Mite (Acari) Communities Associated with 'Ohi'a, Metrosiderospolymorpha (Myrtaceae), at Hono 0 Nil Pall and Kui'a Natural Area

Reserves on Kaua'i Island, Hawaiian Islands1

Sabina F. Swift and M. Lee GofP

Abstract: Native 'Ohi'a trees (Metrosideros polymorpha) were sampled for mites attwo natural area reserves on Kaua'i Island, Kui'a and Hono 0 N:.ii Pali. Ninetysamples of leaves, flowers, bark, leaf litter, and soil under the 'Ohi'a canopieswere taken. Mites were extracted with use of Berlese-Tullgren funnels. Onehundred sixty-four species were found, with the suborder Prostigmata havingthe greatest number of species (74), followed by Mesostigmata (43), Oribatida(43), and Astigmata with the least (4). Leaf litter, leaf litter with soil, and barkhave the most species, composed of predaceous mesostigmatic and prostigmaticmites, but a certain amount of overlap of mite species between the leaf litter andsoil habitats was observed. The predominance of Collembola in the soil andlitter samples indicates a stable food source for the predaceous mites, partly ex­plaining the high number of mites in those habitats. Oribatid mites were col­lected from leaves, but the species composition differs from that on flowers andlitter. Preliminary residency status of identified taxa shows 12% endemic, 17%adventive, and 71 % of unknown status.

MITES OR ACARI are one of the largest andmost biologically diverse groups of the arach­nids, rivaling insects in the extent to whichthey have successfully colonized aquatic andterrestrial habitats (Evans 1992). Mites are animportant part of the Hawaiian ecosystems.Although approximately 630 species havealready been named and reported (Nishida1997, Swift and Norton 1998), mites remainone of the most poorly known arthropodgroups in the Islands. Studies of Acari asso­ciated with particular plants, soils, and litterhabitats are relatively uncommon in the Ha­waiian Islands. Although sampling of litterfrom an area or location to determine its mitefauna has been done, there has been little

1 Funding for this research was provided by Hawai'iDepartment of Land and Natural Resources through aNatural Area Reserves Grant to the authors. Manuscriptaccepted 10 March 2000.

2 Department of Entomology, 3050 Maile Way, 310Gilmore Hall, University of Hawai'i at Manoa, Hono­lulu, Hawai'i 96822.

Pacific Science (2001), vol. 55, no. 1:23-40© 2001 by University of Hawai'i PressAll rights reserved

23

systematic collecting from specific plant hostsor habitats. In the early 1970s, the Interna­tional Biological Program (IBP), through theBishop Museum, sampled mites ~n soil andlitter in an elevational transect of 'Ohi'a foresteast of Mauna Loa Trail on Hawai'i Island.From these collections, work by Radovskyand Tenorio (1981) on the soil Mesostigmata(Parasitiformes) and Collembola gave an in­dication of the diversity of soil and litterf,!una living under the canopies of the native'Ohi'a. Gagne (1979) _sampled arthropods in­cluding mites from 'Ohi'a and Koa canopieswith pyrethrum fogging. Unfortunately, themites recoveresi from the canopy-samplingtechnique on 'Ohi'a and Koa and most mitescollected from the Maunaloa Trail transectcurr~ntly remain unstudied.

'Ohi'a, Metrosideros polymorpha (Gaud.)(Myrtaceae), is a dominant native Hawaiiantree found from sea level on wetter slopes upto 2590 m elevation. It is distributed on thesix largest islands- in areas with annual averagerainfall ranging fro_m 75 cm to 11.5 m (Cornand Heisy 1973). 'Ohi'a was the subject of in­tensive research for over 20 yr because of pe­riodic breakdown of stands known as " 'Ohi'arainforest dieback" (Mueller-Dombois 1985).

Mite (Acari) Communities Associated with 'Ohi'a, Metrosiderospolymorpha (Myrtaceae), at Hono 0 Nil Pall and Kui'a Natural Area

Reserves on Kaua'i Island, Hawaiian Islands1

Sabina F. Swift and M. Lee GofP

Abstract: Native 'Ohi'a trees (Metrosideros polymorpha) were sampled for mites attwo natural area reserves on Kaua'i Island, Kui'a and Hono 0 N) Pali. Ninetysamples of leaves, flowers, bark, leaf litter, and soil under the 'Ohi'a canopieswere taken. Mites were extracted with use of Berlese-Tullgren funnels. Onehundred sixty-four species were found, with the suborder Prostigmata havingthe greatest number of species (74), followed by Mesostigmata (43), Oribatida(43), and Astigmata with the least (4). Leaf litter, leaf litter with soil, and barkhave the most species, composed of predaceous mesostigmatic and prostigmaticmites, but a certain amount of overlap of mite species between the leaf litter andsoil habitats was observed. The predominance of Collembola in the soil andlitter samples indicates a stable food source for the predaceous mites, partly ex­plaining the high number of mites in those habitats. Oribatid mites were col­lected from leaves, but the species composition differs from that on flowers andlitter. Preliminary residency status of identified taxa shows 12% endemic, 17%adventive, and 71 % of unknown status.

MITES OR ACARI are one of the largest andmost biologically diverse groups of the arach­nids, rivaling insects in the extent to whichthey have successfully colonized aquatic andterrestrial habitats (Evans 1992). Mites are animportant part of the Hawaiian ecosystems.Although approximately 630 species havealready been named and reported (Nishida1997, Swift and Norton 1998), mites remainone of the most poorly known arthropodgroups in the Islands. Studies of Acari asso­ciated with particular plants, soils, and litterhabitats are relatively uncommon in the Ha­waiian Islands. Although sampling of litterfrom an area or location to determine its mitefauna has been done, there has been little

I Funding for this research was provided by Hawai'iDepartment of Land and Natural Resources through aNatural Area Reserves Grant to the authors. Manuscriptaccepted 10 March 2000.

2 Department of Entomology, 3050 Maile Way, 310Gilmore Hall, University of Hawai'i at Manoa, Hono­lulu, Hawai'i 96822.

Pacific Science (2001), vol. 55, no. 1:23-40© 2001 by University of Hawai'i PressAll rights reserved

23

systematic collecting from specific plant hostsor habitats. In the early 1970s, the Interna­tional Biological Program (IBP), through theBishop Museum, sampled mites ~n soil andlitter in an elevational transect of 'Ohi'a foresteast of Mauna Loa Trail on Hawai'i Island.From these collections, work by Radovskyand Tenorio (1981) on the soil Mesostigmata(Parasitiformes) and Collembola gave an in­dication of the diversity of soil and litterf'!una living under the canopies of the native'Ohi'a. Gagne (1979) _sampled arthropods in­cluding mites from 'Ohi'a and Koa canopieswith pyrethrum fogging. Unfortunately, themites recoveresi from the canopy-samplingtechnique on 'Ohi'a and Koa and most mitescollected from the Maunaloa Trail transectcurr~ntly remain unstudied.

'Ohi'a, Metrosideros polymorpha (Gaud.)(Myrtaceae), is a dominant native Hawaiiantree found from sea level on wetter slopes upto 2590 m elevation. It is distributed on thesix largest islands- in areas with annual averagerainfall ranging fro_m 75 cm to 11.5 m (Cornand Heisy 1973). 'Ohi'a was the subject of in­tensive research for over 20 yr because of pe­riodic breakdown of stands known as " 'Ohi'arainforest dieback" (Mueller-Dombois 1985).

24

Although the effects of arthropods (e.g., woodborers) were studied extensively, the pos~ibil­

ity of mites as a causal organism for 'Ohi'adieback was never considered. This studys~eks to identify mite species associated with'Ohi'a and to expand the general knowledgeof arthropods associated with this native tree.It is well known that sustainable managementof any forest requires reliable knowledge ofarthropod species and their ecological roles inthe forest ecosystem.

Because of the current incomplete stateof acarine systematics in many parts of theworld, there has been a reluctance to desig­nate residency for mite species describedfrom Hawai'i (Goff 1987, Swift and Norton1998). The preliminary designations forspecies associated with 'Ohi'a in this reportare for the primary purpose of assessments ofthe conservation status of the natural areareserves. For the purposes of this report,endemic refers to taxa found only in theHawaiian Islands; adventive refers to thosetaxa that are immigrant species but not inten­tionally introduced; and introduced refers tothose taxa intentionally brought in for use inbiological control programs. A "?" for statusof a species in the Appendix indicates anidentification only to genus level and statusis unknown. Undescribed species from theHawaiian Islands and species identified as"near" a previously described species (e.g.,Neoeypholaelaps TIr. lindquisti Prasad, 1968) areconsidered here to be endemic until shownotherwise.

MATERIALS AND METHODS

Sampling Sites

KUI'A NATURAL AREA RESERVE: Kui'a Nat­ural Area Reserve (Figure 1) occupies 662 hain the Waimea District of Kaua'i Island(Department of Land and Natural Resources,State of Hawai'i· 1989b). Elevation rangesfrom 600 to 1170 m. Rainfall averages 102­203 cm annually, with December being thewettest month and June the driest (Giambel­luca et al. 1986). Gradual to moderate moun­tain slopes cut by intermittent streams indrier sections of northeastern Koke'e are the

PACIFIC SCIENCE· January 2001

reserve's main features. The lowland sectionhas dry and semiwet shrublands with 'Ohi'aand Koa forests, and rare plants.

Kui'a Reserve lies partially within the NaPali Kona Forest Reserve and adjacent toKoke'e State Park. The reserve is traversedby Nu'alolo Trail; north of the reserve is theAwa'awapuhi Trail, and south is Miloli'iRidge. Outside the reserve's eastern boundaryare Koke'e Lodge, Koke'e Natural HistoryMuseum, and cabins of Koke'e State Park,accessed by the two-lane Highway 55.

The MetrosideroslAcacia koa mixed forestsin the higher half of the reserve along theNu'alolo Trail are over 9 m in height. Treesin the lower elevation consist of younger andshorter stands (1.5-4 m) of Metrosideros.Samples of accessible leaves and fl<:?wers weretaken from these lower stands of 'Ohi'a.

Three sampling sites along Nu'alolo Trailwere marked: top (1100 m), middle (975 m),and bottom (750 m). Samples were takenfrom these three sites during the duration ofthe study. Samples were collected on 27 No­vember 1990 and 29 April 1991.

HONO 0 NA PALl NATURAL AREA RESERVE:

The Hono 0 Na Pali Natural Area Reserve(Figure 1) occupies 1275 ha in the HanaleiDistrict (Department of Land and NaturalResources, State of Hawai'i 1989a). Itstretches from sea level along the Na Palicoast crossing Kalalau Trail and rises south tothe highest point at Pihea (1285 m) to thenorthern edge of Alaka'i Swamp. In the driercoastal lowlands, annual rainfall averages203 cm, increasing to more than 406 cm inthe upland rain forests (Giambelluca et al.1986). Along the coast, the reserve encom­passes parts of Hanakapl'ai and HanakoaStreams and all of Waiahuakua Stream.These intermittent and continuous streamsand riparian and ridgeline lowland and mon­tane semiwet and wet forests are the mainfeatures of the reserve. Rare plants, rarestream animals, and rare forest bird habitatsare prevalent in the reserve.

Upper Hono 0 Na Pali Reserve is com­posed of an e!oded plateau wi~ a series ofridges with 'Ohi'a/mixed and 'Ohi'a/Uluhemontane wet forest. Along the upper south­west corner of the reserve boundary

24

Although the effects of arthropods (e.g., woodborers) were studied extensively, the pos~ibil­

ity of mites as a causal organism for 'Ohi'adieback was never considered. This studys~eks to identify mite species associated with'Ohi'a and to expand the general knowledgeof arthropods associated with this native tree.It is well known that sustainable managementof any forest requires reliable knowledge ofarthropod species and their ecological roles inthe forest ecosystem.

Because of the current incomplete stateof acarine systematics in many parts of theworld, there has been a reluctance to desig­nate residency for mite species describedfrom Hawai'i (Goff 1987, Swift and Norton1998). The preliminary designations forspecies associated with 'Ohi'a in this reportare for the primary purpose of assessments ofthe conservation status of the natural areareserves. For the purposes of this report,endemic refers to taxa found only in theHawaiian Islands; adventive refers to thosetaxa that are immigrant species but not inten­tionally introduced; and introduced refers tothose taxa intentionally brought in for use inbiological control programs. A "?" for statusof a species in the Appendix indicates anidentification only to genus level and statusis unknown. Undescribed species from theHawaiian Islands and species identified as"near" a previously described species (e.g.,Neoeypholaelaps TIr. lindquisti Prasad, 1968) areconsidered here to be endemic until shownotherwise.

MATERIALS AND METHODS

Sampling Sites

KUI'A NATURAL AREA RESERVE: Kui'a Nat­ural Area Reserve (Figure 1) occupies 662 hain the Waimea District of Kaua'i Island(Department of Land and Natural Resources,State of Hawai'i· 1989b). Elevation rangesfrom 600 to 1170 m. Rainfall averages 102­203 cm annually, with December being thewettest month and June the driest (Giambel­luca et al. 1986). Gradual to moderate moun­tain slopes cut by intermittent streams indrier sections of northeastern Koke'e are the

PACIFIC SCIENCE· January 2001

reserve's main features. The lowland sectionhas dry and semiwet shrublands with 'Ohi'aand Koa forests, and rare plants.

