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Accepted by J. Sparks: 20 Jun. 2016; published: 27 Jul. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4144 (2): 227
–
242
http://www.mapress.com/j/zt/
Article
227
http://doi.org/10.11646/zootaxa.4144.2.5
http://zoobank.org/urn:lsid:zoobank.org:pub:5E49500D-AEEB-495A-9A2E-2BB0429CCB99
A new genus and species of cardinalfish (Percomorpha, Apogonidae,
Sphaeramiini) from the coastal waters of Vietnam: luminescent or not?
THOMAS H. FRASER
1
& ARTEM M. PROKOFIEV
2
1
Florida Museum of Natural History, University of Florida, Gainesville, Florida, 32611‒7800 USA, and Mote Marine Laboratory,
1600 Ken Thompson Parkway, Sarasota, FL 34236‒1096 USA. E-mail: cardinalfish@comcast.net
2
Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, pr. Leninskii 33, Moscow, 119071 [address for correspon-
dence] and Shirshov Institute of Oceanology, Russian Academy of Sciences, pr. Nakhimovsky 36 Moscow 117218, Russia.
E-mail: prokartster@gmail.com
Abstract
Xeniamia atrithorax is a diminutive new genus and new species. The following combination of melanophore patterns is
unique among known apogonids: a large cluster of melanophores in the skin anterior to the insertion of the pelvic-fin base,
then extending forward along the sides of the isthmus; posterior portion of the oral chamber is black with melanophores
extending below the gill apparatus forward along the base of the oral chamber thence along the basibranchials ending pos-
terior to and below the level of the tongue; a line of large melanophores extends along the inner side of the ceratohyals;
black to blackish stomach; and the anterior portion of the intestine is mostly pale with few black spots becoming more
densely spotted with melanophores past the first bend and black from the second intestinal turn to the anus. The peritone-
um is silvery with scattered melanophores that are more densely distributed along the lower portion of the abdominal cav-
ity. There are two pale, large pyloric caeca at the connection between the stomach and intestine. The combination of eight
first-dorsal spines, two supernumerary dorsal spines, one supraneural, lacking an ossified basisphenoid, two epurals and
fused hypurals 1+2, preopercle with a smooth ridge and edges, fourth dorsal spine longer than third distinguish this car-
dinalfish from all other genera. This new genus may be related to Jaydia in the Sphaeramiini rather than with the Osto-
rhinchini or Siphamiini. Possible luminescent activity is inferred from anatomy and unique melanization patterns versus
suppression of light from luminescent prey in the alimentary canal. A table lists suspected and known luminescent
apogonids.
Key words: Xeniamia atrithorax n.g. & n. sp., Ostorhinchus, Siphamia, luminous cardinalfishes, South China Sea, oste-
ology, luminescent morphology
Introduction
Morphological diversity of extant cardinalfishes is broad. The number of Recent genera and species peak on coral
reefs. Other known habitats include freshwater streams known from Australia, New Guinea and the West Pacific,
estuaries, bays and to nearly 300 meters depth. Within the water column habitats range from midwater to use of
burrows by an undescribed species (personal communication, Mark Erdmann & Gerry Allen). Fossil otolith- and
skeleton-based records of apogonids are definitely known from Early Eocene (Ypressien and Lutetien) indicating a
rather long geological history (Samant & Bajpai 2001; Bannikov 2008). The Late Cretaceous (Santonian and
Maastrichtian) otolith-based records (Nolf & Stringer 1996; Nolf 2003) later were reclassified as the members of
Myctophiformes and Beryciformes (Schwarzhans 2010). There are least 40 valid genera/subgenera and 356 valid
species (Eschmeyer & Fong 2016). The number of valid genera in the Apogonidae described per year is beginning
to plateau (Mabuchi et al. 2014). Deeper marine realms and geologic strata are likely sources of more undescribed
genera. We add the description of another, possibly luminous new genus and species to the Apogonidae as a result
of trawling in 40–119 m along coastal Vietnam.
All known luminous cardinalfish emit light associated within the alimentary canal or through open connections
to the alimentary canal (see Karplus 2014; Thacker & Roje 2009). In the type species of Jaydia Smith 1961 and
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several close relatives the anterior light-emitting location is associated with a pouch at the second bend of the
intestine followed by a pair of small oval organs just anterior to the anus (Kato 1947; Iwai & Asano 1958; Haneda
& Johnson 1962). Three other species in Jaydia lack the posterior oval organs (Gon 1997). Two other species in
Jaydia have an anterior luminous organ like other Jaydia and a post-pelvic diffuser organ somewhat similar to
Siphamia Weber 1909 (Weber 1909, 1913; Gon & Allen 1998). In Ver ul u x Fraser 1972 the light organs appear at
the ends of two pyloric caeca (Haneda et al. 1969). Species of Siphamia have one light organ as a pouch or disc
with a duct to the second bend of the intestine (Iwai 1958, 1959, 1971; Haneda 1965; Tominaga 1964) much like
the anterior organ of Jaydia. All species of Siphamia have two pairs of diffuser organs from the floor of the mouth
and from the isthmus to/or past the anal fin (see Gon & Allen’s 2012 discussion). The additional light organs
present in the floor of the mouth are connected with the pharynx for some species of Siphamia (Fishelson et al.
2005; Thacker & Roje 2009). Three of 12 species in Tae ni am i a Fraser 2013 all previously in Archamia Gill 1863
have the light organ similar in morphology to Jaydia according to Haneda et al (1969). Thacker & Roje (2009)
described the light organs for these as: “In the Archamia species examined, the light organ is manifested as the
anterior portion of a bend in the intestine, without constriction or other delimitation of the luminescent from the
non-luminescent portion (Fig. 3A).”
