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''Ptychandra'' is a
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
genus in the subfamily
Satyrinae The Satyrinae, the satyrines or satyrids, commonly known as the browns, are a subfamily of the Nymphalidae (brush-footed butterflies). They were formerly considered a distinct family, Satyridae. This group contains nearly half of the known diver ...
within the family
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a re ...
. ''Ptychandra'' was first described by
Cajetan Felder Baron Cajetan von Felder (german: link=no, Cajetan Freiherr von Felder; 19 September 1814 – 30 November 1894) was an Austrian lawyer, entomologist and liberal politician. He served as mayor of Vienna from 1868 to 1878. Life and career Felde ...
and
Rudolf Felder Rudolf Felder (2 May 1842 in Vienna – 29 March 1871 in Vienna) was an Austrian jurist and entomologist. He was mainly interested in Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and ...
in 1861. The genus comprises eight species, seven found in the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
with an additional species found on the island of
Borneo Borneo (; id, Kalimantan) is the List of islands by area, third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java Isl ...
. It is remarkable for the fact that it is one of a few genera to have undergone evolutionary radiation in the Philippine archipelago, with most other species being migrants from the Asian mainland.


Taxonomy

''Ptychandra'' was described in 1861 by
Cajetan von Felder Baron Cajetan von Felder (german: link=no, Cajetan Freiherr von Felder; 19 September 1814 – 30 November 1894) was an Austrian lawyer, entomologist and liberal politician. He served as mayor of Vienna from 1868 to 1878. Life and career Felde ...
and
Rudolf Felder Rudolf Felder (2 May 1842 in Vienna – 29 March 1871 in Vienna) was an Austrian jurist and entomologist. He was mainly interested in Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and ...
. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
is ''Ptychandra lorquinii''. Previously described as "difficult to place satisfactorily in the higher classification of the Satyrinae", it has been previously grouped with the
Elymniini The Elymniini is one of the tribes of the subfamily Satyrinae. If the subfamily Satyrinae elevate to family status, this tribe shall be treated as subfamily Elymniinae. Elymniini was formerly a large group, but recently, it is considered to be ...
and
Melanitini The Melanitini (evening browns and relatives) are one of the smaller tribes of the Satyrinae in the Nymphalidae (brush-footed butterfly) family. They contain the following genera: * '' Aeropetes'' Billberg, 1820 - sometimes placed in Satyrini * ...
. Fruhstorfer (1908) noted similarities with the characteristics of ''
Mycalesis ''Mycalesis'', the bushbrowns, are a genus of brush-footed butterflies. They are common in the warm regions from Central Asia to Australia, and have a high diversity in South Asia and the Wallacea. They are notably polymorphic, with wet- and d ...
'' and ''
Orsotriaena ''Orsotriaena medus'' is a butterfly found in south Asia, southeast Asia, and Australia. It is the only species in the genus ''Orsotriaena'', first described by Hans Daniel Johan Wallengren in 1858. Historical name The butterfly has historic ...
'' rather than ''
Lethe In Greek mythology, Lethe (; Ancient Greek: ''Lḗthē''; , ), also referred to as Lemosyne, was one of the five rivers of the underworld of Hades. Also known as the ''Ameles potamos'' (river of unmindfulness), the Lethe flowed around the cav ...
''. Reuter (1897) instead puts ''Ptychandra'' with '' Neorina'' in his Lethidi. Niklas Wahlberg as part of The Nymphalidae Systematics Group considers ''Ptychandra'' in the subtribe Lethina in the
Satyrini The Satyrini is one of the tribes of the subfamily Satyrinae. It includes about 2200 species and is therefore the largest tribe in the subfamily which comprises 2500 species. Distribution Satyrini butterflies have a worldwide distribution, ...
. ''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. Four of the species are
allopatric Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
, 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.


