Protoblepharon Rosenblatti, a New Genus And

Home , Anomalopidae, Kryptophanaron, Monocentridae, Photoblepharon

10 October 1997 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1l0(3):373-383. 1997. Protoblepharon rosenblatti, a new genus and species of flashlight fish (Beryciformes: Anomalopidae) from the tropical South Pacific, with comments on anomalopid phylogeny

Carole C. Baldwin, G. David Johnson, and John R. Paxton (CCB, GDJ) Department of Vertebrate Zoology, MRC 159, National Museum of Natural History, Smithsonian Institution, Washington, DC. 20560, U.S.A.; (JRP) Division of Vertebrate Zoology, The Australian Museum, Sydney, New South Wales, Australia

Abstract. — Protoblepharon rosenblatti is described from a single large specimen collected at 274 m off Rarotonga, Cook Islands. It differs from other anomalopids most notably in having a low number of gill rakers on the first arch (21 vs. 24 or more), high number of body scale rows (ca. 145 vs. 130 or fewer), no postorbital papillae, and a very small gap between the lacrimal and nasal for passage of the fibrocartilaginous stalk, which is twisted and not broadly exposed posteriorly. Protoblepharon is a primitive member of the lineage of flashlight fishes characterized by a shutter mechanism for light-organ occlusion.

The Anomalopidae comprise a small Phthanophaneron + Kryptophanaron + group of nearly circumtropically distribut- Photoblepharon clade. ed, marine beryciform fishes characterized We have examined a very large (229 mm most conspicuously by a subocular lumi- SL) flashlight fish from the preserved col- nous organ in which symbiotic luminous lections at the Australian Museum that can- bacteria are cultured (e.g., Harvey 1922, not be assigned to any known species. The Haygood & Cohn 1986). Since a review of specimen was collected on hook and line in the family by McCosker & Rosenblatt deep water off Rarotonga, Cook Islands. A (1987), in which five species were recog- comparison of the single known specimen nized in three genera, Johnson & Rosenblatt of the new species with other anomalopids (1988) erected a new genus, Phthanophan- suggests that it is a primitive member of the eron, for the eastern Pacific Kryptophan- group of flashlight fishes characterized by a aron harveyi Rosenblatt & Montgomery, shutter mechanism for light-organ occlu- and Rosenblatt & Johnson (1991) described sion, i.e., all anomalopid genera except An- a new genus and species from Tahiti, Par- omalops (Johnson & Rosenblatt 1988). Our mops coruscans. With the exception of phylogenetic placement of the new species Photoblepharon, with two species, all gen- is best served by erecting a new genus for era are monotypic. it. Using derived morphological features, Flashlight fishes observed by divers and including aspects of the occlusion mecha- those preserved in fish collections typically nism of the light organ, Johnson & Rosen- are small (< 100 mm SL), and thus the large blatt (1988) hypothesized the following re- size of the holotype of the new species is lationships (in phyletic sequence) among unusual. However, several large specimens four anomalopid genera: Anomalops, (>200 mm SL) are known for Anomalops Phthanophaneron, Kryptophanaron, Pho- (McCosker & Rosenblatt 1987), and toblepharon. Rosenblatt & Johnson (1991) Phthanophaneron harveyi is known from a placed Parmops as the sister group of the 20-mm SL juvenile, the 67.7-mm SL ho- 374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Iotype, and a 204-mm SL specimen (Ro- dae hypothesis, which also is based on a senblatt & Montgomery 1976, McCosker & single character: presence of a separate spi- Rosenblatt 1987, Allen & Robertson 1994). nous dorsal fin in monocentrids and all an- Large flashlight fishes typically are taken in omalopids except Photoblepharon. Recog- deeper water than small ones (e.g., Mc- nizing that neither hypothesis of trachich- Cosker & Rosenblatt 1987), and may be thyoid interrelationships is well supported, more common than their poor representa- we initially included the sister group of tion in fish collections suggests. Further trachichthyoids, the Anoplogastridae + Di- collecting efforts are needed to find small retmidae (Zehren 1979, Moore 1993, Bald- specimens of the new species, the holotype win & Johnson 1995), as an additional out- of which is from an area of the Cook Is- group in our analysis, but this inclusion had lands where small specimens of both An- no effect on the topology of the tree and omalops and Photoblepharon occur. Our will not be discussed further. All characters purposes here are to describe the new spe- included in the phylogenetic analysis were cies and discuss its relationships in the con- weighted equally, and multistate characters text of an existing phylogenetic hypothesis were treated as unordered. of anomalopid genera. Protoblepharon, new genus Methods Diagnosis.—An anomalopid with 21 gill Measurements were made with needle- rakers on the first arch, about 145 body point dial calipers or an ocular micrometer scale rows, a small (14.5% HL) rotatable to the nearest 0.1 mm. Terminology of light organ, no postorbital papillae, and a structures associated with the light organ very small gap between the lacrimal and follows Johnson & Rosenblatt (1988). Gill- nasal for passage of the stalk, which is raker counts include all rudiments. Scale twisted and not broadly exposed posterior- bars in illustrations represent 1 mm. We fol- ly. The following combination of characters low Johnson & Rosenblatt (1988) in using also is useful in distinguishing Protobleph- the name "Trachichthyoidei" for the clade aron: a separate spinous dorsal fin, a pel- comprising the Anomalopidae, Monocentri- vic-fin spine, two anal-fin spines, 59—60 en- dae, and Trachichthyidae, the monophyly of larged lateral-line scales, an elastic shutter, which was proposed by Zehren (1979) and and a broad strap-like ethmomaxillary lig- Moore (1993). The Monocentridae and ament with no groove or swelling. Trachichthyidae were considered the first Type species. — Protoblepharon rosen- and second outgroups, respectively, for the blatti, new species. phylogenetic analysis of anomalopid genera Etymology. — From the Greek protos, based on the hypothesized monophyly of first, and blepharon, eyelid, in reference to the Trachichthyoidei and a proposed sister- the cladistic position of the genus as first in group relationship between the Monocen- the lineage of flashlight fishes that occlude tridae and Anomalopidae (Konishi & Oki- the light organ with an erectable shutter. yama 1997). Following Zehren (1979), Moore (1993) suggested a sister-group re- Protoblepharon rosenblatti, new species lationship between the Monocentridae and Fig. 1 Trachichthyidae, but the single character cited as evidence involves the infraorbital Holotype.—AMS 1.24275-001. A series, which is modified in all anomalopids 229-mm SL female specimen caught by to accommodate the light organ. Further hook and line northwest of Matavera, Rar- study is needed to test Konishi & Okiya- otonga, Cook Islands (21°12'S, 159°45'W), ma's (1997) Anomalopidae + Monocentri- at 274 m on 30 Nov. 1983, and donated by VOLUME 110, NUMBER 3 375