The origin of the chemicals needed to produce light has generated confirmed hypotheses: 1) luminous enteric,
symbiotic bacteria present in seawater are acquired by larvae (Leis & Bullock 1986; Dunlap et al. 2012; Karplus
2014); 2) feeding on luminous invertebrates capturing their chemicals (Haneda et al. 1969). The presence of
luminous enteric bacteria was found in 22 species of fishes (Ruby & Morin 1979). One of the species examined
was Apogon parri Breder 1936, a synonym of Apogon pacificus Herre 1935 and a non-luminous species with a
pale alimentary canal and pale peritoneum known only from the Eastern Pacific Ocean. The enteric contents from
four specimens of Apogon pacificus were cultured resulting in 1–42% of colonies being luminous. This kind of
survey, if done for more apogonids, would likely show that luminous enteric bacteria are present more widely in
cardinal fish then indicated by present number of presumed and known luminous cardinal fish (Table 1). Fishelson
et al. (1997) surveyed 78 species of cardinal fish for color patterns of the alimentary canal and peritoneum: 22 had
black guts, 5 partly black guts and 51 were unpigmented or had dispersed melanophores. Some had the peritoneum
with melanophores lightly to densely scattered. They suggested that nocturnal predators may conceal luminous
prey. This likely interpretation also may be the first step to developing systems for the emission of light useful to a
species. Species of Siphamia may have silvery to a darkish peritoneum with melanophores variably along the
alimentary canals (Fishelson et al. 1997; Gon & Allen 2012). In their revision, Gon & Allen discuss two type of
color patterns for the ventral light organs for species of Siphamia and the condition of the light organs adjacent to
the tongue of some species. All of these species are listed in Table 1. Those species known to use external chemical
sources generally will have black or partly black alimentary canals or melanophores in the peritoneum (Haneda et
al. 1970; Gon 1997, fig. 2D; Fishelson et al. 1997). All luminous apogonids have light passing though translucent
tissue of the mouth/isthmus/thorax/pelvic/anal region and/or enhanced by using reflectors and diffusers spreading
indirect light along the ventral part of the body.
Thacker & Roje (2009) concluded that each instance of a luminous representative in a genus evolved
independently based on a molecular tree that included 18 genera and 32 species of cardinal fish. A comparison with
the molecular trees reported by Mabuchi et al. (2014) show a distribution of the genera with luminous species
supporting the initial conclusion of Thacker & Roje. Five tribes proposed by Mabuchi et al. (2014) in the
Apogoninae have species known or suspected of being luminous (Table 1): Archamiini, Ostorhinchini, Siphamiini,
Sphaeramiini, and Veruluxini. Another three tribes having species with black guts or dark peritoneum are yet to be
explored for luminous activity: Apogonini, Gymnapogonini and Rhabdamiini.
Here, we describe unique patterns of melanophores for this new genus and species that may be associated with
luminescence or have the ability to concealing luminous prey. Tribal position is suggested by morphological
character since no specimens had been preserved for molecular information. We examine selected osteological and
luminescent characters for some species of Jaydia driven by some shared characters present in the new genus. We
refer to Gon & Allen’s (2012) revision of Siphamia for luminescent characters and to cleared and stained
specimens, radiographs and Fraser’s (1972) osteological comparisons.
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XENIAMIA ATRITHOR AX NEW GENUS & SPECIES
TABLE 1. Known to produce light or suspected (*) luminescent apogonids with the author first describing either
bioluminescence or inferred by morphology. For Siphamia enough species have been observed to assume those with the
ventral organ and diffusers are also luminous (see Haneda & Johnson 1962; Gon & Allen 2012).
1
permutata & versicolor are synonyms;
2
misidentified as ellioti;
3
elloiti is a synonym.
Tribe Genus Species Literature
Archamiini Taeniamia fucata Haneda, Tsui & Sugiyama 1969
Taeniamia lineolata Haneda, Tsui & Sugiyama 1969
Taeniamia zosterophora Haneda, Tsui & Sugiyama 1969
Ostorhinchini Ostorhinchus gularis* Fraser & Lachner 1984
Siphamiini Siphamia arabica* Gon & Allen 2012
Siphamia argentea* Lachner 1953
Siphamia brevilux* Gon & Allen 2012
Siphamia cephalotes Castelnau 1875
Siphamia corallicola* Allen 1993
Siphamia cuneiceps Whitley 1941
Siphamia cyanophthahma* Gon & Allen 2012
Siphamia elongata* Lachner 1953
Siphamia fistulosa* Weber 1909
Siphamia fraseri* Gon & Allen 2012
Siphamia fuscolineata* Lachner 1953
Siphamia goreni* Gon & Allen 2012
Siphamia guttulata* Alleyne & Macleay 1877
Siphamia jebbi* Allen 1993
Siphamia majimai Matsubara & Iwai 1958
Siphamia mossambica* Smith 1955
Siphamia papuensis* Gon, Allen, Erdmann & Gouws 2014
Siphamia randalli* Gon & Allen 2012
Siphamia roseigaster Ramsay & Ogilby 1887
Siphamia senoui* Gon & Allen 2012
Siphamia spinicola* Gon & Allen 2012
Siphamia stenotes* Gon & Allen 2012
Siphamia tubifer
1
Weber 1909
Siphamia tubulata* Weber 1909
Sphaeramiini Jaydia argyrogaster* Weber 1909
Jaydia hungi* Gon 1996
Jaydia photogaster* Gon & Allen 1998
Jaydia poeciloptera Haneda, Tsui & Lynch 1970
Jaydia smithi
2
Iwai & Asano 1958
Jaydia striata Haneda, Tsui & Sugiyama 1969
Jaydia striatodes* Gon 1996
Jaydia truncata
3
Kato 1947
Veruluxini Verulux cypselurus Haneda, Tsui & Sugiyama 1969
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Methods
Methods of taking and recording meristic data and measurements are as follows:
Standard length. Symphysis of upper jaw to base of hypural plate.
