Bulletin of the 

British Museum (Natural History) 


The cranial muscles and ligaments of 
macrouroid fishes (Teleostei: Gadiformes); 
functional, ecological and phylogenetic 
inferences 


Gordon J. Howes 


Zoology series Vol 54 No 1 25 February 1988 


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World List abbreviation: Bull Br. Mm. not. Hist. (Zool.) 


Trustees of the British Museum (Natural History), 1988 

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ISSN 565050370 

ISBN 0007 1498 Zoology series 

Vol 54 No. 1 pp 1-62 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD Issued 25 February 1988 


The cranial muscles and ligaments of macrouroid fishes 
(Teleostei: Gadiformes); functional, ecological and 
phy logenetic inferences 2 5 F E B 1988 

Gordon J. Howes 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 


Contents 


Introduction 


List of specimens used 

Abbreviations used in the text figures 


Cranial ligaments 


Ligaments of the upper jaw and pterygoid bones 
Ligaments of the lower jaw and opercular bones 


Cranial muscles 


The adductor mandibulae and muscles of the suspensorium in macrouroids 


IBRITISH MUSEUI 

3 (NATURAL HISTORY) 

6 25FE8S938 


6 

8 PRESENTED 
10T 


10 

Macrourinae 12 

Macrouroidinae 15 

Comparisons with gadoids 16 

Summary and discussion of the muscles associated with the jaws and suspensorium 32 

Muscles of the hyoid region 42 

Ventral gill-arch muscles 46 

Dorsal gill-arch muscles 48 

Eye muscles 48 

Functional and ecological inferences 49 

Jaw protrusion mechanisms 49 

Hyoid-opercular mechanisms 53 

Pharyngeal mechanisms 54 

Trophic strategies 55 

Taxonomic and phylogenetic inferences 57 

IntrarelationshipsoftheMacrouroidei 57 

Conclusions 59 

Acknowledgements 60 

References 60 


Introduction 

If overall oceanic diversity could be expressed in numbers of individuals and species, rat-tails 
would surely emerge as the most diverse family of benthopelagic fishes. 

So wrote Marshall ( 1 979) of the Macrouridae, a group assigned to that ill-defined assemblage of 
higher euteleostean fishes known as the Paracanthopterygii. 

Marshall (1965) had earlier noted the sparsity of knowledge of the morphology and biology of 
macro urid fishes. Since then, the studies by Okamura ( 1 91Qa & b) have contributed substantially to 
filling this gap. However, since Okamura studied only Japanese macrourid fishes, the work is 
taxonomically limited because morphologically more diverse taxa occur outside Japanese waters. 
The data Okamura has published, nevertheless are substantial enough to provide a foundation for 
further anatomical studies, particularly those aimed at producing cladistic analyses, which so far, 
have not been applied to macrourid taxa. 


Bull. Br. Mm. not. Hist. (Zool.) 54(1): 1-62 Issued 25 February 1988 


2 G. J. HOWES 

Since Gilbert & Hubbs (1916) published their subfamilial arrangement of macrourid fishes, 
there have been few major taxonomic changes. Apart from the recognition of additional sub- 
families (Parr, 1946) and the elevation of the Macrouroidinae to family level (Okamura, 19700 & 
b), the most noticeable rearrangement was that of Marshall (1966) who recognised the non- 
monophyletic nature of the family. Marshall (op. cit.) found that three taxa, previously recognised 
as macrourids, viz: Steindachneria, Lyconus and Macruronus, shared more characters in common 
with the gadoid family, the Merlucciidae. Although not all of Marshall's chosen characters are 
synapomorphic for merlucciids, there is every reason to agree that these three genera do not belong 
with the macrourids (see Cohen, 1984; Howes, 1988). 

Most recent authors recognise four macrourid subfamilies (see Cohen, 1984), viz: the 
Bathygadinae, Trachyrincinae, Macrouroidinae and Macrourinae, the latter containing the 
majority (ca 30) of genera. Howes (1988), based largely on the data presented here, has challenged 
this concept of the Macrouridae (see below). 

Most macrourid taxonomy has been based on ecological-evolutionary premises. For example, 
Okamura (19706) viewed his hypothesised phylogenetic polarity of primitive to derived taxa as 
reflecting both ecological groups and an evolutionary sequence. McLellan (1977: 1034) devised an 
evolutionary scenario, based on her study of macrourid morphology and ecology, that reflected the 
invasion of continental slopes and deep ocean basins by taxa derived from pelagic ancestors; again, 
a sequence supposedly reflected in extant ecological groupings and ontogenetic development. 

As pointed out by Marshall (1965; 1979) most macrourids are found in tropical seas but some 
areas contain more speciose taxa than others, e.g. the Sulu Sea, Gulf of Mexico and Caribbean. 
Most species are confined to continental slopes and few are common to more than one ocean. 
Marshall (1973; 1979) has hypothesised that the amphi-Atlantic distribution of 33% of the total 
Atlantic species and subspecies, might be attributable to continental drift. Merrett et. al. (1983) 
also note that species common to the Atlantic and Indian Oceans are inhabitants of continental 
slope, abyssal and pelagic regions, a distribution seeming to indicate a more general underlying 
factor than simply one of random dispersal. 

In attempting to explain macrourid ecology and distribution, none of these authors has asked a 
fundamental taxonomic question; how closely related are the taxa under consideration? As yet, 
there exists no rigorously structured hypothesis of macrourid relationships. 

Since previous studies of macrourid morphology have, to a large extent, been concerned with 
feeding mechanisms and because phylogenetic interpretations have been based on comparisons of 
those mechanisms, it is the objective of this study to re-assess the morphological basis of those 
ideas. The jaw musculature of some macrourid taxa has been described by McLellan (1977) and 
Casinos (1978), these descriptions were, however, made more from a functional rather than a 
taxonomic and phylogenetic viewpoint, and a taxonomically restricted range of taxa were used. A 
comparative analysis of macrouroids and other paracanthopterygians would, it was hoped, reveal 
morphological patterns that might indicate both related groups within the suborder and the 
relationships of macrouroids with other gadiforms. The limitations of using a single character 
complex for this purpose are well realised by the author, but past experience with basal euteleosts 
(Howes, 1984; 1985) has indicated that cranial muscles can provide rewarding information on 
which to base phylogenetic interpretations. 

Where availability of material allowed, at least three specimens of each taxon were dissected to 
check the variability of the character described. 

Okamura (19706) has described other character complexes (osteology, scales, brain mor- 
phology, structure of light organs) and these have been used to a certain extent to test the con- 
gruency of relationships arrived at through the myological study. However, these other characters 
must themselves be evaluated by out-group comparisons, and it is evident in Okamura's analysis 
that many characters used to define sub-groups are plesiomorphic for Gadiformes. It remains, 
therefore, for future studies to make polarity assignments to osteological and other characters in 
order to produce a more refined hypothesis of macrourid interrelationships. 

Howes (1988) in an account based principally on the findings presented here has reviewed the 
relationships of macrouroids and gadoids, and shown that the Macrouroidei (sensu Cohen, 1984) 
and the Macrouridae are non-monophyletic groups. Although, in that previous study, clades were 


MACROUROID FISHES 3 

identified they were not given formal taxonomic status. In this account two of those clades are 
recognised as families, namely, the Bathygadidae and the Trachyrincidae. More complete diag- 
noses and taxonomic reviews of both are in preparation. Since the Macrouroidei was restricted to 
containing a single family the Macrouridae (Howes, 1988), the terms macrouroid and macrourid 
are interchangeable. However, in this text the term macrouroid is used when making coordinate 
comparisons with Gadoidei (i.e. gadoids). 

Classification used in this text 
Suborder: MACROUROIDEI 
Family: Macrouridae 

Subfamilies: Macrourinae & Macrouroidinae 

Suborder: GADOIDEI 

Family: Trachyrincidae, Bathygadidae, Moridae, Melanonidae, Steindachneriidae, Euclichthyidae, 
Merlucciidae, Gadidae, Ranicepitidae, Lotidae, Phycidae, Muraenolepididae& Bregmacerotidae 

List of specimens used 

All the specimens used in this study are in the collections of the British Museum (Natural History). Type of 
preparation is indicated as CS = cleared and stained; D = dissected; SK = dry skeleton. 

MACROUROIDEI: Abyssicola macrochir, 1938.6.23: 12-13 (D); Cetonurus globiceps, 1986.4.22: 4-5 (D); 
Chalinura mediterranea, 1986.4.22: 3 (D); Chalinura profundicula, 1986.4.22: 9; Chalinura cf. Simula, 
1967.12.11: 2 (D); Coelorinchus caribbaeus, 1963.2.25: 244-250 (D; CS, 185mmTL); Coelorinchus 
coelorincus, 1905.2.2: 18 (SK); Coryphaenoides rupestris, 1897.12.9: 82 (SK); Coryphaenoides 
anguilliceps, 1981.7.14: 1-4 (D); Coryphaenoides mexicanus, 1971.10.22: 24-25 (D); Cynomacrurus piriei, 
1930.1.12: 952; Echniomacrurus mollis, 1967.12.11: 3^4 (D); Hymenocephalus italicus, 1973.3.5: 7-10 (D); 
Kumbadentoni, 1961.1.30: 6 (Holotype; superficial examination); Lionurus carapinus, 1934.12.19: 33-34 (D); 
Macrosmia phalacra, 1980.12.31: 2 (Paratype, D); Macrouroides inflaticeps, 1939.5.24: 684 (D); Macrourus 
berglax, 1965.6.22: 8-9 (D); Malacocephaluslaevis, 1960.12.20: 2-3 (D); 1904.1 1.30: 33 (SK); Mataeocephalus 
microstomus, 1939.5.24: 723-24 (D); Nematonurus armatus, 1986.4.22: 1-2 (D); Nezumia aequalis, 1973.3.5: 
60-64 (CS, 130mm, tail broken); Nezumia hildebrandi, 1963.2.25: 138-153 (D); Odontomacrurus murrayi, 
1967.12.11: 5 (D); Sphagemacrurus hirundo, 1934.12.19: 30 (D); 1986.4.22: 6-7 (D); Squalogadusmodificatus, 
1963.2.1: 10 (D); Trachonurusvillosus, 1963.2.25: 226-228 (D); Ventr if ossa accident alls, 1965.2.25:61-71 (D; 
CS, 190mmTL). 

GADOIDEI: Antimorarostrata, 1903.9.29: 7 (D); 1986.4.22: 10-11 (CS); Austrophycis marginata, 1936.8.26: 
424-431 (D; CS); Bathygadus favosus, 1963.2.25: 28-30 (D); Bathygadus macrops, 1973.3.5: 3-6 (D); 
Bathygadus melanobranchus, 1969.6.26: 3227-3231 (D, CS); Bathygadus vaillanti, 1963.2.2: 31-35 (D); 
Bregmaceros atlanticus, 1984. 1 1 . 14: 4 (D); Bregmaceros madellandi, 1939.5.24: 792, 799 (D); Brosme brosme, 
1892.6.8: 9 (SK); Ciliata mustela, 1983.8.3: 13-26 (D); Enchelyopus cimbrius, 1980.12.18: 3-12 (D); 
Eudichthys polynemus, 1986.5.14: 1-3; 4-9 (D); Gadomus longifilis, 1963.2.25: 7-17 (D; CS, 190mmTL; 
1890.6.16: 43 (SK); Gadus morhua morhua, 1971.2.16: 634-635 (D); 1971.2.16: 628-633 (CS, 81 mmSL); 
Gadus morhua callaris, 1985.9.6: 7-14 (D); Gaidrospsarus mediterraneus , 1971.10.7: 65-77 (D); uncat. (CS, 
122, 54mmSL); Halargyreus affinis, 1973.10.29: 384-440 (D; CS, 117mmTL); Lepidion eques, 1981.3.16: 
422-427 (D); 11981.3.16: 437^40 (CS, HOmmTL); 1902.10.30: 6 (SK); Lota lota, 1953.6.26: 15-18 (D); 
168.6 (SK, skulls only); Lotilla marginata, 1974.9.28: 6-7 (CS, 1 18 mm TL); Lyconus brachycolus, 1907.6.20: 
15 (Holotype, partly dissected); Macrur onus mage llanicus, 1936.8.26: 352-357 (D); Macruronusnovozealandi, 
25, 120 (SK); Merlangius merlangus, 1971.2.16: 329-331 (D); Melanonus gracilis, 1930.1.12: 933 (D); 
Me lanonus zugmayeri, 1981.3.16: 377 (D); 1986.4.22: 8 (CS, HOmmTL); Merluccius merluccius, 1963.5.14: 
94-109 (D); 1971.7.21: 44-57 (CS, 130mmTL); Merluccius productus, 1896.9.25: 6 (SK); Molva molva, 
1976.6.29: 2-5 (D); Mora moro, 25.370 (SK); Muraenolepis microps, 1937.7.12: 24-29 (D); 1937.7.12: 11-17 
(CS, 95mmTL); 1937.7.12: 24-29 (skull); Physiculus argyropastus , 1901.1.30: 22 (SK); Phycis blennoides, 
1973.10.29: 4411-448 (D); 1976.7.30: 119 (CS); 1898.4.30: 14 (SK); Pseudophydsbrevisculus, 1873.12.13: 30 
(SK); Phycis phycis, 25.400 (SK); Raniceps raninus 1967.1.1: 4 (D); 1893.7.6: 2 (D) 1971.2.16: 640 (CS, 
40mmSL); 1864.8.26: 3 (SK); Salilota australis, 1936.8.26: 394-404 (CS, 58mmTL); Steindachneria 
argentea, 1963.2.25: 335-339 (D); 1963.2.25: 344-354 (CS, 130mmTL); Trachyrincus trachyrincus , 
1904.11.30: 34-35 (D); 1976.7.30: 42-53 (D, CS, HOmmTL); 1888.6.15: 7 (SK); Urophycis regia, 1985.6.6: 
109-11 9 (D). 


4 G. J. HOWES 

OUT-GROUP SPECIES: Atherina presbyter, 1983.4.21: 28-37; Aulopus filamentosus , 1953.11.1: 10-13; 
Brotula jayakari, 1891.2.9: 30 (SK); Cataetyx messieri, 1936.8.26: 1060-61 (D); Centropomus ensiferus, 
1984.8.8: 85-95 (D); Cynoscion jamaicensis, 1961.9.1: 107-113 (D); Dicrolene introniger, 1939.5.24: 
1441-1444 (D); Diplacanthopoma brachysoma, 1972.10.24: 4 (D); Electrona antarctica, 1948.5.14: 128-138; 
Eleotris obscurus, 1903.5.14: 93-99 (D); Esox lucius, 1971.1 1.19: 45^*6 (D); Genypterus blacodes, 1936.8.26: 
1052-57 (D); 1898.6.17: 73 (SK); Glyptophidiummacropus, 1939.5.24: 1456-1465; Gobiesoxnudus, 1985.3.18: 
110-114 (D); Gobius guineensis, 1984.7.29: 1021-22 (D); Harpadon nehereus, uncat. (D); Hoplostethes 
melanopus, 1939.5.24: 817-8 (D); Lampanyctus crocodilus, 1976.7.30: 26-33 (D); Lamprogrammus niger, 
1939.5.24: l4S3-S7(D);Lophiodesmutilus, 1939.5.24: 1 869-75 (D);Lycodesfrigidus, 1969.6.26:3145^9(0); 
Monomitopus metriostoma, 1964.8.6: 43-46 (D); Ophidian rochei, 1971.12.17: 6-8 (D); Percichthys trucha, 
1981.10.14: 28 (D); Percopsis omiscomayus , 1973.3.20: 468 (D); Photichthys argenteus, 1930.1.12: 299-306; 
Plagioscion squamosissimum, 1970.4.2: 5-8 (D); Pogonias chromis, 1886.1.21: 1 1-13 (D); Polymixia nobilis, 
1862.4.22: 17-18 (D); Porichthys porosissimum, 1948.8.6: 1460-72 (D); Senanus cabrilla, 1960.6.10: 6-8 (D); 
Siniperca knerii, 1981.2.3: 1-4 (D); Stephanoberyxmonae, 1972.10.24: 2-3; Tilapia mariae, uncat. (D). 

Abbreviations used in the text figures 

NB. Scale bars in all figures are in divisions of 1 mm. 

Al, Ala, Alp, Aly, A2, A2d, A2v, A3, Aa> Divisions of the adductor mandibulae musculature 

Aa Anguloarticular 

aap adductor arcus palatini muscle 

ad adductores muscle 

AH Anterohyal 

bf buccalis facialis of trigeminal nerve 

bpm bucco-pharyngeal membrane of 1 st gill-arch 

Bsr Branchiostegal membrane 

Cb Ceratobranchial 

ce chondroid element 

Cmb Coronomeckelian bone 

Cmc Coronomeckelian cartilage 

ct connective tissue 

De Dentary 

Dh Dorsohyal 

do dilatator operculi muscle 

Eb Epibranchial 

Ent Entopterygoid 

epx epaxialis muscle 

ey eyeball 

fA2 fascia of muscle A2 

Hb Hypobranchial 

ht heart 

hyab hyohyoideus abductores muscle 

hyad hyohyoideus adductores muscle 

Hyo Hyomandibula 

H yop Opercular process of hyomandibula 

ica infracarinalis anterior muscle 

i A 1 P internal aponeurosis of muscle A 1 P 

10 Interoperculum 

im intermandibularis muscle 

lap levator arcus palatini muscle 

le levator externus muscle 

Let Lateral ethmoid 

Icdh ceratobranchial-dorsohyal ligament 

lee lateral ethmoid-entopterygoid ligament 

lei entopterygoid-infraorbital ligament 

lep lateral ethmoid-palatine ligament 

les lateral ethmoid-suspensorial ligament 

11 levator internus muscle 

lip interopercular-preopercular ligament 


MACROUROID FISHES 

lla labial ligament 

Imh mandibulo-hyoid ligament 

Imi mandibulo-interopercular ligament 

1mm maxillo-mandibular ligament 

Imn maxillary-nasal ligament 

Imp maxillary-premaxillary ligament 

Imq mandibulo-quadrate ligament 

lo levator operculi muscle 

Ipl palatine-lachrymal ligament 

Ism supramaxillary ligament 

Isc semicircular ligament connecting 3rd hypobranchials 

ludh urohyal-dorsohyal ligament 

1VII maxillary-rostral cartilage ligament 

1IX maxillary-premaxillary ligament 

IX palatine-maxillary ligament 

1X1 ethmoid-maxillary ligament 

1XII palatine-premaxillary ligament 

Men Meniscus 

Met Metapterygoid 

Mmc Mentomeckelian cavity 

Mvp Maxillary ventromedial process 

MX Maxilla 

Mxh Maxillary head 

nm neuromast 

Nil Optic nerve 

NV Trigeminal nerve trunk 

NVII Facial (hyomandibularis) nerve 

NVllh Hyoid branch of facial nerve 

NVIIm Mandibular branch of facial nerve 

obd obliqui dorsales muscle 

obp obliquus posterior muscle 

obs obliquus superior muscle 

obv obliqui ventrales muscle 

oi obliquus inferior muscle 

Op Operculum 

Pal Palatine 

Pb Pharyngobranchial 

pee pharyngoclavicularis externus muscle 

pci pharyngoclavicularis internus muscle 

Ph Posterohyal 

phy protractor hyoideus muscle 

Pmx Premaxilla 

Po Preoperculum 

Pro Prootic 

Ps Parasphenoid 

Pte Pterotic 

Ptt Posttemporal 

Q Quadrate 

Re Rostral cartilage 

rd retractor dorsalis muscle 

Ra Retroarticular 

Rbv Buccal branch of trigeminal nerve 

re rectus communis muscle 

rd retractor dorsalis muscle 

re rectus externus muscle 

rei rectus inferior muscle 

ri rectus internus muscle 

RmV Mandibular branch of trigeminal nerve 

RmxV Maxillary branch of trigeminal nerve 


6 G. J. HOWES 

rs rectus superior muscle 

rv recti ventrales muscle 

Scl Supracleithrum 

sh sternohyoideus muscle 

shl lateral segment of sternohyoideus 

So Suboperculum 

Tp Toothplate 

tv transversus muscle 

tvd transversi dorsalis muscle 

tA 1 a, 1 1 , t2 insertion tendons of adductor mandibulae A 1 muscles 

tA2 insertion tendon of adductor mandibulae A2 muscle 

Vo Vomer 


Cranial ligaments 
Ligaments of the upper jaw and pterygoid bones 

In the following account the terminology and numbering system for ligaments follows that of 
Stiassny(1986). 

Stiassny (1986) recognised two synapomorphic arthrological characters uniting the 
acanthomorph lineages 'Paracanthopterygii' and Acanthopterygii, namely: 

the absence of a median palato-maxillary ligament (ligament IV) and 

the subdivision of the palato-vomerine ligament (ligament VI). 

I would confirm Stiassny's findings that a median palato-maxillary ligament (IV) is absent in all 
paracanthopterygian taxa examined. 

In macrouroids there is a single, undivided palato-vomerine ligament, which, from its points of 
attachment to the centre of the palatine and the head of the vomer, corresponds with Stiassny's 
ligament V (the posterior palato-vomerine ligament). The ligament runs parallel to the medial 
face of the palatine and varies in size from a long slender strap to a broad band. In the latter case 
the palatine is deep and is closely applied to the ethmo-vomerine bloc (e.g. Coryphaenoides, 
Hymenocephalus) . 

In Gadoidei the palato-vomerine ligament is also single. The presence of a single rather than a 
double ligamentous connection in gadoids and macrouroids may indicate that there has either 
been a derived loss of the anterior palato-vomerine ligament (ligament VI) or that it represents 
the plesiomorphic condition found in non-acanthomorph fishes. A broad investigation of the 
condition among paracanthopterygians is necessary to support one or other of these hypotheses. 

The maxillo-rostroid ligament (ligament VII) is well-developed in all macrouroids. As in other 
acanthomorphs it runs from the medial portion of the folded maxillary head to the dorsolateral 
face of the rostral-cartilage. In all macrouroids ligament VII appears to be continuous across the 
dorsal surface of the cartilage. In gadoids, a similar situation obtains in Bathygadus and the 
Moridae where the ligament lies in a groove in the cartilage. In many other gadoids, however, 
ligament VII is broader and inserts on the lateral face of the rostral cartilage (Figs 13 & 17). 

In some macrouroids, ligament VII runs parallel to the palato-maxillary ligament (XII), e.g. 
Coryphaenoides (Fig. 1) whereas in others it runs at ca. 45 to that ligament (Fig. 3). Ligament VII 
passes medial to the tips of the premaxillary ascending processes and is not attached to them. 
According to Stiassny (1986) in acanthomorphs ligament VII inserts on the premaxillary ascend- 
ing processes. I have not found this attachment in any gadiform and the condition she reports is 
probably a derived one for acanthopterygians. Gosline (1981) has commented on the functional 
significance of ligament VII (Gosline's ligament re) believing it to be the primary cause of upper jaw 
protrusion in at least some acanthomorphs (see p. 50). 

Casinos (1978) although identifying ligament VII in macrouroids incorrectly states that it is 
absent in the Gadidae. In fact the ligament is present in all gadoid taxa (see comments on p. 5 1 
concerning function). 

An anterior maxillo- premaxillary ligament (ligament IX of Stassny, 1986; ligament 'am' of 
Gosline, 1981) is present in all macrouroids and other gadiforms examined. In macrouroids 


MACROUROID FISHES 7 

however, the ligaments of either side meet ventroposteriorly to the rostral-cartilage forming an 
X-shaped ligament connecting the maxillary heads (their menisci) and the premaxillary ascending 
processes (Figs 28B & C). 

In gadoids ligament IX is variously developed and attached. In Bathygadus and Trachyrincus, 
there is a complex attachment of the ligament to the maxillary head via a cylindrical chondroid or 
fibrous element whose posterior tip joins a thin ligament stretching caudally, which becomes 
incorporated with the connective tissue stretching between the maxilla and premaxilla (Figs 29 A & 
B). In melanonids and merlucciids, ligament IX attaches directly to the medial process of the 
maxillary head, although it may be associated with a thick wedge of fibrous connective tissue (Fig. 
29C). In advanced gadoids, there is sometimes no discrete ligament but only tough connective 
tissue (e.g. Euclichthys) although in the majority there is a short ligament and a thin meniscus 
between the medial maxillary process and the premaxillary ascending process (Fig. 29D); see 
further comments on p. 39. 

An anterior palato-maxillary ligament (ligament X) is present in all macrouroids and gadoids 
examined. It generally connects the base of the palatine prong with the inner central portion of the 
maxillary head. However, in the macrouroids Coryphaenoides and Hymenocephalus, the ligament 
attaches to the medial aspect of the maxillary head then passes forward to attach to the anterior 
process of the premaxilla. 

An ethmo-maxillary ligament (ligament XI) is well-developed in all macrouroids and passes 
beneath the palato-premaxillary ligament (XII). Its attachments are to the lateral prong of the 
mesethmoid and the anterolateral face of the maxilla. 

In two macrourid genera, Cetonurus (Fig. 4) and Echinomacrurus, a ligament extends trans- 
versly from the ethmoid to the palatine. In this respect, the situation corresponds with that in the 
percomorph Morone illustrated by Stiassny (1986, fig. 10). According to Stiassny the additional 
ligament is a branch of a bifurcated ligament XI. Such may also be the case in the two macrourid 
taxa. It is noted that in both these genera the dorsal palatine process is higher than in others and 
that a lateral ethmo-palatine ligament is absent. Thus the 'additional' ligament may serve to brace 
the palatine against too great a lateral movement. 

In all Macrourinae there is a short ligament running from the head of the maxilla to the inner 
face of the extended nasal bone (Fig. 2). The ligament branches from the base of ligament XI; it is 
absent in Bathygadus, Gadomus, Trachyrincus and all other gadiform fishes. A maxillary-nasal 
ligament is apparently present in some acanthopterygians (Cichlidae, P. H. Greenwood, pers. 
comm.). I have not found the ligament in other paracanthopterygians examined, nor in berycoids 
or polymixiids. 

A palato-premaxillary ligament (ligament XII of Stiassny, 1986) is present in all macrouroids 
and runs from the base of the palatine prong to the contralateral premaxillary ascending process. 
Often, the ligament attaches to the antero-dorsal surface of the rostral cartilage prior to its 
insertion on the premaxillary process. Gosline (1963, fig. 5 A) shows a similar situation in the 
percopsiform Aphredoderus where ligament XII as well as attaching to the rostral cartilage is 
united with its antimere in the midline. In Percopsis, however, the ligament of each side attaches 
to its respective premaxillary ascending process, there being no contralateral attachment. The 
percopsiform situation may represent the plesiomorphic condition of ligament XII. 

A lateral ethmoid-palatine ligament is present in all macrouroids examined. This ligament, 
commonly present in nearly all teleosts, connects the posterior face of the lateral ethmoid wing with 
the dorsomedial surface of the palatine. In macrouroids, there are often two ligaments, the medial 
occupying the usual position, while the lateral ligament connects the outer margin of the lateral 
ethmoid to the lateral surface of the palatine. In the macrourines Nezumia and Ventrifossa the 
medial ligament extends posteriorly to the entopterygoid. Cynomacrurus and Odontomacrurus are 
exceptional among macrouroids in lacking a lateral ethmoid-palatine ligament. 

In the gadoid families Melanonidae, Merlucciidae and Steindachneriidae there is a single, 
stout lateral ligament connecting the lateral ethmoid with the palatine, which in Gadomus 
(Bathygadidae) extends medially to attach to the entopterygoid. 

In the Euclichthyidae there is a unique form of ligamentous connection between the lateral 
ethmoid and suspensorial elements. The lateral ethmoid ligament fans out to attach along the 


8 G. J. HOWES 

dorsolateral surface of the palatine; it continues forward as a broad band along the dorsolateral 
border of the entopterygoid, enters to adductor arcus palatini muscle, curves ventrolaterally, leaves 
the muscle and attaches to the antero-medial face of the hyomandibula (Fig. 15). 

In the Moridae there are both separate lateral and medial lateral ethmoid-palatine ligaments, 
and a lateral ethmoid-entopterygoid ligament. In the Gadidae, however, there are no definite 
ligamentous connections between the posterior face of the lateral ethmoid and the palatine. In 
most gadids, the palatine's only ligamentous connection with the neurocranium is with the vomer 
(ligament V; see above). The lateral ethmoid wing of gadids is often reduced and the palatine 
articulates not with the wing but with the anterior part of the lateral ethmoid where it contacts the 
ethmovomerine bloc. In the Muraenolepididae, for example, the palatine bears a high dorsal 
process which contacts the dorsomedial face of the (considerably reduced) lateral ethmoid. The 
palatine process is tightly bound by connective tissue to the lateral ethmoid but is not connected to 
it by a discrete ligament. 

Among more 'advanced' gadoids there is a noticeable shift in the articulation of the palatine 
toward a more anteromedial position. Among macrouroids and plesiomorphic gadoids 
(Bathygadidae; Melanonidae), the palatine articulates with the ventral surface of the lateral 
ethmoid wing to which it is also ligamentously attached. In other gadoid taxa, however, the 
palatine articulates with the anterior, ethmoidal part of the lateral ethmoid and there is a correlated 
loss of ligamentous connection between the bones. In acanthopterygians, the lateral ethmoid- 
palatine connection may be via one or more ligaments (see for example, Stiassny, 1981: 74; 
Greenwood, 1985: 158). The widespread occurrence of discrete ligamentous connections between 
the lateral ethmoid and palatine in teleosts indicates that their absence, often coupled with that of 
an intimate articulation between the two bones (Howes, 1987) represents a derived condition. 

Ligaments of the lower jaw and opercular bones 

There is a single, strong mandibular-interopercular ligament present in all macrouroids. The 
ligament is variable in length and width, from long and strap-like to short and triangular. The 
mandibular attachment of the ligament is the retroarticular, which is usually dorso-ventrally 
elongate. Okamura (19706) has drawn attention to the varying types of retroarticular among 
macrouroids. 

Casinos (1978) refers to a 'circumbuccal' ligament in macrouroids and gadids which he describes 
as a '. . . tendon that contours all the mouth'. Casinos postulates that this ligament plays an 
important role in protrusion of the upper jaw (see p. 52). The 'circumbuccal' ligament of Casinos is 
present in some form or other in all gadiform fishes examined. It does not surround the perimeter of 
the jaw as is implied by Casinos, but is attached anteriorly to each dentary. I thus refer to it as the 
'labial ligament'. The ligament varies in degree of thickness and complexity of anterior attachment, 
among gadoids being least in the Gadidae and most in the Bathygadidae, Moridae, Melanonidae 
and Merlucciidae. In macrouroids the ligament is also well-developed, but less so than in the four 
gadoid families. 

In Bathygadus (Fig. 9) where it is most highly developed, the labial ligament is a thick rope-like 
element having a bifurcate attachment on the anterior aspect of the dentary. At the rictus of the 
jaws, the ligament curves around to attach to the premaxilla, at the point of curvature sending off a 
posterior branch which anchors to the maxillary rim. 

A separate element, with the same gross consistency as the main ligament, forms a stump on the 
posteromedial surface of the maxilla, rising above the border of the bone. Rosen & Patterson 
(1969: 425) refer to this non-osseous structure in Melanonus (Melanonidae) as resembling a 
supramaxilla. I therefore refer to it as the 'supramaxillary ligament'. 

Histological sections of the labial and supramaxillary ligament, stained specifically for elastin, 
reveal the 'ligaments' to consist of a collagenous core surrounded by an elastin coat. This tissue is 
ligament-like in the nature of its attachments (it is free from the dentary, although closely adhering 
to it by a sheet of connective tissue, which is highly innervated by subranches of the ramus 
mandibularis facialis (VII) nerve. 

The distribution of the labial ligament among euteleosts is yet to be ascertained but is possibly a 


MACROUROID FISHES 

lee 


lo 


ao 


Fig. 1 


Imq 


Coryphaenoides mexicanus; cranial muscles and ligaments. Above, in lateral view. Below, medial 
view of the lower jaw adductor musculature and ligamentous connections. 


eurypterygian character (Stiassny, pers. comm.). However, its complex posterior ramification in 
the gadoids listed above appears to be a derived specialization, whose functional significance is 
commented upon elsewhere (p. 52). 

The ligament which in euteleosts connects the posterior tip of the interoperculum to the anterior 
border of the suboperculum is, in Bathygadus and Gadomus reduced and supplemented by another 
ligament stretching from the dorsal midpoint of the interoperculum to the preoperculum and 
hyomandibula. In most gadoids the interoperculum and suboperculum are connected by thin 
connective tissue, the dorsally directed ligament spanning the two bones and attaching to the 
preoperculum and hyomandibula. In the Merlucciidae, the dorsal ligament is a broad band attach- 
ing the interoperculum to the preoperculum. The Trachyrincidae have a unique condition whereby 
the interoperculum is connected by dorsally directed ligaments to the preoperculum and opercu- 


10 


G. J. HOWES 


lee 


lep 


Imn 


Fig. 2 Coelorinchus caribbaeus; cranial muscles and ligaments. In lateral (above) and dorsal (below) 

views. 


lum (see Howes, 1988, figs 2 & 3). The ligamentous connection between the interoperculum and 
hyomandibula/preoperculum is considered to be a derived condition for gadoid fishes; its taxo- 
nomic and phylogenetic implications are discussed more fully in Howes (1988). The functional 
aspects of this linkage are discussed below, p. 54. 


Cranial muscles 
The adductor mandibulae and muscles of the suspensorium in macrouroids 

The muscles of the jaws and suspensorium in macrourids have been described for some taxa by 
Dietz, 1921, McLellan , 1977 and Casinos, 1978; 1981. Dietz gave a brief description of the 
muscles in Coleorinchus coelorinchus; McLellan referred to, and illustrated the adductor muscles of 
Bathygadus and Coelorinchus, and Casinos those of Coryphaenoides and Trachyrincus. The two 


MACROUROID FISHES 


11 


Fig. 3 Ventrifossa occidentalism cranial muscles and ligaments in lateral view. 


lo ao Hyop 


So 


Imm 


lo 


Fig. 4 Cetonurus globiceps; cranial muscles and ligaments in lateral view. 


12 G.J.HOWES 

latter authors were concerned with describing the musculature in a functional context, although 
Casinos (1978) made some observations regarding the homology and evolution of certain 
adductor muscles. The taxonomic range of these authors' works is limited and the applicability of 
their functional conclusions to macrouroids in general requires a reappraisal in the light of 
morphological variations of which they were unaware. 

NB. In the following descriptions the dorsal muscle of the adductor element (levator maxillaris 
superioris of authors) is referred to as A 1 P; its homology is discussed later, p. 34. 

MACROURINAE 

Type 1 morphology: two subgroups are recognised, (a) Coryphaenoides , Abyssicola, Nezumia, 
Coelorinchus, Lionurus (synonymised with Coryphaenoides by Iwamoto & Stein, 1973); 
Nematonurus, Chalinura; (b) Macrourus, Trachonurus. 

The overall morphology of the adductor musculature is similar in the two subgroups, the only 
difference being the presence of an additional adductor element, Aly, in subgroup (a). 

In all the taxa included in the Type I group the mouth is inferior or subinferior and the jaws 
relatively short; the premaxillary ascending process is at least 80% the length of the dentigerous 
ramus; the maxilla is a deep, stout bone with a markedly convex dorsal border. 

The outer adductor muscle is thin but relatively deep and divisible into upper and lower parts 
which are either entirely separated (e.g. Coryphaenoides, Fig. 1), or partially so (e.g. Coelorinchus, 
Fig. 2). The lower part (Ala) originates from the preopercular limb, and in Coryphaenoides from a 
prominent lateral flange of that bone (Fig. 1). The muscle inserts tendinously along the lower part 
of the maxillo-mandibular ligament. The upper part of the adductor (A1P) originates from the 
preopercular limb and inserts via a stout tendon on to a ventromedial process of the maxilla; it is 
not joined to the maxillo-mandibular ligament. 

Running dorsomedially to Al P is a long spindle-shaped muscle here designated Aly (Fig. 3); see 
below. The fibrous part of the muscle originates from a long tendon which in turn stems from the 
fascia of Al p. Insertion is via a cord-like tendon on the same medial process of the maxilla as Al p. 
In Lionurus, there is a marked difference in the relative proportion of the fibrous part to the 
posterior tendinous part of the muscle between small and large-sized specimens. In a specimen of 
150 mm TL, the muscle is 50% tendinous and 50% musculose, whereas in a specimen of 225 mm 
TL, 75% of the muscle is fibrous. In Abyssicola macrochir, muscle Aly is a larger and deeper 
element than in any other taxon examined. Also, unlike other taxa of this group the muscle 
originates from a broad tendinous sheet stemming from the rim of the hyomandibula. 

Muscle A2 is a deep, broad element whose medial fibres originate from the frontal, and those 
more lateral in position from the prootic and hyomandibula. A2 has a complex insertion in the 
lower jaw. Its posteromedial fibres insert into an aponeurosis which bifurcates into a vertical and a 
horizontal tendon. The vertical tendon inserts onto the coronomeckelian bone and continues to 
the dorsomedial surface of the narrow retroarticular, while the horizontal tendon runs forward 
into the mandibular cavity. The majority of fibres of A2 insert on the horizontal tendon, from 
which also originate those of the mandibularis section of the adductor Aco (Fig. 1). Muscle AGO lies 
mostly outside the mentomeckelian cavity, but with a small bundle of lateral fibres running 
forward into it. 

The levator arcus palatini (Figs 1 & 2) is a long, pyramidical muscle running between the 
sphenotic and the lateral face of the hyomandibula; its outermost fibres insert on the edge of the 
preopercular limb. 

The dilatator operculi (Figs 1 & 2) originates from the lateral hyomandibular fossa and inserts on 
the rim of the opercular facet. The adductor and levator opercularis muscles extend from the lateral 
border of the pterotic, the adductor inserting on the opercular process of the hyomandibula and the 
levator on the anteromedial face of the operculum (Figs 1 & 2). 

The adductor arcus palatini occupies the floor of the orbit, its anterior fibres inserting on the 
broad concave surface of the palatine (Figs 1 & 2). Posteriorly, the muscle runs between the 
parasphenoid and the lateral faces of the entopterygoid and metapterygoid and the medial face of 
the hyomandibula. 


MACROUROID FISHES 


13 


Type II morphology. Macrourinae (part): Ventrifossa, Cetonurus, Echinomacrurus , Malaco- 
cephalus, Hymenocephalus, Odontomacrurus, Sphagemacrurus , Cynomacrurus , Mataeocephalus. 

Taxa of this group have a terminal or subterminal mouth, with the exception of Echinomacrurus 
in which it is inferior. The ratio of premaxillary ascending process to dentigerous ramus length 
varies from 30-33% in Odontomacrurus and Chalinura to 50% in Ventrifossa. An opposite extreme 
is Mataeocephalus where the premaxillary ramus is 50% of the length of the ascending process (cf. 
Macrouroidinae, p. 1 5); in Echinomacrurus and Cetonurus, the ramus and ascending process are of 
almost equal length. 

The characteristic myological feature of this morphotype is that Al is a single, or incompletely 
divided, deep element. Ventrifossa occidentalis is taken to illustrate the morphotype, representative 
of the majority of taxa (Fig. 3). Echinomacrurus and Cetonurus which differ somewhat in detail 
from Ventrifossa are considered below. 


Ptt 


Fig. 5 Cynomacrurus piriei; levator arcus palatini and opercular muscles in lateral view. 


Muscle A 1 originates posterodorsally from the upper part of the preopercular limb and ventrally 
from the preoperculum and quadrate (Fig. 3). The dorsal border of the muscle is almost horizontal, 
there being a slight concavity and tendinous area below the orbit. Dorsal fibres of the muscle insert 
on the inner aspect of the maxilla, while the remainder of the muscle, separated from the upper part 
by an internal aponeurosis, inserts into the anterior third of the maxillo-mandibular ligament. 
Along the centre of the muscle is an aponeurosis which is marked laterally by a change in muscle 
fibre direction from almost horizontal (dorsally) to oblique (ventrally). The position of the 
aponeurosis is marked in taxa belonging to morphotype I by a complete or partial division of the 
muscle. In this respect Cetonurus resembles the latter taxa (Fig. 4). 

Muscle A2 is a deep element originating from a cavity formed between the prootic and frontal, 
with fibres stemming from both bones. In Cynomacrurus and Odontomacrurus the anterior muscle 
fibres are vertically aligned or posteroventrally angled, in contrast to the more usual anteroventral 
angle present in Ventrifossa (Fig. 3). The insertion of muscle A2 in the lower jaw is via a cord-like 
tendon carrying outer fibres to the coronomeckelian bone and into a broad aponeurosis from 
which originates muscle AGO. The muscle is short, barely extending halfway along the mandible; 
only its dorsal fibres enter the mentomeckelian cavity. 


14 


G. J. HOWES 


lap 


do 


A 2 


Fig. 6 Macrouroidinae; above, Squalogadus modificatus, preoperculum and associated muscles; below, 
Macrour aides inflaticeps, anterior of adductor musculature. 


The levator arcus palatini is a large muscle in all Type II genera, extending from the sphenotic and 
pterotic to the lateral face of the hyomandibula, its anteroventral portion covering the postero- 
dorsal margin of Al . In Cynomacrurus the levator is angled forward to a greater degree than in the 
other included taxa (Fig. 5). 

The dilatator, adductor and levator operculares muscles are all well-developed. The dilatator 
operculi originates from a lateral hyomandibular fossa and the adductor from the ventral surface of 
the pterotic. Some fibres of the adductor operculi insert with those of the dilatator on the rim of the 
opercular condyle, but the majority insert on the opercular process of the hyomandibula. In 
Cetonurus the separate insertions of the muscle are further marked by the complete division of its 
body. The levator operculi is a long, deep muscle originating from the ventral surface of the pterotic 
and inserting along the anteromedial border of the operculum. In Cynomacrurus the adductor 
operculi inserts entirely on the opercular process of the hyomandibula and the levator operculi is 
divided. The anterior segment of the levator shares a common origin with the adductor operculi, but 
the posterior segment originates from the posttemporal (Fig. 5); both segments insert together on 
the anteromedial face of the operculum. With respect to its posttemporal origin, the levator of 
Cynomacrurus is similar to that of the gadoid Lota (see p. 32 and Howes, 1988). 


MACROUROID FISHES 


15 


The adductor arcus palatini extends the length of the parasphenoid and anteriorly inserts on 
the palatine; posteriorly it inserts on the ento- and metapterygoids and the medial face of the 
hyomandibula. In Cetonurus the muscle extends only halfway along the length of the parasphenoid 
(Fig. 4). 


MACROUROIDINAE 

Two monotypic genera are included in this subfamily, Macrouroides and Squalogadus. Of the 
former, only a single, poorly preserved specimen of Macrouroides inflaticeps was available for 
examination. The specimen has a damaged and partially disarticulated skull and it has been 
impossible to ascertain precisely the configuration and insertions of the adductor muscles. 
Likewise, only a single specimen of Squalogadus modificatus is available for examination and only a 
partial dissection of the posterior region of the cheek musculature has been possible (Fig. 6). 

Muscle Al is a single element originating from the preopercular limb and inserting on to the 
upper part of the maxilla via the maxillo-mandibular ligament. 

Muscle A2 is a thick, crescentic muscle stemming from the frontal and prootic; its insertions in 
the lower jaw and the extent of muscle Aco have not been ascertained in either taxon. 


Table 1 Grouping of Macrourinae based on jaw and ventral gill-arch muscle morphotypes (see 
text, p. 58), compared with Okamura's (19706) groupings 


Jaw muscles 


Gill-arch muscles 


Okamura's 
groups 


Type \a 


Abyssicola 
Coryphaenoides 
Nezumia 


(a) Abyssicola 
Coryphaenoides 
Coelorinchus 


Abyssicola 
Coelorinchus 
Macrourus 


Lionurus 


Lionurus 


Nematonurus 


Nematonurus 


Chalinura 


Chalinura 


Macrourus 


Type \b 


Macrourus 
Trachonurus 


Trachonurus 
Malacocephalus 
Mataeocephalus 
Cetonurus 


Echinomacrurus 


Type II 


Ventrifossa 
Malacocephalus 
Hymenocephalus 
Odon tomacrurus 


(b) Ventrifossa 
Hymenocephalus 
Nezumia 
Odontomacrurus* 


Ventrifossa 
Malacocephalus 

Odontomacrurus 


Cynomacrurus 
Mataeocephalus 
Macrosmia 


Cynomacrurus* 


Cynomacrurus 


Cetonurus 


Cetonurus 


Echinomacrurus 


Echinomacrurus 


Sphagemacrurus 


Sphagemacrurus 


NB. * these two genera lack a palatine-lateral ethmoid ligament. According to Iwamato & Stein (1973), 

Lionurus, Nematonurus and Chalinura should be treated as subgenera of Coryphaenoides. 

The following macrourine genera have not been examined, Astenomacrurus , Cetonurichthys , Haplomac- 
rourus, Lepidorynchus, Paracetonurus, Parakumba, Pseudocetonurus, Pseudonezumia. 


16 


lo 


So 


Imm 
Fig. 7 Trachyrincidae: Trachyrincus trachyrincus; cranial muscles in lateral view. 


The levator arcus palatini and operculi muscles are missing from the specimen of Macrouroides, 
but in the Squalogadus specimen there is a small levator arcus palatini, lying posterolaterally to the 
adductor mandibulae complex (Fig. 6). 

The adductor arcus palatini is a thick element flooring the orbital cavity, inserting anteriorly on 
the palatine and posteriorly on the lateral surface of the ento- and metapterygoids. 

Comparisons with gadoids 

The presence of a dorsal section of the adductor mandibulae originating from the palatoquadrate 
was considered by Rosen & Patterson (1969: 361 et seq.) to be a specialisation of paracanthoptery- 
gian fishes. Fraser (1972) commented that a muscle of this type had developed in several acanthop- 
terygian groups and could not be used as a character indicating phyletic relationships. He also 
pointed out that the number of taxa examined for this character by Rosen and Patterson was too 
few to make generalisations as to its occurrence and homology. Marshall & Cohen (1973) in 
referring to the so-called levator maxillaris superior is muscle note that '. . . character has received 
little comment; its distribution and taxonomic significance are at present in need of fuller survey'. 

There has been much confusion concerning the identity of the dorsal adductor muscle in para- 
canthopterygians. Rosen (1962) and Rosen & Patterson (1969: 341 et seq.} referred to the element 
as a levator maxillaris superioris (i.e. the homologue of that muscle in the halecomorph Amia; see 
Allis, 1897). Previous authors, viz. Holmquist (191 1) and Dietz (1921) had referred respectively to 
the muscle as A4 and Alp. Later, Rosen (1973: 417) reformulated his ideas and, following Dietz, 
referred to the muscle as A 10, a view supported by Winterbottom (19740) and most subsequent 
authors. Casinos ( 1 978: 443) continued to use the term levator maxillaris superioris '. . . because of 
functional reasons'. 

Because the muscle in question lies lateral to the mandibular branch of the trigeminal (V) nerve, I 
concur with Winterbottom (19740) in recognising it as part of muscle Al. Allis (1897: 581-2) 
comments that the '. . . course and position of the inferior maxillary nerve . . . seems to lie always 
between Al and A2 . . .'; see similar remarks of Freihofer (1978: 17) and Howes (1985: 275). 


MACROUROID FISHES 


17 


In all gadiform fishes I have examined, apart from some macrouroids noted above and the 
Trachyrincidae (see below), a dorsal division of muscle Al is present. The various conditions of 
this, and other cranial muscles are as follows: 


TRACHYRINCIDAE 

(Figs 7 & 8) 

The family (formerly recognised as a macrouroid subfamily) contains two genera Trachyrincus 
and Idiolophorynchus. Species are characterised by their unique adductor muscle arrangement 
(described below), interopercular-preopercular-opercular ligamentous arrangement, nasal mor- 
phology, caudal skeleton (see Howes, 1988) and other features such as dorsal scutes (given in 
diagnosis for subfamily by Marshall, 1973). In trachyrincids the jaws are long, the length of the 
premaxillary ascending process being 50% of the ramus. 

Muscle Al is a single, narrowly triangular element which extends from the anterolateral face of 
the preoperculum to insert via a double tendon on the maxilla. The upper tendon (t2, Fig. 7) passes 
medially to insert close to the maxillary head while the lower (tl, Fig. 7) joins the maxillo- 
mandibular ligament to insert on the lateral face of a maxillary dorsal process. Casinos (1978: 443) 
is incorrect in stating that a maxillo-mandibular ligament is absent in Trachyrincus. A ventral 
tendon runs from the aponeurosis to the coronomeckelian bone; muscle AGO extends from the 
anterior part of the aponeurosis, the majority of fibres lying medial (outside) the mentomeckelian 
cavity. 

The levator arcus palatini in contrast to that of macrouroid taxa, is extended posteriorly and its 
anteroventral part is covered by the adductor mandibulae. The muscle originates dorsally from the 
sphenotic and pterotic, and medially from the hyomandibula; insertion is across the extensive 
dorsal face of the preoperculum. 

The dilatator operculi originates from a common aponeurosis with the levator arcus palatini and 
has a strong tendinous insertion on the rim of the opercular condyle. The adductor operculi is a long 
muscle originating from the ventral surface of the pterotic and inserting for much of its length on 
the opercular process of the hyomandibula. Only the posterior fibres insert on the operculum, just 
medial to the insertion of the dilatator muscle. The levator operculi is a thin, narrow element 
running postero-laterally and inserting on the medial face of the small operculum. 

The adductor arcus palatini, although extending the length of the parasphenoid is weakly 
developed anteriorly (of single fibre thickness) and does not insert on the palatine. 


A 2 v 


Po 


Q 


Ra 


Aa Cmb Cmc 

Fig. 8 Trachyrincus trachyrincus; medial view of inner adductor muscle and lower jaw. 


18 


G. J. HOWES 


do 


Rt 


'al 


Re 


lo 


De 


Imm 


Fig. 9 Bathygadidae: Bathygadus melanobranchus; cranial muscles and ligaments in lateral view. 


BATHYGADIDAE 

(Figs 9 & 10) 

Formerly recognised as a subfamily of macrouroids, Howes (1988) referred the 'bathygadine' 
genera Bathygadus and Gadomus to the Gadoidei as a clade, here recognised as a family. A 
complete taxonomic diagnosis is in preparation but it can be stated here that the family is dis- 
tinguished from other gadoid families by its lack of a caudal skeleton, derived RLA pectoral nerve 
pattern, reduced gill-filaments, reticulate scale pattern and myological synapomorphies detailed 
here. 

Bathygadus and Gadomus have a terminal mouth with a wide gape, the jaws are long and slender. 
The outer adductor muscle is a thin, shallow sheet of fibres originating from the preoperculum and 
posterior margin of the hyomandibula. In most Bathygadus species it is clearly divided into ventral 
(A 1 a) and dorsal (A 1 P) parts. However, in B.favosus, the two muscles can only be distinguished by 
their separate tendinous insertions on the maxilla. In the species where the Ala and Alp parts 
remain separated, Ala joins a broad aponeurosis with the maxillo-mandibular ligament halfway 


MACROUROID FISHES 


19 


along the length of the maxilla. Muscle Al(3 originates from the medial fascia of the levator arcus 
palatini muscle and is divided into anterior and posterior segments, the division being brought 
about by the muscle's tendinous constriction below the orbit. The anterior segment of A 1(3 inserts 
on the medial face of the maxillary process. 

In Gadomus muscle Ala has a definite insertion on the outer aspect of the anterior part of the 
maxilla, and the maxillo-mandibular ligament is longer and narrower than in Bathygadus. 

Muscle A2 is a large element originating from the hyomandibula, prootic and frontal and 
inserting on the lower jaw. Insertion is partly via a vertical tendon stretching down the medial face 
of the anguloarticular, and partly on a tendon inserting on the coronomeckelian bone and medial 
face of the anguloarticular. From these tendinous insertions stem the fibres of the mandibularis 
part of the adductor muscle (Aco). The anterior half of Aco enters a long mentomeckelian cavity. 

The levator arcus palatini is a large, deep element which inserts halfway down the preopercular 
limb and entirely covers the origin of muscle Al . The lateral posterodorsal fibres of the levator are 
inseparable from those of the dilatator operculi. 

The adductor arcus palatini extends nearly the entire length of the parasphenoid, but anteriorly it 
is feebly developed, with widely spaced, tendinous bands of fibres; posteriorly the muscle inserts on 
the lateral face of the metapterygoid and the medial face of the hyomandibula. 

As noted above, the anterior fibres of the dilatator operculi intermesh with the dorsoposterior 
fibres of the levator arcus palatini; those respective groups of fibres of both muscles originate from 
the hyomandibula, inserting together with the adductor operculi on the opercular condyle. The 
levator operculi is a well-developed element stemming from the pterotic and inserting along the 
posteromedial border of the operculum. 


ao 


lo 


A,P 


NVII 


Imm 
Fig. 10 Bathygadidae: Gadomus longifilis; cranial muscles and ligaments in lateral view. 


20 


G. J. HOWES 

MORIDAE 

(Fig. 11) 


In their overall morphology, the cranial muscles of morids are most similar to those of the 
Bathygadidae and Gadomus (Melanonidae). In Halargyreus (Fig. 1 1), Pseudophycis and Antimora, 
muscles A 1 a and A 1 P originate laterally to A2; A 1 P is tendinously constricted below the orbit, its 
anterior expansion joining the insertion tendon of A 1 a. 

The levator arcus palatini is large, its ventral tip extending to a point halfway down the pre- 
opercular limb (cf. Bathygadus). The adductor arcus palatini is divided into posterior and anterior 
parts, the latter inserting on the entopterygoid and not the palatine as in most other gadoids. 

In Lepidion (Fig. 1 0), muscles A 1 a and A 1 P are incompletely separated; both segments originate 
from a single body. The dorsal (A 1 P) and ventral (A 1 a) bundles insert on separate tendons beneath 
the orbit, which then join into a single muscle body lateral to the palatine before separating into 
their respective medial and lateral maxillary insertions. 

The levator arcus palatini covers the upper part of A2, dorsally it joins the dilatator operculi along 
an aponeurosis. The adductor arcus palatini is feebly developed in its central portion; the anterior 
fibres insert on the palatine. 


aap 


Fig. 1 1 Moridae: above, Halargyreus affinis; below, Lepidion eques; upper jaw and suspensorial muscles 

in lateral view. 


MACROUROID FISHES 

MELANONIDAE 

(Fig. 12) 


21 


In Melanonus muscles Ala and Alp are inseparable at their origins which is from the fascia of 
muscle A2. The individual elements only become apparent above the jaw articulation. A 1(3 is 
constricted into a tendon halfway along its length at the point where it is crossed transversely by a 
ligament running from the posterolateral edge of the palatine and the entopterygoid to the medial 
face of the second infraorbital. The anterior expansion of Alp inserts on the inner part of the 
maxillary head. The outer element, Ala, inserts via a separate tendon on to the outer face of the 
maxilla. 


do 


lip 


NVIlm 


Imm 


Fig. 12 Melanonidae: Melanonus zugmayeri; cranial muscles in lateral view (suboperculum removed). 


Muscle A2 is strongly developed, its anterior fibres being almost vertical. The muscle is partially 
divided by a hypertrophied ramus mandibularis interims facialis of the hyomandibularis VII nerve 
trunk (Fig. 12). 

The levator arcus palatini shares an aponeurotic origin with the dilatator operculi. The adductor 
arcuspalatiniis well-developed, flooring the orbital cavity, and inserting anteriorly on the palatine. 

The adductor and levator operculares muscles share a common origin from the ventral surface of 
the pterotic shelf. The adductor inserts both on the dorsomedial rim of the opercular condyle and 
the opercular process of the hyomandibula; the levator inserts along the dorsomedial border of the 
operculum: 

STEINDACHNERIIDAE 

(Fig. 13) 

In Steindachneria muscle Al is large, originating from the lower half of the preopercular limb. Its 
fibres are angled anterodorsally, and dorsally the muscle is divided. The posterodorsal group of 
fibres insert on an aponeurosis from which stems a sausage-shaped segment of fibres running 


22 


G. J. HOWES 


forward to meet, laterally, the maxillo-mandibular ligament. From this point, the muscle becomes 
separated from the ligament and almost immediately inserts on the medial aspect of the maxillary 
head. This part of the muscle is identified as A 1(3. The ventrolateral group of fibres inserts directly 
on the maxillo-mandibular ligament and is identified as Ala. 

Muscle A2 originates from the prootic, the sphenotic process and the upper part of the 
hyomandibula, its anterior fibres running almost vertically. 

Insertion in the lower jaw is via a strong vertical tendon to the coronomeckelian bone and a 
broad aponeurosis from which originates Aco. No fibres of A2 insert on the anguloarticular. 
Muscle Aco is lanceolate, the majority of its fibres filling the mentomeckelian cavity. 

The levator arcus palatini is moderately developed, originating from the sphenotic process and 
pterotic, and inserting in a lateral cavity of the hyomandibula. The muscle lies lateral to A2, but its 


lo ao do 


Fig. 13 Steindachneriidae: Steindachneria argentea: cranial muscles in lateral (above) and dorsolateral 
(below) views. NB. Not all the upper jaw ligaments are shown in the lower drawing. 


MACROUROID FISHES 

lap 


23 


lap 


Fig. 14 


A,. 


Euclichthyidae: Euclichthys polynemus; above, cranial muscles in lateral view; below, posterior 
associations of muscle A t p; A t a reflected. 


ventral tip does not reach as far as the origin of muscle Al. The adductor arcus palatini is well- 
developed posteriorly and anteriorly, where it inserts on the palatine but its central portion is 
reduced to a few widely spaced fibres which are well-separated from the dorsal margin of the 
pterygoid series. 

Rosen & Patterson (1 969, fig. 44a) depict the adductor musculature of Steindachneria. However, 
my observations are not completely in accord with theirs, since they show muscles Ala and Al|3 
separated for their entire lengths, and a fully developed adductor arcus palatini. 

The dilatator, adductor and levator operculares muscles are as described for Melanonus. 

EUCLICHTHYDIDAE 

(Figs 14 & 15) 

The adductor mandibulae muscle is a thick, deep element originating from the upright limb of the 
preoperculum; it comprises superficial, Ala, and medial, Al|3, elements which have complex 
associations posteriorly. 

Muscle Al p is, posteriorly, a shallow, band-like muscle, having its origins aponeurotically from, 
dorsally, the levator arcus palatini, and ventrally, the dorsomedial surface of Ala, thus partially 
dividing the latter. Anteriorly, muscle Alp becomes bulbous and transversely expanded, joining 
with Ala before separating from it to insert on the ventromedial surface of the maxillary head. 


24 


G. J. HOWES 


Muscle Ala is tendinous anteriorly and joins the maxillo-mandibular ligament together with A 1 0; 
its insertion is on the dorsolateral surface of the maxilla. 

Muscle A2 is well-developed, originating from the sphenotic and the dorsolateral surface of the 
hyomandibula. In the lower jaw, A2 joins a band-like aponeurosis, from which originates Aco; a 
strong vertical tendon runs from the aponeurosis to the coronomeckelian bone. Muscle Aco is long 
and shallow, lying within the mentomeckelian cavity for most of its length. 

The levator arcus palatini originates from the sphenotic and pterotic; it bifurcates ventrally, the 
anterior branch inserting on the hyomandibula and having an aponeurotic connection with muscle 
Alp (see above and Fig. 14); the posterior branch inserts on the preoperculum and overlaps the 
posterodorsal edge of Ala. 

The adductor arcus palatini is confined to the posterior part of the parasphenoid; it inserts on the 
lateral faces of the ento- and metapterygoids. A unique feature of this muscle is that it is divided 
by a strong ligament running from the lateral ethmoid and palatine to the medial face of the 
hyomandibula (see p. 7 and Fig. 1 5). 

The dilatator operculi is a spindle-shaped muscle extending from the pterotic to the opercular 
process. The adductor operculi runs almost laterally from the underside of a pterotic shelf to insert 
entirely on the opercular process of the hyomandibula; the levator operculi is an extensive muscle 
whose insertion extends along the entire dorsal border of the operculum. 


NSDh 


Met 
Fig. 15 Euclichthyidae: Euclichthys polynemus; adductor arcus palatini muscle and associated elements. 


MERLUCCIIDAE 

(Figs 16-18) 

The following descriptions are based on three genera, Merluccius, Macruronus and Lyconus; 
Lyconodes has not been examined. 

In Merluccius (Fig. 16) muscle Ala is a thin, shallow element, stretching from a tendinous origin 
on the preoperculum across the face of muscle A2 to insert via a cord-like tendon halfway along the 
maxilla where it joins the maxillo-mandibular ligament. Muscle Alp is a deep element having its 
origin from the meta- and entopterygoid and the palatine. It passes medial to the ramus mandibu- 
laris of the trigeminal nerve. The part of the muscle originating from the palatine is thick and 
bolster-like (Fig. 16). Insertion of Al p is across a wide area of the medial face of the maxilla. 


MACROUROID FISHES 

lap aa P 


25 


lo 


Fig. 16 Merlucciidae: Merluccius merluccius; cranial muscles in lateral view; above, entire; centre, 
palatine portion of muscle A t p (superficial); below, after removal of superficial muscle. 


The levator arcus palatini is well-developed and lies lateral to the adductor mandibulae A2, but 
does not cover the origin of A 1 . The adductor arcus palatini floors the orbital cavity and inserts on 
the palatine. An unusual feature is the presence of a separate, small muscle running from the medial 
face of the palatine to the lateral face of the ethmoid bloc (Fig. 16). The muscle stems from the 
fascia of the adductor arcus palatini; in some specimens there are no muscle fibres, but only a 
narrow sheet of connective tissue. 

The opercular muscles are well-differentiated, although the dilatator operculi shares an 
aponeurotic origin with the levator arcus palatini. 

In Macruronus (Fig. 1 7), muscle A 1 a is a thick, bulky element almost covering the lateral face of 
A2. In the pinnate arrangement of its fibres, the muscle differs from that in the taxa so far 
considered. Insertion is on the dorsal maxillary process via a thick tendon. Muscle A10 originates 
from the outer rim of the quadrate, the entopterygoid and a lateral cavity of the palatine; insertion 
is one the medial face of the maxillary head. The position of the ramus mandibularis of the 
trigeminal nerve lies posterior to the origin of Al (3. 

Muscle A2 originates from the dorsomedial face and anterior rim of the hyomandibula and from 
the prootic. Insertion is via a lateral tendon to the coronomeckelian bone and a medial aponeurosis 
from which stems Aco; the latter lies entirely within the mentomeckelian cavity. 

