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Free-living spiders of the genus Ariamnes (Araneae, Theridiidae) in Hawaii.

ABSTRACT. This study examines species in a relatively diverse lineage of Ariamnes spiders in the Hawaiian archipelago, where they appear to have undergone adaptive radiation, with several species generally co-occurring at any one locality. The lineage was initially described by the single species A. corniger Simon 1900. The Hawaiian representatives of this lineage are characterized by variably elongate abdomens and are mostly free-living, at least as adults, although a number have been found to occur, at least facultatively, as kleptoparasites on the webs of other spiders. These spiders are all nocturnal. Their phylogenetic affinities with other representatives of the genus outside the Hawaiian Islands are unclear. Ten new species are described here: A. kahili, A. huinakolu, A. makue, A. uwepa, A. poele, A. melekalikimaka, A. alepeleke, A. laau, A. waikula, and A. hiwa. These species, together with the single described species Ariamnes corniger, include all known representatives of the lineage in the Hawaiian Islands. Different species occur in middle and high elevations and in wet and dry habitats.

Keywords: Descriptions, new species, taxonomy, Argyrodes, islands, Pacific Ocean

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The spider subfamily Argyrodinae (Theridiidae) comprises six genera, Argyrodes Simon 1864, Faiditus Keyserling 1884, Neospintharus Exline 1950, Ariamnes Thorell 1869, Rhomphaea L. Koch 1872, and Spheropistha Yaginuma 1957, the latter five genera having recently been removed from synonymy with Argyrodes (Agnarsson 2004). The original genus Argyrodes was considered to be distinct from two related genera, Rhomphaea and Ariamnes, based on differences in eye arrangement, clypeal modification, and relative length of metatarsi (Simon 1893). These characters were found not to separate groups reliably and so Rhomphaea and Ariamnes, together with Faiditus, Neospintharus, and Spheropistha, were incorporated into the composite genus Argyrodes (Exline & Levi 1962). However, a number of authors did not accept the synonymy of all of these genera, and Ariamnes and Rhomphaea (the two most distinct genera), in particular, continued to be used as generic descriptors (Forster & Forster 1999). Finally, arguing that the distinction between genera was quite clear-cut, Yoshida (2001) resurrected the genera Rhomphaea, Ariamnes, and Spheropistha. Moreover, recent work by Agnarsson (2004) also resurrected Faiditus and Neosphintharus on the basis that all of these genera are required to render the subfamily Argyrodinae monophyletic. The diversity represented by the current subfamily delimitation includes some of the most extraordinary behaviors known in spiders, in particular kleptoparasitism, a rather unusual form of species interaction in which one partner steals food from the other (Elgar 1993), and araneophagy, or preying on other spiders. Although the monophyly of the subfamily is now accepted (Agnarsson 2004), relationships between genera are poorly understood.

Until recently, taxonomic studies on native spiders in the Hawaiian Islands were derived almost entirely from the collections of R.C.L. Perkins (1913), based upon which Simon (1900) described 77 species endemic to the islands. Simon also recognized the speciose nature of one or a few genera in 4 spider families: Theridiidae, Salticidae, Thomisidae, and Tetragnathidae. More recently, taxonomic studies have been conducted on thomisids (Suman 1970), and the genera Tetragnatha Latreille 1804 (Tetragnathidae, Gillespie 1991, 1992, 1994, 2002, 2003) and Orsonwelles Hormiga 2002 (Linyphiidae, Hormiga 2002), and a number of recent studies have provided insights into patterns and processes underlying diversification in the genera Tetragnatha (Gillespie 2004; Blackledge & Gillespie 2004), Orsonwelles (Hormiga et al. 2003), and Havaika Proszynski 2002 (Salticidae, Arnedo & Gillespie 2006) in the Hawaiian Islands.

Ariamnes (Theridiidae) have been collected consistently in the Hawaiian Islands over the years, and Rivera (unpublished data) has studied Hawaiian members of the kleptoparasitic genus Argyrodes (initially known only by the single species Argyrodes hawaiiensis Simon 1900). Within the free-living Ariamnes, Simon (1900) recognized a single species, which he described as A. corniger Simon 1900. Since that time, there have been no further studies on this group.

Hawaiian representatives of the subfamily Argyrodinae were initially described in the two genera mentioned above, Argyrodes hawaiiensis and Ariamnes corniger. Argyrodes hawaiiensis is a short-bodied kleptoparasite; recent studies indicate that it belongs to a small clade of three species in Hawaii (Rivera unpublished data). Ariamnes corniger falls within a clade of multiple species that forms the focus of the current paper. All species show similarity in genitalic structure, yet considerable ecological, and associated morphological, differentiation. The Hawaiian lineage shows considerable morphological difference from other representatives of the genus Ariamnes as described in Agnarsson (2004). Rather, the eye arrangement might place it within the genus Neospintharus. However, the abdominal shape and genitalic structure (in particular that of the male palp) place it closer to Rhomphaea: The arrangement of the primary sclerites (embolus, conductor, median apophysis, theridiid tegular apophysis) of the male palp of Rhomphaea (Agnarsson 2004) is similar to that of the Hawaiian Ariamnes (Fig. 96). Simon (1900) commented on A. corniger as follows (in translation): "This remarkable species falls between the genera Ariamnes and Rhomphaea; it is related to the first by its narrow and vertical clypeus and the cylindrical femur and tibia of the palps; but the tibia of the forelegs is at least as long as the metatarsus, resembling more those of Rhomphaea. Its ocular area is provided, even in the female (only known sex), with a conical median tubercle, a character common to several Rhomphaea but up to now unknown in Ariamnes." Upon discovery of a male, Simon (1904) added "A. corniger, E Simon, is intermediate between the genera Ariamnes and Rhomphaea and could with almost as much justification be assigned to the one as to the other. The sexual characters also resemble those of unquestionable Argyrodes." Preliminary molecular data (M.A.J. Rivera and R.G. Gillespie, unpublished data) from specimens in the genera Argyrodes, Rhomphaea, Ariamnes, and Neospintharus indicate no tight affinity with any of these groups. In summary, the affinities of the long-bodied Hawaiian Argyrodinae are currently unknown; accordingly, we continue with the currently accepted generic assignment to the genus Ariamnes.

Ariamnes in the Hawaiian Islands are exclusively nocturnal and most frequently found free-living. However, it is not uncommon also to find them in kleptoparasitic association with a host spider, with members of the endemic Hawaiian genus Orsonwelles being the only known hosts to date. The following species have been found as kleptoparasites on the webs of different species of Orsonwelles: A. kahili, A. makue, A. melekalikimaka and A. laau. These species are also found as free-living individuals. It appears, therefore, that kleptoparasitism is facultative, unlike representatives of the Argyrodes and Cancellatus species groups in which it appears to be obligate, but perhaps more like members of the Trigonum (= Neospintharus) species group in which at least A. trigonum shows remarkable versatility (Cangialosi 1997). In terms of preycatching behavior, the Hawaiian Ariamnes have been observed to catch their spider prey with a net held in their back legs (M. Whitehouse, pers. comm.) similar to the manner reported for Rhomphaea (Whitehouse 1987) and Ariamnes attenuatus (Eberhard 1979).

