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

The Bulletin of the British Museum (Natural History], instituted in 1949, is issued in four scientific
scries. 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
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
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,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
England.

World List abbreviation: Bull Br. Mm. not. 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

ISSN 0 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

A2

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.

A2v

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 Atp; Ata 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 aaP

25

lo

Fig. 16 Merlucciidae: Merluccius merluccius; cranial muscles in lateral view; above, entire; centre,
palatine portion of muscle Atp (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.

tA2

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 Atp 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 AtP and its associations in: A, Ophidian rochei (muscle Aj + A2 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 A2 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 '9dh

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.

References

Alexander, R. McN. 1969. The functions and mechanisms of protrusible jaws of some acanthopterygian
fishes. Journal of Zoology, London 151: 43-64.

Allis, E. P. 1897. The cranial muscles and cranial and first spinal nerves in Amia calva. Journal of Morphology
12: 487-772.

Anker, G. Ch. 1974. Morphology and kinetics of the head of the stickleback Gasterosteus aculeatus. Trans-
actions of the Zoological Society of London 32: 311-416.

Casinos, A. 1978. The comparative feeding mechanism of Gadidae and Macrouridae. 1. Functional
morphology of the feeding apparatus. Gegenbaurs Morphologisches Jahrbuch, Leipzig, 124 (3): 434-449.

- 1981. The comparative feeding mechanism of Gadidae and Macrouridae. II. Mechanics of the feeding
action. Gegenbaurs Morphologisches Jahrbuch, Leipzig, 127 (2): 246-262.

Cohen, D. M. 1984. Gadiformes: overview. In Ontogeny and systematics of fishes Special Publication No. 1 of

the American Society of Ichthyologists and Herpetologists: 259-265.
Dietz, P. A. 1921 . Uber die systematische Stellung der Gadidae. Zugleich Nr. 2 der 'Beitrage zur Kenntnis der

Kiefer- und Kiemenbogenmuskulatur der Teleostier'. Anatomische Anzeiger 144: 56-64.
Fahay, M. P. & Markle, D. F. 1984. Gadiformes: development and relationships. In Ontogeny and systematics

of fishes Special Publication No. 1 of the American Society of Ichthyologists and Herpetologists: 265-283.
Fink, W. L. & Weitzman, S. H. 1982. Relationships of the stomiiform fishes (Teleostei) with a description of

Diplophus. Bulletin of the Museum of Comparative Zoology, Harvard 150: 31-93.
Fraser, T. H. 1972. Some thoughts about the teleostean fish concept — the Paracanthopterygii. Japanese

Journal of Ichthyology 19 (4): 232-242.
Freihofer, W. C. 1978. Cranial nerves of a percoid fish, Polycentrus schomburgkii (Family Nandidae), a

contribution to the morphology and classification of the order perciformes. Occasional Papers of the

California Academy of Sciences 128: 1-78.
Geistdoerfer, P. 1975. Ecologie alimentaire des Macrouridae (Telesteens. Gadiformes). These de Doctorat

D'Etat Es-Sciences Naturelles, 1'Universite de Paris VI, 315 pp.
Gilbert, C. H. & Hubbs, C. L. 1916. Report on the Japanese macrouroid fishes collected by the United States

fisheries Steamer 'Albatross' in 1906, with a synopsis of the genera. Proceedings of the United States

National Museum 51: 135-214.
Gosline, W. A. 1963. Considerations regarding the relationships of the percopsiform, cyprinodontiform and

gadiform fishes. Occasional Papers of the Museum of Zoology, University of Michigan No. 629: 1-38.

- 1981. The evolution of the premaxillary protrusion system in some teleostean fish groups. Journal of
Zoology, London 193: 1 1-23.

Greenwood, P. H. 1977. Notes on the anatomy and classification of elopomorph fishes. Bulletin of the British
Museum of Natural History (Zoology) 32: 65-102.

MACROUROID FISHES 61

- 1985. Notes on the anatomy and phyletic relationships of Hemichromis Peters, 1858. Bulletin of the

British Museum (Natural History) (Zoology) 48 (3): 131-171.
Holmquist, O. 1911. Studien in der von den Nerven trigeminus und facialis innervierten Muskulatur der

Knochen fische. 1. Die Trigemino-Facialis-Muskulatur bei Gadus callarias L. II. Zur vergleichende

Morphologic der Mm. Intermandibularis, protractor hyoidei und hyohyoideus. Acta Universitatis

Lundensisl (7): 1-79.
Howes, G. J. 1984«. A review of the anatomy, taxonomy and biogeography of the African neoboline cyprinid

fishes. Bulletin of the British Museum (Natural History) (Zoology) 47 (3): 151-185.

— 19846. The cranial muscles of loricarioid catfishes, their homologies and value as taxonomic characters
(Teleostei: Siluroidei). Bulletin of the British Museum (Natural History) (Zoology) 45 (6): 309-345.

— 1985. Cranial muscles of gonorynchiform fishes, with comments on generic relationships. Bulletin of the
British Museum (Natural History) (Zoology) 49 (2): 273-303.

— 1987. The palatine bone and its associations in gadoid fishes. Journal of Fish Biology 31: 625—637.
1988. Phylogenetic relationships of macrouroid and gadoid fishes based on cranial myology and

arthrology. WOGADS Symposium. Contributions in Science. Los Angeles County Museum. (In press).
Inada, T. 1 98 1 . Studies on the merlucciid fishes. Bulletin of the Far Seas Fisheries Research Laboratory, no. 1 6:

1-172.