The levator arcus palatini is small, its ventral portion covered by Ala; insertion is into a small 
lateral hyomandibular cavity. The adductor arcus palatini is divided into anterior and posterior 


26 


G. J. HOWES 


lap 


aap 


Fig. 17 Merlucciidae: Macruronus magellanicus; cranial muscles in lateral view. 


portions, the former inserting into the medial cavity of the palatine. The adductor operculi inserts 
entirely on the opercular process of the hyomandibula. 

In Lyconus (Fig. 18), muscle Ala is a narrowly triangular element whose area of origin extends 
from the central to the upper part of the preopercular limb. Fibre direction in the anteroventral 
section of the muscle is at almost 45 to that of the dorsal part. Insertion is via a long tendon on to 
the dorsal aspect of the maxilla. Muscle Alp resembles that of Merluccius in that it originates 
medially to the ramus mandibularis of the trigeminal nerve. The muscle's origins and insertions are 
complex; a posterior segment originates from the metapterygoid, a narrow, medial segment from 
the rim of the entopterygoid, and a long anterior segment from the concave lateral face of the 
palatine. The posterior segment runs into a long tendon, separate from that of the single tendon 
shared by the medial and anterior segments. The two tendons run forward to share a common 
insertion on the dorso-medial part of the maxillary head. 

Muscle A2 is large, originating from the sphenotic and pterotic, its anterior fibres running 
almost vertically into the lower jaw. 

The levator arcus palatini is a small unipinnate muscle originating from the sphenotic and 
pterotic; its insertion on the preopercular limb is above the origin of Ala. The adductor arcus 
palatini is well-developed, flooring the orbital cavity and, anteriorly, inserting on the palatine. 

As only the type specimen of Lyconus brachycolus was available it has not been possible to make 
a sufficiently extensive dissection to ascertain the morphology of the other cranial muscles. 


GADIDAE 

(Fig. 18) 

The following descriptions are based on three genera, Gadus and Merlangius (Fig. 18). These taxa 
differ from all those previously described in that muscle Ala is merely a thin, flat sheet of fibres 
stemming from the lateral body of A2 (as reported for Microgadus by Rosen, 1 962). The separation 
of Ala from A2 is marked by the course of the ramus mandibularis of the trigeminal nerve, which 


MACROUROID FISHES 


27 


passes medial to the segment; the separated fibres insert on the dorsal aspect of the maxilla via the 
maxillo-mandibular ligament. 

Muscle A 1(3 is a noticeably stout muscle and also differs from the previously described con- 
ditions in that it originates tendinously from the lateral face of the hyomandibula, passing forward 
between muscles A2 and A3 and lateral to the ramus mandibularis nerve (cf. medial in Merluccius). 
The muscle is deep and parallel fibred, running against, but not attaching to, the palatine; insertion 
is on the ventral medial edge of the maxillary head. 

Holmquist (191 1: 12-17) has adequately described and illustrated the origins and insertions of 
the deeper adductor and the suspensorial and opercular muscles in Gadus; I find little variation 
from this condition in other gadid genera examined. It should be noted here, however, that the 
levator arcus palatini inserts on a lateral shelf or slope of the hyomandibula. Although in Gadus, the 
levator is, for the most part, covered laterally by muscle A2, in Merlangius, A2 originates from 
below the hyomandibular shelf, thus leaving the levator exposed laterally and its outermost fibres 
lying in the same lateral plane as those of A2. The adductor muscles A2 and A3 are further 
discussed below (p. 41). 


Fig. 18 Merlucciidae: above, Lyconus brachycolus, jaw and suspensorial muscles in dorsolateral view. 
Gadidae; below, Merlangius merlangus, jaw and suspensorial muscles in lateral view. 


28 


G. J. HOWES 

RANICEPITIDAE 

(Fig. 20) 


Howes (1987) recognised Raniceps as belonging to a distinct family on the basis of its sharing with 
certain phycids and the Muraenolepididae a tendinous attachment of the rectus communis muscle 
and the derived arrangement of the adductor musculature now described. 

Muscle Ala is a small, spindle-shaped element, originating from the anterolateral face of A2. It 
runs alongside A 1 p, to which it is closely applied, and inserts via a long tendon on the dorsolateral 
face of the maxilla. 

Muscle A 1 p is a broad, band-like element having its origin tendinously from the hyomandibula 
and passing between muscles A2 and A3; anteriorly, the muscle becomes bulbous and inserts on the 
ventral surface of the maxillary head. 


tA 2 


Fig. 19 Gadidae: Molva molva; jaw and suspensorial muscles in lateral view; above, entire; below, with 
outer adductor element removed to expose origin of AiP. Mandibularis branch of trigeminal nerve in 
solid black. 


MACROUROID FISHES 


29 


aap 


ao 


do 


lo 


epx 


Imm 


Fig. 20 Ranicepitidae: Raniceps raninus; cranial muscles in dorsolateral view. 


Muscles A2 and A3 are thick, bulbous elements, the former originating from the preoperculum 
and hyomandibula, and the latter from the pterosphenoid and prootic. Both muscles join a 
common aponeurosis in the lower jaw from which Aco originates. 

The levator arcus palatini is also a thick, bulbous element, stemming from the sphenotic and 
pterotic to insert on the hyomandibula and preoperculum. The lateral fibres of the levator meet 
those of adductor A2 along a near vertical raphe. The adductor arcus palatini is well-developed and 
floors the orbital cavity. The anterior fibres do not, however, insert on the palatine but remain 
within the confines of the entopterygoid. 

The dilatator operculi is a narrow, ribbon-like muscle sharing a common origin with the pos- 
terior fibres of the levator arcus palatini; it inserts tendiously on the rim of the opercular facet. The 
adductor and levator operculares muscles share a common origin from beneath the pterotic and are 
separable only because of their insertions. The adductor inserts entirely on the opercular process of 
the hyomandibula; the levator along the dorso-medial surface of the operculum. 


PHYCIDAE 

(Fig. 21) 

Urophycis is taken as the taxon representing this family but in at least one myological character 
both it and Phycis differ from other genera regarded as belonging to the family (see below). 

In Urophycis (Fig. 2 1 ), fibres of muscle A 1 share a common origin from the preopercular margin 
with those of A2. Al separates from the body of A2 above the jaw articulation, its fibre direction 
varying from horizontal to 45, and inserts via a long tendon on the anterodorsal process of the 
maxilla. Ventromedially, the insertion of Ala joins the maxillo-mandibular ligament. 

Muscle Alp is a thick, cylindrical element originating, medially to A2, from the anterior rim of 
the hyomandibula. The muscle passes laterally to the ramus mandibularis of the trigeminal nerve, 
becoming slightly indented on its medial face below the orbit, and inserting musculously on to the 
ventral surface of the maxillary head. 


30 


G. J. HOWES 


Muscles A2 and A3 join a common aponeurosis medial to the anguloarticular (only a few fibres 
insert on the dorsal rim of the bone); the aponeurosis divides into medial and lateral tendons, the 
lateral one inserting on the coronomeckelian bone, while the medial branch forms the site of origin 
for muscle AGO. This muscle fills the mentomeckelian cavity with only a thin layer of fibres passing 
outside the cavity along the medial face of the dentary. 

The levator arcus palatini is extensive and lies between A2 and A3; its dorsoposterior part can 
only be distinguished as a dilatator operculi by the insertion of those fibres on the rim of the 
opercular facet. The adductor operculi runs from the pterotic to the opercular process of the 
hyomandibula; the levator operculi inserts on the medial rim of the operculum. Urophycis is 
unusual in that a segment of epaxial muscle runs anteroventrally from the supracleithrum to insert 
on the medial face of the operculum. The posteroventral border of the muscle meets a part of the 


epx 


lap 


hyad 


hyad 


Fig. 21 Phycidae: above Urophycis regia; cranial muscles in lateral view. Lotidae: below, Lota lota; 
cranial muscles in lateral view; extent of levator arcus palatini and pathway of ramus mandibularis are 
indicated by dashed lines. 


MACROUROID FISHES 

a 


Fig. 22 Bregmacerotidae (above): Bregtnaceros atlanticus. Muraenolepididae (below): Muraenolepis 
microps. Cranial muscle in lateral view (posterior extent of A t p indicated by dashed lines). 


hyohyoidei adductores which extends from the last branchiostegal ray almost to the dorsal margin 
of the operculum. The epaxialis segment is less well-developed in Phycis and such an arrangement 
is absent in other phycids (see Howes, 1988). 

LOTIDAE 

(Figs 19; 21) 

In Lota (Fig. 21) muscle Ala is only differentiable from A2 anteriorly by the separation of a bundle 
of lateral fibres which insert, via a long tendon, on the dorsal process of the maxillary bone. Muscle 
A 1 P originates from the anterior rim of the hyomandibula, medial to A2. A ventral tendon of A 1 (3 
meets A 1 a at the anterior border of A2; the fibres of muscles A 1 a and A 1 P are indistinguishable for 
a short distance prior to separation. Alp inserts on the ventromedial face of the maxillary head. 
The ramus mandibularis of the trigeminal nerve passes medial to Al P, crosses above the lower jaw 
insertion of A2 + 3, and then passes medial to Ala. 

Muscle A3 is separated dorsally from A2 by Alp, the two former elements joining in a common 
aponeurosis from which Aco originates. 

The levator arcus palatini is not covered by A2 as in the Phycidae, its lateral fibres lying in the 


32 G. J. HOWES 

same plane and meeting aponeurotically those of A2. The ventral surface of the levator is bevelled 
to accommodate the medial surface of A2. 

The opercular muscles are similar to those described for Urophycis and, as in that taxon, a 
segment of epaxial muscle inserts on the medial border of the operculum. Its site of origin, 
however, is the posttemporal rather than the supracleithrum as in Urophycis (but cf. Muraenolepis, 
below). 

In Molva (Fig. 19), muscle Alp occupies a position similar to that in Gadus and Merlangius but 
comprises a long tendon stemming from the point of origin on the hyomandibula and expanding 
anteriorly into a thick bundle of fibres which inserts on the maxillary head. 

Muscle A2 runs from the lateral face of the hyomandibula and preoperculum to insert with A3 
on an aponeurosis from which muscle Aco also originates. 

MURAENOLEPIDIDAE 

(Fig. 22) 

In Muraenolepis (Fig. 22) muscle Al has its origins lateral and medial to A2. Its lateral origin is 
from a thin tendinous sheet covering the face of A2; its medial origin is from a tendinous fascia on 
the inner aspect of that muscle. The two bodies of the muscle join into a single element anterior to 
the border of A2. The lateral part of Al (Ala) inserts tendinously on the dorsal aspect of the 
maxilla; the ventral border of the insertion tendon joining the maxillo-mandibular ligament. The 
main portion of Al (A 10) inserts on the ventromedial aspects of the maxillary head. 

Muscle A2 is large, covering most of the levator arcus palatini laterally. It converges with muscle 
A3 into a thick tendon medial to the anguloarticular. A stout subbranch of the tendon inserts on 
the coronomeckelian bone. Muscle AGO extends from the principal part of the tendon; only its 
anterior tip enters the small mentomeckelian cavity. Muscle A3 is divided from A2 by the levator 
arcus palatini which is a large, laterally bulbous muscle, originating from the sphenotic and 
inserting into a shallow cavity on the lateral face of the hyomandibula. The adductor arcus palatini 
is well-developed, flooring the orbital cavity and inserting anteriorly on the palatine. 

The opercular muscles are well-differentiated from one-another; the dilatator is a narrow, 
spindle-shaped element inserting on the long anterior process of the operculum; the adductor and 
levator operculares originate from the underside of the pterotic and insert close together on the 
anteromedial face of the operculum (see Howes, 1988, fig. 5). As in Urophycis and Lota, a segment 
of epaxial musculature inserts on the medial face of the operculum, being narrowly separated from 
the hyohyoidei adductores. As in Lota, the site of origin of the epaxial segment is the posttemporal 
(see Howes, 1988, fig. 5). 

BREGMACEROTIDAE 

(Fig. 22) 

In Bregmaceros (Fig. 22) muscle A 1 is thin and shallow and incompletely divided. However, below 
the eye there is a strong, dorsal tendon and medial aponeurosis with a slight separation of the 
lateral fibres. The muscle insertion covers a long area of the maxilla, being tendinous anteriorly and 
musculose posteriorly. Muscle A2 is large with a deep concave anterior border. 

The levator arcus palatini is a small thin element lying posterodorsally to the adductor. The 
adductor arcus palatini is divided, the anterior part inserting on the palatine, the posterior on the 
medial rim of the hyomandibula. 

Summary and discussion of the muscles associated with the jaws and suspensorium 

Certain features of these muscles are common to all macrouroids, namely: 

1 . Muscle Al is never separated by A2, as is the case in other gadiforms and divisions Ala and 
Al P lie in the same vertical plane. 

2. Muscle A 1 P always lies lateral to the ramus mandibularis of the trigeminal nerve and, because 
of this relationship, is homologous with that muscle in other gadiforms. 


MACROUROID FISHES 


33 


Fig. 23 Muscle A t P and its associations in: A, Ophidian rochei (muscle Aj + A 2 cut away, its borders 
indicated by dashed lines; B, Glyptophidium macropus; C, Ly codes frigidus. 


3. Muscle Al p never originates from the palatine, in contrast to the condition in some gadoids. 

4. Muscle Alp in the majority of macrouroids, is not constricted below the orbit, nor has it an 
anterior expansion. 

5. The adductor arcus palatini is continuous and never divided as in some gadoids. 

6. The levator arcus palatini lies lateral to muscle A 1 and A2, a feature shared with some gadoids 
but few other teleosts. 

7. Muscle A3 is absent, in contrast to most other gadiforms and acanthopterygians. 


34 G. J. HOWES 

Because of the often contrasting conditions in these features between macrouroids and other 
gadiforms, it is necessary to examine each in detail. 

1. In macrouroids both muscles A Jo. and Alfi always lie lateral to muscle A 2 and the division 
between the Al element is in the vertical rather than the sagittal plane. 

In the Macrouroidinae (Macrouroides and Squalogadus) and some genera of Macrourinae (Type 
II morphotype, see p. 12), A 1 is undivided, or incompletely so. Incomplete separation of Ala and 
Al P occurs in some gadoids (i.e. Bregmaceros, Euclichthys, Lyconus, Steindachneria, Lepidion). In 
other paracanthopterygians (ophidioids, zoarcids and percopsids) there is no Ala, the single 
muscle inserting on the lower jaw. However, it is questionable whether this muscle is the homologue 
of A2 since the ramus mandibularis of the trigeminal nerve runs medial to it, and in some taxa the 
upper portion of the muscle has a close association with the maxillo-mandibular ligament. For 
example, in the ophidiiform Ophidian rochei (Fig. 23 A) the ramus mandibularis runs medial to the 
upper part of the muscle, which is attached to the maxillo-mandibular ligament, the nerve then 
piercing the element and running laterally into the lower jaw. In the zoarcid Lycodesfrigidus (Fig. 
23C) the entire length of the nerve branch runs medial to the outer muscle bloc, but there is a 
distinct lateral myocomma which marks an abrupt change in fibre direction; the dorsal, parallel 
fibres insert directly onto the maxillo-mandibular ligament. In the percopsiform Percopsis (Fig. 
24), although the nerve runs medial to the outer adductor muscle, there is no sign of any fibres 
running onto the maxillo-mandibular ligament or the upper jaw. 

Thus, on the basis of the position of the ramus mandibularis and on what has been said above 
(p. 1 6) concerning its topographical position, the outer muscle in the above mentioned ophidioids, 
zoarcids and percopsids must be construed as the homologue of the element identified as Al in 
macrouroids and gadoids (and various other teleosts) despite the fact that in some cases it does not 
insert on the upper jaw. 

Whether the lack of an upper jaw insertion is the plesiomorphic condition or whether attach- 
ment of the muscle to the upper jaw has been lost, may only be assessed through congruence with 
other synapomorphies. 

In the majority of acanthopterygians the ramus mandibularis of the trigeminal nerve consis- 
tently lies medial to muscle Al (and so by its position signifies the identity of that element) even 
though in some taxa it follows the anterior border of A2. 

2. Muscle Al$ (andAlj). A brief account was given above (p. 16) of the nomenclatural history of 
muscle Al P and it is now treated in detail. 

Rosen (1973), realised, correctly, that the muscle in question is not the homologue of the levator 
maxillaris superioris of halecomorphs, which muscle comprises several sections having their origins 
from the infraorbitals and lateral ethmoid as well as from the palatine and hyomandibula. 

In macrouroids muscle Al P lies lateral to the ramus mandibularis and in some taxa it is incom- 
pletely separated from A 1 a posteriorly. It always inserts on to the medial face of the maxillary shaft 
and is never attached to an element of the suspensorium. 

In assumed primitive gadoids (Bathygadidae; see Howes, 1988), muscle Alp has a similar 
morphology to that in macrouroids and similarly lies in the same vertical plane as muscle Ala. 
However, in progressively more advanced gadoids Alp shifts medially; in Melanonus and 
Steindachneria the shift concerns only the anterior part of the muscle, but in the Euclichthyidae and 
other gadoids (apart from the Bregmacerotidae) the posterior part of Alp also shifts medially so 
that the entire muscle comes to lie mesiad not only to A 1 a, but also (in more advanced gadoids) to 
A2. 

In the Trachyrincidae although there is a single adductor muscle, it has a double insertion on the 
maxilla (p. 17) suggesting that Ala and Alp are fused. According to Casinos (1978) Trachyrincus 
has lost muscle Ala and it has been 'replaced' by a '. . . displacement outwards of the levator 
maxillae superioris' (i.e. A1P). There is, however, no evidence to suggest that such a loss and 
subsequent displacement has occurred. On the contrary, I would advocate that the opposite is the 
case and that muscle A 1 P has shifted medially in gadoids (see above). The situation in Trachyrincus 
is simply a derived specialisation of that taxon. 

Casinos (op. cit.) in studying a limited taxonomic range of macrouroids has failed to take into 


MACROUROID FISHES 


35 


Fig. 24 Percopsis omiscomayus. Upper jaw and suspensorial muscles in lateral view. Medial pathway of 

ramus mandibularis indicated by dashed lines. 


account the varying morphology of the adductor muscle. A single element also occurs in the 
Macrouroidinae and in a group of macrourines (see p. 13), and is probably also the result of fusion 
between A 1 a and A 1 p. 

Lauder & Liem (1983: 148) state that it is only in more advanced paracanthopterygians that 
muscle Al p alone inserts on the maxilla. These authors do not justify that statement by providing 
examples, nor do they include it as a synapomorphy in their cladogram of paracanthopterygians. 
Accepting their statement means that no gadiform can be considered 'advanced'. But, on the 
basis of this character ophidiiforms and percopsiforms must form a monophyletic assemblage. If 
lophiiforms are taken to be 'advanced paracanthopterygians' then Lauder & Liem's hypothesis is 
rejected because muscle A 1(3 is absent in these fishes (muscle Al is a single element inserting on a 
broad ribbon-like maxillary tendon and posteriorly is undifferentiated from A2). 

As noted above, in macrouroids and gadoids muscle Al p lies lateral to the ramus mandibularis 
of the trigeminal nerve. In other paracanthopterygians, however, the muscle lies medial to the 
nerve. A variation of this condition is illustrated in the ophidiiform Ophidian rochei where the nerve 
loops medially around the muscle's origin on the hyomandibula (Fig. 23 A). In another ophidioid, 
Glyptophidium macrops, the nerve also passes medially to Al p, but Ala is lacking (Fig. 23B). in the 
neobythitine ophidiiform Lamprogrammus niger the ramus mandibularis follows a convoluted 
path. In this taxon the outer muscle bloc comprises two elements, the dorsal of which inserts on the 
maxillo-mandibular ligament and the ventral on the mandible. There is a well-developed Alp 
which is divided posteriorly by the levator arcus palatini muscle. The mandibularis nerve passes 
between the divisions of Alp, then medially to the dorsal adductor element. The nerve then runs 
laterally across the upper part of the ventral adductor segment, after which it turns inward to 
course medially to the lower segment. Thus, using the nerve pathway as the criterion of muscle 
identification, not only is the upper segment Al a, but so is the ventral portion of the lower segment, 


36 


G. J. HOWES 


Imm 


Fig. 25 


Lamprogrammus niger. Muscle A , P and its associations. In A the upper part of muscle A 2 is cut 
away, and in B the superficial part is removed with AjCi reflected. 


despite its insertion on the lower jaw. Muscle A2 is that small dorsal portion of the lower segment 
inserting on the mandible (Fig. 25). 

Muscle A I P is reported to occur widely in neoteleosts; according to Lauder & Liem (1 983: 143) it 
occurs in stomiiforms, some acanthopterygians, some paracanthopterygians and some aulopi- 
forms but not in atheriniforms or neoscopelids. This statement appears, in part, to be based on the 
work of Rosen (1973). According to Rosen (op. cit.) there is, in stomiiform fishes, an Alp and 
sometimes an Ala. Fink & Weitzman (1984) maintained that Alp was a neomorphic feature 
independently derived in stomiiforms, myctophiforms and acanthomorphs (Fig. 26C). I agree that 
A 1 P in gadiforms is not homologous with the so-called A 1 P in stomiiforms or myctophiforms, nor 
indeed with that in lower groups in which it has been reported (e.g. halosauran elopomorphs; 
Greenwood, 1977). My reasoning for this assumption is that in those latter groups the muscle in 
question always lies medial to the ramus mandibularis of the trigeminal nerve (see for example 
Tchernavin, 1 953, fig. 29). In higher neoteleosts an A 1 P muscle is recorded by Rosen ( 1 973) in some 
beryciforms. He notes that stephanoberycids have entirely separate internal and external maxillary 
muscles similar to the gadiform condition. However, I find that in Stephanoberyx monae, an 
example illustrated by Rosen, muscles he labels as A2 and A 1 p are united at their origin (Fig. 26B). 
Rosen's A 1 p, which in his figure appears to have a double insertion on the maxilla, is equivalent to 
my Ala, and his Ala corresponds to the upper part of what I interpret as muscle A2 since this 
element inserts on the lower jaw; as shown in Rosen's figure. In the beryciform Hoplostethes, 
muscle A 1 lies laterally to A2 (Fig. 26A). 


MACROUROID FISHES 


37 


In the myctophiforms examined (Lampanyctus, Electrond) and that illustrated by Rosen (1973), 
(Protomyctophum), only an Alp is present and this also lies medial to the ramus mandibularis 
branch of the trigeminal nerve. 

Lauder & Liem's (1983: 143) statement that muscle Alp is absent in atheriniforms needs 
confirmation. In the small sample examined I find that in some taxa (e.g. Orestias) the man- 
dibularis nerve branch passes medially to a segment of adductor muscle which inserts on the 
lower jaw. This could be muscle A 1(3 having secondarily acquired a mandibular insertion. In 
Hemiramphus muscle A 1 appears to be entirely lacking. If the presence of A 1 P is an acanthomorph 
synapomorphy then its absence in various atheriniform lineages must be viewed as derived losses. 

In Polymixia the arrangement of adductor muscles is complex (Fig. 27B). Rosen (1973: 420) 
suggested that the polymixiid pattern was '. . . transitional between the Al and Al|3 systems'. My 
interpretation is somewhat different, however, since the specimen of Polymixia nobilis examined 
differs from that of P. japonica illustrated by Rosen. The principal lateral muscle of P. nobilis 
appears to be a combined Ala and A2 (Rosen's A2) since a group of fibres inserts via a narrow 
tendon on to the posterodorsal part of the maxillo-mandibular ligament and passes laterally to the 
ramus mandibularis nerve branch. A large anteromedial muscle, corresponding to Rosen's Al|3, 
inserts on to the dorsal surface of the maxilla. Although adhering to the maxillo-mandibular 
ligament by connective tissue along most of its length, its fibres remain separated from that 
ligament (Rosen's figure of P. japonica shows them inserting on to the maxillo-mandibular liga- 
ment). As shown by Rosen, there are two sections of muscle Al (3 which are separated by the course 
of the ramus mandibularis branch. The inner section is connected to the outer anteriorly along an 
internal aponeurosis which is evident laterally as a tendinous band (lalb, Fig. 27B). I am unable to 
locate in P. nobilis a muscle corresponding the Rosen's labelled A3 in P. japonica. 


Fig. 26 The superficial upper jaw adductor musculature of the beryciformes: A, Hoplostethes 
melanopus; B, Stephanoberyx monae; and the stomiiform, C, Photichthys argenteus. 


38 G. J. HOWES 

It seems that, at least in P. nobilis, muscle Alp is present, although poorly differentiated, and 
muscle A 1 P corresponds in topographical position to the so-called A 1 P in myctophiforms and 
stomiiforms. This observation supports Stiassny's ( 1 986) phylogenetic positioning of Polymixia as 
the sister-group of the Paracanthopterygii plus Acanthopterygii. 

Rather surprisingly, the adductor muscle arrangement in the Sciaenidae (a family currently 
placed in Acanthopterygii) greatly resembles that of Polymixia and certain gadoids. In Cynoscion 
for example (Fig. 27 A) muscle A 1 P is divided as in Polymixia by the ramus mandibularis branch of 
the trigeminal nerve, the outer section running into the maxillo-mandibular ligament; the inner 
section, which originates from the quadrate and hyomandibula, also joins the maxillo-mandibular 
ligament for part of its length. The anterior, expanded part of the muscle originates from the 
palatine as in merlucciid gadoids (see above). Interestingly, Freihofer (1978: 45) draws attention 
to the resemblances between sciaenid and percopsiform musculature, including the innervation 
pattern. 

One group of macrouroids, designated as morphotype la possesses an additional adductor 
muscle, termed Aly (p. 12). This muscle is usually a weak spindle-shaped element originating via a 
thin tendon from the medial face of Alp and having a common insertion with that muscle on the 
maxilla. In Abyssicola, the tendon of origin stems from the anterior rim of the hyomandibula and in 
this respect resembles the condition of Alp in certain gadoids (see above). I have not found a 
muscle corresponding to Aly in any other acanthomorph taxon and regard it as synapomorphic 
for the macourine genera Coryphaenoides, Abyssicola, Nezumia, Coelorinchus, Lionurus, Nemato- 
nurus and Chalinura. 

3. Origin of muscle A 1$ from the palatine. Among gadoids this feature occurs in the Merlucciidae 
(Merluccius, Macururonus and Lyconus). The anterior part of muscle Al P originates from a lateral 
palatine cavity (complexly so in Lyconus; see p. 26 above). The origin of part of Alp from the 
palatine also occurs in some sciaenids (see above) in which taxa it is considered to have arisen 
independently from that in merlucciids (see Howes, 1988 concerning the phylogenetic position of 
the Merluccidae). 

4. Suborbital constriction of muscle Al$ and its insertion. There is no suborbital constriction of 
muscle A 1 P in macrouroids. In gadoids, however, a tendinous constriction of the muscle occurs in 
the Bathygadidae, Moridae, Melanonidae (sensu Howes, 1988) and Euclichthyidae. Constriction 
of the muscle below the orbit leads to an anterior expansion of the muscle. 

The occurrence of a suborbital constriction of the dorsal part of A 1 in such unrelated groups as 
Cichlidae (Otten, 1981) and Cyprinidae (Howes, 1984a) casts doubt on the feature having any 
phylogenetic significance; it seems to be a functional means of accommodating the eye. That this is 
so is indicated bythe long and obliquely angled jaws of the gadoid taxa possessing the constriction, 
which necessitates a sharp change in angle to pass around the eyeball. In contrast, the jaws of 
macrouroids are short, so that the fibres of muscle Al P are directed downward and their course is 
uninterrupted by the eye. The Trachyrincidae are exceptional in that although possessing long jaws 
they are horizontally aligned and the muscle remains unconstricted (Fig. 7), although the eye is not 
relatively smaller than that in macrouroids. 

Although in primitive gadoids such as Bathygadus and Gadomus (Figs 9 & 1 1 ) the anterior 
'expansion' of muscle Alp follows as a consequence of suborbital constriction, in more advanced 
gadoids this expansion has apparently a functional nature in that it is bulbous and transversely 
expanded, and fills the palatine cavity; in merlucciids this section of the muscle is even attached to 
the palatine (see p. 12). Thus, I have considered the anterior development of muscle Alp to be a 
synapomorphy for gadoids including and above Bathygadus (Howes, 1988). 

The variability of the site of attachment of muscle Al P to the maxilla demands some comment. 
In macrouroids the anterior part of the maxilla bears a prominent ventromedial process which 
contacts the ascending process of the premaxilla (Mvp, Figs 2 & 3). On the inner side of the 
maxillary ventral process is a small depression into which inserts the tendon of Alp (Fig. 28 A). A 
similar ventromedial process occurs in the Bathygadidae and Moridae but is not so well-developed 


MACROUROID FISHES 


39 


Let 


Fig. 27 Jaw adductor muscles in A, a sciaenid Cynoscion jamaicensis and B, a polymixiid Polymixia 

nobilis. 


as that in macrouroids and the tendon of A1J3 inserts on the posterior rim of the ventromedial 
process (Fig. 29 A). 

In the Trachyrincidae, the inner insertion tendon (presumably representing A 1 P; see above, p. 1 7) 
inserts on the dorsal aspect of a medial maxillary shelf (Fig. 29B). This is similar to the situation in 
other gadoids where the muscle insertion is shifted forward to insert along the dorsomedial ledge of 
the maxilla (e.g. Euclichthyidae). In the Melanonidae and Merlucciidae Alp inserts on the medial 
aspect of the maxillary head (Fig. 29C) whereas in the more advanced gadoids (Gadidae, Lotidae, 
Phycidae, Ranicepitidae, Muraenolepididae; see Howes, 1 988) its insertion is on the ventral limb of 
the maxillary head (Fig. 29D). 

The anterior shift of A 1 p insertion appears to be correlated with the transverse expansion of that 
muscle and its shift from a lateral to medial position with respect to the outer adductor muscle (see 
above). It is supposed that the anterior placement of the muscle's insertion site is a derived 
condition for those families listed above. 

5. The adductor arcus palatini is, in some gadoids divided into anterior and posterior parts. 
According to Winterbottom (19740: 238) the adductor arcus palatini in its plesiomorphic state is 


40 


G. J. HOWES 


Mxh 


Fig. 28 A, Nezumia hildebrandi, insertion of muscle AjP on maxilla (ventral view); B, Coelorinchus 
caribbaeus, ventral view of maxillary and premaxillary ligamentous connections; C, Ventrifossa 
occidentalis, lateral view of maxillary-premaxillary-rostral cartilage associations. 


confined to the posterior region of the orbit between the skull and hyomandibula; its derived 
condition is to floor the orbital cavity and extend posteriorly. Part of the adductor in gadoids 
usually originates from the parasphenoid, only a small portion stemming from the prootic; a 
separate adductor hyomandibulae is not recognisable. The division of the adductor arcus palatini in 
some gadoids (Bregmaceros , Macruronus} is, on the basis of in- and out-group distribution, a 
derived condition and a 'precursor' can be recognised in some morids (p. 20) and Steindachneria 
(p. 22) where the central portion of the muscle is thin and weakly developed. 

Apart from Bathygadus, Gadomus and Euclichthys an anterior insertion of the adductor arcus 
palatini on to the palatine is a feature of all gadiform fishes examined, and is also present in some 
ophidiiforms. Among acanthopterygians a palatine attachment of the adductor arcus palatini is 
present in Sciaenidae, Eleotridae and some Cichlidae (see, for example, Greenwood, 1985). Of 
other families checked for this feature, it is lacking in Pomacentridae, Labridae, Atherinidae, 
Nototheniidae, Stephanoberycidae, Polymixiidae, Scombridae, Sparidae and Lutjanidae. 
Admittedly this is not an exhaustive survey of acanthopterygian taxa but it does not indicate that a 
palatine attachment of the adductor arcus palatini is an unusual acanthopterygian condition; such 
an attachment has been treated as derived (Greenwood, 1985: 156; 165). To treat a palatine 
attachment of the adductor as a synapomorphy uniting the majority of gadiforms and some 
ophidiiforms, would conflict with the pattern of relationships arrived at through other synapo- 
morphies (see Howes, 1988). It is more parsimonious to assume an independent derivation of the 
feature in the various lineages in which it occurs. 

6. In the Trachyrincidae, the levator arcus palatini is extensive posteriorly, covering the upper 
part of the plate-like preoperculum and lying medial to Al (Fig. 7). In the Melanonidae and 
Steindachneriidae the levator is small, its insertion being high on the preopercular limb (Figs 12 & 
1 3). A similar situation occurs in the Bregmacerotidae, where the levator is much reduced and inserts 
on the dorsal margin of the preoperculum (Fig. 22), a feature also present in the percopsiform 
Percopsis (Fig. 24). 


MACROUROID FISHES 


41 


The Euclichthyidae has an autapomorphic arrangement of the levator arcus palatini. Although 
extensive, the muscle does not lie laterally to the superficial adductor musculature as in lower 
gadoids and macrouroids, but mostly posteromedial. Near its insertion the levator is bifurcate, the 
posterior segment inserting on the preoperculum and just overlapping the posterodorsal edge of 
Al, the anterior one inserting on the hyomandibula and joining an aponeurosis from which 
originates muscle Al P (Fig. 14). 

The morphology of the levator in Euclichthys could, in evolutionary terms, be construed as the 
'precursor' of the situation found in other gadoids where the entire muscle lies medial to the outer 
part of the adductor mandibulae. 

If the acanthomorph condition of the levator arcus palatini occurring medially to the outer 
adductor muscle be regarded as the plesiomorphic condition, then the similarly placed muscle in 
higher gadoids is a phylogenetic reversal from the laterally placed levator which characterises the 
macrouroids and majority of gadoids. One may then interpret as 'intermediates' between these 
taxa and higher gadoids the posterior shifts of the levator found in the Melanonidae and 
Bregmacerotidae and partial lateral overlap of the adductor in the Euclichthyidae. 

7. Muscle A3 is lacking in macrouroids. This muscle is usually defined as the most medial of the 
adductor complex, having its insertion in the lower jaw (according to Winterbottom 1 914a: 234) on 
the 'medial face of the dentary, in the Meckelian fossa, or both.' Allis (1897: 581) identifies the A3 
as that muscle lying medial to the adductor ramus of the maxillaris inferioris nerve. 

In most acanthomorphs, the ventral fibres of A3 converge with those of A2 on to a common 
aponeurosis which attaches to the inner face of the anguloarticular. Dorsally, A2 and A3 are 
separated by the levator arcus palatini. In macrouroids and many gadoids, the levator lies outside 
the adductor complex (see above) and so the medial adductor bloc comprises a single element 
dorsally . Only in the more advanced gadoids (Gadus, Lota etc.) is it possible to distinguish an A3 on 
the grounds of its dorsal separation from A2 by a levator arcus palatini. In macrouroids and lower 
gadoids, the insertion of the medial adductor element (A2) on the lower jaw is a simple one; all 
fibres converge into an aponeurosis from which departs a ventrally directed tendon. The tendon 
inserts on the coronomeckelian bone; no fibres are associated with the tendon. 

In advanced gadoids the situation is more complex, with the inner fibres of A2 inserting on the 
coronomeckelian tendon, while those of A3 cross over to insert on the lateral tendinous sheet. 

According to Casinos (1978) muscle A3 in gadoids is homologous with the inner part of the 
macrouroid adductor complex. Furthermore, Casinos regards the gadoid ( = my higher gadoids, 
e.g. Gadus, Merluccius, Pollachius) condition of separate A2 and A3 muscles to be plesiomorphic 


,Pmx 


Men 


Fig. 29 Maxillary insertion of muscle A l (3 and the ligamentous connection between the maxilla and 
premaxilla in: A, Bathygadidae, Bathygadus favosus; B, Trachyrincidae, Trachyrincus trachyrincus; 
C, Melanonidae; Melanonus zugmayeri; D, Gadidae, Gadus morhua. 


42 


G. J. HOWES 


a 


lo 


hyad 


Fig. 30 Hyoid musculature of A, Macrouroidinae: Squalogadus modiftcatus; B, Macrourinae: 

Coryphaenoides mexicanus. Ventral views. 


and to have given rise to derived states in macrouroids either by the amalgamation of A2 and A3 or 
the loss of A2. 

Casinos' assumptions are based on the recognition of the gadoids as being plesiomorphic but on 
the contrary they appear to be the derived sister-group of macrouroids (see Howes, 1988). I would 
suggest that the 'amalgamation' of A2 and A3 in macrouroids (and in out-groups) is the plesio- 
morphic state representing an undifferentiated muscle bloc. The differentiation of a medial 
muscle element (A3) is a derived condition that has undoubtedly occurred independently in several 
teleostean lineages. 

Muscles of the hyoid region 

Among macrouroids there is little variability of the hyoid muscles, such variation as does occur 
concerns the points of attachment of the protractor hyoidei and the degree of development of the 
hyohyoidei abductores. 

The protractor hyoidei originates from the anterohyal at the articulation of, and usually attach- 
ing to the 2nd, 3rd or 4th branchiostegal ray. Most frequently the muscle attaches to the proximal 
stem of the 3rd and 4th rays. Only in Lionurus is there a single attachment to the 2nd branchiostegal 
ray. The protractor hyoidei is usually well-developed. In most taxa the left and right parts of the 
muscle continue forward, narrowly separated from one another in the midline. Anteriorly the parts 
diverge slightly to insert on their respective dentary. Echinomacrurus is unusual among the 
Macrourinae in having a ribbon-like protractor hyoidei with the right and left parts separated in 
the midline. In this respect, Echinomacrurus resembles taxa of the Macrouroidinae. In both 
macrouroidine genera the protractor hyoidei is a rope-like element extending from the anterohyal 


MACROUROID FISHES 


43 


anterior to the articulation of the 4th branchiostegal ray; insertion is close to the symphysial tip of 
the dentary (Fig. 30). The left and right parts of the muscle are separated for their entire lengths. In 
Cetonurus the left and right parts of the protractor meet only beneath the ventral hyoids, remaining 
separated for the remainder of their lengths. 

An intermandibularis is present in all taxa examined with a single exception, namely the 
macrouroidine Squalogadus. The muscle is a thin, narrow band of fibres with the protractor hyoidei 
inserting below it. 

The hyohyoidei abductores run from the 1st branchiostegal ray to insert tendinously on the 
contralateral dorsohyal. 

The hyohyoidei adductores are weakly developed in all taxa and usually comprise a few widely 
spaced fibres arranged in narrow bands connecting the branchiostegal rays (Fig. 32). Posteriorly, 
those fibres connecting the last branchiostegal ray with the suboperculum and operculum are 
stronger and more numerous. 

In gadoids the hyoid musculature is generally more strongly developed than in macrouroids. For 
example, in the Muraenolepididae (Fig. 31), the protractor hyoidei is well-developed, attaching to 
the base of the 3rd and the upper part of the 2nd branchiostegal ray; an anterior segment of the 
muscle attaches tendinously to the ventromedial border of the dentary. The parts of either side 
meet in the midline and run forward as a single muscle inserting at the symphysis beneath a small 
intermandibularis. The hyohyoidei abductores and adductores are also well-developed. 

The morphology of the hyoid muscles in the Gadidae, Lotidae and Phycidae is similar to that in 
the Muraenolepididae except that the intermandibularis is more strongly developed in the former. 
Holmquist (1911) has described and figured the hyoid musculature ofGadus in which he identifies 
two sections of the intermandibularis. Winterbottom (19740: 245) concluded that the anterohyal 
section should properly be referred to as the protractor hyoidei. 

The morphology of the hyoid musculature is rather uniform and the often recognised taxonomic 
grouping of macrouroids based on the number of branchiostegal rays, viz. 6 or 7 is not reflected 
by different muscle morphotypes. The most noticeable differences are those between the Macro- 
uroidinae and Macrourinae where in the former the two parts of the protractor hyoidei are ribbon- 
like and separated in the midline and the intermandibularis is absent (at least in Squalogadus). A 
similar separation of the protractor also occurs in the macrourine, Echinomacrurus (see above). The 
osteology of Echinomacrurus is unknown, but its external morphology would indicate that it is a 


hyad 


Fig. 31 Hyoid musculature of Muraenolepis microps; ventrolateral view, position of intermandibularis 

indicated by dashed lines. 


44 


G. J. HOWES 


hyad 


Imh 


Fig. 32 Trachyrincus trachyrincus: above, hyoid musculature seen in oblique ventrolateral view; below, 
ligamentous connections of the lower jaw and hyoid bar seen in ventral view. 


member of the Macrourinae, although a highly derived one. The similarity in the morphology of its 
hyoid muscles with that of the macrouroidines has possibly been independently derived. The 
complete separation of the protractor hyoideiin the midline is an unusual feature amongst teleosts 
and is known elsewhere only in the otophysan Loricariidae (Howes, 1983) and the Gyrinocheilidae 
(pers. obs.). 

The sternohyoideus muscle, connecting the pectoral girdle with the hyoid bar occurs among 
gadiforms in two conditions long and compressed or deep, broad and short. Usually, a long, 
compressed sternohyoideus attaches to a urohyal whose posterior margin is widely separated from 
the cleithrum, whereas a short, broad sternohyoideus is associated with a urohyal whose posterior 
border is in contact with, or narrowly separated from the cleithrum. The posterior limit of the 
sternohyoideus is usually well-defined ventrally by its attachment to the cleithra, but its lateral 
fibres are continuous with those of the body musculature (obliquus infer ioris). 


MACROUROID FISHES 


45 


In macrourids a long, compressed sternohyoideus occurs in Odontomacrurus, Hymenocephalus, 
Chalinura, Echinomacrurus , Cynomacrurus and Lionurus. A stout, broad muscle is present in 
Macruorus, tiezumia, Trachonurus, Malacocephalus and the macrouroidine, Squalogadus. The 
latter is unusual in possessing a stout sternohyoideus associated with a urohyal that is widely 
separated from the pectoral girdle, a feature also possessed by the Trachyrincidae. 

Among gadoids, the Bathygadidae and Moridae have a long compressed sternohyoideus. 
However, in bathygadids the tendons of the paired infracarinalis anterior stretch forward from the 
pelvic girdle to attach via connective tissue to the ventral tips of the cleithra, and continue forward 
into the ventral body of the sternohyoideus. The tendons of the infracarinalis finally insert on either 
side of the urohyal keel (Fig. 35). 

In other gadoids a long compressed sternohyoideus is present in the Merlucciidae where there is a 
well-defined ventral division of the muscle to which the infracarinalis anterior is tendinously linked. 


obv1-3 


Dh 


shl 


rv4 


Fig. 33 Ventral gill-arch musculature of: A, a macrourid, Macrourus berglax; B, a gadoid, Trachyrincus 
trachyrincus. Ventrolateral views. In A the anterior border of the cleithrum is indicated by a dashed 
line. 


46 G. J. HOWES 

In all gadiforms and other paracanthopterygians examined a dorsolateral segment of the 
sternohyoideus inserts on a stout tendon which runs anterodorsally to insert on the 3rd hypo- 
branchial (see Figs 33-36). In acanthopterygians (?all) a similar tendon runs from the dorsomedial 
part of the muscle. 

Ventral gill-arch muscles 

Macrouroids, in common with almost all teleosts possess well-developed obliqus ventrales on the 
1st through 3rd ventral gill-arches (Fig. 33 A; 34). In contrast, gadoids have reduced (or even lack) 
obliqus ventrales on either the 1st or the 1st and 2nd gill-arches (Fig. 35). The Bathygadidae, 
Steindachneriidae, Melanonidae, Moridae and Trachyrincidae have reduced (or in some 
bathygadids lack) the muscles from only the 1st arch (Figs 33B; 34); see Howes, 1988. Reduction 
takes the form of a tendon attaching to the proximal tip of the ceratobranchial and with only a 
minute muscular element being present. In the Bregmacerotidae, the muscles appear to be lacking 
entirely on the 1 st and 2nd gill-arches. 

Winterbottom (\914a: 263) notes that there are a variable number ofrecti ventrales in teleosts. 
The usual acanthomorph condition is for rectus ventrales IV to run between the semi-circular 
ligament connecting the 3rd hypobranchials across the midline, and the 4th ceratobranchial. This 
is also the condition present in the majority of macrouroids, with the exception of Hymenocephalus 
where the anteroventral part of the muscle inserts on the urohyal and in Macrourus, where the 
entire muscle inserts on the bone. Squalogadus is also exceptional in that the rectus ventrales IV 
joins posteriorly to the tendon of the rectus communis and so by-passes the 4th ceratobranchial, 
inserting instead on the 5th. 

Among gadoids, the rectus ventrales IV attaches to the urohyal in Lyconus (Merlucciidae), the 
Ranicepitidae, Phycidae and Muraenolepididae. In other merlucciids, Bathygadidae, Moridae, 
Steindachneriidae, Melanonidae and Lotidae, the muscle attaches together with the rectus 
communis to a dorsal aponeurosis of the sternohyoideus muscle (Fig. 35). In the merlucciid 
Macruronus, the rectus ventrales IV has a long tendon which inserts on the 5th ceratobranchial; the 
muscle itself is separated from the 4th arch, and anteriorly attaches to a complex aponeurosis of the 
sternohyoideus (Fig. 36). 


Table 2 Insertion sites of rectus communis and rectus ventralis IV muscles in macrouroids and gadoids. 
Uh = urohyal; Sh = sternohyoideus; 3Hy = third hypobranchial 


Rectus communis 
Ur Sh 


Rectus ventralis IV 
3Hy Sh 


Uh 


Macrourinae 

(a, majority) 

(b, 5 genera) 

(see Table 1) 

Trachyrincidae 

Bathygadidae 

Moridae 

Euclichthyidae 

Merluccidae 

Melanonidae 

Steindachneriidae 

Gadidae 

Phycidae (part) 

Phycidae (part) 

Muraenolepididae 

Ranicepitidae 

Lotidae 


* and 3Hy 


*(Lyconus) 


MACROUROID FISHES 


47 


Uh 


sh 


Isc 


rv 


Fig. 34 Ventral gill-arch musculature of: A, a gadoid, Bathygadus melanobranchus; B, a macrourid, 

Nezumia hildebrandi. Ventral views. 


In other paracanthopterygians examined, rectus ventrales IV extends from the 3rd hypobranchial 
ligament to the 4th ceratobranchial, namely, the plesiomorphic acanthomorph condition. 

In nearly all macrouroids, as in the majority of acanthomorphs, the rectus communis runs 
from the urohyal to the 5th ceratobranchial. Exceptional taxa are Nezumia, Ventrifossa, Hymeno- 
cephalus, Odontomacrurus and Cynomacrurus , where the rectus communis runs from a dorsal 
tendon of the sternohyoideus to the 5th ceratobranchial. 

Among gadoids, the rectus communis almost always has a direct attachment to the sternohyoi- 
deus; in some taxa the rectus communis joins a tendinous aponeurosis (Merlucciidae, Fig. 36); in 
others, the fibres run into the body of the sternohyoideus and insert on an internal myocomma 
(Trachyrincidae, Fig. 33); and yet others, the rectus communis attaches directly to the urohyal 
(some Phycidae, Ranicepitidae, Muraenolepididae and Lotidae). However, unlike the condition in 
macrouroids and acanthomorphs it attaches to the anterior tip of the bone rather than to the lateral 
face of the keel. This different insertion site on the urohyal suggests that the attachment has been 
secondarily derived to that in acanthomorphs rather than representing a plesiomorphic condition 
(see Howes, 1988). 

The various conditions of ventral gill-arch muscles in macrouroids and gadoids are summarised 
in Table 2. 

Lauder (1983) considered a urohyal attachment of the rectus communis a synapomorphy for the 
ctenosquamates (Myctophiformes, Paracanthopterygii and Acanthopterygii). Because of its 
urohyal attachment, Lauder prefers the term 'pharyngohyoideus' for ctenosquamates rather than 
rectus communis. I have, however, continued the use of rectus communis for gadoids since here there 
is no, or at best, an indirect urohyal insertion. It could be argued that 'pharyngohyoideus' should 
be used for macrouroids, but here too there are exceptions to a urohyal insertion (see above and 
Table 2). 


48 G. J. HOWES 

Following Lauder's assumption that a urohyal attachment for the rectus communis is a derived 
state it would appear that a direct linkage with the sternohyoideus represents a further derived 
condition. If it be assumed that this condition represented a less derived state, i.e. an evolutionary 
'intermediate' position between a hypobranchial and urohyal attachment then it must also be 
assumed that the gadoids are less derived than myctophiforms, a conclusion unjustified on the 
basis of other synapomorphies (see Lauder & Liem, 1983). 

Howes (1987) considered the rectus communis-sternohyoideus linkage to be a synapomorphy 
uniting gadoids. It is now apparent that the feature also occurs in some macrouroids (see above). 
The possession by macrouroids of other synapomorphies lacking in gadoids makes it reasonable to 
assume, however, that the rectus communis-sternohyoideus linkage in the five macrouroid genera 
(see p. 47) is homoplastic. 

Lauder (1983: 26) notes that in euteleosts a rectus ventralis IV commonly originates from the 
urohyal, but that the muscle is mosaically distributed and has probably evolved independently in 
several lineages through the subdivision of the rectus communis. 

The lability of the rectus ventralis IV casts doubt upon its usefulness as a phylogenetic indicator 
and I have preferred to regard its association with the sternohyoideus in the Moridae, Merlucciidae 
and Lotidae as having been derived independently in those lineages; certainly there are no other 
synapomorphies that would suggest a close relationship of these taxa (see Howes, 1988). Further 
discussion of functional aspects of the ventral gill-arch musculature is given on pages 54-55. 

Dorsal gill-arch muscles 

Unlike the ventral gill-arch musculature, there is little variability in the dorsal gill-arch muscles 
among macrouroids apart from the angles at which the levatores are aligned between the gill-arches 
and the cranium, and in the size differences of some muscles. 

The basic pattern present in all macrouroid and gadoid taxa examined is: three levatores externi, 
two attaching to the 1 st and 2nd and the third to the 4th epibranchials (levator IV crosses the otic 
region of the cranium anteroventrally, medial to levatores I and II); two levatores interni, one 
attaching to the 2nd infraphayngobranchial, the other to the 3rd or 4th (Fig. 37); all these muscles 
originate from the intercalar and/or the upper part of the prootic. 

Transversi dorsales run from the 2nd, 3rd and 4th epibranchials to a midline raphe (that serving 
the 2nd and 3rd arches is a single element); an obliqus posterior connects the postero-medial surface 
of the 4th epibranchial with the 5th cerate-branchial (pharyngeal tooth-plate). 

In macrouroids the retractor dorsalis stems from the 3rd and 4th centra to insert tendinously on 
the medial rim of the 3rd pharyngobranchial. In some taxa, e.g. Nezumia (Fig. 38C), a ventral part 
of the retractor inserts on the rim of the 4th pharyngobranchial tooth-plate. 

Lauder (1983) noted that in the gadoid Pollachius, the retractor dorsalis inserts on both 
pharyngobranchials 3 and 4; I find similar sites of insertion in all 'advanced' gadoids, but in 
merlucciids, melanonids, morids and bathgadids the muscle inserts, as in most macrouroids on the 
medial rim of pharyngobranchial 3. 

In the Muraenolepididae, the retractor dorsalis inserts only on pharyngobranchial 4 (Howes, 
1988). Lauder's (1983) and my own observations on acanthopterygians suggest that insertion on 
pharyngobranchial 3 is the plesiomorphic site of attachment and that one involving the 4th 
pharyngobranchial is the derived state. In this respect, the Muraenolepididae is the most derived 
group of gadoid taxa. 

Eye muscles 

Macrouroids and gadoids lack a posterior myodome, apparently a secondary loss (see Patterson, 
1975: 544). The posterior eye muscles originate from a medial septum close to the floor of the 
parasphenoid and run almost lateral to their insertions on the eyeball. The accompanying figure of 
Gadomus (Fig. 39) exemplifies the condition in all macrouroids and gadoids examined (the eye 
muscles of Gadomus are narrower than in most other taxa). In some gadoids the posterior eye 
muscles originate from a small ossified protruberance of the parasphenoid. In Gaidropsarus , 


MACROUROID FISHES 


49 


Merluccius and Brosme the eye muscles pass medial to a vertical parasphenoid-pterosphenoid strut 
bordering the optic fenestra. 

The origin of the posterior eye muscles from the centre of the parasphenoid and their transverse 
orientation is considered a further synapomorphy uniting the Macrouroidei and Gadoidei (see 
Howes, 1988). 


rv4 


Cb2 


Hb1 


Cb5 


pci 


ica 


Fig. 35 Ventral gill-arch and hyoid muscles of a gadoid, Gadomus longifilis. Above, lateral; below, 

ventral view. 


Functional and ecological inferences 
Jaw protrusion mechanisms 

Recent recoveries of live macrouroids (Wilson & Smith, 1985) while encouraging, also indicate the 
difficulties of maintaining these fishes under laboratory conditions. Results so far discount the 
possibility of obtaining direct experimental data on jaw function. One is therefore obliged to derive 
hypotheses of functional mechanisms from morbid anatomical investigation. 

Those who have studied macrouroid anatomy are agreed that the jaws of most taxa are highly 
protrusible, the degree of protrusibility being a corollary of the length of the premaxillary ascending 
process (Okamura, 1970a,6; Marshall & Iwamoto, 1973: 479; Geistdorfer, 1975; McLellan, 1977; 


50 


G. J. HOWES 


Casinos, 1978; 1981). Several models of acanthomorph upper jaw protrusion mechanisms have 
been proposed, but most authors are in agreement that muscle A 1 plays a predominant part in this 
function (see Lauder, 1982: 280; Motta, 1984 for references to, and review of previous literature). 
Rosen ( 1 973) disagrees, however, believing that the development of an A 1 a or A 1 p division of the 
adductor mandibulae* . . . is not dependent on, or even correlated with, the existence of a protrusible 
jaw mechanism . . .'. 

According to Anker (1974) muscle Al serves not only to keep the mouth closed but possibly 
forces protrusion of the premaxilla. Gosline (1981: 15) thought the most likely cause of jaw 
protrusion in acanthomorph fishes is ligament IX, viz. that connecting the rostral cartilage with the 
maxilla. He hypothesised that as the maxilla twists around its articulation with the cranium, it pulls 
ligament IX anteroventrally, thus protruding the premaxilla which is attached to the rostral 
cartilage. This idea does not entirely explain protrusion, however, since the initial twisting of the 
maxilla must be explained in terms of muscular control. Is the maxilla pulled downward and 
inward passively by abduction of the lower jaw, or through direct action of muscles Ala and Al (3? 

McLellan (1977) accounted for upper jaw protrusion in macrouroids by the action of muscle A 1 , 
since she found that pulling on the maxillo-mandibular ligament along the line offeree exerted by 


Cb1 '9 dh 


Vh 


rv4 


re 


pci 


rv4 


Fig. 36 Merlucciidae; ventral gill-arch and sternohyoid muscles of Merluccius merluccius in A, 
lateral and B, ventral views; the latter showing the partitions of the sternohyoideus. C, Macruronus 
magellanicus in lateral view. 


MACROUROID FISHES 


51 


tvd 


Eb1 


obd 


Ie4 


Fig. 37 Dorsal gill-arch muscles of a macrourid, Coelorinchus caribbaeus (dorsal view). 


the fibres of Al produced premaxillary protrusion. However, McLellan hypothesised that the 
macrouroid type of mechanism was '. . . a fundamentally different means of protruding the upper 
jaw from that of Bathygadus* , which she supposed to be produced by rotation of the maxillary head 
(i.e. as in Gosline's hypothesis). 

Geistdorfer (1975) gave a theoretical account of jaw movements in the macourine Ventrifossa 
occidentalis. For the most part, he considered jaw action to be similar to that in acanthopterygians 
and although stating that protrusion is limited by ligaments, he gave no account of arthrology. 

Casinos (1981) attempted an explanation of jaw protrusion in gadids (Gadidae) based on 
observation by high-speed cinematography, and in macrouroids by the extrapolation of these 
data. According to Casinos, there is no or little jaw protrusion in gadids, except Pollachius, whereas 
there is pronounced protrusion in macrouroids. 

Casinos devised a protrusion index (not to be confused with the protrusion index of Okamura, 
1 9706; see below) which showed that although the gadid Pollachius has a high degree of protrusion, 
comparable with that of macrouroids, the 'macrouroid' Trachyrincus has a low protrusion index 
comparable with that of the gadids. 

Casinos' explanations of these apparent anomalies are confusing and rely on different sizes and 
moments of muscle Alp, and are also based on incorrect anatomical data. For example, Casinos 
explains the higher degree of protrusibility in macrouroids as due to an 'additional' rostro- 
maxillary ligament. However, a rostro-maxillary ligament (ligament VII, see p. 6) is present in all 
gadiforms and indeed all acanthomorphs (Stiassny, 1986). 

I would agree with Casinos (1981) that the restriction of muscle Al |3 to the same vertical plane as 
Ala gives the macrouroid upper jaw a degree of freedom greater than that of gadoids where the 


52 G. J. HOWES 

vertical movement of the maxilla appears to be restricted by the obliquely and transversely angled 
A 1 P (particularly so in merlucciids where a short-fibred A 1 (3 runs from the palatine to the maxillary 
head thus affording the maxilla little downward movement). 

As noted above (p. 38) in some gadoids, seemingly those with restricted jaw protrusibility, the 
insertion of A 1 (3 is sited further anteriorly on the maxilla than in those with a greater degree of jaw 
freedom. In macrouroids the insertion of muscle A 1 P is on the ventromedial prominence posterior 
to the maxillary head. As noted above (p. 39) in gadoids the ventromedial process is less prominent 
and in more advanced taxa is reduced to a medial shelf, with the insertion of A 1 P shifted anteriorly 
to what is regarded as the most derived situation, namely to the symphysial border of the maxillary 
head. It was also noted above (p. 39) that the anterior insertion of A 1 P is correlated with the medial 
shift and enlargement of the entire muscle. 

What is possibly an important factor concerning differences in protrusion between macrouroids 
and gadoids is that in macrouroids muscle Ala is attached to the maxilla via the maxillo- 
mandibular ligament (Figs 1-4), whereas in gadoids Ala, although often associated with the 
maxillo-mandibular ligament is fastened to the jaw independently by its own tendon (Figs 9-20). 

Another factor which may affect the degree of protrusibility is the ligamentous connection 
between the maxillary head and the premaxillary ascending process. It was noted above (p. 7) that 
there are different forms of attachment between these bones in gadoids. The ligament (lig. XI) 
either attaches directly to the bone or via a cylindrical chondroid or fibrous element. The latter 
form of attachment (confined to the more plesiomorphic gadoids) suggests a greater degree of 
separation between the bones. In macrouroids the meniscus is a thick disc, loosely interposed 
between the maxillary head and the premaxilla, with ligament IX also attaching to the rostral 
cartilage (Fig. 28B & C). Thus, the only check on a total downward release of the premaxilla is 
ligament XI, which connects the maxilla to the ethmoid. If this ligament is cut in preserved 
specimens, there is a dramatic and passive jaw protrusion. Such is not the case in gadoids where 
muscle Al P runs obliquely from the suspensorium to the maxilla and acts as a brake, but there is a 
pronounced protrusion in those gadoids where A 1 P, like the macrouroids, lies in the same vertical 
plane as Ala (e.g. Bathygadus, Gadomus). Thus, it is suggested that muscle Alp plays an active 
role in both holding and rotating the maxilla and so effecting protrusion of the premaxilla as 
hypothesised by Alexander (1963) and Gosline (1981). 

A further point to be considered is that suggested by Casinos (1981) concerning the role of the 
labial 'ligament' (see p. 8). In Casinos' view in macrouroids the '. . . depression of the lower jaw 
transmits the force by means of the circumbuccal ligament' ( = labial ligament). Since a labial 
ligament is also highly developed in some gadoids (Bathygadus, Melanonus, Merluccius) presum- 
ably the same function applies in these taxa. Casinos' statement is somewhat ambiguous, however, 
particularly as he regards the ligament as extending around the mandibles whereas in fact, there is a 
separate ligament attached to each dentary (see Fig. 9). Nonetheless, the idea that the ligament 
plays a role in protruding the upper jaw appears valid when one considers the direct linkage of the 
ligament from the mandible to the premaxilla and maxilla and that it appears to be a contractile 
element (see p. 8). In this regard, some attention should be paid to the work of Otten (1983) 
who points out the importance of the maxillo-dentary, and the posterior premaxillary-maxillary 
ligaments in jaw protrusion. 

Otten (1983) recognised two groups of acanthopterygian fishes on the basis of jaw protrusion 
morphotypes: 

l.in which the maxilla rotates about its long axis and pushes the premaxilla anteriorly 
(exemplified by Percd). 

2. in which the maxilla pushes, pulls and retains the premaxilla in protruded position (occurs in 
cichlids). He notes, however, that these two groups show some degree of overlap. 

Otten also makes the point that a shortening and steepening of the premaxillary ascending 
process coupled with a caudo-ventral shift in the insertion of muscle A 1 are factors responsible for 
increased biting force. Although Otten's hypotheses were directed toward cichlid jaw mechanisms, 
these principles also apply to macrouroids. Most macrouroids differ noticeably from gadoids in 
their long, steep premaxillary ascending processes and posterior insertion of Al P (see p. 38). 


MACROUROID FISHES 


53 


Motta ( 1 984) reviewed the history of ideas concerning the mechanics of teleost jaw protrusion and 
presented a classification of protrusion types but unfortunately disregarded paracanthopterygians 
in his account. Following Motta's classification the macrouroids and gadoids would seem to fall 
into three of his four categories, namely, type A: protrusion as a result of mandibular depression; 
type B: as a result of maxillary twist, and type C: as a result of neurocranial elevation (the degree of 
development ofepaxialis musculature in many gadoids suggest this). 

Motta also believes that the 'twisting maxilla model' has been overemphasised and that the 
'mandible depression model' of jaw protrusion is probably the dominant type. Motta emphasised 
that the protruded mouth forms a circular orifice which is the most efficient configuration for 
employing suction feeding (see also Osse & Muller, 1980). A circular, protruded mouth profile is 
probably produced by all macrouroids and plesiomorphic gadoids (i.e. Bathygadus, Gadomus). 


Ie2 


Fig. 38 Dorsal gill-arch muscles of: A, a gadoid, Gadomus longifilis; B, a macrourid, Nezumia 
hildebrandi (lateral views); C, Nezumia hildebrandi showing retractor dorsalis insertion (dorsal view). 


Hyoid-opercular mechanisms 

In discussing Bathygadus, McLellan (1977: 1026-7) states that '. . . the motion of the maxilla is 
mediated through a ligament of the interopercle . . .' and '. . . ligamentous connections between the 
interopercle, subopercle and opercle, and the contraction of the levator operculi muscle which runs 
from the cranium to the opercle.' 

