ae ee T feet \1S . 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*, 50 Pee ia Pe Wie Ee ACD) REFERENCES Anon. 1904. 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Fische. Wien: 433 pp. LacEPEDE, 1803. AHistoive naturelle des poissons, 5. Paris, i-lxviii, 803 pp. Liane, Y-S. 1948. Notes on a collection of fishes from the Pescadores Islands, Taiwan. Quart. J. Taiwan Mus. 1 (2) : 1-20. LinnarEus, K. 1754. Chinensia Lagerstvémiana (dissertatio Amoen. Acad.) : 1-36. 1758. Systema Naturae, ed. 10 : 1-824. 1759. Chinensia LagerstroOmiana. Amoenitates Academiae, 4 (61) : 230-260. Liu, F. H. & SHEN, S.C. 1957. A preliminary report on the activity of Wen-fishes (Herring- like fishes) along the coast of Taiwan. Rept. Fish. Bioi. Nat. Taiwan Mus. 1 (2): 24-32. LONNBERG, E. 1896. Linnaean type-specimens of birds, reptiles, batrachians and fishes in the Zoological Museum of the University in Uppsala. Bihang K. Svenska Vet.-Akad. Handlingar. 22 (4) : 3-45. MartsuBara, K. 1955. Fish morphology and hievachy. Pt. 1. Ishizaki, Tokyo: 605 pp. Mort, T. 1952. Check list of the fishes of Korea. Mem. Hyogo Univ. Agric. 1 (3) : 1-228. Nicuots, T. 1943. The freshwater fishes of China. Nat. Hist. Central Asia, 9 : 1-322. Norman, J. R. 1923. A revision of the clupeid fishes of the genus Jlisha and allied genera. Ann. Mag. nat. Hist. (9) 11: 1-22. OsBEcCK, P. 1757. Dagbok Gfver en Ostindisk Resa aven 1750-52. Stockholm: 376 pp. Reean, C. T. 1917a. A revision of the clupeid fishes of the genera Pomolobus, Brevoortia and Dorosoma, and their allies. Ann. Mag. nat. Hist. (8) 19 : 297-316. ——1917b. A revision of the clupeid fishes of the genera Sardinella, Harengula, etc. Ibid. (8) 19 : 377-395. RICHARDSON, J. 1842. Contributions to the ichthyology of Australia, 1. Ibid. (1) 9: 15-31. 1843. Contributions to the ichthyology of Australia, 63. Jbid. (1) 11: 489-498. —— 1844-45. The zoology of the voyage of the Sulphur, 1: Ichthyology. Smith, Elder & Co., London : 53-150 (see Rept. Ichth. China Japan, p. 316 for dating of three ‘“ Sulphur ”’ fasciculi). —— 1845-46. The zoology of the voyage of H.M.S. Erebus & Terror: Ichthyology. E. W. Jansen, London: (see Rept. Ichth. China Japan, p. 272 for clue to dating). —— 1846. Report on the ichthyology of the seas of China and Japan (from Rept. Brit. Ass. Adr. Sci., 1845). Richard & John E. Taylor, London : 187-320. RussEL1, P. 1803. Description and figures of two-hundred fishes collected at Vizagapatam on the coast of Covomandal. W. Bulmer & Co., London, 2: 1-85. SEALE, A. 1914. Fishes of Hong Kong. Philipp. J. Sci. 9 (1) Sect. D. : 59-79. SHIPLEY, A. E. 1913. “ J” A Memoir of John Willis Clark. Smith, Elder, London. VALENCIENNES, M. A. 1847. Histoive Naturelle des Poissons, Paris, 20: 1-472. 1848. Ibid. 21 : 1-536. WHITEHEAD, P. J. P. 1962a. <A review of the Indo-Pacific gizzard shad genera Nematalosa, Clupanodon and Konosirus. Bull. Brit. Mus. nat. Hist. (Zool.) 9 (2) : 87-102. 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 ATE PL (N.H.) Zool. 2 Bull. B.M. 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 ae, 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)}. ® x ¥ 4 A o EE ee AA ee Aa. a Bull. B.M. (N.H.) Zool. 14, 2 PLATE -4 f 0 Chahelors choy soplers fie A PLATE 5 Fic. t. Chatoessus maculatus, Reeves drawing No. 109 [= Clupanodon thrissa (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 Bull. B.M. (N.H.) Zool. 14, 2 Px a “ > ie ‘ 2 Pipaparp re» an. PLATE on FiG. I. Fic. 2. Fic. 3. \ PLATE 6 Thryssa mystax, Reeves drawing No. 138 |= Thryssa mystax (Schneider). Megalops setipinnis, Reeves drawing No. 96 [= Megalops cyprinoides (Broussonet) }. Megalops curtifilis, Reeves drawing No. 136 [= Megalops cvprinotides (Broussonet) |. Bull. B.M. (N.H.) Zool. 14, 2 laf Celine Z alefee 4, ae 3 PLATE 7 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) }. : ii ‘ Bull. B.M. (N.H.) Zool. 2 M. ( ) Zool. 14, 2 PLATE 7 peek In in ee hee one ce. 2 Sh Lhe for fi £48 fABABt terre , LE L f Berpeanit tics stk, Covewar fas ty 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 BASSINDALE, R. 1940. The Invertebrate Fauna of the Southern Shores of the Severn Estuary. Proc. Bristol Nat. Soc., (4) 9, 2 : 143-201. BassLER, R. S. 1935. Bryozoa (Generum et Genotyporum Index et Bibliographia) 7 Quen- stedt, W., Fossilium Catalogus 1: Animalia. Pars LXVII: 1-229, ’s-Gravenhage. 1953. Tveatise on Invertebrate Paleontology, part G: Gi—G253, University of Kansas. BERTHELSEN, O. 1962. Cheilostome Bryozoa in the Danian Deposits of East Denmark. Danm. geol. Unders. ser. II, 83 : 3-290. 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The Pliocene Bryozoa of the Low Countries and their bearing on the marine stratigraphy of the North Sea region. Meded. Geol. Sticht. C5 (5) : 5-233. LanpsporouGH, D. 1852. A Popular History of British Zoophytes, oy Corallines, London. Lrvinsen, G.M. R. 1902. Studies on Bryozoa. Vidensk. Medd. dansk naturh. Foren. Kbh. : I-31. 1909. Morphological and Systematic Studies on the Cheilostomatous Bryozoa: 1-431, Copenhagen. MacGitiivray, P. H. 1889. Polyzoa im McCoy, F., Prodvomus of the Zoology of Victoria, Melbourne, 1879-1890. Decade XIX : 307-323. Manzont, A. 1870. Bryozoi fossiliItaliani. (4acontr.). S.B. Akad. Wiss. Wien, 61, 1 : 323- 349. 1877. Briozoi fossili del Miocene d’Austria ed Ungheria, Part II... Denkschy. K. Akad. Wiss. Math.-Nat. Cl., 37 : 1-30. Marcus, E. 1940. Mosdyr (Bryozoa eller Polyzoa). Danmarks Fauna, 46 : 1-401. 1950. Systematical remarks on the bryozoan fauna of Denmark. Vidensk. Medd. dansk naturh. Foren. Kbh., 112 : 1-34. MiLNE-Epwarps, H. 1836. Observations sur les Polypiers fossiles du genre Eschare. Ann. Sci. nat., sér. 2, 6 : 321-345. Moore, H. B. 1937. Marine Fauna of the Isle of Man. Proc. Lpool. biol. Soc., 50 : 5-293. Neviant, A. 1896. Briozoi Neozoici di alcune localita d'Italia. Part III. Boll. Soc. Rom. Stud. Zool., 5 : 102-125. 1900. Briozoi neogenici delle Calabrie. Paleontogr. ital., 6 : 115-266. 78 A. B. HASTINGS O’DonoGHuE, C. H. & DE WATTEVILLE, D. 1939. The fishery grounds near Alexandria. 20. Bryozoa. Note & Mém. Fouad I Inst. Hydvobiol. & Fish. 34: 1-58. Purcuon, R. D. 1937. An ecological study of the beach and the dock at Portishead. Pvoc. Bristol Nat. Soc., (4) 8: 311-329. Reuss, A. E. 1848. Die fossilen Polyparien des Wiener Tertiarbeckens. Haidinger’s Natur- wiss. Abhandl., II, Vienna (1847 sep.). [Vol. II was published 21 Aug. 1848, see Vol. III, ““Vorbericht ’’.] Sttmn, L. 1941. Cheilostomata Anasca (Bryozoa) collected by Prof. Sixten Bock’s expedition to Japan... Ark. zool. 33A, 12 : I-130. Smitt, F. A. 1867. Kritisk forteckning 6fver Skandinaviens Hafs-Bryozoer. 3. Ofvers Vetensk-Akad. Forh. Stockholm, 24, 5 : 279-429. 1873. Floridan Bryozoa, Part 2. K. svenska VetenskAkad. Handl., 9, 4: 3-83. THORNELY, L. R. 1905. Report on the Polyzoa...im Herdman, W. A., Rep. Pearl Oyster Fisheries, Gulf of Manaar, 4, Suppl. Rep. 26 : 107-130, London. VIGNEAUX, M. 1949. Révision des Bryozoaires néogenes du Bassin d’Aquitaine... Mém. Soc. géol. Fr. n.s. 28, mém. 60 : 1-155. Voict, E. 1959. Revision der von F.v.Hagenow 1939-50 aus der Schreibkreide von Riigen verdffentlichen Bryozoen. Geologie, 8, 25 : 1-80. 1960. Sur les différents stades de l’astogénese de certaines Bryozoaires cheilostomes. Bull. Soc. géol. Fr., sér. 7, 1, 7 : 697-704. Waters, A. W. 1888. Supplementary Report on the Polyzoa. Rep. Zool. Chall. Exp., 21, pt. 79. 1898. Observations on Membraniporidae. J. Linn. Soc. London (Zool.), 26 : 654-693. —— 1924. The Ancestrula of Membranipora pilosa, L., and of other Cheilostomatous Bryozoa. Ann. Mag. nat. Hist. (9) 14: 594-612. 1926. Ancestrulae and Frontal of Cheilostomatous Bryozoa. Jbid. (9) 17 : 425-4309. WIESEMANN, G. 1963. Untersuchungen an der Gattung Bevsselina Canu 1913 und ahnlichen Bryozoen (Maastrichtien, Danien, Montien). Mitt. min.-geol. (St.) Inst. Hamb., 32 : 5-7o. 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. » | PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING RINE NEMATODES FROM RBAN, SOUTH AFRICA - WILLIAM G. INGLIS SEUM Pe a HISTORY) Vol. 14 No. 4 SOE Te <etisH “MUSEaS vx & SN MARINE NEMATODES FROM DURBAN, SOU LEE PAFRIGA BY; WILLIAM G. INGLIS Pp. 79-106 ; 47 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 14 No. 4 LONDON: 1966 Issued 23 March, 1966 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted 1m 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 1s Vol. 14, No. 4 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 HiS TORY) Price Eleven Shillings MARINE NEMATODES FROM DURBAN, SOUTH. APRICA By WILLIAM G. INGLIS 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 SECTION Family LEPTOSOMATIDAE Platycoma sudafricana sp. nov. (Text-figs. I-6) MATERIAL STUDIED. 2,4. Beach sand in the surf zone. Near Durban, Republic of South Africa (B8n8). B.M. (N.H.), Reg. Nos. 1965.961—962. A B C Body length (mm.) 146-38 8-21 56:10 1o0'T 144°74 8°53 68°75 I'l _ MEASUREMENTS (in mm., in order of body lengths) Body breadth: 0-069; 0-076. Oecesophagus length: 1:23; 1:29. Diameter of ‘head : 0:031 ; 0-029. Length of cephalic setae (long/short) : 0-023/0-018 ; 0-023/ 0-018. Distance from anterior end of body to posterior edge of amphid: 0-026; 0-027. Length of cervical setae: 0-0r4—-0-018 ; 0-014-0-017. Distance of nerve ring from anterior end of body: 0:30; 0-29. Length of spicules: 0-163; 0-165. Length of lateral piece of gubernaculum: 0-033; 0:035. Length of apophosis of gubernaculum: 0-015; 0-or4. Distance of first pre-cloacal supplement anterior _to cloacal opening: 0-087; 0-122. Distance of second pre-cloacal supplement _ anterior to cloacal opening : not measured, tail too curved ; 0-198. Length of tail : | 0-18; 0-16. Cloacal diameter: 0-066; 0-060. The anterior end of the body is set off as a constricted region starting about the “level of the posterior edge of the amphids (Text-fig. 1). The mouth opening is simple, _tri-radiate and leads almost immediately into the oesophagus which carries three _ small wholly cuticular onchia on each sector (Text-fig. 1). The cephalic sense organs _ lie in two circles of which the inner consists of six prominent papillae while the outer consists of ten setae of which six are long and four are shorter (Text-fig. 1). There _ are many long cervical setae. The amphids are prominent with a large arrow-head- like flap of cuticle developed from their anterior edge (Text-figs. r and 4). The oesophagus is long and simple in outline. The tail bears a pair of prominent spike-like setae about half-way along its length and a long but slim seta just posterior to the cloacal opening (Text-fig. 2). There is no spinerette and there are no caudal glands. On the mid-ventral surface of the _ body anterior to the cloacal opening there are two pre-cloacal supplements (Text-fig. 2) of which the more posterior (Text-fig. 6) is a rounded cap-like structure resembling a Type-I campaniform organ (Inglis, 1963) while the more anterior (Text-fig. 5) is a raised area on which are located three pairs of short, stoutish setae. The spicular : gubernacular complex is double. That is, all the structures shown in the figure (Text-fig. 3) are represented by exactly equivalent structures on the other side of the body and they are in no way connected across the mid-line of the body. Further, the entire complex appears to form one functional unit. Here I | interpret the structures as Gerlach (1955) does and not as he re-interprets them later 84 WILLIAM-G.INGLIS (1959). That is I consider the largest L-shaped structure to be the spicule which is enfolded posteriorly by a slightly triangular-shaped gubernacular apophosis. The most interesting feature is the relationship between the spicule and the rod-like structure bordering it anteriorly (considered a spicule by Gerlach (1959)). This rod swells towards its distal end where it bears a lateral flange which enfolds the distal Fics. 1-6. Platycoma sudafricana sp. nov. Fig. 1. Lateral view of head. Fig. 2. General lateral view of male tail. Fig. 3. Detail of spicular : gubernacular complex from right side. Note the articulation between the spicule and the anterior part of the gubernaculum. Fig. 4. Variant form of amphid. Fig. 5. Ventral detail of more anterior pre-cloacal supplement. Fig. 6. Ventral detail of more posterior pre-cloacal supplement. end of the spicule while the same rod swells to form a slight knob proximally which fits into a socket in the spicule. Discussion This genus appears to be characterized by the peculiar modification of the spicular apparatus, the presence of pre-cloacal supplements, the presence of the pair of stout, spine-like setae on the tail and in lacking caudal glands. In addition the head is highly characteristic with three small onchia and the marked sexual dimorphism of the amphids. In this respect the present species resembles P. africanus (Gerlach, ea 4 | | MARINE NEMATODES FROM DURBAN 85 1959) but differs from it in the arrow-head-like form of the amphidial flap. P. sudafricana is also, possibly, characterized by the number and form of the pre-cloacal supplements and the form of the spicular : gubernacular complex. Platycoma is very similar to Platycomopsts Ditlevsen, 1926 but this latter genus does not show any sexual dimorphism of the male amphids, and the male apparatus is more typical with long spicules and a short anterior part to the gubernaculum. This does, however, appear to represent a simpler version of the apparatus found in Platycoma and is one of the reasons for the interpretation I advance above. In addition Platycomopsis does not have the pair of spine-like setae on the male tail which is such a characteristic feature of the species referred to Platycoma. Trileptium longisetosum sp. nov. (Text-figs. 7-9) MATERIAL STUDIED. 3 4, I 9, 12 larvae. Beach sand in the surf zone. Near Durban, Republic of South Africa (B8n8). B.M. (N.H.) Reg. Nos. 1965.963-972. Body length A B Cc Vv (mm.) Males . ; 3 : 10°64 5°63 22:69 — 4°68 10°65 5:10 24°50 — 4°90 Female : : F 8-10 6-51 22°05 52-90 2°07 Larva . : : . 8-54 5-09 ZANATZ — 4°10 MEASUREMENTS (in mm. in order of body lengths) Mates: Body breadth: 0-044; 0-046. Oesophagus length: 0-83; 0-96. - Diameter of head: 0-017; 0-018. Length of cephalic capsule: 0-036; 0-034. Diameter of cephalic capsule at posterior end: 0-019; 0-022. Length of longest cephalic setae: 0-040; 0-039. Length of cervical setae: 0-018-0-020 ;_ broken. Distance of nerve ring from anterior end of body: 0-19; 0-21. Length of spicules : 0°047; 0:052. Length of gubernaculum: 0-018; 0-016. Length of pre-cloacal supplement : 0:0065; 0:0065. Distance of pre-cloacal supplement anterior to cloacal opening: 0-038; 0-040. Length of tail: 0-21; 0-20. Cloacal diameter : 0:036 ; 0-040. FEMALE: Body breadth: 0-049. Oesophagus length: 0-61. Diameter of head : 0-020. Length of cephalic capsule : 0-022. Diameter of cephalic capsule at posterior end: 0-025. Length of longest cephalic setae: 0-066. Length of cervical setae : 0-008. Distance of nerve ring from anterior end of body: 0-16. Length of tail : 0-18. Anal diameter: 0-036. Distance of vulva from anterior end of body: 2:1. Larva: Body breadth: 0-048. Oesophagus length: 0-72. Diameter of head: 0-020. Length of cephalic capsule : 0-023. Diameter of cephalic capsule at posterior end: 0-021. Length of longest cephalic setae : 0-042. Nocervicalsetae. Distance of nerve ring from anterior end of body: o-1g. Length of tail: 0-17. Anal dia- | Meter: 0-043. ZOOL. 14, 4 78$§ 86 WILLIAM G. INGLIS Fics. 7-9. Trileptium longisetosum sp. nov. Fig. 7. Ventro-lateral view of head. Fig. 8. Lateral view of male tail. Fig. 9. Spicule and gubernaculum from the left. Figs. 10-12. Mesacanthion frica sp. nov. Fig. 10. Dorsal view of head. The cephalic setae are not drawn in full because of their great length. Fig. rr. Lateral view of male tail. Fig. 12. Detail of spicule and gubernaculum (?) from the left. : MARINE NEMATODES FROM DURBAN 87 The structure of the head is the same as that in Tvileptiwm ayum Inglis, 1964 with three small onchia enclosed by three small mandibles. The setae of the inner circle of cephalic sense organs are very stout and prominent and the pouch of the cephalic slit is rounded and very prominent (Text-fig. 7). The “‘cephalic setae’ are numerous and consist of six very long setae followed by two shorter setae on the dorso- and ventro-lateral positions and by one on the lateral positions. In the males there is one circle of ten cervical setae (Text-fig. 7) while there are four, much shorter setae in the equivalent position in the females. In addition there are a number of long, thin setae on the anterior end of the body which become sparser posterior to the nerve ring. Such setae measure 0-009—0-013 mm. in length. The tail is long and narrow in both sexes and a very small rod-like pre-cloacal supplement is present on the ventral surface of the male anterior to the cloacal opening (Text-fig. 8). The spicules are equal and identical with slightly cephalate proximal ends and bluntly rounded distal tips. The gubernaculum is simple and lies very close to the _ spicules (Text-fig. 9). Discussion This species is most similar to 7. ayum Inglis, 1964 but differs from it in the number and distribution of the cervical setae and the short spicules. It appears to differ most markedly from all the other known species in having three onchia and a pre- cloacal supplement in the male. Mesacanthion frica sp. nov. (Text-figs. 10-12) MATERIAL STUDIED. I g. Sand in surf zone. Near Durban. Republic of South Africa (B8n8). B.M. (N.H.) Reg. No. 1965.1o1r1. A B C Body length (mm.) 42°31 3775 0°07 1705 _ MEASUREMENTS (in mm.) Body breadth: 0-039. Oesophagus length: 0-44. Length of cephalic setae (anterior/longer posterior/shorter posterior) : 0-013/0-:096/0-051. Depth of cephalic capsule (?) : 0-026. Diameter of cephalic capsule at posterior edge: 0-026. Dis- tance of nerve ring from anterior end of body: 0-139. Length of body setae about _ level of nerve ring: o-o16. Length of spicules: 0-040. Length of pre-cloacal supplement : 0-012. Distance of pre-cloacal supplement anterior to cloacal opening : 0-078. Length of tail: 0-182. Cloacal diameter: 0-036. The single specimen is in a rather poor condition so that some details cannot be reliably assessed. Nevertheless sufficient detail can be made out to warrant its description in view of the peculiar features it shows. The cephalic sense organs are 88 WILLIAM G. INGLIS very long and those of the posterior circle appear to lie about the posterior edge of the cephalic capsule. This is, however, uncertain and is suggested largely on the basis of the shape of the anterior end of the body (see Text-fig. 10). The mandibles are prominent but fairly lightly developed and show no indication of any mandibular rods as in the species I refer elsewhere to Mesacanthion (Inglis, 1964). In addition, the shape of the anterior edge of the mandibles is different resembling the conditions in Africanthion in that their ends are directed towards the inter-labial spaces rather than inwards towards the mandibular cavity. The onchial plate is very poorly developed with the onchia projecting just anterior to the mandibular ring. The tail is long and slim and a small, rod-like, pre-cloacal supplement is present (Text-fig. 11). The most remarkable feature of the specimen is the form of the spicules (Text-fig. 12), which possess a long narrow ventrally-posteriorly directed process on their anterior ends; are in addition stout with a lateral bump near their posterior ends and appear to have a median central ridge running their full length. There is (?) a small gubernaculum, although its presence is uncertain. Discussion This species is characterized by the form of the spicules with the massive process on their anterior ends. Otherwise the spicules resemble those in Enoplolaimus crassidens Ditlevsen, 1930. Epacanthion oliffi sp. nov. (Text-figs. 13-16) MATERIAL STUDIED. 2,9. Beachsandin surf zone. Near Durban, Republic of South Africa (B8n8). B.M. (N.H.) Reg. Nos. 1965.1004-1010. Body length A B C V (mm.) Males 34°81 3:19 10°33 — 1-88 38°36 3°LO 10-14 — ZANE Females 25°34 3°49 T1-64 58:92 1-85 B207, 2°88 12-10 61°73 I-96 32°58 3°16 8-98 58-42 2-02 MEASUREMENTS (in mm., in order of body lengths). Mates: Body breadth: 0:054; 0-055. Oecsophagus length: 0:59; 0-64. Diameter of cephalic capsule at posterior end: 0-034; 0-041. Depth of cephalic capsule : 0:033 ; 0:039. Length of cephalic setae (anterior/long posterior) : 0-015/ 0:036 ; 0:016/0-039. Length of cervical setae : in range 0-035—0-:039 on both speci- mens. Distance of nerve ring from anterior end of body: 0-131; o-r4o. Length of spicules: 0:028; 0-029. Length of tail: 0-182; 0-208. Cloacal diameter : 0-044; 0:046. Length of setae on tail and on general body surface : up to 0-039 mm, FEMALES: Body breadth: 0-073; 0-058; 0-062. Oesophagus length: 0°53 5 | | MARINE NEMATODES FROM DURBAN 89 0:68 ; 0-64. Diameter of cephalic capsule at posterior end: 0-038 ; 0-039; 0-038. Depth of cephalic capsule: 0-039; 0-041; 0-042. Length of cephalic setae (anterior/long posterior/short posterior): 0-017/0-:034/0-015 ; 0:016/0-036/0-016 ; 0:017/0:033/0-015. Length of cervical setae: 0°032; 0°028; 0°035. Distance of nerve ring from anterior end of body: 0-162; 0-138; 0-165. Length of tail: 0-159; 0-162; 0-225. Anal diameter: 0-048; 0-044; 0-052. Distance of vulva from anterior end of body: I-09; 1-21; 1-18. The head is typically enoplid with three high, slim lips bearing distinct semi-lunar striations lying on an area bounded anteriorly by a curved line running posteriorly from the tips of the mandibles (Text-fig. 13). The major part of the remainder of the lips is also marked by striations which are much further apart. These conditions are exactly as described for Enoploides species (see Inglis, 1964, fig. 30). The mandibles are typical of the genus with what appear to be two parallel rods joined by a thin sheet of cuticle. As usual this is due to studying a curved solid structure in optical section. The anterior ends of the mandibles form well developed projecting hooks. The three equal onchia are rather small and lie posterior to the posterior edge of the mandibles and do not lie close within them as in Hyalacanthion multi- papillatum Wieser, 1959. The cephalic setae are long and slim in both sexes with the more anterior pairs lying about the anterior end of the mandibles. Squarish blocks of specialized cuticle developed from the outer cuticle of head are closely associated with them (Text-fig. 13). In the females the outer setae are represented by six long setae and four short but in the males there are groups of three dorso- and ventro-laterally and two pairs laterally. In the females there are four long cervical setae but in the male the region of the body posterior to the cephalic capsule carries a very large number of long setae (Text-fig. 15). Setae become scarcer on the body posterior to about the level of the nerve ring but long setae occur sporadically over the entire length of the body. The tail is somewhat blunt and stout in both sexes (Text-fig. 14). In the male there is no pre-cloacal supplement or gubernaculum. The spicules are short and stout with a series of serrations on their posterior ends (Text-fig. 16). Discussion Wieser (1959) erects a new genus Hyalacanthion for three species, H. multipapilla- tum Wieser, 1959 (type species) ; H. pellucidus (Savaljev, 1912) and H. murmanicus (Savaljev, 1912). This genus, as he points out, is very similar to Epacanthion ; the major differences between the two being that the mandibles in Epacanthion consist of two plates, separated by a lacuna, which are joined anteriorly by a narrow rod and the spicules are at least 2:5 anal diameters long while the mandibles in Hyalacanthion are plates joined by a thin transparent lamella and the long spicules are never less than 2:5 anal diameters long. However Mawson (1g58a) describes a new species, Epacanthion brevispiculosum, in which the spicules are considerably less than 2-5 anal diameters in length, and the species described above also has very