SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

Emphasis upon publication as a means of “diffusing knowledge” was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.” This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Annals of Flight

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoology

Smithsonian Studies in Air and Space

Smithsonian Studies in History and Technology

In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world.

Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover.

S. Dillon Ripley
Secretary
Smithsonian Institution

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 236

Four New Species of the Fish Genus Ecsenius with Notes on Other Species of the Genus (Blenniidae: Salariini)

James F. McKinney
and Victor G. Springer

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976

A B S T R A C T

McKinney, James F., and Victor G. Springer. Four New Species of the Fish Genus Ecsenius with Notes on Other Species of the Genus (Blenniidae: Salariini). Smithsonian Contributions to Zoology, number 236, 27 pages, 11 figures, 12 tables, 1976.—Four new species of Ecsenius are described: E. isos (from the New Hebrides), E. melarchus (from islands in the Java Sea, Celebes, and Borneo), E. pictus (from the Moluccas and the Solomon Islands), and E. schroederi (from the Moluccas). New information (locality records and geographic and individual morphological variation) is given for 13 previously described species, one of which, E. oculus, is represented by 6 allopatric populations, each completely separable from the others only by features of its color pattern. A key to all Ecsenius species (25) is presented.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

McKinney, James F.

Four new species of the fish genus Ecsenius with notes on other species of the genus (Blenniidae, Salariini)

(Smithsonian contributions to zoology : no. 236)

Bibliography: p.

Supt. of Docs. no.: SI 1.27:236

1. Ecsenius. I. Springer, Victor Gruschka, 1928– joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology; no. 236.

QL1.S54 no. 236 [QL638.B6] 591′.08s [597′.58] 76–608086

Four New Species of the Fish Genus Ecsenius with Notes on Other Species of the Genus (Blenniidae: Salariini)

James F. McKinney
and Victor G. Springer

____________

James F. McKinney and Victor G. Springer, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560.

Introduction

Numerous specimens of the genus Ecsenius McCulloch, including representatives of four new species, have come to our attention since Springer’s (1971, 1972) studies of this genus. The purpose of this paper is to describe the new species (all in the subgenus Ecsenius) and present new data for thirteen other species of Ecsenius. A key to the 25 species of Ecsenius (probably the most speciose genus of blenniids) is included. Two species, E. bandanus and E. trilineatus, hypothesized by Springer (1971:42–43) to occur in the Banda Sea, based on collections lacking locality data, are now demonstrated to occur in that area; a collection of E. oculus also hypothesized by Springer to have come from the Banda Sea, now appears more probably to have been collected at Samoa (see “Remarks” under each species). We are of the opinion that there are more undescribed species of Ecsenius, particularly in areas (for instance the Philippines) that have not been subjected to intensive collecting.

Methods of reporting data are the same as thos given by Springer (1971). Material lists include ai abbreviated locality, followed by institutional abbreviation, catalog number, and (except for E. bicolor, E. nalolo, and E. yaeyamaensis) the number of specimens and size range, in parentheses, as mm SL.

ABBREVIATIONS.—The following institutional abbreviations are used to indicate the location of specimens studied: AM = Australian Museum, Sydney; ANSP = Academy of Natural Sciences of Philadelphia; BMNH = British Museum (Natural History), London; BPBM = Bernice P. Bishop Museum, Honolulu; CAS = California Academy of Sciences, San Francisco; DASF = Department of Agriculture, Stock and Fisheries (Fisheries Research), Papua; FMNH = Field Museum of Natural History; NMNH = National Museum of Natural History (Division of Fishes), Smithsonian Institution, Washington; UG = University of Guam, Agana; USNM = former United States National Museum (specimens in NMNH); WAM = Western Australian Museum, Perth.

ACKNOWLEDGMENTS.—We wish to thank the following individuals for loans or exchanges of specimens and for providing information on material housed in their institutions: J. R. Paxton and D. F. Hoese (AM); J. E. Böhlke and W. F. Smith-Vaniz (ANSP); J. E. Randall (BPBM); W. N. Eschmeyer and T. Iwamoto (CAS); P. Kailola (formerly of DASF); L. P. Woods (FMNH); H. T. Kami (UG); G. R. Allen (WAM); B. Carlson, formerly of the University of South Pacific, Suva, Fiji; P. Fourmanoir, New Caledonia. Allen, Randall, and Carlson also provided color transparencies of freshly preserved specimens of some species.

The manuscript was critically reviewed by W. D. Anderson, Jr. (Grice Marine Biological Laboratory, College of Charleston), W. F. Smith-Vaniz, and R. H. Gibbs, Jr. (NMNH).

Illustrations were prepared by S. L. Chambers (Figure 5), P. K. Hollingsworth (Figures 2 and 7), and J. R. Schroeder (Figures 1, 4, 10, and 11).

Many of the Indonesian specimens included in our study were obtained on an expedition funded by a National Geographic Society research grant to Springer. Springer also wishes to thank Dr. Aprilani Soegiarto, National Institute of Oceanology of Indonesia, for his generous help during Springer’s visits to Indonesia.

Ecsenius bandanus Springer

FIGURE 1

Since publication of the original description, based on a single specimen (from a Longley collection of dubious origin), 124 specimens of E. bandanus from various localities in Indonesia (see “New Material”) have been obtained. On the basis of these specimens the species is redescribed.

DESCRIPTION (Table 1 gives frequency distributions for certain meristic characters; other characters, with frequencies in parentheses, are as follows).—Dorsal-fin spines 11(2), 12(119), 13(1); pectoral-fin rays 12(2), 13(70); segmented caudal-fin rays 13(39); dorsal procurrent caudal-fin rays 6(2), 7(24), 8(13); ventral procurrent caudal-fin rays 6(7), 7(30), 8(2); total caudal-fin elements 25(2), 26(3), 27(23), 28(9), 29(2); gill rakers 11(16), 12(21), 13(8); pseudobranchial filaments 5(8), 6(41), 7(2); lower jaw (one side) posterior canine teeth 0(1), 1(84), 2(1); total lower jaw posterior canine teeth 1(1), 2(84), 3(1); precaudal vertebrae 10(111); epipleural ribs 11(4), 12(36), 13(16), 14(1). Lateral line with no paired pores, extending posteriorly to below level of 8th(5), 9th(30), or 10th(27) dorsal-fin spine. Dorsal fin notched seven-ninths (2) or eight-ninths(44) length of first segmented dorsal-fin ray.

Color Pattern: In preservative, specimens are as described and figured by Springer (1971, fig. 35), or the dorsum of the head and body are considerably darker (Figure 1); dark head stripe may end at posterior margin of opercle or extend on to body, gradually fading out beneath spinous dorsal fin. Dorsal half of body either with faint, broad saddles or uniformly dusky, usually darker than ventral half of body.

In life, body pale buff and pink. Head stripe black, margined ventrally by bright yellowish stripe. Yellow stripes present on snout and dorsum of head.

Sexual dichromatism of the ventral side of the head is present and concerns the presence and nature of spots (Springer, 1971:39). The following tabulation indicates the frequencies (with size ranges in mm SL in parentheses) of the four types of pattern found on the underside of the head. A color transparency of a freshly preserved male specimen (BPBM 18054, 22.6 mm SL) shows the ventral head spots to be yellow; the yellow color is now faded and the spots are unpigmented.

The limited material of three of the other four species in the prooculis subgroup (of which E. bandanus is a member) shows similar dichromatism. Two of the three specimens of E. bimaculatus (all males) have dark spots on the ventral surface of the head (the third specimen is uniformly dusky); five of six E. collettei males have dark spots (the sixth is uniformly dusky), whereas the two females have unpigmented spots; seven of the nine E. prooculis males have dark spots; the other two, and the only known female, have unpigmented spots. The remaining species, E. isos, is characterized by having dark ventral head spots in both males and females. None of the other species of Ecsenius exhibit this type of spotting on the ventral surface of the head.

