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Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species.

INTRODUCTION

Bollmannia Jordan, 1890, is one of five genera that make up the Microgobius group of the tribe Gobiosomatini (Birdsong et al. 1988; Van Tassell & Baldwin 2004). The Gobiosomatini is a monophyletic group within the subfamily Gobiinae (sensu Pezold 1993; Thacker 2003; Thacker & Hardman 2005) and the tribe is often referred to as the American seven-spined gobies. Bollmannia currently comprises 14 nominal species, five of which were described from the Atlantic Ocean: Bollmannia eigenmanni (Garman, 1896), B. boqueronensis Evermann & Marsh, 1899, B. jeannae Fowler, 1941, B. communis Ginsburg, 1942, and B. litura Ginsburg, 1935. While attempting to identify specimens of Bollmannia collected from Panama in 2004, several problems were noted in the descriptions of the Atlantic species. The characters described for each of the species are not uniform across each of the descriptions. Several of the characters described as diagnostic are of little value as these characters are commonly destroyed when Bollmannia are trawled or dredged from considerable depths. In most cases the descriptions are based on a small number of specimens and do not adequately summarize the intraspecific variation seen among populations. Furthermore, there is a discrepancy between the description of B. litura and the corresponding holotype. As a result of these issues, the validity of several species is questioned. Bollmannia jeannae is not listed by Nelson et al. (2004), McEachern & Fechhelm (2005) or Murdy & Hoese (2002). Both B. jeannae and B. eigenmanni are omitted by Smith (1997). Eschmeyer (2011) lists all five nominal species as valid. This paper attempts to clarify these discrepancies and resolve the status of the Atlantic members of the genus.

No formal review of western Atlantic Bollmannia has been published to this point. In an unpublished master's thesis, Bedenbough (1988) reviewed the Pacific members of the genus. Ginsburg addressed the genus in two incomplete, unpublished manuscripts, the drafts and notes of which have been found at the Smithsonian Institution's National Museum of Natural History. Work on these manuscripts likely began between 1939 and 1942, as some of his notes included the eastern Pacific Bollmannia species described in Ginsburg (1939) but not B. communis, which he described in 1942. Subsequent drafts by Ginsburg contained B. communis and additional specimens of each of the species discussed in prior drafts. In his unpublished works Ginsburg described and illustrated sensory papillae and head pore patterns for several species of Bollmannia. We describe these characters for each of the valid species in the western Atlantic.

In addition to the five nominal species of western Atlantic Bollmannia, we also address a previously undescribed deepwater goby from the Caribbean. This species was originally documented as the "filamentous goby" by Colin (1974) and subsequently referenced by Dennis et al. (2004), where it was referred to as the "sabre goby". The molecular phylogenies of Ruber et al. (2003) and Ruber & Van Tassell (unpublished, presented in 2006) suggested that this species belongs within the monophyletic Microgobius group of the tribe Gobiosomatini, specifically as the sister group to a monophyletic clade containing four species of Bollmannia. Despite the apparent close relationship between this species and Bollmannia, there are several prominent differences between the two in terms of morphology and habitat. Therefore, we describe this species and place it in a new genus, Antilligobius, within the Microgobius group of the Gobiosomatini. The affinities between Antilligobius n. gen. and the allied genera Bollmannia, Akko, Parrella, Palatogobius and Microgobius are discussed. Identification keys are provided for the Atlantic species of the Microgobius group of the Gobiosomatini.

METHODS

Methods for counts and measurements follow Bohlke & Robins (1968) in most regards. The last two elements in the second dorsal and anal fin stem from a single pterygiophore and are counted as a single ray. The upper and lower segmented caudal rays are frequently short with only one or two striae but are stouter than adjacent procurrent rays and are included in the counts of segmented caudal rays. Counts of branched caudal rays include rays with only slight terminal branching. Lateral scale counts are the number of diagonal rows beginning with the scale above and behind the posterodorsal corner of the operculum and ending at the posterior edge of the hypural plate. Transverse scale counts are the number of horizontal scale rows counted from the origin of the second dorsal fin diagonally to the origin of the anal fin. Caudal peduncle scales are counted beginning at the first full sized scale on top of the caudal peduncle immediately in front of the caudal fin, and following the scale rows down and forward to the ventral edge of the peduncle, then around and back to the original scale. Gill raker counts are total counts for the outer edge of the first gill arch.

The following measurements are straight-line point to point measurements taken with digital calipers: standard length, snout tip to posterior edge of hypural plate; head length, snout tip to posterior edge of operculum, not including fleshy membrane; predorsal length, distance from the insertion point of the first dorsal spine to the tip of the snout; snout length,snout tip to the anterior rim of eye; upper jaw length, anterior tip of upper jaw to posterior margin of maxillary; eye diameter, horizontal diameter of orbit; pupil diameter, horizontal diameter of pupil; postorbital length, posterior rim of eye to posterior edge of operculum, not including fleshy membrane; body depth at first dorsal fin origin,vertical distance from origin of spinous dorsal fin to ventral profile, not including pelvic fin; least caudal peduncle depth, vertical distance at shallowest point between base of last anal ray to posterior edge of hypural plate; and caudal fin length, distance from origin of ray on hypural plate to tip of longest ray.

Dorsal pterygiophore formula is that of Birdsong et al. (1988), patterns of sensory papillae are described according to Sanzo (1911) and cephalic canal pore terminology follows Akihito et al. (1988). In some cases papillae and pore patterns were more easily observed by staining with cyanine blue (Saruwatari et al. 2006). Osteological observations were made from radiographs and specimens cleared with trypsin and counter stained for cartilage and bone (Dingerkus & Uhler 1977). Institutional acronyms follow Sabaj Perez (2010).

In many cases scale counts of Bollmannia were determined by counting scale pockets or remnants thereof, as scales in this genus are highly deciduous and specimens are typically in poor condition as a result of collection via trawl or dredge. In many other specimens, including some holotypes, scale pockets were missing entirely. For this reason, the scale count ranges presented here are trends from a composite of specimens and should be regarded as approximations. Similarly, the fins are heavily damaged on most specimens of Bollmannia. The relative lengths of rays and spines described here should also be regarded as approximations. Papillae and pore patterns shown here are composite drawings from several specimens. In nearly every specimen of Bollmannia, portions of the lateralis canals are destroyed and in many cases entire rows of papillae are missing as a result of damage or poor preservation. In particular, the predorsal region, the posterior half of the preoperculum and the operculum are typically more heavily damaged than other regions. Thus the absence of extensive papillae or cephalic canals in these regions (specifically the canal between K' and L') on our illustrations may be subject to correction if specimens are examined in which papillae and canals are more perfectly preserved. Despite these issues there is some taxonomic value in the aforementioned characters. However, this value, and consequently the ability to confidently identify specimens of Bollmannia, is largely determined by the overall physical condition of the specimen.

SYSTEMATICS

Antilligobius, n. gen.

Van Tassell & Tornabene

Diagnosis: Antilligobius can be distinguished from other Gobiosomatini by the following combination of characters, not listed in order of taxonomic importance: first dorsal fin very elongate in both sexes, extending posteriorly at least to base of last anal fin ray when depressed, sometimes reaching middle of caudal fin; caudal fin elongate but with upper rays 3,4,5 shorter then corresponding lower rays, forming notch in upper margin of fin; pelvic fins united to form disk with well-developed frenum, frenum with smooth posterior margin, not supported with collagenous thickenings; trunk scales ctenoid, lateral scale rows 21-28; predorsal region, cheek, pectoral fin base and breast with cycloid scales; predorsal scales 8-10; first caudal vertebra with expanded haemal arch and tiny haemal spine; haemal arch with elongate parapophyses, "riblike" in appearance in lateral view; abdominal cavity elongated and extending through first haemal arch.

Description: Body slightly elongate and laterally compressed; first dorsal spines VII; pterygiophore formula of first dorsal fin 3(221110); second dorsal fin I,11-12; first and second dorsal fins separate from each other and from caudal fin; pectoral rays 18-21; anal fin I,11-13, first two anal pterygiophores inserted anterior to first haemal spine, remaining pterygiophores in approximately 1:1 ratio with caudal vertebrae; total vertebrae 27, 11 precaudal and 16 caudal; pelvic fins with one spine and five rays; vomerine teeth absent; mouth large, 10.0-11.2% SL and inclined 30[degrees] from horizontal; eye large, diameter 10.2-12.0% SL; gill opening extending from upper edge of pectoral fin to ventral body midline; gill rakers on lower limb of first arch thin and elongate, rakers on upper arm short, stubby; urogenital papilla thin and conical in males, thick and bulbous in females; cephalic lateralis canal and pore system on head well-developed; sensory papillae in transverse pattern; ventral postcleithrum present; no obvious sexual dimorphism, except for urogenital papilla.

Comparison: Antilligobius has a number of characters not present in other American seven-spined genera that warrant its placement in a new genus. These include: a very narrow, splinter-like metapterygoid; short slender gill rakers on the lower limb of the first arch; a short papillae row b, never extending to posterior of the pupil; a modified first haemal arch; the caudal fin elongate but with the upper rays 3,4,5 shorter than the corresponding rays on the ventral half of the fin, forming a notch in the upper margin of the fin.

Antilligobius is most similar morphologically to Bollmannia and Parrella, sharing with those genera the presence of scales on the breast, belly, pectoral fin base and predominately ctenoid scales on the trunk. Bollmannia and Antilligobius also both possess scales on the cheek and a spatulate basihyal. The bright yellow stripe and blue colors of Antilligobius easily distinguish it from Bollmannia and Parrella which are drab brown with darker brown markings.

Etymology: The generic epithet Antilligobius is formed from the Dutch "Antillen," which refers to the region now known as the Antilles or Caribbean Sea, and the Latin gobius meaning small fish or gudgeon. To be treated as masculine. The name is given in reference to its distribution within the ancient "Sea of the Antilles".

Antilligobius nikkiae, n. sp.

Van Tassell & Colin

Sabre goby (Figs 1-2, 4-11)

Holotype: AMNH 253403, 30.2 mm SL, female, Curacao Sea Aquarium slope, Curacao, 12.08715[degrees]N, 68.896576[degrees]W, quinaldine collection, collected by Adriaan "Dutch" Schrier, July 2004, depth 90 m, loose rock slope (Fig. 3).

Paratypes: AMNH 253404, 2 males, 25.8-26.7 mm SL, 1 female, 26.7 mm SL, collected with holotype; ANSP 147645, 29.7 mm SL, female, cleared and stained specimen, shelf edge of reef at La Parguera, Puerto Rico, 28 August 1979, Pat Colin; ANSP 150387, 1 male 21.7 mm SL, 1 female 25.7 mm SL, shelf edge of reef at La Paruera, 17.949005[degrees]N, 67.045345[degrees]W, 87-90 m, 45[degrees] slope, collected among small indentations in rocky structure of slope face, Puerto Rico, 22 November 1978, Pat Colin; AMNH 233129, 1 male, 23.19 mm SL, 90 m, JVT-01-012, Genbank AF491066, Curacao Sea Aquarium slope, 12.08715[degrees]N, 68.896576[degrees]W, Curacao, January 2001, Adriaan "Dutch" Schrier; USNM 402717, 1 female, 31.6 mm SL, 168-198 m, west coast of Isla de la Juventud, 21.78405[degrees]N, 83.23625[degrees]W, Cuba, 27 December 1997, R. Grant Gilmore.

Description: Morphometric data given in Table I. Counts of the holotype (Fig. 1) indicated by an asterisk, followed by number of specimens with each count in parentheses.

Table I. Morphometries for type specimens of Antilligobius nikkiae
(excluding cleared and stained specimen). Measurments other than
SL are in % SL.

                   AMNH        AMNH        AMNH        AMNH
                  253403      233129      253404      253404
                (holotype)  (paratype)  (paratype)  (paratype)

Sex                 female      female        male        male

SL (mm)               30.2        23.0        26.7        25.8

Eye diameter          10.6        10.6        10.7        10.2

Upper jaw             10.6        10.0        11.2        10.8
length

Head length           24.5        25.7        27.3        25.9

Postorbital           11.9        10.9        12.0        11.2

length

Depth at DI           19.5        18.3        21.0        21.5
origin

Least caudal          10.7        10.2        11.2        11.2
peduncle depth

Snout                  6.3         5.7         6.0         6.2
length

                   AMNH        ANSP        ANSP        USNM
                  253404      150387      150387      402717
                (paratype)  (paratype)  (paratype)  (paratype)

Sex                 female        male      female      female

SL (mm)               26.7        21.7        25.7        31.6

Eye diameter          12.0        10.6        10.5        10.4

Upper jaw             11.2        10.1        10.5        10.7
length

Head length           33.3        30.0        26.8        30.3

Postorbital           14.6        13.4        13.6        14.8
length

Depth at DI           18.4        17.1        17.9        18.3
origin

Least caudal          10.5        10.6        10.1        11.7
peduncle depth

Snout                  6.7         5.5         4.7         5.3
length


Median and paired fins: first dorsal VII*(9); first four dorsal spines very elongate, extending to base of last anal ray when depressed, extending to middle of caudal fin in some specimens; second dorsal I,11(4), I,12*(5); anal fin I,11(1), I,12*(1), I,13(7); pectoral rays 18(2), 20*(6), 21(1); pectoral fin elongate, reaching vertical through anterior end of anus; caudal fin elongate but with upper rays 3,4,5 shorter than corresponding lower rays, forming notch in upper margin of fin; segmented caudal rays 16(3), 17* (6), branched rays 14*(6), 15(3); pelvic fin I,5*(9); pelvic fins united to form disk, disk extending 2/3 to 3/4 distance from insertion of spine to anterior of anus; anterior margin of frenum extending halfway along pelvic spine, posterior margin of frenum smooth, without fimbriae and lacking collagenous thickenings.