short spicules, I do not think that Epacanthion 90 WILLIAM G. INGLIS Figs. 13-16. Epacanthion oliffi sp. nov. Fig. 13. Dorsal view of female head. Fig. 14. Lateral view of male tail. Fig. 15. Dorsal view of anterior end of male body. Fig. 16. — Detail of spicule from the left. i 1 | iy MARINE NEMATODES FROM DURBAN 91 can be separated from Hyalacanthion but the question is simplified by a study of the type male of Enoplus butschlii Southern, 1914 which Wieser (1953) selects as type species of Epacanthion. The structure of the lips and mandibles of this species is exactly the same as that described above for E. oliffi. The same semi-lunar striations are present on the lips, the same masses of dense cuticle on the outer surface of the lips, the onchia are in the same positions relative to the mandibles (that is, posterior to them) and the mandibles have exactly the same appearance. There is no indica- tion of a distinct bar joining the so-called lateral bars of the mandibles as is illustrated by Wieser (1953) and by Southern (1914). As a result Hyalacanthion must fall as a synonym of Epacanthion and I can see no reason to recognize two genera for the species previously grouped in the two genera. I am quite sure that the differences reported in the structure of the man- dibles simply reflect the condition of the specimens when they were studied. But if later careful study shows a need for two genera a new name will be required. Hyalacanthion multipapillatum Wieser, 1959 is, therefore referred to the genus Epacanthion as a new combination. E. oliffi is distinct from all the species with short spicules in lacking a gubernaculum and a pre-cloacal supplement and in the presence of such a large number of cervical setae. The species most similar to it appears to be E. multipapillatum (Wieser, 1959) in which the spicules are short and serrated but this species possesses a gubernaculum and a series of pre-cloacal supplements. Rhandodemania dura sp. nov. (Text-figs. 23-25) MATERIAL STUDIED. II 3, 6 9, r larva. Polluted sediments in the mouth of Durban Harbour, Republic of South Africa (M23n4, collected 20.vi.1964). B.M. (N.H.) Reg. Nos. 1965 .994-1003. Body length A B Cc Vv (mm.) Males . ; ; : 2:02 7:01 24°38 — 3217 30°38 6°92 20°77 _ 3°22 33°57 6-97 24°19 =e 32 36-22 6-93 22°90 — 3°55 Females ‘ ; : 34°32 F377 222770 54°30 3°02 34°40 (jae 26-26 57°56 3°44 2°84 7°09 23°87 53°63 3°58 40°32 7°01 23°01 54°67 3°79 Larva . : 5 : 35°00 4°93 15°98 -— 1°47 MEASUREMENTS (in mm., in order of body lengths) MALEs: Body breadth: 0-099; 0-106; 0-098; 0-098. Oecsophagus length : 0-452; 0:465; 0:472; 0-512. Diameter of head: 0-016; 0-018; 0-017; 0-018. Length of cephalic setae (long/short) : 0-0084/0-0063 ; 0:0083/0:0069 ; 0:0085/ 92 WILLIAM G. INGLIS 0:0064 ; 0:0087/0-:0065. Depth of buccal cavity: 0-020; 0-017; 0-019; 0-018. Length of spicules: 0:055; 0:045; 0°:052; 0-048. Length of gubernaculum : 0°020;; - 0°026'; 0:028.5-"0:031~ Length of tail 2: 0-130; 0°155 ;40°130%, lOsm5e Cloacal diameter : 0:058 ; 0:058 ; 0:057; 0:056. FEMALES: Body breadth: 0-088; 0-100; 0-109; 0-093. Oesophagus length : 0°474; 0°458; 0:505; 0°535. Diameter of head: 0-018; 0-016; 0-017; 0-0IQ. Length of cephalic setae (long/short) : 0-0082/0-0063 ; 0:0085/0-0062 ; 0-0086/ 0:0061 ; 0:0084/0:0065. Depth of buccal cavity: 0-021; 0-022; 0:022; 0-020. Length of tail: 0-133; 0-131; 0-150; 0-163. Anal diameter: 0-057; 0-066; 0-065 ; 0:054. Distance of vulva from anterior end of body: 1-64; 1:98; 1-92; 2:05. Size of eggs: 0:045 X 0-165; 0-04I X 0°142; 0:054 X 0-123 (as examples). Larva: Body breadth: 0-042. Oesophagus length: 0-298. Diameter of head: o-o12. Length of cephalic setae (long/short) : 0:0064/0-0037. Depth of buccal cavity: 0-016. Length of tail: 0-092. Anal diameter: 0-028. The species is typical of the genus with the usual three pairs of long tooth-like structures which represent modified mandibles (see Inglis, 1964) and with the lip region of the head marked by striations (Text-fig. 23). There is a fairly prominent dorsal onchium and no ventro-lateral onchia. The tail is bluntly rounded with a series of long stout setae, in the male (Text-fig. 25). The spicules are slim with prominent caps on their proximal ends and are doubled at their distal ends (Text-fig. 24). They are roughly the same width along the whole of their lengths when viewed from the lateral aspect but have a ridge developed on their inner surfaces which can only be easily seen when the specimen being studied is viewed from a slightly ventral aspect. The gubernaculum is relatively long and slim with a very slight expansion towards the proximal end (Text-fig. 24). Discussion This species is characterized by the presence of only a dorsal onchium, by the slim shape of the spicules and gubernaculum, and by the cephalic setae being arranged in one circle only. In many ways it is similar to R. nancyae Inglis, 1964 which is also from South African waters but the shape of the spicules and gubernaculum is totally different while ventro-lateral onchia are totally lacking, although very reduced in R. nancyae, Family IRONIDAE The next species to be described differs from all others referred to the family in so many characters as to warrant reference to a new genus, thus : PHERONOUS gen. noy. Ironidae : cephalic sense organs all papillose ; two dorsal onchia and one on each ventro-lateral sector of the oesophagus ; no caudal glands. TYPE SPECIES : Pheronous ogdent sp. nov. MARINE NEMATODES FROM DURBAN PHERONOUS OGDENI gen. et. sp. nov. (Text-figs. 17-22) MATERIAL STUDIED. 12 4, 16 9, 4 larvae. Durban, Republic of South Africa (B8n8). Males . : : j 24 34° 33° 38 Females ; : : 27 390 28 31 34° A 03 71 58 83 50 85 ‘81 °45 04 *43 “48 93; Beach sand in the surf zone. Near B.M. (N.H.) Reg. Nos. 1965.974-993. Body length Cc V (mm.) 22°24 — 1*49 26-03 —— L777 22°82 1°98 25°38 — 1°98 T4°59 52°45 1°43 14°36 57°93 T°45 13°94 56-58 I°52 16:02 54°76 173 15°13 53°67 1°77 _ MEASUREMENTS (in mm., in order of body lengths) Males: Body breadth: 0-062; 0-051 ; 0:053; 0:051. Ocsophagus length : O-20%= 0°300 ; 0:330; 0-317. Diameter of head: 0-031; 0:023; 0-026; 0:023. Length of pharyngeal rods: 0:056; 0:062; 0:058; 0:057. Distance of nerve ring , from anterior end of body: 0-128; 0-147; 0-140; 0-168. Length of 0°047 ; 0:041; 0:039; 0-035. Length of gubernaculum: 0-016; 0-019; 0-018 ; FEMALES: Body breadth: 0:052; 0:047; 0:0543; 0:055; 0:052. i 0-020. Length of tail: 0-033; 0-032; 0:033; 0:033. - 0:068 ; 0:076; 0-078. imlengthi= 0-268 ; 0-291; 0:251; 0:269; 0-273. 0:027 ; 0:025; 0-024. Length of pharyngeal rods: 0-055; 0-059; 0:057; 0:054; 0:056. Distance of nerve ring from anterior end of body: 0-121; 0-140; 0-123; Gaye 102124. Length of tail: 0-098; 0-101; 0-109; 0°093; 0:123. MIetel = 07020 ; 0-029 ; 0:032; 0-033; 0-028. of body: 0-75; 0°84; 0:86; 0:93; 0°95. spicules : Cloacal diameter: 0-067 ; Oesophagus Diameter of head: 0-023; 0-019; Anal dia- Distance of vulva from anterior end The head is distinctly set-off from the remainder of the body by a constriction about the level at which the prominent oval shaped amphids open by narrow slits (Text-figs. 17 and 18). The mouth opening is bounded by three lip-lobes and is The outer cephalic sense organs form one circle surrounded by six small papillae. _ of stout papillae of which six are more prominent than the remaining four (Text-figs. 17,18 and1g). The buccal cavity is lined by a large number of small wholly cuticular denticles and the anterior end of the oesophagus bears a series of wholly cuticular onchia. There is a pair of small onchia on the dorsal sector of the oesophagus and a single larger onchium on each ventro-lateral sector (Text-fig. 20). The anterior end of the oesophagus is modified as a distinct oesophastomal region in which the 94 WILLIAM G. INGLIS Figs. 17-22. PHERONOUS ogdeni gen. et sp. nov. Fig. 17. Lateral view of head and oesophastomal region of body. Fig. 18. Dorsal view of head. Note double onchia. Fig. 10. | En face view of head. Fig. 20. Sketch of onchia en face. Fig. 21. Detail of spicule and — gubernaculum from the right. Fig. 22. Lateral view of male tail. : Fics. 23-25. Rhabdodemania dura sp. nov. Fig. 23. Lateral view of head with the dorsal — surface to the right. Fig. 24. Detail of spicule and gubernaculum from the right. Fig. 25. Lateral view of male tail, : MARINE NEMATODES FROM DURBAN 95 - musculature runs slightly anteriorly (Text-fig. 17) and in which the cuticular lining is very prominent. The slim oesophagus expands evenly towards its posterior end. The tail tapers to a sharp point and there are no caudal glands. In the male the tail bears a series of three pairs of small ventro-lateral papillae and a series of four pairs of similar papillae anterior to the cloacal opening (Text-fig. 22). The spicules are stout and clumsy in appearance without any particularly characteristic features although a dorsal strengthened rib sometimes appears to be present. The guber- naculum is small and enfolds the distal ends of the spicules (Text-fig. 21). — Discussion This species is unusual in lacking caudal glands, otherwise it would be referable _ to the genus Dolicholaimus. It is possible that it also differs from the species gener- ally referred to that genus in possessing a pair of small dorsal onchia but this is still - uncertain (see Inglis, 1961). Family ONCHOLAIMIDAE Metoncholaimus murphyi sp. nov. (Text-figs. 26-32) | MATERIAL STUDIED. 3 9,19. Beach sandin surf zone. Near Durban, Republic | of South Africa (B8n8). B.M. (N.H.), Reg. Nos. 1965.1012—-ror5. Body length A B Cc V (mm.) Males . : : é 61°05 5°31 62°71 _ 2°32 64°30 5°86 60°05 — 2:50 61°05 5°31 62-71 — 2232, Female. : : F 52°31 5°55 59°13 70:+22 2°72 | MEASUREMENTS (in mm., in order of body lengths) Mates: Body breadth: 0:038; 0:039; 0-038. Oecesophagus length: 0-437 ; | 0:428 ; 0-419. Diameter of head: 0:027; 0-024; 0:026. Length of cephalic setae _ (long/short) : 0-012/0-008 ; 0-orr/0-009; 0-012/0-008. Buccal cavity (greatest | length/greatest width): 0-035/0-:017; 0:032/0-018 ; 0:035/o-019. Distance of | nerve ring from anterior end of body: o-18r; 0-179; 0-198. Length of spicules : 0:082 ; 0:069; 0-077. Length of tail: 0:037; 0-038; 0-043. Cloacal diameter : | 0-029; 0-022; 0-026. Lengths of circum-cloacal setae (p/ar/a2/a3) : 0°012/0-017/ 0:013/0-007 ; 0:013/0-018/0-016/0-008 ; 0°016/0-021/0-017/0-009 (see Text-fig. 29 for | details). FEMALE: Body breadth: 0-052. Oesophagus length: 0-490. Diameter of head : | 0-036. Length of cephalic setae (long/short) : 0-012/0-009. Buccal cavity (greatest length/greatest width) : 0:044/0:023. Distance of nerve ring from anterior end of body: 0:22. Distance of excretory pore from anterior endof body : 0-026. Length 96 WILLIAM G. INGLIS of tail: 0-046. Anal diameter: 0-036. Distance of vulva from anterior end of body: 1-91. Distance of demanian pore anterior to posterior end of body: 0-072. Distance of uvette anterior to posterior end of body: 0-26. Distance of osmosium anterior to uvette: 0:36. Length of uterine efferent: 0-66. The head is typical with a fairly long buccal cavity with roughly parallel sides (Text-figs. 26, 27 and 28). There is an inner circle of six papillae and an outer circle of cephalic setae of which six are slightly longer than the remaining four. The left ventro-lateral onchium is considerably larger than the smaller and equal dorsal and right ventro-lateral onchia, both of which have somewhat blunt, slightly doubled anterior ends (Text-figs. 26 and 27). The amphids are prominent with slightly different degrees of development of the anterior concavity (Text-figs. 26, 27 and 31). Fics. 26-28. Metoncholaimus murphyi sp. nov., head. Fig. 26. Left-ventro-lateral view. Fig. 27. Wholly lateral view. Fig. 28. Wholly dorsal view. The male tail is short and sharply curved, almost hook-like, with a pair of long stout post-cloacal setae (p in Text-fig. 29), and two similar pairs flanking the cloacal opening, of which the more posterior is longer than the more anterior (aI, a2 in Text- fig. 29). In addition there is a further pair of shorter setae (a3 in Text-fig. 29) which appear to be slightly longer than the setae occurring on the general body surface except that, while these latter body setae vary somewhat in position from specimen to specimen, this pair (a3) is always found in the same position. The equal and identical spicules are long and slim and a small gubernaculum may be present (Text-fig. 29) but this is uncertain. There is a marked thickening of the cuticle to form a bump on the body, just anterior to the cloacal opening, into which a duct appears to open. The structure is definitely present but its details could not be established with certainty (Text-figs. 29 and 32). MARINE NEMATODES FROM DURBAN 97 The female tail is short and stout (Text-fig. 30) and the demanian system empties through a single pore which opens slightly to the left of the mid-ventral line of the body. The details of the demanian system could not be wholly established but (using the terminology of Cobb, 1930) the wvette is surrounded by leaf-like elements, the uterine efferent is long, stout and easily seen and the osmostum could be identified 0-364 mm. anterior to the wvette. The relative measurements of the various parts that could be identified are given above. Discussion This is a typical member of the genus Metoncholatmus which can be easily distin- guished by the presence of only one demanian pore in the female (otherwise known only from M. haplotretos Mawson, 1958a), by the extreme length of the spicules in association with the form of the male tail, by the small number of long circum- cloacal setae, and by the length of the cephalic setae. WIESONCHOLAIMUS Gen. Nov. The specimens described below resemble most closely those referred to the genus Metaparoncholaimus De Coninck & Schuurmans Stekhoven, 1933 in the unpaired female reproductive system, the two long, equal ventro-lateral onchia and the presence of a demanian system. It differs markedly, however, in the presence of the large gubernaculum, the form of the tail and the long, slim spicules. It is, therefore, referred to a new genus, thus: WIESONCHOLAIMUS gen. nov. Oncholaimidae : ventro-lateral onchia equal and larger than dorsal ; tail long and _ stout. MALE: spicules very long and slim ; large gubernaculum present. FEMALE : _ Demanian system present. | TYPE SPECIES: Wiesoncholaimus mawsonae sp. nov. | WIESONCHOLAIMUS MAWSONAE gen. et sp. nov. (Text-figs. 33-35) | MATERIAL STUDIED. 6 3, 12 9. Polluted sediments in the mouth of Durban | Harbour, Republic of South Africa (M23nr11). B.M. (N.H.) Reg. Nos. 1965. 1o16- 1030. (Holotype selected from this sample.) 25,11 9,8larvae. From same locality on same date at a depth of 40 feet (M23n4). B.M. (N.H.) Reg. Nos. 1965 . 1031-1040. 98 WILLIAM G. INGLIS Body length A B c AVE (mm.) Males . : - ‘ 70-16 L204 44°02 — 4°49 64°93 11-29 41°22 = 4°74 4Oi27 12:09 40°94 — 5°20 Females : : j 51-41 9°85 31-96 67°75 4°73 54°95 10°87 33°84 64°38 5711 54°76 13°76 34°81 64°72 5°64 MEASUREMENTS (in mm., in order of body lengths. All ex M23n11r) Mares: Body breadth: 0-064; 0-073; 0:074. Oesophagus length: 0:37; 0:42; 0°43. Diameter of head: 0-033; 0:030; 0-030. Length of cephalic setae (long/short) : 0-0090/0-0051 ; 0:0088/0-0053 ; 0-0091/0-0052. Dimensions of buccal cavity (length/breadth): 0-032/0-018; 0:038/0-016; 0-033/o0-018. Distance of excretory pore from anterior end of body: 0-095; 0-103; 0-096. Length of spicules: o-189; 0-211 ;|0-183- “Length of tail: 0-102; 0:115 > 40-1274. Cloacat diameter : 0:032 ; 0:032; 0°035. FEMALES: Body breadth: 0-092; 0-093; 0-103. Oecesophagus length: 0-48 ; 0°47; 0-41. Diameter of head: 0-032; 0-030; 0:030. Length of cephalic setae (long/short) : 0-:0086/0-0049 ; 0:0092/0-:0051 ; 0-:0087/0-0050. Dimensions of buccal cavity (length/breadth): 0-033/0:020; 0-038/0:020; 0:035/0-01g9. Distance of excretory pore from anterior end of body: 0-089; 0-121; 0-107. Length of tail: 0-148 ; 0-151; 0-162. Anal diameter: 0-038; 0-040; 0:036. Distance of vulva from anterior end of body: 3-11; 3:29; 3°65. Size of eggs: 0-083 x 0-0067 ; 0:097 X 0:078 ; 0:070 X 0-106 (as examples). The head bears an inner circle of six papillae surrounding the six-lobed mouth opening and an outer circle of ten setae of which six are longer than the remaining four. The amphids are smallish and he about the level of the posterior quarter of the buccal cavity. The buccal cavity contains two large, equal ventro-lateral onchia and a smaller dorsal onchium (Text-fig. 33). The tail is long and stout in both sexes (Text-fig. 34) and in the male the spicules are long, slim and equal in length. The gubernaculum is relatively massive (Text-fig. 34). There is a doubled papilla-like structure on the anterior lip of the cloacal opening in the male (Text-figs. 34 and 35) but no details of any nerves or ducts could be seen. The male tail bears a series of long, stout setae which continue anteriorly to surround the cloacal opening (Text-fig. 34). The number of these setae varies shghtly from eight to ten and they are not always wholly symmetrical. Nevertheless there is always a pair of smaller setae flanking the cloacal opening (Text-figs. 34 and 35). Inaddition there is a variable number of smaller, thorn-like setae on the surface of the tail. The demanian system is well developed in the females and opens by two dorso- lateral pores which lie 0:23 mm. anterior to the anus in a specimen 4:9 mm. long (from sample M23n4). A most characteristic feature of the system is that the wvedte, $y MARINE NEMATODES FROM DURBAN 99 29 oe ae iy, 31 ~L | 32 ni ~~ f Fics. 29-32. Metoncholaimus murphyi sp. nov. Fig. 29. Lateral view of male tail. Fig. 30. Lateral view of female tail. Fig. 31. Variant form of amphid. Fig. 32. Ventral view of male tail. (Labelling on Figs. 29 and 32 is defined in the text.) Fics. 33-35. WIESONCHOLAIMUS mawsonae gen. et sp. nov. Fig. 33. Lateral view of head with dorsal surface to the right. Fig. 34. Lateral view of male tail. Note that for convenience the whole length of the spicule is not drawn. Fig. 35. Ventral view of male tail. 100 WILLIAM G. INGLIS which lies 0-52 mm. anterior to the anus, is a simple spherical organ from which a stout uterine efferent runs anteriorly and a thin duct runs posteriorly. This single duct is very easily seen and is a distinct structure as I have established by dissection. It has not, however, been possible to establish any further details of the structure of the demanian system in spite of several attempts. Discussion See above, page 97. Family LEPTOLAIMIDAE PLECTOLAIMUS gen. nov. This species described below is clearly related to those generally referred to the family Leptolaimidae (sensu de Coninck, 1965) and is most similar to the members of the genera Chronogaster Cobb, 1913 and Halaphanolaimus Southern, 1914. It differs from them both in the form of the posterior end of the oesophagus and the modifica- tion of the anterior end of the oesophagus (in which it is convergent with the Rhabdi- toid genera Odontorhabditis Timm, 1959 and Cheilorhabditis Timm, 1959). In addition it appears to differ in the presence of very distinct lips. The genus may be diagnosed thus : PLECTOLAIMUS gen. nov. Leptolaimidae : mouth bounded by six distinct lips; onchia-like structures developed at anterior end of oesophagus from ventro-lateral sectors; post- oesophageal bulb-like region present with small tri-valvulate structure ; pre-cloacal rod-like supplements present on ventral surface of male. Type species: Plectolarmus juliani sp. nov. PLECTOLAIMUS JULIANI gen. et sp. nov. (Text-figs. 36-43) MATERIAL STUDIED. TI 3. Beach sand in surf zone. Near Durban, Republic of South Africa (B8n8). B.M. (N.H.) Reg. No. 1965.973. A B C Body length (mm.) 55°58 20727 30279 2-89 MEASUREMENTS (in mm.) Body breadth: 0-052. Oesophagus length: 0-143. Body breadth at level of amphids: 0-017. Diameter of amphids: 0-0065. Length of cephalic setae: 0-013. Distance of excretory pore from anterior end of body: 0-058. Length of spicules: 0-062. Length of gubernaculum: o-or8. More posterior pre-cloacal supplement (distance anterior to cloacal opening/length) : 0-059/0:033. More anterior pre-cloacal supplement (distance anterior cloacal opening/length) : 0-124; 0-044. Length of tail: o-ogr. Cloacal diameter : 0-065. MARINE NEMATODES FROM DURBAN IOI The cuticle is marked by distinct annules separated from each other by narrow, shallow striations which start at the base of the lips. The annules are marked by distinct elongate punctations over the lateral region of the body from about the middle of the oesophagus to the tip of the tail (Text-fig. 38). There are two files of pores on each lateral surface of the body over its whole length. The mouth opening is bounded by six very distinct lips each of which bears two papillae (Text-fig. 36). More posteriorly, about the level of the anterior end of the oesophagus, there are four prominent, stout setae. The amphids are circular pits into which the nerves enter on the dorsal side. There is a distinct cheilostome. The anterior end of the oesophagus is modified as an elongate region with thickened walls, at the anterior end of which are two flat onchial-like modifications. These structures are at different levels so that from the ventral view the structure on the left ventro- lateral sector of the oesophagus hes anterior to the corresponding structure on the right ventro-lateral sector. In addition both project dorsally to le ina pocket formed in the dorsal sector of the oesophagus (Text-figs. 36 and 40). Posterior to the strongly cuticularized oesophastomal region is a region in which the dorsal sector of the oesophagus develops a slightly rounded bulge which displaces the ventro-lateral sectors. This region does not bear any tooth like structures. Posterior to this region the lumen of the oesophagus develops radial tubes which continue posteriorly for about two-thirds of the length of the oesophagus when they disappear. It has not been possible to make this out in detail but there is definitely no indication of a median bulb. The posterior end of the oesophagus is slightly club shaped and no definite bulb is present. Nevertheless internally it is modified to form a distinct small tri-valvulate structure at the posterior end of a long region in which the cuticular lining of the lumen of the oesophagus is markedly thickened (Text-fig. 37). The tail is short and stout and is curved strongly ventrally (Text fig. 43) so that the measurements of this region are probably rather low. On the tail roughly half way between the tip and the cloacal opening is a distinct papilla like structure while on the mid-ventral line of the body anterior to the cloacal opening there are three supplementary organs. Of these the most posterior lies just anterior to the cloacal opening and is a prominent papilla-like organ. The details of its structure are uncertain as the body is in a rather poor condition in this region but such a structure is definitely present and a nerve supplies it (Text figs. 39 and 43). The remaining pre-cloacal supplements are more typical. They are long massively cuticularized rods of which the more posterior (Text fig. 42) is rather simple in structure with a shght pit in its distal, protruding end. The more anterior is more massive and the distal end is expanded into a cap covered by many small denticles (Text fig. 41). The equal and identical spicules are simple in structure being roughly the same width along their full lengths. They terminate distally in slightly rounded tips which are enclosed by a small arrow like gubernaculum (Text fig. 39). DISCUSSION. See above, p. 100, 102 WILLIAM G. INGLIS wy LL MUO Le 38 42 Se : : Ties. 36-43. PLECTOLAIMUS juliani gen. et sp. nov. Fig. 36. Anterior end of body from lateral aspect with dorsal surface to the right. Fig. 37. Posterior end of oesophagus. Fig. 38. Sketch showing detail of punctation markings on lateral surface of body. Fig. 39. Lateral view of spicules and gubernaculum. Fig. 40. Sketch of modifications at anterior end of oesophagus from the ventral aspect. Fig. 41. Detail of anterior pre-cloacal supplement. Fig. 42. Detail of posterior pre-cloacal supplement, Fig. 43. Lateral view of male tail, MATERIAL S MARINE NEMATODES FROM DURBAN 103 Family TRIPYLOIDIDAE Bathylaimus deconincki sp. nov. (Text-figs. 44-47) TUDIED. I 3. Beach sand in surf zone. Near Durban, Republic of South Africa (B8n8). B.M. (N.H.) Reg. No. 1965.994. 