Isaacs, J.D. & Schwartzlose, R. 1975. Active animals of the deep-sea floor. Scientific American 233 (4): 84-91 .
lawmoto, T. & Stein, D. L. 1973. A systematic review of the rattail fishes (Macrouridae: Gadiformes) from

Oregon and adjacent waters. Occasional Papers of the California Academy of Sciences 111: 1-79.
Lauder, G. V. 1981. Intraspecific functional repertoires in the feeding mechanism of the characoid fishes

Lebiasina, Hopliasand Chalceus. Copeia 1981: 154-168.

— 1982. Patterns of evolution in the feeding mechanism of actinopterygian fishes. American Zoologist 22:

275-285.

1983. Functional design and evolution of the pharyngeal jaw apparatus in euteleostean fishes. Zoological

Journal of the Linnean Society 77 (1): 1-38.
Lauder, G. V. & Liem, K. F. 1 980. The feeding mechanism and cephalic myology of Salvelinusfontinalis; form,

function and evolutionary significance. In: E. K. Balon (Ed.) Charrs: Salmonid fishes of the genus Salvelinus:

365-390.
- 1983. The evolution and interrelationships of the Actinopterygian fishes. Bulletin of the Museum of

Comparative Zoology, Harvard 150 (3): 95-197.
Liem, K. F. 1980. Acquisition of energy by teleosts: Adaptive mechanisms and evolutionary patterns. In:

M. A. Ali (Ed.) Environmental physiology of fishes: 299-334.

McLellan, T. 1977. Feeding strategies of the macrourids. Deep Sea Research 24: 1019-1036.
Marshall, N. B. 1965. Systematic and biological studies of the Macrourid fishes (Anacanthini-Teleostii) Deep

Sea Research 12: 299-322.

— 1966. The relationships of the anacanthine fishes Macruronus, Lyconus and Steindachneria. Copeia 1966:
275-280.

— 1973. Genus Coryphaenoides. Family Macrouridae. Memoir, Sears Foundation for Marine Research 1.
Fishes of the western North Atlantic (6): 565-580.

— 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,
Sears Foundation for Marine Research 1. Fishes of the western North Atlantic (6): 479-495.

& Merrett, N. R. 1977. The existence of a benthopelagic fauna in the deep-sea. In: A voyage of Discovery

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

Mauchline, J. & Gordon, J. D. M. 1 985. Trophic diversity in deep-sea fish. Journal of Fish Biology 26: 527-535.
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.

Motta, P. J. 1984. Mechanics and functions of jaw protrusion in teleost fishes: a review. Copeia 1984: 1-18.
Okamura, O. 1970a. Macrourina. Fauna Japonica. Japanese Biogeographical Society, Tokyo, 216 pp.

- 19706. Studies on the Macrouroid fishes of Japan— morphology, ecology and phylogeny. Reports of the

Usa Marine Biological Station, Kochi University 17 (1-2): 1-179.
Osse, J. W. M. & Muller, M. 1980. A model of suction feeding in teleostean fishes. In: M. A. Ali (Ed.)

Environmental Physiology of fishes: 335-352.
Otten, E. 1981. Vision during growth of a generalized Haplochromis species: H. elegans Trewavas 1933

(Pisces, Cichlidae). Netherlands Journal of 'Zoology 31 (4): 650-700.

62 G. J. HOWES

- 1982. The development of a mouth-opening mechanism in a generalized Haplochromis species:
H. elegans Trewavas 1933 (Pisces, Cichlidae). Netherlands Journal of Zoology 32 (1): 31-48.

1983. The jaw mechanism during growth of a generalized Haplochromis species: H. elegans Trewavas

1933 (Pisces, Cichlidae). Netherlands Journal of Zoology 33(1): 55-98.
Parr, A. E. 1946. The Macrouridae of the western North Atlantic and central American seas. Bulletin of the

Bingham Oceanographic Collection 10 article 1: 1-93.
Patterson, C. 1975. The braincase of pholidophorid and leptolepid fishes, with a review of the actinopterygian

braincase. Philosophical Transactions of the Royal Society of London B 269 (899): 275-579.
Peabody, E. B. 193 1 . Scales of fishes of the order Anacanthini. University of Colorado Studies 18 (3): 1 33-1 50.
Rosen, D. E. 1962. Comments on the relationships of the North American cave fishes of the family

Amblyopsidae. American Museum Novitates 2109: 1-35.
- 1973. Interrelationships of higher euteleostean fishes: In: Interrelationships of fishes. Eds Greenwood,

P. H., Miles, R. S. & Patterson, C. Supplement No. 1 to the ZoologicalJournal of the Linnean Society 53:

397-513.

& Patterson, C. 1969. The structure and relationships of the paracanthopterygian fishes. Bulletin of the

American Museum of Natural History 141 (3): 357-474.
Smith, K. L. & Messier, R. R. 1974. Respiration of benthopelagic fishes: In situ measurements at 1230 Metres.

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

Lakes Malawi and Tanganyika. Bulletin of the British Museum (Natural History) (Zoology) 40 (3): 67-101 .
- 1986. The limits and relationships of the acanthomorph teleosts. Journal of Zoology, London (B) 1:

411-460.
Tchernavin, V. V. 1953. The feeding mechanism of a deep-sea fish Chauliodus sloani Schneider. British Museum

(Natural History) 101 pp.
Verraes, W. 1 977. Postembryonic ontogeny and functional anatomy of the ligamentum mandibulo-hyoideum

and the ligamentum interoperculo-mandibulare, with notes on the opercular bones and some other cranial

elements in Salmo gairdneri Richardson, 1836 (Teleostei: Salmonidae). Journal of Morphology 151 (1):

111-120.

Weihs, D. 1980. Hydrodynamics of suction feeding offish in motion. Journal of Fish Biology 16: 425-433.
Wilson, R. R. & Smith, K. L. 1985. Live capture, maintenance and partial decompression of a deep-sea

genadier fish (Coryphaenoides acrolepis) in a hyperbaric trap-aquarium. Deep Sea Research 32 (12):

1571-1582.
Winterbottom, R. 19740. A descriptive synonymy of the striated muscles of the Teleostei. Proceedings of the

Academy of Natural Sciences, Philadelphia 125: 225-317.

