Corresponding author: Diana M. Percy (
Academic editor: J.N. Zahniser
The Hawaiian psyllids (
Percy DM (2017) Making the most of your host: the
The Hawaiian Islands are renowned for exemplary species radiations (e.g.
The Hawaiian Islands are home to several endemic psyllid lineages that represent multiple independent colonizations, yet an enduring puzzle is why all of these lineages are from a single psyllid family. The native psyllid fauna is composed entirely of species from the family
Many of the Hawaiian triozid species have been ascribed to endemic genera rather than placed within more widespread and established generic groupings; exceptions include taxa placed in
All Hawaiian psyllid lineages appear to exhibit high host specificity that has been conserved during in situ diversification (e.g.
species group |
authority | forms | island | habit |
---|---|---|---|---|
|
(Crawford, 1918) | L, Mo, H | free-living | |
|
sp. n. | Ma | ? | |
|
sp. n. | H | ? | |
|
sp. n. | H | ? | |
|
sp. n. | H | ? | |
|
sp. n. | H | free-living | |
|
sp. n. | H | ? | |
|
sp. n. | Mo | ? | |
|
sp. n. | O | ? | |
|
(Crawford, 1918) | O, Mo, Ma | free-living | |
|
sp. n. | H | pit galls on leaves | |
|
sp. n. | O | pit galls on leaves | |
|
(Crawford, 1918) | H | pit galls on leaves | |
|
sp. n. | Ma | ? | |
|
Kirkaldy, 1902 | K | ? enclosed galls | |
|
sp. n. | K | mainly enclosed galls on leaves: domed or donut type; occasionally stem galls | |
|
sp. n. | K | ? | |
|
sp. n. | K | ? | |
|
sp. n. | K | thin-walled cup galls on stems | |
|
sp. n. | K | thick-walled cup galls on stems | |
|
Crawford, 1925 | K | ? | |
|
sp. n. | K | ? | |
|
sp. n. | K | ? | |
|
sp. n. | K | ? | |
|
sp. n. | O | mainly enclosed galls on stems/buds; occasionally galls leaves: cone type | |
|
Crawford, 1918 | O | enclosed galls on leaves: flat type | |
|
Crawford, 1918 | L, Mo | enclosed galls on stems | |
|
Crawford, 1918 | L | ? | |
|
Crawford, 1927 | Mo | enclosed galls on stems | |
|
sp. n. | Mo | enclosed galls on leaves | |
|
sp. n. | Ma | ? | |
|
sp. n. | Ma | enclosed galls on stems | |
|
sp. n. | Ma | enclosed galls on leaves: flat type | |
|
Crawford, 1918 | H | enclosed galls on stems | |
|
sp. n. | H | enclosed galls on leaves: flat or donut type | |
|
sp. n. | H | mainly enclosed galls on leaves: cone type; occasionally stem galls |
In order to understand how the
The patterns of diversity observed in
When considering the origins of Hawaiian
Multiple shifts in galling biology on the same plant species have rarely been documented. One of the few known systems is the gall-inducing
The objectives of this study are, a) to describe the diversity of species feeding on
Over
species group |
material | n | collection information | lat N, long W | GenBank |
---|---|---|---|---|---|
|
|||||
|
holotype | f | Niulii, Hawaii, USA, ex Ohia Lehua, |
− | − |
other |
|
same data as holotype | − | ||
other |
|
Olaa Forest, Hawaii, USA, ex |
|
− | |
other |
|
Puu O Umi, Kohala, Hawaii, USA, ex |
|
− | |
|
holotype | m | Olinda Flume Road, Makawao, East Maui, USA, ex |
|
− |
paratypes |
|
same data as previous except: |
|
||
other |
|
Kamakou Preserve, Molokai, USA, on |
|
||
other |
|
Puu Kukui, boardwalk trail, West Maui, USA, ex |
|
||
other |
|
same data as previous except: “Hi49-14” D. Percy leg. ( |
|
||
other | Olinda Flume Road, Makawao, East Maui, USA, ex |
|
|||
other | same data as previous except: |
|
|||
other | same data as previous except: “Hi53-14” D. Percy leg. ( |
|
|||
|
holotype | m | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|
paratype |
|
same data as holotype | |||
other |
|
Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
||
|
other |
|
same data as previous except: ex |
|
|
other | same data as previous except: ex |
|
|||
other | same data as previous except: “Hi12-14” D. Percy leg. ( |
|
|||
other |
|
Puu Makaala, Hawaii, USA, ex |
|
||
other |
|
Gulch below Puu O Umi, Kohala, Hawaii, USA, ex |
|
||
other |
|
Humuula trail, Hawaii, USA, ex |
|
/ |
|
other |
|
Kehena, 3200ft, Kohala, Hawaii, USA, on |
− | − | |
|
holotype | f | 1868 lava flow, near Kahuku Ranch, SW Hawaii, USA, ex |
|
|
paratypes |
|
same data as holotype | − | ||
|
holotype | f | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|
|
holotype | m | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|
paratypes | Puu O Umi, Kohala, Hawaii, USA, ex |
|
|||
other |
|
Saddle Rd., 4000ft, Hawaii, USA, ex |
− | − | |
other |
|
same data as previous except: 4500-5000ft, |
− | − | |
other |
|
Olaa Forest, Hawaii, USA, ex |
|
− | |
other |
|
Kipuka off Saddle Road, Hawaii, USA, ex |
|
||
other | Koloko Drive, Honuaula, Hualalai, Hawaii, USA, ex |
|
|||
|
other | Kau, Hawaii, USA, ex |
|
||
other |
|
Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
||
other |
|
same data as previous except: ex |
|
||
|
holotype | m | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|
paratypes | same data as holotype | ||||
other |
|
same data as holotype except: ex |
|
||
other |
|
same data as holotype except: ex |
|
||
other |
|
Gulch below Puu O Umi, Kohala, Hawaii, USA, ex |
|
||
|
holotype | m | Kamakou Preserve, Molokai, USA, on |
|
− |
paratype |
|
same data as holoptype | − | ||
paratype |
|
same data as holotype except: ex |
|
− | |
paratype |
|
same data as holotype except: ex |
|
||
|
holotype | m | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|
paratypes |
|
same data as holotype | |||
|
holotype | f | Kuliouou, Oahu, USA, ex |
− | − |
other |
|
Pahoa Flats, Oahu, USA, ex |
− | − | |
other | Palolo Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− | ||
|
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
||
other | Puu Kaeo, Honolua Valley, West Maui, USA, ex |
|
|||
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− | ||
other | same data as previous except: ex |
|
− | ||
other | Wiliwilinui Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: ex |
|
− | ||
other |
|
Kamakou Preserve, Molokai, USA, ex |
|
− | |
other | same data as previous except: “Hi26-03” D. Percy leg. ( |
|
|||
other | Mnt Kaala road (culvert 55), Waianae Mnts, Oahu, USA, ex |
|
− | ||
other | Mnt Kaala summit, Waianae Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: ex |
|
− | ||
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
other | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|||
other | Mnt Kaala summit, Waianae Mnts, Oahu, USA, ex |
|
|||
other | Pupukea, N. Koolau Mnts, Oahu, USA, ex |
|
|||
|
other | Manoa Cliff trail to Pauoa Flats trail, S. Koolau Mnts, Oahu, USA, ex |
|
||
other | same data as previous except: “Hi83-14” D. Percy leg. ( |
|
|||
other |
|
Mnt Kaala summit, Waianae Mnts, Oahu, USA, |
|
− | |
other |
|
same data as previous except: on |
|
− | |
|
holotype | m | Puu O Umi NAR, Kohala, Hawaii, USA, ex |
|
− |
paratype |
|
same data as holotype | |||
paratypes | same data as holotype except: ex |
|
|||
other |
|
Donkey Mill Rd., Kona, Hawaii, USA, ex |
|
− | |
|
Olaa Forest, Hawaii, USA, ex |
|
− | ||
other |
|
Puu Pili, 3900ft, Kohala, Hawaii, USA, on |
− | − | |
other |
|
Kau, Hawaii, USA, ex |
|
||
other | Alili Spring trail, Kau, Hawaii, USA, ex |
|
|||
other | Humuula trail, Hawaii, USA, ex |
|
|||
other | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|||
other |
|
same data as previous except: ex |
|
||
other |
|
same data as previous except: “Hi12-14” D. Percy leg. ( |
|
||
|
other | Puu Makaala, Hawaii, USA, ex |
|
||
other | same data as previous except: “Hi16-14” D. Percy leg. ( |
|
|||
other |
|
same data as previous except: “Hi17-14” D. Percy leg. ( |
|
||
|
holotype | m | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|
paratypes | same data as holotype | − | |||
|
− | − | |||
|
holotype | m | Kilauea, Hawaii, USA, ex Ohia Lehua, |
− | − |
other | Southern Belt road (93 mile marker to Hilo), Hawaii, USA, |
|
− | ||
other |
|
Saddle Road (near 22 mile marker), Hawaii, USA, ex |
|
− | |
other | Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
|||
other | Olaa Forest near Solid Waste Dump, Hawaii, USA, ex |
|
− | ||
other |
|
same data as previous except: |
|
/ |
|
other | Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
|||
other | Kilauea Iki Crater, Hawaii, USA, ex |
|
|||
other | Puu Oo Trail, off Saddle Road, Hawaii, USA, ex |
|
− | ||
other | Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
|||
|
other | Kau, Hawaii, USA, ex |
|
||
other |
|
Tree Planting Road (1850-1880 flows), off Saddle Road, Hawaii, USA, ex |
|
− | |
other | Kipuka off Saddle Road, Hawaii, USA, ex |
|
|||
other | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|||
other | same data as previous except: ex |
|
|||
other | same data as previous except: ex |
|
|||
other | same data as previous except: “Hi12-14” D. Percy leg. ( |
|
|||
other | Puu Makaala, Hawaii, USA, ex |
|
− | ||
other |
|
Kona Hema TNC, Hawaii, USA, ex |
|
||
other |
|
Donkey Mill Rd., Kona, Hawaii, USA, ex |
|
− | |
|
holotype | m | Olinda Flume Road, Makawao, East Maui, USA, ex |
|
|
paratypes | same data as holotype | ||||
paratypes |
|
same data as previous except: “Hi52-14” D. Percy leg. ( |
