marta, dry forest plot, (11.2219°N -74.1865°W), 10.x.2015, leg. R. Guerrero, CBUM, Catalogue # CBUMAG:ENT:08463]. 9–10) Full face view of the head and profile of the gyne [Colombia: Magdalena, Santa Marta, dry forest plot, (11.2219°N -74.1865°W), 10.x.2015, leg. R. Guerrero, CBUM, Catalogue # CBUMAG:ENT:08464]. 11–12) Full face view of the head and profile of the male [Paraneotype: Colombia: Magdalena, Santa Marta, Calabazo, roadside to Pueblito-PNN Tayrona, nest in dead branch on the roadside, secondary dry forest (11.2825°N, - 74.0239°W), 22.i.2017, leg. R. Guerrero, nest series, RJG01286, MZSP]. All images by Mayron E. Escárraga and Tania K. Franco.

Petiole smooth and shiny, with few short, decumbent hairs near posteroventral margin. Gaster smooth to finely reticulated, relatively shiny, covered with fine dense golden yellowish pubescence; 5–6 erect hairs near posterior margin of fourth tergum, other terga without erect hairs; sternites with two erect hairs per segment. Coloration. Bicolored, variable in color tone. Head and mesosoma intense dark brown to brown with some regions yellowish brown on the propleura and the dorsum of the promesonotum. Mandibles, maxillary and labial palps, scapes, antennal segments, and legs pale yellow; masticatory margin reddish brown. Gaster brown to pale yellow as legs, but sometimes with contrasting brown bands on posterior margins of each tergum. Gyne measurements (n= 20): HL: 0.52 (0.43–0.60) mm; HW: 0.51 (0.41–0.60) mm; SL: 0.45 (0.35–0.52) mm; WL: 0.74 (0.59–0.83) mm; CI: 93–104; SI: 81–93. Gyne diagnosis: Cephalic capsule diverging posteriorly, much wider posterior to the compound eyes than anterior to them. Second segment of maxillary palps triangular, flattened, longer than wide, 3rd and 5th segments much broader and flattened, 3rd and 4th segment almost fused at the ends where they meet, the basal end of the 5th segment inserted into the ventral face of the 4th segment. Gyne description (Figs 15–16): Head. Capsule cephalic square to rectangular (CI: 93–104), cuneiform; sides

