Comprehensive Description
provided by Smithsonian Contributions to Zoology
Nannosalarias nativitatus (Regan)
Blennius nativitatus Regan, 1909, pp. 404–405, pl. LXVI, fig. 2 [original description; type-locality: Christmas Island].
MATERIAL EXAMINED.—BNNH 1909.3.4.42 (adult male, 30.1 mm SL), lectotype of Blennius nativitatus, Christmas Island, lat. 10°30′S, long. 105°40′E, 1909, C. W. Andrews; BMNH 1909.3.4.43-52 (5 of 15 paratopotypes, examined, 25.2–33.7); USNM 157372 (2 paratopotypes, 22.2–33.0); USNM 203098 (10, 24.9–33.0); Australia, Queensland, Capricorn group, One Tree Island, south reef face about 1.6 km from island, lat. 23°30′S, long. 153°08′E, depth 0–4.6 meters, 22 November 1966, Victor G. Springer, VGS 66–7; AMS IB 8207 (5, 26.6–30.4), same data as USNM 203098; USNM 201369 (9, including 1 cleared and stained, 22.5–34.8), One Tree Island, off reef crest on west side of island just above drop off, depth 7.1–12.2 meters, 7 December 1966, Victor G. Springer, VGS 66–16; AMS uncat. (1, 22.7), sta. FT 96, Australia, One Tree Island, south face, depth 10 meters, 26 November 1966, F. Talbot, et al.; AMS uncat. (4, 26.3–34.3), sta. FT 204, Australia, One Tree Island, reef slope on south face, depth 8 meters, 22 September 1967, F. Talbot and B. Goldman; ANSP 109688 (2, 23.5–26.3), Australia, Queensland, Endeavour Reef, depth 1.3–2.4 meters, 13 January 1969, J. C. Tyler and C. L. Smith, TS.A–13; ANSP 109687 (8, 15.7–27.7), Australia, Queensland, Endeavour Reef, depth 1.5–4.6 meters, 16 January 1969, J. C. Tyler and C. L. Smith, TS,A-17; USNM 195700 (1, 25.5), presumably Dutch East Indies; CAS 24156 (5, 25.4–30.1), same data as USNM 201369; USNM 203097 (2, 28.5–32.3), original no. GVF 1737, sta. HK 60, South China Sea, Pratas Reef, about 0.6 km west by north of Southwest Horn Island, lat. 20°36′40"N, long. 116°43′50"E, 21 May 1958, Rolph L. Bolin; USNM 203767 (2, 19.3–23.8), Te Vega Cruise 7, sta. 301, Tonga Islands, Nivatobutabu Island, lat. 16°00′S, long. 175°53′W, depth 0–7 meters, 31 May 1965, Rolph L. Bolin.
DESCRIPTION.—(Characters for holotype in parentheses.) Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Canine on each dentary posteriorly. Premaxillary and dentary teeth movable, but fairly rigid; premaxillary teeth 76 to 80 (80); dentary teeth 42 to 52 (42). Vomer with 5 or 6 small, conical teeth. Terminal vertebra with two epurals and autogenous hypural 5 and ventral hypural plate. Vertebrae 10 + 24 or 25 = 34 or 35; epipleural ribs 14 to 16; last pleural rib on vertebra 10. Circumorbital bones 4.
Dorsal spines 12 (rarely 13); last spine noticeably reduced; segmented rays 15 to 16(16) [18 in one specimen]; basal two-thirds of terminal dorsal ray bound by membrane to caudal peduncle; dorsal fin moderately incised between spinous and rayed portions. Anal spines 2; anal fin rays 16 to 18 (17); basal two-thirds of terminal anal ray bound by membrane to caudal peduncle. Caudal fin with 13 segmented rays, middle 9 each branched once; dorsal procurrent rays 6 to 8 (7), ventral rays 5 to 7 (6). Pelvic rays I, 3. Pectoral rays 15, rarely 14. Pectoral radial formula 2–1–1 (based on 1 cleared and stained specimen). Gill-rakers 12 to 14. Pseudobranchial filaments 5. Nuchal, supraorbital, and nasal cirri short and simple. Upper and lower lips entire. Some pores of infraorbital series in pairs. Lateral line of simple pores, incomplete, continuous to point below 7th or 8th dorsal spine.
PIGMENTATION.—The following description is a composite based on Australian specimens. Background color of head and body pale. Sides with concentrations of melanophores tending to form bands. Melanophores becoming more numerous on dorsum and head, especially beneath eye and at corners of mouth. Throat with two large, dark gular spots extending posteriorly from lower lip. (In the Tonga specimens the gular spots have become united as a single large spot.) Gular spots well defined and intensified in some males, coalesced and diffuse in others, and even less well defined in females. Spinous dorsal fin of some specimens heavily pigmented on basal one-fifth, otherwise unmarked. Soft dorsal with a heavy concentration of pigment on basal one-fifth and on distal margin of fin, rays otherwise largely unmarked. Anal fin with interradial membranes heavily pigmented, especially distally. Caudal fin rays faintly outlined with melanophores; interradial membranes of unbranched rays heavily pigmented in some males. Paired fins unmarked. Females are usually paler than males, and in some specimens the fins are completely devoid of pigment.
