Strumigenys membranifera

Strumigenys membranifera
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Myrmicinae
Tribe:	Attini
Genus:	Strumigenys
Species group:	membranifera
Species:	S. membranifera
Binomial name
	Strumigenys membranifera; Emery, 1869 Specimen Label
Synonyms
	Strumigenys foochowensis Wheeler, W.M., 1928; Strumigenys membranifera marioni Wheeler, W.M., 1933; Strumigenys membranifera santschii Forel, 1904; Strumigenys membranifera simillima Emery, 1890; Strumigenys membranifera williamsi Wheeler, W.M., 1933; Strumigenys silvestriana Wheeler, W.M., 1928; Strumigenys vitiensis Mann, 1921;

Common Name

Tokara-uroko-ari
Language:	Japanese

Membraniferous dacetine ant
Language:	English

This species, globally introduced to all continents except Antarctica, is apparently easily introduced to new areas by human activities. It has a remarkably wide distribution, occurring in both mainland and island regions throughout the tropics and subtropics, and occasionally occurring in the temperate zone. Brown & Wilson (1959) suggested an African origin, but this was questioned by Bolton (1983) and its phylogenetic position places it as Indomalayan in origin (Booher et al. 2021). Colonies typically occur in highly disturbed or cultivated areas, beneath debris or dead wood on the ground (Longino, Ants of Costa Rica) and are often found in grass clippings, mulch piles, and litter debris in more open and disturbed areas (Booher, 2021). It forms relatively large colonies of about 250 individuals (up to 350 individuals; Ito et al. 2010).

Like several other Strumigenys tramp species, this species reproduces asexually through thelytokous parthenogenesis (Ito et al. 2010) and produces alate queens over a much longer period when compared with natives. In southeastern USA states, alate queens have been collected in all months between June and November, whereas in California alates have been collected in February and late July. A confirmed male identification for this species suggests it may also be capable of reproducing sexually. This male specimen was identified genetically (Booher et al. 2021; Booher, 2021). Wilson (1954) reported it feeding on a wide range of small soft-bodied arthropods.

At a Glance

• Highly invasive • Parthenogenetic • Polygynous

Identification

The only member of the Strumigenys membranifera-group.

Longino (Ants of Costa Rica) - Mandible short and curved downward; dorsal mesosoma smooth or nearly so; pronotum sharply marginate anteriorly, marginate dorsolaterally throughout its entire length; with the head in full-face view and the mandibles fully closed the dorsal surface of each mandible basally with a very distinct sharp transverse edge or rim that extends across the width of the blade, parallel to and in front of the anterior clypeal margin; pronotal humeral hair absent; dorsal surfaces of middle and hind tibiae with decumbent to appressed short spatulate hairs.

Tang & Guenard (2023), Fig. 30. New species records of Strumigenys in full-face, profile and dorsal views. A–C. Worker of S. membranifera from Vietnam (NN-S56-sp01). D–F. Worker of S. minutula from Hainan, mainland China (HNA-01641). G–I. Worker of S. mitis from Hainan, mainland China (HNA-00275).

Keys including this Species

Distribution

Native: Afrotropical realm (Ghana, Sierra Leone, South Africa) (Tang & Guenard, 2023).

Introduced: Oriental realm (mainland China: Yunnan; India; Singapore; Taiwan; Vietnam: Đắk Nông, Đồng Nai), Sino-Japanese realm (Bhutan, mainland China: Fujian, Guangdong, Hong Kong, Macau, Sichuan; Japan: Honshu, Kyushu, Ryukyu Islands, Shikoku; Nepal). For a full global account refer to antmaps.org (Janicki et al. 2016; Guénard et al. 2017) (Tang & Guenard, 2023).

Bolton (2000) - P. membranifera is mainly a pantropical tramp species, but it also occurs outside the tropics in favourable locations. Material examined subsequent to that listed in Bolton (1983) is from Sierra Leone, Egypt and U.S.A.. Type-data of the various synonyms record the species from Italy, Tunisia, St Thomas I. in the West Indies, Fiji Is, China, U.S.A. and Hawaii. Wilson & Taylor (1967) note its presence in Samoa. Kempf (1972) has Neotropical records from Puerto Rico, Costa Rica and Venezuela. In the U.S.A. membranifera has been recorded from Alabama, California, Florida (moderately common throughout the state (Deyrup, Davis & Cover, 2000)), Georgia, Louisiana, Mississippi and Texas (M.R. Smith, 1931; D.R. Smith, 1979; Deyrup, Johnson, et al., 1989; Cokendolpher, 1990). Deyrup (1997) records it from Bahamas.

