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A review of the genus Hermatobates (Heteroptera: Hermatobatidae), with descriptions of two new species.

Abstract--The genus Hermatobates is revised and considered to contain 12 valid species, including two new species described herein: H. schuhi from the Ryukyu and Tokara Islands, and H. palmyra from the Line Islands. A morphological redescription is provided for H. djiboutensis from the Indian Ocean, and a neotype is designated for H. singaporensis from Singapore. All previous records of H. marchei from Australia are considered to instead represent H. weddi. A revised key is provided to all known species of Hermatobates, and scanning electron micrographs are provided for certain key characters of the legs, thoracic venter, and male genitalia in H. schuhi, H. palmyra, H. djiboutensis, H. weddi, H. marchei, and H. kula. Additional photographic illustrations are provided for key characters in H. haddoni, H. bredini, H. hawaiiensis, H. tiarae, H. palmyra, H. djiboutensis, H. singaporensis, H. marchei and H. armatus, and line drawing illustrations are provided for endosomal structures in H. weddi, H. haddoni, H. djiboutensis, and H. bredini. Distribution maps are also provided for all species. It is suggested that characters of the male endosomal and metasternal morphology provide an initial basis for delineation of five intraspecific species groupings. It is hypothesized that repeated episodes of vicariance related to alternating high and low Quaternary sea level stands have led to the current segregation of morphologically similar species within the H. weddi group in the region between Australia and Southeast Asia.

Key words: Hermatobates, taxonomy, new species, distribution.

INTRODUCTION

The Hermatobatidae are an obligatorily marine family of surface-dwelling water bugs, and were considered by Andersen (1991) to possibly represent the oldest extant group of marine Gerromorpha, with their phylogenetic affiliations obscure within the infraorder. The morphology and ground plan of the group were analyzed in detail by Andersen (1982), accompanied by many excellent illustrations. A general review of the biology and ecology of the genus was provided by Andersen and Polhemus (1976), with regional treatments of ecology and behavior by Foster (1989, 1996) for Fiji, D. Polhemus (1990) for Aldabra Atoll in the Western Indian Ocean, and Andersen and Weir (2000) for Australia.

Due to the marine habits of its constituent species, their uniformly wingless morphology, and other reductional character states, the taxonomic status of Hermatobates has been unsettled for over 100 years. The genus was originally described by Carpenter (1892), who placed it in the Gerridae. Coutiere and Martin (1901a) retained this familial assignment, and described the subfamily Hermatobatinae to hold the group. China (1957) and Miyamoto (1957) also retained the genus within the Gerridae, but three years later Hungerford and Matsuda (1960) and Matsuda (1960) excluded the family from Gerridae on morphological grounds, although they offered no alternative higher group placement. Poisson (1965) subsequently proposed the family Hermatobatidae to hold the genus, a course of action endorsed by Stys and Kerzhner (1975). Since that time, Azuma and Kinjo (1987) and Gross et al., (1991) have treated Hermatobates as a genus within Gerridae, while all other recent catalogs and taxonomic treatments (including Andersen, 1982, 1995; Cassis and Gross, 1995; Andersen and Weir, 2000; J. Polhemus and D. Polhemus, 2006) have accepted the separate family status, which is also recognized herein.

MATERIAL AND METHODS

The collections upon which the current study is based were obtained flora a series of expeditions to the Pacific Ocean and Indian Ocean regions over the past 20 years, coupled with generous donations from other researchers working in these areas and in the Caribbean. A detailed discussion of some of these collecting activities, dealing with far eastern New Guinea and adjacent archipelagoes, may be found in J. Polhemus and D. Polhemus (2006), along with color photographs of representative habitats.

Later collections in this time series (post-2000) were geo-referenced from GPS readings taken using Garmin GPS 48 and Garmin GPSmap 76S hand held GPS units calibrated to a WGS 84 datum. Earlier collections (pre-2000) have in some cases been retroactively geo-referenced using web-based geospatial browsers; these latter data are given in brackets to indicate they were not directly taken in the field. Salinity values are given in parts per thousand (ppt) and were taken at the water surface using a Fischer Scientific hand refractometer. Temperatures were similarly taken at the water surface using a hand-held thermometer. The CL numbers indicate collection locality numbers used by the authors to catalog sampling sites and cross-reference other metadata including photographs and field notes. Alternate names for geographical features are provided in brackets.

Our specimens of Hermatobates were collected using a wide array of techniques. Although other researchers have reported success with collecting Hermatobates in neuston tows run during both daylight and nighttime hours (Cheng and Schmitt, 1982), many of our specimens were instead taken by visual searching and hand netting on exposed reef flats at low tide during the day. The second author also collected specimens offshore of reef crests in the Indian Ocean during daylight hours by trolling nets from fiberglass dinghies moving at moderate speed, and a similar technique was used by both authors in the New Guinea region while riding in inflatable Zodiac vessels. Many collections were also made at night by hand netting of insects attracted to a incandescent lamps attached to docks or to the sterns of research vessels. In the latter case, a researcher would stand at water level on a duckboard made of metal grating anal attached to the stern of the vessel, netting insects from the surface as they skated through the circle of light cast by the lamp. Specimens collected by all the above methods were preserved in 75% ethanol, then transported to the Bishop Museum and Smithsonian Institution for detailed analysis and identification.

In order to assess character sets not easily observed with optical microscopy, specimens of various Hermatobates species were examined with a scanning electron microscope (SEM) in cases where the number of specimens available permitted such treatment. Specimens selected for SEM analysis were first cleaned in distilled water in an ultrasonic cleaner to remove salt residues, then desiccated in a critical point drier. The specimens thus prepared were then sputter coated with gold-palladium before being digitally imaged using a Philips XL30 SEM. The advantage of this approach is that it reveals fine morphological structures not otherwise visible; the drawbacks are that specimens committed to SEM are irrevocably altered by the sputter coating process (a disincentive to using this method on rare species), and that the thick setiferation covering much of the body on Hermatobates species serves to some extent obscure and confuse interpretation of underlying hard structures. Specimens of rare Hermatobates species not suitable for SEM treatment were instead examined using an AutoMontage digital imaging system linked to a Leica M 165 C dissecting stereomicroscope, which generates composite digital images with enhanced depth of field.

The majority of the specimens treated in this work are deposited in the U. S. National Museum of Natural History, Washington, DC (USNM), with synoptic series held in the Bishop Museum, Honolulu (BPBM), and the Polhemus Collection, Colorado Entomological Institute, Englewood, Colorado (JTPC); the latter will eventually be placed in the USNM. Duplicates of species collected in eastern New Guinea will also eventually be provided to an appropriate specimen depository specified by the Papua New Guinea Department of Environment and Conservation (DEC). Additional material from the Museum of Zoology, Cambridge University (CUMZ) was also examined.

In the following taxonomic section the descriptions of new taxa of Hermatobates follow the basic format of Andersen and Weir (2000). The generic description given in that work is not repeated here, so the reader is directed to that publication for guidance regarding species concepts and discrimination. Our objective in following the format established by Andersen and Weir is to facilitate comparison between the new taxa described herein and those previously described. All measurements in are given in mm. Based on character states revealed by use of SEM, we use the term "papillae" for small structures on the metasternum referred to as "denticles" by previous authors (i.e., Andersen and Weir, 2000).

TAXONOMY

FAMILY HERMATOBATIDAE POISSON 1965 GENUS HERMATOBATES CARPENTER 1892

DISCUSSION: The family Hermatobatidae is one of the few obligately marine insect groups on the planet, and specimens are rarely collected except by a few individual researchers who understand the specialized habits and ecology of this group. The putative rarity of these insects in collections was emphasized by Herring (1965) who noted that "The genus Hermatobates is certainly the rarest and least known genus of Marine Hemiptera. It was founded by Carpenter in 1892 for the reception of a new species, H. haddoni, based on a single specimen from Muibag Island in the Torres Straits, northern Australia. Since that time, five other species have been described. The descriptions of the six species were based on a total of nine specimens and several of the species are still known only from the type material. Females for the most part are either unknown or unassociated with the males. In 2-1/2 years of entomological work in the South Pacific, I was able to collect only three specimens; so it is not surprising that very few museums have representatives of this remarkable genus."

Over a decade later, Cheng (1977) echoed similar sentiments: "Although the genus Hermatobates was first established in 1892, the nine known species have hitherto been represented by only 14 type specimens and the females of only two species have been described ... In no case has a species been described from more than three specimens. Several sea-going entomologists have sought these elusive sea-bugs, but few have been successful and it is quite apparent that these insects are rare."

In the years since, the authors have made a point of searching for these fascinating insects whenever possible, working both individually and together across the Indo-Pacific region, and now possess collections from approximately 50 localities scattered across the seas of 20 countries worldwide, ranging from the coasts of Tanzania, Aldabra Atoll, Madagascar and Mauritius in the Indian Ocean, through the Malay Archipelago, the Philippines, Australia, and New Guinea, to the remote Pacific Islands including New Caledonia, the Solomons, Fiji, the Ryukyus, Palau, Micronesia, Hawaii, Tahiti and the Marquesas. Our collections now total more than 670 adult specimens plus hundreds of immatures, totalling approximately 1,000 specimens in all, and comprising nearly all of the known species. In addition to our own collections, kind gifts from many colleagues, especially Dr. Lanna Cheng, have permitted us to assemble a mostly unreported base of material that contains adequate series of every described species plus two more, described herein, that are new to science.

The evolution of our understanding of Hermatobates biology, and the consequent ability to locate individuals in the field, can be gauged by the fact that in the same year that the paper of Cheng (1977) cited above appeared in print, the senior author was able to collect 87 adults and 30 immatures of H. haddoni on East Point Reef at Darwin, Australia, effectively increasing the known number of specimens in world collections six rimes over. Similarly, the two most recently described species of Hermatobates are H. armatus (Andersen and Weir, 2000), based on 28 adults and 48 immatures, and H. kula (J. Polhemus and D. Polhemus, 2006), based on 103 adults and 22 immatures. Clearly, by the start of the 21st century the myth of excessive rarity in regard to Hermatobates (Herring, 1958) had been conclusively dispelled, and it is now understood that members of this genus are typical components of marine neuston assemblages throughout the tropical Pacific and Indian Oceans, and to some degree in the Caribbean as well, and that they can be effectively sampled by skilled collectors.

A key to the species of Hermatobates as known at the time, accompanied by a review of information regarding biology and ecology, was provided by Andersen and Weir (2000). These authors also provided an excellent redescription of the genus, not repeated here, and the reader should consult this work for the general plan and morphology of the group, the details of which are also not repeated here. Unfortunately, these authors also confused several species concepts, leading to listing of Australian material under an incorrect species name, as explained in greater detail below in the discussions under H. weddi and H. marchei.

The everted male endosoma in Hermatobates has a complex structure (Figs. 7, 18) which is useful in defining species groups, although this structure is difficult to extract and inflate, and is best examined in specimens were it has been extruded at the time of initial collection and preservation. All species of the weddi group (5 species) have a similar structure (Figs. 7A, 18A-D), and the djiboutiensis and haddoni groups (1-2 species each) a different but somewhat similar plan (Fig. 7B, C). By contrast, the structure in the hawaiiensis group (2 species) is much simplified, unlike those of the other species in the genus, and putatively plesiomorphic (Fig. 7D). No male specimens with everted genitalia are known for the Eastern Pacific species H. tiarae or H. palmyra, although these two species share a distinctive morphology of the posterior male metasternum, which is strongly depressed and bisulcate, indicating a sister species relationship.

On the basis of putative homologies of the male endosoma (Figs. 7, 18), male posteromedial process on the metasternum (Fig. 3), metasternal papillae (Fig. 4) and armature of the male forelegs (Fig. 5), we can propose the following species groups within Hermatobates, which are subject to future test using phylogenetic methods:

1) Hermatobates haddoni group. Included species: H. haddoni. Diagnostic characters: small size; anteriorly protruding head (Fig. 8); endosoma bearing lateral projections on the distal lobe (Fig. 7B). Distribution: Australia.

2) Hermatobates hawaiiensis group. Included species: H. hawaiiensis, H. bredini. Diagnostic characters: small size; male endosoma simple (Fig. 7D); male forelegs not heavily armed (Fig. 10B). Distribution: Hawaiian Islands, Caribbean Sea.

3) Hermatobates djiboutensis group. Included species: H. djiboutensis. Diagnostic characters: male metasternum broadly concave and depressed on posterior half; male metasternal process absent (Fig. 2A), but with papillae present in the area where such a process arises in other species (Fig. 3A); male endosomal structure complex and distinctive, similar in some repects to the character state seen in the H. weddi group (Fig. 7C). Distribution: Western Indian Ocean.

4) Hermatobates tiarae group. Included species: H. tiarae, H. palmyra. Diagnostic characters: male metasternum with posterior portion strongly excavate, forming a deep cavity immediately anterior to the genital complex; male metasternal process recessed and greatly reduced, if present then broad and bi-laterally bidentate, with small papillae (Figs. 2B, 3B); male endosomal characters unknown; male fore tibia basally incised (Fig. 12A, B). Distribution: Eastern Tropical Pacific.

5) Hermatobates weddi group. Included species: H. weddi, H. marchei, H. singaporensis, H. kula, H. armatus, H. schuhi. Diagnostic characters: well developed trapezoidal to quadrate posterior metasternal process (2C-F, 19) with prominent papillae (Fig. 3C-F); male fore tibia with variations on a common basal armature consisting of multiple large teeth (Fig. 5 A-D); complex male endosoma with prominent lateral lobes (Figs. 7A, 18A-D). Distribution: Australia, Melanesia, Philippines, Caroline Islands, Ryukyu Islands.

The species of the first three groups possess conspicuous individually diagnostic characters and are easily keyed. The two species of the H. tiarae group are geographically proximal sister taxa that can be separated by a number of somewhat more subtle character states. By contrast, the species of the H. weddi group, although separable on the basis of the everted endosoma and fine scale character systems revealed by SEM, are more challenging to distinguish using consistently available characters visible under optical microscopy; as a result, we have devoted particular attention to this group in the present work. Even so, we consider our current treatment of the H. weddi group to be a work in progress, given the degree of evident intraspecific variability, and the large gaps in geographic coverage still remaining in the Western Pacific. We would not be surprised if a future analysis of the H. weddi group, including use of molecular characters, reveals additional cryptic species, particularly within the broad species concept we have adopted for H. weddi per se.

KEY TO SPECIES OF HERMATOBATES

Modified from China (1957), Andersen and Weir (2000), and J. Polhemus and D. Polhemus (2006).