— 1974£. The familial phylogeny of the Tetraodontiformes (Acanthopterygii: Pisces) as evidenced by their

comparative myology. Smithsonian Contributions to Zoology 155: 1-210.

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. 0 565 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
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
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,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
England.

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 0 565 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 0

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

333 3

or 1---1

550 3

femur 2—2 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 0

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 plt 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 0 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

Athias-Henriot, C. 1968. L'appareil d'insemination laelapo'ide (Acariens anactinotriches: Laelapoidea, $?).
Premieres observations. Possibilite d'emploi a des fins taxonomiques. Bull, scient. Bourgogne 25: 229-274.

— 1969. Notes sur la morphologic externe des gamasides (Acariens anactinotriches). Acarologia 11:
609-629.

Axtell, R. C. 1961. New records of North American Macrochelidae (Acarina: Mesostigmata) and their
predation rates on the housefly. Ann. ent. Soc. Am. 54: 748.

— 1963. Acarina occurring in domestic animal manure. Ann. ent. Soc. Am. 56: 628-633.

Balogh, J. 1958. Macrocheliden aus Bulgarien (Acari, Mesostigmata). Acta ent. Mus. natn. Pragae 32:

247-256.
Berlese, A. 1904. Acari Nuovi. Redia 2: 10-32.

- 1918. Centuria quarta di Acari nuovi. Redia 13: 1 13-190.
Bregetova, N. G. & Koroleva, E. V. 1960. The macrochelid mites (Gamasoidea, Macrochelidae) in the USSR.

Parazit. 56.19:32-154.
Bregetova, N. G. et al. 1977. Key to soil-inhabiting mites. Mesostigmata. Akad. Nauk SSSR, Leningrad, 718

pp. [Russian].

Canestrini, G. & Canestrini, R. 1882. 1 gamasi Italiani. AttiSoc. ven.-trent. sci. 8: 31-82.
Canestrini, G. & Fanzago, F. 1877. Intorno agli Acari italiani. Atti R. 1st. veneto Sci. (5) 4: 69-208.
Costa, M. 1963. The mesostigmatic mites associated with Copris hispanus (L.) (Coleoptera, Scarabaeidae) in

Israel. J. Linn. Soc. (Zool.) 45: 25-45.

— \966a. Notes on macrochelids associated with manure and coprid beetles in Israel. I. Macrocheles
robustulus (Berlese, 1904), development and biology. Acarologia 8: 532-548.

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

- 19666. The present stage of knowledge of mesostigmatic mites in Israel (Acari, Mesostigmata). IsraelJ.
Zool. 15: 69-82.

- 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

Ent. 5: 118-127.

— 1973/>. Additions to the macrochelid mites in New Zealand (Acarina: Mesostigmata: Macrochelidae)

N.Z. Ent. 5: 294-302.
Evans, G. 0. 1 956. On the classification of the family Macrochelidae with particular reference to the subfamily

Parholaspinae (Acarina: Mesostigmata). Proc. zool. Soc. Land. 127: 345-377.

- 1963. Observations on the chaetotaxy of the legs in the free-living Gamasina (Acari: Mesostigmata).
Bull. Br. Mus. not. Hist. (Zool.) 10: 277-303.

- 1 964. Some observations on the chaetotaxy of the pedipalps in the Mesostigmata (Acari). Ann. Mag. nat.
Hist. (13)6:513-527.

- & Browning, E. 1956. British mites of the subfamily Macrochelinae Tragardh (Gamasina, Macro-
chelidae). Bull. Br. Mus. nat. Hist. (Zool.) 4: 1-55.

- & Till, W. M. 1965. Studies on the British Dermanyssidae (Acari: Mesostigmata). Part I. External
morphology. Bull. Br. Mus. nat. Hist. (Zool.) 13: 247-294.

& - - 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.
Falconer, W. 1923. Two British mites new to science and a new subgenus of Macrocheles Latr. Naturalist, Hull

1923: 151-153.
Filipponi, A. & Ilardi, A. 1958. Sulla validita di tre specie del sottogenere Berlesiano Macrocheles (Acarina,

Mesostigmata). Riv. Parassit. 19: 1 17-130.
Filipponi, A. & Pegazzano, F. 1960. Acari del genere Glyptholaspis nom. nov. pro Macrocheles (Macrocheles)

Berl. 1918 (Mesostigmata, Macrochelidae). Redia45: 133-171.

— & 1962. Specie Italiane del gruppo-glaber (Acarina, Mesostigmata, Macrochelidae, Macrocheles).

Redia 47: 211-23%.

— & — — 1963. Specie Italiane del gruppo-subbadius (Acarina, Mesostigmata, Macrochelidae). Redia 48:
69-91.

— & 1967. Contribute alia conoscenza del genere Holostaspella Berlese, 1903. (Acari: Mesostigmata:

Macrochelidae). Redia 50: 219-259.

Franz, H. 1954. Die Nordost-Alpen. 15. Ordnung Acarina, in Spiegel Land-Tierwelt, Innsbruck 1:

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.
M albert. J. N. 1915. Clare Island Survey, Part 39 ii Acarinida: Section II — Terrestrial and marine Acarina.

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.
suppl.: 91-146.

— 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

1918. (Eine systematische Studie aus dem Jahre 1960). Acarologie 14: 2-43.

