Conway et al.: A new genus of clingfish from Indonesia
Taxonomy & Systematics
RAFFLES BULLETIN OF ZOOLOGY 69: 428–437

Date of publication: 13 October 2021
DOI: 10.26107/RBZ-2021-0062
http://zoobank.org/urn:lsid:zoobank.org:pub:5F24A27F-642A-433D-8B60-8A9B426F626E

Erdmannichthys, a new genus of Gobiesocidae (Teleostei: Gobiesociformes),and notes on the rare clingfish E. alorensis (Allen & Erdmann, 2012),
new combination
Kevin W. Conway1,2*, Kyoji Fujiwara3, Hiroyuki Motomura2,3,4 & Adam P. Summers5,6
Abstract. Erdmannichthys, a new genus of the family Gobiesocidae, is introduced to include a species previously
assigned to Aspasmichthys. This act is supported by morphological characters and the results of a recent molecularphylogenetic study that placed the two currently described species of Aspasmichthys as distant relatives within thesubfamily Diademichthyinae (i.e., polyphyletic). New details on live colouration and skeletal anatomy are provided
for Erdmannichthys alorensis, new combination, based on photographs and one additional specimen collected from
Padar Island, Komodo Group (Indonesia).
Key words. Diademichthyinae, Aspasmichthys, computer tomographic scanning, miniature

INTRODUCTION

clingfishes compiled by Briggs (1955). In a recent molecularphylogenetic study of Gobiesocidae, Conway et al. (2020)
identified Aspasmichthys as one of several non-monophyletic
genera in need of further taxonomic study (together with
“Cochleoceps”, “Lepadichthys” and “Lepadogaster”). In thatstudy, the Japanese/South Korean Aspasmichthys ciconiae
was obtained in a sister group relationship with Flexor incus(Fig. 2), a species known to date from a small number of
specimens collected from Australia (Lord Howe Island), New
Caledonia (Loyalty Islands) and New Zealand (Kermadec
Islands) (Conway et al., 2018; Fujiwara et al., 2021a).
The Indonesian Aspasmichthys alorensis was obtained by
Conway et al. (2020) in a sister group relationship with anundescribed species of Pherallodichthys (Fig. 2).

The genus Aspasmichthys was introduced by Briggs (1955)
to accommodate the East Asian species, Aspasma ciconiae,
described earlier by Jordan & Fowler (1902) based onmaterial from Japan. Aspasmichthys ciconiae differs from its
previous congener Aspasma ubauo (previously A. minima; see
Fujiwara & Motomura, 2019) in a number of morphologicalfeatures, including most notably a longer snout and featuresof adhesive disc papillae, justifying the generic reassignmentof Briggs (1955). Aspasmichthys remained monotypic until,
over sixty years later, Allen & Erdmann (2012) described asecond species, Aspasmichthys alorensis, based on a single
specimen (8.2 mm SL) collected from the Alor Strait,
East Nusa Tenggara Province, Indonesia (Fig. 1). The
assignment of this species to Aspasmichthys was describedas “provisional” by Allen & Erdmann (2012: 1162), whoreached this conclusion based on their interpretation of thecharacters listed in the keys to the subfamilies and genera of

The report of non-monophyly for Aspasmichthys by Conway
et al. (2020) encouraged us to take a closer look at the two
species currently assigned to this genus. Our examination of
specimens revealed a number of morphological differences
between A. ciconiae and A. alorensis, including majordifferences in the configuration of the gill membranes(united to the isthmus vs. free from isthmus) that would haverequired these two taxa to be artificially placed into different
subfamilies following the long-standing classification system
of Briggs (1955) (A. ciconiae to the Aspasminae and A.
alorensis to the Diplocrepinae). Based on our findings andthe results of Conway et al. (2020), we transfer A. alorensis
from Aspasmichthys to a new monotypic genus describedherein. We also take this opportunity to report additional
details of live colouration and skeletal anatomy for A.
alorensis based on additional specimens photographed and/or collected from Padar Island, Indonesia (Fig. 1) in 2016.