TABLE 1.—Frequency distributions for certain characters in populations of Ecsenius bandanus

FIGURE 1.—Ecsenius bandanus, USNM 209767, female, 25.8 ram SL, Ambon Island, Moluccas. (Drawn by J. R. Schroeder.)

FIGURE 2.—Distribution of the species of the Ecsenius prooculis subgroup. (Irregular broken line indicates approximate position of Andesite Line.)

SEXUAL DIMORPHISM.—In addition to the sexual dichromatism noted above, females of E. bandanus have higher mean dentary tooth counts than males (Table 1). Significant differences occur in the following populations: Seribu (t = 2.07; df = 17; 0.1 > p > .05); Kabaena (t = 6.21; df = 22; p = < .001); and Ceram (t = 6.73; df = 20; p = < .001).

The upper and/or lower segmented caudal-fin rays of males, but not females, tend to become elongated with increase in SL. The longest caudal-fin ray of males ranges from 22.3–43.8 percent SL; the longest caudal-fin ray of females ranges from 20.1–25.6 percent SL.

Males tend to exceed females in maximum standard length. The largest male examined is 33.5 mm SL, and the largest female is 29.2 mm SL. There are 11 collections containing both males and females, and the largest specimen in 9 of these is a male.

GEOGRAPHIC VARIATION.—Other than slightly higher averages for dorsal-fin, anal-fin, and caudal vertebral counts (Table 1) in the eastern end of the known range of E. bandanus, no geographic trends are noted.

REMARKS.—The holotype of E. bandanus, collected by W. H. Longley, was hypothesized to have come from the Banda Sea (Springer, 1971:42–43). The recent collection of E. bandanus from Ambon (= Amboina) in the Banda Sea supports Springer’s hypothesis that Longley’s collection was from the Banda Sea area rather than from Samoa or the Hawaiian Islands, the only other Pacific areas where Longley is known to have collected.

Ecsenius bandanus is known to occur in the following biogeographical areas as defined by Thorne (1963): the Javan district of the Malayan province and the Celebesian province (both in the Indo-Malayan subregion) and the Moluccan province of the Papuan subregion. No other member of the prooculis subgroup has been collected in these areas (Figure 2).

NEW MATERIAL.—INDONESIA. JAVA SEA: Seribu Islands, USNM 211988 (1 specimen: 21.2), 211993 (17:13.3–29.2), 211996 (18:15.6–33.5), BPBM 10854 (5:15.9–22.6); Karimundjawa Islands, USNM 211979 (11:23.7–31.6). CELEBES: Kabaena Island, AM I.18490-001 (10:17.3–28.6), USNM 211916 (14:22.5–29.2). MOLUCCAS: Ambon Island, USNM 209767 (3:25.8–29.2), 211958 (1:25.4); Ceram Island, USNM 209660 (2:19.4–20.2), 210171 (24:13.4–29.2), CAS 34243 (6:14.9–22.6), BMNH 1975.11.19.1–3 (3:22.0–24.6); Saparua Island, USNM 209991 (7:17.0–31.2), 210111 (1:26.5), 210374 (1:25.2).

Ecsenius bicolor (Day)

FIGURE 3

New locality records of E. bicolor are given in “New Material.” Springer (1971:27) reported that two basic color patterns are found in preserved specimens and noted also that some specimens exhibit a broad longitudinal stripe on the body. Inasmuch as only the “bicolored” pattern has been illustrated (Springer, 1971, figs. 20, 21), we present all three in Figure 3.

FIGURE 3.—The 3 color patterns of Ecsenius bicolor: a, bicolored pattern, USNM 214812, female, 43.3 mm SL, Efate, New Hebrides; b, uniformly dark pattern, WAM P25111-008, male, 54.0 mm SL, Dampier Archipelgao, Western Australia; c, striped pattern, ANSP 128012, male, 43.2 mm SL, Horsburgh Island, Cocos Keeling Islands.

NEW MATERIAL.—COCOS KEELING ISLAND. ANSP 128009, 128010, 128012. INDONESIA. JAVA SEA: Seribu Islands, USNM 211991, 211997; Karimundjawa Islands, USNM 211980; Bawean Island, USNM 211972. CELEBES: Kabaena Island, USNM 211905, 211914; Boeton (= Butung) Island, USNM 211970. MOLUCCAS: Ambon Island, USNM 210262, 211950; Haruku Island, USNM 209563; Saparua Island, USNM 211927; Banda Island, USNM 211932. WESTERN AUSTRALIA. Dampier Archipelago, WAM P25111-008. QUEENSLAND. Lizard Island, BPBM 15550. NEW GUINEA. Trobriand Islands, USNM 214734, 214735. SOLOMON ISLANDS. Florida Island, BPBM 15629; Guadalcanal, USNM 212031. NEW HEBRIDES. Efate Island, USNM 214812. AMERICAN SAMOA. Tutuila Island, BPBM 17516. MARIANA ISLANDS. Tinian Island, UG 5145. CAROLINE ISLANDS. Truk Island, BPBM 9073.

Ecsenius isos, new species

FIGURE 4

Ecsenius yaeyamaensis (Aoyagi).—Springer, 1971:32 [in part].

Ecsenius trilineatus Springer, 1972:2 [in part].

DESCRIPTION.—Dorsal-fin spines 12(4); segmented dorsal-fin rays 14(3), 15(1); segmented anal-fin rays 16 (4); pectoral-fin rays 13(4); segmented caudal-fin rays 13(4); dorsal procurrent caudal-fin rays 6(1), 7(2), 8(1); ventral procurrent caudal-fin rays 6(2), 7(1), 8(1); total caudal-fin elements 25(1), 26(1), 27(1), 29(1); gill rakers 13(1), 14(2); pseudobranchial filaments 5(4); dentary incisor teeth 43(1), 45(1), 46(1), 50(1); lower jaw (one side) posterior canine teeth 0(3), 1(1); total lower jaw posterior canine teeth 0(3), 2(1); precaudal vertebrae 10(4); caudal vertebrae 22(3), 23(1); epipleural ribs 11(3), 12(1). Lateral line with no paired pores, extending posteriorly to below level of 8th(1), 9th(1) or 10th(2) dorsal-fin spine. Dorsal-fin notched eight-ninths (3) length of first segmented dorsal-fin ray. Data for the holotype are given in Table 2.

Color Pattern: Head with series of diffuse, dusky markings extending from upper portion of opercle anteroventrally to underside of head (similar in position to dark head stripe of E. trilineatus), ventralmost mark in series is a prominent spot. Scattered dusky pigment and very diffuse spots elsewhere on side of head. Lower lip dusky anteriorly. Body with two rows of diffuse, dusky blotches, which grade into smaller, more discrete spots posteriorly; spots in the two rows faintly connected as if to form bars; slight amount of dusky pigment extending from posteriormost spots of both rows out on proximal portion of caudal fin; caudal fin otherwise immaculate. Fleshy pectoral-fin base with diffuse dusky marks; upper mark along dorsal margin of fin base, lower mark at mid-base and extending onto base of middle ray as dark spot; pectoral fin otherwise unmarked. Narrow dusky stripe on both spinous and segmented-ray portions of dorsal fin just distal to fin base (prominent on males, faint on females). Distal margin of segmented-ray portion of dorsal fin with dusky stripe. Anal fin with broad dusky stripe, tips of rays pale. Pelvic fins unmarked.

RELATIONSHIPS.—The specimen designated below as the holotype of E. isos was tentatively identified by Springer (1972) as E. trilineatus in his description of that species. The paratypes were treated by Springer (1971) as color pattern variants of E. yaeyamaensis (as were specimens of the then undescribed E. trilineatus) and were overlooked in the 1972 study. The specimens of E. isos were confused with E. yaeyamaensis and E. trilineatus because the specimens have diffuse, but similar, markings on the fleshy pectoral-fin base. However, E. isos belongs in the prooculis subgroup of the yaeyamaensis species group by virtue of its having a short lateral line, short nasal cirri and discrete dark spots on the ventral surface of the head. E. isos is the only member of the prooculis subgroup that has dark spots beneath the head in both sexes. In overall coloration, E. isos most closely resembles E. collettei but differs from the latter by having diffuse dusky marks on the pectoral-fin base, a narrow dusky stripe in the spinous portion of the dorsal fin and a series of dusky markings extending from the upper opercle anteroventrally to the underside of the head. Also, the anterior markings on the body of E. isos are much less discrete than those of E. collettei.