Body depth. Origin of first dorsal spine to insertion of pelvic spine.
Head length. Front of symphysis of upper jaw to posterior-most membrane of opercular flap.
Upper jaw length. Front of symphysis of upper jaw to mid-posterior edge of maxilla.
Snout length. Front of symphysis of upper jaw to anterior edge of orbit.
Eye diameter. Horizontal orbit distance to edge of bony borders.
Pectoral fin length. From the dorsal anterior base to tip of longest ray.
Pelvic fin length. From the origin of pelvic fin spine to tip of longest ray.
Caudal peduncle depth. Least depth in vertical plane.
Caudal peduncle length. Horizontal distance from end of the base of the anal fin to lower hypural base.
Spine length. Anterior base of spine to its tip.
Interorbital width. Least distance between the dorsal bony edges of the eyes.
Dorsal and anal fin-rays. All elements with the last ray, a double element with a single support, counted as one.
Pectoral fin-rays. All elements counted with no differentiation between branched and unbranched elements.
Pectoral radials with formula suggested by Starks (1930).
Gillrakers. All elements counted and divided into rudiments and well-developed rakers. A single gillraker in
the angle included as part of the lower arch count and separate from the upper arch gillrakers. Rudiments are small,
undeveloped structures about as wide as high.
Lateral-line scales. Pored scales from posttemporal bone to base of hypural plate.
Longitudinal rows of scales above lateral line. Same method as lateral-line count starting with scale in
transverse row just above first pored scale.
Transverse scale rows. Rows of scales from origin of first dorsal fin (but not median row) counting downward
and backward to but not including lateral line, and rows of scales from anal fin origin counting upward and forward
to but not including lateral line.
Predorsal row of scales. Median row of scales on nape from anterior-most one to origin of first dorsal spine,
including last scale at spine.
Circumpeduncular scales. Rows around peduncle at narrow portion divided into those above lateral line, the
two lateral line rows and those below the lateral line.
Vertebrae. Precaudal from base of skull to last open arch. Caudal from first closed arch including terminal
centrum.
Caudal-fin rays. Longest unbranched dorsal caudal ray plus branched caudal rays to longest unbranched
ventral caudal ray for total principal rays. Procurrent rays anterior most to first unbranched dorsal and ventral rays.
All measurements are in millimeters to the nearest 0.1mm. All proportions are based on standard length and all
material is reported by standard length (SL) rounded to the nearest millimeter, except for the primary type material.
Radiographs, film or scanned from film by the first author, and digital images were examined to obtain
vertebral counts including the terminal centrum, number of hypurals, number of epurals, status of the anterior
paired uroneurals, status of a terminal sheath with respect to the fourth hypural, status of an eighth dorsal spine and
supporting elements, number of supraneurals (see Springer & Smith-Vaniz 2008 for formula), number of dorsal
and anal spines on the first proximal-middle element (pterygiophores), number of proximal-middle radials anterior
to first haemal spine and between the first and second haemal spines. Radiographs have been accumulated by the
first author beginning in1966, taken at museum facilities at the USNM and at RUSI (now SAIAB). Many other
radiographs came from a program initiated by E. A. Lachner during his studies of apogonids at the USNM. The
second author provided film radiographs of the new genus and species taken at the museum in Russia.
Sources of photographs are listed in the captions. Acronyms used in the lists of materials for institutions and
collections cited, follow usage given in Fricke & Eschmeyer (2016) except SAIAB replaces RUSI as a result of a
name change and ZMH replaced IOES. Field abbreviations are as follows: D - Albatross Expedition (Dredge).
Field data taken during the trawling surveys of 2005–2012 off Khanh Hoa Province, Vietnam used here for
habitat and ecology comments were from records by the second author.
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XENIAMIA ATRITHOR AX NEW GENUS & SPECIES
Xeniamia new genus
Type species: Xeniamia atrithorax new species
Diagnosis. A genus of Sphaeramiini with the following unique morphological character combination: two epurals,
fused hypurals 1+2, no terminal bony sheath over third and fourth hypurals, lacking uroneurals, one supraneural,
two supernumerary dorsal spines, visible eighth dorsal spine (may appear hidden when damaged), fourth dorsal
spine longest, smooth vertical edge and smooth unossified ventral edge of preopercle, third infraorbital without
suborbital shelf, basisphenoid absent, scattered melanophores on silvery peritoneum, anterior intestine speckled
with black spots to past first bend of intestine, two pale pyloric caeca at the anterior end of intestine, stomach black,
intestine black from second bend to anus, black lining of the ventral and posterior oral cavity, and black spots on
outer isthmus lining from external thoracic area.
Description. See species description.
Color in alcohol. See species description.
Distribution. South China Sea off Khanh Hoa Province, Vietnam.
Etymology. Xenos, Greek meaning stranger and Amia, Greek referring to a type of fish, often used with
cardinalfish names, treated as feminine noun.
Remarks. Mabuchi et al. (2014) proposed new subfamilies and tribal classification for the Apogonidae. The
subfamily key places Xeniamia in the Apogoninae. Using their generic key, Xeniamia falls out in the couplet
segment with Jaydia and Ostorhinchus Lacepède 1802 excepting the reported eight first dorsal spines in Jaydia
erythrophthalma Gon, Liao & Shao 2015. No specimens were preserved for DNA analyses to help provide support
for tribal placement.
