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Pleistocene range expansion throughout the Mediterranean and back-colonization from the Canary Islands in the legume Bituminaria bituminosa
<p><strong>Aim:</strong> Species with widespread distributions offer excellent opportunities for investigating recent biogeographical patterns across broad scales. Here, we tested the hypothesis that, due to its intermediate geographical location, NW Africa is pivotal in explaining the phylogeographical patterns of taxa with Mediterranean-Macaronesian distributions using a legume species with short generation times.</p> <p><strong>Region:</strong> Mediterranean, with a focus on NW Africa and the Canary Islands</p> <p><strong>Taxon:</strong> Pitch trefoil ( <em>Bituminaria bituminosa</em>)</p> <p><strong>Methods:</strong> We generated genetic data and performed phylogeographical and demographic analyses at two geographical scales: Mediterranean Basin (MB), using plastid sequences (115 individuals), and Macaronesia, using plastid sequences (182 individuals) and 10 nuclear microsatellite loci (220 individuals). We also performed a literature survey focusing on phylogeographical studies of other circum-Mediterranean taxa.</p> <p><strong>Results:</strong> North-west Africa was identified as a center of genetic diversity (19 out of 38 haplotypes) and demographic expansion of <em>B. bituminosa</em> in the MB during the Pleistocene. Our literature review revealed two main phylogeographical patterns in widespread species: pre-Mediterranean evergreen sclerophylls vs. Pleistocene facultative-deciduous (including <em>Bituminaria</em>) taxa, but on average both functional groups show a similar, large genetic diversity (c. 40% of haplotypes) in NW Africa. At the Macaronesian scale, we found that Canarian <em>Bituminaria </em>is composed of two genetic sublineages that coexist and hybridize on the central islands and in the mainland Macaronesian enclave (Anti-Atlas region). Demographic analyses rejected the progression rule as the model of island colonization, but strongly suggested that Anti-Atlas populations are the result of back-colonization from the easternmost islands before the Last Glacial Maximum (LGM).</p> <p><strong>Conclusions: </strong><em>Bituminaria</em> displays a pattern of Quaternary eastward expansion in the MB that appears to be paralleled by several members of its functional plant group. Thus, our study reveals a previously undescribed dual role of NW Africa in plant biogeography, acting both as a source of species expansion to the rest of the MB and a LGM refugium of plant populations with a Macaronesian island origin.</p>
FIGURE 14 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 14. Type locality and habitat of Collartida phantasma sp. nov. A) View of the eastern sector of the Federica mine, where all specimens were collected B) Entrance to the eastern sector of the Federica mine.
FIGURE 13 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 13. Details of thorax and abdomen of female Collartida phantasma sp. nov. A) Ventral view, white arrows indicating the location of the egg; B) dorsal view, temporarily pinned specimen; C) apex of abdomen and genitalia, ventrolateral view; D) closeup of meso-and-metathorax.
FIGURE 12 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 12. Dorsal views of Collartida phantasma sp. nov. female. A) Head, legs, and prothorax; B) closeup of previous. Abbreviations: apl = anterior pronotal lobe; ppl = posterior pronotal lobe.
FIGURE 11 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 11. Morphology of female of Collartida phantasma sp. nov. A) Head, legs, and prothorax, lateral view; B) head, lateral view; C) foreleg structure and armature, lateral view; D) fore tarsus, ventrolateral view; E) detail of fore tarsal apex, ventral view, arrow indicating location of campaniform sensilla; F) magnification of area indicated in previous photograph, showing the structure and location of the campaniform sensilla. Abbreviations: cs = campaniform sensilla.
FIGURE 9 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 9. Male genitalia of Collartida phantasma sp. nov. (holotype). A) Pygophore and parameres, ventral view; B) same, dorsal view; C) same, lateral view; D) detail of articulatory apparatus and base of struts.
FIGURE 10 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 10. Male genitalia of Collartida phantasma sp. nov. and congeners. A) Collartida phantasma sp. nov. (holotype), left paramere, dorsal view; B) same individual, right paramere, dorsal view; C) Collartida anophthalma, left paramere, dorsal view; D) same species, right paramere, dorsolateral view; E) Collartida phantasma sp. nov., male internal genitalia, dorsal view (drawn from holotype); F) same, closeup photograph. Abbreviations: aa = articulatory apparatus; dps = dorsal phallothecal sclerite; end = endosoma; pd = pedicel; str = endosomal struts.
FIGURE 8 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 8. Wing of male Collartida phantasma sp. nov. (holotype). The dashed line corresponds to parts of the wing that are membranous and visible only under special illumination. Blue arrow indicates crossvein that links the basal cell to Pcu.