FIGURE 4.—Ecsenius isos, BPBM 12122, holotype male, 26.2 mm SL, Efate, New Hebrides, lateral view and ventral view of anterior region. (Drawn by J. R. Schroeder.)

TABLE 2.—Selected counts and measurements (percent SL) from the holotypes of some species of Ecsenius

REMARKS.—E. isos, the fifth species of the prooculis subgroup to be discovered, occupies the New Hebridean district of the Fijian province of the Polynesian subregion (as defined by Thorne, 1963). All five species of the prooculis subgroup are allopatric and relatively restricted in distribution (Figure 2).

ETYMOLOGY.—The specific name, from the Greek isos (equal or like) alludes to the presence of dark spots beneath the head on specimens of both sexes.

HOLOTYPE.—BPBM 12122, male, 26.2 mm SL, New Hebrides, Efate, British Bathing Beach, Pango Peninsula, collected by R. H. Snider, 22 November 1964.

PARATYPES.—USNM 195787, male, 20.6 mm SL and two females, 22.5 and 26.0 mm SL, New Hebrides, around coral heads in Espiritu Santo Harbor, collected by W. M. Chapman and H. Cheyne, 24 April 1944.

Ecsenius lineatus Klausewitz

New records (see “New Material”) extend the range of E. lineatus considerably. Also, Masuda, Araga, and Yoshino (1975) reported E. lineatus from Okinawa, Taketomi Island, the Bonin Islands, and the Izu Islands, all northern range extensions. On the basis of the specimen they illustrated (pl. 86 Q) we accept their records.

Although there are few specimens from any locality, geographic variation in frequency distributions for some meristic characters, especially precaudal vertebrae (Table 3), is apparent. Geographic variation in number of precaudal vertebrae has been reported also for Ecsenius namiyei (Springer, 1971) and some species of Omobranchus (Springer and Gomon, 1975).

NEW MATERIAL.—CEYLON. USNM 213493 (1 specimen: 19.7), BPBM 18056 (3:35.3–10.1). MAURITIUS. BPBM 15943 (4:44.9–54.0), 15946 (1:25.9), 15947 (2:19.7–56.5). REUNION ISLAND. BPBM 16276 (1:22.3). WESTERN AUSTRALIA. North West Cape, WAM P25371-006 (2:45.2–53.9).

Ecsenius lividinalis Chapman and Schultz

Nine additional specimens of E. lividinalis representing new locality records have been procured (see “New Material”). Ranges for meristic characters, where different from those given by Springer (1971), are as follows: dorsal procurrent caudal-fin rays 5–7; ventral procurrent caudal-fin rays 5–7 (count of 4, was recorded by Springer from specimen less than 25 mm SL, and should have been excluded); total caudal-fin elements 23–27; dentary incisor teeth 29–37; precaudal vertebrae 11; caudal vertebrae 18–19 (see below); epipleural ribs 11–14; lateral line ending posteriorly below level of 8th–11th dorsal-fin spine.

TABLE 3.—Frequency distributions for certain characters in populations of Ecsenius lineatus

Ecsenius lividinalis was reported by Springer (1971) to have 10 precaudal vertebrae in 9 of 10 specimens examined. Re-evaluation of his radiographs and radiographs of the new material show 11 to be the characteristic precaudal count.

Color Pattern: An Ektachrome slide of a freshly preserved Florida Island specimen (BPBM 16141, 21.4 mm SL) shows the following: top and sides of head dark brown, underside of head lighter with metallic blue tint. Body beneath spinous dorsal fin brown, grading into dirty yellow posteriorly; belly and ventrolateral part of body yellow. Black spot encompassing anus surrounded by brilliant white ring. Basal halves of dorsal and anal fins yellow, distal portions clear; lower two or three pectoral-fin rays black-tipped; scattered dark pigment along pectoral-fin and caudal-fin rays; interradial membranes clear; pelvic fins creamy white. Color in preservative: no distinct markings other than black anal spot; body dusky, darker anterodorsally.

GEOGRAPHIC VARIATION.—Specimens from the Solomon Islands have higher mean numbers of dentary incisor teeth and caudal vertebrae than do specimens from other localities (Table 4).

NEW MATERIAL.—INDONESIA. CELEBES: Boeton (= Butung) Island, USNM 211966 (5 specimens:27.6–37.1). MOLUCCAS. Saparua Island, USNM 209992 (2:18.0–26.1), 210105 (1:22.1). SOLOMON ISLANDS. Florida Island, BPBM 16141 (1:21.4).

TABLE 4.—Frequency distributions for certain characters in populations of Ecsenius lividinalis

Ecsenius melarchus, new species

FIGURE 5

Ecsenius stigmatura Fowler,—Springer, 1971:30 [in part].

DESCRIPTION (see Table 5 for frequencies of certain counts; other counts, with frequencies in parentheses, as follows).—Dorsal-fin spines 12(73); pectoral-fin rays 13(3), 14(57); segmented caudal-fin rays 13(5), 14(33); dorsal procurrent caudal-fin rays 7(22), 8(12); ventral procurrent caudal-fin rays 6(16), 7(16), 8(2); total caudal-fin elements 26(1), 27(13), 28(9), 29(10); pseudobranchial filaments 6(34), 7(1); lower jaw (one side) posterior canine teeth 0(13), 1(44), 2(1); total lower jaw posterior canine teeth 0(3), 1(12), 2(41), 3(1), 4(1); precaudal vertebrae 10(63), 11(1); caudal vertebrae 22(6), 23(50), 24(8); total vertebrae 32(6), 33(54), 34(10); epipleural ribs 11(2), 12(14), 13(4). Lateral line with no paired pores, extending posteriorly to below level of 8th(35), 9th(22) or 10th(1) dorsal-fin spine. Dorsal fin notched six-ninths (4), seven-ninths (26), or eight-ninths (2) length of first segmented dorsal-fin ray. Data for the holotype are given in Table 2.

Color Pattern: In preservative, head and body dusky brown, darker dorsally. Indistinct, narrow, dark stripe extends from posteroventral margin of eye to posterior margin of opercle at upper pectoral-fin base. Prominent dark spot extends anteriorly from anus. Dorsal fin dusky basally and along margins of rays; interradial membranes clear. Anal fin dusky with indistinct light stripe proximally. Basal portion of all caudal-fin rays and entire length of middle four rays dusky, remainder of fin immaculate. Pectoral and pelvic fins with dark pigment along rays; interradial membranes clear.

Ground color is variable; usually it is quite dark, but some specimens appear faded. Delay in fixation of specimens in the field, however, often results in specimens having a faded appearance.

One male specimen (34.5 mm SL) from Karimundjawa has a light stripe beneath the dusky head stripe (not as distinct as the pale stripe on the opercle of the closely related E. stigmatura). One male specimen (33.1 mm SL) from Kabaena exhibits a row of seven small, pale spots on its body beneath the dorsal fin, a row of six slightly larger, pale spots mid-laterally, and a pale triangular patch just anterior to the first dorsal-fin spine. Another male (29.5 mm SL) from Kabaena shows similar but more irregular spots on its body.

A Kodachrome slide of a freshly preserved specimen from the Seribu Islands shows a brilliant white stripe extending from beneath eye to upper base of pectoral fin and a faint orange stripe beneath the white stripe. Otherwise, the color shown is similar to that of alcohol-preserved specimens.

GEOGRAPHIC VARIATION.—No significant meristic differences between populations were noted.

SEXUAL DIMORPHISM,—The upper and/or lower caudal-fin rays of males generally become longer with increase in SL, a condition not found to any appreciable extent in females (males with longest caudal-fin ray ranging from 23.1–48.7 percent SL; females with longest ray 21.3–31.9 percent SL). No other significant meristic or morphometric differences between sexes were noted.