N. Sims of the Cook Islands Fisheries De- pillae. Distinct notch at symphysis of pre- partment. maxillae, presumably accommodating small Description. — Counts and measure- dentigerous knobs at symphysis of dentaries ments, in mm, of the holotype: Dorsal-fin when mouth closed. Premaxillae, including rays VII, 14; anal-fin rays II, 11; pectoral- most of lateral and medial surfaces, covered fin rays iil5i; pelvic-fin rays 1,5; caudal-fin with bands of villiform teeth; no teeth at rays 10, 10+9, 9 (all procurrent rays spi- symphyseal notch. Each dentary with nar- nous except the posteriormost in the upper row band of villiform teeth posteriorly, and lower caudal-fin lobes); branchiostegals patch of slightly larger teeth near symphy- 8; gill rakers on first arch 21 (5+ 12 rakers, sis extending onto lateral and medial sur- plus two flat plates at dorsal end of epi- faces. Vomer edentulous, palatines with branchial and two at anterior end of cera- well-developed bands of villiform teeth. tobranchial); pored lateral-line scales 59 (60 Bones of head and pectoral girdle cov- on right side); scale rows above lateral line ered with numerous, rugose to minutely ser- ca. 18; abdominal scutes 9; vertebrae rate ridges. Cleithrum with large exposed 14+16. Head length 83.4; predorsal length surface posteriorly, margin smooth. Supra- 96.9; prepelvic length 112; body depth at cleithrum almost completely beneath oper- origin of dorsal fin 81.9; caudal-peduncle cle, only posterodorsal corner exposed, depth 23.3; caudal-peduncle length 50.5; margin smooth. Anterior infraorbitals en- snout length 23.7; eye diameter 19.4; orbit larged, covering anteroventral corner of or- diameter 20.5; light-organ length 12.1; pec- bit, and slightly flared laterally forming a toral-fin length 51.9; pelvic-fin length 41.2; medially sloping plate. Laterosensory ca- first dorsal-spine length 12.2; third dorsal- nals of head appearing as channels of dark spine length 17.0; sixth dorsal-spine length skin surrounded by bone, skin covered with 6.8; seventh dorsal-spine length 20.0; first small black papillae and perforated fre- anal-spine length 5.8; second anal-spine quently by pores. length 12.3. Eye small, diameter 4.1 in head. No Body compressed (width 1.9 in depth) fleshy papillae on posterior rim of orbit. Lu- and deep (depth at origin of dorsal fin 1.8 minous organ below eye small, length 6.9 in length without head). With mouth open, in head. Light organ free posteriorly, sup- profile sloping gradually from occiput to ported by fibrocartilaginous cup anteriorly, snout, somewhat convex in region of mes- which is ligamentously bound to a fibro- ethmoid, then dropping slightly to symphy- cartilaginous stalk. Organ capable of being sis of upper jaw; upper-jaw symphysis at rotated downward into pocket below eye level of horizontal through middle of eye. and medial to infraorbitals. When occluded, Nostrils anterior and completely dorsal to dorsal margin of light organ well below in- eye with mouth open, the anterior with fraorbital rim. Black elastic shutter mem- thickened posterior rim. With jaws forced brane attached along outer margin of sub- closed, mouth oblique, lower jaw originat- orbital pocket. ing anteriorly near horizontal through mid- Numerous small, spinoid scales (cf. Rob- dle of eye, and maxilla extending posteri- erts 1993, = Cf of Johnson 1984) covering orly to vertical through middle of eye. Pos- body, about 145 lateral body rows but dif- terior supramaxilla ovoid, anterodorsal sur- ficult to count because of irregular distri- face with small pointed process extending bution of scales. Lateral line covered by en- anteriorly along posterodorsal edge of small larged scales, and abdomen with series of anterior supramaxilla. Posterior supramax- about 9 enlarged, keeled scutes. Head most- illa covering most of posterior portion of ly scaleless, a few thick, heavily sculptured