Research Article |
Corresponding author: Henri W. Herrera ( henri.herrera@espoch.edu.ec ) Corresponding author: Gianpiero Fiorentino ( gf6@njit.edu ) Academic editor: Sebastian Salata
© 2024 Henri W. Herrera, María C. Tocora, Gianpiero Fiorentino, Charlotte E. Causton, Wouter Dekoninck, Frederik Hendrickx.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Herrera HW, Tocora MC, Fiorentino G, Causton CE, Dekoninck W, Hendrickx F (2024) The ants of the Galápagos Islands (Hymenoptera, Formicidae): a historical overview, checklist, and identification key. ZooKeys 1191: 151-213. https://doi.org/10.3897/zookeys.1191.107324
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The Galápagos ant fauna has long been understudied, with the last taxonomic summary being published almost a century ago. Here, a comprehensive and updated overview of the known ant species of the Galápagos Islands is provided with updated species distributions. The list is based on an extensive review of literature, the identification of more than 382,000 specimens deposited in different entomological collections, and recent expeditions to the islands. The ant fauna is composed of five subfamilies (Dolichoderinae, Dorylinae, Formicinae, Myrmicinae, and Ponerinae), 22 genera, 50 species, and 25 subspecies, although three species (Crematogaster crinosa Mayr, 1862, Camponotus senex (Smith, 1858), and Solenopsis saevissima (Smith, 1855)) are considered dubious records. Finally, an illustrated identification key of the species found in the archipelago is presented.
Checklist, distribution, Galápagos ants, taxonomy
Until recently, the ant fauna of the Galápagos Islands was poorly studied. Early expeditions to the Galápagos collected only a few specimens at specific sites, primarily in the arid zones, which were more accessible (
Delimitation of the three periods in the study of Galápagos ants. Period I: 1877–1933; Period II: 1933 until the end of the 1990’s; Period III: 2000 onwards. The following references correspond to the years listed in the figure: 1877 =
This paper is based on literature reviews and the study of 382,023 specimens deposited mostly in the
Terrestrial Invertebrates Collection of the Charles Darwin Research Station
(
Glossary of terminology labeled from left to right A lateral view of a major worker of Camponotus planus B profile view of terminal portion of gaster of Paratrechina longicornis (Latreille, 1802) C frontal view of Tetramorium bicarinatum (Nylander, 1846) D lateral view of Tetramorium bicarinatum. Abbreviations: ac = antenna club; acp = acidopore; c = clypeus; cx = coxa; ey = eye; fc = frontal carina; fl = frontal lobe; fu = funiculus; GA = gaster; G1, 2, 3, 4, 5 = gastral segments 1–5; HD = head; lc = lateral portion of clypeus; man = mandible; mb = basal margin of mandible; mc = median portion of clypeus; MES = mesosoma; mg = metanotal groove; mn = mesonotum; mpl = mesopleuron; mtp = metapleuron; om = occipital margin; pe = petiole; pms = promesonotal; pn = pronotum; pro = propodeum; ppt = post-petiole; ps = propodeal spine; py = pygidium; s = sting; sc = scape scb = scrobe; sp = spiracle.
Five subfamilies of Formicidae can be found in the Galápagos: Dolichoderinae, Dorylinae, Formicinae, Myrmicinae, and Ponerinae, representing 22 genera, 50 species and 25 subspecies. The subfamily Myrmicinae is the largest with 32 species, while only one species on the islands represents Dorylinae, Cylindromyrmex whymperi (Cameron, 1891). The introduced species Solenopsis globularia (Smith, 1858) (on 35 islands, islets, and/or rocks), Tetramorium bicarinatum (Nylander, 1846) (on 33), Cardiocondyla emeryi Forel, 1881 (on 30), Monomorium floricola (Jerdon, 1851) (on 27), Camponotus zonatus Emery, 1894 (on 24), Tetramorium lanuginosum Mayr, 1870 (on 24), Wasmannia auropunctata (on 21), Solenopsis geminata (Fabricius, 1804) (on 20), and Tapinoma melanocephalum (Fabricius, 1793) (on 18), are the most widely distributed species in the archipelago. Among the putative endemic species (8,
We report 50 species and 25 subspecies of ants from 22 genera from the Galápagos Islands. The number of new species and locality records in the last 15 years combined with the fact that many islands are still highly understudied demonstrates that considerable work still needs to be done to identify and understand the islands’ ant diversity.