RELATIONSHIPS.—E. melarchus, E. stigmatura, and E. lividinalis constitute the stigmatura species group, which is characterized by having a black anal spot and typically 14 segmented caudal-fin rays. E. melarchus has higher numbers of dorsal-, anal-, and pectoral-fin rays and vertebrae than E. lividinalis. The depth of the dorsal-fin notch is typically greater for E. melarchus than E. lividinalis. E. melarchus can be distinguished from E. stigmatura primarily by the absence of the large, dark spot on the caudal peduncle. Additionally, the fresh color of E. melarchus is strikingly different from that of E. stigmatura (see above and color pattern description for E stigmatura). Although all the counts for E. melarchus and E. stigmatura overlap (Table 5), significant differences between the species exist in the means for segmented dorsal-fin rays (t = 3.84; df = 95; p = < .001), segmented anal-fin rays (t = 2.89; df = 94; .01 > p > .001), dentary incisor teeth (t = 4.35; df = 73; p = < .001), and gill rakers (t = 4.34; df = 44; p = < .001).

TABLE 5.—Frequency distributions for certain characters of Ecsenius melarchus and E. stigmatura

FIGURE 5.—Ecsenius melarchus, USNM 201815, paratype, male, 37.7 mm SL, Pulau Gaya, Darvel Bay, Borneo (anterior two dorsal-fin spines damaged), lateral view and ventral view of anal region. (Drawn by S. L. Chambers.)

REMARKS.—Springer (1971) had a single specimen (of E. melarchus) which he questionably assigned to E. stigmatura. The considerable additional material now available shows that the differences between the single specimen and E. stigmatura noted by Springer are constant and that the two forms are allopatric.

E. melarchus occurs on coral reefs at depths as shallow as 1.5 m, and was taken at one station where the maximum depth recorded was 30.5 m.

ETYMOLOGY.—The specific epithet, from the Greek mela (black) + archos (anus), refers to the characteristic black mark that extends from the anus and is herein used as a noun in apposition.

HOLOTYPE.—USNM 212229, male, 39.0 mm SL, Indonesia, off southwest coast of Karimundjawa, somewhat north of Greater Mendjangan Island, coral reef, ca. 05°52′30″S, ca. 110°25′40″E, depth 1.0–12.2 m, collected by V. G. Springer, et al, VGS 74-28, 29 March 1974.

PARATYPES.—INDONESIA. JAVA SEA. USNM 211896, 6 males, 3 females, 3 unsexed, 16.4–44.4 mm SL, taken with the holotype. USNM 212893, 10 males, 18 females, 4 unsexed, 17.5–35.9 mm SL, Seribu Islands, off west side of Pulau Ajer, ca. 05°46′S, ca. 106°35′E, depth to 18.3 m, VGS 74-32, 4 April 1974. CAS 34244, 2 males, 1 female, 30.2–32.0 mm SL, Seribu Islands, off west side of Pulau Tikus, Pulau Pari Group, ca. 05°51′S, ca. 106°34′E, depth 0–30.5 m, VGS 74-35, 6 April 1974. BPBM 18053, 2 males, 2 females, 22.4–25.2 mm SL, Seribu Islands, Pulau Putri, depth 1–1.5 m, collected by J. E. Randall, 16 February 1975. USNM 211894, 3 males, 5 females, 24.3–38.4 mm SL, Karimundjawa Islands, just south of Lesser Mendjangan Island, ca. 05°52′50″S, ca. 110°24′40″E, depth to 18.3 m, VGS 74-30, 30 March 1974. Celebes: USNM 211897, 2 males, 4 females, 24.0–36.1 mm SL, Kabaena Island, Tallabassi Bay, ca. 05°17′20″S, ca 122°04′E, depth 4–8 m, VGS 74-1 24 February 1974. AM I.18491-001, 5 males, 1 female, 25.0–45.3 mm SL, Kabaena Island, close to locality of USNM 211897, depth 2–15 m, VGS 74-2, 25 February 1974. Borneo: USNM 201815, male, 37.7 mm SL, Darvel Bay, Pulau Gaya, E. end of Borneo, depth to 3 m, TE VEGA Sta. 6, 1 February 1965.

Ecsenius midas Starck

New locality records for E. midas are given in the list below. Also, we have been informed of one specimen (CAS 34384, 40.2 mm SL) from Grand Comore Island. The eight specimens from Maré were reportedly collected at a depth of 55 m, the deepest known for any species of Ecsenius. The single specimen from Kabaena Island has a segmented dorsal-fin ray count of 18, one less than previously reported for this species.

NEW MATERIAL.—MAURITIUS. BPBM 15941 (1 specimen: 86.4), 15944 (3:35.2–77.3). COCOS KEELING ISLANDS. ANSP 128011 (2:42.0–48.6). INDONESIA. CELEBES: Kabaena Island, USNM 211913 (1:46.3). SOLOMON ISLANDS. Florida Island, AM I.17532-001 (1:75.5). NEW HEBRIDES, Efate Island, USNM 214736 (1:52.4). FIJI ISLANDS. USNM 214786 (1:67.2). LOYALTY ISLANDS. Maré Island, USNM 213727 (3:33.5–36.3), BPBM 19701 (5:31.2–40.5).

Ecsenius nalolo Smith

Thirty-five additional specimens of E. nalolo (FMNH 71367, 71368, BPBM 18055) collected in the Maldive Islands have the characteristic single dark stripe on the pectoral-fin base that distinguishes E. nalolo from its closest relative, E. yaeyamaensis, which has a Y-shaped stripe. The two species are allopatric: nalolo occurs only west of, and yaeyamaensis only east of, 75°E longitude. It would be important for determining the validity of the distinction between E. nalolo and E. yaeyamaensis to have specimens from points around the southernmost coast of India, if the species occur there.

Ecsenius namiyei (Jordan and Evermann)

Since Springer’s (1971) account of E. namiyei, we have obtained 57 specimens that represent new locality records, and 7 additional specimens from Taiwan (see “New Material”). Examination of these and the previously recorded specimens indicates the existence of significant differences in meristics between the sexes and among the populations (Table 6).

SEXUAL DIMORPHISM.—In addition to the earlier report of higher dorsal- and anal-fin ray and caudal vertebral counts for males from Taiwan, we found that Taiwan females have a higher mean number of dentary teeth than do Taiwan males (t = 2.62; df = 28; .02 > p > .01). There are indications (Table 6) that these sexual differences also occur in the other populations of E. namiyei.

TABLE 6.—Frequency distributions for certain characters in populations of Ecsenius namiyei

GEOGRAPHIC VARIATION.—Specimens from Taiwan exhibit considerably higher numbers of segmented dorsal- and anal-fin rays, caudal vertebrae, and dentary teeth than do specimens from other populations. Additionally, the majority of specimens from areas other than Taiwan have no paired lateral-line pores, while all specimens from Taiwan have at least one pair of pores. The Solomon Islands population of E. namiyei has more precaudal vertebrae and fewer caudal vertebrae than any of the other populations. Increase in number of precaudal vertebrae also occurs in the Mauritius population of E. lineatus, and Springer and Gomon (1975) have reported similar increases in western Australian populations of three species of Omobranchus. The factors causing this type of variation are unknown.

NEW MATERIAL (all represent new locality records except the Taiwan specimens).—TAIWAN: USNM 211323 (7 specimens:53.3–65.7). INDONESIA. CELEBES: Boeton (=Butung) Island, USNM 211969 (14:17.0–63.3); Kabaena Island, USNM 211904 (3:32.7–48.2), 211915 (18:25.6–60.1). MOLUCCAS: Ambon Island, USNM 209736 (1:43.6), 211951 (1:51.7), 211957 (1:52.3); Saparua Island, USNM 210088 (2:21.9–33.9), 210345 (4:23.3–51.4). NEW GUINEA. Port Moresby, BPBM 15930 (1:35.8), DASF uncataloged (4:45.5–63.0). SOLOMON ISLANDS. Florida Island, AM I.17500-002 (2:45.5–52.7), BPBM 15644 (4:49.1–60.8); Guadalcanal, USNM 212038 (1:32.0), 212040(1:50.8).

