Dinoponera

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Lenhart P, Dash S, Mackay W (2013) A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae). Journal of Hymenoptera Research 31 : 119–164, doi. Versioned wiki page: 2013-03-20, version 32079, https://species-id.net/w/index.php?title=Dinoponera&oldid=32079 , contributors (alphabetical order): Pensoft Publishers.

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@article{Lenhart2013JournalofHymenopteraResearch31,
author = {Lenhart, Paul A. AND Dash, Shawn T. AND Mackay, William P.},
journal = {Journal of Hymenoptera Research},
publisher = {Pensoft Publishers},
title = {A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae)},
year = {2013},
volume = {31},
issue = {},
pages = {119--164},
doi = {10.3897/JHR.31.4335},
url = {http://www.pensoft.net/journals/jhr/article/4335/abstract},
note = {Versioned wiki page: 2013-03-20, version 32079, https://species-id.net/w/index.php?title=Dinoponera&oldid=32079 , contributors (alphabetical order): Pensoft Publishers.}

}

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TY - JOUR
T1 - A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae)
A1 - Lenhart P
A1 - Dash S
A1 - Mackay W
Y1 - 2013
JF - Journal of Hymenoptera Research
JA -
VL - 31
IS -
UR - http://dx.doi.org/10.3897/JHR.31.4335
SP - 119
EP - 164
PB - Pensoft Publishers
M1 - Versioned wiki page: 2013-03-20, version 32079, https://species-id.net/w/index.php?title=Dinoponera&oldid=32079 , contributors (alphabetical order): Pensoft Publishers.

M3 - doi:10.3897/JHR.31.4335

Wikipedia/ Citizendium:

<ref name="Lenhart2013Journal of Hymenoptera Research31">{{Citation
| author = Lenhart P, Dash S, Mackay W
| title = A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae)
| journal = Journal of Hymenoptera Research
| year = 2013
| volume = 31
| issue =
| pages = 119--164
| pmid =
| publisher = Pensoft Publishers
| doi = 10.3897/JHR.31.4335
| url = http://www.pensoft.net/journals/jhr/article/4335/abstract
| pmc =
| accessdate = 2022-10-02

}} Versioned wiki page: 2013-03-20, version 32079, https://species-id.net/w/index.php?title=Dinoponera&oldid=32079 , contributors (alphabetical order): Pensoft Publishers.</ref>

See also the citation download page at the journal.


Taxonavigation

Ordo: Hymenoptera
Familia: Formicidae

Name

Dinoponera Roger, 1861Wikispecies linkPensoft Profile

==

Family Formicidae, subfamily Ponerinae, tribe Ponerini. Described by Perty (1833)[1] as Ponera gigantea. Defined as a genus by Roger (1861)[2] (Type species: Dinoponera gigantea)

Diagnosis

Size (TBL > 2.5cm) can easily distinguish Dinoponera from other worker ants. Two laterally projecting clypeal teeth (Fig. 1A) and rows of spines on the pygidium and hypopygidium will further distinguish this genus. The gamergates of Dinoponera are not distinct from workers in their external morphology (Haskins and Zahl 1971[3], Araujo et al. 1990[4], Paiva and Brandão 1995[5], Monnin and Peeters 1998[6]). True gynes have not been found in this genus.