502 · Zootaxa 4410 (3) © 2018 Magnolia Press GUERRERO of the capsule relatively straight, lateral margin interrupted by the outer margin of the eyes; well-defined posterolateral corners, rounded, continuing to straight posterior cephalic margin, with shallow and wide concavity. Mandibles as in workers. Palpal formula 6:4; first segment of maxillary palps cylindrical and thickened, last segment thickened, digitiform, rounded at tip; segments of labial palps as in workers, but relatively broader and flattened. Dorsal face of clypeus as in workers, except for anterior margin relatively straighter and entire, with a slight mesal concavity. Frontal carinae divergent posteriorly. Scapes long, surpassing posterolateral corners of head by distance similar to length of pedicel; outer and inner margins straight in most of the length but slightly curved distally, wider mid-length; antennal segments as in workers. Compound eyes large, located at middle of length of cephalic capsule, relatively close to clypeal insertion; eyes located laterally, external margin breaking border of the cephalic capsule. Ocelli small, near level of posterior border of head; posterior ocelli near vertex but never breaking vertexal line. Mesosoma. In profile, pronotum broad, propleura relatively broader than anterior part of pronotum; dorsal surface of mesonotum flattened; metanotum consisting of very thin strip, separated by deep mesonotal groove; dorsopropodeum without distinctive face, bending abruptly on a relatively inclined slope; mesopleuron divided into anepisternum and katepisternum, those divided by marked and relatively deep anapleural sulcus, anepisternum inverted U-shape, katepisternum subquadrate, almost as long as wide; propodeal spiracle circular, with an outer rim like a ring, directed posterad, and located on lateropropodeal face; metapleuron protruding, forming slight depression below propodeal spiracle. In dorsal view, mesoscutum strongly convex anteriorly, with parallel sides, diverging slightly posteriorly; mesonotal groove straight, slightly marked; mesoscutellum wider than long, posterior margin convex; dorsopropodeum restricted to very narrow band, posterior margin rounded bended abruptly on a slope, propodeum with the lateral margins, almost parallel. Metasoma. Petiole relatively large, longer than wide, petiolar scale very low, restricted to small dorsal projection, inclined, directed anteriorly, with poorly developed lateral carinae; developed ventral lobe, most prominent posteriorly. Gaster as long as the mesosoma, flattened dorsally and slightly convex on ventral face. Body surface and pilosity. Surface of the head dense and finely punctate; with dense whitish decumbent hairs relatively separate, distance between hairs less than individual length. Dorsal face of mandibles densely covered with short golden hairs, on almost entire surface, except near area behind masticatory margin where they are long and inwardly directed. Maxillary palps covered with very short pilosity; labial palps with short erect and suberect hairs. Clypeus with pubescence as remainder of head, pattern of hairs on anterior margin as described in workers. Mesosoma opaque, abundantly and finely punctate; lateral regions of mesosoma covered with whitish pubescence, relatively longer than those of head, more abundant on katepisternum and the metapleuron; dorsum of the mesosoma covered with short pubescence, widely spaced, leaving surface partially exposed. Legs covered with short and decumbent hairs. Metasomal surface finely punctate as mesosoma, but relatively brighter; covered with abundant pubescence; fourth tergite with six long erect hairs near posterior margin, one central pair, and other two towards sides of the tergite. Color. Head brown to dark brown, mandibles, maxillary and labial palps and antennae pale yellow; mesosoma bicolored, part of mesoscutum, mesoscutellum and legs yellowish brown to pale yellow, dorsum of pronotum and central portion of mesoscutum as spot, mesopleuron, inferior part of propodeum, and metapleuron brown. Petiole yellowish; gaster brown with yellowish brown bands on posterior margin. Male measurements (n= 17): HL: 0.44 (0.43–0.46) mm; HW: 0.42 (0.41–0.45) mm; SL: 0.39 (0.35–0.43) mm; WL: 0.66 (0.59–0.73) mm; CI: 93–104; SI: 81–93. Male diagnosis: Third and fourth segment of the maxillary palps widened, oblong in shape, much longer than wide; the basal end of the 5th segment inserted into the ventral face of the 4th segment, like in the female caste. Male description (Figs 17–18): Head. In dorsal view, head as long as broad, largest width posterior to compound eyes, with lateral margin convex, cephalic capsule converging anteriorly, posterolateral corners rounded, continuous with straight posterior cephalic margin; ventral surface of cephalic capsule relatively flattened. Mandibles long, semifalcate; external margin slightly sinuous, curving inwards distally; masticatory margin straight, with apical tooth relatively large, obliquely directed downwards, subapical denticle overhanging from margin, remainder of margin with very tight denticles, same size, continuing towards the undifferentiated basal margin. Palpal formula 6:4; 1st segment cylindrical, 2nd segment two times longer than first, cuneiform, 4th segment much wider than 3rd, 5th and 6th segments finger-shaped, the apical with rounded tip. Clypeus broad,

TAXONOMY OF THE GHOST ANT Zootaxa 4410 (3) © 2018 Magnolia Press · 503 relatively high, dorsal face outwardly projected; anteromedial border slightly concave, narrow, with rounded corners continuing toward relatively straight anterior margin, anterolateral margins inclined posteriorly; posterior margin convex but posterior vertex relatively flattened. Frontal carinae straight and semi-parallel at base, curved and diverging distally. Scape long, surpassing posterior cephalic margin by distance slightly less than distance between frontal carinae, scape with lateral margins parallel, outwardly curved from base to medial length, straight for rest of length, pedicel cuneiform, narrower at base, remainder of antennal segments cylindrical, longer than broad, last segment almost twice length of others. Compound eyes very large, oval, two times longer than broad, occupying almost entire lateral portion of head, breaking outline of lateral margins; anterior margin of eye separated from clypeal insertion by distance shorter than minimal width of scape. Ocelli large, located on wide base and relatively higher than dorsal surface of head, slightly below posterior cephalic vertex; anteromedial ocellus relatively smaller than lateral ocelli.

FIGURES 19–22. Head of Tapinoma melanocephalum castes showing the shape of the maxillary palps. 19–20) Polymorphism in the maxillary palps of the workers [Colombia: Magdalena, Santa Marta, dry forest plot, (11.2219°N -74.1865°W), 10.x.2015, leg. R. Guerrero, CBUM, CBUMAG:ENT:08464 and CBUMAG:ENT:08465]. 21–22) Widened segments of the maxillary palps of queen and male [both reproductive castes with same data as workers above, CBUM, CBUMAG:ENT:08466 and CBUMAG:ENT:08467, respectively]. Black arrows indicate fifth maxillary palp inserted in the inner face of fourth maxillary palp. All images by Mayron E. Escárraga.

