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BULLETIN OF 
THE BRITISH MUSEUM 
(NATURAL HISTORY) 


ZOOLOGY 
VOL<14 


1966 


TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 


LONDON: 10968 


DATES OF PUBLICATION OF THE PARTS 


No. I 1 February 1966 
No, 2 1 February 1966 
No. 3 11 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 


No. 


CONTENTS 
ZOOLOGY VOLUME I4 


The Oedura tryont complex: east Australian rock-dwelling geckos. 
(Reptilia: Gekkonidae). By H. R. BusTarpD 


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. HAsTINGs 


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 Piuiletor (Chiroptera: Vespertilionidae). By 
te. HIT 


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 


107 


371 


389 


441 


487 


THE OEDURA TRYONI COMPLEX: 
EAST AUSTRALIAN ROCK-DWELLING 
GECKOS. (REPTILIA: GEKKONIDAE) 


H. ROBERT BUSTARD 


BULLETIN OF | 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 140 No; 71 
LONDON : 1966 


fae OFDURA FRYONI 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. 14 No. 1 
LONDON: 1966 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), tnstituted 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 
ready. 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. 1 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 i February, 1966 Price Ten Shillings 


~ 


ee 


THE OEDURA TRYONI COMPLEX: EAST 
AUSTRALIAN ROCK-DWELLING GECKOS. 
(REPTILIA : GEKKONIDAE) 


By H. ROBERT BUSTARD 


SYNOPSIS 


Following an historical review of the genus Oedura Gray the specific distinctness of O. tryont 
De Vis from O. marmorata Gray is confirmed. O. ivyoni 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. lesueurit (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. tvyont De Vis. The geckos previously referred to as O. trvyont form 
a discrete group of predominantly rock-dwelling Oedura which can be divided into 
two forms on the basis of morphology, size, coloration and geographic distribution. 
O. tryom as recognized by Cogger (1957) is herein split into two species and the 
variation within them is discussed. 

Collections of Oedura 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 Oeduratryont. Before describing the Oedura tryomt 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. tvyont De Vis, 1884; O. monilis De Vis, 1887; O. 
vobusta Boulenger, 1885 ; O. lesueuri lesueurts (Dum. & Bibr., 1836) and O. lesueuri 
rhombifer Gray, 1845. 


MATERIALS AND METHODS 


At sexual maturity in Oedura 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 H. ROBERT BUSTARD 


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 Oeduva 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 
O. monilis and only one with O. tryoni. A lectotype will be selected from the five 
specimens conspecific with monzlis and the full implications of this will be explained 
in a later paper. 

Phyllodactylus ( = Oedura) castelnaui Thominot (1889) was synonymized with 
O. tryoni by Zietz (1920). The author has examined the three syntypes of Oedura 
castelnaui and has found Zietz’s action to be incorrect. Oedura castelnaui belongs to 
the O. marmorata complex and is being reinstated as a full species (Bustard, in prepn.). 

Loveridge (1934) synonymized O. tryonit 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 tryont. 

Cogger (1957) drew attention to a differentiated population of O. tryont in North 
Queensland and showed that it differed in interorbital and mid-body scale row 


EEE ee —_ 


—_ 


LIZARDS OFTHE OLRDURA PRYONI COMPEEX 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. 


ELE SA LU SiO OF) DURA T Rey ONE DE} NaLS 


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. [X, 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 
tryomt rather than to instigate a redescription of marmorata. 

Cogger (1957) has shown that tryoni is a valid species, on the basis of ecological 
information and meristic characters. A glance at living material is sufficient to 
separate marmorata and tryont 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. tryom 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 marmovata 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 tvyont were examined 
and in every case there was a single postanal tubercle, 


6 H. ROBERT BUSTARD 


O. tryont 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. tryont 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. tryont is selected as neotype. 


Oedura tryoni De Vis 
(Plates 1, 2) 


NeEotTyPE. AM. R 21601, 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. 
First 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 


Li Zee SeOrR IME ODM ATRYONT COMPLEX, ~ a 


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. 
R 15092), 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 2ogg91 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 (OM. J 2294), Woodford, N.C. Line (QM. J 3870), 
Warwick (QM. J 1354), 5 miles N. of Warwick (BMNH. 1964.1509-1510, Gympie 
(OM. 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. (I) Subdigital lamellae. On the fourth toe of the hind foot of 
tvyont there are 4-6 divided subdigital lamellae (75°% possess 4), followed by 1-4 
(usually 3) undivided ones. That is, omitting the distal plates, tvyont 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 


Interorbitals 


80 90 100 110 120 130 


e O6cdura coggeri 


e | 
Mid-body scale rows © Od6cdura tryoni 


Fic. 1. Counts of mid-body and interorbital scales in O. tryoni 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 tvyoni does not resemble coggert 
more closely than that of the southern populations. Although populations of tryoni 
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 tryont (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 g—Io 
(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 tyonz is 
52-95 (77) mm. The smallest and largest males have a snout-vent length of 52 and 


EIZAR DS OFTHE OLDURA TRYONT 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 consis of O. tryont 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. tryoni. 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. 1, 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 tryont 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) 


HototypeE. AM. Riu7791—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), 81 miles N.W. of Cairns on road to Laura via Mt. Carbine (BMNH. 
1964.1495), Stannary Hills (OM. J 1293-5), Ravenshoe (BMNH. 1923.10.13.1-2). 


DESCRIPTION OF HoLotypPEe. 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 (11); 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 coggeri 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, coggeri possesses 5-6 
enlarged subdigital lamellae. The overlap in number with évyont, 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. tryoni 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- 


EIZARDS OFVIHE OLDURA 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 10-12 (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 coggert can be separated from tvyont 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. 


DISTRIBUTION AND HABITAT 


The distribution of Oedura coggert and O. tryont is shown in text-fig. 2. Neither 
species penetrates far inland and both are probably restricted to Eastern Australia. 


Queensland 


New South Wales 


® Oedura coggeri 


o Oedura tryoni 


—l 
100 miles 


Fic. 2, Distribution of O. tvyoni and O. coggeri. 


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 tryont has been examined recently (Cogger, 1957) and 
Oedura coggerit 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 
coggert 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. coggert 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, monilis 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 Oedura charac- 
terized by spots. O. tryonit and some coggeri possess numerous, irregularly placed, 
small, light-coloured spots. The distribution and size of these spots is diagnostic as 
pointed out by Cogger. In tryont and coggeri the spots are numerous and much 
smaller than the orbit whereas in monilis 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 tryoni and coggert, 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 monilis 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 tryont and 
coggeri. 

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 tryont. O. tryoni 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 tryoni has been considered strictly rock 


Ei ZAn DS OERNDHE OL DURA DRYONT 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 tvyont and coggert populations mainly 
occur in rock crevices. At Mount Garnet the northern population of O. marmorata 
and O. lesueuri rhombifer were common below the bark of dead trees (Eucalyptus spp.) 
in the area where the specimen of O. coggert was collected. The three forms are 
also known to be sympatric in a area between Atherton and Mt. Garnet. 

The distribution of tryont 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 castelnaut and mayer: 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 coggert 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 tryont 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. coggert differentiated 
from O. tryoni following geographic isolation. The syntypes of O. tryoni have been 
lost and a neotype is proposed in the interests of stability. 


ACKNOWLEDGEMENTS 


I am grateful to Mr. Joe Bred! 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 


BoucenGcer,G. A. 1885 & 1887. Catalogue of Lizards in the British Museum (Natural History). 
Vols. 1 and 3. 2nded. London. 

Coccer, H. G. 1957. Investigations in the gekkonid genus Oedura 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. Proc. Roy. Soc. Qld. 1: 53-56. 

————- 1887. A contribution to the herpetology of Queensland. Proc. 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. 1st ed. London. 
LonGMAN, H. A. 1915. Reptiles from Queensland and the Northern Territory. Mem. Qld, 
Mus. 3: 30-34. 2 plates. | 
LoveripGe, 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. 

TuHominot, A. 1889. Observations sur quelques reptiles et batraciens de la collection du 
Muséum d'Histoire naturelle de Paris. Bull. Soc. Phil. Paris. 8 (1) : 21-30. 

Zietz, 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 coggeri n. sp. 
LoweER Row : O. tryoni habitat at Mr. Marlay, Stanthorpe. The neotype was collected from 
under the slab at bottom centre. 


Bull. B.M. (N.H.) Zool. 14, I PLATE 1 


PLATE 2 


Top row (left to right): O. tryoni juvenile from type locality (Stanthorpe), adult Ac 
dwelling variant from Tara, S. Queensland and juvenile from 13 m. north of Bendemeer. 
MIDDLE AND BOTTOM ROWS: Variation among adult topotypic O. tryoni. 


Bull. B.M. (N.H.) Zool. 14, 1 Ril wie. 2 


PLATE 3 
Variation among O. coggeri. Tor Row: Petford. MIDDLE Row: Hartley’s Creek, near Cairns. 
BOTTOM ROW: Lappa Junction. 


PLATE 3 


Bull. B.M. (N.H.) Zool. 14, I 


RICHARDSON’S ELOPOID AND CLUPEOID FISHES Bi 


only 15 branched anal rays (18-20 in H. ilisha) and the anal origin is equidistant 
between the caudal base and pelvic tzps (pelvic base in H. tlsha). 

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 tolt (Valenciennes) also occurs in this region (see Table 2), but it has a narrow 
operculum (as in H. ilisha) and the caudal lobes exceed head length (about equal to 
head length in H. ilisha and H. reevesit). 


mee SCRIPTION OF THE TYPE OF CEUPEA REEVESTIEI 


Standard length : 295 mm. 
Total length : 369 mm., or 144 inches (approx. I5 inches according to Richardson). 


mm. Ofek 
Body depth : é ; : : : : gI-o 30°7 
Head length 2 , ; ‘ : : ; 84°5 28°79 
Snout length P > : : : : : 21°5 Ve 
Eye diam. : . : : : : : 12°8 4°3 
Upper jaw l. : ; , : : 2 35°0 EEeO 
Lower jaw l. ‘ : : : : : : 43°2 I4°7 
Pectoral 1. ; ‘ : . . : : 47°0 15°9 
Pelvic 1. .. . j ' ‘ . : F 24°5 8°3 
Operculum height : : : : : : 35°5 12-0 
width ‘ 3 ; : : : 24°0 Sz 
Pre-dorsal , : ; ; ‘ ; 149°4 50°5 
Pre-pelvic . ' ‘ 2 , ‘ . ’ 143°8 48°5 
Pre-anal_ . : q : ‘ ' : ‘ 239°0 81:0 
Dorsal iii 14 (or iv 13, tips broken) ; pectorali15; anal ili 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 reevesur 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 would be expected. 


Note. Richardson identified this fish rather tentatively with Russell’s Palasah 
and at the same time distinguished it from his own Alosa reevesit because of differences 
ZOOL: 14, 2 


4 


32 P;jJ.P. WHITEHEAD 


in head length, pectoral length and body outline. The specimen is clearly H. reevesii 
and not the Palasah of Russell (1803) (i.e. Hilsa ilisha (Ham. Buch.)—see Whitehead 
1965a), having a broad operculum, its width contained only 1} times in its length. 
The differences from H. reevesii 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. reevesi ; cf. 45-48 in H. tlisha— 
see Whitehead 1965a). 


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 abnormis Richardson, 1846, Ichth. China Japan : 306. 

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 
“ Ilisha 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. 


IiGuRE. 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 23-3} in 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. abnormis). 

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 /lisha 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). 


PEC hAR DSON Ss LOPOLD AND CLUPEOLD FISHES 33 
meee oChEPiLON- OF THE TYPEOF ILI SHA ABNORMIS 


Standard length: 295 mm. 
Total length : 356 mm. or 14 inches (approx., caudal tips damaged). 


mm. yon Os 
Body depth ’ ; : : ; < 76-0 25°8 
Head length : : : : , ; 67°38 23:50 
Snout length ‘ : : ; : , : 19:6 6°5 
Eye diam. ; ‘ ' ‘ : : 16-6 5°4 
Upper jaw l. ; ; : é é : 34°0 II°5 
Lower jaw l. : “ ; : ; ; 30°1 E262 
Pectoral length . ; ; i i : : 37°5 IZ. % 
Pelvicilengta. .. : : ‘ ‘ ‘ ; TE4 32°90 
Length of anal base . 2 ‘ : 118-0 40°0 
Pre-dorsal : ‘ : : : : : 150°5 51-0 
Pre-pelvic * : : : : ; FEOrS 30°5 
Pre-anal . : : , ; ; ‘ : E775 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 ili 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). 


) 


9g. ‘‘ 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. 


FIGURE. 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 Konosirus 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. %S.L. 
Body depth 57°0 30°0 
Head length A ‘ - : : ‘ 47°4 24°7 
Snout length ‘ ; : : ; ; : 9°7 51 
Eye diam. R : ; : ‘ ; : 9°3 4°9 
Upper jaw I. : ° ; : ; ; R 15°0 79 
Lower jaw l. - : d . f : ; 190 10°0 
Pectoral 1. : ° : : : ; . 33°0 17°*4 
Pelvic]. . : ‘ . ” . F ‘ 17°0 8-9 
Pre-dorsal : ° , : ‘ ; f 92°9 48°7 
Pre-pelvic . ° ° ; , ; ‘ ; 97°2 51°1 
Pre-anal . ° ° ; ; ‘ 5 : 146°2 78-0 


Dorsal with iii 14 (or iv 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 1} 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 aguosus 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. 


10. ‘“‘ Chatoessus triza Linnaeus ”’ 
= Clupanodon thrissa (Linnaeus) 


“Mystus altus Linnaeus, 1754, Chinensia Lagerstrémiana—“ dissertatio’’ : 26—China, on 
Lagerstrém. | 

Clupea thrissa Osbeck, 1757, Dagbok Ostind, Resa : 257—Canton area. ] 

Clupea thrissa Linnaeus, 1758, Syst. Nat., ed. 10: 318 (name from Osbeck; diagnosis after 
Lagerstrém). 

Clupanodon thrissa: Fowler, 1930, Proc. Acad. nat. Sci. Philad. : 599 (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. Mus. (nat. Hist.) Zool. 9 


(2) : 100 (generic review). 


ReCHARDSION “Ss BLOPOID AN DCLUPEOID- FISHES 35 


Clupea triza 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 osbeckit Valenciennes, 1848, Hist. Nat. Potss. 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 (PI. 4, fig. 1) 
and is a fair representation of the blunter snout found in C. ¢tivissa compared with 
that in K. punctatus. 

b. Operculum. In C. thrissa the operculum is slightly 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 PI. 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 K. 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 2§—3 in standard length ; cf. 3-341n K. punctatus according to Regan, 19174). 


There can be little doubt, therefore, that the illustration refers to C. thvissa 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 tviza 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 (Int. 
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 Lagerstrom 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 triza from the Lagerstrém 
collection at Uppsala; he too, does not mention the filamentous last dorsal, which 
was presumably broken off. 


36 P. J. P. WHITEHEAD | 
1r. “ Chatoessus chrysopterus Richardson ”’ 
== Nematalosa sp., ? N. nasus (Bloch) 


Clupea nasus Bloch, 1795, Nat. Ausl. Fische, 9 : 429, fig. 1. 

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). 

Konosirus 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 2} in standard length) 
with an elongated last dorsal ray and an inferior mouth with a short maxilla (PI. 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 2?—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. Igo). 

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 th 
dermal head bones are not accurately drawn in any of the drawings. A mistake o 


RICHARDSON’ SELOPOID AND CLUPEOID FISHES 37 


_ this kind seems much more likely than one in body depth, and the drawing agrees 


with N. masus 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 Gitinther (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). 

Gitinther (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. 1) 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 (Pl. 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. iviza (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 commersoni Lacepéde, 1803, Hist. Nat. Poiss. 5 : 381, 382, pl. 12, fig. 1. 

Engraulis commersonianus : Richardson, 1846, Ichth. China Japan: 308 (Reeves specimens ex 
“China ’’) ; Fowler, 1930, Proc. 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 Engraulis chinensts Giinther, 1880. 


Figure. 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, 19634), considerable confusion existed in the 
nomenclature of the Chinese and Japanese anchovies. However, Richardson 
correctly identified the British Museum specimens with Lacepéde’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 Engraulis). 
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, 
1964) in his synonymy of E. 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 (Giinther, 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 Giinther’s catalogue 
(1868, p. 390) as: 

a, b,c-f Adult and half grown China 
g-i 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 japonicus 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 Ginther (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, E. japonicus certainly occurs in the 
Philippines and is also found in Hong Kong waters (B.M. specimens). Closely 
related to S. commersonii 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 maxilla (to front edge of operculum, not to 
gill-opening). Fowler (1931, p. 201) lists only one other Chinese record for S. indicus, 
namely Engraulis encrasicholus of Giinther (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. 99-101). 


mele H Ak SON SckhLOP Ol DIAN D CLUPEOLD FISHES 39 


J 


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). 

Cotlia gvayi 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 (jide 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. grayu 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 74 inches. 


SPECIMENS. Giinther (loc. cit.) lists five specimens under C. grayu. 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 e 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. 


FIGURE. 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 104 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 Pp. J. P. WHITEHEAD 


(1754) had originally named this fish Mystus enstformis, giving a figure (fig. 12) which 
was later reproduced in the Amoenitates Academiae (1759). As in 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. grayii 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. 
Lénnberg 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. 1855.9.19.1581; Reeves fish: BMNH. 1964.11.6.2. 
Standard length: 243 mm. (TYPE) ; 238 mm. (Reeves) 
Total length : 275 mm. (TYPE) ; 262 mm. (Reeves) 


mm. %S.L. 
Body depth ; ; ‘ - 45°2 41°3 18°5 17°3 
Head length : ; : . ‘eere 40°2 18:1 16°9 
Snout length : : ; ; 9°6 g°I 3°7 3°8 
Eye diam. . ; ; , 8-9 8-6 3°6 3°6 
Upper jaw 1. , ; R i ee 51°1 21-9 21°5 
Lower jaw l. : > ; me SONG 29°6 I2°7 I2°*4 
Pectoral 1. : . : » 79°5 88-2 32°7 37°0 
Pelvic l. . . : - : 19°*4 15°9 8-0 6°7 
Pre-dorsal ° ; 73°7 65°4 30°2 27°4 
Pre-pelvic . ‘ , ‘ : 69°2 66-0 28°5 27°9 
Pre-anal . : ; : » 1050 100°2 43°0 42°0 


Body compressed, depth about equal to head length, posterior portion elongated, 
caudal peduncle about }? eye at caudal base. Maxilla 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 


micCriewenh DSO s ELOPOTD AND CLUPEHOID FISHES 41 


coronoid process. Pseudobranch exposed, almost equal in length to eye diameter, 
filaments about twenty, short. 

Dorsal (preceded by small spine) with iii 10 rays. Pectoral with vii ro rays, the 
first seven filamentous and unbranched, reaching to about base of 10o-14th branched 
anal ray, well beyond tip of depressed dorsal; branched rays of pectoral reaching 
beyond pelvic base. Pelvic i 6, its base below anterior dorsal rays and nearer to 
pectoral base than to anal origin. Anal iii 88 and 111 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 II. 


15. “ Coilia playfairii McClelland ”’ 
= Coilia playfairii (McClelland) 

Choetomus playfaivit McClelland, 1844, Calcutta J. nat. Hist. 4: 405, pl. 24, fig. 3 (China on 
Playfair specimen(s)). 

Cotlia playfaivi: Richardson, 1845, Voy. Sulphur Ichth.: 100, pl. 54, figs. 3-4 (Hong Kong, 
China seas) ; Idem, 1846, Ichth. China Japan : 309 (Japanese specimen). 4 

Coilia grayt Kner, 1865, Reise Novara, Fische : 335 (Hong Kong). 

Cotlia clupeoides : Giinther (parvt.), 1868, Cat. Fish. Brit. Mus. 7: 404 (Richardson specimen ev 
China). 

Cotlia nasus: Giinther, 1868, Cat. Fish. Brit. Mus. 7: 405 (Japanese, Chinese specimens) ; Nichols, 
1943, Nat. Hist. Central Asia, 9: 19 (Anhwei, nr. Canton). 

? Cotlia 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 “ Cola 
playfairu Type’’. In fact this specimen may well be that on which the figure of 
C. playfairu in the “ Voyage of the Sulphur ’”’ was based (pl. 54, fig. 3, stated to be 
natural size). However itis not atype. 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 Cozlza 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 (9-11), or a short maxilla not 


42 P. Jj. P. WHITEHEAD 


reaching the gill opening, there are three species known from Chinese or Japanese 
waters (C. playfairit, 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. playfairii. 

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. playfairit. 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. Ichth. 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 hamiltoni 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. 


' Referred to as‘ hairy process behind gills '’ in the Chinese Chrestomathy under species No. 4 (Bridg- 
man 1541, p. 450 


mich wk DsSON Ss ELOPOrID AND CLUPEOTID FISHES 43 


Note. Fowler (1931) recorded both T. hamiltonu and T. mystax from Hong Kong. 
T. hamiltoni is distinguished from 7. mystax by its shorter maxilla (to gill opening 
or just beyond; cf. to or beyond pectoral base in I’. mystax). It is suspicious, 
however, that the specimens of T. hamultoni in our collections are mostly large fishes, 
whereas those of 7. mystax are small. Amongst the smaller specimens labelled 
T. hamilton (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 7hrissocles Jordan & Evermann, widely but wrongly used for this genus 
as a senior synonym, should be replaced by Thryssa Cuvier, 1829 (see Whitehead, 
19655). Fowler (1931, p. 203) placed both 7. mystax and T. hamiltona in Setipinna 
Swainson, although he had (correctly) characterized that genus as possessing a 
filamentous upper pectoral ray. 


DESCRIPTIONOFP 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. Tiel. 

ao fa ae aN 
Body depth ; : , : . 46* 4 ~ I> 2 2573, 2s 
Head length : : : : , AO-E AEF 22 O22 x8 
Snout length 4 : : : : yee Fe BZ aS 
Eye diam. : : 3 3 J S6n 1, Sto A746 
Upper jaw length . : : AEs, swe 22 228 
Lower jaw length : ; . y 320°5.. 29°5 how, 115° 7, 
Pectoral |. : : : : : sae. i776 = 
Pmewicd’ |. : } . 3 : He Eco) ea 7-5 
Anal base I. ; i ; 4 : 54°I 63:8 20°06 33:8 
Pre-dorsal : is : : ; 95°4 102-9 52°2 54°5 
Pre-pectoral : : : : , AEOG 2 OM 
Pre-pelvic . : : : : : ia AIR 
Pre-anal_ . : . : : : I15°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 ii 11 rays, preceded by a minute spine, its origin equidistant between 
caudal base and anterior half of eye. Pectoralizo; pelvici6. Anal with iii 37 
rays, its origin slightly behind last dorsal ray. 

Abdominal scutes keeled, with sharp spines, 18 pre-pelvic (first minute), Ir 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. 


17. ‘ Megalops setipinnis Forster ’’ 
== Megalops cyprinoides (Broussonet) 


Clupea cyprinoides Broussonet, 1782, Tableau Ichth.: no pagination, pl. 9 (Oceans between the 
tropics). 
Pict wi 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 Giinther (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. 1843.8.10.1r 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. 961s 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 
znd 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 M. setipinnis 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. 


BiIcCHAR DSON Ss ELOPOLTD 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 (PI. 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 M. 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). 


19. “ Elops machnata Forskal”’ 
= Elops machnata (Forsskal) 
Argentina machnata Forsskal, 1775, Descriptiones Animal. : 13, 68 (Red Sea). 
Elops machnata : Richardson, 1846, Ichth. 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. 1064. 
nr. 0. 3. 

FIGURE. Reeves No. 137 is a fair drawing and easily recognizable as Elops 
by its small scales, elongate body, pored lateral line scales, etc. (Pl. 7, fig. 1). 


Notes. The specimen has approximately 87 pored lateral line scales, which 
accords with EF. machnata, and the lower jaw (now set open) probably covered the 
pre-maxillary tooth band when the jaw was closed (lower jaw included in F.. hawaien- 
sis Regan). On the basis of the most recent key (Whitehead, 19620) the specimen is 
evidently FE. 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. 


46 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. 


Nore. 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 E. machnata. Only a single 
species of Elops is recognized from the Western Pacific (see Whitehead, 19620). 


21. ‘“‘ Chirocentrus dorab Forskal”’ 
= Chirocentrus dorab (Forsskal) 


Ciupea dorab Forsskal, 1775, Descriptiones Animal. : 72 (Red Sea). 

Chirocentrus dorab: Richardson, 1846, Ichth .China Japan: (Canton, seas of China); Giinther, 
1868, Cat. Fish. Brit. Mus. 7: 475 (Reeves specimen) ; Fowler, 1930, Proc. Acad. nat. Sci. 
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. 


FicurE. As Richardson noted, the Reeves figure (Reeves No. 47) hints at pun- 
gent ventral scutes (Pl. 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 (10} 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. dorab (54 and beyond 
respectively in C. hypselosoma). The systematic position of Bleeker’s C. hypselosoma 
has been examined elsewhere (Whitehead, Boeseman & Wheeler, in press) and the 
conclusion reached that there may indeed be two species of Chirocentrus 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*ELOPOLDLAND 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. zollingert (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, 19640), a species unlikely to be 
misidentified by Japanese workers. But published descriptions of S. zollingert 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, 1965) but not from inter- 
vening areas. 

The list of species of Thryssa, Cotlia 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, 14, 2 5A 


48 P. J. P. WHITEHEAD 


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 dorab (Forssk.) 


DUSSUMIERIIDAE 
Etrumeus teres (DeKay) 


Dussumiteria acuta Valenc. 
Spratelloides gracilis (Schlegel) 
CLUPEIDAE 
(Clupeinae) 
Sardinella aurita Valenc. 
+Sardinella brachysoma Blkr. 


+Sardinella bulan (Bleeker) 
Sardinella jussieu (Lac.) 


Sardinella fimbriata (Valenc.) 


Sardinella leiogaster Valenc. 


tSardinella clupeoides (Blkr.) 


+Herklotsichthys schrammi (Bleeker) 
+Herklotsichthys punctatus (Riippell) 
+ Herklotsichthys zunasi (Bleeker) 


(Alosinae) 
Hilsa (Tenualosa) reevesti (Rich.) 


Hilsa (Tenualosa) toli (Valenc.) 


(Pristigasterinae) 
+Pellona ditchela Valenc. 


* Specimen in British Museum from this locality. [ ] references from other areas. 
+ Species may occur in Hong Kong waters, but no record or specimen. 


TABLE 2 


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, 1963, 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); [Kishinouye, 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-vittata; ? Jouan, 1867—Hong 
Kong, as Harengula moluccensis. 

[Kishinouye, 1907—Japan, as C. okinawensis ; 
Fowler, 1941—Philippines; Chan, 1965— 
Thailand, Philippines]. 

[Chu & Tsai, 1958—Formosa]. 

(Chu & Tsai, 1958—Formosa] ; Philippines*. 

(Fowler, 1951—Japan, Philippines]; Japan*, 
East Indies*. 


Richardson, 1846; Whitehead, 1965a (Hon 
Kong*). 

Fowler, 1931 (Hong Kong, as Macrura sinensis) 
{Jordan & Evermann, 1902—Formosa]. 


(Liu & Shen, 1957—-Taiwan, as Ilisha hoevent) 


RECHARDSON SE LOPOID AND CLUPEOID FISHES 49 


Ilisha elongata (Bennett) 


Ilisha brachysoma (Bleeker) 
Ilisha indica (Swainson) 


Ilisha novacula (Valenc.) 


Opisthopterus tardoore (Cuvier) 
Opisthopterus valenciennest Blkr. 


(Dorosomatinae) 
Konosirus punctatus (Schlegel) 


Clupanodon thrissa (Linn.) 


Nematalosa nasus (Bloch) 


Nematalosa japonica Regan 


+t Anodontostoma chacunda (Ham. Buch.) 


ENGRAULIDAE 
Engraulis japonicus (Schlegel) 


Stolephorus commersonii Lac. 


Stolephorus indicus (van Hass.) 


Stolephorus buccaneert Strasburg 
Stolephorus tri (Bleeker) 
Stolephorus heterolobus (Riipp.) 


Thryssa mystax (Schneider) 


tThryssa setivostris (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 Kone); [Eu &~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,, to61— 
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 P. J. P. WHITEHEAD 
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pIeCHARDSON SH DOPOTDEAND GLUPEOID FISHES 51 


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| 


52 P. J. P. WHITEHEAD 


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INDEX TO SPECIES 


abnormis, Ilisha, 32, Pl. 3 gracilis, 29 
acuta, Dussumieria, 48 isingleena, 24, Pl. I 
Alausa reevesti, 30 melanosticta, 27 
Albula vulpes, 48 mystax, 42 
Alosa elongata, 32 mystus, 39 
palasah, 31, Pl. 3 nasus, 36 
reevesti, 30, Pl. 2 nymphaea, 26, Pl. 1 
altus, Mystus, 34 okinawensis, 48 
Anodontostoma chacunda, 49 thrissa, 34 
aquosus, Chatoessus, 33, Pl. 4 triza, 35 
Argentina, machnata, 45 Clupea (Amblygaster) melanosticta, 28 
aurita, Sardinella, 26, 48 clupeoides, Coilia, 41 
Sardinella, 48 
brachysoma, Ilisha, 32, 49 Coitlia clupeoides, 41 
Sardinella, 48 ectenes, 41 
buccaneeri, Stolephorus, 47, 49 grayii, 39, Pl. 5 
bulan, Sardinella, 48 mystus, 39, 49 
nasus, 41 
caeruleo-vittata, Clupea, 28, Pl. 2 playfairii, 41, 49, Pl. 5 
chacunda, Anodontostoma, 49 commersonianus, Engraulis, 37 
Chatoessus aquosus, 33, Pl. 4 commersonii, Engraulis, 37 
chrysopterus, 36, Pl. 4 Stolephorus, 37, 49 
maculatus, 35, 37, Pl. 5 curtifilis, Megalops, 44, 45, Pl. 6 
osbeckii, 35 cyvprinoides, Clupea, 44 
punctatus, 33 Megalops, 44, 45, 48 
triza, 34, Pl. 4 
chinensis, Engraulis, 37 delicatulus, Spratelloides, 30 
Chirocentrus dorab, 46, 48, Pl. 7 ditchela, Pellona, 48 
hypselosoma, 46 dovab, Chirocentrus, 46, 48, Pl. 7 
Choetomus playfairii, 41 Clupea, 46 
chrysopterus, Chatoessus, 36, Pl. 4 dussumieri, Thryssa, 49 
Clupanodon thrissa, 34, 49 Dussumieria acuta, 48 
Clupea caeruleo-vittata, 28, Pl. 2 
cyprinoides, 44 ectenes, Coilia, 41 
dovab, 46 elongata, Alosa, 32 


flosmaris, 29, Pl. 2 Ilisha, 32, 49 


Elops machnata, 45, 48, Pl. 7 
purpurescens, 45, Pl. 7 
saurus, 45 

Engraulis chinensis, 37 
commersonianus, 37 
commersonit, 37 
hamiltoni, 42 
japonica, 37, 49 
mystax, 42 

ensiformis, Mystus, 39 

Etrumeus teres, 48 


fimbriata, Sardinella, 24, 48 
Spratella, 24 
flos-maris, Clupea, 29, Pl. 2 


gracilis, Clupea, 29 
Spratelloides, 29, 48 

grayana, Pellona, 32 

grayu, Cotlia, 39, Pl. 5 


hamiltonu, Engraulis, 42 
Thryssa, 42, 49 

Harengula moluccensis, 28 
nymphaea, 24 

Herklotsichthys punctatus, 48 
schrammti, 48 
zunast, 48 

heterolobus, Stolephorus, 49 

Ailsa veevesii, 30, 31 

Hilsa (Tenualosa) reevesii, 48 
tolt, 48 

hoevemt, Ilisha, 48 

hypselosoma, Chirocentrus, 46 


Ilisha abnormis, 32, Pl. 3 
brachysoma, 32, 49 
elongata, 32, 49 
hoeveni, 48 
indica, 49 
novacula, 32, 49 

indica, Ilisha, 49 

indicus, Stolephorus, 49 

isingleena, Clupea, 24, Pl. 1 


japomica, Engraulis, 37, 49 
Nematalosa, 36, 49 


jussieu, Sardinella, 48 


Konosivus punctatus, 33, 49 
thrissa, 36 


lecogaster, Sardinella, 48 


INDEX 


machnata, Argentina, 45 
Elops, 45, 48, Pl. 7 
Macrura sinensis, 48 


maculatus, Chatoessus, 35, 37, Pl. 5 
Megalops, curtifilis, 44, 45, Pl. 6 


cyprinoides, 44, 45, 48 
setipinnis, 44, Pl. 6 
melanosticta, Clupea, 27 
Clupea (Amblygaster), 28 
moluccensis, Harengula, 28 
mystax, Clupea, 42 
Engraulis, 42 
Setipinna, 42 
Tiwyssa, 42, 49; Ph. 6 
Mystus altus, 34 
ensiformis, 39 
mystus, Clupea, 39 
Cotlia, 39, 49 


nasus, Clupea, 30 
Coilia, 41 
Nematalosa, 36, 49 
Nealosa punctata, 33 
Nematalosa japonica, 36, 49 
nasus, 36, 49 
novacula, Ilisha, 32, 49 
nymphaea, Clupea, 26 
Harengula, 24 


okinawensis, Clupea, 48 

Opisthopterus tardoore, 49 
valenciennest, 49 

osbecki, Chatoessus, 35 


palasah, Alosa, 31, Pl.3 
Pellona ditchella, 48 
grayana, 32 
vimbella, 32 
playfairi, Choetomus, 41 
Cotlia, 41, 49, Pl. 5 
punctata, Nealosa, 33 
punctatus, Chatoessus, 33 
Herklotsichthys, 48 
Konosirus, 33, 49 
purpurescens, Elops, 45, Pl. 7 


veevestt, Alausa, 30 
Alosa, 30, Pl. 2 
ilsay 30, 31 
Hilsa (Tenualosa), 48 


Sardinella aurita, 26, 48 
brachysoma, 48 


53 


54 


Sardinella (contd.) 
bulan, 48 
clupeoides, 48 
fimbriata, 24, 48 
jussieu, 48 
letogaster, 28, 48 
sirm, 28 
saurus, Elops, 45 
schrammi, Herklotsichthys, 48 
Selipinna mystax, 42 
taty, 49 
setipinnis, Megalops, 44, Pl. 6 
setirostris, Thryssa, 49 
sinensis, Macrura, 48 
sirm, Sardinella, 28 
Spratella fimbriata, 24 
Spratelloides delicatulus, 30 
gracilis, 29, 48 
Stolephorus buccaneeri, 47, 49 
commersonit, 37, 49 
heterolobus, 49 
indicus, 49 
iri, 49 
zollingert, 47, 49 


INDEX 


tardoore, Opisthopterus, 49 

taty, Setipinna, 49 

teres, Etrumeus, 48 

thrissa, Clupanodon, 34, 49 
Clupea, 34 
Konosirus, 36 

Thryssa dussumieri, 49 
hamiltonii, 42, 49 
mystax, 42, 49, Pl. 6 
setivostris, 49 

toli, Hilsa (Tenualosa), 48 

tri, Stolephorus, 49 

triza, Chatoessus, 34, Pl. 4 
Clupea, 35 


valenciennest, Opisthopterus, 49 
vimbella, Pellona, 32 ‘a 
vulpes, Albula, 48 


zollingeri, Stolephorus, 47, 49 
zunast, Herklotsichthys, 48 


THE ELOPOID AND CLUPEOID 
FISHES IN RICHARDSON’S 
“TCHTHYOLOGY OF THE SEAS OF 
CHINA AND JAPAN” 1846 


P. J. P. WHITEHEAD 


¥ BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vok “4 ING. 72 
~ LONDON: 1966 


THE ELOPOID AND CLUPEOID FISHES IN 
RICHARDSON’S “ICHTHYOLOGY OF THE SEAS 
OF CHINA AND JAPAN” 1846 


BY 


P. J. P. WHITEHEAD 


British Museum (Natural History) 


Pp. 15-54; 7 Plates 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY WOl EA. INO. 2 
LONDON: 1066 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 


Parts will appear at irregular intervals as they become 
ready. 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. 


fee ELOPOID AND CLUPEOID. FISHES. IN 
RICHARDSON’S “ICHTHOLOGY OF THE SEAS 
OF CHINA AND JAPAN” 1846 


Wl el as ee a lh D 


CONA ENTS 
Page 
EXESTRACT . j ‘ : ‘ ‘ ; F : : : 17 
INTRODUCTION. ; : : : : j P 4 : : LF 
RICHARDSON’S REPORT ‘ : ‘ : ! ; : : 18 
THE REEVES ILLUSTRATIONS 2 : : ‘ : : ; : 19 
RICHARDSON’S SPECIES : ; : : : : y ‘ , 2 
List oF HONG KONG SPECIES 4 : . ‘ : ; oe Ae 
REFERENCES : : : 3 : “ : ; : : : 50 
INDEX : : 2 d : : : : : ; : : 52 
ABSTRACT 


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. 


INTRODUGIION 


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 PrP. J.P. WHITEHEAD 
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 Japomica 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 nomina 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 Giinther’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 


PICA DSON' S ELOPOID AND CLUPEHOID 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 
nymphaea (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 cyprinotdes, 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. (Lacepéde 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. 


¢ 


Tb ee VES TV LALS TRATIONS 


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 (941) is 
omitted in the Richardson set. 

According to one source (Anon., 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 


RICHAKDSON:S ELOPOID AND CLUPEOLD. 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 Miller 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. 


D> fhe™ 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 P.jJ.P. WHITEHEAD 


A list of the elopoid and clupeoid species mentioned by Richardson (1846) showi 


Species 
ELOPIDAE 


Elops machnata (Forssk.) 


MEGALOPIDAE 
Megalops cyprinoides (Brouss.) 


CHIROCENTRIDAE 
Chirocentrus dorab (Forssk.) 


DUSSUMIERIIDAE , : 
Spratelloides gracilis (Schlegel) 


CLUPEIDAE 
(Clupeinae) 
Sardinella aurita Val. 


Sardinella fimbriata (Val.) 
?Sardinella leiogaster Val. 


Sardinella or Herklotsichthys sp. 


(Alosinae) 
Hilsa reevesti (Rich) 


(Pristigasterinae) 
Ilisha elongata (Ham. Buch.) 
(Dorosomatinae) 
Clupanodon thrissa (Linn.) 
Konosirus punctatus (Schlegel) 
Nematalosa, ?N. nasus (Bloch) 


ENGRAULIDAE 
(Stolephorinae) 


Stolephorus commersonii Lacepéde 


Thryssa mystax (Schn.) 


(Coiliinae) 
Cotlia mystus (Linn.) . 
Cotlia playfairii (Rich.) 


Richardson, 1846 


(name) 


Elops machnata Forssk. 
Elops purpurescens Rich. 


2 egalops setipinnis Forster 


Megalops curtifilis Rich. 


Chirocentrus dorab Forssk. 


Clupea gracilis Schl. 


Clupea nymphaea Rich. 
Clupea isingleena Rich. 
Clupea caeruleo-vittata Rich. 
Clupea flosmaris Rich. 


[4 losa veevesii Rich. 


ee palasah Russell 


Llisha abnormis Gray 


Chatoessus maculatus Gray 
Chatoessus aquosus Rich. 
Chatoessus chrvsopterus Rich. 


ere triza Linn. 


Engraulis commersonianus Lac. 
Thryssa mystax Schn. 


Coilia grayii Rich, 
Coilia playfairii (McClelland), 


mec HARDSON-S!:ELOPOLID AND CLUPEOLD FISHES 


ibers in the three sets of Reeves drawings now in the British Museum (Natural History). 


Illustrations—page nos. 
oo 


la eer ae 
Reeves Richardson Hardwicke Reeves specimens 
97, 99 142 30 BMNH. 1964.11.6.3 
46 142 31 = 
68 141 234 BMNH. 1964.11.6.14. TYPE* 
BMNH. 1853.1.4.20 
27,97 I41 tiga ce 
33 142 237 BMNED, 19064: 13.6.1 
ai 136 222 BMNH. 1964.11.6.6—7 (TYPE lost) 
29, 95 136 219 BMNH. 1963.6.17.1 TYPE 
29, 95 136 223 ia 
29, 95 136 Ted i; 
fae : a BMNH. 1963.8.20.2 TYPE 
‘ 37 a BMNH. 1963.8.20.3 
96 127 22% BMNH. 1963.8.20.1 
| f 29, 48 138 upper = BMNH. 1964.11.6.4. TYPE 
100 138 lower 240 
53, 54 139 232 ei 
gI 139 223 Vachell specimen (TYPE lost) 
29, 95 139 230 BMNH. 1964.11.6.5. TYPE 
29, 95 139 231 —- 
= == == BMNH. 1964.11.6.8—11 
92/97 I4I 236 BMNH. 1064 oT 1.6.12 
8 140 252 f BMNH. 1855.9.19.1581. TYPE* 
‘\ BMNH. 1964501026 2e* 
31 140 — BMNH. 1847.5.10.5. TYPE* 


material, ex Haslar collection. 


24 P. J. P. WHITEHEAD 
RICHARDSON’S SPECIES 
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. 


rt. ‘‘ Clupea isingleena Richardson ”’ 
= Sardinella fimbriata (Valenciennes) 


Clupea isingleena Richardson, 1846, Ichth. China Japan : 304 (China Seas). 
Spratella fimbriata Valenciennes, 1847, Hist. Nat. Poiss. 20 : 359, pl. 600. 
Clupea (Harengula) 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 Hi/sa kelee (Cuvier) in shape but lacking any 
spots or marks along the flanks (PI. 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. 1), 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. 7singleena, 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. wymphaea ; 
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. isingleena the 
body depth is contained 2-75 times in standard length, against 3-65 times in the figure 
of C. nymphaea (3-1 in the actual specimen, “ thrice in the length” according to 
Kichardson’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 


RCH ALD SONS ELOPOID AND CLUBEOID 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 Gitinther’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. wymphaea) is known only in one species of Sardinella, 
namely S. auvita, which in fact is a slender species such as is described and figured for 
C.nymphaea. In S.auriia too, the anal count is low (15-19 fide Regan, 1917) and thus 
agrees with Richardson’s C. nymphaea (A.15 vel 16) not with his C. zsengleena (A.21). 

One slight anomaly, however, is in Richardson’s scute count of 16 + Io 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. zsingleena 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 Havengula by Regan (1917)) and 
subsequent workers (e.g. Fowler, 1941; Chu & Tsai, 1958). But it is clearly a 
member of Sardinella, having 8-10 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 (19170) 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 Savdinella in the Taiwan area, as well as Harengula 
nymphaea. Of the four with 8 pelvic rays and sharply keeled scutes (i.e. excluding 
S. aunta and S. sirm 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 jussieui 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 Lacepéde 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, %S.L. 

Body depth d R : ; ; t ; 35°2 32°4 
Head length . : é h ; ; ; 28 +3 26:2 
Snout length ‘ ; ; P . ; : 5°8 5°3 
Eye diam. ; ; ; . : : : 8-0 7°*4 
Upper jaw 1. ; . ; ; : ; . I1°9 Il-o 
Lower jaw l. ; ; ‘ ; : : 4 12°4 II*5 
Pectoral length . : ‘ ‘ i t damaged — 

Pelvic length . ‘ : : : , ; 11°8 10°9 
Pre-dorsal : ‘ ; i ‘ ; ' 49°1 45°3 
Pre-pelvic . : , : . : : / 59:2 54°5 
Pre-anal . ; ; ‘ : : : i 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 ; Ist 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 1 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 iii 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 : wpper } 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. 
yardinella aurita Valenciennes, 1847, Hist. Nat. Poiss. 20 : 263, pl. 594 (name retained for reasons 
of stability—-see below) ; Regan, 1917, Ann. Mag. nat. Hist. (8) 19 : 578 (including Giinther’s 


BRLiCHARDSON'S ELGCPOID 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 I juvenile 
ex China). 
Type. No British Museum specimen. As shown under C. zsingleena, the speci- 
men hitherto labelled as type of C. nymphaea (see Giinther, 1868, p. 428; Regan, 
IQI7, p. 392) 1s in fact the type of C. zsingleena. 


FIGURE. Reeves No. $25 shows a rather slender clupeoid resembling Sardinella 
sim or S. clupeordes (Pl. 1, 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 
15 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 6? 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 9g pelvic 
rays and this immediately excludes all species of Herklotsichthys (Harengula auct., 
see Whitehead, 1964a) 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. sym 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 Sardinella 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. 

Gunther (loc. cit.) placed C. caeruleovittata Richardson in his synonymy of 
C. melanosticta Schlegel. Richardson lists no specimens of C. caeruleovittata, so the 


28 P. J.P. WHITEHEAD 


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 leiogaster Valenciennes, 1847, Hist. Nat. Poiss. 20: 270; Kner, 1865, Reise Novara, 
Fische : 327 (Hong Kong). 
? Harengula 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. caeruleo- 
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 is none in 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. sirm 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 (g) from S. siym and S. leiogaster (8). But since 
‘. aurita is already represented in Richardson’s list (as C. mymphaea) 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 leiogaster seems the more likely. It is a slightly deeper fish 
and lacks the row of dark blue spots along the flanks described in S. sirm (see Ch 
& Tsai, 1958). 


Pichi win SON Ss fLOPOrD AND CLUPEOID FISHES 29 


Another possibility is S. clupeoides (Bleeker, 1849). Unlike 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 moluccensis of Jouan (1867) from Hong Kong may refer to the present 
species, although “le ventre non caréné”’ is suggestive of Etvumeus 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 Etrvuwmeus nor 
Sardinella. 

Richardson’s name caeruleovittata predates Valenciennes’ name lezogaster. 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. 


d 


4. “ Clupea flosmaris Richardson ’ 
= ?Herklotsichthys sp. or Sardinella sp. 


Clupea flosmaris Richardson, 1846, Ichth. 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 S#ratelloides 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: D13, A1g,C 14, P to, 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 9g, 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. 


Nore: 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. 


6. ‘“ Alosa reevesii Richardson ”’ 
= Hilsa reevesii (Richardson) 


Alosa reevesii 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 reevesii : 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: Richardson, 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, ex China, presented by J. R. Reeves, 
until recently unregistered, but now BMNH. 1963.8.20.2. Giinther (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. } 


Ficure. 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 (PI. 2, 
hg. 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. reevesii, not H. ilisha, having the broad oper- 
culum characteristic of the former (see Whitehead, 1965a). In addition, there are 


PLATE 1 


Fic. 1. Clupea isingleena Richardson, holotype (108-5 mm. S.L., BMNH. 1963. 

6.17.1). Formerly believed holotype of C. nymphaea, see text, p. 24. 
Fic. 2. Clupea isingleena, Reeves drawing No. 60 [= Sardinella fimbriata (Val.)]. 
Fic. 3. Clupea nymphaea, Reeves drawing No. £25 [= Sardinella aurita Val.). 


Bu BM .(N.1.) Zool. 14, 2 


Pah or 


Fic. I. 
Fic. 2. 


Fic. 3. 


PLATE 2 


Clupea caeruleo-vittata, Reeves drawing No. 59 [= Sardinella, probably 
S. leiogaster Val.}. 
Clupea flosmaris, Reeves drawing No. 64 |= Herklotsichthys or Sardinella 


sp.|. 
Alosa reevesii, Reeves drawing No. a8 [= Hilsa reevestt (Rich.)]}. 


Bull. B.M.(N.H.) Zool. 14, 2 PUA, 2 


Mii Rhian oe bagi «BBE a 2 


PLATE 3 
Fic. 1. Alosa palasah, Reeves drawing No. P51 [= Hilsa reevesti (Rich.)}. 
Fic. 2. Ilisha abnormis, Reeves drawing No. 81 [= Ilisha elongata (Bennett)}. 
Fic. 3. IJlisha abnormis, Reeves drawing No. 67 | = Ilisha elongata (Bennett) |. 


Bull. B.M. (N.H.) Zool. 14, 2 RATE: 3 
: 


* as me S oe 
oh Extn & i PAE eh Sacto 
5 


e ¥* 


Mk, abr Gah 306 


Hike Fe Sen tre g.. 3 


FIG. I. 


FIG. 2. 
FIG. 3. 


PLATE 4 
Chatoessus aquosus, Reeves drawing No. 63 |= Konosirus punctatus 


(Schl.)). 
Chatoessus triza, Reeves drawing No. 224 |= Clupanodon thrissa (L.)]. 
Chatoessus chrysopterus, Reeves drawing No. 61 |= Nematalosa, probably 


N. nasus (Bloch)}. 


Bull. B.M. (N.H.) Zool. 14, 2 PRATE 4 


: ‘ : fet 
Chushe fous tbo ye opel CAiaed : i) J 


i 
“s 7 


PLATE 5 
Fic. 1. Chatoessus maculatus, Reeves drawing No. 109 [= Clupanodon thrissa 


(L.)] 

Fic. 2. Coilia grayti, Reeves drawing No. a14 |= Coilia mystus (L.)). 

Fic. 3. Coilia playfairii, Reeves drawing No. B26 [= Coilia plavfairii (Mc- 
Clelland)). 


PLATE 5 


Bull. B.M. (N.H.) Zool. 14, 2 


FIG. I. 
FIG. 2. 


FIG. 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 cyprinoides 
(Broussonet) |. 


Bull. B.M. (N.H.) Zool. 14, 2 PEATE -6 


Magalife tliprerrris Qf: se 


ae 


Fic. I. 
FIG. 2. 


FG. 3. 


aki § 
4s 


af 
y 


? 
16.7 a wy 


a 


? 


4 ‘a ; 
a 


ao 
‘a 

me 'Y. 

- ‘ 
¢ 


74 
ae 
> 


me 


PLATE 7 ae 
Elops machnata, Reeves drawing No. 137 [= Elops machnata (For sskal)]. 
Elops purpurescens, Reeves drawing No. 53 [= Elops machnata— ex 


4 
se 
Pte 


us se 


a 
“i; 


y 


2 us 
sskal)}. ; 


Chirocentrus dorab, Reeves drawing No. 47 [= Chirocentrus dorab- 
(Forsskal)}. ~. Raye 


ay » 


+ 


a @ 


Bull. B.M: (N.H.) Zool. 14, 2 PLATE 7 


“3 
thujes P06 ts he Mall Ga Che, fe fa . By? 


Ele fon fr vs fit 22 ewe Lf ‘ 7: oo 


9 & Wd] 


ORespenet Fine ire, Ciiccx fro ker, 


'SPRESENTED> 
< ot, 
OGy \\ 


OBSERVATIONS ON THE TYPE- 
MATERIAL OF SOME GENERA AND 
SPECIES OF POLYZOA 


ANNA B. HASTINGS 


; BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
“ZOOLOGY Vol. 14 No. 3 
} LONDON: 1966 


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 Vol. 14 No. 3 
LONDON: 10966 


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 
ready. 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. 3 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 11 March, 1966 Price Thirteen Shilli 


OBSERVATIONS ON THE TYPE-MATERIAL 
eee ov GENERA AND: SPECIES. OF POLYZOA 


By ANNA B. HASTINGS! 


Page 

I. INTRODUCTION : : : ; ‘ ; . ; F 57 
2. CHALLENGER CoLtEction : é , : . : : sig 
3. SOME TYPE-SPECIMENS . : : : Bens) 
Bifaxaria corrugata Busk aad B. Aaa Busk : ; : « +58 
Harmer’s elucidation of the Challenger Report : : eo 
Lectotypes . : ; ; ‘ i : ‘ : 5 50 
Systematic position ‘ , : ‘ : ‘ : esc 
Comparison of the species ‘ : ‘ ; : : = ~ 59 

Identity of the Siboga material : : ‘ : ‘ S ROE 

Flustra octodon Busk . ; : ; ‘ ‘ Af 62 
Lepralia melolontha Denrenocoden: : : : : : OS 
Lepralia multidentata Thornely . : : : : : alae. c 
Lunulites owenii Gray . : ; : : : ; ; i) "Om 
Membranipora amplectens Hincks : ; é é 2p aOR 
Membranipora crassimarginata var. erecta Busk : . ‘ sre yO5 
Membranipora hexagona Busk : : i ‘ ; «2, » OS 
Membranipora pilosa var. multispinata Hineks : : ; owe OF 
Membraniporella agassizii Smitt . ; : : ‘ : . 68 
Menipea fuegensis Busk : : : ' : : ; 468 
Micropora norman Levinsen : : : ‘ é ; iN" 6G 
Salicornaria clavata Busk . b ; ‘ ‘ ‘ : ; 71 
Siphonoporella nodosa Hincks . : Ge 

4. THE GENUS HIPPOPLEURIFERA AND SOME OF ITS SPECIES . eee 
Hippopleurifera Canu & Bassler . ; : : : : eS 

H. biauriculata (Reuss) : : ; : : : : BN aed 2) 

H. pulchra (Manzoni) . : : 3 : , ; 5h FS 

H. sedgwicki (Milne- davaads) ' 3 3 ; ; : sre ES 

5. ACKNOWLEDGEMENTS . : : : : : ; : ee a 
6. SUMMARY : ; ‘ : ; ; ‘ : ; ; ‘ 75 
7. REFERENCES . ; : : : : 4 4 : : : 76 


tr IN PRODUCTION 


In this paper I have gathered together the results of the examination of the type- 
specimens of a number of species. 


2: (EA UDENGER 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 


* 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 B. 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. 196. 

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). 

* It 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, 


OBsSERVATAZONS ONS TYPE-MATERIAL, POLYZOA 59 


eta 12.9.370, 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 
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 Bb. 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. Ihave 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. Im 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). 

I 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 zoartum 
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. 1c), 
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. IA), 
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. papillatus the pores are peripheral, 
arranged in one or two series. Sutures or cracks run outward from the median line, 
most of them ending in a pore. In S, corrugatus the pores are central, arranged on 

ZOOL, 14, 3 6§ 


60 A. B. HASTINGS 


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. 1A). In S. papillatus it is the develop- 


| 050mm | 


Scale for fig B 


' 0 50mm 


Scale for fig A.C 


Fic. 1. 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. 
\. S. corrugatus, 1887.12.9.379A, lectotype. B, c. S. papillatus, 1964.2.2.4, Siboga colle¢- 
tion, specimen 583A", B. is the large avicularium which is seen obliquely in c, 


OBSERVATIONS ON’ TY PE-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 exact1, 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). 


__* Miss Cook (in litt.), “‘ I have made several sets of measurements and the agreement is quite startling. 
Exact ’ is certainly the right word ”’, 


62 A. B. HASTINGS 


Flustra octodon Busk 


Flustra octodon Busk, 1852 : 49, pl. LVI, fig. 4, pl. LVIII, fig. 5. 
Hincksinoflustra octodon Bobin & Prenant, 1961: 167 (as Flustra octodon), 169, 170, text-fig. 

4I-IV. 
Hincksina (Flustra) octodon (Busk) Gautier, 1962: 51. (Hincksinoflustra 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. 1963.3.12.1 (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), Serupocellaria macandret (p. 24) and Flustra octodon (p. 49) ; from the ‘ Bay 
of Gibraltar ’’, Membranipora rossellii (p. 59), M. calpensis (p. 60), Lepralia violacea 
(p. 69), L. spinifera (p. 70) ; from the “ Mediterranean ’’, Lepralia linearis (p. 71), 
Eschara foliacea (p. 89). 

The slide of A. lafontii (1899.7.1.3752) has labels superimposed on the old ones, 
as was Busk’s custom'. The original label reads “‘ Coast of Spain’, 
‘ 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, 1899.7.1.2310. 
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 Flustra octodon, illustrated in pl. LVI, fig. 
(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 Hincksinoflustra Bobin & Prenant (1961). 


' His specimens were relabelled, sometimes more than once, in accordance with changes in the style 0 
label in use in his collection ; for further details see Hastings, MS, note in Busk Register, British Museu 
(Natural History). 


a 


OBSERVATIONS“ ON TYPE-MATERTIAL, POLYZOA 63 


Lepralia melolontha Landsborough 


Lepralia mtida, branched in a fine dendritic manner, Johnston, 1847: 319. 

Leprahia mtida, specimen d., “ Lep. nitida var.’’, Gray, 1848: 121. 

Lepralia melolontha Busk; Landsborough, 1852: 319, pl. XVIII, fig. 70; Busk, 1854: 79, 
pl EXAXRXY, fig. 3. 

Membraniporella melolonitha Busk Hincks, 1880a: 202, pl. X XVII, figs. 9, 10. 

Aspidelectva melolontha Levinsen, 1909 : 160. 

Aspidelectva 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 
Permian Coll), 191l.10.1.741 part, 1919.6.24.25, 25a,: 26); Goodwin Sands, 
Kent (G. Merritt, 1884.5.10.1) ; Kirkwall, Orkney (Johnston Coll., Gray 1ar1d, 
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), is not in the Museum, but would 
not be a syntype. 

Hincks (p. 203, footnote) rican that Mrs. Gatty (in Landsborough) was right in 
identifying Johnston’s dendritic variety of Lefralia nitida from Kirkwall, Orkney 
Islands, with this species. Marcus (p. 201) noted the Orkney record as being un- 
certain but notimprobable. 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 Electra 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., 1911.10.1.48r 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. 10) 
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 Asfidelectra 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. 


DISTRIBUTION. 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. 


HototyPe, 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 owenti Gray 


Lunulites owentt Gray, 1828 : 8, pl. III, fig. 15. 
Cupuladria owenti (Gray) Cook, 1965 : 213, pl. 2, figs. 3A, B, text-fig. 2c. (Synonymy.) 


DistTRIBUTION. Recent, West Africa. Pliocene, South Carolina; Florida. Mzto- 
cene, North Carolina; Florida. (Teste Cook). 


LectTotyPE, chosen here: 1899.7.1.4879, W. Africa. 


The slide 1899.7.1.4879, which forms part of the Busk Collection, is labell 
“ B.M.. J. E. Gray’, indicating that Busk received the specimen from the Briti 
Museum. It is the only representative of Gray’s material of this species now in 
Museum and Kirkpatrick (MS. Catalogue) accepted it as the type-specimen. It i 
the specimen figured by Busk (1854, pl. CXV, fig. 3). 


REMARKS. Cupuladria owenit is discussed by Cook. 
' 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 


Membranipova amplectens Hincks, 1881a: 129, pl. III, fig. 7. 

Electra amplectens (Hincks sp.) MacGillivray, 1889: 322, pl. CLXXXVII, figs. 14, 14@; 
Levinsen, 1909: 66, 146, 147, pl. IX, figs. ta, b. 

Membranipora amplectens sp; n. [sic] Hincks, 1892 : 332. (Named as “ type of a new genus ”’. 
Introduction of Heterooecium follows.) 

Membrampora 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 
Pore tooo.1t.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 
mame (see below). They are: 1887.12.9.310, 311; 1899:7.1.1004, X005; 
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. MM. crassimarginata var. erecta is synonymous with Acan- 
thodesia perfragilis (MacG.), see Hastings (1945). 


. ‘ Salicornaria clavata to be excepted. The Heard Island species is distinct from the Australian, see 
elow. 


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. 


HoLotyPe, the only specimen: r899.7.1.1083. 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 (1881): 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 7. hexagona has been recorded from various localities : 

Busk himself gave Flustra coritacea Johnston (not Esper) as a synonym of M, 
hexagona. IF. 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 (Miill.). 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 (1g21 : 204) recorded Pliocene fossils from Sicily as Membranipora exagona 

' No other evidence has, however, been produced for thinking that Johnston confused two species 
under Flustra coriacea. _\t appears that only the Sana Island specimens came to the British Museum i 
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 Flustra coriacea Johnston not Esper. The exist- 


ence of a second British species of true Micropora (M. normani see below) does not affect this question of 
whether Johnston may have confused a Membraniporine species with M, coriacea. 


OBSERVATIONS ON 'TYPE-MATERIAL, POLYZOA 67 


[sec] 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 Membranipora 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. 1897.5.1.49I, 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 E. 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 


Membraniporella agassizit Smitt, 1873: 11, pl. V, figs. 103-106 ; Levinsen, 1909: 17 ; Hastings, 
1964 : 258, 262. 


HoLotyPe, 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. 4 
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 Membraniporella agassizii 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 Membrantporella and the taxonomic significance of the multi- 
plication of avicularia and kenozooecia is still unknown (cf. Hastings, 1964 : 258). 


Menipea fuegensis Busk 


Memipea fuegensis Busk, 1852: 21, pl. XIX, figs. 1-3. 

Menipea patagonica Busk (part), 1852: pl. XXIII, fig. 1 (not p. 22, pl. XXV, figs. 1-3, pl 
XXVII, figs. 1, 2). 

Tricellaria aculeata (d'Orbigny) ; Hastings 1943: 356. (Synonymy and distribution.) 


DISTRIBUTION. Widely distributed in sub-Antarctic waters. ; 
LectotTyPeE, chosen here: 1854.11.15.262. Falkland Islands. H. [Hooker]. 


REMARKS. Busk recorded this species from Tierra del Fuego (Darwin) and the 
balkland Islands (Hooker), but the only type-material in the British Museum is the 
one specimen now chosen as lectotype. This specimen was mounted and labell 
by Busk as Menipea fuegensis, and sent to the Museum as part of the “ Britis 
Museum Catalogue Collection ’’, for which see Hastings (1943 : 304). As noted b 
Hastings (1943 : 359) the ancestrula and first few zooecia of the type were figur 


OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 69 


by Busk (pl. XXIII, fig. 1), but, by some curious error, under the name Menipea 
patagomca. There is no evidence as to the locality of the material figured in pl. XIX. 
Mentpea fuegensis Busk is a junior synonym of Tvricellaria aculeata (d’Orb.). 


Micropora normani Levinsen 


Micropora sp. Levinsen, 1902 : 7 (footnote). 

Micropora normani Levinsen, 1909 : 162, pl. VIII, figs. 3a, 3b (M. cortacea on plate, M. normani 
in explanation of plate). 

Micropova coviacea 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: Igir.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. normani; Norman Coll., 
7O11.40.14.744); 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. coriacea!, 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. normam. 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. normanv. 

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. corvacea 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 


1 M. coriacea Johnston not Esper, see footnote above under Membranipora hexagona. 
* See Hastings (forthcoming paper on Hippothoa). 


x0 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.normani: 
I2—I5, average 20. 

On the evidence available to me, M. normani 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. normam. M. cortacea 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: II), extending 
from Spain to the western and south-western shores of the British Isles. ’ 

CHARACTERS OF THE GENUS Micropora. 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. normanz. 

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 zooeci 
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 
species (undescribed?) from Algeria (1869.10.6.6) has, on many zooecia, a pair 
stout distal spines, rather widely set (being about opposite to the proximal corne 
of the distal zooecium). More rarely there is one median distal spine. 


' The term recumbent was introduced by Canu & Bassler (1917 : 66), see Brown (1954 : 244, footnote) 


O@bsERVALIONS ON*TYPE-MATERIAL, POLYZOA Wisi 


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 australis MacGillivray Waters (part), 1888: 16, pl. 11, figs. 1-4. 

Cellaria clavata (Busk) Hastings, 1947 : 236. 


DISTRIBUTION. 5S. Indian Ocean. 
LECTOTYPE, chosen here : 1887.12.9.395 part. Challenger St. 151, Heard Island. 


EPARALECTOTYPES. 1887.12.9.391, 396; 1809.7.1 71545 3 1034.2.16.34. St. 
men iereuelen. 1682.7.29.75; 1887.12.9.389 ; 1899-7:1:1550 ; 1934.2.16.35. 
pipei5t, Heard Island. 1887.12.9.390, 393; 1899-7.1.1551. Prince Edward 
Island. 


USHER MATERIAL. St..1491) (1063.2.12.200) ; 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 
a lectotype. 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. x 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 0-0g— 
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 0-15 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. 