1. Width of head across eyes greater than 4x the length along the longitudinal midline (Fig. 8); posterior margin of metasternum straight or at most only weakly emarginate, bearing a number of minute black spinules but lacking a medial lobe or prominence (Fig. 9); middle femur of female with a ventral row of short bur distinct dark spines; length of male 2.6-3.2 mm, of female 2.6-2.7 mm; northern coast of Australia (Fig. 24) H. haddoni Carpenter

--Width of head across eyes less than 4x the length along the longitudinal midline 2

2. Fore femur and tibia of male lacking dark teeth or tubercles; posterior margin of male metasternum bearing a large, distally flaring posteromedial process resting against the genital capsule, posterior margin of this process emarginate, curving downward, and bearing numerous short, vertical striae (Fig. 11); male length 2.4-2.5 mm, female length 2.7-2.8; Caribbean Sea (Fig. 22) H. bredini Herring

--Fore femur and tibia of male with at least a few dark teeth on ventral surfaces; posterior margin of male metasternum not modified as above, posteromedial process on posterior margin of metasternum, if present, not resting against genital capsule 3

3. Ventral margin of male fore femur with a row of ~10 evenly sized small black spines, without larger basal and apical spurs flanking these spines (Fig. 10B); fore tibia without teeth or tubercles; posterior margin of male metasternum with two prominent, squared-off medial processes separated by a wide gap centrally (Fig. 10 A); length of male 2.5-3.0 mm, of female 3.0-3.1 mm; Hawaiian Islands (Fig. 20) H. hawaiiensis China

--Ventral margin of male fore femur bearing basal and apical teeth that are larger than the remaining dark ventral teeth; fore tibia with a few dark teeth or tubercles, at least basally; posterior margin of male metasternum either straight or with a single medial process, this process sometimes bidentate in the form of two lateral angles separated by a broad, shallow medial concavity, but never in the form of two prominent quadrate processes as noted above 4

4. Male fore tibia incised basally to accept apex of large distal tooth on fore femur (Fig. 12A, B); posterior portion of male metasternum folded upward and inward across its entire width to form a deep, hair-free cavity anterior to the genital capsule, posterior margin of metasternum either unmodified or bearing a broad, bidentate, centrally concave medial process, this process deeply recessed in pregenital cavity and therefore often difficult to see (Figs. 2B, 3B); fore femur strongly incrassate, maximum width 0.35-0.40x the length, armed with a prominent basal tooth and large, blunt, weakly bifid tooth near the apex plus a row of 6-10 shorter spines or spinules lying between these; Eastern Pacific 5

--Male fore tibia not incised basally (Fig. 5A-D); male metasternum not folded so strongly downward and inward as above, if a pregenital depression present then this cavity occurring only on central section of metasternum and lined with silvery setae; posterior margin of male metasternum either lacking a medial process (Fig. 2A, 3A), or if such a process present then prominent, easily observed, and either apically parallel sided (Figs. 2C, 3C, 19), or trapezoidal (Figs. 2 D-F, 3 D-F), rather than broad, medially concave, and bidentate as above (Figs. 2B, 3B); fore femur of male more or less incrassate, armed with prominent basal and apical teeth and a row of 6-14 short, dark spines between them; Indian Ocean and Western Pacific 6

5. Male fore tibia with a narrow, acutely angulate depression basally (Fig. 12B), inner face of tibia distal to this depression bearing two angular teeth, one immediately distal to depression, the other near midpoint of tibia; middle fore femur with only ventral margin of male fore femur with 3-5 widely separated small, sharp black teeth in area between prominent basal and distal spurs; concave and depressed posterior section of metasternum bearing a low but distinct longitudinal carina medially; French Polynesia (Fig. 20) H. tiarae Herring

--Male fore tibia with a broad, obtusely angulate depression basally (Fig. 12A), inner face of tibia distal to this depression bearing a single tooth lying immediately distal to the depression, with no other prominent teeth centrally; ventral margin of male fore femur with 9-10 very small teeth in area between prominent basal and distal spurs (Fig. 12A); concave and depressed posterior section of metasternum lacking longitudinal carina medially; Line Islands (Fig. 20) H. palmyra n. sp.

6. Posteromedial process on male metasternum either absent or weakly formed, recessed and difficult to see (Figs. 2A, 3A, 15B), bearing few papillae 7

--Posteromedial process on male metasternum prominent, easily visible, quadrate or trapezoidal in shape, bearing numerous papillae (Figs. 2C-F, 3C-F, 4A-D) 8

7. Posterior half of male metasternum strongly depressed centrally, posterior section with numerous small, dark papillae (appearing as denticles), these papillae concentrated towards the posterior margin (Figs. 2A, 3A); fore femur of male armed with 6-9 large, stout, conspicuous black teeth between basal and apical spurs, the largest of these teeth approaching the size of the basal spur (Fig. 14A); length of male 3.1-3.9 mm, of female 3.3-3.6 mm; Western Indian Ocean (Fig. 21) H. djiboutensis Coutiere and Martin

--Male metasternum slightly depressed postero-medially, bearing a weakly formed, somewhat trapezoidal medial process with a slightly sinuate posterior margin and a few papillae (Fig. 15B), this process recessed adjacent to genital segment and often difficult to see; fore femur of male armed with 8-13 small, sharp, relatively inconspicuous, black, triangular teeth between basal and apical spurs, all of these teeth much smaller in size than either the basal or apical spur (Fig. 15A); length of male 3.2 mm, of female 3.2-3.3 mm; Malay Peninsula and Singapore (Fig. 23) H. singaporensis Cheng

8. Male metasternal process parallel sided, at least on apical half (Figs. 2C, 3C, 19) 9

--Male metasternal process trapezoidal, with lateral margins angling inward, never parallel sided (Figs. 2D-F, 3D-F) 10

9. Male metasternal process with margins angling inward on basal half, then parallel sided on distal half (Fig. 19), posterior margin bidentate with small points at posterolateral angles, papillae on process small and inconspicuous; Chesterfield Islands (Fig. 24) H. armatus Andersen and Weir

--Male metasternal process tongue-like and quadrate, with margins parallel along entire length, posterior margin slightly bilobate but not possessing small points at posterolateral angles (Figs. 2C, 3C), papillae on process large and prominent (Fig. 4C); apicolateral lobes of male endosoma prominent, rounded (Fig. 18B); Philippines, Palau and northern New Guinea (Figs. 23, 24) H. marchei Coutiere and Martin

10. Male with metasternum tumid posteriorly; median metasternal process distinctly removed from posterior margin of metasternum and angling outward (Fig. 2E, 3E); fore tibia with a distinct tubercle in basal curve, followed by large gap and then two stout, truncate teeth separated by a notch (Fig. 5A); apicolateral lobes of endosoma large and angulate (Fig. 18A). Female middle femur with ~14 very small black teeth along ventral margin; eastern New Guinea, D'Entrecasteaux Islands and Louisiade Archipelago (Fig. 24) H. kula J. Polhemus and D. Polhemus

--Male with metasternum sloping posteriorly to base of genital segment; median metasternal process on a ridge at apparent posterior margin of metasternum (Figs. 2D, F, 3D, F); fore tibia with at most a low, broadly triangular tubercle in basal curve or with tubercle absent (Fig. 5C, D), basal portion of tibia beyond curve with a low, weakly bidentate protuberance followed by a distinct tubercle (Fig. 5C, D); apicolateral lobes of endosoma smaller and more rounded (Fig. 18B-D). Female middle femur with at most ~7 very small black teeth along posterior margin, or with such teeth reduced to spinules and barely evident; Ryukyu Islands or Australia and southwestern Pacific island archipelagoes (Figs. 20, 23, 25) 11

11. Male fore tibia with largest tooth on inner margin separated from the broader, bifid tooth lying immediately basal to it by a depression 2x with width of the large tooth (Fig. 5D); male fore tibia with inner margin bearing two low tubercles beyond basal curve, followed by a broad, black tooth, then a row of small tubercles leading to a small black tooth about two thirds distance from base to apex; papillae on medial metasternal process large and scattered (Figs. 2F, 3D, 4B); posterolateral lobes of male endosoma not expanded or explanate (Fig. 18D); female with posterior margin of middle femur bearing only a few tiny, inconspicuous black teeth along ventral margin; Australia and southwest insular Pacific (Figs. 20, 23, 25) H. weddi China

--Male fore tibia with largest tooth on inner margin separated from the broader, bifid tooth lying immediately basal to it by a depression equal to the width of the large tooth (Fig. 5C); male fore tibia with inner margin bearing a broad, bifid tubercle just beyond basal curve, followed by a depression, then a large stout tooth (Fig. 5C), then a row of small tubercles extending about two-thirds remaining distance to apex; papillae on medial metasternal process smaller and more numerous (Figs. 2D, 3F, 4A); posterolateral lobes of male endosoma expanded and explanate (Fig. 18C); female with posterior margin of middle femur bearing a row of about 7 small black spines on basal hall; Ryukyu Islands (Figs. 23, 25) H. schuhi n. sp.

Hermatobates haddoni, Carpenter Figures 7B, 8, 9, 24

Hermatobates haddoni Carpenter, 1892: 143.

Hermatobates walkeri China, 1957: 346. Syn. by J. Polhemus, 1982: 10.

DIAGNOSIS: For a detailed morphological redescription of this taxon see Andersen and Weir (2000). Male: length 2.48-2.83 mm, maximum width 1.15-1.40 mm. Female: length 2.55-2.75 mm, maximum width 1.23-1.35 mm.

COLORATION: Color dark brown, dull, body with fine silvery pubescence (Fig. 8). Head dark above, with a transverse line of pale spots at base. Pleural and ventral surfaces pale brownish, dull, genital segments dark brown, shiny. Antennae pale brown. Legs brown, except fore coxae and trochanters, ventral surface of fore femur, fore tibia, middle and hind trochanters pale.

Male legs with fore trochanter bearing single small tooth distally just before apex. Fore femur strongly incrassate, bearing a pointed black tooth at base angled distally, a distally bifid black tooth near apex, between these an almost regular row of 8 small black teeth. Fore tibia moderately curved, inner side with a low tubercle in sharp basal curve, several small teeth toward apex, also a small tooth distally at base of oblique comb of hairs. Trochanter of middle leg with small black tooth on ventral margin near apex. Middle femur slightly thickened centrally, ventral surface armed with ~25 stout black spines, these spines becoming shorter and more slender distally. Hind femur slightly thickened on basal third, weakly dorsoventrally compressed, hind trochanter and femur unarmed.

Female legs with fore femur bearing a row of 10-13 small black teeth along ventral margin, fore trochanter and tibia unarmed. Middle trochanter with a small spine; middle femur with ventral margin bearing a row of 12-15 short, black spines.

Male metasternum lacking a posteromedial process on posterior margin (Fig. 9), but bearing several large papillae centrally; styliform processes of abdominal segment VIII relatively slender with hooked apices; endosoma bearing lateral projections on the distal lobe (Fig. 7B).

DISTRIBUTION: Originally described from Muibag Island, in the Torres Strait (Carpenter, 1892), with subsequent records from Western Australia and the Northern Territory (Walker, 1893; China, 1957; J. Polhemus, 1982; Andersen and Weir, 2000). Based on current collections, this species is endemic to the northern and western coasts of Australia and adjacent small islands (Fig. 24 herein, and see Fig. 15 in Andersen and Weir, 2000). As discussed by Andersen and Weir (2000), records of this species from other parts of the Pacific (China and Usinger, 1950) are in error.

DISCUSSION: Hermatobates haddoni is a small species occurring along the northern along the northern and western coasts of Australia. Both sexes may be readily recognized by their small size, and the anterior projection of the head, which is unique within the genus. Males are also notable in lacking a posteromedial process on the hind margin of the metasternum (Fig. 9), with only a few papillae being present in this area, similar to the character state seen in H. djiboutensis (Figs. 2A, 3A). In addition, the male endosoma of H. haddoni is also unique within the genus, bearing lateral projections on the distal lobe (Fig. 7B). China (1957) provided useful line drawings of the ventral male body habitus and male forelegs. Andersen and Weir (2000) provided additional line drawings of the dorsal body habitus, ventral thorax and abdomen, and leg characters for both males and females.

SPECIMENS EXAMINED: AUSTRALIA: Northern Territory: Darwin, East Point Reef [12[degrees]24'25"S, 130[degrees]48'56"E], low tide, 10 December 1977, CL 904, J. T. Polhemus, 12 [male][male], 28 [female][female] (JTPC).

Hermatobates hawaiiensis, China Figures 10A, B, 20

Hermatobates hawaiiensis China, 1956: 353.

DIAGNOSIS: For a detailed morphological description of this taxon see China (1956). Male: length 2.90-3.00 mm, maximum width 1.30-1.35 mm. Female. length 3.00-3.10 mm, maximum width 1.40-1.50 mm.

COLORATION: Color dark brown, dull, body with short, fine, erect silvery pubescence. Head dark brown above, without transverse pale coloration at base of vertex. Pleural and ventral surfaces brownish, dull, genital segments pale brown, shiny. Antennae uniformly brown. Legs dark brown, dorsal surfaces of all femora and tibiae dark grey, all coxae, trochanters and tarsi brown.

Male legs with fore trochanter bearing a single, erect dark spine near distal end, this spine longer than any on the fore femur. Fore femur weakly incrassate, bearing a regularly spaced row of 11-12 small, sharp black spines of gradually decreasing size going distally (Fig. 10B). Fore tibia nearly straight, slightly expanded distally, lacking teeth or spines, covered with long, distally directed semi-erect gold setae. Middle and hind trochanters unarmed. Middle femur slightly thickened in middle, ventral surface armed with 10 long, straight, sharp spines interspersed with 7 much shorter spines. Hind femur slightly thickened on distal third, dorsoventrally flattened and bearing a narrow longitudinal sulcus ventrally, this sulcus lined with short, pale, semi-erect setae.

Female legs with hind femur bearing ~10 minute black spinules along posterior margin.

Male metasternum with posterior margin bearing a pair (1 + 1) of broad, apically truncate processes separated by a central gap equal to the width of a process (Fig. 10A). Styliform processes of segment VIII strap-like, bearing radiating tufts of long gold setae.

DISTRIBUTION: Endemic to the Hawaiian Islands, and currently recorded from Kauai, Oahu and Hawaii islands within that archipelago (Fig. 20), but undoubtedly occurs along the coasts of other high islands in the group. Originally described from Coconut Island, in Kaneohe Bay on the eastern shore of Oahu (China, 1956), with subsequent records from Kauai and Hawaii islands (Usinger and Herring, 1957; Cheng and Leis, 1980).

DISCUSSION: Easily distinguished by its small size, relatively simple armature on the male fore femur consisting of 10 small, almost evenly sized slender spines (Fig. 10B), the presence of a row of tiny spinules on the posterior margin of the female hind femur, and the pair of broad, apically truncate processes on the male metasternum (Fig. 10A).

Both H. hawaiiensis from the Hawaiian Islands and H. bredini from the Caribbean are distinctive within Hermatobates in having weakly modified male forelegs and a relatively simple male endosoma without elaborated lobes or projections, suggesting a sister species relationship. Cheng (1977) provided scanning electron micrographs of the eye, mid-tarsal claws, and thoracic microtrichae, as well as small-scale black and white dorsal view photos of the adult male and female.

ECOLOGICAL NOTES: Although present throughout the main Hawaiian Islands, this species is uncommon and difficult to locate. The localized nature its occurrence on a given island is illustrated by the experience of Cheng and Leis (1980), who reported that repeated collection efforts involving hundreds of neuston tows run over a period of three years (1971-1973) near the Coconut Island type locality in Kaneohe Bay on the eastern side of Oahu produced no additional specimens, while by contrast 34 specimens were taken in a single night on 14 April 1973 at Kahe Point on the western side of Oahu. We have also captured this species at the Kahe Point locality, where it frequents the warm water discharge plume from the nearby power plant. In addition, we were able to collect a long series attracted to light at night on the calm waters of Honokohau Harbor near Kailua-Kona on Hawaii Island; at this latter locality the insects were living under a carpet-covered fuel dock floating close to the water, and possibly ovipositing in the sections of carpet that hung down into the water. Attempts to collect this species at other, more exposed locations in the Kona area by contrast proved unsuccessful.