57 A B C Body length (mm.) °95 3°48 16:26 2°26 MEASUREMENTS (in mm.) Body breadth: 0-039. Oesophagus length: 0-65. Diameter of head at level of posterior ceph alic setae: 0-030. Diameter of head at level of amphids: 0-034. Lateral diameter of amphid: o0-o10. Antero-posterior diameter of amphid : 0-009. Length of more anterior cephalic setae: 0-014. Length of more posterior cephalic setae: 0:038. onchia : 0:030. Depth of buccal cavity from anterior end to level of ventro-lateral Depth of second part of buccal cavity : 0-029. Length of spicules : 44 Tics. 44-47. Bathylaimus deconincki sp. nov. Fig. 44. Lateral view of head with dorsal surface to the right. Fig. 45. Ventral view of gubernaculum. Fig. 46. Lateral view of spicules and gubernaculum. Fig. 47. Lateral view of male tail. 104 WILLIAM G. INGLIS 0-030. Length of gubernaculum : 0-036. Length of tail: 0-139. Cloacal diameter: 0-051. The cuticle is marked by very fine, close-set transverse striations. The head bears two circles of six setae of which the members of the more posterior circle are typically cup-shaped distally. The four short setae usually associated with this posterior circle were not found but they may have been overlooked. The amphid is a prominent cavity with thin walls into which the nerve enters from the posterior edge (Text fig. 44). The mouth opening is bounded by three lips which meet at a level roughly half way between the two circles of cephalic setae. The buccal cavity is large and the thick cuticle lining it is continued as a lobe into each lip. In each lip, just anterior to the level at which each lobe of the buccal cavity lining stops, the cuticle is marked by a series of fine antero-posterior striations. The buccal cavity itself is tulip-shaped with two prominent ventro-lateral onchia at the level of its widest part and a sight bump in the corresponding dorsal position. In the region of the oesophagus immediately posterior to the buccal cavity the lumen of the oesophagus is slightly expanded to form a cavity which is not lined by thickened cuticle but which does bear three wholly cuticular onchia (Text-fig. 44). The oesophageal musculature is apparently attached to the cuticle of the body at the level of the more posterior circle of cephalic sense organs, The tail is fairly stout and carries a series of long, very thin setae which are about 0-02 mm. in length (Text fig. 47). The spicules are short, equal and identical in structure with very distinctly capitate proximal ends and appear to bear very narrow alae. The gubernaculum is the usual massive structure from the lateral aspect with a well chitinized posterior tip and a somewhat slightly chitinized massive region more anteriorly (Text fig. 46). From the ventral aspect, however (Text fig. 45), the gubernaculum is seen to be in two bilaterally symmetrical pieces consisting of a well chitinized median ridge which projects ventrally from a more expanded dorsal plate. The median ridge terminates posteriorly in a swelling from which two tooth- like structures arise on the inner sides (Text-fig. 45). Discussion The identification of species referable to this genus is, as with so many others, very difficult in view of the superficiality of most of the available descriptions. It can, however, be said that the genus Parabathylaimus De Coninck & Schuurmans Stekhoven, 1933 can certainly be accepted, as Luc & De Coninck (1959) argue, in spite of Wieser’s (1956) suggestion that the recognition of a divided or undivided buccal cavity is too difficult to make a useful character. Delimination of species within this genus appears to be based largely on the use of measurements and the structure of the buccal cavity, in spite of Weiser’s comments. Nevertheless the structure of the gubernaculum may be of value but there are very few useful descriptions available. The figures given by de Man (1922) and by De Coninck & Schuurmans Stekhoven (1933) for B. assimilis de Man, 1922 suggest that the form of the posterior end of the gubernaculum may be valuable since in that MARINE NEMATODES FROM DURBAN 105 species it is very different from the condition in B. deconinckt, in that the tooth-like modifications are much longer and slimmer and arise from the outer surface and not the inner as in my species. This species is otherwise characterized by the posterior position of the large amphid, the form of the tail and, apparently, by the shape and relative lengths of the guber- anculum and spicules. REFERENCES Cops, N. A. 1930. The demanian vessels in nemas of the genus Oncholaimus with notes on four new oncholaims. J. Wash. Acad. Sci. 20 (12) : 225-241. Dr Coninck, Lucien. 1965. Systématique des nématodes, in Tyaité de zoologie. Anatomie, systématique, biologique. Embranchement des Némathelminthes (Nematelmia Carl Vogt 1851—Nemathelminthia Gegenbauy 1859) ow Aschelminthes (Aschelmintha Grobben 1910). 4 (2) : 586-681. De Coninck, Lucien A. P. & SCHUURMANS STEKHOVEN, J. H. 1933. The free-living marine nemas of the Belgian coast. II. With general remarks on the structure and system of nemas. Mém. Mus. Hist. nat. Belg. 58 : 1-163. DITLEVSEN, HystMar. 1926. Freeliving nematodes. Dan. Ingolf-Evped. 4 (6) : 1-42. 1930. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. LII. Marine free- living nematodes from New Zealand. Vidensk. Medd. Dansk. naturh. Foren. 87 : 201-242. GERLACH, SEBASTIAN A. 1955. Zur Kenntnis der freilebenden marinen Nematoden von San Salvador. Z. wiss. Zool. 158 : 249-303. 1959. Drei neue Nematoden aus dem Kiistengrundwasser der Insel Abd al-Kuri (Golf von Aden). Zool. Anz. 163 : 360-364. —— 1962. Freilebende Meeresnematoden von den Maldiven. Sond. Keil. Meevresforsch. 18 : 81-108. Incuis, WILLIAM G. 1961. Free-living marine nematodes from South Africa. Bull. Brit. Mus. nat. Hist. (Zoology) 7 (6) : 291-319. 1963. ‘‘ Campaniform-type ”’ organs in nematodes. Nature 197: 618. 1964. The marine Enoplida (Nematoda) : a comparative study of the head. Bull. Brit. Mus. nat. Hist. (Zoology) 11 (4) : 263-376. Luc, Micuer & DE Conincx, L. A. 1959. Nématodes libre marines de la région de Roscoff. Arch. zool. exp. gén. 98 : 103-165. DE Man, J.G. 1922. Uber einige Marine Nematoden von den Kuste von Walcheren, neu fiir die Wissenschaft und fiir unsere Fauna, unter welchen der sehr merkwiirdige Catalaimus Max Weberi n. sp. Biyjdy. Dierk. 22 : 117-124. Mawson, Patricia M. 1958a. Free-living nematodes. Section 2. Additional Enoploidea from Antarctic stations. Rep. B.A.N.Z, Antarct. Res. Exped. Ser. B. 6 (13) : 293-305. 1958b. Free-living nematodes. Section 3. Enoploidea from Subantarctic stations. Rep. B.A.N.Z. Antarct. Res. Exped. Ser. B, 6 (14) : 307-358. SAVALJEV, S. 1912. Zur Kenntnis der freilebenden Nematoden des Kolafjords und des Relictensees Mogilnoje. Tvav. Soc. Nat. Leningy. 43 : 108-126. SOUTHERN, R. i914. Clare Island Survey. Part 54. Nemathelmia, Kinorhyncha and Chaetognatha. Pyvoc. R. Ivish Acad. 31: 1-80. Timm, RicHarp W. 1959. Cheilorhabditis and Cdontorhabditis, two new genera of soil nema- todes allied to Rhabditis. Nematologica 4 : 198-204. 106 WILLIAM G. INGLIS WIESER, WOLFGANG. 1953. Reports of the Lund University Chile Expedition 1948-1949. 10. Free-living nematodes. I. Enoploidea. Acta Univ. lund. N.S. 49 (6) : 1-155. 1956. Reports of the Lund University Chile Expedition, 1948-49. 26. Free-living marine nematodes. III. Axonolaimoidea and Monhysteroidea. Acta. Univ. lund. N.S. 53 (13) : 1-115. 1959. Free-living nematodes and other small invertebrates from Puget Sound beaches. Univ. Wash. Publ. Biol. 19 : 1-179. 20 ew Ns A BS WE RA A ON Mt WRC. LTR SO aL da PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING STUDIES ON THE BRITISH _-—-« DERMANYSSIDAE G. OWEN EVANS and awe Me TILL 7 BOlMeETIN OF ag SH MUSEUM (NATURAL HISTORY) ce : : Vol. 14 No. § TISH MUSE oR TRING 84, : GTR a MUSEUM N a! 4TuR,, wisto® SHODIES ON THE BRITISH DERMANYSSIDAE (ACARI : MESOSTIGMATA) PARA CLASSIFICATION BY G. OWEN EVANS AND W. M. TILL British Museum (Natural History) Pp. 107-370 ; 101 Text-figures BOLLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 14 No. 5 LONDON : 1966 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), tustituted 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 1s Vol. 14, No. 5 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 3 September , 1966 Price £5 5s. SODIES ON: THE BRITISH DERMANYSSIDAE (ACARI : MESOSTIGMATA) PART II CLASSIFICATION By G. OWEN EVANS & W. M. TILL CONZENTS Page SYNOPSIS : : : ; . F ; : : ; ; . 109 INTRODUCTION : ; : : ; ‘ . 109 CLASSIFICATION OF THE Britisu DERMANYsSID AT | a é : , Dn KKEY TO SUBFAMILIES : : : : : : : 22 Subfamily LAELAPINAE : : : ; : : : cael? Subfamily HAEMOGAMASINAE : : 5 : : 7238 Subfamily PSEUDOLAELAPINAE : : : : : . 265 | Subfamily MyonyssINAE ; : : 3 ; : . 268 Subfamily MELITTIPHINAE : ; : : ; F 273 Subfamily HirRSTIONYSSINAE : s : : : : = 270 | Subfamily MACRONYSSINAE : : ; : : ; . 300 Subfamily DERMANYSSINAE : : : : : : - 346 SPECIES INCERTAE SEDIS : : : : : : 3 + 362 SUMMARY OF NEW TAXA AND SyNoNyMY ; ; ; : : : . 362 ACKNOWLEDGEMENTS). 3 : : : : : ; : . 363 REFERENCES. : ; : 4 : , - 364 ABBREVIATIONS USED IN TEXT AND FIGURES. : : : : - 365 INDEX TO GENERA AND SPECIES ‘ 5 i : ; : : . 366 SYNOPSIS The classification of the Dermanyssidae s. lat. is discussed. The free-living, paraphagic, and ' facultative and obligatory ectoparasitic members of this family in the British fauna are distri- ' buted among eight subfamilies (Dermanyssinae, Haemogamasinae, Hirstionyssinae, Laelapinae, _ Macronyssinae, Melittiphinae, Myonyssinae and Pseudolaelapinae). Keys are given to sub- | families, genera and species. Eighty-one species and subspecies are described and figured. | | INTRODUCTION | THE earliest suprageneric classification of the mites included in the present study may _be attributed to Berlese (1892) who considered them to represent two families, the _Dermanyssidae and the Laelapidae. The Dermanyssidae Kolenati, containing the obligatory ectoparasites, included the genera Ophionyssus Mégnin, Ledognathus Canestrini and Dermanyssus Dugés while the free-living forms, paraphages of arthropods and facultative ectoparasites, comprising the genera Laelaps Koch, | Hy poaspis Can., Seiuws Koch, Iphis Koch, Stylochirus Can., Neoberlesia Berl., | Podocinum Berl. and [phiopsis Berl., were placed in the family Laelapidae Berlese. lin his last comprehensive classification of the Mesostigmata published in Acarotheca | Italica in 1913, Berlese retained his original concepts of the families although, by this EWeZoor. 14, 5. 8 j | | | 110 G. OWEN TEV ANS (& Wo Me irae time, the large number of new genera added to the Laelapidae in the first decade of this century enabled him to attempt a tribal classification of this family. He proposed four tribes: Laelapini, Iphiopsini (based on Iphiopsidae Kramer), Evi- phidini and Phytoseiini. Subsequent classificatory works on these families have dealt almost exclusively with the “ parasitic’? forms and the most significant contributions during the thirty years following Berlese’s classification are incorporated in the system presented by Vitzthum (1943). This author recognized a single family, Laelapidae, with thirteen subfamilies. The Dermanyssidae sensu Berlese was divided into two sub- families, Dermanyssinae and Liponyssinae Ewing (= Macronyssidae Oudemans), and the Laelapidae into six subfamilies, namely, Laelapinae, Hypoaspidinae Vitzthum, Hyletastinae Vitzthum (= Eviphidini Berl.), Iphiopsinae, Phytoseiinae and Rail- lietinae Vitzthum. Three of the remaining five subfamilies, Entonyssinae Ewing, Rhinonyssinae Trouessart and Halarachninae Oudms., contain respectively parasites of the respiratory tract of reptiles, birds and mammals which were included by Berlese (1913) in his family Pteroptidae (= Spinturnicidae Oudms.). Vitzthum further recognized the Haemogamasinae Oudms. and the Podocininae Berl. The subfamilial diagnoses are brief and generalized and, in certain subfamilies such as the Hypoaspidinae and Podocininae, bear little relationship to their generic composition. Nevertheless, Vitzthum’s classification provided an excellent summary of the existing knowledge of the group and a basis for subsequent revisionary studies. Baker & Wharton (1952) modified Vitzthum’s classification by distributing his thirteen subfamilies among nine families. Seven of the subfamilies were included in three families and each of the remainder was raised to family status as follows: Laelapidae (Laelapinae, Hypoaspidinae and Hyletastinae), Dermanyssidae (Der- manyssinae and Liponyssinae), Phytoseiidae (Phytoselinae and Podocininae), Iphiopsidae, Haemogamasidae, Raillietidae, Rhinonyssidae, Entonyssidae and Halarachnidae. No attempt was made to critically examine the classification at generic level. Evans (1957), in the course of a revision of the British free-living and parasitic members of the Laelapidae sensu Vitzthum, adopted in a restricted sense Baker & Wharton’s Phytoseiidae, raised the Eviphidini to family rank and combined the Laelapinae and Hypoaspidinae within the subfamily Laelapinae. Numerous genera previously placed in the Laelapinae, Hypoaspidinae and Podocininae were removed from the complex and distributed among the Aceosejidae and Neoparasitidae. Thus, with the earlier relegation of the Raillietinae into the synonomy (Evans, 1955) and the later elevation of the Podocininae to family status (Evans & Hyatt, 1958) Vitzthum’s “ Laelapidae ”’ was restricted to the following eight subfamilies: Lae- lapinae, Iphiopsinae, Haemogamasinae, Dermanyssinae, Macronyssinae, Rhinonys- sinae, Entonyssinae and Halarachninae. To these must be added Ixodorhynchinae Ewing (1923), Myonyssinae Bregetova (1956), Histrichonyssinae Keegan ef al. (1960) and Allolaelapinae Tipton (1960). This broad concept of the Laelapidae represents one of the two current approaches to the classification of the group. Its protagonists argue that it is premature to THE BRITISH DERMANYSSIDAE (ACARI) III attempt a familial classification of the complex when the morphology and biology of a large proportion of its members are so poorly known. Further, new subfamilial categories resulting from piecemeal revisions of the group can be more readily incorporated, provisionally, in a broad family unit than in a more rigid system, the alternative approach favoured by Baker & Wharton. In this latter system certain subfamilies of vertebrate parasites have been withdrawn from the family unit and given familial rank irrespective of their inter-relationships or their relationships with the free-living forms and invertebrate parasites remaining in the Laelapidae. The familial diagnoses are superficial and difficulties arise in the placing of newsuprageneric taxa; for example, the Myonyssinae are included in the Laelapidae by Bregetova (1956) but in the Dermanyssidae by Strandtmann & Wharton (1958). This juggling with subfamilial and familial categories does little to solve the problem of a natural classification of the “ Laelapidae’’. The basic requirement is a knowledge of the morphology and biology of the group at species level. The present contribution is restricted to a study of the free-living, paraphagic, and facultative and obligatory ectoparasitic members of the group occurring in the British fauna. In adopting a broad family concept, we have used the oldest available family name, Dermanyssidae Kolenati, 1859. It should be pointed out that this family taxon and the Iphiopsidae Kramer, 1886, have priority over Laelapidae? Berlese, 1892. CLASSIFICATION OF THE BRITISH DERMANYSSIDAE The adoption of a parasitic mode of life by certain Dermanyssidae realised an evolutionary potential which has produced a vast assemblage of morphologically and biologically diverse forms. Most students of the group consider the parasitic forms to have evolved, in the first instance, from nest-inhabiting laelapines. The transition from a polyphagous free-living to a monophagous parasitic existence probably involved anon-specialized feeding phase that included occasional meals of tissue fluids exuding from skin abrasions of the “‘ host’. Such a generalized type of feeding has been retained by certain species of the genera Androlaelaps, Laelaps and Haemogamasus. Evans (1955) suggested that the Hypoaspis complex may represent the “ primitive ”’ free-living group from which this transition was made. Its members show various degrees of association with other animals, either as nest inhabitants or paraphages, a wide range of feeding habits and an extensive geographical distribution. The least specialized members of this complex are probably the predatory forms inhabiting soil and humus, and on the basis of their morphology and ontogenetic development it is possible to define, at least for the purpose of discussion, a basic dermanyssid type with the following characteristics : Adults : Chelicerae chelate-dentate ; movable digit bidentate in the female, unidentate in the male ; pilus dentilis short, setiform ; dorsal seta and fissures present ; spermadacty] free distally and grooved ; arthrodial processes simple. Gnathosoma with three pairs of hypostomal and ? The genitive stem of Laelaps is ‘‘ Laelap ” and in accordance with the International Code of Zoological Nomenclature (Articles 29a and 32a (iii)) the family and subfamily taxa become LAELAPIDAE and LAELAPINAE respectively (see Domrow, 1963). 112 GG. OWENE VANS: & Wi Mie Worn one pair of capitular setae; deutosternum with six transverse rows of denticles; corniculi horn-like, parallel, grooved dorsally to accommodate salivary styli; pedipalps with five free segments and a two-tined apotele, chaetotaxy (2-5—6—-14-15) ; tectum capituli with anterior margin denticulate. Entire dorsal shield with 39 pairs of setae, of which 22 pairs (71-76, z1—26, sI-s6, v2-75) occur in the podonotal region and 17 pairs (J1—J5, Z1I—Z5, S1-S5 and px2—p#3) in the opisthonotal region, and 22 pairs of pores. Lateral unsclerotized integument with 7 + F# series of eight setae and UR series. Tritosternum bipartite, base longer than wide, laciniae pilose. Sternal shield in female with three pairs of setae and two pairs of pores ; metasternal setae free. Genital shield flask-shaped with one pair of setae. Anal shield with three setae, euanal setae absent. Opisthogastric cuticle with seven pairs of setae, metapodals small, subcircular. Peritrematal shields free posteriorly, peritreme extending beyond coxa I. Male with holoventral shield bearing ten pairs of setae excluding anals ; genital orifice pre-sternal. All legs six-segmented, excluding pretarsus, and with well developed retractile ambulacra ; chaetotaxy as defined for normal condition by Evans & Till (1965), without marked sexual dimorphism. Lavva ; Chelicerae chelate, weakly dentate ; pilus dentilis simple, dorsal seta and fissures present. Gnathosoma with two pairs of hypostomal setae, capitular setae absent ; deutoster- num with weakly developed denticles; corniculi horn-like, weak; pedipalps with five free segments and a two-tined apotele, chaetotaxy (o—-4—5—12-I1) ; tectum simple. Podonotal region with ten pairs of setae, nine pairs being situated on a weakly sclerotized podonotal shield ; opisthonotum with seven pairs of setae, no pygidial shield. Tritosternum bipartite, laciniae well developed. Sternal shield with three pairs of setae and two pairs of pores; metasternal and genital setae lacking. Anal shield with three setae, no euanal setae. Opisthogastric cuticle with four pairs of setae. Stigmata, peritremes and associated shields lacking. Chaetotaxy of legs normal for this instar of the Gamasina (Evans & Till, 1965). Protonymph :; Chelicerae chelate-dentate, movable digit bidentate, pilus dentilis simple, dorsal seta and fissures present. Gnathosoma with three pairs of hypostomal setae, capitular setae present ; deutosternum with six transverse rows of denticles, corniculi horn-lke ; pedipalpal chaetotaxy (I-4—5-12-15). Tectum with weakly denticulate anterior margin. Podonotal region with 16 pairs of setae, II pairs being situated on the podonotal shield. Pygidial shield large, with eight pairs of setae, and the unsclerotized region of the opisthonotum with six pairs. Mesonotal scutellae present. Tritosternum bipartite. Sternito-genital shield with three pairs of sternal setae and flanked by a pair of genital setae in the region of coxae IV ; metasternal setae absent. Anal shield with three setae, euanal setae lacking ; opisthogastric cuticle with four pairs of setae. Stigmata present, peritremes short, no peritrematal shield. Chaetotaxy of legs as defined for free-living Dermanyssidae (Evans & Till, 1965). Deutonymph ; Gnathosoma with characteristics of the female. Dorsal sclerotization com- prising single shield with lateral incisions indicating partial fusion of podonotal and opisthonotal elements. Chaetotaxy as in female. Sternito-genital shield bearing three pairs of sternal setae and one pair of metasternals ; genital setae flanking the shield in the region of coxae IV. Metapodal shields subcircular, small. Chaetotaxy of opisthogastric cuticle as in adult. Anal shield with normal three setae, euanals absent. Stigmata with well developed peritremes, peritrematal shields weakly developed. Chaetotaxy of legs as in adult. Among the Hypoaspis complex certain members of the “ genera’ Gaeolaelaps and Hypoaspis closely conform to our definition of the basic dermanyssid type, but the group as a whole shows considerable diversity in the external morphology of its THE BRITISH DERMANYSSIDAE (ACARI) 113 post-protonymphal developmental stages, particularly of the female. The general trends in the complex are towards increased sclerotization of the genitalshield (Pseudo- parasitus, Ololaelaps, Laelaspis and Gymnolaelaps), hypertrichy of the opisthonotal and opisthogastric regions of the idiosoma of the deutonymph and adults, hypertrichy of tibia III and genu IV and modification of the chelicerae for specialized feeding. In older classifications of the “‘ Laelapidae ”’ considerable emphasis was placed on the size and chaetotaxy of the genital shield at generic and subfamilial levels, but with our knowledge of the morphology of a wider spectrum of the complex it is evident that this character shows considerable variation between congeneric species and does not warrant the taxonomic importance attributed to it by some authors (see Zumpt & Patterson, 1951). Hypertrichy of the idiosoma and legs commonly occurs in those forms showing some degree of association with other animals. It is first evident at the deutonymphal stage and on the dorsal shield is apparent only in the region of the J series. Considerable inter- and intrageneric variability is evident in the hypertrichy of the opisthogaster and in the marginal series of the setae (R and UR). This is very marked in the Pueuwmolaelaps group. Modifications of the chelicerae mainly affect the dentition and the size of the digits, the chelicerae retaining their basic chelate form. In the British representatives of this complex specialization of the chelicerae is found only in certain members of the myrmecophilous group Laelaspis. There is no evidence of any marked modification of the hypostome for specialized feeding in the complex and the degree of morphological diversity is of the same order as that found in other families which have radiated into a comparable variety of terrestrial habitats, for example, the Ascidae. The Laelaps complex represents a group of the Hypoaspis stock which has evolved in close association with mammals, particularly rodents, and has retained to a remarkable degree the basic morphological features of its free-living, nest-inhabiting progenitors. Members of the