|
||
paratype |
|
same data as previous except: “Hi41-14” D. Percy leg. ( |
|
||
|
|||||
|
holotype | f | Halemanu, Kauai, Sandwich Is., USA, 1933-323, 4000 ft, R. Perkins leg. ( |
− | − |
syntype | f | Kokee, Kauai, USA, on |
− | − | |
|
other | Discovery Center, Kokee State Park, Kauai, USA, ex |
|
− | |
other | Kalalau Valley (close to 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− | ||
other | Kokee State Park (near NASA Geophysical Observatory, ~14 mile marker), Kauai, USA, ex |
|
|||
other |
|
Kokee State Park, Kauai, USA, ex |
|
− | |
other |
|
Alakai Swamp trail, Kokee State Park, Kauai, USA, on |
|
||
|
holotype | m | Kalalau Valley (near 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− |
paratypes | same data as holotype | − | |||
other |
|
Kokee State Park (Puu o Kilo, at Kahuamma Flat), Kauai, USA, ex |
|
||
other |
|
Alakai Swamp trail, Kokee State Park, Kauai, USA, ex |
|
||
other |
|
Alakai Swamp trail (margin of 1st bog, Lehua Maka Noe), Kokee State Park, Kauai, USA, ex |
|
||
other |
|
same data as previous except: “Hi09-05” D. Percy leg. ( |
|
||
other |
|
Puu Ka Pele Forest Reserve, Hikimoe valley, Kauai, USA, ex |
|
||
other |
|
Awaawapuhi trail, Kokee State Park, Kauai, USA, ex |
|
− | |
|
holotype | m | Kalalau Valley (between 1st and 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− |
paratypes | same data as holotype | ||||
|
holotype | m | Kalalau Valley (between 1st and 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− |
|
paratype |
|
same data as holotype | − | |
|
holotype | m | Kalalau Valley (between 1st and 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− |
paratypes | same data as holotype | ||||
other | Discovery Center, Kokee State Park, Kauai, USA, ex |
|
− | ||
other |
|
Kalalau Valley (close to 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− | |
other | Kalalau Valley (near 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− | ||
other | Kokee State Park (near NASA Geophysical Observatory, ~14 mile marker), Kauai, USA, ex |
|
|||
other |
|
Kokee State Park (Puu o Kilo, at Kahuamma Flat), Kauai, USA, ex |
|
||
other |
|
Alakai Swamp trail (margin of 1st bog, Lehua Maka Noe), Kokee State Park, Kauai, USA, ex |
|
||
other |
|
Puu Ka Pele Forest Reserve, Hikimoe valley, Kauai, USA, ex |
|
− | |
|
holotype | m | Alakai Swamp trail, 1st bog (Lehua Maka Noe), Kauai, USA, ex |
|
− |
paratypes | same data as holotype | ||||
|
holotype | m? | Nualolo, Kauai, USA, ex Ohia Lehua, |
− | − |
other |
|
same data as holotype | − | ||
other | Alakai Swamp trail, 1st bog (Lehua Maka Noe), Kauai, USA, ex |
|
− | ||
|
holotype | f | Kalalau Valley (near 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− |
|
holotype | f | Kalalau Valley (between 1st and 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
− |
|
holotype | m | Alakai Swamp trail (margin of 1st bog, Lehua Maka Noe), Kokee State Park, Kauai, USA, ex |
|
− |
|
|||||
|
holotype | m | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− |
paratypes | same data as holotype | ||||
other |
|
Kuliouou Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− | |
other | Palolo Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: ex |
|
− | ||
other | Honouliuli Preserve, Waianae Mnts, Oahu, USA, ex |
|
− | ||
other | Pupukea, Koolau Mnts, Oahu, USA, ex |
|
|||
other |
|
Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
||
other |
|
Pahole NAR, Waianaea Mnts, Oahu, USA, ex |
|
− | |
other |
|
same data as previous except: on |
|
− | |
other | same data as previous except: on |
|
− | ||
other | same data as previous except: ex |
|
|||
other | Wiliwilinui Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
|
other | same data as previous except: ex |
|
− | |
other | Mnt Kaala summit, Waianae Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: ex |
|
− | ||
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
other |
|
Mnt Kaala, Waianae Mnts, Oahu, USA, ex |
− | ||
other | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|||
other | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|||
other |
|
Mnt Kaala road, Waianae Mnts, Oahu, USA, ex |
|
||
other | same data as previous except: “Hi64-14” D. Percy leg. ( |
|
|||
other | Palikea trail, Waianae Mnts, Oahu, USA, ex |
|
|||
other | Mokuleia Forest Reserve, Pahole NAR, Waianae Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: “Hi72-14” D. Percy leg. ( |
|
− | ||
other |
|
Pupukea, Koolau Mnts, Oahu, USA, ex |
|
− | |
other | same data as previous except: “Hi78-14” D. Percy leg. ( |
|
|||
other | Manoa Cliff trail to Pauoa Flats trail, Koolau Mnts, Oahu, USA, ex |
|
− | ||
|
other | same data as previous except: “Hi83-14” D. Percy leg. ( |
|
− | |
other | Mnt Kaala, Waianae Mnts, Oahu, USA, ex |
|
− | ||
other |
|
same data as previous except: “KM11-14” K. Magnacca leg. ( |
|
− | |
|
holotype | m | Mnt Kaala, 1600ft, Oahu, USA, ex |
− | − |
other |
|
Palolo Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− | |
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: “356-02” D. Percy leg. ( |
|
− | ||
other | Honouliuli Preserve, Waianae Mnts, Oahu, USA, ex |
|
− | ||
other | Pupukea, Koolau Mnts, Oahu, USA, ex |
|
|||
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− | ||
other | same data as previous except: ex |
|
− | ||
other | Pahole NAR, Waianaea Mnts, Oahu, USA, ex |
|
− | ||
other | Wiliwilinui Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
− | ||
other | Mnt Kaala road (culvert 55), Waianae Mnts, Oahu, USA, ex |
|
|||
other | Mnt Kaala summit, Waianae Mnts, Oahu, USA, ex |
|
|||
other | Aiea Ridge Trail, Koolau Mnts, Oahu, USA, ex |
|
|||
|
other |
|
Koolau Mnts, Oahu, USA, ex |
− | |
other | Mnt Kaala, Waianae Mnts, Oahu, USA, ex |
− | |||
other |
|
Palikoa, S. Waianae, Oahu, USA, ex |
− | ||
other |
|
Kuliouou, Koolau Mnts, Oahu, USA, ex |
|
||
other |
|
same data as previous except: ex |
|
||
other |
|
same data as previous except: ex |
|
||
other | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: “Hi59-14” D. Percy leg. ( |
|
|||
other | Mnt Kaala, boardwalk through bog, Waianae Mnts, Oahu, USA, ex |
|
|||
other |
|
Mnt Kaala summit, Waianae Mnts, Oahu, USA, ex |
|
||
other | Mnt Kaala road, Waianae Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: “Hi64-14” D. Percy leg. ( |
|
|||
other | Waianae Mnts, Oahu, USA, ex |
|
|||
other | Palikea trail, Waianae Mnts, Oahu, USA, ex |
|
|||
other | Mokuleia Forest Reserve, Pahole NAR, Waianae Mnts, Oahu, USA, ex |
|
|||
|
other | Pupukea, N. Koolau Mnts, Oahu, USA, ex |
|
||
other | Manoa Cliff trail to Pauoa Flats trail, S. Koolau Mnts, Oahu, USA, ex |
|
|||
other | same data as previous except: “Hi 84-14” D. Percy leg. ( |
|
|||
other |
|
Mnt Kaala, Waianae Mnts, Oahu, USA, ex |
|
− | |
other | same data as previous except: “KM09/11-14” K. Magnacca leg. ( |
|
− | ||
|
holotype |
|
3400ft, Lanai, USA, ex Ohia Lehua, |
− | − |
|
holotype | f | Kamiloloa, Molokai, USA, 3200 ft, on |
− | − |
other | Kamakou Preserve, Molokai, USA, on |
|
− | ||
other | same data as previous except: ex |
|
− | ||
other |
|
same data as previous except: on |
|
− | |
other | same data as previous except: ex |
|
− | ||
other | same data as previous except: ex |
|
|||
other |
|
same data as previous except: ex |
|
− | |
other | same data as previous except: ex |
|
|||
|
holotype | m | Kamakou Preserve, Molokai, USA, ex |
|
− |
paratypes |
|
same data as holotype | |||
|
other |
|
Kamakou Preserve, Molokai, USA, on |
|
− |
other | same data as previous except: ex |
|
|||
other | same data as previous except: on |
|
− | ||
other |
|
same data as previous except: ex |
|
− | |
|
holotype | m | Olinda Flume Road, Makawao, East Maui, USA, ex |
|
− |
paratypes | same data as holotype | ||||
other | Puu Kukui, boardwalk trail, West Maui, USA, ex |
|
|||
other |
|
same data as previous except: “Hi49-14” D. Percy leg. ( |
|
||
other |
|
Olinda Flume Road, Makawao, East Maui, USA, ex |
|
||
other | same data as previous except: “Hi53-14” D. Percy leg. ( |
|
|||
|
holotype | m | Puu Kaeo, Honolua Valley, West Maui, USA, ex |
|
− |
paratypes | same data as holotype |
|
|||
other | Puu Kukui, boardwalk trail, West Maui, USA, ex |
|
|||
other |
|
same data as previous except: “Hi48-14” D. Percy leg. ( |
|
||
other |
|
same data as previous except: “Hi49-14” D. Percy leg. ( |
|
− | |
other |
|
Olinda Flume Road, Makawao, East Maui, USA, ex |
|
||
|
other | same data as previous except: “Hi53-14” D. Percy leg. ( |
|
||
|
holotype | m | Puu Kaeo, Honolua Valley, West Maui, USA, ex |
|
− |
paratypes | same data as holotype | ||||
other |
|
Olinda Flume Road, Makawao, East Maui, USA, ex |
|
||
other |
|
Puu Kukui, boardwalk trail, West Maui, USA, ex |
|
||
other | same data as previous except: “Hi46-14” D. Percy leg. ( |
|
− | ||
other | same data as previous except: “Hi47-14” D. Percy leg. ( |
|
|||
other |
|
same data as previous except: “Hi48-14” D. Percy leg. ( |
|
||
other | same data as previous except: “Hi49-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi51-14” D. Percy leg. ( |
|