Regan (1909, p. 405) gives the life colors as follows: “color reddish, with about 8 darker cross-bars, sometimes appearing only as a series of oblong blotches along the middle of the sides; fins pale.”
GEOGRAPHIC VARIATION.—The two South China Sea specimens of Nannosalarias differ in several respects from the Australian and Tongan material, especially in pigmentation. In the male the leading edge of the first dorsal spine is intensely pigmented, and the basal half of the spinous dorsal fin has a heavy concentration of melanophores. The head is uniformly dark and the two gular spots (not apparent in the photograph) are smaller, located farther posteriad, and more widely separated. The female differs in having well-defined concentrations of pigment in approximately the position of several of the preoperculomandibular pores. The gular spots, although not as well defined, are in the same position as those of the male.
The two South China Sea specimens also differ from the other material in having longer supraorbital cirri. In the male (28.5) the ratio of the cirri length into the orbital diameter is 1.0; in 8 Australian males (22.5–34.8) this ratio is 1.6 to 2.7, average 2.0. In the female (32.3) the cirri/orbital ratio is 1.8; in 6 Australian females (25.5–29.5) it is 2.0 to 5.3, average 3.0.
We do not believe that the differences exhibited by the two South China Sea specimens warrant description as a taxon separate from that of the Australian and Tongan specimens. Springer (1967) did recognize a separate subspecies for the South China Sea populations of Entomacrodus thalassinus, based mainly on relative length of the supraorbital cirrus, and the South China Sea population of E. stellifer, based primarily on color pattern. In both instances, however, considerable material was available.
Ophioblennius Gill
Blennophis Valenciennes in Webb and Berthelot, 1843, p. 60 [type-species: B. webbi Valenciennes in Webb and Berthelot, 1843, = Salarias atlanticus Valenciennes in Cuvier and Valenciennes, 1836, by original designation; a junior homonym of Blennophis Swainson, 1839, Clinidae].
Ophioblennius Gill, 1860, p. 103 [a substitute name for Blennophis Valenciennes in Webb and Berthelot, 1843, taking the same type-species: Blennophis webbi Valenciennes in Webb and Berthelot, 1843].
Labroblennius Borodin, 1928, p. 31 [type-species: L. nicholsi Borodin, 1928, = Ophioblennius steindachneri Jordan and Everman, 1896, by monotypy].
Cynoscartes Norman, 1943, p. 31 [type-species: Salarias atlanticus Valenciennes in Cuvier and Valenciennes, 1836, by original designation].
Hepatoscartes Fowler, 1944, p. 230 [type-species: H. unbri-fasciatus Fowler, 1944, = Ophioblennius steindachneri Jordan and Evermann, 1896, by original designation].
DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering 184 to 224 in upper jaw and 102 to 108 in lower jaw. Vomer toothless. Dorsal rays XI to XIII, 19 to 24; anal rays II, 20 to 25; segmented caudal rays 13, middle 9 branched; pectoral rays 14 or 15 (usually 15); pelvic rays I, 4. Terminal anal ray not bound by membrane to caudal peduncle. Lateral line consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculomandibular pores without cirri. Middorsal supratemporal pores 5 to more than 50. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri consisting of a pair of cirri or a patch of 3 to 15 or more cirri on each side of nape; supraorbital cirri simple; nasal cirri multifid. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
The genus Ophioblennius was revised by Springer (1962). Ophioblennius appears to be most closely related to Scartichthys. In addition to the characters given in the key to distinguish these two genera, Ophioblennius differs in having dentary canines that are equal to or greater than the diameter of the pupil, instead of having canines less than one-half the pupil diameter (exclusive of larvae), and the lateral line consisting of two overlapping disconnected portions instead of being continuous.
DISTRIBUTION—Restricted to the Atlantic Ocean and eastern Pacific Ocean.
Pereulixia Smith
Pereulixia Smith, 1959, p. 238 [type-species: Salarias kosiensis Regan, 1908, by original designation].
DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines absent; posterior dentary canines present. Premaxillary teeth freely movable, numbering 155 to 157; dentary teeth fairly rigid, numbering 43 to 53. Vomer with small conical teeth. Dorsal rays XI or XII, 11 or 12; anal rays II, 12 to 14; segmented caudal rays 13, middle 9 branched; pectoral rays 15 or 16 (usually 15); pelvic rays I, 4. Terminal anal ray not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; imbricate scalelike flaps covering at least anterior lateral-line pores. Preoperculomandibular pores without cirri. Middorsal supratemporal pores numerous, relatively small and difficult to count. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri consisting of a series of 4 elongate, transverse, contiguous or slightly overlapping patches of cirri interrupted at midline by a narrow hiatus; supraorbital cirri simple, long, and tapering; nasal cirri multifid. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
Pereulixia is apparently monotypic and rare in museum collections; we know of fewer than 10 specimens.
DISTRIBUTION.—Known only from the western Indian Ocean.
Praealticus Schultz and Chapman
Praealticus Schultz and Chapman, 1960, p. 368 [type-species: Salarias natalis Regan, 1909, by original designation].
DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jaw bone. Anterior dentary canines absent; posterior dentary canines present or absent. Premaxillary and dentary teeth freely movable, numbering 190 to 280 in upper jaw and 128 to 214 in lower jaw. Vomer with or without teeth. Dorsal rays XII to XIV, 18 to 20; anal rays II, 18 to 23; segmented caudal rays 12 or 13, branched rays 0 to 9 (usually 4 to 8); pectoral rays 14 or 15; pelvic rays I, 3 or I, 4. Terminal anal ray not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculomandibular pores without cirri. A single middorsal supratemporal pore. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest present. Nuchal cirri absent; supraorbital cirri pinnately branched; nasal cirri simple, occasionally with weak points. Postcleithra consisting of 1–3 small, fragile, well-separated bones on each side (except P. schmidti in which postcleithra consist of two elongate bones, head of ventral element overlapping ventral end of dorsal element). Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
The fact that this genus, which is widely distributed throughout much of the Indo-Pacific, was not recognized until 1960 attests to its remarkable resemblance to two other (though not necessarily closely related) genera. Praealticus is well named, for it has frequently been confused with species of Alticus. The close superficial similarity of the two genera is well illustrated by the drawing of the holotype of Alticus semicrenatus Chapman, which was published in the same work as the original description of Praealticus and was not recognized by Schultz and Chapman as being a species of their new genus.
Although we consider Praealticus more closely related to Alticus than to Istiblennius, species of Istiblennius are most often confused with Praealticus and are the most difficult to distinguish from it. A comparison of the characters most useful in distinguishing these three genera is given in Table 1.
DISTRIBUTION.—Eastern Indian Ocean; central and western Pacific Ocean.
Rhabdoblennius Whitley
Rhabdoblennius Whitley, 1930, p. 20 [type-species: Blennius rhabdotrachelus Fowler and Ball, 1924, by original designation].
Nixiblennius Whitley, 1930, p. 20 [type-species: Blennius snowi Fowler, 1928, by original designation].
Scartoblennius Fowler, 1946, p. 174 [type-species: Blennius ellipes Jordan and Starks, 1906, by original designation].
DIAGNOSIS.—Dentary a closed capsule (open mesially in R. snowi, but with large gaps and foramina in the wall laterally) with replacement teeth entering functional series through foramina in jaw bone. Anterior dentary canines absent; posterior dentary canines present. Premaxillary and dentary teeth immovable or nearly so, numbering 44 to 50 in upper jaw and 28 to 38 in lower jaw. Vomer with small conical teeth. Dorsal rays XII, 18 to 21; anal rays II, 20 to 22; segmented caudal rays 13, middle 9 branched; pectoral rays 13 or 14 (usually 14); pelvic rays I,3. Terminal anal ray bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculomandibular pores without cirri. Middorsal supratemporal pores 3–10. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri absent; supraorbital and nasal cirri simple. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
Norman (1943) first placed Nixiblennius in synonymy with Rhabdoblennius. Schultz and Chapman (1960) first synonymized Scartoblennius. Whitley (1930) proposed the genus Nixiblennius for Blennius snowi on the same page that he erected the genus Rhabdoblennius for B. rhabdotrachelus, but gave no reasons for erecting the genera and no characters to separate them. Later, Fowler (1946) proposed the genus Scartoblennius for Blennius ellipes, but did not compare it with either Rhabdoblennius or Nixiblennius. Rhabdoblennius rhabdotrachelus does differ from the previously mentioned species in possessing a “pseudobasisphenoid” (Springer, 1968, p. 44), but we do not believe that this difference warrants rhabdotrachelus being placed in its own monotypic genus. The species comprising Rhabdoblennius are few in number (probably only 6 or 7) and, except for the presence of a pseudobasisphenoid in rhabdotrachelus, are quite close osteologically.
DISTRIBUTION.—Central and western Pacific Ocean.
Salarias Cuvier
Salarias Cuvier, 1817, p. 251 [type-species: S. quadripennis Cuvier, 1817, = Blennius fasciatus Bloch, 1786, by monotypy].
Erpichthys Swainson, 1839, p. 275 [type-species: Salarias quadripennis Cuvier, 1817, = Blennius fasciatus Bloch, 1786, by subsequent designation].
Negoscartes Whitley, 1930, p. 20 [type-species: Salarias irroratus Alleyne and Macleay, 1877, by original designation].
Crenalticus Whitley, 1930, p. 21 [type-species: Salarias crenulatus pallidus Whitley, 1926, = Salarias sinuosus Snyder, 1908, by original designation].
DIAGNOSIS—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering 110 to 168 in upper jaw and 72 to 134 in lower jaw. Vomer toothless. Dorsal rays XII or XIII, 16 to 20; anal rays II, 18 to 21; segmented caudal rays 13, middle 9 branched; pectoral rays 13 to 15 (rarely 15); pelvic rays I, 3 (innermost pelvic ray minute, visible only in cleared and stained material). Terminal anal ray bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculomandibular pores without cirri. Middorsal supratemporal pores 1–3. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest present or absent. Nuchal cirri simple or palmate; supraorbital and nasal cirri simple to multifid. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
Whitley (1930, p. 20) erected the genus Negoscartes to include those species of Salarias that have a well-developed notch in the dorsal fin and large mandibular canines. Salarias fasciatus was erroneously thought to lack canines. The degree of notching in the dorsal fin varies from well developed to absent in the species of Salarias and in other genera of Salariini (for example Ecsenius). Norman (1943) also recognized Negoscartes, but the defining characters he used are without substance. Salarias fasciatus and S. ceramensis do differ from the other species of Salarias in attaining a greater size as adults, having more teeth in the upper jaw, and nuchal and supraorbital cirri that are broad and multifid. Several of these characters, however, appear to show a trend, with fasciatus and ceramensis merely the extremes. We believe that the characters used in the key and diagnosis to define the genus are of much greater importance than the differences noted above.