Of the species recorded in this checklist, there are still dubious records. This is the case for Camponotus senex, Crematogaster crinosa, and Solenopsis saevissima (
Regarding the genus Camponotus, our studies suggest that the identification of the introduced ant Camponotus zonatus may have been confused with that of the only known from Galápagos species C. macilentus. This confusion is of particular interest regarding ecological studies that have cited the abundance of C. macilentus, which is typically more cryptic (
Although efforts in the last two decades have substantially increased our knowledge of the ant fauna of the Galápagos Islands, a good portion of the material studied during the last 15 years came from surveys that were not focused on ants. Ants remain poorly studied, and systematic sampling of the archipelago is necessary. Apart from Santa Cruz (in 1982, 1984) and Floreana (in 1997 and 2005) (
1 | Mesosoma attached to the gaster by a single intermediate segment, the petiole (Fig. |
2 |
– | Mesosoma attached to the gaster by two intermediate segments, the petiole, and post-petiole (Fig. |
Myrmicinae |
2 | Gaster with a slight to remarkable constriction between its first and second segments (Fig. |
3 |
– | Gaster without constriction between its first and second segments (Fig. |
4 |
3 | Pygidium with small spines or denticles (Fig. |
(Dorylinae) Cylindromyrmex whymperi |
– | Pygidium without spines or denticles (Fig. |
Ponerinae |
4 | Apex of abdomen with a circular orifice surrounded by a fringe of short setae, the acidopore, formed from the hypopygium (Fig. |
Formicinae |
– | Apex of abdomen without acidopore (Fig. |
Dolichoderinae |
1 | In lateral and dorsal views, petiole visible (Fig. |
Dorymyrmex pyramicus albemarlensis |
– | Petiole squamiform and notably reduced, in lateral and dorsal view, hidden under the first segment of the gaster (Fig. |
2 |
2 | Small (~ 1.5 mm), head mesosoma and antennae pale brown, legs and gaster pale yellow (Fig. |
Tapinoma melanocephalum |
– | Ants measuring ~ 2 mm with body and legs brown-gray (Fig. |
Tapinoma sp. hh07 |
Dorymyrmex pyramicus albemarlensis Wheeler, 1919
Fig.
Remarks. In
Taxonomic history.
Distribution. Possibly endemic: Baltra, Bartolomé, Daphne Mayor, Edén, Española, Fernandina, Genovesa, Isabela (VA, VD, VW), Marchena, Pinta, Rábida, Santa Cruz, Santa Fé, Santiago (
Tapinoma melanocephalum (Fabricius, 1793)
Fig.
Remarks. Originally described as Formica melanocephalum (Fabricius, 1793). First published record
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Albany, Baltra, Champion, Española, Fernandina, Floreana, Genovesa, Isabela (CA, SN, VA), Marchena, Pinta, Plaza Sur, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé (
New record. Mariela Mediana Islet.
Tapinoma sp. hh07
Fig.
Remarks. In
Distribution. Undetermined origin: Santa Cruz (
Cylindromyrmex whymperi (Cameron, 1891)
Fig.
Remarks. Originally described as Holcoponera whymperi (Cameron, 1891). Cited as Cylindromyrmex striatus in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Baltra, Fernandina, Isabela (VA, VW), Santa Cruz (Herrera and al. 2020).
New record. Santiago Island.
1 | Antenna, including scape, with 9 segments (Fig. |
Brachymyrmex heeri |
– | Antenna, including scape, with > 9 segments (Fig. |
2 |
2 | Polymorphic, minor workers > 4 mm (total length); antennal insertions located distantly from posterior margin of the clypeus (Fig. |
3 |
– | Monomorphic, workers of small size, < 4 mm (total length), with antennal insertions located near to posterior margin of clypeus (Fig. |
5 |
3 | In lateral view, promesonotum and dorsum of propodeum flat; propodeal declivity angulate (Fig. |
Camponotus planus |
– | In lateral view, promesonotum and propodeum rounded until the base of the declivity of propodeum, forming a single convexity (Figs |
4 |
4 | Longitudinal carina visible in middle of the clypeus (major workers); head in frontal view with frontal carinae closing towards the middle of eyes; mesosoma with > 10 erect hairs (Fig. |
Camponotus zonatus |
– | Longitudinal carina in the middle of clypeus inconspicuous or absent (major workers); head in frontal view with frontal carinae opening from base of fronto-clypeal suture towards middle of eyes; mesosoma with < 10 erect hairs (Fig. |
Camponotus macilentus |
5 | Scape obviously elongate without erect setae and extending at least twice the length of the head in lateral view (Fig. |
Paratrechina longicornis |
– | Scape with abundant erect setae and never extending twice the length of the head in lateral view (Fig. |
6 |
6 | Head, mesosoma, gaster and legs dark brown with trochanters yellowish; mesopleuron and metapleuron smooth and shiny (Fig. |
Nylanderia steinheili |
– | Species without combination of characteristics described above | Nylanderia spp. |
Brachymyrmex heeri Forel, 1874
Fig.
Remarks. First published record in
Taxonomic history.
Distribution. Nearctic, Neotropical, and Palearctic.
Galápagos distribution. Introduced: Floreana, Isabela (SN, VE), Marchena, San Cristóbal, Santa Cruz (
Camponotus zonatus Emery, 1894
Fig.
Remarks. First published record (
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Bainbridge #1, Bainbridge #3, Bainbridge #4, Bainbridge #5, Bainbridge #6, Baltra, Champion, Cuevas, Daphne Mayor, Eden, Floreana, Genovesa, Isabela (CA, SN, VA, VD, VW), Mao, Marchena, Pinta, Pinzón, Plaza Norte, Plaza Sur, San Cristóbal, Santa Cruz, Santa Fé, Santiago, Seymour Norte (
Camponotus macilentus Smith, 1877
Fig.