Ecsenius oculus Springer

FIGURE 6

A total of 146 specimens of E. oculus, most from collections representing new locality records (see “New Material”), have been examined for this study. Analysis of data obtained from this material together with a re-examination of previously reported specimens has revealed significant variation among and within populations of this wide-ranging species.

Springer (1971) mentioned the possibility that the color-pattern variations he noted were representative of population differences. His specimens embrace five of the six types we recognize, but Springer recognized only four types, confusing our types A and B. All six of the color-pattern types we recognize occur allopatrically (Figure 7) and, in general, are restricted in distribution. The salient characters of the types are described as follows (and briefly compared in Table 7).

TYPE A.—Nape and predorsal area with indistinct pale stripes, no prominent dark spots; prominent postocular stripe extending from posterior margin of eye to or slightly beyond dorsal insertion of pectoral fin; row of 7 or 8 (modally 8) pale-margined dark spots, each about equal in size to eye diameter, usually arranged in pairs on body; anterior body spots at level of postocular stripe; posteriormost body spot forming saddle at rear base of dorsal fin; 4 pairs of small, dark spots along dorsal body profile, 2 pairs beneath spinous dorsal fin and 2 beneath segmented-ray portion of dorsal fin.

TYPE B.—Nape with prominent dark spot on each side of midline; prominent dark postocular stripe present as in type A; row of 7–10 (modally 9) pale-margined dark spots on body, similar to those of type A, but somewhat smaller; anterior body spots above level of postocular stripe; dark spots along dorsal body profile absent or indistinct.

TYPE C.—Dark spots on nape present but less distinct than those of type B; postocular stripe present but not as prominent as that of type B; 6–9 (modally 9) pale-margined dark spots on body, similar in size to those of type B; anterior body spots above level of postocular stripe; dark spots along dorsal body profile absent or indistinct.

TYPE D.—Dark nape spots present but less obvious than those of type B; postocular stripe absent or indistinct; single, large, pale-margined dark spot on body, forming saddle at end of dorsal-fin base; spots along dorsal body profile absent.

TYPE E.—Two dark spots on nape as in type B; no postocular stripe but opercle with vertical pair of dark spots separated by narrow light stripe; 3–5 (modally 4) large, pale-margined, dark spots on body at base of segmented-ray portion of dorsal fin; 4–6 small, pale spots along spinous dorsal-fin base.

FIGURE 6.—The six color pattern types of Ecsenius oculus: a, type A, USNM 203140, holotype, male, 53.8 mm SL, Taiwan; b, type B, USNM 211995, male, 33.7 mm SL, Seribu Islands, Java Sea; c, type C, WAM P25374-018, female, 29.5 mm SL, North West Cape, Western Australia; d, type D, USNM 209577, male, 48.7 mm SL, Haruku Island, Moluccas; e, type E. USNM 206431, male, 37.1 mm SL, Trobriand Island, New Guinea; f, type F, USNM 214520, male, 37.8 mm SL, Viti Levu, Fiji.

FIGURE 7.—Distribution of the Ecsenius oculus subgroup; the 6 color pattern types (A-F) of E. oculus indicated. (Irregular broken line indicates approximate position of Andesite Line.)

TYPE F.—Dark spots on nape absent or indistinct; postocular stripe prominent as in type A; row of 5 or 6 (modally 6) pale-margined dark spots on body at level of postocular stripe plus ventrolateral row of 3 to 5 (modally 4) pale-margined dark spots; 4 or 5 smaller dark spots along dorsal body profile beneath spinous dorsal-fin base and 0–2 beneath segmented-ray portion of dorsal fin.

DISCUSSION.—Meristic data from the new material re-enforces Springer’s (1971, 1972) evidence for the existence of north-south clines (Figure 8 shows means for number of dentary teeth plotted against latitude as an example of a cline; means for segmented dorsal-fin, anal-fin, and caudal vertebral counts exhibit similar clines). These clines do not appear to be correlated with color pattern type (Tables 8 and 9). For example, the B and D color pattern types are least similar, yet these types are very similar in meristics. The least degree of similarity in meristics between contiguous populations exists between types A and B, whereas their color patterns are quite similar. The type F color pattern shows similarities to that of type A (both have prominent postocular stripes and lack dark spots on the nape) and both populations exhibit high counts, yet of all the populations of E. oculus, these are separated by the greatest distance. Clearly, these examples show that expression of the color pattern is independent of meristic variation, although both can be correlated with geography.

The six color pattern types of E. oculus could reasonably be recognized nomenclaturally, especially as no two types occur sympatrically. Our decision not to name some or all of them is subjective and intuitive. On the other hand, we have recognized nomenclaturally other species of the yaeyamaensis species group of Ecsenius solely on the basis of color pattern. For instance E. yaeyamaensis and E. nalolo are distinguishable only on the basis of a slight but consistent difference in color pattern. In contrast to the color-pattern types of E. oculus, however, the two species occupy extensive, allopatric (parapatric?) ranges. The important issue, we believe, is not nomenclatural consistency but a description of the circumstances that will alert the reader and allow him to come to his own conclusions.

TABLE 7.—Comparison of certain aspects of the 6 color pattern types of Ecsenius oculus

REMARKS.—The three specimens collected by Longley (USNM 195716), which were hypothesized by Springer (1971:35, 42–43) to have come from the Banda Sea, have been re-examined. Although quite faded, they most closely resemble specimens with the type F color pattern. Also, they agree with type F specimens in meristics (Table 7). Our specimens from the Moluccas, the same area as that of Longley’s collection sites in the Banda Sea, have lower counts and a strikingly different color pattern. This evidence leads us to believe that Longley collected E. oculus in Samoa rather than in the Banda Sea as proposed by Springer. These are the only Longley specimens that we know of that appear to have come from Samoa.

FIGURE 8.—Variation in mean numbers of dentary incisor teeth of Ecsenius oculus as correlated with latitude (based on data in Table 9).

TABLE 8.—Color pattern types (CPT) and frequency distributions for certain characters in populations of Ecsenius oculus (see text for discussion of CPT)

Springer’s (1972) key differentiated E. oculus from other species of the yaeyamaensis species group by the presence of two dark basi-caudal spots, more than half orbital diameter, extending with little or no decrease in intensity well out on caudal fin. Among our new material, we find some E. opsifrontalis specimens with considerable dusky pigment on the caudal fin, and they could be misidentified as E. oculus using this character. However, E. oculus always has an obvious dark mark covering the proximal portion of the outermost segmented pelvic-fin ray and distal two-thirds of the remainder of the pelvic-fin, while E. opsifrontalis has uniformly pale pelvic fins. These two species are quite dissimilar in overall coloration (compare Figures 6 and 9).

Springer (1971, 1972) believed that E. oculus and E. opsifrontalis were a closely related species pair, and (1971) that their distributions were separated by the Andesite Line. All collections of E. oculus are from localities on or to the west of the Line, and all collections, except two, of E. opsifrontalis have been taken on or to the east of the Line (Figure 7). Both species occur together at only one locality, Fiji, where they were taken in the same collection. It is at Fiji where the most distinctive color patterns of E. opsifrontalis occur (see account of E. opsifrontalis). Although sympatric, E. opsifrontalis and E. oculus appear not to be syntopic. E. oculus occurs at the shallower ends of surge channels (“spur and groove” areas) of coral reefs while E. opsifrontalis tends to be found along vertical walls of the fore-reef, usually in small barren areas surrounded by coral growth (B. Carlson, personal communication). Thus the geographic distributions of E. oculus and E. opsifrontalis are known to coincide at Fiji where their habitat preferences appear to be different.