Accepted by: Zeehan Jaafar
Department of Ecology and Conservation Biology and Biodiversity Research andTeaching Collections, Texas A&M University, College Station, TX 77843, USA;
Email: kevin.conway@tamu.edu (*corresponding author)
1

Research Associate, Ichthyology, Australian Museum Research Institute, 1 William
Street, Sydney, NSW 2010, Australia
2

The United Graduate School of Agricultural Sciences, Kagoshima University, 1-21-24
Korimoto, Kagoshima 890-0065, Japan
3

4

The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan

Friday Harbor Laboratories, University of Washington, Friday Harbor, Washington
98250
5

Research Associate, Burke Museum of Natural History and Culture, University ofWashington, Seattle, Washington, 98105, USA
6© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

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Fig. 1. Map showing locations (red circles) from which specimens of Erdmannichthys alorensis have been collected.

MATERIAL AND METHODS

canal from anterior to posterior or dorsal to ventral (lachrymal
canal). Osteological terminology used herein follows Springer& Fraser (1976), with modifications of Conway et al. (2018,2019). Collection abbreviations follow Sabaj (2020).

Observations and photographs of specimens or their parts
were generally made using a ZEISS SteReo Discovery V20
stereomicroscope equipped with a Zeiss Axiocam MRc5
digital camera. Images were obtained using the stacking
function in the Zeiss Axiovision software and processed using
a combination of Adobe Photoshop 2020 and Illustrator 2020.
Computed tomography (CT) scans of select specimens wereobtained at the Karel F. Liem BioImaging Center (Friday
Harbor Laboratories, University of Washington) using a
Bruker SkyScan 1173 scanner with a 1 mm aluminium filter.
Scans were run at 55–65 kV and 123–143 μA on a 2048×2048
pixel CCD at a resolution of 6.7–12.4 μm. Specimens were
scanned while inside a 15 ml plastic Falcon tube, in which
they were wrapped with cheesecloth moistened with ethanol(70%) to prevent movement during scanning. The resulting
CT data were visualised using 3D Slicer v.4.10.2 (https://www.slicer.org/) and have been deposited on MorphoSource(https://www.morphosource.org/). Adhesive disc papillae
terminology follows Briggs (1955). Cephalic lateral-line poreterminology follows Shiogaki & Dotsu (1983), except that
we also use numbers to refer to individual pores following
Conway et al. (2017), with pores numbered along a particular

TAXONOMY
Erdmannichthys, new genus
Diagnosis. A genus of the subfamily Diademichthyinae sensu
Conway et al. (2020), distinguished from all other genera ofthis subfamily by the presence of a prominent dark reddishbrown oval to triangular marking at base of caudal fin (Figs.
3, 4). The new genus is distinguished from Diademichthys,
Discotrema, Flabellicauda, “Lepadichthys”, Lepadicyathus
and Unguitrema by having a double adhesive disc (Fig. 5) (vs.
single adhesive disc). It is distinguished from the remaining
diademichthyine genera (viz. Aspasma, Aspasmichthys,
Aspasmodes, Briggsia, Flexor, Liobranchia, Lissonanchus,
Pherallodus, Pherallodichthys and Propherallodus), allwith a double adhesive disc, by the presence of three whitelines on the body surface in life (Figs. 3, 4), including a
mid-dorsal stripe (originating on tip of snout posterior to429

Conway et al.: A new genus of clingfish from Indonesia

Fig. 2. Phylogenetic relationships of Gobiesocidae modified from Conway et al. (2020; fig. 3) showing placement of Aspasmichthys ciconiae(red) and Erdmannichthys alorensis (blue). Numbers above branches represent bootstrap support values (values below 65 not shown).