genus Androlaelaps appear to be the least specialized group of the complex and have a wide geographical distribution in the nests of birds and mammals. Morphologically they can only be separated from species of the Hypoaspis complex (Gaeolaelaps) by the specialization of the digits and spermadactyl of the male chelicera and, to a lesser extent, by the tendency towards the enlargement of the pilus dentilis in the feeding stages. The functional significance of the non- chelate chelicerae of the males is not known, but this type of chelicera is by no means confined to the Laelaps complex. The chelicerae of the males of the myrmecophilous genera Laelaspis and Laelaspulus approach this condition and essentially similar chelicerae occur in such widely different ecological groups as Julolaelaps, associated with millipedes, and Hemuilaelaps, ectoparasitic on reptiles. As in the Hypoaspis complex there is a trend towards increased sclerotization of the genital shield which may bear up to six pairs of opisthogastric setae and in the more specialized members of the complex, Hyperlaelaps, Ondatralaelaps and Oryctolaelaps, hypotrichy of certain palpal segments and of the dorsal shield is not uncommon. Hypertrophy of the leg setae occurs in many species of the complex and these stout, spine-like setae probably assist the movement of the mite through the fur of the host. The Laelaps complex appears to be basically polyphagous but with a tendency towards haematophagy in 114 G. (OW ENSEVANS @aw. Mi TIlE the more specialized members. They are usually considered to be facultative ecto- parasites. As the result of the unspecialized type of feeding the gnathosoma shows only minor modifications. Corniculi are present although sometimes reduced in size and degree of sclerotization ; there are five to eight transverse rows of deutosternal denticles with a tendency towards a reduction in the number of denticles in each row ; the labrum is well developed and usually broad and the weakly sclerotized tectum capituli has a smooth anterior margin. The Haemogamasinae (Haemogamasus, Eulaelabs and Brevisterna) appear to represent another branch of the Hypoaspis stock which has evolved in close associa- tion with birds and mammals, but show a higher degree of morphological specialization than the Laelaps complex. A remarkable feature of the haemogamasines is the extensive hypertrichy of the idiosoma. This is not restricted to the opisthonotum and opisthogaster as in the majority of the Hypoaspis-Laelaps group, but extends into the podonotal region and, in some species, to the intercoxalregion. Hypertrichy is first apparent at the protonymphal stage and may be restricted to the opisthosoma, asin Lulaelaps, or also affect the podonotum, asin Haemogamasus. The nature of the sclerotization of the idiosoma in the protonymph, deutonymph and adults of Ewlaelaps and Haemogamasus conforms with the normal Hypoaspis type although there is increased sclerotization of the genital and metapodal shields in the female of Ezdaelaps. In Brevisterna reduced sclerotization of the dorsum of the idiosoma is evident and the deutonymph and some adults have an abbreviated dorsal shield and a small discrete pygidial shield. The gnathosoma has been subjected to considerable specialization in the course of the adoption of haematophagy by some species. Invariably the number of transverse rows of deutosternal denticles shows an increase in com- parison with the Hypoaspis-Laelaps complex and ranges between nine and 16 rows whilst the number of denticles in each transverse row tends to decrease to the extent that the denticles form a single longitudinal file in Brevisterna. Typical horn-like corniculi occur in the polyphagous species of Eulaelaps and Haemogamasus (e.g. H. pontiger), but there is a distinct trend towards their modification into membranous flaps in the obligate ectoparasites (H. liponyssoides, H. hirsutus and Brevisterna). The chelicerae also show considerable diversity inform. Unspecialized feeders retain the typical chelate-dentate form of the free-living Hypoaspis type, but in the blood- feeding deutonymphs and females of the group there is a tendency towards a reduc- tion in the dentition of the digits and the enlargement of the pilus dentilis. In the males the chelate form of the chelicerae is retained and the relatively short sperma- dactyl is never entirely fused with the movable digit. The anterior margin of the tectum may be fimbriated or smooth. The pedipalpal chaetotaxy is normal and the apotele is two- or three-tined. Deviations from the basic Hypoaspis type of leg chaetotaxy occur on genu IV and rarely on femora IT and III. Fain (1962), in his detailed study of the Ixodorhynchinae, has pointed out the close affinities of this subfamily with the Laelapinae (Hypoaspis-Laelaps group). Its members are ectoparasites of snakes and display some adaptive features which have not been observed in the Laelapinae and Haemogamasinae. The main evolutionary trends are towards decreased sclerotization of the dorsum and, to a lesser extent, the ~ THE BRITISH DERMANYSSIDAE (ACARI) II5 sternal regions of the adults, hypotrichy of the idiosoma and appendages (legs and pedipalps), hypertrichy of the idiosoma, hypertrophy of the coxal setae and corniculi, and the reduction of the fixed digit of the chelicera. The sclerotization of the dorsum of the adults shows considerable diversity and may consist of an entire or laterally incised shield partially covering the idiosoma, or two distinct shields, a podonotal and opisthonotal, which allow greater flexibility of the idiosoma for move- ment under the scales of the host. Chaetotactic patterns of the idiosoma and segments of the appendages are extremely diverse even among congeneric species. Holotrichy of the dorsum of the idiosoma and of the opisthogaster is rarely, if ever, present, the primary chaetotaxy being reduced by the retardation in the development of certain setae during ontogeny or obscured by regional hypertrichy at the deutonymphal and adult stages. The segmental chaetotaxy of the legs and pedipalps also shows considerable deviation from the normal laelapine type and is apparent in the varying degrees of reduction in the segmental chaetotaxy of the femora, genua and tibiae. Hypertrophy of the postero-ventral setae of coxae I-III and more rarely the antero- ventral seta of coxa I occurs to a greater or lesser degree in all the genera. It is most marked in the genus Asvatolaelaps where the postero-ventral seta of coxa I is greatly enlarged and resembles the posterior retrose spur on coxa I in certain ixodid ticks. The corniculi are horn-like and subparallel or convergent in the genera Hemulaelaps, Strandtibbettsia and Asiatolaelaps, but in Ixodorhynchus and Ixobiordes they are greatly enlarged and terminate in one or two retrose hooks which probably serve to anchor the mite to its host ina manner reminiscent of the Metastigmata. The hypostome invariably carries the normal three pairs of setae; the deutosternum is provided with five to seven transverse rows of denticles (each row comprising one to 20 denticles) and the tectum has a smooth anterior margin. The chelicerae are typically chelate- dentate in the females of Hemilaelaps and chelate-edentate in Asiatolaelaps. In both genera the pilus dentilis is conspicuous and has a tendency to be enlarged as in the Laelaps group. The chelicerae of the females of Stvandtibbettsia, Ixodorhynchus and Jxobioides show a reduction in the size of the fixed digit. It is present as a rem- nant with a pilus dentilis in Stvandtibbettsia but is completelyl acking in the other two genera. According to Fain (loc. cit.) the chelicerae of the male resemble those of the Laelaps group. The nature of the sclerotization of the venter of the idiosoma is surprisingly constant throughout the subfamily ; the sternal shield in the female usually has three pairs of setae and two pairs of pores ; rarely it is reduced in size and carries only two pairs of setae ; the genital shield is typically flask-shaped and bears the genital setae, and the anal shield is large. Stigmata and peritremes are present in both sexes and the peritrematal shield shows various degrees of develop- ment. The tritosternum is typically laelapine. The males have a narrow holo- ventral shield or separate sternito-genital and anal shields. The post-embryonic developmental stages comprise a larva, two active nymphal stages and the adult. The subfamily Iphiopsinae contains paraphages and parasites of Arthropoda, particularly Insecta and Myriapoda. Little detailed information is available on the morphology and biology of this group and the subfamily at present functions as a depository for the more specialized forms which cannot be accommodated in the sub- 116 G: ‘OWEN EVANS: @&: iW. “M: TILL family Laelapinae. Our somewhat fragmentary knowledge of the species associated with Myriapoda and Formicidae (Insecta) suggests that the extent of adaptive radia- tion with its attendant morphological and biological specialization is considerable. The trends in morphological specialization parallel those in members of the family which have evolved in association with vertebrate hosts: reduction in idiosomal sclerotization (Jacobsonia, Myrmoleichus), hypertrichy of the idiosoma (Dinogamasus), hypotrichy of the idiosoma and the appendages, reduction of the fixed digit of the chelicera (Berlesia), elongation and fusion of the spermadactyl with the movable digit of the chelicera in the male (Julolaelaps), increase in number of transverse rows of deutosternal denticles (Myrmonyssus), reduction in the length of the peritreme and the enlargement of the stigmata (Iphiopsis), reduction of claws and the migration of the anus to a terminal position. The hypostome, on the other hand, does not appear to be affected to the same degree as in the vertebrate parasites. The Myonyssinae probably represent an offshoot of the nest-inhabiting Hypoaspis- Laelaps group inwhich the main evolutionary trend has been towards the specialization of the gnathosoma. This is evident in the shape of the cheliceral shaft, the slender essentially edentate cheliceral digits, the reduction of the pilus dentilis and arthrodial processes, the 9 to 12 rows of deutosternal denticles arranged in a single file and the elongate, distally tapering tectum. On the other hand, the corniculi remain horn- like although often weakly sclerotized. The chaeotaxy of the dorsum of the idiosoma and of the legs and pedipalps is typically laelapine. Varying degrees of hypertrichy of the opisthonotum are present and the genital shield may bear up to 22 setae. The anal shield is characteristically much broader than long. This subfamily contains only the genus Myonyssus whose members are facultative (? obligatory) parasites of mammals. The subfamily Alphalaelapinae is based on the single species Alphalaelap saplodon- tiae (Jellison, 1945), an ectoparasite of the primitive rodent Aflodontia. In general appearance the species resembles Laelaps, but presents a combination of specialized morphological features which precludes its classification in the Laelapinae. The gnathosoma shows considerable specialization : the deutosternal denticles (up to I4 in number) are arranged in a single file, the corniculi are semi-membranous, the tectum is membranous with a fimbriate anterior margin and the pedipalpal chaetotaxy is (2-4-6-14). Both digits of the chelicerae in the female are modified. The weakly sclerotized fixed digit is slender, tapering distally, and has a hyaline, retrose tooth, but apparently no pilus dentilis, while the movable digit is large, strongly sclerotized, hooked distally and unidentate. In the male the chelicera is of the non-chelate Laelaps type with the spermadactyl enormously elongated and about five to six times the length of the cheliceral shaft. The reduced fixed digit lacks a pilus dentilis. Arthrodial processes and dorsal seta are absent. The chaetotaxy of the entire dorsal shield in both sexes is markedly hypotrichous, the podonotal region having 16 pairs of setae (zI present) and the opisthonotal region Io pairs (J1-5 and Z1-5) of which J4 and J5 are reduced to microsetae. The ventral sclerotization of the female comprises a sternal shield with three pairs of setae and two pairs of pores (the metasternals being free), a large flask-shaped genito-ventral shield bearing six pairs of setae of which two THE BRITISH DERMANYSSIDAE (ACART) 17 _ pairs are widely removed from the margins of the shield, anda normalanalshield. A sternito-genito-ventral and discrete anal shield are present in the male. Hypertrichy of the opisthogaster occurs in both sexes and the peritrematal shield is fused with the podal elements of coxa IV. All the legs are strong and terminate in well developed ambulacra. Certain coxal setae are stout and spine-like, and the anterior spine on coxa II is well developed. Coxae III and IV each have a posterior spine as in ety sss. The chaetotaxy of the leg segments is normal except for tibia II (1I—}, }—2), pgen III (2—?, 2—1), ania III (r—+, #—1), genu IV (1—#, 2-1) and tibia IV (a (I ). The deutonymph is an active feeding instar with the general Beton etic of the female. The Raillietinae, parasites of the auditory meatus of ungulates, display yet another grade of morphological specialization which is seen in the sclerotization and chaetotaxy of the idiosoma and in the form of the cheliceral digits. Dorsal sclerotization in both sexes is reduced and comprises a relatively small, entire shield reminiscent of the form in some macronyssines. Hypotrichy of the podonotal and opisthonotal regions is very marked in R. auris and R. hopkinsi, and results from the suppression of a number of setae which normally appear at the protonymphal and deutonymphal instars. The sternal shield in the female is not fused with the podal elements of coxae II and, in the male, the ventral sclerotization consists of a sternito-genital and a discrete anal shield. Both digits of the well sclerotized chelicera are edentate in the female, and the pilus dentilis and dorsal seta are reduced to microsetae or are absent. Well developed arthrodial processes are present at the base of the movable digit. In the male the fixed digit is reduced and the large spermadactyl is fused with the movable digit as in the Laelaps group. The hypostome and labrum also resemble those of the Laelaps group with the cornicul well sclerotized and blade-like. There are six to eight transverse rows of deutosternal denticles. The pedipalps show a deficiency in tibial chaetotaxy, this segment retaining the larval complement of 12 setae. Deviations from the normal chaetotaxy of the legs occur on femur I (only three ventral setae) and femur III (five setae). The immature stages of the railliet- Ines are not known. Domrow (1961) has provisionally placed within the genus Raillietia a species of dermanyssid (R. australis) which inhabits the ears of the wombat, Phascolomis mitchelli (Owen). This species, of which only the female is known, differs from auris and hopkinsi in having little or no hypotrichy of the idio- soma and thus shows a closer relationship with the Laelaps group. The subfamilies Dermanyssinae, Macronyssinae, Histrichonyssinae and Hirstionys- sinae comprise forms which are obligatory, haematophagous ectoparasites of reptiles, birds and mammals. They exhibit, in varying degrees, the following trends in morphological specialization : reduction in idiosomal sclerotization, hypotrichy of the primary chaetotaxy of the dorsum of the idiosoma, hypertrichy of the lateral and marginal series of dorsal setae and of the opisthogaster, modification of the corniculi and chelicerae, reduction in the number of denticles in each transverse row of deutosternal teeth, enlargement of the antero-dorsal spine of coxa II and diversi- fication of the segmental chaetotaxy of the appendages. These trends are also apparent in many of the other groups of parasitic Dermanyssidae (see above), but it is 118 Gi OWEN YVEVASNS & =Wi Mis ee amongst these subfamilies that we find the most morphologically and biologically specialized ectoparasites of vertebrates. Of the four subfamilies the Hirstionyssinae appear in many ways to be the least specialized. The edentate or weakly dentate form of the chelicerae, the modification of the corniculi into membranous flaps and the single file of deutosternal teeth are adaptive features of the gnathosoma which appear to accompany haematophagy. Some degree of hypotrichy of the dorsal chaetotaxy occurs in all members of the subfamily, but the segmental chaetotaxy of the legs is normal except for deficiencies in the chaetotaxy of tibiae III and IV in some genera. The most significant adaptive trend, however, is towards the develop- ment of spur-like structures on the coxae and, to a lesser extent, on the genu and tibia of the anterior pair of legs. These processes, which may be setigerous (hypertrophied setae) or non-setigerous (cuticular protuberances) in origin, undoubtedly assist the mite in its movements through the fur of the host. The lfe-cycle comprises a non- feeding larva, a (?) feeding protonymph and an active, feeding deutonymph and adult. The genus Hzrstzonyssus and its allies have previously been included in the Macronyssinae, but it is apparent that they represent a bio-morphological group which is quite distinct from the Macronyssus complex. Many of the similarities between the two groups in external morphology (gnathosomal features and hypotrichy of the primary setation of the dorsum) are probably the result of convergent or parallel evolution. Morphological specialization is so advanced in the Dermanyssinae that the external morphology of its members shows little resemblance to that of the laelapine stock. The extreme attenuation of the second cheliceral segment and reduction in the size of the digits in the protonymph, deutonymph and adult female are unique, while the high degree of specialization of the hypostome, the grades of reduction in dorsal idiosomal sclerotization, the hypotrichy of the idiosoma and the diverse segmental chaetotactic patterns of the legs and palps provide little tangible evidence as to the origin of the group or to its relationship with other obligatory ectoparasitic dermanys- sids. Within the subfamily the species infesting birds (Dermanyssus) display con- siderably greater morphological plasticity (both inter- and intraspecific), especially as regards chaetotaxy, than do those parasitizing mammals (Liponyssoides). The developmental cycle is normal in that it comprises an inactive, non-feeding larva without the cheliceral modifications of the subsequent nymphal stages, an active feeding deutonymph with the general characteristics of the female, and the adult. The protonymph is apparently a more active feeding stage than the equivalent instar in the free-living laelapines. The Macronyssinae also represent an extremely specialized group of obligatory ectoparasites but, unlike the Dermanyssinae, certain of its members have retained many features of the Laelaps group (particularly of Neolaelaps and Notolaelaps). These features are apparent in Bewsvella and Ichoronyssus, both parasites of Chirop- tera, and may be seen in the general form of the idiosoma and legs, dorsal and ventral sclerotization and slight hypotrichy of the dorsum of the idiosoma and appendages. The occurrence of the more “ primitive ’’ macronyssines on Chiroptera has led Radovsky (1964), in his comprehensive and stimulating study of the bat macro- ” a ae a a a a ee THE BRITISH DERMANYSSIDAE (ACARI) 119 _ nyssines, to postulate that the subfamily “‘ has evolved primarily on bats and secon- darily to have transferred to other mammals, reptiles and birds’’. This seems a logical hypothesis on the basis of the degree of morphological specialization exhibited by the mammal, bird and reptile forms. Certainly, as in the dermanyssines, the - parasites of mammals appear to represent the least specialized members of this sub- family, while the forms associated with birds and with reptiles show considerably greater morphological plasticity. All macronyssines have a strongly modified hypostome with hyaline corniculi which, in the majority, form a pre-oral trough. The chelicerae of the protonymph and female are essentially edentate although there is a tendency for the digits of the bat parasites to have denticulate hyaline pro- cesses. Only rarely is the fixed digit reduced in the male which has a grooved spermadactyl incompletely fused with the movable digit. Reduction in dorsal idiosomal sclerotization is often marked in both protonymph and female and the dorsal chaetotaxy shows some degree of hypotrichy, the paravertical setae 21 rarely being present. Many females and a few protonymphs and males have a conspicuous medio-ventral keel on the trochanter of the pedipalp, a structure which is not known to occur in the Dermanyssinae and is rarely present in the Haemogamasinae (some Haemogamasus) and Laelapinae (Aetholaelaps). The function of the keels is not known, but they may possibly protect and stabilize the somewhat elongated, membranous hypostome or, as Radovsky (1964) suggests, act as guides for the chelicerae. In addition to the adaptive features of the morphology, the macronys- sines display one feature of their biology which is unique amongst the facultative and obligatory ectoparasitic Dermanyssidae, that is, the replacement of the normal, active, feeding deutonymph by a relatively inactive, non-feeding instar which shows considerable degeneration of the feeding organs, tritosternum, idiosomal sclerotiza- tion and ambulacra. The presence of this stage (a quasi-calyptostase) has not been established for the genera Bewsvella and Ichoronyssus. It seems probable that the Dermanyssinae and Macronyssinae represent two distinct bio-morphological groups which have probably evolved quite independently of each other from amongst the Laelapinae. The Hystrichonyssinae, based on the genus Hystrichonyssus whose only known species is an ectoparasite of a Malayan porcupine (Atherurus macrourus) and an elephant tree snake (Acrochordus javanicus), present yet another combination of adaptive characters which includes modification of the hypostome for haematophagy, extreme elongation of the basal cheliceral segment, reduction in dorsal idiosomal sclerotization and marked hypotrichy of the primary chaetotaxy of the dorsum. Although the chelicerae superficially resemble those of the Dermanyssinae in their extreme length and needle-like proportions, they are the result of a different evolutionary trend, the secondsegment in Hystrichonyssus retaining the typical form of a macronyssine mite. Only the female is known. The subfamilies Entonyssinae, Rhinonyssinae and Halarachninae contain the endoparasitic members of the Dermanyssidae s. lat. In general, morphological specialization in these groups is characterized by simplification in structure and this is to be seen in the form of the gnathosoma, in the progressive trend towards extreme 120 G. OWEN EVANS &W. M. TILL reduction in idiosomal sclerotization, in the reduction of the tritosternum and in the high degree of hypotrichy of the idiosoma and appendages due to both larval and post-larval specialization. This contrasts with the tendency towards the elaboration of certain structures in the facultative and obligatory parasites, for example, the complex specialization of the hypostome, the hypertrophy of setae to form attach- ment organs and the hypertrichy of the idiosoma. Each subfamily exhibits grades of morphological specialization and many of the similarities between them are probably the result of convergent or parallel evolution. The least specialized biologically and morphologically are the Entonyssinae which inhabit the respiratory tract of reptiles. They show comparatively little reduction in idiosomal sclerotization and chaetotaxy when compared with the dermanyssids (Ixodorhynchinae) ectoparasitic on reptiles and, in the female, the chelicerae are well developed, chelate-dentate (rarely edentate), and usually have a