Description

Description of the worker. Abundant setae; black integument, ranges from smooth and shiny with no microsculpturing, to finely micropunctate or scaled depending on species (Fig. 12). Head: Mandibles long and curved posteriorly in side view; seven large teeth; erect setae on dorsum. Ventral surface of head with sparse decumbent and subdecumbent setae; may have fine striations depending on species; Papal formula 4, 4; large bilobed labrum. Clypeus with two laterally projecting teeth on anterior edge, clypeus bulging medially, extending posteriorly between frontal lobes, anterior edge with row of long setae; sparse appressed setae from distal edges to medial area of clypeus. Area posterior to clypeus with varying amounts of striation. Tentorial pits apparent. Frontal lobes raised and conspicuous, with striations at posterior constriction. Antennae: geniculate, 12 segments, all with flagellate setae; scape long, extending past posterior border of head; funiculus covered in minute appressed pubescence. Gena depressed medially of eye; dense appressed setae on the antero-lateral sides of the head; covered in conflected punctulate sculpturing. Eyes large, elliptical with slight depression (ocular ring) around circumference. Frons with large pads of long flagellate pubescence (lost in older or poorly curated specimens). Median furrow running from posterior termination of clypeus, between frontal lobes to center of frons, terminates in shallow pit in most specimens. Entire head covered in long flagellate subdecumbent setae (Fig. 1A). Mesosoma: in lateral view weakly convex; covered in long subdecumbent to erect flagellate pilosity and dense pubescence; pronotal disc with slight bulges; promesonotal suture distinct, suture between mesopleuron and propodeum distinct; mesonotum fused with propodeum and episternum, separated by slight furrows; basilar sclerite large, ovaloid; propodeum with broadly rounded dorsal outline, dorsal surface gradually curves into posterior face (Fig. 2); propodeal spiracle forms nearly vertical slit; sulcus running from center of propodeum along lower edge of propodeal spiracle to posterior edge of propodeum at dorsal edge of bulla, patches of short white pubescence at curved posterior border of pronotum and basilar sclerite. Legs long, covered in long setae with short, stiff pubescence. One well-developed, antennae cleaning, comb-like spur on foreleg; one spine-like appendage and one less developed denticular comb on mesothoracic tibia; one spine and one comb-like spur on hind tibia. Posterior side of fore leg basitarsus with dense pads of golden setae; tarsal claws bidentate. Petiole: node large and tabular in lateral view, narrow attachments at base to propodeum and gaster; in dorsal view largest width less than propodeum and gaster, varies from ovate rectangular to ovate triangular in outline; covered in long subdecumbent to erect flagellate pilosity; pubescence on anterior face and ridges of subpetiolar process; subpetiolar process reduced, slightly variable between species. Gaster: typical of ponerines; covered with flagellate setae with short pubescence; small protuberance at articulation of gastric sternite III and the petiole; stridulatory file of varying size on acrotergite of gastral tergum II; posterior edges of the pygidium and hypopygidium with characteristic rows of minute spines. Description of the male. Integument: smooth and nitid; reddish to dark brown/black. Head: Mandibles greatly reduced, rounded, spoon shaped, lacking teeth; palps elongated, maxillary