‘ 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, 


72 A. B. HASTINGS 


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 0-0g-0-1I mm., average 0-10 mm. lo 0-TI-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. 


DISTRIBUTION. Australia. 


LEecTOTYPE, 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 
(E. 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. 


ye, the nodule in the corner next to the other zooecium of the pair, 


@BseivVALTIONS ONvTYPESMATERIAL, POLY ZOA 73 


meaeeeGENUS AIPPOPLEURIFERA AND SOME OF ITS SPECIES 


Hippopleurifera Canu & Bassler 


Hippopleunifera Canu & Bassler, July 1925: 679. (Hippopleurifera biauriculata Reuss, 1847 
[sec] 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 Eschara 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.) 

Hippopleunfera Canu & Bassler, 1927: Bassler, 1935: 127. (Listed with a reference to Canu & 
Bassler, 1929, and with E. sedgwicki as type.) 

Hippopleurifera Canu ; Hastings, 1949: 524. (Discussed with EF. sedgwicki as type.) 

Hippopleunifera Canu & Lecointre, 1925; Vigneaux, 1949:96. (Same French definition and 
type as Canu & Lecointre, 1930.) 

Hippopleunifera Canu & Bassler, 1925; Lagaaij, 1952: 92. (Revised diagnosis with Eschara 
biauriculata Reuss named as type by monotypy.) 

Hippopleurifera Canu, 1927 ; Bassler, 1953: G196. (Brief definition with E. sedgwicki as type.) 

Hippopleurifera Canu & Bassler, 1924; Buge, 1957: 265. (Redefined with EF. biauriculata as 
type.) 

Hippopleurifera Brown, 1958: 65. (Footnote on priority of E. 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. 1-3). 


Eschava biauriculata Reuss (part), 1848 : 66, pl. VIII, fig. 15 ; Manzoni, 1877: 11, pl. IX, fig. 29. 
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). 


LEcTOTYPE, 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: (r) 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. * 3-1 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 zooecium. 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). 


REMARKS. H. biauriculata (P\. 1, figs. 1-3) has more in common with H. pulchra 
Manzoni (see Hastings, 1949, pl. XII, fig. 2) than with H. sedgwicki (see Pl. 1, figs. — 
4, 5, and Hastings, 1949, pl. XII, fig. 1). It resembles H. pulchra 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. pulchra in the smoothness of the frontal wall between 
the pores, which are larger; and in the shape and position of the avicularia. In 
H. pulchra 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. inauriculata, 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 et 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 
I rigonopora which also has fossac., 


OBSERVATIONS ON TYPE-MATERIAL, POLYZOA 75 


Hippopleurifera pulchra (Manzoni) 


Cellepova 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 souliert Calvet ; O’Donoghue & de Watteville, 1939: 28. (Record only.) 

Hippopleurifera pulchra (Manzoni) Hastings, 1949: 521, pls. XII, figs. 2-4, XIII, figs. 1, 1a, 2, 
2a (Synonymy); Gautier, 1962 : 189. 

Hippopleurifera granulosa Canu ; Vigneaux, 1949:97. (Record only.) 
DISTRIBUTION. Recent, Mediterranean. Muocene, 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 Hippodiplosza 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 : 140). 


Hippopleurifera sedgwicki (Milne-Edwards) 


(PI. 1, figs. 45, 5) 
Eschara sedgwicki Milne-Edwards, 1836 : 330 (sep. p. Io), 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). Muzocene, 
Austria. 


REMARKS. The paralectotype of Eschara biauriculata, referred above to Hippo- 
pleurifera sedgwickt, is from Eisenstadt, and extends the known range of H. sedgwick, 
both geologically and geographically. 

Buge redescribed this species. 


5. ACKNOWLEDGEMENTS 

I am grateful to the Manchester Museum, the Naturhistorisches Museum, 
Vienna, and the Riksmuseum, Stockholm, for lending specimens discussed in this 
paper ; to Professor D. A. Brown who, on his own initiative, borrowed the type- 
material of Eschera biauriculata, and brought it to me from Vienna; to Mr. M. G. 
Sawyers for the photographs; to Miss P. L. Cook for her assistance, including 
drawings, measurements and the compilation of the list of references ; and to my 
husband, Dr. H. Dighton Thomas, for his encouragement and counsel. 


6. SUMMARY 
Examination of type-material of a number of species has yielded information on 
their taxonomic status, distribution, structure, etc. The type-species of the genus 
Hippopleurifera is included, and one of the Siboga species of Sclerodomus is recon- 
sidered. Membranipora hexagona Busk was based on a single, mutilated specimen 
of an unidentifiable, non-membraniporine species. 


76 A. B. HASTINGS 


7. REFERENCES 

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Basster, R. S. 1935. Bryozoa (Generum et Genotyporum Index et Bibliographia) in Quen- 
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Brown, D. A. 1949. On the Polyzoan Genus Hippomenella Canu & Bassler and its Genotype 
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—— 1958. Fossil Cheilostomatous Polyzoa from South-West Victoria. Mem. Geol. Surv. 
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Buce, E. 1957. Les Bryozoaires du Néogéne de l’ouest de la France. Mém. Mus. Nat. Hist. 
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Busk, G. 1852. Catalogue of Marine Polyzoa in the ... British Museum, Part I, London. 

—— 1854. Op. cit., Part II. 

—— 1856. Zoophytology. Quart. J. micr. Sci., 4: 308-312. 

1884. Report on the Polyzoa. The Cheilostomata. Rep. Zool. Chall. Exp., 10, pt. 30. 

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Canu, F. 1916. Les Bryozoaires fossiles des terrains du Sud-Ouest de la France, IX. Aqui- 
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—— 1926. Contributions a ]’étude des Bryozoaires fossiles. Jbid., 25: 763-766. [For date 
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Canu, F. & Basster, 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. 
Fr., 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: li—xi, 1-685. 

Canu, F. & Lecorntre, 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. 

CrpoLLa, F. 1921. I briozoi pliocenici di Altavilla presso Palermo. G, Sci. nat. econ. Palermo, 
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Coox, 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. 

FiscHER, 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 l'Université d’Aix-Marseille, 
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Cheb y AVrONSSON TY PE-MATERIAL, POLYZOA 73 


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. 

HaGeEnow, F. von. 1839. Monographie der Ritigenschen Kreide-Versteinerungen. Leonhard 
und Bronn’s Neues Jahrbuch. f. Min., 1839 : 253-206. 

HARMER, S.F. 1900. A Revision of the Genus Steganoporella. Quart. J. micr. Sci., 43 : 225- 

297. 

1902. On the Morphology of the Cheilostomata. Ibid., 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. 

Hastines, 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 crassimarginaia auctt., with 

Remarks on some Genera. Ann. Mag. nat. Hist. (11) 12 : 70-103. 

1947. Op. cit—III. On some species of Cellaria ... . I[bid., 13 : 217-241. 

1949. On the Polyzoan Cellepora pulchra Manzoni and the genus Hippopleurifera. J. 

Linn. Soc. London (Zool.), 41 : 521-528. 

1964. The Cheilostomatous Polyzoa Neoeuthyris woosteri (MacGillivray) and Reginella 

doliaris (Maplestone). Bull. Brit. Mus. (N.H.) Zool., 11, 3 : 245-262. 

Hincks, 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. 

—— 1601a@. Op. cit., VI. Polyzoa from Bass’s Straits. Ibid., (5) 8: I-14,. 122-120. 

—— 1881b. Op. cit., VII. Foreign Membraniporina (third series). Jbid.: 129-132. 

—— 1882. Op.cit.,IX. Foreign Cheilostomata (Miscellaneous). Ibid. (5) 9: 116-127. 

p—— 1892. Op. cit., Appendix. Jbid., (6) 9: 327-334. 

= 1895. [O>. cit.], Index: i—vi, London. 

Jounston, G. 1847. A History of the British Zoophytes, Ed. 2, 2 vols., London. 

LaGaalj, 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. 

LaNDsBoROUGH, D. 1852. A Popular History of British Zoophytes, or Corallines, London. 

LEvinsEN, G.M. R. tI902. Studies on Bryozoa. Vidensk. Medd. dansk naturh. Foren. Kbh. : 


I-31. 
1909. Morphological and Systematic Studies on the Cheilostomatous Bryozoa: 1-431, 
Copenhagen. 


MacGiiiivray, P. H. 1889. Polyzoa in McCoy, F., Prodromus of the Zoology of Victoria, 
Melbourne, 1879-1890. Decade XIX : 307-323. 

Manzoni, A. 1870. Bryozoi fossili Italiani. (4acontr.). S.B. Akad. Wiss. Wien, 61, 1 : 323- 

349. 

1877. Briozoi fossili del Miocene d’Austria ed Ungheria, Part II... Denkschr. 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, Sct. nat., sér. 2, 6 : 321-345. 

Moore, H. B. 1937. Marine Fauna of the Isle of Man. Proc. Lpool. biol. Soc., 50: 5-293. 

Nevianti, 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: 115-2606. 


78 A. B. HASTINGS 


O'DonoGuue, C. H. & bE WATTEVILLE, D. 1939. The fishery grounds near Alexandria. 
Bryozoa. Note & Mém. Fouad I Inst. Hydrobiol. & Fish. 34: 1-58. 
Purcuon, R. D. 1937. An ecological study of the beach and the dock at Portishead. 
Bristol Nat. Soc., (4) 8: 311-329. 
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wiss. Abhandl., IT, Vienna (1847 sep.). [Vol. II was published 21 Aug. 1848, see Vol. III, 
‘“* Vorbericht "’.] 
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to Japan... Ark. zool. 33A, 12 : I-130. 
Smitr, F. A. 1867. Kritisk férteckning 6fver Skandinaviens Hafs-Bryozoer. 3. Ofvers 
Vetensk-Akad. Foérh. Stockholm, 24, 5 : 279-429. 
1873. Floridan Bryozoa, Part 2. K. svenska VetenskAkad. Handl., 9, 4: 3-83. 
Tuorney, L. R. 1905. Report on the Polyzoa...in Herdman, W. tte Rep. Pearl om e7 
Fisheries, Gulf of Manaar, 4, Suppl. Rep. 26 : 107-130, London. z 
ViGNEAUX, M. 1949. Révision des Bryozoaires néogénes 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 Riiger 
veréffentlichen Bryozoen. Geologie, 8, 25 : 1-80. 4 
1960. Sur les différents stades de l’astogénése 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., 
pt. 79. 
——— 1898. Observations on Membraniporidae. J. Linn. Soc. London (Zool.), 26: 654-6 
1924. The Ancestrula of Membranipora pilosa, L., and of other Cheilostomate 
Bryozoa. Ann. Mag. nat. Hist. (9) 14: 594-612. 
—— 1926. Ancestrulae and Frontal of Cheilostomatous Bryozoa. Ibid. (9) 17: 425-439. 
WIESEMANN, G. 1963. Untersuchungen an der Gattung Beisselina Canu 1913 und dhnlick 
Bryozoen (Maastrichtien, Danien, Montien). Mitt. min.-geol. (St.) Inst. Hamb., 32:5 


PLATE 1 
Fics. 1-3. Hippopleurifera biauriculata (Reuss). 


Fics. 1,2. The two faces of the bilaminar lectotype. Fig.1. The face figured by Manzo 
showing the edge of the frond (left) and stages in the secondary calcification of the ovice 


Fic. 3. Part of fig. 1 enlarged, showing two biauriculate fertile zooecia, and two with one 
the avicularia large and spatulate ; variation in older, secondarily calcified ovicells ; and p 
of the outwardly facing zooecia at the edge of the frond. 


Fics. 4, 5. Hippopleurifera sedgwicki (Milne-Edwards). The two faces of the bilam 
paralectotype of H. biauriculata (Reuss.). 


Bigs; 1:2). 4s-Soo MFO Fig. 3: xX 36. 


Pa Aw Ee 


oe) 
+ 
S| 
ns 
js) 
So 
NX 
= 
oS 
= 


Buill. B. 


MARINE NEMATODES FROM 
- DURBAN, SOUTH AFRICA 


WILLIAM G. INGLIS 


> 4 MAR 1966 | 


_ BULLETIN OF 
‘THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY bs Vol. 14 No. 4 
g LONDON: 1966 


MARINE NEMATODES FROM DURBAN, 
SOUTH AFRICA 


BY 


WILLIAM G. INGLIS 


Pp. 79-106 ; 47 Text-figures 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 14 No. 4 
LONDON: 1966 


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(NATURAL HISTORY), instituted in 1949, ts 

issued in five series corresponding to the Departments 

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within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
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This paper ts Vol. 14, No. 4 of the Zoological series. 
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Issued 23, March, 1966 Price Eleven Shillings 


MARINE NEMATODES FROM DURBAN, 
SOUTH BVPRICA 


By WILLIAM G. INGLIS 


SYNOPSIS 


Ten new species and three new genera of free-living marine nematodes are described from the 
Durban area of the Republic of South Africa, thus: Platycoma sudafricana sp. nov. is charac- 
terized by an arrow-head-shaped flap over the amphid and, possibly, by some features of the 
male tail; Tvileptium longisetosum sp. nov. is similar to T. ayum Inglis, 1964 but differs in the 
distribution of the cervical setae and the form of the spicules; Mesacanthion frica sp. nov. is 
characterized by the form of the spicules which have long ventro-posteriorly directed processes 
on their anterior ends ; Epacanthion oliffi sp. nov. is a short spiculed form characterized by lacking 
a gubernaculum and pre-cloacal supplement and having many cervical setae in the male ; Rhab- 
dodemania dura sp. nov. is characterized by a lack of ventro-lateral onchia and the shape of the 
spicules; PHERONOUS ogdeni gen. et sp. nov. of the Ironidae is characterized by a lack of 
caudal glands and papilla-like cephalic sense organs; Metoncholaimus murphyi sp. nov. 1s 
characterized by having one pore to the demanian system, extremely long spicules and a few long 
circum-cloacal setae; WIESONCHOLAIMUS mawsonae gen. et sp. nov. of the Oncholaimidae 
is characterized by long, equal, ventro-lateral onchia and a short dorsal onchium, a long stout 
tail, very long slim spicules and a large gubernaculum, and the presence of a demanian system ; 
PLECTOLAIMUS juliani gen. et sp. nov. of the Leptolaimidae is characterized by six distinct 
lips, ventral onchia-like structures at the anterior end of the oesophagus and a distinct bulb-like 
region with tiny valves at the posterior end of the oesophagus ; Bathylaimus deconinchi sp. nov. 
is characterized by a large relatively posterior amphid and by the shape of the spicules and 
gubernaculum. Hyalacanthion Wieser, 1959 is a Synonym of Epacanthion Wieser, 1953 because 
E. buischlit (Southern, 1914), type species of Epacanthion, has typical Hyalacanthion mandibles. 
As a further consequence it is argued that subdivision of the families of the Enoploidea is not 
yet possible, in spite of De Coninck’s recent attempt, particularly as De Coninck refers Hyala- 
canthion to one subfamily and Epacanthion to another. 


INE RODUCTION 


THREE samples of free-living marine nematodes were sent to me by Dr. W. D. Oliff, 
South African Council for Scientific and Industrial Research, National Institute for 
Water Research. These samples contain ten species, all of which are described below 
as new, referable to ten genera, of which three are new. As usual only those species 
represented by males in good condition are described. The species were collected 
from the following localities : 


Beach sand in the surf zone from the Durban area on I0.viii.1964. (Collection 
number: B8n8.) 


Ig. Bathylaimus deconinchi sp. nov. 
24,99. Epacanthion oliffi sp. nov. 
ZOOL. 14, 4 78 


82 WILLIAM G. INGLIS 


1 3. Mesacanthion frica sp. nov. 

3 5,12. Metoncholaimus murphyi sp. nov. 

12 3, 169, 4 larvae. Pheronous ogdeni sp. et gen. nov. 
2 3. Platycoma sudafricana sp. nov. 

1 3. Plectolaimus juliani sp. et gen. nov. 

3. 5,18, 12 larvae. Trileptium longisetosum sp. nov. 


Sediments in the mouth of Durban Harbour (polluted) on 20.vi.1964. (M23n4.) 


Ir g,6%, rlarva. Rhabdodemania dura sp. nov. 
6 3,12 2. Weesoncholaimus mawsonae sp. et gen. nov. 


Irom the same locality as M23n4 on same date (M23nr1). 


2 3, 11 9, 8 larvae. Wéiesoncholaimus mawsonae sp. et gen. nov. 


SYSTEMATIC CONSIDERATIONS 


De Coninck (1965) has made a heroic attempt to provide a more reasonable classi- 
fication of the free-living nematodes in which he divides the Superfamily Enoploidez 
into six families and fifteen subfamilies. Although some of these divisions appear to 
be reasonable others are certainly not at present acceptable. For example he still 
recognizes Thoracostomopsidae as a distinct family with only one genus Thoracoste 
mopsis. But this genus is only a somewhat modified Enoplidae which is well withi 
the ranges of variation shown by the genera currently referred to that family. 
Further, De Coninck refers Hyalacanthion Wieser, 1959 to one subfamily of the 
Enoplidae (Enoploidinae) and refers Epacanthion Wieser, 1953 to another (Enoplo- 
laiminae). But I show below that the genera cannot be separated. 

I argued recently (Inglis, 1964*) that Enoplus is a highly evolved form, while De 
Coninck treats it as a form ancestral to all the genera he refers to the family Enoplidae, 
but gives no indication of the origins of Enoplus itself. I have further argued, and 
still do, that the Enoplidae can be considered as having arisen from the genera 
commonly referred to the Family Phanodermatidae. In the latter family bucce 
rods appear to be commonly present and it is easy to imagine how, by a process of 
increased sclerotization, the region of the inner surface of the lip between these rods 
could become thickened to produce the mandibles so characteristic of the Enoplidae. 
Within the Enoplidae several lines can be crudely recognized along each of which the 
mandibular : onchial complex has been modified in a different way. The problem 
is not that of interpreting the components of each line but simply that we do not have 
enough information to make such an attempt worth while. We can only await the 
collection of more data and until then I still propose to recognize no groups other 
than Families and Genera. 


* In this paper read cephalic ventricle for cephalic vesicle throughout. 


MARINE NEMATODES FROM DURBAN 83 
DESCRIPTIVE SECTION 
Family LEPTOSOMATIDAE 
Platycoma sudafricana sp. nov. 
(Text-figs. 1-6) 


MATERIAL STUDIED. 2 3. Beach sand in the surf zone. Near Durban, Republic 
of South Africa (B8n8). B.M. (N.H.), Reg. Nos. 1965.961-962. 


A B C Body length (mm.) 
146°38 O° 25 56-10 IO'I 
144°74 8°53 68 +75 LEE 


MEASUREMENTS (in mm., in order of body lengths) 

Body breadth: 0-069; 0:076. Ocesophagus length: 1-23; 1-29. Diameter of 
head: 0-031; 0-029. Length of cephalic setae (long/short) : 0-023/0-018 ; 0:023/ 
0-018. Distance from anterior end of body to posterior edge of amphid: 0-026 ; 
0-027. Length of cervical setae: 0-014—-0:018; 0-014-0-017. Distance of nerve 
ting from anterior end of body: 0:30; 0:29. Length of spicules: 0-163; 0-165. 
Length of lateral piece of gubernaculum: 0-033; 0-035. Length of apophosis of 
gubernaculum: 0-015; 0-014. Distance of first pre-cloacal supplement anterior 
to cloacal opening: 0-087; 0-122. Distance of second pre-cloacal supplement 
anterior to cloacal opening : not measured, tail too curved ; 0-198. Length of tail: 
0:18; 0-16. Cloacal diameter: 0:066; 0-060. 


The anterior end of the body is set off as a constricted region starting about the 
level of the posterior edge of the amphids (Text-fig. 1). The mouth opening is simple, 
tri-radiate and leads almost immediately into the oesophagus which carries three 
small wholly cuticular onchia on each sector (Text-fig. 1). The cephalic sense organs 
lie in two circles of which the inner consists of six prominent papillae while the outer 
consists of ten setae of which six are long and four are shorter (Text-fig. 1). There 
are many long cervical setae. The amphids are prominent with a large arrow-head- 
like flap of cuticle developed from their anterior edge (Text-figs. r and 4). The 
oesophagus is long and simple in outline. 

The tail bears a pair of prominent spike-like setae about half-way along its length 
and a long but slim seta just posterior to the cloacal opening (Text-fig. 2). There is 
no spinerette and there are no caudal glands. On the mid-ventral surface of the 
body anterior to the cloacal opening there are two pre-cloacal supplements (Text-fig. 
2) of which the more posterior (Text-fig. 6) is a rounded cap-like structure resembling 
a Type-I campaniform organ (Inglis, 1963) while the more anterior (Text-fig. 5) is a 
raised area on which are located three pairs of short, stoutish setae. 

The spicular : gubernacular complex is double. That is, all the structures shown 
in the figure (Text-fig. 3) are represented by exactly equivalent structures on the 
other side of the body and they are in no way connected across the mid-line of the 
body. Further, the entire complex appears to form one functional unit. Here I 
interpret the structures as Gerlach (1955) does and not as he re-interprets them later 


84 WILLIAM G. INGLIS 


(1959). That is I consider the largest L-shaped structure to be the spicule which is 
enfolded posteriorly by a slightly triangular-shaped gubernacular apophosis. n ne 
most interesting feature is the relationship between the spicule and the rod-li 
structure bordering it anteriorly (considered a spicule by Gerlach (1959)). This rod 
swells towards its distal end where it bears a lateral flange which enfolds the distal 


Fics. 1-6. Platycoma sudafricana sp. nov. Fig. 1. Lateral view of head. Fig. 2. Gene 
lateral view of male tail. Fig. 3. Detail of spicular : gubernacular complex from right side 
Note the articulation between the spicule and the anterior part of the gubernaculum. Fig. 4 
Variant form of amphid. Fig. 5. Ventral detail of more anterior pre-cloacal supplemen 
Fig. 6. Ventral detail of more posterior pre-cloacal supplement. > 


end of the spicule while the same rod swells to form a slight knob proximally which 
fits into a socket in the spicule. 


Discussion 


This genus appears to be characterized by the peculiar modification of the spicul: 
apparatus, the presence of pre-cloacal supplements, the presence of the pair of stout 
spine-like setae on the tail and in lacking caudal glands. In addition the head 1 
highly characteristic with three small onchia and the marked sexual dimorphism ¢ 
the amphids. In this respect the present species resembles P. africanus (Gerl 


wl 
4 


MARINE NEMATODES FROM DURBAN 85 


1959) but differs from it in the arrow-head-like form of the amphidial flap. P. 
sudafricana is also, possibly, characterized by the number and form of the pre-cloacal 
supplements and the form of the spicular : gubernacular complex. 

Platycoma is very similar to Platycomopsis Ditlevsen, 1926 but this latter genus 
does not show any sexual dimorphism of the male amphids, and the male apparatus 
is more typical with long spicules and a short anterior part to the gubernaculum. 
This does, however, appear to represent a simpler version of the apparatus found in 
Platycoma and is one of the reasons for the interpretation I advance above. In 
addition Platycomopsis does not have the pair of spine-like setae on the male tail 
which is such a characteristic feature of the species referred to Platycoma. 


Trileptium longisetosum sp. nov. 
(Text-figs. 7-9) 


MATERIAL STUDIED. 3 4, I 9, 12 larvae. Beach sand in the surf zone. Near 
Durban, Republic of South Africa (B8n8). B.M. (N.H.) Reg. Nos. 1965.963-972. 


Body 

length 
A B € V (mm.) 

Males . : : F 10°64 5°63 22:69 — 4°68 
10°65 5:10 24°50 — 4°90 

Female é : ; 8-10 6°51 22°05 52°90 3°97 
Larva . : : : 8-54 5°69 24°12 — 4°10 


MEASUREMENTS (in mm. in order of body lengths) 

Mares: Body breadth: 0-044; 0:046. Oesophagus length: 0-83; 0-96. 
Diameter of head: 0-017; 0-018. Length of cephalic capsule: 0:036; 0-034. 
Diameter of cephalic capsule at posterior end: 0-019; 0-022. Length of longest 
cephalic setae: 0-040; 0:039. Length of cervical setae: 0-018-0-020 ; broken. 
Distance of nerve ring from anterior end of body: 0:19; 0-21. Length of spicules : 
0047; 0:052. Length of gubernaculum: 0-018; 0-016. Length of pre-cloacal 
supplement: 0:0065; 0:0065. Distance of pre-cloacal supplement anterior to 
Meacal opening: 0-038; o-040. Length of tail: 0-21; 0-20. Cloacal diameter : 
0:036 ; 0-040. 

FEMALE: Body breadth: 0-049. Ocesophagus length: 0-61. Diameter of head : 
0:020. Length of cephalic capsule : 0-022. Diameter of cephalic capsule at posterior 
end: 0-025. Length of longest cephalic setae: 0-066. Length of cervical setae: 
0-008. Distance of nerve ring from anterior end of body: o-16. Length of tail: 
0-18. Anal diameter: 0-036. Distance of vulva from anterior end of body: 2:1. 


LARVA: Body breadth: 0-048. Oesophagus length: 0-72. Diameter of head: 
0-020. Length of cephalic capsule: 0-023. Diameter of cephalic capsule at posterior 
end: 0-021. Length of longest cephalic setae : 0-042. Nocervicalsetae.. Distance 
of nerve ring from anterior end of body: o-1g. Length of tail: o-17. Anal dia- 
mieter : 0-043. 


ZOOL. 14, 4 788 


WILLIAM G. INGLIS 


i 


lics. 7-9. Trileptium longisetosum sp. nov. Fig. 7. Ventro-lateral view of head. Fig. $ 
Lateral view of male tail. Fig. 9. Spicule and gubernaculum from the left. Figs. 10-1: 
Mesacanthion frica sp. nov. Fig. 10. Dorsal view of head. The cephalic setae are n 
drawn in full because of their great length. Fig. 11. Lateral view of male tail. Fig. 1 


Detail of spicule and gubernaculum (?) from the left. a 


MARINE NEMATODES FROM DURBAN 87 


The structure of the head is the same as that in Tvileptvum ayum Inglis, 1964 with 
three small onchia enclosed by three small mandibles. The setae of the inner circle 
_ of cephalic sense organs are very stout and prominent and the pouch of the cephalic 
slit is rounded and very prominent (Text-fig. 7). The “cephalic setae’ are numerous 
and consist of six very long setae followed by two shorter setae on the dorso- and 
ventro-lateral positions and by one on the lateral positions. In the males there is 
one circle of ten cervical setae (Text-fig. 7) while there are four, much shorter setae 
in the equivalent position in the females. In addition there are a number of long, 
thin setae on the anterior end of the body which become sparser posterior to the 
nerve ring. Such setae measure 0:009—-0-013 mm. in length. 
- The tail is long and narrow in both sexes and a very small rod-like pre-cloacal 
supplement is present on the ventral surface of the male anterior to the cloacal 
opening (Text-fig. 8). 
_ The spicules are equal and identical with slightly cephalate proximal ends and 
bluntly rounded distal tips. The gubernaculum is simple and lies very close to the 
spicules (Text-fig. 9). 


) 
_ Discussion 

| This species is most similar to 7. ayum Inglis, 1964 but differs from it in the number 
and distribution of the cervical setae and the short spicules. It appears to differ 
most markedly from all the other known species in having three onchia and a pre- 
cloacal supplement in the male. 


| 
| 
| 


Mesacanthion frica sp. nov. 
| (Text-figs. 10-12) 


MATERIAL STUDIED. I ¢. Sand in surf zone. Near Durban. Republic of 


| 
‘South Africa (B8n8). B.M. (N.H.) Reg. No. 1965. xorr. 


| A ‘en © Body length (mm.) 
| 42°31 3°75 0-07 1°65 
| 


| MEASUREMENTS (in mm.) 

| Body breadth: 0-039. Oecsophagus length: 0-44. Length of cephalic setae 
_(anterior/longer posterior/shorter posterior) : 0:013/0:096/0-051. Depth of cephalic 
capsule (?): 0-026. Diameter of cephalic capsule at posterior edge: 0-026. Dis- 
| tance of nerve ring from anterior end of body: 0-139. Length of body setae about 
level of nerve ring: 0-016. Length of spicules: o-o40. Length of pre-cloacal 
| supplement : 0-012. Distance of pre-cloacal supplement anterior to cloacal opening : 
(0-078. Length of tail: o-182. Cloacal diameter: 0-036. 


The single specimen is in a rather poor condition so that some details cannot be 
teliably assessed. Nevertheless sufficient detail can be made out to warrant its 
} . . . . . ° . 

| description in view of the peculiar features it shows. The cephalic sense organs are 


' 


88 WILLIAM G. INGLIS 


very long and those of the posterior circle appear to lie about the posterior edge of 
the cephalic capsule. This is, however, uncertain and is suggested largely on the 
basis of the shape of the anterior end of the body (see Text-fig. to). The mandibles 
are prominent but fairly lightly developed and show no indication of any mandibular 
rods as in the species I refer elsewhere to Mesacanthion (Inglis, 1964). In addition, 
the shape of the anterior edge of the mandibles is different resembling the conditions 
in Africanthion in that their ends are directed towards the inter-labial spaces rather 
than inwards towards the mandibular cavity. The onchial plate is very poorly 
developed with the onchia projecting just anterior to the mandibular ring. } 

The tail is long and slim and a small, rod-like, pre-cloacal supplement is present 
(Text-fig. 11). The most remarkable feature of the specimen is the form of the 
spicules (Text-fig. 12), which possess a long narrow ventrally-posteriorly directed 
process on their anterior ends; are in addition stout with a lateral bump 
near their posterior ends and appear to have a median central ridge running their 
full length. There is (?) a small gubernaculum, although its presence is uncertain. 


Discussion 


This species is characterized by the form of the spicules with the massive process. 
on their anterior ends. Otherwise the spicules resemble those in Enoplolaimus 
crassidens Ditlevsen, 1930. 


Epacanthion oliffi sp. nov. 
(Text-figs. 13-16) 


MATERIAL STUDIED. 2 3,9. Beach sand in surf zone. Near Durban, Republic 
of South Africa (B8n8). B.M. (N.H.) Reg. Nos. 1965.1004—-roro0. 


Body 

length 
A B C V (mm.) 

Males . ‘ ; ° 34°81 3°19 10°33 — 1°88 
38-36 3°10 10°14 — 2°11 

Females , : ; 25°34 3°49 11°64 58-92 1°85 
32°07 2°88 12°10 61°73 1°96 

32°58 3°16 8-938 58°42 2°02 


MEASUREMENTS (in mm., in order of body lengths). | 
Mates: Body breadth: 0:054; 0°055. Oecesophagus length: 0-59; 0-6 
Diameter of cephalic capsule at posterior end: 0-034; 0-041. Depth of cephalic 
capsule : 0-033; 0-039. Length of cephalic setae (anterior/long posterior) : 0-015, 
0-036; 0-016/0-039. Length of cervical setae : in range 0:035—0-039 on both s 
mens. Distance of nerve ring from anterior end of body: 0-131; o-r40. Lengt 
of spicules: 0-028; 0-029. Length of tail: o-182; 0-208. Cloacal diameter 
0044 ; 0°046. Length of setae on tail and on general body surface : up to 0-039 


FEMALES: Body breadth: 0-073; 0-058; 0-062. Oesophagus length: 0°53 


MARINE NEMATODES FROM DURBAN 89 


0:68 ; 0:64. Diameter of cephalic capsule at posterior end: 0-038; 0-039; 0-038. 
Depth of cephalic capsule: 0-039; 0-041; 0-042. Length of cephalic setae 
(anterior/long posterior/short posterior): 0-017/0-034/0-015 ; 0-016/0-036/0-016 ; 
0:017/0-033/0-015. Length of cervical setae: 0°032; 0°028; 0°035. Distance of 
nerve ring from anterior end of body: 0-162; 0-138; 0-165. Length of tail: 
0:159; 0-162; 0-225. Anal diameter: 0-048; 0-044; 0-052. Distance of vulva 
fom anterior end of body: 1-09; I-21 ; -1-18. 


The head is typically enoplid with three high, slim lips bearing distinct semi-lunar 
striations lying on an area bounded anteriorly by a curved line running posteriorly 
from the tips of the mandibles (Text-fig. 13). The major part of the remainder of 
the lips is also marked by striations which are much further apart. These conditions 
are exactly as described for Enoploides species (see Inglis, 1964, fig. 30). The 
mandibles are typical of the genus with what appear to be two parallel rods joined 
by a thin sheet of cuticle. As usual this is due to studying a curved solid structure in 
optical section. The anterior ends of the mandibles form well developed projecting 
hooks. The three equal onchia are rather small and lie posterior to the posterior 
edge of the mandibles and do not lie close within them as in Hyalacanthion multi- 
papillatum Wieser, 1959. 

The cephalic setae are long and slim in both sexes with the more anterior pairs 
lying about the anterior end of the mandibles. Squarish blocks of specialized cuticle 
developed from the outer cuticle of head are closely associated with them (Text-fig. 
13). In the females the outer setae are represented by six long setae and four short 
but in the males there are groups of three dorso- and ventro-laterally and two pairs 
laterally. In the females there are four long cervical setae but in the male the region 
of the body posterior to the cephalic capsule carries a very large number of long 
setae (Text-fig. 15). Setae become scarcer on the body posterior to about the level 
of the nerve ring but long setae occur sporadically over the entire length of the body. 

The tail is somewhat blunt and stout in both sexes (Text-fig. 14). In the male there 
is no pre-cloacal supplement or gubernaculum. The spicules are short and stout 
with a series of serrations on.their posterior ends (Text-fig. 16). 


Discussion 


Wieser (1959) erects a new genus Hyalacanthion for three species, H. multipapilla- 
tum Wieser, 1959 (type species) ; H. pellucidus (Savaljev, 1912) and H. murmanicus 
(Savaljev, 1912). This genus, as he points out, is very similar to Epacanthion ; 
the major differences between the two being that the mandibles in Efacanthion 
consist of two plates, separated by a lacuna, which are joined anteriorly by a narrow 
rod and the spicules are at least 2-5 anal diameters long while the mandibles in 
Hyalacanthion are plates joined by a thin transparent lamella and the long spicules 
are never less than 2:5 anal diameters long. 

However Mawson (1958a) describes a new species, Epacanthion brevispiculosum, 
in which the spicules are considerably less than 2:5 anal diameters in length, and the 
species described above also has very short spicules, Ido not think that Epacanthion 


WILLIAM G. INGLIS 


Fics. 13-16. Epacanthion oliffi sp. nov. Fig. 13. Dorsal view of female head. 
Lateral view of male tail. Fig. 15. Dorsal view of anterior end of male body. 


Detail of spicule from the left. 


MARINE NEMATODES FROM DURBAN QI 


can be separated from Hyalacanthion but the question is simplified by a study of the 
type male of Enoplus butschlit Southern, 1914 which Wieser (1953) selects as type 
species of Epacanthion. The structure of the lips and mandibles of this species is 
exactly the same as that described above for E. oliffi. The same semi-lunar striations 
are present on the lips, the same masses of dense cuticle on the outer surface of the 
lips, the onchia are in the same positions relative to the mandibles (that is, posterior 
to them) and the mandibles have exactly the same appearance. There is no indica- 
tion of a distinct bar joining the so-called lateral bars of the mandibles as is illustrated 
by Wieser (1953) and by Southern (1914). 

As a result Hyalacanthion must fall as a synonym of Epacanthion and I can see 
no reason to recognize two genera for the species previously grouped in the two 
genera. I am quite sure that the differences reported in the structure of the man- 
dibles simply reflect the condition of the specimens when they were studied. But 
if later careful study shows a need for two genera a new name will be required. 
Hyalacanthion multipapillatum Wieser, 1959 is, therefore referred to the genus 
Epacanthion as a new combination. 

E.. oliffi is distinct from all the species with short spicules in lacking a gubernaculum 
and a pre-cloacal supplement and in the presence of such a large number of cervical 
setae. The species most similar to it appears to be E. multipapillatum (Wieser, 1959) 
in which the spicules are short and serrated but this species possesses a gubernaculum 
and a series of pre-cloacal supplements. 


Rhandodemania dura sp. nov. 
(Text-figs. 23-25) 


MATERIAL STUDIED. II 4, 6 9, r larva. Polluted sediments in the mouth of 
Durban Harbour, Republic of South Africa (M23n4, collected 20.vi.1964). B.M. 
(N.H.) Reg. Nos. 1965 .994-1003. 


Body 

length 

A B C Vv (mm.) 

Males . : , 32°02 7*O1 24°38 —- Cry 
30°38 6-92 20°77 —- 3°22 

33°57 6°97 24°19 a, 3°29 

36-22 6-93 22°90 5 3555 

Females : ; : 34°32 6-37 222 Gh 54°30 3°02 
34°40 (ae B. 26°26 57° 50 3°44 

32°84 7°09 23°87 53°63 3258 

40°32 7°01 23°01 54°67 3°43 

Larva . : : 35°00 4°93 15°98 — 1°47 


MEASUREMENTS (in mm., in order of body lengths) 

Mates: Body breadth: 0-099; 0-106; 0-098; 0-098. Ocesophagus length : 
0°452; 0-465; 0-472; 0:512. Diameter of head: 0-016; 0-018; 0-017; 0-018. 
Length of cephalic setae (long/short) : 0-0084/0-0063 ; 0:0083/0:0069 ; 0-0085/ 


92 WILLIAM G. INGLIS 


0-0064 ; 0-0087/0-0065. Depth of buccal cavity: 0-020; 0-017; 0-019; 0-018, 
Length of spicules: 0°055; 0°045; 0-052; 0-048. Length of gubernaculum: 
0-029; 0-026; 0-028; 0-031. Length of tail: 0-130; 0-155; 0-136; 0155, 
Cloacal diameter : 0-058 ; 0:058 ; 0°:057; 0-056. 

FEMALES: Body breadth: 0-088; 0-100; 0-109; 0-093. Oesophagus length : 
0°474; 0°458; 0505; 0535. Diameter of head: 0-018; 0-016; 0-017; 0-org. 
Length of cephalic setae (long/short) : 0-0082/0-0063 ; 0:0085/0-0062 ; 0-0086/ 
0-006T ; 0-0084/0-0065. Depth of buccal cavity: 0-021; 0:022; 0022; 0020. 
Length of tail: 0-133; O-r31; 0-150; 0-163. Anal diameter: 0-057; 0-066; 
0-065 ; 0°054. Distance of vulva from anterior end of body: 1-64; 1-98; 1-92; 
2:05. Size of eggs: 0-045 X 0°165; 0°04I X 0-142; 0:054 X 0-123 (as examples). 

LarvA: Body breadth: 0-042. Oesophagus length: 0-298. Diameter of © 
head: o-o12. Length of cephalic setae (long/short) : 0-0064/0-0037. 5 ae of 
buccal cavity: 0-016. Length of tail: 0-092. Anal diameter: 0-028. 


The species is typical of the genus with the usual three pairs of long tooth-like 
structures which represent modified mandibles (see Inglis, 1964) and with the lip 
region of the head marked by striations (Text-fig. 23). There is a fairly prominen 
dorsal onchium and no ventro-lateral onchia. 

The tail is bluntly rounded with a series of long stout setae, in the male (Text-fig.. 
25). g 

The spicules are slim with prominent caps on their proximal ends and are doubled 
at their distal ends (Text-fig. 24). They are roughly the same width along the whole 
of their lengths when viewed from the lateral aspect but have a ridge developed on 
their inner surfaces which can only be easily seen when the specimen being studied is 
viewed from a slightly ventral aspect. The gubernaculum is relatively long and 
slim with a very slight expansion towards the proximal end (Text-fig. 24). 


Discussion 


This species is characterized by the presence of only a dorsal onchium, by the slim 
shape of the spicules and gubernaculum, and by the cephalic setae being arranged in 
one circle only. In many ways it is similar to R. nancyae Inglis, 1964 which is als 
from South African waters but the shape of the spicules and gubernaculum is totally 
different while ventro-lateral onchia are totally lacking, although very reduced in 
R. nancyae. 


Family IRONIDAE 


The next species to be described differs from all others referred to the family in so 
many characters as to warrant reference to a new genus, thus : 


PHERONOUS gen. nov. 


lronidae : cephalic sense organs all papillose ; two dorsal onchia and one on € 
ventro-lateral sector of the oesophagus ; no caudal glands. 
Type species : Pheronous ogdeni sp. nov. 


MARINE NEMATODES FROM DURBAN 93 
PHERONOUS OGDENTI gen. et. sp. nov. 
(Text-figs. 17-22) 


MATERIAL STUDIED. 12 4, 16 9, 4 larvae. Beach sand in the surf zone. Near 
Durban, Republic of South Africa (B8n8). B.M. (N.H.) Reg. Nos. 1965.974-993. 


Body 

length 

fe B C V (mm.) 

Males . ; : ' 24°03 5°30 22°24 — 1°49 
34°71 593 26°03 a wi 

33°58 5°39 22-82 a 1°76 

38°83 5°52 25°38 = 1-98 

Females : : : 27*50 5°34 14°59 52°45 148 
30°85 4°98 14°36 57°93 eA 

28-81 6:06 13°94 56°58 Ta52 

e245 6°43 16-02 54°70 L773 

34°04 6°48 ty 3 5307 B77 


MEASUREMENTS (in mm., in order of body lengths) 

Mares: Body breadth: 0-062; 0:051; 0:053; 0:051. Odcsophagus length : 
osieeO 200; 0:330; 0-317. Diameter of head: 0-031: 0-023; 0-020; 0-023. 
Length of pharyngeal rods: 0-056; 0:062; 0:058; 0:057. Distance of nerve ring 
from anterior end of body: 0-128; 0-147; 0-140; 0-168. Length of spicules: 
0047; 0-041; 0:039; 0:035. Length of gubernaculum: 0-016; 0-019; 0-018 ; 
meen Leneth of tail: 0-033; 0:032; 0:033; 0:033. Cloacal diameter’: 0-067 ; 
B-00G.;, 0-070 ; 0-078. 

FEMALES: Body breadth: 0-052; 0:047; 0°054; 0°:055; 0:052. Oesophagus 
Bee. §0°208 5 0-291 ; 0°251 ; 0:269; 0-273. Diameter of head: 0-023; 0-019; 
0:027 ; 0°025; 0:024. Length of pharyngeal rods: 0-055; 0:059; 0:057; 0:054 ; 
0-056. Distance of nerve ring from anterior end of body: 0-121; 0-140; 0-123 ; 
Be, 0124. Length of tail: 0-098; 0-101; 0-109; 0°093; 0-123. Anal dia- 
tet 0-020 ; 0:0290; 0-032 ; 0-033; 0-028. Distance of vulva from anterior end 
mmpody : 0°75; 0°84; 0-86; 0°93; 0°95. 


The head is distinctly set-off from the remainder of the body by a constriction 
about the level at which the prominent oval shaped amphids open by narrow slits 
(Text-figs. r7 and 18). The mouth opening is bounded by three lip-lobes and is 
surrounded by six small papillae. The outer cephalic sense organs form one circle 
of stout papillae of which six are more prominent than the remaining four (Text-figs. 
17,18 andig). The buccal cavity is lined by a large number of small wholly cuticular 
denticles and the anterior end of the oesophagus bears a series of wholly cuticular 
-onchia. There is a pair of small onchia on the dorsal sector of the oesophagus and a 
single larger onchium on each ventro-lateral sector (Text-fig. 20). The anterior 
end of the oesophagus is modified as a distinct oesophastomal region in which the 


WILLIAM G. INGLIS 


22 


a\ 


_ — 


Fics. 17-22. PHERONOUS ogdeni gen. et sp. nov. Fig. 17. Lateral view of head and 
oesophastomal region of body. Fig. 18. Dorsal view of head. Note double onchia. Fig. 19. 
En face view of head. Fig. 20. Sketch of onchia en face. Fig. 21. Detail of spicule and 
gubernaculum from the right. Fig. 22. Lateral view of male tail. , 

Fics. 23-25. Rhabdodemania dura sp. nov. Fig. 23. Lateral view of head with the dorsal 
surface to the right. Fig. 24. Detail of spicule and gubernaculum from the right. Fig. 25. 
Lateral view of male tail. — 


MARINE NEMATODES FROM DURBAN 95 


musculature runs slightly anteriorly (Text-fig. 17) and in which the cuticular lining is 
very prominent. The slim oesophagus expands evenly towards its posterior end. 
The tail tapers to a sharp point and there are no caudal glands. In the male the 
tail bears a series of three pairs of small ventro-lateral papillae and a series of four 
pairs of similar papillae anterior to the cloacal opening (Text-fig. 22). The spicules 
are stout and clumsy in appearance without any particularly characteristic features 
although a dorsal strengthened rib sometimes appears to be present. The guber- 
naculum is small and enfolds the distal ends of the spicules (Text-fig. 2r). 


Discussion 


This species is unusual in lacking caudal glands, otherwise it would be referable 
to the genus Dolicholaimus. It is possible that it also differs from the species gener- 
ally referred to that genus in possessing a pair of small dorsal onchia but this is still 
uncertain (see Inglis, 1961). 


Family ONCHOLAIMIDAE 
Metoncholaimus murphyi sp. nov. 
(Text-figs. 26-32) 


MATERIAL STUDIED. 3,19. Beach sand in surf zone. Near Durban, Republic 
of South Africa (B8n8). B.M. (N.H.), Reg. Nos. 1965. 1or2—ror5. 3 


Body 

length 

A B C V (mm.) 

Males . : : ; 61°05 Loe 62-71 = 2322 
64°36 5°86 66°05 -— 2°51 

61°05 5+ 3a 62°71 — 229 

Female. : : d 52°31 5°55 59°13 WO 22 2-2 


MEASUREMENTS (in mm., in order of body lengths) 

Mares: Body breadth: 0-038; 0:039; 0:038. Oecsophagus length: 0-437 ; 
0-428 ; 0-419. Diameter of head: 0-027; 0-024; 0-026. Length of cephalic setae 
(long/short) : 0-012/0-008 ; o-orr/o-009; 0-012/0-008. Buccal cavity (greatest 
length/greatest width): 0-035/0-:017; 0:032/0-018 ;  0:035/o0-019. Distance of 
nerve ring from anterior end of body: o-r8r; 0-179; 0-198. Length of spicules : 
0:082 ; 0-069; 0-077. Length of tail: 0-037; 0-038; 0-043. Cloacal diameter : 
0029 ; 0-022; 0:026. Lengths of circum-cloacal setae (p/at/a2/a3): 0°012/0-017/ 
0:013/0-007 ; 0:013/0:018/0-016/0-008 ;_ 0:016/0-021/0-017/0-009 (see Text-fig. 29 for 
details). 

FEMALE: Body breadth: 0-052. Oesophagus length: 0-490. Diameter of head : 
0036. Length of cephalic setae (long/short) : 0-0r2/0-009. Buccal cavity (greatest 
length/greatest width) : 0-044/0-023. Distance of nerve ring from anterior end of 
body: 0-22. Distance of excretory pore from anterior end of body : 0-026. Length 


96 WILLIAM G. INGLIS 


of tail: o-o46. Anal diameter: 0-036. Distance of vulva from anterior end of 
body : 1-91. Distance of demanian pore anterior to posterior end of body: 0-072. 
Distance of uvette anterior to posterior end of body: 0:26. Distance of osmosium 

anterior to uvette: 0°36. Length of uterine efferent: 0-66. 


The head is typical with a fairly long buccal cavity with roughly parallel sides 
(Text-figs. 26, 27 and 28). There is an inner circle of six papillae and an outer circle 
of cephalic setae of which six are slightly longer than the remaining four. The left 
ventro-lateral onchium is considerably larger than the smaller and equal dorsal and 
right ventro-lateral onchia, both of which have somewhat blunt, slightly doubled 
anterior ends (Text-figs. 26 and 27). The amphids are prominent with slightly — 
different degrees of development of the anterior concavity (Text-figs. 26, 27 and 31). 


Fics. 26-28. Metoncholaimus murphyi sp. nov., head. Fig. 26. Left-ventro-lateral view. 
Fig. 27. Wholly lateral view. Fig. 28. Wholly dorsal view. 


The male tail is short and sharply curved, almost hook-like, with a pair of long 
stout post-cloacal setae (p in Text-fig. 29), and two similar pairs flanking the cloac 
opening, of which the more posterior is longer than the more anterior (ar, a2 in Text- 
fig. 29). In addition there is a further pair of shorter setae (a3 in Text-fig. 29) 
which appear to be slightly longer than the setae occurring on the general body 
surface except that, while these latter body setae vary somewhat in position fro 
specimen to specimen, this pair (a3) is always found in the same position. 

The equal and identical spicules are long and slim and a small gubernaculum may 
be present (Text-fig. 29) but this is uncertain. There is a marked thickening of the 
cuticle to form a bump on the body, just anterior to the cloacal opening, into which 
duct appears to open. The structure is definitely present but its details could n 
be established with certainty (Text-figs. 29 and 32). 


MARINE NEMATODES FROM DURBAN 97 


The female tail is short and stout (Text-fig. 30) and the demanian system empties 
through a single pore which opens slightly to the left of the mid-ventral line of the 
body. The details of the demanian system could not be wholly established but 
(using the terminology of Cobb, 1930) the wvette is surrounded by leaf-like elements, 
the uterine efferent is long, stout and easily seen and the osmosium could be identified 
0-364 mm. anterior to the uwvette. The relative measurements of the various parts 
that could be identified are given above. 


Discussion 


This is a typical member of the genus Metoncholaimus which can be easily distin- 
guished by the presence of only one demanian pore in the female (otherwise known 
only from M. haplotretos Mawson, 19584), by the extreme length of the spicules in 
association with the form of the male tail, by the small number of long circum- 
cloacal setae, and by the length of the cephalic setae. 


WIESONCHOLAIMUS Gen. Nov. 


The specimens described below resemble most closely those referred to the genus 
Metaparoncholaimus De Coninck & Schuurmans Stekhoven, 1933 in the unpaired 
female reproductive system, the two long, equal ventro-lateral onchia and the presence 
of a demanian system. It differs markedly, however, in the presence of the large 
gubernaculum, the form of the tail and the long, slim spicules. It is, therefore, 
referred to a new genus, thus: 


WIESONCHOLAIMUS gen. nov. 


Oncholaimidae : ventro-lateral onchia equal and larger than dorsal ; tail long and 
stout. MALE: spicules very long and slim ; large gubernaculum present. FEMALE: 
Demanian system present. 

TYPE SPECIES: Wzaesoncholaimus mawsonae sp. nov. 


WIESONCHOLAIMUS MAWSONAE gen. et sp. nov. 
¢ (Text-figs. 33-35) 


MATERIAL STUDIED. 6 4, 12 9. Polluted sediments in the mouth of Durban 
Harbour, Republic of South Africa (M23nir). B.M. (N.H.) Reg. Nos. 1965. 1o16— 
1030. (Holotype selected from this sample.) 

24,11 9, 8larvae. From same locality on same date at a depth of 40 feet (M23n4). 
B.M. (N.H.) Reg. Nos. 1965. 1031-1040. 


98 WILLIAM G, INGLIS 


Body 

length 
A B Cc V (mm.) 

Males . : . t . gO°r6 12°14 44°02 -—- 4°49 
64°93 I1*29 41°22 —_ 4°74 

70°27 12-09 40°94 = 5°20 

Females . : d 51°41 9°85 31°96 67°75 4°73 
54°95 10°87 33°84 64°38 5°11 

54°76 13°76 34°81 64°72 5°04 


MEASUREMENTS (in mm., in order of body lengths. All ex M23n11) 

MALES: Body breadth: 0-064; 0:073; 0-074. Oesophagus length: 0-37; 
0°42; 0-43. Diameter of head: 0-033; 0-030; 0-030. Length of cephalic setae 
(long/short) : 0:0090/0-0051 ; 0-0088/0-0053 ; 0:0091/0-0052. Dimensions of buccal 
cavity (length/breadth) : 0-032/0-018; 0-038/0-:016; 0-033/0-018. Distance of 
excretory pore from anterior end of body: 0-095; 0-103; 0-096. Length of 
spicules: 0-189; 0-211; 0-183. Length of tail: o-102; 0-115; 0-127. Cloacal 
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 ; 0:0092/0:005r ; 0:0087/0-0050. Dimensions of buccal 
cavity (length/breadth) : 0-033/0-020; 0-038/0-:020; 0:035/o-019. Distance of 
excretory pore from anterior end of body: 0-089; 0-121; 0-107. Length of tail: 
o-r48; O-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 xX 0-0067 ; 
0°097 * 0°078; 0-070 & 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 lie 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 
slightly 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). In addition 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. lo 
(from sample M23n4). A most characteristic feature of the system is that the wvelle, 


MARINE NEMATODES FROM DURBAN 99 


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. 


which lies 0-52 mm. anterior to the anus, is a simple spherical organ from which a_ 
stout wlerine 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 LEP TOLAIMIDAE 
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, 1g14. 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 Chetlorhabditis 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: Plectolaimus juliani sp. nov. 


PLECTOLAIMUS JULIANI gen. et sp. nov. 
(Text-figs. 36-43) 


MATERIAL STUDIED. 1 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°59 20°21 31°76 2°89 


MEASUREMENTS (in mm.) 

Body breadth: 0-052. Oesophagus length: 0-143. Body breadth at level o 
amphids: o-or7. Diameter of amphids: 0-0065. Length of cephalic setae: 
0-013. Distance of excretory pore from anterior end of body: 0-058. Length o 
spicules: 0-062. Length of gubernaculum: o-or8. More posterior pre-clo 
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. 


MAKINE 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 lies anterior to the corresponding structure on the 
right ventro-lateral sector. In addition both project dorsally to he in a 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 
slight 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. 309). 


DISCUSSION. 
See above, p. 100, 


WILLIAM G. INGLIS 


women 


yy oped 


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, | 
| | fe Wife x 
ayia a 39 | 
| \ 37 . 


38 


© ' * 


Pics. 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 en 
of oesophagus from the ventral aspect. Fig. 41. Detail of anterior pre-cloacal supplemen 
Fig. 42. Detail of posterior pre-cloacal supplement. Fig. 43. Lateral view of male tail, 


MARINE NEMATODES FROM DURBAN 103 
Family TRIPYLOIDIDAE 


Bathylaimus deconincki sp. nov. 
(Text-figs. 44-47) 


MATERIAL STUDIED. I 3. Beach sand in surf zone. Near Durban, Republic of 
South Africa (B8n8). B.M. (N.H.) Reg. No. 1965.994. 


A B C Body length (mm.) 
57°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 cephalic setae: 0-030. Diameter of head at level of amphids: 0-034. 
Lateral diameter of amphid: o-o10. Antero-posterior diameter of amphid: 0-009. 
Length of more anterior cephalic setae: 0-014. Length of more posterior cephalic 
setae: 0:038. Depth of buccal cavity from anterior end to level of ventro-lateral 
onchia: 0-030. Depth of second part of buccal cavity : 0-029. Length of spicules : 


ts 


Fics. 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. 


44 


104 WILLIAM G. INGLIS 


0-030. Length of gubernaculum : 0-036. Length of tail: 0-139. Cloacal diameter: — 


O-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 slight 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 1s 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 


| 
i 
| 
| 


MARINE NEMATODES FROM DURBAN 105 


species it is very different from the condition in B. deconinck1, 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 Tvaité de zoologie. Anatomie, 
systématique, biologique. Embranchement des Némathelminthes (Nematelmia Carl Vogt 
1851—Nemathelminthia Gegenbaur 1859) ou Aschelminthes (Aschelmintha Grobben 1910). 
4 (2) : 586-681. 

DE ConrINcK, 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, HJALMAR. 1926. Freeliving nematodes. Dan. Ingolf-Exped. 4 (6) : 1-42. 

1930. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. LI. 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. Meeresforsch. 

18 : 81-108. 

INGLIS, 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. Bujdr. 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. Sev. B. © (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. Leningr. 43 : 108-126. 

SOUTHERN, R. 1914. Clare Island Survey. Part 54. Nemathelmia, Kinorhyncha and 
Chaetognatha. Proc. R. Ivish Acad. 31: 1-80. 


Timm, Richard W. 1959. Cheilorhabditis and Odontorhabditis, 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. 1 
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. a 


"STUDIES ON THE BRITISH 
--—s DERMANYSSIDAE 
(ACARI : MESOSTIGMATA) 
PART II CLASSIFICATION 


G. OWEN EVANS 
and 


Ur Mee TIL 


BULLETIN OF 


| | | oe 14 No. 5 
LONDON: 1966 


STUDIES ON THE BRITISH DERMANYSSIDAE 
(ACARI : MESOSTIGMATA) 
PART II CLASSIFICATION 


BY 


G. OWEN EVANS 
AND 


WM. TILE 
British Museum (Natural History) 


Pp. 107-370 ; 101 Text-figures 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 14 No. 5 
LONDON: 1966 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, 1s 
tssued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. 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. 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 


Sees ON THE BRITISH DERMANYSSIDAE 


(ACARI : MESOSTIGMATA) 
PARA ToC LASSIFICA TION 


By G. OWEN EVANS & W. M. TILL 


CONTENTS 
Page 
SYNOPSIS : ; ; F - 4 ; , : : 5 i) TOO 
INTRODUCTION ‘ : 5 : : s {LOG 
CLASSIFICATION OF THE Britis DERMANYSSIDAE : F : : eerie 
KEY TO SUBFAMILIES. : F : : : ‘ 2 Sesto? 
Subfamily LAELAPINAE . . ; : , : 4 eel 24 
Subfamily HAEMOGAMASINAE . ‘ ; : : . $1238 
Subfamily PSEUDOLAELAPINAE , , : : : 265 
Subfamily MyonyssINAE . : ; : : : 1208 
Subfamily MELITTIPHINAE : : : : : : 27s 
Subfamily H1RSTIONYSSINAE . : : : A ‘ -. 276 
Subfamily MacRONYSSINAE : c ; A ; : . 300 
Subfamily DERMANYSSINAE ‘ : : ; : 3 - 346 
SPECIES INCERTAE SEDIS : d : : f : . n 362 
SUMMARY OF NEW TAXA AND Seco 3 F 5 : ‘ 1362 
ACKNOWLEDGEMENTS . : : ’ 5 , : A ‘ ne6s 
REFERENCES. : , ‘ ; : ‘ ; ‘ ) 304 
ABBREVIATIONS USED IN TEXT AND FIGURES : : : : =, 305 
INDEX TO GENERA AND SPECIES : . : : : : : 2 ‘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, Leiognathus 
Canestrini and Dermanyssus Dugés while the free-living forms, paraphages of 
arthropods and facultative ectoparasites, comprising the genera Laelaps Koch, 
Hypoaspis Can., Seius Koch, Iphis Koch, Stylochirus Can., Neoberlesia Berl., 
Podocinum Berl. and Iphiopsis Berl., were placed in the family Laelapidae Berlese. 
In his last comprehensive classification of the Mesostigmata published in Acarotheca 
Italica in 1913, Berlese retained his original concepts of the families although, by this 
ZOOL. 14, 5. 8 


110 G. OWEN EVANS @& W. M. TILL 


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 (Phytoseiinae 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, Rhinony 
sinae, Entonyssinae and Halarachninae. To these must be added Ixodorhynchinae 
Ewing (1923), Myonyssinae Bregetova (1956), Histrichonyssinae Keegan et 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 


LHe BRITISH DERMANYSSIDAE (ACART) 2 ie 


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? 
iserlese, 1892. 


SlLASsSIPIGATION OF vI RE BRUEISH. DERMANYSSLDAE 


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 
a non-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 

1 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 G. OWEN EVANS & W. M. TILL 


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 (j1-76, z1—z6, s1—s6, y2-75) occur in 
the podonotal region and 17 pairs (J/1—J5, Z1-Z5, St-S5 and px2-—p3) in the opisthonotal region, 
and 22 pairs of pores. Lateral unsclerotized integument with y + R 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. 


Larva; 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 (0-4-5-12- II); 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-like ; pedipalpal 
chaetotaxy (1-4—5-12-15). Tectum with weakly denticulate anterior margin. 

Podonotal region with 16 pairs of setae, 11 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 ; geni 
setae flanking the shield in the region of coxae IV. Metapodal shields subcircular, sm 
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’ Gaeolaelap: 
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 it: 


THRE BRITISH DERMANYSSIDAE (ACART) Ts 


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 Pneumolaelaps 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 Hypoaspfis 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 Hemilaelaps, 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. OWEN EVANS & W. M. TILL 


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, Eulaelapbs 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 intercoxal region. Hypertrichy 
is first apparent at the protonymphal stage and may be restricted to the opisthosoma, 
asin Eudlaelaps, or also affect the podonotum, asin Haemogamasus. The nature of the 
sclerotization of the idiosoma in the protonymph, deutonymph and adults of Eulaelaps 
and Haemogamasus conforms with the normal Hypoaspis type although there is 
increased sclerotization of the genital and metapodal shields in the female of Ewlaelaps. 
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 in form. 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 II 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 hav 
not been observed in the Laelapinae and Haemogamasinae. The main evolutio 
trends are towards decreased sclerotization of the dorsum and, to a lesser extent, th 


ae (bat Fish DERMANYSSTIDAE (\(ACART) I15 


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 Aszatolaelaps 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 Hemuilaelaps, 
Strandtrbbettsia and A siatolaelaps, but in Ixodorhynchus and Ixobioides 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 Asvzatolaelaps. 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 Stvandtibbetisia, Ixodorhynchus 
and Ixobio1des 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 completely] 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 & W. 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 (Berlesta), elongation and fusion of the spermadactyl with the movable digit 
of the chelicera in the male (J«lolaelaps), increase in number of transverse rows of 
deutosternal denticles (Myrmonyssus), reduction in the length of the peritreme and 
the enlargement of the stigmata (/phiopsis), 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- 
trae (Jellison, 1945), an ectoparasite of the primitive rodent Aplodontia. 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 14 
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 ro pairs (J1-5 and Z1-5) of which J4 — 
and /5 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 


Tae BRITISH DERMANYSSIDAE (ACARI) EY? 


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 
a The pertotany of the leg segments is normal except for tibia II 
(1—7, {—2), genu III (2—#, 2—1), tibia III (1—4, #—1), genu IV (1—2, 2-1) and 
tibia [IV (1—4, #—1). The deutonymph is an active feeding instar with the general 
characteristics 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. hopkinst, 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 analshield. 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 Laelafs group. The hypostome and labrum also 
resemble those of the Laelaps group with the corniculi 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 (RF. 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 Laelafs 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 


15 G. OWEN EVANS & W. M. TILL 


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 life-cycle comprises a non- 
feeding larva, a (?) feeding protonymph and an active, feeding deutonymph and 
adult. The genus Hirstionyssus 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 Bewstella 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- 


Lie spa Ttish DERMANYSSIDAE (ACART) 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 zI 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 Bewszella 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 second segment 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 in some 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 Ixodorhynchinae 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 pedipalpal chaetotaxy 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. 

i-ndoparasites 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 BRITiIisH DERMAN YSSIDAE (ACARE) De 


towards fusion of the tibia and tarsus and, in Pueuwmonyssus, 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 (Pueumonyssus) 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 Pueumonyssus simicola Banks appears 
to be non-feeding and of short duration. In Puneumonyssus 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 Pueumonyssus. 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 1s 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 & W. M. TILL 


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. M. 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. 


