Table Of ContentPROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
116(l):52-60. 2003.
Halidesmus socotraensis new species and Haliophis guttatus
(Forsskal), new records of congrogadine fishes from the Socotra
Archipelago (Perciformes: Pseudochromidae)
Anthony C. Gill and Uwe Zajonz
(ACG) Fish Research Group, Department of Zoology, The Natural History Museum,
London SW7 5BD, U.K., e-mail: [email protected];
(UZ) Ichthyology, Senckenberg Research Institute and Natural History Museum,
Senckenberganlage 25, D-60325 Frankfurt am Main, Germany,
e-mail: [email protected].
—
Abstract. Halidesmus socotraensis is described from six specimens, 39.6-
mm
69.5 SL, from the Socotra Archipelago, western Indian Ocean. It is dis-
tinguished from all other congrogadine species in having the following com-
bination of characters: three lateral lines (dorsal, middle and ventral com-
plexes); ventral lateral line branched on abdomen, with unpaired median section
on ventral midline; and upper lateral line without anterodorsal branch on to
nape. Haliophis guttatus is also newly recorded from the Socotra Archipelago
on the basis of 15 specimens.
The Socotra Archipelago in the western tions Development Programme (UNDP)
Indian Ocean at the entrance to the Gulf of Global Environment Facility funded project
Aden consists of four main islands (Fig. 1). "Conservation and Sustainable Use of Bio-
The largest and most easterly island, So- diversity of Socotra Archipelago," execut-
cotra, is located some 400 km south of the ed under the auspices of the Environmental
Arabian Peninsula and 200 km east of Cape Protection Council of Yemen. During two
Guardafui, the Horn of Africa. The other expeditions the second author collected
islands are Abd al-Kuri in the west and the fishes at inter- and subtidal stations
two smaller southern islands of Samha and throughout the archipelago. Among the col-
Darsah, also known as "The Brothers". lections were several specimens of the
The coastal fishes of the Socotra Archi- pseudochromid subfamily Congrogadinae,
pelago were poorly known until recently. a group of eel-like reef fishes commonly
Steindachner (1902, 1903) pubhshed the called eel blennies or snakelets. The sub-
first study of fishes from Socotra, based on family was revised by Winterbottom (1986)
material collected by the Austrian Expedi- who recognised 19 species in eight genera
tion to Socotra and South Arabia in 1898- (one with two subgenera); four additional
1899. He listed 57 species of marine and species (and one new subgenus) have been
&
brackish-water fishes. The first detailed described subsequently (Winterbottom
study of the fish fauna was a sight survey Randall 1994, Winterbottom 1996, Gill et
conducted by Kemp (1998), who provided al. 2000). Among the collections from the
an account of 215 fish species and a prelim- Socotra Archipelago, an additional new
inary zoogeographical analysis. species referable to the western-central In-
In 1999 and 2000 the Senckenberg Re- dian Ocean endemic genus Halidesmus
search Institute, Frankfurt, conducted the Giinther was discovered, and specimens of
Marine Habitat, Biodiversity and Fisheries the widely distributed western Indian Ocean
Surveys in the framework ofthe United Na- species Haliophis guttatus (Forsskal) were
—
VOLUME NUMBER
116, 1 53
]mn
Pakistan
^
Arabia
C^an
Yemen fnd^a
Socotra Archipelago
SomaNa
Indian Ocean
—
52°00' 52°30' 53°00' 53=30' 54=00' 54°30'.
I
12°50'--
12^40'--
12^30' --
12°20• Abd al-Kuri
^^^
^^^^^^ M3
Sia.
12°10'
Sta. 709
Darsah
12^00'
Fig. 1. Map of the Socotra Archipelago showing collection sites.
identified. The purposes of this paper are to G2). Vertebral counts are presented in the
describe the new species and to document form precaudal + caudal; the latter are de-
the new record of H. guttatus from the So- fined as vertebrae bearing a haemal spine,
cotra Archipelago. and include the terminal urostylar complex.