|||
other | Olinda Flume Road, Makawao, East Maui, USA, ex |
|
|||
other | same data as previous except: “Hi53-14” D. Percy leg. ( |
|
|||
|
holotype | m (?) | Kilauea, near Volcano, 4000ft, Hawaii, USA, |
− | − |
other |
|
Kilauea, Hawaii, USA, ex |
− | − | |
other | Southern Belt road (93-95 mile marker to Hilo), Hawaii, USA, |
|
|||
|
other | Southern Belt road (91 mile marker to Hilo), Hawaii, USA, |
|
||
other |
|
Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− | |
other | Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
− | ||
other |
|
Southern Belt road (34 mile marker to Hilo), Hawaii, USA, |
|
− | |
other | 1868 lava flow, near Kahuku Ranch, SW Hawaii, USA, ex |
|
− | ||
other |
|
Olaa Forest, Hawaii, USA, ex |
|
− | |
other |
|
Kipuka Alani, Hawaii, USA, on |
|
− | |
other |
|
Kipuka Ki, Hawaii, USA, on |
|
− | |
other | Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− | ||
other |
|
Olaa Forest, Hawaii, USA, ex |
|
− | |
other |
|
Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
− | |
other | Kehena, 3200ft, Kohala, Hawaii, USA, on |
− | − | ||
other | Puu Oo Trail, off Saddle Road, Hawaii, USA, ex |
|
|||
other | Puu O Umi NAR, Kohala, Hawaii, USA, ex |
|
|||
other | Waiakamali Gulch, Kohala, Hawaii, USA, ex |
|
|||
|
other |
|
RIG Site (E. Stacy), off Saddle Road, Hawaii, USA, ex |
|
|
other |
|
Kau, Hawaii, USA, ex |
|
− | |
other |
|
Gulch below Puu O Umi, Kohala, Hawaii, USA, ex |
|
− | |
other |
|
Olaa Forest (small unit off Wright Rd), Hawaii, USA, ex |
|
− | |
other |
|
Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
||
other | same data as previous except: “Hi13-14” D. Percy leg. ( |
|
|||
other | Kona Hema, Hawaii, USA, ex |
|
|||
other |
|
same data as previous except: “Hi25-14” D. Percy leg. ( |
|
− | |
other |
|
same data as previous except: “Hi26-14” D. Percy leg. ( |
|
||
other | same data as previous except: |
|
− | ||
other | same data as previous except: “Hi33-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi34-14” D. Percy leg. ( |
|
|||
other | Wright Rd (common garden), Volcano, Hawaii, USA, ex |
|
− | ||
|
holotype | m | Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− |
paratypes | same data as holotype | − | |||
other | Southern Belt road (93 mile marker to Hilo), Hawaii, USA, |
|
− | ||
|
other | Southern Belt road (91 mile marker to Hilo), Hawaii, USA, |
|
||
other | Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− | ||
other | Southern Belt road (near entrance to HVNP), Hawaii, USA, ex |
|
− | ||
other |
|
Southern Belt road (34 mile marker to Hilo), Hawaii, USA, |
|
− | |
other | 1868 lava flow, near Kahuku Ranch, SW Hawaii, USA, ex |
|
− | ||
other | Olaa Forest, Hawaii, USA, ex |
|
− | ||
other | Kilauea Crater area, HVNP, Hawaii, USA, ex |
|
− | ||
other |
|
Kipuka Alani, Hawaii, USA, on |
|
− | |
other | Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− | ||
other | Olaa Forest, Hawaii, USA, ex |
|
− | ||
other |
|
same data as previous except: |
|
− | |
other | Manuka NAR, Hawaii, USA, ex |
|
− | ||
other | Kipahoehoe NAR, Hawaii, USA, ex |
|
− | ||
other | same data as previous except: “Hi42-03” D. Percy leg. ( |
|
− | ||
other | Olaa Forest, Hawaii, USA, ex |
|
|||
|
other | Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
− | |
other |
|
Kehena, 2800ft, Kohala, Hawaii, USA, on |
− | − | |
other |
|
Puu Oo Trail, off Saddle Road, Hawaii, USA, ex |
|
||
other |
|
East of Waimea (off Hwy 19), Kohala, Hawaii, USA, ex |
|
||
other | Waiakamali Gulch, Kohala, Hawaii, USA, on |
|
− | ||
other | Puu O Umi NAR, Kohala, Hawaii, USA, ex |
|
|||
other |
|
same data as previous except: ex |
|
− | |
other | Waiakamali Gulch, Kohala, Hawaii, USA, ex |
|
|||
other | Tree Planting Road, off Saddle Road, Hawaii, USA, ex |
|
|||
other | RIG Site (E. Stacy), off Saddle Road, Hawaii, USA, ex |
|
|||
other | Hamakua Coast (21 mile marker), Hawaii, USA, ex |
|
|||
other |
|
Laupahoehoe, Hawaii, USA, |
|
||
other | Tree Planting Road (1850-1880 flows), off Saddle Road, Hawaii, USA, ex |
|
− | ||
other | Puu Lae Lae, Kohala, Hawaii, USA, ex |
|
|||
other |
|
Kohala Forest Preserve, Hawaii, USA, on |
|
− | |
|
other |
|
Gulch below Puu O Umi, Kohala, Hawaii, USA, ex |
|
|
other | Olaa Forest (small unit off Wright Rd), Hawaii, USA, ex |
|
|||
other |
|
Alili Spring trail, Kau, Hawaii, USA, ex |
|
||
other | Humuula trail, Hawaii, USA, ex |
|
|||
other | Kipuka off Saddle Road, Hawaii, USA, ex |
|
|||
other | Upper Hamakua Ditch, Kohala, Hawaii, USA, ex |
|
|||
other |
|
same data as previous except: “Hi09-14” D. Percy leg. ( |
|
||
other |
|
same data as previous except: ex |
|
||
other | same data as previous except: “Hi12-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi13-14” D. Percy leg. ( |
|
|||
other |
|
Puu Makaala, Hawaii, USA, ex |
|
||
other | same data as previous except: “Hi17-14” D. Percy leg. ( |
|
|||
other | Kona Hema, Hawaii, USA, ex |
|
|||
other | same data as previous except: “Hi22-14” D. Percy leg. ( |
|
− | ||
other | same data as previous except: “Hi23-14” D. Percy leg. ( |
|
|||
|
other | same data as previous except: “Hi24-14” D. Percy leg. ( |
|
||
other | same data as previous except: “Hi25-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi26-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi27-14” D. Percy leg. ( |
|
− | ||
other | same data as previous except: |
|
− | ||
other | same data as previous except: “Hi32-14” D. Percy leg. ( |
|
− | ||
other | same data as previous except: “Hi33-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi34-14” D. Percy leg. ( |
|
− | ||
other | Honuaula FR, Makaula Ooma Tract, Hualalai, Hawaii, USA, ex |
|
|||
other |
|
same data as previous except: “Hi29-14” D. Percy leg. ( |
|
||
other | Koloko Drive, Honuaula, Hualalai, Hawaii, USA, ex |
|
|||
other | Wright Rd (common garden), Volcano, Hawaii, USA, ex |
|
− | ||
other |
|
Tree Planting Rd, off Saddle Rd, Hawaii, USA, ex |
|
||
|
holotype | m | Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− |
paratypes | same data as holotype | − | |||
other | Southern Belt road (95 mile marker to Hilo), Hawaii, USA, |
|
|||
|
other | Southern Belt road (91 mile marker to Hilo), Hawaii, USA, |
|
||
other |
|
Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
||
other | Southern Belt road (near entrance to HVNP), Hawaii, USA, ex |
|
− | ||
other |
|
Southern Belt road (34 mile marker to Hilo), Hawaii, USA, |
|
− | |
other | 1868 lava flow, near Kahuku Ranch, SW Hawaii, USA, ex |
|
− | ||
other | Olaa Forest, Hawaii, USA, ex |
|
− | ||
other | Mountain View, Hawaii, USA, |
|
− | ||
other |
|
Puuwaawaa, Hawaii, USA, ex |
|
− | |
other |
|
Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
||
other |
|
same data as previous except: “Hi442-02” D. Percy leg. ( |
|
||
other | Kilauea Crater area, HVNP, Hawaii, USA, ex |
|
− | ||
other | Kipuka Puaulu (Bird Park), Hawaii, USA, ex |
|
− | ||
other | Olaa Forest, Hawaii, USA, ex |
|
− | ||
other | Manuka NAR, Hawaii, USA, ex |
|
− | ||
other | Kipahoehoe NAR, Hawaii, USA, ex |
|
− | ||
|
other | Olaa Forest, Hawaii, USA, ex |
|
− | |
other |
|
Kehena, 3150ft, Kohala, Hawaii, USA, on |
− | − | |
other |
|
Kilauea Iki Crater, Hawaii, USA, ex |
|
||
other | East of Waimea (off Hwy 19), Kohala, Hawaii, USA, ex |
|
|||
other |
|
Keauohana FR, Hawaii, USA, ex |
|
||
other |
|
KMC military camp, HVNP, Hawaii, USA, |
|
||
other |
|
Kohala Forest Preserve, Hawaii, USA, ex |
|
||
other |
|
Humuula trail, Hawaii, USA, ex |
|
||
other |
|
Kona Hema, Hawaii, USA, ex |
|
||
other | same data as previous except: “Hi26-14” D. Percy leg. ( |
|
|||
other | same data as previous except: “Hi27-14” D. Percy leg. ( |
|
− | ||
other | same data as previous except: |
|
|||
other | same data as previous except: “Hi34-14” D. Percy leg. ( |
|
− | ||
other | Honuaula FR, Makaula Ooma Tract, Hualalai, Hawaii, USA, ex |
|
|||
other | same data as previous except: “Hi29-14” D. Percy leg. ( |
|
|||
other | Wright Rd (common garden), Volcano, Hawaii, USA, ex |
|
− |
Outgroup material examined with GenBank numbers for cytochrome oxidase one (
Family |
n | Collection information | lat, long |
|
|
---|---|---|---|---|---|
|
Motu Mautaro, Tubuai, Austral Islands, French Polynesia, ex |
|
|
|
|
|
north of airport, Moorea, Society Islands, French Polynesia, ex |
|
|
|
|
|
|
below Ati Ati Peak, Moorea, Society Islands, French Polynesia, ex |
|
|
|
|
coastal road, western Raiatea, Society Islands, French Polynesia, ex |
|
|
|
|
|
Pouanlotch River, ca. 20km N of Voh, New Caledonia, ex |
-20°51’29”S, 164°36’44”E |
|
|
|
|
|
Airport, Norfolk Island, Australia, ex |
|
|
|
|
Berlayer Creek, Labrador Nature Reserve, Singapore, ex |
|
|
|
|
|
Rapa Island, French Polynesia, ex |
− |
|
|
|
− | Coahuila, Mexico [DNAs sample supplied by Trumble Lab, University of California, Riverside] | − |
|
|
|
|
− | Orange Co., California, USA [DNAs sample supplied by Trumble Lab, University of California, Riverside] | − |
|
|
Bladen Lakes State Forest, North Carolina, USA, ex |
− |
|
|
||
|
|
Louisiana, USA, ex |
− |
|
|
Nualolo Trail, Kokee State Park, Kauai, USA, ex |