dorsal seta, arthrodial processes, fissures and a pilus dentilis while in the male the fixed digit is reduced and the long spermadactyl is almost entirely fused with the slender movable digit. All grades of reduction of the tritosternum are evident, from the large bipartite form in Entonyssus to its complete absence in Hamertonia. The hypostome is simple and weak corniculi may be present or absent. Fusion of the palptibia and tarsus to form a tibiotarsus and a reduction of the apotele occur insome forms. The segmental chaetotaxy of the appendages is diverse. Two active nymphal stages occur in the life-cycle, the deutonymph showing no degenera- tion of the feeding organs (Fain, 1961). Present morphological and _ biological evidence suggests that the Entonyssinae have probably evolved from amongst the Ixodorhynchinae. Fain (1961 and 1962) classifies the [xodorhynchinae within the family Laelapidae but gives familial status to the Entonyssinae. In our opinion this exaggerates the difference between the two groups and has little practical value. The Rhinonyssinae are parasites of the respiratory tract of birds and the degree of reduction in idiosomal sclerotization and hypotrichy of the idiosoma and appendages exceeds that in the Entonyssinae and in any of the groups of ectoparasitic dermanys- sids. Reduction or loss of the tritosternum is a common feature of the group, the peritremes may be strongly reduced or lacking and the stigmata displaced to a dorso- lateral position. Modifications of the chelicerae affect the size of the digits, their dentition and shape of the shaft. The gnathosomaland pedipalpalchaetotaxy shows varying degrees of reduction but the hypostome is invariably simple with the corniculi reduced or absent. As in other endoparasitic dermanyssids, the sensory area on tarsus I is extensive with the eupathidia- or solenidia-like setae dispersed and not aggregated as in the free-living and ectoparasitic forms. The life-cycle of the rhinonyssines resembles that of the macronyssines in having an inactive deutonymph with non-functional trophic appendages. Many of the less specialized members of the subfamily, e.g. Mesonyssoides, show a striking morphological resemblance to the macronyssines and we are inclined to agree with Radovsky (1964) that the rhinonys- sines have probably evolved from the Macronyssinae. Endoparasites of the respiratory tract of mammals are included in the subfamily Halarachninae. The gnathosoma of the adults is relatively simple ; the hypostome lacks corniculi, the deutosternal denticles are reduced, the pedipalps show a trend THE BRITISH DERMANYSSIDAE (ACART) [20 _ towards fusion of the tibia and tarsus and, in Pnewmonyssus, a marked reduction in the size of the segments, and the apotele may be present or absent. The chelicerae may be small with the digits edentate (Puewmonyssus) or strongly developed with the fixed digit reduced or absent in the female (Halarachne). Hypotrichy of the venter - of the gnathosoma, pedipalps and idiosoma is often very marked. The dorsum is - usually partially covered by an entire shield while the genital shield is reduced or absent. Many segments of the legs retain their larval complement of setae. All legs have ambulacra, normally with strong claws. Biologically this group differs from all other known Dermanyssidae in having an extremely active larva which probably functions as the dispersal stage, but no active feeding nymphal stages. The nymphal instar (or instars) described by Hull (1956) for Puewmonyssus simicola Banks appears - to be non-feeding and of short duration. In Puewmonyssus the larval pedipalps _ show considerably less specialization than in the adult and the legs have long, slender - pretarsi with unmodified pulvilli and claws. Recently Domrow (1961) has described a nymphal (protonymph) dermanyssid from the nasal passages of a marsupial mouse (Dasyuridae) and two species of scale-tailed rats (Muridae). On“ purely ecological grounds ”’ this species was placed jn the genus Pneumonyssus (Pneumonyssus dentatus Domrow). The protonymph presents a combination of laelapine and macronyssine characters and is certainly not congeneric with Pnewmonyssus. In our opinion it should not be classified in the Halarachninae. Macronyssine features are seen in the nature of the sclerotization of the dorsum of the idiosoma which comprises a well defined podonotal shield with the normal eleven pairs of setae, mesonotal scutellae and a reduced pygidial shield bearing seven pairs of setae and similar in form to that of the protonymph of certain Macronyssus. The gnathosoma, on the other hand, exhibits certain features of the Laelapinae, horn- like corniculi and up to nine transverse rows of deutosternal denticles, each row being multidentate. The chelicerae are extremely specialized, the basal segment is strongly sclerotized and is about two-thirds the length of the second segment which terminates in a large, hook-like, fixed digit lacking a pilus dentilis while the shorter movable digit is slender and edentate. Hypertrophy of certain gnathosomal and leg setae is also a feature of the species. A decision on the systematic position of P. dentatus must await the discovery of the adult stages, but, in the meantime, the unique combination of morphological characters displayed by the protonymph requires recognition and we propose the genus Domrownyssus (type: Pneumonyssus dentatus Domrow, 1961) for its reception. Provisionally the genus may be placed in the Laelapinae. The above general review of the characteristics of the major subfamilies of the Dermanyssidae illustrates the high degree of morphological and biological specializa- tion which has accompanied the adoption of a parasitic mode of life by certain members of the family. Each subfamily appears to represent a distinct bio-morpho- logical group and exhibits grades of morphological specialization. A classification of the family purely on the basis of the external morphology of its members is fraught with difficulties, the most significant being the difficulty of defining concise practical units. It is only those groups of obligate ectoparasites, for example the Dermanys- 122 G: (OWEN EVANS? @ Wi Me ELIE sinae and Macronyssinae, in which the transition from polyphagy to monophagy (haematophagy) has been universally accomplished and theaccompanying adaptations of the chelicerae, hypostome and life-cycle are more or less stabilized, that a group- diagnosis appears to be at all practicable. In the other groups, the members present grades of morphological specialization depending on the nature of their association with the host. This is clearly seen in the structure of the chelicerae and hypostome of the Haemogamasinae, and in the degree of development of fixation organs (retro- grade spurs) in the Ixodorhynchinae. Invariably, the least specialized members of the group provide a link between its more specialized members and the generalized laelapine forms. In these circumstances biological criteria may prove to be of con- considerable value in defining functional suprageneric categories. The free-living, paraphagic and ectoparasitic members of the Dermanyssidae represented in the British fauna appear to fall, with two exceptions, into six sub- families or biomorphological groups, namely the Laelapinae, Haemogamasinae, Myonyssinae, Hirstionyssinae, Dermanyssinae and Macronyssinae. The two species which cannot be accommodated in these subfamilies, Pseudolaelaps doderoi Berlese and Melittiphis alvearius (Berlese), appear to have greater affinities with the Laela- pinae than with any other subfamily. We have already referred to the systematic position of P. doderoi as being “ problematical ’’ (Evans & Till, 1965). This arises from two unusual features of the external morphology of the adults, the marked hypotrichy of the idiosoma and appendages and the trispinate tectum, which immediately distinguish them from any known free-living laelapine. MM. alvearius, previously classified in the Eviphididae (see p. 273), also presents a combination of morphological characters (form and chaetotaxy of the dorsal shield, sclerotization of the presternal region, nature of the deutosternal denticles etc.) which precludes its classification in the Laelapinae. Provisionally, and as a practical expediency, we are proposing suprageneric categories for each of these species. The following key to the subfamilies is based on the characteristics of the British species of free-living and ectoparasitic Dermanyssidae. A comprehensive diagnosis of these groups is given elsewhere in the text. It is rarely that any one “ character ”’ is sufficiently stable thoughout a subfamily to be entirely diagnostic, therefore the separation of the groups has been based on combinations of characters. IKEY TO THE SUBFAMILIES OF THE BrITISH DERMANYSSIDAE 1 Chelicerae in the nymphal stages and female with the second segment enormously elongated, stylet-like ; digits minute. Males with fixed digit reduced and the elongate spermadactyl entirely fused with the movable digit; chelicerae never chelate. Tarsi III and IV of the male with seta pv, modified into a short, tooth- like projection. Both sexes with paravertical setae absent. Obligatory ecto- parasites of birds . : : : ‘ : ‘ DERMANYSSINAE (p. 346) — Second segment of the chelicerae in nymphal stages and female never conspicuously elongated, digits prominent. Seta pv, on the tarsi of the males never modified into a tooth-lke projection Ny Dorsal shield in deutonymphal and aul stages aricediy) ie per eachons setation often resembling a pelage. Genital shield, and sometimes sternal and anal shields, with unpaired accessory setae. Deutosternum with Io or more transverse rows of denticles, majority of rows compound. Tectum with margins conspicuously nN THE BRITISH DERMANYSSIDAE (ACART) 123 y fimbriated. Pedipalpal chaetotaxy normal. Chelicerae normally chelate-dentate in both sexes, rarely edentate, but pilus dentilis prominent. Associated with birds and mammals as nest inhabitants or ectoparasites . AEMOGAMASINAE (p. 238) _— Hypertrichy of the dorsal shield, when present, restricted to the region of the J series or, if more extensive, deutosternal denticles in a single file and no accessory setae on the genital shield. : : : : : , : 5 3 3 Genu and tibia I with two ventral setae C= 3—2), genu IT with one ventral seta (2—3, 22). Tectum trispinate. Marked hypotrichy of the dorsal shield, podonotal region with 15 pairs (paraverticals absent) and opisthonotal with nine pairs of setae. Podal-peritrematal shields in female greatly enlarged posterior to coxae IV and flanking the genito-ventral shield. Chelicerae in both sexes chelate-dentate. | Free-living. : . PSEUDOLAELAPINAE (p. 265) / - Genu and tibia I sermaily wath fide venta setae (2—3, #—2z),* genu IT with two ventral setae (2—3, 2—2). Without the above combination of characters . : 4 | ' 4 Anal shield in the female considerably wider than long, anterior margin concave | (Text-fig. 648). Chelicerae essentially edentate, digits long and slender in both sexes, spermadactyl entirely fused with movable digit in the male; pilus dentilis, arthrodial processes and dorsal seta apparently absent. Deutosternum with 9-12 denticles arranged in a single file. Paravertical setae (z1) present in both sexes. Genital shield in the female with unpaired accessory setae. Anterior spine of coxaII prominent. Associated with small mammals. : . MyYOoNySSINAE (p. 26 — Without the above combination of characters. 5 5 Corniculi horn-like, strongly or weakly sclerotized, never in the frei of iyaline jokes Chelicerae in the female usually chelate-dentate, rarely edentate but pilus dentilis present. Chelicerae of the male chelate-dentate with free spermadactyl, or with reduced fixed digit and elongate spermadactyl partially or completely fused with movable digit, non-chelate. Dorsal chaetotaxy holotrichous or hypertrichous, if hypotrichous, paravertical setae present or chelicerae in both sexes chelate- | dentate (Ololaelaps sellnicki). Free-living, nest inhabitants and facultative (? obligatory ) ectoparasites of vertebrates ; 3 : : : 6 — Corniculi never horn-like, usually in the form of eatin’ ones Chelicera in the female with digits essentially edentate, often with smooth or denticulate hyaline lobes, pilus dentilis absent. Chelicerae of the male edentate, digits subequal, | fixed digits never markedly reduced in length, spermadactyl free distally, rarely approaching the length of the second cheliceral segment. Primary dorsal chaeto- taxy invariably hypotrichous, a belies setae absent. Obligatory ecto- parasites of vertebrates. . 9 : a , : 7 6 Presternal area in the female with a pee Hike gee) "eaeostenham in both sexes with about eight denticles arranged in a single file. Cheliceral digits in the female chelate, weakly dentate, pilus dentilis prominent, in the male non-chelate, fixed digit reduced, elongate spermadactyl completely fused with movable digit. Dorsal shield with general hypertrichy. Anal shield (female) or ventro-anal shield (male) with posteriorly directed spur anterior to the anus. Male with sternito-genital and ventro-anal shields. In bee-hives : MELITTIPHINAE (p. 273) — Presternal area in the female reticulated or with pre-endopodal shields, never with a ' keel-like ridge. Deutosternum with five to seven (usually six) transverse rows of denticles, rows usually compound, sometimes reduced to a single denticle. Cheli- cerae in the female chelate-dentate or, rarely, chelate-edentate, pilus dentilis present, in the male chelate-dentate with spermadactyl free anteriorly or non- chelate with edentate fixed digit reduced and spermadactyl entirely or partially eo a 2 Ornithonyssus sylviavum is exceptional in having only two ventral setae on tibia and genu I in some | specimens, but the chaetotaxy of genu II is normal (2—, 3—2). ZOOL. 14, 5. 9 124 G: OWEN VEWANSS & We M> Tiley fused with the movable digit. Hypertrichy of the dorsal shield, when present, usually restricted to the region of the J series of setae. Anal shield never with posteriorly directed spur. Male with holoventral shield or with discrete sternito- genito ventral and anal shields : : . LAELAPINAE (p. 124) 7 Both sexes with one or more of the coxae val rounds non-setigerous spurs, excluding the anterior spine of coxa II; genu IV with one ventral seta. Palp- trochanter without medio-ventral keel. Dorsal shield entire in both sexes. Deutonymph active, feeding, and resembling the female in the structure of the gnathosoma and its well developed, bipartite tritosternum HIRSTIONYSSINAE (p. 276) - Both sexes without retrograde non-setigerous spurs on the coxae (excluding anterior spine on coxa II), at the most with weak ridges; genu IV with two ventral setae. Dorsal shield entire in the male but with entire, two subequal shields or with discrete podonotal, mesonotal and pygidial elements in the female. Palptrochanter usually with a medio-ventral keel in the female (weakly developed in Ophionyssus and some Ornithonyssus). Deutonymph inactive, non-feeding, with degenerate (larviform) chelicerae and a simple tritosternum bipartite only at the tip : : : : : é : : : MACRONYSSINAE (p. 300) The measurements given in the descriptions of species apply, in most cases, to a single specimen ; no attempt has been made to assess intraspecific variation. They have been taken as follows, unless otherwise stated in the text : Chelicera: segment I, maximum length; segment II, from base of segment to apex of fixed digit; movable digit, maximum length; spermadactyl, from base of movable digit to tip of spermadactyl. Dorsal (or podonotal) shield: length in midline ; width at level of seta 26. Opisthonotal shield : width at level of seta JT. Pygidial shield : width at level of antero-lateral corners. Tritosternum : base, to point of division into two laciniae ; laciniae, from point of division to tip. Sternal shield: length in midline ; width at level of second pair of sternal setae (si2)e Genital (or genito-ventral) shield: length from level of genital setae to posterior margin ; width at level of genital setae (gen.). Anal shield: length in midline to base of postanal seta ; width through middle of anus. Holoventral shield : length in midline from anterior margin of genital orifice to base of postanal setae ; width at level of second pair of sternal setae (s?2). Distances between setae (sti, st3, c.s., hyp. 2, Jv2) : from centre of setal bases. Segments of appendages (legs and pedipalps) : median dorsal length ; width at base of segment. Subfamily LAELAPINAE Berlese Laelaptidae Berlese, 1892, Acari, Myr. Scorp, Ital. Ordo Mesostigmata : 30. Adults : Chelicerae chelate-dentate in the female, movable digit bidentate ; digits rarely edentate. Male chelicera chelate-dentate with spermadactyl long, grooved and free distally or non-chelate with spermadactyl partially or entirely fused with the THE BRITISH DERMANYSSIDAE (ACARI) 125 movable digit ; digits subequal in length or fixed digit markedly reduced, edentate. Pilus dentilis present, setiform or inflated; dorsal seta, fissures and arthrodial processes usually well developed. Chaetotaxy of venter of gnathosoma normal. Pedipalps with five free segments, apotele two- or three-tined, chaetotaxy of trochanter to tibia (2-5—6-14), rarely femur, genu or tibia unideficient (2-5—5-14 ; 2—4/5-6 13/14). Palptrochanter rarely with a medio-ventral spur-like ridge. Deutosternum with five to seven, usually six, transverse rows of denticles; transverse rows sometimes reduced to a single denticle. Corniculi essentially horn-like, rarely with a hyaline expansion. Internal malae simple or branched, margin smooth or fringed. Tectum capituli well sclerotized with anterior margin denticulate, or membranous with smooth anterior margin. Dorsal sclerotization in both sexes comprising an entire dorsal shield, rarely retain- ing the deutonymphal incisions. Primary chaetotaxy of dorsal shield comprising 39 pairs of setae of which 22 pairs occur in the podonotal region and 17 pairs in the opisthonotal. Variants are due to addition of one pair of posterior accessories (px1I), or a pair of anterior accessories (ax), or to suppression of posterior accessories (px series), 73, 23, J3, J4 and Z3 and rarely z1. Hypertrichy of the shield usually restricted to the region of the J series, rarely affecting the entire shield. Marginal series showing considerable diversity in the number of setae. Dorsal setae simple, pilose, subspinose, clavate or cuneiform. Tritosternum bipartite, laciniae well developed, smooth or pilose. Pre-endopodal shields present or absent. Sternal shield in the female with three, rarely four, pairs of setae and two or three pairs of pores. Metasternal setae, when free, situated on unsclerotized cuticle or on platelets. Genital shield narrow or variously inflated and bearing the genital setae and, in some forms, up to five pairs of opisthogastric setae. Anal shield free or fused with the genito-ventral shield to form a genito-ventro-anal shield (Ololaelaps) ; with the normal three setae, no euanal setae ; aciculated area conspicuous. Opisthogastric cuticle with varying grades of hypertrichy. Metapodal shields usually small, rarely fused with the genito-ventral shield. Peritrematal shields usually free posteriorly, but may be fused with the podal elements behind coxae IV. Podal shields poorly or well developed. Stigmata with well developed peritremes usually extending up to or beyond coxa I. Male with holoventral shield, rarely with sternito-genito-ventral and anal shields. Genital orifice presternal. All legs six-segmented with ambulacra ; claws sometimes reduced or absent. Variants from normal segmental chaetotactic patterns may occur on tibia I (2—3, 3—2), genu III (2—?, 2?—2; 2—2, 2—1), tibia III (2—}, ?—2; 1—4+, ?—1) and 2 ul! le 1 genu IV (2—2, 32; 2-2, 3—2; 1—#?, 2—-1). Anterior spine on coxa II promi- nent, small, or absent. Seta av, on femur II may be modified to form a stout spine or spur in both sexes or in the male only. Larva: Relatively inactive, non-feeding instar. Chelicerae small, chelate, dentate or edentate ; dorsal setae and fissures present ; pilus dentilis present but often reduced ; arthrodial processes weakly developed or absent. Hypostome with two pairs of setae ; corniculi small, horn-like ; internal malae simple. Deutosternum with five to seven transverse rows of denticles. Pedipalps with five free segments, 126 G. OWEN EVANS: & W.-M. PILL apotele two- or three-tined, chaetotaxy (0—4-5-12). Dorsal sclerotization weak or absent, podonotal region with ten, rarely nine, pairs of setae. Chaetotaxy of opis- thonotum variable. Tritosternum bipartite, laciniae usually smooth. Sternal region with three pairs of setae ; opisthogaster with four pairs, excluding the three setae associated with the anus. Stigmata and peritremes absent. Segmental chaetotaxy of the legs normal for this instar. Protonymph : Active (?) feeding stage. Gnathosoma, excluding pedipalps, similar to that of the female but not so well developed. Pedipalps with normal chaetotaxy (1-45-12). Dorsal sclerotization comprising well developed podonotal, mesonotal and pygidial elements. Holotrichous condition of the dorsum consisting of 30 pairs of setae ; podonotal shield usually with ro or 11 pairs, pygidial shield with eight pairs. Trito- sternum asin adult. Sternal shield with three pairs of setae and two pairs of pores ; genital setae free on integument. Anal shield normal. Opisthogastric cuticle with four pairs of setae. Stigmata and short peritremes present. Chaetotaxy of legs normal for this instar. Deutonymph : Active feeding stage. Gnathosoma as in female. Dorsum with a single, incised, rarely entire, dorsal shield. Dorsal chaetotaxy as in adult. Sexual dimorphism affecting marginal series in some forms. Tritosternum as in adult, sternito-genital shield with four pairs of setae (metasternals present) and three pairs of pores; genital setae free on integument. Anal shield normal. Opisthogastric cuticle with seven pairs of setae or displaying hypertrichy. Peritremes as in adult, peritrematal shield always free posteriorly. Metapodal shields present. Segmental chaetotaxy of legs as in corresponding adult. KKEY TO THE GENERA OF THE BRITISH LAELAPINAE 1 Female with anal shield completely fused with genito-ventral shield. Metasternal setae situated on sternal shield. Dorsal shield arched, completely covering dorsum of mite. Apotele with three unequal prongs. 2 : OLOLAELAPS (p. 228) — Female with anal shield free. Metasternal setae not on sternal shield. : 2 2 Genu IV with two postero-lateral setae, pl, present (2—?, 3—2). Pilus dentilis long, slender or inflated. Chelicerae of male strongly modified, never chelate Oo — Genu IV with one postero-lateral seta (2—?, 3—1) or, if two postero-lateral setae present, pilus dentilis short, setiform. Chelicerae of male chelate-dentate . 6 3 Female with genital shield bearing the genital setae only . ANDROLAELAPS . 