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BULLE TINO fie BRITS VEOSEUM (NATURAL HISTORY)
ZOOLOGY VOR, 14
1966
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1968
DATES OF PUBLICATION OF THE PARTS
No. I 1 February 1966 No. 2 1 February 1966 No. 3 t1 March 1966 No. 4 23 March 1966 No. 5 3 September 1966 No. 6 2 September 1966 No. 7 9 September 1966 No. 8
9 September 1966
PRINTED IN GREAT BRITAIN BY ADLARD & SON _ LIMITED BARTHOLOMEW PRESS, DORKING
CONTENTS ZOOLOGY VOLUME I4
The Oedura tryont complex: east Australian rock-dwelling geckos. (Reptilia: Gekkonidae). By H. R. Bustarp
The elopoid and clupeoid fishes in Richardson’s “ Ichthyology of the seas of China and Japan’ 1846. By P. J. P. WHITEHEAD
Observations on the type-material of some genera and species of Polyzoa. By A. B. HAstTinGs
Marine nematodes from Durban, South Africa. By W. G. INGLIs
Studies on the British Dermanyssidae (Acari: Mesostigmata) Part II. Classification. By G. O. Evans-& W. M. TILL
A review of the genus Piuletor (Chiroptera: Vespertilionidae). By |ddehoas
Pontobdellinae (Piscicolidae: Hirudinea) in the British Museum (Natural History) with a review of the subfamily. By L. C.
LLEWELLYN
On the first Halosaur leptocephalus: from Madeira. By C. M. H. HARRISSON
Index to Volume 14
PAGE
441
487
: THE OEDURA TRYONI COMPLEX: EAST AUSTRALIAN ROCK-DWELLING | GECKOS. (REPTILIA: GEKKONIDAE)
|
H. ROBERT BUSTARD
BULLETIN OF "THE BRITISH MUSEUM (NATURAL HISTORY) | ZOOLOGY Volo ig Noo: LONDON: 1966
THE OEDURA TRYONI COMPLEX: EAST AUSTRALIAN ROCK-DWELLING GECKOS. (REPTILIA : GEKKONIDAE)
BY H. ROBERT BUSTARD
Department of Zoology, The Australian National University, Canberra, A.C.T.
Pp. 1-14; 3 Plates; 2 Text-figures
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. ra No, a LONDON: 1966
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 14, No. i of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.
© Trustees of the British Museum (Natural History) 1966
ERUS DEES Ou THE BRITISH MUSEUM (NATURAL HISTORY)
Issued i February, 1966 Price Ten Shillings
THE OEDURA TRYONI COMPLEX: EAST AUSTRALIAN ROCK-DWELLING GECKOS. (REPTILIA : GEKKONIDAE)
By H. ROBERT BUSTARD
SYNORSTS
Following an historical review of the genus Oeduva Gray the specific distinctness of O. tryoni De Vis from O. maymorata Gray is confirmed. O. tvyoni as previously recognized is shown to be a complex comprising two distinct species one of which is described as new. The two species are geographically isolated, the nearest populations being five hundred miles apart. Since the original type material of O. tvyoni has been lost a neotype is proposed for O. tvyoni in the interests of stability. Aspects of the ecology are discussed.
INTRODUCTION
THE genus Oedura is morphologically divided into two groups. One group comprises Oedura robusta Boulenger and the two subspecies of O. lesweurit (Dum. & Bibr.) and is characterized by minute granular dorsal scales. The other possesses moderately large, flat, juxtaposed dorsal scales arranged in regular rows; it includes O. mar- morata Gray which, as currently defined, contains two distinct species, and O. monilis De Vis and O. tryont De Vis. The geckos previously referred to as O. tryont form a discrete group of predominantly rock-dwelling Oeduva which can be divided into two forms on the basis of morphology, size, coloration and geographic distribution. O. tryont as recognized by Cogger (1957) is herein split into two species and the variation within them is discussed.
Collections of Oeduva are expanding rapidly, as is the increase of ecological know- ledge of the genus, and it is now possible to examine the status of the forms grouped together as Oeduratryonit. Before describing the Oedura tryont complex it is necessary to review the status of certain other members of the genus, since absence of clear-cut morphological differences has caused considerable confusion.
Since 1944 when Loveridge placed the African species in a separate genus (A/fro- edura), a decision which has recently been confirmed by osteological studies by Cogger (1964), the genus Oedura has been restricted to Australia and consists of O. marmorata Gray, 1842; O. tryoni De Vis, 1884; O. monilis De Vis, 1887; O. robusta Boulenger, 1885 ; O. lesueurii lesueurii (Dum. & Bibr., 1836) and O. lesweurii rhombifer Gray, 1845.
MATERIALS AND METHODS
At sexual maturity in Oeduva preanal pores develop in male specimens and are accompanied by a conspicious bulge at the base of the tail. Since both of these secondary sexual characters are always lacking in females the sexes can be readily separated. It is not possible to sex juveniles,
ZOOL, 14, I I
4 He ROBE Re Bilis AD
Specimens possessing male characters are considered mature although growth continues for at least one year after attainment of sexual maturity. Adult size for a species is based on the range of size of specimens definitely identifiable as males. Specimens of about mean male size or above not showing male characters are con- sidered as mature females. The size at sexual maturity in females is, therefore, arbitrary except when a field study is in progress, then it is defined as size at first breeding.
Throughout the paper the following abbreviations are used :
BMNH. British Museum (Natural History)
AM. Australian Museum, Sydney OM. Queensland Museum NMV. National Museum of Victoria
CHARACTERS OF THE GENUS OEDURA GRAY (Modified after Boulenger, 1885)
Limbs pentadactyl ; digits free, all clawed, dilated at the base with raised distal joint bearing a discoid dilation which has two large plates inferiorly, separated by a long groove into which the claw is retractile. Proximal to the distal plates are two to six paired and one to six undivided plates. Upper surface covered with homo- genous, juxtaposed scales; abdominal scales juxtaposed or slightly imbricate. Pupil vertical. Body depressed. Femoral and preanal pores in the male.
HISTORICAL REVIEW OF OEDURA TRYONI
Gray (1842) proposed the genus Oedura and described Oedura marmorata. In 1884 De Vis described O. tryont.
Oedura ocellata Boulenger, 1885 was later (1887) synonymized with O. tryont by Boulenger when he received the 1884 description by De Vis based on specimens from Stanthorpe, south-east Queensland. However, examination of the six syntypes of ocellata in the British Museum collection has shown that five are conspecific with QO. monilis and only one with O. tryont. A lectotype will be selected from the five specimens conspecific with monilis and the full implications of this will be explained in a later paper.
Phyllodactylus ( = Oedura) castelnaui Thominot (1889) was synonymized with O. tryont by Zietz (1920). The author has examined the three syntypes of Oeduva castelnaut and has found Zietz’s action to be incorrect. Oedurva castelnaui belongs to the O. maymorata complex and is being reinstated as a full species (Bustard, in prepn.).
Loveridge (1934) synonymized O. tryoni with O. marmorata. Loveridge was influenced by a specimen of O. marmorata from Port Darwin mentioned by Longman (1915) which had the infralabials separated by an azygous scale behind the mental, a character which Boulenger believed to be distinctive of tryoni.
Cogger (1957) drew attention to a differentiated population of O. tryoni in North Queensland and showed that it differed in interorbital and mid-body scale row
ET ZAR DS OR CHE OLDURA ERYONI COMPLEX 5
counts from the southern population of the same species. He considered that they were possibly subspecifically distinct but took no action due to the lack of material at his disposal.
re S LAU StOR OLD URA ERY ONE DE VIS
The species which possess moderately large, flat, juxtaposed, dorsal scales arranged in regular rows are similar in external morphology and differ most obviously in size, pattern of markings and coloration. The confusion concerning the validity of O. tryont which led Loveridge (1934) to synonymize it with O. marmorata was caused in part by the inadequate material available for study. However, the main cause was the partly inaccurate description of Oedura marmorata given by Boulenger (1885) and based on thirteen specimens. He wrote, “‘ mental triangular, shorter than the adjacent labials, which are in contact behind it ’’. This feature is certainly not characteristic of marmorata. Fifty-three specimens definitely identifiable as marmorata were examined and in fifty of these the adjacent labials did not meet behind the mental. In the remaining three specimens (A 4891 and A 4893 (both from Port Essington, N.T.) and R 12366 (one of two specimens from Yirrkala, via Darwin, N.T.), Australian Museum collection, Sydney) the adjacent labials do meet as figured by Boulenger (1885, pl. IX, 2). Boulenger also wrote, “‘ Tail thick, short, much depressed, oval, the end tapering to a point ; its width is contained twice or twice and a half in its length, and equals at least the width of the body’. While this statement is applicable to regrown tails from certain localities, e.g. the Northern Territory, it is untrue for original tails. In this and other genera where the tail is used as a food store, its condition is entirely dependent on the nutritional state of the animal. For this reason gross tail morphology is an unreliable taxonomic character.
The result of Boulenger’s error has been to throw doubt on the specific status of tryom rather than to instigate a redescription of marmorata.
Cogger (1957) has shown that tvyoni is a valid species, on the basis of ecological information and meristic characters. A glance at living material is sufficient to separate marmorata and tryoni on the basis of colour and markings and this also applies to most preserved material. Full details of coloration and its geographic variation are given by Cogger (1957).
Oedura tryont is restricted to eastern Australia (parts of N.S.W. and Queensland) whereas O. marmorata occurs in all States on the mainland of Australia, except Victoria.
O. tryont and O. marmorata can usually be readily separated morphologically by the postanal tubercles. In the case of marmorata there are one to four large tubercles (mostly 2-4) whereas in tvyoni there is a single postanal tubercle. Fifty specimens definitely identifiable as marmorata were examined and in only one was there a single postanal tubercle. Twenty-six specimens possessed two, fifteen had three, and seven had four postanal tubercles. Fifty specimens of tryoni were examined and in every case there was a single postanal tubercle,
6 H. ROBERT BUSTARD
O. tryoni is therefore accepted as a distinct species. As has been indicated above there are no valid synonyms.
All the syntypes of O. tryoni held in the Queensland Museum have been lost. No cataloguing of material was carried out prior to 1911, and it has proved impossible to identify these specimens in spite of a thorough personal examination of all the gekkonid material held at the museum. Since O. tvyont has not found universal acceptance and furthermore since the ¢vyont complex is being divided into two species, it seems desirable that a neotype should be designated in the interests of stability.
The author has visited the type locality of O. tryoni (Stanthorpe, S.E. Queensland) and made a collection of this species on Mount Marlay, elevation 2,850 ft., a rocky hillside within the town. From this series of topotypic material a specimen consistent with De Vis’ original description of O. tryoni is selected as neotype.
Oedura tryoni De Vis (Plates I, 2)
NeotypeE. AM. R2r160r1, Mount Marlay, Stanthorpe, S.E. Queensland (the original type locality). Collected under rock (pl. 1) by the author and Dr. P. Maderson, September, 1964. Adult female.
DESCRIPTION OF NEOTYPE. Head large, greatly depressed, oval, eye large, snout moderately long, ear opening oblique. Body much depressed. Limbs moderate. Digits strongly dilated, less broad than the apical dilation which is moderate, rounded, as broad as long. Four pairs of broad infradigital plates under the third and fourth toes of hind feet with indication of division of the fifth plate on the third toe, followed by three to four undivided lamellae. Head covered with hexagonal or rounded, moderately sized, flattened scales, interorbitals number twenty-one, rostral four- sided, about twice as broad as high, with median cleft above ; nostril pierced between the rostral, first labial, and four nasals. The anterior nasal greatly elongated transversely, its depth contained at least twice in its breadth, in contact with its fellow on the opposite side ; eleven upper and nine lower labials, mental triangular. Virst infralabials deep, separated by a moderate octagonal scale below the mental. Back covered with juxtaposed flat round scales, considerably larger than those on the head and arranged in a regular transverse series, becoming markedly smaller on the flanks ; abdominal scales juxtaposed, flat, rounded, larger than the dorsal scales. Mid-body scale row count about one hundred and eight. Tail regenerated, moderate length, depressed, thick, the end tapering to a point ; the regenerated portion is covered with uniform square scales arranged like the bricks of a wall. Postanal tubercle single.
Dorsal coloration (in spirits) pale grey with interspersed scales of a donkey brown colour. Pale grey ocellate markings edged with donkey brown occur on the head, body and limbs. On the head they are conspicuous on the post-orbital to occipital region, On the body these form transverse bands of ocelli (pl. 1). | In life the darker colour is honey brown ; lower surface dirty white, each scale possesses up to a dozen or more minute donkey brown spots. Dimensions: Absolute dimensions in
|
LIZARDS OF THE OEDURA TRYONI COMPLEX 7 mm. are followed by percentage of snout-vent length in brackets; snout-vent 88 ; tail (regrown) 58 (66) ; head 17 (19) ; snout 9 (10) ; orbit 4-5 (5); eye to ear 8 (9) ; width of head 18 (20) ; axilla to groin 40 (45); left fore limb length 27 (31) ; left fourth finger 5-5 (6) ; left hind limb length 31-5 (36) ; left fourth toe 6 (7).
DIAGNOSIS OF O. TRYONI. A species of Oedura with flat, round, juxtaposed dorsal scales about as large as the ventrals, characterized by numerous irregularly placed spots smaller than the orbit on the dorsal surfaces. The dorsal area of the head possesses a pattern of pale markings on a darker ground colour.
Referred Material in Australian State Museums and the British Museum (Natural History)
AUSTRALIA (no precise locality): (BMNH. 54.11.1.1: syntype of O. ocellata). NEw SoutH WALES: Moonbi Range (AM. R 17676-17681), 13 miles N. of Bendemeer
_ (between Tamworth and Armidale) (BMNH. 1964.1503-1508 ; AM. R 21608-21612), | Inverell (AM. R1947 and R 20952; BMNH. 96.7.1.5), Port Macquarie (AM. _ R15092), 16 miles S. of Tenterfield on the road from Glen Innes (BMNH. 1964. 1502), Tenterfield (AM. R 13118), Bundarra (AM. R 15649), Moree (AM. R 1815).
QUEENSLAND: Dalby (AM. R 17772-17774), Chinchilla (AM. R 2801-2805 ; QM. J 12591-12598 and 12600), Brisbane (AM. R 18310), Eidsvold, Upper Burnett River (AM. R 5404-5408, R 5307-5309, R 5832, R 6210), Maryborough (AM. R 2oggr and R 20994), Cooroomon, near Rockhampton (BMNH. 1926.2.25.21). Emuford (AM. R 15641), Fletcher (NMV. D 131), 3 miles west of Amiens (NMV. D 9314-9319, 9321-9323, 9325-9347, AM. R 21605-21606), near Pozieres (AM. R 21607), Boonah (OM. J 6418), Bell, Darling Downs (QM. J 2294), Woodford, N.C. Line (QM. J 3870), Warwick (OM. J 1354), 5 miles N. of Warwick (BMNH. 1964.1509-1510, Gympie (QM. J 12390, J 12391-12395), Alum Rock Station, via Amiens (OM. J 11975; BMNH. 1964.1511-1512), near Tara (BMNH. 1964.1501 ; AM. R 21613), Mount Marlay, Stanthorpe, (AM. R 21602-21604; BMNH. 1964.1496—1500).
VARIATION. (1) Subdigital lamellae. On the fourth toe of the hind foot of tryont there are 4-6 divided subdigital lamellae (75°, possess 4), followed by 1-4
‘(usually 3) undivided ones. That is, omitting the distal plates, tvyoni possesses
6-8 enlarged subdigital lamellae, in over 70° of those examined 7 or 8.
(2) There is some variation in the number of labials ; upper 10-12, lower 9-12.
(3) The variation in interorbital scale and mid-body scale row counts is illustrated in text-fig. 1. A detailed examination of the geographical variation in interorbital and mid-body scale row counts was made to see if these variations were clinal. No correlation was found to exist between latitude and scale counts.
(4) The variation in the pattern of markings is illustrated in pl. 2. Almost all specimens are characterized by well defined ocelli on the upper surface. Exceptional specimens (for example BMNH. 1964.1501, pl. 2 collected at Tara, South Queens- land) have these replaced by numerous very small spots. There is little variation in coloration the ground colour being brown or grey-brown and the ocelli cream or creamy-yellow. The main variation occurs in the arrangement of the ocelli which
8 H. ROBERT BUSTARD
30 ° ° Oy GOR0 o Oo io ° ° ° ° oo og ° ° ° ° ° e000 fo} oo 95 8 fo} ° 8 ° oo 8 og S Chew) 8°88 8 ©9000 ° ° ° 00000000 9 900 ° o 000 O° 0 0°0 “ 000 ° ° 2 ° ° go o ©0 0° 9 ° ° a re 3 8 au —
80 90 100 110 120 130
e = Oe6cdura coggeri
Mid-body scale rows i y ° Oeduro tryoni
Fic. 1. Counts of mid-body and interorbital scales in O. ‘vyoni and O. coggeri.
are usually placed transversely. Complete fusion forming continuous transverse bands never takes place, the individual ocelli invariably remain distinct, see below. The colour pattern of the northern populations of tvyonz does not resemble coggert more closely than that of the southern populations. Although populations of ¢vyonz can be separated by slight differences of colour and pattern these do not form a north-south cline.
(5) Dimensions. The dimensions of 6 adult topotypic fryont (BMNH. 1964. 1496-1499 and AM. R 21602-21603) are given below. No sexual dimorphism was found in any of the measurements taken. Averages are bracketed : Snout-vent 71-84 (78) mm., tail (where original) 77 (one only) ; head 21-23 (22) ; snout 9-10 (9); orbit 5; eye to ear 8-9 (8:5); width of head 20-21 (21); axilla to groin 41-44 (43); left fore limb 26-31 (29) ; left fourth finger 6-7 (6); left first limb 36-40 (38) ; left fourth toe 7-8 (7). The snout-vent length of 60 adult tryonz is 52-98 (77) mm. The smallest and largest males have a snout-vent length of 52 and
eee = tne
EIZARD:S OMTHE OL DURA TRYONI COMPLEX 9
88 mm. respectively. There is no evidence to suggest a cline of decreasing size with decrease in latitude, the size range being similar in the north and south of the geographic range of the species.
The form previously considered as the northern population of O. tvyont was found to be markedly smaller and of a very variable colour pattern. There is considerable overlap in body colour pattern between the two species, but constant pattern differences occur on the head (see diagnosis). The northern population possess a lower average subdigital lamellae count, and much larger and thus fewer inter- orbital and body scales than the southern population of O. tryont. Text-fig. 1 shows that on the basis of scale number (i.e. number of interorbitals and number of scales in a mid-body scale row around the circumference of the body) the two populations can be completely separated. In general the two forms can be separated on either scalation character. However, there is a region, shown in text-fig. I, where there is overlap in either character used alone but if the two characters are used together a line of difference separating all individuals of the two species can be clearly defined.
The degree of morphological differentiation between tvyont and coggeri in a genus where morphological differences even between sympatric species are slight, the demonstrated fact that the differences in scale counts, pattern of markings and size are not clinal in nature and the marked geographical isolation of the two species (both have discrete geographic ranges, see text-fig. 2) has led the author unhesi- tatingly to describe the northern population as a new species.
Oedura coggeri sp. n. (Pls. I, 3)
HototypE. AM. R17791—Lappa Junction, North Queensland, Australia, collected under rock by Mr. W. Hosmer, 24th January, 1954. Adult female.
PaRATYPES. All localities mentioned are in North Queensland: Lappa Junction (AM. R 17783-17790, R 17793-17803), Petford (AM. R 17629-17631 and R 17771), Herberton (AM. R 16679), Emuford (AM. R 15641), Irvinebank (AM. R 15644), Hartley’s Creek, nr. Cairns (AM. R 17767-17770), Mt. Garnet (BMNH. 1964.1494 ; OM. J 9290), 8x miles N.W. of Cairns on road to Laura via Mt. Carbine (BMNH. 1964.1495), Stannary Hills (QM. J 1293-5), Ravenshoe (BMNH. 1923.10.13.1-2).
DESCRIPTION OF HoLotyPe. Head large, greatly depressed, oval, eye large, snout moderately long, ear opening oblique. Body much depressed. Limbs moderate. Digits strongly dilated, almost as broad as the apical expansion, which is moderate, rounded, as broad as long. Four pairs of broad infradigital plates under the third and fourth toes of hind feet, followed by two or three undivided ones. Head covered with uniform hexagonal or rounded, large flat scales, interorbitals number fifteen, rostral four-sided, at least twice as broad as high, with median cleft above. Nostril pierced between rostral, first labial, and four nasals. The two upper nasals large, the transverse length of the anterior nasal about equal to its depth, the supra- nasal in contact with its fellow on the opposite side. Ten upper and ten lower labials,
ZOOL. 14, I 2
10 H. ROBERT. BUSTARD
mental triangular. First infralabial deep, separated by a large octagonal scale below the mental. Back covered with juxtaposed flat round scales, of a similar size to those on the head, arranged in a regular transverse series, not becoming markedly smaller on the flanks. Abdominal scales juxtaposed, flat, rounded, of a similar size or slightly larger than the dorsal scales. Mid-body scale row count about eighty- eight. Tail regenerated, moderate length, depressed, not noticeably thick, the end tapering to a point; the regenerated portion is covered with equal square scales arranged like the bricks of a wall. Postanal tubercle single. Dorsal coloration fawn with dark brown reticulate markings continued on to the limbs and tail. A dark line along the upper border of the snout immediately above the supralabials, commencing at the rostral, passing through the eye and forming a “ V-shaped ”’ marking on the nape of the neck. | Postorbital to occipital region of head unmarked (pl. 1). Lower surfaces cream.
Dimensions. Absolute dimensions in mm. are followed by percentage of snout- vent length in brackets: snout-vent 65; tail (regrown) 34 (52); head 16-5 (25) ; snout 7 (II); orbit 4 (6); eye to ear 6-5 (10) ; width of head 13-5 (21); axilla to groin 30 (46) ; left fore limb length 20 (31) ; left fourth finger 3-5 (5) ; left hind limb length 26 (40) ; left fourth toe 5 (8). The smallest and largest males of coggeri have a snout-vent length of 44 and 68 mm. respectively.
VARIATION AMONG THE PARATYPES. The following characters are subject to variations :
(1) Subdigital lamellae. On the fourth toe of the hind foot of coggevz there are 3-4 divided subdigital lamellae (both occur in equal proportions), followed by 1-3 (usually 2) undivided ones. That is, omitting the distal plates, coggervi possesses 5-6 enlarged subdigital lamellae. The overlap in number with tryonz, however, prevents the lamellar arrangement being used as a diagnostic feature.
(2) Occasionally (in AM. R 17784-17785 and 17802) there is only one small nasal instead of two. The presence of two large nasals is constant.
(3) There is considerable variation in the labials ; upper 8-12, lower 8-11.
(4) The variation in interorbital scale and mid-body scale row counts is illustrated in text-fig. 1. As with O. tvyont a test was made for a north-south clinal variation in scale counts. No evidence for this was found.
(5) There is considerable variation in the pattern of markings. Juveniles are brown with circular or transverse cream markings. In adults the ground colour is cream to brown. Some specimens possess dark brown reticulate markings. In many specimens these enclose oblong or circular patches of cream colour. In others there are definite cream spots outlined in dark brown on the dorsal surface. The ventral surface is white. This pattern variation is not clinal, even within a single series—14 specimens from Lappa Junction—the range of colour pattern is very great, the three basic types outlined above being present (pl. 3).
(6) Dimensions. No sexual dimorphism was found in any of the measurements taken. Separate figures are given where differences occur between adults and juveniles. Figures other than snout-vent length are given as percentage of snout-
LIZARDS OF THE OEDURA TRYONI COMPLEX II
vent, averages are bracketed: snout-vent : adults 50-70 (61) mm., juveniles 33-43 (37) mm.; tail (where original) 53-78 (66); head: adults 24-28 (26), juveniles 28-30 (29); snout: adults ro-12 (11); juveniles 11-13; orbit: adults 6-7 (6), juveniles 7-8 (7) ; eye to ear 7-11 (10) ; width of head: adults 18—23 (21), juveniles 21-24 (22) ; axilla to groin 34-46 (41) ; left fore limb 24-34 (30) ; left fourth finger : adults 5-7 (6), juveniles 6-7 (7) ; left hind limb 34-44 (40) ; left fourth toe: 6-9 (7).
DIAGNOSIS OF O. COGGERI. A species of Oedura with flat, round, juxtaposed, dorsal scales about as large as the ventrals, characterized by numerous irregularly placed spots much smaller than the orbit. The spots frequently coalesce to form transverse cross bands (pl. 3).
The spotted specimens of coggevt can be separated from tryont by the possession of a pale unspotted area in the postorbital to occipital region of the head. This area is bounded by a line running along the side of the head, passing through the eye, and joining with its neighbour at the nape of the neck.
DEST RUBUETON AND HABITAT
The distribution of Oedura coggeri and O. tryoni is shown in text-fig. 2. Neither species penetrates far inland and both are probably restricted to Eastern Australia.
Queensland
New South Wales
® Oeduro coggeri
o Oedura tryoni
— 100 miles
Fic. 2, Distribution of O. tryoni and O. coggert.
12 H. ROBERT BUSTARD
The two species are separated by a gap of over five hundred miles in which to date no specimens of either species have been collected. As with O. marmorata suitable habitat occurs in this region and this has been searched by the author so far without SUCCESS.
The ecology of Oedura tryoni has been examined recently (Cogger, 1957) and Oedura coggert appears to occupy a similar niche. Both species seem generally to inhabit rock crevices but some individuals live on trees. Of the two specimens of coggeri collected in North Queensland by the author (p. 9) one was under the bark of a dead iron bark tree (Eucalyptus spp.) at a height of four feet from the ground and the other under a slab of a large granite boulder forming part of a low granite outcrop in Eucalyptus savannah-woodland. Prolonged searching failed to locate further specimens at either locality.