Scales: body completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with cycloid scales; lateral scale rows 21(1), 25(4), 26*(2), 28(1); transverse scale rows 6(2), 7*(4); predorsal scales 7(1), 8(4), 9(1), 10*(2), scales extending anteriorly to behind eye; caudal peduncle scales 9(1), 10(3), 11*(2), 12(1); scales on cheek 3(4), 4*(2), 5(2); operculum scales 0*(4), 3(4); no modified basicaudal scales present.

Head: head length 24.5-33.3% SL; mouth angled upwards 30[degrees] from horizontal, reaching posteriorly to vertical through pupil; upper jaw length 10-11.2% SL; teeth in upper jaw in 2-3 rows anteriorly, becoming two rows about midway along premaxilla, ending in single row at posterior end of premaxilla; all teeth conical with slightly recurved tips; teeth in outer row longest; teeth in lower jaw in two rows anteriorly, becoming single row about half way along dentary; teeth conical with slightly recurved tips; single recurved canine located at mid-dentary; tongue slightly emarginate; eye large, 10.2-12.0% SL; interorbital width approximately equal to pupil diameter; snout short, 4.7-6.7% SL; gill opening extending from upper edge of pectoral fin to ventral midline; gill rakers on lower arm of first arch 12-13, upper arm 5; gill rakers on lower arm of first arch thin and elongate, rakers on subsequent arches shorter and thicker; epibranchials present; anterior nostril a short erect tube; posterior nostril with raised rim along anterior edge; no barbels present.

Genitalia: papilla elongate and conical in males, with distinct line of melanophores along dorsal edge present in one specimen, all others with no pigment; papilla short, bulbous, and without melanophores in females.

Sensory papillae and head pores (Fig. 4): oculoscapular pores B', C(s), D(s), E, F, G, H' present; posterior canal (typically between pores K'and L') absent; preopercular pores M', N, O' present; single interorbital canal; papillae in transverse pattern, with 5 short vertical rows below eye, none extending below level of row d; row d long, beginning below anterior margin of eye, ending before transverse row 4; row b short, beginning below eye and posterior to 5th transverse row, ending well short of posterior margin of preopercle; row n continuous across dorsal midline; rows g and m reduced, each represented by 2-3 papillae on each side of dorsal midline.

Pigment: body translucent pearlescent in life, tan in preservation; body with bright yellow stripe (white after preservation) along lateral midline beginning at eye and extending onto caudal fin; width of stripe about 3/4 of eye diameter at widest point, becoming thinner on caudal fin, ending at or near tip of caudal fin; thin iridescent blue stripe along ventral surface of yellow stripe anteriorly, absent in preservation; dorsal midline of body with narrow iridescent blue stripe, absent in preservation; abdomen and gular regions more or less pale; lateral edge of upper and lower jaw reddish-orange; iris of eye iridescent blue along dorsal and ventral margins, with iridescent patches of yellow at the anterior and posterior margins, iris uniformly dark in preservative; first dorsal fin translucent and tinged with yellow on interspinal membranes, uniformly clear in preservation; membranes between spines 4-6 of first dorsal fin with diffuse dark streak, streak more prominent in preservation; second dorsal fin yellow at base, translucent distally, uniformly clear with dusky distal margin in preservation; anal fin translucent with scattered melanophores along dorsal and ventral edges and with horizontal yellow stripe along midline of fin in life, stripe pale in preservation; pelvic fin translucent with yellow dorsal margin in life, pale in preservation; caudal fin translucent with yellow stripes in life, dusky in preservation; two caudal fin stripes continuations of midlateral yellow body stripe and yellow stripe along base of anal fin; two additional yellow stripes on caudal fin, one thin one along dorsal fin margin and another along ventral margin; stripes pale in preservation; upper half of pectoral fin base yellow (white in preservation), lower half unpigmented; pectoral rays translucent.

Osteology: based on ANSP 147645, female, cleared and stained specimen.

Suspensorium and jaws (Fig. 5): anterior arm of suspensorium consisting of palatine, relatively broad dorsally, tapering to point about midway along ectopterygoid; ectopterygoid articulating with quadrate along its anterodorsal edge; metapterygoid very thin, "needle-like", not articulating with quadrate or its small cartilaginous posterior edge; no symplectic process on preopercle; ascending process of premaxilla extending dorsally to anterodorsal edge of rostral cartilage; maxilla narrow, lacking process along shaft, with ventral tip curved anteriorly; dentary reduced, lacking large pocket typically receiving anguloarticular; anguloarticular articulating with dentary along ventral groove in dentary and in small anterior pocket near symphysis of dentary.

Cranial (Fig. 6): median ethmoid forming septum between eyes, anterior and posterior edges ossified; anterior of frontals overlapping median ethmoid; sagittal crest present along dorsal midline where left and right frontal bones fuse; sagittal crest confluent with supraoccipital crest; anterolateral edge of frontals possessing well-developed transverse process; this process forming open tube supporting cephalic sensory canal; sphenotic short, lacking well-developed anterior process.

Pectoral girdle (Fig. 7): left side of specimen with four partly ossified radials and cartilaginous scapula; radials on right side not ossified, appearing fused into single cartilaginous plate along with scapula; posttemporal possessing single reduced lateral shelf; ventral postcleithrum present.

Vertebral column and caudal skeleton (Fig. 8): precaudal vertebrae 11; caudal vertebrae (including terminal element) 16; vertebrae 3-11 with pleural rib articulating with respective parapophyses; twelve pairs of epineurals, articulating with vertebrae as follows: epineurals 1, 2 with corresponding parapophyses, 3-10 with corresponding pleural ribs, epineural 11 reduced and not articulating with pleural rib and 12 highly reduced and embedded in tissue; first caudal vertebra with expanded haemal arch and tiny haemal spine; haemal arch comprising elongate parapophyses, "rib-like" in appearance in lateral view; where parapophyses join ventrally to form haemal arch, a nubbin of bone (haemal spine) projecting posteriorly (Fig. 8B); "rib-like" lateral side of haemal arch easily confused with pleural ribs on radiographs; dorsal fin pterygiophore formula 3(221110); 2 pterygiophores of anal fin preceding first haemal arch; hypurals 1-2 not fused with hypurals 3-4; foramen on neural arches 6-22 singular and very large; neural arches 1, 24 through 27 with no foramen; neural arches 2, 3, and 4 with single small foramen; neural arch 5 with two foramina.

Hyoid series (Fig. 9): branchiostegals 5, first branchiostegal small, thin, and attached to ventral surface of narrow portion of ceratohyal, following three branchiostegals blade-like in appearance and attached at ventrolateral surface of broad portion of ceratohyal; posteriormost branchiostegal broadest, articulating with ventral surface of epihyal; basihyal broad, spatulate, slightly emarginate, with only posterior third ossified (Fig. 9); ventral hypohyals, dorsal hypohyals and urohyal ossified; urohyal extremely thin; basibranchials 2, 3 ossified; basibranchial 4 either absent, unstained or too small to discern; hyobranchials 1, 2, and 3 cartilaginous; ceratobranchials 1, 2, 3, 4 cartilaginous at tips; ceratobranchial 5 completely cartilaginous and fused with lower pharyngeal plate; lower pharyngeal plate ossified, possessing large conical teeth with slightly recurved tips.

Myology (Fig. 10): superficial jaw musculature composed of adductor mandibulae 1, adductor mandibulae 1 and adductor mandibulae 2; adductor mandibulae 1 and 1 combined along most of their length and originating on pterotic and ventrally along anterior edge of preopercle; adductor mandibulae 1 separated anteriorly from adductor mandibulae 1 and inserting on primordial ligament near its anterior attachment to maxilla, then running ventrally along primordial ligament and combining with fibers of adductor mandibulae 1; adductor mandibulae 1 with two insertions, along ventral section of primordial ligament and separately by tendon attached to coronoid process; adductor mandibulae 1 and 2 differentiated by presence of ramus mandibularis V, visible between muscle masses; adductor mandibulae 2 originating along preopercle, ventral to adductor mandibulae 1 and continuing ventrally to end of preopercle; adductor mandibulae 2 inserting via tendon onto coronoid process of dentary, joining with tendon of adductor mandibulae 1 before inserting on coronoid process.

Habitat: Antilligobius nikkiae is known from collections or observations from Curacao, Bahamas, Cuba, Belize, Chinchorro Bank off eastern Yucatan (Quintana Roo) and Puerto Rico. It is a calcareous wall associate most often found in large numbers on the glacial sea level plateau at around 100 m (Grant Gilmore pers. comm.). Records of the habitat and behavior of this species are described below.

Belize: Colin (1974) recorded what is probably A. nikkiae, which he listed as "filamentous goby." His observations were made from a submersible on deep reef escarpments at Tobacco Reef and Glovers Reef, Belize, between 90 and 170 m (Colin 1974). The fish occurred in schools of hundreds of individuals and may have been the most common fish observed between 120 and 150 m in these areas. The gobies were reported as being slender, about 5 cm in length, with a silver body and blue eyes (consistent with the appearance of A. nikkiae in the field) and the first dorsal fin was extremely long and held erect. No specimens were collected or photographed in Belize.

Bahamas: The first series was collected by SCUBA diving at Acklins Island at 73 m in 1973 by one of us (PC). Small groups were found hovering along a steep drop-off in small indentations in the reef face. Specimens were returned to the surface alive, and some were maintained for several months in aquaria. Dennis et al. (2004) reported the species to be common at 90-150 m off Lee Stocking Island, Exuma chain, Bahamas.

Cuba: Antilligobius nikkiae (Fig. 11) was captured using crystalline rotenone in DMSO from the Johnson-Sea-Link II submarine, Dive 3,069 on 27 December 1997 at a depth of 169 to 198 m. It was collected at Cayos los Indios, 21.78405[degrees]N, 83.23625[degrees]W, off the west coast of the Isla de Juventud and Golfo de Batabano (Grant Gilmore pers. comm.). Gilmore observed A. nikkiae in small schools typically on vertical calcareous walls, hovering amidst the gorgonians and sponges, hanging along the wall margin. He has also observed them quite often over the sandy terrace that marks the last glacial sea level beach, typically at a depth of around 100 m. He and his colleagues have observed this species in San Salvador, Abaco, New Providence, Grand Bahama and Cat Island in the Bahama Islands; Chinchorro Bank off eastern Yucatan (Quintana Roo); and Terneffe, Glovers and Lighthouse reefs off Belize. Observations in Cuba were limited to the southern shore west of Cayo Largo to Cabo San Antonio. The Johnson-Sea-Link II observations were made from 1978 to 1997.

Curacao: Antilligobius nikkiae has been observed by technical divers, on talus slopes, starting at depths of 90 m and was quite common at 100 m (Fig. 3). They hovered a few inches to a foot above the substrate in small groups of 3-4, or in larger groups exceeding 24 individuals where they appeared to hunt for copepods or other zooplankton (Forrest Young, pers. comm.).

Puerto Rico: Specimens of A. nikkiae were collected on two occasions at 87-90 m depth on the shelf edge reef at La Parguera, Puerto Rico (Fig. 2). Two fish were taken on 22 November 1978 (ANSP 150387) while four specimens were collected on 28 August 1979 and later lost in the mail. Collection dates for these two fish from November 1978 are incorrectly listed in Dennis et al. (2004) as May 1978. Field notes record one deep dive on May 1978, but no fish were collected on that dive. Additional fish were seen on 21 August 1979 at another area along the shelf edge reef with similar geomorphology. Dennis et al. (2004) recorded the species from south-western Puerto Rico at 91 m. The gobies were found to hover slightly above the bottom, remaining close to one another and often facing in the same direction. The reef slope where the gobies were captured was steep,with an overall slope of about 45[degrees] from 45 to 90 m depth, then becoming near vertical below 90 m. The gobies were found among small indentations in the rocky structure of the reef face where the slope transitions between steep and vertical. They were found with other fish species typical of the deep vertical reef environment, including Ptereleotris helenae, Chromis enchrysura, Chromis scotti, Centropyge argi, Chromis insolata, Liopropoma mowbrayi, Liopropoma carmabi, Gramma linki and juvenile Lutjanus buccanella. The sabre gobies collected from Puerto Rico were captured alive using quinaldine-alcohol solution and returned directly to the surface with the SCUBA divers, where they were maintained in aquaria.

Distribution: Known from Belize, Bahamas, Puerto Rico, Cuba, Mexico and Curacao. The distribution is most likely widespread throughout the Caribbean wherever the glacial sea level plateau occurs.

Etymology: The specific epithet nikkiae is for Nicole Laura Schrier, the daughter of Adriaan "Dutch" Schrier, owner of the Sea Aquarium in Curacao, who collected many of the type specimens. The species is given the common name Sabre goby in reference to its long, filamentous first dorsal fin that resembles a sabre (a long sword with a curved blade).

Bollmannia Jordan, 1890

Bollmannia Jordan, in Jordan & Bollman, 1890: 164 (type species Bollmannia chlamydes Jordan by monotypy).

Diagnosis: Bollmannia can be distinguished from other Gobiosomatini genera by the following combination of characters, not listed in order of taxonomic importance: first dorsal of most species with black spot or blotch on posterior portion of fin; second dorsal I,11-15; anal I,10-15; pelvic fins with well-developed frenum supported with collagenous thickenings, frenum with scalloped posterior margin; caudal fin lanceolate; trunk scales ctenoid; predorsal region, cheek, pectoral fin base and breast with cycloid scales; predorsal scales 7-10.

