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. The history of the collections contained in the Natural History Departments of the British Museum. 1. Trustees British Museum, London: 442 pp. Bertin, L. 1944. Mise au point sur quelques espeéces de clupéidés. Bull. Soc. zool. France, 66 : 18-25. BLEEKER, P. 1865. Notice sur les poissons envoyés de Chine au Musée de Leide par M. G. Schlegel. Ned. Tijdschr. Dierk. 2: 55-62. 1873. Memoire sur la faune ichthyologique de Chine. Ibid. 4: 113-154. BoEsSEMAN, M. 1947. Revision of the fishes collected by Burger and von Siebold in Japan. E. J. Brill, Leiden : 242 pp. BripDGMAN, E.C. 1841. Chinese Chrestomathy in the Canton dialect. 5S. W. Williams, Macao : 698 pp. Cuan, W. L. 1965. A systematic revision of the Indo-Pacific clupeoid fishes of the genus Sardinella (Family Clupeidae). Jap. J. Ichthyol. 12: 104-157; ibid, 13: 1-39. CHEN, T-R. 1961. Contributions to the fishes from Quemoy (Kinmen). Quart. J. Taiwan Mus. 13 (3-4) : 191-213. Cuu, K-Y. & Tsar, C-T. 1958. 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. Bosin, G. & PRENANT, M. 1961. JRemarques sur certaines “‘ Hincksinidae ’”’, Alderinidae et Flustridae (Bryozoaires Chilostomes). Cah. Biol. Mar., 2: 161-175. Brown, D. A. 1949. On the Polyzoan Genus Hippomenella Canu & Bassler and its Genotype Lepralia mucronelliformis Waters. J. Linn. Soc. (Zool.), 41: 513-520. 1952. The Tertiary Cheilostomatous Polyzoa of New Zealand. British Museum (Nat. Hist.), London. 1954. On the Polyzoan Genus Crepidacantha Levinsen. Bull. Brit. Mus. (N.H.) Zool., 2, 7: 243-263. 1958. Fossil Cheilostomatous Polyzoa from South-West Victoria. Mem. Geol. Surv. Victoria, 20, 7-83. Buce, E. 1957. Les Bryozoaires du Néogeéne de l’ouest de la France. Mém. Mus. Nat. Hist. Nat., sér. C., @ : 1-436. Busx, G. 1852. Catalogue of Marine Polyzoa in the ... British Museum, Part I, London. 1854. Op. cit., Part IT. 1856. Zoophytology. Quart. J. micr. Sci., 4: 308-312. 1884. Report on the Polyzoa. The Cheilostomata. Rep. Zool. Chall. Exp., 10, pt. 30. CatvetT, L. 1896. Résultats scientifiques de la Campagne du ‘‘ Caudan’”’ dans le Golfe de Gascogne... Ann. Univ. Lyon, 26 : 251-270. Canu, F. 1916. Les Bryozoaires fossiles des terrains du Sud-Ouest de la France, IX. Aqui- tanien. Bull. Soc. géol. Fr., sér. 4, 15 : 320-334. —— 1926. Contributions a l’étude des Bryozoaires fossiles. Jbid., 25 : 763-766. [For date of publication see Canu, F. 1926. Bryozoaires Bartoniens du Bassin Franco-Belge. Ibid. : 741] Canu, F. & BassLter, R.S. 1917. A Synopsis of American Early Tertiary Cheilostome Bryo- zoa. Bull. U.S. nat. Mus., 96: 1-87. 1920. North American Early Tertiary Bryozoa. Jbid., 106 : v-xx, 1-879. —— 1925. Contribution a l’étude des Bryozoaires d’Autriche et de Hongrie. Bull. Soc. géol. Fry., sér. 4, 24: 672-690. [For date of publication see p. 689.) 1927. Classification of the Cheilostomatous Bryozoa. Proc. U.S. nat. Mus., 69, 14 : I-42. 1929. Bryozoa of the Philippine Region. Bull. U.S. nat. Mus., 9, 100 : 1i—xi, I-685. Canu, F. & LECOINTRE, G. 1925. Les Bryozoaires Cheilostomes des Faluns de Touraine et d’Anjou. Mém. Soc. géol. Fr., (n.s.) 2, 3: 1-18. 1930. Op. cit., Ibid., (n.s.) 6, 1 : 83-130. CIpoLia, F. 1921. I briozoi pliocenici di Altavilla presso Palermo. G. Sci. nat. econ. Palermo, 32 : 163-337. Cook, P. L. 1964. Polyzoa from west Africa. I. Notes on the Steganoporellidae, Thalamo- porellidae and Onychocellidae (Anasca, Coilostega). Ann. Inst. Océanogr. (Calypso, VI), 41 : 43-78. —— 1965. Polyzoa from West Africa. The Cupuladriidae (Cheilostomata, Anasca). Bull. Brit. Mus. (N.H.) Zool., 13, 6 : 189-228. FiscHerR, P. 1870. Bryozoaires, Echinodermes et Foraminiféres marins du Département de la Gironde. Act. Soc. linn. Bordeaux, 27 : 1-70. GauTIER, Y. V. 1962. Recherches écologiques sur les Bryozoaires Chilostomes en Méditerranée occidentale. Théses présenteés a la Faculté des Sciences de Il Université @’Aix-Marseille, 91 : 9-434. OBSERVATIONS ON TYPE-MATERIAL, POLYZOA a7 Gray, J. E. 1828. Spicilegia Zoologica ... part 1, London. 1848. List of the Specimens of British Animals in the... British Museum, part 1, Centroniae ov Radiated Animals, London. HaGENnow, F. von. 1839. Monographie der Riigenschen Kreide-Versteinerungen. Leonhard und Bronn’s Neues Jahrbuch. f. Min., 1839 : 253-296. HarMER, S. F. 1900. A Revision of the Genus Steganoporella. Quart. J. micr. Sci., 43 : 225- 297. 1902. On the Morphology of the Cheilostomata. Jbid., 46 : 263-350. —— 1926. The Polyzoa of the Siboga Expedition. Pt. 2, Cheilostomata, Anasca. Rep. Siboga Exp., 28b : 181-501. 1957. Op. cit. Pt. 4, Cheilostomata Ascophora II. Jbid., 28d : 641-1147. Hastincs, A. B. 1930. Cheilostomatous Polyzoa from the Vicinity of the Panama Canal. Proc. zool. Soc. Lond., 1929 : 697-740. 1943. Polyzoa (Bryozoa) I. “ Discovery’ Rep., 22 : 301-510. 1945. Notes on Polyzoa (Bryozoa).—II. Membranipora crassimarginata auctt., with Remarks on some Genera. Ann. Mag. nat. Hist. (11) 12 : 70-103. 1947. Op. cit—III. Onsome species of Cellavia ... Ibid., 13 : 217-241. 1949. On the Polyzoan Cellepora pulchva Manzoni and the genus Hippopleurifera. J. Linn. Soc. London (Zool.), 41 : 521-528. 1964. The Cheilostomatous Polyzoa Neoeuthyrvis woostevi (MacGillivray) and Reginella doliaris (Maplestone). Bull. Brit. Mus. (N.H.) Zool., 11, 3 : 245-262. Hincxs, T. H. 1880a. A History of the British Marine Polyzoa: i-cxli, 1-601, London. 2 vols. 1880b. Contributions towards a General History of the Marine Polyzoa. II. Foreign Membraniporina. Ann. Mag. nat. Hist. (5) 6: 81-92. —— 1881a. Op. cit., VI. Polyzoa from Bass’s Straits. Jbid., (5) 8: 1-14, 122-129. —— 1881b. Op. cit., VII. Foreign Membraniporina (third series). Jbid.: 129-132. —— 1882. Op.cit., TX. Foreign Cheilostomata (Miscellaneous). Jbid. (5) 9: 116-127. —— 1892. Op. cit., Appendix. Ibid., (6) 9: 327-334. 1895. [Op. cit.], Index: i-vi, London. Jounston, G. 1847. A History of the British Zoophytes, Ed. 2, 2 vols., London. Lacaaty, R. 1952. 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 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) 15 16 i], 24 25 G. OWEN EVANS & W. M. TILL Sternal shield wider than long (L/W = 0-6). Opisthogastric setae flanking genital shield about twice as long as those near the margin of the body. Peritreme relatively wide ; length of posterior part of peritrematal shield not more than twice the width of the peritreme : : : : : . 15 Setae on anterior part of dorsal shield rebels long ; seta 73 reaches base of seta 74 (Text-fig. 294) ; surface of shield with distinct reticulate pattern in addition to sculptured areas for muscle attachment . 6 H. bombicolens (Canestrini) (p. 191) Setae on anterior part of dorsal shield relatively short ; seta 73 extends about half- way to base of seta 74 ; surface of shield without a distinct reticulate pattern H. breviseta sp. nov. (p. 193) Genu IV with two postero-lateral setae (2—?, 2—2) (Text-fig. 44Cc) : ; : 17 Genu IV with one postero-lateral seta (2—2, 2—1) (Text-fig. 448). ; . 19 Antero-ventral seta on femur II and genu II stout and spine-like (Text-fig. ea) : tibia III with two postero-lateral setae (2—+, #—2) ; dorsal shield with 39 pairs of setae (Text-fig. 20a) ; : 3 : H. sardoa Berlese (p. 175) Antero-ventral seta on femur IT and genu Ir not eopenietonely thickened ; tibia III with one postero-lateral seta (2—+, 7?) ; dorsal shield with 37 or 41 pairs of setae. Myrmecophilous : 18 Dorsal shield about 1,100 yw in length ern ai pairs of shore fine: dorsal Bete ext: fig. 43D) ; apotele with three subequal tines. : H. laevis (Michael) (p. 220) — Dorsal shield about 700 yw in length with 37 pairs of medium, relatively stout, dorsal setae (Text-fig. 41A) ; apotele with two subequal tines H. acuta (Michael) (p. 212) — Peritrematal shield fused with the expodal shield in the region of coxa IV. é : 20 Peritrematal shield free posteriorly . : 2 Dorsal setae short, setae J1 reaching about mid- aes bear een the pases ef uf fa and fe 2 (Text-fig. 21a) ; genito-ventral shield sub-rectangular in outline (Text-fig. 21D) with a pair of setae lying between the genito-ventral and anal shields H. heselhausi (Oudemans) (p. 177) Dorsal setae long, seta J1 reaching beyond the bases of /2 by about one-half their length (Text-fig. 224) ; genito-ventral shield flask-shaped (Text-fig. 22B), without setae between the genito-ventral and anal shields : H. nidicorva sp. nov. (p. 179) Apotele with three subequal tines (Text-fig. 42C) . 22 Apotele with two subequal tines. 5 - . : . 23 Larger species, over 1,000 uw ; sternal shield ain iw =0: 8: genito-ventral shield as in Text-fig. 42B : ‘ : H. myrmecophila (Berlese) (p. 214) Smaller species, about 700 wu ; sternal shield almost twice as broad as long (L/W = 0-5) ; genito-ventral shield as in Text-fig. 43B . H. myrmophila (Michael) (p. 218) Dorsum with only one pair of setae in the FR series (Text-fig. 194) ; peritreme extend- ing to about the middle of coxa II (Text-fig. 19F) ; Small species about 500 yu inlength . : : H. praesternalis Willmann (p. 173) Dorsum with five or more noe of gore in the FR series ; Oe aes extending up to or beyondcoxal . . : 24 Majority of dorsal and opisthogastric eee very fone) sinuous , (Text: figs. re A all 1D)\.5 setae J3 extending beyond the bases of setae J4 by at least a quarter of their length. Myemecocnions : : : ; : : : 25 Dorsal and opisthogastric setae normal in length ; setae J3 never reach the bases of Legs roleavely short and stumpy : lee Iv (eochanter to tarsus) about one-half the length of the dorsal shield ; movable digit of the chelicera strongly bidentate (Text-fig. gop)... : : : H. humerata Berlese (p. 212) Legs relatively long and Senden ; leg IV (rochanter to tarsus) scarcely shorter than the dorsal shield ; movable digit of chelicera edentate (Text-fig. 408) H. equitans (Michael) (p. 210) THE BRITISH DERMANYSSIDAE (ACARI) 163 26 Femur II with antero-ventral seta stout, spine-like ; tarsus IV with long, stout, spine-like setae ; genital shield small : : H. aculeifer (Canestrini) (p. 166) — Femur II with antero-ventral seta simple, not conspicuously thickened ; tarsus IV without long spine-like setae. : : : 3 : : : : ; 27 27 Cheliceral digits normal, slender ; movable digit longer than the palpfemur and strongly Bideneate : : : : 2 : : : , : 28 — Cheliceral digits atypical, Same movable digit shorter than the palpfemur, weakly dentate or edentate . ; : : : : 29 28 Genito-ventral shield greatly expanded nestener to coxae ys metapodal shields flanking genito-ventral shield ; marginal setae of dorsal shield strong, serrated H. astronomica (Koch) (p. 208) — Genital shield relatively narrow posterior to coxae IV ; metapodal shields widely removed from its margins (Text-fig. 17E) ; dorsal setae simple H. giffordi sp. nov. (p. 170) 29 Dorsal setae of the J series markedly unequal in length ; setae J4 and /J5 pilose distally and considerably longer and stouter than simple setae /1—/3 (Text-fig. 384). Myrmecophilous . : : : H. oophila (Wasman) (p. 207) = Dorsal setae of the J series subequal j in length : ; 30 30 Dorsal seta Z5 pilose distally and about three times the length of simple seta VE 5 (Text-fig. 374) ; movable digit of the chelicera edentate (Text-fig. 378) H. montana (Berlese) (p. 206) --— Dorsal seta Z5 less than twice the length of seta /5 (Text-fig. 364) ; movable digit of the chelicera dentate (Text-fig. 36D) : : H., isotricha (Wolenati) (p. 203) Hypoaspis krameri (Canestrini) Gamasus kvamervi Canestrini G. & R., 1881, Atti Ist. Veneto (5) 7: 1083, figs. Female: Chelicera with segment I, 72 » ; II, 180 » ; movable digit 80 p, bidentate ; fixed digit with about ten teeth (Text-fig. 13D) ; pilus dentilis short, setiform ; dorsal _ seta about twice as long as the pilus. Four pairs of gnathosomal setae with c.s. about 72 wapart, hyp.2 about 60 w apart. Deutosternum with six transverse rows of denticles (6-7 per row) ; corniculi 66 » ; internal malae as in Text-fig. 3c. Tectum capituli _ with anterior margin denticulate (Text-fig. 13B). Salivary stylus extends nearly to tip of corniculus. Pedipalp (2—-5—6—14) with two-tined apotele. Dorsal shield (684 , x 396 ») with 37 pairs of setae distributed as in Text-fig. 134; seta s3, the humeral seta and seven pairs of marginal setae on the cuticle. _ Setae 72-3, 22-6, considerably longer than the other setae, seta 74 extremely long _ and sinuous ; no hypertrichy. Surface of shield granular. Tritosternum with base 45 p, laciniae 123 p. Sternal shield (117 p» X 150 p) with three pairs of setae and three pairs of pores, the third pair being on the posterior margin of the shield ; metasternal setae free. Between st1, 93 »; between sti and | sf3, 114 p. Genital shield (114 » X 105 p) with genital setae only. Anal shield (93 » X 72 ») with paranal setae 48 p, postanal seta 30 ». Chaetotaxy and sclero- tization of opisthogaster as in Text-fig. 13F. Peritreme extends beyond middle of | coxa I; peritrematal shields free anteriorly and posteriorly. Legs with normal chaetotaxy. Long, whip-like setae present on femur IT (fd,), femur IV (ad,) and tarsus IV (ad, and pd,-3), Text-fig. 144. Tarsus II with two 164 G. OWEN EVANS & W. M. TILL ( Fic. 13. Hypoaspis kvamevi (Canestrini). A. Dorsum of idiosoma of female; B. tectum capituli of female; c. venter of gnathosoma of female; Dp. chelicera of female; THE BRITISH DERMANYSSIDAE (ACARI) 165 stout, blunt setae (a/, and f/,) and several which are stout basally, tapering to fine points (av,-2, Pv,-2, mv) ; relatively thick setae on tibia II (av, and v,) and genu II (av). Length/width (in ») of leg segments : I II III IV genu 93/42 84/57 66/45 90/45 tibia 108/33 84/54 75/42 100/45 tarsus 140/33 120/42 144/36 190/36 Male; Chelicera with segment I, 72 » ; II, 168 » ; movable digit 70 p, unidentate ; | spermadactyl 130 w long, slender (Text-fig. 13E); fixed digit tridentate ; pilus | | | | | | | _ dentilis and dorsal seta asin female. Four pairs of gnathosomal setae with c.s. about 60 » apart, hyp.2 about 50 p apart. Details of gnathosomal structure not clearly visible. Dorsal shield (636 » x 348 pw) as in female. Tritosternum with base 45 1, laciniae 114 ». Holoventral shield (516 » 114 pw) with four pairs of setae in the sternal region, six pairs in the genito-ventral region ; moderately expanded behind _ coxae IV but not extending beyond lateral margins of the coxae. Between sit, 80 p ; _ between str and st3, 123 w. Peritreme extends beyond middle of coxa I. | Legs with normal chaetotaxy ; long whip-like setae on legs IT and IV as in female. Leg II (Text-fig. 148) bears several stout setae : av, and especially av, on the femur ; Fic. 14. Hypoaspis kvameri (Canestrini). a. Leg IV of female; B. leg II of male. 166 G. OWEN EVANS & W. M. TILL av on the genu; av and fv on the tibia; al,, ply, avy, pv2, mv on the tarsus ; setae av, pv, and md on the tarsus are fairly stout basally, tapering to fine points. Legs all bent and not suitable for measurement. Hasitat: Recorded from Lucanus sp. (Coleoptera) in Britain and from Oryctes nasicornis Linn. in the Netherlands. Hypoaspis aculeifer (Canestrini) Laelaps aculeifey Canestrini, 1884, Aiti Ist. Veneto (6) 2: 608. Hypoaspis aculeifer: Bregetova, 1956, Opred. Faune SSSR. 61: 77, figs. 102-106. Female : Chelicera with segment I, 75 » ; I,216 ; movable digit 98 p, bidentate ; fixed digit with about ro teeth (Text-fig. 158) ; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 66 » apart and hyp.2 about 50 p apart. Deutosternum with six transverse rows of denticles ; corniculi 60 p long ; internal malae fringed (Text-fig. 15D). Tectum capituli with denticulate anterior margin (Text-fig. 15F). Salivary styli conspicuous. Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (684 » x 360 ») with normal 39 pairs of simple setae distributed as in Text-fig. 154. Setae of podonotal region conspicuously longer than those on the opisthonotal region. Surface of shield reticulate ; no hypertrichy ; with seven pairs of R setae on the cuticle. Tritosternum with base 48 p, laciniae 105 pw. Sternal shield (190 ph xX 138 2) extending almost to posterior margin of coxa III, with three pairs of setae and two pairs of pores; metasternal setae free with associated pores. Between sfI, 75 “; between sfx and st3, 144 w. Genital shield flask-shaped (120 » & 96 qu) with one pair of setae. Anal shield (90 » x 75 pw), with paranal seta 27 » and postanal seta 36 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. I5¢. Peritreme extends to middle of coxa I ; peritrematal shields free posteriorly, anterior region fused with dorsal shield at level of 71. Legs with normal chaetotaxy. Tarsus II with setae av,—5, pvy-o, aly, ply, md, mz, ad,, stout and spine-like. Tibia II with av and fv stout ; genu II with av stout ; femur II with av, stout. Stout setae are found on genu III (av and fv), tibia III (av and fv), tarsus III (most setae, except al2, plz, ads, pd3), genu IV (av), tibia 1V (az, pv, pl,) and tarsus IV (most setae). Length/width (in ») of leg segments : I II dE IV genu 96/40 105/66 63/50 105/63 tibia 114/30 96/66 66/48 120/63 tarsus 180/28 135/48 156/40 240/48 Male : Chelicera with segment I, 54 »; I, 130 » ; movable digit 60 », unidentate, spermadactyl as in Text-fig. 15G; fixed digit with about 5 teeth; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 50 » apart, yp.2 THE BRITISH DERMANYSSIDAE (ACARI) 167 Fic. 15. Hypoaspis aculeifer (Canestrini). A. Dorsum of idiosoma of female ; B. chelicera of female; c. venter of idiosoma of female; pb. venter of gnathosoma of female; E. dorsum of idiosoma of male; F. tectum capituli of female; G. chelicera of male ; H. venter of idiosoma of male. 168 G. OWEN EVANS & W.M. TILL about 42 » apart. Deutosternum, internal malae, tectum capituli as in female ; corniculi 54 » long. Pedipalps as in female. Dorsal shield (540 » x 276 «) with 39 pairs of setae distributed as in Text-fig. 15E. Tritosternum with base 27 p, laciniae 78 ». Holoventral shield (456 » x 114 yp) with three pairs of sternal, one pair of metasternal setae and three pairs of pores in the sternal region ; between st1, 63 »; between sf1 and st3, 126 w. Genito-ventral region with six pairs of setae. Paranal setae about 24 p, postanal seta about 20 1 long. Chaetotaxy of opisthogaster as in Text-fig. 15H. Peritreme extends nearly to middle of coxa I; anterior part of peritrematal shield fused with dorsal shield ; posterior part free. Chaetotaxy of legs as in female. Length/width (in ») of leg segments : I I Ill IV genu 75/33 80/54 50/42 84/54 tibia 90/27 75/54 50/40 90/54 tarsus 150/24 110/40 130/36 186/45 Protonymph : Chelicera with segment I, 40 »; II, 132 »; movable digit 50 yu. Four pairs of gnathosomal setae with c.s. about 45 » apart, hyp.2 about 36 pw apart. Deutosternum, internal malae and tectum as in female ; corniculi 36 » long. Pedi- palp (1-4-5-12) with two-tined apotele. Podonotal shield (258 » x 190 ») with eleven pairs of setae ; pygidial shield (90 x 120 p) with eight pairs of setae; chaetotaxy and sclerotization of mesonotal region as in Text-fig. 164A. Tritosternum with base 30 p, Jaciniae 60 ». Sternal shield (170 » x 80 pu) with three pairs of setae and two pairs of pores; between sti, about 48 1; between sf1 and st3, about 120 pw. Anal shield (50 » x 42 ») with paranal setae 20 p and post- anal seta 18 » long. Chaetotaxy of opisthogaster as in Text-fig. 16 B. Peritreme extends to middle of coxa ITI. Legs with normal chaetotaxy. Length/width (in ,) of leg segments : I II Th IV genu 60/30 60/36 40/30 57/33 tibia 66/30 50/40 42/30 60/36 tarsus 114/27 78/33 87/27 130/33 Deutonymph : Chelicera with segment I, 40 »; I, 147 «; movable digit 57 u. Deutosternum, hypostomal processes, corniculi and tectum as in protonymph. Pedipalp (2-5—6—14) as in female. Dorsal shield (450 » X 216 ») with lateral incisions, bearing 39 pairs of setae arranged as in Text-fig. 16c ; no hypertrichy; with seven pairs of R setae on the cuticle. THE BRITISH DERMANYSSIDAE (ACARI) 169 a Fic. 16. Hypoaspis aculeifey (Canestrini). a. Dorsum of idiosoma of protonymph ; B. venter of idiosoma of protonymph ; c. dorsum of idiosoma of deutonymph ; bD. venter of idiosoma of deutonymph. 170 G.:OWEN EVANS @°W.-M. TELL Tritosternum with base 40 p, laciniae 65 pw. Sternal shield (243 » x 82 p) bears three pairs of sternal, one pair of metasternal setae and three pairs of pores ; between stI, 60 4; between stx1 and sf3, 114 w. Anal shield (approximately 57 p x 50 p) with par- and postanal setae about 18 » long. Chaetotaxy of opisthogaster as in Text-fig. 16D. Peritreme as in female. Chaetotaxy of legs normal. Tibia IV with av,, pv, and pl, stout. Length/width (in ») of leg segments : I II III IV genu 66/33 72/48 48/42 60/42 tibia 78/30 57/48 48/40 72/50 tarsus 123/24 96/30 102/30 135/36 Hasitat : Common in soil and litter. Recorded from the nest of Riparia riparia (Linn.) in Gloucestershire, Britain ; from the nest of Spalax ehrenbergi Nehring in Israel and the nests of a variety of rodents in the U.S.S.R. Hypoaspis giffordi sp. nov. Female: Chelicera with segment I, 45 »; II, 120 »; movable digit 45 p», biden- tate ; fixed digit with three small teeth, pilus dentilis short, setiform (Text-fig. 17D). Four pairs of gnathosomal setae with c.s. about 45 » apart, hyp.2 about 36 pu apart. Deutosternum with six transverse rows of denticles (3—7 per row) ; corniculi 36 u long ; internal malae reduced (Text-fig. 17B). Tectum capituli with anterior margin smooth (Text-fig. 17c). Pedipalp (2-5—6—14) with two-tined apotele. Dorsal shield (480 » x 276 ») with 39 pairs of setae and four accessory setae distri- buted as in Text-fig. 17A. Surface of shield reticulate. Tritosternum with base 24 p, laciniae 54 ». Sternal shield (100 np x 84 p) weakly reticulate anteriorly and laterally, with three pairs of setae and two pairs of pores. Metasternal setae free with associated pores. Between sfI, 60 »; between sfI and st3, 90 w. Genital shield (84 » x 93 ») with genital setae only. Anal shield (60 x x 72 ») with paranal and postanal setae about 20 » long. Chaetotaxy and sclero- tization of opisthogaster as in Text-fig. 17E. Peritreme extends to middle of coxa [; anterior tip of peritrematal shield fused with dorsal shield ; posterior part free. Chaetotaxy of legs normal except in one specimen in which genu IV has a postero- ventral seta on one side only. Tarsus II with ventral setae, a/,, 61, and md stouter than the other setae ; tarsus IV with seta p/, spine-like. Length/width (in pz) of leg segments : I I] Ill IV genu 63/30 54/36 40/28 60/28 tibia 70/27 48/33 36/28 60/28 tarsus 110/24 80/24 82/42 117/24 Fic. 17. Hypoaspis Oe , female; dorsum of idiosoma ve a utosternum and\ hypo stome (B); t ate ae a ia li i (c); chelicera (D); v She: of idio a (E). ZOOL. 14, 5. 172 G..OWEN EVANS & W.2M. TILE Hasitat: Found on birch timber rotted by Polyporus betulinus, Rannoch, Perthshire. Collected by D. R. Gifford, November, 1962. Holotype 9 (1965 : 12: 29 : 3), paratype 2 (1965 : 12: 29: 4). Hypoaspis lubrica Voigts & Oudemans Hypoaspis lubrica Voigts & Oudemans, 1904, Zool. Anz. 27 : 654. Samsinak, 1960, Acta Soc. ent. Cech. 57 : 278, fig. 2 (1-9). Hypoaspis (Haemolaelaps) inveysus Berlese, 1918, Redia 13 : 127. Hypoaspis compressus Hull, 1925, Ann. Mag. nat. Hist. (9) 15: 209 (syn. nov.) Hypoaspis smithii Hughes, 1948, Mites associated with stored food products : 131. Hypoaspis murinus Strandtmann & Menzies, 1948 (1949), Ann. ent. Soc. Amer. 41: 479, figs. Female : Chelicera with segment I, 66 » ; II, 156 »; movable digit 56 pu, bidentate ; fixed digit tridentate ; pilus dentilis short, setiform (Text-fig. 188). Four pairs of gnathosomal setae with c.s. about 66 apart ; Ayp.2 about 42 papart. Deutosternum with six transverse rows of denticles (7-10 per row) ; corniculi 45 » long ; internal malae fringed. Tectum capituli with fine striations and a smooth anterior margin (Text-fig. 18c). Salivary stylus reaches tip of corniculus. Pedipalp (2-5-6-14) with two-tined apotele. Fic. 18. Hypoaspis lubrica Voigts and Oudemans, female; dorsum of idiosoma (a); chelicera (B) ; tectum capituli (c) ; venter of idiosoma (D). THE BRITISH DERMANYSSIDAE. (ACARI) 173 Dorsal shield (708 » x 420 ») with 39 pairs of setae and 4-9 accessory setae distributed as in Text-fig. r8A. Surface of shield reticulated. Tritosternum with base 30 p, laciniae 90 p. Sternal shield (132 pm X 120 ph) extending to middle of coxa III, with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between stf1, 70 «; between sf1 and st3, 112 p. Genital shield (150 » x 117 ») flask-shaped with one pair of setae. Anal shield (110 » X 105 p) with par- and postanal setae 36 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 18p. Peritreme extends to beyond middle of coxa I; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs differs from typical pattern in having a postero-ventral seta on genu lV. Setae av, pv, al,, pl, on tarsus II are stout and spine-like ; setae avy, pu, and mv are stout and pointed. Tibia II has stout ventral setae (av, and pv,). Legs III and IV bear many stout ventral setae on the genu, tibia and tarsus. Length/ width (in ») of leg segments : I II III IV genu 87/38 84/50 54/40 90/42 tibia 96/36 75/48 57/40 go/40 tarsus 147/33 105/36 = 130/33 174/33 HapitaT: In decaying oats and hay ; in the nest of Riparia riparia (Linn.) ; on a | variety of rodents and in their nests. Recorded from Britain, Europe, the U.S.S.R. and the U.S.A. Hypoaspis praesternalis Willmann | Hypoaspis praesternalis Willmann, 1949, Veroff. Mus. Nat. Bremen no, 1A: 115, figs. Hypoaspis nolli Karg, 1962, Mitt. zocl. Mus. Berlin 38 : 62, figs. (syn. nov.). Female : Chelicera with segment I, 50 »; II, 144 »; movable digit 60 pw, biden- tate ; fixed digit with bifid tip and four teeth, pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 45 p apart, hyp.2 about 42 pw apart. _ Deutosternum with six transverse rows of fine denticles; corniculi 52 p long ; _ internal malae slender, triangular, fringed. Tectum capituli with anterior margin _ denticulate (Text-fig. 19c). Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (492 » 264 ») with 39 pairs of setae distributed as in Text-fig. 19. No hypertrichy. Surface of shield reticulate. Two pairs of R setae on the cuticle. Tritosternum with base 36 p, laciniae 84 » ; presternal area reticulate and granular. | Sternal shield (138 » X 105 ») with reticulate pattern most pronounced laterally ; with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between sf1, 75 »; between sti and s¢3, 100 p. Genital shield (90 » x 72 p) _ with genital setae only. Anal shield (72 « x 70 ») with paranal and postanal setae _ about 33 w long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. IgE. 174 G. OWEN EVANS) @0W: Mo TILE Fic. 19. Hypoaspis praesternalis Willmann. chelicera of male; c. tectum capituli of female ; female ; ©. venter of idiosoma of female. Ae Dorsum of idiosoma of female; B. D. peritreme and peritrematal shield of THE BRITISH DERMANYSSIDAE (ACARI) 175 _ Peritreme extends to about middle of coxa II ; anterior part of peritrematal shield fused with dorsal shield, posterior part free (Text-fig. 19D). Chaetotaxy oflegsnormal. Tarsi II-IV with ventral setae relatively stout basally, - tapering to long, fine points. Tarsus IV with setae f/, and #/; relatively stout and blade-like. Length/width (in ») of leg segments : if i Ill IV genu 68/30 60/36 40/27 63/30 tibia 75/24 54/36 40/27 66/32 tarsus 130/20 80/30 72/24 E17 /27 Male: Chelicera with segment I, 40 » ; II, 105 » ; movable digit 42 pu, unidentate ; _ fixed digit with bifid tip and four teeth, pilus dentilis simple ; spermadactyl large as in Text-fig. 198. Four pairs of gnathosomal setae with c.s. about 42 p apart, hyp.2 . about 36 » apart. Corniculi 36 pw long and set wider apart than in the female. Tectum and pedipalps as in female. Dorsal shield (420 » x 228 ») with chaetotaxy similar to that in the female except for the absence of setae px2 in the opisthonotal region. Surface of shield reticulated. Tritosternum with base 15 p, laciniae 704. Holoventral shield (348 p x 93 1) _ with four pairs of setae and three pairs of pores in the sternal region, five pairs in the genito-ventral region ; anal region with normal three setae. Between s?1, 66 yp; between s¢1 and s¢3, go w. Par- and post-anal setae subequal in length. Peritreme extends to about the middle of coxa II; peritrematal shield free posteriorly. Chaetotaxy of legs as in female. Setae of femur II simple. Length/width (in p) _ of leg segments : I II iy IV genu 66/27 54/36 36/30 57/33 tibia 68 /24 48/33 36/27 60/30 tarsus 117/20 T2127 70/24 105/26 HapitatT: Recorded from soil, grassland and marshes in Britain and Europe. Hypoaspis sardoa (Berlese) Laelaps (Androlaelaps) sardous Berlese, 1911, Redia 7 : 433. Bregetova, 1956, Opred. Faune SSSR. 61 : 83, figs. 120-122. Hypoaspis laevis var. pilifer Oudemans, 1912, Ent. Ber. Amst. 3: 231. Female: Chelicera with segment I, 102 »; II, 228 »; movable digit go p, biden- | tate; fixed digit with three teeth and a row of serrations; pilus dentilis short, _ setiform (Text-fig. 20c). Four pairs of gnathosomal setae with c.s. about 68 apart, hyp.2 about 57 » apart. Deutosternum with six transverse rows of denticles (4-8 | per row); corniculi go p» long; internal malae fringed (Text-fig. 20£). Tectum 176 G. OWEN EVANS @2W. M. TILE Fic. 20. Hypoaspis sardoa (Berlese), female ; dorsum of idiosoma (A) ; tectum (B) ; cheli- cera (c); femur, genu and tibia of leg II (p) ; deutosternum and hypostome (E) ; venter of idiosoma (F). THE BRITISH DERMANYSSIDAE (ACARI) e767, _capituli with denticulate anterior margin (Text-fig. 20B) ; salivary styli reach nearly to tips of corniculi. Pedipalp (2-5—6-14) with two-tined apotele. Dorsal shield (732 » 456 wu) with 39 pairs of setae (Text-fig. 204A). Surface of shield reticulated ; no hypertrichy ; one pair of 7 and six pairs of R setae on the cuticle. Tritosternum with base 48 p, laciniae 120 wp. Sternal shield (170 pw x 156 p) with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between st1, 84 »; between sfz and s#3, 160 ». Genital shield (216 p x 156 ) flask-shaped, with two pairs of setae. Anal shield (80 » x 114 ») with maxi- mum width 140 » ; paranal setae about 66 p, postanal seta about 33 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 20F. Peritreme extends to anterior margin of coxa II; peritrematal shield has anterior part fused with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern in that tibia III and genu IV each have a second postero-lateral seta. Femur II has a stout, blunt seta (av,) ; genu II has one (av,) and tibia II has two (av, and pv,) stout, pointed setae (Text-fig. 20D). On tarsus II setae av,—2, pv,;-, and mv are stout and pointed. Length/width (in p) of leg segments : | I II III IV | genu 108/48 114/66 84/48 120/48 tibia 126/45 105/60 90/48 132/48 tarsus 198/42 150/45 170/42 230/40 | _ Hapirat: In litter and in the nests, more rarely on the bodies, of moles and field mice (Apodemus flavicollis (Melchior) and A. sylvaticus (Linn.)) in Britain, Europe and the U.S.S.R. Recorded by Bregetova from the nest of Phoenicurus phoenicurus » (Linn.). | Hypoaspis heselhausi Oudemans Hypoaspis heselhaust Oudemans, 1912, Ent. Bey. Amst. 3: 216. Bregetova, 1956, Opred. Faune SSSR. 61: 78, figs. Female : Chelicera with segment I, 45 » ; II, 126»; movable digit 48 p, bidentate ; | fixed digit with five teeth, pilus dentilis short, setiform, dorsal seta a little longer and _ stouter than the pilus (Text-fig. 21c). Four pairs of gnathosomal setae with c.s. about 45 pw apart, hyp.2 about 30 » apart. Deutosternum with six transverse rows of | denticles (4-6 per row) ; corniculi 36 » long; internal malae as in Text-fig. 21B. _ Tectum capituli with anterior margin smooth. Pedipalp (2-5—6-14) with two-tined _ apotele. Dorsal shield (480 » x 280 ») with 39 pairs of short, simple setae distributed as in | Text-fig. 21a. Surface of shield with scale-like markings. | Tritosternum with base 36 p, laciniae 72 ys. Sternal shield (108 p x 102 yp) | with three pairs of setae and two pairs of pores ; metasternal setae free with associa- —= 178 G. OWEN EVANS & Wi, Mo rtil Pic Fic. 21. Hypoaspis heselhausi Oudemans, female ; dorsum of idiosoma (A) ; deutosternum _ and hypostome (B) ; chelicera (c) ; venter of idiosoma (Dp). THE BRITISH DERMANYSSIDAE (ACARI) 179 _ ted pores. Between sti, 56 «; between sf1 and st3, 96 uw. Genito-ventral shield (108 » X 84 ») with maximum width (135 ») near posterior margin, bearing a pair of genital and a pair of opisthogastric setae. Anal shield (75 » x 105 ») with maximum width (117 ») near anterior margin ; paranal setae 20 p, postanal seta 18 p. Chaeto- taxy and sclerotization of opisthogaster as in Text-fig. 21D. Peritreme extends to middle of coxa I ; anterior tip of peritrematal shield fused with dorsal shield, posterior part completely or incompletely fused with podal shield of coxa IV. Legs with normal chaetotaxy. Cuticle of legs with warty appearance. Length/ width (in ») of leg segments : I II ett IV genu 60/27 57/38 36/30 57/33 tibia 63/27 48/36 36/30 57/30 tarsus 108/25 W227, 80/24 90/27 | HasitaT: In the nests of Talpa europaea Linn. and Microtus arvalis (Pallas) in ‘Britain, the Netherlands and the U.S.S.R. We have also examined specimens from the nest of Bombus lapidarius (Linn.) at Langley Park, Buckinghamshire (coll. _K. F. Colombo). | Hypoaspis nidicorva sp. nov. Female: Chelicera with segment I, 70 » ; II, 180 7; movable digit 75 ju, biden- tate; fixed digit tridentate ; pilus dentilis short, setiform (Text-fig. 23E) Four pairs of gnathosomal setae with c.s. about 54 » apart, hyp.2 about 42 pw apart. -Deutosternum with six transverse rows of denticles (6—7 per row) ; corniculi 66 long; internal malae with a deep fringe as in Text-fig. 23F. Tectum capituli with anterior margin smooth (Text-fig. 23B). Salivary styli conspicuous, extending a little beyond tips of corniculi. Pedipalp (2-5-6—-14) with two-tined apotele. | Dorsal shield (624 » x 384 ») with 39 pairs of setae and three accessory setae distributed as in Text-fig. 224. Surface of shield reticulate and granular. Tritosternum with base 48 p, laciniae 102 pp. Sternal shield (110 p x 130 p) granular, anterior two-thirds reticulate, with three pairs of setae and two pairs of pores. Metasternal setae free with associated pores. Between st1, 70 »; between st1 and st3, 105 p. Genito-ventral shield (216 » x 120 p) flask-shaped, posterior ‘margin truncate, very close to anal shield ; bearing one pair of genital and three pairs of opisthogastric setae. Between Jv2, 162 ». Anal shield (78 p x 105 p) ‘with maximum width at anterior margin (132 ») ; paranal setae 30 »; postanal seta 45 w. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 22B. Peri- treme extends beyond posterior margin of coxa I; anterior part of peritrematal shield fused with dorsal shield ; posterior part fused with podal shield of coxa IV. 180 G. OWEN EVANS & W. M. TILL Hyvpoaspis nidicorva sp. nov., female; dorsum (A) and venter (B) of idiosoma. FIGY 22: Chaetotaxy oflegsnormal. Dorsalsetaon trochanter lV relatively stout. Length/ width (in ») of leg segments : I II Ill IV genu 84/36 75/42 54/30 78/30 tibia 90/30 70/42 54/30 80/27 tarsus 132/30 102/30 110/24 162/24 Male: Chelicera with segment I, 40 »; II, 117 »; movable digit 45 », as long as | spermadactyl, unidentate ; fixed digit unidentate, pilus dentilis short, setiform (Text-fig. 23c). Four pairs of gnathosomal setae with c.s. about 40 u apart, iyp.2 | about 36 » apart. Corniculi 48 » long. Deutosternum, internal malae, tectum and pedipalps as in female. Dorsal shield (396 » x 267 ) as in female, with 39 pairs of setae and one or two accessory setae. Tritosternum with base 36 jy, laciniae 105 ,. Holoventral shield (360 « X 96 ») with four pairs of setae and three pairs of pores in the sternal THE BRITISH DERMANYSSIDAE (ACART) 181 region, one pair of genital and five pairs of opisthogastric setae in the genito-ventral region (Text-fig. 234). Between sf1, 48 »; between sf1 and st3, 90 w. Paranal setae 24 ; postanal seta 36 w. Peritreme extends beyond posterior margin of coxa T; anterior part of peritrematal shield fused with dorsal shield, posterior part fused with holoventral shield and with podal shield of coxa IV. Fic. 23. Hypoaspis nidicorva sp.nov. A. Venter of idiosoma of male; B. tectum capituli of female; c. chelicera of male; pb. femur II of male ;-£. chelicera of female; F. venter of gnathosoma of female. 182 G. OWEN EVANS & W. M. DILL Chaetotaxy of legs as in female. Femur II with seta av, very stout and spine-like (Text-fig. 23D). Length/width (in ») of leg segments : I II Ill IV genu 63/27 60/36 40/22 60/20 tibia 70/24 50/33 40/24 63/22 tarsus 105/24 75/26 84/21 126/21 Deutonymph : Chelicera with segment I, 50 »; II, 156 »; movable digit 60 uz. Gnathosoma generally as in female ; setae c.s. about 48 p apart, hyp.2 about 36 p apart ; corniculi 45 p long. Dorsal shield (444 » 246 ») with lateral incisions ; with 14 pairs of setae in the podonotal region, the s series and the humeral seta being on the striated cuticle ; opisthonotal region with the usual 17 pairs of setae and two accessory setae. Trito- sternum with base 45 p, laciniae 84 pw. Sternito-genital shield (246 pw X I14 ph) with four pairs of setae and three pairs of pores; between sfI, 70 »; between sf and st3, 108 ». Anal shield (66 » x 75 pw) triangular with maximum width (100 p) at anterior margin; paranal setae 24 p, postanal seta 4o p. Peritreme extends beyond posterior margin of coxa I; peritrematal shield free anteriorly, posterior part not developed. Chaetotaxy of legs as in female. Length/width (in ) of leg segments : I II III IV genu 70/30 63/42 45/27 63/27 tibia 72/30 54/40 45/30 63/27 tarsus 108/27 84/27 90/24 132/24 Hasitat: Nest of a Jackdaw, Wookey Hole, Somerset, Britain, collected by O. C. Lloyd, 11th September, 1957. Holotype female (1965 : 12 : 29 : 27), allotype male (1065 7127-2091: 28). Hypoaspis claviger (Berlese) Laelaps claviger Berlese, 1883, Acari, Myr. Scorp. Ital. fasc. 4, no. 2. Female; Chelicera with segment I, 60 »; II, 156 »; movable digit 60 p, biden- tate; fixed digit with four teeth (Text-fig. 24G); pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 57 apart, hyp.2 about 42 mu apart. Deutosternum with six transverse rows of denticles (2-6 per row) ; corniculi 50 # long; internal malae branched (Evans & Till, 1965; Text-fig. 4A). Tectum capituli with denticulate anterior margin. Salivary stylus reaches tip of corniculus. Pedipalp (2-5-6—14) with two-tined apotele. Dorsal shield (654 « X 366 «) with the normal 39 pairs of setae and four unpaired accessory setae, all spatulate in shape (Text-fig. 24c) and arranged as in Text-fig. 244, Surface of shield reticulate ; seven pairs of & setae on the cuticle. THE BRITISH DERMANYSSIDAE (ACARI) 183 | Fic. 24. Hypoaspis claviger (Berlese). A. Dorsum of idiosoma of female; B. dorsal seta J4 of female; c. dorsal seta Z5 of female; p. venter of idiosoma of female; E. venter of idiosoma of male; F. chelicera of male; G. chelicera of female. 184 G. OWENSEVANS? & WW. M. TILE Tritosternum with base 45 p, laciniae 80 ». Sternal shield (110 » X IIO yw) with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sti, about 70 » ; between sti and sf3, 102 w. Genital shield flask-shaped (105 # X 102 pw), with one pair of setae. Anal shield (102 » X 114 pv) with paranal and postanal setae about 20 w long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 24D. Peritremes reach midpoint at anterior end of idiosoma ; _peri- trematal shields free anteriorly, posterior part fused with podal elements of coxa IV. Legs with normal chaetotaxy. Length/width (in ,) of leg segments : I af eh IV genu 100/40 80/50 63/45 87/45 tibia 100/33 70/48 60/45 90/48 tarsus 153/27 138/40 140/36 216/36 Male: Chelicera with segment I, 42 »; II, 108 »; movable digit 40 p, uni- dentate ; spermadactyl as in Text-fig. 24F; fixed digit bidentate. Four pairs oi gnathosomal setae with c.s. about 48 p apart, hyp. 2 about 36 wapart. Deutosternum with six transverse rows of denticles (2-5 per row) ; corniculi 36 » long. Other gnathosomal features as in female. Dorsal shield (486 » x 282 ) with 39 pairs of setae and five unpaired accessory setae ; seven pairs of & setae on the cuticle. Tritosternum with base 30 yz, laciniae 60 ». Holoventral shield (360 » x 96 pw) with three pairs of sternal, one pair of metasternal setae and three pairs of pores in the sternal region ; between sfI, 54 u; between sti and s¢3, 93 pw. Genito-ventral region with six pairs of setae ; paranal setae 18 p, postanal seta I5 wp. Chaetotaxy of opisthogaster, including R series, as in Text-fig. 24£. Peritremes reach mid-anterior point of idiosoma ; peritrematal shields fused anteriorly with dorsal shield, posteriorly with holoventral shield. Legs with normal chaetotaxy. Length/width (in ») of leg segments : I Il Iil IV genu 80/36 63/42 50/36 78/40 tibia 80/30 56/40 48/36 80/36 tarsus 135/24 17/3 120/30 170/33 Hasitat: Probably predacious, living in soil, litter and rotting wood. Recorded from Britain and Europe. Hypoaspis cuneifer (Michael) Laelaps cuneifer Michael, 1891, Proc. zool. Soc. Lond. : 647. Laelaps (Cosmolaelaps) cuneifer: Berlese, 1904, Redia 1: 416, pl. 15, figs. 123-124, pl. 16, fig. 125. Cosmolaelaps cuneifer: Bregetova, 1956, Opred. Faune SSSR. 61: 75. Female: Chelicera with segment I, 78 »; II, 234 »; movable digit 84 ,, biden- tate ; fixed digit with about seven teeth, pilus dentilis short, setiform (Text-fig. 25F). THE BRITISH DERMANYSSIDAE (ACART) 185 Four pairs of gnathosomal setae with c.s. about 90 uw and hyp.2 about 72 mu apart. Deutosternum with six transverse rows of denticles ; corniculi 50 u long; internal malae with long slender filaments as in Text-fig. 25H. Tectum capituli with denti- culate anterior margin (Text-fig. 25D). Pedipalp (2-5-6—14) with two-tined apotele. Dorsal shield (804 » xX 684 ») with the normal 39 pairs of setae plus 14 unpaired accessory setae; all except setae v1 and S2 are cuneiform (Text-fig. 254 and Cc). Surface of shield with reticulations consisting of rows of tiny spine-like markings. Tritosternum with base 54 pu, laciniae 123 w. Sternal shield (168 » Xx 162 yp) with two pairs of setae and one pair of pores ; first pair of setae and pores situated on granular presternal area. Metasternals free with associated pores. Between s/t, 72 w; between sfI and s/3, 153 w. Genital shield (168 » x 204 ») expanded behind genitalsetae. Analshield (114 X 126 u) with paranal and postanal setae subequal, 24-30 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 25B. Peritreme extends to middle of coxa 1; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern in having two postero-lateral setae on genu IV (2—?, 2—2). Most leg setae simple, some dorsal setae (on femora and trochanters) spatulate. Length/width (in ,) of leg segments : i Il III IV genu 132/54 108/66 90/54 138/57 tibia 150/45 96/66 90/54 = 138/54 tarsus 234/42 180/50 186/45 270/42 Male: Chelicera with segment I, 70 »; II, 192 »; movable digit 66 uv (84 u with spermadactyl), unidentate ; fixed digit with four distinct teeth and a row of fine serrations. Four pairs of gnathosomal setae with c.s. about 75 » and hyp.2 about 66 uw apart. Corniculi 45 pw long; deutosternum, internal malae, tectum and pedipalps as in female. Dorsal shield (636 » x 546 w) with the normal 39 pairs of setae as well as unpaired accessory setae. Surface of shield patterned as in female. Tritosternum with laciniae approximately three times as long as the base. Holo- ventral shield (492 » X 147 ») with four pairs of setae and three pairs of pores in the sternal region, five pairs of setae in the genito-ventral region (Text-fig. 25£). Bet- ween sfI, 84 »; between sf1 and s/3, 123 uw. Par- and postanal setae subequal in length, about 30 uw. Peritreme extends to middle of coxa I; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs asin female. In addition, seta av, on femur IT is large and spine- like (Text-fig. 25G) ; setae av on genu and tibia II are considerably stouter than setae fv. Length/width (in pz) of leg segments : I Il Ill IV genu 120/48 96/57 78/48 120/54 tibia 120/42 78/54 72/45 120/48 tarsus 198/36 120/42 132/36 198/42 186 G. OWEN EVANS & W. M. TILL Fic. 25. Hypoaspis cuneifey (Michael). a. Dorsum of idiosoma of female; 3B. venter of idiosoma of female; c. dorsal seta 26 of female; D. tectum capituli of female ; E. venter of idiosoma of male; F. chelicera of female ; G. femur II of male; H. venter of gnathosoma of female. THE BRITISH DERMANYSSIDAE (ACARI) 187 Hasitat: Myrmecophilous, associated with many ant hosts. Recorded from Europe, the U.S.S.R. and (?) Britain. We have included this species on the basis of the records given by Hull (1918) and by Turk (1953). It is possible that this species does not occur in Britain, but that its identity has been confused with that of H. neocunerfer. Hypoaspis neocuneifer sp. nov. Female: Chelicera with segment I, 75 »; II, 186 »; movable digit 66 », biden- tate ; fixed digit with from five to nine teeth ; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 75 p apart, Ayvp.2 about 57 wm apart. Deutosternum with six transverse rows of small, fine denticles ; corniculi 45 1; internal malae with long, slender filaments as in H. cuneifer. Tectum capituli with denticulate anterior margin. Pedipalp (2—5—6-14) with two-tined apotele. Dorsal shield (740 » X 528 ») with the normal 39 pairs of setae and about nine unpaired accessory setae distributed as in Text-fig. 264. Setae cuneiform in shape (Text-fig. 26D). Surface of shield with reticulations consisting of rows of tiny spine- like markings. Tritosternum with base 45 p, laciniae 110 w. Sternal shield (117 pw x 144 pL) with two pairs of setae and one pair of pores, the first pair of setae and pores being situated on the granular pre-sternal area. Metasternal setae free with associated pores. Between sti, 80 »; between sti and st#3, 126 w. Genital shield (123 p Xx 153 p) flask-shaped, reticulated. Anal shield (110 » x 108 ,) with paranal setae simple, 27 » long; postanal seta spatulate, 24 » long. Chaetotaxy and sclerotiza- tion of opisthogaster as in Text-fig. 26B. Peritreme extends to about anterior third of coxa I ; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Legs with normal chaetotaxy. Most leg setae simple, but rather short and spine- like, except at tip of tarsus I; some dorsal setae (on trochanters I, III, IV, and femora II-IV) spatulate. Length/width (in ») of leg segments : I II Ill IV genu 120/48 102/60 84/50 120/54 tibia 132/42 84/54 78/48 120/48 tarsus 186/36 144/45 156/42 225/42 Male : Chelicera with segment I, 50 » ; II, 165 » ; movable digit 58 », unidentate ; spermadactyl 84 » long from base of movable digit ; fixed digit with about nine small teeth, pilus dentilis short, setiform (Text-fig. 26£). Four pairs of gnathosomal setae with c.s. about 63 » apart, hyp.2 about 50 pw apart. Corniculi 36 » long; deuto- sternum, internal malae, tectum and pedipalps as in female. Dorsal shield (624 » x 408 p) with 39 pairs of setae and about nine unpaired accessory setae. Surface of shield as in female. Tritosternum with base 33 p, laciniae 90 p. Venter bears a sternito-genito- ventral shield (336 » x 126 w) and aseparate anal shield (110 » * 96 p») ; the former ZOOL. 14, 5. 13 188 G.LOWEN EVANS <&. W. Meet LEE Fic. 26. Hvypoaspis neocuneifer sp. nov. A. Dorsum of idiosoma of female; 3B. venter of idiosoma of female; c. venter of idiosoma of male; Db. setae z4 and 26 of female; E. chelicera of male. THE BRITISH DERMANYSSIDAE (ACARI) 189 with four pairs of sternal setae and three pairs of pores, a pair of genital setae and three to five opisthogastric setae (Text-fig. 26c). Between sf1, 70 » ; between sfi and sf3, 102 p. Paranal setae 24 pw, postanal seta 18 p long. Chaetotaxy of legs and form of setae as in female ; in addition, seta av, on femur II stout and spine-like. Length/width (in ,) of leg segments : I II ri LV genu 99/42 84/54 66/45 102/48 tibia 105/38 78/50 66/42 102/45 tarsus 150/33 114/42 126/40 186/38 HasitaT: Nests of Formica rufa Linn., White Hill Wood, Canterbury, 1st Novem- ber, 1942 and 25th April, 1942. Holotype 9 (1965 : 12 : 29 : 23), allotype (1965 : 12: 29: 24), four female paratypes. Ants’ nest, White Hill, Canterbury, 18th October, 1941, one male, four female paratypes. Chip nest, same locality, 29th March, 1943, one female paratype. Allspecimens collected by Dr. E. Warren and deposited in the collection of the British Museum (Natural History). It is possible that this species has been confused with Hypoaspis cunetfer (Michael). The specimens of H. cunezfer in the Donisthorpe collection are referable to the new species. We have not seen material of the true H. cunezfer in Britain. Fic. 27. Hypoaspis vacua (Michael), female ; dorsum (a) and venter (B) of idiosoma. 190 G..OWEN- EVANS & W.M:. TILL Hypoaspis vacua (Michael) Laelaps vacua Michael, 1891, Proc. zool. Soc. Lond.: 651, figs. Laelaps (Cosmolaelaps) vacuus; Berlese, 1904, Redia 1: 4109, figs. Laelaps (Cosmolaelaps) vacuus var. ensigey Berlese, 1904, Redia 1: 420, figs. Female ; Chelicera with segment I, 33 » ; II, 96 »; movable digit 33 p, bidentate ; fixed digit with five teeth, pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 50 u apart, hyp.2 about 36 w apart. Deutosternum with 6 or 7 transverse rows of denticles ; corniculi 27 » long ; internal malae difficult to discern, but appear to consist of at least two pairs of delicate filaments. Tectum capituli with denticulate anterior margin. Salivary styli slender, reaching to tips of cor- niculi. Pedipalp (2—5—6-14) with two-tined apotele. Dorsal shield (420 » 264 ») with the normal 39 pairs of setae and two unpaired accessory setae distributed as in Text-fig. 27A. All the setae except 21 are scimitar- like. Surface of shield reticulate. | | / / hh | i borage Nae VX oe) Sz! We A B Fic. 28. Hypoaspis vacua (Michael), female. and /3 and of the genital shield in (a), the type specimen from Igls (Austria); in (B), a specimen from Eire, and in (c), a specimen from Sussex (England). Variation in the form of dorsal setae /2 THE BRITISH DERMANYSSIDAE (ACARI) 191 Tritosternum with base 30 p, laciniae 72 ». Sternal shield (93 » x 90 ) with anterior margin extremely indistinct, bearing three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sfI, 54 «; between sti and st3, 70 w. Genital shield (90 » x 78 pw) with lateral margins almost parallel ; bearing genital setae only. Anal shield (66 » x 63 ) with paranal setae 18 yu long, postanal seta 21 long. Chaetotaxy and sclerotization of opisthogaster as in Text- fig.27B. Peritremes extend to middle of coxa I; anterior part of peritrematal shield fused with dorsal shield, posterior part free. In the limited material of this species which we have examined there appears to be considerable intraspecific variation in the length of the dorsal setae and in the form of the genital shield (Text-figs. 284—c). Legs with normal chaetotaxy. Length/width (in ,) of leg segments : I I I IV genu 60/30 51/33 36/27 60/33 tibia 66/24 45/33 33/27 63/30 tarsus 117/24 84/27 72,24 Eii/27 Hapitat: Moss on tree stumps, Beckley, Sussex, 19th May, 1951; on Isle of Rhum, Scotland, July, 1964; ants’ nest, Leigh-on-Sea, Essex, 14th May 1951 ; Mulranny, Eire, September, 1913; ants’ nests in Austria and Italy. Hypoaspis bombicolens (Canestrini) Iphis bombicolens Canestrini, 1884, Prosp. Acarof. Ital. (6) 2 : 1634. Female: Chelicera with segment I, 42 »; II, 108 »; movable digit 38 », biden- tate ; fixed digit with one large and one small tooth, pilus dentilis short, setiform ; dorsal seta short, slender (Text-fig. 29F). Four pairs of gnathosomal setae with c.s. about 54 » apart, hyp.2 about 38 » apart. Deutosternum with six transverse rows of denticles (2-4 per row) ; corniculi 27 » ; internal malae as in Text-fig. 29c. Tectum capituli with smooth anterior margin (Text-fig. 29E). Salivary styli slender, nearly reaching tips of corniculi. Pedipalp (2-5—6-14) with two-tined apotele. Dorsal shield (672 » « 384 ») with at least 40 pairs of setae, pxr being present on the opisthonotal part (in the specimen figured seta $x3 is missing on one side). In addition, a variable number of marginal setae are included on both the podonotal and opisthonotal regions of the shield (Text-fig. 29A). Several anterior setae of the j, 2, 8 and yr series are considerably longer than the remaining setae. Surface of shield granular and reticulate ; no hypertrichy. Tritosternum with base 24 p, laciniae 81 p. Sternal shield (84 » x 138 mw) with three pairs of setae and two pairs of pores, metasternal setae free with associated pores. Between st1, 87 » ; between sti and st3, 99 ». Genital shield (120 » * 105 p) with genital setae only. Anal shield (96 » x 70 ») with paranal setae 18 p, postanal seta 21 wu. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 29B. 192 G. OWEN EVANS & W. M. TILL WLI, Ae, \\ U/L ul \y THE BRITISH DERMANYSSIDAE (ACARI) 193 Peritreme extends to anterior third of coxa I; peritrematal shield fused anteriorly with dorsal shield (Text-fig. 29D) ; posterior part free. Chaetotaxy of legs differs from normal pattern in that genu IV has two ventral setae (2—7, ?—1). Length/width (in y) of leg segments : I II III IV genu 75/45 75/57 50/48 72/45 tibia 50/40 66/50 54/42 75/40 tarsus 132/36 102/40 120/36 144/33 HasiTaT: Recorded from Bombus terrestyvis (Linn.) in Essex, Britain, and from Bombus sp. in Italy. Hypoaspis breviseta sp. nov. Female: Chelicera with segment I about 50 »; II, 120 pw. Structural details are not visible as the chelicerae are retracted. Four pairs of gnathosomal setae with c.s. about 60 pw apart, hyp.2 about 45 » apart. Deutosternum with six transverse rows of denticles (I-6 per row) ; corniculi 30 »; internal malae similar to those of H. bombicolens, but with the addition of a pair of slender, inner processes. Tectum capituli with smooth anterior margin. Salivary styli nearly reaching tips of cor- niculi. Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (744 x 456 ») with 42-44 pairs of setae distributed as in Text-fig. 30A. Setae pxI-—3 are present in the opisthonotal region ; additional marginal setae are present both in the podonotal and the opisthonotal region, but there are no accessories between the J series. The setae are short and simple, 71 being relatively stout and Z5 having the form indicated in Text-fig. 30B. Surface of shield granular. Tritosternum with base 27 p, laciniae 84 ». Sternal shield (84 » x 150 ») granular, reticulations very faint except in the antero-lateral corners ; with three pairs of setae and two pairs of pores; metasternals free with associated pores. Between s/I, 75 »; between sfi and st3, 102 w. Genital shield (144 » X 150 pw) granular, with genital setae only. Anal shield (123 » x 108 m) granular; paranal setae 30 p long, postanal seta broken. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 30c. Peritreme extends to middle of coxa I; peritrematal shield fused anteriorly with dorsal shield ; posterior part free. Chaetotaxy of legs differs from free-living pattern in that genu IV bears two ventral setae (pv present). Length/width (in p) of leg segments : il II III IV genu 78/48 75,66 66/54 90/54 tibia 87/42 72/57 66/48 87/48 tarsus 138/40 120/42 130/40 150/40 Fic. 29. Hypoaspis bombicolens (Canestrini), female; dorsum of idiosoma (A); venter of idiosoma (B); venter of gnathosoma (c); peritreme and peritrematal shield (Dp) ; tectum capituli (E) ; chelicera (F). 194 G. OWEN EVANS & W. Mo LLvE Fic. 30. Hypoaspis breviseta sp. nov., female; dorsum of idiosoma (A); seta Z5 (B); venter of idiosoma (c). Hasitat: On Bombus muscorum (Linn.), Scarborough, Yorkshire, October, 1964. Holotype 2 (1965 : 12:29:26) in the collection of the British Museum (Natural History). Hypoaspis colomboi sp. nov. Female: Chelicera with segment I, 54 »; II, 126 »; movable digit 45 p», biden- tate ; fixed digit unidentate, pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 60 pw apart, Ayp.2 about 40 » apart. Deutosternum with six transverse rows of denticles; corniculi 40 » long; internal malae with fringed, elongate, inner lobe and fimbriate outer lobe. Tectum capituli with anterior margin smooth, slightly scalloped. Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (600 » 336 ») with 38 pairs of setae, x2 missing (Text-fig. 314). Surface of shield granular, no hypertrichy ; marginal cuticle densely setose. Tritosternum with base about 24 p, laciniae 80 ». Presternal area with a pair of pre-endopodal shields. Sternal shield (135 » X 120 m) granular, thickened along lateral and anterior margins, with three pairs of setae and two pairs of pores ; meta- sternal setae and their associated pores situated over the endopodal shields. Between stI, 63 4; between str and s#3, 105 pw. Genital shield (120 » X 138 ») with genital setae only. Anal shield (80 » x 78 «) with maximum width 86 »; paranal setae | | | | THE BRITISH DERMANYSSIDAE (ACARI) 195 , postanal seta 30 p. Chaetotaxy and sclerotization of opisthogaster as in Text- B. Peritreme extends to about anterior third of coxa I. Peritrematal shield lM iM et a ta etotaxy of legs differs from normal pattern in that genu IV has two ventral (2—?, 31). Tarsus II with ventral setae and al, and #1, stout basally, ng to fine points. Tarsus IV with ventral setae stout basally, tapering ; seta b1, stout and spine-like. Length/width (in ,) of leg segments : I II III IV genu 66/33 70/45 42/33 60/33 tibia 80/30 60/42 42/30 63/33 tarsus 130/28 98 /33 105/27 135/27 Male: Chelicera with segment I, 45 » ; II, 108 »; movable digit 36 p (57 » with rmadactyl), unidentate ; fixed digit unidentate, pilus dentilis short, setiform. © pairs of gnathosomal setae with c.s. about 54 » apart; hyp.2 about 42 pw apart; j itosternum with six transverse rows of denticles ; corniculi 36 » ; internal malae _ tniangular. Tectum capituli weakly dentate ; pedipalp as in female. ii~,- i a | yo] : Fic. Sr. Hypoaspis colombot sp. nov., female; dorsum (a) and venter (B) of idiosoma. 196 G. OWEN EVANS & W. M. TILL Dorsal shield (546 » x 372 ~) asin female. Holoventral shield (420 p Xx I02 p) granular with four pairs of setae and three pairs of pores in the sternal region, one pair of genitals, and about 26 opisthogastric setae (Text-fig. 32D). Between sft, 68 » ; between sti and st3, 96 ». Paranal setae 42 pu, postanal seta 33 ». Peritreme extends to anterior fourth of coxa I; posterior part of peritrematal shield free. Chaetotaxy of legs as in female. Length/width (in y») of leg segments : I II III IV genu 60/32 63/45 40/27 54/30 tibia 70/27 54/40 42/27 57/27 tarsus 117/27 84/30 84/24 110/24 Hasitat: In nest of Bombus lapidarius (Linn.), Langley Park, Buckinghamshire (K.F. Colombo, 1961). Holotype 2 (1965 : 12 : 29: 1), allotype g (1965 : 12: 29: 2). Hypoaspis fuscicolens Oudemans Hypoaspis fuscicolens Oudemans, 1902, Tijdschr. ned. dierk. Ver. (2) 7: 289 (nom. nud.) ; r904, Abh. nat. ver. Bremen 18 : 83, pl. 4, figs. 23-28. Female ; Chelicerae retracted, details of their structure not clearly visible. Four pairs of gnathosomal setae with c.s. very stout and about 72 » apart, hyp.2 about 48 u apart (Text-fig. 32B). Deutosternum with six transverse rows of denticles (2 to about 8 per row); corniculi 45 »; internal malae triangular, fringed. Tectum capituli subtriangular with a minute antero-median projection, margin smooth. Pedipalp (2—5—6-14) with three-tined apotele. Dorsal shield (720 » x 480 p) with 24 pairs of setae in the podonotal region, an extra pair of v setae being present on the margin and a pair of accessory setae between 76. Opisthonotal region of shield hypertrichous (Text-fig. 32A). Surface of shield granular and reticulate. Tritosternum with base 45 p, laciniae 105 ». Sternal shield (84 » x 186 ,«) with three pairs of setae and two pairs of pores, metasternal setae free with associated pores. Between st1, 75 »#; between sti and sf3, 105 pw. Genital shield (130 p X 150 ») with genital setae only. Anal shield (132 » x 108 ,») with par- and postanal setae about 24 » long. Chaetotaxy and sclerotization of venter as in Text-fig. 32c. Peritreme extends onto dorsum, terminating at a point over posterior margin of coxa I; anterior part of peritrematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern in that genu IV has two ventral setae (2—2, 3—1). Coxae I and II with very stout setae ; genu and tibia II with pv stouter than av; tarus II with seta mv short and stout. Ambulacrum I without claws. Length/width (in «) of leg segments : if Il Ill IV genu go /50 96/60 66/57 93/57 tibia 96/50 80/63 72/50 99/50 tarsus 140/48 126/50 140/48 168 /45 THE BRITISH DERMANYSSIDAE (ACARI) | n= “4 aH coyet| oO sa SS =>2 <3 S Eo om ns Qe Mo tanE s nome) q Bo Zo as 8 v's ce = & 8 n =| EES ay O46 =} Ol eS gl 3: S oc Dae n Se} oo “44 Qo nH S 5 gt a5 m > Oo on eh As} if a 5 8 io) & shield. 198 G. OWEN EVANS & W.M. TILL Hasitat: Found with several species of Bombus and with Psithyrus vestalis (Geoff.) on Wangeroog Island, Germany. Hypoaspis hyatti sp. nov. Female ; Chelicera with segment I, 60 »; II, 120 »; movable digit 40 yp, biden- tate ; fixed digit with one large and two minute teeth ; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 57 » apart ; Ayp.2 about 42 p apart. Deutosternum with six transverse rows of deutosternal teeth (2-6 per row) ; corniculi 33 #3 internal malae as in Hypoaspis bombicolens. Tectum capituli with smooth anterior margin. Salivary styli reach almost to tips of corniculi. Pedipalp (2—5-6-. 14) with two-tined apotele. Dorsal shield (612-642 » X 396-432 ») with 43-44 pairs of setae as well as about 16 unpaired accessory setae between the J series; pxI-3 present in opisthonotal region (Text-fig. 33A). Setae simple, slightly longer in the anterior than in the pos- terior region. Surface of shield granular and reticulate. Tritosternum with base 30 p, laciniae go p. Sternal shield (84—90 » x 132-140 p) granular and reticulate, with three pairs of setae and two pairs of pores; meta- ae Pee a. IN ip, / oS Trl Ss ry / \ / - nN / f i j ey \ \\ a \ | [eee oe See ee We te, ia ] she » J \ : F. ih \Y a ) } } \ i A V ! } j \ N / ) ) \ | a | iN \ ’ Caley \ K fe 7 q- Fic. 33. Hypoaspis hyatti sp. nov., female ; dorsum (aA) and venter (B) of idiosoma. THE BRITISH DERMANYSSIDAE (ACARI) 199 sternal setae free with associated pores. Between sti, 72 ~; between sf1 and s?3, 108 ». A pair of well-defined pre-endopodal shields is situated in the pre-sternal region. Genital shield (130 » x 150 «) with genitalsetae only. Anal shield (93 » x 84 «) reticulate ; par- and postanal setae about 20 » long. Chaetotaxy and sclero- tization of opisthogaster as in Text-fig. 338. Peritreme extends to anterior third of coxa I; anterior part of peritrematal shield fused with dorsal shield ; posterior part free. Chaetotaxy of legs differs from the normal pattern in that genu IV bears two ventral setae (pv present). Length/width (in p) of leg segments : I II Ill IV genu 84/40 75/54 57/42 75/40 tibia 90/36 68/50 57/36 78/36 tarsus 140/33 110/36 117/33 150/33 Hapitat: On Bombus muscorum (Linn.), Scarborough, Yorkshire, October, 1964. Holotype @ (1965 :12: 29:11) and twenty-two female paratypes (1965 : 12: 29: 12-22) in the collection of the British Museum (Natural History). Hypoaspis marginalis (Willmann) Pneumolaelaps marginalis Willmann, 1953, Sitz. Osterr. Akad. Wiss. Abt. 1, 162 (6) : 465, figs. II-12. Female: Chelicera with segment I, 48 » ; II, 120 »; movable digit 42 p, biden- tate ; fixed digit tridentate, pilus dentilis short, setiform (Text-fig. 34c). Four pairs of gnathosomal setae with c.s. about 63 p apart, hyp.2 about 48 w apart. Deuto- sternum with six transverse rows of denticles (3-8 per row) (Text-fig. 34D) ; corniculi 30 #; internal malae with a pair of slender, triangular inner lobes and a pair of fimbriate outer lobes. Tectum capituli with anterior margin smooth. Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (624 » x 408 ,) with 25 pairs of setae in the podonotal region, three pairs of y setae apart from the humerals being included on the shield (Text-fig. 344). Opisthonotal region with 18 pairs of setae, px1 being present, and about 13 unpaired accessory setae between the / series. Setae near anterior tip of shield noticeably longer than those near posterior tip. Surface of shield granular and reticulate. Tritosternum with base 33 p, laciniae about 75 ». Presternal area with a pair of pre-endopodal shields. Sternal shield (93 » x 144 ») with three pairs of setae and two paires of pores. Metasternal setae and associated pores situated over endopodal shields. Between sti, 66 «7; between sf1 and sf3, 110 w. Genital shield (110 p x 147 ») with genital setae only. Anal shield (90 » x 90 ») with paranal setae 21 p, postanal seta 24 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 34B. Peritreme extends to anterior third of coxa I; posterior part of peritrematal shield free. 200 G. OWEN EVANS (& W.-M a TIE oe = ———- ~~ =~ — A ve é Le Rl Le Seal POS so Si = — ~ ™ ~ A A —_ == ae P 2? jp, l?I— 4 Wien gf iN A Fic. 34. Hypoaspis marginalis (Willmann). A. Dorsum of idiosoma of female ; B. venter of idiosoma of female ; c. fixed digit of chelicera of female; Db. deutosternum of female ; rE. chelicera of male; F. venter of idiosoma of male. EE — THE BRITISH DERMANYSSIDAE (ACARI) 201 Chaetotaxy of legs differs from the normal pattern in that genu IV has two ventral setae (2—2, }—1). Tarsi II-IV with ventral setae stouter than the dorsals, tarsus IV with setae p/,-, spine-like. Length/width (in ») of leg segments : I IME Ill IV genu 78/40 75157 60/44 78/42 tibia 84/36 66/50 57/40 78 /38 tarsus 132/33 102/38 117/36 156/33 Male: Chelicera with segment I, 48 » ; II, 110 1; movable digit 36 pu (57 » with spermadactyl), unidentate ; fixed digit unidentate, pilus dentilis short, setiform (Text-fig. 34E) ; groove of spermadactyl granular. Four pairs of gnathosomal setae with c.s. about 60 papart, hyp.2 about 42 wapart. Deutosternum with six transverse rows of denticles (5-6 per row) ; corniculi 36 ». Internal malae, tectum, capituli and pedipalps as in female. Dorsal shield (576 » x 348 p) with chaetotaxy as in female, but only about six accessory setae. Tritosternum with base 24 p, laciniae 84 ». Presternal area witha pair of pre-endopodal shields. Holoventral shield (456 » * 135 ») with four pairs of setae and three pairs of pores in the sternal region, with a pair of genital setae and 37-44 opisthogastric setae in the genito-ventral region (Text-fig. 34F). Between st, 66 1; between sf1 and st3, 102 w. Peritreme extends to about middle of coxa I; anterior part of peritrematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs as in female. Femur II with seta av, very stout and spine-like ; genu II and tibia II with seta av strongly inflated basally ; tarsus II with setae av,_, and mv inflated basally, tapering to long fine points. Length/width (in p) of leg segments : I i (ig i IV genu 78/36 78/60 60/42 78/42 tibia 84/36 66/54 54/36 72/30 tarsus 132/30 96/36 96/30 138/30 Hasitat: Found on Bombus terrestris (Linn.) in Essex, in the nest of B. hortorum (Linn.) in Hertfordshire, and on Bombus mucidus (Gerst.) in Austria. Hypoaspis minutissima sp. nov. Female : Chelicera with segment I, 42 »; II, 102 »; movable digit 36 pu, biden- tate ; fixed digit with bifid tip and three small teeth, pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 50 p apart, hyp.2 about 36 p apart. Deutosternum with six transverse rows of denticles ; corniculi 42 » ; internal malae slender, indistinct. Tectum capituli with anterior margin smooth. Pedipalp (2-5-6-14) with two-tined apotele. 202 G. OWEN EVANS & W.M. TILL Fic. 35. Hypoaspis minutissima sp. nov., female ; dorsum (A) and venter (B) of idiosoma. Dorsal shield (492 » X 312 ») with 39 pairs of setae and about five accessory setae between the J series (Text-fig. 354). Setae simple, subequal. Surface of shield granular and reticulate. Tritosternum with base 24 p, laciniae 84 uw. Presternal area with a pair of pre- endopodal shields. Sternal shield (105 » x 105 «) granular with faint reticulations, bearing three pairs of setae and two pairs of pores. Metasternal setae and associated pores situated over the endopodal shields. Between sf1, 65 « ; between sii and sf3, 80 ». Genital shield (102 » X I14 ») with genital setae only. Anal shield (60 zh X 75 p) with paranal setae 15 », postanal seta 18 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 35B. Peritreme extends to anterior third of coxa I; anterior tip of peritrematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs differs from normal pattern in that genu IV bears two ventral Tarsi II and IV with ventral setae relatively long, stout basally, tapering to setae. Seta fl, on tarsus IV long, spine-like. Length/width (in ,) of leg fine points. segments : | | | | | | | | | | | THE BRITISH DERMANYSSIDAE (ACARI) 203 I II II IV genu 60/30 60/36 42/27 54/27 tibia 72 [27 50/33 42/24 54/24 tarsus 108/27 80/27 96/24 17/24 Hapitat: On a Bombus terrestvis (Linn.) queen, Gosforth, Cumberland, Britain, collected by D. C. Lee, 3rd May, 1960. Holotype 2 (1965 : 12 : 29: 5) and paratype BPR(TOO5 ; 12 : 29 : 6). Hypoaspis isotricha (Kolenati) Holostaspis isotricha Kolenati, 1858, Wien. Ent. Monatschr. 2 : 87, figs. Female : Chelicera with segment II, 66 »; movable digit 18 », with one minute, subterminal tooth ; fixed digit with one small tooth ; pilus dentilis very short and setiform (Text-fig. 36D). Four pairs of gnathosomal setae with c.s. about 50 p apart, hyp.2 about 36 » apart. Deutosternum with six transverse rows of denticles ; corniculi 18 » long ; internal malae asin Text-fig. 36£. Tectum capituli with smooth anterior margin (Text-fig. 368). Pedipalp (2-5—6-14) with two-tined apotele. Dorsal shield (612 » X 432 ») with 39 pairs of setae and two unpaired accessory setae distributed as in Text-fig. 364. All the setae except 71 and z1 are barbed (Text-fig. 36c). Surface of shield sculptured anteriorly, reticulate posteriorly. Tritosternum with base 36 yp, laciniae about 72 ». Sternal shield (96 w x IT4 p) with three pairs of setae and two pairs of pores ; metasternal setae and pores situated over the endopodal shields. Between sf1, 60 « ; between sti and s#3, 108 ». Genital shield (168 » X 150 p) with genital setae only. Anal shield (84 4 x 80 p) concave anteriorly ; paranal setae 36 », postanal seta 60 p. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 36F. Peritreme extends approximately to anterior margin of coxa II ; posterior part of peritrematal shield free. Legs with normal chaetotaxy. Dorsal seta on trochanter I stout and spine-like ; seta ad, on femora II-IV relatively stout and spine-like. Length/width (in p) of leg segments : if II TEL IV genu 63/24 42/40 45/35 60/32 tibia 68/18 45/33 42/30 57/27 tarsus 123/19 80/25 78/24 110/20 Hapitat: Ants’ nests (Formica sp.) in the Black Wood of Rannoch, Perthshire, Scotland, and in Europe. ZOOL. 14, 5. 14 204 G. OWEN EVANS & W. M. TILL Fic. 36. Hypoaspis isotricha (olenti), female ; dorsum of idiosoma (a); tectum (B) ; dorsal seta (c) ; chelicera (D); venter of gnathosoma (E); venter of idiosoma (F). Fig. 37. THE BRITISH DERMANYSSIDAE (ACARI) Hypoaspis montana (Berlese), female ; dorsum of idiosoma (A) ; venter of gnathosoma (c) ; venter of idiosoma (p). chelicera (B) ; 206 G. OWEN. EVANS. & W. M. TILL Hypoaspis montana (Berlese) Laelaps (Oolaelaps) montanus Berlese, 1904, Redia 1 : 430. Female : Chelicera with segment I, 20 » ; II, 75 » ; movable digit 21 p, edentate ; fixed digit unidentate ; pilus dentilis absent, dorsal seta and arthrodial processes present (Text-fig. 37B). Four pairs of gnathosomal setae with c.s. about 36 p apart, hyp.2 about 30 » apart. Deutosternum with six transverse rows of denticles (4-8 denticles per row) ; corniculi horn-like, bifid at tip, about 20 » long ; internal malae fimbriate (Text-fig. 37c). Tectum capituli with anterior margin smooth. Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (468 » < 312 ») with 39 pairs of setae distributed as in Text-fig. 374 ; a pair of accessory setae situated between the J series. Setae Z5 pilose distally and about three times the length of simple seta /5. Anterior part of shield “ sculptured ”, reticulations and striations distinct only in the marginal zone. Tritosternum ‘with base 20 p, laciniae 54 ». Sternal shield (105 » x I00 ») with weak striations laterally, bearing three pairs of setae and two pairs of pores ; meta- sternals free with associated pores. Between sti, 50 »; between sti and s¢3, 87 pu. Genital shield (110 » X 95 p») with genital setae only. Anal shield (60 p x 57.) with paranal setae 24 p, postanal seta 30 w long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 37D. Peritremes extend to middle of coxa I; peri- trematal shields free posteriorly. Chaetotaxy of legs normal. Length/width (in ) of leg segments : iI II IBLE IV genu 63/30 51/36 42/28 60/30 tibia 70/30 45/33 40/26 60/28 tarsus 114/27 78/24 70/24 96/24 approx. Hapitat: Ants’ nest near Canterbury, Kent, 25th April, 1942, coll. Dr. E. Warren. Recorded from nests of Formica rufa Linn., F. fusca Linn. and Lasius niger (Linn.) in Britain and Ireland, and from ants’ nests in Europe. Hypoaspis oophila (Wasman) Laelaps oophilus Wasman, 1897, Zool. Anz. 20: 172. Female: Chelicera with movable digit 18 » long, bidentate ; fixed digit uniden- tate ; pilus dentilis and dorsal seta short, setiform ; arthrodial processes reduced (Text-fig. 388). Four pairs of gnathosomal setae with c.s. about 54 m apart, /ivp.2 about 42 papart. Deutosternum with six transverse rows of denticles (5-7 denticles per row); corniculi horn-like, 21 » long; internal malae weakly serrate basally (Text-fig. 38D). Tectum capituli with smooth margin (Text-fig. 38c). Pedipalp (2-5-6-14) with two-tined apotele. Dorsal shield (600 » x 438 ~) with 39 pairs of setae distributed as in Text-fig. 384 ; THE BRITISH DERMANYSSIDAE (ACARI) Fic. 38. Hypoaspis oophila (Wasman), female ; dorsum of idiosoma (A) ; chelicera (B) ; tectum capituli (c) ; deutosternum and hypostome (p) ; venter of idiosoma (E). 207 208 G. OWEN EVANS & W.M. TILL a single accessory seta present between the J series. Setae markedly unequal in - length; J4-5, Z3-5 and S2-5 pilose distally and considerably longer than the remaining simple setae. Surface of shield reticulated. Tritosternum with base 36 p, laciniae 60 ». Sternal shield (102 w X 117 pw) with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sii, 66 »; between sti and s¢3, I10 ». Genital shield (153 » X 140 p) tapers posteriorly and bears a pair of genital setae only. Anal shield (80 » x 72 py; maximum width 96 ») with paranal setae 33 p», postanal seta 48 wlong. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 38E. Peritremes extend to a point above coxae I ; peritrematal shields free posteriorly. Chaetotaxy of legs normal. Length/width (in ») of leg segments : i II IDEN IV genu 54/21 40/45 38/42 48/36 tibia 60/18 36/33 33/38 48/30 tarsus 108/21 80/24 84/24 102/24 HasitatT: Nests of Formica fusca Linn., Leeds, England (Hull Collection) ; recorded from several Formica species in Britain and Europe. Hypoaspis astronomica (Koch) Iphis astvonomicus Koch, 1839, Deutschl. C.M.A. fasc. 27, t. 18. Laelaspis ovatus Willmann, 1951, S.B. Akad. Wiss. Wien. 160: 115, fig. 7 (Syn. nov.). Female: Chelicera with segment I, 45 »; II, 150 1; movable digit 36 p, biden- tate ; fixed digit with about five teeth, pilus dentilis minute, setiform (Text-fig. 39). Four pairs of gnathosomal setae with c.s. about 45 » apart ; Ayp.2 about 36 pw apart. Deutosternum with six transverse rows of denticles ; corniculi 30 » long ; hypostome with about six pairs of finger-like processes (Text-fig. 39D). Tectum capituli with anterior margin apparently smooth (Text-fig .39B). Salivary styli slender, reaching tips of corniculi. Pedipalp (2-5—6-14) with two-tined apotele. Dorsal shield (552 » 396 ») with 39 pairs of setae and three unpaired accessory setae (Text-fig. 394). Setae of the s and S series and seta Z5 relatively long, stout and barbed. Surface of shield reticulate. Tritosternum with base 20 p, laciniae 72 ». Sternal shield (96 » X 96 ) with heavily sclerotized margins; with three pairs of setae and two pairs of pores; metasternal setae on small shields, associated pores on cuticle. Between siI, 63 «; between sf1 and s#3, 84 ». Genito-ventral shield (186 « x 186 «) greatly expanded behind coxae IV, with three pairs of setae. Anal shield (78 » X 102 «) with maxi- mum width at anterior margin 126 #; paranal setae 2I pu, postanal seta 33 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 39F. Peritreme ex- Fic. 39. Hypoaspis astronomica (Koch), female ; dorsum of idiosoma (aA); tectum (8) ; chelicera (c) ; venter of gnathosoma (pD); venter of idiosoma (E). SH DERMANYSSIDAE (ACARI) 210 G. OWEN EVANS 3& (WME! Did tends to anterior margin of coxa I ; anterior part of peritrematal shield fused with ~ dorsal shield, posterior part free. Chaetotaxy of legs normal. Length/width (in p) of leg segments : I I ul IV genu 70/27 48/33 42/33 60/30 | tibia 75,24 48/33 40/27 60/27 | tarsus 126/20 93/27 93/24 130/24 Hapitat: Ants’ nests in Britain (Cornwall) and in Germany. Hypoaspis equitans (Michael) Laelaps equitans Michael, 1891. Proc. zool. Soc. Lond. : 649. Female: Chelicera with segment I, 42 »; II, 126 1; movable digit 42 1; both digits virtually edentate with two slight irregularities on the internal margin ; pilus dentilis short, setiform (Text-fig. 40s). Four pairs of gnathosomal setae with c.s. about 50 pw apart, hyp.2 about 40 w apart. Deutosternum with six transverse rows of denticles ; corniculi 27 » long; internal malae with inner and outer finger-like processes and shorter fimbriae between these. Salivary styli and tectum capituli not clearly visible. Pedipalp (2-5-—6-14) with two-tined apotele. Dorsal shield (672 » x 486 p) with 39 pairs of setae and six unpaired accessory setae distributed as in Text-fig. 404. Most of the setae, especially in the opisthonotal region, are long and sinuous. Surface of shield reticulate. Tritosternum with base 30 p, lacininae 66 ». Sternal shield (135 » X 144 p) with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between sti, 78 uw; between sf and s¢3, 108 pw. Genito-ventral shield (210 « X 222 m) expanded behind coxae IV, with two pairs of setae. Anal shield (126 » X 105 wv) with maximum width 150 » at anterior margin ; paranal setae 36 p, postanal seta 85 » long. Chaetotaxy and sclerotization of opisthogaster as in Text- fig. 40c. Peritreme extends beyond posterior margin of coxa I. Legs with normal chaetotaxy. Length/width (in ,) of leg segments : I II Ill IV e| genu 102/40 78/48 972/45 go /40 | tibia 120/36 72/42 72/42 93/36 tarsus 198/33 132/30 144/30 180/30 Hapitat: Ants’ nests [Tetramorium caespitum (Linn.)] in Britain (Cornwall, Lundy Island Berkshire). | i THE BRITISH DERMANYSSIDAE (ACARI) Bic. 40. a-c. Hypoaspis equitans (Michael), female ; dorsum of idiosoma (A) ; chelicera (B); venter of idiosoma (c). D-F. Hypoaspis humerata (Berlese), female ; chelicera (D) ; dorsum of idiosoma (E) ; venter of idiosoma (F). 212 G. OWEN EVANS & W.™M. TILL Hypoaspis humerata (Berlese) Laelaps (Laelaspis) humeratus Berlese, 1904, Redia 1: 425, pl. 18, figs. 151-153. Female : Chelicera with segment II, 110 » ; movable digit 36 p, bidentate ; fixed digit tridentate (Text-fig. 40D). No other details of the gnathosomal structure can be described owing to the poor condition of the only available specimen. Dorsal shield (approximately 600 » * 490 ») with 39 pairs of setae distributed as in Text-fig. 4oE, and three unpaired accessory setae. Surface of shield reticulate. Sternal shield (90 » x 120 ) with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between st1, 63 »; between sfi and st3, 84 ». Genito-ventral shield (210 » x 198 ) expanded behind coxae IV with maximum width 282 » ; with two pairs of setae, a third pair being just on or off the margin of the shield. Anal shield (84 » x go pw) with maximum width 123 pu at anterior margin ; paranal setae 30 p long, postanal seta broken. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 4or. Peritreme extends approximately to middle of coxa I; posterior part of peritrematal shield free. Legs apparently with normal chaetotaxy, although this cannot be checked on every segment. Legs rather short and stumpy. Length/width (in ) of leg segments : I II III IV genu 45/27 40/54 33/45 45/42 tibia 48/24 36/42 33/42 45/36 tarsus 100/24 75/30 75/33 102/30 Hapitat: Found in ants’ nests [Tetvamorium caespitum (Linn.)] in Britain (Cornwall) and in Luxemburg. Hypoaspis acuta (Michael) Laelaps acuta Michael, 1891, Pyoc. zool. Soc. Lond. : 652. Female : Chelicera with segment I, 60 »; II, 177 »; movable digit 57 », biden- tate ; fixed digit with four teeth ; pilus dentilis short, setiform ; dorsal seta present (Text-fig. 41F). Four pairs of gnathosomal setae with c.s. about 66 » apart ; /yp.2 about 48 » apart. Deutosternum with five transverse rows of denticles, each row multidenticulate ; corniculi horn-like, 40 » long; internal malae fimbriate (Text- fig. 41E). Tectum capituli with anterior margin apparently smooth. Pedipalp (2-56-14) with two-tined apotele. Dorsal shield (720 » 468 p) with 37 pairs of setae distributed as in Text-fig. 414 ; setae 23 and ri lacking on the podonotal region ; about four accessory setae present on the opisthonotal region between the J series. Setae inflated sub-basally (Text- fig. 4IC). Fic. 41. Hypoaspis acuta (Michael). a. Dorsum of idiosoma of female; 8B. chelicera of male; c. dorsal seta 25 of female; p. holoventral shield of male; ©. deutosternum and hypostome of female ; F. chelicera of female; G. venter of idiosoma of female. { | | | THE BRITISH DERMANYSSIDAE (ACARI) 214 G. OWEN- EVANS & W. M: TILL Tritosternum with base 48 p, laciniae go pw. Sternal shield (140 p X 150 yp) © granular, reticulated laterally, bearing three pairs of setae and two pairs of pores. Metasternal setae and pores situated on cuticle overlying the endopodal shields. Between st1, 80 »; between sti and st3, 129 w. Genital shield (168 » X 170 4) flask-shaped with genital setae only. Anal shield (105 p» x 100 yp) pear-shaped, paranal setae 36 p, postanal seta 48 » long. Chaetotaxy and sclerotization of opis- thogaster as in Text-fig. 41G. Peritremes extend to anterior third of coxae I; per- trematal shields free posteriorly. Chaetotaxy of legs normal except genu IV which has two postero-lateral setae (2—#?, 3—2). Length/width (in p) of leg segments : I Il Ill IV genu 108/36 75/48 70/40 114/40 tibia 120/30 78/40 72 [36 117/36 tarsus 192/30 140/33 147/32 228/33 Male : Chelicera with segment I, 48 » ; II, 123 » ; movable digit dentate, partially fused with a grooved spermadactyl which measures 60 » in length from the base of the movable digit (Text-fig. 41B). Pilus dentilis and dorsal seta as in female. Four pairs of gnathosomal setae with c.s. about 54 w apart; hyp.2 about 45 mw apart. Deutosternum with about four transverse rows of denticles; corniculi horn-like, 32 » long ; internal malae fimbriate. Pedipalps and tectum as in female. Dorsal shield (582 » 354 ») with chaetotaxy as in female. Holoventral shield (450 » X 108 pu) expanded behind coxae IV, bearing nine pairs of setae excluding the anals (Text-fig. 41D). Between sti, 72 «4; between sti and st3, 100 pw. Paranal setae 36 p, postanal seta 30 » long. Peritremes extend to middle of coxae I. Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : I uA Ill IN; genu 87/33 66/40 57/33 93/33 tibia 102/28 63/36 57/30 99/33 tarsus 162/27 114/30 120/24 186/27 Hasitat: In the nests of Myrmica scabrinodis Nylander in Mayo, Ireland, and Camponotus herculeanus (Linn.) in Europe. Hypoaspis myrmecophila (Berlese) Laelaps myrmecophilus Berlese, 1892, Acari, Myr. Scorp. Ital. fasc. 69, no. 2. Laelaps ovalis (Moniez), Berlese, 1904, Redia 1: 409. Female: Chelicera with segment I, 150-190 »; II, 324-336 »; movable digit 135 p, bidentate ; fixed digit with about nine teeth, pilus dentilis short, setiform (Text-fig. 42). Four pairs of gnathosomal setae with c.s. about 84 « apart, /wvp.2 about 70 w apart. Deutosternum with six transverse rows of denticles ; corniculi 98 «; internal malae with slender inner and triangular median, pilose lobes, and | THE BRITISH DERMANYSSIDAE } i (ACARI) 215 Fic. 42. Hypoaspis myrmecophila (Berlese). a. Dorsum of idiosoma of female ; B. venter of idiosoma of female; c. apotele of pedipalp; pb. venter of gnathosoma of female ; E. chelicera of female; r. chelicera of male. ee 216 G. OWEN EVANS & W. Me TILL long, slender, outer lobes (Text-fig. 42D). Tectum capituli with denticulate anterior margin. Salivary styli long, slender, reaching approximately to tips of corniculi. Pedipalp (2-5-6-14) with three-tined apotele (Text-fig. 42c). Dorsal shield (1,116 * 768 ) with 41 pairs of slender, simple setae and five unpaired accessory setae (Text-fig. 424) ; an extra pair of 7 setae is present in the podonotal region and setae pxI are present in the opisthonotal region. Surface of shield granular, with sculpturing anteriorly. Tritosternum with base 60 p, laciniae 168 p» ; pre-endopodal shields large, reticulate and granular. Sternal shield (180 » x 240 p) granular, reticulate antero-laterally, with three pairs of setae and two pairs of pores ; metasternal setae free with associa- ted pores. Between sf1, 126-132 » ; between sf and s¢3, 180-192 ». Genito-ventral shield (290 » X 246 ») with one pair of genital and one pair of opisthogastric setae. Anal shield (198 » X 120 ») with maximum width 186 »; paranal setae 48 np; postanal seta 30 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 42B. Peritreme extends to about posterior third of coxa I; posterior part of peri- trematal shield free. Legs with normal chaetotaxy. Relatively stout setae present on femur I (fd5_,), tarsus II (ventrals, al,, pl,, md), and femur IV (ad, and pd,). Seta p/ on genu IV stout and spine-like (Text-fig. 448). Length/width (in p) of leg segments : I II 12H IV genu 192/66 156/90 126/66 180/78 tibia 222/60 144/80 120/66 180/78 tarsus 300/54 228/63 228/54 306/63 Male : Chelicera with segment I, 108 »; II, 300 1; movable digit 126 yu, uniden- tate ; spermadactyl 140 p, a small tooth near its tip ; fixed digit with one large and about five small teeth ; pilus dentilis short, setiform (Text-fig. 42F). Four pairs of gnathosomal setae with c.s. about 78 » apart, hyp.2 about 75 w apart. Corniculi 90 long, internal malae not clearly visible. Deutosternum, salivary styli, tectum capituli and pedipalps as in female. Dorsal shield (1,080 » X 756 ») asin female. Holoventral shield (816 » x 216 p) with four pairs of setae and three pairs of pores in the sternal region, one pair of genital and five pairs of opisthogastric setae in the genito-ventral region (Text-fig. 44A). Between sfI, 114 »; between sfI and st#3, 195 ». Paranal setae 60 p, post- anal seta 42 ». Peritreme extends to posterior margin of coxa I. Posterior part of peritrematal shield free. Legs with normal chaetotaxy, stout setae asin female. Length/width (in pz) of leg segments : I Il Ill IV genu 180/66 162/90 132/72 192/84 tibia 210/60 150/84 126/66 198/78 7 tarsus 270/54 220/60 240/54 300/60 THE BRITISH DERMANYSSIDAE (ACART) 217 - Hasirat: Found in ants’ nests in Britain (Cornwall, Devon, Wales, Isle of Wight) ~ and Europe. en / i \ : , ae Gs a ee eer eet) Fic. 43. A-B. Hypoaspis myrmophila (Michael), female ; dorsum (a) and venter (8) of idio- soma. C-—D. Hypoaspis laevis (Michael), female ; venter (c) and dorsum (D) of idiosoma. 218 G. OWEN EVANS & W. M. TILL Hypoaspis myrmophila (Michael) Laelaps myrmophila Michael, 1891, Proc. zool. Soc. Lond. : 649. Female ; Chelicera with segment I, 90 » ; II, 246 » ; movable digit 105 yp, biden- tate; fixed digit with four conspicuous teeth and a row of 3-5 tiny teeth; pilus dentilis and dorsal seta normal. Four pairs of gnathosomal setae with c.s. about 66 » apart, hyp.2 about 63 » apart. Deutosternum with six transverse rows of denticles, each row multidenticulate ; corniculi horn-like, 63 » long ; internal malae as in H. myrmecophila, but middle process almost as long as inner, and outer process forkedat tip. Tectum capituli with anterior margin serrated. Pedipalp (2-5-6-14) with three-tined apotele. Dorsal shield (696 x 528 ») with 40 pairs of setae and seven unpaired accessories distributed as in Text-fig. 434 ; an extra pair of 7 setae is present in the podonotal region. Tritosternum with base 40 yp, laciniae 114 y. Pre-endopodal shields present. Sternal shield (96 » x 180 p) reticulated antero-laterally, with three pairs of setae and two pairs of pores; metasternal setae and associated pores situated over the endopodal shields. Between sf1, 99 » ; between sfz and s?#3, 105 ». Genito-ventral shield (210 » X 190 p) reticulated, bearing a pair of genital and two pairs of opistho- gastric setae. Anal shield (114 » X 99 #; maximum width 138 ») with the usual three setae. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 438. Peritreme extends to about middle of coxa I; posterior part of peritrematal shield free. Legs with normal chaetotaxy. Length/width (in ,) of leg segments : I Il INGE IV genu 105/48 96/60 66/42 114/48 tibia 126/45 90/57 69/42 114/48 tarsus 198/42 150/42 140/36 180/38 Male: Chelicera with segment I, 78 »; II, 210 »; movable digit 84 yu, uniden- tate ; spermadactyl relatively short, 102 » from base of movable digit ; fixed digit with one large and two small teeth (Text-fig. 44D). Puilus dentilis and dorsal seta normal. Four pairs of gnathosomal setae with c.s. about 60 » apart, iyp.2 about 57 « apart. Corniculi horn-like, 54 » long; internal malae pilose. Deutosternum, tectum and pedipalps as in female. Dorsal shield (612 » X 420 «) with chaetotaxy as in female. Pre-endopodal shields present. Holoventral shield (516 » X 144 ) with ten pairs of setae, exclud- ing the anals (Text-fig. 44£). Between sti, 87 w; between sti and sf3, I17 jh Peritreme extends to about middle of coxa I. Legs with normal chaetotaxy. Genu and tibia II with seta fv stout ; tarsus II with ventral setae and al, and #/, stout. Length/width (in pz) of leg segments : ————————— oe THE BRITISH DERMANYSSIDAE (ACARI) 219 Fic. 44. a. Hypoaspis myyrmecophila (Berlese), holoventral shield of male ; B. H. myymeco- Phila, genu IV of female ; c. Hypoaspis laevis (Michael), genu LV of female ; p. Hypoaspis myrmophila (Michael), chelicera of male; E. H. myymophila, holoventral shield of male. ZOOL. 14, 5. T5 220 G. OWEN EVANS & W. M. TILL I II Ill IV genu 96/45 90/57 66/38 99/45 tibia 111/38 84/51 63/39 105/45 tarsus 180/36 138/38 126/32 162/36 Hapsitat: The nests of ants [Laszus mixtus Nylander, L. flavus (Fabr.), Formica fusca Linn., Tetramorium caespitum (Linn.)| in Cornwall, Surrey and Northumber- land; ants’ nests in Corsica. Hypoaspis laevis (Michael) Laelaps laevis Michael, 1891, Proc. zool. Soc. Lond. : 648. Female : Chelicera with segment I, 90 »; II, 324 »; movable digit 130 p, biden- tate ; fixed digit with three prominent teeth and a row of about four small ones. Pilus dentilis and dorsal seta short, setiform. Four pairs of gnathosomal setae with c.s. about 102 » apart, hyp.2 about 78 » apart. Deutosternum with five transverse rows of denticles (4-9 denticles per row). Corniculi horn-like, 80 p» long ; internal malae comprise pilose outer, middle and inner processes. Tectum capituli with anterior margin denticulate. Pedipalp (2-5—6-14) with three-tined apotele. Dorsal shield (1,140 » X 770 ») with 41 pairs of setae and about six accessories distributed as in Text-fig. 43D ; an additional pair of 7 setae is present on the podono- tal region and an extra pair of px setae on the opisthonotal region. Surface of shield granular with some striations near the margin. Tritosternum with laciniae about 23 times the length of the base. Pre-endopodal shields present. Sternal shield (204 » x 246 y) reticulated antero-laterally, with three pairs of setae and two pairs of pores ; metasternal setae situated over the endo- podal shields. Between sti, 70 »; between sf and s#3, 192 w. Genito-ventral shield (312 » X 237 ») reticulated, with one pair of genital and two pairs of opistho- gastric setae. Anal shield (186 » X 132 »; maximum width 195 yz) with paranal setae 45 pw, postanal seta 21 » long. Sclerotization of opisthogaster as in Text-fig. 43c. Peritreme extends at least to anterior margin of coxa I ; peritrematal shield fused anteriorly with dorsal shield, posterior part free. Chaetotaxy of legs normal except genu IV which has two postero-lateral setae (2—?, 3—2) as in Text-fig. 44c. Tarsus II with several stout setae (av,, pv,, al, pl,). Length/width (in p) of leg segments : I II Tey IV genu 180/66 144/87 126/66 180/75 tibia 222/57 150/80 126/60 198/75 tarsus 306/54 228/57 240/48 318/57 Hasirat: Nests of ants [Tetramorium caespitum (Linn.)] in Cornwall and in Austria. THE BRITISH DERMANYSSIDAE (ACARI) eee. Fic. 45. Hypoaspis miles (Berlese), female ; dorsum of idiosoma (a); tecta capituli (B) ; chelicera (c) ; venter of gnathosoma (Dp) ; venter of idiosoma (E). G. OWEN EVANS & W. M.-TILL nN N N Hypoaspis miles (Berlese) Laelaps (Iphis) miles Berlese, 1892, Acari, Myr. Scorp. Ital. fasc. 63, no. 9. Cosmolaelaps gurabensis Fox, 1946, J. Parasit. 32 : 449, fig. 1. Bregetova, 1956, Opred. Faune SSSR. 61: 75, figs. (syn. nov.). Female : Chelicera with segment I, 96 »; II, 216 1; movable digit 130 pu long, bidentate ; fixed digit with about four teeth ; pilus dentilis short, setiform (Text-fig. 45C). Four pairs of gnathosomal setae with c.s. about 63 » and hyp.2 about 66 pu apart. Deutosternum with six transverse rows of denticles ; corniculi 90 p long ; outer processes of internal malae fringed (Text-fig. 45p). Tectum capituli with denticulate anterior margin (Text-fig. 458). Salivary styl conspicuous, almost reaching tips of corniculi. Pedipalp (2-5—6—14) with two-tined apotele. Dorsal shield (648 » x 348 ») with 37 pairs of leaf-like setae distributed as in Text-fig. 454 ; setae px2 and px3 absent. Surface of shield reticulated ; no hyper- trichy ; seven pairs of R setae on the cuticle. Tritosternum with base 60 p, laciniae 96 pw. Sternal shield (180 pu x 138 pz) extending almost to posterior margin of coxa III, with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sti, 96 » ; between st1 and st3, 144 ». Genital shield flask-shaped (135 » X 102 pu), with one pair of setae. Anal shield (72 » x 87 u) with paranal setae about 24 m, postanal seta about 15 p. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 45E. Peri- treme extends nearly to anterior margin of coxa I; peritrematal shields free an- teriorly and posteriorly. Legs with normal chaetotaxy. Length/width (in p) of leg segments : I 161 ET IV genu 87/30 84/50 60/33 93/36 tibia 93/24 78/48 60/33 96/36 tarsus 170/27 144/36 135/30 186/30 Hapitat: Found in decaying oat spillage at Leith, Scotland. Recorded in the literature from a variety of rodents and their nests in Europe, the U.S.S.R. and the USA. Hypoaspis oblonga (Halbert) Laelaps (Hypoaspis) oblongus Halbert, 1915, Proc. R. Ivish Acad. 31: 70. Pseudoparasitus angulatus Berlese, 1917, Redia 12: 164 (syn. nov.). Pseudoparasitus (Alloparasitus) angulatus Berlese, 1921, Redia 14: 169. Female: Chelicera with segment I, 60 »; I, 162 » ; movable digit 68 ,, biden- tate ; fixed digit with about six teeth (Text-fig. 468) ; pilus dentilis and dorsal seta short, setiform. Four pairs of gnathosomal setae with c.s. about 42 apart, /iyp.2 about 40 papart. Deutosternum with six transverse rows of denticles (4-6 denticles per row). Corniculi horn-like, 42 » long ; internal malae fimbriate (Text-fig. 465). | | | | | | | THE BRITISH DERMANYSSIDAE (ACARI) Y . = ee a case ; A a ae Sa Van os . : i‘ / ° a7 pa leg daa E F Fic. 46. Hypoaspis oblonga (Halbert), female; dorsum of idiosoma (a) ; chelicera (B) ; tectum capituli (c) ; apotele of pedipalp (D) ; venter of gnathosoma (£); venter of idiosoma (F). 12 VPP PPE ARTE CR NT ae 224 G. OWEN EVANS & W. M. TILL Tectum capituli with anterior margin denticulate (Text-fig. 46c). Pedipalp (2-5-6- - 14) with two-tined apotele (Text-fig. 46D). Dorsal shield (684 » x 408 ) with 39 pairs of setae distributed as in Text-fig. 46a ; no hypertrichy. Surface of shield reticulated. Tritosternum bipartite ; pre-endopodal shields present. Sternal shield (150 » x 132 p) reticulated, with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sti, 70 »; between sti and s#3, 120 p. Genito- ventral shield (204 » X 144 ») expanded behind coxae IV and bearing a pair of genital and a pair of opisthogastric setae. Anal shield (100 » x 100 w; maximum width 156 ) ; paranal setae 36 u long, postanal seta slightly longer. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 46F. Peritreme extends to middle of coxa I; peritrematal shields extend posteriorly well beyond level of coxae IV, nee: Legs with normal chaetotaxy, except genu IV which has only one antero-lateral seta (I—?, 3—1). Hasitat: Under bark of decayed trees and in moss, in Ireland, Scotland, England (Northumberland) and Europe. Genus PSEUDOPARASITUS Oudemans Pseudopayasitus Oudemans, 1902, Tijdschr. Ent. 45 : 29. Hoplolaelaps Berlese, 1903, Zool. Anz. 27: 14. Laelapsoides Willmann, 1952, Vevoff. Inst. Meersforsch. Bremerhaven 1 : 150. Austrogamasellus Domrow, 1957, Proc. Linn. Soc. N.S.W. 81: 204, (syn. nov.). Type: Laelaps meridionalis Canestrini, 1882. This genus is closely related to Ololaelaps Berlese and differs chiefly in the presence of a discrete anal shield in the female. Chelicerae chelate-dentate in the adults wit the movable digit bidentate in the female and unidentate in the male. Pilus dentilis and dorsal seta simple ; fissures normal. Spermadactyl long, slender, groove direc- ted anteriorly. Chaetotaxy of venter of gnathosoma and pedipalps normal. Apotele three-tined, posterior tine reduced. Deutosternum with six transverse rows of denticles; internal malae strong; corniculi horn-like. Tectum capituli with anterior margin denticulate. Dorsal shield entire with 39 pairs of setae (posterior accessories present) ; no unpaired accessory setae in the region of the J series. Tritosternum normal, bi- partite. Pre-endopodal shields conspicuous. Sternal shield in the female with three pairs of setae and two pairs of pores. Genito-ventral shield large with four or more pairs of setae of which two pairs are widely removed from its lateral margins. Anal shield free, paranals at level of anterior margin of anus. Podal shields posterior to coxae IV large, not fused with the peritrematal shield. Male with holoventral shield ; peritrematal shields free posteriorly. Leg chaetotaxy normal, seta av, on femur IIT spine-like in the female, spur-like in the male. Coxa II without distinct anterior spine. Ambulacra with paired claws. We have not studied the immature stages of this genus. | | THE BRITISH DERMANYSSIDAE (ACARI) 225 The females of the two British representatives of Pseudoparasitus may be readily distinguished by the size and chaetotaxy of the genito-ventral shield, P. centralis having five pairs of setae on the shield and P. dentatus four pairs. P. centralis may be a synonym of P. meridionalis, but we are unable to reach a definite conclusion on this since the type material of merzdionalis appears to be lost. Pseudoparasitus centralis Berlese Pseudoparasitus centralis Berlese, 1921, Redia 14: 167. Female : Chelicera with segment I, 66 »; II, 162 « ; movable digit 66 p, biden- tate ; fixed digit unidentate ; pilus dentilis short, setiform (Text-fig. 47D). Four pairs of gnathosomal setae with c.s. about 45 » apart; Ayp.2 about 42 pw apart. Deutosternum with six transverse rows of denticles (I-3 per row) ; corniculi 40 p long ; internal malae finger-like and denticulate (Text-fig. 47F). Tectum capituli with denticulate anterior margin. Salivary styli do not quite reach tips of cornicull. Pedipalp (2-5-6—14) with three-tined apotele. Dorsal shield (588 » 420 ») with 39 pairs of setae distributed as in Text-fig. 474. Surface of shield reticulate and granular ; no hypertrichy. Tritosternum with base 27 p, laciniae about 78 pw. Sternal shield (126 » x I14 p) extends to posterior margin of coxa III, with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sfI, 54 »; between sé and st3, 99 ». Genital shield (180 » x 138 p) flask-shaped, expanded behind coxae IV, with truncate posterior margin, bearing five pairs of setae. Between gen., 120 p ; between Jv2, 114 pw. Anal shield (96 » x 105 ») with a straight anterior margin measuring 180 ». Paranal setae 30 p, postanal seta 10 p. Chaetotaxy and sclero- tization of opisthogaster as in Text-fig. 47B. Peritreme extends to anterior part of coxa II ; peritrematal shield fused anteriorly with dorsal shield, free posteriorly. Legs with normal chaetotaxy. Setae av, on femur II and fd, on femur IV stout and spine-like. Ventral setae on tarsus II stout, pointed. Length/width (in ,) of leg segments : I II III IV genu 75/33 69/45 45/33 63/32 tibia 81/27 57/42 42/35 66/32 tarsus 132/24 96/32 90/28 120/30 Male: Chelicera with segment I, 54 »; II, 132 »; movable digit 54 p», uniden- tate ; spermadactyl as in Text-fig. 47C ; fixed digit bidentate, pilus dentilis slender, setiform. Four pairs of gnathosomal setae with distance between c.s. equal to that between hyp.2 (42 «). Deutosternum with six transverse rows of denticles (2-4 per row) ; corniculi 30 » long. Internal malae elongate, triangular, pilose. Tectum and pedipalps as in female. 226 Gs OWENTEVANS & “W... Me2DIEe Fic. 47. Pseudoparasitus centralis (Berlese). A. Dorsum of idiosoma of female ; B. venter of idiosoma of female; c. chelicera of male; pb. chelicera of female; §E. venter of idio- soma of male; Fr. venter of gnathosoma of female. THE BRITISH DERMANYSSIDAE (ACARI) 227 Dorsal shield (495 » X 300 ») with 39 pairs of setae distributed asin female. Trito- sternum with base 21 p, laciniae 63 ». Holoventral shield (390 » x 99 ») with four pairs of setae and three pairs of pores in the sternal region, six pairs of setae in the genito-ventral region. Between sti, 45 mu; between sft and st3, go pw. Paranal setae 24 p, postanal seta 12 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 47E. Peritreme extends nearly to anterior margin of coxa II; anterior part of peritrematal shield fused with dorsal shield, posterior part free. Chaetotaxy of legs normal. Femur I with setae ad, and pd, short, stout and spine-like ; femur II with seta av, a stout spine ; genu II with seta pv inflated basally ; femur IV with seta pd; inflated. Length/width (in ) of leg segments : I II 1G IV genu 69/27 66/45 42/30 60/36 tibia PGI 54/42 40/32 60/40 tarsus 120/22 84/30 84/24 114/30 HapitatT: Free-living in litter under deciduous trees etc. in Britain and Europe. Pseudoparasitus dentatus (Halbert) Laelaps dentatus Halbert, 1920, Proc. R. Ivish Acad. 25: 123. Female: Chelicera with segment I, 54 II, 162 ~; movable digit 70 p, bidentate ; fixed digit with four teeth ; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 48 papart, hvp.2 42-45 papart. Deutosternum with six transvere rows of denticles (5~7 per row) ; corniculi 45-48 » long. Internal malae as in P. centralis. Tectum capituli with denticulate anterior margin. Pedipalp (2—5-6-14) with three-tined apotele. Dorsal shield (564-678 » x 348-360 ») with 39 pairs of setae distributed as in Text- fig. 48A. Surface of shield reticulate ; no hypertrichy. Tritosternum with base 33 p», laciniae81 pv. Sternalshield (150-180 p & I14-120 p) with convex posterior margin reaching nearly to posterior margin of coxa III ; with three pairs of setae and two pairs of pores, the second pair small and circular ; meta- sternal setae free with associated pores. Between sti, 66-72 » ; between sfI and sf3, 108-132 ». Genital shield (192-216 p x 105 pm) flask-shaped with four pairs of setae ; between gen., 102 4; between Jv2, 84 ». Anal shield (60-68 » x 75-90 p) with maximum width (123-170 ») at anterior margin; paranal setae 30-40 p, postanal seta 18-30 ». Chaetotaxy and sclerotization of opisthogaster as in Text- fig. 488. Peritreme extends to about the anterior fourth of coxa II; anterior part of peritrematal shield fused with dorsal shield, posterior part free. Legs with normal chaetotaxy. Femur II with seta av, stout. Femur IV with seta pd, stout, spine-like. Tarsus II has setae av,, avo, pv, Pz, ply and mv all stout basally with fine tips. Length/width (in yu) of leg segments : 228 G. OWEN EVANS & W. M. TILL I I Ul IV genu 80/30 75/40 45/30 72/36 tibia 87/30 60/42 45/30 72/33 tarsus 138/27 87 /32 90/27 105/30 HABITAT: Free-living (in tidal debris, moss under Calluna, humus under deciduous trees) in Britain, Ireland and the Channel Islands. Genus OLOLAELAPS Berlese Hypoaspis Berlese, 1903, Zool. Anz. 27: 14 (non Hypoaspis Canestrini). Ololaelaps Berlese, 1904, Redia 1 : 260. Pristolaelaps Womersley, 1956, J. Linn. Soc. Lond. Zool. 42: 511. Type - Hypoaspis venetus Berlese, 1903. Medium to heavily sclerotized mites. Chelicerae chelate-dentate ; movable digit bidentate in the female, unidentate in the male ; spermadactyl groove straight or Fic. 48. Pseudoparasitus dentatus (Halbert), female ; dorsum (A) and venter (8) of idiosoma., THE BRITISH DERMANYSSIDAE (ACARI) 229 sinuous. Pilus dentilis and dorsal seta simple, fissures normal. Chaetotaxy of venter of gnathosoma and pedipalps normal; apotele three-tined, posterior tine reduced. Deutosternum with six transverse rows of denticles. Internal malae well- developed ; corniculihorn-like. Tectum capituli with rounded, minutely denticulate anterior margin. Dorsal surface of idiosoma completely covered by an entire shield bearing 37 (zt and z3 lacking), 38 (z3 lacking) or 39 pairs of fine, simple, setae. Unpaired accessory setae may be present in the region of the J series. Tritosternum normal, bipartite. Pre-endopodal shields conspicuous. Sterno- metasternal shield in the female with four pairs of setae and three pairs of pores. Genito-ventro-anal shield large, bearing four to six pairs of setae excluding the anals, and free or fused by way of the metapodal shields with the peritrematal shields. Male with holoventral shield free or fused with the peritrematal shields. Podal shields strongly developed behind coxae IV. Segmental chaetotaxy of the legs normal. Two groups of species appear to be represented within the genus, namely, the venela group in which the peritrematal shields are fused by way of the metapodal _ shields with the genito-ventral shield of the female, the spermathecal ducts are heavily sclerotized and conspicuous and the spermadactyl groove is sinuous, and the placen- tula group in which the peritrematal shields are free posteriorly in the female, there are no distinct spermathecal ducts and the spermadactyl groove is straight. There is also a tendency in the veneta group towards hypotrichy of the podonotum in the adult. Three species are represented in the British fauna and they appear to favour damp habitats. KEY TO FEMALES I Paravertical seta (z1) absent, dorsal shield with 37 pairs of setae (Text-fig. 504) ; spermathecal ducts as in Text-fig. 50E; genito-ventro-anal shield normally with four pairs of setae excluding anals (Text-fig. 50F) O. sellnicki Bregetova & Koroleva( p. 231) — Paravertical setae (z1) present, dorsal shield with 38 or 39 pairs of setae : ; Spermathecal ducts conspicuous (Text-fig. 49c) ; peritrematal shield fused with the podal and genito-ventro-anal shields (Text-fig. 49B and pb) ; with 38 pairs (23 absent) of relatively short dorsal setae (Text-fig. 49A) . O. veneta (Berlese) (p. 231) — Spermathecal duct apparently absent ; peritrematal shield free posteriorly (Text-fig. 52F) ; with 39 pairs of long, fine, dorsal setae (Text-fig. 524) O. placentula (Berlese) (p. 235) Key To MALES I Spermadactyl groove straight (Text-fig. 52B) ; genu III with seta pu modified into a rounded protuberance (Text-fig. 52c) ; dorsal shield with 39 pairs of setae O. placentula (Berlese) (p. 237) - Spermadactyl groove sinuous (Text-fig. 51B and p) ; genu III with seta pv normal ; dorsal shield with 37 or 38 pairs of setae : : : : : : é 2 2 Paravertical setae absent . F é : O. sellnicki Bregetova & Koroleva (p.233 ) Paravertical setae present. : : : ; : : : O. veneta (Berlese) 230 G. OWEN EVANS & W. Se Fic. 49. Ololaelaps veneta (Berlese), female ; podal shields in region of coxa IV (B) ; shield (p). Means dorsum of idiosoma (a) ; ““spermathecal ducts’ (c) ; genito-ventro-anal peritrematal- THE BRITISH DERMANYSSIDAE. (ACART) 231 Ololaelaps veneta (Berlese) Laelaps tumidulus (Koch) Berlese, 1889, Acari, Myr. Scorp. Ital. fasc. 44, no. 5. Hypoaspis venetus Berlese, 1903, Zool. Anz. 27: 14 (nom. nov. pro Laelaps tumidulus Berlese, 1889 nec Koch). Ololaelaps venetus Berlese, 1904, Redia 1: 260. Ololaelaps halaskovae Bregetova & Koroleva, 1964, Pavazit. Sb. 22: 81, figs. (Syn. nov.). Female: Chelicerae with segment I, 75 »; II, 150 »; movable digit 87 yw, bi- dentate; fixed digit tridentate; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 57 » apart; hyp.2 about 48 pw apart. Deuto- sternum with six transverse rows of denticles (3-6 denticles per row) ; corniculi 45 p» long ; internal malae as in O. sellnickt. Tectum capituli with anterior margin finely denticulate. Salivary styli conspicuous. Pedipalp (2-5-6-14) with three-tined apotele. Dorsal shield (672-804 X 444-456 ») with 38 pairs of relatively short, simple setae distributed as in Text-fig. 494, seta 23 being absent. An unpaired accessory seta is present between the / series. Surface of shield granular with sculptured areas in the podonotal region. Tritosternum with base 30 p, laciniae 81 p. Sterno-metasternal shield (102 p x 132 ») with four pairs of setae and three pairs of pores. Between sf1, 66 » ; between sti and st3, 87 pw. Genito-ventro-anal shield (270 » x 258 ) with six pairs of setae, excluding the anals. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 49D. Peritreme extends to posterior fourth of coxa I. Peritrematal shield fused with podal shield and with genito-ventro-anal shield (Text-fig. 49B). Spermathecal ducts as in Text-fig. 49C. Legs with normal chaetotaxy. Length/width (in ») of leg segments : I II III IV genu 81/36 7250 54/38 75/40 tibia 87/32 60/45 54/30 75/38 tarsus 153/30 102/34 105/30 135/33 Male: Described and figured by Bregetova & Koroleva (1964). Hasitat: In moss, Britain and Italy, and in nests of small mammals U.S.S.R. Ololaelaps sellnicki Bregetova & Koroleva Ololaelaps haemisphaericus (Koch), Sellnick, 1940, Géteborgs Vetensk. Samh. Handl. (5B), 6 (14) : 69, fig. 48. Ololaelaps selinicki Bregetova, 1964, Pavazit. Sb. 22: 77, figs. (nom. nov. pro Ololaelaps haemis- phaericus Sellnick, 194¢ non Koch, 1839). Female : Chelicera with segment I, 102 » ; II, 204 «1; movable digit 87 pv, biden- tate; fixed digit tridentate ; pilus dentilis short, setiform (Text-fig. 50c). Four 232 G. OWEN EVANS & W. M. TILL gnathosoma (B); chelicera (c); tectum capituli (Dp); “spermathecal ducts” | } | Fic. 50. Ololaelaps sellnicki Bregetova, female; dorsum of idiosoma (a) ; oe . venter of idiosoma (F). THE BRITISH DERMANYSSIDAE (ACARI) 233 pairs of gnathosomal setae with c.s. about 60 p» apart, hyp.2 about 48 p apart. Deutosternum with six transverse rows of denticles (I-5 per row) ; corniculi 604 long ; internal malae as in Text-fig. 508. Tectum capituli with anterior margin very finely denticulate (Text-fig. 50D). Salivary styliconspicuous. Pedipalp (2—5—6—-14) with three-tined apotele. Dorsal shield (636 » x 480 ») with 37 pairs of simple, subequal setae distributed as in Text-fig. 50A, setae zI and 23 being absent. A single accessory seta is present between the / series. Surface of shield granular with sculptured areas in the podonotal region. Tritosternum with base 24 p, laciniae 66 ». Sterno-metasternal shield (102 p x 132 ») with four pairs of setae and three pairs of pores. Between sf1, 60 » ; between sti and st3, 87 ». Genito-ventro-anal shield (264 » 264 ») with four pairs of setae in the preanal region. Paranal setae 36 pw, postanal seta 21 p. Chaetotaxy of opisthogaster as in Text-fig. 50F. Peritreme extends to middle of coxa I; anterior part of peritrematal shield fused with dorsal shield, posterior part fused with podal shield behind coxa IV and with genito-ventro-anal shield. Spermathecal ducts as in Text-fig. 50E. Legs with normal chaetotaxy. Tarsus II with ventral setae inflated basally, tapering to long, fine points. Length/width (in ») of leg segments : I 10 III by genu 90/40 75/50 63/42 84/45 tibia 102/36 72/48 60/40 80/42 tarsus 168 /32 126/40 114/36 156/36 Male: Chelicera with segment I, 60 »; 282 »; movable digit 63 pw (102 pw with spermadactyl), unidentate ; groove of spermadactyl with sinous margins (Text-figs. 51 B and D); fixed digit unidentate, pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 50 p apart, hyp.2 about 45 p apart. Deuto- sternum with six transverse rows of denticles (2-5 per row) ; corniculi 48 y long ; internal malae fringed (Text-fig. 51c). Tectum, salivary styli and pedipalps as in female. Dorsal shield (516 » x 360 yw) with 37 pairs of setae and one unpaired accessory seta. Tritosternum with base 15 p, laciniae 57 ». Holoventral shield (408 p x 93 ») expanded behind coxae IV and fused with podal shields ; with four pairs of setae and three pairs of pores in the sternal region, six pairs of setae in the genito- ventral region. Between st1, 54 »; between sti and st3, 87 ». Paranal setae 27 p, postanal seta g p. Chaetotaxy of opisthogaster as in Text-fig. 51. Peritreme extends to about middle of coxa I; anterior part of peritrematal shield fused with dorsal shield, posterior part free. Legs with normal chaetotaxy. Tarsus II with seta pv, strongly inflated basally, terminating in a fine point. Length/width (in ») of leg segments : G. OWEN EVANS & W. M. TILL 234 THE BRITISH DERMANYSSIDAE (ACARI) 235 I II IgG IV genu 75/33 66/45 48/36 66/36 tibia 81/30 60/42 48/33 69/36 tarsus 144/27 108/36 99/30 132/30 Deutonymph : Chelicera with segment I, 72 » ; Il about 160 » ; movable digit 66 yp. Four pairs of gnathosomal setae with c.s. about 60 p apart, hvp.2 about 48 p apart. Deutosternum with six transverse rows of denticles (3-6 per row) ; corniculi 40 p long. Internal malae and pedipalps as in female. Dorsal shield (624 » x 396 «) with lateral incisions, bearing 34 pairs of setae ; setae zI and 23 absent; setae s3, s6 and humeral seta on cuticle (Text-fig. 51A). Opisthonotal region with one accessory seta; five pairs of R setae on the cuticle. Tritosternum with base 20 p, laciniae 50 w. Sternal shield (246 » x 105 pw) with four pairs of setae and three pairs of pores. Between sf1, 60 » ; between sfi and sf3, 102 wp. Analshield (96 » X I10 pw) with maximum width 162 » ; paranal setae 27 p ; postanal seta 18 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 5iF. Peritreme extends to middle of coxa I; anterior part of peritrematal shield free, extending a short distance beyond tip of peritreme ; posterior part not developed. Legs with normal chaetotaxy. Tarsus II with setae av and pv stout basally, tapering to long, fine points. Length/width (in ,) of leg segments : M Il Lit IV genu 75/42 70/50 50/42 72/45 tibia 87/36 60/45 48/40 72/42 tarsus 150/33 93/40 105/33 144/40 HABITAT: Found in moss and decaying organic matter in Britain, Europe and Iceland. Note: Bregetova & Koroleva (1964) have figured the dorsum of O. sellnicki with paravertical setae. Dr. Bregetova has kindly re-examined her specimens at our request and has confirmed that the paraverticals are actually absent as in our material. Ololaelaps placentula (Berlese) Laelaps placentula Berlese, 1887, Acari, Myr. Scorp. Ital. fasc. 44, no. 3. Ololaelaps placentula: Sellnick, 1940, Géteborgs Vetensk. Samh. Handl. (58), 6 (14): 69, fig. 49; Bregetova & Koroleva, 1964, Pavazit. Sb. 22: 70, figs. Ololaelaps confinis Berlese, 1904, Redia 1: 261. Female : Chelicera with segment I, go » ; II, 192 » ; movable digit 102 p, biden- tate ; fixed digit with three teeth, two of which are very small and rounded ; pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 66 » apart, Fic. 51. Ololaelaps sellnichi Bregetova. A. Dorsum of idiosoma of deutonymph; B. chelicera of male, internal view ; c. deutosternum and hypostome of male; b. chelicera of male, dorsal view; E. venter of idiosoma of male; F. venter of idiosoma of deuto- nymph. ZOOL. 14, 5. 16 236 FIG. 52. G. OWEN EVANS & W. M. TILL Ololaelaps placentula (Berlese). A. Dorsum of idiosoma of female ; B. chelicera of male; c. genu III of male; pb. tarsus II of male; §E. venter of idiosoma of male; F. venter of idiosoma of female. THE BRITISH DERMANYSSTIDAE (ACARI) 237 hyp.2 about 57 » apart. Deutosternum with six transverse rows each of many denticles ; corniculi 63 #; internal malae and tectum captituli as in J. sellnicki. Salivary styl conspicuous, extending almost to tips of corniculi. Pedipalp (2-56-14) with three-tined apotele. Dorsal shield (approximately 732 » 540 «) with 39 pairs of simple setae distri- buted as in Text-fig. 524; Surface of shield granular, with faint reticulations ; no hypertrichy. Tritosternum with base 27 p, laciniae 84 wp. Sternal shield (80 » x 132 p») with four pairs of setae and three pairs of pores. Between s/1, 80 » ; between sf1 and sé3, 84 p. Genito-ventro-anal shield (288 » x 252 «) expanded behind coxae IV, but not fused with podal or peritrematal shields ; with six pairs of setae in the preanal region. Paranal setae 36 p», postanal seta 30 uw. Chaetotaxy of opisthogaster as in Text-fig. 52F. Peritreme extends to about middle of coxa I; anterior part of peri- trematal shield fused with dorsal shield, posterior part free. Legs with normal chaetotaxy; tarsus II with ventral setae relatively stout basally, tapering to long, fine points. Length/width (in p) of leg segments : I I III IV genu 93/42 75/54 63/45 78/45 tibia 96/36 66/50 54/42 75/45 tarsus 165/33 126/42 120/38 160/38 Male: Chelicera with segment I, 75 »; II, 165 »; movable digit 75 pw (123 uw with spermadactyl), unidentate ; margins of spermadactyl groove smooth (Text-fig. 52B) ; fixed digit unidentate, pilus dentilis short, setiform. Four pairsofgnathosomal setae with c.s. about 60 p apart, hyp.2 about 50 » apart. Deutosternum with seven transverse rows each of many denticles ; corniculi 54 » long ; hypostomal processes consist of a pair of slender, pilose triangles. Tectum capituli, salivary styli and pedipalps as in female. Dorsal shield (570 » x 460 p) with 39 pairs of setae as in female. Tritosternum with base 21 p, laciniae 63 ». Holoventral shield (420 » x 123 ») expanded behind coxae IV, fused with podal but not with peritrematal shields ; with four pairs of setae and three pairs of pores in the sternal region, six pairs of setae in the genito-ventral region. Between sti, 70 » ; between st1 and st3, 93 ». Paranal setae 33 », postanal seta 24 p. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 52E. Peritreme extends beyond posterior margin of coxa I; anterior part of peritrematal shield fused with dorsal shield, posterior part free. Legs with normal chaetotaxy. Genu III with seta pv reduced to a small “ pimple situated on a protuberance of the segment (Text-fig. 52c). Tarsus II with seta pu, inflated basally, tapering to a long, fine point (Text-fig. 52D). Length/width (in p) of leg segments : ” I II III IV genu 75/30 78/50 60/42 72/42 tibia 84/33 66/45 57/40 75/40 tarsus T5OVSOr | 132/35 1 LIZ iB4) A438 238 G. OWEN EVANS *& W. M.. TILE Immature stages: The larva, protonymph and deutonymph of this species are — described and figured by Bregetova & Koroleva (1964). Hasitat: Found in moss and decaying organic matter in Britain, Europe and Iceland. Recorded from the nests of small mammals in the U.S.S.R. Subfamily HAEMOGAMASINAE Oudemans Haemogamasidae Oudemans, 1926, Ent. Ber. Amst. 7 (150) : 120. Adults : Chelicerae chelate, dentate or edentate ; in dentate forms movable digit in the female bidentate, in the male unidentate. In some Brevisterna fixed digit in both sexes weakly sclerotized and transversely striated, movable digit of female with conspicuous hyaline lobe. Pilus dentilis simple or inflated ; dorsal seta conspicuous, rarely reduced to a microseta or absent ; fissures normal. Arthrodial processes present or absent, arthrodial membrane sometimes lobate (Haemogamasus). Sperma- dactyl grooved, relatively short, free distally but closely associated with the movable digit. Chaetotaxy of venter of gnathosoma normal. Pedipalps with five free seg- ments, apotele two- or three-tined ; chaetotaxy normal (2—5-6-14). Palptrochanter rarely with weak medio-ventral ridge ; with sensory organ in Eulaelaps. Deuto- sternum with ten or more transverse rows of denticles ; denticles rarely in single file (Brevisterna). Corniculi horn-like, in the form of flattened lobes or membranous. Internal malae usually bipartite and conspicuous. Tectum capituli fimbriated and overhanging the hypostome, or weakly sclerotized with smooth margins (brevisierna). Dorsum of idiosoma in both sexes with entire shield or with abbreviated shield and a small setae-bearing pygidial shield in the female (Brevisterna). Primary chaetotaxy completely obliterated by secondary hypertrichy of both the dorsal shield and the unsclerotized cuticle. Dorsal setae simple or pilose. Tritosternum normal, laciniae pilose; lateral margins of basal region rarely denticulate (Eulaelaps). Sternal shield in the female normally with three pairs of setae and two pairs of pores, but with accessorysetae in some species of H aemogamasus; shield reduced and bearing only two pairs of setae and two pairs of pores in brevi- sterna. Metasternal setae free. Genito-ventral shield usually with numerous setae. Anal shield normal or with accessory setae, opisthogastric cuticle showing hyper- trichy. Metapodal shields large, subtriangular or subovate (Ezlaelaps) or relatively small. Peritrematal shields free or fused with the podal shields posterior to coxae IV. Stigmata normal, peritremes well developed or reduced. Podal shields weakly developed. Male with holoventral shield, with or without accessory setae in the sternito-genital region, opisthogastric region densely setose; or with sternito- genito-ventral and anal shields (Ischyropoda). Genital orifice presternal. Segmental chaetotaxy of legs normal except for genu [V (2—¥#, 3—2) in all species and rarely (H. irsutosimilis) femur II (2—?—1) and femur III (1—3—1). Anterior spine on coxa II small or absent. Ambulacra with claws sometimes reduced on leg I. Femur II with one or more ventral setae hypertrophied in some males. Larva and Protonymph : Insufficient data are available on these stages for their definition. Both stages have been described for Haemogamasus ambulans : keegan, THE BRITISH DERMANYSSIDAE (ACART) 239 1951, by Furman (1959). Hypertrichy of the idiosoma is apparent at the protonym- phal stage in Haemogamasus. Deutonymph : Gnathosoma with the general characteristcs of the female. Dorsal idiosomal sclerotization comprising a single shield with (Eulaelaps and Haemogamasus) or without (Ischyropoda) lateral incisions, or an abbreviated dorsal shield (podonotal) with mesonotal scutellae and fragments of the pygidial shield (Brevisterna). Dorsal chaetotaxy shows marked hypertrichy. Tritosternum as in female. Sternito-genital shield with three pairs of sternal and one pair of metasternal setae, genital setae free, or with numerous accessory setae. Anal shield normal or with accessory setae ; opisthogastric cuticle showing hyper- trichy. Podal shields usually small. Peritrematal shields free posteriorly ; stig- mata and peritremes as in adult. Leg chaetotaxy as in corresponding adult. KEY TO THE GENERA OF THE BRITISH HAEMOGAMASINAE 1 Female with metapodal shields large, subtriangular, and flanking a broad genito- ventral shield ; palptrochanter with sensory organ (Text-fig. 62F) ; anterior rows of deutosternal denticles with 5 to 7 denticles per row in both sexes EULAELAPS (p. 259) — Female with metapodal shields small, sub-ovate ; genito-ventral shield relatively narrow ; palptrochanter lacking sensory organ ; anterior rows of deutosternal denticles with two to three denticles per row in both sexes HAEMOGAMASUS (p. 239) Genus HAEMOGAMASUS Berlese Liponissus Kolenati, 1858, Wien. ent. Monatschr. 2: 5. Liponyssus Kolenati, 1859, S.B. Akad. Wiss. Wien. 35: 172. Haemogamasus Berlese, 1889, Acay., Mvyr. Scorp. Ital. fasc. 52, nos. 2 and 10. Euhaemogamasus Ewing, 1933, Proc. U.S. nat. Mus. 82 (art. 30) : 3. Groschaftella Samsinak, 1957, Cslka. parasitol. 4: 270 (syn. nov.). Type: Haemogamasus hirsutus Berlese, 1889. Chelicerae chelate-dentate, rarely edentate ; pilus dentilis simple or inflated ; dorsal seta long, stout, rarely absent. Dorsal and anti-axial fissures conspicuous. Spermadactyl short, closely associated with the movable digit. Tectum fimbriated and overhanging the hypostome. Basis capituli with a pair of capitular setae ; hypostome with the normal three pairs of setae. Deutosternum with 10-18 trans- verse rows of denticles. Internal malae bipartite, conspicuous. Corniculi horn-like or in the form of flattened lobes. Salivary styli well developed. Pedipalps with normal chaetotaxy ; apotele two- or three-tined. Dorsum of idiosoma with marked hypertrichy which completely obliterates the primary chaetotaxy. Dorsal shield entire in both sexes; dorsal setae simple or pilose. Tritosternum normal, pilose laciniae about three times the length of the base. Sternal shield in the female with the normal three pairs of setae and two or three pairs of pores ; with accessory setae in some species. Metasternal setae free. Genito- G. OWEN EVANS & W. M. TILL 240 ventral shield flask-shaped with numerous setae. Anal shield pear-shaped, with the usual three setae associated with the anus, and with or without accessory setae. Opisthogastric integument densely setose. Metapodal shields small, peritrematal shield fused with expodal shield in the region of coxa IV. Male with holoventral shield, with or without accessory setae in the sternito-genital region ; opisthogastric region of shield densely setose. Segmental chaetotaxy of legs normal except for genu IV (2—?, 3—2) and tibia IV (2—?, 7—2) in all species and femur II (2—?—1) and femur III (1—3—1) in ZH. hirsutosimilts. The type of the genus Liponyssus Kolenati (Liponyssus setosus Kolenati, 1858) was so poorly described and figured that its identity has always been in doubt. In earlier standard works on the obligatory ectoparasitic “‘ Laelaptidae”’ the genus was widely used to accommodate certain species now included in the Macronyssinae and Ewing (1923) proposed the subfamily Liponyssinae for Liponyssus sensu authors and certain related genera. Oudemans (1936) considered L. setosus to be “a totally different sort of mite ”’ from the other species included in Lifonyssus by authors, the majority of which were placed by him in Macronyssus for which he proposed the family Macronyssidae. Although certain authors have continued to use Liponyssus and Liponyssinae since Oudemans (1936) the general tendency has been for the suppres- sion of Liponyssus sensu authors, especially since Fonseca’s revisionary work in 1948. A single damaged specimen, probably the type, of Liponyssus setosus Kolenati has recently been found in the Kolenati collection in Paris. This species is a male of Haemogamasus horridus Michael. The strict application of the Rules of Priority would necessitate the synonymizing of Haemogamasus Berlese with Liponyssus Kolenati. However this course would be contrary to the interests of nomenclatural stability especially since Haemogamasus and suprageneric taxa based upon it are so well established in the literature. We propose to submit an application to the International Commission on Zoological Nomenclature requesting its use of the plenary powers to suppress the genus Liponyssus Kolenati, 1858, its type species and the suprageneric categories based on the name Liponyssus. KEY TO FEMALES nN £ Sternal shield with the normal three pairs of setae only Sternal shield with accessory setae which are usually shorter seta ihe aralanease sternal setae and vary in size, number and position on the shield . Posterior margin of the sternal shield deeply incised to a level midway bees een the first and second pairs of sternal setae (Text-fig. 60B). Anal shield usually with only the normal three setae. Dorsal and ventral surface of the idiosoma less densely setose than in the following species (Text-figs. 60A—B) . H. pontiger (Berlese) (p. 257) Posterior margin of sternal shield at the most slightly concave (Text-fig. 50B). Anal shield with several accessory setae : : 3 : s 3. First pair of sternal setae and hypostomal setae barbed. Dorsal shield, especially in the anterior and marginal regions, with many barbed setae. Tectum capituli nar- row, pointed, with simple and multiple fimbriae. Pilus dentilis inflated (Text-fig. 590E). . H. nidi Michael (p. 254) wn N 3 THE BRITISH DERMANYSSIDAE (ACARI) 241 Sternal, hypostomal and dorsal setae smooth. Tectum capituli with gently rounded margin of simple and multiple fimbriae. VPilus dentilis short, setiform (Text-figs. 57A andc) . : 4 Postero-ventral margin of the idiosoma ah at as one pair of setae Gonspicueusly, longer than the rest (Text-fig. 588) ; sternal shield about as wide as long ; genito- ventral shield with 60 or more setae ; deutosternal denticles as in Text-fig. 58b. Associated with Talpa (Insectivora) é : H. horridus Michael (p. 252) Postero-ventral margin of the idiosoma with cece subequal (Text-fig. 56B) ; sternal shield about 14 times wider than long ; genito-ventral shield with about 40-45 setae ; deutosternal denticles as in Text-fig. 56Fr. Associated with Muridae (Rodentia) . 4 : 3 : H. arvicolarum (Berlese) ( p. 250) Femur II with 12 setae (2—}—1), femur II with seven setae (I—3—1). One pair of extremely long postero-ventral idiosomal setae about three times the length of the neighbouring setae (Text-fig. 55c) : H, hirsutosimilis Willmann (p. 246) Femur II with 11 setae (2—3—1), femur IIT with six setae (I—3—1). Without very long postero-ventral idiosomal setae ; longest pair at the most twice the length of the neighbouring setae : 6 Chelae dentate ; palptrochanter with hot setae pilose, Por inflated. Genito: Sore shield with accessory setae restricted mainly to the posterior region. Many setae barbed(st1, hyp. 2-3, c.s., many dorsal setae and most setae on the appendages). Dorsal setae uniform in size except at anterior and posterior margins. Anal shield wider than long . 5 : H. ambulans (Thorell) (p. 244) Chelae edentate ; palptrochanter ita inner seta inflated basally. Genito-ventral shield with accessory setae evenly distributed over entire surface. All setae smooth. Dorsal shield with large and small setae interspersed over its entire surface. Anal shield longer than wide . : 6 : é . H. hirsutus Berlese (p. 242) KEry To MALES Sternal region of holoventral shield with no accessory setae anterior to the level of the second pair of pores Holoventral shield with Aceon eetae distributed over ene Bue antbies First pair of sternal setae smooth (Text-fig. 58c). Tectum capituli with ena rounded margin of simple and multiple fimbriae. Chelicera with dorsal seta dis- N tinctly distal to base of fixed digit : : c : : : : : 3 First pair of sternal setae barbed (Text-fig. 59c). Tectum captuli tapering, pointed, with fimbriate margin. Chelicera with dorsal seta proximal to base of fixed digit 4 Femur II with av, short, spinose, conspicuously different from other ventral setae (Text-fig. 56£). Postero-ventral idiosomal setae subequal (Text-fig. 56D). Sperm- dactyl without retrose spur (Text-fig. 57D) . H. arvicolarum Berlese (p. 251) Femur II with av, setiform, similar to other ventral setae. At least one pair of postero-ventral idiosomal setae conspicuously longer than the neighbouring setae (Text-fig. 58c). Spermadactyl with a stout retrose spur (Text-fig. 57B). H. horridus Michael (p. 254) Ventral seta on femur III setiform. Holoventral shield with dense covering of setae (Text-fig. 59c). Peritrematal shields fused posteriorly with holoventral shield and with podal elements of coxa IV. Pilus dentilis inflated (Text-fig. 59F) H. nidi (Michael (p. 255) Ventral seta on femur III short, spinose. Holoventral shield with relatively sparse covering of setae (Text-fig. 60D). Peritrematal shields free posteriorly. Pilus dentilis slender, setiform (Text -fig. 60E) . 5 H. pontiger (Berlese) (p. 259) First pair of sternal setae barbed : : : : : . H.ambulans (Thorell) Allsternal setae smooth . 7 : : : ; : : , : ; 6 G. OWEN EVANS: & W. Mo TEL bd aN Nd 6 Femur II with 12 setae (4 ventrals, of which two are blunt and peg-like). A pair of very long setae near posterior end of idiosoma about three times the length of the neighbouring setae. Setae on dorsal shield subequal in size, except at anterior and posterior margins. Fixed digit of chelicera relatively short and broad, with a short, antero-median projection (Text-fig. 555). H. hirsutosimilis Willmann (p. 248) Femur II with 11 setae (three ventrals, all stout, pointed, spine-like). No very long setae near posterior end of body, longest seta at the most 14 times the length of the neighbouring setae. Dorsal shield with large and small setae interspersed over its entire surface. Fixed digit of chelicera relatively long, slender, hook-like, with a prominent median expansion (Text-fig. 53E) H. hirsutus Berlese (p. 244) Haemogamasus hirsutus Berlese Haemogamasus hirsutus Berlese, 1889, Acari, Myr. Scorp. Ital. fasc. 52, no. 2. Keegan, 1951, Proc. U.S. nat. Mus. 101: 210, fig. 42. Bregetova, 1956, Opred. Faune SSSR. 61: 140, figs. 290-293, 322-325. Female: Chelicera with segment I, 144 » ; I, 300; movable digit 87 »; both digits edentate ; pilus dentilis short, setiform (Text-fig. 53D), dorsal seta apparently absent. Four pairs of gnathosomal setae with c.s. about 120 w apart ; hyp.2 about 84 » apart. Deutosternum with 14 transverse rows of denticles ; corniculi mem- branous, about 63 » long ; internal malae with inner processes slender, pilose, outer processes fimbriate (Evans & Till, 1965, Text-fig. 4c). Tectum capituli narrow, with anterior margin fimbriate. Salivary styli conspicuous, tubular, curved. Pedipalp (2-5—6-14) with three-tined apotele, one being very small; palptrochanter with inner seta inflated basally. Dorsal shield (1,164 » * 636 ») densely covered with simple setae of variable length (Text-fig. 534). Surface of shield granular with some weakly defined reticula- tions. Tritosternum with base 60 p, laciniae 228 w; presternal area reticulate, with minute spines. Sternal shield (174 uw * 234 «) with the usual three pairs of sternal setae and about 25 accessories ; with three pairs of pores, the pair usually associated with the metasternal setae being on the posterior margin of the shield ; metasternal setae free. Between sf1, 150 »; between sti and st3, 180 w. Genito-ventral shield relatively slender with numerous accessory setae distributed over entire shield. Anal shield (168 » x 96 ) with the usual three setae and five accessories ; paranal setae 123 p, postanal seta 108 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 53B. Peritreme extends to posterior margin of coxa II; peritrematal shield free anteriorly, fused posteriorly with podal shield of coxa IV. Chaetotaxy of legs differs from normal pattern in that genu IV has two postero- lateral setae and tibia IV has the dorsal setal pattern (2-2) instead of (1-3). All the A. Dorsum of idiosoma of female; B. venter of Fic. 53. Haemogamasus hirsutus Berlese. D. chelicera of female; E. chelicera idiosoma of female ; c. venter of idiosoma of male ; of male. 243 THE BRITISH DERMANYSSIDAE (ACARI) 244 G. OWEN VEWANS! & W. OM. «DEBE leg setae are smooth. Distal margins of most leg segments denticulate ventrally. Length/width (in ») of leg segments : I II Ill IV genu 222/60 156/100 =: 150/78 204/72 tibia 228/54 156/90 162/66 222/60 tarsus 330/48 264/66 342/54 498/54 Male: Chelicera with segment I, 114 »; II, 222 »; movable digit 78 pu (87 » with spermadactyl) ; both digits edentate (Text-fig. 53£); pilus dentilis short, setiform. Four pairs of gnathosomal setae with c.s. about 99 » apart ; hyp.2 about 72 wapart. Deutosternum with 13 transverse rows of denticles ; corniculi 54 p» long. Tectum capituli with anterior and lateral margins fimbriate. Internal malae, sali- vary styli and pedipalps as in female. Dorsal shield (1,026 » X 520 mw) as in female. Tritosternum with base 50 u, laciniae 165 ». Holoventral shield (840 » x 198 yw) with the usual sternal, meta- sternal and anal setae recognizable ; accessory setae present over entire surface of shield (Text-fig. 53c). Par- and postanal setae subequal, about 96 » long. Peri- tremes and peritrematal shields as in female. Chaetotaxy of legs as in female ; relatively stouter setae on femur II (ventrals), genu and tibia II (av) and tarsus II (mv and av). Length/width (in pz) of leg seg- ments: I Il Ill IV genu 198/54 132/84 114/66 180/60 tibia 204/48 138/78 144/60 204/60 tarsus 264/42 186/54 264/48 414/48 Hasitat: Found in Britain in the nest and on the body of Talpa europaea Linn. and on the field vole Microtus agrestis (Linn.). Also recorded from a variety of rodents in the U.S.S.R. and Europe. Haemogamasus ambulans (Thorell) Dermanyssus ambulans Thorell, 1872, Ofv. Vet.-Akad. Fovh. 2: 164. Hypoaspis ambulans: Tragardh, 1904, Fawna Arctica 4: 33, figs. Haemogamasus ambulans: Bregetova, 1953, Parazit. Sb. 15: 316; and 1956, Opred. Faune SSSR. 61: 147, figs. Haemogamasus alaskensis Ewing, 1925, Proc. biol. Soc. Wash. 38 : 13 Keegan, 1951, Proc. U.S. nat. Mus. 101 : 213, fig. 43. Haemogamasus avisugus Vitzthum, 1930, Zool. Jahrb. 60 : 397. Haemogamasus sternalis Ewing, 1933, Proc. U.S. nat. Mus. 82: 3, pl. Haemogamasus pavlovskyi Bregetova, 1949, Parazit. Sb. 11: 179, figs. Female; Chelicera with segment I, 105 »; II, 160 «; movable digit 54 x, bidentate ; fixed digit tridentate ; pilus dentilis setiform ; dorsal seta long and THE BRITISH DERMANYSSIDAE (ACART) venter of Fig. 54. Haemogamasus ambulans (Thorell), female ; dorsum of idiosoma (A) ; chelicera (D). (c) ; venter of gnathosoma idiosoma (B) ; 246 Gs OWEN UE VANS Sd Wi Mi Mieieede stout (Text-fig. 54D). Four pairs of gnathosomal setae with c.s. about 105 p apart, hyp.2 about 75 » apart; hyp.2-3 and c.s. barbed. Deutosternum with ten trans- ° verse rows of denticles (2-5 per row) ; corniculi 40 » long ; internal malae as in Text- fig. 54c. Tectum capituli narrow, pointed, fimbriate (Evans & Till, 1965 ; Text-fig. 3c). Salivary styli conspicuous, extending beyond tips of corniculi. Pedipalp (2-5-6-14) with two-tined apotele ; both trochantal setae barbed. Dorsal shield (1,008 » * 612 ») densely covered with setae mostly of uniform length (Text-fig. 544) ; some anterior and marginal setae barbed. Surface of shield granu- lar, reticulations most distinct in opisthonotal area. Tritosternum with base 57 pw, laciniae 150 ». Sternal shield (132 » X 190 p») with the usual three pairs of sternal setae and about 14 accessories, and with three pairs of pores ; metasternal setae free. Between st1, 150 »; between sti and sf3, 120 zg; stt barbed. Genito-ventral shield (276 » x 120 p) with a pair of genital setae and about 36 accessories distributed chiefly on the posterior part of the shield. Anal shield (123 X 130 m) with the usual three anal setae and four or five accessories ; paranal setae 68 »; postanal seta 78 ». Chaetotaxy and sclerotization of opis- thogaster as in Text-fig. 548. Peritreme extends to anterior third of coxa II; peritrematal shield free anteriorly, fused posteriorly with podal shields of coxa IV. Chaetotaxy of legs differs from normal pattern in that genu IV bears two postero- lateral setae and tibia IV bears two antero- and two postero-dorsal setae. All leg setae barbed except terminal setae on tarsi (ad,, pdy, av,, pv,, aly, ply, pv, on tarsi II-IV, as well as av, on tarsi III-IV, and mv on tarsus IV). Distal margins of all segments except tarsi serrated. Length/width (in p) of leg segments : I II Ill IV genu 150/54 120/60 114/54 168/50 tibia 168/48 120/57 120/48 180/48 tarsus 228/42 210/45 264/42 372/42 Hasitat: Found in the nest of Reparia riparia (Linn.) in Gloucestershire, Britain. Recorded in the literature from the nests of birds, from insectivores, bats, sables, and a variety of rodents in Germany, the U.S.S.R., Greenland, Spitsbergen and North America. Haemogamasus hirsutosimilis Willmann Haemogamasus hirsutosimilis Willmann, 1952, Z. Pavasitenk. 15 : 403, figs. Bregetova, 1956, Opved. Faune SSSR. 61: 141, figs. Female: Chelicera with segment I, 84 »; II, 156 » ; movable digit 60 », biden- tate ; fixed digit with two large and two very small teeth ; pilus dentilis short, seti- form, dorsal seta long (Text-fig. 558), Four pairs of simple gnathosomal setae with Fic. 55. Haemogamasus hirvsutosimilis Willmann. A. Dorsum of idiosoma of female ; B. chelicera of female ; c. venter of idiosoma of female ; Dp. venter of idiosoma of male ; E. chelicera of male; F. deutosternum and hypostome of female. 247 THE BRITISH DERMANYSSIDAE (ACART) geek 248 G. OWEN EVANS & W. M. TILL c.s. about 96 pw apart, Ayp.2 about 70 w apart. Deutosternum with 13 transverse rows of denticles (2-6 per row) as in Text-fig. 55F. Corniculi 40 » long ; internal malae with long, pointed inner and short, broad, outer lobes, all pilose. Tectum capituli slender, pointed, deeply denticulate. Salivary styli conspicuous. Pedi- palp (2-5—6-14) with two-tined apotele. Dorsal shield (1,032 * 564 ») with a dense covering of setae mostly of subequal length, those on the posterior margin being considerably longer ; vertical setae long and stout, not barbed (Text-fig. 554). Surface of shield granular. Tritosternum with base barbed, 50 » long ; laciniae 150 pw. Sternal shield (186 u 216 pv) with the usual sternal setae, numerous accessories, and three pairs of pores ; metasternal setae free. Between sf1, 60 »; between sti and s#3, 174 ». Genito- ventral shield with numerous setae distributed over its entire surface. Anal shield (126 » X 9O p) with the usual three anal setae and five accessories ; paranal setae I10 »; postanal seta 117 pw. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 55C; a pair of extremely long setae at the posterior end of the body. Peritreme extends nearly to posterior margin of coxa I; peritrematal shield free anteriorly, fused posteriorly with podal shield of coxa IV ; pore on posterior shield situated at tip of small tubercle. Chaetotaxy of legs differs from the normal pattern in that femur II bears 12 setae (4 ventrals), femur III bears 7 setae (2 ventrals), genu IV bears two postero-lateral setae and tibia IV has two antero- and two postero-dorsal setae ; all leg setae simple. Coxae all with rows of denticles on ventral surface ; distal margins of all segments except tarsi serrated. Ambulacrum IV nearly one-and-a-half times as long as the others. Length/width (in p) of leg segments : il LE Ill IV genu 174/54 120/72 108/66 156/60 tibia 180/45 120/63 120/60 168/54 tarsus 240/42 192/50 237/48 318/42 Male; Chelicera with segment I, 60 »; II, 150 w; fixed digit short and broad with an anterior projection; movable digit and spermadactyl strongly curved (Text-fig. 55E). Four pairs of gnathosomal setae with c.s. about 85 u apart, /ivp.2 about 63 w apart. Corniculi 42 » long ; other gnathosomal features as in female. Dorsal shield (780 » x 438 ») asin female. Tritosternum with base 33 p, laciniae 105 #3; presternal area as in female. Holoventral shield 648 , long, with a dense covering of accessory setae (Text-fig. 55D) ; true sternal setae difficult to distinguish. Par- and postanal setae subequal, about 75 » long. Peritreme extends to anterior margin of coxa II ; anterior tip of peritrematal shield terminates very close to dorsal Fic. 56. Haemogamasus arvicolarum Berlese. A. Dorsum of idiosoma of female; B. venter of idiosoma of female; c. corniculus of female, lateral view ; D. venter of idio- soma of male; §£. femur II of male; F. deutosternum of female. 249 THE BRITISH DERMANYSSIDAE (ACARI) 250 GG. -OWEN EV ANS & OW. Mo DIL shield but appears to end freely ; posterior part of shield fused with podal elements of coxa IV. Chaetotaxy and ornamentation of legs as in female. Stout, blunt, peg-like setae are present on femur II (fv.-3), genu II (av), tibia II (av) and tarsus II (mv); seta av, on tarsus IT is very stout but pointed. Seta fd, on tarsus IV is extremely long, about two-thirds the length of the tarsus. Ambulacrum IV relatively shorter than in female. Length/width (in ») of leg segments : I Il MT IV genu 160/48 102/66 96/50 138/54 tibia 156/42 105/57 108/50 150/50 tarsus 207/40 156/42 204/40 270/40 Hapitat: Found in a mole’s nest in Gloucestershire, Britain. Recorded from moles, voles and field mice in Europe and the U.S.S.R. Haemogamasus arvicolarum Berlese Haemogamasus hoyvvidus var. avvicolarum Berlese, 1920, Redia 14: 166. Haemogamasus avvicolarum: Turk, 1945, Ann. Mag. nat. Hist. (2) 12: 785, figs. Female: Chelicera with segment I, 120 »; II, 270 »; movable digit 80 p, biden- tate; fixed digit bidentate with terminal hook; pilus dentilis short, setiform ; dorsal seta long, stout, blunt (Text-fig.57c). Four pairs of simple gnathosomal setae with c.s. about 123 p» apart, hyp.2 about 93 » apart. Deutosternum with 17 transverse rows of denticles (I-4 per row) ; corniculi broad, weakly sclerotized (Text-fig. 56c and F); internal malae as in H. horridus. Tectum capituli with a rounded an- terior margin provided with numerous simple or multiple fimbriae. Salivary styli conspicuous, reaching nearly to tips of corniculi. Pedipalp (2-5—6—14) with two- tined apotele ; inner trochantal seta barbed. Dorsal shield (1,320 » x 840 ) with a dense covering of subequal setae (Text-fig. 56a). Setae 71 considerably stouter than the others but not barbed ; a few marginal setae have a minute barb near the tip. Surface of shield granular with sculpturing anteriorly and reticulations posteriorly. Tritosternum with base 63 4, laciniae 258 « ; a small spine on each side of the base. Presternal area with small spines on the reticulations. Sternal shield (156 «# X 240 ) with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between st1, 90 «; between sfi and sf3, 168 «. Genital shield with about 44 setae distributed mainly on the posterior part. Anal shield (258 « X 138 ») with maximum width 186 », bearing the usual three anal setae and about 13 accessories ; paranal setae 114 .; postanal seta 126 ». Chaetotaxy and sclerotiza- tion of opisthogaster as in Text-fig. 56B ; some marginal setae barbed. Peritreme extends to posterior margin of coxa II; peritrematal shield fused anteriorly with dorsal shield and posteriorly with podal shield of coxa IV. ——eEeEeoEESEE = THE BRITISH DERMANYSSIDAE (ACARI) 251 Chaetotaxy of legs differs from normal pattern in that genu IV has two postero- lateral setae and tibia IV has two antero- and two postero-dorsal setae. All leg setae simple. Coxae bear rows of small denticles ; distal margins of all segments except tarsi serrated ventrally. Length/width (in ,) of leg segments : I II Ill IV genu 207 /72 168/102 150/84 192/78 tibia 222/66 168 /go 168/72 240 /'72 tarsus 306/60 240/66 330/66 450/66 Male: Chelicera with segment I, 84 »; II, 200 »; movable digit with sperma- dactyl, 150 »; pilus dentilis short, setiform; dorsal seta as in female, but situated a considerable distance distal to the base of the fixed digit (Text-fig. 57D). Four pairs of simple gnathosomal setae with c.s. about 105 w apart ; hyp.2 about 93 p apart. Deutosternum with 14 transverse rows of denticles (I-3 per row) ; corniculi 57 long ; internal malae, tectum and pedipalps as in female. Dorsal shield (1,092 » X 600 pv) as in female. Tritosternum with spine on each side of base ; with base 60 p, laciniae 210 p». Presternal area with small spines on the reticulations. Holoventral shield (876 » x 192 ») with the usual sternal and metasternal setae and three pairs of pores in the sternal region, and the usual three anal setae. Numerous accessory setae are distributed on the genito-ventral region Fic. 57. A-B. Haemogamasus horvidus Michael; chelicera of female (A) and male (Bs). C-D. Haemogamasus avvicolarum Berlese ; chelicera of female (c) and male (D). ZOOL. 14, 5. 17, 252 G. OWEN EVANS & W. M. TILL and may extend to the level of the metasternals (Text-fig. 56D). Between sfz, 87 »; between sf and st3, 123 w. Paranal setae 90 » ; postanal seta 80 p. Peri- treme extends approximately to posterior border of coxa II; peritrematal shield fused anteriorly with dorsal shield, posteriorly with podal shield of coxa IV. Chaetotaxy of legs as in female. Femur II with seta av, short and stout (Text-fig. 56£). Length/width (in p) of leg segments : I Il Ill IV genu 186/66 150/87 144/72 180/66 tibia 198/54 144/78 156/60 204/66 tarsus 270/54 192/60 264/54 384/54 HaApitat: Found on moles, voles and field mice and in their nests in Britain and Europe. Haemogamasus horridus Michael Haemogamasus horridus Michael, 1892. Trans. Linn. Soc. Lond. 5 : 312, figs. Bregetova, 1956, Opred. Faune SSSR. 61 : 131, figs. Female; Chelicera with segment I, 96 » ; II, 222 »; movable digit 75 yu, biden- tate ; fixed digit bidentate, with terminal hook; pilus dentilis short, setiform ; dorsal seta long and stout (Text-fig. 57A). Four pairs of simple gnathosomal setae with c.s. about 126 p» apart; hyp.2 about 100 pw apart. Deutosternum with 15 transverse rows of denticles (1-5 denticles per row) ; corniculi broad, membranous, about 45 » long; internal malae fringed, comprising slender inner and broad, triangular, outer processes (Text-fig. 58D). Tectum capituli has a rounded anterior margin provided with numerous simple or multiple fimbriae. Salivary styli con- spicuous. Pedipalp (2—5—6-14) with two-tined apotele; inner trochantal seta barbed. Dorsal shield (1,536 » xX 864 ») with a dense covering of setae for the most part subequal in length, but tending to become longer towards the posterior margin ; vertical setae long and stout (Text-fig. 58a). Surface of shield granular and reticulate. Tritosternum with base 60 p, laciniae 276 » ; a small spine present on each side of — the base. Presternal area with small spines on the reticulations. Sternal shield (234 » X 240 p) with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between sf1, 80 » ; between sf1 and sf3, 222 ». Genito- ventral shield with about 70 setae. Anal shield (234 » X 138 w#; maximum width | 180 ») with the usual three anal setae and about 14 accessories. Paranal setae 132 4, postanal seta 150 wp long. Chaetotaxy and sclerotization of opisthogaster as in Text- fig. 588 ; at least one pair of setae at posterior margin considerably longer than the Fic. 58. Haemogamasus horvvidus Michael. A. Dorsum of idiosoma of female; 8B. venter of idiosoma of female; c. venter of idiosoma of male; Db. venter of gnathosoma of female. 253 THE BRITISH DERMANYSSIDAE (ACARI) 254 G. OWEN EVANS & W. M. TILL neighbouring setae; some setae with fine, sparse barbs. Peritreme extends to anterior margin of coxa III ; anterior tip of peritrematal shield fused with dorsal shield at level of 71 ; posterior part fused with pedal shields behind coxae IV. Chaetotaxy of legs normal except genu IV (2—7, 2—2) and tibia IV (2—?, 22). Leg setae apparently simple ; distal margins of most leg segments serrated ventrally. Length/width (in p) of leg segments : I II II IV | genu 252/75 192/108 —_168/go 222/90 tibia 276/66 192/98 186/90 264/90 tarsus 348/54 288/72, 372/72, == 504/70 Male; Chelicera with segment I, 102 »; II, 168 »; spermadactyl with a stout retrose spur (Text-fig. 57B). Four pairs of simple gnathosomal setae with c.s. about | 102 pw apart, hyp.2 about 84 » apart. Deutosternum with 15 transverse rows of | denticles (2-3 per row). Corniculi 54 » long; other gnathosomal features as in | female. Dorsal shield (1,212 » X 600 p) as in female. Tritosternum with base 48 yp, | laciniae 210 ». Holoventral shield (984 » x 204 ») with the usual four pairs of | sternal and metasternal setae, and also bearing numerous accessory setae which | extend anteriorly to the level of st4. Between str, 80 » ; between sfI and sf3, 168 x. | Paranal setae 110 p, postanal seta 135 » long. Chaetotaxy and sclerotization of | opisthogaster as in Text-fig. 58c. Peritreme extends to anterior margin of coxa IIT; | peritrematal shield fused at its anterior tip with the dorsal shield, and posteriorly © with the holoventral shield. Chaetotaxy of legs as in female. Length/width (in ») of leg segments : I II one IV | genu 222/66 156/90 150/75 192/78 | tibia 240/57 162/84 168/75 234/78 tarsus 300/54 258/63 330/63 450/60 Hapitat: Associated with moles (Talpa europaea Linn.) in Britain, Europe and | | the UiS-S Ake Haemogamasus nidi Michael Haemogamasus nidi Michael, 1892, Tvans. Linn. Soc. Lond. 5 : 314, figs. Bregetova, 1956, Opred. Faune SSSR. 61 : 134, figs. Haemogamasus michaeli Oudemans, 1903, Tijdschr. ned. dierk. Ver. (2) 8: 87. Eulaelaps ambulans (Thorell) Tragardh, 1910, Nat. Unter. Sarekgebirges Swed. Lap. £ (4) : 43 figs. 116-122. on Female: Chelicera with segment I, 90 »; II, 147 »; movable digit 48 pz, bir- dentate ; fixed digit bidentate ; pilus dentilis inflated basally; dorsal seta long and TRE BREMISH DERMANYSSIEDAE. (A CART) 255 stout (Text-fig. 59E). Four pairs of barbed gnathosomal setae with c.s. about 93 mu apart, hyp.2 about 70 w apart. Deutosternum with ten transverse rows of denticles (2-5 per row); corniculi 30 » long; internal malae with long, pointed inner and short, broad, outer processes, all pilose. Salivary styl conspicuous, extending beyond tips of corniculi. Tectum capituli slender, pointed, fimbriate. Pedipalp (2-56-14) with two-tined apotele. Palptrochanter with both setae barbed. Dorsal shield (984 » X 576 mu) with a dense covering of subequal setae (Text-fig. 59A) ; those on posterior margin slightly longer ; setae 71 longer, stouter and barbed. Many marginal and anterior setae barbed. Surface of shield granular. Tritosternum with base 48 p, laciniae 150 pw. Sternal shield (108 » x 168 u) with three pairs of setae and three pairs of pores ; metasternal setae free. Between s/1, 120 »; between sf1 and sf3, 120 p; sti barbed. Genito-ventral shield bears numerous setae distributed mostly on posterior two-thirds of shield. Anal shield (II0 » X 105 ») with five or six accessories in addition to the usual three anal setae ; paranal setae 66 «; postanal seta 84 ». Chaetotaxy and sclerotization of opistho- gaster as in Text-fig. 598. Peritreme extends to middle of coxa II ; anterior tip of peritrematal shield free, but lying very close to dorsal shield ; posterior part fused with podal shield of coxa IV. Chaetotaxy of legs differs from normal pattern in that genu IV has two postero- lateral setae and tibia IV has two antero- and two postero-dorsal setae ; setae ad, and pd, are not visible on tarsi II-IV. Most leg setae barbed. Distal margins of all segments except tarsi serrated. Ambulacrum IV nearly one-and-a-half times as long as ambulacrum III. Length/width (in yp) of leg segments : i! II III 1 genu 144/54 114/72 go/60 138/57 tibia 156/48 108/66 102/48 150/48 tarsus 216/42 168/48 216/42 315/42 Male: Chelicera with segment I, 75 »; II, 140 p. Cheliceral digits and sperma- dactyl as in Text-fig. 59F. Four pairs of gnathosomal setae with c.s. about 84 apart, hyp.2 about 66 » apart. Deutosternum with ten transverse rows of denticles (2-7 per row). Internal malae, salivary styli, tectum and pedipalps as in female. Dorsal shield (840 » x 480 ») asin female. Tritosternum with base 40 p, laciniae 120 pw, presternal area as in female. Holoventral shield (624 w x 150 p) with the usual sternal and metasternal setae, and with numerous accessory setae distributed over almost entire shield and extending anteriorly to between st2 and st3 (Text-fig. 59C). Between stz, 90 »; between sti and sf3, 110 » ; sft barbed. Par- and post- anal setae subequal, about 60 » long. Peritreme extends to middle of coxa II ; anterior tip of peritrematal shield fused with dorsal shield, posterior tip fused with podal shield of coxa IV. Chaetotaxy and ornamentation of legs as in female. Stout setae present on femur II (av), genu II (av), tibia II (av) and tarsus II (av, and mv). Length/width (in ,) of leg segments : G. OWEN EVANS & W. M. TILL 250 x “SSS =—s SSS THE BRITISH DERMANYSSIDAE (ACARI) 257 I II Ill IV genu 132/48 108/66 96/48 132/48 tibia 138/42 108 /60 96/48 144/45 tarsus 192/36 144/42 186/36 282/36 HapitaT: Found in Britain, Europe and the U.S.S.R. chiefly on the bodies and in the nests of a variety of rodents. Also recorded from insectivores, small carni- vores and birds. Haemogamasus pontiger (Berlese) Laelaps (Eulaelaps) pontigey Berlese, 1904, Redia 1: 260. Bregetova, 1956, Opred. Faune SSSR. 61: 1209, figs. Groschaftella pontigey: Samsinak, 1957, Cslka. parasitol. 4: 270, figs. I-2. Haemogamasus oudemansi Hirst, 1914, Bull. ent. Res. 5: 122, pls. 14-16. Keegan, 1951, Proc. U.S. nat. Mus. 101 : 240, figs. 48-49. Eulaelaps mawsoni Womersley, 1937, Austral. Antarctic Exped. 1911-1914: Scientific Rep. ser. C. Zool. & Bot. 10: 19, pl. 12, figs. 4-7. Female: Chelicera with segment I, 102 »; II, 162 »; movable digit 63 p, bi- dentate ; fixed digit bidentate, pilus dentilis setiform, dorsal seta long and stout (Text-fig. 60c). Four pairs of gnathosomal setae with c.s. about 87 pw apart, hyp.2 about 66m apart; /yp.1 simple, hyp.2—-3 and c.s. barbed. Deutosternum with 14 transverse rows of denticles (2-6 per row) ; corniculi 50 » long ; internal malae as in male (Text-fig. 60G). Tectum capituli slender, pointed, fimbriate. Salivary styli conspicuous. Pedipalp (2—5—6-14) with three-tined apotele (Text-fig. 60F) ; both trochantal setae barbed. Dorsal shield (840 » X 495 “) with numerous setae distributed as in Text-fig. 604. Setae 71 considerably stouter than the others, with fine barbs ; some of the other anterior and antero-lateral setae also barbed. Surface of shield granular, with sculpturing on the podonotal and reticulations on the opisthonotal part. Tritosternum with base serrated, 50 » long ; laciniae 126 w. Presternal area with small spines on the reticulations. Sternal shield (42 » x 160 ,) deeply concave posteriorly, with three pairs of setae and three pairs of pores ; metasternal setae free. Between st1, 117 » ; between sti and sf3, 123 »; sti barbed. Genito-ventral shield (300 » X 114 ») enlarged posteriorly, with maximum width 246 » ; bearing a pair of genital setae and about 16 accessories on the posterior part of the shield. Anal shield (99 » X III p) with two small anterior protuberances ; paranalsetae 60 p ; postanal seta 78 p. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 60B. Peritreme extends a little beyond posterior border of coxa II ; peritrematal shield free anteriorly, fused posteriorly with podal shield of coxa IV ; pore in posterior part with thickened rim. Fic. 59. Haemogamasus nidi Michael. A. Dorsum of idiosoma of female; 8B. venter of idiosoma of female; c. venter of idiosoma of male; p. deutosternum of female; E. chelicera of female; F. chelicera of male. G. OWEN EVANS & W. M: TILL 258 20 GOO Son ; NG ™ — > = \ 4 ie \ a ) \ { iS A. Dorsum of idiosoma of female; B. venter Fic. 60. Haemogamasus pontiger (Berlese). of idiosoma of female ; c. chelicera of female ; bp. venter of idiosoma of male ; E. chelicera of male; F. apotele of pedipalp; G. venter of deutosternum and hypostome of male. THE BRITISH DERMANYSSIDAE (ACARI) 259 Chaetotaxy of legs differs from normal pattern in that genu IV has two postero- lateral setae and tibia IV has two antero- and two postero-dorsal setae. Setae ad, and pd, at the extremities of tarsi II-IV are extremely minute and inconspicuous. Tarsus IV with seta fd, very long, three-fifths the length of the tarsus. Most leg setae barbed except terminal ones on tarsi. Distal margins of some segments faintly serrated. Length/width (in p) of leg segments : I II HUGE TV. genu 150/45 108/57 90/50 138/48 tibia 162/40 117/54 105/45 150/42 tarsus 246/36 192/40 210/36 300/36 Male : Chelicera with segment I, 70 » ; II, 126 »; movable digit unidentate, 45 » long (54 » with spermadactyl) ; fixed digit unidentate ; dorsal seta long and stout ; pilus dentilis setiform (Text-fig. 60E). Four pairs of gnathosomal setae as in female with c.s. about 72 » apart, hyp.2 about 54 wapart. Deutosternum with 13 transverse rows of denticles ; corniculi 45 » long ; internal malae as in Text-fig. 60G. Tectum capituli, salivary styli and pedipalps as in female. Dorsal shield (660 » x 384 ») asin female. Tritosternum with base 40 p, laciniae Io2 ». Presternal area as in female. Holoventral shield (552 » x 126 ») with four pairs of setae and three pairs of pores in the sternal region, numerous setae distributed over the genito-ventral region (Text-fig. 60D). Between sfz, 90 »; between sti and s#3, 110 »; sti barbed. Paranal setae 36 »; postanal seta 48 pp. Peritreme extends a little beyond posterior margin of coxa II ; peritrematal shield free anteriorly and posteriorly, although anterior tip lies very close to dorsal shield. Chaetotaxy of legs as in female; short, stout, blade-like setae on femur III (v) and genu III (pv). Length/width (in p) of leg segments : I II Il IV genu 130/40 102/51 90/44 126/42 tibia 138/36 102/48 100/40 138/40 tarsus 222/36 156/36 180/33 270/33 Hapitat: A cosmopolitan, facultative parasite recorded from rodents, a mole (Talpa europaea Linn.) and a bat ; also free-living in a variety of habitats including barley, sugar, wheat and rice straw, rice hulls and flax tow. Genus EULAELAPS Berlese Eulaelaps Berlese, 1903, Zool. Anz. 27 : 13. Hemilaelaps Hull, 1918, Tvans. nat. Hist. Soc. Northumb. 4: 61(lapsus for Eulaelaps). Rhinolaelaps Fonseca, 1960, Acarologia 2 : 17. Type: Gamasus stabularis Koch, 1836. Medium to strongly sclerotized mites up to 1,000 » in length. Chelicerae chelate- dentate, movable digit bidentate in the female, unidentate in the male. _Pilus dentilis 260 G. OWEN EVANS & W. M. TILL relatively long and simple. Dorsal seta and fissures normal. Spermadactyl short, grooved. Arthrodial processes present at base of movable digit in the female. ~ Capitular and hypostomal setae normal. Deutosternum with ten or more transverse rows of denticles, four to seven denticles per row. Corniculi horn-like, internal malae elongate, fringed. Pedipalpal chaetotaxy normal, apotele two-tined. Palp- trochanter with unique sensory organ ventrally. Tectum capituli with anterior and lateral margins deeply denticulate. Dorsum of idiosoma densely hypertrichous except for region of dorsal hexagon (75, 25, 76) in some species. Dorsal setae simple or minutely pilose. Tritosternum normal with lateral margins of basal region denticulate. No distinct pre-endopodal shields. Sternal shield in the female with three pairs of setae and two pairs of pores. Genito-ventral shield large with numerous setae. Anal shield free with normal three setae. Opisthogastric integument hypertrichous. Metapodal shields in the female very large and usually sub-triangular in outline. Peritrematal shields free in the female. Male with holoventral shield ; peritrematal shields free or fused with the shield. Podal shields weakly developed posterior to coxae IV. Segmental chaetotaxy of legs normal except genu IV (2—7, #—2). Coxa I] witha small anterior spine. Ambulacra with claws sometimes strongly reduced on leg I. KEY TO FEMALES Genito-ventral shield of female with lateral incisions posterior to the genital setae, lateral margins angular, bearing 47—52 opisthogastric setae ; no setae on cuticle between genito-ventral and anal shieids ae 61F). Tarsus II with some spine-like setae (Text-fig. 61C) : E. stabularis Koch (p. 260) Genito-ventral shield without lateral incisions, lateral margins rounded, bearing about 26 opisthogastric setae ; setae present on cuticle between genito-ventral and anal shield (Text-fig. 62a). Tarsus II with somestout, spur-like setae (Text-fig. 62E) E. novus Vitzthum (p. 263) Lal { KEY TO MALES 1 Tarsus II with setiform and spiniform setae ; femur II without sclerotized pro- tuberance ventrally ; E. stabularis Woch (p. 262) Tarsus IT with setiform and stout, Spur Ale setae ; femur II with conspicuous pro- tuberance ventrally distal to fissure. E. novus Vitzthum (p. 265) Eulaelaps stabularis (Koch)® Gamasus stabulavis Woch, 1836, Deutschl. C.M.A. fasc. 27, no. I. Eulaelaps stabularis: Vitzthum, 1927, Jen. Z. Naturwiss. 62 : 162. Bregetova, 1956, Opred. Faune SSSR. 61 : 100, figs. Laelaps oribatoides Michael, 1892, Tvans. Linn. Soc. Lond. Zool. 5 : 315, figs. Eulaelaps arvcualis (IWoch) Tragardh, 1912, Arch. zool. exp. gen. 48 : 577, figs. 90-93. Eulaelaps oudemansi Turk, 1945, Parasitology 36 : 137. 6 According to Johnston (1959) Laelaps pedalis Banks, 1909 and L. propheticus Banks, 1909 are also synonyms of E. stabularis. THE BRITISH DERMANYSSIDAE (ACARI) of male ; ) Lee! E is} 25 nH OO Ove Ss go oma n oo yp septs 5 63 g2 vane Ae eS 8 Ovens ae she) SS oon o ret tae oP au 3) oe et ° om i Fic. 61. Eulaelaps stabulavis (Koch). t 262 G. OWEN EVANS & W. M. TIEL Female: Chelicera with segment I, 90 p; II, 180 »; movable digit 75 p», bidentate; fixed digit tridentate, pilus dentilis short, setiform ; dorsal seta stout (Text-fig. 61E). Four pairs of gnathosomal setae with c.s. about 93 p apart, hyp.2 about 66 » apart. Deutosternum (Text-fig. 61B) with ten transverse rows of denticles (4-6 per row) ; corniculi 60 » long ; internal malae elongate, fringed. Tectum capituli with anterior and lateral margins deeply denticulate. Salivary styli conspicuous, reaching tips of corniculi. Pedipalp (2-5-6-14) with two-tined apotele; palptrochanter with sensory organ ventrally. Dorsal shield (960 » [ea] A a = ° o THE BRITISH DERMANYSSIDAE (ACARI) 201 Hirstionyssus latiscutatus (de Meillon & Lavoipierre) Liponyssus latiscutatus de Meillon & Lavoipierre, 1944, J. ent. Soc. S. Afr. 7: 62. Ichoronyssus orcadensis Turk, 1946, Ann. Mag. nat. Hist. (11) 12: 786, figs. 1-7. (Syn. nov.). Hirstionyssus musculi (Johnston) Bregetova, 1956, Opred. Faune SSSR. 61: 185, figs. Female : Chelicera with segment I, 60 »; II, 114 »; movable digit 50 ». Four pairs of gnathosomal setae with c.s. about 54 m apart, hyp.2 about 33 p apart. Deutosternum with about 14 rows of denticles (one to three denticles per row). Corniculi membranous, internal malae slender. Tectum capituli tongue-shaped, anterior margin rounded, fringed. Salivary styliconspicuous. Pedipalp (2-5—6—14) with normal chaetotaxy (Text-fig. 72c) ; apotele two-tined. Dorsal shield (552 » < 294 ») with 26 or more pairs of setae distributed as in Text- fig. 72A. The setae on the opisthonotal region are variable in number and distribu- tion. There may, for example, be one or two setae between the first and second pairs of J setae. Surface of shield granular, with reticulations near the antero-lateral margins ; unsclerotized integument of dorsum with about Ig pairs of setae, including five pairs of 7 setae. Tritosternum with base 45 p, laciniae 78 ». Sternal shield (27 » x 144 p) arched, posterior margin deeply concave ; with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Surface of shield granular with reticula- tions in the antero-lateral corners. Between sti, 70 »; between sti and s¢3, 72 p. Genital shield (108 » x 108 «) tongue-shaped, granular, with one pair of setae. Anal shield (84 » X 72 m) with paranal setae 30 p, postanal setae 36 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 72B. Peritreme extends to middle of coxa I ; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs normal ; tarsus II with setae av, and pv, stout and claw-like. Coxal spur formula (o-2-2-1). Length/width (in y) of leg segments : 1 iB Ill IV. genu 63/48 63/48 45/38 63/38 tibia 63/42 50/45 45/36 60/36 tarsus 96/36 78/36 90/30 110/32 Male: Chelicera with segment I, 45 »; II, 90 »; spermadactyl 42 » long from base of movable digit. Four pairs of gnathosomal setae with c.s. about 45 p apart, _ hyp.2 about 30 w apart. Other features as in female. Dorsal shield (468 » x 294 «) with 28 or more pairs of setae distributed as in Text- _ fig. 72D ; unsclerotized integument of dorsum with 19 pairs of setae including three | pairs of y setae. Surface of shield granular with reticulations antero-laterally. Fic. 71. a-B. Hirstionyssus latiscutatus (de Meillon and Lavoipierre), deutonymph ; dorsum (a) and venter (B) of idiosoma. c—p. Hirstionyssus sovicis (Turk), deutonymph ; dorsum (c) and venter (D) of idiosoma. LL G. OWEN EVANS & W. M. TI THE BRITISH DERMANYSSIDAE (ACARI) 293 Tritosternum with base 33 p, laciniae 72 ». Holoventral shield (360 » x I17 1) with four pairs of setae and three pairs of pores in the sternal region, four pairs of setae in the genito-ventral region. Between sf1, 50 w; between sti and st3, 72 yp. Paranal setae 33 pu, postanal seta 27 pw long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 72E. Peritreme extends to middle of coxa I; peritre- matal shield fused anteriorly with dorsal shield, posteriorly with podal elements of coxa IV. Chaetotaxy of legs normal. Tarsus II with setae av, and pv, stout and claw-like. Stout, spine-like setae present on genu IV (av), tibia IV (av) and tarsus IV (av, and mv). Coxal spur formula (o—2-2-1). Length/width (in p ) of leg segments : I II III IV genu 60/40 60/42 50/38 66/40 tibia 54/36 45/42 45/36 63/36 tarsus 80/30 66/30 75/30 100/30 Deutonymph : Chelicera with segment I, 45 »; II, 72 »; movable digit 40 yp. Four pairs of gnathosomal setae with c.s. about 40 pw apart, hyp.2 about 26 p apart. Tectum capituli with anterior margin weakly denticulate. Other gnathosomal features as in female. Dorsal shield (372 » x 180 ») with 24 pairs of setae distributed as in Text-fig. 71 ; unsclerotized integument with about 20 pairs of setae, including si, s2 and five pairs of 7 setae. Tritosternum with base 30 p, laciniae 50 ». Sternal shield (180 » x I14 ») with four pairs of setae (sternal and metasternal) and three pairs of pores. One pair of genital setae is situated on the integument lateral to the posterior tip of the shield. Between sf1, 66 »; between sti and sf3, 84 ». Anal shield (45 x 45 »); par- and postanal setae subequal in length (about 20 yw). Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 71B. Peritreme extends to anterior fourth of coxa II ; anterior part of peritrematal shield extends to posterior fourth of coxa I, posterior part not developed. Chaetotaxy of legs normal. Tarsus II with setae av, and pv, simple. Coxal spur formula (o-2-1-0). Length/width (in ») of leg segments : I II Il IV genu 50/42 48/42 34/36 45/36 tibia 48/38 42/40 36/34 45/33 tarsus 75/32 60/30 TOl27 80/30 Fic. 72. Hirstionyssus latiscutatus (de Meillon & Lavoipierre). A. Dorsum of idiosoma of female ; B. venter of idiosoma of female ; c. palptibia of female; p. dorsum of idiosoma of male; £. venter of idiosoma of male. THE BRITISH DERMANYSSIDAE (ACARI) 295 HaBitTaAT: Found in Britain on Muicrotus orcadensis Millais, Scotland, and on laboratory mice, Newcastle-upon-Tyne. Also recorded from South Africa (on Rattus vattus (Linn.) and the U.S.S.R. (on Rattus species, Mus musculus Linn., Apodemus species, Micromys minutus (Pallas), Microtus species). Hirstionyssus isabellinus (Oudemans) ?Liponyssus albato-affinis Oudemans, 1903, Tijdschr. Ned. dierk. Vereen. (2) 8: 24. Liponyssus isabellinus Oudemans, 1913, Ent. Ber. Amst. 3: 384, et Arch. Naturgesch. Abt. A 79 (9) : 80, figs. Hirstionyssus isabellinus : Bregetova, 1956, Opred. Faune SSSR. 61 : 181, figs. Hirstionyssus avvicolae Zemskaya, 1955, in Bregetova et al., Opred. Faune SSSR. 59 : 366, figs. 799-792. Female: Chelicera with segment I, 66 »; II, 135 »; movable digit 57 ». Four pairs of gnathosomal setae with c.s. about 60 p apart, yp.2 about 40 pw apart. Deutosternum with 13 rows of denticles (one or two denticles per row). Corniculi membranous, internal malae long and slender. Tectum capituli tongue-shaped, anterior margin fimbriate. Salivary styli conspicuous. Pedipalp (2-5—6-14) with two-tined apotele. Dorsal shield (558 » X 324 ») with 27 pairs of setae distributed as in Text-fig. 734, seta 72 being on the shield. Surface of shield granular with reticulations antero- laterally. Unsclerotized integument with about 16 pairs of setae, including four pairs of 7 setae (73-76). Tritosternum with base 42 p, laciniae 84 ». Sternal shield (42 » x 158 p) arched, strongly concave posteriorly, with three pairs of setae and two pairs of pores ; meta- sternals free with associated pores. Between si1, 84 »; between sf1 and st3, go yp. Surface of shield granular with reticulations antero-laterally. Genital shield (156 pu < 138 ») granular, with one pair of setae. Anal shield (96 » x 87 ) with paranal setae 33 pw, postanal seta 36 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 73B. Peritreme extends to anterior third of coxa I; anterior part of peritrematal shield free, posterior part fused with podal shields of coxa IV. Chaetotaxy of legs normal. Tarsus II with setae av, and pv, simple. Coxal spur formula (o-2-2-0). Length/width (in ,) of leg segments : I II III IV genu 66/45 66/45 45/30 63/36 tibia 66/40 50/45 42/36 60/36 tarsus 99/33 75/34 80/30 = 114/30 Fic. 73. Hirstionyssus isabellinus (Oudemans). A. Dorsum of idiosoma of female; B. venter of idiosoma of female; c. dorsum of idiosoma of male; pb. venter of idiosoma of male. 296 G. OWEN EVANS & W.M. TILL Male: Chelicera with segment I, 48 »; II, 87 »; spermadactyl 50 w. Four pairs of gnathosomal setae with c.s. about 42 pw apart, hyp.2 about 30 p apart. - Other gnathosomal features as in female. Dorsal shield (508 » x 312 ») with 27 pairs of setae distributed as in Text-fig. 73¢ ; unsclerotized integument of dorsum with 16 pairs of setae, including 73-76. Holoventral shield (360 » x 108 ) with four pairs of setae in the sternal region and four pairs in the genito-ventral region. Between sf1, 50 »; between sti and si3, 74 ». Par- and postanal setae about 27 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 73D. Peritreme extends to anterior fourth of coxa I and is fused anteriorly with the dorsal shield; posterior part of peritrematal shield weakly developed. Chaetotaxy of legs normal. Tarsus II with setae av, and pv, stout and claw-like. Coxal spur formula (o-1-2-1). Length/width (in ,) of leg segments : I II Ill IV genu 66/42 66/45 «50/40 «78/45 tibia 60/36 48/42 48/36 78/40 tarsus 90/30 80/32 78/30 120/33 Hapitat: Found in Britain on the weasel (Mustela sp.), Brentwood, Essex ; on Sorex araneus Linn., Meall Greigh, Perthshire, and on Mzcrotus agrestis (Linn.), Isle of Muck, Hebrides. Recorded by Oudemans from Putorius sp., Arvicola terrestris (Linn.) and Mus musculus Linn. in Holland, and by Bregetova from Mustela nivalis Linn., Rattus norvegicus (Berkenhout), Microtus arvalis (Pallas) and other rodents in the US... The above description of the female is based on a specimen from Mustela sp. and that of the male on a specimen from Mzcrotus arvalis presented by Dr. N. G. Bregetova. The male we have described differs from the male attributed to the species by Oude- mans (1913, fig. 302) in lacking a ventral spur on coxa II. Oudemans lists a variety of hosts for this species and it is possible that his male and female are not conspecific. Hirstionyssus blanchardi (Trouessart) Leiognathus blanchardi Trouessart, 1904, Arch. Parasitol. Paris 8 : 558. Liponyssus blanchardi:; Hirst, 1922, Proc. zool. Soc. Lond (1921) : 787, figs. Hirstionyssus blanchardi: Bregetova, 1956, Opred. Faune SSSR. 61: 177. Female: Chelicerae with segment I, 78 » ; II, 138 »; movable digit 60 ». Four pairs of gnathosomal setae with c.s. about 57 mw apart, Ayp.2 about 36 pm apart. Deutosternum with 18 rows of denticles (one or two denticles per row). Corniculi membranous, internal malae slender. Tectum capituli tongue-shaped , somewhat pointed anteriorly, margin smooth. Salivary styli conspicuous. Pedipalp (2-5-6- 13) with unideficient chaetotaxy on palptibia (Text-fig. 748) ; apotele two-tined. Dorsal shield (660 » X 324 ») with 24 pairs of setae distributed as in Text-fig. 744. Unsclerotized integument of dorsum with about 25 pairs of setae, including sI, s2 THE BRITISH DERMANYSSIDAE (ACARI) 297 Af a 1 i} Fic. 74. Hirstionyssus blanchardi (Trouessart), female; dorsum of idiosoma (a) ; palp- tibia (B) ; venter of idiosoma (c). and six pairs of y setae. Surface of shield granular, with a less heavily sclerotized marginal band. Tritosternum with base 45 p, laciniae about 75 ». Sternal shield (87 » x I41 p) rectangular, with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sti, 80 »; between str and st3, 84 uw. Genital shield {132 » X 132 pw) granular except for the weakly sclerotized marginal zone, bearing one pair of setae. Anal shield (99 » x 96 ») with paranal setae 30 p, postanal seta 33 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 74Cc. Peritreme extends to middle of coxa I; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs normal. Tarsus II with setae av, and pv, simple. Coxal spur formula (o-1-1-0). Length/width (in ») of leg segments : I II ag! IV genu 84/48 84/50 66/42 90/42 tibia 80/40 = 60/40 80/40 tarsus 100/36 — 100/33 126/36 298 G. OWEN EVANS & W. M. TILL Male : Chelicera with segment I, 72 »; II, 117 »; spermadactyl 54 ». Four pairs of gnathosomal setae with c.s. 45-52 » apart ; hyp.2 30-36 wapart. Deutosternum with about 15 rows of denticles (one to two denticles per row). Other features as in female. Dorsal shield (580-615 » X 324 ») with 26 pairs of setae distributed as in Text-fig. 754; fifteen pairs of setae on the unsclerotized integument, including six pairs of 7 setae. Surface of shield as in female. Tritosternum with base 30 p, laciniae72 4. Holoventralshield (444 « x 120-138 yp) with four pairs of setae and three pairs of pores in the sternal region, four pairs of setae in the genito-ventral region. Between sti, 60-70 »; between sfr and sf3, 87-98 w. Surface of shield granular. Par- and post-anal setae subequal in length (about 30 »). Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 758. Peritreme extends to middle of coxa I; peritrematal shields free posteriorly. Chaetotaxy of legs normal. Tarsus II with setae av, and fv, stout and claw-like Stout spine-like setae on trochanter I (d), femora I and II (ad, and pd,_.), femur III (al and v). Coxal spur formula (o-1-1-1). Legs bent, not suitable for measure- ment. Deutonymph : Chelicera with segment I, 38 »; II, 96 »; movable digit 40 ux. Four pairs of gnathosomal setae with c.s. about 36-45 pu apart, hyp.2 about 26-30 u apart. Deutosternum with 13-16 rows of denticles (one or two denticles per row). Other features as in female. Dorsal shield (408-444 » * 156-210 ») with 24 pairs of setae distributed as in Text-fig. 75c ; about 20 pairs of setae on the unsclerotized integument, including st, s2 and six pairs of 7 setae. Tritosternum with base 30 p, laciniae 57». Sternalshield (240-252 u x IIO-120 p) with four pairs of setae and three pairs of pores ; one pair of genital setae is situated on the cuticle lateral to the posterior tip of the shield. Between sf1, 70-72 u; between stz and s#3, 100-108 pw. Anal shield (48-54 mw X 50-57 mw) with par- and postanal setae subequal in length (about 15 »). Chaetotaxy and sclerotization of venter as in Text-fig. 75D. Peritreme extends to a point between coxae I and II; anterior part of peritrematal shield short, posterior part not developed. Chaetotaxy of legs normal. Tarsus II with setae av, and pv, simple. Slightly thickened setae present on femora I (ad, and #d,-,) and II (ad). Coxal spur formula (o-1-o-0). Length/width (in ») of leg segments : I II Ill IV genu 48/40 45/42 36/36 50/38 tibia 54/36 42/38 36/34 54/36 tarsus 75/30 — 60/27 93/27 Fic. 75. Hirstionyssus blanchardi (Trouessart). A. Dorsum of idiosoma of male; B. venter of idiosoma of male; c. dorsum of idiosoma of deutonymph ; pb. venter of idio- soma of deutonymph. THE BRITISH DERMANYSSIDAE (ACARI) 299 300 G. OWEN_EVANS .& W.-M... TILL Hapitat: Found on Marmota marmota (Linn.), the Alpine Marmot, Zermatt, Switzerland, and on Cynomys ludovicianus (Ord.) the Black-tailed Prairie Dog (speci-. mens kept in captivity), Horsham, England. Subfamily MACRONYSSINAE Oudemans Liponyssinae Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 2. [see p. 240] Macronyssidae Oudemans, 1936, Kritisch Historisch Overzicht der Acarologie (III) A: 430. Adults; Chelicerae essentially chelate-edentate, digits often with membranous lobes and denticulate processes ; spermadactyl relatively short and free, rarely long and entirely fused with the movable digit ; fixed digit rarely shorter than the movable. Pilus dentilis absent, dorsal seta minute or absent ; fissures normal: arthrodial processes rarely present. Chaetotaxy of venter of gnathosoma normal. Pedipalps with five free segments, apotele two-tined, rarely simple ; chaetotaxy of trochanter to tibia normal (2-5—6-14) or with tibia unideficient, rarely femur and and genu unideficient or tibia bideficient. Palptrochanter usually with medio- ventral keel in female. Deutosternum with denticles in a single file, rarely with two to five denticles in a transverse row. Hypostome somewhat elongated, corniculi membranous, internal malae free or fused. Tectum capituli elongated with margins smooth, weakly denticulated or fimbriated. Dorsal sclerotization reduced in the female, comprising an entire shield, two sub- equal shields or a podonotal shield, mesonotal scutellae and a small pygidial shield ; male with entire dorsal shield. Primary chaetotaxy of the dorsum of the idiosoma showing hypotrichy, paravertical setae (zI) normally absent. Secondary hyper- trichy of the dorsum common, particularly of the lateral and marginal series. Dorsal setae simple, pilose or subspinose. Tritosternum bipartite, laciniae smooth or pilose and often with denticulate, hyaline, lateral margins. Sternal shield in female with two or three pairs of setae and two pairs of pores; metasternal setae free, rarely absent. “‘ Porose areas” present in the region of the first pair of sternal pores or posterior margin of sternal shield with a strongly sclerotized band in some chiropteran parasites. Genital shield usually tapering posteriorly and bearing the genital setae (except in Ophionyssus) : opisthogastric setae may be present on the shield. Anal shield normal, aciculated area conspicuous, no euanal setae. Opisthogastric cuticle invariably hypertrichous. Metapodal shields small, relatively inconspicuous. Peritrematal shield fused with podal shields posterior to coxae IV ; peritemes of varying lengths. Podal shields weakly developed. Male with holoventral shield ; sternito-genital and ventro-anal shields ; sternito-genital, ventral and ventro-anal shields ; sternito-genital, ventral and anal shields ; or sternito-genital and anal shields. Genital orifice presternal. All legs six-segmented and with well developed ambulacra. Segmental chaetotaxy extremely diverse, never entirely normal. Coxae I-IV may have sclerotized ridges of varying degrees of development. Anterior spine of coxa IT usually large, rarely minute or absent. THE BRITISH DERMANYSSIDAE (ACARI) 301 Larva: Inactive non-feeding instar. Chelicerae with digits minute, edentate and lacking setae, fissures and arthrodial processes. Pedipalps five-segmented ; chaeto- taxy of tibia and tarsus deficient. Hypostome with two pairs of setae, corniculi degenerate, deutosternal denticles weak or absent. Idiosoma without distinct sclerotization. Podonotal region with eight or nine pairs of setae, opisthonotal region with three pairs of relatively long setae. Tritosternum bipartite but laciniae strongly reduced. Sternal region with normal three pairs of setae, opisthogastric region with two pairs, normal three setae associated with the non-functional anus. Stigmata and peritremes absent. Leg chaetotaxy normal for this instar. Thelarval stage is known only for a few species (Ornithonyssus, Macronyssus and Ophionyssus) Protonymph: Active feeding instar. Chelicerae, basis capituli and hypostome with the characteristics of the female. Pedipalpal chaetotaxy normal (1-—4—5-12), palptrochanter rarely with medio-ventral keel. Dorsal sclerotization comprising well developed podonotal, mesonotal and pygidial elements. Podonotal shield usually with ro or 11 pairs of setae, pygidial shield with two to seven pairs, hypertrichy of lateral and marginal series of setae of the opis- thonotum common. Tritosternum as in female. Sternal shield with three pairs of setae and two pairs of pores, genital setae present, anal shield normal. Opistho- gastric cuticle usually with four pairs of setae, hypertrichy of opisthogaster rare. Stigmata and short peritremes present. Chaetotaxy of legs normal for this instar except on genu IV (2—2, 2—1) in Ophionyssus. Deutonymph: Inactive, non-feeding instar characteristic of the macronyssines. Chelicerae degenerate, digits minute, hypostome with discrete corniculi and internal malae, deutosternal denticles as in female, gnathosomal setae normal. Pedipalps (2-5-6-12/13). Idiosoma without distinct sclerotization. Sexual dimorphism apparent in the dorsal and opisthogastric chaetotaxy, the male and female deuto- nymphs having similar chaetotaxy to the adult male and female respectively. Tritosternum simple, bipartite only at its tip. Intercoxal region with four (meta- sternals absent) or five pairs of setae. Stigmata, peritremesand segmental chaetotaxy of the legs as in the adult. KEY TO THE GENERA OF THE BRITISH MACRONYSSINAE FEMALES 1 Dorsum of the idiosoma ceed covered by discrete, subequal, podonotal and opisthonotal shields : : STEATONYSSUS (p. 329) — Dorsum of the idiosoma with an Nentize shield or, if with two or more scutal elements, the pygidial shield is relatively minute. ; : 2 2 Genu and tibia III each with one postero- Aateral —s (enn 2—2, 24 5 fibia 2—+, 21) ; tritosternum without hyaline denticulate border ; “‘ porose ’’ areas present in region of first pair of sternal pores ; fixed digit of chelicera with distal acuminate lobe and retrose denticular processes, movable digit with dorsal hyaline flap usually with denticular processes ; sclerotized ridges of varying degrees of development usually present on ventral surfaces of coxae II-IV MACRONYSSUS (p. 303) Cat tS G. OWEN EVANS: & W. M. -TIEL Genu and tibia III each with two postero-lateral setae (genu Zee 2—2; tibia 2—1, 22); tritosternum with hyaline denticulate border ; ‘‘ porose’”’ areas absent on sternal shield ; cheliceral digits with membranous lobes but usually lacking denticular processes ; without sclerotized ridges on coxae II-IV 3 Genu IV with two postero-lateral setae (2—#, 4—-2 or 2—#, 3-2) ; genital setae on integument ; ectoparasites ofreptiles . 2 : OPHIONYSSUS (p. 335) Genu IV with one postero-lateral seta (2—2, 2—1) ; genital setae situated on the genital shield ; ectoparasites of birds and mammals ORNITHONYSSUS (p. 313) MALES Ventral sclerotization comprising discrete sternito-genital and anal shields ; genu IV with two posterolateral setae (2-2,2—2 or 2-2,3—2) ; ‘ OPHIONYSSUS Ventral sclerotization comprising holoventral shield, sternito-genital and ventro-anal shields or sternito-genital, ventral and ventro-anal shields, never with discrete anal shield ; genu IV with one postero-lateral seta (2-?—3—1) : 2 Genu and tibia III each with one postero-lateral seta, segments respectiv ely Ww ith nine and eight setae ; tritosternum with a hyaline denticulate border 5 MACR ON YSSUS Genu and tibia III each with two postero-lateral setae, segments respectively with ten and nine setae ; tritosternum with a hyaline denticulate border Movable digit of chelicera with a membranous lobe between the spermadacty] and the digit ; ectoparasites of Chiroptera Us ‘STEA TON YSSUS Movable digit without membranous lobe between the spermadactyl and the digit ; ectoparasites of birds and mammals : : : ORNITHONYSSUS KEY TO THE PROTONYMPHS OF THE BRITISH MACRONYSSINAE Genu IV with six setae (I—2,2—1) . : : P ; : - é i 2 Genu IV with five setae (I—, 2—o) . d : : : : 3 Pygidial shield with three pairs of setae (Text- fig. 92a) Ophionyssus natricis (Gervais) (p. 340) Pygidial shield with two pairs of setae (Text-fig. 92c) Ophionyssus saurarum (Oudemans) (p. 344) Podonotal seta 73 and opisthonotal seta /5 absent (Text-fig. 844). . : : 4 Podonotal seta 73 and opisthonotal seta /5 present (Text-fig. 884) . : 5 Pygidial shield with four pairs of setae ; eee setae 75 about equal in Teneen to the distance between the bases of setaey5 andz5 Ornithonyssus bacoti (Hirst) (p. 324) Pygidial shield with three pairs of setae ; Fad riotal setae 75 less than one half the length of the distance between the bases of setae 75 and 25 Ornithonyssus sylviarum (Canestrini & Fanzago) (p. 321) Pygidial shield with four pairs of setae (Text-fig. 88a). : : : . ; 6 Pygidial shield with five to seven pairs of setae : 7 Pygidial shield with setae Z4, S5 and Z5 subequal Steatonyssus periblepharus Wolenati (p. 333) Pygidial shield with setae 74, S5 and Z5 increasing in size in that order Steatonyssus occidentalis (Ewing) Pygidial shield with seven pairs of setae : . : : : : : . 8 Pygidial shield with five or six pairs of setae : : : : : : : ) THE BRITISH DERMANYSSIDAE (ACART) 303 8 Unsclerotized integument of idiosoma with 16 pairs of setae (excluding 71), Text-fig. 7QA-B. . . 6 : Macronyssus kolenatii (Oudemans) (p. 314) — Unslcerotized integument of idiosoma markedly hypertrichous (Text-fig. 79c—p) Macronyssus flavus (Kolenati) (p. 310) 9g Pygidial shield with five pairs of setae Macronyssus ellipticus (IKolenati) (p. 307) — Pygidial shield with six pairs of setae : . : : : : : 4 10 to Tritosternum with denticulate, hyaline border. Pygidial shield truncated or slightly concave anteriorly ; its width at least twice its length ; setae J4 situated on the anterior margin of the shield . Ornithonyssus pipistrelli (OQudemans) (p. 329) — Tritosternum without denticulatc, hyaline border. Pygidial shield convex an- teriorly ; its width not more than 1} times its length ; setae J4 well removed from the anterior margin ofthe shield . : 3 Macronyssus uncinatus(Canestrini) Genus MACRONYSSUS Kolenati Macronyssus Kolenati, 1858, Wien. ent. Monatschr. 2: 5. Lepronyssus Kolenati, 1858, Wien. ent. Monatschr. 2: 5. Allonyssus Buitendijk, 1945, Zool. Med. 24 : 307 (syn. nov.). Hirstesia Fonseca, 1948, Proc. zool. Soc. Lond. 118 : 267. Type: Carts longimana Kolenati, 1857. Chelicerae chelate, essentially edentate, fixed digit of the female usually produced distally into an acuminate lobe and with up to three retrose denticular processes in its distal half ; movable digit with a strong dorsal hyaline flap usually with denticular processes in its anterior half. Movable digit of male with a grooved spermadactyl, its free portion extending beyond the tip of the digit by about the length of the digit, rarely longer (M. ellipticus). Pilus dentilis apparently absent, dorsal seta present, fissures normal. Arthrodial processes absent or in the form of microprocesses. Chaetotaxy of venter of gnathosoma normal, pedipalp with five free segments and a _ two-tined apotele. Chaetotaxy of pedipalps, trochanter to tibia (2-5—6-14), rarely (2-5-6-13) as in M. ellipticus. Palptrochanter with a conspicuous ventral keel. Deutosternum with 8-12 rows of denticles arranged in a single file, or some rows comprising two to five denticles. Corniculi membranous, internal malae simple. Tectum capituli elongate, tapering distally to an obtuse point. Entire dorsal shield with 21—28 pairs of setae in the female, 21~29 pairs in the male ; paravertical setae (z1) absent. Tritosternum normal, bipartite, lacking denticulate hyaline border. Sternal shield of the female with three pairs of setae and two pairs of pores. ‘‘ Porose ’’ areas present in the region of the first pair of sternal pores, weakly or strongly defined. _ Genital shield tapering posteriorly and with genital setae only ; weakly sclerotized | area extending from posterior margin of the genital shield with one to several setae _insome species. Anal shield pear-shaped, with normal chaetotaxy ; region posterior to postanal seta often markedly elongated. Metapodal shields poorly defined. | Opisthogastric cuticle shows hypertrichy. Peritrematal shields fused with the podal shields posterior to coxae IV. Podal shields weakly developed. Male with holo- | | | | | | | | 304 G. OWEN EVANS & W. M. TILL ventral shield, with sternito-genital and ventro-anal or with sternito-genital, ventral and ventro-anal shields. Presternal area in both sexes weakly sclerotized. Chaetotaxy of legs normal except genu IV which has two ventral setae (2—2, 31), Coxa II with prominent anterior spine. Sclerotized ridges of varying degrees of development usually present on ventral surfaces of coxae II-IV. All ambulacra with well-developed claws. Protonymph with dorsum of idiosoma normally bearing 29 pairs of setae. Podo- notal shield with ten pairs of setae, verticals on the integument. Pygidial shield with seven pairs of setae. Normal complement of setae reduced by absence of one or two pairs of setae on the pygidial shield, or increased by hypertrichy of lateral unsclerotized integument. Venter with three pairs of sternal setae, one pair of genital and four pairs of opisthogastric setae. Opisthogaster may show hypertrichy. Anal shield with normal complement of setae. Gnathosoma essentially as in female. Chaetotaxy of legs and pedipalps normal. KEY TO FEMALES 1 Opisthonotal region of the dorsal shield with nine pairs of setae (J series deficient, Text-fig. 764) ; majority of deutosternal denticles compound (Text-fig. 76c) . — Opisthonotal region of the dorsal shield with 12 pairs of setae ; deutosternal denticles simple, ina single file (Text-fig. 77B) 5 : : : 3 2 Sternal‘‘ porose ’’ areas weakly defined, irregular (Text- fg. 768) ; peritreme reduced, extending to the level of the posterior third of coxa II ; podonotal region with 12 pairs of setae, 71 on integument (Text-fig. 76a) . ; . M.ellipticus (Kolenati) (p. 306) — Sternal ‘“ porose’”’ areas conspicuous, subcircular, bordered by a sclerotized ring (Text-fig. 81B) ; pertreme extending to the level of the middle of coxalI ; podonotal region with 14 pairs of setae, 71 on shield (Text-fig. 81a) M. uncinatus (Canestrini) (p. 315) 3. Bases of setae 26, Zr and Si lying more or less in a straight line (Text-fig. Soa) ; sternal “‘ porose’’ areas as in gas 80B ; lateral and opisthogastric cuticle moderately setose. A M. kolenatii (OQudemans) (p. 312) — Seta z6, Z1 and Sti forming more or less the points of an equilateral triangle (Text-fig. 78a) ; sternal “‘ porose”’ areas as in Text-fig. 78B ; lateral and opisthogastric cuticle densely setose . : : . : : M. flavus (Kolenati) (p. 308) to Key To MALES I Spermadactyl extremely long, about as long as the second cheliceral segment M. ellipticus (Kolenati) (p. 306) — Spermadactyl obviously shorter in length than the second cheliceralsegment . : 2 2 Three pairs of dorsal marginal setae stout, thorn-lke ; ventral sclerotization com- prising separate sternito-genital, ventral and ventro-anal shields M. uncinatus (Canestrini) — None of the marginal setae conspicuously enlarged ; without separate ventral shield 3 3. Sternito-genital and ventro-anal shield distinctly separated (Text-fig. 78D) ; un- sclerotized dorsal integument with numerous short setae (Text-fig. 78c) M. flavus (Kolenati) (p. 310) — With a holoventral shield ; sternito-genital and ventro-anal regions connected at the level of the posterior margin of coxae IV (Text-fig. Sop) ; unsclerotized dorsal integument with about seven pairs of setae M. kolenatii (Oudemans) (p. 314) THE BRITISH DERMANYSSIDAE (ACARI) venter of idiosoma (c) ; chelicera (D-E). venter of gnathosoma B); ( Fic. 76. Marconvssus ellipticus (Kolenati), female ; dorsal shield (a) ; 306 G. OWEN EVANS & W. M. TILL Macronyssus ellipticus (Kolenati) Caris elliptica Kolenati, 1857, Parasit. Chiropt. : 16. Liponyssus ellipticus: Hirst, 1922, Proc. zool. Soc. Lond. (1921) : 795, fig. 36. Ichoronyssus mohrae Vitzthum, 1932, Z. Parasitenk. 4: 32. Female: Chelicera with segment I, 45 »; II, 170 %; movable digit 42 yp long, hooked terminally, dorsal hyaline flap with two denticulate processes. Fixed digit with two denticulate processes and a pointed distal process which appears variously shaped depending on the orientation of the chelicera (Text-figs. 76D and E). Pilus dentilis absent, dorsal seta present. Four pairs of gnathosomal setae with c.s. about 60 p apart, hyp.2 about 45 pw apart. Deutosternum with 12 transverse rows of denticles (two to five denticles per row). Corniculi membranous, internal malae simple, tectum capituli tapering. Pedipalp (2—5-6-13) with two-tined apotele ; palptrochanter with conspicuous ventral keel (Text-fig. 76c). Dorsal shield (774 4 * 348 ») with about 20 pairs of setae distributed as in Text-fig. 760A; setae 71 off the shield. Surface of shield with sculptured areas and a weak reticulate pattern. Unsclerotized integument of dorsum hypertrichous. Tritosternum with base 36 p , laciniae 114 w. Sternal shield (105 p X 174 pn) with irregular punctate areas in the antero-lateral corners ; with three pairs of setae and two pairs of pores; metasternal setae free. Between sf1, 80 »; between sf1 and sf3, 117 w. Genital shield (135 » X 102 ») with genital setae only. Anal shield (117 # X 100 p) pear-shaped with strongly arched anterior margin ; paranal setae about 40 pw long, postanal seta about 54 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 768. Peritreme extends to posterior third of coxa II ; peritrematal shield free anteriorly ; posterior part fused with podal elements of coxa IV. Chaetotaxy of legs normal except genu IV which has two ventral setae (2—2, 31). Coxa II with prominent anterior spine. Length/width (in yz) of leg seg- ments : I II Ill IV genu 120/60 102/66 84/57 144/60 tibia 114/54 84/60 © g0/54. 132/54 tarsus 192/48 162/54 180/48 240/50 Male: Chelicera with segment I, 30 »; II, 123 »; movable digit 36 pu long, broad, weakly sclerotized ; spermadactyl greatly enlarged, 87 » long from base of movable digit, boat-shaped. Fixed digit edentate. Four pairs of gnathosomal setae with c.s. about 50 m apart, hyp.2 about 42 m apart. Other gnathosomal features as in female. Dorsal shield (648 » Xx 288 ,) with 26 pairs of setae. Sternito-genital shield (270 w X 138 «) with four pairs of long sternal and a pair of short genital setae. Between sti, 63 »; between sfi and sf3, 110 ». Ventro-anal shield with irregular lateral margins ; bearing six asymmetrically arranged setae excluding the anals. Paranal setae 33 p, postanal seta 42 » long. Peritreme extends to about middle of coxa II. —E—————————— THE BRITISH DERMANYSSIDAE (ACARI) 307 Fic. 77. A. Macronyssus ellipticus (Kolenati), protonymph; dorsum of idiosoma. B-c. Macronyssus flavus (Wolenati), female ; venter of gnathosoma (B) ; chelicera (c). Chaetotaxy of legs as in female. Some relatively long, stout setae on femur I (ad, and pd,) and femur II (ad, and pd,-,)._ Legs bent and not suitable for measure- ment. Protonymph : Chelicera with segment I, 30 »; II, 114 »; movable digit 33 p. Gnathosoma essentially as in female. Four pairs of gnathosomal setae with c.s. about 45 » apart, hyp.2 about 36 wapart. Deutosternum with 12 transverse rows of denticles (one to four denticles per row). Pedipalp (1-4-5-12) with two-tined apo- tele ; trochanter with conspicuous ventral keel. Podonotal shield (282 » x 216 ) with ten pairs of setae as in Text-fig. 774 ; setae 71 situated in front of shield. Pygidial shield (114 » x 144 ») with anterior Margin convex ; bearing five pairs of setae. Unsclerotized integument of dorsum bears 11 pairs of setae. 308 G. OWEN EVANS & W. M. TILL Tritosternum with base 30 p, laciniae I00 pw. Sternal shield (180 p xX 126 p) with three pairs of setae and two pairs of pores. Between sf1, 70 1; between sf1 and s#3, 117 ». Anal shield (66 » x 66 ») with paranal setae 36 p, postanal seta 45 p long. Peritreme extends to middle of coxa III. Chaetotaxy of legs normal. Coxa II with conspicuous anterior spine. Length/ width (in p) of leg segments : I II Ill IV genu 70/52 66/54 54/48 75/48 tibia 66/45 51/48 48/45 68/45 tarsus 120/40 90/40 90/40 135/40 Hapitat: Ectoparasitic on Myotis myotis (Borkhausen), M. daubentonit (Kuhl) and M. nattereri (Kuhl) in Suffolk, England. Recorded from Myotis species and also from Rhinolophus ferrumequinum (Schreber), R. hipposideros (Bechstein) and Plecotus auritus (Linn.) in Europe. Macronyssus flavus (Kolenati) Dermanissus flavus Kolenati, 1857, Parasit. Chiropt. : 19. Lepronyssus flavus Kolenati, 1859, S.B. Akad. Wiss. Wien. 35: 184. Liponyssus flavus: Hirst, 1922, Proc. zool. Soc. Lond. : 791, figs. 34-35. Liponyssus lobatus : Oudemans, 1903, Tijdschr. Ned. dierk. Vereen. (2) 8 : 18, figs. (non Kolenati, 1857). Liponyssus britannicus Radford, 1941, Parasitology 33 : 311. Female: Chelicera with segment I, 42 »; II, 144 »; movable digit 45 p long, with a dorsal, hyaline flap which is irregularly denticulate in its anterior half. Fixed digit with an acuminate terminal lobe and three retrose denticular processes in its distal half (Text-fig. 77c). Pilus dentilis absent, dorsal seta minute. Four pairs of gnathosomal setae with c.s. 50 » apart, Ayp.2 about 36 wp apart. Deutosternum with nine denticles arranged in a single file. Corniculi membranous, internal malae simple. Tectum capituli elongate, pointed distally. Pedipalp (2-5-6—14) with two- tined apotele. Palptrochanter with a conspicuous ventral keel (Text-fig. 778). Dorsal shield (624 » 288 p) with 28 pairs of simple setae distributed as in Text- fig. 78A. Surface of shield reticulated. Unsclerotized integument of dorsum shows marked hypertrichy. Tritosternum with base 33 p, laciniae 105 ». Sternal shield (66 « X 168 «) witha distinct reticulate pattern in the antero-lateral corners, bearing three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between s/I, 54 »; between sfx and sf3, 96 ». Genital shield (117 » X 84 ») with an overall reticulate pattern, bearing one pair of genital setae. Anal shield (90 » X 96 ») with par- and postanal setae about 24 » long. Sclerotization and chaetotaxy of opistho- ——— Fic. 78. Macronyssus flavus (iolenati). A. Dorsum of idiosoma of female; B. venter of idiosoma of female ; c. dorsum of idiosoma of male; pb. venter of idiosoma of male. 309 THE BRITISH DERMANYSSIDAE (ACARI) 310 G. OWEN EVANS & W. M. TILL gaster as in Text-fig. 788. Peritreme extends to anterior third of coxa I ; peritrema- tal shield free anteriorly ; posterior part fused with podal elements of coxa IV. Chaetotaxy of legs normal except genu IV which has two ventral setae (2—?, ?—1). Coxae II with large anterior spine. Length/width (in p) of leg segments : if II III IV genu 72/50 7257 48/45 66/45 tibia 72/45 58/54 48/45 60/42 tarsus 105/40 70/48 78/42 126/40 Male: Chelicera with segment I, 24 »; II, 81 »; movable digit 30 p long, strongly hooked terminally ; spermadactyl grooved, 45 » long from base of movable digit. Fixed digit edentate. Four pairs of gnathosomal setae with c.s. about 45 pz apart, hyp.2 about 38 » apart. Other gnathosomal features as in female. Dorsal shield (540 » X 252 p») with 27 or 28 pairs of setae as in Text-fig. 78c. Unsclerotized integument of opisthonotum bears numerous very short, fine setae. Tritosternum with base 20 p, laciniae 96 ». Sternito-genital shield (240 x 129 p) reticulated, with five pairs of setae and three pairs of pores. Between siI, 63 pn; between sf and st#3, 96 w. Ventro-anal shield (204 » x 66 pw) with about fifteen setae excluding the anals ; par- and postanal setae about 20 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 78p. Peritreme extends to middle of coxa I; peritrematal shield fused anteriorly with dorsal shield ; posterior part fused with podal elements of coxa IV. Chaetotaxy of legs asin female. Leg II with some setae situated on small tubercles (anterior seta on coxa ; av on femur ; av,, pv; and mv on tarsus). Setae pd and pi, on genu IV and dorsal setae on femur IV stout and spine-like. Coxa II with anterior spine. Length/width (in ») of leg segments not measurable in the single male we have examined. Protonymph ; Chelicera with segment I, 30 »; II, 110 »; movable digit 33 x. Four pairs of gnathosomal setae with c.s. about 36 » apart ; hypf.2 about 30 p apart. Pedipalp (1-4-5-12) with two-tined apotele. Other gnathosomal features as in female. Podonotal shield (240 » x 162 «) with ten pairs of setae (Text-fig. 79C), 71 situated on the unsclerotized integument. Pygidial shield slightly wider than long, with seven pairs of setae. Two pairs of mesonotal scutellae present between the anterior and posterior dorsal shields. Unsclerotized integument of opisthonotum markedly hypertrichous. Tritosternum with base 26 p, laciniae 84 ». Sternal shield (150 » X I00 p) with three pairs of setae and two pairs of pores. Between sfI, 54 «; between sfI and s?3, 105 #. Anal shield (42 » xX 48 p) with par- and postanal setae about I5 4 long. Fic. 79. A-B. Macronyssus kolenatit (Oudemans), protonymph; dorsum (A) and venter (B) of idiosoma. c—p. Macronyssus flavus (Kolenati), protonymph; dorsum (c) and venter (D) of idiosoma. 311 THE BRITISH DERMANYSSIDAE (ACARI) 312 G. OWEN EVANS & W. M. TILL Chaetotaxy of opisthogaster as in Text-fig. 79D. Peritreme extends to about middle of coxa III. Chaetotaxy of legs normal. Length/width (in p) of leg segments : I II III IV genu 50/50 45/50 36/45 42/40 tibia 48/45 40/48 36/42 42/40 tarsus 60/40 60/38 54/36 72/36 Hasitat : Anectoparasite of the common Noctule Bat, Nyctalus noctula (Schreber), in Britain and Europe. Macronyssus kolenatii (Oudemans) Liponyssus kolenatii Oudemans, 1902, Ent. Ber. Amst. 1 (8) : 48 et 1903, Tijdschr. Ned. dierk. Vereen. (2) 8: 25, figs. Liponyssus pipistrelli Oudemans, 1906, Ent. Ber. Amst. 2 (27) : 37 (non Oudemans, 1904). Liponyssus cyclaspis Oudemans, 1906, Ent. Ber. Amst. 2 (28): 61 (nom. nov. pro L. pipistyelli Oudemans, 1906) et 1915, Arch. Naturgesch. 81 (A) : 165, figs. 128-134. Hirstesia brittent Radford, 1953, Parasitology 42 : 239. Female: Chelicera with segment I, 33 »; II, 130 »; movable digit 42 p, distal end of dorsal hyaline flap thorn-like and usually bent at right angles to chela ; fixed digit with slender denticular processes. Pilus dentilis absent. Four pairs of gnatho- somal setae with c.s. about 42 » apart, hyp.2 about 33 wapart. Deutosternum with eight denticles arranged in a single file. Corniculi membranous, internal malae simple, tectum capituli tapering to an obtuse point. Pedipalp (2-5-6—14) with two- tined apotelele; palptrochanter with a conspicuous ventral keel. Dorsal shield (540 » X 252 ») with 28 pairs of setae distributed as in Text-fig. Soa. Surface of shield reticulated. Unsclerotized integument of dorsum hypertrichous. Tritosternum with base 21 p, laciniae 84 ». Presternal area reticulated. Sternal shield (57 # X 129 #) with a pair of elliptical, finely striated areas in the antero- lateral corners ; with three pairs of setae and two pairs of pores ; metasternal setae free with associated pores. Between sfI, 45 «; between sf1 and sf3, 78 uw. Genital shield (90 » X 70 p) reticulated, bearing a pair of genital setae and with an unpaired seta situated on a weakly sclerotized posterior extension of the shield. Anal shield (75 w X 75 ») with anterior margin almost straight ; paranal setae 2I p, postanal seta 24 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 8oB. Peritremes extend to middle of coxae I; peritrematal shields free anteriorly, fused posteriorly with podal elements of coxae LV. Fic. 80. Macronyssus kolenatii (Qudemans). A. Dorsum of idiosoma of female; B. venter of idiosoma of female ; c. dorsum of idiosoma of male; D. venter of idiosoma of male. S53 THE BRITISH DERMANYSSIDAE (ACARI) 314 G. OWEN EVANS & W. M. TILL Chaetotaxy of legs normal except genu IV which has two ventral setae (2—2, —1). Femur I with setae ad, and pd, long and stout. Coxa II with large, pointed 3 z anterior spine. Length/width (in ») of leg segments : I II III IV genu 63/48 60/54 38/45 54/45 tibia 54/45 45/50 36/42 50/42 tarsus 72/40 78/40 60/40 80/38 Male: Chelicera with segment I, 36 »; II, 87 1; movable digit 30 » long, eden- tate ; spermadactyl grooved, about 45 » long from base of movable digit. Fixed digit edentate. Four pairs of gnathosomal setae with c.s. about 40 pu apart, hyp.2 about 33 » apart. Deutosternum with ro denticles arranged in a single file ; cor- | niculi about 36 » long, membranous. Other gnathosomal features as in female. Dorsal shield (504 » 270 ») with 28 or 29 pairs of setae distributed as in Text- fig. 80c. Unsclerotized integument of dorsum with seven pairs of setae. Tritosternum with base 27 p, laciniae 90 ». Holoventral shield (380 p Xx I05 p) with five pairs of setae in the sternito-genital region and about 20 setae in the ventral region. Between sf1, 45 »; between sti and s¢3, 81 ». Paranal and postanal setae about 18 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 80D. Peritremes extend to middle of coxae I; peritrematal shields fused anteriorly with | dorsal shield, posteriorly with podal elements of coxae IV. Chaetotaxy of legs as in female. Length/width (in pz) of leg segments : I II III IV genu 66/45 60/50 40/40 54/40 tibia 54/40 45/45 36/36 48/38 tarsus 69/33 50/36 54/33 75,134 Protonymph : Chelicera with segment I, 15 »; II, 72 »; movable digit 27 zu. Gnathosomal features essentially as in female. Capitular setae about 30 p apart, hyp.2 about 27 w apart. Pedipalp (I-4-5-12) with two-tined apotele. Podonotal shield (180 » x 147 ») with Io pairs of setae as in Text-fig. 79A ; setae jz in front of shield. Pygidial shield (72 » x 99 ») with seven pairs of setae. Two pairs of mesonotal scutellae present. Unsclerotized integument of dorsum with eleven pairs of setae (excluding 71). Tritosternum with base 21 p, laciniae 54 ». Sternal shield (110 p X 75 ») with three pairs of setae and two pairs of pores. Between sfI, 42 «; between sfI and sf3, | 75 p. Anal shield (36 » X 39 ») with par- and postanal setae about 15 4 long. Opisthogaster with five pairs of setae (Text-fig. 79B). Peritremes extend to middle of coxae III. Chaetotaxy of legs normal. Femur I with setae ad, and fd, long and thick. | Length/width (in ) of leg segments : THE BRITISH DERMANYSSIDAE (ACARI) 315 I II III IV genu 36/42 36/45 24/34 30/34 tibia 36/36 30/39 21/34 30/34 tarsus 45/30 42/30 39/27 51/27 HapitaT: An ectoparasite on bats of the genus P7pistrellus. Recorded from P. pipistrellus (Schreber) in Britain and Europe and from P. kuhli (Kuhl) in Egypt. Macronyssus uncinatus (Canestrini) Leiognathus uncinatus Canestrini, 1885, Acarvofauna ital. 1:122; Berlese, 1889, Acar. Mvr. Scorp. Ttal. fasc. 53, no. 7. Liponyssus rhinolopht Oudemans, 1902, Tijdschr. Ned. dievk. Vereen. (2) 7: 65. Liponyssus sternalis Hirst, 1922, Proc. zool. Soc. Lond. (1921) : 781, figs. 25-26. Female: Chelicera with segment I, 54 »; II, 150 »; movable digit 4o yu long, truncate tip with acute dorsal angle. Fixed digit with two slender, subequal, denticular processes. Pilus dentilis absent. Four pairs of gnathosomal setae with c.s. about 63 » apart; Ayp.2 about 45 » apart. Deutosternum with 10-12 rows of denticles (1-4, usually one or two denticles per row). Corniculi membranous, internal malae simple. Pedipalp (2-5—6-14) with two-tined apotele ; palptrochanter with conspicuous ventral keel. Dorsal shield (798 » x 372 ») with 25 pairs of setae distributed as in Text-fig. 81. Marginal setae of podonotal region of shield conspicuously longer than setae 74-j6 and 25-26. Surface of shield granular and weakly reticulated ; unsclerotized integu- ment of opisthonotum hypertrichous. Tritosternum with base 54 pu, laciniae 132 wp. Sternal shield (84 » x 180 pu) witha pair of rounded porose areas each surrounded by a heavily sclerotized margin. The shield bears three pairs of setae and two pairs of pores; metasternals free with associated pores. Between sti, 72 »; between sfi and sf3, 105 p. Genital shield (150 » X go p) with a weakly sclerotized extension of its tip bearing from one to a maximum of five opisthogastric setae. Anal shield (130 » x 96 ») with paranal setae about 45 » long, postanal seta 60 » long. Chaetotaxy and sclerotization of opistho- gaster as in Text-fig. 818. Peritreme extends to middle of coxa I; peritrematal shield free anteriorly, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs normal except genu IV which has two ventral setae (2—}7, ?—1). Coxa II with pronounced anterior spine. Length/width (in yp) of leg segments : I II ELT IV genu 114/78 120/78 102/72 150/72 tibia 114/70 96/72 90/72 144/66 tarsus 210/60 174/60 198/63 288 /60 Male and protonymph : Described by Dusbabek (1964). ZOOL. 14, 5. 21 G. OWEN EVANS & W. M. TILL Fic, 81. Macronyssus uncinatus (Canestrini), female; dorsum (A) and venter (8) of idiosoma. Hasitat: An ectoparasite of bats. Reported from Rhinolophus hipposideros (Bechstein) in Britain and from R. ferrumequinum (Schreber), R. ewryale Blasius, Pipistrellus pipistrellus (Schreber), Myotis emarginatus (Geoffroy) and Miniopierus schreibersi (Natterer) in Europe. Genus ORNITHONYSSUS Sambon Leiognathus Canestrini, 1884, Atti ist. Veneto (6) 2: 1573 (nom. preocc.). Ornithonyssus Sambon, 1928, Ann. trop. Med. Parasit. 22: 105. Bdellonyssus Fonseca, 1941, Cienca 2 : 262. Fonsecaonyssus Radford, 1950, Parasitology 40 : 373. Type: Dermanyssus sylviarum Canestrini and Fanzago, 1877. Chelicerae chelate, edentate, without denticular processes; fixed digit in the female terminating in a hyaline boss. Spermadactyl grooved, free distally, relatively short. Pilus dentilis absent, dorsal seta minute, fissures present. Arthrodial THE BRITISH. DERMANYSSIDAE (ACART) 317 processes absent. Chaetotaxy of venter of gnathosoma normal, pedipalps with five free segments and a two-tined apotele. Chaetotaxy of palptrochanter to tibia (2-5-6-13), rarely (2-5-6-12). Seta p/ on palpfemur of male usually situated on a tubercle. Palptrochanter usually with an antero-ventral keel in the females, rarely in the males. Deutosternum with 9-11 denticles arranged in a single file. Corniculi membranous, internal malae simple, free or fused. Tectum elongate, tapering to an obtuse or acute point, lateral margins smooth or weakly denticulate. Dorsal idiosomal sclerotization reduced in the female and normally comprising an entire dorsal shield strongly tapering behind the level of 76 and 26, rarely with podonotal shield bearing ten pairs of podonotal setae and one pair of opisthonotal setae (JI), and a separate pygidial shield with two pairs of setae (O. avidus). Podo- notal region with 10-12 pairs of setae, 73 present or absent. Opisthonotal region with 6-11 pairs of setae, J/5 present or absent. Lateral cuticle hypertrichous. Dorsal idiosomal sclerotization in the male more extensive, podonotal region with 10-14 pairs of setae, opisthonotal region with 7-11 pairs. Lateral cuticle less setose than in the female. Dorsal setae smooth or sparsely barbed. Tritosternum bipartite with hyaline denticulate border. Sternal shield normally with three pairs of setae and two pairs of pores, rarely with two pairs of setae and two pairs of pores (O. sylviarum). Genital shield strongly tapering posteriorly with genital setae only. Anal shield normal. Metapodals poorly developed. Opistho- gastric cuticle shows hypertrichy. Peritrematal shield fused with podals of coxae IV, free anteriorly. Podalshields weakly developed. Males with slender holoventral shield. Peritrematal shield free or fused anteriorly with the dorsal shield. Chaetotaxy of legs normal except genu III (2—?, {—2), tibia III (2—1+, 2—2) and genu IV (2—2, 2—1). Rarely genu and tibia I with two ventrals only and genua II and III with one ventral in some populations of O. syluiarum. Coxa II with anterior spine sometimes minute. Coxa III with retrograde spur only in O. pereiva; no sclerotized ridges. Protonymph with dorsum of idiosoma provided with a podonotal shield bearing 10 (73 lacking) or I1 pairs of setae, two or more pairs of mesonotal scutellae, and a pygidial shield bearing from three to six pairs of setae. The unsclerotized integu- ment of the dorsum usually bears 15 or 16 pairs of setae. Venter with three pairs of sternal setae, one pair of genital and four to five pairs of opisthogastric setae. Anal shield with normal chaetotaxy. Gnathosoma essentially asinfemale. Chaetotaxy of legs and pedipalps normal. Palptrochanter with or without ventral keel. The above concept of Ovnithonyssus is considerably wider than that of Furman & Radovsky (1963). These authors restrict this genus to New World forms, except for four species, O. bacoti, ondatrae, syluiarum and bursa, “ which have been widely dis- persed by human activity’. They correctly point out that the species included in their concept of Orvmithonyssus differ from Old World forms in both sexes having certain idiosomal setae, particularly the caudal and marginal setae, barbed and seta pl on the palpfemur of the male situated on a tubercle. It is possible that the New and Old World species of Ovnithonyssus s. lat. are not congeneric but their status must await a comprehensive revision of the complex. 318 G. OWEN EVANS & W. M. TILL KEY TO FEMALES 1 Dorsal shield with setae 73 present ; opisthonotal region of shield with eight pairs of setae. Dorsal setae smooth. Peritremes extend to middle of coxae II. First pair of sternal setae about half as long as the second. On bats (Eptesicus and Plecotus) . : O. pipistrelli (Oudemans) (p.326) — Dorsal shield with sete J 3 absent ; opisthonotal region of shield with not more than seven pairs of setae. Some dorsal setae barbed. Peritremes extend to posterior margin of coxae I. Sternal setae subequal . : ; : : . 5 2 2 Dorsum with setae sq4 and S5 situated on the shield ; setae of J series relatively long, Jt reaching base of J2 (Text-fig. 834). Sternal shield with three pairs of setae. Opisthogastric cuticle relatively densely setose (Text-fig. 83p). Palptibia with 13 setae. Onsmallmammals_ . : : O. bacoti (Hirst) (p. 321) — Dorsum with setae s4 and S5 not on the Shicidl: setae of J series relatively short, Jt extending less than half way to base of J2 (Text-fig. 82a). Sternal shield with two pairs of setae. Opisthogastric cuticle relatively sparsely setose (Text-fig. 828). Palptibia with 12 setae. On birds O. sylviarum (Canestrini & Fanzago) (p. 318) KEY To MALES 1 Dorsal shield with 14 pairs of setae in the podonotal region. Coxae II and III with anterior setae stout and spine-like. Palpfemur with seta p/ normal, not situated on atubercle. On bats (Eptesicus and Plecotus) O. pipistrelli (Oudemans) (p. 326) — Dorsal shield with 10 or 11 pairs of setae in podonotal region. Coxae II and III with anterior setae normal. Palpfemur with seta #/ situated onatubercle . : 2 Dorsum with seta S5 on shield. Setae of 7 series relatively long, seta j74 reaching base of75. Holoventral shield with eight pairs of setae, excluding anals. Palptibia with 13 setae. Onsmall mammals ; : : O. bacoti (Hirst) (p. 323) - Dorsum with seta S5 not on the shield. Sere i the 7 series relatively short, seta 74 reaching half-way to base of 75. Holoventral shield with six pairs of setae, excluding anals. Palptibia with 12 setae. On birds O. sylviarum (Canestrini & Fanzago) (p. 320) is) Ornithonyssus sylviarum (Canestrini & Fanzago) Dermanyssus syluiavum Canestrini & Fanzago, 1877, Atti Ist. Veneto 5: 124. Liponyssus americanus Banks, 1906, Proc. ent. Soc. Wash. 8 : 136. Liponyssus canadensis Banks, 1909, Proc. ent. Soc. Wash. 11 : 134. Liponyssus pacificus Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 19 Female: Chelicera with segment I, 20 »; II, 150 »; movable digit 36 u long, with a dorsal “trough”; both digits edentate; pilus dentilis and arthrodial processes absent. Four pairs of gnathosomal setae with c.s. about 50 mw apart, Ayp.2 about 36 » apart. Deutosternum with about ro denticles arranged in a single file. Corniculi membranous ; internal malae with inner processes fused, outer leat-like. Tectum capituli tapers to a sharp point, margin weakly denticulate. Pedipalp (2-5-6-12) with two-tined apotele ; palptrochanter with small ventral Keel. Fic. 82. Ovnithonyssus sylviavum (Canestrini and Fanzago). A. Dorsum of idiosoma of female; B. venter of idiosoma of female; c. venter of idiosoma of male; pD. dorsum of idiosoma of protonymph. 319 THE BRITISH DERMANYSSIDAE (ACARI) 320 G. OWEN ENX-ANS*& IWi. Me fiir Dorsal shield (534 » X 228 pu) usually with 17 pairs of setae distributed as in Text- fig. 82A ; one or more of the setae in the opisthonotal region may be absent. Un- sclerotized integument of dorsum hypertrichous. Tritosternum with hyaline denticulate membrane, base 42 yp, laciniae 75 p. Sternal shield (36 » x I00 ») reticulated and bearing two pairs of setae and two pairs of pores ; s¢3 and stq on unsclerotized cuticle. Between sf1, 66 »; between str and st3, 52 pw. Genital shield (132 » x 60 uw) tapering, with genital setae only. Anal shield (96 » x 66 ) with par- and postanal setae subequal, about 30 wlong. Chaeto- taxy and sclerotization of opisthogaster as in Text-fig. 82B. Peritremes extend to posterior margin of coxae I; peritrematal shields free anteriorly, fused posteriorly with podal elements of coxae IV. Chaetotaxy of legs generally normal except genu III (2—, 57;—2), tibia III (2—1, 2—2) and genu IV (2—#, #—1). In addition, one of the ventral pclae pay be lacking on genu I (2—,4,, #2), tibia I (2-745, 2) and genu IT (2—#, ,7,—2) in some specimens. Coxa II without anterior spine. Length/width (in yp) of leg segments : I II Ill IV genu 60/38 54/45 48/36 60/33 tibia 60/34 50/38 48/33 63/33 tarsus 110/34 90/36 96/30 130/30 Male: Chelicera with segment I, 20 »; II, 90 w»; movable digit 27 » long ; both digits edentate ; spermadactyl 36 » long from base of movable digit. Four pairs of gnathosomal setae with c.s. about 50 w apart, Wvp.2 about 36 wapart. Deutosternum with ten denticles arranged in a single file. Corniculi membranous ; internal malae blade-like. Palptrochanter without a ventral keel. Other gnathosomal features as in female. Dorsal shield (444 » X 210 ») with chaetotaxy as in female, except that it may incorporate one or more marginal setae in the opisthonotalregion. The short postero- lateral seta (x) varies in position ; it may be posterior to seta Z4 as in the female, level with it or anterior toit. Unsclerotized integument of dorsum hypertrichous. Tritosternum with base 24 p, laciniae 57 ». Holoventral shield (360 « X 93x) narrow, with six pairs of setae excluding the anals. Between stI, 54 »; between st1 and st3, 70 w. Par- and postanal setae subequal, about 24 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 82c. Peritreme extends to middle of coxa II; peritrematal shields free anteriorly, fused posteriorly with podal shields of coxae IV. Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : I i Ill IV genu 46/32 45/38 40/33 54/30 tibia 48/30 42/33 38/27 52/28 tarsus 87/27 75/30 75/27 102/27 THE BRITISH DERMANYSSIDAE (ACARI) 321 Protonymph ; Chelicera with segment I, 15 w; II, 105 w; movable digit 27 uz. Gnathosomal featues generally as in female. Four pairs of gnathosomal setae with c.s. about 45 p apart, hyp.2 about 30 » apart. Deutosternum with 8 denticles arranged in a single file. Pedipalp (1-45-12) with two-tined apotele. Podonotal shield (162-174 w 144-156 ) with ten pairs of setae distributed as in Text-fig. 82D. Pygidial shield (50 » x 100 ») with three pairs of setae, the terminal pair (Z5) being at least four times as long as the other setae on the shield. Un- sclerotized integument of dorsum with 16 pairs of setae. Tritosternum with base 24 p, laciniae 60 w. Sternal shield (120 p x 96 pv) with three pairs of setae and two pairs of pores. Between sti, 63 «; between sfi and s¢3, 75 ». Genital setae very short, about half as long as the sternal and the four pairs of opisthogastric setae. Anal shield (42-45 » x 50-54 pu) with paranal setae about 24 p, postanal seta about 18 » long. Peritreme extends to posterior third of coxa III. Chaetotaxy of legs normal. Length/width (in p) of leg segments : I II III IV genu 40/36 36/36 32/32 40/30 tibia 42/33 83108 30/30 42/30 tarsus 78 /30 57/27 63/27 84/26 HABITAT: A common parasite of poultry and wild birds in Britain, Europe, U.S.S.R. and North America. It has also been recorded from a number of wild birds in South Africa (Zumpt & Till, 1961). Ornithonyssus bacoti (Hirst) Leiognathus bacott Hirst, 1913, Bull. ent. Res. 4: 122 et 1914, Bull. ent. Res. 5: 225, figs. 12-14. Haemogamasus sanguineus Ewing & Stover, 1915, Ent. News 26: 109. Liponyssus tenuiscutatus Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 22. Liponyssus nagayot Yamada, 1930, Proc. eighth Congr. Far East Ass. trop. Med. 2: 258. Female: Chelicera with segment I, 33 »; II, 228 »; movable digit 50 p» long, with an elliptical “‘ trough” in its dorsal surface. Fixed digit slender, terminating in a hyaline boss. Pilus dentilis and arthrodial processes absent (Text-fig. 838). Four pairs of gnathosomal setae with c.s. about 57 » apart, Wyp.2 about 42 p apart. Deutosternum with about ten denticles arranged in a single file. Corniculi mem- branous ; internal malae fused. Tectum capituli tapering to a point. Pedipalp (2-5-6-13) with two-tined apotele ; palptrochanter with an antero-ventral keel. Dorsal shield (660 » x 198 p») with 18 pairs of setae distributed as in Text-fig. 834. Unslcerotized integument of dorsum hypertrichous. Tritosternum with hyaline denticulate membrane; base 66 p, laciniae 108 p. Sternal shield (50 » X 120 y) reticulated, with three pairs of setae and two pairs of pores ; metasternals free. Between sti, 70 »; between sf1 and st3, 66 pw. Genital shield (174-180 » X 70-75 p) tapering, with genital setae only. Anal shield (123- 135 » X 78-87 u) with paranal setae 42 pu, postanal seta 45 wlong. Chaetotaxy and EN EVANS & W. M. TILL Seen THE BRITISH DERMANYSSIDAE (ACARI) 323 sclerotization of opisthogaster as in Text-fig. 83p. Peritreme extends to posterior margin of coxa I ; peritrematal shield free anteriorly, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs normal except genu III (2—?, ?—2), tibia III (2—+, ?—2) and genu IV (2—?, 3—1). Coxa II with small anterior spine. Length/width (in ,) of leg segments : I 10 Ill IV genu 96/40 66/45 60/42 84/38 tibia 96/33 66/42 60/40 84/33 tarsus 162/33 120/36 126/34 168/33 Male: Chelicera with segment I, 27 »; II, 120 uw; movable digit 36 uw; both digits edentate. Spermadactyl relatively short, 45 » from base of movable digit (Text-fig. 838). Dorsal seta minute, pilus dentilis and arthrodial processes absent. Four pairs of gnathosomal setae with c.s. about 54 » apart, hyp.2 about 38 p apart. Corniculi membranous ; internal malae blade-like, free. Pedipalp as in female, but palptrochanter without ventral keel; seta f/ on palpfemur situated on a small tubercle. Dorsal shield (528 » 270 p) with 21 pairs of setae distributed as in Text-fig. 83¢. Unsclerotized integument of dorsum much less densely setose than in the female. Tritosternum with base 36 p, laciniae 75 ». Holoventral shield (444 # x 100 p) narrow, with eight pairs of setae excluding the anals. Paranal setae 30 p, postanal seta 33 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 83F. Peritremes extend to posterior margin of coxae I; peritrematal shields fused an- teriorly with dorsal shield and posteriorly with podal elements of coxae IV. Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : I II III IV genu 75/40 60/48 60/45 72/36 tibia 78/33 54/48 54/42 75/36 tarsus 130/30 80/40 87/34 120/30 Larva: Chelicera with segment I, 10 »; II, 63 » ; movable digit 8 ~ ; both digits simple and edentate. Two pairs of gnathosomal setae (yp.1 and 2) with hyp.2 about 24 m apart. Deutosternal teeth not visible. Corniculi degenerate. Pedipalp (0-3/4-5-10) with two-tined apotele ; palptrochanter without keel. Dorsum with no visible sclerotization ; with eight pairs of setae in the podonotal region and three pairs in the opisthonotal region. Tritosternum bipartite, laciniae strongly reduced, without visible denticulate membrane ; base 36 p, laciniae 24 ». Sternal region with three pairs of setae, but Fic. 83. Ovrnithonyssus bacoti (Hirst). a. Dorsum of idiosoma of female; B. chelicera of male; c. dorsum of idiosoma of male; pb. venter of idiosoma of female; E. chelicera of female ; F. venter of idiosoma of male. 324 G. OWEN EVANS & W.M. TILL no visible sclerotization. Opisthogaster with two pairs of setae (Jv and Jv2) ; three anal setae present, but no definite shield. Stigma and peritreme absent. Chaetotaxy © of legs as in free-living Dermanyssidae. Protonymph : Chelicera with segment I, 24 »; II, 135 »; movable digit 33 xz. Gnathosomal features generally as in female. Four pairs of gnathosomal setae with c.s. about 36 » apart, hyp.2 about 27 » apart. Deutosternum with nine denticles arranged in a single file. Pedipalp (1-4-5-12) with two-tined apotele, no keel on palptrochanter. Podonotal shield (228 » x 174 pw) with ten pairs of setae as in Text-fig. 844; pygidial shield (75 » < 123 ») with four pairs of setae. Unsclerotized integument of dorsum with 16 pairs of setae. Tritosternum bipartite, with hyaline denticulate membrane ; base 27 py, laciniae 75 pw. Sternal shield (123 » x 90 p) with three pairs of setae and two pairs of pores. Between sti, 50 »; between sti and st3, 90 pw. A pair of short genital setae is situated posterior to the sternal shield. Anal shield (60 » x 42 ») with paranal setae 30 »; postanal seta 36 » long. Opisthogastric cuticle with four pairs of setae, Jvi-2, Jv5 and Zv2 (Text-fig. 848). Peritreme extends to anterior margin of coxa UOe Chaetotaxy of legs normal. Length/width (in ,) of leg segments : I II III IV genu 48/33 42/39 33/33 45/33 tibia 54/30 40/36 33/33 48/30 tarsus 93/27 60/30 60/27 84/27 Deutonymph : Male and female deutonymphs have been distinguished which differ chiefly in their setation, the female being more densely hypertrichous than the male. In both forms the chelicerae have simple edentate digits as in the larva. Deutoster- num with ten transverse rows of denticles (one to three denticles per row) ; internal malae not fused ; pedipalp (2-5—6—13) with two-tined apotele. Dorsum very weakly sclerotized, so that distinct shields are not recognizable. Male with 15 pairs of setae in the podonotal and 22 pairs in the opisthonotal region ; female with 23 pairs of setae in the podonotal region, opisthonotal region markedly hypertrichous. Tritosternum simple, bipartite only at its tip. Area between the coxae with three pairs of relatively long sternal setae, a pair of short metasternal and a pair of short genital setae. The margins of a shield are weakly defined in the male (Text-fig. 84¢), but not visible in the female (Text-fig. 84D). Opisthogastric cuticle with 13 pairs of setae in the male, markedly hypertrichous in the female. Par- and postanal setae subequal in length. Peritremes extend to anterior margin of coxae III. Fic. 84. Ornithonyssus bacoti (Hirst). A. Dorsum of idiosoma of protonymph ; B. venter of idiosoma of protonymph ; c. venter of idiosoma of deutonymph (male) ; D. venter of idiosoma of deutonymph (female). THE BRITISH DERMANYSSIDAE (ACARI) 325 oe A Woe anve aN 326 G: OWEN EVANS & W. M. TIEL Chaetotaxy of legs as in female. Hasitat: O. bacoti, the tropical rat mite, is apparently of recent introduction to Britain where its distribution is centred mainly around ports. It is essentially a parasite of rodents, especially Rattus vattus (Linn.), but is also known to bite man (Evans et al., 1961). Ornithonyssus pipistrelli (Qudemans) Liponyssus muscult (Koch) Oudemans, 1902, Tijdschr. Ned. dierk Vereen. (2) 8:17, pl. I, figs. I-2. Liponyssus pipistrelli Oudemans, 1903, Tijdschr. Ent. 46: 111 [nom. nov. pro Liponyssus musculi (Koch) Oudemans, 1902]. Female : Chelicera with segment I, 54 » ; II, 174 »; movable digit 50 p, edentate with dorsal “trough”; fixed digit with terminal hyaline boss. Four pairs of gnathosomal setae with c.s. about 66 p» apart, hyp.2 about 42 wapart. Deutosternum with nine denticles arranged in a single file. Corniculi membranous ; internal malae with inner processes fused, outer large and leaf-like. Tectum capituli taper- ing, with serrated margin. Pedipalp (2—-5-6-13) with two-tined apotele ; palp- trochanter with ventral keel. Dorsal shield (912 » x 294 ») narrow, reticulated, with 20 pairs of setae distributed as in Text-fig. 854. Unsclerotized integument of dorsum hypertrichous. Tritosternum with denticulate hyaline border ; base 54 p, laciniae 110 p. Sternal shield (75 » X 144 p) reticulated, with three pairs of setae and two pairs of pores ; metasternals free with associated pores. Between sti, 87 »; between sf1 and s/3, 93 p. Genital shield (180 » x go p) tapering, with genital setae only. Anal shield (75 » X 63 ») with paranal setae 18 p, postanal seta 27 ». Integument of opistho- gaster hypertrichous (Text-fig. 85c). Peritreme extends to middle of coxa II. Anterior part of peritrematal shield extends to middle of coxa I, ending freely ; posterior part fused with podal elements of coxa IV. Chaetotaxy of legs normal except genu III (2—?, 2?—2), tibia III (2—}, }—2) and genu IV (2—#, ?—1). Coxa II with prominent anterior spine. Length/width (in 4) of leg segments : I II Ill IV genu 144/54 117/66 114/66 150/60 tibia 144/51 114/63 114/60 150/54 tarsus 258/48 192/54 174/48 240/54 Male: Chelicera with segment I, 36 »; IJ, 126 »; movable digit edentate, 42 4 long ; spermadactyl relatively short, 54 , from base of movable digit. Four pairs of gnathosomal setae with c.s. about 51 » apart, Ayp.2 about 39 mw apart. Deuto- sternum with ten denticles arranged in a single file. Corniculi membranous, internal malae leaf-like. Tectum capituli and pedipalps as in female. — Fic. 85. Ornithonyssus pipistrelli (OQudemans). A. Dorsal shield of female ; 8. dorsum of idiosoma of male; c. venter of idiosoma of female; pb. venter of idiosoma of male 327 THE BRITISH DERMANYSSIDAE (ACARI) 328 G. OWEN EVANS & W.M. TILL Dorsal shield (696 4 x 360 ») considerably broader than in the female, covering almost the entire dorsum (Text-fig. 858) ; bearing about 25 pairs of setae. Surface - of shield strongly reticulated ; unsclerotized integument of dorsum with about 12 pairs of setae. Tritosternum with base 30 yp, laciniae 78 ». Holoventral shield (588 » x 123 p) narrow, reticulated, with 19 setae excluding the anals. Between sf1, 70 » ; between sti and sf3, 100 ». Paranal setae about 33 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 85D. Peritreme extends to anterior margin of coxa III; anterior part of peritrematal shield fused with dorsal shield, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs asin female. Coxa II with prominent anterior spine. Anterior setae on coxae II and III large and spine-like. Length/width (in p) of leg segments : I II III IV genu 120/48 96/60 99/57. 132/57 tibia 126/48 84/57 93/54 126/54 tarsus 204/42 126/48 126/45 192/45 Fic. 86. Ornithonyssus pipistrelli (OQudemans), protonymph ; dorsum (A) and venter (8) of idiosoma. | THE BRITISH DERMANYSSIDAE (ACARI) 329 Protonymph :; Chelicera with segment I, 30 »; II, 99 »; movable digit 30 up. Gnathosomal features essentially as in female. Four pairs of gnathosomal setae with c.s. about 42 mw apart, Ayp.2 about 30 mw apart. Deutosternum with nine denticles arranged in a single file. Pedipalp (1-4-5-12) with two-tined apotele ; palptrochanter with small ventral keel. Podonotal shield (210 » X 204 ») with 11 pairs of setae (Text-fig. 864) ; pygidial shield (66 » < 180 p») very broad, with six pairs of setae. Unsclerotized integument of dorsum with about 15 pairs of setae. Tritosternum with base 36 p, laciniae 45 ww. Sternal shield (150 » x 117 pw) with three pairs of setae and two pairs of pores. Between sf1, 66 » ; between séi and s¢3, 96 pw. Anal shield (60-63 » x 66-69 ») with paranal setae about 30 yp, postanal seta about 36 plong. Opisthogaster with a pair of genital and five pairs of opisthogastric setae distributed as in Text-fig. 868. Peritreme extends to posterior third of coxa II. Chaetotaxy of legs normal. Length / width(in w) of leg segments : I iu III nV, genu 63/39 54/45 48/40 60/36 tibia 63/36 54/42 51/38 60/33 tarsus T7383 87/33 84/32 110/30 HapitaT: Found in Dorset, England, on Eptesicus serotinus (Schreber) and Plecotus auritus (Linn.). _ Oudemans described the species from ‘‘ Vespertilio pipi- strellus’’, but does not give any locality. Lange (1959) describes a closely related species (O. pavlovskit) from Muscardinus avellanarius (Linn.) in Rumania. The number and relative positions of the setae on the dorsal shield of the female appear to be identical with the condition in O. pipistrellz. Genus STEATONYSSUS Kolenati Steatonyssus Kolenati, 1858, Wien. ent. Monatschy. 2:6; et 1859, S.B. Akad. Wiss. Wien. 35: 186. Till & Evans, 1964, Bull. Brit. Mus. (nat. Hist.) Zool. 11: 513. Ceratonyssus Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 6. Type: Steatonyssus periblepharus Kolenati, 1858. Cheliceral digits edentate, lacking denticulate processes ; fixed digit with distal hyaline boss. Movable digit of male not hooked distally ; spermadactyl grooved, relatively short, a membranous lobe present between spermadactyl and digit. Pilus dentilis absent, dorsal seta minute or absent, fissures normal. Arthrodial processes absent. Chaetotaxy of venter of gnathosoma normal; pedipalps with five free segments and a two-tined apotele ; chaetotaxy of trochanter to tibia (2—-5—6-13). Palptrochanter with well-developed keel in the female. Deutosternum with six to ten denticles arranged in a single file. Corniculi membranous, internal malae simple. Tectum capituli elongate, apex fimbriated. 330 G. OWEN EVANS & W. M. TILL Dorsal sclerotization of the female comprising well-developed podonotal and opisthonotal shields. Podonotal shield with ten (73 lacking), 11 or 13 pairs of setae, paraverticals (z1) absent. Opisthonotal shield normally with seven pairs of setae, but this number may be reduced by the absence of J5 or J/5 and Z5. Unpaired accessory setae sometimes present in the / region. Lateral integument hyper- trichous. Male with entire dorsal shield ; podonotal region with II or 12 pairs of setae ; opisthonotal region with similar chaetotaxy to the opisthonotal shield of the female, but with a tendency for the addition of Z and S setae to the shield. Tritosternum normal, bipartite, with hyaline denticulate border. Sternal shield with three pairs of setae and two pairs of pores ; posterior margin often with a more strongly sclerotized band. Genital shield tapering posteriorly, with genital setae only. Anal shield normal. Opisthogastric cuticle usually shows hypertrichy. Metapodal shields weak. Peritrematal shields fused with podal shields of coxae IV. Male with holoventral shield. Leg chaetotaxy typically—teg I normal, leg II normal, leg III normal except genu I ?—2) and tibia III (2—}, #—2) and leg IV normal except genu IV (2— Other exceptions occurring rarely are tibia I (2—3, {—2), tibia III (2— —2) and tibia IV (2—?, #—2). Coxa II with strong anterior spine in both sexes. Protonymph with dorsum of idiosoma bearing 28 ( : lacking) or 29 pairs of setae. Podonotal shield with Io or 11 pairs of setae, pygidial shield normally with four pairs. According to chaetotactic patterns of certain adults whose protonymphs have not been found, it is possible that the number of dorsal setae may be reduced to 26 (72, J5 and Z5 lacking) or 28 (/5 missing). Venter of idiosoma with three pairs of sternal setae, one pair of genital setae and five pairs of opisthogastric setae. Anal shield normal. Gnathosoma as in female, but palptrochanter lacking ventral keel. Chaetotaxy of palps and legs normal. Si Ibo RIO TS I | iS eiuele KEY TO FEMALES rt Sternal setae 1 very short, not more than one-half the length of sternal setae 2 (Text- fig. 87B) ; peritrematal shield entire anteriorly S. periblepharus Kolenati (p. 330) — Sternal setae 1 and 2 long, subequal in length ; anterior portion of peritrematal shield interrupted. ‘ : : ; ; S. occidentalis Ewing (p. 334) Steatonyssus (Steatonyssus) periblepharus Wolenati Steatonyssus periblepharus WKolenati, 1858, Wien. ent. Monatschy.2:6; et 1850, S.B. Akad. Wiss. Wien 35: 186, pl. 8, fig. 34. Till & Evans, 1964, Bull. Brit. Mus. (nat. Hist.) Zool. 11: 562, figs. 62-67. Female; Chelicera with segment I, 40 w; II, 168 »; movable digit 54 »; both digits edentate. Four pairs of gnathosomal setae with c.s. about 45 « apart, /yp.2 about 33 p» apart. Deutosternum with eight denticles arranged in a single file. Corniculi membranous ; internal malae simple. Tectum capituli tapering, with anterior margin fimbriated. Pedipalp (2-5-6-13) with two-tined apotele ; palp- trochanter with prominent antero-ventral keel. — ae THE BRITISH DERMANYSSIDAE (ACART) 331 Fic. 87. Steatonyssus periblepharus Kolenati. A. Dorsum of idiosoma of female; B. venter of idiosoma of female ; c. dorsum of idiosoma of male; bD. venter of idiosoma of male. ZOOL. 14, 5. 22 332 G. OWEN EVANS & W. M. TILL Podonotal shield (294-312 » * 252-276 p) bears II pairs of setae distributed as in Text-fig. 874. Opisthonotal shield (348-396 » x 180-198 yw) with seven pairs of setae. Seta JI relatively stouter than the other setae on the shield. Surface of both shields reticulated ; unsclerotized integument of dorsum with numerous rather stout, spine-like setae. Tritosternum with hyaline denticulated mambrane ; base 38 yp laciniae 102 yz. Sternal shield (45-57 » < 130-144 yw) reticulated, not sharply demarkated from presternal area ; posterior border of shield heavily sclerotized ; bearing three pairs of setae and two pairs of pores; metasternal setae free. Between sti, 45-50 yp; between sf1 and st3, 70-72 pw. Sternal seta I relatively short, not more than half as long as st2. Genital shield (144-156 » x 75-93 p») tapering, with genital setae only. Anal shield (114-135 » < 80-94 ») with par- and postanal setae subequal, about 40 u long. Integument of opisthogaster bears numerous setae, those near the margin being relatively longer (65 ») and stouter than the others (Text-fig. 878). Peritreme extends to middle of coxa II ; anterior part of peritrematal shield continues to middle of coxa I; posterior part fused with podal Smelioks of coxa IV. Chaetotaxy of legs normal except genu III (2—7, #—2), tibia III (2+, 2—2) and genu IV (2—#?, $—1). Coxa II with prominent anterior spine. Length/width (in p) of leg segments : I i Ill IV genu 102/54 93/69 84/60 = 108/54 tibia 102/48 84/60 78/54 102/48 tarsus 177/42 156/48 156/48 192/48 Male: Chelicera with segment I, 27 »; II, 99 »; movable digit 34 » ; sperma- dactyl relatively short, grooved, 45 » long from base of movable digit ; a mem- branous lobe is present between the spermadactyl and the digit. Fixed digit edentate with distal hyaline boss. Four pairs of gnathosomal setae with c.s. about 36 mw apart, Ayp.2 about 36 w apart. Pedipalp trochanter without ventral keel ; other features as in female. Dorsal shield (504 » x 228 ») with 19 pairs of setae distributed as in Text-fig. 87c. Surface of shield reticulated. Unsclerotized integument of dorsum hy pertrichon Tritosternum with base 21 p, laciniae 66 pz. Holoventeal shield (420 ~ X 105 p) narrow, bearing six pairs of setae excluding the anals ; metasternal setae on integu- ment beside shield. Between sf1, 48 «: between sf1 and s#3, 75 ». Par- and post- anal setae subequal, about 24 » long. Chaetotaxy and sclerotization of opistho- gaster as in Text-fig. 87p. Peritreme extends to anterior margin of coxa III; anterior part of peritrematal shield short, free; posterior part fused with podal elements of coxa IV. Chaetotaxy of legs as in female. Anterior seta on coxa III short, spinose, with blade-like extension on proximo-anterior margin. Length/width (in x) of leg | segments : THE BRITISH DERMANYSSIDAE (ACARI) 333 I II III IV genu 75/38 66/48 50/45 72/40 tibia 75,34 54/42 50/40 69/33 tarsus 130/32 go /36 90/33 120/30 Protonymph : Chelicera with segment I, 20 »; II, 100 w; movable digit 30 up. Four pairs of gnathosomal setae with c.s. about 33 w apart ; hyp.2 about 27 p apart. Pedipalp (I-4-5—12) without keel on trochanter. Podonotal shield (180 » X 150 yp) with 11 pairs of setae as in Text-fig. 88a ; pygidial shield with four pairs of setae ; unsclerotized integument of dorsum with 14 pairs of setae. Tritosternum with base 15 p, laciniae 60 pw. Sternal shield (140 » X 114 p) with three pairs of setae and two pairs of pores. Between sf1, 60 »; between sfi and s¢3, 96 w. Analshield (48 » x 54-57 ») with par- and postanal setae subequal, about 27 u long. Integument of venter bears a pair of genital and five pairs of opisthogastric setae (Text-fig. 88B). Peritreme extends nearly to middle of coxa III. Fic. 88. Steatonyssus periblepharus Kolenati, protonymph ; dorsum (A) and venter (B) of idiosoma. G. OWEN EVANS & W. M. TILL 334 Chaetotaxy of legs normal. Length/width (in y) of leg segments : I iB EE IV genu 48/40 45/42 36/39 48/36 tibia 45/34 36/39 36/36 48/33 tarsus 90/30 66/30 66/32 102/30 Hapitat: Found in Britain on Prpistrellus pipistrellus (Schreber) and Myotis mystacinus (Kuhl). Recorded from Europe, North Africa and the Near East, especially on bats of the genera Prpistrellus and Myotis. Steatonyssus (Steatonyssus) occidentalis (Ewing) Ceratonyssus occidentalis Ewing, 1933, Proc. U.S. nat. Mus. 82 (art. 30) : 10, figs. Steatonyssus occidentalis : Radovsky & Furman, 1963, Ann. ent. Soc. Amer. 56 : 272, figs. Till & Evans, 1964, Bull. Brit. Mus. (nat. Hist.) Zool. 11: 5509, figs. Female: Chelicera with segment I, 27 »; II, 140 »; movable digit 40 »; both digits edentate. Four pairs of gnathosomal setae with c.s. about 45 » apart ; hyp.2 about 36 » apart. Deutosternum with six denticles arranged in a single file. Cor- Tectum capituli tapering, anterior palp- niculi membranous ; internal malae simple. margin weakly serrated. Pedipalp (2-5-6-13) with two-tined apotele ; trochanter with prominent antero-ventral keel. Podonotal shield (288 » x 282 ») with 11 pairs of setae distributed as in Text-fig. 89A ; opisthonotal shield (372 » x 204 ») with seven pairs of setae. Surface of both shields reticulated, opisthonotal shield with striations posteriorly. Unsclerotized integument of dorsum hypertrichous. Tritosternum with hyaline, denticulated membrane. Sternal shield (40 » X 132 p) reticulated, with thickened posterior border ; bearing three pairs of setae and two pairs of pores; metasternal setae free. Between sf1, 68 «; between sf1 and s¢3, 72 p. Genital shield (144 » X 93 ») tapering, with genital setae only. Anal shield (120 » X 96 ») with par- and postanal setae about 40 » long. Integument of opisthogaster hypertrichous (Text-fig. 898). Peritreme extends to anterior margin of coxa III; anterior part of peritrematal shield interrupted, a separate leaf-like portion lying over coxae I-II ; posterior part of peritrematal shield fused with podal elements of coxa IV. Chaetotaxy oulegs normal except genu III (2— 2), tibia III (2—+, ?—2) and 4, Cpl genu IV (2—#?, ?—1). Coxa II with anterior spine. Length/width (in ,) of leg segments : I Il Ill I\ genu 96/57 84/75 72/60 = 102/54 tibia 96/45 78/60 75/52 102/48 tarsus 162/40 150/45 147/42 198/42 THE BRITISH DERMANYSSIDAE (ACARI) 235 ( = ~ Co en a ae ek ae te ) 1 | | oe \ | / \ \ | ; \ Fic. 89. Steatonyssus occidentalis (Ewing), female ; dorsal shields (A) ; ventral shields (B). Male and Protonymph ; Described by Radovsky & Furman (1963) and by Till & Evans (1964). HABITAT: Found on Eptesicus serotinus (Schreber), Wickham Market, Suffolk, 24th June, 1964 (coll. Lord Cranbrook). This is the first record of S. (S.) occidentalis from Britain. Previously the species was known only from North America, where it Hy occurs on bats of the genera Eptesicus, Myotis, Tadarida, Corynorhinus and Lasiurus. Genus OPHIONYSSUS Meégnin Hi i Ophionyssus Mégnin, 1884, C.R. Soc. Biol. Paris 35 : 614. i} Serpenticola Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 6. Nal Sauronyssus Sambon, 1928, Ann. trop. Med. 22 : 106. Neoliponyssus Ewing, 1929, Manual of External Parasites : 187. Hy Oudemansiella Fonseca, 1948, Proc. zool. Soc. Lond. 118 : 269. Type: Dermanyssus natricis Gervais, 1844. 3360 G. OWEN EVANS & W.M. TILL Cheliceral digits edentate, movable digit in the female with a well-defined hyaline flap; fixed digit grooved, rounded anteriorly. Movable digit of male chelicera - completely fused with the deeply grooved spermadactyl ; fixed digit not hooked. Dorsal seta and pilus dentilis absent ; arthrodial membrane at base of chelicera without setiform processes. Chaetotaxy of venter of gnathosoma normal, pedipalps with five free segments and a two-tined apotele. Chaetotaxy of pedipalps (2-4/5- 5/6-13). Palptrochanter with a shallow, inconspicuous, weakly sclerotized keel. Deutosternum with 6-9 denticles in a single longitudinal row. Corniculi membran- ous, typically macronyssine. Tectum capituli elongate, obtuse, anterior margin fimbriated. Dorsal sclerotization of the female of three basic types; one consisting of a podonotal shield bearing nine or ten pairs of setae, two pairs of mesonotal scutellae and a small sub-circular pygidial shield normally lacking setae ; the second with a posterior extension of the podonotal shield incorporating opisthonotal setae /I and Jj2, and with a discrete pygidial shield lacking setae ; the third consisting of a single dorsal shield incorporating Io or II pairs of setae in the podonotal region and three pairs of J setae in the opisthonotal region, with setae 75 on or off the shield. In the male the dorsal shield is entire, bearing 17-24 pairs of setae. Paravertical setae (z1) absent. Tritosternum bipartite, with a denticulate hyaline border. Sternal shield in female with two pairs of setae and two pairs of pores, the third pair of sternal setae and the metasternals being situated on unsclerotized integument. Genital shield narrow, tapering, the genital setae placed on the integument. Anal shield pear- shaped with normal chaetotaxy. Unsclerotized integument of opisthogaster hypertrichous. Peritrematal shields fused posteriorly with podal shields of coxae IV. Metapodal and podal shields weakly developed. Male with a sternito-genital shield bearing two or three pairs of setae ; anal shield free. Chaetotaxy of legs shows several deviations from the normal pattern: femur I (2—3—2), genu I and tibia I (2—#, 7—2) or (2—?, 7—2), femur II (1—3—1), 2__2 genu II (2—?, 17); Hb II (2—}4, 12), oa III (r—%—»), genu III (2—4, ¢—2), tibia III (2—}, 3-2), genu IV (2—7, }-—2) or (2—4, $2) and tibia IV 2—z, #2). Coxa Tl Sous an oe spine. All ambulacra with a pair of Protonymph with general gnathosomal characteristics of the female. Podonotal region with 16 pairs of setae (podonotal shield with ten pairs) ; opisthonotal region with ten pairs (pygidial shield with two pairs, /4 and Z5) or It pairs (pygidial shield with three pairs, J4, S5 and Z5) of setae. Sternal shield with three pairs of setae and two pairs of pores, genital setae present. Opisthogastric cuticle with four pairs of setae. Leg chaetotaxy normal except for genu IV (1—3, 2—1). Strandtmann & Wharton (1958) recognize Sauvonyssus as a distinct genus differing from Ophionyssus in the nature of the idiosomal sclerotization of the female. This appears to be the only distinguishing feature between them. The exclusion of setae Z5 from the entire dorsal shield of certain Sauvonyssus suggests that a reduction of the pygidial shield also occurs between the protonymphal and adult female stages of THE BRITISH DERMANYSSIDAE (ACARI) 337 these species and that the elongate dorsal shield may be formed by the posterior prolongation of the podonotal shield to incorporate setae J1, J/3 and /4. In the males of both Ophionyssus and Sauronyssus on the other hand there appears to be no reduction of the pygidial shield which forms (with its protonymphal complement of setae) part of the dorsal shield. The degree of specialization of the idiosomal sclero- tization exhibited by the Ophionyssus-Sauronyssus complex is no greater than that occurring in certain other genera of highly specialized macronyssines and dermanys- sines, for example, compare Dermanyssus gallinae and D. triscutatus Krantz, and Ormthonyssus bacott and O. aridus Furman and Radovsky. In view of this we consider Sauronyssus to be synonymous with Ophionyssus. The distinction based on the host preferences of Ophionyssus (snakes) and Sauronyssus (lizards) is not valid. Ophionyssus monodi (Hirst) recorded from Acanthodactylus scutellatus and Ophionys- sus mabuyae Till from the skink Mabuya striata have the idiosomal sclerotization of the natricis type. In fact, the dorsal sclerotization of O. mabuyae is intermediate between the conditions in O. natricis and O. saurarum, having a short posterior extension of the podonotal shield which incorporates two pairs of J setae, but retain- ing a small pygidial shield. KEY TO FEMALES 1 Dorsal sclerotization consisting of a podonotal shield, two pairs of mesonotal scutellae and a pygidial shield. : : : O. natricis (Gervais) (p. 337) Dorsal sclerotization consisting of a single dorsal shield . : ° 2 2 Tibia IV with two postero-dorsal setae (2—+, 2—2). Dorsal shield inte ten pairs of setae in the podonotal region . : : E O. saurarum (Oudemans) (p. 342) — Tibia IV with three postero-dorsal setae (2—+, $—2). Dorsal shield with rr pairs of setae in the podonotal region . ‘ : : O. lacertinus (Berlese) (p. 345) KrEy To MALES I Dorsal shield with 23-24 pairs of setae. Femur III with ventral seta greatly enlarged, spur-like : ' O. lacertinus (Berlese) (p. 346) — Dorsal shield with 16-18 pairs of setae. F Ces III with ventralsetanormal . : 2 2 Tibia IV with three postero-dorsal setae (2—+, 3—2) . O. natricis (Gervais) (p. 340) Tibia IV with two postero-dorsal setae (2-1, 2—2) O. saurarum (Oudemans) (p. 342) Ophionyssus natricis (Gervais) Dermanyssus natricis Gervais, 1844, Hist. Nat. Ins. Apt. 3 : 223. Ophionyssus natricis : Camin, 1953, Chicago Acad. Sci. no. 10: 1, figs. Bregetova, 1956, Opred. Faune. SSSR. 61 : 160, 223, figs. 556-562. Ichoronyssus serpentium Hirst, 1915, Proc. zool. Soc. Lond. : 383. ?Serpenticola easti Ewing, 1925, Ent. News 36: 18 Female ; Chelicera with segment I, 30 »; II, 165 »; movable digit 45 ». Both digits edentate, pilus dentilis and dorsal seta absent (Text-fig. g1c). Four pairs of gnathosomal setae with c.s. about 60 pw apart, Wyp.2 about 42 wapart. Deutosternum with eight or nine denticles arranged in a single file. Corniculi membranous, internal malae slender, elongated (Text-fig. gta). Tectum capituli with anterior margin G. OWEN EVANS & W. M. TILL V Za ph GPF (A ZZ 2 \ « 2 5 \ \ ‘ a \ V 4 \ Fic. 90. Ophionyssus natricis (Gervais). A. Dorsum of idiosoma of female; B. venter of idiosoma of female; c. dorsum of idiosoma of male; bp. venter of idiosoma of male. THE BRITISH DERMANYSSIDAE (ACART) 339 fimbriated. Pedipalp (2-5-6-13) with two-tined apotele ; palptrochanter with a shallow, inconspicuous, weakly sclerotized keel. Podonotal shield (300 » X 276 w) with ten pairs of setae distributed as in Text-fig. goa ; two pairs of mesonotal scutellae le posterior to the podonotal shield. Surface of podonotal shield granular, weakly reticulated antero-laterally. Pygidial shield (54 » X 45 ») without setae. Unsclerotized integument of dorsum shows marked hypertrichy. Tritosternum with hyaline membrane ; base 54 p, laciniae 96 pp. Sternal shield (45 # X 114 ) granular, with two pairs of setae and two pairs of pores ; the third pair of setae and the metasternals are situated on the striated integument. Between stI, 42 »; between sti and s#3, 102 w. Genital shield (about 135 w x 45 yp) slender and tapering with the genital setae on the integument beside the shield. Anal shield (78 » X 72 ») with paranal setae 33 » postanal seta 36 w long. Sclerotization and chaetotaxy of venter as in Text-fig. goB. Peritreme extends to posterior margin of coxa II; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs shows deviations from the normal pattern on most segments : femur I (2—3—2), femur II (1—3—1) femur III (1—?—o) ; genu I (2—3, #—), genu III (2—?, 7—2), genu IV (2—?, 2—2) ; tibia I (2—3, ?—2), tibia III (2—4, 22). Coxa II without anterior spine. Length/width (in ») of leg segments : A Fic. 91. Ophionyssus natricis (Gervais). A. Venter of gnathosoma of female ; B. chelicera of male; c. chelicera of female. G. OWEN EVANS & W.M. TILL 340 I II III IV genu 75,45 60/48 60/45 72/40 tibia 75/40 60/45 54/40 66/36 tarsus 123/36 96/40 90/36 I10/33 Male ; Chelicera with segment I, 18 »; II, 96 ». Fixed digit edentate ; movable digit completely fused with grooved spermadactyl (Text-fig. 91B). Four pairs of gnathosomal setae with c.s. about 42 » apart, hyp.2 about 36 w apart. Other features generally as in female. Dorsal shield (492 » X 240 ») with 17 pairs of setae distributed as in Text-fig. goc. Surface of shield mainly granular with reticulations antero-laterally. Tritosternum with hyaline membrane ; base 20 p, laciniae 70 p. Sternito-genital shield (204 » x 81 ») with sfx and st2 on the shield, st#3, st4 and the genital setae on the cuticle lateral to the shield. Between sf1, 51 » ; between sti and s¢3, 84 ». Anal shield (66 » Xx 60 ») with par- and postanal setae about 27 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. gop. Peritreme extends to anterior margin of coxa III ; peritrematal shield fused posteriorly with podal shield of coxa IV. Chaetotaxy of legs as in female. Length/width (in p) of leg segments : I I Ul IV genu 60/36 51/39 48/36 60/33 tibia 60/33 51/38 48/33 60/33 tarsus 102/33 75/30 75/30 93/30 Protonymph ; Chelicera with segment I, 15 »; II, 110 ~%; movable digit 28 u. Four pairs of gnathosomal setae with c.s. about 39 apart, hyp.2 about 27 pu apart. Deutosternum with nine denticles arranged in a single file. Pedipalp (1-4~-5-12) with two-tined apotele. Other gnathosomal features as in female. Podonotal shield (204 » 192 ») with eleven pairs of setae distributed as in Text- fig. g2A ; setae s6 and 72, 73, 75 and 76 are situated on the unsclerotized integument. Pygidial shield (63 » 75 ») bears three pairs of setae. Tritosternum with hyaline border ; base 20 p, laciniae 75 ». Sternal shield (110 « x 78 pu) with three pairs of setae and two pairs of pores. Between sf1, 48 » ; between st1 and st3, 80 pw. Anal shield (48 » x 48 ») with paranal setae 2I », postanal seta 24 p long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 92B. Peritremes extend to anterior margin of coxa III. Chaetotaxy of legs differs from the normal pattern in the presence of a postero- lateral seta on genu IV (1—2, 2—1). Length/width (in yz) of leg segments : i Il Ill IV genu 42/36 2/36 40/33 48/30 tibia 48/33 2/36 36/30 48/30 ? tarsus 84/30 60/30 60/27 66 THE BRITISH DERMANYSSIDAE (ACARI) 341 Fic. 92. a-B. Ophionyssus natricis (Gervais), protonymph ; dorsum (A) and venter (B) of idiosoma. c—D. Ophionyssus sauravum (Oudemans), protonymph; dorsum (c) and venter (D) of idiosoma. 342 G. OWEN EVANS?& WM. bE Hasitat: A common ectoparasite of snakes and lizards in vivaria all over the world. It has also been found in natural conditions on a wide variety of snakes in . Africa. Ophionyssus saurarum (Oudemans) Liponyssus sauvarum Oudemans, 1901, Tijdschr. Ned. dierk. Vereen. (2) 7: 66. Sauronyssus sauvavum: Bregetova, 1956, Opred. Faune SSSR. 61 : 218, figs. Female: Chelicera with segment I, 36 »; II, 147 »; movable digit 40 p. Both digits edentate, pilus dentilis and dorsal seta absent.Four pairs of gnathosomal setae with c.s. about 60 » apart, hyp.2 about 42 » apart. Deutosternum with six denticles arranged in a single file. Corniculi membranous ; internal malae elongate. Tectum capituli with anterior margin fimbriated. Pedipalp (2-5-6-13) with two-tined apotele ; palptrochanter with a low ventral keel. Dorsal shield (670 » x 192 p) entire, slender, bearing 13 pairs of setae distributed as in Text-fig. 93A. Surface of shield with a reticulated pattern which tends to become punctate posterior to setae74. Unsclerotized integument of dorsum markedly hypertrichous. Tritosternum with denticulate, hyaline border; base 45 p, laciniae 78 yp long. Sternal shield (42 » 120 ») with two pairs of setae and two pairs of pores ; the third pair of setae and the metasternals are situated on unsclerotized integument. Between stl, 57 #; between sti and s#3, 96 ». Genital shield (about 150 » X 60 ,) tapering ; genital setae on the unsclerotized integument. Anal shield (105 » xX 75 p) with par- and postanal setae about 33 » long. Chaetotaxy and sclerotization of opistho- gaster as in Text-fig. 93B. Peritreme extends to about anterior third of coxa IT; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs shows deviations from the normal pattern on most segments, as in O. natricis. In addition, tibia IV bears only three dorsal setae (2—}, {—2). Relatively stout setae are present on trochanter I (d), femur I (ad, and fd,) and femur II (ad, and pd,-,)._ Coxa II without anterior spine. Length/width (in pz) oi leg segments : if 1 Ill IV genu 102/45 80/48 78/40 93/40 tibia 100/42 75/48 75/40 90/36 tarsus 180/36 110/40 120/36 120/36 Male: Chelicera with segment I, 20 »; II, 93 »; fixed digit edentate ; movable digit completely fused with short, grooved spermadactyl. Four pairs of gnathosomal setae with c.s. about 42 » apart, hyp.2 about 39 wapart. Other gnathosomal features as in female. Dorsal shield (528 « x 264 ») with 16 pairs of setae distributed as in Text-fig. 93C ; one or more additional unpaired setae may be situated on the margins of the podo- or 343 THE BRITISH DERMANYSSIDAE (ACARI) A. Dorsum of idiosoma of female; B sauvarum (Oudemans). ; of female ; c. dorsum of idiosoma of male; bD. venter of idiosoma of oma venter of idios Fic. 93. Ophionyssus male. 344 G. OWEN EVANS & W. M. TILL opisthonotal regions. Surface of shield with a reticulated pattern which has a slight tendency to become punctate near the posterior tip. Unsclerotized integument of dorsum with Io or 12 pairs of setae. Tritosternum with base 24 p, laciniae 48 ». Sternito-genital shield (228 » x 80 p) reticulated, with three pairs of setae and two pairs of pores ; metasternal and genital setae situated on the unsclerotized integument. Anal shield (72 » x 63 p) with paranal setae 36 » long, postanal seta 30 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 93D. Peritreme extends to posterior margin of coxa II ; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs as in female. Relatively stout spine-like setae present on coxae II and III, trochanter I (d), femora I and II (ad, and fd,_,)._ Seta v on femora III and IV is stout basally, tapering distally. Coxa II without anterior spine. Length/width (in ») of leg segments : i II III IV genu 70/40 60/45 60/40 75/40 tibia 70/30 57/42 60/40 75/40 tarsus 120/36 93/36 84/34 110/34 Protonymph: Chelicera with segment I, 20 »; II, 90 »; movable digit 24 pz. Four pairs of gnathosomal setae with c.s. about 38 u apart, Ayp.2 about 27 pu apart. Pedipalp (1-4-5-12) with two-tined apotele. Other gnathosomal features as in female. Podonotal shield (213 » x 180 p) with eleven pairs of setae distributed as in Text- fig.g2c. Surface of shield reticulated. Pygidial shield (84 » x 81 4) bears two pairs of setae. Tritosternum with hyaline border; base 32 yp, laciniae 60 pw. Sternal shield (126 » X 74 ») with three pairs of setae and two pairs of pores ; between sf1, 48 p between sti and st3, 87 ». Anal shield (36 » x 40 ») with par- and postanal setae about 20 pw long. Chaetotaxy of opisthogaster as in Text-fig. 92D. Peritreme extends to anterior margin of coxa III ; peritrematal shields not developed. Chaetotaxy of legs normal, except genu IV which has a postero-lateral seta (1—2, 2—1). Relatively stout setae present on the dorsal surfaces of femora I (ad, and pd,), II (ad, and pd,-,), II (ad,) and IV (ad,-,)._ Length/width (in ,) of leg segments : I II iba IV genu 48/33 42/30 45/33 57/30 tibia 54/33 45/36 45/33 57/32 tarsus 102/30 70/30 72/28 75/27 Hasitat: Recorded from lizards [Lacerta agilis Linn. and L. viridis (Laurenti)] in the Netherlands, Hungary and Italy. In Britain specimens have been obtained from L. agilis in Dorset, and from lizards at Albury, Surrey, and near Aberdeen, Scotland. | THE BRITISH DERMANYSSIDAE (ACART) 345 Ophionyssus lacertinus (Berlese) Leiognathus lacertinus Berlese, 1892, Acari, Myr. Scorp. Ital. fasc. 70, no. 3, figs. Female: Chelicera with segment I, 30 »; II, 180 »; movable digit 50 ». Four pairs of gnathosomal setae with c.s. about 66 p apart, hyp.2 about 38 pm apart. Deutosternum with about seven denticles arranged in a single file. Corniculi membranous ; internal malae elongated. Pedipalp (2-5-6-13) with two-tined apotele ; palptrochanter with a very shallow, inconspicuous keel. Dorsal shield (696 » * 264 ») with at least 14 pairs of setae distributed as in Text- fig.94A. There may be one or two terminal setae, or none, at the posterior tip of the shield. Surface of shield with a reticulate pattern which tends to become punctate. Tritosternum with a hyaline border; base 72 p, laciniae 80 ». Sternal shield (40 » X 120 p) with two pairs of setae and two pairs of pores. Between si1, 54 »; between sf and s#3, 100 w. Genital shield (144 » x 54 ) tapering. Anal shield (106 » x 84 ») with paranal setae about 38 p, postanal seta about 30 » long. Peri- Fic. 94. Ophionyssus lacertinus (Berlese). A. Dorsal shield of female; sB. femur III of male, ventral view ; c. dorsum of idiosoma of male. 340 G. OWEN EVANS & W.M. TILL treme extends to anterior margin of coxa II ; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs as described for O. natricis. Relatively stout setae present on - trochanters I, III and IV (d), femora I-II (ad, and pd,-,), femora III-IV (ad,_,). Coxa II without anterior spine. Length/width (in p) of leg segments : I II Ill IV genu 102/45 84/51 84/48 105/42 tibia 102/40 78/48 78 |42 105/39 tarsus 186/38 123/42 135/36 170/36 Male : Chelicera with segment I, 18 » ; II, 96 » ; fused movable digit and sperma- dactyl about 40 »long. Four pairs of gnathosomal setae with c.s. about 40 pu apart, hyp.2 about 30 papart. Deutosternum with eight denticles arranged in a single file. Other gnathosomal features as in female. Dorsal shield (516 ~ 250 ») broader than in female, with 13 pairs of setae distri- buted as in Text-fig. 94c ; one or more additional unpaired setae may be present on the margin of the opisthonotal region. Unsclerotized integument of dorsum with nine or ten pairs of setae. Tritosternum with base 20 p, laciniae 54 ». Sternito-genital shield (216 up Xx 75 ») reticulated, with s¢z and s¢2 on the shield, s¢3, st4 and genital setae on the unsclerotized integument. Between sii, 45 »; between sf1 and st3, go uw. Anal shield (60 p» x 57 ) with paranal setae 36 p, postanal seta 30 » long. Opisthogaster with about seven pairs of setae. Peritreme extends to posterior margin of coxa II ; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs and setal form as in female. In addition, coxa III bears a relatively stout anterior seta and femur III has a greatly enlarged, spur-like ventral seta (Text-fig. 94B). Length/width (in ») of leg segments : I II III IV genu 66/38 60/40 60/42 70/36 tibia 70/36 57/40 60/40 72/34 tarsus T1732 80/33 96/33 105/30 HaBItTaT: Parasitic on Lacerta muralis (Laurenti) in Italy and on Lacerta viridis _ (Laurenti) imported into Britain from Italy. Subfamily DERMANYSSINAE Kolenati Dermanyssidae Kolenati, 1859, S.B. Akad. Wiss. Wien 35: 171. Adults: Chelicerae of female strongly modified, resembling stylets ; second seg- | Sais : ‘ : sya | ment enormously elongated, slender ; digits minute, weakly dentate. Pilus dentilis, | dorsal seta and arthrodial processes absent. Male chelicera with second segment | THE BRITISH DERMANYSSIDAE (ACARI) 347 normal in length, spermadactyl long and entirely fused with the movable digit ; fixed digit slender, usually shorter than the spermadactyl, rarely equal in length. Chaetotaxy of venter of gnathosoma normal. Pedipalps with five free segments, apotele two-tined ; chaetotaxy of trochanter to tibia (2-5-6-14) in Liponyssoides and some Dermanyssus, but showing deficiency on femur and tibia, rarely genu, in most Dermanyssus. Palptrochanter without medio-ventral keel. Deutosternum with nine or more denticles in a single file. Hypostome compact, elongate ; cor- niculi never horn-like, usually membranous ; internal malae small, simple. Tectum capituli in the form of a triangular lobe. Dorsum of idiosoma of female with reduced sclerotization comprising an entire shield formed by the posterior extension of the podonotal shield ; mesonotal scutellae sometimes present ; small pygidial shield retained in some Liponyssoides. Male with entire dorsal shield. Primary chaetotaxy of the dorsum of the idiosoma showing hypotrichy, paravertical seta (z1) absent, dorsal series (J) often reduced. Hyper- trichy of lateral and marginal series and of the opisthogaster apparent in some species, particularly Liponyssoides. Dorsal setae simple. Tritosternum well developed, biramous, laciniae smooth or pilose ; denticulate hyaline margin present or absent. Sternal shield in the female with two, rarely three, pairs of setae and two pairs of pores; metasternal setae present or absent. Genital shield broadly rounded or tapering posteriorly and bearing one pair of genital setae. Anal shield normal, aciculated area conspicuous ; no euanal setae. Meta- podal shields small, inconspicuous. Peritrematal shield fused with the podal shields posterior to coxae IV ; peritremes variable in length. Podalshields weakly developed. Male with holoventral shield, or sternito-genital and ventro-anal shields, or sternito- genital and anal shields. Genital orifice presternal. All legs six-segmented with well developed ambulacra. Segmental chaetotaxy extremely diverse and often showing intraspecific variation. Tarsi III and IV in male with seta pv, modified into a short, tooth-like projection. Coxae II-IV without sclerotized ridges. Anterior spine of coxa II usually absent, present in some Liponys- soides. Larva: Relatively inactive, non-feeding instar. Chelicerae without elongated second segment ; digits minute, edentate, lacking setae, fissures and arthrodial processes. Pedipalps in Dermanyssus apparently four-segmented (tibia and tarsus fused), chaetotaxy of basal three segments (0-4-5). Hypostome with two pairs of setae ; corniculi and internal malae degenerate ; deutosternal denticles weak or absent. Idiosoma without distinct sclerotization. Podonotal region with seven to nine pairs of short setae, opisthonotal region with variable number of microsetae, J5 always present. Tritosternum biramous, relatively small. Sternal region with three pairs of setae, opisthogastric region with two pairs of setae excluding the normal three setae associated with the non-functional anus. Stigmata and peritremes absent. Leg chaetotaxy normal for this instar. Larval stage known only for a few species. ZOOL. 14, 5. 23 348 G. OWEN EVANS & W. M. TILL Protonymph : Active, feeding instar. Chelicerae, basis-capituli and hypostome as in the female. Pedipalpal chaetotaxy (I—4-5-12), rarely genu and tibia deficient ; palptrochanter without medio-ventral keel. Dorsal sclerotization comprising well developed podonotal shield with marke” posterior extension; mesonotal scutellae conspicuous, pygidial shield minute, bearing at the most one pair of setae, or represented by small scutellae or absent. Podonotal shield with 7-11 pairs of setae. Opisthonotum with four or five pairs of setae in the J series, never markedly hypertrichous. Tritosternum as in female. Intercoxal region with three pairs of sternal setae on a distinct sternal shield ; genital setae present (Liponyssoides) or suppressed (Dermanyssus). Opisthogastric cuticle with six pairs of setae. Anal shield well-defined and with normal three setae. Stigmata and short peritremes present. Chaetotaxy of legs normal for this instar. Deutonymph: Active feeding instar. Gnathosoma, dorsal sclerotization and chaetotaxy as in the female. Tritosternum biramous, laciniae long, smooth or pilose. Sternito-genital shield well sclerotized and normally bearing four pairs of sternal setae, rarely three pairs due to the suppression of the metasternals ; genital setae present, on or flanking the posterior extremity of the sternito-genital shield. Chaetotaxy of opisthogaster as in female ; anal shield well developed and with three setae. Metapodal shields small, stigmata and peritremes as in female. Leg chaeto- taxy as in corresponding adults. Genus DERMANYSSUS De Geer Dermanyssus De Geer, 1778, Mém. Hist. Ins. 7 : 106 Type: Acarus gallinae De Geer, 1778. Chelicerae of female resembling stylets, second segment greatly elongated, digits minute, weakly dentate. Pilus dentilis, dorsal seta and arthrodial processes absent. Male chelicera with segments normal in size, fixed digit reduced, spermadacty] long, grooved, completely fused with movable digit. Chaetotaxy of venter of gnathosoma normal ; pedipalp with five free segments and a two-tined apotele. Chaetotaxy oi palptrochanter to tibia normal (2-5—6-14) in some Dermanyssus, but usually showing deficiency on the femur and tibia (2~-4-6-12) and occasionally also on the genu (2-4-5-7/8). Deutosternum with nine or more denticles in single file. Hypostome elongate, corniculi membranous ; internal malae small, simple; tectum capituli elongate, triangular, with smooth margin. Dorsal shield in female entire but reduced, mesonotal scutellae sometimes present ; podonotal region of shield with 7 to 11 pairs of setae (setae 73 and zI absent) ; opis- thonotal region with from one to four pairs of setae. Male with entire dorsal shield incorporating a greater number of setae than in the female, usually 18—20 pairs (13-I4 pairs in the podonotal and 5-6 pairs in the opisthonotal region). Tritosternum normal, bipartite, with denticulate, hyaline margin. Sternal shield usually with two pairs of setae and one or two pairs of pores ; metasternal setae THE BRITISH DERMANYSSIDAE (ACARI) 349 present or absent. Genital shield with genital setae only ; anal shield with normal chaetotaxy ; metapodal shields inconspicuous. Chaetotaxy of opisthogastric cuticle variable. Peritremes variable in length; peritrematal shields fused with podal shields posterior to coxae IV. Male with holoventral shield, or with sternitogenital and ventro-anal shields. In most species the segmental chaetotaxy of the legs shows deviations from the normal complement and protonymphal patterns are often retained, especially on tibiae II-IV. Intraspecific variation is not uncommon. The ad and fd series on tibia I each comprises only two setae, and the ad series on tibia [V and fd series on genu IV also only two setae. Tarsi III and IV in male with seta fv, modified to form a tooth-like projection. Ambulacra and claws well developed, coxa II without anterior spine. Immature stages have the features described in the diagnosis of the subfamily. Kry To FEMALES 1 Dorsal shield with at least eight pairs of setae in the podonotal region (74 present) ; no conspicuous pore on shield posterior to seta s4. cea with 12 setae. Meta- sternal setae usually present . : : : : — Dorsal shield with seven pairs of setae in fhe podonoeall region (74 absent) ; a con- spicuous pore present on each side of the shield posterior to seta s4. Palptibia with N 7 or8setae. Metasternalsetae absent . ‘ : D. alaudae (Schrank) (p. 359) 2 Dorsal shield with four pairs of setae in / series (/2 present) ; with 9 or 11 pairs of setae in podonotal region (setae z6 present) : : : ‘ : 3 — Dorsal shield with three pairs of setae in J series (/2 absent) ; with eight pas of setae in podonotal region (z6 absent or on unsclerotized cuticle) . : 4 3. Peritreme extends at least to middle of coxa II. Setae 71 and si situated on the dorsal shield. Genu IV with one postero-lateral setae —_-D.. gallinae (De Geer) (p. 350) — Peritreme extends to middle of coxa III. Setae j1 and st off the dorsal shield. Genu IV with two postero-lateral setae : D. chelidonis Oudemans (p. 353) 4 Opisthogaster with arc of stout, spine-like setae. Anal shield with maximum width about one and one-half times its length to base of postanal seta. Setae z2 and 24 on dorsal shield considerably longer than setaj4. Tibia I with two ventral setae D. quintus Vitzthum (p. 361) - Opisthogaster without arc of stout spines. Anal shield with maximum width approximately equal to its length. Setae 22, 24 and 74 on dorsal shield subequal. Tibia I with three ventral setae. : ; D. hirundinis (Hermann) (p. 356) Kry TO PROTONY MPHS!?® 1 Podonotal shield with nine pairs of setae, 71 on the shield ; opisthonotal region of dorsum with J series complete (J/1—/5) : ; : 2 - Podonotal shield with seven or eight pairs of eee jt off the shield : epiceonoc! region of dorsum with setae J2 lacking. 3 2 Posterior extension of podonotal shield SENSE, Bore roundes: reaching ice of setae ]1 (Text-fig.97A) . : : D. gallinae (De Geer) (p. 353) - Posterior extension of podonotal Shiela relatively long, pointed, extending beyond level of setae J2 (Text-fig. 978) : ; : D. chelidonis Oudemans (p. 356) 10 Protonymph of D. quintus Vitzthum not known. 