|
|
|
||
Nualolo Trail, Kokee State Park, Kauai, USA, ex |
|
|
|
||
Mnt Kaala, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
Mnt Kaala, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
TNC Honouliuli Preserve, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
Pahole NAR, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
Pahole NAR, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
|
Motu Motihia, Tubuai, Austral Islands, French Polynesia, |
|
|
|
|
|
Forêt Sèche, Parc Forestier, Noumea, New Caledonia, ex |
|
|
|
|
|
Singapore Botanical Garden, Singapore, ex |
|
|
|
|
|
National Cheng Kung University campus, Tainan, Taiwan, ex |
|
|
|
|
Kalalau Valley (close to 2nd lookout), Kokee State Park, Kauai, USA, ex |
|
|
|
||
Pohakuloa Training Area, Hawaii, USA, ex |
|
|
|
||
Burnt Pine, Norfolk Island, Australia, ex |
|
|
|
||
Adelaide Hills, South Australia, Australia, reared from galls ex |
− |
|
|
||
Adelaide Hills, South Australia, Australia, reared from galls ex |
− |
|
|
||
Adelaide Hills, South Australia, Australia, reared from galls ex |
− |
|
|
||
South Mohiakea, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
|
Mnt Kaala road (culvert 32), Waianae Mnts, Oahu, USA, ex |
− |
|
|
|
Mokuleia Forest Reserve, Pahole NAR, Waianae Mnts, Oahu, USA, ex |
|
|
|
||
|
Puu Hapapa, Waianae Mnts, Oahu, USA, ex |
|
|
|
|
Auckland, New Zealand, ex |
− |
|
|
||
Mt. Ootua, Hiva Oa, Marquesas, French Polynesia, ex |
− |
|
|
||
Dur-West Farm, Sumpango, Sacatepequez, Guatemala, ex |
|
|
|
||
|
|
Las Cuevas, Chiquibul Forest Reserve, Cayo District, Belize, ex |
− |
|
|
|
Santa Cruz, California, USA, ex |
− |
|
|
|
|
Santa Monica hills, Los Angeles, California, USA, |
|
|
|
|
|
|
Queensland, Australia, ex |
− |
|
|
|
|
University of California, Berkeley, California, USA, ex |
|
|
|
Kenting National Park, Pingtung, Taiwan, ex |
|
|
|
||
Jonathon Dickson State Park, Florida, USA, ex |
− |
|
|
||
Rafting Ground Reserve, Queensland, Australia, ex |
-27°31’17”S, 152°55’30”E |
|
|
||
Pingtung, Taiwan, ex |
|
|
|
||
Pingtung, Taiwan, ex |
|
|
|
||
|
Rafting Ground Reserve, Queensland, Australia, ex |
|
|
|
|
West of Murray Bridge, South Australia, Australia, ex |
− |
|
|
||
Cawdor, Scotland, ex |
− |
|
|
||
|
Bookham Common, Surrey, UK, on |
|
|
|
|
Cradle Mnt, Tasmania, Australia, ex |
− |
|
|
||
West of Murray Bridge, South Australia, Australia, ex |
− |
|
|
||
Kew, London, UK, ex |
− |
|
|
||
|
Bookham Common, Surrey, UK, ex |
|
|
|
|
|
Mouaputa cascade, Vaioro River, Moorea, Society Islands, French Polynesia, ex |
|
|
|
|
ridge between Tonarutu and Tavaetu, Tubuai, Austral Islands, French Polynesia, ex |
|
|
|
||
22i | Pingtung, Taiwan, dissected from galls ex |
|
|
|
The molecular analysis includes 537 individuals, 479 in
To interpret the putative ancestral biology (i.e. galling versus non-galling) for
Ethanol preserved material was cleared in 10% potassium hydroxide followed by clove oil and slide mounted in Canada balsam as described in Natural History Museum, London, UK Bishop Museum, Honolulu, USA
As there is a large degree of size variation in some
Data resources. The collections and specimen data underpinning the analyses reported in this paper are deposited in the Natural History Museum Data Portal as Diana Percy (2017) Dataset:
Adult
Group | Species | n |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
|
1.7–2.30 | 0.7–0.97 | 0.47–0.52 | 0.56–0.80 | 0.03–0.05 | 0.06–0.08 | 0.19 | 0.16 | 0.16 | 0.30–0.38 | 0.20–0.28 | 0.14–0.20 | 0.06–0.08 | 0.26 | |
|
1.61–2.21 | 0.67–0.91 | 0.46–0.55 | 0.62–0.73 | 0.03–0.05 | 0.05–0.06 | 0.14 | 0.12–0.13 | 0.12 | 0.28–0.34 | 0.18–0.26 | 0.13–0.15 | 0.06–0.07 | 0.26–0.28 | ||
|
1.70–2.38 | 0.76–1.12 | 0.48–0.55 | 0.61–0.82 | 0.03–0.06 | 0.06–0.08 | 0.17–0.20 | 0.16–0.18 | 0.16–0.20 | 0.34–0.42 | 0.23–0.30 | 0.15–0.22 | 0.07–0.09 | 0.30–0.34 | ||
|
1.85–1.94 | 0.77–0.82 | 0.48–0.52 | 0.58–0.61 | 0.03–0.05 | 0.10–0.11 | − | − | − | 0.42–0.45 | 0.30–0.33 | 0.11–0.13 | 0.08–0.09 | 0.27–0.28 | ||
|
2.67 | 1.12 | 0.58 | 0.94 | 0.08 | 0.11 | − | − | − | 0.62 | 0.54 | 0.18 | 0.10 | − | ||
|
1.61–2.11 | 0.58–0.89 | 0.43–0.49 | 0.64–0.76 | 0.04–0.07 | 0.12–0.15 | 0.18–0.21 | 0.17–0.19 | 0.16–0.18 | 0.55–0.58 | 0.37–0.50 | 0.18–0.20 | 0.08–0.10 | 0.28 | ||
|
1.67–2.24 | 0.68–0.95 | 0.47–0.52 | 0.61–0.73 | 0.03–0.05 | 0.05–0.06 | 0.16–0.17 | 0.13–0.14 | 0.17–0.19 | 0.34–0.36 | 0.23–0.26 | 0.17–0.18 | 0.06–0.08 | 0.26 | ||
|
1.74–2.03 | 0.68–0.83 | 0.47–0.52 | 0.71–0.73 | 0.05–0.08 | 0.11–0.12 | 0.19 | 0.17 | 0.18 | 0.51–0.55 | 0.40–0.42 | 0.15–0.16 | 0.09 | 0.22 | ||
|
1.89–2.51 | 0.71–0.94 | 0.52–0.55 | 0.74–0.76 | 0.03–0.06 | 0.05–0.08 | 0.16 | 0.14 | 0.13 | 0.36–0.37 | 0.25 | 0.18–0.19 | 0.11 | 0.30 | ||
|
1.24–1.92 | 0.53–0.80 | 0.46–0.58 | 0.50–0.59 | 0.03–0.07 | 0.06–0.08 | 0.14–0.17 | 0.12–0.14 | 0.12–0.17 | 0.31–0.42 | 0.26–0.34 | 0.14–0.18 | 0.06–0.10 | 0.20–0.26 | ||
|
2.03–2.65 | 0.83–1.03 | 0.47–0.55 | 0.53–0.73 | 0.05–0.08 | 0.06–0.08 | 0.17–0.20 | 0.16–0.20 | 0.14–0.16 | 0.32–0.37 | 0.36–0.40 | 0.12–0.14 | 0.08 | 0.24–0.28 | ||
|
2.30–2.58 | 0.82–0.97 | 0.44–0.48 | 0.85 | 0.05–0.08 | 0.06–0.08 | 0.22–0.23 | 0.19–0.20 | 0.16 | 0.39 | 0.44 | 0.13 | 0.08 | − | ||
|
|
1.42–1.98 | 0.62–0.88 | 0.41–0.50 | 0.45–0.59 | 0.05–0.07 | 0.07–0.11 | 0.14–0.17 | 0.14–0.17 | 0.14–0.17 | 0.38–0.45 | 0.28–0.39 | 0.14–0.18 | 0.06–0.07 | 0.19–0.23 | |
|
1.71–2.01 | 0.73–0.86 | 0.48–0.52 | 0.48–0.55 | 0.02–0.04 | 0.07–0.08 | 0.16 | 0.16 | 0.18 | 0.31–0.40 | 0.26–0.35 | 0.13–0.16 | 0.06–0.07 | − | ||
|
|
2.62–3.66 | 1.15–1.60 | 0.67–0.75 | 1.26–1.40 | 0.12–0.14 | 0.08–0.11 | 0.22–0.25 | 0.21–0.23 | 0.23–0.24 | 0.43–0.45 | 0.38–0.41 | 0.10–0.12 | 0.13–0.14 | 0.29 | |
|
1.96–2.93 | 0.88–1.28 | 0.60–0.68 | 0.95–1.20 | 0.10–0.13 | 0.09–0.11 | 0.25–0.26 | 0.25–0.27 | 0.25–0.26 | 0.47–0.48 | 0.39–0.44 | 0.10–0.12 | 0.09 | 0.25–0.27 | ||
|
2.73–3.60 | 1.17–1.53 | 0.69–0.78 | 1.17–1.45 | 0.12–0.15 | 0.09–0.11 | 0.26–0.29 | 0.25–0.27 | 0.20–0.25 | 0.46–0.52 | 0.51–0.55 | 0.11 | 0.09–0.12 | 0.29 | ||
|
2.95–4.08 | 1.28–1.78 | 0.74–0.79 | 1.40–1.55 | 0.13 | 0.10–0.12 | 0.31 | 0.27 | 0.27 | 0.85 | 0.84 | 0.10 | 0.11 | 0.34 | ||
|
1.61–2.24 | 0.54–0.90 | 0.45–0.59 | 0.60–0.81 | 0.05–0.08 | 0.09–0.12 | 0.21–0.24 | 0.19–0.21 | 0.18–0.23 | 0.46–0.55 | 0.37–0.47 | 0.15–0.19 | 0.08–0.09 | 0.18–0.24 | ||
|
1.62–2.18 | 0.64–0.85 | 0.45–0.55 | 0.64–0.76 | 0.06–0.08 | 0.08–0.11 | 0.20–0.22 | 0.18–0.20 | 0.18–0.20 | 0.46–0.48 | 0.42–0.45 | 0.10–0.16 | 0.08–0.09 | 0.24–0.26 | ||
|
1.94–2.09 | 0.79–0.82 | 0.50 | 0.73–0.89 | 0.08–0.09 | 0.08–0.09 | 0.21 | 0.19 | 0.18 | 0.50 | 0.43 | 0.14 | 0.08 | 0.25 | ||
|
2.15 | 0.89 | 0.58 | 0.82 | 0.08 | 0.12 | − | − | − | 0.73 | 0.62 | 0.18 | 0.09 | 0.26 | ||
|
2.38 | 1.00 | 0.55 | 0.75 | 0.05 | 0.09 | − | − | − | 0.50 | 0.41 | 0.19 | 0.08 | 0.24 | ||
|
1.70 | 0.67 | 0.44 | 0.55 | 0.08 | 0.06 | 0.15 | 0.14 | 0.15 | − | − | − | − | − | ||
|
|
2.38–3.55 | 0.94–1.52 | 0.62–0.73 | 1.16–1.46 | 0.14–0.27 | 0.08–0.14 | 0.15–0.28 | 0.22–0.30 | 0.20–0.27 | 0.60–0.98 | 0.49–0.68 | 0.08–0.13 | 0.08–0.10 | 0.23–0.28 | |
|
1.86–3.42 | 0.67–1.30 | 0.47–0.70 | 0.78–1.26 | 0.10–0.23 | 0.07–0.09 | 0.17–0.26 | 0.18–0.28 | 0.16–0.25 | 0.47–0.88 | 0.36–0.56 | 0.09–0.17 | 0.08–0.11 | 0.17–0.24 | ||
|
2.50–3.40 | 1.00–1.45 | 0.64–0.66 | 1.09–1.60 | 0.15–0.28 | 0.09–0.12 | 0.22–0.26 | 0.26–0.29 | 0.22–0.27 | 0.57–0.78 | 0.49–0.69 | 0.09–0.13 | 0.10–0.11 | 0.23–0.30 | ||
|
1.94–2.68 | 0.76–1.06 | 0.52–0.63 | 0.94–1.16 | 0.15–0.25 | 0.07–0.09 | 0.17–0.18 | 0.18–0.20 | 0.15–0.17 | 0.38–0.43 | 0.29–0.31 | 0.09–0.12 | 0.11–0.12 | 0.18–0.20 | ||
|
2.70–3.20 | 1.04–1.27 | 0.60–0.67 | 1.25–1.45 | 0.24–0.31 | 0.09–0.11 | 0.22–0.24 | 0.25–0.26 | 0.20–0.21 | 0.55–0.59 | 0.41–0.50 | 0.11–0.14 | 0.12–0.13 | 0.24–0.26 | ||
|
2.68–3.39 | 1.03–1.30 | 0.61–0.71 | 1.18–1.48 | 0.20–0.32 | 0.09–0.12 | 0.25–0.26 | 0.28–0.32 | 0.25–0.27 | 0.62–0.75 | 0.55–0.68 | 0.14–0.19 | 0.10–0.11 | 0.25 | ||
|
2.08–2.88 | 0.80–1.15 | 0.53–0.65 | 0.90–1.21 | 0.08–0.13 | 0.08–0.09 | 0.20–0.23 | 0.19–0.23 | 0.18–0.22 | 0.55–0.70 | 0.41–0.50 | 0.10–0.15 | 0.09–0.11 | 0.20–0.22 | ||
|
2.61–3.73 | 1.03–1.50 | 0.53–0.78 | 1.21–1.50 | 0.18–0.27 | 0.10–0.12 | 0.23–0.31 | 0.24–0.28 | 0.21–0.25 | 0.54–0.65 | 0.43–0.63 | 0.10–0.17 | 0.10–0.12 | 0.26–0.28 | ||
|
1.70–2.98 | 0.67–1.15 | 0.43–0.65 | 0.76–1.23 | 0.11–0.22 | 0.07–0.11 | 0.17–0.25 | 0.18–0.24 | 0.16–0.22 | 0.37–0.54 | 0.31–0.44 | 0.09–0.15 | 0.10–0.12 | 0.19–0.22 | ||
|
1.83–2.93 | 0.73–1.18 | 0.43–0.65 | 0.93–1.20 | 0.09–0.14 | 0.08–0.11 | 0.22–0.24 | 0.24–0.30 | 0.21–0.25 | 0.55–0.73 | 0.49–0.63 | 0.10–0.13 | 0.09 | 0.23–0.24 |
Adult
Group | species |
|
|
||||||
---|---|---|---|---|---|---|---|---|---|
|
|
2.38–2.50 | 0.96–1.19 | 0.86–0.93 | 1.82–1.89 | 3.61–6.00 | 0.68–0.78 | 1.19–1.56 | 0.96–1.11 |
|
2.35–2.54 | 1.04–1.30 | 0.78–1.07 | 1.78–2.00 | 3.33–7.00 | 0.56–0.75 | 1.28–1.48 | 0.94–1.13 | |
|
1.92–2.52 | 0.81–1.20 | 0.77–1.00 | 1.78–2.00 | 3.25–6.50 | 0.63–0.78 | 1.14–1.59 | 0.95–1.21 | |
|
2.37–2.40 | 1.00 | 0.79–1.00 | 1.60–1.73 | 4.33–6.50 | 0.65–0.71 | 1.12–1.19 | 1.06–1.10 | |