FIGURE 7 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 7. Tarsal structure of male Collartida phantasma sp. nov. (holotype). A) Fore tarsus; B) mid tarsus; C) hind tarsus.
FIGURE 6 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 6. Foreleg structure and armature of male Collartida phantasma sp. nov. (holotype). A) Overall leg structure; B) details of coxal, trochanteral and femoral armature; C) closeup of trochanteral armature and bases of femoral spines. Abbreviations: cx = coxa; fm = femur; tb = tibia; trc = trochanter.
FIGURE 3 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 3. Antennal structure of male Collartida phantasma sp. nov. (holotype). A) Long, erect hairs on scape; B) setation at the pedicellar-basiflagellar articulation; C) setation of basiflagellomere. Abbreviations: bsf = basiflagellomere; pd = pedicel; scp = scape.
FIGURE 5 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 5. Wing muscles and scutellum of male Collartida phantasma sp. nov. (holotype). Abbreviations: IIdlm1 = musculus prophragma-mesophragmalis; IIdvm1 = musculus mesonoto-sternalis; sct = post scutellar spine.
FIGURE 4 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 4. Thorax of male Collartida phantasma sp. nov. (holotype). A) Dorsal view, with emphasis on flight musculature; dashed line indicates margin of anterior lobe; B) lateral view. Abbreviations: apl = anterior pronotal lobe; IIdlm1 = musculus prophragma-mesophragmalis; IIdvm1 = musculus mesonoto-sternalis; ppl = posterior pronotal lobe.
FIGURE 2 in A new cavernicolous assassin bug from the Canary Islands (Hemiptera: Reduviidae: Emesinae: Collartidini)
FIGURE 2. Head morphology of male Collartida phantasma sp. nov. (holotype). A) Dorsal view; B) lateral view.
Subspecies and Distribution. B.b.barbastellusSchreber,1774—EuropefromIrelandandIberianPeninsulaEtoLaivia,Belarus,Ukraine,andtheCauca-sus;alsomajorMediterraneanIs,Moroc-co,andTurkey(Anatolia);possiblyextinctinBelgium,Netherlands,andNorway. B. b. guanchae Trujillo, Ibanez & Juste, 2002 — Canary Is (Tenerife and La Gomera). in Vespertilionidae
Subspecies and Distribution. B.b.barbastellusSchreber,1774—EuropefromIrelandandIberianPeninsulaEtoLaivia,Belarus,Ukraine,andtheCauca-sus;alsomajorMediterraneanIs,Moroc-co,andTurkey(Anatolia);possiblyextinctinBelgium,Netherlands,andNorway. B. b. guanchae Trujillo, Ibanez & Juste, 2002 — Canary Is (Tenerife and La Gomera).
Subspecies and Distribution. E.i.isabellinusTemminck,1840—NEAlgeria,Tunisia,andNWLibya. E. i. boscai Cabrera, 1904 — S Portugal, S Spain, N & C Morocco, NW Algeria, and Canary Is (Lanzarote I). in Vespertilionidae
Subspecies and Distribution. E.i.isabellinusTemminck,1840—NEAlgeria,Tunisia,andNWLibya. E. i. boscai Cabrera, 1904 — S Portugal, S Spain, N & C Morocco, NW Algeria, and Canary Is (Lanzarote I).
Subspecies and Distribution. H.s.saviiBonaparte,1837—SC&EEuropefromItalytoBalkans,Anatolia,RhodesI,Levant,Crete,andCyprus.DistributionhasbeenexpandingintoCEuropeinthelast30years. H.s.austenianusDobson,1871—knownonlyfromtypelocalityinNEIndia(KhasiHills,Meghalaya). H.s.caucasicusSatunin,1901—CrimeaandCaucasusEtoWKazakhstan,WChina,andNWIndia. H. s. darwini Tomes, 1859 — SW Europe from Iberia to Switzerland including Balearics, Corsica, Sicily, Sardinia, and Montecristo I, and N Africa from N Morocco to N Tunisia along with Canary and Cape Verde Is (Sao Vicente and Santiago). in Vespertilionidae
Subspecies and Distribution. H.s.saviiBonaparte,1837—SC&EEuropefromItalytoBalkans,Anatolia,RhodesI,Levant,Crete,andCyprus.DistributionhasbeenexpandingintoCEuropeinthelast30years. H.s.austenianusDobson,1871—knownonlyfromtypelocalityinNEIndia(KhasiHills,Meghalaya). H.s.caucasicusSatunin,1901—CrimeaandCaucasusEtoWKazakhstan,WChina,andNWIndia. H. s. darwini Tomes, 1859 — SW Europe from Iberia to Switzerland including Balearics, Corsica, Sicily, Sardinia, and Montecristo I, and N Africa from N Morocco to N Tunisia along with Canary and Cape Verde Is (Sao Vicente and Santiago).