TABLE 9.—Color pattern types (CPT) and frequency distributions for number of dentary incisor teeth in populations of Ecsenius oculus (see text for discussion of CPT)

NEW MATERIAL (asterisk denotes previously reported localities; all others are new locality records).—RYUKYU ISLANDS. *Ishigaki, BPBM 8694 (1 specimen: 18.5). INDONESIA. JAVA SEA: Seribu Islands, USNM 211995 (19:14.5–33.7): Bawean Island, USNM 211975 (87:12.8–38.3). MOLUCCAS: Ambon Island, BPBM 18046 (1:27.1), 18048 (1:19.7), 18049 (1:28.2), 18052 (4:19.3–32.7): Saparua Island, USNM 211925 (7:13.2–17.0); Haruku Island USNM 209577 (7:13.1–48.7). SOLOMON ISLANDS. Savo Island, AM I.17490-001 (1:40.5). FIJI ISLANDS. *Viti Levu, BPBM 11270 (3:34.0–42.9), 14584 (2:45.0–46.4), USNM 214520 (3:31.8–37.8), 214787 (1:18.1); Moala Island, USNM 214788 (4:30.8–39.7). AMERICAN SAMOA. Tutuila Island, BPBM 17514 (5:22.0–36.8). WESTERN AUSTRALIA. North West Cape, WAM P25374-018 (9:26.3–33.4).

Ecsenius opsifrontalis Chapman and Schultz

FIGURE 9

Fifty-nine specimens of E. opsifrontalis have been obtained; all but two (from Guam) represent new locality records (see “New Material”). These specimens show that E. opsifrontalis varies geographically as well as individually in color pattern and meristics.

FIGURE 9.—Atoll, Gilbert Islands: b, BPBM 17958, female, 29.6 mm SL, Eniwetok; c, UG 1746 female 26 8 mm SL, Guam; d, ANSP 128013, male. 37.5 mm SL, Viti Levu, Fiji; e, USNM 211285, male, 35.3 mm SL, Kandavu, Fiji.

TABLE 10.—Frequency distributions for certain characters in populations of Ecsenius opsifrontalis

The basic color pattern of E. opsifrontalis is that of 2 stripes on the side of the body (a third is present along the dorsal body contour) crossed by a series of up to 10 bars that extend to the dorsal contour. Springer (1971) reported that these markings were usually faint and of equal intensity, but we find considerable individual and population variation in the intensity of their pigmentation. While some specimens from Oceania are faintly marked (Figure 9a and Springer, 1971, fig. 32), others have the bars more intensely pigmented than the stripes (Chapman and Schultz, 1952, fig. 93). A few specimens have the pigment most intense at the points where the bars cross the stripes, thus presenting the appearance of a series of dark spots with diffuse horizontal and vertical connections (Figure 9b–c). The specimens from Fiji (second record from west of the Andesite Line) have the most intense pigment and the most distinctive patterns (Figure 9d–e). In the Fijian specimens the points of intersection are darker and the connections less evident than in specimens from other localities.

Some specimens from Oceania have a considerable amount of dark dusky pigment extending out on the caudal fin, and, thus, could be misidentified as E. oculus in Springer’s (1972) revised key to Ecsenius. E. opsifrontalis, however, always has uniformly pale pelvic fins, whereas E. oculus has very darkly pigmented pelvic fins. These two species are known to occur together at only one locality, Fiji (see “Remarks” under E. oculus).

The Fijian forms of E. opsifrontalis superficially resemble E. collettei in color pattern but are easily distinguished. E. collettei lacks dark marks extending to the dorsal body contour and has a shorter lateral line and shorter nasal cirri than does E. opsifrontalis. In addition, E. collettei males have dark spots on the underside of the head, which are absent in E. opsifrontalis.

The frequency distributions for numbers of segmented dorsal-fin and anal-fin rays and caudal vertebrae (Table 10) in the Fijian population show a closer similarity to those of the Australian population (reported by Springer, 1972) than to those of other populations. The Australian specimens, however, do not show the intensification of pigment typical of the Fijian specimens.

NEW MATERIAL.—CAROLINE ISLANDS. Ulithi Atoll, BPBM 11487 (3 specimens:22.9–35.1). MARIANA ISLANDS. Guam, UG 1542 (1:28.3), 1746 (1:26.8); Saipan, UG 4892 (7:18.8–32.5), 5097 (3:27.2–35.5); Tinian, UG 5146 (16:15.9–29.5). MARSHALL ISLANDS. Eniwetok, BPBM 12896 (1:23.3), 17958 (1:29.6); Majuro Atoll, BPBM 17740 (1:25.2). GILBERT ISLANDS. Abiang Atoll, AM I.18044-004 (16:18.2–39.1). FIJI ISLANDS. Viti Levu, ANSP 128013 (4:33.2–39.3), 128014 (1:30.9), USNM 214521 (1:37.0); Kandavu, USNM 210562 (1:32.0), 211285 (2:34.2–35.2).

Ecsenius pictus, new species

FIGURE 10

DESCRIPTION (see Table 11 for frequencies of certain counts; other counts, with frequencies in parentheses, as follows).—Dorsal-fin spines 12(8); pectoral-fin rays 13(8, 1 specimen with 14 on right side only); segmented caudal-fin rays 13(5); dorsal procurrent caudal-fin rays 7(3), 8(1); ventral procurrent caudal-fin rays 6(2), 7(1), 8(1); total caudal-fin elements 26(2), 27(1), 29(1); pseudobranchial filaments 6(6); lower jaw (one side) posterior canine teeth 1(7); total lower jaw posterior canine teeth 1(1), 2(6); precaudal vertebrae 10(7); total vertebrae 31(3), 32(4). Lateral line with no paired pores, extending to below level of 9th(l), 10th(6), or 11th(1) dorsal-fin spine. Dorsal fin notched seven-ninths(6) length of first segmented dorsal-fin ray. Data for the holotype are given in Table 2.

FIGURE 10.—Ecsenius pictus, BPBM 15612, paratype, male, 41.6 mm SL, Alite Reef off Malaita, Solomon Islands. (Drawn by J. R. Schroeder.)

Color Pattern: In preservative, body dark brown with 7 or 8 pale pinstripes, mid-lateral stripe with series of 2–6 pale enlargements (spots) along its length; irregular row of numerous light spots dorsal to dorsalmost pinstripe; stripes and dorsal row of spots end posteriorly below level of 7th to 10th segmented dorsal-fin ray, anterior to area of alternating pale and dark irregular bars that extends to caudal-fin base. Mid-lateral stripe and second stripe above it extend anteriorly on head to posterior margin of orbit. Top of head and cheeks as dark as body or considerably lighter; underside of head less dusky than dorsal surface of head; light area present along lower lip. Pectoral-fin base with prominent light stripe; irregular light markings may be present dorsal to pectoral-fin base stripe. Pectoral and caudal fins with dusky pigment along rays, membranes immaculate. Anal fin dusky with indistinct light basal stripe. Dorsal fin dusky at base and along proximal half of rays; diffuse light stripe may interrupt dusky basal area. Pelvic fins unmarked 01 faintly pigmented proximally.

Color transparencies of specimens from the Moluccas and the Solomon Islands show the following: ground color chocolate-brown; pinstripes on head and body, and pectoral-fin base stripe brilliant white; area around mouth and underside of head yellow; light irregular areas posteriorly on body yellow interspersed with brown. Fins clear.

TABLE 11.—Frequency distributions for certain characters in populations of Ecsenius pictus

SEXUAL DIMORPHISM.—We found no significant differences in meristic or morphometric characters between the sexes.

GEOGRAPHICAL VARIATION.—Specimens from the Moluccas have lower means for counts of segmented dorsal-fin and anal-fin rays, caudal vertebrae, dentary teeth, epipleural ribs, and gill rakers than do specimens from the Solomon Islands (Table 11).