350 G. OWEN EVANS & W.M. TILL 3. Podonotal shield with eight pairs of setae (74 present) ; palptibia with 12 setae D. hirundinis (Hermann) (p. 358) — Podonotal shield with seven pairs of setae (j4 absent) ; palptibia with eight setae D. alaudae (Schrank) (p. 360) Dermanyssus gallinae (De Geer) Acarus gallinae De Geer, 1778, Mém. Hist. Ins. 7: 111. Dermanyssus gallinae: Evans & Till, 1962, Ann. Mag. nat. Hist. (13) 5: 283. Dermanyssus avium Duges, 1834, Ann. Sci. nat. zool. (2) 1: 18. Dermanyssus evotomydis Ewing, 1933, Proc. U.S. nat. Mus. 82 (art. 30): 11. [For detailed synonymy see Oudemans (1936), p. 299.] Female ; Chelicera with segment I, 45 » ; II, approximately 276 » ; digits minute, weakly dentate (Text-fig. g6c). Four pairs of gnathosomal setae with c.s. about 57 « apart; Ayp.2 about 45 w apart; Ayp.1 relatively stout. Deutosternum with 11 denticles arranged in single file. Corniculi membranous, with sclerotized bars (Text-fig. 96B). Tectum capituli elongate, triangular, with smooth margin. Pedipalp (2-4-6-12) with two-tined apotele. / ON 4. lism SB Fic. 95. Dervmanyssus gallinae (De Geer), female ; dorsum (aA) and venter (B) of idiosoma. THE BRITISH DERMANYSSIDAE (ACARI) 351 Dorsal shield (756 « * 300 ») with 15 pairs of setae distributed as in Text-fig. g5A ; setae J/5 situated posterior to the shield ; setae /3 and J4 sometimes asymmetrically arranged ; /3 may be unpaired. Surface of shield reticulated. Tritosternum biramous, with denticulate, hyaline margin. Sternal shield (24 » x 150 p) reticulated, with two pairs of setae and one pair of pores ; setae sf3, metasternal setae and two pairs of pores situated on the unsclerotized cuticle. Between sft, 78 «; between sti and s¢3, 88 ». Genital shield (198 » x 126 p) reticulated ; with genital setae only. Anal shield (140 » & 123 »; maximum width 150 u) with paranal setae 36 » long, postanal seta 20 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 958. Peritreme extends to anterior half of coxa II; peritrematal shield fused posteriorly with podal elements of coxae IV. Chaetotaxy of legs shows deviations from the normal on most segments, the protonymphal patterns being retained on femur III and tibiae I-IV : I II III IV i 7 39 3 3 femur 2—ya 2 2—3—I I——0 I—#—I 3 2 2 2 “ : 2 2 2 2 OO 2, 2 5 f 2 ii 2 al 2 tibia 2—$5, 7-2 I—z, 7—I I—}, 7—I I—;, {—I Coxa II without anterior spine. Length/width (in ,) of leg segments : ii Il iG IV genu 114/66 96/60 84/54 108/54 tibia 108/60 90/57 84/52 102/48 tarsus 186/48 150/45 150/42 192/42 Male ; Chelicera with segment I, 54 1; II, 84»; fixed digit reduced, spermadactyl long (105 »), grooved, completely fused with movable digit (Text-fig. 96D). Four pairs of gnathosomal setae with c.s. about 50 » apart; hyp.2 about 45 pw apart ; hyp. normal. Other gnathosomal features as in female. Dorsal shield (636 » x 276 p) relatively broader than in the female, bearing I9 or 20 pairs of setae ; three additional pairs (s3, 73 and s5) in the podonotal and one pair (J5) in the opisthonotal region ; in some specimens 71 is also on the shield. Tritosternum with hyaline denticulate margin. Holoventral shield (528 p x 126 ») with five pairs of setae in the sternito-genital region, two pairs in the ventral region and the usual three anal setae ; sometimes one or more additional opistho- gastric setae may be incorporated. Between sf1, 63 » ; between sf1 and sé3, I10 p; paranal setae 27 », postanal seta 18 p long. Peritreme extends to anterior margin of coxa II ; peritrematal shield fused anteriorly with dorsal shield and posteriorly with podal elements of coxa IV. Unsclerotized cuticle of opisthogaster with six pairs of setae (Text-fig. 964). Chaetotaxy of legs as in female ; seta pv, on tarsi III and IV modified to form a backwardly directed tooth-like projection (Text-fig. g6E). Coxa II without anterior spine. Length/width (in p) of leg segments : 352 G. OWEN EVANS & W. M. TILL I II III IV genu 87/54 72/48 66/45 81/45 tibia 90/50 70/45 72/42 87/42 tarsus 126/42 102/40 108/36 135/36 Prrr ere per e pp E Fic. 96. Dermanyssus gallinae (De Geer). A. Venter of idiosoma of male; 8B. venter of gnathosoma of female; c. chelicera of female; pb. chelicera of male; E. distal half of tarsus IV of male. THE BRITISH DERMANYSSIDAE (ACARI) 353 Protonymph ; Gnathosoma, excluding pedipalps, essentially as in the female, but not so well developed. Four pairs of gnathosomal setae with c.s. about 36 » apart, hyp.2 about 33 m apart. Pedipalps with normal chaetotaxy (1-4-5-12) for this instar. Podonotal shield (276 » x 120 »; maximum width 204 ») with a short posterior prolongation as in Text-fig. 97A ; bearing nine pairs of setae. Opisthonotal region of dorsum with J series complete. Tritosternum with base 45 p, laciniae 70 wp. Sternal shield (168 » x 108 «) with three pairs of setae and two pairs of pores. Between sf1, 50 w; between sf1 and sf3, 120 w. Genital setae absent. Anal shield (66 » x 66 ~; maximum width 75 p) with paranal setae 15 », postanal seta about 6 » long. Opisthogastric cuticle with six pairs of setae. Peritreme extends to middle of coxa III. Chaetotaxy of legs normal for this instar except tibia 1V which sometimes lacks the postero-lateral seta (I—}, }—0/1). Coxa II without anterior spine. Length/ width (in ») of leg segments : I LOE EP IV genu 60/48 50/40 42/36 56/36 tibia 63/42 50/38 42/34 60/36 tarsus 114/36 96/30 96/30 130/30 Hasitat: D. gallinae is primarily an avian parasite of world-wide distribution which will attack mammals in the absence of its normal host. The hosts most commonly recorded are domestic fowl, turkey, duck, pigeon (Columba livia Gmelin), English sparrow [Passer domesticus (Linn.)]|, starling (Sturnus vulgaris Linn.) and canaries [Sevinus canarius (Linn.)]. Dermanyssus chelidonis Oudemans Dermanyssus hirundinis: Berlese, 1889, Acay., Myr. Scorp. ital. fasc. 53, no. 3. Dermanyssus chelidonis Oudemans, 1939, Zool. Anz. 126: 306 (nom. nov. pro D. hirundinis Berlese, 1889, non Hermann, 1804). Dermanyssus chelidonis; Evans & Till, 1962, Ann. Mag. nat. Hist. (13) 5: 283. Female ; Chelicera with segment I, 60 »; II, 420 »; digits minute (about 6 4), weakly dentate. Four pairs of gnathosomal setae with c.s. about 70 » apart, hyp.2 about 57 w apart; hyp.1 relatively stout. Deutosternum with 12 denticles in a single file. Corniculi membranous. Tectum capituli elongate, triangular, with smooth margin. Pedipalp (2-4-6-12) with two-tined apotele. Dorsal shield (816 » x 372 ») with 13 pairs of setae distributed as in Text-fig. 98A ; setae 71 and J5 off the shield. Surface of shield patterned as in Text-fig. 98a. Tritosternum with denticulate hyaline margin. Sternal shield (45 » Xx 150 yp) with two pairs of setae and two pairs of pores; s¢3 and metasternals free on un- G. OWEN EVANS & W. M. TILL Fic. 97. Dorsum of idiosoma of protonymphs of four species of Dermanyssus | D. gailinae (A) ; D. chelidonis (B); D. alaudae (c); D. hirundinis (D). iy THE BRITISH DERMANYSSIDAE’ (ACARI) 355 sclerotized cuticle. Between sti, 100 »; between sti and sf3, 93 w. Genital shield (222 » X 144 ») with genital setae only. Anal shield (168-174 » X 150 wu; maxi- mum width 180-210 ) with paranal setae 40 p, postanal seta 27 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 98B. Peritreme extends to anterior half of coxa III ; peritrematal shield short and free anteriorly, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs with deviations from the normal pattern affecting most seg- ments, with retention of the protonymphal pattern on femur III and tibiae II-IV : I i Ill IV femur 2—3—2 2—3 1 ee) ones genu js lay ue, a ae ae ee ee ae a oy ee tibia 2—3, 72 ee eee eee a Coxa II without anterior spine. Length/width (in ,) of leg segments : I PE III IV genu 120/66 108/60 114/54 135/54 tibia 120/60 102/60 108/54 132/48 tarsus 198/54 162/48 174/45 222/48 —~S Se aD \ A a. a cs ag . ix vA c t LR | Vi ‘) DS ya ee All Fic. 98. Dermanyssus chelidonis Oudemans, female ; dorsum (A) and venter (B) of idiosoma. ZOOL. 14, 5. 24 356 G. OWEN EVANS & W.M- TILL Protonymph : Dorsal shield markedly attenuated posteriorly, extending to a point behind the level of setae /2 (Text-fig. 97B) ; bearing nine pairs of setae, 71 being on the shield. Opisthonotal region of dorsum with J series of setae complete. Hapsitat: In the nests of swallows (Hirundo rustica Linn.) in England and Italy. Dermanyssus hirundinis (Hermann) Acarus hivundinis Hermann, 1804, Mémoire A ptérologique Strasbourg : 83. Dermanyssus hivundinis: Evans & Till, 1962, Ann. Mag. nat. Hist. (13) 5: 285. Female ; Chelicera with second segment greatly elongated, digits minute, weakly dentate. Four pairs of gnathosomal setae with c.s. about 57 » apart, hyp.2 about 42 pw apart, hyp.1 relatively stout. Deutosternum with ten denticles arranged in single file ; corniculi membranous with sclerotized bars. Tectum capituli elongate, triangular, with smooth margin. Pedipalp (2-4—6-12) with two-tined apotele. Fic. 99. Dermanyssus hirvundinis (Hermann), female ; dorsum (a) and venter (B) of idiosoma. THE BRITISH DERMANYSSIDAE (ACARI) 357 Dorsal shield (680 » x 294 ») with II pairs of setae distributed as in Text-fig. gga ; setae jI and /5 off the shield, seta J2 missing. Surface of shield reticulated. Tritosternum with base 60 p, laciniae 84 , with hyaline denticulate margin. Ster- nal shield (42 » * 130 ») weakly reticulated, with two pairs of setae and two pairs of pores ; s¢f3 and metasternals free on unsclerotized integument. Between sf1, 63 p; between sti and st3, 84 uw. Genital shield (180 » x I1I0 ») weakly reticulated, with genital setae only. Anal shield (130 w x 105 w; maximum width 132 yp) with paranal setae 33 mw, postanal seta 18 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 99B. Peritreme extends to posterior third of coxa II ; anterior part of peritrematal shield not developed, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs shows deviations affecting most segments, with some intra- specific variability ; protonymphal patterns retained on femur III and on tibiae II-IV : I II Ill LV femur 2—3—2 2—3—I1 I—3—o o—Z—1 genu geet 2a ip! Fs ot Ee, gt tibia 2—}, 72 1—i, #—1 I—+, #—1 I—}, ?—1 Coxa II without anterior spine. Length/width (in ,) of leg segments : I II III IV genu go/48 75/45 60/42 80/40 tibia 90/45 72/45 66/40 90/38 tarsus 165/40 126/40 126/40 162/36 Male ; Chelicera with segment I, 50 »; II, 90»; fixed digit reduced, spermadactyl long (110 »), grooved, completely fused with movable digit. Four pairs of gnathoso- mal setae with c.s. about 50 p apart, hyp.2 about 50 pw apart, hyp. normal. Other gnathosomal features as in female. Dorsal shield (570 » x 282 ) bearing 18 or 19 pairs of setae. In the podonotal region setae jI, SI, s3, 73 and s5 are on the shield ; in the opisthonotal region Z1, J5 and sometimes Z4 are on the shield. Surface of shield reticulated. Tritosternum with base 42 p, laciniae 66 p, with hyaline denticulate border. Holoventral shield (468 » x 108 ,) reticulated, with five pairs of setae in the sternito- genital region, two pairs in the ventral region and the usual three anal setae. Bet- ween stI, 52 »; between sti and sf3, 108 ». Paranal setae 27 pw, postanal seta 15 p long. Peritreme extends to middle of coxa II ; peritrematal shield fused posteriorly with podal elements of coxa IV. Chaetotaxy of legs as in female ; seta pv, on tarsi III and IV modified to form tooth-like structures. Coxa II without anterior spine. Length/width (in ,) of leg segments : 358 G. OWEN EVANS & W.. M. TILL I Il Ill IV genu 75/42 60/45 60/42 78/45 tibia 75/40 51/42 60/42 78/42 tarsus 114/34 75/36 — -- Protonymph : (Gnathosomal structures, excluding pedipalps, essentially as in female, but not so well developed. Four pairs of gnathosomal setae with c.s. about 36 pw apart, hyp.2 about 32 w apart. Pedipalps with normal chaetotaxy (I-4—5-12) for this instar. Podonotal shield (312 » * 120 #4; maximum width 198 ,) with slender posterior prolongation of variable length (Text-fig. 97D), bearing eight pairs of setae, 71 being off the shield. Opisthonotal region of dorsum with four pairs of J setae, /2 being absent. Tritosternum with base 36 p, laciniae 70 ». Sternal shield (180 p x I02 ,») with three pairs of setae and two pairs of pores. Between sf1, 48 » ; between sfi and sf3, 108 ». Genital setae absent. Anal shield (84 » x 57 #; maximum width 75 ,) ; paranal setae 24 pw, postanal seta 16 » long ; Opisthogastric cuticle with six pairs of setae. Peritreme extends to middle of coxa III. Chaetotaxy oflegsnormal forthisinstar. CoxaII without anteriorspine. Length/ width (in 2) of leg segments : I II III IV genu 51/39 45/38 42/34 51/34 tibia 54/36 45/36 42/34 51/34 tarsus 98/32 93/30 90/30 98/28 Deutonymph : Gnathosoma essentially as in female. Four pairs of gnathosomal setae with c.s. about 45 p apart, Wyp.2 about 36 p» apart. Dorsal shield (522 » xX 198 ») with chaetotaxy as in female. Tritosternum with base 50 p, laciniae 70 pw. Sternito-genital shield (270 » x I08 u) with three pairs of sternal setae, one pair of metasternal and one pair of genital setae, the metasternals and genitals being relatively short and fine. Between sfI, 60 «; between sf and st3, 110 pw. Anal shield (105 » X 80 #; maximum width 96 xz) with paranal seta 24 », postanal seta 12 plong. Integument of opisthogaster with 10 pairs of setae. Peritreme extends to posterior margin of coxa II ; peritrematal shields apparently not developed. Chaetotaxy of legs as in female. Coxa II without anterior spine. Length/width (in x) of leg segments : I iu III I\ genu 69/45 60/42 51/40 63/36 tibia 72/42 57/40 54/36 72/36 tarsus 135/36 98 /34 96/33 114/32 HaBitat: Recorded in Britain from Hivundo rustica Linn. THE BRITISH DERMANYSSIDAE (ACARI) 359 Dermanyssus alaudae (Schrank) Acarus alaudae Schrank, 1781, Enum. Ins, Austriae : 515. Dermanyssus alaudae: Evans & Till, 1962, Ann. Mag. nat. Hist. (13) 5: 279. Acayus truncatus von Olfers, 1816, De Vegetativis etc. Commentarius 1 (5): 71. Female : Chelicera with segment II greatly elongated, digits minute. Four pairs of gnathosomal setae with c.s. about 36 w apart, hyp.2 about 36 wapart ; Wyp.1 relatively stout. Deutosternum with denticles arranged in a single longitudinal file. Corniculi membranous with sclerotized bars. Tectum capituli elongate, triangular, with smooth margin. Pedipalp (2—4-5-7/8) with two-tined apotele. Dorsal shield (576 » X 300 ») with ten pairs of setae distributed as in Text-fig. IOOA ; setae 71 and /J5 lie off the shield; setae 73, 74 and J2 are absent. Seta s4 considerably longer than the setae of the hexagon (75, 25, 76). A well defined pore is located on each side of the shield shghtly posterior to seta s4. Surface of shield without a well defined pattern. Tritosternum with base 30 p, laciniae 66 »; with hyaline denticulate margin. Sternal shield (approximately 24 » * 120 ») weakly sclerotized, usually with one pair of setae; sf2 and s¢3 free on unsclerotized integument, metasternals absent. Fic. 100. Dermanyssus alaudae (Schrank), female; dorsum (A) and venter (B) of idiosoma. 360 G. OWEN EVANS? & W. Moe TIEL Between sii, 48 1; between sti and sf3, 57 w. Genital shield (162 » X 120 p) with no definite pattern and bearing genital setae only. Analshield (108 pw x I20n; maximum width 135 ») with paranal setae 20 pw, postanal seta 9g yp long. . Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 100s. Peritreme very short, extending to middle of coxa III; peritrematal shield not developed anteriorly, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs shows deviations from the normal pattern on most segments, including trochanter I which lacks a dorsal seta (I—{—1). Protonymphal patterns are retained on femora I and III, genua II and III and tibiae II-IV: it III IV femur 2—$—2 747 i? 6 Te 2s 6 eon 2 I—9, o—1 py Ge I—}, 2-1 tibia 2a—?, 7-2 i—i, 2— — en pot ey Coxa II without anterior spine. Length/width (in p) of leg segments : I II PE IV genu 57/50 60/48 50/48 60/45 tibia 54/48 54/48 50/45 60/45 tarsus 102/42 102/40 96/40 120/38 Protonymph ; Gnathosoma, excluding pedipalps, as in female, but not so well developed. Four pairs of gnathosomal setae with c.s. about 27 » apart, hyp.2 about 27 w apart. Pedipalp (1-4—4/5-8) with minute, two-tined apotele. Podonotal shield attenuated posteriorly to form a fairly acute point ; with seven pairs of setae distributed as in Text-fig. 97c ; 71 is off the shield and 73 and 74 are absent. Opisthonotal region with several small, ill-defined scutellae; setae /2 absent. Tritosternum biramous; sternal shield sometimes with a weakly sclerotized posterior extension, with three pairs of setae and two pairs of pores. Between s/I, 56 »; between sti and st3, 102 w. Genitalsetae absent. Anal shield (63 » X 72 pn; maximum width 80 y), paranal setae 15 », postanal seta 10 » long. Opisthogastric cuticle with six pairs of setae. Stigma at level of middle of coxa IV, peritreme extending to anterior margin of coxa IV. Chaetotaxy of legs normal for this instar except for femora II and III which have seven and five setae respectively. Coxa II without anterior spine. Length/width (in «) of leg segments : I Il Ill I\ genu 33/42 33/33 24/36 30/36 tibia 30/40 30/36 27/33 30/33 tarsus 72/30 63/30 63/27 72/27 Hasitat: Recorded from Alauda arvensis Linn. in Scotland and Europe (Austria). j | THE BRITISH DERMANYSSIDAE (ACART) 301 Dermanyssus quintus Vitzthum Dermanyssus quintus Vitzthum, 1921, Arch. Naturgesch. (A) 86: 7. Evans & Till, 1962, Ann. Mag. nat. Hist. (13) 5: 286. Female: Chelicera with segment II greatly elongated, digits minute, weakly dentate. Four pairs of gnathosomal setae with c.s. about 42 » apart, hyp.2 about 36 w apart ; Ayp.1 relatively stout. Deutosternum with ten denticles arranged in a single longitudinal file. Corniculi membranous with sclerotized bars. Tectum capituli elongate, triangular, with smooth margin. Pedipalp (2-4—6-12) with two- tined apotele. Dorsal shield (672 » X 336 mu) with I1 pairs of setae distributed as in Text-fig. to1A; setae J5 off the shield, J2 absent. Setae 72, z2 and z4 considerably longer than the remaining setae on the shield. Surface of shield weakly reticulate, with sculpturing anteriorly. Tritosternum biramous, with hyaline denticulate margin. Sternal shield (approxi- mately 40 » X 140 u) with a more strongly sclerotized central portion ; bearing two pairs of setae and one pair of pores; st3 and metasternals free on unsclerotized integument. Between sti, 72 »; between sti and s#3, 90 ». Genital shield (174 » x 60 ») with weakly defined linear markings and bearing genital setae only. Anal shield (112 » * 138 w; maximum width 174 ,») with paranal setae about 27 p long. Opisthogastric cuticle bears a prominent arc of stout setae (Text-fig. rors). Peri- Fic. 101. Deymanyssus quintus Vitzthum, female; dorsum (A) and venter (B) of idiosoma. 362 G. OWEN EVANS & W. M. TILL treme extends to middle of coxa II ; peritrematal shield not developed anteriorly, posterior part fused with podal elements of coxa IV. Chaetotaxy of legs shows deviations from the normal on most segments, with - retention of the protonymphal patterns on tibiae II-IV : I II Ill IV femur 2—3—2 I—3—I I—3—_1 i genu 2—3, 2-1 2—?, ?—1 I—?, 21 iT tibia 2—i, 7-1 I—j, {—1 I—i, {}—1 I—}, ?—1 Coxa II without anterior spine. Length/width (in yp) of leg segments : I II III IV genu 78/60 78/66 69/60 75/57 tibia 78/57 75/63 66/57 72/54 tarsus 128/51 126/52 120/50 138/48 Hapsitat : Recorded from Dendrocopos major (Linn.) in Britain, and from Dendro- copos leucotos (Bechstein) and D. major in Germany and the U.S.S.R. SPECIES INCERTAE SEDIS We have been unable to examine the type material of the following species of Dermanyssidae recorded from Britain and Eire. The original descriptions are inadequate for their certain identity. Eulaelaps affinis Hull, 1925 (g and 9), possibly a synonym of Eulaelaps stabularis (Koch). Eulaelaps lapidarius Hull, 1925 (2), possibly a synonym of Pseudolaelaps doderoi Berlese. Hypoaspis fucorum Hull, 1925 (3), referable to Hyfoaspis (Pneumolaelaps). Hypoaspis latifrons Hull, 1925 (9), referable to Hypoaspis (Gaeolaelaps). Hypoaspis nitidissimus Hull, 1918 (2), possibly a synonym of Haemogamasus pontiger (Berlese). Hypoaspis soricinus Hull, 1925 (3). Laelaps (Hypoaspis) longipes Halbert, 1915 (9), possibly a synonym of Haemo- gamasus horridus Michael. Laelaps (Hypoaspis) ovatulus Halbert, 1915 (9), referable to Pseudoparasitus. Laeliphis birksianus Hull, 1925 (Q), referable to Hypoaspis (Gymnolaelaps) . Laeliphis fuscipes Hull, 1925 ($ and 9), referable to Hypoaspis (Gymnolaelaps). SUMMARY OF NEW TAXA AND SYNONYMY 1. The following new taxa are proposed : subfamilies : Hirstionyssinae nov., Melittiphinae nov. and Pseudolaelapinae nov. THE BRITISH DERMANYSSIDAE (ACARI) 363 genus: Domrownyssus nov., type: Pneumonyssus dentatus Domrow, 1961 (Laelapinae) species and subspecies : Hirstionyssus oryctolagi nov., Hypoaspis breviseta nov., Hypoaspis colombot nov., Hypoaspis giffordi nov., Hypoaspis hyatti nov., Hypoaspis neocuneifer nov., Hypoaspis nidicorva nov., Hypoaspis casalis myrmecophila nov. 2. The following new synonymy is presented : Allonyssus Buitendijk, 1945, a synonym of Macronyssus Kolenati Austrogamasellus Domrow, 1957, a synonym of Pseudoparasitus Oudemans Daviseilla Zampt & Patterson, 1951, a synonym of Stratiolaelaps Berlese Groschaftella Samsinak, 1957, a synonym of Haemogamasus Berlese Sauronyssus Sambon, 1928, a synonym of Ophionyssus Mégnin Cosmolaelaps gurabensis Fox, 1946, a synonym of Hypoaspis miles (Berlese) Eulaelaps kolpakovae Bregetova, 1950, a synonym of Eulaelaps novus Vitzthum Hypoaspis compressus Hull, 1925, a synonym of Hypoaspis lubrica Voigts & Oude- mans Hypoaspis nolli Karg, 1962, a synonym of Hypoaspis praesternalis Willmann Hypoaspis soarianus Hull, 1925, a synonym of Androlaelaps casalis casalis (Berlese) Ichoronyssus orcadensis Turk, 1946, a synonym of Hurstionyssus latiscutatus (de Meillon & Lavoipierre) Laelaspis ovatus Willmann, 1951, a synonym of Hypoaspis astronomica (Koch) Laelaps pachypus: Oudemans, 1927 (male), a synonym of Hyperlaelaps microti (Ewing) Laelaps pachypus: Oudemans, 1927 (female) a synonym of Hyperlaelaps amphibia Zachvatkin Laelaps (Hyperlaelaps) arvalis Zachvatkin, 1948, a synonym of Hyperlaelaps microti (Ewing) Pseudoparasitus angulatus Berlese, 1917 a synonym of Laelaps (Hypoaspis) oblonga Halbert, 1915. Liponyssus rhinolophi Oudemans, 1902, a synonym of Macronyssus uncinatus (Canestrini) Ololaelaps halaskovae Bregetova & Koroleva, 1964, a synonym of Ololaelaps veneta (Berlese). ACKNOWLEDGEMENTS We are extremely grateful to Dr. F. Radovsky, George Williams Hooper Founda- tion, San Francisco, U.S.A. for allowing us to read his unpublished dissertation on “The Macronyssidae and Laelapidae (Acarina: Mesostigmata) parasitic on bats”’, and to Dr. R. E. Crabill, Dr. F. A. Turk and Dr. L. van der Hammen, for the loan of type material. 364 G. OWEN EVANS & W.-M, DILL Baker, E. W. & Wuarton, G. W. 1952. An introduction to Acarology. New York. BeERLESE, A. 1889. Acari, Myriopoda et Scorpiones hucusque in Italia veperta. Fasc. 53, No. 8. 1892. Acari, Myriopoda et Scorpiones hucusque in Italia vepevta. Ordo Mesostigmata (Gamasidae) : I-143. 1903 [1904]. Illustrazione iconographica degli acari mirmecofili. Redia 1 : 299-474. 1913. Acayvotheca Italica, Firenze, 1 and 2: 1-221. Brecetova, N. G. 1956. 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An introduction to the British Mesostigmata (Acarina) with keys to families and | genera. J. Linn. Soc. (Zool.) 43 : 203-259. — & Hyatt, K.H. 1958. The genera Podocinum Berl. and Podocinella gen. nov. (Acarina : Mesostigmata). Ann. Mag. nat. Hist. (12) 10 : 913-932. ——, SHEALS, J. G. & MacFARLANE, D. 10961. The terrestrial Acari of the British Isles. Vol. 1. | Introduction and biology. London, pp. 1-219. —— & Tir, W. M. 1965. Studies on the British Dermanyssidae (Acari: Mesostigmata). | Part 1. External Morphology. Bull. Brit. Mus. (nat. Hist.) Zool. 13 : 249-204. | Ewinc, H. E. 1923. The dermanyssid mites of North America. Proc. U.S. nat. Mus. 62 | (art. 13) : I-26. | | | REFERENCES | | | | if Farn, A. 1961. Les acariens parasites endopulmonaires des serpentes (Entonvssidae : Meso- | stigmata). Bull. Inst. voy. Sci. Nat. Belg. 37 (6) : 1-135. 1962. Les Acariens mesostigmatiques ectoparasites des serpentes. Buill. Inst. voy. Sci. Nat. Belg. 38 (18) : 1-149. Furman, D. 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ABREVIATIONS USED IN TEXT AND FIGURES ad,_, antero-dorsal setae JviI-5 internal ventral setae Gis antero-lateral setae lat.Lf. lateral lyriform fissure WW y_5 antero-ventral setae m.d. movable digit 6.5. capitular setae pi-3 sternal pores dys dorsal setae p.d. pilus dentilis d.l.f. dorsal lyriform fissure pai, postero-dorsal setae a.s. dorsal seta ply_5 postero-lateral setae fd. fixed digit Pry. postero-ventral setae gen. genital setae pxi-3 posterior accessory setae hyp. 1-3 hypostomal setae v2-6 marginal series of podonotal setae j1-6 dorsal series of podonotal setae Ri-7 marginal series of opisthonotal Ji-5 dorsal series of opisthonotal setae setae 366 G. OWEN sI-6 lateral series of podonotal setae SI-5 lateral] series of opisthonotal setae Sp. spermadactyl stI-3 sternal setae st4 metasternal setae UR submarginal series of opisthonotal setae EVANS & W. M. TILL Uae ventral setae ZI-6 median series of podonotal setae Z1-5 median series of opisthonotal setae ZvI-5 medio-lateral ventral setae INDEX TO GENERA AND SPECIES Names printed in italics refer to synonyms ; numbers in bold face refer to figures in the text. aculeifer, Hypoaspis, 15, 16, 159, 163, 166 aculeifer, Laelaps, 166 acuta, Hypoaspis, 41, 159, 162, 212 acuta, Laelaps, 212 AETHOLAELAPS, 119 affinis, Eulaelaps, 362 agilis, Laelaps, 1, 2, 127, 128, 132, 137 alaudae, Acarus, 359 alaudae, Dermanyssus, 100, 349, 350, 359 alaskensis, Haemogamasus, 244 albatoafinis, Liponyssus, 295 albatus, Dermanyssus, 278 ALEONYSSUS, 303 ALLOPARASITUS, 159, 160 alvearius, Laelaps (Iphis), 275 alvearius, Melittiphis, 66, 122, 275 ambulans, Dermanyssus, 244 ambulans, Eulaelaps, 254 ambulans, Haemogamasus, 54, 238, 241, 244 ambulans, Hypoaspis, 244 americanus, Liponyssus, 318 amphibia, Hyperlaelaps, 6, 7, 140, 141, 144 ANDROLAELAPS, III, 113, 126, 150 angulatus, Pseudoparasitus, 222 aplodontiae Alphalaelaps, 116 aycualis, Eulaelaps, 260 arvcuatus, Dermanyssus, 277, 278 arcuatus, Hiystionyssus, 277, 278, 279 arvcuatus, Leiognathus, 278 avcuatus, Liponyssus, 278, 281 aridus, Ornithonyssus, 317, 337 avvalis, Laelaps (Hyperlaelaps), 140, 141 avuicola, Laelaps, 137 avvicolae, Hirstionyssus, 295 arvicolarum, Haemogamasus, 56, 57, 241, 250 avuicolarum, Haemogamasus horrvidus, 2 ASIATOLAELAPS, 115 astronomica, Hypoaspis, 39, 159, 163, astvonomicus, Iphis, 159, 208 ATRICHOLAELAPS, 150 auris, Raillietia, 117 australis, Raillietia, 117 AUSTROGAMASELLUS, 224 avisugus, Haemogamasus, 244 avium, Dermanyssus, 350 bacoti, Letognathus, 321 bacoti, Ornithonyssus, 83, 84, 279, 302, 317, BES; 321, 320,5337 BDELLONYSSUS, 316 BERLESIA, 116 berlesei, Laelaps, 134 BEWSIELLA, 118, 119 birksianus, Laeliphis, 362 blanchardi, Hirstionyssus, 74, 75, 280, 296 blanchardi, Leiognathus, 296 blanchardi, Liponyssus, 296 bombicolens, Hypoaspis, 29, 159, 162, 193, 198 bombicolens, Iphis, 159, 191 breviseta, Hypoaspis, 30, 159, 162, 193 BREVISTERNA, 114, 238, 239 britannicus, Liponyssus, 308 brittent, Hirstesia, 312 bursa, Ornithonyssus, 317 Tol, californicus, Laelaps, 156 canadensis, Liponyssus, 318 casalis, Androlaelaps casalis, 10, 11, 151, 152, 154 casalis, Iphis, 152 centralis, Pseudoparasitus, 47, 225, 227 CERATONYSSUS, 329 ceylonicus, Echinolaelaps hirsti, 134 chelidonis, Dermanyssus, 97, 98, 340, 353 claviger, Hypoaspis, 24, 159, 161, 182 claviger, Laelaps, 182 clethrionomydis, Laelaps, 128 coleoptratorum, Gamasus, 279 colomboi, Hypoaspis, 31, 32, 159, 161, 104 EE compressus, Hypoaspis, 172 confinis, Ololaelaps, 235 COSMOLAELAPS, 158, 159, 160 crassipes, Laelaps, 137 cricetophilus, Haemolaelaps, 156 cuneifer, Cosmolaelaps, 184 cuneifer, Hypoaspis, 25, 159, 161, 184, 187, 189 cunetfer, Laelaps, 184 cyclaspis, Liponyssus, 312 CYCLOLAELAPS, 150 cynomys, Hirstionyssus, 279, 280 DAVISIELLA, 159 decumani, Myonyssus, 269, 271 dentatus, Laelaps, 227 dentatus, Pneumonyssus, 121 dentatus, Pseudoparasitus, 48, 225, 227 DERMANYSSUS, tog, 118, 347, 348 DINOGAMASUS, 116 doderoi, Laelaps (Hoplolaelaps), 266 doderoi, Pseudolaelaps, 63, 122, 266, 362 DOMROWNYSSUS, 121 easti, Serpenticola, 337 echidnina, Laelaps, 3, 4, 128, 134 ECHINOLEAPS, 127 ECHINONYSSUS, 276 elliptica, Caris, 306 ellipticus, Liponyssus, 306 ellipticus, Macronyssus, 76, 77, 303, 304, 306 ensigey, Laelaps (Cosmolaelaps) vacuus, 190 ENTONYSSUS, 120 eos, Hypoaspis (Haemolaelaps), 156 equitans, Hypoaspis, 40, 159, 162, 210 equitans, Laelaps, 210 EUBRACHYLAELAPS, 150 EUHAEMOGAMASUS, 239 EULAELAPS, 114, 238, 239, 259 eusoricis, Hirystionyssus, 286 evansi, Ondatralaelaps, 147 evotomydis, Dermanyssus, 350 fahrenholzi, Androlaelaps, 12, 151, 156 fenilis, Gamasus, 152 festinus, Laelaps, 132 flaviot, Echinolaelaps, 134 flavus, Deymanissus, 308 flavus, Liponyssus, 308 flavus, Macronyssus, 77, 78, 79, 303, 304, 308 FONSECAONYSSUS, 316 freemani, Hypoaspis, 152 fucorum, Hypoaspis, 362 INDEX fuscicolens, Hypoaspis, 32, 161, 196 fuscipes Laeliphis, 362 GAEOLAELAPS, 112, 113, 159, 160 gallinae, Acarus, 348, 350 gallinae, Dermanyssus, 95, 96, 97, 337, 359 giffordi, Hypoaspis, 17, 159, 163, 170 gigas, Liponyssus, 271 gigas, Myonyssus, 64, 65, 269, 271 glasgowt, Androlaelaps, 156 glasgowi, Laelaps, 156 GROSCHAFTELLA, 239 gurabensis, Cosmolaelaps, 222 GYMNOLAELAPS, 113, 159, 160 haemisphaericus, Ololaelaps, 231% HAEMOGAMASUS, 111, 114, 119, 238, 239, 240, 279 HAEMOLAELAPS, 150 haemorrhagicus, Haemolaelaps, 152 HALARACHNE, 121 halaskovae, Ololaelaps, 231 HALBERTIA, 159 HAMERTONIA, 120 HEMILAELAPS, 113, 115 HEMILAELAPS, 259 hermaphrodita, Laelaps, (Iphis), 150 heselhausi, Hypoaspis, 21, 159, 162, 177 hilaris, Laelaps, 1, 2, 127, 128 hilavoides, Laelaps, 132 HIRSTESIA, 303 hirsti, Echinolaelaps, 134 HIRSTIONYSSUS, 118, 276, 277, 279 hirsutosimilis, Haemogamasus, 55, 238, 241, 242, 246 hirsutus, Haemogamasus, 53, I14, 239, 242 hivundinis Acarus, 356 hirundinis, Dermanyssus, 97, 99, 349, 356 hivundinis, Dermanyssus, 353 HISTRICHONYSSUS, 119 HOLOSTASPIS, 159, 160 hopkinsi, Raillietia, 117 HOPLOLAELAPS, 224, 266 horridus, Haemogamasus, 57, 58, 240, 250, 252, 362 humerata, Hypoaspis, 40, 159, 162, 212 humeratus, Laelaps (Laelaspis), 212 hyatti, Hypoaspis, 33, 159, 161, 198 HYPERLAELAPS, 113, 126, 140 367 349, 240, 241, 359, 241, 368 INDEX HYPOASPIS, t09, 111-4, 127, 158, 159, 160 HY POASPIS, 228 ICHORONYSSUS, 118, 119 enversus, Hypoaspis (Haemolaelaps), 172 IPHIOPSIS, 109, 116 IPHIS, to9 isabellinus, Hirstionyssus, 73, 280, 281, 295 isabellinus, Liponyssus, 295 ISCHNOLAELAPS, 150 ISCHYROPODA, 238, 239 wsotvicha, Holostaspis, 159, 203 isotricha, Hypoaspis, 36, 159, 163, 203 IXOBIOIDES, 115 IXODORHYNCHUS, 115 JACOBSONIA, 116 johnstoni, Hirstionyssus, 279 johnstoni, Macronyssus, 279 JULOLAELAPS, 113, 116 kocht, Laelaps, 141 kolenatit, Liponyssus, 312 kolenatii, Macronyssus, 79, 80, 303, 304, 312 kolpakovae, Eulaelaps, 263 hrameri, Gamasus, 158, 159, 163 krameri, Hypoaspis, 13, 14, 159, 161, 163 lacertinus, Leiognathus, 345 lacertinus, Ophionyssus, 94, 337, 345 LAELAPS, 109, 111, 113-8, 126, 127, 140, 145, 147 LAELAPSOIDES, 224 LAELASPIS, 113, 159, 160 LAELASPULUS, 113 laevis, Hypoaspis, 43, 44, 159, 162, 220 laevis, Laelaps, 220 lapidarius, Eulaelaps, 362 latifrons Hypoaspis, 362 latiscutatus, Hirstionyssus, 71, 72, 279, 280, 281, 201 latiscutatus, Liponyssus, 291 LEIOGNATHUS, 109, 316 LEPRONYSSUS, 303 LIPONISSUS, 239 LIPONYSSOIDES, 118, 347, 348 liponyssoides, Haemogamasus, I14 LIPONYSSUS, 239, 240 lobatus, Liponyssus, 308 longimana, Caris, 303 longipes, Laelaps (Hvpoaspis), 362 lubrica, Hypoaspis, 18, 159, 161, 172 mabuyae, Ophionyssus, 337 MACROLAELAPS, 127 MACRONYSSUS, 118, 121, 240, 278, 301,- 302, 303 marginalis, Hypoaspis, 34, 159, I61, 199 marginalis, Pneumolaelaps, 199 mawsoni, Eulaelaps, 257 megaventralis, Atricholaelaps, 152 MELITTIPHIS, 275 meridionalis, Laelaps, 224 meridionalis, Pseudoparasitus, 225 MESONYSSOIDES, 120 michaelt Haemogamasus, 254 microti, Hyperlaelaps, 6, 7, 141, 144, 145 microti, Hypoaspis, 156 microti, Laelaps, 137 microti, Tetvagonyssus, 140, 141 MICROTILAELAPS, 127 miles, Hypoaspis, 45, 159, 160, 222 miles, Laelaps (Iphis), 159, 222 minutissima, Hypoaspis, 35, 159, I6I, 201 mohrae, Haemolaelaps, 156 mohrae, Ichoronyssus, 306 molestus, Haemolaelaps, 152 monodi, Ophionyssus, 337 montana, Hypoaspis, 37, 159, 163, 206 montanus, Laelaps (Oolaelaps), 206 montanus, Myonyssus, 269 multispinosa, Ondatralaelaps, 8, 9, 147 multispinosus, Laelaps, 145, 147 murinus, Hypoaspis, 172 muris, Laelaps, 5, 128, 137 musculi, Acarus, 277, 278, 2 muscult, Dermanyssus, 277, : musculi, Hirstionyssus, 277, 279, 280, 201 muscult, Liponyssus, 326 MYOLAELAPS, 127 MYONYSSUS, 269 myrmecophila, Androlaelaps casalis, 11, 151, 154 myrmecophila, Hypoaspis, 42, 44, 159, 162, 214, 218 myrmecophilus, Laelaps, 159, 214 MYRMOLEICHUS, 116 MYRMONYSSUS, 116 myrmophila, Hypoaspis, 43, 44, 159, 162, 218 myrmophila Laelaps, 218 nagayoi, Liponyssus, 321 natricis, Dermanyssus, 335, 337 natricis, Ophionyssus, 90, 91, 92, 302, 337, 340 NEOBERLESIA, tog | | | INDEX neocuneifer, Hypoaspis, 26, 159, 161, 187 NEOLAELAPS, 118 NEOLIPONYSSUS, 335 nidi, Haemogamasus, 59, 240, 241, 254 nidicorva, Hypoaspis, 22, 23, 159, 162, 179 nitidissimus, Hypoaspis, 362 noctula, Dermanyssus, 278 nolli, Hypoaspis, 173 NOTOLAELAPS, 118 novus, Eulaelaps, 62, 260, 263 oblonga, Hypoaspis, 46, 159, 160, 222 oblongus, Laelaps (Hypoaspis), 159 occidentalis, Ceratonyssus, 334 occidentalis, Steatonyssus, 89, 302, 330, 334, 335 oculatus, Hypoaspis, 152 @LOLAELAPS, 113, 125, 126, 224, 228 ondatrae, Ornithonyssus, 317 ONDATRALAELAPS, 113, 126, 145 oophila, Hypoaspis, 38, 159, 163, 206 oophila, Laelaps, 206 orcadensis, Hirstionyssus, 280 orcadensis, Ichoronyssus, 291 OPHIONYSSUS, 109, 124, 300, 301, 302, 335, 336, 337 ovibatoides, Laelaps, 260 ORNITHONYSSUS, 124, 278, 279, 301, 302, 316, 317 ORYCTOLAELAPS, 113 oryctolagi, Hirstionyssus, 67, 69, 280, 286 otomys, Hirstionyssus, 279 oudemansi, Eulaelaps, 260 oudemansi, Haemogamasus, 257 OUDEMANSIELLA, 335 ovalis, Laelaps, 214 ovatulus, Laelaps (Hypoaspis), 362 ovatus, Laelaps, 208 pachypus, Laelaps, 140, 141, 144 pacificus, Liponyssus, 318 pallidus, Laelaps, 134 parvanalis, Laelaps, 147 PATRINYSSUS, 276 pavlovskii, Ornithonyssus, 329 pavilovskyi, Haemogamasus, 244 pedalis, Laelaps, 260 pereira, Ornithonyssus, 317 periblepharus, Steatonyssus, 87, 88, 278, 302, 329, 330 pilifer, Hypoaspis laevis, 175 pipistrvellae, Dermanyssus, 278 Pipistrelli, Liponyssus, 312, 326 369 pipistreli, Ornithonyssus, 85, 86, 303, 318, 326, 329 placentula, Laelaps, 235 placentula, Ololaelaps, 52, 229, 235 PNEUMOLAELAPS, 113, 159, 160 PNEUMONYSSUS, 121 PODOCINUM, 109 pontiger, Groschaftella, 257 pontiger, Haemogamasus, 60, 114, 240, 241, 257, 362 pontiger, Laelaps (Eulaelaps), 257 praesternalis, Hypoaspis, 19, 159, 162, 173 PRISTOLAELAPS, 228 propheticus, Laelaps, 260 PSEUDOLAELAPS, 266 PSEUDOPARASITUS, 113, 127, 224, nN nN nn quintus, Dermanyssus, 101, 349, 361 RAT RIELAP EARS. 027, vhabdomys, Ischnolaelaps, 156 RHINOLAELAPS, 259 vhinolophi, Liponyssus, 315 sanguineus, Haemogamasus, 321 sardoa, Hypoaspis, 20, 159, 162, 175 savdous, Laelaps (Androlaelaps), 175 saurarum, Liponyssus, 342 saurarum, Ophionyssus, 92, 93, 302, 337, 342 SAURONYSSUS, 335, 336, 337 scalopi, Haemolaelaps, 156 SEIUS, 109 sellnicki, Ololaelaps, 50, 51, 123, 229, 231, 235, 237 SERPENTICOLA, 335 serpentium, Ichoronyssus, 337 setiger, Liponyssus, 156 setosus, Liponyssus, 240 sigmodoni, Atricholaelaps, 156 simicola, Pneumonyssus, 121 smith, Hypoaspis, 172 soarianus, Hypoaspis, 152 soricinus, Hypoaspis, 362 soricis, Hirstionyssus, 70, 71, 280, 281, 286 sorvicis, Ichoronyssus, 286 spinigey, Liponyssus, 147 spiniger, Tetvagonyssus, 147 stabularis, Eulaelaps, 61, 260, 362 stabularis, Gamasus, 259, 260 STEATONYSSUS, 278, 279, 301, 302, 329 stegemant, Laelaps, 156 sternalis, Haemogamasus, 244 379. INDEX steynalis, Liponyssus, 315 STRANDTIBBETTSIA, 115 stvandtmanni, Atricholaelaps, 156 STRATIOLAELAPS, 159, 160 STYLOCHIRUS, 109 syluiarum, Deymanyssus, 316, 318 sylviarum, Ornithonyssus, 82, 123, 302, 317, | 318 talpae, Hirstionyssus, 67, 68, 69, 277, 278, 280, 281 tenuiscutatus, Liponyssus, 321 TETRAGONYSSUS, 269 TRICHOSAUROLAELAPS, 276 triscutatus, Dermanyssus, 337 truncatus, Acurus, 359 tumidulus, Laelaps, 231 TURKIELLA, 150 uncinatus, Leiognathus, 315 uncinatus, Macronyssus, 81, 303, 304, 315 vacua, Hypoaspis, 27, 28, 159, 161, I90 vacua, Laelaps, 190 vacuus, Laelaps (Cosmolaelaps), 190 veneta, Ololaelaps, 49, 229, 231 venetus, Hypoaspis, 228, 231 virginianus, Laelaps, 156 vitethumt, Echinolaelaps echidninus, 134 ZYGOLAELAPS, 150 | | { | { | i} | i | i} { = = \ | | ip — i 8 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING J. E. HILL _ 4 BULLETIN OF | THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY 3 LONDON: 1966 Na x ee TURAL wistoe mee Viv OP THE GENUS: PHILETOR (CHIROPTERA; VESPERTILIONIDAE) BY J. E. HILL Department of Zoology, British Museum (Natural History) Pp. 371-387 ; 3 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 14 No. 6 LONDON: 1066 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), istituted 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 @ separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 14, No. 6 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 2 September 1966 Price Seven Shillings Ren eEVIEW OP THE GENUS PHILETOR (CHIROPTERA : VESPERTILIONIDAE) By J. E. HILL SYNOPSIS The genus Philetoy is reviewed and its geographical distribution extended from New Guinea to Malaya by the inclusion of Eptesicus verecundus Chasen, 1940. Particular attention is given to features considered of value in determining the affinities of the genus within the Vesper- tilioninae and its relationships are examined in detail. INTRODUCTION THE vespertilionid genus Philetor with type species P. rvohui was described by Thomas (1902) from a series of ten specimens obtained in the Albert Edward Mountains, Papua, New Guinea. Since that time, additional specimens of P. vohui from New Guinea have been received by the British Museum (Natural History) and, with the original series, these form the basis of this brief review. At the same time, an examination has been made of the holotype of Eptesicus verecundus, described by Chasen in 1940 from Malaya and apparently so far known only from this and one other specimen. The holotype proves referable not to Eptesicus but to Philetor, Chasen having failed to note the salient features of the external genitalia and skull. The major features of Philetor as thus understood have been studied with a view to establishing the relationships of this small but interesting genus. PHILETOR Thomas Philetoy Thomas, 1902: 220. Type species Philetoy vohui Thomas. A study of the genera Pipistrellus, Nyctalus, Eptesicus, Tylonycteris and Mimetillus indicates that apart from its curiously specialized external genitalia (which are approached in some respects by Tylonycteris and Mimetillus) there are no other exclusive diagnostic characters which serve to isolate Philetor as sharply as was thought by Thomas or by Miller (1907: 213). However, it may be distinguished readily by a number of features in combination. The wings are narrow, with the fifth digit much reduced, its total length equal approximately to the combined length of the metacarpal and one-half of the length of the first phalange of the fourth digit. The braincase is high and rounded, the supraorbital region inflated with prominent supraorbital tubercles. The inner upper incisor (i?) is bifid and is long and narrow in contrast to the outer upper incisor (i%), which is small and conical. There is a prominent secondary posterior upper canine cusp and the small anterior upper premolar (pm?) is lacking. The sole upper premolar (pm*) and the second lower premolar (pm,) are very much shortened so that their length at the cingulum is equal to approximately one-half of their width and at the alveolus is equal approximately to one-third of their width. The third upper molar (m‘) is not reduced and has three well-defined commissures. ZOOL. 14, 6. 25 374 .o ESI Philetor rohui Thomas The head is short and broad, with a wide, flattened muzzle, the narial openings sublateral, widely separated and not projecting. The anterior part of the muzzle is sparsely clothed with hair, with a distinct fringe of hairs on the upper lip. The ears are short, triangular in outline and rounded at the tip. There is a small lobe at the base of the anterior margin of the ear, above which the anterior margin is strongly convex near its base but is otherwise straight or nearly so, the posterior margin of the ear being slightly convex in its upper and lower parts, these separated by a shallow concavity. There is a small antitragal lobe, the posterior margin of the ear terminating behind and slightly below the angle of the mouth. There are a few sparse hairs on the anterior part of both the internal and external faces of the ear. The tragus is short, thick and fleshy, its anterior margin straight for its basal half but otherwise slightly convex, the upper part of the tragus slightly prolonged anteriorly. The upper margin of the tragus is slightly convex and its posterior margin slightly so or straight, with a small triangular lobe near the base. This lobe is absent in the holotype of verecundus but apparently has been destroyed during the extraction of the skull from the specimen in alcohol. The wing is narrow, with the fifth digit much reduced, its total length exceeding the length of the meta- carpal of the fourth digit only by one-half of the length of the first phalange of the fourth digit. The metacarpal of the fifth digit is by far the shortest of the meta- carpals and the combined lengths of the two terminal phalanges of the fifth digit are equal only to the length of the first phalange of the third digit. The thumb is short and stout. There is a distinct post-calcarial lobe supported by a robust calcar which extends along approximately one-half of the length of the posterior margin of the interfemoral membrane. The genus is remarkable in the extraordinary complexity of the structure of the external genitalia. The penis (Text-fig. 1a—b) is long, with a stout shaft and large, prominent glans. It is densely pilose only at its base. Distally, the dorsal surface of the shaft (Text-fig. Ia) bears a cushion-like pad, on occasion divided to some extent by a longitudinal median depression. For much of its area this pad is moderately clothed with short, stiff bristles. The ventral surface of the shaft (Text-fig. 1b) is prolonged distally by a projecting preputial flap or fold, the slender stem of the glans penis emerging between its lateral lips which each bear a tuit of moderate, rather stiff hairs. According to Thomas (1902 : 221) the penis of Phzletor lacks a prepuce: however, it seems evident that the flap or fold projecting from the lower part of the penial shaft is the lower or ventral lip of the prepuce, its upper or dorsal lip forming the anterior margin of the curious cushion-like pad which embellishes the distal part of the dorsal surface of the shaft. The glans penis is a swollen, approximately triangular structure pierced on its dorsal surface near the tip by the urethral opening. It is supported by a slender, upwardly curved stem which emerges from the tip of the shaft a little below its centre. Ventrally, the stem extends beneath the swollen glans penis almost to the tip and it is bordered by lateral fissures in the body of the glans: the underside of the stem has a median AVE VEE WO (EEE GENUS PHITLE TOR 375 longitudinal groove extending almost to the point of emergence from the prepuce (Text-fig. 1b). The os penis or baculum (Text-fig. Ice) is strongly curved dorso- ventrally (Text-fig. rd) and has a wide, deep flanged base, slender shaft and an expanded tip. Viewed dorsally (Text-fig. 1c) the base of the os penis forms a Cc 2 mm. Fic. 1. Philetoy vohuivohui. a, dorsal aspect of penis ; 6, ventral aspect of penis ; c, dorsal aspect of os penis; d, lateral aspect of os penis ; e, ventral aspect of os penis. 376 Jess. EEL solid, bifid structure: ventrally (Text-fig. re) the base is hollowed, the medial groove thus formed extending to the base of the stem. As suggested by Thomas, ~ the paired basal flanges evidently support the cushion-like pad on the dorsal surface of the penial shaft and the enlarged tip of the os penis supports the glans. The external genitalia of the female (Text-fig. 2a—b) are similarly complex. The vulval opening is separated from the anus by paired, swollen perineal cushions or pads (Text-fig. 2a) which are divided by a median longitudinal groove and which have their surfaces sparsely covered with short hairs. Anterior to the vulval opening and normally partially concealing it there is a wide, subtriangular pad, partially divided longitudinally by a shallow median trough, its base anteriorly situated and with its apical part directed posteriorly and immediately above the vulval opening. This fleshy pad clearly corresponds to the prepuce of the male and a well-developed glans clitoris emerges from the fleshy fold forming its apical part (Text-fig. 2a). It is separated from the perineal pads by deep lateral fissures. There is a slightly swollen area forming a low cushion immediately posterior to the anus. The vulval opening is transverse, but also extends anteriorly as a narrow longitudinal slit (Text-fig. 2b) between fleshy lips, each with a small projecting spur posteriorly where they form the anterior rim of the transverse part of the vulval opening. No doubt this longitudinal slit led Thomas (1902 : 221) to his statement that the vulva was longitudinal instead of transverse : in fact the actual opening is transverse but is prolonged longitudinally. Posteriorly, the rim of the vulval opening is formed by the anterior walls of the perineal pads, which extend inwards in shelf-like fashion to form its posterior margin. Normally, the apical part of the subtriangular anterior pad fits into the shallowly V-shaped margin formed by the anterior part of the two perineal pads, thus concealing the vulval opening. The glans clitoris is normally concealed within the apical folds of this pad, its tip only being exposed. Thomas (1902 : 221) suggested that it appears probable that the slender projecting tip of the glans penis alone enters the vulva and the arrangement of cushion-like pads on the dorsal surface of the penis and on the perineum of the female supports this conclusion. The bristly pad on the dorsal surface of the penial shaft evidently engages with the perineal pads of the female during copulation while the tip of the glans clitoris engages in the groove beneath the stem supporting the glans penis. All of the available specimens of P. vohwi are preserved in alcohol. From these it appears that the dorsal surface of the body is uniformly dark brown, as is the head and nape: the ventral surface is similar in colour to the back but is slightly paler. As suggested by Thomas (1902 : 221) from the original series, which was even then in a bad state of preservation, the fur is short and close. It does not extend appreci- ably on to the flight membranes except for a sprinkling of fine hairs on the ventral surface of the wing from the body to a line joining the elbow and knee and for a few sparse hairs at the root of the tail. The skull is short and wide, with short, wide, rounded braincase, its frontal part elevated. There is a slight occipital helmet with sharp lambdoidal ridges in older specimens, The sagittal crest is weak, dividing anteriorly into weak supraorbital 377 A REVIEW OF THE GENUS PHILETOR ‘pasodxe Sutuedo yearna ‘ q : SI[C}IUSS [eUI9}xXo oTeWoT "uw 7 faa Sa oe at ¢ D ‘INYO INYOA AOJO}1Y T ZO 378 Ji Es BL ridges which terminate in prominent supraorbital tubercles. The interorbital region is broad and the rostrum wide and rather high, the anteorbital foramen enclosed by a comparatively wide bar of bone. The zygomata are slender, in older specimens with a small inferior projection anteriorly, external to m*. They are slightly widened posteriorly. The narial emargination is deep, extending posteriorly to a line joining the supraorbital tubercles and almost halfway to the interorbital con- striction. The anterior palatal emargination is deep and is wide posteriorly, slightly chordate in outline, in older specimens with a median anterior palatal spine. The palate is short, wide and domed, with a large, blunt, ligulate post-palatal spine. The meso-pterygoid fossa is wide and there are moderate basial pits: the bullae are high and are slightly inflated. The inner upper incisor (i?) is elongate, narrow, and bicuspid, the posterior or outer cusp a little smaller and lower than the anterior or inner cusp. The outer upper incisor (13) is small and conical, its height only slightly exceeding the cingulum height of 1?, with a well-developed cingulum in contact or nearly so with the inner tooth. It is separated from the canine by a moderate diastema. The upper canine has a prominent posterior secondary cusp extending for one-third to one-half of the height of the tooth. The upper premolar (pm%) is short, its length at the cingulum equal to approximately one-half of its width and it is tightly compressed between the canine and the first upper molar (m1). The upper molars exhibit no especial peculiarities : m* is not reduced and has a prominent metacone and three well- defined commissures. The lower incisors are tricuspid and are not imbricated. They are situated in the line of the toothrow and are not turned at all transversely to it. The outer lower incisor (ij) is very slightly wider than 1, or i,. The lower canine has small anterior and posterior cingulum cusps. The anterior lower premolar (pm,) is equal in height to the second lower premolar (pm,) and is comparatively unreduced, its length at the cingulum equal approximately to its width. The second lower premolar (pm,) is much reduced, its length at the cingulum rather less than one-half of its width and with its posterior face hollowed where it engages with the anterior face of pm*. Its crown area is equal to or barely exceeds the crown area of pm,. The lower molars have no unusual features. Philetor rohui rohui Thomas Philetoy rohui Thomas, 1902: 220. Albert Edward Mountains, Papua, New Guinea, 6,000 ft. Narial emargination not abruptly widened just above the roots of 1?-?. Minimum, maximum and (in parentheses) mean measurements (in millimetres) of fourteen specimens, except where stated: length of forearm (twenty specimens) 31-3—35°5 (33:4) ; greatest length of skull 13-4-14:8 (14:3); condylobasal length (thirteen specimens) 12-9-14-0 (13:6) ; least width of interorbital constriction 4-3-4:8 (4-6) ; zygomatic width (four specimens) I0-3-10-7 (10-5); width of braincase 7-3—-8-2 (7-8) ; c-m$ (front of canine to crown of m§) 4:4-4:9 (4:7). This subspecies is known so far only from New Guinea. It is recorded from speci- mens in the Archbold Collections of the American Museum of Natural History from AUREVIEW OF THE GENUS. PHITEETOR 379 Oomsis, Morobe District and Kassam, Highlands District, in the Territory of New Guinea by Brass (1964: 180, 204) who also (1956: 136) records it from Biniguni Camp, Gwariu River, Papua, again from the results of the Archbold Expeditions. Tate (1942 : 265) records a specimen in the Archbold Collections from the Idenburg River, West Irian. Laurie (1952 : 313) recorded a series of specimens from Enaena, on the north-eastern slopes of Mount Simpson, eastern Papua, in the collection of the British Museum (Natural History). This collection also contains specimens from Dinawa, Owen Stanley Range, Papua and from Madeu, inland from Port Moresby, Papua. These are the specimens mentioned by Tate (1942 : 265). Philetor rohui verecundus (Chasen) Eptesicus verecundus Chasen, 1940: 53. Mount Kladang, Perak, Federation of Malaya, 2,646 ft. Narial emargination abruptly widened just above the roots of i?-*. Chasen described Eptesicus verecundus from two specimens originally in the collection of the Raffles (now National) Museum, Singapore. That designated as the holotype is preserved in alcohol, with the skull extracted and was collected in November, 1916 by Dr. R. Hanitsch. Originally Raffles Museum No. 199, it was transferred to the British Museum (Natural History) in 1947 and is now B.M. 47.1437. The second example, also in alcohol with the skull extracted, came from an unspecified locality in Perak and remains in the National Museum, Singapore (Gibson-Hill, 1949: 171). So far as I am aware, no further specimens have been obtained. The original description compared Eptesicus verecundus with E. pachyotis (Dobson) and FE. demissus Thomas, Chasen noting that it was smaller than either of these species. He remarked also that compared with pachyotis the body of verecundus is larger in relation to the wings, “‘ asin demissus’”’. He had available for comparison specimens of pachyotis from the Khasia Hills, Assam, “‘ very kindly sent to Singapore from the Indian Museum, Calcutta as the types of pachyotis some years ago...” and gives measurements of the length of forearm and lower leg with foot in these specimens. He states that the skulls had not been extracted, and his brief descrip- tion of verecundus omits any mention of the structure of the external genitalia or of cranial features beyond noting that the inner incisor is much the larger and is tricuspid and that the upper canines have a small posterior secondary cusp. The description is otherwise concerned with external features such as the point of insertion of the wing membrane on the leg, and with the ears and tragus: he draws attention to the presence of a distinct post-calcarial lobe. Tate (1942 : 279) says “‘ Verecundus obviously has peculiarities not seen elsewhere in Eptesicus’”’ apparently solely on _ the basis of the published description. An examination of the holotype of Eptesicus verecundus shows that in fact it is _ not an Eptesicus but is referable to Philetor, agreeing in almost every respect with | P. vohwi and differing only in a few minor and relatively insignificant points. The wing is reduced to almost exactly the same extent, with the fifth digit extending as far as a point slightly more than half-way along the first phalange of the fourth | digit. The external genitalia of the female holotype are exactly as described for P, rohui, Cranially, there is a high degree of agreement but the braincase is a little 380 j. Es AL more inflated than in P. vohwi and the narial emargination is more abruptly widened just above the roots of i?