Mesosoma. In profile, propleura broad, flattened, with dorsal border thin, projecting dorsally; mesonotum anteriorly curved, with hump in lateral view, dorsal face relatively straight and high; mesopleuron divided into anepisternum and katepisternum, both delimited by wide and relatively deep anapleural sulcus; anepisternum inverted U-shaped, relatively wider than long, with dorsal margin slightly inclined posteriorly, anterior margin longer than posterior; katepisternum semiquadrate. Dorsopropodeum below level of mesonotum, with long surface, posteropropodeum slightly bulging decline of 90° toward base; propodeal spiracle oval, longer than broad, located on lateropropodeum, opening posteriorly. In dorsal view, anterior line of pronotum visible; mesoscutum slightly triangular with inferior margin truncated, longer than broad; dorsal face of propodeum visible.

504 · Zootaxa 4410 (3) © 2018 Magnolia Press GUERRERO FIGURES 23–28. Forewings of Tapinoma spp males. 23) Tapinoma melanocephalum [Ecuador: Galápagos, Santa Cruz, Bella Vista, 0°41’38.1’’S 90°19’16.8’’W, 180 m, 13.x.2007, H.Herrera#HWH197, ICCDRS0002606 on www.AntWeb.org. 24) Tapinoma luffae male (= T. melanocephalum senior syn.) modified from Kurian (1955). 25) Tapinoma sessile [Mexico: 26 Km N J. María Aguirre, no more data, 20.v.1988. leg. W.P. Mackay, WEMC#10289]. 26) Tapinoma nigerrimum [Spain: Pozuelo, syntype specimen at Naturhistorisches Museum Basel (NHMB, Basel, Switzerland-), CASENT0911578]. 27) Tapinoma sp. [Brazil: Bahia, Ilhéus, Teotônio Vilela, 06.v.1998, leg. S.R.M. Santos, CPDC, no catalogue number]. 28) Tapinoma subtile [Madagascar: Antsiranana, Rés. Analamerana, 16.7 km 123° Anivorano-Nord, -12.80467°S, 49.37383°E, 225m, 4.xii.2004, leg., B.L. Fisher, BLF11379, CASENT0053839]. Black arrow indicates 1m-cu closing to form the discal cell-1; white arrow indicates 1m-cu absence. Note the wide reduction of venation in T. subtile. Photos 23 and 26 by April Nobile and Zach Lieberman, respectively, from www.AntWeb.org. Photos 27–28 by Mayron E. Escárraga. Name of veins and cells are those of Yoshimura & Fisher (2011).

Metasoma. In lateral view, petiole with petiolar scale well developed, anterior face relatively high and straight, forming angle with oblique dorsal face that descends posteriorly to declivity; ventral lobe developed, slightly convex. Cupula weakly sclerotized, easily collapsible when specimen pinned; telomeres dorsoventrally broad, obliquely truncated at apex; digitus long and narrow to tip, latter ventrally curved, cuspis located well inside genital capsule, semi-rectangular in lateral/medial view. Body surface and pilosity. Head surface opaque, softly punctulate; abundant, whitish, appressed hairs. Mandibles smooth, shining, bearing scattered short and long hairs, subdecumbent from base to behind masticatory margin; some hairs long, surpassing external margin. Anterior margin of clypeus with same pattern of hairs as described in worker; more lateral portions with relatively larger curved hairs; dorsal face densely hairy. Scape surface opaque, slightly punctate, covered by very short pilosity, with some hairs relatively longer and suberect in