Leitner, E. 1946. Zur Kenntnis der Milbenfauna auf Diingerstatten. Zentbl. Gesamtgeb. Ent. 1: 75-156.
Lindquist, E. E. & Evans, G. 0. 1965. Taxonomic concepts in the Ascidae, with a modified setal nomenclature

for the idiosoma of the Gamasina (Acarina: Mesostigmata). Mem. ent. Soc. Can. 47: 1-64.
Petrova, A. D. 1960. Materialien iiber den Bau des inneren Sackchens des Receptaculum seminis der

gamasoiden Milben Macrochelidae Vitz. Zool. Anz. 165: 393-400.
- & Taskaeva, E. Z. 1964. Gamasoid mites (Parasitiformes, Gamasoidea) from southern China (1st

Communication). Byull. Mask. Obshch. Ispyt. Prir. (Biol.) 69, 5: 47-61.
Sellnick, M. 1931. Acari. In Max Beier, Zoologische Forschungreise nach den Jonischen Inseln und dem

Peloponnes. XVI Teil. Sher. Akad. Wiss. Wien 140: 693-776.
Smith, K. G. V. 1975. The faunal succession of insects and other invertebrates on a dead fox. Entomologist's

Gaz. 26: 277-287.
Valle, A. 1953. Revisione di generi e sottogeneri Berlesiani di Acari (Primo contributo). Redia 38: 316-360.

— 1955. Revisione dell' Acaroteca Canestrini. Atti Memorie Accad. patavina67: 67-101.
Willmann, C. 1951. Untersuchungen iiber die terrestrische Milbenfauna in pannonischen Klimagebiet

Osterreichs. Sber. Akad. Wiss. Wien 160: 91-176.

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
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. 0 565 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

Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester. Dorset

- f J

GEN

Bulletin of the

British Museum (Natural History)

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

Alan Warren

Zoology series Vol54 No 3 26 May 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
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
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,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
England.

World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.)

ustees of the British Museum (Natural History), 1988

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 0 565 05039 7

ISSN 0007-1 498 Zoology series

Vol54 No. 3 pp 127-1 52
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 26 May 1 988

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.

134

A. WARREN

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 34—49 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.

136

A. WARREN

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.

138

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.

140

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.

142

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.

144

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.

146

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

148

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. 0 565 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
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
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,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
England.

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 : MrJ. F. Peake
Editor of Bulletin : Dr C. R. Curds
Assistant Editor : Mr C. G. Ogden

ISBN 0 565 05040 0

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 AW»ri*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. 50°N,
1 5°55'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

o°w

Bathymetry

200 m
1000 m

2000 rn

••— 3000 IT
AOOOrn

.....

\ A V X y-^Tu'^^ k^rfi

...... ,

^/V/^-X ^

18°W 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. 50°N. 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. 54°N, 12°30'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 0 OTSB( 1 ), 3/85/ 1 0 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. 56°N 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 36°34'N, 73°48'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. 56°N.

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. 56°N 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. 57°N 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 m2 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

175

0 O
ON NO

O O O — O OO — O

O O
oo m

OOOfNO OOOO

O O

OOOrNO rN O — ' fN

_

SO
rn

£

O O — OO O fN rN — f>

a

O O
in rN

2

O O O n-> O <N NO — NO

O O O

u

OmtNrO— >n NO O fN — OOOOOOOO —
rxi — _ _< rs]

•o
u

u

O O O
fNNOooNom ON^>n— inoot^o — ^f — o —

q3

O O O
rN ON OO

O

^^^SIC ^"^^ ^2^^^^^r^°S

«J

"o

O O O
— oo (^

U

£-£^5 ?Q S^r^r^^^So^^

(M

O
T3

C

O O O

f*^ £ — NO

NO^fN >n — ' TJ- ONfNin

0

o

0 0 O

ON NO m

c
<u

u

o

r»i m <N

0 00
oo in Tt

jo

E

u

•^J-OOOrn (NOt^O — 'O — OOOOOoOO

0 00
i-~ Tt m

(NOOOO — O O O OOOOOOO — ^O

1

o o o
NO m tN

— OOOO — O fN O — OOOOOO — OO

«M

c«

o o o
in rN —

"f

3

rs

3

a,
u

a

in ln i/">»n*nm inmino inm^n

ooooo oomo oinooooo'nino
oorj-NOfNO fioorso rot^ooinm^NOmrNO
ON ON ON t — f*> •— ' ON <N <O ^^ t — ON r^ t^ ON ON ON fN ^D

o

_c

f 1

^-frequencies of e<

c
on

ffloaoooQOQ oQCQoacQ cQoacQoaoQoapQcnaacQ
onon&oonon onononon ononononononononon&n
HHHHH HHHH HHHHHHHHHH
OOOOO OOOO OOOOOOOOOO

c*"if*~)^iO O1^ r*"ir*"ir~-OlO
\^Q ~ — — fsj r^ ^o CN c^ ^J cs *o *o r — ON ^^ ^^ ^~* 0s) rs

•"* r*^ ^f to */") i/^ is co ro >O */^ St rn r*"i ro r*^ ^~ ^f l/") *O */~i v^
i: oooooooooo -2 oooooooo ^ op^op^oo^oo^oo^oo^oo^op^op^op^

N

~5 *s ' ^

S3

—

R

H

U

Q

i g 1

•^ N^ O ..'. ^1 O U-. U-. (J-. t-~.