The Hawaiian Ariamnes are generally elongate, though there is some variability in the relative elongation of the abdomen which can be almost triangular in side view (A. alepeleke, A. huinakolu and A. melekalikimaka) with no great extension behind the spinnerets, to thin with considerable elongation behind the spinnerets (all other species described). The legs are also quite variable in length, though all show the leg formula of 1423 characteristic of most argyrodines (Agnarsson 2004). For example, in A. corniger, the female leg I measures 24 mm (3.8 X length of abdomen), leg II 14 mm, leg III 8 mm, and leg IV 20 mm.

METHODS

Characters examined.--Morphological measurements taken were the same as those described in Exline & Levi (1962): the shape of the head and clypeus of the male, and the shape of the abdomen and genitalia. Species are described in order of geographic locality from Kauai (the oldest island) through Oahu, Molokai, Maui, and Hawaii (youngest island).

Terminology.--Unless indicated otherwise, all measurements are in mm. All holotypes and allotypes have been deposited in the Bishop Museum, Honolulu, Hawaii (BPBM) and all paratypes will be deposited in the Essig Museum of Entomology of the University of California, Berkeley, California (EMUC).

TAXONOMY

Family Theridiidae Sundevall 1833 Subfamily Argyrodinae Exline and Levi 1962

Genus Ariamnes Thorell 1869

Ariadne Doleschall 1857:410 (preoccupied by Ariadna Audouin 1826).

Ariamnes Thorell 1869:37 (replacement name for Ariadne Doleschall 1859).

Type species.--Ariadne flagellum Doleschall 1857, by monotypy.
KEY TO HAWAIIAN LONG-BODIED ARIAMNES SPECIES

 1. Males 2

 Females 12

 2. Embolus with large, looping, distal undulation
 (Figs. 89-91, 94, 100) 3

 Embolus straight or with very shallow
 undulations (Figs. 92, 93, 95-99) 7

 3. Abdomen short, extension behind spinnerets
 shorter than section in front; high, height
 approximately 50% length; shape triangular
 (Fig. 11) Argyrodes
 huinakolu new species

 Abdomen longer, extension behind spinnerets
 much greater than section in front (Fig. 3) 4

 4. Abdomen much higher at spinnerets, height
 about a third of length (Fig. 43);
 rectangular area separating cephalic and
 clypeal processes (Fig. 42); conductor less
 projecting, much shorter than embolus (Fig.
 44) Argyrodes
 melekalikimaka
 new species

 Maximum abdominal height [less than or equal
 to] 20% length; "U" or "V" shaped area
 separating cephalic and clypeal processes;
 conductor as long or longer than embolus 5

 5. Conductor long, thread-like, distally
 projecting, much longer than embolus and
 not closely aligned with it at tip (Figs.
 84, 100) Argyrodes hiwa
 new species

 Conductor and embolus similar in length 6

 6. Cephalic process short (13% carapace
 length), thick, straight, not distended
 at tip (Fig. 18) Argyrodes makue
 new species

 Cephalic process longer (16% carapace
 length), thinner stalk, curved over and
 distended at tip (Fig. 2) Argyrodes kahili
 new species

 7. Embolus with shallow undulations
 (Figs. 92, 95-97); abdomen covered
 with densely packed silver blocks 8

 Embolus without noticeable
 undulations, at most shallow,
 screw-like twists (Figs. 93, 98,
 99) 10

 8. Abdomen length > 2.5 x carapace
 length (Fig. 27); proximal and
 distal undulations of embolus of
 similar height with tip angled
 parallel to conductor (Figs. 28, 92) Argyrodes uwepa
 new species

 Abdomen length < 2.3 x carapace
 length; proximal undulation of
 embolus of greater height than
 distal, and/or tip angled away
 from conductor 9

 9. Opening between cephalic and
 clypeal process greatest at distal
 end with cephalic and clypeal setae
 angled towards each other (Fig. 58);
 total length approximately 5.0 mm Argyrodes
 alepeleke new species

 Opening between cephalic and
 clypeal process similar at distal
 and proximal end with cephalic and
 clypeal setae approximately at right
 angles to each other (Fig. 50); total
 length approximately 6.5 mm Argyrodes corniger
 Simon

10. Cephalic process very long
 (approximately 20% length of
 carapace), curved over and not noticeably
 distended at tip (Fig. 74); abdomen
 densely covered in silver blocks; carapace
 pale with dark lines on either side of
 midline Argyrodes waikula
 new species

 Cephalic process short (approximately
 15% length of carapace), straight or very
 slightly curved over; abdomen dark with
 variable scattered silver blocks; carapace
 dark with lighter midline 11

11. Embolus almost straight (Figs. 68, 98);
 abdomen with silver blotches on black
 blotches on dorsal surface as well as on
 sides Argyrodes laau
 new species

 Twists in embolus quite pronounced
 (Figs. 36, 93); abdomen very dark black
 on dorsal surface Argyrodes poele
 new species

12. Abdominal height > 45% length (Fig. 15,
 47, 63) 13

 Abdominal height approximately < 45%
 length (Fig. 55) 15

13. Abdominal height approximately 60-65%
 length, short, approximately 2.5-3.5 mm
 (Fig. 15); cephalic and clypeal processes
 similar in size (Fig. 14); coloration of
 abdomen in dorsal area very dark, tan below
 with numerous silver and black flecks Argyrodes
 huinakolu new
 species

 Abdominal height approximately 45-55%
 length, short, approximately 3.5-6.0
 mm; cephalic process > 3 x size of
 clypeal process (Figs. 46, 62). Abdomen
 fairly uniformly colored with silver
 blocks 14

14. Cephalic process projecting well in
 front of AMEs by almost width of AMEs
 (Fig. 46); abdomen brown fairly loosely
 covered with silver blocks Argyrodes
 melekalikimaka new
 species

 Cephalic process barely projecting at
 all (by < 50% width of AMEs) in front
 of AMEs (Fig. 62); abdomen densely
 covered with silver blocks, sometimes
 with red superimposed Argyrodes alepeleke
 new species

15. Abdomen much higher at spinnerets than
 half way between spinnerets and
 posterior end (2.3-3.5 x as high)
 (Figs. 55, 79); abdomen covered all
 over with large silver blocks 16 Abdomen
 only slightly higher at spinnerets than
 half way between spinnerets and posterior
 end (1.4-2.0 as high) (Fig. 7) 17