Description: Body slightly elongate and laterally compressed; first dorsal VII; pterygiophore formula of first dorsal fin 3(221110); spines of dorsal fin frequently elongate in both sexes; second dorsal I,11-15; first and second dorsal fins separate from each other and from caudal fin; pectoral rays 19-26; anal I,10-15; trunk scales ctenoid; lateral scale rows 24-31; predorsal region, cheek, pectoral fin base and breast with cycloid scales; predorsal scales 6-10; scales highly deciduous; first pterygiophore of second dorsal fin inserted behind ninth neural spine, remaining pterygiophores arranged in 1:1 ratio with underlying vertebrae; first two anal pterygiophores inserted anterior to first haemal spine, remaining pterygiophores in 1:1 ratio with caudal vertebrae; total vertebrae 27, 11 precaudal, 16 caudal; caudal fin long, 30-50% SL, and lanceolate; pelvic fin with one spine and five rays, fins united, forming disk with well-developed frenum, frenum with thickened collagenous supports and scalloped margin (Fig. 12); teeth in both jaws arranged in multiple rows, teeth in inner and outer rows typically enlarged, evenly spaced, and more caninoid; teeth in midde rows smaller and more irregularly spaced; vomerine teeth absent; mouth large, 12-17% SL and inclined vertically; eye large, diameter 7.5-11% SL; gill opening extending entire length of pectoral fin base; gill rakers on lower arm of first arch elongate, rakers on upper arm short, stubby and poorly ossified; urogenital papilla thin and conical in males, thick and bulbous in females; cephalic lateralis canal and pore system on head well-developed (Fig. 13); sensory papillae pattern in transverse pattern (Fig. 13); ventral post-cleithrum present; nearly all species with dark conspicuous spot on posterior portion of first dorsal fin and sometimes on caudal peduncle; sexual dimorphism generally absent, with the exception of slightly larger dorsal fins in males of B. boqueronensis, and subtle pigmentation differences in B. communis.

Bollmannia boqueronensis Evermann & Marsh, 1899

White-eye goby (Figs 13B, 14)

Bollmannia boqueronensis Evermann & Marsh, in Evermann & Marsh 1899: 356 (type locality Puerto Real, Puerto Rico).

Bollmannia jeannae Fowler, in Fowler 1941: 95, Figs7-9 (type locality Key West, Florida).

Material examined: USNM 49366, holotype of B. boqueronensis, 15.5 m, Puerto Real, Puerto Rico; USNM 204311, paratype, 3, 15.5 m, Puerto Real, Puerto Rico; UF 172806, 2, 55-64 m, Cordoba, Colombia, 12 July 1966; AMNH 253405, 2 plus 1 cleared and stained, 29-32.2 mm SL, 8.4 m, JVT-04-306, Bocas del Toro, Laguna de Chiriqui, Panama, 9 June 2004, Robertson et al.; AMNH 253406, 4, 20-42.5 mm SL, 14.3 m, JVT-04-317, Bocas del Toro, 09.422[degrees]N, 81.910[degrees]W, Panama, 12 June 2004, Robertson et al.; UF 223444, 12, 36 m, Gulfo de Morrosquillo, Colombia, 13 July 1966; UF 223664, 5, 29-42 m, Caribbean Sea, N of Cabo Tiburon, Panama, 18 July 1966; USNM 116363, 1, 14.17 m, Key West, Florida, 13 February 1902; USNM 290282, 2, 37 m, Gulf of Mexico, Silver Bay Station 54, 28.150[degrees]N, 83.830[degrees]W, Florida, 16 July 1957; USNM 116366, 1, 49 m, Gulf of Mexico, 26.5583N[degrees], 83.2583[degrees]W, Florida, 18 March 1885; USNM 118088, 2, Tortugas Islands, Florida, 24 July 1924; USNM 186228, 3, 73 m, Gulf of Venezuela, 12.2417[degrees]N, 70.33[degrees]W, Venezuela, 10 November 1958; AMNH 239100, 1, 47.7 mm SL, 1521 m, Spaanse Water, 12.069[degrees]N, 68.8519[degrees]W, Curacao, 11 February 2005, Robertson et al.; AMNH 239103, 1, 49.4 mm SL, 15-21 m, Spaanse Water, 12.0697[degrees]N, 68.8519[degrees]W, Curacao, 11 February 2005, Robertson et al.; AMNH 239083, 1, 45.22 mm SL, 15 m, Spaanse Water, 12.0697[degrees]N, 68.8519[degrees]W, Curacao, 10 February 2005, Robertson et al.; ANSP 69718, Bollmannia jeannae holotype, 58.8 mm SL, 34.7 m, Key West, Florida, 12 April 1940; ANSP 69719, B. jeannae paratype, 52.7 mm SL, 34.7 m, Key West, Florida, 12 April 1940; UPRM 3392, 1, 45.8 mm SL, 18 m, east end of Laurel Reef, Puerto Rico, 7 September 1975; UPRM 3788, 3, 37.3-44.0 mm SL, no collection data (possibly collected with UPRM 3790); URPM 3791, 1, 51.8 mm SL, no collection data (possibly collected with UPRM 3790); UPRM 3790, 49.0-57.0 mm SL, 18 m, Laurel Reef, La Parguera, Puerto Rico; UF 226784, 3, 29.8-32.8 mm SL, Golfo de Morrosquillo, 09.521667[degrees]N, 76.256667[degrees]W, Colombia, 13 July 1966; UF 231124, 24.5-27.5 m, field stations PLC-J19, Caribbean Sea, Jamaica, 13 February 1971, P. L. Colin and N. Copland.

Diagnosis: Second dorsal fin typically I,12; anal fin typically I,12; eye diameter 7.7-10% SL; spines of first dorsal fin not distinctly elongate; first and second dorsal fins with 3-5 brown or tan horizontal stripes; upper lip lightly pigmented to pale, no dark stripe above upper lip; pelvic disk lightly pigmented to pale; body with 4-5 dark brown blotches along lateral midline; blotches often connected by slightly lighter horizontal band of pigment along lateral midline; longitudinal papillae row b originating at vertical through anterior margin of pupil.

Description: Body somewhat laterally compressed; depth at origin of first dorsal fin 17.5-21.8% SL; least depth of caudal peduncle 9.9-10.5% SL; size to 100 mm TL (Robins & Ray 1986).

Median and paired fins: first dorsal VII, spines not notably elongate; second dorsal I,11(8), I,12*(35), I,13(2); anal I,11*(7), I,12 (36), I,13(2); pectoral rays 19(3), 20(2), 21*(7), 22(5), 23(1), longest rays reaching posteriorly to vertical through anus; pelvic I,5*(10), fifth ray longest, reaching posteriorly to or just falling short of anus; pelvic fin with well-developed frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(1), 17(7), 19*(1), branched rays 14(6), 15(1).

Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; lateral scale rows 27-31; transverse scale rows 7; predorsal scales 9-10, scales extending anteriorly to vertical behind eye; caudal peduncle scales 11-13; no modified basicaudal scales present.

Head: head length 27.0-31.0% SL; mouth large, angled upwards approximately 20-30 degrees from horizontal; mouth reaching posteriorly to vertical through pupil; upper jaw length 11.0-13.0% SL; teeth in both jaws arranged in three rows; teeth in inner and outer rows enlarged, regularly spaced and caninoid; teeth in middle row smaller and irregularly spaced; eye diameter 7.7-10.0% SL; interorbital narrow, width less than diameter of pupil; snout short, 4.8-5.5% SL; postorbital length 12.8-16.5% SL; gill rakers on lower arm of first arch 10, upper arm 3; gill rakers on lower limb thin and elongate, rakers on subsequent arches shorter and thicker.

Genitalia: papilla elongate and conical in males, no melanophores present; papilla short and bulbous in females.

Sensory papillae and head pores (Fig. 13B): oculoscapular pores B', C(s), D(s), E, F, G, H', K' and L' present; in approximately half of specimens examined, anterior canal terminating with pore H', other half of specimens with anterior canal terminating with pore posterior to H, herein termed J', which may not be homologous with J sensu Akihito (1984); preopercular pores M', N, O' present; papillae in transverse pattern with 4-5 rows present below eye; first transverse row beginning approximately below anterior margin of eye; last transverse row beginning below posterior margin of pupil; transverse rows often extending below longitudinal row b; row b beginning at vertical through anterior margin of pupil; row d beginning slightly posteroventral of first transverse papillae row, at vertical through anterior one-third of eye and continuing posteriorly to below posterior margin of eye, sometimes slightly further posteriorly; row n not continuous across dorsal midline, stopping slightly before pore D; row x beginning at ventral termination of row n and continuing posteriorly to just before posterior margin of operculum; two horizontal rows of papillae on operculum, one beginning slightly above and behind pore N, the other slightly below and behind pore O'.

Pigmentation: following description based on recently collected specimens or preserved specimens with well-preserved pigmentation. Overall body background color pale; body with 4-5 dark brown blotches along lateral midline; blotches typically connected by slightly lighter horizontal stripe of pigment along lateral midline; thin, light brown or tan horizontal stripe of pigment below dorsal midline and above lateral midline, beginning behind eye and continuing to base of caudal fin; this stripe often broken or incomplete and becoming less distinct posteriorly; abdomen and gular regions pale; in life, head often with bright iridescent orange vertical markings on preoperculum and operculum and orange stripes on upper and lower jaws; dorsal fins with 3-5 brown or tan horizontal stripes separated by pale or translucent stripes; posterior portion of first dorsal fin with distinct black spot ranging in size from as small as pupil to as large as eye; anal fin completely pale or lightly pigmented at base; pelvic fin lightly pigmented to pale; upper half of caudal fin often with 2-3 horizontal or slightly diagonal brown or tan stripes similar to those on dorsal fins; midline of caudal fin often with brown horizontal stripe continuing from stripe on lateral midline of body; lower half of caudal fin pale; pectoral fin base and rays pale.

Habitat: Specimens in this study were collected over sand and sandy-mud bottoms and flat areas with fine sediment adjacent to coral reefs at depths from 8.4-73.0 meters.

Distribution: Specimens examined in this study were collected from the eastern Gulf of Mexico, Florida Keys, Tortugas, Puerto Rico, Jamaica, Colombia, Panama, Curacao and Venezuela. Additional museum lots exist, but have not been examined, from Guatemala (UF 15661), Honduras (UF 288135-288136), Martinique (ANSP 108832), and Bahia, Brazil (MNHN 1989-0786). MorenoMendoza et al. (2011) also report B. boqueronensis from Alacranes Reef Marine Park, Mexico.

Remarks: Fowler's description of B. jeannae stated that B. jeannae differs from B. boqueronensis in coloration, pectoral fin length and cheek scale size (Fowler 1941). Ginsburg (1942) stated that the differences do not hold up when B. jeannae is compared to actual specimens of B. boqueronensis and that the illustrations of B. boqueronensis that accompany Evermann & Marsh's (1899) description are inaccurate. Ginsburg concluded that B. jeannae, which is known from only the holotype and one paratype, is almost certainly a synonym of B. boqueronensis. He offered no formal synonymy of the two species. Bollmannia jeannae falls within the range of B. boqueronensis in all measurements taken here. The coloration differences between the two species as described by Fowler (1941) do not hold when live or freshly preserved specimens of B. boqueronensis are compared, nor do the differences in scale size. The B. jeannae types differ from the B. boqueronsensis holotype in second dorsal and anal fin ray counts. Both type specimens of B. jeannae have I,11 in the second dorsal fin and I,12 in the anal fin, whereas B. boqueronensis was described as having I,12 in the second dorsal and I,13 in the anal fin. Based on our analysis, both I,11 and I,12 are counts that have been observed in both fins of B. boqueronensis. For these reasons, we agree with Ginsburg (1942) and consider B. jeannae to be a synonym of B. boqueronensis.

Evermann & Marsh (1899) used the presence of a series of 3-5 large scales on the lower margin of the cheek as a unique and distinguishing characteristic for B. boqueronensis. Although this row of scales is rare or nonexistent in other species of Bollmannia, the deciduous nature of the scales on all species of Bollmannia diminishes the diagnostic utility of this character. Some specimens of B. boqueronensis examined in this study were missing some or all of these scales. Because scale pockets on the face are also frequently destroyed, it is difficult to determine if cheek scales were present at some point and lost during collection, or if scales were absent and thus this character is somewhat variable.

Robins & Ray (1986), Humann & Deloach (2002) and McEachran & Fechhelm (2005) report B. boqueronensis as sometimes having orange and yellow spots on the dorsal fins, but these observations were almost certainly based on misidentified specimens of B. litura (which has been collected with B. boqueronensis), or B. communis.

Ginsburg (unpublished manuscript) described B. boqueronensis as having an anterior oculoscapular lateralis canal ending with pore H' (missing pore J'). Ginsburg used the absence of this character to divide Bollmannia into two distinct groups (proposed subgenera), with B. litura and B. communis both possessing pore J'. Our study shows this character to be highly variable in B. boqueronensis, as approximately half of the specimens examined have pore J'. The variation in this character does not appear to be correlated with geographic distribution, sexual dimorphism or size.

Two specimens of Bollmannia from Alacranes Reef, Mexico were identified as B. boqueronensis by Moreno-Mendoza et al. (2011) despite possessing one spine and 13 rays in the anal and second dorsal fins. This combination of counts is more typical of B. communis, which is commonly reported from the western Gulf of Mexico; however, we agree with the identification of Moreno-Mendoza et al. (2011). Their specimens possess all of the diagnostic pigmentation features of B. boqueronensis described here and were also collected from sandymud sediment at the base of a coral reef, which is a common habitat for B. boqueronensis. Bollmannia communis is common over thick mud and is not typically associated with reef habitats.

Comparisons: Bollmannia boqueronensis can be distinguished from B. communis in usually having fewer anal and second dorsal elements (typically I,12 in B. boqueronensis vs. typically I,13 in B. communis). It differs from B. litura and B. communis in having a lightly pigmented pelvic fin (vs. a dark pelvic fin), in having tan or brown stripes on the dorsal fin (vs. having yellow or orange stripes or spots) and in having dark spots and a stripe of pigment along the lateral midline (vs. dark spots that are rarely connected by a stripe). Bollmannia boqueronensis can be distinguished from B. eigenmanni in lacking a distinct dark stripe of pigment on or above the upper lip, having a slightly deeper caudal peduncle (9.9-10.5% SL in B. boqueronensis vs. less than 10% SL in B. eigenmanni), having three pores in the preopercular series (vs. two pores in the peropercular series) and in having longitudinal papillae row b originating at a vertical through the anterior margin of pupil (vs. originating more posterior in B. eigenmanni).

