Pachycondyla apicalis (Latreille)

Wild, A. L., 2005, Taxonomic revision of the Pachycondyla apicalis species complex (Hymenoptera: Formicidae)., Zootaxa 834, pp. 1-25 : 5-9

Pachycondyla apicalis (Latreille)

(Figs. 1, 2, 7, 10)

Formica apicalis Latreille 1802: 204.

Formica flavicornis Latreille 1802: 202. Synonymy by Brown (1957).

Pachycondyla apicalis (Latreille 1802); Mayr 1863: 439. First combination in Pachycondyla .

Pachycondyla flavicornis (Latreille 1802); Emery 1890: 58.

Neoponera flavicornis (Latreille 1802); Emery 1901: 47. First combination of flavicornis sensu Emery in Neoponera ; also first combination of apicalis sensu Emery (a misidentification of verenae Forel 1922) in Neoponera .

Neoponera latreillei Forel 1905: 161. Replacement name for Formica flavicornis Latreille 1802, j. hom. of Formica flavicornis Fabr. 1798. Synonymy by Brown (1957).

Neoponera obscuricornis r. latocciput Forel 1921: 132. NEW SYNONYMY

Neoponera obscuricornis latreillei (Forel 1905); Wheeler and Wheeler 1952: 613-615. Description of larva.

Neoponera apicalis (Latreille 1802); Brown 1957: 230; Kempf 1972: 161 (part); Fresneau 1985: 109-166; Fresneau and Dupuy 1988: 1389-1399.

Pachycondyla obscuricornis ; Reiskind 1977: 2-6. Not Emery (1890). Misidentification.

Pachycondyla apicalis (Latreille 1802); Goss et al 1989: 65-69. Revived combination in Pachycondyla , first use of implicit combination by Brown (1973).

Pachycondyla apicalis (Latreille 1802); Hölldobler and Wilson 1990: 385; Oliveira and Hölldobler 1990: 383-393; Soroker et al 1998: 1077-1090; Deitemann and Peeters 2000: 223-228; Longino2004.

Pachycondyla apicalis (Latreille 1802); Brown, in Bolton 1995: 302. Stated as “revived combination.”

Type data: Formica apicalis Latreille South America [type not located] . Formica flavicornis Latreille French Guiana. “ Cayenne ” [type not located] . Neoponera obscuricornis r. latocciput Forel : Ecuador. “Quito” (approx. loc.) [3w SYNTYPES, MHNG, examined] .

Other material examined:

Bolivia. Santa Cruz: 10k NW Terevinto [ PSWC] . Brazil. Amazonas: Ilha de Curari [ LACM] ; Ypiranga, R. Ica-Putomayo [ MCZC] ; 300k E Humaita, Transamazonica Hwy [ PSWC] ; Ponta Negra, N of Manaus [ MCZC] . Bahia: CEPEC/CEPLEC, Rodovia Ilheus / Itabuna [ ALWC, MCZC] . Mato Grosso: Chapada dos Guimaraes [ PSWC] . Pará: Santarem, Taperinha [ MCZC] ; Tucurui, Margem esq. [ LACM] ; Utinga tract, nr. Belem [ MCZC] ; “ Para ” (s. loc.) [ MCZC] . Rondônia: Porto Velho, Madeira [ MCZC] ; Rio Madeira, Madeira Mamore R. R. Camp #39 [ MCZC] ; Rio Madeira, Madeira Mamore R. R Co. Camp #41 [ LACM] . Rio de Janeiro: Ilha Grande [ ALWC] . São Paulo: Res. Florestal Caraguatatuba [ MCZC] . Colombia. Chocó 10 km SW S. Jose de Palmar, Rio Torito, Finca Los Guaduales [ MCZC] . Guajira: R. Don Diego [ MCZC] . Magdalena: Tayrona Park, S park boundary above Calabasos [ MCZC] . Valle: km 98, old road Cali to Buenaventura [ MCZC] . Costa Rica. Cartago: "Natrolista Platanillo", 1mi S Tuis [ UCDC] ; Turrialba [ MCZC] . Guanacaste: Guanacaste Cons. Area, Pitilla Research Station [ UCDC] . Heredia: La Selva Biol. Sta. [ LACM, MCZC, PSWC] ; P. N. Braulio Carrillo [ LACM] . Limón: R. Toro Amarillo, vic. Guapiles [ MCZC] ; Sarapiqui R., Oro Verde Lodge [ MCZC] ; Zent [ MCZC] . Puntarenas: Corcovado Nat. Park, nr. Rio Nino [ MCZC] ; Corcovado Nat. Park, Sirena [ MCZC] ; Palmar [ MCZC] . Ecuador. Guayas: 10 mi. N. Manglar Alto [ MCZC] ; 3 km SW Bucay [ MCZC] . Los Rios: Rio Palenque Biol. Sta. [ LACM] . Morona-Santiago: Sucúa [ LACM] . Napo: Misahualli [ MCZC] . Pastaza: 2-8 mi. N. Puyo [ MCZC] . Pichincha: ENDESA Forest Reserve [ ALWC, UCDC] . Sucumbios: Limon Cocha & vic. [ MCZC] . French Guiana. Cayenne: Paracou Experimental Forest, 45k W of Karou [ MCZC] . Guatemala. Petén: Nacum [ MCZC] . Retalhuleu: El Asintal [ UCDC] . Suchitepéquez: Finca Los Tarrales [ ALWC] . Guyana. Cuyuni-Mazaruni: Cuyani R. [ MCZC] ; Kamakusa [ MCZC] ; Kartabo [ MCZC] . Upper Takutu-Upper Essequibo: N. Side Acari Mts. [ PSWC] . Honduras. Atlántida: 14 km S La Ceiba [ MCZC] ; Lancetilla, nr. Tela [ MCZC] . “Portillo Grande” (loc. indet.) [ MCZC] . Mexico. Campeche: 10 Km E Campeche [ MCZC] . Chiapas: Ocosingo, Laguna Ocotal Grande [ MCZC] ; Ruinas Palenque [ LACM] . Oaxaca: Temescal [ LACM] . Quintana Roo: 13 km S Senor , Cenote de Tos Viriol [ LACM] ; Cobá [ LACM] ; Felipe Carillo Puerto, Cenote de Juan Coh [ LACM] . San Luis Potosí: 18 mi S. Tamazunchale [ MCZC] ; Huichihuayan [ MCZC] . Veracruz: 2 mi W. Fortin , park canon HWY 150 [ MCZC] ; Cueva de la Sala de Agua [ MCZC] ; El Palmar, 16 k W. Tezonapa [ MCZC] ; Laguna Encantada [ MCZC] ; Las Hamacas, 17k N Santiago, nr. Tuxtla [ MCZC] ; Los Tuxtlas [ ALWC] ; Presidio, Trail above Presidio [ LACM] ; Pueblo Nuevo nr. Tezonapa [ MCZC] ; Sa. Teoviscocla, nr. Cuichapa [ MCZC] ; Tlacotalpan (as "Tapalcapan") [ MCZC] . Yucatan: Chichen Itza [ MCZC] ; 1 km NE Tixcancal [ LACM] ; Actun Xpukil, 3k S Calcehtok [ LACM] ; Grutas de Balankanche 4 km E Chichen Itza [ LACM] . Nicaragua. Chinandega: (s. loc.) [ MCZC] . Indio Mais Res., San Juan and Sarapiqui Rivers [ MCZC] . “Tuli Creek” (loc. indet.) [ MCZC] . Panama. Darien: Cana [ PSWC] . Panamá: Barro Colorado I. [ LACM, MCZC, UCDC] . Peru. Huánuco: 12 km SW Tingo Maria [ LACM] ; Cueva de Castillo nr. Tingo Maria [ LACM] ; Monson Valley, Tingo Maria [ MCZC] ; Tingo Maria & vic. [ MCZC] . Loreto: Previsto [ LACM] ; Quebrada Yanayaco, NE Iquitos [ LACM] . Madre de Dios: Est. Biol. Cocha Cashu [ LACM, MCZC] ; 15 k NE Puerto Maldonado [ MCZC] . San Martín: Davidcillo, 30k NNE Tarapoto [ PSWC] . Surinam. Raleigh Vallen-Voltzberg Res. Voltzberg camp [ MCZC] ; “Surinam” (s. loc.) [ MHNG, 1w labeled “latreille Forel type ”] . Trinidad and Tobago. St. George: Caura [ MCZC] ; Guanapo Valley Quarry Guanapo Rd. [ MCZC] . Venezuela. Amazonas? ("Terr. Amazonas"): 3 km N. of San Carlos de Rio Negro [ MCZC] . Araguá: Rancho Grande [ MCZC] . Barinas: 17k SSW Ciudad Bolivia [ PSWC] . Bolívar: 1k S confl. R. Nichare & Rio Caura [ PSWC] ; Nichare Field Stn., Rio Tawadu [ PSWC] . Delta Amacuro: 140k NE Barrancas, Cano Mariusa [ LACM] .

Worker measurements: (n = 19) HL 2.40-3.19, HW 1.96-2.72, SL 2.72-3.61, WL 3.96-5.17, FL 2.63-3.66, LHT 3.05-4.38, PL 1.04-1.36, PH 1.36-1.78, CI 0.77-.86, SI 1.28-1.47.

Worker diagnosis: A large species (WL> 3.9 mm) with the apical antennomeres colored bright yellow. Head somewhat longer than broad (CI 0.77-.86); mandibles elongatetriangular and bearing 12-15 teeth. Antennal scape longer than head length. Posterolateral margins of the propodeum rounded. Posterior and lateral faces of the petiole usually meeting at a rounded or an indistinct angle. Petiolar node relatively tall (PH> 1.35mm). Abdominal tergite 3 usually with erect setae, abdominal tergite 4 always with at least a posterior row of setae and often with additional setae. Hypopygium coarsely punctate posteriorly with shining interspaces in area adjacent to sting, bearing moderate to sparse subdecumbent pubescence not completely obscuring integument (Fig. 7). Body and appendages dark brown to black, except for yellow apical antennomeres.