Although the opercular series-jaw linkage serves to depress the mandible, its effect on moving 
the maxilla is doubtful. Furthermore, McLellan has overlooked the fact that Bathygadus, in 


54 G. J. HOWES 

common with other gadoids, possesses a different jaw-opercular linkage system from that in other 
acanthomorphs (see p. 9). The interoperculum, instead of being the mediator between the 
mandible and suboperculum as in most acanthomorphs mediates between the mandible and 
preoperculum/hyomandibula. It should be noted that in most macrouroids and all gadoids the 
adductor operculi muscle inserts principally or entirely on the horizontal, opercular process of the 
hyomandibula; although the levator retains some attachment to the operculum, often its area of 
insertion is small and confined to the anterodorsal margin of the bone. 

The adductor operculi normally acts as an antagonist to the dilatator operculi and together 
they elevate the operculum. The consequences of the redirection of the adductor force to the 
hyomandibula are difficult to evaluate, but a rotational component directly coupled via the hyo- 
mandibular-interopercular ligament to the lower jaw is suggested. Another feature to note is that 
because of the anterior shift of the adductor operculi and the reduced insertion area of the levator 
operculi, there is a greater medial surface area available for the insertion of the hyohyoidei adduc- 
tores, and in the Phycidae, Lotidae and Muraenolepididae, for epaxial muscle as well (see p. 32). 

The presence in the Trachyrincidae of a mandibulohyoid ligament (Fig. 32) may be another 
indication of this taxon's closer relationships with gadoids than macrouroids (see Howes, 1987). 
However, as yet, too few comparative data exist on the distribution of a mandibulohyoid ligament 
to comment on its phylogenetic value. It is certainly present in most, if not all basal euteleosts and 
has been reported in clupeomorphs, stomiiforms and percomorphs (see Verraes, 1977; Otten, 1982 
for specific examples). The ligament has also been reported by Holmquist (191 1) in the gadoid 
Gadus, a condition which I can confirm. Furthermore, the ligament is also present in other Gadidae 
examined and the Lotidae and Phycidae, but not in the Muraenolepididae. 

According to Verraes ( 1 977) a mandibulohyoid ligament is a feature of fishes having a long lower 
jaw and short interoperculum. In Trachyrincus the ligament attaches to the central part of the 
anterorhyal (Fig. 32) but in other gadoids it attaches to the posterior part of the anterohyal as in the 
percomorph Perca (see Verraes op. cit.). According to Lauder & Liem (1980: 389) in the salmonid 
Salvelinus the presence of the mandibulohyoid ligament possibly allows for another coupling to 
depress the lower jaw. Such may also be the case in Trachyrincus and other gadoids where it 
is present. Otten (1982: 47) believes that the occurrence of the mandibulohyoid ligament in 
various teleosts is homoplastic. 'Undoubtedly, ligaments are products of evolutionary pathways. 
Theoretically, redundancies may occur along these pathways, but it is more likely that ligaments 
evolve together with the whole apparatus in which they are functional . . .'. 

Finally, it should be noted that macrouroids and gadoids possess an elongate interhyal. Lauder 
& Liem (1980) drew attention to two functional roles of the inferhyal in feeding mechanics, namely: 

giving the hyoid an increased dorso ventral rotation and so providing greater orobranchial 

expansion 

giving the hyoid a posterodorsal movement. 

Although the above discussion has concentrated on the jaw mechanism in terms of feeding, the 
various modifications of the jaws and their couplings in gadoids possibly have a greater significance 
in respiratory function. After all, Smith & Hessler (1974) have pointed out that the respiratory rate 
for cod (Gadus) is over twenty times greater than that for a macrourid (Coryphaenoides). 
Experimental and functional analytic data are needed to assess the significance of the gadoid 
type of lower jaw coupling in which an interopercular-preopercular-hyomandibular ligament is 
introduced. 

Pharyngeal mechanisms 

Geistdorfer (1975) described and commented on the pharyngeal dentition of various macrouroid 
genera but paid no attention to the pharyngeal musculature. There is little variation in both the 
upper and lower pharyngeal muscles in the taxa examined (see p. 48). In general the posterior 
levatores are long, deep and angled at 45, suggesting a high degree of forward movement of the 
upper pharyngeal apparatus. 
The functional significance of a sternohyoideus link with the rectus communis and recti ventrales 


MACROUROID FISHES 55 

IV muscles is speculative. According to Lauder ( 1 983: 25) the shift of the anterior attachment of the 
rectus communis from hypobranchial 3 to the urohyal is a key specialisation in the evolution of 
euteleostean pharyngeal manipulation. A urohyal attachment of the muscle provides an axial 
rotation to the pharyngeal tooth plates. In gadoids the degree of axial rotation allowed by a direct 
sternohyoideus-rectus communis link would seem to be minimal but might facilitate asymmetrical 
activity of the lower pharyngeal tooth-plates. 

The reduction or absence oiobliqus ventrales muscles associated with the 1st and/or 2nd gill arch 
is probably correlated with a strong ligamentous attachment of the 1st gill-arch to the hyoid bar. 


n 


rei 


re 


ey 


Fig. 39 Eye muscles of a gadoid, Gadomus longifilis. Dorsal view of left eye. 


Trophic strategies 

The trophic ecology of macrouroids is poorly known, but it would seem that the majority of taxa 
feed on a broad spectrum of organisms (see Mauchline & Gordon, 1985). This trophic diversity is 
a consequence of the rather unspecialised organisation of the adductor and ventral gill-arch 
musculature described in this text. 

Okamura (1 9706) classified macrouroids into four groups according to their jaw and mouth type 
and devised a protrusion index (a percentage ratio of premaxillary ascending process length to that 
of the ramus), the application of which gives a higher figure to the smaller mouth. Okamura 
attempted to relate the index to ecological categories, those taxa with a low index being predomi- 
nantly nekton feeders, those with a high index being mostly benthic feeders. Okamura's groupings 
included Bathygadus and Gadomus, which are here considered to be gadoids and which according 
to his formula have a low protrusion index. Trachyrincus was not studied by Okamura, but 
application of his formula to this taxon gives it a protrusion index closer to Bathygadus and 
Gadomus than to any macrouroid. 


56 G. J. HOWES 

McLellan (1977) also attempted to correlate head shape with trophic specialisations and 
believed she could match those taxa with long rostra (i.e. elongate nasals) to benthic habitats, 
whereas taxa with large, terminal mouths were associated with benthopelagic feeding. In 
McLellan's view Bathygadus and Coelorinchus represent two extremes of the Macrouridae with 
regard to ratio of mouth to head length, degree of jaw movement and expansion of orobranchial 
cavity. She considered Bathygadus to possess a prey-capture activity similar to that of 'other 
teleosts' with large, terminal mouths. However, as Lauder & Liem (1980: 387) point out, the 
feeding mechanisms of acanthopterygians (e.g. Percd) and basal euteleosts (e.g. Salvelinus) are 
quite different, involving different patterns of muscle activity. McLellan's reference to 'other 
teleosts' presumably refers to acanthopterygians, but the anatomical observations made in this text 
on the mode of jaw coupling in gadoids suggest that a somewhat different type of feeding pattern to 
that in other paracanthopterygians and in acanthopterygians might be operating. 

McLellan (1977: 1034) hypothesised that Coelorinchus and taxa with a protracted rostrum feed 
by using the rostrum as a sediment probe, as did Casinos (1978) for Trachyrincus. McLellan argues 
that this method of feeding prevents rapid forward swimming necessary for creating suction. As 
such she sees a high degree of muscular control in varying the protrusion angle of the upper jaw 
(elsewhere, however, she noted that Coelorinchus has 'weak musculature'). McLellan's assumption 
that rapid swimming is the only means of generating suction is not entirely correct, however, and 
she seems to be confusing inertial suction and ram feeding. For example, Lauder & Liem (1980) in 
their study of the salmonid Salvelinus note that although this fish is not primarily a suction feeder, 
inertial suction is created by varying the sequence of muscle activity, i.e. the activation of the 
levator arcus palatini prior to that of the levator operculi and the hyoid musculature. McLellan 
appears to be working from similar assumptions made by Weihs (1980) that higher swimming 
speeds extend the suction field further forward. However, this strategy would seem applicable only 
to predatory fish using strike tactics rather than hovering fish sucking food from the substrate 
whose principal feeding method would seem more likely to be inertial suction (see Liem, 1980). 

Both McLellan (1977) and Casinos (1978) have hypothesised that the rostrum of macrouroids 
and trachyrincids is used as a probe. Both authors have based their assumptions on the character- 
istic swimming mode of macrourids, shown in Marshall & Bourne (1964) to be head down and 
forming an angle of approximately 45 with the substrate. Isaacs & Schwartzlose (1975) have 
reported, from cinematographic evidence, macrouroids thrusting into the sediment and 'throwing 
a cloud of sediment through the gills'. Marshall (1979) believes this method of feeding enables 
macrourids to screen ooze-laden water by forcing it through the restricted first gill-slit. McLellan 
also supposed that the shorter snout of some species is probably more a sensory device than a 
mechanical probe. It should be emphasised that the rostrum is not part of the snout (i.e. ethmoid) 
but comprises the medially joined nasal bones which are trough-like, house many neuromasts and 
are fluid filled in life. Thus, a sensory function of the 'rostrum' seems a feasible idea. Casinos (1978) 
in his reconstruction of Trachyrincus probing and feeding from the substrate appears to overlook 
the fact that although the mouth may appear to be inferior in relation to the extended nasals, it is, in 
relation to the ethmoid, terminal. Furthermore, the lack of extended premaxillary ascending 
processes, the elongate jaws and the wide gape of the mouth (see p. 17) all point to Trachyrincus 
being benthopelagic rather than benthic in its feeding habits. Marshall & Merrett (1977: 489) point 
out that Trachyrincus '. . . has a marked preference for pelagic food'. 

As mentioned above, macrouroids display little morphological specialisation in their jaw 
musculature but from their trophic diversification (see Mauchine & Gordon, 1985) it would seem 
that they are capable of employing some specialised feeding habits. In this respect their feeding 
mechanisms resemble those of cichlids recognised by Liem (1980) as being suboptimilised. 

The hypothesis presented here and in Howes ( 1 988) that Bathygadus, Gadomus and Trachyrincus 
are not members of the Macrouroidei, but represent clades within the Gadoidei, negates to a 
certain degree the functional hypotheses advanced by McLellan (1977) and Casinos (1978; 1981). 
To those authors, the Macrouroidei was unquestionably a monophyletic group and so functional 
mechanisms identified among any of the included taxa would be considered homologous. The 
identification of a different jaw coupling in Bathygadus, Gadomus and Trachyrincus that is shared 
with gadoids necessitates a reappraisal of macrouroid trophic strategies. 


MACROUROID FISHES 57 

Taxonomic and phylogenetic inferences 

The results of the character analyses in this study have already been utilised in another paper 
(Howes, 1988) to demonstrate the paraphyletic nature of the Macrouroidei. It has been established 
here and in Howes (1987) that four taxa formerly recognised as macrouroids, namely, Euclichthys, 
Bathygadus, Gadomus, and Trachyrincus share myological and arthrological synapomorphies with 
the Gadoidei, viz.: 

anterior expansion of the jaw adductor muscle Al 

reduction ofobliqui ventrales on the 1st and 2nd gill-arches 

rectus communis attaching aponeurotically to the sternohyoideus muscle 

possession of an interopercular-hyomandibular-preopercular ligament. 

As yet, there exists no comprehensive osteological account of Bathygadus and Gadomus, and their 
close affinities among gadoids are obscure. Howes (1988) tentatively recognised the Bathygadidae 
as the sister-group to the Moridae. The basis for this arrangement is the scale pattern shared 
between the two families. The scales are cycloid, with a reticulate pattern of sulci. According to 
Peabody (1931) the scale pattern of Bathygadus '. . . show no affinity for the Macrouridae and could 
easily be classified with either or both of the other families' (i.e. Gadidae and Bregmacerotidae). In 
Okamura's (19706) opinion, the 'bathygadine' scales are not secondarily derived from the ctenoid 
scales present in macrourids but are 'essentially primitive' and he draws attention to the '. . . 
striking resemblance to morids . . . which are characterised by the reticulate structure of ridges on 
the exposed area'. There are, regrettably, no derived myological features shared between the 
Bathygadidae and Moridae, the shared myomorphology being plesiomorphic for gadoids. 

The Bathygadidae was considered by Howes (1988) to represent a plesiomorphic lineage of 
gadoids. Work in progress has identified further synapomorphies that support the sister-group 
relationship of Bathygadus and Gadomus and the distinctiveness of the family (see also p. 18), 
but no further evidence has come to light which would suggest that the family is anything but 
plesiomorphic. 

Trachyrincus and Euclichthys each represent gadoid lineages, the former being recognised as the 
sister-group to all other gadoids, the latter as the sister-taxon of more advanced gadoid families 
(Howes, 1988). 

Intrarelationships of the Macrouroidei 

The attrition of the Macrouroidei by the removal of the Euclichthyidae leaves a single family in 
the suborder, the Macrouridae. In turn, this family has been reduced by the removal of the 
'Bathygadinae' and Trachyrincinae' to two subfamilies, the Macrouroidinae and Macrourinae. 
The former contains two genera, Macrouroides and Squalogadus, the latter some 30 genera of 
diverse external morphology. 

The monophyly of the Macrouridae has been discussed by Howes (1988) and is supported by 
three derived characters; levator arcus palatini muscle enlarged and lying lateral to the jaw adductor 
musculature; trough-like nasal bones meeting in the midline; compressed, plate-like ethmoid. 

The laterally placed levator arcus palatini is a feature shared with the gadoids and is discussed in 
Howes (1988). 

The nasals of macrouroids are large, with a noticeable anteroventral curvature, the anterior 
border is often notched and the medial surface raised, contacting its partner along the midline. 
Enlarged, medially united nasals also occur in the gadoid family Trachyrincidae. Arguments for 
recognising the macrouroid and trachyrincid conditions as homoplastic are given in Howes (1988). 

The ethmoid region of macrouroid taxa comprises a deep, vertical, plate-like bone with an 
expanded base capping a cartilaginous bloc. The plate-like part of the ethmoid divides the posterior 
borders of the nasals and the anterior borders of the frontals. The dorsal margin of the ethmoid is 
confluent with the dorsomedial margin of the nasals, thus forming a long crest. The identity of the 
ossified crest-like cap of the ethmoid is doubtful, but it is easily detached from the ethmoid 
(mesethmoid) cartilage and, as there is no sign of perichondral ossification, it would appear to be a 
rostrodermosupraethmoid. 


58 G. J. HOWES 

Apart from the derived nature of the levator arcus palatini, which is also shared with the 
Gadoidei, no myological synapomorphies have been identified which corroborate the monophyly 
of the Macrouridae. 

MACROUROIDINAE 

Three myological apomorphies have been identified in Squalogadus, one is also known to occur in 
Macrouroides but it has not been possible, with the nature of the material available (see p. 15), to 
observe the other two features. These characters are: 

protractor hyoidei entirely separated in the midline (present in Squalogadus and 

Macrouroides) 

intermandibularis lacking 

rectus ventralis IV joins rectus communis, by-passes and inserts on ceratobranchial 5. 

Okamura (19706) treated the group as a family on the basis of. . . notable differences from other 
macrouroids'. Indeed, from the characters he enumerates there is little doubt that the Macro- 
uroidinae possess a number of uniquely derived features, e.g. anterior and posterior ascending 
processes of the ethmoid (? rostrodermethmoid); division of the orbital fontanel; reduced, 
filamentous lateral ethmoid; deep 1st infraorbital; enlarged 5th infraorbital. Other features listed 
by Okamura (op. cit.} as defining the group, however, appear to be plesiomorphic, viz.: unmodified 
gill-rakers, unrestricted buccobranchial aperture, dorsal fin lacking spinous rays, flattened parietal, 
bar-like parasphenoid. It is more difficult to ascribe polarity to certain other characters, e.g. 
'rostral' cartilage between ethmoid and parasphenoid (Okamura appears to confuse the rostral 
and mesethmoidal cartilages); lachrymal lacking ascending process (this could be a plesiomorphic 
condition or a reversal). 

MACROURINAE 

The characters given by Okamura (\91Qa,b) and Marshall (1973) as defining macrourines are, for 

the most part synapomorphic; they are: 

aperture between operculum and 1st gill-arch restricted by bucco-pharyngeal lining 

(Fig. 38B). 

olfactory bulbs lying close to nasal sac and within the nasal cavity 

spinule-bearing scales 

swimbladder (often) with drumming muscles and high number of retia mirabilia 

presence of light organs (in some taxa). 

The first three characters are common to all genera, but the other two only to some (see Fahay & 
Markle, 1984 for generic distribution of the light organs). No myological synapomorphies have 
been identified that support monophyly of the group. 

Within the Macrourinae three groups of genera can be distinguished on the basis of their jaw 
adductor muscle morphology; see pp. 12-13. These groups largely correspond with Okamura's 
(19706) scheme of generic relationships. Okamura's Coelorinchus group includes Coelorinchus , 
Coryphaenoides and Abyssicola and thus corresponds to my group la (p. 12). He also relates 
the genera Ventrifossa and Malacocephalus; Odontomacrurus and Cynomacrurus, and 
Echinomacrurus, Cetonurus and Sphagemacrurus; groupings which correspond with my group II 
(Table I). This group, however, is probably based on symplesiomorphies. 

Two groups of genera can be distinguished on the basis of their ventral gill-arch musculature, 
namely those where the rectus communis has an aponeurotic attachment to the sterno-hyoideus (as 
in gadoids) and where the muscle inserts on the urohyal. The latter group comprising the majority 
of macrourine taxa, the former includes Nezumia, Ventrifossa, Hymenocephalus, Odontomacrurus 
and Cynomacrurus. All these genera except for one (Nezumia) belong to the jaw muscle morphotype 
II. 


MACROUROID FISHES 59 

Conclusions 

The salient points to emerge from this study of macrouroid and gadoid cranial muscles are: 

Anatomical 

The jaw adductor muscles of macrouroids are, in comparison with those of other 'paracanthoptery- 
gians', unspecialised. Muscle Alp is homologous with that so-called element in other acantho- 
morphs. M uscles A 1 a and A 1 p lie in the same vertical plane, an arrangement which seems to afford 
a large degree of jaw protrusion. 

The arrangement of macrouroid hyoid muscles are typically those of acanthopterygians, viz.: 
anterior attachment of the rectus communis to the urohyal (with the exception of five genera), rectus 
ventralis IV to the 3rd hypobranchial, and a well-developed complement of obliqui ventrales 
muscles. In gadoids, the rectus communis and rectus ventralis IV attach directly to the sternohyoideus 
and the obliqui ventrales are reduced on the 1 st or 1 st and 2nd gill-arches (in some bathygadids they 
are absent from the 1st arch). 

In common with acanthopterygians, macrouroids possess a mandibular-interopercular- 
subopercular ligamentous connection. In gadoids the linkage runs from the interoperculum to the 
hyomandibula, preoperculum or both. In both macrouroids and gadoids the adductor operculi 
muscle inserts wholly, or principally on the opercular process of the hyomandibula. 

Macrouroids, in common with 'lower' gadoids have the levator arcus palatini situated postero- 
lateral to the jaw adductor muscles. This is considered a derived condition and one allowing for a 
high degree of orobranchial expansion. 

Taxonomic and phylogenetic 

The Macrouroidei comprises a single family, the Macrouridae and two subfamilies, Macrourinae 
and Macrouroidinae. Although the monophyly of the Macrouridae is unsupported by myological 
characters, the presence of a maxillary-nasal ligament and a rostral cartilage attachment of ligament 
IX corroborates other synapomorphies. 

Monophyly of the Macrouroidinae is attested by ventral gill-arch and hyoid muscle synapo- 
morphies (pp. 43; 46). No myological characters have been identified as synapomorphic for the 
Macrourinae, although two jaw adductor muscle morphotypes are identified, one of which, pos- 
sessed by the genera Coelorinchus , Coryphaenoides, Abyssicola, Nezumia, Lionurus, Nematonurus 
and Chalinura is considered to be derived. 

Functional 

Previous functional hypothesis of macrouroid feeding mechanisms were based on the assumption 
that the group is monophyletic. The studies of McLellan ( 1 977) and Casinos ( 1 978; 1 98 1 ) included 
taxa which properly belong to the Gadoidei. Indications from jaw-opercular linkages and muscle 
insertions are that gadoids employ a different feeding mechanism from that of macrouroids. 
The extrapolation of data gleaned from functional studies of acanthopterygian fishes to 
'paracanthopterygians' is a flawed approach. 

Ecological 

Hypotheses of trophic strategies have also suffered by the tacit assumption of macrouroid 
monophyly. The ecological and evolutionary scenarios of McLellan (1977) and Casinos (1978) 
must be reassessed in the light of the revised classification of macrouroids and gadoids (Howes, 
1988; Howes & Crimmen, in prep.). 

That the monophyly of a group so seemingly highly characterised as the Macrouroidei should be 
questioned is a warning that functional and ecological hypotheses must be used guardedly and are 
valid only for groups whose monophyly is well corroborated. 

According to Lauder's (1981) 'decoupling hypothesis' there is a phylogenetic increase in the 
number of biomechanical components and their pathways. Thus the derived sister taxon of a group 


60 G. J. HOWES 

displays greater diversity and 'constructional flexibility' than its plesiomorphic sister taxon. The 
Gadoidei are hypothesised to be the derived sister group of the Macrouroidei (see Howes, 1988), 
and as such Lauder's hypothesis is borne out (in part) since the macrouroids lack what might be the 
more manipulative functions of the upper jaws possessed by gadoids. There is also a more complex 
arrangement of the hyoid and ventral gill-arch musculature in gadoids, although it is arguable 
whether a greater range of function is achieved (see p. 54). Only a complete comparative functional 
analysis of feeding mechanisms of taxa in the two groups will support Lauder's claim. 

The intrarelationships of the morphologically diverse genera assigned to the Macrourinae have 
yet to be worked out cladistically. Myological characters have not been rewarding in this regard, 
and synapomorphies must be sought in other soft-anatomical (particularly in the structure of the 
light organs) and skeletal features. 


Acknowledgements 

I am most grateful to Humphry Greenwood, N. B. Marshall, Nigel Merrett and Alwyne Wheeler for their 
critical reading of the manuscript and their many helpful suggestions for its improvement. 

I am particularly indebted to Nigel Merrett of the Institute of Oceanographic Sciences, for supplying 
several macrouroid specimens for dissection and for much helpful discussion. Special thanks are due to my 
colleagues Mandy Holloway and Chris Sanford for preparing radiographs and cleared and stained specimens. 


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1966. The relationships of the anacanthine fishes Macruronus, Lyconus and Steindachneria. Copeia 1966: 
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1979. Developments in deep-sea biology. 566 pp. 

& Bourne, D. W. 1964. A photographic survey of the benthic fishes of the Red Sea and Gulf of Aden. 
Bulletin of the Museum of Comparative Zoology, Harvard 132: 223-244. 

& Cohen, D. M. 1973. Order Anacanthini (Gadiformes). Characters and synopsis of families. Memoir, 
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& Merrett, N. R. 1977. The existence of a benthopelagic fauna in the deep-sea. In: A voyage of Discovery 


(Ed. M. Angel): 483-497. 

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Merrett, N. R., Sazonov, Y. I. & Scherbachev, Y. N. 1983. A new genus and species of rattail fish (Macrouridae) 

from the eastern North Atlantic and eastern Indian Ocean, with notes on its ecology. Journal of Fish Biology 

22:549-661. 

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Usa Marine Biological Station, Kochi University 17 (1-2): 1-179. 
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Amblyopsidae. American Museum Novitates 2109: 1-35. 
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P. H., Miles, R. S. & Patterson, C. Supplement No. 1 to the ZoologicalJournal of the Linnean Society 53: 

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Stiassny, M. L. J. 1981 . Phylogenetic versus convergent relationship between piscivorous cichlid fishes from 

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Manuscript accepted for publication 18 November 1986 


British Museum (Natural History) 

The birds of Mount Nimba, Liberia 


Peter R. Colston & Kai Curry-Lindahl 

For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland ram 
forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Labc 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 


Titles to be published in Volume 54 


The cranial muscles and ligaments of macrouroid fishes (Teleostei: Gadiformes); 
functional, ecological and phylogenetic inferences. By Gordon J. Howes 

A review of the Macrochelidae (Acari: Mesostigmata) of the British Isles. 

By Keith H. Hyatt & Rowan M. Emberson 

A revision of Haplocaulus Precht, 1935 (Ciliophora: Peritrichida) and its 
morphological relatives. By Alan Warren 

Other titles to follow 


r. Dorset 


28APJ 

Bulletin of the [,. 

British Museum (Natural History) 


A review of the Macrochelidae (Acari: 
Mesostigmata) of the British Isles 

Keith H. Hyatt & Rowan M. Emberson 


Zoology series Vol 54 No 2 28 April 1988 


The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
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specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 


Trustees of the British Museum (Natural History), 1988 


The Zoology Series is edited in the Museum's Department of Zoology 

Keeper of Zoology : Mr J. F. Peake 
Editor of Bulletin : Dr C. R. Curds 
Assistant Editor : Mr C. G. Ogden 


ISBN 565 05038 9 

ISSN 0007 1 498 Zoology series 

Vol54 No. 2 pp 63-1 25 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 28 April 1988 


A review of the Macrochelidae (Acari: Mesostigmata) 

of the British Isles / 2 8 APS 1988 j 

Keith H. Hyatt k 

Department of Zoology, British Museum (Natural History), Cromwell Road, London, SW7 5BD 

Rowan M. Emberson 

Department of Entomology, Lincoln College, Canterbury, New Zealand* 


This paper is dedicated to Ernest Browning, M.B.E., who died on 
the 1st September 1987, aged 91 


Contents 

Synopsis. 

Introduction ....... 

Material examined ...... 

Morphology 

Classification 

Genus Geholaspis Berlese s. lat. 

Genus Dissoloncha Falconer. 

Genus Macrocheles Latreille 

Genus Glyptholaspis Filipponi & Pegazzano . 

Genus Holostaspella Berlese .... 
Taxonomic Summary ..... 

Acknowledgements 

References 


BRITISH MUSED! 
(NATURAL HISTORY) 

28 APR ^983 

PRESENTED 


63 


63 

64 

64 

68 

69 

76 

78 

114 

118 

122 

123 

123 


Synopsis 

Thirty-two species of mites of the family Macrochelidae are now known to occur in the British Isles. 
Descriptions are given for the nine species, including one new to science, recorded for the first time, and for 
other selected species. Habitat and distributional data are given and keys to the genera and species for adults 
are provided. Two neotypes and four lectotypes are newly designated. The genus Dissoloncha Falconer, 1923 
is resurrected and four specific names are newly synonymized. 


Introduction 

It is over thirty years since the publication by Evans & Browning (1956) of their work on the British 
Macrochelinae (now generally treated as the family Macrochelidae). This work was a turning point 
in the taxonomic study of the Macrochelidae. For the first time it was relatively easy to identify 
many of the common macrochelids, not only of the British Isles but also of northern Europe 
and to some extent further afield. It was also the first substantial revisionary work on European 
macrochelids since that of Berlese (1918). 

In the intervening time our knowledge of all aspects of macrochelids has increased rapidly. Their 
role as predators of the eggs and larvae of synanthropic flies has been fully realised, and steps taken 
to make practical use of this knowledge. In connection with this, detailed studies of the biology of 


"The first part of this work was done whilst on leave based at the British Museum (Natural History). 


Bull. Br. Mm. nat. Hist. (Zool.) 54 (2): 63-125 


Issued 28 April 1988 


63 


64 K. H. HYATT & R. M. EMBERSON 

some species have been undertaken. Taxonomic work has proceeded at all levels within the family 
both in Europe and elsewhere. Generic concepts have changed, many new species have been 
described and the type specimens of old species re-examined. Groups of closely related species have 
been recognised and in some cases distinguished with the help of breeding experiments. 

It therefore seems appropriate to apply this new understanding of the group to the fauna of the 
British Isles, particularly as part of the acarine collection of the Rev. J. E. Hull, containing some of 
his type material of British macrochelids, has been found and is now housed in the Arachnida 
collections of the British Museum (Natural History). 

Evans & Browning (1956) recorded and gave descriptions of twenty-three species of 
Macrochelidae. It now seems likely that at least one and probably two of the species recorded by 
them are not members of the British fauna. The inclusion of these species was based on old records 
and no British material was seen by Evans & Browning or in the extensive collections examined in 
the present study. The names of seven previously recorded species must be changed. In all, nine 
species, one previously undescribed, are reported from the British Isles for the first time. 

Since the main purpose of this work is to bring our knowledge of the British Macrochelidae up to 
date, species described in detail by Evans & Browning are not redescribed here although amend- 
ments are made where necessary. However, species not previously recorded are described fully. All 
the species are figured; those previously recorded are illustrated, with some minor amendments, 
from the original figures of Evans & Browning, since that paper has been out of print for many 
years, whilst those recorded for the first time are newly illustrated. Notes on morphology and 
classification are included and keys to genera, species groups and species are provided. 


Material examined 

The primary source of material on which this study is based is the large collection of Macrochelidae 
preserved in the collections of the British Museum (Natural History). Over five and a half thousand 
specimens have been examined. All the 'locality information for the British Isles is based on this 
material, except where otherwise noted. 

In addition, macrochelid material in the Berlese Collection, Istituto Sperimentale per la 
Zoologia Agraria, Firenze, was examined. This was mainly for type material of species occurring in 
the British Isles, but much other material, particularly of European species, was also examined. 

Type material of species described by Bregetova & Koroleva (1960) was borrowed from the 
Institute of Zoology of the Academy of Sciences, in Leningrad and parts of the extensive collection 
of North American and tropical macrochelids in the Department of Entomology, Oregon State 
University, Corvallis, were also examined. 

The following abbreviations appear in the text: 

BMNH: British Museum (Natural History) 

ISZA: Berlese Collection, Istituto Sperimentale per la Zoologia Agraria, Firenze, 

Italy. 

ZINL: Institute of Zoology, Academy of Sciences, Leningrad, U.S.S.R. 

OSUC: Entomology Department, Oregon State University, Corvallis, U.S.A. 


Morphology 

A detailed study of the morphology of Glyptholaspis confusa (Foa) has been published by van 
der Hammen (1964), but not all of his interpretations would be accepted by all workers on 
Mesostigmata (Evans & Till, 1965). In general the terminology used here follows Evans & Till 
(1979). 


MACROCHELIDAE OF THE BRITISH ISLES 


65 


Idiosoma 

Dorsum 

The system of setal nomenclature used here was first developed by Lindquist & Evans (1965) for 
the Ascidae and has since been widely applied to other groups of gamasine Mesostigmata. The 
relationships between the setal nomenclature used by Evans & Browning ( 1 956) and that used here 
are shown in Figure 1A and further compared in Table 1 with those of Bregetova & Koroleva 
(1960) and Hirschmann (1957), both of which have been used by various authors. More recently 
Halliday ( 1 986) has compared in detail the various systems of dorsal setal nomenclature used in the 
Macrochelidae and has advocated that of Lindquist & Evans (1965) for general application in this 
family. 

The number of setae on the dorsal shield in British species is remarkably constant, varying from 
56-60, but usually expressed as 28 regularly arranged pairs (Fig. 1 A). The major exception to this 
arrangement is in the opacus species group of Macrocheles (the former genus Macrholaspis 
Oudemans) which always have setae J3 present instead of J2 and, in addition have 1-4 setae, often 
asymmetrically arranged between j6 and J3. M. montanus (Willman) has both J2 and J3 present 
giving 29 pairs and Glyptholaspis confusa typically has one or two small asymmetric setae between 
j6 and J2. The situation in Geholaspis Berlese is more difficult to interpret. It has the usual 28 pairs 
of setae buty'5 andy'6 appear to have migrated posteriorly from their usual positions. 

Venter 

The form and ornamentation of the sternal shield have been used as one of the main characters to 
distinguish the genus Glyptholaspis Fillipponi & Pegazzano from Macrocheles, whilst the pattern 
of lines described by Berlese (1918) is very useful in distinguishing species and species groups of 
Macrocheles (Fig. 1 B). 

The setal nomenclature system of Lindquist & Evans (1965) is used for the opisthogaster. The 
setation of the ventrianal shield is now known to be more variable than previously realised. 
Holostaspella Berlese may have three or four pairs of setae, Zvl lying either on or off the ventrianal 
shield. In all British species it is on the shield (Fig. 22D). The opacus species group of Macrocheles, 
previously recognised as a separate genus, Macrholaspis, largely on the basis of only having Jv2 


Table 1 Chaetotaxy of the dorsal shield. 


Dorsocentral series 


L&E E&B B&K H 


Mediolateral series 


L&E E&B B&K H 


Lateral series 


L&E E&B B&K H 


Marginal series 


L&E E&B B&K H 


J2 
J3 
J4 
J5 
J6 


Dl 
D2 
D3 
D4 
D5 
D6 


J2 D7 
J3 


Fl 
F3 
V 
Dl 
D2 
D4 


D6 
D7 


il 
si 
12 
i3 

14 
i5 


Jl 
J2 


zl 

z2 

z4 
z5 
z6 


Zl 
Z2 


Anterior region (podonotum) 


Ml 

M2 

L2 
M3 
M4 


F2 rl 

Tl s2 

Sc zl 

D3 z2 

II z3 


s2 


LI SI 


Posterior region (opisthonotum) 


L4 
L5 


S4 
S6 


Zl 
Z2 


J5 D8 S8 J5 


Z4 L6 S7 Z3 
Z5 MglO Mil S5 


r3 


s4 Mg3 S2 s5 
s5 L3 S3 s6 
s6 Mg5 M4 s7 


51 Mg6 M6 SI 

52 Mg7 M8 S2 

54 Mg8 M9 S3 

55 Mg9 M10 S4 


r2 
r3 

r4 


Mgl Ml r4 
Mg2 M2 r5 
Mg4 M3 r7 


L & E = Lindquist & Evans ( 1 965) B & K = Bregetova & Koroleva ( 1 960) 
E & B = Evans & Browning (1956) H = Hirschmann (1957) 


66 


K. H. HYATT & R. M. EMBERSON 


Evens t Browning 
1956 


Prtstnt work, after 
Lindquist I Evans, 1965 


Mg1 


2 


pore 1 

linea Eingulata 

linea arcuata 

linea oblique anterior 

linea media transverse 

pore 2 

linea oblique posterior 

areae punctatae 

B 

Fig. 1 A dorsal chaetotaxy of Macrocheles sp. comparing the systems of Evans & Browning (1956) and 
the present work, after Lindquist & Evans (1965); B structure and ornamentation of the sternal shield 
in the Macrochelidae, based on Berlese (1918). 


MACROCHELIDAE OF THE BRITISH ISLES 67 

and Jv3 on the ventrianal shield, has been found to be much more variable. Species exist without 
any of the Jv series on the sclerotised part of the venter (Fig. 1 3B) and others with Jv3, or Jv2 and 3 
or Jvl-3 on the ventrianal shield (Fig. 3B). 

The distribution and type of pore-like structures in the Gamasina has been studied by Athias- 
Henriot (1969). The inguinal pores occur in the anterolateral corners of the ventrianal shield in 
Geholaspis and Dissoloncha Falconer, but are free on the membrane in Macrocheles, Glyptholaspis 
and Holostaspella. 

In the genus Glyptholaspis all males have holoventral shields, whilst in Dissoloncha the males 
have separate sternogenital and ventrianal shields. In Macrocheles and Holostaspella the males 
have either separate sternogenital and ventrianal shields or holoventral shields. 

Gnathosoma 

The terminology used is that of Evans & Till (1979). The chaetotaxy of the pedipalps has been 
investigated by Evans (1964). The numbers and positions of setae in the Macrochelidae are quite 
typical for the free-living Gamasina having 2-5-6-15-15 setae on the trochanter, femur, genu, 
tibia and tarsus respectively. 

The form and length of the brush-like outgrowths of the cheliceral arthrodial membrane are 
important at the generic level, although they do not vary much in the British species. The form of 
the cheliceral dorsal seta is of use in distinguishing species groups in Macrocheles. The shape of the 
gnathotectum is of importance at the generic level and for species groups in Macrocheles (Figs 2B, 
7B, 13B). 

Legs 

Leg I is without ambulacral apparatus in all British species although species with pulvilli in the 
adults and pulvilli and claws in the immature stages have been assigned to the macrochelid genus 
Neopodocinum Oudemans by Krantz (1965). 

The leg setation of the Gamasina has been studied by Evans (1963). Macrochelids have a 
remarkably constant pattern of leg setation (Table 2). The only variation known in species from the 


Table 2 Numbers of setae on leg segments in the Macrochelidae. 

legs I II III IV 

coxa 222 1 

000 

trochanter 1---1 1---1 1---1 1---1 

333 3 

or 1---1 

550 3 

femur 22 2---1 1---1 1---1 

432 1 

32 32 22 22 

genu 2--, --2 2--, --2 1--, --1 1--, --0 

11 11 10 10 

2 2 

or 1-- --1 
1 

32 22 12 12 

tibia 2--, --2 2--, --2 1--, --1 1--, --1 

21 11 11 11 

tarsus 18 18 18 


68 K. H. HYATT & R. M. EMBERSON 

British Isles is in genu IV which has seven setae in M. subbadius, seta/?/, being present, instead of 
the usual six setae. This condition is also found in a few other species of Macrocheles and in the 
mainly tropical genera Holocelaeno Berlese and Neopodocinum. The genus Neopodocinum also has 
only four setae on trochanter III instead of the usual five due to the absence of one of the ventral 
setae. 

The distribution of spurs on the legs of male macrochelids is useful at the generic level; 
Glyptholaspis having spurs on legs II, III and IV, Macrocheles on legs II and often on legs IV and 
Holostaspella only on femur II, if at all. The form and precise distribution of these spurs is also 
useful in distinguishing closely related species, as are the spurs on leg II of female Holostaspella. 

Spermathecal structures 

Recently there have been a number of studies on the method of insemination of gamasine mites and 
of the spermathecal structures. The information derived from this work has proved to be useful at 
all levels of classification in the Gamasina. 

There appear to be two basic types of insemination (Athias-Henriot, 1968), vaginal, or toco- 
spermic, in which insemination is via a median endogynal, cuticular sac and podospermic where it 
is via tubular cuticular invaginations associated with the bases of legs III or IV. In the British 
Gamasina, tocospermic insemination is found only in the families Parasitidae, Epicriidae and 
Zerconidae. Podospermic insemination is characteristic of other gamasine groups. 

Within the podospermic group two main variations of the spermathecal structures (Michael's 
organ) are found, the tubular cuticular invaginations may lead either to paired terminal organs 
(phytoseiid type) or to a single median organ (laelapid type). Paired spermathecal structures are 
found in the Phytoseiidae and some genera of the Ascidae (sensu Lindquist & Evans, 1965), 
median spermathecal structures are found in most other families of the Gamasina, including the 
Macrochelidae. 

The spermathecal complex has been described for a number of species of European Macro- 
chelidae (Petrova, 1960; Costa, 1966a, 1967; Athias-Henriot, 1968) and provides good taxonomic 
characters at the generic and species level. 

The opening, or solenostome, of the tubulus annulatus, is always on the posterior basal margin of 
coxa III. In Macrocheles the infundibulum is well developed, the rami are usually short and the 
sacculus generally consists of two, more or less, spherical lobes broadly joined by a tubular section 
which gives rise to the corniculum posteriorly (Fig. 14F). In a few species the sacculus is unlobed, 
spherical and merges into the corniculum to give an overall pear-shaped median organ (Costa, 
1967). Dissoloncha superbus has a very different sacculus from all other species of Macrochelidae: it 
is large, more or less spherical and strongly convoluted. 


Classification 

Evans & Browning (1956) recognised two subfamilies of Macrochelidae, the Macrochelinae 
Tragardh, 1949 (sic) and the Areolaspinae Tragardh, 1952, based mainly on the patterns effusion 
of the ventral shields. Subsequently Evans (1956), in a radical reappraisal of the classification, 
taking into account the great variation shown in the ventral shields, proposed a new classification 
based on characters of the peritreme, gnathotectum, genital sclerotisation and gnathosoma. In this 
classification the family was split into the Macrochelinae Tragardh (sic) and the Parholaspinae 
Evans 1956 (Areolaspis having been shown to be closely allied to other macrocheline genera). 

Krantz ( 1 969) regarded the two groups as distinct families, as have most subsequent authors, but 
see Karg (1971) and Krauss ( 1 970). Evans' ( 1 956) definition of the group, however, remains almost 
unchanged. 

About sixteen genera of Macrochelidae are distinguished at present, of which only five occur in 
the British Isles. There has been some change in generic limits affecting these genera during the last 
thirty years. 

Macrholaspis is now generally regarded as a synonym of Macrocheles (Krantz, 1962) following 


MACROCHELIDAE OF THE BRITISH ISLES 69 

recognition of species closely related to the type species, Gamasus opacus C. L. Koch, with three 
pairs of setae on the ventrianal shield and lacking denticulate margins to the dorsal shield. 

The genus Glyptholaspis has been split off from Macrocheles to contain several species, including 
two formerly confused under the name M. plumiventris Hull, in the British fauna. The main 
distinguishing features of the genus are the posterior extension of the sternal shields, the crenulate 
reticular pattern of the main shields and the presence of spurs on legs II, III and IV in the male. 
The genus Dissoloncha Falconer is here resurrected for M. superbus Hull, which is shown to 
share characters of the gnathosoma and other features with Geholaspis and to be isolated from 
Macrocheles s. str. 

The generic limits of Holastaspella have been widened to include species without seta Zvl on the 
ventrianal shield* and a greater range of variation in the form of the ventral sclerotisation. 

Geholaspis remains essentially unchanged although some authors give the subgenus Longicheles 
Valle full generic status (Athias-Henriot, 1968). 

A number of species groups have been distinguished in the genus Macrocheles following the 
work of Filipponi & Pegazzano (1962, 1963) on closely related species. This concept has been 
extended by Krantz (1972) and is also used here. 

Key to the genera of Macrochelidae occurring in the British Isles 

1 Femur II armed with a sclerotised spur in the female, seta mv of tarsus II modified into a thick spine 
(Fig. 22B); vertical setae inserted on an anterior projection of the dorsal shield (Fig. 22A). British 
species with four pairs of preanal setae on the female ventrianal shield 

HOLOSTASPELLA Berlese (p. 1 18) 

Femur II unarmed in the female, seta mv of tarsus II unmodified; without anterior projection of the 
dorsal shield. British species never with four pairs of preanal setae 2 

2 Ventrianal shield with inguinal pores on anterolateral corners; gnathotectum lacking lateral pro- 
cesses (Figs 2B, 4E); corniculi three or more time longer than broad (Fig. 3C) .... 3 
Inguinal pores free on post-coxal membrane; gnathotectum with free or fused lateral processes 
(Fig. 7B, D); corniculi no more than twice as long as broad, or if gnathotectum without lateral 
processes and corniculi elongate then with anal shield only (Fig. 13B) 4 

3 Ventrianal shield with five pairs of preanal setae; terrestrial litter species 

GEHOLASPIS Berlese (p. 69) 

Ventrianal shield with three pairs of preanal setae; seashore species 

DISSOLONCHA Falconer (p. 76) 

4 Sternal shield with characteristic reticulate pattern (PI. 3C), extending posterolaterally to level of 
posterior margins of coxae III; legs II-IV of males armed with spurs 

GL YPTHOLASPIS Filipponi & Pegazzano (p. 1 14) 

Sternal shield variously ornamented but never similar to the above, not produced posterolaterally 
beyond the middle of coxae III; legs II and sometimes IV, but not III, armed with spurs and 
tubercles in the male MACROCHELES Lutreitte (p. 78) 


Genus GEHOLASPIS Berlese 
Geholaspis Berlese, 1918. Redia 13: 145. 
TYPE SPECIES. Gamasus longispinosus Kramer, 1876. 

The dorsal shield has 28 pairs of setae which are mostly pilose or plumose distally. The ventral setae 
are mostly simple except towards the posterior lateral margins. The sternal and genital shields are 
similar to those of Macrocheles, but the metasternal plates may be free or fused to the endopodal 
shields (subgenus Cyrtocheles Valle). The ventrianal shield has five pairs of preanal setae and bears 
the inguinal pores (Athias-Henriot, 1969) in the anterolateral corners. Males, where known, have 


*It should be noted that contrary to the statement and figures of Krantz (1967), H. sculpta Berlese, the type species of 
Holoxtaxpella, in fact lacks setae Zvl on the ventrianal shield (Filipponi & Pegazzano, 1967, and personal observation of 
R.M.E.). 


70 K. H. HYATT & R. M. EMBERSON 

holoventral shields. The gnathotectum has an elongate median process that may be toothed or 
bifurcate distally and dentate laterally. The chelicerae may be either short (Geholaspis s. str. and 
Cyrtocheles) with basically tridentate fixed chelae and bidentate movable chelae, or very elongate 
(Longicheles Valle) and multidentate. The spermatodactyl is short and dorsally directed. The 
corniculi are elongate, more than three times as long as broad and the external hypostomal setae 
are anterior to the internals. The males have a small spur on femur II. 

The setation of the dorsum varies from the usual condition as seen in Macrocheles (Fig. 1 A) in 
that setaey'5 are displaced posteriorly, so as to lie mesad and only slightly anterior of/6 in subgenera 
Geholaspis s. str. (Figs 2A, 3A) and Cyrtocheles and considerably posterior to setaey'6 in subgenus 
Longicheles (Figs 4A, 5A). This latter position is associated with a posterior projection of the 
podonotal shield in the protonymphs and is no doubt a consequence of a posterior migration of the 
cheliceral retractor muscles, associated with the massive development of the chelicerae in this 
subgenus. 

Key to subgenera and species of Geholaspis s. lat. recorded from the British Isles 

1 Cheliceral digits short, with not more than 5 teeth (Fig. 2C); gnathotectum more or less triangular, 
median process with prominent lateral projections (Fig. 2B) Subgenus GEHOLASPIS s. str. 2 
Cheliceral digits prominently elongate, multidentate, movable digit with 10 or more teeth (Fig. 

4F); gnathotectum with median process parallel sided, bifurcate distally (Fig. 4E) 

Subgenus LONGICHELES Valle 3 

2 Dorsal setae less than 100 \xm in length; ventrianal shield only slightly wider than long (PI. 4A). 

Geholaspis (G.) longispinosus (Kramer) (p. 70) 

Dorsal setae generally exceeding 1 50 um in length; ventrianal shield conspicuously wider than long 
(Fig. 3B) Geholaspis (G.) aeneus Krauss (p. 71) 

3 Setae z5 simple (Fig. 4A); median process of gnathotectum smooth or minutely denticulate behind 
terminal bifurcation (Fig. 4E); ventrianal shield almost as broad as or broader than long 

(Fig.4B-D) Geholaspis (L.) mandibularis (Berlese) (p. 74) 

Setae z5 plumosej'6 serrate or simple (Fig. 5 A); median process of tectum strongly toothed behind 
terminal bifurcation (Fig. 5C); ventrianal shield noticeably longer than broad (Fig. 5B) 

. Geholaspis (L.) hortorum (Berlese) (p. 74) 

Geholaspis (Geholaspis) longispinosus (Kramer) 
(Fig. 2A-C, PI. 4A) 

The description and synonymy given by Evans & Browning (1956) are unchanged. 

MATERIAL EXAMINED. 127 collections 4 PNN, 16 DNN, numerous 99- 

ENGLAND: Isles of Scilly, Cornwall, Devon, Somerset, Dorset, Gloucestershire, Hampshire, 
Sussex, Surrey, London, Kent, Essex, Cambridgeshire (including Huntingdonshire), Norfolk, 
Suffolk, Hertfordshire, Berkshire, Buckinghamshire, Worcestershire, Warwickshire, Lincolnshire, 
Yorkshire, Cumbria (Cumberland and Westmorland), Northumberland. 

SCOTLAND: Strathclyde (Argyllshire), Dumfries & Galloway (Wigtownshire), Tayside 
(Perthshire), Highland (Inverness-shire, Wester Ross), Inner Hebrides (Mull, Ulva), Shetland. 

WALES: Glamorgan, Gwent, Gwynedd (Merionethshire and Caernarvonshire), Clwyd 
(Denbighshire). 

IRELAND: Clare, Westmeath, Galway, Mayo, Leitrim. 

CHANNEL ISLANDS: Jersey. 

HABITATS. Found in all sorts of forest leaf litter, among dead grass and other decaying vegetation, 
also in moss. Bregetova & Koroleva (1960) also have records from small mammal nests. 

DISTRIBUTION. One of the commonest European macrochelids, found throughout the British Isles 
and Europe generally (Valle, 1953; Balogh, 1958; Bregetova & Koroleva, 1960; Halaskova & 
Kunst, 1960; Johnston, 1970; Krantz, 1972). Emberson (\913a) has reported the species from New 
Zealand where it is presumably adventive. 


MACROCHELIDAE OF THE BRITISH ISLES 


71 


Fig. 2 Geholaspis (G.) longispinosus (Kramer): female A dorsal shield; B gnathotectum; C chelicera. 

After Evans & Browning (1956). 


Geholaspis (Geholaspis) aeneus Krauss 
(Fig. 3A-C) 

Geholaspis (Geholaspis) aeneus Krauss, 1970. Acarologie 14: 38. 

FEMALE. The dorsal shield (Fig. 3 A) measures 11 30 (am long x 840 um wide (Krauss gives 
1050 um x 750 um) and is finely granular. The posterior half is covered by a finely regular reticu- 
lated pattern, whilst anteriorly it is punctate-reticulate towards the lateral margins. There are 28 
pairs of setae. With the exception of setae jl, zl and J5, all exceed 150 um in length. The majority 
are finely pilose, at least in their distal halves. The dorsal pores are conspicuous. 

The ventral ornamentation and chaetotaxy are shown in figure 3B. The sternal shield has a 
characteristic reticulate pattern. The metasternal plates are free. The genital and ventrianal shields 
have strong reticulate ornamentation. The ventrianal shield (460 um long x 630 um wide) is 
conspicuously wider than long and the preanal setae appear to be simple. 

The venter of the gnathosoma is shown in figure 3C. The corniculi measure c. 140 um in length, 
and the setae appear simple. The chelicerae and gnathotectum are not visible in the only specimen 
available for study. The leg setae are normal for the genus. The majority are pilose, whilst those on 
tarsus I and distally on the remaining tarsi are simple. 


72 


K. H. HYATT & R. M. EMBERSON 


Fig. 3 Geholaspis (G.) aeneus Krauss: female A dorsal shield; B ventral sclerotisation; C venter of 

gnathosoma. 


MATERIAL EXAMINED. 1 collection 19, in moss. 
IRELAND: Mayo. See below. 
This is the first British record. 

REMARKS. This species was described from protonymph, deutonymph and female stages collected 
at Valle de Lozera, Puento de Lozera (600 m), Lugo Province, in northwest Spain (Krauss, 1970). 
The habitat is given as the foot of an old sweet chestnut Castanea saliva tree and an oak Quercus 
toza tree in a dry river-bed. We have tried to obtain specimens, but have been informed by Dr W. 
Hirschmann that Dr Krauss (pers. comm.) has no specimens in his possession. 

The Halbert collection contains a single slide preparation labelled "Holostaspis longispinosus 
(Kram.), 19, Clare Island, in moss, HI/1910'. The idiosoma of this specimen has been damaged on 
the slide and it is felt at present inadvisable to dismount it. However, the dorsal and ventral 
chaetotaxy and the venter of the gnathosoma are clearly discernible. 


MACROCHELIDAE OF THE BRITISH ISLES 


73 


Fig. 4 Geholaspis ( Longicheles ) mandibularis (Berlese): female A dorsal shield; B-D variation in the 
form of the ventrianal shield; E gnathotectum, F chelicera. After Evans & Browning (1956). 


74 K. H. HYATT & R. M. EMBERSON 

Geholaspis (Longicheles) longulus (Berlese) 

The presence of this species in the British Isles is now doubtful. The discovery of a specimen in the 
Hull Collection with the data 'Macrocheles longulus dead fowl' is probably the one mentioned 
(Hull, 1 9 1 8) as being caught 'at one of my carrion traps'. This is a specimen of G. longispinosus. The 
specimens recorded by Halbert (1915) from Clare Island, Mulranny and Castlebar, all in Co. 
Mayo, have been examined and are G. mandibularis. 

Geholaspis (Longicheles) mandibularis (Berlese) 
(Fig. 4A-F) 

The description and synonymy given by Evans & Browning (1956) do not require amendment. 
TYPE MATERIAL. I lolotvpc 9, Cansiglio. Slide 2/34 [ISZA]. 

MATERIAL EXAMINED. 1 14 collections 3 PNN, 29 DNN, many $9. 

ENGLAND: Isles of Scilly, Cornwall, Somerset, Dorset, Gloucestershire, Hampshire, Surrey, 
Sussex, London, Kent, Middlesex, Hertfordshire, Suffolk, Cambridgeshire (including 
Huntingdonshire), Berkshire, Lancashire, Cumbria (Westmorland), Northumberland. 

SCOTLAND: Strathclyde (Argyllshire), Dumfries & Galloway (Wigtownshire), Inner Hebrides 
(Mull, Ulva, lona), Highland (Ross & Cromarty), Shetland. 

WALES: Glamorgan, Gwent, Gwynedd (Merionethshire and Caernarvonshire). 

IRELAND: Clare, Sligo, Galway, Mayo, Leitrim, Westmeath. 

HABITATS. A wide variety of litter habitats, also turf, soil, moss, ants' nests and nests of small 
mammals. 

DISTRIBUTION. Found throughout the British Isles and widespread in Europe (Valle, 1953). The 
specimens recorded by Halbert (1915) from Co. Mayo, as Holostaspis longulus Berlese, have been 
examined and are G. mandibularis. 

Geholaspis (Longicheles) hortorum (Berlese) 
(Fig. 5A-D, PL 5A) 

Holostaspis longulus var. hortorum Berlese, 1904. Redia 1: 265. 
Macrocheles (Geholaspis) hortorum: Berlese, 1918. Redia 13: 145. 
Geholaspis (Longicheles) mandibularis hortorum: Valle, 1953. Redia 38: 349. 

FEMALE. Generally very similar to G. mandibularis but differing in numerous details. -The dorsal 
shield (Fig. 5A), which measures 770-880 um long x 440-500 um wide, is more tapered posteriorly 
than in G. mandibularis. The setae are arranged as in G. mandibularis, except that z5 is plumose and 
z6 is occasionally dentate; these setae are simple in G. mandibularis. Setae J5,j6, J2 and J5 are 
simple, all other dorsal setae are plumose. In the figured specimen there is, on the left side, an 
additional seta between and slightly below SI and Zl. The dorsal shield is ornamented with 
conspicuous, dense, small denticles which diminish towards the centre of the shield, becoming fine 
granulation. The lateral margins are crenate. 

With the exception of setae Zvl and Zv2 the setae of the ventral shields are simple. The sternal 
shield has a characteristic reticulate pattern with fine punctures (PI. 5A). The metasternal plate- 
lets are free and ovate. The genital and ventrianal shields have reticulate ornamentation. The 
ventrianal shield (Fig. 5B) is longer than broad (290-360 um long x 260-305 um wide) and has 
prominent pores at its anterolateral corners and on its posterolateral margin. The ventrianal setae 
are noticeably shorter than in G. mandibularis. Setae Zvl are pilose and Zv2 are plumose. The 
shield is ornamented with clear reticulation. Only the tubuli and rami of the spermathecal 
apparatus are normally visible. 

The gnathotectum (Fig. 5C) has the median process bifurcate and almost fimbriate distally; 
posterior to the bifurcation the process is distinctly toothed while basally there is a series of 
irregular lateral teeth. The cheliceral dorsal seta is simple. The chelae are elongate and multidentate 
(Fig. 5D). The fixed chela has a main row of about 10-15 teeth with the third or fourth tooth from 


MACROCHELIDAE OF THE BRITISH ISLES 


75 


Zv1 


Fig. 5 Geholaspis (Longicheles) hortorum (Berlese): female A dorsal shield; B ventrianal shield; C 

gnathotectum; D chelicera. 


76 K. H. HYATT & R. M. EMBERSON 

the distal end noticeably larger than the others and a subsidiary row of 6 or 8 teeth on the exterior 
face extending posteriorly from the large tooth. The movable chela (c. 198 urn) has a main row of 
about nine teeth with the second noticeably larger, whilst there is a subsidiary outer row of about 
5-6 teeth. 

Most leg setae are pilose, except for those on tarsus I, the distal part of tarsi II-IV, and the setae 
on coxae I and II, trochanters I and II and ventrally on femora I and II. 

MALE. Unknown. 

TYPE MATERIAL. Not seen, identification based on Valle's (1953) redescription of Berlese's type 
material. 


MATERIAL EXAMINED. 8 collections 1 PN, 1 DN, 13 9$. 
ENGLAND: Yorkshire. 

HABITAT. The first British record. From semi-natural grasslands in the Yorkshire Wolds. 

DISTRIBUTION. Valle (1953) lists material from Italy, Switzerland, Austria, Belgium, Iceland and 
Germany, however it was not reported from Germany by Karg (1971) or Krauss (1970). 

REMARKS. This species is most clearly separated from G. mandibularis on the setation of the dorsal 
shield, the longer than broad ventrianal shield, details of the gnathotectum and the dentition of the 
chelicerae, which in British specimens of G. mandibularis are characterised by more numerous 
(fixed chela 1 8-20 teeth, movable chela 14-15 teeth) and smaller teeth, the fixed chela also lacks the 
clear subsidiary row of teeth found in G. hortorum. There seems to be considerable variation in the 
cheliceral dentition in continental specimens of both species and a complex of forms could be 
involved. Since G. hortorum and G. mandibularis occur together in Yorkshire, and appear to overlap 
in much of their continental ranges, they must be regarded as distinct species rather than subspecies 
as suggested by Valle (1953). 


Genus DISSOLONCHA Falconer 
Dissoloncha Falconer, 1923. Naturalist, Hull 1923: 151. 
TYPE SPECIES. Macrocheles super bus Hull, 1918. 

The dorsal shield has 28 pairs of setae which are mainly pilose distally; it has a distinct border and 
crenulate lateral margins. The ventral setae are all pilose distally with the exception of the paranals. 
The sternal and genital shields are similar to those of Macrocheles, but have distinctive patterns. 
The ventrianal shield has three pairs of preanal setae and bears the inguinal pores (Athias, 1969) 
in the anterolateral corners. There may be up to three pairs of muscle apodomes between the ven- 
trianal and genital shields, usually two pairs of these adjoin the ventrianal shield. Males have 
separate sternogenital and ventrianal shields. The gnathotectum tapers into an elongate median 
process, dentate laterally and bifurcate distally. The female chelae are elongate and basically 
bidentate; the male chelae are shorter, the fixed chela has four to five teeth and the movable chela is 
unidentate, the spermatodactyl is short, blunt and dorsally directed. The corniculi are elongate, 
more than three times as long as broad at the base and the external hypostomal setae are anterior to 
the internals. The males have major spurs on legs II and IV. The spermathecal sacculus is large, 
spherical and strongly convoluted, its diameter is greater than the distance between coxae IV. 

The most distinctive features of Dissoloncha are the shape of the gnathotectum, the elongate 
corniculi and the more distally placed external hypostomal setae, characters which are all shared 
with Geholaspis s. lat. The placement of the inguinal pores on the anterior lateral corners of the 
ventrianal shield, a feature also found in Geholaspis, is not simply a reflection of the lateral 
expansion of the ventrianal shield as species of Macrocheles in which the shield is strongly 
expanded still have the inguinal pores free on the membrane. The structure of the sacculus is unique 
within the Macrochelidae, as is the habitat of rotting seaweed and tidal wrack. 


MACROCHELIDAE OF THE BRITISH ISLES 


77 


Fig. 6 Dissoloncha superbus (Hull): female A dorsal shield; B lateral margin of dorsal shield; C 
gnathotectum; D chelicera: male E chelicera; F distal end of gnathotectum; G leg II; H leg IV. After 
Evans & Browning (1956). 


78 K. H. HYATT & R. M. EMBERSON 

Dissoloncha super bus (Hull) 
(Fig. 6A-H, PI. 3D) 

Mac rochele s superbus Hull, 1918. Trans, nat. Hist. Soc. Nor thumb. 5, 1: 71. 

Dissoloncha superbus: Falconer, 1923. Naturalist, Hull 1923: 151. 

Macrocheles superbus: Evans & Browning, 1956. Bull. Br. Mus. nat. Hist. (Zool.) 4: 38. 

The description of this species given by Evans & Browning (1956) remains adequate. 

MATERIAL EXAMINED. 25 collections 3 PNN, approximately 45 DNN, 28 <$<$ and 200 99. 

ENGLAND: Cornwall, Dorset, Kent, Essex, Yorkshire, Northumberland, Durham. 

SCOTLAND: Highland (Inverness-shire), Outer Hebrides (Lewis), Fife, Shetland, Fair Isle, 
Dumfries & Galloway (Wigtownshire). 

WALES: Menai Straits, Milford Haven. 

HABITATS. Common in rotting seaweed on beaches, also found in a salt marsh and in rotten grass 
on a beach. Bregetova & Koroleva (1960) reported it from gulls' nests. 

DISTRIBUTION. Probably around the entire coast of the British Isles; also northern Europe, 
Germany (Krantz, 1972), N. America (Krantz, 1972) and Kuril Islands (Bregetova & Koroleva, 
1960). This is quite possibly a holarctic seashore species. 


Genus MACROCHELES Latreille 

Macrocheles Latreille, 1829. In Cuvier, Regne animalium 2nd ed. 4: 282. Type species: Acarus marginatus 

Hermann, 1804 = Acarus muscae domesticae Scopoli, 1772. 

Coprholaspis Berlese, 1918. Redia 13: 146. Type species: Holostaspis glabra Miiller, 1860. 
Nothrholaspis Berlese, 1918. Redia 13: 169. Type species: Holostaspis tridentinus G. & R. Canestrini, 1882. 
Monoplites Hull, 1925. Ann. Mag. nat. Hist. (9) 15: 215. Type species: Macrocheles (Monoplites) oudemansii 

Hull, 1925 = Macrocheles marginatus Oudemans, 1901 nee Hermann, 1804. 

Macrholaspis Oudemans, 1 93 1 . Ent. Ber. 8 No. 1 80: 272. Type species: Gamasus opacus C. L. Koch, 1 839. 
Andrholaspis Turk, 1948. Proc. zool. Soc. Lond. 118: 103. Type species: Andrholaspis trinitatus Turk, 1948. 