RELATIONSHIPS.—Ecsenius pictus belongs to the yaeyamaensis species group, which is characterized by a deeply notched dorsal fin and low counts of vertebrae and vertical fin rays. E. pictus does not easily fit into any of the three subgroups of the yaeyamaensis group; it does not have the reduced thirteenth dorsal-fin spine or long lateral line typical of the oculus subgroup, nor does it have the short nasal cirri characteristic of the prooculis subgroup. The pectoral-fin base of E. pictus has a distinct pale stripe, whereas the species of the yaeyamaensis subgroup have dark stripes or marks on the pectoral-fin base. The distinctive color pattern of E. pictus is also quite unlike that of any of the other species of the yaeyamaensis group. We, therefore, consider E. pictus to represent a fourth subgroup, the pictus subgroup, within the yaeyamaensis group of species.

REMARKS.—Ecsenius pictus is one of the deeper dwelling species of Ecsenius, having been taken (on coral reefs) at depths of 1–38 m; however, most specimens have come from depths greater than 10 m.

ETYMOLOGY.—The specific name, from the Latin pictus (painted or colored), refers to the unique and striking color pattern of this species.

HOLOTYPE.—USNM 213853, female 33.3 mm SL, Indonesia, Moluccas, just west of northernmost tip of Great Banda Island, reef and rock slope area, 04°30′30″S, 129°56′10″E, depth 10.7–18.3 m, collected by V. G. Springer and M. F. Gomon, VGS 74-11, 9 March 1974.

PARATYPES.—INDONESIA. MOLUCCAS: CAS 34245, male, 22.7 mm SL, and USNM 211895, female, 30.9 mm SL, collected with the holotype. USNM 210056, male, 17.7 mm SL, Saparua Island, off Kampungmaha, coral heads, depth 13.7–16.8 m, VGS 73-12, 17 January 1973. SOLOMON ISLANDS. BPBM 15612, male, 35.1 mm SL, Alite Reef off Malaita, lagoon reef in 12 m, collected by G. R. Allen and J. E. Randall, 24 July 1973. BPBM 15624, female, 35.1 mm SL, Alite Reef, outer reef slope in 38 m, J. E. Randall and B. Goldman, 25 July 1973. BPBM 15940, male, 37.3 mm SL, Alite Reef, from stomach of Lethrinus sp, collected by J. E. Randall, 25 July, 1973. BPBM 19016, male, 20.3 mm SL, Guadalcanal, 7 miles (11.7 km) W. of Honiara, reef in 1–2 m, J. E. Randall, G. R. Allen and M. McCoy, 11 July 1973.

Ecsenius prooculis Chapman and Schultz

Five additional specimens of E. prooculis have been obtained from New Britain, one of which is the only female known for this species. The female specimen has 15 segmented anal-fin rays (1 male with 15, 8 males with 16 anal-fin rays). The lateral line terminates below the tenth dorsal-fin spine (4 males with lateral line terminating below eighth, 5 males with lateral line terminating below ninth dorsal-fin spine). Additionally, where 7 males have concentrations of dark pigment on the underside of the head, the female and 2 males have pale areas devoid of pigment (see account of sexual dichromatism under “Color Pattern” in E. bandanus).

NEW MATERIAL.—New Britain: AM 17503-001 (1 specimen: 27.2), BPBM 15719 (4:23.0–36.0).

Ecsenius schroederi, new species

FIGURE 11

DESCRIPTION (certain counts and measurements for the holotype and only known specimen are given in Table 2; other data, with counts indicated for both sides, are as follows).—Gill rakers 14–14; pseudobranchial filaments 5–6; lower jaw posterior canine teeth 5–4. Lateral line with no paired pores.

Color Pattern: Head dusky, darker dorsally and anteriorly; distinct dusky postocular stripe extends from posterior margin of orbit to upper edge of opercle; an indistinct, thin dusky stripe extends across lower cheek onto opercle at level of mid-base of pectoral fin. Body pale with three wavy, dark pinstripes; dorsalmost pinstripe extends from supra-temporal canal posteriorly to base of twelfth segmented dorsal-fin ray; mid-lateral pinstripe is a continuation of postocular head stripe and extends posteriorly to caudal-fin base; ventralmost pinstripe extends from upper pectoral-fin insertion posteriorly to caudal-fin base. Three or four indistinct, pale spots present between mid-lateral and ventral pinstripes. Dorsal fin with concentration of dusky pigment along base, indistinct dusky stripe proximally, and dusky pigment extending outward along rays. Anal fin with dusky stripe on outer third of fin, tips of rays pale. Caudal fin with scattered dusky pigment proximally and along rays, interradial membranes clear. Pectoral fin with scattered dark pigment proximally; tips of five lowermost rays dark. Pelvic fin with scattered dusky pigment distally, otherwise unmarked.

FIGURE 11.—Ecsenius schroederi, USNM 209743, holotype, male, 24.0 mm SL, Ambon Island, Moluccas. (Drawn by J. R. Schroeder.)

RELATIONSHIPS.—Ecsenius schroederi and E. mandibularis are the only species of Ecsenius with more than two posterior canine teeth on each side of the lower jaw. E. schroederi can be distinguished from E. mandibularis by its very different color pattern and by having fewer segmented dorsal-fin and anal-fin rays (see “Key to the Species of Ecsenius”).

ETYMOLOGY.—Named for Jack R. Schroeder, whose excellent illustrations have so greatly enhanced the usefulness of our blennioid studies.

HOLOTYPE.—USNM 209743, male, 24.0 mm SL, Indonesia, Moluccas, Ambon Island, off Tandjung Suli, shallow coral reef, depth to 2.4 m, collected by V. G. Springer and M. F. Gomon, VGS 73-8, 11 January 1973.

Ecsenius stigmatura Fowler

New material (and records) of E. stigmatura expands the ranges of meristics for this species as given by Springer (1971). The ranges are as follows: segmented anal-fin rays 17–19; dorsal procurrent caudal-fin rays 6–8; ventral procurrent caudal-fin rays 5–7; total caudal-fin elements 26–28; gill rakers 13–17; pseudobranchial filaments 5–6; dentary incisor teeth 34–40; epipleural ribs 12–13; lateral line extending posteriorly to below level of 7th to 9th dorsal-fin spine.

A color transparency taken shortly after the capture of a specimen of E. stigmatura shows color similar to that described by Springer (1971), which was taken from a color sketch of the holotype. However, the transparency shows the body to be dusky gray anteriorly and bright orange-red posteriorly. The orange opercular stripe is prominent but does not extend anteriorly on the cheek. The black spot on the caudal peduncle is bordered anteriorly by a white semicircle.

REMARKS.—The specimen from Darvel Bay, Borneo, questionably assigned to E. stigmatura by Springer (1971), has been found to represent a new species, E. melarchus. Comparisons of E. melarchus and E. stigmatura are given in Table 5 and under “Relationships” for E. melarchus.

NEW MATERIAL—INDONESIA. MOLUCCAS: Ambon Island, BPBM 18613 (I specimen:25.1), USNM 210445 (1:42.8); Ceram Island, USNM 209696 (2:19.6–21.8), 209851 (12:16.4–315); Saparua Island, USNM 209993 (1:24.2), 210146 (5:26.2–36.6).

Ecsenius trilineatus Springer

Since the description of E. trilineatus (Springer, 1972), 31 additional specimens representing new locality records have been obtained (see “New Material”). Springer (1971) had available only one specimen of E. trilineatus, which he questionably identified as E. yaeyamaensis and hypothesized (1971, “Appendix”) as having come from the Banda Sea. Our new material includes the first definite records of E. trilineatus from Banda Sea localities.

TABLE 12.—Frequency distributions for certain characters in populations of Ecsenius trilineatus

The new material expands the ranges for the meristics of certain characters. Some of these characters, which hint at possible geographic variation, are given in Table 12; other characters are as follows: pectoral-fin rays 12–14 (3 counts of 12 recorded); dorsal procurrent caudal-fin rays 6–9; ventral procurrent caudal-fin rays 6–8; total caudal-fin elements 26–30; epipleural ribs 11–13. The depth of the dorsal-fin notch of the holotype was printed erroneously as 0 (Springer, 1972, table 1); the correct figure is 9 (nine-ninths the length of the first segmented dorsal-fin ray).