The genus Crenalticus (Whitley, 1930, p. 21) was proposed for Salarias crenulatus (Weber) [= S. sinuosus Snyder], which differs from the other species of Salarias in possessing a crenulate upper lip. We consider this character to be of only specific significance. Springer (1967) found that the degree or presence of crenulation of the upper lip was only of specific importance in the salariinine genus Entomacrodus.
DISTRIBUTION.—Red Sea; Indian Ocean; central and western Pacific Ocean.
Scartichthys Jordan and Evermann
Scartes Jordan and Evermann, 1896, p. 471 [type-species: Salarias rubropunctatus Valenciennes in Cuvier and Valencinnes, 1836, by original designation; a junior homonym of Scartes Swainson, 1835, Mammalia].
Scartichthys Jordan and Evermann, 1898, p. 2395 [type-species: Salarias rubropunctatus Valenciennes in Cuvier and Valenciennes, 1836, by original designation].
DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines absent; posterior dentary canines present. Premaxillary and dentary teeth freely movable, numbering 144 to 256 in upper jaw and 67 to 123 in lower jaw. Vomer toothless. Dorsal rays XII, 17 or 18; anal rays II, 18 to 20; segmented caudal rays 13, middle 9 branched; pectoral rays 14; pelvic rays I,4. Terminal anal ray not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; no scalelike flaps covering lateral-line pores. Preoperculamandibular pores without cirri. Middorsal supratemporal pores 5–12. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal cirri consisting of a patch of cirri on each side of nape separated by a wide hiatus equal to or greater than length of base of either patch of cirri; supraorbital and nasal cirri multifid. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
Specimens housed in the National Museum of Natural History that we have examined indicate that the genus consists of at least two well-defined species (also see discussion under Cirripectes).
The species of Scartichthys constitute the largest members of the family Blenniidae, although not the longest. (Xiphasia Swainson, of the Nemophidinae, is eel-like and attains a length over 500 mm SL.) Adults frequently exceed 175 mm in standard length.
DISTRIBUTION.—Restricted to the eastern Pacific Ocean.
Stanulus Smith
Stanulus Smith, 1959, p. 246 [type-species: S. seychellensis Smith, 1959, by original designation].
Fallacirripectes Schultz and Chapman, 1960, p. 362 [type-species: F. minutus Schultz and Chapman, 1960, = Stanulus seychellensis Smith, 1959, by original designation].
DIAGNOSIS.—Dentary an open capsule with replacement teeth entering functional series through excavated area in jawbone. Anterior dentary canines absent; posterior dentary canines present. Premaxillary and dentary canines freely movable, numbering 118 to 138 in upper jaw and 62 to 82 in lower jaw. Vomer with or without teeth. Dorsal rays XII, 9 to 12; anal rays II, 10 to 13; segmented caudal rays 13, middle 9 branched; pectoral rays 14 to 16 (usually 15); pelvic rays I, 4. Terminal anal ray not bound by membrane to caudal peduncle. Lateral line not consisting of two overlapping disconnected portions; minute, imbricate, scalelike flaps covering anterior lateral-line pores. Preoperculomandibular pores without cirri. Middorsal supratemporal pores 3–22. Upper lip without free dorsal margin. No cup-shaped fleshy disk or appendage behind lower lip. Gill membranes free. Occipital crest absent. Nuchal, supraorbital, and nasal cirri simple. Postcleithra consisting of two elongate bones, head of ventral element overlapping ventral end of dorsal element. Lateral extrascapular not fused with pterotic. Median ethmoid ossified. Circumorbital bones 5.
Relationships are discussed under section titled “Recognition of Genera.”
This genus, consisting of two species, has been revised recently by Springer (1968).
DISTRIBUTION.—Indian Ocean; central and western Pacific Ocean.
Comparison of Cephalic Sensory Pores
Although the sensory canal system has generally been neglected in the classification of blenniid fishes, Springer (1967) used the nature of the sensory pores as a specific character to distinguish certain species of the salariinine genus Entomacrodus. In our study we find that the cephalic sensory pores have utility as generic characters and may indicate phyletic relationships. We have illustrated the pore patterns of representative species of each genus and subgenus of Salariini (Figures 17–50) as an adjunct to our generic discussions and to encourage future students of the Blenniidae to survey the character more fully.
Correlation between development of cephalic pores and the ecology and/or behavior of blenniid fishes is not readily apparent. Blennius ocellaris (subfamily Blenniinae), the deepest dwelling (to at least 400 meters) member of the family, has simple pores, as do species of the salariinine genera Alticus and Andamia, which spend much of their existence out of water. Unlike the previously mentioned genera, all species of the subfamily Nemophidinae spend most of their time actively swimming, yet they share in common a reduced sensory pore system. The only general observation that we can make regarding sensory pores is that there seems to be a strong trend for the pores to be reduced in number in the more specialized forms.