KEY 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 pu, modified into a short, tooth- 
like projection. Both sexes with Sams setae absent. Obligatory ecto- 
parasites of birds . : DERMANYSSINAE (p. 346) 

- Second segment of the chalcone’ in chal dma and female never conspicuously 
elongated, digits prominent. Seta pv, on the tarsi of the males never modified 
into a tooth-like projection . . ; a 

2 Dorsal shield in deutonymphal and adult papi AES PR he ha, setation 
often resembling a pelage. Genital shield, and sometimes sternal and anal shields, 
with unpaired accessory setae. Deutosternum with 10 or more transverse rows 
of denticles, majority of rows compound. Tectum with margins conspicuously 


LE BRIDISH DEKMANYSSIDAE (ACARI) 123 


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 . HAEMOGAMASINAE (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 genitalshield . ; ; ‘ ‘ ‘ ‘ ; : : 3 
3 Genu and tibia I with two ventral setae (z2—3, 3—2), genu II 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. d ; . PSEUDOLAELAPINAE (p. 265) 
- Genu and tibia I ene) sate vite sree sone (2—3, #—2),? genu II with two 
ventral setae (2—3, ?—-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 
coxa lI prominent. Associated with small mammals. ; . MYONYSSINAE (p. 268) 
— Without the above combination of characters. 5 
5 Corniculi horn-like, strongly or weakly sclerotized, never in ee iad of teyaine pee 
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 : ‘ : 5 : 6 
— Corniculi never horn-like, usually in the form of eee robes 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, oe aca, setae absent. Obligatory ecto- 
parasites of vertebrates... : : : : : Gi) 
6 Presternal area in the female with a sical: ie didge “Del@astenniain 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 
2 Ornithonyssus sylviarum 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 EVANS & W. M. TILL 


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 4 : : . LAELAPINAE (p. 124) 
7 Both sexes with one or more of the coxae with retrograde 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 ; : ; . i : é i 3 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 
(st2). | 

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 (st, 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 


Pe Bibi Ssk DE RIAN Y Sol DAE (ACA RT) 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 (x1), 
or a pair of anterior accessories (ax), or to suppression of posterior accessories (px 
series), 73, 23, J/3, J/4 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, 2—1), tibia III (2—}, #—2; 1—+4, }—1) and 
genu IV (2—2, 3—2; 2-2, 32; 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. TILL 


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 
(I-4-5-I2). 

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 10 or I1 pairs, pygidial shield with eight pairs. Trito- 
sternum as in 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. 


KEY 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 . . 4 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 : 3 
~ Genu IV with one postero-lateral seta (2—?, §—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 (p. 150) 
~ Female with genito-ventral shield RAPERE three pairs of hi alin setae in 
addition to the genitals. : 4 
4 Medio-dorsal series of setae hypotrichous, j3, pi 8 th lacking. Femur I with seta bd, 
arising from a strong, sclerotized Ful toeoet a Associated with Ondatra, 
introduced . : . . ONDATRALAELAPS (p. 145) 
Medio-dorsal series of setae normal, ‘comprising It pairs of setae (j1-76; J1-—J5). 
Seta pd, on femur I not arising from a strong protuberance ; 5 
5 Genu III withnine setae. Dorsal setae rarely short, Wi al px2 and (x3 present. 
Male with holoventral shield . : LAELAPS (p. 127) 


~- Genu III with eight setae. Dorsal setae short, subspinoee, never more than one pair 
of px setae. Male with sternito-genito-ventral and anal shields 
HYPERLAELAPS (p. 140) © 


Pie SSRIs DEKRMAN YSSIDAE (ACART) i277 


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 oe 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 AYPOASPIS s. lat. (p. 158) 


Genus LAELAPS Koch 


Laelaps Koch, 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 lilaris Koch, 1836. 


Medium sclerotized mites ranging from 500 « to 1,500 w# 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-lke, 
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 toit. 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 
Ly. 

Chaetotaxy of legs normal except for genu IV which has two postero-laterals 
(2—+, 2—z2) 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 


I Genu IV with two ventral setae (pv present) ; paranal setae short, their length equal 
approximately to width of anus (Text-fig. 1B) . : : L. agilis Koch (p. 132) 


128 G. OWEN EVANS & W. M. TILL 


— Genu IV with one ventral seta (pu absent) ; ei setae Dbiicsd aie their 
length at least twice the width of the anus __. 2 
Genito-ventral shield greatly expanded behind coxae IV, ‘with, posterior margin 
deeply concave ; distance between genital setae (gen.) less than that between /Jv2. 
Dorsal setae relatively long ; length of 75 distinctly greater than distance between 75 
and s5 (Text-fig. 3A). 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 /v2. Dorsal setae relatively short ; length of J 5 distinctly 
less than distance between 75 and s5. Pilus dentilis inflated a 3 
3 Sternal setae relatively long and slender, str reaching posterior margin of sternal 
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 st2. Dorsal setae stout and spine- 
like ; marginal opisthogastric setae simple ‘ ; L. muris (Ljungh) (p. 137) 


te 


KEY TO MALES 


1 Genu IV with two ventral setae (pu present) ; tarsus 1V 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 hale approximately equal to width of anus. 
Chelicera as in Text-fig. 2E_ . : : L. agilis Koch (p. 132) 
— Genu IV with one ventral seta (pu absent) ; eatwen IV with setae ad, and ad, setiform 
or blade-like but not exceeding 14 times the width of the tarsus. Peritrensiih 
longer, extending at least to anterior margin of coxa II. Paranal setae longer, 
their length at least 14 times the width oftheanus . 2 
2 Holoventral shield with five pairs of stout setae in genito- ventral region (Text- fig. sD). 
Tarsus IV with setae ad, and ad, blade-like ; seta pv, short, stout and spine-like 


(Text-fig. 5F). Chelicera as in Text-fig. 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- Mike 3 tarsus lV withsetae al, and pu, 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 pv, slender, 
pointed. Pilus dentilis setiform ; chelicera as in Text-fig. 3E 
L. echidnina Berlese (p. 136) 


Laelaps hilaris Koch 


Laelaps hilaris Koch, 1836, Deutschl. C.M.A.4:20. Tipton, 1960, Univ. Calif. Publ. Ent. 16: 
271, figs. 


Female : Chelicera with segment I, 54 »; II, 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 ; hyp. 2 about 54 » apart. Deutoster- 
num with six transverse rows of denticles (1-2 per row) ; corniculi 27 , long ; internal 


» Laelaps clethrionomydis Lange (see Bregetova, 1956) commonly occurs on the bank vole, Clethrionomys 
clareolus (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 pve short and spine-like and setae 
adg 9 and fde-s long and blade-like. These tarsal setae are simple, setiform in the male of L. hilaris. 


THE BRITISH DERMANYSSIDAE (ACARI) 


d venter 
of idiosoma. 


um (D) 


nter (c) and dors 


Laelaps agilis Koch, female ; dorsum (a) an 


IG. I. A-B. 
Laelaps hilaris Koch, female ; ve 


130 G. OWEN EVANS & W. M. TILL 


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 r 
and seven pairs of R setae on the cuticle, all with bifurcate tips or small barbs. 

Tritosternum with base 45 p», laciniae 102 ». Sternal shield (102 px 186 ») with 
three pairs of setae and two pairs of pores ; metasternal setae on small platelets, 
their associated pores on the cuticle. Between sf1, 80 » ; between sfI and st3, 100 p. 
Genital shield (144 » X 135 ») flask-shaped, with four pairs of setae. Between gen., 
114 «; between /v2, 80 pw. Anal shield (105 » X 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 pv, 
is slender. Coxal setae as in Text-fig. 1c. Length/width (in yp) of leg segments : 


I II III IV 
genu 66/57 72/60 50/45 80/48 
tibia 70/40 57/57 48/45 78/48 


tarsus 100/36 90/38 105/36 150/40 


Male : Chelicera with segment I, 48 »; II, 105 »; movable digit 50 p» (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 w 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 » * 456 ») with 39 pairs of setae distributed as in Text-fig. 2G. 
Surface of shield reticulated ; no hypertrichy ; two pairs of r and five pairs of R setae 
on the cuticle. 

Tritosternum with base 45 p, laciniae 108 ». MHoloventral shield (516 » & 144 p) 
with four pairs of setae and three pairs of pores in the sternal region ; six pairs of 
setae in the genito-ventral region. Between sf1, 30 »; between sti and st3, 102 p. 
Paranal setae 45 p, postanal seta 84 ». Peritreme extends to posterior margin of 
coxa | ; peritrematal shield fused anteriorly with dorsal shield ; posterior part free. 

Chaetotaxy of legs asin female. Tarsi II and III with setae al,, pl,, av,, and also 
av», relatively shorter and stouter than the corresponding setae in the female. 
Tarsus IV has two short, stout, blunt setae, al, and pv,. Coxal setae as in Text-fig. 
2H. Length/width (in ») of leg segments : 


] II III IV 
genu 78/66 84/72 63/57 102/66 
tibia 75/48 66/64 60/50 96/63 


tarsus 102/36 go/40 102/36 147/42 


Tie BRITISH DEKMANYSSIDAE (ACART) 131 


Hapitat: 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; 3B. Laelaps hilaris Koch, chelicera of 
male; c. L. agilis, tarsus IV of male; pb. L. hilaris, chelicera of female; k. L. agilts, 
chelicera of male; F. L. hilaris female, venter of gnathosoma ; G. L. hilaris male, dorsum 
of idiosoma ; H. L. hilaris male, venter of idiosoma. 


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 hilaroides OQudemans, 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 » apart; hyp. 2 about 45 » apart. Deuto- 
sternum with seven transverse rows of denticles (I-2 per row) ; corniculi 27 y» 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 « x 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 r and seven pairs of R setae. 

Tritosternum with base 57 p, laciniae 100 p. Sternal shield (123 » X 192 w) with 
three pairs of setae and two pairs of pores; metasternal setae situated on small 
shields, their associated pores on the cuticle. Between st1, 90 1»; between sti and 
st3, 114 ww. Genital shield (147 » x 168 ,) flask-shaped, with four pairs of setae. 
Between gen., 126 »; between Jv2,72 w. Anal shield (110 » X 144 ») with paranal 
setae 27 w, postanal seta 105 p. Chaetotaxy and sclerotization of opisthogaster as in 
Text-fig. 1p. 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 pd,); on tarsus II, setae al, and #l,, and on tarsus III, 
setae al,_, are stout ; on tarsus IV setae al, and pv, are stouter than setae p/, and a7. 
Length/width (in ) of leg segments : 


I II III IV 
genu 87/63 78/68 50/50 75/48 
tibia 72/54 60/60 45/48 72/42 
tarsus 90/40 84/42 114/42 153/36 


Male; Chelicera with segment I, 50 » ; II, 120 »; spermadactyl about 84 p long, 
separate movable digit not present (Text-fig. 2E). Four pairs of gnathosomal setae 
with c.s. about 48 apart ; hyp.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 » * 432 ») with 39 pairs of setae as in female. Tritosternum 
with base 42 u, laciniae 90 ». Holoventral shield (516 » * 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 st1, 72 »; between sf1 and sf3, 114 ». Paranal setae 18 p, 


THE BRITISH DERMANYSSIDAE (ACARI) 


A. Dorsum of idiosoma of female ; 
of female ; p. venter of idiosoma of male; 


F. tectum capituli of female; G. venter of gnathosoma of female. 


Fic. 3. Laelaps echidnina Berlese. 


soma of female; c. 


male ; 


chelicera 


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,, p/, 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 pd, very long and thick. Length/ 
width (in ,») of leg segments : 


I II Ill IV 
genu 80/70 80/66 63/57 87/60 ; 
tibia 66/56 60/63 50/34 80/54 ; 
tarsus 75/40 80/45 96/42 147/42 


HABITAT: Found on Afodemus 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. j 


Laelaps echidnina Berlese 


Laelaps (Iphis) echidninus Berlese, 1887, Acari, Myr. Scorp. Ital. fasc. 39, no. 1. 

Echinolaelaps echidninus : Strandtmann & Mitchell, 1963, Pacific Insects 5 : 547, fig. 1. 

Laelaps pallidus Tragardh, 1931, Nat. Hist. Juan Fernandez 3: 616, figs. 145-147. 

Laelaps berlesei Fonseca, 1939, Mem. Inst. Butantan 12: 104. 

Echinolaelaps echidninus vitzthumi Turk, 1950, Parasitology 40: 71, fig. 7. 

Echinolaelaps hirsti Turk, 1950 (nom. nov. pro L. echidninus Hirst, 1913, non Berlese, 1887), 
Parasitology 40 : 72, fig. 6. 

Echinolaelaps hirsti cevlonicus 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 »; movable digit 72 yp, bi- 
dentate ; fixed digit unidentate (Text-fig. 3c) ; pilus dentilis long and slender. Four 
pairs of gnathosomal setae with c.s. about 96 yw apart; Ayp.2 about 84 mw apart. 
Deutosternum with six transverse rows of denticles (2-3 per row) ; corniculi 54 
long; internal malae fringed (Text-fig. 3c). 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. 3A; 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 pw. Sternal shield (234 p 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 
stl, 132 «; between str and st3, 204 p. Genital shield (258 » x 216 yp) greatly 
expanded behind coxae IV (maximum width 396 ,), bearing four pairs of setae. 


¢ 


136 G. OWEN EVANS & W. M. TILL 


Between gen, 180 «x; between Jv2, 276 w. Anal shield (168 » xX 190 p), paranal 
setae 72 mw, 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 genu IV. Setae on tarsi II-IV thick and spine-like ; the following setae are 
stout and blunt: al, and pi, 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 1; 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 » X 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 ». 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 st1, 84 » ; between sti and st3, 150 ». Paranal setae 72 » ; postanal seta 
140 ». 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 : 


I II 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 y« part, hyp.2 about 70 pw 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 » * 372 ») with eleven pairs of setae ; pygidial shield 
(192 « & 350 m) with eight pairs of setae, the marginal ones being barbed. Chaeto-— 
taxy and sclerotization of mesonotal region as in Text-fig. 4A. 


THRE BRITISH DERMANYSSIDAE (ACARI) 137 


Tritosternum with base 60 yp, laciniae 135 p. Sternal shield (255 mw x 162 yp) 
with three pairs of setae and two pairs of pores ; between sf1, 96 » ; between sti and 
st3, 102 w. Anal shield (130 » X 123 ») with paranal setae 78 pw, 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 pz) of leg segments : 


I II Ii! IV 
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 w; II, 168 »; chelae as in female, 
movable digit 54 ». Four pairs of gnathosomal setae with c.s. about 78 yp 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 p, laciniae 150 pw. Sternal shield (432 » x 168 p) 
with four pairs of setae and three pairs of pores. Between sf1, 114 w; between sf1 
and s#3, 192 w. Anal shield (150 » X 156 p») with paranal setae 75 pu, postanal seta 
168 y. Peritreme extends to posterior third of coxa I (Text-fig. 4D). 

Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : 


I II Il 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 


HaBITAT: Cosmopolitan, chiefly on rodents of the genus Rattus. Also recorded 
from the genera Mus and Bandicota and from the tree-shrew, Tupaza 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 arvicola George, 1889, Science Gossip 25: 6, figs. I-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. 


Evmaie: Chelicera with segment I, 70 #7; II, 110 7; 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 


a. Dorsum of idiosoma of female; B. chelicera of mal 


Fic. 5. Laelaps murts (Ljungh). 
of idiosoma of male; £. pilus dentilis 


c. venter of idiosoma of female; D. venter 
female ; F. tarsus 1V of male, ventral view. 


THE BRITISH DERMANYSSIDAE)(ACART) 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. 5A. 
Most setae short and spine-like, setae 75 and S3-5 at least twice as long as seta /TI. 
Surface of shield granular, weakly reticulate laterally ; no hypertrichy ; two pairs of 
vy and ten pairs of F& setae on the cuticle. 

Tritosternum with base 60 p, laciniae 126 w. Sternal shield (144 w x 222 wu) with 
three pairs of short, spine-like setae and two pairs of pores ; metasternal setae on 
small shields, associated pores on cuticle. Between sti, 93 » ; between sfI and s?3, 
120 w. Genital shield (180 » x 174 p») flask-shaped with four pairs of setae ; between 
gen., 140 w; between Jv2, 84 u. Anal shield (96 » X I10 p») with paranal setae 63 p, 
postanal seta 168 w. 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 fl, being considerably shorter than #/, ; tarsi II-IV with setae ad, 
and #d,. Femur I with setae ad, and fd, stout and sinuous, the latter very long. 
Setae av, al, and #1, 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 II III IV 
genu 90/70 90/70 60/54 102/54 
tibia 75/58 66/63 50/50 90/50 
tarsus 96/45 90/48 = 114/42 153/42 


Male: Chelicera with segment I, 48 »; II, 87 »; both digits edentate ; movable 
digit about 70 p» 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 pw long ; 
salivary styli extend approximately to tips of corniculi. Deutosternum, internal 
malae, tectum and pedipalps as in female. 

Dorsal shield (636 » x 462 ») with 39 pairs of setae ; one pair of 7 and six pairs of 
R setae on the cuticle. Tritosternum with base 45 yp, laciniae 114 ». Holoventral 
shield (546 » X 180 ») with four pairs of setae and three pairs of pores in the sternal 
region ; five pairs of setae in the genito-ventral region. Between sti, 78 »; between 
ieand siz, 108 ». Paranal setae 36 p, postanal seta 130 wp. 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 asin female. In addition, seta av, on tarsus 
Il is stout and blunt. Tarsus IV with setae fv, and fv, 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 p) of leg segments : 


ZOOL. 14, 5. 10 


140 G. OWEN EVANS & W. M. TILL 


I II Ill IV 

genu 90/63 90/78 66/60 114/66 
tibia 75/54 66/66 57/57 99/63 
tarsus 93/44 84/45 105/45 140/45 


HasiTat: Found in Britain, Europe and the U.S.S.R., mainly on Arvicola 
terrestris (Linn.). Also recorded from Microtus 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 (fx2 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—#, 4—1), tibia III in H. amphibia 
(1—}, }—1) and genu IV (2—?, $—2). Femora I and II in both sexes with ad, at 
least three times the length of #d,. 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 (‘‘ Lemnus arvalis”), — 
believing it to be conspecific with Acarus pachypus Hermann, 1804. The first 


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 Microtus arvalis) being 
referable to Laelaps pachypus sensu Hirst, and the female (from Arvicola amphibius) 
to a species which has subsequently been described as Laelaps (Hyferlaelaps) 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 
koch, 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 kocht Oudemans. 

Recently we have had the opportunity, through Dr. Crabill, Smithsonian Institution, 
of examining the “ cotype ’”’ material of Tetvagonyssus microti Ewing from Mucrotus 
californicus, Los Angeles, California, and from Microtus sp., Alaska. We are unable 
to separate morphologically Ewing’s species from Laelaps kochi Oudemans taken froiu 
Microtus arvalts in Britain. We therefore consider L. kochi to be synonymous with 
T. microti. 


Kery TO MALES AND FEMALES 


1 Tibia III with chaetotactic formula (2—1, 2—1). Palpfemur with five setae. 


I dE 
Dorsal shield with setae px2 present or absent. In the female, seta s6 short 
and spinose, seta SI relatively long and slender : H. microti (Ewing) (p. 141) 


- Tibia III with chaetotactic formula (1—}, #—1). Palpfemur with four setae. 
Dorsal shield usually with setae pv¥2 present. Inthe female, setas6andS1 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) arvalis Zachvatkin, 1948, Parazit. Sb. 10: 74, figs. 25-26 (syn. nov.). 


Female: Chelicera with segment I, 54 »; II, 140 w; movable digit 36 p, biden- 
tate; fixed digit unidentate, pilus dentilis elongate, broad, not inflated distally 
(Text-fig. 78). Four pairs of gnathosomal setae with c.s. about 66 p 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 » x 444 ») 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. OWEN EVANS & W. M. TILL 


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. eae 

Tritosternum with base 45 », laciniae 110 ». Sternal shield (70 » x 192 p) with 
posterior margin deeply concave, lateral margins heavily sclerotized ; with three 


Fic. 6. A-B. Hyperlaelaps microti (Ewing), female ; dorsum (a) and venter (8) of idiosoma. 
c-D. Hyperlaelaps amphibia Zachvatkin, female ; opisthonotal region (c) ; sternal and 
genito-ventral shields (p). 


THE BRITISH 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 sti, 70 u; between sti and sf3, 117 p. 
Genital shield flask-shaped (135 » X 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%; between /v2, 50 ». Anal shield 
(70 » X 78 uw) with maximum width (87 uw) at anterior margin. Paranal setae 78 pn, 
postanal seta 108 » 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 fv ; genu IV has seta #/, ; tarsi II-IV lack setae ad, and fd,. Femur 
I with seta ad, at least three times as long as pd, ; femur II with seta ad, about three 
times as long as #d,._ 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 al 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 I UI 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 »; spermadactyl 150 » ; movable 
digit not visible (Text-fig. 7c). Four pairs of gnathosomal setae with c.s. about 57 yp 
apart ; hyp.2 about 42 w apart. Corniculi about 40 yp long, weakly sclerotized, 
membranous. Internal malae difficult to distinguish. Deutosternum, tectum and 
pedipalps as in female. 

Dorsal shield (504 » X 360 p) with 37 pairs of setae as in female ; seven pairs of R 
setae on the cuticle. | 

Tritosternum with base 36 p; laciniae 90 pw. Sternito-genito-ventral shield (324 pw 
x 150 ») with four pairs of setae and three pairs of pores in sternal region, six pairs 
of setae in genito-ventral region. Between si1, 54 »; between sti and sf3, 102 up. 
Anal shield (60 » x 70 ») with maximum width (80 ») at anterior margin. Paranal 
setae 48 », postanal seta 75 uw. 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 p) of leg segments : 


I I II 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 


Hasirat: Found in Britain on Microtus 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 (Koch) Oudemans, 1927, Tijdschr. Ent. 70: 179 (in part). 


Female ; Chelicera with segme1t I, 50 » ; II, 153 » ; movable digit 36 pu, bidentate ; 
fixed digit unidentate, pilus dentilis elongate, band-like, with pointed tip. Four 
pairs of gnathosomal setae with c.s. about 66 » apart, Ayp.2 about 54 pw apart. 
Deutosternum with six rows of denticles (1-2 per row) ; corniculi 30 »long. 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 St both short and spine-like. Surface of shield granular, reticulate 
laterally ; seven pairs of R setae on the cuticle. 

Tritosternum with base 42 yp, laciniae 117 pw. Sternal shield (84 » x 192 ») with 
setae 2 and 3 longer and more pointed than in H. microti (Text-fig. 6D) ; between sft, 


THE BRITISH DERMANYSSIDAE (ACART) 145 


78 u; between sfi and sf#3, 123 w. Metasternal setae on small platelets, associated 
pores on cuticle, very close to sternal shield. Genital shield (150 w X 162 wu) with 
maximum width (216 yp) at level of Zv1 ; between gen., 102 w; between Jv2, 60 up. 
Genital setae (gen.) more pointed than in H. microti (Text-fig. 6D). Anal shield 
(60 » X 78 p) with maximum width (102 ») at anterior margin. Paranal setae 60 pz, 
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 p») of leg segments : 


I II a0! IV 

genu 90/75 90/84 54/63 90/54 
tibia 66/60 60/72 40/54 84/45 
tarsus 72/45 93/48 87/45 150/42 


Male : Chelicera with segment I, 42 1; II, 120 w; spermadactyl 168 », movable 
digit not visible. Four pairs of gnathosomal setae with c.s. about 63 yp apart ; 
hyp.2 about 48 w apart Corniculi about 54 » long, weakly sclerotized. Internal 
malae difficult to distinguish. Deutosternum, tectum capituli and pedipalps as in 
female. 

Dorsal shield (552 » X 396 ») with 38 pairs of setae asin female. Tritosternum 
with base 33 p, laciniae 90 ». Sternito-genito-ventral shield (372 » x 168 p) as in 
H. microti. Between sti, 63 »; between sti and sf3, 102 p. Anal shield (63 p x 
75 ») with maximum width (90 ») at anterior margin. Paranal setae 45 p, postanal 
seta 90 » long. Chaetotaxy and sclerotization of opisthogaster as in H. mucrotu. 
Peritreme extends to middle of coxa II. 

Chaetotaxy of legs and form of setae as in female. Length/width (in yp) of leg 
segments : 


I II III 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 


Hasitat: 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: 2609. 


Type: Laelaps multispinosa Banks, rgo9. 


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 


146 


G. OWEN EVANS & W. M. TILL 


ais 
A 
"tt hy dhe ie 
’ q i -" 
a eee asi 


‘\ - 


rN 


Fic. 8. Ondatralaelaps multispinosa (Banks). A. Dorsum of idiosoma of female; B. 
venter of idiosoma of female; c. dorsum of idiosoma of male; b. venter of idiosoma of 


male. 


roHE- Beliseh 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 
per row. Corniculi horn-like ; internal malae fimbriated. Tectum capitul 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, /3, /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 
(c and A) 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—2, 
32). Coxa II with well-developed anterior spine. Femur I with ad, and fd, very 
stout and long, fd, arising from sclerotized protuberance (Text-fig. 9D). 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 evanst (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 spinigey Ewing & Stover, 1915, Ent. News 26 (3) : 109. 

Tetragonyssus spiniger: Ewing, 1922, Proc. U.S. nat. Mus. 62 (art. 13) : 10. 

Laelaps parvanalis Willmann, 1952, Z. Parasitenk. 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. 9£). Four pairs of gnathosomal setae with c.s. about 66 p apart, hyp.2 
about 60 w apart. Deutosternum with six transverse rows of denticles (one to three 
denticles per row) ; corniculi horn-like, 30 » long; internal malae fimbriate (Text- 
fig. oF). Tectum capituli 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 » 432 #) with 32 pairs of setae distributed as in Text-fig. 8A ; 
3, 23, /3, /4, 23, 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 w. Sternal shield (150 » x 207 p») with 
three pairs of setae and two pairs of pores. Metasternal setae on distinct shields. 
Between st1, 80 «%; between sf and st3, 120 pw. Genital shield (174 » X 126 p) 
flask-shaped, bearing four pairs of setae ; between gen., 105 «; between Jv2, 99 um. 
Anal shield (140 » X 93 p») elongate pear-shaped, with paranal setae situated 
posterior tothe anus. Paranals 18 » long, postanal seta 57 np long. Chaetotaxy and 
sclerotization of opisthogaster as in Text-fig. 8B. Peritreme extends to middle of 
coxa |. Peritrematal shield fused anteriorly with dorsal shield, posterior part free. 

Chaetotaxy of legs normal except genu III, which lacks seta pu (2—?, §—1) and 
genu IV, which has two postero-lateral setae (2—}, $—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 go/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. gc). Four pairs of gnathosomal setae with 
c.s. about 57 w apart ; Ayp.2 about 48 » apart. Corniculi 30 » long ; other gnatho- 
somal structures as in female. 

Dorsal shield (612 » * 360 ) 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 yu, laciniae 66 ». Holoventral shield (528 » x 168 p) 
with nine pairs of setae, excluding the anals. Between st1, 57 »; between sti and 
st3, 117 ». 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 II II IV 
genu 90/75 75/75 60/63 == 90/60 
tibia 72/60 54/60 50/54 88/54 


tarsus go/42 93/42 93/42 135/40 


: 
1 
: 
; 
bi 


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. TILL 


Deutonymph (male) : Chelicera with segment I, 36 » ; II, 80 »; chelae asin female, 
movable digit 27 ». Four pairs of gnathosomal setae with c.s. about 54 » apart ; 
hyp.2 about 45 » apart. Gnathosomal structures generally as in female. 

Dorsal shield (540 «# 228 ») with 16 pairs of setae in the podonotal region, 11-12 
pairs in the opisthonotal region (Text-fig. ga). 

Tritosternum with base 42 p, laciniae 100 »; Sternal shield (240 » X II7 p) 
with four pairs of setae and three pairs of pores. Between sf1, 60 » ; between sti and 
st3, 110 ». Anal shield (96 » x 66 pw); par- and postanal setae very short, about 
8 » and 10 » 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 III 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 p; 
movable digit 36 ». Dorsal shield (588 » x 240 p) ; sternal shield (252 » X 126 p) ; 
anal shield (96 » X 75 p). 


_ Hasrrat: 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 : 186. 
Ischnolaelaps Fonseca, 1936, Mem. Inst. Butantan S. Paulo 10: 19. 
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 » in length. Cheli- 
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 


THE BRITISH DERIMAN YSSIDAE (ACARI) 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-like, 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 J 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 z3 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 aseparate anal shield. Podal shields poorly developed posterior to 
coxae IV. 

Chaetotaxy of legs normal except genu IV which has two postero-laterals come 
*—2). Insome ethiopian species ek patterns are found on tibia I (2—3, 3—2), 
genu IIT (2—?, 22) and tibia III (2—+, 22) and in one species genu IV is normal. 
Seta av, on saaaie 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 


I Both sexes with pilus dentilis inflated basally, distal portion slender, curved (Text- 
fig. 12B). Greatest width of genital shield in female at the level of the first pair 
(Zvi) 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 approximately at level of second pair (Jv1) of opisthogastric 
setae (Text-fig. 108) : : ; : , A. casalis (Berlese) s. lat. 


2 Dorsal shield 620 uw-7I0 uw long in female, 466 fit 5 # long in male ; seta Z5 slender, 
setiform, about one and one-half times as long as J5 (Text-fig. 118), Associated 
with birds and rodents . : : ; ; ; : A. casalis s. str. (p. 152) 
— Dorsal shield 504 u long in female, 402 w long in male ; seta Z5 stout, subspiniform, 


scarcely longer than J5 (Text-fig. 11c). Associated with ants 
A. casalis myrmecophila ssp. nov. (p. 154) 


152 G. OWEN EVANS @& W. M. TILL 


Androlaelaps casalis casalis (Berlese) 


?Gamasus fenilis Mégnin, 1876', J. Anat. Physiol. Paris 12 : 332. 

I phis 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. 

Altricholaelaps 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. 1. 


Female : Chelicera with segment I, 63 » ; II, 108 » ; movable digit 36 », bidentate ; 
fixed digit with two or three teeth ; pilus dentilis slender, setiform (Text-fig. ropD). 
Four pairs of gnathosomal setae with c.s. about 60 p apart ; hyp.2 about 42 pw apart. 
Deutosternum with six transverse rows of denticles (2-6 per row). Corniculi 36 pw 
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 » x 376-429 p) with the normal 39 pairs of simple setae 
and a variable number of unpaired accessory setae distributed as in Text-fig. I0A. 
Surface of shield reticulate ; two pairs of ry and seven pairs of R setae on the cuticle. 

Tritosternum with base 54 p», laciniae 80 ». Sternal shield (100 » xX 126 p) with 
three pairs of setae and two pairs of pores; metasternal setae free with associated 
pores. Between sf1, 80 » ; between sti and st3,96 ». Genitalshield (177 » X 105 p) 
flask-shaped with one pair of setae. Anal shield (g0 » x 102 ») with par- and post- 
anal setae about 42 » long. Chaetotaxy and sclerotization of opisthogaster as in 
Text-fig. 10s. 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 genu lV. Length/width (in p) of leg segments : 


I 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, 40 »; II, 75 »; movable digit 40 p, 54 » with 
spermadactyl ; both digits edentate (Text-fig. 10£). Four pairs of gnathosomal 
setae with c.s. about 48 pw apart; Ayp.2 about 33 » apart. Corniculi 27 yp long ; 
deutosternum, internal malae, tectum and pedipalps as in female. 

Dorsal shield (466-485 » * 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 ». MHoloventral shield (384 » 95 p) — 


with four pairs of setae and three pairs of pores in the sternal region, six pairs of setae _ 


“It is not possible to be certain of the identity of this species from its short and inadequate original 
description. 


s 


) 


PE -SERITISH DEKMANYSSIDAE) (ACARI 


154 G. OWEN EVANS @& W. M. TILL 


in the genito-ventral region. Between sti, 60 » ; between sti and st3, 80 ». Paranal 
setae 18 «, 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 p) of leg segments : 


I II 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 


Hasitrat : Cosmopolitan, in the nests of birds, in poultry litter and also on rodents. 


Androlaelaps casalis myrmecophila ssp. nov. 


In all instars 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. I1C). 

Female: Dorsal shield (504 » * 288 ,») with the usual 39 pairs of setae and two to 
three unpaired accessory setae distributed as in Text-fig. 11a. Sternal shield (80 p x 
105 ) not sharply demarkated from pre-sternal area. Between st1, 72 1; between 
str and st3, 78 pw. Genital shield (140 » x 84 w) with genital setae only. Anal 
shield (72 » & 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 II Ill IV 
genu 70/32 54/36 42/27 63/27 
tibia 75/27 50/36 45/27 66/27 
tarsus 130/26 84/24 go/22 120/22 


Male: Gnathosoma as in A. casalis s. str. Dorsal shield (402 » * 222 mp) 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 sfI, 57 p»; 
between sft and st3, 66 yw. Par- and postanal setae about 20 p long. 

Chaetotaxy of legs asin female. Length/width (in ,) of leg segments : 


I II III 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 


a0 


Fic. 11. A. Androlaelaps casalis myrmecophila 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 w; movable digit 22 up. 
Four pairs of gnathosomal setae with c.s. about 45 p» apart, hyp.2 about 33 p apart. 
Corniculi 27 » long ; pedipalp (1-4-5-12) with two-tined apotele. 

Podonotal shield (210 » X 186 pw) 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 yp, laciniae 54 w. Sternal shield (about 144 p x 93 p) 
with anterior margin indistinct. Between stz, 63 ~; between sfx and st3, 102 p. 
Anal shield (48 » x 60 ») with par- and postanal setae about 24 » long. Peritreme 
extends to middle of coxa III. 

ZOOL. I4, 5. BE 


156 G. OWEN EVANS @& W. M. TILL 


Chaetotaxy of legs normal. Length/width (in ,») of leg segments : 


I Il 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 p. 
Four pairs of gnathosomal setae with c.s. about 42 p» apart, hyp.2 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 sf1, 54 « ; between sf1 and st3, 78 ». Anal shield (48 » x 45 »; maximum 
width 60 »); par- and postanal setae about 20 p long. Peritreme extends to 
posterior third of coxa 1; peritrematal shield weakly developed. 

Chaetotaxy of legs as in female. Length/width (in yp) 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 


HABITAT: From the nests of Formica rufa Linn. Holotype 2 (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 glasgour : 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 mohrae Oudemans, 1928, Ent. Ber. Amst. 7 : 374. 

Haemolaelaps cricetophilus Vitzthum, 1930, Zool. Jb. Jena, Abt. Syst. 60: 417. 
Laelaps stegemani Hefley, 1935, |. Kansas ent. Soc. 8: 22. 

Ischnolaelaps rhabdomys Radford, 1939, Parasitology 31 : 249. 

Haemolaelaps scalopi Keegan, 1946, Trans. Amer. Micr. Soc. 65: 71. 


Atricholaelaps stranditmanni Fox, 1947, Ann. ent. Soc. Amer. 40: 580. 
Hypoaspis (Haemolaelaps) eos Zumpt & Patterson, 1951, J. ent. Soc. S. Afr. 14: 


Atnricholaelaps sigmodoni Strandtmann, 1946, J]. Parasit. 32 : 164. 
79 


LHE BROLLISoH DERMANYSSIDAE. (ACARI) 157 


Female ; Chelicera with segment I, 72 » ; II, 126 1; 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 w apart ; hyp.2 
about 54 wapart. 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 » X 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 7 and seven pairs of # setae on 
the cuticle. 


Fic. 12. Androlaelaps fahrenholzi (Berlese), female ; dorsum of idiosoma (a); chelicera 
(B) ; venter of idiosoma (c). 


Tritosternum with base 45 p, laciniae 135 pw. Sternal shield (96 » x 168 p) with 
three pairs of setae and two pairs of pores. Metasternals free with associated pores. 
Between sti, 96 » ; between sti and st3, 102 w. Genital shield tongue-shaped (156 pu 
X I14 p»), with genital setae only. Anal shield (90 w X 112 pw) with paranal setae 
54 pw, postanal seta 66 uw. 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 in having two postero-lateral setae 
on genulV. Length/width (in p) of leg segments : 


155 G. OWEN EVANS & W. M. TILL 


I II 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, Acarofauna ital. part I: 55. 


Type: Gamasus kramerit 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 norma], 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 #x3 (37 pairs), 
r2 and r4 on lateral integument (37 pairs), 23 and rr absent (37 pairs), presence of 
extra 7 seta (40 pairs) or extra ry 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 pl, 
on tibia III (2—4+, ?—2), seta pl, or pu on genu IV (2—#, $—2 or 2—#, #—1) and 
the absence of al, on genu IV (1—#, §—1). Anterior spine on coxa II small or 
absent. Ambulacra usually with paired claws ; claws on leg I occasionally reduced 


THE BRELISH DER IMANYSSIDAE (ACARI) 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. kramerz (Canestrini). 
ALLOPARASITUS Berlese, 1920 (type: Laelaps (Hypoaspis) oblongus Halbt., 
1915) syn: Halbertia, Hullin 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 aculerfer 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 ; 
Hi. 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 Zampt & 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. 

® 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 


2 


G. OWEN EVANS & W. M. TILL 
KEY TO FEMALES 


Genu IV with eight setae, an antero-lateral seta lacking (t—}?, 3—1) ; peritrematal 
shields free posteriorly and Se well beyond posterior margins of coxae IV 
(Text-fig 46F) ‘ ; ‘ : ; H. oblonga (Halbt.) (p. 222) 

Genu IV with 9 or 10 setae, twoantero- Jaiebilatine present (2—#, §—1 ; 2—?, #—1 or 
2—#, 3—2); Rag ay re not Serne as to coxae IV, free or fused with 
the podal shields . ‘ : 2 

Dorsal shield lacking setae px on the eideenaial region a never with om 
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 . ° ; : . : 3 


(Footnote continued from p. 159) 


I 


te 


Genu IV with eight setae (1—}?, §}—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 ped 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 
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 
Dorsal setae spatulate, cuneiform or scimitar-like. ' Free-living or myrmecophilous 
Cosmolaelaps Berlese 
Dorsal setae simple or pilose, never markedly flattened 
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 never, long and whip-like : F : ‘ 
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 (2-2, #—1/2), if with two ventral setae, then ara ta 
narrow 
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) . 
Digits of the chelicerae in the female stumpy, dentate or edentate and modified for oophagy, 
movable digit about 20 4 in length. Myrmecophilous 
Holostaspis \Kolenati 
Digits of the chelicerae in the female normal in form, dentate or edentate ; movable digit at 
least 35 mw in length 
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 
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) 


4 
i 
1 


ln 


13 


14 


THE BRITISH DERMANYSSTDAE (ACARI) 161 


Dorsal setae j2—3, 22-4, S4—5 and 24 considerably longer than other dorsal setae, Z4 
very long and whip-like (Text-fig. 13a) ; seta pd, on femur II, ad, on femur IV, 
ad,, and pd, on tarsus IV long and os -like. Associated with Lucanus 
feeteopicra) ‘ H. krameri (Canestrini) (p. 163) 

Dorsal chaetotaxy otherwise ; on Sets never — and whip-like : : : 

Dorsal setae spatulate (Text- ms 24c), cuneiform ae me 2 a or scimitar-like 
(Text-fig. 28A—c) . 

Dorsal setae simple or pilose, never sn aaebeatee finttoned 

Majority of dorsal setae cuneiform (Text-fig. 25c) . 

Majority of dorsal setae spatulate or scimitar-like. : : : ; : 

Genu IV with two postero-lateral setae (2—2?, 3—2); dorsal seta S2 clavate, 
considerably shorter than cuneiform seta S1 (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, 2—1) ; dorsal setae S1—2 cuneiform, 
subequal (Text-fig. 26a) ; proximal region of cuneiform seta conspicuously 
enlarged (Text-fig. 26D). : 4 : 5 . 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. 


aN 


N Don 


27B) .. i : H. vacua (Michael) (p. 190) 
Genu IV with nie el See Pe eee ee 3—1T) , : : ; , 9 
Genu IV with one ventral seta, pu absent (2—?, 3—1 or 2—?, 3—2) : 16 
Capitular setae and setae on coxae I—II stout, spinelike. Associated with ne MS 

(Hymenoptera) . : " : H fuscicolens (OQudemans) (p. 196) 
Capitular setae and setae on coxae me IT slender, hopaal ; ste) 
Dorsal shield shows hypertrichy between the J series ; surface of qn aa a clearly 

defined reticulate pattern ; II 
Dorsal shield without hypertichy ce ee the J series ; ones of shield 8h or 

without a distinct reticulate pattern : ; ‘ ; ‘ : 14 
Sternal shield distinctly wider than long (L/W = o: 7); ; 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 ie ade one == 0 1) ; : 
deutosternum with six rows each of at least six small, uniform, denticles. : 13 


Opisthogaster densely setose, region between genital and anal shields with eee 
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 yw in length ; seit “ Ste at least three-quarters of the 
distance between 72 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 72 extending to about one-half the distance 
between j2 and 73 ; setae on distal third of tarsus II not distinctly thickened nor 
spine-like. Associated with Bombus ; H. minutissima sp. nov. (p. 201) 

Sternal shield longer than wide (L/W = 1:1). JSpcinoecseds 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. 318) 

H. colomboi sp. nov. (p. 194) 


162 


15 


ne 
~J ’ 


2! 


22 


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 oe ee shield not more than twice 
the width of the peritreme __. : : . ; . 15 
Setae on anterior part of dorsal shield relatively long ; seta 73 reaches base of seta 
j4 (Text-fig. 29a) ; surface of shield with distinct reticulate pattern in addition to 
sculptured areas for muscle attachment . : 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—}?, 3—2) (Text-fig. 44c) : : : 17 
Genu IV with one postero-lateral seta (2—}?, 3—1) (Text- -fig. 44B). ‘ ; 19 
Antero-ventral seta on femur II and genu II stout and spine-like (Text-fig. ‘20D) ; 
tibia III with two postero-lateral setae (2—}, #—2) ; dorsal shield with 39 pairs of 
setae (Text-fig. 20a)  . . H.sardoa Berlese (p. 175) 
Antero-ventral seta on femur II and genu Il not conspicuously thickened ; tibia III 
with one postero-lateral seta (2—}, ae ; dorsal shield with 37 or 41 pairs of 


setae. Myrmecophilous . 18 
Dorsal shield about 1,100 yu in length with 41 pairs of short, fine, dorsal setae (Text: 
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. : : 20 
Peritrematal shield free posteriorly . : 21 
Dorsal setae short, setae J/1 reaching about mid- “way between the bases of Ji and pP 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 J2 by about one-half their 

length (Text-fig. 22A) ; 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. ; 23 
Larger species, over 1,000 wu ; sternal shield with L/W = 0° 8: genito- -ventral shield 

as in Text-fig. 42B : > H. myrmecophila (Berlese) (p. 214) 
Smaller species, about 700 ys ; 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 RF series (Text-fig. 19A) ; peritreme extend- 
ing to about the middle of coxa II (Text-fig. 19E) ; Small species about 500 4 


inlength . ‘ ; : H. praesternalis Willmann (p. 173) 
Dorsum with five or more pairs of setae in the RF series ; Bierirere gc up to or 
beyond coxa I ; 24 


Majority of dorsal and opisthogastric setaé very long, sinuous  (Text- figs. 40 A and E) ; 
setae ]3 extending beyond the bases of setae J4 by at least a quarter of their 


length. Myrmecophilous . ; . : ‘ : : 25 
Dorsal and opisthogastric setae normal in length ; setae ]3 never reach the bases of 
d4 ; _ : : ; “ 26 


Legs rel: ativ ely short and stumpy : leg IV (trochanter to tarsus) about one-half the 
length of the dorsal shield ; movable digit of the chelicera strongly bidentate 
(Text-fig.40p). . F ; H. humerata Berlese (p. 212) 

Legs relatively long and slender leg IV (trochanter to tarsus) scarcely shorter than 
the dorsal shield ; movable digit of chelicera edentate (Text-fig. 40B) 

H. equitans (Michael) (p. 210) 


LOE BVT isn 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. : : ‘ ‘ : : : : : 27 
27 Cheliceral digits normal, slender ; movable digit longer than the palpfemur and 
strongly bidentate , . d : : : : . ; ; 28 
— - Cheliceral. digits atypical, Sonny movable ar shorter than the gear 
weakly dentate or edentate . : 29 


28 Genito-ventral shield greatly expanded siete to coxae IV: Sosees ie 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 
Hi, 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 J1—J3 (Text-fig. 38a). 
Myrmecophilous . ‘ i H. oophila (Wasman) (p. 207) 
— Dorsal setae of the J series abeanele in length : ; 30 
30 Dorsal seta Z5 pilose distally and about three times the lenath ei ae seta ae 5 
(Text-fig. 374A) ; 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. 36a) ; movable digit of 
the chelicera dentate (Text-fig. 36D) ‘ H. isotricha (Kolenati) (p. 203) 


Hypoaspis krameri (Canestrini) 


Gamasus kvameri Canestrini G. & R., 1881, Atti Ist. Veneto (5) 7: 1083, figs. 


Female: Chelicera with segment I, 72 » ; II, 180 » ; movable digit 80 yp, bidentate ; 
fixed digit with about ten teeth (Text-fig. 13D) ; pilus dentilis short, setiform ; dorsal 
seta about twice as long asthe pilus. Four pairs of gnathosomal setae with c.s. about 
72 wapart, hyp.2 about 60 » apart. Deutosternum with six transverse rows of denticles 
(6-7 per row) ; corniculi 66 » ; internal malae as in Text-fig. 13c. 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 Z4 extremely long 
and sinuous ; no hypertrichy. Surface of shield granular. 

Tritosternum with base 45 p, laciniae 123 p. Sternal shield (117 pw 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 sf1, 93 »; between sf1 and 
st3, 114 p. Genital shield (114 » X 105 pw) with genital setae only. Anal shield 
(93 » X 72 ») with paranal setae 48 p, postanal seta 30 w. Chaetotaxy and sclero- 
tization of opisthogaster as in Text-fig. 13F. Peritreme extends beyond middle of 
coxa 1; peritrematal shields free anteriorly and posteriorly. 

Legs with normal chaetotaxy. Long, whip-like setae present on femur II (d,), 
femur IV (ad,) and tarsus IV (ad, and fd,-;), Text-fig. 14a. Tarsus II with two 


G. OWEN EVANS & W. M. TILL 


“wit 


qi 


Fic. 13. Hypoaspis krameri (Canestrini). A. Dorsum of idiosoma of female; 8B. tectum 
capituli of female; c. venter of gnathosoma of female; D. chelicera of female; 
x. chelicera of male; F. venter of idiosoma of female. > 


THE BRITISH DERMANYSSIDAE (ACARI) 165 


stout, blunt setae (a/, and #/,) and several which are stout basally, tapering to fine 
points (av,~», Pv;-», mv) ; relatively thick setae on tibia II (av, and pv,) and genu II 
(av). Length/width (in ») of leg segments : 


I Il 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 », unidentate ; 
spermadactyl 130 pw long, slender (Text-fig. 13E); fixed digit tridentate ; pilus 
dentilis and dorsalseta asinfemale. Four pairs of gnathosomal setae with c.s. about 
60 » apart, dyp.2 about 50 pw apart. Details of gnathosomal structure not clearly 
visible. 

Dorsal shield (636 » x 348 pw) as in female. Tritosternum with base 45 uz, 
laciniae 114 ». Holoventral shield (516 » x 114 p) 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 sfI, 80 wp; 
between s¢i and s?#3, 123 pw. Peritreme extends beyond middle of coxa I. 

Legs with normal chaetotaxy ; long whip-like setae on legs II and IV as in female. 
Leg II (Text-fig. 148) bears several stout setae : av, and especially a v, on the femur ; 


Fic. 14. Hypoaspis kramevi (Canestrini). a. Leg IV of female; sB. leg II of male. 


166 G. OWEN EVANS & W. M. TILL 


av on the genu ; av and pv on the tibia ; al,, pl,, v2, pug, 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 
nasicornts Linn. in the Netherlands. 


Hypoaspis aculeifer (Canestrini) 


Laelaps aculeifer Canestrini, 1884, Altt Ist. Veneto (6) 2 : 608. 
Hypoaspis aculeifer: Bregetova, 1956, Opred. Faune SSSR. 61: 77, figs. 102-106. 


Female : Chelicera with segment I, 75 » ; II,216; movable digit 98 », bidentate ; 
fixed digit with about Io 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 » 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. I5A. 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 yp, laciniae 105 pw. Sternal shield (190 » X 138 yp) 
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 p; 
between sf1 and sf3, 144 w. Genital shield flask-shaped (120 » x 96 u) with one 
pair of setae. Anal shield (go » x 75 m), with paranal seta 27 » and postanal seta 
36 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. I5C. 
Peritreme extends to middle of coxa I ; peritrematal shields free posteriorly, anterior 
region fused with dorsal shield at level of rr. 

Legs with normal chaetotaxy. Tarsus II with setae av,—», pv,-2, al,, ply, md, mv, 
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 pv), tibia III (av 
and pv), tarsus III (most setae, except al,, pl,, ads, pd), genu IV (av), tibia IV (avy, 
pv,, pl,) and tarsus IV (most setae). Length/width (in p) of leg segments : 


I II Ill IV 
genu 96/40 105/66 63/50 105/63 
tibia 114/30 96/66 66/45 120/63 


tarsus 180/28 135/48 156/40 240/48 


Male ; Chelicera with segment I, 54 »; Il, 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 p apart, hyp.2 


THE BRITISH DERMANYSSIDAE (ACART) 167 


Fic. 15. Hypoaspis aculeifey (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 y) with 39 pairs of setae distributed as in Text-fig. 15F. 
Tritosternum with base 27 p, laciniae 78 p. Holoventral shield (456 » x 114 p) 
with three pairs of sternal, one pair of metasternal setae and three pairs of pores in 
the sternal region ; between st1, 63 »; between sti and st3, 126 ». Genito-ventral 
region with six pairs of setae. Paranal setae about 24 p, postanal seta about 20 p 
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 II 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 up. 
Four pairs of gnathosomal setae with c.s. about 45 » apart, Ayp.2 about 36 pw apart. 
Deutosternum, internal malae and tectum as in female ; corniculi 36 » long. Pedi- 
palp (I-4-5-12) with two-tined apotele. 

Podonotal shield (258 » * 190 ») with eleven pairs of setae ; pygidial shield (go » 
120 p) with eight pairs of setae; chaetotaxy and sclerotization of mesonotal 
region as in Text-fig. 164A. 

Tritosternum with base 30 yp, Jaciniae 60 p. Sternal shield (170 » x 80 pw) with 
three pairs of setae and two pairs of pores ; between sf1, about 48 »; between s?r 
and st3, about 120 ». 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 III. 

Legs with normal chaetotaxy. Length/width (in ,) of leg segments : 


I II Ill 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 »; II, 147 »; movable digit 57 uz. 
Deutosternum, hypostomal processes, corniculi and tectum as in protonymph. 
Pedipalp (2—5—6-14) as in female. 

Dorsal shield (450 » * 216 yw) with lateral incisions, bearing 39 pairs of setae 
arranged as in Text-fig. 16¢ ; no hypertrichy; with seven pairs of R setae on the 
cuticle. 


THE BRITISH DERMANYSSIDAE (ACART) 


NaZ, 


Fic. 16. Hypoaspis aculeifer (Canestrini). A. Dorsum of idiosoma of protonymph; B. 


venter of idiosoma of protonymph; c. dorsum of idiosoma of deutonymph ; D. venter 
of idiosoma of deutonymph. 


169 


170 G. OWEN EVANS & W. M. TILL 


Tritosternum with base 40 p, laciniae 65 ». Sternal shield (243 » x 82 p») bears 
three pairs of sternal, one pair of metasternal setae and three pairs of pores ; between 
stt, 60 w; between sft and s¢3, 114 w. Anal shield (approximately 57 » x 50 ,) 
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 fl, stout. Length/width 
(in ») of leg segments : 


I Il 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 


HaBiTAT: 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 pu, 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 p» apart, hyp.2 about 36 p apart. 
Deutosternum with six transverse rows of denticles (3-7 per row) ; corniculi 36 p» 
long ; internal malae reduced (Text-fig. 178). Tectum capituli with anterior margin 
smooth (Text-fig. 17c). Pedipalp (2—5—6—14) with two-tined apotele. 

Dorsal shield (480 » * 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 » x 84 ») weakly 
reticulate anteriorly and laterally, with three pairs of setae and two pairs of pores. 
Metasternal setae free with associated pores. Between sti, 60 »; between str and 
st3, go w. Genital shield (84 » x 93 ») with genital setae only. Anal shield (60 » 
72 mw) 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 I ; 
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, al,, pl, and md stouter 
than the other setae ; tarsus 1V with seta fl, spine-like. Length/width (in ,) of leg 
segments : 


I Il 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 


THE BRITISH DERMANYSSIDAE (ACARI) 


| Bic. 17. Hypoaspis giffordi sp. nov., female 

| and hypostome (B) ; tectum capituli (c) ; chel 
| ZOOL..14, 5. 

| 


; dorsum of idiosoma (aA); deutosternum 
icera (D) ; venter of idiosoma (E). 


172 G. OWEN EVANS &@ W.M. TILL 7 


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 (1-0). 
Hypoaspis (Haemolaelaps) inversus 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. 


eS ee. 


Female ; Chelicera with segment I, 66 » ; II, 156 » ; movable digit 56 p, bidentate ; 
fixed digit tridentate ; pilus dentilis short, setiform (Text-fig. 188). Four pairs of 
gnathosomal setae with c.s. about 66 p apart ; hyp.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 (8); tectum capituli (c) ; venter of idiosoma (D). 


THE BRITISH DERMANYSSIDAE’ (ACART) ‘73 


Dorsal shield (708 » X 420 ») with 39 pairs of setae and 4-9 accessory setae 
distributed as in Text-fig. 18A. Surface of shield reticulated. 

Wmrosternum with base 30 p, laciniae go p. Sternal shield (132 » xX 120 2) 
extending to middle of coxa III, with three pairs of setae and two pairs of pores ; 
metasternal setae free with associated pores. Between sf1, 70 4; between sti and 
st3, 112 p. Genital shield (150 w x 117 p) flask-shaped with one pair of setae. Anal 
shield (II0 » X 105 yp) 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 IV. Setae av,, pvj, al,, pl, on tarsus II are stout and spine-like ; setae avg, pu, 
and mv are stout and pointed. Tibia II hasstout ventral setae (av, and fv,). 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 


HasitaT: In decaying oats and hay ; in the nest of Reparia riparia (Linn.) ; ona 
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. zool. Mus. Berlin 38 : 62, figs. (syn. nov.). 


Female: Chelicera with segment I, 50 »; II, 144 »; movable digit 60 p, biden- 
tate ; fixed digit with bifid tip and four teeth, pilus dentilis short, setiform. Four 
pairs of gnathosomal setae with c.s. about 45 pw 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. IgA. 


_ Nohypertrichy. 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 p) with reticulate pattern most pronounced laterally ; 


| 
{ 
; 
} 
j 


| 


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 pw. Genital shield (90 pw X 72 p) 


_ with genital setae only. Anal shield (72 » x 70 ») with paranal and postanal setae 
_ about 33 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. I9E. 


174 


G. OWEN EVANS & W. M. TILL 


Fic. 19. Hypoaspis praesternalis Willmann. A. 


chelicera of male; c. tectum capituli of female ; 
fermale: &. venter of idiosoma of female. 


Dorsum of idiosoma of female; B. 
p. peritreme and peritrematal shield of 


BEE SRETISH 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-LV with ventral setae relatively stout basally, 
tapering to long, fine points. Tarsus IV with setae £/, and f/; relatively stout and 
blade-like. Length/width (in ») of leg segments : 


I II Ng. TV 
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 LEG 27 


Male ; Chelicera with segment I, 40 »p ; II, 105 » ; movable digit 42 w, 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 pw apart, hyp.2 
about 36 m apart. Corniculi 36 w 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 #x2 in the opisthonotal region. Surface of shield reticulated. 

Tritosternum with base 15 p, laciniae 704. Holoventral shield (348 pw xX 93 p) 
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 sf1, 66 yp; 
between sti and s#3, 90 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 asin female. Setae of femur II simple. Length/width (in p) 
of leg segments : 


I iT III IV 
genu 66/27 54/36 36/30 ayia 
tibia 68 /24 48/33 36/27 60/30 
tarsus 117/20 Gai27 70/24 105/26 


HasitatT: 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 «1; movable digit 90 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 yu apart, 
hyp.2 about 57 » apart. Deutosternum with six transverse rows of denticles (4-8 
per row); corniculi 90 » long; internal malae fringed (Text-fig. 20F). Tectum 


G. OWEN EVANS & W. M. TILL 


um of idiosoma (a) ; tectum (B) ; cheli- 
tosternum and hypostome (£) ; venter 


Fic. 20. Hypoaspis sardoa (Berlese), female ; dors 
cera (c); femur, genu and tibia of leg II (p); deu 
of idiosoma (F). 


DoE British, DERMANYSSIDAE ,(ACART) 177 


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 » X 456 ») with 39 pairs of setae (Text-fig. 20A). 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 p. Sternal shield (170 uw xX 156 yp) 
with three pairs of setae and two pairs of pores ; metasternal setae free with associated 
pores. Between sf1, 84 »; between sti and st3, 160 pw. Genital shield (216 p x 
156 p) 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,-,, fv;-, and mv are stout and pointed. Length/width (in p) 
of leg segments : 


I II Ill 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 


HABITAT: 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.S5.S.R. Recorded by Bregetova from the nest of Phoenicurus phoentcurus 
 (Linn.). 


Hypoaspis heselhausi Oudemans 


_ Hypoaspis heselhausi Oudemans, 1912, Ent. Ber. Amst. 3 : 216. 
Bregetova, 1956, Opred. Faune SSSR. 61 : 78, figs. 


_ Female: Chelicera with segment I, 45 » ; II, 126»; movable digit 48 py, 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 w apart. Deutosternum with six transverse rows of 
| denticles (4-6 per row); corniculi 36 » long; internal malae as in Text-fig. 218. 
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 yw, laciniae 72 p. Sternal shield (108 p X 102 p) 
with three pairs of setae and two pairs of pores ; metasternal setae free with associa- 


| 
| 


178 


G. OWEN EVANS & W. M. TILL 


pi =e hy 
‘ An NOV, vy 
seca, Sandee D 


Fic. 21. Hypoaspis heselhausi Oudemans, female ; dorsum of idiosoma (A) ; deutostern n 
and hypostome (B); chelicera (c) ; venter of idiosoma (D). : 


Tae ERVrish DERMANYSSIDAE: (ACARI) 179 


ted pores. Between sii, 56 »; between sf1 and sf3, 96 w. Genito-ventral shield 
(108 » X 84 ») with maximum width (135 ») near posterior margin, bearing a pair of 
genital and a pair of opisthogastric setae. Analshield (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 p) of leg segments : 


I II Ul IV 
genu 60/27 57/38 36/30 57/33 
tibia 63/27 48/36 36/30 57/30 


tarsus 108/25 G2 27 80/24 90/27 


HapitaT: 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 »; movable digit 75 p», biden- 
tate; fixed digit tridentate ; pilus dentilis short, setiform (Text-fig. 23E) Four 
pairs of gnathosomal setae with c.s. about 54 mw apart, hyp.2 about 42 wp apart. 
Deutosternum with six transverse rows of denticles (6-7 per row) ; corniculi 66 pu 
long ; internal malae with a deep fringe as in Text-fig. 23F. Tectum capituli with 
anterior margin smooth (Text-fig. 238). 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. 22A. Surface of shield reticulate and granular. 

_ Tritosternum with base 48 yp, laciniae 102 pw. Sternal shield (110 mw 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 sf1, 70 w; between 
_stI and st3, 105 mw. Genito-ventral shield (216 w X 120 pw) flask-shaped, posterior 
margin truncate, very close to anal shield; bearing one pair of genital and three 
pairs of opisthogastric setae. Between /v2, 162 w. Anal shield (78 mw X 105 p) 
with maximum width at anterior margin (132 ») ; paranal setae 30 » ; postanal seta 
45 p. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 228. 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. 


j 


180 G. OWEN EVANS & W. M. TILL 


Fic. 22. Hypoaspis nidicorva sp. nov., female; dorsum (A) and venter (B) of idiosoma. 


Chaetotaxy oflegsnormal. Dorsalsetaon trochanter IV relatively stout. Length/— 
width (in ») of leg segments : 


I II II] IV 
genu 84/36 75/42 54/30 78/30 
tibia go/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 p», 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 p apart, hyp.2 
about 36 » apart. Corniculi 48 » long. Deutosternum, internal malae, tectum and — 
pedipalps as in female. 

Dorsal shield (396 » * 267 ) as in female, with 39 pairs of setae and one or two 
accessory setae. Tritosternum with base 36 yw, laciniae 105 pw. Holoventral 
shield (360 » x 96 «) with four pairs of setae and three pairs of pores in the ste 


THE BRITISH DERMANYSSIDAE (ACARI) 181 


region, one pair of genital and five pairs of opisthogastric setae in the genito-ventral 
region (Text-fig. 23a). Between stz, 48 »; between sf and st3, 90 p. Paranal 
setae 24 w; postanal seta 36 w. Peritreme extends beyond posterior margin of coxa 
I; 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; Bs. tectum capituli of 


female; c. chelicera of male ; D. femur II of male; £. chelicera of female ; F. venter 
of gnathosoma of female. 


182 G. OWEN EVANS & W. M. TILL 


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 p. 
Gnathosoma generally as in female ; setae c.s. about 48 p apart, hyp.2 about 36 pw 
apart ; corniculi 45 » long. 

Dorsal shield (444 » X 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 p. Sternito-genital shield (246 mw X II4 p) ~ 
with four pairs of setae and three pairs of pores; between stI, 70 »; between sf1 
and st3, 108 ». Anal shield (66 » Xx 75 m) triangular with maximum width (100 ,) 
at anterior margin; paranal setae 24 p, postanal seta 40 pw. 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 
(1965 : 12 : 29: 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 p 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. 3 

Dorsal shield (654 » * 366 ») with the normal 39 pairs of setae and four un 
accessory setae, all spatulate in shape (Text-fig. 24c) and arranged as in Text-fig. 2 
Surface of shield reticulate ; seven pairs of R setae on the cuticle. 


2HE BRITISH DERMAN YSSIDAE (ACARI) 183 


Fic. 24. Hypoaspis clavigey (Berlese). A. Dorsum of idiosoma of female; B. dorsal seta 
J4 of female; c. dorsal seta Z5 of female; D. venter of idiosoma of female; E. venter 
of idiosoma of male; F. chelicera of male; G. chelicera of female. 


184 G. OWEN EVANS & W. M. TILL 


Tritosternum with base 45 », laciniae 80 ». Sternal shield (110 » x II0 pw) with 
three pairs of setae and two pairs of pores ; metasternals free with associated pores. 
Between sf1, about 70 »; between sf and st3, 102 ». Genital shield flask-shaped 
(105 # X 102 p), with one pair of setae. Anal shield (102 » x 114 ») with paranal 
and postanal setae about 20 » 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 p») of leg segments : 


I II Ill 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 of 
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 p) with 39 pairs of setae and five unpaired accessory 
setae ; seven pairs of RX setae on the cuticle. Tritosternum with base 30 p, laciniae 
60 ». Holoventral shield (360 » x 96 ») with three pairs of sternal, one pair of 
metasternal setae and three pairs of pores in the sternal region ; between sfI, 54 p»; 
between sf1 and st3, 93 ». Genito-ventral region with six pairs of setae ; paranal 
setae 18 », postanal seta 15 ». 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 II III IV 
genu 80/36 63/42 50/36 78/40 
tibia 80/30 56/40 48/36 80/36 


tarsus 135/24 117/30 120/30 170/33 


Hapitat: 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 cunetfer; Bregetova, 1956, Opred. Faune SSSR. 61: 75. 


Female: Chelicera with segment I, 78 »; II, 234 »; movable digit 84 p», biden- 
tate ; fixed digit with about seven teeth, pilus dentilis short, setiform (Text-fig. 25F). 


THE BRITISH DERMANYSSIDAE (ACARI) 185 


Four pairs of gnathosomal setae with c.s. about 90 » and hyp.2 about 72 apart. 
Deutosternum with six transverse rows of denticles ; corniculi 50 pu long ; internal 
malae with long slender filaments as in Text-fig. 25H. Tectum capituli with denti- 
culate anterior margin (Text-fig. 25D). Pedipalp (2-56-14) with two-tined apotele. 

