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PIMELIA (AMBLYPTERA) FORNICATIFORMIS KOCH, 1941 MOROCCO: Casablanca-Settat: Morocco Casablanca Reitter// Coll. Reitter, Pimelia scabrosa fornicatiformis det. dr. Kaszab: 10 exx. (MTM) (Fig. 10C); Casablanca // Cotypus P. sculptilis Reitt. Coll. Reitter // P. rotundipen. v. simonis Sen. det. dr. Kaszab: 6 exx (MTM); Bolhaut 3-4-35 Marokko // Sammlung Georg. Frey// Cotypus Pimelia scabrosa fornicatiformis C. Koch: 14 exx. (MTM). Marrakesh-Safi: Oualidia, Road R301, 10-IV- 1992, 32º46’N- 8º58’W, J.L. Ruiz leg.: 3 exx (MNCN)., Published as part of Mas-Peinado, Paloma, García-París, Mario, Ruiz, José L & Buckley, David, 2022, The Strait of Gibraltar is an ineffective palaeogeographic barrier for some flightless darkling beetles (Coleoptera: Tenebrionidae: Pimelia), pp. 1147-1180 in Zoological Journal of the Linnean Society 195 (4) on page 1161, DOI: 10.1093/zoolinnean/zlab088, http://zenodo.org/record/6985775

PIMELIA (MAGREBMELIA) ESCALERAI CASTRO TOVAR & FERRER, 2012 Type material: ‘ Ceuta // Paratipo// Pimelia escalerai det. Castro Tovar & J. Ferrer 2011’: (MNCN) (Fig. 10 E); ‘ De Tetuán a Ceuta, Escalera leg.// Paratipo// Pimelia escalerai det. Castro Tovar & J. Ferrer 2011’: 15 exx (MNCN). MOROCCO: Tanger-Tétouan-Al Hoceima: Tanger, (M. Escalera leg.): 30 exx (MNCN) (Fig. 10F); Embalse Dar Chaoui, 35º32’19”N- 5º50’34”W, 133 m, 6-V-2006, J.L. Ruiz leg.: 1 ex (MNCN). SPAIN (North Africa): Ciudad Autónoma de Ceuta: Cercanías Mirador Isabel II, 35º53’30”N- 5º21’48”W, 270 m, 2/ 23-V-1993, J.L. Ruiz leg.: 23 exx (MNCN); Falda Mirador Isabel II, 35º53’36”N- 5º21’44”W, 270 m, 22-VI-1993, J.L. Ruiz leg.: 3 exx (MNCN); Torre Fco. de Asís, 35º53’13”N- 5º22’03”W, 267 m, 29-IX- 2007, J.L. Ruiz leg.: 1 ex (MNCN); Cala Desnarigado, Monte Hacho, 35º53’44”N- 5º16’57”W, 15 m, 30-V-1993, J.L. Ruiz leg.: 1 ex (MNCN); Castillo del Desnarigado (cercanías), Monte Hacho, 35º53’47”N- 5º16’52”W, 65 m, 1-V-2019, J.L. Ruiz leg.: 7 exx (MNCN); Cercanías Posición A-García Aldave, 35º53’30”N- 5º22’06”W, 258 m, J.L. Ruiz leg.: 5 exx (MNCN); Pista Torre Piniers- Perímetro Fronterizo, 35º52’54”N- 5º22’07”W, 157 m, 2-VI-2020, J.L. Ruiz leg.: 2 exx (MNCN)., Published as part of Mas-Peinado, Paloma, García-París, Mario, Ruiz, José L & Buckley, David, 2022, The Strait of Gibraltar is an ineffective palaeogeographic barrier for some flightless darkling beetles (Coleoptera: Tenebrionidae: Pimelia), pp. 1147-1180 in Zoological Journal of the Linnean Society 195 (4) on page 1161, DOI: 10.1093/zoolinnean/zlab088, http://zenodo.org/record/6985775, {"references":["Castro Tovar A, Ferrer J. 2012. Nuevas especies y claves de las Pimelia Franco-Ibericas (Coleoptera, Tenebrionidae) y notas sobre Pimelia (Amblyptera) de Marruecos. Boletin de la Sociedad Entomologica Aragonesa 50: 219 - 253."]}

PIMELIA (MAGREBMELIA) ESCALERAI CASTRO TOVAR & FERRER, 2012 Type material: ‘ Ceuta // Paratipo// Pimelia escalerai det. Castro Tovar & J. Ferrer 2011’: (MNCN) (Fig. 10 E); ‘ De Tetuán a Ceuta, Escalera leg.// Paratipo// Pimelia escalerai det. Castro Tovar & J. Ferrer 2011’: 15 exx (MNCN). MOROCCO: Tanger-Tétouan-Al Hoceima: Tanger, (M. Escalera leg.): 30 exx (MNCN) (Fig. 10F); Embalse Dar Chaoui, 35º32’19”N- 5º50’34”W, 133 m, 6-V-2006, J.L. Ruiz leg.: 1 ex (MNCN). SPAIN (North Africa): Ciudad Autónoma de Ceuta: Cercanías Mirador Isabel II, 35º53’30”N- 5º21’48”W, 270 m, 2/ 23-V-1993, J.L. Ruiz leg.: 23 exx (MNCN); Falda Mirador Isabel II, 35º53’36”N- 5º21’44”W, 270 m, 22-VI-1993, J.L. Ruiz leg.: 3 exx (MNCN); Torre Fco. de Asís, 35º53’13”N- 5º22’03”W, 267 m, 29-IX- 2007, J.L. Ruiz leg.: 1 ex (MNCN); Cala Desnarigado, Monte Hacho, 35º53’44”N- 5º16’57”W, 15 m, 30-V-1993, J.L. Ruiz leg.: 1 ex (MNCN); Castillo del Desnarigado (cercanías), Monte Hacho, 35º53’47”N- 5º16’52”W, 65 m, 1-V-2019, J.L. Ruiz leg.: 7 exx (MNCN); Cercanías Posición A-García Aldave, 35º53’30”N- 5º22’06”W, 258 m, J.L. Ruiz leg.: 5 exx (MNCN); Pista Torre Piniers- Perímetro Fronterizo, 35º52’54”N- 5º22’07”W, 157 m, 2-VI-2020, J.L. Ruiz leg.: 2 exx (MNCN).