Kui'a Reserve lies partially within the NaPali Kona Forest Reserve and adjacent toKoke'e State Park. The reserve is traversedby Nu'alolo Trail; north of the reserve is theAwa'awapuhi Trail, and south is Miloli'iRidge. Outside the reserve's eastern boundaryare Koke'e Lodge, Koke'e Natural HistoryMuseum, and cabins of Koke'e State Park,accessed by the two-lane Highway 55.

The Metrosideros/Acacia koa mixed forestsin the higher half of the reserve along theNu'alolo Trail are over 9 m in height. Treesin the lower elevation consist of younger andshorter stands (1.5-4 m) of Metrosideros.Samples of accessible leaves and fl<:?wers weretaken from these lower stands of 'Ohi'a.

Three sampling sites along Nu'alolo Trailwere marked: top (1100 m), middle (975 m),and bottom (750 m). Samples were takenfrom these three sites during the duration ofthe study. Samples were collected on 27 No­vember 1990 and 29 April 1991.

HONO 0 NA PALl NATURAL AREA RESERVE:

The Hono 0 Na Pali Natural Area Reserve(Figure 1) occupies 1275 ha in the HanaleiDistrict (Department of Land and NaturalResources, State of Hawai'i 1989a). Itstretches from sea level along the Na Palicoast crossing Kalalau Trail and rises south tothe highest point at Pihea (1285 m) to thenorthern edge of Alaka'i Swamp. In the driercoastal lowlands, annual rainfall averages203 cm, increasing to more than 406 cm inthe upland rain forests (Giambelluca et al.1986). Along the coast, the reserve encom­passes parts of Hanakapl'ai and HanakoaStreams and all of Waiahuakua Stream.These intermittent and continuous streamsand riparian and ridgeline lowland and mon­tane semiwet and wet forests are the mainfeatures of the reserve. Rare plants, rarestream animals, and rare forest bird habitatsare prevalent in the reserve.

Upper Hono 0 Na Pali Reserve is com­posed of an e!oded plateau wit4. a series ofridges with 'Ohi'a/mixed and 'Ohi'a/Uluhemontane wet forest. Along the upper south­west corner of the reserve boundary

Mite Communities Associated with 'Ohi'a . Swift and Goff

Hono 0 Na Pali

25

KAVA'I

FIGURE 1. Topographic map of Kaua'i Island showing locations of Kui'a and Hono 0 Nii Pali Natural Area Reserves.

where samples from habitats were collected,the 'Ohi'a trees were mixed with Lapalapa(Cheirodendron platyphyllum subsp. kauaiense),'Olapa (Cheirodendron trigynum), Kawa'u (Ilexanomala), Kolea (Myrsine lessertiana and M.alyxifolia), and 'Ohe (Tetraplasandra spp.).The _mixed shrub layer contained speciesof 'Ohelo kau la'au (Vaccinium calycinum),patches of Uluhe (Dicranopteris linearis), treefern (Cibotium), Naupaka (Scaevola procera),Styp~elia, and Dodonaea. Trunks and twigsof 'Ohi'a were well covered with epiphyticmosses and liverworts. Diverse ground coverof mosses, ferns, and a thick layer ofleaflitteron spongy humus dominates the forest floor.No signs of animal disturbance were ob­served.

The coastal section of Hono 0 Na PaliReserve is traversed by Kalalau Trail withsparse 'Ohi'a communities. These trees weregrowing either below or above the trail andthus barely accessible for sampling. Duringthe one-time sampling of the coastal Hono 0Na Pali Reserve, it had not rained for over 2weeks, rendering the area extremely dry.

The upper Hono 0 Na Pali Reserve wassampled on 19 November 1990,18 February1991, and 13 July 1991. The coastal section ofthe reserve was sampled on4 September 1991.

Sampling and Extraction Techniques

A total of 90 samples of flowers, leaves, bark,bark with moss, litter, litter with soil, and soil

Mite Communities Associated with 'Ohi'a . Swift and Goff

Hono 0 Na Pali

25

KAVA'I

FIGURE I. Topographic map of Kaua'i Island showing locations of Kui'a and Hono 0 Nii Pali Natural Area Reserves.

where samples from habitats were collected,the 'Ohi'a trees were mixed with Lapalapa(Cheirodendron platyphyllum subsp. kauaiense),'Olapa (Cheirodendron trigynum), Kawa'u (Ilexanomala), Kolea (Myrsine lessertiana and M.alyxifolia), and 'Ohe (Tetraplasandra spp.).The _mixed shrub layer contained speciesof 'Ohelo kau la'au (Vaccinium calycinum),patches of Uluhe (Dicranopteris linearis), treefern (Cibotium), Naupaka (Scaevola procera),Styp~elia, and Dodonaea. Trunks and twigsof 'Ohi'a were well covered with epiphyticmosses and liverworts. Diverse ground coverof mosses, ferns, and a thick layer of leaf litteron spongy humus dominates the forest floor.No signs of animal disturbance were ob­served.

The coastal section of Hono 0 Na PaliReserve is traversed by Kalalau Trail withsparse 'Ohi'a communities. These trees weregrowing either below or above the trail andthus barely accessible for sampling. Duringthe one-time sampling of the coastal Hono 0Na Pali Reserve, it had not rained for over 2weeks, rendering the area extremely dry.

The upper Hono 0 Na Pali Reserve wassampled on 19 November 1990,18 February1991, and 13 July 1991. The coastal section ofthe reserve was sampled on 4 September 1991.

Sampling and Extraction Techniques

A total of 90 samples of flowers, leaves, bark,bark with moss, litter, litter with soil, and soil

26

TABLE 1

Number of Habitat Samples Collected from Hono 0Nil Pali and Kui'a Natural Area Reserves

Sample Type Hono 0 Nil Pali" Kui'ab

Flower 3 3Leaves 14 10Bark 6 14Bark with moss 8 1Leaf litter 9 10Leaf litter with soil 9 3Soil 0 5

Total 49 41

"Three collections from upper elevation on Pihea, and onecollection from coastal area along Kalalau Trail.

b Two collections along Nu'alolo Trail.

was taken (Table 1). A litter sample consistedof leaves and small twigs lying on top of themineral soil showing signs of decomposition.A soil sample was the mineral soil under thelitter scooped to desired depth. Each littersample was taken with a lightweight plastictrowel, approximately three trowelsful (ap­proximately three to four cups [700-950 cm J]

dry weight). When mineral soil was in­advertently included in the litter during sam­pling, the category became "litter with soiL"At Hono 0 Na Pali, moss and liverwortgrowth on trunks and twigs of 'Ohi'a waspredominant, making it difficult to samplebark without moss or liverworts. Few flowersamples were taken because they were located4 to 10m up in the canopy. Flowering seasonwas also a factor in the low number of flowersamples. Leaves were picked off singly fromtree branches. Efforts to standardize leaf sam­ples at 20-30 leaves per sample were main­tained during sampling. Samples wereindividually bagged in half-gallon (1893 cmJ)

plastic bags and labeled. Plastic bags were leftopen in the canvas carrier to provide a con­tinuous supply of oxygen to the live micro­organisms in the bag and to reduce con­densation inside the bag, deemed lethal to thetiny mites. Closing the bags was unnecessarybecause most mites, unlike ants and spring­tails, will not wander off from the substrate.Samples were transported the same day to theAcarology Laboratory at the University of

PACIFIC SCIENCE· January 2001

Hawai'i at Manoa for extraction. Mites wereextracted using Tullgren-Berlese funnels(Krantz 1978).

Extracted mites in 70% ethyl alcohol weresorted to suborders. Samples of each mor­phospecies were mounted on microslides withHoyer's mounting medium, then later driedin an oven for 2 weeks at 45-50°C and sealedwith Glyptal electric paint before examina­tion. A phase-contrast microscope (Wild) wasused in mite identification. Voucher speci­mens were kept at the J. Linsley GressittCenter for Research in Entomology of theBishop Museum, and the Acarology Labora­tory, University of Hawai'i at Manoa.

RESULTS AND DISCUSSION

Acarine Population, Composition, and Densities

The suborders Mesostigmata (Gamasida),Prostigmata (Actinedida), Astigmata (Acari­dida), and Oribatida (Cryptostigmata) wereall represented at Kui'a and Hono 0 Na PaliNatural Area Reserves. Approximately 33,400mites were extracted from 90 habitat sam­ples. Of the mite total, 67% (22,500) wereOribatida, 23% (7,711) Prostigmata, 19%(2,568) Mesostigmata, and 2% (592) Astig­mata. Of the 164 species found, 74 were ofthe suborder Prostigmata, 43 were Meso­stigmata, 4 were Astigmata, and 43 wereOribatida (Appendix). Irnrnatures composed35-40% of the entire mite collection.

MESOSTIGMATA: (Figure 2). A large andcosmopolitan suborder, mites in this grouphave adapted successfully to a remarkablywide range of habitats. Many are free-livingpredators in the soil, in litter, or on plants,where they prey on other small invertebrates(Krantz and Ainscough 1990). Many speciesare parasites of mammals, birds, reptiles, orinvertebrates (Strandtrnann and Wharton1958, Yunker 1973, Treat 1975). Mesostig­matic mites are generally well sclerotized.They range in size from 200 to 2500 /lm.

A total of 43 species of mesostigmaticmites was recorded on flowers, leaves, bark,leaf litter, leaf litter and soil, and soil. Mostspecies were found in leaf litter, bark, and thecombination of leaf litter with soil (Table 2)

26

TABLE 1

Number of Habitat Samples Collected from Hono 0Nil Pali and Kui'a Natural Area Reserves

Sample Type Hono 0 Nil Pali" Kui'ab

Flower 3 3Leaves 14 10Bark 6 14Bark with moss 8 1Leaf litter 9 10Leaf litter with soil 9 3Soil 0 5

Total 49 41

"Three collections from upper elevation on Pihea, and onecollection from coastal area along Kalalau Trail.

b Two collections along Nu'alolo Trail.

was taken (Table 1). A litter sample consistedof leaves and small twigs lying on top of themineral soil showing signs of decomposition.A soil sample was the mineral soil under thelitter scooped to desired depth. Each littersample was taken with a lightweight plastictrowel, approximately three trowelsful (ap­proximately three to four cups [700-950 cm J]

dry weight). When mineral soil was in­advertently included in the litter during sam­pling, the category became "litter with soil."At Hono 0 Na Pali, moss and liverwortgrowth on trunks and twigs of 'Ohi'a waspredominant, making it difficult to samplebark without moss or liverworts. Few flowersamples were taken because they were located4 to 10m up in the canopy. Flowering seasonwas also a factor in the low number of flowersamples. Leaves were picked off singly fromtree branches. Efforts to standardize leaf sam­ples at 20-30 leaves per sample were main­tained during sampling. Samples wereindividually bagged in half-gallon (1893 cm3

)

plastic bags and labeled. Plastic bags were leftopen in the canvas carrier to provide a con­tinuous supply of oxygen to the live micro­organisms in the bag and to reduce con­densation inside the bag, deemed lethal to thetiny mites. Closing the bags was unnecessarybecause most mites, unlike ants and spring­tails, will not wander off from the substrate.Samples were transported the same day to theAcarology Laboratory at the University of

PACIFIC SCIENCE· January 2001

Hawai'i at Manoa for extraction. Mites wereextracted using Tullgren-Berlese funnels(Krantz 1978).

Extracted mites in 70% ethyl alcohol weresorted to suborders. Samples of each mor­phospecies were mounted on microslides withHoyer's mounting medium, then later driedin an oven for 2 weeks at 45-50°C and sealedwith Glyptal electric paint before examina­tion. A phase-contrast microscope (Wild) wasused in mite identification. Voucher speci­mens were kept at the J. Linsley GressittCenter for Research in Entomology of theBishop Museum, and the Acarology Labora­tory, University of Hawai'i at Manoa.

RESULTS AND DISCUSSION

Acarine Population, Composition, and Densities

The suborders Mesostigmata (Gamasida),Prostigmata (Actinedida), Astigmata (Acari­elida), and Oribatida (Cryptostigmata) wereall represented at Kui'a and Hono 0 Na PaliNatural Area Reserves. Approximately 33,400mites were extracted from 90 habitat sam­ples. Of the mite total, 67% (22,500) wereOribatida, 23% (7,711) Prostigmata, 19%(2,568) Mesostigmata, and 2% (592) Astig­mata. Of the 164 species found, 74 were ofthe suborder Prostigmata, 43 were Meso­stigmata, 4 were Astigmata, and 43 wereOribatida (Appendix). Irnrnatures composed35-40% of the entire mite collection.