16. Cephalic process projecting well in front
 of AMEs by almost width of AMEs (Fig. 54);
 duct of vulva loosely coiled (Fig. 56) Argyrodes
 corniger Simon

 Cephalic process less projecting (by < 50%
 width of AMEs) in front of AMEs (Fig. 78);
 duct of vulva in a tight loop (Fig. 80) Argyrodes
 waikula new species

17. Abdomen very long and thin, height only
 approx 15% length (Figs. 7, 31, 87) 18

 Abdomen height > 20% length (Figs. 23,
 39, 71) 20

18. Spinnerets close to anterior, > 75% of
 abdomen behind spinnerets (Fig. 31);
 abdomen covered all over with blocks
 of silver Argyrodes uwepa
 new species

 Spinnerets not as close to anterior, <
 70% of abdomen behind spinnerets (Figs.
 7, 87); abdomen with some black flecks 19

19. Posterior eye row straight (Fig. 5);
 abdomen covered all over with small
 silver blocks and black flecks; epigynal
 aperture almost round (Fig. 101) Argyrodes kahili
 new species

 Posterior margin of posterior eye row
 procurved (Fig. 85); abdomen with dark
 dorsal area, remainder covered all over
 with small silver blocks and black flecks;
 epigynal aperture transverse ellipse (Fig.
 111) Argyrodes hiwa
 new species

20. Spinnerets close to anterior,
 approximately 75% of abdomen behind
 spinnerets (Fig. 71); abdomen dark
 streaked with silver blotches Argyrodes laau
 new species

 Spinnerets further down abdomen, < 65%
 of abdomen behind spinnerets 21

21. Cephalic process pronounced and
 pointed (Fig. 22); abdomen dark on
 dorsal surface, packed with silver
 blocks with black flecks below Argyrodes makue
 new species

 Cephalic process short and rounded
 (Fig. 38); abdomen silver and black
 but very dark Argyrodes poele
 new species


[FIGURES 1-8 OMITTED]

[FIGURES 9-16 OMITTED]

[FIGURES 17-24 OMITTED]

[FIGURES 25-32 OMITTED]

[FIGURES 33-40 OMITTED]

Ariamnes kahili new species Figs. 1-8, 89, 101, 112

Type specimens.--USA: Hawaiian Islands: holotype male, allotype female, Kauai, Mt. Kahili, 670 m elev., 21.99[degrees]N, 159.49[degrees]W, 20 December 1997, A. Asquith (BPBM); 1 paratype male, Kauai, Mohihi Ditch, 1067 m elev., 22.12[degrees]N, 159.63[degrees]W, 19 February 1991, R. Gillespie (EMUC); 1 paratype female, Kokee, 1097 m elev., 22.13[degrees]N, 159.65[degrees]W, in vegetation, 26 June 1964, D.E. Hardy, (BPBM).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the type locality of the species on Kauai.

Diagnosis.--Ariamnes kahili can be distinguished from most other species based on the deep undulations of the embolus of the male palp, projecting above the cymbium in parallel with the conductor (Fig. 4, 89). Similar palps are found in A. huinakolu and A. makue. Compared to A. makue, the cephalic process in A. huinakolu is longer and distended at tip; the abdomen is much longer (2.3 x length of carapace, compared to approximately 1.4 x in A. huinakolu and 1.7 x in A. makue).

Description.--Holotype male (Figs. 1-4, 89): Carapace dark brown with light band running longitudinally down middle. Abdomen dull light brown with numerous very loosely packed small silver blocks and black streaks running longitudinally in a line along side from front, turning into scattered black specks behind spinnerets (Fig. 3). Posterior eyes separated by 1.5 x diameter (Fig. 1). Cephalic process 16% length of carapace, tip distended with numerous setae (Fig. 2). Clypeal process 80% length of cephalic process, narrowing at tip where there are several long setae. Abdomen uniformly elongate, only ~1.5 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 3). Abdomen 5.0 mm length, 70% of this posterior to spinnerets. Carapace 2.1 mm length, 1.1 mm width. Palp (Figs. 4, 89): Embolus with deep undulations, projecting well above cymbium, closely aligned with conductor along length; conductor (membranous) same length as embolus.

Allotype female (Figs. 5-8, 101, 112): Color similar to male. Posterior eyes separated by 1.9 x diameter (Fig. 5). Cephalic process 5% length of carapace, clypeal process indistinct, rounded area (Fig. 6). Abdomen uniformly elongate, only ~1.5 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 7). Abdomen 8.6 mm length, 70% of this posterior to spinnerets. Carapace 2.1 mm length, 1.1 mm width. Vulva (Fig. 8): Receptacles spherical, ducts looped in figure-eight. In external view, epigynal area almost round with larger dark ring around central area (Fig. 101) with hooked scape (Fig. 112).

Variation.--Total length varies 6.5-7.1 mm in males, 10.7-11.3 mm in females, due to variable elongation in the abdomen. Color in life is quite variable; most individuals collected had a bright iridescent gold abdomen, but some had extensive dark markings.

Natural history.--Ariamnes kahili is found in wet forest habitats on the island of Kauai. The species appears to be largely free-living, but has been found on the webs of Orsonwelles.

Ariamnes huinakolu new species Figs. 9-16, 90, 102, 113

Type specimens.--USA: Hawaiian Islands: holotype male, Kauai, Makaleha Mountains, 950 m elev., 22.13[degrees]N, 159.42[degrees]W, 28 April 2000, M. Arnedo, G. Hormiga (BPBM); allotype female, Kauai, Makaleha Mountains, 950 m elev., 22.13[degrees]N, 159.42[degrees]W, 19 April 1999, M. Arnedo, G. Hormiga, N. Scharff (BPBM); 1 paratype male, Kauai, Kahili Mountain, 920 m elev., 21.98[degrees]N, 159.50[degrees]W, March 1998 (EMUC); 1 paratype female, Haupu, 700 m elev., 21.93[degrees]N, 159.40[degrees]W, 30 April 2000, M. Arnedo, G. Hormiga, A. Asquith (EMUC).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian word "huinakolu" meaning triangle, and refers to the almost equilateral triangular shape to the abdomen.

Diagnosis.--Ariamnes huinakolu can be readily distinguished from all other species based on the very high and short abdomen (height approximately 52% length in males, 63% in females) (Figs. 11, 15).

Description.--Holotype male (Figs. 9-2, 90): Carapace dark brown with creamy elongate oval area running longitudinally down middle. Abdomen dark dorsally, tan below midline with silver flecks at border between dark and tan areas (Fig. 11). Lower tan area with black lines and flecks together with sparse silver blocks. Posterior eyes separated by 1.6 x diameter (Fig. 9). Cephalic process 15% length of carapace, not noticeably distended, with numerous setae at tip (Fig. 10). Clypeal process 64% length of cephalic process. Abdomen short, triangular from side. uniformly elongate, only ~1.5 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 11). Abdomen 2.2 mm length, 40% of this posterior to spinnerets. Carapace 1.5 mm length, 0.9 mm width. Palp (Fig. 12, 90): Embolus with deep undulations, projecting well above cymbium, parallel to conductor along length; conductor (membranous) thin, same length as embolus.