The geographic distribution of a species is shaped by its biology and by environmental and palaeogeographic factors that interact at different spatial-temporal scales, which leads to distributions and diversification patterns observed between and within lineages. The darkling beetle genus Pimelia has been diversifying for more than 31.2 Mya showing different colonization patterns after the opening of the Gibraltar Strait 5 Mya. Three of the 14 subgenera of Pimelia have populations on both sides of the Strait. Through extensive sampling and the analysis of three molecular markers, we determine levels of intra- and interspecific genetic variation, identify evolutionary lineages in subgenera, estimate their temporal origin and distribution ranges and discuss the historical basis for the geographic and diversification patterns of Pimelia around the Strait. This single geographical feature acted both as a barrier and as a dispersal route for different Pimelia species. The Strait has represented a strong barrier for the subgenus Magrebmelia since the Middle Miocene. However, the subgenera Amblyptera and Amblypteraca share repetitive signatures of post-Messinian colonization across the Strait, possibly driven by stochastic or ‘catastrophic’ events such as tsunamis. Our demographic analyses support Wallace’s hypothesis on insect dispersal stochasticity. Some taxonomic changes, including the designation of a lectotype for Pimelia maura, are also proposed.

Systematics of Amblypteraca The analysis of the molecular and morphological data sets yielded quite similar views, with populations of Larache, Assilah, and C��diz representing a fairly homogeneous unit corresponding to the mitochondrial group V, different to a point from the populations of Troia (mtDNA group IV), Tleta Rissana (mtDNA group IV) and La Mamora (mtDNA group III). All these populations are well differentiated from those from Sidi Boughaba (mtDNA group II), and also from Sidi Ifni ���Sidi Boughaba (mtDNA group I). Morphological differentiation of Tleta Rissana and La Mamora specimens is relatively well marked, also compatible with the subdivision represented in their mtDNA and mtDNA- ITS2 (Fig. 6). In these groupings, however, there were some incongruences between the nuclear (ITS2) and mtDNA markers, suggesting the potential existence of gene exchange and/or mitochondrial introgression between some of those mitocondrial-morphological groups, or potential issues of incomplete lineage sorting (ILS) among markers. The nuclear tree showed that one specimen from Troia was nested among Assilah and C��diz specimens, and was not sister to the other specimens from Troia. In addition, the specimen from Larache clustered together with specimens from Tleta Rissana, instead of being clustered with Assilah and C��diz as they were in the mtDNA tree. Discordances between mtDNA and nuclear markers in a few specimens may suggest some gene exchange between mtDNA groups III, IV and V (Larache-Tleta Rissana; Troia-C��diz/ Assilah). Specimens of La Mamora from mtDNA group III seem to retain their morphological and genetic identity, but they could differ by up to three mutations (ITS2) with respect to other populations of mtDNA groups IV and V. Since samples from La Mamora consisted of only three specimens, it is not unlikely that gene exchange could be effective among neighboring populations of clades IV and V. Consequently, we tentatively retain in a single evolutionary unit (species) all populations included in mtDNA groups III, IV and V. There is complete marker congruence, and no evidence of gene flow between clades A (I), B (II), and C (III-IV-V), and therefore with the data at hand we consider the existence of three main evolutionary units (species) within Amblypteraca: Clades A, B and C. Geographic correspondence between patterns of morphological and mtDNA variability suggests that the potential gene exchange within Clade C must have been very limited or that it is relatively recent. The close geographic proximity between Larache and Tleta Rissana, and the apparent absence of environmental barriers, makes quite likely the existence of current gene flow between them. These two population groups were probably separated by the Lukus (or Loukkos) river during the Pleistocene. Nowadays, the Lukus river forms an estuary in the area between Larache and Lixus which may have favoured the connection between both populations (Gharbaoui 1981; El Morhit et al. 2013). Specimens of Amblyteraca that inhabit sandy soils might be easily changing river shores as the flowing patterns changed when filled with sediments. On the other hand, the morphological differentiation of the Troia specimens with respect to other sequenced individuals is quite evident; however, these