MESOSTIGMATA: (Figure 2). A large andcosmopolitan suborder, mites in this grouphave adapted successfully to a remarkablywide range of habitats. Many are free-livingpredators in the soil, in litter, or on plants,where they prey on other small invertebrates(Krantz and Ainscough 1990). Many speciesare parasites of mammals, birds, reptiles, orinvertebrates (Strandtrnann and Wharton1958, Yunker 1973, Treat 1975). Mesostig­matic mites are generally well sclerotized.They range in size from 200 to 2500 /lm.

A total of 43 species of mesostigmaticmites was recorded on flowers, leaves, bark,leaf litter, leaf litter and soil, and soil. Mostspecies were found in leaf litter, bark, and thecombination of leaf litter with soil (Table 2)

Mite Communities Associated with 'Ohi'a . Swift and Goff 27

FIGURE 2. Suborder Mesostigmata, Pergamams sp., family Parasitidae. SEM photomicrograph by S.F.S.

in both reserves. Gamasiphis, undescribedsp. (Ologamasidae), Gamasholaspis gamasoides(Parholaspididae), and Veigaia planicola (Vei­gaiidae) were the predominant species in bothreserves. In Kui'a, Athiesella, undescribed sp.(Rhodacaridae), was numerous in all thesamples, but was not found at Hono 0 NilPali. Radovsky and Tenorio (1981) reportedthe ubiquitous distribution of this species insoil on the lower half of Mauna Loa transectin Hawai'i Island. In Kui'a, this species wasmore frequently represented in litter and barkthan in soil. Gamasolaelaps whartoni (Veigaii­dae) was found only on Hono 0 Nil Pali inlitter and litter with soil. The family Veigaii­dae was well represented with seven species inHono 0 Nil Pali: one species in the genusGamasolaelaps (G. whartoni) and six species inthe genus Veigaia (V. planicola, V. ce1-va, V.

uncata, and Veigaia species 1, 2, and 3. InKui'a, V. planicola and V. uncata were the twospecies represented.

Mes~stigmatic mite species found in dif­ferent 'Ohi'a habitats were mostly predaceous(except species in the genus Neocypholaelaps,family Ameroseiidae), feeding on small ar­thropods like Collembola and other smallmites. Their presence and abundance in bark,soil, and litter indicate the availability of food(prey) in these habitats as well as the opti­mum temperature and moisture they requirefor survival.

PROSTIGMATA: (Figure 3). Among theAcari, prostigmatic mites exhibit the great­est biological and morphological diversity.Members of this group are delicate, small,and, unlike the mesostigmatic mites, almostfree of sclerotization. They are free-living

Mite Communities Associated with 'Ohi'a . Swift and Goff 27

FIGURE 2. Suborder Mesostigmata, Pergamams sp., family Parasitidae. SEM photomicrograph by S.F.S.

in both reserves. Gamasiphis, undescribedsp. (Ologamasidae), Gamasholaspis gamasoides(Parholaspididae), and Veigaia planicola (Vei­gaiidae) were the predominant species in bothreserves. In Kui'a, Athiesella, undescribed sp.(Rhodacaridae), was numerous in all thesamples, but was not found at Hono 0 NilPali. Radovsky and Tenorio (1981) reportedthe ubiquitous distribution of this species insoil on the lower half of Mauna Loa transectin Hawai'i Island. In Kui'a, this species wasmore frequently represented in litter and barkthan in soil. Gamasolaelaps whartoni (Veigaii­dae) was found only on Hono 0 Nil Pali inlitter and litter with soil. The family Veigaii­dae was well represented with seven species inHono 0 Nil Pali: one species in the genusGamasolaelaps (G. whartoni) and six species inthe genus Veigaia (V. planicola, V. ce1va, V.

uncata, and Veigaia species 1, 2, and 3. InKui'a, V. planicola and V. uncata were the twospecies represented.

Mes~stigmatic mite species found in dif­ferent 'Ohi'a habitats were mostly predaceous(except species in the genus Neocypholaelaps,family Ameroseiidae), feeding on small ar­thropods like Collembola and other smallmites. Their presence and abundance in bark,soil, and litter indicate the availability of food(prey) in these habitats as well as the opti­mum temperature and moisture they requirefor survival.

PROSTIGMATA: (Figure 3). Among theAcari, prostigmatic mites exhibit the great­est biological and morphological diversity.Members of this group are delicate, small,and, unlike the mesostigmatic mites, almostfree of sclerotization. They are free-living

28 PACIFIC SCIENCE· January 2001

TABLE 2

Number of Mite Families, Genera, and Species for Each Sample Habitat at Hono 0 Na Pall andKui'a Natural Area Reserves

Hono 0 Na Pall Kui'a

Habitat Major Taxa Family Genus Species Family Genus Species

Flower Mesostigmata 4 4 5 2 3 3Prostigmata 5 5 5 5 5 5Astigmata 1 1 1 1 2 2Oribatida 1 1 1 0 0 0

Leaf Mesostigmata 3 6 10 3 3 3Prostigmata 8 13 15 8 8 12Astigmata 1 1 1 1 2 2Oribatida" 7 7 7 5 5 5

Bark Mesostigmata 6 14 15 6 9 10Prostigmata 10 17 24 12 20 28Astigmata 1 1 1 1 2 2Oribatida 7 7 7 7 7 7

Bark with moss Mesostigmata 6 10 14 6 7 7Prostigmata 12 18 25 8 10 14Astigmata 1 1 1 1 1 1Oribatida 13 13 13 10 10 10

Leaf litter Mesostigmata 9 12 16 9 13 16Prostigmata 15 26 34 15 22 30Astigmata 2 3 3 1 3 3Oribatida 22 23 23 15 15 15

Leaf litter with soil Mesostigmata 9 13 22 9 12 17Prostigmata 13 22 29 12 15 17Astigmata 1 2 2 1 2 2Oribatida 12 12 12 9 12 12

Soil" Mesostigmata 0 0 0 7 10 11Prostigmata 0 0 0 11 12 13Astigmata 0 0 0 1 2 2Oribatida 0 0 0 10 10 10

a Soil samples taken at Hono a Nii Pali mixed with leaf litter.

predators, fungivores/algaevores, and obligate sensilla (Rhagidiidae), Ereynetes sp. (Ereyne-plant feeders. A few species are parasitic on tidae), and three species in the superfamilyvertebrates and invertebrates and are vectors Raphignathoidea, Neog;nathus spectabilis (Cali-of diseases like scrub typhus. gonellidae), Favog;nathus distinctus and F. pic-

This group was represented by 74 species tus (Cryptognathidae), and Raphig;nathusin both reserves. Leaf litter habitat had the sp. (Raphignathidae) were widespread alonggreatest number of species at both Hono 0 Nu'alolo Trail and Hono 0 Nil Pali. Al-Nil Pali and Kui'a (34 and 30, respectively), though raphignathoid mites are primarilyfollowed by bark habitat (24 and 28, respec- predaceous, the Favog;nathus species are mosttively). Predaceous species such as Spinibdella likely nonpredaceous; instead they feed onthori (Bdellidae), Eupodes hawaiiensis and Eu- fungal hyphae and other microorganisms inpodes sip;moidensis (Eupodidae), Rhagidia longi- their habitat (Swift 1996). An undescribed

28 PACIFIC SCIENCE· January 2001

TABLE 2

Number of Mite Families, Genera, and Species for Each Sample Habitat at Hono 0 Nii Pali andKui'a Natural Area Reserves

Hono ONiiPali Kui'a

Habitat Major Taxa Family Genus Species Family Genus Species

Flower Mesostigmata 4 4 5 2 3 3Prostigmata 5 5 5 5 5 5Astigmata 1 1 1 1 2 2Oribatida 1 1 1 0 0 0

Leaf Mesostigmata 3 6 10 3 3 3Prostigmata 8 13 15 8 8 12Astigmata 1 1 1 1 2 2Oribatidaa 7 7 7 5 5 5

Bark Mesostigmata 6 14 15 6 9 10Prostigmata 10 17 24 12 20 28Astigmata 1 1 1 1 2 2Oribatida 7 7 7 7 7 7

Bark with moss Mesostigmata 6 10 14 6 7 7Prostigmata 12 18 25 8 10 14Astigmata 1 1 1 I 1 1Oribatida 13 13 13 10 10 10

Leaf litter Mesostigmata 9 12 16 9 13 16Prostigmata 15 26 34 15 22 30Astigmata 2 3 3 1 3 3Oribatida 22 23 23 15 15 15

Leaf litter with soil Mesostigmata 9 13 22 9 12 17Prostigmata 13 22 29 12 15 17Astigmata 1 2 2 1 2 2Oribatida 12 12 12 9 12 12

Soila Mesostigmata 0 0 0 7 10 11Prostigmata 0 0 0 11 12 13Astigmata 0 0 0 1 2 2Oribatida 0 0 0 10 10 10

a Soil samples taken at Hono a No Pali mixed with leaf litter.

predators, fungivores/algaevores, and obligate sensil/a (Rhagidiidae), Ereynetes sp. (Ereyne-plant feeders. A few species are parasitic on tidae), and three species in the superfamilyvertebrates and invertebrates and are vectors Raphignathoidea, Neog;nathus spectabilis (Cali-of diseases like scrub typhus. gonellidae), Favog;nathus distinctus and F. pic-

This group was represented by 74 species tus (Cryptognathidae), and Rtzphig;nathusin both reserves. Leaf litter habitat had the sp. (Raphignathidae) were widespread alonggreatest number of species at both Hono 0 Nu'alolo Trail and Hono 0 Nil Pall. Al-Nil Pali and Kui'a (34 and 30, respectively), though raphignathoid mites are primarilyfollowed by bark habitat (24 and 28, respec- predaceous, the Favog;nathus species are mosttively). Predaceous species such as Spinibdella likely nonpredaceous; instead they feed onthori (Bdellidae), Eupodes hawaiiensis and Eu- fungal hyphae and other microorganisms inpodes sigmoidensis (Eupodidae), Rhagidia longi- their habitat (Swift 1996). An undescribed

Mite Communities Associated with 'Ohi'a . Swift and Goff 29

FIGURE 3. Suborder Prostigmata, Stigmaeus, undescribed sp., family Stigmaeidae. SEM photomicrograph by S.F.S.

species of Spinibdella (Bdellidae) (one speci­men) was found on 'Ohi'a bark at 1100 m.The species diversity of the families Cunax­idae, Eupodidae, Stigmaeidae, and Tydeidaewas unexpectedly high (see Appendix).

ASTIGMATA: Tyrophagus putrescentiae(Acaridae) was the most widespread of theAstigmata. The species was represented on allsample habitats in the Nu'alolo Trail transectand the upper fringe of Hono 0 Na Pali, butnot in the low coastal area. The absence ofthis species and the entire group in the lowcoastal area was probably a sampling artifactbecause few samples were taken along KalalauTrail. It was also very dry when samples werecollected, suggesting that conditions wereunfavorable for the mite group, althoughAstigmata represents one-third of the mitefauna in California desert soil (Wallwork

1972). Three other astigmatic species col­lected from the reserves were Suidasia pon­tifica (Acaridae), Rhizoglyphus sp. (Acaridae),and Glycyphagus sp. (Glyeyphagidae).

The hypopus stage (phoretic deuto­nymph), which represents an adaptationunique to astigmatic mites (OConnor 1982),was the most common stage collected in thesamples. Attempts to identify them were notmade in the study.

ORIBATIDA: (Figure 4). Often called"beetle mites" because of their usuallyrounded shape and dark, hard, and shinyexoskeleton, oribatid mites are usually themost abundant and diverse arthropods intemperate forest soils and subtropical envi­ronments (Swift and Norton 1998). Highdensities surpassing 100,000/m2 are com­monly reported (Norton 1994). Arboreal

Mite Communities Associated with 'Ohi'a . Swift and Goff 29

FIGURE 3. Suborder Prostigmata, Stigmaeus, undescribed sp., family Stigmaeidae. SEM photomicrograph by S.F.S.

species of Spinibdella (Bdellidae) (one speci­men) was found on 'Ohi'a bark at 1100 m.The species diversity of the families Cunax­idae, Eupodidae, Stigmaeidae, and Tydeidaewas unexpectedly high (see Appendix).

ASTIGMATA: Tyrophagus putrescentiae(Acaridae) was the most widespread of theAstigmata. The species was represented on allsample habitats in the Nu'alolo Trail transectand the upper fringe of Hono 0 Na Pali, butnot in the low coastal area. The absence ofthis species and the entire group in the lowcoastal area was probably a sampling artifactbecause few samples were taken along KalalauTrail. It was also very dry when samples werecollected, suggesting that conditions wereunfavorable for the mite group, althoughAstigmata represents one-third of the mitefauna in California desert soil· (Wallwork

1972). Three other astigmatic species col­lected from the reserves were Suidasia pon­tifica (Acaridae), Rhizoglyphus sp. (Acaridae),and Glycyphagus sp. (Glyeyphagidae).

The hypopus stage (phoretic deuto­nymph), which represents an adaptationunique to astigmatic mites (OConnor 1982),was the most common stage collected in thesamples. Attempts to identify them were notmade in the study.