^ oooooooooo o oooooooo S oooooooooooooooooooo
^ ON ^— Tj* "^ ^f ^ ONON^Tt ^ ON ON ON ON — ^ ' — '^Tt^^f^f
Q rN "^ oo oo m .^ — fN rN m ;? — -^ rN fN " . oo — rs c*i
<O fMin^-^^rN Co rxifNfNrN "-. (NfN(NfN-<t'n — fNfNrs

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

179

0

o
•o

o.

o
o

OH C

° 1^

"O lyj

V3 O
O S)

0 o\

"5 cd

|1
fr|

0 03

s^

— 03

n »

•_— M

'C "fi

u i

o>

N S

01 »

1 s

H2

.00 JS
Jd 00

o

ex

fc «

J _§" S rt Q

^ t/3 S II rrj

•^ t. N *y^

.2 J. oo C

fe • C ?3

^3 6

^•?5
2 o oo c

<u S

Qo

CXD
ON

Ch

Cha

# * * #

****

OfNOON^O^O

«— c i— c 1/1

ob — < ON >— i r^ ON

__

.00 cj ac:

^ -S -S 5

^ oo oi oo

v> C C C
(L) (U (U <U

«-> O

2 .2-n

J-H <D

C .*_» c/5 QJ _£^

s .... e e s s -o .2

e;ro3a3cdcdJ3-2

EOt-i^^-i^j^s
ol5«^.22-23 g ««

1 i *^ ^~ -7-1 FT— * . _

^ .c .c 22^ o
.§• -B -a o o^-
fe rt rt c c •§>

±2 S.D.

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. 58°N, 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 57°N 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 7«go//"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 39°N 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. 65°N. 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 0 [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

0

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 2970—2980 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