Japanese records are generally from the south, ranging northwards to southern Honshu. The most northern record is from Tokyo (Kubota & Terayama, 1988).

Within the United States it is more commonly collected in eastern states. It has been collected infrequently in the western states of California (Los Angeles and Yolo counties), Arizona (Maricopa County) and Texas (Brazos and Robertson counties) (Booher, 2021).

Taheri & Reyes-Lopez (2023) report that this species was first discovered in Morocco beginning in 2011 (Taheri & Reyes-López, 2011; Taheri & Reyes-López, 2018). In all cases, the records were from urban areas. The importance of their discovery is that this species has been recorded for the first time in a natural habitat which is the largest cork oak forest in the world. It should also be noted that the forest area where the species has been found is heavily frequented by families for entertainment. Moreover, a continuous urbanization expansion is also remarkable such as: the construction of sports complexes, roads and gardens, which can probably explain the introduction of this tramp species.

Tang & Guenard (2023) - This record in southern Vietnam is the second in the Indochinese Peninsula, besides its recent collection in Singapore (Wang et al. 2022). Given the widespread distribution of this Afrotropical tramp species across almost all biogeographic realms, and its presence from India to Japan in the north, and from Indonesia (Lesser Sunda Islands) to the Solomon Islands in the south, this new record is not surprising. It also indicates the possibility of its presence in nearby countries such as Laos, Cambodia, Thailand and Myanmar, and Guangxi and Hainan provinces of mainland China.

Latitudinal Distribution Pattern

Latitudinal Range: 16.884° to -33.81667°.


North Temperate	North Subtropical	Tropical	South Subtropical	South Temperate

Source: AntMaps

Distribution based on Regional Taxon Lists

Afrotropical Region: Cameroun, Comoros, Saudi Arabia, Sierra Leone, South Africa, United Arab Emirates.
Australasian Region: Australia.
Indo-Australian Region: Fiji, Guam, Hawaii, Indonesia, Marshall Islands, Micronesia (Federated States of), New Guinea, Northern Mariana Islands, Samoa, Singapore, Solomon Islands, Tonga, Wallis and Futuna Islands.
Malagasy Region: Mayotte, Seychelles.
Nearctic Region: United States.
Neotropical Region: Barbados, British Virgin Islands, Colombia, Cuba, Dominican Republic, Ecuador, Galapagos Islands, Greater Antilles, Mexico, Puerto Rico, Turks and Caicos Islands.
Oriental Region: Bhutan, India, Nepal, Sri Lanka, Taiwan, Vietnam.
Palaearctic Region: Algeria, Balearic Islands, China, Egypt, Greece, Iberian Peninsula, Israel, Italy (type locality), Japan, Malta, Oman, Qatar, Republic of Korea, Spain, Tunisia, Türkiye.

Distribution based on AntMaps

Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.

Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.

Habitat

From moist forests to disturbed open areas, such as lawns and pastures.

Biology

Wilson (1953) studied two colonies in laboratory nests over an eight-month period. He reported this species accepted a wide range of small invertebrates. The following is a modified synopsis of additional observations:

These ants are the most active of the dacetines studied and show unexpected pugnacity under certain conditions. Japyx and termites were frequently attacked in the artificial nests with a viciousness approaching that of some of the more aggressive larger myrmicines. The little dacetines actually rushed at these insects when they contacted them, seizing hairs or appendages and doubling up in an effort to sting.

Although they lacked long mandibles to be used as a weapon of shock, they were able to panic fairly large individuals and dispose of them as quickly as dacetines with long mandibles. Since insects combated in this fashion were rarely eaten, it is assumed that such aggressive behavior is used in the defense of the nest.

Stalking of Collembola is in an entirely different mood and is usually executed cautiously and with deliberation. Workers rarely recognize an insect until they come in contact with it, but are very sensitive to nearby movement and probably depend a great deal on this in spotting prey.