REMARKS.—The single specimen from Efate, New Hebrides (BPBM 12122), tentatively identified as E. trilineatus by Springer (1972), has proven to be of a new species, E. isos. Comparisons of this new species with E. trilineatus are given in the account of E. isos.

NEW MATERIAL.—INDONESIA. MOLUCCAS: Saparua Island, USNM 211926 (17 specimens: 14.0–27.2); Banda Islands, Goenoeng (= Gunung) Api Island, USNM 211930 (5:13.8–25.6), Naira Island, USNM 211941 (1:14.8), Great Banda Island, USNM 211945 (7:21.5–27.1). SOLOMON ISLANDS. Florida Island, AM I.17497-001 (1:26.0).

Ecsenius yaeyamaensis (Aoyagi)

New locality records for E. yaeyamaensis are given in “New Material.” All specimens in the list, including the Western Australian specimen, do not have the fine dark spots posteriorly on the body that are typical of specimens from the Great Barrier Reef, but instead have the common color pattern.

Springer (1971) reported three specimens of E. yaeyamaensis from New Hebrides with variant color patterns similar to that of the species he subsequently described as E. trilineatus. The recent collection of normally patterned E. yaeyamaensis from New Hebrides has prompted us to reassess the identification of the earlier specimens, and we find them to represent a new species, E. isos (see “Relationships” for E. isos).

NEW MATERIAL.—WESTERN AUSTRALIA. Dampier Archipelago, WAM P25107-021. INDONESIA. JAVA SEA: Seribu Islands, USNM 211994; Karimundjawa Islands, USNM 211981; Bawean Island, USNM 211974. CELEBES: Boeton (= Butung) Island, USNM 211967. MOLUCCAS: Ambon Island, BPBM 18051, USNM 209735; Haruku Island, USNM 209603; Saparua Island, USNM 211924. SOLOMON ISLANDS. Guadalcanal, AM I.17486-002; Florida Island, AM I.17497-002. NEW HEBRIDES. Efate Island, USNM 214819, BPBM 19577.

Key to the Species of Ecsenius

Literature Cited

Chapman, W. M., and L. P. Schultz

1952. Review of the Fishes of the Blennioid Genus Ecsenius, with Descriptions of Five New Species. Proceedings of the United States National Museum, 102(3310):507–528.

Masuda, H., C. Araga, and T. Yoshino

1975. Coastal Fishes of Southern Japan. 379 pages. Tokai University Press.

Smith-Vaniz, W. F., and V. G. Springer

1971. Synopsis of the Tribe Salariini, with Description of Five New Genera and Three New Species (Pisces: Blenniidae). Smithsonian Contributions to Zoology, 71:1–72.

Springer, V. G.

1968. Osteology and Classification of the Fishes of the Family Blenniidae. United States National Museum Bulletin, 284:1–85.

1971. Revision of the Fish Genus Ecsenius (Blenniidae, Blenniinae, Salariini). Smithsonian Contributions to Zoology, 72:1–74.

1972. Additions to Revisions of the Blenniid Fish Genera Ecsenius and Entomacrodus, with Descriptions of Three New Species of Ecsenius. Smithsonian Contributions to Zoology, 134:1–13.

Springer, V. G., and M. F. Gomon

1975. Revision of the Blenniid Fish Genus Omobranchus with Descriptions of Three New Species and Notes on Other Species of the Tribe Omobranchini. Smithsonian Contributions to Zoology, 177:1–135.

Thorne, R. F.

1963. Biotic Distribution Patterns in the Tropical Pacific. Pages 311–350 in J. L. Gressitt, editor, Pacific Basin Biogeography. Honolulu: Bishop Museum Press.

REQUIREMENTS FOR SMITHSONIAN SERIES PUBLICATION

Manuscripts intended for series publication receive substantive review within their originating Smithsonian museums or offices and are submitted to the Smithsonian Institution Press with approval of the appropriate museum authority on Form SI–36. Requests for special treatment—use of color, foldouts, casebound covers, etc.—require, on the same form, the added approval of designated committees or museum directors.

Review of manuscripts and art by the Press for requirements of series format and style, completeness and clarity of copy, and arrangement of all material, as outlined below, will govern, within the judgment of the Press, acceptance or rejection of the manuscripts and art.

Copy must be typewritten, double-spaced, on one side of standard white bond paper, with 1¼″ margins, submitted as ribbon copy (not carbon or xerox), in loose sheets (not stapled or bound), and accompanied by original art. Minimum acceptable length is 30 pages.

Front matter (preceding the text) should include: title page with only title and author and no other information, abstract page with author/title/series/etc., following the established format, table of contents with indents reflecting the heads and structure of the paper.

First page of text should carry the title and author at the top of the page and an unnumbered footnote at the bottom consisting of author's name and professional mailing address.

Center heads of whatever level should be typed with initial caps of major words, with extra space above and below the head, but with no other preparation (such as all caps or underline). Run-in paragraph heads should use period/dashes or colons as necessary.

Tabulations within text (lists of data, often in parallel columns) can be typed on the text page where they occur, but they should not contain rules or formal, numbered table heads.

Formal tables (numbered, with table heads, boxheads, stubs, rules) should be submitted as camera copy, but the author must contact the series section of the Press for editorial attention and preparation assistance before final typing of this matter.

Taxonomic keys in natural history papers should use the alined-couplet form in the zoology and paleobiology series and the multi-level indent form in the botany series. If cross-referencing is required between key and text, do not include page references within the key, but number the keyed-out taxa with their corresponding heads in the text.

Synonymy in the zoology and paleobiology series must use the short form (taxon, author, year:page), with a full reference at the end of the paper under “Literature Cited.” For the botany series, the long form (taxon, author, abbreviated journal or book title, volume, page, year, with no reference in the “Literature Cited”) is optional.

Footnotes, when few in number, whether annotative or bibliographic, should be typed at the bottom of the text page on which the reference occurs. Extensive notes must appear at the end of the text in a notes section. If bibliographic footnotes are required, use the short form (author/brief title/page) with the full reference in the bibliography.

Text-reference system (author/year/page within the text, with the full reference in a “Literature Cited” at the end of the text) must be used in place of bibliographic footnotes in all scientific series and is strongly recommended in the history and technology series: “(Jones, 1910:122)” or “… Jones (1910:122).”

Bibliography, depending upon use, is termed “References,” “Selected References,” or “Literature Cited.” Spell out book, journal, and article titles, using initial caps in all major words. For capitalization of titles in foreign languages, follow the national practice of each language. Underline (for italics) book and journal titles. Use the colon-parentheses system for volume/number/page citations: “10(2):5–9.” For alinement and arrangement of elements, follow the format of the series for which the manuscript is intended.

Legends for illustrations must not be attached to the art nor included within the text but must be submitted at the end of the manuscript—with as many legends typed, double-spaced, to a page as convenient.

Illustrations must not be included within the manuscript but must be submitted separately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.) can be intermixed throughout the printed text. They should be termed Figures and should be numbered consecutively. If several “figures” are treated as components of a single larger figure, they should be designated by lowercase italic letters (underlined in copy) on the illustration, in the legend, and in text references: “Figure 9b.” If illustrations are intended to be printed separately on coated stock following the text, they should be termed Plates and any components should be lettered as in figures: “Plate 9b.” Keys to any symbols within an illustration should appear on the art and not in the legend.

A few points of style: (1) Do not use periods after such abbreviations as “mm, ft. yds, USNM, NNE, AM, BC.” (2) Use hyphens in spelled cut fractions: “two-thirds.” (3) Spell out numbers “one” through “nine” in expository text, but use numerals in all other cases if possible. (4) Use the metric system of measurement, where possible, instead of the English system. (5) Use the decimal system, where possible, in place of fractions. (6) Use day/month/year sequence for dates: “9 April 1976.” (7) For months in tabular listings or data sections, use three-letter abbreviations with no periods: “Jan, Mar, Jun,” etc.

Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front matter and ALL legends, etc., at the back of the text—in the following order: (1) title page, (2) abstract, (3) table of contents. (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.