Tribes we excluded for placement using morphologic characters are: Apogonichthyini (thin supramaxilla, third
dorsal spine longest, stomach and intestine pale); Apogonini (large unossified preopercular flap, six first dorsal
spines, second dorsal-fin spine longest, pale stomach and intestine except one species); Cheilodipterini (large
canine or caninoid teeth, small supramaxilla, six first dorsal spines, second dorsal-fin spine longest, pale stomach
and intestine); Glossamiini (large supramaxilla, six first dorsal spines, second dorsal-fin spine longest, pale
stomach and intestine); Gymnapogonini (six first dorsal-fin spines, second dorsal-fin spine longest, narrow band of
teeth with some caninoid teeth, terminal centrum fused with hypurals 3+4, parhypural fused with hypurals 1+2);
Lepidamiini (small body scales, third dorsal-fin spine longest, pale stomach and intestine), Pristiapogonini
(preopercular edges and ridge serrate, pale stomach and intestine, third dorsal-fin spine longest); Rhabdamiini (6–7
first dorsal-fin spines, second or third spine longest, anal fin with 11–13 soft rays, 24–31 gill rakers, terminal
centrum fused with hypurals 1+2+3+4, pale stomach and intestine); Veruluxini (six first dorsal-fin spines, second
spine longest, terminal centrum fused with hypurals 1+2+3+4, modified pyloric caeca) and Zoramiini (six first
dorsal-fin spines, second spine longest, five free hypurals, three epurals). Characters of three other tribes are
discussed in more detail.
The Ostorhinchini is known to have more than 91 species in at least four unresolved molecular lineages that
lack distinctive independent morphologic support (Mabuchi et al. 2014). Almost all known species have blackish
stomachs and intestines. Many species have single stripe to multiple arrangements on the head and or body. All
have a serrate edges of the preopercle, a very few species have six first dorsal-fin spines, most have seven first
dorsal-fin spines, most have the third first dorsal-fin spine longest, a few have second first dorsal-fin spine longest,
preopercle ridge smooth, edges serrate, 2–3 supraneurals, supramaxilla absent, basisphenoid present, shelf present
on third infraorbital, one pair of reduced uroneurals or absent, three epurals, five free hypurals or 1+2 fused.
Ostorhinchus moluccensis (Valenciennes 1832), O. monospilus (Fraser, Randall & Allen 2002) and O. oxina
(Fraser 1999) variably have the fourth dorsal spine longer, equal or shorter. Ostorhinchus margaritophorus
(Bleeker 1854) is the only known species in the genus with fused hypurals 1+2. The only know species suspected
of having luminous activity is Ostorhinchus gularis (Fraser & Lachner 1984). Xeniamia differs from all members
of Ostorhinchini by the presence of eight first dorsal-fin spines, smooth ridge and edges of preopercle, absence of
basisphenoid and suborbital shelf, single supraneural, melanistic patterns and absence of spotted, striped or banded
pattern on flanks.
Xeniamia appears similar to species in the monogeneric Siphamiini but lacks the luminous apparatus along the
ventral side of the body present in all Siphamia (Gon & Allen 2012). Species of Siphamia have six or seven first
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dorsal-fin spines; third dorsal spine longest; terminal centrum fused with upper hypural plate (3+4 hypurals fused)
and free but fused 1+2 hypurals, or terminal centrum fused with hypurals 1+2+3+4 (Fraser 1972). Xeniamia shares
with some species of Siphamia a single supraneural, lacking an ossified basisphenoid, two epurals, no paired
uroneurals, no terminal bony sheath over third and fourth hypurals and small size but differs from that genus by the
presence of eight first dorsal-fin spines, fourth spine longest; presence of uroneurals, free upper hypurals 3 and 4,
lacking a suborbital shelf, and melanistic patterns.
The Sphaeramiini are represented by seven genera and 55 species lacking molecular resolution for several
genera (Mabuchi et al. 2014). This tribe has genera with the following characters: no supramaxilla, three epurals,
two or three supraneurals, two supernumerary dorsal spines, seven or eight first dorsal spines, some species with
luminous organs, paired uroneurals, stomach and intestine pale, speckled or black, peritoneum pale or with
speckled melanophores. All species of Nectamia Jordan 1917 have high gill raker counts (22-30), bars on body,
seven first dorsal-fin spines, third dorsal spine longest, and both stomachs and intestine blackish. Sphaeramia
Fowler & Bean 1930 and Pterapogon Koumans 1933 are deep bodied, with at least one bar on body, have seven
first dorsal-fin spines, third dorsal spine longest and spinous procurrent rays. Quinca Mees 1966 is black and deep-
bodied (Vagelli 2014), possibly belonging in Apogonichthyoides (see Fraser & Allen 2010). Apogonichthyoides
Smith 1961 has serrated hind edge of preopercle, seven to eight first dorsal-fin spines, with third spine always
longest, mostly dark body color with spotted or barred pattern and/or ocellus-like mark in some species, and cheek
bars, peritoneum, stomach and intestine pale or spotted. All species of Jaydia are characterized by the fourth
dorsal-fin spine longest, truncated or rounded caudal fin, ventral surface of head and/or isthmus bearing some
melanophore pigmentation, light organs associated with the intestine present in some species. These characters
suggest a likely relationship with our new genus than any other genus in the Sphaeramiini.
Xeniamia differs from known species of Jaydia by having a single supraneural, lacking an ossified
basisphenoid, infraorbital without a shelf, two epurals instead of three, fused hypurals 1+2 instead of being free,
lacking a bony sheath as part of the terminal centrum that covers the third and fourth hypurals, except present in
Jaydia photogaster (Gon & Allen 1998) and J. argyrogaster (Weber 1909), and a short visible eighth first dorsal-
fin spine (except present in Jaydia erythrophthalma Gon, Liao & Shao 2015) instead of a tiny nubbin hidden under
the skin or absent. Like all Jaydia. the fourth first dorsal-fin spine in Xeniamia is longer than the third.