SPECIMENS EXAMINED: USA: Hawaii: Oahu Is., Kahe Point [21[degrees]21'21"N, 158[degrees]07'51"W], calm sea 200 m. offshore, 18:00 hrs., 2 August 1991, D. A. & J. T. Polhemus, 4 [female][female] (JTPC); Oahu Is., Waikiki, Gray's Raft, thriving in salt water, 19 March 1922, E. H. Bryan, 1 immature (BPBM); Hawaii Is., Honokohau Harbor, N. of Kailua Kona, 19[degrees]40'7"N, 156[degrees]01' 32"W, at night light from dock, 21 July 2009, CL 8518, D. A. & J. T. Polhemus, 4 [male][male], 15 [female], 11 immatures (JTPC); Kauai Is., Hanalei Bay, 28 April 1972, L. Cheng, 2 [male][male] (JTPC); same data as preceding but 5 June 1972, 1 [male] (JTPC).

Hermatobates bredini, Herring Figures 7D, 11, 22

Hermatobates bredini Herring, 1965: 124.

HOLOTYPE: DOMINICA: Woodbridge Bay, BWI, 25 March 1956, W. L. Schmitt, [male], Type No. 66875 (USNM).

DIAGNOSIS: For a detailed morphological description of this taxon see Herring (1965). Male: length 2.45-2.55 mm, maximum width 1.10-1.20 mm. Female: length 2.70-2.80 mm, maximum width 1.30-1.40 mm.

COLORATION: Color dark reddish brown, dull, body with very short, inconspicuous silvery pubescence. Head dark reddish brown above, without pale markings at base of vertex. Pleural and ventral surfaces dark brown, dull, acetabulae and central thoracic venter orange brown, medium brown, shining. Antennae with segments I-III medium brown, segment IV dark brown. Legs dark brown above, medium brown below, except all trochanters yellowish brown.

Male legs with fore trochanter, femur and tibia lacking spines or teeth, ventral surface of fore femur bearing long, distally-directed, semi-erect pale setae. Middle and hind trochanters unarmed. Middle femur slightly thickened on basal half, becoming more slender and gently curving distally, ventral surface armed with 14 moderately long, straight, sharp spines. Hind femur straight, slightly thickened and dorsoventrally flattened on distal third.

Female legs with fore, middle and hind femora lacking spines or teeth.

Male metasternum with posterior margin bearing a single broad posteromedial process, this process slightly expanded distally, angling downward and resting against genital capsule, posterior border broadly concave, emarginate, forming a downward curved lip bearing numerous parallel vertical striae (Fig. 11). Styliform processes of segment VIII recurved distally, apices slightly bulbous. Male endosoma bulbous, lacking lateral lobes or projections (Fig. 7D).

DISTRIBUTION: Originally described from Woodbridge Bay, Dominica (Herring, 1965), with subsequent records from the Bahamas and Cuba (Polhemus and Herring, 1979). Spangler and Polhemus (1988) noted the presence of this species in Belize, and we provide specific records for that country herein (Fig. 22). Based on this distribution, the species is likely to occur in suitable habitats throughout the Caribbean Sea islands.

DISCUSSION: We have examined the holotype, a male specimen in moderately good condition, held in the Smithsonian Institution's National Museum of Natural History. The specimen is glued to a point, is missing antennal segments III and IV on both sides, and has been crudely dissected, with much of the terminal abdomen missing, and the genitalia in a small glass vial below on the same pin holding the specimen. Herring (1965) provided line drawings of the ventral habitus and male foreleg.

This species is easily distinguished by small size, lack of armature on the foreleg of the male or the hind femur of the female, and the distinctively expanded male posteromedial metasternal process, which is flared against the base of the genital segments in a striate lip (Fig. 11). The male endosoma is bulbous but simple in form, lacking significant modifications in the form of lateral lobes or projections (Fig. 7D), a character state shared with H. hawaiiensis.

OTHER SPECIMENS EXAMINED: BELIZE, Stann Creek Dist.: Carrie Bow Cay [16[degrees]48'09"N, 88[degrees]04'55"W], 20 May 1986, collection 12, P. J. Spanger & R. Faitoute, 2 [male][male], 3 [female][female] (JTPC). BAHAMAS: Long Island, night light, 23 August 1995, L. Cheng, 1 [female] (JTPC); San Salvador Is., 27 August 1995, L. Cheng, 3 immatures (JTPC). CUBA: Pinar del Rio Prov.: Corrientes Bay [Gunanahacabibes Peninsula], 9 April 1937, P. Bartsch, 1 [male] (JTPC).

Hermatobates tiarae, Herring Figures 12B, 20

Hermatobates tiarae Herring, 1965: 126.

HOLOTYPE: FRENCH POLYNESIA: Tuamoto Archipelago: 14[degrees]56'S, 146[degrees]01'W, 21 January 1953, dip net, lite, "Capricorn," 107 Cap, M. W. Johnson, [male], Type No. 66876 (USNM).

DIAGNOSIS: For a detailed morphological description of this taxon see Herring (1965). Male: length 3.20-3.45 mm, maximum width 1.40-1.60 mm. Female: length 3.05-3.10 mm, maximum width 1.50-1.60 mm.

COLORATION: Color dull bluish-grey, body with fine, erect, silvery pubescence, and longer setae. Head dark grey above, with a transverse line of brown spots at posterior margin of vertex. Pleural and ventral surfaces brownish-grey, dull, genital segments blackish-brown, shining. Antennae uniformly black. Legs black above, dark brown ventrally, fore coxae brown, all trochanters brown ventrally.

Male legs with fore femur incrassate, bearing a sharp, slender pointed black tooth at base, angled distally, a broad, distally bifid black tubercle before apex, between these an almost regular row of 3-5 small, sharp black teeth. Fore tibia moderately curved, inner side with a distinct triangular tubercle in sharp basal curve, followed immediately by a deep, V-shaped notch (Fig. 12B), then a low, moderately large, triangular tubercle, 4-5 very small tubercles, and another slightly larger blunt tubercle at near midpoint, femur becoming progressively more slender and curved on distal half, lacking additional armature. Fore trochanter with small, sharp spine near middle on ventral margin, middle trochanter with a very small spine distally on ventral margin, hind trochanter lacking spines or teeth. Middle femur slightly thickened in middle, ventral surface armed with a line of 9 stout spines, these spines intermixed with 12 smaller spines, length of smaller spines less than half the length of the larger spines. Hind femur distinctly thickened distally; middle and hind femora thickly set with long silvery setae.

Middle femur slightly thickened in middle, unarmed, central portion of posterior margin bearing 4 very long, slender dark setae.

Male metasternum with posterior strongly depressed, folded upward into body, creating a distinct concavity immediately anterior to genital segment, posteromedial metasternal process not evident. Styliform processes of segment VIII relatively slender, apices slightly spatulate, not hooked.

DISTRIBUTION: Originally described from the Tuamotu Archipelago of French Polynesia (Herring, 1965); in this same publication he hypothesized that an immature specimen taken in the Marquesas and attributed to H. haddoni by China and Usinger (1950) in fact represented H. tiarae. We can now definitively document the presence of this species in the Marquesas Islands, thus confirming Herring's hypothesis, and also provide records from Moorea, in the Society Islands. Based on known collections, H. tiarae occupies an elongate east-west distribution south of the equator in the Eastern Pacific (Fig. 20).

DISCUSSION: We have examined the holotype, a male specimen in moderately good condition, held in the Smithsonian Institution's National Museum of Natural History. The specimen has been pinned through the center of the body, is missing antennal segments II-IV on the left side and segments III and IV on the right, and has the right foreleg removed and placed in a small glass vial below the labels on the same pin holding the specimen. The male genitalia of the type specimen have not been dissected. Herring (1965) provided line drawings of the ventral habitus and male foreleg.

Hermatobates tiarae may be easily recognized by the strongly depressed postrior portion of the metasternum which lacks an evident metasternal process, the distinctively notched male fore tibia (Fig. 12B), and the armature of the male fore femur, the ventral margin of which bears only 4-5 widely separated moderately small, sharp black spines between the large basal and apical spurs.

OTHER SPECIMENS EXAMINED: FRENCH POLYNESIA: Society Islands: Moorea Is., Cook's Bay [17[degrees]30'14"S, 149[degrees]49'14"W], at light, 10-11 June 1999, L. Cheng, 1 [male], 8 [female][female] (JTPC). Marquesas Islands: Tahuata Is., Vaitahu, harbor at boat landing, 9[degrees]56'11"S, 139[degrees]06'33"W, on ocean surface at night, at light, 27 October 1999, CL 4250, D. A. & J. T. Polhemus, 4 [male][male], 5 [female][female] (JTPC).

Hermatobates palmyra, new species Figures 2B, 3B, 12A, 20

HOLOTYPE: USA: Palmyra Atoll: Station 134, 2025-2054, 12 September 1990, G. Boehlert, [male] (BPBM).

DESCRIPTION: Male: length 3.30-3.45 mm, maximum width 1.50-1.60 mm. Female: length 3.10-3.30 mm, maximum width 1.50-1.60 mm.

COLORATION: Ground color dark grey with a bluish tinge, dull, body with fine silvery pubescence, and longer setae laterally (0.10 in males; 0.05-0.11 in females). Head dark above, with a transverse line of reddish brown irregular spots at base. Pleural and ventral surfaces bluish-grey, dull, thoracic and abdominal sternites embrowned centrally, genital segments shining brown. Antennae black. Legs shining black dorsally, dark brown ventrally.

Apterous male: body fusiform, length more than 2.0x greater than greatest width across thorax (3.40 : 1.60). Head length 0.27x greatest head width across eyes (0.35 : 1.30); eyes rather small, about 0.29x interocular width (0.23 : 0.80).

Antennal formula I : II : III : IV; 0.60 : 0.75 : 0.45 : 0.42; segment I less than 2x head length (0.60:0.35), stout, slightly thicker than segment IV; combined length of all antennal segments 0.65x total length of insect.

Pronotum very short, median length (0.12) slightly greater than half of an eye width; posterior broadly curved backwards. Meso- and metanotum simple. Metasternum posteriorly depressed, meeting depressed basal abdominal sterna to form a deep depression anterior to genital segment, this depression bearing a bidentate medial process with small papillae (Figs. 2B, 3B).

Proportions of legs as follows: Femur, tibia, tarsal 1, tarsal 2, tarsal 3 of fore leg, 1.10 : 1.10 : 0.05 : 0.05 : 0.20; of middle leg, 1.50 : 0.90 : 0.10 : 0.45 : 0.40; of hind leg, 1.45 : 1.90 : 0.10 : 0.48 : 0.55. Trochanter of fore leg with single small black spine before distal end. Fore femur incrassate, width 0.40, with a sharply pointed black tooth at base, angled distally, a broad distally truncate black tubercle before apex, between these an almost regular row of 9-10 small black teeth (Fig. 12A). Fore tibia (Fig. 12A) sharply curved basally, gently curved distally, central portion straight, inner side with a broad, weakly bidentate tubercle near basal curve, a row of small black spinules extending about two-thirds remaining distance to apex. Middle trochanter unarmed. Middle femur slightly thickened in middle (0.20), ventral surface armed with 8 widely spaced stout, sharp, black spines, spaces before, after, and between these spines each occupied by 1-3 shorter sharp black spinules. Hind femur slightly thickened centrally (0.15), not dorsoventrally compressed; trochanter and femur unarmed. All leg segments thickly set with long, erect, silvery pilose setae.

Abdomen very short, length from abdominal scent orifice to tip 0.70; lengths of abdominal terga 4-7: 0.10, 0.10, 0.15, 0.18; genital segments large, length 0.70; styliform processes of segment VIII with apices slightly spatulate, not hooked.

Apterous female: body fusiform, length about 2.0x greater than greatest width across thorax (3.00 : 1.50). Head length 0.25x head width across eyes (0.30 : 1.20); eyes rather small, width approximately 0.25x interocular width (0.20 : 0.80).

Antennae relatively shorter than in male, 0.6x total length of insect, antennal formula I : II : III : IV; 0.60 : 0.65 : 0.40 : 0.40; segment IV about 1.3x head length (0.40:0.30), subequal in thickness to segment I.

Pronotum very short, median length (0.10) about half of an eye width; metasternum not modified.

Legs relatively shorter than in male. Proportions of legs as follows: Femur, tibia, tarsal 1, tarsal 2, tarsal 3 of fore leg, 0.70 : 0.75 : 0.03 : 0.07: 0.20; of middle leg, 1.10 : 0.60 : 0.09 : 0.40: 0.35; of hind leg, 1.30 : 0.80 : 0.01 : 0.30 : 0.50. Fore femur slightly swollen, greatest width 0.20; trochanter and femur unarmed except for a row of about 13 small black teeth ventrally on femur; fore tibia gently curving on outer margin, nearly straight along inner margin, unarmed. Middle femur slightly thickened in middle (0.18), unarmed, central portion of posterior margin bearing 3 very long, slender dark setae; trochanter unarmed. Hind femur slightly thickened in middle (0.14), unarmed. All leg segments bearing erect, slender silvery setae, these setae shorter and less dense than in male.

Abdomen length from abdominal scent orifice to tip 1.40; dorsal segmentation obscure. Abdominal sterna II-VII forming a large quadrate plate, length 0.40, posterior margin slightly emarginate. Genital segments, except for cone-shaped proctiger, hidden beneath abdominal plate.

ETYMOLOGY: The name "palmyra" is a noun in apposition and refers to the Palmyra Aton type locality.

DISTRIBUTION: At present known only from Palmyra Atoll in the northern Line Islands (Fig. 20), and provisionally from Pagan Island in the northern Mariana Islands.

DISCUSSION: Hermatobates palmyra is morphologically similar to the geographically proximal H. tiarae from French Polynesia on the basis of the strongly depressed posterior margin of the metasternum that forms a deep concavity immediately anterior to the genital segments. Hermatobates palmyra, however, has a broad but very short metasternal process with small, slightly produced angles at the lateral corners (Figs. 2B, 3B); such a process is not evident in any male specimen of H. tiarae that we have examined. The structure and armature of the male fore femur and tibia are also somewhat different, with H. palmyra having a shallower anal more open notch basally on the tibia (compare Fig. 12A, B) and 9-10 small teeth on the ventral surface of the fore femur between the basal and apical spurs (Fig. 12A), as compared to an almost regular row of 3-5 small, sharp black teeth in H. tiarae. The type series was taken from neuston tows conducted offshore of Palmyra Atoll (5[degrees]53'19"N, 162[degrees]04'42"W) in the Line Islands by the United States National Marine Fisheries Service, so adults of this species clearly range well seaward of the reef crest.

We have also provisionally assigned our specimens from Pagan Island in the northern Mariana Islands to H. palmyra for the present, because they possess the distinctively concave posterior metasternum, a small and bidentate posteromedial metasternal process, and a basally incised fore tibia, but have not designated them as paratypes because of the significant geographic distance separating the two populations, which suggests that the Pagan population may eventually prove to be a distinct species once additional material is available for a more detailed evaluation. Collections from the intervening Marshall Islands would be very useful in resolving this question.