ORIBATIDA: (Figure 4). Often called"beetle mites" because of their usuallyrounded shape and dark, hard, and shinyexoskeleton, oribatid mites are usually themost abundant and diverse arthropods intemperate forest soils and subtropical envi­ronments (Swift and Norton 1998). Highdensities surpassing 100,000/m2 are com­monly reported (Norton 1994). Arboreal

30 PACIFIC SCIENCE· January 2001

FIGURE 4. Suborder Oribatida, Galumna sp., family Galumnatidae. SEM photomicrograph by S.F.S.

forms of oribatid mites are found on treebark, flowers, leaves, mosses, liverworts,lichens, and algae. Adult oribatid mite sizeranges from 150 ~m to 1.5 mm. Luxton(1972) categorized oribatid mites by feedinghabits as macrophytophages (those that feedstrictly on higher plant material after somedecomposition has taken place); microphy­tophages (those that feed on certain soilmicroflora such as fungi, algae, and bacteria);and panphytophages (those that utilizeboth microbial and higher plant materials).The latter includes perhaps the majority oforibatid mites.

Thirty-three families and 43 species ofOribatida were recorded on the Kui'a tran­sect and on both coastal and upper elevationsof Hono 0 Na Pali. Numerous species infamilies such as Galumnatidae and Schelori­batidae are reported only to generic level.

Several species known only from the nat­ural area reserves of Kaua'i were found onleaves and flowers of 'Ohi'a. Adhaesozetessp. (Adhaesozetidae) and Campbellobates sp.(<;;:ampbellobatidae) were reported from'Ohi'a leaves at Hono 0 Na Pali (Swift andNorton 1998). An undescribed genus in thefamily Oribatulidae was recorded fromflowers (Swift and Norton 1998). The familyPhthiracaridae had three species, Atropacarus(A.) striculus, Phthiracarus anonymus, and P.curiosus, recorded for Kaua'i and the Hawai­ian Islands by Niedbala (1998). Phthiracaruscuriosus Niedbala, 1998, an endemic species,was collected from Kui'a (Niedbala 1998). Adefinite introduction from along the Atlanticand Gulf coastal plains of southeastern NorthAmerica is the unusually unsclerotized Nehy­pochthonius porosus Norton & Metz, 1980,found for the first time in Kaua'i (Hono 0

30 PACIFIC SCIENCE· January 2001

FIGURE 4. Suborder Oribatida, Galumna sp., family Galumnatidae. SEM photomicrograph by S.F.S.

forms of oribatid mites are found on treebark, flowers, leaves, mosses, liverworts,lichens, and algae. Adult oribatid mite sizeranges from 150 ~m to 1.5 mm. Luxton(1972) categorized oribatid mites by feedinghabits as macrophytophages (those that feedstrictly on higher plant material after somedecomposition has taken place); microphy­tophages (those that feed on certain soilmicroflora such as fungi, algae, and bacteria);and panphytophages (those that utilizeboth microbial and higher plant materials).The latter includes perhaps the majority oforibatid mites.

Thirty-three families and 43 species ofOribatida were recorded on the Kui'a tran­sect and on both coastal and upper elevationsof Hono 0 Na Pali. Numerous species infamilies such as Galumnatidae and Schelori­batidae are reported only to generic level.

Several species known only from the nat­ural area reserves of Kaua'i were found onleaves and flowers of 'Ohi'a. Adhaesozetessp. (Adhaesozetidae) and Campbellobates sp.(<;;:ampbellobatidae) were reported from'Ohi'a leaves at Hono 0 Na Pali (Swift andNorton 1998). An undescribed genus in thefamily Oribatulidae was recorded fromflowers (Swift and Norton 1998). The familyPhthiracaridae had three species, Atropacarus(A.) striculus, Phthiracarus anonymus, and P.curiosus, recorded for Kaua'i and the Hawai­ian Islands by Niedbala (1998). Phthiracaruscuriosus Niedbala, 1998, an endemic species,was collected from Kui'a (Niedbala 1998). Adefinite introduction from along the Atlanticand Gulf coastal plains of southeastern NorthAmerica is the unusually unsclerotized Nehy­pochthonius porosus Norton & Metz, 1980,found for the first time in Kaua'i (Hono 0

Mite Communities Associated with 'Ohi'a . Swift and Goff 31

Na Pali) and in five other islands (O'ahu,Moloka'i, Lana'i, Maui, and Hawai'i) (Swiftand Norton 1998).

Distribution Trends

Mites were widely distributed on both re­serves. Different species were collected, pos­sibly related to elevation, at the three sitesalong Nu'alolo Trail on Kui'a Reserve. Threedistribution groups could be distinguished asfollows: group I, composed of species dis­tributed throughout the transect; group II,distributed from 975 to 1100 m; and groupIII, distributed from 762 to 974 m. In theMesostigmata, Gamasiphis, undescribed sp.,and Athiesella, undescribed sp., were two offive species found along the Nu'alolo tran­sect. Podocinum paeificum (Podocinidae) andPseudoparasitus anneetans (Laelapidae) weretwo species distributed above 975 m in soiland leaf litter. These two species were notfound at Hono 0 Na Pali Reserve. Asea aphi­doides (Ascidae), Veigaia uncata (Veigaiidae),Amblyseius sp. 1 (Phytoseiidae), and Pseudo­parasitus trineisus (Laelapidae) were in thegroup III category found in the lower ele­vations of Nu'alolo Trail.

In the Prostigmata, Eupodes sigmoidensisand E. hawaiiensis, Spinibdella thori, and Erey­netes sp. were four of nine species that werewidespread along the transect. Found in highelevations (group III) were two species ofCunaxidae (Cunaxa llf. veracruzana and Neo­cunaxoides sp. 2) and three species of Eupodi­dae (Eupodes, undescribed sp., Eupodes sp. 8,and Coeeeupodes llf. triscutatus). Two mitespecies in the supercohort Endeostigmata,Oehserehestes sp. (Oehserchestidae) and Bimi­ehaelia sp. 1 (Bimichaeliidae), were collectedin leaf litter and soil at Kui'a Reserve.

Habitats

Leaf litter, litter with soil, and bark habitatshad the most diverse mite fauna in the re­serves. The remarkable species diversity fromleaf litter was not unexpected. The presenceof tremendous number of microhabitats,availability of optimum environmental re­quirements, and decomposing plant parts and

other debris provide ideal growth environ­ment both for mites and for the microflorathat serve as mite food. The large, sclerotized,predatory mesostigmatic mites such as Perga­masus sp., Veigaia spp., and Pseudoparasitusspp. generally move freely chasing after preyanimals in the upper litter layer. The Vei­gaiidae, which are common in woodlandand forest soils, probably feed extensively onoribatid mites (Wallwork 1957) and Collem­bola. The smaller and more delicate Pro­stigmata in the families Cunaxidae, Bdellidae,and Stigmaeidae occur in the lower litterlayer near the mineral soil where most mem­bers in the suborder flourish (Wallwork 1972).The large number of predaceous Mesostig­mata and Prostigmata can be partly explainedby the presence of large population of preyavailable in the form of Collembola and othermites, particularly immature Oribatida (whichmake up approximately 30-40% of the prey).

The inner bark of trees provides habitatsfor a great variety of burrowing insects andtheir mite associates. In a review of foodhabits and host relationships of mites foundunder bark, Lindquist (1970) reported sym­biotic relationships between insects andmites. In view of this, the presence of scolytidbeetles occasionally seen in the bark samplesraises the question of whether the beetleshave_ any relationship to Acari groups foundin 'Ohi'a bark. Lindquist also observed thatcoexistent congeneric species have differenthabits and are not directly competitive withinthe bark habitat. Perhaps this is also true ofmite groups found in other habitats, whenseveral species of the same family are found inthe same microhabitat. In a study on apples,the mite fauna found on bark at the base ofthe trunk is apparently richer than on anyother part of the tree (Garman 1948). A largeportion of the fauna were oribatid mites,which could be partly explained by theirmovement from the soil and litter layer dur­ing the winter season or during adverse envi­ronmental conditions. Fauna of soil and barkcrevices overlaps to some extent, as observedin pseudoscorpions (Wallwork 1970).

Leaves and flowers had the least number ofmite associates. They also harbor an entirelydifferent mite group. Phytophagous Prostig-

Mite Communities Associated with 'Ohi'a . Swift and Goff 31

Na Pali) and in five other islands (O'ahu,Moloka'i, Lana'i, Maui, and Hawai'i) (Swiftand Norton 1998).

Distribution Trends

Mites were widely distributed on both re­serves. Different species were collected, pos­sibly related to elevation, at the three sitesalong Nu'alolo Trail on Kui'a Reserve. Threedistribution groups could be distinguished asfollows: group I, composed of species dis­tributed throughout the transect; group II,distributed from 975 to 1100 m; and groupIII, distributed from 762 to 974 m. In theMesostigmata, Gamasiphis, undescribed sp.,and Athiesella, undescribed sp., were two offive species found along the Nu'alolo tran­sect. Podocinum pacificum (Podocinidae) andPseudoparasitus annectans (Laelapidae) weretwo species distributed above 975 m in soiland leaf litter. These two species were notfound at Hono 0 Na Pali Reserve. Asca aphi­doides (Ascidae), Veigaia uncata (Veigaiidae),Amblyseius sp. 1 (Phytoseiidae), and Pseudo­parasitus trincisus (Laelapidae) were in thegroup III category found in the lower ele­vations of Nu'alolo Trail.

In the Prostigmata, Eupodes sigmoidensisand E. hawaiiensis, Spinibdella thori, and Erey­netes sp. were four of nine species that werewidespread along the transect. Found in highelevations (group III) were two species ofCunaxidae (Cunaxa nr. veracruzana and Neo­cunaxoides sp. 2) and three species of Eupodi­dae (Eupodes, undescribed sp., Eupodes sp. 8,and Cocceupodes nr. triscutatus). Two mitespecies in the supercohort Endeostigmata,Oehserchestes sp. (Oehserchestidae) and Bimi­chaelia sp. 1 (Bimichaeliidae), were collectedin leaf litter and soil at Kui'a Reserve.

Habitats

Leaf litter, litter with soil, and bark habitatshad the most diverse mite fauna in the re­serves. The remarkable species diversity fromleaf litter was not unexpected. The presenceof tremendous number of microhabitats,availability of optimum environmental re­quirements, and decomposing plant parts and

other debris provide ideal growth environ­ment both for mites and for the microflorathat serve as mite food. The large, sclerotized,predatory mesostigmatic mites such as Perga­masus sp., Veigaia spp., and Pseudoparasitusspp. generally move freely chasing after preyanimals in the upper litter layer. The Vei­gaiidae, which are common in woodlandand forest soils, probably feed extensively onoribatid mites (Wallwork 1957) and Collem­bola. The smaller and more delicate Pro­stigmata in the families Cunaxidae, Bdellidae,and Stigmaeidae occur in the lower litterlayer near the mineral soil where most mem­bers in the suborder flourish (Wallwork 1972).The large number of predaceous Mesostig­mata and Prostigmata can be partly explainedby the presence of large population of preyavailable in the form of Collembola and othermites, particularly immature Oribatida (whichmake up approximately 30-40% of the prey).

The inner bark of trees provides habitatsfor a great variety of burrowing insects andtheir mite associates. In a review of foodhabits and host relationships of mites foundunder bark, Lindquist (1970) reported sym­biotic relationships between insects andmites. In view of this, the presence of scolytidbeetles occasionally seen in the bark samplesraises the question of whether the beetleshave_ any relationship to Acari groups foundin 'Ohi'a bark. Lindquist also observed thatcoexistent congeneric species have differenthabits and are not directly competitive withinthe bark habitat. Perhaps this is also true ofmite groups found in other habitats, whenseveral species of the same family are found inthe same microhabitat. In a study on apples,the mite fauna found on bark at the base ofthe trunk is apparently richer than on anyother part of the tree (Garman 1948). A largeportion of the fauna were oribatid mites,which could be partly explained by theirmovement from the soil and litter layer dur­ing the winter season or during adverse envi­ronmental conditions. Fauna of soil and barkcrevices overlaps to some extent, as observedin pseudoscorpions (Wallwork 1970).

Leaves and flowers had the least number ofmite associates. They also harbor an entirelydifferent mite group. Phytophagous Prostig-

32

mata species such as Tuckerella sp. (Tuc­kerellidae) and Brevipalpus sp. (Tenuipalpi­dae) fed on living plant parts. The familyAmeroseiidae, with two species in the genusNeocypholaelaps, 'Yas reported only from leavesand flowers of 'Ohi'a. Members of this genusare known to live in flowers feeding on pollenand nectar using bees and butterflies forphoretic transport (Eickwort 1994). It is alsopossible that these mites use Hawaiian nativebirds feeding on nectars and pollen of 'Ohi'aflowers to move from flower to flower.Tyrophagus putrescentiae, Asca pineta, fourcunaxid species (Cunaxa sp., Cunaxoides sp.,Neocunaxoides andre, and Dactyloscirus iner­mis), three stigmaeid species (Eryngiopus sp. 1,Stigmaeus sp. 2, Stigmaeus sp. 3), two tydeidspecies (Lorryia sp. 1, Tydeus sp. 2), and ninefamilies in the Oribatida (Haplochthoniidae,Malaconothridae, N othridae, Adhaesozetidae,Campbellobatidae, Oribatulidae, Scheloriba­tidae, Ceratozetidae, and Galumnatidae) weresome of the taxa collected from leaves andflowers. The predaceous cunaxid and stig­maeid species probably feed on small insectsand their immatures and other small mites.The oribatid mite families collected fromleaves and flowers were examples of arborealpanphytophagous Oribatida, which feed onmicroflora found in plant parts and also onhigher plant materials (Luxton 1972, Walterand Behan-Pelletier 1999).