head in Aspasma, Aspasmichthys, Aspasmodes, Briggsia,
Liobranchia, Lissonanchus and Pherallodichthys); papillae
of disc region A continuous across centre (Fig. 5A, B)(vs. centre of region A without papillae in Aspasma,
Aspasmichthys [Fig. 5C, D] and Pherallodus); mandibular
portion of preoperculo-mandibular lateral-line canal absent(Fig. 6C) (vs. present on anguloarticular in Aspasmichthys);
preopercular portion of preoperculo-mandibular lateral-line
canal with two pores (PR1 and PR3) only (Fig. 6B, C) (vs.
three pores, PR1–3, in Aspasma, Aspasmichthys, Flexor
and Propherallodus, preopercle without canal in Briggsia,
Lissonanchus and Pherallodus); presence of a small oval gap
in dorsal view between anterior tips of premaxillae formed bya semicircular indentation along medial edge of premaxilla(Fig. 6A) (vs. minute square gap in Aspasma, medial edgeof premaxilla straight, without gap between premaxillae inupper lip and extending to dorsal-fin origin) and a pair of
dorsolateral stripes (originating on dorsolateral part of snout
adjacent to anterior nostril and extending posterior, throughupper part of eye, to area on upper body side close to caudal
fin base) (vs. colour pattern in life largely uniform, withsmaller indistinct dark markings [Aspasma, Aspasmodes,
Briggsia, Flexor and Lissonanchus] or with prominent
darker stripes, saddles or other markings [Aspasmichthys,
Pherallodus, Pherallodichthys and Propherallodus] but
without prominent white lines along long axis of body; life
colour unknown for Liobranchia). It is further distinguishedfrom the other diademichthyine genera with a double
adhesive disc by the following combination of characters:
gill membranes united and free from isthmus forming single
continuous gill opening (Figs. 4, 5A) (vs. gill membranes
connected to isthmus restricting gill opening to side of430

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Fig. 3. Five individuals of Erdmannichthys alorensis photographed by M.V. Erdmann at a submerged reef (10 metres depth) off south
east coast of Padar island, Nusa Tenggara Province, Indonesia. A–C. Not collected, not measured. D. Potentially WAM P.34629-001,
13.3 mm SL; E–F. Potentially WAM P.34629-005, 9.1 mm SL.

Aspasmodes, Liobranchia and Pherallodichthys); subopercle
poorly ossified, posterior margin rounded (Fig. 6B, C) (vs.
subopercle well ossified, posterior edge with spine-likeposterior process in Aspasmichthys).

org/10.17602/M2/M388066); Indonesia: Nusa Tenggara
Province: submerged reef off south east coast of Padar island,
Komodo Group, 10 metres depth, 08°41.284′S, 119°34.528′E,
M.V. Erdmann and A. Sianipar (MVE-16-051).

Etymology. Named for Mark V. Erdmann, in honour of his
work on Indo-Pacific reef fishes. Gender masculine.

Notes on Erdmannichthys alorensis, new combination.
Allen & Erdmann (2012) described Erdmannichthys alorensis
based on a single individual (MZB 20588, holotype, 8.2
mm SL) collected in 2011 from a submerged reef on thewestern side of Alor Island (Nusa Tenggara Province) (Fig.
1). The small size of this specimen presented challenges tothe examination of its features, including the configurationof the gill membranes, which are free from the isthmus(Figs. 4, 5A) but were interpreted as fused to the isthmus by

Type species. Aspasmichthys alorensis Allen & Erdmann,
2012, by present designation.
Material examined. WAM P.34629-001, 1 (photographonly), 13.3 mm SL; WAM P.34629-005, 1 (preservedin ethanol [DNA voucher]), 9.1 mm SL (CT, https://doi.
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Fig. 4. Erdmannichthys alorensis, WAM P.34629-005, 9.1 mm SL; Indonesia, Nusa Tenggara Province. Dashed white circles highlight
location of cephalic lateral line canal pores. Abbreviations: LC1, 2, lachrymal pore 1, 2; NC1, 2, nasal pore 1, 2; PO1, 2, postorbital pore
1, 2; PR1, 3, preopercular pore 1, 3.