maxilla, the posteroventral corner of max- scales at anterodorsal corner of opercle, illa exposed and covered with tiny black pa- scales coalescing on cheek to form strong 376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Holotype of Protoblepharon rosenblatti, AMS 1.24275-001, 229 mm SL. Photograph by C. Bento, AMS. bony covering. Gular region naked, with supporting the light organ is connected to a low, pigmented, transverse ridges. fibrocartilaginous stalk that is continuous Most rakers on first gill arch long and with its contralateral member across the lath-like, length of first raker below angle snout, with no attenuation at the commis- nearly two-thirds diameter of eye. Four an- sure. Anteriorly, the stalk lies flat against teriormost and two dorsalmost rakers on the snout, with the broad surface facing out- first arch and rakers on remaining arches ward; posteriorly, where the stalk passes less than half that long. Pseudobranch well through a small gap between the lacrimal developed, about 27 filaments. and nasal, it twists such that its broad sur- Spinous dorsal fin lower than soft dorsal face lies nearly in the horizontal plane (Fig. fin, the longest spine (seventh) 1.7 in length 2B). The posteroventral portion of the stalk of longest unbroken soft ray (the fifth). and anteroventral corner of the cup each Length of first spine 1.2 in second, second terminate in a short ventral process, and 1.1 in third and fourth, fifth 1.3 in first, these are loosely joined dorsally by a short sixth 1.7 in first, and seventh 0.6 in first. ligament. There is no ventral stalk hook. First anal spine shorter than any dorsal The shutter is slightly thickened near its an- spine, second anal spine about equal in terodorsal corner, but there is no discrete length to first dorsal spine. Longest anal shutter knob. The cup supports the anterior soft ray (second) about equal to longest dor- end of the light organ and extends posteri- sal soft ray. Caudal fin deeply forked. orly along about two-thirds of its ventral Pectoral-fin base nearly horizontal, fin di- surface (Fig. 2B). A large section of it is rected posterodorsally. Pectoral-fin length exposed anterolaterally between the light about 1.6 in head, third through sixth rays organ and stalk; anteromedially, the cup the longest. Pelvic fin shorter than pectoral, forms a medially projecting shelf. The stalk 2.0 in head, and falling well short of anus. muscle is not differentiated into dorsal and Color in alcohol dark brown to black; fins, ventral bundles and inserts on the ligament posterior trunk, and cheek darkest. Lateral- connecting the stalk and cup. The Ligament line scales pale. of Diogenes originates on the rostral carti- Occlusion mechanism (Fig. 2).—The cup lage, curves around the ethmomaxillary lig- VOLUME 110. NUMBER 3 377 ament as a broad strap, and then narrows stalk hook. These features are important posteriorly before inserting broadly on the in the erection of the shutter in Phthano- ventral process and anteromedial shelf of phaneron, Kryptophanaron, and Photo- the cup. The ethmomaxillary ligament de- blepharon (Johnson & Rosenblatt 1988), scends anteroventrally from its origin on and it is thus unclear how the shutter the mesethmoid to insert on the maxilla mechanism operates in either Parmops or with a short branch to the palatine. This lig- Protoblepharon. Protoblepharon lacks ament is broad and flat, lacks a groove the primary diagnostic feature of Par- where it is crossed by the Ligament of Di- mops, expansion of the first four infraor- ogenes, has no pronounced forward flexure, bital bones to form a medially sloping and bears no swellings. shelf that protrudes laterally well beyond Etymology.