As soon as the worker becomes aware of the presence of a collembolan, it "freezes" in a lowered, crouching position and holds this stance briefly. If the collembolan is to its back or side, the worker now turns very slowly to face it. Once it is aligned with the collembolan, it begins a forward movement so extraordinarily slow that it can be detected only by persistent and careful observation. Several minutes may pass before the ant finally maneuvers over less than a millimeter's distance to come into a striking position, and it may be remain in this position for as much as a minute or more. During this time the collembolan may move away for one reason or another, in which case the ant rarely attempts to follow but begins to forage anew. Workers open their mandibles only to about a 60° angle. Tactile labral hairs are present and eventually come to touch the prey. The mandibular strike is sudden but lacking a long mandible it is usually directed at an appendage and does not have the same stunning effect on the collembolan. Captured insects often struggle vigorously to escape, but the ants are very tenacious and retain a fast grip until they are able to sting their prey into immobility.

Unusual behavior in foraging workers was sometimes observed. Occasional bold workers were seen to approach their prey rapidly and attempt to seize a hair or appendage, and these were frequently successful. Sometimes workers would "freeze" when Collembola were moving rapidly around them and wait without moving, apparently in anticipation of ambushing the animals as these blundered into their mandibles.

Workers were never observed to participate in any sort of trophallaxis, either between workers or between workers and larvae.

Development times (constant temperature of 27 C) in days were found to be: egg-13, larvae-19 and pupa-19

This species is relatively adaptable in its nesting requirements being found in many different situations that vary from forest litter to under bits of slate in a cultivated field. Its moisture requirements are not as strict as for many congeners, and membranifera venture into relatively dry, open soil.

In Florida this species often occurs in open areas such as pastures and lawns. Although a form of this species was described from specimens apparently collected in the southeastern U.S. in the 1800's (Brown 1948), the first published Florida record seems to be that of Smith 1951; earlier specimens: 1943. (Deyrup, Davis & Cover, 2000.)

Atchison & Lucky (2022) found that this species, as expected, does not remove seeds.

Behavior

Masuko (1984) observed and reported on what he called "body smearing behavior" in S. membranifera and other Strumigenys species (see Strumigenys behavior).

Regional Information

Hong Kong

Tang et al. (2019): This is a species restricted to habitats with frequent disturbance, particularly within lowland areas (Hong Kong Airport, Disneyland, and Mai Po Nature Reserve) covered with grasslands or remnants of forests within urbanized matrices. One alate female was collected in a Malaise trap between the June 13–27, suggesting potential swarming during this period in Hong Kong. Elevation ranged from 1 to 29 m, with the exception of an alate gyne collected at an elevation of 142 m.

The record of this tramp species in Hong Kong is not surprising considering its widespread range in the region and previous records in the nearby provinces of Guangdong and Fujian as well as from Macau for 90 years (Wheeler 1928), while extending further west in China to Yunnan. Considering the widespread range of this species in tropical and subtropical regions, it appears likely that this species is already present in two other provinces of China: Guangxi and Hainan. Future sampling efforts, in particular within urban habitats, could support this hypothesis.

Qatar

Sharaf et al. (2015) - A single specimen was found in a semidry soil at the edge of an irrigated area which was planted with Bermuda grass (Cynodon sp.) and some trees of Pritchardia lanaiensis Becc. & Rock (Arecaceae). The single specimen was collected by sifting soil. Despite hours of observing, sifting, and digging soil in this area, no additional specimens were detected.

The only available study on the invasive ant species known from the Arabian Peninsula was by Collingwood et al. (1997) treating those for the UAE. These authors listed 15 introduced ant species. The discovery of S. membranifera in nearly dry soil located adjacent to an irrigated field of cultivated Bermuda grass is similar with other by reports by Wilson & Hunt (1967), Derup (1997), and Ordóñez-Urbano et al. (2008), who indicated that the preferred habitat of the species was semidry soil of exposed lawn and pastures. However, in Saudi Arabia, Sharaf et al. (2014) reported this ant foraging in date palm orchards with relatively dry soils with decaying dates.

Flight Period

	X	X	X	X	X	X	X	X	X	X
Jan	Feb	Mar	Apr	May	Jun	Jul	Aug	Sep	Oct	Nov	Dec

Source: Booher et al. 2021; Booher, 2021.

Check details at Worldwide Ant Nuptial Flights Data, AntNupTracker and AntKeeping.

Explore: Show all Flight Month data or Search these data. See also a list of all data tables or learn how data is managed.

Association with Other Organisms

Explore: Show all Associate data or Search these data. See also a list of all data tables or learn how data is managed.