The dorsal shield has 28-30 pairs of setae and smooth or dentate lateral margins; the dorsum lacks 
an anterior extension bearing setae jl. The sternal shield does not extend posteriorly beyond the 
middle of coxae III. The metasternal platelets are free, usually small, rounded and bear the 
metasternal setae. The ventrianal shield has 0-3 pairs of preanal setae, depending on the extent to 
which it is reduced. If it is reduced there are 1-3 pairs of platelets (muscle apodemes) between it and 
the genital shield. The peritrematic shield is not fused to the expodal shields. The males either have 
holoventral shields or separate sternogenital and ventrianal shields. The gnathotectum is usually 
tripartite, the lateral processes may be free, fused basally or strongly reduced. The chelicerae are 
strong, the dentition is variable, the cheliceral brushes are shorter than the movable digit, the 
dorsal seta may be simple, spatulate or pectinate. The leg chaetotaxy is normal for the family 
(except M. subbadius Berlese which has seta pl t present on genua IV). The legs of the females are 
without spurs, the males have spurs on leg II and often on leg IV. 

The main change to the definition of Evans & Browning (1956) has been its widening to include 
species formerly placed in Macrholaspis Oudemans (Krantz, 1962) following the realisation that 
there are species closely related to M. opacus (C. L. Koch), its type, with three pairs of preanal 
ventrianal setae. 

Ecological and morphological grouping of the species of Macrocheles 

The species of the genus Macrocheles fall into two broad categories on ecological grounds which 
correlate with certain morphological features. There are those species that are usually found in leaf 
litter, moss, nests of birds and small mammals and other habitats not predominantly associated 
with coprophilic insects, and there are those species that are usually coprophilic, but also found 
in compost heaps, rotting grass clippings, carrion and similar habitats, generally favoured as 
breeding grounds by synanthropic muscoid flies. 


MACROCHELIDAE OF THE BRITISH ISLES 79 

Species of the latter grouping are often found phoretic, as females, on coprophilic and necro- 
philic insects, for instance dung beetles, burying beetles and synanthropic flies. Males of these 
species are rarely found, but when they are, they are usually strongly dimorphic in the shape of the 
dorsal shield and in the number of pilose dorsal setae, which tends to increase. Well developed 
spurs are usually found on legs IV of the males of this group as well as on legs II. In the European 
fauna species of this group tend to have a preponderance of simple setae that are only faintly pilose, 
but this feature is not constant, particularly in the tropical macrochelid fauna. 

Species of the former grouping are generally not phoretic on coprophilic insects, and in some, 
males are commonly found. The males are not strongly dimorphic, although smaller and with a 
more tapered dorsal shield. The number of pilose setae remains approximately the same in the 
males. Well developed spurs are usually only found on legs II although there may be minor 
tubercles and ridges on legs IV. There tends to be a greater number of strongly pilose setae than in 
the former grouping. 

A few species combine the characters of both these categories, M. penicilliger is found phoretic on 
insects, is not strongly dimorphic and has well-developed spurs on legs IV of the male. M. matrius is 
very often associated with chicken manure and compost heaps, but is not usually phoretic on 
coprophilic insects; it is not strongly dimorphic but has well developed spurs on legs IV of the male. 
Both these species have a preponderance of strongly pilose setae. 

The species of Macrocheles found in the British Isles may therefore be grouped as follows: 

Leaf-litter species Coprophilic species 

M. decoloratus M. muscaedomesticae 

M. punctoscutatus M. robustulus 

M. rotundiscutis glaber species group 

carinatus species group subbadius species group 
opacus species group 

Intermediate species 

M. matrius 
M. penicilliger 

It is interesting to note that Krantz (1981) has shown that the glaber species group, the subbadius 
species group, and M. robustulus, share characters of the ambulacra in the immature stages not 
found in other species of Macrocheles, and that M. penicilliger is intermediate between the two 
main types of ambulacral structures. 

Most of the coprophilic species have been shown to be specialised predators on eggs and young 
larvae of muscid flies and also on the nematodes and small enchytraeid worms found in their 
habitat. The leaf-litter group are more likely to be general predators on small anthropods and other 
animals in their habitat, although biological data are much more fragmentary for these species. 

The coprophilic way of life and the characters associated with it are probably derived from the 
more generalised leaf-litter species. This is supported by comparison with the genus Geholaspis 
which is all litter dwelling and is the most plesiomorphic group of macrochelids. However, males 
are very rare or unknown in most species of the genus Geholaspis and in the opacus species group 
which makes it difficult to draw positive conclusions about their relationships. 

Key to the females of the species of Macrocheles occurring in the British Isles 

1 All setae on dorsal shield simple, needle-like, setae jl short, spine-like; sternal shield with lineae 
oblique anteriores joined by four or five transverse lines, the most posterior of which is the linea 
media transversa (PI. IF) ........ subbadius species group 2 

Some dorsal setae, always including jl, pilose, at least distally; sternal shield variously 
ornamented, sometimes without regular lines and never as above 4 

2 Sternal shield with lineae oblique anteriores connected by five lines (PI. 2A), linea media transversa 
straight or slightly curved posteriorly; genu IV with six or seven setae. ..... 3 


Plate 1 Sternal, genital and ventrianal shields of the females of: A Macrocheles muscaedomesticae 
(Scopoli); B M. robustulus (Berlese); C M. glaber (Muller); D M. punctoscutatus Evans & Browning; E 
M. scutatus (Berlese); F M. insignitus Berlese. After Evans & Browning (1956). 


Plate 2 Sternal, genital and ventrianal shields of the females of: A Macrocheles merdarius (Berlese); B 
M. montanus (Willmann); C M. carinatus (C. L. Koch); D M. penicilliger (Berlese); E M. submotus 
Falconer; F M. tardus (C. L. Koch). After Evans & Browning (1956). 


82 K. H. HYATT & R. M. EMBERSON 

Sternal shield with lineae oblique anteriores connected by four lines (PI. 1 F), linea media transversa 
arched forwards, all sternal shield lines with a strong edging of punctures; genu IV with six setae, 
pi, absent .......... M. insignitus Berlese (p. 1 1 3) 

3 All lines of the sternal shield with a strong edging of punctures (PI. 6E); reticulations of dorsal 
shield densely covered with minute punctures (Fig. 19A); genu IV with seven setae, pi, present 

. . . . M.subbadius (Berlese) (p. 110) 

Transverse lines of sternal shield poorly marked and punctured (PI. 2A); dorsal shield reticu- 
lations more or less unmarked with punctures; genu IV with six setae, />/, absent 
M. merdarius (Berlese) (p. 113) 

4 With 1-3 pairs of postgenital platelets or apodemes; ventrianal shield sometimes reduced in size, 

sometimes with less than three pairs of preanal setae (PI. 4B) 5 

Without separate postgenital platelets, muscle attachments on ventrianal shield; ventrianal shield 

of normal size, always with three pairs of preanal setae . 12 

5 All dorsal setae strongly pilose throughout their entire length, setae J3 always present, either with 
J2 also present or with unpaired setae betweeny'6 and J3\ ventrianal shield with two or three pairs 
of setae, or reduced to an anal shield (PI. 4B); lateral elements of gnathotectum free (Fig. 12D), 
reduced in size or absent (Fig. 1 3D) . . . . . opacus species group (partim) 6 
Some dorsal setae, including j6, z5, z6, J2 and J5, simple, others only pilose in their distal half 

or two thirds, setae J3 absent (except M. montanus (Willmann)); ventrianal shield always with 
three pairs of preanal setae; lateral elements of gnathotectum fused basally (Fig. 1 1 B) 

carinatus species group 9 

6 Opisthogastric sclerotisation reduced to an anal shield, without preanal setae (Fig. 13B); lateral 
elements of gnathotectum absent, with single median process bifurcate distally (Fig. 13D) 

M. emails sp. nov. (p. 96) 

Opisthogaster with a ventrianal shield with two or three pairs of preanal setae; gnathotectum with 
reduced lateral elements present (Fig. 12D) 7 

7 Ventrianal shield with two pairs of preanal setae 8 

Ventrianal shield with three pairs of preanal setae M. terreus (Canestrini & Fanzago) (p. 103) 

8 Anterior portion of dorsal shield with a network of minute spicules (Fig. 12B), lateral margins 
with small rounded serrations (Fig. 12C), with a pair of setae in the J2 position (Fig. 12A) 

M. opacus (C. L. Koch) (p. 96) 

Anterior portion of dorsal shield without spicules (Fig. 14B), lateral margins with sharp pointed 

serrations (Fig. 14C), with three or four unpaired median setae between j6 and J3 (Fig. 14A) 

M. dentatus (Evans & Browning) (p. 101) 

9 Dorsal setae zl smooth, nearly as long as or longer than setaey'7, and always extending beyond the 

bases of setaey'2 (Fig. 11 A, E) 10 

Dorsal setae zl smooth or pilose, much shorter than setae jl and never reaching the bases of setae 
y2(Fig. 10A,E) . . 11 

10 Dorsal shield never with more than six pairs of smooth setae (j6, zl, z5, z6, J2, J5), setae j5 as long 

as setae j4 and lightly pilose M. tardus (C. L. Koch) (p. 94) 

Dorsal shield with more than six pairs of smooth setae, setae j2,j5, s2, r3, r4 in addition also 
smooth (Fig. 11 A) M. submotus Falconer (p. 92) 

11 Dorsal shield with 29 pairs of setae, J3 present (Fig. 10E) . M. montanus (Willmann) (p. 92) 
Dorsal shield with 28 pairs of setae, J3 absent, setae j5 shorter than setaey 

M. carinatus (C. L. Koch) (p. 90) 

12 Some dorsal setae, at least a group in the middle of the dorsal shield, including setaey'6, z5, z6 and 

J2, simple needle-like, not pilose (Fig. 9) 13 

All dorsal setae pilose (Fig. 14A) 20 

1 3 Most setae on the dorsal shield simple, setae s6, SI and S2 and usually others on the shield margin 

simple 14 

All setae on the dorsal shield margins, including all the s-S series, pilose (Fig. 15C). 19 

14 Dorsal setae generally long, curved or wavy, setae 7.4 reaching beyond bases of setae J5, setae j4, 
z2, z4, s2, r2, r3 pilose (Fig. 9); ventrianal shield subcircular, slightly truncate anteriorly, 
ornamented with lines and punctures . M. rotundiscutis Bregetova & Koroleva (p. 88) 
Dorsal setae generally short, straight or slightly curved; distal ends of setae 7.4 fall well short of 
bases of setae J5 (Fig. 16A); ventrianal shield usually pentagonal, truncate anteriorly (PI. IB), if 
strongly rounded then densely covered with minute punctations (PI. 1 D) and with only setaey'7 and 

r3 pilose on anterior part of dorsal shield (Fig. 8G) 1 5 

1 5 Setaey'7 elongate, over twice as long as setaey'2 and noticeably longer than setaey'J, minutely pilose 


MACROCHELIDAE OF THE BRITISH ISLES 


83 


D 


Plate 3 Sternal, genital and ventrianal shields of the females of: A Macrocheles decoloratus (C. L. 
Koch); B M. matrius (Hull); C Glyptholaspis confusa (Foa); D Dissoloncha superbus (Hull). After 
Evans & Browning (1956). 


84 


K. H. HYATT & R. M. EMBERSON 


D 


Plate 4 Sternal, genital and ventrianal shields of the females of: A Geholaspis (G.) longispinosus 
(Kramer); B Macrocheles opacus (C. L. Koch); C M. dentatus Evans & Browning; D Holostaspella 
orngta (Berlese). After Evans & Browning (1956). 


MACROCHELIDAE OF THE BRITISH ISLES 85 

distally (Fig. 1 6B), setae J5 simple, about half as long as setae Z5 (Fig. 1 6A); sternal shield without 
distinct lines but with a more or less symmetrical pattern of punctures some of which may be 

linearly arranged (PI. IB) M. robustulus (Berlese) (p. 106) 

Setae jl short and stout, little longer than setae j2 and/?, distinctly pilose in distal third to half, 
setae J5 simple or pilose, as long as or little shorter than setae Z5 (Fig. 8D); sternal shield with a 
distinct pattern of lines (PI. ID) 16 

16 All sclerotised regions densely covered with minute punctations (Fig. 8D); ventrianal shield 
strongly rounded, almost subcircular in outline (PI. ID). 

M. punctoscutatus Evans & Browning (p. 88) 
Sclerotised regions not densely punctate as above; ventrianal shield pentagonal in outline (PI. 1C) 

glaber species group 1 7 

17 Posterior series of dorsal setae with five pairs (J5, Z4, Z5, S4, 55) of setae on the dorsal shield 
pilose, at least distally (Fig. 1 7E); sternal shield with linea arcuata faintly impressed impunctate, all 
sternal shield punctures minute (PI. 6D) . . M. nataliae Bregetova & Koroleva (p. 109) 
Posterior series of dorsal setae with only one pair of pilose setae (/5) on the dorsal shield (Fig. 
17A); sternal shield with several well developed punctures along linea arcuata (PI. 1C), sternal 
shield punctures of different sizes, some minute, some larger 18 

1 8 Linea arcuata on sternal shield more or less straight but having its ends directed posteriorly, all 
lines on sternal shield well developed, not strongly punctured (PI. 1C) 

. . . M. glaber (Mutter) (p. 107) 

Linea arcuata strongly concave, having its ends directed anteriorly, lines on sternal shield not 
strongly impressed, punctures well developed (PI. IE). M. scutatus (Berlese) (p. 1 10) 

19 Dorsal setae only pilose in their distal third; setae J5 pilose (Fig. 15C); lateral elements of gnatho- 

tectum free (Fig. 15D) M. muscaedomesticae (Scopoli) (p. 105) 

Dorsal setae pilose in their distal half to two thirds, setae J5 simple (Fig. 7C); lateral elements of 
gnathotectum fused basally (Fig. 7D) .... M . penicilliger (Berlese) (p. 86) 

20 Dorsal setae pilose along their entire length, with an extra unpaired median seta present in the J2 
position between j6 and J3 (Fig. 14E); sternal shield densely covered with punctures (PI. 6B) 

M. punctatissimus Berlese (opacus species group) (p. 101) 

Dorsal setae only pilose in their distal halves, setae J2 paired, setae J3 absent, i.e. with 28 pairs of 
setae (Fig. 8 A); sternal shield with a pattern of lines and small areas of punctures (PI. 3 A) . . 21 

2 1 Setae J5 approximately equal in length to setae Z5 and little more than half as long as setae 55 (Fig. 
8 A); outer margins of lateral elements of gnathotectum smooth (Fig. 8B) 

M. decoloratus (C. L. Koch) (p. 88) 

Setae J5 shorter than setae Z5, setae Z5 approximately equal in length to setae 55 (Fig. 7A); outer 
margins of lateral elements of gnathotectum serrated (Fig. 7B) . M. matrius (Hull) (p. 85) 


Macrocheles matrius (Hull) 
(Fig. 7A, B, PI. 3B) 

The description of Evans & Browning (1956) requires no amendment. 

TYPE MATERIAL. Lectotype 9 'n.v.' [? Ninebanks vicarage, Northumberland], poultry manure, Hull 
collection. Here designated. Paralectotypes 18 $$, same data as lectotype [BMNH]. 

MATERIAL EXAMINED. 1 1 collections 4 <$<$, 194 ??. 

ENGLAND: Gloucestershire, Surrey, Cambridgeshire (Huntingdonshire), Worcestershire, 
Staffordshire, Derbyshire, Northumberland. 

SCOTLAND: Grampian (Aberdeenshire), Inner Hebrides (Pabay). 

HABITATS. Four collections associated with poultry, also with mink Mustela vison, water shrews 
Neomysfodiens, in a canary cage and from grass clippings and compost. 

DISTRIBUTION. Widespread in Europe; Germany (Krantz, 1972), Austria (Johnston, 1970), 
Bulgaria (Balogh, 1958), U.S.S.R. (Bregetova & Koroleva, 1960). Also known from the U.S.A. 
(Axtell, 1963), Israel (Costa, 19666) and New Zealand (Emberson, 1973a). 


86 


K. H. HYATT & R. M. EMBERSON 


Fig. 7 Macrocheles matrius (Hull): female A dorsal shield; B gnathotectum. Macrocheles penicilliger 
(Berlese): female C dorsal shield; D gnathotectum; E chelicera. After Evans & Browning (1956). 


Macrocheles penicilliger (Berlese) 
(Fig. 7C-E, PI. 2D) 

The description of Evans & Browning (1956) requires no amendment. 

TYPE MATERIAL. S\ nt\ pes 2 9? only, Italy, Cison di Valmarino, Treviso, sotto legni [under wood] 
slide 3/38 [ISZA]. 


MACROCHELIDAE OF THE BRITISH ISLES 


87 


Fig. 8 Macrocheles decoloratus (C. L. Koch): female A dorsal shield; B gnathotectum; C chelicera. 
Macrocheles punctoscutatus Evans & Browning: female D dorsal shield; E anterior margin of dorsal 
shield; F seta s2; G seta r3. After Evans & Browning (1956). 


88 K. H. HYATT & R. M. EMBERSON 

MATERIAL EXAMINED. 13 collections 3 DNN, 1 <$, 125 ??. 

ENGLAND: Cornwall, Surrey, Kent, London, Middlesex, Essex, Bucks, Oxfordshire, Cheshire. 
WALES: Gwynedd (Caernarvonshire). 
IRELAND: Clare. 
CHANNEL ISLANDS: Jersey. 

HABITATS. Three collections associated with Trox scaber (L.), also found in insect cultures, decay- 
ing leaves, compost and sewage culture. 

DISTRIBUTION. Apparently widespread in Europe; Germany (Karg, 1970), U.S.S.R. (Bregetova & 
Koroleva, 1960), also known from New Zealand (Emberson, 1973<z). 

Macrocheles decolor atus (C. L. Koch) 
(Fig. 8A-C, PI. 3A) 

Evans & Browning's (1956) description and synonymy remain unchanged 

MATERIAL EXAMINED. 8 collections 1 g, 60 $?. 
ENGLAND: Gloucestershire, Kent. 

SCOTLAND: Tayside (Perthshire), Fife, Highland (Inverness-shire). 
IRELAND: Cork. 

HABITATS. Six out of eight collections are from sand martin Riparia riparia nests, also collected 
from cow dung and moss and known from small mammal nests in U.S.S.R. 

DISTRIBUTION. Widespread in the British Isles and Europe; Germany (Karg, 1970), U.S.S.R., 
(Bregetova & Koroleva, 1960). 

Macrocheles punctoscutatm Evans & Browning 
(Fig. 8D-G,P1. ID) 

Macrocheles punctoscutatus Evans & Browning, 1956. Bull. Br. Mus. nat. Hist. (Zool.) 4: 18. 

The description of this species remains adequate. Bregetova & Koroleva (1960) have published a 
description of a male reputed to be this species. It appears to be very different from the female 
having only setae J5 on the dorsal shield pilose, also the complete covering of fine punctures 
appears to be lacking. The male has a holoventral shield, leg II is armed with a strong spur on the 
femur, small tubercles are present on other segments and on leg IV. 

TYPE MATERIAL, llolotypc ?, England, Glos., Churcham, from nest of mole Talpa europaea, 
16.V.1954. R. S. George. [BMNH]. 

MATERIAL EXAMINED. 4 collections 12 9?. 
ENGLAND: Gloucestershire, Wiltshire, Oxfordshire. 
SCOTLAND: Inner Hebrides (Eigg). 

HABITATS. Associated with small mammals; three collections from nests of moles Talpa europaea, 
one collection from Microtus agrestis. Bregetova & Koroleva (1960) record this species from a 
variety of small mammals. 

DISTRIBUTION. Probably throughout the British Isles. Widespread in U.S.S.R. (Bregetova & 
Koroleva, 1960), Germany (Karg, 1970). 

Macrocheles rotundiscutis Bregetova & Koroleva 
(Fig. 9, PI. 6A) 

Macrocheles rotundiscutis Bregetova & Koroleva, 1960. Parazit. Sb. 19: 1 16. 
Macrocheles bombophilus Gotz, in Krauss, 1970. Acarologie 14: 28, Syn. nov. 

FEMALE. The dorsal shield has 28 pairs of setae, most of which are long and curved or slightly wavy 
(Fig. 9). SetaeyV are short, broad and pilose; eight pairs of setae (j4, z2, z4, s2, r2, J5, Z5, S5) are 
long and strongly pilose over most of their length, setae r3 are faintly pilose and the other setae are 
simple. The setae on the dorsal shield are much longer than those off the shield. The dorsal shield 


MACROCHELIDAE OF THE BRITISH ISLES 


89 


Fig. 9 Macrocheles rotundiscutis Bregetova & Koroleva: female dorsal shield. 


is ornamented with reticulations and fine dense punctation, the lateral margins are irregularly 
serrate. 

With the exception of the postanal setae, the setae on the ventral shields are simple. The sternal 
shield has a characteristic reticulate pattern (PI. 6A) of ridges and punctures within the reticu- 
lations. The metasternal plates are rounded ovate. The genital shield is well marked with fine ridges 
and punctures. The ventrianal shield is broader than long and subcircular in outline, truncate 
anteriorly and only slightly angled laterally. The ornamentation is characteristic, consisting of 
curved transverse lines with a broad band of punctuation along the posterior edge of each line, and 
the punctures in each band decreasing in size posteriorly. The spermathecal apparatus has the 
normal two-lobed sacculus with the lobes widely separated, the corniculum is small and narrow. 

The gnathotectum has free lateral processes, the bifurcate median process is spiculate basally. 
The fixed chela is bidentate and the movable chela tridentate. The cheliceral dorsal seta is simple or 
faintly serrate distally. 

The setae on coxae III and IV are pilose; those on coxa, tibia and tarsus I are simple; other leg 
segments have a mixture of pilose and simple setae. 

MALE. The male is unknown in the British Isles and was not known to Bregetova & Koroleva 
(1960). However, Gotz (in Krauss, 1970), described the male from Germany (as M. bombophilus). 
The distribution of pilose dorsal setae is similar to the female. There is a holoventral shield. The 
fixed chela is bidentate and the movable chela unidentate; the spermatodactyl is about one-and-a- 
half times the length of the movable chela and has an anteriorly projecting basal tubercle. Leg II is 
armed with a spur on the femur and leg IV with tubercles and ridges on the trochanter and femur. 


90 K. H. HYATT & R. M. EMBERSON 

TYPE MATERIAL. M. rotundiscutis: 1099 syntypes, Lithuanian S.S.R., Ventes-ragas, litter under 
leaves in cemetery, 27.vii. 1957, Ivanov [ZINL]. 

The type material of M. bombophilus was not available for study, but there can be little doubt of 
the synonymy based on the description and drawings of Gotz (in Krauss, 1970). 

MATERIAL EXAMINED. 1 collection 1 9- 
ENGLAND: Surrey. 

HABITAT. A single female in leaf litter at Kew Gardens, Surrey is the first British record. Bregetova 
& Koroleva (1960) record this species from leaf litter, hay and compost, and Gotz (in Krauss, 1970) 
records it from bumble-bee nests. 

DISTRIBUTION. Germany (Gotz in Krauss, 1970); U.S.S.R., Leningrad region, Lithuanian S.S.R. 
and Caucasus (Bregetova & Koroleva, 1960). 

REMARKS. There are slight variations between the British specimen and the Lithuanian syntypes, 
the sternal shield pattern is slightly more complex and the punctate bands on the ventrianal shield 
wider in the British specimen. Setae J5 are longer than shown in Bregetova & Koroleva's drawing, 
both in the British and Lithuanian material examined, almost reaching the posterior margin of the 
dorsal shield. Also setae z2 are pilose in all the material examined and are shown in Bregetova & 
Koroleva's drawing, but said to be simple by Gotz. 

car 'mat ua group 

Krantz (1972) intimated that M. carinatus and other similar species formed a species group, 
although the existence of a group of very similar species was implicitly recognised by Bregetova & 
Koroleva (1960) and others. The members are medium to large sized (females 1000-1500 urn in 
length) heavily sclerotized, litter-dwelling species ofMacrocheles. The dorsal shield has 28-29 pairs 
of setae, J3 being present in addition to J2 in some species. Most of the dorsal setae are pilose in 
their distal half to two thirds. There is always a group of more or less simple setae in the middle of 
the dorsal shield that includesytf, z5, z6, J2 and J3 if present. In addition other setae, includingy'2,/5, 
zl, s2, r3, r4 and J5, are simple in some species. The dorsal shield is generally ornamented with 
a reticulate pattern and sometimes with a punctate microsculpture. The posterior and lateral 
margins may be irregularly crenulate or smooth. The setae on the ventral shields may be simple or 
pilose. The sternal shield pattern consists of a reticulate-punctate network which is more pro- 
nounced and more punctate in the posterior third. The metasternal plates are small and oval with 
both seta and pore. The genital shield is ornamented with a punctate lineal pattern. It is generally 
rounded posteriorly and the associated pores are free on the membrane. The ventrianal shield is 
reduced, subtriangular or rounded and has a pattern of punctate lines. There are three pairs 
of free postepigynal apodemes in the membranous cuticle. Males have separate sternogenital and 
ventrianal shields. The gnathotectum has the lateral processes fused and the stem of the median 
process strongly spiculate. The dentition of the chelae varies from species to species, but the 
spermatodactyl is short, about half the length of the movable chela, strongly tapered and directed 
postero-dorsally; the dorsal seta is pectinate latero-distally. The leg setation is normal for the genus 
and is a mixture of simple and pilose setae. In the male leg II is armed with a simple spur on the 
femur and small tubercles on the genu and tibia. The spermathecal structures are indistinct. 

INCLUDED SPECIES: M . carinatus (C. L. Koch), M. montanus (Willmann), M. submotus Falconer, M. 
tardus (C. L. Koch). 

Macrocheles carinatus (C. L. Koch) 
(Fig. 10A-D,P1.2C) 

The description and synonymy given by Evans & Browning (1956) need no amendment. 

MATERIAL EXAMINED. 53 collections several DNN, 49 dd, 90 99- 

ENGLAND: Isles of Scilly, Cornwall, Somerset, Dorset, Surrey, London, Kent, Hertfordshire, 
Berkshire, Buckinghamshire, Cambridgeshire, Leicestershire, Nottinghamshire, Yorkshire, 
Cumbria (Westmorland), Northumberland. 


MACROCHELIDAE OF THE BRITISH ISLES 


91 


B 


Fig. 10 Macrocheles carinatus (C. L. Koch): female A anterior region of dorsal shield; B postero- 
lateral margin of dorsal shield; C gnathotectum; D chelicera. Macrocheles montanus (Willmann): 
female E dorsal shield. After Evans & Browning (1956). 


92 K. H. HYATT & R. M. EMBERSON 

SCOTLAND: Tayside (Perthshire), Inner Hebrides (Mull), Shetland. 

WALES: Dyfed (Pembrokeshire), Glamorgan, Gwent, Gwynedd (Caernarvonshire). 

IRELAND: Clare, Westmeath, Leitrim. 

HABITATS. A variety of litter habitats including flood debris, dead grass, seaweed, moss, etc., also 
from Microtus agrestis. Probably mainly found in damp places. 

DISTRIBUTION. Throughout the British Isles, widespread in Europe, recent records include: 
U.S.S.R. (Bregetova & Koroleva, 1960), Germany (Krantz, 1972), Austria (Johnston, 1970), 
Bulgaria (Balogh, 1958). 

Macrocheles montanus (Willmann) 
(Fig. 10E,P1.2B) 

The description and synonymy given by Evans & Browning (1956) remain unaltered. 

MATERIAL EXAMINED. 40 collections 1 ^, 82 99- 

ENGLAND: Cornwall, Somerset, Dorset, Sussex, Surrey, London, Kent, Hertfordshire, 
Berkshire, Buckinghamshire, Cambridgeshire (including Huntingdonshire), Northamptonshire, 
Lancashire, Durham, Northumberland, Cumbria (Cumberland). 

SCOTLAND: Tayside (Angus, Perthshire), Inner Hebrides (Mull). 

WALES: Gwent. 

IRELAND: Westmeath. 

HABITATS. Mainly leaf litter, also cow dung and under wet oak bark. Three collections from small 
mammals (Sorex araneus and Microtus agrestis). 

DISTRIBUTION. Similar to M. carinatus, recent records include U.S.S.R. (Bregetova & Koroleva, 
1960), and Germany (Krantz, 1972). 

REMARKS. A very similar species, also with setae J3 present, has been described by Bregetova & 
Koroleva ( 1 960) from the Caucasus region of the U.S.S.R. One of us (R.M.E.), having examined the 
types of M. montivagus Berlese, follows Krantz (1972) in rejecting the synonymy of M. montanus 
with that species as proposed by Krauss (1970). 

Macrocheles submotus Falconer 
(Fig. 11A-D,P1. 2E) 

The description of this species in Evans & Browning (1956) is satisfactory, except thaty'3 (D3) and 
z2 (M2) are usually less pilose than described and figured. 

It is evident that Falconer had several collections before him when he described this species as M. 
cognatus, a junior homonym of M. cognatus Berlese 1918. He states (Falconer, 1923: 152) 'Found 
amongst dead leaves, sphagnum and heaps of cut grass in a field at Slaithwaite, first examples [our 
italics] May, 1919'. He also mentions both males and females. No type was designated. 

The only material of these collections now known to exist is two females and two males labelled 
'2nd lot' from the Falconer collection and now housed in the Arachnida collection of the British 
Museum (Natural History). 

TYPE MATERIAL. Lectotype 9, here designated, [England, ? Slaithwaite, Yorkshire] '2nd lot', 
Falconer coll. [BMNH]. Paralectotypes, 1 9, 2 3d, same data as lectotype, [BMNH]. 

MATERIAL EXAMINED. 122 collections c. 26 PNN, 44 DNN, 134 <^, 225 99. 

ENGLAND: Devon, Somerset, Gloucestershire, Dorset, Hampshire, Surrey, Sussex, Kent, 
London, Essex, Middlesex, Berkshire, Leicestershire, Lancashire, Yorkshire, Durham, 
Northumberland, Cumbria (Westmorland). 

SCOTLAND: Strathclyde (Argyllshire), Tayside (Perthshire), Dumfries & Galloway 
(Wigtownshire), Highland (Inverness-shire, Ross & Cromarty), Inner Hebrides (Skye, Scalpay, 
Mull, Ulva), Shetland. 


MACROCHELIDAE OF THE BRITISH ISLES 


93 


H 


Fig. 1 1 Macrocheles submotus Falconer: female A dorsal shield; B gnathotectum; C chelicera: male 
D chelicera. Macrocheles tardus (C. L. Koch): female E anterior region of dorsal shield; F postero- 
lateral margin of dorsal shield; G gnathotectum; H chelicera. After Evans & Browning (1956). 


94 K. H. HYATT & R. M. EMBERSON 

WALES: Glamorgan, Gwent, Gwynedd (Merionethshire, Caernarvonshire), Clwyd 
(Denbighshire). 

IRELAND: Clare, Mayo, Galway, Sligo, Leitrim, Westmeath. 
CHANNEL ISLANDS: Jersey. 

HABITATS. A wide variety of leaf litter habitats, also mosses; some evidence of association with 
damp habitats, i.e. Sphagnum, flood debris, saltwater marsh, etc. Twelve collections from small 
mammals (Sorex minutus, S. araneus and Microtus agrestis). 

DISTRIBUTION. Common throughout the British Isles. No positive records for this species outside 
the British Isles are known to the authors. The records by Halbert (191 5) of Holostaspis tridentinus 
G. & R. Can. refer to this species, but other records, where it has been possible to check, refer to 
other species of the carinatus group. 

REMARKS. As mentioned above, the correct name for this species has been very confused in the 
European literature. The situation has not been helped by Krauss (1970) who synonymizes M. 
submotus Falconer and M. tardus, sensu Evans & Browning, with M. tridentinus (G. & R. 
Canestrini). Whatever decision is made as to the status of M. tardus (Koch) the fact remains that 
M. submotus is a very different species from that described and illustrated by Krauss (1970) as 
M. tridentinus. 

Macrocheles tardus (C. L. Koch) 
(Fig. 11E-H,P1.2F) 

Gamasus tardus C. L. Koch, 1841. Deutschlands Crustaceen, Myriapoden und Arachniden. Regensburg, Heft 

39, no. 14. 
Holostaspis tridentinus G. & R. Canestrini, 1882. Atti Soc. ven.-trent. sci. 8: 28. 

The treatment of this species by Evans & Browning (1956) remains adequate. 

TYPE MATERIAL. M. tridentinus: Neotype <$, here designated. Labelled as follows: yellow-bordered 
printed label, Coll. Berlese. Macrocheles tmdentinus (G. & R. Can.), $ Tiarno (Trentino) musco 
[moss], slide 187/43 [ISZA]. 

MATERIAL EXAMINED. 1 8 collections 9 <$<$, 29 ??. 

ENGLAND: Gloucestershire, Cambridgeshire (including Huntingdonshire), Yorkshire, Cumbria 
(Westmorland). 

WALES: Glamorgan. 

IRELAND: Leitrim, Westmeath. 

HABITAT. A variety of habitats with a tendency towards damp situations, i.e. flood debris, moss, 
peat and Molinia tussocks in the fens. 

DISTRIBUTION. A widespread species in the British Isles and Europe. Recent records include: 
U.S.S.R. (Bregetova & Koroleva, 1960), Germany (Krantz, 1972). 

REMARKS. The present authors agree with Krantz (1972) that the name M. tardus Koch is best used 
for this species rather than for species of Glyptholaspis as proposed by Krauss (1970). 

The identity of M. tridentinus has long remained a problem because of the inadequate original 
description and figures (G. & R. Canestrini, 1882). One of us (R.M.E.) has recently had the 
opportunity to examine parts of the Canestrini Collection in Padova and hopes that the problem 
may here be resolved. 

The original description was based mainly on a male specimen, and, to a lesser extent, on what 
the Canestrini brothers referred to as juvenile females. These latter were probably specimens of an 
unrelated and now entirely unrecognisable species. No types were specifically designated but 
Trentino was mentioned as the locality from which the material came. In the Canestrini Collection 
there is a tube (jar no. XX, vial no. 556) labelled Holostaspis tridentinus Can., Trentino, which 
could be taken as containing the type series of M. tridentinus. This material was evidently seen by 
Valle (1955). Unfortunately this tube, together with several others examined, was empty. The 


MACROCHELIDAE OF THE BRITISH ISLES 95 

late Dr A. Filipponi also encountered this problem when he examined parts of the Canestrini 
Collection (Pegazzano, personal communication). 

The only other material in the Collection labelled as H. tridentinus is a badly damaged, over- 
cleared, transparent male of M. carinatus (jar no. XX, vial no. 604) from an unknown locality. 

H. tridentinus appears to be conspecific with either M. tardus or M. carinatus as understood here 
and interpreted by Evans & Browning (1956). For reasons given below synonymy with the former 
species is favoured. M. submotus can be almost totally discounted from consideration in spite 
of its frequent synonymisation with tridentinus on the grounds that no authentic material of 
M. submotus is known to exist in central or southern Europe. 

In the Berlese collection there are two specimens identified as M. tridentinus from Tiarno 
(Trentino) which can be considered topotypical. They date from Berlese's Padova period (slide 
labels with yellow borders) and may have been collected when Berlese was a student of G. 
Canestrini. These specimens are both M. tardus as here understood. One of these (187/43) has been 
selected as a neotype of M. tridentinus. 

The only discrepancy between the neotype and the original description of tridentinus is the length 
of the body. The following are measurements based on G. & R. Canestrinis' (1882) description, the 
neotype, and typical values for M. carinatus and also ratios of body length to the other values: 

Canestrini neotype typical values 

tridentinus tridentinus carinatus 

Body length urn 860 1050 870-910 

Body width um 640(1:0.74) 660(1:0.63) 500-540(1:0.58) 

LeglVlength um 1250(1:1.45) 1300(1:1.24) 980-1000(1:1.11) 

It is apparent that there is good agreement in body width and leg length with the neotype and that 
these measurements fall outside the normal range for M. carinatus. Also the ratios of body length 
to body width and body length to length of leg IV derived from the Canestrini measurements would 
be abnormal for this type of mite. 

The most likely explanation for these inconsistencies is that there is an error in the body length 
measurement in the original description, and that the Canestrinis had before them a specimen of 
M. tardus. For these reasons synonymy with M. tardus is favoured and a suitable neotype chosen. 

opacus group 

The opacus species group was distinguished by Johnston (1970) for a characteristic group of litter- 
and soil-dwelling Macrocheles. The dorsal shield has a basic complement of twenty-eight pairs of 
setae plus an additional one to four setae that may be paired or unpaired. Setae J3 are always 
present (absent in most other Macrocheles) and between setaey'6 and J3 there may be a single seta, a 
pair of setae or three or four unpaired setae. Setae jl are short and laterally expanded, other setae 
are usually long, tapered and pilose along their entire length; setae zl and J5 may be considerably 
shorter than the others. The dorsal shield usually tapers from the shoulders and often does not 
cover the entire dorsum. The lateral and posterior margins are often finely dentate or crenulate. 
The sternal shield is reticulate-punctate without a strong pattern of lines. The genital shield is 
usually reduced and rounded posteriorly. The ventrianal shield is reduced to a greater or lesser 
extent so that three pairs of apodemes are usually exposed; the number of ventrianal shield setae 
may be reduced to two, one, or no preanal pairs. Males have separate sternogenital and ventrianal 
shields, which may also be reduced. The lateral processes of the gnathotectum are usually reduced 
to narrow spikes and the median bifurcate process may be dissected distally. The fixed chela has an 
offset subterminal tooth and a central tooth, whilst the movable chela of the female has two central 
teeth; the male has a unidentate movable chela and a short, strongly tapered, posterodorsally 
directed spermatodactyl. The cheliceral dorsal seta is pectinate distally. The legs are highly rugose 
and have mainly pilose setae on most segments except coxa and tarsus I. The form of the spermath- 
ecal complex varies from the typical bilobed Macrocheles type through a stage with slightly 
enlarged rami and increasingly delicate membranous median organs to an extreme form with very 
elongate rami and indiscernible median organs similar to Geholaspis (Longicheles). 


96 K. H. HYATT & R. M. EMBERSON 

INCLUDED SPECIES. M. opacus (C. L. Koch), M. analis sp. nov., M. dentatus (Evans & Browning), 
M. punctatissimus Berlese, M. terreus (Canestrini & Fanzago). 

The inclusion of M. punctatissimus in this species group is provisional. The group includes a 
series of species increasingly specialised by reduction of sclerotization and gnathotectal form, of 
which M. punctatissimus appears to represent the least specialised end. It shows little evidence of 
reduction of sclerotization except that the ventrianal shield is rather small and rounded. However, 
the dorsal chaetotaxy is the same as other members of the group and the chelicerae are of the opacus 
type. 

Macrocheles opacus (C. L. Koch) 
(Fig. 12A-E,P1.4B) 

Macrholaspis opacus (C. L. Koch): Evans & Browning, 1956. Bull. Br. Mus. not. Hist. (Zool.) 4: 46. 
Macrocheles (M.) opacus var. aciculatus: Krauss, 1970. Acarologie 14: 23. 

Evans & Browning placed this species in the genus Macrholaspis Oudemans, mainly on account of 
it having only two pairs of preanal setae on the ventrianal shield and crenulate margins to the 
dorsal shield. Krantz (1962) showed that there are closely related species having three pairs of 
preanal ventrianal setae and synonymised Macrolaspis with Macrocheles. The present authors also 
follow Krantz ( 1 972) in synonymising M. opacus var. aciculatus, Krauss with M. opacus. 

MATERIAL EXAMINED. 47 collections 7 PNN, 17 DNN, 88 ??. 

ENGLAND: Dorset, Gloucestershire, Berkshire, Surrey, Kent, Hertfordshire, Suffolk, 
Warwickshire, Worcestershire, Cumbria (Westmorland), Northumberland. 

SCOTLAND: Strathclyde (Argyllshire), Dumfries & Galloway (Wigtownshire), Highland (Ross & 
Cromarty), Inner Hebrides (Mull, Ulva). 

WALES: Glamorgan, Gwent, Gwynedd (Caernarvonshire), Clwyd (Denbighshire). 

IRELAND: Clare, Galway, Mayo, Westmeath. 

HABITATS. Collected in a variety of litter and soil habitats, but mainly in moss. Also found in rotten 
wood and an ants' nest. 

DISTRIBUTION. Widespread in Europe, recent records include Germany (Krantz, 1973) and Spain 
(Athias-Henriot, 1968). 

Macrocheles analis sp. nov. 
(Fig. 13A-F,P1. 5B) 

FEMALE. The dorsal shield tapers from the shoulders posteriorly (Fig. 1 3 A). It is ornamented with a 
raised polygonal reticular network which breaks up into small blunt tubercles anteriorly. The 
ornamentation is more rugose towards the shoulders. Between the main ornamentation there is 
fine granular microsculpture. The edges of the shield are smooth. There are twenty-nine pairs of 
pilose setae on the dorsal shield including both J2 and J3. 

The shields of the ventral surface are weakly sclerotized and reduced in size. The sternal shield 
has a fine granular texture and distinct reticulations posteriorly (PI. 5B). The sternal and genital 
setae are pilose. The metasternal plates are small and round, only just large enough for the setae; 
the pores are separate in the unsclerotized membrane. The genital shield has a porous surface 
without reticulation. It is almost semicircular posteriorly and the genital pores are on the unscler- 
otized membrane. The postgenital apodemes are weakly sclerotized, barely discernible. The 
opisthogastric sclerotization is reduced to a small subcircular shield bearing the anal setae (Fig. 
1 3B). The adanal setae are simple, longer than the postanal which is pilose. The Jv setae are simple, 
the other opisthogastric setae are strongly pilose. The spermathecal apparatus has elongate rami 
and indistinct median organs (Fig. 13C). 

The gnathotectum has a bifid median process with the arms dissected and the lateral processes 
absent, the anterior edge is dentate (Fig. 13D). The fixed chela is basically bidentate with a central 
tooth and an offset subterminal tooth, it is truncate distally; the movable chela is bidentate (Fig. 
13E). The dorsal seta is pectinate laterodistally. The corniculi are elongate (Fig. 13F), one-and-a- 


MACROCHELIDAE OF THE BRITISH ISLES 


97 


Fig. 12 Macrocheles opacus (C. L. Koch): female A dorsal shield; B anterior region of dorsal shield; C 
lateral margin of dorsal shield; D gnathotectum; E chelicera. After Evans & Browning (1956). 


98 


K. H. HYATT & R. M. EMBERSON 


x 


Plate 5 Sternal shields of the females of: A Geholaspis (Longicheles) hortorum (Berlese); B Macro- 
cheles analis sp. nov.; C Holostaspella ornata (Berlese); D H. exornata Filipponi & Pegazzano; E 
Glyptholaspis fimicola (Berlese). 


k of 

?- 'I. 


B 


Plate 6 Sternal shields of the females of: A Macrocheles rotundiscutis Bregetova & Koroleva; B M. 
punctatissimus Berlese; C M. terreus (Canestrini & Fanzago); D M. nataliae Bregetova & Koroleva; E 
M. subbadius (Berlese); F Glyptholaspis americana (Berlese). 


100 


K. H. HYATT & R. M. EMBERSON 


Jv1 


Fig. 13 Macrocheles analis sp. n.: female A dorsal shield; B anal shield; C tubulus and ramus of 
spermathecal apparatus; D gnathotectum; E chelicera; F venter of gnathosoma. 


half times as long as the pedipalp trochanter. The internal hypostomal setae are posterior to the 
external hypostomal setae. The hypognathal groove has two rows of strong denticles (G5 and G6 in 
Hirschmann's notation) and three fine rows (G2, G3, G4). The hollow seta on the external part of 
the palp tarsus, which is elongate in all known species of Macrocheles, is barely longer than the 
surrounding setae. 

The legs have mainly pilose setae on all segments except coxa and tarsus I which have all setae 
simple. 

TYPE MATERIAL. Holotype ?, England, Berks, Silwood Park, 'light sandy loam with winter wheat', 
23.xi.1963. [BMNH]. 

MATERIAL EXAMINED. Known only from the holotype in the British Isles, but two specimens from 
the U.S.A., apparently referrable to this species, have been seen; 1 ?, Corvallis, Oregon, rotten bulb 
[OSUC] and 1 9, Lexington, Kentucky, duff. 

REMARKS. This species represents the end of a series of species showing increasing reduction of the 
ventral shields and increasing specialisation of the gnathosoma and spermathecal apparatus in the 
direction of the condition found in Geholaspis (Longicheles). It is probable that the reduction of 
sclerotization is an adaption to life in deeper soil layers. 


MACROCHELIDAE OF THE BRITISH ISLES 1 1 

Macrocheles dent at us (Evans & Browning) 
(Fig. 14A-D, PI. 4C) 

Macrholaspis dentatus Evans & Browning, 1956. Bull. Br. Mus. nat. Hist. (Zool.) 4: 49. 
Macrocheles ( M.) multisetosus Gotz in Krauss, 1970. Acarologie 14: 29. Syn. nov. 

Evans & Browning (1956) described this species in the genus Macrholaspis, which was 
synonymised by Krantz ( 1 962) with Macrocheles. 

The type material of M. multisetosus Gotz has not been examined, but the description and 
illustrations fail to show any significant difference between it and the holotype of M. dentatus, with 
the exception of four rather than three unpaired median dorsal setae. This character has been 
found to vary in material from collections in the British Isles and is not correlated with other 
characters. However, the original description of M. dentatus is misleading in that the setae on the 
membrane between the genital and ventrianal shields are stated to be plumose when in fact there 
are two pairs of simple setae, Jvl and Zvl ( VI and V5 in Hirschmann's notation), in this position in 
all specimens examined, including the holotype. 

In their key to species of Macrholaspis Evans & Browning (1956) state that there is a strong 
tubercle on the posterior margin of trochanter IV. This is an error for femur III. 

The Hull Collection includes a specimen labelled 'Macrocheles pannosus 9 Bks.' [Ninebanks, 
Northumberland], which is a specimen of M. dentatus. Reluctantly the present authors have 
concluded that this cannot be the type of M. pannosus Hull, 1925, described from West Allendale, 
as it does not agree with the description either as to size or in the characters described. 

TYPE MATERIAL. M. dentatus; holotype 9, Wales, Cardiganshire, Dolybont, Leri Valley, in litter 
under bracken, viii.1954, G. O. Evans. [BMNH]. 

MATERIAL EXAMINED. 19 collections 2 DNN, 22 99- 

ENGLAND: Cornwall, Devon, Somerset, Surrey, Kent, Cambridgeshire, Yorkshire, 
Northumberland. 

SCOTLAND: Tayside (Perthshire). 

WALES: Gwent, Dyfed (Cardiganshire). 

IRELAND: Clare, Gal way, Westmeath. 

HABITATS. Bryophytes, leaf litter and grassland soil, also one collection from Microtus agrestis. 

DISTRIBUTION. Evidently widespread but rarely collected in the British Isles; also known from 
Germany (Gotz in Krauss, 1970, as M. multisetosus). 

Macrocheles punctatissimus Berlese 
(Fig. 14E-G, PI. 6B) 

Macrocheles (Nothrholaspis) punctatissimus Berlese, 1918. Redial3: 171. 
Nothrholaspis pulcherrimus Willmann, 1951. Sber. Akad. Wiss. Wien. 160: 104. 

FEMALE. The dorsal shield (Fig. 14E) completely covers the dorsal surface. It is strongly orna- 
mented with reticulations, the sides of the reticulations are crenate and within them there is a 
porose microsculpture. There are twenty-eight pairs of setae plus an unpaired median seta between 
j6 and J3 on the dorsal shield. Setae// are slightly flattened, all other setae are of the usual tapering 
pilose variety. The lateral margins of the dorsal shield are smooth. 

The sternal shield is strongly punctured all over; the punctures are arranged in groups to form a 
symmetrical pattern (PI. 6B). The metasternal plates are rounded and include the pore. The genital 
shield is truncate posteriorly and not noticeably reduced. It is ornamented with a series of minute 
tubercles arranged in rows. The sternal and genital setae are simple. The ventrianal shield is 
rounded and closely abuts the genital, it bears three pairs of simple preanal setae. The postanal seta 
is pilose. The spermathecal complex is of the normal type within the genus Macrocheles with short 
rami joining a two-lobed sacculus (Fig. 14F). 

The gnathosomal structures are typical of the genus Macrocheles. The gnathotectum has a 
median bifid process (Fig. 14G) and free lateral processes expanded and serrate at the distal end. 


102 


K. H. HYATT & R. M. EMBERSON 


B 


f- I \ \ 


Fig. 14 Macrocheles dentatus Evans & Browning: female A dorsal shield; B anterior region of dorsal 
shield; C lateral margin of dorsal shield; D dorsal seta z4. After Evans & Browning (1956). Macro- 
cheles punctatissimus Berlese: female E dorsal shield; F spermathecal apparatus; G gnathotectum. 


MACROCHELIDAE OF THE BRITISH ISLES 1 03 

The fixed chela is basically bidentate, including an offset terminal tooth. The movable chela is 
bidentate and the dorsal seta is slightly pectinate distally. The hypognathal groove has the usual 
five rows of denticles and hypostomal setae 2 and 3 are opposite each other. 

The legs are normal for the species group, rugose and with the usual mixture of pilose and simple 
setae on most segments. Setap/j on femur III is conspicuous and blade shaped. 

TYPE MATERIAL. Macrocheles punctatissimus: Syntypes 2 $9 'Castions distruda (Udine), musco 
[moss]', slide 13 1/46 [ISZA]. 

MATERIAL EXAMINED. 2 collections 3 $$. 
ENGLAND: Cambridgeshire. 

HABITATS. The first British record. From the bases ofMolinia tussocks in Wicken Fen. Apparently 
found in similar places in Austria (as pulcherrimus). 

DISTRIBUTION. Germany (Krauss, 1970), Austria (Willmann, 1951 as pulcherrimus). 

REMARKS. No type material of Nothrholaspis pulcherrimus has been examined so the synonymy 
follows Krauss (1970). The species present in Britain is clearly conspecific with that described by 
Krauss (1970) and probably with that described by Willmann (1951). 

Macrocheles terreus (Canestrini & Fanzago) 
(Fig. 15A-B,P1. 6C) 

Gamasus terreus Canestrini & Fanzago 1877. Atti R. 1st. veneto Sci. (V) 4: 48. 

Holostaspis echinatus Berlese, 1904. Redia 2: 20. 

Holostaspis terreus (Canestrini & Fanzago), Halbert, 1915. Proc. R. Ir. Acad. 31: 66. 

Macrocheles (M.) opacus Krauss, 1970. Acarologie 14: 22. 

Macrocheles terreus (Canestrini & Fanzago); Krantz, 1972. Ent. Mitt. zool. Mus. Hamburgh: 270. 

FEMALE. The dorsal shield has twenty-nine pairs of setae (Fig. 15A) including pairs in both the J2 
and J3 positions. The setae are generally shorter and blunter than in other members of the species 
group and pilose or plumose almost to the base; setae// are strongly flattened, almost fan-shaped. 
The dorsal shield ornamentation consists of faint reticulations and a dense microsculpture of fine 
pores. The lateral margins are bluntly dentate. 

All the ventral shields are finely punctured, many of the punctures are arranged in lines. The 
setae on the ventral shields are basically simple, except the genital setae, which may appear finely 
pilose, and the pilose post anal seta. The sternal shield has a symmetrical pattern of punctate areas 
and lines of fine punctures. The metasternal platelets are elongate oval. The genital shield is semi- 
circular behind and the associated pores are free on the membrane. The subcircular ventrianal 
shield is well separated from the genital with three pairs of apodomes between. The median 
structures of the spermathecal complex are indistinct. 

The gnathosoma is typical of the species group with free spike-like lateral processes on the 
gnathotectum and the chelae with offset subterminal and central teeth on the fixed chela and two 
central teeth on the movable chela (Fig. 15B). The hypognathal groove has six rows of denticles. 

The legs are normal for the species group except that many setae are somewhat flattened or 
blade-like instead of pilose. 

TYPE MATERIAL. H. echinatus: Lectotype ? (here designated), Italy, Firenze, Boboli, Foglia marce 
palme, 'tipico M. echinatus Berl.' slide 22/38 [ISZA]. G. terreus: Neotype 9 (here designated), the 
lectotype of//, echinatus, from Firenze, Italy, slide 22/38 [ISZA]. 

Although Berlese indicated a type designation on slide 22/38 and in his notebooks, this desig- 
nation was not published. 

MATERIAL EXAMINED. 6 collections 6 ??. 

ENGLAND: Hampshire, Kent, Suffolk, North Yorkshire, Humberside. 
IRELAND: Mayo (Halbert, 1915). 

HABITAT. From arable and grassland soils and beech forest litter, also recorded from moss and leaf 
litter by Krauss (1970) and from moss and under bark by Halbert (1915). 


104 


K. H. HYATT & R. M. EMBERSON 


Fig. 15 Macrocheles terreus (Canestrini & Fanzago): female A dorsal shield with right seta S4 
enlarged; B chelicera. Macrocheles muscaedomesticae (Scopoli): female C dorsal shield; D 
gnathotectum; E chelicera. After Evans & Browning (1956). 


MACROCHELIDAE OF THE BRITISH ISLES 1 05 

DISTRIBUTION. According to Krauss (1970) it is known from Germany and Spain (as opacus). Since 
Krauss's concept of M. opacus varies from that of most other authors, the literature records he cites 
are probably for several species. 

REMARKS. The original material of Gamasus terreus was stated (Canestrini & Fanzago, 1877) to 
have come from Padova. The type material is not present in the Canestrini Collection in Padova 
(see remarks under M. tardus) and must be presumed lost. 

Valle (1955) records two vials listed under terreus in the catalogue of the Canestrini Collection. 
The first vial (jar no. XX, vial no. 703) apparently contained specimens of terreus from Rome, when 
seen by Valle, but now only contains two feather-mites. The other vial (jar no. XIII, vial no. 504) 
contained Geholaspis longispinosus when seen by Valle, but is now empty. It is labelled 'Messina'. 
Neither of these vials could have contained the types as these came from Padova. 

In the absence of Canestrini and Fanzago's type material of G. terreus a neotype has been 
selected, this is necessary in view of the likelihood of there being a complex of closely similar species 
centred on M. terreus. 

The lectotype specimen of echinatus has been chosen as neotype of terreus. This will have the 
effect of making echinatus an objective synonym of terreus and confirming a synonymy indicated 
by Berlese on the slide of echinatus and in his notebooks. 

Berlese's (1904) original drawing of echinatus (Plate II, Fig. 33) differs from his later drawing in 
his notebook and from the designated type of echinatus in having a truncate posterior margin to the 
genital shield and in having a larger, more triangular ventrianal shield. 

Macrocheles terreus evidently belongs to a complex of very closely related species within the 
opacus species group of which M. beieri Johnston from Austria may be a member. It is possible that 
the species described above may also represent a different taxon based on the simple setae of the 
ventral shields, the six rather than five rows of hypognathal denticles and fine differences in the 
sculpture of the shields. However, a decision on this must await the discovery of more British 
material and direct comparison of the various forms involved. 

The present authors have followed Krantz (1972) in the synonymy of M. opacus, Krauss, with 
M. terreus (Canestrini & Fanzago). 

Macrocheles muscaedomesticae (Scopoli) 
(Fig. 15C-E,P1. 1A) 

The description and synonymy of this species as given by Evans & Browning (1956) remain correct. 

MATERIAL EXAMINED. 27 collections 76 99, see below. 

ENGLAND: Cornwall, Devon, Wiltshire, Hampshire, Sussex, Surrey, Kent, London, Essex, 
Middlesex, Hertfordshire, Cambridgeshire, Lancashire, Cheshire. 

A sample from a deep-litter house in which partridges Perdix perdix were being reared in 
Cambridgeshire in 1978 contained thousands of females of this species. 

HABITATS. Mostly associated with synanthropic flies, i.e. Musca domestica, Fannia spp., Stomoxys 
calcitrans, also with fly breeding places, compost heaps, rotting seaweed and artificial nests of rats. 

DISTRIBUTION. Apparently cosmopolitan wherever suitable phoretic hosts exist, although surpris- 
ingly, the only British material examined is from England. 

Macrocheles pisentii (Berlese) 

This species almost certainly does not occur in the British Isles. Its inclusion by Evans & Browning 
(1956) was based on three records of Hull (1918), but they were unable to examine his specimens. 
The present authors have examined material labelled as M. pisentii in the Hull Collection, but 
without locality data, and found it to be M. glaber. 

Costa (1967) has shown M. pisentii to be very closely associated with Scarabaeus semipunctatus 
F., even to the exclusion of other species of Scarabaeus. Since 5. semipunctatus is not present in the 
British fauna it seems that M. pisentii is also unlikely to be present. 

Costa (1967) redescribed M. pisentii and distinguished it from other closely related species. 


106 


K. H. HYATT & R. M. EMBERSON 


Fig. 16 Macrocheles robustulus (Berlese): female A dorsal shield; B setay'7; C chelicera. After Evans & 

Browning (1956). 


Macrocheles robustulus (Berlese) 
(Fig. 16A-C,P1. IB) 

Holostaspis subbadius var. robustulus Berlese, 1904. Redia 1: 264. 

Holos t aspis humeratus Berlese 1908. Redia 5: 13. 

NothrholaspispunctillatusWi\\mann, 1939. Abh. naturw. Ver. Bremen3\: 176. 

Macrocheles coprophila Womersley, 1942. Trans. Roy. Soc. S. Aust. 66: 167. 

Macrocheles rothamstedensis Evans & Browning, 1956. Bull. Br. Mus. nat. Hist. (Zool.) 4: 15. 

Macrocheles robustulus: Axtell, 1961. Ann. ent. Soc. Am. 54: 748. 


MACROCHELIDAE OF THE BRITISH ISLES 1 07 

The first British record of this species was that of Evans & Browning (1956) who described it 
under the name M. rothamstedensis. This was synonymised with M. punctillatus (Willmann) by 
Bregetova & Koroleva (1960) and with M. robustulus (Berlese) by Axtell (1961). For the complete 
synonymy see Krantz & Filipponi (1964). 

Evans & Browning's (1956) description needs no amendment. 

MATERIAL EXAMINED. 3 collections 3 <$<$, 64 $?. 
ENGLAND: Essex, Hertfordshire, Bedfordshire. 

HABITATS. Cattle dung, 'colony on cucumber leaves'. Elsewhere this species is most often collected 
from cattle manure (Axtell, 1961), and phoretic on scarabaeine beetles (Costa, 1966a). 

DISTRIBUTION. Widespread in temperate regions of the world; Italy (Filipponi & Pegazzano, 1962), 
Germany (Krantz, 1972), U.S.S.R. (Bregetova & Koroleva, 1956), Israel (Costa, 1966<z), U.S.A. 
(Axtell, 1961), Australia (Krantz & Filipponi, 1964), New Zealand (Emberson, unpublished). 

glaber group 

This species group was defined by Filipponi & Pegazzano (1962) and is here slightly modified to 
include another obviously related species. The species are medium sized with the 28 pairs of setae 
on the dorsal shield being mainly simple, but there are from four to ten pairs that are pilose. These 
usually include jl, j4, z3 which are pilose distally and J5 pilose over their entire length. Males 
usually have additional pilose setae. The dorsal shield is lightly ornamented with transverse reticu- 
lations and, in the females, is divided by a procurved line passing close to the bases of setae z6 and 
extending laterally towards setae r4 (Fig. 17A). The lateral margins are smooth. The male dorsal 
shield is strongly tapered posteriorly with crenulate lateral and posterior margins. The type of 
sternal shield pattern is characteristic, with lines often edged with punctures. The linea media 
transversa is well developed and straight, the linea angulata is similar in all species and there are one 
or two lineae arcuatae of varying form. Behind the linea media transversa the lineae obliquae 
posteriores are bifurcate distally and the areae punctatae posteriores are well developed. The 
ventrianal shield is usually more or less pentagonal and ornamented with concentric lines and 
reticulations, at least anteriorly. The males have separate sternogenital and ventrianal shields. All 
the ventral setae are simple, except the postanal which is finely serrated. The gnathotectum has free 
lateral processes and a bifurcate median process which is minutely spiculate medially. The fixed 
chela is bidentate, with a proximal major tooth and a distal minor one, it is bidentate or tridentate 
in the male. The movable chela is basically tridentate with two adjoining proximal teeth, lacking in 
the male, and a minor distal tooth. The spermatodactyl extends dorsally then posteriorly and is 
about the length of the movable chela. The cheliceral dorsal seta is simple to slightly lanceolate. 
The legs have mainly simple setae with some pilose setae on trochanters, femora, genua and tibiae. 
The male leg armature consists of complex spurs on trochanter and femur IV, a strong spur on 
femur II and a series of other small spurs and tubercles on the genua, tibiae and tarsi of legs II and 
IV. The spermathecal sacculus is of the normal two-lobed variety with the lobes close together and 
broadly joined, the corniculum is wide, almost cup-shaped. 

INCLUDED SPECIES. M. glaber (Miiller), M. nataliae Bregetova & Koroleva, M. scutatus (Berlese). 

M. nataliae, here included in the glaber group, is obviously closely related to the other species 
although it differs slightly from them in having setae j4 and z3 simple, and the lineae obliquae 
posteriores are not obviously bifurcate as they are in other species. Krantz (198 1), in describing M. 
eurygaster, assigned it to the glaber group in spite of setaey'4 and z4 being simple in the female. They 
are pilose distally in the immatures and males. 

Macrocheles glaber (Miiller) 
(Fig. 17A-D,P1. 1C) 

The description and synonymy given by Evans & Browning (1956) remain adequate although 
Filipponi & Pegazzano (1962) have redescribed this species and give an extended synonymy, 
including M. alecto Berlese and its varieties. 


108 


K. H. HYATT & R. M. EMBERSON 


Fig. 17 Macrocheles glaber (Muller): female A dorsal shield; B setay'7; C gnathotectum; D chelicera. 
After Evans & Browning (1956). Macrocheles nataliae Bregetova & Koroleva: female E dorsal 
shield. 


MACROCHELIDAE OF THE BRITISH ISLES 1 09 

TYPE MATERIAL. M. alecto: Syntype 9 only, Italy, Firenze, Giarbino R. Stazione, humus. Slide 
1 86/ 1 5 [ISZA]. Holostaspis marginatus var. littoralis Halbert, 1915: Lectotype 9 here designated, 
Westport, Co. Mayo, Ireland, on seashore, July 1910. Slide 55 1923 [National Museum of 
Ireland, Dublin]. Paralectotypes 2 $9 on same slide, same data, 1 <$ (dissected) on two separate 
slides, both 55 1923, also same data. 

MATERIAL EXAMINED. 170 collections 9 DNN, 42 $$, c. 3, 100 99. 