features of the live colour pattern of E. alorensis not reported
in the original description, including the presence of a black
marking on the posterior part of the dorsal fin in one of theindividuals photographed (Fig. 3C), and the presence of darkbrown melanocytes on the lower abdomen of another (Fig.
3F; potentially WAM P.34629-005). Dark brown melanocytes
are also visible over the surface of the head, around the anus,
and the surface of the adhesive disc in a 9.1 mm SL specimen(WAM P.34629-005; Figs. 4, 5) and are also visible on thesurface of the adhesive disc in the 8.2 mm SL holotype (Allen& Erdmann, 2012: 1163; fig. 5). Dark brown melanocytes
are common in larval and juvenile stages of gobiesocids(reviewed by Allen, 1984) but are typically absent from adultstages. The largest specimen of E. alorensis recorded to date

Allen & Erdmann (2012). This misinterpretation of a single
character lead Allen & Erdmann (2012) to assign their new
species to the subfamily Aspasminae (sensu Briggs, 1955, in
which the gill membranes are fused to the isthmus) insteadof the Diplocrepinae (sensu Briggs, 1955, in which the gillmembranes are free), and ultimately generic assignment to
Aspasmichthys. The sole specimen of E. alorensis available
to Allen & Erdmann (2012) also precluded the descriptionof internal characters, including features of the skeleton.
In 2016, five additional specimens of E. alorensis were
photographed by M. V. Erdmann off the eastern coast of
Padar Island (also Nusa Tenggara Province). The resulting
series of underwater photographs (Fig. 3) reveals additional
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Fig. 5. A. Photograph of adhesive disc of Erdmannichthys alorensis, WAM P.34629-005, 9.1 mm SL; anterior margin of disc inserted
under gill membrane; dashed grey line highlights posterior margin of gill membrane. B. Schematic line drawing of adhesive disc in A.
C. Photograph of adhesive disc of Aspasmichthys ciconiae, KAUM–I. 147568, 43.1 mm SL; disc treated with Cyanine blue (following
Saruwatari et al., 1997). D. Schematic line drawing of disc in C. Abbreviations: A–C, adhesive disc regions A–C; GM, gill membrane;
LM, larval melanocyte.

E. alorensis is a small-bodied species (i.e., miniature sensu
Weitzman & Vari, 1988) and the presence of melanocytes
could represent a paedomorphic character. Examination of
additional specimens will be needed to confirm both size atmaturity and maximum size of E. alorensis.

is only 13.3 mm SL (WAM P.34629-001). If melanocytes
are lost ontogenetically in E. alorensis, as they appear to
be in other gobiesocids, the presence of melanocytes in the
aforementioned specimens may suggest that these individuals
are juveniles, and perhaps only recently settled on the reef
prior to collection. An alternative explanation could be that433