PARATYPES: USA: Palmyra Atoll: same data as holotype 2 [male][male], 4 [female][female] (JTPC, BPBM); Station 77, TC 9007, Manta, 2240-2301, neuston tow, 5 September 1990, G. Boehlert, 1 [male], 9 [female][female] (JTPC): same data as above except Station 123, 2021-2045, 11 September 1990, G. Boehlert, 5 [male][male] (JTPC); same data as above except Station 124, 2325-2349, 11 September 1990, G. Boehlert, 2 [male][male], 1 [female] (JTPC); same data as above except Station 127, 0837-0901, 12 September 1990, G. Boehlert, 2 [male][male], 5 [female][female] (JTPC); same data as above except Station 131, 1617-1641, 12 September 1990, G. Boehlert, 1 [female] (JTPC); same data as above except Station 135, 2255-2340, 12 September 1990, G. Boehlert, 2 [male][male], 2 [female][female] (JTPC); same data as above except Station 136, 0203-0227, 13 September 1990, G. Boehlert, 1 [male], 1 [female] (JTPC); same data as above except TC 9201, Station 229, 1541-1605, 27 February 1992, G. Boehlert, 4 [female][female] (JTPC).

OTHER SPECIMENS EXAMINED: USA: Commonwealth of the Northern Mariana Islands: Pagan Island, west coast, bay N. of Bandeera Peninsula, 18[degrees]07'48"N, 145[degrees]45'32"E, sea surface temp. 30[degrees]C., salinity 38 ppt., 18 July 2010, 10:30-11:30 hrs., CL 7471, D. A. Polhemus 1 [male] 3 [female][female] (BPBM).

Hermatobates djihoutensis, Coutiere and Martin Figures 1, 2A, 3A, 7C, 13, 14A, B, 21

Hermatobates djiboutensis Coutiere and Martin, 1901b: 172. Type locality: Red Sea, Djibouti.

SUPPLEMENTAL REDESCRIPTION: Male: length 3.35-3.90 mm, maximum width 1.55-1.90 mm. Female: length 3.30-3.60 mm, maximum width 1.50-1.85 mm.

COLORATION: Ground color blackish grey with a bluish tinge, dull, body with fine silvery pubescence, and longer setae (Fig. 1). Head dark above, posterior margin of vertex brown. Pleural and ventral surfaces blackish grey, dull, genital segments black, slightly shining. Antennae brown, basal third of segment I yellow, increasingly embrowned distally. Legs blackish brown; all coxae medium brown; trochanters yellowish brown, darker dorsally and distally; fore femur uniformly brown ventrally, dark brown to black dorsally; mid- and femora yellowish brown at extreme bases, otherwise dark brown; all tibiae and tarsi brown.

Apterous male: body fusiform, length more than 2.0x greater than greatest width across thorax (3.62 : 1.72). Head length 0.28x greatest head width across eyes (0.35 : 1.25); eyes rather small, width about 0.24x interocular width (0.22 : 0.90).

Antennal formula I : II : III : IV; 0.80 : 0.80 : 0.50 : 0.55; segment I almost 3x head length (1.00:0.35), slightly thicker than segments II-IV; combined length of all antennal segments 0.65x total length of insect.

Proportions of legs as follows: Femur, tibia, tarsal 1, tarsal 2, tarsal 3 of fore leg, 1.40 : 1.30 : 0.05 : 0.05 : 0.20; of middle leg, 1.90 : 1.05 : 0.10 :

0.70 : 0.45; of hind leg, 2.00 : 1.00 : 0.10 : 0.60 : 0.50.

Male legs with fore femur incrassate, bearing a stout pointed black tooth at base, angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of 9-10 relatively large, pointed black teeth (Fig. 14A). Fore tibia weakly and gently curved, inner side with a small tubercle in the sharp basal curve, followed at some distance by 2 widely separated teeth, then one additional small, low, broad tooth centrally. Fore and hind trochanters each with 1 small, sharp, ventrally-directed black spine ventrally near distal apices. Middle femur slightly thickened in middle, ventral surface armed with 16-18 equally sized, straight, stout, erect black spines with truncate apices, these spines not interspersed with smaller teeth. Hind femur slightly thickened centrally, hind trochanter and femur unarmed. Middle and hind femora and tibiae bearing numerous long, erect, fine silvery setae, the lengths of these setae exceeding the diameters of the segments from which they arise.

Male metasternum with posterior margin simple, weakly and broadly depressed centrally, lacking posteromedial process, extreme posterior margin bearing a few small, dark papillae (appearing as denticles under all but highest magnification) (Fig. 2A, 3A). Abdomen length from abdominal scent orifice to tip 1.15; genital segments large, length 0.70; styliform processes of segment VIII relatively stout with curved apices (Fig. 14B); male endosoma expanded, with large apicolateral lobes and short medial lobe (Fig. 7C).

Apterous female: Dorsal habitus as in Fig. 13; antennal formula I : II : III : IV; 0.70 : 0.70 : 0.48 : 0.50; segment I less than 1.5x head length (0.70:0.50), slightly thicker than segments II and III, segment IV expanded, thicker than segment I; combined length of all antennal segments 0.69x total length of insect.

Proportions of legs as follows: Femur, tibia, tarsal 1, tarsal 2, tarsal 3 of fore leg, 0.90 : 0.95 : 0.05 : 0.05 : 0.20; of middle leg, 1.50 : 0.90 : 0.10 : 0.50 : 0.45; of hind leg, 1.50 : 0.80 : 0.10 : 0.50 : 0.40. Middle femur with row of ~10 short, sharp black spines along posterior margin.

DISTRIBUTION: Originally described from the Red Sea (Coutiere and Martin, 1901b), with subsequent records from Somalia (China, 1957), Israel (Linnavouri, 1973), the Maldives (Phillips, 1959), Djibouti, Tanzania, Madagascar, Mauritius, Aldabra and Cosmoledo (D. Polhemus, 1990). The broad range of this species encompasses the entire western Indian Ocean (Fig. 21).

DISCUSSION: This is the only species of Hermatobates inhabiting the Western Indian Ocean. Males may be recognized by their moderately large size, the armature of the fore femur with numerous relatively large teeth in the central section (Fig. 14A), the presence of single small black spine ventroapically on both the fore and middle trochanters, the presence of 16 similarly sized stout black spines on the posterior margin of the middle femur with no gaps or smaller spine groups interrupting this row, and the absence of a posteromedial process on the metasternum (Figs. 2A, 3A). Females possess a line of short, sharp dark spines along the posterior margin of the middle femur.

ECOLOGICAL NOTES: This species shelters among coral blocks on tidally emergent reef crests, with adults skating on the deeper fore reef waters beyond, where they are often syntopic with various Halobates species (D. Polhemus, 1990). Because of these habitar preferences, adults are usually best captured by trolling a net from the bow of a moving boat.

A detailed discussion of the habitat preferences and behavior of H. djiboutensis at Aldabra Atoll was provided by D. Polhemus (1990). At Aldabra, it was noted that adults and late instar immatures were typically encountered in the deep water over the fore reef between 500 and 1,000 meters offshore, in areas subject to moderate swell. Individuais were observed running across the sea surface in company with Halobatesflaviventris, and appeared to be more abundant in areas where seaweed and other organic debris formed loose floating mats on the sea surface. Label data on the series from the Maldives notes that the specimens were taken from under a lighted dock, indicating that this species, like many other Hermatobates, is attacted to lights set near the water surface at night.

SPECIMENS EXAMINED: MADAGASCAR: Diego Suarez Prov.: rocky coast along Courrier Bay, 40 km. NW of Diego Suarez, sea level, [12[degrees]13'31"S, 49[degrees]09'18"E], 14 November 1986, CL 2279, D. A. & J. T. Polhemus, 11 [male][male], 4 [female][female] (JTPC). SEYCHELLES: Aldabra Atoll: Picard Is.: calm sea 1,000 m. off of Anse Var, 19 March 1989, D. A. Polhemus, 2 [female][female] (USNM); offshore intertidal reef at Aldabra Research Station, 9[degrees]24'08"S, 46[degrees]12'09"E, 9 March 1989, CL 8024, D. A. Polhemus, 1 [male], 1 [female]) (USNM). Malabar Is.: calm sea 1,000 m. offshore, 12 March 1989, CL 8032, D. A. Polhemus, 2 [male][male], 3 99 (USNM). Grande Terre Is.: 1,000 m. offshore nr. Dune Jean Louis, 9[degrees]27'56"S, 46[degrees]25'55"E, 11:30 hrs., sea surface temp. 28[degrees]C., 8 March 1989, CL 8018, D. A. Polhemus, 1 [male], 8 [male][male] (USNM). Polymnie Is.: calm sea 1,000 m offshore of north coast, 19 March 1989, D. A. Polhemus, 1 [male] (USNM). Cosmoledo Atoll: Menai Is.: calm sea offshore of NE tip, 9[degrees]42'04"S, 47[degrees]32'00"E, 27 March 1989, D. A. Polhemus, 1 [male] (USNM). MAURITIUS: Black River Dist.: Black River Bay [20[degrees]21'49"S, 57[degrees]22'06"E], 19 October 1986, CL 2224, D. A. & J. T. Polhemus, 7 [male][male], 2 [female][female] (JTPC). REPUBLIC OF THE MALDIVES: Maldive Archipelago, Kuramathi Is., under landing stage lights, 15 March 1996, D. Kovac, 7 [male][male], 7 [female][female] (JTPC). TANZANIA, Tanga Prov.: Mkwaja, Sange Island, 10 March 1979, CL 994, J. T. Polhemus, 2 [male][male], 1 [female] (JTPC).

Hermatobates singaporensis, Cheng

Figures 15A, B, 23, 25

Hermatobates singaporensis Cheng, 1976: 209. Type locality: Singapore, Siglap Coast.

DIAGNOSIS: For a detailed morphological description of H. singaporensis see Cheng (1976). Male: length 3.50-3.55 mm, maximum width 1.72.0 mm. Female: length 3.50-3.60 mm, maximum width 1.80-1.90 mm.

COLORATION: Color black with a bluish tinge, dull, body with fine, moderately long silvery pubescence. Head dark above, transversely brown at posterior margin of vertex. Pleural and ventral surfaces dark grey, dull, meso- and metasternum longitudinally brownish centrally, genital segments brown, shiny. Antennae dark brown, basal quarter of segment I pale, increasingly embrowned beyond. Legs dark brown, all femora nearly black dorsally, coxae and trochanters without pale coloration.

Male legs with fore trochanter bearing small tooth ventrally at about two-thirds distance to apex. Fore femur strongly incrassate, ventral surface bearing a slender, pointed black tooth at base angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of 10-12 small triangular black teeth (Fig. 15A). Fore tibia slightly angled on distal quarter, inner margin bearing two low tubercles beyond basal curve, followed by a gap the width of the tubercle, then a broad, black tooth; a small tooth at base of oblique subapical comb of hairs. Trochanter of middle leg with a small, sharp ventral spine near apex. Middle femur slightly thickened centrally, ventral surface bearing a row of ~20 almost straight stout spines, more basal spines with tips spatulate, more distal spines becoming progressively smaller in size. Hind femur slightly thickened centrally, somewhat dorsoventrally compressed on distal half; hind trochanter, femur and tibia unarmed.

Female legs with fore femur bearing a longitudinal row of ~20 tiny black triangular teeth along ventral margin, this margin also set with numerous long, erect, pilose pale setae; fore trochanter and tibia unarmed; middle femur with posterior margin bearing at most 8 very small, inconspicuous dark spinules, or unarmed.

Male metasternum with posterior section angled gently inward toward base of genital segment, posterior margin with a broad, trapezoidal posteromedial process bearing scattered large papillae, this process recessed (Fig. 15B), hidden by genital segment when latter retracted; styliform processes of abdominal segment VIII slender, with apices strongly hooked; male endosoma with expanded lateral lobes and an elongate medial lobe.

DISTRIBUTION: Originally described from Singapore, the specimens having been taken near a floating fishing platform about 5 kilometers offshore of the Siglap coast, on the eastern side of the island (Cheng, 1976). We provide additional records below from Peninsular Malaysia (Fig. 23).

This is a rarely collected species that appears to have a restricted range in the vicinity of the Straits of Malacca and northward along the coasts of Peninsular Malaysia. The absence of Hermatobates collections from the western portion of the Malay Archipelago currently precludes evaluation of potential intergrading morphoclines between this species and H. weddi, which has a range lying further to the southeast, in the Lesser Sunda Islands, Australia and the southwest Pacific. In addition, Andersen (1989: 206) mentions the presence of Hermatobates immatures and exuviae at Phuket Island, Thailand, and we consider it likely that this record may represent H. singaporensis, given that it would be a logical northwestern extension of the species' range from currently known localities, but this must be confirmed by future collection of adult specimens from Phuket.

DISCUSSION: Hermatobates singaporensis is a western segregate of the H. weddi group; it is most easily separated from other members of the H. weddi group by the median process of the male metasternum, which is very weakly formed and slightly sinuate along its posterior edge, located at the posterior margin of the depressed metasternum (Fig. 15B), and not nearly as prominent as in H. weddi, marchei, kula, armatus or schuhi (Figs. 2A-D, 19). The dentition of the fore femur (Fig. 15A) and tibia, which were used by Cheng (1976) as key characters for this species, are very similar to the character states seen in H. weddi, the latter taxon being somewhat variable in regard to this character state over its wide range; as such, they are unreliable characters for species separation. Females of H. singaporensis also bear 8 very small black spinules along the posterior margin of the middle femur, whereas such spinules are absent or incipient in H. weddi, which possesses at most only a few extremely tiny, barely visible spinules. Hermatobates weddi is also a slightly larger species, with males having a body length of 3.40-4.00 mm, versus 3.50-3.60 mm in H. singaporensis. Further collections from intervening localities on the islands of Bali, Java and Sumatra in the western Malay Archipelago may eventually indicate that this species is a synonym of 14. weddi, but such a conclusion is deferred herein.

Cheng (1976) provided line drawings of the dorsal body habitus, and of the male and female forelegs, as well as small scale black-and-white photographs of the male in dorsal and ventral view. Her holotype of H. singaporensis from which these photographs were taken was deposited in the Smithsonian Institution by Cheng, but after extensive unsuccessful search for it in that museum's collections is considered lost. A neotype is herein designated from St. John Island, Singapore, not far from the original Siglap type locality.

ECOLOGICAL NOTES: Our neotype female from Singapore was collected at St. John Island, a small, rocky island immediately adjacent to the Singapore Straits. The specimen was taken on northern side of the island, where a rocky shore exposes at low tide, in company with other immature Hermatobates, Halobates hayanus White, Halovelia lannae Andersen, and a few Haloveloides sundaensis Andersen. The samples here were made from late afternoon until dusk, with the tide falling throughout the sampling period.