|
2.38 | 1.19 | 0.92 | 2.13 | 2.80 | 0.78 | 1.61 | 0.87 | |
|
2.36–3.05 | 0.95–1.15 | 0.79–0.95 | 1.72–1.82 | 2.95–4.67 | 0.69–0.82 | 1.47–1.58 | 0.86–0.94 | |
|
2.35–2.45 | 0.95–1.09 | 0.76–0.95 | 1.82–1.94 | 4.00–6.00 | 0.61–0.75 | 1.30–1.55 | 1.00–1.11 | |
|
2.24–2.56 | 1.00–1.12 | 0.78–0.96 | 1.81–1.88 | 2.20–4.33 | 0.58–0.76 | 1.37–1.40 | 0.89–1.01 | |
|
2.66–2.73 | 1.14–1.44 | 0.75–0.89 | 1.79 | 3.67–5.50 | 0.56–0.63 | 1.47 | 1.09–1.16 | |
|
2.22–2.54 | 0.86–1.17 | 0.83–1.00 | 1.70–1.95 | 3.25–8.00 | 0.61–0.84 | 0.94–1.10 | 1.16–1.45 | |
|
2.44–2.63 | 1.08–1.24 | 0.78–0.92 | 1.74–2.00 | 2.40–5.33 | 0.58–0.89 | 1.13–1.33 | 0.85–0.97 | |
|
2.66–2.81 | 1.43–1.68 | 0.43–0.88 | 1.82–1.93 | 2.20–2.50 | 0.59–0.73 | − | 0.78 | |
|
|
2.21–2.48 | 0.95–1.12 | 0.67–1.00 | 1.79–2.07 | 2.17–4.33 | 0.59–0.86 | 1.00–1.26 | 0.87–1.13 |
|
2.24–2.36 | 0.95–1.05 | 0.86–1.06 | 1.88–2.00 | 5.42–11.00 | 0.65–0.88 | 0.94–1.13 | 1.06–1.14 | |
|
|
2.28–2.40 | 1.08–1.19 | 0.71–0.83 | 1.71–1.87 | 1.88–2.20 | 0.58–0.70 | 1.70–2.00 | 0.94–1.10 |
|
2.15–2.30 | 1.10–1.27 | 0.75–0.92 | 1.73–1.86 | 1.80–2.25 | 0.60–0.71 | 1.58–1.85 | 0.84–0.98 | |
|
2.31–2.41 | 1.37–1.62 | 0.88–0.95 | 1.72–1.87 | 1.67–2.13 | 0.56–0.67 | 1.66–2.07 | 0.85–0.89 | |
|
2.30–2.31 | 1.22–1.25 | 0.72–0.79 | 1.70–1.79 | 1.90–2.20 | 0.58–0.59 | 1.90–1.97 | 0.83–0.89 | |
|
2.44–3.21 | 1.00–1.26 | 0.78–0.95 | 1.82–2.05 | 2.33–4.00 | 0.57–0.78 | 1.20–1.63 | 0.95–1.16 | |
|
2.47–2.71 | 1.11–1.29 | 0.79–1.13 | 1.72–1.94 | 2.20–3.64 | 0.67–0.80 | 1.22–1.61 | 1.18–1.45 | |
|
2.37–2.65 | 1.32–1.33 | 0.83–0.85 | 1.74 | 2.00–2.60 | 0.63–0.68 | 1.45–1.79 | 1.14–1.32 | |
|
2.41 | 1.09 | 0.96 | 1.90 | 3.00 | 0.75 | 1.42 | 0.93 | |
|
2.38 | 1.26 | 0.88 | 2.00 | 3.50 | 0.64 | 1.36 | 0.94 | |
|
2.55 | 1.12 | 0.86 | 1.93 | 2.00 | 0.67 | 1.24 | 1.04 | |
|
|
2.28–2.79 | 0.97–1.29 | 0.73–1.27 | 1.68–1.93 | 1.13–2.12 | 0.60–0.86 | 1.79–2.25 | 0.80–1.00 |
|
2.48–3.41 | 1.08–1.57 | 0.55–1.04 | 1.71–2.08 | 1.07–2.25 | 0.58–0.83 | 1.63–1.88 | 0.86–1.24 | |
|
2.05–2.58 | 1.05–1.43 | 0.83–1.06 | 1.76–2.00 | 0.90–1.63 | 0.67–0.89 | 1.85–2.38 | 0.85–1.02 | |
|
2.45–2.56 | 1.10–1.28 | 0.81–1.09 | 1.81–2.00 | 1.05–1.50 | 0.72–0.90 | 1.73–1.94 | 0.96–1.17 | |
|
|
2.51–2.60 | 1.09–1.24 | 0.74–0.91 | 1.76–1.93 | 0.80–1.05 | 0.71–0.83 | 1.92–2.23 | 0.96–1.05 |
|
2.51–2.61 | 1.18–1.30 | 0.82–0.93 | 1.82–1.93 | 0.86–1.13 | 0.64–0.77 | 1.86–2.11 | 0.90–1.00 | |
|
2.47–2.66 | 1.12–1.52 | 0.74–1.00 | 1.80–2.09 | 1.90–2.86 | 0.73–0.86 | 1.71–2.07 | 0.95–1.11 | |
|
2.42–2.63 | 1.00–1.24 | 0.70–0.96 | 1.50–1.93 | 0.94–1.43 | 0.65–0.80 | 1.90–2.22 | 0.84–1.07 | |
|
2.40–2.65 | 0.96–1.36 | 0.77–0.95 | 1.67–2.00 | 1.04–2.00 | 0.64–0.92 | 1.63–1.95 | 0.96–1.19 | |
|
2.48–2.62 | 1.06–1.27 | 0.81–0.94 | 1.70–2.10 | 1.64–2.25 | 0.69–0.90 | 1.64–1.85 | 0.92–1.02 |
Adult
Group | Species | |||||||||
---|---|---|---|---|---|---|---|---|---|---|
|
|
0.31 | 1.20 | 1.00 | 2.00 | 0.95 | 0.63–0.72 | 1.88–2.53 | 1.34–1.48 | 1.88–2.29 |
|
0.26 | 1.16–1.17 | 0.97–1.03 | 1.88–2.21 | 0.75–0.78 | 0.54–0.67 | 1.84–2.24 | 1.28–1.70 | 2.23–3.05 | |
|
0.31–0.35 | 0.96–1.10 | 0.84–1.00 | 2.08–2.27 | 0.89–1.05 | 0.66–0.73 | 1.92–2.24 | 1.35–1.57 | 2.85 | |
|
− | − | − | − | − | 0.87 | 3.31–3.71 | 1.33–1.37 | 3.18–3.78 | |
|
− | − | − | − | − | 1.06 | 3.50 | 1.15 | − | |
|
0.37–0.40 | 1.04–1.21 | 0.95–1.12 | 2.28–2.63 | 0.91–1.05 | 1.12–1.19 | 2.82–3.09 | 1.16–1.50 | 2.12 | |
|
0.29 | 1.21–1.24 | 0.72–0.79 | 2.10–2.68 | 0.76–0.77 | 0.67–0.73 | 1.87–2.14 | 1.30–1.55 | 3.00 | |
|
0.36 | 1.14 | 0.91 | 2.30 | 0.84 | 1.00–1.06 | 3.37–3.45 | 1.27–1.33 | 2.33 | |
|
0.26 | 1.18 | 1.03 | 2.20 | 0.76 | 0.66–0.67 | 1.92–2.05 | 1.48 | 3.36 | |
|
0.25–0.27 | 1.12–1.38 | 0.78–0.97 | 2.35–3.57 | 0.69–0.76 | 0.63–0.74 | 2.05–2.82 | 1.11–1.48 | 2.78–3.57 | |
|
0.33–0.39 | 0.90–1.07 | 1.15–1.31 | 1.90–2.27 | 0.92–1.07 | 0.60–0.71 | 2.50–2.80 | 0.84–1.02 | 2.67–2.92 | |
|
0.44 | 1.13 | 1.17 | 2.41 | 1.00 | 0.83 | 3.06 | 0.89 | − | |
|
|
0.33–0.39 | 0.86–1.11 | 0.98–1.09 | 2.27–2.73 | 0.87–1.00 | 0.85–0.97 | 2.55–2.78 | 1.08–1.35 | 1.81–3.11 |
|
0.32 | 1.03 | 0.89 | 2.75 | 0.85 | 0.75–0.78 | 2.50–2.53 | 1.09–1.16 | − | |
|
|
0.31–0.34 | 0.97–1.15 | 0.87–1.00 | 2.10–2.31 | 0.60–0.73 | 0.59–0.60 | 3.83–4.50 | 1.06–1.18 | 2.16–2.40 |
|
0.41–0.42 | 0.96 | 1.02–1.04 | 1.95–2.07 | 1.00–1.04 | 0.70–0.74 | 4.00–4.67 | 1.09–1.23 | 1.91–2.07 | |
|
0.36–0.39 | 1.00–1.07 | 1.04–1.32 | 1.92–2.66 | 0.93–1.05 | 0.67–0.72 | 4.27–4.82 | 0.90–0.95 | 1.84–1.93 | |
|
0.37 | 1.14 | 1.00 | 2.14 | 0.93 | 1.08 | 8.70 | 1.01 | 2.14 | |
|
0.35–0.42 | 1.08–1.20 | 0.89–1.08 | 2.32–3.13 | 0.95–1.08 | 0.82–1.02 | 2.76–3.20 | 1.07–1.24 | 2.60–3.37 | |
|
0.38–0.40 | 1.09–1.22 | 0.94–1.02 | 2.33–2.67 | 1.05–1.05 | 0.85–0.97 | 3.33–4.62 | 1.04–1.15 | 2.13 | |
|
0.38 | 1.08 | 1.04 | 2.30 | 0.96–1.04 | 1.01 | 3.50 | 1.17 | 2.07 | |
|
− | − | − | − | − | 1.26 | 3.96 | 1.17 | 2.36 | |
|
− | − | − | − | − | 0.91 | 2.68 | 1.21 | 2.94 | |
|
0.31 | 1.12 | 0.92 | 2.18 | 0.77 | − | − | − | − | |
|
|
0.35–0.43 | 0.61–1.02 | 1.06–1.30 | 2.13–3.00 | 1.00–1.19 | 0.87–1.31 | 5.71–9.38 | 1.12–1.46 | 1.45–2.06 |
|
0.33–0.44 | 0.78–0.96 | 1.00–1.29 | 2.01–2.68 | 1.04–1.26 | 0.78–1.32 | 3.18–6.47 | 1.18–1.67 | 1.20–1.51 | |
|
0.42–0.44 | 0.79–0.93 | 1.04–1.20 | 1.94–2.37 | 0.92–1.07 | 0.95–1.12 | 5.27–7.50 | 1.14–1.17 | 1.66–1.80 | |
|
|
0.33–0.34 | 0.88–0.95 | 1.12–1.26 | 1.95–2.21 | 0.92–1.10 | 0.64–0.73 | 3.58–4.56 | 1.26–1.43 | 1.23–1.53 |
|
0.41–0.42 | 0.86–0.94 | 1.21–1.29 | 1.93–2.21 | 1.00–1.04 | 0.83–0.89 | 4.21–5.09 | 1.10–1.40 | 1.68–1.94 | |
|
0.46–0.50 | 0.76–0.94 | 1.13–1.24 | 2.07–2.32 | 1.03–1.14 | 0.98–1.06 | 3.92–4.59 | 1.11–1.13 | 1.91 | |
|
0.35–0.41 | 0.87–1.03 | 1.00–1.10 | 2.45–2.89 | 1.00–1.24 | 1.02–1.07 | 4.47–6.00 | 1.30–1.40 | 1.32–1.54 | |
|
0.36–0.52 | 0.88–1.10 | 1.09–1.26 | 2.01–2.32 | 0.94–1.17 | 0.83–0.93 | 3.72–5.46 | 0.97–1.31 | 1.83–1.92 | |
|
0.33–0.43 | 0.88–1.14 | 0.95–1.19 | 2.10–2.50 | 0.94–1.09 | 0.71–0.93 | 3.64–4.82 | 1.19–1.30 | 1.44–1.80 | |
|
0.47–0.55 | 0.76–0.92 | 1.09–1.25 | 1.96–2.45 | 1.04–1.15 | 0.95–1.17 | 5.15–7.30 | 1.12–1.23 | 1.76–1.94 |
Group | Species | n |
|
|
|
|
|
|
|
|
|||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
|
2 | 1.36–1.48 | 1.15 | 0.79 | 0.42–0.48 | 0.97–1.03 | 0.19–0.21 | 0.56–0.57 | 0.15–0.16 | 1.18–1.29 | 3.55–3.68 | 4.95–5.05 |
|
5 | 1.41–1.55 | 0.98–1.12 | 0.73–0.76 | 0.58–0.73 | 0.73–0.88 | 0.21–0.22 | 0.51–0.64 | 0.21–0.23 | 1.32–1.45 | 2.24–3.08 | 3.37–4.08 | |
|
2 | 1.36–1.70 | 0.82–1.00 | 0.61–0.76 | 0.55–0.65 | 0.70–0.85 | 0.15–0.20 | 0.50–0.56 | 0.16–0.18 | 1.67–1.70 | 3.15–3.18 | 4.24–4.58 | |
|
6 | 1.33–1.52 | 1.03–1.27 | 0.76–1.00 | 0.55–0.64 | 0.91–1.06 | 0.22–0.25 | 0.47–0.54 | 0.17–0.19 | 1.15–1.32 | 2.68–2.86 | 4.06–4.42 | |
|
5 | 1.30–1.48 | 1.03–1.15 | 0.76–0.88 | 0.50–0.55 | 0.88–0.97 | 0.21–0.24 | 0.46–0.50 | 0.18–0.21 | 1.26–1.35 | 2.31–2.59 | 3.85–4.22 | |
|
|
5 | 1.00–1.18 | 0.85–0.94 | 0.61–0.64 | 0.36–0.45 | 0.64–0.73 | 0.13–0.16 | 0.42–0.45 | 0.16–0.17 | 1.13–1.39 | 2.52–2.75 | 4.55–4.97 |
|
|
6 | 1.45–2.00 | 1.03–1.36 | 0.61–0.88 | 0.76–1.06 | 0.82–1.15 | 0.03–0.05 | 0.53–0.70 | 0.22–0.32 | 1.41–1.50 | 2.15–2.50 | 20.45–35.98 |
|
1 | 1.18 | 1.00 | 0.64 | 0.48 | 0.85 | 0.14 | 0.51 | 0.17 | 1.18 | 3.05 | 5.89 | |
|
5 | 1.15–1.18 | 0.79–0.88 | 0.61–0.64 | 0.45–0.45 | 0.70–0.76 | 0.14–0.16 | 0.44–0.49 | 0.16–0.18 | 1.34–1.46 | 2.39–2.95 | 4.73–5.26 | |
|
|
6 | 1.85–2.33 | 1.33–1.52 | 0.82–0.91 | 0.91–1.24 | 1.12–1.36 | 0.04–0.06 | 0.72–0.78 | 0.39–0.44 | 1.36–1.60 | 1.70–1.94 | 20.02–28.79 |
|
3 | 1.21–1.61 | 0.97–1.09 | 0.61–0.70 | 0.61–0.79 | 0.73–0.91 | 0.03–0.06 | 0.54–0.56 | 0.34–0.36 | 1.25–1.47 | 1.56–1.67 | 15.15–28.41 | |
|
1 | 1.82 | 1.18 | 0.82 | 0.76 | 1.03 | 0.06 | 0.70 | 0.39 | 1.54 | 1.80 | 18.40 | |
|
5 | 1.70–2.00 | 1.12–1.30 | 0.70–0.85 | 0.76–1.06 | 0.94–1.12 | 0.16–0.17 | 0.62–0.72 | 0.32–0.42 | 1.42–1.58 | 1.54–2.25 | 5.59–7.01 | |
|
2 | 2.33–2.61 | 1.55–1.73 | 0.94–1.06 | 0.97–1.09 | 1.09–1.36 | 0.06 | 0.80 | 0.46–0.48 | 1.51 | 1.67–1.75 | 19.48–21.30 | |
|
7 | 1.45–1.88 | 1.00–1.24 | 0.67–0.85 | 0.70–0.97 | 0.73–1.03 | 0.03–0.06 | 0.56–0.70 | 0.36–0.44 | 1.43–1.51 | 1.54–1.82 | 15.15–32.19 | |
|
6 | 1.45–2.12 | 1.06–1.39 | 0.73–0.91 | 0.64–1.06 | 0.76–1.06 | 0.05–0.06 | 0.58–0.78 | 0.32–0.42 | 1.37–1.56 | 1.76–1.92 | 11.84–22.09 |
Combined analysis of the two mitochondrial regions provides two robustly supported key results: a) confirmation of the monophyly of the genus
Results from outgroup sampling. Majority-rule consensus of maximum likelihood analysis (combined
Parsimony reconstruction analysis resolved the ancestral state for
Maximum inter-specific molecular divergence within
Based on morphological and molecular analyses, four species groups are recognized (Figs
Best maximum likelihood topology (-lnL = -33769.25, combined
The
Recognized morphological forms provide information about the extent and distribution of morphological variation within, in some cases, relatively broad species concepts, and a number of the more distinct forms may eventually require recognition at species level.