Fungi

This species is a host for the fungus Myrmicinosporidium durum (a pathogen) in United States (Pereira, 2004; Espadaler & Santamaria, 2012).

Life History Traits

Mean colony size: 250
Maximum colony size: 350 (Ito et al., 2010)

Castes

Worker

Images from AntWeb


Worker. Specimen code casent0104518. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by ABS, Lake Placid, FL, USA.


Worker. Specimen code casent0133285. Photographer Erin Prado, uploaded by California Academy of Sciences.	Owned by CAS, San Francisco, CA, USA.


Worker. Specimen code casent0173252. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by CDRS, Galapagos, Ecuador.


Worker. Specimen code casent0005675. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by UCDC, Davis, CA, USA.


Syntype of Strumigenys membranifera. Worker. Specimen code casent0102081. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by MSNG, Genoa, Italy.

Queen

Images from AntWeb


Queen (alate/dealate). Specimen code casent0104474. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by ABS, Lake Placid, FL, USA.


Queen (alate/dealate). Specimen code casent0173251. Photographer April Nobile, uploaded by California Academy of Sciences.	Owned by CDRS, Galapagos, Ecuador.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

membranifera. Strumigenys (Trichoscapa) membranifera Emery, 1869b: 24, fig. 11 (w.) ITALY. Emery, 1916b: 205 (q.); Wheeler, G.C. & Wheeler, J. 1991b: 93 (l.). Combination in S. (Cephaloxys): Emery, 1916b: 205; in Trichoscapa: Brown, 1948e: 113; in Pyramica: Bolton, 1999: 1673; in Strumigenys: Baroni Urbani & De Andrade, 2007: 123. Senior synonym of foochowensis, marioni, santschii, silvestriana, simillima, vitiensis, williamsi: Brown, 1948e: 114. See also: Brown, 1949d: 6; Wilson, 1954: 483; Bolton, 1983: 319; Bolton, 2000: 322.
simillima. Strumigenys membranifera r. simillima Emery, 1890b: 69, pl. 8, fig. 5 (w.) ANTILLES. Combination in S. (Cephaloxys): Emery, 1924d: 325. Junior synonym of membranifera: Brown, 1948e: 114.
santschii. Strumigenys membranifera var. santschii Forel, 1904c: 6 (w.) TUNISIA. Junior synonym of membranifera: Brown, 1948e: 114.
vitiensis. Strumigenys (Cephaloxys) vitiensis Mann, 1921: 461, fig. 22 (w.) FIJI IS. Junior synonym of membranifera: Brown, 1948e: 114.
foochowensis. Strumigenys (Cephaloxys) foochowensis Wheeler, W.M. 1928c: 28 (q.) CHINA. Junior synonym of membranifera: Brown, 1948e: 114.
silvestriana. Strumigenys (Cephaloxys) silvestriana Wheeler, W.M. 1928c: 27 (w.) CHINA. Junior synonym of membranifera: Brown, 1948e: 114.
marioni. Strumigenys (Cephaloxys) membranifera var. marioni Wheeler, W.M. 1933h: 276 (w.) U.S.A. Junior synonym of membranifera: Brown, 1948e: 114.
williamsi. Strumigenys (Cephaloxys) membranifera var. williamsi Wheeler, W.M. 1933h: 276 (w.q.) HAWAII. Junior synonym of membranifera: Brown, 1948e: 114.

Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.

Bolton (2000) - The single species of this Strumigenys membranifera-group previously constituted the genus Trichoscapa in the sense of Brown (1948b) and Bolton (1983), now abandoned (Bolton, 1999).

Description

Worker

Bolton (2000) - TL 1.9 - 2.1, HL 0.46 - 0.50, HW 0.40 - 0.44, CI 84 - 90, ML 0.08 - 0.10, MI 16 - 20, SL 0.22 - 0.24, SI 51 - 57, PW 0.23 - 0.26, AL 0.50 - 0.53 (10 measured; from Bolton, 1983). As there is only one species in the Strumigenys membranifera-group its diagnosis will also serve to characterise the species.

Mandibles in full-face view and at full closure short triangular, serially dentate and the masticatory margins engaging throughout their length. Dorsal surface of mandible sharply depressed immediately behind the level of the basal tooth, the rim of the depression appearing as a sharp transverse edge that traverses the width of the mandible immediately behind the level of the basal tooth, and paralleling the anterior clypeal margin. The transverse edge appears to be followed proximally by a deep transverse groove or impression immediately in front of the anterior clypeal margin. In profile dorsal surface of mandible arched-downcurved from clypeal margin to apex. In ventral view outer margin of mandible with a shallowly rounded prebasal inflected angle. MI 16 - 20.