— It is our pleasure to name the margin of the orbit (Rosenblatt & this species in honor of Dr. Richard H. Ro- Johnson 1991). It shares with Parmops, senblatt, a mentor to one of us (GDJ), Phthanophaneron, Kryptophanaron, and friend and valuable colleague to all of us. Photoblepharon many features recognized His contributions to the systematics and as synapomorphies of that lineage by functional morphology of flashlight fishes Johnson & Rosenblatt (1988): two supra- have shed much light on the evolution and maxillae, transverse ridges on the gular biology of the Anomalopidae. isthmus, large v-shaped lateral dentary Remarks. — States for P. rosenblatti of tooth patches, over 100 lateral body scale characters not associated with the light or- rows, reflective lateral-line scales, an gan used by Johnson & Rosenblatt (1988) erectable (or at least elastic) shutter, and and Rosenblatt & Johnson (1991) to recon- a stalk that is continuous across the snout. struct the evolutionary history of anomal- It lacks another derived feature of that opid genera are given in Table 1, those as- group, a groove in the ethmomaxillary sociated with the light organ in Table 2. In ligament (secondarily absent in Phthano- Table 3, we list states of characters among phaneron). It is thus most parsimonious to anomalopids and outgroups not considered hypothesize that Protoblepharon is the in previous publications that are useful in sister group of Parmops + Phthanophan- diagnosing P. rosenblatti: number and rel- eron + Kryptophanaron + Photoblephar- ative size of lateral-line scales, number of on. A previously undescribed character gill rakers, relative size of light organ, least corroborating the placement of Proto- distance between nasal and lacrimal, and blepharon below Parmops involves the orientation of fibrocartilaginous stalk. The configuration of the lacrimal, nasal, and relatively small size of the light organ of P. stalk. In most anomalopids, the stalk is rosenblatti could be a function of the large broadly exposed at the commissure (Figs. size of the holotype, as comparisons with 2B, C; 3B, C), and there is no attenuation Phthanophaneron haweyi indicate an in- where it joins its contralateral member. In verse relationship between relative size of Protoblepharon, the lacrimal and nasal light organ and body (31.2% HL in the are separated by only a small gap (least 67.7-mm SL holotype and 22.7% HL in the distance between them ca. 1.6% HL; Fig. 204-mm SL specimen). McCosker (1982) 3B), and the stalk twists before passing noted a similar trend in Kryptophanaron al- through this gap such that the broad sur- fredi (44.7% HL in a 25-mm SL specimen, face is nearly in the horizontal plane, and 34.9% HL in an 89-mm SL fish). the narrow margin formerly in a ventral Relationships.— Protoblepharon is position is exposed laterally (Fig. 2B). In most similar to Parmops in the primitive the Parmops clade, the lacrimal is sepa- nature of the occlusion mechanism, es- rated from the nasal by a large gap (4.0% pecially the absence of a shutter knob and HL or more, Fig. 3C), and the broad sur- 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Light organ and associated structures in (A) Anomalops katoptron. USNM 293340, (B) Proto- blepharon rosenblatti, AMS 1.24275-001, (C) Kryptophanaron alfredi, USNM 343635. Right side. face of the stalk is exposed along its entire blepharon. A small gap is autapomorphic length (Figs. 2C, 3C). Anomalops also has for the Anomalopidae. Although both a very small gap between the lacrimal and have small gaps, the conditions in An- nasal (ca. 1.0% HL, Fig. 3A), and the two omalops (figs. 2A, 3A) and Protoblephar- bones are in contact with one another in on (figs. 2B, 3B) are different in that the the outgroups. A large gap between the stalk is never broadly exposed in Anom- lacrimal and nasal is thus an additional alops (it is attenuated at the commissure) synapomorphy of Parmops, Phthan- and does not twist posteriorly but nearly ophaneron, Kryptophanaron, and Photo- disappears from view in the region of the VOLUME 110, NUMBER 3