Conway et al.: A new genus of clingfish from Indonesia

Fig. 6. CT scanned anterior skeleton, including cranium, paired-fin girdles and abdominal region of vertebral column, of Erdmannichthys
alorensis WAM P.34629-005, 9.1 mm SL. A. Dorsal view. B. Lateral view of right side (image mirrored). C. Ventral view. Right side of
specimen damaged. Cephalic lateral line canals highlighted in green. Pores of cephalic lateral line system orange. Anteriormost epicentral(epicentral 2) highlighted in aqua blue. Dorsal postcleithra highlighted in pink. Ventral postcleithra highlighted in yellow. Outline ofsubopercle highlighted with thin white line in B, C. Abbreviations: ACh, anterior ceratohyal; Ana, angulo-articular; Apa, autopalatine;
Boc, basioccipital; Bp, basipterygium; Br, branchiostegal rays; Cb, ceratobranchials; Cl, cleithrum; Cor, coracoid; DHh, dorsal hypohyal;
DPcl, dorsal postcleithrum; Ec2, 6, epicentral 2, 6; Ect, ectopterygoid; EPC, epiphyseal commissure; Epoc, epiotic; Exoc, exoccipital; Fr,
frontal; Hy, hyomandibular; I, pelvic-fin spine; Iop, interopercle; LC, lachrymal canal; LC1, 2, lachrymal pore 1, 2; LE, lateral ethmoid;Na, nasal; NC1, 2, nasal pore 1, 2; NS1, 5, neural spine of vertebral centrum 1, 5; Op, opercle; Pa, parietal; PecR, pectoral radial; PecFR,
pectoral-fin ray; PelFR1–4, pelvic-fin ray 1–4; PO1, 2, postorbital pore 1, 2; Pop, preopercle; PRC, preopercular canal; PR1, 3, preopercular
canal pore 1, 3; Pro, prootic; Psph, parasphenoid; Pt, posttemporal; Pte, pterotic; Q, quadrate; Ra, retroarticular; Ri, ribs; Sc, scapula; Scl,
supracleithrum; Soc, supraoccipital; Sop, subopercle; SPC, supraorbital canal; Sph, sphenotic; STC, stomach content; Ur, urohyal; V1,
vertebral centrum 1; Vo, vomer; VPcl, ventral postcleithrum.

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Fig. 7. CT scanned skeleton of Erdmannichthys alorensis WAM P.34629-005, 9.1 mm SL. A. Dorsal view. B. Lateral view. C. Ventral
view. D. Caudal-fin skeleton, lateral view. Right side of specimen damaged. Abbreviations: AFR, anal-fin rays; DFR, dorsal-fin rays;
DPCR, dorsal procurrent rays; HS1, hemal spine 1; NS1, 10, neural spine 1, 10; PCR, principal caudal-fin rays; PU2, preural centrum 2;
VPCR, ventral procurrent rays.

Two of the five specimens photographed by M. V. Erdmann(Fig. 3) were collected and we have had the opportunity to
examine and also CT scan one of these (WAM P.34629005, 9.1 mm SL; Fig. 4). From the rendering of the CT scan(Figs. 6, 7) we are able to report the following osteologicalinformation: Dorsal-fin rays 7. Anal-fin rays 7. Total
caudal-fin rays 20 (17 reported by Allen & Erdmann, 2012),
including 5+5 principal rays, 5 dorsal procurrent rays, and5 ventral procurrent rays (anteriormost procurrent rays arepoorly ossified and difficult to observe in the scan rendering
presented in Fig. 7D). Pectoral-fin rays 24 (21 reported byAllen & Erdmann, 2012). Branchiostegal rays 6. Total numberof vertebrae 32, comprising 15 abdominal and 17 caudal.
Ribs 12, associated with vertebrae 3–14. Epicentrals 14,
associated with vertebrae 2–15. The anteriormost epicentral(epicentral 2) is a thin rod (Fig. 6).

also visible lateral to the functional teeth on the outer marginof the premaxilla (Fig. 6C, D). A single row of ca. 9 small
conical teeth, each with slightly recurved tip, is present along
the dorsal ridge of the dentary on each side of the lower jaw.
A small oval opening is present between the premaxillae,
formed by a semicircular indentation along the medial edgeof each premaxilla (Fig. 6A). Ceratobranchials 1–4 arerepresented by poorly ossified rods of bone. Ceratobranchial
5 is approximately half the length of ceratobanchial 4. Other
elements of the gill arch skeleton could not be observed in
the available scan.
The cephalic lateral-line system comprises the following:supraorbital canal on nasal (pores NC1, NC2) and frontal
bone; infraorbital canal on lachrymal (LC1, 2); otic canalon sphenotic (PO1 at junction of frontal and sphenotic) andpterotic (PO2); and preopercular canal on preopercle (PR1,
PR3). The second pore of the preopercular canal (PR2)present in most gobiesocids (e.g., see Shiogaki & Dotsu,
1983) is absent in E. alorensis. The subopercle of E. alorensis
is poorly ossified and difficult to detect in the rendering of
the CT scan (Fig. 6B, C). The same is likely the case for