There are 12 species in the
There are two species in the
There are 10 species in the
There are 12 species in the
Notable morphological variation is evident for several species, which has resulted in the recognition, in this treatment, of a number of morphological forms. These forms are usually also identifiable in genotypic clusters, but in two widespread species on Oahu,
Neighbour-joining analysis (combined
Repeated patterns of intraspecific morphological variation in the
When considering the age of
The closest sister group of the Hawaiian
The question of long distance dispersal to the Hawaiian Islands, whether from another Pacific archipelago or from a continental source, remains an enduring enigma for many insect groups with no obvious means of trans-ocean dispersal.
Some aspects of
There still remain many questions regarding the underlying mechanisms responsible for morphological variation in
Examples of variability in
There are many examples suggesting a strong role for localized isolation by distance in
Complex patterns of diversification observed on the youngest island of Hawaii (e.g. “
A comparison of the patterns of local divergence in
The diversity in
An ancestral pit-galling habit may have acted as an evolutionary springboard to both other galling and non-galling habits, with potentially multiple shifts to closed galling or free-living biologies. Independent shifts to closed galling biologies in the
As mentioned above, stem/bud gallers have generally larger body size and paler body colour than leaf gallers in the
Because gall position is one of the factors determining gall structure, maternal selection of an oviposition site will be important in determining gall type and potentially promoting diversification (
Species diversity in
General body colour either entirely dark (black, brown, or red), entirely or mostly pale (cream, yellow or orange), or distinctly bicolored (pale/dark) (e.g. Fig.
Extremely variable immature morphology reflects variation in biologies and galling habits. Immatures may be free-living or gall forming (open and closed galls). Galling species appear to have one immature per gall/gall chamber, but in some galling species dense aggregations of galls result in clusters of chambers in close proximity. Morphologically consistent characters for 5th instars are antennae with 8-9 segments bearing 4 rhinaria (on segments 4, 6, 8 and 9, or 2 on segment 8) and two short to medium long terminal simple setae, tarsi with broad crescent arolia and each terminal tarsus bearing a long simple or weakly capitate seta, and anus situated ventrally.
The host plant of all
The original placement of some taxa in
The original placement of some of the
In this study, broad species concepts are combined with the recognition of morphological forms (infrasubspecific names as per ICZN) to convey the extent and distribution of morphological variation. Some forms recognized here may warrant subsequent recognition at species level.
1 | On Kauai |
|
– | On other islands |
|
2 | Generally smaller species (often bicolored cream and black, or entirely black, or entirely pale) with short genal processes (ratio |
|
– | Generally larger species (often red-brown or yellow-green), genal processes either long (ratio |
|
1 | Male with paramere shape triangular, broad at base and tapering to narrow apex, female subgenital plate truncate apically, ovipositor valvulae dorsalis strongly convex dorsally |
|
– | Male with paramere more or less parallel sided before constricting below apex, or tapering to apex, female subgenital plate not truncate either acute or bluntly acute, ovipositor valvulae dorsalis typically moderately or not convex dorsally, rarely strongly convex dorsally (e.g. |
|
2 | Fore wing narrow (ratio |
|
– | Fore wing broader (ratio |
|
3 | Male with longer paramere ( |
|
– | Male with shorter paramere ( |
|
4 | Male with longer distal aedeagus segment ( |
|
– | Male with shorter distal aedeagus segment ( |
|
5 | Male with paramere length less than 0.80 height of subgenital plate, shape of male and female terminalia as in Fig. |
|
– | Male with paramere length greater than 0.80 height of subgenital plate, shape of male and female terminalia as in Fig. |
|
6 | Female terminalia with relatively small anal ring (ratio |
|
– | Female terminalia with relatively large anal ring (ratio |
|
7 | Larger species ( |
|
– | Smaller species ( |
|
8 | Antenna longer (>0.70 mm), female proctiger longer ( |
|
– | Antenna shorter (<0.70 mm), female proctiger shorter ( |
|
9 | Length of female proctiger and subgenital plate subequal (ratio |
|
Female proctiger longer than subgenital plate (ratio |
|
|
10 | Fore wing narrower ( |
|
– | Fore wing broader ( |
|
11 | Long distal proboscis segment ( |
|
– | Shorter distal proboscis segment ( |
|
12 | Male with short paramere (ratios |
|
– | Male with longer paramere (ratios |
|
13 | Male with paramere apical hook inward pointing, longer distal aedeagus segment (ratio |
|
– | Male with paramere apical hook upward pointing, shorter distal aedeagus segment (ratio |
|
A group of mostly diminutive species, though the largest are similar in size to the smaller members of other groups (Fig.
Typically bicoloured, generally pale cream-yellow to green on thorax and abdomen, head darker, with a dark dorsal stripe from the head extending part or all the length of the body. Fore wing membrane clear or slightly fuscous.
Fore wing apex rounded; surface spinules dispersed, usually in all cells but may be reduced or absent in r1 and c+sc; setae on margins and veins short to minute (Fig.
Unpigmented or light brown, broad and moderately long with uninterrupted striations over entire surface, short pedicel 1/4 length from base, tail moderately long (Fig.
(note: immature association for
Unconfirmed, but may prefer more glabrous morphotypes.
Molokai, Lanai, Hawaii.
Although treated broadly as occurring on three islands (based on previous records summarized in
The immatures described here are considered free-living but develop under a domed casing which remains after the adult has enclosed (see note on need to confirm adult-immature association).
Belongs to a complex of species within the
Holotype, female (slide mounted,
Typically bicoloured, generally pale cream-yellow to green on thorax and abdomen, head darker, with or without a dark dorsal stripe from the head extending part or all the length of the body (Fig.
Fore wing apex rounded to bluntly acute; surface spinules well dispersed, present in all cells but reduced coverage in r1 and c+sc; setae on margins and veins short to minute (Fig.
Unpigmented or light brown, long to moderately long with striations over entire surface, but in form
Unknown.
Morphotype preference unknown.
Molokai, Maui.
Of the three recognized forms, form
Unknown.
Named after Hina, a goddess of the moon in Hawaiian mythology (noun in the nominative singular standing in apposition to the generic name).
Three forms are recognized (Figs
Holotype male (slide mounted,
Variable, often strikingly bicoloured with black or dark brown head and pale cream or yellow-green thorax and abdomen, with or without a dark dorsal stripe from the head extending part or all the length of the body, but can also be completely pale throughout (Fig.
Fore wing apex rounded; surface spinules well dispersed in all cells but reduced or absent in r1 and c+sc; setae on margins and veins short to minute (Fig.
(only known for form
Unknown.
Unconfirmed, but may prefer more glabrous morphotypes.
Hawaii.
All three forms are found in Kohala, with form
Unknown.
Named after “wyvern”, a mythical winged creature in Medieval mythology, in reference to the rarity and acknowledged taxonomic puzzle this taxon presents (noun in the nominative singular).
Three forms are recognized (Fig.
Holotype male (slide mounted,
Typically bicoloured, generally pale cream-yellow thorax and abdomen, head darker and a dark dorsal stripe from the head extending part or all the length of the body (Fig.
Fore wing apex rounded; surface spinules dispersed in all cells but reduced coverage in r1 and c+sc; setae on margins and veins short to minute (Fig.
Unpigmented or light brown, narrow, slender, entire surface with narrowly spaced uninterrupted striations no visible pedicel, tail short (Fig.
Unknown.
Collected from semi-glabrous morphotype growing on lava flow (dated to 1880s) in south western Hawaii.
Hawaii.
Known from only one locality.
Unknown, but may be free-living as for
In reference to the dark dorsal stripe that is frequently present in this species and signifies affiliation with the
Currently known only from females; females appear particularly similar to
Holotype female (slide mounted,
General body colour yellow to brown. Head darker than the rest of the body, apparently not distinctly bicoloured (e.g. without distinct dorsal stripe). Fore wing membrane clear, or slightly fuscous.
Fore wing apex rounded; surface spinules dispersed in all cells but reduced or none in r1 and c+sc; setae on margins and veins short to minute (Fig.
Unknown.
Unconfirmed, but 1st instars recovered on the surface of leaves at the collection locality have a setal arrangement similar to that illustrated for
Collected from pubescent morphotypes.
Hawaii.
Only known from Kohala.
Unconfirmed, but this species was collected from low growing pubescent forms in upland bog; eggs and 1st instar immatures were recovered from the plant surface among the trichomes along the mid-rib (upper leaf surface) and petiole, these eggs have widely spaced interrupted surface striations, a short pedicel and a long tail, however, two other
Named after Kapo, a goddess of fertility in Hawaiian mythology (noun in the nominative singular standing in apposition to the generic name).
Currently known from only one female; this is the largest species in the
Holotype female (slide mounted,
Typically bicoloured, generally pale cream-yellow thorax and abdomen, head brown or black, apparently lacking dorsal stripe. Fore wing membrane slightly fuscous.
Fore wing apex rounded; surface spinules dispersed, usually in all cells except may be reduced or absent from cell r1 and c+sc; short setae on margins and veins (Fig.
Unpigmented, broad, surface covered with long uninterrupted striations, short pedicel positioned 1/4 length from base, tail moderately long (Fig.
Colour and structure 5th instar: Appearance is white and spikey (hedgehog-like) due to coverage of stiff white filaments produced from sectasetae (Fig.
On pubescent and tomentose morphotypes.
Hawaii.
Widespread on Hawaii: DNA analysis indicates distinct clusters of individuals from (a) Kohala as basal and sister to (b) Kau, (c) Saddle Road, and (d) Hualalai.
This species is free-living on the undersides of pubescent leaves.
Named for the distinctly longer distal proboscis segment (adjective in the nominative singular).
Holotype male (slide mounted,
Typically bicoloured, generally pale yellow to green on thorax and abdomen, head darker black or brown, and a dark dorsal stripe extending part or all the length of the body. Fore wing membrane slightly fuscous.
Fore wing apex rounded; surface spinules distributed in all cells except few or none in r1 and c+sc; short setae on margins and veins. Antennae short (av. length 0.67; ratio
Unpigmented, slender, and apparently without striations, pedicel or tail.
Unknown.
Collected from mixed glabrous and pubescent morphotypes.
Hawaii.
Only known from the Kohala region of Hawaii.
Unknown.
Named for Poliahu, a goddess of snow in Hawaiian mythology, in reference to a concept that each snowflake is unique, as many individuals sampled for this species have highly divergent genetic haplotypes (noun in the nominative singular standing in apposition to the generic name).
Holotype male (slide mounted,
Typically bicoloured, generally pale yellow to green thorax and abdomen, head darker black or brown, and a dark dorsal stripe extending from head down the thorax. Fore wing membrane slightly fuscous.
Fore wing apex bluntly acute; surface spinules dispersed, usually in all cells, but may be limited or absent from cells c+sc and r1; short setae on margins and veins (Fig.
Unpigmented to light brown, short, broad and with striations, mostly uninterrupted, on the dorsal surface, pedicel and tail short (Fig.
Unknown.
Collected from glabrous morphotype.
Molokai.
Known from only one location in Kamakou Preserve.
Unknown.
Named for Lona, a lunar deity in Hawaiian mythology (noun in the nominative singular standing in apposition to the generic name).
Holotype male (slide mounted,
Typically bicoloured, generally pale cream-yellow to green thorax and abdomen, head darker, with a dark dorsal stripe from the head extending part or all the length of the body. Fore wing membrane fuscous.
Fore wing moderately narrow, apex bluntly acute; surface spinules distributed in all cells, but limited in c+sc; short setae on margins and veins (Fig.
Light brown, extremely long, slender and with striations, mostly uninterrupted, over entire surface, no pedicel apparent, tail moderately long (Fig.
Unknown.
Collected from glabrous morphotypes.
Oahu.
Only known from one locality, the high elevation bog area on Mnt Kaala.
Unknown.
Named for Kuini Liliha, a High Chiefess who served the Kingdom of Hawaii as royal governor of Oahu (noun in the nominative singular standing in apposition to the generic name).
Holotype male (slide mounted,
General body colour typically dark brown to black throughout, but occasionally partly or entirely pale cream or yellow throughout (Fig.