ENGLAND: Isles of Scilly, Cornwall, Devon, Dorset, Gloucestershire, Somerset, Isle of Wight, 
Hampshire, Sussex, Surrey, Kent, London, Essex, Middlesex, Hertfordshire, Buckinghamshire, 
Suffolk, Norfolk, Cambridgeshire, Warwickshire, Derbyshire, Yorkshire, Lancashire, Cheshire, 
Cumbria (Cumberland, Westmorland), Northumberland. 

SCOTLAND: Dumfries & Galloway (Kirkcudbrightshire), Strathclyde (Argyllshire), Inner 
Hebrides (Mull), Outer Hebrides (Shillay), Shetland. 

WALES: Dyfed (Cardiganshire), Gwynedd (Anglesey, Caernarvonshire, Merionethshire), Clywd 
(Denbighshire). 

IRELAND: Cork, Clare, Galway, Mayo. 

HABITATS. Most often collected associated with coprophagous beetles (79 collections), records 
include: Geotrupes mutator Marsham, G. pyrenaeus Charpentier, G. spiniger Marsham, G. 
stercorosus (Scriba), G. vernalis (L.), Typhaeus typhoeus (L.), Aphodius rufipes (L.) and A. 
scybalarius (F.). M. glaber has also been collected from burying beetles Nicrophorus humator F., (2 
collections), from bumble bees Bombus agrorum F., from a fox corpse (Smith, 1975), from small 
mammals and their nests, and from a variety of manure and rotting vegetation habitats, including 
occasionally seaweed. 

DISTRIBUTION. Found everywhere in the British Isles where suitable habitats occur and probably in 
most temperate regions of the world. Filipponi & Pegazzano (1962) report it as widespread in 
Europe and the Meditteranean area. Outside these areas it has been recorded throughout the 
U.S.S.R. (Bregetova & Koroleva, 1960), U.S.A. (Axtell, 1963) and New Zealand (Emberson, 
1973a). 

Macrocheles nataliae Bregetova & Koroleva 
(Fig. 17E,P1. 6D) 

Macrocheles nataliae Bregetova & Koroleva, 1960. Parazit. Sb. 19: 140. 
Macrocheles (Macrocheles) melisii Krauss, 1970. Acarologie 14: 24. Syn. nov. 

FEMALE. The dorsal shield (Fig. 17E) is ornamented with mainly transverse reticulations and a 
microsculpture of very fine regular punctations. There are twenty-eight pairs of setae on the dorsal 
shield of which//, z2, s2, r2, r3, r4, J5, Z4, Z5, S4 and S5 are pilose distally, other dorsal setae are 
simple. 

The sternal shield (PI. 6D) has a characteristic pattern of lines and fine punctures, there are no 
coarse punctures. The linea media transversa is well defined, there is one linea arcuata which tends 
to be concave anteriorly and meet with polygonal reticulations laterally. There is an area of fine 
punctures just posterior to the linea angulata. The lineae obliquae posterior es are poorly defined and 
not noticeably bifurcate. The areae punctatae posteriores are represented mainly by a series of fine 
edging punctures. The metasternal plates are small elongate-ovate. The genital shield is weakly 
ornamented, with fine lines and punctures. The ventrianal shield is noticeably longer than broad 
and ornamented with more or less concentric reticulations anteriorly which tend to become 
polygonal posteriorly. 

The gnathosoma is typical of the species group, the gnathotectum having free lateral processes 
and a bifurcate median process which is minutely spiculate in the fork. The fixed chela has one 
major tooth and one minor distal tooth, the movable chela is tridentate, the two proximal teeth 
arising from a common base; the cheliceral dorsal seta is slightly flattened and leaf-shaped. 

The legs are normal for the group with a mixture of simple setae and stubby, faintly pilose setae 
on all segments except the coxae, trochanters, and on leg I, the tibia and tarsus. 


110 K. H. HYATT & R. M. EMBERSON 

MALE. The male is unknown in the British Isles. Krauss (1970) described a male which is probably 
this species (as melisii), it differs from other males of the species group mainly in the distribution of 
pilose dorsal setae. 

TYPE MATERIAL. M. nataliae: 1 $ syntype, Lithuanian S.S.R., Plunge, on the corpses of Microtus 
arvalis, 1 l.vii. 1954, coll. Likyavichene[ZINL]. 1 $ syntype, Tatar A.S.S.R., Kuibyshev region, in 
soil at experiment station, under grass 1 l.vi.1957, coll. Aleinikova [ZINL]. 
The type material of M. melisii was not available for study, but there can be little doubt of the 
synonymy based on the description and drawing of Krauss (1970). 

MATERIAL EXAMINED. 6 collections 20 9?. 
ENGLAND: Cornwall, Hampshire, London, Lincolnshire. 

HABITATS. The first record for the British Isles. On burying beetles Nicrophorus humator F., N. 
investigator Zett. and N. vespillo L. Bregetova & Koroleva (1960) report it from a variety of small 
mammals and also from soil samples, whilst Bregetova et al. (1977) record it from Geotrupes. 

DISTRIBUTION. Widespread in U.S.S.R., from Lithuanian SSR, to the Maritime region of Siberia 
(Bregetova & Koroleva, 1960); also reported from Germany (Karg, 1970; Krauss, 1970). 

REMARKS. There are several small inconsistencies in the details of the description and drawings of 
Bregetova & Koroleva (1960) both when compared with their syntypes and with British material. 
They do not describe setae z2, s2, Z4 and S3 as pilose and seta r4 is described as pilose but drawn 
simple, these setae are only very faintly pilose in British material. They mention that there is some 
variation in the presence of pilose setae and certainly in two syntypes examined z2 was simple, 
although it is pilose in British material. Krauss (1970) also records it as simple. The other setae 
mentioned were pilose in both syntypes from the U.S.S.R., in British material and in that illus- 
trated by Krauss (1970). It appears that there may be some slight variation in the distribution of 
pilose dorsal setae but this is perhaps not surprising in a species distributed across the whole of the 
Palaearctic region. 

Macrocheles scutatus (Berlese) 
(Fig. 18, PI. IE) 

Holostaspis subbadius var. scutatus Berlese, 1904. Redia 1: 264. 

Macrocheles subbadius: Evans & Browning, 1956, non Berlese, 1904. Bull. Br. Mus. not. Hist. (Zool.) 4: 19. 

Macrocheles scutatus (Berlese): Filipponi & Pegazzano, 1962. Redia 47: 228. 

The female of this species is adequately described by Evans & Browning (1956) under the name 
M. subbadius (Berlese) which Filipponi & Pegazzano (1963) have shown to be a completely 
different species. 

The male has been described on the basis of reared material by Filipponi & Pegazzano (1962). It 
differs from related species chiefly in the distribution of pilose dorsal setae, in details of the shape 
and ornamentation of the ventral sclerotisation and in the leg armature. 

MATERIAL EXAMINED. 5 collections 17 33, 24 $?. 
ENGLAND: Worcestershire, Derbyshire, Yorkshire. 
WALES: Glamorgan. 

HABITATS. Farmyard manure. Filipponi & Pegazzano (1962) record this species from dung and 
compost habitats, also associated with coprophagous beetles, but not muscid flies. 

DISTRIBUTION. Italy (Filipponi & Pegazzano, 1962), U.S.S.R. (Bregetova & Koroleva, 1960 as 
M. subbadius and Bregetova et al., 1977), New Zealand (Emberson, 19736). The continuing 
confusion of closely related species, including M. vicinus Leitner (1946), makes assessment of the 
distribution of this species problematical. 

subbadius group 

This species group was reviewed by Filipponi & Pegazzano (1963) who defined it as consisting of 
small species, with 28 pairs of simple needle-like setae in the females, with setaey'7 short, spine-like 


MACROCHELIDAE OF THE BRITISH ISLES 


111 


Fig. 18 Macrocheles scutatus (Berlese): female dorsal shield. After Evans & Browning (1956). 


and diverging. The dorsal shield of males is more or less strongly tuberculate posteriorly, and has at 
least two pairs of large blunt, slightly spatulate setae (j4, z4). The ventral shields are ornamented 
with a series of punctate lines; in the females the sternal shield has the lineae oblique anteriores 
connected transversely by 4 or 5 lines, the most posterior of which is the linea media transversa. All 
the ventral setae are simple. The males have a holoventral shield. The gnathotectum has the lateral 
processes free and the median process bifurcate and minutely spiculate distally. Both chelae are 
basically bidentate in the female, but in the male the fixed chela is tridentate and the movable chela 
unidentate; the dorsal seta is simple and the spermatodactyl is elongate, tapering, strongly 
recurved and almost twice as long as the movable digit. All the leg setae are simple, although 
spinose on tarsi II-IV. The males have the femora, genua, tibiae and tarsi of legs II armed with 
spines and tubercles, leg III is unarmed and the armature of leg IV is more variable. 

INCLUDED SPECIES: M. subbadius (Berlese), M. insignitus Berlese and M. merdarius (Berlese). 

Macrocheles subbadius (Berlese) 
(Fig. 19 A, PI. 6E) 

Holostaspis subbadius Berlese, 1904. Redia 1: 264. non Macrocheles subbadius, Evans & Browning, 1956. Bull. 
Br. Mus. not. Hist. (Zool.) 4: 19. 

The species described as M. subbadius by Evans & Browning (1956) is in fact M. scutatus (vide 
Filipponi & Pegazzano, 1962). However, M. subbadius does occur in the British Isles and a short 
description is given below. For a detailed description and synonymy see Filipponi & Pegazzano 
(1963). 


112 


K. H. HYATT & R. M. EMBERSON 


Fig. 19 Macrocheles subbadius (Berlese): female A dorsal shield. Macrocheles merdarius (Berlese): 
female B dorsal shield; C gnathotectum; D chelicera. B-D after Evans & Browning (1956). 


FEMALE. The dorsal shield (700-750 um long x 400-440 um wide) is punctate-reticulate, the lateral 
margins are slightly dentate and the posterior margin has three small incisions between setae Z5 
(Fig. 19 A). 

The sternal shield is strongly patterned with punctate lines (PI. 6E) and the lineae oblique 
anteriores are connected by five transverse lines, the most posterior of which, the linea media 


MACROCHELIDAE OF THE BRITISH ISLES 1 1 3 

transversa, is straight. Genu IV has seven setae, seta/?/ ; being present. Other characters similar to 
those of M. insignitus. 

MALE. Unknown from the British Isles, but distinguished from M . insignitus and M. merdarius by 
having only two pairs of enlarged dorsal setae and three pairs of dorsal tubercles, also tarsus II 
lacks a strong medial spine and femur IV has a strong spur (Filipponi & Pegazzano, 1963). 

TYPE MATERIAL. Lectotype ?, Italy, Padova, letamai [manure], 1882, Vecchia coll. Designated by 
Filipponi & Pegazzano, 1963, slide 19/41 [ISZA]. 

MATERIAL EXAMINED. 7 collections 40 ?$. 
ENGLAND: Surrey, Oxfordshire, Northumberland, Cheshire. 
IRELAND: Clare. 

HABITATS. The first record for the British Isles. Phoretic on houseflies and in garden refuse. It has 
also been found on other species of flies and in manure. Thirteen females were removed from 1 1 
specimens of the dipteran Meroplius minutus Wiedemann (Sepsidae) from Goring-on-Thames, 
Oxfordshire, coll. A. C. and B. Pont, 26.8.1985. Ten of the flies were carrying one apiece and the 
eleventh was carrying three when examined by K.H.H., although a few of the mites had been lost 
during the setting of the flies. They were attached to the ventral side of either the abdomen or the 
thorax. 

DISTRIBUTION. Widespread in temperate regions. Italy (Filipponi & Pegazzano, 1963), Germany 
(Karg, 1971), U.S.A. (Axtell, 1961), New Zealand (Emberson, 19736). 


Macrocheles insignitus Berlese 
(PI. IF) 

The description of the female of this species as given by Evans & Browning (1956) needs no 
amendment. The male has been described from rearing experiments by Filipponi & Pegazzano 
(1963). 

TYPE MATERIAL. Holotype 9, France, Longuy Orne, musco [moss] Cordier coll., slide 58/50 [ISZA]. 

MATERIAL EXAMINED. 4 collections 1 cJ, 3 ?$. 
ENGLAND: Norfolk, Warwickshire, Lancashire. 
WALES: Gwynedd (Caernarvonshire). 

HABITAT. Grass cuttings, compost, nest of bumble-bees Bombus sp. and wet mosses. 

DISTRIBUTION. France, Italy (Filipponi & Pegazzano, 1963). Apparently a European species that is 
only rarely collected. 

Macrocheles merdarius (Berlese) 
(Fig. 19B-D,P1. 2A) 

Holostaspis merdarius Berlese, 1889. Acari, Myriapoda et Scorpiones, etc. Fasc. 52, T.I. 

Holostaspis adulescens Berlese, 1910. Redia 6: 252. 

Macrocheles merdarius: Sellnick, 1940. Goteborgs K. Vetensk.-o. vitterhSamh. Handl. B6, 14: 86. 

The description of the female given by Evans & Browning (1956) remains unchanged. The male 
remains unknown in collections from the British Isles, but has been redescribed by Filipponi & 
Pegazzano (1963). 

TYPE MATERIAL. Neotype $, Italy, Vittorio, Veneto, letamai [manure]. Designated by Filipponi & 
Pegazzano (1963). Not seen, but slide material from the same collection has been examined, slides 
89/15, 17, 46, 48 [ISZA]. 

MATERIAL EXAMINED. 3 collections c. 30 ??. 
ENGLAND: Surrey, Hertfordshire. 


114 K. H. HYATT & R. M. EMBERSON 

HABITATS. Two collections from cattle dung and stable manure and one collection from a guinea- 
pig breeding unit. Elsewhere this species has been collected from flies. 

DISTRIBUTION. M. merdarius appears to be practically cosmopolitan and has been recorded from 
many countries in Europe, the Americas, Africa and Australia (vide Filipponi & Pegazzano, 1963). 
In addition to these records it has been reported from Israel (Costa, 19666) and New Zealand 
(Emberson, 19730). 


Genus GLYPTHOLASPIS Filipponi & Pegazzano 

Glyptholaspis Filipponi & Pegazzano, 1960. Redia 45: 136. Type species: Nothrholaspis fimicola Sellnick, 
1931 =Gamasus tardus Berlese, 1882. non C. L. Koch, 1841. 

All the shields are strongly reticulate. The posterior and lateral margins of the dorsal shield are 
dentate, with 28 pairs of setae, supernumerary setae are sometimes present between setaeytf and J2. 
The sternal shield has a characteristic raised polygonal pattern (PI. 3C) and extends posteriorly to 
the level of the posterior margins of coxae III, where it abuts rounded metasternal platelets. The 
genital and ventrianal shields also have a raised polygonal pattern; the ventrianal shield is truncate 
and angled anteriorly and rounded posteriorly. The median process of the gnathotectum is 
strongly bifid and spiculate, the lateral processes are sometimes partially fused basally. The fixed 
chela has four or five teeth in the female and three in the male, the movable chela usually has three 
teeth in the female and one in the male, the dorsal seta is dentate distally and the male spermato- 
dactyl is simple, dorsally directed, tapering and recurved distally. The male has a holoventral shield 
and spurs on legs II, III and IV. 

Filipponi & Pegazzano (1960) erected the genus for a clearly defined group of species formerly 
included in the genus Macrocheles under a variety of names. There is however some doubt as to the 
rank that should be assigned to this group (Krantz, 1962). 

The recent synonymy of the three species of Glyptholaspis found in the British Isles with 
Macrocheles tardus (C. L. Koch) by Krauss (1970) cannot be supported, whether the genus 
Glyptholaspis is accepted or not. Filipponi & Pegazzano (1960) have shown the three species to be 
consistently different morphologically, in samples from various parts of the world, and Filipponi & 
Ilardi (1958) showed them to be reproductively isolated. The present authors also follow Sellnick 
(1931) and most other recent European authors in regarding Gamasus tardus C. L. Koch, 1841, as 
being a common European Macrocheles species in moss and leaf litter, whereas the Glyptholaspis 
species, including Berlese's 1882 concept of Gamasus tardus, are usually found in compost heaps, 
dung and associated with flies. Three species are known from the British Isles, two of which were 
previously confused under the name Macrocheles plumiventris Hull. All species are described in 
detail with full synonymies by Filipponi & Pegazzano (1960). 


Key to the species of Glyptholaspis known to occur in the British Isles 

1 Posterior margin of the dorsal shield between setae Z5 with five large teeth, and, in the females, 
numerous minute teeth, setaey'6 and z6 more or less in a straight line, seta J5 no longer than setae 

Z5; trochanter II of male with a spur 

Posterior margin of the dorsal shield between setae Z5 with two large teeth and numerous minute 
teeth (Fig. 20A), setae j6 anterior to setae z6, setae J5 distinctly longer than setae Z5; trochanter II 
of male unarmed Glyptholaspis americana (Berlese) (p. 116) 

2 Setae j5 strongly pilose and erect, setae J5 shorter than Z5; gnathotectum with lateral processes 
fused (Fig. 20E); female sternal shield pattern symmetrical, with a posterior median ridge (PI. 3C); 
male trochanter IV with three subequal ventral spurs, femur IV unarmed 

Glyptholaspis confusa (Foa) (p. 1 16) 

Setae j5 lightly pilose and adpressed, setae J5 about equal in length to Z5 in the female; gnatho- 
tectum with lateral processes free. Female sternal pattern asymmetrical, without a distinct 
posterior median ridge (PI. 5E); male trochanter IV and femur IV each with a posteriorly directed 
spur . Glyptholaspis fimicola (Sellnick) (p. 117) 


MACROCHELIDAE OF THE BRITISH ISLES 


115 


Fig. 20 Glyptholaspis americana (Berlese): female A dorsal shield. Glyptholaspis confusa (Foa): 
female B anterior region of dorsal shield; C lateral margin of dorsal shield; D ornamentation of 
dorsal shield; E gnathotectum; F chelicera; G dorsal seta of chelicera. B-G after Evans & Browning 
(1956). 


116 K. H. HYATT & R. M. EMBERSON 

Glyptholaspis americana (Berlese) 
(Fig. 20A, PI. 6F) 

Holostaspis marginatus var. americanus Berlese, 1888. Boll. Soc. ent. ital. 20: 195. 
Holostaspis vagabundus Berlese, 1889. Acari, Myriopoda et Scorpiones, etc. Fasc. 52, T. 9. 
Macrocheles tardior Hull, 1925. Ann. Mag. nat. Hist. (9) 15: 126. Syn. nov. 
Glyptholaspis americana (Berlese): Filipponi & Pegazzano, 1960. Redia45: 148. 
Macrocheles (M.) tardus: Krauss, 1970. Acarologie 14: 2Q,partim, necC. L. Koch, 1841. 

FEMALE. The dorsal setae 76, J2 and z5 are glabrous, setae j5, z6 and J5 are slightly pilose, other 
setae are strongly pilose. Setae J5J6, z6 and J2 are relatively long and slender, of similar length; 
setae J5 are distinctly longer than setae Z5. The posterior and lateral margins of the dorsal shield 
are strongly dentate, the posterior margin between setae Z5 has two large teeth and an even row of 
minute ones (Fig. 20A). 

All ventral shields are strongly reticulated with raised polygonal pattern. Sternal shield pattern 
(PI. 6F) slightly asymmetric without a distinct median posterior ridge. Sternal and metasternal 
setae simple, genital setae moderately plumose, setae Jvl-3 plumose, adanal setae simple, postanal 
seta short and plumose. 

The gnathotectum has fused lateral processes and a bifurcate median process. The fixed chela 
has four teeth, the movable chela has three teeth. 

The legs are normal for the genus with plumose setae on all segments except tarsus I. 

MALE. The male is unknown from the British Isles, but has been adequately described by Filipponi 
& Pegazzano (1960). The main distinguishing features are the chaetotaxy of the dorsum which is 
similar to the female and the form of the spurs on legs II, III and IV. 

TYPE MATERIAL. Gamasus americanus: Holotype 9, South America, Balzan, slide 33/21 [ISZA]. M. 
tardior. Syntype ?, Oxfordshire [? R. S. Bagnall] [BMNH]. 

MATERIAL EXAMINED. 4 collections 1 1 99- 
ENGLAND: Isles of Scilly, Sussex, Oxfordshire. 
IRELAND: Clare. 

HABITATS. Geotrupes spiniger, also known elsewhere from compost heaps and associated with 
synanthropic flies. 

DISTRIBUTION. Apparently practically cosmopolitan. Filipponi & Pegazzano (1960) report it from 
Brazil, Argentina, Uruguay, Australia, South Africa, Italy and France. It has also been reported 
from the U.S.S.R. (Bregetova & Koroleva, 1960, as M. vagabundus), Bulgaria (Balogh, 1958, as M. 
vagabundus), Germany (Krantz, 1972), Israel (Costa, 1963) and New Zealand (Emberson, 1973#). 

REMARKS. In examining the Hull Collection, Professor G. O. Evans (in litt.) did not believe the 
specimen of M. tardior to be the type as it is larger than the dimensions stated by Hull. However, it 
is labelled from Oxfordshire, the type locality, and agrees fairly well with the description, which 
undoubtedly refers to a species of Glyptholaspis, so we see no good reason to doubt its status. 

Glyptholaspis confusa (Foa) 
(Fig. 20B-G, PI. 3C) 

Holostaspis confusa Foa, 1900. Boll. Soc. ent. Ital. 32: 137. 

Macrocheles plumi ventris Hull, 1925. Ann. Mag. nat. Hist. (9) 15: 216. 

Macrocheles plumiventris: Evans & Browning, 1956. Bull. Br. Mus. nat. Hist. (Zool.) 4: 32. partim, female 

only. 
Glyptholaspis confusa: Filipponi & Pegazzano, 1960. Redia45: 154. 

The female of this species has been adequately described by Evans & Browning (1956) under the 
name Macrocheles plumiventris Hull. The male described under that name is in fact a specimen of 
G.fimicola. 

The only necessary additions to the description of the female are to note that there are usually 
one or two small glabrous or finely pilose supernumerary setae betweeny'6 and J2, that setae J5 are 


MACROCHELIDAE OF THE BRITISH ISLES 1 1 7 

shorter than setae Z5 and that the posterior margin of the dorsal shield between setae Z5 is 
irregularly dentate with about five major teeth and many small teeth. 

MALE. The sculpturing and chaetotaxy of the dorsal shield are similar to that of the female. The 
reticulate pattern on the holoventral shield is interrupted in the region of coxae IV. Setae st 1 are 
plumose, st 2-3 simple, st 4-5 plumose. All the ventrianal setae except the adanals are plumose. 

The gnathotectum is similar to the female. The fixed chela has three teeth and the movable chela 
one tooth. The spermatodactyl is simple, tapered and recurved towards the tip, about as long as the 
movable digit. 

Leg I is similar to that of the female. Leg II has sclerotized spurs on the trochanter, femur, genu 
and tibia. The spur on trochanter II is towards the posterior lateral surface and bears a plumose 
seta distally. Femur II has a number of sclerotised protuberances, including a small ventral spur 
with a simple seta and another spur on the posterior lateral surface at the distal margin. Genu II has 
a ventral spur and three other small protuberances and tibia II has a ventral spur on the distal 
margin and trochanter IV has three strong ventral spurs. 

TYPE MATERIAL. Macrocheles plumiventris: A single female without habitat data in the Hull 
Collection [BMNH]. Hull (1925) gave 'a manure-heap in West Allendale' [Northumberland] for 
his solitary example. 

MATERIAL EXAMINED. 8 collections 8 <$<$, 31 $9- 

ENGLAND: Sussex, Essex, Hertfordshire, Middlesex, Berkshire, Buckinghamshire, Bedfordshire. 
WALES: Gwynedd (Merionethshire). 

HABITATS. Manure, granary refuse, leaf litter, also known from compost and associated with 
synanthropic flies. 

DISTRIBUTION. G. confusa has been reported from Italy, Argentina and Australia (Filipponi & 
Pegazzano, 1960), Bulgaria (Balogh, 1958 as M. plumiventris), U.S.S.R. (Bregetova & Koroleva, 
1960 as M. plumiventris), Germany (Krantz, 1972) and New Zealand (Emberson, 19730). 

Gfyptholaspis fimicola (Sdlnick) 

(PI. 5E) 

Gamasus tardus: Berlese [non C. L. Koch, 1841], 1882 Boll. Soc. ent. Ital. 14: 108. 

Holostaspis marginatus: Berlese [non Hermann, 1804], 1889, Acari, Myriopoda et Scorpiones, etc. Fasc. 52, 

T. 4-5. 
Nothrholaspis fimicola Sellnick, 1931. Sber. Akad. Wiss. Wien 140: 765. Nom. nov. pro Holostaspis 

marginatus: Berlese [non Hermann, 1804,] 1889. 

Macrocheles plumiventris'. Evans & Browning, 1956. Bull. Br. Mus. nat. Hist. (Zool.) 4: 38, male only. 
Glyptholaspis fimicola (Berlese) [sic]: Filipponi & Pegazzano, 1960. Redia45: 139. 

As suggested by Filipponi & Pegazzano (1960) the specimen described as the male of M. 
plumiventris by Evans & Browning (1956) from the Michael Collection is not conspecific with the 
females ascribed to that species and is the only known specimen of G. fimicola from the British Isles. 

FEMALE. The following brief description is taken from Filipponi & Pegazzano (1960) supplemented 
by examination of Italian material. 

Generally most similar to G. confusa but differing as follows. On the dorsal shield many of the 
setae are longer than their counterparts in G. confusa: j5, j6, J2 and z6 are similar in form and 
length, lightly pilose; in G. confusa j5 is more strongly pilose than the others and erect rather than 
adpressed;y'6 is longer than the distance between the bases ofytf and z6. J2 is about equal in length to 
the distance between the bases of seta 72, without supernumerary setae between setaey<5 and J2. J5 
is about equal in length to Z5. 

Setae st 1 on the sternal shield are simple, not pilose, the sculpturing of the sternal shield (PI. 5E) 
is very similar to that of G. americana, lacking the median posterior ridge found in G. confusa. 

The gnathotectum has the two lateral processes free, not fused as in G. confusa and G. americana. 
The fixed chela has four teeth and the movable chela three teeth. 

The legs are very similar to those of G. confusa. 


118 K. H. HYATT & R. M. EMBERSON 

MALE. The male of this species has been described by Evans & Browning (1956) under the name 
M. plumiventris Hull. 

The chaetotaxy of the dorsum is generally similar to that of the female, except that J5 tends to be 
shorter than Z5. Between setae Z5 there are five large teeth and no small teeth (Italian specimens). 
The pattern of the holo ventral shield is not interrupted in the region of coxae IV. 

The gnathotectum is similar to that of the female. The fixed chela has four teeth and the movable 
chela a single tooth, the spermatodactyl is longer than the movable digit, strongly recurved and 
pointed distally. 

Leg II has spurs on the trochanter, femur, genu and tibia, the distal spur on the femur has a pilose 
seta terminally. There are posteriorly directed spurs on genu III, trochanter IV and femur IV, the 
spur on trochanter IV has a terminal pilose seta. 

MATERIAL EXAMINED. 1 collection 1 $, A. D. Michael Collection (1930.8.25.2215). 
ENGLAND: Warwickshire: Austrey. May 1892. No habitat given. 

HABITATS. Known from a variety of sorts of dung and compost (Filipponi & Pegazzano, 1960). 

DISTRIBUTION. Apparently only previously recorded from southern Europe, Italy (Filipponi & 
Pegazzano, 1960) and Greece (Sellnick, 1931). 


Genus HOLOSTASPELLA Berlese 

Holostaspella Berlese, 1903. Redia 1: 241. Type species: Holostaspis (Holostaspella) sculpta Berlese, 1903. 
Prholaspina Berlese, 1918. Redia 13: 174. Type species: Holostaspella (Prholaspina) micrarrhena Berlese, 

1916. 
Areolaspis Tragardh, 1952. Ark. Zool. 4 (3): 60. Type species: Areolaspis bifoliatus Tragardh, 1952. 

Most species are heavily sclerotised with strong ornamentation of the main shields. Dorsal shield 
with 28 pairs of setae, jl short and pectinate, usually borne on a tuberculate anterior extension of 
the shield; lateral region of the dorsal shield often with a series of depressions. The sternal shield 
nearly always has a strong median ridge; the metasternal platelets are often enlarged and fused to 
the endopodal sclerites. The ventrianal shield may be narrow or broad with 2-4 pairs of preanal 
setae (four pairs in species from the British Isles). The peritrematic shield may be fused with the 
expodal sclerites. Males have either a holoventral shield or separate ventrianal shields. The sper- 
mathecal apparatus is of the normal bilobed Macrocheles type. The gnathosoma is normal for the 
family. The lateral processes of the gnathotectum are free and the cheliceral brushes are shorter 
than the movable digit of the chelicera. Legs II in the females are armed with one or more spurs on 
the femora. Spurs or protuberances may also be present on the coxae, trochanters and genua, seta 
mv of tarsus II is also developed into a strong spine in all British species. Male leg armature is 
limited to femur II. Leg chaetotaxy is normal for the family. 

Following Krantz (1962) and Petrova & Taskaeva ( 1 964) the concept of Holostaspella has been 
considerably broadened. Filipponi & Pegazzano (1967) and Krantz (1967) have both revised the 
genus and split it into a number of species groups. The species from the British Isles fit into the 
ornata group of Filipponi & Pegazzano and the sculpta group of Krantz. 

Key to species of Holostaspella known to occur in the British Isles 

1 Dorsal shield lacking distinct depressions, setae zl more than half as long as j2, j3 distinctly 
anterior to z2 (Fig. 22C); female sternal shield lacking cruciform pattern 

Holostaspella subornata Bregetova & Koroleva (p. 122) 

Dorsal shield with lateral depressions, setae zl less than half as long as/2,/3 and z2 more or less 
level (Fig. 21 A); female sternal shield with cruciform pattern (PI. 5C, D) 2 

2 Seta zl pilose, setae j3 and z2 as long as or longer than the distance between the bases ofj3 andj4 
(Fig. 2 1 A); female sternal shield as in Pis 4D, 5C . Holostaspella ornata (Berlese) (p. 1 20) 
Setae zl simple, not pilose, setae j3 and z2 shorter than the distance between the bases ofj3 andj4 
(Fig. 22A); female sternal shield as in PI. 5D 

Holostaspella exornata Filipponi & Pegazzano (p. 120) 


Fig. 21 Holostaspella ornata (Berlese): female A dorsal shield; B anterior region of dorsal shield; C 
gnathotectum; D chelicera; E leg II, ambulacrum omitted. After Evans & Browning (1956). 


1 20 K. H. HYATT & R. M. EMBERSON 

Holostaspella ornata (Berlese) 
(Fig. 21A-E, Pls4D, 5C) 

The description and synonymy of this species are as given by Evans & Browning (1956), the 
description being based on the type in the Oudemans Collection, although the British specimen 
they mentioned belongs in fact to the next species, H. exornata. 

MATERIAL EXAMINED. 7 collections 12 99. 

ENGLAND: Isles of Scilly, Surrey, Kent, Yorkshire, Cumbria (Cumberland). 
IRELAND: Clare. 

HABITATS. The first record for the British Isles. Under a board, in leaf litter, dead reeds, compost 
and manure. 

DISTRIBUTION. A European species, Holland (Berlese, 1904), Russia (Bregetova & Koroleva, 
1960), Austria (Franz, 1954), Germany (Krauss, 1970; Krantz, 1972), also recorded from Zaire 
(Krantz, 1967), but this could be a closely related species. 

REMARKS. Filipponi & Pegazzano ( 1 967) have suggested that some of the material assigned to this 
species by Bregetova & Koroleva (1960) belongs in fact to the following species. Their illustrations 
of the dorsal and ventral surfaces certainly appear to be of specimens of H. exornata and the size 
range overlaps those of both species as given by the former authors. 

Holostaspella exornata Filipponi & Pegazzano 
(Fig. 22A-B, PI. 5D) 

Holostaspella exornata Filipponi & Pegazzano, 1967. Redia 50: 230. 

Recorded by Evans & Browning (1956) as Holostaspella ornata (Berlese). 

This species is morphologically extremely similar to the preceding species but differs from it 
consistently in a number of ways, most obviously in size, female dorsal shield length 649-800 urn as 
against 917-948 um for H. ornata (Filipponi & Pegazzano, 1967). 

FEMALE. Setae of the dorsal shield (Fig. 22 A) generally shorter than in H. ornata J3 not reaching 
bases oij4, s5 shorter than distance between s5 and 26, J5 shorter than the distance between its base 
and the posterior margin of the dorsal shield. Setae jl broadly pectinate, zl smooth, simple, all 
other setae finely pilose. 

Sternal shield divided by cross-shaped ridges into quadrants, the point where the arms of the 
cross meet is not expanded and reticulated (PI. 5D). Anterior margin of sternal shield with four 
tooth-like projections medially (smoothly concave in H. ornata), st 1 lightly pilose, other sternal 
setae simple. Metasternal platelets elongate, abutting endopodals. The ventrianal shield has four 
pairs of simple preanal setae and is ornamented with a series of small reticulations within large 
reticulations. 

The gnathosoma is very similar to that of H. ornata. The fixed chela has two teeth and the 
movable chela one tooth. The lateral processes of the gnathotectum are free. 

The legs are strongly rugose, with all segments finely granular. On leg II the femur and tro- 
chanter are armed with small spurs and tibia II has seta mv produced into a spine (Fig. 22B). 

MALE. Not known from the British Isles, but described by Filipponi & Pegazzano (1967) from 
reared material. 

The chaetotaxy is generally similar to the female. The holoventral shield has a strong median 
longitudinal ridge in the sternal region. The spermatodactyl is elongate, tapering, more than twice 
as long as the movable digit. Legs unarmed. 

MATERIAL EXAMINED. 5 collections 9 99- 
ENGLAND: Kent, London, Berkshire, Warwickshire. 

HABITATS. Phoretic on Sphaerocera spp. (Diptera) in rotting vegetation. It is of interest to note that 
in two of the five collections the specimens were associated with Sphaerocera spp. Filipponi & 


MACROCHELIDAE OF THE BRITISH ISLES 


121 


Fig. 22 Holostaspella exornata Filipponi & Pegazzano: female A dorsal shield; B leg II. Holostaspella 
subornata Bregetova & Koroleva: female C dorsal shield; D ventral sclerotisation. D after Bregetova 
&Koroleva(1960). 


1 22 K. H. HYATT & R. M. EMBERSON 

Pegazzano ( 1 967) mention a specimen in the Berlese Collection from the nest of an ant Acromymex 
lundi in Argentina. 

DISTRIBUTION. Italy, Argentina, ? U.S.S.R. Filipponi & Pegazzano (1967) suspect that some of the 
material reported from U.S.S.R. by Bregetova & Koroleva (1960) as H. ornatamay be referrable to 
this species. 

Holostaspella subornata Bregetova & Koroleva 
(Fig. 22C, D) 

Holostaspella subornata Bregetova & Koroleva, 1960. Parazit. Sb. 19: 60 

This species is generally similar to the preceding two but differs from them most noticeably in the 
lack of strong lateral sculpturing on the dorsal shield and in details of the dorsal chaetotaxy. 

FEMALE. Dorsal shield (Fig. 22C) lacking lateral depressions, ornamented with densely packed fine 
punctures. Setae of the dorsal shield relatively long, as in H. ornata, but simple to minutely barbed 
rather than conspicuously pilose. Setae jl are broadly pectinate and in close -proximity. Setae j2 
reach the bases of/3,/3 are displaced substantially anterior to z2, so that they do not reach the bases 
of/4, zl are elongate, more than half as long asy'2. 

The sternal shield (Fig. 22D) is punctate reticulate, lacking the cruciform pattern of H. ornata 
and H. exornata, and the anterior margin is smoothly concave. All the ventral setae are simple. The 
metasternal platelets are barely separated from the posterior corners of the sternal shield and the 
endopodals. The genital and ventrianal shields are punctate reticulate. 

The gnathosoma is very similar to the other species of Holostaspella. The fixed chela has one 
major and two minor teeth and the movable chela one tooth. The lateral processes of the 
gnathotectum are free. 

The legs are rugose and all segments covered in fine granulations. Leg II has small spurs on the 
trochanter and femur and seta mv on the tibia is produced into a spine. 

MALE. Unknown. 

MATERIAL EXAMINED. 1 collection 1 $. 
IRELAND: Clare. 

HABITAT. The first British record. Amongst grass roots. Bregetova & Koroleva report this species 
from leaf litter and associated with the musk-rat Ondatra zibethicus. 

DISTRIBUTION. Only otherwise known from the Leningrad and Omsk regions of the U.S.S.R. 
(Bregetova & Koroleva, 1960). 


Taxonomic Summary 

(a) Macrocheles analis sp. nov. is described. 

(b) Neotypes are designated for Macrocheles tridentinus (G. & R. Canestrini, 1882) and 

Macrocheles terreus (Canestrini & Fanzago, 1877). 

(c) Lectotypes are selected for Macrocheles submotus Falconer, 1924, Macrocheles matrius Hull, 

1925, Holostaspis echinatus Berlese, 1904, and Holostaspis marginatus var. littoralis Halbert, 
1915. 

(d) The genus Dissoloncha Falconer, 1923, is resurrected. 

(e) The following new synonymy is proposed: 

Macrocheles melisii Krauss, 1970, is a junior synonym of Macrocheles nataliae Bregetova & 

Koroleva, 1960. 
Macrocheles bombophilus Gotz, 1970, is a junior synonym of Macrocheles rotundiscutis 

Bregetova & Koroleva, 1960. 


MACROCHELIDAE OF THE BRITISH ISLES 1 23 

Macrocheles multisetosus Gotz, 1970, is a junior synonym of Macrocheles dentatus (Evans & 

Browning, 1956). 

Macrocheles tardior Hull, 1925, is a junior synonym of Glyptholaspis americana (Berlese, 
1888). 

(/) Species new to the British Isles: 

Geholaspis (Geholaspis) aeneus Krauss, 1970. 

Geholaspis (Longicheles) hortorum (Berlese, 1904). 

Macrocheles rotundiscutis Bregetova & Koroleva, 1960. 

Macrocheles analis sp. nov. 

Macrocheles punctatissimus Berlese, 1918. 

Macrocheles nataliae Bregetova & Koroleva, 1960. 

Macrocheles subbadius (Berlese, 1904). 

Holostaspella ornata (Berlese, 1904), nee Evans & Browning, 1956. 

Holostaspella subornata Bregetova & Koroleva, 1960. 


Acknowledgements 

Numerous acarologists have helped in this study with advice, access to material in their charge, and the loan of 
valuable specimens for which we are most grateful. Our special thanks go to Dr Fausta Pegazzano for the 
benefit of her wide experience with the Macrochelidae and her generous help to one of us (R.M.E.) with the 
Berlese collection; to Dr Nina Bregetova for the loan of valuable type material in her charge; to Dr Jerry 
Krantz for his superb hospitality to R.M.E. and for the benefit of his broad knowledge and valuable collection 
of Macrochelidae. We also wish to thank Professor P. Omodeo and Dr A. Minelli, Institute di Biologia 
Animale dell' Universita di Padova, for access to and help with the Canestrini Collection, Dr J. P. O'Connor, 
National Museum of Ireland, for the loan of material from the Halbert collection, and Dr M. V. Hounsome, 
Manchester Museum, for the loan of Harry Britten specimens. 

R.M.E. also wishes to express his sincere thanks to Dr J. G. Sheals, former Keeper of Zoology, and to the 
Trustees of the British Museum (Natural History) for providing working facilities and access to the collec- 
tions on which this work is based. Thanks are due also to Mr Peter V. York for photomicrography and Mr D. 
Macfarlane, CAB International Institute of Entomology, for critical comments on the manuscript. 


References 

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- 1967. Notes on macrochelids associated with manure and coprid beetles in Israel. II. Three new species 
of the Macrocheles pisentii complex with notes on their biology. Acarologia 9: 304-329. 

Emberson, R. M. 1973a. Macrochelid mites in New Zealand (Acarina: Mesostigmata: Macrochelidae). N.Z 

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1973/>. Additions to the macrochelid mites in New Zealand (Acarina: Mesostigmata: Macrochelidae) 

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- & Browning, E. 1956. British mites of the subfamily Macrochelinae Tragardh (Gamasina, Macro- 
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& - - 1979. Mesostigmatic mites of Britain and Ireland (Chelicerata: Acari-Parasitiformes). An 


introduction to their external morphology and classification. Trans, zool. Soc. Lond. 35: 139-270. 
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& 1962. Specie Italiane del gruppo-glaber (Acarina, Mesostigmata, Macrochelidae, Macrocheles). 

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& 1963. Specie Italiane del gruppo-subbadius (Acarina, Mesostigmata, Macrochelidae). Redia 48: 
69-91. 

& 1967. Contribute alia conoscenza del genere Holostaspella Berlese, 1903. (Acari: Mesostigmata: 

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329^52. 
Halaskova, V. & Kunst, M. 1 960. Uber einige Bodenmilben-gruppen aus dem Moorgebeit "Soos" in Bohmen. 

(Acari: Gamasina, Zerconina, Oribatei). Ada Univ. Carol. Biol. Suppl. 1960:1 1-58. 
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Proc. R. Ir. Acad. 31: 45-136. 
Halliday, R. B. 1986. On the systems of notation used for the dorsal setae in the family Macrochelidae 

(Acarina). Internal. J. Acarol. 12: 27-35. 
Hammen, L. van der. 1964. The morphology of Glyptholaspis confusa (Foa, 1900) (Acarida, Gamasina). Zool. 

Verh. Leiden No. 71: 1-56. 
Hirschmann, W. 1957. Gangsystematik der Parasitiformes. Teil 1. Rumpfbehaarung und Riickenflachen. 

Acarologie 1: 1-20. 

Hull, J. E. 1918. Terrestrial Acari of the Tyne Province. Trans, nat. Hist. Soc. Northumb. 5, 1: 13-88. 
Johnston, D. E. 1970. Notes on a collection of Austrian Macrochelidae with a description of Macrocheles 

beierin. sp. Annln. naturh. Mus. Wien 74: 145-150. 
Karg. W. 1971. Acari (Acarina), Milben Unterordnung Anactinochaeta (Parasitiformes). Die freilebenden 

Gamasina (Gamasides), Raubmilben. Tierwelt Dtl. 59: 475 pp. 
Krantz, G. W. 1960. A re-evaluation of the Parholaspinae Evans, 1965 (Mesostigmata: Macrochelidae). 

Acarologia 2: 293-433. 

1962. A review of the genera of the family Macrochelidae Vitzthum, 1930 (Acarina: Mesostigmata). 
Acarologia 4: 143-173. 

1965. A review of the genus Neopodocinum Oudemans, 1902 (Acarina: Macrochelidae). Acarologia 7: 
139-226. 

- 1967. A review of the genus Holostaspella Berlese, 1904 (Acarina: Macrcochelidae). Acarologia 9, fasc. 
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1972. Macrochelidae from Hamburg (Acari Mesostigmata), with descriptions of two new species. Ent. 
Mitt. zool. Mus. Hamburg 4: 263-275. 


MACROCHELIDAE OF THE BRITISH ISLES 1 25 

1 981. Two new glaber group species of Macrocheles (Acari: Macrochelidae) from southern Africa. Int.J. 
Acarol.l:3-\6. 

& Filipponi, A. 1964. Acari della famiglia Macrochelidae (Mesostigmata) nella Collezione del South 


Australian Museum. Riv. Parassit. 25: 35-54. 
Krauss, W. 1970. Die europaischen Arten der Gattungen Macrocheles Latreille 1829 und Geholaspis Berlese 

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Leitner, E. 1946. Zur Kenntnis der Milbenfauna auf Diingerstatten. Zentbl. Gesamtgeb. Ent. 1: 75-156. 
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for the idiosoma of the Gamasina (Acarina: Mesostigmata). Mem. ent. Soc. Can. 47: 1-64. 
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gamasoiden Milben Macrochelidae Vitz. Zool. Anz. 165: 393-400. 
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Gaz. 26: 277-287. 
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Manuscript accepted for publication 8 April 1987 


British Museum (Natural History) 

The birds of Mount Nimba, Liberia 

Peter R. Colston & Kai Curry-Lindahl 


For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
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archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
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human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland rain- 
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1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 


Titles to be published in Volume 54 


The cranial muscles and ligaments of macrouroid fishes (Teleostei: Gadiformes); 
functional, ecological and phylogenetic inferences. By Gordon J. Howes 

A review of the Macrochelidae (Acari: Mesostigmata) of the British Isles. 

By Keith H. Hyatt & Rowan M. Emberson 

A revision of Haplocaulus Precht, 1935 (Ciliophora: Peritrichida) and its 
morphological relatives. By Alan Warren 

Echinoderms of the Rockall Trough and adjacent areas. 3. Additional records. 

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Bulletin of the 

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A revision of Haplocaulus Precht, 1935 
(Ciliophora: Peritrichida) and its 
morphological relatives 

Alan Warren 


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v 

,f 

'.' 9 / MAY 1OQ ( 

A revision of Haplocaulus Precht, 1935 (Ciliophora: * 

Peritrichida) and its morphological relatives % 


Alan Warren 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 ^JBD '"Tirr"" * 


BRITISH MUSEUI 
Contents 

Q . I1T7 - 1JUW988 

Synopsis. .............. j^ 127 

Introduction . 
Genus Haplocaulus . 

Genus Baikalonis . . . L143 

Genus Cotensita ............. 145 

Genus Parazoothamnium . ........... 145 

Genus Piesika 147 

Genus Pseudohaplocaulus 147 

Incertae sedis 149 

References . . . . . . . . . . . . . .150 

Index to species 152 


Synopsis 

The species of Haplocaulus and of five closely related genera have been revised. A diagnosis for each genus is 
given with a key to its constituent species. Two new genera, Piesika n. gen. and Pseudohaplocaulus n. gen. are 
described. All extant species are described and figured. These include Haplocaulus of which 26 species are 
recognised; Baikaloms 3 species; Cotensita 1 species; Parazoothamnium 2 species; Piesika 1 species; 
Pseudohaplocaulus 2 species. Two other genera, Monintranstylum and Tucolesca, are also considered. 


Introduction 

In two previous papers (Warren, 1986, 1987) the peritrich genera Vorticella and Pseudovorticella 
were revised. Both are solitary forms the stalks of which coil up in a helical fashion upon contrac- 
tion. The genera dealt with in this paper are also solitary and borne on unbranched contractile 
stalks but, upon contraction, the stalks do not coil helically. 

Two of the principal generic characters used here are the mode of stalk contraction and the 
presence of pellicular tubercles with their underlying reticulate silverline system. Nevertheless for 
some species, details concerning these characters are not available. For example, with the excep- 
tion of Haplocaulus terrenus, no species of the genera reviewed have been impregnated with 
silver the possession of reticulate silverline systems has been assumed from the presence of 
pellicular tubercles. However, although pellicular tubercles are usually associated with reticulate 
silverline systems, this may not be true in every case (Foissner, pers. comm.). 

The major genus included here is Haplocaulus Precht, 1935 which is found in marine, fresh- 
water and terrestrial habitats attached to plant, animal and inanimate substrates. The only 
previous revision of Haplocaulus was that of Stiller (1971); seven new species have since been 
described and several species have been transferred from other genera. Baikalonis, Cotensita and 
Parazoothamnium, three genera closely related to Haplocaulus, are revised and two new genera, 
Piesika and Pseudohaplocaulus, are described. All extant species are described and figured and keys 
to their identification are provided. A brief account of the morphological structures important in 
the taxonomy of the Vorticellidae is given in Warren (1986). 


Bull. Br. Mus. nat. Hist. (Zool.) 54(3): 127-152 Issued 26 May 1988 

127 


128 A. WARREN 

Key to genera 

1 Zooid with pellicular tubercles ............. 2 

Zooid without pellicular tubercles 3 

2 Stalk contracts in two stages, initially shortening in a concertina-like fashion and then bending in a 

zigzag manner PIESIKA 

Stalk contracts in zigzag manner only PSEUDOHAPLOCAULUS 

3 Stalk sheath folded below zooid . COTENSITA 
Stalk sheath not folded below zooid 4 

4 Upon contraction stalk shortens longitudinally and is enveloped by zooid . BAIKALONIS 
Contraction does not result in zooid enveloping stalk 5 

5 Stalk contracts in two stages, initially shortening in a concertina-like fashion and then bending in a 

zigzag manner PARAZOOTHAMNWM 

Stalk contracts in zigzag manner only HAPLOCAULUS 

Genus HAPLOCA ULUS Precht, 1 935 

The genus Haplocaulus was erected by Precht ( 1 935) to include solitary vorticellids whose stalks are 
circular in cross-section and which contract in a zigzag rather than helical fashion. Impregnation 
by silver reveals a pattern of equally spaced horizontal lines or striations which encircle the body. 
These striations may or may not be visible in the living zooid. 

Two species were originally described by Precht (1935), H. nicoleae and H. furcellariae, 
although he failed to designate either as the type. H. nicoleae is here transferred to the genus 
Pseudohaplocaulus. H. furcellariae is designated the type species. Stiller (1971) transferred ten 
species to Haplocaulus all of which had previously belonged to the genus Vorticella. Three more 
vorticellids are here assigned to Haplocaulus for the first time. 

DIAGNOSIS. Zooid borne upon an unbranched stalk which is circular in cross-section and contracts in a zigzag 
manner. Zooids usually oval, cylindrical or inverted bell-shaped. Impregnation by silver reveals a transverse 
silverline system. Spasmoneme lies either parallel to the walls of the stalk sheath or is slightly twisted in the 
form of a shallow helix. 

Key to the species of Haplocaulus 

\ Diameter of peristomial lip greater than or equal to maximum body width . 

Diameter of peristomial lip less than maximum body width 15 

2 Pellicular striations visible on live zooid . 

Pellicular striations not visible on live zooid .10 

3 Zooid elongate and cylindrical in shape 4 

Zooid either cone- or inverted bell-shaped 8 

4 Pellicle with knob-like projections H.eforianus 

Pellicle without knob-like projections 5 

5 Zooid with two contractile vacuoles H. dipneumon 

Zooid with one contractile vacuole 6 

6 Zooid with distinct, broadly spaced striations H.fluviatilis 

Zooid with fine, narrow-spaced striations 

7 Macronucleus C-shaped and situated in anterior part of zooid .... H.procerus 
Macronucleus irregular and situated in centre of zooid .... H. furcellariae 

8 Zooid inverted bell-shaped with constriction beneath peristome 9 

Zooid cone-shaped, not constricted beneath peristome H. conosomus 

9 Macronucleus lies horizontally in anterior part of zooid H.epizoicus 

Macronucleus lies longitudinally in centre of zooid H. distinguendus 

10 Zooid elongate, length at least x 3 maximum body width .... 

Zooid length less than x 3 maximum body width .12 

1 1 Upon contraction, zooid assumes characteristic nodding position . . H. crassicaulis 
Upon contraction, zooid remains vertical ........ H. extensa 

12 Zooid with endosymbiotic zoochlorellae H. sertulariarum 

Zooid without endosymbiotic zoochlorellae .13 

1 3 Zooid cone-shaped, not constricted beneath peristome H. pelagicus 

Zooid inverted bell-shaped, usually with constriction beneath peristome 14 


HAPLOCAUL US MORPHOLOGICAL RELATIVES 


129 


14 Pellicular granules present 

Pellicular granules absent 

1 5 Pellicular striations visible on live zooid 

Pellicular striations not visible on live zooid 

16 Striations distinct and widely spaced 

Striations fine with narrow spacing 

17 Pellicle with concave ribbing between striations 

Pellicle with convex ribbing between striations 

18 Zooid elongate, length at least x 2 maximum body width .... 
Zooid rotund, length less than x 2 maximum body width .... 

19 Macronucleus U-shaped; spasmoneme extends full length of stalk 
Macronucleus C-shaped; spasmoneme does not extend complete stalk length 

20 Marine; epizoic on the echinoderm Amphiurae squamata .... 
Freshwater; epizoic on the amphibian Rana temporaria .... 

21 Zooid elongate, length at least x 2 maximum body width .... 
Zooid length less than maximum body width 

22 Macronucleus spirally twisted and lies longitudinally in zooid . 
Macronucleus C-shaped and lies horizontally in zooid .... 

23 Zooid 1 34 155 um long; epizoic on the amphibian Rana temporaria . 
Zooid 60 um long; epizoic on the crustacean Gammarus pulex . 

24 Macronucleus C-, S-, or irregular in shape; infundibulum reaches at least one- 
Macronucleus J-shaped; infundibulum less than one-third zooid length 

25 Macronucleus S-shaped or irregular; stalk about equal to body length 
Macronucleus C-shaped or irregular; stalk up to x 2 body length 


//. leanderi 
H. elegans 


H. terrenus 
H. kahili 


H.fusiformis 

H. claudlcans 

H. amphlurae 

H. longinucleatus 


H. macronucleatus 

H. amphiblarum 

H. stlllerl 

third zooid length . 

H. walteri 

H. brehml 

H. carinogammarl 


16 

21 
17 
18 


19 
20 


22 
24 

23 


25 


Species descriptions 
Haplocaulus furcellariae Precht, 1935 

DESCRIPTION (Fig. 1). The zooid of this, the type species is elongate, almost cylindrical in shape, approxi- 
mately 98 urn long x 40 um wide. Peristomial lip 50-55 um in diameter. Disc convex. Contractile vacuole 
small and situated in the peristomial region. Macronucleus elongate and irregular in shape. Pellicle with 
distinct transverse striations. Stalk up to 400 um long. 

HABITAT. Marine, originally found as an epizoite of Furcellaria. 


Fig. 1 Haplocaulus furcellariae, after Precht, 1935. Bar = 50 um. 


130 


A. WARREN 

Haplocaulus amphibiarum Banina, 1 982 


DESCRIPTION (Fig. 2). Zooid 134- 155 urn long x 52-65 um wide, tapering at both ends and widest in the 
central region. Diameter of peristomial lip less than maximum body width. Disc cone-shaped and prominent. 
Infundibulum reaches one-third body length. Contractile vacuole situated in upper half of zooid. 
Macronucleus C-shaped and lies horizontally across centre of body. Food vacuoles often numerous and 
spindle-shaped. Stalk less than body length. 

HABITAT. Freshwater, originally found near Leningrad, U.S.S.R. attached to the amphibian Rana 
temporaria. 


Fig. 2 Haplocaulus amphibiarum, after Banina, 1982. Fig. 3 Haplocaulus amphiurae, after Cuenot, 1891 
Bar = 50 um. (called Vorticella amphiurae) . Bar = 25 urn. 


Fig. 4 Haplocaulus brehmi, after Liipkes, 1 975. Bar = 50 urn. 


HAPLOCAULUS MORPHOLOGICAL RELATIVES 


131 


NOTE. H. amphibiarum bears a strong rr^rphological similarity to H. fusiformis (Nenninger, 1948) Stiller, 
1971 and to Carchesium amphibiarum Nenninger, 1948 which are also epibionts of amphibians. It is possible 
that these three species may be synonymous. 

Haplocaulus amphiurae (Cuenot, 1891) n. comb. 
Vorticella amphiurae Cuenot, 1891 

DESCRIPTION (Fig. 3). Zooid 40 urn long x 25-30 um wide, oval in shape, rounded posteriorly and with a 
narrow peristome. Contractile vacuole situated in upper one-third of body. Macronucleus C-shaped or 
occasionally irregular. Pellicle with fine transverse striations. Stalk less than body length. 

HABITAT. Marine, originally isolated from the Bay of Naples as an epizoite of the echinoderm Amphiurae 
squamata. 

NOTE. H. amphiurae is morphologically similar to Baikalonis. The original description of H. amphiurae, 
however, was made from partially contracted specimens. In order to determine the correct taxonomic 
position of this species, a redescription based on observations of healthy living specimens is required. 

Haplocaulus brehmi Liipkes, 1975 

DESCRIPTION (Fig. 4). Zooid oval in shape, 80 um long x 70 um wide. Peristomial lip well developed, 50 um in 
dhmeter. Disc slightly convex. Infundibulum broad and reaches two-thirds body length. Contractile vacuole 
situated just below the peristome. Macronucleus S-shaped or irregular, and lies longitudinally with respect to 
the major body axis. Pellicular striations not observed. Stalk about equal to body length. 

HABITAT. Freshwater, originally found attached to the gills of larvae of the caddis-fly Agapatus. 

Haplocaulus carinogammari Stiller ( 1 963), 1 97 1 
Vorticella carinogammari Stiller, 1963 

DESCRIPTION (Fig. 5). Zooid inverted bell-shaped, rounded, 45-75 um long. Peristomial lip well developed, 
diameter about equal to maximum body width. Disc convex. Infundibulum narrow and reaches centre of 
zooid. Contractile vacuole situated just beneath peristome. Macronucleus elongate and irregular in shape. 
Pellicular striations visible on fixed specimens. Stalk up to x 2 body length. 

HABITAT. Freshwater, originally found attached to the crustaceans Carinogammarus roeselii var triacanthus 
and Gammarus fossarum; also occurs on Rivulogammarus (Piesik, 1975) and Asellus aquaticus 
(Szczepanowski, 1978). 


Fig. 5 Haplocaulus carinogammari (a) contracted; (b) relaxed, after Stiller, 1963 (called Vorticella 
carinogammari); (c) & (d) showing variation of macronucleus, after Szczepanowski, 1978. 
Bar = 50um. 


132 


A. WARREN 


Fig. 6 Haplocaulus claudicans, (a) relaxed zooid; (b) contracted zooid; (c) detail of stalk, after Penard, 

1922 (called Vorticella claudicans). Bar = 25 urn. 


Fig. 7 Haplocaulus conosomus, (a) after Stokes, 1 889 (called Vorticella conosoma) ; (b) relaxed zooid; (c) 
contracted zooid, after Gajewskaja, 1933 (called Vorticella conesoma). Bar = 50 urn. 


Haplocaulus claudicans (Penard, 1922) Stiller, 1971 
Vorticella claudicans Penard, 1922 

DESCRIPTION (Fig. 6). Zooid elongate, almost cylindrical in shape, 40-55 urn long x 30 urn wide. Diameter of 
peristomial lip less than or occasionally equal to maximum body width. Contractile vacuole situated in upper 
one-third of body. Macronucleus C-shaped and lies horizontally across centre of zooid. Pellicle with fine 
transverse striations. Stalk about equal to body length. Spasmoneme terminates about half-way down the 
stalk and is connected to the base of the stalk by a fibre. 


HAPLOCAULUS MORPHOLOGICAL RELATIVES 


133 


Fig. 8 Haplocaulus crassicaulis, (a) relaxed zooid; (b) contracted zooid, composite after Kent, 1881 
(called Vorticella crassicaulis) and Nenninger, 1948. Bar = 25 um. 


HABITAT. Freshwater, originally found attached to mosses. 

NOTE. It is unclear whether the 'fibre' in the lower part of the stalk is an extension of the spasmoneme, or a 
separate structure. In the case of the latter, it may be necessary to transfer this species to the genus 
Monintranstylum. 

Haplocaulus conosomus (Stokes, 1889) n. comb. 

Vorticella conosoma Stokes, 1889 
Vorticella conesoma Gajewskaja, 1933 

DESCRIPTION (Fig. 7). Zooid conical and elongate, 75 um long x 30 um wide, and with a distinct ridge in the 
region of the telotroch band. Peristomial lip 35 urn in diameter. Contractile vacuole situated just beneath 
peristome. Macronucleus C-shaped or irregular, and situated either centrally or in the anterior part of the 
body. Pellicular striations visible on zooid. Stalk 150-200 um long. 

HABITAT. Freshwater. 

NOTE. In his original description, Stokes (1889) did not describe the spasmoneme. According to Gajewskaja 
(1933), however, the spasmoneme terminates in the upper part of the stalk. If spasmoneme length is accepted 
as a generic character, it may be necessary to transfer this species to the genus Monintranstylum. 

Haplocaulus crassicaulis (Kent, 1881) Stiller, 1971 
Vorticella crassicaulis Kent, 1881 

DESCRIPTION (Fig. 8). Zooid elongate, 45-50 um long x 25 um wide. Peristomial lip 20 um in diameter. 
Contractile vacuole situated in upper one-third of body. Macronucleus C-shaped and situated in centre of 
zooid. Stalk x 1 x 2 body length. Stalk sheath has several transverse folds. 

HABITAT. Freshwater, originally isolated as an epizoite of the crustacean Asellus aquaticus. 


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Fig. 9 Haplocaulus dipneumon, after Penard, 1922 (called Vorticella dipneumon). Bar = 25 um. 


Fig. 10 Haplocaulus distinguendus , (a) after Sommer, 1951; (b) after Bierhof & Roos, 1976 (called 
Haplocaulus distinguendis ) ; (c) after Liipkes, 1976 (called Haplocaulus hengsti). Bar = 50 um. 


Haplocaulus dipneumon (Penard, 1922) Stiller, 1971 
Vorticella dipneumon Penard, 1922 

DESCRIPTION (Fig. 9). Zooid almost cylindrical in shape, 50-56 urn long x 25 urn wide. Peristomial lip well 
developed, 25-30 urn in diameter. Disc convex and raised centrally to a point. Infundibulum almost reaches 
centre of zooid. Two contractile vacuoles situated in upper one-third of body. Macronucleus elongate and lies 


HAPLOCA ULUS MORPHOLOGICAL RELATIVES 135 

longitudinally with respect to the major body axis. Transverse striations visible on pellicle. Stalk less than 
body length. Spasmoneme flared at either end. 

HABITAT. Freshwater, originally found as an epizoite of the crustacean Gammarus pulex. 

NOTE. This species bears a strong morphological resemblance to Baikalonis. Until its mode of contraction has 
been described, however, H. dipneumon should remain in the genus Haplocaulus. 

Haplocaulm distinguendus Sommer, 1951 

Haplocaulus distinguen dis (Sommer, 1951) Bierhof & Roos, 1976 
H. hengsti Liipkes, 1976 

DESCRIPTION (Fig. 10). Zooid ovoid, 60-79 um long x 3449 um wide, and constricted beneath peristome. 
Peristomial lip well developed, 27-40 um in diameter. Disc convex. Infundibulum broad and reaches almost 
to centre of body. Contractile vacuole situated in upper half of zooid. Macronucleus irregular or C-shaped. 
Transverse pellicular striations visible on zooid. Stalk up to 190 um long x 7-0-9-0 um wide. 

HABITAT. Freshwater, attached to the river weed Enteromorpha intestinalis (Sommer 1951), to inanimate 
substrates (Liipkes, 1976), and to the crustaceans Asellus aquations (Sommer, 1951) and Gammarus spp. 
(Bierhof & Roos, 1976). 

