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Review of the Lepadichthys lineatus complex (Gobiesocidae: Diademichthyinae) with descriptions of three new species

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Journal of Fish Biology
Authors:
Kyoji Fujiwara at National Museum of Nature and Science
Kyoji Fujiwara
Hiroyuki Motomura at Kagoshima University Museum
Hiroyuki Motomura
  • Kagoshima University Museum
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Abstract and Figures

The Lepadichthys lineatus complex (Gobiesocidae: Diademichthyinae) is defined by three unique characters within Lepadichthys: (a) upper‐jaw lip fused with snout skin, usually lacking a distinct groove between the dorsal lip margin and snout (if present, very weak, restricted to posterior portion of jaw); (b) snout tip well extended, distinctly beyond lower‐jaw tip; and (c) inner surface of both lips with oral papillae. A taxonomic review of the complex recognized four valid species: Lepadichthys geminus sp. nov. (southern Japan and Indonesia), Lepadichthys heemstraorum sp. nov. (southwestern Indian Ocean), Lepadichthys polyastrous sp. nov. (southwestern Indian Ocean) and L. lineatus Briggs, 1966 (Red Sea, Arabian Sea, Seychelles and Sri Lanka). L. geminus and L. lineatus are distinct from L. heemstraorum and L. polyastrous in having a circular (vs. elliptical) disc and more posteriorly located anus [L. geminus and L. lineatus with disc length and width 15.0–18.7 (mean 16.9) and 12.9–16.5 (14.6) % LS, respectively, and length to width ratio 1.03–1.25 cf. L. heemstraorum and L. polyastrous, 17.0–21.5 (18.9) and 11.6–15.2 (13.0) % LS, respectively, and 1.26–1.61; pre‐anus length and disc to anus length 65.1–73.6 (68.7) and 25.7–31.6 (28.6) % LS, respectively vs. 60.2–68.3 (65.3) and 21.6–28.9 (25.5) % LS, respectively]. Body depth (as % of LS) is also useful to distinguish L. geminus and L. polyastrous from L. heemstraorum and L. lineatus [viz., 12.7–16.1 (14.4) in L. geminus and 10.8–14.9 (13.1) in L. polyastrous vs. 15.0–17.1 (15.9) in L. heemstraorum and 14.6–18.9 (16.8) in L. lineatus]. L. geminus differs distinctly from other species in the complex as follows: snout tip directed upward, usually on same horizontal level with lower margin of eye lens (lateral view) (vs. directed somewhat downward, horizontal level usually between lower margins of eye and eye lens in L. heemstraorum and L. lineatus, lower margin of eye in L. polyastrous); and lower abdomen with two yellow stripes (vs. a single stripe along ventral midline in L. polyastrous and L. lineatus, unknown in L. heemstraorum). L. polyastrous has unique patterns of yellow dots on the dorsal and ventral body surfaces, forming c. six to eight and three to five longitudinal rows, respectively [vs. usually forming c. three to five longitudinal rows and a single broken line, respectively, in L. geminus and L. lineatus; yellow dots usually absent in L. heemstraorum]. A poorly known species, Lepadichthys caritus Briggs, 1969, is regarded as a junior synonym of L. lineatus.