-?, with a broadly V-shaped apex. Although the supra- orbital region has been damaged on both sides of the skull, the left side retains a © prominent supraorbital tubercle: the corresponding tubercle on the right side of the skull has been lost. The inner upper incisor (i?) is elongate and narrow, bicuspid, with a faint trace of a third posterior cusp which doubtless led Chasen to describe it as tricuspid. Measurements (in millimetres) of the holotype of verecundus : length of forearm 34:0; greatest length of skull —; condylobasal length 14:5 ; least width of interorbital constriction 4-7; zygomatic width —; width of braincase 7:9; c-m (front of canine to crown of m%) 4-7. The very close measure of agreement in structure and size between verecundus and vohwi indicates that specific separation is unwarranted and for the present I consider them to be but subspecifically related. The allocation of verecundus to Philetor clearly raises some question as to the status of Eptesicus pachyotis and E. demissus. The precise status of E. pachyotis seems uncertain (Tate, 1942: 277) and unfortunately the collections of the British Museum (Natural History) contain no specimens referable to it. The descriptions by Dobson (1871: 211, 1876: 104, figs., 1878 : 206) are insufficient to enable any firm conclusions to be drawn, and the question must remain in abeyance until the holotype (Dobson, 1878 : 206) can be re-examined. It is presumably in the Indian Museum, Calcutta, whence Dobson described it. The collection of the British Museum (Natural History), however, does include the holotype of EF. demissus. Externally, there is some similarity between this species and verecundus but demissus is considerably larger and the wings are not reduced, the fifth digit not conspicuously shortened but with its metacarpal nearly as long as the metacarpal of the fourth digit and with its tip reaching almost to the end of the first phalange of the fourth digit. The external genitalia of the female holotype lack the specialisations of verecundus. Cranially, the two are markedly dissimilar. The skull of the holotype of demissus is damaged but sufficient remains to demonstrate the presence of an occipital ““helmet”’ and of a prominent sagittal crest. As Tate (1942: 277) pointed out, the roof of the narial canal and of the anterior part of the mesopterygoid fossa is raised: in verecundus these lie deep in the skull. The inner upper incisor (i*) of demissus is massive and wide, quite unlike the elongate, narrow i? of verecundus. The outer upper incisor (i) is wide, its width exceeding its length, and has a large central cusp flanked by small lateral cusps. There is a low secondary posterior canine cusp but pm‘ is not shortened: i, 3 are considerably imbricated and pm, is not reduced, its length at the cingulum nearly equal to its width and its crown area exceeding that of pm,. There seems no doubt that demissus has no close relationship to verecundus and thus to Philetoy but instead apparently constitutes a distinct group within Eptesicus. RELATIONSHIPS Thomas (1902 : 220) considered Philetor allied to Vespertilio (= presumably to Eptesicus as understood by Miller (907 : 207)), Tylonycteris and Hesperoptenus, also drawing attention to its resemblances to Pterygistes (= Nyctalus). Miller (1907 : 214) A REVIEW ‘OF DHE GENUS PHILETOR 381 ll remarked that Philetor appeared related to Tylonycteris although lacking the flatten- ing of the skull found in that genus. Subsequently, Tate (1942 : 266) thought that Philetor and Tylonycteris might be independently derived from near the joffrez group of Prpistrellus, and noted (pp. 252, 253, 265) the resemblances between the members of this group and Philetor. Evidently the relationships of Philetor are likely to be found within this complex of genera and their associated species: the absence of the small pm? in Philetory no doubt has influenced earlier authors in suggest- ing relationship to those genera in which this evanescent tooth is absent rather than to those in which it is present. There appears to be no close relationship to Hesperoptenus, which has wings of normal proportions with the fifth digit unreduced in length. Although some modi- fication of the penis has occurred in this genus, it is of a different nature to the | specialisation of the penis in Philetor, as Thomas (1902: 221) pointed out. In ' Hesperoptenus the prepuce is much developed and although the os penis is similarly divided at the base to that of Philetoy, it is a much longer structure, straight and not upwardly curved and not expanded at its tip. Cranially, the braincase of Hesperoptenus is not elevated anteriorly and there are no supraorbital tubercles. The inner upper incisor (i?) is a massive, unicuspid tooth, not elongate and narrow as in Philetor and i° in some species is displaced inwards to such an extent that it is situated behind the inner tooth. There is no secondary posterior canine cusp and pm are not shortened antero-posteriorly as in Philetor, with pm, much reduced. Externally, there is a close resemblance between Philetor and Eptesicus but in that genus the wing is not reduced as it is in Philetor. The frontal part of the brain- case 1s not elevated and there is little or no inflation of the supraorbital region. Prominent supraorbital tubercles are not developed in Eptesicus. Although bicuspid, i? is not elongate and narrow as in Philetoy and 1° is not especially reduced although smaller than the inner tooth. There is no secondary posterior canine cusp and pm are not shortened. The lower incisors are usually imbricated and pm, is reduced as in Hesperoptenus. There is stronger evidence for a degree of relationship with Tylonycteris. As Tate (1942 : 266) has remarked, if the flattening of the skull in this genus be dis- regarded, then cranially and dentally it has a very close resemblance to Philetor. Externally, it differs from Philetor in the presence of pads on the thumbs and feet and the wing is not reduced but the penis has some similarity to that of Philetor and consists of a strong shaft with a large, expanded terminal pad like that of Philetor, similarly embellished with short, bristly hairs. On the dorsal surface of the shaft the pad is divided by a median longitudinal fissure to form swollen lateral cushions but ventrally it is less swollen although extending completely across the width of the shaft. There is no preputial fold such as is found in Philetoy and the _ small glans penis emerges directly from a terminal perforation. The female external _ genitalia display none of the peculiarities of Philetor. There are no perineal pads _ although the vulval area is slightly swollen, and the vulval opening is wholly trans- verse without any median longitudinal extension. Cranially, there is some expansion _ of the supraorbital region in Tylonycteris, coupled with a varying degree of develop- 382 fe RIL bAL, ment of supraorbital tubercles, minimal in T. pachypus and its allies, maximal in T. vobustula and T. malayana. There is on occasion a small, rather poorly-defined | inferior zygomatic projection external to m* and the post-palatal spine is ligulate as in Philetor. The inner upper incisor (i?) is elongate and narrow as in Philetor. It is similarly bicuspid, the posterior cusp as high or almost as high as the anterior cusp. The outer upper incisor (i%) differs from that of P/iletor and has a large central cusp equal in height or nearly equal in height to the posterior cusp of 12, flanked by two smaller lateral cusps. It is slightly hollowed posteriorly. In Philetoy this tooth is peg-like and conical: a faint trace of lateral cusps can be found only in the holotype of verecundus. The upper canine of Tylonycteris has a strong secondary posterior cusp and pm? is slightly shortened much as in Philetor, its length at the cingulum equal approximately to one-half of its width. The lower incisors (i,_3) are slightly imbricated and pm, is but slightly reduced and is almost equal in height to pm,, which is shortened in the same way as it is in Philetor, its length at the cingulum equal to one-half or to a little less than one-half of its width and its crown area equal to or only slightly exceeding the crown area of pmg. Great reduction of the wing is found in the Ethiopian genus Mimetillus which resembles Tylonycteris in the presence of pads on the thumbs and feet and to which it has been considered (Allen, 1939 : 194) to be related. In Mzmetillus the reduction of the wing is not confined only to shortening of the fifth digit as in Philetory but extends also to shortening of the third digit, a feature unremarked in any other bat. The penis of Mzmetillus does not resemble that of Tylonycteris or of Philetor at all closely. It is long and is wide at the base, tapering to a narrow tip. The shaft has no terminal pad such as occurs in these genera, its conical distal half instead being only slightly expanded and covered with short, bristly hairs. There is a small terminal opening through which protrudes the glans penis. The female external genitalia display some similarity to those of Piiletor. The vulval opening is separated from the anus by paired, small poorly-developed perineal pads and there is a prominent posteriorly directed fold anterior to the opening which covers it and is slightly triangular in outline, a fissure at the apex indicating the glans clitoris. The vulval opening is wholly transverse and has no median longitudinal extension as in Philetor. The skull of Mimetillus is flattened as is the skull of Tylonycteris and the supraorbital region is much widened by the great degree oi inflation of the maxillaries above the anteorbital foramina: as Tate (1942 : 266, footnote) has pointed out, these swellings are not strictly homologous with the supraorbital tubercles of Tylonycteris (and Philetor) but instead are “ swellings of the area anterior to that part of the orbit which encloses the anteorbital foramen, but posterior to the foramen ’’. It is the upper part of the bar enclosing the foramen and the part of the maxilla immediately adjacent which is swollen. The zygomata are massive, in contrast to the slender zygomata of Philetor and Tylonycteris. The inner upper incisor (i?) is less markedly elongated than in Philetor or Tylonycterts but is bicuspid as it is in those genera. The outer upper incisor (i’) is almost exactly like the corresponding tooth in Tylonycteris. There is no secondary posterior canine cusp but pm! is reduced to about the same extent as it is in Philetor and Tylonycteris, a — fa —— oat — ASREVGEEW OF TE GENUS PHILETOR 383 its length at the cingulum equal approximately to one-half of its width. The lower incisors are imbricated. There is a marked difference between Mimetillus, Philetor and Tylonycteris in the proportions of the lower premolars: in Mimetillus pm, is very much reduced, almost peg-like, its height not exceeding one-half of the height of pm, and its crown area approximately one-quarter of the crown area of that tooth, which is not especially shortened, its length at the cingulum rather more than one- half of its width. Despite some similarities to Philetor and to Tylonycteris, a number of features of Mimetillus indicate that it has no very close relation to either of these genera, and Tate (1942 : 266) has remarked that certain of its characteristics suggest that it may be of independent origin. In the original description Thomas (1g02: 220) noted that Philetor resembled Nyctalus in its general appearance and in its much shortened fifth finger. It must be remembered, however, that at the time that Thomas wrote, the genus Nyctalus was held to include the species joffrez, stenopterus and brachypterus subsequently removed by Tate (1942 : 252) to form the joffre: group of Pipistrellus. There is no immediate connection between Philetor and Nyctalus as thus restricted. As under- stood by Tate, Nyctalus has a high, convex frontal region and high rostrum as in Philetor but the rostrum is less widened and supraorbital tubercles are lacking. There is no inferior zygomatic projection and the post-palatal process is spine-like. The inner upper incisor (i?) is massive, its length equal approximately to its width, and its posterior cusp is obsolete The outer upper incisor (i%) is wider than long and is deeply concave posteriorly, with a small secondary posterior cusp. There is no secondary posterior canine cusp and pm? is present, pm{ being only slightly shortened. The lower incisors are imbricated and pm, is much reduced, its crown area one-half or less than one-half of the crown area of pm,. The nearest relatives of Philetor appear to be found in the joffrer group of Pipi- strellus (more correctly the brachypterus group), created by Tate (1942 : 251); to include the three species joffret, stenopterus and brachypterus, all formerly included within Nyctalus, together with a fourth, anthony1, which he described in that paper (p. 252). Ellerman and Morrison-Scott (1951 : 159), however, retain joffrev: in Nyctalus but (p. 173) leave anthonyi in Pipistrellus. Of these four species, only joffrez and stenopterus are available to me. The members of the joff/rez group have the wing reduced to a greater or lesser extent by shortening of the fifth digit. The rostrum in these species is short and wide, with incipient or moderately developed supra- orbital tubercles and on occasion there is an inferior descending zygomatic process external to m?. The outer upper incisor (13) is small and a secondary posterior canine cusp is present: pm? is minute, pm, unreduced and pm¥ shortened antero- _ posteriorly. There is considerable justification for the removal of this group of species from Nyctalus since apart from the reduced wing and the presence of a minute pm? these features are anomalous in that genus as otherwise understood. In fact, the joffrer group of species displays a number of the features characteristic of Philetor. Tate (1942 : 252) notes that the group approaches the Oriental members _ of the saviz group of Pipistrellus in which pm? is minute and the palate shortened _ but which have not developed supraorbital tubercles. 384 1 De SU con So far as | am able to determine from the material available to me and from the literature Pipistrellus joffrec and P. anthonyi appear to be the most greatly modified - members of the group. The collection of the British Museum (Natural History) contains the male holotype of P. joffrer B.M. 88.12.1.37, from the Kachin Hills, Burma, and also three female examples, B.M. 16.3.26.2, 83, 84, from 50 miles west of Kindat, Chindwin, Burma The ear and tragus closely resemble those of Philetor and the fifth digit is correspondingly reduced, its metacarpal conspicuously shorter than the metacarpal of the fourth digit and its tip reaching a point approxiately half-way along the first phalange of the fourth digit The external genitalia (Text- fig. 3), however, differ from those of Phzletor although in some respects those of the male conform closely to the same pattern. The penis (Text-fig. 3a—-b) is shorter than that of Philetor and the dorsal surface of the shaft bears a similar but less developed bristly pad in its distal part. This pad is less swollen and less extensive than in Philetor. The preputial fold is well developed and originates a little below this rudimentary pad, from which it is clearly demarcated. It is sparsely scattered with short hairs and has a shallow median longitudinal fissure in its dorsal surface and a relatively large terminal opening. Through this protrudes the small glans penis, which is perforated near its tip by the urethral opening. So far as can be discovered, an os penis, if present, must be very small and rudimentary. The female external genitalia (Text-fig. 3c) are not specialized as in Philetor. There are no perineal pads as in that genus, the vulval area being only slightly swollen, and the vulval opening is wholly transverse with no median longitudinal extension. The lips of the vulva protrude slightly and the glans clitoris is represented by a small protuberance immediately anterior to the vulval opening. The skull of Pipzstrellus joffre: is very like that of Philetor in its general appearance, but the braincase is more elevated posteriorly and a little less so anteriorly. The rostrum is rather less elevated and usually the supraorbital tubercles are not as much developed. There is a small inferior descending zygomatic process external to m*. The palate is relatively a little narrower than in Philetoy and has a similarly ligulate post-palatal spine. The inner upper incisor (i?) is elongate as in Philetoy but is relatively very slightly wider. It is bicuspid as in that genus, the posterior cusp nearly as high as the anterior cusp. The outer upper incisor (i%) is wide, less reduced than in Philetor, and is much hollowed posteriorly. It has a large central cusp flanked by smaller lateral cusps, the postero-external cusp obsolescent, the antero- internal cusp with a small subsidiary cusp below it. There is a well-developed secondary posterior canine cusp. The anterior upper premolar (pm?) is very small and is situated in the angle or recess formed by the base of the posterior canine cusp, the postero-internal part of the canine cingulum and the anterior face of pm‘, which is slightly shortened antero-posteriorly, its length at the cingulum a little more than one-half of its width. The third upper molar is unreduced, with a well-developed metacone and three commissures. The lower incisors (i,;_3) are not imbricated and pm, is not reduced, its height nearly equal to that of pm, and its crown area equal approximately to the crown area of that tooth, which is slightly shortened antero-posteriorly, its length at the cingulum slightly exceeding one-hali a A REVIEW OF THE GENUS PHILETOR ‘PITeIIUIS [eUIO}xe syeuroy ‘9 { stuod jo yoodse Terjzu9A “g ! stuad fo yoodse jesiop ‘» “wu Z eee ee gq ‘waaffol snqjaagsidig € oly 386 i, 2 Eee of its width. There seems little doubt from the description by Tate (1942 : 252) that Piprstrellus anthonyi is very closely related to P. joffrei, differing from this species only in colour and in minor cranial details. It appears to be known only from the male holotype, a dry skin with damaged skull, and no details of the external genitalia are available. The description and measurements suggest that it may approach even more closely to Philetor than does Pipistrellus joffrei in the widening of the rostrum, the degree of development of the supraorbital tubercles and the extent of the shortening of pm#. The remaining species allocated to the joffrec group seem much less closely related to Philetor than are Prpistrellus joffrer or P. anthonyi. The ears and tragus of P. stenopterus are similar to those of P. joffrec but the wing is less reduced with the fifth digit less shortened, its metacarpal nearly as long as the metacarpal of the fourth digit and its tip reaching almost to the distal end of the first phalange of the fourth digit. The penis, although about the same length as in P. joffrei, lacks any rudimentary pad on the dorsal surface of its shaft and there is no preputial fold. It consists instead of a simple shaft with a median longitudinal fissure along its dorsal surface extending to a terminal opening. The female external genitalia are similar to those of P. joffret. The skull is much lke that of P. joffre: but the supra- orbital tubercles are less developed. The anterior upper premolar (pm?) is relatively larger than in P. joffrer and pm, is unreduced, its height equal to that of pm,, which it exceeds in both length and width. The crown area of pm, is twice that of pmy, which is more reduced than in P. joffrer. From the description by Tate (1942 : 253) based on an alleged “co-type’”’, Pipistrellus brachypterus seems very near to P. stenopterus, and, if the specimen described by Dobson (1876: 92, 1878 : 223) from the Berlin Museum is correctly identified as brachypterus, then possibly these are conspecific or even synonymous. Of particular significance is the remark by Dobson concerning the Berlin specimen that its first lower premolar (pm,) is slightly longer than and in transverse diameter nearly double the second (pm,) and is also nearly equal to the canine in vertical extent. The Vespertilioninae comprise a complex of closely interrelated genera separated in some instances by comparatively slender or even rather arbitrary distinctions and the pattern of relationship within the subfamily is often obscured by parallelism or convergence. However, there is substantial evidence to indicate a relationship between Philetor and the joffrer group of Prpistrellus and also between Philetor and Tylonycteris, although Philetor seems sufficiently removed from both Piprstrellus and Tylonycteris to justify its retention as a distinct genus, presenting as it does a combination of features not met with elsewhere. In a suggested phylogeny of the pipistrelloid genera, Tate (1942 : 233, fig.) derived Philetor, Tylonycteris and Mimeitl- lus from a point on the P7pistrellus stem. I would endorse the view expressed by this author elsewhere in the same paper (p. 266) that Phzletor and Tylonycteris are derived from near the Pipistrellus joffrei group and have lost the small pm? independ- ently. Ina number of respects the two genera display further extensions of trends evidently inherent in this group: in Philetor the external genitalia have become very much modified and Tylonycteris has developed additional specializations oi Ay ARGES Vile Wor Oi RISES | GubsNuW Ss: =e ueer eae Te Our 387 the thumbs, feet and braincase. There seems little doubt that Philetor and Tylonyc- teris are Closely related but the affinities of Mimetillus are less certain. Although in some features it approaches Philetor and Tylonycteris, in others it differs markedly from both genera and, if not an indication of independent origin, these characteristics suggest no more than a remote relationship. The exact status of the joffrec group is open to some doubt. As Tate (1942 : 252) suggests, it may warrant subgeneric recognition within Pzpistrellus but it has not been possible to examine all of the species allocated to it and until this can be done its status must remain uncertain. There is insufficient evidence to justify its transfer to Philetoy and in any event, only joffrer and anthony: among its included species show any near approach to that genus. SUMMARY The genus Philetor is considered to remain monotypic but two subspecies of P. rvohui are now recognized, the nominate subspecies being so far known only from New Guinea. Eptesicus verecundus Chasen, 1940, from Malaya, is not an Eptesicus but is referable to Philetor. It is very similar to specimens from New Guinea and is allocated to P. vohui as the second subspecies. A review of the structural features of Philetor and of other genera and species to which relationship has been postulated hitherto indicates that its affinities are with the joffrez group of Pipistrellus and with the genus Tylonycteris. REFERENCES ALLEN, G.M. 1939. Bats. Cambridge, Massachusetts. Brass, L. J. 1956. Results of the Archbold Expeditions. No. 75. Summary of the fourth Archbold Expedition to New Guinea (1953). Bull. Amer. Mus. nat. Hist. 111: 77-152, 1 pls., 1 fig. 1964. Results of the Archbold Expeditions. No. 86. Summary of the sixth Archbold Expedition to New Guinea (1959). Jbid. 127 : 145-216, 12 pls., 1 fig., 1 tab. Cuasen, F. N. 1940. A Handlist of Malaysian mammals. Bull. Raffles Mus. No. 15 :i-xx, I-209, map. Dogpson, G. E. 1871. Notes on nine new species of Indian and Indo-Chinese Vespertilionidae, with remarks on the synonymy and classification of some other species of the same family. Proc. Asiat. Soc. Beng. 210-215. —— 1876. Monograph of the Asiatic Chiroptera and Catalogue of the species of bats in the collection of the Indian Museum, Calcutta. London. 1878. Catalogue of the Chiroptera in the collection of British Museum. London. ELLERMAN, J. R. & Morrison-Scort, T. C. S. 1951. Checklist of Palaearctic and Indian mammals, 1758-1946. London. Gipson-Hir1, C. A. 1949. Bird and mammal type specimens formerly in the Raffles Museum collections. Bull. Raffles Mus. No. 19 : 133-198. Lauri£, E. M. O. 1952. Mammals collected by Mr. Shaw Mayer in New Guinea 1932-1949. Bull. By. Mus. nat. Hist. Zool. 1: 269-318, 1 fig. Mitter, G.S. 1907. The families and genera of bats. Bull. U.S. natn. Mus. No. 57 : i-xvii, 1-282, 14 pls., 49 figs. Tate, G. H. H. 1942. Review of the Vespertilionine bats, with special attention to genera and species of the Archbold Collections. Bull. Amer. Mus. nat. Hist. 80: 221-2907, 5 figs. Tuomas, O. 1902. On a new genus of Vespertilionine bat from New Guinea. Ann. Mag. nat. Hist. (7), 9 : 220-222. ZOOL 14, 6. 206 a ‘1 PRINTED IN GREAT BRITAIN BY ADLARD & SON LIMITED BARTHOLOMEW PRESS, DORKING _ PONTOBDELLINAE = (PISCICOLIDAE : HIRUDINEA) IN THE BRITISH MUSEUM (NATURAL HISTORY) ‘WITH A REVIEW OF THE SUBFAMILY L. C. LLEWELLYN . BULLETIN OF | BRITISH MUSEUM (NATURAL HISTORY) OGY Vol. 14 No. 7 LONDON: 1966 En MED i Ki Boas f PONTOBDELLINAE (PISCICOLIDAE : HIRUDINEA) IN THE BRITISH MUSEUM (NATURAL HISTORY) WITH A REVIEW OF THE SUBFAMILY BY ESC, LEEWELEYN Department of Zoology, University College of Swansea Present address: Inland Fisheries Research Station, Narrandera, New South Wales Pp. 389-439 ; 28 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 14 No. 7 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 @ separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 14, No. 7 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 9 September 1966 Price £1 PONTOBDELLINAE (PISCICOLIDAE : HIRUDINEA) IN THE BRITISH MUSEUM (NATURAL HISTORY), WITH A REVIEW OF THE SUBFAMILY BY: By LC oLLEWELEY N CONTENTS Page SYNOPSIS . : , : : : : : 5 : : . 301 INTRODUCTION . : : ; ‘ : : : : : 708302 MATERIAL . : 393 ANATOMICAL COMPARISON OF Si banoldella ‘macrothela AND " Pontobdella muricata : : : . . : : : . . : 393 Gut. : : : ; : ; : - ‘ : 393 Reproductive aon ; : : : : 5 : ‘ 397 Nervous system and ponaiacon c : : : : : : 401 PONTOBDELLINAE SUBFAM. NOV.—DIAGNOSIS : ‘ : : - 404 Genus Stibarobdella—Diagnosis . : ‘ : ; é - 405 Stibarobdella macrothela : j : ; : : - 405 Stibarobdella lovicata . A : : : : : a 413 Stibarobdella bimaculata : : . : : : » 415 Stibayvobdella planodiscus. 3 : : : : - 418 *(Stibarobdella tasmanica) . : : ‘ : ‘ : 420 (Stibarvobdella taprobanensis) : : , : ; a AZT (Stibarobdella mooret) . : 422 (Stibarobdella australiensis) . 422 Genus Pontobdella—Diagnosis 423 Pontobdella muricata . 423 Pontobdella vosmaeri 427 (Pontobdella aculeata) . : : : 3 : - - Agi (Pontobdella vagosa) . : : : : : 2 432 Genus Pentabdella gen. nov. —Diagnosis. : ; : : - 433 (Pentabdella dispar). : ‘ : : : : 2 5433 Genus et species inquivendae . ‘ : : ‘ = 7435 KEY TO THE SPECIES IN THE SUBFAM. PONTOBDELLINAE : ‘ : 487 ACKNOWLEDGEMENTS . ; ; : : F : : : sso REFERENCES : : : : : é : : : : = +438 * Species not examined during this study, but well described 7m Jitteyis, appear above in brac kets. | SYNOPSIS The conspicuously tuberculate marine leeches may conveniently be separated in a new sub- family Pontobdellinae containing the genera Stibarobdella, Pontobdella, and Pentabdella gen. nov., _ for species in which the mid-body somites bear threef, four and five annuli respectively. | The internal anatomy of Stibarobdella macyrothela is described and is found to be similar to that of Pontobdella muricata, except that the former has a somewhat longer proboscis, a pair of lateral diverticula near the origin of the caecum, traces of caecal fenestrae, more elongated |} orno more than three (p. 439). ZOOL. I4, 7. 27 392 Li. C. LLEWELLYN | seminal vesicles and ovaries and the abdominal ganglia less widely separated from one another. The extra annulus in each abdominal somite of Pontobdella seems to be innervated from the ganglion anterior to it. Of the Stzbarobdella examined, S. macrothela is particularly characterized by a very larg posterior sucker, amorphous tubercles with large, more or less square basal areas and a pair of ocular patches; S. lovicata by a large anterior sucker with a fringe; S. bimaculata by small suckers and tubercles and a pair of ocular patches ; and S. variegata by a mid-ventral tubercle on each a, annulus. Of the Pontobdella examined, P. muricata is distinguished by a large anterior sucker with a fringe and P. vosmaeri by a small anterior sucker lacking a fringe and by a mid-ventral tubercle on each a, annulus. Complete descriptions of external characters are given for these species, all the literature relating to the Pontobdellinae is reviewed and attention is drawn to a number of species inquivendae. A key is constructed for the identification of the species reviewed. INTRODUCTION THE group of closely related Piscicolid marine leeches, which are characterized by possessing numerous tubercles on each body somite, are often placed in the single genus Pontobdella Leach. Many of the descriptions of these leeches are inadequate or based on characters which vary individually in those species which have been more thoroughly studied. The literature about them is scattered and needs reviewing. In the absence of convenient works of identification, the British species are not well | known. Examination of material in the collection of the Plymouth Laboratory, | for instance, revealed the presence of two species, preserved together under the same | name, Pontobdella muricata (L.), the second, previously unrecognized species being | Pontobdella vosmaeri Apathy. A preliminary examination of fifty specimens of Pontobdella muricata, the most | examined from one species, showed that the characters of the suckers were a reliable guide to identification. Papillation and relative size always appeared to be reason- | ably constant, though slight variations in size due to contraction obviously had to be | taken into account. The number of annuli in the clitellar constriction, however, | may be cited as an example of a character which occasionally varies slightly. The number and distribution of the smaller tubercles is also variable, particularly on the smaller annuli. Some of the specimens examined appeared to be rather starved and | contracted, having a slender anterior region separated by the clitellum from a broad, flattened posterior region. These unsatisfactory characters were used by Harding (1927) as diagnostic of the subgenus Pontobdellina, which he erected for the single species Pontobdella macrothela Schmarda. Harant (1929), too, proposed a questionable new genus Parapontobdelia, for Pontobdella tateyjamensis, which was described rather briefly by Oka (1g1o). He separated it on the grounds that it was said to possess some indication of lateral | vesicles. According to Selensky (1915), however, it is quite usual for a species of | Pontobdella to show a few somewhat inconspicuous vesicles, so although there are insufficient reasons for retaining fateyamensis in Pontobdella (see p. 436), Para- pontobdella may have to be separated on other characters. REVIEW OF LHE PONTOBDELLINAE 393 As my studies progressed it became evident that one set of characters, which was always constant in any species, was the number of annuli per somite. This seems important because, in the past, closely related Piscicolidae, such as Piscicola and Calliobdella, have been divided on the basis of the number of annuli per somite, together with secondary differences. It is convenient, therefore, to use a similar criterion of classification for those species usually assigned to Pontobdella. All the species in this group are either, tri-, tetra- or pentameric, i.e. each somite of the testicular and caecal (blind-gut) regions is composed of three, four or five annuli respectively. The additional annuli per somite necessitate modification of the nervous system to innervate them. I propose, therefore, to follow Harant (1929) in restricting Pontobdella, to include only tetrameric species, and redefining Stibarob- della, to include all trimeric species, and to name a third genus for the only known pentameric species. The three genera can then be placed conveniently in a new subfamily, the Pontobdellinae, when the new classification becomes as follows :— Subfamily Pontobdellinae Genus Stibarobdella Leigh-Sharpe, 1925, for trimeric species (but see p. 439) Genus Pontobdella Leach, 1815, for tetrameric species Genus Pentabdella gen. nov., for pentameric species The sections which follow aim to give: r. An introduction to the internal anatomy of these forms by comparing dissections of the type species of the genera Stibarobdella and Pontobdella. 2. A systematic account of the genera and species placed in the sub-family, accompanied by details of several species inquirendae. MATERIAL About seventy-five specimens in the British Museum (Natural History) were examined and about twenty-five from Plymouth. They comprised seven species, two of which occur at Plymouth. The number examined of a particular species is given later, at the beginning of the description of the species. ANATOMICAL COMPARISON OF STIBAROBDELLA MACROTHELA AND PONTOBDELLA MURICATA The internal anatomy of Pontobdella muricata is fairly well known from the work of Bourne (1884), Vaillant (1870) and Leydig (1851), but that of Stcbarobdella macro- thela is hardly known at all. The general dissections depicted in Text-figs. 1 and 2 show the relationship of the organs to the somites of the body. The following description applies to both species unless stated otherwise. Gut. The proboscis in Pontobdella muricata extends from somite VII to the middle of somite IX, while in Stibarobdella macrothela it extends from somite VII into the anterior part of somite X, at least a specific taxonomic difference according to Selensky (1915). Clearly visible protractor muscles are attached to the posterior end of the proboscis sheath. Along each side of the oesophagus and pharynx le salivary glands which penetrate the proboscis at its base. The oesophagus is short and its precise limits are difficult to determine. The anterior portion of the intestine, 394 Tes 6G. see Wa oe yaNi f. H.R, P.C.R. C.A. sep. B.G.R. b.g. AR, Fic. 1. General dissection of Pontobdella muricaia to show relationship of organs to somites of the body. I-XXVII, numbering of somites; am., anus; A.R., anal region; 4@s., anterior sucker ; }.g., caecum (blind-gut) ; B.G.R., caecal region (blind-gut region) ; C.R., clitellar region ; d.l/., dorsal blood vessel; e.d., ejaculatory duct; f., fringe; g., salivary gland cells ; H.R., head region; m.i., mid-intestine; 0., ovary; p., papilla; P.c.R., pre- clitellar region ; $.7., posterior intestine ; ).m., protractor muscle ; #r., proboscis ; /.s., posterior sucker; sep., septa; t,-ts, testes; J.R., testicular region. REVIEW OF THE PONTOBDELLINAE a.s. P.C.R. CyR. AR who 395 Fic. 2. General dissection of Stibavobdella macrothela to show relationship of organs to somites of the body. p.c.d., diverticula. See Fig. 1 (p. 394) for other abbreviations. é., pigmented eye patches ; /.p., lateral projections of mid-intestine ; 396 L. Cc. LLEWELLYN which stretches from the posterior edge of the oesophagus to the XIIth or last clitellar somite, is fairly thick-walled in St:barobdella, while it appeared to be more membranous in Pontobdella. A very small pair of diverticula come out from the gut in somite XI, though Selensky (1915) found these in somite X. It should be noted that Selensky in fact called this somite VIII, but believed the head complex to contain four somites ; it is now regarded as containing six somites (Mann, 1953), sO appropriate amendments are made here and throughout the text following. | m. Y c.0.g. —— Frm, oe an. Fie. 3. Gut of Stibarobdella macrothela: c.o.g., circum-oesophageal ganglion; /e., fenestra ; m., mouth; oe., oesophagus ; 7.m., retractor muscle. See Figs. 1 & 2 (p. 394 & p. 395) for other abbreviations. REVIEW OF THE PONTOBDELLINAE 397 The mid-intestine, which extends throughout the testicular region, is divided by constrictions into six chambers in Pontobdella muricata, corresponding to the six somites (XIII to XVIII) of this region. In Stibarobdella macrothela there are instead small lateral projections on the dorsal surface of the mid-intestine (Text-fig. 3). From the posterior end of the mid-intestine arises the caecum, which in Pontob- della is entirely unpaired and extends ventrally from somites XVIII to XXVI. In Stibarobdella it extends from somites XIX to XXVI and shows signs of fenestrae in somites XX and XXII, although these were not particularly clear and seem unimportant in distinguishing genera. Selensky (1915) regarded the degree of fusion of caeca as generally constant within a genus, but Moore (1938) found in Pontobdella rugosa that the last pair of caeca are only partially fused with five fenestrae. The caecum in Pontobdella muricata shows slight constrictions corresponding to the increased musculature or rudimentary septa found between annuli a,—ay, a,—b; and within the small annulus b,, but these cannot easily be distinguished in Strbarobdella macrothela. The labelling of annuli is explained by Mann (1953) and can be seen in Text-fig. 9 (p. 403). The posterior intestine extends from somites XIX to XXVII and is surrounded for most of its length by a blood sinus, which anterior to this forms the dorsal blood vessel. In Stibarobdella macrothela there is a large pair of diverticula in somite XIX, but in Pontobdella muricata there is only a slight swelling here. Beyond this the intestine winds backwards. Numerous septa occur within it, but these are not obvious from the exterior. Finally, the rectum is short and rather inconspicuous and terminates at the anus between somites XXVI and XXVIII, or in Pontobdella muricata between the annuli of somite X XVI, or in the anterior annulus of this somite. REPRODUCTIVE SYSTEM. Both species possess six pairs of oval testes (Text-figs. 4 and 5), situated posteriorly in somites XIII to XVIII. When fully ripe each overlaps into the somite behind, becoming virtually intersegmental. Richardson (1950) described Pontobdella benhami (=Stibarobdelia macrothela) as having three pairs of testes and according to his diagram the vas deferens looped posteriorly for another two somites after the posterior testes. It seems, however, that it is quite possible for this leech to possess six pairs of testes, as is normal in Piscicolidae. In Pontobdella muricata each of the vasa efferentia leaves the testis antero-laterally, while in Stibarobdella macrothela it leaves postero-ventrally. This character, however, may perhaps vary with contraction and during the ontogeny. The vasa efferentia proceed laterally to join the vas deferens on each side. This proceeds anteriorly, following a more wavy course in Pontobdella muricata, and joins the ejaculatory duct in somite XI in this form, but in somite XII in Strbarobdella macrothela. The ejaculatory duct communicates posteriorly with a long coiled vesicula seminalis and extends anteriorly as far as somite XI, where it coils postero-ventrally and finally enters the dorsal portion of the genital atrium, which is divided dorso-laterally into two pouches. The male pore opens in somite XI, between the second and third annulus in Pontobdella muricata, and between the two annuli of this somite in t | Stibarobdella macrothela. The paired ovaries are tubular but comparatively short, leading into short oviducts L. CC. LEEWELLYN 398 ‘SUOTVEIAGIGGe Ioy}O 107 ‘Z6E ‘d ‘bs anNyT ooG *SI}SO4 podojosopun “771 5 jJONPIAO “10 “H]aYJOAIDU DIJAPYOAMAYS JO wioyshs oatjonpoadeary =*S “By ‘suorayjo sea “aa { suarojop sea “pa { soysoy ‘7-7 { SITPULUIOS B[NOISOA “A's ‘ Areao “O ! XT XK-K sepruros jo eysures oAs9u ‘VIX-N Su { wntye yeyued “ws! yonp Aroyepnoele “pia “wypaianw vyjapqojuog Jo uieysAs aAtyonpoideryy “V diy °S V ‘sory / Si) F| r Ol REVIEW OF THE PONTOBDELLINAE H.R. P.C.R. C.R. TR. B.G.R. A.R. Fic. 6. Dissection to show the arrangement of ganglia in the neurosomites of Pontobdella muricata. a.g., accessory ganglion (or lateral glia cells); an.g., anal ganglion; A.R., anal region; a.s. anterior sucker; B.G.R., caecal region (blind-gut) ; cm., commissure ; c.0.g., Circum-oesophageal ganglion; c.R., clitellar region; f., fringe; gn., ganglion ; H.R., head region; i.., segmental nerve; p., papilla; pP.c.R., preclitellar region; p.s., posterior sucker; sep., septa; ¢,-t,, testes; T.R., testicular region. 399 400 L. C. LLEWELLYN e H.R. N e- | | P.C.R. C.R | Ma li TA. nt tha | XVIII ( | im XIX ( | hy ( ile Xx fall and ral B.G.R. XXI | any X XXII | il | itt X XH ( \ 2 XXIV { AR. XXVI Th Fic. 7. Dissection to show the arrangement of ganglia in the neurosomites of Sizbarobdella macrothela. e., pigmented eye patches. See Fig. 6 (p. 399) for other abbreviations. REVIEW OF THE PONTOBDELLINAE 401 which terminate in the female pore, between the two annuli of somite XII in Stzibarob- della macrothela, and between the first and second annulus of somite XII in Pontobdella muricata. NERVOUS SYSTEM AND ANNULATION. Between the circum-oesophageal and anal ganglionic masses, lie twenty-one ganglia joined by commissures, which mark a corresponding number of somites (Text-figs. 6 and 7). These are grouped by Johansson (1896) and others as follows :— Preclitellar region Clitellar region Testicular region Caecal (blind-gut) region Anal region W DAW W Following Mann (1953), we may provisionally assume that the circum-oesophageal mass consists of six neurosomites, in spite of older views that it consists of four (Livanow, 1903 and 1904, and others). In Pontobdella muricata it innervates the anterior sucker plus the adjoining three narrow annuli, which together constitute the head region. The widths of these three annuli are in the ratio 1: 1:2. In Stibarobdella macrothela the arrangement of the head region is very similar, except that occasionally only two neck annuli can be distinguished. In both species, the next three neuro-somites of the preclitellar region VII-IX are trimeric, with their annular widths varying in the ratio r:2:1. The anterior limit of the clitellar region is rather ambiguous in all Pontobdellinae. The most obvious external clue to the position of the clitellum is a constriction of the body and a reduction in width of the annuli there. The size and number of tubercles are also reduced. The most anterior of these narrow annuli is the last annulus of somite X and the most posterior is the first annulus of somite XIII. The term “ clitellar region’ generally given to the three somites X—XII does not therefore coincide exactly with the narrow and modified part of the body. Posterior to the clitellar region lies the testicular region which is made up of six somites, XIII to XVIII. In Stibarobdella macrothela these are trimeric, with their widths varying in the ratio 1 : 2: 1, while in Pontobdella muricata they are tetrameric with their widths varying in the ratio 2:3:2:1. The ganglia always appear to be situated within the limits of the largest annulus. Behind this region lies the caecal region, which is also made up of six somites XIX to XXIV, having an arrangement similar to that in the previous region. The anal region is made up of three somites, XXV to XXVII, containing three ganglia which are much closer together. In Stibarobdella macrothela somite XXV consists of two annuli and XXVI and XXVII of one annulus each. Finally, the posterior ganglionic mass of the posterior sucker is regarded as consisting of seven fused ganglia, making a total of thirty-four in all. The distance between ganglia varies considerably and is least towards the extremities of the nerve chain and in the clitellar region (Text-fig. 8). In Pontobdella 402 L. ¢. LLEWELLYN muricata the distance between ganglia in the caecal region is relatively greater than in Stibarobdella macrothela. Each ganglion consists of ganglia cells arranged into six cell packets or capsules, with one pair on each side and one pair mid-ventrally, making three longitudinal rows. Through the middle of the ganglion run two fibrous tracts, which fuse at the centre of the ganglion (Harant & Grassé, 1959) and continue as the commissures linking successive ganglia, each tract possessing a single ‘“ commissural cell’ making a total of two between adjacent ganglia. From the fibrous tract at the centre of the ganglion the main segmental nerve runs out on each side. In Pontobdella muri- 36 32 28 Okie € 2) D oa s ~ “SS ny 404 L.., C°-LLEWELLYN cata, in the testicular and caecal region, this nerve divides near the lateral glia cells or accessory ganglia (Scriban & Autrum, 1934) into five main branches (Text-fig. 9). Nerve I, the most anterior branch, is comparatively small. It turns ventrally and innervates the ventral portion of annulus a,. Nerve II curves anteriorly and innervates annulus a, laterally. Nerve III runs laterally and terminates in the lateral region of annulus ag. Nerve IV runs laterally for a short distance and then posteriorly until it reaches the boundary between annulus b,; and bg. There it divides, the main branch turning laterally again and running along this boundary It seems likely that one of these nerve branches innervates annulus bs. Nerve V, which is the largest of these nerves, runs along the boundary between annuli a, and b; until it subdivides dorso-laterally into four branches (i-iv), innervating the lateral and dorsal regions of annulus a, and b,. Stibarobdella macrothela differs from this condition in several ways (Text-fig. Io). The main segmental nerve trunk gives off a branch as it leaves the ganglion, which terminates ventrally in the region of the border between annulus ag and a3. As in Pontobdella muricata there are five main branches at the lateral glia cells, of which I, II and III run anteriorly. Nerve IV appeared to terminate laterally in annulus ag. The main branch, nerve V, had a small branch in the region of the lateral line, which ran posteriorly and terminated at the border between annulus a, anda,. The main branch continued dorsally with two very small branches running posteriorly, until in the dorso-lateral region of annulus a, it divided, the posterior branch ii terminating in annulus ag, the anterior branch i in annulus ay. It can be seen that the arrangement of nerves I, II, III and V is somewhat similar in the two species. Only nerve IV is strikingly different, being elongated posteriorly in Pontobdella muricata and apparently innervating annulus bg. This supports the view put forward by Oka (1917), who stated that the fourth annulus of a tetrameric somite is derived from the a, annulus. It seems that annulus bg, is indeed the posterior annulus of the somite and that the modifications are for the innervation of this fourth annulus. Family PISCICOLIDAE Subfamily PONTOBDELLINAE, Subfam. nov. Diacnosis. Characterized particularly by the tubercles, which occur on at least the major (a,) annulus of each somite and usually on most annuli of the testicular and caecal regions. The latter regions are divided into trimeric, tetrameric or pentameric somites. The body is circular or slightly flattened in cross section. The clitellar constriction does not coincide exactly with the three clitellar somites (X, XI, and XII) of the “ clitellar region’ defined by Johansson (1896) for most Hirudineae, but extends from Xa, to XIIIa, inclusive (Text-figs. 11-28). TYPE GENUS. Pontobdella Leach, 1815. Remarks. Pending further studies the remaining Piscicolidae should perhaps be grouped as the Piscicolinae and Branchelliinae (Caballero, 1956) and may eventually be divided further. Leigh-Sharp (1916) suggested that those possessing pulsatile REVIEW OF THE PONTOBDELLINAE 405 vesicles or other respiratory appendages might be separated off as the Branchiob- dellinae, but this name is unacceptable, since Branchiobdella is an oligochaete, and there are intermediate forms, such as Johanssonia and Austrobdella, which make such separation difficult. Genus STIBAROBDELLA Leigh-Sharpe, 1925* Stibarobdella Leigh-Sharpe, 1925: 417; Harant, 1929: 651. Pontobdella (Pontobdellina) Harding, 1927 : 44. TYPE SPECIES. Pontobdella macrothela Schmarda. D1aGNosis. Somites of the testicular and caecal regions trimeric (but see p. 439) Stibarobdella macrothela (Schmarda, 1861). Pontobdella macrothela Schmarda, 1861 : 6. Pontobdella afva Baird, 1869 : 312. Pontobdella papillata Grube, 1871 : 56. Pontobdella zonata Weber, 1915 : 20. Pontobdella (Pontobdellina) macrothela : Harding, 1927 : 45. Pontobdella benhami Richardson, 1950 : 97. Trvachelobdella cavajbica Dequal, 1917 : 2. Stibarobdella superba Leigh-Sharpe, 1925 : 417. DESCRIPTION. General characters. About thirty specimens were examined. The testicular and caecal regions are somewhat flattened, except when fully gorged, and are distinctly broader than and clearly separated from those regions anterior to them. The body varies from yellowish brown to bluish green, even in preserved material, with the tubercles generally white or cream, and sometimes with bands or patches of pigment. The length may reach 16:5 cm., excluding suckers. The anterior sucker is slightly oval, elongated transversely and attached eccentrically, with the dorsal surface longer than the ventral. It lacks a marginal fringe (Text-fig. IIA and 12), but possesses three pairs of tentacular papillae situated marginally and one pair laterally nearer the base of the sucker. These are occasionally rather difficult to see. All specimens examined had coronet-shaped (ocular?) pigment patches on the dorsal surface of the anterior sucker, although in one long preserved specimen these had become extremely faint. The posterior sucker, which is attached centrally, is between two and three times the size of the anterior sucker and is gener- ally almost twice the diameter of its attachment and broader than the broadest part of the body. When contracted, however, it may be little more than half the greatest breadth of a fully gorged specimen. Sometimes it bears dark and light radiating bands or patches. ANNULATION. Uniannulate somites z j Biannulate somites XI, XII XXV, XXVI XXVII Triannulate somites VII-X, XIII-XXIV (Text-fig. 12) The first three annuli of the body, the anterior two of which are particularly narrow, constitute the neck, which together with the anterior sucker makes up the ZOOL. 14, 7. 28 * Including Stibarobdella biannulata (Moore), see p. 439 Es €. LEEWELLYN XIII d. di 1. v.i. m.y. 500 erele O0009 a, SoC) ia Gears OO forkeleKeleric O00! d.m. d.p. dm. v.m. d.i. CO @ Fic. 11. Stibarobdella macrothela. A. Section through anterior sucker, showing oral surface. B. Ventral surface of clitellar constriction, showing positions of genital pores and tubercles. c. (a, bandc). Glandular or furrowed tubercles. Db. Tuberculation of a typical triannulate somite split mid-ventrally. For designations of annuli (i.e. ay, ay and ay) see Mann, 1953. X—XJII, numbering of somites; Ant., anterior end; d.i., dorsal intermediate line; d.m., dorso-para-marginal line; d.p., dorso-para-median line; /-p., female pore ; g.t., furrowed tubercle ; /., lateral line ; m., mouth ; .d., mid-dorsal line ; m.p., male pore; m.v., mid-ventral line ; p., papilla; s-.e., scalloped edge; ss., sensilla ; v.i., ventral intermediate line ; v.m., ventro-para-marginal line ; v.p., ventro-para-median line. REVIEW OF THE PONTOBDELLINAE 407 oXeKeMeNekKe Ome an. Fic. 12. Dorsal view of Stibavobdella macrothela showing annulation, tuberculation and suckers. J-~XXVJIJ, numbering of somites; an., anus; A.R., anal region; a.s., anterior sucker; B.G.R., caecal region (blind-gut region); c.N., clitellar constriction; C.R., clitellar region ; e., pigment eye patches; H.R., head region; p., papilla; P.c.R., pre- clitellar region ; p.s., posterior sucker ; s., small white tubercle ; T.R., testicular region. 408 ESC. LEE WEEE YN head region. Then come three trimeric somites forming the “‘preclitellar region ”’. As already remarked (page 401) most of the first somite of the “ clitellar region ”’ also lies anterior to the clitellar constriction, which extends from Xa, to XIIla,. The male and female pores open on the ventral side of this constriction, between the two annuli of somites XI and XII respectively (Text-fig. 118). In some specimens the male pore is surrounded by tumid lips. Annulus XIIla, is extremely small and generally lacks tubercles, whilst the other annuli of the clitellar constriction are also rather small, of uniform size, and with tubercles of reduced size. Sometimes an annulus is absent from the posterior part of the clitellar constriction and this is probably annulus XIIJa,, rather than one of the annuli of somite XII. If so, it appears that the first testicular somite XIII is then bi-annulate. The remaining five somites of the testicular region XIV to XVIII and the six of the caecal region XIX to XXIV are all trimeric and their annular widths vary in the ratio 2 : 3-4: 2. The annulation and tuberculation of the anal region XXV to XXVII are subject to variation. The anus is situated in the furrow between somites XXV and XXVI. These are biannulate, whilst somite X XVII is usually uniannulate, but occasionally shows signs of becoming biannulate, as also observed by Cordero (1937-38). Moore (1927) described similar annulation in this region, but later (1958) described a further specimen with somite XXIV biannulate and XXV, XXVI and XXVII uniannulate. In all other respects annulation is generally the same as that described by Cordero (1937-38) and Moore (1927 and 1958). TUBERCULATION. Text-fig. 13 illustrates the tuberculation of a single specimen. There is considerable individual variation, particularly affecting the smaller tubercles. The big tubercles, however, are fairly constant in number and arrangement. The tubercles on the first two annuli of the neck, which are minute, are often absent ventrally. On the ventral surface of the a, and a, annuli of the preclitellar region the number of tubercles varied from 4 to 7. Where odd numbers are indicated in Text-fig. 13, this was due to bilateral asymmetry and not to median tubercles. Somite XXVI and XXVII appear to have no sensillae or small white tubercles, and their annuli have more tubercles dorsally and few or none ventrally. Text-fig. 11D shows a typical triannulate somite of the testicular or caecal region. Here and generally throughout the body, the dorsal tubercles are larger than the ventral ones. Numbers and sizes of tubercles are reduced towards the extremities of the body. These vary considerably in shape and size depending on extent of gorging, but the four large tubercles on the dorsal surface of the a, annuli differ from those of any other species examined (Text-figs. r1ca, b and cc). They are generally more or less square, occasionally with lateral indentations making them somewhat “HH” shaped (Text-fig. r1ca). They generally appear rather like cotton wool in texture and somewhat furrowed, and sometimes bear an apical ring of small sensillae. In addition to these large tubercles there are generally four small white tubercles without papillae, which are difficult to see in starved specimens, two situated dorso- laterally and two ventro-laterally. The tubercles of a, and a; are generally more rounded and more like tubercles of other Pontobdellinae, although they too may possess rather irregular surfaces and small apical sensillae. The tubercles on the RE Vie W OF GEE PONTOBDELLINAE 409 clitellar constriction are very small and rounded. Annulus XIIIa, lacks tubercles, but the other annuli of the clitellar constriction may possess up to eight both dorsally and ventrally. The size, position and number of these vary considerably. REMARKS. Hzirudo indica Linnaeus (=Pontobdella indica: de Blainville, 1827) may well have been this species (Harding, 1927). According to Cordero (1937-38) Pontobdella mooret, Pontobdella bimaculata, Pontobdella zonata and Trachelobdella carajbica are synonymous with Pontobdella macrothela. However, Oka (1929), in his comparison of the external morphology of P. moorei, P. bimaculata and Pontob- della muricata, never mentioned irregularly shaped tubercles, and these are known to be absent from P. muricata and were not illustrated in the diagrams of his other two species. Furthermore Cordero’s view appears to require an undue degree of variation Somitas. |-vI Vil Vill XII XH XIV Dorsal. Bclelalele le ealelebole lg islels[olele|olsendshobal Ventral. fees eso siaei| 5 al A sol Sze ehselkels (eho Size 1-3. 24 3°12 Somites. Dorsal. Ventral. Size |-3. Fic. 13. Diagram illustrating the tuberculation of a single specimen of Stibarobdella macrothela. The anterior half is shown above, reading from left to right, H.R., head region ; P.C.R., preclitellar region ; C.R., clitellar region; T.R., testicular region showing first two somites ; N.R., neck region; C.N., clitellar constriction where 3 and 2 show the position of the respective pores. The posterior half is shown below including, reading from left to right, B.G.R., caecal region (blind-gut region); a.R., anal region; an., anus. The numbers of tubercles dorsally and ventrally on each annulus are indicated in the squares. Where mid-lateral tubercles are present this is indicated by a small arrow pointing down- wards, to indicate that two of the tubercles counted as dorsal are actually mid-lateral. In this species numbers in brackets indicate small secondary tubercles which are situated para-marginally (terms such as this are illustrated in Fig. 11p)._ Numbers in a circle indicate the presence of small white tubercles which are in the intermediate line of the dorsal and ventral region of the annulus and usually visible laterally. There are no median tubercles in normal annuli. The size range 1-3, 3 being the largest, indicates the relative size of the annulus (and tubercles) compared with the annulus next toit. It does not attempt to indicate sizes of annuli from different parts of the body, since there is a decrease in annular size towards the body’s extremities. 410 LC, LLEWELLYN in the characters of the suckers, such as papillae and eyes, which appear to be fairly constant in other species. It is extremely doubtful whether either P. moovei or P. bimaculata is synonymous with P. macrothela. On examining the type specimen of Pontobdella afra it was found to be the same as Stibarobdella macrothela. According to Baird (1869) tubercles are absent from the a, and a, annuli of the preclitellar region, and the clitellar region, but in fact they are present though much reduced in size. Ringuelet (1944) made Pontobdella papillata Grube (1871) synonymous with Stibarobdella macrothela. Stibarobdella superba Leigh-Sharpe (1925), which was the type by monotypy of St¢barobdella Leigh-Sharpe, also appears to by synonymous with the latter so this older generic name is used in place of Pontobdellina Harding (1927). The earlier generic diagnosis seems, incidentally, to be more useful. On examining the type specimen of Stibarobdella superba, which is in a rather poor condi- tion, it was found that the median papilla on the anterior sucker figured by Leigh- Sharpe was absent. Some of the primary tubercles on the dorsal surface bore sensillae arranged diagonally and the lateral tubercles on the a, and a, annuli often bore three small sensillae arranged longitudinally. However, since sensillae are highly variable and in some specimens apparently absent, there seem to be no grounds for separating this from Stibarobdella macrothela. Richardson (1950), in his description of Pontobdella benhamt, regarded the number of tubercles on the a, annulus of the testicular and caecal region as different in his leech and Pontobdella macrothela, i.e. eight and twelve respectively. Actually S. macrothela has eight primary tubercles, with an additional mid-ventral pair of medium size, making a possible ten primary tubercles. In addition to these there are secondary tubercles situated latero-dorsally and latero-ventrally, which are often obscured by the larger tubercles on each side. These small latero-dorsal tubercles were mentioned by Ringuelet (1944), Goddard (1909), Harding (1924) and Cordero (1937-38), but those lying latero-ventrally were not mentioned. The tuberculation agrees well with Richardson’s description, which states that the a, annulus, in addition to the eight primary tubercles, may possess small tubercles without sensillae which may show clearly only on gorged specimens, and that a pair of mid-ventral secondary tubercles occur near the anterior border of the a, annulus. So the information available suggests that Pontobdella benhami is a synonym of Stibarobdella macrothela. Goddard (1909) described a specimen of Pontobdella macrothela, found in Australia, which lacked any external markings and was possibly a preserved specimen which may have lost the ocular pigment patches on the anterior sucker. Cordero (1937-38) stated that Pontobdella macrothela may or may not possess tentacular papillae and pigmented ocular patches, but this variation seems unlikely within a single species. It may be noted that the dimensions of the suckers varied greatly amongst the forms grouped by Cordero, which suggests that different species were involved. DISTRIBUTION. Widely distributed since it is found on numerous species of sharks. Specimens examined were from Gambia, Queensland, Japan, China, Barrier Reef Australia, Freemantle, British Guinana, Malindi (Kenya) and Coiba Island (Pacific). REVIEW OF THE PONTOBDELLINAE 411 They have also been recorded from Tandjong, Sumatra, Jamaica, New South Wales and the Bay of Bengal. Hosts. Black-tipped shark, Carcharinus melanopterus ; Tiger shark, Galeocerdo arcticus ; Hammer-head shark, Sphyrna sp. (Moore, 1927) ; Eulamia sp. ; Scoliodon sp. Found attached to fins, mouth and claspers. xt Fic. 14. Stibarvobdella loricata. A. Section through anterior sucker, showing oral surface. B. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. c. (a) Tubercle with sensillae. (0) Tubercle on annulus showing striae. vb. Tuberculation of a typical triannulate somite split mid-ventrally. /., fringe; st., stria; fu., tubercle. See Fig. 11 (p. 406) for other abbreviations. Fic. 15. L.- CC. LLEWELLYN Dorsal view of Stibavobdella lovicata showing annulation, tuberculation and suckers. /f., fringe. See Fig. 12 (p. 407) for other abbreviations. REVIEW OF THE PONTOBDELLINAE 413 Stibarobdella loricata (Harding, 1924) Pontobdella lovicata Harding, 1924 : 39. Stibarobdella loricata : Harant, 1929 : 651. DESCRIPTION. General Characters. About a dozen specimens were examined. The body is fusiform, tapering gradually towards the anterior end, up to 16-6 cm. long and circular in cross-section when fully gorged, but somewhat flattened dorso- ventrally in starved specimens. Although presumably faded, due to preservation in alcohol, and without external markings, the colour varied between pale yellow, green and brown, The anterior sucker is cup-shaped and attached eccentrically so that the dorsal surface is longer than the ventral. It is generally circular but may be slightly flattened laterally, so that its opening becomes slit-like. It possesses a marginal fringe and generally three pairs of inconspicuous papillae on each side (Text-figs. 144 and 15). The anterior sucker is often about equal in size to the posterior sucker, but when both are fully expanded the latter is slightly larger. The anterior sucker is usually three times the diameter of its point of attachment but never exceeds the greatest diameter of the body. The posterior sucker is cup-shaped when contracted and its diameter is then less than the greatest diameter of the body, but when expanded it is approxi- mately equal to it. It is attached centrally and is greater in diameter than its point of attachment. ANNULATION. Uniannulate somites XXVII (Text-fig. 15) Biannulate somites XXV; XXVI , XI Triannulate somites VII-XI ; XIII-XVXI eas The first three annuli, the anterior two of which are smallest, constitute the neck. The three preclitellar somites, VII-IX, are all trimeric, with the annular and tuber- cular sizes of each somite in the ratio 1:2: T. All eight annuli of the clitellar constriction, i.e. Xa; to XIIJa, inclusive, are greatly reduced and approximately equal in size (Text-figs. 148 and 15). Somite XI always appears to be triannulate and includes the male pore, which is situated mid-ventrally in the furrow between a, and ay. Somite XII is clearly divided dorsally into three annuli, but the anterior annulus is small, non-tuberculate and indistinguishable ventrally, where the female pore lies between a, and a,;. The testicular somite XITI-XVIII are all trimeric, with annular widths and tubercular sizes in the ratio I:2:1. The somites of the caecal region XIX—XXIV are similar, and the body is generally broadest at about somite XXII. The annulation of the three anal somites XXV-XXVII varies considerably, with annular widths decreasing towards the posterior end. Somites XXV and XXVI are biannulate and reduced ventrally. Somite X XVII may be uniannulate or biannulate. If the latter, the last annulus of the somite is generally unrecognizable ventrally. The anus is situated dorsally in the furrow between somite XXV and XXVI. Annulation and tuberculation vary towards the extremities of the body, where there is some ventral curvature. TUBERCULATION (Text-fig. 16). Harding (1924 and 1927) gave the normal number of tubercles on a, and a; as fourteen, six dorsal, six ventral and a pair lateral, but in 414 LC. ELEWEEL YN my specimens the more usual number was twelve, six dorsal and six ventral, with lateral tubercles occurring rather irregularly. The number varied from ten to fourteen, a, tending to have fewer than a,. The first two annuli of the neck region were extremely small and tubercles were never obvious on their ventral surfaces. In one specimen there were only five tubercles, instead of the normal six, on the dorsal surfaces of a, and a, annuli in the preclitellar region. The tuberculation of the clitellar constriction varied considerably and the tubercles were small and irregularly placed. Xaz, XIa, and XIIIa, were often free of tubercles, but occasionally XIa, bore a minute pair dorsally. Most of the remaining annuli of the clitellar constriction bore between twelve and fourteen tubercles per annulus. Tuberculation of a typical trimeric somite is shown in Text- fig. 14D, the larger tubercles being dorsal in position. Small tubercles were present dorsally on the last annulus of somite X XVII, which is generally indistinguishable ventrally. In many of the specimens each annulus possessed between four and seven striae dividing it up into smaller rings (Text-fig. 14c), but these were variable and absent in some specimens, particularly the smaller ones. Selensky (1915), in discussing the annulation of some Pontobdellinae, stated that the somite is divided into fourteen or fifteen secondary annuli. Out of the six species which I have examined, including nearly one hundred specimens, Stibarobdella loricata is the only species showing such subdivisions. Moreover, their presence depends on whether the specimen is gorged or starved, whereas true annuli are more permanent. Minute tubercles are never found on these small rings and a large tubercle may cover four or five of them. The H.R. P.C.R. C.R. eRe (ieee Pen ai Somites, xl XIII XIV j Dorsal. ptelelelalelelelslolaialelalem Ventral. HME Amor np tela Size !-3. 2)2 Dili sheesace Somites. Dorsal. Ventral. Size |-3. Fic. 16. Diagram illustrating the tuberculation of a single specimen of Stibavobdella loricaia, anterior extremity above, posterior below. The numbers in brackets in this diagram indicate the presence of small secondary tubercles which are generally situated para- marginally. ‘‘ }’’ in the size column indicates the presence of an annulus dorsally only. See Fig. 13 (p. 409) for other abbreviations. REVIEW OF TE PON TOBDELLINAE 415 tubercles (Text-fig. r4ca and 6b) are generally quite prominent and bear sensillae, which are usually concentrated at the summits but may be present over the whole tubercle. DISTRIBUTION. Off Cabinda, West Coast of Central Africa ; West Coast of Mexico and Peru; India; Sierra Leone and Philippine Islands. Hosts. Unrecorded but the wide distribution suggests oceanic sharks. Stibarobdella bimaculata (Oka, 1910) Pontobdella bimaculata Oka, 1910 : 171. DESCRIPTION. General characters. One specimen examined. This species is very close to Stibarobdella tasmanica (p. 420), but may provisionally be regarded as separate. Only one specimen was found in the British Museum collection and this resembled Oka’s (1910 and 1927) description very closely. Harant (1929) seems to be wrong in assuming this to be a quadriannulate form. In this specimen the body is elongated, distended, flattened and brown, but in Oka’s it was cylindrical and yellow. The specimen was 3 cm. long and 5 mm. wide. The anterior sucker (Text-figs. 17A and 18) is circular, attached eccentrically, and separated from the body byaneck. The diameter of this sucker in the contracted state is slightly broader than the preclitellar region, but according to Oka it may be nearly twice as broad when expanded. It carries three pairs of marginal elongated papillae, a smaller pair situated near the base of the sucker, and a pair of large rectan- gular eye patches of a dark reddish brown colour. A few concentric grooves can be seen at the base of the sucker. The posterior sucker is bell-shaped, scarcely larger than the anterior sucker and separated from the body by a constriction. ANNULATION. Uniannulate somites XXVII (Text-fig. 18) Biannulate somites XXV, XXVI ) € ? Triannulate somites VIII-X, XIII-XXIV \xr, aE: The posterior limit of the neck is doubtful in this species, since Oka (1927) called the third and fourth annulus a double annulus because they were usually poorly separated. But it seems doubtful whether both these two annuli should be regarded as lying in the neck region, making a total of four annuli there, for that would involve regarding the first preclitellar somite VII as biannulate, with the anterior annulus the largest, whereas it is triannulate in other Stibarobdella. The other two preclitellar ~ somites VIII and IX are triannulate with their annular widths 1: 2:1. Of the annuli in the clitellar constriction, Xa, is very small. Somites XI and XII may be regarded as biannulate, with the anterior annulus double, forming a smaller anterior ring, which can scarcely be regarded as a separate annulus. The male pore is situated mid-ventrally in the furrow between the two major annuli of somite XI, while the female pore is similarly situated in somite XII. Annulus XIIIa, is very small. Somites XIV to XXIV are all triannulate, with annular widths in the ratio 2:3:2. The outline of the nerve chain could be made out on the ventral surface, with the ganglia on the a, annulus of each somite. Somites XXV and XXVI are 416 L. €. LLEWELLYN biannulate, with the anus situated dorsally on the a, annulus of somite XXVI. Somite XXVII is very small and indistinguishable ventrally, but according to Oka (1927) may be double. TUBERCULATION (Text-fig.19). In the clitellar constriction, annuli Xa, and XIIIa, possess no tubercles and all the other tubercles of this region are small. The tuber- culation on somites VIII, IX and XIV—XXIV is fairly constant (see also Text-fig. 17c). The a, annulus of a typical somite has four large dorsal tubercles and four smaller ventral ones, and the a, and a, annuli generally have six dorsal and four slightly smaller ventral ones. The tubercles are conical, generally with between six and ten apical sensillae. gn: ' i Sc. ° e) Oo @) e) re) ° <0, i a ° ° ° ° ° ° ° ° ° ° w Fic.17. Stibarobdella bimaculata, a. Section through anterior sucker, showing oral surface. B. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. c. Tuberculation of a typical triannulate somite split mid-ventrally. g7., outline of gang- lion on ventral surface; ¢u., tubercle; See Fig. rz (p. 406) for other abbreviations. 1 Fie. 18. REVIEW OF THE PONTOBDELLINAE Dorsal view of Stibavobdella bimaculata showing annulation, tuberculation and suckers. See Fig. 12 (p. 407) for abbreviations. 417 418 L. C. LLEWELLYN REMARKS. Both Stibarobdella bimaculata and S. moorei (p. 422) differ from S. macrothela in having the posterior sucker only slightly wider than the anterior sucker and only eight tubercles on the ag annulus. Their tubercles are also very different in appearance. They should not be regarded as synonymous as suggested by Cordero (1937-38). DistRIBuTION. Pearl Island, Panama ; Hondo (Coast of Sagami Awa). Host. Sharks. Stibarobdella planodiscus (Baird, 1869) Pontobdella planodiscus Baird, 1869 : 312. Pontobdella variegata Baird, 1869 : 313. DESCRIPTION. General characters. Three specimens were examined, 40-55 mm. in length and 5-7 mm. in breadth. The body is elongated, yellowish and cylindrical or flattened. Baird (1869) named the cylindrical specimens Pontobdella variegata, but in fact they seem to be the same species as the flattened one, which he described as P. planodiscus and which has page priority. The anterior sucker (Text-fig. 204) is circular and attached eccentrically with its dorsal surface elongated. It is less than twice as broad as the neck at its narrowest point. It bears four pairs of papillae, of which the ventral pair are very small and often difficult to see, and no marginal fringe. The posterior sucker when contracted is equal in width to its point of attachment and separated from the body only by a shallow groove, but when expanded is half Somites. Dorsal. BAO OOoooOoooodG Ventral potas tafe tet state tate [a te| 3} 2 2 Size I-3. Somites. XXIII XXTV XXV XXVI] pXXVil 1 Dorsal. fa[s[ela[7]4{4]4]2]oli ' Ventral. ES ea RSE lel : 4 Size |-3. Fic. 19. Diagram illustrating the tuberculation of a single specimen of Siibarobdella bimaculata, anterior extremity above, posterior below. ? indicates where the limits of the somites are uncertain. ‘‘}’’ in the size column indicates the presence of an annulus dorsally only. See Fig. 13 (p. 409) for other abbreviations. REVIEW OF THE PONTOBDELLINAE 419 as broad again (Text-fig. 20s). The anterior : posterior sucker ratio when contracted 1S 10 35 Ge oy ANNULATION. Uniannulate somites XXVI, XXVII. Biannulate somites XI, XII, XXV. Triannulate somites VII-X, XIII-XXIV. The three preclitellar somites VII-IX are triannulate, with annular widths in the ratio 2:3:2. The clitellar constriction (Text-fig. 20c) is made up of small annuli with somites XI and XII biannulate. The male and female pores lie in the grooves between the two annuli of somites XI and XII respectively. Somites XIV to XXIV are all triannulate, with their annular widths in the ratio 2:3:2. The first anal somite X XV is biannulate, with the anus situated mid-dorsally on the first annulus. Somites XXVI and XXVII consist of single small annuli which are reduced further ventrally. m.v. 1 m.d. | mv. bio ©: .O.O.© (O30) © OOo o70 6s me OPO OC) O-O 00 ts DMO MOLOrtO). ©) © 20). 21O..O: 0. G4, D Fic. 20. Stibarvobdella planodiscus. A. Anterior sucker. 8. Posterior sucker. c. Dorsal surface of clitellar region. pb. Tuberculation of a typical triannulate somite split mid- ventrally. an., anus; a.s., anterior sucker; g.f.p., groove in which female pore is situated ; g.m.p., groove in which male pore is situated ; m.v.t., mid-ventral tubercle ; p.s., posterior sucker ; fw., tubercle. See Fig. 11 (p. 406) for other abbreviations. 420 TiC? Se WEE Te Yon TUBERCULATION. The normal triannulate somite (Text-fig. 20D) has nine tubercles on the a, annulus, four large tubercles situated dorsally and five smaller ventrally, of which one is mid-ventral. The a, annulus possesses between fourteen and sixteen tubercles and the a, annulus generally eleven. The ventral tubercles in both cases — are smallest. Tubercles on annuli in the clitellar constriction are much smaller, more irregular in number and absent from the XIIIa, annulus. Tubercles are also small and irregular on the last three annul of the anal region and are often indistinguishable on the ventral side of the annulus of somite XX VII. The tubercles are mammiform, with sensillae at their summit. REMARKS. It appears that the two species described by Baird (1869) as Pontob- della variegata and Pontobdella planodiscus are the same. According to Baird the major differences were that P. planodiscus had a flattened body, apical sensillae on the tubercles and three pairs of papillae on the anterior sucker, while P. variegata had a cylindrical body, rounded tubercles and no papillae. However, examination of his material, including the cotypes, showed that the flattening of P. planodiscus was similar to that seen as a preservation artefact in otherspecies andthat the anterior sucker had three pairs of large papillae and one pair of small ones. Three pairs of retracted papillae were also discernible on P. variegata. Since the presence of sensillae at the summit of tubercles is also highly variable, there seems from the material available to be no reason for regarding these as separate species. The sucker characters and mid-ventral tubercles of St:barobdella planodiscus are similar to those of the quadriannulate form Pontobdella vosmaert, indicating parallel evolution. DISTRIBUTION. Straits of Magellan ; Possession Bay, Patagonia. Host. Not recorded. Stibarobdella tasmanica (Hickman, 1946) Pontobdella tasmanica Hickman, 1946:27, nom. nov. pro Pontobdella verrucosa Hickman, 1941 : 41 (non Hivudo verrucosa Fleming, 1811 : 245). DESCRIPTION. General characters. (Specimens not examined. Apparently originally described from a single specimen, although Ingram (1957) described further specimens.) Body 37 mm. long, fusiform, circular in section and narrowing anteriorly. Colour light brown, becoming mustard yellow on preservation. Single dark brown triangular patch on the dorsal side of the anterior sucker. Large tuber- cles tipped with white and apparently mammiform. Anterior sucker 2°26 mm. in diameter, cup-shaped, eccentrically attached and with four pairs of submarginal papillae and twelve radial furrows on its oral surface. No eyes. Posterior sucker cup-shaped, 3°54 mm. in diameter, centrally attached and slightly wider than the greatest width of the body. Anterior : posterior sucker ratio between I: I and I: 2 ANNULATION. Uniannulate somites XXVIT rarty Ser, (SOMULeS nuio bene Biannulate somites X—NII KAV, LAVI- as in S. macrothela, Triannulate somites VII-IX, NIJTI-XNNIV. Text-fig. 12.) | | | | REVIEW OF THE PONTOBDELLINAE 421 Somites of the testicular and caecal regions have annular widths in the ratio 2:3:2. Clitellar constriction between a, annuli of somites X and XIII, consisting of five narrow rings. Male pore originally described as lying in furrow between somite XI and XII, but according to Ingram (1957) opening between XIa, and as, suggesting that this somite is triannulate. Female pore in furrow between somite XII and XIII. Somite XXV divided into two annuli only ventrally and somite X XVI divided into two only dorsally. Anus near posterior edge of annulus XXVIa,. TUBERCULATION. A, annuli each possess four large dorsal tubercles and four smaller ventral ones. A, and a, annuli have six dorsal and six ventral tubercles with the two dorso-median tubercles smallest, and a ventro-median tubercle often present. Interposed tubercles common. DISTRIBUTION. Sandy Bay, Hobart; Kingston, Jamaica; Brig Rocks, King Island. Host. Skate (Ingram, 1957). Stibarobdella taprobanensis (de Silva, 1963) Pontobdella taprobanensis de Silva, 19636 : 39. DESCRIPTION. General characters. (Specimens not examined.) Body up to 29 mm. long (including suckers). Preclitellar region translucent (enabling underlying organs to be seen). Rest of body yellowish brown, with pairs of brown spots dorsally on testicular and caecal region. Anterior sucker deeply cupped and slightly wider than preclitellar region, with dorsal margin three times the length of ventral margin, incised, shovel shaped, and bearing dorsally a pair of vermilion coloured blotches which meet anteriorly and two pairs of eyes, but no papillae. Anterior : posterior sucker ratio between 1: 1-3 and 1:2. Posterior sucker with dark radial bands and a smooth margin, oval, sometimes folded dorso-ventrally, eccentrically attached, so that the dorsal portion is twice the ventral portion, and equal to or less than the maximum diameter of the body. ANNULATION AND TUBERCULATION. Uniannulate somites (XXV-XXVII)? Biannulate somites XI, XII. Triannulate somites VII-X, XIII-XXIV. (Somites numbered as in S. macrothela, Text-fig. 12.) No true tubercles, but only slightly protuberant opaque patches, segmentally arranged. Two small neck annuli. Preclitellar annuli VII—X triannulate, a, annuli largest and possessing two opaque patches dorso-laterally and two ventro- laterally, composed of groups of small opaque spots, which are distributed sparsely elsewhere. Annuli a, and a; possess a pair of opaque patches dorsally only. Pre- clitellar constriction in usual position, between Xa, and XIIIa,. Somite XI and XII without opaque patches. Male pore mid-ventral, in the middle of somite XI. Female pore mid-ventral, in middle third of somite XII. A, annuli of somites XIII- XXIV with two pairs of opaque patches (rosettes) dorsally, situated para-marginally ZOOL. 14, 7. 29 422 IEC. ACTS ES Webs Iva and para-medially, and composed of two to six triangular opaque areas. Mid- dorsal ellipsoidal umber coloured clear areas on all annuli. No rosettes ventrally or on somites posterior to somite XXIV. Anus mid-dorsal between somites XXVI and — XXVII. DISTRIBUTION. Wadge Bank, Ceylon. Host. Unknown. Stibarobdella moorei (Oka, I9I0) Pontobdella moovet Oka, 1910: 171. Stibavobdella moorei : Harant, 1929: 651. DESCRIPTION. General characters. (Specimens not examined.) Body up to 14 cm. long and 14 mm. broad, fusiform, yellowish and thickest in the middle of the posterior half. Anterior sucker smooth, hemispherical, rather large, and without fringe, eyes, papillae or annular marks. Posterior sucker bell-shaped, scarcely larger than anterior sucker and separated from the body by a constriction. ANNULATION. Uniannulate somites—none. Biannulate somites XI, XII, XXV-XXVII. (Somite numbering as Triannulate somites VII-X, XIIJI-XXIV. in S. macrothela, Text-fig. 12.) Triannulate somites have annular widths in the ratio 3:4:3. Neck of three annuli, the posterior largest and the anterior two often hidden by sucker. Pre- clitellar region of three triannulate somites. Clitellar constriction as usual between a, annuli of somites X and XIII. Somites XI and XII biannulate and bearing between their annuli the male and female pores respectively. Somites XXV—XXVII biannulate, becoming smaller posteriorly. Anus on a, or between a, and a, of somite XXVI. TUBERCULATION. A, annuli with four prominent tubercles dorsally and four ventrally, each bearing 7—I10 apical sensillae. A, and ag annuli with ten smaller tubercles, four dorsally, four ventrally and two laterally. Tubercles on clitellar constriction are smaller still and absent from annulus Xa, and sometimes from XITTa,. DISTRIBUTION. Hondo (Coast of Sagami, Awa). Coast of Tokyo. Host. Shark. Stibarobdella australiensis (Goddard, 1909) Pontobdella austvaliensis Goddard, 1909 : 724. DESCRIPTION. General characters. (Specimens not examined.) Body up to 20 mm. long and 3 mm. broad, attenuated anteriorly and circular in cross-section, extremities yellowish brown, the rest bluish grey. Anterior sucker I mm. in diameter, with four or five pairs of papillae (the posterior pair inconspicuous or absent), two faint annuli on dorsal surface and no fringe or pigmented markings. Posterior sucker I°5 mm. in diameter, with five or six annuli on dorsal surface. Anterior: posterior sucker ratio I: 1°5. tl REVIEW OF THE PONTOBDELLINAE 423 ANNULATION. Uniannulate somites Biannulate somites XI, XXV, San XXIT, XXVIT (Somite numbering as for S. macrothela, Text- Triannulate somites VII-X, XIII-XXIV fig. 12.) In triannulate somites a, is slightly larger than annuli a, and a, and possesses more prominent tubercles. TUBERCULATION. Tubercles largest in the testicular and caecal regions, where the a, annuli each bear six prominent conical papillae, four dorsal, situated para-medially and para-marginally, and two ventral, situated para-marginally. These ventral tubercles are smaller and similar in size to tubercles of a, and ag annuli. Dorsal tubercles are larger than a, and a, tubercles. DISTRIBUTION. Not recorded. Host. Not recorded. Genus PONTOBDELLA Leach, 1815 Pontobdella Leach, 1815: 9. TYPE SPECIES. Hirudo muricata Linnaeus. D1iAGnosis. Somites of the testicular and caecal regions tetrameric. Pontobdella muricata (Linnaeus, 1758) Hirudo muricata Linnaeus, 1758 : 650. Hirudo verrucosa Fleming, 1811 : 245. Pontobdella verrucata Leach, 1815: II. Pontobdella aveolata Leach, 1815: 10. Pontobdella spinulosa Leach, 1815 : 12. Pontobdella muricata: de Blainville, 1818: 293; Harding, 1910: 143. Pontobdella laevis de Blainville, 1827 : 243. Pontobdella verrucosa : Leydig, 1851 : 318. DESCRIPTION. General characters. (About fifty specimens examined.) The body is club-shaped, generally circular in cross-section or slightly flattened dorso-ventrally in some starved specimens, and up to 19°0 cm. long (excluding suckers). The colour varies from dull yellow to olive green or occasionally pinkish. The cup-shaped anterior sucker (Text-fig. 2rA and 22) is between two and three times the diameter of the neck and is generally as large as or larger than the posterior sucker. It is circular and fixed eccentrically, so that the dorsal surface is longer than the ventral. Around its edge is a very noticeable fringe, bearing three pairs of lateral papillae, which are sometimes retracted and obscure. The sucker may be somewhat flattened laterally and then appears to have a slit-like opening. The posterior sucker is cup-shaped when contracted, centrally fixed and generally just broader than its point of attachment, although considerably larger when ex- panded. It never exceeds the maximum diameter of the body. ZOOL. I4, 7. 29§ 424 LE. CC. DEEWELLYN ANNULATION. Uniannulate somites — Biannulate somites XI, XII, XXV, XXVI, XXVII. Triannulate somites VII-X. Text-fig. 22. Quadriannulate somites XITI-XXIV. The first three annuli constitute the neck and the anterior two of these are much smaller than the other. The three preclitellar somites VII-IX are all triannulate with their annular and tubercular sizes in the ratio: 2:1. The clitellar constriction includes the small annuli Xa, and XIIIa,, between which are from four to six annuli, Fic. B. Cc. 21. #0 30 400 pOhe is @l#0%2 (Oy Os = wings) 4 Os On Our Ee yO” MOR (ON norm =O) Gre O Ole! eu OuRCt Borge Re (ito OL atiol ko ~3PcOke 6 ° ° ° SEES D Pontobdella muricata. A. Section through anterior sucker, showing oral surface. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. a, band c: Tubercles in various stages of contraction. pb. Tuberculation of a typical quadriannulate somite split mid-ventrally. For designation of annuli, i.e. a,, aj, b; and b,, see Mann 1953 ; f., fringe; fz., tubercles ; See Fig. rr (p. 406) for other abbreviations. 6 953i. J 8 Ss 4 Fic. REVIEW OF THE PONTOBDELLINAE 22. Dorsal view of Pontobdella muricata showing annulation, tuberculation and suckers. f., fringe; See Fig. 12 (p. 407) for other abbreviations. 4 3D 420 ES CG: LE EWELEY N two or three per somite (see dotted lines in Fig. 21p). The appearance of these extra subdivisions may depend on the stage of gorging or contraction of the specimen, The male pore is situated in the furrow between a, and ay, or a, and az of somites XI (depending upon whether this somite is bi- or triannulate), while the female pore is situated between a, and a, of somite XII. Somites XIII to XXIV are all quadriannulate and their annular widths and tubercular sizes vary in the ratio of 2 AROnuAy ie, 2 The annulation of the three anal somites XXV—X XVII is rather indistinct, but it appears that there are two annuli per somite, decreasing in size posteriorly. The last two small annuli are often unrecognizable and nearly always absent from the ventral surface. The anus is mid-dorsal in somite XXVI, either between the two annuli, or in the middle of the anterior annulus. TUBERCULATION. Text-fig. 23 shows the arrangement of tubercles towards the anterior and posterior extremities of the body. Text-fig. 21D shows the tuberculation of a typical quadriannulate somite. Dorsal tubercles tend to be larger than ventral ones. The a, annulus has eight primary tubercles, between which may lie 0-6 secondary tubercles. The a, annulus has generally about twelve primary tubercles and up to six secondary tubercles. The b; annulus has about ten primary tubercles and up to six secondary tubercles, and the b, annulus about fourteen small tubercles. The tubercles on the first two small annuli of the neck region are much reduced and often absent from the ventral surface. In the clitellar constriction the tubercles are much reduced and irregular in arrangement. Annuli Xa, and XIIIa, often have H.R. P.C.R. C.R. T.R. Somites. Dorsal. alele|s[elelalelalale]a [ata [a]a| 1 (aialalsl oe Ventral. EEE Size 1-3. Somites. XXIII XXIV XXXVI) XXVII rs[aT Tal s]eTo balls [Tale or Dorsal. Ventral. eee a a a aa a SS Size |-3. 2/3] 2 2/3] 2 fr Fic. 23. Diagram illustrating the tuberculation of a single specimen of Porfobdella muri- cata, anterior extremity above, posterior below. The numbers in brackets in this diagram indicate a regular increase in numbers of tubercles found in this region in other specimens. Secondary tubercles are not indicated here because of their irregularity. “‘}’’ in the size column indicates the presence of a small annulus dorsally only. See Fig. 13 (p. 409) for other abbreviations. REVIEW OF THE PONTOBDELLINAE 427 no tubercles, and when present these are more usually found dorsally. In the anal region too, tubercles are few or absent on the ventral surfaces of the annuli. ReMARKS. Harding (1910) observed that the shape of the tubercles can vary considerably depending on whether the leech is relaxed, gorged or otherwise (Fig. 21ca, b,c). According to Selensky (1915), muscle fibres can pull down the tips of the tubercles, so that the skin becomes smooth. This has led to considerable confusion and the erection of numerous invalid species. Pontobdella muricata was described as having tubercles possessing a rosette of sensillae on their summit, P. verrucata as having mammiform tubercles without a rosette of sensillae, P. aveolata as having tubercles forming low irregular prominences and P. laevis as having entirely retracted tubercles. It is possible, however, that P. /aevis is not one of the Pontobdellinae (Moquin-Tandon, 1846). Pontobdella verrucosa, according to Moquin-Tandon (1846) was distinguished by each quadriannulate somite having one large annulus and three small annuli, com- pared with three large annuli and one small in P. muricata. There are no grounds for the former species, since in fact P. muricata somites usually have one large annulus ay, two medium sized annuli a, and b;, and one small annulus bg. DISTRIBUTION. Mediterranean Sea, North Sea and most waters in north-east Atlantic. Hosts. Many species of rays and sometimes plaice. Pontobdella vosmaeri Apathy, 1888. Pontobdella vosmaeri Apathy, 1888 : 59. Pontobdella bvumpti Riviere, 1925 : 292. DESCRIPTION. General characters. (Five specimens examined.) The body is club-shaped, circular in cross-section and may reach a length of 7°5 cm. (excluding suckers). The colour is dull yellow and usually uniform. The neck region is only slightly smaller than the diameter of the anterior sucker, the ratio in size being about I:1-3. The anterior posterior sucker ratio is about 1:2. The anterior sucker (Text-fig. 244 and 25) is circular, cup-shaped and fixed eccentrically, so that the dorsal surface is somewhat larger than the ventral. The margin is without a fringe but generally bears three pairs of papillae. The posterior sucker is cup-shaped when contracted and fixed to the body centrally. It is generally slightly thicker than the body at its point of attachment, although when expanded it may be considerably larger, but never exceeding the maximum diameter of the body. ANNULATION. Uniannulate somites XXVI, XXVII. Biannulate somites De OIE XV. VU Triannulate somites VIII, X ; Quadriannulate somites Gea EX. The dorsal surface of the anterior sucker is sometimes faintly marked by three transverse furrows. The neck region includes the first three annuli, the anterior two of which are much smaller than the other. The annulation of the three pre- Text-fig..25. 428 Loc. LLEWELLYN clitellar somites is quite different from Pontobdella muricata. Somite VII is generally biannulate, but occasionally there is a much reduced anterior annulus. Somite VIII is triannulate with the annular widths in the ratio of 1: 3:2; whilst somite IX, although generally divided by two furrows to form three annuli with widths in the ratio 2:3: 2, has an extra row of tubercles on the aj annulus, suggesting incipient subdivision. Somite X (Text-fig. 24B) is triannulate with its annular widths in the ratio of 2: 3:1, the Xa, annulus being the first annulus of the clitellar constriction, Somites XI and XII are both biannulate and the annuli are generally of equal size. The male pore is situated mid-ventrally in the furrow between the two annuli of somite XI, while the female pore is similarly situated just posterior to the furrow dividing the two annuli of somite XII. Annulus XII1a,;, which is small and sometimes difficult to see, is the last of the clitellar constriction. Somite XIII is quadriannulate, the ratio of widths being 1:3:2:1. Somites XIV to XXIV are also tetra-annulate, s.@. A P- \\ \ t t r 1 m.v.¢, ——? ° ° ‘Or re ° ° fe) ° ° qa, rT ' So — HERG ° oe Oo ° Cymr-) ° ° ° ° 1 de ' ! tere o 8° © © ° ° eo 0 ° ° ° ° tbe Fic. 24. Pontobdella vosmaervi. A. Section through anterior sucker, showing oral surface. B. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. c. Tuberculation of typical quadriannulate somite split mid-ventrally. m.v.¢., mid-ventral tubercle ; ¢w., tubercle; See Fig. rr (p. 406) for other abbreviations. REVIEW OF THE PONTOBDELLINAE Hig. 25. Dorsal view of Pontobdella vosmaeri showing annulation, tuberculation and suckers. See Fig. 12 (p. 407) for abbreviations. 430 LOC. LEE WELL VN but with the ratio of widths and tubercular sizes 2:3:2:1. The anal somites XXV-XXVII are much reduced. Somite XXV contains two annuli of which the posterior one is largest. Both somite XXVI and XXVII appear to consist of a single annulus each, the annuli being practically absent from the ventral surface. The anus occurs either in the furrow between somite XXV and X XVI, or near the posterior edge of the a, annulus of somite XXV. TUBERCULATION (Text-fig. 26). The dorsal surface of the anterior sucker is sometimes marked by faint grooves, between which may occur seven to eight extremely small papillae. Tubercles on the first two annuli of the neck region are much reduced and generally absent from the ventral surface. Annulus IXa, has a double row of tubercles, in which the posterior row is by far the smallest. According to Riviere (1925) annulus Xag, the first annulus of the clitellar region, is non-tubercu- late, but in some specimens it bears tubercles dorsally. On the other hand somite XIIlai was according to Riviere tuberculate, but in some specimens it is non-tuber- culate. As these annuli are small it may be expected that their tuberculation will vary. Text-fig. 24c shows the tuberculation of a typical quadriannulate somite. Although the secondary tubercles on the a, annulus are subject to variation, the number of primary tubercles is constant. This species differs from Pontobdella muricata in having nine primary tubercles on the a, annulus, including one situated mid-ventrally. The a, annulus possesses about fourteen primary tubercles and a varying number of secondary tubercles, while b; has about twelve primary tubercles Somites. Dorsal. Ventral. Size 1-3. Somites. Dorsal. Ventral. Size 1-3. Fic. 26. Diagram illustrating the tuberculation of a single specimen of Ponfobdella vosmaeri, anterior extremity above, posterior below. A.s., region of anterior sucker. The numbers in brackets in this diagram indicate a regular increase in numbers of tubercles 5 5 found in this region in other specimens. Secondary tubercles are not indicated here 5 - : 3 because of their irregularity. ‘“‘}’’ in the size column indicates the presence of small =) 9 2S . annulus dorsally only. Dotted lines indicate where additional annuli may or may not be present. See Fig. 13 (p. 409) for other abbreviations. REVIEW OF THE PONTOBDELLINAE 431 and up to six secondary ones, rarely having a total of more than fifteen. Annulus b, has about fourteen small tubercles. The large tubercles are generally conical, with up to eight apical sensillae, whereas small tubercles generally lack sensillae. REMARKS. Herter (1935) suggested that Pontobdella brumpti Riviere (1925) appears to be synonymous with Pontobdella vosmaert Apathy (1888). Although Riviere referred to Apathy, he did not mention the brief but adequate description of P. vosmaert, which is placed in an addendum to the paper. It seems that Riviere overlooked this description, since he referred only to a brief description given by Blanchard (1893). DISTRIBUTION. Plymouth, Roscoff, Capri. Host. Rays? Pontobdella aculeata Harding, 1924 Pontobdella aculeata Harding, 1924: 491. DESCRIPTION. General characters. (Specimens not examined.) Body fusiform, attenuated anteriorly and circular in cross-section. Length 35-64 mm., breadth 6-8 mm. Colour dull grey to reddish brown, occasionally with linear spots on the a, annulus of each somite. Anterior sucker attached eccentrically, small, circular, with a corrugated edge, without papillae and apparently with no fringe. Anterior : posterior sucker ratio 1: I or former slightly larger. Posterior sucker circular, with corrugated edge and attached centrally. Somites. Dorsal. "aI 08 aces EATEN CE EMCACAES PANERA RAIS Ventral. | fofo|2[of2|2]o[2[2[2 [2] 2/2 EARIRIEARIIEIRIEI Lied Size 1-3. Somites. XXIII Dorsal. HRA Aan AnRE Ventral. == 4 Size |-3. rm Fic. 27. Diagram illustrating the tuberculation of a single specimen of Pontobdella aculeata, anterior extremity above, posterior below. Compiled from the details in Harding (1924 and 1927). A.S., region of anterior sucker; p.s., region of posterior sucker. Small number in top right hand corner of square, with a black dot, denotes number of brown spots on dorsal surface of leech. See Fig. 13 (p. 409) for other abbreviations. 432 L. iC LEEWELLYN ANNULATION. Uniannulate somites XXVI, XXVIII. Biannulate somites XXV. Triannulate somites Ville VA eile xe Quadriannulate somites IX, X, XIII-XXIV. Annuli of quadriannulate somites approximately equal in width but becoming smaller towards the extremities of body. Dorsal surface of anterior sucker may possess faint furrows. Neck possesses three annuli with ratio of widths from anterior end r: 1:2. Clitellum inconspicuous, probably comprising annuli Xb, to XIIIa,. Male pore opens mid-ventrally in furrow between XIa, and XIa,;. Female pore opens mid-ventrally in furrow between XIIa, and XIIa;. Anus opens mid-dorsally on the annulus of somite XXVI. TUBERCULATION. Text-fig. 27 shows tuberculation near the anterior and posterior extremities, derived from details given by Harding (1924 and 1927). All annuli of a quadriannulate somite possess lateral tubercles with the tubercles on b; largest. In addition, on the dorsal surface, the a, and b,; annuli possess a paramedian pair, whilst the a, and b, annuli possess a para-marginal pair and a median tubercle. Ventrally there is only a para-marginal pair present on all annuli. DISTRIBUTION. Bassein River, Burma; Gregory Isles, Mergui Archipelago, Burma ; Wadge Bank, Ceylon (de Silva), 19634). Host. Teleost Harpodon nehereus. (This fish is found in fresh water and estuaries as well as the sea.) Pontobdella rugosa Moore, 1938 Pontobdella vugosa Moore, 1938 : 5. DESCRIPTION. General characters. (No specimens examined.) Body slender, tapering gradually from the anterior to posterior end, length about 40°5 mm., maximum breadth 2°3 mm. and slightly flattened. Striking colour pattern meta- merically blotched and annulated, with ground colour of ferruginous brown and pale yellow markings or vice-versa. The pattern varies considerably. Anterior sucker sub-hemispherical, eccentrically attached so that oral surface faces ventrally, bearing a fringe (welt according to Moore), with fifty-two small papillae, two circles of sub-marginal papillae, and marked postero-dorsally by faint annulations, with rows of small papillae and pigment spots with vestigial eyes. Mouth situated centrally. Anterior: posterior sucker ratio 1:1°7. Posterior sucker cup-shaped when contracted, generally not exceeding the diameter of the body. Margin furrowed, cavity of sucker shallow, and externally rayed with brown rays. ANNULATION. Uniannulate somites XXVI, XXVII. Biannulate somites AXXV : XT Triannulate somites (VI?) XI eee xxiv Quadriannulate somites NIJI-XNIII ,, = vie & : VII-X Sexannulate somites i REVIEW-OF THE PONTOBDELLINAE 433 Neck made up of four annuli, the first being small and reduced ventrally, the other three equal in size. Somites VII-X quadriannulate, with a, and a, tending to be biannulate. This tendency becomes more prominent posteriorly, as does the com- plexity and size of the somites. Clitellum ill-defined, probably extending from Xb, to XIIIa,. Somite XI triannulate with annular size becoming slightly smaller posteriorly. Somite XII bi- or triannulate. Male pore on annulus XIa, and female pore in furrow between annuli XIla, and XIla, or as far forward as middle of annulus XIla, (i.e. in furrow between XIIb, and XIIb,). Somites XIII to XXIII are quadriannulate and the annular sizes are usually a, > a, > b;—b,. In somite XXIV the furrow between b; and bg is often indistinct. Somite XXV is biannulate, with the anterior annulus largest. Anus mid-dorsal in furrow between somite XXVI and XXVII. TUBERCULATION. Text-fig. 28 shows tuberculation of the anterior and posterior regions from details given by Moore (1938), but not including small secondary tubercles, although they are said to be present in varying numbers. Tubercles are generally largest dorsally, Particularly on a; annuli. They are small or absent on neck annuli, somite VII, the clitellar constriction (Xb,—XIIIaj,), the anal region and b, of somites VIII-X. There are no tubercles on the small indistinct b, and b, annuli of somites VII-X. Otherwise dorsally, on all a,, b; and be annuli, there are two pairs of tubercles, outer paramedians and supra-marginals, while on a, annuli there are a lateral pair of tubercles and a pair near the inter- mediate line. Ventrally, a,, b; and b, bear two pairs of (para-median and sub- marginal) tubercles, while a, bears a single pair on the intermediate lines. The tubercles are large, pointed, conical and with one or more apical papillae. DISTRIBUTION. Commonwealth Bay, King George V Land; 66° 32’ S., 141° 39’ E., 157 fathoms ; 65° 42’ S., 92° 60’ E., 60 fathoms. Host. Not recorded. Genus PENTABDELLA gen. nov. TYPE SPECIES. Pontobdella dispar Cordero, 1937. Diacnosis. Tuberculate Piscicolidae in which most of the somites of the testicular and caecal regions are pentameric. Pentabdella dispar (Cordero, 1937) Pontobdella dispar Cordero, 1937 : 13. DESCRIPTION. General characters. The following description is based on that by Cordero. The body is fusiform, elongated, of circular section and a uniform pale yellow. It may be up to 160 mm. long (including suckers) and 16 mm. wide (about annulus XXIa,). Anterior sucker hemispherical, smooth, convex externally and somewhat flattened laterally, with about five faint annulations near its base dorsally. 434 L.' OC. ALBEE WELL YN It possesses a marginal fringe bearing three pairs of papillae. The internal surface is concave and smooth. The anterior : posterior sucker ratio varies between I : 0-8— I: 1:6, presumably depending upon contraction. The posterior sucker is constricted at the base, convex and smooth externally, with a thick margin but no fringe. It does not generally exceed the maximum diameter of the body. ANNULATION. Uniannulate somites XXVII XXVI Biannulate somites XXV i Triannulate somites VII-XII, XXIV. Quadriannulate somites XIII. Ouinqueannulate somites XIV—-XXIII. The neck is made up of three annuli, of which the first is smallest while the other two increase in size posteriorly. The preclitellar somites are triannulate, increasing in size posteriorly with the central annulus largest. The first two annuli of somite X are similar to annuli in the preclitellar region, but annulus Xa, is smooth and much thinner, being the first annulus of the clitellar constriction. This is composed of eight annuli, including Xa, and XIIIa,, all uniformly small. The male pore opens between annulus XIa, and XIas, and the female pore betwen XIla, and XIlIas3. Annulus XIIa, is sometimes unrecognizable ventrally. The abdomen, from somite XIII to XXIV, is five times the length of the preclitellar and clitellar regions com- bined. Somite XIII is quadriannulate, with a, the largest annulus. Somites XIV to XXIII are quinqueannulate, with the ratios of the annuli b,, by, a, b; and b, approximately 1:4:6:4:1. Somite XXIV consists of three annuli which are Somites. Dorsal. Ventral, Size |-3. B.G.R. A.R. iF SAREE Somites. XxXttl XXIV XXV oy XXXVI pox Dorsal. lofsts]pfolels[ele]s [4] 3 [3 | Ventral. lof4]4]4fo]4[4][4[4]4[4/3]3| Les Weer At ce Sel Pe 3a PTV Vine 2 Pak ba) be ee So et bd bi a Size 1-3. an. Fic. 28. Diagram illustrating the tuberculation of a single specimen of Pentabdella dispar, anterior extremity above, posterior below. Compiled from the details in Cordero (1937). Size numbers 1—3 indicate smallest, middle and largest annuli respectively, of any somite, but they are related only to adjacent somites since size of annuli generally decreases towards the extremities. ‘‘ }’’ in size column indicates the presence of a small annulus dorsally only. The dotted line indicates where annuli may or may not be present, while blank squares indicate where no information is given. See Fig. 13 (p. 409) for other abbreviations. Pon REVIEW OF THE PONTOBDELLINAE 435 similar to the central three annuli of previous somites. Somite XXV, the first of the three anal somites, consists of two annuli, equivalent to a, and a, of previous somites, while the two annuli of somite X XVI are much reduced. Somite XXVII is uniannulate and very small, and the anus opens mid-dorsally on the a, annulus of somite XXVI. TUBERCULATION. Text-fig. 29 is derived from the description by Cordero (1937). No details of tuberculation are given for the neck region. Each a, annulus of the preclitellar region has four pairs of mammilate tubercles with apical rosettes, one pair para-medianly on the dorsal surface, one pair laterally and two slightly smaller pairs ventrally. Annuli a, and a; possess five pairs of conical warts with apical rosettes, which are not as conspicuous as those on the a, annulus. In the clitellar constriction, annuli Xa;, Xla,, XIla, and XIIla, are generally smooth, while the other annuli are more wrinkled and bear six pairs of small sharply-pointed tubercles each. The b, and b, annuli of somites XIV to XXIII are smooth, while b, and b,; possess ten mammilate tubercles each, four dorsal, two lateral and four ventral, the centro-ventrals being smaller than their neighbours. The a, annulus possesses eight large conical tubercles with rosettes at their summits, two dorsally, two later- ally and four ventrally. Annuli XIIIa,, b; and b, have similar tuberculation to somite XIV. The tuberculations of annuli a, to a, of somite XXIV are similar to b,, a, and b; of previous somites and the first and second annuli of somite X XV are similar to a, and a; of somite XXIV. The a, and a, annuli of somite XXVI have eight and six tubercles respectively, while the single annulus of somite X XVII has about six tubercles. Small secondary tubercles may be dispersed between the larger tubercles and may even occur in the furrows. DISTRIBUTION. 34° 50’ S., 52° 20’ W., 58-65 fathoms; Ilha Rasa, 80 metres depth. Host. Not recorded. GENUS ET SPECIES INQUIRENDAE* Pontobdella rayneri Baird, 1869 Pontobdella vayneri Baird, 1869 : 313. DEscrRIPTION after Baird. Body cylindrical, attenuated anteriorly. Annuli with tubercles which have sensillae at summit. Preclitellar region twelve tuberculate annuli. Clitellum five tuberculate annuli. Anterior sucker small, circular, with three pairs of papillae on margin and large brown triangular pigment patches. Posterior sucker largest, rayed with brown and with puckered margin. On body, every third row of tubercles is largest. Length r inch. On re-examination of this material it was also found that the posterior sucker was joined to the body by quite a narrow waist ; however the material was in too poor a condition to improve on this description. This species is almost certainly a member of the genus Stzbarobdella. * Including Ovientobdella japonica (Vasiliev), see p. 439. 436 L. C. LLEWELLYN Pontobdella leucothela Schmarda, 1861 Pontobdella leucothela Schmarda, 1861 : 6. DESCRIPTION after Schmarda. Body circular with tubercles on back and sixteen trimeric somites. Neck fifteen annuli. A, annuli broadest, with four semi-conical tubercles. Colour yellowish grey, tubercles white. Length 18 mm. This species is apparently a member of the genus Stibarobdella. DISTRIBUTION. Port Jackson, New South Wales. Host. Not recorded. Pontobdella prionodiscus Schmarda, 1861 Pontobdella prionodiscus Schmarda, 1861 : 7. DEscrRIPTION after Schmarda. Body cylindrical, length 48 mm. and breadth Io mm., attenuated at each end, tuberculate and a dirty green with a little yellow and brown coloration. Anterior sucker with sixteen rounded tubercles and with eight papillae on the rim. Anterior : posterior sucker ratio 1: 1-6. Posterior sucker bell-shaped. Said to have some similarities to the quadriannulate form Pontobdella spinulosa, which is now a synonym of Pontobdella muricata. Pontobdella tatejamensis Oka, 1910 Pontobdella tatejamensis Oka, 1910: 171. Parapontobdella tatejamensis : Harant, 1929 : 650. DESCRIPTION after Oka. Body elongated, fusiform, posterior middle half of body expanded like abdomen. Red brown to dark green. Tubercles weakly developed. Suckers small. Lateral vesicles indistinctly recognizable externally. Length of body up to 2 cm., breadth up to 3 mm. This description gives no sound evidence for placing the leech in the genus Pontobdella. DistriBpuTION. Hondo; Awa, Suruga, Sagami. Host. Teleost. Pontobdella oligothela Schmarda, 1861 Pontobdella oligothela Schmarda, 1861 : 6. This species does not belong to the Pontobdellinae since it possesses numerous salient pulsatile vesicles which the latter lack. The rough white patches described on the dorsal surface of somite 10, Ir, 14 and 17 on fresh specimens seem to indicate, according to Blanchard (1894) and Harding (rg1to), that the species is probably identical with Trachelobdella lubrica Grube, 1840 : 60, (Pontobdella littoralis : John- ston, 1865 and Pontobdella campanulata : Johnston, 1865, pp. 42 and 304 were also probably Trachelobdella lubrica). REVIEW OF THE PONTOBDELLINAE 437 Pontobdella vittata de Blainville, 1828 Pontobdella vittata de Blainville, 1828 : 557. Not one of the Pontobdellinae because it possesses no tubercles and probably has biannulate somites. OPHIBDELLA Beneden & Hesse, 1863 Ophibdella labvacis Beneden & Hesse, 1863 : 25 The specimen described under this generic name had no tubercles, yet Apathy (1888) believed it was a young Pontobdellinid. However, the hatching young of Pontobdella muricata are similar to the adults except in size, so presumably Ophibdella should remain a separate genus. WwW | | on) NI | ae) It KEY TO THE SPECIES IN THE SUBFAM. PONTOBDELLINAE Abdominal somites triannulate Gen. STIBAROBDELLA 2 Abdominal somites quadriannulate Gen. PONTOBDELLA 9 Abdominal somites quinqueannulate . . Pentabdella dispar (p. 433) Anterior sucker with a marginal fringe Stibarobdella loricata (p. 413) Anterior sucker without a marginal fringe . : : ; : : 3 Anterior sucker with a pigment patch or patches 4 Anterior sucker without pigment patches ; ; : 3 é : 7 Anterior sucker with no papillae Stibarobdella taprobanensis (p. 421) Anterior sucker with papillae 4 : ‘ : ‘ : 5 Anterior/posterior sucker ratio between 1 : 2 and I : 3 Stibarobdella macrothela (p. 405) Anterior /posterior sucker ratio between 1 : I find 1: 2 : Anterior sucker with two distinct pigment patches and anterior fposteriot sucker ratio approximately 1 : 1 or latter slightly larger : Stibarobdella bimaculata (p. 415) Anterior sucker with single triangular pigment patch and anterior/posterior sucker ratio approximately I: 1.5 Stibarobdella tasmanica (p. 420) Anterior sucker with papillae Anterior sucker without papillae : Stibarobdella moorei (pe. 422) 9 tubercles including a mid-ventral tibetcle: on the a, annulus Stibarobdella planodiscus (p. 418) 6 tubercles on a, annulus 4 dorsally and 2 ventrally Stibarobdella australiensis (p. 422) Anterior sucker with fringe or welt . : : : : : 6 : : Io Anterior sucker without fringe . WT Anterior sucker without pigment patches but swith 3 pairs of papillae, aafiseony| posterior sucker ratio 1 : 1 or former slightly larger . Pontobdella muricata (p. 423) Anterior sucker with a pair of pigment patches and 52 marginal papillae and 2 sub- marginal rows of papillae, anterior/posterior sucker ratio approximately 1 : 1-7 Pontobdella rugosa (p. 432) ae sucker with 3 pairs of papillae, anterior/posterior sucker ratio between wUsSeand ten 2. . Pontobdella vosmaeri (p. 427) perso: sucker without ‘papilla, anterior posterior sucker ratio I: I or former slightly larger. : : : : : : Pontobdella aculeata (p. 431) 438 L. C. LLEWELLYN ACKNOWLEDGEMENTS In presenting this review I must thank the D.S.I.R. for the award of a Research Studentship, the Trustees of the British Museum (Natural History) for allowing me to study the collection, Mr. R. W. Sims, for much help with the nomenclature and presentation, Professor E. W. Knight-Jones and Dr. K. H. Mann for advice and criticism, and the Director of the Plymouth Laboratory, for access to the collections there and to the library. REFERENCES ApatHy, S. 1888. Systematische Streiflichter. I, Marine Hirudineen. Arch. Naturgesch. (1888) : 43-61. Bairp, W. 1869. Descriptions of some new suctorial annelides in the collection of the British Museum. Pyvoc. Zool. Soc. Lond., (1869) : 310-318. BENEDEN, P. J. van, & Hesse, C. E. 1863. Reécherches sur les Bdellodes ou Hirudinées et les Trématodes marins. Mém. Acad. R. Belg. 34: 11-450. BLANCHARD, R. 1893. Révision de Hirudinées du Musée de Turin. Boll. Musei Zool. Anat. comp. R. Univ. Torino, 8 (145) : I-32. 1894. Hirudinées de l’Italie continentale et insulaire. Boll. Musei Zool. Anat. comp. Rk. Univ. Torino. 9 (192) : 1-84. Bourne, A. G. 1884. Contributions to the anatomy of the Hirudinea. Quart. J. micr. Sci. 24 : 419-500. CABALLERO, E. 1956. Hirudineosde Mexico. XX, Taxa y nomenclatura de la clase Hirudinea hasta generos. An. Inst. Biol. Univ. Méx. 27 : 279-302. CorpDERO, E. H. 1937. Hirudineos neotropicales y subantarticos nuevos, criticos 0 ya cono- cidos del Museo Argentino de Ciencias Naturales. An. Mus. Argent. Cience. Nat. 39 (4): I-77. GopDARD, E. J. 1909. Contributions to our knowledge of Australian Hirudinea. Pyvoc. Linn. Soc. N.S.W. 34 (4) : 721-732. Harant, H. 1929. Essai sur les Hirudinées. = Ss co SS 7 cs al —s = a ond ee a ne oa ee ‘ssoydiourejour ye syuourdos 7$1-EET 1940 soyeasror snwe oy} yeyy suises snyy yf “Ss WUNWTUTUT oY) ‘FZ SVM AOR S}[UPe OT} M1 PUNOF OVA OFAOA yeurosd yo soquind TantENXeUd otf, ‘sourojoAw jeuvoid ZZ Sey VAIPL OL, OVLY TINS SUISSTUL quoso.1d SUOT}POOT[O9O 10 yuosqe pjoy-uy ALOAOODSICY +087z uy [vuy LOZ A[UO ge L ob! 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