Terminology of ribs and intermuscular
Materials and Methods bones follows Gill (1998). Other methods
The nomenclature of cephalic laterosen- of counting and measuring follow Winter-
sory pores follows Gill et al. (2000). Al- bottom (1982). Institutional codes follow
phabetical codes for lateral lines in the new Leviton et al. (1985), except for NHCY-F,
Halidesmus generally follow Winterbottom which is for the fish section of the Natural
(1982), except that the structure of the ven- History Collection of Yemen. Osteological
tral lateral line necessitates the following details were determined from x-radio-
modifications (Fig. 2): the apparent homo- graphs. In the description of the new spe-
G
log of line in H. scapularis is represented cies, counts are given as values or value
in H. socotraensis (and H. polytretus) by a ranges for all type specimens, followed,
median anterior section (termed Gl) and a where variation was noted, by values for
bilaterally paired posterior section (termed the holotype in parentheses. Where counts
.
54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
B + C
Fig. 2. Diagram of Halidesmus socotraensis in lateral and ventral views showing lateral-line systems and
lettering scheme.
were recorded bilaterally, both counts are rotenone, U. Zajonz, F.N. Saeed, M. Apel
presented for the holotype, separated by a and E. Zandri, 4 April 2000 (station 709).
slash; the first count given is the left count. Paratypes.—Bymn 2002.1.19.3, 1, 63.3
Ratios for morphometric characters and fre- mm SL, ROM 72697, 1, 60.0 mm SL, SMF
mm
quency distributions for selected meristic 29293, 3, 39.6-69.5 SL, all collected
characters are given in Tables 1 and 2, re- with holotype.
—
spectively. Comparisons with other Hali- Diagnosis. The following combination
desmus species are based primarily on data of characters distinguishes H. socotraensis
provided by Winterbottom (1982, 1986) from all other congrogadines: three lateral
&
and Winterbottom Randall (1994). lines (dorsal, middle and ventral complex-
es); ventral lateral line branched on abdo-
Halidesmus socotraensis, new species
men, with unpaired median section on ven-
Socotran Snakelet
tral midline; and upper lateral line without
Figs. 2-4, Tables 1-3
anterodorsal branch on to nape.
—
Holotype.—SM¥ 29223, 64.3 mm SL, Description. Data for morphometric
Socotra Archipelago, Abd al-Kuri Island, characters appear in Table 1
SW coast, ca. 12°13'N 52°04'E, 6-15 m. Dorsal fin I + 58-61 (I + 61), all seg-
—
Table 1. Morphometric values forHalidesmus socotraensis expressed as percentages ofstandardlength (SL).
Paratypes
Holotype
mm SL 64.3 39.6 48.7 60.0 63.3 69.5
Soft dorsal-fin base 78.2 80.0 78.2 78.3 78.2 81.7
Anal-fin base 61.6 59.8 60.6 6L7 63.0 61.2
Snout to first dorsal 17.1 20.2 19.7 17.7 18.3 17.7
Snout to soft dorsal 19.8 22.5 22.2 19.2 19.6 20.1
Snout to anal origin 37.2 39.1 39.4 38.0 35.2 37.4
Head length 13.2 15.4 14.4 13.5 13.1 12.7
Depth at parietal 7.0 8.1 7.6 8.0 7.4 7.5
Depth at anal origin 7.9 8.8 8.8 8.7 8.5 7.5
Eye diameter 2.8 4.0 3.3 3.0 3.0 2.9
Snout length 2.8 3.3 2.9 2.8 3.0 3.2
Interorbital width 1.2 1.3 1.4 1.3 1.3 1.3
Upper jaw length 4.5 4.8 4.7 4.5 4.4 4.6
Lower jaw length 6.2 7.3 6.8 6.3 6.3 6.5
Pectoral length 5.6 5.1 4.9 4.7 5.1 9*
fin damaged.