Anterior portion of snout of (a, b) Lepadichthys geminus sp. nov. (KAUM–I. 145207, 21.2 mm LS), (c, d) Lepadichthys polyastrous sp. nov. (SAIAB 69396, 18.7 mm LS), (e, f) Lepadichthys misakius (KAUM–I. 40010, 22.6 mm LS) and (g, h) Discotrema crinophilum (KAUM–I. 145204, 18.3 mm LS), showing oral features (stained with cyanine blue). (a, e, g) dorsal views; (b, f, h) lateral views; (c) and (d) inner surface of upper‐ and lower‐jaw lips, respectively. Red arrows indicate oral papillae. Yellow arrows indicate groove between upper‐jaw lip and snout. Orange and green broken lines indicate tips of snout and lower jaw, respectively. () oral papillae and () groove between upper‐jaw lip and snout
Anterior portion of snout of (a, b) Lepadichthys geminus sp. nov. (KAUM–I. 145207, 21.2 mm LS), (c, d) Lepadichthys polyastrous sp. nov. (SAIAB 69396, 18.7 mm LS), (e, f) Lepadichthys misakius (KAUM–I. 40010, 22.6 mm LS) and (g, h) Discotrema crinophilum (KAUM–I. 145204, 18.3 mm LS), showing oral features (stained with cyanine blue). (a, e, g) dorsal views; (b, f, h) lateral views; (c) and (d) inner surface of upper‐ and lower‐jaw lips, respectively. Red arrows indicate oral papillae. Yellow arrows indicate groove between upper‐jaw lip and snout. Orange and green broken lines indicate tips of snout and lower jaw, respectively. () oral papillae and () groove between upper‐jaw lip and snout
… 
Fresh (a–c) and preserved (d–h) holotype of Lepadichthys geminus sp. nov. (KAUM–I. 145214, 22.3 mm LS, Okinoerabu Island, Amami Islands, Japan). (a, d) lateral view; (b, e) dorsal view; (c, f) ventral view; (g) and (h) lateral and dorsal views of head, respectively. Red broken line indicates horizontal level of snout tip. Blue and orange triangles indicate lower margins of eye and eye lens, respectively. Abbreviations: AN and PN: anterior and posterior nostrils, respectively; LC: lacrimal canal pores; NC: nasal canal pores; PO: postorbital canal pores
Fresh (a–c) and preserved (d–h) holotype of Lepadichthys geminus sp. nov. (KAUM–I. 145214, 22.3 mm LS, Okinoerabu Island, Amami Islands, Japan). (a, d) lateral view; (b, e) dorsal view; (c, f) ventral view; (g) and (h) lateral and dorsal views of head, respectively. Red broken line indicates horizontal level of snout tip. Blue and orange triangles indicate lower margins of eye and eye lens, respectively. Abbreviations: AN and PN: anterior and posterior nostrils, respectively; LC: lacrimal canal pores; NC: nasal canal pores; PO: postorbital canal pores
… 
Adhesive discs and disc papillae of species of the Lepadichthys lineatus complex and Discotrema (stained with cyanine blue). (a) Lepadichthys geminus sp. nov. (KAUM–I. 145214, holotype, 22.3 mm LS); (b) Lepadichthys heemstraorum sp. nov. (USNM 260552, 26.2 mm LS); (c) Lepadichthys polyastrous sp. nov. (SAIAB 70904, holotype, 27.7 mm LS); (d) L. lineatus (USNM 360166, 27.8 mm LS); (e) Discotrema crinophilum (KAUM–I. 128566, 40.0 mm LS); (f) Discotrema monogrammum (BPBM 40504, paratype, 16.8 mm LS)