48°27'N, 10°21'W 1310

50°55'N, 12°21'W 2036

50°55'N, 12°20'W 2030

56°44'N,09°13'W 1206

54°42'N, 12°12'W 2898

57°17'N, 10°16'W 2195

57°17'N, 10°12'W 2180
57°12'N,09°54'W c. 2000

57°07'N, 09°23'W 1047

57°23'N, 10°20'W 2150

57°21'N, 10°21'W 2165

59°02'N, 10°55'W 2084

59°59'N, 10°33'W 1150

59°44'N, 12°36'W 1265

59°43'N, 12°33'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

58°52'N, 12°56'W

1530

ES252

30 July 1983

58°52'N, 12°53'W

1510

AT 254

31 July 1983

58°26'N, 12°35'W

1595

ES255

31 July 1983

58°26'N, 12°42'W

1595

AT 256

31 July 1983

57°56'N, 12°21'W

1705

ES257

31 July 1983

57°55'N, 12°18'W

1700

RD258

1 Aug. 1983

57°56'N, 13°24'W

135

AT 259

1 Aug. 1983

57°27'N, 12°52'W

1041

ES261

lAug. 1983

57°24'N, 12°05'W

1824

ES264

2 Aug. 1983

56°26'N, 13°31'W

2144

ES266

3 Aug. 1983

56°24'N, 1 1°59'W

2591

AT 267

3 Aug. 1983

56°24'N, 1 1°58'W

2605

AT 271

4 Aug. 1983

56°39'N, 10°35'W

2255

SBC 272

5 Aug. 1983

56°40'N, 10°30'W

2250

AT 273

5 Aug. 1983

56°05'N, 10°28'W

2185

SBC 275

6 Aug. 1983

56°13'N, 10°06'W

1961

SBC 276

6 Aug. 1983

56°14'N,09°51'W

1792

SBC 278

6 Aug. 1983

56°15'N,09°46'W

1631

AT 282

14 April 1985

55°06'N, 11°22'W

c. 2760

ES283

15 April 1985

54°39'N, 12°15'W

2946

AT 284

15 April 1985

54°40'N, 12°12'W

2906

ES285

15 April 1985

54°39'N, 12°14'W

2906

AT 286

16 April 1985

54°44'N, 12°17'W

2896

AT 287

18 April 1985

56°43'N,09°21'W

1383

AT 288

20 April 1985

57°18'N, 10°22'W

2190

ES289

21 April 1985

57°19'N, 10°25'W

2190

AT 290

26 April 1985

56°28'N,09°16'W

970

AT 291

27 April 1985

56°22'N,09°12'W

775

AT 292

27 April 1985

56°23'N, 09°08'W

525

Fishing stations

Starting position

Depth range

13/83/1OTSB

20 Sept. 1983

56°33'N, 09°40'W

1540-1550

1 3/83/2 OTSB

21 Sept. 1983

56°46'N,09°15'W

1130-1265

1 3/83/3 GT

21 Sept. 1983

56°36'N, 09°02'W

220-270

1 3/83/4 GT

21 Sept. 1983

56°32'N, 09°05'W

230-380

13/83/5 OTSB

21 Sept. 1983

56°41'N,09°47'W

1775-1835

13/83/6 OTSB

22 Sept. 1983

56°36'N,09°17'W

980-1005

13/83/7 OTSB

22 Sept. 1983

56°27'N,09°10'W

750-800

13/83/8 OTSB

22 Sept. 1983

56°20'N, 09°08'W

500-560

9/84/1 OTSB

2 Nov. 1984

56°32'N,09°13'W

760-815

9/84/2 OTSB

2 Nov. 1984

56°34'N,09°16'W

910-960

9/84/9 OTSB

4 Nov. 1984

56°47'N, 09°36'W

1750-1770

9/84/10 OTSB

4 Nov. 1984

56°16'N,09°13'W

580-630

9/84/1 3 OTSB

5 Nov. 1984

56°25'N,09°16'W

940-975

3/85/5 OTSB

15 April 1985

54°27'N, 12°25'W

2970-2980

3/85/7 OTSB

16 April 1985

55°47'N, 10°52'W

2500-2455

3/85/8 OTSB

17 April 1985

56°27'N,09°17'W

970-1010

3/85/9 OTSB

17 April 1985

56°43'N,09°11'W

945-1010

3/85/10 OTSB

17 April 1985

56°31'N,09°13'W

795-805

3/85/11 OTSB

18 April 1985

56°33'N, 09°26'W

1250-1270

3/85/13 OTSB

18 April 1985

56°28'N,09°17'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

56°30'N,09°12'W

720-775

3/85/17 OTSB

21 April 1985

56°54'N, 10°00'W

1955-1995

3/85/18 MBA

21 April 1985

56°27'N,09°17'W

990-1020

3/85/19 MBA

22 April 1985

56°34'N,09°18'W

1030-1035

3/85/20 OTSB

22 April 1985

56°34'N, 09°26'W

1225-1245

3/85/21 OTSB

22 April 1985

56°31'N,09°39'W

1480-1500

3/85/22 MBA

22 April 1985

56°25'N,09°17'W

995-1020

3/85/23 OTSB

23 April 1985

56°25'N,09°18'W

995-1000

3/85/24 OTSB

23 April 1985

56°26'N,09°17'W

980-990

3/85/25 OTSB

23 April 1985

56°25'N,09°18'W

1000-1005

3/85/26 OTSB

23 April 1985

56°25'N,09°16'W

940-985

3/85/27 OTSB

24 April 1985

56°24'N,09018'W

990-1000

3/85/28 OTSB

24 April 1985

56°35'N,09°18'W

990-1075

3/85/29 OTSB

24 April 1985

56°50'N,09°31'W

1690-1740

3/85/30 OTSB

24 April 1985

56°29'N, 09°38'W

1420-1480

3/85/31 OTSB

25 April 1985

56°24'N,09°17'W

995-1020

3/85/32 OTSB

25 April 1985

56°25'N,09°19'W

1055-1060

3/85/33 OTSB

25 April 1985

56°26'N,09°18'W

985-1000

3/85/34 OTSB

25 April 1985

56°24'N,09°18'W

980-990

3/85/36 OTSB

26 April 1985

56°24'N,09°19'W

1000-1025

3/85/37 OTSB

26 April 1985

56°25'N,09018'W

945-985

3/85/38 OTSB

26 April 1985

56°23'N, 09°08'W

410-490

3/85/43 OTSB

27 April 1985

56°17'N,09°12'W

565-700

3/85/44 OTSB

27 April 1985

56°18'N,09°11'W

545-600

3/85/45 OTSB

28 April 1985

56°30'N, 09°38'W

1470-1500

3/85/46 OTSB

28 April 1985

56°25'N,09°17'W

960-985

References

Backus, R. H., Mead, G. L., Haedrich, R. L. & Ebeling, A. W. 1965. The mesopelagic fishes collected during

cruise 1 7 of the R/V Chain, with a method for analysing faunal transects. Bulletin of the Museum of

Comparative Zoology Harvard 134: 139-158.
Billett, D. S. M., Hansen, B. & Huggett, Q. J. 1985. Pelagic Holothurioidea (Echinodermata) from the

northeast Atlantic, pp. 399^1 1 1 In Keegan, B. F. & O'Connor, B. D. S. [Eds] Echinodermata. Proceedings of

the 5th International Echinoderm Conference, Galway, 1984 Balkema, Rotterdam.
Blake, D. B. 1987. A classification and phylogeny of post-Palaeozoic sea stars (Asteroidea: Echinodermata).

Journal of Natural History 21: 48 1-528.
Brattstrom, H. 1946. Obervations on Brissopsis lyrifera (Forbes) in the Gullmar Fjord. Arkivfor Zoologi 37a

(18): 1-27.
Cherbonnier, G. 1953. Recherches sur les synaptes (holothuries apodes) de Roscoff. Archives de Zoologie

experimentale et generate 90: 163-1 86.

1963. Note sur Leptosynapta bergensis (Ostergren) espece critique d'holothurie apode. Bulletin du

Museum d'Histoire Naturelle (2) 35 (4): 429-440.

Chesher, R. H. 1968. The systematics of sympatric species of West Indian spatangoids: a revision of the genera

Brissopsis, Plethotaenia, Paleopneustes, and Saviniaster. Studies in tropical oceanography No. 7. viii+ 168

pp. Institute of Marine Sciences, University of Miami.
Clark, A. H. 1913. On a collection of recent crinoids from the waters about Ireland. Scientific Investigations,

1912. Fisheries Branch, Department of Agriculture for Ireland, Dublin (24): 1-5.
Clark, A. M. 1970. Echinodermata: Crinoidea. Marine Invertebrates of Scandinavia. No. 3. 55 pp.

Universitetsforlaget, Oslo.

1977. Notes on deep-water Atlantic Crinoidea. Bulletin of the British Museum (Natural History)

Zoology 31: 157-186.

1 96 HARVEY, GAGE, BILLETT, CLARK & PATERSON

— 1980. Crinoidea collected by the Meteor and Discovery in the North-East Atlantic. Bulletin of the British
Museum (Natural History) Zoology 38: 187-210.

- 1981. Notes on Atlantic and other Asteroidea. 1. Family Benthopectinidae. Bulletin of the British
Museum (Natural History) Zoology 41(3): 91-135.

1984. Notes on Atlantic and other Asteroidea. 4. Families Poraniidae and Asteropseidae. Bulletin of the

British Museum (Natural History) Zoology 47(1): 19-51.
Clark, H. L. 1907. The apodous holothurians. Smithsonian Contributions to Knowledge 35, No. 1732, 231 pp.
Clark, H. L. 1941. Reports on the scientific results of the Atlantis expeditions to the West Indies, under the

joint auspices of the University of Havana and Harvard University. The Echinoderms (other than

Holothurians). Memoiras de la Sociedad Cubana de Historia Natural Felipe Poey 15( 1 ): 1 54 pp.
Dartnall, A. J., Pawson, D. L., Pope, E. C. & Smith, D. J. 1 969. Replacement name for the preoccupied genus

name Odinia Perrier, 1 885 (Echinodermata: Asteroidea). Proceedings of the Linnaean Society NSW 93: 2 1 1 .
Deacon, M. 1977. Staff-Commander Tizard's journal and the voyages of H.M. Ships Knight Errant and Triton

to the Wyville Thomson Ridge in 1880 and 1882. pp. 1-14. In Angel, M. [Ed.] A Voyage of Discovery.