The cephalic sensory canals in the tribe Salariini agree with the typical perciform pattern described by Branson and Moore (1962). Some of the major sensory canals and associated pores of a representative salariinine genus are shown in Figure 16. The sensory pore series of the head that we found of importance are defined below.
Infraorbital (IFO) and supraorbital (SO) series: Pores in the positions occupied by these canals.
Supratemporal series (ST): Only pores in the region anterior to the first dorsal spine and between the bases of the nuchal cirri (if present), extending anteriorly across the top of the head, are discussed below. In some genera, for instance Entomacrodus, pores anterior to the level of the nuchal cirri frequently spread laterally and cover an expanse broader than the distance between the nuchal cirri; all pores anterior to the nuchal cirri in this mid-region are included in our counts. In some specimens this count may be difficult to make. Springer (1967, p. 6) termed pores in this series “predorsal commissural pores.”
Mandibular series (MDB): The number of mandibular pores on each side of the head beginning with the pore nearest the symphysis of the lower jaw and terminating posteriorly with the pore immediately behind or slightly dorsal to the membranous corner of the mouth. In some genera it is difficult to determine by external examination alone where the break occurs between the MDB series and the POP series. In the following discussion of pore patterns the reader should experience little difficulty determining which pores we include in either series if reference is made to the illustrations.
Preopercular series (POP): In this series we consider the first pore position (depending on the genus or species, a position may be occupied by one or more pores, all connected by a common canal with a single opening in the preopercular bone) as the dorsalmost, which occurs slightly above the level of the greatest distance between the posterior margins of the preopercle and opercle, excluding the branchiostegal membrane.
In the Salariini there may be either five or six positions in the POP series, depending on the genus, describing an arc between the first and last positions. The last pore position is the ventralmost of the series and is posterior to the last MDB pore. In some genera the last POP position is always occupied by a vertical pair of pores, in other genera by a single pore. On the figures we have placed an “x” below the last POP position.
In the following account of pore patterns the reader should refer to Figures 17–50. Unless stated otherwise, the species illustrated is considered typical for all species of the genus. To facilitate comparison of genera the sequence of presentation follows that of Figure 51 (from top to bottom). In many species the pores are difficult to see and are easily overlooked. Overlying mucus must be carefully removed and drying of the area in which pores occur greatly facilitates examination.
Ecsenius (Figures 17–18). All pore positions occupied by simple pores. MDB pores 4. While the basic pore pattern is the same for both subgenera, there has been a posterior displacement of the mandibular pores in Anthiiblennius, perhaps correlated with changes in the structure of the lower jaw. Anthiiblennius also differs in having a more anterior placement of the middorsal ST pore. This shift in position is apparently the result of the more anterior placement of the first dorsal spine and its pterygiophore.
Andamia (Figure 20). All pore positions occupied by simple pores. A single MBD pore. Reduction of MDB pores is probably correlated with the presence of a prominent cup-shaped fleshy disk behind the lower lip (Figure 10). A single middorsal ST pore present.
Alticus (Figure 19). All pore positions occupied by simple pores. MDB pores 2 or 3. Reduction of MDB pores in Alticus and Andamia (4 or more in other salariinine genera except group 21 of Figure 51) is another reason for placing Alticus as we do in Figure 51. A single middorsal ST pore present.
Praealticus (Figure 21). All pore position occupied by simple pores. MDB pores 4. Distribution of ST pores is identical to that found in the related Alticus and Andamia, and is characterized primarily by the presence of a single middorsal ST pore.
Istiblennius (Figures 22–25). This large assemblage of species, which shows the greatest range of variation in its other characters, likewise exhibits much variation in pore development. IFO positions occupied by multiple or simple pores, simple in most species. MDB pores 5 or 6, 5 in most species. POP positions consisting of simple pores in all species except I. gibbifrons in which the pores at each position are double. Also, since an ophioblennius larval stage is known only for gibbifrons among the Istiblennius species, placement of gibbifrons in a separate genus may be indicated (see discussion under general synoptic account). Middorsal ST pores 0–12 or more, 3 or more in all except I. steinitzi.
The genera illustrated in Figures 29–34 (group 7 of Figure 51) are very difficult to characterize individually because of the complex nature of their sensory pores. In Pereulixia (Figure 29), Exalhas (Figure 30), and Cirripectes (Figures 31–32) most or all of the pore positions are occupied by relatively small and numerous pores arranged in clusters. Each pore in a cluster at a given position is united by a minute branch to a common canal. In most species of Cirripectes these pores are very minute and easily overlooked. In Scartichthys (Figure 33) and Ophioblennius (Figure 34) the pores are larger and less numerous. In all of these genera there tends to be an increase in the number of pores with increase in standard length.
Stanulus (Figure 37). IFO typically containing positions with some multiple pores. POP positions typically with simple pores. MDB pores 4 or 5. Middorsal ST pores 3–22, the number increases with increase in standard length (Springer, 1968a).
Entomacrodus (Figures 35–36). IFO typically containing several positions with multiple pores. POP positions variable, consisting of simple, paired or mulitple pores. MDB pore positions (some positions may have multiple pores) 4–6. ST pores in middorsal region 3 to more than 50. In those species that have a relatively high number of ST pores, the number increases with increase in standard length.