palps 4 segmented, labial palps 3 segmented; labrum reduced, rounded to truncate, emarginated distal margin in Dinoponera snellingi and Dinoponera longipes covered with setae. Clypeus large, triangular, bulging medially; anterior tentorial pits large; frontal lobes absent; antennal sockets almost touching, located at posterior apex of clypeus. Antennae: geniculate, 13–segmented, pilosity varies from fine pubescence to long setae in different species; scape shorter than second funicular segment, but shorter than 1st, 1st funicular segment reduced. Compound eyes large, along lateral side of head, deeply emarginated medially. Three ocelli at posterior margin of head, bulging beyond margin of head in all species except Dinoponera australis. Entire head immaculate, covered in fine pubescence and long erect setae (Fig. 3). Mesosoma: pronotum triangular, exposed narrowly dorsally anterior to scutum; scutum large, bulging antero-dorsally, with 3 longitudinal carina; small tegula over insertion of forewing; scutellum domed, side with vertical carina, dorsal surface smooth; basilar sclerite under hind wing reduced; fused mesopleuron, separated by furrow into anepisternum and katepisternum; metanotum exposed between scutellum and propodeum, reduced; dorsal face of propodeum shorter than posterior face, rounded into posterior face; coxa large, conical (Fig. 3). Wings: covered in minute pubescence, venation as shown in Figure 5. Legs: one well-developed, antennae cleaning, pectinate spur on foreleg; one spine-like and one less developed denticular comb on mesothoracic tibia; one spine and one comb-like spur on hind tibia. Posterior side of fore basitarsus with dense pads of golden setae; tarsal claws bidentate. Petiole: narrow attachments at base to propodeum and gaster; petiolar node humped dorsally, subpetiolar process anteriorly triangular. Gaster: large, cylindrical, covered in fine silvery pubescence; pygidium terminating in spine posteriorly, with short cerci; hypopygidium with long fine erect setae, tabular subgenital plate with posterior end truncated, often emarginated. Genitalia (Figs 6–11): basal ring with dorso-anterior loop structures; parameres long, rounded, with emarginated ventro-basal edge (Fig. 9); volsella articulated with basiparamere along ventral edge, lateral finger-like cuspis volsellaris, medial digitus volsellaris with distal wide toothed cusp, basal medial lobe with tooth-like structures varying with species (Fig. 10); penis valve of aedeagus roughly triangular and rounded, aedeagal apodeme curved horn-like antero-lateral arm structure arising from mid-valve ridge, terminating at interior surface of basiparamere (Fig. 11). Description of the larvae. A basic description of the larva of Dinoponera quadriceps (cited as Dinoponera grandis mutica) is present in Mann (1916)[7]. A detailed description of the egg and all larval stages of Dinoponera gigantea are present in Wheeler and Wheeler (1985)[8]. The following generic description of Dinoponera larvae is from their work:
"Profile pogonomyrmecoid (i.e., diameter greatest near the middle of abdomen, decreasing gradually toward anterior end and more rapidly toward posterior end, which is rounded; thorax more slender than abdomen and forming a neck, which is curved ventrally). Body with numerous (114–160) mammiform tubercles, each with 2–25 short simple hairs; body hairs lacking elsewhere. Cranial hairs lacking. Mandible dinoponeroid (i.e. narrowly subtriangular in anterior view; anterior portion curved posteriorly; with or without medial teeth.)"