Pergamon, Oxford.
Deichmann, E. 1930. The holothurians of the western part of the Atlantic Ocean. Bulletin of the Museum of

Comparative Zoology Harvard 71(3): 43-226.
1940. Report on the holothurians, collected by the Harvard-Havana Expeditions 1938 and 1939, with a

revision of the Molpadonia of the Atlantic Ocean. Memoiras de la Sociedad Cubana de Historia Natural

'Felipe Poey' 14: 183-240.

1954. The holothurians of the Gulf of Mexico. Fishery Bulletin of the United States Fish and Wildlife

Service 55: 381-410.
Doderlein, L. 1912. Die gesteilten Crinoiden der Deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse

derDeutschen Tiefsee— aufdem DampferValdivia 1898-1899, 17(1): 1-34.
Downey, M. E. 1973. Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions to

Zoology 126: 1-158.
1986. Revision of the Atlantic Brisingida (Echinodermata: Asteroidea), with description of a new Genus

and Family. Smithsonian Contributions to Zoology 435: 1-57.
Duineveld, G. C. A. & Jenness, M. I. 1 984. Differences in growth rates of the sea urchin Echinocardium

cordatum as estimated by the parameter of the von Bertalanffy equation applied to skeletal rings. Marine

Ecology Progress Series, 19: 65-72.

Einarsson, H. 1948. Echinoderma. The Zoology of Iceland 4(70): 1-67.
Ellett, D. J. & Roberts, D. G. 1973. The overflow of Norwegian deep-sea water across the Wyville Thomson

Ridge. Deep-Sea Research 20: 819-835.
Farran, G. P. 1913. The deep-water Asteroidea, Ophiuroidea and Echinoidea of the west coast of Ireland.

Scientific Investigations, 1912. Fisheries Branch, Department of Agriculture Ireland, Dublin 6: 1-66.
Fell, F. J. 1982. Echinodermata. pp. 785-818 In Parker, S. P. [ed.] Synopsis and Classification of Living

Organisms. McGraw-Hill, New York.
Fisher, W. K. 1911. Asteroidea of the North Pacific and adjacent waters Part 1 . Phanerozonia and Spinulosa.

Bulletin of the United States National Museum 76: 419 pp.
Gage, J. D. (1987). Growth of the deep-sea irregular sea urchins Hemiaster expergitus and Echinosigra phiale

in the Rockall Trough (N. E. Atlantic). Marine Biology 96: 19-30.
& Billett, D. S. M. 1986. The family Myriotrochidae Theel (Echinodermata: Holothurioidea) in the deep

northeast Atlantic Ocean. ZoologicalJournal of the Linnaean Society 88: 229-276.

Jensen, M. & Tyler, P. A. 1985a. Echinoderms of the Rockall Trough and adjacent areas 2.

Echinoidea and Holothurioidea. Bulletin of the British Museum (Natural History) Zoology 48(4): 1 73-2 13.
— , Pearson, M., Billett, D. S. M., Clark, A. M., Jensen, M., Paterson, G. L. J. & Tyler, P. A. 19856.
Echinoderm zonation in the Rockall Trough (NE Atlantic), pp. 31-36 In Keegan, B. F. & O'Connor, B.
[eds] Echinodermata. Proceedings of the Fifth International Echinoderms Conference Galway, 1984.
Balkema, Rotterdam.

Clark, A M., Paterson, G. L. J. & Tyler, P. A. 1983. Echinoderms of the Rockall Trough 1.

Crinoidea, Asteroidea and Ophiuroidea. Bulletin of the British Museum (Natural History) Zoology 45(5):
263-308.

— & Tyler, P. A. 1985. Growth and recruitment of the deep-sea urchin Echinus affinis. Marine Biology 90:
41-53.

& Nichols, D. ( 1 986). Reproduction and growth of Echinus acutus var. norvegicus and E. elegans on

the continental slope off Scotland. Journal of experimental marine Biology and Ecology, 101: 61-83.
Gordon, J. D. M. 1986. The fish populations of the Rockall Trough. Proceedings of the Royal Society of
Edinburgh 88B: 191-204.

ROCKALL TROUGH ECHINODERMS 3 1 97

Gray, I. E., Downey, M. E. & Cerame-Vivas, M. J. 1968. Sea stars of North Carolina. Fishery Bulletin of the

United States Fish and Wildlife Service 67( 1 ): 1 27-1 63.
Halpern, J. A. 1972. Pseudarchasterinae (Echinodermata: Asteroidea) of the Atlantic. Proceedings of the

Biological Society of Washington #5(30): 359-384.
Hansen, B. 1975. Systematics and biology of the deep-sea holothurians. Part 1 . Elasipoda. Galathea Report 13:

1-262.
Harvey, R. & Gage, J. D. 1984. Observations on the reproduction and postlarval morphology of pourtalesiid

sea urchins in the Rockall Trough area (N. E. Atlantic Ocean). Marine Biology 82: 181-190.
Heding, S. G. 1935. Holothurioidea. Part 1. Apoda, Molpadioidea, Gephyrothurioidea. Danish Ingolf

Expedition 4(9): 1-84.

1942. Holothurioidea. Part 2. Aspidochirota, Elasipoda, Dendrochirota. Danish Ingolf Expedition

4(13): 1 39.

Herouard, E. 1902. Holothuries provenant des campagnes de la Princess Alice 1892-1897). Resultats des
Campagnes Scientifique accomplies par le Prince Albert I Monaco 21: 1—61.

1923. Holothuries provenant des campagnes des yachts Princess Alice et Hirondelle II (1898-1915).

Resultats des Campagnes Scientifiques accomplies par le Prince Albert I Monaco 66: 1-1 63.