Bollmannia communis Ginsburg, 1942

Ragged goby (Figs 13D, 15)

Bollmannia communis Ginsburg, in Ginsburg 1942: 634 (type locality Gulf of Mexico, Louisiana).

Materials examined: USNM 119873, holotype, 1 male, 52.5 mm SL, 18 m, Gulf of Mexico, NW of Southwest Pass entrance, of Mississippi Delta, 28.983333[degrees]N, 89.483333[degrees]W, Louisiana, 7 September 2006; USNM 119879, paratype, 40 m, Gulf of Mexico, 28.4667[degrees]N, 91.2250[degrees]W, Louisiana, 30 January 1939; USNM 119880, paratype, 27 m, Gulf of Mexico, NW of Southwest Pass entrance, off Mississippi River Delta, 28.970[degrees]N, 89.475[degrees]W, Louisiana, 10 November 1938; USNM 119881, paratype, 68 m, Gulf of Mexico, 27.7333[degrees]N, 96.2917[degrees]W, Louisiana, 23 January 1939; USNM 119882, paratype,55 m, Gulf of Mexico, 28.800[degrees]N, 89.850[degrees]W, Louisiana, 13 May 1938; USNM 119883, paratypes, 7, 77 m, Gulf of Mexico, 27.6667[degrees]N, 96.5667[degrees]W, Texas, 22 March 1938; USNM 119884, paratype, 46 m, Gulf of Mexico, 26.7167[degrees]N, 96.8500[degrees]W, Texas, 4 February 1939; USNM 119885, paratypes, 6, 64 m, Gulf of Mexico, 27.083333[degrees]N, 96.825[degrees]W, Texas, 31 January 1939; USNM 119886, paratypes, 3, 64 m, Gulf of Mexico, 28.825[degrees]N, 89.750[degrees]W, Louisiana, 13 May 1938; USNM 119887, paratypes, 5, 64 m, Gulf of Mexico, east of Mustang Island, 27.7083[degrees]N, 96.3583[degrees]W, Texas, 23 January 1939; USNM 119888, paratype, 31 m, Gulf of Mexico, east of Mustang Island, 27.725[degrees]N, 96.730[degrees]W, Texas, 22 April 1938; USNM 119890, paratype, 82 m, Gulf of Mexico, 26.5667[degrees]N, 96.5333[degrees]W, Texas, 4 February 1939; USNM 119892, paratypes, 7, 46.4-68.5 mm SL, 60 m, Gulf of Mexico, 27.2167[degrees]N, 29.7833[degrees]W, Texas, 30 January 1939; USNM 119893, paratypes, 50.3 m, Gulf of Mexico, east of Mustang Island, 27.6667[degrees]N, 96.5667[degrees]W, Texas, 22 April 1938; USNM 197731, 20, Gulf of Mexico, 26.2700[degrees]N, 82.0500[degrees]W, Florida, 21 March 1962; USNM 337773, 6, 44 m, Gulf of Mexico, Campeche Bank, 110 Miles of (SW) Campeche, Yucatan, 19.600[degrees]N, 91.7917[degrees]W, Mexico, 19 August 1951; USNM 364236, 11 of 12, 64 m, 6.90000[degrees]N, 53.96667[degrees]W, French Guiana, 30 June 1972; TCWC 6829.10, 5, 29.3-54.0 mm SL, 47.5 m, Gulf of Mexico, Mississippi/Alabama, 12 March 1988; AMNH 253407, 16, Gulf of Mexico, NOAA Station 022, Texas, October 2006; AMNH 253408, 12, Gulf of Mexico, Texas, NOAA Station 032, October 2006; AMNH 253409, 12, Gulf of Mexico, Texas, NOAA Station 045, October 2006; UPRM 2196, 1, 40 mm SL, off Anasco, Puerto Rico, 7 August 1964, J. Randall; UPRM 3788, 2, 45-48 mm SL, off Rio Anasco, Puerto Rico, 11 August 1987, B. Yoshioka; UPRM 1551, 42 mm SL, Anasco, Puerto Rico, 29 March 1963, Randall et al.; GCRL 17540, 58, 41.0-49.0 mm SL, Gulf of Mexico, 28.952167[degrees]N, 89.683833[degrees]W, Louisiana, 26 August 1978; GCRL 28344, 2, 64.2-71.5 mm SL, 35 m, Gulf of Mexico, 29.300333[degrees]N, 88.902833[degrees]W, Louisiana, 9 November 1997, W. E. Aguirre.

Diagnosis: Second dorsal fin typically I,13; anal fin typically I,13; eye diameter 8.1-10% SL; middle spines of first dorsal slightly elongate; first and second dorsal fins with dark circles or ellipses with orange or yellow centers (centers pale in preservation); upper lip lightly pigmented to pale, no dark stripe above upper lip; pelvic fin heavily pigmented, black or nearly so; body with 4-5 dark brown blotches along lateral midline; blotches typically not connected by slightly lighter horizontal stripe of pigment along lateral midline; longitudinal papillae row b originating at vertical through anterior margin of pupil.

Description: Body somewhat laterally compressed, depth at origin of first dorsal fin 19.2- 21.1% SL; least depth of caudal peduncle 9.4-11.6% SL; size to 100 mm SL (Robins & Ray 1986).

Median and paired fins: first dorsal VII, middle spines slightly elongate in some specimens; in males, fourth spine reaching posteriorly to base of fourth to sixth ray of second dorsal fin when depressed; spines slightly shorter in females; second dorsal I,12(23), 1,13*(82), I,14(3); anal I,11(1), I,12(15), I,13*(84), I,14(8); pectoral rays 19(3), 20(4), 21(4), 22(8), 23*(4), longest rays extending posteriorly to vertical through anus; pelvic I,5*(10), fifth ray longest, extending to anus or falling just short of anus; pelvic fin with well-developed frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(1), 17*(5), branched rays 14*(6).

Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; scales in lateral series 24-29; transverse scale rows 6-7; predorsal scales 8-9, scales extending anteriorly to vertical behind eye; caudal peduncle scales 10-13; no modified basicaudal scales present.

Head: head length 31.0-33.5% SL; mouth large, angled upwards approximately 20-30 degrees from horizontal; mouth reaching posteriorly to vertical through pupil; upper jaw length 12.7-14.5% SL; teeth in both jaws arranged in three rows; teeth in inner and outer rows enlarged, regularly spaced and caninoid; teeth in middle row smaller and irregularly spaced; eye diameter 8.1-10.0% SL; interorbital narrow, width less than diameter of pupil; snout short, 5.8-7.3% SL; postorbital length 14.2-16.5% SL; gill rakers on lower arm of first arch 10, upper arm 3; gill rakers on lower limb thin and elongate, rakers on subsequent arches shorter and thicker.

Genitalia: papilla elongate and conical in males, occasionally with melanophores on lateral edges; papillae short and bulbous in females.

Sensory papillae and head pores (Fig. 13D): oculoscapular pores B', C(s), D(s), E, F, G, H, J' and K', L' present; anterior canal terminating in pore J'; preopercular pores M', N, O' present; papillae in transverse pattern, with 4-5 transverse rows present below eye; first transverse row beginning approximately below anterior margin of eye; last transverse row beginning below posterior margin of pupil or slightly more posterior; middle transverse rows highly variable, frequently disjunct, sometimes extending below longitudinal row b; transverse rows on opposite sides of head often varying in a single individual; row b originating at vertical through anterior margin of pupil and continuing posteriorly to below posterior margin of eye, sometimes extending slightly further posteriorly; row d beginning slightly posteroventral of bottom of first transverse row, at vertical through anterior margin of pupil and continuing posteriorly to below posterior margin of eye, sometimes extending slightly further posteriorly; row n continuous across dorsal midline; row x beginning at ventral termination of row n and continuing posteriorly to above pore L'.

Pigmentation: following description based largely on recently preserved specimens. Comments on coloration in life based on color photographs of B. communis specimens from Gulf of Mexico (Fig. 15) which we have not examined. Overall body background color pale; body with 4-5 faint brown blotches along lateral midline; blotches rarely connected by slightly lighter horizontal stripe of pigment along lateral midline; abdomen and gular regions pale; top of head, snout, interorbital and area above jaw uniformly pigmented with scattered small dark melanophores; occasionally faint elongate patches of pigment present on preoperculum directly beneath and behind eye; base of first dorsal fin with two yellowish orange stripes in life (pale in preservation), stripes becoming broken distally and forming distinct ellipses with dark margins and bright yellow or orange centers in life; yellow and orange pigment on fins fading to pale in preservation; posterior portion of first dorsal fin with distinct black spot ranging in size from as small as pupil to as large as eye; distal margin of first dorsal fin with black stripe in males; second dorsal fin pigmented with ellipses similar to those of first dorsal fin; anal fin nearly pale at base, becoming more heavily pigmented distally; pelvic fin heavily pigmented, black or nearly so; upper half of caudal fin pigmented with ellipses similar to dorsal fins; lower half of caudal fin with two faint yellowish-orange diagonal stripes in life, fading in preservation; pectoral fin base pale; pectoral rays dusky to pale.

Habitat: Specimens in this study were collected from mud bottoms at depths of 18-77 m.

Distribution. Known primarily from the Gulf of Mexico, with one confirmed record from French Guiana. Several lots of Bollmannia from Puerto Rico, (UPRM 1551, UPRM 2196, ANSP 147645) were tentatively listed as B. communis by Dennis et al. (2004); however, these specimens are in very poor condition and it is unclear as to whether these specimens are B. communis or B. litura.

Comparisons: Bollmannia communis and B. litura are the most similar of all western Atlantic Bollmannia species. The two species both have similar sensory papillae and sensory pore/canal patterns, nearly identical median fin pigmentation and lack a black band on the upper lip. Bollmannia communis can be distinguished from B. litura in having a smaller eye (typically less than 10% SL in B. communis vs. typically more than 10% SL in B. litura) and from B. litura and all other species in having higher modal counts in the second dorsal and anal fins (typically I,13 in B. communis vs. typically I,10-12 in other western Atlantic Bollmannia species). Male B. communis have dark distal margins on the first dorsal fin whereas male B. litura do not. Bollmannia communis can be further distinguished from B. boqueronensis by pigmentation, as B. boqueronensis has tan or brown stripes on the dorsal fins, a pale pelvic disk and usually a brown stripe of pigment on the lateral midline and B. communis typically has yellow or orange (pale in preservation) stripes and ellipses on the dorsal fins, a dark pelvic disk and no stripe of pigment on the lateral portion of the body. Bollmannia communis can be distinguished from B. eigenmanni in having three pores on the preoperculum (M', N, O') vs. having two (M', O'), having a dark pelvic fin (vs. a pale pelvic fin), lacking a distinct black stripe on or above the upper jaw (black stripe present in B. eigenmanni) and in having longitudinal papillae row b originating at a vertical through anterior margin of the pupil (vs. originating more posterior to this point in B. eigenmanni).

Bollmannia eigenmanni (Garman, 1896)

Shelf Goby (Figs 13C, 16)

Gobius eigenmanni Garman, in Garman 1896:88, Pl. 3 (type locality Key West, Florida).

Materials examined: USNM 231318, holotype, 1 male, 26.12 mm SL, 110 m, Key West, Florida, 1893; USNM 116368, 1, 37.6 mm SL, 55-110 m, Gulf of Mexico, 28.74167[degrees]N, 85.1500[degrees]W, Florida, 15 March 1885; UF 210982, 1, 23.3 mm SL, 36.5-40 m, Gulf of Mexico, South of Cailou Bay, 28.441667[degrees]N, 91.008333[degrees]W, Louisiana, 12 September 1962; UF 228128, 1, 57-68 m, north of Isla de Margarita, 11.48333[degrees]N, 63.85000[degrees]W, Venezuela, 20 July 1968; AMNH 83384, 1, 20.1 mm SL, Gulf of Mexico, 29.805[degrees]N, 86.691667[degrees]W, Florida, 29 August 1976; UF 153662, 1, 36.2 mm SL, 41 m, Gulf of Mexico, 29.75[degrees]N, 86.00[degrees]W, United States, 5 June 1974; UF 228129, 1, 23.8 mm SL, 43-46 m, Golfo Triste, N of Puerto Cabello, 10.755[degrees]N, 68.138333[degrees]W, Venezuela, 26 July 1968; UF 230139, 1, 27.8 mm SL, 150-170 m, W of Cartagena, 10.365[degrees]N, 75.788333[degrees]W, Colombia, 1 August 1968; AMNH 85997, 1, 24.75 mm SL, Gulf of Mexico, 29.5000[degrees]N, 86.418667[degrees]W, United States, 7 February 1978; AMNH 83254, 1, Gulf of Mexico, 29.90500[degrees]N, 87.11667[degrees]W, United States, 28 August 1976; YPM 3987, 1, 84 m, Gulf of Mexico, 29.117[degrees]N, 88.717[degrees]W, Louisiana, 10 April 1937, R/V Atlantis; YPM 3986, 4, 51 m, Gulf of Mexico, 28.317[degrees]N, 90.983[degrees]W, Louisiana, 25 March 1937, R/V Atlantis; YPM 3983, 4, 79 m, Gulf of Mexico, 29.117[degrees]N, 88.717[degrees]W, Louisiana, 10 April 1937, R/V Atlantis; YPM 3921, 3, 79 m, 29.117[degrees]N, 88.717[degrees]W, Louisiana, 10 April 1937; TCWC 6829.32, 3, 24.7-30.1 mm SL, 47.5 m, Gulf of Mexico, 29.454722[degrees]N, 88.595000[degrees]W, Mississippi and Alabama, 12 March 1988; TCWC 6829.33, 1 cleared and stained, 28.3 mm SL, 47.5 m, Gulf of Mexico, 29.454722[degrees]N, 88.595000[degrees]W, Mississippi and Alabama, 12 March 1988; GCRL V69:3460, 1, 34 mm SL, 15.41667[degrees]N, 61.20000[degrees]W, Dominica, 5 March 1966, M/V Oregon. GCRL V81:17057, 1, 27.6 mm SL, 68.5 m, off Port O'Conner, Gulf of Mexico, 27.83333[degrees]N, 95.6667[degrees]W, Texas, 8 September 1980, R. Mulcahy.