Dorsal shield (804 » x 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 C) 
Surface of shield with reticulations consisting of rows of tiny spine-like markings. 

Tritosternum with base 54 yw, laciniae 123 p. 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 SHI, 
72 »; between sfx and st3, 153 w. Genital shield (168 p» x 204 «) expanded behind 
genitalsetae. Anal shield (114 » x 126 ») with paranal and postanal setae subequal, 
24-30 w long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 25B. 
Peritreme extends to middle of coxa I; 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—{7, 52). Most leg setae simple, some dorsal setae (on femora and 
trochanters) spatulate. Length/width (in y) of leg segments : 


I II 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 p (84 » 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 » apart. Corniculi 45 » long; deutosternum, internal malae, tectum and 

-pedipalps as in female. 

Dorsal shield (636 » xX 546 «) 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. 25E). Bet- 
ween siI, 84 »; between sf1 and sf3, 123 p. Par- and postanal setae subequal in 
length, about 30 ». Peritreme extends to middle of coxa I ; peritrematal shield 
fused anteriorly with dorsal shield, posterior part free. 

_ Chaetotaxy of legs as in female. In addition, seta av, on femur II is large and spine- 
like (Text-fig. 25G); setae av on genu and tibia II are considerably stouter than 


‘Setae pv. Length/width (in yp) of leg segments : 


| I II I 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 


G 


Fic. 25. Hypoaspis cuneifer (Michael). a. Dorsum of idiosoma of female; B. venter of 
idiosoma of female; c. dorsal seta 26 of female; pb. 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 


HABITAT: 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 1; movable digit 66 p, biden- 
tate ; fixed digit with from five to nine teeth ; pilus dentilis short, setiform. Four 
pairs of gnathosomal setae with c.s. about 75 pw apart, hyp.2 about 57 pw apart. 
Deutosternum with six transverse rows of small, fine denticles; corniculi 45 p; 
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 mw X 144 3B) 
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 sf1, 80 w; between sf1 and sf3, 126 w. Genital shield (123 p x 
153 p) flask-shaped, reticulated. Anal shield (110 » x 108 yw) 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 III 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 pu, 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 p apart, typ.2 about 50 w apart. Corniculi 36 » long; deuto- 
sternum, internal malae, tectum and pedipalps as in female. 

Dorsal shield (624 » X 408 yp) with 39 pairs of setae and about nine unpaired 
accessory setae. Surface of shield as in female. 

Tritosternum with base 33 p, laciniae 90 pw. Venter bears a sternito-genito- 
ventral shield (336 » X 126 ») and a separate anal shield (110 » X 96 ») ; the former 

ZOOL. 14, 5. 13 


188 G. OWEN EVANS & W. M. TILL 


Fic. 26. Hypoaspis neocuneifer sp. nov. A. Dorsum of idiosoma of female; B. venter of 
idiosoma of female; c. venter of idiosoma of male; pb. setae z4 and 26 of female; E. 
chelicera of male. "a 


tee 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 pw. Paranal setae 24 p, 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 p) of leg segments : 


I II III IV 
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 


HABITAT: Nests of Formica rufa Linn., White Hill Wood, Canterbury, 1st Novem- 
ber, 1942 and 25th April, 1942. Holotype 9 (1965 : 12 : 29: 23), allotype 9 (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 cunezfer (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. ensiger Berlese, 1904, Redia 1 : 420, figs. 


Female : Chelicera with segment I, 33 » ; II, 96 » ; movable digit 33 », bidentate ; 
fixed digit with five teeth, pilus dentilis short, setiform. Four pairs of gnathosomal — 
setae with c.s. about 50 m apart, Ayp.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. 274. All the setae except zI are scimitar- 
like. Surface of shield reticulate. 


J2 J3 J2 J3 J2 J3 


ry 


Fic. 28. Hypoaspis vacua (Michael), female. Variation in the form of dorsal setae J2 
and /3 and of the genital shield in (a), the type specimen from Igls (Austria) ; in 
(8), a specimen from Eire, and in (c), a specimen from Sussex (England). , 


1Be ERIGLISH DERMANYSSIDAE \(ACART) IQI 


Tritosternum with base 30 p, laciniae 72 w. Sternal shield (93 » x 90 p») with 
anterior margin extremely indistinct, bearing three pairs of setae and two pairs of 
pores ; metasternals free with associated pores. Between sf1, 54 »; between s?f1 
and st3, 70 pw. Genital shield (90 » x 78 yu) with lateral margins almost parallel ; 
bearing genital setae only. Anal shield (66 » < 63 yw) with paranal setae 18 yp long, 
postanal seta 21 wlong. 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 yp) of leg segments : 


I I II IV 
genu 60/30 51/33 36/27 60/33 
tibia 66/24 45/33 33/27 63/30 


tarsus i724 84/27 72,24 LIT /27 


HABITAT: 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 wapart. Deutosternum with six transverse rows of 
denticles (2-4 per row) ; corniculi 27 w; 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, x1 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, S and 7 series are considerably longer than the remaining setae. Surface of 
shield granular and reticulate ; no hypertrichy. 

Tritosternum with base 24 p, laciniae 81 pw. Sternal shield (84 » x 138 ») with 
three pairs of setae and two pairs of pores, metasternal setae free with associated 
pores. Between st1, 87 » ; between sti and st3, 99 w. Genital shield (120 » Xx 105 p) 
with genital setae only. Anal shield (96 » x 70 ») with paranal setae 18 yp, postanal 
seta 21 pw. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 20B. 


192 


G. OWEN EVANS & W. M. TILL 


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—?, 3—1). Length/width (in ,) of leg segments : 


I II II IV 
genu 75/45 75/57 50/48 72/45 
tibia 50/40 66/50 54/42 75/40 


tarsus 192/360 * 10a/4o ~~ 120/30, 144/33 


HABITAT: 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 w. Structural details are 
not visible as the chelicerae are retracted. Four pairs of gnathosomal setae with c.s. 
about 60 p» 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 pu) with 42-44 pairs of setae distributed as in Text-fig. 
30A. Setae px1-3 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 75 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 sti, 
75 »; between sf1 and st3, 102 pw. Genital shield (144 w X 150 yp) granular, with 
genital setae only. Anal shield (123 » xX 108 uw) granular; paranal setae 30 pw 
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 (fv present). Length/width (in y») of leg segments : 


I 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 (D) ; 
tectum capituli (£); chelicera (F). 


194 G. OWEN EVANS & W. M. TILL 


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 » apart, hyp.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 » x 336 yw) with 38 pairs of setae, px2 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 y») 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 
stt, 63 «; between sti and st3, 105 w. Genital shield (120 » x 138 y) with genital 
setae only. Anal shield (80 » * 78 ») with maximum width 86 »; paranal setae 


Tae br hiSt DER MAN YSSTDAE (ACARI) 195 


36 pw, postanal seta 30 p. Chaetotaxy and sclerotization of opisthogaster as in Text- 
fig. 31 B. Peritreme extends to about anterior third of coxa I. Peritrematal shield 
free posteriorly. 

Chaetotaxy of legs differs from normal pattern in that genu IV has two ventral 
setae (2—?, 3—1). Tarsus II with ventral setae and al, and #1, stout basally, 
tapering to fine points. Tarsus IV with ventral setae stout basally, tapering ; seta 
pl, stout and spine-like. Length/width (in ) of leg segments : 


I II Tit 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 pv (57 » with 
spermadactyl), unidentate ; fixed digit unidentate, pilus dentilis short, setiform. 
Four pairs of gnathosomal setae with c.s. about 54 u apart; hyp.2 about 42 p apart; 
Deutosternum with six transverse rows of denticles ; corniculi 36 »; internal malae 
triangular. Tectum capituli weakly dentate ; pedipalp as in female. 


Fic. 31. Hypoaspis colomboi sp. nov., female; dorsum (A) and venter (B) of idiosoma. 


196 G. OWEN EVANS & W. M. TILL 


Dorsal shield (546 » X 372 m) as in female. Holoventral shield (420 » X 102 yp) 
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 sf1 and st3, 96 ». Paranal setae 42 p, 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 ,) of leg segments : 


I II Ill 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 9 (1965 : 12 : 29: I), allotype ¢ (1965 : 12 : 29: 2). 


Hypoaspis fuscicolens Oudemans 


Hypoaspis fuscicolens Oudemans, 1902, Tijdschr. ned. dierk. Ver. (2) 7: 289 (nom. nud.) ; 1904, 
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 p apart, hyp.2 about 48 p 
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 y 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 pw. Sternal shield (84 » x 186 p») with 
three pairs of setae and two pairs of pores, metasternal setae free with associated 
pores. Between st1, 75 »; between sf1 and st3, 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 |; 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—7, {—1). Coxae I and II with very stout setae ; genu and tibia II with 
pu stouter than av; tarus II with seta mv short and stout. Ambulacrum I without — 
claws. Length/width (in ,) of leg segments : 


I 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 


tee BREPISH DERMANYSSIDAE (ACART) 
| 


: basis 
male ; holoventral 


dorsum of idiosoma (A) ; 


D. Hypoaspis colombo sp. nov., 


(e): 


olens Oudemans, female ; 


rt of idiosoma 


A-C. Hypoaspis fuscic 
té 


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 p», 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 ; hyp.2 about 42 p apart. 
Deutosternum with six transverse rows of deutosternal teeth (2-6 per row) ; corniculi 
33 #; 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 / 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 90 ». Sternal shield (84-90 » X 132-140 p) 
granular and reticulate, with three pairs of setae and two pairs of pores; meta- 


Pic. 33. Hypoaspis hyatti sp. nov., female; dorsum (a) and venter (B) of idiosoma. 


SHE BRITISH DEKRMANYSSIDAE: (ACARI) 199 


sternal setae free with associated pores. Between sti, 72 w; between sti and s/3, 
108 p. 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 p 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 ») of leg segments : 


I iu III IV 
genu 84/40 75/54 57/42 75/40 
tibia 90/36 68/50 57/36 78/36 


tarsus 140/33 110/36 Li a3 150/33 


HABITAT: On Bombus muscorum (Linn.), Scarborough, Yorkshire, October, 1964. 
Holotype 2 (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, Siz. Osterr. Akad. Wiss. Abt. 1, 162 (6) : 465, figs. 
II-12. 


Female: Chelicera with segment I, 48 »; II, 120 »; movable digit 42 », 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 » 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 4 xX 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, x1 being present, and about 13 unpaired 
accessory setae between the J 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 uw. Presternal area with a pair of 
pre-endopodal shields. Sternal shield (93 » X 144 p») with three pairs of setae and 
two paires of pores. Metasternal setae and associated pores situated over endopodal 
shields. Between st1, 66 »; between sti and s#3, 110 w. Genital shield (110 p x 
147 ») with genital setae only. Anal shield (90 » X 90 pw) with paranal setae 21 yp, 
postanal seta 24 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 34B. 
Peritreme extends to anterior third of coxa 1; posterior part of peritrematal shield 
free. 


200 G. OWEN EVANS & W. M. TILL 


E 


Fic. 34. Hypoaspis marginalis (Willmann). A. Dorsum of idiosoma of female; B. venter — 
of idiosoma of female; c. fixed digit of chelicera of female; p. deutosternum of female ; 
gE. chelicera of male; F. venter of idiosoma of male. . 


THE BRITISH DERMANYSSIDAE (ACART) 201 


Chaetotaxy of legs differs from the normal pattern in that genu IV has two ventral 
setae (2—?, 3—1). Tarsi II-IV with ventral setae stouter than the dorsals, tarsus 
IV with setae #/,-, spine-like. Length/width (in yp) of leg segments : 


I II ITI IV 
genu 78/40 / Bs tai 60/44 78/42 
tibia 84/36 66/50 57/40 78/38 


tarsus £32/33 102/38 117/36 156/33 


Male: Chelicera with segment I, 48 »; II, 110 »; movable digit 36 pw (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 p apart, hyp.2 about 42 wapart. Deutosternum with six transverse 
rows of denticles (5-6 per row) ; corniculi 36 w. 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 yp, laciniae 84 ». Presternal area witha 
pair of pre-endopodal shields. Holoventral shield (456 » xX 135 m) 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 
stI, 66 »; between sti and st3, 102 p. 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 py) of leg 


| 


/ 


} 
| 
' 
; 
’ 
| 


segments : 
I II Ill IV 
genu 78/36 78 /60 60/42 78/42 
tibia 84/36 66/54 54/36 —-72/36 


tarsus 132 /30 96/36 96/30 138/30 


HaAsBiTaT: 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 1»; movable digit 36 p, 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 pw 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 » 312 ») with 39 pairs of setae and about five accessory setae 
between the / series (Text-fig. 354A). Setae simple, subequal. Surface of shield 
granular and reticulate. 

Tritosternum with base 24 p, laciniae 84 ». 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 st1, 65 »; between sti and st¢3, 
80 ». Genital shield (102 » * 114 ») with genital setae only. Anal shield (60 p x 
75 ») with paranal setae 15 pw, postanal seta 18 ». Chaetotaxy and sclerotization of 
opisthogaster as in Text-fig. 358. 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 
setae. Tarsi Il and IV with ventral setae relatively long, stout basally, tapering to 
fine points. Seta p/, on tarsus IV long, spine-like. Length/width (in p) of leg 
segments : 


PRE BEITISH DERMANYSSIDAE (ACART) 203 


I II III 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. 117/24 


HABITAT: On a Bombus terrestris (Linn.) queen, Gosforth, Cumberland, Britain, 
collected by D. C. Lee, 3rd May, 1960. Holotype @ (1965 : 12 : 29: 5) and paratype 


Eaprdo> 712 : 20 : 6). 


Hypoaspis isotricha (Kolenati) 
Holostaspis isotricha Kolenati, 1858, Wien. Ent. Monaischr. 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 u apart, 
hyp.2 about 36 mw 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. 36B). 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 zr are barbed 
(Text-fig. 36c). Surface of shield sculptured anteriorly, reticulate posteriorly. 

Tritosternum with base 36 yp, laciniae about 72 w. Sternal shield (96 w x I14 pn) 
with three pairs of setae and two pairs of pores ; metasternal setae and pores situated 
over the endopodal shields. Between sti, 60 uw ; between sti and s#3, 108 ». Genital 
shield (168 » X 150 p») with genital setae only. Anal shield (84 w x 80 p) concave 
anteriorly ; paranal setae 36 », postanal seta 60 w. 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 yp) of leg 
segments : 


I II III 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. I4, 5. 14 


G. OWEN EVANS & W. M. TILL 


dorsum of idiosoma (A); tectum (B) ; 
thosoma (E); venter of idiosoma (F).' 


Fic. 36. Hypoaspis isotricha (Kolenti), female ; 
dorsal seta (c) ; chelicera (p); venter of gna 


THE BRITISH DERMANYSSIDAE (ACART) 


i 2 os eae ( 


\ al ail, y 


Hic. 37. Hypoaspis montana (Berlese), female ; dorsum of idiosoma (A) ; chelicera (s) ; 
venter of gnathosoma (c) ; venter of idiosoma (p). 


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 » 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 yp, laciniae 54 ». Sternal shield (105 » x 100 ») with 
weak striations laterally, bearing three pairs of setae and two pairs of pores ; meta- 
sternals free with associated pores. Between sfI, 50 »; between sf1 and s#3, 87 p. 
Genital shield (110 » X 95 p») with genital setae only. Anal shield (60 p x 57,) 
with paranal setae 24 », postanal seta 30 » 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 : 


I II Ill IV 
genu 63/30 51/30 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 p apart, hyp.2 
about 42 »apart. Deutosternum with six transverse rows of denticles (5—7 denticles 
per row); corniculi horn-like, 21 » long; internal malae weakly serrate basally 
(Text-fig. 38p). Tectum capituli with smooth margin (Text-fig. 38c). ro | 
(2-5-6-14) with two-tined apotele. 

Dorsal shield (600 » * 438 ») with 39 pairs of setae distributed as in Text-fig. 3845 ; 


ime BRITISH DERMANYSSIEDAEs(ACART) 


Fic. 38. Hypoaspis oophila (Wasman), female ; dorsum of idiosoma (a) ; chelicera (B) ; 


tectum capituli (c) ; deutosternum and hypostome (p) ; venter of idiosoma (E). 


208 G. OWEN EVANS & W. M. TILL 


a single accessory seta present between the / 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 », laciniae 60 ». Sternal shield (102 » X 117 ») with 
three pairs of setae and two pairs of pores ; metasternals free with associated pores. 
Between sti, 66 «; between sf1 and s¢3, 110 ». Genital shield (153 » X 140 p,) 
tapers posteriorly and bears a pair of genital setae only. Anal shield (80 p X 72 p; 
maximum width 96 ») with paranal setae 33 », 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 Ill 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 


HABITAT: Nests of Formica fusca Linn., Leeds, England (Hull Collection) ; 
recorded from several Formica species in Britain and Europe. 


Hypoaspis astronomica (Koch) 


Iphis astronomicus 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 »; movable digit 36 pu, biden- 
tate ; fixed digit with about five teeth, pilus dentilis minute, setiform (Text-fig. 39C). — 
Four pairs of gnathosomal setae with c.s. about 45 » apart ; Ayp.2 about 36 p 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. 39A). 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 w. Sternal shield (96 » x 96 p») with 
heavily sclerotized margins; with three pairs of setae and two pairs of pores; 
metasternal setae on small shields, associated pores on cuticle. Between st1, 63 »; 
between str and st3, 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 21 p, postanal seta 33 p» long. 
Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 39r. Peritreme ex- 


Fic. 39. Hypoaspis astronomica (Koch), female; dorsum of idiosoma (a); tectum (B) ; 
chelicera (Cc); venter of gnathosoma (p) ; venter of idiosoma (E). 


THE BRITISH DERMANYSSIDAE (ACARI) 


210 G. OWEN EVANS @ W. M. TILL 


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 ») of leg segments : 


I II III 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 


HasitaTt: 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 »; movable digit 42 »; both 
digits virtually edentate with two slight irregularities on the internal margin ; pilus 
dentilis short, setiform (Text-fig. 4oB). Four pairs of gnathosomal setae with c.s. 
about 50 » apart, Ayp.2 about 40 » 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 » * 486 ») with 39 pairs of setae and six unpaired accessory 
setae distributed as in Text-fig. 40A. 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 ») with 
three pairs of setae and two pairs of pores ; metasternal setae free with associated 
pores. Between sti, 78 »; between sti and st3, 108 pw. Genito-ventral shield 
(210 w & 222 pw) expanded behind coxae IV, with two pairs of setae. Anal shield 
(126 w & 105 ») 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 III IV 
genu 102/40 78/48 72/45 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). 


iE BRITISH, DERMANYSSIDAEB, (ACART) 


Fic. 40. A-c. Hypoaspis equitans (Michael), female ; dorsum of idiosoma (A) ; chelicera 
(B); venter of idiosoma (c). D—F. Hypoaspis humevata (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 », 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 » x 490 p) with 39 pairs of setae distributed as 
in Text-fig. 40£, and three unpaired accessory setae. Surface of shield reticulate. 

Sternal shield (go » xX 120 ,») with three pairs of setae and two pairs of pores ; 
metasternal setae free with associated pores. Between st1, 63 »; between sti and 
st3, 84 #. Genito-ventral shield (210 » xX 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 mu) with maximum width 123 p at 
anterior margin ; paranal setae 30 » long, postanal seta broken. Chaetotaxy and 
sclerotization of opisthogaster as in Text-fig. 4or. Peritreme extends approximately 
to middle of coxa 1; 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 Ill 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 


HaBiITAT: Found in ants’ nests [Tetramorium caespitum (Linn.)| in Britain 
(Cornwall) and in Luxemburg. 


Hypoaspis acuta (Michael) 
Laelaps acuta Michael, 1891, Proc. zool. Soc. Lond. : 652. 


Female: Chelicera with segment I, 60 »; II, 177 »; movable digit 57 pu, 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 ; Ayp.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—5-6-14) with two-tined apotele. 

Dorsal shield (720 » * 468 ,) with 37 pairs of setae distributed as in Text-fig. 41A ; 
setae 23 and rt lacking on the podonotal region ; about four accessory setae present 
on the opisthonotal region between the J series. Setae inflated sub-basally (Text- 
fig. 41C). 


Fic. 41. Hypoaspis acuta (Michael). a. Dorsum of idiosoma of female; 8. chelicera of 
male; c. dorsal seta 25 of female; p. holoventral shield of male; £. deutosternum and 
hypostome of female; Fr. 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 p. Sternal shield (140 p» X I50 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 sf1, 80 »; between sti and sf3, 129 pw. Genital shield (168 » x 170 yp) 
flask-shaped with genital setae only. Anal shield (105 » X 100 ,) pear-shaped, 
paranal setae 36 », 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—}, $—2). Length/width (in ,») of leg segments : 


I II 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 p» in length from the base of 
the movable digit (Text-fig. 418). Pilus dentilis and dorsal seta as in female. Four 
pairs of gnathosomal setae with c.s. about 54 mw apart; hyp.2 about 45 p 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 » X 354 ») with chaetotaxy as in female. Holoventral shield 
(450 » & 108 «) expanded behind coxae IV, bearing nine pairs of setae excluding the 
anals (Text-fig. 41D). Between sti, 72 »; between sti and st3, 100 wp. Paranal 
setae 36 yw, postanal seta 30 » long. Peritremes extend to middle of coxae I. 

Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : 


I II III IV 
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 », 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, hyp.2 
about 70 » apart. Deutosternum with six transverse rows of denticles ; corniculi 
g$ ; internal malae with slender inner and triangular median, pilose lobes, and 


THE BRITISH DERMANYSSIDAE (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; rF. chelicera of male. 


216 G. OWEN EVANS & W. M. 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 » X 768 «) with 41 pairs of slender, simple setae and five 
unpaired accessory setae (Text-fig. 42A) ; 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 yp, laciniae 168 » ; pre-endopodal shields large, reticulate 
and granular. Sternal shield (180 » X 240 yp) granular, reticulate antero-laterally, 
with three pairs of setae and two pairs of pores ; metasternal setae free with associa- 
tedpores. Between st1, 126-132 » ; between sfI 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 yp; 
postanal seta 30 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 
428. 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 (fd,-,), 
tarsus II (ventrals, al,, pl,, md), and femur IV (ad, and fd,). Seta p/ on genu IV 
stout and spine-like (Text-fig. 448). Length/width (in ,) of leg segments : 


I II Ill 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 »; movable digit 126 p», uniden- 
tate ; spermadactyl 140 yp, 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, Ayp.2 about 75 wapart. Corniculi 90 p 
long, internal malae not clearly visible. Deutosternum, salivary styli, tectum capituli 
and pedipalps as in female. 

Dorsal shield (1,080 » * 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 sti, 114 »; between str and st3, 195 pw. 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 as in female. Length/width (in ,) of leg 
segments : 


I II III IV 
genu 180/66 162/90 132/72 192/84 
tibia 210/60 150/84 126/66 198/78 


tarsus 270/54 220/60 240/54 300/60 


THRE BRITISH DERMANYSSIDAE (ACARI) Ae 


HaBitTaT: Found in ants’ nests in Britain (Cornwall, Devon, Wales, Isle of Wight) 
and Europe. 


Fic. 43. A-—B. Hypoaspis myrmophila (Michael), female ; dorsum (a) and venter (B) 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. 43A ; an extra pair of 7 setae is present in the podonotal 
region. | 

Tritosternum with base 40 yp, laciniae 114 p. Pre-endopodal shields present. 
Sternal shield (96 » x 180 ») reticulated antero-laterally, with three pairs of setae | 
and two pairs of pores; metasternal setae and associated pores situated over the 
endopodal shields. Between st1, 99 » ; between sfi and st3, 105 ». Genito-ventral — 
shield (210 « & 190 ,) 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. 43B. 
Peritreme extends to about middle of coxa I; posterior part of peritrematal shield 
free. 

Legs with normal chaetotaxy. Length/width (in p») of leg segments : 


I II Ill 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 y», uniden- 
tate ; spermadacty]l relatively short, 102 » from base of movable digit ; fixed digit 
with one large and two small teeth (Text-fig. 44D). Pilus dentilis and dorsal seta 
normal. Four pairs of gnathosomal setae with c.s. about 60 » apart, hyp.2 about 57 
apart. Corniculi horn-like, 54 p» long; internal malae pilose. Deutosternum, 
tectum and pedipalps as in female. 

Dorsal shield (612 » * 420 yw) with chaetotaxy as in female. Pre-endo 
shields present. Holoventral shield (516 » * 144 ») with ten pairs of setae, exclud 
ing the anals (Text-fig. 44£). Between st1, 87 »; between sti and st3, 117 p. 
Peritreme extends to about middle of coxa I. 

Legs with normal chaetotaxy. Genu and tibia II with seta pv stout ; tarsus | 
with ventral setae and al, and pl, stout. Length/width (in ,) of leg segments : 


THE BRITISH DERMANYSSIDAE (ACART) 219 


ie.44. a. H ypoaspis myrmecophila (Berlese), holoventral shield of male; B. H. myrmeco- 
phila, genu IV of female: c. Hypoaspis laevis (Michael), genu IV of female ; D. Hypoaspis 
myrmophila (Michael), chelicera of male ; E. H. myrmophila, holoventral shield of male. 
ZOOL. 14, 5. 


T5 


220 G. OWEN EVANS @& W. M. TILL 


I Il 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 


Hapitat: The nests of ants [Lastus 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 pu, 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 » 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 r 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 2} times the length of the base. Pre-endopodal 
shields present. Sternal shield (204 » X 246 p) 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 sti and st3, 192 pw. 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 ») 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—}, 4—2) as in Text-fig. 44c. Tarsus II with several stout setae (av,, pv,, aly, 
pl,). Length/width (in p) of leg segments : 


I II III 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 


Hapitat: Nests of ants [Tetramorium caespitum (Linn.)| in Cornwall and in 
Austria. 


THE BRITISH DERMANYSSIDAE (ACART) 


Fic. 45. Hypoaspis miles (Berlese), female ; dorsum of idiosoma (A); tecta capituli (B) ; 
chelicera (c) ; venter of gnathosoma (D) ; venter of idiosoma (g). 


222 G. OWEN EVANS & W. M. TILL 


Hypoaspis miles (Berlese) 


Laelaps (I phis) miles Berlese, 1892, Acari, Myr. Scorp. Ital. fasc. 63, no. 9. 
Cosmolaelaps gurabensis Fox, 1946, ]. Parasit. 32: 449, fig. 1. 
Bregetova, 1956, Opred. Faune SSSR. 61: 75, figs. (syn. nov.). 


Female: Chelicera with segment I, 96 »; II, 216 %; movable digit 130 yp 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 go yp long ; 
outer processes of internal malae fringed (Text-fig. 45D). Tectum capituli with 
denticulate anterior margin (Text-fig. 45B). Salivary styli 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. 45A ; setae x2 and px3 absent. Surface of shield reticulated ; no hyper- 
trichy ; seven pairs of X setae on the cuticle. 

Tritosternum with base 60 yp, laciniae 96 pw. Sternal shield (180 p x 138 ph) 
extending almost to posterior margin of coxa III, with three pairs of setae and two 
pairs of pores ; metasternals free with associated pores. Between st1, 96 » ; between 
str and st3, 144 ». Genital shield flask-shaped (135 » X I02 ), with one pair of — 
setae. Anal shield (72 » x 87 ») with paranal setae about 24 p, postanal seta about | 
15 ». 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 ») of leg segments : 


I II lll 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 


HaBitAt: 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 
U.S.A. 


Hypoaspis oblonga (Halbert) 


Laelaps (Hypoaspis) oblongus Halbert, 1915, Proc. R. Irish 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 »; II, 162 7; movable digit 68 », biden- 
tate ; fixed digit with about six teeth (Text-fig. 468) ; pilus dentilis and dorsal 
short, setiform. Four pairs of gnathosomal setae with c.s. about 42 » apart, hyp. 
about 40 » apart. Deutosternum with six transverse rows of denticles (4—6 denticle 
per row). Corniculi horn-like, 42 » long ; internal malae fimbriate (Text-fig. 468) 


THE BRITISH DERMANYSSIDAE: (ACA RT) 


Fic. 46. Hypoaspis oblonga (Halbert), female; dorsum of idiosoma (a) ; 
tectum capituli (c); apotele of pedipalp (D); venter of gnathosoma (E); venter of 
idiosoma (F). 


chelicera (B) ; 


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 p x 
132 ») reticulated, with three pairs of setae and two pairs of pores ; metasternals free 
with associated pores. Between st1, 70 »; between sti and st3, 120 p. Genito- 
ventral shield (204 » * 144 m») expanded behind coxae IV and bearing a pair of 
genital and a pair of opisthogastric setae. Anal shield (100 » X I00 »; maximum 
width 156 «); paranal setae 36 » 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, 
free. 

Legs with normal chaetotaxy, except genu IV which has only one antero-lateral 
seta (I—}, §—1). 


Hasirat : Under bark of decayed trees and in moss, in Ireland, Scotland, England 
(Northumberland) and Europe. 


Genus PSEUDOPARASITUS Oudemans 


Pseudoparasitus Oudemans, 1902, Tijdschr. Ent, 45 : 29. 

Hoplolaelaps Berlese, 1903, Zool. Anz, 27: 14. 

Laelapsoides Willmann, 1952, Veroff. 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 with 
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 II 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 meridionalis appears to be lost. 


Pseudoparasitus centralis Berlese 


Pseudoparasitus centralis Berlese, 1921, Redia 14: 167. 


Female: Chelicera with segment I, 66 »; II, 162 1»; 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 pw apart; hyp.2 about 42 wm 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 » x 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 ». Sternal shield (126 w xX 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 stI, 54 »; between sf 
and st3, 99 ». Genital shield (180 » xX 138 yp) flask-shaped, expanded behind coxae 
IV, with truncate posterior margin, bearing five pairs of setae. Between gen., 120 p ; 
between /v2, 114 pw. Anal shield (96 » X I05 p) with a straight anterior margin 
measuring 180 ». Paranal setae 30 p, postanal seta 10 pw. 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 p) 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 pu, 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 


G. OWEN EVANS & W. M. TILL 


Fic. 47. Pseudoparasitus centralis (Berlese). A. Dorsum of idiosoma of female ; B. venter 
of idiosoma of female; c. chelicera of male; D. chelicera of female; &. venter of idio- 


soma of male: F. venter of gnathosoma of female. 


SRE BRETVTISH DEKMANYSSIDAE'(ACART) 227 


Dorsal shield (495 » X 300 u) with 39 pairs of setae distributed asinfemale. Trito- 
sternum with base 21 p, laciniae 63 ». Holoventral shield (390 » x 99 p) 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, 45 mu; between sti and st3, 90 pw. Paranal 
setae 24 p, postanal seta 12 w. 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 fv inflated basally ; 
femur IV with seta pd, inflated. Length/width (in yp) of leg segments : 


I II III IV 
genu 69/27 66/45 42/30 60/36 
tibia AEG | 54/42 40/32 60/40 


tarsus 120/22 84/30 84/24 114/30 


HABITAT: Free-living in litter under deciduous trees etc. in Britain and Europe. 


Pseudoparasitus dentatus (Halbert) 
Laelaps dentatus Halbert, 1920, Proc. R. Irish Acad. 25: 123. 


Female: Chelicera with segment I, 54 w 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 wapart, hyp.2 42-45 wapart. 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 p) with 39 pairs of setae distributed as in Text- 
fig. 484. Surface of shield reticulate ; no hypertrichy. 

Tritosternum with base 33 p, laciniae8I pu. Sternalshield (150-180 w X 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 st1, 66-72 » ; between sti and 3/3, 
108-132 pw. Genital shield (192-216 mw X 105 yp) flask-shaped with four pairs of 
setae ; between gen., 102 4; between /v2, 84 ». Anal shield (60-68 » X 75-90 yz) 
with maximum width (123-170 pw) at anterior margin; paranal setae 30-40 yp, 
postanal seta 18-30 p. 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,, avy, pv1, Pie, pl, and mv all stout 
basally with fine tips. Length/width (in ) of leg segments : 


228 G. OWEN EVANS & W. M. TILL 


I Il Ill 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 


HasiraT: 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, Redta 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 (B) 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 
(zi and 23 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 


= 


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 ‘ ‘ 2 
2 Spermathecal ducts conspicuous (Text-fig. 49c) ; peritrematal shield fused with the 
podal and genito-ventro-anal shields (Text-fig. 49B and D) ; with 38 pairs (23 
absent) of relatively short dorsal setae (Text-fig. 494). 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. 528) ; 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 D) ; genu III with seta pv normal ; 
dorsal shield with 37 or 38 pairs of setae : : : : : : : 2 
2 Paravertical setae absent . ‘ : : O. sellnicki Bregetova & Koroleva (p.233 ) 
- Paravertical setae present. ; ; : : : . : O. veneta (Berlese) 


230 G. OWEN EVANS & W. M. TILL 


= 
Fic. 49. Ololaelaps veneta (Berlese), female; dorsum of idiosoma (A); perit mata 


podal shields in region of coxa IV (s); “ spermathecal ducts " (c); genito-ventro-al 
shield (p). : 


DEE SELL ISH DERMANYSSI DAE) (AC ART) 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, Parazit. Sb. 22 : 81, figs. (syn. nov.). 


Female: Chelicerae with segment I, 75 w; II, 150 w; movable digit 87 yw, bi- 
dentate ; fixed digit tridentate; pilus dentilis short, setiform. Four pairs of 
gnathosomal setae with c.s. about 57 w apart; hyp.2 about 48 p» apart. Deuto- 
sternum with six transverse rows of denticles (3-6 denticles per row) ; corniculi 45 pu 
long ; internal malae as in O. selinicki. 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 pu) with 38 pairs of relatively short, simple 
setae distributed as in Text-fig. 494, seta z3 being absent. An unpaired accessory 
seta is present between the J series. Surface of shield granular with sculptured areas 
in the podonotal region. 

Tritosternum with base 30 yp, laciniae 81 wp. Sterno-metasternal shield (102 p x 
132 ») with four pairs of setae and three pairs of pores. Between sf1, 66 1; between 
sti and st3, 87 ». Genito-ventro-anal shield (270 w 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 72/50 54/38 75/40 
tibia 87/32 60/45 54/36 75/38 


tarsus 153/30 102/34 105/30 135/39 


Male: Described and figured by Bregetova & Koroleva (1964). 
Hapirat: 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. (58), 6 (14) : 
69, fig. 48. 

Ololaelaps selinicki Bregetova, 1964, Parazit. Sb. 22: 77, figs. (nom. nov. pro Ololaelaps haemis- 
phaericus Sellnick, 1940 non Koch, 18309). 


Female : Chelicera with segment I, 102 »; II, 204 »; movable digit 87 », biden- 
tate; fixed digit tridentate ; pilus dentilis short, setiform (Text-fig. 50c). Four 


232 


G. OWEN EVANS & W. M. TILL 


D 


Fic. 50. Ololaelaps sellnicki Bregetova, female; dorsum of idiosoma (A); venter of 


gnathosoma (B); chelicera (c); tectum capituli (D) ; 
venter of idiosoma (F). 


‘‘spermathecal ducts ’’ (E) ; 


THE BRITISH DERMANYSSIDAE (ACARI) 233 


pairs of gnathosomal setae with c.s. about 60 pw apart, hyp.2 about 48 pw apart. 
Deutosternum with six transverse rows of denticles (I—-5 per row) ; corniculi 60 
long ; internal malae as in Text-fig. 508. Tectum capituli with anterior margin very 
finely denticulate (Text-fig. 50D). Salivary styli conspicuous. Pedipalp (2—-5-6-14) 
with three-tined apotele. 

Dorsal shield (636 » xX 480 pz) 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 J series. Surface of shield granular with sculptured areas in the 
podonotal region. 

Tritosternum with base 24 yp, laciniae 66 ». Sterno-metasternal shield (102 p x 
132 ») with four pairs of setae and three pairs of pores. Between st1,60 w; between 
sti and st3, 87 ». Genito-ventro-anal shield (264 » x 264 ») with four pairs of setae 
in the preanal region. Paranal setae 36 pw, postanal seta 21 pw. 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! II III ry 
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 w; movable digit 63 p (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 pw apart, hyp.2 about 45 pw apart. Deuto- 
sternum with six transverse rows of denticles (2-5 per row) ; corniculi 48 yp long ; 
internal malae fringed (Text-fig. 51c). Tectum, salivary styli and pedipalps as in 
female. 

Dorsal shield (516 » xX 360 p») with 37 pairs of setae and one unpaired accessory 
seta. Tritosternum with base 15 p, laciniae 57 pw. MHoloventral 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 9 wp. Chaetotaxy of opisthogaster as in Text-fig. 51E. 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 : 


234 


G. OWEN EVANS & W. M. TILL 


THE BRITISH DERMANYSSIDAE (ACARI) 235 


I II ELT IV 
genu 75/33 66/45 48/30 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 p. 
Four pairs of gnathosomal setae with c.s. about 60 pw apart, hyp.2 about 48 p apart. 
Deutosternum with six transverse rows of denticles (3-6 per row) ; corniculi 40 pu 
long. Internal malae and pedipalps as in female. 

Dorsal shield (624 » X 396 pw) with lateral incisions, bearing 34 pairs of setae ; 
setae zI and 23 absent; setae s3, sod and humeral seta on cuticle (Text-fig. 514A). 
Opisthonotal region with one accessory seta; five pairs of Fk setae on the cuticle. 
Tritosternum with base 20 yp, laciniae 50 pw. Sternal shield (246 w Xx 105 p») with 
four pairs of setae and three pairs of pores. Between sf1, 60 »; between sf1 and s/3, 
102 ». Anal shield (96 » X 110 ») with maximum width 162 » ; paranal setae 27 pw; 
postanal seta 18 ». Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 
51F. 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 fv stout basally, 
tapering to long, fine points. Length/width (in ») of leg segments : 


I II Ill IV 
genu 75/42 70/50 50/42 7245 
tibia 87 /36 60/45 48/40 72/42 


tarsus 150/33 93/40, "105/53 144/40 


HABITAT: Found in moss and decaying organic matter in Britain, Europe and 
Iceland. 


Note: Bregetova & Koroleva (1964) have figured the dorsum of O. sellnickhi 
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. (5B), 6 (14): 69, fig. 49; 
Bregetova & Koroleva, 1964, Parazit. Sb. 22 : 70, figs. 

Ololaelaps confinis Berlese, 1904, Redia 1 : 261. 


Female : Chelicera with segment I, 90 » ; 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 selinickt Bregetova. A. Dorsum of idiosoma of deutonymph; B. 
chelicera of male, internal view ; c. deutosternum and hypostome of male; bD. chelicera 
of male, dorsal view; E. venter of idiosoma of male; F. venter of idiosoma of deuto- 
nymph. 

ZOOL. 14, 5. 16 


236 


G. OWEN EVANS & W. M. TILL 


Fic. 52. Ololaelaps placentula (Berlese). A. Dorsum of idiosoma of female ; 8. chelicera 
male; c. genu III of male; p. tarsus II of male; E. venter of idiosoma of male ; 1 


venter of idiosoma of female. 


PTE BSiPish DERMANYSSiDAL (ACARI) 237 


hyp.2 about 57 mw apart. Deutosternum with six transverse rows each of many 
denticles ; corniculi 63 »; internal malae and tectum captituli as in I. sellnicki. 
Salivary styli conspicuous, extending almost to tips of corniculi. Pedipalp 
(2-5-6-14) with three-tined apotele. 

Dorsal shield (approximately 732 u X 540 ») with 39 pairs of simple setae distri- 
buted as in Text-fig. 524A; Surface of shield granular, with faint reticulations ; no 
hypertrichy. 

Tritosternum with base 27 yp, laciniae 84 w. Sternal shield (80 » xX 132 p») with 
four pairs of setae and three pairs of pores. Between sf1, 80 » ; between sf and s/3, 
84 p». Genito-ventro-anal shield (288 » Xx 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 pw, postanal seta 30 w. 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 yp) of leg segments : 


I I Ill IV 
genus (93/42 75/54 63/45 75/45 
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 w; movable digit 75 pw (123 p 
with spermadactyl), unidentate ; margins of spermadactyl groove smooth (Text-fig. 
52B) ; fixed digit unidentate, pilus dentilisshort, setiform. Fourpairsofgnathosomal 
setae with c.s. about 60 pw 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 » xX 460 ») 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 st1, 70 w; between sii and st3, 93 w. Paranal setae 33 p, postanal 
seta 24 w. 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 pv, 
inflated basally, tapering to a long, fine point (Text-fig. 52D). Length/width (in ,) 
of leg segments : 


I II i IV 
genu 75/36 78/50 60/42 72/42 
tibia 84/33 66/45 57/40 75/40 


tarsus. 150/30 132/35 «117/34 144/38 


238 G. OWEN EVANS & W. M. TILL 


Immature stages: The larva, protonymph and deutonymph of this species are 
described and figured by Bregetova & Koroleva (1964). 


Hapitat: 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 (Brevisterna). 

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 accessory setae in some species of Haemogamasus; 
shield reduced and bearing only two pairs of setae and two pairs of pores in Brevt- 
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 (Eulaelaps) 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 IV (2—?, %—2) in all species’ 
and rarely (H. hirsutosimilis) 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 
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. Bie lish DE RMANYSSIDAE)( ACARI) 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, Acar., Myr. Scorp. Ital. fasc. 52, nos. 2 and Io. 
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- 


240 G. OWEN EVANS & W. M. TILL 


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—7, §—2) and tibia IV © 
(2—?, ?—2) in all species and femur II (2—}—1) and femur III (1——1z) in H., 
hirsutosimilits. 

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 Liponyssus 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 nomenclature 
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 anc 
the suprageneric categories based on the name Liponyssus. 


KEY TO FEMALES 


1! Sternal shield with the normal three pairs of setae only . 

Sternal shield with accessory setae which are usually shorter than the primary 
sternal setae and vary in size, number and position on the shield . 

2 Posterior margin of the sternal shield deeply incised to a level midway between the 
first and second pairs of sternal setae (Text-fig. 608). 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. 59B). Anal 
shield with several accessory setae . ; 

3 First pair of sternal setae and hypostomal setae barbed. Dorsal shield, especially. is 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. 
59E). . ; 4 : : : ; ‘ : : H. nidi Michael (p. 254) 


THE SSE Lish  DERMANYSSIDAE «ACA RTI) 241 


Sternal, hypostomal and dorsal setae smooth. Tectum capituli with gently rounded 

margin of simple and multiple fimbriae. Pilus dentilis short, setiform (Text-figs. 

57A andc) . , 4 
Postero-ventral margin of the idiosoma with at least one pair of setae conspicuously 

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. 58p. 

Associated with Talpa (Insectivora) : , : H. horridus Michael (p. 252) 
Postero-ventral margin of the idiosoma with setae 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. 56F. Associated with Muridae 

(Rodentia) . : , : ‘ H. arvicolarum (Berlese) ( p. 250) 
Femur II with 12 setae (2—{—1), fend 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 II] 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 both =e “cies ee aed. eon ote 

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 . , , H. ambulans (Thorell) (p. 244) 
Chelae edentate ; Paleirodauter ne 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 . : , : i .  H. hirsutus Berlese (p. 242) 


KEy To MALES 


Sternal region of holoventral shield with no accessory setae anterior to the level of the 
second pair of pores ‘ 3 . : 2 

Holoventral shield with eeeor setae dietributed over ne pdeire Sree : 5 

First pair of sternal setae smooth (Text-fig. 58c). Tectum capituli with sett 
rounded margin of simple and multiple fimbriae. Chelicera with dorsal seta dis- 


tinctly distal to base of fixed digit ‘ : 3 . : : : ' 2 
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. 56E). 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. 578). 

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) . 4 H. pontiger (Berlese) (p. 259) 

First pair of sternal setae barbed : ; : ; : . H.ambulans (Thorell) 

Allsternalsetae smooth . ‘ : : : : ‘ : : é 6 


G. OWEN EVANS & W. M. TILL 


we 
— 
te 


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. 558). 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 1} 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 » ; II, 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 p» 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 yw) densely covered with simple setae of variable 
length (Text-fig. 53A). Surface of shield granular with some weakly defined reticula- 
tions. 

Tritosternum with base 60 yp, laciniae 228 »; presternal area reticulate, with 
minute spines. Sternal shield (174 uw x 234 m) 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 pang on the posterior margin of the shield ; metasternal 
setae free. Between st1, 150 1; between sf1 and st3, 180 yp. Genito-venteal shield 
relatively slender with numerous accessory setae distributed over entire shield. Anal 
shield (168 » & 96 «) with the usual three setae and five accessories ; paranal setae 
123 w, postanal seta 108 » long. Chaetotaxy and sclerotization of opisthogaster as 
in Text-fig. 538. Peritreme extends to posterior margin of coxa II; peritremata 
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 


Pic. 53. Haemogamasus hirsutus Berlese. A. Dorsum of idiosoma of female; 8B. venter of 
idiosoma of female; c. venter of idiosoma of male; pb. chelicera of female; £. chelicera — 
of male. 


243 


THE BRITISH DERMANYSSIDAE (ACARI) 


a SSS 


244 G. OWEN EVANS & W. M. TILL 


leg setae are smooth. Distal margins of most leg segments denticulate ventrally. 
Length/width (in ») of leg segments : 


I Il 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 pv (87 p 
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 «apart. Deutosternum with 13 transverse rows of denticles ; corniculi 54 » 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 pw) as in female. Tritosternum with base 50 
laciniae 165 ». Holoventral shield (840 » x 198 pw) 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 p» long. Peri- 
tremes and peritrematal shields as in female. 

Chaetotaxy of legs as in female ; relatively stouter setae on femur II (ventre 
genu and tibia II (av) and tarsus II (mv and av,)._ Length/width (in ,) of leg 
ments : 


I II 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 Lir = 
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. Forh. 2: 164. 

Hypoaspis ambulans; Tragardh, 1904, Fauna Arctica 4 : 33, figs. 

Haemogamasus ambulans: Bregetova, 1953, Parazit. Sb. 15: 316; and 1956, Opred. Faun 
SSSR. 61: 147, figs. 

Haemogamasus alaskensis Ewing, 1925, Proc. biol. Soc. Wash. 38 : 138. 
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. 1, fig. 2. 

Haemogamasus paviovskyi Bregetova, 1949, Parazit. Sb. 11: 179, figs. 


Female; Chelicera with segment I, 105 »; II, 160 »; movable digit 54 | 
bidentate ; fixed digit tridentate; pilus dentilis setiform ; dorsal seta long @ 


THE BRITISH DERMANYSSIDAE (ACART) 


Fic. 54. Haemogamasus ambulans (Thorell), female ; dorsum of idiosoma (A) ; venter of 


chelicera (D). 


idiosoma (B) ; venter of gnathosoma (c) ; 


246 G. OWEN EVANS & W. M. TILL 


stout (Text-fig. 54D). Four pairs of gnathosomal setae with c.s. about 105 p» apart, 
hyp.2 about 75 » apart; Ayp.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 » x 612 ») densely covered with setae mostly of uniform length 
(Text-fig. 54A) ; some anterior and marginal setae barbed. Surface of shield granu- | 
lar, reticulations most distinct in opisthonotal area. 

Tritosternum with base 57 p», laciniae 150 ». Sternal shield (132 » x 190 ») with 
the usual three pairs of sternal setae and about 14 accessories, and with three pairs of 
pores ; metasternal setae free. Between sf1, 150 »; between sf1 and st3, 120 p; 
stt barbed. Genito-ventral shield (276 » x 120 ») with a pair of genital setae and 
about 36 accessories distributed chiefly on the posterior part of the shield. Anal 
shield (123 » X 130 ») 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. 54B. Peritreme extends to anterior third of coxa IT; 
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,, pd,, avy, pry, al,, 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 III IV 
genu 156/54 120/60 114/54 168/50 
tibia 168/48 120/57 120/48 180/48 


tarsus 228/42 210/45 264/42 372/42 
Hasitat: Found in the nest of Riparia riparia (Linn.) in Gloucestershire, Britain, 
Recorded in the literature from the nests of birds, from insectivores, bats, sables, anc 


a variety of rodents in Germany, the U.S.S.R., Greenland, Spitsbergen and North 
America. 


Haemogamasus hirsutosimilis Willmann 


Haemogamasus hirsutosimilis Willmann, 1952, Z. Parasitenk. 15 : 403, figs. 
Bregetova, 1956, Opred. Faune SSSR. 61: 141, figs. 


Female: Chelicera with segment I, 84 »; II, 156 »; movable digit 60 p, biden- 
tate ; fixed digit with two large and two very small teeth ; pilus dentilis short, sett 
form, dorsal seta long (Text-fig. 558), Four pairs of simple gnathosomal setae wit 


4 


np. chelicera of female; c. venter of idiosoma of female; pb. venter of idiosoma of male; 
&. chelicera of male; F. deutosternum and hypostome of female. 


247 


THE BRITISH DERMANYSSIDAE (ACART) 


248 G. OWEN EVANS & W. M. TILL 


c.s. about 96 » apart, hyp.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. 55A). Surface of shield granular. 

Tritosternum with base barbed, 50 » long ; laciniae 150 ». Sternal shield (186 » 
216 «) with the usual sternal setae, numerous accessories, and three pairs of pores ; 
metasternal setae free. Between sf1, 60 »; between s/1 and st3, 174 p. Genito- 
ventral shield with numerous setae distributed over its entire surface. Anal shield 
(126 «% X go ») with the usual three anal setae and five accessories ; paranal setae 
II0 «; postanal seta 117 ». 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 ,) of leg segments : 


I II 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 »; fixed digit short and broac 
with an anterior projection; movable digit and spermadactyl strongly curvec 
(Text-fig. 55£). Four pairs of gnathosomal setae with c.s. about 85 apart, hyp. 
about 63 » apart. Corniculi 42 » long ; other gnathosomal features as in female. 

Dorsal shield (780 » x 438 ») asin female. Tritosternum with base 33 p, laciniae 
105 #; presternal area as in female. Holoventral shield 648 p» 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 ; 
venter of idiosoma of female; c. corniculus of female, lateral view; pb. venter of idio 
soma of male; E. femur II of male; F. deutosternum of female. 


249 


THE BRITISH DERMANYSSIDAE (A CART) 


Y 
4 
, 
Jo 


250 G. OWEN EVANS & W. M. TILL 


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 (fvg-5), genu II (av), tibia II (av) and tarsus II (mv); seta av, 
on tarsus I] is very stout but pointed. Seta pd, 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 II Ill 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 


Hapirat: 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 horridus var. arvicolarum Berlese, 1920, Redia 14: 166. 
Haemogamasus arvicolarum: 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 » apart, hyp.2 about 93 » apart. Deutosternum with 17 transverse 
rows of denticles (1-4 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 » * 840 pv) with a dense covering of subequal setae (Text-fig. 
560A). 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 », 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 wi 
associated pores. Between sf1, 90 »; between sf1 and st3, 168 ». Genital shiel 
with about 44 setae distributed mainly on the posterior part. Anal shield (258 » x 
135 ») with maximum width 186 yp, 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. Peritre 
extends to posterior margin of coxa II ; peritrematal shield fused anteriorly with 
dorsal shield and posteriorly with podal shield of coxa IV. 


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 p) of leg segments : 


I Id OE IV 
genu 207 |'72 168/102 150/84 192/78 
tibia 222/66 168 /go 168/72 240/72 


tarsus 306/60 240/66 330/66 456/66 


Male: Chelicera with segment I, 84 »; I, 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. 57p). Four pairs 
of simple gnathosomal setae with c.s. about 105 uw apart ; Ayp.2 about 93 p apart. 
Deutosternum with 14 transverse rows of denticles (I-3 per row) ; corniculi 57 p 
long ; internal malae, tectum and pedipalps as in female. 

Dorsal shield (1,092 » X 600 ») as in female. Tritosternum with spine on each 
side of base ; with base 60 yp, laciniae 210 w. 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 


=a = = 
. 


‘e 


Fic. 57. a-B. Haemogamasus horvidus Michael; chelicera of female (A) and male (s). 
C-D. Haemogamasus arvicolarum Berlese ; chelicera of female (c) and male (p). 
ZOOL. TA; 5< 


saa 


252 G. OWEN EVANS & W. M. TILL 


and may extend to the level of the metasternals (Text-fig. 56D). Between sér, 
87 «; between sf and st3, 123 w. Paranal setae 90 »; postanal seta 80 pw. Peri- 
treme extends approximately to posterior border of coxa II; peritrematal shiel¢ 
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, 
56E). Length/width (in ») of leg segments : 


I II 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 


HABITAT: 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 py, 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 setas 
with c.s. about 126 » apart; Ayp.2 about 100 pw apart. Deutosternum with 1 
transverse rows of denticles (1-5 denticles per row) ; corniculi broad, membranous 
about 45 » long; internal malae fringed, comprising slender inner and _ broac 
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 » x 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 | 
(234 » * 240 p) with three pairs of setae and two pairs of pores ; metasternal setat 
free with associated pores. Between st1, 80 » ; between str and st3, 222 yp. 
ventral shield with about 70 setae. Anal shield (234 » x 138 »; maximum \ 
180 ») with the usual three anal setae and about 14 accessories. Paranal setae 132 # 
postanal seta 150 plong. Chaetotaxy and sclerotization of opisthogaster as in Text 
fig. 588; at least one pair of setae at posterior margin considerably longer than the 


Cs 
So 
Tie 


—— ee  t— ee 


Fic. 58. Haemogamasus horridus Michael. a. Dorsum of idiosoma of female; B. venter 
of idiosoma of female; c. venter of idiosoma of male; pb. venter of gnathosoma of female. 


253 


THE BRITISH DERMANYSSIDAE (ACART) 


254 G. OWEN EVANS & W. M. TILL 


neighbouring setae ; some setae with fine, sparse barbs. Peritreme extends to 
anterior margin of coxa II]; anterior tip of peritrematal shield fused with dorsal 
shield at level of 71 ; posterior part fused with podal shields behind coxae IV. 

Chaetotaxy of legs normal except genu IV (2—{, §—2) and tibia IV (2—#, #—2). 
Leg setae apparently simple ; distal margins of most leg segments serrated ventrally. 
Length/width (in ,) of leg segments : 


I II III lV 
genu 252/75 192/108 168/90 222/90 
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 mw apart, hyp.2 about 84 pw 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 » xX 600 pw) as in female. Tritosternum with base 48 uz, 
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 s/1, 80 » ; between sti and s#3, 168 p. 
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 III ; 
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 III 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 U.S.S.R. 


Haemogamasus nidi Michael 


Haemogamasus nidi Michael, 1892, Trans. 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 (4) : 435, 
figs. 116-122. 


Female: Chelicera with segment I, 90 »; II, 147 »; movable digit 48 pn, bi- 
dentate ; fixed digit bidentate ; pilus dentilis inflated basally; dorsal seta long and 


a a : 


THE BRITISH DERMANYSSIDAE (ACARI) 255 


stout (Text-fig. 59E). Four pairs of barbed gnathosomal setae with c.s. about 93 p» 
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 styli conspicuous, extending 
beyond tips of corniculi. Tectum capituli slender, pointed, fimbriate. Pedipalp 
(2-5—6-14) with two-tined apotele. Palptrochanter with both setae barbed. 

Dorsal shield (984 » X 576 ») 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 yp, laciniae 150 w. Sternal shield (108 » x 168 yp) with 
three pairs of setae and three pairs of pores ; metasternal setae free. Between s/1, 
120 »; between sfx and si3, 120 »; sti barbed. Genito-ventral shield bears 
numerous setae distributed mostly on posterior two-thirds of shield. Anal shield 
(110 » X 105 p) with five or six accessories in addition to the usual three anal setae ; 
paranal setae 66 w; postanal seta 84 pw. Chaetotaxy and sclerotization of opistho- 
gaster as in Text-fig. 59B. 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 Ti Ill IV 
genu 144/54 114/72 GO (60, .. 4138/57 
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 w. Cheliceral digits and sperma- 
dactyl as in Text-fig. 59F. Four pairs of gnathosomal setae with c.s. about 84 pu 
apart, hyp.2 about 66 w 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 p, presternal area as in female. Holoventral shield (624 » x 150 y) with the 
usual sternal and metasternal setae, and with numerous accessory setae distributed 
over almost entire shield and extending anteriorly to between séz and st3 (Text-fig. 
59C). Between sti, 90 »; between sfI and sf3, 110 »; sti barbed. Par- and post- 
anal setae subequal, about 60 pw 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 asin female. Stout setae present on femur 
II (av), genu II (av), tibia II (av) and tarsus II (av, and mv). Length/width (in yp) 
of leg segments : 


256 


G. OWEN EVANS & W. M. TILL 


oe on 


LE 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 


HasiTaT: 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) pontiger Berlese, 1904, Redia 1: 260. 
Bregetova, 1956, Opred. Faune SSSR. 61 : 1209, figs. 

Groschaftella pontiger: 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 mawsont 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 w; movable digit 63 pn, 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 wu apart, hyp.2 
about 66m apart; hyp.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 w long ; laciniae 126 pw. Presternal area with 
small spines on the reticulations. Sternal shield (42 » x 160 p) deeply concave 
posteriorly, with three pairs of setae and three pairs of pores ; metasternal setae free. 
Between sf1, 117 » ; between sti and st3, 123 w; st1 barbed. Genito-ventral shield 
(300 » X 114 p) 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 ») with two small anterior protuberances ; paranalsetae 60 » ; postanal 
seta 78 pw. 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; pb. deutosternum of female; E. 
chelicera of female; F. chelicera of male. 


Fic. 60. Haemogamasus pontiger (Berlese). A. Dorsum of idiosoma of female; B. venter — 
of idiosoma of female ; c. chelicera of female ; p. venter of idiosoma of male; £. chelicera — 
of male; F. apotele of pedipalp; G. venter of deutosternum and hypostome of male. — 


fie BVatsH DERMANYSSILDAE: (ACARD) 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 pd, 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 yp) of leg segments : 


I II Ill IV 
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 p» 
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 wapart, 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 
102 p. Presternal area asin 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 sf1, 90 »; between sti 
and s#3, 110 »; sti barbed. Paranal setae 36 w; postanal seta 48 pw. Peritreme 
extends a little beyond posterior margin of coxa II ; peritrematal shield free anteriorly 
and posteriorly, although anterior tip hes very close to dorsal shield. 

Chaetotaxy of legs as in female ; short, stout, blade-like setae on femur III (v) 
and genu III (fv). Length/width (in ») of leg segments : 


I II III yi 
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 


HaBitaT: 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, Trans. 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 LV. 

Segmental chaetotaxy of legs normal except genu IV (2—#, #—2). Coxa II witha 
small anterior spine. Ambulacra with claws sometimes strongly reduced on leg I. 


KEY TO FEMALES 


1 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 shields Sore 61F). Tarsus II with some 
spine-like setae (Text-fig. 61¢) ‘ ; 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) 


KEY TO MALES 


1 Tarsus II with setiform and — setae ; femur II without sclerotized pro- 


tuberance ventrally 3 : ; E. stabularis Koch (p. 262) | 
~ Tarsus II with setiform and stout, spur- like aie ; femur II with conspicuous pro- 
tuberance ventrally distal to fissure. ‘ , ; E, novus Vitzthum (p. 265) 


Eulaelaps stabularis (iKoch)*® 


Gamasus slabularis Koch, 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, Trans. Linn. Soc. Lond. Zool. 5 : 315, figs. 
Eulaelaps arcualis (Koch) Tragardh, 1912, Arch. zool. exp. gen. 48: 577, figs. 90-93. 
Eulaclaps oudemansi Turk, 1945, Parasitology 36 : 137. 


* According to Johnston (1959) Laelaps pedalis Banks, 1909 and L. propheticus Banks, 1909 are also 
synonyms of E. stabularis. 


THE BRITISH DERMANYSSIDAE (ACARI) 


Eee 
=} | (2b) 
fa pate) 
~ — 
£3 
5 0 
JD) 
> 
o 
Li@ esc 
c= fab) 
Aen 
= 
ml 
e) 


able digit of chelicera 
of female. 


Dorsum of idiosoma of female ; 


egion 


s II of female; D. mov 


F. opisthogastric r 


f female ; 
of female ; 


Fic. 61. Eulaelaps stabularis (Koch). A. 
oI female; c. tarsu 


262 G. OWEN EVANS & W. M. TILL 


Female: Chelicera with segment I, 90 »; II, 180 «; movable digit 75 », bidentate; 
fixed digit tridentate, pilus dentilis short, setiform ; dorsal seta stout (Text-fig. 618). 
Four pairs of gnathosomal setae with c.s. about 93 » apart, hyp.2 about 66 p» 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 » X 612 ») shows marked hypertrichy ; setae simple, mostly 
subequal, shorter towards anterior margin (Text-fig. 61A). Surface of shield 
granular and reticulate. 

Tritosternum with base 45 p, laciniae 160 »; presternal area well sclerotized. 
Sternal shield (156 » x 186 ») with three pairs of setae and two pairs of pores ; 
metasternal setae free with associated pores. Between s/1, 96 «; between sti and 
sf3, 138 pw. Genito-ventral shield (408 » x 180 ,») with lateral incisions behind the 
genital setae ; widely expanded behind coxae IV with maximum width 456 1; 
bearing a pair of genital setae and about 47-52 opisthogastric setae (Text-fig. 61F). 
Anal shield (78 » & 114 ») with maximum width at anterior margin 222 »; paranal 
setae 63 pw, postanal seta 78-84 » long. Metapodal shields very large, subtriangular. 
Peritreme extends to middle of coxa 1 ; anterior tip of peritrematal shield fused with 
dorsal shield at level of seta z1, posterior part free. 

Chaetotaxy of legs differs from normal pattern in having two postero-lateral setae 
on genu 1V (2—j;, §—2). Tarsus II with some spine-like setae (Text-fig. 61C). 
Some leg segments with distal margins serrate. Length/width (in ,) of leg segments 


I II Ill IV 
genu 138/60 120/78 96/60 132/54 
tibia 150/48 114/72 96/54 138/54 


tarsus 228/42 156/54 180/48 270/48 


Male; Chelicera with segment I, 48 »; II, 130 »; movable digit 63 », with a 
large, subterminal, tooth-like projection (Text-fig. 61D); spermadactyl slightly 
surpasses tip of movable digit ; fixed digit edentate, pilus dentilis short, setiform. 
four pairs of gnathosomal setae with c.s. about 54 » apart, Ayp.2 about 50 » apart. 
Deutosternum with nine transverse rows of denticles (4-8 per row) ; corniculi 50 p 
long. Internal malae, tectum, salivary styli and pedipalps as in female. 

Dorsal shield (582 » * 348 ») asin female. Tritosternum with base 30 yp, laciniae 
108 ». Holoventral shield (468 » * 126 ») granular and reticulate ; with four pairs 
of setae and three pairs of pores in the sternal region ; a pair of genitals and 36-43 
opisthogastric setae in the genito-ventral region. Between st1, 66 »; between sfI 
and sf3, 102 4. Paranal setae 48 p, postanal seta 60 » long. Peritreme extends to 
anterior margin of coxa II; anterior part of peritrematal shield fused with dorsal 
shield, posterior part free, 


. 
’ 


foe BRITISH DERMANYSSIDAE (ACARI) 263 


Chaetotaxy of legs asin female. Femur II with seta v, relatively stout, tapering. 
Setae av, and mv on tarsus II, seta av on genu and tibia II, stout basally, tapering to 
fine points. Length/width (in ,) of leg segments : 


I II III IV 
genu 96/39 75/48 60/36 78/30 
tibia 100/36 72/45 57/36 84/30 


tarsus 150/30 96/36 105/30 140/27 


HABITAT: Foundin Britain and Europe on the bodies and in the nests of rodents 
and insectivores and in the nests of the Wheatear, Oenanthe oenanthe Linn. and the 
Sandmartin, Riparia riparia (Linn.). 