Residency Status/Endemism

Typically, island biotas exhibit a high degreeof endemism resulting from isolation andgeologic history. In Hawai'i, well over 90% ofplants and terrestrial arthropods are endemic(Simon et al. 1984). Currently, this level doesnot appear to be reflected in the acarinef~una. Of the 164 mite species associated with'Ohi'a, 19 are endemic (12%) and 28 adven­tive (17%). However, there are 117 species(71 %) for which we are currently unable todetermine their status.

CONCLUSIONS

The mite fauna associated with 'Ohi'a treesin the two Kaua'i natural area reserves was

PACIFIC SCIENCE· January 2001

clearly a rich one, both in number of speciesand number of individuals. The numbers arecomparable to numbers of mites found oncultivated apples. In a study of mite associatesin an apple orchard in southwestern Quebec,71 species were collected from ba~k, leaves,and litter (Forest et al. 1982). On 'Ohi'a, 127species were determined from bark, leaves,and litter. In Wisconsin, apple foliage is hometo 31 mostly pr~daceous mite species (Oat­man 1963); on 'Ohi'a, 33 species were foundon foliage, not all of them predaceous. Thespecies composition, however, was quite dif­ferent in the two studies.

A certain amount of overlap exists betweenlitter and mineral soil mite populations. Litterand mineral soil layers represent two distinctmicrohabitats for the soil fauna, and althoughthere is appreciable faunal movement be­tween the two, the presence of an associationof mite species characteristic of each suggeststhat some habitat separation is occurring(Wallwork 1972). A closer look at the verticaldistribution of certain focal taxa is necessaryto clear up this uncertainty and establish thevertical distribution pattern for Hawaiian rainforest soil and litter fauna. Although thereappeared to be a distribution trend amongsome mite groups on the Kui'a Reserve tran­sect, distribution overlap among taxa was alsoprevalent. Perhaps factors such as relativehumidity and temperature that accompanyelevation changes did not have any limitingeffect or the elevation difference in the studywas not enough to allow recognition of dif­ferences in the mite faunal communities.

Factors that influence the distribution ofpredatory Mesostigmata and Prostigmata arebasically unknown in the Islands. The distri­bution of prey animals as food sources is def­initely important for these mites to survivein particular habitats. The predominance ofCollembola probably plays an important roleas preferred food in the maintenance ofcommunities of predaceous mites inhabitingthe forest floor. From a sample of 'Ohi'a litterwith soil collected along the Pihea Trail onKaua'i, the Collembola-Acari compositionshowed 67% Collembola and 25% Acari(S.F.S., unpubl. data). It is probable that dis­tribution of predaceous mites is governed

32

mata species such as Tuckerella sp. (Tuc­kerellidae) and Brevipalpus sp. (Tenuipalpi­dae) fed on living plant parts. The familyAmeroseiidae, with two species in the genusNeorypholaelaps, ~as reported only from leavesand flowers of 'Ohi'a. Members of this genusare known to live in flowers feeding on pollenand nectar using bees and butterflies forphoretic transport (Eickwort 1994). It is alsopossible that these mites use Hawaiian nativebirds feeding on nectars and pollen of 'Ohi'aflowers to move from flower to flower.Tyrophagus putrescentiae, Asca pineta, fourcunaxid species (Cunaxa sp., Cunaxoides sp.,Neocunaxoides andre, and Dactyloscirus iner­mis), three stigmaeid species (Eryngiopus sp. 1,Stigmaeus sp. 2, Stigmaeus sp. 3), two tydeidspecies (Lorryia sp. 1, Tydeus sp. 2), and ninefamilies in the Oribatida (Haplochthoniidae,Malaconothridae, Nothridae, Adhaesozetidae,Campbellobatidae, Oribatulidae, Scheloriba­tidae, Ceratozetidae, and Galumnatidae) weresome of the taxa collected from leaves andflowers. The predaceous cunaxid and stig­maeid species probably feed on small insectsand their immatures and other small mites.The oribatid mite families collected fromleaves and flowers were examples of arborealpanphytophagous Oribatida, which feed onmicroflora found in plant parts and also onhigher plant materials (Luxton 1972, Walterand Behan-Pelletier 1999).

Residenry Status/Endemism

Typically, island biotas exhibit a high degreeof endemism resulting from isolation andgeologic history. In Hawai'i, well over 90% ofplants and terrestrial arthropods are endemic(Simon et al. 1984). Currently, this level doesnot appear to be reflected in the acarinef~una. Of the 164 mite species associated with'Ohi'a, 19 are endemic (12%) and 28 adven­tive (17 %). However, there are 117 species(71 %) for which we are currently unable todetermine their status.

CONCLUSIONS

The mite fauna associated with 'Ohi'a treesin the two Kaua'i natural area reserves was

PACIFIC SCIENCE· January 2001

clearly a rich one, both in number of speciesand number of individuals. The numbers arecomparable to numbers of mites found oncultivated apples. In a study of mite associatesin an apple orchard in southwestern Quebec,71 species were collected from ba.t::k, leaves,and litter (Forest et al. 1982). On 'Ohi'a, 127species were determined from bark, leaves,and litter. In Wisconsin, apple foliage is hometo 31 mostly pr~daceous mite species (Oat­man 1963); on 'Ohi'a, 33 species were foundon foliage, not all of them predaceous. Thespecies composition, however, was quite dif­ferent in the two studies.

A certain amount of overlap exists betweenlitter and mineral soil mite populations. Litterand mineral soil layers represent two distinctmicrohabitats for the soil fauna, and althoughthere is appreciable faunal movement be­tween the two, the presence of an associationof mite species characteristic of each suggeststhat some habitat separation is occurring(Wallwork 1972). A closer look at the verticaldistribution of certain focal taxa is necessaryto clear up this uncertainty and establish thevertical distribution pattern for Hawaiian rainforest soil and litter fauna. Although thereappeared to be a distribution trend amongsome mite groups on the Kui'a Reserve tran­sect, distribution overlap among taxa was alsoprevalent. Perhaps factors such as relativehumidity and temperature that accompanyelevation changes did not have any limitingeffect or the elevation difference in the studywas not enough to allow recognition of dif­ferences in the mite faunal communities.

Factors that influence the distribution ofpredatory Mesostigmata and Prostigmata arebasically unknown in the Islands. The distri­bution of prey animals as food sources is def­initely important for these mites to survivein particular habitats. The predominance ofCollembola probably plays an important roleas preferred food in the maintenance ofcommunities of predaceous mites inhabitingthe forest floor. From a sample of 'Ohi'a litterwith soil collected along the Pihea Trail onKaua'i, the Collembola-Acari compositionshowed 67% Collemboia and 25% Acari(S.F.S., unpubl. data). It is probable that dis­tribution of predaceous mites is governed

Mite Communities Associated with 'Ohi'a . Swift and Goff 33

more by the distribution of prey animals thanby the type of organic material or groundvegetation in the particular habitat.

Another dominant tree in Hawaiian forestc~mmunities, Koa (Acacia koa), together with'Ohi'a, provides the essential matrix of thenatural forest ecosystems in the HawaiianIslands (Mueller-Dombois 1981). Can we ex­pect similar mite species diversity on Koatrees? How do the findings from these tworeserves compare with data from reserves inother islands? Knowledge of Acari diversity iscrucial as loss of hosts and habitats speeds uptheir decimation and possible extinction. Al­though we doubt that tJ1eir loss will have animmediate impact on 'Ohi'a communities orany ecosystem in general, distinguishing en­demic mites is becoming even more difficultbecause of the continuous and frequent in­troduction of arthropods to the Islands.

Findings from this study show that futuremonitoring of Hawaiian ecosystems shouldinclude the acarine fauna, which can contrib­ute substantially to our knowledge of bothfauna and ecosystems that could be used inconservation assessments of forest reserves.

ACKNOWLEDGMENTS

Sincere thanks to Adam Asquith, Pam Cairy,and Todd Swift for their help in the fieldand to Karin Kami for her assistance in thelaboratory. Roy A. Norton, State Universityof New York, and Wocjiech Niedbala, A.Mickiewicz University, Poland, assisted in theidentification of the oribatid species. S.F.Sacknowledges use of facilities at Bishop Mu­seum during the study.

Literature Cited

Corn, C. A., and W. M. Heisey. 1973. Alti­tudinal ecotypes in Hawaiian Metrosideros.Island Ecosyst. IRP Tech. Rep. 18:1-19.

Department of Land and Natural Resources,State of Hawai'i. 1989a. Hono 0 Na PaliNatural Area Reserve Management Plan.31 pp.

---. 1989b. Kuia Natural Area ReserveManagement Plan. 33 pp.

Eickwort, G. C. 1994. Evolution of life-

history patterns of mites associated withbees. Pages 218-251 in M. Houck, ed.Mites: Ecological and evolutionary anal­yses of life-history patterns. Chapman &Hall, New York.

Evans, G. O. 1992. Principles of acarology.CAB International, Wallingford, UnitedKingdom.

Forest, J., ].-G. Pilon, and R. O. Paradis.1982. Acariens des vergers de pommiersdu sud-ouest du Quebec. Ann. Soc. En­tomol. Queb. 27:7-67.

Gagne, W. 1979. Canopy-associated arthro­pods in Acacia koa and Metrosideros treecommunities along an altitudinal transecton Hawaii Island. Pac. Insects 21:56-82.

Garman, P. 1948. Mite species from appletrees in Connecticut. Conn. Agric. Exp.Stn. Bull. 520:19-20.

Giambelluca, T. W., M. A. Nullet, and T. A.Schroeder. 1986. Rainfall atlas of Hawaii.Water Resources Research Center/Department ofMeteorology, University ofHawai'i at Manoa, State of Hawai'i,DLNR, Division of Water and Land De­velopment, Report R76. Honolulu.

Goff, M. L. 1987. A catalog of Acari of theHawaiian Islands. Univ. Hawaii Res. Ext.Ser.075:1-75.

Krantz, G. W. 1978. A manual of acarology.OSU Bookstore, Corvallis, Oregon. 500pp.

Krantz, G. W., and B. D. Ainscough. 1990.Acarina: Mesostigmata (Gamasida). Pages583-665 in D. L. Dindal, ed. Soil biologyguide.]. Wiley & Sons, New York.

Lindquist, E. E. 1970. Relationships betweenmites and insects in forest habitats. Can.Entomol. 102:978-984.

Luxton, M. 1972. Studies on the oribatidmites of the Danish beech wood soil. I.Nutritional biology. Pedobiologia 12:434­463.

Mueller-Dombois, D. 1981. Island ecosys­tems: What is unique about their ecology?Pages 485-501 in D. Mueller-Dombois,K. W. Bridges, and H. L. Carson, eds.Island ecosystems. Hutchinson Ross Pub­lishing Co., Woods Hole, Massachusetts.

---. 1985. Ohia dieback and protectionmanagement of the Hawaiian rainforest.

Mite Communities Associated with 'Ohi'a . Swift and Goff 33

more by the distribution of prey animals thanby the type of organic material or groundvegetation in the particular habitat.

Another dominant tree in Hawaiian forestc~mmunities, Koa (Acacia koa), together with'Ohi'a, provides the essential matrix of thenatural forest ecosystems in the HawaiianIslands (Mueller-Dombois 1981). Can we ex­pect similar mite species diversity on Koatrees? How do the findings from these tworeserves compare with data from reserves inother islands? Knowledge of Acari diversity iscrucial as loss of hosts and habitats speeds uptheir decimation and possible extinction. Al­though we doubt that tJ1eir loss will have animmediate impact on 'Ohi'a communities orany ecosystem in general, distinguishing en­demic mites is becoming even more difficultbecause of the continuous and frequent in­troduction of arthropods to the Islands.

Findings from this study show that futuremonitoring of Hawaiian ecosystems shouldinclude the acarine fauna, which can contrib­ute substantially to our knowledge of bothfauna and ecosystems that could be used inconservation assessments of forest reserves.

ACKNOWLEDGMENTS

Sincere thanks to Adam Asquith, Pam Cairy,and Todd Swift for their help in the fieldand to Karin Kami for her assistance in thelaboratory. Roy A. Norton, State Universityof New York, and Wocjiech Niedbala, A.Mickiewicz University, Poland, assisted in theidentification of the oribatid species. S.F.Sacknowledges use of facilities at Bishop Mu­seum during the study.