specimens are similar to those from populations of Ksar es Seghir (= Alcazarseguer), located at the beaches between Tangier and Ceuta (northern tip of the Tingitanian Peninsula). Ksar es Seghir was taken by the Portuguese in 1458 and remained for about 100 years as a main center for communications and transport between Lisbon and the Northern African Portuguese settlements (Braga 1998). An appealing explanation for the current presence of this Amblypteraca lineage in continental Portugal, near Lisbon, is that it is the result of a fortuitous human-mediated introduction from populations located near Ksar es Seghir. Unfortunately, these populations are mostly gone because of the construction of Tanger-Med, a mega-portuary infrastructure, and we were not able to get specimens to test this hypothesis. Lack of congruence between nuclear and mtDNA in one specimen from Troia may suggest that the Portuguese specimens were in contact with coastal Moroccan populations, likely before to their arrival to their present location in Portugal. Considering that gene flow is apparently limited, and that populations still retain their morphological singularity, we consider that the evolutionary unit corresponding to Clade C (mtDNA groups III, IVa, IVb and V), can be subdivided in four subspecific entities. By doing this, we retained an evolutionary species concept within an integrative taxonomy framework (Padial et al. 2010) for the combined mtDNA groups III, IV and V, but we also considered that the population units of La Mamora, Troia, Tleta Rissana, and Larache-Assilah-C��diz, might become isolated and independent from each other in the future. The consideration of these morphological groupings as subspecific units was already proposed by Koch (1941) and partially by Antoine (1949, 1951)., Published as part of Mas-Peinado, Paloma, Ruiz, Jos�� L., Merkl, Ott��, Buckley, David & Garc��a-Par��s, Mario, 2021, Taxonomy of the North Moroccan and Iberian species of the subgenus Amblypteraca (Coleoptera: Tenebrionidae: Pimeliinae: Pimelia), pp. 457-482 in Zootaxa 4963 (3) on pages 469-471, DOI: 10.11646/zootaxa.4963.3.4, http://zenodo.org/record/4704365, {"references":["Pallas, P. S. (1781) Icones Insectorum praesertim Rossiae Sibiriaeque quae collegit et descriptionibus illsutravit. Part IV. Erlangae: Sumtu Wolfgangi Waltheri, 97 - 104 + pl. G-H. [Published in 4 parts in 1781, 1782, 1798 and 1806? teste Hagen] https: // doi. org / 10.5962 / bhl. title. 15809","Gharbaoui, A. (1981) La terre et l'homme dans la Peninsule Tingitane. Etude sur l'homme et le milieu naturel dans le Rif occidental. Travaux de l'Institut scientifique, Rabat. Serie Geologie et Geographie physique, 15, 1 - 439.","El Morhit, M., Fekhaoui, M., Serghini, A., El Blidi, S., El Abidi, A. & Yahyaoui, A. (2013) Typology of water quality in the Loukkos river estuary (Morocco). Larhyss Journal, 12, 7 - 24.","Braga, P. D. (1998) A expansao no Norte de Africa. In: Nova Historia da Expansao Portuguesa. Vol. II, A Expansao Quatrocentista. Serrao, de. J. & Marques, A. H. de. O. (Eds.), Editorial Estampa, Lisboa, 237 - 360.","Padial, J. M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 1 - 14. https: // doi. org / 10.1186 / 1742 - 9994 - 7 - 16","Koch, C. (1941) Die Verbreitung und Rassenbildung der marokkanischen Pimelien (Col. Tenebr.) (Eine biogeographische Studie). Eos, 16, 7 - 123.","Antoine, M. (1949) Notes d'entomologie marocaine XLVIII. Tableaux de determination des Pimelia du Maroc (Col. Tenebrionidae). Annales de la Societe Entomologique de France, 116, 17 - 58.","Antoine, M. (1951) Notes d'entomologie marocaine LIV. Captures et observations nouvelles (Coleopteres). Bulletin de la Societe des Sciences Naturelles du Maroc, 31 (1950), 87 - 101."]}

Systematics of Amblypteraca The analysis of the molecular and morphological data sets yielded quite similar views, with populations of Larache, Assilah, and Cádiz representing a fairly homogeneous unit corresponding to the mitochondrial group V, different to a point from the populations of Troia (mtDNA group IV), Tleta Rissana (mtDNA group IV) and La Mamora (mtDNA group III). All these populations are well differentiated from those from Sidi Boughaba (mtDNA group II), and also from Sidi Ifni —Sidi Boughaba (mtDNA group I). Morphological differentiation of Tleta Rissana and La Mamora specimens is relatively well marked, also compatible with the subdivision