Eulaelaps novus Vitzthum 


Eulaelaps novus Vitzthum, 1925, Jen. Z. Naturw. 62 : 165, fig. 
Samsinak, 1960, Acta Soc. ent. Cech. 57 : 276, figs. 
Eulaelaps kolpakovae Bregetova, 1950, Parazit. Sb. 12 : 13, fig. (Syn. nov.). 


Female : Chelicera with segment I, 84»; II, 186 1»; movable digit 70 p, biden- 
tate ; fixed digit as in Text-fig. 62c ; pilus dentilis and dorsal seta short, setiform. 
Four pairs of gnathosomal setae with c.s. about 72 » apart, hyp. 2 about 66 p apart. 
Deutosternum with eleven transverse rows of denticles (5-7 per row) ; corniculi 50 u 
long ; internal malae elongate, fringed (Text-fig. 62F). Tectum capituli with an- 
terior and lateral margins deeply denticulate (Text-fig. 62B). Salivary styli con- 
spicuous, reaching tips of corniculi. Pedipalp (2—-5-6-14) with two-tined apotele ; 
palptrochanter with sensory organ ventrally. 

Dorsal shield (936 » X 588 ,) hypertrichous, surface of shield granular and reti- 
culate. Setae simple, mostly subequal, slightly longer in the opisthonotal region. 

Tritosternum with base 63 p, laciniae 150 pw. Sternal shield (120 w Xx 168 y») with 
three pairs of setae and two pairs of pores; metasternal setae free with associated 
pores on medial margin ‘of setal bases. Between sti, 102 »; between sf and sf3, 
132 p. Genito-ventral shield (324 » x 144 ») expanded posteriorly with maximum 
width 300 »; bearing a pair of genital and about 26 opisthogastric setae. Anal 
shield (100 » X 132 ») with paranal setae 75 p, postanal seta 50 wlong. Chaetotaxy 
and sclerotization of opisthogaster as in Text-fig. 624 ; metapodal shields large, sub- 
triangular. Peritreme extends to anterior margin of coxa II ; anterior part of peri- 
trematal shield fused with dorsal shield, posterior part free, its pore apparently 
situated at the bottom of a wide cavity. 

Chaetotaxy of legs differs from the normal pattern in that genu IV has two postero- 
lateral setae (2—?, 2—2) and tarsi II-IV lack setae ad, and pd,. Tarsus II (Text- 
fig. 62E) bears several stout, spur-like setae (av,, al,, ply, aV2, Pv2, mv). Ambulacrum 
I without claws (Text-fig. 62G).  Length/width (in ») of leg segments : 


204 


G. OWEN EVANS & W. M. TILL 


G 


a. Venter of idiosoma of female; 8B. tectum capituli — 
r. tarsus Ilof female; F. venter 
al shield of male. 


Fic. 62. Eulaelaps novus Vitzthum. 
of female: c.cheliceraof female ; p.chelicera of male ; 
of gnathosoma of female; G.ambulacrum of leg lof female; H. holoventr 


PE BRITISH DERMANYSSIDAE (ACART) 265 


I II III IV 
genu 126/54 426/72 90/54 132/52 
tibia 138/50 108/72 96/54. 144/48 


tarsus 204/42 150/52 162/45 276/42 


Male; Chelicera with segment I, 75 »; II, 162 »; movable digit 66 p (81 » with 
spermadactyl) ; both digits edentate ; pilus dentilis and dorsal seta as in female 
(Text-fig. 62D). Fou pairs of gnathosomal setae with c.s. about 75 » apart, hyp.2 
apart 60 » apart. Deutosternum with 12 transverse rows of denticles (6-10 per 
row) ; corniculi 36 » long. Internal malae, tectum, salivary styli and pedipalps as 
in female. 

Dorsal shield (732 » X 492 ») asinfemale. Tritosternum with base 42 yp, laciniae 
132 wp. Holoventral shield (576 » X 135 ») asin Text-fig. 62H. Between sf1, 84 yp; 
between sti and sf3, 110 w. Paranal setae 54 »; postanal seta 40 pw. Peritreme 
extends to middle of coxa II ; anterior part of peritrematal shield fused with dorsal 
shield ; posterior part free on one side of the only available male, on the other side 
fused with podal elements of coxa IV and with holoventral shield. 

Chaetotaxy of legs asin female. Femur II with large ventral protuberance distal 
to fissure. Genu and tibia II with setae av stout basally. Tarsus II with several 
stout, blunt setae (av,, al,, pl, aV2, Py) ; seta mv stout basally with sharp, tapering 
point. Length/width (in ,) of leg segments : 


I II III IV 
genu 114/48 114/70 84/50 120/48 
tibia 120/45 102/60 87/48 126/45 


tarsus 186/42 126/40 126/40 234/38 


HaBiTaT: Found in Britain and Europe in the nests of Riparia riparia (Linn.). 
Recorded by Bregetova from the nests of a variety of rodents in the U.S.S.R. 


Subfamily PSEUDOLAELAPINAE nov. 


Adults: Chelicera of both sexes chelate-dentate ; pilus dentilis short, setiform ; 
dorsal seta and arthrodial processes present ; fissures normal. Male chelicera with 
spermadactyl grooved, free distally. Chaetotaxy of venter of gnathosoma normal. 
Pedipalps with five free segments ; chaetotaxy of trochanter to tibia (2-5—6-—14) ; 
apotele two-tined. Deutosternum with seven transverse rows of denticles. Cor- 
niculi horn-like, internal malae inflated basally. Tectum captuli trispinate. 

Dorsal sclerotization in both sexes comprises an entire shield ; primary chaetotaxy 
reduced, consisting of 24 pairs of setae. 

Tritosternum bipartite, laciniae pilose. Pre-endopodal shields present. Sternal 
shield in the female with three pairs of setae and two pairs of pores; metasternal 
setae on small platelets. Genito-ventral shield inflated, bearing the genital and two 
pairs of opisthogastric setae. Anal shield with normal chaetotaxy, no euanal setae. 


266 G. OWEN EVANS & W. M. TILL 


Opisthogastric cuticle sparsely setose. Metapodal shields small, ovoid. Peritrema- 
tal shields fused with large podal shields posterior to coxae IV ; peritremes extend 
beyond coxae I. Male with holoventral shield fused with peritrematal shields ; 
genital orifice presternal. 

All legs six-segmented, with ambulacra and claws. Deviations from normal 
chaetotaxy occur on genu and tibia I (2—}, }—2), genu II (2—%, ?—2) and genu III 
(2— 7, 3—1). Coxa II with rounded anterior protuberance. 

Immature stages not known. 

A single genus. 


Genus PSEUDOLAELAPS Berlese 


Hoplolaelaps Berlese, 1910, Redia 6 : 259 (non Hoplolaelaps Berlese, 1903). 
Pseudolaelaps Berlese, 1916, Redia 12 : 30. 


Type: Laelaps (Hoplolaelaps) doderoi Berlese, 1910. 


With the characters of the subfamily. 


Pseudolaelaps doderoi (Berlese) 


Laelaps (Hoplolaelaps) doderoi Berlese, 1910, Redia 6 : 259. 


Female ; Chelicera with segment I, 54 »; II, 130 »; movable digit 50 », biden- 
tate; fixed digit with about nine teeth, pilus dentilis short, setiform ; dorsal seta 
nearly half-way towards apex of digit (Text-fig. 63E). Four pairs of gnathosomal 
setae with c.s. about 42 » apart, hyp.2 about 36 » apart. Deutosternum with seven 
transverse rows of denticles (3 to 7 denticles per row) ; a row of small denticles 
postero-lateral to the capitular setae ; corniculi 33 » long, horn-like ; internal malae 
broad basally, tapering to fine points (Text-fig. 63F). Tectum capituli trispinate, 
lateral margins denticulate (Text-fig. 63B). Pedipalps (2—5-6-14) with two-tined 
apotele. 

Dorsal shield (474 » & 318 ») with 24 pairs of setae distributed as in Text-fig. 634, 
some posterior setae paddle-shaped. Surface of shield reticulate. 

Tritosternum with base 27 p, laciniae 70 w. Sternal shield (100 » X IIO ») with 
three pairs of setae and two pairs of pores; metasternal setae on small shields, 
associated pores not visible. Between sf1, 48 » ; between sf1 and st3, 78 ». Genito- 
ventral shield (135 » * 100 m) greatly expanded posteriorly, with a pair of genital 
and two pairs of opisthogastric setae ; greatest width of shield about 204 ». Anal 
shield (48 » & 108 yw) with paranal setae 18 p, postanal seta 15 » long. Chaetotaxy 


hic.63. Pseudolaelaps doderoi Berlese. a, Dorsum of idiosoma of female ; B.tectum capituli 
offemale ; c. cheliceraof male, internal view ; Db. venter of idiosomaof male; E. chelicera 
of female, internal view; F. venter of gnathosoma of female; G. venter of idiosoma of 
female. 


THE BRITISH DERMANYSSIDAE (CEC Ait 


) 


267 


ZOOL. 14, 5. 


18 


268 G. OWEN EVANS @.W. M. TILL 


and sclerotization of opisthogaster as in Text-fig. 63G. Peritreme extends beyond 
lateral margin of gnathosoma ; peritrematal shield fused with large metapodal shield. 

Chaetotaxy of legs differs from normal pattern on genu and tibia I (2—}, {—2), 
genu II (2—%, }—2), and genu III (2—7, §—1). Coxa II has a rounded anterior 
protuberance. Length/width (in ,) of leg segments : 


I II Ill IV 
genu 66/24 54/30 42/24 60/27 
tibia 63/18 54/30 45/27 72/24 
tarsus 108/15 117/24 108/24 170/20 


Male: Chelicera with segment I, 42 »; II, 100 »; movable digit 42 pw (54 » with 
spermadactyl), unidentate ; fixed digit with two teeth and a minute, setiform, pilus 
dentilis on a small, tooth-like projection (Text-fig. 63c). Four pairs of gnathosomal 
setae with c.s. about 36 w apart ; hyp.2 about 30 » apart. Deutosternum with six 
transverse rows of denticles (two to several per row) ; a row of small denticles postero- 
lateral to setae c.s. Corniculi 30 » long ; internal malae, tectum and pedipalps as in 
female. 

Dorsal shield (354 » X 246 «) with 24 pairs of setae as in female. Tritosternum 
with base 12 p», laciniae 57 ». Holoventral shield (282 » x go ») with four pairs of 
setae and three pairs of pores in the sternal region, four pairs of setae in the genito- 
ventral region (Text-fig. 63D). Between sti, 40 »; between sti and st3, 66 pz. 
Paranal setae 15 », postanal seta 12 » long. Peritreme as in female, peritrematal 
shield fused with holoventral shield. 

Chaetotaxy of legs as in female. Length/width (in ,) of leg segments : 


I II Ill IV 
genu 54/20 48/27 40/24 50/20 
tibia 57/18 45/27 40/24 60/20 
tarsus go/15 go /20 go /20 144/18 


Hasitat : Recorded in Britain from the soil of a grassy bank, London, and from an 
arable field on clay/chalk, Winchester, Hampshire. Originally found in soil in Italy. 


Subfamily MYONYSSINAE Bregetova 


Myonyssinae Bregetova, 1956, Opred. Faune SSSR. 61: 52. 


Adults ; Chelicerae essentially edentate, fixed digit long and slender in both sexes 
and in the female it may have a small dorsally directed tooth. Spermadactyl 
relatively short, grooved, and entirely fused with the movable digit. Pilus dentilis 
and dorsal seta apparently absent, fissures inconspicuous, arthrodial processes absent. 
Chaetotaxy of venter of gnathosoma normal; pedipalps with five free segments 


THRE BRITISH DERMANYSSIDAE (ACARI) 269 


and a two-tined apotele, chaetotaxy normal (2~5-6-14). Deutosternum with 9-12 
denticles arranged in a single file. Corniculi horn-like, relatively weakly slcerotized ; 
internal malae long, simple. Salivary styli massive. Tectum capituli elongate, 
tapering distally and with a smooth margin. 

Dorsal shield entire in both sexes ; in the female typically with 38 pairs of setae 
comprising 21 pairs of podonotals (z2 lacking) and 17 pairs of opisthonotals (px2 and 
px3 present). Sometimes normal complement decreased (MM. montanus) by the 
absence of x2, 6x3 and S3 on the opisthonotal region. Dorsal shield in the males of 
two main types, one in which the chaetotaxy resembles that of the female (MV. 
montanus group), and the other in which the female complement is increased by the 
incorporation of certain of the 7 and KF setae on the lateral margins of the shield 
(M. gigas group). Sexual dimorphism is evident in the second type, by the extreme 
length of certain of the setae on the posterior region of the shield. 

Tritosternum bipartite, base lacking hyaline denticulate border. Sternal shield 
well-sclerotized, reticulate, slightly wider than long and bearing three pairs of setae 
and two pairs of pores. Genito-ventral shield large, flask-shaped, with from 4-22 
setae excluding the genitals. Anal shield with normal three setae, rarely with an 
additional unpaired accessory seta, and considerably wider than long. Metapodals 
present. Opisthogastric cuticle with varying degrees of hypertrichy. Peritrematal 
shields free posteriorly. Male with holoventral shield free from peritrematal shields 
and with hypertrichy of the ventral region. 

Chaetotaxy of legs normal. Coxa II usually with prominent anterior spine. All 
ambulacra with well-developed claws. 


Larva and Protonymph : Apparently unknown. 


Deutonymph :; Active feeding instar. Gnathosoma with characteristics of female. 
Dorsal shield with lateral incisions ; chaetotaxy of the dorsum as in the female, but 
fewer setae situated on the dorsal shield. Tritosternum bipartite, pilose. Sternito- 
genital shield with sternal setae 1-3 ; metasternals and genitals flanking the shield. 
Anal shield similar in shape to that of female; chaetotaxy of opisthogaster as in 
adult. Peritremes shorter than in the adult. Leg chaetotaxy normal. 


Genus MYONYSSUS Tiraboschi 
Myonyssus Tiraboschi, 1904, Arch. Parasit. 8 : 337. 
Tetvagonyssus Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 6. 
Type: Myonyssus decumami Tiraboschi, 1904. 


The genus has the characters of the subfamily as diagnosed above. 


KEY TO FEMALES 


Genito-ventral shield with less than 12 setae ; anal shield about 14 times as broad as 

long . . M.decumani Tiraboschi (p. 271) 
Genito- Pama, shield au aeud 20 ae (Text fi 64B) ; anal shield about twice as 

broad as long , d . : : M. gigas (Oudemans) (p. 271) 


G. OWEN EVANS & W. M. TILL 


Poin BRET IiSoH DERMANYSSIDAE: (ACART) Pighii 


Myonyssus decumani Tiraboschi 


Myonyssus decumani Tiraboschi, 1904, Arch. parasit. 8 : 337; Hirst, 1916, J. zool. Res. 1: 64, 
figs. : 


We have not examined specimens of this species which is adequately described and 
figured by Hirst (1916). 

HapitatT: The only British records of the species are from a house mouse (Mus 
musculus Linn.) at Ollaberry, Shetland Islands (Hirst, 1916). 


Myonyssus gigas (Oudemans) 


Liponyssus gigas Oudemans, 1912, Ent. Ber. Amst. 3 : 231. 
Myonyssus gigas; Bregetova, 1956, Opred. Faune SSSR. 61 : 123, figs. 243, 251. 


Female: Chelicera with segment I, 48 »; II 189 » ; movable digit 57 yw, edentate; 
fixed digit unidentate (Text-fig. 65A and B); pilus dentilis absent. Four pairs of 
gnathosomal setae with c.s. about 72 wapart, hyp.2 about 54 wapart. Deutosternum 
with eleven denticles arranged in a single file; corniculi 45 p» long, weakly 
sclerotized ; internal malae elongate (Text-fig. 65c). Salivary styli conspicuous, 
extending well beyond tips of corniculi. Tectum capituli triangular with smooth 
margin (Text-fig. 65D). Pedipalp (2-5-6-14) with two-tined apotele. 

Dorsal shield (1,140 » X 636 ») with 38 pairs of setae distributed as in Text-fig. 
644A ; seta 22 is absent. Setae Z5 and S5 considerably longer than the other setae 
and very stout and blade-like. Surface of shield reticulate with a granular pattern 
posteriorly ; no hypertrichy. 

Tritosternum with base 70 p, laciniae 170 p. Sternal shield (210 w x 252 p») with 
three pairs of setae and two pairs of pores ; metasternal setae free with associated 
pores ; between sf1, 120 » ; betweensf1 andsi3,174. Genitalshield (282 » x 160 p) 
flask-shaped with one pair of genital setae and about 22 opisthogastric setae which 
are not strictly symmetrical in their arrangement (Text-fig. 648). Anal shield 
(186 w X 294 ») with deeply concave anterior margin and maximum width 408 pz. 
Paranal setae 60 pw, postanal seta 66 uw. Chaetotaxy and sclerotization of opistho- 
gaster as in Text-fig.64B. Peritreme extends to posterior third of coxa II ; anterior 
part of peritrematal shield fused with dorsal shield, posterior part free. 

Legs with normal chaetotaxy. Coxae II-IV are provided with spurs, as shown in 
Text-fig. 648. Length/width (in ») of leg segments : 


I II Ill IV 
genu 144/96 144/102 120/90 150/84 
tibia 135/84 108/96 108/84 150/84 


tarsus 174/70 144/72 168/70 216/72 


Fic. 64. Myonyssus gigas (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. 


272 G. OWEN EVANS & W. M. TILL 


Male ; Chelicera with segment I, 60 » ; II, 180 » ; movable digit 80 », both digits 
edentate (Text-fig. 65£). Four pairs of gnathosomal setae with c.s. about 72 » apart, 
hyp.2 about 57 » apart. Deutosternum, internal malae, salivary styli, tectum and ‘ 
pedipalps as in female. Corniculi 54 » long, slender, weakly sclerotized. 


lic. 65. Myonyssus gigas (Oudemans). A-B. Chelicera of female; c. venter of gnatho- 
soma of female; bp. tectum capituli of female _ £. chelicera of male. 


Dorsal shield (1,188 » * 684 «) with 21 pairs of setae in the podonotal region, as in — 
the female, whereas the opisthonotal region bears 22 pairs of setae, five marginal 
pairs being included on the shield (Text-fig. 64c). The setae are relatively longer 
than in the female and some, particularly the Z and S series and J4—5, are sinuous. 

Tritosternum with base 36 yp, laciniae 174 ». Holoventral shield (984 » x 270 p) 
with four pairs of setae and three pairs of pores in the sternal region and numerous 
setae in the genito-ventral region. Between sf1, 114 » ; between sf1 and s¢3, 198 p. 
Paranal setae 60 «, postanal seta 78 yw. Chaetotaxy and sclerotization of opistho- 


THE BRITISH DERMANYSSIDA® (ACARI) 2793 


gaster asin Text-fig.64D. Peritreme extends to posterior third of coxa II ; anterior 
part of peritrematal shield fused with dorsal shield, posterior part free. 

Legs with normal chaetotaxy. Coxae I-IV provided with spurs, as shown in 
Text-fig. 64D. Length/width (in p» ) of leg segments : 


I ‘al III IV 
genu 105/168 ,/ WGB/132 " TAA/TI4- ¥O2/117 
tibia 162/96 138/120 - 126/102 ~ 180/ErO 


tarsus 192/84 174/84 180/78 264/87 


HABITAT: Found on moles (Lalpa europaea Linn.) and in their nests, and on field 
mice [A podemus flavicollis (Melchior) and Apodemus sylvaticus (Linn.)] in Britain and 
Europe. 


Subfamily MELITTIPHINAE nov. 


Adults : Chelicerae in the female chelate, fixed digit essentially edentate, movable 
digit weakly bidentate. Chelicerae of male strongly modified ; fixed digit reduced ; 
movable digit fused with long, slender, grooved spermadactyl. Pilus dentilis present 
in female ; dorsal seta and fissures normal ; arthrodial processes present, at least in 
female. Chaetotaxy of venter of gnathosoma and pedipalps normal. Apotele two- 
tined. Deutosternum with about eight denticles in a single file. Corniculi slender, 
horn-like ; internal malae long, bipartite anteriorly. Tectum capituli with smooth, 
rounded, anterior margin. 

Dorsum of idiosoma entirely covered by a single shield, hypertrichy obscuring the 
primary chaetotaxy. Certain marginal setae subspinose. 

Tritosternum bipartite, normal. Presternal area moderately sclerotized, with a 
median keel-like ridge. Sternal shield in female with three pairs of setae and two 
pairs of pores. Genital shield flask-shaped with genital setae only. Anal shield 
ovoid, with aciculated portion bent dorsally, resulting in posterior margin of shield 
appearing concave. Anus situated in posterior third of shield ; a posteriorly directed 
spur present immediately anterior to the anus. Opisthogastric cuticle hypertrichous. 
Peritrematal shields free posteriorly. Male with a sternito-genital and a ventro-anal 
shield. 

Chaetotaxy of legs normal except tibia III (2—+, ?—2z) and genu IV (2—?, 3—1). 
Coxa II with complex anterior spine (Text-fig. 66c). All ambulacra with a pair of 
claws. 

The genus Melittiphis was previously placed in the family Eviphididae, on the basis of 
Vitzthum’s (1930) description of the tectum capituli, namely : “‘ Epistom dem [p/is- 
Typus entsprechend eine langgezogene Spitze, beiderseits und dorsal fein gezahnelt ”. 
This description refers to the labrum and not to the tectum (see Text-fig. 668). 

Immature stages not known. 


G. OWEN EVANS & W. M. TILL 


274 


- Pa 
: 
- 
\~— 


Te. ' + 
——3 mm eT ee a re rr qe 2 8 em 


Se me paabecesqubtles 40 goCaunes Serna oS cere re ne 
err z 
- [oa) 
> : 
- > 2 ae : 
\ = . %, | . 
rl Pr F 
~ > “< : ; 
: j 
- 


male ; dorsum of idiosoma (a) ; venter of gnatho- . 


venter of idiosoma (F). 


tectum capituli (E) ; 


coxa II (c); chelicera (D) ; 


Melittiphis alvearius (Berlese), fe 


soma (B) ; 


Fic. 66. 


22 BRITISH DE RIAN YSSIDAE:(ACART) 275 
Genus MELITTIPHIS Berlese 
Melittiphis Berlese, 1918, Redia 13 : 117. 


Type: Laelaps (Iphis) alvearius Berlese, 1896. 


The genus has the characters of the subfamily as diagnosed above. 


Melittiphis alvearius (Berlese) 
Iphis alvearius Berlese, 1896, Atti Soc. Veneto-Tvent. (2) 2 : 318. 


Female ; Chelicera with segment I, 48 w; II, 115 » ; movable digit 45 », weakly 
bidentate ; fixed digit edentate, pilus dentilis and dorsal seta short, setiform (Text- 
fig.66D). Four pairs of gnathosomal setae with distance between c.s. slightly exceed- 
ing distance between hyp.2. Deutosternum with eight denticles arranged in a single 
file ; corniculi 36 w long, converging; internal malae elongate, bipartite anteriorly 
(Text-fig. 668). Tectum capituli with anterior margin smooth (Text-fig. 66E). 
Pedipalp (2-5—6-14) with two-tined apotele. 

Dorsal shield (780 ~ < 660 ») shows marked hypertrichy ; marginal setae relatively 
stout and spine-like (Text-fig. 664A). Surface of shield reticulate. 

Tritosternum with laciniae about three times as long as the base. Sternal shield 
(75 » X 80 pw) with three pairs of setae and two pairs of pores; metasternal setae 
situated over the endopodal shields. Between sf1, 30 »; between sfr and sf3, 78 wp. 
Genital shield (90 » < 147 ») with genital setae only. Anal shield (204 » x 138 p) 
with paranal setae 40 pw, postanal seta 48 pw. Chaetotaxy and sclerotization of 
opisthogaster as in Text-fig. 66F. Peritreme extends nearly to anterior margin of 
coxa I; anterior part of peritrematal shield fused with dorsal shield at level of seta 
22, posterior part free. 

Chaetotaxy of legs differs from normal pattern in that tibia III has two postero- 
lateral setae (2—+, ?—2), genu IV has two ventral setae (2-7, #—1), tarsi II-IV lack 
setae ad, and fd,. Stout spine-like setae are present on femur II (ad,, pd,, pl, and 
vs) ; trochanter III (al) ;, femur III (distal dorsal) ; genu III (ad,) ; trochanter IV 
(al) ; femur IV (distal dorsal) ; genu IV (ad,_,). Length/width (in p) of leg seg- 
ments * 


I II III IV 
genu 72/45 84/70 63/57 73/54 
tibia 72,36 60/54 54/48 66/48 


tarsus 138/36 135/42 144/42 168/42 


Male: Certain details of the morphology of the male, given in the subfamilial 
diagnosis, have been obtained from a specimen in the Berlese collection, Florence. 


HasitatT: Found in bee-hives in Britain and Europe. 


G. OWEN EVANS @& W. M. TILL 


te 
J 
o 


Subfamily HIRSTIONYSSINAE nov. 


Adults ; Chelicerae in female chelate-edentate, movable digit rarely bidentate or 
weakly dentate ; fixed digit with hyaline process which may be produced distally 
into a slender lobe (Patrimyssus). Male with relatively short, grooved spermadactyl 
usually completely fused with the movable digit; fixed digit edentate. Pilus 
dentilis apparently absent, dorsal seta reduced to microseta or absent, fissures present, 
arthrodial processes reduced or absent. Chaetotaxy of venter of gnathosoma normal. 
Pedipalpal chaetotaxy normal or femur unideficient, genu unideficient, or tibia uni- 
or bideficient ; apotele simple (Patrimyssus) or two-tined. Palptrochanter rarely 
with ventral keel produced posteriorly into a spur (7 7vichosurolaelaps). Deutosternum 
with five to 18 transverse rows of denticles, transverse rows comprising one to four 
denticles. Hypostome elongate, internal malae simple, often long ; corniculi mem- 
branous, rarely with horn-like apex. Tectum capituli tongue-like, with anterior 
margin smooth or fimbriated. 

Dorsum with entire shield in both sexes, bearing 22-27 pairs of setae in the female, 
never holotrichous ; setae z1I rarely present, J series showing hypotrichy, J/5 present 
or absent ; opisthonotal region of shield with less than 15 pairs of setae in the female. 
Sexual dimorphism apparent in the relative lengths of the lateral and marginal setae, 
those of the male being considerably longer than in the female. Dorsal setae simple. 


Tritosternum normal, bipartite, laciniae smooth or pilose, without hyaline den- 
ticulate margins. Sternal shield in female with three pairs of setae and two pairs of 
pores ; metasternals free. Genital shield tongue-shaped, normally bearing only the 
genital setae. Analshield normal, opisthogastric cuticle with a variable number of setae 
which may be affected by sexual dimorphism. Metapodal shields weakly developed 
or absent. Peritrematal shields free or fused posteriorly with the podal shields. 
Stigmatal atrium normal or greatly enlarged, peritremes often reduced. Podal 
shields poorly developed posterior to coxae IV. Male with holoventral shield not 
fused with peritrematal shields ; rarely with division or weaker sclerotized region 
between the ventral and anal portions of the shield. Genital orifice presternal. 


Chaetotaxy of legs normal (Hirstionyssus) or with deficiencies in the chaetotaxy of 
genua and tibiae III and IV. Anterior spine of coxa II well developed or absent 
(Patrinyssus), that of Echinonyssus greatly enlarged and hook-like. One or more of 
the coxae with a distinct, non-setigerous spur or spurs (excluding anterior spine of 
coxa Il). Certain coxal setae and ventral setae of tibiae and genua may be hypertro- 
phied and form retrograde spurs. Ambulacra with well-developed claws. Male with 
thumb-like ventral spur on femur II in some Echinonyssus. 

Immature stages: The larval and protonymphal stages have been described only 
for Htrstionyssus (Bregetova, 1956). The larva is apparently non-feeding and has 
weakly developed chelicerae with minute digits. On the other hand, the proto- 
nymph is probably a feeding stage although the form of the chelicerae is not shown in 
detail by Bregetova. It lacks distinct dorsal and ventral sclerotization. All known 
deutonymphs are feeding stages with the general characteristics of the female except 


fHE BRITISH DERMANYSSIDAE (ACARI) 277 


for the intercoxal region which has a sternito-genital shield bearing four pairs of 
setae ; the genital setae flank the shield in the region of coxae IV. 
This subfamily is represented in the British fauna by the genus Hzrstionyssus. 


Genus HIRSTIONYSSUS Fonseca 
Hirstionyssus Fonseca, 1948, Proc. zool. Soc. Lond. 118 : 266. 


Type: Hurstionyssus talbae Zemskaya, 1955 (= Huirstionyssus arcuatus (Koch) 
Fonseca, 1948). 


Medium sclerotized mites ranging from 450 » to 950 winlength. Chelicerae chelate- 
edentate in both sexes ; fixed digit in the female with membranous sheath. Movable 
digit in male completely fused with the deeply grooved spermadactyl. Pilus dentilis 
apparently absent ; dorsal seta reduced to microseta or absent ; fissures present. 
Arthrodial processes at base of movable digit reduced or absent. Chaetotaxy of 
venter of gnathosoma normal ; pedipalpal chaetotaxy normal (2—5—6—-14), or palp- 
tibia unideficient (2-5—6—-13) or bideficient (2—5-6-12), apotele two-tined. Palp- 
trochanter without ventral keel. Deutosternum with 11 to 18 rows of denticles, 
each row comprising one to four denticles. Internal malae simple, variable in length ; 
corniculi membranous, never horn-like. Salivary styli strong. Tectum capituli 
elongate, tongue-shaped, with anterior margin smooth or fimbriated. 

Dorsum of idiosoma with entire shield bearing 23 to 27 pairs of setae; zI never 
present. Chaetotaxy of opisthonotal region may show intraspecific variation. 
Unpaired accessory setae never present. 

Tritosternum normal, bipartite. Sternal shield in female with three pairs of setae 
and two pairs of pores. Genital shield tongue-shaped, bearing one pair of genital 
setae only. Pear-shaped anal shield with normal chaetotaxy, paranal setae approxi- 
mately level with or behind anterior margin of anus. Opisthogastric cuticle with 
a variable number of setae which may be affected by sexual dimorphism. Peri- 
trematal shields free or fused posteriorly with the podal shields. Metapodal shields 
weakly developed or absent. Male with holoventral shield not fused with peri- 
trematal shields, and bearing eight pairs of setae excluding the anals. Podal shields 
poorly developed posterior to coxae IV. 

Chaetotaxy of legs normal. At least one of the coxae provided with non-setigerous 
spurs, apart from the anterior spine of coxa II. All ambulacra with a pair of well- 
developed claws. 

As in many other genera of obligatory ectoparasitic dermanyssid mites, there is 
considerable confusion in the literature concerning the nomenclature and systematic 
position of certain of the “ older’ species. This is particularly evident in the genus 
Hirstionyssus and concerns, in the present study, the species Hirstionyssus arcuatus 
(Koch) and musculi (Johnston). The confusion has arisen largely from Oudemans’ 
concept of the species Dermanyssus arcuatus Koch, Acarus muscult Schrank and 
Dermanyssus muscult Koch. 

The type host of D. arcuatus is the chiropteran “ Vespertilio noctula’’, but the 
original description and illustration of the species are inadequate for its specific 


278 G. OWEN EVANS & W. M. TILL 


identity. Berlese (1889), following Canestrini 1885, attributed arcuatus {Letognathus 
arcuatus (IX.) Can.) to a species of dermanyssine mite commonly found on vesper- 
tilionid bats (particularly “‘ V. noctula "’) and his figures and description leave one in 
no doubt that he considered arcuatus to be a member of the genus Steatonyssus 
(probably S. periblepharus Kolenati) in its present concept. However, Oudemans 
(1913), without reference to Berlese or Canestrini, considered Dermanyssus arcuatus 
Koch, which he transferred to the genus Liponyssus, to be conspecific with a dermanys- 
sine mite associated with Talpa and gave a comprehensive description, with figures, 
of this mole mite. Unfortunately subsequent workers, excluding Zemskaya and 
Bregetova in the U.S.S.R., have accepted Oudemans’ concept of arcuatus and Fonseca 
(1948)? proposed the genus Hirstionyssus to accommodate the type Hirstionyssus 
arcuatus (Koch, 1839) and several congeneric species. Hirstionyssus arcuatus (Koch) 
Fonseca, 1948, andits congeners are essentially ectoparasites of Insectivora, Carnivora, 
Lagomorpha and Rodentia; there are no authenticated records of any Hirstionyssus 
species from Chiroptera. Thus, there can be no doubt that H. arcuatus sensu 
Fonseca 1948 and Oudemans 1913, is neither conspecific nor congeneric with Der- 
manyssus arcuatus Koch, 1839 and, in our opinion, Zemskaya (see Bregetova, 1956) 
was correct in proposing the name Hirstionyssus talpae for Liponyssus arcuatus sensu 
Oudemans, 1913 [= Hirstionyssus arcuatus (Koch) Fonseca}. Huzrstionyssus talpae 
Zemskaya, 1955, becomes the type of the genus Hzrstzonyssus Fonseca. 

Strandtmann & Wharton (1958) have continued to use Hirstionyssus arcuatus 
(Koch) sensu Oudemans 1913 and their synonymy of the species and list of hosts 
demonstrate the utter confusion surrounding thisname. The synonyms Dermanyssus 
albatus Koch, 1839, Dermanyssus noctula Koch, 1839 (teste Oudemans, 1936) and 
Dermanyssus pipistrellae Gervais, 1841, are all bat mites and probably belong in the 
Macronyssinae whilst the host list contains several species of Chiroptera, Putorius 
erminea, as well as Talpa europaea and many Rodentia. There is little point in 
attempting to conserve Huirstionyssus arcuatus under these conditions. We 
strongly advise for the sake of nomenclatural stability the acceptance of Hirstionyssus 
talpae for Hirstionyssus arcuatus Oudemans non Koch, 1839, and the removal of 
Dermanyssus arcuatus, albatus (based on a protonymph), noctulae and pipistrellae 
from Hirstionyssus to the subfamily Macronyssinae where they can remain as species 
incertae sedis. Reference to Chiroptera as hosts of Hzrstionyssus should in future 
be excluded unless, of course, the record can be authenticated. 

Till & Evans (1964) have already commented on Oudemans’ concepts of Dermanys- 
sus muscult Koch 1839 and have shown that at one time or another he considered the 
species to be congeneric with Steatonyssus, Macronyssus and Ornithonyssus. His 
final opinion appears to be expressed in the “‘ Kritisch Historisch Oversicht der Acaro- 
logie ’’ (1936) where he considered D. musculi to be synonymous with Acarus muscult 
Schrank 1803 and referred both to the genus Steatonyssus. Both Schrank and Koch 
recorded their species from the house mouse (Mus musculus) and since there have 


’ Fonseca states: ‘ Hirstionyssus, gen. nov., here created to include numerous species related to 
Dermanyssus arcuatus Koch, 1839 (= Liponyssus arcuatus auct.) parasites on Chiroptera, Rodentia 
etc....Genotype Hirstionyssus arcuatus (Koch, 1839) ”’. 


THE BRITISH DEKMANYSSIDAE (ACARI) 279 


never been any confirmed records of species of the genus Steatonyssus from this host 
(or from any other rodent for that matter), it seems almost certain that Schrank’s 
and Koch’s mites do not belong to this genus. Zemskaya (1955) and Bregetova 
(1956) who were the first to examine critically the old ‘‘ Hirstionyssus arcuatus 
complex ”’ showed that the species occurring on Talpa was not conspecific with the 
species parasitizing Mus, Rattus and Apodemus, for which they adopted the name 
Hirstionyssus musculs (Johnston). In our opinion this name cannot be used for this 
species for the following reasons : 


ig Johnston (1849) described his species under the name Dermanyssus musculi 
and indicated that he considered it to be conspecific with D. muscult Koch 
by referring to “ Dermanyssus musculi, Koch Uebers. Arachnid p. 81 
tab. 9 fig. 46”’ immediately after his Latin diagnosis of the species. If 
Johnston’s species is not conspecific with D. musculi Koch, as Zemskaya and 
Bregetova infer, then they should have used the combination Hzrstionyssus 
johnstont (Oudemans, 1936) [= Macronyssus johnstont Oudemans nom. nov. 
pro Dermanyssus musculi Johnston 1849 non Koch 1836] and not Hirstionys- 
sus musculs (Johnston). 

z. Itis clear from Johnston’s description that his species was not a Hirstionyssus ; 
for example, its size is stated to be equal to Gamasus coleoptratorum (whose 
nymphs measure about 1,000 p in length), the body is thickly covered with 
very short, adpressed, somewhat curved bristles and the “ bristles ”’ of the 
legs are stated to be “setaceous, pointed downwards, longer than the 
diameter of the joint, barbed with minute spinules on one side’”’. John- 
ston’s mite was probably a species of Haemogamasus. 


The original descriptions of the colour of the idiosoma of Acarus muscult Schrank 
and its synonym Dermanyssus muscult Koch indicate that the species is a haemato- 
phagous, obligatory ectoparasite. Schrank’s “braunroth ... und das Vorder- 
ende weisslicht ’’ and Koch’s “ Der Vorderleib rein weiss, ein Schieffleck, von den 
Seiten bis zum Hinterrande reichend dunkelroth, dazwischen ein herzf6rmiges 
Fleckchen hellroth ”’ could refer to engorged forms of both Hzrstionyssus and Orm- 
thonyssus in which the general brownish-red colouration of the idiosoma does not 
extend to the anterior region, the colour pattern being determined by the blood- 
containing diverticula of the “stomach ’”’. The description of the first pair of legs— 
‘““ Fusspaar etwas langer und schmaler ’’—by Schrank and the form of these appen- 
dages in Koch’s illustration of the species excludes the possibility that A. muscult is a 
Hirstionyssus and supports our previous suggestion that the species ““ might possibly 
be conspecific with O. bacoti’’ (Till and Evans, 1964). 

Hirstionyssus musculi (Johnston) Bregetova appears to be conspecific with at least 
two, possibly three, presently considered valid species of Hirstionyssus,® namely, H. 
cynomys (Radford, 1941) from Cynomys species, H. latiscutatus (de Meillon & Lavoi- 


8 Hirstionyssus otomys (Radford, 1942) also bears a superficial resemblance to this complex but may 
be readily distinguished by having only 12 setae on the palptibia and the tendency for the third pair 
of sternal setae to be off the sternal shield. We have recently found this species on Arvicanthis abys- 
sinicus from the Congo. 


280 G. OWEN EVANS @& W. M. TILL 


pierre, 1944) from Rattus rattus and H. orcadensis (Turk, 1946) from Microtus orca- 
densis. We are unable to distinguish morphologically the type specimens of the 
females of H. latiscutatus and orcadensis from our material (including specimens 
presented by Dr. Bregetova) of “ H. musculi”’ from Mus musculus and Apodemus 
sylvaticus ; all occur on murines. H. cynomys, of which we have examined only the 
‘“ type ’’ specimen, is undoubtedly closely related to “ musculi ’’, but shows differences 
in the form of setae av, and pv, on tarsi Il and IV ; on tarsus II these setae are claw- 
like but longer and less distinctly curved while on tarsus IV seta av, is stout and spine- 
like and fv, is relatively slender and whip-like distally. In view of these slight 
morphological differences and the systematic position of the host (Cynomys : Sciuridae) 
we are reluctant to assign ‘‘H. musculi (Johnston) Bregetova’’ to this taxon and 
propose using the next available combination, Hirstionyssus latiscutatus (de Meillon 
& Lavoipierre). 


KEY TO FEMALES 


1 Sternal shield rectangular. Podonotal region with setae st and s2 on integument. 
Coxae Il and III each with one spur (including anterior spurof coxa II). Palptibia 
with 13 seta ; _ H., blanchardi (Trouessart) (p. 296) 
— Sternal shield arched, posterior border markedly concave. Podonotal region with 
setae st and s2 on shield. Coxae II and III each with two spurs. Palptibia with 
12 to 14 setae ; ‘ : 
Opisthonotal region of dorsal shield with 10 Or more pairs of setae. Palptibia with 
14 setae é ; ‘ 
— Opisthonotal region ‘of dorsal shield ‘with eight pairs ‘of setae. Palptibia with 12 
or 13 setae : ; . : 
3 Tarsus II with setae avy and pv, stout, claw-like ; coxa IV with a small spur on its 
postero-distal margin. Podonotal region of dorsal shield with 16 pairs of setae (v2 
off the shield) : : . . H. latiscutatus (de Meillon & Lavoipierre) (p. 
~ Tarsus II with setae av, and pv, simple ; coxa IV without spur. Podonotal region 
of dorsal shield with 17 pairs of setae (r2 off the shield) 
H., isabellinus (Qudemans) (p. 
4 Podonotal region with 15 pairs of setae on the dorsal shield, s6 lacking ; three pairs of 
ysetaeonthecuticle. : ‘ H. soricis (Turk) (p. 
~— Podonotal region with 16 pairs of setae on the dorsal shield, s6 present ; four pairs of 
ry setae on the cuticle , ; “ 
5 Coxa II with anterior seta short, stout and blunt ; posterior spur large and pointed. 
Dorsal shield with seta z2 at least twice the length of seta sI 
H. talpae Zemskaya (p. 
~ Coxa II with anterior seta simple ; posterior spur relatively smaller. Dorsal shield 
with setae z2 and st subequal in length H., oryctolagi sp. nov. (p. 


ty 


KEY TO MALES?® 


1 CoxallI withonespur. Palptibia with 13 setae. H, blanchardi (Trouessart) (p. 
Coxa III with two spurs. Palptibia with 12 or 14 setae , : : ; 
Opisthonotal region of dorsal shield with 10 or more pairs of setae. Palptibia with 14 


te : 


setae : : : : - 
Opisthonotal region ‘of dorsal shield with eight pairs ‘of setae. Palptibia with 12 
setae 


* Male of H. oryctolagi not known. 


tee SRI isoH DERMANYSSIDAE )(ACART) 281 


3 CoxaII with anteriorspuronly. Opisthogastric setae relatively short 
H, isabellinus (Oudemans) (p. 296) 
— CoxalII withtwospurs. Opisthogastric setae extremely long 
H. latiscutatus (de Meillon & Lavoipierre) (p. 291) 
4 Dorsal shield with postero-lateral setae considerably longer than terminal (Z5) 
setae ; podonotal region of shield with 18 pairs of setae. Anterior seta on coxa II 
simple 3 é : : ‘ : : H. talpae Zemskaya (p. 283) 
— Dorsal shield with postero-lateral and terminal (Z5) setae subequal in length ; 
podonotal region of shield with 17 pairs of setae. Anterior seta on coxa II short 
and stout ; . : ; ; : : H. soricis (Turk) (p. 288) 


Hirstionyssus talpae Zemskaya 


Liponyssus arcuatus (Koch) Oudemans, 1913, Arch. Naturgesch. Abt. A 79 (9) : 68, figs. 
Hirstionyssus talbae Zemskaya, 1955, in Bregetova ef al., Opred. Faune SSSR. 59 : 360, figs. 


Female: Chelicera with segment I, 50 »; II, 93 »; movable digit 40 1; both 
digits edentate (Text-fig. 69c). Four pairs of gnathosomal setae with c.s. about 48 » 
apart ; hyp.2 about 30 w apart. Deutosternum with 12 rows of denticles (one to 
four denticles per row). Corniculi membranous, internal malae slender, elongate ; 
salivary styli conspicuous (Text-fig. 68D). Tectum capituli tongue-shaped, anterior 
margin fimbriate (Text-fig. 69D). Pedipalp (2-5-6—12) with bi-deficient chaetotaxy 
on palptibia (Text-fig. 69B) ; apotele two-tined. 

Dorsal shield (576 » xX 324 ») with 24 pairs of setae distributed as in Text-fig. 67C ; 
seta 22 about twice as long as setae sI and s2. Surface of shield granular, with 
reticulations anteriorly and laterally. About 13 pairs of setae on the unsclerotized 
integument, including 72, 73, 75 and 76; 74 is lacking. 

Tritosternum with base 36 p, laciniae 75 w. Sternal shield (24 w X 135 p) arched, 
strongly concave posteriorly, with three pairs of setae and two pairs of pores ; 
metasternals free with associated pores. Between sf1, 68 «1; between sfi and sf3, 
80 p. Surface of shield granular, reticulated in antero-lateral corners. Genital 
shield (138 » X 114 ») granular, with one pair of setae. Anal shield (96 » x 78 p) 
with par- and post-anal setae subequal in length (27 »). Chaetotaxy and sclerotiza- 
tion of opisthogaster as in Text-fig. 67D. Peritreme extends about to middle of coxa 
I ; anterior part of peritrematal shield free ; posterior part fused with podal elements 
of coxa IV. 

Chaetotaxy of legs normal. Tarsus II with setae av, and fv, stout and claw-like 
(Text-fig. 69F), tarsus IV with these setae long and slender (Text-fig. 69G). Anterior 
seta on coxa II short and stout (Text-fig. 694). Coxal spur formula (o—2-2-1). 
Length/width (in ») of leg segments : 


I II II IV 
genu 60/48 60/54 45/42 60/45 
tibia 60/45 48/48 42/40 54/42 


tarsus 90/36 72/40 72/36 90/36 


G. OWEN EVANS & W. M. TILL 


THE BRITISH DERMANYSSIDAE (ACARI) 283 


Male (large form) : Chelicera with segment I, 45 w; II, 844; spermadactyl 45 p 
long from base of movable digit (Text-fig. 69E). Four pairs of gnathosomal setae 
with c.s. about 48 » apart, yp.2 about 30 wapart. Other gnathosomal features as in 
female. 

Dorsal shield (552 » X 348 ») with 26 pairs of setae distributed as in Text-fig. 68a ; 
v2 and 73 situated on the shield. Setae 22, 24, s3-s6, 73 and postero-lateral setae 
considerably longer than any of the othersetae. Surfaceofshield granular, reticulated 
anteriorly and laterally. Unsclerotized integument of dorsum with five pairs of 
setae, including 75 and 76, 74 being absent. 

Tritosternum with base 33 p, laciniae 78 wu. Holoventral shield (444 w x 117 p) 
with four pairs of setae and three pairs of pores in the sternal region, four pairs of 
setae in the genito-ventral region. Between siI, 50 »; between siz and sf3, 98 up. 
Par- and postanal setae subequal in length (30 w). Chaetotaxy and sclerotization of 
opisthogaster as in Text-fig. 688. Peritreme extends to middle of coxa I; fused 
anteriorly with dorsal shield. Posterior part of peritrematal shield very small and 
free. 

Chaetotaxy of legs normal. Tarsus II with setae av, and fv, stout and claw-like. 
Stout, spine-like setae present on a number of segments: trochanter I (d), femur I 
(ad,, pd,-.) ; femur II (pd,) ; genu and tibia II (fv) ; femur III (v and al); genu 
and tibia III (av) ; femur IV (a/ and v) ; genu and tibia IV (av) ; tarsus IV (av, and 
mv). All the ventral setae on tarsus III and seta mv on tarsus II are stout basally, 
tapering to long fine points. Coxa II with anterior seta relatively longer than in 
female. Coxal spur formula (o—-2-2-1). Length/width (in p) of leg segments : 


I II III Ly 
genu 60/48 70/54 57/48 75) 57 
tibia 60/40 50/50 50/45 70/48 
tarsus 84/33 70/38 80/36 105/36 


Male (small form) : A small male has been found which closely resembles the large 
form except in its body measurements and the relative length and thickness of the 
setae. 

Dorsal shield (468 » xX 258 yw) with chaetotaxy as in large form. Holoventral 
shield (372 » X I14 p) a little more slender than in large form ; opisthogastric setae 
relatively shorter (Text-fig. 68c). Between sf1, 50 »; between siti and sf3, 78 uw. 
Par- and postanal setae about 20 vp long. 

Chaetotaxy of legs as in large form, but the stout setae are not as greatly enlarged. 
The spur on coxa IV is a rather small, rounded process. Length/width (in p) of leg 
segments : 


Fic. 67. A-B. Hirstionyssus oryctolagi sp. nov., female; dorsum (A) and venter (B) of 
idiosoma. cC-—D. Hirstionyssus talbae Zemskaya, female ; dorsum (c) and venter (D) of 
idiosoma. 


ZOOL. 14, 5. 19 


284 G. OWEN EVANS & W. M. TILL 


Pic. 68. Hirstionyssus talpae Zemskaya. A. Dorsum of idiosoma of male; B. venter of 
idiosoma of male, large form; c. venter of idiosoma of male, small form; pb. venter of 
gnathosoma of female. 


PoE BRITISH DEKRMANYSSIDAE (ACART) 


285 
I II III IV 
genu 52/42 54/45 40/36 50/40 
tibia 50/36 42/42 36/36 48/36 
tarsus 75/30 60/33 66/30 78/30 


HABITAT: Common in the nests of Talpa europaea Linn. in Britain, Europe and 
qae.U.S.5.R. 


F G 


Fic. 69. A-G. Hirstionyssus talbae Zemskaya ; coxa II of female (a) ; palptibia of female 
(B) ; chelicera of female (c) ; tectum capituli of female (D) ; chelicera of male (E) ; venter 
of distal third of tarsus II of female (F); venter of distal third of tarsus IV of female 


(G). H. Hirstionyssus oryctolagi sp. nov. ; coxa II of female. 


286 G. OWEN EVANS &_W. M. TILL 


Hirstionyssus oryctolagi sp. nov. 


Female: Chelicera with segment I, 60 »; II, 108 »; movable digit 50-54 un. 
Four pairs of gnathosomal setae with c.s. about 48 » apart, hyp.2 about 24-33 » apart. 
Deutosternum with 14-15 rows of denticles (one to three denticles per row). Cor- 
niculi membranous ; internal malae slender, elongate ; salivary styli conspicuous. 
Tectum capituli tongue-shaped, anterior margin apparently smooth. Pedipalp 
(2-5-6-12/13) with uni- or bideficient chaetotaxy on palptibia ; apotele two-tined. 

Dorsal shield (666-672 » x 366 «) with 24 pairs of setae distributed as in Text- 
fig. 607A ; setae 22, 24 and the s series subequal in length. Surface of shield granular 
with reticulations near the margin, especially anteriorly and laterally. Unsclerotized 
integument of dorsum with about 17 pairs of setae, including 72, 73, 75 and 76; 74 
is absent. 

Tritosternum with base 40 p», laciniae 75 pw. Sternal shield (32 » X 153-156 1) 
with three pairs of setae and two pairs of pores ; metasternals free with associated 
pores. Between sfI, 75-78 »; between sti and st3, 84-87 pw. Surface of shield 
granular, reticulated in antero-lateral corners. Genital shield (150-153 pw X 123- 
135 ) granular, with one pair of setae. Anal shield (102-105 » X 75-78 mw) with 
paranal setae 30 p, postanal seta 36 pw» long. Chaetotaxy and sclerotization of 
opisthogaster as in Text-fig. 67B. Peritreme extends to middle of coxa I; posterior 
part of peritrematal shield fused with podal elements of coxa IV. 

Chaetotaxy of legs normal. Tarsus II with setae av, and fv, stout ; many leg 
setae robust but not greatly thickened. Coxa II with anterior seta simple (Text-fig. 
69H). Coxal spur formula (o-2—2-1). Length/width (in ») of leg segments : 


I II II IV 
genu 72/54 72/54 48/42 66/45 
tibia 63/45 54/50 45/40 63/44 
tarsus 98/36 84/40 84/36 105/38 


HABITAT: Found on a young rabbit, Truro, Cornwall, April, 1949 (coll. Dr. F. A. 
Turk). Holotype female (1965 : 12 : 29 : 30) and female paratype in the collection 
of the British Museum (Natural History) ; four female paratypes in the collection of 
Dr. F. A. Turk. 


Hirstionyssus soricis (Turk) 


Ichoronyssus soricis Turk, 1945, Parasitology 36 : 133, figs. 
?Hirstionyssus eusoricis Bregetova, 1956, Opred. Faune SSSR. 61: 184, figs. (nom. nov. pro. 
H. soricis: Zemskaya, 1955). 


Female; Chelicera with segment I, 40 »; II, 90 »; movable digit 42 ». Four 


Fic. 70. Hirstionyssus soricis (Turk). A. Dorsum of idiosoma of female; 8B. venter of 


idiosoma of female; c. venter of idiosoma of male; pb. dorsum of idiosoma of male. 


THE BRITISH DERMANYSSIDAE (ACARI) 


288 G. OWEN EVANS & W. M. TILL 


pairs of gnathosomal setae with c.s. about 40 » apart, hyp.2 about 30 papart. Deuto- 
sternum with eleven transverse rows of denticles (one or two denticles per row). 
Corniculi membranous, internal malae slender, elongate ; salivary styli conspicuous. 
Tectum capituli tongue-shaped, anterior margin fimbriate. Pedipalp (2—5-6—12) 
with bideficient chaetotaxy on palptibia ; apotele two-tined. 

Dorsal shield (490-504 # X 252 «) with 23 pairs of setae distributed as in Text-fig. 
70A, seta s6 being absent. Surface of shield with an overall reticulate pattern, but 
from about the level of setae 73, the lines show a tendency to break up into rows of 
punctations. About ten pairs of setae on the unsclerotized integument, including r2, 
r3 and 75; r4 and 76 are lacking. 

Tritosternum with base 36 p, laciniae 66 ». Sternal shield (30 » x 114 ») arched, 
strongly concave posteriorly, with three pairs of setae and two pairs of pores ; 
metasternals free with associated pores. Between st1, 60 »; between sti and s#3, 
70-75 mw. Surface of shield with reticulations in antero-lateral corners. Genital 
shield (114-123 » X 96-98 ») with one pair of setae. Anal shield (84 » X 70 yp) 
with par- and post-anal setae subequal in length (20-24 »). Chaetotaxy and sclero- 
tization of opisthogaster as in Text-fig. 70B. Peritreme extends to middle of coxa 
1; anterior part of peritrematal shield free, posterior part fused with podal elements 
of coxa IV. 

Chaetotaxy of legs normal. Tarsus II with setae av, and pv, stout and claw-like. 
Fairly stout setae on trochanter I (d), femur I (ad, and pd,_.), femur II (ad, and pd,), 
femora III and IV (ad,-,)._ Coxa II with anterior seta short and stout. Coxal spur 
formula (o-2-2-1). Length/width (in ,) of leg segments : 


I II III IV 
genu 54/45 54/45 38/38 48/40 
tibia 50/40 42/42 36/36 48/36 
tarsus 80/34 60/36 63/33 78/33 


Male ; Chelicera with segment I, 45 »; II, 80 »; spermadactyl 45 » from base of 
movable digit. Four pairs of gnathosomal setae with c.s. about 42 p apart, hyp.2 
about 27 w apart. Other gnathosomal features as in female. 

Dorsal shield (480 » x 276 pw) with 25 pairs of setae distributed as in Text-fig. 70D ; 
r2 and r3 are situated on the shield. Setae 22, 24, s3, s4 and r3 are considerably 
longer than the other setae on the shield. Marginal areas of shield reticulated, the 
central region being mainly granular. Unsclerotized integument with eight pairs of 
setae including 75 ; r4 and 76 are absent. 

Holoventral shield (408 « * 102 mw) with four pairs of setae and three pairs of pores 
in the sternal region, four pairs or setae in the genito-ventral region. Between sfI, 
48 »; between sti and st3, 87 w. Paranal setae 20 p, postanal seta 25 p long. 
Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 7oc. Peritreme 
extends to middle of coxa I, and is fused anteriorly with dorsal shield ; peritrematal 
shield free posteriorly. 


| 
| 


THE BRITISH DERMANYSSIDAE (ACARI) 289 


Chaetotaxy of legs normal. Tarsus II with setae av, and fv, stout and claw-like. 
Several thickened dorsal setae present as in female. In addition, very stout, spine- 
like setae are present on the following segments: tibia II (fv) ; femur IV (al and v) ; 
genu and tibia IV (av) ; tarsus IV (av, and mv). Moderately stout setae are present 
on femur III (a/ and v) and genu III (av). Seta mv on tarsus II and all the ventral 
setae on tarsus III are thickened basally, tapering to long, fine points. Coxa II with 
anterior seta short and stout. Coxal spur formula (o-2-2-1). Length/width (in p) 
of leg segments : 


fi II III IV 

genu 54/40 63/45 45/36 66/42 
tibia 50/33 45/42 42/33 57/42 
tarsus 70/30 57/33 75/30 84/30 


Deutonymph : Chelicera with segment I, 30 »; II, 75 »; movable digit 33 up. 
Four pairs of gnathosomal setae with c.s. about 36 w apart, hyp.2 about 24 p apart. 
Deutosternum with ten rows of denticles (one to two denticles per row). Tectum 
capituli with anterior margin smooth. Other gnathosomal features as in female. 

Dorsal shield (336 » <X 144-162 ») with 21 pairs of setae distributed as in Text-fig. 
71c; stand s2are off the shield. Terminal setae (75) more than twice as long as the 
other setae on the shield. Surface of shield reticulated, the “lines ”’ consisting of 
rows of punctations. Unsclerotized integument of dorsum with about Ig pairs of 
setae, including sI, s2, 72, 3 and 76. 

Tritosternum with base 27 p, laciniae 54 w. Sternal shield (153-168 » x 93-98 p) 
with four pairs of setae and three pairs of pores ; metasternal setae shorter than the 
sternals. Between sti, 48 w; between sti and st3, 81-84 pw. One pair of genital 
setae situated lateral or postero-lateral to posterior tip of shield. Anal shield 
(40 » X 40 p) with paranal setae 15-18 » long, postanal seta 15 w long. Chaetotaxy 
and sclerotization of opisthogaster as in Text-fig. 71D. Peritreme extends nearly to 
anterior margin of coxa II. Anterior part of peritrematal shield free, posterior part 
not developed. 

Chaetotaxy of legs normal. Tarsus II with setae av, and fv, simple. Coxal spur 
formula (o—2-1-0). Length/width (in ») of leg segments : 


I II III IV 

genu 42/42 42/42 36/34 36/33 
tibia 42/38 40/40 33/30 38/33 
tarsus 63/32 60/30 68 /27 75/27 


HABITAT: Found on Sorex minutus Linn., Camborne, Cornwall (type material 
from collection of Dr. F. A. Turk); Sovex avaneus Linn., Craighouse, Jura, Inner 
Hebrides (coll. Dr. G. B. Corbet, 6th July, 1961), and a shrew, West Horsley, Surrey 
(coll. Dr. M. Burton, 22nd August, 1957). , 


4 
= 
_ 
e 
2 
z 
2 
n 
Zz 
< 
- 
“ 
Zz 
(63) 
z 
° 
o 


THE BRITISH DERMANYSSIDAE (ACARI) 291 


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. I-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 mw apart, Ayp.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 » X 294 ») with 26 or more pairs of setae distributed as in Text- 
fig. 724. 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 19 pairs of setae, including 
five pairs of 7 setae. 

Tritosternum with base 45 p, laciniae 78 pw. 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 w; between sii and sf3, 72 wu. 
Genital shield (108 » x 108 ») tongue-shaped, granular, with one pair of setae. Anal 
shield (84 » X 72 ) 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 fv, stout and claw-like. 
Coxal spur formula (o-2-2-1). Length/width (in p) of leg segments : 


I II Ei IV 
genu 63/48 63/48 45/38 63/38 
tibia 63/42 50/45 45/30 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 pw apart, 
hyp.2 about 30 » 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—D. Hirstionyssus sorvicis (Turk), deutonymph ; 
dorsum (c) and venter (D) of idiosoma. 


THRE BRITISH DERMANYSSIDAE<(ACARTI) 293 


Tritosternum with base 33 py, laciniae 72 ». Holoventral shield (360 » x 117 p) 
with four pairs of setae and three pairs of pores in the sternal region, four pairs of 
setae in the genito-ventral region. Between st1, 50 uw; between sti and sf3, 72 yp. 
Paranal setae 33 p, postanal seta 27 mw 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 fv, 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 i} 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 w; II, 72 »; movable digit 40 pz. 
Four pairs of gnathosomal setae with c.s. about 40 pw apart, hyp.2 about 26 pw 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. 714 ; 
unsclerotized integument with about 20 pairs of setae, including sI, s2 and five pairs 
of 7 setae. 

Tritosternum with base 30 yp, laciniae 50 w. Sternal shield (180 w x I14 p») 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 sii, 66 »; between sti and s#3, 84 w. Anal shield (45 X 45 »); par- and 
postanal setae subequal in length (about 20 »). 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 HT IV 
genu 50/42 48/42 34/30 45/30 
tibia 48/38 42/40 36/34 45/33 
tarsus 75/32 60/30 70/27 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; Dp. dorsum of idiosoma 
of male; E. venter of idiosoma of male. 


G. OWEN EVANS & W. M. TILL 


294 


THE BRITISH DEKMANYSSIDAE (ACART) 295 


HABITAT: Found in Britain on Muicrotus orcadensis Millais, Scotland, and on 
laboratory mice, Newcastle-upon-Tyne. Also recorded from South Africa (on 
Rattus rattus (Linn.) and the U.S.S.R. (on Rattus species, Mus musculus Linn., 
Apodemus species, Micromys minutus (Pallas), Muicrotus 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 tsabellinus : Bregetova, 1956, Opred. Faune SSSR. 61: 181, figs. 

Hirstionyssus arvicolae Zemskaya, 1955, in Bregetova et al., Opred. Faune SSSR. 59 : 366, figs. 
799-792. 


Female: Chelicera with segment I, 66 »; II, 135 w; movable digit 57 uw. Four 
pairs of gnathosomal setae with c.s. about 60 mw apart, hyp.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. 73a, 
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 w. Sternal shield (42 » x 158 yp) arched, 
strongly concave posteriorly, with three pairs of setae and two pairs of pores ; meta- 
sternals free with associated pores. Between st1, 84 w; between siz and s¢3, 90 pz. 
Surface of shield granular with reticulations antero-laterally. Genital shield (156 p 
x 138 ») granular, with one pair of setae. Anal shield (96 » x 87 yw) 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 fv, simple. Coxal spur 
formula (o—2—2-0). Length/width (in ») of leg segments : 


I II III IV 

genu 66/45 66/45 45/36 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; D. venter of idiosoma of 
male. 


296 G. OWEN EVANS & W. M. TILL 


Male: Chelicera with segment I, 48 »; II, 87 »; spermadactyl 50 p. Four 
pairs of gnathosomal setae with c.s. about 42 p apart, Ayp.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. 73C¢ ; 
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 stI, 50 »; between sf1 and s/f3, 
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 fv, 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 


HaBiItaT: Found in Britain on the weasel (Mustela sp.), Brentwood, Essex ; 
on Sorex araneus Linn., Meall Greigh, Perthshire, and on Microtus 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 U.S.S.R. 

The above description of the female is based on a specimen from Mustela sp. and 
that of the male on a specimen from Microtus 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 oncoxa 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 » apart, hyp.2 about 36 p 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. 74B) ; apotele two-tined. 

Dorsal shield (660 » * 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 


ST" was. Ye se ee 


‘ 


THE BRITISH DERMANYSSIDAE (ACARI) 


297 


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 w. Sternal shield (87 w X I4I p) 
rectangular, with three pairs of setae and two pairs of pores ; metasternals free with 
associated pores. Between sti, 80 w; between sti and sf3, 84 pw. Genital shield 
(132 w X 132 pw) granular except for the weakly sclerotized marginal zone, bearing 
one pair of setae. Anal shield (99 » X 96 pw) with paranal setae 30 pw, postanal seta 
33 p long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 74C. 
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 fv, simple. Coxal spur 
formula (o-1-1-0). Length/width (in p) of leg segments : 


I II III 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 ; Ayp.2 30-36 » apart. Deutosternum 
with about 15 rows of denticles (one to two denticles per row). Other features as 
in female. 

Dorsal shield (580-615 »# * 324 ») with 26 pairs of setae distributed as in Text-fig. 
75A; fifteen pairs of setae on the unsclerotized integument, including six pairs of r 
setae. Surface of shield as in female. 