Description

Males sport an iridescent blue-purple upperside wings with sometimes white or blueish subapical or antemarginal markings. This blue color is the result of the presence of scutes featured by the wing scales, which are layered and reflect blue light. The underside of the wings are brown in color, with marbled patterning of a darker brown, in addition to wavy transverse lines in the discal area. Some species sport silvery lines on their wing undersides. The underside sport a series of submarginal ocelli between the wing cells R5 to 1A + 2A. The ocelli patterning feature "imaginary outermost rings", scales which border the ocelli proper. The R5 cell often larger and inwardly displaced. The curbital branches of the forewing are displaced by modified scales which are covered by a
Hair-pencil Hair-pencils and coremata are pheromone signaling structures present in lepidopteran males. Males use hair-pencils in courtship behaviors with females. The pheromones they excrete serve as both aphrodisiacs and tranquilizers to females as well a ...
between veins CuA1 and M3, with the exception of ''Ptychandra schadenbergi'', and a hair pencil pouched between veins CuA1 and CuA2 one third of the length of the wing and found in the interneural furrow, with the exception of ''P. talboti''. The eyes of the butterfly are hairy, as are the palpi on the second segment. Antennae are half the length of the costa. The butterfly exhibits tibial spurs. The forewing cell of the male is one third of the length of the costa. Vein M3 is colinear with the lower discocellular vein. The median discocellular vein is almost straight between the origins for veins M2 and M3, slightly angled towards the origin of vein M2. This makes it meet with the lower discocellular vein, at an approximate right angle with it. The origins of both the subcostal and cubital veins are both slightly swollen at the base, and the anal vein is slightly less swollen. The hindwing cell is closed and is around half the length of the wing. The vein Sc + R1 is slightly inflated towards the humeral vein. The M3 vein extends past the margin of the wing to form a tail, with lobes at the edges of veins CuA1 and CuA2 for some species. The origins for veins CuA1 and M2 are separated from each other. Female ''Ptychandra'' butterflies have a light brown wing upperside, with white or whitish patterns as well as a broad white subapical band on the forewing. The underside of the wings are brown and marked with darker brown wavy lines. There are submarginal ocelli akin to the males, but often larger in size. The wing venation resembles the male butterflies, with displaced cubital branches despite the lack of any brands of pouches that the males have. The females' bodies are similar to the males, but with antennae shorter than the length of the forewing.


Life history

The larval stages of ''Ptychandra'' are poorly known. The larvae of ''Ptychandra lorquinii'' are known to feed on grasses in the genera ''
Bambusa ''Bambusa'' is a large genus of clumping bamboos. Most species of ''Bambusa'' are rather large, with numerous branches emerging from the nodes, and one or two much larger than the rest. The branches can be as long as 11 m (35 ft). The ...
'' and ''
Dinochloa ''Dinochloa'' is a genus of tropical clumping high-climbing bamboos in the grass family. These species bear zigzag culms and fleshy fruits. They are found in the hill forests and lowland dipterocarp forest of southern China, Southeast Asia, a ...
''. The species are primarily woodland and forest butterflies found at an elevation between sea level and , with additional records of a ''Ptychandra lorquinii lorquinii'' at . 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 ''Neptis'' is a large genus of butterflies of Old World tropics subtropics. They are commonly called sailer butterflies or sailers, or more precisely typical sailers to distinguish them from the related blue sailer ('' Pseudoneptis''). Descri ...
'' 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 In Greek mythology, Lethe (; Ancient Greek: ''Lḗthē''; , ), also referred to as Lemosyne, was one of the five rivers of the underworld of Hades. Also known as the ''Ameles potamos'' (river of unmindfulness), the Lethe flowed around the cav ...
'' or ''
Zemeros ''Zemeros'' is a genus of butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, ...
''.