Nannosalarias (Figure 38). IFO positions occupied by multiple pores. POP positions with simple pores. MDB pores usually 5, occasionally 4. Middorsal ST pores typically 3.
Crossosalarias (Figure 26). Some IFO positions typically with multiple pores. POP positions with simple pores. MDB pores 5. Middorsal ST pores 3.
Atrosalarias (Figure 27). All pore positions occupied by simple pores. MDB pores 4. A single middorsal ST pore present.
Salarias (Figures 16 and 28). All pore positions occupied by simple pores. MDB pores 4. Middorsal ST pores 1–3.
Glyptoparus (Figure 41). All pore positions occupied by simple pores. MDB pores 4. Mid-dorsal ST pores 1 or 2, usually 2.
Medusablennius (Figure 44). All pore positions occupied by simple pores. MDB pores 3. Mid-dorsal ST pores 2.
Mimoblennius (Figures 39–40). IFO with some positions occupied by multiple pores. POP positions with simple pores except for the last position, which consists of a vertical pair of pores. MDB pores 3. Mid-dorsal ST pores 3.
Litobranchus (Figure 42). All pore positions occupied by simple pores. MDB pores 3. Mid-dorsal ST pores 2–4, usually 2.
Rhabdoblennius (Figures 43 and 45). IFO positions occupied by simple or multiple pores. POP positions with simple pores except for the last position, which consists of a vertical pair of pores. MDB pores 3 or 4. Mid-dorsal ST pores 3–10.
Hirculops (Figure 46). IFO with some positions occupied by multiple pores. POP pores simple except for last position, which consists of a vertical pair of pores. MDB pores 3. Middorsal ST pores 8–15.
Antennablennius (Figures 47–48). IFO with some positions occupied by multiple pores. All POP positions occupied by simple pores. MDB pores 3. Middorsal ST pores 1–4, typically 3 in subgenus Antennablennius and usually 2 in subgenus Croaltus (see general comments under Antennablennius).
Alloblennius (Figures 49–50). IFO positions usually occupied by simple pores or with relatively few paired pores. All POP positions occupied by simple pores. MDB pores 3. Mid-dorsal ST pores 3–8.
Recognition of Genera
In any study embracing as many genera and species as in our study, it behooves the authors to discuss their generic concept. The only rule that can be consistently applied to the genus-group category is that the species of a particular taxon be more closely related to each other than to species in another taxon of equal rank. Recognition of monotypic taxa, above the species level, simply infers a high degree of differentiation in a single species. Obviously, the level of differentiation that one chooses to regard as meriting generic (as opposed to subgeneric) recognition will depend upon the number and stability of characters within each family of animals considered and the subjective bias of the taxonomist involved. In the final analysis the genus-subgenus system of nomenclature is subjective.
For every genus that we recognize we have spent much time in discussion and debate challenging each other to defend an action, whether it be to synonymize a genus, recognize one in synonymy, erect a new genus, or not erect a genus where one might be recognizable. We have taken special note of the fact that some of these genera are as yet poorly known. From these discussions has evolved the dendrogram shown in Figure 51, which is a schematic representation of our concept of the relationships of the salariinine genera that we recognize. The dendrogram does not imply that one genus is more primitive than, or ancestral to, another. The arrangement simply places those genera that we consider to be most closely related together in brackets. In the following discussion we attempt to explain the reasons for our arrangement of the genera of Salariini in Figure 51 and comment on why we recognize some of them. Numbers given in the following discussion correspond to those shown on the figure.
1. Ecsenius is the most distinct genus in the Salariini. Indeed, it occupies a unique position in the Blenniidae and perhaps merits a separate tribe. The general nature of its dentition, however, at least places Ecsenius as an offshoot of the Salariini. The following information is the result of a revision of the genus by Springer. Ecsenius differs from all other blenniids in having dorsal processes extending from the proximal portions of the epipleural ribs on at least the third through fifth vertebrae; in having both anterior and posterior canines on the dentary (the anterior canines are difficult to distinguish from the incisoriform teeth in most species, but their nature is apparent in osteological preparations); and in lacking an ossified median ethmoid. In addition to the above characters, Ecsenius also differs from the other salariinine genera in having the lateral extrascapular fused with the pterotic.
2. Excluding Ecsenius, there appear to be at least four major phyletic lines within the Salariini. Their relationships to each other, however, are not clear. Alticus and its allies comprise a group of slender, elongate species that closely approach each other in general physiognomy. They share in common high numbers of dentary and premaxillary teeth, a reduced pore system, a fleshy occipital crest (secondarily lost in all except one species of Andamia), and loss of nuchal cirri. There has been a tendency for the ventralmost rays of the caudal fin to become thickened in most species, probably in response to their habit of frequently leaving the water and using their tails to hop from one pool to another. They have either unbranched caudal rays or the number of branched rays is reduced, usually 4 to 8 (typically 9 in those Salariini with branched rays). With the exception of one species of the unrelated genus Ecsenius (E. midas) and one species of Praealticus, this group of genera is unique among the Salariini in having the postcleithra consisting of one to three small, fragile, well-separated bones on each side (Figures 15a, b, c).