150) — Female with genito-ventral shield bearing three pairs of opisthogastric setae in addition to the genitals. : : : : : : 4 4 Medio-dorsal series of setae nyeouichons 3, Bs va ela Femur I with seta Jd, arising from a_ strong, sclerotized protuberance. Associated with Ondaiva, introduced . : : . ONDATRALAELAPS (p. 145) — Medio-dorsal series of setae normal, ‘comprising II pairs of setae (j1-j6; J1-—J5). Seta pd, on femur I not arising from a strong protuberance : : . : 5 5 Genu III withnine setae. Dorsal setae rarely short, subspinose ; $¥2 and )¥3 present. Male with holoventral shield . : 3 LAELAPS (p. 127) — Genu III with eight setae. Dorsal setae Shore subspinose, never more than one pair of px setae. Male with sternito-genito-ventral and anal shields HYPERLAELAPS (p. 140) THE- BRITISH DERMANYSSIEDAE: (ACART) 12 6 Female with large genito-ventral shield bearing four or more pairs of setae of which two pairs are widely removed from its lateral margins. Pedipalpal apotele three- tined, posterior tine small : : : . PSEUDOPARASITUS (p. 224) — Female with genital shield bearing genital cae only or, if with genito-ventral shield, the additional setae are situated on its lateral margins. Pedipalpal apotele normally two-tined, rarely with three subequal tines BYPOASPIS s. lat. (p. 158) Genus LAELAPS Koch Laelaps Woch, 1836, Deutschl. C.M.A.4:19; Koch 1843, Avachnidensystems 3: 88. Echinolaelaps Ewing, 1929, Manual of External Parasites : 10 and 185. Macrolaelaps Ewing, 1929, Manual of External Parasites : 185. Myolaelaps Lange, 1955, Opred. Faune SSSR. 59: 328. Rattilaelaps Lange, 1955, Opred. Faune SSSR. 59 : 328. Microtilaelaps Lange, 1955, Opred. Faune SSSR. 59 : 329. Type: Laelaps hilaris Koch, 1836. Medium sclerotized mites ranging from 500 y to 1,500 yz in length. Chelicerae chelate-dentate in the female with the movable digit bidentate. Fixed digit in the male markedly reduced, normally edentate ; movable digit edentate and partially or entirely fused with the elongated, grooved spermadactyl. Pilus dentilis in both sexes normally large and inflated distally. Dorsal seta and fissures normal. Arthro- dial processes present at base of movable digit. Chaetotaxy of venter of gnathosoma and pedipalps normal, apotele two-tined. Deutosternum normally with six, rarely seven, rows of denticles, each row with one to three denticles. Corniculi horn-like, weak to medium sclerotization. Internal malae simple or fringed. Labrum broad, striated. Tectum capituli membranous, rounded anteriorly, never denticulate. Dorsum of idiosoma with entire shield bearing 39 pairs of setae ; no unpaired accessory setae. Tritosternum normal, bipartite. No definite pre-endopodal shields. Sternal shield in the female with three pairs of setae and two pairs of pores. Genito-ventral shield variously inflated and bearing four pairs of setae. Pear-shaped anal shield with normal chaetotaxy, paranal setae in line with posterior margin of anus or posterior to it. Opisthogastric cuticle with variable number of setae and affected by sexual dimorphism which is first apparent at the deutonymphal stage. Peritrematal shields free posteriorly, often weakly developed. Metapodal shields present. Male with holoventral shield not fused with peritrematal shields and bearing nine or ten pairs of setae excluding the anals. Podal shields poorly developed posterior to coxae IV. Chaetotaxy of legs normal except for genu IV which has two postero-laterals (2—}, 23—2) and sometimes two ventrals (2—?, #—2). Setae ad, and pd, on femur I and less conspicuously on femur II enlarged and elongated. Coxal setae often thickened to form stout spine-like structures. Coxa II with anterior spine. Am- bulacra of legs II-IV with paired claws. Claws of leg I often reduced. KEY TO FEMALES 1 Genu IV with two ventral setae (pu present) ; paranal setae short, their length equal approximately to width of anus (Text-fig. 1B) . . ° L. agilis Koch (p. 132) 128 G:' OWEN- EVANS &4W.. M.) TIED — Genu IV with one ventral seta (pu absent) ; Eee setae relatively oe their length at least twice the width of the anus : Genito-ventral shield greatly expanded behind coxae IV, ee BOsterios angen deeply concave ; distance between genital setae (gen.) less than that between Jv2. Dorsalsetae relatively long ; length of 75 distinctly greater than distance between 75 and s5 (Text-fig. 34). Pilus dentilis setiform : : L. echidnina Berlese (p. 134) — Genito-ventral shield only moderately expanded behind coxae IV, posterior margin rounded or very slightly concave ; distance between genital setae (gen.) greater than distance between Jv2. Dorsal setae relatively short ; length of 75 distinctly less than distance between 75 and s5. Pilus dentilis inflated ; : 3 3 Sternal setae relatively long and slender, s¢r reaching posterior margin of octave shield. Dorsal setae slender ; marginal opisthogastric setae with bifid tips L. hilaris Koch* (p. 128) — Sternal setae short and stout, s¢#1 reaching base of s¢f2. Dorsal setae stout and spine- like ; marginal opisthogastric setae simple : : L. muris (Ljungh) (p. 137) N No Kery To MALES r Genu IV with two ventral setae (pu present) ; tarsus IV with setae ad, and ad, very long, thick, sword-like, their length at least twice the basal width of the tarsus (Text-fig. 2c). Peritremes relatively short, extending to posterior margin of coxae II. Paranal setae short, their ea approximately equal to width of anus. Chelicera as in Text-fig. 2E_ . 5 L. agilis Koch (p. 132) — Genu IV with one ventral seta (pu alpeent) B nests IV qth setae ad, and ad, setiform or blade-like but not exceeding 14 times the width of the tarsus. Bentremes longer, extending at least to anterior margin of coxa II. Paranal setae longer, their length at least 14 times the width of the anus : : : Holoventral shield with ‘eae pairs of stout setae in genito- _ventral region (Text-fig. 5D). Tarsus IV with setae ad, and ad, blade-like ; seta pv, short, stout and spine-like iS) iS) (Text-fig. 5F). Chelicera as in Text-fie. 5B : : * LL. muris (Ljungh) (p. 139) — Holoventral shield with six pairs of relatively slender setae in the genito-ventral region. Tarsus IV with setae ad, ad, and pv, normal ; : 3 3 Coxa III with posterior seta stout, spine-like ; tarsus 1V withsetaea/,and a Stout and spine-like. Pilus dentilis inflated near its distal end ; chelicera as in Text-fig. 2B L. hilaris Koch’ (p. 130) — Coxa III with posterior seta normal ; tarsus IV with setae al, and uv, slender, pointed. Pilus dentilis setiform ; chelicera as in Text-fig. 3E L. echidnina Berlese (p. 136) Laelaps hilaris Koch Laelaps hilavis Koch, 1836, Deutschl. C.M.A.4:20. Tipton, 1960, Univ. Calif. Publ. Ent. 16: 271, figs. Female : Chelicera with segment I, 54 w; Il, 110 « ; movable digit 33 », biden- tate ; fixed digit unidentate ; pilus dentilis elongate with swollen tip. Four pairs of gnathosomal setae with c.s. about 60 » apart ; vp. 2 about 54 w apart. Deutoster- num with six transverse rows of denticles (I-2 per row) ; corniculi27 «long ; internal 3 Laelapsclethrionomydis Lange (see Bregetova, 1956) commonly occurs on the bank vole, Clethrionomys glaveolus (Schreber) in Europe and may possibly parastize this host in Britain. It is closely related to L. hilaris but may be distinguished from it by the larger number of shorter, more spinose setae of the opisthogaster of the female and by tarsus IV of the male having seta pug short and spine-like and setae ad2-3 and pde-3 long and blade-like. These tarsal setae are simple, setiform in the male of L. hilaris. elaps agilis Koch, female ; dorsum (A) and venter (B) of idiosoma. c—pD enter (Cc) and dorsum (Dp) of idiosoma IG. 1. A-B. Laelap Laelaps hilaris Koch, female ; venter (c) and 130 G. OWEN EVANS & We Mi TD ise malae with inner processes fringed, outer smooth (Text-fig. 2F). Tectum capituli with smooth anterior margin. Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (636 » * 456 ») with 39 pairs of setae arranged as in Text-fig. ID ; marginal setae longer than the others, especially in the opisthonotal region, and most of them are barbed. Surface of shield reticulated ; no hypertrichy ; two pairs of 7 and seven pairs of R setae on the cuticle, all with bifurcate tips or small barbs. Tritosternum with base 45 p, laciniae 102 p. Sternal shield (102 Xx 186 p) with three pairs of setae and two pairs of pores; metasternal setae on small platelets, their associated pores on the cuticle. Between si1, 80 » ; between sf1 and sf3, 100 p. Genital shield (144 » * 135 p) flask-shaped, with four pairs of setae. Between gen., 114 »; between Jv2, 80 p. Anal shield (105 4 * I14 p), paranal setae 66 yw, post- anal seta 96 w. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. Ic. Peritreme extends to posterior margin of coxa I; peritrematal shield free. Chaetotaxy of legs differs from normal pattern in having two postero-lateral setae on genu IV. On tarsi II and III, setae al,, pl, and av, are relatively stout, seta fv, is slender. Coxal setae as in Text-fig. 1c. Length/width (in yp) of leg segments : I 1 ry IV genu 66/57 7260 50/45 80/48 tibia 70/40 VAY 48/45 78/48 tarsus 100/36 90/38 105/36 150/40 Male: Chelicera with segment I, 48 »; I, 105 »; movable digit 50 pu (70 » with spermadactyl) ; both digits edentate ; pilus dentilis as in female, Four pairs of gnathosomal setae with c.s. about 48 » apart ; hyp.2 about 40 pw part. Deutosternum with six transverse rows of denticles (I-3 per row) ; corniculi 33 » long. Internal malae, tectum and pedipalps as in female. Dorsal shield (648 » x 456 ») with 39 pairs of setae distributed as in Text-fig. 26. Surface of shield reticulated ; no hypertrichy ; two pairs of and five pairs of R setae on the cuticle. Tritosternum with base 45 p, laciniae 108 ». Holoventral shield (516 w X 144 ph) with four pairs of setae and three pairs of pores in the sternal region ; six pairs of setae in the genito-ventral region. Between sfI, 30 »; between sf1 and sf3, 102 ux. Paranal setae 45 p, postanal seta 84 w. Peritreme extends to posterior margin of coxa I; peritrematal shield fused anteriorly with dorsal shield ; posterior part free. Chaetotaxy of legs asin female. Tarsi II and III with setae al,, ply, av,, and also AV», relatively shorter and stouter than the corresponding setae in the female. Tarsus IV has two short, stout, blunt setae, a/, and pv,. Coxal setae as in Text-fig. 2H. Length/width (in ,) of leg segments : I IME Ill IV genu 78/66 84/72 63/57 102/66 tibia 75/48 66/64 60/50 96/63 tarsus 102/36 90/40 102/36 147/42 THE BRITISH DERMANYSSIDAE (ACART) 131 HasitaT: Found mainly on species of the genus Microtus in Britain, Europe and the U.S.S.R. Also recorded from a weasel in Germany. Fic. 2. a. Laelaps agilis Koch, venter of male; B. Laelaps hilavis Koch, chelicera of male; c. L. agilis, tarsus IV of male; pb. L. hilarvis, chelicera of female; E. L. agilis, chelicera of male; F. L. hilavis female, venter of gnathosoma ; G. L. hilavis male, dorsum of idiosoma ; H. L. hilavis male, venter of idicsoma. 132 G. OWEN EVANS & W. M. TILL Laelaps agilis Koch Laelaps agilis Koch, 1836, Deutschl. C.M.A. 4: 19. Tipton, 1960, Univ. Calif. Publ. Entomology 16 : 265, figs. Laelaps festinus Koch, 1838, Deutschl. C.M.A. 24: 7. Laelaps hilavoides Oudemans, 1928, Ent. Ber. Amst. 7 : 375. Strandtmann, 1963, J. Kansas ent. Soc. 36: 5, figs. 17-18. Female: Chelicera with segment I, 66 »; II, 123 »; movable digit 33 yp, bi- dentate ; fixed digit unidentate ; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 50 w apart; hyp. 2 about 45 w apart. Deuto- sternum with seven transverse rows of denticles (I-2 per row) ; corniculi 27 p long ; internal malae with smooth margins. Tectum capituli with free margin rounded, smooth ; salivary styli conspicuous, extending a little beyond tips of corniculi. Pedipalp (2-5-6—14) with two-tined apotele. Dorsal shield (636 » * 456 ) with 39 pairs of setae distributed as in Text-fig. IA. Setae Z5 considerably longer and stouter than J5 or S5. Surface of shield reticulate ; no hypertrichy. Cuticle with two pairs of 7 and seven pairs of R setae. Tritosternum with base 57 p, laciniae 100 p. Sternal shield (123 » X 192 ») with three pairs of setae and two pairs of pores; metasternal setae situated on small shields, their associated pores on the cuticle. Between sf, 90 » ; between sti and st3, 114 w. Genital shield (147 » x 168 yp) flask-shaped, with four pairs of setae. Between gen., 126 «7; between Jv2, 72 w. Anal shield (110 w X 144 ») with paranal setae 27 », postanal seta 105 pw. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 1B. Peritreme extends to anterior third of coxa II; anterior part of peri- trematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern in that genu IV bears a postero- ventral seta and two postero-lateral setae (2—?, }—2). Femur I bears extremely long dorsal setae (ad, and fd,) ; on tarsus II, setae al, and #/,, and on tarsus II], setae al,_, are stout ; on tarsus IV setae al, and pv, are stouter than setae J, and az. Length/width (in ») of leg segments : I II Ill IV. genu 87/63 78/68 50/50 75/48 tibia 972/54 60/60 45/48 72/42 tarsus 90/40 84/42 114/42 153/36 Male : Chelicera with segment I, 50 »; II, 120 w; spermadactyl about 84 u long, separate movable digit not present (Text-fig. 2E). Four pairs of gnathosomal setae with c.s. about 48 papart ; hypf.2 about 36 papart. Deutosternum with six rows of denticles (1-2 per row) ; corniculi 33 » long, weakly sclerotized. Internal malae, salivary styli, tectum and pedipalps as in female. Dorsal shield (612 »# X 432 «) with 39 pairs of setae as in female. Tritosternum with base 42 p, laciniae 90 p. Holoventral shield (516 « X 162 «) with four pairs of setae and three pairs of pores in the sternal region ; five pairs of setae in the genito- ventral region. Between sf1, 72 »; between sfI and sf3, 114 «. Paranal setae I8 4, THE BRITISH DERMANYSSIDAE (ACARI) A. Dorsum of idiosoma of female ; Fic. 3. Laelaps echidnina Berlese soma of female; c. chelicera male; F. tectum capituli of female; G. venter of gnathosoma of female. 134 G. OWEN EVANS. & W. M. TILL postanal seta 70 » long. Chaetotaxy and sclerotization of opisthogaster as in Text- fig. 2A. Peritreme extends to posterior margin of coxa II; anterior part of peri- trematal shield not developed ; posterior part very short, free. Chaetotaxy of legs as in female. Setae al,, pl, and av, on tarsus II, setae al, and av, on tarsus III, short, stout and blunt. Tarsus IV has setae al, and pv, stout and blunt ; al, and al, stout and spine-like ; ad, and ad, very long, thick and sword- like (Text-fig. 2c). Femur I with setae ad, and fd, very long and thick. Length/ width (in ) of leg segments : I i Ill IV genu 80/70 80/66 63/57 87/60 tibia 66/56 60/63 50/54 80/54 tarsus 7540 80/45 96/42 147/42 Hapitat: Found on Apodemus flavicollis (Melchior) and Apodemus sylvaticus (Linn.) in Britain, Europe, Iceland and the U.S.S.R. L. hilaroides was recorded from a bat, Eptesicus serotinus (Schreber) in Germany. Laelaps echidnina Berlese Laelaps (Iphis) echidninus Berlese, 1887, Acari, Myr. Scorp. Ital. fasc. 39, no. I. Echinolaelaps echidninus : Strandtmann & Mitchell, 1963, Pacific Insects 5 : 547, fig. I. Laelaps pallidus Tragardh, 1931, Nat. Hist. Juan Fernandez 3 : 616, figs. 145-147. Laelaps berleset Fonseca, 1939, Mem. Inst. Butantan 12 : 104. Echinolaelaps echidninus vitzthumi Turk, 1950, Parasitology 40: 71, fig. 7. Echinolaelaps hiysti Turk, 1950 (nom. nov. pro L. echidninus Hirst, 1913, non Berlese, 1887), Parasitology 40 : 72, fig. 6. Echinolaelaps hirsti ceylonicus Turk, 1950, Parasitology 40 : 72, figs. 9-10. Echinolaelaps flavioi Tipton 1960, Univ. Calif. Publ. Ent. 16: 288. Female: Chelicera with segment I, 120 »; II, 228 w; movable digit 72 p, bi- dentate ; fixed digit unidentate (Text-fig. 3c) ; pilus dentilis long and slender. Four pairs of gnathosomal setae with c.s. about 96 “ apart; hyp.2 about 84 mw apart. Deutosternum with six transverse rows of denticles (2-3 per row) ; corniculi 54 / long; internal malae fringed (Text-fig. 3G). Tectum capituli with smooth anterior margin (Text-fig. 3F). Salivary styli conspicuous, reaching tips of corniculi. Pedi- palp (2-5—6-14) with two-tined apotele. Dorsal shield (1008 » 648 ») with the normal 39 pairs of setae distributed as in Text-fig. 34 ; no hypertrichy. Surface of shield reticulate ; with eleven pairs of R setae on the cuticle. Marginal setae on scutum and cuticle barbed. Tritosternum with base 84 p, laciniae 192 mw. Sternal shield (234 pu X 240 p) extends to middle of coxa III, with three pairs of setae and two pairs of pores. Metasternal setae situated on small shields, associated pores on cuticle. Between stI, 132 w; between stx and st3, 204 p. Genital shield (258 » X 216 p») greatly expanded behind coxae IV (maximum width 396 «), bearing four pairs of setae. ‘ ot yee 136 G. OWEN EVANS & W. M. TILL Between gen, 180 »; between Jv2, 276 w. Anal shield (168 » x 190 yp), paranal setae 72 pw, postanal seta 162 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 3B. Peritreme extends to posterior margin of coxa I; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern only in having two postero-lateral setae on genulIV. Setae on tarsi II-IV thick and spine-like ; the following setae are stout and blunt: al, and pl, on tarsus II, al,, on tarsus III, al, on tarsus IV. Length/width (in ») of leg segments : I II Ill IV genu 120/75 120/90 99/72 117/66 tibia 120/57 108/78 102/66 129/63 tarsus 165/48 150/54 192/50 270/50 Male: Chelicera with segment I, 75 »; II, 126 w; both digits edentate, movable digit 102 » ; spermadactyl as in Text-fig. 3E. Four pairs of gnathosomal setae with c.s. about 66 » apart; Ayp.2 about 72 » apart. Deutosternum, tectum and pedi- palps as in female ; corniculi longer and more attenuate. Dorsal shield (840 » * 492 ») with the normal 39 pairs of setae ; no hypertrichy. Surface of shield reticulated ; marginal setae on shield and cuticle barbed. Tritosternum with base 60 p, laciniae 150 w. Holoventral shield (684 » x 174 p) greatly expanded behind coxae IV, with four pairs of setae and three pairs of pores in the sternal region, six pairs of setae in the genito-ventral region (Text-fig. 3D). Between sti, 84 » ; between sz and s#3, 150 ». Paranal setae 72 » ; postanal seta 140 p. Peritreme extends to anterior margin of coxa II ; anterior part of peritrema- tal shield fused with dorsal shield from a point a little posterior to the tip of the peri- treme ; posterior part free. Chaetotaxy of legs as in female, but stout setae on tarsi II-IV are all pointed. Length/width (in ) of leg segments : Ik ime Ill IV genu 120/72 120/84 102/66 114/66 tibia 120/60 108/72 102/60 126/60 tarsus 174/54 144/50 180/48 246/48 Protonymph : Chelicera with segment I, 75 »; II, 135 »; movable digit 45 p, bidentate ; fixed digit unidentate. Four pairs of gnathosomal setae with c.s. about 66 » part, hyp.2 about 70 w apart. Corniculi 45 » long; deutosternum with six transverse rows of denticles (I—3 per row) ; internal malae and tectum as in female. Pedipalp (1-4—5—12) with two-tined apotele. Podonotal shield (444 » X 372 «) with eleven pairs of setae ; pygidial shield (192 w X 350 p) with eight pairs of setae, the marginal ones being barbed. Chaeto- taxy and sclerotization of mesonotal region as in Text-fig. 4A. THE BRITISH DERMANYSSIDAE (ACARI) 137 Tritosternum with base 60 p, laciniae 135 mw. Sternal shield (255 mw x 162 p) with three pairs of setae and two pairs of pores ; between sf1, 96 » ; between sti and st3, 102 pw. Anal shield (130 » X 123 ») with paranal setae 78 », postanal seta 144 p long. Peritreme extends at least to middle of coxa III (Text-fig. 48). Chaetotaxy of legs normal ; many of the setae stout and spine-like ; particularly long setae on femora I and II (distal dorsals) and on dorsal surface of genu, tibia and tarsus IV. Length/width (in p) of leg segments : I IT III 1G genu 96/78 96/84 78/66 84/60 tibia 100/57 84/72 72/60 84/54 tarsus 168/42 150/48 162/48 207/45 Deutonymph : Chelicera with segment I, 90 »; II, 168 »; chelae as in female, movable digit 54 ». Four pairs of gnathosomal setae with c.s. about 78 p apart ; hyp.2 about 78 » apart. Corniculi, deutosternum, internal malae and tectum as in protonymph. Pedipalps as in female. Dorsal shield (876 » X 504 ») with the normal 39 pairs of setae ; six pairs of R setae on the cuticle. Marginal setae on shield and cuticle barbed (Text-fig. 4c). Tritosternum with base 90 yp, laciniae 150 pw. Sternal shield (432 p x 168 yp) with four pairs of setae and three pairs of pores. Between sfI, 114 »; between sf1 and st3, 192 ». Anal shield (150 » * 156 yw) with paranal setae 75 pu, postanal seta 168 ». Peritreme extends to posterior third of coxa I (Text-fig. 4D). Chaetotaxy of legs as in female. Length/width (in yp) of leg segments : I II ih IV genu 108/70 105/78 87/66 100/60 tibia 114/60 90/72 90/63 108/54 tarsus 174/48 138/54 186/50 228/50 HaBirat : Cosmopolitan, chiefly on rodents of the genus Rattus. Also recorded from the genera Mus and Bandicota and from the tree-shrew, Tupaia glis (Diard). Laelaps muris (Ljungh) Acarus muris Ljungh, 1799, Nova Act. Reg. Soc. Sci. Uppsala 6: to. Laelaps muris: Tipton, 1960, Univ. Calif. Publ. Ent. 16 : 277, figs. Laelaps avvicola George, 1889, Science Gossip 25: 6, figs. 1-2. Laelaps crassipes Schrank, Oudemans, 1897, Tijdschr. Ent. 39 : 135. Laelaps microti Oudemans, 1916, Ent. Ber. Amst. 4 : 309. Laelaps agilis (Koch) Hirst, 1916, J. Zool. Res. 1 : 67. Female: Chelicera with segment I, 70 u; II, 110 »; movable digit 32 p, bi- dentate ; fixed digit unidentate, pilus dentilis inflated, elongate, tapering abruptly at the tip (Text-fig. 5E). Four pairs of gnathosomal setae with c.s. about 78 p apart, hyp.2 about 54 » apart. Deutosternum with six rows of deutosternal denticles (1-2 138 G. (OWEN EVANS: @°W. Me Tlie Fic. 5. Laelaps muris (Ljungh). a. Dorsum of idiosoma of female ; B. chelicera of male ; c. venter of idiosoma of female; D. venter of idiosoma of male; £. pilus dentilis of female ; F. tarsus IV of male, ventral view. THE BRITISH DERMANYSSIDAE (ACARI) 139 per row) ; corniculi 30 » long ; internal malae with inner processes fringed, outer smooth. Tectum capituli with free margin rounded, smooth. Salivary styli extend slightly beyond tips of corniculi. Pedipalp (2-5—6-14) with two tined apotele. Dorsal shield (660 » X 492 ) with 39 pairs of setae distributed as in Text-fig. 54. Most setae short and spine-like, setae 75 and S3-5 at least twice as long as seta /T. Surface of shield granular, weakly reticulate laterally ; no hypertrichy ; two pairs of y and ten pairs of RF setae on the cuticle. Tritosternum with base 60 p, laciniae 126 ». Sternal shield (144 » 222 pv) with three pairs of short, spine-like setae and two pairs of pores ; metasternal setae on small shields, associated pores on cuticle. Between sf1, 93 » ; between sf1 and s?3, 120 p. Genital shield (180 » x 174 p) flask-shaped with four pairs of setae ; between gen., 140 »; between Jv2, 84 ». Anal shield (96 » x 110 ») with paranal setae 63 p, postanal seta 168 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 5c. Peritreme extends beyond posterior margin of coxa I; anterior part of peri- trematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern in that genu IV has two postero- lateral setae, seta #/, being considerably shorter than p/, ; tarsi II-IV with setae ad, and pd,. Femur I with setae ad, and pd, stout and sinuous, the latter very long. Setae av,, al, and fl, on tarsi II and III stout and blunt. Setae al, and al, on tarsus III and the ventral and lateral setae on tarsus IV stout and spine-like. Length/ width (in ») of leg segments : I 18 III IV genu 90/70 90/70 60/54 102/54 tibia 75/58 66/63 50/50 go /50 tarsus 96/45 go/48 114/42: 153/42 Male: Chelicera with segment I, 48 »; II, 87 »; both digits edentate ; movable digit about 70 pu long, 84 » with spermadactyl (Text-fig. 5B). Four pairs of gnatho- somal setae with c.s. about 63 wapart ; hyp.2 about 48 wapart. Corniculi 33 » long ; salivary styli extend approximately to tips of corniculi. Deutosternum, internal malae, tectum and pedipalps as in female. Dorsal shield (636 » 462 «) with 39 pairs of setae ; one pair of y and six pairs of R setae on the cuticle. Tritosternum with base 45 yp, laciniae 114 ». Holoventral shield (546 » 180 p) with four pairs of setae and three pairs of pores in the sternal region ; five pairs of setae in the genito-ventralregion. Between sti, 78 «; between sit and st#3, 108 w. Paranal setae 36 pw, postanal seta 130 w. Surface of shield reticulate and granular. Chaetotaxy and sclerotization of opisthogaster as in Text- fig. 5D. Peritreme extends to posterior margin of coxa I; anterior part of peri- trematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs and form of setae as in female. In addition, seta av, on tarsus - ILis stout and blunt. Tarsus IV with setae fv, and pv, stout and blunt ; setae al,, ad, and ad, stout and blade-like (Text-fig. 5F), genu and tibia IV each with a stout blade-like seta al,. Length/width (in pu) of leg segments : ZOOL. 14, 5. 