Adhesive discs and disc papillae of species of the Lepadichthys lineatus complex and Discotrema (stained with cyanine blue). (a) Lepadichthys geminus sp. nov. (KAUM–I. 145214, holotype, 22.3 mm LS); (b) Lepadichthys heemstraorum sp. nov. (USNM 260552, 26.2 mm LS); (c) Lepadichthys polyastrous sp. nov. (SAIAB 70904, holotype, 27.7 mm LS); (d) L. lineatus (USNM 360166, 27.8 mm LS); (e) Discotrema crinophilum (KAUM–I. 128566, 40.0 mm LS); (f) Discotrema monogrammum (BPBM 40504, paratype, 16.8 mm LS)
… 
Lateral view of heads of (a–e) Lepadichthys geminus sp. nov., (f–j) Lepadichthys heemstraorum sp. nov., (k–o) Lepadichthys polyastrous sp. nov. and (p–t) Lepadichthys lineatus. Red broken line indicates horizontal level of snout tip. Blue and orange triangles indicate lower margins of eye and eye lens, respectively
Lateral view of heads of (a–e) Lepadichthys geminus sp. nov., (f–j) Lepadichthys heemstraorum sp. nov., (k–o) Lepadichthys polyastrous sp. nov. and (p–t) Lepadichthys lineatus. Red broken line indicates horizontal level of snout tip. Blue and orange triangles indicate lower margins of eye and eye lens, respectively
… 
Fresh colouration of (a–g, red circles) Lepadichthys geminus sp. nov., (h, orange diamond) Lepadichthys heemstraorum sp. nov., (i–m, blue squares) Lepadichthys polyastrous sp. nov. and (n, green triangle) Lepadichthys lineatus. (a, b) BSKU 55004, 27.0 mm LS (photos by BSKU); (c, d) KAUM–I. 145208, 23.2 mm LS; (e, f) KAUM–I. 145215, 16.5 mm LS; (g) KAUM–I. 128500, 22.0 mm LS; (h) specimen (from South Africa) not available for this study (photo by J, Randall); (i) SAIAB 63007, 30.9 mm LS, (j) 1 of ROM 72740 (photo by R. Winterbottom); (k) SAIAB 59104, 23.0 mm LS; (l, m) SAIAB 43241, 22.8 mm LS; (n) SAIAB 79365, 21.9 mm LS. (a, c, e, g, h, i, l, n) lateral views; (b, d, j, k) dorsal views; (f, m) ventral views. (h–k, m) Photos by SAIAB. () L. geminus sp. nov., () L. heemstraorum sp. nov., () L. polyastrous sp. nov. and () L. lineatus
+6
Fresh colouration of (a–g, red circles) Lepadichthys geminus sp. nov., (h, orange diamond) Lepadichthys heemstraorum sp. nov., (i–m, blue squares) Lepadichthys polyastrous sp. nov. and (n, green triangle) Lepadichthys lineatus. (a, b) BSKU 55004, 27.0 mm LS (photos by BSKU); (c, d) KAUM–I. 145208, 23.2 mm LS; (e, f) KAUM–I. 145215, 16.5 mm LS; (g) KAUM–I. 128500, 22.0 mm LS; (h) specimen (from South Africa) not available for this study (photo by J, Randall); (i) SAIAB 63007, 30.9 mm LS, (j) 1 of ROM 72740 (photo by R. Winterbottom); (k) SAIAB 59104, 23.0 mm LS; (l, m) SAIAB 43241, 22.8 mm LS; (n) SAIAB 79365, 21.9 mm LS. (a, c, e, g, h, i, l, n) lateral views; (b, d, j, k) dorsal views; (f, m) ventral views. (h–k, m) Photos by SAIAB. () L. geminus sp. nov., () L. heemstraorum sp. nov., () L. polyastrous sp. nov. and () L. lineatus
… 
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REGULAR PAPER
Review of the Lepadichthys lineatus complex (Gobiesocidae:
Diademichthyinae) with descriptions of three new species
Kyoji Fujiwara
1
| Hiroyuki Motomura
2
1
The United Graduate School of Agricultural
Sciences, Kagoshima University, Kagoshima,
Japan
2
The Kagoshima University Museum,
Kagoshima, Japan
Correspondence
Kyoji Fujiwara, The United Graduate School of
Agricultural Sciences, Kagoshima University,
1-21-24 Korimoto, Kagoshima 890-0065,
Japan.