Allotype female (Figs. 13-16, 102, 113): Color similar to male. Posterior eyes separated by 2.9 x diameter (Fig. 13). Cephalic process 3% length of carapace, clypeal process similar in size and shape (Fig. 14). Abdomen short and high, ~1.5 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 15). Abdomen 2.7 mm length, 30% of this posterior to spinnerets. Carapace 1.7 mm length, 1.1 mm width. Vulva (Fig. 16): Receptacles spherical, ducts looped in figure-eight. In external view, epigynal area almost round with dark patch surrounding anterior area (Fig. 101) and with small pointed scape (Fig. 112).

Variation.--Total length varies 3.4-3.7 mm in males, 3.5-4.4 mm in females. Specimens have been found from a single locality only, and little is known as to how much variation exists.

Natural history.--Ariamnes huinakolu has been found mainly on the summit of the Makleha Mountains on Kauai. It is largely restricted to this area, a site of wet forest on the summit of an isolated mountain. Kleptoparasitic tendencies are unknown.

Ariamnes makue new species Figs. 17-24, 91, 103, 114

Type specimens.--USA: Hawaiian Islands: holotype male, Oahu, Waianae Mountains, Puu Kaua, South Gulch, Honouliuli Preserve, 580 m elev., 21.45[degrees]N, 158.10[degrees]W, 6 August 1998, M. Arnedo, C. Ewing (BPBM); allotype female, Oahu, Waianae Mountains, Palikea, Honouliuli Preserve, 610 m elev., 21.43[degrees]N, 158.83[degrees]W, 20 August 1995, R. Gillespie, G. Roderick (BPBM); 1 paratype female, Oahu, Waianae Mountains, Puu Kaua, South Gulch, Honouliuli Preserve, 580 m elev., 21.45[degrees]N, 158.10[degrees]W, 6 August 1998, M. Arnedo, C. Ewing (EMUC); 1 paratype male, Oahu, Mount Kaala, 550 m elev., 21.52[degrees]N, 158.15[degrees]W, on large webs of Orsonwelles sp. (Linyphiidae), 23 September 1996, M. Rivera & G. Roberts (EMUC).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian word "maku'e" meaning a dark color, and refers to the very dark color of the abdomen of this species.

Diagnosis.--Ariamnes makue can be distinguished from all other species based on the deep undulations of the embolus (Fig. 91) and the relatively short posterior end of the abdomen (Figs. 19, 23).

Description.--Holotype male (Figs. 17-20, 91): Carapace dark with lighter longitudinal band running down midline. Abdomen with dark band running along dorsal surface (Fig. 19). Brownish below with loosely packed silver blocks and black longitudinal flecks along sides. Posterior eyes separated by 1.6 x diameter (Fig. 17). Cephalic process 13% length of carapace, thick but not noticeably distended, with numerous setae at tip (Fig. 18). Clypeal process 84% length of cephalic process. Abdomen uniformly elongate, only ~1.1 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 19). Abdomen 3.6 mm length, 60% of this posterior to spinnerets. Carapace 2.1 mm length, 1.0 mm width. Palp (Figs. 20, 91): Embolus with very deep distal undulation, projecting above cymbium, parallel to conductor along length; conductor same length as embolus.

Allotype female (Figs. 21-24, 103, 114): Color similar to male. Posterior eyes separated by 2.3 x diameter (Fig. 21). Cephalic process 5% length of carapace, similar in shape to (but 3.7 x as long as) clypeal process (Fig. 22). Abdomen elongate, ~1.8 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 23). Abdomen 4.6 mm length, 63% of this posterior to spinnerets. Carapace 1.7 mm length, 0.9 mm width. Vulva (Fig. 24): Receptacles elongate-oval, ducts looped in figure-eight. In external view, receptacles visible, epigynal area indistinct (Fig. 103) and with scape curved over (Fig. 114).

Remarks.--A male specimen of this species was initially described as a co-type of A. corniger. Simon (1904) stated (in translation): "The frontal tubercle already indicated in the female is much longer, resembling that of Rhomphaea projiciens Cambr., from Central America, though it is less dilated and not at all pilous at the end; the second lower tubercle, about half the size, is pointed and directed upward, reminiscent of Ariamnes bicornis Cambr., from Brazil." This specimen was examined, and the genitalia found to be very similar to that of A. makue. However, the frontal tubercle is considerably longer than any of the Hawaiian Ariamnes examined. Without any additional material, it is impossible to determine whether this specimen warrants a separate description.

Variation.--Total length varies 5.3-5.7 mm in males, 5.8-6.3 mm in females, due to variable elongation in the abdomen. Color in life is very dark with little apparent variability.

Natural history.--Ariamnes makue is found in mesic forest habitats on the island of Oahu. It has been found on the webs of Orsonwelles but can also be free-living; however, microhabitat affinities are unknown.

Ariamnes uwepa new species Figs. 25-32, 92, 104, 115

Type specimens.--USA: Hawaiian Islands: holotype male, Oahu, Koolau Mountains, Poamoho Trail, 600 m elev., 21.52[degrees]N, 157.95[degrees]W, 18 January 1980, W.C. Gagne (BPBM); allotype female, Oahu, Koolau Mountains, Poamoho Trail, 600 m elev., 21.53[degrees]N, 157.95[degrees]W, 31 January 1999, R. Gillespie, G. Roderick (BPBM); 1 paratype male, Oahu, Waianae Mountains, Kamaileunu Ridge, north of Puu Kawiwi, 760 m elev., 21.48[degrees]N, 158.18[degrees]W, 19 October 1975, F.G. Howarth (BPBM); 1 paratype female, Oahu, Waianae Mountains, summer 1966, W. Hay (BPBM).

Etymology.--The specific epithet, regarded as a noun in apposition, comes from the Hawaiian word "uwepa" meaning whip, and refers to the common name often used for the genus Ariamnes, with reference to the very elongate and flexible abdomen of these spiders.

Diagnosis.--Ariamnes uwepa can be distinguished from other species based on the shallow undulations of the embolus (Figs. 28, 92) and the very long posterior end of the abdomen (Figs. 27, 31).