Discussion

Dinoponera’s status as a genus is validated as several characters differentiate it from other genera. Size (TBL>2.3cm) is the most obvious character distinguishing Dinoponera. The only other ants with a worker caste approaching this size are Paraponera clavata (Fabricius)and the larger Pachycondyla such as Pachycondyla crassinoda (Latreille 1802[9]), Pachycondyla impressa Roger 1861[2] and Pachycondyla villosa (Fabricius 1804[10]). Paraponera clavata is easily separated by its anvil shaped petiole with a spine on the ventral surface, highly sculptured body and deep antennal scrobes. Pachycondyla is regarded as the sister taxa to Dinoponera (Kempf 1971[11], Schmidt 2010[12]). Dinoponera, in addition to their size, are distinguishable from Pachycondyla by the presence of two laterally projecting clypeal teeth (Fig. 1A) and rows of spines on the pygidium and hypopygidium. Several (n=6) specimens have been observed to have a single ocelli in the pit at the termination of the median furrow. These anomalous specimens were previously thought to be queens (Borgmeier 1937[13]) but as it has been shown that Dinoponera lacks queens, the presence of the ocelli is hypothesized to be the result of a Mermis Dujardin 1842 nematode parasite (Kempf 1971[11]).

Dinoponera biology

Dinoponera is one of the roughly 10 ponerine genera in which some species have secondarily lost the typical morphologically specialized queen caste for a reproductive worker known as a gamergate (Haskins and Zahl 1971[3], Araujo et al. 1990[4], Paiva and Brandão 1995[5], Monnin and Peeters 1998[6], Peixoto et al. 2008[14]). Conflict over dominance is intense in colonies with younger workers usually joining a linear hierarchy of one to five workers depending on colony size. The gamergate, or alpha female has the highest ranking (Monnin and Ratnieks 1999[15], Monnin et al. 2003[16]). The alpha female mates with non-nestmate males at night at the entrance of the nest (Monnin and Peeters 1998[6], Monnin and Peeters 1999[17]). After copulation the female bites through the male’s gaster to release herself and pulls out the genital capsule which acts as a temporary sperm plug (Monnin and Peeters 1998[6]). After mating the female is unreceptive to other males and remains monandrous (Monnin and Peeters 1998[6]). The gamergate maintains dominance with ritualized behaviors such as antennal boxing and biting, ‘blocking’, as well as gaster rubbing and curling (Monnin and Peeters 1999[17]). Lipid stores within Dinoponera australis females were found to be strongly related to foraging activity and reproductive status within the colony, ranging from 1–39% of an individual’s dry mass (Smith et al. 2011[18]). It is uncertain, however, whether nutritional differences between females is a cause or consequence of rank. Gamergate females possess a higher concentration of a cuticular hydrocarbon (9-hentriacontene, 9-C31:1) that indicates rank and is passed onto gamergate-laid egg cuticles (Monnin and Peeters 1997[19], Monnin et al. 1998[6], Peeters et al. 1999[20]). Additionally, alpha females may ‘sting smear’ a competing female with secretions from the Dufour’s gland, triggering the lower ranking workers to immobilize the marked female (Monnin and Ratnieks 2001[21]). Subordinate females (beta, gamma, or delta) may produce unfertilized eggs but these are usually consumed by the alpha female in a form of “queen policing” (Monnin and Peeters 1997[19]). Egg recognition in Dinoponera quadriceps was found to be due to differences in cuticular hydrocarbons, and only workers engaged in brood care could distinguish non-nestmate eggs (Tannure-Nascimento et al. 2009[22]). Cuticular hydrocarbons are also used to distinguish adult nestmates from non-nestmates, however, this is only effective with non-nestmate foragers (Nascimento et al. 2012[23]). Nascimento et al. (2012)[23] found that brood-caring workers from different colonies had very similar hydrocarbon profiles and were more often accepted into alien colonies.
Males are born throughout most of the year in tropical species (Araujo and Jaisson 1994[24], Monnin and Peeters 1998[6]), however Dinoponera australis which lives in the more temperate south was found to only produce males in May-July (Paiva and Brandão 1995[5]). When the alpha declines reproductively or dies, she is replaced by a high-ranking worker (Monnin and Peeters 1999[17]).
New colonies are founded by fission, a process in which a beta female is fertilized in the natal nest (Monnin and Peeters 1998[6]). This new alpha female then leaves the nest with a cohort of workers to found an incipient colony, sometimes employing tandem running (Overal 1980[25]).