NOTE. H. hengsti is synonymysed with H. distinguendus because of their morphological similarity. The main 
differences between the two are the size of the zooid, H. hengsti (79 jim) being slightly longer than H. 
distinguendus (60-75 um), and stalk length (H. distinguendus up to 190 um long: H. hengsti less than body 
length). Neither of these characters are particularly reliable for separating species, and there are insufficient 
differences here to recognise the two as separate taxa. 

Haplocaulus eforianus (Tucolesco, 1962) n. comb 
Vorticella eforiana Tucolesco, 1 962 

DESCRIPTION (Fig. 1 1). Zooid 55-60 um long x 20 um wide. Upper part of body cylindrical in shape, lower 
part conical. Peristomial lip 20-25 urn in diameter. Pellicle with transverse striations, and convex ribbing 
between the striations. Pellicle also ornamented with numerous knob-like projections. Stalk x 1-x 2 body 
length. 

HABITAT. Marine 


Fig. 11 Haplocaulus eforianus, (a) relaxed zooid; (b) contracted zooid; (c) pellicle, showing convex 
ribbing and knob-like projections, after Tucolesca, 1962 (called Vorticella eforiana). Bar = 25 um. 


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Fig. 12 Haplocaulus elegans (a) showing macronucleus; (b) relaxed zooid, after Nenninger, 1948 (called 
Vorticella elegans); (c) after Szczepanowski, 1978. Bar = 50 urn. 


Fig. 13 Haplocaulus epizoicus (a) after Sramek-Husek, 1948 (called Vorticella epizoica), bar = 25 urn; 

) after Piesik, 1975, bar = 25 urn. 


Haplocaulus elegans (Wenninger, 1948) Stiller, 1971 

Intranstylum elegans Nenninger, 1948 
Haplocaulus elegans f. gammari Piesik, 1975 

DESCRIPTION (Fig. 1 2). Zooid 42-84 um long, constricted beneath the peristome and tapering posteriorly 
towards the stalk. Diameter of peristomial lip less than maximum body width. Disc convex. Contractile 
vacuole situated in upper one-third of zooid. Macronucleus C-shaped and lies horizontally in the anterior part 
of the body. Pellicular striations not observed. Stalk x 1- x 2 body length. 

HABITAT. Freshwater, originally found attached to the crustaceans Asellus aquaticus and Cyclops sp. 


HAPLOCAULUS MORPHOLOGICAL RELATIVES 


137 


Fig. 14 Haplocaulus extensus (a) relaxed zooid; (b) contracted zooid, after Kahl, 1 935 (called Vorticella 

extensa) . Bar = 50 urn. 


Haplocaulus epizoicus (Sramek-Husek, 1948) Stiller, 1971 
Vorticella epizoica Sramek-Husek, 1948 

DESCRIPTION (Fig. 13). Zooid 25-46 um long x 18-37 um wide, inverted bell-shaped and rounded posteriorly. 
Diameter of peristomial lip about equal to maximum body width. Contractile vacuole situated just beneath 
peristome. Macronucleus C-shaped and lies either horizontally or obliquely in the anterior part of the zooid. 
Pellicle with fine transverse striations. Stalk x 1- x 2 body length. 

HABITAT. Freshwater, found as an epizoite of the crustaceans Megacyclopsis viridis (Sramek-Husek, 1948) 
and Gammamspulexfossarum (Piesik, 1975). 

Haplocaulus extensus (Kahl, 1935)Sommer, 1951 
Vorticella extensa Kahl, 1935 

DESCRIPTION (Fig. 14). Zooid elongate, 50-100 um long xl 5-27 urn wide (Kahl (1935) described two 
morphological forms of this species, one 90-100 urn the other 50-70 urn long). Peristomial lip 35 um in 
diameter. Contractile vacuole situated just beneath peristome. Macronucleus C-shaped and lies transversely 
in anterior part of zooid. Pellicle with fine transverse striations. Stalk up to 200 um long. 

HABITAT. Freshwater, found by Sommer (1951) attached to the river weed Enteromorpha intestinalis. 

Haplocaulus fluviatilis Shubernetskij & Chorik, 1977 

DESCRIPTION (Fig. 15). Zooid barrel-shaped with broadened scopular region, 28-48 um (mean 38 um) 
long x 14-30 jim (mean 22 um) wide. Peristomial lip 17-25 um (mean 21 um) in diameter. Disc elevated at 
angle to peristome and raised centrally to a point. Infundibulum reaches one-third body length. Contractile 
vacuole situated just beneath peristome. Macronucleus C-shaped and lies horizontally across upper part of 
zooid. Pellicle with distinct transverse striations, and convex ribbing between the striations. Stalk up to 
50 um long. 

HABITAT. Freshwater, originally found attached to the ostracod Heterocypris. 


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A. WARREN 


Fig. 15 Haplocaulusfluviatilis, after Shubernetskij & 
Chorik, 1977. Bar = 25 urn. 


Fig. 16 Haplocaulus fusiformis, after Nenninger, 
1948 (called Vorticellafusiforma). Bar = 75 um. 


Fig. 17 


Haplocaulus kahlii (a) & (b) after Stiller, 1931 (called Vorticella kahlii) ; (c) after Bierhof & Roos, 

1976. Bar = 25 urn. 


Haplocaulus fusiformis (Nenninger, 1948) Stiller, 1971 
Vorticellafusiforma Nenninger, 1948 

DESCRIPTION (Fig. 16). Zooid 140-162 um long x 60 um wide, spindle-shaped with prominent central bulge. 
Peristomial lip 45 um in diameter. Disc prominent and conical in shape. Infundibulum reaches one-quarter 
body length. Contractile vacuole situated in upper one-third of zooid. Macronucleus U-shaped and lies in 
anterior half of body. Food vacuoles large, dark and hexagonal in shape. Fine transverse striations visible on 
pellicle. Stalk less than body length. 

HABITAT. Freshwater, originally found attached to the tails of tadpoles. 

NOTE. Food vacuoles are not accepted as a taxonomic character among the Vorticellidae (Noland & Finley, 
1931; Warren, 1986) or, indeed, the rest of the phylum Ciliophora. 


HAPLOCAULUS MORPHOLOGICAL RELATIVES 

Haplocauluskahlii (Stiller, 1931) Sommer, 1951 


139 


yorticellakahlii Stiller, 1931 

DESCRIPTION (Fig. 1 7). Zooid pyriform with prominent bulge in anterior region, 32-44 um long x 24-36 um 
wide. Peristomial lip well developed, occasionally with central furrow giving the appearance of a double lip. 
Diameter of lip less than maximum body width. Macronucleus elongate, variable in shape. Pellicle distinctly 
striated with convex ribbing between striations. Stalk equal to or less than body length. 

HABITAT. Freshwater, found as an epizoite of the crustaceans Leptodora kindlii, Daphnia longispina var 
hyalina and Salpinia sp. (Stiller, 1931), Megacyclopsis viridis (Sramek-Husek, 1948), and Gammarus tigrinus 
(Bierhof & Roos, 1976). 

Haplocaulus leanderi Stiller, 1968 

DESCRIPTION (Fig. 18). Zooid inverted bell-shaped, 30-32 um long x 22-24 um wide. Peristomial lip well 
developed, 26-28 um in diameter. Infundibulum reaches one-third body length. Macronucleus C-shaped and 
lies horizontally in anterior half of body. Pellicle furnished with numerous small granules. Stalk x 1- x 2 body 
length. 

HABITAT. Marine, originally isolated as an epizoite of the crustacean Leander sp. 

Haplocaulus longinuclei Banina, 1982 

DESCRIPTION (Fig. 19). Zooid 43-61 um long x 41-49 um wide, somewhat variable in shape but usually oval 
or spherical and tapering sharply towards the stalk. Diameter of peristomial lip less than maximum body 
width. Infundibulum reaches centre of zooid. Macronucleus elongate and irregular in shape. Fine transverse 
striations visible on pellicle. Stalk less than body length. 

HABITAT. Freshwater, originally isolated as epizoites of tadpoles of the amphibian Rana temporaria. 

Haplocaulus macronucleatm (Nenninger, 1948) Stiller, 1971 
Vorticella extensa var macronucleata Nenninger, 1948 

DESCRIPTION (Fig. 20). Zooid elongate almost cylindrical in shape, 91-5 um long x 35 um wide. Diameter of 
peristomial lip about equal to maximum body width. Disc convex and raised centrally to a point. Contractile 
vacuole small and situated in upper one-third of body. Macronucleus broad, S-shaped and lies longitudinally 
with respect to major body axis. Pellicular striations not visible. Stalk about equal to body length. 

HABITAT. Freshwater, originally isolated as an epizoite of the hexapod Ephemera vulgata. 


Fig. 18 Haplocaulus leanderi, after Stiller, 
Bar = 25um. 


1 968. Fig. 19 Haplocaulus longinuclei (a) relaxed zooid; (b) 
contracted zooid, after Banina, 1982. Bar = 25 um. 


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A. WARREN 


Fig. 20 Haplocaulus macronucleatus, after 
Nenninger, 1948 (called Vorticella extensa var. 
macronucleata) . Bar = 50 urn. 


Fig. 21 Haplocaulus pelagicus, after Gajewskaja, 
1933 (called Vort icella pelagica) . Bar = 25 um. 


Fig. 22 Haplocaulus procerus , after Nenninger, 1948 
(called Vorticella procera). Bar = 50 urn. 


Fig. 23 Haplocaulus sertulariarum, after Entz, 1884 
(called Spastostyla sertulariarum). Bar = 50 urn. 


Haplocaulus pelagicus (Gajewskaja, 193 3) Stiller, 1971 
Vorticella pelagica Gajewskaja, 1933 

DESCRIPTION (Fig. 21). Zooid conical or inverted bell-shaped, 30-40 um long x 50 urn wide. Peristomial lip 
70 um in diameter. Contractile vacuole situated just below peristome. Macronucleus elongate, irregular in 
shape and lies longitudinally with respect to major body axis. Pellicular striations not visible. Stalk usually less 
than body length. 

HABITAT. Marine 


HAPLOCA UL US MORPHOLOGICAL RELATIVES 1 4 1 

Haplocaulm procerus (Nenninger, 1948) Stiller, 1971 
Vorticella procera Nenninger, 1948 

DESCRIPTION (Fig. 22). Zooid elongate, almost cylindrical in shape, 103 um long x 40 um wide. Peristomial lip 
50 ^m in diameter. Disc flat. Contractile vacuole situated just beneath peristomial lip. Macronucleus C- 
shaped and lies either horizontally or obliquely in the anterior part of the zooid. Food vacuoles typically large. 
Fine transverse striations visible on pellicle. Stalk x 2- x 3 body length. 

HABITAT. Freshwater, originally found attached to the tails of tadpoles. Typically forms pseudocolonies. 
Haplocaulus sertulariarum (Entz, 1884)Banina, 1982 

Spastostyla sertulariarum Entz, 1884 
Vorticella sertulariarum (Entz, 1884)Kahl, 1935 

DESCRIPTION (Fig. 23). Zooid inverted bell-shaped, 60-109 um long x 34-48 um wide. Diameter of 
peristomial lip greater than maximum body width. Disc flat. Infundibulum reaches centre of zooid. 
Contractile vacuole situated just beneath peristome. Macronucleus C-shaped and lies horizontally across 
centre of anterior part of zooid. Zoochlorellae may be present in cytoplasm. Stalk broad, and equal to or less 
than body length. Stalk sheath frequently with distinct transverse folds. 

HABITAT. Freshwater or marine, originally isolated from the Bay of Naples attached to polyps and sea urchins 
(Entz, 1884); also found as epizoites of tadpoles ofRana temporaria. 

NOTE. Entz (1884) considered this organism to be identicle to Rhabdostyla sertularium Kent, 1881. 
Rhabdostyla sertularium, however, does not possess a spasmoneme and the stalk is non-contractile whereas 
Entz's organism clearly has a spasmoneme. S. sertulariarum Entz, 1884 was transferred to Vorticella by Kahl 
(1935) and, more recently, to Haplocaulus by Banina (1982). 

Haplocaulus stilleri Piesik, 1975 

DESCRIPTION (Fig. 24). Zooid spindle-shaped, 60 um long x 27 um wide, with prominent central bulge. 
Peristomial lip 20 um in diameter. Disc convex. Contractile vacuole situated just beneath peristome. Macro- 
nucleus C-shaped and lies horizontally across centre of zooid. Pellicular striations not observed. Stalk up to 
60 um long and 12 um wide. 

HABITAT. Freshwater, originally described as an epibiont attached to the crustaceans Gammarus pulex pulex 
(L) and Gammarus pulex fossarum (Koch). 


Fig. 24 Haplocaulus stilleri, after Piesik, 1 975. Bar = 50 um. 


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A. WARREN 


Fig. 25 Haplocaulus terrenus, showing variability in relaxed and contracted zooids, (a) relaxed; (b) 
contracted, bar = 25 ^irn; (c) relaxed, bar = 25 ^im; (d) contracted, bar= 15 nm, after Foissner, 1981. 


Fig. 26 Haplocaulus walteri, (a) typical zooid; (b & c) variations in zooid shape under poor conditions; 

(d) telotroch, after Guhl, 1985. Bar = 25 urn. 


HAPLOCA ULUS MORPHOLOGICAL RELATIVES 1 43 

Haplocaulus terrenm Foissner, 1 98 1 

DESCRIPTION (Fig. 25). Zooid somewhat variable in shape, 40-60 um long x 20-26 nm wide. Peristomial lip 
20 urn in diameter. Infundibulum reaches centre of zooid. Contractile vacuole large, situated in anterior part 
of zooid and empties into infundibulum via a short channel. Macronucleus C-shaped and lies either horizon- 
tally or obliquely across centre of body. Zooid has 30 transverse striations with concave ribbing between 
striations. Stalk 2-5-3-5 um wide, and one-half x 3 body length. Upon contraction posterior end of zooid 
may become folded, overlapping the stalk. 

HABITAT. Terrestrial, originally isolated from Alpine soils. 

Haplocaulus vealteri Guhl, 1985 

DESCRIPTION (Fig. 26). Zooid 55-75 um long x 40-60 um wide, cylindrical to inverted bell-shaped. Diameter 
of peristomial lip usually less than maximum body width. Infundibulum short. Contractile vacuole situated 
just below peristome. Macronucleus J- or 7-shaped. Pellicular striations not observed. Stalk up to 158 um 
long, and with rod-shaped bacteria attached to the stalk sheath. 

HABITAT. Freshwater, originally found in activated sludge. 

Genus BAIKALONIS Jankowski, 1982 

The main distinguishing feature of this genus is the mode of contraction during which the stalk 
shortens longitudinally and is enveloped by the zooid. When Baikalonis was first described 
(Jankowski, 1982) just one species, B.foissneri, was known. B.foissneri thus became the type 
species by monotypy. Two more species, Vorticella undulata (Dons, 1918) Noland & Finley, 1931 
and Haplocaulus sp. Bierhof & Roos, 1976 are here transferred to Baikalonis. Three other species of 
Haplocaulus, H. amphiurae, H. dipneumon and H.fusca, also bear a strong morphological resemb- 
lance to Baikalonis although their modes of contraction have yet to be described. 

DIAGNOSIS. Solitary, borne upon an unbranched stalk which is typically much shorter than the 
body. Spasmoneme straight and extends the entire length of the stalk. Upon contraction, the zooid 
envelopes the stalk which shortens longitudinally. 

Key to the species of Baikalonis 

1 Freshwater; zooid with a contractile vacuole 2 

Marine; zooid without a contractile vacuole B. undulata 

2 Infundibulum broad and reaches centre of zooid. Contractile vacuole centrally located. 

Peristomial lip with central furrow B.foissneri 

Infundibulum lies diagonally and reaches one-third body length. Contractile vacuole situated just 
below peristome. Peristomial lip without a furrow B. gammari 

Species descriptions 
Baikalonis foissneri Jankowski, 1982 

DESCRIPTION (Fig. 27). This the type species is cylindrical in shape, 56-60 um long x 20 um wide. Peristomial 
lip well developed with central furrow giving the appearance of a double lip. Disc prominently arched above 
peristome. Infundibulum broad and reaches centre of zooid. Contractile vacuole situated centrally in body. 
Macronucleus cylindrical in shape and slightly curved. Stalk 12-14 um long. Spasmoneme broad. 

HABITAT. Freshwater, originally found as an epizoite of the larvae of the caddis-fly Baicalina bellicosa in Lake 
Baikal, U.S.S.R. 

Baikalonis gammari n. sp. 

Haplocaulus sp. Bierhof & Roos, 1976 

DESCRIPTION (Fig. 28). Zooid cylindrical in shape, narrowing slightly towards the stalk, 75 um long x 45 um 
wide. Peristomial lip well developed, diameter about equal to maximum body width. Infundibulum reaches 
one-third body length. Contractile vacuole situated just below peristome. Macronucleus C-shaped and lies 
longitudinally in centre of zooid. Fine transverse striations visible on pellicle. Stalk 10 um long x 8 um wide. 

HABITAT. Freshwater, originally isolated as an epizoite of the crustacean Gammarus pulex. 


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A. WARREN 


Fig. 27 Baikalonis foissneri, (a) relaxed; (b) 
contracted, after Jankowski, 1982. Bar = 25 um. 


Fig. 28 Baikalonis gammari, after Bierhof & Roos, 
1976 (called Haplocaulus sp.). Bar = 50 um. 


Fig. 29 Baikalonis undulata, (a) relaxed; (b) contracted, after Dons, 1918 (called Vorticellopsis 

undulata) . Bar = 50 um. 


Baikalonis undulata (Dons, 1918) n. comb. 

Vorticellopsis undulata Dons, 1918 

Vorticella undulata (Dons, 1918)Noland& Finley, 1931 

DESCRIPTION (Fig. 29). Zooid pyriform, 1 10 um long x 72 um wide. Peristomial lip 57 um in diameter. Two 
distinct constrictions present on body, one beneath the peristomial lip and the other in the region of the 
telotroch band. Contractile vacuole not observed. Macronucleus C-shaped, 30 um long x 20 nm wide and lies 
longitudinally with respect to major body axis. Transverse striations clearly visible on pellicle. Stalk 1 15 um 
long x 19 um wide. Spasmoneme 12 um in diameter. 

HABITAT. Marine, originally found attached to the alga Desmaresita viridis. 


HAPLOCAULUS MORPHOLOGICAL RELATIVES 


145 


NOTE. Both Noland & Finley (1931) and Kahl (1935) considered that Vorticellopsis undulata Dons, 1918 
should belong to the genus Vorticella. In a recent revision of Vorticella, Warren (1986) noted that the stalk of 
V. undulata does not contract spirally and should not, therefore, be included in the genus Vorticella. The mode 
of contraction and the morphology of the stalk both indicate that this species should belong to the genus 
Baikalonis. 

Genus COTENSITA Jankowski, 1982 

The main distinguishing feature of the genus Cotensita is the stalk sheath which is folded in a 
characteristic fashion just beneath the zooid. Although some colonial peritrichs such as Craspedo- 
myoschiston have ornamentation of the stalk sheath, this feature has not previously been recorded 
among solitary vorticellids. The spasmoneme of Cotensita is straight and extends to about two- 
thirds of the stalk length. C. commensalis Jankowski, 1982 is the type species by monotypy. 

DIAGNOSIS. Spasmoneme straight, not extending the full length of the stalk. Stalk sheath twisted in 
the region just below the stalk. Upon contraction the zooid bends over to one side and assumes a 
characteristic 'nodding' position. 

Species description 

Cotensita commensalis Jankowski, 1982 

DESCRIPTION (Fig. 30). This the type species is inverted bell-shaped, 46 um long x 36 um wide. Diameter of 
peristomial lip about equal to maximum body width. Contractile vacuole situated just beneath peristome. 
Macronucleus C-shaped. Stalk 85-90 um long. Stalk sheath twisted to form 1-2 helical coils just below zooid. 
Spasmoneme reaches two-thirds stalk length with the posterior end tapering to a point. 

HABITAT. Freshwater, originally isolated as an epizoite of the larvae of the caddis-fly Baicalina bellicosa from 
Lake Baikal, U.S.S.R. 

Genus PARAZOOTHAMNIUMPiesik, 1975 

The main distinguishing feature of this genus is the mode of stalk contraction which takes place in 
two stages: initially the stalk shortens longitudinally in a concertina-like fashion but, if the stimulus 
is sufficiently strong, it loops and bends in the more usual manner. 


Fig. 30 Cotensita commensalis, (a) relaxed zooid, bar = 25 um. (b) partially contracted; (c) detail of 

stalk, after Jankowski, 1982. 


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A. WARREN 


Fig. 31 Parazoothamnium stenotica, after Piesik, 
1975. Bar = 50 urn. 


Fig. 32 Parazoothamnium claparedei, (a) relaxed 
zooid; (b) contracted, after Andrussowa, 1886 
(called Vorticella claparedei). Bar = 50 nm. 


Fig. 33 Piesika gammari, after Piesik, 1975 (called Parazoothamnium gammari) . Bar = 50 urn. 


Piesik (1975) originally described two species of Parazoothamnium, P. stenotica and P. gammari. 
The latter species, however, has rows of regularly arranged tubercles and, since this is recognised as 
a generic character among the Vorticellidae, P. gammari is transferred to the new genus Piesika. P. 
claparedeiis included in the genus Parazoothamnium for the first time. P. stenotica is designated the 
type species. 

DIAGNOSIS. Solitary, borne upon an unbranched stalk. Spasmoneme extends the complete length 
of the stalk. Stalk contraction takes place in two stages; initially it shortens longitudinally in a 


HAPLOCA ULUS MORPHOLOGICAL RELATIVES 1 47 

concertina-like fashion but, if the stimulus is sufficiently strong, it loops and folds. Zooid has a 
transverse silverline system. 

Key to the species of Parazoothamnium 

1 Macronucleus J-shaped; contractile vacuole situated in anterior half of body . P. stenotica 
Macronucleus C-shaped; contractile vacuole situated in posterior half of body . P. claparedei 

Species descriptions 

Parazoothamnium stenotica Piesik, 1975 

DESCRIPTION (Fig. 31). This the type species is inverted bell-shaped, somewhat rotund, 70 um long x 50 um 
wide. Peristomial lip well developed, diameter about equal to maximum body width. Contractile vacuole 
situated just beneath peristome. Macronucleus J-shaped. Pellicle with fine transverse striations. Stalk up to 
9-0 um wide. 

HABITAT. Freshwater, originally isolated as an epizoite of the crustacean Gammarus pulex. 

Parazoothamnium claparedei (Andrussowa, 1886) n. comb. 
Vorticella claparedei Andrussowa, 1 886 

DESCRIPTION (Fig. 32). Zooid campanulate, 85 urn long x 60 um wide and with a distinct swelling just below 
the peristome. Peristomial lip 60 um in diameter and with a central furrow to give the appearance of a double 
lip. Disc raised obliquely above peristome. Infundibulum reaches centre of zooid. Single contractile vacuole 
situated in posterior half of zooid. Macronucleus C-shaped and lies longitudinally with respect to major body 
axis. Pellicular striations not observed. Upon contraction stalk sheath assumes a wrinkled appearance and 
stalk coils into two or three unevenly spaced loops. 

HABITAT. Marine, attached to filamentous algae. 

Genus PIESIKA n. gen. 

The possession of regularly arranged pellicular tubercles along with an underlying reticulate 
silverline system is regarded as a generic character among peritrichs (Foissner & SchifTmann, 1 974; 
Warren, 1986). The genus Piesika is erected to include solitary vorticellids which both possess 
pellicular tubercles, and exhibit the two stage stalk contraction process seen in Parazoothamnium. 
P. gammari is the type species by monotypy. 

DIAGNOSIS. Solitary zooids borne upon an unbranched stalk. Spasmoneme extends the complete 
length of the stalk. Contraction of the stalk takes place in two stages; initially the stalk shortens 
longitudinally in a concertina-like fashion but, if the stimulus is sufficiently strong, it loops and 
folds. Zooid with rows of regularly aligned pellicular tubercles. 

Species description 

Piesika gammari (Piesik, 1975) n. comb. 
Parazoothamnium gammari Piesik, 1 975 

DESCRIPTION (Fig. 33). This the type species is inverted bell-shaped, 70 um long x 50 um wide. Diameter of 
peristomial lip about equal to maximum body width. Disc convex. Contractile vacuole situated just below 
peristome. Macronucleus elongate and irregular in shape. Stalk up to 240 um long x 10 um wide. Cysts nearly 
spherical in shape, 49 um x 43 um. 

HABITAT. Freshwater, originally found as an epizoite of the crustacean Gammarus pulex pulex. 

Genus PSEUDOHAPLOCA ULUS n. gen. 

Foissner & SchifTmann ( 1 974) reported that two types of silverline system are found among 
vorticellids; (i) transverse, in which the lines encircle the body in one direction only, and (ii) 
reticulate, which consists of a network of vertical and horizontal lines. Furthermore, it has been 
established that vorticellids with regularly arranged rows of pellicular tubercles also possess reticu- 
late silverline systems (Foissner, 1979, 1981; Carey & Warren, 1983; Warren, 1987). The genus 
Pseudohaplocaulus is erected to include Haplocaulus-\ike peritrichs which possess rows of regularly 
aligned pellicular tubercles and, therefore, reticulate silverline systems. There are two species of 


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A. WARREN 


Fig. 34 Pseudohaplocaulus nicoleae, after Precht, 
1935 (called Haplocaulus nicoleae). Bar = 25 um. 


Fig. 35 Pseudohaplocaulus anabaenae, after Stiller, 
1940 (called Vorticella anabaenae). Bar = 25 um. 


Pseudohaplocaulus, both formerly belonging to the genus Haplocaulus. Pseudohaplocaulus nicoleae 
(Precht, 1935) n. comb, is the type species. 

DIAGNOSIS. Solitary, borne upon an unbranched stalk that is circular in cross-section and contracts 
in a zigzag manner. Zooid with rows of regularly aligned pellicular tubercles and a reticulate 
silverline system. 

Key to species of Pseudohaplocaulus 

1 Marine. Zooid with one contractile vacuole ....... P. nicoleae 

Freshwater. Zooid with two contractile vacuoles P. anabaenae 

Species descriptions 

Pseudohaplocaulus nicoleae (Precht, 1935) n. comb. 
Haplocaulus nicoleae Precht, 1935 

DESCRIPTION (Fig. 34). This the type species is inverted bell-shaped, 35-40 urn long x 25 um wide. Peristomial 
lip 30 urn in diameter. Contractile vacuole situated just below the peristome. Macronucleus J-shaped. Stalk 
x 1- x 2 body length. Sites of previous zooid division may be visible as short lateral extensions of the stalk 
sheath (see Fig. 34). 

HABITAT. Marine, originally found as an epizoite of the polychaete, Nicolea zostericola. 
Pseudohaplocaulus anabaenae (Stiller, 1940) n. comb. 

Vorticella anabaenae Stiller, 1940 
Haplocaulus anabaenae Stiller, (1940) 1971 

DESCRIPTION (Fig. 35). Zooid inverted bell-shaped, 40-45 urn long x 40 um wide. Peristomial lip 45 urn in 
diameter. Disc flat. Infundibulum reaches one-third body length. Two contractile vacuoles, one situated 
near the base of the infundibulum, the other just below the peristome. Macronucleus J-shaped. Stalk x 1- x 2 
body length. 

HABITAT. Freshwater, originally isolated as an epibiont attached to the Cyanobacterium Anabaena. 


HAPLOCA ULUS MORPHOLOGICAL RELATIVES 1 49 

Incertae Sedis 
Genus MONINTRANSTYLUM Banina, 1977 

Monintranstylum is a solitary peritrich the stalk of which contracts in a zigzag rather than helical 
fashion. The main distinguishing feature of Monintranstylum is the spasmoneme which is short and 
terminates above the base of the stalk, unlike that of Haplocaulus and other related genera where 
the spasmoneme is usually about the same length as the stalk. 

Spasmoneme length has long been recognised as a generic character among colonial peritrichs. 
Intranstylum Faure-Fremiet, 1904, for example, differs from Carchesium Ehrenberg, 1830 by the 
absence of the spasmoneme in the central trunk of the stalk. Myoschiston Precht, 1935 may 
similarly be recognised from Zoothamnium Bory, 1826. Among the solitary peritrichs, however, 
the situation is less clear and several species belonging to genera in which the whole stalk is 
normally contractile, have short or otherwise incomplete spasmonemes. The spasmoneme of 
Vorticella intermissa Nenninger, 1948, for example, begins a short distance below the scopula and 
is therefore absent in the uppermost part of the stalk; and in Haplocaulus claudicans a small 'fibre' 
connects the short spasmoneme to the stalk base (see p. 132). Furthermore, the author has fre- 
quently observed both solitary and colonial peritrichs in which the spasmoneme appears to have 
partially degenerated and lost its contractility (unpublished data). Clearly, further research is 
necessary before spasmoneme length can be accepted as a generic character among solitary 
peritrichs. 

NOTE. The name Monintranstylum was first mentioned in an abstract (Banina, 1976). A full 
description of Monintranstylum, however, did not appear until the following year (Banina, 1977a), 
the date from which the genus should properly be recognised. The first named species of 
Monintranstylum were described in a second article later in the same journal (Banina, 1977b). No 
type species was designated. 

DIAGNOSIS. Zooid borne upon a stalk which contracts in a zigzag manner. Solitary, never colonial, 
and carried upon an unbranched stalk. Spasmoneme present only in the upper part of the stalk. 

Species of Monintranstylum 

Three species of Monintranstylum were described by Banina (19776), M. rotundus, an epibiont of Daphnia 
longispina; M. stammeri, an epibiont of Polyphemus pediculus, Daphnia pulex, Ceriodaphina quadrangulata 
and Cyclops sp.; and M. sommeri on the abdomen of Eucydops serrulatus and the shell of Cypris sp. The 
original descriptions of these species were not available for this revision. 

Banina (1982) also transferred Intranstylum ranae Stiller, 1953 to the genus Monintranstylum on the basis 
that /. ranae is solitary rather than colonial, and that its spasmoneme terminates above the base of the stalk. 
However, for the reasons given above, M. ranae and the three other species of Monintranstylum may perhaps 
more properly belong to the genus Haplocaulus. 

Monintranstylum ranae (Stiller, 1953) Banina, 1982 
Intranstylum ranae Stiller, 1953 

DESCRIPTION (Fig. 36). Zooid 65-90 um long x 35-50 um wide. Diameter of peristomial lip usually less than 
the maximum body width. Contractile vacuole situated just below the peristome. Macronucleus variable, 
usually S-shaped. Pellicle with fine transverse striations. Stalk up to x 3 body length. Stalk base flattened to 
form a large attachment area, the diameter of which often exceeds that of the body. 

HABITAT. Freshwater, attached to the tail fins of frog tadpoles. 

Genus TVCOLESCA (Tucolesco, 1962) Lorn in Corliss, 1979 
Leptodiscus Tucolesco, 1962. 

The main distinguishing features of this genus are (i) the reported absence of a peristomial lip, and 
(ii) upon contraction, the peristome remains open with the cilia extended while the stalk folds into 
several (usually three) loose coils. T. mirabilis is the type species by monotypy. 
The name originally assigned to this genus was Leptodiscus, a homonym of the dinoflagellate 


150 


A. WARREN 


Fig. 36 Monintranstylum ranae, (a) typical form; Fig. 37 Tucolesca mirabilis, after Tucolesco, 1962 
showing variability of (b) zooid, and (c) macro- (called Leptodiscus mirabilis). Bar = 25 um. 

nucleus, after Stiller, 1953 (called Intranstylum 
ranae). Bar = 50 urn. 

Leptodiscus Hartwig, 1877 (Corliss, 1979). The genus was renamed Tucolesca by Lorn (unpub- 
lished see Corliss, 1979). The taxonomic status of the genus and validity of the name are both 
considered doubtful. 

DIAGNOSIS. Zooid with long, prominent cilia and without a peristomial disc. Upon contraction, 
peristome remains open with the cilia extended while the stalk is folded into several loose coils. 

Species description 

Tucolesca mirabilis (Tucolesco, 1962) Lom 
Leptodiscus mirabilis Tucolesco, 1962 

DESCRIPTION (Fig. 37). Zooid 27-30 urn long x 19 um wide. Peristomial lip 20 urn in diameter and obliquely 
orientated with respect to major body axis. Cilia 16-19 um long. Infundibulum reaches two-thirds of the body 
length. Contractile vacuole located near base of infundibulum. Macronucleus S-shaped and moniliform. 
Stalk up to 80 um long. 

HABITAT. Originally isolated from Lake Tekirghiol, a saline lake in Romania. 

References 

Banina, N. N. 1976. (Some consideration of the systematics of the families of the Aloricata (Peritricha 

Sessilina Kahl, 1935). Abstr. in Russian). Mat. 11 All-Union Congress of Protozoology Pt. 1: 20-22. 
1977a. (Morphological systematical descriptions of Peritricha Sessila. in Russian) Trudy Gosniorh 


119:5-11. 

19776. (New species of Peritricha on planktonic organisms of Ropsha ponds.- 


-in Russian) Trudy 


Gosniorh 119: 24-38. 
1982. (Ciliates of Peritricha, Sessilina in amphibians from Leningrad environs. in Russian). 

Parazitologiya 16: 1 44 - 1 5 1 . 
Bierhof, M. J. & Rons, P. J. 1976. Sedentary ciliates from two Dutch freshwater Gammarus species. Bijdragen 

tot de Dierkunde 46: 1 5 1-1 70. 
Bory de St. Vincent 1826. Essaid'une Classification des Animaux Microscopiques. Paris, 104 pp. 


HAPLOCA ULUS MORPHOLOGICAL RELATIVES 1 5 1 

Carey, P. G. & Warren, A. 1983. The role of surface topography in the taxonomy of peritrich ciliates. 

Protistologica 19: 73-89. 
Corliss, J. O. 1979. The Ciliated Protozoa: Characterisation, Classification and Guide to the Literature, 2nd 

edition, Pergamon Press, Oxford, 455 pp. 
Cuenot, L. 1891. Protozoaires commensaux et parasites des echinodermes. Revue Biologique du Nordde la 

France 3: 292-293. 
Curds, C. R., Roberts, D. McL & Gates, M. A. 1983. British and other freshwater ciliated protozoa. Part 2. 

Ciliophora: Oligohymenophora and Polymenophora. Synopsis of the British Fauna (NS), 22: pp.387, 

London, Linnean Society. 
Dons, C. 1918. Neue marine Ciliaten und Suctorien. Tromso Museums Aarshefter 38-39 (yr 1915-1916)- 

75-100. 
Ehrenberg, C. G. 1830(1832). Beitrage zur kenntnis der organisation der infusorien und ihrer geographischen 

verbeitung, besonders in Siberien. Abhandlungen der Akademie der Wissenschaften der DDR. Berlin Year 

1832 1-88. 
Entz, G. 1 884. Uber infusorien des golfes von Neapel. Mittheilungen aus der Zoologischen Station zu NeapelS: 

289-444. 
Faure-Fremiet, E. 1904. Sur la structure du pedoncle des Vorticellidae. Compte Rendu Hebdomadaire des 

Seances de I'Academie des Sciences. Paris 57: 506-508. 
Foissner, W. 1979. Peritriche ciliaten (Protozoa: Ciliophora) aus Alpinen kleingewassern. Zoologische 

Jahrbucher (Systematik) 106: 529-558. 
1981 . Morphologic und taxonomie einiger heterotricher und peritricher ciliaten (Protozoa: Ciliophora) 

aus alpinen boden. Protistologica 17: 29-43. 
& Schiffmann, H. 1974. Vergleichende studien an argyrophilen strukturen von vierzehn peritrichen 

ciliaten. Protistologica 10: 489-508. 
Gajewskaja, N. 1933. Zur oekologie, morphologic und systematik der infusorien des Baikalsees. Zoologica. 

Stuttgart 32: 1-298. 
Guhl, W. 1985. Beitrag zur kenntnis der ciliatenfauna verschiedener belebtschlamme mit besonderer 

berucksichtigung der fruherkennung von blah- und schwimmschlammbildung an der variabilitat peri- 
tricher ciliaten. Archivfur Protistenkunde 129: 203-238. 
Jankowski, A. V. 1982. (New genera of protozoan symbionts for the fauna of Lake Baikal, Part 3. in 

Russian). Izdalelistvo 'Nauka ' Sibirskoe otdelenie Novosibirsk p. 25-32. 
Kahl, A. 1935. Urtiere oder Protozoa. I: Wimpertiere oder ciliata (infusoria), eine bearbeitung der freileben- 

den und ectocommensalen infusorien der erde, unter ausschluss der marinen Tintinnidae. 4 Peritricha und 

Chonotricha. In: F. Dahl, ed., Die Tierwelt Deutschlands, Teil 30: 651-864. 
Kent, W. S. 1881. A Manual of the Infusoria vol II, Part V, pp. 577-720, London, David Brogue. 
Ltipkes, G. 1 975. Beitrag zur kenntnis der symphoriontenfauna auf trichopterenlarven: Haplocaulus brehmi n. 

sp. und Epistylis daxi n. sp., zwei neue peritrichen auf Agape tus fuscipes Larven. Symphoriontstudien 2. 

Protistologica 11: 295-296. 
1976. Die vertikale verteilung von ciliaten im stygorhithral der fulda (beitrag zur kenntnis mesop- 

sammaler ciliaten in fliessgewassern). InternationalJournal of Speleology 8: 127-133. 
Nenninger, U. 1948. Die peritrichen der umgebung von erlangen mit besonderer berucksichtigung ihrer 

wirtsspezifitat. Zoologischer Jahrbucher (Systematik) 77: 169-266. 
Noland, L. E. & Finley, H. E. 1931. Studies on the taxonomy of the genus Vorticella. Transactions of the 

American Microscopical Society 50: 81-123. 

Penard, E. 1922. Etudes sur les Infusoires d'Eau Douce. George & Cie, Geneve, 331 pp. 
Piesik, Z. 1975. Orzeski epizoiczne kielzach w podrodzaju Rivulogammarus Karaman strumieni okolic 

Poznania. Badania Fizjogrqficzne nad Polska Zachodnia (Zoologica) 28: 41-77. 
Precht H. 1935. Epizoen der kieler bucht. Nova Acta Leopoldina Halle NF3: 405-474. 
Shubernetskij, I. V. & Chorik, F. P. 1977. (Epibiont peritricha infusoria (Ciliata: Peritricha) of the lower 

crustaceans. in Russian). Izvestiya Akademii Nauk Moldavskoi SSR (Biologia) Year 1977: 61-66. 
Sommer, G. 1 95 1 . Die peritrichen ciliaten des grossen ploner sees. Archivfur Hydrobiologie 44: 349-440. 
Sramek-Husek, R. 1948. K Poznani planktonu a epizoonu stfedoceskych rybniku. Casopis Narodniho Musea 

117: 67-81. 
Stiller J. 1940. Beitrag zur peritrichenfauna des grossen ploner sees in Holstein. Archivfur Hydrobiologie 36: 

263-285. 
1951. Die limnologischen verhaltnisse des naturschutzgebietes von batorliget in ungarn nebst beschrei- 

bung einiger neuer peritrichen-arten (Ciliata, Protozoa). Archivfur Hydrobiologie 56: 186-260. 
1963. Zur limnologie der natrongewasser ungarns. I Der natronsee nagyszek und seine peritrichenfauna. 

Internationale Revue der Gesamten Hydrobiologie und Hydrographie 48: 603-612. 


152 


A. WARREN 


1968. Peritriche ciliaten okologisch verschiedener biotope von rovinj und umgebung. Acta Zoologica 


Academiae Scientiarum Hungaricae 14: 185-21 1. 

1971. Szajkoszorus Csillosok Peritricha. Fauna Hungariae 105: 1-245. 


Stokes, A. C. 1 889. Notice on new peritrichous infusoria from the fresh waters of the United States. Journal of 

the Royal Microscopical Society 12: 477-482. 
Szczepanowski, P. 1978. (Epizoic Ciliata on Asellus aquaticus (L) of Poznah and surroundings. in Polish). 

Prace Komisji Matematyczno- Przyrodniczej Poznanskie Towarszystwo Prvjacio Nank Poznan 47: 1-72. 
Tucolesco, J. 1962. 1. Especes nouvelles d'Infusoires de la Mer Noire et des bassins sales paramins. Archivfur 

Protistenkunde 106: 1-36. 
Warren, A. 1986. A revision of the genus Vorticella (Ciliophora: Peritrichida). Bulletin of the British Museum 

(Natural History). Zoology. London 50: 1-57. 
1987. A revision of the genus Pseudovorticella Foissner & Schiffmann, 1974 (Ciliophora: Peritrichida). 

Bulletin of the British Museum (Natural History). Zoology. London 52(1); 1-12. 

Manuscript accepted for publication 14 July 1987 


Index to species 


(Names given in roman refer to synonyms) 


Baikalonis foissneri 143 
gammari 143 
undulata 144 

Cotensita commensalis 145 

Haplocaulus amphibiarum 1 30 
amphiurae 1 3 1 
anabaenae 148 
brehmi 1 3 1 
carinogammari 131 
claudicans 1 32 
conosomus 133 
crassicaulis 133 
dipneumon 134 
distinguendis 135 
distinguendus 135 
eforianus 135 
elegans 1 36 

elegans f. gammari 1 36 
epizoicus 137 
extensus 137 
fluviatilis 137 
fur cellar iae 129 
fusiformis 138 
hengsti 135 
kahlii\39 
leanderi 1 39 


longinuclei 1 39 
macronucleatus 139 
nicoleae 148 
pelagicus 140 
procerus 141 
sertulariarum 141 
stiller i 141 
terrenus 143 
waiter i 143 

Intranstylum elegans 136 
ranae 149 

Leptodiscus mirabilis 1 50 

Monintranstylum ranae 149 
rotundus 149 
sommeri 149 
stammeri 149 

Parazoothamnium claparedei 147 
gammari 147 
stenotica 147 

Piesika gammari 147 
Pseudohaplocaulus anabaenae 148 


nicoleae 148 

Spastosty la sertulariarum 141 
Tucolesca mirabilis 1 50 

Vorticella amphiurae 131 
anabaenae 148 
carinogammari 131 
claparedei 147 
claudicans 132 
conesoma 133 
conosoma 133 
crassicaulis 133 
dipneumon 134 
eforiana 135 
epizoica 137 
extensa 137 

extensa var. macronucleata 139 
fusiforma 138 
kahlii 139 
pelagica 140 
procera 141 
sertulariarum 141 
undulata 144 

Vorticellopsis undulata 144 


British Museum (Natural History) 

The birds of Mount Nimba, Liberia 


Peter R. Colston & Kai Curry-Lindahl 


For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located within the rain-forest biome, is rapidly being destroyed by 
human exploitation. 

This book is the first major work on the birds of Mount Nimba and surrounding lowland rain- 
forests. During 20 years (1962-1982) of research at the Nimba Research Laboratory in 
Grassfield (Liberia), located at the foot of Mount Nimba, scientists. from three continents have 
studied the birds. In this way Mount Nimba has become the ornithologically most thoroughly 
explored lowland rain-forest area of Africa. 

The book offers a comprehensive synthesis of information on the avifauna of Mount Nimba 
and its ecological setting. During the 20 years period of biological investigations at Nimba this in 
1962 intact area was gradually opened up by man with far-reaching environmental consequences 
for the rain-forest habitats and profound effects on the birds. Therefore, the book provides not 
only a source of reference material on the systematics, physiology, ecology and biology of the 
birds of Mount Nimba and the African rain-forest, but also data on biogeography in the African 
context as well as conservation problems. Also behaviour and migration are discussed. At 
Nimba a number of migrants from Europe and/or Asia meet Afrotropical migratory and 
sedentary birds. 

Professor Kai Curry-Lindahl has served as Chairman of the Nimba Research Laboratory and 
Committee since its inception in 1962. Peter Colston is from the Subdepartment of Ornithology, 
British Museum (Natural History), Tring, and Malcolm Coe is from the Animal Ecology 
Research Group, Department of Zoology, Oxford. 

1986, 129pp. Hardback. 565 00982 6 17.50. 


Titles to be published in Volume 54 


The cranial muscles and ligaments of macrouroid fishes (Teleostei: Gadiformes); 
functional, ecological and phylogenetic inferences. By Gordon J. Howes 

A review of the Macrochelidae (Acari: Mesostigmata) of the British Isles. 

By Keith H. Hyatt & Rowan M. Emberson 

A revision of Haplocaulus Precht, 1935 (Ciliophora: Peritrichida) and its 
morphological relatives. By Alan Warren 

Echinoderms of the Rockall Trough and adjacent areas. 3. Additional records. 

By R. Harvey, J. D. Gage, D. S. M. Billet, A. M. Clark & G. L. J. Paterson 


Primed in Cireat Britain by Henry Ling Ltd., at the Dorset Press, Dorchester. Dorset 


(NATURA! 

- 1AI 

PRESENTED 
GEN ; 


Bulletin of the 

British Museum (Natural History) 


Echinoderms of the Rockall Trough and 

adjacent areas 

3. Additional records 


R. Harvey, J. D. Gage, D. S. M. Billett, 
Ailsa M. Clark & G. L J. Paterson 


Zoology series Vol 54 No 4 28 July 1988 


The Bulletin of the British Museum (Natural History], instituted in 1949, is issued in four 
scientific series. Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
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specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 


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The Zoology Series is edited in the Museum's Department of Zoology 

Keeper of Zoology : MrJ. F. Peake 
Editor of Bulletin : Dr C. R. Curds 
Assistant Editor : Mr C. G. Ogden 


ISBN 565 05040 

ISSN 0007-1498 Zoology series 

Vol54 No 4 pp 153- 198 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 28 July 1 988 


Echinoderms of the Rockall Trough and adjacentT^ril^ 
3. Additional records ., , 2 8 JUL )988 

R. Harvey & J. D. Gage 1 

Dunstaffnage Marine Research Laboratory, Scottish Marine Biological AWri*tirm rito. Jkv* -V.J 
Oban, Argyll PA34 4AD 

D. S. M. Billett 

Institute of Oceanographic Sciences, Brook Road, Wormley, Godalming, Surrey GU8 5UIT 


Ailsa M. Clark & G. L. J. Paterson 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD 

- 1AUGS98* 


Introduction 

This paper is the third part of a study primarily on the distribution of the echinoderm 


PRESENTED 
aunaofthe 


Rockall Trough. Part 1 dealt with the crinoids, asteroids and ophiuroids, while Part 2 covered the 
echinoids and holothurians (Gage et al. 1983, 19850). Taxonomic descriptions and discussion of 
new or problematic species in these collections are given separately in papers cited in the text. 

The present paper results from sampling undertaken from RRS Challenger by the Scottish 
Marine Biological Association (SMBA) since the publication of Parts 1 and 2. This has resulted in 
the recovery of four additional species of crinoids, ten asteroids, seven ophiuroids, one echinoid 
and eight holothurians. Of these, one asteroid, the goniasterid Mediaster bairdi, is a new record for 
the NE. Atlantic while another, a pterasterid, appears to be undescribed. Seven of the species 
mentioned in Parts 1 and 2 (one crinoid, one ophiuroid, one asteroid, one echinoid and three 
holothurians) are identified or re-identified as a result of further research. Additional records of 
species included in Parts 1 and 2 are given, together with a summary of the zoogeographic and 
bathymetric distribution with details of any range extension within the Trough provided by the 
new records. These data have mainly resulted from a greater intensity of sampling effort in the 
depth range 500-2000 m than in the period 1973 to 1982 covered by the previous papers. The 
majority of new records are from the Feni Ridge and Hebridean Slope, the latter having been 
obtained largely from fishing cruises by Dr J. D. M. Gordon of SMBA using a semi-balloon otter 
trawl (Gordon, 1986). 

In addition to records from new stations, an updated total is given for the number of specimens 
recovered from old stations where sorting of additional subsamples have provided more material. 
The depth range at the end of the list of stations for each species is the new range within the Rockall 
Trough area as indicated by our samples. 

The format of Part 3 broadly follows that of the two previous papers. Details of the gears 
employed in the sampling programme may be found in Part 1, while lists of sampling stations 
worked are distributed among all three parts. 

List of Species 

Only species additional to those included in Parts 1 and 2 are listed below. An asterisk denotes 
species included in Parts 1 and 2 under another name. 

Class Crinoidea 

Order Millericrinida 
Family Bathycrinidae 

Bathycrinus gracilis Wyville Thomson 
*Democrinus parfaiti Perrier 


Bull. Br. Mm. not. Hist. (Zool.) 54(4): 153-198 Issued 28 July 1988 

153 


1 54 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

Order Comatulida 
Family Antedonidae 

Trichometra cubensis (Pourtales) 

Poliometra prolixa (Sladen) 
Family Atelecrinidae 

Atelecrinus balanoides (P. H. Carpenter) 

Class Asteroidea 
Order Paxillosida 

Family Astropectinidae 

* Persephonaster patagiatus (Sladen) 
Order Notomyotida 

Family Benthopectinidae 

Cheiraster sepitus (Verrill) 
Order Valvatida 
Family Asterinidae 

Anseropoda placenta (Pennant) 
Family Goniasteridae 

Ceramaster granularis (Retzius) 
Mediaster bairdi (Verrill) 
Family Poraniidae 

Chondt -aster grandis (Verrill) 
Poraniomorpha hispida rosea Danielssen & Keren 
Order Spinulosida 
Family Pterasteridae 

Pteraster ( Apterodon) sp. 
Diplopteraster multipes (M. Sars) 
Hymenaster regalis Verrill 
Order Brisingida 
Family Brisingidae 

Novodinia pandina Sladen 
Order Forcipulatida 
Family Asteriidae 
Neomorphaster talismani E. Perrier 

Class Ophiuroidea 
Order Phrynophiurida 
Family Ophiomyxidae 

Ophiomyxa serpentaria Lyman 
Ophioscolex glacialis Miiller & Troschel 
Ophiophrixus spinosus (Storm) 
Order Myophiurida 
Family Ophiacanthidae 
Subfamily Ophiacanthinae 

Ophiolebes bacata Koehler 
Subfamily Ophiotominae 

Ophiotoma coriacea Lyman 
Subfamily Ophioplinthacinae 

Ophiomitrella clavigera (Ljungman) 
Family Amphiuridae 

Amphiura tritonis Hoyle 
Family Ophiuridae 
Subfamily Ophiurinae 
*Ophiura scomba Paterson 

Class Echinoidea 
Order Spatangoida 
Family Spatangidae 

*Brissopsis ?lyrifera (Forbes) 
Brisaster fragilis (Diiben & Koren) 


ROCKALL TROUGH ECHINODERMS 3 1 55 


Class Holothurioidea 
Order Dendrochirotida 
Family Paracucumidae 

Paracucumaria hyndmani (Thompson) 
Family Sclerodactylidae 

Pseudothyone raphanus (Duben & Koren) 
Family Cucumariidae 

Thyonefusus(O. F. Muller) 
Order Aspidochirotida 
Family Synallactidae 

Mesothuria intestinalis (Ascanius & Rathke) 
Mesothuria verrilli (Theel) 
Order Elasipodida 
Family Elpidiidae 

Ellipinion delagei (Herouard) 
Order Apodida 
Family Synaptidae 

Leptosynapta decaria (Ostergren) 
Family Myriotrochidae 

*Myriotrochus clarki Gage & Billett 
* Prototrochus zenkevitchi rockallensis Gage & Billett 
* Parvotrochus belyaevi Gage & Billett 
Order Molpadiida 
Family Caudinidae 

Hedingia albicans (Theel) 


Systematic Account 

A chart showing the localities of all records covered by the three papers is given in Fig. 1 . 
Classification of the Ophiuroidea follows Fell (1982). Treatment of the other four classes follows 
Parts 1 & 2 with the following exceptions: Asteriidae Blake ( 1 987), Brisingidae Downey ( 1 986), and 
Dendrochirotida Panning (1949) as amended by Pawson & Fell (1965). References to works 
describing species have only been given for those species which are listed here for the first time or 
where a new work has appeared since the publication of Parts 1 & 2, as in the case of the monograph 
of the deep North Atlantic Ophiuroidea (Paterson, 1985). 

Taxonomic responsibility is shared as follows: Crinoids, A.M.C.; Asteroids, A.M.C. & R.H.; 
Ophiuroids, G.L.J.P.; Echinoids, J.D.G.; Holothurians, D.S.M.B. & J.D.G. 

Class CRINOIDEA 
Order MILLERICRINIDA 
Family BATHYCRINIDAE 

Bat hycrinus gracilis Wyville Thomson, 1872 

See: A. M. Clark, 1977: 164-167, fig. 1; A. M. Clark, 1980: 206-207, fig. 5. 
SAMPLE. ES 27 (1). [c. 2900m] 

DISTRIBUTION. Previously known from the West European and Iberian Basins in 4430-5275 m; the 
northernmost and also the least deep record was from the Porcupine Abyssal Plain, c. 50N, 
1 555'W (A. M. Clark, 1977). There is also an unpublished intermediate record in the collections of 
the Institute of Oceanographic Sciences (IOS), Wormley, from the Porcupine Seabight (Sta. 51 109 
#2) in c. 3985 m depth. The new record, from the southern Rockall Trough, is somewhat shallower 
and farther north. 

REMARKS. The specimen consists of a crown including proximal parts of some arms to the sixth 
brachial. The total height is c. 6 mm. The appearance is similar to that of the specimen shown in 
Fig. 5 in A. M. Clark (1980), except that the knob-like fused basals and short uppermost columnals 


ow 


Bathymetry 


200 m 
1000 m 

2000 rn 

3000 IT 
AOOOrn 


..... 

\ A V X y-^Tu'^^ k^rfi 

...... , 


^/V/^- X ^ 


18W 16 U 

Fig. 1 Bathymetric chart of the area sampled s 


10 8 6 

.howing location of all stations yielding echinoderms. 


ROCKALL TROUGH ECHINODERMS 3 1 57 

are missing. The division series have sharp lateral flanges and a median keel, unlike the medially 
rounded ossicles of B. carpenteri (Danielssen & Keren), known from the deep basin of the 
Norwegian Sea in 1 360-28 1 5 m. 

In addition the collections include a very young specimen of Bathycrinus from SBC 21 1 on the 
Hebridean Slope in the anomalous depth of 402 m. This is at the same stage of development as the 
small B. gracilis figured by A. M. Clark (1977, fig. le) from the Porcupine Abyssal Plain, with 
rudimentary arms and an inverted conical calyx with all the sutures distinct and the basal ring 
'integrated' with the radial ring, so that the suture between the two rings forms a zigzag. In larger 
specimens of Bathycrinus the articulation becomes almost a straight line. Only two division series 
remain attached to the calyx and these are too poorly developed to exhibit the specific characters. It 
is just possible that the specimen may be a juvenile Bathycrinus carpenteri that has been carried over 
the Wyville Thomson Ridge. 

Democrinus parfaiti Perrier, 1 883 

See: Rom, 1977: 39^0, figs 4, 9, 10, 11, 16, pi. 2, figs 6-8, pi. 5, figs 1-6; A. M. Clark, 1977: 172-177, fig. 3a. 
Also Gage et al., 1983: 270 (as Rhizocrinus lofotensis). 

SAMPLES. SBC 66 (? frag.), SBC 67(2), SBC 2 1 6( 1 ), AT 230( 1 5), SBC 280 ( 1 ) [also ES 1 8(3), ES 20(8) in Part 1 as 
Rhizocrinus lofotensis}. [c. 1000-2200 m] 

DISTRIBUTION. The most northerly record hitherto is from SW. Ireland at c. 50N. The discovery of 
these specimens since the completion of Part 1 suggests that the species represented in the Rockall 
Trough is not Rhizocrinus lofotensis, as thought by A.M.C. in 1983, but Democrinus parfaiti. The 
depths of more than 1000 m are also indicative of this species which ranges from 870-2500 m (and 
possibly 2959 m) off Western Europe and NW. Africa to the Azores. The specimen from Sta. 10 of 
the Ingolf, off SW. Iceland, that was figured by A.M.C. (1970, fig. 4e) as Rhizocrinus lofotensis, is 
likely to be referable to Democrinus parfaiti on the basis of the Rockall Trough records, thus 
extending the range even further. Records from Knight Errant, Lightning and Porcupine are also 
likely to be D. parfaiti. However the fragment described by Doderlein (1912) from the Wyville 
Thomson Ridge in 547 m under the name of/?, rawsoni may well have been R. lofotensis, the latter 
species occupying relatively shallow depths of 140-700 m in the southern part of the Norwegian 
Basin. It has however been recorded from down to 3475 m off Greenland. An unpublished recent 
IOS record of D. parfaiti in the Porcupine Seabight (Sta. 101 1 1 #8) in c. 1630 m helps to confirm 
the Helga record of A. H. Clark (1913), that specimen now being in a badly decalcified condition. 
The distinction between D. parfaiti and Rhinzocrinus lofotensis is discussed in Clark (1970: 21). 

REMARKS. The calyx of the largest of the present specimens, from SBC 67, is only 2.5 mm high, the 
same size as the smallest one given in the table of measurements of/), parfaiti in A. M. Clark ( 1 977). 
Study of the material indicates that the supposition made in 1977 that the calyx is consistently 
narrow, even in young specimens of this species, was incorrect. Judging from the smaller specimens 
from the Rockall Trough, the initial shape is inverted conical as in the other species of the genus for 
which ontogenic series have been observed. 

Order COMATULIDA 
Family ANTEDONIDAE 

Tnchometra cubensis (Pourtales, 1 869) 
See: A.M. Clark, 1970: 46-^8; 1980: 195-197. 
SAMPLES. AT 219(1), AT248(41), AT249(1), ES 250(3). [1 150-1991 m] 

DISTRIBUTION. North Atlantic from the Gulf of Mexico to the Davis Strait and from Morocco to 
Portugal and NW. Spain; 210-2380 m (72432 m). Also recorded from SW. of Iceland in 31 1 m. The 
present records from W. of the Anton Dohrn Seamount and from the North Feni Ridge represent 
an extension of range in the deeper NE. Atlantic. It is possible that the smaller specimens recorded 
from Helga stations in the Bay of Biscay and W. of Ireland under the name T. delicata A. H. Clark, 
1911, will also prove to be conspecific with T. cubensis. 


1 58 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

REMARKS. The colour in life (AT 248) was brownish except for the very flared arm joints which were 
white. 

Poliometra prolixa (Sladen, 1881) 
See: A. M. Clark, 1970: 42-^5, figs 15, 16. 
SAMPLES. ES 87( 1 ), AT 226( 1 1 ). [c. 1 050- 1 1 1 8 m] 

DISTRIBUTION. Known only from the Arctic and Norwegian Sea, from Greenland to the seas off 
western Siberia. The present records from the Faeroe Bank Channel and Faeroe-Shetland Channel 
represent the southernmost limit of the range. The bathymetric range is 20-1960 m, but all of the 
more southern records exceed 500 m. 

Family ATELECRINIDAE 
Atelecrinus balanoides P. H. Carpenter, 1881 
See: A. H. Clark, 1913: 45 (as A. helgae; A. M. Clark, 1970: 49-51, fig. 19 (as A. balanoides). 

SAMPLES. AT 223(1), AT 230(2), AT 248(3), AT 249(1), ES 250(1). 13/83/6 OTSB(l), 3/85/20 OTSB(2). 
[980-1 005m to 1270m] 

DISTRIBUTION. Known in the western tropical Atlantic from Florida to NW. Brazil, and in the 
NE. Atlantic from the southern Rockall Trough (Helgd) c. 54N, 1230'W and south west of the 
Faeroes (Thor). The present records are intermediate in position between these last two. The 
recorded bathymetric range is 532-1256 m. 

REMARKS. The very long straight delicate and easily lost cirri of this species contrast with the 
relatively short, curly ones of Trichometra cubensis which was often collected in the same haul, both 
in the Rockall Trough and in the western tropical Atlantic where both species extend. They are also 
distinguished by their habitats, T. cubensis being epizoic on other organisms such as the gorgonian 
Acanella, while A. balanoides is self-supporting on muddy substrates with its widely spread cirri. 
The colour in life (AT 249) was a yellowish-buff. 

Class ASTEROIDEA 

Order PAXILLOSIDA 

Family LUIDIIDAE 

Luidia ciliaris (Philippi, 1837) 

SAMPLES. AT 29 1(2), AT 292(4). 13/83/3 GT(ll), 13/83/4GT(7), 1 3/83/7 OTSB(6), 1 3/83/8 OTSB(23), 3/85/14 
OTSB(IO), 3/85/38 OTSB(l), 3/85/43 OTSB(9), 3/85/44 OTSB(l). [220-270 m to 650-805 m] 

DISTRIBUTION. The new shallower records from the Hebridean Slope are more typical of the 
distribution of this common bathyal species than the single record from 650-805 m given in Part 1 . 

Family ASTROPECTINIDAE 
Astropecten irregularis (Pennant, 1 777) 
SAMPLES. RD 258(1), 13/83/7 OTSB(5). [135 m to 650-805 m] 

DISTRIBUTION. The new shallow records from Rockall Bank and the Hebridean Slope are not 
unexpected given the sublittoral and bathyal distribution of this species. 

Bathybiaster vexillifer (Thomson, 1873) 

SAMPLES. ES 34(?[juveniles] 4), ES 197(2), ES 200(3,?[juvenile] 1, AT 201(17), AT 218(1), AT 219(15), 
AT 228( 1 ), ES 232(2), AT 233( 1 4), ES 244( 1 ), AT 245(8), AT 247(juveniles 2), AT 248( 1 ), AT 267(juveniles 2), 
AT 271(10), AT 273(2), AT 288(22), 13/83/5 OTSB(l), 3/85/7 OTSB(3), 3/85/17 OTSB( 16). [992-2600 m] 

DISTRIBUTION. The record from AT 267 provides a slight increase in the lower bathymetric range 
within the Rockall Trough. 


ROCKALL TROUGH ECHINODERMS 3 1 59 

Plutonaster bifrons (Thomson, 1873) 

SAMPLES. ES 184(2, ?[juvenile] 1), ES 190(?[juveniles] 2), AT 201(11), ES 202(5), ES 218(l;?[juvenile] 1), 
AT 2 1 9( 1 8), AT 22 1 ( 1 ), AT 223( 1 5), AT 228 (?[juveniles] 3), AT 229( 1 , juvenile 1 ), AT 230( 1 03, ?[juvenile] 1 ), 
ES 232(2), AT 233(1), AT 239(5), ES 244(2), AT 245(4), AT 247(5), AT 249(4), ES 250(6), AT 251(7), ES 
252(juvenile 1), AT 254(2), ES 255Guvenile 1), AT 256(23), ES 264(juveniles 6), AT 267(7), AT 271(3), 
AT 273(1), ES 285(?[juvenile] 1), AT 286Guvenile 1), AT 287(?[juvenile] 1), AT 288(8), ES 289(1), 13/83/1 
OTSB(?[juvenile] 1), 13/83/2 OTSB(38), 13/83/5 OTSB(71), 13/83/6 OTSB(13), 13/83/7 OTSB(l), 9/84/9 
OTSB(22), 9/84/10 OTSB(l), 9/84/13 OTSB(12), 3/85/9 OTSB(l), 3/85/17 OTSB(77), 3/85/18 MBA(ll), 
3/85/19 MBA(l), 3/85/20 OTSB(IO), 3/85/25 OTSB(2), 3/85/29 OTSB(IO), 3/85/30 OTSB(55). [580-630 m to 
2965 m] 

DISTRIBUTION. The new records from the Hebridean Slope give an upward extension of bathymetric range by 
c. 400 m. 