10 140 G. OWEN EVANS & W.M. TILL I II Il IV genu 90/63 90/78 66/60 114/66 tibia 7554 66/66 57/57 99/63 tarsus 93/44 84/45 105/45 140/45 HapitatT: Found in Britain, Europe and the U.S.S.R., mainly on Arvicola terrestris (Linn.). Also recorded from Mzucrotus arvalis (Pallas) and Talpa europaea Linn. Genus HYPERLAELAPS Zachvatkin Hyperlaelaps Zachvatkin, 1948, Parazit. Sb. 10: 61. Type: Tetragonyssus microti Ewing, 1933. Small, medium sclerotized mites ranging from 500 to 600 » in length. Chelicerae chelate-dentate in the female with the movable digit bidentate. Fixed digit in the male markedly reduced, edentate ; movable digit entirely fused with the extremely elongated spermadactyl. Pilus dentilis long, not inflated distally. Dorsal seta and fissures normal. Arthrodial processes at base of movable digit in female few in number and long. Chaetotaxy of venter of gnathosoma normal; chaetotaxy of pedipalp deficient (2—-4/5-6-13/14) ; apotele two-tined. Deutosternum with six or seven rows of denticles, one or two denticles per row. Corniculi horn-like. Tectum membranous, with rounded or lobate anterior margin, never denticulate. Dorsal shield entire and with 37 (px2 and x3 lacking) or 38 (fx3 lacking) pairs of setae. Majority of dorsal setae short and subspinose. Tritosternum normal, bipartite. No definite pre-endopodal shields. Sternal shield in female with three pairs of setae and two pairs of pores ; metasternal setae on distinct shields. Genito-ventral shield inflated posterior to coxae IV and bearing four pairs of setae. Anal shield truncate anteriorly with normal setation. Opistho- gastric cuticle with a maximum of twelve pairs of setae ; male with fewer setae than the female. Peritrematal shield poorly developed, free posteriorly. Metapodal shields present. Male with sternito-genito-ventral shield bearing ten pairs of setae, anal shield free. Podal shields weakly developed posterior to coxae IV. Chaetotaxy of legs normal except genu III (2—#, 3—1), tibia III in H. amphibia (I—t, 7-1) and genu IV (2—?, 3—2). Femora I and II in both sexes with ad, at least three times the length of fd,. Coxa II with anterior spine. Zachvatkin (1948) proposed Hyperlaelaps as a subgenus of Laelaps with Laelaps (Hyperlaelaps) arvalis Zachvatkin as the type. In recent revisionary works on this group (Strandtmann & Wharton, 1958 and Tipton, 1960) this subgenus has been synonymized with Laelaps. Detailed studies of the chaetotaxy of the species included in Hyperlaelaps have shown that they are not congeneric with the species of Laelaps. There is some confusion, at present, concerning the name of the type species. Koch (1839) described Laelaps pachypus from the field mouse (‘‘ Lemus arvalis”’), believing it to be conspecific with Acarus pachypus Hermann, 1804. The first bi | | THE BRITISH DERMANYSSIDAE (ACARI) 141 accurate description and figure of Laelaps pachypus sensu Koch may be attributed to Hirst (1916). Later Oudemans (1927), without reference to Hirst, re-described both sexes of what he considered to be Laelaps pachypus sensu Koch. Unfortunately two related species are confused in his description, the male (from Mzcrotus arvalis) being referable to Laelaps pachypus sensu Hirst, and the female (from Arvicola amphibius) to a species which has subsequently been described as Laelaps (Hyperlaelaps) am- phibius by Zachvatkin (1948). We have examined Oudemans’ material of L. pachypus and can confirm that these two species were represented in the material he had before him in 1927. In 1936 Oudemans correctly pointed out that Koch (1839) has misidentified Acarus pachypus Hermann and proposed a new name, Laelaps kocht, for Laelaps pachypus sensu Koch. On the basis of the type host and of the figures of pachypus by Hirst (1916) we consider L. (H.) arvalis Zachvatkin to be a synonym of Laelaps kochi Oudemans. Recently we have had the opportunity, through Dr. Crabill, Smithsonian Institution, of examining the “ cotype ”’ material of Tetragonyssus microti Ewing from Mzcrotus californicus, Los Angeles, California, and from Microtus sp., Alaska. We are unable to separate morphologically Ewing’s species from Laelaps kocht Oudemans taken from Microtus arvalis in Britain. We therefore consider L. kochi to be synonymous with T. microti. KeEy TO MALES AND FEMALES 1 Tibia III with chaetotactic formula (2—+, #—1). Palpfemur with five setae. Dorsal shield with setae px2 present or absent. In the female, seta s6 short and spinose, seta St relatively long and slender : H. microti (Ewing) (p. 141) - Tibia III with chaetotactic formula (1—+, 2—1). Palpfemur with four setae. Dorsal shield usually with setae px2 present. Inthe female, setas6and Str bothshort and spinose. . : : : : : H. amphibia Zachvatkin (p. 144) Hyperlaelaps microti (Ewing) Laelaps pachypus Koch, 1839, Deutschl. C.M.A. 24:8 (non Acarus pachypus Hermann, 1804). Hirst, 1916, J. Zool. Res. 1 (2) : 68, figs. 5-6. Oudemans, 1927, Tijdschr. Ent. 70: 179 (in part). Tetvagonyssus microti Ewing, 1933, Proc. U.S. nat. Mus. 82 (art. 30) : 9. Laelaps kochi Oudemans, 1936, Krit. Hist. Over. Acar. 3 (A): 244 (nom. nov. for L. pachypus Koch). [in part]. Laelaps (Hyperlaelaps) avvalis Zachvatkin, 1948, Pavazit. Sb. 10: 74, figs. 25-26 (syn. nov.). Female : Chelicera with segment I, 54 »; I, 140 »; movable digit 36 pw, biden- tate; fixed digit unidentate, pilus dentilis elongate, broad, not inflated distally (Text-fig. 7B). Four pairs of gnathosomal setae with c.s. about 66 » apart, hyp.2 about 48 » apart. Deutosternum with seven rows of denticles (1-2 per row) ; corniculi 42 » long; internal malae with smooth margins. Tectum capituli with free margin rounded or slightly lobed in appearance. Pedipalp (2-5-5-13) with two-tined apotele. Dorsal shield (588 . 444 4) with 37 pairs of setae distributed as in Text-fig. 64, the setae of the px series being absent, more rarely px2 present paired or unpaired. 142 G..OW-EN EVANS) & W.-M. “TLE ~ Surface of shield granular, reticulate laterally ; no hypertrichy ; six pairs of R setae on the cuticle. Most of the setae are short and spine-like ; those of the S series and seta Z5 are considerably longer. Seta s6 short and spine-like. Tritosternum with base 45 p, laciniae 110 pw. Sternal shield (70 » xX I9g2 p#) with © posterior margin deeply concave, lateral margins heavily sclerotized ; with three | Fic. 6. A-B. Hyperlaelaps microti (Ewing), female ; dorsum (a) and venter (B) of idiosoma. c-p. Hyperlaelaps amphibia Zachvatkin, female ; opisthonotal region (c) ; sternal and genito-ventral shields (pD). THE BRATISH DERMANYSSIDAE (ACARI) 143 pairs of setae and two pairs of pores. Setae 2 and 3 are short, with unevenly truncate tips which give the appearance of being broken ; metasternal setae situated on small shields, associated pores on cuticle. Between sf1, 70 » ; between sf1 and sf3, 117 wp. Genital shield flask-shaped (135 » 135 pm), with four pairs of setae, expanded behind the genital setae and reaching its maximum width (216 p) at level of Zvr. Between the short, truncate gen., 96 4; between Jv2, 50 w. Anal shield (70 » X 78 ») with maximum width (87 ») at anterior margin. Paranal setae 78 p, postanal seta 108 w long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 6B, the setae being situated on small projections of the integument. Peritreme extends to posterior margin of coxa 1; anterior part of peritrematal shield fused with dorsal shield ; posterior part not visible. The chaetotaxy of the legs differs from the normal pattern in three respects : genu III lacks seta pv ; genu IV has seta p/, ; tarsi II-IV lack setae ad, and pd,. Femur I with seta ad, at least three times as long as pd, ; femur II with seta ad, about three times as long as pd,._ Many of the leg setae are reduced to short spines. Trochanter I with pv, short and stout; genu I with ad, and fd, relatively long and thick ; trochanters III and IV with a/ and d stout ; femur IV with ad, long and stout ; tarsi II-IV with ventral and lateral setae relatively stout, tapering. Length/width (in ») of leg segments : I iar III IV genu 80/84 84/72 54/57 90/57 tibia 69/63 57/69 45/54 80/50 tarsus 78/45 96/45 93/42 144/42 Male: Chelicera with segment I, 42 »; II, 75 w; spermadactyl 150 »; movable digit not visible (Text-fig. 7c). Four pairs of gnathosomal setae with c.s. about 57 apart; Ayp.2 about 42 » apart. Corniculi about 40 p long, weakly sclerotized, membranous. Internal malae difficult to distinguish. Deutosternum, tectum and pedipalps as in female. Dorsal shield (504 » x 360 ) with 37 pairs of setae as in female ; seven pairs of R setae on the cuticle. Tritosternum with base 36 4; laciniae 90 ». Sternito-genito-ventral shield (324 pu x 150 ») with four pairs of setae and three pairs of pores in sternal region, six pairs of setae in genito-ventral region. Between sti, 54 »; between sti and st3, 102 uw. Anal shield (60 » x 70 ») with maximum width (80 ) at anterior margin. Paranal setae 48 4, postanal seta 75 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 7A. Peritreme extends about to anterior margin of coxa II ; anterior part of peritrematal shield fused with dorsal shield ; posterior part not developed. Chaetotaxy of the legs as in female. Length/width (in yp) of leg segments : I il Ul IV genu 78/72 75/63 50/56 80/50 tibia 60/57 54/63 42/48 75/45 tarsus 72,/42 80/42 84/40 132/40 144 G. OWEN EVANS & W. M. TILL Hapirat: Found in Britain on Mzcrotus arvalis (Pallas), M. agrestis (Linn.) and M. orcadensis Millais. Recorded from Europe and the U.S.S.R. from the genus Microtus, from Rattus rattus (Linn.), Apodemus sylvaticus (Linn.) and “ Mustela vulgaris ’’. Fic. 7. a-c. Hyperlaelaps microti (Ewing) ; venter of idiosoma of male (a) ; pilus dentilis of female (B); chelicera of male (c). Db. Hyperlaelaps amphibia Zachvatkin, female ; right pedipalp (trochanter to tibia), dorsal view. Hyperlaelaps amphibia Zachvatkin Laelaps (Hyperlaelaps) amphibius Zachvatkin, 1948, Parazit. Sb. 10: 74, figs. 22-24. Laelaps pachypus (Ikoch) Oudemans, 1927, Tijdschr. Ent. 70: 179 (in part). Female : Chelicera with segment I, 50 » ; II, 153 » ; movable digit 36 », bidentate ; fixed digit unidentate, pilus dentilis elongate, band-like, with pointed tip. Four pairs of gnathosomal setae with c.s. about 66 mw apart, hyp.2 about 54 4 apart. Deutosternum with six rows of denticles (I-2 per row) ; corniculi 30 nlong. Internal malae and tectum capituli as in H. microti. Pedipalp (2-4-5-13) with two-tined apotele (Text-fig. 7D). Dorsal shield (600 » 408 ») with 38 pairs of setae, px2 being present (Text-fig. 6c). Setae s6 and Sr both short and spine-like. Surface of shield granular, reticulate laterally ; seven pairs of & setae on the cuticle. Tritosternum with base 42 p, laciniae 117 w. Sternal shield (84 » X 192 ») with setae 2 and 3 longer and more pointed than in H. mzcrot: (Text-fig. 6D) ; between sfI, THE BRITISH DERMANYSSIDAE (ACART) 145 78 «; between sti and sf3, 123 w. Metasternal setae on small platelets, associated pores on cuticle, very close to sternal shield. Genital shield (150 w * 162 ) with maximum width (216 yp) at level of Zvr ; between gen., 102 «4; between Jv2, 60 uw. Genital setae (gen.) more pointed than in H. microti (Text-fig. 6D). Anal shield (60 » X 78 ») with maximum width (102 “) at anterior margin. Paranal setae 60 p, postanal seta 96 wlong. Peritremes and chaetotaxy and sclerotization of opisthogaster as in H. micrott. Chaetotaxy of legs as in H. microti except that tibia III bears only one antero- lateral seta. Length/width (in ») of leg segments : I ai III IV genu 90/75 90/84 54/63 90/54 tibia 66/60 60/72 40/54 84/45 tarsus 7245 93/48 87/45 150/42 Male : Chelicera with segment I, 42 »; II, 120 »; spermadactyl 168 », movable digit not visible. Four pairs of gnathosomal setae with c.s. about 63 pw apart ; hyp.2 about 48 » apart Corniculi about 54 pu long, weakly sclerotized. Internal malae difficult to distinguish. Deutosternum, tectum capituli and pedipalps as in female. Dorsal shield (552 » 396 ») with 38 pairs of setae as in female. Tritosternum with base 33 p, laciniae 90 ». Sternito-genito-ventral shield (372 » x 168 pw) as in H. microti. Between sti, 63 »; between sti and st3, 102 yw. Anal shield (63 pp x 75 ») with maximum width (go ») at anterior margin. Paranal setae 45 », postanal seta 90 » long. Chaetotaxy and sclerotization of opisthogaster as in H. micrott. Peritreme extends to middle of coxa II. Chaetotaxy of legs and form of setae as in female. Length/width (in ,) of leg segments : I II TT IV genu 78/72 84/70 60/63 80/48 tibia 66/63 54/66 45/54 80/42 tarsus 72148 84/45 90/45 135/38 HasitaTt: Recorded from Britain, Europe and the U.S.S.R. on Arvicola terrestris (Linn.) Genus ONDATRALAELAPS Evans & Till Ondatralaelaps Evans & Till, 1965, Bull. Brit. Mus. (nat. Hist.) Zool. 13: 269. Type : Laelaps multispinosa Banks, 1909. Medium sclerotized mites ranging from 650 » to 750 win length and with the general facies of Laelaps. Chelicerae chelate-dentate in the female with the broad movable digit bidentate and the shorter fixed digit unidentate. Fixed digit in the male markedly reduced, edentate, movable digit partially fused with elongate grooved G. OWEN EVANS & W. M. TILL — a ees i [— —_ thy ¢ es € € A. Dorsum of idiosoma of female; B. venter of idiosoma of female ; c. dorsum of idiosoma of male; D. venter of idiosoma of male. (Banks). Fic. 8. Ondatralaelaps multispinosa THE BRITISH DERMANYSSIDAE (ACARI) 147 _ spermadactyl. Pilus dentilis long, hooked distally. Dorsal seta and fissures normal. Arthrodial processes at base of movable digit present. Chaetotaxy of the venter of the gnathosoma normal; chaetotaxy of pedipalp deficient (2—-5-5-14) ; apotele two-tined. Deutosternum with six transverse rows of denticles, one to three denticles - perrow. Corniculi horn-like ; internal malae fimbriated. Tectum capituli tongue- shaped, margin smooth. Dorsal shield entire with typically 32 pairs of setae, podonotal region with 20 pairs . of setae, opisthonotal region with 12 pairs (73, 23, J/3, J/4, 23, px2 and x3 lacking in comparison with the typical form in Laelaps). Intraspecific variation of the opis- thonotal chaetotaxy common. Dorsal setae in the female shorter and stouter than in the male. Sexual dimorphism affecting form and number of marginal setae » (cand &) in the adult. Tritosternum normal, bipartite. No definite pre-endopodal shields. Sternal _ shield in the female with three pairs of setae and two pairs of pores, metasternal setae _ on discrete platelets. Genito-ventral shield typically Laelaps with four pairs of setae. Anal shield elongate-oval in outline, paranal setae situated behind the anus. Opis- thogastric cuticle with up to ten setae, variable. Peritrematal shields free posteriorly. Small metapodal shields present. Male with holoventral shield bearing nine pairs of setae excluding anals. Opisthogastric cuticle with about two pairs of setae. Podal - shields weakly developed posterior to coxae IV. Chaetotaxy of legs normal except for genu III (2—?, 2—1) and genu IV (2—?, 32). Coxa II with well-developed anterior spine. Femur I with ad, and pd, very stout and long, fd, arising from sclerotized protuberance (Text-fig. 9p). Certain _ setae on other segments tend to be stout. Ambulacra with claws reduced or absent. Two species are included in this genus, the type species occurring on the Musk Rat, | Ondatra zibethica (Linn.) in North America and Europe (introduced), and Ondatra- laelaps evansi (Tipton) comb. nov. from the Southern Musk Rat, Neofiber alleni True ) in Florida, U.S.A. Ondatralaelaps multispinosa (Banks) Laelaps multispinosus Banks, 1909, Proc. ent. Soc. Wash. 11:136; Bregetova, 1956, Opred. Faune SSSR. 61 : 105. Liponyssus spiniger Ewing & Stover, 1915, Ent. News 26 (3) : 109. Tetragonyssus spinigey: Ewing, 1922, Proc. U.S. nat. Mus. 62 (art. 13) : 10. Laelaps pavvanalis Willmann, 1952, Z. Pavasitenk. 15 (5) : 398. Female : Chelicera with segment I, 57 » ; II, 117 » ; movable digit 36 p, bidentate ; fixed digit reduced in size, with a terminal hook, pilus dentilis extremely elongate (Text-fig. gE). Four pairs of gnathosomal setae with c.s. about 66 w apart, hyp.2 about 60 » apart. Deutosternum with six transverse rows of denticles (one to three denticles per row) ; corniculi horn-like, 30 » long ; internal malae fimbriate (Text- fig. 9F). Tectum capitul? tongue-shaped with anterior margin rounded, smooth. Pedipalp (2-5-5-14) with two-tined apotele. 148 G. OWEN EVANS *&-W.M. TILL Dorsal shield (696 » x 432 ») with 32 pairs of setae distributed as in Text-fig. 84 ; 13, 23, J3, J4, Z3, px2 and px3 lacking. Chaetotaxy of opisthonotal region variable. Most of the setae short and spinose. Surface of shield granular. Tritosternum with base 48 p, laciniae 132 ». Sternal shield (150 » X 207 p) with three pairs of setae and two pairs of pores. Metasternal setae on distinct shields. Between si, 80 «4; between sti and s#3, 120 w. Genital shield (174 u X 126 p) flask-shaped, bearing four pairs of setae ; between gen., 105 4; between Jv2, 99 pu. Anal shield (140 p» X 93 p) elongate pear-shaped, with paranal setae situated posterior tothe anus. Paranals 18 » long, postanal seta 57 wp long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 8B. Peritreme extends to middle of coxa I. Peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs normal except genu III, which lacks seta pu (2—?, 2—1) and genu IV, which has two postero-lateral setae (2—?, 3—2). Both setae on coxa I and posterior setae on coxae II-III stout and spine-like ; coxa II with well-developed anterior spine. Many of the leg setae stout and spinose. Length/width (in ,) of leg segments : I II Ill IV genu 96/82 90/78 57/63 80/54 tibia 72/60 63/63 51/54 82/48 tarsus 90/48 105/50 105/45 147/42 Male : Chelicera with segment I, 40 » ; II, 66 » ; movable digit 48 » long, partially fused with spermadactyl. Fixed digit greatly reduced, bearing an elongate pilus dentilis ; both digits edentate (Text-fig. 9G). Four pairs of gnathosomal setae with c.s. about 57 » apart ; Ayp.2 about 48 w apart. Cornicul 30 p long ; other gnatho- somal structures asin female. _ Dorsal shield (612 » X 360 p) with 32 pairs of setae distriuted as in Text-fig. 8c, the setae being relatively longer and more slender than in the female. Unsclerotized integument of dorsum bears seven to eight pairs of setae, one pair at the posterior margin and six to seven pairs on the mid-lateral margin. Tritosternum with base 33 p, laciniae 66 ». Holoventral shield (528 p x 168 x) with nine pairs of setae, excluding the anals. Between sf1, 57 «; between sf and st3, 117 pw. Paranal setae 12 » long, postanal seta at least three times this length. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 8p. Peritreme extends to middle of coxa I; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs asin female. Stout, spine-like coxal setae relatively longer and more slender than in female. Tarsus IV of male with three stout, spur-like setae (Text-fig. 9B). Length/width (in p) of leg segments : i) Il Ill IV genu 90/75 975/75 60/63 90/60 tibia 72/60 54/60 50/54 88/54 tarsus 90/42 93/42 93/42 135/40 THE BRITISH DERMANYSSIDAE (ACARI) 149 Fic. 9. Ondatralaelaps multispinosa (Banks). A. Dorsum of idiosoma of deutonymph (male) ; B. tarsus IV of male; c. venter of idiosoma of deutonymph (male) ; Dp. femur I of male, dorsal view; £. chelicera of female; F. venter of gnathosoma of female ; G. chelicera of male. 150 G. OWEN EVANS & W. M. TIEL Deutonymph (male) : Chelicera with segment I, 36 »; I, 80 »; chelae as in female, movable digit 27 ». Four pairs of gnathosomal setae with c.s. about 54 pw apart ; hyp.2 about 45 » apart. Gnathosomal structures generally as in female. Dorsal shield (540 » x 228 ») with 16 pairs of setae in the podonotal region, 11-12 pairs in the opisthonotal region (Text-fig. 9A). Tritosternum with base 42 p, laciniae 100 »; Sternal shield (240 pw X 117 p) with four pairs of setae and three pairs of pores. Between sfI, 60 » ; between sf1 and st3, 110 p. Anal shield (96 » x 66 »); par- and postanal setae very short, about 8 » and 10 p long respectively. Opisthogastric cuticle with seven pairs of setae (Text-fig. 9c). Peritreme extends to middle of coxa I; peritrematal shields free anteriorly, poorly developed posteriorly. Chaetotaxy of legs as in female, but setae not so markedly enlarged. Length/ width (in ») of leg segments : I II 1881 IV genu 78/72 66/66 45/57 63/50 tibia 63/58 50/57 45/50 63/45 tarsus 90/45 84/40 96/40 120/36 Deutonymph (female) : Slightly larger than male deutonymph and with eleven pairs of setae on the opisthogastric cuticle. Chelicera with segment I, 45 »; II, 96 up; movable digit 36 ». Dorsal shield (588 » 240 p) ; sternal shield (252 » X 126 2) ; anal shield (96 » X 75 p). Hasitat: The musk rat, Ondatra zibethica (Linn.) in North America, Britain and Germany. Genus ANDROLAELAPS Berlese Androlaelaps Berlese, 1903, Zool. Anz. 27: 14. Haemolaelaps Berlese, 1910, Redia 6: 216; and 1916, Redia 12: 170. Atricholaelaps Ewing, 1929, Manual of External Parasites : 186. Eubrachylaelaps Ewing, 1929, Manual of External Parasites : 180. Ischnolaelaps Fonseca, 1936, Mem. Inst. Butantan S. Paulo 10: to. Cyclolaelaps Ewing 1933, Proc. U.S. nat. Mus. 82 (art. 30) : 5. Turkiella Zumpt & Till, 1953, An. Inst. Med. trop. Lisboa 10: 215. Zygolaelaps Tipton, 1957, J. Parasit. 43 : 367. Type: Laelaps (Iphis) hermaphrodita Berlese, 1887. Medium sclerotized mites ranging from about 400 » to 1,600 win length. Cheh- cerae usually chelate-dentate in the female with the movable digit bidentate ; occasionally this digit is edentate except for a terminal hook, and may have an additional cutting process with outwardly directed teeth. Fixed digit in the male markedly reduced, edentate ; movable digit usually edentate and partially fused with the elongated, grooved spermadactyl. Dorsal seta and fissures normal. Arthrodial THER RITISH DERMAN YSSIDAE (A CART) I51 processes present at base of movable digit. Chaetotaxy of venter of gnathosoma and pedipalps normal, apotele two-tined. Deutosternum with 6 rows of denticles, each row with from one to six denticles. Corniculi usually horn-hke, occasionally reduced ; internal malae fringed or fimbriated. Tectum capituli membranous, rounded anteriorly, never denticulate. Dorsal shield entire, usually with the basic 39 pairs of simple or barbed setae. This number may be increased by the addition of a pair of setae on the podonotal region (anterior accessory—ax) and/or a pair of setae (posterior accessory—px) on the opisthonotal region, giving patterns with 40 and 41 pairs of setae respectively. There may also be varying degrees of hypertrichy ranging from a few accessory setae between the / series to a dense covering of setae which completely obscures the basic pattern. The basic number of setae may be reduced by the absence of 23 or the px series, or one or more pairs of marginal setae may be situated on the unsclerotized integument. Tritosternum normal, bipartite. No definite pre-endopodal shields. Sternal shield in the female with three pairs of setae and two pairs of pores. Genital shield narrow or variously inflated, bearing one pair of setae. Anal shield pear-shaped or subtriangular with normal setation. Opisthogastric cuticle with variable number of setae. Peritrematal shields free posteriorly. Metapodal shields present. Male usually with holoventral shield not fused with peritrematal shield; bearing from seven to ten pairs of setae, excluding the anals. A few species have a sternito-genito- ventral shield and a separate analshield. Podal shields poorly developed posterior to coxae IV. Chaetotaxy of legs normal except genu IV which has two postero-laterals (2—7, 32). Insome pauonien species Cs patterns are found on tibia I (2—3, 3—2), genu III (2—2, ?—2) and tibia III (2—+, ?