Table 1.—States in Protoblepharon rosenblatti of which is probably smaller in Proto- Johnson & Rosenblatt's (1988) characters not associ- blepharon than in the Parmops lineage ated with the light-organ complex. because of the closer association of the 1. Vertebrae bearing epineurals* 1, 2, 10-14 lacrimal and nasal (and hence the lateral 2. Branchiostegals spiny ethmoid). Additional specimens of Proto- 3. Openings in pars jugularis ? blepharon that can be cleared and stained 4. Parasphenoid flanges ? are needed to examine osteology. 5. Swimbladder stay A cladogram depicting relationships 6. Postorbital papillae 0 7. Cephalic sensory canal covering Papillose among the six anomalopid genera is shown 8. Lateral-line tubes Closed in Figure 4. We constructed the tree using 9. Midventral scutes Continuous the Branch and Bound option of Swofford's 10. Dorsal Fin VII, 14 (1991) PAUP 3.0 with the matrix in Table Supraneurals 0/0/1 + 11 11. Supramaxillae 2 4. All of the characters (Tables 1, 2) used 12. Transverse ridges on gular isthmus + by Johnson & Rosenblatt (1988) and Ro- 13. Lateral dentary tooth patch Large "V" senblatt & Johnson (1991) initially were in- 14. Body scale rows ca. 145 cluded in the analysis, but many are not in- 15. Reflective or transparent lateral- line scales + formative and were eliminated from the 16. Pelvic spine + matrix. Johnson & Rosenblatt (1988) and 17. Anal spines II Rosenblatt & Johnson (1991) included all 18. Vertebrae 14 + 16 characters in Tables 1 and 2 on their dado- 19. Comer of maxilla Papillae grams, but noted that many features asso- Epipleurals of Johnson & Rosenblatt (1 ciated with the light organ could not be po- epineurals (Patterson & Johnson 1995). larized by outgroup comparison because of the absence of comparable conditions in the small lacrimal/nasal gap as it turns abrupt- outgroups. They surmised that the fully ro- ly ventrad (Johnson & Rosenblatt 1988). tatable light organ of Anomalops and the The different associations between the complex shutter mechanism of the non-ro- lacrimal and nasal possibly are also re- tatable light organ of Photoblepharon rep- flected in the morphology of the C-shaped resent highly specialized conditions within process of the lacrimal (Zehren 1979), the family, and interpreted the less special-