Psilaster andromeda (Muller & Troschel, 1842) 

SAMPLES. ES 23(3), AT 223(12), AT 230(juvenile 1), AT 239(8), AT 291(20), GT 2(1), GT 7(1), GT 11(5), 
GT 14(2), GT 15(1), GT 16(4), AT 1(3), 13/83/6 OTSB(15), 13/83/7 OTSB(16), 9/84/1 OTSB(2), 9/84/13 
OTSB(4), 3/85/9 OTSB(8), 3/85/10 OTSB(2), 3/85/13 OTSB(1 1 1), 3/85/1 4 OTSB(5), 3/85/18 MBA(27), 3/85/ 
19 MBA(2), 3/85/25 OTSB(2), 3/85/28 OTSB(l). [640-780 m to 990-1075 m] 

DISTRIBUTION. Further studies on this and the related Persephonaster patagiatus (formerly 
Psilaster) (see remarks below and under that species) have shown that in the Rockall Trough on the 
Hebridean Slope Psilaster andromeda has a more restricted distribution than hitherto recorded, 
being common in the 700-1000 m zone. 

REMARKS. Following additional studies of the Astropectinidae by A.M.C., the distinction of 
Psilaster andromeda and Persephonaster patagiatus by the relative breadth of the superomarginal 
plates in dorsal view as shown by Mortensen (1927) was found to be fallacious. This character is 
variable in both species. A better distinction is the more convex contours of individual marginals 
in dorsal view in Persephonaster patagiatus compared with the flat surface but sharply-cut inter- 
marginal fascioles visible in Psilaster andromeda, when specimens are denuded with bleach. 
In section, the marginals of Psilaster andromeda tend to form continuously rounded arcs. In 
Persephonaster patagiatus however, the superomarginals in particular are more abruptly bent, 
making the sides of the arms flatter. Other differences are the more attenuated arms of P. patagiatus 
with somewhat longer and fewer marginal plates, usually 25-30 at R 60-90 mm as opposed to 35 + 
in P. andromeda at this size. The armament of the marginals is also different, the inferomarginal 
spines being more needle-like in P. patagiatus, and the armament at the apex of the jaw projecting 
horizontally below the mouth is very different. In Persephonaster patagiatus there is a pair of inset 
fascicles of blunt spines partly concealed above the apical spines, which are rounded in section, 
whereas in P. andromeda there is only a line of 3 or 4, very flat spade-like apical spines. Indeed this 
last character coupled with the absence of well-defined intermarginal fascioles in P. patagiatus 
justifies generic isolation of the species in the genus Persephonaster, which has previously been 
confused to some extent with Psilaster but can now be distinguished by these characters. The 
nomenclature is therefore restored to the combinations used by Mortensen (1927). 

Use of the above characters has resulted in the re-identification of some of the samples recorded 
in Part 1 . 

Persephonaster patagiatus (Sladen, 1889) 

SAMPLES. ES 15(2), ES 18(3), AT 68A(1), AT 107A(5), AT 186(3), AT 192(30,?[juveniles] 5), AT 221(4,? 
[juvenile] 1), AT 229(63), AT 254(1), AT 256(37), AT 287(27,?[juvenile] 1), SWT 18(8), SWT 27(4?), 13/83/1 
OTSB(274), 13/83/2 OTSB(IO), 13/83/5 OTSB(15), 9/84/9 OTSB(3), 3/85/20 OTSB(?[juvenile] 1), 3/85/29 
OTSB(13), 3/85/30 OTSB(66). [1265-1 130 m to 1809 m (?2965 m)] 

DISTRIBUTION. This species is widely distributed in the N. Atlantic (but see notes in Part 1. for 
possible complications). On the Hebridean Slope it occurs slightly deeper than Psilaster andromeda. 


1 60 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

Curiously there are no samples in which both species occurred. Following the reassignment of this 
species to the genus Persephonaster, as explained above, further clarification of the depth limits of 
the two species was sought by recourse to the collections from the Porcupine Seabight held at IOS, 
Wormley. These were found to include 21 samples of Psilaster andromeda identified by D.S.M.B. 
with positive depths ranging from 700 to 1490m, compared with 1 1 samples of Persephonaster 
patagiatus from 1360-2000 m. The deepest sample of/*, andromeda also included seven specimens 
of P. patagiatus. The SMBA record from SWT 27 (a fishing station) in 2965 m therefore seems 
doubtful, and may be a contaminant from an earlier haul. 

Family PORCELLANASTERIDAE 
Porcellanaster ceruleus Wyville Thomson, 1877 

SAMPLES. ES 4(5, juveniles 2), ES 10(1 01, juveniles 7), ES 27(99), ES 57(1, juveniles 7), ES 1 1 1(75), ES 1 18(43), 
ES 129(75), ES 152(61 juvenile 1), SBC 174(juveniles 2), ES 184(5), ES 185(330), ES 190(165), ES 197(2, 
juveniles 3), ES 204(90), ES 207(344), ES 2 1 8(3), ES 23 1 (20 1 ), ES 266(7), AT 267(37), AT 282(4), ES 283(31 7), 
AT 284(6), ES 285(21), AT 286(3). [1993 m to 3425-3500 m] 

DISTRIBUTION. No change. 

Order NOTOMYOTIDA 
Family BENTHOPECTINIDAE 

Benthopecten simplex Perrier, 1 88 1 

SAMPLES. ES 1 05(?[juvenile] 1 ), ES 1 84( 1 7, ?[juvenile] 1 ), ES 1 97(37), ES 200( 1 1 , ?[juveniles] 8), AT 20 1 (6 1 ), ES 
202(3,?[juveniles] 8), AT 218(44), AT 219(103), AT 228(132), ES 232(12), AT 233(95), ES 244(12), AT 
245(70), AT 247(4), ES 255(juvenile 1), AT 256(41), ES 257(1), ES 264(9), AT 271(82), AT 273(60), AT 
288(102), ES 289(32), 13/83/5 OTSB(75), 13/83/6 OTSB(4), 9/84/9 OTSB(3), 3/85/7 OTSB(2), 3/85/17 
OTSB(239), 3/85/29 OTSB(13). [1595 m to 3425-3500 m] 

DISTRIBUTION. The new records raise the upper bathymetric limit in the Rockall Trough from 
1785-1 845m to 1595m. 

Pectinasterfilholi Perrier, 1885 

SAMPLES. ES 34(8), ES 197(juveniles 20), AT 201(1), AT 219(4), AT 233(3), AT 267(3), AT 288(4), 3/85/7 
OTSB(34). [1752-2909 m] 

DISTRIBUTION. No change. 

Cheiraster sepitus (Verrill, 1 885) 

See: Sladen, 1889: 52-55, pi. 8, figs 5 & 6, pi. 12, figs 5 & 6 (as Pontaster venustus); A. M. Clark, 198 1 : 1 1 7-1 1 8, 
figs 4i-r & 5c. 

SAMPLES. AT 229(7), AT287(9), 13/83/1 OTSB(45), 3/85/29 OTSB(4), 3/85/30 OTSB(23), 3/85/45 OTSB(2). 
[1383m to 1690- 1740m] 

DISTRIBUTION. Nova Scotia south to the Caribbean, Azores and Bay of Biscay south to Cape 
Verde; 485-3703 m, but mainly 1 000-2000 m. The present records from the Hebridean Slope 
extend the known distribution in the NE. Atlantic to c. 56 30'N. confirming the prediction of 
Mortensen (1927) that Pontaster venustus Sladen, a synonym of Cheiraster sepitus according to 
A. M. Clark (1981), would probably be found in British waters. 

REMARKS. The 45 specimens measured from Sta. 1 3/83/1 OTSB range from R 45 mm to R 22 mm, 
R/r 4.8/1 to 3.0/1 (mean 3.7/1). Arm length varied within individuals and several specimens had 
regenerating arm tips. There was little tendency for the arm tips to curl dorsally in preserved 
specimens unlike those of Pontaster tenuispinus. The specimens are of a robust appearance due to 
the encroachment of the superomarginal plates on to the abactinal surface of the arms, and the 
tumidity of the inferomarginals. Sladen (1889) states that no pedicellariae of any kind are to be 


ROCKALL TROUGH ECHINODERMS 3 161 

found in Cheiraster, whereas A. M. Clark (1981) found some to be present but only rarely. At least 
8 of the Rockall specimens have small pectinate pedicellariae on the actinal interradii formed by 
the opposition of short spines on the plates. Their occurrence varies between interradii but where 
present they are distinctive. 

Pontaster tenuispinus (Diiben & Koren, 1 846) 

SAMPLES. AT 226( 1 ), AT 239(2), AT 27 1 (2), AT 273(2), AT 29 1 (22), 1 3/83/2 OTSB( 1 ), 1 3/83/7 OTSB( 1 59), 
1 3/83/8 OTSB(6), 9/84/ 1 OTSB(9), 9/84/2 OTSB( 1 ), 9/84/ 1 OTSB( 1 ), 3/85/ 1 OTSB( 1 74), 3/85/ 1 1 OTSB(2), 
3/85/14 OTSB(l), 3/85/43 OTSB(l), 3/85/44 OTSB(2). [500-560 m to 2255 m] 

DISTRIBUTION. The record from AT 271 extends the lower bathymetric limit to 2255 m. 

Order VALVATIDA 
Family ODONTASTERIDAE 

Hoplaster spinosus Perrier, 1 882 
SAMPLES. ES 266(1), AT 267(1). [2300-29 10m] 

DISTRIBUTION. The new records from the Feni Ridge are intermediate in depth between those of the 
two specimens recorded in Part 1. R was 12 mm in the specimen from ES 266. 

Family RADIASTERIDAE 
Radiaster tizardi (Sladen, 1 882) 
SAMPLES. AT 223(22), 13/83/2 OTSB(2), 3/85/20 OTSB(l). [1075 m to 1 130-1265 m] 

DISTRIBUTION. The new records confirm the occurrence of this species in the Rockall Trough, the 
record in Part 1 being from a single juvenile. They are all within the previously known bathymetric 
range. 

REMARKS. The specimens from AT 223 and 3/85/20 OTSB have R 60-1 10 mm. 

Family ASTERINIDAE 
Anseropoda placenta (Pennant, 1777) 
See: Mortensen, 1927: 99-101, fig. 57. 
SAMPLES. 1 3/83/3 GT( 14), 1 3/83/4 GT(4), 1 3/83/8 OTSB(4). [220-270 m to 500-560 m] 

DISTRIBUTION. Known from the Shetlands to the Mediterranean around both the west and east 
coasts of Britain in 10-200 m, but previously down to 600 m only in the eastern Mediterranean. 
The present records from the Hebridean Shelf and Slope provide an extension of bathymetric 
range in northern waters. 

REMARKS. This species is thought to prefer sandy ground, which may explain its failure to penetrate 
far beyond the shelf edge. 

Family GONIASTERIDAE 
Ceramast er granularis (Retzius, 1783) 
See: Mortensen, 1927: 81-82, fig. 44. 
SAMPLES. AT 229(1), AT 248(1), AT 259(1), AT 273(1), AT 287(1). [1 150-2185 m] 

DISTRIBUTION. Widely distributed on both sides of the North Atlantic. Previously recorded from 
British waters from the Faeroe Channel, Lousy Bank, and on the Irish slope; 20-1400 m. The 
present records from the Feni Ridge and Hebridean Slope extend the lower bathymetric limit by 
nearly 800 m. Some of the specimens appear to intergrade morphologically with the more southern 
species C. grenadensis (recorded as C. balteatus by Mortensen, 1927). 


1 62 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

Pseudarchaster parelii (Diiben & Koren, 1846) 

SAMPLES. AT 20 1 ( 1 ), ES 202(juvenile 1 ), AT 223(1 ), AT 229( 1 ), AT 230(2), AT 248(3), AT 25 1 (juvenile 1 ;?3), 
AT 259(1), AT 267(74), AT 287(74), AT 288(4), AT 291(1), 13/83/8 OTSB(3), 3/85/10 OTSB(?1), 3/85/14 
OTSB(7), 3/85/20 OTSB(2), 3/85/29 OTSB(l), 3/85/30 OTSB(4), 3/85/43 OTSB(2), 3/85/44 OTSB(2). 
[225-2965 m] 

DISTRIBUTION. No change. 

REMARKS. The distinction between P. parelii and P. gracilis is not clear in the Rockall samples. 
Halpern (1972) has separated the species on the basis of the armament of the actinal and infero- 
marginal plates, those of P. gracilis bearing cylindrical spines 3 to 5 times longer than wide, while in 
P. parelii the spines, if any, are short and flattened. Some of the specimens identified as P. parelii 
from the Rockall area have a few longer cylindrical spines. Mortensen (1927) states that P. parelii 
grows to nearly 200mm R, and Farran (1913) recovered specimens from the west of Ireland 
reaching R 192 mm, the majority of the Irish specimens measured being > 80 mm R. None of the 
Rockall specimens with R > 60 mm appear to be P. parelii, having very thorny actinal plates and a 
conspicuous pectinate pedicellaria-like arrangement of subambulacral spines said to be character- 
istic of P. gracilis (Halpern, 1972). It is possible therefore that the 'P. parelif specimens from the 
Rockall area are merely the young of 'P. gracilis'. Until more work can be done on the distinction 
of these two nominal species, the Rockall records of P. parelii refer to the small specimens with few 
spines on the actinal plates and less obvious 'pectinate pedicellariae'. 

Pseudarchaster gracilis (Sladen, 1889) 

SAMPLES. AT 219(1), AT 223(3), AT 229(2), AT 233(1, ?[juvenile] 1 ), AT 254(71), AT 287(1 , 71), AT 288(2), ES 
289(1), 13/83/1 OTSB(6), 13/83/2 OTSB(l), 1 3/83/5 OTSB(21), 1 3/83/6 OTSB(5), 1 3/83/7 OTSB(7), 13/83/8 
OTSB(l), 9/84/9 OTSB(l), 9/84/13 OTSB(l), 3/85/7 OTSB(5), 3/85/8 OTSB(l), 3/85/17 OTSB(14), 3/85/29 
OTSB(l), 3/85/30 OTSB(2), 3/85/36 OTSB(l). [7500-560 m to 2190 m] 

DISTRIBUTION. The new records extend the known distribution northwards to just south of the 
Wyville Thomson Ridge and the North Feni Ridge c. 59 40'N (but see remarks under P. parelii 
and below). 

REMARKS. The above records relate to the larger specimens of Pseudarchaster with more thorny 
actinal plates and prominent 'pectinate pedicellariae' along the subambulacrals. The colour in life 
was a bright brick red. The largest specimen measured R 165 mm. 

Paragonaster subtilis (Perrier, 1881) 

SAMPLES. ES 204(2), AT 267(18), AT 282(1), ES 283(1), AT 284(3), ES 285Guvenile 1), AT 286(18), 51301 
OTSB(6), 3/85/5 OTSB(4), 3/85/7 OTSB(2). [1785-1845 m to 2970-2980 m] 

DISTRIBUTION. The new records slightly extend the maximum recorded depth for this species in the 
Rockall Trough. 

Plinthaster dentatm (Perrier, 1884) 

SAMPLES. ES 129(?[juvenile] 1), AT 223(1), AT 287(17), 13/83/1 OTSB(37), 13/83/5 OTSB(l), 3/85/17 
OTSB(8), 3/85/29 OTSB(l), 3/85/30 OTSB(30), 3/85/37 OTSB(l), 3/85/45 OTSB(4). [945-985 m to 2910 m] 

DISTRIBUTION. The new records raised the upper bathymetric limit on the Hebridean Slope by 
almost 400 m. 

Mediaster bairdi (Verrill, 1 882) 
See: Gray, Downey & Cerame- Vivas, 1968: 150-151, fig. 24. 

SAMPLES. AT 229(14), ES 252(5), AT 287(49), 13/83/1 OTSB(22), 3/85/30 OTSB(72), 3/85/45 OTSB(2). 
[1383-1587 m] 


ROCKALL TROUGH ECHINODERMS 3 1 63 

DISTRIBUTION. Previously known only in the W. Atlantic from Newfoundland to New Jersey, in the 
lesser Antilles and off Guyana; 642-1446 m. The Rockall records from the Hebridean Slope and 
North Feni Ridge are the first from the NE. Atlantic. Downey (pers. comm.) has found very little 
difference between N. American M. bairdi and M. capensis from S. Africa, and believes that any 
taxonomic distinction between them is infraspecific. 

REMARKS. The specimens from AT 287 ranged from R53 mm to R 21 mm, R/r = 2.4-3. 1/1. Gray et 
al. (1968) state that the specimen in their figure 24 has R/r 3.3/1, but measurements from this figure 
suggest a ratio nearer 2.5/1 . The specimens from AT 287 were a pale orange-ochre when collected 
fading to off-white in spirit. 

Family PORANIIDAE 
Porania pulvillus (O . F. Miiller, 1766) 

SAMPLES. ES 113(1, juveniles 2), AT 292(8), 13/83/3 GT(69), 13/83/4 GT(43), 13/83/8 OTSB(52), 3/85/38 
OTSB(4), 3/85/43 OTSB(3). [148 m to 565-700 m] 

DISTRIBUTION. The new records are all from the Hebridean Slope and suggest that this species is 
more common below 300 m than was thought previously, four of the above samples having come 
from >400m. 

Poraniomorpha hispida rosea Danielssen & Koren, 1 88 1 
See: Mortensen, 1927: 92-93, fig. 53; A. M. Clark, 1984: 34, fig. 1 1 B, C. 
SAMPLES. ES 112(?[juvenile] 1), AT 230(1), AT 287(1). [1210-1383(?1900)m] 

DISTRIBUTION. This subspecies was previously recorded from just south of the Wyville Thomson 
Ridge under the name Lasiaster villosus Sladen, 1889 (Porcupine Sta. 47A, 990 m). The synonymy 
of Poraniomorpha hispida and P. rosea is complicated but rosea has usually been treated as a stellate 
variety of the more nearly pentagonal P. hispida. However, in 1984 A.M.C. distinguished it 
subspecifically on account of the isolation of rosea for much of its range, both geographically and 
bathymetrically. Whereas P. hispida hispida is found all round the coast of Norway extending 
north to the southern Barents Sea, with positive depths of 100-350 m, P. hispida rosea seems to 
be essentially an upper bathyal taxon extending south from the Norwegian Basin along the 
Norwegian Trench to the Skaggerak in the east and down the Rockall Trough to the Bay of Biscay 
further west in 290m to c. 1400m. If the small specimen (R only 3-2 mm) from ES 1 12 is a young 
P. hispida rosea, as its already stellate form suggests, than the depth range extends further to 
1900m. 

Chondraster grandis (Verrill, 1878) 
See: A. M. Clark, 1984: 27, figs 4A, B, 5A, 6, 7d. 
SAMPLES. AT 229 (1), AT 247(2), 3/85/9 OTSB(l). [945-1010 m to 2084 m] 

DISTRIBUTION. This species was recorded from the NE. Atlantic for the first time by A. M. Clark 
(1984) on the basis not only of the two deeper SMBA samples but also of six others ranging from 
the Lousy Bank south to the southern Bay of Biscay (BIOGAS), including Helga and more recent 
IOS material from the Porcupine Seabight. The small holotype of Marginasterfimbriatus Sladen, 
1889 from Porcupine Sta. 31 (c. 56N ll'W) in 2487m is almost certainly conspecific with C. 
grandis, though this depth exceeds by c. 300 m the positive maximum from AT 247. The type 
locality of C. grandis is in the vicinity of Cape Cod in c. 400 m but other records from N. America 
extend down to 1640 m, while E. Atlantic records range from 840 to 2070 (?2487)m. 

REMARKS. The colour in life of the specimens from AT 247 was light red midradially on the dorsal 
surface paling laterally to cream below. The specimen collected from 3/85/9 OTSB retained 
vermilion red colour in formalin on the upper side and around the ventral margin, the rest of the 


1 64 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

lower side being off-white. Distinct areas of white papulae were present along each arm of the latter 
specimen, with a central narrow naked zone and naked interradii. The madreporite was off-white 
in colour. 

Order SPINULOSIDA 
Family PTERASTERIDAE 
Pterastermilitaris(O. F. Muller, 1776) 
SAMPLES. 1 3/83/6 OTSB( 1 ), 3/85/ 1 3 OTSB( 1 ), 3/85/28 OTSB( 1 ). [934- 1 054 m to 990-1 075 m] 

DISTRIBUTION. The new records bring the total number of specimens recorded from the Rockall 
Trough to only 6, all from the Hebridean Slope at around 1000 m. 

Pteraster pulvillus M. Sars, 1861 
SAMPLES. AT 230(1), 13/83/3 GT(1), 3/85/14 OTSB(l). [168-1210 m] 

DISTRIBUTION. The new records, all from the Hebridean Slope, supplement the two previous 
records to provide a more continuous bathymetric distribution. 

Pteraster reductus Koehler, 1907 
SAMPLE. AT 251(1). [1530-1 900m] 

DISTRIBUTION. Only two other specimens of this species have been recorded from the NE. Atlantic, 
both from the Feni Ridge. The new record is almost 400 m shallower than the record in Part 1 . 

Pteraster (Apterodon) sp. aff. P. acicula (Downey, 1970) 

See: Downey, 1973: 79, pi. 34 C, D (for P. acicula). Also Gage et al, 1983: 282 (For P. sp. aff. P. acicula). 
SAMPLE. 3/85/13 OTSB(2). [958-995 m] 

DISTRIBUTION. The new specimens were recovered from almost the identical position and depth on 
the Hebridean Slope from which a specimen subsequently confirmed as P. acicula (M. Downey, 
pers. comm.) was recorded (see Part 1). These are the only records for the NE Atlantic, the type 
locality being in the Gulf of Mexico. 

REMARKS. The specimens are plump and pentagonal, both with R 16mm, rlOmm and in good 
condition with blunt arms curling dorsally at the tips. There is no webbing at all between the oral 
spines, an absence characteristic of the subgenus Apterodon. There are six spines on each oral plate, 
the apical spine being the largest and at least twice as long and thick as the distalmost spine. The 
single stout suboral spine is longer and slightly broader than the apical oral spine, glassy through- 
out its length and distincitively tricarinate, ending in an acute point. The grooves on the sides of 
these spines which give rise to their tricarinate form commence at about one third of the length 
from the spine base. The paxillae of the dorsal surface consist of a peripheral ring of c. 10 spinelets 
each around 0-06 mm thick with an imperforate portion immediately above the base which 
becomes regularly trabeculate distally without becoming broader, as in many echinoid spines. This 
structure continues to the spine tip. Within this ring are > 1 5 spinelets with more slender rod-like 
bases only 0-02 mm thick which become spatulate and trabeculate in their distal half and 0- 14 mm 
in width. The tips of both types of spinelet protrude through the dorsal membrane, those of the 
spatulate type having trifid tips. This feature, combined with the large number of spinelets, gives the 
dorsal surface a dense prickly appearance with only small inter-paxillar spaces. As this description 
is slightly at variance with that for P. acicula (Downey, 1 973), some doubt remains as to the specific 
identity of these new specimens. 

Pteraster (Apterodon) sp. 

See: Downey, 1973: 77, pi. 33, figs A, B (for P. caribbaeus). 
SAMPLES. AT 247(2), ES 264(10). [2084-2144 m] 


ROCKALL TROUGH ECHINODERMS 3 1 65 

DISTRIBUTION. North Feni Ridge at the foot of Rosemary Bank and at the foot of Rockall Bank. 

REMARKS. This species is distinguished from P. acicula by having more attenuated arms and a 
distinctly hispid appearance as collected due to the protrusion through the dorsal membrane of the 
paxillar spinelets which end in a single point. There are c. 18-24 spinelets on each paxilla, all of 
uniform thickness for most of their length. The interpaxillar spaces are relatively large and the 
membrane is almost transparent. The unwebbed oral spines decrease evenly in size from the apical 
to the most distal sixth or occasionally seventh spine. Each oral plate bears one or sometimes two 
suboral spines which are larger than the apical oral spine and have a hyaline tip where the outer 
opaque sheath has worn away. They are trabeculate for most of their length with the exception of 
the slightly tricarinate distal portion. This species is superficially similar to P. caribbaeus Perrier, 
1881, but the paxillae have almost twice as many spinelets and there are other small differences 
between the spines on the jaw plates and the musculature of the dorsal membrane. Seven of the 
specimens are in good condition. 

The records of this species and P. sp. aff. P. acicula suggest that there may be some separation on 
the basis of habitat, given the differences in depth and geographic distribution within the Trough. 

Diploptemster multipes (M. Sars, 1865) 
See: Fisher, 191 1: 371, pi. 107, figs 1,2. 
SAMPLES. 3/85/20 OTSB(l), 3/85/34 OTSB(l), 3/85/36 OTSB(l). [980-990 m to 1225-1245 m] 

DISTRIBUTION. This species is circumarctic extending south in the NE. Atlantic along the 
Norwegian coast to the Skagerrak, in the NW. Atlantic to the latitude of Chesapeake Bay, and in 
the Pacific to California in the east and Japan in the west. These are the first records from the 
Rockall Trough and they also provide an extension of the previous known bathymetric range of 
91-1 170 m. A further sample has been taken by IOS in the Porcupine Seabight (Sta. 50602 #4) in 
1080-1 120 m. The specimen from 3/85/34 OTSB has six arms. 

REMARKS. This is the largest pterasterid occurring in British waters, R max can reach c. 1 10 mm. In 
life the colour is pale mauve dotted with white dorsally continuing to the ventral interradii. The 
wide ambulacra are emphasised by red colouration along the subambulacral plates. 

Hymenaster membranaceus Wyville Thomson, 1 887 

SAMPLES. ES 184(82), ES 185(?[juveniles] 5), ES 197(37, ?[juveniles] 4), AT 201(324), ES 202(12), ES 218(9), 
AT 219(277), ES 232(15), AT 233(324), ES 244(15), AT 245(421), AT 271(104), AT 273(? 1), AT 287(1), AT 
288(367), ES 289(25), 3/85/5 OTSBGuvenile 1). [1383-2909 m] 

DISTRIBUTION. The record from AT 287 on the Hebridean Slope raises the upper bathymetric limit 
within the Trough by some 600 m, although this is still within the known distribution of this species 
(1000-3000 m). 

Hymenaster regalis Verrill, 1 895 
See: Verrill, 1895: 203-204; H. L. Clark, 1941: 64. 
SAMPLE. AT 195(1) [in Part 1 as H. ?gennaeus], AT 288(1). [2190 m] 

DISTRIBUTION. Hymenaster regalis is a rare species known only from the holotype taken in the 
NW. Atlantic off N. Carolina at 3634'N, 7348'W; 2521 m, and from another single specimen 
taken off Cuba in 1847 m. 

REMARKS. The specimen from AT 195 measures 75mm R, 40mm r, while that from AT 288 
measures c. 75 mm R, c. 55 mm r, the measurements being approximate due to the arched dorsal 
surface and recurved arm tips. Both of the previously known specimens were of a similar size. The 
dorsal surface is firm and opaque with distinct muscle fibres between the paxillae which consist of 
only a single stout spine raising the dorsal membrane into firm peaks. The membrane is perforated 


1 66 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

by small spiraculae numbering > 50 in the space within a ring of paxillae. In the life the specimen 
from AT 288 was a pale red dorsally. 

These are the first records of this species from the eastern Atlantic, and also represent a 
considerable extension of range northwards. 

Hymenaster gennaeus H. L. Clark, 1923 
SAMPLE. AT 233(1). [2 180-29 10m] 

Family SOLASTERIDAE 
Crossaster squamatus (Doderlein, 1900) 
SAMPLE. AT 287(3). [1050-1383 m) 

DISTRIBUTION. Previously unknown south of the Faeroe Channel, this new record extends the 
range south to the Hebridean Slope at c. 56N. 

Family ECHINASTERIDAE 
Henricia Gray 

Madsen is currently revising this genus (pers. comm.). The synonymy is complex and rather than 
confuse the literature further by publishing new records at this stage, it is felt that this should await 
the revision. Two species with slightly different depth distributions appear to be represented (see 
Gage et al. , 1 983: 284), that from the deeper stations being conspecific with Henricia abyssicola sensu 
Mortensen, 1 927 (non Cribrella sanguinolenta var. abyssicola Norman, 1 869) and distinguished by 
more attenuated arms and abactinal spinelets with a prolonged glassy point. The shallower species 
has less attenuated arms and abactinal spinelets ending in three points all at the same level. This 
taxon is conspecific with Norman's variety abyssicola. 

Order BRISINGIDA 

Family BRISINGIDAE 

Brisinga endecacnemos Asbjornsen, 1 856 

SAMPLES. AT 201(5), AT 219(2), AT 233(1), AT 245(1), AT 254(1), ES 264(?[juvenile] 1), 13/83/5 OTSB(24), 
9/84/9 OTSB(19), 3/85/17 OTSB(43), 3/85/29 OTSB(IO). [1690-1740 m to 2220 m] 

DISTRIBUTION. The new records raise the upper bathymetric limit on the Hebridean Slope although 
this is still much deeper than the shallowest known record of 286 m from Trondheim Fjord. 

REPRODUCTION. Tyler et al. (1984) have described the reproductive biology of this species. Up to 
60 000 eggs may be produced by each individual and there was no clear evidence of any seasonality 
in breeding in the limited number of samples available. The oocytes reach a maximum diameter of 
c. 1250 um suggesting a direct form of demersal development. 

Brisingella coronata (G. O. Sars, 1871) 
SAMPLE. ES 289(1). [992-2450 m] 

REMARKS. This specimen is unusual in having only eight arms instead of the usual 9-13. 
DISTRIBUTION. No change. 

REPRODUCTION. This appears to follow a similar pattern to that in Brisinga endecacnemos (Tyler 
etal., 1984). 

Novo dinia pandina (Sladen, 1889) 

See: Sladen, 1889: 597-601, pi. 109, figs 1-5; Mortensen, 1927: 123-125, fig. 72 (as O dinia pandina); Downey, 
1986: 27-29, fig. 13. 

SAMPLE. 3/85/28 OTSB(l). [990-1075 m] 


ROCKALL TROUGH ECHINODERMS 3 1 67 

DISTRIBUTION. Known from the 'cold' area of the NE. Atlantic from the Faeroe Channel c. 790- 
900 m (Lightning and Porcupine), to Iceland in 225 m (Einarsson, 1948). A single specimen was 
recently recovered from the western Atlantic off N. Carolina (Downey, 1986). The occurrence of 
this rarely found species at c. 56N on the Hebridean Slope represents a southerly extension of 
range in the eastern Atlantic into the 'warm' area. Mortensen (1927) speculated however that O. 
pandina may be synonymous with O. semicoronata E. Perrier, 1885 recorded from the Denmark 
Strait and south of the Canaries in 1000-1435 m, and also O. robusta E. Perrier, 1885 known from 
the Bay of Biscay and south of the Canaries c. 880-1445 m. Following the discovery that the name 
Odinia is preoccupied, all these nominal species were referred to Novodinia by Dartnall et al., 
( 1 969). In her revision of the Atlantic brisingids, Downey ( 1 986) synonymises N. semicoronata with 
N. robusta, while retaining N. pandina as a separate entity. She suggests that Novodinia is a genus of 
only moderately deep water c. 250-1 500 m. 

REMARKS. The specimen, in common with most brisingids recovered, is incomplete, consisting of a 
disc of radius 18mm with one partly detached arm and an arm fragment. There are sixteen 
ambulacral furrows. The attached arm measures 60 mm from the disc edge and 23 mm in height at 
the point of maximum gonadal swelling. The specific characters agree with Sladen's precise 
description and the presence of papulae on the disc and arms in particular distinguishes this 
specimen from all brisingids hitherto recorded from the Rockall Trough. This specimen is a male 
so no information can be given on the possible mode of development. 

Order FORCIPULATIDA 

Family ASTERIIDAE 
Stichastrella rosea (O. F. Miiller, 1776) 
SAMPLES. RD 258(1). [135m] 

DISTRIBUTION. Further study of the bathymetric limits of Stichastrella suggests that in the Rockall 
Trough area, S. rosea is confined to the shelf in depths of less than 200 m. The record from GT 1 4 in 
713-788 m listed in Part 1 is therefore reassigned to the variety ambigua. 

Stichastrella rosea var. ambigua (Farran, 1913) 

SAMPLES. AT 259(2), AT 291(7), AT 292(21), GT 14(3) [listed as S. rosea in Part 1], 13/83/3 GT(44), 13/83/4 
GT(45), 13/83/7 OTSB(IO), 13/83/8 OTSB(165), 3/85/38 OTSB(15), 3/85/43 OTSB(21), 3/85/44 OTSB(5). 
[220-270 m to 1632m] 

DISTRIBUTION. No change. 

Neomorphaster talismani Perrier, 1 894 
See: Mortensen, 1927: 134-135, fig. 76. 
SAMPLES. AT 259(1), 3/85/30 OTSB(l). [1041m to 1420-1480 m] 

DISTRIBUTION. Known from SW. Ireland in 1350 m and south to Morocco mainly c. 400-2000 m 
but exceptionally found at 54 1 3 m. The above records from the Rockall Bank and Hebridean Slope 
extend the known geographic range of this species to the Rockall Trough. 

Family ZOROASTERIDAE 
Zoroaster fulgens Wyville Thomson, 1873 

SAMPLES. AT 198(3), AT 201(1), AT 219(1), AT 223(8), AT 229(5), AT 230(2), ES 232(3), AT 233(3), AT 
239(6; ?[juvenile] 1 ), AT 245( 1 ), AT 256( 1 3), AT 257( 1 ), AT 259( 1 4), ES 26 1 ( 1 ), ES 264(juvenile, 1 ), AT 267(3), 
AT 273(3), AT 287(190), AT 288(4), 13/83/1 OTSB(4), 13/83/2 OTSB(l), 13/83/5 OTSB(33), 13/83/6 
OTSB(21), 9/84/9 OTSB( 14), 9/84/10 OTSB(3), 9/84/1 3 OTSB(30), 3/85/7 OTSB( 1 82), 3/85/8 OTSB(3), 3/85/ 
9 OTSB(l), 3/85/17 OTSB(4), 3/85/18 MBA(IO), 3/85/20 OTSB(ll), 3/85/25 OTSB(43), 3/85/29 OTSB(49), 
3/85/30 OTSB(lOl). [940-975(7580-630) m to 4810 m] 


1 68 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

DISTRIBUTION. The records from 9/84/10 OTSB approach the minimum depth known for this 
species, but it is possible that these specimens were contaminants from the previous samples in 
1 750-1 770 m. This is supported by their small size, similar to the specimen figured as Z.fulgens var. 
ackleyiby Farran (1913 pi. 1, fig. 3). On the Hebridean Slope it is noticeable that individuals of Z. 
fulgens from depths greater than c. 1 500 m are nearly always of this smaller slender-armed form, 
whereas those from around 1000 m are larger and more robust. 

Class OPHIUROIDEA 
Order PHRYNOPHIURIDA 
Family ASTERONYCHIDAE 

Asteronyx loveni Miiller & Troschel, 1 842 

See: Paterson, 1985: 13-15, fig. 9. 

SAMPLE. 3/85/32 OTSB(l). [1055 m to 1995-2020 m] 

DISTRIBUTION. All three specimens recovered in our samples have been from the Hebridean Slope. 

Family ASTEROSCHEMATIDAE 
Asteroschema inornatum Koehler, 1906 
See: Paterson, 1985: 16, fig. 10. 
SAMPLE. ES 264(1). [1900-2144 m] 

DISTRIBUTION. This and the previous specimen are from the west side of the Trough and both were 
entwined in the branches of gorgonians. 

Family GORGONOCEPHALIDAE 
Gorgonocephalus caputmedusae (Linnaeus, 1 758) 
See: Paterson, 1985: 1 1-13, fig. 8. 

SAMPLES. AT 239(1), ES 250(?[juvenile] 1), GT 2(1), 13/83/2 OTSB(l), 13/83/6 OTSB(l), 9/84/13 OTSB(l), 
3/85/13 OTSB(l), 3/85/19 MBA(4), 3/85/23 OTSB(l), 3/85/24 OTSB(2), 3/85/25 OTSB(4), 3/85/26 OTSB(6), 
3/85/27 OTSB(4), 3/85/28 OTSB(2), 3/85/31 OTSB(3), 3/85/33 OTSB(l), 3/85/34 OTSB(2), 3/85/36 OTSB(6), 
3/85/46 OTSB(l). [940-985 m to 1 130-1265(71270)] 

DISTRIBUTION. All specimens, with the exception of the queried one from the Feni Ridge are from 
the Hebridean Slope where this species is apparently common. The new records extend the 
bathymetric range in this area. 

Family OPHIOMYXIDAE 
Ophiomyxa serpentarla Lyman, 1883 
See: Paterson, 1985: 18-20, fig. 1 1. 
SAMPLE. AT 259(1 5). [1041 m] 

DISTRIBUTION. Previously recorded in the eastern Atlantic from the Faeroe Channel and SW. 
Ireland to the Azores; 450-2440 m. Its occurrence on the Feni Ridge is therefore not unexpected. 

Ophioscolex glacialis Miiller & Troschel, 1842 
See: Paterson, 1985: 20-21, fig. 11. 

SAMPLES. AT 248(1), AT 249(1), AT 251(1), AT 259(2), 9/84/2 OTSB(l), 3/85/28 OTSB(l). [91 0-960 m to 
1530m] 

DISTRIBUTION. Previously recorded from both sides of the N. Atlantic and from Arctic seas; 
50-2727 m. These specimens, mainly from the N. Feni Ridge but also from the Hebridean Slope, 
provide a link between a specimen recovered by IOS at the extreme SW. end of the Trough and the 
Arctic populations. 


ROCKALL TROUGH ECHINODERMS 3 1 69 

Ophiophrixus spinosus (Storm, 1881) 
See: Paterson, 1985: 21-22, fig. 12. 

SAMPLES. AT 287(1), 3/85/9 OTSB(l), 3/85/10 OTSB(l), 3/85/14 OTSB(l), 3/85/28 OTSB(l), 3/85/46 
OTSB(l). [720-775 m to 1383 m] 

DISTRIBUTION. Known from the Denmark Strait and SE. Iceland to the Azores; 40-1310 m. These 
specimens are all from the Hebridean Slope and provide a slight increase in bathymetric range. 

Order MYOPHIURIDA 

Family OPHIACANTHIDAE 

Subfamily OPHIACANTHINAE 

Ophiacantha abyssicola G. O. Sars, 1871 
See: Paterson, 1985: 47-48, fig. 20. 

SAMPLES. AT 239(? 1), AT 259(2), AT 290(1), GT 1(?1), 13/83/5 OTSB(2), 13/83/13 OTSB(l), 3/85/17 
OTSB(9), 3/85/28 OTSB(5). [7650-805 m to 1955-1995 m] 

DISTRIBUTION. The new records from the Rockall Bank and Hebridean Slope provide a consider- 
able extension of bathymetric range in the Trough from the maximum of c. 1000 m given in Part 1, 
although still well within the range known for this species. 

Ophiacantha bidentata (Retzius, 1 805) 
See: Paterson, 1985: 34-36, fig. 15. 

SAMPLES. ES 129(2,?[juvenile] 1),ES 184(61),ES 185(?[juvenile] 1),ES 1 97(1 60), ES 200(56), AT 20 1(265), ES 
202(1 1), ES 207(?[juvenile] 1), AT 218(20), AT 219(308), AT 221(17), SBC 222(?[juvenile] 1), AT 228(6), AT 
229(1), ES 232(23; ?[juvenile] 1), AT 233(95), ES 244(1, ?[juveniles] 3), AT 245(1 10), AT 247(32), AT 248(1), 
AT 251(4), ES 252(1), ES 255(3), AT 256(8), ES 257(1), ES 264(27), ES 266(2), AT 267(6), AT 271(93), AT 
273(144), ES 283Guvenile 1), AT 288(1), ES 289(7), 13/83/6 OTSB(l), 3/85/7 OTSB(l), 3/85/17 OTSB(8). 
[980- 1005m to 2946m] 

DISTRIBUTION. The new records extend the upper bathymetric limit in the Trough from 1 330 m to 
c. 1000m. 

Ophiacantha crassidens Verrill, 1 885 
See: Paterson, 1985: 40-41, fig. 17. 
SAMPLES. AT 223(4), AT 230(1). [1075 1862 m] 

DISTRIBUTION. The new records are from just south of the Wyville Thomson Ridge and Hebridean 
Slope and indicate a wider distribution within the Trough than the few previous samples suggested. 

Ophiolebes bacata Koehler, 1921 
See: Paterson, 1985: 51, fig. 22. 
SAMPLE. AT 259(4). [1041m] 

DISTRIBUTION. Eastern Atlantic from the Bay of Biscay and off Madeira; 1 300-2034 m. This record 
from the North Feni Ridge is a considerable extension of range north to c. 57N and to slightly 
shallower depths. 

Subfamily OPHIOTOMINAE 
Ophiotoma coriacea Lyman, 1883 
See: Paterson, 1985: 57, fig. 23. 
SAMPLES. AT 22 1(1), AT 256(1). [160 5-1 705m] 


170 


HARVEY, GAGE, BILLETT, CLARK & PATERSON 


DISTRIBUTION. Recorded from both the western and eastern Atlantic; Cape Cod 1242m and 
the Azores to Iceland 1765-4106 m including a Helga record from SW. of Ireland. The present 
specimens from east of Rosemary Bank and the North Feni Ridge provide a small upward 
extension of bathymetric range in the eastern Atlantic and the first records from the Rockall Trough. 

Ophiolimna bairdi (Lyman, 1883) 
See: Paterson, 1985: 60, fig. 24. 
SAMPLE. ES 264(3). [2144-2910 m] 

DISTRIBUTION. This second record from the the foot of the Rockall Bank is shallower than the 
previous record by 750 m. The flow of cold water along the Feni Ridge and east side of the Rockall 
Bank may allow this circumpolar arctic species to extend into shallower depths on this side of the 
Trough. 

Subfamily OPHIOPLINTHACINAE 
Ophiomitrella clavigera (Ljungman, 1864) 
See: Paterson, 1985: 71, fig. 28. 
SAMPLE. AT 259(1). [104 1m] 

DISTRIBUTION. Known from both sides of the N. Atlantic: Davis Strait and W. Greenland; the Azores 
to the Faeroes; 1 66- 1 348 m. The single specimen was taken from the east side of Rockall Bank. 

Family OPHIACTIDAE 
Ophiactisabyssicola(M. Sars, 1861) 
See: Paterson, 1985: 76-78, fig. 32. 

SAMPLES. ES 990uvenile 1), ES 112(783; ?[juveniles 22), AT 153(2), ES 197(22), AT 201(2), ES 202(5), ES 
218(1), AT 219(2), AT 223(18), AT 226(juvenile 1), AT 228(9), AT 229(33), AT 230(39), AT 233(1), AT 
239(3), AT 247(19), AT 248(47), AT 249(1), ES 250(16), AT 251(6), ES 255(6), AT 256(2), AT 259(62), ES 
261(1), ES 264(237), AT 271(32), AT 273(10), AT 287(21), AT 290(1), 13/83/1 OTSB(54), 3/85/28 OTSB(32), 
3/85/29 OTSB(l), 3/85/30 OTSB(59). [168-3000 m] 

DISTRIBUTION. No change. 

Family AMPHIURIDAE 
Amphiura tritonis Hoyle, 1884 

See: Mortensen, 1927: 213; Paterson, 1985: 85-86, fig. 33. 
SAMPLE. SBC 222(juvenile 1). [1 101 m] 


Fig. 2 Amphiura tritonis (a) ventral (b) dorsal view of disk; (c) ventral (d) dorsal view of the ends of the 

arms. Scale bars= 1 mm. 


ROCKALL TROUGH ECHINODERMS 3 171 

DISTRIBUTION. A rare species having been recorded once from a position very near the current 
record just south of the Wyville Thomson Ridge and twice from the Bay of Biscay; 627-1290 m 

REMARKS. The undamaged specimen, which was recovered from a 0-25 m 2 box core, is considerably 
smaller than the holotype from the same locality (disc diameters 2-5 and 12 mm respectively). The 
arm tips bear numerous newly added arm segments (Fig. 2c, d) which lack spines. This small 
specimen is characterised by the ventral interradial area being scaled; by large, broadly triangular 
oral papillae and by three arm spines, of which the middle one tends to be the largest (Fig. 2a). 
Of the three N. Atlantic species with similar characteristics, A. tritonis, A. richardi Koehler and 
A. abyssorum Norman, it appears to be closest to A. tritonis. The oral shield lacks the pronounced 
distal lobe seen in A. tritonis, but this might be due to the small size of the specimen. Both A. 
richardi and A. abyssorum lack scaling on the ventral interradial area and the oral papillae of 
A. abyssorum are spine-like. These features readily distinguish them from the Rockall specimens. 

Amphiura otteri Ljungman, 1871 
See: Paterson, 1985: 86-87, fig. 33. 
SAMPLES. ES 1 12(1), SBC 205(1), ES 207(1), ES 231(3), ES 285(1). [1000-2906 m) 

DISTRIBUTION. The new records, while mainly from the SMBA Permanent Station in c. 2900m, 
include a record from 1 900 m on the South Feni Ridge. This, together with a 1 000 m record listed in 
Part 1 from the Hebridean Slope and its wide distribution in the N. Atlantic suggests that this 
species is tolerant of a variety of bottom conditions. The bathymetric range is 198-3200 m. 

Amphipholis squamata (Delle Chiaje, 1829) 
See: Paterson, 1985: 91, fig. 36. 
SAMPLES. ES 1 12(17), SBC 21 IGuvenile 1), ES 250(1). [402-1900 m] 

DISTRIBUTION. The record from ES 1 12 gives a further surprising extension of nearly 600 m to the 
known bathymetric range of this species; 0-1900 m. 

Amphilepis ingolfiana (Mortensen, 1933) 
See: Paterson, 1985: 93-94, fig. 37. 

SAMPLES. ES 27(2, ?[juveniles] 2), ES 118(2, juveniles 2), ES 129(1), ES 135(1, ?1), SBC 160(juvenile 1), ES 
184(15, ?[juvenile] 1), ES 185(21), ES 197(76), ES 200(16), AT 201(1), ES 202(3), ES 204(5), ES 207(9), 
ES 218Guveniles 5), AT 219(5), SBC 220Guveniles 2), AT 221(1), ES 231(7), ES 232(6), AT 239(1), ES 
244(juveniles 17), ES 255(1), AT 267(7), AT 271(1), SBC 272(3), AT 273(1), SBC 275(?[juvenile] 1), SBC 
276Guvenile 1), AT 282(20), ES 283(9), ES 285(1), AT 286(4), ES 289Guveniles 31). [1047-2946 m] 

DISTRIBUTION. The record from AT 239 raises the upper limit of this species in the Rockall Trough 
to near the minimum of 957 m quoted by Mortensen (1933). 


Family OPHIOCHITONIDAE 
Ophiochiton ternispinus Lyman, 1883 
See: Paterson, 1985: 96-97, fig. 39. 

SAMPLES. AT 221(5), AT 223(3), AT 228(4), AT 229(1), AT 230(1), AT 239(1), AT 251(1), ES 255(1), AT 
256(2), AT 287(1), 13/83/1 OTSB(2), 1 3/83/2 OTSB(2), 13/83/5 OTSB(l), 9/84/9 OTSB(l), 3/85/17 OTSB(l), 
3/85/29 OTSB(7), 3/85/30 OTSB(6). [1047-2200 m] 

DISTRIBUTION. The new records from the Hebridean Slope raise the known upper bathymetric limit 
within the Trough by 700 m. 


1 72 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

Family OPHIURIDAE 
Subfamily OPHIURINAE 

Ophiopleura inermis (Lyman, 1 878) 
See: Paterson, 1985: 128, fig. 48. 

SAMPLES. AT 223(6), AT 230(5), AT 239(3), AT 248(4), AT 249( 1 ), ES 250(4), AT 259(6), AT 290( 1 0), 1 3/83/2 
OTSB(3), 13/83/6 OTSB(182), 9/84/13 OTSB(3), 3/85/8 OTSB(3), 3/85/9 OTSB(28), 3/85/13 OTSB(35), 
3/85/20 OTSB(18), 3/85/23 OTSB(2), 3/85/24 OTSB(52), 3/85/25 OTSB(8), 3/85/26 OTSB(ll), 3/85/27 
OTSB(3), 3/85/28 OTSB(119), 3/85/31 OTSB(l), 3/85/32 OTSB(2), 3/85/33 OTSB(21), 3/85/36 OTSB(6), 
3/85/46 OTSB(3). [650-805 m to 1271 m] 

DISTRIBUTION. No change. 

Homophiura tesselata (Verrill, 1 894) 
See: Paterson, 1985: 135-138. 

SAMPLES. AT 181(4), AT 228(1). [1785-1 845m to 2264m] 
DISTRIBUTION. No change. 

Amphiophiura saurura (Verrill, 1894) 
See: Paterson, 1985: 134, fig. 50. 
SAMPLE. 3/85/20 OTSB(4). [1225-1245 m to 1900 m] 

DISTRIBUTION. This record extends the known distribution within the Trough to the Hebridean 
Slope and into shallower water. 

Ophioctengracilis(G. O. Sars, 1871) 
See: Paterson, 1985: 130, fig. 50. 

SAMPLES. ES 6(juveniles 1583), ES 14(juveniles 88), ES 15(juveniles 2908), ES 27(juveniles 5), ES 57(juveniles 
76), ES 105(juveniles 3362), ES 129(juveniles 5, ?[juvenile] 1), SBC 150(juveniles 39), SBC 160Guveniles 3), 
SBC 163(juveniles 5), SBC 174(juvenile 1), ES 178(juveniles 789), ES 184(juveniles 654), SBC 188(]uvenile 1), 
ES 197(juveniles 2700), ES 200(juveniles 9), ES 202(juvenile 1), ES 204(juvenile 1), ES 2070uveniles 38), SBC 
2 1 5(juveniles 2), SBC 2 1 6(?[juvenile] 1 ), ES 2 1 8(juveniles 1 768), SBC 220(juvenile 1 ), AT 226(3), AT 239( 1 62), 
ES 244Guveniles 285), ES 264(9), SBC 276(juveniles 4), SBC 278(juveniles 3), ES 283(juveniles 10), AT 
290(11), 13/83/6 OTSB(118), 13/83/7 OTSB(95), 9/84/2 OTSB(27), 9/84/13 OTSB(54), 3/85/10 OTSB(68), 
3/85/24 OTSB(IO), 3/85/26 OTSB(3), 3/85/28 OTSB(25), 3/85/36 OTSB(l). [704-2946 m] 

DISTRIBUTION. No change. 

Ophiocten hastatum Lyman, 1878 
See: Paterson, 1985: 129, fig. 49. 

SAMPLES. SBC 64(1), ES 111(2), ES 129(2), ES 185(8), ES 204(1), ES 207(1, ?1), ES 266(1), AT 267(1), ES 
283(2), 3/85/5 OTSB(9). [2000 m to 2970-2980 m] 

DISTRIBUTION. The new records slightly extend the bathymetric range within the Trough of this 
esentially abyssal species. 

Ophiura carnea Liitken, 1 858 
See: Paterson, 1985: 117, fig. 42. 
SAMPLE. 3/85/9 OTSB(l). [630-2857 m] 

DISTRIBUTION. The new record extends the known distribution within the Trough to the Hebridean 
Slope. 


ROCKALL TROUGH ECHINODERMS 3 1 73 

Ophiura scomba Paterson, 1985 
See: Paterson, 1985: 125-127, figs 46 & 56; Gage et al., 1983: 297-298 (as O. irrorata). 

SAMPLES. AT 201(1), ES 218(1), AT 219(1), AT 245(1), AT 247(28), ES 252(5), AT 254(3), ES 255(1 1), AT 
256(24), ES 257(7; ?[juvenile] 1), ES 261(17), ES 264(13), 3/85/17 OTSB(l). [1510-2220 m] 

DISTRIBUTION. O. scomba is known from Rockall south to Morocco; 1 595-4406 m. The new 
records are from the North Feni Ridge and also in the vicinity of a 2200 m repeat station (Sta. 'M') 
in the northern Rockall Trough and provide a new slightly shallower upper bathymetric limit. 

REMARKS. This species was recorded as O. irrorata in Part 1 . Subsequent studies have shown that 
the Rockall specimens should be referred to O. scomba. The swollen oral shields, rounded ventral 
arm plates and pointed oral papillae distinguish this species from its congener O. ljungmani, which 
has an overlapping bathymetric distribution in the Trough. 

Ophiura ljungmani (Lyman, 1878) 
See: Paterson, 1985: 1 18-120, fig. 44. 

SAMPLES. ES 27(172), ES 57(927), SBC 58(2), ES 105(juveniles 13), ES 111(623), ES 118(401), ES 129(951), 
SBC 159(juveniles 3), SBC 160(juvenile 1), SBC 163(juveniles 2), ES 180(305), ES 184(359), ES 185(634), ES 
197(807), ES 200(juveniles 70), AT 201(8), ES 202(69), ES 204(106), SBC 205Guvenile 1), ES 207(232), ES 
218(1 94), AT 2 1 9(8), SBC 220(2), AT 228(4 1 ), ES 23 1 (286), ES 232(26), AT 233( 1 6), ES 244( 1 70), AT 245( 1 7), 
AT 247(3), ES 264(67), ES 266(2 1 ), AT 267(22), AT 27 1 (6), AT 273( 1 4), SBC 278(juvenile 1 ), ES 283(4 1 4), ES 
285(39), AT 288(4), ES 289Guveniles 111), 13/83/5 OTSB(2). [1050 m to 3425-3500 m] 

DISTRIBUTION. No change. 

Ophiura ophiura (Linnaeus, 1 758) 

See: Sussbach & Breckner, 191 1: 234-241 (as O. ciliaris); Mortensen, 1927: 236-238, fig. 128 (as O. texturata). 
SAMPLES. SBC 210(71), AT 292(2), 13/83/3 GT(8), 3/85/43 OTSB(2). [220-270 m to 704 m] 

DISTRIBUTION. These new records confirm the presence of this sublittoral to bathyal species at 
depths > 500 m on the Hebridean Slope as reported from a single record in Part 1 . 

Subfamily OPHIOLEPIDINAE 
Ophiomusium lymani Wyville Thomson, 1 873 
See: Paterson, 1985: 147-148, fig. 58. 

SAMPLES. ES 105(juveniles2), ES 184(1 196), ES 185(juvenile 1), AT 186(1015), ES 197(451), ES 200(240), AT 
20 1 (948), ES 202(3 1 ), ES 2 1 8(juveniles 1 2), AT 2 1 9( 1 458), SBC 220(juvenile 1 ), AT 22 1 (44), AT 228( 1 224), ES 
232( 1 2 1 ), AT 233( 1 039), AT 239(3), ES 244(48), AT 245(392), AT 247(39 1 ), ES 250(2), AT 25 1 ( 1 7), ES 252(6), 
AT 254(12), AT 256(86), ES 257(12), ES 261(3), ES 264(10), AT 271(860), AT 273(572), ES 285(20), AT 
287(3), AT 288(1787), ES 289(171), 13/83/5 OTSB(417), 3/85/7 OTSB(44), 3/85/17 OTSB(1800), 3/85/29 
OTSB(1614), 3/85/30 OTSB(29). [810-2921 m] 

DISTRIBUTION. No change. 

Class ECHINOIDEA 
Order CIDAROIDA 
Family CIDARIDAE 

Cidaris cidaris (Linnaeus, 1 758) 

SAMPLES. AT 223(1), AT 292(3), 13/83/8 OTSB(l), 9/84/10 OTSB(l), 3/85/28 OTSB(4), 3/85/31 OTSB(l), 
3/85/43 OTSB(155), 3/85/44 OTSB(52). [500-560 m to 1075 m] 

DISTRIBUTION. No change. 


1 74 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

Poriocidaris purpurat a (Wyville Thomson, 1872) 
SAMPLES. AT 223(3 1 ), AT 230(4), AT 239( 1 ), AT 248(3), AT 259( 1 ), 1 3/83/2 OTSB(3). [ 1 04 1 - 1 296 m] 

DISTRIBUTION. The record from AT 223 is near the type locality of this species. The new records 
raise the known upper bathymetric limit for the Rockall Trough by c. 1 50 m. 

Order ECHINOTHURIOIDA 
Family ECHINOTHURIIDAE 

Araeosomafenestratum (Wyville Thomson, 1869) 
SAMPLES. 1 3/83/2 OTSB(l), 1 3/83/7 OTSB(l). [631 m to 1265-1 130m] 

DISTRIBUTION. These new records from the Hebridean Slope provide an extension of the lower 
bathymetric range of this species in the NE. Atlantic. 

Calveriosoma hystrix (Wyville Thomson, 1869) 

SAMPLES. AT 223(19), AT 239(1), AT 248(2), AT 259(2), AT 291(11), AT 3(1), 13/83/2 OTSB(6), 13/83/6 
OTSB(180), 13/83/7 OTSB(5), 9/84/1 OTSB(2), 9/84/2 OTSB(3), 9/84/10 OTSB(l), 9/84/1 3 OTSB(45), 3/85/8 
OTSB(l), 3/85/9 OTSB(3), 3/85/10 OTSB(8), 3/85/13 OTSB(183), 3/85/14 OTSB(126), 3/85/18 MBA(109), 
3/85/19 MBA(16), 3/85/25 OTSB(103), 3/85/28 OTSB(l). [580-630 m to 1265-1 130m] 

DISTRIBUTION. The new records extend both the upper and lower bathymetric limits of this species 
within the Trough. The numbers taken in fish trawls suggest that it is abundant on the Hebridean 
Slope at around 1000 m. 

REMARKS. The specimen from AT 239 was in excellent condition when recovered, with most of its 
long red spines intact and the body wall still supported by coelomic fluid. Trawled specimens are 
generally almost completely devoid of spines and give little impression of the magnificence of this 
species in life. 

The size-frequencies (Table 1) from semi-balloon trawls (OTSB) suggest that the largest 
individuals occur at the upper end of the depth range on the Hebridean Slope, whereas recruitment 
occurs towards the lower end of the depth range. The occurrence of juveniles at a greater depth 
than adults has been noted in a number of echinoderm taxa in the Rockall Trough (Gage et al., 
1983, 1985<2). The smallest individual recovered measured 55mm, considerably larger than the 
smallest echinoids taken with this trawl, suggesting that recruitment may be sporadic in this 
non-seasonally reproducing species (Tyler & Gage, 19840). 

Hygrosomapetersii(A. Agassiz, 1880) 

SAMPLES. AT 230(2), AT 233(2), AT 239( 1 ), AT 282( 1 ), ES 283(2), AT 286(3), AT 287(2), SWT 27(2), 1 3/83/ 1 
OTSB(l), 13/83/5 OTSB(l), 3/85/5 OTSB(IO), 3/85/21 OTSB(l), 3/85/30 OTSB(9), 3/85/45 OTSB(3). 
[11 60m to 2970-2980 m] 

DISTRIBUTION. The new records provide a small extension of the known lower bathymetric limit, 
the total range now being 730-2980 m. 

Sperosoma grimaldii Koehler, 1897 

SAMPLES. AT 239(9), AT 248(2), AT 249(72), 13/83/2 OTSB(167), 13/83/5 OTSB(l), 13/83/6 OTSB(40), 
9/84/13 OTSB(6), 3/85/8 OTSB(2), 3/85/9 OTSB(l), 3/85/1 1 OTSB(4), 3/85/13 OTSB(3), 3/85/18 MBA(88), 
3/85/19 MBA(37), 3/85/20 OTSB(62), 3/85/23 OTSB(l), 3/85/24 OTSB(19), 3/85/25 OTSB(58), 3/85/28 
OTSB(7), 3/85/33 OTSB(17), 3/85/43 OTSB(2). [565-700 m to 2910 m] 

DISTRIBUTION. Intensive trawlings on the Hebridean Slope suggest that this species is common at 
around 1000 m where it is frequently recovered with Calveriosoma hystrix. The upper bathymetric 
limit within the Trough is raised by around 500 m. 

REMARKS. Size-frequency data (Table 1) suggest that this species attains a smaller size on the 
Hebridean Slope than Calveriosoma hystrix. It appears also that as in C. hystrix the smallest 


ROCKALL TROUGH ECHINODERMS 3 


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1 76 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

individuals are found in depths greater than 1000m, with no juveniles recovered from the 
shallower stations sampled at or about the same time. Sperosoma grimaldii appears to have a 
non-seasonal reproductive cycle (Tyler & Gage, 19840). 

Family PHORMOSOMATIDAE 
Phormosoma placenta Wyville Thomson, 1 872 

SAMPLES. AT 223(88), AT 228(55), AT 230(4), ES 23 1 ( 1 ), AT 239(34), AT 248(47), AT 249(8), ES 250(3), AT 
251(2), AT 254(40), ES 255(64), AT 256(7), AT 257(1), AT 259(1), ES 261(1), AT 287(1), AT 288(2), AT 
291(2), AT 292(1), GT 7(2), GT 8(1), 13/83/2 OTSB(232), 13/83/5 OTSB(753), 13/83/6 OTSB(76), 13/83/7 
OTSB(91), 9/84/9 OTSB(71), 9/84/13 OTSB(104), 3/85/9 OTSB(1 1), 3/85/10 OTSB(2), 3/85/13 OTSB(46), 
3/85/14 OTSB(3), 3/85/17 OTSB(262), 3/85/18 MBA(17), 3/85/19 MBA(8), 3/85/20 OTSB(69), 3/85/25 
OTSB(353), 3/85/29 OTSB( 14), 3/85/34 OTSB(l), 3/85/46 OTSB(l). [525-2898 m] 

DISTRIBUTION. The new records extend both the upper bathymetric limit within the Trough, and 
the known lower bathymetric limit of this species. The total range is now 260-2898 m. 

REMARKS. The size-frequency data (Table 1) indicate that as in Calveriosoma hystrix and 
Sperosoma grimaldi juveniles tend to occur at a greater depth than adults. 