SPECIMENS EXAMINED: SINGAPORE: Siglap, 1[degrees]17'N, 103[degrees]52'E, 3 October 1963, L. Cheng, 2 99 (allotype and paratype designated by Cheng, USNM); St. John Island, NE side, sea level, sea surface temp. 31[degrees]C., salinity 30 ppt, 1[degrees]13'22"N, 103[degrees]50'37"E, 25 October 2010, 15:00-18:00 hrs., falling tide to daily low, J. T. & D. A. Polhemus 1 [female], 2 immatures (neotype, USNM). MALAYSIA: Negiri Sembilan: 12 km. S. of Port Dickson, in lagoon pools at low tide, 16 March 1983, R. T. Schuh & B. Massie, 3 [male][male], 1 [female] (JTPC); 10 km. S. of Port Dickson [2[degrees]27'45"N, 101[degrees]50'56"E], 15 August 1985, CL 2060, D. A. & J. T. Polhemus, 2 [male][male], 2 [female][female]) (JTPC). Pahang: East Coast, Pulau Tioman, rock pool at low tide, 90ctober 1985, L. Cheng, 2 [male][male], 2 [female][female] (JTPC).

Hermatobates weddi, China

Figures 2F, 3D, 4B, 5D, 6B, 7A, 16, 17A, B, 18D, 20, 23, 25

Hermatobates weddi China, 1957: 346. Syn by Andersen and Weir, 2000: 335. Restored from synonymy by J. Polhemus and D. Polhemus, 2006: 948

DIAGNOSIS. For a detailed morphological description of this taxon see China (1957), and Andersen and Weir (2000, as H. marchei). Male length 3.40-4.00 rum, maximum width 1.70-2.00 mm; female length 3.3-3.50 mm, maximum width 1.70-1.80 mm.

Color black with a bluish tinge, dull, body with fine silvery pubescence (Fig. 16). Head dark above, transversely pale at posterior margin of vertex. Pleural and ventral surfaces brownish, dull, genital segments dark brown, shiny. Antennae blackish brown, basal third of segment I pale, increasingly embrowned beyond. Legs brown, except fore coxae and trochanters, ventral surface of fore femur, fore tibia other than extreme base, basal halves of middle and hind trochanters pale.

Male legs with fore trochanter bearing a small tooth ventrally near apex. Fore femur strongly incrassate (Fig. 16), ventral surface bearing a stout pointed black tooth at base angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of 10-16 small black teeth. Fore tibia with inner margin bearing two low tubercles beyond basal curve, followed by a broad, black tooth (Fig. 5D), then a row of small tubercles leading to a small black tooth about two thirds distance from base to apex; several small tubercles present near apex; a small tooth at base of oblique subapical comb of hairs. Trochanter of middle leg with a small ventral spine near apex. Middle femur slightly thickened centrally, ventral surface bearing a row of ~20 socketed, spatulate, paddle-like spines (17A, B). Hind femur slightly thickened centrally, somewhat dorsoventrally compressed in larger specimens; hind trochanter, femur and tibia unarmed.

Female legs with fore femur bearing a row of 15-18 small black teeth along ventral margin, fore trochanter and tibia unarmed. Middle trochanter unarmed; middle femur with a few tiny, barely visible black teeth basally on ventral margin.

Male metastemum with a broad, trapezoidal posteromedial process arising from center of posterior margin (Figs. 2F, 3D), bearing scattered large papillae (Fig. 4B); styliform processes of abdominal segment VIII broad, with hooked apices (Fig. 6B); male endosoma with moderately developed basolateral and apicolateral lobes, medial lobe elongate with apex bidentate (Figs. 7A, 18D).

DISTRIBUTION: Originally described from the Monte Bello Islands, Western Australia (China, 1957), with subsequent records from elsewhere in Australia (Northern Territory; Queensland), New Caledonia, Papua New Guinea (south coast; D'Entrecasteaux Islands; Engineer Group), and East Timor (Andersen and Weir, 2000; J. Polhemus and D. Polhemus, 2006). Mention was also made by J. Polhemus and D. Polhemus (2006) of additional occurrences in Indonesia (Lesser Sunda Islands), Tonga, Fiji, and the Solomon Islands, but without supporting data, which we now provide.

The documented range of H. weddi now extends from the Lesser Sunda Islands and the Indian Ocean coast of Australia eastward through the Arafura Sea, Torres Strait, and Coral Sea to southern New Guinea, the Solomon Islands, and New Caledonia, and then eastward into the insular Pacific to Fiji, Tonga, and Samoa (Figs. 20, 23, 25). This broad distribution lies generally to the south of that occupied by H. marchei, which occurs from the Philippines through the Carolines to the northern coasts of New Guinea (J. Polhemus and D. Polhemus, 2006), with the two species occupying a syntopic contact zone in far eastern New Guinea, where they also co-occur with H. kula (J. Polhemus and D. Polhemus, 2006). Previous records of H. weddi from the islands of Okinawa, Ishigaki, Iriomote, Amami-oshima, and the Tokara Islands by Azuma and Kinjo (1987) refer instead to the new species H. schuhi described herein. The detailed Australian range plot for H. marchei in Andersen and Weir (2000) actually depicts the distribution of H. weddi, and all their Australian records for H. marchei in fact refer to H. weddi. We have not repeated this extensive set of Australian records herein, and readers are instead referred to Andersen and Weir (2000) for detailed information regarding the distribution of H. weddi around the coasts of Australia (see Fig. 29 in Andersen and Weir, 2000).

DISCUSSION: All species of the Hermatobates weddi group have very similar facies, differing primarily in the morphology of the metasternal process, male fore femur and tibia, and male genitalia. This subtlety of differentiation resulted in H. weddi being synonymized under H. marchei by Andersen and Weir (2000), presumably because these authors had no proper specimens of H. marchei for comparison.

Hermatobates weddi males can be separated from both H. marchei and H. armatus by the trapezoidal rather than parallel-sided shape of the posteromedial metasternal process (compare Fig. 2C, F), and by the more angular and less developed apicolateral lobes on the male endosoma (compare Fig. 18B, D). It may be separated from H. kula by the shape of the male endosoma, with H. weddi having more weakly developed and less angulate lateral lobes but a longer medial lobe (compare Fig. 18A, D); by the structure of the fore tibia, which lacks a tubercle in the basal curve, has more broadly separated basal teeth beyond the basal curve (compare Fig. 5A, D), and often possesses a small tooth distally ar about two thirds the distance to the apex. The metasternal processes are similar in both H. weddi and H. kula, but that of H. kula arises slightly anterior to the posterior margin of the metasternum, rather than at the posterior margin as seen in H. weddi (compare Fig. 2E, F). The species is close in many respects to H. schuhi, bur can be separated on the basis of the male endosoma, which has the basolateral lobes far more expanded and explanate in H. schuhi (compare Fig. 18C, D); by papillae on the male metasternal process, which are far larger and less numerous in H. weddi (compare Fig. 4A, B); and by geography, with H. weddi occurring around Australia and in the Southwest Pacific (Figs. 20, 23, 25), and H. sehuhi occurring in the Ryukyu Archipelago (Figs. 23, 25). Females in the H. weddi group are difficult to identify without associated males, although the pale coloration of the trochanters will often distinguish those of H. kula within this assemblage, at least for populations occurring in eastern New Guinea and adjacent archipelagoes (see additional discussion under H. kula).

Because Hermatobates weddi has a very broad geographic range, extending from the Lesser Sunda Islands to Tonga, with consequent intraspecific variation in morphology, future research, particularly using molecular characters, may indicate that our current species concept in fact encompass several geographically segregated species.

China (1957) provided useful line drawings of the ventral male body habitus and male forelegs for H. weddi. The subsequent illustrations of this species in Andersen and Weir (2000) are a mix of character states for both H. weddi and H. marchei, which these authors considered synonymous, and should be used with caution. The illustrations in J. Polhemus and D. Polhemus (2006), by contrast, clearly depict certain morphological differences between the two species, and these have been further supplemented herein.

ECOLOGICAL NOTES. The ecology and behavior of H. weddi were investigated in detail by Foster (1989), based on work in Fiji, Samoa, Tonga and the Cook Islands. He concluded that H. weddi occupied a zone approximately 20 cru to either side of Mean Low Water Neaps along suitable shorelines with coral rock containing cavities to which the insects could retreat at high tide. At low tide the insects would emerge from these refugia to skate on the reef crest and waters immediately seaward, with activity centered on a period from one hour before to one hour after dead low tide. This general behavior pattern was also observed by the authors for H. weddi on the reef offshore of Fakfak in western New Guinea.

By contrast, the second author found immatures of H. weddi to be abundant on calm, shallow backreef waters at Fatumafuti Rock near Pago Pago, on the island of Tutuila in American Samoa, but no adults were captured at this locality despite several hours of effort at low tide. This, coupled with the overwhelming predominance of immatures in samples from adjacent Upolu, indicates that adults of the Samoan populations may skate on deeper waters beyond the reef crest, similar to the pattern documented for H. djiboutensis at Aldabra Atoll (D. Polhemus, 1990). These observations are congruent with other results summarized by Andersen and Weir (2000) who concluded that H. weddi and H. armatus were, like H. djiboutensis, capable of remaining active between low and high tide cycles if sea surface conditions on waters beyond the fore reef remained sufficiently calm.

SPECIMENS EXAMINED: AUSTRALIA: Queensland: Iron Range, Chili Beach, nr. Portland Roads [12[degrees]37'00"S, 143[degrees]25'39"E], 24 August 1983, CL 1954, D. A. & J. T. Polhemus, 2 [male][male] (JTPC); Magnetic Is., Cockle Bay, 9 July 1976, L. Cheng, 1 [female]) (JTPC); Prince of Wales Island, Torres Strait, intertidal reef, 30 August 1983, CL 1769, D. A. & J. T. Polhemus, 13 [male][male], 12 [female][female] (JTPC). Terr. of Aslunore & Cartier Islands: Ashmore Reef, West Is., anchorage, at night, 20 September 1987, H. Larson, 2 [male][male] (JTPC). INDONESIA, Nusa Tenggara Barat Prov.: Sumbawa Is., Kab. Bima, coast 5 km. E. of Nipa [8[degrees]17'15"S, 118[degrees]47'00"E], low tide, 21 October 1985, CL 2176, D. A. & J. T. Polhemus, 13 [male][male], 11 [female][female] (JTPC). West Papua Prov.[=former Irian Jaya Prov.]: New Guinea, offshore reef below Fakfak airport, 5 km. N. of Fakfak [2[degrees]55'42"S, 132[degrees]15'00"E], sea surface temp. 30.5[degrees]C., low tide, 11 October 1991, CL 2636, D. A. & J. T. Polhemus, 19 [male][male], 13 [female][female] (JTPC). EAST TIMOR, Dili Dist.: exposed reef crest at Bekarit, Areia Branca Bay, 4.5 km. E. of Dili, 0 m. [sea level], 8[degrees]32'21"S, 125[degrees]36'48"E, sea temp. 27[degrees]C., 1 September 2004, 17:30-18:30 hrs., CL 7265, D. A. Polhemus, 2 [male][male], 1 [female] (USNM). PAPUA NEW GUINEA, Milne Bay Prov.: D'Entrecasteaux Islands, Fergusson Island, east coast, sea 100 m. offshore of N. side of Mebulibuli Point, 9[degrees]30'18"S, 150[degrees]52'53"E, sea surface temp. 29[degrees]C., salinity 36 ppt., 26-27 January 2003, 18:30-00:30 hrs. each night (low tide near midnight), CL 7223, D. A. & J. T. Polhemus, 1 [male] (JTPC); D'Entrecasteaux Islands, Normanby Island, Bunama Bay, sea 60 m. offshore, 10[degrees]08'38"S, 151[degrees]09'12"E, salinity 36 ppt., sea surface temp. 29[degrees]C., 28 January 2003, 18:30-13:00 hrs., CL 7227, D. A. & J. T. Polhemus, 11 [male][male], 27 [female][female], 112 nymphs (JTPC); D'Entrecasteaux Islands, Normanby Island, east coast, Kasikasi Bay, sea 200 m. offshore, 10[degrees]09'04"S, 151[degrees]13'56"E, salinity 36 ppt., sea surface temp. 28[degrees]C., 29 January 2003, 19:00-00:30 hrs., CL 7229, D. A. & J. T. Polhemus, 3 [male][male], 6 [female][female], 15 nymphs (JTPC); Tubetube Is., Engineer Group, bay on NE side, 10[degrees]35'13"S, 151[degrees]12'12"E, sea surface temp. 29[degrees]C., salinity 36 ppt., 18 January 2004, 20:00-24:00 hrs., CL 7298, D. A. & J. T. Polhemus, 3 [female][female]), 1 nymph (JTPC); Nuakata Is., Dudawali Bay, 10[degrees]17'36"S, 151[degrees]00'33"E, sea surface temp. 30[degrees]C., salinity 36 ppt., 19 January 2004, 20:00-24:00 hrs., CL 7301, D. A. & J. T. Polhemus, 1 [male], 4 [female][female], 2 nymphs (JTPC). SOLOMON ISLANDS, Western Prov.: New Georgia Is., Munda, night light, 1 April 1983, L. Cheng, 1 [male], 1 [female] (JTPC); Gizo Is., mangrove-lined bay on north coast, W. of Gizo, 8[degrees]02'41"S, 156[degrees]48'32"E, sea temp. 31[degrees]C., salinity 36 ppt., 13 Match 2005, 20:00-22:00 hrs., CL 7380, D. A. Polhemus, and J. T. Polhemus, 1 [male], 1 immature (JTPC); Mbanga Is., western shore near Goldie College, 8[degrees]16'27"S, 157[degrees]12'36"E, sea surface temp. 31[degrees]C., salinity 36 ppt., 20 March 2005, CL 7398, D. A. Polhemus, and J. T. Polhemus, 1 [female], 12 immatures (JTPC). Choiseul Prov.: Choiseul Is., Choiseul Bay near Parama Island, 6[degrees]41' 18"S, 156[degrees]24'20"E, sea surface temp. 31[degrees]C., salinity 34 ppt., 16 March 2005, 19:30-23:00 hrs., CL 7388, D. A. Polhemus and J. T. Polhemus, 1 [female], 2 immatures (JTPC); Choiseul Is., south coast, Mole harbor, 6[degrees]49'34"S, 156[degrees]31'13"E, sea surface temp. 31[degrees]C., salinity 34 ppt., 17 March 2005, 19:00-22:00 hrs., CL 7391, D. A. Polhemus and J. T. Polhemus, 1 [male], 13 [female][female], 1 immature (JTPC). NEW CALEDONIA, South Prov.: Noumea Bay, day, low tide, 18 April 1976, L. Cheng, 2 [male][male], 1 [female] (JTPC); Noumea, rocky beach, 27 September 1983, CL 1867, D. A. & J. T. Polhemus, 1 [male] (JTPC). FIJI, Navosa Prov.: Viti Levu Is., Naviti Resort Hotel, 28 July 1978, L. Cheng, 3 [female][female] (JTPC). Rewa Prov.: Viti Levu Is., Suva, main reef, 16 February 1969, J. S. Tobler, 1 [male], 1 [female] (JTPC); Viti Levu Is., Suva Bay, incoming tide, 9 July 1983, W. A. Foster, 7 [female][female] (CUMZ); same data as preceding except 4 July 1983, 7 [male][male], 1 [female] (JTPC). Ba Prov.: 1 [male], 1 [female], Viti Levu Is., Vuda, low tide, coral, 19 August 1986, W. A. Foster, 5 [male][male], 2 [female][female] (CUMZ); same data as preceding except 18 August 1986, 2 [male][male], 2 [female][female], 2 immatures (CUMZ). Kadavu Prov.: Dravuni Is., Astrolabe Reef, 25 July 1978, collector unknown, 1 [male] (JTPC). Lau Prov.: Lakeba Is., Tubou, 4 July 1983, W. A. Foster, 1 [male] (CUMZ); Vanua Balavu, Lomaloma Dist., reef flat S. of Wailevu, 17[degrees]18"13'S. 178[degrees]59'20"E, sea surface temp. 26.5[degrees]C., 21 August 2005, 12:30-14:30 hrs., CL 7450, J. T. Polhemus, 1 [male], 1 [female] (JTPC). TONGA: Nuka'alofa, Jetty 22, 17 April 1976, L. Cheng, 3 99 (JTPC); reef side of Jetty #2, 17 April 1976, L. Cheng, 1 [male], 8 [female][female] (JTPC); Nukualofa Is., 21:00 hrs, at light, 12 August 1986, W. A. Foster, 5 [male][male], 1 [female] (CUMZ).