DISCUSSION
O. coggeri is by far the smallest species of the group possessing flat, round, juxta- posed dorsal scales, which also includes marmorata, monilis and tryont. The species marmorata, mons and tryont are most readily separated by coloration which Cogger (1957) has convincingly shown to be a reliable character. O. marmorata possesses about five or six light-coloured cross bands. In many geographical areas these bands break up in adults and the resultant light patches are irregular in shape and size. O. monilis, tryont and some specimens of coggeri are the only species of Oeduva charac- terized by spots. O. tvyont and some coggert possess numerous, irregularly placed, small, light-coloured spots. The distribution and size of these spots is diagnostic as pointed out by Cogger. In ¢tvyoni and coggeri the spots are numerous and much smaller than the orbit whereas in monzlis they are much fewer in number (typically only five or six pairs which may show various degrees of fusion), and are at least as large as the orbit. O. monilis possesses a conspicuous, pale-coloured ‘“‘ V-shaped ”’ marking or nuchal band on the nape of the neck, formed by the fusion of two ocelli. This characteristic band is usually absent in tvyoni and coggeri, and is to be distin- guished from the dark lines along the upper border of the snout which may join to form a “‘V’’ in some specimens of coggeri (e.g. the holotype, pl. 1). Another useful character, Cogger (pers. comm.), is the presence in monzlis of a light area on the side of the head with its upper edge extending from the angle of the mouth to the ear and thence to about the insertion of the fore limb. This is typically absent in fvyoni and coggert.
Morphological differences in relative scale size are summarized in text-fig. I.
The relationship of O. coggeri to other members of the genus can be considered only tentatively at present. The author considers that coggeri is probably the northern representative of tvyon?. O. fryont may at one time have had a continuous distribu- tion from New South Wales through South-East and East-Central Queensland to the Cape York Peninsula. Geographic isolation of the two populations at a later time led to the observed speciation, Although ¢vyoni has been considered strictly rock
Se
5 enon
LIZARDS OF THE OEDURA TRYONT COMPLEX 13
dwelling (Cogger, 1957), in several localities the author has found specimens on trees (Eucalyptus spp. and Acacia harpophyla) and it is possible that formerly this habitat was much more extensively used. Competition may have prevented the continued exploitation of this habitat and present day tryont and coggert populations mainly occur in rock crevices. At Mount Garnet the northern population of O. marmorata and O. lesueurit rhombifer were common below the bark of dead trees (Eucalyptus spp.) in the area where the specimen of O. coggeri was collected. The three forms are also known to be sympatric in a area between Atherton and Mt. Garnet.
The distribution of tvyoni and coggeri is closely paralleled by the distribution of Oedura marmorata in Eastern Australia where there is a geographical gap of some 500 miles between northern and southern populations. The northern population of O. marmorata has twice received specific recognition (as castelnaw and mayert Garman (1901)) and is to be separated from O. marmorata occurring elsewhere in Australia and restored to specific status following an Australia-wide study on the O. marmorata complex (Bustard, in prepn.).
Oedura coggeri is named after Mr. Harold G. Cogger, Curator. of Reptiles and Amphibians in the Australian Museum, Sydney, in recognition of his pioneer work in applying the results of ecological investigation to the systematics of this difficult genus.
The author’s collection of twenty-two specimens has been deposited in the British Museum (Natural History) and the Australian Museum, Sydney.
SUMMARY
A detailed examination of museum material together with ecological field collect- ing has confirmed the specific distinctness of Oedura tryoni De Vis. O. tryont as currently defined is shown to comprise two species. O. coggeri n. sp. is described from North Queensland. It is tentatively suggested that O. coggeri differentiated from O. tryoni following geographic isolation. The syntypes of O. tvyont have been lost and a neotype is proposed in the interests of stability.
ACKNOWLEDGEMENTS
I am grateful to Mr. Joe Bredl my companion on my trip to Cape York for his skill and energy in collecting material; Dr. R. D. Hughes, Mr. R. E. Barwick and Mr. H. G. Cogger for reading the MS. and for valuable comments and advice ; Miss A. G. C. Grandison for constructive criticism and for editing the MS.; Mr. I. Grant for preparing the plates and Mr. A. A. Argyle for ready assistance in lengthy preparation for field trips. The following people kindly lent me material under their care: Miss A. G. C. Grandison, British Museum (Natural History), Mr. H. G. Cogger, Australian Museum, Mr. J. T. Woods, Queensland Museum, Mr. J. McNally, National
_ Museum of Victoria, Mr. F. J. Mitchell, South Australian Museum, Dr. G. M. Storr,
Western Australian Museum, and Mr. K. Slater, Northern Territory collection.
14 H. ROBERT BUSTARD
REFERENCES
BouLENGER, G. A. 1885 & 1887. Catalogue of Lizards in the British Museum (Natural History). Vols. 1 and 3. 2nd ed. London.
Coccer, H. G. 1957. Investigations in the gekkonid genus Oeduva Gray. Proc. Linn. Soc.
N.S.W. 82 (2): 167-179. 2 plates, 6 text-figures.
1964. The comparative osteology and systematic status of the gekkonid genera A froedura
Loveridge and Oedura Gray. Proc. Linn. Soc. N.S.W. 89: 364-372.
De Vis, C. W. 1884. On new Australian lizards. Pyvoc. Roy. Soc. Qld. 1: 53-56.
1887. A contribution to the herpetology of Queensland. Pyvoc. Linn. Soc. N.S.W.
12 : 811-826.
GARMAN, S. 1901. Some reptiles and batrachians from Australasia. Bull. Mus. comp. Zool. Harv. 39: 1-14.
Gray, J. E. 1842. Description of some new species of reptiles chiefly from the British Museum
collection. Zool. Miscell. 57-59.
1845. Catalogue of Lizards in the British Museum. ist ed. London.
Loneman, H. A. 1915. Reptiles from Queensland and the Northern Territory. Mem. Qld. Mus. 3: 30-34. 2 plates.
LoveERIDGE, A. 1934. Australian reptiles in the Museum of Comparative Zoology, Cambridge,
Mass. Bull. Mus. comp. Zool. Harv. 77 : 243-383.
1944. New Geckos of the Genera Afroedura new genus and Pachydactylus from Angola.
Amer. Mus. Novit. No. 1254: I-4.
TuHomrinot, A. 1889. Observations sur quelques reptiles et batraciens de la collection du Muséum d’Histoire naturelle de Paris. Bull. Soc. Phil. Pavis. 8 (1) : 21-30.
Ziftz, F. R. 1920. Catalogue of Australian lizards. Rec. S. Aust. Mus. 1: 181-228.
PLATE 1 UPPER ROw (left to right) : neotype of Oedura tryoni De Vis, holotype of Oedura coggervi n. sp. LowER ROW : O. tvyoni habitat at Mr. Marlay, Stanthorpe. The neotype was collected from under the slab at bottom centre.
Bull. B.M. (N.H.) Zool. 14, 1 PAA ES
Pace?
Top row (left to right): O. tvyont juvenile from type locality (Stanthorpe), adult Acacia. dwelling variant from Tara, S. Queensland and juvenile from 13 m. north of Bendemecr. i MIDDLE AND BOTTOM ROWS: Variation among adult topotypic O. tryont.
C
PLATE
Bull. B.M. (N.H.) Zool. 14, 1
PLATE 3
Variation among O. coggeri. Top row: Petford. MIDDLE Row: Hartley’s Creek, near Cairns. BOTTOM ROW: Lappa Junction.
Bull. B.M. (N.H.) Zool. 14, 1 PLATE 3
PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING
THE ELOPOID AND CLUPEOID FISHES IN RICHARDSON’S “ICHTHYOLOGY OF THE SEAS OF CHINA AND JAPAN” 1846
Pee) WV LE BRE AD
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Wiel, tut ING, 2 LONDON: 1966
fae ELOPOID, AND CLUPEOID FISHES IN RICHARDSON’S “ICHTHYOLOGY OF THE SEAS OF CHINA AND JAPAN” 1846
BY P. J--P.. WHITEREAD
British Museum (Natural History)
Pp. 15-54; 7 Plates
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 14 No. 2 LONDON: 1966
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL “HISTORY),. wnstituted in: 1940, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a@ separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 14, No. 2 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.
© Trustees of the British Museum (Natural History) 1966
TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 1 February, 1966 Price £1 6s.
ak ELOPOID AND CLUPEOW FISHES: IN RICHARDSON’S “ICHTHOLOGY OF THE SEAS OF CHINA AND JAPAN” 1846
ByuP ls Po Wir EEE AD
CONTENDS
Page ABSTRACT . : 1 é : ‘ ‘ ‘ . . : 3 17 INTRODUCTION . : ; , : : ; g : : : 7, RICHARDSON’S REPORT ; : : : : : : : . 18 THE REEVES ILLUSTRATIONS : ; : : : : : : 19 RICHARDSON’S SPECIES : : : ; : : : : nn 24. List oF HonG KoNnG SPECIES : : ; : : : : + 46 REFERENCES : : : z : : : 2 : : é 50 INDEX : : : : : : E : : : : : 52
A Bis TRAC Th
THE twenty-one elopoid and clupeoid fishes included by Richardson in his Report on the Ichthyology of the Seas of China and Japan are critically examined in the light of the specimens and the hitherto unpublished Reeves illustrations in the British Museum (Natural History). Sixteen of these species are considered valid, and the type status of certain of the specimens is established. A list is given of thirty-four elopoid and clupeoid species recorded from the Hong Kong area, and ten further species likely to occur there.
ENTRODUCTION
While studying certain herring and anchovy species, it became necessary to examine those species included by Richardson in his Report on the Ichthyology of the Seas of China and Japan, published in 1846. Since Richardson’s “ Report’ was based partly on specimens in the British Museum (Natural History) and partly on a collection of coloured illustrations compiled by John Reeves, also now in the British Museum (Natural History), the opportunity was taken to make a critical assessment of all twenty-one of the elopoid and clupeoid species reported by Richardson. The results have shown that a similar study of other groups would clear up many of the doubts and errors which have surrounded some of Richardson’s species.
Over three hundred of the species listed by Richardson in the “ Report ’’ were represented by a Reeves illustration based on specimens from the markets at Macao and Canton. The Reeves illustrations are thus an important, sometimes decisive, factor in the identification of certain of Richardson’s species. Unfortunately, the illustrations have never been published and ichthyologists have not always had the opportunity for consulting them. The twenty drawings covered by the present work are therefore reproduced here.
A full list of the herring-like fishes in the Hong Kong area has not yet been pub- lished. Since identification of the Richardson fishes has entailed an assessment of all Hong Kong—Canton records, I have compiled a tentative list of forty-four species for this area (Table 2). Richardson mentioned twenty-one species but four of these are here considered synonyms, and one cannot be identified (Table 1).
ZOOL. 14, 2 3
18 Pe JR Wer LE eR ALD RICHARDSON’S “REPORT”
For various historical reasons, knowledge of the fishes of the seas of China and Japan lagged far behind that of European waters by the beginning of the nineteenth century. In the Chinensia Lagerstrémiana, Linnaeus (1754) listed only 12 species of fish from China. In 1750, Pieter Osbeck, chaplain to a Swedish East Indiaman and a pupil of Linnaeus, examined fishes in the vicinity of Canton and he mentioned g species (Osbeck, 1757). Forster (1771), in an English translation of Osbeck’s “ Dagbok”’, added a further 9 species given by Linnaeus in the Systema Naturae (or in the Amoenitates Academicae), to make a total of only 18 Chinese species described in the Linnaean method at that time. Boeseman (1947) has pointed to the equally sparse knowledge of Japanese fishes during the eighteenth century and in fact up until the publication of the ichthyological volumes of the Fauna Japonica in 1842-50.
Richardson’s “‘ Report ’’ was one of a number of important ichthyological works which appeared at the middle of the nineteenth century. The works of Bleeker, Temminck & Schlegel, Giinther, and Cuvier & Valenciennes, all dealing (in part at least) with fishes from the western Pacific, appeared at this time. The “ Report ”’ was published in the same year as that part of the Fauna Japonica dealing with clupeoid fishes, and it pre-dates the clupeoid volumes of Cuvier & Valenciennes (vols. 20 and 21) and also Giinther (vol. 7), as well as most of Bleeker’s papers. Unfortunately, due to curatorial errors and the fact that the Reeves illustrations were never published, some of Richardson’s species have been ignored for so long that they are now nomuina oblita.
Richardson based the “ Report ’”’ chiefly on the coloured illustrations of John Reeves (1774-1856), supplemented by specimens from various collections. Of the latter, the fishes sent by John Reeves himself from Macao and Canton must be considered the most important since some of these were the actual models from which the drawings were made. Reeves’ son, John Russell Reeves (1804-77), also resident in Macao, sent further specimens to the British Museum, although certain of the latter were “not figured in his father’s drawings’ according to Richardson (““ Report ’’, p. 189). The British Museum specimens are mostly labelled J. R. Reeves, even where the “ Report’ implies that they were collected by the father not the son. The remainder are merely labelled “ Reeves’’. Richardson based 26 of his new species on specimens sent by John Reeves (or his son).
Unfortunately, the elopoid and clupeoid specimens presented by Reeves (and no doubt this is true of other groups also) were not registered, although most were listed in Gunther’s catalogue. The consistent omission of registration numbers on the Reeves clupeoid specimens suggests that they were presented prior to the adoption of the present registration system (in 1837). The three specimens listed here which were registered were all from the Haslar Hospital Museum. Richardson was Medical Inspector of Naval Hospitals at about this time.
A second collection of fishes studied by Richardson was that made by the Rev. George Vachell, who was Chaplain to the India Company in Macao in about 1830. The Vachell collection, of about a hundred fishes, was deposited with the Cambridge Philosophical Institution. According to Shipley (1913) the Philosophical Society Collection was transferred in 1865 to the Museum of Comparative Anatomy and
”
REC AR ID SONS ELOPOLD AND CLUPEOID FISHES 19
Zoology, but for several years was maintained as a separate collection. In 1866, time was spent in “ overhauling these collections!, eliminating useless or decayed specimens ’’. In 1866-67 Giinther examined the Vachell, Darwin and Lowe collec- tions of fishes at Cambridge, and he included a note on at least some of the species represented in the subsequent volumes of the Catalogue of Fishes in the British Museum, e.g. Clupea fuegensis and C. arcuata, but not the Darwin type of C. sagax (although it was evidently there since it was later (1917) transferred to the British Museum).
Richardson based 22 of his new species solely on Vachell specimens. Only one such type is involved in the present study, Chatoessus maculatus. However, this specimen appears to have been lost (see p. 37).
In addition to the Reeves and Vachell specimens, Richardson based a further 16 new species on specimens already in the British Museum (mainly presented by the Haslar Hospital, but a few from other sources).
Richardson’s descriptions are often brief and one could wish for additional notes, especially when type designations are required. There is an interleaved copy of the ‘“ Report ”’ in this museum, but it contains only a few short alterations or additions by Gray, who was then Keeper of Zoology. At about this time Gray had compiled a manuscript catalogue of the British Museum fish specimens. Unfortunately, only the “‘ Chondropterygii’’ section was published (Gray, 1851), and the remainder appears to be lost. The Reeves specimens would have been listed here and this might well have resolved such puzzles as the disappearance of the type of Clupea nymvphaea (see below, p. 24).
On Richardson’s death, a bundle of his notes, drawings and some published figures were offered to the British Museum by his son. The notes were rejected, but the figures kept. However, the latter have little relevance to the present study, except perhaps for some tracings of certain of Forster’s drawings of Australian fishes (from the second of Cook’s voyages) (see below, Megalops cyprinoides, p. 44).
Compared with other ichthyological works of the time, Richardson’s “ Report ”’ is a slender volume of a little over a hundred pages. However, some 665 species are listed from Chinese (or Japanese) waters, an enormous increase on any previous list. (Lacepede and Schneider knew about fifty Chinese species at the beginning of the nineteenth century.) Of these, 142 were described as new species or varieties. The “ Report’? would undoubtedly have been enriched by reproduction of the Reeves illustrations since 83 (or over half) of Richardson’s new species were based solely on a Reeves drawing. But, although the “ Report ’’ was in many ways over- shadowed by the Fauna Japonica, it dealt with certain tropical species not encoun- tered in Japanese waters, and many of Richardson’s species are still accepted today.
THE REEVES TLLUSTRATIONS
In the Zoological Library of the British Museum (Natural History) are certain original sets of drawings, some of which were published (e.g. the Hardwicke illus- trations of Indian fishes by Gray, 1830-35). Others, such as the drawings by Forster, Parkinson and Ellis, made during Cook’s voyages, have never been published. The
1 This statement from Shipley (1913) does not refer specifically to the fish collections, although it is likely that they too were overhauled at this time.
20 P. J. P. WHITEHEAD
Reeves illustrations, of which the Zoological Library has three sets, have also remained unpublished.
Richardson (‘‘ Report ’’, p. 188) states that ““ John Reeves, Esq., who was long resident at Macao, filling an important office in the employ of the India Company, with an enlightened munificence, caused beautiful coloured drawings to be made of no fewer than 340 species of fish which are brought to the markets at Canton ”’ Richardson then praises the drawings ‘‘ which are not surpassed in the plates of any large European work of the present day’”’. The praise is justified: some of the illustrations would not be out of place in a modern ichthyological paper. The paintings of clupeoid fishes, often finely dusted with silver and gold, are both aestheti- cally satisfying and at the same time are strongly indicative of painting from “ life ”’ rather than compilation from colour notes.
According to Richardson, Reeves had four sets of these illustrations made. Three of these sets are now in the Zoological Library of this museum. Comparing the three sets, it is not possible to judge whether one particular set contains the original draw- ings from which the other two were copied. The standard of the individual paintings varies somewhat within each set, and good and bad figures occur in each. Neither is there any indication that the same artist was responsible for all or the majority of figures in any one set. It seems that a number of Chinese artists were employed by Reeves over a period of several years.
Also in the Zoological Library is a bound volume containing a number of lists in Reeves’ hand giving dates of completion of many of the drawings, and in some cases a few brief notes and occasionally the name of the artist (Akut, Akew and Asung are mentioned). The lists are written on East India Company notepaper, watermarked 1827 and 1828, and the dates given cover the period 1828-30. Also included are some receipts listing a number of fishes by their Chinese vernacular names (in Chinese script) and a pencilled note of the artist ’s name and the amount paid (e.g. 2-5 dollars for a dozen or more paintings).
The four sets of Reeves illustrations can be commented on briefly.
A. The Reeves Set
A bound volume of paintings with four or more fishes on each page. The pages are numbered 1-124 and each fish bears a small number, usually in red ink, sometimes in pencil. On each page these Reeves numbers run consecutively, but the order of the pages has evidently been altered, perhaps in an initial attempt to place them in systematic order. The Reeves numbers correspond with those in the lists mentioned above. The figures also bear the Chinese vernacular (presumably Cantonese) names in Chinese script, but there are no latin names (although such are used in the lists). Several illustrations are repeated (e.g. see Table 1), and in one case at least (pp. 3 and 6) almost the entire page is duplicated. At least one figure (94l) is omitted in the Richardson set.
According to one source (Anov., 1904), Miss Reeves, sister of John Russell Reeves, presented her father’s collection of Chinese drawings to this museum on her brother’s death in 1877. In the “ Report ’’, Richardson states that ‘“‘ Another copy, left by Mr. Reeves at Macao with Mr. Beale, formed the groundwork of the enumeration of
RICHARDSON’S ELOPOID AND CLUPEOID FISHES 21
Chinese fish in Bridgman’s ‘ Chrestomathy’”’. This might be the copy which Miss Reeves presented, since Richardson only listed two copies in the British Museum in 1846 (but see also below).
B. The Richardson Set
A bound volume of paintings inscribed on the fly-sheet “ Sir John Richardson’s set of drawings of Chinese fish by native artists (one of four) prepared under the supervision of J. Reeves (used by Richardson for his ‘ Report ’) ’’.
The drawings have been cut out from their original pages and have been remounted in the systematic order adopted by Richardson in the “‘ Report’’. The pages are numbered 1-146 and several fishes are mounted on each page. Each drawing bears a small number in red ink, corresponding with the Reeves numbers in the preceding set, and the Chinese vernacular name (in Chinese script). Each drawing is also given its latin name in accordance with Richardson’s text.
The figures reproduced here are taken from the Richardson set. All are reduced to the same size.
C. The Hardwicke Set
Four bound folio volumes amongst the Hardwicke collection of drawings contain, in addition to illustrations of Indian fishes, a set of Reeves drawings of Chinese fishes. Richardson states that these drawings were presented to General Hardwicke by Reeves and had been examined “by many English and foreign ichthyologists ’’, including Miiller and Henle. The drawings have been cut out and remounted, and they are individually numbered in pencil (1-165, 166-313, in vol. 20a and b; 1-174, 175-317 in vol. 21a and b). In some cases the Reeves number (in red ink) and the Chinese vernacular ideogram are present. All the Reeves drawings are named and have a page reference to Richardson’s “‘ Report’’. Of the elopoid and clupeoid species, three found in the previous two sets are here missing (see Table 1). In the inter-leaved copy of the “ Report” in the Zoological Library, Gray has listed about 45 Reeves drawings not represented in the Hardwicke set.
The Hardwicke drawings were presented to this museum in 1835.
De the Fourth” Set
I have been unable to trace the fourth set mentioned by Richardson nor any other reference to such a set other than those based on Richardson’s statement. It is possible that the fourth set is indeed the Beale set, i.e. the Reeves set given to Beale and lent to Samuel Wells Williams, who was responsible for the Natural History section of the Chinese Chrestomathy (Bridgman, 1841). In this work, 245 fishes are listed (15 clupeoids), but their serial numbers do not tally with the numbers on the Reeves drawings. Cantonese vernacular names are given (English and Chinese script) and also the generic names; the latter may have been derived from the drawings, but if so, then the drawings will indeed constitute the fourth set, since the Reeves set in this museum does not bear generic names.
The fourth set is not amongst the several series of paintings of Chinese fishes in the Department of Prints and Drawings nor in the Department of Oriental Printed books and Manuscripts of the British Museum ; nor is it in the Library of the India Office in London.
22 B. J. P; WHITERE AD.
Tz
A list of the elopoid and clupeoid species mentioned by Richardson (1846) showing pz
Species ELOPIDAE
Elops machnata (Forssk.)
MEGALOPIDAE Megalops cyprinoides (Brouss.)
CHIROCENTRIDAE
Chirocentyvus dovab (¥orssk.)
DUSSUMIERIIDAE A é Spratelloides gracilis (Schlegel)
CLUPEIDAE (Clupeinae) Sardinella aurita Val.
Sardinella fimbriata (Val.) ?Sardinella leiogastey Val.
Sardinella or Herklotsichthys sp.
(Alosinae) Hilsa veevesit (Rich)
(Pristigasterinae) Ilisha elongata (Ham. Buch.)
(Dorosomatinae)
Clupanodon thrissa (Linn.)
Konosivus punctatus (Schlegel) Nematalosa, ?N. nasus (Bloch)
ENGRAULIDAE (Stolephorinae)
Stolephorus commersonti Lacepede
Thryssa mystax (Schn.)
(Coiliinae)
Coilia mystus (Linn.) Coilia plavfairit (Rich.)
Richardson, 1846 ve
(name)
eee machnata Forssk.
Elops purpurescens Rich.
( Megalops setipinnis Forster
| Megalops curtifilis Rich.
Chirocentrus dorab Forssk.
Clupea gracilis Schl.
Clupea nymphaea Rich. Clupea isingleena Rich. Clupea caeruleo-vittata Rich. Clupea flosmaris Rich.
ie er Rich.
Ns losa palasah Russell
Ilisha abnormis Gray
Chatoessus triza Linn.
ee ee maculatus Gray Chatoessus aquosus Rich. Chatoessus chryvsopterus Rich.
Engraulis commersonianus Lac. Thryssa mystax Schn.
Coilia grayti Rich. Coilia playfairii (McClelland).
RICHARDSON’ S ELOPOITD AND CLUPEOID FISHES
yn numbers in the three sets of Reeves drawings now in the British Museum (Natural History).
Illustrations—page nos.
es (aw na er Reeves Richardson Hardwicke
97, 99 142 30 40 142 31
68 141 234 27,97 I41 = 33 142 237
31 136 222 29, 95 136 219 29, 95 136 223 29, 95 136 oT 102 137 220 96 137 221 29, 48 138 upper — 100 138 lower 240 53, 54 139 232 gI 139 233 29, 95 139 230 29, 95 139 231 92, 97 141 236 8 40 252
31 140 —
‘eeves material, ev Haslar collection.
{
{
Reeves specimens
BMNH. 1964.11.6.3
BMNH BMNH.
BMNH.
BMNH. BMNH BMNH.
BMNH.
1964. 1853.
1964.
1964. 1963.