—2) and in one species genu IV is normal. Seta av, on ane II may be modified to form a stout, spur-like structure in both sexes, or in the male only, of certain species. Ambulacra of all legs with paired claws. Coxa II has a small anterior spine. Key To MALES AND FEMALES 1 Both sexes with pilus dentilis inflated basally, distal portion slender, curved (Text- fig. 128). Greatest width of genital shield in female at the level of the first pair (Zv1) of opisthogastric setae (Text-fig. 12C) : A. fahrenholzi (Berlese) (p. 156) — Both sexes with pilus dentilis slender, setiform (Text-fig. top). Greatest width of genital shield in female ed aa at levelof second pair (Jv1) of opisthogastric setae (Text-fig. 10B) : < : : ; A. casalis (Berlese) s. lat. iS) Dorsal shield 620 w—710 uw long in female, 466 u-485 w long in male ; seta Z5 slender, setiform, about one and one-half times as long as /5 (Text-fig. 118). Associated with birds and rodents . é : : : : : A. casalis s. str. (p. 152) — Dorsal shield 504 « long in female, 402 « long in male ; seta Z5 stout, subspiniform, scarcely longer than /5 (Text-fig. 11c). Associated with ants A. casalis myrmecophila ssp. nov. (p. 154) 152 G. OWEN EVANS: & W.-M: DIE Androlaelaps casalis casalis (Berlese) ?Gamasus fenilis Mégnin, 18761, J. Anat. Physiol. Paris 12 : 332. Iphis casalis Berlese, 1887, Acari, Myr. Scorp. Ital fasc. 38, no. 8, figs. 3-5. Androlaelaps casalis: Till, 1963, Bull. Brit. Mus. (nat. Hist.) Zool. 10 : 23, figs. 6-9, 25-29. Hypoaspis oculatus Oudemans, 1915, Arch. Naturgesch. (A) 81: 134. Hypoaspis soarianus Hull, 1925, Ann. Mag. nat. Hist. (9), 15 : 209 (syn. nov.). Haemolaelaps molestus Oudemans, 1929, Ent. Ber. Amst. 8: 13. Atricholaelaps megaventralis Strandtmann, 1947, Proc. ent. Soc. Wash. 49: 112. Hypoaspis freemani Hughes, 1948, Mites associated with stored food products : 129, figs. 173-179. Haemolaelaps haemorrhagicus Asanuma, 1952, Misc. Rep. Res. Inst. Nat. Res. no. 25 : 87, fig. r- Female : Chelicera with segment I, 63 » ; II, 108 » ; movable digit 36 pu, bidentate ; fixed digit with two or three teeth ; pilus dentilis slender, setiform (Text-fig. Ion). Four pairs of gnathosomal setae with c.s. about 60 » apart ; Wyp.2 about 42 p» apart. Deutosternum with six transverse rows of denticles (2-6 per row). Corniculi 36 p long, internal malae fringed. Tectum capituli with smooth anterior margin. Salivary styli conspicuous, not reaching tips of corniculi. Pedipalp (2-5—6-14) with two-tined apotele. Dorsal shield (620-710 » < 376-429 ») with the normal 39 pairs of simple setae and a variable number of unpaired accessory setae distributed as in Text-fig. Ioa. Surface of shield reticulate ; two pairs of v and seven pairs of RF setae on the cuticle. Tritosternum with base 54 p, laciniae 80 ». Sternal shield (100 » x 126 ») with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between sf1, 80 » ; between sf1 and s#3, 96 w. Genitalshield (177 w x 105 p) flask-shaped with one pair of setae. Anal shield (90 » x 102 u) with par- and post- anal setae about 42 w long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 10B. Peritreme extends to middle of coxa I ; peritrematal shield free post- eriorly ; anterior tip almost touches, but is not fused with, dorsal shield. Chaetotaxy of legs differs from normal pattern in having two posterolateral setae on genulV. Length/width (in ) of leg segments : iI II Ill IV genu 87/38 72/45 54/36 78/33 tibia 90/33 70/42 57/33 80/33 tarsus 147/30 120/30 120/30 162/27 Male: Chelicera with segment I, go w; II, 75 w; movable digit 40 p, 54 » with spermadactyl; both digits edentate (Text-fig. IoE). Four pairs of gnathosomal setae with c.s. about 48 w apart; hyp.2 about 33 w apart. Corniculi 27 » long; deutosternum, internal malae, tectum and pedipalps as in female. Dorsal shield (466-485 » X 276-322) with the usual 39 pairs of setae and three unpaired accessory setae. Surface of shield reticulated. Tritosternum with base 33 p, laciniae 66 ». Holoventral shield (384 u xX 95 4) with four pairs of setae and three pairs of pores in the sternal region, six pairs of setae 4It is not possible to be certain of the identity of this species from its short and inadequate original description. THE BRITISH DERMANYSSIDAE (ACART) e). Fic. oo — Reete oF aS oO ees Gy ones (2) a Es n°? ©), us 2a Ae) oY one fj go a8 .& ta LS cs “A 66 Te oo ~Y Os oO 2 > 18. 6 VU 8 Se 28 Sg n sa S o 6 Sa ~s 3 & ANS) So 2 § 8 eg NS 6 5 Be oO > of male. E. chelicera 154 G. OWEN EVANS & W.M. TILL in the genito-ventral region. Between sf1, 60 » ; between sti and sf3, 80 pw. Paranal setae 18 uw, postanal seta 27 » long. Chaetotaxy and sclerotization of opisthogaster _ as in Text-fig. roc. Peritreme extends to middle of coxa I; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : I iO Ill IV genu 72/36 70/45 50/33 70/33 tibia 78/30 60/40 50/30 72/30 tarsus 120/28 87/30 87/27 130/24 Habitat : Cosmopolitan, in the nests of birds, in poultry litter and also on rodents. Androlaelaps casalis myrmecophila ssp. nov. In allinstars this subspecies closely resembles Androlaelaps casalis s. str. from which it differs in certain of its measurements and in having setae Z5 at the posterior margin of the dorsal shield relatively short and stout (Text-fig. r1c). Female: Dorsal shield (504 » * 288 ») with the usual 39 pairs of setae and two te three unpaired accessory setae distributed as in Text-fig. 11a. Sternal shield (80 » x 105 ») not sharply demarkated from pre-sternal area. Between sif1, 72 w; between stt and st3, 78 wp. Genital shield (140 » x 84 pw) with genital setae only. Anal shield (72 # X 72 ») with par- and postanal setae subequal, about 30 p long. Chaetotaxy of legs as in A. casalis s. str. Tibia and tarsus II with relatively stout, spinose, ventral setae. Length/width (in ,) of leg segments : I I] MH IV genu 70/32 54/36 42/27 63/27 tibia 75/27 50/30 45/27 66/27 tarsus 130/26 84/24 90/22 120/22 Male: Gnathosoma as in A. casalis s. str. Dorsal shield (402 » X 222 p) with chaetotaxy and setal form asin female. Holoventral shield (324 » x 90 ») expanded behind coxae IV, with ten pairs of setae excluding the anals. Between s?fI, 57 p; between sii and sf3, 66 ». Par- and postanal setae about 20 pu long. Chaetotaxy of legs asin female. Length/width (in ») of leg segments : I Jit Ill IV genu 66/30 54/36 38/27 60/27 tibia 69/26 48/33 40/26 60/26 tarsus 110/24 75/24 75/21 108/21 THE BRITISH DERMANYSSIDAE (ACARI) 155 A | | Fie. 11. a. Andyolaelaps casalis myrvmecophila ssp. nov.; dorsum of idiosoma of female. B-C. Relative lengths of dorsal setae J/5, 75, and S5 of A. casalis casalis (B) and A. casalis myrmecophila (c). Protonymph : Chelicera with segment I, 36 »; II, 66 »; movable digit 22 uw. Four pairs of gnathosomal setae with c.s. about 45 » apart, Avp.2 about 33 p» apart. Corniculi 27 » long ; pedipalp (1-4-5-12) with two-tined apotele. | Podonotal shield (210 » x 186 ,) with eleven pairs of setae; pygidial shield (96 » X 156 «) with eight pairs of setae ; mesonotal region with four pairs of scutellae. _ Tritosternum with base 40 p, laciniae 54 . Sternal shield (about 144 p x 93 p) with anterior margin indistinct. Between sti, 63 «; between str and sf3, 102 wp. Anal shield (48 » x 60 4) with par- and postanal setae about 24 » long. Peritreme _extends to middle of coxa III. ZOOL. 14, 5. i 156 G. OWEN EVANS & W. M. TILL Chaetotaxy of legs normal. Length/width (in ,) of leg segments : I II III IV genu 54/33 45/36 33/27 45/28 tibia 54/30 42/33 33/27 48/27 tarsus 93/26 63/24 69/24 93/22 Deutonymph : Chelicera with segment I, 40 »; II, 68-70 »; movable digit 21 yp. Four pairs of gnathosomal setae with c.s. about 42 » apart, hyp.z about 30 p apart. Corniculi 24 » long ; other gnathosomal features as in female. Dorsal shield (348 » x 198 ) with lateral incisions and bearing the usual 39 pairs of setae and two or three accessories. Tritosternum with base 40 p, laciniae 54 ». Sternal shield (about 174 p x 72 p) with anterior margin indistinct ; bearing four pairs of setae and three pairs of pores. Between st1, 54 » ; between sti and st3, 78 uw. Anal shield (48 » x 45 »; maximum width 60 p); par- and postanal setae about 20 » long. Peritreme extends to posterior third of coxa I; peritrematal shield weakly developed. Chaetotaxy of legs as in female. Length/width (in p) of Jeg segments : I II III IV genu 54/30 45/34 30/27 48/27 tibia 57/26 42/30 33/24 48/24 tarsus 96/24 66/24 70/21 93/21 Hasitat: From the nests of Formica rufa Linn. Holotype 92 (1965 : 12 : 29: 7) and female paratype (1965 : 12 : 29 : 8) from the Black Wood of Rannoch, Perthshire (coll. R. M. Emberson, 13 April 1964) ; male allotype (1965: 12 : 29: 9), male and female paratypes (1965 : 12 : 29: 10), protonymph and deutonymph from Ufton Park, Berkshire (coll. C. A. Collingwood). The specimens are deposited in the collection of the British Museum (Natural History), London. Androlaelaps fahrenholzi (Berlese) Laelaps (Haemolaelaps) fahrenholzi Berlese, 1911, Redia 7 : 432. Liponyssus setiger Ewing, 1920, Ent. News 31: 290, figs. 6-7. Laelaps glasgowi Ewing, 1925, Proc. ent. Soc. Wash. 27: 6. Androlaelaps glasgow : Till, 1963, Bull. Brit. Mus. (nat. Hist.) Zool. 10: 40, figs. 65-68. Laelaps californicus Ewing, 1925, Proc. ent. Soc. Wash. 27: 5. Laelaps virginianus Ewing, 1925, Proc. ent. Soc. Wash. 27: 6. Hypoaspis microti Oudemans, 1926, Ent. Ber. Amst. 7: 101. Haemolaelaps mohrvae Oudemans, 1928, Ent. Bey. Amist. 7 : 374. Haemolaelaps cricetophilus Vitzthum, 1930, Zool. Jb. Jena, Abt. Syst. 60 : 417. Laelaps stegemani Hefley, 1935, J. Kansas ent. Soc. 8: 22. Ischnolaelaps rhabdomys Radford, 1939, Parasitology 31 : 240. Haemolaelaps scalopi Keegan, 1946, Trans. Amer. Micr. Soc. 65: 71. Atricholaelaps sigmodont Strandtmann, 1946, J. Parasit. 32 : 164. Atricholaelaps strandtmanni Fox, 1947, Ann. ent. Soc. Amer. 40 : 580. Hypoaspis (Haemolaelaps) eos Zumpt & Patterson, 1951, J. ent. Soc. S. Afr. 14: 79. THE BRITISH DERMANYSSIDAE (ACARI) 157 Female : Chelicera with segment I, 72 » ; II, 126; movable digit 36 p, bidentate ; fixed digit bidentate ; pilus dentilis inflated basally, distal portion slender, curved (Text-fig. 12B). Four pairs of gnathosomal setae with c.s. about 72 » apart ; hyp.2 about 54 «apart. Deutosternum with six transverse rows of denticles (3-5 per row) ; corniculi 30 » long ; internal malae fringed. Tectum capituli with smooth anterior margin. Salivary styli slender, extending to tips of corniculi. Pedipalp (2—-5—6—14) with two-tined apotele. Dorsal shield (714 » 432 ») with the normal 39 pairs of setae distributed as in Text-fig. 124; some anterior setae barbed. Surface of shield reticulated; no hypertrichy in British specimens ; with two pairs of y and seven pairs of & setae on the cuticle. Fic. 12. Androlaelaps fahrenholzi (Berlese), female ; dorsum of idiosoma (A) ; chelicera (B) ; venter of idiosoma (Cc). | Tritosternum with base 45 p, laciniae 135 w. Sternal shield (96 » x 168 yp) with _ three pairs of setae and two pairs of pores. Metasternals free with associated pores. Between st1, 96 » ; between sf1 and s#3, 102 pw. Genital shield tongue-shaped (156 pw _X 114 p), with genital setae only. Anal shield (90 » x 112 ») with paranal setae 54 p, postanal seta 66 ». Chaetotaxy and sclerotization of opisthogaster as in Text- _fig.12c. Peritreme extends to middle of coxa I ; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern jn having two postero-lateral setae on genu IV. Length/width (in yp) of leg segments : 158 G. OWEN EVANS & W.M. TILL if el Ill IV genu 96/45 go /60 70/45 105/45 tibia 93/36 80/57 70/42 110/42 tarsus 162/36 126/42 144/36 240/36 Male: Described and figured by Till (1963). Hasitat: Cosmopolitan, chiefly on rodents and in their nests. Also recorded from birds, bats and small carnivores. Genus HYPOASPIS Canestrini Hypoaspis Canestrini, 1884, Aiti. Ist. Veneto (6) 2: 1569; 1885, Acarvofauna ital. part 1: 55. Type: Gamasus krameri Canestrini, 1881. Chelicerae typically chelate-dentate with the movable digit bidentate in the female, rarely digits edentate or specialized for egg-feeding in certain myrmecophilous forms. Pilus dentilis normally short, setiform ; dorsal seta and fissures present. Chelicerae of male chelate, movable digit usually unidentate, spermadactyl] free, distally grooved and never completely fused with the movable digit. Arthrodial processes simple, rarely absent. Chaetotaxy of the venter of the gnathosoma and of the pedipalps normal, apotele two or three-tined, the latter with subequal tines. Deutosternum with six, rarely five, transverse rows of denticles. Corniculi horn- like, salivary styli present. Tectum capituli with anterior margin smooth or den- ticulate. Entire dorsal shield typically with 39 pairs of setae, px2 and px3 being present. Variations in the basic pattern are the result of absence of px2 and px3 (37 pairs), v2 and 74 on lateral integument (37 pairs), 73 and v1 absent (37 pairs), presence of extra v seta (40 pairs) or extra v seta and px (41 pairs). Unpaired accessory setae in the region of the J series may be present or absent. Tritosternum normal, bipartite. Pre-endopodal shields occasionally present. Sternal shield in the female usually with three pairs of setae and two pairs of pores ; first pair of setae may be placed on the pre-sternal area (Cosmolaelaps group). Genital shield flask-shaped with one pair of setae, or, genito-ventral shield variously inflated and bearing one to three pairs of opisthogastric setae situated on its lateral margins. Anal shield usually pear-shaped or subtriangular, with normal chaetotaxy. Opisthogastric cuticle with variable number of setae. Peritrematal shields free posteriorly or fused with podal shields. Metapodal shields present. Male usually with holoventral shield which may or may not be fused with the peritrematal shields, and which bears 9 or 10 pairs of setae excluding the anals. Anal shield occasionally free. Podal shields poorly or well developed posterior to coxae IV. Chaetotaxy of legs usually normal. Variations result from addition of seta fl, on tibia III (2—1, 2—2), seta pl, or pu on genu IV (2—?, 3—2 or 2—?, 3—1) and the absence of al, on genu IV (1—?, 3—1). Anterior spine on coxa II small] or absent. Ambulacra usually with paired claws ; claws on leg I occasionally reduced THE BRELISH DERMANYSSIDAE: (A CART) 159 -or absent. A ventralseta on femur II in the male, or both sexes, sometimes enlarged | | } and spine-like. The present classification of the Hypoaspis complex is largely based on palaearctic forms and requires radical revision. It appears to be well represented in the nearctic, tropical and subtropical regions and we feel that a knowledge of the extra-palaearctic forms is an essential prerequisite to any attempt at a “ generic ’’ classification of the complex. According to the current classification of Hypoaspis, the majority of the British species may be distributed among the following of its subgenera : HY POASPIS s. str. (type: Gamasus kramert G. & R. Canestrini, 1881). H. kvameri (Canestrini). ALLOPARASITUS Berlese, 1920 (type: Laelaps (Hypoaspis) oblongus Halbt., 1915) syn: Halbertia, Hull in Turk & Turk, 1952. H. oblonga (Halbt). COSMOLAELAPS Berlese, 1903 (type: Laelaps claviger Berlese, 1883). H. claviger (Berlese); H. cuneifer (Michael); H. vacua (Michael); H. neo- cunetfer sp. Nov. GAEOLAELAPS Tragardh, 1952 (type: Laelaps aculeifer Canestrini, 1884). H. aculeifer (Canestrini) ; H. giffordi sp. nov.; H. lubrica Voigts & Oudemans ; H. praesternalis Willmann and (?) H. sardoa Berlese. GYMNOLAELAPS Berlese, 1903 (type: Laelaps myrmecophilus Berlese, 1892). H. myrmecophila (Berlese) ; H. laevis (Michael) and H. myrmophila (Michael). HOLOSTASPIS Kolenati, 1858 (type: Holostaspis isotricha Kolenati, 1858). H. isotricha Kolenati; H. montana (Berlese) and H. oophila (Wasman). LAELASPIS Berlese, 1903 (type: Iphis astronomicus Koch, 1839). H. astronomica (Koch) ; H. humerata (Berlese) and H. equitans (Michael). PNEUMOLAELAPS Berlese, 1920 (type: Iphis bombicolens Canestrini, 1885). H. bombicolens (Canestrini) ; H. marginalis Willmann ; H. breviseta sp. nov. ; H. colomboi sp. nov. ; H. hyatti sp. nov. and H. minutis- sima sp. Nov. STRATIOLAELAPS Berlese, 1916 (type: Laelaps (Iphis) miles Berlese, 1882). syn. Davisiella Zumpt & Patterson, 1951 (Syn. nov.) H. miles (Berlese). Hypoaspis heselhausi Oudms. and H. nidicorva sp. nov., and H. acuta (Michael) appear to represent distinct species-groups. There is little doubt that certain of these subgenera represent natural bio-morpho- logical groups, although the boundaries between them cannot be satisfactorily defined on the basis of their morphological characteristics alone, even in such apparently distinct groups as Pneumolaelaps, Holostaspis and Gymnolaelaps.’ In view of this we are disregarding the subgeneric divisions in the following key to the British species of the complex. 5 There are, at present, no comprehensive definitions or keys to the subgenera of Hypoaspis. The placing of species within the subgenera has largely been by rule of thumb, generally following Berlese’s (1903) concepts. The following key, based on eco-morphological characteristics provides a guide to the subgenera or species-groups represented in the British fauna. (Footnote continued on p. 160) 160 G. OWEN EVANS & W. M. TILL KrEy TO FEMALES 1 Genu IV with eight setae, an antero-lateral seta lacking (I—?, 3—1) ; peritrematal shields free posteriorly and extending well beyond posterior margins of coxae IV (Text-fig 46F) 5 : : : : : : H, oblonga (Halbt.) (p. 222) - Genu IV withg or 10 setae, twoantero-lateral setae present (2—2, 3—1 ; 2—#2, 3—1 or 2—2,3—2); peritrematalshield not extending PESO to coxae IV, free or fused with the podal shields ; : é Dorsal shield lacking setae re on the oniethonotal region and never vit ihpaived accessory setae ; dorsal setae leaf-like, spatulate (Text-fig. 45a) ; corniculi long, slender, extending almost to the level of the anterior margin of the palpfemur H. miles (Berlese) (p. 222} - Dorsal shield with setae px present, with or without unpaired accessory setae; dorsal setae various ; corniculi relatively shorter, if reaching beyond the level of the middle of the palpfemur then dorsal setae simple . : é : é . 3 bo nN (Footnote continued from p. 159) 1 Genu IV with eight setae (1—?, 3—1) ; peritrematal shield in the female free posteriorly and extending well beyond the posterior margins of coxae IV (Text-fig. 46F) Alloparasitus Berlese — Genu IV with nine or ten setae ; peritrematal shield in the female not extending ea to coxae IV, free or fused with podal shields . ‘ 2 Dorsal shield lacking setae px in the opisthonotal region ‘and never with unpaired accessory setae ; dorsal setae leaf-like, spatulate. Corniculi long, slender, extending almost to the level of the anterior margin of the palpfemur nN Stratiolaelaps Berlese — Dorsal shield with setae px present, with or without unpaired accessories; dorsal setae various. Corniculi relatively shorter, if ee to the middle of the palpfemur then dorsal setae simple . : : ; : ; : : : 3 3. Dorsal setae spatulate, cuneiform « or scimitar- like. " Free-living or myrmecophilous Cosmolaelaps Berlese — Dorsal setae simple or pilose, never markedly flattened 4 Certain setae of the dorsum and of the segments of the legs (femur II, femur and tarsus IV) long and whip-like. Associated with Coleoptera Hypoaspis s. str. — Setae of the dorsum rarely, and legs” NSS long and whip-like : : : é 2 5 5 Genu IV with two ventral setae (2—?, #—1) ; peritremes usually broad ; lateral and ventral cuticle often densely setose. In nests of Bombus Pneumolaelaps Berlese — Genu IV with one ventral seta age %—1/2), if with two ventral setae, then peritremes narrow : 3 : : : : . : : : 6 6 Apotele with three subequal tines. Discrete, well-sclerotized pre-endopodal shields present ; dorsal shield with 40-41 pairs of setae (excluding unpaired accessories). _Myrmecophilous Gymnolaelaps Berlese — Apotele two-tined. Usually without discrete pre-endopodal shields ; dorsal shield with not more than 39 pairs of setae (excluding unpaired accessories) . 7 Digits of the chelicerae in the female stumpy, dentate or edentate and modified for oophagy, movable digit about 20 w in length. Mvyrmecophilous Holostaspis Wolenati — Digits of the chelicerae in the female normal in form, dentate or edentate ; movable digit at least 35 « in length 8 Genito-ventral shield in the female markedly inflated posterior to coxae IV, ” flanked by linear metapodals and characteristically ornamented. Dorsal setae long, whip- -like, or marginal setae serrated. Myrmecophilous or free-living 8 Laelaspis Berlese - Genital or genito-ventral shield otherwise ; dorsal setae never long or whip-like. Free-living or inhabitants of nests of birds and mammals Gaeolaelaps Trigardh (including heselhausi-nidicorva group) Hiln 13 14 THE BRITISH DERMANYSSIDAE (ACARI) 161 Dorsal setae 72-3, 22-4, S4—5 and Z4 considerably longer than other dorsal setae, 274 very long and whip-like (Text-fig. 134) ; seta pd, on femur II, ad, on femur IV, ad,_, and pd, on tarsus IV long and ee ogy Associated with Lucanus (Coleoptera) . ; H. krameri (Canestrini) (p. 163) Dorsal chaetotaxy otherwise ; fee setae never long and whip-like : : Dorsal setae spatulate (Text- ae 24C), cuneiform soe fig. 25C) or scimitar-like (Text-fig. 28a—c) . Dorsal setae simple or pilose, never markedly flattened Majority of dorsal setae cuneiform (Text-fig. 25c) . Majority of dorsal setae spatulate or scimitar-like. : : : : : Genu IV with two postero-lateral setae (2—%, 3—2) dorsal seta S2 clavate, considerably shorter than cuneiform seta Sr (Text-fig. 254) ; proximal region of cuneiform seta only slightly enlarged (Text-fig. 25c) . H. cuneifer (Michael) (p. 184) Genu IV with one postero-lateral seta (2—#?, 2—1) ; dorsal setae S1—2 cuneiform, subequal (Text-fig. 26a) ; proximal region of cuneiform seta conspicuously enlarged (Text-fig. 26D). S : : : . HH. neocuneifer sp. nov. (p. 187) Dorsal setae spatulate, clavate (Text-fig. 24c) ; peritremes extending to the level of the vertical setae (71) ; sternal shield withstrong reticulate pattern (Text-fig. 24D) H. claviger (Berlese) (p. 182) Dorsal setae scimitar-like (Text-fig. 28A—c) ; peritremes extending to the level of the middle of coxae I ; sternal shield without conspicuous ornamentation (Text-fig. ND Oun 271B)\ es : H. vacua (Michael) (p. 190) Genu IV with no ventral pes $e teste C= = I) : : : ; : 9 Genu IV with one ventral seta, pv absent (2—?, 3—1 or 2—#, 3—2) 16 Capitular setae and setae on coxae I—II stout, spinelike. Associated ath Bombus (Hymenoptera) . ; H fuscicolens (Oudemans) (p. 196) Capitular setae and setae. on coxae ie II slender, poral : : 10 Dorsal shield shows hypertrichy between the J series ; surface of shield nn a clearly defined reticulate pattern : II Dorsal shield without hypertichy between. the J series ; surface of shield tn or without a distinct reticulate pattern : : : : é : 14 Sternal shield distinctly wider than long (L/W = o- ae ; deutosternum with six rows each consisting of 2-8 unequally sized denticles. : : : 12 Sternal shield about as wide as long (L/W = 1:0) or longer than idle (L/W = == Tn) < deutosternum with six rows each of at least six small, uniform, denticles. é 13 Opisthogaster densely setose, region between genital and anal shields with about twenty setae (Text-fig. 348) ; ratio of distance between genital setae to the maximum width of the genital shield about 1-15 H. marginalis (Willmann) (p. 199) Opisthogaster not densely setose, region between genital and anal shields with four setae (Text-fig. 33B) ; ratio of distance between genital setae to maximum width of genital shield about 1-3. : H. hyatti sp. nov. (p. 198) Dorsal shield about 700 y in length ; setae Up extending at least three-quarters of the distance between j2 and 73 ; setae on distal third of tarsus II distinctly thickened, spine-like. Associated with nests of birds and small mammals H. lubrica Voigts & Oudemans (p. 172) Dorsal shield about 500 win length ; setae j2 extending to about one-half the distance between 72 and 73 ; setae on distal third of tarsus IT not distinctly thickened nor spine-like. Associated with Bombus : H. minutissima sp. nov. (p. 201) Sternal shield longer than wide (L/W = 1:1). " Opisthogastric setae subequal in length. Peritreme relatively narrow ; length of posterior part of peritrematal shield at least three times the width of the peritreme (Text-fig. 31B) H. colomboi sp. nov. (p. 194)