OTHER SPECIMEYS EXAMINED: The following series are also provisionally assigned to H. weddi for the present in order to note the potential presence of this species in the Samoa and Cook Island groups, but with the caution that these determinations require subsequent validation by examination of males from the localities involved. COOK ISLANDS: Rarotonga: Rarotonga, 4 July 1986, W. A. Foster, 1 9 (CUMZ). SAMOA: Upolu: Apia, 11:00 hrs., low tide, rock, 31 August 1986, W. A. Foster, 1 9 (CUMZ); same data as preceding except 21 July 1986, 3 99, 20 immatures (JTPC). AMERICAN SAMOA: Tutuila: rocky coast at Fatu Rock, nr. Fatumafuti, 14[degrees]17'45"S, 170[degrees]40'38"W, 22 March 2009, CL 7465, D. A. Polhemus, 13 immatures (BPBM).

Hermatobates marchei, Coutiere and Martin

Figures 2C, 3C, 4C, 5B, 6D, 18B, 23, 24, 25

Hermatobates marchei Coutiere and Martin, 1901c: 215.

DIAGNOSIS. For previous discussion of this species see J. Polhemus and D. Polhemus (2006). Male: length 3.30-3.70 mm, maximum width 1.80-1.90 mm. Female: length 3.50-3.70 mm, maximum width 1.80-2.00 mm.

COLORATION: Color dark grey, dull, body with fine silvery pubescence, and longer setae. Head dark grey above, narrowly brown in transverse band across extreme posterior vertex. Pleural areas grey, dull, central portion of thoracic venter trending to brown, genital segments brown, shining. Antennae black, antennal segment I with basal third yellowish brown. Legs uniformly black dorsally, unformly brown ventrally.

Male legs with fore femur incrassate, bearing a stout pointed black tooth at base, angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of about 14 tiny black teeth. Fore tibia moderately curved, inner side with a distinct tubercle in sharp basal curve, followed after a short distance by a broad, stout, bidentate tooth, then a short space, then a larger apically truncate tooth (Fig. 5B). Trochanters of fore and middle legs each with small ventral spine distally. Middle femur slightly thickened in middle, ventral surface armed with about 20 almost straight stout spines, those near base with tips hooked, those lying more distally interspersed with shorter spines. Hind femur without dorsoventral flattening. Middle and hind femora and tibiae bearing numerous long, erect, fine silvery setae, the lengths of these setae exceeding the diameters of the segments from which they arise.

Female legs with fore femur bearing a row of about 17 small dark denticles on the ventral margin; middle femur with a row of about 12 very small dark spines on basal two-thirds.

Male metasternum with posterior margin bearing a quadrate process centrally (Figs. 2C, 3C) bearing prominent conical papillae (Fig. 4C); styliform processes of abdominal segment VIII relatively slender with strongly hooked apices (Fig. 6D); male endosoma with strongly rounded and inflated apicolateral lobes (Fig. 18B).

DISTRIBUTION: Originally described from a mixed series of poorly labelled specimens taken in the Philippines, some of which came from Palawan (Coutiere and Martin, 1901; Andersen and Weir, 2000). Subsequent records were provided for other areas in the Philippine Islands (Mindanao), Palau, New Guinea (Morobe, Oro and Milne Bay Provinces of Papua New Guinea), and the D'Entrecasteaux Islands (Fergusson) by J. Polhemus and D. Polhemus (2006). As currently understood, the range of this species encompasses the Philippines, the Caroline Islands, and the New Guinea region, but does not extend further southeast into the insular Pacific or to Australia (Figs. 23, 24, 25). Previous records of H. marchei from the Ryukyu Archipelago by Takara 0957) refer instead to the new species H. schuhi described herein. The Australian range plot provided for H. marchei by Andersen and Weir (2000) in fact represents the distribution of H. weddi around the margins of that continent.

DISCUSSION: Hermatobates marchei falls within our H. weddi species group. Males are most easily separated from the other members of that group by the spatulate, nearly quadrate metastemal process which is distinctly separated from the rounded posterior margin of the metasternum (Figs. 2C, 3C) and bears large, conical papillae (Fig. 4C); by the slender styliform process of the male genital complex with a hooked tip (Fig. 6D); and by the endosoma with its strongly expanded and rounded apicolateral lobes (Fig. 18B). The middle femur is also not dorsoventrally flattened as in H. weddi, and antennal segment I is light colored only on the basal 1/4 to 1/3, darkening rapidly toward the middle, whereas in H. weddi the first antennal segrnent is light colored except distally, and in H. armatus only the base is light colored.

As noted by J. Polhemus and D. Polhemus (2006), this species was originally described from the Philippines, but the types appear to be lost. Although reportedly in the Paris Museum, searches for them in that collection by the authors proved unsuccessful. A male from Mindanao was therefore designated as a neotype and is held in the USNM (J. Polhemus and D. Polhemus, 2006).

The illustrations of this species in Andersen and Weir (2000) actually depict H. weddi, which these authors incorrectly considered a synonym of H. marchei (J. Polhemus and D. Polhemus, 2006). By contrast, the illustrations in J. Polhemus and D. Polhemus (2006) and herein unambiguously depict the morphological differences between the two species.

SPECIMENS EXAMINED: PHILIPPINE ISLANDS, Zamboanga deL Sur Prov.: Santa Cruz Island off Zamboanga [Mindanao], coral reef [6[degrees]52'06"N, 122[degrees]04'06"E], 21 July 1985, CL 1997, J. T. & D. A. Polhemus, 1 [male], neotype, plus 1 [male], 1 [female], 5 immatures (JTPC). PALAU, Koror State: Koror Island, T-Dock at Koror waterfront, 7[degrees]21'07"N, 134[degrees]28'42"E, sea surface temp. in water pockets amid coral rubble 30[degrees]C., 8 August 1999, 11:00-12:00 hrs. (falling tide); 9 August 1999, 12:00-13:00 hrs. (falling tide); 10 August 1999, 14:00-15:30 hrs. (falling tide); 11 August 1999, 15:00-16:30 hrs. (falling tide), CL 7126, D. A. and J. T. Polhemus, 17 [male][male], 11 [female][female], 2 immatures (USNM). INDONESIA, West Papua Prov.: Roon Island, Cenderawasih Bay, calm sea 200 m. offshore of Yende village, 2[degrees]21'42"S, 134[degrees]31'49"E, sea surface temp. 30[degrees]C., salinity 35 ppt., 13 November 2008, 20:30-22:00 hrs., CL 7544, D. A. Polhemus, 1 [male], 3 [female][female] (BPBM). PAPUA NEW GUINEA, Milne Bay Prov.: New Guinea, sea offshore of W. side of Lihitabu Point, N. sitie of Milne Bay, E. of Alotau, 10[degrees]21'13'S, 150[degrees]35'05"E, sea surface temp. 29[degrees]C., salinity 33 ppt., 8 April 2002, 16:00-16:30 hrs., CL 7168, D. A. & J. T. Polhemus, 1 [male], 3 [female][female], 2 nymphs (USNM, BPBM, JTPC); D'Entrecasteaux Islands, Fergusson Island, east coast, sea 100 m. offshore of N. side of Mebulibuli Point, 9'30'18"S, 150 52'53"E, sea surface temp. 29[degrees]C., salinity 36 ppt., 26-27 January 2003, 18:30-00:30 hrs. each night (low tide near midnight), CL 722L D. A. & J. T. Polhemus, 18 [male][male], 67 [female][female], 10 nymphs (USNM, BPBM, JTPC). Morobe Prov.: 1 [male], 4 [female][female], Cape Roon, 150 m. offshore, 7[degrees]20'45"S, 147[degrees]09'33"E, sea surface temp. 30[degrees]C., salinity 29 ppt., 2 May 2003, 10:00-10:30 hrs., CL 7237, D. A. Polhemus (USNM); rocky point N. of Cape Dinga [=Yaraku Point], 7[degrees]18'04"S, 147[degrees]08'35"E, sea surface temp. 30[degrees]C., salinity 30 ppt., 4 May 2003, 10:00-12:30 hrs., CL 7242, D. A. Polhemus, 1 [male] (USNM). Oro Prov.: Tufi Harbor, cliff-bound rocky shore near village dock, 9[degrees]04'57"S, 149[degrees]19'15"E, 24 January 2004, 23:00-24:00 hrs., CL 4468, J. T. Polhemus, 3 [male][male], 1 [female], 4 nymphs (JTPC).

Hermatobates kula, J. Polhemus and D. Polhemus Figures 2E, 2E, 4D, 5A, 6C, 18A, 24

Hermatobates kula J. Polhemus and D. Polhemus, 2006: 950.

DIAGNOSIS: For detailed description of this taxon see J. Polhemus and D. Polhemus (2006). Male: length 3.44-4.00 mm maximum width 1.61-1.94 mm. Female: length 3.61-3.83 mm, maximum width 1.89-2.11 mm.

COLORATION: Color black with a bluish tinge, dull, body with fine silvery pubescence, and longer setae. Head dark above, with a transverse line of pale spots at base. Pleural and ventral surfaces brownish, dull, genital segments blackish, shiny. Antennae blackish brown, basal third to half luteous, increasingly embrowned beyond. Legs blackish brown, fore coxae partly luteous; fore trochanters, basal part of fore femora luteous; mid- and hind trochanters partly testaceous in males, mostly luteous in females, slightly darkened dorsally and posteriorly on distal part; fore tibia yellowish brown to luteous.

Male legs with fore femur incrassate, with a stout pointed black tooth at base, angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of 11-14 small black teeth. Fore tibia moderately curved, inner side with a distinct tubercle in sharp basal curve, followed at some distance by a short stout tooth separated by a short distance from a larger tooth (Fig. 5A), then a row of small tubercles ending with a step-like declevity at about two thirds of tibia length from base, this declivity lacking any small tooth, several small tubercles also present on inner surface near apex; a small tooth present on outer margin near tip at base of oblique comb of hairs. Trochanter of middle leg with distinct ventral spine distally. Middle femur slightly thickened in middle, ventral surface armed with 19-21 almost straight stout spines, those near base with tips hooked, interspersed by shorter teeth. Hind femur slightly thickened on distal third, dorsoventrally compressed in large specimens; trochanter and femur unarmed. Middle and hind femora thickly set with long silvery setae (0.22).

Female legs with fore femur bearing a row of 16 very small black teeth along ventral margin, fore trochanter and tibia unarmed. Middle trochanter unarmed; middle femur with ~14 very small black teeth along ventral margin.

Metasternum with posterior margin bearing a trapezoidal posteromedial process arising slightly anterior to poserior margin of the metasternum (Figs. 2E, 3E). Abdomen very short, length from abdominal scent orifice to tip 0.97; genital segments large, length 0.71; styliform processes of segment VIII relatively slender with small, hooked apices (Fig. 6C); male endosoma with expanded, angulate lateral lobes (Fig. 18A).

DISTRIBUTION: Originally described from specimens taken in the Louisiade Archipelago at Woa Island, in the Rossel Island lagoon (J. Polhemus and D. Polhemus, 2006), with paratypes from elsewhere in the Louisiades (Misima, Panatinanae), the D' Entrecasteaux Islands (Normanby, Fergusson), and other small islands near China Strait. Based on current collections, this species appears to be endemic to far eastern New Guinea and the small archipelagoes lying immediately to the east (Fig. 24), a set of land masses composing the tectonically defined East Papua Composite Terrane, which is a known area of endemism for other genera and species of aquatic Heteroptera (D. Polhemus and J. Polhemus, 2004).

DISCUSSION: Males of Hermatobates kula may be separated from those of H. marchei by the trapezoidal rather than quadrate metasternal process (compare Fig. 2C, E). The trapezoidal shape of the process is similar to the character state in H. weddi, but in H. kula the posterior portion of the metasternum is tumid and the process arises ahead of the posterior metasternal margin (Figs. 2E, 3E), whereas in H. weddi the posterior half of the metasternum forms a flat plate tilted evenly toward the base of the genital cavity, with the metasternal process arising at the posterior margin (compare Figs. 6 and 7, and Figs. 27 and 31 in J. Polhemus and D. Polhemus, 2006). In addition, the fore tibia of H. kula bears a tubercle in the basal curve and two stout teeth in close proximity on the basal hall (Fig. 5A), and the styliform abdominal processes are slender with small, hooked tips (Fig. 6C), whereas in H. weddi the tooth in the inner curve is incipient or absent

(Fig. 5D), and the shafts of the styliform processes are stouter, with the apices more strongly hooked (compare Fig. 6B). The male fore tibia in H. weddi also frequently possesses a small tooth at about two-thirds the distance to the apex on the inner margin, which is absent in H. kula. The everted endosoma also shows clear differences, with that of H. kula having more highly expanded and angulate lateral lobes, and a shorter medial lobe (compare Fig. 18A, D).

Females of H. kula may be separated from those of H. weddi by the presence of ~14 very small black teeth along posterior margin of the middle femur, whereas in H. weddi the middle femur bears at most only a few tiny, inconspicuous black teeth. Separation from females of H. marchei is more equivocal, with the middle femur in H. marchei also bearing a row of about 12 very small dark spines on basal two-thirds, but in H. marchei only the extreme base of the first antennal segment is luteous, rather than the basal half, and the trochanters are dark rather than pale. Female color characters were also used by J. Polhemus and D. Polhemus (2006) to distinguish H. kula from H. weddi, but as noted in that publication these characters are only applicable in the eastern New Guinea region where the geographic ranges of H. kula, H. weddi, and H. marchei co-mingle in confusing multi-species assemblages. Females of more eastern populations of H. weddi, such as those occurring in Tonga, by contrast display pale coloration on the basal leg segments similar to that seen in H. kula, and therefore require association with males for definitive identification, although all of these more pale colored H. weddi morphs originate from areas far to the east of the known range of H. kula.