Diagnosis: Second dorsal fin typically I,11; anal fin I,10-12; anterior spines of first dorsal fin elongate; eye diameter 6.9-10.0% SL; upper lip with distinct dark stripe of pigment on or directly above lip; pelvic disk lightly pigmented to pale; side of body with no distinct markings; longitudinal papillae row b originating at vertical through posterior margin of pupil.

Description: Body somewhat laterally compressed, depth at origin of first dorsal fin 16.0-20.0% SL; body depth at caudal peduncle typically 7.8-10% SL (two specimens reaching 10.5% SL); size to 37 cm SL.

Median and paired fins: first dorsal VII, first three spines elongate, third spine longest, sometimes reaching origin of eighth second dorsal ray when placed flat against back; second dorsal I,10(1), I,11*(12); anal I,10(6), I,11(3), I,12*(4); pectoral rays 19(2), 20(3), 21*(4), rays extending posteriorly to vertical through anus; pelvic I,5, fifth ray longest, reaching posteriorly to or just falling short of anus; pelvic fin with well-developed frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(1), 17(7), 18(1), branched rays 14(4).

Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; lateral scales 27-29; transverse scale rows 7; predorsal scales 7-9, extending anteriorly to behind eye; caudal peduncle scales 10-11; no modified basicaudal scales present.

Head: head length 25.5-32.5% SL; mouth large, angled upward approximately 20-30 degrees from horizontal; mouth reaching posteriorly to vertical through pupil; upper jaw length 12.0-14.0% SL; teeth in both jaws arranged in three rows; teeth in inner and outer rows enlarged, regularly spaced and caninoid; middle row of teeth smaller and irregularly spaced; eye diameter 7.0-10.0% SL; interorbital narrow, width less than diameter of pupil; snout short, 5.2-6.0% SL; postorbital length 13.7-17.1% SL; gill rakers in upper arm 3-4, lower arm 9-10; gill rakers on lower limb thin and elongate, rakers on subsequent arches shorter and thicker.

Genitalia: papilla elongate and conical in males, no melanophores present; papilla short and bulbous in females.

Sensory papillae and head pores (Fig. 13C): oculoscapular pores B', C(s), D(s), E, F, G, H' present; no enclosed posterior canal present over operculum; preopercular pores M', O' present; papillae in transverse pattern, with 4-5 rows present below eye; first transverse row beginning at vertical just anterior to anterior margin of pupil; last transverse row beginning below posterior margin of eye; middle transverse rows somewhat variable, with anterior rows sometimes extending ventrally to row d; row b beginning at vertical through posterior margin of pupil and continuing posteriorly to posterior margin of preopercle; row d beginning slightly posteroventral of first transverse row, at vertical through anterior one-third of eye and continuing posteriorly to vertical through posterior margin of eye and sometimes slightly further posteriorly; row n nearly continuous across dorsal midline, stopping before pore D; row x sometimes broken into two segments, beginning over pore F and ultimately terminating over operculum.

Pigmentation: no fresh or recently preserved specimens of B. eigenmanni were available for description. The museum specimens used in this study had no prominent pigmentation on the body and fins. For the most part the general color on the body and fins of these specimens is pale or dark brown, depending on the method and quality of preservation. The only pigmentation that is apparent after prolonged preservation is the distinct dark band of pigment on or directly above the upper jaw(diagnostic for this species) and a dark spot on the posterior portion of the first dorsal fin, and, in some specimens, a small patch of melanophores at the base of the caudal fin.

Habitat: Some specimens in this study were taken over muddy bottoms. It has been collected at depths of 36.5-170.0 m.

Distribution: Bollmannia eigenmanni is fairly uncommon in collections and is primarily known from the Florida Keys (type locality) and throughout the Gulf of Mexico, where it has been collected in trawls with B. communis. Records also exist from Venezuela. A single specimen from Dominica was previously identified as B. eigenmanni by C. E. Dawson, however, this record is questionable (see Remarks below).

Remarks: When describing B. litura, Ginsburg (1935) noted that specimens of B. eigenmanni were not available for comparison, thus he was forced to rely on the original description of B. eigenmanni for comparison of the two species. In an unpublished manuscript, Ginsburg later noted that the type of B. eigenmanni had been lost. The Gobius eigenmanni holotype (USNM 231318) has since been found at USNM.

A single specimen from Dominica (GCRL 69:3460) was identified by C. E. Dawson as B. eigenmanni, likely due to the second dorsal and anal fin counts (I,11 and I,12 respectively), elongate first dorsal spines and a faint area of pigment above the upper jaw, which is restricted to the very tip of the snout on this specimen. There is no pigment on the remainder of the upper jaw. When compared with other B. eigenmanni this specimen has more pectoral rays (22) and gill rakers on the lower arm of the first arch (12) and also has a slightly shorter jaw, larger eye and shorter postorbital length. In addition, there are 3 pores in the preopercular canal series, whereas all other B. eigenmanni have two. For these reasons we do not consider this specimen to be B. eigenmanni. The identity of this specimen remains unclear and additional specimens are needed to clarify what species occurs in Dominica.

Comparisons: Bollmannia eigenmanni is the most distinctive species of western Atlantic Bollmannia. It differs from all other genera in having 2 pores in the preopercular series (M', O') versus having 3 (M', N, O'), a trait that Ginsburg (unpublished manuscript) used to separate B. eigenmanni into its own subgenus. Longitudinal papilla row b originates at a vertical through the posterior margin of the pupil in B. eigenmanni, whereas row b originates more anteriorly in the other Atlantic species. Bollmannia eigenmanni can also be distinguished from its Atlantic congeners by possessing a distinct dark stripe of pigment on or directly above the upper jaw and in having a shallower caudal peduncle (typically 7.8-10% SL in B. eigenmanni, with two specimens reaching 10.5% SL, vs. typically < 10% SL in other species). It further differs from B. communis in averaging fewer rays in the second dorsal and anal fins (I,10-12, in B. eigenmanni vs. I,12-14 in B. communis) and from B. boqueronensis in having prolonged first dorsal spines.

Bollmannia litura Ginsburg, 1935

Citrine goby (Figs 13A, 17)

Bollmannia litura Ginsburg, in Ginsburg 1939: 1, Pl. 1 (type locality Samano Bay, Dominican Republic).

Materials examined: USNM 93797, holotype, 1 male, 38.4 mm SL, 33-35 m, Caroline station 59, Samana Bay, 19.175[degrees], -69.509[degrees], Dominican Republic, 16 February 1933; USNM 116365, 2 females, 35.9-38.5 mm SL, 31 m, Caroline station 59, Samana Bay, 19.170[degrees]N, 69.451[degrees]W, Dominican Republic, 16 February 1933; UF 141435, 4 of 14, 40.4-55.8 mm SL, 33-35 m, Gulf of Venezuela, Venezuela, 27 July 1968; AMNH 248373, 1, Portobello, 09.68319[degrees]N, 79.61935[degrees]W, Panama, 1 June 2007, Robertson et al.; AMNH 253410, 6 females, 2 males, 31.3-43.4 mm SL, 31.5 m, JVT-04-283, Bocas del Toro, 09.005367[degrees]N, 81.97682[degrees]W, Panama, 6 June 2004, Robertson et al.; AMNH 253411, 3 females, 7 males plus 1 male cleared and stained, 33.4-44.0 mm SL, 34.2 m, JVT-04-284, Bocas del Toro, 09.005367[degrees]N, 81.97682[degrees]W, Panama, 6 June 2004, Robertson et al.; UF 172809, 1, 53-54 m, Caribbean Sea, Colombia, 11 July 1966; USNM 338585, 1 female, field number IU 8849, St. Lucia; UF 141435, 15 of 16, 33-35 m, Golfo de Venezuela, PIL 762, Venezuela, 27 July 1968; UF141434, 10, 55-64 m, Cordoba, PIL 362, Colombia, 12 July 1966; UF 228131, 75, 19.3-41.5 mm SL, 12.8 m, NE of Cabo Res Puntas, 16.02500[degrees]N, 88.70833[degrees]W, Guatemala, 19 March 1968; UF 226730, 4 (labeled as 5), 25.0-49.3 mm SL, 18 m, Golfo de Morrosquillo, 09.59500[degrees]N, 75.62666[degrees]W, Colombia, 13 January 1966; UF 230660, 31 (labeled as 30), 26.5-57.4 mm SL, 35 m, Caribbean Sea, 1186667[degrees]N, 70.366667[degrees]W, Venezuela, 27 July 1968; AMNH 253412, 1, 32.6 mm SL, 25.2 m, JVT-04-280, Bocas del Toro, Panama, 2004, Robertson et al.; UF 223880, 2, 22.2-35.3 mm SL, 36 m, Golfo de Morrosquillo, 09.520000[degrees]N, 75.68500[degrees]W, Colombia, 13 July 1966; UF 230134, 1, 41.3 mm SL, 60-66 m, off Cartagene, 10.34500[degrees]N, 75.65167[degrees]W, Colombia, 1 August 1968; UF 223967, 8 of 14 (2 cleared and stained removed under same number), 28.5-48.3 mm SL, 60-71 m, Caribbean Sea, station PIL 723, Venezuela, 21 July 1968; AMNH 253413, JVT-04-282, 1, 35.5 mm SL, 20.7 m, JVT-04-282, Bocas del Toro, Laguna de Chiriqui, 8.9755[degrees]N, 81.94413[degrees]W, Panama, 6 June 2004, Robertson et al.

Diagnosis: Second dorsal fin typically I,11 or I,12; anal fin typically I,11 or I,12; eye diameter 10.0-12.2% SL (9.5% in two specimens); middle spines of first dorsal fin elongate in some specimens; first and second dorsal fins with dark circles or ellipses with orange or yellow centers (centers pale in preservation); upper lip lightly pigmented to pale, no dark stripe above upper lip; pelvic disk heavily pigmented to black; body with 4-5 dark brown blotches along lateral midline; blotches very rarely connected to one another by narrow stripe of lighter pigment; longitudinal papilla row b beginning at vertical through anterior margin of pupil. Description: Body and head somewhat laterally compressed, depth at origin of first dorsal fin 21.8-23.4% SL; least depth of caudal peduncle 10.0-12.0% SL; size to 57.4 mm SL.

Median and paired fins: first dorsal VII, middle spines slightly elongate in some specimens, fourth spine reaching posteriorly to base of fifth to seventh ray of second dorsal fin with placed flat along back, fifth and sixth spines somewhat shorter, typically not reaching beyond base of fourth ray of second dorsal fin when placed flat along back; second dorsal I,11(17), I,12(36), I,13*(1); anal I,10(1), I,11(13), I,12(41), I,13*(1); pectoral rays 18(1), 19(1), 20(5), 21(4), 22*(2), 23(2), reaching posteriorly to vertical through anus; pelvic I,5*(10), fifth ray longest, extending posteriorly to or just falling short of anus; pelvic fin with welldeveloped frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(3),17*(10), branched rays 13*(3), 14(10).

Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; lateral scale rows 24-28; transverse scale rows 6-7; predorsal scales 7-9, scales extending anteriorly to vertical behind eye; caudal peduncle scales 11-12; and no modified basicaudal scales present.

Head: head length 31.0-33.5% SL; mouth large, angled upward approximately 20-30 degrees from horizontal; mouth extending posteriorly to vertical through pupil; upper jaw length 13.0-15.0% SL; teeth in both jaws arranged in three rows; inner and outer rows of teeth enlarged and caninoid; middle row of teeth smaller than other rows and irregularly spaced; eye large, 10.0-12.2 (9.5 in two specimens) % SL; interorbital narrow, width less than diameter of pupil; snout length 5.5-7.8% SL; postorbital length 12.6-16.0% SL; gill rakers on lower arm 10*(5), upper arm 3*(5); gill rakers on lower first arch thin and elongate, rakers on subsequent arches shorter and thicker.

Genitalia: papilla elongate and conical in males, occasionally with melanophores on lateral edges; papilla short and bulbous in females.

Sensory papillae and head pores (Fig. 13A): oculoscapular pores B', C(s), D(s), E, F, G, H, J'and K', L' present; anterior canal terminating posterior to H in a pore herein termed J', which may not be homologous with J sensu Akihito (1984); preopercular pores M', N, O' present; papillae in transverse pattern, with 4-5 transverse rows present below eye; first transverse row beginning below anterior margin of eye; last transverse row beginning below and slightly behind posterior margin of pupil; middle transverse rows highly variable, often differing on each side of head of a single individual, frequently broken and sometimes extending below longitudinal row b; row b beginning at vertical through anterior margin of pupil, continuing posteriorly to vertical through posterior margin of eye, sometimes slightly further; row d beginning slightly posteroventral of termination of first transverse row, at vertical through anterior one-third of eye and continuing posteriorly to below posterior margin of eye, sometimes slightly further; row n continuous across dorsal midline; row x beginning at ventral termination of row n and continuing posteriorly to above pore L'.

Pigmentation: following description based on recently collected specimens or preserved specimens with well-preserved pigmentation. Overall body background color pale; body with 4-5 faint brown blotches along lateral midline, most conspicuous at base of caudal fin; blotches never connected by slightly lighter horizontal stripe of pigment along lateral midline; abdomen and gular regions pale; in life, head often with bright orange markings vertically on preoperculum and operculum and horizontal orange stripes on upper and lower jaws; dorsal fins with distinct circles or ellipses with dark margins and bright yellow or orange centers in life, with yellow and orange becoming pale in preservation; posterior portion of first dorsal fin with distinct black spot ranging in size from as small as pupil to as large as eye; anal fin nearly pale, in life with 1-2 yellowish orange stripes continuing onto lower half of caudal fin; pelvic fin heavily pigmented, considerably darker than other fins; upper half of caudal fin pigmented with ellipses similar to dorsal fins; lower half of caudal fin with two yellowish orange diagonal stripes in life, stripes fading in preservation; pectoral fin base pale; pectoral rays lightly pigmented, appearing dusky to pale.