We place Xeniamia in the Sphaeramiini pending molecular information.
Xeniamia atrithorax new species
Figures 1–8, Table 2
Diagnosis. A miniature species of Xeniamia (Figs. 1–3) attaining less than 30 mm SL with the following color
pattern and morphologic characters: a large melanophore patch anterior to insertion of pelvic fin, extending
forward on each side of isthmus, progressively thinning out (Fig. 2), black stomach, anterior intestine pale with
black spots black intestine from second bend, interior of lower gill chamber blackish with concentrated
melanophores extending forward to dorsal part of isthmus (Fig. 3), dorsal fins VIII-I,9 eighth dorsal spine visible,
fourth dorsal spine longer than third, anal fin II,8, nine gill rakers.
Description. Holotype (Figs. 1–2): proportions as percentage of SL in Table 2. All scales missing on head and
body; dorsal fins VIII–I,9; anal fin II,8, damaged; pectoral fins 14–14; pelvic fins I,5; outer row of first gill arch
with 2 rudiment, 1 raker on upper arch, and 2 rudiments and 8 rakers on lower arch, 6 pseudobranch filaments.
Villiform teeth in band on premaxilla & dentary; one row on vomer; one row (long) on palatine; none on
basihyal or ectopterygoid.
Damaged preopercle lower edge with smooth, unossified flap, vertical bony edge smooth, bony ridge edge
smooth.
Vertebrae 10+14, one ossified supraneural as /0/1/, two epurals, no paired uroneurals, no terminal bony sheath
over third and fourth hypural, fused hypural 1+2, parhypural free, crest on preural centrum 2; principal caudal fin-
ray 17, upper and lower unbranched.
Preserved color pattern (Figs 1 & 2): head with scattered melanophores dorsally from between eyes posteriorly
to anterior edge of nape, a few melanophores in a patch near posttemporal; scattered melanophores at insertion of
pelvic fins and on pale region posterior to a large melanophore patch extending forward on each side of urohyal,
progressively thinning out; no melanophores on any fins or at fin-bases.
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XENIAMIA ATRITHOR AX NEW GENUS & SPECIES
TABLE 2. Proportions as a percentage of standard length for the holotype of Xeniamia atrithorax with proportion ranges
for two subsamples of all of the paratypes.
* = one or more element broken
. = missing measurement because element broken
Internal melanophore organization (Figs. 3 & 4): Peritoneum silvery with speckled black spots becoming more
dense in distribution ventrally, anterior wall black at connection with pharynx, black tissue extending forward
along the floor of the oral cavity to below tongue; swim bladder whitish, no black spots; gonad pale with diffuse
black spots; dorsal vein to sinus with black spots; intestine pale with black spots becoming more densely spotted
past the first intestinal turn to completely black by the second intestinal turn thence black to anus along ventral
portion of abdominal cavity; two large, pale pyloric caeca extending along the lower portion of the stomach;
stomach black on upper half and with dense black spots on lower half; liver and heart pale.
Live color pattern unknown.
Gravid female, ova well-developed, extruding posteriorly.
Paratypes. Proportions in Table 2. Pectoral-fin rays 14–15; lateral line scales 20–22 (8 specimens);
circumpeduncular scales about 8; predorsal scales about 4; transverse row of scales above lateral line and first
dorsal fin 3; transverse rows of scales below lateral line and anal fin 5 (transverse count based on one specimen);
existing scales on body ctenoid except cycloid around paired-fin bases; and two cleared and stained specimens with
one supraneural as /0/1/, fused hypurals 1+2, no terminal centrum sheath over the third fourth hypural, no evidence
of paired uroneurals, 10+14 vertebrae; one cleared and stained specimen has a single epural (Fig. 5); radiographs of
36 specimens, one shown (Fig. 6). Many specimens have the first and/or eighth dorsal spines missing.
Cranial crests Y-shaped, well-developed, supraoccipital crest moderate, basisphenoid absent, otic bulla
incompletely ossified.
Six infraorbitals (Fig. 7) with upper and lower edges smooth, lachrymal and second infraorbital expanded,
third infraorbital without suborbital shelf; single large extrascapular; posttemporal edge smooth.
Holotype Paratypes
USNM 436743 ZMMU
23156
ZMMU
23157
Nha Trang Bay N=8 Van Phong Bay N=10
25.1 mm SL 22.5–29.5 mm SL 22.0–29.0 mm SL
Percentages
Body depth 38.7 38.0–40.4 35.7–43.2
Head length 37 38.9–42.0 37.9–44.4
Eye diameter 8.76 10.0–10.2 9.26–11.4
Snout length 5.98 7.20–9.62 7.72–11.4
Interorbit width 9.16 8.93–10.0 9.26–11.6
Upper jaw 22.3 20.0–23.0 21.1–25.0
First dorsal spine 1.2 1.48–2.69* 1.11–3.46*
Second dorsal spine 6.77 4.80–10.4 7.56–13.5*
Third dorsal spine 15.9 12.5–14.2 12.8–18.2*
Fourth dorsal spine 16.7 14.0–16.4 14.8–18.2*
Second dorsal fin spine . 17.3–26.0* 20.4–25.5*
First anal spine 3.57 3.05–4.80 2.76–4.56
Second anal spine 9.16 8.47–10.4 9.33–11.4
Pectoral fin length . 25.0–30.0* 24.6–28.9*
Pelvic fin length . 20.0–23.0 20.7–24.1*
Caudal peduncle length 21.1 20.0–23.1 19.0–25.0
Caudal peduncle depth 13.2 12.0–14.3 13.1–14.9
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FIGURE 1. Xeniamia atrithorax, holotype, left side, USNM 436743, 25.1mm SL, female, by T.H. Fraser, scale= 4mm.