VOLUME NUMBER
116, 1 55
PlOP
SOTP
AlOPx
LLP
NASP
DENP
POPP
ATP
Fig. 3. Halidesmus socotraensis, holotype, SMF 29223, 64.3 mm SL; detail of head showing laterosensory
pores, scalation and preserved pigmentation details. Abbreviations: ATOP, anterior interorbital pore; AN, anterior
nostril; ATP, anterior temporal pore; DENP, dentary pores; ITP, intertemporal pore; LLP, lateral-line scale pores;
NASP, nasal pores; PARP, parietal pores; PIOP, posterior interorbital pore; PN, posteriornostril; POPP. preopercle
pores; POTP, posterior otic pore; PTP, posttemporal pore; SCLP, supracleithral pore; SOBP, suborbital pores;
SOTP, supraotic pore. Scale bar = 2 mm.
mented rays branched; anal fin 45-48 (48), and, variably, to parhypural; epurals 1; su-
all rays branched; pectoral fins 9-10 (10/ praneurals 2.
10), upper 1-2 (2/1) and lower 0-2 (1/1) Cephalic sensory pore openings (Fig. 3;
rays unbranched, other rays branched; pel- all pores bilateral unless otherwise indicat-
vic fin absent, though small rod-like pelvis ed): nasal two, one pore just posterior to
present internally; caudal fin with five dor- upper lip, second pore just above posterior
sal and five ventral principal rays, and one nostril; anterior interorbital usually one,
dorsal and one ventral procurrent rays; small second pore present unilaterally in
branched caudal fin rays 5 + 4. Caudal fin one paratype, about two-thirds distance
fully connected by membrane to last ray of from typical anterior interorbital pore to
dorsal and anal fins. Vertebrae 17-18 + 46- posterior nasal pore; median (unpaired)
49 = 63-66 (17 + 49); ribs present on pre- posterior interorbital one, positioned above
caudal vertebrae 3 through 4-5 (3 through or slightly behind vertical through posterior
5); epineural bones present on precaudal edge of eye, additional posterior interorbital
vertebrae 1 through 3-5? (1 through 3?; pore present in one paratype, displaced lat-
bones difficult to distinguish on x-radio- erally in line with right side anterior inter-
graphs); hypural 5 absent; hypurals 3-4 orbital pore; supraotic one; posterior otic
fused to one another and to urostylar com- usually one, two pores present unilaterally
plex; hypurals 1-2 fused to one another in one paratype; suborbital usually eight.
56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Halidesmus socotraensis, holotype, SMF 29223, 64.3 mm SL, Abd al-Kuri Island, Socotra Archi-
pelago.
nine unilaterally in one paratype; preoper- 14 (11/14) pored scales, connecting poste-
cular seven; dentary usually four, three uni- rior terminus of line Gl with junction of
laterally in one paratype; intertemporal usu- lines F and H; line H (bilateral) with 125-
ally one, two unilaterally in two paratypes; 152 (152/147) pored scales, extending from
parietal two, canals from left and right side junction of lines F and G2 along lower part
of head not communicating in midline; an- of body to vertical through base of anal ray
terior temporal one; posttemporal usually 36-42 (41/42).
one, two unilaterally in one paratype; su- Olfactory capsule with two openings; an-
pracleithral one. terior opening a short tube, positioned about
Gill membranes fused to one another midway between posterior opening and edge
ventrally but free posteriorly from isthmus; of lip; posterior opening with slightly raised
branchiostegal rays 6; outer (anterior) gill rim, positioned near anterodorsal rim of or-
rakers on first arch 2 + 6-7 (examined in bit (Fig. 3). Cheek, upper edge of operculum
three largest paratypes only); pseudobranch (above horizontal through opercular spine)
filaments 6-7 (examined in three largest and body with small cycloid scales, those on
paratypes only). Lateral-line system com- upper part of operculum imbedded and in-
plex (Fig. 2), with pores on either dorsal or conspicuous; predorsal scales extending an-
ventral margins of lateral-line scales (or in teriorly to parietal commissure (Fig. 3).
the case of median lateral lines, on either Teeth in outer row ofpremaxilla and den-
left or right margins of lateral-lines scales): tary conical, recurved anteriorly, decreasing
line A absent; line B + C (bilateral) with in size and becoming laterally compressed
186-210 (210/206) pored scales, extending posteriorly, numbering 11-12 in the pre-
from shoulder to beneath vertical through maxilla and 13-15 in the dentary; single
base of segmented dorsal ray 49-54 (51/ row of small conical teeth present anteriorly
D
51); line (bilateral) with 177-205 (202/ on premaxilla and dentary immediately be-
197) pored scales, extending from just be- hind outer row teeth; vomer edentate; pal-
hind upper edge of pectoral-fin base along atine edentate.