Colonies vary in size depending upon species. Dinoponera australis colonies have an average of 14 workers (range 3–37) (Paiva and Brandão 1995[5], Monnin et al. 2003[16]), Dinoponera gigantea average 41 workers (range~30–96) (Overal 1980[25], Fourcassié and Oliviera 2002[26], Monnin et al. 2003[16]) and Dinoponera quadriceps has the largest colonies with an average of 80 workers (range 26–238) (Monnin and Peeters 1999[17], Monnin and Ratnieks 2001[21]). Morgan (1993)[27] excavated two Dinoponera longipes nests, a possible incipient colony with 7 workers and another mature colony of 120 workers.
The nest consists of large chambers and tunnels in the soil possibly with an earthen mound and can be 0.10–1.2m deep (Araujo et al. 1990[4], Morgan 1993[27], Fourcassié and Oliviera 2002[26], Vasconcellos et al. 2004[28]). Nests are deeper in Dinoponera australis and Dinoponera quadriceps than in Dinoponera gigantea, Monnin et al. (2003)[16] suggests that deeper nests are a possible adaptation to seasons and aridity. Dinoponera gigantea nests may have up to eight entrances and can be weakly polydomous (Fourcassié and Oliviera 2002[26]), whereas 1–30 openings with an average of 11 were recorded for Dinoponera longipes (Morgan 1993[27]). Nesting density and spatial distribution varies depending on habitat (Fowler 1985[29], Vasconcellos et al. 2004[28]). Density ranges from 15–40 nests per ha-1 (Vasconcellos et al. 2004[28]) to 80 nests per ha-1 (Paiva and Brandão 1995[5]). Morgan (1993)[27] measured a spacing between nests for Dinoponera longipes with a median of 35m (n=22, range 14–69.5m). Dinoponera australis and Dinoponera gigantea usually nest at the base of trees (Paiva and Brandão 1995[5], Fourcassié and Oliviera 2002[26]). Observations of Dinoponera quadriceps nests show that in more arid Caatinga and Cerrado habitats, nests are predominantly constructed under trees, whereas in Atlantic forest 60% of nests were 3m away from any tree (Vasconcellos et al. 2004[28]).
Workers lower in the hierarchy forage individually for food items on the substrate and do not recruit other nestmates to assist with food transport (Fowler 1985[29], Fourcassié et al. 1999[30], Fourcassié and Oliviera 2002[26], Araújo and Rodrigues 2006[31]). Although foraging workers do not recruit nestmates, Nascimento et al. (2012)[23] found a positive feedback between incoming food and stimulation of new foragers as well as task partitioning once food was brought into the nest. Lower ranking females processed protein resources while higher ranking females handled small food pieces and distributed them to the larvae. Fourcassié and Oliviera (2002)[26] found Dinoponera gigantea foraging to be concentrated in the early morning and afternoon but did not sample at night. Morgan (1993)[27] observed the highest activity at night in Dinoponera longipes. Dinoponera quadriceps has a marked seasonal pattern in activity. It is most active in May-August , the late rainy season to early dry season in the semiarid Caatinga (Medeiros et al. 2012[32]). Activity was strongly negatively correlated to temperature and positively correlated to prey abundance (Medeiros et al. 2012[32]). The diets of both Dinoponera gigantea and Dinoponera quadriceps have been shown to be predominantly scavenged invertebrates, but include live prey, seeds and fruits (Zahl 1959[33], Fourcassié and Oliviera 2002[26], Araújo and Rodrigues 2006[31]). Araújo and Rodrigues (2006)[31] state that the taxonomic diversity of prey is comparable to other tropical ponerines, but has an optimal prey size of 2–3 cm in Dinoponera. Diet seems to be very similar across the genus, regardless of habitat (Araújo and Rodrigues 2006[31]).
Despite their large size and strong venom, Dinoponera are likely preyed on by many vertebrate and invertebrate species across South America. Like many other ant species, Dinoponera can be infected by the entomopathogenic fungi Codyceps sp. (Evans 1982[34]). Buys et al. (2010)[35] discovered a Kapala sp. eucharitid wasp emerging from the puparia of Dinoponera lucida.
Anatomy has been described several times. Marques-Silva et al. (2006)[36] studied of the sensilla and glands of the antennae. Anatomy of the venom apparatus and mandibular glands of Dinoponera gigantea is presented inHermann et al. (1984)[37]. Further studies of the mandibular glands and its contents were presented by Oldham and Morgan (1993)[38] and Oldham et al. (1994)[39]. Oldham et al. (1994)[39] found that the mandibular gland secretions of workers differed greatly from those of gamergates, which were 98% dimethylalkylpyrazine and lacked the four other pyrazines and 50 times more volatiles than those found in worker secretions. The post-pharyngeal gland morphology was examined by Schoeters and Billen (1997)[40]. The cuticular hydrocarbons used in nestmate recognition may be produced by epidermal glands which Serrão et al. (2009)[41] found in the epidermis of abdominal sternites in Dinoponera lucida.