Dentition. Basally with a dental row of 7 sharply triangular teeth of which 1 - 5 are slightly taller than 6 - 7, followed distally by 4 minute denticles and a slightly enlarged apical tooth, giving a total dental count of 12.

Basal lamella of mandible truncated-triangular, not in the same plane as the principal basal dental series but located below and slightly behind the basal tooth; not visible in full-face view with the mandibles closed.

Labrum terminates in a pair of narrow elongate triangular lobes.

Clypeus with anterior margin broad, transverse to extremely shallowly convex. Lateral margins of clypeus approximately straight and more or less parallel. Anterolateral clypeal angles located much further laterally than the point at which the outer margins of the mandibles intersect the anterior clypeal margin.

Clypeal margins and dorsum with minute appressed pubescence only, appearing naked under normal magnifications. In ventral view the lateral clypeal margins extending well beyond the outer margins of the fully closed mandibles through the basal third or more of their length.

Preocular carina visible anteriorly in full-face view but behind level of frontal lobes concealed from view by frontal carina.

Ventrolateral margin of head between eye and mandible marginate, the margination curving shallowly upward anteriorly from the eye; not terminating anteriorly in a tooth. Postbuccal impression small and inconspicuous.

Cuticle of side of head within scrobe finely shagreenate or punctulate.

Scape short, SI 51 - 57, dorsoventrally flattened, the dorsum and ventre converging anteriorly so that the leading edge is a sharp flange or even a thin lamella. Ventral surface of scape behind the leading edge concave.

Leading edge of scape with a row of large spatulate hairs, one or more of which are curved toward the base of the scape.

Pronotum sharply marginate laterally, without a median longitudinal carina.

Propodeum with a broad lamella running down the declivity; lamella incorporating the propodeal teeth which are indistinguishable except for a minute apical point.

Spongiform appendages massively developed on waist segments. Base of first gastral stemite in profile with a distinct pad of dense spongiform tissue.

Pilosity. Pronotal humeri without projecting hairs of any description. A single pair of feebly clavate hairs present at highest point of vertex, otherwise dorsolateral and dorsal surfaces of head without standing pilosity; apicoscrobal hair absent. Dorsal surfaces of alitrunk, waist segments and first gastral tergite without standing pilosity. Ground-pilosity of head minute, sparse and appressed, very inconspicuous.

Sculpture. Dorsum of head behind clypeus finely reticulate-punctate. Dorsum and sides of alitrunk mostly to entirely smooth and shining; disc of postpetiole smooth.

Type Material

Bolton (2000):

Holotype worker, ITALY: Napoli, Portici (Museo Civico di Storia Naturale, Genoa) [examined].
Strumigenys membranifera r. simillima Emery, 1890: 69, pl. 8, fig. 5. Holotype worker, ANTILLES IS: St Thomas I. (MCSN) [not seen].
Strumigenys membranifera var. santschii Forel, 1904b: 6. Syntype workers, TUNISIA: Kairouan (F. Santschi) (Musee d'Histoire Naturelle Genève) [examined].
Strumigenys (Cephaloxys) vitiensis Mann, 1921: 461, fig. 22. Syntype workers, FIJI IS: Vanua Levu, Lasema (W'M. Mann) (Museum of Comparative Zoology) [examined].
Strumigenys (Cephaloxys) silvestriana W.M. Wheeler, 1928a: 27. Syntype workers, MACAO: (F. Silvestri) (MCZ) [examined].
Strumigenys (Cephaloxys) foochowensis W.M. Wheeler, 1928a: 28. Holotype queen, CHINA: Foochow (F. Silvestri) (location of holotype unknown, not in MCZ, presumed lost).
Strumigenys (Cephaloxys) membranifera var. marioni W.M. Wheeler, 1933: 276. Syntype workers, U.S.A.: Mississippi (M.R. Smith) (MCZ) [examined].
Strumigenys (Cephaloxys) membranifera var. williamsi W.M. Wheeler, 1933 : 276. Syntype workers, HAWAII: S of Olaa, off the road to Puna, iv.1932, under moss etc. (F. X. Williams) (MCZ) [examined].

References

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