Distribution

A majority of the species in ''Ptychandra'' are found in the Philippines, with the remainder being found in Borneo. Five of the seven then known species are found in the
Philippine archipelago As an archipelago, the Philippines comprises about 7,641 islands clustered into three major island groups: Luzon, the Visayas, and Mindanao. Only about 2,000 islands are inhabited,
, an additional species being found in
Palawan Palawan (), officially the Province of Palawan ( cyo, Probinsya i'ang Palawan; tl, Lalawigan ng Palawan), is an archipelagic province of the Philippines that is located in the region of Mimaropa. It is the largest province in the country in ...
, and the final species being found in Northern
Borneo Borneo (; id, Kalimantan) is the List of islands by area, third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java Isl ...
. It is considered one of the few butterfly genera to have undergone
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rap ...
in the Philippine archipelago, with most species found in the Philippines being migrants from Mainland Asia. ''Ptychandra leucogyne'' is found on the islands of Camiguin de Luzon,
Cebu Cebu (; ceb, Sugbo), officially the Province of Cebu ( ceb, Lalawigan sa Sugbo; tl, Lalawigan ng Cebu; hil, Kapuroan sang Sugbo), is a province of the Philippines located in the Central Visayas region, and consists of a main island and ...
,
Leyte Leyte ( ) is an island in the Visayas group of islands in the Philippines. It is eighth-largest and sixth-most populous island in the Philippines, with a total population of 2,626,970 as of 2020 census. Since the accessibility of land has be ...
,
Luzon Luzon (; ) is the largest and most populous island in the Philippines. Located in the northern portion of the Philippines archipelago, it is the economic and political center of the nation, being home to the country's capital city, Manila, ...
,
Masbate Masbate, officially the Province of Masbate ( Masbateño: ''Probinsya san Masbate''; tl, Lalawigan ng Masbate), is an island province in the Philippines located near the midsection of the nation's archipelago. Its provincial capital is Masbate ...
,
Mindanao Mindanao ( ) ( Jawi: مينداناو) is the second-largest island in the Philippines, after Luzon, and seventh-most populous island in the world. Located in the southern region of the archipelago, the island is part of an island group of t ...
,
Negros Negros is the fourth largest and third most populous island in the Philippines, with a total land area of . Negros is one of the many islands of the Visayas, in the central part of the country. The predominant inhabitants of the island region a ...
,
Panay Panay is the sixth-largest and fourth-most populous island in the Philippines, with a total land area of and has a total population of 4,542,926 as of 2020 census. Panay comprises 4.4 percent of the entire population of the country. The City o ...
,
Samar Samar ( ) is the third-largest and seventh-most populous island in the Philippines, with a total population of 1,909,537 as of the 2020 census. It is located in the eastern Visayas, which are in the central Philippines. The island is divided in ...
, and
Sibuyan Sibuyan is a crescent-shaped island, the second largest in an archipelago comprising Romblon Province, Philippines. Located in the namesake Sibuyan Sea, it has an area of and has a total population of 62,815 as of 2020 census. The island has t ...
. ''P. lorquinii'' is found on
Basilan Basilan, officially the Province of Basilan ( cbk, Provincia de Basilan; yka, Wilayah Basilanin; tsg, Wilaya' sin Basilan; fil, Lalawigan ng Basilan), is an island province of the Philippines located primarily in the Bangsamoro Autonomous Re ...
,
Biliran Biliran, officially the Province of Biliran ( Waray-Waray: ''Probinsya han Biliran''; ceb, Lalawigan sa Biliran; tl, Lalawigan ng Biliran), is an island province in the Philippines located in the Eastern Visayas region (Region VIII). Biliran is ...
,
Bohol Bohol (), officially the Province of Bohol ( ceb, Lalawigan sa Bohol; tl, Lalawigan ng Bohol), is an island province of the Philippines located in the Central Visayas region, consisting of the island itself and 75 minor surrounding islands. It ...
, Camiguin de Luzon, Leyte, Luzon, Marinduque, Mindanao,
Mindoro Mindoro is the seventh largest and eighth-most populous island in the Philippines. With a total land area of 10,571 km2 ( 4,082 sq.mi ) and has a population of 1,408,454 as of 2020 census. It is located off the southwestern coast of Luz ...
,
Palawan Palawan (), officially the Province of Palawan ( cyo, Probinsya i'ang Palawan; tl, Lalawigan ng Palawan), is an archipelagic province of the Philippines that is located in the region of Mimaropa. It is the largest province in the country in ...
, Samar,
Sarangani Sarangani, officially the Province of Sarangani ( ceb, Lalawigan sa Sarangani; hil, Kapuoran sang Sarangani; Maguindanaon: ''Dairat nu Sarangani'', Jawi: دايرت نو سارڠني; fil, Lalawigan ng Sarangani), is a province in the Philipp ...
, and
Siargao Siargao is a tear-drop shaped island in the Philippine Sea situated 196 kilometers southeast of Tacloban. It has a land area of approximately . The east coast is relatively straight with one deep inlet, Port Pilar. The coastline is marked by a s ...
. Both ''P. mizunumai'' and ''P. nakamotoi'' are found on
Mount Apo Mount Apo, also known locally as Apo Sandawa, is a large solfataric, dormant stratovolcano on the island of Mindanao, Philippines. With an elevation of above sea level, it is the highest-mountain in the Philippine Archipelago, Mindanao and 24 ...
in Mindanao. ''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. ''P. talboti'' is found on
Sarawak Sarawak (; ) is a state of Malaysia. The largest among the 13 states, with an area almost equal to that of Peninsular Malaysia, Sarawak is located in northwest Borneo Island, and is bordered by the Malaysian state of Sabah to the northeast, ...
's
Mount Dulit Mount Dulit is a mountain in Borneo. It peaks at above sea level and stands at the head of the Baram River in northern Sarawak, Malaysia. It is a western outlier of the Bornean cordillera and is largely covered with tropical rainforest. It h ...
, in addition to
Mount Kinabalu Mount Kinabalu ( ms, Gunung Kinabalu, Dusun: ''Gayo Ngaran or Nulu Nabalu'') is the highest mountain in Borneo and Malaysia. With an elevation of , it is third-highest peak of an island on Earth, and 20th most prominent mountain in the worl ...
in
Sabah Sabah () is a state of Malaysia located in northern Borneo, in the region of East Malaysia. Sabah borders the Malaysian state of Sarawak to the southwest and the North Kalimantan province of Indonesia to the south. The Federal Territory ...
. There exist records of ''P. leucogyne'' on Halmahera or Batjan, but Semper (1889) states that it was probably erroneous.


Species

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


References

{{Taxonbar, from=Q13846329 Taxa described in 1861 Taxa named by Rudolf Felder Butterflies of Borneo Satyrinae Satyrini