SPECIMENS EXAMINED: PAPUA NEW GUINEA: Milne Bay Prov.: Louisiade Archipelago, Station 13, Wola Island, Rossel Lagoon, north side, 11[degrees]18'05"S, 154[degrees]00'51"E, sea surface temp. 25.5[degrees]C., 1 September 2002, 19:00-22:30 hrs., at light, CL 7199, D. A. & J. T. Polhemus, 27 [male][male], 33 [female][female], 11 immatures (holotype and paratypes, USNM, JTPC, BPBM); Louisiade Archipelago, Calvados Chain, Panatinane Island, Huniye Bay, 11[degrees]16'22"S, 153[degrees]10'50"E, sea surface temp. 26[degrees]C., salinity 36 ppt., 28 August 2002, 19:00-20:00 hrs., at light, CL 7189, D. A. & J. T. Polhemus, 4 [male][male], 10 [female][female], 2 immatures (USNM, JTPC, BPBM); Louisiade Archipelago, Tagula [Sudest] Island, sea offshore of Mohuwo Point [Nepenthes Point], 11[degrees]21'59"S, 153[degrees]16'29"E, sea surface temp. 25.5[degrees]C., salinity 36 ppt., 29 August 2002, 19:00-20:00 hrs., at light, CL 7193, D. A. & J. T. Polhemus, 2 [male][male], 4 [female][female] (USNM, JTPC, BPBM); Louisiade Archipelago, Misima Island, Bwagoia harbor, 10[degrees]41'16"S, 152[degrees]50'46"E, sea surface temp. 26[degrees]C., salinity 36 ppt., 2 September 2002, 19:00-20:00 hrs., at light, CL 7200, D. A. & J. T. Polhemus, 2 [male][male], 8 [female][female], 4 immatures (USNM, JTPC, BPBM); Conflict Group, Irai Island, north side, 10[degrees]45'57"S, 151[degrees]42'29"E, sea surface temp. 26[degrees]C., salinity 36 ppt., 4 September 2002, 15:00-20:00 hrs., CL 7213, D. A. & J. T. Polhemus, 9 [male][male] 6 [female][female], 2 immatures (USNM, JTPC, BPBM); D'Entrecasteaux Islands, Fergusson Island, east coast, sea 100 m offshore of N side of Mebulibuli Point, 9[degrees]30'18"S, 150[degrees]52'53"E, Station 25, sea temp. 29[degrees]C, salinity 36 ppt, 26-27 January 2003, 18:30-00:30 hrs. each night (low tide near midnight), night light from boat, CL 7223, D.A. & J.T. Polhemus, 2 [male]male] (USNM); D'Entrecasteaux Islands, Normanby Island, Bunama Bay, sea 60 m. offshore, 10[degrees]08'38"S, 151[degrees]09'13"E, salinity 36 ppt., sea surface temp. 29[degrees]C., 28 January 2003, 18:30-13:00 hrs., CL 7227, D. A. & J. T. Polhemus, 1 [male], 1 [female] (USNM,); Basilaki Is., Hemoe Bay, 10[degrees]35'41"S, 150[degrees]59'57"E, sea surface temp. 29.5[degrees]C., salinity 36 ppt., 17 January 2004, 20:30-23:00 hrs., CL 7295, D. A. & J. T. Polhemus, 1 [female] (USNM). Central Prov.: Motupore Island, [9[degrees]31'21"S, 147[degrees] 17'03"E], 21 September 1983, CL 1839, D. A. & J. T. Polhemus, 1 [male], 5 [female][female] (JTPC).

Hermatobates armatus, Andersen and Weir

Figures 19, 24

Hermatobates armatus Andersen and Weir, 2000: 340.

DIAGNOSIS: For a detailed morphological description of this taxon see Andersen and Weir (2000). Male: length 3.33-3.65 mm, maximum width 1.50-1.68 mm. Female: length 3.51-3.70 mm, maximum width 1.65-1.80 mm.

COLORATION: Color black with a bluish tinge, dull, body with fine silvery pubescence. Head dark above, with transverse set of pale spots at posterior margin of vertex. Pleural and ventral surfaces brownish, dull, genital segments black, shiny. Antennae dark blackish brown, basal fifth of segment I pale, increasingly embrowned beyond. Legs brown, except ventral surfaces of trochanters, fore tibia, and tarsi pale.

Male legs with fore trochanter bearing small tooth ventrally near apex. Fore femur incrassate, ventral surface bearing a stout pointed black tooth at base angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of 12-14 small black teeth. Fore tibia moderately curved, inner margin with two very low tubercles beyond basal curve, followed by a broader, black tooth, then several small tubercles more distally; a small tooth also present at base of oblique subapical comb of hairs. Trochanter of middle leg with a small ventral spine near apex. Middle femur slightly thickened centrally, ventral surface bearing a row of 7-9 almost straight black spines interspersed with ~20 much smaller spines. Hind femur slightly thickened on distal third, not dorsoventrally compressed; hind trochanter, femur and tibia unarmed.

Female legs with fore femur bearing a row of ~20 small black teeth along ventral margin, fore trochanter and tibia unarmed. Middle trochanter unarmed; middle femur with ~18 small black spines along ventral margin.

Male metasternum with an apically truncate posteromedial process arising from center of posterior margin, this process with lateral margins converging inward on basal halves, then parallel on distal halves, posterior portion of process bearing scattered moderate sized papillae (Fig. 19); styliform processes of abdominal segment VIII slender with curved apices.

DISTRIBUTION: Described from the Chesterfield Islands, in the Coral Sea (Andersen and Weir, 2000), and currently known only from that small archipelago (Fig. 24). The type locality is given in Andersen and Weir (2000) as "Bennett and Long Island"; in fact, both Bennett Island and Long Island are separate islands within the Chesterfield Reefs, and are several nautical miles apart. Long Island is the largest of Chesterfield Islands, and is approximately 1,500 m long but only 100 m wide; Bennett Island is slightly smaller and lies south of Long Island. Both islands are less than 15 m in elevation. It appears from the label data on the type series of H. armatus that the specimens were taken from a boat at night, lying offshore somewhere in the sector between these two islands along the northwestern side of the Chesterfield Islands complex.

DISCUSSION: Hermatobates armatus is similar to H. marchei, bur can be separated by the shape of the male metasternal process, the lateral margins of which are inwardly angling on the basal half and parallel on the distal half (Fig. 19), rather than being parallel throughout in as in H. marchei (Figs. 2C, 3C), and by the dentition of the male fore tibia, with a relatively low bi-tumescent tubercle immediately distal to the basal curve, followed by a larger blunt tooth. Andersen and Weir (2000) provided a discussion of additional characters separating H. armatus from other Hermatobates species, although their comparison to H. marchei actually refers to H. weddi. These authors also provided excellent illustrations of the male thoracic and abdominal venter, male styliform processes, male foreleg, and male middle and hind femora.

SPECIMENS EXAMINED: NEW CALEDONIA: North Prov.: Chesterfield Islands, Bennett and Long Island [area between Bennett Island and Long Island], 7-9 May 1979, 100 m outside reef, at light on sea surface [19[degrees]53'27"S, 158[degrees]19'28"E], N. Svennevig, 2 [male][male], 2 [female][female] (paratypes, JTPC).

Hermatobates schuhi, new species

Figures 2D, 3F, 4A, 5C, 6A, 18C, 23, 25

HOLOTYPE: JAPAN: Ryukyu Islands: Iriomote Is., south side, beach and rocks inside reef [24[degrees]16'17"N, 123[degrees]49'51"E], 22 March 1995, CL 2934, J. T. Polhemus, [male] (BPBM).

DESCRIPTION: Male: length 4.00-4.20 mm, maximum width 1.70-2.00 mm. Female: length 3.55-3.60 mm, maximum width 2.00-2.10 mm.

COLORATION: Ground colour black with a bluish tinge, dull, body with fine silvery pubescence, and longer setae laterally (lengths of these setae approximately 0.20 in males; 0.10 in females). Head dark bluish-black above, posterior margin narrowly brown; thorax bluish black centrally, silvery-grey laterally, abdomen with tergites bluish-black, connexiva brown. Pleural and ventral surfaces grey, dull, thoracic and abdominal sternites browned centrally, genital segments brown, shiny. Antennae blackish brown, basal two-thirds of segment I luteous, increasingly embrowned beyond. Legs medium brown, with dorsal surfaces and entire middle femur darker, fore coxae partly luteous; fore coxae and trochanters yellowish brown.

Apterous male: body fusiform, length more than 2.0x greater than greatest width across thorax (4.00 : 1.70). Head length 0.25x greatest head width across eyes (0.35 : 1.38); eyes rather small, width about 0.28x interocular width (0.25 : 0.90).

Antennal formula I : II : III : IV; 1.00 : 1.00 : 0.75 : 0.60; segment I almost 3x head length (1.00:0.35), slightly thicker than segments II-IV; combined length of all antennal segments 0.84x total length of insect.

Pronotum very short, median length (0.10) less than half of an eye width; posterior margin almost straight in middle, curved backwards laterally. Meso- and metanotum simple. Metastemum with relatively short, semi-trapezoidal posteromedial process arising at posterior margin of metasternum (Figs. 2D, 3F), this process bearing numerous small, stout, conical papillae (Figs. 3F, 4A).

Proportions of legs as follows: Femur, tibia, tarsal 1, tarsal 2, tarsal 3 of fore leg, 1.30 : 1.40 : 0.03 : 0.03 : 0.10; of middle leg, 2.10 : 1.30 : 0.10 : 0.80 : 0.50; of hind leg, 2.10 : 1.40 : 0.10 : 0.55 : 0.50. Trochanter of fore leg with single small black spine before distal end. Fore femur incrassate, with a stout pointed black tooth at base, angled distally, a broad distally bifid black tubercle before apex, between these an almost regular row of 14-16 small black teeth. Fore tibia gently curving basally and distally, central portion straight, basal curve with a low, triangular process on inner margin, ventral margin of tibia with a broad, bifid tubercle just beyond basal curve, followed by a depression, then a large stout tooth (Fig. 5C), then a row of small tubercles extending about two-thirds remaining distance to apex. Middle trochanter with distinct slender, black ventral spine distally. Middle femur slightly thickened in middle (0.27), ventral surface armed with 20-24 almost straight stout spines of relatively equal size, the most distal of these progressively shorter, those near base with tips hooked, interspersed by shorter teeth. Hind femur slightly thickened centrally (0.20), dorsoventrally compressed on distai hall; trochanter and femur unarmed. All leg segments thickly set with long, erect, silvery pilose setae.

Abdomen short, length from abdominal scent orifice to tip 1.30; lengths of abdominal terga 4-7: 0.10, 0.16, 0.20, 0.25; genital segments large, length 0.70; styliform processes of segment VIII basally broad, tips curved over at appoximately 90[degrees], with a conspicuous subapical concavity (Fig. 6A); male endosoma with basolateral lobes highly expanded, inner margins explanate (Fig. 18C).

Apterous female: body fusiform, length about 2.0x greater than greatest width across thorax (3.55: 1.80). Head length 0.28x head width across eyes (0.40 : 1.40); eyes rather small, slightly less than 0.22x interocular width (0.20 : 0.90).

Antennae relatively shorter than in male, 0.6x total length of insect, antennal formula I : II : III : IV; 0.40 : 0.70 : 0.50 : 0.55; segment IV about 1.4x head length (0.55:0.40), subequal in thickness to segment I.

Pronotum very short, median length (0.10) about half of an eye width; metasternum not modified.

Legs relatively shorter than in male. Proportions of legs as follows: Femur, tibia, tarsal 1, tarsal 2, tarsal 3 of fore leg, 0.95 : 0.95 : 0.03 : 0.07: 0.28; of middle leg, 1.30 : 0.90 : 0.09 : 0.60: 0.52; of hind leg, 1.60 : 1.00 : 0.07 : 0.53 : 0.50. Fore femur moderately thickened, greatest width 0.20; trochanter and femur unarmed except for a row of about 16 small black teeth ventrally on femur; fore tibia nearly straight, unarmed. Middle femur slightly thickened in middle (0.19), ventrally with a row of ~7 small black spines on basal half; trochanter unarmed. Hind femur slightly thickened in middle (0.14), unarmed. All leg segments bearing erect, slender silvery setae, these setae shorter and less dense than in male.

Abdomen length from abdominal scent orifice to tip 2.10; lengths of abdominal terga I-VII: 0.90; 0.20; 0.15; 0.15; 0.20; 0.25; 0.20. Abdominal sterna II-VII forming a large quadrate plate, length 0.50, posterior margin slightly indented centrally. Genital segments, except for cone-shaped proctiger, hidden beneath sternal plate.

ETYMOLOGY: The name "schuhi" honors Randall "Toby" Schuh for his many contributions to the study of Heteroptera, and his long friendship with the authors.

DISTRIBUTION: Apparently endemic to the Tokara and Ryukyu archipelagoes, lying south of the main islands of Japan (Figs. 23, 25). We have records from Takarajima Island in the Tokara group, recent collections from Okinawa, Iriomote and Yonaguni islands in the Ryukyus, and additional literature records from AmamiOshima, the Kerama Isles (Tokashiki), Kumejima, and Ishigaki, also in the Ryukyu chain (Hayashi, 1997). Given that Yonaguni Island is within sight of Taiwan, it seems logical that this species might occur along the coasts of the latter island as well. Specimens taken from coral reefs off the coast of Vietnam may also represent this species (Tran, in litt.) but we have not yet examined these specimens to verify their identity.

DISCUSSION: The presence of Hermatobates in the waters surrounding the island groups immediately south of Japan has been noted for many decades (Esaki 1934, 1935; Miyamoto 1961; Sato 1994; Hayashi 1997). These preceding authors all provided records of the family Hermatobatidae or the genus Hermatobates, but the precise identity of the species in question remained confusing. Esaki (1947) eventually concluded that his specimens taken off Ishigaki Island in the Ryukyu Islands represented H. haddoni, Takara (1957) subsequently recorded H. marchei from Ishigaki, and Miyamoto (1961), followed by Kawai (1985) while Azuma and Kinjo (1987), recorded H. weddi from various localities in the Ryukyus and the adjacent Tokara Islands. All of these previous records are in fact referable to our new species H. schuhi.

Hermatobates schuhi is a northern segregate of the H. weddi group and is close to H. weddi, but may be separated from that species by the dentition of the fore tibia, which has the prominent sub-basal teeth distal to the basal curve closer together, with the rounded notch between them equal in width to the width of largest, most distal of these teeth (compare Fig. 5C, D). The posteroventral processes of the male metasterna, although broadly trapezoidal in shape in both species, are also different, with that of H. schuhi being slightly larger and more emarginate (compare Figs. 2F, 3D to 2D, 3F), and bearing smaller and more numerous papillae, the latter only visible in detail using SEM imagery (compare Figs. 3D, 4B to 3F, 4A). The apices of the male styliform processes also show slight differences between the two taxa, in particular the presence of a subapical concavity in H. schuhi (compare Fig. 6A, B). The everted male endosoma shows distinct differences, particularly in the form of the basolateral lobes, which are far larger and plate-like in H. schuhi (compare Fig. 18C, D). Finally, in overall size H. schuhi is the largest species in the genus, with the body length of males equal to or exceeding 4.0 mm.

The geographic range of H. schuhi in the Ryukyus is also highly geographically disjunct from that of H. weddi in Australia and the adjacent southwest Pacific, with the ranges of H. marchei and H. singaporensis intervening between the two.