Tritosternum with base 30 p, laciniae72 . Holoventralshield (444 x 120-138 ,) 
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 ». Surface of shield granular. Par- and post-anal setae subequal in length 
(about 30 «). Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 75B. 
Peritreme extends to middle of coxa I; peritrematal shields free posteriorly. 

Chaetotaxy of legs normal. Tarsus II with setae av, and pv, 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—1-1I-1). Legs bent, not suitable for measure- 
ment. 

Deutonymph : Chelicera with segment I, 38 »; II, 96 »; movable digit 40 x. 
Four pairs of gnathosomal setae with c.s. about 36-45 pu apart, hyp.2 about 26-30 uw 
apart. Deutosternum with 13-16 rows of denticles (one or two denticles per row). 
Other features as in female. 

Dorsal shield (408-444 » X 156-210 p) 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 r setae. 

Tritosternum with base 30 p, laciniae 57». Sternalshield (240-252 ~  II0-I20 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 st1, 70-72 pp; 
between sft and st3, 100-108 yp. Anal shield (48-54 » 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 fd,-,) and II (ad). Coxal spur formula 
(o-1-0-0). 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. 


(ACARI) 


THE BRITISH DERMANYSSIDAE 


AH, 


Vv" @ 
py ae | 
vib E | 
\ = P « 
; - ° —, 6 


See siae 


ZOOT. 14, 5. 


300 G. OWEN EVANS & W. M. TILL 


Hasitat: 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 (z1) 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 II-IV may have sclerotized ridges 
of varying degrees of development. Anterior spine of coxa II 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 peritremesabsent. Leg chaetotaxy normal for this instar. Thelarval 
stage is known only for a few species (Orvnithonyssus, 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 Io 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 asin 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 


t Dorsum of the idiosoma partially covered by discrete, subequal, podonotal and 


opisthonotal shields : : : STEATONYSSUS (p. 329) 
— Dorsum of the idiosoma with an gece pera or, if with two or more scutal elements, 

the pygidial shield is relatively minute . : ; : 2 
2 Genu and tibia III each with one postero-lateral seta (genu 28, i; tibia 

2—i, 21) ; tritosternum without hyaline denticulate border ; ‘ porose ’’ areas 


eat 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) 


302 


be 


G. OWEN EVANS & W. M. TILL 


Genu and tibia III each with two postero-lateral setae (genu 2—}, #—2; tibia 
2—}, #?—2); 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 ‘ie 
Genu IV with two postero-lateral setae (2—#, #—2 or wees ?—2) ; genital setae on 
integument ; ectoparasitesofreptiles  . OPHIONYSSUS (p. 335) 
Genu IV with one postero-lateral seta (2—#, 9—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 or 2-7,#-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—7—}—1) ‘ 2 

Genu and tibia III each with one postero-lateral seta, segments respectively with 
nine and eight setae ; tritosternum with a hyaline denticulate border 

MACRON 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 

‘Ss TEA 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—%, §—I) . ; ; ; ; ‘ : . : 2 
Genu IV with five setae (I—g, §—0) . . . , ; : 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 j 3 and opisthonotal seta J5 absent (Text-fig. 84a) . ° . : 4 
Podonotal seta 73 and opisthonotal seta ]5 present (Text- fig. 88a) . 
Pygidial shield with four pairs of setae ; podonotal setae 75 about equal in length to 
the distance between the bases of setaej75 andz5 Ornithonyssus bacoti (Hirst) (p. 324) 
Pygidial shield with three pairs of setae ; podonotal 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) ; 
Pygidial shield with five to seven pairs ofsetae ‘ : ‘ ; : . 7 
Pygidial shield with setae 74, S5 and Z5 subequal 
Steatonyssus periblepharus Kolenati (p. 333) 
Pygidial shield with setae Z4, 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 : ; ; : ‘ ; ; 9 


THE BRITISH DERMANYSSIDAE (ACARI) 303 


8 Unsclerotized integument of idiosoma with 16 pairs of setae (excluding 71), Text-fig. 


79A-B : Macronyssus kolenatii (OQudemans) (p. 314) 
- Miiecrotized Bie ament 2 *ahogocne markedly hypertrichous (Text-fig. 79c—D) 


Macronyssus flavus (Kolenati) (p. 310) 
9 Pygidial shield with five pairs of setae 


Macronyssus ellipticus (Kolenati) (p. 307) 


— Pygidial shield with six pairs of setae 10 


10 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 (Oudemans) (p. 329) 

— Tritosternum without denticulate, hyaline border. Pygidial shield convex an- 
teriorly ; its width not more than 14 times its length ; setae J4 well removed from 
the anterior margin of the shield . : § 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: Cams 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—}, 
}—1). 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. 76A) ; majority of deutosternal denticles compound (Text-fig. 76c). ; 2 
— Opisthonotal region of the dorsal shield with 12 pairs of setae ; deutosternal denticles 
simple, ina single file (Text-fig. 77B) ; . ‘ 
2 Sternal“ porose ’’ areas weakly defined, irregular (Text- fig. 768) ; peritreme reduced, 
extending to the level of the posterior third of coxa II ; podonotal region with 12 pairs 
of setae, 7I on integument (Text-fig. 764) . ; . 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 coxal ; podonotal 
region with 14 pairs of setae, 71 on shield (Text-fig. 81a) 
M. uncinatus (Canestrini) (p. 315) 
3 Bases of setae 26, Zt and Sr lying more or less in a straight line (Text-fig. 804) ; 
sternal ‘‘ porose’’ areas as in pon, > 80B ; lateral and opisthogastric cuticle 
moderately setose. ; M. kolenatii (OQudemans) (p. 312) 
— Seta 26, Z1 and S1 forming more or less the points of an equilateral triangle (Text-fig. 
78a) ; sternal “‘ porose’’ areas as in Text-fig. 788 ; lateral and opisthogastric 
cuticle densely setose . ; ; : ‘ : M. flavus (Kolenati) (p. 308) 


KEY TO MALES 


1 Spermadactyl extremely long, about as long as the second cheliceral segment 
M. ellipticus (Kolenati) ®. 300) 
— Spermadacty] obviously shorter in length than the second cheliceral segment 
2 Three pairs of dorsal marginal setae stout, thorn-like ; 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 
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. 80p) ; unsclerotized 
dorsal integument with about seven pairs of setae M. kolenatii (Oudemans) (p. 314) 


THE BRITISH DERMANYSSIDAE (ACARI) 


venter of idiosoma 


Fic. 76. Marconyssus ellipticus (Kolenati), female ; dorsal shield (a) ; 


(D—E). 


chelicera 


venter of gnathosoma (c) ; 


(B)s 


306 G. OWEN EVANS & W. M. TILL 


Macronyssus ellipticus (\Xolenati) 


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 p 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 » apart, Ayp.2 about 45 p» 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 »# * 348 ») with about 20 pairs of setae distributed as in Text-fig. 
760A; setae j1 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 pw. Sternal shield (105 pw X 174 p) 
with irregular punctate areas in the antero-lateral corners ; with three pairs of setae 
and two pairs of pores; metasternal setae free. Between st1, 80 »; between séI 
and sf3, 117 w. Genital shield (135 » x 102 ») with genital setae only. Anal shield 
(117 « X 100 pw) pear-shaped with strongly arched anterior margin ; paranal setae 
about 40 » 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—7, 
}—1). Coxa II with prominent anterior spine. Length/width (in p) of leg seg- 
ments : 


I Il III IV 
genu 120/60 102/66 84/57 144/60 
tibia 114/54 84/60 90/54 132/54 


tarsus 192/48 162/54 180/48 240/50 


Male: Chelicera with segment I, 30 »; II, 123 »; movable digit 36 wp 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 p apart, hyp.2 about 42 p apart. Other gnathosomal 
features as in female. 

Dorsal shield (648 » * 288 yp) with 26 pairs of setae. Sternito-genital shield 
(270 « * 138 yw) with four pairs of long sternal and a pair of short genital setae. 
Between st1, 63 »; between str and st3, 110 ». Ventro-anal shield with irregular 
lateral margins; bearing six asymmetrically arranged setae excluding the anals. 


Paranal setae 33 yw, postanal seta 42 » long. Peritreme extends to about middle of 
coxa II. 


THE BRITISH DERMANYSSIDAE (ACART) 407 


A 
A 
uy 
A 
N 
A 
A 


=> => 


Fic. 77. A. Macronyssus ellipticus (Kolenati), protonymph; dorsum of idiosoma. B-c. 
Macronyssus flavus (Kolenati), female ; venter of gnathosoma (B) ; chelicera (c). 


Chaetotaxy of legs as in female. Some relatively long, stout setae on femur I 


(ad, and #d,) and femur II (ad, and fd,_,)._ Legs bent and not suitable for measure- 
ment. 


Protonymph : Chelicera with segment I, 30 »; II, 114 »; movable digit 33 uz. 
Gnathosoma essentially as in female. Four pairs of gnathosomal setae with c.s. 
about 45 » apart, hyp.2 about 36 w apart. Deutosternum with 12 transverse rows of 
denticles (one to four denticles per row). Pedipalp (1-45-12) with two-tined apo- 
tele ; trochanter with conspicuous ventral keel. 

Podonotal shield (282 » X 216 p) with ten pairs of setae as in Text-fig. 774 ; 
setae 71 situated in front of shield. Pygidial shield (114 » x 144 pw) with anterior 


Margin convex ; bearing five pairs of setae. Unsclerotized integument of dorsum 
bears II pairs of setae. 


308 G. OWEN EVANS & W. M. TILL 


Tritosternum with base 30 p, laciniae 100 p. Sternal shield (180 p» X 126 p) 
with three pairs of setae and two pairs of pores. Between stI, 70 »; between st 
and sf3, 117 «. Anal shield (66 » x 66 ») with paranal setae 36 pu, postanal seta 45 u 
long. Peritreme extends to middle of coxa III. 

Chaetotaxy of legs normal. Coxa II with conspicuous anterior spine. Length/ 
width (in ,») of leg segments : 


I Il III IV 
genu 70/52 66/54 54/48 75/48 
tibia 66/45 51/48 48/45 68/45 


tarsus 120/40 go /40 90/40 135/40 


HasitaT: Ectoparasitic on Myotis myotis (Borkhausen), M. daubentonit (Kuhl) 
and M. natterert (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, hyp.2 about 36 » 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 » x 288 ») 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 », laciniae 105 p. 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 sft, 
54 »; between sfr and st3, 96 ». Genital shield (117 » x 84 ») with an overall 
reticulate pattern, bearing one pair of genital setae. Anal shield (go » x 96 p) with | 
par- and postanal setae about 24 » long. Sclerotization and chaetotaxy of opistho- _ 


Fic. 78. Macronyssus flavus (Kolenati). 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 (ACART) 


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 : 


I Il Ill IV 
genu 72/50 72/57 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 pu 
apart, Ayp.2 about 38 » apart. Other gnathosomal features as in female. | 

Dorsal shield (540 » X 252 ») 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 sf1, 63 p; 
between sf1 and sf3, 96 ». Ventro-anal shield (204 » x 66 ») with about fifteen 
setae excluding the anals; par- and postanal setae about 20 p» long. Chaetotaxy 
and sclerotization of opisthogaster as in Text-fig. 78D. Peritreme extends to middle 
of coxa 1; 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, fv, and mv on tarsus). Setae pd and pv, 
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 pz. 
Four pairs of gnathosomal setae with c.s. about 36 » apart ; Ayp.2 about 30 p apart. 
Pedipalp (1-4-5-12) with two-tined apotele. Other gnathosomal features as in 
female. 

Podonotal shield (240 » 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 w. Sternal shield (150 » x 100 p) with 
three pairs of setae and two pairs of pores. Between sf1, 54 »; between sf1 and s#3, 
105 #. Anal shield (42 » & 48 pw) with par- and postanal setae about 15 yp long. 


Fic. 79. AB. Macronyssus kolenatii (Oudemans), protonymph; dorsum (A) and venter 
(8) of idiosoma. c-p. Macronyssus flavus (Kolenati), protonymph; dorsum (c) and 
venter (bp) of idiosoma. 


THE BRITISH DERMANYSSIDAE (ACART) 


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 ,») of leg segments : 


I II II 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. pipistrelli 
Oudemans, 1906) et 1915, Arch. Naturgesch. 81 (A) : 165, figs. 128-134. 

Hirstesia britteni 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, Ayp.2 about 33 w apart. 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 » * 252 ») with 28 pairs of setae distributed as in Text-fig. 80A. 
Surface of shield reticulated. Unsclerotized integument of dorsum hypertrichous. 

Tritosternum with base 21 p, laciniae 84 ». Presternal area reticulated. Sternal 
shield (57 » X 129 pw) 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 stI, 45 »; between sti and st3, 78 ». Genital 
shield (90 « * 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 » * 75 ») with anterior margin almost straight ; paranal setae 21 pu, postanal 
seta 24 » long. Chaetotaxy and sclerotization of opisthogaster as in Text-fig. 80B. 
Peritremes extend to middle of coxae I; peritrematal shields free anteriorly, fused 
posteriorly with podal elements of coxae IV. 


Fic. 80. Macronyssus kolenatii (OQudemans). A. Dorsum of idiosoma of female; B. venter 
of idiosoma of female ; c. dorsum of idiosoma of male; pb. venter of idiosoma of male. 


313 


THE BRITISH DERMANYSSIDAE (ACARI) 


314 G. OWEN EVANS & W. M. TILL 


Chaetotaxy of legs normal except genu IV which has two ventral setae (2—}, 
}—1). Femur I with setae ad, and pd, long and stout. Coxa II with large, pointed 
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 »; 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 p» apart, Ayp.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 » X 270 «) with 28 or 29 pairs of setae distributed as in Text- 
fig. 8oc. Unsclerotized integument of dorsum with seven pairs of setae. 

Tritosternum with base 27 p, laciniae 90 ». Holoventral shield (380 » X 105 yp) 
with five pairs of setae in the sternito-genital region and about 20 setae in the ventral 
region. Between stI, 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 ,) 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 75/34 


Protonymph : Chelicera with segment I, 15 »; II, 72 »; movable digit 27 up. 
Gnathosomal features essentially as in female. Capitular setae about 30 pw apart, 
hyp.2 about 27 » apart. Pedipalp (1-4-5-12) with two-tined apotele. 

Podonotal shield (180 » x 147 ») with Io pairs of setae as in Text-fig. 79A ; setae 
jt 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 yp, laciniae 54 p. Sternal shield (110 » x 75 mu) with 
three pairs of setae and two pairs of pores. Between st1, 42 »; between sti and s#3, 
75 ». Anal shield (36 » * 39 ») with par- and postanal setae about 15 p 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 pd, 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 Prpistrellus. Recorded from 
P. pipistrellus (Schreber) in Britain and Europe and from P. kuAli (Kuhl) in Egypt. 


Macronyssus uncinatus (Canestrini) 


Letognathus uncinatus Canestrini, 1885, Acavofauna ital. 1:122; Berlese, 1889, Acar. Myr. Scorp. 
Ital. fasc. 53, no. 7. 

Liponyssus rhinolophi Oudemans, 1902, Tijdschr. Ned. dierk. 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 40 pu 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 w apart; hyp.2 about 45 w 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. 81A. 
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 p, laciniae 132 w. Sternal shield (84 » x 180 ») 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 st1, 72 4; between sf and st3, 105 pw. Genital shield 
(150 » X 90 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 p» long, postanal seta 60 » long. Chaetotaxy and sclerotization of opistho- 
gaster as in Text-fig. 81B.. 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—#?, 
3—1), Coxa II with pronounced anterior spine. Length/width (in yp) of leg 
segments : 


I II III 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 


316 G. OWEN EVANS & W. M. TILL 


Fic. 81. Macronyssus uncinatus (Canestrini), female; dorsum (A) and venter (B) of 
idiosoma. 


HasitatT: An ectoparasite of bats. Reported from Rhinolophus hipposideros 
(Bechstein) in Britain and from R. ferrumequinum (Schreber), R. euryale Blasius, 
Pipistrellus pipistrellus (Schreber), Myotis emarginatus (Geoffroy) and Miniopterus 
schreiberst (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. Arthrodia 


fee BiNlisk DERMWAN YSsIDAEB (ACA RT) Sii7 


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 fl 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 (/1), 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, /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—?, 4—2), tibia III (2—+, ?—2) and 
genu IV (2—#, 3—1). Rarely genu and tibia I with two ventrals only and genua IT 
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. pereira; no 
sclerotized ridges. 

Protonymph with dorsum of idiosoma provided with a podonotal shield bearing 
10 (73 lacking) or 11 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 asin female. Chaetotaxy of 
legs and pedipalps normal. Palptrochanter with or without ventral keel. 

The above concept of Ornithonyssus is considerably wider than that of Furman & 
Radovsky (1963). These authors restrict this genus to New World forms, except for 
four species, O. bacotr, 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 Ornithonyssus 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 Ovmithonyssus 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. VPeritremes 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 (Qudemans) (p.326) 
- Dorsal shield with setae 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 . : : . : : . 2 
2 Dorsum with setae s4 and S5 situated on the shield ; setae of J series relatively long, 

Jt reaching base of J2 (Text-fig. 83a). Sternal shield with three pairs of setae. 

Opisthogastric cuticle relatively raced setose Agar -fig. 83D). Palptibia with 13 

setae. Onsmallmammals . O. bacoti (Hirst) (p. 321) 
- Dorsum with setae s4 and S5 not on ‘the shield ; 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. 82B). 

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 // normal, not situated on 
atubercle. On bats (Eptesicus and Plecotus) 

O. pipistrelli (OQudemans) (p. 326) 

— Dorsal shield with ro or 11 pairs of setae in podonotal region. Coxae II and III 

with anterior setae normal. Palpfemur with seta p/ situated onatubercle . : 2 

Dorsum with seta S5 on shield. Setae of series relatively long, seta 74 reaching base 

of75. Holoventral shield with eight pairs of setae, excludinganals. Palptibia with 
13 setae. Onsmall mammals : ; : O. bacoti (Hirst) (p. 323) 

- Dorsum with seta S5 not on the shield. Setae of 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) 


to 


Ornithonyssus sylviarum (Canestrini & Fanzago) 


Dermanyssus sylviarum 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): 1 


Female: Chelicera with segment I, 20 »; II, 150 »; movable digit 36 yp long, 
with a dorsal “trough’’; both digits edentate; pilus dentilis and arthrodial 
processes absent. Four pairs of gnathosomal setae with c.s. about 50 p» apart, hyp.2 
about 36 » apart. Deutosternum with about ro denticles arranged in a single file. 
Corniculi membranous ; internal malae with inner processes fused, outer leaf-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. Ornithonyssus sylviarum (Canestrini and Fanzago). A. Dorsum of idiosoma of 
female; B. venter of idiosoma of female; c. venter of idiosoma of male; pb. dorsum of 
idiosoma of protonymph. 


ne 


319 


LHE BRITISH DERMANYSSIDAE (ACARI) 


320 G. OWEN EVANS & W. M. TILL 


Dorsal shield (534 # X 228 ») 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 p», laciniae 75 ». Sternal 
shield (36 » X 100 ,) reticulated and bearing two pairs of setae and two pairs of 
pores ; sf3 and sf4 on unsclerotized cuticle. Between st1, 66 ~; between str and 
st3, 52 «. Genital shield (132 » X 60 ») tapering, with genital setae only. Anal 
shield (96 « & 66 «) with par- and postanal setae subequal, about 30 » long. 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—7, »7;—2), tibia III (2—4, 
7—2) and genu IV (2—}, #—1). In addition, one of the ventral setae may be 
lacking on genu I (2—;#,, }—2), tibia I (2— jy, }—2) and genu II (2—#, 57;—2) in 
some specimens. Coxa II without anterior spine. Length/width (in p) of leg 
segments : 


I Il 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 »; 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 » apart, hyp.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 pw) with chaetotaxy as in female, except that it may 
incorporate one or more marginal setae in the opisthonotal region. The short postero- 
lateral seta (x) varies in position ; it may be posterior to seta 74 as in the female, 
level with it or anterior to it. Unsclerotized integument of dorsum hypertrichous. 

Tritosternum with base 24 p, laciniae 57 ». Holoventral shield (360 p» X 93) 
narrow, with six pairs of setae excluding the anals. Between sti, 54 »; between 
stt and st3, 70 ». Par- and postanal setae subequal, about 24 » long. Chaetotaxy 
and sclerotization of opisthogaster as in Text-fig. 82c. Peritreme extends to middle 
of coxa Il ; 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 II III 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 »; II, 105 »; movable digit 27 u. 
Gnathosomal featues generally as in female. Four pairs of gnathosomal setae with 
c.s. about 45 p apart, hyp.2 about 30 mw apart. Deutosternum with 8 denticles 
arranged in a single file. Pedipalp (1-4-5-12) with two-tined apotele. 

Podonotal shield (162-174 » X 144-156 pu) with ten pairs of setae distributed as in 
Text-fig. 82p. Pygidial shield (50 » x 100 pu) 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 pw. Sternal shield (120 » X 96 p») with 
three pairs of setae and two pairs of pores. Between sfI, 63 »; between sfI and sf3, 
75 ». Genital setae very short, about half as long as the sternal and the four pairs of 
opisthogastric setae. Analshield (42-45 » xX 50-54 ») 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 ») of leg segments : 


I II III IV 

genu 40/36 36/36 32/32 40/30 
tibia 42/33 33/33 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) 


Letognathus bacoti 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 yp 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. 83£). 
Four pairs of gnathosomal setae with c.s. about 57 w apart, dyp.2 about 42 pw 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 » xX 198 yp) with 18 pairs of setae distributed as in Text-fig. 834. 
Unslcerotized integument of dorsum hypertrichous. 

Tritosternum with hyaline denticulate membrane; base 66 yp, laciniae 108 wp. 
Sternal shield (50 » X 120 yp) reticulated, with three pairs of setae and two pairs of 
pores; metasternals free. Between st1, 70 »; between sf and st3, 66 uw. Genital 
shield (174-180 » X 70-75 p) tapering, with genital setae only. Anal shield (123- 
135 » X 78-87 ») with paranal setae 42 pu, postanal seta 45 w long. Chaetotaxy and 


G. OWEN EVANS & W. M. TILL 


THE BRATISH DERMANYSSIDAE (ACART) 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—#, 4—2), tibia III (2—+, 2—2) and 
genu IV (2—#?, 3—1). Coxa II with small anterior spine. Length/width (in 1) of 
leg segments : 


I 1 II 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 »; 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 pw apart, hyp.2 about 38 p apart. 
Corniculi membranous ; internal malae blade-like, free. Pedipalp as in female, but 
palptrochanter without ventral keel; seta #/ on palpfemur situated on a small 
tubercle. 

Dorsal shield (528 » x 270 ») 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 I00 pz) 
narrow, with eight pairs of setae excluding the anals. Paranal setae 30 yp, 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 p) of leg segments : 


I II III IV 
genu 7540 60/48 60/45 72/30 
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 (hyp.1I and 2) with hyp.2 about 
24 mw apart. Deutosternal teeth not visible. Corniculi degenerate. Pedipalp 
(o-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. Ovnithonyssus bacoti (Hirst). A. Dorsum of idiosoma of female; B. chelicera of 
male; c. dorsum of idiosoma of male; p. 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 (Jv1 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 x. 
Gnathosomal features generally as in female. Four pairs of gnathosomal setae with 
c.s. about 36 » apart, Ayp.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 ») with ten pairs of setae as in Text-fig. 844 ; 
pygidial shield (75 » X 123 ») with four pairs of setae. Unsclerotized integument of 
dorsum with 16 pairs of setae. | 

Tritosternum bipartite, with hyaline denticulate membrane ; base 27 yp, laciniae | 
75 . Sternal shield (123 » X 90 ») with three pairs of setae and two pairs of pores. 
Between sti, 50 »; between str and st3, 90 w. 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, 
Jvit-2, Jv5 and Zv2 (Text-fig. 848). Peritreme extends to anterior margin of coxa 
III. 

Chaetotaxy of legs normal. Length/width (in ,») of leg segments : 


I II Ill 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. 84c), 
but not visible in the female (Text-fig. 84p). 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. 


biG. 84. Ornithonyssus bacoti (Hirst). A. Dorsum of idiosoma of protonymph ; B. venter 
of idiosoma of protonymph ; c. venter of idiosoma of deutonymph (male) ; pb. venter of 
idiosoma of deutonymph (female). 


THE BRITISH DERMANYSSIDAE (ACART) 


gen... 


a 


325 


326 G. OWEN EVANS & W. M. TILL 


Chaetotaxy of legs as in female. 


HABITAT: QO. dacoti, 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 rattus (Linn.), but is also known to bite man 
(Evans et al., 1961). 


Ornithonyssus pipistrelli (Qudemans) 


Liponyssus musculi (Koch) Oudemans, 1902, Tijdschr. Ned. dierk Vereen. (2) 8:17, pl. 1, figs. 
1-2. 

Liponyssus pipistrelli Oudemans, 1903, Tijdschr. Ent. 46: 111% [nom. nov. pro Liponyssus 
muscult (Koch) Oudemans, 1902]. 


Female: Chelicera with segment I, 54 »; II, 174 »; movable digit 50 », edentate 
with dorsal “trough’’; fixed digit with terminal hyaline boss. Four pairs of 
gnathosomal setae with c.s. about 66 » 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 » & 294 ») narrow, reticulated, with 20 pairs of setae distributed 
as in Text-fig. 85a. Unsclerotized integument of dorsum hypertrichous. 

Tritosternum with denticulate hyaline border ; base 54 pw, laciniae 110 wp. Sternal 
shield (75 « X 144 p») reticulated, with three pairs of setae and two pairs of pores ; 
metasternals free with associated pores. Between sf1, 87 »; between sf and s#3, 
93 ». Genital shield (180 » x go ») tapering, with genital setae only. Anal shield 
(75 » & 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—?, 7—2), tibia III (2—}, }—2) and 
genu IV (2—4, #}—1). Coxa II with prominent anterior spine. Length/width (in p) ~ 
of leg segments : 


I II III 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 »; II, 126 »; movable digit edentate, 42 p 
long ; spermadactyl relatively short, 54 » from base of movable digit. Four pairs 
of gnathosomal setae with c.s. about 51 » apart, hyp.2 about 39 » apart. Deuto- 
sternum with ten denticles arranged in a single file. Corniculi membranous, internal 
malae leaf-like. Tectum capituli and pedipalps as in female. 


FIG. 85. Ornithonyssus pipistrelli (Oudemans). A. Dorsal shield of female ; 8. dorsum of 
idiosoma of male; c. venter of idiosoma of female; p. venter of idiosoma of male. 


327 


THE BRITISH DERMANYSSIDAE (ACART) 


328 - G. OWEN EVANS & W. M. TILL 


Dorsal shield (696 » 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 p, laciniae 78 ». Holoventral shield (588 » x 123 p) 
narrow, reticulated, with 19 setae excluding the anals. Between s¢1, 70 1; between 
stt and sf3, 100 ». Paranal setae about 33 » long. Chaetotaxy and sclerotization of 
opisthogaster as in Text-fig. 85p. 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 ,) 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 (B) 
of idiosoma. ‘ 


foe BREIISH 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 pw apart, Ayp.2 about 30 pw 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 p) with 11 pairs of setae (Text-fig. 864) ; pygidial 
shield (66 » x 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 w. Sternal shield (150 » X 117 p) with 
three pairs of setae and two pairs of pores. Between si, 66 » ; between séz and s/3, 
96 pw. Anal shield (60-63 » xX 66-69 ») with paranal setae about 30 yp, postanal seta 
about 36 » long. Opisthogaster with a pair of genital and five pairs of opisthogastric 
setae distributed as in Text-fig. 86B. Peritreme extends to posterior third of coxa ITI. 

Chaetotaxy of legs normal. Length / width(in yw) of leg segments : 


I II III IV 
genu 63/39 54/45 48/40 60/36 
tibia 63/36 54/42 51/38 60/33 


tarsus 117/33 87 /33 84/32 110/30 


HABITAT: Found in Dorset, England, on Eftesicus 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. piprstrellr. 


Genus STEATONYSSUS Kolenati 


Steatonyssus Kolenati, 1858, Wien. ent. Monatschr.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 (zr) absent. Opisthonotal shield normally with seven pairs of setae, 
but this number may be reduced by the absence of J/5 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 I1 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—tleg I normal, leg II normal, leg II] normal except genu III 
(2—}, {—2) and tibia III (2—4, 3—2) and leg IV normal except genu IV (2—#, 
—1). 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 (72 lacking) or 29 pairs of setae. 
Podonotal shield with Io or I1 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, /5 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. 


ioe 


KEY TO FEMALES 


1 Sternal setae 1 very short, not more than one-half the length of sternal setae 2 (Text- 
fig. 878) ; peritrematal shield entire anteriorly 
S. periblepharus Kolenati (p. 330) — 
— Sternal setae 1 and 2 long, subequal in length ; anterior portion of peritrematal 
shield interrupted. : F ‘ : ‘ S. occidentalis Ewing (p. 334) 


Steatonyssus (Steatonyssus) periblepharus Kolenati 


Steatonyssus periblepharus Kolenati, 1858, Wien. ent. Monatschr.2:6; et 1859, 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 »; II, 168 »; movable digit 54 »; both 
digits edentate. Four pairs of gnathosomal setae with c.s. about 45 » apart, hyp.2 
about 33 » 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. 


THE BRITISH DERMANYSSIDAE (ACARI) 


Fic. 87. Steatonyssus periblepharus Kolenati. a. Dorsum of idiosoma of female ; B. venter 
of idiosoma of female ; c. dorsum of idiosoma of male; D. venter of idiosoma of male. 


_ ZOOL. 14, 5. 


332 G. OWEN EVANS & W. M. TILL 


Podonotal shield (294-312 « X 252-276 p) bears II pairs of setae distributed as in 
Text-fig. 87a. Opisthonotal shield (348-396 p x 180-198 ,») with seven pairs of 
setae. Seta _/1 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 yw laciniae 102 up. 
Sternal shield (45-57 » X 130-144 p) 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 p; 
between sft and st3, 70-72 yw. Sternal seta I relatively short, not more than half as 
long as sf2. Genital shield (144-156 » X 75-93 ») tapering, with genital setae only. 
Anal shield (114-135 » 80-94 ») with par- and postanal setae subequal, about 40 p 
long. Integument of opisthogaster bears numerous setae, those near the margin 
being relatively longer (65 ) and stouter than the others (Text-fig. 87B). Peritreme 
extends to middle of coxa II ; anterior part of peritrematal shield continues to middle 
of coxa I; posterior part fused with podal shields of coxa IV. 

Chaetotaxy of legs normal except genu III (2—}, {—2), tibia III (2—+, ?—2) and 
genu IV (2—?, }—1). Coxa II with prominent anterior spine. Length/width (in p) 
of leg segments : 


I II 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 pw apart, Ayp.2 about 36 pw 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. 87¢. 
Surface of shield reticulated. Unsclerotized integument of dorsum hypertrichous. 

Tritosternum with base 21 p, laciniae 66 ». Holoventral shield (420 » x 105 p) 
narrow, bearing six pairs of setae excluding the anals ; metasternal setae on integu- 
ment beside shield. Between st1, 48 »: between st1 and st3, 75 w. Par- and post- 
anal setae subequal, about 24 p» 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 yp) of leg 
segments : 


EERE BRICISH 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 90/36 90/33 120/30 


Protonymph : Chelicera with segment I, 20 »; II, 100 »; movable digit 30 pn. 
Four pairs of gnathosomal setae with c.s. about 33 » apart ; hyp.2 about 27 p apart. 
Pedipalp (1-4-5-12) without keel on trochanter. 

Podonotal shield (180 » X 150 p) with I1 pairs of setae as in Text-fig. 884 ; 
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 sii, 60 » ; between si and si3, 
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. 888). Peritreme extends nearly to middle of coxa III. 


Fic. 88. 


Steatonyssus periblepharus Kolenati, protonymph; dorsum (A) and venter (B) 
of idiosoma. 


334 G. OWEN EVANS & W. M. TILL 


Chaetotaxy of legs normal. Length/width (in ,) of leg segments : 


I Il Ill 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 


Hapirat: 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) : Io, figs. 
Steatonyssus occidentalis: Radovsky & Furman, 1963, Ann. ent. Soc. Amer. 56: 272, figs. 
Till & Evans, 1964, Bull. Brit. Mus. (nat. Hist.) Zool. 11: 559, figs. 


Female: Chelicera with segment I, 27 »; II, 140 1; 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- 
niculi membranous ; internal malae simple. Tectum capituli tapering, anterior 
margin weakly serrated. Pedipalp (2-5-6-13) with two-tined apotele; palp- 
trochanter with prominent antero-ventral keel. 

Podonotal shield (288 » * 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 » * 132 p) 
reticulated, with thickened posterior border ; bearing three pairs of setae and two 
pairs of pores; metasternal setae free. Between sti, 68 1; between sti and s#3, 
72 w. Genital shield (144 » X 93 p») tapering, with genital setae only. Anal shield 
(120 » * 96 w) with par- and postanal setae about 40 p» long. Integument of 
opisthogaster hypertrichous (Text-fig. 89B). 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 of legs normal except genu III (2—#, ?;—2), tibia III (2—}, }—2) and 
genu IV (2—%, {—1). Coxa II with anterior spine. Length/width (in ,) of leg | 
segments : 


I Il Ill IV 
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 


TRE BRATISI DERMANYsSI DAE (ACARI) 


Fic. 89. Steatonyssus occidentalis (Ewing), female ; dorsal shields (A) ; ventral shields (s). 


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 
occurs on bats of the genera Eptesicus, Myotis, Tadarida, Corynorhinus and Lasiurus. 


Genus OPHIONYSSUS Meégnin 


Ophionyssus Mégnin, 1884, C.R. Soc. Biol. Paris 35: 614. 
Serpenticola Ewing, 1923, Proc. U.S. nat. Mus. 62 (art. 13) : 6. 
Sauronyssus Sambon, 1928, Ann. trop. Med. 22 : 106. 
Neoliponyssus Ewing, 1929, Manual of External Parasites : 187. 
Oudemansiella Fonseca, 1948, Proc. zool. Soc. Lond. 118 : 269. 


Type: Dermanyssus natricis Gervais, 1844. 


336 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 /1 and 
J2,and with a discrete pygidial shield lacking setae ; the third consisting of a single 
dorsal shield incorporating 10 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—%, ?—2) or (2—#, 7—2), femur II (1—3—1), 
genu II (2—?, #—2), tibia II (2—+, ?—2), femur III (1—}—0), genu III (2—4, — 
i—2), tibia III (2—}, #—2), genu IV (2—}, #—2) or (2—}, #—2) and tibia IV 
(2—}, }—2). Coxa II without an anterior spine. All ambulacra with a pair of 
claws. 

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, J4 and Z5) or 11 pairs (pygidial shield 
with three pairs, J/4,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—s, 5—1). 

Strandtmann & Wharton (1958) recognize Sauronyssus 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 
75 from the entire dorsal shield of certain Sauronyssus suggests that a reduction 0 
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, /3 and /4. In the 
males of both Opiionyssus 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 Opionyssus. The distinction based on 
the host preferences of Ophionyssus (snakes) and Sauronyssus (lizards) is not valid. 
Ophionyssus monodi (Hirst) recorded from Acanthodactylus scutellatus and Opmonys- 
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 QO. 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 


I Dorsal sclerotization consisting of a podonotal shield, two pairs of mesonotal 


scutellae and a pygidial shield. : ; : O. natricis ghia (p. oT 
Dorsal sclerotization consisting of a single dorsal shield. 
2 Tibia IV with two postero-dorsal setae (2—4, 2—2). Dorsal ela ay ten panes of 
setae in the podonotal region . ‘ : . O. saurarum (Oudemans) (p. 342) 
— Tibia IV with three postero-dorsal setae (2—+1, 3—2). Dorsal shield with 11 pairs of 
setae in the podonotal region . : ‘ ; O. lacertinus (Berlese) (p. 345) 


KeEy To MALES 


1 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 & Cae Fer emur III with ventralsetanormal . : 2 
2 Tibia IV with three postero-dorsal setae ey 32). 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 w; movable digit 45 w. Both 
digits edentate, pilus dentilis and dorsal seta absent (Text-fig. g1c). Four pairs of 
gnathosomal setae with c.s. about 60 uw apart, hyp.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 


Fic. 90. Ophionyssus natricis (Gervais). A. Dorsum of idiosoma of female; B. venter 


idiosoma of female; c. dorsum of idiosoma of male; pb. venter of idiosoma of male. — 


PRE BRITISH DEKMANYSSIDAE? (ACARI) 339 


fimbriated. Pedipalp (2—-5-6-13) with two-tined apotele ; palptrochanter with a 
shallow, inconspicuous, weakly sclerotized keel. 

Podonotal shield (300 » x 276 ») with ten pairs of setae distributed as in Text-fig. 
goA ; two pairs of mesonotal scutellae lie posterior to the podonotal shield. Surface 
of podonotal shield granular, weakly reticulated antero-laterally. Pygidial shield 
(54 » X 45 pw) without setae. Unsclerotized integument of dorsum shows marked 
hypertrichy. 

Tritosternum with hyaline membrane ; base 54 p, laciniae 96 pw. Sternal shield 
(45 » X 114 p) 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 
stI1, 42 w; between sti and s#3, 102 pw. Genital shield (about 135 » x 45 ») 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, #—2), 
genu III (2—?, 2—2), genu IV (2—#, #—2) ; tibia I (2—3, #—2), tibia III (2—1+, 
#—2). Coxa II without anterior spine. Length/width (in y) of leg segments : 


A 


Fic. 91. Ophionyssus natricis (Gervais). A. Venter of gnathosoma of female ; B. chelicera 
of male; c. chelicera of female. 


340 G. OWEN EVANS & W. M. TILL 


I II ll IV 
genu 75/45 60/48 60/45 72/40 
tibia 75/40 60/45 54/40 66/36 


tarsus 123/36 96/40 90/36 110/33 


Male ; Chelicera with segment I, 18 » ; II, 96 w. 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 wapart. 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 yp, laciniae 70 ». Sternito-genital 
shield (204 » & 81 mw) with str and sf2 on the shield, s¢3, st4 and the genital setae on 
the cuticle lateral to the shield. Between sf1, 51 » ; between sf1 and st3, 84 ». Anal 
shield (66 »% x 60 ») with par- and postanal setae about 27 w 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 ,) of leg segments : 


I II Ill 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 p. 
Four pairs of gnathosomal setae with c.s. about 39» apart, hyp.2 about 27 p 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 » * I9g2 p) with eleven pairs of setae distributed as in Text- 
fig. 922A; setae s6 and r2, r3, 75 and 76 are situated on the unsclerotized integument. 
Pygidial shield (63 » 75 p) bears three pairs of setae. 

Tritosternum with hyaline border ; base 20 p, laciniae 75 ». Sternal shield (110 » 
* 78 w) with three pairs of setae and two pairs of pores. Between st1, 48 » ; between 
sti and st3, 80 », Anal shield (48 » x 48 ») with paranal setae 21 p», postanal se 
24 » 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—1). Length/width (in ,) of leg segments : 


I II Ill IV 
genu 42/36 42/36 40/33 48/30 
tibia 48/33 42/36 36/30 48/30 


tarsus 84/30 60/30 60/27 66/27 


THE BRITISH DERMANYSSIDAE (ACARI) 


341 


Fic. 92. a-B. Ophionvssus natricis (Gervais), protonymph ; dorsum (a) and venter (B) 


of idiosoma. c—p. Ophionyssus sauvarum (Oudemans), protonymph; dorsum (Cc) and 
venter (D) of idiosoma. 


342 G. OWEN EVANS .& W. M. TILL 


Hasirat: 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 saurarum Oudemans, 1901, Tijdschr. Ned. dierk. Vereen. (2) 7 : 66. 
Sauronyssus saurarum: Bregetova, 1956, Opred. Faune SSSR. 61: 218, figs. 


Female: Chelicera with segment I, 36 »; II, 147 1; movable digit 40 ». Both 
digits edentate, pilus dentilis and dorsal seta absent.Four pairs of gnathosomal setae 
with c.s. about 60 » apart, hyp.2 about 42 w 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 ») 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 setae 74. Unsclerotized integument of dorsum markedly 
hypertrichous. 

Tritosternum with denticulate, hyaline border; base 45 yp, laciniae 78 p long. 
Sternal shield (42 » x 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 sf1 and st3, 96 ». Genital shield (about 150 » Xx 60 yp) tapering ; 
genital setae on the unsclerotized integument. Anal shield (105 p x 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 II; 
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 pd,) and 
femur II (ad, and fd,-,)._ Coxa II without anterior spine. Length/width (in p) of 
leg segments : 


I Il III 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 papart. 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- 


THE BRITISH DERMANYSSIDAE (ACARI) 343 


AS 


= 


Fic. 93. Ophionyssus saurarum (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. 


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 10 or 12 pairs of setae. 

Tritosternum with base 24 », 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 m) 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 pd,-,)._ 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 Ill IV 
genu 70/40 60/45 60/40 75/40 
tibia 70/36 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 uw. 
Four pairs of gnathosomal setae with c.s. about 38 » apart, hyp.2 about 27 p» apart. 
Pedipalp (1-4-5-12) with two-tined apotele. Other gnathosomal features as in 
female. 

Podonotal shield (213 » x 180 ») with eleven pairs of setae distributed as in Text- 
fig.g2c. Surface of shield reticulated. Pygidial shield (84 » x 81 yp) bears two pairs 
of setae. 

Tritosternum with hyaline border; base 32 yp, laciniae 60 pw. Sternal shield 
(126 » & 74 pw) with three pairs of setae and two pairs of pores ; between sf1, 48 pu 
between sti and st3, 87 ». Anal shield (36 » x 40 ») with par- and postanal setae 
about 20 » 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,-,), III (ad,) and IV (ad,-,). Length/width (in ,) of leg” 


segments : 


I Il Ill IV 
genu 48/33 42/36 45/33 57/30 
tibia 54/33 45/36 45/33 57/32 


tarsus 102/30 70/30 72/28 75/27 


Hapitat: 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. 


TRE BRIPTISH DERMANYSSIDAE (ACARI) 345 


Ophionyssus lacertinus (Berlese) 
Leiognathus lacertinus Berlese, 1892, Acar1, Myr. Scorp. Ital. fasc. 70, no. 3, figs. 


Female : Chelicera with segment I, 30 »; II, 180 1»; movable digit 50 wp. Four 
pairs of gnathosomal setae with c.s. about 66 pw apart, hyp.2 about 38 wp 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 » X 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 yp, laciniae 80 p. Sternal shield 
(40 » X 120 p) with two pairs of setae and two pairs of pores. Between sfI, 54 p; 
between sf and s#3, 100 pw. Genital shield (144 » X 54 p) tapering. Anal shield 
(106 » X 84 p) with paranal setae about 38 p, postanal seta about 30 w long. Peri- 


Fic. 94. Ophionyssus lacertinus (Berlese). A. Dorsal shield of female; 8. 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 ,») 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 p apart, 
hyp.2 about 30 wapart. Deutosternum with eight denticles arranged in a single file. 
Other gnathosomal features as in female. 

Dorsal shield (516 « X 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 yp, laciniae 54 ». Sternito-genital shield (216 » x 75 p) 
reticulated, with s¢1 and s¢2 on the shield, s¢3, s¢4 and genital setae on the unsclerotized 
integument. Between sfI, 45 »; between sti and st3, 90 w. 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. 948). 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 117/32 80/33 96/33 105/30 


HaBITAT: 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- 


ment enormously elongated, slender ; digits minute, weakly dentate. Pilus dentilis, 
dorsal seta and arthrodial processes absent. Male chelicera with second segment 


fTHE 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 Lifonyssoides. 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 (o—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. I4, 5. 23 


348 G. OWEN EVANS & W. M. TILL 


Protonymph : Active, feeding instar. Chelicerae, basis-capituli and hypostome as 
in the female. Pedipalpal chaetotaxy (1-4—5—12), rarely genu and tibia deficient ; 
palptrochanter without medio-ventral keel. 

Dorsal sclerotization comprising well developed podonotal shield with marked 
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 (Liponyssotdes) 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, spermadactyl long, 
grooved, completely fused with movable digit. Chaetotaxy of venter of gnathosoma 
normal ; pedipalp with five free segments and a two-tined apotele. Chaetotaxy of 


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 z1 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 (13-14 
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 


, i oe 


J 


§ 
; 


; 
7 


PRE BRILTISH DEKMANYSSIDAE (ACART) 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 pd series on 
tibia I each comprises only two setae, and the ad series on tibia IV and fd series on 
genu IV also only two setae. Tarsi III and IV in male with seta pv, 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. 


KEY TO FEMALES 


I Dorsal shield with at least eight pairs of setae in the podonotal region (74 present) ; no 
conspicuous pore on shield posterior to seta s4. Pe aee with 12 setae. Meta- 
sternal setae usually present . : : 3 2 

— Dorsal shield with seven pairs of setae in the Podanoral ea (7 a pre ; a con- 
spicuous pore present on each side of the shield posterior to seta s4. Palptibia with 


7or8setae. Metasternalsetaeabsent . : : D. alaudae (Schrank) (p. 359) 
2 Dorsal shield with four pairs of setae m j series (/2 present):;, with 9 or 11 pairs 

of setae in podonotal region (setae 26 present) : ‘ : : : 3 
— Dorsal shield with three pairs of setae in J series (J2 gene) ; with eight So of 

setae in podonotal region (z6 absent or on unsclerotized cuticle) : : 4 


Se Eetiureme extends at least to middle of coxa Il. Setae j1.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 sti 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 74 
on dorsal shield considerably longer than setaj4. Tibia I with two ventral setae 
D. quintus Vitzthum (p. 361) 
-- QOpisthogaster without arc of stout spines. Anal shield with maximum width 
approximately equal to.its length. Setae 22, z4 and 74 on dorsal shield subequal. 
Tibia I with three ventral setae . ; : D. hirundinis (Hermann) (p. 356) 


KEy TO PROTONY MPHS!9 


1 Podonotal shield with nine pairs of setae, 71 on the shield ; opisthonotal region of 


dorsum with J series complete (J1—J5) ; 2 

~ Podonotal shield with seven or eight pairs of ao a1 off fhe aya Bpicthaatd 
region of dorsum with setae J2 lacking . 3 

2 Posterior extension of podonotal shield ere aie eaceded saat evel of 
setae J1 (Text-fig. 974) . : D. gallinae (De Geer) (p. 353) 

— Posterior extension of podonotal opicld een: 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 Dugeés, 1834, Ann. Sct. nat. zool. (2) 1: 18. 

Dermanyssus evotomydis Ewing, 1933, Proc. U.S. nat. Mus. 82 (art. 30): II. 


[Por detailed synonymy see Oudemans (1936), p. 299.] 


Female ; Chelicera with segment I, 45 »; II, approximately 276 » ; digits minute, 
weakly dentate (Text-fig. 96c). Four pairs of gnathosomal setae with c.s. about 57 » 
apart; Aypf.2 about 45 » apart; hAyp.1 relatively stout. Deutosternum with 11 
denticles arranged in single file. Corniculi membranous, with sclerotized bars 
(Text-fig. 968). Tectum capituli elongate, triangular, with smooth margin. Pedipalp 
(2—4-6-12) with two-tined apotele. 


THE BRITISH DERMANYSSIDAE (ACARI) 351 


Dorsal shield (756 » xX 300 ») with 15 pairs of setae distributed as in Text-fig. 95a ; 
setae /5 situated posterior to the shield ; setae J/3 and J/4 sometimes asymmetrically 
arranged ; /3 may be unpaired. Surface of shield reticulated. 

Tritosternum biramous, with denticulate, hyaline margin. Sternal shield (24 p x 
150 ») reticulated, with two pairs of setae and one pair of pores ; setae st3, metasternal 
setae and two pairs of pores situated on the unsclerotized cuticle. Between sf1, 
78 »; between sti and st3, 88 ». Genital shield (198 » X 126 yp) reticulated ; with 
genital setae only. Anal shield (140 » X 123 »; maximum width 150 yp) with 
paranal setae 36 u long, postanal seta 20 » long. Chaetotaxy and sclerotization of 
opisthogaster as in Text-fig. 95B. 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 II-IV: 


: if III IV 
femur 2—34g—2 2—3—I t—3_9 i 
Su we, 2, 2, 2 2 2 
ea 2—F> Ti 2 ee ae ame ie 2—4, oO a 
tibia 2—%, 7-2 I—j, 7—I I—i, 21 i—i, 21 


Coxa II without anterior spine. Length/width (in p) of leg segments : 


I II Ill 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 w; 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 w apart; Ayp.2 about 45 p apart ; 
hyp.t normal. Other gnathosomal features as in female. 

Dorsal shield (636 » X 276 yz) 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 Z1 is also on the shield. 

Tritosternum with hyaline denticulate margin. Holoventral shield (528 pw 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 sf3, 110 wp; 
paranal setae 27 p, postanal seta 18 » 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. 964A). 

Chaetotaxy of legs as in female ; seta pv, on tarsi II] and IV modified to form a 
backwardly directed tooth-like projection (Text-fig. 96£). Coxa II without anterior 
spine. Length/width (in p) of leg segments : 


G. OWEN EVANS & W. M. TILL %\ 


352 
I Il Ill IV 

genu 87/54 72/48 66/45 81/45 

tibia go/50 70/45 72/42 87/42 

tarsus 126/42 102/40 108/36 135/36 


rPPrrr errr Pp pep -p 


@ 


= 


bic. 96. Dermanyssus gallinae (De Geer). a. Venter of idiosoma of male; B. venter of a 
gnathosoma of female; c. chelicera of female; bp. chelicera of male; £. 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 p» apart, 
hyp.2 about 33 mw apart. Pedipalps with normal chaetotaxy (1-4-5-12) for this 
instar. 

Podonotal shield (276 » X 120 w»; maximum width 204 ») with a short posterior 
prolongation as in Text-fig. 974 ; bearing nine pairs of setae. Opisthonotal region 
of dorsum with / series complete. 

Tritosternum with base 45 p, laciniae 70 ». Sternal shield (168 » x 108 p) with 
three pairs of setae and two pairs of pores. Between sf1, 50 w; between sfz and sf3, 
120 wp. Genital setae absent. Anal shield (66 » X 66 »; maximum width 75 yp) 
with paranal setae 15 p», 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 ITV which sometimes lacks the 
postero-lateral seta (1—+, #—0o/1). Coxa II without anterior spine. Length/ 
width (in ») of leg segments : 


I II III 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 


& 

HapitatT: 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 [Serimus canarius (Linn.)]. 


Dermanyssus chelidonis Oudemans 


Dermanyssus hirundinis: Berlese, 1889, Acar., 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 yw), 
weakly dentate. Four pairs of gnathosomal setae with c.s. about 70 pw apart, hyp.2 
about 57 » 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 /5 off the shield. Surface of shield patterned as in Text-fig. 98a. 

Tritosternum with denticulate hyaline margin. Sternal shield (45 w X 150 p) 
with two pairs of setae and two pairs of pores; st3 and metasternals free on un- 


354 


FiG. 97. 


G. OWEN EVANS & W. M. TILL 


Dorsum of idiosoma of protonymphs of four species of Dermanyssus ; 
(a); D. chelidonis (B); D. alaudae (c); D. hirundinis (D). 


D. gallinae 


‘ 


tne BRITISH DEKRMANYSSIDAE (ACARI) 355 


sclerotized cuticle. Between sii, 100 w; between sti and sf3, 93 uw. Genital shield 
(222 w X 144 p) with genital setae only. Anal shield (168-174 » X 150 w; maxi- 
mum width 180-210 ») with paranal setae 40 p, postanal seta 27 uw. 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 IJ-IV : 


I II III lV 
femur 2—3z—2 2—3—I I—3—o0 — 
3 2 2 2 2 2 2 
Spt 2—F> tT 2 2-=7» a re Za eae Za) pa 
tibia 2—3%, 7-2 i—i,2—r 1—1, 2-1 a 


Coxa II without anterior spine. Length/width (in yp) of leg segments : 


I II Ill IV 
genu 120/66 108 /60 114/54 135/54 
tibia 120/60 102/60 108/54 132/46 


tarsus 198/54 162/48 174/45 222/48 


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 / series of setae complete. 

Habitat: In the nests of swallows (Hirundo rustica Linn.) in England and Italy. 


Dermanyssus hirundinis (Hermann) 


Acarus hirundinis Hermann, 1804, Mémoire A ptérologique Strasbourg : 83. 
Dermanyssus hirundinis: 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 m» apart, Ayp.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 hirundinis (Hermann), female ; dorsum (a) and venter (B) of idiosoma. 


TRE BRITISH DEKMANYSSIDAE (ACARI) 357 


Dorsal shield (680 » X 294 ») with 11 pairs of setae distributed as in Text-fig. 99A ; 
setae 71 and /5 off the shield, seta /2 missing. Surface of shield reticulated. 

Tritosternum with base 60 p, laciniae 84 », with hyaline denticulate margin. Ster- 
nal shield (42 » X 130 ») weakly reticulated, with two pairs of setae and two pairs of 
pores ; st3 and metasternals free on unsclerotized integument. Between s/1, 63 p; 
between sf1 and st3, 84 w. Genital shield (180 » x I1IO p) weakly reticulated, with 
genital setae only. Anal shield (130 » X 105 w; maximum width 132 yp) with 
paranal setae 33 p, postanal seta 18 pw 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 III IV 
femur 2—3—2 2—2—I I—23—o o—3—I 
genu a—3, 2-2 1/242, #51 1-3, 2-1 ey, 
tibia 2—3, 7-2 I—i, #—1 I—}j, {—I I—j, {—I 


Coxa II without anterior spine. Length/width (in p») of leg segments : 


I I III IV 
genu 90/48 7545 60/42 80/40 
tibia 90/45 7245 66/40 90/38 


tarsus 165/40 126/40 126/40 162/36 


Male : Chelicera with segment I, 50 » ; II, 90 p; fixed digit reduced, spermadactyl 
long (110 »), grooved, completely fused with movable digit. Four pairs of gnathoso- 
mal setae with c.s. about 50 pw apart, Ayp.2 about 50 pw apart, Ayp.1 normal. Other 
gnathosomal features as in female. 

Dorsal shield (570 » x 282 ») bearing 18 or Ig 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 yp, laciniae 66 », 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 sfI, 52 »; between sti and st3, 108 uw. Paranal setae 27 p, postanal seta 15 pw 
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 II] and IV modified to form 
tooth-like structures. Coxa II without anterior spine. Length/width (in y) of leg 
segments : 


I II III IV 


genu 75/42 60/45 60/42 78/45 
tibia 75/40 51/42 60/42 78/42 
tarsus 114/34 75/30 -— —- 


. 
Protonymph ; Gnathosomal structures, excluding pedipalps, essentially as in — 
| 
| 


358 G. OWEN EVANS & W. M. TILL 1 
; 
| 
‘ 


female, but not so well developed. Four pairs of gnathosomal setae with c.s. about 
36 w apart, Ayp.2 about 32 w apart. Pedipalps with normal chaetotaxy (1I-4—5-12) 
for this instar. 

Podonotal shield (312 « X 120 «%; 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 p. Sternal shield (180 » Xx 102 p) with 
three pairs of setae and two pairs of pores. Between st1, 48 » ; between sti and sf3, 
108 «. Genital setae absent. Anal shield (84 » X 57 »; maximum width 75 yp) ; 
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 ) of leg segments : 


I II II 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 » apart, hyp.2 about 36 p apart. 

Dorsal shield (522 » x 198 ») with chaetotaxy as in female. 

Tritosternum with base 50 p, laciniae 70 p. Sternito-genital shield (270 « x 108 p) 
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 sf1, 60 py; 
between sft and st3, 110 w. Anal shield (105 » X 80 »; maximum width 96 yp) 
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 «) of leg segments : 


I Il lI IV 
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 


HapitaAt: Recorded in Britain from Hirundo rustica Linn. 


THE BRITISH DERMANYSSIDAE (ACART) 359 


Dermanyssus alaudae (Schrank) 


Acarus alaudae Schrank, 1781, Enum. Ins. Austriae : 515. 
Dermanyssus alaudae: Evans & Till, 1962, Ann. Mag. nat. Hist. (13) 5: 279. 
Acarus 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 wapart, hyp.2 about 36 wapart ; hyp.t 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. 
meen setae 71 and /5 lie off the shield; setae 73,74 and /2 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 slightly posterior to seta s4. Surface of shield 
without a well defined pattern. 

Tritosternum with base 30 yp, laciniae 66 »; with hyaline denticulate margin. 
Sternal shield (approximately 24 » X 120 pu) weakly sclerotized, usually with one 
pair of setae; st2 and s¢3 free on unsclerotized integument, metasternals absent. 


Fig. 100. Deyvmanyssus alaudae (Schrank), female; dorsum (A) and venter (B) of idiosoma. 


360 G. OWEN EVANS @& W. M. TILL 


Between str, 48 «; between sf1 and st3, 57 mw. Genital shield (162 » X 120 pn) 
with no definite pattern and bearing genital setae only. Analshield (108 » x I20p; 
maximum width 135 mw) with paranal setae 20 p, postanal seta 9 wp long. 
Chaetotaxy and sclerotization of opisthogaster as in Text-fig. loos. 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: 


I II III IV 
femur af? 4-3 {3-6 I—Sne 

~~ hd > me ee 
genu 2—t, {2 I—z, §—t OF oe Ti oe 
tibia 2—?, j-—2 I—}, j—I Py I—}; 2 


Coxa II without anterior spine. Length/width (in ,) of leg segments : 


I II Ill 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, Ayp.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/1, 
56 »« ; between s/r and st3, 102 w. Genital setae absent. Anal shield (63 » x 72 »; 
maximum width 80 »), 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 I] Ill IV 
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 


Hapirat: Recorded from Alauda arvensis Linn. in Scotland and Europe (Austria). 


THE BRITISH DERMANYSSIDAE (ACARI) 361 


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 pw apart, hyp.2 about 
36 w apart ; hyp.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 w X 336 mw) with II pairs of setae distributed as in Text-fig. 
ro1A; setae /5 off the shield, J/2 absent. Setae 72, 22 and 24 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 p») 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 st1, 72 w; between sf1 and st3, 90 w. Genital shield (174 » x 
60 pu) with weakly defined linear markings and bearing genital setae only. Anal 
shield (112 » X 138 w; maximum width 174 ») with paranal setae about 27 wu long. 
Opisthogastric cuticle bears a prominent arc of stout setae (Text-fig. 1r01B). Peri- 


Fic. tor. Deyvmanyssus 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—;—I I—}—1 I—}—I 
genu 2—#, i—I 2—i, j—I I—j, 5—I I—}, 5—I 
tibia 24,91 «=I-4, fr OI, C+}, Gt 


Coxa II without anterior spine. Length/width (in p) of leg segments : 


I II Ill 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 


Hasitat : 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 (3 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 Hypoaspis (Pneumolaelaps). 

Hypoaspis latifrons Hull, 1925 (9), referable to Hypoaspis (Gaeolaelaps). 

Hypoaspis nitidissimus Hull, 1918 (9), possibly a synonym of Haemogamasus pontiger 
(Berlese). 

Hypoaspis soricinus Hull, 1925 (3). 

Laelaps (Hypoaspis) longipes Halbert, 1915 (3), possibly a synonym of Haemo- 
gamasus horridus Michael. 

Laelaps \Hypoaspis) ovatulus Halbert, 1915 (9), referable to Pseudoparasitus. 

Laeliphis birksianus Hull, 1925 (9), referable to Hypoaspis (Gymnolaelaps) . 

Laeliphis fuscipes Hull, 1925 (gf 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 colombo nov., Hypoaspis giffordi nov., Hypoaspis hyatti nov., 
Hypoaspis neocunetfer 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 Ofhionyssus 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 nolla 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 Hyferlaelaps microta 
(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. 