Fore wing apex rounded; surface spinules densely distributed throughout all cells; short to minute setae on margins and veins (Fig.
Unpigmented, elongate, slender, slightly sinusoidal, dorsal surface with widely spaced interrupted striations, medium-short pedicel positioned 1/4-1/3 length from base, tail long (Fig.
Colour and structure: Orange or cream. 5th instar: Narrowly ovoid in outline with wing buds protruding and with distinct humeral lobes (Fig.
Predominantly on pubescent and tomentose morphotypes.
Oahu, Molokai, Maui.
A common species on Oahu. Four clusters can be recognized in the DNA analysis: (a) individuals from Molokai and Maui; these in turn group with (b) a population from Oahu’s southern Koolau Mnts that have more developed genal processes (Fig.
The immatures are free-living, usually on the lower leaf surface of pubescent and tomentose morphotypes, this host morphotype preference is also noted on slide specimens collected in 1973 by Beardsley (BISH). Eggs appear to be laid mostly singly and sparsely distributed amongst the leaf trichomes.
One of the most commonly encountered species in the
Holotype female (dry mounted,
Variable, usually bicoloured with orange-brown, cream or yellow to greenish-yellow thorax and abdomen, and a dark dorsal stripe from the head extending part or all the length of the body. Fore wing fuscous, especially around anal margin.
Fore wing apex bluntly acute to rounded; dispersed spinules present in all cells, but reduced or absent in cell r1; setae on margins and veins short to minute (Fig.
Unpigmented, narrowly oval, marginally sinusoidal, entire surface with interrupted striations, medium-short pedicel positioned 1/4 length from base, tail short (Fig.
Colour and structure: Pale cream, yellow to green. 5th instar: Broadly ovoid and ventro-dorsally flattened wing buds only slightly protruding and distinct humeral lobes (Fig.
Apparently prefers glabrous morphotypes, with pit galls mostly on the lower leaf surface, occasionally on the upper leaf surface.
Hawaii.
Appears to be widespread; collected from four regions that group into distinct clusters in the DNA analysis: (a) Puu Makaala, (b) Alili plus Kau, (c) Humuula (Hamakua Coast), and (d) the Kohala region (Fig.
Immatures make pit galls, typically on the lower leaf surface that are initially shallow, becoming deeper with older instars (Fig.
Named for the dark dorsal stripe that is frequently present in this species and signifies its affiliation with the
One of the largest species in the
Holotype male (slide mounted,
Variable, either entirely pale cream or yellow to greenish-yellow, or with head darker; a partial or weakly marked dorsal stripe extends from the head part or all the length of the body. Fore wing membrane clear or slightly fuscous.
Fore wing narrow, apex bluntly acute; surface spinules distributed in all cells, but limited in c+sc; short setae on margins and veins (Fig.
Unknown.
Colour and structure: Pale cream to orange-yellow. 5th instar: Ovoid and ventro-dorsally flattened wing buds only slightly protruding and distinct humeral lobes (Fig.
Collected on glabrous morphotypes.
Oahu.
Only known from one locality, the high elevation bog area on Mnt Kaala.
Immatures make pit galls on the lower leaf surface (Fig.
Named after Namaka, a sea goddess or water spirit in Hawaiian mythology, in reference to the type locality in the wet bog on top of Mnt Kaala (noun in the nominative singular standing in apposition to the generic name).
A pit-galling habit on Oahu was only recently discovered, previously pit-gallers were only known from younger islands and their presence on Oahu together with subfossils on Kauai (see Discussion) supports an older and more widespread status for the
Holotype male (slide mounted,
Currently this group includes only two species. These are small species, similar to those in the
Variable, typically mid- to dark brown throughout, recently emerged adults can be completely pale cream, head often darker than the rest of the body. A population in the Kilauea Iki caldera (form
Fore wing apex rounded; surface spinules fairly densely distributed in all cells; setae on margins and veins minute (Fig.
Unpigmented to light brown, elongate ovoid, not sinusoidal, smooth, without striations, short pedicel 1/5 length from base, tail short to moderately long (Fig.
Colour and structure: Smaller immatures are orange and cream, or yellow-brown, larger become blue-green or remain orange (e.g. Kilauea Iki population). 5th instar: Broadly ovoid in outline and ventro-dorsally flattened with wing buds protruding and distinct humeral lobes (Fig.
Usually on thick leaved, pubescent or semi-pubescent morphotypes.
Hawaii.
Widespread on Hawaii. The DNA analysis indicates distinct population clusters, but also more dispersal than for other taxa: two distinct clusters in Kohala, a mixed cluster from Olaa, Kilauea, Kau, and Kona Hema, and another cluster almost entirely from Saddle Road with a single Kau sample.
Makes pit galls on upper leaf surface. The leaf tissue often forms into a thickened rim of red or yellow around the immature (Fig.
Two forms are recognized (Fig.
Holotype, male (slide mounted,
Most specimens examined are almost entirely dark brown to black, however, as with
Fore wing apex rounded; surface spinules fairly densely distributed in all cells; setae on margins and veins minute (Fig.
Unknown.
Unknown.
Collected from pubescent morphotypes.
Maui.
Only known from eastern Maui, in the Makawao area.
Unknown, but likely to be pit galling given the biology of the sister taxon,
Named for the more dorsally humped (
Adults of this species are easily confused with
Holotype male (slide mounted,
The
1 | Fore wing membrane with distinct brown patches bordering wing veins, shape of male and female terminalia as in Fig. |
|
– | Fore wing membrane without distinct brown patches bordering wing veins, membrane either clear or slightly fuscous |
|
2 | Male with paramere extremely broad, with prominent posterior shoulder below apex, female terminalia short (≤0.60 × head width), female subgenital plate strikingly concave ventrally, shape of male and female terminalia as in Fig. |
|
– | Male with paramere narrower, more or less parallel sided or tapering, without or with only moderate posterior shoulder below apex, female terminalia ≥0.70 × head width, female subgenital plate more or less straight or slightly convex ventrally |
|
3 | Large species (antennal length >1.30 mm, head width >0.70 mm, wing length ≥2.95), female terminalia long (>0.80 mm) with apex acute and extremely small anal ring (ratio |
|
– | Smaller species (antennal length <1.30 mm, head width <0.70 mm, wing length <2.95), female terminalia shorter (<0.80 mm) with apex either acute, blunt or truncate, but ring relatively large (ratio |
|
4 | Larger species (antennal length >0.90 mm, head width ≥0.60 mm), with broader wings (ratio |
|
– | Smaller species (antennal length <0.90 mm, head width ≤0.60 mm), with narrower wings (ratio |
|
5 | Male with short proctiger and paramere (both ≤0.15 mm, ratio |
|
– | Male with longer proctiger and paramere (both >0.15 mm, ratio |
|
6 | Female with shorter terminalia ( |
|
– | Female with long terminalia ( |
|
7 | Female anal ring relatively small (ratio |
|
– | Female anal ring relatively large (ratio |
|
8 | General body colour brown to red, paramere tapering to slender neck below apex, egg with long sinuous tail, shape of male and female terminalia as in Fig. |
|
– | General body colour orange, yellow or yellow-brown, paramere tapering to apex or more parallel sided and constricted just below apex, egg with short bulbous tail |
|
9 | Male with paramere tapering gradually to apex, aedeagus hook less well developed, shape of male and female terminalia as in Fig. |
|
– | Male with paramere more parallel side, constricting just below apex, aedeagus hook more well developed, shape of male and female terminalia as in Fig. |
|
General body colour green, yellow-green or yellow-orange, often with brown on legs, thorax and abdomen. Females may have a darker abdomen due to darkly pigmented egg load. Fore wing membrane clear.
Fore wing apex rounded; surface spinules sparsely distributed, usually in all cells except limited or absent from cell r1; long setae on margins and particularly dense on the ventral margin, sparse long setae on veins (Fig.
Mid- to light brown, elongate oval, with longitudinal medial suture entire length of egg (coffee bean-like), surface with microsculpturing and granular in appearance, extremely short pedicel 1/3 length from base, tail lacking (Fig.
Unknown.
Collected predominantly from glabrous and semi-pubescent morphotypes.
Kauai.
Collected in several locations in Kokee State Park, including Alakai, Kalalau and Nu Alolo.
Based on phylogenetic closeness to
The female lectotype (dry mounted,
Two forms are recognized on Kauai (Fig.
Lectotype, female (dry mounted,
General body colour red-brown or brown. Fore wing membrane clear.
Fore wing apex rounded; surface spinules with limited distribution, few or none in cells r1, r2, m2; long setae on margins and veins (Fig.
Light brown, smooth, apparently without microsculpturing but with a granular appearance, short pedicel 1/4 length from base, tail lacking (Fig.
Colour and structure: Smaller instars orange, larger becoming yellow or blue-green with grey thorax and head. 5th instar ovoid in outline with wing buds protruding and nondistinct humeral lobes (Fig.
Collected predominantly from glabrous and semi-pubescent morphotypes.
Kauai.
Collected in two locations in Kokee State Park.
This species forms enclosed galls on leaves that resemble flat leaf galls in the
Named after Hiiaka in Hawaiian mythology, the favoured sister of Pele, who dwelled in a sacred Lehua grove and journeyed to Kauai (noun in the nominative singular standing in apposition to the generic name).
Some variation in paramere shape, particularly in development of anterior shoulder, is illustrated in Fig.
Holotype male (slide mounted,
General body colour mid- to dark brown. Fore wing membrane with brown pigmentation around wing base and patches of brown pigmentation bordering veins resulting in a distinct wing pattern (Fig.
Fore wing apex rounded; surface spinules with limited distribution in cells c+sc and cu2, and absent or very limited in all other cells; long setae on margins and veins (Fig.
Light brown, smooth, apparently without microsculpturing but with a slight granular appearance, short pedicel 1/4 length from base, tail lacking (Fig.
Unknown.
Morphotype preference unknown, adults collected on both glabrous and pubescent types.
Kauai.
Known from only one location in Kokee State Park.
Unknown, but its close relationship with
Named for the dark pigmentation around the fore wing veins (adjective in the nominative singular).
This species is the only member of
Holotype male (slide mounted,
General body colour is either brown to yellow-brown, or green. Fore wing membrane clear.
Fore wing apex rounded; surface spinules with limited distribution in all cells except absent from r1; long setae on margins and medium long on veins (Fig.
Light brown, smooth, apparently without microsculpturing, short pedicel 1/4 length from base, tail lacking (Fig.
Unknown.
Morphotype preference unknown, adults collected on both glabrous and pubescent types.
Kauai.
Known from only one location, Kalalau Valley in Kokee State Park.
Unknown, but morphological affinities with
The name refers to the large size of the species, it is the largest of the
This is the largest of the
Holotype male (slide mounted,
General body colour is either brown to dark brown, or light red to red-brown. Fore wing membrane clear or slightly fuscous.
Fore wing apex bluntly acute; surface spinules distributed in all cells except few or none in r1; short setae on margins and veins (Fig.
Unpigmented to light brown, elongate and sinusoidal, no microsculpturing, short pedicel 1/4 length from base, long tail with slightly inflated tip (Fig.
Colour and structure: Black or brown dorsally, cream to pale orange ventrally. 5th instar: Broadly ovoid in outline, wing buds only slightly protruding with distinct humeral lobes (Fig.
Found predominantly on glabrous and semi-pubescent morphotypes.
Kauai.
The two recognized forms (
This species forms thin-walled cup galls on stems, often clustered together, with one immature per gall chamber (Fig.
The name refers to the gall type which is a cup (
Two forms are recognized (Fig.
Holotype male (slide mounted,
General body colour pale yellow to orange (Fig.
Fore wing apex bluntly acute; surface spinules dispersed, usually present in all cells but may be limited or absent; short setae on margins and veins (Fig.
Light brown, short, slightly sinusoidal, surface granular in appearance, no microsculpturing, short pedicel 1/4 length from base, tail bulbous (Fig.
Colour and structure: Brown to orange-brown dorsally, cream to orange ventrally. 5th instar: Broadly ovoid in outline (but narrower than
Only known from one locality where it galls densely pubescent bog morphotypes.
Kauai.
Known only from Alakai Swamp, Kokee State Park.
Forms thick-walled cup galls on stems, galls are often clustered together, with one individual per gall chamber (Fig.
The name refers to the gall type which is a thick (
This species is a sister taxon to the other known cup gall maker,
Holotype male (slide mounted,
General body colour yellow or orange. Fore wing membrane clear.
Fore wing apex rounded; surface spinules distributed in all cells; short setae on margins and veins (Fig.
Unpigmented, not sinusoidal, smooth, no microsculpturing, short pedicel 1/4 length from base, tail lacking.
Unknown (see comment under
Unknown.
Kauai
The type location is recorded only as “Nualolo”.