Habitat: Occurs at depths of 12.8-71.0 m. Specimens in this study from Panama were collected over mud, mixed mud, shell and leaf litter bottoms.

Distribution: This species has been recorded from Dominican Republic, Venezuela, Panama, Colombia, Guatemala, St. Lucia and also may occur off Puerto Rico (see Remarks on B. communis above).

Remarks: The description of B. litura (Ginsburg 1935) states that B. litura differs from western Atlantic congeners primarily in numbers of second dorsal and anal fin elements. The species is described as having one spine and 11 rays in the second dorsal and anal fins, however, the specimen cataloged as the holotype (USNM 93797) has one spine and 13 rays in both fins. In addition to the differences in dorsal and anal fin elements, there are also discrepancies between the holotype and the description in eye diameter, body depth at the caudal peduncle, maximum body depth, least depth of caudal peduncle, depth and width of head at the posterior margin of the eye and in the length of the caudal fin, which, along with nearly every other fin, was in poor condition in the holotype when we examined it. The number of pectoral rays is also higher in the holotype than in the description (22 vs. 20), although the fins are heavily damaged and it was difficult to count the rays in the dorsalmost and ventralmost parts of the fin. The standard length, head length, predorsal distance and postorbital length were similar in the description and the holotype. Only one specimen of B. litura was known at the time of description; however, a second pair of Bollmannia litura (USNM 116365) was collected during the same expedition in which the holotype was collected, less than 5 miles from the type locality (Samana Bay, Domincan Republic) at similar depths (~20-30 m). These specimens were later identified as B. litura by Ginsburg and were included in notes from one of Ginsburg's unpublished manuscripts, making a total of three B. litura specimens analyzed by Ginsburg. One specimen (38.5 mm SL female) from USNM 116365 has counts of I,11 in both the second dorsal and anal fin and the second specimen (35.9 mm SL female) has counts of I,12 in both fins. Similar discrepancies in morphometics were noted in the two USNM 116365 specimens, as the standard lengths, head lengths, and post orbital lengths measured in this study were similar to those recorded in Ginsburg's notes, but other features measured in this study were smaller than those recorded in Ginsburg's notes.

The differences in eye, depth and width measurements between Ginsburg's observations and ours may be due to specimen shrinkage over time, mainly in width and depth relative to length. But shrinkage cannot account for the discrepancies in counts of anal fin and second dorsal fin elements, the most notable inconsistency between the original description and the holotype. Ginsburg may have miscounted the fin and recorded the incorrect counts in the description as well as in his unpublished records. Alternatively, it is possible that at some point the holotype was switched with another specimen, with the original holotype now being lost. One reason for the former option being unlikely is Ginsburg's reputation of being meticulous and highly repetitive in his meristic counts. As noted by Myers (1976), "Ginsburg excelled in the meticulous measuring and counting of meristic characters ... he demanded of himself the utmost accuracy. I have seen him spend the morning measuring and counting the same small sample two or three times over, and then return after lunch to repeat the process on the same specimens once or twice again".

A second line of evidence supporting the scenario that the type was switched is that the description of B. litura, although inaccurate in its account of what now is catalogued as the holotype, is altogether quite accurate in its description of a morphologically distinctive species of Bollmannia. We have observed the species from multiple localities and note that several of its distinctive morphological characters were described by Ginsburg himself. We consider B. litura a valid species. If the holotype was indeed switched with another specimen, it is unlikely that it was one of the specimens from USNM 116365, as these specimens are female, and the holotype was described as being male. The identification of the specimen currently cataloged as the B. litura holotype remains unclear. Counts and measurements from the cataloged holotype (USNM 93797) and the original description are compared in Table II.

Table II. Comparison of Bollmannia litura holotype and original
description. Measurements other than SL are in % SL.

                      B. litura    Description of B.
                    holotype USNM  litura (Ginsburg
                        93737            1935)

Sex                          male               male

SL                        38.4 mm              39 mm

Eye diamter                   9.1                 12

Postorbital length           15.5                 16

Head length                  30.7                 31

Maximum body depth           18.1                 25
(base of first
dorsal fin)

Least depth of               10.0                 13
caudal peduncle

Head depth behind            16.0                 20
eye

Head width behind            15.4                 17
eye

Predorsal distance           34.0                 36

Caudal fin length          broken                 49

Second dorsal fin            I,13               I,11
elements

Anal fin elements            I,13               I,11

Pectoral fin rays              22                 20


Comparisons: Bollmannia litura is most similar to B. communis but can be distinguished from that species in having a larger eye (typically greater than 10% SL in B. litura vs. less than 10% SL in B. communis) and in having lower modal counts in the second dorsal and anal fins (I,11-12 for B. litura vs. typically I,13 for B. communis). It can be distinguished from B. boqueronensis by lacking tan or brown stripes on the dorsal fins and brown blotches along the lateral midline. Bollmannia litura also typically has yellow or orange (pale in preservation) stripes and ellipses on the dorsal that are lacking in B. boqueronensis, and it has a much darker pelvic fin than B. boqueronensis. Bollmannia litura differs from B. eigenmanni in having three pores on the preoperculum (M', N, O') vs. having two (M', O'), having a dark pelvic fin (vs. a pale pelvic fin), lacking a distinct black stripe on or above the upper jaw (stripe present in B. eigenmanni), having longitudinal papillae row b originating at a vertical through the anterior margin of the eye (originating more posteriorly in B. eigenmanni) and in having a deeper caudal peduncle (10.0-12.0% SL vs. 7.8-10.0% SL).

Etymology: The common name of "citrine goby" is given in reference to the bright yellowish-orange spots on the dorsal and caudal fins of this species, which are reminiscent of the yellow-orange colored gemstone citrine.

Key to genera and species of the Microgobius group of the
Gobiosomatini (Atlantic species only) Below is an artificial
identification key to the Atlantic species of the genera Akko,
Bollmannia, Antilligobius, Microgobius, Palatagobius and Parrella.
The characters used in the generic key do not necessarily apply to
eastern Pacific members of the genus. The key to western Atlantic
Microgobius is modified from Birdsong (1981) and the characters
used in the key to Palatogobius are from Greenfield (2002).

1a)  Second dorsal fin and anal fin           Akko (A. dionea)
     confluent with caudal fin

1b)  Second dorsal fin and anal fin           2
     distinctly separate from caudal fin

2a)  Vomerine teeth present; pelvic fin       Palatogobius
     narrow, lacking a well-developed
     frenum; preopercular canal pores absent

2b)  Vomerine teeth absent; pelvic fin        3
     rounded or oval, with a well-developed
     frenum; preopercular canal pores
     present or absent

3a)  Top of head naked; second dorsal         Microgobius
     typically I,14-17

3b)  Top of head scaled; second dorsal        4
     typically 1,11-13, rarely 1,14

4a)  Preopercular canal pores absent, head    Parrella (P.
     dorsoventrally depressed, pelvic fin     macropteryx)
     round

4b)  Preopercular canal pores present, head   5
     laterally compressed, pelvic fin oval

5a)  Dorsal spines 1-4 greatly elongate,      Antilligobius
     longest spine reaching to or extending   (A. nikkiae)
     beyond posterior margin of second
     dorsal fin when depressed; body with
     distinct broad yellow band along
     lateral midline (white in preserva-
     tion); pelvic frenum with smooth
     posterior margin, frenum lacking
     thickened collagenous supports

5b)  Dorsal spines 1-4 may or may not be      Bollmannia
     elongate, but never extending to
     posterior margin of anal fin when
     depressed; body not pigmented as stated
     above; pelvic frenum with scalloped
     margin, frenum supported with
     collagenous thickenings (Fig. 12)

Key to Atlantic species of Palatogobius (characters from
Greenfield 2002)

1a)  Eye 9.7-11.3% SL; interorbital 1.6-1.9%  P
     SL; snout 3.1-3.9% SL; pectoral rays 21  grandoculus

1b)  Eye 7.4-8.7% SL; interorbital 3.2-4.6%   P.
     SL; snout 5.8-7.8% SL; pectoral rays     paradoxus
     typically 19 (sometimes 18 or 20)

Key to Atlantic species of Microgobi'us (modified from Birdsong 1981)

la.  Three pores in preopercular sensory      2
     canal; sec ond dorsal fin elements
     typically I,17 or more; anal fin
     elements typically I,l8 or more; great
     than 65 lateral scale rows

lb.  Two pores in preopercular sensory        3
     canal; sec ond dorsal fin elements
     typically I,16 or less; anal fin
     elements I,17 or less; fewer than 65
     lateral scale rows

2a.  Soft dorsal-fin elements I,19-20; anal   M. signatus
     fin ele ments I,20 (occasionally I,19);
     lateral scale rows about 77-90; scales
     mostly cycloid; females with pale bar
     edged in black on body above pectoral
     fin

2b.  Soft dorsal fin elements I,17-18;       M.
     anal fin elements I,18 (occasionally    microlepis
     I,19); lateral scale rows 68-78; scales
     mostly ctenoid; no dark markings on
     body in either sex

3a.  A fleshy median crest present on nape;   M. meeki
     a prominent dark spot on body below
     spinous dorsal fin origin; caudal fin
     usually greater than 40% of SL

3b.  Fleshy median crest absent or poorly     4
     devel oped on nape; body with no dark
     spot below spinous dorsal origin or
     with many dark spots; caudal fin
     usually less than 40% of SL

4a.  Scales mostly ctenoid; about 4 enlarged  M. carri
     cani noid teeth in outer row of each
     dentary; interorbital width broad
     (about 4% SL); a broad yellow stripe
     on side with 2 narrow yel low stripes
     above

4b.  Scales mostly cycloid; about 8           5
     enlarged cani noid teeth in outer row
     of each dentary; interorbital width
     narrow (less than 3% SL); no yellow
     stripe on body

5a.  Three pores in lateral cephalic sensory  M. gulosus
     canal; body with numerous dark
     blotches; mouth of males greatly
     enlarged (greater than 15% SL in males
     larger than 25 mm)

5b.  Two pores in lateral cephalic sensory    M.
     canal; body without dark spots; mouth    thalassinus
     of males little enlarged (less than 15%
     SL in males)

Key to the Atlantic species of Bollmannia

la)  Upper jaw with distinct black stripe of  B. eigenmanni
     pig ment on or above upper lip; sensory
     lateralis canal on posterior margin of
     preopercle with 2 pores (M', O'- Fig.
     13C); longitudinal papil lae row b
     originating at vertical through poste
     rior margin of pupil

lb)  Upper jaw dusky or lacking pigment com-  2
     pletely; sensory lateralis canal on
     posterior margin of preopercle with 3
     pores (M', N, O'- Fig. 13A, B, D);
     longitudinal papillae row b originating
     at vertical through anterior margin of
     pupil

2a)  Pelvic fin lightly pigmented to pale;    B.
     dorsal fins with 3-4 light brown to tan  boqueronensis
     longitudinal stripes often continuing
     onto caudal fin; lat eral midline of
     body with 4-5 brown blotches often
     connected with a slightly lighter hori
     zontal stripe of pigment

2b)  Pelvic fin heavily pigmented to black;   3
     dorsal fins and upper half of caudal
     fin with dark ellipses with bright
     orange or yellow centers in life,
     center of ellipses pale in
     preservation; lat eral midline of body
     with 4-5 faint brown blotches that are
     rarely connected by horizontal stripe
     of pigment

3a)  Eye diameter greater than 10% SL;        B. litura
     second dorsal and anal fin elements
     typically 1,11-12 (rarely I,13)

3b)  Eye diameter less than 10% SL; second    B. communis
     dorsal and anal fin elements typically
     I,13 (occasionally I,12 or I,14 in
     either the second dorsal or anal, but
     rarely in both)


Comments on the tribe Gobiosomatini

(Materials examined listed in Appendix I) Birdsong (1975) first proposed the tribe Gobiosomatini (see Smith & Baldwin (1999)) for discussion on spelling of tribe name) as a monophyletic assemblage to include 19 genera of New World gobiids based on the presence of the following characters: (1) vertebral count of 11 precaudal, 16-17 caudal; (2) first dorsal pterygiophore pattern of 3(221110); (3) scapula unossified; and (4) first 2 anal pterygiophores inserted anterior to the first haemal arch. Several of the sponge-dwelling genera do not possess those characters but were included based on previously suggested relationships (Bohlke & Robins 1969; Bohlke 1969) and the presence of hypural fusion in those genera and most genera within the Gobiosomatini - a character rarely found in other genera of Gobiiformes. The tribe was later separated into the Gobiosoma and Microgobius groups (Birdsong et al. 1988). The Gobiosoma group is united based on: (1) fusion of hypurals 1-2 with 3-4 and the terminal vertebral element; (2) dorsal pterygiophore pattern of 3(221110); (3) vertebral counts of 11 precaudal and 16-17 caudal vertebrae; (4) one epural; and (5) two anal pterygiophores anterior to the first haemal arch. The Microgobius group lacks hypural fusion and possesses the remainder of characters uniting the Gobiosoma group. The genera Robinsichthys Birdsong, 1988, Akko Birdsong & Robins, 1995 and Ophiogobius Gill, 1863, were later added to the tribe.

Genera currently in the Gobiosoma group include Aboma, Aruma, Barbulifer, Elacatinus, Chriolepis, Eleotrica, Evermannichthys, Ginsburgellus, Gobiosoma, Gobulus, Gymneleotris, Nes, Ophiogobius, Pariah, Psilotris, Pycnomma, Risor, Robinsichthys, Tigrigobius and Varicus. Aboma, Elacatinus and Tigrigobius were included in Birdsong's genus Gobiosoma based on their subgeneric status as defined by either Bohlke & Robins (1969) for Elacatinus and Tigrigobius or Hoese (1971) for Aboma. The Microgobius group includes Akko, Bollmannia, Microgobius, Palatogobius and Parrella.