Literature Cited

Corn, C. A., and W. M. Heisey. 1973. Alti­tudinal ecotypes in Hawaiian Metrosideros.Island Ecosyst. IRP Tech. Rep. 18:1-19.

Department of Land and Natural Resources,State of Hawai'i. 1989a. Hono 0 Na PaliNatural Area Reserve Management Plan.31 pp.

---. 1989b. Kuia Natural Area ReserveManagement Plan. 33 pp.

Eickwort, G. C. 1994. Evolution of life-

history patterns of mites associated withbees. Pages 218-251 in M. Houck, ed.Mites: Ecological and evolutionary anal­yses of life-history patterns. Chapman &Hall, N ew York.

Evans, G. O. 1992. Principles of acarology.CAB International, Wallingford, UnitedKingdom.

Forest, ]., ].-G. Pilon, and R. O. Paradis.1982. Acariens des vergers de pommiersdu sud-ouest du Quebec. Ann. Soc. En­tomol. Queb. 27:7-67.

Gagne, W. 1979. Canopy-associated arthro­pods in Acacia koa and Metrosideros treecommunities along an altitudinal transecton Hawaii Island. Pac. Insects 21:56-82.

Garman, P. 1948. Mite species from appletrees in Connecticut. Conn. Agric. Exp.Stn. Bull. 520:19-20.

Giambelluca, T. W., M. A. Nullet, and T. A.Schroeder. 1986. Rainfall atlas of Hawaii.Water Resources Research Center!Department ofMeteorology, University ofHawai'i at Manoa, State of Hawai'i,DLNR, Division of Water and Land De­velopment, Report R76. Honolulu.

Goff, M. L. 1987. A catalog of Acari of theHawaiian Islands. Univ. Hawaii Res. Ext.Ser.075:1-75.

Krantz, G. W. 1978. A manual of acarology.OSU Bookstore, Corvallis, Oregon. 500pp.

Krantz, G. W., and B. D. Ainscough. 1990.Acarina: Mesostigmata (Gamasida). Pages583-665 in D. L. Dindal, ed. Soil biologyguide.]. Wiley & Sons, New York.

Lindquist, E. E. 1970. Relationships betweenmites and insects in forest habitats. Can.Entomol. 102:978-984.

Luxton, M. 1972. Studies on the oribatidmites of the Danish beech wood soil. I.Nutritional biology. Pedobiologia 12:434­463.

Mueller-Dombois, D. 1981. Island ecosys­tems: What is unique about their ecology?Pages 485-501 in D. Mueller-Dombois,K. W. Bridges, and H. L. Carson, eds.Island ecosystems. Hutchinson Ross Pub­lishing Co., Woods Hole, Massachusetts.

---. 1985. Ohia dieback and protectionmanagement of the Hawaiian rainforest.

34

Pages 403-417 in C. P. Stone and J. M.Scott, eds. Hawaii's terrestrial ecosystems:Preservation and management. Coopera­tive National Park Resources Studies Unit,University of Hawai'i at Manoa, Hono­lulu.

Niedbala, W. 1998. Euptyctima (Acari, Ori­batida) in the Hawaiian Islands. Proc.Hawaii. Entomol. Soc. 33:125-146.

Nishida, G. M. 1997. Hawaiian terrestrial ar­thropod checklist, 3d ed. Bishop MuseumPress, Honolulu.

Norton, R. A. 1994. Evolutionary aspectsof oribatid mite life-histories and conse­quences for the origin of Astigmata. Pages99-135 in M. Houck, ed. Mites: Ecologi­cal and evolutionary analysis of life-historypatterns. Chapman & Hall, New York.

Oatman, E. R. 1963. Mite species on applefoliage in Wisconsin. Adv. Acarol. 1:22­24.

OConnor, B. M. 1982. Acari:Astigmata. Pages 146-169 in S. B. Parker,ed. Synopsis and classification of living or­ganisms. Vol. 2. McGraw-Hill, New York.

Radovsky, F. J., and J. M. Tenorio. 1981. Soilarthropods. Pages 156-165 in D. Mueller­Dombois, K. W. Bridges, and H. L. Car­son, eds. Island ecosystems. HutchinsonRoss Publishing Co., Woods Hole, Mas­sachusetts.

Simon, C. M, W. C. Gagne, F. G. Howarth,and F. J. Radovsky. 1984. Hawai'i: A nat­ural entomological laboratory. Bull. En­tomol Soc. Am. 30:9-17.

PACIFIC SCIENCE· January 2001

Strandtmann, R. W., and G. W. Wharton.1958. Manual of mesostigmatid mites par­asitic on vertebrates. Institute of Acarol­ogy, University of Maryland, Contribution4. College Park, Maryland.

Swift, S. F. 1996. Hawaiian Raphignathoidea:Family Cryptognathidae (Acariformes:Prostigmata), with descriptions of threenew species of the genus Favognathus. Int.J. Acarol. 22:83-99.

Swift, S. F., and R. A. N. Norton. 1998. Pre­liminary report on oribatid mite (Acari:Oribatida) diversity in the HawaiianIslands. Bishop Museum Occas. Pap., ASpecial Publication of the Hawaii Biologi­cal Survey, No. 57.

Treat, A. E. 1975. Mites of moths and but­terflies. Cornell University Press, Ithaca,New York.

Wallwork, J. A. 1957. The Acarina of ahemlock-yellow birch forest floor. Ph.D.diss., University of Michigan, Ann Arbor.

---. 1970. Ecology of soil animals.McGraw-Hill Publishing Co., London.

---. 1972. Distribution patterns and pop­ulation dynamics of the micro-arthropodsof a desert soil in southern California. J.Anim. Ecol. 41:291-310.

Walter, D. E., and V. Behan-Pelletier. 1999.Mites in forest canopies: Filling the sizedistribution shortfall? Annu. Rev. Entomol.44:1-19.

Yunker, C. E. 1973. Mites. Pages 425-492 inR. J. Flynn, ed. Parasites of laboratoryanimals. Iowa State University Press, Ames.

Appendix

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii PaIi (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

Taxa Status' Reserve HabitatbEcological

Note'

Suborder MESOSTIGMATASuperfamily Ascoidea

Family AscidaeAsea aphidoides (Linnaeus, 1758)Asea pineta De Leon, 1967Blattisocius sp. 1Blattisocius sp. 2Cheiroseius sp.Gamase/lodes bie%r complex

Adv H K Ii Is b MiAdv H K Is b Ie f Mi? H Ii b bm Ie Mi? H b Ie f Mi? H Is b bm MiAdv H K Ii Is b bm Mi

34

Pages 403-417 in C. P. Stone and J. M.Scott, eds. Hawaii's terrestrial ecosystems:Preservation and management. Coopera­tive National Park Resources Studies Unit,University of Hawai'i at Manoa, Hono­lulu.

Niedbala, W. 1998. Euptyctima (Acari, Ori­batida) in the Hawaiian Islands. Proc.Hawaii. Entomol. Soc. 33:125-146.

Nishida, G. M. 1997. Hawaiian terrestrial ar­thropod checklist, 3d ed. Bishop MuseumPress, Honolulu.

Norton, R. A. 1994. Evolutionary aspectsof oribatid mite life-histories and conse­quences for the origin of Astigmata. Pages99-135 in M. Houck, ed. Mites: Ecologi­cal and evolutionary analysis of life-historypatterns. Chapman & Hall, New York.

Oatman, E. R. 1963. Mite species on applefoliage in Wisconsin. Adv. Acarol. 1:22­24.

OConnor, B. M. 1982. Acari:Astigmata. Pages 146-169 in S. B. Parker,ed. Synopsis and classification of living or­ganisms. Vol. 2. McGraw-Hill, New York.

Radovsky, F. J., and J. M. Tenorio. 1981. Soilarthropods. Pages 156-165 in D. Mueller­Dombois, K. W. Bridges, and H. L. Car­son, eds. Island ecosystems. HutchinsonRoss Publishing Co., Woods Hole, Mas­sachusetts.

Simon, C. M, W. C. Gagne, F. G. Howarth,and F. J. Radovsky. 1984. Hawai'i: A nat­ural entomological laboratory. Bull. En­tomol Soc. Am. 30:9-17.

PACIFIC SCIENCE· January 2001

Strandtmann, R. W., and G. W. Wharton.1958. Manual of mesostigmatid mites par­asitic on vertebrates. Institute of Acarol­ogy, University of Maryland, Contribution4. College Park, Maryland.

Swift, S. F. 1996. Hawaiian Raphignathoidea:Family Cryptognathidae (Acariformes:Prostigmata), with descriptions of threenew species of the genus Favognathus. Int.J. Acarol. 22:83-99.

Swift, S. F., and R. A. N. Norton. 1998. Pre­liminary report on oribatid mite (Acari:Oribatida) diversity in the HawaiianIslands. Bishop Museum Occas. Pap., ASpecial Publication of the Hawaii Biologi­cal Survey, No. 57.

Treat, A. E. 1975. Mites of moths and but­terflies. Cornell University Press, Ithaca,New York.

Wallwork, J. A. 1957. The Acarina of ahemlock-yellow birch forest floor. Ph.D.diss., University of Michigan, Ann Arbor.

---. 1970. Ecology of soil animals.McGraw-Hill Publishing Co., London.

---. 1972. Distribution patterns and pop­ulation dynamics of the micro-arthropodsof a desert soil in southern California. J.Anim. Ecol. 41:291-310.

Walter, D. E., and V. Behan-Pelletier. 1999.Mites in forest canopies: Filling the sizedistribution shortfall? Annu. Rev. Entomol.44:1-19.

Yunker, C. E. 1973. Mites. Pages 425-492 inR. J. Flynn, ed. Parasites of laboratoryanimals. Iowa State University Press, Ames.

Appendix

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

Taxa StatusU Reserve HabitatbEcological

Note'

Suborder MESOSTIGMATASuperfamily Ascoidea

Family AscidaeAsea aphidoides (Linnaeus, 1758)Asea pineta De Leon, 1967Blattisocius sp. 1Blattisocius sp. 2Cheiroseius sp.Gamase/lodes bie%r complex

Adv H K Ii Is b MiAdv H K Is b Ie f Mi? H Ii b bm Ie Mi? H b Ie f Mi? H Is b bm MiAdv H K Ii Is b bm Mi

Mite Communities Associated with 'Ohi'a . Swift and Goff 35

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii PaIi (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status' Reserve Habitatb Note'

Lasioseius sugawarai Ehara, 1964 Adv H bm Pr,MiLasioseius nr. sugawarai Ehara, 1964 End? H bm Ie Pr,MiLasioseius sp. 2 ? H bm Ie Pr,MiLasioseius sp. 3 ? H bm Ie Pr,Mi

Superfamily DermanyssoideaFamily Laelapidae

Hypoaspis (Geolaelaps) queenslandica(Womersley, 1956) Adv H b Pr,Mi

Pseudoparasitus (Gymnolaelaps)annectans (Womersley, 1955) Adv K Is bm Pr,Mi

Pseudoparasitus trincisus (Hunter,1966) Adv H K Ii Is b Pr,Mi

Superfamily EviphiidoideaFamily Parholaspididae

Gamasholaspis gamasoides (Berlese,1904) Adv H K Ii Is b bm Pr

Holaspina sp. 1 ? H bm PrHolaspina sp. 2 ? H Is PrHolaspina sp. 3 ? H Is b bm PrParholaspulus (Parholaspidellus)

maunaloaensis Tenorio & Marshall,1977 End K b Pr

Superfamily ParasitoideaFamily Parasitidae

Pergamasus sp. H K Is PrVulgarogamasus sp. 1 H K Ii Is b PrVulgarogamasus sp. 2 H K Ii Is PrVulgarogamasus sp. 3 H K Ii Is PrVulgarogamasus sp. 4 H Is Pr

Family VeigaiidaeVeigaia cerva (Kramer, 1987) Adv H Is PrVeigaia planicola (Berlese, 1892) Adv H K Ii Is b bm PrVeigaia uncata Farrier, 1957 Adv H K Ii Is PrVeigaia sp. 1 ? H Is PrVeigaia sp. 2 ? H Is PrVeigaia sp. 3 ? H Is PrGamasolaelaps whartoni (Farrier,

1957) Adv H Ii Is PrFamily Ameroseiidae

Neocypholaelaps nr. lindquisti Prasad,1968 End? H Ie f Mi

Neocypholaelaps sp. ? H K Ie f MiFamily Phytoseiidae

Amblyseius ovatus (Garman, 1958) Adv H Ii b PrAmblyseius nr. ovatus (Garman, 1958) End? H K Ii Is bm Ie f PrAmblyseius sp. 1 ? K Ii PrPhytoseius nr. hawaiiensis Prasad, 1968 End? K Ii PrPhytoseius sp. ? H Ie PrTyphlodromus nr. haramotoi Prasad,

1968 End? H b PrFamily Podocinidae

Podocinum pacificum (Berlese, 1896) Adv K Ii Is PrPodocinum sagax (Berlese, 1882) Adv H K Ii Is Pr