Description.--Holotype male (Figs. 25-28, 92): Carapace quite pale, slightly darker along sides. Abdomen almost solid silver with thin lines of reticulation between blocks (Fig. 27). Posterior eyes separated by 1.5 x diameter (Fig. 25). Cephalic process 16% length of carapace, slightly rounded and blunt at tip, very slightly distended (Fig. 26). Clypeal process 75% length of cephalic process. Abdomen very elongate and narrow, only ~2.0 x higher at spinnerets than half-way between the spinnerets and posterior end (Fig. 27). Abdomen 5.5 mm length, 67% of this posterior to spinnerets. Carapace 1.7 mm length, 0.7 mm width. Palp (Fig. 28, 92): Embolus with two shallow undulations of similar height, projecting slightly above cymbium, parallel to conductor along length; conductor same length as embolus.

Allotype female (Figs. 29-32, 104, 115): Color similar to male. Posterior eyes separated by 1.4 x diameter (Fig. 29). Cephalic process 4% length of carapace, similar in shape to, but 3.7 x as long as, clypeal process (Fig. 30). Abdomen very long and thin, ~2.0 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 31). Abdomen 8.1 mm length, 76% of this posterior to spinnerets. Carapace 1.9 mm length, 1.0 mm width. Vulva (Fig. 32): receptacles elongateoval, ducts looped in figure-eight. In external view, receptacles visible, epigynal area indistinct (Fig. 104) and with scape curved over (Fig. 115).

Variation.--Total length varies 6.3-7.2 mm in males, 10.0-10.5 mm in females, due to variable elongation in the abdomen. The abdomen is bright gold in life, sometimes with red marks.

Natural history.--Ariamnes uwepa is found in wet and mesic forest habitats on the island of Oahu. Individuals have been collected from beneath leaves in the forest, though little else is known of its natural history.

Ariamnes poele new species Figs. 33-40, 93, 105, 116

Type specimens.--USA: Hawaiian Islands: holotype male, Molokai, Pepeopae boardwalk at night, 1150 m elev., 21.12[degrees]N, 156.90[degrees]W, 12 May 2000, M. Arnedo, G. Hormiga, I. Agnarsson (BPBM); allotype female, Molokai, Kamakou, 1280 m elev., 21.12[degrees]N, 156.87[degrees]W, 4 June 1997, C.P. Ewing (BPBM); 1 paratype male, 1 paratype female, Maui, Puu Kukui, 1067 m elev., 20.90[degrees]N, 156.63[degrees]W, 15 January 1998, R. Gillespie, K. Shaw (EMUC); 1 paratype female, Molokai, Upper Kawela Gulch, 1200 m elev., 21.10[degrees]N, 156.93[degrees]W, 5 January 1981, W.C. Gagne (BPBM).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian word "po'ele" meaning black or dark, and refers to the dark color of the abdomen.

Diagnosis.--Male Ariamnes poele can be distinguished (especially from A. corniger and A. alepeleke) based on the embolus which has very shallow undulations and is similar in length to the conductor (Figs. 36, 93). The cephalic projection is relatively short and thinner than in A. corniger (Fig. 34 as compared to 50). In both sexes the abdomen is black above with a silver blotched bar along midline of side (Figs. 35, 39).

Description.--Holotype male (Figs. 33-36, 93): Carapace dusky black with light line running down midline. Abdomen dark brown/ black with silver bar along midline of side (Fig. 35). Legs banded. Posterior eyes separated by 2.0 x diameter (Fig. 33). Cephalic process 19% length of carapace, very slightly curved over at tip, not distended and with numerous robust setae at tip (Fig. 34). Clypeal process 68% length of cephalic process. Abdomen elongate and only 1.4 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 35). Abdomen 3.4 mm length, 67% of this posterior to spinnerets. Carapace 2.3 mm length, 1.2 mm width. Palp (Figs. 36, 93): embolus fairly long with shallow undulations, projecting slightly above distal margin of cymbium; conductor (membranous) broad, tapering to point, similar in length to embolus.

Allotype female (Figs. 37-40, 105, 116): Color similar to male. Posterior eyes separated by 1.7 x diameter (Fig. 37). Cephalic process 4% length of carapace, slightly more pointed and larger (3.7 x as long) than clypeal process (Fig. 38). Abdomen fairly long, not high, ~1.7 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 39). Abdomen 5.9 mm length, 72% of this posterior to spinnerets. Carapace 2.1 mm length, 1.1 mm width. Vulva (Fig. 40): receptacles large and spherical, ducts looped. In external view, receptacles visible, epigynal area indistinct (Fig. 105) and with scape short and slightly curved (Fig. 116).

Variation.--Total length varies 3.9-4.6 mm in males, 6.6-8.2 mm in females, due to variable elongation in the abdomen. The abdomen is very dark in color with little apparent variability.

Natural history.--Ariamnes poele is found in wet forest habitats on the islands of Molokai and Maui (West Maui only). It appears to be largely free-living.

Ariamnes melekalikimaka new species Figs. 41-48, 94, 106, 117

Type specimens.--USA: Hawaiian Islands: holotype male, allotype female, Maui, Puu Kukui, 1067 m elev., 20.90[degrees]N, 156.63[degrees]W, 15 January 1998, R. Gillespie, K. Shaw (BPBM); 1 paratype female, Molokai, Pepeopae boardwalk at night, 1150 m elev., 21.12[degrees]N, 156.90[degrees]W, 12 May 2000, M. Arnedo, G. Hormiga, I. Agnarsson (EMUC).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian phrase "Mele Kalikimaka" meaning "Merry Christmas". The name refers to the color of the animal in life, which is sparkling bright green and red.

Diagnosis.--Male Ariamnes melekalikimaka can be distinguished based on the embolus which has a large distal undulation and is much longer and well separated from the conductor (Fig. 44, 94). The form of the cephalic projection is unique, the area separating the cephalic and clypeal processes almost rectangular (usually triangular shaped) (Fig. 42). Females (and to a lesser extent males) are recognized by the relatively short and high abdomen (height 35% length) (Fig. 47). The coloration of the abdomen is also unique, being distinctly marked with a background color of brown (bright green in life), covered all over by loosely packed silver blocks and with red lines and flecks.

Description.--Holotype male (Figs. 41-44, 94): Carapace dusky dark with light line running down midline. Abdomen brown (greenbrown in life) with rust diamond on the dorsal surface (Fig. 43). Posterior eyes separated by 1.7 x diameter (Fig. 41). Cephalic process 19% length of carapace, angular on inside border, curved over, thick but not noticeably distended, and with numerous robust setae at tip (Fig. 42). Clypeal process 87% length of cephalic process. Abdomen relatively short and high, 3.4 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 43). Abdomen 4.9 mm length, 64% of this posterior to spinnerets. Carapace 2.4 mm length, 1.2 mm width. Palp (Figs. 44, 94): embolus very long with deep distal undulation, projecting well above distal margin of cymbium; conductor (membranous) blunt-ended and terminating at edge of cymbium.