The biology of this species is unknown, it may form cup galls on stems as morphologically it is close to the two stem cup-gallers,
Holotype, male (?) (dry mounted, damaged, abdomen and fore wings missing,
General body colour brown. Fore wing membrane clear.
Fore wing apex bluntly acute; surface spinules sparsely distributed, few or none in cells r1, cu2, c+sc; short setae on margins and veins (Fig.
Unpigmented, short, not sinusoidal, no microsculpturing, pedicel not visible, tail lacking (Fig.
Unknown.
Collected from glabrous morphotype.
Kauai.
Only known location is Kalalau Valley, Kokee State Park.
Unknown.
The name refers to the small and elegant appearance with slender elongate female terminalia and long, slender tibiae (adjective in the nominative singular).
Known from only one female specimen; the distinctly long, slender terminalia is unlike any other described species.
Holotype female (slide mounted,
General body colour yellow with darker yellow-brown dorsally. Fore wing membrane clear or slightly fuscous basally.
Fore wing apex bluntly acute; surface spinules sparsely distributed, absent from r1 and c+sc; short to minute setae on margins and veins (Fig.
Unpigmented to light brown, long and narrow, not sinusoidal, surface with broadly spaced longitudinal striations that are either continuous or interrupted, pedicel appears to be absent, tail lacking (Fig.
Unknown.
Morphotype preference unknown.
Kauai.
Only known location is Kalalau Valley, Kokee State Park.
Unknown.
Named after Betsy Gagné to honour her role in promoting biodiversity research, entomology, and conservation in the Hawaiian Islands (noun in the genitive case).
Known from only one female specimen; the distinctly shaped female terminalia and egg characteristics are not found in other species.
Holotype female (slide mounted,
General body colour yellow to pale brown. Fore wing membrane clear or slightly fuscous.
Fore wing apex bluntly acute to almost rounded; surface spinules sparsely distributed, absent from r1 and c+sc; short to minute setae on margins and veins (Fig.
Unknown.
Unknown.
Collected from glabrous forest tree.
Kauai.
Only known location is Alakai, Kokee State Park, in forest near Alakai bog area.
Unknown.
Named after Haumea in Hawaiian mythology, the Hawaiian goddess of fertility and mother of Pele (noun in the nominative singular standing in apposition to the generic name).
Known from only one male specimen; the distinctly shaped male paramere is not found in other species; morphologically it appears most closely related to
Holotype male (slide mounted,
The
Due to a high degree of intraspecific variation and inter-island convergence in the
1 | Generally larger species ( |
|
– | Generally smaller species ( |
|
1 | Generally larger species, male with longer, broader paramere (ratio |
|
– | Generally smaller species, male with shorter, narrower paramere (ratio |
|
1 | Smaller species with short genal processes ( |
|
Larger species with long genal processes ( |
|
|
2 | Female terminalia shorter ( |
|
Female terminalia longer ( |
|
1 | Intermediate sized species, male with shorter paramere ( |
|
Larger or smaller species, male with longer paramere ( |
|
|
2 | Larger species, longer antennae ( |
|
Smaller species, shorter antennae ( |
|
General body colour yellow-green to yellow-brown. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear.
Fore wing apex rounded; spinules distributed in all cells, but few in r1; medium to long setae on margins and veins (Fig.
Short, broad, pigmented brown to dark brown (except tip of pedicel and tail) with surface microsculpturing, either with long pedicel and tail, the pedicel with an inflated tip (forms
Colour and structure: 5th instar: Cream to orange. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (Fig.
Primarily on more pubescent morphotypes.
Oahu.
The distribution ranges of the three recognized forms overlap, all three are found in both Waianae and Koolau ranges, but rarely at the same collection site suggesting microecological divergence may play a role in explaining this diversity; form
Usually galls stems, buds and petioles resulting in irregular swellings (Fig.
Named for its distribution on the island of Oahu (noun in the genitive case).
One of the commonest species on Oahu. It can be distinguished most easily from other Oahu species by its typically much larger size and predominantly yellow-green or yellow-brown colour. This species encompasses
Holotype male (slide mounted,
General body colour red-brown to orange-brown. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear or fuscous.
Fore wing apex acute to bluntly acute; spinules distributed in all cells; short to medium-short setae on margins and veins (Fig.
Colour and structure: 2nd-5th instars: Orange or orange-red with cream wing buds. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (Fig.
Mostly associated with glabrous morphotypes.
Oahu.
A widespread taxon and probably the most commonly encountered on Oahu, but as with
Makes flat leaf galls. 1st instars are found in very shallow pits (Fig.
Four forms are recognized (Figs
Holotype, male (dry mounted,
No new material was collected during this study. Below is a summary of the description from
General body colour yellow to brown. Fore wing membrane clear or slightly fuscous, short setae on margins and veins. Reported size is similar to
Unknown.
Lanai, Molokai.
Apparently common on Lanai (
Unknown, but it likely makes enclosed galls, and if Crawford’s hypothesis of parallel divergence to that on Oahu is correct, then this may be a stem galling sister taxon to a leaf galling
Holotype, female (dry mounted,
No new material was collected during this study. Below is a summary of the description from
Generally body colour dark brown to black, probably the darkest of the
Unknown.
Probably
Lanai.
Known from two localities on Lanai: “Waiopao” (“Waiopaa, west side” in
Unknown, but it likely makes enclosed galls, and if Crawford’s hypothesis of parallel divergence to that on Oahu is correct (see comment for
No type material was found at
General body colour green, or yellow-green to yellow-brown. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear.
Fore wing apex rounded; spinules distributed in all cells, except few or none in r1; medium-short setae on margins and veins (Fig.
Short, broad, pigmented brown to dark brown (except tip of pedicel and tail) with surface microsculpturing, medium-long pedicel with slightly inflated tip, tail short (Fig.
Unknown.
Collected from glabrous and pubescent morphotypes.
Molokai.
Kamakou Preserve (this study), Kamiloloa, Kawela (
Unknown, but morphology suggests it may gall stems/buds and may have made the bud gall in Fig.
Two forms are recognized (Fig.
Holotype, female (dry mounted,
General body colour dark red to red-brown, or yellow-brown. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear.
Fore wing apex bluntly acute; spinules distributed in all cells; short to medium-short setae on margins and veins (Fig.
Short, broadly ovoid to almost circular, pigmented brown (except tip of pedicel and tail), surface with microsculpturing, short pedicel with slightly inflated tip, tail extremely short (Fig.
Unknown.
Preference unknown; collected from glabrous and pubescent morphotypes.
Molokai.
Known only from Kamakou Preserve.
Unknown; but morphology suggests it may be a leaf galler.
Named after Hualani, a High Chiefess of Molokai in ancient times (noun in the nominative singular standing in apposition to the generic name).
Molecular data recovers this taxon as sister to
Holotype male (slide mounted,
General body colour yellow-brown to green. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear, or slightly fuscous.
Fore wing apex rounded; spinules distributed in all cells; short setae on margins and veins (Fig.
Short, broad, pigmented brown to dark brown (except tip of pedicel and tail), surface with coarse microsculpturing, and either with distinct medium-long pedicel with slightly inflated tip, or pedicel obscured and an unsclerotized patch at the base of the egg, tail medium-short (Fig.
Unknown.
Collected from glabrous morphotypes.
Maui.
Known from east and west Maui; there is some geographical clustering in the molecular data but the east/west divergence of haplotypes is not as distinct as in
Unknown; but morphology suggests it may gall stems/buds.
Named for the distribution on the island of Maui (noun in the genitive case).
Two forms are recognized (Fig.
Holotype male (slide mounted,
General body colour yellow-brown to green. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear, or slightly fuscous.
Fore wing apex bluntly acute to rounded; spinules sparsely distributed in all cells except r1; short setae on margins and veins (Fig.
Short, broadly ovoid to almost circular, pigmented brown (except tip of pedicel and tail), surface with microsculpturing, short pedicel with slightly inflated tip, tail extremely short.
Colour and structure: 5th instar: Cream to orange. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (similar to
Collected from glabrous morphotypes.
Maui.
Known from east and west Maui; the molecular analysis clearly distinguishes eastern from western haplotypes.
Galls stems and occasionally petioles, resulting in irregular swellings.
Named after the
Molecular data recovers this taxon as sister to
Holotype male (slide mounted,
Generally body colour orange-red to red-brown. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear, or slightly fuscous.
Fore wing apex bluntly acute to rounded; spinules distributed in all cells but few in r1; medium-short setae on margins and veins. Antennae medium-long (av. length 1.06; ratio
Short, broad, pigmented brown to dark brown (except tip of pedicel and tail) with fine surface microsculpturing, medium-long pedicel with slightly inflated tip, tail extremely short or absent (Fig.
Colour and structure: 5th instar: Cream to orange. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (Fig.
Collected from glabrous morphotypes.
Maui.
Known from east and west Maui; molecular data indicates very distinct eastern and western populations that are characterized by the two recognized forms.
Makes flat leaf galls (Fig.
Named after Steve Montgomery, an extraordinary field biologist who made a substantial contribution to this study (noun in the genitive case).
Two forms are recognized (Fig.
Holotype male (slide mounted,
General body colour green or yellow-green, but often with brown on legs, thorax and abdomen. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear.
Fore wing apex rounded; spinules with limited distributed in all cells; medium-short to short setae on margins and veins (Fig.
Short, broad, pigmented brown to dark brown (except tip of pedicel and tail), surface with microsculpturing, medium-long pedicel with slightly inflated tip, tail short (Fig.
Colour and structure: 5th instar: Cream to orange. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (Fig.
Known from both glabrous and pubescent morphotypes, but mostly associated with pubescent and semi-pubescent types.
Hawaii.
Widely distributed on the island of Hawaii; there are some distinct clusters in the molecular analysis, with the southern and western populations sister to and nested within members from Kohala (north), and these are distinct from two individuals from the central Saddle Rd area.
Galls stems and buds, resulting in irregular swellings, and in the case of buds, inhibits bud growth (Fig.
Individuals from Kona Hema have a notably broader paramere.
Holotype, male (?) (dry mounted, damaged, abdomen missing,
General body colour dark brown to red-brown, the head and genal processes are often darker or black. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear, or slightly fuscous.
Fore wing apex rounded; spinules distributed in all cells, except few or none in r1; medium-short setae on margins and veins (Fig.
Short, broad (almost circular in Kohala form), pigmented brown to dark brown (except tip of pedicel and tail) with surface microsculpturing, medium-long pedicel with slightly inflated tip, tail extremely short or absent (Fig.
Colour and structure: 5th instar: Cream to orange. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (Fig.
Known from both glabrous and pubescent morphotypes, but mostly associated with glabrous and semi-pubescent forms.
Hawaii.
The most common species on Hawaii. In addition to a distinct form “Kohala” only known from the Kohala region, within the “common” form there are two groups both of which are widespread: group 1 includes distinct populations from Kona Hema (south west), Kohala (north east), Saddle Rd (central), and Hualalai (north west); group 2 is more mixed with individuals from south, central, east and west, but not from Kohala.
Makes flat galls on leaves (Fig.
Named after Pele, the volcano and fire goddess in Hawaiian mythology (noun in the nominative singular standing in apposition to the generic name).
Two forms are recognized (Figs
Holotype male (slide mounted,
General body colour brown or yellow-brown, or yellow-green, the genal processes are often paler than the head. Females often appear to have a dark abdomen due to darkly pigmented egg load. Fore wing membrane clear, or slightly fuscous.
Fore wing apex rounded; spinules distributed in all cells; short setae on margins and veins (Fig.
Short, broad, pigmented brown to dark brown (except tip of pedicel and tail) with surface microsculpturing, long pedicel with inflated tip, tail long (Fig.
Colour and structure: 5th instar: Cream to orange. Elongate ovoid in outline, wing buds protruding with moderate humeral lobes (similar to
Known from both glabrous and pubescent morphotypes.
Hawaii.
This species is widely distributed on Hawaii. The molecular data suggests an initial diversification in the south western part of the island and subsequent spread north, west, and east.
Although predominantly makes cone galls on leaves (Fig.
Named for the shape of the pyramid-like cone gall produced on leaves (adjective in the nominative singular).
This species is sister to
Holotype male (slide mounted,
Map indicating sampling sites for
Map indicating sampling sites for
Map indicating sampling sites for
Financial support that assisted in fieldwork is gratefully acknowledged from a Smithsonian Institution Postdoctoral Fellowship (2004–2005), and an NSF Dimensions of Biodiversity award [DEB 1241253] to the University of California, Berkeley. Molecular sequencing was partly supported by a Smithsonian Institution Fellowship, and the Natural History Museum, London Molecular Taxonomy Fund (DIF). I am grateful to the Bernice
Figure S1
PNG image file
Illustration of adult characters and measurements referred to in the text.