Table 2.—States in Protoblepharon rosenblatti of Johnson & Rosenblatt's (1 characters associated with the light organ.

I Attachment of Ligament of Diogenes on cup Medial II Attachment of Ligament of Diogenes anteriorly Rostral Cartilage III Cup with medial shelf Moderate IV Insertion of stalk muscle dorsally Ligament V Stalk with inward flexure at cup articulation VI Rotation pad VII Postocular skin flap Villa Erectile shutter Vlllb Shutter knob rx Stalk hook X Stalk continuous across snout XIa Ethmomaxillary ligament with groove Xlb Ethmomaxillary ligament with medial swelling XII Hook and shutter knob intimately associated NA XIII Cup process attached to stalk hook by ligament NA XIV Organ rotatable + 380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 3.—Comparison of some diagnostic features of Protoblepharon rosenblatti among anomalopids and other trachichthyoids. Data are from the literature or specimens examined in this study (single values represent counts or measurements of a single individual). For light organ length and least distance between nasal and lacrimal, range of values given only for taxa where data from a wide range of adult sizes was available. HL = head length, ant. = anteriorly, NA = not applicable.

Least distance between Light organ nasal & Gill rakers length lacrimal Taxon Lateral-line scales on 1st arch <% HL) (% HL) Orientation of stalk

Anomalops 57-59 28-32' 35.4 1.0 Not broadly exposed (Not enlarged) Protoblepharon 59-60 21" 14.5 1.6 Broadly exposed ant., (Enlarged) twisted posteriorly Parmops 30 30 35.6 4.8 Broadly exposed (Enlarged) Phthanophaneron 38 22-24" 22.7-31.2 4.0-4.6 Broadly exposed (Enlarged) Kryptophanaron 32-34' 24-28' 36.3^4.7" 5.7 Broadly exposed (Enlarged) Photoblepharon 39 25-30' 48.6 5.6 Broadly exposed (Enlarged) Monocentridae 12-15' 19-21' NA 0 NA (Not enlarged) Trachichthyidae 25-63» 15^4" NA 0 NA (Enlarged in some)

"Shimizu (1984). this study. h Counts include flat plates at dorsal end of epibranchial, anterior end of ceratobranchial, or both. c Colin et al. (1979), this study. ''McCosker (1982). 'Abe & Haneda (1973). ' Smith (1986). sGomon (1994). h Kotlyar (1980).

NASAL

LACRIMAL

Fig. 3. Diagrammatic illustrations of the anterior snout region in three anomalopids, showing different con- figurations of the nasal, lacrimal, and fibrocartilaginous stalk. (A) Anomalops katoptron, USNM 293340, (B) Protoblepharon rosenblatti, AMS 1.24275-001, (C) Kryptophanaron alfredi, USNM 343635. Dotted line in Anomalops shows anterior extent of skin covering stalk, that in Protoblepharon represents region where stalk is twisted. VOLUME 110, NUMBER 3 381

• J0<" /

_ _ Hv Elhmomaxillary ligament with swelling

"13,: Shutter knob present 14,: stalk hook present

4,: One postorbltal papilla 17,: Large gap between lacrimal & nasal 15.*:Ethmomaxluary ligament with groove 18,: 15 or note dorsal rays 5,: 2 Supramaxlllae 6,: Gular ridges present 7,: Large V-shaped dentary tooth patch &,: >100 body scale rows 9,: Pale Lateral-he scales 1,: Light organ present 2,: Neural & haemal spines support procurrent m p 3,: Interarcual cartilage present 17,: Lacrimal & nasal separate but close