A single row of teeth is present along the outer margin of thepremaxilla, comprising ca. 4 slightly recurved conical teeth
anteriorly at and adjacent to symphysis, and ca. 12 strongly
laterally compressed incisiviform teeth, each with strongly
recurved cusp. Several replacement incisiviform teeth are435

Conway et al.: A new genus of clingfish from Indonesia
MA655401, 2, 18.0–24.0 mm SL; AIM MA655316, 1, 23.0mm SL (CT; https://doi.org/10.17602/M2/M56344); AMS
I.45807-001, 1, 19.7 mm SL; AMS I.45823-010, 1, 19.1 mm
SL; NMNZ P.060717, 1, holotype, 20.8 mm SL; NMNZ
P.049965, 1, 24.3 mm SL; NMNZ P.049966, 1, 27.2 mm
SL; NMNZ P.017760, 1, 14.0 mm SL; NMNZ P.041114,
1, 22.4 mm SL; NMNZ P.024500, 1, not measured; NMNZ
P.025315, 1, 19.4 mm SL; NMNZ P.028570, 1, 27.2 mm
SL; NMNZ P.029570, 1, 27.1 mm SL; NMNZ P.050069,
1, 21.1 mm; New Zealand, Kermadec Islands. — AMS
I.17424-003, 1, 20.8 mm SL; Australia, Lord Howe Island.
— USNM 322448, 1, 29.1 mm SL; New Caledonia.
Liobranchia stria: USNM 113992, 1, 15.3 mm SL; MarshallIslands.
Lissonanchus lusheri: SAIAB 342, holotype, 19.1 mm SL;
Mozambique.
Pherallodichthys meshimaensis: KAUM–I. 131060, 25.7mm SL; Japan, Yaku-shima island. — NSMT-P 46752,
holotype, 17.6 mm SL; NSMT-P 46753, paratype (C&S),
19.2 mm SL; Japan, Kyushu. — NSMT-P 72561, 1, 13.0mm SL; Indonesia, Ambon. — ROM 74264, 1, 12.5 mm
SL; Vietnam. — ROM 55170, 1, 18.6 mm SL; Philippines.
Pherallodus indicus: NSMT-P 114703, 1, 25.0 mm SL(CT; https://doi.org/10.17602/M2/M56337); Japan, Honshu.
— TCWC 16462.01, 2, 16.5–18.1 mm SL; Japan, Kochi.
Propherallodus briggsi: NSMT-P 46749, holotype, 20.8mm SL; Japan, Kyusyu.
Propherallodus longipterus: ANSP 193493, 1, 22.3 mm
SL; India, Kerala — KUAM–I. 114615, paratype, 17.8mm; Philippines.
Propherallodus smithi: CAS 131349, holotype, 18.5 mm
SL; South Africa, Durban.