The outgroup to the Gobiosomatini is currently unknown and may be composed of more than one species (Ruber et al. 2003). Several eastern Atlantic and Mediterranean genera, namely Odondebuenia, Corcyrogobius, Vanneaugobius and Gorogobius have been proposed as possible outgroups (Miller & Tortonese 1969; Miller 1972, 1978; Van Tassell et al. 1988) along with Egglestonichthys and Callogobius from the Indo-west Pacific (Miller & Wongrat 1979) based primarily on sensory papillae patterns. The presence of modified basicaudal scales in many of the Gobiosomatini along with similar pterygiophore insertion patterns suggest an eastern Atlantic/ Mediterranean origin for the outgroup (Van Tassell et al. 1988; Birdsong 1988; Birdsong et al. 1988).

Molecular studies of the tribe have added additional support for the monophyly and proposed relationships of the tribe to eastern Atlantic and Mediterranean genera (Ruber et al. 2003; Ruber & Van Tassell, unpublished, presented in 2006). They indicate additional subgroups within the Gobiosoma group and areas in need of further investigation. Four subgroups can be recognized: Barbulifer group (Aruma, Barbulifer, Elacatinus, Enypnias, Evermannichthys, Ginsburgellus, Gobiosoma, Risor, Tigrigobius); Nes group (Nes, Psilotris, Gobulus, Gymneleotris, Pycnomma, Chriolepis, Varicus); Aboma; and Robinsichthys. Aboma is sister to the Barbulifer Group and lacks fusion of hypural 1-2 with 3-4, adductor mandibulae 1 gamma and adductor mandibulae 2 gamma, characters present in the Barbulifer group (Van Tassell 1998). Most genera within the Nes group examined to date (Nes, Psilotris, Gobulus, Gymneleotris, Pycnomma, Chriolepis, Varicus) have pelvic fins that are separate to at least some extent (never with complete disks), lack adductor mandibulae 1, possess no head pores or canals, but have hypural fusion. Aboma and Robinsichthys lack hypural fusion, but this was unknown when the Gobiosoma Group was first proposed and their relationship to the remaining genera requires further investigation. These subgroups are used to facilitate comparisons among genera in the Microgobius group below.

Comments on the Microgobius group

The Microgobius group proposed by Birdsong et al. (1988) consisted of Microgobius, Bollmannia, Parrella and Palatogobius. This group was phenetically associated with the Gobiosoma group on the basis of a dorsal pterygiophore pattern of 3(221110), vertebral count of 11+16, one epural and two anal pterygiophores anterior to the first haemal arch. The Microgobius group was distinguished from the Gobiosoma group by lacking fusion of hypural 1-2 with 3-4 and the terminal caudal element (Birdsong et al. 1988). Birdsong & Robins (1995) added Akko to the Gobiosomatini, but did not comment further on relationships due to the large number of autapomorphies it possesses. Additional studies of mtDNA (Ruber et al. 2003) and morphological characters (Van Tassell & Baldwin 2004) placed Akko in the Microgobius group. Additional characters that unite the group include dorsal papillae row n elongate, uniting across dorsum or nearly so, a feature present in all genera except Parrella; caudal fin lanceolate; body laterally compressed; metapterygoid process not overlapping the quadrate; and modified basicaudal scales absent.

We place Antilligobius nikkiae within the Microgobius group of the Gobiosomatini based on the presence of the following characters: vertebrae 11 +16; dorsal pterygiophore formula 3(221110); anal pterygiophores anterior to the first haemal arch 2; hypurals 1-2 and 3-4 not fused; dorsal papillae row n elongate, uniting across dorsum; caudal fin lanceolate; body laterally compressed; metapterygoid process not overlapping the quadrate; and modified basicaudal scales absent. A total of 57 morphological characters, including papillae and head pore patterns (Fig. 18), are summarized for the 6 genera of the Microgobius group in Table III.

Table III. Comparison of morphological characters across the
Microgoblus-group genera

                      Antilligobius        Bollmannia       Parrella

Type species        Antilligobius     Bollmannia        Parrella
                    nikkiae           chlamydes         maxillaris
                    Van Tassell &     Jordan, 1890      Ginsburg,
                    Colin                               1938

Precaudal           11                11                11
vertebrae

Caudal vertebrae    16                16                16

Hypurals 1-2 fused  No                No                No
to 3-4 and to
urostyle

DI                  7                 7                 7

DI pterygiophores   3(221110)         3(221110)         3(221110)

DI elongate         Yes, both sexes   Yes, both sexes   Yes, females
                                      of some species   of some
                                                        species

DII                 I,11-12           I,11-15           I,11-12

DII position        Post 9th neural   Post 9th neural   Post 9th
                    spine             spine             neural spine

A                   I,11-13           I,10-15           I,10-12

Anal fin position   First two         First two         First two
                    elements          elements          elements
                    anterior to       anterior to       anterior to
                    haemal spine on   haemal spine on   haemal spine
                    12th vertebrae    12th vertebrae    on 12th
                                                        vertebrae

C segmented         17                16-18 (Atlantic   17
                                      species only)

C branched          15                14-16 (Atlantic   16
                                      species only)

C shape             Lanceolate        Lanceolate        Lanceolate

P                   20                19-26             19-20

V                   I,5               I,5               I,5

V shape             Ovate             Ovate             Rounded

Pelvic frenum       Well developed,   Well developed,   Well
                    smooth margin     scalloped         developed,
                                      margin            scalloped
                                                        margin

Cephalic            B' C(s) D(s) E F  B' C(s) D(s) E F  B' D(s) F,
lateral-line canal  H' (sometimes     G H' (or H, J')   H', or B'
pores               G)                K' L' (K'L'       D(s) F G H'
                                      absent in B.      or B' C(s) G
                                      eigenmanni)       H', sometimes
                                                        K' L'

Opercular canal     M' N O' Mainly    M' N O' or M' O'  Absent or
pores Scale type    ctenoid, some     Mainly ctenoid,   M',N' Ctenoid
                    cycloid           some cycloid      with reduced
                                                        ctenii

Predorsal scales    6                 6-10              18

Scales cheek        2-3 scales        Few large         Absent
                                      scales

Scales opercular    Absent            Present           Absent

Scales breast       Present           Present           Present

Scales belly        Present           Present           Present

Scales pectoral     Present           Present           Present
fin base

Scales transverse   6-8               6-8               8
scale rows

Lateral scale       21-28             24-29             32
rows

Modified            Absent            Absent            Absent
basicaudal scales

Vomerine teeth      Absent            Absent            Absent

Angle of mouth      30 deg            30 deg            35 deg

Extent of gill      Upper pectoral    Upper pectoral    Upper
openings            fin base to       fin base to       pectoral fin
                    ventral surface   ventral surface   base to
                    of body           of body           ventral
                                                        surface of
                                                        body

Gill rakers shape   Short slender     Moderately long,  Short stout
                                      slender

Gill rakers         5+12              3-4+9-12          4+8
upper+lower

Posterior           Raised rim        Slight raised     No rim
nostrils                              rim

Anterior nostrils   Short tube        Short tube        Short tube

Fleshy nape crest   Absent            Absent            Absent

Male papillae       Conical           Conical           Conical

Female papillae     Bulbous           Bulbous           Bulbous

Color sexually      No                Only B.           No
dimorphic                             communis

Head sexually       No                No                Mouth and eye
dimorphic                                               larger in
                                                        males

Gas bladder         Well developed    Well developed    Absent

Bony support of     Canal supported   Portions of       Canal

supraorbital        by open trough    canal enclosed    supported by
sensory canal                         in bony tube      open trough

Sagittal crest on   Present           Present           Absent
frontal
bone

Basihyal shape      Spatulate         Spatulate         Bifid

Process on          Absent            Absent            Absent
metapterygoid
overlapping
quadrate

Metapterygoid       Very narrow,      Broad, 2-3 times  Very broad,
shape               splinter like     width of          at least 3
                                      symplectic        times width
                                      (male)            of
                                                        symplectic

Symplectic process  Absent            Slightly          Absent
on preopercle                         developed to
                                      absent

Lateral shelf on    Present (1        Present (2)       Present (1)
posttemporal        reduced)
(number of
shelfs)

Sphenotic length    Moderate          Short             Short

Ventral             Present           Present           Absent
postcleithrum

Shape of braincase  Circular          Circular          Circular
in dorsal view

Sensory papillae    Short, not        Medium to long,   Very long
row b               forward of last   extending to      extending
                    transverse row    anterior of eye   almost to 1st
                    but B.            in all but B.     row
                    eigenmanni        eigenmanni

Sensory papillae    Long, continuous  Long, continuous  Short, not
row n               across dorsal     across dorsal     continous
                    midline           midline or        across dorsal
                                      nearly so         midline

Sensory papillae    Not continuous    Almost            Almost
row x (1)
                            continuous        continuous
Sensory papillae    Continuous        Continuous        Continuous
row d

Sensory papillae    None              None              None
transverse rows
extending below d

                     Microgobius       Akko        Palatogobius

Type species        Microgobius    Akko dionaea   Palatogobius
                    signatus                      paradoxus
                    Poey, 1876     Birdsong &     Gilbert, 1971
                                   Robins, 1995

Precaudal           11             11             11
vertebrae

Caudal vertebrae    16-17          16-17          16
Hypurals 1-2 fused  No             No             No
to 3-4 and to
urostyle

DI                  7              7              7

DI pterygiophores   3(221110)      3(221110)      3(221110)

DI elongate         Yes, males     No             No

DII                 I,14-19        I,14-15        I,17-20

DII position        Post 9th       Post 9th       Post 9th
                    neural spine   neural spine   neural spine

A                   I,14-20        I,14-15        I,19-21

Anal fin position   First two      First two      First two
                    elements       elements       elements
                    anterior to    anterior to    anterior to
                    haemal spine   haemal spine   haemal spine
                    on 12th        on 12th        on 12th
                    vertebrae      vertebrae      vertebrae

C segmented         17             16-17          17

C branched          15             15-16          14

C shape             Lanceolate     Lanceolate     Lanceolate

P                   20-23          17-21          18-21

V                   I,5            I,5            I,5

V shape             Ovate          Ovate          Ovate

Pelvic frenum       Well           Well           Absent
                    developed,     developed,
                    smooth         smooth
                    margin         margin

Cephalic            B' D F G H'    B' G'          B' C(s) D(s) E
lateral-line canal                                F'
pores

Opercular canal     M' N O' or M'  Absent         Absent
pores Scale type    N' Cycloid +   Cycloid        Cycloid
                    ctenoid

Predorsal scales    0              Some           0

Scales cheek        Absent         Absent         Absent

Scales opercular    Absent         Absent         Absent

Scales breast       Absent         Absent         Absent

Scales belly        Absent         Absent         Absent

Scales pectoral     Absent         Absent         Absent
fin base

Scales transverse   20             19-20          None
scale rows

Lateral scale       35-90          53-115         12-17
rows

Modified            Absent         Absent         Absent
basicaudal scales

Vomerine teeth      Absent         Absent         Present

Angle of mouth      Highly         50 deg         35 deg
                    variable
                    between
                    species

Extent of gill      and sex        Base of 4th    Upper margin
openings            Length of      pectoral to    of orbit to
                    pectoral       ray to pelvic  just below
                    base           fin base       lower margin
                                                  of orbit

Gill rakers shape   Long slender   Papilloid      Moderately
                                                  long, slender

Gill rakers         3-5+10-18      6+6            4-5+12-13
upper+lower

Posterior           No rim         No rim         Raised rim
nostrils

Anterior nostrils   Short tube     Short tube     Short tube

Fleshy nape crest   Present -      Absent         Absent
                    some species

Male papillae       Conical        Conical        Short, bluntly
                                                  pointed

Female papillae     Bulbous        Short,         Short blunt
                                   rounded        conical

Color sexually      Some species   No             No
dimorphic

Head sexually       Some species   No             No
dimorphic

Gas bladder         Well           Small -        Absent
                    developed      thickened

Bony support of     Canal          Canal          Canal
supraorbital        supported by   supported by   supported by
sensory canal       open trough    open trough    open trough

Sagittal crest on   Present        Absent         Absent
frontal
bone

Basihyal shape      Bifid          Minute,        Narrow
                                   reduced

Process on          Absent         Absent         Absent
metapterygoid
overlapping
quadrate

Metapterygoid       Slightly       Very broad,    Very narrow
shape               broadened,     at least 3
                    less than 2    times width
                    times width    of
                    of symplectic  symplectic
                    (sexually
                    dimorphic)

Symplectic process  Slightly       Absent         Present
on preopercle       developed to
                    absent

Lateral shelf on    Absent         Present (1)    Absent
posttemporal
(number of
shelfs)

Sphenotic length    Short          Long,          Short
                                   extending to
                                   vertical
                                   through
                                   posterior
                                   margin of
                                   jaw

Ventral             Absent         Absent         Present
postcleithrum

Shape of braincase  Circular       Elongate       Circular
in dorsal view

Sensory papillae    Short,         Very long      Long extending
row b               anterior of    extending      forward to
                    pupil to       almost to      under middle
                    posterior of   posterior of   of eye
                    eye or         pupil
                    longer

Sensory papillae    Long,          Long,          Long, nearly
row n               continuous     continuous     continuous
                    across dorsal  across dorsal  across dorsal
                    midline        midline        midline

Sensory papillae    Continuous     Continuous     Almost
row x (1)
                                      continuous
Sensory papillae    Continuous     Continuous     Continuous
row d

Sensory papillae    None           Last           None
transverse rows                    transverse
extending below d                  row


DISCUSSION

Antilligobius nikkiae has a number of characters not present in other genera of the Microgobius group that warrant its placement in a new genus. These include a very narrow, splinter-like metapterygoid; short slender gill rakers on the lower limb of the first arch; a short papillae row b, never extending posterior of the pupil, and the modified first haemal arch. The modified haemal arch and its elongate parapophyses, "rib-like" in appearance in lateral view, where parapophyses join ventrally to form the haemal arch, is easily confused with pleural ribs in radiographs.