11.6.14. TYPE* T4220
Tt On dr
11.6.6—7 (TYPE lost) 6.17.1 TYPE
1963.8.20.2 TYPE 1963.8.20.3 1963.8.20.1 1964.11.6.4. TYPE
Vachell specimen (TyPE lost) BMNH. 1964.11.6.5. TYPE
1964.11.6.8-I1 1964.11.6.12
1855.9.19.1581. TYPE* T.964,.shT 62:2,4 1847.5.10.5. TYPE*
24 P. J. P. WHITEHEAD
RICHARDSON ‘*S SPECGLES
The twenty-one species are listed here in the order adopted by Richardson (see Table 1 for correct systematic list). Synonymies list only Hong Kong, Canton and Macao references. Type specimens are redescribed, and also certain species whose systematic position is uncertain or controversial.
t. “ Clupea isingleena Richardson ” = Sardinella fimbriata (Valenciennes)
Clupea isingleena Richardson, 1846, I[chth. China Japan : 304 (China Seas). Spratella fimbriata Valenciennes, 1847, Hist. Nat. Poiss. 20 : 359, pl. 600. Clupea (Harvengula) kowala : Bleeker, 1873, Ned. Tijdschr. Dierk. 4: 147 (on Richardson). Harengula nymphea : Regan, 1917, Ann. Mag. nat. Hist. (8) 19: 392; Fowler, 1941, Bull. U.S. nat. Mus., No. 100: 599, fig. 15; Chu & Tsai, 1958, Quart. J. Taiwan Mus. 11 (1 and 2): 115, pl. 3, fig. 1 [non Clupea nymphaea Richardson—see below}. Type. A fish, 108-5 mm. standard length, until recently unregistered and labelled “ Clupea nymphaea TYPE China’’ (Pl. 1, fig. 1). Now recognized as the lost type of C. isingleena Richardson and registered BMNH. 1963.6.17.1 (see discussion below).
FIGURE. Reeves No. 60 shows a rather deep-bodied clupeoid (depth 2-75 in standard length) not unlike a juvenile Hilsa kelee (Cuvier) in shape but lacking any spots or marks along the flanks (Pl. 1, fig. 2). Dorsal rays vary between 12 and 14 in the three sets of illustrations, and anal rays from 9 to 12. However, the number of finrays shown in these drawings seems to conform more to aesthetic than to scientific standards. The figure shows a fish slightly deeper than either Richardson describes (3 times in length) or is the case with the actual specimen (3-1 times). But there is otherwise sufficient conformity between the figure, the description and the specimen for the three to relate to the same species. Since the drawing and the specimen are both 54 inches, it is quite possible that the specimen was the actual model for the illustration used by Reeves’ artist.
Notes. There is no specimen labelled Clupea isingleena in the British Museum and Giinther (1868, p. 429) reported the same. However, Giinther (loc. cit.) recog- nized a specimen (of 54 inches) as the type of Clupea nymphaea (Pl. I, fig. I), pre- sumably because it was then labelled as such, and until now this has been accepted as the type of C. nymphaea (e.g. by Regan (1917) in his revision). However, Giinther noted that the anal ray count (21) in this specimen tallied not with the description and figure of C. nymphaea but with that of C. zsingleena, as also did the size of the specimen. He then states that the “ one example in the British Museum
belongs, on account of its oblong form, to the figure named Cl. nymphaea”’, a statement which is certainly not true of the present specimen labelled C. nymphaea ; neither does it accord with Giinther’s own description of this specimen (“ the height of body thrice and one sixth ”’ in standard length). In the figure of C. zsvvgleena the body depth is contained 2-75 times in standard length, against 3-65 times in the figure of C. nymphaea (3°r in the actual specimen, “ thrice in the length’’ according to Richardson’s description of C. isingleena).
Since Richardson gave an adequate description of C. isingleena, based on a specimen (of 54 inches) and on a drawing (also 54 inches), and at the same time indicated the
RICHARDSON’S ELOPROID AND CLUPEOID FISHES 25
museum in which the specimen was deposited ; and since there is a British Museum specimen (of 54 inches) which conforms with both drawing and description in almost all respects ; then Giinther’s type designation must be recognized as wrong, being based on a curatorial error. The specimen in question has 8 pelvic rays. A pelvic count of g (described for C. nymphaea) is known only in one species of Sardinella, namely S. aurita, which in fact is a slender species such as is described and figured for C.nymphaea. In S.aurita too, the anal count is low (15-19 fide Regan, 19170) and thus agrees with Richardson’s C. nymphaea (A.15 vel 16) not with his C. 1semgleena (A.21).
One slight anomaly, however, is in Richardson’s scute count of 16 + 10 for C. isingleena ; the specimen has 18 + 12, although one post-pelvic scute is very small. But even in S. aurita there are at least 13 post-pelvic scutes.
It must be accepted, therefore, that the specimen long known as the type of Clupea nymphaea Richardson is in fact the lost type of C. tsimgleena Richardson. However, the latter name, although pre-dating all other names for this species, has not been used as a senior synonym since Richardson’s time and is therefore a nomen oblitum.
The type specimen was considered a species of Harengula by Regan (19170) and subsequent workers (e.g. Fowler, 1941; Chu & Tsai, 1958). But it is clearly a member of Sardinella, having 8—1o fronto-parietal striae and upper and lower parts of the 2nd supra-maxilla similar in shape and size (see Whitehead, 1964a, 1964c for diagnosis). The vertical striae on the scales (a character used by Regan, 19170) resemble those in Harengula or Herklotsichthys, appearing to be continuous across the scale, but in fact in most cases the inner ends of the striae do not meet in the centre but overlap each other (Chan, 1965, fig. 8). The anal, however, is too poorly preserved to judge whether the antepenultimate ray is significantly shorter than the final two rays (a Sardinella character). It is interesting to note that in neither Fowler’s drawing (1941, fig. 15) nor in the description of this species by Chu & Tsai (1958) are the last two anal rays indicated as enlarged (‘‘ somewhat larger ’’ according to Chan, 1965). Regan (1917b) placed emphasis on this character in his differentia- tion between Sardinella and Harengula, but in this species at least, it does not appear to be diagnostic.
The search for the correct name for the present species is by no means simple. Chu & Tsai (1958) list six species of Sardinella in the Taiwan area, as well as Harengula nymphaea. Of the four with 8 pelvic rays and sharply keeled scutes (i.e. excluding S. aurita and S. sivm respectively), none has a gillraker count above 63, whereas the specimen in question has 71 (or 69, Chan, 1965). A count of 69-81 and a body depth of 2-99-3°53 is given for S. fimbriata (Valenciennes) by Chan (loc. cit.) in his revision of the genus, and until further studies on this genus are published, this should be considered the next available junior synonym. Bertin (1944) concluded that the Valenciennes types of Clupeonia jussieut and Spratella fimbriata represented a single species (both with about 70 gillrakers). The former name has page priority over the latter, but until the identity of Clupanodon jussieu Lacepéede is established (or the name rejected), it is best that the name fimbriata be retained for the present since authors have used the name jusszew for another species (i.e. for Bleeker’s species gibbosa ; see also discussion in Whitehead, 19650).
26 P. J. P. WHITEHEAD REDESCRIPTION OF TYPE OF CLUPEA ISINGLEENA
Standard length : 108-5 mm. Total length : 140 mm. or 54 inches (estimated since caudal tips damaged).
mm. ois Body depth : : : : : : : 35°2 32°4 Head length : : , : : , 28-3 26:2 Snout length : : ‘ : ‘ : : 5°8 5°3 Eye diam. . : ‘ : . : 8-0 7*4 Upper jaw 1. : : : ¥ : : ; pomre) II-'o Lower jaw 1. : : : : : : : I2°4 II°+5 Pectoral length . : : : : : 3 damaged — Pelvic length . ° ; : ‘ F 3 TIS 10-9 Pre-dorsal é é : : : : : 49°1 45°3 Pre-pelvic . : : : : ; : : 59°2 54°5 Pre-anal_. : : : : : : ; 88-5 81-5
Body strongly compressed, its depth greater than head length, snout less than eye diameter. Maxilla, reaching to eye centre but not to articulation of lower jaw, lower edge with a few minute denticulations ; cxposed portion of maxilla with 4-5 longitudinal ridges ; 2nd supra-maxilla “ paddle-shaped ’’, upper and lower parts of expanded portion similar in size and shape, length of expanded portion equal to depth ; rst supra-maxilla slender. Pseudobranch short, its length about ? of eye diameter, ventral border without prominent ridge.
Dorsal surface of head with well-defined cuneiform fronto-parietal areas with 8—ro longitudinal striae. Bilobed dermal outgrowth on vertical portion of cleithrum and well-developed cleithral lobe along lower border of gill opening. Gillrakers 41 + 71 on first arch, about equal to gill filaments, neither upper nor lower series overlapping the other ; gillrakers present on posterior face of 3rd epibranchial.
No teeth in jaws but a single longitudinal series of fine teeth along tongue and numerous fine papillae on rest of tongue ; fine teeth present on pterygoids.
Dorsal origin nearer to snout than to base of caudal, but entire dorsal base equi- distant between snout and caudal base ; dorsal rays iv 14, tips damaged. Pectoral with i 15 rays, tips damaged. Pelvic, with i 7 rays, its base under middle of dorsal and equidistant between pectoral base and anal origin. Anal with ii 18 rays, its origin a little nearer to caudal base than to pelvic base.
Scutes sharply keeled, 18 pre-pelvic, 12 post-pelvic. Many scales missing, ex- posed portions with fine perforations and longitudinal ridges leaving an almost pectinated posterior border (two scales from type figured by Chan, 1965, fig. 8).
Colour in alcohol : upper + of body brown, flanks silver, fins hyaline ; no indication of dark markings.
2. ‘ Clupea nymphaea Richardson ”’
= Sardinella aurita Valenciennes
Clupea nymphaea Richardson, 1846, Ichth. China Japan : 304. Sardinella aurita Valenciennes, 1847, Hist. Nat. Poiss. 20 : 263, pl. 504 (name retained for reasons of stability—see below) ; Regan, tot7, dan. Mag. nat. Hist. (8) 19 : 578 (including Giinther’s
RICHARDSON’S ELOPOID AND CLUPEOID FISHES 27
two specimens of C. melanostictus from China—see below) ; Fowler, 1931, Hong Kong Nat. 2 (2) : 116 (compiled, no Chinese specimens).
Clupea melanosticta : Giinther, 1868, Cat. Fish. Brit. Mus. 7 : 430 (2 Reeves fishes and 1 juvenile
ex China). Type. No British Museum specimen. As shown under C. tsingleena, the speci-
men hitherto labelled as type of C. nymphaea (see Giinther, 1868, p. 428; Regan,
IQI7, p. 392) is in fact the type of C. ¢s¢ngleena.
FicurRE. Reeves No. $25 shows a rather slender clupeoid resembling Sardinella sirm or S. clupeoides (Pl. I, fig. 3). It lacks any black spots or marks along the flanks such as occur in Sardinops. Pre-pelvic scutes are not shown, but there are 14 or I5 post-pelvic scutes (only 12 in the Hardwicke illustration). The anal base is moderate, about equal to dorsal base, and 14 rays are shown. The figure is 62 inches in total length (140 mm. standard length).
Notes. The identity of this species has hitherto remained doubtful, partly due to the confusion over the type specimen. However, Richardson describes 9 pelvic rays and this immediately excludes all species of Herklotsichthys (Harengula auct., see Whitehead, 19642) and also all species of Sardinella except S. aurita Valenciennes, 1847. Of clupeoid species with rather elongate bodies, Sardinops melanosticta (Schlegel) can be ruled out since it has only 8 pelvic rays and the series of black spots along the flanks are obvious even in long preserved material. Sardinella sirm (Walbaum) is similar in form to the Reeves illustration, but it has only 8 pelvic rays and there is a series of dark spots along the flank. I have found no record of S. sivm from the Hong Kong area. Finally, Clupea harengus pallasi can be considered. But although the check list given by Liang (1951) of specimens in the Provincial Fisheries Institute in Taiwan suggests that this species is present in the area, Chu & Tsai (1958), in a review of Taiwan clupeoids, found no evidence of C. harengus and believed that Liang’s specimens were from Japan. It is most unlikely that C. harengus would penetrate as far south as Macao.
By elimination, therefore, Richardson’s C. nymphaea can only be Sardinella aurita. Richardson’s name, however, pre-dates that of Valenciennes, and should strictly replace it; it is not a nomen oblitum, having been in constant (mis)use. But the species is the most widespread and commercially important of all Savdinella species. In the interests of stability, therefore, it will be recommended to the International Commission that Richardson’s name should be suppressed, and the name S. aurita Valenciennes retained.
There are two specimens of S. aurita in the British Museum which are labelled “ Clupea melanosticta (TYPES) China Reeves’”’. They are adult fishes (160 mm. S.L.), until now unregistered, but now BMNH. 1964.11.6.67. They were listed, as Reeves specimens, by Giinther (1868, p. 430) under the name Clupea melanosticta Schlegel. This was a misidentification, since C. melanosticta Schlegel is a species of Sardinops. Giinther included a third specimen, a juvenile of 65 mm. S.L. also from China. The latter is too small and was registered too late (1851) to have been the missing Richardson type of C. nymphaea.
Giinther (loc. cit.) placed C. caeruleovittata Richardson in his synonymy of C. melanosticta Schlegel. Richardson lists no specimens of C. caeruleovittata, so the
28 Pia SRS WwWAIEEE READ
present specimens cannot be types of that species. The original label on the bottle containing these two Reeves specimens is now missing (the bottle was relabelled after the war, presumably copied from the old label which had become detached). The designation of these specimens as types seems to have occurred after Giinther had listed them. They are not types of C. nymphaea since Richardson only refers to a single “‘ Specimen in Br. Mus.”’ ; neither are they the types of Clupea melanosticta Schlegel, which are in Leiden (Boeseman, 1947). They may, perhaps, have been sent to the British Museum by Reeves’ son after the “‘ Report ’’ was written.
,
3. “ Clupea caeruleo-vittata Richardson ’ = Sardinella, probably S. leiogaster Valenciennes
Clupea caeruleo-vittata Richardson, 1846, Ichth. China Japan : 305 (on Reeves illustr.). Sardinella leiogastey Valenciennes, 1847, Hist. Nat. Poiss. 20: 270; Kner, 1865, Reise Novara,
Fische : 327 (Hong Kong).
? Harvengula moluccensis : Jouan, 1867, Mém. Soc. Imp. Sci. nat., Cherbourg, 13 (2° ser.) (3) : 272
(Hong Kong).
Clupea (Amblygaster) melanosticta : Bleeker, 1873, Ned. Tijdschr. Dierk. 4: 147 (on C. caervuleo- vittata Richardson).
Sardinella sirm : Fowler, 1931, Hong Kong Nat. 2 (2) : 119 (solely on Kner, 1865 for Hong Kong record).
SPECIMENS. Nospecimens mentioned by Richardson. There isnonein the British Museum, except the two Reeves specimens labelled ‘‘ Clupea melanosticta TyPEs”’ mentioned under the previous species. Since Giinther placed C. caeruleovittata Rich. in his synonymy of C. melanosticta, and since he listed two Reeves specimens, it is odd that these two fishes are not labelled as types of C. caeruleovittata. However, in certain cases, the most recent bottle label has been a copy, not of the original name on the old label, but of the name as later amended (by Giinther, Regan, Norman, etc.). Unfortunately the original label has gone.
FIGURE. Reeves No. 59 shows an even more elongate clupeoid than the previous species, a fact commented on by Richardson (PI. 2, fig. 1). In appearance it suggests a gravid female with distended abdomen. It resembles Etrumeus teres (DeKay), which occurs in Hong Kong waters (see Table 2), but the pelvics are set below the dorsal base, not well behind it. Of the remaining elongate clupeoids recorded from this area, the following can be considered : Sardinops melanostictus, Clupea harengus, Sardinella aurita, S. siym and S. leiogaster. The first can be eliminated on grounds of coloration, there being no spots shown on the flanks in Reeves’ illustration. The second can also be ruled out on geographical grounds (see discussion under C. nymphaea). The Reeves drawing is not accurate enough for a pelvic finray count, which would distinguish S. aurita (9) from S. stym and S. leiogaster (8). But since S. aurita is already represented in Richardson’s list (as C. #ymphaea) under the Chinese vernacular name Chang yaou lin (“ long-waisted scale ’’ or “ long fine waist ’’), it can be argued that C. caeruleovittata, the Huang-tsih (‘ yellow glossy’) of the “ Report ’’, must therefore be another species. Of the two remaining elongate clupeoids, Sardinella letogaster seems the more likely. It is a slightly deeper fish and lacks the row of dark blue spots along the flanks described in S. sivm (see Chu & Tsai, 1958).
RICHARDSON’ S ELOPOID-AND CLUPEOID FISHES 29
Another possibility is S. clupeoides (Bleeker, 1849). Unhke Bertin (1944), Chan (1965) recognised this species as distinct from S. levogaster ; it has the dorsal origin nearer to snout tip than to caudal base, a feature well shown in the Reeves drawing (Pl. 2, fig. 1). However, there are no records of this species from the Hong Kong area.
Harengula moluccensts of Jouan (1867) from Hong Kong may refer to the present species, although “le ventre non caréné”’ is suggestive of Etrumeus teres. There are records of E. teres from Hong Kong (Whitehead, 1963a p. 374), and Jouan (loc. cit.) states that his fish is common (in October) (4 Hong Kong juveniles in British Museum sent by Chan). However, his pelvic count of 7 fits neither Etvwmeus nor Sardinella.
Richardson’s name caeruleovittata predates Valenciennes’ name lerogaster. How- ever, it has now become a nomen oblitum, and in view of the difficulty in making a correct identification of Richardson’s species, no purpose would be served in attempt- ing to resurrect this name.
4. “ Clupea flosmaris Richardson ”’ = ?Herklotsichthys sp. or Sardinella sp.
Clupea flosmaris Richardson, 1846, I[chth. China Japan : 305 (on Reeves’ illustr.).
SPECIMENS. None mentioned by Richardson, and none in British Museum.
FIGURE. Reeves No. 64 shows a clupeoid of moderate body depth in which the scales appear to have been lost (Pl.2, fig. 2). Fowler (1931, p. 112) identified this species with Spratelloides delicatulus (Bennett) although admitting that the figure (6 inches) is much too large “ due to an exageration of the artist’s drawing ’’. The fish shown is too deep for S. delicatulus, and is most likely a juvenile Sardinella or Herklotsichthys. Richardson compared this fish with one described and figured in the “ Description of Animals ’’, p. 201, fig. 149 (see ‘‘ Report ”’ for note on this work.) This latter description gives the following finray counts: D 13, A 19, C 14, P 10, Vg. But the anal count, and the serrated belly in the figure, rule out a round herring ; the pelvic count is virtually diagnostic of Sardinella aurita. However, the Reeves illustration shows an anal count of only 9, and there is no real evidence that the Reeves drawing is of the same fish as that in the “ Description of Animals ”’
The most that can be said of this Reeves illustration is that it is of a juvenile clupeoid, probably a species of either Sardinella or Herklotsichthys.
5. ‘ Clupea gracilis Temm. et Schl. F. J. Sieb.” = Spratelloides gracilis (Schlegel)
Clupea gracilis Schlegel, 1846, Faun. Japon. Poiss., pt. 5, pl. 108, fig. 2 (Japan) ; Richardson, 1846, Ichth. China Japan : 305 (? Japan).
SPECIMENS. Richardson examined a British Museum specimen (in bad condition) labelled “‘ Clupea gracilis’ and concluded that he could not “ identify it with any
of the preceding species’”’. I have been unable to determine which specimen this
30 P. J. P. WHITEHEAD
might be, but Giinther (1868, p. 465) lists three Japanese specimens (registered BMNH.4.6.8134). There are Hong Kong specimens in our collection, and three Taiwan specimens have been described (Whitehead, 19630, p. 343).
Ficure. No Reeves illustration.
Note: I can find no published reference to this species from Hong Kong, although it is well recorded from Japan, the Philippines and also Taiwan. Fowler (1931, p- 112) reports the closely related S. delicatulus (Bennett) from China, but as noted earlier, this is based solely on a misidentification of Richardson’s C. flosmaris. There are now a number of specimens of S. gracilis in the British Museum (BMNH. 1965.7.5-49-70) sent by W. L. Chan from Hong Kong and the species is probably not uncommon there. Whereas S. gracilis and S. delicatulus occupy roughly the same range in the Indian Ocean and along the shores of the Indo-Malayan Archi- pelago, in the western part of the Pacific their ranges diverge. S. delicatulus has the more southerly distribution, extending southwards to Tasmania, while S. gracilis reaches further north (to Japan). S. delicatulus is recorded from the Philippines (see Fowler, 1941, p. 562), but probably does not reach Hong Kong or Taiwan.
5)
6. “ Alosa reevesii Richardson ’ = Hilsa reevesii (Richardson)
Alosa veevesit Richardson, 1846, Ichth. China Japan : 305.
Alausa reevesii : Valenciennes, 1847, Hist. Nat. Poiss. 20: 437 (dry specimen from Macao) ; Jouan, 1867, Mém. Soc. Imp. Sci. nat. Cherbourg, 13 (2) (3) : 271 (Hong Kong).
Hilsa veevesti : Fowler, 1931, Hong Kong Nat. 2 (2): 115 (China, compiled; no Hong Kong specimens) ; Whitehead, 1964, Bull. Brit. Mus. (nat. Hist.) Zool. 12 (4) : 141 (revision; type and Reeves and Hong Kong specimens).
Alosa palasah: Wichardson, 1846, Ichth. China Japan : 306 (Reeves specimen).
Alausa palasah: Jouan, 1867, Mém. Soc. Imp. Sci. nat. Cherbourg, 13 (2) (3) : 271 (Hong Kong).
Type. A fish, 295 mm. standard length, ev China, presented by J. R. Reeves, until recently unregistered, but now BMNH. 1963.8.20.2. Gziinther (1868, p. 447) regarded this specimen as the type and mentioned a smaller Reeves specimen (140 mm. standard length, now BMNH. 1963.8.20.3). The latter, not the former, has until now been labelled as the type in our collections, but this is wrong: the smaller specimen is not mentioned by Reeves. Both are mounted skins.
Figure. Of the three copies of Reeves’ figure «8 in the British Museum, that found in the Hardwicke set is the best, but all are adequate to identify the species (Pl. 2, fig. 3). Richardson states that ‘“ Mr. Reeves deposited a specimen in the British Museum which still retains the original label numbered in reference to his (Reeves’) drawing ’’. It is not clear from this whether the drawing (17 inches) was made from this actual specimen (15 inches). There is no label now attached to the fish, but a pencilled note on the underside of the stand on which the specimen is mounted gives the Reeves number, name, locality and collector.
Note. The holotype is clearly H. veevesit, not H. tlisha, having the broad oper- culum characteristic of the former (see Whitehead, 1965@)._ In addition, there are
RICHARDSON’S ELOPOID AND CLUPEOID FISHES 31
only 15 branched anal rays (18-20 in H. alisha) and the anal origin is equidistant between the caudal base and pelvic tzps (pelvic base in H. tisha).
Richardson records too few scales in lateral series (““ Thirty of them compose a longitudinal row’’), since there are at least 40 in the specimen, probably more. Hilsa toli (Valenciennes) also occurs in this region (see Table 2), but it has a narrow operculum (as in H. tlisha) and the caudal lobes exceed head length (about equal to head length in H. ilisha and H. reevesit).
RIDE SCRIPTION OF THE fYPE OF CLUPEA REEVESTII Standard length : 295 mm. Total length : 369 mm., or 144 inches (approx. I5 inches according to Richardson).
mm. Caorles
Body depth : : : : : : : gI-o 30:7 Head length : : : : : : : 84°5 28°7 Snout length ‘ ‘ : : F ; ; 21°5 73 Eye diam. : : : : F : : 12°8 4°3 Upper jaw 1. ‘ : : : : : : 35°0 TL+9 Lower jaw 1. ; : : : : : : 43°2 14:7 Pectoral 1. ‘ 3 F : : : : 47°90 15°9 Relvic l= ©. : : : F : ; : 24°5 8-3 Operculum height : : : : : : 35°5 12-0 width : : : : : : 24°0 8-2
Pre-dorsal : : : : : : : 149°4 50°5 Pre-pelvic . : : ; : : : F 143°8 48°5 Pre-anal_ . : : ‘ : : : , 239°0 81-0
Dorsal ili 14 (or iv 13, tips broken) ; pectorali 15; anal iii 15; scales in lateral series about 41. In all other respects, this specimen conforms to the description given by Whitehead (19652).
7. “ Alosa palasah Russell ”’ = Hilsa reevesii (Richardson) (See previous synonymy.)
SPECIMEN. A fish, 140 mm. standard length, ex China, collected and/or presented by J. Reeves. Hitherto unregistered, now BMNH. 1963.8.20.1. The bottle still bears two of the original labels, the first reading “‘ Clupea reevesii China J. Reeves Esq.”’ and the second “‘ Reeves B51 Hard. 221’’. Richardson mentions a British Museum specimen of 7 inches ; the present specimen is 7} inches in total length and is undoubtedly that examined by Richardson.
FIGURE. Reeves No. /51 shows a fish of 12 inches (305 mm.) and since the Reeves illustrations were usually life-size, the British Museum specimen is probably not the model for the drawing. The illustration shows well the broad operculum character- istic of this species (Pl. 3, fig. 1). The lower third of the body is shown as dark bronze, the upper part silver—the reverse of what wouid be expected.
Note. Richardson identified this fish rather tentatively with Russell’s Palasah and at the same time distinguished it from his own Alosa reevesii because of differences ZOOL. I4, 2 4
32 Puy. PW EE Ee AD
in head length, pectoral length and body outline. The specimen is clearly H. veevesii and not the Palasah of Russell (1803) (i.e. Hilsa alisha (Ham. Buch.)—see Whitehead 1965a), having a broad operculum, its width contained only 14 times in its length. The differences from H. rveevesii noted by Richardson arise from a comparison of juveniles and adults and can be accounted for by allometric growth. Richardson records 40 scales in longitudinal series (42-45 in H. reevesii ;_ cf. 45-48 in H. tlisha— see Whitehead 19652).
8. “ Ilisha abnormis Gray ”’ = Ilisha elongata (Bennett)
Alosa elongata Bennett, 1830, Mem. Life of Raffles : 691.