3. Alticus and Andamia are clearly more closely related to each other than to Praealticus. In adults of Praealticus, unlike Alticus and Andamia, the tips of some of the caudal rays are always branched and the terminal anal pterygiophore frequently bears two rays (one in Alticus and Andamia). With the exception of Alticus simplicirrus (see text discussion under Alticus), the general nature of the supraorbital cirri is different in the two groups.
4. Andamia is unique among the genera of Salariini in having a cup-shaped fleshy disk or appendage behind the lower lip (Figures 10 and 20).
5. The precise placement of Istiblennius is uncertain, but if difficulty in distinguishing the genus is an indication, then it is closely related to Praealticus (see text discussion under Praealticus). Since an ophioblennius larval stage is known for only one species of Istiblennius (gibbifrons), for none of the presumed closest relatives of Istiblennius (group 2), and for most (if not all) of the unrelated genera of group 6, Istiblennius as here defined may be polyphyletic.
6. Included are those genera that share the following combination of characters: ophioblennius larvae, the anal spines of mature males enveloped in conspicuous fleshy rugosities, multiple pores on the head tending to increase in number with increase in standard length, I, 4 pelvic rays (except for one species of Cirripectes, which has I, 3), and the terminal anal ray not bound by a membrane to the caudal peduncle. Most of the species typically have the last anal pterygiophore bearing two rays.
7. Included are those genera with complete lateral lines (extending almost to the caudal base) and nuchal cirri consisting of a transverse series across the nape or a patch of cirri on each side of the nape. One species of Entomacrodus (E. marmoratus) overlaps this group.
8. Ophioblennius differs from its nearest relatives in having the lateral line consisting of two overlapping, disconnected portions (instead of being continuous to the caudal base), the dorsal fin without a notch between spinous and segmented portions (but notched somewhat in larvae), and in having more segmented dorsal and anal rays. It is the only genus of the group that has become established in the Atlantic Ocean.
9. Pereulixia differs from its nearest allies in having small, imbricate, scalelike flaps covering at least the anterior lateral-line pores, vomerine teeth, simple supraorbital cirri that are long and tapering (simple but short in Cirripectes jenningsi), and the upper lip entire.
10. Exallias differs from both Cirripectes and Scartichthys in having about one-third as many teeth in the lower jaw as in the upper (instead of about one-half as many teeth in the lower jaw as in the upper), no dentary canines, a pair of barbels on each side of the chin just behind the lower lip, and fewer segmented dorsal and anal rays.
11. We know of no single character that alone will distinguish Cirripectes and Scartichthys from their respective relatives. This, it seems to us, is not too surprising when dealing with a relatively large number of genera and species that have evolved along similar lines.
12. As indicated by the dendrogram, Cirripectes and Scartichthys are quite closely related. We considered treating them as subgenera but believe that it would be premature to do so until the species of these genera are better known. For the present we recognize both genera because they are well entrenched in the literature (Strasburg and Schultz, 1953; Cohen, 1956), and we encountered no difficulty in assigning species to one or the other of these genera (see discussion under Cirripectes).
13. Both Entomacrodus and Stanulus have recently been revised by Springer (1967; 1968). There is considerable overall similarity between the two species of Stanulus, especially S. talboti, and the species of Entomacrodus.
14. Stanulus differs from Entomacrodus primarily in having minute, imbricate scalelike flaps covering the lateral-line pores, typically one less dorsal spine, one more pectoral ray, and fewer segmented dorsal and anal rays.
15. Nannosalarias does not appear to show especially close affinities to any other genus. We tentatively place it near the base of those genera centering around Cirripectes and Entomacrodus because it shares with that group a distinctive type of sexual dimorphism in which the anal spines of mature males become enveloped in conspicuous fleshy rugosities. Nannosalarias differs from those genera in the general nature of its head pore distribution, and in having one less circumorbital bone, one less segmented pelvic ray, and a membranous attachment of the terminal anal ray to the caudal peduncle.
16. Such a wide range of characters is shown between the species of Crossosalarias, Atrosalarias, and Salarias that it is difficult to define them collectively as a group. The various combinations of characters shared between the three genera, together with their general physiognomy, however, leads us to believe that they are more closely related to each other than to other genera of Salariini.
17. Crossosalarias differs from all other genera of Salariini in having restricted gill openings (the gill membranes are bound to the isthmus at about, or slightly below, the level of the ventralmost pectoral ray) and cirri associated with 3 or 4 preoperculomandibular pores on each side. It differs from Salarias and Atrosalarias in having the innermost pelvic ray well developed, as opposed to rudimentary, and a fleshy flap at the base of the first dorsal spine.
18. Atrosalarias differs from both Salarias and Crossosalarias in having no branched caudal rays, typically one more pectoral ray in each fin (usually 16), one to three less dorsal spines, and in having the skin of the anal spines and first several segmented anal rays rugose in mature males (see illustration in Springer and Smith-Vaniz, 1968).
19. Salarias differs from all other genera of Blenniidae in having the anterior anal segmented rays elongated in large, mature males.