Email: k2043960@kadai.jp
Abstract
The Lepadichthys lineatus complex (Gobiesocidae: Diademichthyinae) is defined by three
unique characters within Lepadichthys: (a) upper-jaw lip fused with snout skin, usually lac-
king a distinct groove between the dorsal lip margin and snout (if present, very weak,
restricted to posterior portion of jaw); (b) snout tip well extended, distinctly beyond
lower-jaw tip; and (c) inner surface of both lips with oral papillae. A taxonomic review of
the complex recognized four valid species: Lepadichthys geminus sp. nov. (southern Japan
and Indonesia), Lepadichthys heemstraorum sp. nov. (southwestern Indian Ocean),
Lepadichthys polyastrous sp. nov. (southwestern Indian Ocean) and L. lineatus Briggs,
1966 (Red Sea, Arabian Sea, Seychelles and Sri Lanka). L. geminus and L. lineatus are dis-
tinct from L. heemstraorum and L. polyastrous in having a circular (vs. elliptical) disc and
more posteriorly located anus [L. geminus and L. lineatus with disc length and width 15.0–
18.7 (mean 16.9) and 12.9–16.5 (14.6) % L
S
, respectively, and length to width ratio 1.03–
1.25 cf. L. heemstraorum and L. polyastrous,17.0–21.5 (18.9) and 11.6–15.2 (13.0) % L
S
,
respectively, and 1.26–1.61; pre-anus length and disc to anus length 65.1–73.6 (68.7)
and 25.7–31.6 (28.6) % L
S
, respectively vs.60.2–68.3 (65.3) and 21.6–28.9 (25.5) % L
S
,
respectively]. Body depth (as % of L
S
)isalsousefultodistinguishL. geminus and
L. polyastrous from L. heemstraorum and L. lineatus [viz., 12.7–16.1 (14.4) in L. geminus and
10.8–14.9 (13.1) in L. polyastrous vs.15.0–17.1 (15.9) in L. heemstraorum and 14.6–18.9
(16.8) in L. lineatus]. L. geminus differs distinctly from other species in the complex as fol-
lows: snout tip directed upward, usually on same horizontal level with lower margin of
eye lens (lateral view) (vs. directed somewhat downward, horizontal level usually between
lower margins of eye and eye lens in L. heemstraorum and L. lineatus, lower margin of eye
in L. polyastrous); and lower abdomen with two yellow stripes (vs. a single stripe along
ventral midline in L. polyastrous and L. lineatus, unknown in L. heemstraorum).
L. polyastrous has unique patterns of yellow dots on the dorsal and ventral body surfaces,
forming c. six to eight and three to five longitudinal rows, respectively [vs. usually forming
c. three to five longitudinal rows and a single broken line, respectively, in L. geminus and
L. lineatus; yellow dots usually absent in L. heemstraorum]. A poorly known species,
Lepadichthys caritus Briggs, 1969, is regarded as a junior synonym of L. lineatus.
KEYWORDS
cryptobenthic fish, Indo-West Pacific, Lepadichthys caritus, morphology, taxonomy
Received: 28 June 2021 Accepted: 30 September 2021
DOI: 10.1111/jfb.14919
FISH
62 © 2021 Fisheries Society of the British Isles. J Fish Biol. 2022;100:62–81.wileyonlinelibrary.com/journal/jfb
... The genus Rhinolepadichthys (previously known as Lepadichthys lineatus complex; see Fujiwara and Motomura, 2021), including four valid species, e.g., Rhinolepadichthys lineatus (Briggs, 1966), Rhinolepadichthys geminus (Fujiwara and Motomura, 2021), Rhinolepadichthys heemstraorum (Fujiwara and Motomura, 2021), and Rhinolepadichthys polyastrous (Fujiwara and Motomura, 2021), was recently defined by Fujiwara et al. (2024). The genus is most similar to the genus Discotrema Briggs, 1976 in having the presence of a hardened cap on the surface of at least some disc papillae and the anterolateral part of the ventral postcleithrum extended anteriorly as a well-developed rod-like process (Fujiwara et al., 2024). ...
... The genus Rhinolepadichthys (previously known as Lepadichthys lineatus complex; see Fujiwara and Motomura, 2021), including four valid species, e.g., Rhinolepadichthys lineatus (Briggs, 1966), Rhinolepadichthys geminus (Fujiwara and Motomura, 2021), Rhinolepadichthys heemstraorum (Fujiwara and Motomura, 2021), and Rhinolepadichthys polyastrous (Fujiwara and Motomura, 2021), was recently defined by Fujiwara et al. (2024). The genus is most similar to the genus Discotrema Briggs, 1976 in having the presence of a hardened cap on the surface of at least some disc papillae and the anterolateral part of the ventral postcleithrum extended anteriorly as a well-developed rod-like process (Fujiwara et al., 2024). ...
... The genus Rhinolepadichthys (previously known as Lepadichthys lineatus complex; see Fujiwara and Motomura, 2021), including four valid species, e.g., Rhinolepadichthys lineatus (Briggs, 1966), Rhinolepadichthys geminus (Fujiwara and Motomura, 2021), Rhinolepadichthys heemstraorum (Fujiwara and Motomura, 2021), and Rhinolepadichthys polyastrous (Fujiwara and Motomura, 2021), was recently defined by Fujiwara et al. (2024). The genus is most similar to the genus Discotrema Briggs, 1976 in having the presence of a hardened cap on the surface of at least some disc papillae and the anterolateral part of the ventral postcleithrum extended anteriorly as a well-developed rod-like process (Fujiwara et al., 2024). ...