FIGURE 2. Xeniamia atrithorax, holotype, ventral view, USNM 436743, 25.1 mm SL, female, showing the thoracic
melanophore patch, by T.H. Fraser, scale=3 mm.
FIGURE 3. Xeniamia atrithorax, paratype, USNM 436744, right side of 25.5 mm SL specimen, dissection showing oral to
anal region. AIN=anterior portion of the intestine; AL=anterior portion of the liver; B1=first bend of intestine; B2 second bend
of intestine; G=gonad; H=heart; I=right side of isthmus; IN=straight potion of intestine from second bend; LO=lower oral
stratum; P=pelvic fin base; PC=pyloric caeca; S=black stomach; SB=swim bladder; T=location of external thoracic
melanophore patch; V=blood vessel to sinus, by T.H. Fraser, scale=3 mm.
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FIGURE 4. Xeniamia atrithorax, paratype, ZMMU 23157, 27.0 mm SL, canted view of the pyloric caeca, stomach and
dorsally flipped intestine to show the caeca. B1=first bend of intestine; B2=second bend of intestine; P=pyloric caeca;
S=stomach; by A.M. Prokofiev, scale=2 mm.
Maxilla with short expanded posterior plate; supramaxilla absent; junction between metapterygoid and
hyomandibula interdigitated; seven branchiostegal rays (4 + 3); junction between ceratohyal and epihyal smooth;
upper margin of ceratohyal deeply but smoothly concave without foramen; urohyal low and long, incised on
posterior margin; opercular bones incompletely ossified; lowermost pectoral bony radial much larger than three
upper ones, 2–1–1; pelvis with short and broad posterior processes, slightly divergent and broadly separated from
each other medially; basiventral processes short (approximately equal in length to posterior processes), rod-like,
strongly divergent; proximal-middle element of first anal-fin pterygiophore ending before first haemal spine, with
2/3 or 1/4 interdigitating pterygiophores; last pair of pleural ribs modified, expanded and shortened.
Many specimens with fin and body damage caused by trawls.
Otolith (Fig. 8). Oval (immature specimens) to sub-rhomboidal (adults), moderately deep otolith up to 2.5 mm
length. Dorsal rim weakly convex, entire, with indistinct predorsal angle and strongly pronounced broad postdorsal
angle. Ventral rim strongly convex, entire to indistinctly lobed; posterior rim strongly oblique, with more or less
developed incision at junction with ventral rim; this incision is less developed in youngest specimens. Sulcus ostial,
heterosulcoid; ostium and cauda of equal length; cauda narrow, straight bearing a distinct crest just above the crista
inferior; colliculi heteromorphy; anterior colliculum oval, raised towards dorsal edge; posterior colliculum narrow,
raised towards ventral edge; collum and pseudocolliculi absent; ostium possessing a conspicuous widening
between anterior colliculum and crista superior; cristae superior and inferior moderately expressed. Rostrum large
and massive, bluntly pointed, median; antirostrum moderately to well-developed, broad, rounded to bluntly
pointed; excisura sharply angled. Dorsal depression large but shallow; ventral furrow distinct along whole length of
ventral rim. Outer and inner surfaces weakly convex, slightly more on outer surface. Length:height ratio = 1.4–1.6;
length:width ratio = 4.5–4.9.
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FIGURE 5. Xeniamia atrithorax, paratypes, left side of caudal complex: A. with one epural and fin rays, ZMMU 23156, 24.5
mm SL. B. with two epurals and fin rays detached, ZMMU 23156, 25.0 mm SL. Specimens cleared and stained. e = epural; ph
= parhypural; tc = terminal centrum; 1+2, 3–5 = hypurals, by A.M. Prokofiev, scale = 1 mm.
FIGURE 6. Xeniamia atrithorax, paratype, ZMMU 23157, 24.2 mm SL, left side, a positive image from a radiograph. sn =
supraneural; e = epural; I = first dorsal spine; VIII = eighth dorsal spine; 10 = tenth vertebra, by A.M. Prokofiev, scale = 5 mm.
Distribution. South China Sea off Khanh Hoa Province, Vietnam (Nha Trang and Van Phong Bays).
Etymology. Atrithorax from ater, atra, atrum Latin adjective meaning black and thorax Greek masculine noun
meaning breastplate, the compound referring to the position of numerous melanophores as a dark spot in advanced
of the pelvic-fin base, a noun in apposition.
Habitat. This species has been taken in shrimp trawls on muddy bottom. Most of specimens were collected
from depths 70–119 m, but few samples were made at depths 40–60 m (currently in the unsorted part of
Vietnamese collection in IOM and not listed here in the material examined) in the main waterway of the Bay of
Nha Trang. The latter samples might have been pumped through canyons cutting out the outer shelf as proposed for
other fish taxa collected from the unusually shallow depths in this area (Nielsen & Prokofiev, 2010). There were no
specimens or fragments of sea urchins commonly trawling together with Siphamia specimens. Year-to-year
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fluctuations in density of population were observed – this species was present at 18 of 22 stations trawling eastward
and southward of Pyramid Island in 2005 and 2007 and two of 17 stations sampled in 2006 and 2009–2012. All
samples were made between April and July, and no specimens were collected during a winter monsoon season
(October–January). This species always appeared in groups of about 10-60 specimens per trawl, with many
specimens having ripe ova or brooding eggs.
FIGURE 7. Xeniamia atrithorax, paratype, ZMMU 23156, 25.0 mm SL cleared and stained, left view of the six infraorbitals,
by A.M. Prokofiev, scale = 1 mm.