the mesethmoid and the central part of the frontal, neitherof which are visible in the optimal rendering of the scan.
The dorsal postcleithrum of the left side is a shield-shapedbone (Fig. 6). The dorsal postcleithrum of the right sideis severed into two pieces in the individual scanned (Fig.
6). The ventral postcleithrum exhibits a facet on theanteromedial edge which accommodates the posterior tipof the basipterygium. The lateral margin of the ventral
postcleithrum is weakly serrated, as is common in otherdiademichthyine taxa (e.g., see Hayashi et al., 1986: fig.
21). No poorly ossified fimbrae could be observed along theposterior edge of either the dorsal or ventral postcleithra in
the renderings of the CT scan.
The new information presented herein on Erdmannichthys
alorensis, though scant, complements that available from theoriginal description provided by Allen & Erdmann (2012).
Importantly, we have also been able to confirm that E.
alorensis exhibits all three of the morphological charactersproposed as putative synapomorphies for a monophyletic
group of diademichthyine clingfishes (viz. Aspasmichthys,
Diademichthys, Flexor, Flabellicauda, “Lepadichthys”,
Lepadicyathus, Pherallodichthys and Pherallodus) by
Conway et al. (2018, 2020) and Fujiwara et al. (2021b),
including: (1) a unique type of laterally compressed
incisiviform tooth with a hooked tip; (2) an oval opening(variable in size) visible in dorsal view at the tip of thepremaxillae, formed by a semicircular indentation along
the medial edge of each premaxilla; and (3) a complex
articulation between the posterior tip of the basipterygium
and the anteromedial edge of the ventral postcleithrum. Basedon the aforementioned characters, and the results of Conway
et al. (2020) (Fig. 2), we are confident that Erdmannichthys
also is a member of this putative monophyletic group.
The results of Conway et al. (2020) suggest that within
this larger group, Erdmannichthys is a close relative of
Pherallodichthys (Fig. 2). We are not currently awareof any morphological characters that could potentiallyrepresent synapomorphies in support of a clade including
Erdmannichthys and Pherallodichthys and the comparative
work to identify such characters, and refine relationships
among the diademichthyine clingfishes, is ongoing.

ACKNOWLEDGEMENTS
We thank M.V. Erdmann for generously sharing his
underwater photographs and knowledge of E. alorensis andfor organising access to material. Thanks also to T. Trnski
and S. Hannam (AIM), M. McGrouther and A. Hay (AMS),
M. Sabaj (ANSP), D. Catania and M. Hoang (CAS), A.
Stewart (NMNZ), G. Shinohara (NSMT), E. Holm (ROM),
R. Bills, O. Gon, M. Dwani, and E. Heemstra (SAIAB), J.
Williams, S. Raredon, K. Murphy, and D. Pitassy (USNM)
and M. Allen and G. Moore (WAM) for granting accessto material under their care. We also thank J. Huie fordouble checking fin ray counts, and R. Britz and G. Allenfor providing critical comments on an earlier draft of themanuscript. This study was supported in part by JSPS
KAKENHI Grant Numbers 20H03311 and 21H03651; theJSPS Core-to-core CREPSUM JPJSCCB2020009; and the“Establishment of Research and Education Network in the
Amami Islands” project of Kagoshima University adopted
by the Ministry of Education, Culture, Sports, Science andTechnology, Japan. CT scanning was possible via funding
from NSF (DBI 1702442 to KWC; DBI 1701665 to A.
Summers). KWC acknowledges the financial support ofTexas A&M Agrilife Research (TEX09452). KF was funded
by a Grant-in-Aid from the Japan Society for the Promotionof Science for JSPS Fellows (DC1: 19J21103). This isComparative material examined
Aspasma ubauo: KAUM–I. 128774, holotype, 32.6 mm SL;
NSMT-P 114699, 1, 29.9 mm SL; Japan, Honshu.
Aspasmichthys ciconiae: CAS-SU 7136, syntype, 49.9 mm
SL; USNM 50760, syntype, 56.0 mm SL; Japan, Honshu.
— KAUM–I. 147708, 1, 26.7 mm SL; KAUM–I. 147568,
1, 43.1 mm SL; Japan, Kyushu. — TCWC 16461.02, 1,
26.0 mm SL (CT; https://doi.org/10.17602/M2/M30821);
Japan, Kochi.
Aspasmodes briggsi: ANSP 150414, 1, 17.1 mm SL; ANSP
150415, 5 (1C&S), 7.9–15.1 mm SL; Seychelles. — ROM
55927, 1, 6.6 mm SL; Chagos Archipelago.
Briggsia hastingsi: Information on this species obtained
from Craig & Randall (2009).
Flabellicauda: See material listed in Fujiwara et al. (2021b).
Flexor incus: AIM MA655142, 1 (C&S), 20.0 mm SL; AIM
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