[FIGURES 41-48 OMITTED]

Allotype female (Figs. 45-48, 106, 117): Color similar to male. Posterior eyes separated by 2.3 x diameter (Fig. 45). Cephalic process 5% length of carapace, similar in shape but larger (3.4 x as long) than clypeal process (Fig. 46). Abdomen relatively short and high, ~1.7 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 47). Abdomen 4.5 mm length, 57% of this posterior to spinnerets. Carapace 2.2 mm length, 1.4 mm width. Vulva (Fig. 48): receptacles large and spherical, ducts in loose figure-eight. In external view, receptacles visible, epigynal area distinct, urn-shaped (Fig. 106), scape almost straight (Fig. 117).

Variation.--Total length varies 6.7-7.3 mm in males, 6.6-7.2 mm in females. The abdomen is brightly colored in life with red marks on iridescent gold.

Natural history.--Ariamnes melekalikimaka is found in wet forest habitats at middle elevations on the volcano of Puu Kukui, West Maui, and on Molokai. This species, appears to be largely free-living, but has been found on the webs of Orsonwelles sp.

Ariamnes corniger Simon Figs. 49-56, 95-96, 107, 118

Ariamnes corniger Simon 1900:447, plate XV, fig. 2.

Type specimens.--USA: Hawaiian Islands: 1 female syntype, Maui, Haleakala (MNHN); 1 female syntype, Lanai, Koele (MNHN).

Other material examined.--USA: Hawaiian Islands: Maui, Upper Waikamoi Preserve, 1860 m elev., 20.78[degrees]N, 156.22[degrees]W, 21 November 1997, M. Rivera, B. Thorsby, A. Bohonak, (EMUC); 2 [??], 2 [??], Maui, Auwahi, south slope, Haleakala, 1250 m, 20.65[degrees]N, 156.35[degrees]W, 20 August 1997, R. Gillespie, J. Garb, M. Heddle (EMUC); 2 [??], 2 [??], Maui, Auwahi, south slope, Haleakala, 1524 m elev., 20.65[degrees]N, 156.35[degrees]W, 4 March 1999, R. Gillespie, A. Medeiros, K. Teramura (EMUC); 2 [??], 2 [??], Maui, Kipahulu Valley, 1830 m elev., 20.72[degrees]N, 156.13[degrees]W, 27 April 1988, A.C. Medeiros (EMUC); 2 [??], 2 [??], Maui, bogs on NE. rift of Haleakala, 1676 m elev., 20.73[degrees]N, 156.10[degrees]W, 15 January 1988, R. Gillespie, A. Medeiros (EMUC).

Diagnosis.--Male Ariamnes corniger can be distinguished based on the shallow undulations of the embolus which is the same length as the conductor (Figs. 52, 95-96), and the robust and slightly curved cephalic process (Fig. 50). Both males and females can be recognized based on the anvil-shaped abdomen, much higher at the spinnerets, and the coloration of the abdomen, densely packed with silver blocks (Figs. 51, 55).

Description.--Male (Figs. 49-52, 95, 96): Carapace uniformly pale. Abdomen very densely packed all over with silver blocks on a fairly dark background (Fig. 51). Posterior eyes separated by 2.3 x diameter (Fig. 49). Cephalic process robust, 18% length of carapace, slightly curved over, not noticeably distended, and with numerous setae at tip (Fig. 50). Clypeal process 69% length of cephalic process. Abdomen elongate, much higher, 2.8 x, at spinnerets than half way between the spinnerets and posterior end (Fig. 51). Abdomen 4.5 mm length, 62% of this posterior to spinnerets. Carapace 2.1 mm length, 1.0 mm width. Palp (Figs. 52, 95, 96): embolus with shallow undulations, projecting above cymbium, roughly parallel to conductor along length; conductor (membranous) broad, tapering to point, same length as embolus.

Female (Figs. 53-56, 107, 118): Color similar to male. Posterior eyes separated by 2.0 x diameter (Fig. 53). Cephalic process 5% length of carapace, similar in shape to, but much more projecting (4.6 x as long) than clypeal process (Fig. 54). Abdomen anvil-shaped, ~2.8 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 55). Abdomen 6.2 mm length, 74% of this posterior to spinnerets. Carapace 2.0 mm length, 1.2 mm width. Vulva (Fig. 56): receptacles spherical, ducts tightly looped in figure-eight. In external view, receptacles fairly distinct, epigynal area indistinct (Fig. 107), scape slightly hooked (Fig. 118).

[FIGURES 49-56 OMITTED]

Variation.--Total length varies 6.2-6.9 mm in males, 8.0-8.9 mm in females. The abdominal coloration is somewhat variable, ranging from pale gold in the wet forest, to pure white in the dry forest, where it is camouflaged against the white lichen that is found throughout the area.

Natural history.--Ariamnes corniger is found on East Maui, where it occurs in both wet forest habitats on the north east slope, and dry forest on the south slope. To date, individuals have been found free living only.

Ariamnes alepeleke new species Figs. 57-64, 97, 108, 119

Type specimens.--USA: Hawaiian Islands: holotype male, allotype female, Maui, Waikamoi Flume, 1250 m elev., 20.90[degrees]N, 156.28[degrees]W, 8 February 1999, M. Rivera, & A. Rivera (BPBM); 1 paratype female, Maui, Waikamoi Flume, 1250 m elev., 20.82[degrees]N, 156.23[degrees]W, 13 June 1994, A.M. Tan (EMUC); 1 paratype female, Waikamoi, 1341 m elev., 20.80[degrees]N, 156.25[degrees]W, 13 August 1994, A.M. Tan (EMUC); 1 paratype male, Makawao Forest Reserve, 1463 m elev., 20.80[degrees]N, 156.25[degrees]W, under living leaves, 5 October 1979, S.L. Montgomery (BPBM).

Etymology.--The specific epithet, regarded as a noun in apposition, is named after Alepeleke Rivera, the collector of the type species and father of M.A.J. Rivera. Alepeleke is the Hawaiian name for Alfred.

Diagnosis.--Male Ariamnes alepeleke can be distinguished based on the shallow undulations of the embolus which barely projects above cymbium (Figs. 60, 97). In both males and females the abdomen is broadly triangular and solid silver (gold in life), sometimes with red superimposed.

Description.--Holotype male (Figs. 57-60, 97): Carapace fairly dark at sides, broad, almost white band running down midline. Abdomen very tightly packed all over with silver blocks with little room for reticulation between the blocks (Fig. 59). Ventral area above and anterior to spinnerets flat brownish. Legs uniformly yellow/brown. Posterior eyes separated by 2.5 x diameter (Fig. 57). Cephalic process 20% length of carapace, slightly curved over, not noticeably distended, and with a number of fairly short setae at tip (Fig. 58). Clypeal process 70% length of cephalic process. Abdomen relatively short, slightly higher, 1.2x, at spinnerets than half way between the spinnerets and posterior end (Fig. 59). Abdomen 3.2 mm length, 50% of this posterior to spinnerets. Carapace 1.8 mm length, 0.9 mm width. Palp (Figs. 60, 97): Embolus with shallow undulations, barely projecting above cymbium, curving away from conductor at tip; conductor broad, blunt-tipped, terminating at edge of cymbium.