Ilisha elongata: Norman, 1923, Ann. Mag. nat. Hist. (9) 11:7 (revision; Reeves’ specimen examined) ; Fowler, 1930, Proc. Acad. nat. Sci. Philad.: 599 (Hong Kong); Idem, 1931, Hong Kong nat. 2 (2) : 121 (Hong Kong specimens).
Llisha abnoymis Richardson, 1846, Ichth. China Japan : 300.
Pellona vimbella Valenciennes, 1847, Hist. Nat. Poiss. 20: 317 (specimen from Macao).
Pellona grayana Kner, 1865, Reise Novara, Fische : 328 (Hong Kong).
Type. A mounted skin (right side), 295 mm. standard length (total length about 14 inches, caudal tips damaged), ex China Seas, presented by J. R. Reeves. Hitherto unregistered, now BMNH. 1964.11.6.4. Underside of base of wooden stand marked “ Tlisha abnorma H 240 R 81’’. The specimen is in poor condition, the anterior part of the flank lacking scales and having been sewn up across a large split in the skin and across the gill opening ; pectoral fin detached and sewn loosely to body; a specimen of 144 inches in British Museum (“‘ dried and varnished ’’) mentioned by Richardson.
TiGurRE. Reeves Nos. 81 and 67 are respectively a little larger and a little smaller than the type specimen. Both are recognizably J. elongata (Pl. 3, figs. 2 and 3). Detail of head poor in both illustrations.
Note. From the keys and descriptions given by Norman (1923) and Chu & Tsai (1958), there is no doubt regarding the identity of the type specimen, assuming of course that Richardson’s counts were reasonably accurate. Unfortunately, scale and scute counts are no longer possible. Three other species are reported from China. I. indica (Swainson) is a deeper-bodied fish (depth 23-34 1n length ; cf. 3$in the type of I. abnormis) with 39-45 scales in lateral series (50-55 in J. elongata, and “ about fifty scales ’’ described by Richardson in the holotype of J. abnorntis).
A second possible species is J. novacula (Valenciennes), again a rather deeper-bodied species (34 in length) with fewer scales (45). J. brachysoma (Bleeker) is also a deeper bodied species with fewer scales in lateral series (42-45). In all three cases, the more slender body in Richardson’s specimen distinguishes it, even if the meristic counts are not completely accurate,
The names Jlisha and abnormis were created by Gray in an unpublished catalogue of British Museum specimens. This catalogue is not now in the Zoological Library of this museum, and appears to have been lost. It was a continuation from the “ Chondropterygii ”’ of Gray (1851).
REGHAR DSONS ELOPOLD AND CLUPEOLD FISHES 33 RE DES CR LP RION OF TRE TYPE OF TILISHA ABNORMIS
Standard length : 295 mm. Total length: 356 mm. or 14 inches (approx., caudal tips damaged).
mm. Sr Body depth : : ; : ‘ : : 76:0 25°8 Head length ; 3 ; : : : : 67°8 230 Snout length ; ‘ P ; ‘ F : I9°6 6°5 Eye diam. : : E ; : 3 : 16-0 5°4 Upper jaw l. : : : F ‘ ; : 34°70 II-5 Lower jaw l. : : ; : : : ; 36°1 I2°2 Pectoral length . : : . : : E 37°5 12°7 Pelvic length . ; ; ; : : : 1-4 3°9 Length of anal base. : : : : ‘ 118-0 40-0 Pre-dorsal ; : : : : : ; 150°5 51-0 Pre-pelvic . < : : ‘ 2 : : TO 39°5 Pre-anal . : : ‘ : : ‘ : L775 60-0
Lower jaw strongly projecting, upper jaw with two supra-maxillae but no hypo- maxilla, extending to middle of eye if mouth closed, but not reaching to articulation of lower jaw.
Dorsal with iv 15 rays, its origin equidistant between snout tip and caudal base. Pectoral i 15. Pelvic (rays damaged) shorter than eye, its base a little nearer to pectoral base than to anal origin. Anal with i 45 rays, its origin under last dorsal ray. Scales—no count possible. Scutes, 21 pre-pelvic, no count possible on post- pelvic (Richardson gives 14 + 13, but evidently missed seven pre-pelvic scutes).
’
g. ‘‘ Chatoessus aquosus Richardson ’ =Konosirus punctatus (Schlegel)
Chatoessus punctatus Schlegel, 1846, Fauna Japon. Poiss., pt. 5: 240, pl. 109, fig. 1.
Nealosa punctata : Herre & Myers, 1931, Lingnan Sci. J. 10 (2 and 3) : 236 (3 Hong Kong speci- mens).
Konosivus punctatus : Whitehead, 1962, Bull. Brit. Mus. (nat. Hist.) Zool. 9 (2): 100 (generic
review).
Chatoessus aquosus Richardson, 1846, Ichth. China Japan : 307.
Type. A dried skin (left side) mounted on wood, 190 mm. standard length, ex China, presented by Reeves. Hitherto unregistered, now BMNH. 1964.11.6.5 The specimen is in poor condition : pectoral detached, dorsal and anal fins damaged, caudal entirely missing. On the reverse side it is marked “ H 230 R 63’. This is the only specimen mentioned by Richardson, who gives its length as 7? inches.
FicurRE. Reeves No. 63 shows a clupeoid with a filamentous last dorsal ray, but with a rather rectangular suboperculum (PI. 4, fig. 1). No scutes are shown, and there are about 45 scales in lateral series. Either Konosivus punctatus or Clupanodon thrissa are possible, but it must be presumed that the figure agrees with the specimen.
Note. The specimen conforms to the diagnosis of the monotypic genus Konosirus (Whitehead 1962a, p. 100) in the following characters :
34 P. J. P. WHITEHEAD
a. Last dorsal ray, although now broken, sufficiently stout to have been fila- mentous.
b. Suboperculum with (exposed) anterior and upper margins meeting at obtuse angle, posterior margin rounded (exposed part of suboperculum rectangular in Nematalosa).
c. Outer edge of dentary not strongly flared outwards.
d. Post-pelvic scutes 14 or more likely 15 (cf. 11-12 in Clupanodon thrissa).
REDESCRIPTION OF THE TYPE OF CHATOESSUS AQUOSUS
Standard length : 190 mm.
mm. ous Body depth 2 : ; : : 6 : 57°0 30-0 Head length 5 : : : : : ; 47°4 24°7 Snout length ‘ F : ; F ‘ : 9°7 5:1 Eye diam. : : : é : : : 9°3 4°9 Upper jaw l. ; : : : : : : 15°0 7°9 Lower jaw 1. : c : ; ; : ; 19-0 10-0 Pectoral 1. . é : . : 3 : 33:0 17°4 Pelvicl. . ; : : : ; : : 17°0 8-9 Pre-dorsal : : : : : ; : 92-9 48-7 Pre-pelvic . : : : ; ; i : 97°2 51-1 Pre-anal . : ¢ : : : : : 146°2 78-0
Dorsal with ii 14 (or 1v 13) rays, its origin slightly nearer to caudal base than to snout tip. Pectoral with i 14 rays. Pelvic with i 7 rays, its base below first un- branched dorsal ray, and slightly nearer to pectoral base than to anal origin. Anal (count impossible) moderate, its base slightly longer than dorsal base, about I} in head length. Caudal missing.
Scutes, Ig pre-pelvic, 15 post-pelvic (about 15 + 13 according to Richardson). Scales in lateral series, about 43 or 44 (Richardson, 46).
In all other features (jaws, opercular bones, etc.) this specimen conforms to the diagnosis for Konosirus given by Whitehead (1962a, p. 100).
The priority of Richardson’s aquosus over Schlegel’s punctatus need not be adhered to since the former is a nomen oblitum and there would be little value in resurrecting the name.
to. ‘“‘ Chatoessus triza Linnaeus ”’ = Clupanodon thrissa (Linnaeus)
China, on
[Mystus altus Linnaeus, 1754, Chinensia Lagerstriémiana— dissertatio’’ : 26 Lagerstrom. |
[Clupea thrissa Osbeck, 175
Clupea thrissa Linnaeus, 1 Lagerstrom).
Clupanodon thrissa: Fowler, 1930, Proc. Acad. nat. Sci. Philad. : 5909 (Hong kong) ; Idem, 1931, Hong Kong Nat. 2 (1) : 76 (Hong Kong specimens) ; Herre, 1934, Hong Kong Nat. Supplement, No. 3: 26 (3 Hong Kong specimens) ; Whitehead, 1962, Bull. Brit. ATus. (nat. Hist.) Zool. 9 (2) : 100 (generic review).
7, Dagbok Ostind. Resa : 257—Canton area. ] 758, Syst. Nat., ed. 10: 318 (name from Osbeck; diagnosis after
RICHARDSON’S ELOPOID AND CLUPEOID FISHES 35
Clupea tviza Linnaeus, 1759, Amoen. Acad. 4: 251. Chatoessus triza: Richardson, 1846, Ichth. China Japan : 307 (on Reeves illustration, Canton or Macao). Chatoessus maculatus Richardson, 1846, Ichth. China Japan : 308 (specimen from Canton). Chatoessus osbeckii Valenciennes, 1848, Hist. Nat. Poiss. 21: 106. SPECIMENS. None are mentioned by Richardson and there are no Reeves speci- mens in the British Museum.
FiGuRE. Reeves No. 224 shows a fairly deep fish (deeper than the preceding species), with an elongated last dorsal ray, a slender maxilla reaching almost to eye centre, and a dentary which is barely flared (Pl. 4, fig. 2). Mouth shape clearly eliminates Nematalosa, and two possibilities remain, the monotypic genera Konosirus and Clupanodon.
The illustration can be definitely identified on three characters.
a. Snout. The snout is less pointed than in the preceding figure (Pl. 4, fig. I) and is a fair representation of the blunter snout found in C. thrissa compared with that in Kk. punctatus.
b. Operculum. In C. thrissa the operculum is shghtly broader than in Kk. puncta- tus and its lower edge is a little higher up on the body. Although the opercular series in Reeves’ drawings 63 and 224 are not completely accurate, yet the overall impression gives a good illustration of the differences in opercular shape between the two genera (compare Pl. 4, figs. 1 and 2).
c. Anal base. Again the artist has not made an accurate drawing, but the longer anal base in Reeves No. 224 (just over head length ; about head length or a little less in specimens of C. thrissa) can be contrasted with that shown in Reeves No 63 (about # of head in both figure and in specimens of x. punctatus).
In addition, Reeves’ illustration No. 224 shows a slightly deeper fish than in the illustration of K. punctatus (No. 63). C. thrissa is indeed a slightly deeper fish (depth 22-3 in standard length ; cf. 3-34 1n K. punctatus according to Regan, 19174).
There can be little doubt, therefore, that the illustration refers to C. thrissa and not to K. punctatus, although Reeves himself was apparently dissatisfied with the painting. In his notes (list of illustrations, dates, etc.—see section on Reeves’ illustrations), he states “7th June [1828] Clupea sp. now transparent as glass—this badly painted.”
Note. Richardson took the name éviza from Linnaeus’ Chinensia Lagerstrémiana (1759), not from the Systema Naturae (1758), but that was incorrect. The earliest names, Mystus altus L., as well as Clupea thrissa Osbeck, were inadmissible (/nt. Code Zool. Nomen. 1961, Art. 3).
The synonymy is cited here because the species was largely based on Osbeck’s good description which was written (in November 1750) after examining (a) speci- men(s) from the Canton area. In the two Lagerstrém descriptions by Linnaeus there is no mention of a filamentous last dorsal ray, but this is given in the Systema Naturae, presumably on Osbeck’s description (‘‘ quorum ultimo duplo longior ’’). Lonnberg (1896) was able to identify a specimen of Clupea tviza from the Lagerstrom collection at Uppsala ; he too, does not mention the filamentous last dorsal, which was presumably broken off.
36 P. J. P. WHITEHEAD
11. “ Chatoessus chrysopterus Richardson ”’ = Nematalosa sp., ? N. nasus (Bloch)
Clupea nasus Bloch, 1795, Nat. Ausl. Fische, 9: 4209, fig. I.
Nematalosa nasus : Herre, 1934, Hong Kong Nat. Supplement, No. 3 : 26 (1 Hong Kong specimen).
? Chatoessus chrysopterus Richardson, 1846, Ichth. China Japan : 308 (on Reeves’ illustration— Canton or Macao).
Konosivus thrissa: Jordan & Seale, 1905, Proc. Davenport Acad. Sci. 10: 2 (Hong Kong) (non Clupea thrissa L.).
SPECIMENS. None mentioned by Richardson and no Reeves specimens in British Museum.
FiGuRE. Reeves No. 61 shows a deep-bodied fish (depth 24 in standard length) with an elongated last dorsal ray and an inferior mouth with a short maxilla (Pl. 4, fig. 3). This is clearly a species of Nematalosa, and three species can be considered, N. nasus, N. japonica Regan and N. come (Richardson). There are two principle features shown in the illustration which may help to identify this drawing.
a. Body depth: the body shape shown in the illustration strongly suggests N. come (depth 2—24 in length according to Regan, 1917a). In N. japonica, a more slender species judging from the type specimens in this museum, the depth is con- tained 3 times in length and the head 14 times in body depth (1# in the illustration). Nematalosa nasus is also rather deep-bodied (depth 22—24 times in length according to Regan, 19174).
b. 2nd suborbital : in the illustration, the anterior border of the 2nd suborbital is shown as slightly oblique (rather than vertical) and the edge is concave (rather than straight or convex). In N. nasus alone, the anterior border is vertical and slightly convex (see Whitehead, 1962a, fig. 4). In all other species the anterior border is oblique, and leaves exposed a small triangular area above the anterior part of the lower limb of the pre-operculum. Such a naked area is not, however, shown in the drawing.
Note. Nematalosa japonica can be eliminated because of its more slender body. The only record of N. japonica from the Hong Kong area seems to be the single specimen examined by Herre & Myers (1931). The Reeves figure probably best fits N. come, but this species is not known from so far north (Indo-Australian Archipelago according to Regan). It differs from N. japonica in having a deeper body and a lower scute count (11-14 post-pelvic scutes ; cf. 13-16—see Whitehead, 1962a@), but a more strongly flared dentary. Unfortunately, neither of these two features can be determined from the drawing. Finally, it must be wondered whether Richardson would not have recognized his own species (i.e. N. come), especially since he comments on the close correspondence between the ichthyofauna of the northern and southern parts of the western shores of the Pacific (“ Report ’’, p. 190).
Nematalosa nasus, the remaining possibility, is recorded from the Philippines, China and Japan, and two Hong Kong references appear in the literature (see syn- onymy). There is a Hong Kong specimen in this museum. The discrepancy in shape of 2nd suborbital might be misinterpretation by the artist, for certainly the dermal head bones are not accurately drawn in any of the drawings. A mistake of
RICHARDSON’S-ELOPOID AND CLUPEOID FISHES 37
this kind seems much more likely than one in body depth, and the drawing agrees with N. nasus and not with other species of Nematalosa in failing to show the small naked, triangular area above the anterior part of the pre-operculum. Nematalosa nasus is therefore chosen as the most likely species.
12. “ Chatoessus maculatus Gray ”’ = Clupanodon thrissa (Linnaeus) (see synonymy under Chatoessus triza.)
SPECIMENS. Richardson states that a single specimen was presented by Vachell to the Cambridge Philosophical Institution. There is now no such specimen in the Cambridge collections, and Giinther (1868, p. 409) does not list any Vachell material for this species. In 1893, S. F. Harmer, then director of the Museum in Cambridge, listed all fish specimens, but C. maculatus does not appear on the list.? It may have been one of the specimens destroyed in 1866, during the overhaul of the collections. An account of the Vachell collection is given by Whitehead (in press) ..
Giinther (loc. cit.) listed three specimens of C. maculatus collected by Swinhoe from Formosa; these are Clupanodon thrissa. Richardson believed C. maculatus to be close to C. chrysopterus, both sharing the same Chinese vernacular name, but the Reeves figure (Pl. 5, fig. I) shows upper and lower jaws typical of Clupanodon.
Figure. According to the Reeves notebook, this figure was painted at Canton in November 1828. Reeves notes 6 or 7 black spots on the flanks, and these are well shown in Reeves No. 109 (PI. 5, fig. 1). But for the elongated last dorsal ray, this figure resembles Hilsa kelee (Cuvier). Apart from the spots it is otherwise similar to the figure of C. triza (Reeves No. 224, see Pl. 4, fig. 2).
Notes. The name maculatus was first used by Gray in a manuscript list of fishes in the British Museum. As in the case of other manuscript names (by Forster and one by Broussonet), Richardson accredited the name to Gray even though the list had not been published. As stated earlier, this list never was published and now appears to be lost.
13. “ Engraulis commersonianus Lacepéde ”’ = Stolephorus commersonii Lacepéde
Stolephorus commersonii Lacepeéde, 1803, Hist. Nat. Poiss. 5 : 381, 382, pl. 12, fig. 1.
Engraulis commersonianus ;: Richardson, 1846, Ichth. China Japan: 308 (Reeves specimens e% “ China ’’) ; Fowler, 1930, Pyvoc. Acad. nat. Sci. Philad.: 600 (Hong Kong, as commersonit).
Engraulis japonica : Giinther (part.), (non E. japonica Schlegel) 1868, Cat. Fish. Brit. Mus. 7: 390 (Reeves specimens ev “‘ China’’).
Engraulis chinensis Giinther, 1880, Rep. Voy. Challenger, 1: 73 (Reeves specimens and 4 other Chinese specimens—see below).
SPECIMENS. Four fishes, 79-82 mm. standard length, ex China, presented by J. R. Reeves, hitherto labelled “‘ Engraulis japonica” and unregistered, now BMNH.
1 Harmer’s Catalogues (two manuscript volumes) are now in the University Museum of Zoology in Cambridge.
38 P. J. P. WHITEHEAD
1964.11.6.8.11. These are the specimens listed as c-f in Giinther’s Catalogue (1868, p. 390). All are in good condition. For reasons given below, these fishes should not be regarded as syntypes of Engvaulis chinensis Giinther, 1880.
FicureE. There is no Reeves figure of this species.
Notes. Asa result of the poor description given by Houttuyn (1782) of a species, Atherina japonica (variously interpreted as one of two species of anchovy or a species of round herring—see Whitehead, 19630), considerable confusion existed in the nomenclature of the Chinese and Japanese anchovies. However, Richardson correctly identified the British Museum specimens with Lacepeéde’s Stolephore commersonien. Richardson noted that Cuvier had ranged the latter species “‘ among the anchovies, whose bellies are not toothed”’ (i.e. the modern genus Engrvaulis). But he observed that the Reeves specimens “ show six teeth before the ventrals as fine hairs’’ (characteristic of the modern genus Stolephorus). Unfortunately, Richardson placed Atherina australis Shaw (a true member of Engraulis—Whitehead, 19646) in his synonymy of FE. commersonianus.
Giinther (1868, p. 390), overlooking Schlegel’s Engraulis japonica (a true Engraulis), placed the Reeves specimens and some further Chinese specimens in Engraulis japonica (Houttuyn). Later (Ginther, 1880), finding a difference in finray counts between his E£. japonica and Schlegel’s, he proposed the name Engraulis chinensis for the Reeves and other specimens. These specimens appear in Gunther’s catalogue (1868, p. 390) as:
a, b, c-f Adult and half grown China gt Adult. Amoy. Purchased of Mr. Stevens
Specimen a is registered BMNH. 1831.12.27.207. It is now an alizarin prepara- tion (standard length 60 mm.). Specimen b is a juvenile (46 mm. S.L.); it is Engraulis gaponicus Schlegel and was evidently misidentified. Specimens c—f are labelled “ J. R. Reeves” but it is not clear why Giinther did not record them as such. Specimens g—i are registered BMNH. 1860.7.20.103.6.
Since the Reeves specimens are not positively identified as such in Giinther’s catalogue, it seems best to regard the three Stevens specimens as the syntypes of Engraulis chinensis Giinther (four fishes registered but one missing).
Fowler (1931, p. 199 ; 1941, p. 695) included Engraulis commersonianus of Richard- son in his synonymy of Engraulis japonicus Schlegel, overlooking Richardson’s reference to abdominal scutes. However, EF. japonicus certainly occurs in the Philippines and is also found in Hong Kong waters (B.M. specimens). Closely related to S. commersonit is S. indicus (van Hasselt), reported by Seale (1914) and Herre & Myers (1931) from Hong Kong. The latter species can be distinguished from S. commersonii chiefly by its shorter manilla (to front edge of operculum, not to gill-opening). Fowler (1931, p. 201) lists only one other Chinese record for S. indicus, namely Engrvaulis encrasicholus of Gtinther (1874) from Chefoo. However, these Giinther specimens (collected by Swinhoe) are true Engraulis japonicus. Stolephorus indicus is well known in the Philippines, and is reported from Formosan and japanese waters (Hayashi & Tadokoro, 1962). I have examined five Hong Kong specimens of S. indicus deposited in the Zoologiske Museum in Copenhagen (Nos. 99-IoT).
|
RICHARDSON ’S ELOPOLD AND CLUPEOID FISHES 39
’
14. ‘‘ Coilia grayii Richardson ’ = Coilia mystus (Linnaeus)
[Mystus ensiformis Linnaeus, 1754, Chinensia Lagerstrémiana— dissertatio’’: 26, fig. 12— China, on Lagerstrém. |
[Clupea mystus Osbeck, 1757, Dagbok Ostind. Resa : 256—Canton area. |
Clupea mystus Linnaeus, 1758, Syst. Nat., ed. 10 : 319 (name from Osbeck ; description mostly after Lagerstr6m) ; Idem, 1759, Amoen. Acad. 4 (61) : 252, fig. 12 (repeat of 1754 description and figure.)
Coilia mystus : Jordan & Seale, 1926, Bull. Mus. Comp. Zool. 67 (11) : 359 (Hong Kong); Fowler, 1931, Hong Kong Nat. 2 (3) : 206 (China, compiled) ; Herre, 1934, Hong Kong Nat. Supple- ment, No. 3: 26 (2 Hong Kong fishes).
Coilia grayit Richardson, 1845, Ichth. Voy. Sulphur: 99, pl. 54, figs. 1-2 (China seas) ; Idem, 1846, Ichth. China Japan : 309 (on type and Reeves illustr.) ; Jordan & Seale, 1926, Bull. Mus. Comp. Zool. 67 (11) : 361 (4 Hong Kong specimens) ; Herre & Myers, 1931, Lingnan Sci. J. 10 (2-3) : 238 (4 Hong Kong specimens).
Type. A fish, 243 mm. standard length (about 104 inches in total length, caudal tip damaged), registered BMNH. 1855.9.19.1581, presumed ex China Seas, sent to British Museum from the Haslar Hospital Museum to which (fide Richardson 1844, p- 100) it had been presented by Captain Dawkins, R.N. Although this specimen was listed as type by Giinther (1868, p. 405), the jar containing it (with original label) has not been marked as containing a type. Instead a second jar has been labelled C. grayut Type. This jar contains two smaller specimens, also from the Haslar collection, registered BMNH. 1855.9.19.1157. Richardson (1844) clearly states the length of the specimen from which the description was made (11 inches) ;_ his plate (pl. 54, fig. 1), stated to be life size, shows a fish of 250 mm. standard length. The two smaller Haslar specimens are barely 7} inches.
SPECIMENS. Gitinther (loc. cit.) lists five specimens under C. grayii. The first, indicated as ‘‘a’’, is the type; 0, is an adult (in alcohol) presented by J. R. Reeves ; cis another adult (stuffed) also presented by J. R. Reeves but subsequently destroyed; d and ¢ are the two small Haslar specimens. Curiously enough, Richardson does not mention these Reeves specimens, although the old label on the jar of the surviving (alcohol) specimen has the Hardwicke illustration number on it (H 252 R). This fish, 235 mm. standard length, is now registered BMNH. 1964.11.6.2.
FicurE. Reeves No. «14 (134 inches total length) shows a species with seven filamentous pectoral rays and a blunt maxilla reaching only to the pectoral base (Pl. 5, fig. 2). However, the maxilla has the appearance of having been broken off at its tip. Although the finrays of the anal and caudal are shown (correctly) as con- tiguous, the two fins are strongly demarcated by colour (anal grey/green, caudal orange/yellow). Scales and scutes are rather vaguely shown, and the drawing is far inferior to that given in the Voyage of the Sulphur (Richardson, 1844, pl. 54, fig. I). The surviving Reeves specimen, of 10} inches, is too small to have been model for the Reeves illustration.
Note. This is the second of the two clupeoid species listed by Osbeck (1757),
and reference is made to this early description in the synonymy since it was based on a Canton record and was used by Linnaeus (1758) in describing the species. Linnaeus
40 P. J. P. WHITEHEAD
(1754) had originally named this fish Mystus ensiformis, giving a figure (fig. 12) which was later reproduced in the Amoenitates Academiae (1759). Asin the case of Mystus altus, Linnaeus evidently decided to give priority to the name used by his pupil Osbeck during the latter’s voyage in 1770, although that name did not appear in print until 1757.
The genus Cozlia is badly in need of revision. It is not known, for example, to what extent small variations in numbers of pectoral filaments or gillrakers truly indicate specific differences. Authors have been divided on whether C. grayi1 is a distinct species or whether it is conspecific with C. mystus. Fowler (1931) distin- guished the two on gillraker counts (C. mystus 22-25 ; C. grayit 28-30) and anal rays (7o-86 and 86-92 respectively), but later (Fowler, 1941) he increased the range of anal rays in C. mystus to include C. gray. Lonnberg (1896) identified a specimen at Uppsala (labelled Clupea encrasicolus Mus. Lin.) as the type of C. mystus, claiming that it was really one from the Lagerstrém collection which had been mislabelled. Lonnberg gave no gillraker count (if such a count is indeed possible), but on pectoral filament numbers placed C. grayii in the synonymy of C. mystus (7 free filaments) ; he distinguished C. clupeoides Lacepéde (with 6 free filaments) as a separate species. It can be noted, however, that Lacepéde (1803, pp. 466, 467) does not refer to fila- mentous pectoral rays and based his description on Clupea mystus of Linnaeus and Osbeck. The single specimen of C. mystus of Jordan & Seale (1926, p. 359) had 6 free pectoral rays and 24 gillrakers. Giinther (1868) and Fowler (1941) list C. clup- eoides as a Synonym of C. mystus. The status of those species with only 6 pectoral filaments is discussed under the next species.