Reproduction. The smallest specimen, 14.5 mm SL was immature. Males with buccal enlargement suggestive
of brooding eggs were 22.0–24.0 mm SL and showed loss of teeth on the vomer and palatines. A 23.0 mm SL male
had approximately 280 orangish eggs, 0.5 mm in diameter. Females with developed ova were as small as 23.0 mm
SL. The largest gravid female was 29.5 mm SL. No possible indications for commensal relationships of this species
were present.
Remarks. The remarkable organization of melanophores raises questions about what these patterns might play
in the life of this species: 1) suppressing the emission of light cause by luminous prey or 2) the use of chemicals
partially from prey and reactions to create light in the pyloric caeca and intestine. Parsimony suggests that a single
completely black alimentary canal or some variation of the length is all that is needed for the suppression of light.
The complicate system of melanophore patterns and speckled anterior intestine coupled with well-developed pale
caeca may allow for light to emanate from the anterior speckled isthmus region. It is not clear why the sinus vein or
the artery below the swim bladder have melanophores. The dense melanophores forming a blackish spot in advance
of the pelvic-fin base is unique among all known species of apogonids.
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FIGURE 8. Xeniamia atrithorax, lateral view of sagittae, paratypes, ZMMU 23158. A. SL 17.5 mm. B. SL 27.0 mm. C. SL
28.0 mm. D. Lateral and dorsal view of sagitta, SL 28.5 mm, by A.M. Prokofiev, scale = 0.5 mm.
We use the long fourth dorsal spine as an indication of relationship with Jaydia rather than with the few species
of Ostorhinchus that share this characteristic.
Gon (1997) revised species of Jaydia Smith (1961) as a subgenus of Apogon Lacepède (1801) with little
osteological information, referring to Fraser (1972) who treated many species in the broad non-monophyletic
subgenus Nectamia Jordan 1917 including five of those now in Jaydia. There are differing luminous systems
described for Jaydia: 1) Weber (1909, 1913, plate 10 figure 7) and Weber & de Beaufort (1929) partially described
for Jaydia argyrogaster (Weber 1909); Gon & Allen (1998) a more completely described this unique system for
Jaydia photogaster (Gon & Allen 1998). 2) Kato (1947), as Apogon marginatus (an unavailable name by Döderlein
in Jordan & Snyder 1901, see Eschmeyer et al. (2016), now J. truncata (Bleeker 1855) and Iwai & Asano (1958),
as Apogon ellioti Day 1875, now Jaydia smithi Kotthaus 1970. A list of known and likely luminous apogonids is
given in Table 1.
There are about 17–18 valid species of Jaydia (see Eschmeyer et al. 2016). Gon (1997) recognized ten species
and nine synonyms. Nine more nominal species (three or four appear valid) were not treated in Gon’s revision.
Three additional species have been described since Gon’s revision. We examined radiographs for about 11–12 valid
species (see comparative material) of Jaydia yielded the following: all had three supraneurals as 0/0/0–1/, all had
three epurals, all had five free hypurals. An eighth dorsal spine was reported for Jaydia erythrophthalma by Gon et
al. (2015) but provided limited information about other osteological characters. Radiographs confirm the 3
supraneurals described by Gon et al. and we observed three epurals, five free hypurals. All other Jaydia have seven
visible first dorsal-fin spines.
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The otoliths of Apogonidae are not sufficiently known, though preliminary data based on study of the various
Vietnamese taxa indicate morphologic diversity in the sagittal structures. Xeniamia shares a marked dorsal
widening of ostium and a narrow cauda bearing a collicular crest, both of which are characteristic for the apogonid
sagittae, but the poorly developed predorsal angle and a conspicuous excision at posterior ventral margin appear to
be peculiar for the new taxon. All small specimens less than 23 mm SL examined have otoliths strongly corroded
(by formalin fixation) and partially disintegrated. However, it is clear, that immature otoliths are more oval in
shape, with feebly developed ventroposterior incision. Variations in shape of otolith in adult specimens close to
maximal known size (SL 27.0–28.5 mm) are considerable, particularly in shape of dorsal and ventral rim, rostrum
and antirostrum and in expression of ventroposterior excision. Three otoliths shown in Figure 7 look like growth
changes rather than individual variations; however, they were taken from fishes of similar size, all of which are
females with ripe eggs close to spawning. Together with poor ossification of some skeletal elements this may
suggests a neotenic nature of Xeniamia.
Material examined. Holotype USNM 436743, 25.1 mm SL, female, Van Phong Bay,12°40.944'–12°36.924'N
109°29.752'–109°30.002'E,Vietnam, 30 May–1 Jun 2007, 97–119 m, 6 m shrimp trawl, 2247–0039 hrs, A.M.
Prokofiev, x-ray.
Paratypes: CAS 23808 (2, 21.5–24.3), ROM 98985 (2, 25.1–27.2), UF 237134 (3, 20.4–24.1) & USNM
436744 (2, 20.5–25.5), all same data as the holotype, 5 specimens x-ray. ZMMU 23156 (20, 22.0–29.5) all same
data as the holotype, x-ray, (2, 24.5–25.0) cleared & stained. ZMMU 23157 (19, 14.5–29.0), Van Phong Bay,
12°35.044'–12°29.927'N 109°30.004'–109°30.018'E, Vietnam, 30 May–1 Jun 2007, 95–120 m, 6 m shrimp trawl,
0144–0402 hrs, x-ray, A.M. Prokofiev. ZMMU 23158 (14, 16.0–29.0), Nha Trang Bay, off Hon Dun (Pyramid
Island), 12°13.200'–12°11.329'N 109°24.161'–109°23.944'E, Vietnam, 21 May 2005; 70-80 m, 6 m shrimp trawl,
2045–2145 hrs, x-ray, D.A. Astakhov & A.M. Prokofiev.