TAXONOMY OF THE GHOST ANT Zootaxa 4410 (3) © 2018 Magnolia Press · 505 lateral margins; antennal segments with surface granulate, densely covered by short decumbent hairs, with some short suberect hairs. Mesosoma relatively shining, with some areas notably opaque, puncticulate; surface covered by relatively short dense golden scattered pubescence, more abundant on katepisternum and metapleuron. Gaster reticulate, with small alveoli; covered by dense golden pubescence, but relatively sparse, leaving uncovered surface; telomeres densely covered by relatively long pilosity, surpassing margin. Color. Ants completely yellowish brown, except for mandibles, scape and antennal segment, and legs which are whitish, contrasting with the remainder of body. Larvae: Dolichoderine larvae described by Wheeler and Wheeler (1951), and examined in detail by Jesus et al. (2010). Distribution: Species of worldwide distribution, especially in humid tropical and subtropical regions in both hemispheres, but also in temperate places (Wetterer 2009). It is widely distributed within the Neotropical region, except for Chile. Biology: The ghost ant biology is reviewed in Wetterer (2009), however, I provide here some additional information. Tapinoma melanocephalum exhibits a wide altitudinal distribution. Populations can be found from environments at sea level to almost 2700 m. Some records in Central America and Caribbean islands suggest that this species is generally distributed below 300 m, except for a record from Panama at 1220 m. In contrast, the distribution of populations established in South America is wider, ranging from 0 m to 2700 m; the highest elevations are recorded for the Colombian Andes, especially towards the center of that country (Antioquia, Cundinamarca, Risaralda, and others). In the center and south of South America, most records correspond to altitudes below 600 m (e.g., São Paulo and Rondônia in Brazil). Tapinoma melanocephalum is a polygynous species with 1–25 queens per colony (Bustos & Cherix 1998). In some colonies studied in Santa Marta (Colombia) 2–5 queens were found, typical of the reproductive behavior of other invasive dolichoderine species such as Linepithema humile, Technomyrmex albipes, and other widely distributed Tapinoma species (Guerrero, unpublished data). One of these colonies (collected in October 2015, RJG01263) harbored two queens, 86 workers, 22 larvae and 10 pupae (3 males and 7 workers). Recently, I collected two colonies of T. melanocephalum near the Tayrona National Park (Santa Marta, Colombia: RJG01286 and RJG01289). One of those colonies (RJG01286, neotype series) was nesting in a dead branch on the roadside to Pueblito-Parque Nacional Natural Tayrona, and another one inside a dry branch hanging from the vegetation on the riverbank of a stream. In both cases, the colonies contained dozens of workers, with two queens, many eggs, larvae and worker pupae, but few males. In addition to multiple queens, colonies can also be composed of workers with relatively modified morphology. A colony collected in Honduras (Cortés, La Lima: MZSP, catalogue # 6878) contained intercaste females. One of intercastes has a pair of vestigial mesonotal wings; one has a forewing on the right side; two have an enlarged mesonotum and wing scars; one has wing primordia on the mesonotum.

Discussion

Tapinoma melanocephalum is distinguishable from other species of Tapinoma in all biogeographic regions, although there are other species with similar coloration (e.g. Tapinoma atriceps Emery, 1888). Both workers and reproductive castes (gynes and males) share the widening of the 3rd and 4th segment of maxillary palps (Figs 19, 21–22). However, in the worker caste, this character may be intra-colonially dimorphic (Fig. 19–20), that is, some nest-mates can have all segments of the maxillary palps elongate, cylindrical to subcylindrical in shape, while most nest-mate workers have the typical state (widened segments). Nevertheless, in either of the two states the 5th segment of the maxillary palp is inserted into the inner face of the 4th segment, and not at the apex as in any of the other known species of Tapinoma. The latter is constant throughout the species (i.e. in examined specimens from anywhere in the world). Despite the polymorphism in the workers, gynes and males have both the 3rd to the 4th segment widened and flattened (Figs 21–22). Moreover, males have a conspicuous discal cell in the forewing (Figs 23–24), a feature shared with males of Nearctic species (e.g. T. sessile) and Palearctic species (e.g. T. madeirense). In T. madeirense the cell is rectangular (Figs 25–26), relatively longer than wide, while in T. melanocephalum it is relatively square. Additionally, males of T. sessile and T. madeirense are much larger and more robust than T.