REDESCRIPTION OF TYPE AND REEVES SPECIMEN
TypE: BMNH. 1855.9.19.1581r ; Reeves fish: BMNH. 1964.11.6.2. Standard length: 243 mm. (TYPE) ; 238 mm. (Reeves) Total length : 275 mm. (TYPE) ; 262 mm. (Reeves)
mm. WASHES Body depth ; : : 3 4552 ALS 18°5 17°3 Head length : : ; 2 .A4r2 40°2 18-1 16-9 Snout length : : ; : 9:6 Q:I 3°7 3°8 Eye diam. : ; : : 8-9 8-6 3:6 3:6 Upper jaw 1. : : : . 54°3 51°1 21-9 21°5 Lower jaw l. : : F ; 30°8 29°76 12°7 I2°4 Pectoral 1. ‘ F : ‘ 79°5 88-2 32°7 37:0 Pelvicl. . : : . : I9*4 15°9 S-o 6°7 Pre-dorsal ‘ : : ; 73°7 65-4 30:2 27°4 Pre-pelvic . : : : ‘ 69-2 66-0 28-5 27°97 Pre-anal . : : : 1, LOS SO" We LOOs2 43°90 42:0
Body compressed, depth about equal to head length, posterior portion elongated, caudal peduncle about ? eye at caudal base. Manilla pointed posteriorly, reaching well beyond pectoral base, with fine conical teeth in a single series along entire lower edge becoming larger posteriorly ; two supramaxillae. Fine teeth present on pre- maxillae, vomer, pterygoids, palatines and on dentaries, the latter with well-developed
RICHARDSON’S ELOPOID AND CLUPEOID FISHES 4I
coronoid process. Pseudobranch exposed, almost equal in length to eye diameter, filaments about twenty, short.
Dorsal (preceded by small spine) with iii Io rays. Pectoral with vii Io rays, the first seven filamentous and unbranched, reaching to about base of 1o—14th branched anal ray, well beyond tip of depressed dorsal; branched rays of pectoral reaching beyond pelvic base. Pelvic 1 6, its base below anterior dorsal rays and nearer to pectoral base than to anal origin. Anal ii 88 and iii 86, final rays joined to lower rays of caudal.
Scales caducous, no count possible. Abdominal scutes trenchant, 12 and 15 pre-pelvic, 24 and 24 post-pelvic, the latter with slender ascending arms, alternately long and short.
Gillrakers moderate, about eye diameter, strongly armed with serrae along inner edge ; 22 and 22 rakers on upper arm of Ist arch, 30 and 27 on lower arm.
Branchiostegal rays Ir.
15. “ Coilia playfairii McClelland ”’ = Coilia playfairii (McClelland) Choetomus playfaivii McClelland, 1844, Calcutta J. nat. Hist. 4: 405, pl. 24, fig. 3 (China on
Playfair specimen(s)).
Coilia playfaivi : Richardson, 1845, Voy. Sulphur Ichth.: 100, pl. 54, figs. 3-4 (Hong Kong,
China seas) ; Idem, 1846, Ichth. China Japan : 309 (Japanese specimen).
Coiha grayt Kner, 1865, Reise Novara, Fische : 335 (Hong Kong). Coilia clupeoides : Giinther (part.), 1868, Cat. Fish. Brit. Mus. 7: 404 (Richardson specimen e+
China).
Cotlia Bee Giinther, 1868, Cat. Fish. Brit. Mus. 7: 405 (Japanese, Chinese specimens) ; Nichols,
1943, Nat. Hist. Central Asia, 9: 19 (Anhwei, nr. Canton).
? Coilia ectenes Fowler, 1930, Proc. Acad. nat. Sci. Philad.: 601 (Hong Kong); Idem, 1931,
Hong Kong Nat. 2 (3) : 208 (Kong Hong specimens).
SPECIMENS. No Reeves specimens listed by Richardson and none in British Museum collections. There is, however, a Japanese specimen (173 mm. standard length, labelled ‘“‘ Adara Japan ’’) which Richardson states was “‘ labelled ‘ Adara ’ by the authors of the ‘Fauna Japonica’’’. This fish was identified by Giinther (1868, p. 406, specimen “a’’) as C. nasus. There is also a Chinese specimen (150 mm., BMNH. 1847.5.10.5) presented by Richardson which Giinther (loc. cit., p. 404) included under Cozlia clupeoides Lacepede. The jar was later marked “ Cozlia playfair TyPE”’. In fact this specimen may well be that on which the figure of C. playfairi in the “ Voyage of the Sulphur ”’ was based (pl. 54, fig. 3, stated to be natural size). However it is not a type. Finally, there is a Vachell fish of 268.5 mm. S.L. at Cambridge (Whitehead, in press).
FIGURE. Reeves No. £26 shows a smaller fish than the figure for C. grayi, with a steeply rising dorsal profile (Pl. 5, fig. 3). The drawing is poor compared to that given in the “‘ Voyage of the Sulphur’. The number of pectoral filaments shown is 6.
Note. Several nominal species of Cozlia are stated to have 6 (or 5-6) free filamen- tous pectoral rays. Excluding those with pearly spots along the flanks (light organs, see Haneda, 1961), or with few post-pelvic scutes (g-11), or a short maxilla not
42 P. J. P. WHITEHEAD
reaching the gill opening, there are three species known from Chinese or Japanese waters (C. playfairi1, C. nasus and C. ectenes). Jordan & Starks (1906) and Jordan & Herre (1906) distinguished C. ectenes Jordan & Seale from C. nasus Schlegel mainly because of its greater number of anal rays and abdominal scutes and its more elongate form. The Japanese specimen labelled “‘ Adara’”’ (see above) has a markedly elon- gate body compared with other Chinese or Japanese specimens examined, and it has 49 scutes (48-49 in C. ectenes : cf. 42-43 in C. nasus, according to Jordan & Starks, 1906), and go anal rays (96-113 in C. ectenes; cf. 80-82 in C. nasus). However, Boeseman (1947, p. 178) describes 46 scutes and 85-88 anal rays in the type material of C. nasus, while Richardson (1844) gives 42-47 scutes and 70-80 anal rays for C. playfairit.
Thus the meristic differences separating these three nominal species are slight, and further material may well show that only a single small species is present in Chinese and Japanese waters, viz. C. playfairii. Fowler (1941) placed all three in the synonymy of C. mystus, but the latter differs in pectoral count and, on the basis of the specimens in the British Museum, appears to be a larger species. As noted already, Jordan & Seale (1926) found 6 pectoral filaments and 24 gillrakers in the specimens they considered to be C. mystus ; they record 7 filaments and 30 gillrakers in their specimens of C. grayit.
’
16. “‘ Thryssa mystax Bl. Schn.’ = Thryssa mystax (Schneider)
Clupea mystax Schneider, 1801, Syst. I[chth. Bloch. : 426, pl. 83 (Malabar).
Thryssa mystax : Richardson, 1846, Ichth. China Japan : 309 (on Reeves specimens).
Engraulis mystax : Fowler, 1930, Proc. Acad. nat. Sci. Philad. : 600 (Hong Kong).
Setipinna mystax : Fowler, 1931, Hong Kong Nat. 2 (3) : 203 (Hong Kong specimens).
Engraulis hamiltonii : Giinther, 1868, Cat. Fish. Brit. Mus. 7: 395 (Reeves and other Chinese specimens).
SPECIMENS. A fish, 183 mm. standard length (in alcohol) ex China presented by J. R. Reeves, with a metal tag sewn to caudal peduncle “ R138”; jar labelled “ Clupea H 236 306 R 138’, hitherto unregistered, but now BMNH. 1964.11.6.12. There is also a mounted skin (left side), labelled “ Engraulis hamiltont: China J. R. Reeves, Esq.”’ and with the Hardwicke and Reeves illustration numbers pencilled on the wooden base ; hitherto unregistered, now BMNH. 1964.11.6.13. Both speci- mens are mentioned by Richardson.
FIGURE. Reeves No. 138 might well have been drawn from the dry specimen in this museum, being only slightly larger than the illustration. It shows a rather deep- bodied compressed engraulid with post-pelvic scutes, a maxilla reaching beyond the posterior margin of the operculum, a long anal fin and no filamentous pectoral rays (Pl. 6, fig. 1). The black venulose supra-scapular area seen in the specimens is clearly shown in the illustration’, and the position of the fins closely correspond with those in the dried specimen. Richardson notes “ an indistinct stripe along the middle of the anal’’, but this does not appear in the preserved material.
1 Referred to as ‘‘ hairy process behind gills *’ in the Chinese Chrestomathy under species No. 4 (Bridg- man 1841, p. 480).
RICHARDSON’S-ELOPOID AND CLUPEOID FISHES 43
Note. Fowler (1931) recorded both T. hamiltoni and T. mystax from Hong Kong. T. hamiltonii is distinguished from T. mystax by its shorter maxilla (to gill opening or just beyond; cf. to or beyond pectoral base in 7. mystax). It is suspicious, however, that the specimens of T. hamultonii in our collections are mostly large fishes, whereas those of 7. mystax are small. Amongst the smaller specimens labelled T. hamiltonii (100 mm. and below), the maxilla reaches almost to the pectoral base. In all other respect these two nominal species are similar and their meristic counts overlap. The genus is currently under revision by Dr. S. Dutt.
The name Thrissocles Jordan & Evermann, widely but wrongly used for this genus as a senior synonym, should be replaced by Thryssa Cuvier, 1829 (see Whitehead, 19650). Fowler (1931, p. 203) placed both 7. mystax and T. hamultoniai in Setipinna Swainson, although he had (correctly) characterized that genus as possessing a filamentous upper pectoral ray.
DESCRIPTION, OF REEVES SPECIMENS
Standard lengths: 183 mm. BMNH. 1964.11.6.12 (alcohol) 188 mm. BMNH. 1964.11.6.13 (skin) (Figures for the alcohol specimen are given first.)
mm. Oe Sil
eS (aN Body depth : : . : ; 46°4 51-2 253) 273 Head length : : : : : 40°I 41°7 22:0 22:3 Snout length t : : : : 518 72 3:2 3°8 Eye diam. : F : 2 : 8:6 8-6 As 426 Upper jaw length 3 : : : AT 5. SATO, 227/122 <3 Lower jaw length : ; : : 30°5 29°5 to°7 15"7 Pectoral 1. : : : . ‘ Zi 1776 — Pelvic. . ‘ , : : i 13°8 75 Anal base 1. : : : ‘ ; 54°I1 63:8 29°6 33:8 Pre-dorsal ‘ : . : : 95°4 102°9 52°2 54°5 Pre-pectoral : : : : : A330) —— 24°70 — Pre-pelvic . : : : : a GAS 40-5 — Pre-anal_. : : F ; ‘ II5°9 122°5 63°5 65°5
Body compressed, its width almost three times inits depth. Maxilla long, reaching beyond posterior border of operculum to a point half way between operculum border and base of first pectoral ray. Pectoral fins just reaching base of pelvics, the tips of the latter just before dorsal origin. The following counts apply to the alcohol specimen only.
Dorsal with iii 11 rays, preceded by a minute spine, its origin equidistant between caudal base and anterior half of eye. Pectoraliito; pelvici6. Anal with iii 37 rays, its origin slightly behind last dorsal ray.
Abdominal scutes keeled, with sharp spines, 18 pre-pelvic (first minute), II post- pelvic (Richardson, 13 + 9).
Gillrakers 9 + 14 on first arch, each raker bearing serrae of approximately even length, not ranged into clumps.
Scale counts not possible (Richardson, 38).
44 P. J. P. WHITEHEAD
Venulose supra-scapular area with the venules dotted with small, linearly arranged melanophores. Flanks silver, except for upper ¢ which is brown. All fins hyaline.
d
17. ‘‘ Megalops setipinnis Forster ’ = Megalops cyprinoides (Broussonet)
Clupea cyprinoides Broussonet, 1782, Tableau Ichth.:no pagination, pl. 9 (Oceans between the tropics).
Megalops setipinnis Richardson, 1843, Ann. Mag. nat. Hist. 11: 493 (Port Essington) ; Idem, 1846, Ichth. China Japan : 310 (Seas of China).
Megalops curtifilis Richardson, 1846, Ichth. China Japan : 310 (on Reeves illustrations, Chinese Seas).
Type. Askin (right side), 180 mm. standard length, labelled “ Fresh water swamp near Victoria, Port Essington ’’ and on the reverse side “‘ Sept. 20.1840.3.”. A second label, pasted on the inside of the specimen, reads “‘ 3. Megalops setipinna Forster.’ This is the smaller of two skins, the other of which is registered BMNH. 1853.1.4.20; both are listed by Gtinther (1868, p. 472, d-e).
Richardson (1842 and subsequent papers on Australian fishes) described some dried skins numbered I-37, presented to this museum by J. Gould and collected in the Port Essington area by Gould’s assistant, Gilbert. In the original description of M. setipinnis, Richardson (1843, p. 493) describes a single Gilbert specimen of 9 inches 2 lines total length, ‘‘ No. 3, Mr. Gilbert’s list’”’. The smaller of the two British Museum skins, although now with damaged caudal tips, corresponds with the measurements given by Richardson. This fish is certainly the holotype of M. setipinnis Richardson, 1843, and it is now registered BMNH. 1964.11.6.14.
SPECIMENS. Richardson states that he had seen no Chinese (or Indian) specimens, and there are no Reeves specimens in the British Museum. There are two alcohol specimens from Port Essington (BMNH. 1843.8.10.11 and 1855.9.19.1142-3). The first of these was presented by Gould and was no doubt one of those referred to
by Richardson in the “ Report ”’.
FIGURE. Reeves No. 96 is a fair illustration (PI. 6, fig. 2), showing well the anasto- mising canals on the lateral line pore scales. There is, however, a single canine shown in the upper jaw ; Richardson remarks on this anomaly.
Notes. Richardson based his name for this species on a pencilled title “ Clupea setipinna’’ written underneath the uncoloured and only partly finished drawing by J. G. Forster (No. 242 in Forsters drawings from Cooks 2nd voyage, 1772-75, the 2nd of two volumes in the Zoological Library of the British Museum (Natural History)). Since Forster’s drawings were unpublished, Richardson was wrong to place Clupea cyprinoides Broussonet, 1782 as a junior synonym of this species.
Amongst the drawings, figures, etc. belonging to Richardson and left to this museum by his son, there is a tracing of Forster’s “ Kundinga ’’ with a pencilled note underneath “ not Gilbert’s fish’. In his description of JZ. setzpimnis Richardson notes that the Reeves drawing corresponds “ exactly in profile and size of fins, shape of head, etc.’ with the figures of both Forster and Broussonet, but that Forster's colours are different.
RICHARDSON Ssh OPOLD AND CLUPEOID FISHES 45
,
18. “ Megalops curtifilis Richardson ’ = Megalops cyprinoides (Broussonet)
(See previous species for synonymy.) SPECIMEN. No specimens mentioned by Richardson, and none in British Museum.
FiGuRE. Reeves No. 136 shows a smaller fish than in No. 96, but clearly referable to Megalops cyprinotdes (Pl. 6, fig. 3). The dorsal filament is a little shorter and the upper jaw a little longer than in No. 96, but the pored lateral line scales with their radiating canals are well shown.
Note. This is evidently MW. cyprinoides, the differences found by Richardson being attributable to poor drawing (scales fewer) and the fact that it was most likely a juvenile (dorsal filament shorter, body more slender).
1g. “ Elops machnata Forskal”’ = Elops machnata (Forsskal) Argentina machnata Forsskal, 1775, Descriptiones Animal. : 13, 68 (Red Sea). Elops machnata : Richardson, 1846, [chth. China Japan : 311 (Canton, Seas of China). Elops purpurescens Richardson, 1846, Ichth. China Japan : 311 (Chinese Seas). Elops saurus : Giinther, (part), 1868, Cat. Fish. Brit. Mus. 7: 470 (Reeves specimen, Indo-Pacific material).
SPECIMEN. Richardson states that Reeves deposited a specimen from Canton in this museum. Amongst the stuffed specimens is one, of 390 mm. standard length, originally labelled “ Elops saurus’’ but with the name “ machnata ’’ added in another hand. Unlike the other stuffed Reeves specimens, no details are painted on the wooden base, but a pencilled note underneath reads “ Elops machnata Canton J. R. Reeves Esq.’”’. The specimen was hitherto unregistered but is now BMNH. 1964. TC. OLS
FIGURE. Reeves No. 137 is a fair drawing and easily recognizable as Flops by its small scales, elongate body, pored lateral line scales, etc. (Pl. 7, fig. I).
Notes. The specimen has approximately 87 pored lateral line scales, which accords with FE. machnata, and the lower jaw (now set open) probably covered the pre-maxillary tooth band when the jaw was closed (lower jaw included in EF. hawaien- sis Regan). On the basis of the most recent key (Whitehead, 19620) the specimen is evidently E. machnata. Richardson rightly distinguished his fish from the Mugil salmoneus (Forster) Schneider figured in his Ichthyology of the Erebus and Terror (Richardson, 1896, pl. 36, figs. 1, 2) ; that fish was Chanos chanos (Forsskal), a species apparently not encountered by Reeves.
20. “ Elops purpurescens Richardson ”’ = Elops machnata (Forsskal) (See previous species for synonymy.) Type. Richardson based this name solely on the Reeves illustration. The name is now a nomen oblitum but it is very unlikely that a distinct Chinese or Western Pacific species or subspecies of Elops will ever be recognized.
40 P. J.P. WHITEHEAD
FIGURE. Reeves No. 53 appears to have been drawn from a specimen long out of water (Pl. 7, Fig. 2). The fins have darkened, the flanks are paler and the back is darker than in the figure of E. machnata. In addition, the body is twisted, giving a more convex lower profile than in the preceding species.
Notre. Richardson distinguished this fish from E. machnata, but added “ This drawing does not differ very greatly from the preceding one in form’’. He notes its more irregular and less arched dorsal outline, more convex belly and slightly decurved lateral line. He also notes a slight difference in the Chinese names given by Reeves : Chuh Keaou, “ Bamboo——’’ for E. machnata ; Chuh Kin, “ variegated Bamboo ”’ for E. purpurescens.
The figure suggests merely a twisted specimen of FE. machnata. Only a single
species of Elops is recognized from the Western Pacific (see Whitehead, 19625).
21. “ Chirocentrus dorab Forskal ”’ = Chirocentrus dorab (Forsskal)
Clupea dorab Forsskal, 1775, Descriptiones Animal. : 72 (Red Sea).
Chirocentvus dovab: Richardson, 1846, Ichth .China Japan: (Canton, seas of China); Giinther, 1868, Cat. Fish. Brit. Mus. 7: 475 (Reeves specimen) ; Fowler, 1930, Proc. Acad. nat. Sct. Philad. : 598 (Hong Kong) ; Idem, 1931, Hong Kong Nat. 2 (1) : 75 (Hong Kong specimens) ; Herre, 1931, Hong Kong Nat. Supplement, No 3 : 26 (2 Hong Kong specimens).
SPECIMEN. Richardson mentions an alcohol specimen from Canton presented by Reeves of 10} inches. This specimen (220 mm. S.L.; hitherto unregistered, but now BMNH. 1964.11.6.1) is labelled “‘ Chirocentrus dorab China J. R. Reeves Esq. H 237 R”’. It lacks scales and the fins are damaged slightly, but otherwise the specimen is in fair condition.
Figure. As Richardson noted, the Reeves figure (Reeves No. 47) hints at pun- gent ventral scutes (PI. 7, fig. 3), but these spines are in fact the tips of the ribs, a common artifact in preserved specimens of Chirocentrus. The figure shows no scales, but it is not possible to judge whether the present specimen (104 inches) served as model for the illustration (15 inches).
Notes. This specimen has 5 + 16 gillrakers on the first arch, a count which places it in C. hypselosoma Bleeker according to Hardenberg (1930) (modal count 14-15 on lower part of first arch for C. dorab). On the other hand, the body depth is contained 6 times in standard length in the Reeves specimen and the maxilla does not reach the front border of the preoperculum, which accords with C. dorvab (54 and beyond respectively in C. hypselosoma). The systematic position of Bleeker’s C. iypselosoma has been examined elsewhere (Whitehead, Boeseman & Wheeler, in press) and the conclusion reached that there may indeed be two species of Chivocentrus present in the Indo-Pacific. However, for the present, the Reeves fish is identified with C. dorab until the two species can be more trenchantly defined.
LIST OF HONG KONG ELOPOID AND CLUDEOID SPECIES For a list of the elopoid and clupeoid species recorded from the Hong Kong area (see Table 2) I have relied chiefly on Fowler (1930, 1931), Herre & Myers (1931)
RICHARDSON’S ELOPOID:AND CLUPEOID FISHES 47
and Herre (1934). There have been few subsequent records. In addition to published records, the list of species given has been augmented by inclusion of specimens represented in the British Museum collections, and especially by Hong Kong speci- mens generously donated to the museum by Mr. W. Chan.
Hong Kong lies just within the tropics, and many of the species found there are common both to the Philippines and to the sea around Taiwan (Formosa). Where species have been reported from near Taiwan (Chen, 1961, and a useful review of clupeoids by Chu & Tsai, 1958), or from Korea (Mori, 1952), or from Japan (Matsu- bara, 1955), and at the same time are also known from the Philippines (Fowler, 1941 ; Herre, 1953), then I have assumed their probable occurrence in Hong Kong waters. Such an assumption is usually justified in clupeoid fishes, the marine species, at least in the Indo-Pacific region, being for the most part wide-ranging.
Forty-four species are listed here, more than twice as many as were known to Richardson. However, in eleven cases there is no actual Hong Kong record or speci- men, and the list for certain genera can only be tentative. This is particularly true for the genera Herklotsichthys and Sardinella. The species of Stolephorus of this area are also poorly known, but the Chan collection contained three species and showed, perhaps surprisingly, that one of the commonest is S. buccaneert Strasburg, a species closely related to S. purpureus Fowler, both of which were believed confined to the Hawaiian Islands. The presence of S. buccaneert in the Hong Kong region may explain the reports of S. zollingeri (Bleeker) from Japanese waters (Hayashi & Tadokoro, 1962). Thus the types of S. zollingert are not members of Stolephorus at all, but are Engraulis japonicus Schlegel (Whitehead, 1964), a species unlikely to be misidentified by Japanese workers. But published descriptions of S. zollingeri in Japanese waters (e.g. Hayashi & Tadokoro, loc. cit.) strongly suggest S. buccaneert, although none have mentioned the characteristic diamond-shaped urohyal plate (see Whitehead, 1965), fig. 4a). Specimens of S. buccaneert have also been recorded from the Red Sea region and from Durban (Whitehead, 19656) but not from inter- vening areas.
The list of species of Thryssa, Coilia and Ilisha given here must also be considered tentative, all three genera badly needing revision.
ACKNOWLEDGEMENTS
It is a pleasure to record the assistance given me by Mr. P. H. Hulton, Assistant Keeper, Department of Prints and Drawings, British Museum, during the search for the fourth Reeves set of illustrations ; similarly, I must acknowledge the help of Dr. K. Gardiner, Keeper of the Department of Oriental Printed Books and Manu- scripts, British Museum ; and Mrs. M. Archer, India Office Library, London.
My thanks are also due to Mr. W. Chan, Fisheries Research Station, Hong Kong, for so generously donating an excellent collection of Hong Kong clupeoids and for notes on the Hong Kong species; also to Dr. K. Joysey, University Museum of Zoology, Cambridge, for help with the Vachell collection.
The manuscript was read by Dr. P. H. Greenwood and I gratefully acknowledge his useful comments and criticisms.
ZOOL, I4, 2 5A
48 P. J. P. WHITEHEAD
TABLE 2
Elopoid and clupeoid species recorded from the vicinity of Hong Kong.
Species ELOPIDAE Elops machnata (Forssk.) MEGALOPIDAE
Megalops cyprinoides (Brouss.) ALBULIDAE tAlbula vulpes (Linn.)
CHIROCENTRIDAE Chirocentrus dovab (Forssk.)
DUSSUMIERIIDAE Etvumeus teres (DeKay)
Dussumieria acuta Valenc. Spratelloides gracilis (Schlegel) CLUPEIDAE (Clupeinae) Sardinella aurita Valenc.
tSardinella brachysoma Blkr.
tSardinella bulan (Bleeker) Sardinella jussieu (Lac.)
Sardinella fimbriata (Valenc.)
Sardinella leiogastey Valenc.
tSardinella clupeoides (Blkr.)
}Herklotsichthys schrammi (Bleeker) }Herklotsichthys punctatus (Riippell) tHerklotsichthys zunasi (Bleeker)
(Alosinae) Hilsa (Tenualosa) veevesti (Rich.)
Hilsa (Tenualosa) toli (Valenc.)
(Pristigasterinae) {Pellona ditchela Valenc,
Reference
Richardson, 1846 (Canton*). Richardson, 1846 (on Reeves Illustr.)
[Fowler, 1941—Japan, East Indies; Liu & Shen, 1957—Taiwan.|
Richardson, 1846 (Canton*); Fowler, 1930, 1931, (Hong Kong).