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Full-text available
  • Oct 2020
Assembled here is a reasonably complete list of annotated codes for historical and modern natural history collections associated with lost and extant specimens of fossil and Recent fishes, amphibians, and reptiles. A total of 3,845 codes are anchored to about 2,064 distinct collections and/or institutions in 155 countries. At least 633 of those collections are exclusively paleontological or include fossil specimens. The list is primarily derived from the scientific literature and may serve as a resource for plainly citing specimens in publications and for linking such citations to records in online databases. © 2020 by the American Society of Ichthyologists and Herpetologists
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Coastal fish diversity of the Socotra Archipelago, Yemen
Article
Full-text available
  • Jul 2019
The Socotra Archipelago, located in the eastern Gulf of Aden, has a unique marine environment which combines tropical and ‘pseudo-temperate’ elements. Studies on the fish biogeography of the archipelago, partially framed in regional studies, have substantially outpaced critical elementary research on the archipelago’s fish diversity. The present study seeks to close this gap and identifies the Socotra Archipelago as a major hotspot of coastal fish diversity in the Indian Ocean. The archipelago supports unique coastal fish assemblages which are predominantly composed of coral-associated (“reef”) species, in spite of the limited biogenic reef frameworks. A Preliminary Checklist comprises 682 species with confirmed records and a “Working List” includes an additional 51 records, totalling 733 faunal records in 108 families. The family Labridae is the most speciose, followed by Gobiidae, Pomacentridae, Serranidae and Chaetodontidae. The species richness of the archipelago is the highest when compared to adjacent Arabian ecoregions. The richness of the Acanthuridae, Chaetodontidae, Labridae, Pomacentridae and Pseudochromidae stand out as particularily high, and the richness of several families is as high as or higher than in the entire Red Sea. The total archipelagic richness is extrapolated at up to 875 species based on incidence-based richness models and expert opinion. Inshore fish inventories, covering 497 species, found between 14 and 132 species per site (x̄ = 66). Site diversity decreased across the archipelago from west to east and from north to south. Total fish diversity was highest around Socotra Island, followed by Abd al-Kuri & Kal Farun and Darsa & Samha. Occurrence frequencies were very unevenly distributed and dominated by Pomacentrus caeruleus and Thalassoma lunare, whilst many species were infrequent. The fish assemblages are dominated by species from the Indo-West Pacific and the north-western Indian Ocean. The assemblages are rich in rare species and hybrids, and include a low number of endemics (4–5), and a high number of species with far-reaching and Western Indian Ocean ranges.
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Redescription of Lepadichthys coccinotaenia Regan 1921 and description of Lepadichthys trishula sp. nov. from southern Japan (Gobiesocidae: Diademichthyinae)
Article
  • Feb 2020
Lepadichthys coccinotaenia Regan 1921 (Gobiesocidae: Diademichthyinae) is redescribed on the basis of 60 specimens, including the holotype, and a new species of Lepadichthys, previously misidentified as L. coccinotaenia, described from Japan. Lepadichthys trishula sp. nov. is characterized by the following combination of characters: 10–12 (modally 11) dorsal-fin rays; 9–11 (10) anal-fin rays; 23–26 (23) pectoral-fin rays; upper end of gill membrane level with base of 8th to 10th (usually 10th) pectoral-fin ray in lateral view; 6–8 (7) gill rakers on each arch; 32 or 33 (32) vertebrae; snout somewhat short and slender, its length and depth 7.4–9.4 (mean 8.4) and 4.0–6.5 (5.1) % SL, respectively; upper jaw not sharply pointed or projected, its length 5.5–7.7 (6.5) % SL; side of lower-jaw teeth with hook-like tips; head depth 7.2–10.4 (8.5) % SL; disc size moderate, its length 14.0–18.2 (16.8) % SL; dorsal and anal fins with weak membranous connections to caudal fin; head sensory canal pores well developed, including 2 nasal, lacrimal and postorbital, and 3 preopercular pores; and three narrow white stripes from snout to vertical through posterior margin of disc, including two lateral and one on the dorsal midline. The new species is currently known only from Japan, whereas L. coccinotaenia occurs only in the southwestern Indian Ocean.
