Phanaeus (Notiophanaeus) malyi Arnaud, 2002

Kohlmann, Bert, Arriaga-Jimenez, Alfonsina & Roes, Matthias, 2018, Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae), ZooKeys 743, pp. 67-93 : 73-76

Phanaeus (Notiophanaeus) malyi Arnaud, 2002 Fig. 8 a–b

Phanaeus pyrois malyi: Arnaud (2002: 96-97).

Phanaeus malyi Arnaud, Solís & Kohlmann (2012: 10).

Phanaeus pyrois Bates, Edmonds & Zídek (2012: 1, 3, 13, 52-53, 57)

Remarks.

Solís and Kohlmann (2012) elevated Phanaeus pyrois malyi Arnaud to species status based on the results of mitochondrial DNA studies of Costa Rican specimens that clearly separated Caribbean from Pacific populations at a genetic distance concordant with species level (average Kimura-2-parameter [K2P] = 3.8 %, Solís and Kohlmann 2012). Moreover, Ph. malyi has short and rounded postero-lateral pronotal angles (Fig. 8 a–b), whereas they are long and slender in Ph. pyrois (Fig. 8 c–d). Subsequently, Edmonds and Zídek (2012) synonymised Ph. pyrois malyi with Ph. pyrois , pending a comparative analysis of black varieties of Panamanian and South American populations. Phanaeus malyi was compared with material from J. Blackaller’s private collection and photos sent by François Génier from the Canadian Museum of Nature (Suppl. material 1: Figure S1) of black specimens of Ph. pyrois from Cerro Campana, Province of Panama, Panama (formerly treated as Ph. pyrois olsoufieffi Balthasar) and Playa de Oro, Esmeralda, Ecuador (formerly treated as Ph. pyrois funereus Balthasar). Ph. malyi and Ph. p. olsoufieffi are different taxa. While Ph. malyi has short and rounded posterolateral angles of the pronotum, similar to Ph. endymion , Ph. p. olsoufieffi has the typical pyrois long and slender posterolateral angles of the pronotum. Ph. p. olsoufieffi also exists in Costa Rica, distributed along the foot of the mountains and not in the lowlands like the typical metallic-red pyrois . A more detailed analysis would be required in order to determine if it should again be resurrected into subspecific status. In relation to Ph. p. funereus, these specimens have an intermediate form of the postero-lateral angles of the pronotum between malyi and olsoufieffi . Moreover funereus is distinctly dull black whereas malyi is a shiny black. A more detailed analysis is required for resurrecting funereus , but it is a different taxon from malyi .

We therefore revalidate Ph. malyi to full species status based on the previously mentioned mitochondrial DNA analysis ( Solís and Kohlmann 2012); the characteristic ecology and distribution of this species in tropical rainforest along the southern Pacific coast of Costa Rica, probably extending to the northern Panamanian Pacific coast ( Solís and Kohlmann 2012); and the different shapes of the pronota (Suppl. material 1: Figure S1).

Phanaeus malyi follows a characteristic distribution pattern described by Kohlmann and Wilkinson (2007) and by Solís and Kohlmann (2012), where many species living in the Costa Rican southern Pacific tropical forest are sister species of taxa found along the tropical forest of the Caribbean coast. This vicariant speciation pattern (apparently mediated by the emergence of the Talamanca Cordillera that has divided a formerly continuous tropical forest stretching from the Caribbean to the Pacific) has been observed to occur in species of dung beetles, fishes, amphibians, reptiles, and birds ( Kohlmann and Wilkinson 2007).