ECOLOGICAL NOTES: On Yonaguni, at Katabara-Lama near Higawa, the first author took several specimens of H. schuhi in the lee of large, porous boulders near low tide, a typical Hermatobates habitat, but the majority of specimens were by contrast taken from the glassy smooth surface of the bay beyond, in shallow water along the farthest inland shore. Here the slanting light was fortuitous and by using the glare on the water, individuals could occasionally be seen cruising in a sinuous course with incredible speed. In order to capture these, it was crucial to move quickly so as to intercept their course, then when the insects were close at hand and visual contact was lost, to strike downward with the net along the estimated trajectory of the speeding insect, leading ir as a trap shooter might lead a target. This "glare shot" method of capture is often very effective at revealing Hermatobates in areas where they might not otherwise be detected.

PARATYPES: same data as holotype, 3 [male][male], 1 [female] (JTPC, BPBM); Okinawa Is., Bise-zaki [26[degrees]42'32"N, 127[degrees]52'45"E], 27 March 1995, CL 2949, J. T. Polhemus, 1 [male] (JTPC); Yonaguni Is., Katabara-Lama, nr. Higawa [24[degrees]26' '18"N, 122[degrees]58'25"E], 21 March 1995, CL 2929, J. T. Polhemus, 2 [male][male], 3 [female][female] (JTPC); Tokara Islands, Takarajima [29[degrees]08'38"N, 129[degrees]13'03"E], 26 May-1 June 1953, Osaka Municipal Museum Tokara Islands Expedition, S. Miyamoto, 2 [male] (JTPC).

ZOOGEOGRAPHIC NOTES

The genus Hermatobates seems to be definitely restricted to [+ or -] 30[degrees] latitude (see Fig. 2 in Andersen, 1991), and is most prevalent between the Tropic of Capricorn and the Tropic of Cancer ([+ or -] 23[degrees]), with a few notable exceptions. For instance, the entire known range of H. schuhi lies above the Tropic of Cancer (ca. 23.5[degrees]N) in the northwestern Pacific Ocean, with the northernmost records from the Tokara Islands (28[degrees]N). In the Indian Ocean, H. djiboutiensis has been taken from the coast of Israel at 28[degrees]N, and in the Atlantic, H. bredini is known from the Bahamas at 25[degrees]N.

Based on the information presented in this paper and references cited herein, Hermatobates species are clearly reef-associated to varying degrees, and although individuais of some species may be taken at considerable distances seaward of the reef crest, none can be considered truly pelagic. Despite this, the fact that the genus has colonized many isolated island groups, including Hawaii, the Marquesas, and the Line Islands, indieates that Hermatobates has a certain capacity for intermittent long distante ocean dispersal. Such dispersai would seem to be infrequent, given that the known species occupy generally allopatric ranges and are rarely taken in multi-species assemblages, although exceptions to this general rule occur. For instance, H. marchei, H. weddi, and H. kula have all been collected in the same night light sample at Mebulibuli Point on Fergusson Island in the D'Entrecasteaux group, off eastern New Guinea. This anomalous site would appear to lie in a narrow contact zone at the margins of several larger core ranges occupied by each of the species involved.

Hermatobates weddi is the most widespread species in the genus, distributed from northern Australia and southern New Guinea eastward to at least Fiji, and perhaps Tonga and Samoa; its range thus involves the Arafura Sea, Coral Sea and North Fiji Basin (Figs. 20, 23, 25). Hermatobates djiboutiensis also has a similarly wide range in the western Indian Ocean, occurring from the Red Sea and Gulf of Aden to the coasts of Tanzania, Madagascar and Mauritius, and eastward to the Maldive Archipelago (Fig. 21). It is interesting that no other species of Hermatobates is known to have colonized the Indian Ocean westward of the coast of Thailand, although this biogeographic disjunction between the Western Pacific and the Western Indian Ocean is also seen in other demonstrably monophyletic marine groups, including nearshore Halobates water striders (Andersen and Cheng, 2004) and multiple subgenera of cowries (Meyer, 2003), and therefore represents a typical and repetitive pattern of marine vicariance.

By contrast to the above, all other Hermatobates species have more restricted ranges, as far as they are known. The only species occurring in the Atlantic Ocean, H. bredini, is confined to the Caribbean Sea, with records from Dominica in the east to Belize in the west and the Bahamas in the north, and so far including Cuba and Puerto Rico, but with no records yet from the Yucatan Peninsula of Mexico, the Cayman Islands, Jamaica or Hispaniola, although it is likely present in all these areas (Fig. 22). Its putative sister species, H. hawaiiensis, occupies an extremely isolated position in the Hawaiian Islands. The two other eastern Pacific species, H. palmyra and H. tiarae, are also putative sister taxa with ranges lying north and south of the equator respectively (Fig. 20), suggesting possible latitudinal stratification by equatorial currents. Another restricted distribution is that of the small and morphologically distinctive species H. haddoni, known only from northern and western Australia (Fig. 24).

Whereas the distributions of most Hermatobates species, with the exception of H. haddoni and to some extent H. djiboutensis, occupy largely archipelagic distributions, the ranges of species in the H. weddi group are by contrast largely associated with Asian and Australian continental platform coastlines that were significantly altered by sea level fluctuations linked to Quaternary glaciation (Voris, 2000). In particular, unless one assumes that the current distribution of H. weddi is the result of very recent, wide-ranging dispersal from a single source area, then the populations of this species currently occurring in western Australia and the Lesser Sunda Islands were isolated from those occurring in the Coral Sea and archipelagoes to the east (the Solomons, Fiji, Tonga, Samoa) as recently as 20,000 years ago during the last Pleistocene glacial maximum (Fig. 25). At this time, sea levels were over 100 m lower than at present, and the Torres Strait and Sahul Shelf were emergent land. This same low sea level stand would also have isolated any populations of the H. weddi group occurring north of the Sunda Shelf from those occurring to the south, which would explain the present day disjunction between H. schuhi in the Ryukyu Islands and the closely allied H. weddi in the southwest Pacific.

This recent Pleistocene event was in turn only one in a series of sea level low and high stands that have alternated in the Southeast Asian region throughout the Quaternary. Given that the Sunda and Sahul shelves would be almost entirely exposed at sea levels 50 m below present that prevailed between 70-20 K years BP, and that extensive areas would be emergent at even 25 m below present sea level, such potential vicariance has occurred at least 10 times in the past 800 K years. Maximum low stands occurred at 630 K, 430 K, 140 K and 20 K years BP (Bintanja et al., 2005), all of which would have represented significant vicariance events for Southeast Asian marine biota. Within this period of multiple low stands, sea level was at least 25 m lower than present on a continuous basis from 100 K-10 K years BP, a period of 90 K years. By contrast, marine transgression onto the Sunda Platform comparable to present sea levels occurred at 400 K (Holsteinian), 325 K, and 125 K (Eemian) years BP; the two highest of these episodes, the Holsteinian and Eemian, saw sea levels perhaps 5 m higher than present. All other high stands were at least 25 meters lower than these peaks (Bintanja et al., 2005). The two highest stands may have been sufficient to isolate the Malay Peninsula as an island by flooding the Tasek Berah lowlands, although such isolation would only have persisted for perhaps 10 K years. Each of these high stands would have represented a potential dispersai opportunity during which previously separated populations of marine organisms could be reunified. Such alternating episodes of potential marine vicariance and reconsolidation are very much a Holocene phenomenon, since sea level in the preceding Pliocene was 25 m higher than present, which would have promoted broad distribution of shallow water tropical marine species and precluded vicariance of such biotas, because there would have been no possibility for interposition of exposed continental shelves (Dowsett el al., 1994).

We hypothesize that one result of these multiple Quaternary episodes of minimum and maximum sea level has been the repeated isolation and re-integration of populations in the H. weddi group, leading to the current confusing situation in terras of clearly delimiting species concepts. As alluded to above, we consider it likely that one population in this group, which we have now described as H. schuhi, was isolated north of the Sunda Shelf, where it differentiated in the Ryukyu and Tokara Islands. In the west, the Gulf of Thailand formed a sheltered, island-enclosed tropical sea basin during the Holocene low stands, separated by thousands of kilometers of emergent Sundaland from the similarly sheltered Banda and Timor seas bordering Australia in the west (Voris, 2000). We consider it possible that local differentiation may also have occurred in this western sector, giving rise to the populations we currently treat as H. singaporensis. The Hermatobates populations oceurring along the coasts of Australia were similarly divided by the emergent Sahul Shelf, with one set isolated in western Australia, and the other in the Coral Sea basin (Fig. 25), but neither we nor previous authors (i.e., Andersen and Weir, 2000) have yet found a basis for discriminating separate taxa as a result of this vicariance event. All of the above populations in the H. weddi group have now been re-integrated to some degree during the current Holocene high stand, and the present distributions of species in this group are probably extremely recent in geologieal terms; in particular, all known localities for H. singaporensis must have been re-colonized within the past 10 K years because they lie on the formerly emergent Sunda platform (Fig. 25).

Of additional interest is a clear segregation of generally allopatric species along the ares bordering the western margin of the Pacific Plate; from north to south these are H. schuhi in the Ryukyu Arehipelago (Fig. 23); H. marchei inthe Philippines, Carolines and northern New Guinea (Figs 23, 24); H. kula in the Louisiade and D'Entrecasteaux island groups offshore of far eastern New Guinea (Fig. 24), and H. armatus in the New Caledonia sector (Fig. 24). These Western Pacific distribution patterns suggest some degree of older, tectonically-mediated speciation in Hermatobates along the western margin of the Pacific Plate that operated independently of the Quaternary vicariance events hypothesized above.

Despite our improved understanding of Hermatobates distributions in the insular Pacific and Indian oceans, many areas remain to be adequately surveyed. Prime among these are the coasts of Sumatra, Java, Borneo, Sulawesi, and the Moluccas in the Malay Archipelago, which represent a potential contact zone for H. singaporensis, H. marchei and H. weddi, and are likely to be critical to clarifying species concepts in the H. weddi group and assessing putative Quaternary vicariance. It is possible that additional new species may occur here as parapatric segregates, as seems to be the case with H. kula and H. armatus in the complex tectonic and faunal contact zone of eastern Melanesia. In the Indian Ocean, we have seen no collections of Hermatobates from the Bay of Bengal or the Gulf of Thailand, although the genus clearly occurs in the latter area based on records of immatures and exuviae at Phuket Island (Andersen, 1989). As such, we do not have a good understanding of the eastern limits of 11. djiboutensis, which probably does not cross the central Indian Ocean barrier, or of the northwestern range limit of H. singaporensis, the latter once again bearing on evaluation of potential Holocene vicariance in the H. weddi group. Similarly, we have a poor understanding of Hermatobates distributions along the coasts of Vietnam and China, around Taiwan, and in the Marianas, or of the furthest northward distribution of Hermatobates along the island arc systems running north from the Mariana Islands to Japan in the Ogasawara Island group. AH of these areas provide excellent prospects for further collecting and research, and specimens from these areas are criticai to resolving current taxonomic uncertainties in the genus.

ACKNOWLEDGMENTS

It is impossible to adequately thank all of the persons who contributed to our research on Hermatobates over the past three decades, they include other scientists, in-country counterparts at all levels, from science ministries to local villagers, as well as those who skillfully skippered the boats large and small from which we collected specimens off many islands in the tropical oceans of the world. The list below provides a partial acknowledgement of all this generous assistance; if there are omissions of those who made significant contributions, we apologize in advance for the oversight.

The staff of Conservation International's Melanesia Program, specifically Bruce Beehler and Todd Stevenson in Washington, DC, Gai Kula in Port Moresby, and David Mitchell, Bena Sena, Neal Lima in Alotau, and Dr. Gerald Allen of Perth, Australia, ali of whom provided invaluable support for surveys in Papua New Guinea and the Solomon Islands. Dr. Ron Englund, Dr. Allen Allison, and Dr. Neal Evenhuis, all of the Bishop Museum, Honolulu, Hawaii, who provided logistical support in both the lab and the field. Dr. Brian Kensley, Dept. of Invertebrate Zoology, Dr. Thomas J. Henry, USDA ARS, and the late Jon Herring, all of the Smithsonian Institution, Washington, DC (USNM). Jean-Yves Meyer of the Delegation a la Recherche, Papeete, Tahiti, who permitted research in the Society and Marquesas Islands. Dr. Peter K. L. Ng, National University of Singapore Kent Ridge and Lanna Cheng, Scripps Institute of Oceanography, La Jolla, who facilitated surveys around Singapore. Drs. Rosichon Ubaidillah, Dr. Soetikno Wiroatmodjo, Dr. Mohammed Amir, and other personnel from the Indonesian Institute of Sciences (LIPI), Bogor; Mrs. Moertini, Mr. Sanchoyo, the other staff members of the LIPI office in Jakarta; Mr. Benny Lesomar, Natrabu Corp., Biak; Mr. Lewis Zeigler, Mr. Jan Beiers, Mr. T. Loebis, Mr. Franz Petrusz and many others from the Petromer Trend Oil Company, Jakarta and Sorong; Mr. Neal Blackburn, Mobil Oil Indonesia Inc., Jakarta; Mr. James Carpenter, Mobil Oil Indonesia Inc., Fakfak; Mr. Petrus Manggai, Fakfak; Mr. Jacob Bakabessy and Mr. Bernardus Sambery, PHPA, Manokwari; all of whom facilitated surveys in Indonesia. Dr. M. Sato of Nagoya Womens University, Nagoya for his assistance with field work in the Ryukyu Islands. Burt Barrion, International Rice Research Institute, Los Banos, who organized surveys in the Philippines. Dr. Vincent Razafimahatratra, Mme. Laia Rakotovao, Dr. Voara Randrianasolo and Dr. Evariste Randrianasolo, all of Tananarive, who made possible our survey work in Madagascar, Dr. Damir Kovac, for gift of specimens he collected in the Maldives; and Dr. William Foster of Cambridge University, for specimen loan and kind donation to JTPC of specimens collected in the Southwest Pacific by Lanna Cheng. Dr. Cheng also separately donated additional material collected in various localities worldwide.

Special thanks are also due to the crew of the sailing yacht El Gringo, which served as a floating base of operations during the 1989 surveys by the second author in the Western Indian Ocean, and to Wayne Thompson, his wife Lee, and the various Melanesian crew members of the research vessel

Marlin 1, which served as a floating laboratory during marine surveys in Melanesia from 2002 to 2005.

This research was sponsored by a series of grants from the National Geographic Society, Washington, D. C. (2698-83, 3053-85, 3398-86, 4537-91); by grants BSR-9020442 and DEB 952-8025 from the National Science Foundation, Washington, D. C.; by a Melanesian Freshwater Ecosystems grant from Conservation International; anal by the Smithsonian Institution's Drake Fund. This paper is a research contribution from Colorado State University, where JTP is adjunct faculty, and with which the Colorado Entomological Museum is affiliated. We thank all these organizations for their continued support of research into the systematics and zoogeography of aquatic Heteroptera.

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JOHN T. POLHEMUS (1) AND DAN A. POLHEMUS (2)

(1) Colorado Entomological Institute, 3115 South York Street, Englewood, CO 80113, USA

(2) Dept. of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, HI 96817; bugman@bishopmuseum.org

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