midside to or almost to caudal-fin base, not Live coloration: Not recorded.
connecting anteriorly with line B + C; line Preserved coloration (Figs. 3, 4): Head
E (median) with 6-12 (9) pored scales, ex- and body dark brown, with indistinct to dis-
tending from just beneath branchiostegal tinct paler mottling, which may align to
membranes to vertical beneath pectoral-fin form diffuse bars or reticulations; lower
base; line F (bilateral) with 78-90 (90/89) part of head and abdomen pale brown; dark
pored scales, extending posterodorsally grey-brown stripe extending from midside
from junction of lines E and Gl almost to of lower lip through midside of upper lip to
pectoral fin, then along lower part of ab- mid-anterior edge of eye, then behind mid-
domen to junction with line G2; line Gl posterior edge of eye to upper edge of oper-
(median) with 59-63 (62) pored scales, ex- culum; dark grey-brown stripe edged ven-
tending from junction with lines E and F trally with creamy white stripe; large pale-
along ventral edge of abdomen to just in edged, dark grey-brown spot on shoulder;
front of anus; line G2 (bilateral) with 10- creamy white median stripe or series of
VOLUME NUMBER
116, 1 57
spots extending from tip of upper lip to or- bases only); and total vertebrae 63-66 (ver-
igin of dorsal fin; dorsal fin dark greyish sus 66-69).
brown with series of alternating small dark Halidesmus socotraensis differs from H.
grey-brown and pale brown spots on base polytretus in the following: lateral line A
of fin, spots encroaching slightly on to absent (versus present); pores in lateral line
D
body; anal and caudal fins dark greyish 177-205 (versus 204-216); cheek, nape
brown, sometimes with several pale brown and opercular scales present (versus ab-
or dark grey-brown spots; pectoral fin pale sent); preopercular pores 7 (versus 9); sub-
brown, with small dark grey-brown spot on orbital pores 8-9, usually 8 (versus 11-13);
middle of fin bas—e. posterior interorbital pores usually 1, rarely
Comparisons. The presence of three 2 (versus 2); principal caudal fin rays 5 +
lateral lines on the body (dorsal, middle and 5 (versus 6 + 5); and gill rakers 2 + 6-7
ventral complexes) places the new species (versus 1 + 5).
in the genus Halidesmus. There are four Relationships among H. socotraensis, H.
other species currently recognised in the ge- scapularis and H. polytretus are ambiguous.
nus (Winterbottom, 1982, 1986; Winterbot- The presence of lateral-line A is unique
tom & Randall, 1994): H. scapularis Giin- among congrogadines to H. scapularis and
ther from South Africa; H. polytretus Win- H. polytretus and suggests a sister relation-
terbottom from Kenya; H. coccus Winter- ship between the two species. Conversely,
&
bottom Randall from southern Oman; the presence of a median section of the low-
and H. thomaseni (Nielsen) from Masirah er lateral line is unique to H. polytretus and
Island (southeastern Oman) to Karachi H. socotraensis, and suggests relationship
(Pakistan) and the Bay of Bengal. Charac- between those two species. These relation-
ters distinguishing the four species are sum- ships will be tested in a forthcoming study
marized in Table 3. of the phytogeny and historical biogeogra-
Halidesmus is divisible into two apparent phy of Halidesmus by the se—cond author.
monophyletic groups, which are diagnosed Habitat and distribution. All known
by characters that are uniquely derived specimens were collected at a single station
SW
among pseudochromids. The first, which in a small enclosed bay on the coast of
consists of H. coccus and H. thomaseni, is Abd al-Kuri (Fig. 1, Sta. 709). The site is
diagnosed by a single synapomorphy characterised by highly diverse assemblag-
(fleshy crest on the snout and interorbital es of large massive and encrusting corals,
area). The other, which consists of H. so- and an associ—ated diverse fish community.
