Ptychandra

Ptychandra lorquinii, the type species

Ptychandra was described in 1861 by Cajetan von Felder and Rudolf Felder. The type species is Ptychandra lorquinii.[1][2]

Previously described as "difficult to place satisfactorily in the higher classification of the Satyrinae", it has been previously grouped with the Elymniini[2] and Melanitini. Fruhstorfer (1908) noted similarities with the characteristics of Mycalesis and Orsotriaena rather than Lethe. Reuter (1897) instead puts Ptychandra with Neorina in his Lethidi.[1] Niklas Wahlberg as part of The Nymphalidae Systematics Group considers Ptychandra in the subtribe Lethina in the Satyrini.[3]

Ptychandra talboti shows morphological differences compared to the remainder of the species found on the Philippines. It is considered as an early offshoot from the remainder of the genus.[1] Four of the species are allopatric, with the remainder P. lorquinii overlapping with P. leucogyne on the island of Luzon and the central Philippines, as well as with P. schadenbergi on the southern part of the island. It is thought that sea level changes and the subsequent isolation of butterfly populations resulted in the butterfly distribution patterns in the region. Two theories exist for the evolutionary history of Ptychandra. One theory with P. mindorana, P. negrosensis, and P. schadenbergi have staying on their original isolation sites, P. leucogyne being isolated on the central portion of the Philippines and the southern peninsular region of Luzon, and P. lorquinii being isolated on Luzon. These areas of isolation match up with the appearance of the Philippine archipelago during times of lowered sea level. A secondary hypothesis, based on the morphology of the secondary sexual characteristics (coremata) of the butterflies, postulates three ancestral spread phases. The first spread of an ancestor over the Philippines and Borneo, resulting in three types emerging that would become the various species over time with differing morphologies.[1]

Ptychandra larvae feed on grasses such as members of the genus Bambusa

The larval stages of Ptychandra are poorly known. The larvae of Ptychandra lorquinii are known to feed on grasses in the genera Bambusa and Dinochloa.[6]

The species are primarily woodland and forest butterflies found at an elevation between sea level and 700 metres (2,300 ft), with additional records of a Ptychandra lorquinii lorquinii at 2,000 metres (6,600 ft). They prefer to avoid direct sunlight, preferring dappled glades of forest instead. Males are seen in clusters settling on leaves obscured by shade and occasionally entering into flight. Males have been seen feeding on bird droppings and rotting fruit, with an observation of P. negrosensis feeding on cut grass stems with members of Amathusiid and Neptis butterflies. Different species are separated by their preferred habitat when their ranges overlap, with P. lorquinii and P. leucogyne being found in advanced secondary woodland and mature forest respectively. P. talboti is found in montane primary forest and is active during overcast weather. In flight, individuals of the genus resemble butterflies in the genus Lethe or Zemeros.[1]

A majority of the species in Ptychandra are found in the Philippines, with the remainder being found in Borneo.[7][8] Five of the seven then known species are found in the Philippine archipelago, an additional species being found in Palawan, and the final species being found in Northern Borneo.[7] It is considered one of the few butterfly genera to have undergone evolutionary radiation in the Philippine archipelago, with most species found in the Philippines being migrants from Mainland Asia.[1]

Ptychandra leucogyne is found on the islands of Camiguin de Luzon, Cebu, Leyte, Luzon, Masbate, Mindanao, Negros, Panay, Samar, and Sibuyan. P. lorquinii is found on Basilan, Biliran, Bohol, Camiguin de Luzon, Leyte, Luzon, Marinduque, Mindanao, Mindoro, Palawan, Samar, Sarangani, and Siargao.[6] Both P. mizunumai and P. nakamotoi are found on Mount Apo in Mindanao.[9][10] P. negrosensis is found on Bohol, Masbate, Negros, and West Panay. P. ohtanii is found on Leyte, Eastern Mindanao, and Sarangani. P. schadenbergi is found on Basilan and Mindanao.[6] P. talboti is found on Sarawak's Mount Dulit,[9] in addition to Mount Kinabalu in Sabah.[1] There exist records of P. leucogyne on Halmahera or Batjan, but Semper (1889) states that it was probably erroneous.[1]

There are six species of the genus Ptychandra found in the Philippines, with Badon and Nyuda, as well as Racheli and Biondi,[8] listing Ptychandra mindorana Semper, 1892 as a distinct species.[6] Funet recognizes eight total species in the genus, this interpretation is backed up by Wahlberg's species list:[9][11]