Allotype female (Figs. 61-64, 108, 119): Color similar to male. Posterior eyes separated by 2.8 x diameter (Fig. 61). Cephalic process 3% length of carapace, similar in shape to, but larger (4.3 x as long) than clypeal process (Fig. 62). Abdomen broadly triangular, ~2.0 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 63). Abdomen 4.7 mm length, 67% of this posterior to spinnerets. Carapace 1.8 mm length, 1.1 mm width. Vulva (Fig. 64): receptacles spherical, ducts with single tight loop. In external view, receptacles and ducts fairly distinct (Fig. 108), scape slightly hooked (Fig. 119).

Variation.--Total length varies 4.8-5.0 mm in males, 5.5-6.6 mm in females. This species is bright iridescent gold in life, with variable saddle-shaped red markings on the dorsal surface of the abdomen.

Natural history.--Ariamnes alepeleke is found in wet forest habitats at middle elevations on East Maui. To date the species has been found only in free-living state, and usually hangs at night below large leaves in the forest.

Ariamnes laau new species Figs. 65-72, 98, 109, 120

Type specimens.--USA: Hawaiian Islands: holotype male, Maui, Hanawi Natural Area Reserve, Poouli Cabin, 975 m elev., 20.83[degrees]N, 156.13[degrees]W, 3 May 1998, J. Liebherr (BPBM); allotype female, Maui, Upper Waikamoi, 1860 m elev., 20.78[degrees]N, 156.22[degrees]W, 21 November 1997, M. Rivera, B. Thorsby, A. Bohonak (BPBM); 1 paratype male, Maui, Hanawi Natural Area Reserve, Poouli Cabin, 975 m elev., 20.83[degrees]N, 156.13[degrees]W, 3 May 1998, J. Liebherr (EMUC); 1 paratype female, Maui, Makawao Forest Reserve, 1463 m elev., 20.80[degrees]N, 156.25[degrees]W, under living leaves, 5 October 1979, S.L. Montgomery (BPBM); 1 paratype male, 1 paratype female, Maui, gulch east of Nianiau, 2130 m elev., 20.78[degrees]N, 156.25[degrees]W, swept from Elaphoglossum hirtum on shaded stream bank, 6 July 1975, W.C. Gagne (BPBM).

[FIGURES 57-64 OMITTED]

[FIGURES 65-72 OMITTED]

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian word "la'au" meaning stick, and refers to the common name often used for this group of Ariamnes: "stick spiders."

Diagnosis.--Male Ariamnes laau can be distinguished based on the embolus which is almost straight (without undulations) and shorter than the conductor (Figs. 68, 98), the latter barely projecting above the cymbium. Compared to A. corniger and A. alepeleke the cephalic process is shorter and straight (Fig. 66). In females the extension of the abdomen behind the spinnerets is greater than in A. alepeleke, and the difference in height of the abdomen (at spinnerets compared to behind) is much less pronounced than in A. corniger. In both sexes the abdomen is black with silver blotches, compared to silver and gold in A. corniger and A. alepeleke respectively.

Description.--Holotype male (Figs. 65-68, 98): Carapace almost black with light line running down midline. Abdomen dark, almost uniformly black on dorsal surface, with silver blotches along sides, decreasing posteriorly (Fig. 67). Legs dark and banded. Posterior eyes separated by 1.4 x diameter (Fig. 65). Cephalic process 14% length of carapace, straight, and slightly distended at tip with numerous robust setae (Fig. 66). Clypeal process 57% length of cephalic process. Abdomen relatively long, 2.4 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 67). Abdomen 4.2 mm length, 68% of this posterior to spinnerets. Carapace 1.7 mm length, 0.9 mm width. Palp (Figs. 68, 98): embolus almost straight and short, terminating well below distal margin of cymbium; conductor terminating at edge of cymbium, longer than embolus.

Allotype female (Figs. 69-72, 109, 120): Color similar to male. Posterior eyes separated by 2.0 x diameter (Fig. 69). Cephalic process 4% length of carapace, slightly more pointed and larger (3.4 x as long) than clypeal process (Fig. 70). Abdomen elongate, ~1.5 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 71). Abdomen 5.4 mm length, 74% of this posterior to spinnerets. Carapace 1.8 mm length, 1.0 mm width. Vulva (Fig. 72): receptacles spherical, ducts with single tight loop. In external view, receptacles indistinct, epigynal area marked with elongate transverse "[omega]" (Fig. 109), scape almost straight (Fig. 120).

Variation.--Total length varies 5.2-6.0 mm in males, 7.0-8.3 mm in females, due to variable elongation in the abdomen. The abdomen is very darkly colored with little apparent variability.

Natural history.--Ariamnes laau is found in wet forest habitats at middle and high elevations on East Maui. This species appears to be largely free-living, but has been found on the webs of Orsonwelles sp; habitat associations are unknown.

Ariamnes waikula new species Figs. 73-80, 99, 110, 121

Type specimens.--USA: Hawaiian Islands: Holotype male, allotype female, Hawaii, Thurston, Hawaii Volcanoes National Park, 1067 m elev., 19.42[degrees]N, 155.23[degrees]W, 29 May 1998, M. Rivera, A. Vandergast (BPBM); 1 paratype female, Hawaii, Honomalino, 1036 m elev., 19.17[degrees]N, 155.87[degrees]W, 5 March 1999, J. Garb, K. Fay, S. Steven (EMUC).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian word "waikula" meaning gold-colored, and refers to the solid gold color of the abdomen.

Diagnosis.--Male Ariamnes waikula can be distinguished based on the embolus which has very shallow undulations, bent up at tip to distal margin of cymbium, slightly shorter than the conductor (Figs. 76, 99). Cephalic process differs from A. alepeleke by being long and markedly curved over at distal end (Fig. 74). In females, copulatary ducts a simple loop (Fig. 80). In both sexes the entire abdomen is covered with large silver blocks with red superimposed on top.

Description.--Holotype male (Figs. 73-76, 99): Carapace pale with dark lines running down either side of midline. Abdomen dull brown covered all over with large irregular silver blocks, with red superimposed on top (Fig. 75). Posterior eyes separated by 2.0 x diameter (Fig. 73). Cephalic process long, 19% length of carapace, curved over at distal end, not noticeably distended at tip with numerous robust setae (Fig. 74). Clypeal process 70% length of cephalic process. Abdomen relatively long, 1.3 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 75). Abdomen 3.2 mm length, 60% of this posterior to spinnerets. Carapace 1.8 mm. Palp (Figs. 76, 99): embolus with very shallow undulations, bent up at tip, terminating at distal margin of cymbium; conductor (membranous) narrowing towards tip and terminating just above distal edge of cymbium, slightly longer than embolus.