Fig. 4. Cladogram showing hypothesized relationships among anomalopid genera, = ambiguous character resolved using ACCTRAN. Treelength = 29 CI = 0.83 RI = 0.83. ized states in Parmops and Phthanophan- the ethmomaxillary ligament, is ancestral. eron as ancestral for the family. They po- Relative to outgroups Monocentridae and larized features of the light organ based on Trachichthyidae, other features of Proto- these assumptions. blepharon that might corroborate a Proto- The PAUP analysis did not make such blepharon-like anomalopid ancestor include assumptions, and recognized that the occlu- the small size of the light organ (14.5% HL sion mechanisms in either Anomalops or vs. 22.7-48.6% HL in all other anomalopid cladistically primitive members of the Pro- genera), absence of postorbital papillae toblepharon lineage could be ancestral for (eight in Anomalops, one in other anomal- the family. Nevertheless, the phylogeny of opids, none in the outgroups), and low Johnson & Rosenblatt (1988) and Rosen- number of gill rakers (21 or fewer in Pro- blatt & Johnson (1991) emerged as the sin- toblepharon, monocentrids, and some gle most parsimonious tree based on 18 in- trachichthyids, 24 or more in other flash- formative characters (Table 4). We concur light fishes). with Johnson & Rosenblatt (1988) that the occlusion mechanisms of Anomalops and Acknowledgments Photoblepharon probably represent the most derived conditions within the family We thank N. Sims of the Cook Island and that the occlusion mechanism of Pro- Fisheries Department for donating the spec- toblepharon, which lacks a well defined imen of the new flashlight fish to the Aus- shutter knob, a stalk hook to engage the tralian Museum. J. E. McCosker, J. A. shutter knob, and a groove and swelling in Moore, R. H. Rosenblatt, V. G. Springer, 382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.—Character matrix used in constructing opidae) from the Red Sea.—Sea Fisheries Re- cladogram in Figure 4, followed by brief description search Station, Haifa 60:57—62. of character states. See text and Johnson & Rosenblatt Allen, G R., & D. R Robertson. 1994. Fishes of the (1988) for further descriptions of characters. ? = miss- tropical eastern Pacific. Crawford House Press, ing data, 9 = not applicable, % = polymorphism. Bathurst, Australia, 332 pp. Baldwin, C. G, & G D Johnson. 1995. A larva of 1-5 6-10 11-13 16-18 the Atlantic flashlight fish. Kryptophanaron al- Trachichthyidae 00000 00100 1%999 90% fredi (Beryciformes: Anomalopidae), with a Monocentridae 00000 00000 00999 900 comparison of beryciform and stephanoberyci- Anomalops 11120 00101 11000 010 form larvae.—Bulletin of Marine Science 56:1— Protoblepharon 11701 11210 00000 010 24. Parmops 11711 11210 21001 021 Colin, P. L., D. W. Ameson, & W. E Smith-Vaniz. Phthanophaneron 11711 11210 OHIO 021 1979. Rediscovery and redescription of the Ca- Kryptophanaron 11111 11210 01111 121 ribbean anomalopid fish Kryptophanaron alfre- Photoblepharon 11111 11211 11111 121 di Silvester and Fowler (Pisces: Anomalopi- dae).—Bulletin of Marine Science 29:312-319. 1. Light organ absent (0), present (1). Gomon, M. E 1994. Family Trachichthyidae. Pp. 2. Neural and haemal spines of fourth preural ver- 399-410 in M. F. Gomon, J. C. Glover, & R. tebra do not support procurrent caudal rays (0), H. Kuiter, eds., The fishes of Australia's south support procurrent caudal rays (1). coast. State Print, Adelaide, 992 pp. 3. Interarcual cartilage absent (0), present (1). Harvey, E. N. 1922. The production of light by the 4. Postorbital papillae zero (0), one (1), eight or fishes Photoblepharon and Anomalops.—Car- nine (2). negie Institute of Washington Publication 312: 5. Supramaxillae one (0), two (1). 45-60. 6. Transverse gular ridges absent (0), present (1). Haygood, M. G, & D. H. Cohn. 1986. Luciferase 7. Lateral dentary tooth patch small (0), large v- genes cloned from the unculturable luminous shaped (1). bacteriod symbiont of the Caribbean flashlight 8. Lateral body scale rows <50 (0), 50-100 (1), fish, Kryptophanaron alfredi.—Gene 45:203- >100 (2). 209. 9. Pale (reflective?) lateral-line scales absent (0), Johnson, G. D. 1984. Percoidei: development and represent (1). *10. Pelvic-fin spine present (0), absent (1). lationships. Pp. 464-498 in H. G Moser, W. J. *11. Comer of maxilla papillose (0), with bony or- Richards, D. M. Cohen, M. P. Fahay, A. W namentation (1), smooth (2). Kendall, Jr., & S. L. Richardson,eds., Ontogeny *12. Gill rakers 21 or fewer (0), 24 or more (1). and systematics of fishes.—American Society 13. Shutter knob absent (0), present (1). of Ichthyologists and Herpetologists Special 14. Stalk hook absent (0), present (1). Publication 1:1-760. **15. Efhmomaxillary ligament without groove (0), , & R. H Rosenblatt. 1988. Mechanisms of with groove (1). light organ occlusion in flashlight fishes, family 16. Efhmomaxillary ligament without swelling (0), Anomalopidae (Teleostei: Beryciformes), and with discrete swelling (1). the evolution of the group. —Zoological Journal 17. Nasal connected to lacrimal (0), not connected of the Linnean Society 94:65—96. but close (least distance between bones 1.0— Konishi, Y, & M. Okiyama. 1997. Morphological de- 1.6% headlength)—(1), separated by large velopment of four trachichthyoid larvae (Pisces: space (least distance between bones >4.0% Beryciformes), with comments on trachich- headlength)—(2). thyoid relationships. — Bulletin of Marine Sci- 18. Dorsal-fin rays 14 or fewer (0), 15 or more (1). ence 60:66-88. Kotlyar, A. N. 1980. Systematics and distribution of * Informative only as autapomorphy of one or more trachichthyid fishes (Trachichthyidae, Beryci- genera; ambiguous character. formes) of the Indian Ocean.—Transactions of the P, P, Shirshova Institute of Oceanology 110: and H. J. Walker made helpful comments 177-224. on a draft of this manuscript. McCosker, J. E. 1982. Discovery of Kryptophanaron alfredi (Pisces: Anomalopidae) at San Salvador, Literature Cited Bahamas, with notes on anomalopid light or- gans.—Re vista De Biologia Tropical 30:97—99. Abe, T, & Y. Haneda. 1973. Description of a new fish , & R. H. Rosenblatt. 1987. Notes on the bi- of the genus Photoblepharon (Family Anomal- ology, taxonomy and distribution of anomalopid VOLUME 110, NUMBER 3 383