Mite Communities Associated with 'Ohi'a . Swift and Goff 35

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Na Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa StatusO Reserve Habitatb Note'

Lasioseius sugawarai Ehara, 1964 Adv H bm Pr,MiLasioseius ill. sugawarai Ehara, 1964 End? H bm Ie Pr,MiLasioseius sp. 2 ? H bm Ie Pr,MiLasioseius sp. 3 ? H bm Ie Pr,Mi

Superfamily DermanyssoideaFamily Laelapidae

Hypoaspis (Geolaelaps) queenslandica(Womersley, 1956) Adv H b Pr,Mi

Pseudoparasitus (Gymnolaelaps)annectans (Womersley, 1955) Adv K Is bm Pr,Mi

Pseudoparasitus trincisus (Hunter,1966) Adv H K Ii Is b Pr,Mi

Superfamily EviphiidoideaFamily Parholaspididae

Gamasholaspis gamasoides (Berlese,1904) Adv H K Ii Is b bm Pr

Holaspina sp. 1 ? H bm PrHolaspina sp. 2 ? H Is PrHolaspina sp. 3 ? H Is b bm PrParholaspulus (Parholaspidellus)

maunaloaensis Tenorio & Marshall,1977 End K b Pr

Superfamily ParasitoideaFamily Parasitidae

Pergamasus sp. H K Is PrVulgarogamasus sp. 1 H K Ii Is b PrVulgarogamasus sp. 2 H K Ii Is PrVulgarogamasus sp. 3 H K Ii Is PrVulgarogamasus sp. 4 H Is Pr

Family VeigaiidaeVeigaia cerua (Kramer, 1987) Adv H Is PrVeigaia planicola (Berlese, 1892) Adv H K Ii Is b bm PrVeigaia uncata Farrier, 1957 Adv H K Ii Is PrVeigaia sp. 1 ? H Is PrVeigaia sp. 2 ? H Is PrVeigaia sp. 3 ? H Is PrGamasolaelaps whartoni (Farrier,

1957) Adv H Ii Is PrFamily Ameroseiidae

Neorypholaelaps nr. lindquisti Prasad,1968 End? H Ie f Mi

Neorypholaelaps sp. ? H K Ie f MiFamily Phytoseiidae

Amblyseius ovatus (Garman, 1958) Adv H Ii b PrAmblyseius ill. ovatus (Garman, 1958) End? H K Ii Is bm Ie f PrAmblyseius sp. 1 ? K Ii PrPhytoseius ill. hawaiiensis Prasad, 1968 End? K Ii PrPhytoseius sp. ? H Ie PrTyphlodromus ill. haramotoi Prasad,

1968 End? H b PrFamily Podocinidae

Podocinum pacificum (Berlese, 1896) Adv K Ii Is PrPodccinum sagax (Berlese, 1882) Adv H K Ii Is Pr

36 PACIFIC SCIENCE· January 2001

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Statusfl Reserve Habitatb Note C

Superfamily PolyaspidoideaFamily Polyaspididae

Unidentified genus H K IiSuperfamily Rhodacaroidea

Family OlogamasidaeGamasiphis, undescribed sp. End H K Ii Is b bm Pr

Family RhodacaridaeAthiesella, undescribed sp. End K Ii Is b bm PrRhodacarus sp. ? H Ii bm Pr

Superfamily UropodoideaFamily Uropodidae

Fuscuropoda sp. H K Is Sa, PrSuborder PROSTIGMATA

Superfamily AnystoideaFamily Anystidae

Anystinae, unknown genus K Ii b Ie f PrErytbracarinae, unknown genus H K Ii Is b Ie f Pr

Superfamily BdelloideaFamily Bdellidae

Edelfa nihoaensis Swift & Goff, 1987 End H Ii Is b bm PrCyta kauaiensis Swift & Goff, 1987 End H Ii Is PrSpinibdella thori (Meyer & Ryke,

1959) Adv H K Ii Is b bm Ie PrSpinibdella, undescribed sp. End K b Pr

Family CunaxidaeCunaxa nr. veracruzana Baker &

Hoffmann, 1948 End? H K b bm PrCunaxa sp. ? H Ii b bm Ie PrCunaxoides sp. ? H K Ii b bm Ie PrDactyloscirus inermis (Baker &

Hoffmann, 1948) Adv H Ii Is bm Ie PrNeocunaxoides andre (Baker &

Hoffmann, 1948) Adv H K Ii Is b bm Ie PrNeocunaxoides sp. 1 ? H bm PrNeocunaxoides sp. 2 ? H K Ii bm PrPseudobonzia sp. ? H Ii Is PrPseudocunaxa sp ? H Is Pr

Superfamily CheyletoideaFamily Cheyletidae

Eueheyletia sp. H f PrSuperfamily Eupodoidea

Family EupodidaeCocceupodes triscutatus Strandtmann

& Prasse, 1977 Adv K Is PrCocceupodes nr. triscutatus

Strandtmann & Prasse, 1977 End? H K Ii Is b PrEupodes hawaiiensis (Strandtmann &

Goff,1978) End H K Ii b bm Ie PrEupodes sigmoidensis (Strandtmann &

Goff,1978) Adv H K Ii Is b bm Ie PrEupodes, undescribed sp. End H Ii Is bm PrEupodes sp. 2 ? H Is PrEupodes sp. 3 ? H Is Pr

36 PACIFIC SCIENCE· January 2001

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Statusfl Reserve Habitatb Note<

Superfamily PolyaspidoideaFamily Polyaspididae

Unidentified genus H K IiSuperfamily Rhodacaroidea

Family OlogamasidaeGamasiphis, undescribed sp. End H K Ii Is b bm Pr

Family RhodacaridaeAthiesella, undescribed sp. End K Ii Is b bm PrRhodacarus sp. ? H Ii bm Pr

Superfamily UropodoideaFamily Uropodidae

Fuscuropoda sp. H K Is Sa, PrSuborder PROSTIGMATA

Superfamily AnystoideaFamily Anystidae

Anystinae, unknown genus K Ii b Ie f PrErythracarinae, unknown genus H K Ii Is b Ie f Pr

Superfamily BdelloideaFamily Bdellidae

Bdella nihoaensis Swift & Goff, 1987 End H Ii Is b bm PrCyta kauaiensis Swift & Goff, 1987 End H Ii Is PrSpinibdella thori (Meyer & Ryke,

1959) Adv H K Ii Is b bm Ie PrSpinibdella, undescribed sp. End K b Pr

Family CunaxidaeCunaxa nr. veracruzana Baker &

Hoffmann, 1948 End? H K b bm PrCunaxa sp. ? H Ii b bm Ie PrCunaxoides sp. ? H K Ii b bm Ie PrDactyloscirus inermis (Baker &

Hoffmann, 1948) Adv H Ii Is bm Ie PrNeocunaxoides andre (Baker &

Hoffmann, 1948) Adv H K Ii Is b bm Ie PrNeocunaxoides sp. 1 ? H bm PrNeocunaxoides sp. 2 ? H K Ii bm PrPseudobonzia sp. ? H Ii Is PrPseudocunaxa sp ? H Is Pr

Superfamily CheyletoideaFamily Cheyletidae

Eueheyletia sp. H f PrSuperfamily Eupodoidea

Family EupodidaeCocceupodes triscutatus Strandtmann

& Prasse, 1977 Adv K Is PrCocceupodes nr. triscutatus

Strandtmann & Prasse, 1977 End? H K Ii Is b PrEupodes hawaiiensis (Strandtmann &

Goff,1978) End H K Ii b bm Ie PrEupodes sigmoidensis (Strandtmann &

Goff, 1978) Adv H K Ii Is b bm Ie PrEupodes, undescribed sp. End H Ii Is bm PrEupodes sp. 2 ? H Is PrEupodes sp. 3 ? H Is Pr

Mite Communities Associated with 'Ohi'a . Swift and Goff 37

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status" Reserve Habitatb Note<

Eupodes sp. 4 H K Is PrEupodes sp. 5 H Is PrEupodes sp. 6 K Ii PrEupodes sp. 7 K b PrEupodes sp. 8 K Ii PrProtereunetes sp. H K Ii Pr

Family RhagidiidaeCoccorhagidia nr. clavifrons

(G. Canestrini, 1886) End? K Is PrRhagidia longisensilla Shiba, 1969 Adv K Ii Is PrRhagidia sp. ? H Ii bm Pr

Superfamily ErythraeoideaFamily Erythraeidae

Hauptmannia sp. H bm PaSuperfamily Pygmephoroidea

Family PygmephoridaeBakerdania sp. H K Ii Is

Family ScutacaridaeImparipes sp. H K Ii Is

Superfamily RaphignathoideaFamily Caligonellidae

Neognathus spectabilis (Summers &Schlinger, 1955) Adv K b Pr

Family CryptognathidaeFavognathus distinctus Swift, 1996 End H K Ii b bm MiFavognathus pictus (Summers &

Chaudhri, 1965) Adv H K Ii bm MiFamily Raphignathidae Pr

RDphignathus sp. K Ii b PrFamily Stigmaeidae

Agistemus sp. H K b bm PrEryngiopus sp. 1 H Is b Ie PrEryngiopus sp. 2 H Is PrEustigmaeus segnis group H K Ii b PrLedermuelleriopsis plumosus

Willmann, 1951 Adv K Ii b PrStigmaeus sp. 2 ? H Ii b bm Ie PrStigmaeus sp. 3 ? H K Ii Is b Ie PrStigmaeus sp. 4 ? K b Pr

Superfamily TarsonemoideaFamily Tarsonemidae

Hemitarsonemus sp. H K Ii Is b bm Ph?Heterotarsonemus sp. H K Ii Is Ph?

Family TenuipalpidaeBrevipalpus sp. H K f Ph

Superfamily TetranychoideaFamily Tuckerellidae

Tuckerella sp. H K Ie f PhSuperfamily Tydeoidea

Family EreynetidaeEreynetes sp. H K Ii Is b bm Ie f Pa?

Family ParatydeidaeNeotydeus? sp. K b Mi

Mite Communities Associated with 'Ohi'a . Swift and Goff 37

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nil PaIi (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status· Reserve Habitatb Note'

Eupodes sp. 4 H K Is PrEupodes sp. 5 H Is PrEupodes sp. 6 K Ii PrEupodes sp. 7 K b PrEupodes sp. 8 K Ii PrProtereunetes sp. H K Ii Pr

Family RbagidiidaeCoccorhagidia nr. clavifrons

(G. Canestrini, 1886) End? K Is PrRhagidia longisensilla Shiba, 1969 Adv K Ii Is PrRhagidia sp. ? H Ii bm Pr

Superfamily ErythraeoideaFamily Erythraeidae

Hauptmannia sp. H bm PaSuperfamily Pygmephoroidea

Family PygmephoridaeBakerdania sp. H K Ii Is s

Family ScutacaridaeImparipes sp. H K Ii Is

Superfamily RaphignathoideaFamily Caligonellidae

Neognathus spectabilis (Summers &Schlinger, 1955) Adv K b Pr

Family CryptognathidaeFavognathus distinctus Swift, 1996 End H K Ii b bm MiFavognathus pictus (Summers &

Chaudhri, 1965) Adv H K Ii bm MiFamily Raphignathidae Pr

&lphignathus sp. K Ii b PrFamily Stigmaeidae

Agistemus sp. H K b bm PrEryngiopus sp. I H Is b Ie PrEryngiopus sp. 2 H Is PrEustigmaeus segnis group H K Ii b PrLedermuelleriopsis plumosus

Willmann, 1951 Adv K Ii b PrStigmaeus sp. 2 ? H Ii b bm Ie PrStigmaeus sp. 3 ? H K Ii Is b Ie PrStigmaeus sp. 4 ? K b Pr

Superfamily TarsonemoideaFamily Tarsonemidae

Hemitarsonemus sp. H K Ii Is b bm Ph?Heterotarsonemus sp. H K Ii Is Ph?

Family TenuipalpidaeBrevipalpus sp. H K f Ph

Superfamily TetranychoideaFamily Tuckerellidae

Tuckerella sp. H K Ie f PhSuperfamily Tydeoidea

Family EreynetidaeEreynetes sp. H K Ii Is b bm Ie f Pa?

Family ParatydeidaeNeotydeus? sp. K b Mi

38 PACIFIC SCIENCE· January 2001

Appenmx(Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Qhi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa StatusO Reserve Habitatb Note'

Family TydeidaeTydaeolus sp. H Ii b Pr?Lorryia sp. 1 H K Ii Ie Pr?Lorryia sp. 2 H K Ii Pr?Lorryia sp. 3 H Is Pr?Tydeus sp. 1 K b Pr?Tydeussp.2 H K Ii Is b bm Ie Pr?Tydeussp.3 K Ii b Pr?Tydeussp.4 H Ii Pr?Tydeus sp. 5 K b Pr?Tydeus sp. 6 H K Ii b bm Pr?Tydeussp.7 H b Pr?Tydeus sp. 8 H b Pr?Tydeussp.9 H K b Pr?