represented in their mtDNA and mtDNA- ITS2 (Fig. 6). In these groupings, however, there were some incongruences between the nuclear (ITS2) and mtDNA markers, suggesting the potential existence of gene exchange and/or mitochondrial introgression between some of those mitocondrial-morphological groups, or potential issues of incomplete lineage sorting (ILS) among markers. The nuclear tree showed that one specimen from Troia was nested among Assilah and Cádiz specimens, and was not sister to the other specimens from Troia. In addition, the specimen from Larache clustered together with specimens from Tleta Rissana, instead of being clustered with Assilah and Cádiz as they were in the mtDNA tree. Discordances between mtDNA and nuclear markers in a few specimens may suggest some gene exchange between mtDNA groups III, IV and V (Larache-Tleta Rissana; Troia-Cádiz/ Assilah). Specimens of La Mamora from mtDNA group III seem to retain their morphological and genetic identity, but they could differ by up to three mutations (ITS2) with respect to other populations of mtDNA groups IV and V. Since samples from La Mamora consisted of only three specimens, it is not unlikely that gene exchange could be effective among neighboring populations of clades IV and V. Consequently, we tentatively retain in a single evolutionary unit (species) all populations included in mtDNA groups III, IV and V. There is complete marker congruence, and no evidence of gene flow between clades A (I), B (II), and C (III-IV-V), and therefore with the data at hand we consider the existence of three main evolutionary units (species) within Amblypteraca: Clades A, B and C. Geographic correspondence between patterns of morphological and mtDNA variability suggests that the potential gene exchange within Clade C must have been very limited or that it is relatively recent. The close geographic proximity between Larache and Tleta Rissana, and the apparent absence of environmental barriers, makes quite likely the existence of current gene flow between them. These two population groups were probably separated by the Lukus (or Loukkos) river during the Pleistocene. Nowadays, the Lukus river forms an estuary in the area between Larache and Lixus which may have favoured the connection between both populations (Gharbaoui 1981; El Morhit et al. 2013). Specimens of Amblyteraca that inhabit sandy soils might be easily changing river shores as the flowing patterns changed when filled with sediments. On the other hand, the morphological differentiation of the Troia specimens with respect to other sequenced individuals is quite evident; however, these specimens are similar to those from populations of Ksar es Seghir (= Alcazarseguer), located at the beaches between Tangier and Ceuta (northern tip of the Tingitanian Peninsula). Ksar es Seghir was taken by the Portuguese in 1458 and remained for about 100 years as a main center for communications and transport between Lisbon and the Northern African Portuguese settlements (Braga 1998). An appealing explanation for the current presence of this Amblypteraca lineage in continental Portugal, near Lisbon, is that it is the result of a fortuitous human-mediated introduction from populations located near Ksar es Seghir. Unfortunately, these populations are mostly gone because of the construction of Tanger-Med, a mega-portuary infrastructure, and we were not able to get specimens to test this hypothesis. Lack of congruence between nuclear and mtDNA in one specimen from Troia may suggest that the Portuguese specimens were in contact with coastal Moroccan populations, likely before to their arrival to their present location in Portugal. Considering that gene flow is apparently limited, and that populations still retain their morphological singularity, we consider that the evolutionary unit corresponding to Clade C (mtDNA groups III, IVa, IVb and V), can be subdivided in four subspecific entities. By doing this, we retained an evolutionary species concept within an integrative taxonomy framework (Padial et al. 2010) for the combined mtDNA groups III, IV and V, but we also considered that the population units of La Mamora, Troia, Tleta Rissana, and Larache-Assilah-Cádiz, might become isolated and independent from each other in the future. The consideration of these morphological groupings as subspecific units was already proposed by Koch (1941) and partially by Antoine (1949, 1951).