AntTRAILS

The genomic legacy of major human transitions in Mediterranean ants across history

Marie Skłodowska-Curie ActionsGrant agreement No 101206623Museum für Naturkunde Berlin

Pheidole pallidulaTepelenë, Albania · Hjalte KjærbyCC BY-NC · iNaturalist
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Pheidole species recorded per 300 km cellKass et al. 2022

01The ant

The big-headed ants

Pheidole is one of the largest genera of ants, more than 1,100 species are known. A colony raises two worker castes from the same mother: small minors, which do most of the work, and larger majors, whose oversized heads are packed with mandible muscle, a bite strong enough to open food the minors cannot. In some species the majors also serve as living reservoirs, storing food in their gasters.

The range of Pheidole pallidula as it was mapped before 2016

02The name

The Mediterranean big-headed ants

Around the Mediterranean the common native one is small, running from pale to dark brown, and it is everywhere: under stones, along paths, in the cracks of every city in the basin.

For more than a century and a half it was considered a single species. Everything from Portugal to the Caucasus was called Pheidole pallidula.

  • Pheidole pallidula
Ranges of the four species in the Pheidole pallidula group

03The revision

From one to four cryptic species

In 2016 Bernhard Seifert measured 287 major workers from 112 nests across seventeen morphometric characters and found four species inside that one name. Three of them overlap broadly in the Balkans and Asia Minor.

Pheidole pallidula group

  • P. pallidula
  • P. cicatricosa
  • P. balcanica
  • P. koshewnikovi
Trade routes across the Mediterranean, 600 BCE to 1500 CE

04The question

Everything moved

Since the rise of civilisation people have carried other species with them, occasionally on purpose and far more often by accident. Ants travel with us particularly well: they like the places we build, they eat what we produce, and they are small enough to arrive unnoticed.

Some of the oldest and busiest of those routes run along the same coasts where these species meet.

A distribution map cannot say whether a population has always been there. A genome can, and that is what AntTRAILS sets out to read.

  • 600–500 BCEThe Levant and the Red Sea
  • 350 BCEThe Hellenistic world
  • 150–200 CERoman sea lanes
  • 200 CERoman roads and sea
  • 1300–1500 CEEurope, the Levant and North Africa

Routes after OWTRAD (Ciolek 1999), CC BY-NC.
Straight lines between ports.

Ranges of the four species in the Pheidole pallidula group
Pheidole pallidula group
  • P. pallidula
  • P. cicatricosa
  • P. balcanica
  • P. koshewnikovi

05The aim

What AntTRAILS asks

  1. How many, and whereFour species were told apart by linear morphometry. A genome can test whether they really are four, and show where each one lives.
  2. Which were carried, and whenSome populations have always been there, others came with people. Their genomes can tell the two apart, and roughly when the newcomers arrived.
  3. What happens where they meetThree of the four live side by side across wide areas. Do they still interbreed where they meet, and how has that shaped them?

06The model

Why the Mediterranean big-headed ants

  1. CommonAmong the most widespread ants in the region, and easy to find and collect.
  2. At home with usThey live in the places we build and eat what we grow, so they have always had the chance to travel with us.
  3. CrypticThe four are told apart by morphometry rather than by eye, so one arriving where another already lives can pass for it.
  4. Together and apartThe species meet in some areas and not in others, which is what makes gene flow possible to measure.
  5. Deeply dividedMore than fifteen percent divergence between mitochondrial lineages, apparently even within the same species: enough resolution to read populations, not only species.
  6. Its own controlOther Pheidole live here too: recent arrivals like P. megacephala and P. indica, and one native from another group, P. sinaitica.
Pheidole pallidulaTepelenë, Albania · Hjalte KjærbyCC BY-NC · iNaturalist
Collection localities across the Mediterranean

07The sampling

What we collected

All the specimens going through the lab, plotted over the four supposed native ranges. Including many introduced populations.

08The people

The team

  • Mattia MenchettiMuseum für Naturkunde, Berlin, Germany
  • Bonnie BlaimerMuseum für Naturkunde, Berlin, Germany
  • Pablo LibradoInstitute of Evolutionary Biology (CSIC-UPF), Barcelona, Spain
  • Jelena BujanRuđer Bošković Institute, Zagreb, Croatia
  • Lidya HamechaDepartment SNV, University of Algiers, Algeria
  • Celal KaramanTrakya University, Edirne, Türkiye
  • Albena Lapeva-GjonovaSofia University, Bulgaria
  • Wala OueslatiUniversity of Tunis El Manar, Tunisia
  • Sebastian SalataUniversity of Wrocław, Poland
  • Elisabetta SbregaInstitut Botànic de Barcelona (IBB), CSIC-CMCNB, Spain
  • Enrico SchifaniInstitut de Biologia Evolutiva (CSIC-UPF), Barcelona, Spain
  • Ahmed TaheriAbdelmalek Essaâdi University, Tétouan, Morocco
  • Roger VilaInstitut de Biologia Evolutiva (CSIC-UPF), Barcelona, Spain
  • Jody VogesMuseum für Naturkunde, Berlin, Germany
Pheidole pallidulaHimarë, Albania · Hjalte KjærbyCC BY-NC · iNaturalist

09Funding

AntTRAILS is funded by the European Union under the Marie Skłodowska-Curie Actions, grant agreement No 101206623, and hosted by Bonnie Blaimer's lab at the Museum für Naturkunde Berlin, with a secondment to Pablo Librado's group at the Institute of Evolutionary Biology in Barcelona.

  • Marie Skłodowska-Curie Actions, European Commission
  • Institut de Biologia Evolutiva, CSIC-UPF

Seifert, B. (2016) Inconvenient hyperdiversity: the traditional concept of Pheidole pallidula includes four cryptic species (Hymenoptera: Formicidae). Soil Organisms 88(1): 1–17.

Kass, J.M. et al. (2022) The global distribution of known and undiscovered ant biodiversity. Science Advances 8(31): eabp9908. 10.1126/sciadv.abp9908

Ciolek, T.M. (1999–) Old World Trade Routes (OWTRAD) Project. Asia Pacific Research Online, Canberra. ciolek.com/owtrad

Photograph: Pheidole pallidula, Himarë, Albania. Hjalte Kjærby, CC BY-NC, iNaturalist.