Remarks. Cited as Camponotus (Myrmamblys) macilentus in
Taxonomic history.
Distribution. Endemic: Baltra, Champion, Española, Fernandina, Floreana, Genovesa, Isabela (SN, VA, VD, VW), Marchena, Pinta, Pinzón, Plaza Norte, Rábida, Santa Cruz, Santa Fé, Santiago, (Herrera and al. 2020).
Citations.
Camponotus macilentus albemarlensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. albemarlensis Wheeler, 1919: 284. Camponotus (Pseudocolobopsis) macilentus var. albemarlensis in
Taxonomic history.
Distribution. Endemic: Isabela Island.
Camponotus macilentus altinotus Stitz, 1932. Cited as Camponotus (Pseudocolobopsis) macilentus var. altinota Stitz, 1932: 370. Camponotus (Pseudocolobopsis) macilentus altinotus in
Taxonomic history.
Distribution. Endemic: Floreana Island (
Camponotus macilentus barringtonensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. barringtonensis Wheeler, 1919: 282. Camponotus (Pseudocolobopsis) macilentus var. barringtonensis in
Taxonomic history.
Distribution. Endemic: Santa Fé Island (
Camponotus macilentus bindloensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. bindloensis Wheeler, 1919: 286. Camponotus (Pseudocolobopsis) macilentus var. bindloensis in
Taxonomic history.
Distribution. Endemic: Marchena Island (
Camponotus macilentus castellanus Wheeler, 1924. Cited as Camponotus (Myrmamblys) macilentus var. castellanus Wheeler, 1924: 116. Cited as Camponotus (Pseudocolobopsis) macilentus castellanus in
Taxonomic history.
Distribution. Endemic: Genovesa Island (
Camponotus macilentus duncanensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. duncanensis Wheeler, 1919: 283. Camponotus (Pseudocolobopsis) macilentus var. duncanensis in
Taxonomic history.
Distribution. Endemic: Floreana, Pinzón Islands (
Camponotus macilentus hoodensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. hoodensis Wheeler, 1919: 285. Cited ad Camponotus (Pseudocolobopsis) macilentus var. hoodensis in
Taxonomic history.
Distribution. Endemic: Española Island (
Camponotus macilentus jacobensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. jacobensis Wheeler, 1919: 280. Camponotus (Pseudocolobopsis) macilentus var. jacobensis in
Taxonomic history.
Distribution. Endemic: Santiago Island (
Camponotus macilentus narboroensis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. narboroensis Wheeler, 1919: 286. Camponotus (Pseudocolobopsis) macilentus var. narboroensis in
Taxonomic history.
Distribution. Endemic: Fernandina Island (
Camponotus macilentus pervicus Wheeler, 1924. Cited as Camponotus (Myrmamblys) macilentus var. pervicus Wheeler, 1924: 115. Camponotus (Pseudocolobopsis) macilentus pervicus in
Taxonomic history.
Distribution. Endemic: Santa Cruz Island (
Camponotus macilentus sapphirinus Wheeler, 1924. Cited as Camponotus (Myrmamblys) macilentus var. sapphirinus Wheeler, 1924: 114. Cited as Camponotus (Pseudocolobopsis) macilentus sapphirinus in
Taxonomic history.
Distribution. Endemic: Santa Cruz, Baltra Islands (
Camponotus macilentus vulcanalis Wheeler, 1919. Cited as Camponotus (Myrmamblys) macilentus var. vulcanalis Wheeler, 1919: 284. Camponotus (Pseudocolobopsis) macilentus var. vulcanalis in
Taxonomic history.
Distribution. Endemic: Isabela Island (
Camponotus macilentus wollebaeki Stitz, 1932. Cited as Camponotus (Myrmamblys) macilentus var. wollebaeki Stitz, 1932: 371. Camponotus (Pseudocolobopsis) macilentus wollebaeki in
Taxonomic history.
Distribution. Endemic: Floreana Island (
Camponotus planus Smith, 1877
Fig.
Remarks. Cited as Camponotus (Myrmorhachis) planus in
Taxonomic history.
Distribution. Endemic: Bainbridge #1, Baltra, Bartolomé, Cousin, Fernandina, Floreana, Isabela (CA, SN, VA, VD, VE, VW), Logie, Marchena, Pinzón, Plaza Sur, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé (
Citations.
Camponotus planus fernandinensis Wheeler, 1919. Cited as Camponotus (Myrmorhachis) planus var. fernandinensis Wheeler, 1919: 296. Camponotus (Myrmocladoecus) planus fernandinensis in
Taxonomic history.
Distribution. Endemic: Fernandina Island (
Camponotus planus fidelis Wheeler, 1919. Cited as Camponotus (Myrmorhachis) planus var. fidelis Wheeler, 1919: 295. Camponotus (Myrmocladoecus) planus var. fidelis in
Taxonomic history.