Comparative material. All in Jaydia with holotypes and syntypes listed by original genus, alphabetical by
species: Amia albomarginata Holotype, USNM 68402, 83.6 mm SL, Cavite, Luzon I., Philippines, x-ray. Apogon
argyrogaster Syntypes, ZMA 101.075, 2, 34.9–47.5 mm SL, near western coast of New Guinea, Siboga
Expedition. Jaydia carinata USNM 175735, 1, 97.0 mm SL, Marinduque I., Philippines, Albatross 3009, 2 Mar
1909, x-ray. USNM 71527, 5, 81.8–94.4 mm SL, Shimizu Suruga, Japan, Albatross, 1906, x-ray. Apogon ellioti
Lectotype, ZSI F1904, 77.5, Madras, India, x-ray. AMS Paralectotype, B8226, 77.3, Madras, India, x-ray. Jaydia
erythrophthalma Paratypes CAS 236504, 2, 50.3–57.0 between Luzon & Mindoro, Philippines, 1 June 2011, 115–
144 m, x-ray. Apogon fuscomaculatus Holotype, NTM S. 13284–014, 48.2 mm SL, NE of Charles Point, Beagle
Gulf, Northern Territory, Australia, 2 Sep 1992, 18–24 m, trawl, photograph, x-ray. Mionorus heraldi Holotype,
SU 38263, 104.5 mm SL, Ragay Gulf, Luzon, Philippines, x-ray. Jaydia hungi SAIAB 3116, 2, 87.6–96.5 Comoro
Ids., Jun 1959, x-ray. SAIAB 3117, 3, 88.2–100, Pemba I., Tanzania, x-ray. Jaydia lineata USNM 71108, 4, 64.3–
709, Nanao, Honshu I., Japan, x-ray. Jaydia novaeguinae USNM 177675, 2, 53.8–55.9 Peitaiho, China, x-ray.
USNM 51965, Southern Negros, Philippines, online x-ray. USNM 163229, Cavite, Luzon, online x-ray Apogon
photogaster Holotype WAM P.31213–057, 49.0 mm SL, Padoz Reef, Madang Lagoon, Papua New Guinea 21 Oct
1996, 18–23 m. Paratype USNM 348213, 43.6 mm SL, same data as holotype, photograph, x-ray. Paratype BPBM
38301, 42.0 mm SL same data as holotype, x-ray. Paratype WAM P. 30358–001, 29.8 mm SL, Madang Lagoon,
Padoz Reef, 28 Oct 1991, 26–30 m, x-ray. Paratype SAIAB (RUSI) 57597, 47.4 mm SL, Madang Lagoon, Padoz
Reef, 28 Oct 1991, 26–30 m, cleared & stained, data via Ofer Gon. BPBM 32628, 1, 49.0 mm SL, Tripod Reef,
lagoon off of Nagada Harbor, Papua New Guinea, 18 Nov 1987, 30 m, photograph, x-ray. Apogon quartus
Holotype, USNM 307688, 49.8 mm SL, Saya de Malha Bank, 11°05'00"S 62°02'00"E, x-ray. Apogon queketti
Syntype, SAM 11657 1, 75.1, Natal, South Africa, 40 fms. Syntypes, SAM 11658 5, 44.1–77.4, Natal, South
Africa, 40 fms. SAIAB (RUSI) 3177, 2, 64.4–68.5, Durban, Natal, South Africa. Jaydia smithi Paratypes, ZMH
5034 (ex IOES 120B,E, F), 5, 12.7–46.3, Gulf of Aden, 11°17.9'N 49°01.3E, Meteor Sta 97, 42 m, 18 Dec 1964, x-
ray. USNM 149365, 4, Manila Bay, Philippines, online x-ray. Amia striata Holotype, USNM 68403, 69.8 mm SL,
Manila Bay/Lingayen Gulf, Luzon, Philippines, D 5442, 82 m, x-ray. Paratypes, USNM 93410 11, 31.5–66.6, same
data as holotype, x-ray. Apogon tchefouensis Paratype, MNHN 41–148, 46.0, China, x-ray. Paratype, MNHN 41–
149 36.2 mm SL, China, damaged, x-ray. Jaydia truncata USNM 71275, 5, 80.4–86.3, Kagoshima, Satsuma,
Kyushu I., Japan, x-ray. USNM 59635, 2, 50.9–52.2, Matsushima Bay, Japan, x-ray. USNM 59636, Yamagawa,
Japan, online x-ray.
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Acknowledgments
We thank the following people for providing assistance with information prior to and for this study. Radiographs by
Ofer Gon, SAIAB, Ross Langston, Windward Community College, Helen Larson, NTM, Mark McGouther and
Sally Reader, AMS, Glenn Moore, WAM, Sandra Raredon, USNM, Arnold Suzumoto, BPBM, D. Catania and Jon
Fong, CAS were examined. Adolf Kotthaus ZMH, loaned type specimens of Apogon smithi and P. Hulley SAM
loaned type specimens of Apogon queketti while the first author was at the formerly known J.L.B. Smith Institute
of Ichthyology. Dmitry Astakhov IOM, assisted the second author with collecting Vietnamese fishes. Tilman
Alpermann SMF, Jørgen Nielsen ZMC and Jeff Williams USNM, provided logistic assistance with preserved fishes
sent by the second author. David Catania CAS, Erling Holm ROM and Shirleen Smith USNM, provided curatorial
assistance with type material. Larry Page, Rob Robins, Bill Smith-Vaniz and Bill Eschmeyer of the Florida
Museum of Natural History assisted with visits, curation and nomenclature. The first author’s acknowledges
support while at the former J.L.B. Smith Institute of Ichthyology, a Smithsonian Fellowship at the Natural History
Museum and a Research Fellowship at the Australian Museum that helped make this article possible. The work of
the second author was partly supported by a grant (16-04-00365) of the Russian Foundation for Basic Research.
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