304 G. OWEN EVANS & W. M. TILL 


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ABREVIATIONS USED IN TEXT AND FIGURES 


ad,_. antero-dorsal setae Jvi-5 internal ventral setae 

al,_. antero-lateral setae lat.l.f. lateral lyriform fissure 

QVy_5 antero-ventral setae m.a. movable digit 

C.S. capitular setae pi-3 sternal pores 

Bi) dorsal setae pd. pilus dentilis 

a.l.f. dorsal lyriform fissure pay_s postero-dorsal setae 

d.s. dorsal seta pl postero-lateral setae 

jd. fixed digit Pv 4-2 postero-ventral setae 

gen. genital setae pxI-3 posterior accessory setae 

hyp. 1-3 hypostomal setae 72-6 marginal series of podonotal setae 
JI-6 dorsal series of podonotal setae R1-7 marginal series of opisthonotal 


J I—5 dorsal series of opisthonotal setae setae 


366 G. OWEN EVANS & W. M. TILL 


s1-6 lateral series of podonotal setae 

S1-5 lateral series of opisthonotal setae 

Sp. spermadactyl 

sti—3 sternal setae 

sts metasternal setae 

UR submarginal series of opisthonotal 
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 
albatoaffiinis, Liponyssus, 295 

albatus, Dermanyssus, 278 
ALLONYSSUS, 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, 111, 113, 126, 150 
angulatus, Pseudoparasitus, 222 
aplodontiae Alphalaelaps, 116 

arcualis, Eulaelaps, 260 

arcuatus, Dermanyssus, 277, 278 

arcuatus, Hirstionyssus, 277, 278, 279 
arcuatus, Leiognathus, 278 

arcuatus, Liponyssus, 278, 281 

aridus, Ornithonyssus, 317, 337 

arvalis, Laelaps (Hyperlaelaps), 140, 141 
arvicola, Laelaps, 137 

arvicolae, Hirstionyssus, 295 

arvicolarum, Haemogamasus, 56, 57, 241, 250 
arvicolarum, Haemogamasus horridus, 250 
ASIATOLAELAPS, 115 

astronomica, Hypoaspis, 39, 159, 163, 208 
astronomicus, Iphis, 159, 208 
ATRICHOLAELAPS, 150 


Vi- ventral setae 

z1-6 median series of podonotal setae 

Z1-5 median series of opisthonotal 
setae 


ZvI-5 medio-lateral ventral setae 


auris, Raillietia, 117 

australis, Raillietia, 117 
AUSTROGAMASELLUS, 224 
avisugus, Haemogamasus, 244 
avium, Dermanyssus, 350 


bacoti, Leiognathus, 321 

bacoti, Ornithonyssus, 83, 84, 279, 302, 317, 
318, 321, 326, 337 

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, I91, 
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 


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, 349, 353 

claviger, Hypoaspis, 24, 159, 161, 182 

claviger, Laelaps, 182 

clethrionomydis, Laelaps, 128 

coleoptratorum, Gamasus, 279 

colomboi, Hypoaspis, 31, 32, 159, 161, 194 


INDEX 367 


compressus, Hypoaspis, 172 
confinis, Ololaelaps, 235 
COSMOLAELAPS, 158, 159, 160 
crassipes, Laelaps, 137 
cricetophilus, Haemolaelaps, 156 
cunetfer, Cosmolaelaps, 184 
cuneifer, Hypoaspis, 25, 159, 161, 184, 187, 
189 
cunetfer, Laelaps, 184 
cyclaspis, Liponyssus, 312 
BeCLOLAELAPS, 150 
cynomys, Hirstionyssus, 279, 280 


DWAVISTELLA, 159 

decumani, Myonyssus, 269, 271 

dentatus, Laelaps, 227 

dentatus, Pneumonyssus, I21 

dentatus, Pseudoparasitus, 48, 225, 227 
DERMANYSSUS,; 109, 118,347, 348 
DINOGAMASUS, 116 

dodevot, Laelaps (Hoplolaelaps), 266 
doderoi, Pseudolaelaps, 63, 122, 266, 362 
DOMROWNYSSUS, 121 


easti, Serpenticola, 337 

echidnina, Laelaps, 3, 4, 128, 134 
mCHINOLEAPS, 127 
ECHINONYSSUS, 276 

elliptica, Caris, 306 

ellipticus, Liponyssus, 306 

ellipticus, Macronyssus, 76, 77, 303, 304, 306 
ensiger, Laelaps (Cosmolaelaps) vacuus, 190 
ENTONYSSUS, 120 

eos, Hypoaspis (Haemolaelaps), 156 
equitans, Hypoaspis, 40, 159, 162, 210 
equitans, Laelaps, 210 
MUBRACHYLAELAPS, 150 
EUHAEMOGAMASUS, 239 
BPMLAELAPS, 114, 238, 239, 259 
eusorvicis, Hirstionyssus, 286 

evansi, Ondatralaelaps, 147 

evotomydis, Dermanyssus, 350 


fahrenholzi, Androlaelaps, 12, 151, 156 
fenilis, Gamasus, 152 

festinus, Laelaps, 132 

flaviot, Echinolaelaps, 134 

flavus, Dermanissus, 308 

flavus, Liponyssus, 308 

flavus, Macronyssus, 77, 78, 79, 303, 304, 308 
mONSECAONYSSUS, 316 

freemani, Hypoaspis, 152 

fucorum, Hypoaspis, 362 


fuscicolens, Hypoaspis, 32; 161, 196 
fuscipes Laeliphis, 362 


GArOLABRLAPS,112)i10%, 050, 160 

gallinae, Acarus, 348, 350 

gallinae, Dermanyssus, 95, 96, 97, 337, 349, 
35° 

giffordi, Hypoaspis, 17, 159, 163, 170 

gigas, Liponyssus, 271 

gigas, Myonyssus, 64, 65, 269, 271 

glasgow, Androlaelaps, 156 

glasgowt, Laelaps, 156 

GROSCHAFTELLA, 239 

gurabensis, Cosmolaelaps, 222 

GYMNOLAELAPS; £13,559; 160 


haemisphaericus, Ololaelaps, 231 

HAEMOGAMASUS, 111, 114, E19, 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, Laclaps, 15:2, 127,128 

hilaroides, Laelaps, 132 

HIRSTESIA, 303 

hivsti, Echinolaelaps, 134 

HERSTIONY SSWS; 1135 276; 277,279 

hirsutosimilis, Haemogamasus, 55, 238, 240, 
2AT; 242. 246 

hirsutus, Haemogamasus, 53, 114, 239, 241, 
242 

hirundinis Acarus, 356 

hirundinis, Dermanyssus, 97, 99, 349, 350, 
356 

hivundinis, Derymanyssus, 353 

HIST RICHONY SSUSsti9 

HOLOSTASPIS, 159,. 160 

hopkinsi, Raillietia, 117 

HOPLOLAELA PS, 224.266 

horridus, Haemogamasus, 57, 58, 240, 241, 
250) 252, 362 

humerata, Hypoaspis, 40, 159, 162, 212 

humeratus, Laelaps (Laelaspis), 212 

hyatti, Hypoaspis, 33, 159, 161, 198 

HYPERLAELAPS,. 113, 126, 140 


368 INDEX 


HYPOASPIS, 109, 111-4, 127, 158, 159, 160 
HYPOASPIS, 228 


ICHORONYSSUS, 118, 119 

inversus, Hypoaspis (Haemolaelaps), 172 
IPHIOPSIS, 109, 116 

IPHIS, tog 

isabellinus, Hirstionyssus, 73, 280, 281, 295 
isabellinus, Liponyssus, 295 
ISCHNOLAELAPS, 150 

ISCHY ROPODA, 238, 239 

isotricha, Holostaspis, 159, 203 
isotricha, Hypoaspis, 36, 159, 163, 203 
IXOBIOIDES, 115 
IXODORHYNCHUS, 115 


JACOBSONIA, 116 
johnstoni, Hirstionyssus, 279 
johnstoni, Macronyssus, 279 
JULOLAELAPS, 113, 116 


kochi, Laelaps, 141 

kolenatii, 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, 291 

latiscutatus, Liponyssus, 291 

LEIOGNATHUS, tog, 316 

LEPRONYSSUS, 303 

LIPONISSUS, 239 

LIPONYSSOIDES, 118, 347, 348 

liponyssoides, Haemogamasus, 114 

LIPONYSSUS, 239, 240 

lobatus, Liponyssus, 308 

longimana, Caris, 303 

longipes, Laelaps (Hypoaspis), 362 

lubrica, Hypoaspis, 18, 159, 161, 172 


mabuyae, Ophionyssus, 337 

MACROLAELAPS, 127 

MACRONYSSUS, 118, 121, 240, 278, 301, 
302, 303 

marginalis, Hypoaspis, 34, 159, 161, 199 

marginalis, Pneumolaelaps, 199 

mawsoni, Eulaelaps, 257 

megaventralis, Atricholaelaps, 152 

MELITTIPHIS, 275 

meridionalis, Laelaps, 224 

meridionalis, Pseudoparasitus, 225 

MESONYSSOIDES, 120 

michaeli Haemogamasus, 254 

microti, Hyperlaelaps, 6, 7, 141, 144, 145 

microti, Hypoaspis, 156 

microti, Laelaps, 137 

microti, Tetragonyssus, 140, 141 

MICROTILAELAPS, 127 

miles, Hypoaspis, 45, 159, 160, 222 

miles, Laelaps (Iphis), 159, 222 

minutissima, Hypoaspis, 35, 159, 161, 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, 279 

musculi, Dermanyssus, 277, 278, 279 

muscult, Hirstionyssus, 277, 279, 280, 291 

musculi, 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 ’ 


a SS 


nagayoi, Liponyssus, 321 
natricis, Dermanyssus, 335, 337 
natricis, Ophionyssus, 90, 91, 92, 302, 337, 
340 
NEOBERLESIA, tog 


INDEX 369 


neocuneifer, Hypoaspis, 26, 159, 161, 187 
NEOLAELAPS, 118 
MEOLTPONYSSUS, 335 

nidi, Haemogamasus, 59, 240, 241, 254 
nidicorva, Hypoaspis, 22, 23, 159, 162, 179 
nitidissimus, Hypoaspis, 362 

noctula, Dermanyssus, 278 

nol, 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, 
33D 

oculatus, Hypoaspis, 152 

BeOrAr LAPS, 113, 125, 126, 224, 228 

ondatrae, Ornithonyssus, 317 

SNDATRALAELAPS, 113, 126,145 

oophila, Hypoaspis, 38, 159, 163, 206 

oophila, Laelaps, 206 

orcadensis, Hirstionyssus, 280 

orcadensis, Ichoronyssus, 291 

BEEIONYSSUS, 100, 124, 300, 301, 302, 
43997 330; 337 

oribatoides, Laelaps, 260 

SOE NETHONYSSUS, 124, 278, 279, 301, 302, 
316, 317 

OY CLOLAELAPS, 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 
pavvanalis, Laelaps, 147 
Pet KRINYSSUS, 276 
pavlovskii, Ornithonyssus, 329 
paviovskyi, Haemogamasus, 244 
pedalis, Laelaps, 260 
pereira, Ornithonyssus, 317 
periblepharus, Steatonyssus, 87, 88, 278, 302, 
329, 330 
pilifer, Hypoaspis laevis, 175 
Pipistrellae, Dermanyssus, 278 
Pipistrelli, Liponyssus, 312, 326 


pipistrelli, Ornithonyssus, 85, 86, 303, 318, 
2265. 320 

placentula, Laelaps, 235 

placentula, Ololaelaps, 52, 229, 235 

PNEUMOWAEB TAPS, 113) 159, 160 

PNEUMONYSSUS,-121 

PODOCINUM, tI09 

pontiger, Groschaftella, 257 

pontiger, Haemogamasus, 60, 114, 240, 241, 
257; 362 

pontiger, Laelaps (Eulaelaps), 257 

praesternalis, Hypoaspis, 19, 159, 162, 173 

PRISTOLAELAPS, 226 

propheticus, Laelaps, 260 

PSEUDOLAELAPS; 266 

PSEUDORPABASIIUS, 113, 127, 224, 225 


quintus, Dermanyssus, 101, 349, 361 


RAL UE ATICAP S «127 
rhabdomys, Ischnolaelaps, 156 
RHINOLAELAPS, 259 
vhinolophi, Liponyssus, 315 


sanguineus, Haemogamasus, 321 

sardoa, Hypoaspis, 20, 159, 162, 175 

savdous, Laelaps (Androlaelaps), 175 

sauravum, Liponyssus, 342 

saurarum, Ophionyssus, 92, 93, 302, 337, 
342 

SAURON VWSSIUS, 335, 330; 337 

scalopi, Haemolaelaps, 156 

SEIUS, 109 

sellnicki, Ololaclaps, 50; 51, 123, 220,, 231, 
235, 237 

SERPENTICOLA, 335 

serpentium, Ichoronyssus, 337 

setiger, Liponyssus, 156 

setosus, Liponyssus, 240 

sigmodont, Atricholaelaps, 156 

simicola, Pneumonyssus, 121 

smith, Hypoaspis, 172 

soarianus, Hypoaspis, 152 

soricinus, Hypoaspis, 362 

soricis, Hirstionyssus, 70, 71, 280, 281, 286 

sorvicis, Ichoronyssus, 286 

spiniger, Liponyssus, 147 

spiniger, Tetvagonyssus, 147 

stabularis, Eulaelaps, 61, 260, 362 

stabularis, Gamasus, 259, 260 

SLEATONYSSUS, 278; 279, 301, 302, 320 

stegemani, Laelaps, 156 

sternalis, Haemogamasus, 244 


370 INDEX 
sternalis, Liponyssus, 315 truncatus, Acurus, 359 
STRANDTIBBETTSIA, 115 tumidulus, Laelaps, 231 
strandimanni, Alricholaelaps, 156 TURKIELLA, 150 
STRATIOLAELAPS, 159, 160 
STYLOCHIRUS, 109 uncinatus, Leiognathus, 315 
sylviarum, Dermanyssus, 316, 318 uncinatus, Macronyssus, 81, 303, 304, 315 
sylviarum, Ornithonyssus, 82, 123, 302, 317, vacua, Hypoaspis, 27, 28, 159, 161, 190 

318 vacua, Laelaps, 190 

vacuus, Laelaps (Cosmolaelaps), 190 

talpae, Hirstionyssus, 67, 68, 69, 277, 278, veneta, Ololaelaps, 49, 229, 231 

280, 281 venetus, Hypoaspis, 228, 231 
tenuiscutatus, Liponyssus, 321 virginianus, Laelaps, 156 
TETRAGONYSSUS, 269 vitzthumi, Echinolaelaps echidninus, 134 
TRICHOSAU ROLAELAPS, 276 
triscutatus, Dermanyssus, 337 ZYGOLAELAPS, 150 


OW. Hat Z 
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_ A REVIEW OF THE GENUS PHILETOR 
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ZOOLO GY | Vol. 14 No. 6 


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A REVIEW OF THE GENUS PHILETOR 
(CHIROP TERA: .VESPERTILIONIDAE) 


BY 
J. E. HILL 


Department of Zoology, British Museum (Natural History) 


Pp. 371-387 ; 3 Text-figures 


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BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY | Vol. 14 No. 6 
LONDON : 1966 


THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, 1s 
issued in five series corresponding to the Departments 
of the Musewm, and an Historical series. 


Parts will appear at irregular intervals as they become 
ready. 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. 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 


A REVIEW OF THE GENUS PHILETOR 
(CHIROPTERA : VESPERTILIONID AE) 


By J. E. HILL 


SYNOPSIS 


The genus Philetor 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 P/uletor with type species P. rohui 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 Piletor as thus understood have been studied with a view to 
establishing the relationships of this small but interesting genus. 


PHILETOR Thomas 
Philetor Thomas, 1902: 220. Type species Philetor vohu1 Thomas. 


A study of the genera Prfistrellus, 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 P/iletor 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 (1°), 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 


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374 ee bg me 


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 isnarrow, _ 
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. ra—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. 1a) 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 tuft of 
moderate, rather stiff hairs. According to Thomas (1902 : 221) the penis of Philetor _ 
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 


= = —————————E—EEO-llS 


Mace VIEW ORCS GENUS PHIEETOR 375 


longitudinal groove extending almost to the point of emergence from the prepuce 
(Text-fig. 1b). The os penis or baculum (Text-fig. 1c—e) is strongly curved dorso- 
ventrally (Text-fig. 1d) 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 


sutinieaiics 


Wirth h et: 


ae] snebeeiven 


2 mm. e 


Fic. 1. Philetor rohui vohui. a, dorsal aspect of penis ; b, ventral aspect of penis ; c, dorsal 
aspect of os penis; d, lateral aspect of os penis ; e, ventral aspect of os penis. 


37° j..@. BIL 


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—) 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. rohwi 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 


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PALE EDP OR 


A REVIEW OF THE GENUS 


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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 (i*) is small and conical, its height only slightly exceeding the cingulum 
height of i*, 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 (m'). 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 (is) is very slightly wider than i, or iz. 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 
Philetor rohui Thomas, 1902: 220. Albert Edward Mountains, Papua, New Guinea, 6,000 ft. 


Narial emargination not abruptly widened just above the roots of i?-*._ 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) 10-3—10-7 (10-5); width of braincase 7-3-8-2 
(78) ; c-m® (front of canine to crown of m4) 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 


| 
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| 
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A weEVIEW OF THE GENUS PAIEETOR 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 i2-?. 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 fachyotis 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 Epiesicus’’ 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. rohm 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. rohm, Cranially, there is a high degree of agreement but the braincase is a little 


380 J. E. HILL 


more inflated than in P. rohut 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 rohui 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 E. 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,_, 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 Philetor 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 (1907 : 207)), Tylonycteris and Hesperoptenus, also — 
drawing attention to its resemblances to Pterygistes (= Nyctalus). Miller (1907 : 214) _ 


SaRE VIEW OF GH "GENUS PHITLETOR 381 


remarked that Philetor appeared related to Tylonycterts 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 joffrei 
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 P/iletor 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 Piiletor, 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 Philetor, 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 Piiletor and Eptesicus but in 
that genus the wing is not reduced as it is in Philetor. The frontal part of the brain- 
case is 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 Philetor and i° 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 Tylomycteris. 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 Philetor 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 $387 BILL 


ment of supraorbital tubercles, minimal in 7. fachypus and its allies, maximal in 
T. robustula 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 Philetor and has a large 
central cusp equal in height or nearly equal in height to the posterior cusp of i, 
flanked by two smaller lateral cusps. It is slightly hollowed posteriorly. In 
Philetor 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,_,) are slightly imbricated and pm, is but slightly reduced and is almost 
equal in height to pmy, 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 Mimetillus the reduction _ 
of the wing is not confined only to shortening of the fifth digit as in Philetor but _ 
extends also to shortening of the third digit, a feature unremarked in any other bat. 
The penis of Mimetillus 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 Philetor. 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 of 
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 Tylonycteris 
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, 


_ _ ————————_——_——_ESrls” 
————— nines 


aie VIEW OF HE 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 (1902: 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 joffrer, stenopterus and brachypterus subsequently 
removed by Tate (1942 : 252) to form the joffrer group of Pipisivellus. There is no 
immediate connection between P/iuletor 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 pmy. 

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)j 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 joffrez: in 
Nyctalus but (p. 173) leave anthony1 in Pipistrellus. Of these four species, only joffrei 
and stenopterus are available to me. The members of the joffrez 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 m3. The outer upper incisor (1°) 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 joffre: 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 savii group of Pipistrellus in which pm? is minute and the palate shortened 
but which have not developed supraorbital tubercles. 


354 J. BE. HILL 


So far as I am able to determine from the material available to me and from the 
literature Pipistrellus joffret and P. anthonyt 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. joffret 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 Philetor 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 Pipistrellus joffret 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 Philetor and has a similarly ligulate 
post-palatal spine. The inner upper incisor (i?) is elongate as in Philetor 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 (1%) 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-half 


PREVIEW OF THE GENUS PHILETOR 


mm. 


la. 


c, female external genital 


° 
’ 


1S 


; 6, ventral aspect of pen 


a, dorsal aspect of penis 


Di 


trellus joffve 


4s 


ye) 


FIG. 3. 


385 


386 1k at. WEL. 


of its width. There seems little doubt from the description by Tate (1942 : 252) 
that Prptstrellus anthonyt is very closely related to P. joffret, 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 joffret group seem much less closely related 
to Philetor than are Pipistrellus joffrei or P. anthonyi. The ears and tragus of 
P. stenopterus are similar to those of P. joffret 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 like that of P. joffrec but the supra- 
orbital tubercles are less developed. The anterior upper premolar (pm?) is relatively 
larger than in P. joffret 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 pm,, 
which is more reduced than in P. joffre:. 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 joffret group of Pipistrellus and also between Philetor 


: 
; 


and Tylonycteris, although Philetor seems sufficiently removed from both Pipistrellus _ 


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 Mimetil- 4 
lus from a point on the Pipistrellus stem. I would endorse the view expressed by — 


this author elsewhere in the same paper (p. 266) that Philetor 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 of © 


woe wie WwW OF Tie GENUS PHIELETOR 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 Tylonycterts, 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 joffre: 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 Philetor 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. rohut are now recognized, the nominate subspecies being so far known only from 
New Guinea. LEptesicus 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 joffre: group of Pipistrellus and with 
the genus Tylonycterts. 


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, 

a2 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. 

CHASEN, F. N. 1940. A Handlist of Malaysian mammals. Bull. Raffles Mus. No. 15: i-xx, 
I—209, map. 

Dosson, 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-Scott, T. C. S. 1951. Checklist of Palaearctic and Indian 
mammals, 1758-1946. London. 

GiBSON-HiLL, C. A. 1949. Bird and mammal type specimens formerly in the Raffles Museum 
collections. Bull. Raffies Mus. No. 19: 133-198. 

Lauriz£, 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. 

MiLtLER, 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: 221-297, 5 figs. 

Tuomas, O. 1902. On a new genus of Vespertilionine bat from New Guinea. Ann. Mag. 
wat. Fist. (7), 9 : 220-222. 


ZOOL 14, 6. 26 


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. PONTOBDELLINAE > 
__ (PISCICOLIDAE : HIRUDINEA) 


IN THE BRITISH MUSEUM (NATURAL HISTORY) 
WITH A REVIEW OF THE SUBFAMILY 


b Gadocew BLOYN 


a _ BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
me oey ape mes i Vol rm Noe 7 


LONDON > 1966 


PONTOBDELLINAE (PISCICOLIDAE : HIRUDINEA) 


IN THE BRITISH MUSEUM (NATURAL HISTORY) 
‘WITH A REVIEW OF THE SUBFAMILY 


BY 


L. C. LLEWELLYN 


Department of Zoology, University College of Swansea 
Present address: Inland Fisheries Research Station, Narrandera, New South Wales 


Ph. 389-439 ; 28 Text-figures 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY ~ Vol. 14 No. 7 
LONDON : 1966 


9 THE BULLETIN OF THE BRITISH MUSEUM if) 1 

| (NATURAL HISTORY), instituted in 1949, is > aan 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. 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. 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 g September 1966 


PONTOBDELLINAE (PISCICOLIDAE : HIRUDINEA) 
IN THE BRITISH MUSEUM (NATURAL HISTORY), 
WITH A REVIEW OF wail SUBFAMILY 


# mah ’ 
By L. Cf LLEWELLYN 
CONTENTS 

\ og, ee I Page 

SYNOPSIS . ‘ : : : —_ ow” : : : : ~ aso 

INTRODUCTION . ; , : , : : ; ‘ . : 392 

MATERIAL . i 393 

ANATOMICAL COMPARISON OF Sa eeacie ‘daerothela AND ' Pontobdella 

muricata : : : : : : - : . . : 393 

Gurt.y. - : ‘ ‘ - ‘ : : : _ ee, 

Reproductive oe : , ‘ : ; : ot bys Beng 7 

Nervous system and sina cel . : : ; : : -. = 401 

PONTOBDELLINAE SUBFAM. NOV.—DIAGNOSIS 6 : : : 2 4404 

Genus Stibarobdella—Diagnosis_ . * ; : : ; sh) 1405 

Stibarobdella macrothela : , ‘ ; . , i) 405 

Stibarobdella loricata . , : : : : , sue ALD 

Stibarobdella bimaculata : ' : ; , ‘ aw. bs 

Stibarobdella planodiscus . ; : : : : et EG 

*(Stibarobdella tasmanica) . : : : : : pee", 

(Stibavobdella taprobanensis) d : : : : Amer 42 x 

(Stibarobdella mooret) . é i : ; i ; - eee 

(Stzbarobdella australiensis) . : : , : é ae 225. 

Genus Pontobdella—Diagnosis : : : : : : ma gs 423 

Pontobdella muricata . : : : : ‘ : Pm 423 

Pontobdella vosmaent . ; i ‘ ’ : : MOL, 427 
(Pontobdella aculeata) . : ; : ; , ‘ iameage * 

(Pontobdella vagosa) . : ‘ i : é : softies 

Genus Pentabdella gen. nov.—Diagnosis. : . : : aa 

(Pentabdella dispar) . : ! 3 ‘ : ‘ Pe 

Genus et species inquirendae . . : : : Pd 

KEY TO THE SPECIES IN THE SUBFAM. PONTOBDELLINAE 5 ; =o) OEE S9 

ACKNOWLEDGEMENTS . : é d : d : é 4 “a 4a8 

REFERENCES ; . é ‘ z ‘ ‘ 3 3 : : 438 


* Species not examined during this study, but well described in litteris, appear above in brac kets. 


SYNOPSIS 


The conspicuously tuberculate marine leeches may conveniently be separated in a new sub- 
family Pontobdellinae containing the genera Stzbarobdella, Pontobdella, and Pentabdella gen. nov., 
for species in which the mid-body somites bear threef, four and five annuli respectively. 

The internal anatomy of Stzbarobdella macrothela 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 


t or no more than three (p. 439). 
ZOOL. 14,7. 27 


392 L. CC. 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 Stibarobdella examined, S. macrothela is particularly characterized by a very large 
posterior sucker, amorphous tubercles with large, more or less square basal areas and a pair of 
ocular patches; S. loricata 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. vosmaert 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 
inguirendae. A key is constructed for the identification of the species reviewed. 


INTRODUCTION 


Tue 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 vosmaert 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 Parapontobdella, for 


Pontobdella tatejamensis, which was described rather briefly by Oka (rgro). 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 tatejamensis in Pontobdella (see p. 436), Para- — 


pontobdella may have to be separated on other characters. 


REVIEW OF THE 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 Prscicola 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: 1. An introduction to the internal anatomy 
of these forms by comparing dissections of the type species of the genera Siibarobdella 
and Pontobdella. 2. A systematic account of the genera and species placed in the 
sub-family, accompanied by details of several species inguirendae. 


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 St:barobdella macro- 
thela is hardly known at all. The general dissections depicted in Text-figs. I 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 lie 
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, 


304 L. Cc. LLEWELLYN 


f. 


pam, Pp . 


ut 


HR, 


P.C.R. 


Cc.R. 


B.G.R, 


AR, 


ric. 1. General dissection of Pontobdella muricaia to show relationship of organs to somites 
of the body. I-XXVII, numbering of somites; an., anus; A.R., anal region; 4@.s., 
anterior sucker ; b.g., caecum (blind-gut) ; B.G.R., caecal region (blind-gut region) ; C.R., 
clitellar region ; d./., dorsal blood vessel; e.d., ejaculatory duct; /., fringe; g., salivary 
gland cells; H.R., head region; m.i., mid-intestine; 0., ovary; p., papilla; P.c.R., pre- 
chitellar region ; ~.i., posterior intestine ; p.m., protractor muscle; pr., proboscis; p.s., 
posterior sucker; sep., septa; t,-t,, testes; 7 R., testicular region. 


REVIEW OF THE PONTOBDELLINAE 395 


B.G.R. 


S35 
XXV 
a< XXVI-KX VIL, ae aye aS 


Fic. 2. General dissection of Stibarobdella macrothela to show relationship of organs to 
somites of the body. ¢., pigmented eye patches ; /.p., lateral projections of mid-intestine ; 
p.c.d., diverticula. See Fig. 1 (p. 394) for other abbreviations. 


396 L. C. LLEWELLYN 


which stretches from the posterior edge of the oesophagus to the XIIth or last 
clitellar somite, is fairly thick-walled in Stibarobdella, 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. 


€.0.g. 


p.c. 4. 


fe. 


an. 


bic. 3. Gut of Stibarobdella macrothela: c.0.g., circum-oesophageal ganglion; /fe., 
fenestra ; m., mouth; oe., oesophagus; r.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 (1038) 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,—a,, a,—b, and 
within the small annulus be, but these cannot easily be distinguished in Stzbarobdella 
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 XXVII, 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 (=Sitibarobdella 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 1s 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 Stibarobdella 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 
Stibarobdella macrothela. 

The paired ovaries are tubular but comparatively short, leading into short oviducts 


LLEWELLYN 


C. 


L. 


398 


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REVIEW OF THE PONTOBDELLINAE 399: 


H.R. 


P.C.R. 


C.R. 


T.R, 


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; gm., ganglion ; 
H.R., head region; 2.”., segmental nerve; p., papilla; p.c.R., preclitellar region; p.s., 
posterior sucker ; sep., septa; ¢,-¢,, testes; T.R., testicular region. 


400 L. C. LLEWELLYN 


H.R. 


P.C.R. 


TR. 


B.G.R. 


AR. 


lic. 7. Dissection to show the arrangement of ganglia in the neurosomites of Stibarobdella 
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 Sézbarob- 
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 3 
-Clitellar region 3 
Testicular region 6 
Caecal (blind-gut) region 6 
Anal region 3 


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 1:2:1. 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 Stbarobdella macrothela these are trimeric, with their 
widths varying in the ratio I : 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 X XVII, containing three 
ganglia which are much closer together. In Stbarobdella 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 


DISTANCE BETWEEN GANGLIA 


402 L. C. LLEWELLYN 


murtcata 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 


24 


20 


---O--- Pontobdella muricata. 


~--@--- Stibarobdella macrothela. 


25 50 75 100 125 15sO 175 200 225 250 275 300 325 ‘ 


DISTANCE FROM ANTERIOR END TO MIDOLE OF COMMISSURE 


Fic. 8. Graph to show relative distances between ganglia in Pontobdella muricata and 
Stibarobdella macrothela and the elongation of the caecal (blind-gut) region in the former. 
The scales for the axis and abscissa are in arbitrary units which differ slightly between 
species. The measurements were taken from one specimen of each species. 


403 


REVSEW OF LE PON TOBDELELNAE 


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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 b,. 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 bg. 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 bs. 

Stibarobdella macrothela differs from this condition in several ways (Text-fig. 10). 
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 aj. As in 
Pontobdella muricata there are five main branches at the lateral glia cells, of which I, 
Il and III run anteriorly. Nerve IV appeared to terminate laterally in annulus ay. 
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, and a,;. 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 ag. 

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 b,. 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 b, 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. 


DiaGnosis. 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. 
DIAGNOSIS. 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. 

Tvachelobdella 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 
TE sATIIFICR WCC Same 4 Ged WAM (Gelapahuaaieinesonie bea eiasibt 
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 Stibavobdella biannulata (Moore), see p. 439 


406 


L. C. LLEWELLYN 


Ant. 


x 
x! 
all 
WHY ea 
A all 
a 
m.¥. vi ! d.i m.d. 4.i I oi oan 


v.p. v.m. dm. d.p. D d.p. d,m. v.m. v.p. 


Fic. 11. Stibarobdella macrothela. a. Section through anterior sucker, showing oral 


surface. w. Ventral surface of clitellar constriction, showing positions of genital pores 
and tubercles. c.(a,bandc). Glandular or furrowed tubercles. pb. Tuberculation of a 
typical triannulate somite split mid-ventrally. For designations of annuli (i.e. ay, a, and 
a,) see Mann, 1953. X—XJJJ, numbering of somites; Ant., anterior end ; d.i., dorsal 
intermediate line; d.m., dorso-para-marginal line; d.p., dorso-para-median line; f-p., 
female pore ; g.t., furrowed tubercle ; /., lateral line ; m., mouth; m.d., mid-dorsal line ; 
m.p., male pore; m.v., mid-ventral line; #., papilla; s.e., scalloped edge; ss., sensilla ; 
»i., ventral intermediate line ; v.m., ventro-para-marginal line ; v.p., ventro-para-median 
line. 


REVIEW OF TITE,PONTOBDELRLIN AE 407 


H.R. 


: : 
PCR 0.000 Vill 


ENS eee: i ee a ii 


ola OneIm el Ome ae 
T.R. DmIOnONOUOENG 


etd dS 


So 


B.G.R. 


OOO) xx 
a 
ie) }(E) Boy 


qn. 


Fic. 12. Dorsal view of Stibarobdella macrothela showing annulation, tuberculation and 
suckers. J-XXVITI, numbering of somites; av., 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; ., papilla ; P.c.R.)’ pre= 
clitellar region ; p.s., posterior sucker; s., small white tubercle; T.R., testicular region. 


408 L. C. LLEWELLYN 


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 XIIIa,. 
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 XIIIa, 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 XIIla,, 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 c). They are generally 
more or less square, occasionally with lateral indentations making them somewhat 
“ H” 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 


REVIEW OF THE’ PONTOBDELLINAE 409 


clitellar constriction are very small and rounded. Annulus XIIJa, 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. AHirudo 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 (19209), 
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 


Somites. 


Dorsal. refers taleelteleelstobote le olels [oe] <[o eehal «oh 
Ventral, Ce eee eee 


Size |-3. 


Somites. 


Dorsal. [6 [42612] 6 ojo alaa|eta| e | @ | 4 | 
Ventral. i a 


Size 1-3. 


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 ¢ and 9 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; amn., 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 L. C. 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. moorei 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 Stbarobdella 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 benhami, 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 4Il 


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. 


Fic. 14. Stibarobdella lovicata. a. Section through anterior sucker, showing oral surface. 
B. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. 
Cc. (a) Tubercle with sensillae. (6) Tubercle on annulus showing striae. Db. Tuberculation 
of a typical triannulate somite split mid-ventrally. f., fringe; st., stria; ¢w., tubercle. 
See Fig. 11 (p. 406) for other abbreviations. 


Fic. 


15. 


L.C. LLEWELLYN 


H.R, 


P.C.R, 


C.R, 


TR, 


B.G.R. 


Dorsal view of Stibarobdella loricata 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 toit. 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 XII 
Triannulate somites VII-XI; XIII-XVXI ; 


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 I: 2:1. 

All eight annuli of the clitellar constriction, i.e. Xa, to XIIla, 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, anda 3. 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 ag. 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 XX VII 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 L. C. LLEWELLYN 


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. Xa,, XIa, and XIIIa, were 
often free of tubercles, but occasionally Xla, 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 


Somites. | 
Dorsal. pif <8 ie 6 [0] 4 [e@lec| 4 [oo 
Ventral. WOON Ao 
Size '-3. 2/2 

Somites. xxIII XKIV XXV ,XXVI pXXVII 

Dorsal. 6] 4 bal o[s[ols}elo]s] 4 nd 

Ventral. BNO RD RIF Ld RSE ~ Oo} 

Size 1-3. iid} 

an. 
bic. 16. Diagram illustrating the tuberculation of a single specimen of Stibarobdella loricata, 


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. ‘4°’ in the size column indicates the presence of an annulus dorsally only. 
See Fig. 13 (p. 409) for other abbreviations, 


REVIEW OF THE PONTOBDELLINAE 415 


tubercles (Text-fig. 14ca and 0) 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 Stubarobdella 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 (I910 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, XITI-X XIV bx, a 


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 Stcbarobdella. The other two preclitellar 
somites VIII and IX are triannulate with their annular widths 1: 2: TI. 

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 XIIla, 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. C. 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 ag annuli generally have six dorsal and four 
slightly smaller ventral ones. The tubercles are conical, generally with between six 
and ten apical sensillae. 


my 1 m 1 my. 


. 
° 
° 
° 
° 
° 
° 
° 
° 
° 
° 
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fe) 
Oo 
oO 
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O 
fe) 
° 
en be © oe 
2 


° 
° 
° 
° 
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° 
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mI 


Cc 


Fic. 17. Stibarobdella bimaculata. a. Section through anterior sucker, showing oral surface. 
». Ventral surface of clitellar constriction, showing position of genital pores and tubercles. 
c. Tuberculation of a typical triannulate somite split mid-ventrally. gv., outline of gang- 
lion on ventral surface: tu., tubercle; See Fig. 11 (p. 406) for other abbreviations. 


REVIEW Of THE/PONTOBDELLINAE 417 


H.R. 


P.C.R. Sas 


C.R. pea ee XL 


B.G.R. Pp 


A.R. —- 
6 Sho ae XXVI 


Dorsal view of Stibarobdella bimaculata showing annulation, tuberculation 


See Fig. 12 (p. 407) for abbreviations. 


Fic. 18. 
and suckers. 


418 L. C. LLEWELLYN 


REMARKS. Both Sétbarobdella 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 a, 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. || 3] 6 is] 4] elo] ololale| 6 
jolo| 2ia/4|a[4ja]s|a]4]4[a | 
1} 4 3 


Ventral, 


Size 1-3. 


Somites, 
Dorsal. 
Ventral, 


Size |-3. 


' 
' 
J 
' 
' 
' 
’ 
' 
i 


ric. 19, Diagram illustrating the tuberculation of a single specimen of Stibarobdella 
bimaculata, anterior extremity above, posterior below. ? indicates where the limits 
of the somites are uncertain. ‘‘ 4” in the size column indicates the presence of an 
annulus dorsally only. See Fig. 13 (p. 409) for other abbreviations. 


— ae 


REVIEW OF THE, PONTOBDELLINAE 419 


as broad again (Text-fig. 20B). The anterior : posterior sucker ratio when contracted 
en 511: 6. 


_ANNULATION. Uniannulate somites XXVI, XXVIT. 
Biannulate somites XI, XII, XXV. 
Triannulate somites VII-X, XITI-XXIV. 


The three preclitellar somites VIJ-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 XXV 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 1 mV. 


boo 1 Go. OO: OOo pi 74 


eee Hy lt © Ono ha Re eek: OQ wwibxs 


De Onn 4G) LOO 0 Oo: Ga 


D 


Fic. 20. Stibarobdella planodiscus. a. Anterior sucker. 8B. Posterior sucker. c. Dorsal 
surface of clitellar region. ob. 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 ; fu., tubercle. See Fig. 11 (p. 406) for other abbreviations. 


420 L. Cc. LLEWELLYN 


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 XIIla, annulus. Tubercles are also small 
and irregular on the last three annuli of the anal region and are often indistinguishable 
on the ventral side of the annulus of somite XXVII. 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 and that 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 Stibarobdella 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 Hirudo 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 XXVII : (Somites numbered 
Biannulate somites X—XII AAV, XAVI. as in S. macrothela, 


Triannulate somites VII-IX, XIII-XXIV. 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 az, 
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, 1963b : 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 1z: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, XITI-XXIV. 
(Somites numbered as in S. macrothela, 
Text-figs 72.) 


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 L. C,. LLEWELLYN 


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 XX VI and 
XXVII. 

DISTRIBUTION. Wadge Bank, Ceylon. 

Host. Unknown. 


Stibarobdella moorei (Oka, Igo) 


Pontobdella mooret Oka, 1910: 171. 
Stibarobdella 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, XIII-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—10 apical sensillae. A, and a, 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 XIIIa,. 

DisTRIBUTION. Hondo (Coast of Sagami, Awa). Coast of Tokyo. 


Host. Shark. 


Stibarobdella australiensis (Goddard, 1909) 
Pontobdella australiensis Goddard, 1909 : 724. 
DESCRIPTION. General characters. (Specimens not examined.) Body up to 


ir 


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. 


REVIEW OF THE PONTOBDELLINAE 423 


ANNULATION. Uniannulate somites RO MRT 


Biannulate somites XI, XXV, XXVI . : 
(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 a, 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. 
DiaGnosis. 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: 11. 

Pontobdella areolata Leach, 1815: Io. 

Pontobdella spinulosa Leach, 1815: 12. 

Pontobdella muricata: de Blainville, 1818: 293; Harding, I910: 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. 21A 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. 14, 7. 29§ 


424 L.C. LLEWELLYN 


ANNULATION. Uniannulate somites — 
Biannulate somites XI, XII, XXV, XXVI, XXVII. 
Triannulate somites VII-X. Text-fig. 22. 
Quadriannulate somites XIII-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 ratio1: 2:1. The clitellar constriction 
includes the small annuli Xa, and XIIIa,, between which are from four to six annuli, 


bic. 21. Pontobdella muricata. a. Section through anterior sucker, showing oral surface. 
®. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. 
c. a, band c: Tubercles in various stages of contraction. p. Tuberculation of a typical 
quadriannulate somite split mid-ventrally. For designation of annuli, i.e. a, ag, bs and 
b,, see Mann 1953; /,, fringe ; tu., tubercles ; See Fig. 11 (p. 406) for other abbreviations. 


REVIEW OF THE PONTOBDELLINAE 425 


H.R. { 
a.s. 


P.C.R. Gea eee 


TR. 90079 xvi 


B.G.R. 6... 0.05.0 
Sar es mes 
OOO; © 
G6 6.46 
aeons a 
©) C) xxi 
Goo Oo 
° ° oe = ae ee = 
YO OL FO | KKUY 
ia one. oy 
Se on grees 9s 
() 
ie Se Sane 
= | —— Sees 0 


ar. p.s, 


Fic. 22. Dorsal view of Pontoddella muricata showing annulation, tuberculation 
and suckers. /f., fringe; See Fig. 12 (p. 407) for other abbreviations. 


426 L.C. LLEWELLYN 


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 ag, or a, and a, 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:(30r 4):2 

The annulation of the three anal somites XXV—XXVII 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 XIIla, often have 


H.R. P.C.R. C.R. Teh 


Somites. 


Dorsal rae letelelalotsleleleletelelittatets lal s1¢ [= elanaeraa 
Ventral. opetore tate tats tote tas bate T ete tet2) olen s bey stat stole 


Size !-3. 


Somites. xXxIll XXIV XXV XXVI ,XXVII "| 
Dorsal. DAON ON Gc: 
Ventral. oO} 

-< 
Size 1-3. +3 


hic. 23. Diagram illustrating the tuberculation of a single specimen of Pontobdella 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. ‘‘ 4”’ 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 annulli. 


REMARKS. Harding (1910) observed that the shape of the tubercles can vary 
considerably depending on whether the leech is relaxed, gorged or otherwise (Fig. 
21Cca, 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. /aevis as having entirely retracted 
tubercles. It is possible, however, that P. laevis 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 smallin P. muricata. There are no grounds for 
the former species, since in fact P. muricata somites usually have one large annulus 
a,, 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 brumpti 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 
mer-4. Ihe. 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 ROO LOX VEE. 
Biannulate somites Pele A: XO VI 
Triannulate somites VIII, X 
Quadriannulate somites Soa, 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 L. C. 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 ag 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 XIIIa,, which is small and sometimes 
difficult to see, is the last of the clitellar constriction. Somite XIII is quadriannulate, 
the ratio of widths being I:3:2:1. Somites XIV to XXIV are also tetra-annulate, 


3 
= 
3 
a 
3 
< 


3 
= 
| 
. 
° 
° 
° oO 
° 
° 
° o 
° 
° 
°° 
° 
°o . 
12) 
° 
° 
° ‘ 
° 
oO 
ee ee) ee ee 
eo a 
“ N 


hic. 24. Pontobdella vosmaeri. a. Section through anterior sucker, showing oral surface. 
&. Ventral surface of clitellar constriction, showing position of genital pores and tubercles. 
©. Tuberculation of typical quadriannulate somite split mid-ventrally. _m.v.t., mid-ventral 
tubercle; éu., tubercle; See Fig. 11 (p. 406) for other abbreviations. 


REVIEW OF THE PONTOBDELLINAE 429 


H.R. aT vow 


P.C.R. 
p coco dy 


C.R. errr 
C.N. ee 


TR. Sass 


CIO BO Bea 


A.R. 5 


Fic. 25.: Dorsal view of Pontobdella vosmaeri showing annulation, tuberculation 
and suckers. See Fig. 12 (p. 407) for abbreviations. 


430 L. C. LLEWELLYN 


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 XX VI 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 Riviére (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 
XI Ila; was according to Riviére 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 


H.R. P.C.R. CR. a; 

‘ea re Se Sa ae i ae 
Somites. I-VI vil Vill Ix x XI xi XH XIV 

"ache pave & ce 
corsa. {7isial[s[e[e: [4 bolo] «ow si) 4 [6] 6]7 | 6 omlol4|6]4 | 6] 4 
vera £1 f Tofolst hal [«[s [+ sfol«fa[«tsfolstets[s 
Sizet-3% + fof Puprpappap2yr]3 3 2;2;i1};3]2]: 

ee ee 

AS. R. 


Somites. 
Dorsal. 
Ventral, 


Size 1-3. 


ric. 26. Diagram illustrating the tuberculation of a single specimen of Pontobdella 
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 
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 small 
annulus dorsally only. Dotted lines indicate where additional annuli may or may not be 
present. See Fig. 13 (p. 409) for other abbreviations. 


4. §ild bye 


REVIEW OF THE PONTOBDELLINAE 431 


and up to six secondary ones, rarely having a total of more than fifteen. Annulus oy: 
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 Riviére 


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: 4901. 


DESCRIPTION. General characters. (Specimens not examined.) Body fusiform, 
attenuated anteriorly and circular in cross-section. Length 35-64 mm., breadth 
6-8mm. 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: r or former slightly larger. Posterior sucker circular, with 
corrugated edge and attached centrally. 


H.R P.C.R. C.R. T.R. 

eae” SS ee Ge Se (ae Faia Ren Beene 
Somites. =| I-VI Vil vill Ix x XI XI XIII 
Pera 1010 ibe 4l4 bis lial eels lelaigtislelglalslaleietsiteis. 
Sear i 104° }ofo|2jofal2iol2f2|2]2} aya] 2{2[2[2]2] 2) 2] 2] 2/2 [2 
Size 1-3. } | 

ESS Saale 

A.S. wR, 


Somites. 
Dorsal. 
Ventral. 


Size [-3. 


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). aA.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. C. LLEWELLYN 


ANNULATION. Uniannulate somites XXVI, XXVII. 
Biannulate somites XXV. 
Triannulate somites Vi, VI, XI, Xl: 
Quadriannulate somites IX, X, XITI-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:1r:2. Clitellum inconspicuous, probably comprising annuli Xb, to XIIIa,. 
Male pore opens mid-ventrally in furrow between Xla, and XIa,;. Female pore 
opens mid-ventrally in furrow between XIIla, 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 bg, 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 rugosa 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 XAV 

lets , XII. 
rriannulate somites (VI?) XI xxiv 
Ouadriannulate somites XIII-XXIII vii-x } ; 


Sexannulate somites 


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 XIIa, and XIIa, 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, > bs; —b,. In somite 
XXIV the furrow between b; and b, 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 ai annuli. They are small or 
absent on neck annuli, somite VII, the clitellar constriction (Xb,—XIIIa,), 
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,, bs; and b, 
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. 


DiaGNosis. Tuberculate Piscicolidae in which most of the somites of the 
testicular and caecal regions are pentameric. 


Pentabdella dispar (Cordero, 1937) 
Pontobdella dispay 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. C. LLEWELLYN 


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— 
1: 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 xxv 
Biannulate somites XXV ‘ ; 
Triannulate somites VII-XII, XXIV. 
Quadriannulate somites XIII. 
OQuinqueannulate 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 Xla, and XIa,, and the female pore betwen XIla, and XIlIag. 
Annulus XIla, 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,, bs, ay, b; and b, 
approximately 1:4:6:4:1. Somite XXIV consists of three annuli which are 


Somites 
Dorse! 
Vertreo! 


Site 1-3 


Somites. 


Dorset lol sta| plolalsisieis isis 
Ventral. CORIRIRSCORSLILILILILIEIED 


Size i-3. 


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. ‘‘ 4 "’ 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 


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 X XVII 
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 XIIIa, are generally smooth, while the 
other annuli are more wrinkled and bear six pairs of small sharply-pointed tubercles 
each. The b, and bs 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 XXV 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 vayner1 Baird, 1869 : 313. 


DESCRIPTION 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 1 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 Orientobdella 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 Strbarobdella. 


DISTRIBUTION. Port Jackson, New South Wales. 
Host. Not recorded. 


Pontobdella prionodiscus Schmarda, 1861 
Pontobdella prionodiscus Schmarda, 1861 : 7. 

DescrIPTION after Schmarda. Body cylindrical, length 48 mm. and breadth 
10 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, Ig10 
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. 

DistrisuTion. 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 ro, rr, 14 and 17 on fresh specimens seem to indicate, 
according to Blanchard (1894) and Harding (1910), that the species is probably 
identical with Tvachelobdella lubrica Grube, 1840 : 60, (Pontobdella littoralis : John- 
ston, 1865 and Pontobdella campanulata : Johnston, 1865, pp. 42 and 304 were also 
probably Trachelobdella lubrica). 


~=ieltl 


+176 en Ty Rae eh yee 44 Lined Oe bate 


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 labracis 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. 


KEY TO THE SPECIES IN THE SUBFAM. PONTOBDELLINAE 


1 Abdominal somites triannulate , : ‘ : Gen. STIBAROBDELLA 
— Abdominal somites quadriannulate . ; ‘ : . Gen. PONTOBDELLA 
— Abdominal somites quinqueannulate . : : : . Pentabdella dispar (p. 43 


2 Anterior sucker with a marginal fringe . ; .  Stibarobdella loricata (p. 41 
— Anterior sucker without a marginal fringe . : : 
3 Anterior sucker with a pigment patch or patches 
— Anterior sucker without pigment patches : : : ‘ : : 
4 Anterior sucker with no papillae ; : . Stibarobdella taprobanensis (p. 42 
— Anterior sucker with papillae s : : , ; 
5 Anterior/posterior sucker ratio between 1 : 2 and I: 3 

Stibarobdella macrothela (p. 49) 
— Anterior/posterior sucker ratio between 1 : 1 find 1 : 2 : 
6 Anterior sucker with two distinct pigment patches and anterior feastenion ence tio 


Cewe Aa @o) Eee, ee 


approximately 1 : I 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) 
7 Anterior sucker with papillae ° ; : 
— Anterior sucker without papillae ‘ é ; Stibarobdella moorei io: 422) 


8 9g tubercles including a mid-ventral Paberle. on the a, annulus 
Stibarobdella planodiscus (p. 418) 
6 tubercles on a, annulus 4 dorsally and 2 ventrally Stibarobdella australiensis (p. 422) 


9 Anterior sucker with fringe or welt . : é é : j ‘ ‘ ; se) 

— Anterior sucker without fringe . 4 si 
to Anterior sucker without pigment patches but an 4 ae of pois amtenion! 

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 I: 1-7 
Pontobdella rugosa (p. 432) 

1r Anterior sucker with 3 pairs of papillae, anterior/posterior sucker ratio between 
ibe 5 ANG 1-S 2) ss : . Pontobdella vosmaeri (p. 427) 

— Anterior sucker without spapillae, aitterion/posterion sucker ratio 1:1 or former 
slightly larger. ; ‘ : ‘ of 5 Pontobdella aculeata (p. 431) 


435 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 

Apatuy, 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. Proc. Zool. Soc. Lond., (1869) : 310-318. 

BENEDEN, P. J. vAN, & Hesse, C. E. 1863. Récherches 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. Museit Zool. Anat. comp. 

R. 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. 

Corpero, 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. 

Gopparp, E. J. 1909. Contributions to our knowledge of Australian Hirudinea. Proc. Linn. 


Soc. N.S.W. 34 (4) : 721-732. 
Harant, H. 1929. Essai sur les Hirudinées. Archs. Soc. Sci. méd. biol. Montpellier, 10 : 615- 


Traité de Zoologie, 5, prem. Fasc : 471-593. 

Harpinc, W. A. 1910. A revision of the British leeches. Parasitology, 3 : 130-201. 

1924. Descriptions of some new leeches from India, Burma and Ceylon. Ann. Mag. nat. 

Hist. (9) 14: 489-499. 

——- 1927. Introduction to the Rhynchobdellae. Hirudinea. Fauna of British India. 
London, 13-56. 

Herter, K. 1935. Hirudinea. Tierwelt N.u. Ostsee, 29 (6c) : 45-106. 

ICKMANN, V. V. 1941. A new Ichthyobdellid leech and its egg capsules. Pap. Roy. Soc. 


‘ 4 Tasm. (1941), 41-44. 
_ Aya —— 1946. Pontobdella tasmanica nom. nov. (Hirudinea). Pap. Roy. Soc. Tasm. (1946), 27. 

SG kaM, D. M. 1957. Some Tasmanian Hirudinea. Pap. Roy. Soc. Tasm. 91: 191-232. 

- Jonansson, L. 1896. Bidrag till kannedomen om Sveriges Ichthyobdellider. Uppsala. 119 pp. 
“" [5e1GH-SHARPE, W. H. 1925. Stibarobdella superba n.g. et sp. A new Ichthyobdellid leech. 
Cy ..e Parasitology 17 (4) : 417-420. 

~Leypic, F. 1851. Anatomisches uber Branchellion und Pontobdella. Z. wiss. Zool. 3: 315- 
324. 
Mann, K. H. 1953. The segmentation of leeches. Biol. Rev. 28: 1-15. 
Moore, J. P. 1927. The segmentation (metamerism and annulation) of the Hirudinea. 
fauna of British India. Hirudinea. London. 


: 

’ 

682 i 
Harant, H. & Grassé, P-P. 1959. Classe des Annélides achétes ou Hirudinées ou sangsues. | 


jt 4—hee ~ FBP dimeng 


re dc 


REN LEW OF THE PONTOBDELLINAE 439 


Moore, J. P. 1938. Leeches. Aust. Antarctic Exped. 1911-1914. 10 (3) : 5-16. 

1958. The leeches (Hirudinea) in the collection of the Natal Museum (Pontobdella 

(Pontobdellina) macrothela). Ann. Natal Mus. 14: 303-340. 

Moguin-Tanpon, A. 1846. Monographie de la famille des Hivudinées. Paris, J.-B. Bailliére. 

Oxa, A. 1910. Synopsis der Japanischen Hirudineen, mit Diagnosen der Neuen Species. 
Annotnes zool. japan. 7 : 165-183. 

1917. MHirudinea. In: Zoological results of a tour in the far East by N. Annandale. 

Mem. Asiat. Soc. Beng. 6: 1817. 

1929. Sur la morphologie externe de Pontobdella. Proc. Imp. Acad. Japan. 3: 90-93. 

1935. Aper¢u de la faune hirudineenne japonaise. Pvoc. Imp. Acad. Japan 11 (6) : 240- 

241. 

RicHarpson, L. R. 1950. Studies on New Zealand Hirudinea. Part (1). Pontobdella 

benhami n. sp. Trans. Roy. Soc. N.Z. 78 (1) : 97—I00. 

1953. Studies on New Zealand Hirudinea. Part (III). Bdellamaris eptatreti n. g., n. sp. 

and notes on other Piscicolidae. Tvans. Roy. Soc. N.Z. 81 (2) : 283-204. 
RINGUELET, R. 1944. Sinopsis sistematica y zoogeografica de los Hirudineos de la Argentina, 
Brazil, Paraguay y Uruguay. Rev. Mus. La Plata (N.S.). 3. Sec. Zool: 163-232. 
RIVIERE, R. 1925. Description d’une nouvelle Pontobdelle Pontobdella brumpti n. sp., Annls 
Parasit. hum. comp. 3 (3) : 292-300. 

SCHMARDA, L. K. 1861. Neue Wirbellose Thiere. I. Turbellarien, Rotatorien und Anneliden. 
Hit. 2. Leipzig: Engelman. 

ScRIBAN, I. & AutTruM, H. 1934. MHirudinea. Kukenthal-Krumbach. Handb. Zool. 2, Teil 8, 
Lief. 17, 240-251 and 338. 

SELENSKY, W. D. 10915. Etudes morphologiques et systématiques sur les Hirudinées. I. 
L’organisation des Ichthyobdellides. Petrograd (in Russian). 

Sirva, P. H. D. H. de. 1963a. The occurrence of Pontobdella (Pontobdellina) macrothela, 

Schmarda, and Pontobdella aculeata, Harding in the Wadge Bank. Spholia zeylan. 30 (1): 

35-38. 

19630. A new leech Pontobdella taprobanensis sp. nov. from the Wadge Bank. Spolia 
zeylan. 30 (1) : 39-46. 

VAILLANT, L. 1870. Contribution a l'étude anatomique de genre Pontobdella. Anmnls. Sct. nat. 
£3: I-71. 


ADDENDUM 


In preparing this review I unfortunately overlooked two species which are impor- 
tant for problems of annulation and affinities of the Pontobdellinae. 

The first, Pontobdella biannulata Moore (1957. MHirudinea. Rep. B.A.N.Z. 
antarct. Res. Exped. 1929-1931. Ser.B, 6 (6) : 99-105) has two annuli per urosome 
somite. It may now be placed tentively in Stibarobdella if the definition of that 
genus is widened to include tuberculate leeches with either three or two annuli per 
urosome somite. 

The other species, Pontobdella japomca Vasiliev (1939. The Ichthyobdellidae of 
the Far East. Works of the Karelian State Pedological Institute, 1, Biol. Ser., (1) : 
25-08), has fourteen annuli per urosome somite and is now the type of Orentobdella 
Epshtein (1962. A survey of the fish leeches (Hirudinea, Piscicolidae) of the Bering 
Sea, the Sea of Okhotsk and the Sea of Japan. Dokl. Akad. Nauk SSSR, 144 
(5) : 181-1184). Although tuberculate, it has quite large lateral vesicles and it 
seems doubtful whether it should be included in the Pontobdellinae. 


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ON THE FIRST 
HALOSAUR LEPTOCEPHALUS: 
FROM MADEIRA 


C. M. H. HARRISSON 


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BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
| EZOOLOGY Vol. 14 No. 8 
a LONDON: 1966 


eee tHE FIRST HALOSAUR LEPTOCEPHALUS: 
FROM MADEIRA 


BY 


CoM. ELCrARRISSON 


National Institute of Oceanography 


Php. 441-486 ; Plate 1; 6 Text-figures 


BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 14 No. 8 


LONDON : 1966 


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(NATURAL HISTORY), instituted in 1949, 1s 
issued in five series corresponding to the Departments 
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Parts will appear at irregular intervals as they become 
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within one calendar year. 

In 1965 @ separate supplementary series of longer 
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This paper is Vol. 14, No. 8 of the Zoological series. 
The abbreviated titles of periodicals cited follow those of 
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© Trustees of the British Museum (Natural History) 1966 


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THE BRITISH MUSEUM (NATURAL HISTORY) 


Issued g September, 1966 Price {I 35. 


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Fic. 1. Whole leptocephalus of an Aldrovandia sp. captured off Madeira. 


ON THE FIRST HALOSAUR LEPTOCEPHALUS: 
FROM MADEIRA | 


By CA Be HARRISSON © UNG Sy 
Ne L\* 


INTRODUCTION 

DurinG the autumn of 1961 the R.R.S. Discovery made a series of collections in 
the Canary Basin of the North Atlantic. Among the fishes examined in the spring 
of 1964 was an elongate larva taken by an Isaacs-Kidd midwater trawl towed for 
2 hours at a mean depth of I100 m. (601-5 fths.) and a position between 29° 50’ N., 
22° 56’ W. and 29° 50’N., 22°57’ W. This larva strongly resembled one of the 
larger ribbon-like eel leptocephali, though after preservation for 2} years in formalin 
and subsequent transfer to 70% alcohol, it was somewhat less transparent and of a 
yellowish tinge. The presence of a pair of small ventral fins combined with the 
highly characteristic pattern of opercular bones and head canals indicate that it 
can only be a halosaur. It is indeed the earliest developmental stage yet known 
for this curious group of largely benthic deep sea fishes. 


DESCRIPTION 

The larva (Text-fig. 1) is 190 mm. in length though the tip of the “ tail ”’ is missing. 
The original length must have been at least 3 or 4mm. more. The head is small, 
9-5 mm. from snout to basis cranil, 3-5 mm. in maximum depth. The body is 
flat and ribbon-like, with a gradual dorsoventral broadening behind the head, 
reaching its greatest depth (7-3 mm.) well behind the ventrals, then tapering away 
again gradually to the tip of the tail. There are some 250 myotomes, but an exact 
count is made difficult as the tail is damaged, and the myotomes become less distinct 
posteriorly. Each myotome consists of a simple V whose apex points forwards 
(Text-fig. 1). Damage also leaves intact epidermis only on the head, and for a 
short distance along the body beyond the level of the pectorals (Text-fig. 2). Over 
this whole area the skin is lightly speckled with black pigment. Summarizing the 
body proportions, the head length is contained some 20 times in the length and 
3 times in the distance, from the snout to the origin of the ventral fins, while the 
maximum body depth is twice the maximum head depth. 

The fins consist of (i) a pair of pedunculate pectorals, set somewhat below the 
mid-line of the lateral profile, and with ro rays, (ii) a pair of ventrals with 8 rays, 
and (ii) a larval fin-fold commencing a long distance behind the insertion of the 
ventrals and confined entirely to the dorsum. The “anal papilla’’ is close to the 
tip of the tail, at the level of the 207th myotome, and there is no sign of an anal 
fin-fold whatsoever, though it must be remembered the tip of the tail is missing. 
The branchiostegal rays are Io in number, and there are g rakers, I on the upper, 
and 8 on the lower limb of the rst gill arch. The ray formula so far as can be ascer- 
tained is thus: Bro; D—; P10; V8; A—. 

The head is roughly conical, tapering forwards to a pointed snout that overhangs 


ZOOL. F4, 8. 30 


440 Cc. M. H. HARRISSON 


‘ 
fefe gatas 
eee 7 46 


lom. 


Head of Aldvovandia leptocephalus enlarged to show details of lateral line canals and opercular structure. 


THE HALOSAUR LEPTOCEPHALUS 447 


the mouth. There are a number of tiny rounded mucus pores on the snout, while 
a series of pale elongate ovals marks the position of the supraorbital lateral-line 
canal. A pore just anterior to the nasal capsule marks the posterior limit of the 
adnasal canal. Much more conspicuous is the very large suborbital canal. Seen 
by transmitted light the neuromast cushions are visible as opaque dots and above 
each of these organs in the lateral canal wall is a pale elongate elipse, representing 
the curled dorsal edge of a developing lateral line scale. Seven such scales are 
visible, the most posterior, the smallest, lies at the point where the suborbital canal 
turns upwards behind the eye. Alternating with the cushions are large oval pores 
connecting the exterior with the canal lumen. A similar series of pores (likewise 
difficult to see owing to the transparency of the membrane they perforate), marks 
the position of the mandibular canal. There are no signs of the lateral line along 
the body. The opercular apparatus (Text-fig. 2) consists of a small shoulder-blade 
shaped operculum, partly overlain by a very large preoperculum which is perforated 
by endings of the VIIth cranial nerve. The suboperculum is a narrow splint of 
glassy transparency lying along the lower border of the operculum. The inter- 
operculum is a slender rod passing from the suboperculum to the hind border of the 
mandible. It is connected to the “epihyal’’. The suspensorium slopes obliquely 
upwards to the otic capsule, the dorsal end of the hyomandibular being roughly 
triangular and having a horizontal edge. The head was somewhat damaged on the 
right side and the operculum torn outwards allowing an internal view of the opercular 
apparatus. The lens of the right eye is missing. The left eye remains in better 
condition, and the diameter of the spherical lens closely approximates to the inter- 
orbital width across the frontals. The nasal capsules are placed immediately 
anterior to the orbits. The nasal rosette has 8 leaflets arranged in pairs. After 
clearing in glycerine, the structure of the auditory capsule became distinct. Three 
pale zones, and 2 clearly marked dark zones with the beginnings of a third, were 
seen in the otoliths. 

Turning to details of the body characters there is a conspicuous row of large 
ventral melanophores spaced at intervals of about I pair to every third pair of 
myotomes, though more closely spaced at the anterior end and more widely separated 
at the posterior end of the series (Text-fig. 1). These pigment spots discontinue 
close to the level of the insertion of the dorsal fin-fold. Along the ventral edges 
of the myotomes is a series of small black dots, while similar minute dots of pigment 
are arranged close to the myotome septa above the mid-line of the body. This 
pigmentation is apparently subepidermal (vide supra). There are no signs of 
developing scales. It is difficult to make out accurate details of the course of the 
gut, of the kidneys and blood vessels, due to the semi-opacity of the myotomes. 
Although the viscera are still largely displaced below the body segments, the upper 
surface of the gut is partly hidden. A thorough examination could not therefore 
be undertaken without doing what was considered as excessive damage to the 
specimen. From the partial details visible it seems probable that the duct opening 
on to the anal papilla is the renal duct. The gut appears to end blindly some 22 
segments more anteriorly. 


448 Cc. M. H. HARRISSON 


LEPTOCEPHALUS FEATURES AND METAMORPHOSIS 


Having described the chief morphological characters, the general importance of 
the discovery of this larval halosaur may now be considered. Following a discussion 
comparing modes of development in various groups of fishes, the characters of 
adult halosaurs will be compared with features in the larva, with a view to inter- 
preting structures from a morphogenetic viewpoint, as well as to consider the 
larva’s probably systematic position. 


There is a series of “ soft-finned ”’ fishes, including the Tarpons and Lady-fishes 
(Tarpon, Elops) ; Pterothrissus ; the banana fish and bone fishes (Albula, Dixonina) ; 
the eels (Apodes) ; and the gulper eels (Lyomert), all of which have a larval stage 
referred to as a Leptocephalus. The question is, whether the halosaurs also have 
a larva of this rather special type. To enlarge on this, one must first establish 
what particular features distinguish a Leptocephalus from other sorts of larva, 
then see which of these characters the only available halosaur larva has. 


The first description of a Leptocephalus was that given by Gronovius in his 
Zoophylacium of 1763. He describes specimens taken in the Irish sea near Holyhead, 
Anglesey, by William Morris and sent to him by Thomas Pennant. The characters 
Gronovius gives in his latin diagnosis are: a scaleless body and head, laterally 
flattened, large eyes and mouth, as well as a long dorsal fin fold. Subsequent 
to the studies of Delage (1886), and Grassi & Calandruccio (1893) it was realized 
that the leptocephalus described by Gronovius was in fact a larval eel, while the 
classical work of Johannes Schmidt made known in great detail the developmental 
history of Anguilla. Later, it was found that the fishes of the groups mentioned 
above also have transparent larvae, with small pointed heads which are dorso- 
ventrally much narrower than the greatly flattened body. Like eel leptocephali, 
such larvae also have a long dorsal fin-fold, an anus close to the tip of the tail and 
large larval teeth borne by the membrane-bones of the jaws. Probably in all, the 
space between the two lateral myotome sheets, above and below the notochord and 
bounded ventrally by the low-slung viscera, is filled with an acellular gelatinous 
tissue, as found by Rasquin (1955) in larval Albula. The above, then, may be taken 
as basic morphological characters common to all leptocephalus larvae. 


The halosaur larva has a long and greatly flattened body. Like established 
leptocephali, it too has a small pointed head. Its anal papilla is close to the tip 
of the tail, and there is a long dorsal fin fold, while the body broadens to a maximum 
dorsoventral extent of more than twice the head depth. The simple myotomes 
form a ribbon down each flank of the body. Above the notochord, and between 
the nerve chord and the gut, the space is occupied by a gelatinous mass. Only 
two leptocephalic characters are absent: the transparency of the body and the 
large larval teeth. The specimen under consideration was yellowish and translucent, 
but not transparent or glassy-hyaline as are typical leptocephali. However, many 
Indo-Pacific eel leptocephali are similar to this in appearance, and moreover resemble 
the halosaur larva in having a ventral series of dark spots. The absence of teeth 


ae ee ee ef ee 


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ie 


THE HAvOsSAUR LEPTOCEPHALUS 449 


seems readily understandable if it is assumed that the halosaur larva has just 
reached the stage at which it is beginning metamorphosis. This contention is, 
greatly strengthened by the following evidence. 