Supercohort ENDEOSTIGMATAFaruily Alicorhagiidae

Alicorhagia sp. H Ii MiFamily Birnichaeliidae

Bimichaelia sp. 1 K Ii Is MiBimichaelia sp. 2 H Is bm MiBimichaelia sp. 3 H Is MiGenus Asp. 1 H Is MiGenus A sp. 2 H bm Mi

Faruily NanorchestidaeNanorchestes sp. H Ii Is bm Mi

Faruily OehserchestidaeOehserchestes sp. K Mi

Suborder ASTIGMATASuperfaruily Acaroidea

Faruily AcaridaeRhizoglyphus sp. ? H K Ii Is s Sa,MiSuidasia pontifUa (Oudemans, 1905) Adv K Ii b Ie f Sa,MiTyrophagus putrescentiae (Schrank,

1781) Adv H K Ii Is b bm Ie f Sa,MiFaruily Glycyphagidae

Glycyphagus sp. H Ii Sa,MiSuborder ORIBATIDA

Superfaruily CtenacaroideaFaruily Aphelacaridae

Aphelacarus sp. H Ii bm Sa, MiSuperfaruily Hypochthonoidea

Faruily HypochthoniidaeEohypochthonius sp. H Is Sa, Mi

Superfaruily ProtoplophoroideaFaruily Haplochthoniidae

Haplochthonius sp. H K Ii Ie Sa,MiFaruily Cosmochthoniidae

Cosmochthonius spp. K Ii b Sa,MiSuperfaruily Brachychthonoidea

Faruily BrachychthoniidaeLiochthonius sp. K Is Sa,MiSellnickoethonius nr. zelawaiensis

(Sellnick, 1928) End? H K Is bm Sa, Mi

38 PACIFIC SCIENCE· January 2001

Appenmx(Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii PaIi (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa StatusO Reserve Habitatb Note'

Family TydeidaeTydaeolus sp. H Ii b Pr?Lorryia sp. 1 H K Ii Ie Pr?Lorryia sp. 2 H K Ii Pr?Lorryia sp. 3 H Is Pr?Tydeus sp. 1 K b Pr?Tydeus sp. 2 H K Ii Is b bm Ie Pr?Tydeus sp. 3 K Ii b Pr?Tydeus sp. 4 H Ii Pr?Tydeus sp. 5 K b Pr?Tydeus sp. 6 H K Ii b bm Pr?Tydeus sp. 7 H b Pr?Tyde'lts sp. 8 H b Pr?Tydeus sp. 9 H K b Pr?

Supercohort ENDEOSTIGMATAFamily Alicorhagiidae

Alicorhagia sp. H Ii MiFamily Bimichaeliidae

Bimichaelia sp. 1 K Ii Is MiBimichaelia sp. 2 H Is bm MiBimichaelia sp. 3 H Is MiGenus Asp. 1 H Is MiGenus A sp. 2 H bm Mi

Family NanorchestidaeNanorchestes sp. H Ii Is bm Mi

Family OehserchestidaeOehserchestes sp. K Mi

Suborder ASTIGMATASuperfamily Acaroidea

Family AcaridaeRhizoglyphus sp. ? H K Ii Is s Sa,MiSuidasia pontifica (Oudemans, 1905) Adv K Ii b Ie f Sa,MiTyrophagus putrescentiae (Schrank,

1781) Adv H K Ii Is b bm Ie f Sa,MiFamily Glycyphagidae

Glycyphagus sp. H Ii Sa,MiSuborder ORIBATIDA

Superfamily CtenacaroideaFamily Aphelacaridae

Aphelacarus sp. H Ii bm Sa, MiSuperfamily Hypochthonoidea

Family HypochthoniidaeEohypochthonius sp. H Is Sa,Mi

Superfamily ProtoplophoroideaFamily Haplochthoniidae

Haplochthonius sp. H K Ii Ie Sa,MiFamily Cosmochthoniidae

Cosmochthonius spp. K Ii b Sa, MiSuperfamily Brachychthonoidea

Family BrachychthoniidaeLiochthonius sp. K Is Sa,MiSellnickoethonius nr. zelawaiensis

(Sellnick, 1928) End? H K Is bm Sa, Mi

Mite Communities Associated with 'Ohi'a . Swift and Goff 39

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 N:i Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status' Reserve Habitatb Note'

Superfamily AtopochthonioideaFamily Pterochthoniidae

Pterochthonius angelus (Berlese, 1910) Adv H Ii Is Sa,MiSuperfamily Phthiracaroidea

Family PhthiracaridaeAtropacarus (A.) striculus (c. L. Koch,

1836) Adv K Is Sa,MiPhthiracarus anonymus (Grandjean,

1933) Adv K Is Sa,MiPhthiracarus curiosus Niedbala, 1998 End K Is Sa,Mi

Superfamily EuphthiracaroideaFamily Euphthiracaridae

Microtritia sp. K Ii Sa, MiFamily Oribotritiidae

Oribotritia sp. H K Ii Sa,MiSuperfamily Epilohmannioidea

Family EpilohmaniidaeEpilohmannia sp. K Ii Sa,Mi

Superfamily NehypochthonioideaFamily Nehypochthoniidae

Nehypochthonius porosus Norton &Metz,1980 Adv H Ii Sa, Mi

Superfamily CrotonioideaFamily Camisiidae

Platynothrus sp. H K Ii b bm Sa,MiFamily Malaconothridae

Malaconothrus sp. H K Ii b bm Ie Sa,MiFamily Nothridae

Nothrus sp. H K Ii Is b bm Ie Sa,MiFamily Trhypochthoniidae

Trhypochthoniellus sp. H Ii Sa,MiTrhypochthonius sp. H Ii b Sa,Mi

Superfamily NanhermannioideaFamily Nanhermanniidae

Masthermannia sp. H Ii Sa,MiNanhermannia sp. H Ii Sa,Mi

Superfamily HermannioideaFamily Hermanniidae

Phyllhermannia sp.? H Ii Sa,MiSuperfamily Liodoidea

Family LiodiidaeLiodes sp. H Ii Sa,Mi

Superfamily DamaeoideaFamily Damaeidae

Damaeus sp. H K Ii Is b bm Sa,MiSuperfamily Carabodoidea

Family CarabodidaeAustrocarabodes sp. H Ii Is Sa,Mi

Superfamily CepheidoideaFamily Cepheidae

"CepheZlS" pustulatZlS, undescribed g. End K Ii Sa,MiSuperfamily Amerobelboidea

Family EremobelbidaeEremobelba sp. H Ii Sa,Mi

Mite Communities Associated with 'Ohi'a . Swift and Goff 39

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status" Reserve Habitatb Note'

Superfamily AtopochthonioideaFamily Pterochthoniidae

Pterochthonius angelus (Berlese, 1910) Adv H Ii Is Sa,MiSuperfamily Phthiracaroidea

Family PhthiracaridaeAtropacarus (A.) striculus (c. L. Koch,

1836) Adv K Is Sa,MiPhthiracarus anonymus (Grandjean,

1933) Adv K Is Sa,MiPhthiracarus curiosus Niedbala, 1998 End K Is Sa,Mi

Superfamily EuphthiracaroideaFamily Euphthiracaridae

Microtritia sp. K Ii Sa, MiFamily Oribotritiidae

Oribotritia sp. H K Ii Sa,MiSuperfamily Epilohmannioidea

Family EpilohmaniidaeEpilohmannia sp. K Ii Sa,Mi

Superfamily NehypochthonioideaFamily Nehypochthoniidae

Nehypochthonius porosus Norton &Metz,1980 Adv H Ii Sa, Mi

Superfamily CrotonioideaFamily Camisiidae

Platynothrus sp. H K Ii b bm Sa,MiFamily Malaconothridae

Malaconothrus sp. H K Ii b bm Ie Sa,MiFamily Nothridae

Nothrus sp. H K Ii Is b bm Ie Sa,MiFamily Trhypochthoniidae

Trhypochthoniellus sp. H Ii Sa,MiTrhypochthonius sp. H Ii b Sa,Mi

Superfamily NanhermannioideaFamily Nanhermanniidae

Masthermannia sp. H Ii Sa,MiNanhermannia sp. H Ii Sa,Mi

Superfamily HermannioideaFamily Hermanniidae

Phyllhermannia sp.? H Ii Sa,MiSuperfamily Liodoidea

Family LiodiidaeLiodes sp. H Ii Sa,Mi

Superfamily DamaeoideaFamily Damaeidae

Damaeus sp. H K Ii Is b bm Sa,MiSuperfamily Carabodoidea

Family CarabodidaeAustrocarabodes sp. H Ii Is Sa,Mi

Superfamily CepheidoideaFamily Cepheidae

"Cepheus" pustulatus, undescribed g. End K Ii Sa,MiSuperfamily Amerobelboidea

Family EremobelbidaeEremobelba sp. H Ii Sa,Mi

40 PACIFIC SCIENCE . January 2001

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status· Reserve Habitatb Note'

Family TectocepheidaeTectocepheus sp. H K Ii bm Sa,Mi

Superfamily OppioideaFamily Oppiidae

Machuella sp. H K Is bm Sa,MiMicroppia sp. K Ii Sa,MiMultioppia sp. K Is Sa,Mi

Family QuadropiidaeQuadroppia sp. H Is Sa,Mi

Family SuctobelbidaeSuctobelbella sp. H Ii Is bm Sa, Mi

Superfamily LicneremaeoideaFamily Adhaesozetidae

Adhaesozetes sp. H Ie Sa,MiSuperfamily Oripodoidea

Family CampbellobatidaeCampbellobates acanthus hawaiiensis

Balogh, 1985 End H bm Sa,MiCampbellobates sp. ? H Ie Sa,Mi

Family HaplozetidaeRostrozetes sp. H K Ii Is Sa,Mi

Family OribatulidaeUndescribed genus End H f Sa,Mi

Family ScheloribatidaeScheloribates spp. H K Ii Is b bm Ie Sa,Mi

Superfamily CeratozetoideaFamily Ceratozetidae

Ceratozetes sp. H K bm Sa,MiFuscozetes sp. H K Ii Sa,Mi

Superfamily AchipterioideaFamily Tegoribatidae

Tegoribates sp. H Ii Sa,MiSuperfamily Galumnatoidea

Family GalumnatidaeGalumna spp. H K Ii Is b bm Ie Sa,Mi

n Adv, adventive; End, endemic; 1, unknown.b b, bark; bm, bark with moss; f, flower; Ie, leaves; Ii, leaf litter; Is, leaf litter with soil; s, soil.'Mi, microphytophage (fungivore, lichenovore, pollinovore, and phycophage of Luxton [1972]; Pa, parasite; Ph, phytophage

(feeding on living plant material); Pr, predator; Sa, saprophage (ingesting decaying plant or animal matter); 1, unknown ecological role.

40 PACIFIC SCIENCE . January 2001

Appendix (Continued)

Systematic Position, Status, and Ecological Notes of Mites Associated with 'Ohi'a in Hono 0 Nii Pali (H) andKui'a (K) Natural Area Reserves on Kaua'i Island

EcologicalTaxa Status" Reserve Habitatb Note'

Family TectocepheidaeTectocepheus sp. H K Ii bm Sa,Mi

Superfamily OppioideaFamily Oppiidae

Machuella sp. H K Is bm Sa,MiMicroppia sp. K Ii Sa, MiMultioppia sp. K Is Sa,Mi

Family QuadropiidaeQuadroppia sp. H Is Sa,Mi

Family SuctobelbidaeSuetobelbella sp. H Ii Is bm Sa, Mi

Superfamily LicneremaeoideaFamily Adhaesozetidae

Adhaesozetes sp. H Ie Sa,MiSuperfamily Oripodoidea

Family CampbellobatidaeCampbellobates acanthus hawaiiensis

Balogh, 1985 End H bm Sa,MiCampbellobates sp. ? H Ie Sa, Mi

Family HaplozetidaeRostrozetes sp. H K Ii Is Sa,Mi

Family OribatulidaeUndescribed genus End H Sa,Mi

Family ScheloribatidaeScheloribates spp. H K Ii Is b bm Ie Sa, Mi

Superfamily CeratozetoideaFamily Ceratozetidae

Ceratozetes sp. H K bm Sa,MiFuscozetes sp. H K Ii Sa,Mi

Superfamily AchipterioideaFamily Tegoribatidae

Tegoribates sp. H Ii Sa,MiSuperfamily Galumnatoidea

Family GalumnatidaeGalumna spp. H K Ii Is b bm Ie Sa,Mi

n Adv, adventive; End, endemic; 1, unknown.b b, bark; bm, bark with moss; f, flower; Ie, leaves; Ii, leaf litter; Is, leaf litter with soil; s, soil.'Mi, microphytophage (fungivore, lichenovore, pollinovore, and phycophage of Luxton [1972]; Pa, parasite; Ph, phytophage

(feeding on living plant material); Pr, predator; Sa, saprophage (ingesting decaying plant or animal matter); 1, unknown ecological role.