Distribution. Endemic: Santa Fé (
Camponotus planus hephaestus Wheeler, 1933. Cited as Camponotus (Myrmorhachis) planus var. hephaestus Wheeler, 1933: 59. Camponotus (Myrmocladoecus) planus hephaestus in
Taxonomic history.
Distribution. Endemic: Isabela Island (
Camponotus planus indefessus Wheeler, 1919. Cited as Camponotus (Myrmorhachis) planus var. indefessus Wheeler, 1919: 294. Camponotus (Myrmocladoecus) planus indefessus in
Taxonomic history.
Distribution. Endemic: Santa Cruz Island (
Camponotus planus isabelensis Wheeler, 1919. Cited as Camponotus (Myrmorhachis) planus var. isabelensis Wheeler, 1919: 293. Camponotus (Myrmocladoecus) planus isabelensis in
Taxonomic history.
Distribution. Endemic: Isabela Island (
Camponotus planus peregrinus Emery, 1893. Cited as Camponotus peregrinus Emery, 1893: 91. Camponotus (Myrmorhachis) planus peregrinus in
Taxonomic history.
Distribution. Endemic: Floreana, San Cristóbal Island (
Camponotus planus pinzonensis Wheeler, 1919. Cited as Camponotus (Myrmorhachis) planus var. pinzonensis Wheeler, 1919: 297. Camponotus (Myrmocladoecus) planus var. pinzonensis in
Taxonomic history.
Distribution. Endemic: Pinzón Island (
Camponotus planus sansalvadorensis Wheeler, 1924. Cited as Camponotus (Myrmorhachis) planus var. sansalvadorensis Wheeler, 1924: 119. Cited as Camponotus (Myrmocladoecus) planus sansalvadorensis in
Taxonomic history.
Distribution. Endemic: Santiago Island (
Camponotus planus santacruzensis Wheeler, 1919. Cited as Camponotus (Myrmorhachis) planus var. santacruzensis Wheeler, 1919: 294. Camponotus (Myrmocladoecus) planus var. santacruzensis in
Taxonomic history.
Distribution. Endemic: Santa Cruz, Baltra Island (
Camponotus senex (Smith, 1858). Originally described as Formica senex (Smith, 1858). Cited in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Uncertain: San Cristóbal Island (
Nylanderia fulva nesiotis (Wheeler, 1919)
Remarks. Cited as Prenolepis fulva nesiotis in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Española, Isabela, Santiago, San Cristobal, Santa Cruz (
Nylanderia guatemalensis itinerans (Forel, 1901)
Remarks. Cited as Prenolepis vividula guatemalensis itinerans in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Floreana, San Cristobal, Santa Cruz (
Nylanderia steinheili (Forel, 1893)
Fig.
Remarks. Cited as Prenolepis steinheili in (Forel, 1893). First record in
Taxonomic history.
Distribution. Malagasy, Nearctic, Neotropical.
Galápagos distribution. Introduced: Floreana, Gardner (next to Floreana), Isabela (CA), Pinzón, San Cristóbal, Santa Cruz (
New record. Santiago Island.
Nylanderia vaga (Forel, 1901)
Remarks. Cited as Prenolepis vaga in (Forel, 1901). Cited as Paratrechina vaga in
Taxonomic history.
Distribution. Australasia, Indomalaya, Neotropical, Oceania.
Galápagos distribution. Introduced: Floreana, Santa Cruz, Pinta (
Paratrechina longicornis (Latreille, 1802)
Fig.
Remarks. Cited as Formica longicornis in (Latreille, 1802). Cited as Prenolepis longicornis (Latreille, 1802) in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Baltra, Bartolomé, Champion, Española, Fernandina, Floreana, Gardner (next to Española), Isabela (SN, VA), Marchena, Pinta, Rábida, Santiago, San Cristóbal, Santa Cruz, Santa Fé, Seymour Norte (
1 | Postpetiole attached to the dorsal surface of the first segment of the gaster (Crematogaster) (Fig. |
Crematogaster JTL – 022 |
– | Postpetiole attached on anterior surface of the first segment of the gaster (Figs |
2 |
2 | Antenna with 12 segments, scape included (Fig. |
3 |
– | Antenna with < 12 segments: scape included (Figs |
19 |
3 | Antennal club of two segments (Fig. |
Adelomyrmex longinoi |
– | Antennal club diffuse or with 2 or 3 segments (Fig. |
4 |
4 | Propodeum without spines (Figs |
5 |
– | Propodeum armed with spines (Figs |
8 |
5 | Posterior surface of the head and propodeal dorsum transversely striate (Fig. |
Trichomyrmex destructor |
– | Posterior surface of the head and propodeal dorsum not transversely striate (Figs |
6 |
6 | Head, mesosoma and gaster smooth and shiny (Fig. |
Monomorium floricola |
– | Head and mesosoma neither smooth nor shiny (Figs |
7 |
7 | In lateral or dorsal view, pronotum with a pair of erect setae (Fig. |
Monomorium pharaonis |
– | In lateral or dorsal view, pronotum without a pair of erect setae, only appressed pubescence present (Fig. |
Monomorium cf. pharaonis |