Distribution. Madeira Archipelago (Madeira and Porto Santo) and W Canary Is (La Palma, La Gomera, El Hierro, and Tenerife). Individuals classified as Pipistrellus sp. from the Azores have been suggested to be Madeira Pipistrelles. in Vespertilionidae
Distribution. Madeira Archipelago (Madeira and Porto Santo) and W Canary Is (La Palma, La Gomera, El Hierro, and Tenerife). Individuals classified as Pipistrellus sp. from the Azores have been suggested to be Madeira Pipistrelles.
Subspecies and Distribution. P.h.hesperidusTemminck,1840—EAfricainEritrea,Djibouti,Ethiopia,SWSudan,SouthSudan,andNE&SSomalia. P.h.fuscatusThomas,1901—EAfricainEDRCongo,Uganda,Kenya,Rwanda,Burundi,andTanzania. P. h. subtilis Sundevall, 1846 — S Africa in NC, C & W Angola, Zambia, Malawi, Mozambique, Zimbabwe, NC Botswana, E & South Africa, Swaziland, Lesotho, and along the W coast of Madagascar. There are also records on Canary and Cape Verde Is and in W Africa in W & SE Senegal, N Liberia, SE Ivory Coast, SC Burkina Faso, SE Ghana, SW Niger, NC Nigeria, W Cameroon, and Bioko I, although exact placement of these populations among taxa currently recognized under the P. hesperidus species complex is uncertain. A thorough investigation of records of this species is needed. in Vespertilionidae
Subspecies and Distribution. P.h.hesperidusTemminck,1840—EAfricainEritrea,Djibouti,Ethiopia,SWSudan,SouthSudan,andNE&SSomalia. P.h.fuscatusThomas,1901—EAfricainEDRCongo,Uganda,Kenya,Rwanda,Burundi,andTanzania. P. h. subtilis Sundevall, 1846 — S Africa in NC, C & W Angola, Zambia, Malawi, Mozambique, Zimbabwe, NC Botswana, E & South Africa, Swaziland, Lesotho, and along the W coast of Madagascar. There are also records on Canary and Cape Verde Is and in W Africa in W & SE Senegal, N Liberia, SE Ivory Coast, SC Burkina Faso, SE Ghana, SW Niger, NC Nigeria, W Cameroon, and Bioko I, although exact placement of these populations among taxa currently recognized under the P. hesperidus species complex is uncertain. A thorough investigation of records of this species is needed.
Subspecies and Distribution. A.l.lerviaPallas,1777—Morocco,NA.l.,andNTunisia. A.l.angusiRothschild,1921—NWNiger(Air&TermitMassifs). A.l.blaineiRothschild,1913—SELybia,NEChad,andNW&NESudan(probablynowrestrictedtoRedSeahills). A.l.fassiniLepri,1930—NWLibya,extremeSTunisia. A.l.ornatus1.GeoffroySaint-Hilaire,1827—SE&SWEgypt. A. l. sahariensis Rothschild, 1913 — S Morocco, Western Sahara, NW Mauritania, S A.l ria, extreme S Libya, NE Mali, SE Niger, and NW Chad. Introduced, free-ranging populations occur in S Spain, the Canary Is, USA (California, New Mexico, and Texas), and NE Mexico. Subspecies of free-ranging introduced populations are unknown because they originate from zoo animals of uncertain origin or from hybrids. Most introduced populations are probably from subspecies lervia, derived from European zoos. The Aoudad has become a widespread invasive species. in Bovidae
Subspecies and Distribution. A.l.lerviaPallas,1777—Morocco,NA.l.,andNTunisia. A.l.angusiRothschild,1921—NWNiger(Air&TermitMassifs). A.l.blaineiRothschild,1913—SELybia,NEChad,andNW&NESudan(probablynowrestrictedtoRedSeahills). A.l.fassiniLepri,1930—NWLibya,extremeSTunisia. A.l.ornatus1.GeoffroySaint-Hilaire,1827—SE&SWEgypt. A. l. sahariensis Rothschild, 1913 — S Morocco, Western Sahara, NW Mauritania, S A.l ria, extreme S Libya, NE Mali, SE Niger, and NW Chad. Introduced, free-ranging populations occur in S Spain, the Canary Is, USA (California, New Mexico, and Texas), and NE Mexico. Subspecies of free-ranging introduced populations are unknown because they originate from zoo animals of uncertain origin or from hybrids. Most introduced populations are probably from subspecies lervia, derived from European zoos. The Aoudad has become a widespread invasive species.
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