cotraensis, H. scapularis and H. polytretus, Etymology. The specific epithet refers
is diagnosed by two synapomorphies (ven- to the known distribution of the species.
tral lateral line with complex branching on
abdomen; lateral-line pores opening to ei- Haliophis guttatus (Forsskal)
ther ventral or dorsal margins of scales). Barred Snakelet
Halidesmus socotraensis differs from H. Fig. 5, Table 2
scapularis in the following features: pelvic
fins absent (versus pelvic-fin rays usually Haliophis guttatus is known from
1,2-3, fins rarely absent); lateral line A ab- throughout the Red Sea to the northern Gulf
sent (versus present); lateral line D with of Aden, southern Oman, the east coast of
177-205 pores, not joining C (versus with Africa between Kenya and Mozambique,
157-179 pores, joining C); lateral lines E and the west coast of Madagascar (Winter-
G &
and median (versus bilateral); medial pa- bottom 1984, Winterbottom Randall
rietal pores bilaterally paired (versus single 1994). Fifteen specimens were collected by
median pore); caudal, anal and dorsal fins the second author and associates in the So-
fully confluent (versus united near fin-ray cotra Archipelago. The morphology of
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
58
Fig. 5. Haliophis guttatus, SMF 29198, 59.5 mm SL, Abd al-Kuri Island, Socotra Archipelago.
these specimens agrees well with published reported by Winterbottom are small (often
descriptions of the species (e.g., Winterbot- five or fewer specimens).
tom, 1984, 1986; Randall, 1995). Winter- Material—SM¥ 29224, 1: 64.8 mm SL,
bottom (1984) noted that the species exhib- Socotra Archipelago, Abd al-Kuri Island,
SW
its geographic variation in several meristic coast, ca. 12°13'N 52°04'E, 6-15 m,
and morphometric characters, which he in- rotenone, U. Zajonz, F.N. Saeed, M. Apel
terpreted as a step cline, with the step oc- and E. Zandri, 4 April 2000 (station 709);
curring between about 27°N and 20°N in SMF 29198, 7: 38.0-79.5 mm SL, NHCY-
the northern Red Sea. Frequency distribu- F uncataloged, 7: 43.2-66.0 SL, Socotra
tions of counts relevant to this phenomenon Archipelago, Abd al-Kuri Island, E coast,
are provided for the Socotran specimens in Ras Anjara bay, ca. 12°10'N 52°22'E, 3-9
Table 2. Unfortunately, the specimens are m, rotenone, U. Zajonz and M. Apel, 8
somewhat distorted and difficult to measure April 1999 (station 173).
accurately; therefore, we do not provide rel-
evant morphometric details. Generally, me- Discussion
ristic values agree well with data presented
by Winterbottom for specimens from the The fish surveys of 1999 and 2000 com-
southern Arabian area (central and southern prised 18 large sampling stations scattered
Red Sea and Gulf of Aden), except for all around the Socotra Archipelago. Sam-
counts of precaudal vertebrae, which more ples of congrogadine species were obtained
closely approached values for specimens only at Abd al-Kuri, whereas pseudochrom-
from Mozambique and Madagascar. How- ids of the subfamilies Pseudochrominae (six
ever, sample sizes for many of the localities species of Pseudochromis Riippell) and
—
Table 2. Frequency distributions for selected meristic characters of Halidesmus socotraensis and Haliophis
guttatus from the Socotra Archipelago. Bilateral counts of pectoral rays are included.