This species may be separated from P. obscuricornis by the longer antennal scape and from P. verenae by the lack of posterolateral margination of the petiole.

Geographic variation: The shape of the petiole changes noticeably between localities. In one specimen from Guyana the petiolar form approaches the marginate condition of P. verenae . Specimens vary considerably as well in the development of abdominal pilosity. Ants from Central America often lack erect setae on abdominal tergite 3, while specimens from elsewhere in the range commonly have anywhere between 2 and 25 erect setae on tergite 3, with Peruvian specimens being the most pilose; a few of the most pilose Peruvian specimens have erect setae on the mesosomal dorsum and petiolar node. Eye size appears to vary slightly between localities as well.

Distribution: Southern Mexico to southeastern Brazil.

Biology: Pachycondyla apicalis is a common and conspicuous insect in many Neotropical forests. Most observations and collection records are of single foragers on the ground or on low vegetation. According to the collection data associated with museum specimens, P. apicalis occurs from sea level to 1600 meters (n = 40, median = 380 meters, mean = 642 meters). 14 collections were from primary or secondary rainforest or other kind of tall, moist forest. Two records were from rainforest edges and clearings, three from coffee plantations, one from cloud forest, and one from second growth thorn forest. This species has been observed nesting in rotting wood on or near the ground (Levings & Franks 1982, Dietemann & Peeters 2000, Longino 2004), in the ground (Levings & Franks 1982), and in the root mass of large Ficus trees within one meter of the ground (Fresneau 1985). One Colombian record in MCZC is from a Guadua sp.

Colonies are small, containing fewer than 200 workers (Fresneau 1985, Goss et al 1989, Dietemann & Peeters 2000), and monogynous (Dietemann & Peeters 2000). Dietemann and Peeters (2000) investigated the interactions between queens and workers, finding that workers can lay both trophic eggs and reproductive male eggs, some switching to reproductive male eggs in the absence of physical contact with the queen. Workers are apparently incapable of mating (Dietemann & Peeters 2000) and exhibit overt dominance interactions as well as egg-policing (Oliveira & Hölldobler 1990). Age polyethism in P. apicalis is described by Fresneau and Dupuy (1988). The formation of colony odor was studied by Soroker et al (1998), who tagged lipid precursors with radioactive tracers, injected them into ants, and conducted a series of experiments demonstrating that the molecules were spread through the colony by allogrooming of nestmates and not by trophallaxis.

Fresneau (1985) describes foraging behavior in a field population in Chiapas, Mexico. He found apicalis to be generalist predators and scavengers, collecting “20%… fruit debris and vertebrate carcasses and the remaining 80%…an assortment of 12 arthropod orders half of which were dead, and other half of which were living Lepidoptera and Coleoptera larvae.”(Fresneau 1985, pg 110.) Longino (2004) observed that foragers readily take crushed tabanid flies and lepidopteran larvae. A collection by E. O. Wilson from Veracruz, Mexico, records P. apicalis preying on termites, and two apicalis workers in the MCZC collection were found at a tuna bait in Guanacaste, Costa Rica.

Foraging is done individually, without recruitment, and individual foragers over time show strong fidelity to a particular area (Fresneau 1985). Tandem-running has been observed during nest relocation (Fresneau 1985). Orientation is probably visual (Fresneau 1985). Goss et al (1989) test an optimal foraging model using P. apicalis , concluding that foraging in the observed colonies is sub-optimal. Interestingly, a group of computer scientists have used the foraging behavior of P. apicalis as a model for creating an internet search algorithm ( Monmarché et al 2000).

Pachycondyla apicalis , as in other ponerine ants, subdues its prey by injecting venom through a sting. The venom may also have a defensive purpose and is described as tasting “bitter and burning” (Schmidt 1986). Cruz and Morgan (1997) investigate venom chemistry, Schmidt (1980) looks at venom toxicity, and Schmidt et al (1984) score P. apicalis sting-induced pain in humans as a “two” on a standardized ascending scale of one to four.

Pavan et al (1997) report on the auditory emissions of P. apicalis . As in most stridulating ants, the stridulatory organ is composed of a file on the fourth abdominal tergite and a scraper on the preceding tergite (Giovannotti 1996, Pavan et al 1997). Abdominal glands in the male are described by Hölldobler and Engel-Siegel (1982).

In Panama, P. apicalis serves as a model for the ant-mimic spider Castianeira memnonia(Koch) (Reiskind 1977). Reiskind (1977) reports the identity of the model as P. obscuricornis , but the voucher specimen at LACM is clearly P. apicalis . Additionally, his description of the yellow antennal apices and the photographs in the article unambiguously identify the ant as P. apicalis .

There is one record in MCZC of Pachycondyla apicalis in the gut contents of a leptodactylid frog, Eleutherodactylus biporcatus (Peters) , in Nicaragua.