8 | Antennal scrobes very well marked, extending posteriorly past the eyes (Fig. |
9 |
– | Antennal scrobes absent (Figs |
13 |
9 | Propodeal spines long, strong, and acute (Figs |
10 |
– | Propodeal spines short and not acute (Figs |
12 |
10 | Sculpture on the cephalic dorsum of the head strigose (Fig. |
Tetramorium lucayanum |
– | Sculpture on the cephalic dorsum of the head alveolate or areolate (Fig. |
11 |
11 | Anterior clypeal margin with a distinct median notch or impression; median portion of the clypeus with 3 longitudinal carinae (Fig. |
Tetramorium bicarinatum |
– | Anterior clypeal margin without a median notch or impression (Fig. |
Tetramorium lanuginosum |
12 | Frontal carinae very well marked (Fig. |
Tetramorium simillimum |
– | Frontal carinae not well marked, scrobes vestigial, feebly developed (Fig. |
Tetramorium caldarium |
13 | Head in full-face view and mesosoma in dorsal view strigose (Fig. |
Rogeria curvipubens |
– | Head in full-face view and mesosoma in dorsal view with variable sculpturing, but never uniformly strigose; mesosoma with notopropodeal suture present and grooved in lateral view (Slightly reduced to absent in Cardiocondyla minutior); number of ommatidia variable | 14 |
14 | Monomorphic worker caste; dorsal view of the head and mesosoma densely foveolate with small appressed hairs (Figs |
15 |
– | Polymorphic worker caste; dorsal view of the head with the occipital corners smooth and shiny (major workers) (Fig. |
16 |
15 | Metanotal groove not impressed on the dorsum of mesosoma (Fig. |
Cardiocondyla minutior |
– | Metanotal groove impressed on the dorsum of mesosoma (Fig. |
Cardiocondyla emeryi |
16 | Major workers orange to reddish; total length ~ 2 mm (Figs |
17 |
– | Major workers dark brown to brown; total length ~ 2.5 mm (Figs |
18 |
17 | Major workers: head in frontal view with antennal scrobe weakly developed and alveolate (Fig. |
Pheidole flavens |
– | Major workers: head in frontal view with antennal scrobe absent (Fig. |
Pheidole hh01 |
18 | Promesonotum in lateral view convex until it reaches the metanotal groove (Fig. |
Pheidole megacephala |
– | Promesonotum in lateral view forming two convexities, truncated before reaching the metanotal groove (Fig. |
Pheidole williamsi |
19 | Antenna with 10 or 11 segments (Figs |
20 |
– | Antenna with < 6 segments (Fig. |
24 |
20 | Antenna with 10 segments; funiculus with 2-segmented club (Fig. |
21 |
– | Antenna with 11 segments (Figs |
27 |
21 | Large (Fig. |
Solenopsis geminata |
– | Smaller (Figs |
22 |
22 | Postpetiole greatly dilated, wider than petiole (seen from above), globose; eye with 15–25 ommatidia (Fig. |
Solenopsis globularia |
– | Postpetiole not dilated nor globose (Fig. |
23 |
23 | In full face view, occipital margin of the head slightly concave (Fig. |
Solenopsis gnoma |
– | In full face view, occipital margin of the head relatively straight to convex rather than concave (Fig. |
Solenopsis cf. basalis (hh06) |
24 | Mandibles long and straight (Figs |
25 |
– | Mandibles short and curved downwards in profile, otherwise triangular (Figs |
26 |
25 | Mandibles with a small preapical tooth, without denticles on inner border (Fig. |
Strumigenys louisianae |
– | Mandibles armed with small denticles on inner border (Fig. |
Strumigenys eggersi |
26 | Triangular mandibles armed with denticles (Fig. |
Strumigenys membranifera |
– | Mandibles in appearance curved and short, armed with an apical fork (Fig. |
Strumigenys emmae |
27 | Head in frontal view with the antennal insertions hidden under the frontal lobes, which are exceptionally broad or expanded (Figs |
28 |
– | Head in full face view with the antennal insertions partly visible (Fig. |
Wasmannia auropunctata |
28 | Pair of tubercles absent in the anterior median region of the pronotum (Fig. |
Cyphomyrmex nesiotus |
– | Pair of tubercles present in the anterior median region of the pronotum (Figs |
29 |
29 | In dorsal view, propodeal declivity with a pair of tubercles located at the level of spiracles (Fig. |
Cyphomyrmex sp. hh04 |
– | In dorsal view, propodeal declivity without a pair of tubercles situated at level of spiracles (Fig. |
Cyphomyrmex rimosus |
Adelomyrmex longinoi Fernández, 2003
Fig.
Remarks. Misidentification in
Taxonomic history.
Distribution. Central America.
Galápagos distribution. Introduced: Isabela Island (
Cardiocondyla emeryi Forel, 1881
Fig.