The modified first haemal arch and anterior curvature of the ventral tip of the premaxilla warrant further investigation. A similar modified haemal arch occurs in several species of Trimma, the expanded area being utilized by the swim bladder (Winterbottom 1984, 2007). The swim bladder of A. nikkiae also occupies this expanded area in the specimen examined. The anterior curvature of the ventral tip of the premaxilla occurs in Antilligobius and several species of Microgobius. Birdsong (1975) observed sexually dimorphic differences in the curvature of the ventral tip of the premaxilla, with the tip curving anteriorly in females of several species of Microgobius. Differences were also observed between small and large-mouthed species of Microgobius, regardless of sex (Birdsong 1975). The significance and distribution of these characters within the Gobiosomatini and Gobiiformes awaits further studies.

Within the Microgobius group Antilligobius is most similar morphologically to Bollmannia and Parrella. Antilligobius shares with those genera I,11-15 elements in the second dorsal fin (I,14-20 in Microgobius, Akko, Palatogobius); I,10-15 elements in the anal fin (I,14-21 in Microgobius, Akko, Palatogobius); mostly ctenoid scales on the trunk (few ctenoid scales present in Microgobius, and cycloid scales present in Palatogobius and Akko); scales present on the breast, belly and pectoral fin base (no scales in Microgobius, Akko, Palatogobius); 21-29 scales in the lateral series (35-115 in Microgobius and Akko, 12-17 in Palatogobius); and 6-8 transverse scale rows (19-20 in Microgobius, Akko and none in Palatogobius). Antilligobius and Bollmannia both possess scales on the cheek and a spatulate basihyal, features not present in the other genera. In Parrella sensory papillae row n is not continuous across the nape, whereas it is continuous or almost so in the other genera. The foramen on each anterior vertebrae of Antilligobius (Fig. 8C) is much larger than that in B. litura, B. communis and B. boqueronensis, but is similar in size to that of B. eigenmanni. Parrella maxillaris vertebrae are unlike those of Antilligobius and Bollmannia in that there are numerous small foramina on the anterior vertebrae. Antilligobius also lacks the collagenous supports on the pelvic frenum that are present in Bollmannia (Fig. 12), which may be homologous with the extensive fimbriae found on the posterior margin of the pelvic frenum of Parrella. Antilligobius also differs from Parrella and Bollmannia ecologically, as it inhabits rock-reef talus slopes while both Bollmannia and Parrella are burrow dwellers in soft mud-sand substrate.

Elongate first dorsal filaments are found in nearly all species of Bollmannia, variable within Microgobius and Parrella and absent in Palatogobius. In Antilligobius the first four dorsal spines are all very elongate, giving the fin a sabre-like appearance. Microgobius, Bollmannia and Antilligobius all possess 2-3 pores in the preopercular canal (pores absent in Parrella maxillaris, Akko, Palatogobius) and have a sagittal crest on the frontal bone (absent in Parrella, Akko, Palatogobius). The sagittal crest was considered a specialized character within the Microgobius group by Birdsong (1975). Antilligobius shares the presence of a ventral postcleithrum with Bollmannia and Palatogobius. The very narrow metapterygoid in Antilligobius, somewhat similar but wider in Palatogobius, is broad in the remaining genera.

Molecular phylogenies by Ruber et al. (2003) and Ruber & Van Tassell (unpublished, presented in 2006) suggest a sister relationship between Antilligobius and Bollmannia. Ruber et al. (2003) recovered a strongly supported clade comprising Antilligobius + a monophyletic group of four Bollmannia species. That clade is sister to a clade containing eight species of Microgobius and Akko. Parrella and Palatogobius were not included in their study. With additional species of Microgobius and Parrella lucretia incorporated in the tree, the phylogeny reconstructed by Ruber & Van Tassell (unpublished, presented in 2006) showed an identical relationship between Bollmannia and Antilligobius (labeled as New Genus in the tree), but Parella lucretia and Akko now formed a distinct clade sister to Microgobius.

While several key morphological characters support a close relationship between Antilligobius, Bollmannia and Parrella, the existence of other characters shared between Antilligobius and other Microgobius group genera cloud this relationship. The large number of autapomorphic specializiations and losses within the group coupled with the absence of DNA sequence data for many species in the Microgobius group adds to the ambiguity regarding the relationships among species. A robust phylogenetic analysis of all Microgobius group species using morphological and molecular data will help clarify the relationships between Antilligobius and allied species.

Appendix I

Gobiosomatini material examined:

Akko dionaea - USNM 32524, holotype, 1, male, 78.1 mm SL, east of Isla de Maraca, Mapa Province, Brazil, 25 May 1976, Oregon II.

Akko brevis - EL-01-017 (JVT collection), 2 cleared and stained, 20.0 m, shrimp trawler, 13.159[degrees]N, 087.805[degrees]W to 13.140333[degrees]N, 087.8[degrees]W, El Salvador, J.L. Van Tassell.

Aruma histrio - CAS 79602, 1, Guaymas, 2-3 miles north of Bahia San Carlos, Mexico, 25 May 1954, C. Limbaugh.

Barbulifer pantherinus - SU 49929, 52, Ensenada San Francisco, flat reef 0.3 miles north-west of - tip of Punta de las Cuevas, Puerto San Carlos, Mexico, 6 April 1955, Dickinson et al.

Barbulifer ceuthoecus - ANSP 110660, 7, shore at International Airport, Key West, Florida, 21 March 1958, Cohen et al.

Bollmannia chlamydes - CAS 42777, 11, off Ballenas Bay, Gulf of Nicoya, 9.686111[degrees]N, 84.850833[degrees]W, Costa Rica, 25 February 1938, New York Zoological Society Eastern Pacific Expedition.

Chriolepis minutillus - LACM 20148, 4, Gulf of California, Angel de la Guarda Island, Puerto Refugio, Mexico, 27 January 1940, R/V Velero; UF 72110, 3 plus 1 cleared and stained, Gulf of Mexico, Florida, 3 October 1970, R/V Tursiops.

Chriolepis vespa - UF 72110, 1 cleared and stained, Gulf of Mexico, Florida, 3 October 1970, R/V Tursiops.

Elacatinus horsti - UF 26237, 4, Isla de Providencia, Colombia, 29 August 1971, Gilbert and Anderson. Elacatinus oceanops - UF 16169, 23, Hawk Channel, Florida Keys, Florida, 6 August 1967, Gilbert and Starck. Elacatinus xanthiprora - UF 24797, 2, Isla de Providencia, Colombia, western Atlantic, 30 August 1971, P. Colin.

Eleotrica cableae - USNM 278977, 47, Indefatigable Island, Galapagos Islands, 0.7647[degrees]S, 90.3414[degrees]W, 18 May 1966, H. A. Fehlmann.

Enypnias seminudus - USNM 123641, 2 cleared and stained, Miraflores Locks, Panama Canal, Panama, 28 April 1937, S.F. Hildebrand; FMNH 8457, 1, tidepools, Panama, 24 March 1912, Meek and Hildebrand.

Evermannichthys spongicola - AMNH 82393, 3 cleared and stained, Gulf of Mexico, 25 July 1975, S. Bortone.

Ginsburgellus novemlineatus - UF 201602, 8, Bethel Channel, Andros Islands, Bahama Islands, June 1951, E.C. Jones.

Gobiosoma chiquita - SU 66911, 1 cleared and stained, 1.5 m, Outer San Carlos Bay, shore of island, Gulf of California, Mexico, 2 June 1950, Wilimovsky et al.

Gobiosoma homochroma - USNM 293555, 9, Rio Pirre above tidal zone, Darien Prov., Panama, 17 February 1985, B. Chernoff et al.

Gobiosoma longipala - GCRL 815, 26, station 896, Gulf of Mexico, Mississippi, 3 November 1960.

Gobiosoma nudum - GCRL V73:11523, 11, station 1607, Venado Beach, Pacific Panama, 4 April 1973, Dawson et al.

Gobulus hancocki - SIO61-265, 4, 18.5-25.0 mm SL, Espiritu Santo Islands, El Cardonal Cove, 24.61[degrees]N, 110.38833[degrees]W, Mexico, 23 June 1961, R/V Orca, B. Walker et al.

Gymneleotris seminudus -MCZ 44759, 2, Rey Island, Perlas Islands, Panama, 27 September 1964, I. Rubinoff.

Microgobius miraflorensis -FMNH 8489, 2, Rio Culebra, Culebra, Panama, 19 March 1911, Meek and Hildebrand.

Microgobius signatus - USNM 35164, 1 of 12, Havana, Cuba; MCZ 30602, 3, Havana, Cuba, 1913, T. Barbour.

Nes longus - ANSP 133224, 2 cleared and stained, Castle Harbor, Bermuda, 10 August 1975, W. F. Smith-Vaniz.

Ophiogobius jenynsi - USNM 176505, 1, Ancud, Golo de Quatalmaue, Pullingua, Chile, 7 February 1945, M. J. Lobell.

Palatogobius grandoculus - USNM 367311, holotype, 1, male, 33.0 mm SL, off Cozumel, 20.4475[degrees]N, 87.2553[degrees]W, Mexico, 11 April 1976, Marine Biomedical Institute.

Palatogobius paradoxus [degrees] ANSP 109182, holotype, female, 25.5 mm SL, St. John, Virgin Islands, 30 June 1961, J.E. Randall; UF 163509, 2, Gulf of Mexico, 29.833333[degrees]N, 86.108333[degrees]W, Florida, 20 July 1975, SA Bortone and Nester; UF 148027, 7, Gulf of Mexico, 29.933[degrees]N, 86.1[degrees]W, USA, 3 June 1974, R/V Mafla.

Pariah scotius [degrees] AMNH 26096, 3, Conception Island, Bahamas, 6 March 1966, C.L. Smith.

Parrella macropteryx [degrees] UF 167272, cleared and stained, PIL751 Field number, Caribbean Sea, Venezuela, 26 July 1968; UF 223879, 1, 36 m, Caribbean Sea, 9.52[degrees]N, 75.685[degrees]W, Colombia, 13 July 1966, R/V Pilsbury.

Parrella maxillaris [degrees] USNM 119901, Paratype, 1, La Paz Bay, Gulf of California, Mexico, 21 February 1936, Hancock Expedition; USNM 322735, 82, Punta San Ignacio, Sonora, 25[degrees] 35' N, 110[degrees], 25'W, Mexico, 19 June 1990, G. Allen and D. Robertson; UF 167275, 22.9mm SL, 1 female, cleared and stained, 22-25 m, 7 miles north of Manta, bearing 270 deg, along 20 fm curve, 0.716667[degrees]S, 80.683333[degrees]W, Ecuador, 1 October 1961, R/V Argosy.

Robinsichthys arrowsmithensis [degrees] UF 234392, paratype, 1 female, cleared and stained, 23.5 mm SL, 242-320 m, Quintana Roo, station 893, 21.166667[degrees]N 86.35[degrees]N, Mexico, 10 Sept 1967, R/V Gerda.

Tigrigobius dilepis [degrees] UF 14032, 2, Exuma Sound, Exuma Cays, Bahama Islands, 29 August 1966, C.R. Gilbert. Tigrigobius gemmatum [degrees] AMNH 26076, 6, cleared and stained, Hog Cay, Ragged Islands, Bahamas, 9 July 1965, C. L. Smith & H. Tischler.

Tigrigobius janssi [degrees] LACM 32524-45, 1, Bahia Herradura, at tip of outer reef on N side of bay, Costa Rica, 9 March 1972, Bussing et al.

Tigrigobius macrodon [degrees] ANSP 147847, 36, 11.9-21.5 mm, Baileys Bay, Bermuda, 23 July 1981, W. F. Smith-Vaniz et al.

ACKNOWLEDGEMENTS

Thanks to D. Ross Robertson, Ben Victor, and Ernesto A. Pena E. along with the crew of the RV Urraca (Smithsonian Tropical Research Institute) for their aid in obtaining the Bollmannia litura specimens. Frank Pezold provided valuable comments throughout this study. Funding for collections in Panama was provided by Smithsonian Tropical Research Institute to D. Ross Robertson. B. Brown and R. Arrindell at AMNH, M. S. Perez at ANSP, S. LeCroy at GCRL, K. Conway and H. Prestridge at Texas A&M University - College Station, R. Robins at UF, M. Craig at UPRM, J. Williams, D. Pitassy and S. Raredon at USNM, and D. Skelly at YPM provided assistance with museum specimens. G. D. Johnson provided assistance with digital photography of cleared and stained specimen and helpful discussions on osteology. D. Smith provided assistance with unpublished Ginsburg material at USNM. C. Baldwin provided photographs of Antilligobius habitat. F. Young provided habitat information for Antilligobius specimens in Curacao. A. Schrier provided critical comments, photographs and specimens of Antilligobius. R. G. Gilmore provided the specimen from Cuba, illustrations and detailed records of Antilligobius from Cuba and other localities.

Received: 20 May 2011 - Accepted: 30 September 2011

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James L. Van Tassell (1)*, Luke Tornabene (2), Patrick L. Colin (3)

(1.) American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A.

* Corresponding author: jvantassell@gobiidae.com

(2.) Texas A&M University - Corpus Christi, 6300 Ocean Drive, Corpus Christi, TX 78412, U.S.A.

(3.) Coral Reef Research Foundation P.O. Box 1765 Koror, Palau 96940
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Author:Van Tassell, James L.; Tornabene, Luke; Colin, Patrick L.
Publication:aqua: International Journal of Ichthyology
Article Type:Report
Geographic Code:0ATLA
Date:Apr 15, 2012
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