fishes (Anomalopidae: Beryciformes).—Japa- , & W. L. Montgomery. 1976. Kryptophan- nese Journal of Ichthyology 34:157—164. eron harveyi, a new anomalopid fish from the Moore, J. A. 1993. The phylogeny of the Trachi- eastern tropical Pacific, and the evolution of the chthyiformes (Teleostei: Percomorpha). Pp. Anomalopidae.—Copeia 1976:510-515. 114-135 in G. D. Johnson & W. D. Anderson, Shimizu, T. 1984. Order Beryciformes. Pp. 108-110 Jr., eds., Phylogeny of the Percomorpha.—Bul- in H. K. Masuda, K. Amaoka, C. Araga, T letin of Marine Science 52:1-629. Uyeno, & T. Yoshino, eds., The fishes of the Patterson, C, & G. D. Johnson. 1995. The intermus- Japanese Archipelago. Tokai University Press: cular bones and ligaments of teleostean fish- 1-437. es. — Smithsonian Contributions to Zoology Smith, M. M. 1986. Family No. 128: Monocentridae. 559:1-83. Pg. 413 in M. M. Smith and P. C. Heemstra, eds., Smiths' Sea Fishes. Macmillan South Af- Roberts, C. D. 1993. Comparative morphology of rica (Publishers) (Pty) Ltd., Johannesburg, 1047 spined scales and their phylogenetic signifi- pp. cance in the teleostei.—Bulletin of Marine Sci- Swofford, D. L. 1991. PAUP: Phylogenetic analysis ence 52:60-113. using parsimony, version 3.0s. Computer pro- Rosenblatt, R. H., & G. D. Johnson. 1991. Parmops gram distributed by the Illinois Natural History coruscans, a new genus and species of flashlight Survey, Champaign, Illinois. fish (Beryciformes: Anomalopidae) from the Zehren, S. J. 1979. Pisces: Teleostei. The comparative South Pacific.—Proceedings of the Biological osteology and phylogeny of the Berycifor- Society of Washington 104:328-334. mes.—Evolutionary Monographs I, 389 pp.