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An annotated checklist of fishes of Amami-oshima Island, the Ryukyu Islands, Japan.
Article
  • Mar 2018
A comprehensive list of fishes from Amami-oshima Island, the Ryukyu Islands, Japan, is reported for the first time on the basis of collected specimens and literature surveys. A total of 1615 species (618 genera, 175 families and 35 orders) are recorded with specimen registration numbers (if present), localities and literature references. Preliminary comparisons of fish faunas of the island with those of four island regions (Kashiwa-jima Island, the Ogasa wara Islands, Okinawa-jima Island and Yaku-shima Island) in southern Japan show that the fish fauna of Amamioshima Island is most similar to that of Okinawa-jima Island.
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Validity of Lepadichthys misakius (Tanaka 1908) and redescription of Lepadichthys frenatus Waite 1904 (Gobiesocidae: Diademichthyinae)
Article
  • Jan 2019
The clingfishes Lepadichthys frenatus Waite 1904 and Lepadichthys misakius (Tanaka 1908) are both redescribed as valid species, although the latter has previously been regarded as a junior synonym of the former. Lepadichthys frenatus and L. misakius are easily distinguished from their congeners by the following combination of characters: 14–19 dorsal-fin rays; 12–14 anal-fin rays; disc size moderate, its length 15.1–20.8 % of standard length (SL); and dorsal, anal and caudal fins connected by membranes. Lepadichthys misakius can be distinguished from L. frenatus by having 25–28 (modally 26 or 27) pectoral-fin rays [vs. 27–29 (29) in L. frenatus]; the upper end of the gill membrane level with the 5th to 8th (usually 6th) pectoral-fin ray base in lateral view [vs. 6th to 8th (7th)]; the lower 8th to 11th (9th) pectoral-fin ray base attached to the disc base by membrane [vs. 10th to 12th (11th)]; 7–11 (9) gill rakers on each arch [vs. 11–14 (12)]; anterior, posterior and least interorbital widths 9.0–11.8 (mean 10.7), 13.4–16.9 (15.3) and 3.2–8.8 (6.2) % SL, respectively [vs. 8.4–10.2 (9.4), 12.1–14.7 (13.5) and 2.8–5.9 (4.8) % SL, respectively]; and NC2 (nasal canal pore) usually located between the anterior and posterior margins of the posterior nostril (vs. usually located before the posterior nostril anterior margin). Morphological changes with growth of the two species are described in detail and a lectotype designated for L. frenatus.
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Checklist of Red Sea fishes with delineation of the Gulf of Suez, Gulf of Aqaba, endemism and Lessepsian migrants
Article
  • Nov 2018
The current checklist provides for each species of the Red Sea its records in the Gulf of Suez, Gulf of Aqaba, Red Sea main basin and its general distribution.This new checklist of Red Sea fishes enumerates 1207 species, representing 164 families. Of these, 797 species were recorded from the Gulf of Aqaba and 339 from the Gulf of Suez. The number of species from the Gulf of Suez is evidently lower than the actual number not including 27 Lessepsian (Red Sea) migrants to the Mediterranean that most likely occur in the Gulf. The current list includes 73 species that were newly described for science since the last checklist of 2010. The most specious Osteichthyes families are: Gobiidae (134 species), Labridae (66), Apogonidae (59), Serranidae (including Anthiadinae) (44), Blenniidae (42), Carangidae (38), Muraenidae (36), Pomacentridae (35), Syngnathidae (34), Scorpaenidae (24) and Lutjanidae (23). Among the families of Chondrichthyes, the most specious families are the Carcharhinidae (18 species) and Dasyatidae (11). The total number of endemic species in the Red Sea is 174 species, of these, 34 species are endemic to the Gulf of Aqaba and 8 to the Gulf of Suez.
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