Order ECHINOIDA 
Family ECHINIDAE 

Echinus acutus var. norvegicus Diiben & Koren, 1844 

SAMPLES. ES 115(?[juveniles] 4), ES 178(72), SBC 210(juveniles 2), AT 223(65), AT 230(juveniles 7), AT 
239(48, ?[juveniles] 3), AT 248(?[juveniles] 2), AT 291(214), 13/83/2 OTSB(juveniles 16), 13/83/4 GT(?1), 
13/83/7 OTSB(6), 13/83/8 OTSB(134), 9/84/1 OTSB(49), 3/85/10 OTSB(26), 3/85/13 OTSB(l), 3/85/14 
OTSB(IOOO), 3/85/18 MBA(l), 3/85/20 OTSB(4, ?1), 3/85/24 OTSB(2), 3/85/36 OTSB(2). [401m to 
1225-1 245m] 

DISTRIBUTION. The new records raise the upper bathymetric limit by over 300 m in the Trough and 
provide a small extension of the lower bathymetric limit. Intensive trawlings on the Hebridean 
Slope suggest that this species is most common in the 500-800 m depth interval. 

REMARKS. The maximum size in these samples was found to be 77 mm test diameter, which is in 
approximate agreement with Mortensen (1943) who thought that it does not reach a size larger 
than c. 70 mm. Morphometric data from these large samples is given by Gage et /., (1986), these 
data showing good agreement with measurements given by Mortensen (1943) from material 
collected from various locations. These authors found size frequency distributions for this sea 
urchin to be quite variable in the hauls from the Hebridean Slope, this variability apparently being 
unrelated to bathymetry or time of year. This species occurs at depths similar to those yielding large 
numbers of its congener E. elegans yet analysis of these records shows that in most trawlings either 
one or the other species is overwhelmingly dominant, and in many only one of the two species 
occurs (see under Remarks for E. elegans). 

LIFE HISTORY. Gage et al. (1986) describe the seasonal cycle in oogenesis in this species on the 
Hebridean Slope; spawning occurring in about March. These authors, in interpreting growth 
banding in the plates of the test as reflecting a seasonal pattern of growth, infer that specimens 
> 50 mm in test diameter are 6-1 1 yr old. The estimated age of specimens of around 30 mm is 6 yr 
and agrees fairly well with results of Sime & Cranmer ( 1 985) from a study of populations at depths 
of 108-149 m in the northern North Sea. 

Echinus affinis Mortensen, 1903 

SAMPLES. ES 1120uveniles 94 [not 1073 as stated in Part 2]), AT 154(116), ES 176(7, ?[juveniles] 73), AT 
1 77(548), ES 1 84(58), AT 1 92(? 1 7), ES 1 97( 1 1 0, ?[juveniles] 8), ES 200(62, ?[juvenile] 1 ), ES 2 1 8(6, juveniles 2), 
ES 232(11), ES 244(10, ?0uveniles] 5), AT 245(92), AT 247(291), ES 264(130, ?[juveniles] 4), AT 267(7), 
AT 271(154), AT 273(708), AT 288(122), ES 289(14), 13/83/5 OTSB(191), 9/84/9 OTSB(78), 3/85/7 
OTSBGuvenile 1 ), 3/85/1 7 OTSB(204), 3/85/29 OTSB(22), SWT 1 3(none [listed as 3 in Part 2]). [1 605-2605 m] 


ROCKALL TROUGH ECHINODERMS 3 1 77 

DISTRIBUTION. The new records extend the known lower bathymetric limit of this species slightly. 

LIFE HISTORY. In the northern Rockall Trough at Sta 'M' in 2200 m where this species is abundant, 
E. affinis has a seasonal reproductive cycle and probably produces a planktotrophic larva. This 
seasonal reproduction is thought by Tyler & Gage ( 1 9846) possibly to be tuned to a pulsed annual 
fallout of phyto-detrital food to the sea floor in this area. Additional studies at this station have 
revealed skeletal banding which is thought to result from a seasonal growth pattern controlled by 
the same factor (Gage & Tyler, 1985). Adults were inferred to be up to c. 28 years old. These latter 
authors also indicate that postlarval survivorship is probably very low; a markedly uneven 
representation of ages in a large sample aged by means of growth rings was interpreted by them as 
probably reflecting multi-year cycles in recruitment success. 

Echinus alexandri Danielssen & Koren, 1883 

SAMPLES. ES 112(12, ?[juveniles] 108), AT 248(7), AT 251Guvenile 1), AT 256(3), AT 259(4), AT 288(1), 
13/83/1 OTSB(3), 3/85/30 OTSB(?[juvenile] 1). [1041-2300 m] 

DISTRIBUTION. The new records raise the upper bathymetric limit in the Trough by c. 230 m. 

Echinus elegans Diiben & Koren, 1 844 

SAMPLES. AT 287(718), AT 291(63), 13/83/3 GT(3), 13/83/6 OTSB(7), 13/83/7 OTSB(l), 13/83/8 OTSB(l), 
9/84/2 OTSB(15), 9/84/13 OTSB(3), 3/85/9 OTSB(241), 3/85/13 OTSB(77), 3/85/14 OTSB(5), 3/85/18 
MBA(9), 3/85/20 OTSB(?10), 3/85/23 OTSB(l), 3/85/24 OTSB(21), 3/85/25 OTSB(l), 3/85/28 OTSB(205), 
3/85/31 OTSB(2), 3/85/36 OTSB(l). [220-270 m to 1 210(71 383) m] 

DISTRIBUTION. The new records extend the known distribution within the Trough onto the shelf to 
considerably shallower depths than reported in Part 2, although still within the overall recorded 
range for this species. 

REMARKS. The maximum size of the specimens was 93 mm in test diameter, which is close to the 
maximum of 80 mm given by Mortensen (1927). Morphometric data from these large samples is 
given by Gage et al. (1986), these measurements showing good agreement with more limited data 
given by Mortensen (1943) from material collected from various other locations. The size 
frequencies of the Rockall samples are variable from haul to haul, with from one to three modes 
present; these appeared to show no relationship to either time of year or bathymetry. This species 
occurs at depths similar to those of trawlings yielding large numbers of its congener E. acutus var. 
norvegicus; yet, as pointed out above for the latter species, analysis of these hauls shows that in all 
of them either one or the other species is overwhelmingly dominant, while in many only one species 
occurs. Gage et al. (1986) suggest that this may result from differing habitat requirements. The 
muddy gut contents of E. acutus var. norvegicus indicate deposit feeding, while the varied particu- 
late remains found in E. elegans, that include both sediment and small prey, suggest a more 
omnivorous diet (Mortensen, 1943; Gage et al., 1986). 

LIFE HISTORY. The sexes are separate and equal in number. Females show a seasonal cycle in 
oogenesis, spawnout probably occurring in March. An egg size up to 60 um diameter is indicative 
of planktotrophic development as in other species of this genus (Gage et al., 1986). These authors 
interpret growth banding visible in the skeletal plates as reflecting a seasonal growth pattern. 
Counts of these growth zones and a fitted growth curve indicate that a size of 70 mm test diameter is 
reached at an age of 12-20 yr. 

Order SPATANGOIDA 
Family HEMIASTERIDAE 

Hemiaster expergitus Loven, 1874 

SAMPLES. ES 15(juveniles 2), ES 18(juveniles 3), SBC 61 (juveniles 3), SBC 156(juveniles 3), SBC 160(juveniles 
2), SBC 168(1), ES 172Guveniles 2), ES 176(juveniles 3), AT 177(?[juvenile] 1), ES 197(5), ES 204(juvenile 1), 
ES 207(1, juvenile 1), ES 218(juveniles 52), SBC 220(juveniles 2), SBC 222(juvenile 1), ES 232(2, juveniles 3), 


1 78 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

AT 239(juveniles 15), ES 244(juveniles 19), AT 245(9), ES 250(juvenile 1), AT 251(2), ES 252(40), ES 
255(juveniles 4), ES 261(1), AT 271(1), AT 282(1), ES289(juvenilel). [1047-2910 m] 

DISTRIBUTION. These records raise the upper bathymetric limit slightly within the Trough. 

LIFE HISTORY. Counts of growth bands present in the test plates indicate a test length of 30 mm 
might be reached by c. 16 yr (Gage, 1987). 

Family SPATANGIDAE 
Spatangus raschi Loven, 1869 

SAMPLES. SBC 210(?[juveniles] 4), AT 239(juvenile 1), AT 291(36), 13/83/7 OTSB(54), 13/83/8 OTSB(2), 
9/84/1 OTSB(l), 3/85/10 OTSB(13), 3/85/13 OTSB(l), 3/85/14 OTSB(40), 3/85/18 OTSB(3), 3/85/26 
OTSB(l), 3/85/43 OTSB(37). [225 m to 990-1020 m] 

DISTRIBUTION. The new records are all from the Hebridean Slope and extend the lower bathymetric 
limit within the Trough by some 200 m. This species appears to be most common in the 500-800 m 
depth zone in this area. 

Brissopsis llyrifera (Forbes, 1 84 1 ) 

See: Mortensen, 1907: 152-160, pi. 3, figs 2, 3, 7, 1 1, 12, 18, 20-23, pi. 4, figs 2, 3, 9, 14-17, pi. 18, figs 1, 6, 12, 
18, 25-26, pi. 19, figs 3, 6, 10, 15, 18-21,29,34; 1927: 338-340, figs 200, 201; 1951: 380-390, pi. 30, figs 1-4, 
7-13, pi. 32, figs 15, 20, 22, pi. 57, fig. 15; Gage et al., 1985a: 187, fig. 3 (as Brissopsis sp.). 

SAMPLES. ES 99(1), AT 239(31 + fragments). [1047-1 160 m] 

DISTRIBUTION. NE. Atlantic from Lofoten, S. and W. Iceland along the European coasts and the 
Mediterranean to S. of the Canaries; c. 5-1400 m; in soft mud. 

REMARKS. Except for the specimen from ES 99, the present material is all from a large haul (AT 
239) of small, very fragile specimens, most of which were broken when examined. In this haul the 
cod-end of the trawl was full of a muddy deposit containing Brissopsis, specimens of Hemiaster 
expergitus (see above), and a single specimen of Brisaster fragilis (see below). The largest intact 
specimen measures 22-9 mm and the smallest 9 mm in test length. It is likely that smaller specimens 
had been lost through the 10 mm-wide meshes of the trawl. 

B. lyrifera is typically an inshore species, with only a few, somewhat doubtful records from deep 
water (Mortensen, 1927). For example, it seems very unlikely that it is this species that Thomson 
(1874) records from depths to 2090 fathoms (3873 m) from the Porcupine in 1869. However, the 
absence of any confluence in the posterior petals of the present material from deep water immedi- 
ately distinguishes it from the species Brissopsis atlantica and B. mediterranea which are known to 
occur to bathyal depths in the western Atlantic (Chesher, 1968). Furthermore, B. mediterranea, 
although known from the eastern Atlantic and Mediterranean, is thought there to be restricted to 
relatively shallow water. 

The present specimens are similar in size and appearance to two lots, each consisting of two 
partly broken specimens about 1 5 mm in length, labelled as Brissopsis lyrifera, in the collections of 
the British Museum (Natural History) (BMNH). These were dredged by the Triton (presumably on 
her voyage in 1882, see Deacon, 1977) from 942m and 1170m depth, respectively, from the 
Wyville Thomson Ridge and Faroe Bank Channel. A larger specimen at BMNH measuring 
27-5 mm length was collected by the Helga in 1909 from the Porcupine Seabight at 956-1088 m 
depth (see Farran, 1913). The fragmented specimen recorded under 'Brissopsis sp.' in Part 2 that 
was taken from the same area and depth on the upper Hebridean Slope as the specimens from AT 
239 is now identified as Brissopsis llyrifera. On the basis of the lengths and widths of the anterior 
and posterior petals it is estimated that this specimen was about 30 mm in test length. The 
pedicellaria in Fig. 3a, b of Part 2 from this specimen is a rostrate pedicellaria and not globiferous 
as labelled. Similar rostrate pedicellariae were found on inshore B. lyrifera. Examples of the 
double-pronged globiferous pedicellariae typical of B. lyrifera were not found on any of the deep- 
water specimens from BMNH. The tridentate pedicellaria in figure 3c of Part 2 that was also 
obtained from the large fragmented specimen from ES 99 agrees somewhat with Mortensen's 


ROCKALL TROUGH ECHINODERMS 3 


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1 80 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

(1907) third form of tridentate of B. lyrifera, examples of the other two kinds, although described 
by Mortensen as richly developed, not being found on the present specimens. However, a typical 
form of the largest, narrow-bladed variety of tridentate pedicellaria, with the valves rather widely 
separated along their lower length, was found on one of the B. lyrifera from the Triton. Not 
surprisingly, no pedicellariae were found on the somewhat denuded specimens from AT 239. 

Measurements of 21 of the morphometric characters of the test, established by Chesher (1968), 
were made on the Triton and Helga specimens (although the accuracy of many of these is reduced 
as a result of not removing their spines) and on the present material from AT 239. In Table 2 these 
data are compared to similar measurements on inshore specimens dredged in 1968 from c. 165 m 
depth in the Arran Deep, Firth of Clyde, in the possession of J. D. G. The relatively small number 
and limited size range of the sample of inshore specimens is insufficient to adequately define 
the natural variation in this species, in which monstrosities in test form are known to occur 
(Brattstrom, 1946, Mortensen, 1951). However, measurements on specimens in BMNH trawled 
from stations in the Minch (c. 58N, 06'W) by the Scottish Fisheries Board in 1927, nearly all lie 
within 2 S.D. of the mean of the measurements on those from the Arran Deep, only measurements 
of the span of the posterior petals on two specimens in one of the hauls falling slightly below this 
range. In contrast, values of 9 out of 21 characters measured on the small deep-water urchins fall 
outside 2 S.D. of the mean for the Arran Deep sample for some specimens while for 5 of these 
characters measurements on all specimens fall outside this range (Table 2). 

The present, and our previous, finds of this urchin in the SMBA samples concur with Thomson 
(1874) who noted that at great depths specimens of B. lyrifera decrease in size, having '. . . all the 
appearance of being very young . . .'. However, on the present material genital pores were present 
on specimens as small as 1 5 mm test length, which also possessed small gonads, indicating that 
sexual maturity is reached at a small size. 

Such differences might suggest that the deep water urchins constitute a distinct taxonomic entity. 
However, Brissopsis lyrifera is known to have a planktotrophic echinopluteus that may be 
dispersed over wide areas. Since such larvae are likely to stay in the plankton for some time, it seems 
likely that the deep-water populations off the W. of Scotland are in genetic continuity with inshore 
stocks. The small size of the deep-water individuals possibly then results from dwarfing as a result 
of resource limitation in the deep sea. The differences in test characters may possibly reflect other 
differences in bottom conditions as well as their smaller size. Until the range of variation amongst 
larger inshore samples, covering both a wider range in size and bottom environment, becomes 
better understood the possible separate identity of the deep-water urchins must remain uncertain. 


Brisaster fragilis (Diiben & Koren, 1844) 

See: Mortensen, 1907: 108-123,pl. I,figs6-7,pl. 13,pl. 14, fig. 3, 7, 11, 13, 16, 18,20,24-25,31,37,39,43,46, 
50-51; 1927: 325-326, figs 187, 2-3, fig. 188, 2, fig. 189, 1-2. 

SAMPLES. SBC 224(1), AT 239Guvenile 1). [903-1047 m] 

DISTRIBUTION. NE. Atlantic from Finmark to Bergen, to north and west of the Shetlands, the 
Faroe-Shetland Channel, and from south and west of Iceland. In the western Atlantic from the 
Davis Strait to Florida; c. 65-1300 m on soft mud. 

REMARKS. The single specimen recovered in perfect condition from a box-core sample (SBC 224) 
measures 38 mm in overall test length. It was collected from 903 m depth in the Faroe Shetland 
Channel, where this species was previously taken by the Porcupine at about the same depth. The 
juvenile specimen from AT 239 measures 10-5 mm test length and was recovered from 1047 m on 
the upper continental slope west of Barra in a haul containing a mixed sample of small Brissopsis 
llyrifera and juvenile Hemiaster expergitus. As shown in the growth series figured and described by 
Mortensen (1907, pi. 13) the posterior petals at this size are relatively undeveloped. Genital pores, 
indicating sexual maturity, were not found, although Mortensen (1907) remarks that they appear 
at a size of 9-1 1 mm. 


ROCKALL TROUGH ECHINODERMS 3 181 

The present records extend the recorded range of this species in the NE. Atlantic from the Faroe 
Shetland Channel to a latitude of 57N in the Rockall Trough. 

REPRODUCTION. The large yolky eggs of Brisaster fragilis are thought to indicate direct 
development (Mortensen, 1907, 1927). 

Order POURTALESIOIDA 
Family POURTALESIIDAE 

Pourtalesia miranda A. Agassiz, 1869 

SAMPLES. ES 59(8), ES 1 18(2), ES 120(2), ES 122(4), ES 143(3), ES 147(2), ES 152(3), ES 164(3), ES 204(9 [not 
54 as stated in Part 2], ?[juvenile] 1), ES 207(6), ES 266(2), AT 282(1), ES 283(1). [2245-2946 m] 

DISTRIBUTION. The additional records extend the lower bathymetric limit slightly within the 
Trough. 

Echinosigra phiale (Wyville Thomson, 1874) 

SAMPLES. ES 27(24), ES 59(2), ES 111(2), ES 118(7), AT 121(14), ES 122(10), ES 129(9), AT 153(2), ES 
172(23), ES 180(3), ES 184(1), ES 185(37), ES 197(5), ES 204(19), ES 207(15), ES 218(4), ES 231(6), AT 
245(2), ES 257(1), AT 267(5), AT 282(6), ES 283(14), ES 285(3), AT 286(2), ES 289(2). [1993-2946 m] 

DISTRIBUTION. The lower bathymetric limit within the Trough is extended slightly and predictably, 
given the abyssal distribution of this species. 

LIFE HISTORY. Counts of growth zones present in the plates of the test indicate a size of 50 mm 
length may be reached by an age of 5-8 yr (Gage, 1987). 

Class HOLOTHURIOIDEA 

Order DENDROCHIROTIDA 

Family PSOLIDAE 

P solus pourtalesi Theel, 1886 
SAMPLE. AT 247(233). [2084-2190 m] 

DISTRIBUTION. This sample, like the first described in Part 2 (Gage et al., 1985) was taken from the 
west side of the Trough. The specimens were recovered from the base of Rosemary Bank close to 
areas where this species was found during the 7go//"Expedition (Heding, 1942). Gage et al. (1985#) 
followed Mortensen (1927) in reporting P. pourtalesi from the West Indies. Mortensen (1927) cited 
specimens from the Blake Expedition (Theel, 18866), but these came from the eastern seaboard of 
North America, not the West Indies. P. pourtalesi has not been recorded south of 39N in the 
western Atlantic. The lower bathymetric limit reported in Part 2 as 2271 m should be increased 
slightly to 2341 m to include the records of Deichmann (1940). The total range is therefore 
1096-234 1m. 

REMARKS. Heding (1942) and Gage et al. (1985a) refer to this species as P. pourtalesii but it should 
be referred to as P. pourtalesi as in the original description (Theel, 18866). 

The present specimens were found attached to a collection of small, probably ice-rafted, cobbles. 
The specimens range from 12-33 mm in length. There is some suggestion of bimodality in the size 
distribution, with a small mode around 16 mm and a much larger mode around 25 mm (Table 3). 

Table 3 Psolus pourtalesi length frequencies from station AT 247 

Lower bound of size class (mm) 
12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 

Frequency 1 3 2 2 5 4 3 8 13 12 13 32 28 33 24 16 13 5 7 3 2 1 (n = 230) 


1 82 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

Family PARACUCUMIDAE 
Paracucumana hyndmani (Thompson, 1 840) 

See: Mortensen, 1927: 400-401, figs 237, 3; 239, 1 (as Cucumaria hyndmani); Madsen, 1942: 395-406, figs 1-6 
(as Cucumaria hyndmani). 

SAMPLE. 13/83/8 OTSB(l). [500-560 m] 

DISTRIBUTION. The coasts of Europe from the Mediterranean north to the Shetland Isles and the 
coast of Norway including Rockall Bank. Mortensen (1927) notes that it is usually found in deep 
water, c. 20-1 150 m. The present specimen was taken at the top of the Hebridean Slope. 

REMARKS. Madsen (1942) recognised two forms of the species which he considered 'would prove to 
be of two geographical races of the same species': a more northern P. (Cucumaria) hyndmani f. 
typica and a more southern P. (Cucumaria) hyndmani f. robusta. Our single specimen agrees with 
the northern form and is close to the locality of a deep-water specimen examined by Madsen ( 1 942, 
p. 405). 


Family SCLERODACTYLIDAE 
Pseudothyone raphanus (Diiben & Koren, 1844) 

See: Mortensen, 1927: 407^*08, fig. 242: 2, 245: 1 (as Thyone raphanus); Panning, 1949: 456-457, fig. 52. 
SAMPLE. SBC 222(4, ?[juveniles] 2), AT 248(1 1). [1 101-1 150 m] 

DISTRIBUTION. P. raphanus occurs around the coasts of the British Isles and Norway as far north as 
c. 65N. It is also known to occur in the Mediterranean. Bathymetric range c. 10-1050 m, the 
present records extending this slightly to 1 150 m. 

REMARKS. The two juvenile specimens from SBC 222, both 1 mm long, have only primary cross 
deposits which differ from the plate deposits normally found in adult P. raphanus. However, 
primary cross deposits are the precursors to plates, and in specimens intermediate in size, 3 to 4 mm 
long, both types of deposits occur. 


Family CUCUMARIIDAE 
Thyone fusus (O. F. Muller, 1776) 
See: Madsen, 1941: 17-26, Figs 3b, 4g-h, 7b, 12-16. 
SAMPLES. SBC 21 1 (juvenile, 1). [402 m] 

DISTRIBUTION. Northeast Atlantic from Trondheim Fjord to the Mediterranean, including the 
North Sea and adjacent Scandinavian Seas. There is some uncertainty in the bathymetric range of 
this species since T. fusus has been confused in the past with other species, such as T. gadeana 
Perrier and T. wahrbergi Madsen. From his analysis of Scandinavian holothurians, Hansen con- 
siders that T. fusus generally occurs between 10 and 200m (Hansen, pers. comm.). The present 
specimen from 402 m increases the lower bathymetric limit. However, since the specimen is a 
juvenile it is possible that as with several asteroids, ophiuroids and echinoids (Gage et al. 1983, 
19850) the lower bathymetric limit is greater for juveniles than for adults. 

REMARKS. The specimen is only 2-5 mm long. The deposits of the body wall are irregular 
polyporous plates similar to those described for T. fusus f. subvillosa (Herouard, 1890; Madsen, 
1941) with a spire made of two slender columns as in T. fusus and unlike those of T. wahrbergi and 
T. gadeana. The tubefeet are large in comparison to the size of the body and have terminal plates 
100 to 125 um in diameter. These are smaller than the plates found in adult specimens but possess 
the concentric circles of large holes typical of T. fusus. The size of the specimen precluded an 
examination of the tentacle deposits. 


ROCK ALL TROUGH ECHINODERMS 3 183 

Order DACTYLOCHIROTIDA 
Family YPSILOTHURIIDAE 

YpsilothuriatalismanitalismaniE. Perrier, 1886 

SAMPLES. SBC 168(3), SBC 222(10, juvenile 1), AT 226(1), AT 230(2), AT 239(37), AT 248(8), ES 250(38). 
[c. 1000-1271 m] 

DISTRIBUTION. These additional records extend the distribution of this species to the North Feni 
Ridge and provide a geographical link between our previous records from the Rockall Trough and 
those of the Ingolffrom Iceland. Furthermore they confirm the essentially bathyal distribution of 
this species. 

Ypsilothuria bitentaculata attenuata R. Perrier, 1902 

SAMPLES. ES 27(79, juvenile 1), ES 56(15), ES 111(23), ES 118(37), ES 129(30), ES 184(31, juvenile 15), ES 
1 85(22 1 ), ES 1 90(48), ES 200( 1 1 3), ES 204(33 [not 78 as stated in Part 2]), ES 2 1 8(92), ES 23 1 (97), ES 232(1 8), 
AT 233(2), ES 244(20), AT 245(268), ES 26 1 ( 1 ), ES 266(4), AT 267(80), AT 27 1 (4), AT 273( 1 0), AT 282(5), ES 
283(67), ES 285(5), AT 286(7), AT 288(6), ES 289(41). [1862-2951 m] 

DISTRIBUTION. No change. 

Echinocucumis hispida (Barrett, 1857) 
SAMPLES. SBC 222(juveniles, 3), AT 248(8), ES 250(13), 3/85/9 OTSB(l). [945-1270 m] 

DISTRIBUTION. These records from the Feni Ridge and Hebridean Slope broaden the known 
geographic and bathymetric range of this species within the Trough. 

Order ASPIDOCHIROTIDA 
Family SYNALLACTIDAE 

Bathyplotes natans (M. Sars, 1868) 

SAMPLES. AT 239(37), AT 248(3), AT 249(1), AT 259(1), 13/83/2 OTSB(25), 3/85/36 OTSB(2). [1000-1025 m 
to 11 30-1 265m] 

DISTRIBUTION. No change. 

REMARKS. These specimens were in better condition than those described in Part 2, and had 
spicules present in both the skin and papillae. A mid ventral groove was a distinctive feature of the 
better preserved specimens. The colour in spirit was off-white with the orange body organs 
showing faintly through the skin. 

Benthothuriafunebris R. Perrier, 1902 

SAMPLES. ES 204(2), AT 284(1), ES 285(1), AT 286(9), 3/85/5 OTSB(9). [2890-2996 m] 
DISTRIBUTION. No change. 

Paelopatides grisea R. Perrier, 1902 

SAMPLES. AT 219(1), AT 273(7), AT 288(1), 13/83/5 OTSB(35), 9/84/9 OTSB(18), 3/85/29 OTSB(3). 
[1690-1 740m to 2190m] 

DISTRIBUTION. These are the first records from the Rockall Trough, the single sample recorded in 
Part 2 having been taken in the Porcupine Seabight. They are all from the area between the lower 
Hebridean Slope and the Anton Dohrn Seamount and fall within the known bathymetric range. 

Mesothuria lactea (Theel, 1886) 

SAMPLES. AT 254(2), ES 255(7), 13/83/5 OTSB(l), 9/84/9 OTSB(13), 3/85/29 OTSB(86). [1595m to 
1775-1 835m] 


1 84 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

DISTRIBUTION. These records from the North Feni Ridge and Hebridean Slope confirm the 
bathymetric distribution reported by Herouard (1923) for the NE. Atlantic. The Rockall records 
suggest a ribbon-like distribution along the margins of the Trough. 

Mesothuria intestinalis (Ascanius & Rathke, 1 767) 
See: Mortensen, 1927: 381, figs 225, 228: 3; Heding, 1942: 7, text-fig. 6. 
SAMPLES. AT 292(1), 3/85/38 OTSB(l). [41 0-490 m to 525 m] 

DISTRIBUTION. Widely distributed in the NE. Atlantic from off NW. Africa (Herouard, 1 923) to the 
coasts of Norway, although nowhere in the deep sea does it appear to be particularly common. A 
few specimens are known from the Mediterranean (see Perrier, 1902; Koehler, 1927; Sibuet, 1974) 
and the western Atlantic (Deichmann, 1930). The species has a wide bathymetric range c. 
18-1445 m, but reports of this species occurring as deep as 2000 m , appear to be the result of 
confusion with Mesothuria verrilli. The shallowest records come only from cold waters off 
Norway. The present records are from the upper Hebridean Slope. 

Mesothuria verrilli (Theel, 1886) 

See: Mortensen, 1927: 381, 382, fig. 224: 4-5; Deichmann, 1930: 93-94, pi. 6, figs 1-8. 
SAMPLES. AT 287(2), 3/85/30 OTSB(l). [1383 m to 1420-1480 m] 

DISTRIBUTION. Widely distributed in the north Atlantic with some records from the Mediterranean. 
In the western Atlantic it is known from the Caribbean Sea, eastern Gulf of Mexico (Deichmann, 
1954; Miller & Pawson, 1984) and off the Bahamas (Pawson, 1982). In the eastern Atlantic it is 
previously known from NW. Africa, the Canary Islands (Perrier, 1902), the Azores (Herouard, 
1902, 1923; Perrier, 1902), the Bay of Biscay (Sibuet, 1977) and as far north as the Porcupine 
Seabight, southwest of Ireland (Mortensen, 1927). The present records extend the distribution 
northwards into the Rockall Trough on the Hebridean Slope. 

The bathymetric range is not known with certainty partly as a result of confusion with M. 
intestinalis. Hansen (1975) gives a wide bathymetric range of 6 18^1 65m for M. verrilli in the 
Atlantic and 280 to 1 103 m in the Mediterranean. With the aid of detailed bathymetric charts 
available today it is possible to see that all reliable records of this species deeper than 2000 m come 
from areas where the seabed is particularly steep e.g. on the continental slope off NW. Africa or in 
the area of the King's Trough (see Perrier (1902) for details of some stations). The accuracy of the 
depths from which some samples were taken must therefore be called into question. This also 
applies to a shallow sample of M. verrilli on the Magazan escarpment in 550m (Perrier, 1902). 
Mortensen (1927) gives a reduced range of c. 990-1765 m for M. verrilli sampled to the southwest 
of Ireland. Recent intensive sampling in the same area, the Porcupine Seabight, confirms this total 
range but indicates that M. verrilli is only common in a reduced range between 1250 and 1500 m 
(Billett, in preparation). The present samples from the Hebridean Slope fall within this narrower 
range, as do most of those taken off the Azores (Marenzeller, 1893; Herouard, 1902, 1923; Perrier, 
1902). 

The samples from the eastern Atlantic fall within the total range proposed for M. verrilli in 
the western Atlantic of 699-3720 m (Deichmann, 1954; Miller & Pawson, 1984; Pawson, 1982; 
Suchanek et al., 1985). The study of Suchanek et al. (1985) increased the lower bathymetric limit 
from 2141 to 3720 m. The shallowest record, from 618m, quoted by Hansen (1975) for material 
from the Atlantic should be referred to specimens collected within the Mediterranean. 

REMARKS. In the western Atlantic M. verrilli is known to cover itself with detrital seagrass and shell 
material such as the seagrass Thalassia and brachiopod shells (Suchanek et at., 1985). 

Family STICHOPODIDAE 

Stichopus tremulus (Gunnerus, 1 767) 

SAMPLES. AT 292(4), 13/83/3 GT(14), 13/83/4 GT(29), 13/83/7 OTSB(l), 13/83/8 OTSB(91), 3/85/9 OTSB 


ROCKALL TROUGH ECHINODERMS 3 185 

(juvenile 1), 3/85/14 OTSB(6), 3/85/18 OTSB(l), 3/85/38 OTSB(2), 3/85/43 OTSB(18), 3/85/44 OTSB(4). 
[168m to 990- 1020m] 

DISTRIBUTION. The new records from the Hebridean Slope extend the lower bathymetric limit 
within the Trough by some 300 m. 

Order ELASIPODIDA 

Family LAETMOGONIDAE 

Laetmogone violacea Theel, 1879 

SAMPLES. AT 239(11), AT 248(9), AT 249(5), ES 250(1), AT 259(37), GT 8(6), 13/83/2 OTSB(5), 13/83/6 
OTSB(157), 13/83/7 OTSB(9), 9/84/13 OTSB(54), 3/85/18 MBA(4), 3/85/19 MBA(7), 3/85/20 OTSB(112), 
3/85/22 MBA(7), 3/85/23 OTSB(18), 3/85/24 OTSB(35), 3/85/25 OTSB(99), 3/85/28 OTSB(3), 3/85/33 
OTSB(126). [940-975 m to 1400(3000) m] 

DISTRIBUTION. The new records from the North Feni Ridge and Hebridean Slope indicate the 
presence of large populations of this species in suitable localities. 

REPRODUCTION. A non-seasonal cycle with abbreviated development is indicated (Tyler et al., 
1985/j). 

Benthogone rosea Koehler, 1 896 
SAMPLES. AT 271(10), 1 3/83/5 OTSB(3). [1745-2255 m] 
DISTRIBUTION. No change. 

REPRODUCTION. Direct development is indicated by the large oocyte size and there is no evidence of 
seasonality in oogenesis (Tyler et al., 19856). 

Family PSYCHROPOTIDAE 
Psychropotes longicauda Theel, 1 882 
SAMPLE. SWT 15(3) (incorrectly listed in Part 2 as SWT 5(3)). [2910-4810 m] 

REMARKS. P. longicauda has the largest known egg size for a holothurian at 4-4 mm diameter 
(Hansen, 1975). This size of egg probably leads to a prolonged form of early development within 
the deep-sea plankton, and juveniles of this species 13-23 mm in length have been recovered in 
pelagic nets fished at between 17 and 1500m above the seabed at abyssal depths (Billett et al., 
1985). 

Family ELPIDIIDAE 
Peniagone azorica von Marenzeller, 1893 
SAMPLES. AT 121(79), AT 271(23). [1991-3463 m] 
DISTRIBUTION. No change. 

REPRODUCTION. Additional information to that given for this species in Part 2 is provided by Tyler 
etal. (1985a). 

Kolga hyalina Danielssen & Koren, 1879 
SAMPLES. ES 266(61). [2591-2910m] 

DISTRIBUTION. These records extend the geographic range within the Rockall Trough of this 
patchily distributed species to the north central part of the Trough and Feni Ridge. 

Ellipinion delagei (Herouard, 1 896) 

See: Herouard, 1902: 39^0, pi. 6 figs 1-3, pi. 8 figs 8-9; Hansen, 1975: 163, fig. 122. 
SAMPLE. AT 247(18). [2084 m] 


186 


HARVEY, GAGE, BILLETT, CLARK & PATERSON 


DISTRIBUTION. An extremely rare species found previously only around the Azores and Cape Verde 
Islands close to steep slopes (Herouard, 1902, 1923). The presence of specimens on the Rosemary 
Bank indicates a preference for steep topography by this species, particularly around seamounts 
and oceanic islands in the N. Atlantic. The total bathymetric range is 1 165-2478 m. 

REMARKS. Only one specimen is in good condition. It widens towards the posterior end and has 12 
pairs of tubefeet bordering the entire ventral surface. Those tubefeet at the anterior end are spaced 
more widely than those at the posterior end. The posterior few pairs of tubefeet are slightly smaller 
than the rest. The deposits include large rods (up to 550 \im long), regular c-shaped deposits (about 
1 30 urn long) and small irregularly curved rods (40-50 urn in size) which are usually c-shaped and 
are often developed into tripartite deposits (Herouard, 1902). 

Order APODIDA 

Family SYNAPTIDAE 

Labidoplax southwardorum Gage, 1985 

SAMPLES. ES 1 0( 1 ), ES 34(?2), ES 53( 1 ), SBC 66(?3), ES 1 29(8, ?3), ES 1 37(3), ES 1 72(6), ES 1 76( 1 [not 22 as 
stated in Part 2]), ES 185(21), ES 190(7), ES 197(720), ES 200(1), ES 204(1), ES 218(16), SBC 220(2), ES 
244(10), ES 283(37), ES 285(1), ES 289(6). [1000-2946 m] 

DISTRIBUTION. The new records provide a slight extension of the lower bathymetric limit. 


B 


Fig. 3 (A) Labidoplax similimedia from SBC 222: entire, uncontracted specimen showing pinnate 
tentacles and position of plates and anchors in body wall; (B) Prototrochus zenkevitchi rockallensis 
from SBC 220: entire specimen showing sac-like body distended with sediment. Scale bar = 1 mm. 


ROCKALL TROUGH ECHINODERMS 3 1 87 

Labidoplax similimedia Gage, 1985 

SAMPLES. ES 118(1), ES 129(2), ES 137(4), ES 143(71), ES 164(87), ES 169(4), ES 185(4), ES 218(72), SBC 
222(8), ES 244(1), ES 283(6), ES 285(1), ES 289(2). [1 101-2946 m] 

DISTRIBUTION. The new records provide an upward extension of the bathymetric range by c. 
1 100 m, and also an extension of the lower bathymetric limit. 

REMARKS. Several of the specimens from SBC 222 are complete (Fig. 3A), with the skin deposits 
scattered in the body wall rather than crowded together and overlapping as described by Gage 
(1985), and as found in the other material from epibenthic sled hauls. This is no doubt because the 
specimen is not severely contracted as material usually is when collected by the epibenthic sledge. 
The plates and anchors have a lateral orientation in the body wall (Fig. 3 A) similar to that of other 
synaptids. The tentacles are clearly pinnate with two pairs of rounded lateral, and a single terminal, 
digit (Fig. 3A). 

Leptosynapta decaria (Ostergren, 1905) 

See: Ostergren, 1905: 146-148, fig. IB; Clark, 1907: 93; Mortensen, 1927: 431, fig. 262, 3. 
SAMPLE. SBC 210(1 + frag.). [401 m] 
DISTRIBUTION. Hitherto known only from the Trondheim Fjord to the Kattegat, 40-70 m depth. 

REMARKS. The material consists of an anterior end with the oral ring measuring 1.41 mm in 
diameter along with a posterior end, quite possibly from the same specimen, around 0.6 mm wide. 
Both fragments were whitish in colour with no trace of pigmentation when examined from spirit. 
There are 10 tentacles, each with a short terminal digit and three pairs of short rounded digits, 
increasing in size distally. Deposits consist of plates and anchors, which are found in the skin of 
both fragments, and rods present only in the tentacles. The rods, which are slightly curved and 
possess enlarged, branched ends (Fig. 4), are numerous and present throughout the tentacles, 
increasing in size distally. Skin deposits (Fig. 4) consist of plates typical in form of Leptosynapta, 
with nine toothed holes, the outermost and central holes being the largest. In the anterior fragment 
the plates have a mean length of 124-8 urn, range 1 16-138 ( = 28) and mean width of 99-1 um, 
range 90-1 1 3 (n = 14). This agrees well with the values of 126-5 and 97-5 um, respectively, given by 
Ostergren (1905) for the deposits from the anterior of the animal. The anchors agree well with 
Ostergren's description; resembling those of Leptosynapta inhaerens but being smaller and possess- 
ing up to 5 (usually 3) teeth on each fluke. Anchors are slightly longer but narrower than the plates, 
mean length 144 um, range 127-160 um ( = 23), mean width 73-1 um, range 70-75 ( = 8). These 
values are also reasonably close to the values given by Ostergren (149 x 97 um). Besides the fully 
developed plates and anchors there are a number of developmental stages of both deposits present 
(Fig. 4). 

The deposits in the posterior end are similar to those in the anterior fragment, but slightly larger 
(plates, mean length 130 um, mean width 98 um; anchors, mean length 143, mean width 72 um 
n = 4). Assuming that the two fragments belong to the same animal, it is pertinent to note that in 
Leptosynapta, it is usual for the deposits to increase in size posteriorly (Ostergren, 1905; 
Cherbonnier, 1953, 1963). Both the whitish colour and diameter of the head-end fragment 
( 1 -4 mm) is similar to that ( 1 -5-3 mm) described by Ostergren for L. decaria, and the three pairs of 
tentacle digits present is within the range of 2-4 pairs on each tentacle described by Ostergren 
(1905). 

Only two European species of Leptosynapta are known to possess 10 rather than the 12 tentacles 
typical of other species of the genus; L. minuta (Becher), a tiny viviparous species known from 
shallow water off NW Europe, and L. decaria. Both species are small and appear to lack pigmen- 
tation. The present specimen differs from L. minuta in possessing tentacle digits, in having anchors 
clearly longer than the plates (Fig. 4), and in having at the basal (articular) end of the plate small 
irregularly arranged holes, typical of most Leptosynapta species, rather than the regular slit-like 
holes of L. minuta. Ostergren (1905) found specimens of L. inhaerens with 10, 1 1 or 13 tentacles, 
this possibly influencing Clark (1907) to suggest that '. . . it is not impossible that this [L. decaria} is 


188 


HARVEY, GAGE, BILLETT, CLARK & PATERSON 


Fig. 4 Skin deposits from anterior fragment of Leptosynapta decaria from SBC 210. Fully developed 
(middle and lower right) and developing plates (upper left); fully developed (lower left) and developing 
(centre) anchor; rods from tentacles, lower left. Scale bar = 50 um. 


only the young of inhaerens. There is a striking similarity in the calcareous particles and in the 
tentacles, the differences in number of digits being simply a matter of age.' Dr Bent Hansen of the 
Zoological Museum, University of Copenhagen has confirmed (unpublished communication to 
J.D.G) that there appears to be no published record of L. decaria subsequent to Ostergren's (1905) 
paper. Dr Hansen very kindly re-examined Norwegian material from Tromso and Oslo, including 


ROCKALL TROUGH ECHINODERMS 3 


189 


A 


C 


u50 
o> 

D 
CT 
O) 


CU 
Q_ 


100 '2 

Oocyte diameter pm 


100 200 300 

Oocyte diameter jum 


400 


Fig. 5 (A) Ovary dissected from a female Myriotrochus bathybius from ES 207 with a calcareous ring 
diameter of 4-5 mm; (B) oocyte diameter frequencies from this specimen (lower) and from a smaller M. 
bathybius from ES 207 (upper); (C) oocyte diameter frequencies from a female of Myriotrochus 
giganteus from ES 207. 


1 90 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

specimens examined by Ostergren, held in the Zoological Museum and also informs us that he 
considers that L. decaria is a valid species. He has also drawn our attention to a single record of L. 
decaria from 1 fathom depth in Port Erin Harbour in the Isle of Man dated 1933 that he was also 
able to re-examine and confirm as conforming to Ostergren's description of L. decaria. With the 
present record from 401 m depth we conclude that Leptosynapta decaria exists as a distinct species 
with a rather wide bathymetric distribution in muddy sediments of the continental shelf and 
adjacent upper slope of the more northerly areas of Europe. It is a small form that may well have 
been overlooked or lost when sieving benthic samples. 

Protankyra brychia (Verrill, 1885) 
SAMPLE. ES 1 18(1). [2871-2925 m] 
DISTRIBUTION. No change. 

Family MYRIOTROCHIDAE 
Myriotrochus bathybius H. L. Clark, 1920 
See: Gage & Billett, 1986: 234-239, figs 1, 3-6, 7A, B, 9A, B, 18B. 
SAMPLES. ES 152(2), ES 185(3), ES 231(6), AT 267(1), AT 282(3), ES 283(2), ES 289(1). [1800-2946 m] 

DISTRIBUTION. The combined bathymetric data for samples of M. bathybius taken in the Rockall 
Trough and on the Porcupine Abyssal Plain gives a total bathymetric range of 1800 to 4310m 
(Gage & Billett, 1986). 

REPRODUCTION. The reference in Part 2 to Gage & Billett (in press) for details of the reproduction 
of this species was incorrect, and further information is therefore provided here. One of the pair of 
branched ovaries is shown in Fig. 5A from an incomplete female specimen with a calcareous ring 
diameter of 4- 5 mm. It contained previtellogenic oocytes measuring up to 160 urn in longest 
diameter (Fig. 5B). In a smaller specimen from the same sample (ES 207) the maximum oocyte size 
did not exceed 60 um (Fig. 5B). 

Myriotrochus giganteus H. L. Clark, 1920 
See: Gage & Billett, 1986: 239-247, figs 1, 7C, 8, 9C, 10-12, 24B. 
SAMPLES. ES 169(1), ES 231(1), ES 283(1). [2898-2946 m] 

DISTRIBUTION. The new records provide a slight extension of the lower bathymetric limit within the 
Trough. The total known bathymetric range is 2898 to 3800 m. 

REPRODUCTION. The maximum dimension of the irregularly shaped, vitellogenic oocytes of the 
specimen from ES 207 referred to by Gage et al. (19850) is 320 um and not 211 x 169 um as given. 
However, the overall size distribution of the oocytes was, like those for M. bathybius, markedly 
skewed to the left, with the size frequencies peaking in the intervals between 20-60 um (Fig. 5C). 

Myriotrochus clarki Gage & Billett, 1986 

See: Gage & Billett, 1986: 247-252, figs 1, 7D, 9D, 13-17, 18 A; also Gage et al. 1985a: 203 (as Myriotrochus 
sp.) 

SAMPLES. ES 90(1), ES 185(1), ES 218(1), SBC 222(71), ES 231(2, ?1), ES 232(1), ES 255(1). [c. 1040-2907 m] 

DISTRIBUTION. These new records increase the previously known bathymetric range of 1605- 
2515m given in part 2. 

Prototrochus zenkevitchi rockallensis Gage & Billett, 1986 

See: Gage & Billett, 1986: 252-259, figs 1,7E, F, 18C-E, 19-23,24A;Gagee/a/., \985a:2Q4(asP.zenkevitchi 
subsp.) 


ROCKALL TROUGH ECHINODERMS 3 191 

SAMPLES. ES 56(1), ES 1 18(3), ES 129(7), ES 135(15), ES 147(2), SBC 156(2), SBC 159(1), SBC 160(1), SBC 
163(1), SBC 168(3), ES 176(23), ES 184(1), ES 185(3), ES 197(36), ES 204(3), SBC 215(71), SBC 216(2), ES 
218(8), SBC 220(3), SBC 222(2), ES 231(1), ES 232(3), SBC 275(1), SBC 276(1), ES 283(15), ES 285(2), ES 
289(5). [c. 1 000-2946 m] 

DISTRIBUTION. The new records provide a slight increase in the lower bathymetric limit. Other 
subspecies of P. zenkevitchi, however, occur much deeper at between 7400 and 9735 m. A bathy- 
metric range of 1000-9735 m is unprecedented in deep-sea holothurians. Gage & Billett (1986) 
suggest that the wide geographic and bathymetric separation of such records of myriotrochid 
species results from the problems of sampling infauna in the deep sea. Possibly, characterisation of 
separate species will be possible given more material. 

REMARKS. Developing wheels, still lacking the rim, and with developing, spike-like spokes, have 
been found amongst the fully developed wheels on a specimen from SBC 220. The largest of the 
specimens from SBC 220 is complete, the sac-like body, length 2-36 mm, being distended with 
sediment (Fig. 3B); some of these being calcareous particles that are quite large compared to the 
size of the animal. 

REPRODUCTION. Gonads have not been found in any of the specimens examined by us, perhaps 
indicating that they are immature. 

Parvotrochm belyaevi Gage & Billett, 1986 

See: Gage & Billett, 1986: 262-266, figs 1, 24C-F, 26, 27; Gage et al., 1985a: 204 (as Myriotrochidae gen. et 
sp.) 

SAMPLES. ES 99(2), ES 147(2), ES 152(3), ES 172(3), ES 204(2), SBC 220(1), ES 285(4). [1 160-2921 m] 

DISTRIBUTION. These new records are mainly from the type locality at c. 2900m depth in the 
southern Rockall Trough. The record from ES 99 in 1 160 m extends the known bathymetric range 
upwards by more than 500 m. 

Order MOLPADIIDA 

Family MOLPADIIDAE 
Cherbonniera utriculm Sibuet, 1974 

SAMPLES. ES 27(242), ES 56(218), ES 111(183), ES 118(74), ES 129(151), SBC 174(1), ES 185(544), ES 
190(161), ES 204(144), ES 23 1(667), 8283(1180), ES 285(1 16). [25 15-2946 m] 

DISTRIBUTION. The records from ES 283 provide a slight extension of the lower bathymetric range 
within the Trough. The total bathymetric range is 2039-4251 m. 

REPRODUCTION. The oocytes grow to a maximum diameter of 200 um, and Tyler et al. (1987) 
suggest that development may be planktotrophic. Too few specimens were examined to determine 
any possible periodicity in reproduction. 

Molpadia blakei (Theel, 1886) 
SAMPLES. ES 283(1), AT 288(2), 3/85/5 OTSB(l). [1991 m to 29702980 m] 

DISTRIBUTION. These records extend the lower bathymetric limit slightly within the Trough and 
also provide a slight extension northwards. 

Molpadia borealis M. Sars, 1858 
SAMPLE. AT 107A(1 [No. of specimens omitted in Part 2]). [c. 2000m] 

Family CAUDINIDAE 
Hedingia albicans (Theel, 1886) 

See: Heding, 1935: 65-67, figs 18, 19, pi. 4, fig. 9, pi. 5, fig. 17, pi. 8, fig. 10 (as Haplodactyla albicans). 
SAMPLES. ES 252(1), ES 255(2), AT 256(1). [1510-1706 m] 


1 92 HARVEY, GAGE, BILLETT, CLARK & PATERSON 

DISTRIBUTION. Eastern and western Atlantic from offNW. Africa (3200 m), southwest of Iceland 
( 1 590- 1 628 m) and the eastern seaboard of N. America (1600-2423 m). Also known in the 
Mediterranean, off southern India and in the Bay of Bengal (484-8 14m). A variety, var. glabra 
Theel, 1886a is known from off New Zealand (1280 m). The present material increases the upper 
bathymetric limit in the Atlantic to 1510m, with the total worldwide range 494-3200 m. 

REMARKS. Only a few specimens are known from each of these widely separated localities. This may 
be the result of inefficient sampling of large infaunal animals by trawls and epibenthic sledges. The 
specimen from AT 256 measures c. 27 mm in length. Specimens measuring up to 45 mm in length 
were found by Heding ( 1 935) to have undeveloped gonads and were hence probably juveniles. The 
deposits in the body wall of our 27 mm specimen lacked both the trilobed appearance and the 
thorny spines on the column of some of those figured by Heding (1935, fig. XVIII), possibly 
because of the relatively small size of the present specimen. It is possible however, that distinct 
sub-species may be recognisable when more material becomes available. 

Discussion 

Gage et al. (1983, 19850) found a higher species richness amongst the relatively small number of 
samples taken on the western side of the Trough, despite the much higher sampling effort on the 
eastern side. This was thought to be related to stronger currents in the west (Jones et al., 1 970; Ellett 
& Roberts, 1973; Roberts, 1975; Lonsdale & Hollister, 1979) favouring microphagous suspension 
feeders. Many of the species found only in the west, particularly ophiuroids and asteroids, were 
inferred to feed on current-borne particles. In general these distributions have been confirmed by 
subsequent sampling but the following species have now been found to have a wider distribution: 
Hoplaster spinosus, Psilaster andromeda, Ophiacantha crassidens and Amphiophiura saurura. 

Bathymetric zonation for the most abundant echinoderm species in the samples covered by Parts 
1 & 2 is summarised by Gage et al. 19856. These authors also employed the coincidence-of-range 
statistic of Backus et al. ( 1 965) to describe the rate of change in the joint bathymetric ranges of these 
species at 100-m depth intervals along a notional transect encompassing most of the sampling 
effort in the east of the Trough. Peaks in the value of this statistic occur at around 800-1200 and 
1800 m depth and were thought to be related to discontinuities in hydrodynamic and water-mass 
properties. 

Despite the frequent sampling in the Rockall Trough since 1973, the echinoderm fauna is still 
imperfectly known, particularly on the southern Feni Ridge on the west side of the Trough from 
which only one sample has been taken. Interestingly, seven of the taxa recorded in this series of 
papers were only found at this station. From a total of 164 echinoderm taxa listed in Parts 1-3, 66 
are recorded from less than 10 specimens and 22 of these are from single specimens. Some of these 
'rarities' can be explained by the wide depth and geographic distribution of our samples, coupled 
with a low sampling effort at most stations. It is clear, however, from the more heavily sampled 
stations where the bottom is thought to be relatively homogeneous, that there are a number of 
relatively rare species which are widely distributed in the Atlantic z.g.Hymenaster gennaeus. The 
additional species recovered on the Hebridean Slope in 1985 with the semi-balloon otter trawl may 
reflect the greater catching power of this gear, with an estimated swept path width of 8-5 m 
compared with either the Agassiz Trawl (3 m) or the epibenthic sledge (1 m). 

Given the high diversity in the echinoderm fauna from the Feni Ridge revealed by the relatively 
low sampling effort so far, it is clear that further sampling is necessary. The clearer picture of 
echinoderm distribution resulting from this would help in evaluating the relative importance of 
sedimentary and hydrographic features in bringing about the contrasts in the echinoderm faunas 
of the eastern and western margins of the Trough. Until this can be accomplished, the echinoderm 
fauna, and its zonation on the relatively well sampled and comparatively gently sloping Hebridean 
Slope, should not be viewed as being typical of the margins of the Trough as a whole. 

Summary 

Five species of crinoids, eleven asteroids, eight ophiuroids, two echinoids and eight species of 


ROCKALL TROUGH ECHINODERMS 3 


193 


holothurian are identified, mainly from sampling carried out between 1983 and 1985 in the deep 
water areas to the west of the British Isles. The following echinoderm species have not been 
recorded previously from this area of the NE. Atlantic. 

Cheiraster sepitus 
Mediaster bairdi 
Pter aster (Apterodon) sp. 
Hymenaster regalis 
Ellipinion delagei 
Leptosynapta decaria 

Of these, Mediaster bairdi and Hymenaster regalis are new records for the eastern Atlantic, while 
the Pteraster (Apterodon) sp. is probably new to science. 

This study confirms the generally higher species richness along the western margin of the Trough 
noted in earlier papers in this series and the tendency for juveniles of many species to be distributed 
at the lower end of the bathymetric range of those species. Extensions to the 
bathymetric and geographic ranges of several species are provided by these records. 

Acknowledgements 

We again express our gratitude to the officers and crew of RRS Challenger, and to colleagues too numerous to 
name individually who participated in cruises and helped with the processing of samples. We are particularly 
indebted to Miss B. Rae of Dervaig, Isle of Mull, for her help in sorting sled and box-core samples; to Mr G. 
Davies and Mr A MacArthur for the sorting of other samples as summer students in the years 1983-85, and to 
Mrs Margaret Pearson for her valuable help in sorting and careful curation of the collections up to 1 983. Dr J. 
D. M. Gordon of SMBA kindly provided additional material from fishing cruises in which the authors did not 
participate. We also thank Dr J. M. Graham of SMBA for constructing a digitising caliper used to measure 
echinoids. The Scottish Marine Biological Association is grant-aided by the Natural Environment Research 
Council who have been generous in allocating cruise time on which the present records were made. 


Station List 

Only details of stations yielding records of echinoderms not listed by Gage et al. (1983, 19850) are 
given below. 


Station No. 


Date 


Position 

(at mid-point of 

track on bottom 

if applicable) 


Depth 
(m) 


Benthic stations 

SBC 156 
SBC 159 
SBC 160 
SBC 168 

ES231 

ES232 
AT 233 
RMT 234 
AT 239 

ES244 
AT 245 
AT 247 
AT 248 
AT 249 
ES250 


5 Aug. 1979 

8 Aug. 1979 

8 Aug. 1979 

13 Aug. 1979 

17 May 1983 

19 May 1983 

19 May 1983 

20 May 1983 

24 July 1983 

25 July 1983 

26 July 1983 

27 July 1983 

27 July 1983 

28 July 1983 
28 July 1983 


4827'N, 1021'W 1310 

5055'N, 1221'W 2036 

5055'N, 1220'W 2030 

5644'N,0913'W 1206 

5442'N, 1212'W 2898 

5717'N, 1016'W 2195 

5717'N, 1012'W 2180 
5712'N,0954'W c. 2000 

5707'N, 0923'W 1047 

5723'N, 1020'W 2150 

5721'N, 1021'W 2165 

5902'N, 1055'W 2084 

5959'N, 1033'W 1150 

5944'N, 1236'W 1265 

5943'N, 1233'W 1270 


194 


HARVEY, GAGE, BILLETT, CLARK & PATERSON 


Station No. 


Date 


Position 
(at mid-point of 
track on bottom 
if applicable) 


Depth 

(m) 


AT 251 


30 July 1983 


5852'N, 1256'W 


1530 


ES252 


30 July 1983 


5852'N, 1253'W 


1510 


AT 254 


31 July 1983 


5826'N, 1235'W 


1595 


ES255 


31 July 1983 


5826'N, 1242'W 


1595 


AT 256 


31 July 1983 


5756'N, 1221'W 


1705 


ES257 


31 July 1983 


5755'N, 1218'W 


1700 


RD258 


1 Aug. 1983 


5756'N, 1324'W 


135 


AT 259 


1 Aug. 1983 


5727'N, 1252'W 


1041 


ES261 


lAug. 1983 


5724'N, 1205'W 


1824 


ES264 


2 Aug. 1983 


5626'N, 1331'W 


2144 


ES266 


3 Aug. 1983 


5624'N, 1 159'W 


2591 


AT 267 


3 Aug. 1983 


5624'N, 1 158'W 


2605 


AT 271 


4 Aug. 1983 


5639'N, 1035'W 


2255 


SBC 272 


5 Aug. 1983 


5640'N, 1030'W 


2250 


AT 273 


5 Aug. 1983 


5605'N, 1028'W 


2185 


SBC 275 


6 Aug. 1983 


5613'N, 1006'W 


1961 


SBC 276 


6 Aug. 1983 


5614'N,0951'W 


1792 


SBC 278 


6 Aug. 1983 


5615'N,0946'W 


1631 


AT 282 


14 April 1985 


5506'N, 1122'W 


c. 2760 


ES283 


15 April 1985 


5439'N, 1215'W 


2946 


AT 284 


15 April 1985 


5440'N, 1212'W 


2906 


ES285 


15 April 1985 


5439'N, 1214'W 


2906 


AT 286 


16 April 1985 


5444'N, 1217'W 


2896 


AT 287 


18 April 1985 


5643'N,0921'W 


1383 


AT 288 


20 April 1985 


5718'N, 1022'W 


2190 


ES289 


21 April 1985 


5719'N, 1025'W 


2190 


AT 290 


26 April 1985 


5628'N,0916'W 


970 


AT 291 


27 April 1985 


5622'N,0912'W 


775 


AT 292 


27 April 1985 


5623'N, 0908'W 


525 


Fishing stations 


Starting position 


Depth range 


13/83/1OTSB 


20 Sept. 1983 


5633'N, 0940'W 


1540-1550 


1 3/83/2 OTSB 


21 Sept. 1983 


5646'N,0915'W 


1130-1265 


1 3/83/3 GT 


21 Sept. 1983 


5636'N, 0902'W 


220-270 


1 3/83/4 GT 


21 Sept. 1983 


5632'N, 0905'W 


230-380 


13/83/5 OTSB 


21 Sept. 1983 


5641'N,0947'W 


1775-1835 


13/83/6 OTSB 


22 Sept. 1983 


5636'N,0917'W 


980-1005 


13/83/7 OTSB 


22 Sept. 1983 


5627'N,0910'W 


750-800 


13/83/8 OTSB 


22 Sept. 1983 


5620'N, 0908'W 


500-560 


9/84/1 OTSB 


2 Nov. 1984 


5632'N,0913'W 


760-815 


9/84/2 OTSB 


2 Nov. 1984 


5634'N,0916'W 


910-960 


9/84/9 OTSB 


4 Nov. 1984 


5647'N, 0936'W 


1750-1770 


9/84/10 OTSB 


4 Nov. 1984 


5616'N,0913'W 


580-630 


9/84/1 3 OTSB 


5 Nov. 1984 


5625'N,0916'W 


940-975 


3/85/5 OTSB 


15 April 1985 


5427'N, 1225'W 


2970-2980 


3/85/7 OTSB 


16 April 1985 


5547'N, 1052'W 


2500-2455 


3/85/8 OTSB 


17 April 1985 


5627'N,0917'W 


970-1010 


3/85/9 OTSB 


17 April 1985 


5643'N,0911'W 


945-1010 


3/85/10 OTSB 


17 April 1985 


5631'N,0913'W 


795-805 


3/85/11 OTSB 


18 April 1985 


5633'N, 0926'W 


1250-1270 


3/85/13 OTSB 


18 April 1985 


5628'N,0917'W 


960-995 


ROCKALL TROUGH ECHINODERMS 3 


195 


Station No. 


Date 


Position 
(at mid-point of 
track on bottom 
if applicable) 


Depth 
(m) 


3/85/14 OTSB 


18 April 1985 


5630'N,0912'W 


720-775 


3/85/17 OTSB 


21 April 1985 


5654'N, 1000'W 


1955-1995 


3/85/18 MBA 


21 April 1985 


5627'N,0917'W 


990-1020 


3/85/19 MBA 


22 April 1985 


5634'N,0918'W 


1030-1035 


3/85/20 OTSB 


22 April 1985 


5634'N, 0926'W 


1225-1245 


3/85/21 OTSB 


22 April 1985 


5631'N,0939'W 


1480-1500 


3/85/22 MBA 


22 April 1985 


5625'N,0917'W 


995-1020 


3/85/23 OTSB 


23 April 1985 


5625'N,0918'W 


995-1000 


3/85/24 OTSB 


23 April 1985 


5626'N,0917'W 


980-990 


3/85/25 OTSB 


23 April 1985 


5625'N,0918'W 


1000-1005 


3/85/26 OTSB 


23 April 1985 


5625'N,0916'W 


940-985 


3/85/27 OTSB 


24 April 1985 


5624'N,09 18'W 


990-1000 


3/85/28 OTSB 


24 April 1985 


5635'N,0918'W 


990-1075 


3/85/29 OTSB 


24 April 1985 


5650'N,0931'W 


1690-1740 


3/85/30 OTSB 


24 April 1985 


5629'N, 0938'W 


1420-1480 


3/85/31 OTSB 


25 April 1985 


5624'N,0917'W 


995-1020 


3/85/32 OTSB 


25 April 1985 


5625'N,0919'W 


1055-1060 


3/85/33 OTSB 


25 April 1985 


5626'N,0918'W 


985-1000 


3/85/34 OTSB 


25 April 1985 


5624'N,0918'W 


980-990 


3/85/36 OTSB 


26 April 1985 


5624'N,0919'W 


1000-1025 


3/85/37 OTSB 


26 April 1985 


5625'N,09 18'W 


945-985 


3/85/38 OTSB 


26 April 1985 


5623'N, 0908'W 


410-490 


3/85/43 OTSB 


27 April 1985 


5617'N,0912'W 


565-700 


3/85/44 OTSB 


27 April 1985 


5618'N,0911'W 


545-600 


3/85/45 OTSB 


28 April 1985 


5630'N, 0938'W 


1470-1500 


3/85/46 OTSB 


28 April 1985 


5625'N,0917'W 


960-985 


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Manuscript accepted for publication 3 June 1987 


British Museum (Natural History) 

The birds of Mount Nimba, Liberia 


Peter R. Colston & Kai Curry-Lindahl 


For evolution and speciation of animals Mount Nimba in Liberia, Guinea and the Ivory Coast is 
a key area in Africa representing for biologists what the Abu Simbel site in Egypt signified for 
archaeologists. No less than about 200 species of animals are endemic to Mount Nimba. Yet, this 
mountain massif, entirely located