Remarks. Cited in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Albany, Bainbridge #1, Bainbridge #3, Bainbridge #4, Bainbridge #5, Bainbridge #6, Bainbridge #8, Bar, Cousin, Darwin, Eden, Fernandina, Floreana, Gardner (next to Floreana), Genovesa, Gran Felipe, Isabela (CA, SN, VA, VD, VE, VW), Mariela Grande, Mao, Marchena, Pinta, Pinzón, Plaza Sur, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé, Wolf (
Cardiocondyla minutior Forel, 1899
Fig.
Remarks. Cited as Cardiocondyla nuda in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania.
Galápagos distribution. Introduced: Bainbridge #1, Cousin, Daphne Mayor, Darwin, Fernandina, Floreana, Gardner (next to Floreana), Isabela (CA, SN, VA, VD, VE), Mariela Grande, Marchena, Pinta, Santiago, San Cristóbal, Santa Cruz, Santa Fé, Wolf (
Crematogaster crinosa Mayr, 1862
Remarks. Cited as Crematogaster (Orthocrema) brevispionsa chatamensis in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Uncertain: San Cristóbal (
Crematogaster JTL-022
Fig.
Remarks. First published record
Distribution. Introduced: San Cristóbal Island (
Cyphomyrmex nesiotus Snelling & Longino, 1992
Fig.
Remarks. Cited in
Taxonomic history.
Distribution. Probably endemic: Isabela (
Cyphomyrmex rimosus (Spinola, 1851)
Fig.
Remarks. First published record
Taxonomic history.
Distribution. Nearctic and Neotropical.
Galápagos distribution. Introduced: Gardner (next to Floreana), Isabela (SN), San Cristóbal, Santa Cruz (
Cyphomyrmex sp. hh04
Fig.
Remarks. First published record as dark form of C. rimosus in
Distribution. Origin uncertain: Isabela (SN) Pinzón, Santa Cruz (
New record. Santiago Island.
Monomorium floricola (Jerdon, 1851)
Fig.
Remarks. Originally cited as Atta floricola in (Jerdon, 1851). Cited as Monomorium floreanun in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Bainbridge #5, Baltra, Bartolomé, Bayas, Bowditch South, Champion, Cousin, Daphne Mayor, Española, Fernandina, Floreana, Gardner (next to Floreana), Genovesa, Isabela (CA, SN, VA, VD), Mariela Grande, Mariela Mediana, Marchena, Pinta, Plaza Norte, Plaza Sur, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé (
New record. Sombrero Chino.
Monomorium pharaonis (Linnaeus, 1758)
Fig.
Remarks. Originally cited as Formica pharaonis in (Linnaeus, 1758). Galápagos first published record in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Baltra, Isabela (SN), Pinta, Santa Cruz (
Monomorium cf. pharaonis
Fig.
Remarks. First record in
Distribution. Undetermined origin: Baltra, Fernandina, Floreana, Isabela (SN), Marchena, Pinta, San Cristóbal, Santa Cruz, Santa Fé (
Pheidole flavens Roger, 1863
Fig.
Remarks. Cited in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Isabela (CA, SN, VA, VD, VW), San Cristóbal, Santa Cruz (
Pheidole megacephala (Fabricius, 1793)
Fig.
Remarks. Originally cited as Formica megacephala (Fabricius, 1793). Cited in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Isabela (SN), San Cristóbal, Santa Cruz (
Pheidole williamsi Wheeler, 1919
Fig.
Remarks. Cited as Pheidole williamsi in (
Taxonomic history.
Distribution. Possibly endemic: Albany, Bainbridge #1, Bainbridge #2, Bainbridge #3, Bainbridge #4, Bainbridge #5, Bainbridge #6, Baltra, Bowditch South, Daphne Mayor, Fernandina, Floreana, Gardner (next to Floreana), Isabela (SN, VA, VD, VW), Mariela Grande, Mariela Mediana, Pinta, Plaza Sur, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé, Tortuga (
New records. Bartolomé and Beagle.
Pheidole sp. hh01
Fig.
Remarks. In
Distribution. Origin uncertain: Bowditch South, Eden, Floreana, Isabela (CA, SN, VA, VD, VE, VW), Logie, Pinzón, Santiago, San Cristóbal, Santa Cruz (
Rogeria curvipubens Emery, 1894
Fig.
Remarks. Galápagos first published record (
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Isabela (SN), San Cristóbal, Santa Cruz (
Solenopsis geminata (Fabricius, 1804)
Fig.
Remarks. Originally cited as Atta geminata in (Fabricius, 1804). Cited as Solenopsis geminata in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Albany, Bainbridge #1, Baltra, Bayas, Champion, Cuevas, Eden, Enderby, Fernandina, Floreana, Gardner (next to Floreana), Isabela (CA, SN, VA), Mariela Grande, Mao, Mariela Mediana, Plaza Sur, Santa Fé, Santiago, San Cristóbal, Santa Cruz, Seymour Norte (
Solenopsis globularia (Smith, 1858)
Fig.
Remarks. Originally cited as Myrmica globularia pacifica in (Smith, 1858). Cited as Solenopsis globularia pacifica in
Taxonomic history.
Distribution. Afrotropical, Nearctic, Neotropical.