506 · Zootaxa 4410 (3) © 2018 Magnolia Press GUERRERO melanocephalum, and dark brown to reddish brown in color. The males of Neotropical and Malagasy species have no discal cell in the forewing (Figs 27–28) (Guerrero unpublished). Tapinoma melanocephalum may be confused with other Tapinoma species in Asia and Oceania. Clouse (2007) found that in Micronesia there are intermediate forms between the species T. melanocephalum and Tapinoma minutum Mayr, 1862, which he referred to as Tapinoma A (color like minutum, scapes like melanocephalum) and Tapinoma B (color like melanocephalum, scapes like minutum). Despite the intermediate forms between T. melanocephalum and T. minutum in Micronesia, Sarnat & Economo (2012) found that in Fiji these two species can be distinguished by color and differences in the length of the scape. I studied several T. melanocephalum and T. minutum from Fiji (Viti Levu), finding that some measures both in T. minutum non-type specimens (n=4, HL= 0.44–0.46 mm, HW= 0.38–0.40 mm, SL= 0.32–0.34 mm) and the T. minutum type (HL= 0.53 mm, HW= 0.46 mm, SL= 45 mm; measured in the image on www.AntWeb.org: CASENT0915549) fit the measurement ranges of Tapinoma melanocephalum, consequently, the length of the scape would not be useful to separate the species. Despite this overlap in measurements, the shape and length of the maxillary palps, and the colour between both species (concolour in Tapinoma minutum) are very different, hence I used those characters to assign them unequivocally to either T. minutum or T. melanocephalum. On the other hand, Wetterer (2009 and references there) suggests that T. melanocephalum could also be confused with Tapinoma indicum Forel, 1895, pointing out that the descriptions of T. melanocephalum malesianum and T. indicum are very similar. I did not get access to specimens of T. indicum, therefore I could not directly study this species, only the images on www.AntWeb.org (CASENT0249761, CASENT0909774-Type, CASENT0903073-Type, and CASENT0911571-Type), and thereby I have no measurements to compare. However, the diagnostic characteristics of T. melanocephalum worker (shape of the 3rd and 4th segment of the maxillary palps and the insertion of the 5th segment with respect to the anterior segment) resolve the ambiguity in the identification with these other species (T. minutum and T. indicum) or intermediate forms. Tapinoma melanocephalum workers exhibit some degree of variability between and within the colonies. In general, the cephalic capsule is ovoid, however, some workers have a relatively straight vertex with a shallow concavity, or sometimes with a slightly emarginated vertex. The pattern of coloration is variable within the workers. While most of the workers in a population have completely brown heads and dark brown mesosomata, it is possible to find workers with a wide stain or yellowish brown "crown" on the dorsum of promesonotum (such as T. melanocephalum coronatum Forel), sometimes extending variably towards the mesopleuron. In these ants it is common to find brown bands on the gaster, which alternate with pale yellow bands. Occasional specimens (CASENT0903063) have the head and mesosoma lighter colored, which could be due to workers being callow, or fading of aged or exposed specimens, or some other factor causing intraspecific variation. The reproductives (gynes and males) are more homogeneous in metric characters and relatively variable in the coloration of the body, especially the gynes. The latter have rectangular heads (CI >93), and the scapes always exceed the vertices by a similar or slightly greater length than the pedicel of the antenna; the gynes may vary from yellowish brown to reddish brown (Figs 15–16). In males, the head posterior to the compound eyes is always semicircular in full-face view, slightly longer than wide, and the body is completely light brown (Figs 17–18). The phenotypic variability of workers, especially associated with body color, was used by several authors to propose infraspecific taxa, which are abandoned here and proposed as junior synonyms of Tapinoma melanocephalum. Prior to this study, five of these names were proposed as synonyms (see Taxonomic synopsis of T. melanocephalum). In this revision, I propose that T. melanocephalum coronatum, T. melanocephalum malesianum and Tapinoma luffae are junior synonyms of T. melanocephalum. Tapinoma melanocephalum coronatum and T. melanocephalum malesianum type series (both of MHNG) were examined and measured. I did not find differences between the workers of these nominal subspecies (i.e. type specimens) and the workers identified as Tapinoma melanocephalum. The head size, scape length and mesosoma length of both T. melanocephalum coronatum and T. melanocephalum malesianum fit within range of variation of Tapinoma melanocephalum. No other morphological differences were found. The type specimens of each of these subspecies do not have elongated and dilated maxillary palps, however the 5th segment of the maxillary palp is inserted into the inner face of the 4th segment; cylindrical maxillary palps in T. melanocephalum coronatum and T. melanocephalum malesianum is considered here as part of the polymorphism of that trait in T. melanocephalum workers (see discussion above). Both forms have workers with a light colored promesonotal crown, much more