Whitehead, 19636, p. 374 (Hong Kong) ; Hong Kong*
Herre & Myers, 1931 (Hong Kong); Hong Kong.*
[Whitehead, 1963a, p. 375—Formosa*; Fowler, 1941—Philippines] ; Hong Kong.*
Richardson, 1846 (on Reeves illustr. of C. nymphaea); [WKishinouye, 1907—Amoy, Swatow; Chan, 1965—Taiwan] ; China*.
[Chu & Tsai, 1958—Formosa, as S. albella (Val.).]
[Bleeker, 1873—Amoy]; Amoy*.
Fowler, 1931 (Hong Kong); Amoy*, Hong Kong*.
Richardson, 1846—as C. isingleena; Hong Kong*.
Kner, 1865 (Hong Kong); Richardson, 1846— as C. caeruleo-vitiata; ? Jouan, 1867—Hong Kong, as Harengula moluccensis.
[Kishinouye, iI907—Japan, as C. okinawensis ; Fowler, 1941—Philippines; Chan, 10965 Thailand, Philippines].
[Chu & Tsai, 1958—Formosa].
[Chu & Tsai, 1958—Formosa] ; Philippines*.
[Fowler, 1951—Japan, Philippines]; Japan*, East Indies*.
Richardson, 1846 ; Ixong*).
Fowler, 1931 (Hong Kong, as MWacruva sinensis); [Jordan & Evermann, t902—Formosa].
Whitehead, 1965a@ (Hong
[Liu & Shen, 1957—Taiwan, as Ilisha hoeveni].
* Specimen in British Museum from this locality. [ ] references from other areas. t Species may occur in Hong IKkong waters, but no record or specimen.
RICHARDSON’S ELOPOID AND CLUPEOID FISHES 49
Ilisha elongata (Bennett)
Ilisha brachysoma (Bleeker) Ilisha indica (Swainson)
Ilisha novacula (Valenc.)
Opisthopterus tardoore (Cuvier) Opisthopterus valenciennest Blkr.
(Dorosomatinae) Konosivus punctatus (Schlegel)
Clupanodon thrissa (Linn.)
Nematalosa nasus (Bloch)
Nematalosa japonica Regan
tAnodontostoma chacunda (Ham. Buch.)
ENGRAULIDAE Engraulis japonicus (Schlegel)
Stolephorus commersonii Lac.
Stolephorus indicus (van Hass.)
Stolephorus buccaneert Strasburg Stolephorus tri (Bleeker) Stolephorus hetevolobus (Riipp.)
Thryssa mystax (Schneider)
tThryssa setivostvis (Brouss.) Thryssa dussumiert (Valenc.)
tThryssa hamiltoni (Gray) Setipinna taty (Valenc.)
Cotlia mystus (Linn.)
Cotlia playfairit (McClelland)
Richardson, 1846 (China Seas*) ; Valenciennes, 1847 (Macao); Kner, 1865 (Hong Kong) ; Fowler, 1931 (Hong Kong); Chen, 1961 (Quemoy) ; Amoy*.
Fowler, 1931 (Hong Kong) ; Hong Kong*.
Norman, 1923 (Hong Kong*); (Chen, 1961— Quemoy].
Norman, 1923 (China*).
Fowler, 1931 (Hong Kong).
Foochow*, Hong Kong*.
Richardson, 1846 (Chinese sea*); Herre & Myers, 1931 (Hong Kong) ; Amoy*.
Osbeck, 1757 (Canton); Richardson, 1846 (on Reeves illustr. and specimen from Canton*); Fowler, 1930, 1931 (Hong Kong); Herre, 1934 (Hong Kong); [Liu & Shen, 1957— Taiwan].
? Richardson, 1846 (on Reeves illustr.) ; Jordan & Seale, 1905 (Hong Kong); Herre, 1934 (Hong Kong) ; Hong Kong*.
Herre & Myers, 1931 (Hong Kong); Hong Kong*.
[Fowler, 1941—Hainan, Philippines}.
Chefoo*, Hong Kong* [Whitehead, 1964— Japan*, East Indies].
Richardson, 1846 (China*); Fowler, 1930 (Hong Kong) ; Hong Kong*.
Seale, 1914 (Hong Kong); Herre & Myers, 1931 (Hong Kong); [Chen, 1961—Quemoy] ; Hong Kong specimens, Zool. Mus. Copen- hagen.
Formosa*; [? Chen, 1961, as S. zollingeri— Quemoy]; Hong Kong*.
(Bleeker, 1865—Amoy; |Liu & Shen, 1957— Taiwan]; Hong Kong*.
Hong Kong*; [Liu & Shen, 1957—Taiwan, as S. pseudoheterolobus}.
Osbeck, 1757 (Canton); Richardson 1844-5, 1846 (China Seas); Jordan & Seale, 1926 (Hong Kong); Herre & Myers, 1931 (Hong Kong); Herre 1934 (Hong Kong).
[Fowler, 1941—Philippines] ; Amoy*.
Seale, 1914 (Hong Kong); ([Chen, 1961— Quemoy].
(Mori, 1952—Japan; Fowler, 1931—Philip- pines] ; China*.
Fowler, 1931 (Hong Kong).
Osbeck, 1757 (Canton); Richardson, 1844-5, 1846 (China Seas*); Jordan & Seale, 1926 (Hong Kong).
Richardson, 1845 (Hong Kong); Kner, 1865 (Hong Kong); Fowler, 1930, 1931 (Hong Kong) ; Nichols, 1943 (Anhwei, nr. Canton) ; China*,
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1962b. The species of Flops. Ann. Mag. nat. Hist. (13) 5 : 321-329.
1963a. A revision of the recent round herrings (Pisces: Dussumieriidae). Bull. Brit.
Mus. nat. Hist. (Zool.) 10 (6) : 305-380.
19636. Atherina japonica Houttuyn, 1782: proposed suppression as being a nomen
dubium, Bull. zool. Nomencl. 20 (4) : 281-284.
52 P.jJ.P. WHITEHEAD
WHITEHEAD, P. J. P. 1964a. Herklotsichthys Whitley, 1951 to replace Havengula Valenciennes, 1847 for Indo-Pacific species (Pisces : Clupeidae). Ann. Mag. nat. Hist. (13) 6: 273-284. 1964b. New data extending the range of the bipolar antitropical anchovy genus Engraulis. (in Russian). Zool. Zh. 43 (6) : 879-888.
1964c. A redescription of the holotype of Clupalosa bulan Bleeker, and notes on the genera Herklotsichthys, Sardinella and Escualosa (Pisces: Clupeidae). Ann. Mag. nat. Hist. (13) : 273-284.
1965a. A preliminary revision of the Indo-Pacific Alosinae (Pisces: Clupeidae). Bull. Brit. Mus. nat. Hist. (Zool) 12 (4) : 117-156. ——— 1965). A review of the elopoid and clupeoid fishes of the Red Sea region. Jbid. 12 (7) : 225-281.
(in press). The Vachell collection of Chinese fishes at Cambridge. WHITEHEAD, P. J. P., Bo—eseman, M. & WHEELER, A. C. (in press). The types of Bleeker’s Indo-Pacific elopoid and clupeoid fishes. Zool. Verhandel. Leiden.
INDEX TO. SPECIES
abnoymis, Ilisha, 32, Pl. 3 gracilis, 29 acuta, Dussumieria, 48 isingleena, 24, Pl. I Alausa reevesii, 30 melanosticta, 27 Albula vulpes, 48 mystax, 42 Alosa elongata, 32 mystus, 39 palasah, 31, Pl. 3 nasus, 30 yeevesit, 30, Pl. 2 nymphaea, 26, Pl. 1 altus, Mystus, 34 okinawensis, 48 Anodontostoma chacunda, 49 thrissa, 34 aquosus, Chatoessus, 33, Pl. 4 tviza, 35 Argentina, machnata, 45 Clupea (Amblygaster) melanosticta, 28 aurita, Sardinella, 26, 48 clupeotdes, Cotlia, 41 Sardinella, 48 brachysoma, Ilisha, 32, 49 Cotlia clupeoides, 41 Sardinella, 48 ectenes, 41 buccaneert, Stolephorus, 47, 49 grvayit, 39, Pl. 5 bulan, Sardinella, 48 mystus, 39, 49 nasus, 41 caeruleo-vittata, Clupea, 28, Pl. 2 playfaivit, 41, 49, Pl. 5 chacunda, Anodontostoma, 49 commersonianus, Engraulis, 37 Chatoessus aquosus, 33, Pl. 4 commersonit, Engraulis, 37 chrysopterus, 36, Pl. 4 Stolephorus, 37, 49 maculatus, 35, 37, Pl. 5 curtifilis, Megalops, 44, 45, Pl. 6 osbeckit, 35 cvprinoides, Clupea, 44 punctatus, 33 Megalops, 44, 45, 48 tyiza, 34, Pl. 4 chinensis, Engraulis, 37 delicatulus, Spratelloides, 30 Chirocentrvus dovab, 46, 48, Pl. 7 ditchela, Pellona, 48 hypselosoma, 46 dorab, Chirocentrus, 46, 48, Pl. 7 Choetomus playfaivit, 41 Clupea, 46 chrysopterus, Chatoessus, 36, Pl. 4 dussumiert, Thryssa, 49 Clupanodon thrissa, 34, 49 Dussumieria acuta, 48 Clupea caeruleo-vittata, 28, Pl. 2 cyprinoides, 44 ectenes, Coilia, 41 dovab, 46 elongata, Alosa, 32
flosmaris, 29, Pl. 2 Ilisha, 32, 49
INDEX
Elops machnata, 45, 48, Pl. 7 machnata, Argentina, 45 purpurescens, 45, Pl. 7 Elops, 45, 48, Pl. 7 saurus, 45 Macrura sinensis, 48
Engraulis chinensis, 37 maculatus, Chatoessus, 35, 37, Pl. 5 commersonianus, 37 Megalops, curtifilis, 44, 45, Pl. 6 cOMMEYSONI1, 37 cyprinoides, 44, 45, 48 hamiltonit, 42 setipinnis, 44, Pl. 6 japonica, 37, 49 melanosticta, Clupea, 27 mystax, 42 Clupea (Amblygaster), 28
ensiformis, Mystus, 39 moluccensis, Harengula, 28
Etrumeus teres, 48 mystax, Clupea, 42
Engraulis, 42
jfimbriata, Sarvdinella, 24, 48 Setipinna, 42 Spratella, 24 Thryssa, 42, 49, Pl. 6
flos-maris, Clupea, 29, Pl. 2 Mystus altus, 34
enstformis, 39
gracilis, Clupea, 29 mystus, Clupea, 39 Spratelloides, 29, 48 Cotlia, 39, 49
grayana, Pellona, 32
grayu, Coilia, 39, Pl. 5 nasus, Clupea, 36
Coilia, 41
hamiltonu, Engraulis, 42 Nematalosa, 36, 49 Thryssa, 42, 49 Nealosa punctata, 33
Harengula moluccensis, 28 Nematalosa japonica, 36, 49 nymphaea, 24 nasus, 30, 49
Herklotsichthys punctatus, 48 novacula, Ilisha, 32, 49 schrammi, 48 nymphaea, Clupea, 26 zunast, 48 Harengula, 24
heterolobus, Stolephorus, 49
Hilsa veevesii, 30, 31 okinawensis, Clupea, 48
Hilsa (Tenualosa) reevesii, 48 Opisthopterus tardoore, 49 toli, 48 valenciennest, 49
hoevemi, Ilisha, 48 osbeckii, Chatoessus, 35
hypselosoma, Chirvocentrus, 46 palasah, Alosa, 31, Pl. 3
Ilisha abnorymis, 32, Pl. 3 Pellona ditchella, 48 brachysoma, 32, 49 grvayana, 32 elongata, 32, 49 vimbella, 32 hoevem, 48 playfaivit, Choetomus, 41 indica, 49 Coilia, 41, 49, Pl. 5 novacula, 32, 49 punctata, Nealosa, 33 indica, Ilisha, 49 punctatus, Chatoessus, 33 indicus, Stolephorus, 49 Herklotsichthys, 48 isingleena, Clupea, 24, Pl. 1 Konosivus, 33, 49
purpurescens, Elops, 45, Pl. 7 japonica, Engraulis, 37, 49
Nematalosa, 36, 49 veevestt, Alausa, 30 jussieu, Sardinella, 48 Alosa, 30, Pl. 2 Hilsa, 30, 31 Konosivus punctatus, 33, 49 Hilsa (Tenualosa), 48
thrissa, 36
Sardinella aurita, 26, 48 leiogaster, Savdinella, 48 brachysoma, 48
54 INDEX
Sardinella (contd.) taydoove, Opisthopterus, 49 bulan, 48 taty, Setipinna, 49 clupeoides, 48 teres, Etrumeus, 48 fimbriata, 24, 48 thrissa, Clupanodon, 34, 49 jussieu, 48 Clupea, 34 leiogaster, 28, 48 Konosivus, 36 sym, 28 Thryssa dussumieri, 49
saurus, Elops, 45 hamiltoni, 42, 49
schvammi, Herklotsichthys, 48 mystax, 42, 49, Pl. 6
Setipinna mystax, 42 setivostris, 49 taty, 49 toli, Hilsa (Tenualosa), 48
setipinnis, Megalops, 44, Pl. 6 tvt, Stolephorus, 49
setivostvis, Thryssa, 49 tviza, Chatoessus, 34, Pl. 4
sinensis, Macrura, 48 Clupea, 35
sivm, Sarvdinella, 28
Spratella fimbriata, 24.
Spratelloides delicatulus, 30 gracilis, 29, 48
Stolephorus buccaneeri, 47, 49 commersonii, 37, 49 heterolobus, 49 indicus, 49 tvt, 49 zollingeri, Stolephorus, 47, 49 zollingert, 47, 49 zunast, Herklotsichthys, 48
valenciennest, Opisthopterus, 49 vimbella, Pellona, 32 vulpes, Albula, 48
Fic. 1.
Fic. 2. Fie. 3.
PLATE r
Clupea isingleena Richardson, holotype (108-5 mm. S.L., BMNH. 1963. 6.17.1). Formerly believed holotype of C. nymphaea, see text, p. 24. Clupea isingleena, Reeves drawing No. 60 [= Sardinella fimbriata (Val.)]. Clupea nymphaea, Reeves drawing No. [25 [= Sardinella aurita Val.).
I
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TA, 2
PLATE 2
Fic. 1. Clupea caeruleo-vittata, Reeves drawing No. 59 [= Sardinella, probably S. leiogastey Val.]. Fic. 2. Clupea flosmaris, Reeves drawing No. 64 [= Herklotsichthys or Sardinella
sp.]. Fic. 3. Alosa reevesii, Reeves drawing No. a8 [= Hilsa veevesii (Rich.)}.
Bull. B.M. (N.H.) Zool. 14, 2 PLATE 2
PLATE 3
Fic. 1. Alosa palasah, Reeves drawing No. (651 [= Hilsa reevesti (Rich.)]}. Fic. 2. Llisha abnormis, Reeves drawing No. 81 [= Ilisha elongata (Bennett)]. Fic. 3. Ilisha abnovmis, Reeves drawing No. 67 [ = Ilisha elongata (Bennett) |.
PbAT E
Bull. B.M. (N.H.) Zool. 14, 2
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PLATE 4 Fic. 1. Chatoessus aquosus, Reeves drawing No. 63 [= Konosivus punctatus (Schl.)]. Fic. 2. Chatoessus triza, Reeves drawing No. 224 [= Clupanodon thrissa (L.)|. —
Fic. 3. Chatoessus chrysopterus, Reeves drawing No. 61 [= Nematalosa, probably N. nasus (Bloch)}.
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(L.)]
Fic. 2. Coilia gravii, Reeves drawing No. at4 [= Coilia mystus (L.)]}.
Fic. 3. Coilia playfairii, Reeves drawing No. 626 [= Coilia plavfairia (Mc- Clelland)]. eS
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Fic. 1. Elops machnata, Reeves drawing No. 137 [= Elops machnata (Forsskal)].
Fic. 2. Elops purpurescens, Reeves drawing No. 53 [= Elops machnata (For- sskal)].
Fic. 3. Chivocentrus dovab, Reeves drawing No. 47 [= Chirocentrus dorab (Forsskal) }.
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PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING
OBSERVATIONS ON THE TYPE- MATERIAL OF SOME GENERA AND SEE@IES OF -POLYZOA
ANNA B. HASTINGS
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY WO wi INO, 4 LONDON: 1966
ogee MUSE 7 TRING Vig
x
OBSERVATIONS ON THE TYPE-MATERIAL OF SOME GENERA AND SPECIES, OF POLYZOA
BY
ANNA B. HASTINGS
Pp. 55-78 ; 1 Plate ; 1 Text-figure
BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Voki No, 2 LONDON : 1966
THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted im 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical series.
Parts will appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year.
In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department.
This paper is Vol. 14, No. 3 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals.
RUSE E E'S: Os THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 11 March, 1966 Price Thirteen Shillings
© Trustees of the British Museum (Natural History) 1966
OBSERVATIONS: ON THE TYPE-MATERIAL @OEFSSOME GENERA AND SPECIES OF POLYZOA
By ANNA B. HASTINGS!
Page
1. INTRODUCTION ; ‘ 3 : c : : : : 57 2. CHALLENGER COLLECTION : : 5 ‘ ‘ . é ‘ 57 3. SOME TYPE-SPECIMENS . : : - - 58 Bifaxaria corrugata Busk ane 18 papillata Busk : : 2 . 58 Harmer’s elucidation of the Challenger Report : 2 . 58 Lectotypes . : 2 : : ; : : : : 59 Systematic position : : 5 ; : : ; : 59 Comparison of the species 4 é ; : . c : 59
Identity of the Siboga material é : : é : 61
Flustva octodon Busk . : : : ; 2 : 5 62 Lepralia melolontha Landsborough : : : : : =) 63 Lepralia multidentata Thornely . : : : : - 64 Lunulites owen Gray . ‘ : : ; : : : - 64 Membranipora amplectens Hincks 2 E : : . 65 Membranipora crassimarginata var. evecta Busk : : ; ee O5 Membranipora hexagona Busk é é : : : = “160 Membranipora pilosa var. multispinata Finck ‘ : : 2, 107 Membraniporella agassizit Smitt . é : : : : . 68 Menipea fuegensis Busk : : : : : : é = 68 Micropora normani Levinsen : , : : : : . 69 Salicornaria clavata Busk . : : ; ‘ : : ; 7. Siphonoporella nodosa Hincks F : m2
4. THE GENUS HIPPOPLEURIFERA AND SOME OF ITS SPECIES . : 73 Hippopleurifera Canu & Bassler. é i A : é eS
Hi. biauriculata (Reuss) Q ; : : : : ; : 73
H. pulchva (Manzoni) . : F : : : : 5
H. sedgwicki (Milne- Edwards) : . . : : ; - 75
5. ACKNOWLEDGEMENTS : : : : Z : ‘ ? : 75 6. SUMMARY , : F ; 5 ‘ F : F : ; 75 7. JREFERENCES . ; : : : : : : : : : 76
i INTRODUCTION
In this paper I have gathered together the results of the examination of the type- specimens of a number of species.
2. CHALLENGER COLLECTION
The two main sets of Challenger Polyzoa (received by the Museum in 1887 and 1899, see Hastings, 1943 : 304) have been supplemented by further material from the Busk Collection, received in 1943 (registration 1944.1.8.1-406), among which
1 Address: 28, Kew Gardens Road, Richmond, Surrey, England. ZOOL. 14, 3 6
58 A. B. HASTINGS
some figured, and otherwise informative, specimens have been found. There is also a series of preparations of chitinous parts, made by Busk from Challenger specimens, received by the Museum in 1882. All these collections include syntypes of Busk’s Challenger species.
3. SOME TYPE-SPECIMENS
It has seemed best to arrange these notes, on the examination of sundry type- specimens, alphabetically according to the names under which the species were originally described. An indication of the present generic attribution of each species is given.
Bifaxaria corrugata Busk and B. papillata Busk (Text-fig. IA, B, C)
HARMER’S ELUCIDATION OF THE CHALLENGER REporT. Harmer (1957 : 868) eluci- dated Busk’s confusion between Bifaxaria corrugata and B. papillata, both of which (pp. 867, 868) he referred to Sclerodomus Levinsen. Harmer did not see Busk’s drawings (published or unpublished), nor his copy of the Challenger Report. They have yielded evidence, in Busk’s hand-writing, confirming Harmer’s conclusions. The details are filed in the British Museum (Nat. Hist.).
Harmer’s conclusions may be summarized thus: Busk’s pl. XIII, figs. 4, 4a, des- cribed as B. papillata, represent B. corrugata; and his pl. XXIV, fig. 6, described as B. corrugata, represents 6. papillata; the attributions of pl. XIII, figs. 3, 3a to B. corrugata and pl. XXIV, figs. 4 A-D to B. papillata are correct. B. corrugata was obtained solely at St. 122, and B. papillata solely at St. 106.
With only four exceptions (pl. XIII, fig. 4, one zoarium, and pl. XXIV, figs. 4B, 4C and 6, see below) all the figured specimens are definitely recognizable by their detailed agreement with the figures. Harmer’s identification of these specimens agrees with Busk’s original labelling. In the subsequent confusion Busk altered the name on one slide (1887.12.9.379A changed from B. corrugata to B. papillata, presumably because the specimen is shown in pl. XIII, fig. 4, left hand figure, under that name). Kirkpatrick (MS. Catalogue) accepted this, and added a label to the slide. These are the alterations mentioned by Harmer.
To sum up, the specimens are mounted on three slides, and are figured as follows: 1887 .12.9.377, B. corrugata, St. 122.
Left hand zoarium is represented in pl. XIII, fig. 3 (nat. size), and fig. 4a.
Right hand zoarium is represented in pl. XIII, fig. 4, right-hand figure (nat. size).* 1887.12.9.379A, B. corrugata, St. 122, is represented in pl. XIII, fig. 4, left-hand
figure (nat. size), and fig. 3a.
! He should have excluded one of the three zoaria in fig. 4 from B. corrugata (see below).
2 Tt will be noticed that the figures of the zoaria are mostly placed with magnified figures of the same species, but not always with those of the same specimen. The original drawings of the zoaria were (with one exception) not on the same cards as the magnified drawings, which perhaps led to the errors in
arranging them on the plate. None of the original drawings of zoaria have been preserved ; even the one on the same card as other drawings was cut out, and a note made beside the hole,
| | 1887.12.9.379, B. papillata, St. 196, is represented in pl. XXIV, figs. 4A and 4D. Figs. 4B, 4C and 6 (pl. XXIV) show no peculiarities by which the figured
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 59
zooecia can be recognized, but they closely agree with the zooecia of comparable age in this specimen.
The pieces mounted on this slide (1887.12.9.379) are evidently parts of one colony, which appears to have been the central zoarium of three figured in pl. XIII, fig. 4; but I have not proof of this. The presence of secondary branches does, however, support the conclusion that this drawing represents b. papillata | (see footnote I on p. 58).
LECTOTYPES :
a. B. corrugata. Harmer’s choice (1957 : 868) of the specimen 1887.12.9.379A / as ‘‘ type’’, in his list of specimens, and “ type-specimen ”’ in the text, constitutes a _ valid choice of lectotype.
b. B. papillata. J have said above that the fragments on the slide 1887.12.9.379 are evidently parts of one colony. If this could be proved the whole slide would be holotype by monotypy. In the absence of proof, I have chosen as lectotype the _ lower (larger) of the two pieces figured in pl. XXIV, fig. 4A.
| SYSTEMATIC PosiTION. The frontal wall of Sclerodomus corrugatus and S. papilla- tus is arched over a frontal membrane with a membraniporine operculum. The _ wall is thus of the nature of a frontal shield, and these species belong to the Ascophora _ imperfecta of Harmer (1957 : 645, 651).
_ [am not confident that Harmer was right in referring these two species to Sclero- domus, the type-species of which, S. denticulatus (Busk), was well described by _Levinsen (1909 : 302). Levinsen (p. 304) examined a small fragment of Busk’s material of Bifaxaria corrugata and had “‘ no doubt that this species belongs not only _ to another genus but even to another family than Sclerodomus denticulatus”’. Ihave | not seen Levinsen’s fragment, but it evidently came from an old part of the zoarium and may in some ways have misled Levinsen, as also did Waters’s description of the | ovicell (1888 : 15)—"‘ formed by a swelling of the superjacent zooecia’’. Both
species have normal ovicells (cf. Harmer, 1957: 869). Nevertheless, there are _ differences (e.g. the absence of an avicularian cross-bar, which is sometimes thought to be important).
| COMPARISON OF THE SPECIES. S. papillatus and S. corrugatus differ markedly in | the details of their primary frontal walls (Text-fig. ra, c). In both there is a median _ longitudinal line (‘‘ keel ’’) which is a little more thickly calcified (cf. Harmer, 1957 : 866). In S. papillatus this line runs up into a prominent, mucro-like lip (Text-fig. Ic), except in fertile zooecia where the edge of the frontal shield is transverse (Harmer, | pl. LVII, fig. 16 as S. corrugatus, see below). In S. corrugatus the longitudinal line ‘is less pronounced. The edge of the shield is more or less transverse (Text-fig. 1A), or slightly prominent, in the non-fertile zooecia, and in the fertile zooecia it forms a broad rounded lip (Busk, pl. XIII, fig. 3a). In S. fapillatus the pores are peripheral, arranged in one or two series. Sutures or cracks run outward from the median line, ‘most of them ending ina pore, In S, corrugatus the pores are central, arranged on 1) zoon, 14, 3 6§
60 ASB. HCA STEEL NiG:s
each side of the median line in several longitudinal series, and there are no sutures or cracks. Laterally there are longitudinal ridges, and the extension and thickening of this ridged calcification forms the striated thickening which gradually obliterates the original frontal wall and transforms the appearance of the older parts of the colony by enveloping the branches. The pores are continued upwards as oblique tubes in this thickening, the oblique transection of the tubes and the wall giving the appearance of oval or elongate pores (Text-fig. ra). In S. papillatus it is the develop-
om +4
&
ep AERA TEM
| 050mm |
Scale for fig B
| 0:50mm
Scale for fig A.C
Fic. t. Sclerodomus corrugatus and S, papillatus. Zooecia near the growing point for compari- son of their characters before obliteration of the primary walls by secondary developments. A. S. corrugatus, 1887.12.9.379A, lectotype. B, Cc. S. papillatus, 1964.2-2.4, Siboga collec- tion, specimen 583A", 8B, is the large avicularium which is seen obliquely in c,
OBSERVADIONS ON DYPE-MATERIAL, POLYZOA 61
ment of a covering of avicularia which leads to marked differences in the appearance of the branches at different stages. The youngest branches are slender with almost hyaline, glistening zooecia, in which the lateral frontal-pores and the fine sutures can quite readily be seen. As they get older they become opaque, and the covering of avicularia begins to develop down each side of the branch. The oldest branches are covered with a mosaic of these avicularia, which makes them considerably thicker than the younger branches; and a large avicularium is frequently developed at the bifurcations (but not in the axils). The first avicularia to appear are single lateral ones on each side of each zooecium, forming a zig-zag series down each side of the branch. An occasional one of these is enlarged. In fig. 1c the first lateral avicularia have appeared, the youngest zooecia still being without them. One is large (figs. 1B and C) and the most proximal is not fully developed.