[FIGURES 73-80 OMITTED]

Allotype female (Figs. 77-80, 110, 121): Color similar to male. Posterior eyes separated by 1.9 x diameter (Fig. 77). Cephalic process 5% length of carapace, slightly more pointed and larger (5.0 x as long) than clypeal process (Fig. 78). Abdomen elongate, ~2.3 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 79). Abdomen 6.6 mm length, 69% of this posterior to spinnerets. Carapace 1.8 mm length, 1.0 mm width. Vulva (Fig. 80): receptacles spherical, ducts with single tight loop. In external view, receptacles and ducts clearly visible (Fig. 110), scape short and slightly curved (Fig. 121).

Variation.--Total length 4.8-5.0 mm in males, 8.0-9.3 mm in females, due to variable elongation in the abdomen. The abdomen is shiny, iridescent gold in life, sometimes with red marks.

Natural history.--Ariamnes waikula is found in wet forest habitats at middle elevations on Hawaii island. It lives under larger leaves in the forest, and to date has been found to be exclusively free-living.

Ariamnes hiwa new species Figs. 81-88, 100, 111, 122

Type specimens.--USA: Hawaiian Islands: holotype male, Hawaii, Thurston, Hawaii Volcanoes National Park, 1067 m elev., 19.42[degrees]N, 155.23[degrees]W, 29 May 1998, M. Rivera, A. Vandergast (BPBM); allotype female, Hawaii, Puu Makaala, 1067 m elev., 19.55[degrees]N, 155.22[degrees]W, October 1997, D. Preston (BPBM); 1 paratype female, Hawaii, Puna Forest Reserve, 640 m elev., 19.44[degrees]N, 155.03[degrees]W, 11-14 February 1983, Y. Ching, S. Gon III (BPBM); 1 paratype male, 1 paratype female, Hawaii, Thurston, Hawaii Volcanoes National Park, 1067 m elev., 19.42[degrees]N, 155.23[degrees]W, 29 May 1998, M. Rivera, A. Vandergast (EMUC).

Etymology.--The specific epithet, regarded as a noun in apposition, is from the Hawaiian word "hiwa" meaning black and refers to the black dorsal surface of the abdomen of these spiders.

Diagnosis.--Male Ariamnes hiwa can be distinguished based on the embolus which has a deep distal undulation, coupled with long conductor drawn out into thread-like tip extending beyond distal margin of cymbium, much longer than conductor (Figs. 84, 100). Cephalic process very slightly curved over and distended at distal end (Fig. 82). In females, copulatory ducts in open figure-eight (Fig. 88). In both sexes the entire abdomen is dark black/brown on top, abdomen greatly elongated beyond spinnerets.

Description.--Holotype male (Figs. 81-84, 100): Carapace dark, lighter band running down midline. Abdomen dark on dorsal surface, brown below with scattered silver blocks and black flecks (Fig. 83). Posterior eyes separated by 1.1 x diameter (Fig. 81). Cephalic process 15% length of carapace, curved over very slightly and somewhat distended at distal end with numerous robust setae (Fig. 82). Clypeal process 70% length of cephalic process. Abdomen relatively long, 2.1 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 83). Abdomen 4.0 mm length, 63% of this posterior to spinnerets. Carapace 1.9 mm length, 0.9 mm width. Palp (Fig. 84, 100): embolus with very deep distal undulation, terminating just above distal margin of cymbium; conductor (membranous) narrowing distally and drawn out into long thread-like extension, extending well above distal margin of cymbium.

Allotype female (Figs. 85-88, 111, 122): Color similar to male. Posterior eyes separated by 1.4 x diameter (Fig. 85). Cephalic process 5% length of carapace, similar in shape and larger (3.7 x as long) than clypeal process (Fig. 86). Abdomen elongate, only ~1.5 x higher at spinnerets than half way between the spinnerets and posterior end (Fig. 87). Abdomen 5.2 mm length, 66% of this posterior to spinnerets. Carapace 1.9 mm length, 0.9 mm width. Vulva (Fig. 88): receptacles spherical, ducts coiled in open figure-eight. In external view, upper portion of receptacles distinct, making them appear semicircular in shape, epigynal area indistinct (Fig. 111), scape tightly hooked (Fig. 122).

Variation.--Total length varies 5.2-5.9 mm in males, 7.0-7.4 mm in females. The abdomen is very darkly colored with little apparent variability.

[FIGURES 81-88 OMITTED]

[FIGURES 89-100 OMITTED]

Natural history.--Ariamnes hiwa is found in wet forest habitats at middle elevations on Hawaii island. This species is very dark in color, and has been collected exclusively in the free-living state, where it appears to occur low down near the ground, often in rocky sites and inside rocky crevices.

[FIGURES 101-122 OMITTED]

ACKNOWLEDGMENTS

We are grateful to the following for help in collecting specimens: Miquel Arnedo, Adam Asquith, Andy Bohonak, Curtis Ewing, Gustavo Hormiga, Jim Liebherr, Art Medeiros, David Preston, Alfred (Alepeleke) Rivera, George Roderick, AnMing Tan, Kerry Shaw, Karen Teramura, Bonnie Thorsby, and Amy Vandergast. Thanks also to the Museum National d'Histoire Naturelle, in particular Christine Rollard, and the Bishop Museum, in particular Neal Evenhuis and Tino Gonsalves, for assistance in obtaining specimens. We are grateful to the following organizations for providing access to collection localities: State of Hawaii Department of Land and Natural Resources (especially Betsy Gagne and John Giffin), The Nature Conservancy of Hawaii, West Maui Land and Pineapple (especially Randy Bartlett), Haleakala National Park (especially Art Medeiros and Lloyd Loope) and Hawaii Volcanoes National Park. We owe much thanks to Jennifer Kane for help with the illustrations. This work has been supported by funds from the NSF and the Schlinger Foundation (to RGG) and from the Ecology, Evolution, and Conservation Biology Program and the Haumana Graduate Program at the University of Hawaii at Manoa (to MAJR).

Manuscript received 15 January 2004, revised 14 November 2006.

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R.G. Gillespie and M.A.J. Rivera: Division of Insect Biology, University of California Berkeley, 137 Mulford Hall, Berkeley, CA 94720-3114, USA. E-mail: gillespi@nature.berkeley.edu
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Author:Gillespie, R.G.; Rivera, M.A.J.
Publication:The Journal of Arachnology
Article Type:Report
Geographic Code:1U9HI
Date:Jan 1, 2007
Words:9147
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