Galápagos distribution. Introduced: Albany, Bainbridge #1, Bainbridge #3, Bainbridge #5, Bainbridge #7, Bainbridge #8, Baltra, Bowditch South, Champion, Daphne Mayor, Darwin, Eden, Enderby, Española, Fernandina, Floreana, Gardner (next to Española), Gardner (next to Floreana), Genovesa, Isabela (CA, SN, VA, VD, VE, VW), Mariela Grande, Mao, Mariela Pequeña, Marchena, Pinta, Pinzón, Plaza Sur, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé, Tortuga (
New record. Sombrero Chino.
Solenopsis gnoma Pacheco, Herrera & Mackay, 2007
Fig.
Remarks. Cited also in
Taxonomic history.
Distribution. Probably endemic: Albany, Bowditch South, Española, Floreana, Isabela (SN, VA, CA), Marchena, San Cristóbal, Santa Cruz (
New record. Santiago.
Solenopsis saevissima (Smith, 1855)
Remarks. Originally cited as Myrmica saevissima (Smith, 1855). Doubtful record for Galápagos (
Taxonomic history.
Distribution. Nearctic, Neotropical.
Galápagos distribution. Uncertain: Santa Cruz Island (
Solenopsis cf. basalis (hh06)
Fig.
Remarks. First record in
Distribution. Origin uncertain: Bainbridge #5, Santa Cruz, Santiago (
Strumigenys eggersi Emery, 1890
Fig.
Remarks. Galápagos first published record in
Taxonomic history.
Distribution. Indomalaya, Nearctic, Neotropical.
Galápagos distribution. Introduced: Santa Cruz (
Strumigenys emmae (Emery, 1890)
Fig.
Remarks. Originally cited as Epitritus emmae in (Emery, 1890). Cited as Quadristruma emmae in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Floreana, Isabela (SN, VA), San Cristóbal, Santa Cruz (
Strumigenys louisianae Roger, 1863
Fig.
Remarks. Cited in
Taxonomic history. In
Distribution. Nearctic, Neotropical.
Galápagos distribution. Introduced: Floreana, Isabela (CA, SN, VA), San Cristóbal, Santa Cruz (
New record. Santiago.
Strumigenys membranifera Emery, 1869
Fig.
Remarks. Galápagos first published record in
Taxonomic history. In
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Isabela (VA, VW), Santiago (
Tetramorium bicarinatum (Nylander, 1846)
Fig.
Remarks. Originally cited as Myrmica bicarinatum in (Nylander, 1846). Cited as Tetramorium guineense in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Bainbridge #1, Bainbridge #2, Bainbridge #3, Bainbridge #4, Bainbridge #5, Bainbridge #6, Bainbridge #8, Baltra, Bar, Bayas, Caldwell, Daphne Mayor, Española, Fernandina, Floreana, Gardner (next to Floreana), Gardner (next to Española), Genovesa, Guy Fawkes, Isabela (CA, SN, VA, VD, VE, VW), Mariela Grande, Mariela Mediana, Marchena, Pinzón, Plaza Norte, Plaza Sur, Rábida, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé, Sombrero Chino (
New records. Beagle #2, Beagle #3, Santiago Island.
Tetramorium caldarium (Roger, 1857)
Fig.
Remarks. Originally cited as Tetrogmus caldarium in (Roger, 1857). Cited in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Floreana, Santa Cruz (
Tetramorium lanuginosum Mayr, 1870
Fig.
Remarks. First published record (
Taxonomic history. In
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Bainbridge #3, Bainbridge #8, Baltra, Floreana, Gardner (next to Española), Isabela (VD), Pinzón, Plaza Norte, Plaza Sur, Rábida, San Cristóbal, Santa Cruz, Santa Fé, Seymour Norte, Wolf (
New records. Bainbridge #1, Bartolomé, Beagle #2, Beagle #3, Champion, Mao, Marchena, Santiago, Sombrero Chino.
Tetramorium simillimum (Smith, 1851)
Fig.
Remarks. Originally cited as Myrmica simillimum in (Smith, 1851). First published record in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Bainbridge #6, Baltra, Bar, Cousin, Daphne Mayor, Floreana, Gardner (next to Floreana), Isabela (SN, VA), Marchena, Mariela Grande, Santiago, San Cristóbal, Santa Cruz, Tortuga (
New record. Mariela Mediana.
Tetramorium lucayanum Wheeler, 1905
Fig.
Remarks. First published record in
Taxonomic history.
Distribution. Afrotropical, Neotropical, Palearctic.
Galápagos distribution. Introduced: Isabela (CA) (
New record. Isabela (SN).
Trichomyrmex destructor (Jerdon, 1851)
Fig.
Remarks. Originally cited as Atta destructor in (Jerdon, 1851). Cited as Monomorium destructor, in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Malagasy, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Baltra, Floreana, Isabela (SN), Santiago (
Wasmannia auropunctata (Roger, 1863)
Fig.
Remarks. Originally cited as Tetramorium auropunctata in (Roger, 1863). First published record in
Taxonomic history.