IDENTITY OF THE SIBOGA MATERIAL. The difference in age of the specimens probably accounts for the rather surprising fact that, having correctly elucidated Busk’s work on these two species, Harmer (1957, p. 869) was in doubt as to the identity of the Siboga species, and finally placed it in S. corrugatus. The Siboga specimens agree perfectly with S. papillatus, and differ from S. corrugatus, in the shape of the zooecia and peristome, the distribution of the pores, the presence of a lateral avicularium on each zooecium and of enlarged ones beside some of the ovicells, the presence of secondary branches, and the covering of the older parts by avicularia. This covering was not described by Busk, but its earlier stages are present on the proximal part of the main axis of the Challenger specimen of S. papillatus. Busk’s pl. XXIV, fig. D resembles Harmer’s pl. LVII, fig. 10 with remarkable closeness. Only the large lateral avicularium is absent in the Siboga figure, but these are present in the Siboga material.
The apparent differences between the Siboga material and the type-specimen of S. papillatus lie wholly in their age. The type-specimen consists almost entirely of branches in the younger phases, the most proximal having reached the phase with a zig-zag line of avicularia, and no older phases being represented. The greater part of the Siboga material is in the oldest phase, and there are only few young branches. It therefore looks decidedly stouter than the type-specimen, and different in surface sculpture, etc. ; but if branches in the same phase are compared the agreement in all characters, including dimensions, is found to be exact!, except for the absence of enlarged avicularia at the bifurcations in the Challenger specimen. As they only appear in the older, thicker parts of the Siboga material, the difference in the age of the material probably also accounts for this.
Station 122, type-locality of S. corrugatus, is off Brazil, and Station 196, type- locality of S. papillatus, is off Celebes. Thus the geographical probabilities also support my conclusion that the Siboga material belongs to S. papillatus.
The synonymies of the two species given by Harmer (pp. 867, 868) are, as already indicated, correct ; but the one to be applied to the Siboga specimens is that given for Sclerodomus papillatus (p. 868).
1 Miss Cook (in litt.), ‘‘ I have made several sets of measurements and the agreement is quite startling. ‘Exact ’ is certainly the right word ”’,
62 AS. 1B) EVA'S DEINIGS
Flustra octodon Busk
Flustva octodon Busk, 1852: 49, pl. LVI, fig. 4, pl. LVIII, fig. 5.
Hincksinoflustva octodon Bobin & Prenant, 1961: 167 (as Flustyva octodon), 169, 170, text-fig. 4L-IV.
Hincksina (Flustya) octodon (Busk) Gautier, 1962: 51. (Hincksinoflustya in text. Synonymy.)
DISTRIBUTION. Coast of Spain (probably Mediterranean) ; Mediterranean ; Atlantic coast of Morocco.
LECTOTYPE, chosen here: 1899.7.1.112B. PARALECTOTYPES. 1899.7.1.5789. 10963.3.12.1 (previously unregistered).
LOCALITY OF THE TYPE-MATERIAL. This material consists of two spirit specimens, figured by Busk (1852, pl. LVI, fig. 4; left-hand figure, 1899.7.1.5789; right hand figure, 1899.7.1.112B), aslide from which pl. LVIII, fig. 5 was drawn (1963 .3.12.1), and two preparations made in comparatively recent years from 112B. All these specimens came from M’Andrew’s Spanish collection, and the locality published by Busk is “‘ Coast of Spain, M’Andrew”’.
The question arises as to whether the specimens came from the Atlantic or the Mediterranean side of the peninsula. Busk (1852, 1854) published descriptions of M’Andrew’s Polyzoa as follows: from the “ Coast of Spain’”’, Alysidium lafontit (p. 14), Scrupocellaria macandret (p. 24) and Flustra octodon (p. 49) ; from the “ Bay of Gibraltar ’’, Membranipora rossellit (p. 59), M. calpensts (p. 60), Lepralia violacea (p. 69), L. spinifera (p. 70); from the “ Mediterranean ’’, Lepralia linearis (p. 71), Eschara foliacea (p. 80).
The shde of A. dafontit (1899.7.1.3752) has labels superimposed on the old ones, as was Busk’s custom!. The original label reads “‘ Coast of Spain ’’, the newer one “Mediterranean ’’. The original drawings of S. macandrei (pl. XXIV, fig. 1), and F’. octodon (pl. LVIII, fig. 5) are both inscribed ‘‘ Mediterranean ”’.
Conversely, three of the four slides of the two species recorded as from the Mediterranean, are labelled “‘ Coast of Spain’’. The four slides are: L. linearis, S. Coast of Spain, 72-128 fms., 1899.7.1.2330; Mediterranean, 1899.7.1.23109. E. foliacea, Coast of Spain, 1899.7.1.1417; S. Coast of Spain, 1899.7.1.1412.
There is thus evidence that suggests that all the M’Andrew Spanish material described by Busk in the British Museum Catalogue came from the Mediterranean side of the peninsula.
The locality given on the specimen of Flustva octodon, illustrated in pl. LVI, fig. 4 (right-hand specimen), is “‘ Between Vigo Bay and Tunis’’. Unless this is an error, it must be a generalization covering the area visited by M’Andrew, and would not be incompatible with a Mediterranean origin for the specimen.
Flustra octodon is the type-species of Hincksinofiustva Bobin & Prenant (1061).
1 His specimens were relabelled, sometimes more than once, in accordance with changes in the style of
label in use in his collection ; for further details see Hastings, MS, note in Busk Register, British Museum (Natural History).
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 63 Lepralia melolontha Landsborough
Lepralia nitida, branched in a fine dendritic manner, Johnston, 1847: 319.
Lepralia nitida, specimen d., “ Lep. nitida var.”’, Gray, 1848: 121.
Lepralia melolontha Busk; Landsborough, 1852: 319, pl. XVIII, fig. 70; Busk, 1854: 79, pl. LXXXV, fig. 3.
Membraniporella melolontha Busk Hincks, 1880a: 202, pl. XXVII, figs. 9, to.
Aspidelectva melolontha Levinsen, 1909 : 160.
Aspidelectra melolontha Busk: Marcus, 1940: 199, text-fig. 103.
DISTRIBUTION. This species is only known from localities of more or less reduced salinity bordering the North Sea. The distribution given by Marcus is Denmark (one locality, Horns Rev!); Heligoland; Belgium; southern England [eastern England would be more correct].
LECTOTYPE, chosen here: 1854.11.15.140, British Museum Catalogue Coll. Thames Estuary, Lieut. Thomas.
PARALECTOTYPES. 1899.7.1.1363, Busk Coll. Thames Estuary, Lieut. Thomas. 1899.7.1.1364, Busk Coll. Thames Estuary.
OTHER MATERIAL. River Roach, Essex (Dr. Baird, 1865.6.28.1) ; River Roach (Norman Coll., I9I1.10.1.741 part, I919.6.24.25, 25a, 26); Goodwin Sands, Kent (G. Merritt, 1884.5.10.1) ; Kirkwall, Orkney (Johnston Coll., Gray ra1d, 1847.9.16.130) ; Locality ? (Hincks Coll., 1899.5.1.727.)
Remarks. The first published account of this species is by Landsborough. He gave credit to Busk for the discovery of the species, and published a description and drawings supplied by Mrs. Gatty, who had recognized it independently (see Busk). Busk has hitherto been named as the authority for the species, but strictly Lands- borough’s name should be given. Under these slightly confused circumstances it seems desirable to choose a lectotype.
The Busk material in the Museum consists of three slides from the Thames Estuary material mentioned by Landsborough. These are syntypes, and, in the absence of any of the material figured by Mrs. Gatty, I have chosen one of them as lectotype. The material from Bawdsey, figured by Busk (1854), 1s not in the Museum, but would not be a syntype.
Hincks (p. 203, footnote) thought that Mrs. Gatty (in Landsborough) was right in identifying Johnston’s dendritic variety of Lepralia mitida from Kirkwall, Orkney Islands, with this species. Marcus (p. 201) noted the Orkney record as being un- certain but not improbable. It is confirmed by Johnston’s specimen (1847.9.16.130, Kirkwall). This consists of several small colonies of L. melolontha on pieces of Mytilus shell. They are growing with Electva monostachys (Busk) and Conopeum veticulum (Linn.), an association which suggests a brackish-water habitat. The same three species are, for example, found together from the River Roach, Essex (Norman Coll., 191r.10.1.48r and 483), where one shell in particular shows all three intermingled as in the Orkney specimen.
One of the Johnston colonies of L. melolontha has its membraniporine ancestrula
1 Horns Rev, a shoal to the west of Jutland.
64 A. B. HASTINGS
intact. It has formed a pair of distal zooecia, each of which has budded a further pair. Only the left-hand zooecium of each of these two pairs has budded, and from these buds the two arms of the colony fan out and diverge—see also Hincks (fig. Io) and Marcus (text-fig. 103) which both show fan-shaped branches starting from a single zooecium.
L. melolontha is the type species, by monotypy, of Aspidelectra Levinsen (1909).
Lepralia multidentata Thornely
Lepralia multidentata Thornely, 1905 : 120, pl. [unnumbered], fig. 9. Hippoporella multidentata (Thornely) Harmer, 1957: 1099, pl. LX XIII, figs. 9-12. Lepralia purpurea Thornely, 1905 : 120, pl. [unnumbered], fig. 13.
DistTRIBUTION. Ceylon; India; Sulu Archipelago. LectotyPE, of L. multidentata, chosen here: 1906.12.3.4. Ceylon.
PARALECTOTYPE, of L. multidentata: Cambridge Museum, Feb. 22, 1906.9. Ceylon.
HotoryPe, of L. purpurea, the only specimen: 1906.12.3.6. Ceylon.
Remarks. Under this species Harmer listed three specimens from the Thornely Collection in the British Museum. They are the one chosen here as lectotype, and two (not named by Thornely) which he had himself found, accompanying specimens of other species in the Thornely collection. The Cambridge specimen was named by Thornely and is a syntype.
Harmer recognized Lepralia purpurea Thornely as a synonym of L. multidentata, but did not list the specimen.
Lunulites owenii Gray
Lunulites owenit Gray, 1828: 8, pl. III, fig. 15. Cupuladria owenit (Gray) Cook, 1965 : 213, pl. 2, figs. 34, B, text-fig. 2c. (Synonymy.)
DISTRIBUTION. Recent, West Africa. Pliocene, South Carolina; Florida. JJio- cene, North Carolina ; Florida. (Teste Cook).
LECTOTYPE, chosen here: 1899.7.1.4879, W. Africa.
The slide 1899.7.1.4879, which forms part of the Busk Collection, is labelled “B.M.. J. E. Gray’, indicating that Busk received the specimen from the British Museum. It is the only representative of Gray’s material of this species now in the Museum and Kirkpatrick (MS. Catalogue) accepted it as the type-specimen. It is the specimen figured by Busk (1854, pl. CXV, fig. 3).
REMARKS. Cupuladria owenti is discussed by Cook.
1 Their numbers, incorporating numbers in Harmer’s lists, are:
1936.12.30.41A with Cleidochasma protrusum 1936.12.30.41D. Ceylon. See Harmer, 1957 : 1040. 1936.12.30.117B on Celleporaria columnaris 1936.12.30.117A. India. See Harmer, 1957 : 677.
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 65
Membranipora amplectens Hincks
Membranipora amplectens Hincks, 1881a: 129, pl. III, fig. 7.
Electva amplectens (Hincks sp.) MacGillivray, 1889: 322, pl. CLXXXVII, figs. 14, 14a; Levinsen, 1909 : 66, 146, 147, pl. IX, figs. 1a, b.
Membranipora amplectens sp. n. [sic] Hincks, 1892 : 332. (Named as “ type of a new genus ”’ Introduction of Heterooecium follows.)
Membranipora amplectens (Heterooectum) Hincks: Waters, 1898 : 671.
Heterooecium amplectens Hincks : Harmer, 1902 : 284.
DISTRIBUTION. Australia. LECTOTYPE, chosen here: 1899.5.1.702, Hincks Coll. W. Australia. PARALECTOTYPE : 1899.5.1.703, Hincks Coll. W. Australia.
OTHER MATERIAL. Australia (1897.5.1.497, 498, mounted by Jelly) ; Australia (Vine Coll., 1934.10.20.85, 91) ; Port Phillip Heads, Victoria (Bracebridge Wilson CollPers888e 11-14. 117, 283).
All the specimens listed are growing on algae.
REMARKS. WM. amplectens is the type-species, by original designation, of Hetero- oectum Hincks (1892 : 332).
Membranipora crassimarginata var. erecta Busk
Membranipora crassimarginata var. erecta Busk, 1884 : 63, pl. XIV, fig. 3. Acanthodesia perfragilis (MacGillivray) Hastings, 1945: 98. (Synonymy.)
DISTRIBUTION. Australia. LECTOTYPE, chosen here: 1944.1.8.160. Challenger St. 162, Bass Strait.
PARALECTOTYPES. 1887.12.9.309. Challenger, St. 162, Bass Strait.
The other paratypes are all marked with the dubious locality, St. 151, Heard iis¥andy (see below). They are: 1887.12.9.310, 311; 1899.7.1.1004, 1005; 1944.1.8. 161-164.
REMARKS. Harmer (1926: 224) accepted a specimen from St. 162, Bass Strait, 1887.12.9.309, as the original of Busk’s figure (1884, pl. XIV, fig. 3). The actual figured specimen (1944.1.8.160) had not been deposited in the British Museum at that time. It likewise comes from St. 162 and is recognizable by its detailed agreement with the figure. It was found in the supplementary Busk Collection, received by the Museum in 1943 (see section 2, above). It is chosen here as lecto- type.
There is evidence (Hastings, paper in preparation) that Busk’s record of Australian species (including this one) from Heard Island (Busk, 1884: X, those species! from St. 151 marked D) may have been due to an error in sorting before the collections were sent to specialists. M. crassimarginata var. erecta is synonymous with Acan- thodesia perfragilis (MacG.), see Hastings (1945).
1 Salicornaria clavata to be excepted. The Heard Island species is distinct from the Australian, see below.
66 A. B. HASTINGS Membranipora hexagona Busk
Membranipora hexagona Busk, 1856: 308, pl. XII, fig. 4; Hincks, 1880a: 143, pl. 18, fig. 7 (after Busk) ; 1881a: 6, pl. IV, fig. 6.
HorotyPE, the only specimen: 1899.7.14.1083. Devon, Miss Cutler.
REMARKS. Membranipora hexagona has remained an enigma. Hincks (1880a: 143), for example, remarked that he knew “ nothing of M. hexagona but what may be gathered from the brief description and figure in the ‘ Zoophytology’”’. He reproduced part of Busk’s figure. Subsequently (18815: pl. IV, fig. 6) he figured a specimen lent to him (see 1881a: 6) by Busk, presumably the holotype.
Examination of the holotype shows that it is not membraniporine, and that Busk based the name on a mutilated specimen of a species with a calcareous frontal- shield. This shield has been completely broken away in most of the zooecia, and even the lateral walls are somewhat abraded, sometimes exposing the cavities
of pore-chambers. On a concave part of the substratum, a few zooecia have been |
less worn. In these the lateral walls stand higher and in two zooecia small parts of the broken frontal-shield project at the margins of the aperture. The floor of a hyperstomial ovicell also remains. The frontal membranes (which were below the frontal shields) have adhered to the floors of the zooecia, and remained intact, and with them the opercula. The opercula are widely open as in Busk’s and Hincks’s figures. It is thus clear that M. hexagona was based on a misconception.
Supposed material of M. hexagona has been recorded from various localities :
Busk himself gave Flustra coriacea Johnston (not Esper) as a synonym of WM. hexagona. I. coriacea was recorded by Johnston (1847: 348, pl. LVI, fig. 8) from specimens from Sana Island, Fowey and the Isle of Man. Busk listed all three localities in the distribution of M. hexagona, but Hincks considered that only the specimen! from the Isle of Man (collected by Forbes, whose description was used by Johnston) should be identified with Busk’s species. This was the first supposed record of M. hexagona from the Isle of Man. Two subsequent records were included in the first edition of the Fauna List (Moore, 1937: 199). The specimens are not available and their identity cannot be surmised.
Smitt (1867 : 371) tentatively included M. hexagona in the synonymy of Mem- branipora pilosa forma membranacea (Mill.). The latter is a synonym of Electra crustulenta (Pallas), and Smitt evidently misinterpreted Busk’s figure as showing the calcified opercula characteristic of that species.
Calvet (1896 : 253) identified material from 180 m. in the Gulf of Gascony as M. hexagona. There is no clue to its true identity.
Cipolla (1921 : 204) recorded Pliocene fossils from Sicily as Wembranipora exagona
‘No other evidence has, however, been produced for thinking that Johnston confused two species under Flustra coriacea. It appears that only the Sana Island specimens came to the British Museum in the Johnston Collection, for they alone are listed by Gray (1848 : 115), and no others have been traced in the collection. These specimens belong to Micropora coriacea as generally understood, and include the specimen chosen by Brown (1952 : 126) as lectotype of Flustrva coriacea Johnston not Esper. The exist-
ence of a second British species of true Micropora (AI. normani see below) does not affect. this question of whether Johnston may have confused a Membraniporine species with J. coriacea.
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 67
[stc] and quoted earlier uses of the name by Fischer (1870) and Neviani (1896 and 1900).
Bassindale (1940 : 194) included Busk’s original record in his list of Polyzoa of the Severn Estuary. He noted (p. 197) that a later record from that region (Purchon, 1937 : 329 as M. hexagonia [sic|) was based on specimens of Alcyonidium polyoum (Hassall), one of which I have seen.
Cellepora hexagona von Hagenow (1839) was referred to Membranitpora by Voigt (1959 : 29, see also Berthelsen, 1962: 51), and thus became a senior homonym of M. hexagona Busk.
Membranipora pilosa var. multispinata Hincks Membranipora pilosa var. multispinata Hincks, 1882: 117, pl. V, fig. 6. Membranipora multispinata Hincks, 1892 : 334; 1895: 11 footnote; Waters, 1924: 607, 608, pl. XIX, fig. 9 (as multispina on p. 607, and as multispinosa in expl. pl.). DISTRIBUTION. Western Australia.
LECTOTYPE, chosen here: 1899.5.1.700, Hincks Coll., mounted by Jelly. W. Australia.
OTHER MATERIAL. 1897.5.1.491, specimen figured by Waters; 1897.5.1.492. Both from W. Australia and mounted by Jelly. There is no evidence as to whether Hincks saw these two specimens which, being closely similar to the lectotype and also mounted by Jelly, may be parts of the original material. Waters’s figure of 1897.5.1.491 is not quite accurate as regards the arrangement of the zooecia in the region which appears to be ancestrular.
Remarks. There is little to add to Hincks’s description. The walls are extremely delicate, glistening and transparent. As described by Hincks, there are five long, more or less stout, socketed spines on each zooecium. They are pointed and may be very long and curved. It appears that the opesia does not extend to the distal end of the zooecium and that the paired spines, described by Hincks as “ near the bottom ”’, are in fact in the distal corners. In addition there may be one to three small, almost thread-like spines, similarly socketed, on the lateral gymnocyst alongside, but at a distance from, the opesia.
The proximal gymnocyst may have a few bright spots, either scattered or arranged in one or two transverse rows near the proximal end. These are the bases of tiny, sharp, thorn-like spines projecting from the inner surface of the transparent wall.
Waters considered that the “ articulated’ spines distinguished this species from Electra and related it to the “ M. corbula group’’. He probably had particularly in mind the series of incurved marginal spines which differ, as he said, from those of E. pilosa. The erect spines seem to be remarkably like those seen in some forms of E. pilosa and E. verticillata, and both the erect and the incurved spines resemble
those of E. monostachys (Busk), see for example the figure given by Marcus (1950 : 8,
text-fig. 1, as Membranipora (E.) hastingsae!). E. multispinata differs from E.
1 Examination of type-material has shown that E. monostachys is a senior synonym of E. hastingsae (Marcus), not a junior synonym of EF. crustulenta as supposed by Marcus (1950).
68 A. B. HASTINGS
monostachys in its additional gymnocystal spines, its internal spinules, its dimensions, fragile texture and transparency ; but to me they seem clearly to be congeneric.
Membraniporella agassizii Smitt
Membrantporella agassizii Smitt, 1873: 11, pl. V, figs. 103-106 ; Levinsen, 1909: 17; Hastings, 1964 : 258, 262.
Ho.otyre, the only specimen: Riksmuseum, Stockholm, 262. Off Cojima,
Cuba, 450 fms., March 1869, Pourtales. This is recognizably the zoarium figured by Smitt.
REMARKS. Each kenozooecium has a small round opesia, closed by a membrane, as shown in Smitt’s figure.
At two points an autozooecium has been budded in the kenozooecial crust. In one instance it lies in a plane parallel to the surface of the branch and forms part of the crust. The other one projects at right angles to the parent branch, and, together with some kenozooecia, appears to form the base of an incipient branch.
The changes with age in the zoaria of Polyzoa, particularly in the erect zoaria of some Ascophora, are well-known and may be very marked. Some striking examples in Cretaceous species have been discussed by Voigt (1960) and Wiesemann (1963). In Membrantporella agassizit the secondary layer is built up of both kenozooecia and avicularia. In Sclerodomus papillatus (see above under Bifaxaria) it is mostly, per- haps entirely, made up of a profusion of avicularia. In Sclerodomus corrugatus (above) and Foveolaria elliptica Busk (Hastings MS.) the original surface is obliterated by massive calcification, with characteristic texture—striated in the one species, perhaps best described as “ fibrous-looking ”’ in the other.
No other generic placing has been proposed for M. agassizii. The zooecial characters are those of Membraniporella and the taxonomic significance of the multi- plication of avicularia and kenozooecia is still unknown (cf. Hastings, 1964 : 258).
Menipea fuegensis Busk Mentrpea fuegensis Busk, 1852: 21, pl. XIX, figs. 1-3. Menipea patagonica Busk (part), 1852: pl. XXIII, fig. 1 (not p. 22, pl. NXV, figs. 1-3, pl. XXVII, figs. x, 2). Tricellavia aculeata (d’Orbigny) ; Hastings 1943 : 356. (Synonymy and distribution.)
DISTRIBUTION. Widely distributed in sub-Antarctic waters.
Lectotype, chosen here: 1854.11r.15.262. Falkland Islands. H. [Hooker).
REMARKS. Busk recorded this species from Tierra del Fuego (Darwin) and the Valkland Islands (Hooker), but the only type-material in the British Museum is the one specimen now chosen as lectotype. This specimen was mounted and labelled by Busk as Menipea fuegensis, and sent to the Museum as part of the “ British Museum Catalogue Collection ’’, for which see Hastings (1943 : 304). As noted by Hastings (1943 : 359) the ancestrula and first few zooecia of the type were figured
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 69
by Busk (pl. XXIII, fig. 1), but, by some curious error, under the name Menipea patagonica. ‘There is no evidence as to the locality of the material figured in pl. XIX. Mentpea fuegensis Busk is a junior synonym of Tvicellaria aculeata (d’Orb.).
Micropora normani Levinsen
Micropora sp. Levinsen, 1902 : 7 (footnote).
Micropora noyvmani Levinsen, 1909 : 162, pl. VIII, figs. 3a, 3b (IM. coriacea on plate, M. normani in explanation of plate).
Micropora coriacea Hincks (part) 1880a:174 (specimens with avicularia).
DISTRIBUTION. Southern and western coasts of Great Britain and Ireland from Hastings to Shetland; west coast of Spain.
NEOTYPE, chosen here: IgirI.10.1.623, Norman Coll. Antrim.
Levinsen stated that he had only seen “a little fragment ’’ of this species, and had lost it. A neotype is therefore needed. The specimen chosen was labelled by Norman as “ Micropora n sp [sic] with avicularia, see Levinsen’’. It clearly shows the characters of Levinsen’s species, including those of the ovicell.
OTHER MATERIAL. Hastings, Sussex, type-locality (1897.5.1.597 mounted by Jelly) ; Antrim, neotype-locality (Hincks Coll., 1899.5.1.737, slide bearing three specimens of Micropora, of which two (on shell) are M. norvmani; Norman Coll., HOLM TOnt.7A4); Liverpool Bay (Thornely Coll., 1936.12.30.349 B); Guernsey (Norman Coll., 1965.1.2.1 and 1919.6.25.82) ; West coast of Spain (1872.2.3.146).