Segmented dorsal rays Anal rays
43 44 // 581159 360 61 34 35 36 // 45 46 47 48
—
socotraensis 1
guttatus 2 9 4
Pectoral rays Total caudal rays Precaudal vertebrae
10 I 1 12 13 14 15 13 14 18
— — — —
12
socotraensis 4 8 —6 —
guttatus 27 4 11 13 2
Caudal vertebrae Total vertebrae
36 37 II 46 47 48 49 49 50 51 II 63 64 65 66
socotraensis 1 —3 2 1 2 3
guttatus 4 11 3 11 1
1 < ' ' > t
VOLUME NUMBER
116, 1 59
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———
60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pseudoplesiopinae (two species of Chlid- congrogadine fishes (Perciformes: Pseudochro-
ichthys Smith) were collected from other is- midae).—Copeia 1998:1041-1045.
lands as well. It seems possible that Abd al- -tioRn. oDf.aMonoeiw, s&ubJ.geBn.usHuatcnhdinssp.ec2i0e0s0.ofDtehsecrfiips-h
Kuri receives a higher influx of pelagic genus Congrogadus Giinther from Western
eggs and larvae from East Africa through Australia (Perciformes: Pseudochromidae).
the Somali current compared to the other Records of the Western Australian Museum 20:
69-79.
islands, enriching the fish communities. In
particular along the south and east coasts Kemp, J. M. 1998. Zoogeography o—f coral reef fishes
of the Socotra Archipelago. Journal of Bio-
there are isolated biodiversity "hotspots," geography 25:919-933.
which may act like traps for recruits by pro- Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. Daw-
viding shelter and suitable habitats (U. Za- son. 1985. Standards in herpetology and ichthy-
jonz, unpubl. data). Both sites where con- ology: Part 1. Standard symbolic codes for in-
stitutional resource collections in herpetology
grogadines were collected host particularly
and ichthyology.—Copeia 1985:802-832.
rich fish and coral assemblages as com- Randall, J. E. 1995. Coastal Fishes ofOman. Crawford
pared to other areas visited at Socotra. The House Publishing, Bathurst, 439 pp.
fish species compositions were exception- Steindachner, F. 1902. Wissenschaftliche Ergebnisse
ally rich, suggesting that there is limited der siidarabischen Expedition in den Jahren
1898 bis 1899. Fische von Siidarabien und So-
faunal exchange between these patches and —
cotra. Anzeiger der Akademie der Wissen-
the wider archipelago, at least of closely schaften Wien 39(24):316-318.
reef-associated species. 1903. Fische aus Siidarabien und Sokotra.
.
Denkschriften der Kaiserlichen Akademie der
Acknowledgments Wissenschaften, Wien 71:123-168. [dated 1902,
but actually published in 1903]
We would like to thank the staff of the Winterbottom, R. 1982. A revision ofthe congrogadid
Project Implementation Unit (PIU), espe- fish genus Halidesmus (Pisces: Perciformes),
cially C. Cheung and E. Zandri, the repre- with the description ofa new species from Ken-
sentatives of the Environmental Protection ya a—nd a list of the species included in the fam-
ily. Canadian Journal ofZoology 60:754-763.
Council (EPC) of Yemen, and the United
1984. Revision of the congrogadid Haliophis
.
Nations Office for Project Services (UN- (Pisces: Perciformes), with the description of a
OPS YEM/96/G32) for their assistance and new species from Indonesia, and comments on
support. All team members that helped in the —endemic fish fauna of the northern Red
Sea. Canadian Journal of Zoology 63:209-
the field are cordially thanked. J. Seigel
217.
checked details on the lateral-line system of
1986. Revision and vicariance biogeography
.
the holotype of Halidesmus polytretus, for of the subfamily Congrogadinae (Pisces: Perci-
—
which we are grateful. P. Hurst photo- formes: Pseudochromidae). Indo-Pacific Fish-
graphed the specimens in Figs. 4 and 5. R. es 9:1-34, pi. I. [dated 1985, but actually pub-
Winterbottom kindly discussed details of lished 1986]
1996. A new species of the congrogadin ge-
congrogadine systematics, reviewed a draft n.us Rusichthys from southern Oman (Percifor-
of the manuscript and provided useful sug- mes; Ps—eudochromidae), with notes on its os-
gestions. teology. Canadian Journal of Zoology 74:
581-584.
Literature Cited , & J. E. Randall. 1994. Two new species of
congrogadins (Teleostei; Pseudochromidae),
Gill, A. C. 1998. Homology of the anterior vertebrae, with range extensions for four other species.
ribs, and dorsal fin pterygiophores and rays in Canadian Journal of Zoology 72:750-756.