Distribution. Afrotropical, Australasia, Indomalaya, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Albany, Bainbridge #1, Baltra, Bowditch South, Champion, Cousin, Eden, Española, Floreana, Gran Felipe, Isabela (SN, VA, VD, VE, VW), Mao, Marchena, Pinzón, Rábida, Santiago, San Cristóbal, Santa Cruz, Seymour Norte, Santa Fé, Tortuga (
1 | Mandible elongate and linear (Figs |
2 |
– | Mandibles not elongate (Figs |
3 |
2 | Entirely dark brown (Fig. |
Odontomachus bauri |
– | Somewhat tricolored: head, antennae and legs orangish, mesosoma reddish brown and gaster dark brown (Fig. |
Odontomachus ruginodis |
3 | Mandibles falcate with apical tooth; anterior margin of clypeus triangular with carina conspicuously or slightly visible in median portion (Figs |
4 |
– | Triangular with dentate mandibles; anterior margin of clypeus without median carina (Fig. |
5 |
4 | Mandibles with basal margin distant from the anterior margin of clypeus when closed (Fig. |
Leptogenys santacruzi |
– | Mandibles with basal margin almost flush with the anterior border of clypeus when closed (Fig. |
Leptogenys cf. gorgona |
5 | Color dark red brown to black (Fig. |
Hypoponera opaciceps |
– | Color red brown to dark brown (Figs |
6 |
6 | Lateral surface of petiole relatively coarse with the dorsum somewhat rounded, not totally covered with fine appressed hairs (Fig. |
Hypoponera beebei |
– | Lateral surface of petiole somewhat more thick than coarse, with the dorsum somewhat triangular, sometimes covered by many fine appressed hairs (Fig. |
Hypoponera opacior |
Hypoponera beebei (Wheeler, 1924)
Fig.
Remarks. Originally cited as Ponera beebei in
Taxonomic history.
Distribution. Possibly endemic: Fernandina, Floreana, Isabela (CA, SN, VA, VW), Marchena, San Cristóbal, Santa Cruz, Seymour Norte, Genovesa (
Hypoponera opaciceps (Mayr, 1887)
Fig.
Remarks. Originally cited as Ponera opaciceps in (Mayr, 1887). First published record in (
Taxonomic history.
Distribution. Australasia, Indomalaya, Nearctic, Neotropical, Oceania, Palearctic.
Galápagos distribution. Introduced: Baltra, Fernandina, Floreana, Isabela (CA, SN, VA, VD) Marchena, San Cristóbal, Santa Cruz, Santiago (
Hypoponera cf. opacior (Forel, 1893)
Fig.
Remarks. Originally cited as Ponera opacior (Forel, 1893). In
Taxonomic history.
Distribution. Nearctic, Neotropical.
Galápagos distribution. Introduced: Fernandina, Floreana, Isabela (CA, SN, VA, VD, VW), San Cristóbal, Santa Cruz (
Leptogenys santacruzi Lattke, 2011
Fig.
Remarks. Cited in
Taxonomic history.
Distribution. Endemic: Isabela (VA), Santa Cruz Islands (
Leptogenys sp. gorgona (hh03)
Fig.
Remarks. Cited in
Distribution. Native: Santa Cruz.
New record. Isabela Island (SN).
Odontomachus bauri Emery, 1892
Fig.
Remarks. Cited as O. bauri in
Taxonomic history.
Distribution. Neotropical.
Galápagos distribution. Introduced: Floreana, Isabela (CA, SN), San Cristóbal, Santa Cruz (
Odontomachus ruginodis Wheeler, 1908
Fig.
Remarks. First published record in
Taxonomic history.
Distribution. Nearctic, Neotropical.
Galápagos distribution. Introduced: Santa Cruz (
We thank the Galápagos National Park Directorate (GNPD) for providing research and export permits (PC-32-14; PC-37-14; PC-27-15; PC-67-18; PC-41-19), along with the Charles Darwin Foundation (CDF) for their support and providing facilities. We are grateful to all the contributors cited in
The authors have declared that no competing interests exist.
No ethical statement was reported.
This research was supported by Swiss Friends of Galápagos (Freunde der Galápagos Inseln), Lindblad Expeditions-National Geographic Fund, Galápagos Conservancy, the Lakeside Fund of the California Academy of Sciences, the Leopold III funding, Department Entomology of
Conceptualization: HWH, FH, WD. Data curation: MCCT, HWH, GF. Formal analysis: MCCT, GF. Investigation: HWH. Methodology: MCCT, GF. Supervision: HWH. Visualization: GF, MCCT. Writing – original draft: HWH. Writing – review and editing: FH, GF, CEC, WD, MCCT.
Henri W. Herrera https://orcid.org/0000-0002-1417-4349
María C. Tocora https://orcid.org/0000-0002-5304-4010
Gianpiero Fiorentino https://orcid.org/0000-0001-6948-5032
Charlotte E. Causton https://orcid.org/0000-0002-2861-0025
Wouter Dekoninck https://orcid.org/0000-0002-7065-7310
Frederik Hendrickx https://orcid.org/0000-0002-1176-0318
All of the data that support the findings of this study are available in the main text.