TAXONOMY OF THE GHOST ANT Zootaxa 4410 (3) © 2018 Magnolia Press · 507 extended laterally in T. melanocephalum coronatum. This coloration has been observed in specimens collected elsewhere, typically within series that also have workers with the typical coloration pattern (head and mesosoma dark brown), and the typical character-state of the maxillary palps of T. melanocephalum. Tapinoma luffae (Kurian) was previously described as a species of Bethylidae of the genus Neoclystopsenella. Brown (1988) determined on the basis of Kurian's (1955) description that the males correspond to ants of the genus Tapinoma, proposing Neoclystopsenella as synonymous with Tapinoma and the combination Tapinoma luffae (Kurian). The Tapinoma luffae (Kurian) type is lost, but the illustrations of the forewings (Figure 255 in Kurian 1995:155) show the small typical discal cell in T. melanocephalum males (Fig 24). Tapinoma luffae is thus proposed as a junior synonym of T. melanocephalum. Tapinoma melanocephalum (Fabricius, 1793) is a species described more than 220 years ago, and apparently the Fabricius types are lost. Although most types of ants of the species described by Fabricius are hosted in the “Zoological Museum” (Copenhagen, Denmark), those of T. melanocephalum are not (Dr. Lars Vilhelmsen, pers. comm.). Dr. Vilhelmsen indicated that this material was deposited in the ‘Mus. Dom. Bosc.’, one collection that was later incorporated into the Museum of Paris (today, Muséum national d'histoire naturelle, Paris, French- MNHN). Since there are only a few types of Hymenoptera in the MNHN from the Bosc collection (Zimsen 1964:17), the database of the MNHN arthropod collection was consulted to search for the “Formica melanocephala” type, but the results were unsuccessful. Subsequently the types were searched for directly by the Collection Manager at MNHN (Agnièle Touret-Alby) and were not found. The type material of T. melanocephalum is thus assumed lost, necessitating the designation of a neotype. Tapinoma melanocephalum was described from Cayenne, French Guiana, while the neotype is from a coastal region of Colombia. This is counter to a strict following of article 75.3.6 of the ICNZ (ICNZ 1999), but there is a reason for the selection of the Colombian material. In the museums visited, there was very little material of T. melanocephalum from French Guiana, and no large series containing multiple castes. In contrast, a large series collected in northern Colombia, with all castes, was available for examination and distribution.

Acknowledgments

Many thanks to the two anonymous reviewers and Jack Longino (subject editor) for their comments and suggestions that greatly improved this manuscript. I am very grateful to each of the curators who kindly sent or allowed me to study the Tapinoma ants in the collections they hold: Ana Harada, Bernard Landry, Beto Brandão, Bill Mackay, Claudia Martínez, Fernando Fernández, Inge Armbrecht, Itana Fernandes, Jack Longino, Jacques Delabie, James Montoya, James K. Wetterer, John E. Lattke, Marta Wolff, Miguel Vásquez Bolaños, Patricia Chacón, Phil S. Ward, Rodrigo M. Feitosa, Rogerio R. da Silva and Ted S. Schultz. Thanks to Bill Mackay and John E. Lattke by helping me to improve both the English and early version of this manuscript. Fernando Fernandez helped me to improve the first draft. I want to thank James K. Wetterer and Brendon Boudinot for helping me to improve the English of the final version; they also suggested changes that improved the content of the manuscript. I am so grateful to Lars Vilhelmsen (Zoological Museum, Copenhagen, Denmark) and Agnièle Touret-Alby (Collection Manager, MNHN) for collaborating in the look for the types of Tapinoma melanocephalum. Thanks to Bernard Landry for sending some of the ant types of Forel from the MHNG to the lab of Phil S. Ward (UC at Davis, CA, USA). I also give thanks to Phil S. Ward and Ted S. Schultz for allowing me to use the optic equipment in their labs and other resources. Thanks to the Multifocus Image Team for the T. melanocephalum neotype images. Emira García measured many of the specimens examined here, and Lina María Ramos-Ortega helped by editing the examined material and literature cited sections. James Wetterer kindly sent me numerous T. melanocephalum specimens and geographic data. My visit to USNM was partially supported by The Ernst Mayr Animal Systematics Travel Grants of the Museum of Comparative Zoology of Harvard University and University of Magdalena. This is a contribution derived from the research project “Systematics and Evolution of the Ant Genus Tapinoma Föerster (Formicidae: Dolichoderinae) in the Neotropical Region” supported by Colciencias-Universidad del Magdalena through agreement #FP44842-008-2015 (code 1117-658-42796). This is Scientific Contribution No. 12 from the Centro de Colecciones Biológicas—Universidad del Magdalena.

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