In the great majority of fishes the larval period is a short one, lasting a matter 
of a few weeks. Prolonged “infantilism ’’ is perhaps a general feature of fishes 
with leptocephalus larvae. At all events, in Anguilla anguilla the leptocephali are 
in their third year when metamorphosis begins (Schmidt, 1935). They then lose 
their larval teeth, while the body, following changes in the head, becomes more 
slender and less leaf-like. It is thus of great interest to find in this halosaur larva 
with a head of rather adult appearance, and strangely larval body, that the otoliths 
have three rings. If these are indeed annual rings, then not only does the develop- 
mental pattern seem similar, its timing too is alike in both eels and halosaurs. 
To recapitulate, there is evidence to suggest that the larva to hand was just beginning 
metamorphosis after a life span of three years. Probably a younger larva would 
have had large teeth as well as a more transparent and even more leaf-like body. 
There thus seems excellent justification for stating that the larval halosaur is a 
leptocephalus. 


In the general description, it was remarked that the gut may have ended blindly. 
This may seem surprising. However one of the features of metamorphosis from 
the leptocephalus both in eels and elopoids (though probably not in Lyomeri) is 
the remarkable shift forward of the position of the anus. Bertin (1926), for example, 
records a shift of some 245-345 myotomes in Nemichthys scolopaceus. Now at this 
period the leptocephali stop feeding. Rasquin (1955) thinks that in Albula the 
gelatinous tissue between the myotome sheets provides the necessary reserves at 
metamorphosis, and until feeding begins again. This is probably true for other 
leptocephali. It would not therefore seem strange in a metamorphosing larva to 
find that the non-functioning gut pinched off a posterior section before retracting, 
and re-establishing the anus in its definitive position. It seems possible that this 
is what is happening in the halosaur larva. Alternatively, what appears to be the 
blind ending of the gut may ultimately prove to be merely a gastric caecum or 
hepatic lobe. Further material is required before this can be satisfactorily decided. 
Through the kindness of Mr. Alwyne Wheeler I was able to examine a number of 
X-ray photographs of adults of 7 species of Halosaur and in all the anus occurs at 
the level of the 55th-74th vertebra (see Table I, which is further discussed on p. 458). 
Assuming that the larval halosaur had an anus opening on the anal papilla, at the 
level of the 207th myotome, there must be a shift in the position of the anus of 
some 140 myotomes in amplitude during the change from larva to adult. Similar 
hypermetamorphic phenomena must therefore occur in both nemichthyid eels and 
halosaurs. 


A fuller discussion of the processes occurring at metamorphosis in the halosaurs 
can only be made after a comparison of larval and adult features. The topic will 
be resumed after an account of the morphological characters which serve to identify 
the larva. 


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THE HAVOSAURK LEPTOCEPHALUS 451 
foe OPERCULAR STRUCTURE AND,-1TS; DEVELOPMENT 


In comparing characters shown by the larva with those of adult halosaurs, one 
may begin with features of the opercular structure and the head canals of the lateral 
line system. These are the most salient features indicating that this is indeed a 
halosaur. Adult characters seem well established in the head region long before 
the body assumes its definitive form. 

Taking first the opercular structure. Marshall (1962) has disentangled from a 
century's terminological controversy the true relation of the opercular bones in 
halosaurs by examining an Aldrovandia macrochir. He showed that all the usual 
opercular bones are present in halosaurs, but that the preoperculum grows back 
in the opercular fold to cover the suboperculum and the interoperculum more or 
less completely. Superficially only two bones are visible, a small upper disc, the 
operculum, and a large lower flange, the preoperculum (Text-fig. 4c). This condition, 
clearly visible in the larva (Text-fig. 2), is unique among fishes. It seems worth a 
short digression to discuss it in detail. 

The fish operculum is a functional unit acting as a respiratory valve and pump 
(Hughes, 1960), and as a linking mechanism allowing small muscles of the hyoid 
plate (adductor operculi, Edgeworth, 1935) to help open the lower jaw (van Dobben, 
1935). Ihe (dermal) bones are, however, of two sorts: (i) the preoperculum 
which develops in relation to the lateral line and (ii) the “ truly opercular ’”’ bones 
related solely to “ cartilage bones”’ of the hyoid arch. The operculum articulates 
to a process on the hyomandibular. The suboperculum is often attached to the 
operculum, but in Mormyrids it is hard to distinguish it from a branchiostegal ray 
and in Engraulis it is even connected to the epihyal, increasing its resemblance to 
a branchiostegal ray (Ridewood, 1904: 75). This raises the question of the develop- 
ment of the bones. If branchiostegal rays develop as procartilaginous rudiments 
can they be equated with plates of bone held not to do so? The operculum first 
appears as a cartilage in eels (Norman, 1926), and the evanescent rudiment in 
lyomerous larvae is also cartilaginous (Orton, 1963), which suggests they can. 
There is then the interopercular : commonly it is displaced anteriorly as a triangle 
of bone. According to van Dobben (l.c.) it is often connected to the interhyal 
(stylohal). In Elofs (Ridewood, 1904) the branchiostegal rays all develop into 
flattened plates of bone so there is a continuous series of similar ossifications from 
the operculum downwards, but the anteriorly displaced interoperculum does not 
appear to belong with this series of bones. It is thus interesting to note that in 
the halosaur larva the interoperculum (Text-fig. 5) develops with all the appearance 
of a branchiostegal ray. It is attached to the epihyal, with its upper end connected 
to the suboperculum, and linked to the hind end of the lower jaw by a slip of muscle, 
and a tendon which is shown in Gtinther, 1887, pl. IX, fig. 2, labelled “lig”’. In 
adult Halosaurs the interoperculum flattens out and enlarges to become a paddle 
shaped bone (Text-fig. 4a). Morphologically then, the interoperculum seems 
equivalent to a branchiostegal ray. Its forward displacement in many fishes is 
explained if one accepts that it most often belongs to that part of the hyoid arch 
kinked anteriorly as the interhyal. The connection of the interoperculum with the 


452 Cc. M. H. HARRISSON 


epihyal in halosaurs is thus unusual. In Anguilla elvers, too, the interoperculum 
is paddle-shaped (Text-fig. 3). Further, in eels the opercular fold is supported 
largely by the branchiostegal rays (Regan, 1912), while the operculum and sub- 
operculum are small. Except for the large preoperculum, the halosaur opercular 
structure thus resembles the condition found in the eels. The details seen in the 
larva and described above, correspond closely with those given by Marshall (1962), 
for an adult halosaur. 


Pop | Sop 
iop 
bic. 3. Head of an Anguilla elver showing the bone structure. Amd. = hyomandibular, 
op. = operculum, sop. = suboperculum, iop. = interoperculum, pop. = preoperculum, 


q. quadrate. 


THE PREOPERCULUM AND THE HEAD CANALS 

The preoperculum may best be considered in relation to the head canals of the 
lateral line series. Its great size in halosaurs (and in these fishes it has secondarily 
become the chief support of the opercular fold) seems related to the enormously 
enlarged mandibular canal (and not to the infraorbital canal as Marshall, 1962, 
stated). There is no connection between infraorbital and mandibular canals. 
Figures (pl. LXXXIV : 3-6) by Garman (1899) are misleading in this respect ; they 
suggest a jugular connection, present in Dipnoi but not known in any Actinoptery- 
gian. 

The thin walled and greatly expanded canals in the preopercular region are 
delicate and often damaged in preserved material. In the larva the infraorbital 
canal stops short of the preoperculum, terminating in a backwardly directed bulge 
(Text-fig. 2). In several adult Aldrovandia examined, the infraorbital canal just 
extends on to the preoperculum to end blindly in a series of finger-like processes, 
idhering closely to the preopercular membrane and the wall of the mandibular canal 


PE Eee Osa Re EE PTOCEPHALUS 453 


Sor lI. 


Fic. 4. Diagrams of adult Aldvovandia showing features of the lateral line and opercular 
structure (after Gilbert, 1905). (a) head with preoperculum as stippled outline. iop = 
interoperculum, sop. = suboperculum, op. = operculum. (b) head with inset vertical 
section xX x of the preoperculum = pop. (c) head and anterior part of body 
md.=mandibular canal, sov.=suborbital canal, md. 1.=preopercular loop of mandibular 
canal, /./.=lateral line along body. 


below (Text-fig. 4). The larval mandibular canal is evident from the series of large 
pores along the underside of the lower jaw. It curls round the end of the jaw on 
to the preoperculum, but further detail is obliterated by damage. In adult Aldro- 
vandia the mandibular canal curves back in a broad loop, over the main preopercular 
flange, and is partly overlain by a thin lamina of bone. This lamina is connected 
by a delicate strut to the main preopercular flange. The preoperculum thus has 
an I-girder cross-section with the outer lamina smaller than the main flange (Text- 
fig. 4b). Probably the lamina arises from the fusion back-to-back of two series of 
curled scale-like ossifications (similar to those in the suborbital canal), formed in 
the wall of the preopercular loop of the mandibular canal, and additionally fused 
basally to the flange. The lamina is clearly visible in the larva, as is a part of the 


454 Cc. M. H. HARRISSON 


canal which, in the adults, was seen to narrow above the loop, and open to the 
surface by a large pore near the upper margin of the preoperculum. It is evident 
that the extraordinary size of the preoperculum may be correlated in part with the 
unusually developed lateral-line canal normally associated with this bone. The 
outline of the bone corresponds with the shape of the canal-loop. Neuromast 
cushions of the lateral-line system develop in relation to endings of the facial, 
glossopharyngeal and vagus nerves. The backward growth in the opercular fold 
of numerous branches of the ramus hyomandibularis of the facial, may explain 
the origin of this loop. The differences in opercular structure between halosaurs 
and eels can thus be related to the development of the giant lateral line canals and 
the unusually rich innervation of the opercular fold in the former group of fishes. 

Little more detail of the larval head-canal pattern can be seen than has been 
described above. The suborbital and mandibular canals are clear, and the rostral 
commissure is apparent from pores on the snout. Gosline’s account (1961) of the 
arrangement in an adult Aldrovandia, may be compared with Text-fig. 2 showing the 
larval head. 


THE GENERIC AND SPECIFIC IDENTITY OF THE LARVA 


Beyond characters general to the family Halosauridae, there remains a restricted 
range of features useful at the generic and specific level for attempting to determine 
the larva. A consideration of the significance of certain adult features seems a 
necessary corollary in the following discussion. 

Since Johnson’s description in 1863 of the first halosaur, 24 other forms have 
been named, and published records and descriptions of very various excellence 
have appeared covering a total of over 400 specimens, more than 300 of which are 
from the Atlantic. The rest are from localities scattered through the tropical and 
subtropical regions: The Prince Edward Islands (Giinther, 1878), the Indian 
Ocean, (Alcock, 1889-98 ; Brauer, 1908; Norman, 1939), the Malay Archipelago 
(Weber, 1913), South Australia (McCulloch, 1926), The Philippines (Fowler, 1933), 
Japan (Giinther, 1877, 1887), Hawaii (Gilbert, 1905), and the Gulf of Panama 
(Garman, 1899). Additional observations have been made from bathyscaphes and 
by deep-sea photography [e.g. Pérés (1956), Houot (1958)]. Further details are 
given in Table III, in the map, Text-fig. 6, and in the Gazetteer (Appendix 
pp. 475-486). There has, however, been norecent review of the family Halosauridae. 

Vaillant in 1888 after studying the material brought back by the “ Talisman ” 
divided the genus Halosaurus into 2 groups according to whether or not the inter- 
orbital width was greater than the horizontal diameter of the eye. Vaillant’s group 
of species with a large interorbital width was also characterized by lacking scales 
on the vertex of the head. His other group, those with a narrow interorbital width, 
contained, apart from his new species phalacrus, only species with a scaly vertex. 
(,oode & Bean (1895) used the criterion of scaled as against scaleless vertices to 
divide the halosaurs, placing Vaillant’s phalacrus in their new genus Aldrovandia, 
along with the other smooth-headed species. Now the type of Aldrovandia phalacra 


=. 


eh eee ee 


THE HALOSAUR LEPTOCEPHALUS 455 


was slightly smaller (430 mm.) than previously described species and is perhaps 
a species generally characterized by small size. One is led, further, to wonder 
whether interorbital width in Aldrovandia does not increase with age. The levator 
arcus-palatini/hyoidei muscles which in Aldrovandia (as in Polypterus) slant forward 
and upwards to insert on the frontals, are placed more vertically in Halosaurus. 
The greater interorbital width seems related to the larger surface required for the 
muscle insertions in Aldrovandia. Perhaps, too, the wider spacing of the supraorbital 
lateral-line canals, concomitant with wider frontals, explains the differences in 
squamation. Scales are developed in Halosaurus in which the canals are close 
together, but not in Aldvovandia in which they are wider apart. 

The larva under discussion has developing scales only in the giant suborbital 
and mandibular canals, so that one cannot rely on this character here. The levator 
arcus palatini muscles (Text-fig. 5) are placed almost vertically and originate on 
the posterior border of the orbit and the lateral wall of the cranium. The supra- 
orbital canals, on the other hand, are not very close together, running almost along 
the upper rims of the orbits. One might perhaps expect a broadening of the head 
from the compressed state pre-supposed in a leptocephalus head. Also, the origins 
of the levator arcus palatini muscles are narrow crescents on the frontals of a small 
syntype of A. phalacra, suggesting that the muscle may increase in bulk during 


fap: 


dil. op. 


Fic. 5. Head of Aldvovandia leptocephalus showing musculature. m.c. = nasal capsule, 
l.a.p. = levator arcus palatini, dil. op. = dilator operculi, /.0.b. = levator operculi, 
op. = operculum, s.o.p. = suboperculum, p.f. = preopercular flange, pop. = pre- 
operculum, 2. op. = interoperculum, sov. = suborbital lateral line. canal. 


456 Cc. M. H. HARRISSON 


development. A shift forward of the origin of the muscle from the wall of the brain 
case to the dermal bones roofing the orbits, may occur at metamorphosis, and 
presumably at a time when the larva is not feeding. On the basis of criteria currently 
used to separate genera it is not possible to decide whether the larva is a Halosaurus 
or an Aldrovandia. The facts, that the interorbital width is narrow, and that the 
levator arcus palatini muscle is placed nearly vertically, seem insufficient for assigning 
it to the genus Halosaurus. 

The characters used to separate the species of the family Halosauridae, as adults, 
are largely valueless when applied to a larva in which the body proportions are 
altering to such an indisputably major extent. This is best emphasized by a com- 
parison of the body proportions assembled in Table II. Measurements for the 7 
species of Halosaur examined are set out, using the same material as detailed in 
Table I. For specimens that had obviously lost the tip of the tail, the length is 
given, followed by a plus sign. Since many had stood on their snouts in jars for 
almost a century, their rostra were bent, so that it is impossible to give reliable 
length data, irrespective of tail-truncation. Standard lengths are given to the 
nearest millimetre, but should be interpreted with caution. Lastly there is the 
factor of shrinkage following preservation. In the last century much material was 
placed in strong spirit, and “‘ hardened ’’. Johnson (1863) in his description of the 
holotype of H. oventz quotes the standard length as 18 # in. (= 464 mm.). Giinther 
(1887) gives the length of the same specimen as 17}in. (= 444mm.), and my 
measurement in 1965 showed it to be c. 440mm. Assuming comparable levels of 
accuracy in measurement, in the first 24 years after its preservation, the type had 
shrunk by c. 20 mm., and shortened by another 4mm. in the following 78 years. 
Halosaurs are rather weakly ossified fishes, with more bone in the head region 
than elsewhere, so the head is probably least subject to shrinkage except in the 
mucous-filled snout region. Shrinking along the vertebral column, offset by good 
fixation of the nervous tissue, perhaps explains Vaillant’s observation that the 
spinal-cord in a specimen of H. johnsonianus was bent forward in a pleat, beneath 
the cerebrum (1888: 182). It is plain that no great reliance can be placed on the 
proportions given in Table II, but they provide a convenient and uniform series 
of data for rough comparison. The material has been arranged in order of decreasing 
standard length, with the larval measurements placed at the bottom, so that allo- 
metric phenomena should be more readily apparent for species with adults of 
comparable size. Three features seem worthy of comment. Firstly the standard 
length: head-length (Sy: Hy), and the preventral-length: head-length (Vz: Hx) 
ratios are much higher in the larva, 20 and 3 respectively, than in any of the adults 
(maxima of 9°8 and 2-6), while the body-depth: head depth (Bp: Hp) shows the 
same trend (2-1 in the larva, and a maximum of 1:9 in the adults). This is the 
equivalent to stating that the larva is a leptocephalus (= ‘‘small head ’’, in Greek), 
The larval head is smaller in all dimensions, relative to the body, than in the adults. 
Secondly the larval body depth : head-depth ratio is closer to that of the largest, 
but more than twice the value for the smaller ‘‘ adults’’ measured. It seems that 
the larva becomes more shallow-bodied following metamorphosis, and the adults 


457 


THE BALOSAUR LEPTOCEPHALUS 


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455 C. M. H. HARRISSON 


become deep-bodied again after the abdomen has been distended by the gonads. 
Lastly, the head-length: head-depth (Hy,: Hp) ratios seem to show specific differences. 
The larval ratio appears comparable with the figures for the adults ; the precocity 
of the larval head is noted above. But as the adult snouts have been crumpled to 
such a varying degree, one can only suggest that these figures make it less likely 
that the larva is one of the following: A. machrochir, A. mediorostris or H. ovenit. 

Returning to Table I, the possible systematic value of the segmentation may 
be considered briefly. Two prefatory remarks are necessary. There is no absolute 
correspondence between myotomes and vertebral numbers. The myotomes act 
across the vertebrae, and the first vertebra counted in the radiographs appeared to 
be a hemicentrum (the basioccipital, fused to the basisphenoid?). One might 
expect at least one myotome fewer than the vertebral number in the front trunk- 
region. Also, the thickness and curvature of the body-wall means that the position 
of the pectoral girdles and the paired-fin insertions, relative to the vertebral column 
is subject to some variation dependent upon the angle at which the radiograph was 
taken, the posture in which the specimen was preserved and the state of contraction 
of the body musculature. For these reasons a direct comparison between vertebral 
numbers in the adults and myotomes in the larva, seems unjustified where the 
differences in segmentation over the range of species examined, is so slight. Secondly 
it should be noted that the lectotype of A. macrochir is that chosen by Giinther 
himself, in his final report on the Challenger fishes (1887). In view of the lack of 
exact information on the effect of muscular contraction on the position of the fins 
relative to the vertebrae, the information in Table I may simply be regarded as 
showing a general conformity of the larval segmentation with that in adult halosaurs, 
in addition to indicating the magnitude of the anal migration at metamorphosis 
(see p. 449). Perhaps, though, the rather small number of vertebrae anterior to 
the ventrals makes it less probable that the larva is either A. phalacra or A. medio- 
rostris. The data in Table I have been arranged with the “ adults ”’ of the different 
species placed in descending order for the number of preanal segments. It will be 
seen that for the adults the number of segments anterior to the dorsal and ventral 
fins decreases in this same order, with the exception that the dorsal fin is further 
from the head in A. phalacra. The present standard lengths in this Table are placed 
above; below them the first published figures, converted to mm., are given in 
parentheses. The only characters that really might seem available for both adults 
and larvae are those of the ray formula. Unfortunately almost nothing is known 
about the range of intraspecific variation, and in relying on published data which 
have not been adequately reviewed, one is apt to be engulfed in the quicksands of 
error amassed unfathomed over a century of time. Assuming all published synony- 
mies to be correct, Table III presents the available data for comparison with the 
larval fin-ray formula and branchiostegal numbers given above. Certain previously 
unpublished details could be added thanks to Dr. P. H. Greenwood and Mr. N. B. 
Marshall who allowed me to see type and other material kept in the British Museum, 
Such details are shown by an asterisk in the Table. Before looking more closely at 
the ray formulae it may be remarked that the ray count for the larval pectoral fin 


Lie Ab ossurk LEPTOCEPHALUS 459 


given above is probably a conservative figure. The radials and their rays become 
so small towards the lower border of the fin that they are exceedingly difficult to see. 

The species in Table III are arranged in descending order of branchiostegal and 
fin-ray numbers, except that with Aldrovandia the species with a narrow interorbital 
width (Hawaiian forms, and A. phalacra) are placed first, while the species affnis, 
gracilis and rostrata, which seem to form a natural group, are placed after A. medzo- 
vostvis. Where an author gives an incomplete description of a type specimen, but 
gives supplementary details for cotypes, such data are given in parentheses. Sizes 
originally cited in inches have been converted to millimetres. In general only the 
largest size is given where more than one specimen is treated. Under the head 
“ origin of material’’ only approximate information is given, as this is often all 
that is offered by early authors. Full data have been assembled in the Gazetteer 
(see Appendix). The column “nominal species and genus’ shows in brief what 
changes have occurred in nomenclature. For the sake of brevity and to avoid 
tedious repetition the full history of transfer from genus to genus is omitted. The 
next column on the right indicates the author for final recognitions of synonymy 
and gives additional brief notes where these seem necessary. Where published details 
are available, all the species of Halosaurus have a branchiostegal ray count of 12 or 
more, whereas in Aldvrovandia the count is generally smaller, with the exception of 
Hawaiian forms. Likewise in Halosaurus the number of pectoral fin rays tends to be 
higher, though apparently less markedly so. With respect to the ventral fins the 
species of both genera have between 7 and ro rays. The larva has 10 branchios- 
tegal rays (not including the interoperculum) resembling Aldvovandia. The 
remaining ray numbers would fit A. affinis, A. gracilis or A. phalacra. A. phalacra 
has been eliminated on evidence given above, so the larva may probably be referred 
to as close to the species A. affimis. Further than this it seems unwise to venture. As 
an additional comment one may note as a curiosity that in Aldrovandia the enlarged 
scales of the lateral line are spaced at intervals of roughly 1 to every 3 rows of body 
scales. The same periodicity was noted above in the distribution of ventral melano- 
phores: ir about every 3 myotomes, in the larva. 

Summarizing the data given above, the larva described is probably a metamor- 
phosing leptocephalus of some species of Aldrovandia. Its exact identity will only 
become clear when more precise accounts of the head morphology of the different 
halosaurs become available, and when the family has been reviewed. The present 
tentative determination relies largely on fin-ray and branchiostegal numbers, whose 
systematic value has not been investigated for this group of fishes. 


ON THE RECORD OF A HETEROMOUS LARVA 
7 FROM THE INDIAN OCEAN 
One may turn at this point to an interesting record published by Mead, 1965 
while the present work was in preparation. The title of the paper “ The larval 
form of the Heteromi”’ is, perhaps, misleading, as the material treated consists 
of only one larva, stated to be an Aldrovandia, and of a juvenile (postmetamorphosis) 
halosaur. Both specimens were taken in the Indian Ocean. The juvenile specimen 


C. M. H. HARRISSON 


400 


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402 C. M. H. HARRISSON 


was tenatively identified as ‘‘ Halosaurus nigerrimus "’. The latter was synonymized 
by Weber (191%3) with Aldrovandia affinis (see Table III), but Dr. S. McDowell 
(in litt.) suggests an alternative synonymy with A. phalacra. This agrees with the 
fact that Dr. A. G. K. Menon who examined the specimen of H. nigerrimus finds a 
much higher number of gill rakers than in A. affinis (see Table III). Thus on first 
appearance, the material would consist of two specimens (only one a larva), of a 
single genus of halosaurs, i.e. Aldrovandia. No evidence whatsoever is presented 
concerning the type of larva that either the lipogenyids or the notacanths possess. 

The larva is offered as being a leptocephalus, though apart from the comment 
that it bears a general resemblance to a larval eel, no mention is made of what 
specifically leptocephalus characters it has. Concerning its ordinal determination, 
one may remark that the specimen does not appear to possess pelvic fins, which all 
Heteromi do. Further, the drawings made for Mead show the nasal capsule with 
one aperture—halosaurs, notacanths, and lipogenyids have two nasal openings. 
An interesting observation by Mead is that his larva appears to have luminous 
organs. These organs are probably not present in any of the Heteromi—Brauer, 
1908 (p. rrg) considers that luminous organs reported by some early observers 
(e.g. Giinther), are in fact only neuromast cushions. The only character given for 
assigning the specimen to the genus Aldrovandia is an obscure reference to jaw 
characters said to be generically specific. As these have not been used by previous 
authors, it must be hoped that this point may be clarified when McDowell's revision 
of the Heteromi (viz. Marshall, 1962 : 261), finally appears in the Sears Foundation 
series The Fishes of the Western North Atlantic. Until then it is not possible to 
give a proper assessment of this point. In view of the fact that none of the opercular 
characters typical of halosaurs were found, and that no giant lateral line canals 
were shown to be present, the statement that the head structure was unmistakably 
that of a halosaur (Mead, p. 1) thus comes as a surprise. An incomplete ray formula 
is given, and since details of segmentation are obscured in the available figure by 
the stipple-shading, there appear to be no reliable characters from which one might 
settle the identity of Mead’s larva. It therefore seems wisest to reject the record, 
for the present, as representing a heteromous larva. Should it ultimately prove to 
be that of a halosaur, it would be of great interest, as when fresh it was transparent, 
and it is a larva with at least one enlarged tooth! (cf. p. 449 above). Further 
discussion must however, be waived, until a better description becomes available, 
and the identity of the material is properly established. 


METAMORPHOSIS IN THE HALOSAURIDAE 
After the brief review given above comparing adult characters with those in the 
larva from Madeira, one may return to the process of metamorphosis. It is not 
possible on the basis of a single specimen to give a full account of this transition 


* Since writing tus Dr. Mead has kindly allowed me to see the specimen. It is larger than the halosaur 
leptocephalus described above, and is badly damaged. This is perhaps to be expected of material from an 
!saacs-Kidd tow of nearly twelve hours! Many specimens from hauls lasting two hours are almost unrecog- 


nizable. I would suggest that the transparency of the specimen when fresh was due to the almost complete loss 
{ (probably dark black) shin. I was unable to find any enlarged teeth. Other details might have been seen ad 
leaving and dissection been carried out. 


THE HALOSAUR LEPTOCEPHALUS 463 


in the Halosauridae, but additional information may be sought from two other 
specimens recorded in the literature. These further specimens are Alcock’s “ H. 
nigerrimus ’’ (1898) of I90 mm. from 905 m. off the Maldives, and Gilchrist’s (1908) 
120 mm. specimen from North of Cape Point, South Africa, which he provisionally 
assigned to “ H. niger’’. The former was tentatively referred to A. affinis by Weber 
(1913), the latter, in like fashion, to A. macrochir by Barnard (1925), (see Table ITI). 
Alcock’s specimen is apparently an Aldrovandia as in his catalogue (1899) he says 
that there are about 30 much enlarged lateral line scales between the gill opening 
and the vent. As there are upwards of 60 myotomes anterior to the vent (Table I), 
this implies r lateral line scale per 2 myotome segments and referring back the 
comment on the usual frequency of lateral line scales (p. 459) suggests that Alcock’s 
“about 30” should be treated with caution. 

The forward shift of the anus and renal pore at metamorphosis has been discussed 
(p. 449, and Table I), as has the proportional increase in the size of the head relative 
to the body (p. 456, and Table II). Further, the body changes from its laterally 
compressed ribbon-like leptocephalus shape, to the cylindrical body typical of the 
adults. The pigmentation of the leptocephalus is confined almost completely to the 
ventral series of dark spots. Gilchrist’s 120 mm. specimen had a black head, and 
a white body. It seems quite possible that in halosaurs generally, the head darkens 
before the rest of the body, in contrast to Anguilla where the pigmentation spreads 
forward from the tail, though the ease with which halosaurs seem to lose body skin 
must be remembered. Alcock’s specimen of 190 mm. is described as being uniformly 
jet black. The small size of both these juvenile halosaurs, and the relatively large 
size (190 mm.) of the only known leptocephalus, tempts one to wonder whether 
initially these fishes “‘ grow by shrinking ”’ such as Hollister (1936) has shown to 
occur in Albula. As however, the identity of all three halosaurs concerned is un- 
certain, since the different species of Aldvovandia appear to vary greatly in adult 
size, and because of the possibility that as in eels (Bellini, 1907, etc.), so in halosarus, 
small leptocephali may give rise to males and large leptocephali to females, the matter 
can only be decided when much more material of the young stages becomes available. 

Even more striking are the changes in the arrangement of the fins. Consideration 
of the dorsal and anal fins in the discussion of the adult ray formula above, was 
omitted. The larva had no anal, and the larval dorsal is merely a long dorsal 
fold. The changes at metamorphosis are thus considerable in this respect. An 
adult dorsal fin with a short base and from ro-12 rays (see Table III) appears in 
front of the larval fin fold. The fold may disappear comparatively late in develop- 
ment in Aldvovandia—Alcock’s 190 mm. “ H. nigerrimus’’ retains a low fold of 
skin which begins at an enlarged scale two-thirds of a head length behind the dorsal 
fin and is not continued to the end of the tail—or, in Halosaurus, may be retained 
in the adult. The 394mm. holotype of Halosaurus carinicanda Alcock, 1889 is 
described as having “a low median fold of skin, (not much more than half a milli- 
metre high) . . . enclosing distant, thin, sharp, irregular indurations ’’, (auct. cit., 
P- 455). The Aldvovandia larva described here did not appear to have “ indurations ”’ 
in the fin fold ; this may be a character to be expected in larval Halosaurus. From 


404 C. M. H. HARRISSON 


Alcock’s data it seems likely that the larval fin fold is resorbed from the tail end first. 
After the anus had shifted forward, a long anal fin must develop. No signs of an 
anal fin could be seen in the present leptocephalus (the tip of the tail was damaged), 
but in the adults the anal occupies 156-256 segments in Aldrovandia (198-201 in 
Halosaurus), on the basis of evidence presented in Table I. Further changes in the 
fins appear to include an alteration in the “set ’’ of the pectorals; they are set 
rather high in the adults, and low in the larva. The change seems concomitant 
with the development of the swimbladder (cf. Harris, 1937), of which no sign could 
be detected in the larva. In eels the swimbladder first appears in the elvers. 

It is hard to guess at what time the body scales appear in halosaurs. In eels 
they appear long after metamorphosis has been completed; 2-4 years later in 
Anguilla, where they first form along the lateral line and in the mid trunk-region 
(Hornyold, 1937). Alcock (1899), describes his 190 mm. specimen as having scales 
on the temples and cheeks’’. Those of the lateral line he notes as adherent, 
the other scales he dismisses as “ deciduous ’’, which might have meant that they 
had not yet developed were it not for a figure published by Alcock, and Dr. Menon’s 
assurance that body scales are indeed still present. It may still indicate the develop- 
ment of the body lateral-line scales before the others, and in this case the sequence 
of the appearance of scales would resemble the pattern in Anguilla. 

Finally there are some metamorphic changes in the arrangement of muscle and 
bone. In the body the muscles thicken and fold forward at their dorsal and ventral 
ends so that the myotomes lose their simple V-shape noted in the larva. At the 
same time the ribs and 2 sets of intermuscular bones (see Plate 1) develop, the 
space between the larval myotomes disappears as the jelly-filling is resorbed (see 
p. 449), and the muscles meet ventrally below the viscera as these move up to 
their definitive position closer to the vertebral column which forms along the noto- 
chord. In the head, changes appear to involve the opercular bones, teeth, the 
dermal roofing of the skull and the musculature of the mandibular-plate. The 
interoperculum broadens from the narrow branchiostegal-like splint of the larva, 
to give the paddle-shaped bone seen in the adults. It is likely that a set of larval 
teeth is shed before the development of the granular adultiform dentition. The 
present specimen had no teeth, so presumably had already lost its larval set. It 
is suggested above (p. 456), that the levator arcus palatini muscles shift their insertion 
at metamorphosis. Dramatic changes including the fusion, degeneration, and the 
alteration of insertions and origins of various muscles are known to occur in amphibian 
metamorphosis (Nieuwkoop & Faber, 1956: 100), which makes it seem probable 
that analogous changes could occur in fishes. It is also suggested here that the 
frontals may broaden in Aldrovandia, though whether this happens at metamorphosis 
or after it seems uncertain. Of the smallest known halosaurs all appear to have a 
narrow interorbital width, and three at least (the Madeira larva, “ H. nigerrimus ”’ 
and Weber's (1913) “ H. carinicauda’’ of 280 mm.), have long median dorsal fin 
folds. It seems distinctly possible that a narrow interorbital width in Aldrovandia, 
and a dorsal fin fold in Halosaurus may occur as larval features retained in some 
adults by that process commonly called neoteny. 


ee 


Tae HAarosA OR LEPTOCEPHALUS 4605 


FOOD, HEE DING ANDACLASSIPICATORY FEATURES IN 
HETEROMOUS FISHES 


In a broader setting, the discovery of a metamorphosing halosaur leptocephalus 
makes clearer the ties between the Apodes and “ the Lyopomi”’ (Berg’s Halosauri- 
formes), but what of the notacanths? The latter share many osteological peculi- 
arities in common with the halosaurs. Likewise the notacanths have an eel-like 
swimbladder (Marshall, 1962). There seem good grounds for believing that nota- 
canths eels and halosaurs derive from a common stock, and all 3 may be regarded 
as members of a natural group of eel-like fishes. The bone structure of eels perhaps 
serves to isolate them somewhat from notacanths and halosaurs. A third family 
of heteromous fishes, the Lipogenyidae, is omitted from the following discussion : 
the author can add no new information on its status. 

The major differences between notacanths and halosaurs lie in the structure of 
the lateral-line canal-system and the operculum. As Marshall (1962) has shown 
the intergradation between spines and soft rays in the notacanths, and there are 
frequent references in the literature to spines in halosaurs, the justification disappears 
for separating the notacanths as spiny fishes from the halosaurs as soft-finned. 
Gosline (1961 : 36) states that the pelvic structure of notacanths is unique. This 
is not clear from his previous discussion in the same paper (pp. 17-21). If based 
on the “‘ pungent ”’ elements in notacanths, it should be borne in mind that halosaurs 
are also reported as possessing pelvic spines by many authors. Halosaurs have 
lateralis canals lying external to the scales and the opercular apparatus (cf. Gunther, 
1887 : 238-239: “luminous organs’’) and have large free branchiostegal folds. 
Notacanths have a larger operculum, suboperculum and interoperculum, and the 
preoperculum small, while the lateral line canals lie internal to the scales, both on 
the head and the body. These are considerable differences, but one may ask how far 
they are related to functional requirements. 

Baglioni (1907) divided marine fishes into four main groups according to their 
habits and noticed accompanying differences, chiefly in the branchiostegal apparatus, 
when considering respiratory mechanisms. Bottom living fishes tend to have a 
large branchiostegal apparatus, while in pelagic fishes the opercular apparatus is 
large and the branchiostegal flap small. These differences parallel those between 
notacanths and halosaurs. The following is offered as a possible interpretation, 
considering the differences in relation to feeding requirements in the two groups. 

Actively swimming pelagic fishes pursue their prey, and whether or not they 
catch it may be thought of as depending largely “‘ on who swims fastest ’’. Assuming 
this is the predator, all that is required is for it to open its mouth at the right moment, 
when, if the victim is of a suitable size, it will pass down the gullet of the oncoming 
pursuer. Water can flow over the gills automatically during swimming, and no 
extra pumping is needed. For a bottom living fish the situation is different. An 
excess of guile over muscle may be advantageous, but of no less importance is the 
possession of a large branchiostegal flap. Potential food animals crawling over the 
bottom may disappear into places not accessible to the predator. A rapid gulp, 
involving a sudden intake of water through the mouth, is thus important, and is 


4006 C. M. H. HARRISSON 


one function of a well developed flexible branchiostegal flap. The fold can be fanned 
forwards and down by the hyohyoid muscles in the web between the rays, so that the 
extrabranchial cavity expands. A flick of the web, produced by relaxation of the 
muscles joining the tips of the rays, and by a rotation of the hyoid bar by muscle 
joining it to the mandible, suddenly pushes back water from outside and behind the 
gill cavity resulting in an inrush of water through the rictus as the mouth-floor 
sinks. The development of the branchiostegal flap may therefore be important as 
part of a complex mechanism, [also involving the mouth, the shoulder girdle and 
the operculum (Tchernavin, 1953)], for the purpose of catching moving prey, in 
addition to the need for pumping water over the gills (Hughes, loc. cit.). Where 
the opercular fold is supported chiefly by the branchiostegal rays a highly flexible 
and much more readily expansible structure is achieved than when the opercular 
bones are larger and the branchiostegal flap smaller. 

Both halosaurs and notacanths are benthic fishes, but whereas notacanths can 
browse at leisure on banks of sessile sea anemones, the halosaurs appear to feed 
almost exlusively on benthic microcrustacea. Notacanth stomachs are packed with 
fragments of actinians (Tucker & Jones, 1951 ; Wheeler, personal communication). 
Records of food from halosaur stomachs have suggested that they were catholic 
feeders. Collett (r896:151) records 2 Rossia of 28mm. from the stomach of 
one A. macrochir; mud, sand, foraminifera, sponge spicules and a Cleodora shell 
from the intestine of another. The Rossia were described as “ well preserved ”’ 
and may have been swallowed in the trawl-bag as it was hauled in by the yacht 
Hirondelle. Giinther (1887 : 233) records “‘ shrimp-like crustaceans’”’ from the 
stomach of a third A. macrochiy apparently from the Marion Islands, while Bell 
(1887) describes a trematode parasite from the ureter of a halosaur and gives the 
provenance of the material as “‘ off Cape St. Vincent . . . 1,090 fths. ’’, (= 1,993 m.). 
This seems to be an error, as only one halosaur was taken by the Challenger at 
Station V, off Cape St. Vincent, and this was Giinther’s lectotype of A. macrochir, 
which shows no signs of having been dissected. Bell’s apparent mistake has been 
reproduced elsewhere (Manter, 1934: 262; Dogiel, 1964: 285). Gtinther more 
probably dissected one of the 4 A. macrochiry from the Marion Islands (taken at 
Stn. 146—1,365 fths. —2,515m.) prior to its preparation as a skeleton! Mr. 
Prudhoe, who has kindly examined the material of ‘“‘ Distomum halosauri”’, Bell, 
kept in the British Museum, suggests that it may be a Phyllodistomum, and in this genus 
those species whose life history is known (species from freshwater fishes etc.) always 
have a larval stage whose host is a lamellibranch. This parasite record thus suggests 
that A. macrochiy may also feed on bivalve molluscs. It is likely that the sloping 
levator-arcus-palatini muscles in Aldrovandia help rock the palatoquadrate back 
and forth, and a grinding mechanism of this type would seem well suited to triturat- 
ing lamellibranchs. Zugmayer (rgrr) records crustaceans and sand from the stomach 
of another A. macrochir. An Aldrovandia among material collected by the Rosaura 
expedition from the Atlantic had its stomach packed with fragments of Cumacea, 
tanaids etc. recalling Giinther’s “ shrimp-like crustacea’”’. A remarkable number 
of recognisably crustacean fragments could also be seen in the radiographs of a 


THE HALOSAURSLEPTOCEPHALUS 467 


series of halosaurs. An astonishingly clear image of a whole tanaid is shown in 
the hind gut of the type specimen of A. vostvata (Plate 1). This evidence suggests 
that crustacea form, at least, the basis of the halosaurs’ diet. The Tanaidacea live 
in tubes which they spin for themselves. While the Cumacea are highly active 
little crustaceans which swim for short distances, then burrow back rapidly into 
the silt in which they live. The great development of the sense organs of the 
lateral line, and the large contribution to the support of the opercular fold by the 
branchiostegal rays, may thus be features connected in halosaurs with catching 
moving prey, and the chief differences (vide supra) between the two groups of 
heteromous fishes would then resolve themselves as functional devices related to 
their markedly divergent food requirements. A possible difference remains in the 
mode of development. Whether or not notacanths have a leptocephalus larva, 
remains an unanswered question. N. B. Marshall (pers. comm.) has found in the 
British Museum collections a 115 mm. notacanth from Messina which is laterally 
flattened and is perhaps a young post-larva. Earlier stages remain unknown. 


Re Pin O DU ClTON. LN AALOSAURS AND OTHER FISHES WITH 
LEP LOCKE PHALUS- LARVAE 


A prolonged period of development implies a small number of generations over 
a long time-span, hence provides less material for genetic variation or natural 
selection to act upon than would be so were development and maturation more 
rapid, given in both cases a similar level of fecundity. It is thus not surprising to 
find in the groups of fishes with slow developing leptocephalus larvae a range of 
morphological oddities otherwise associated with extinct or ancient forms. (The 
gular plate of Elops, the rostral commisures of Pterothrissus, Elopidae, Megalopidae, 
Albula and halosaurs, the valved conus arteriosus also in Albula and Megalopidae 
and perhaps the extra gill bars in Saccopharynx, may serve as examples). The 
simple myotomes of the larva resemble those of the Acrania, and perhaps too the 
anomalous Silurian fossil Jamoytius kerwood: White (1946), in which Ritchie (1960) 
shows that the V-shaped smears are probably scales. (Presumably they nonetheless 
correspond to the underlying myotomes). Nor is it strange that fossil halosaurs, 
very like the living forms, are known from the Cretaceous. Balancing the long 
life-span, it appears that in those fishes with leptocephalus larvae, for which data 
are available, prodigious numbers of eggs are produced. Thus estimates of the egg 
numbers in Anguilla anguilla vary between 20 millions (quoted without reference 
in Bigelow & Schroeder, 1953) and 5-10 millions (Bertin, 1956: 77), while J. T. 
Nichols counted 12,201,94 eggs from a 142 1b. Tarpon atlanticus (cited by S. F. 
Hildebrand, 1963: 115). This must allow a maximum of genetic recombination at 
meiosis, for gametes from a single parent, but the effect will be enhanced if, as in 
Anguilla, the adults congregate to spawn, since then the recombination possible in 
the zyogote may be as between a larger number of adults assuming that the freely 
shed eggs of any female may be fertilized by sperm from a number of different males. 
This may explain how the eels which congregate to spawn have acquired many 


468 C. M. H. HARRISSON 


striking adaptive modifications of basic teleost body form in the adult in spite of 
their longevity, whereas Tarpon which probably spawns in pairs, retains many 
“ primitive '’ unmodified characters, in addition to a few advanced ones such as 
its secondary lung-like swim bladder. Acting against the variation to be expected 
from such enormous fecundity, is the length of vulnerable larval life. Only a very 
few of the immense numbers of leptocephali produced will survive to adulthood. 
Indeed the sunfishes find leptocephali sufficiently palatable to feed solely upon them 
when opportunity allows, preferring them to their more usual diet of jellyfishes. 
Grassi (1896 : 263) found sunfishes with their stomachs packed with eel-leptocephali 
in the Straits of Messina. The low “ survivorship ’’ and slow rate of development 
are probably the two most important factors producing the assortment of “ ad- 
vanced ’ and “ primitive ’”’ characters in the fishes with leptocephalous larvae. 
Now these fishes seem in the main to be of sedentary habit. Gosline (1959), for 
instance, considers that the chief characters of eels are related to their living in 
crevices. Halosaurs are benthic, gulper eels are probably not powerful swimmers. 
The whole complex appears to have renounced higher rates of evolution, and acquired 
instead, pelagic larvae that act largely as a distributive phase in the life cycle. It 
is remarkable how widespread many of the fishes with leptocephalus larvae are. 
What little information is available on breeding in halosaurs is scattered through 
the literature. It seems worth summarizing it briefly. All the records of halosaurs 
fall between the latitudes of 40° North and South of the equator, except for Giinther’s 
record of A. macrochir from the Prince Edward Islands (Marion Islands), reports of 
H. giintherit and A. gracilis from the North West Atlantic where the Gulf Stream 
carries northward water masses of more southerly characteristics, and a specimen 
of A. macrochir from off Ireland, (see Map, Text fig 6). Within these boundaries of 
latitude, the halosaurs are world-wide. The depth range for the group appears to be 
between a record maximum of 5029 m. (= 2750 fths.) for an A. rostrata (N. Atlantic), 
and a record minimum of 383 m. for an A. affinis (Timor Sea). The author is not 
aware of records of halosaurs in nets fished at shallower levels. Most of these fishes 
were not found in hauls taken above 900 m. or below 3,000 m. and are distributed 
around the lower edges of the continental shelves and along oceanic ridges. Bathy- 
scaphe observations and deep-sea photographs show that halosaurs normally swim 
just above the bottom (e.g. Pérés l.c., Marshall & Bourne, 1965). There are records 
of more than twenty-four halosaurs with eggs. At least five of these females can 
be referred to the genus Halosaurus, and some nineteen to Aldrovandia. In Halo- 
saurus Johnson's 465 mm. genotype of H. oveniit 1863, collected in February had 
eggs, some of which measure 1-0 mm. in diameter after more than a century in spirit. 
The specimen came from off Madeira, i.e. at a latitude of c. 32° N. Poll (1953) 
records a 375 mm. specimen of the same species taken on October r4th, 1948 at 
5° 39'S., which also had ripe eggs, whereas Vaillant, 1888 whose samples covered 
the period from June to August records that all the females of H. ovenii taken by 
the Talisman had small eggs. A specimen of H. johnsonianus, taken on 18th August, 
i558 is reported by Collett as having eggs at different stages of development. In 
some other fishes this condition is indicative of a prolonged spawning period. The 


469 


PRE HALOSAUR LEPTOCEPHALUS 


Fic. 6. Map showing the distribution of halosaurs from published records. 
water of a depth of 2,000 fathoms (= 3658 m.) or less. Circles indicate 
of members of the genus Aldrovandia. 


Land shown in black. The stippled area indicates 
records of Halosaurus species, triangles show records 


470 C. M. H. HARRISSON 


picture for the Indian Ocean and the Pacific is similar. A gravid female of H. 
parvipennis was taken by the Investigator at Station 122 on October 21st, 1891, 
while in the Pacific, an H. radiatus with well developed eggs was taken by the 
Albatross expedition in February, or March of 1891. Poll’s record is the only one 
for the southern hemisphere (see Gazetteer). Records of females with ripe eggs in 
the Northern hemisphere are thus grouped in the period October to March. 

Turning to the genus Aldrovandia one finds there are no records at all of females 
with ripe eggs. Two authors apparently report material of A. affinis: Alcock, 
(1889) says that his two “ H. anguilliformis ”’ collected on May 5th, 1886 had eggs, 
and Grey (1958) tells us that twelve of her “ A. pallida’ had tiny eggs when collected 
on May 26th, 1955. Collett, r896 had two A. macrochir, with unripe eggs, the largest 
of which were 0-5 mm. in diameter, collected on 31st July/rst August, 1888, and 
Zugmayer, (IgI1) examined two specimens collected on 18th August, Igto, one he 
describes as possessing eggs that were not at all ripe, the other bore eggs considered 
as not fully ripe. The same author describes an A. phalacra collected on the last 
mentioned date: it has half-ripe ovaries. 

In contrast to the genus Halosaurus, species of the genus Aldrovandia mostly 
live at greater depths, lower temperatures and higher pressures. Thus on the map, 
the triangles symbolizing records for this genus, fall in a belt closer to the deep 
ocean basins, while the circles indicating Halosaurus records, are almost all close 
to the continents. Aldrovandia has some very widely distributed species, for 
example, A. macrochir and A. phalacra (Atlantic and Indian oceans), and A. affinis 
(all oceans), whereas there are different species of Halosaurus for each ocean, and 
these are often of limited known distribution. The most widely dispersed Halosaurus 
appears to be H. ovenit known from both sides of the North Atlantic, and reported 
from points reaching from Morocco to Cape Town. 

There seem then to be differences in the reproductive biology within the family 
Halosauridae. Either the species of Aldrovandia have very much smaller eggs, or 
they migrate to particular spawning-grounds, where they have not yet been caught. 
Possibly, too, if the eggs of Halosaurus species are indeed larger, their mode of 
development is different. It may ultimately prove no coincidence that leptocephali 
of Aldrovandia are those first known for the family. In any case one may expect a 
shorter larval life span for Halosaurus species on the grounds of their more limited 
distribution. The data available suggest that female halosaurs mature early in 
life. Thus Alcock’s type of Halosaurus parvipennis was 381mm. long, while 
Garman’s H. radiatus was only 356mm. S.L. An example for Aldrovandia is 
given by Alcocks “ H. anguilliformis’”’ (=A. affinis see Table III) which were 
ovigerous at c. 356mm. S.L. (this is not a precise length: both specimens were 
brought aboard in fragments). It seems justifiable to compare the eggs of the 
genus Halosaurus, at least, with those of other fishes. Presuming that the eggs 
swell after oviposition and the formation of the perivitelline fluid they might be 
expected to be comparable in size to the pelagic eggs of eels, where diameters of 
2°40 to 2-70 mm. are recorded by Schmidt (1930) for Nessorhamphus, and 3:3 mm. 
for an unidentified eel by Beebe (1936). The present record of a larva from 1,110 m. 


Pur, BPALOSAUR LEPTOCEPHALUS 471 


(bottom at c. 3,000 m.) suggests that oviposition may be followed by a larval existence 
within the horizontal plane inhabited by the adults. 

If distribution is associated with length of larval life in fishes not otherwise 
thought to be powerful swimmers or of migratory habit, perhaps the almost ubiquit- 
ous occurrence of the notacanths may be taken as weak circumstantial evidence 
that they, too, have leptocephalus larvae. Be that as it may, the discovery of a 
metamorphosing halosaur larva adds another tessera to the mosaic showing the 
lower teleosts, as a diverse group that has at the same time frequently retained a 
basic similarity in the pattern of development from egg to adult. 


ACKNOWLEDGEMENTS 


I am grateful to my colleagues at the National Institute of Oceanography, and 
especially to Mr. R. I. Currie and Mr. P. Foxton both for enabling me to work on 
the material and for their helpful advice. Dr. N. A. Mackintosh kindly provided 
me with working-space during the preparation of this paper. In addition it is a 
pleasure to be able to thank Dr. P. H. Greenwood for his detailed reading of the 
manuscript, and Mr. N. B. Marshall who has discussed many points and corrected 
several errors. Mr. A. Wheeler has earned my gratitude with his editorial skills 
in addition to placing at my disposal the series of fine X-radiographs, from among 
which Plate I has been selected. Mr. R. H. Harris kindly lent me one of his beautiful 
alizarin preparations from which I have drawn Text-fig. 3. To Dr. S. McDowell 
my thanks are due for his stimulating correspondence. Dr. Giles Mead and Mr. 
M. J. Penrith have both lent me precious specimens, while Dr. A. G. K. Menon 
carried out an investigation of some material in the Indian Museum. Miss S. 
Hiddleston has patiently typed and checked my manuscript. To all these people 
I am most grateful. There remains finally to record my indebtedness to the library 
staffs of the British Museum at Bloomsbury, and the British Museum (Natural 
History) at South Kensington. 


SUMMARY 


1. A single specimen of a halosaur leptocephalus is described, and its characters 
compared with those of the leptocephali of other fishes. 

2. The distinctive features of the specimen are compared with details observable 
in adult halosaurs. 

3. A review is made of such systematic characters in the adults as can be observed 
in the leptocephalus. Notes of previously unpublished details based on a re-examina- 
tion of type material are given. It is concluded that the leptocephalus is an Aldro- 
vandia, close to A. affinis. 

4. An attempt is made to outline the processes occurring at metamorphosis. 

5. The feeding of adult Heteromi is considered in relation to their systematic status. 

6. The reproductive biology and distribution of the halosaurs is discussed. 

7. An effort has been made to gather together as full a series of data as possible, 
relating to published records of halosaurs. These are given in the form of a gazetteer. 


472 C. M. H. HARRISSON 


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Bexiiini, A. 1907. Expérience sur l’élevage de l’anguille en stabulation 4 Comacchio. Bull. 
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Bertin, L. 1926. Les migrations de l’anus au cours de la métamorphose chez les poissons 

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1956. Eels: a biological study. Cleaver-Hume Press, London, 192 pp. 

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Bussinc, W. A. 1965. Studies of the midwater fishes of the Peru-Chile Trench. Antarctic 
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Cottett, R. 1896. Poissons provenants des Campagnes du yacht l’Hirondelle 1885-1888. 
Rés. Camp. Sci. Albert I** Monaco 10: 198 pp. 

De.ace, Y. 1886. Sur les relations de parenté du Congre et du leptocephale. C. yr. hebd. 
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Docrer, V. A. 1964. General Parasitology. Oliver & Boyd: Edinburgh and London. 
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EpcewortH, F. H. 1935. The Cranial muscles of vertebrates. Cambridge University 
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FowLer, H.W. 1933. Description of new fishes obtained 1907-1910, chiefly in the Philippine 
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GARMAN, S. 1899. Reports on an exploration off the west coasts of Mexico Central and S. 
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Gitpert, C. H. 1905. The aquatic resources of the Hawaiian Islands, Section II. The 
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THE HALOSAUR LEPTOCEPHALUS 473 


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Gitt, T. N. 1881. Diagnosis of new genera and species of deep-sea fish-like vertebrates. 
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— & CALANDRUCCIO, S. 1893. Ancora sullo sviluppo dei Muraenoidei. Terza nota pre- 
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ZOOL. 14, 8: 32 


474 C. M. H. HARRISSON 


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Albulidae ... Proc. zool. Soc. London 1904: 35-81. 
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Scumipt, E. J. 1930. Nessorhamphus, a new cosmopolitan genus of oceanic eels. Vidensk 
Meddr. dansk. naturh. Foren. 90 : 371-375. | 
—— 1935. Danish Eel Investigations. Carlsberg Foundn. Copenhagen; 16 pp.(Trans. fm. 
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TCHERNAVIN, V. V. 1953. The feeding mechanisms of a deep-sea fish Chauliodus sloam 
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Sci. Albert I* Monaco 35, 174 pp. 


SUPPLEMENTARY REFERENCES 


(Selected references not quoted in the text and used solely for data on distribution). 


Atcockx, A. W. 1902. A naturalist in Indian Seas. London. 

EscuMEyer, W. A. 1965. Station data errors for fishes collected by the steamer Blake. 
Copeia 1965 (No. 2) : 236-238. 

Korrorep, E. 1927. Fishes from the sea bottom. Rep. ‘‘ Michael Sars”’ N. Atl. Deep-Sea 
Exped. 1910, 4(1) : 1-148. 

Parrait, J. T. 1884. Mission Scientifique du Talisman dans l’océan Atlantique Annls 
hydrager. Paris. 1883. 5 (2°™* Sev): 258-311. 

Parr, A. E. 1937. Concluding report on fishes. Bull. Bingham oceanogr. coll. 3 (7): 79 pp. 

RICHARD, J. 1939. Liste générale des stations des campagnes scientifiques du Prince Albert 
de Monaco. Rés. Camp. Sci. Albert I** Monaco 89. 348 pp. 

Roucu, J. 1948. Stations Hydrologiques des Campagnes Scientifiques du Prince Albert 
Ir, Rés. Camp. Sci. Albert I** Monaco 108. 26 pp. 


THE HALOSAUR LEPTOCEPHALUS 475 


Route, L. 1919. Poissons provenant des campagnes du yacht Princesse-Alice 1891-1913, 
et du yacht Hirondelle II (1914). és. Camp. Sci. Albert I*° Monaco 52: 190 pp. 

—— & ANGEL, F. 1933. Poissons provenant de Campagnes du Prince Albert I¢ de 
Monaco. Rés. Camp. Sci. Albert I°* Monaco 86: 115 pp. 

SMITH, S. (et al.). 1889. Appendix E. pp. 1-144. Lists of the dredging stations . . . in the 
Atlantic and Arctic oceans. Rep. U.S. Fish Comm. 1886. 

SPRINGER, S. & BuLiis, H. R. 1956. Collections by the “‘ Oregon ’”’ in the Gulf of Mexico 
1950-1955. Spec. scient. Rep. U.S. Fish & Wildlife Serv. 196: 134. pp. 

TANNER, Z. 1886. Report of the work of the U.S.F.C. steamer Albatross for the year ending 
December 31st 1884. Rep. U.S. Fish Comm. 1884. 

TOWNSEND, C. H. & NicnHots, J. T. 1925. Deep-sea fishes of the ‘‘ Albatross ’’ Lower Cali- 
fornia Expedition. Bull. Am. Mus. nat. Hist. 52: 20 pp. 

TuckER, D. W. 1954. 4. Report on the fishes collected by S.Y. “‘ Rosaura’”’ in the North 
and Central Atlantic, 1937-38. Bull. Br. Mus. nat. Hist. (Zool.) 2 : 163-214. 


APPENDIX 


A GAZETTEER. FTO.HALOSAUR RECORDS 


In order to summarize the fragmentary information on the reproductive biology 
of the Halosauridae it was found desirable to gather together as full a series of data 
as possible from published records. The immense labour of piecing together these 
results from station lists, hydrographic papers, studies on bottom deposits, annual 
reports and charts, leads to the conclusion that it would be valuable to print what 
has been gathered in this way, if only to save others from this onerous task in the 
future. Considerations of space preclude the listing of all the papers consulted for 
this part of the work. A selection of the most important works is given above after the 
main body of references. Special notice should however be given to the following 
points. As Eschmeyer (1965) has shown, many of the stations assigned to Blake 
material by Goode & Bean (1895) and printed in roman numerals in “‘ Oceanic 
Ichthyology ’’, are in fact erroneous. Probably the same applies to specimens 
from “‘ Albatross”’ stations. It has not been possible at present to trace all such 
errors. In general the roman numerals have simply been sought against the 
equivalent arabic numbers given in the lists of Smith (1899). An additional 
record of A. macrochir which does not appear in Goode & Bean, has been added 
from Tanner’s (1886) report on the work of the ‘‘ Albatross’’. Gill’s records of 
“ Halosaurus gooder’’ have been combined with Goode & Bean’s data for A. macro- 
chy: Gill states how many specimens he saw, Goode & Bean do not. This means 
that the numbers of macrochir for the stations concerned are minimal figures, as in 
some cases specimens formerly separated into the supposedly different forms macro- 
chr and “ goodet’’, occurred together, but only Gill’s data for numbers of “ goodez ”’ 
are available. It should be noted that the Talisman station-numbers given by the 
biologists concerned with working up the material from the 1883 expedition, are 
printed in roman type, and include secondary substations. The numbering given 
by the hydrographers (Parfait, 1884) is, on the other hand, in a simple continuous 
series of arabic numerals. Caution is therefore necessary in tracing data when 
referring to the hydrographic lists, from a series of biological records. In spite of 


476 Cc. M. H. HARRISSON 


the agreement at the Washington meridian conference in 1884 to quote longitudes 
from Greenwich, the publications on this expedition, even though they appeared 
some years later than this date, still quote longitude from Paris. Smith gives correct 
longitude data for both conventions in most (not all) instances, but follows the 
French hydrographers in printing simplified station numbers which thus differ from 
those given in Vaillant’s (1888) account of the fishes. The early reports on the 
expeditions of Prince Albert | of Monaco, also quote station-position longitudes from 
Paris, but corrected data quoted from Greenwich are to be found in Richard (1934). 
It will be seen that the species occur in the tables in the order of their first discovery 
in the relevant ocean, with data for species of Aldrovandia following records for the 
genus Halosauras. Finally, the asterisks against some bottom temperatures given 
in parentheses indicate information cited from a different, but closely adjacent 
station, of comparable depth. 


KEY TO ABBREVIATIONS 


S red cl. clay 

br. brown m. mud 

ye. yellow for foraminifera 
gn. green glob. oz. globigerina ooze 
ol. olive Oz. ooze 

blu. blue r radiolaria 
gy. grey bar. nods. _ barytes nodules 
bl. black mang. manganese 
wh. white bm. bottom 

-sh. ish hd. hard 

spkd. speckled shay. shaley 

It. light bkn. broken 

dk. dark shlly. shelly 

rks. rocks cs. coarse 

sts. stones th. thick 

sha shale sy. sandy 

shr. deb shore debris my. muddy 

shs. shells sft. soft 

cor. S. coral sand fn. fine 

vol. s. volcanic sand reg region 

s. sand w. with 


7, 


BA Siete eh 


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PLATE 1 


X-radiograph of type of Aldrovandia rostrata, showing intermuscular bones and the clear 
image of a tanaid (=?), in the hind-gut. 


Pi A haa 


Bull. B.M. (N.H.) Zool. 14, 8 


INDEX-1TO° VOLUME 2a 


The page numbers of the principal references and the new taxonomic names are printed in bold type. 


For index to Number 2 see page 52 and to Number 5 see page 3606. 


aculeata, Pontobdella ¢ . 431-432 
agassizzii, Membraniporella_ . : . 68 
Aldrovandiasp. . : . 441-486 
amplectens, i ioraapors : : « “65 
australiensis, Stibarobdella ; . 422-423 
biannulata, Pontobdella ; : -. 430 
biauriculata, Hippopleurifera . : 73-74 
bimaculata, Stibarobdella A . 415-418 
clavata, Salicornaria : ‘ : 71-72 
corrugata, Bifaxaria E ; : ' 58-61 
crassimarginata, Membranipora, vay. erecta 65 
deconiniki, Bathylaimus s ‘ 103-105 
dispar, Pentabdella ; . 433-435 
dura, Rhandodemania : : 91-92 
frica, Mesacanthion . ; é 87-88 
fuegensis : : : 68—69 
hexagona, Me pabbranipora. ; : 66-67 
Hippopleurifera. : ‘ ; 73-75 
Ironidae ‘ : ‘ ‘ 92-95 
japonica, Pontobdella : ; ‘ 2" 436 
juliani, Plectolaimus . : . 100-102 
Leptolaimidae : 5 . 100-102 
Leptosomatidae . ; ; : 83-92 
leucothela, Pontobdella . 3 : . 436 
longisetosum, Trileptium . ; 85-87 
loricata, Stibarobdella_. : . 413-415 
macrothela, Stibarobdella 393-404, 405-411 
mawsonae, Weisoncholaimus : 97—100 
melolontha, Lepralia P : : 63-64 
moorei, Stibarobdella P : ; . 422 
multidentata, Lepralia_ . 64 
muricata, Pontobdella oe oh 423-427 
murphyi, Metoncholaimus . : 95-97 
nodosa, Siphonoporella_ . : ; uae 
normani, Micropora 2 : y 69-70 


octodon, Flustra . : : : 62 


ogdeni, Pheronous 93-95 
oliffi, Epacanthion 88-91 - 
oligothela, Pontobdella . 436 
Oncholaimidae 95-100 
Ophibdella 437 
owenii, Lunulites é 64 
papillata, Bifaxaria 58-61 
Pentabdella 433-435 
pilosa, Membranipora. var. maltiseiats 67-68 
Piscicolidae 389-439 
Pheronous 92-95 
Philetor 371-387 
planodiscus, Srnaropdeln 418-420 
Plectolaimus ; . 100-102 
Pontobdella . Vora 33, 435-437 
Pontobdellinae : 389-404-439 
prionodiscus, Pontobdella 436 
pulchra, Hippopleurifera 75 
rayneri, Pontobdella "N45 
rohui, Philetor 374-378 
rohui, Phitetor rohui 378-379 
rugosa, Pontobdella 432-433 
sedgwicki, Hippopleurifera Fens 
Stibarobdella 405-423 
sudafricana, Platycoma 83-85 
taprobanensis, Stibarobdella 421-422 
tasmanica, Stibarobdella 420-421 
tetajamensis, Pontobdella . 436 
Tripyloididae 103-105 
tryoni, Oedura 1-14 
verecundus, Philetor fone 379-380 
Vespertilionidae 371-387 
vittata, Pontobdella . 437 
vosmaeri, Pontobdella 427-431 
Wiesoncholaimus 97-100 


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