RemARKS,_ In addition to the differences noted by Levinsen (presence of avicula- ria, the characters of the ovicell, “‘ calcified ’’ opercula) these specimens have smaller zooecia and ovicells than are seen in British specimens of typical M. corzacea!, and the pores in the cryptocyst are few and irregularly scattered. Levinsen’s figure shows this latter character. Hincks included specimens with avicularia in M. coriacea and, except for its more numerous and evenly distributed pores, his fig. 7 (pl. XXIII) appears to represent M. normani. His slide 1899.5.1.737 is evidently one of the comparative mounts that he made of British forms? and this supports the conclusion that Hincks’s specimens with avicularia represented M. normani.
I have not confirmed that the opercula are calcified, but they are whitish and evenly granular, in contrast to the clear, brown, chitinous opercula of M. coriacea. The avicularia are sporadic in their distribution, but a few are present in every colony which shows the other characteristics. The granulation of the cryptocyst is a distinctive feature, being fine and close and very even. That of M. coriacea is somewhat coarser and less close, and, over the greater part appears less regular owing to interruption by the more numerous pores.
The differences in dimensions and number of pores are evident to direct examina- tion. Sample measurements and counts made by Miss Cook, as a trial, support these observations. They are based on a total of 30 zooecia and 16 ovicells from 4 colonies
1M. coriacea Johnston not Esper, see footnote above under Membranipora hexagona. 2 See Hastings (forthcoming paper on Hippothoa).
70 A. B. HASTINGS
of M. coriacea from 4 different British localities ; and 40 zooecia and 20 ovicells from 5 colonies of M. normani from 3 different British localities. Many more would be needed to obtain statistically reliable figures.
MEASUREMENTS OF ZOOECIA AND OVICELLS. M. coriacea: Lz 0:40-0:70 mm., average 0-50 mm. lz 0-30-0-48 mm., average 0:39 mm. Lov o-18-0-25 mm., average 0:22 mm. lov 0:24-0:32 mm., average 0:28 mm. M. normani: Lz 0-30- 0-47 mm., average 0-39 mm. lz 0-2I-0-40 mm., average 0:32 mm. Lov 0:12-0:17 mm., average 0-15 mm. lov 0-15—0-26 mm., average 0-2I mm.
NUMBER OF CRYPTOCYSTAL PORES. M.coriacea: 30-60, average 45. M.normant: I2-15, average 20.
On the evidence available to me, M. normanz is distinct from M. coriacea. The two are easily recognizable and there is no intergrading.
Species of Micropora with avicularia from (a) Port Phillip, Victoria (Bracebridge Wilson Collection) and (b) Madeira (Norman Collection) and the Mediterranean (Busk Collection) have been identified with M. coriacea, but it seems likely that both species are distinct from both M. coriacea and M. normant. M. coriacea s. lat. needs full revision.
To sum up, M. normani differs from British M. coriacea s. str. in the presence of avicularia, the smaller zooecia, the smaller number of cryptocystal pores, the whitish, granular opercula and the absence of the ridge (sometimes umbonate) on the ovicell which may, however, have a small umbo.
The distribution of M. normani is Armorican (see Lagaaij, 1952 : I1), extending from Spain to the western and south-western shores of the British Isles.
CHARACTERS OF THE GENUS Muicropora. M. coriacea is the type-species of Micropora Gray (1848 : 115), see Brown (1952: 125). If it is confirmed that British M. coriacea s. str. is without avicularia, generic definitions, e.g. those of Brown (1952) and Bassler (1953), will have to be emended by adding the words “‘ when present ”’ to the characterization of the avicularia.
Brown (1952 : 126) noted that the avicularia in Micropora are not adventitious, as described by Harmer, but are what we now term interzooecial. This can be seen in the neotype and other specimens of M. norman.
The ovicells of both M. coriacea and M. normani are not endozooecial (as stated by Levinsen, 1909 : 162), but recumbent. This can be observed by examining the growing edge of the colony where the complete ovicell projects at the distal end of the fertile zooecium before the distal and distal-lateral zooecia appear. As these zooecia develop the ovicell becomes surrounded by them and thus imbedded in the zoarium.
Spines are usually stated to be absent in Micropora, but a single specimen of a species (undescribed?) from Algeria (1869.10.6.6) has, on many zooecia, a pair of stout distal spines, rather widely set (being about opposite to the proximal corners of the distal zooecium). More rarely there is one median distal spine.
' The term recumbent was introduced by Canu & Bassler (1917 : 66), see Brown (1054: 244, footnote).
meh
OBSEHERVATVONS ONE TYPE-MATERIAL, POLYZOA 71
Salicornaria clavata Busk
Salicornaria clavata Busk (part), 1884 : 88, pl. XII, figs. 8, 8c, text-fig. 5 (not figs. 8a, b, St. 162 Bass Strait = C. australis).
Cellaria austvalis MacGillivray Waters (part), 1888: 16, pl. 11, figs. 1-4.
Cellaria clavata (Busk) Hastings, 1947 : 236.
DISTRIBUTION. S. Indian Ocean. LECTOTYPE, chosen here : 1887.12.9.395 part. Challenger St. 151, Heard Island.
PARADE CIOLYPES.. 1887.12.9.391, 396;. 1899:7-1-1545; 1934.2.16.34. St. 149D, Kerguelen. 1882.7.29.75 ; 1887.12.9.389 ; 1899.7.1.1550; 1934.2.16.35. St. 151, Heard Island. 1887.12.9.390, 393; 1899.7.1.1551. Prince Edward Island.
OTHER MATERIAL. St. 149D (1963.2.12.200); St. 151 (1963.2.12.243). Both specimens received from Dundee Museum. No evidence as to whether named by Busk.
REMARKS. Waters accepted Busk’s treatment of the Australian and South Indian Ocean forms as one species, and used the earlier name Cellaria australis MacGillivray. Hastings distinguished Cellaria clavata (Busk) from C. australis! but did not choose alectotype. A colony from St. 151 is now chosen, this being the station from which Busk’s figured material of this species was obtained.
The colony from St. 149D (1887.12.9.396) is large, has a mass of anchoring
_ rootlets forming a short stalk, and very numerous branches which, in its dry state, _ are close together and more or less parallel to each other,” as if growing vertically. _ The rootlets attaching the lateral branches to the parent branches are brittle in their _ dry state, and many branches have fallen off, but the remaining colony is 93 mm. tall and 42 mm. =< 35 mm. at its thickest point. The longest branch (broken at the
tip) measures 78 mm. The average diameter of the branches is 2 mm.
The figures of the chitinous parts given by both Busk and Waters represent true C. clavata. The opercula of C. australis differ from those of C. clavata in their less conspicuous ‘ foramina ’’, which are not visible at all focal levels, and in the absence of the granulations shown in Busk’s and Waters’s figures. In C. australis the oper- cula are very slightly shorter and distinctly narrower. Miss Cook has measured chitinous parts mounted by Busk (the figures being necessarily based on rather few
measurements) :
C. australis, 1899.7.1.1546. Lo 0-08-0-09 mm., average 0:085 mm. lo 0-09— 0-105 mm., average 0-10 mm.
C. clavata, 1899.7.1.1545. Lo 0-085-0-10 mm., average 0-095 mm. lo 0:13-0:16 mm., average 0:I5 mm.
C. clavata, 1882.7.29.75. Lo 0-12-0:I4 mm., average 0-13 mm. lo 0:12-0:14 mm., average 0-13 mm.
1 They differ in areolation and jointing, and in the presence of cryptocyst ridges in C. clavata. * The flexibility of the rootlet attachment of the lateral branches, before drying, has made this possible,
2 AY. B.S ELAS TD ENG's:
The third of these preparations, in which the opercula are as long as wide, was made from a fertile branch and has remarkably large embryos throughout.
The mandibles of C. australis are considerably wider than those of C. clavata, being wider than the operculum :
C. clavata, | mand. 0-10-0-II mm., average 0-105 mm. C. australis, | mand. 0:14-0:17 mm., average 0-155 mm.
The specimens of C. clavata from Prince Edward Island differ in their longer zooecia and in the areolation of the fertile zones (Hastings, 1947: 237). The opercula, measured from those mounted by Busk, resemble those of C. clavata from the other stations in width, but are shorter :
1899.7.1.1551. Lo 0-09-0-:1I mm., average 0-10 mm. lo 0-II-0-13 mm., average o-II5 mm.
The width of the mandibles is the same as for those from the other stations quoted above.
Siphonoporella nodosa Hincks
Siphonoporella nodosa Hincks, 1880b: 90, pl. XI, fig. 10; Harmer, 1900: 231; Levinsen, 1909 : 170, pl. VI, figs. 2a, b.
DiIstRIBUTION. Australia.
Lectotype, chosen here: Specimen figured by Hincks, E.C. Jelly Collection, Manchester Museum. Australia.
PARALECTOTYPE. 1899.5.1.685, Hincks Coll., mounted by Jelly. Australia.
OTHER MATERIAL. Australia (Bracebridge Wilson Coll., 1897.5.1.582, mounted by Jelly, but not known to have been examined by Hincks) ; Western Australia (EK. Lempriere Coll., 1948.3.12.2, on Metamastophora plana).
REMARKS. The figured zooecia are recognizable in the Manchester Museum specimen, and I have therefore chosen it as lectotype. Levinsen figured a specimen resembling the lectotype in having a well developed gymnocyst with usually more than 2 nodules.
In the British Museum specimens the gymnocyst of nearly all the zooecia is much narrower, usually with only 2 nodules, one in each proximal corner. In the paired daughter-zooecia, at the bifurcation of the series, one or both the inner! nodules is reduced or absent. In each Jelly specimen there are, however, a few zooecia with a wider gymnocyst and more numerous, more irregular nodules, and these are more frequent in 1948.3.12.2. In the latter specimen the zooecia with narrow gymno- cyst often have a small median nodule in addition to the two in the corners.
The most recent comments on Siphonoporella have been made by Silén (1941 : 62) and Cook (1964: 57). References to some earlier work are to be found in both these papers. Canu & Bassler (1929: 149) described a species from the Philippines. S. nodosa is the type-species of the genus.
'j.e. the nodule in the corner next to the other zooecium of the pair,
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 73
Zocor GENUS AEPPOPLEURTFERA AND SOME OF ITS SPECIES
Hippopleurifera Canu & Bassler
Hippopleurifera Canu & Bassler, July 1925: 679. (Hippopleurifera biauriculata Reuss, 1847 [stc] listed, with full reference.) Hippopleurifera Canu nov. Canu & Lecointre, 1925 : 9 (Genus only, listed.)
_ Hippofleurifera {sic| Milne-Edwards, 1836; Canu, 1926: 764. (H. sedgwicki Milne-Edwards,
1836 listed.)
Hippopleurifera new genus Canu & Bassler, 1927: 7. (Defined, with Eschava sedgwicki Milne- Edwards designated as type.)
Hippopleurifera Canu & Bassler, 1927; Canu & Bassler, 1929: 326. (Same definition and type as in 1927.)
Hippopleurifera nov. Canu & Lecointre, 1930: 86. (Same definition, in French, and type as in Canu & Bassler, 1927.)
Hippopleurifera Canu & Bassler, 1927: Bassler, 1935: 127. (Listed with a reference to Canu & Bassler, 1929, and with EF. sedgwicki as type.)
| Hippopleurifera Canu ; Hastings, 1949: 524. (Discussed with E. sedgwichi as type.)
Hippopleurifera Canu & Lecointre, 1925; Vigneaux, 1949: 96. (Same French definition and type as Canu & Lecointre, 1930.)
Hippopleurifera Canu & Bassler, 1925; Lagaaij, 1952: 92. (Revised diagnosis with Eschara biauriculata Reuss named as type by monotypy.)
Hippopleurifeva Canu, 1927 ; Bassler, 1953: G196. (Brief definition with EF. sedgwicki as type.)
Hippopleurifera Canu & Bassler, 1924; Buge, 1957: 265. (Redefined with E. biauriculata as
type.)
| Hippopleurifera Brown, 1958: 65. (Footnote on priority of FE. biauriculata as type-species.)
TYPE-SPECIES. It appears that the use of the name Hippopleurifera by Canu & Bassler in 1925 was a valid generic introduction, and that Lagaaij and Buge were right in taking Eschara biauriculata Reuss as the type by monotypy. It is therefore
important to ascertain the characters of Reuss’s species.
Hippopleurifera biauriculata (Reuss) (Plate 1, figs. I-3).
Eschara biauriculata Reuss (part), 1848 : 66, pl. VIII, fig. 15 ; Manzoni, 1877: 11, pl. IX, fig. 209. Hippopleurifera biauriculata Reuss Canu & Bassler, 1925 : 679. (Listed.)
DISTRIBUTION. Miocene, Austria and Hungary. (Leithakalk at Eisenstadt,
_ Morbisch and Kroisbach bei Oedenburg, see bed no. 6, Reuss p. 4).
LeEcToTYPE, chosen here: 1859.L.748, Mineralien-Kabinet of Naturhistorisches Museum, Vienna. Bryozoen Sand, Eisenstadt. Specimen figured by Manzoni.
EXAMINATION OF SYNTYPES. The syntype material, kindly lent by the Natur- historisches Museum, Vienna, consists of two small, escharan fragments, together in one tube, with an original label. They belong to distinct species. One of them (Pl. 1, figs. 4, 5) is referable to Hippopleurifera sedgwicki (Milne-Edwards, see below). The other (Pl. 1, figs. 1-3) is evidently the form figured by Reuss, as it shows the
74 A. B. HASTINGS
“ biauriculate ’’ character. This specimen is now chosen as the lectotype of Eschara biauriculata.
Comparison with my PI. 1, fig. 1 will show that the detailed agreement (zooecium by zooecium, and ovicell by ovicell) leaves no possible doubt that the lectotype specimen is represented (in mirror image) in Manzoni’s figure. There are some points for criticism in his figure: (1) the scattered pores on the marginal zooecium are shown as arranged in regular rows, (2) the areola-like pores round the ovicell are exaggerated, and the frontal sculpture omitted, (3) although some variation in the size of the avicularia is shown, their shape is not satisfactorily represented.
DESCRIPTION OF LECTOTYPE (PI. I, figs. 1-3). Specimen a bilaminar fragment (5-0 mm. < 3:r mm.) including part of edge of frond. Zooecia alike on both flat surfaces and continuous over marginal surface.
Zooecia with median, suboral mucro and porous frontal wall, pores separated by smooth bars. Peripheral pores round, grading into a few oblique groove-like pores which converge on mucro (evidently a porous pleurocyst.)
Spines absent. Orifice round with a very slight demarcation of a large anter and a small poster.
Avicularia one on each side of orifice, either both small (as in Reuss’s figure) or one enlarged. Small avicularia on rim of orifice directed outwards with bluntly pointed rostrum. Large avicularia lying beside orifice and directed distally, mostly, but not all, on fertile zooecia, rostrum spatulate with straight or (in the largest avicularia) concave sides, proximal] end of avicularium raised.
Ovicells hyperstomial, their secondary covering continuous with pleurocyst of distal zooeclum. Secondary cover with two large fossae, or with irregular smaller pits and protuberances (see two ovicells side by side in PI. 1, figs. 1 and 3).
RemarRKs. H. brauriculata (Pl. 1, figs. I-3) has more in common with H. pulchra Manzoni (see Hastings, 1949, pl. XII, fig. 2) than with H. sedgwickz (see Pl. I, figs. 4, 5, and Hastings, 1949, pl. XII, fig. 1).. It resembles H. pulchva in the greater extent of the porous pleurocyst (H. sedgwicki usually having a small area without pores behind the orifice) ; in the less pronounced division of the orifice into anter and poster ; and in the details of the secondary calcification of the ovicell.
H, biauriculata differs from H. pulchva in the smoothness of the frontal wall between the pores, which are larger; and in the shape and position of the avicularia. In H. pulchva the granulations and projections on the areas between the pores are the noticeable feature of the frontal wall. Spines, which are absent in the small specimen of H. biauriculata, are conspicuous in both the other species. The three species appear to be congeneric.
A phase with two fossae is also seen in the ovicells of most of the American Early Tertiary species referred to Hippomenella by Canu & Bassler (1920 : 379 ef seq.). Brown (1949 : 517) rejected these species from Hippomenella. Hastings (1949 : 525) transferred some of them to Hippopleurifera, and Brown (1958 : 65) placed some in Trigonopora which also has fossae,
OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 75
Hippopleurifera pulchra (Manzoni) Cellepora pulcra [sic] Manzoni (Michelotti MS.), 1870 : 336, 338, pl. IV, fig. 20. Hippodiplosia granulosa Canu, 1916: 327, pl. III, figs. 7, 8. Lepralia soulieri Calv.; Waters, 1926: 429. (Development of pleurocyst.) Not Mucronella soulievi Calvet ; O’Donoghue & de Watteville, 1939: 28. (Record only.) Hippopleurifera pulchva (Manzoni) Hastings, 1949: 521, pls. XII, figs. 2-4, XIII, figs. 1, fa, 2, 2a (Synonymy); Gautier, 1962 : 189. Hippopleurifera granulosa Canu ; Vigneaux, 1949:97. (Record only.) DISTRIBUTION. Recent, Mediterranean. Miocene, Italy; Sicily; Austria ; France.
REMARKS. Hippodiplosia granulosa Canu is clearly recognizable, from Canu’s figure, as a synonym of Hippopleurifera pulchra. It is one of the two genosyntypes of Hippodiplosia Canu (1916 : 326). As Hastings (1930 : 724) chose the other geno- syntype as genolectotype, the status of Hippodiplosia is not affected by this synonymy.
The specimen from Alexandria (1963.8.2.27), recorded by O’Donoghue & de Watteville as Mucronella souliert, is referable to Schizoporella errata (Waters), for which see Gautier (1962 : 149).
Hippopleurifera sedgwicki (Milne-Edwards)
(BIE 1, tes-A.. 5) Eschara sedgwickt Milne-Edwards, 1836 : 330 (sep. p. 10), pl. Io, figs. 5, 5a. Eschara biauriculata Reuss (part), 1848: 66. (Not pl. VIII, fig. 15.) Hippopleurifera sedgwicki (Milne-Edwards) ; Buge, 1957 : 265. (Synonymy.) DISTRIBUTION. Pliocene, Europe and North Africa (see Buge). Mzocene, Austria.
REMARKS. The paralectotype of Eschara biauriculata, referred above to Hippo- pleurifera sedgwicki, is from Eisenstadt, and extends the known range of H. sedgwicki, both geologically and geographically.
Buge redescribed this species.
5. ACKNOWLEDGEMENTS
I am grateful to the Manchester Museum, the Naturhistorisches Museum, Vienna, and the Riksmuseum, Stockholm, for lending specimens discussed in this paper ; to Professor D. A. Brown who, on his own initiative, borrowed the type- material of Eschera biauriculata, and brought it to me from Vienna; to Mr. M. G. Sawyers for the photographs; to Miss P. L. Cook for her assistance, including drawings, measurements and the compilation of the list of references ; and to my husband, Dr. H. Dighton Thomas, for his encouragement and counsel.
6 SUMMARY Examination of type-material of a number of species has yielded information on their taxonomic status, distribution, structure, etc. The type-species of the genus Hippopleurifera is included, and one of the Siboga species of Sclerodomus is recon- sidered. Membranipora hexagona Busk was based on a single, mutilated specimen of an unidentifiable, non-membraniporine species.
76 A. B. HASTINGS
7. REFERENCES
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PLATE 1 Fics. 1-3. Hippopleurifera biauriculata (Reuss).
Fics. 1, 2. The two faces of the bilaminar lectotype. Fig. 1. The face figured by Manzoni, showing the edge of the frond (left) and stages in the secondary calcification of the ovicells.
Fic. 3. Part of fig. 1 enlarged, showing two biauriculate fertile zooecia, and two with one of the avicularia large and spatulate ; variation in older, secondarily calcified ovicells ; and parts of the outwardly facing zooecia at the edge of the frond.
Fics. 4, 5. Hippopleurifera sedgwickt (Milne-Edwards). The two faces of the bilaminar paralectotype of H. biauriculata (Reuss.).
ISS Ts 245 leaks Rig. igs ~xX 36.
.) Zool. 14, 3
Bull. B.M. (N.
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- WILLIAM G. INGLIS
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a HISTORY) Vol. 14 No. 4 SOE
Te <etisH “MUSEaS vx & SN
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BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 14 No. 4 LONDON: 1966
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MARINE NEMATODES FROM DURBAN, SOUTH. APRICA
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SYNOPSIS
Ten new species and three new genera of free-living marine nematodes are described from the Durban area of the Republic of South Africa, thus: Platycoma sudafricana sp. nov. is charac- terized by an arrow-head-shaped flap over the amphid and, possibly, by some features of the male tail; Tvileptiwm longisetosum sp. nov. is similar to T. ayum Inglis, 1964 but differs in the distribution of the cervical setae and the form of the spicules; Mesacanthion frica sp. nov. is characterized by the form of the spicules which have long ventro-posteriorly directed processes on their anterior ends ; Epacanthion oliffi sp. nov. is a short spiculed form characterized by lacking
_ a gubernaculum and pre-cloacal supplement and having many cervical setae in the male ; Rhab- | dodemania dura sp. nov. is characterized by a lack of ventro-lateral onchia and the shape of the | spicules; PHERONOUS ogdeni gen. et sp. nov. of the Ironidae is characterized by a lack of
caudal glands and papilla-like cephalic sense organs; Metoncholaimus murphyi sp. nov. is characterized by having one pore to the demanian system, extremely long spicules and a few long circum-cloacal setae; WIESONCHOLAIMUS mawsonae gen. et sp. nov. of the Oncholaimidae is characterized by long, equal, ventro-lateral onchia and a short dorsal onchium, a long stout tail, very long slim spicules and a large gubernaculum, and the presence of a demanian system ; PLECTOLAIMUS juliani gen. et sp. nov. of the Leptolaimidae is characterized by six distinct lips, ventral onchia-like structures at the anterior end of the oesophagus and a distinct bulb-like
i region with tiny valves at the posterior end of the oesophagus ; Bathylaimus deconincki sp. nov. _ is characterized by a large relatively posterior amphid and by the shape of the spicules and ' gubernaculum. Hyalacanthion Wieser, 1959 is a synonym of Epacanthion Wieser, 1953 because
E. butschlit (Southern, 1914), type species of Epacanthion, has typical Hyalacanthion mandibles. As a further consequence it is argued that subdivision of the families of the Enoploidea is not
_ yet possible, in spite of De Coninck’s recent attempt, particularly as De Coninck refers Hyala-
canthion to one subfamily and Epacanthion to another.
INTRODUCTION
THREE samples of free-living marine nematodes were sent to me by Dr. W. D. Oliff,
- South African Council for Scientific and Industrial Research, National Institute for
Water Research. These samples contain ten species, all of which are described below as new, referable to ten genera, of which three are new. As usual only those species represented by males in good condition are described. The species were collected from the following localities :
Beach sand in the surf zone from the Durban area on 10. viii.1g64. (Collection number: B8n8.)
I 3. Bathylaimus deconincki sp. nov. 25,99. Epacanthion oliffi sp. nov. ZOOL. 14, 4 a
82 WILLIAM.G.-INGLIS
1g. Mesacanthion frica sp. nov. 34,12. Metoncholaimus murphyi sp. nov. 12 g, 169, 4 larvae. Pheronous ogdeni sp. et gen. nov.
2 5. Platycoma sudafricana sp. nov. 1g. Plectolaimus julian sp. et gen. nov. 34,19, 12 larvae. Trileptium longisetosum sp. nov.
Sediments in the mouth of Durban Harbour (polluted) on 20.vi.1964. (M23n4.)
11 g,69, larva. Rhabdodemamia dura sp. nov. 6 3,122. Wriesoncholaimus mawsonae sp. et gen. nov.
Irom the same locality as M23n4 on same date (M23n1ITI).
2 g, 11 2, 8 larvae. Weesoncholaimus mawsonae sp. et gen. nov.
SYSTEMATIC CONSIDERATIONS
De Coninck (1965) has made a heroic attempt to provide a more reasonable classi- fication of the free-living nematodes in which he divides the Superfamily Enoploidea into six families and fifteen subfamilies. Although some of these divisions appear to be reasonable others are certainly not at present acceptable. For example he still recognizes Thoracostomopsidae as a distinct family with only one genus Thoracosto- mopsis. But this genus is only a somewhat modified Enoplidae which is well within the ranges of variation shown by the genera currently referred to that family. Further, De Coninck refers Hyalacanthion Wieser, 1959 to one subfamily of the Enoplidae (Enoploidinae) and refers Epacanthion Wieser, 1953 to another (Enoplo- laiminae). But I show below that the genera cannot be separated.
I argued recently (Inglis, 1964*) that Enoplus is a highly evolved form, while De Coninck treats 1t as a form ancestral to all the genera he refers to the family Enoplidae, but gives no indication of the origins of Enoplus itself. I have further argued, and still do, that the Enoplidae can be considered as having arisen from the genera commonly referred to the Family Phanodermatidae. In the latter family buccal rods appear to be commonly present and it is easy to imagine how, by a process of increased sclerotization, the region of the inner surface of the lip between these rods could become thickened to produce the mandibles so characteristic of the Enophdae. Within the Enoplidae several lines can be crudely recognized along each of which the mandibular : onchial complex has been modified in a different way. The problem is not that of interpreting the components of each line but simply that we do not have enough information to make such an attempt worth while. We can only await the
collection of more data and until then I still propose to recognize no groups other than Families and Genera.
* In this paper read cephalic ventricle for cephalic vesicle throughout.
MARINE NEMATODES FROM DURBAN 83 DESCRIPTIVE