20. We place Glyptoparus as we do because it does not appear to show any obvious affinities with the other groups of salariinine genera. As in one species of Ecsenius (E. mandibularis), Glyptoparus frequently has more than one (usually two or three) posterior canines on each dentary, arranged in linear series. It does not, however, have both anterior and posterior dentary canines or any of the other numerous distinguishing osteological characters of Ecsenius.
21. Included here are those genera that have relatively few premaxillary (18 to 50) and dentary (16 to 38) teeth, and have the dentary replacement teeth generally making their appearance through foramina in the bone (Figure 7, except Rhabdoblennius snowi, Figure 6, in which the dentaries are moderately excavated). All of the species belonging to this group of genera are relatively small, have the terminal anal ray bound by a membrane to the caudal peduncle, and usually 14 pectoral rays (except Medusablennius, which has 13).
22. Medusablennius differs from its nearest relatives in having considerably fewer teeth, only two circumorbital bones (no other salariinine has less than 4), typically 13 pectoral rays, and attaining maturity at a size when the sex of the other species is hardly distinguishable. While its complex cirri are to some extent duplicated in Mimoblennius cirrosus, no salariinine species has such well-developed cirri at so small a size. Simple caudal rays are found in group 21 only in Medusablennius and Litobranchus; Medusablennius, however, has 13 segmented rays and Litobranchus has only 12. Except for Alloblennius pictus, Medusablennius is the only genus of salariinines in which the ventral hypural plate is fused to the urostylar centrum.
23. Mimoblennius differs from all other genera in group 21 in having 13, instead of typically 12 dorsal spines. Although Mimoblennius has no unique character that will separate it from all other genera, when compared with individual genera in group 21 there are numerous characters that set it apart.
24. Litobranchus differs from all members of group 21, except Medusablennius, in having simple caudal rays. It differs from all members of group 21 in having only 12 instead of 13 segmented caudal rays.
25. We believe that the genera of group 25 are more closely related to each other than to any of the other genera in group 21 because they appear to be relatively unspecialized when compared with the other genera. They exhibit none of the obvious specializations of cirri or caudal fin structure found in the other three genera. In his review of the Blenniidae, Norman (1943) regarded Rhabdoblennius and Antennablennius as the most primitive members of the subfamily Salariinae. He considered these two genera so closely related that he accorded them only subgeneric rank.
26. Rhabdoblennius and Hirculops differ from Antennablennius and Alloblennius in having vomerine teeth, well-developed canines in the lower jaw (instead of lacking canines or having them only weakly developed), and in having the last POP pore position with a vertical pair of pores (instead of with a single pore). Our concept of the relationship of these four genera to each other has been influenced by their geographic distributions (see general accounts). The members of each pair of genera are closely related and occupy adjacent geographic areas with little or only slight overlap. In each pair one genus is relatively speciose and the other is either monotypic (Hirculops) or known to comprise only two species (Alloblennius).
27. Hirculops differs from Rhabdoblennius and all other genera included in group 21 in consistently having 11 (instead of 10) precaudal vertebrae and in having the premaxillary closed laterally with replacement teeth entering the functional row through foramina in the bone (Figure 4). Hirculops can also be distinguished from Rhabdoblennius by its extremely long supraorbital cirri (Figure 46) and by its small nuchal cirri (absent in Rhabdoblennius). The mutually exclusive geographic distributions of these two taxa reinforces their generic separation.
28. Antennablennius differs from Alloblennius in having 5, instead of 4, circumorbitals, and in not having supraorbital cirri. Alloblennius is apparently restricted to the Red Sea and Antennablennius is known from the Red Sea and Indian Ocean.
Annotated List of Nominal Species, Tribe Salariini
The following list gives in order: (1) the scientific name in alphabetical order by species as it originally appeared (hyphens omitted and capitalized specific names in lower case, however); (2) the author or authors (Cuvier and Valenciennes, 1836, is abbreviated to C & V, 1836); (3) date of publication (some species were described as new by the same author on two occasions); (4) page reference (see literature cited section for complete reference); (5) the genus to which we currently assign the species, if such is possible; (6) one or two letters of the alphabet that indicate the basis on which generic allocation was made (or other pertinent information), as follows: A—holotype, lectotype, neotype, or syntypes examined; B—para-type examined; C—literature or other information obtained sufficient for determination; D—probably belongs in this genus, but should be verified (in some instances this is impossible); E—questionable, could belong in a different tribe of Blenniidae but probably not; F—name unavailable by reason of not having been presented properly, a nomen nudum; (7) one or more museum abbreviations (see methods section for corresponding museum name) and catalog numbers indicating holotypes, lectotypes, neotypes, or syntypes. (Parentheses around information in 7 signifies that either we are not certain of the information or have not seen the specimens in question). Occasionally, comments referring to a particular name have been included immediately below that name. Noteworthy misspellings have been included, as well as some names of questionable validity.
Following this list is a list of available names in the genus Salarias that refer to species belonging in blenniid tribes other than the Salariini. We know of no other species originally described in genera of the Salariini that do not belong in the Salariini.
Tables of Generic Characters
- bibliographic citation
- Smith-Vaniz, William F. and Springer, Victor G. 1971. "Synopsis of the tribe Salariini, with description of five new genera and three new species (Pisces: Blenniidae)." Smithsonian Contributions to Zoology. 1-72. https://doi.org/10.5479/si.00810282.73