For subduing large live prey and defense (Morgan 1993[27]), workers possess a sting that has been known to cause severe pain lasting up to 48 hours, lymphaedenopathy, edema, tachycardia and fresh blood to appear in human victim feces are common symptoms (Haddad et al. 2005[42]). In gamergates the venom sac is empty (Monnin et al. 2002[43]). Workers may have 60–75 unique proteinaceous components in the venom (Morgan et al. 2003[44], Johnson et al. 2010[45]). The convoluted gland within the venom system of Dinoponera australis has been found to possess close similarities to those of vespine wasps (Schoeters and Billen 1995[46]). The contents of Dinoponera australis venom have been found to be similar to those of Pachycondyla spp. (Cruz López 1994[47], Johnson et al. 2010[45]). Billen et al. (1995)[48] studied the morphology and ultrastructure of the pygidial gland of Dinoponera australis. Due to the high diversity of compounds and systemic effects found by Haddad et al. (2005)[42], venom of Dinoponera could be of use to the pharmaceutical industry. For instance, Sousa et al. (2012)[49] demonstrated in mice that venom from Dinoponera quadriceps had antinociceptive properties. The authors note that the local population of northeast Brazil uses dry crushed Dinoponera quadriceps ants to treat earaches, and the stings of live ants are administered for back pain and rheumatism.
Several studies of the cytogenetics of Dinoponera species have been conducted. Dinoponera lucida may have the highest number of chromosomes within the Hymenoptera however the karyotype is variable between populations (2n=106–120) (Mariano et al. 2004[50], Mariano et al. 2008[51], Barros et al. 2009[52]). Mariano et al. (2008)[51] interpreted the karyotype differences between populations as being due to a division of the species into allopatric populations during the Quaternary. Variability in the karyotype within a described species has been found in the Pachycondyla as well, and may represent cryptic species (Mariano et al. 2012[53]). Descriptions of the banding patterns on Dinoponera chromosomes are provided by Barros et al. (2009)[52] and de Aguiar et al. (2011)[54].
Dinoponera belongs to the tribe Ponerini in the subfamily Ponerinae. The evolutionary position of the genus within Ponerinae was resolved by Schmidt (2010)[12]. Based on the phylogenetic analysis of Schmidt (2010)[12] and karyotype analysis by Mariano et al. (2012)[53], Dinoponera’s closest living relatives are in the Pachycondyla species group consisting of Pachycondyla crassinoda, Pachycondyla harpax (Fabricius 1804[10]), Pachycondyla impressa, Pachycondyla metanotalis Luederwaldt 1918[55], and Pachycondyla striata Smith 1858[56]. Prior to the generation of well supported phylogenies other associations had been proposed. Carpenter (1930)[57] suggested that the fossil Archiponera wheeleri Carpenter from the Miocene Florissant shale of Colorado may be an ancestor of Dinoponera and Streblognathus aethiopicus Smith 1858[56]. Molecular data has shown that Carpenter’s (1930) hypothesis is false (Schmidt 2010[12]). Streblognathus is not closely related to Dinoponera, and its morphological similarity is purely convergence. The placement of Archiponera wheeleri is still questionable.

Key to the workers of Dinoponera

Key to the known males of Dinoponera (couplets 1 and 2 are included to easily separate males of other genera which are likely confused with Dinoponera)


Clave para la identificación de las obreras de Dinoponera
Clave para la identificación de los machos conocidos de Dinoponera
Chave para identificação de operários de Dinoponera
Chave para identificação de machos de Dinoponera

Taxon Treatment

  • Lenhart, P; Dash, S; Mackay, W; 2013: A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) Journal of Hymenoptera Research, 31: 119-164. doi

Other References

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  2. 2.0 2.1 Roger J (1861) Die Ponera-artigen Ameisen (Schluss). Berliner Entomologische Zeitschrift 5: 1-54.
  3. 3.0 3.1 Haskins C, Zahl P (1971) The reproductive pattern of Dinoponera grandis Roger (Hymenoptera, Ponerinae) with notes on the ethology of the species. Psyche 78: 1-11.
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