Jones, E. J. W., Ewing, M., Ewing, J. I. & Eittreim, S. 1970. Influences of Norwegian sea overflow water on

sedimentation in the northern North Atlantic and Labrador Sea. Journal of Geophysical Research 7:

1655-1680.

Koehler, R. 1927. Echinodermes des mers d 'Europe 2. 339 pp. Doin Paris.
Lonsdale, P. & Hollister, C. D. 1979. A near bottom traverse of Rockall Trough: hydrographic and geologic

inferences. Oceanologica Acta 2: 91-105.
Madsen, F. J. 1941. On Thyone wahrbergi n. sp., a new holothurian from the Skagerrak with remarks on T.

fusus (O. F. M.) and other related species. Goteborgs Kungl. Vetenskaps — och Vitterhets-Samhalles

Handlingar F.6, Ser. B., 1(1), 1-31.

1942. Cucumaria hyndmani. The variation of its calcareous deposits. Videnskabelige Meddelelser fra

Dansk Naturhistorisk Forening i Kjobenhavn 105: 395-406.

Maranzeller, E. von 1893. Contribution a 1'etude des holothuries de 1'Atlantique Nord. Resultats des

Campagnes Scientifiques accomplies par le Prince Albert I Monaco 6: 1-22.
Miller, J. E. & Pawson, D. L. 1 984. Holothurians (Echinodermata: Holothuroidea). Memoirs of the Hourglass

Cruises 7(1): 1-79.
Mortensen, T. 1907. Echinoidea (Part 2). Danish Ingolf Expedition. 4(2): 1-200.

1927. Handbook of the Echinoderms of the British Isles. 471 pp. Oxford University Press, London.

1933. Ophiuroidea. Danish Ingolf Expedition 4(2): 1-121.

1943. A Monograph of the Echinoidea Vol. iii(3). Camarodonta. 2. Echinidae, Strongylocentrotidae,

Paraseleniidae, Echinometridae. 446 pp. C.A. Reitzel, Copenhagen.

1951. A Monograph of the Echinoidea Vol. V(2). Spatangoida 2. Amphisternata. 2. Spatangidae,

Loveniidae, Percosmidae, Schizasteridae, Brissidae. 593 pp. C. A. Reitzel, Copenhagen.
Ostergren, H. 1905. Zur kenntnis der skandinavischen und arktischen Synaptiden. Archives de Zoologie

Experimentale et Generate , Ser. 4, 3, Notes et revue, No. 17: 133-164.
Panning, A. 1949. Versuch einer Neuordnung der familie Cucumariidae (Holothurioidea, Dendrochirota).

Zoologische Jahrbucher Jena 78: 404-470.
Paterson, G. L. J. 1985. The deep-sea Ophiuroidea of the North Atlantic Ocean. Bulletin of the British

Museum (Natural History) Zoology 49(1): 1-162.
Pawson, D. L. 1982. Deep-sea echinoderms in the Tongue of the Ocean, Bahama Islands: a survey, using the

research submersible Alvin. Memoirs of the Australian Museum No. 16: 129-145.

& Fell, H. B. 1965. A revised classification of the dendrochirote holothurians. Breviora No. 214: 1-7.

Perrier, R. 1902. Holothuries. Expeditions Scientifiques du Travailleur et du Talisman 5: 273-554.

Roberts, D. G. 1975. Marine geology of the Rockall Plateau and Trough. Philosophical Transactions of the

Royal Society Series A 278: 447-509.
Roux, M. 1 977. Les Bourguetcrinina du Golfe de Gascogne. Bulletin du Museum National d'Histoire Naturelle

Paris y Serie. Zoologie No. 426. 296: 25-83.
Sibuet, M. 1974. Echinodermes de la mer d'Alboran. Bulletin du Museum National d'Histoire Naturelle Paris

3e Serie. Zoologie No. 231. 155: 789-798.
1977. Repartition et diversite des Echinodermes (Holothurides-Asterides) en zone profunde dans le

Golfe de Gascogne. Deep-Sea Research 24: 549-563.
Sime, A. A. T. & Cranmer, G. J. 1985. Age and growth of North Sea echinoids. Journal of the Marine

Biological Association of the United Kingdom 65: 583-588.

1 98 HARVEY, GAGE, BILLETT, CLARK & PATERSON

Sladen, W. P. 1 889. Asteroidea. Report of the Scientific Results of the voyage of H. M.S. Challenger 1873-1876.

Zoology 30: 1-935.
Suchanek, T. H., Williams, S. L., Ogden, J. C, Hubbard, D. K. & Gill, I. P. 1985. Utilization of shallow- water

seagrass detritus by Caribbean deep-sea macrofauna: 613C evidence. Deep-Sea Research 32: 201-214.
Siissbach, S. & Breckner, A. 1911. Die Seeigel, Seesterne und Schlangensterne der Nord und Ostsee.

Wissenschaftliche Meeresuntersuchungen (Kiel) N.S. 12: 167-300.
Theel, H. 1882. Report on the Holothurioidea dredged by H.M.S. Challenger during the years 1873-1876.

Part 1 . Report of the Scientific Results of the voyage of H.M.S. Challenger 1873-1876. Zoology 4( 1 3): 1-1 76.
1886a. Report on the Holothurioidea dredged by H.M.S. Challenger during the years 1873-1876. Part 2.

Report of the Scientific Results of the voyage of H.M.S. Challenger 1873-1876. Zoology 14(39): 1-290.
18866. Report on the Holothurioidea of the Blake expedition. Bulletin of the Museum of Comparative

Zoology Harvard 13( 1 ): 1-2 1 .
Thomson, W. 1874. On the Echinoidea of the Porcupine deep-sea dredging expedition. Philosophical

Transactions of the Royal Society 164: 719-756.
Tyler, P. A., Billett, D. S. M. & Gage, J. D. (1987). The ecology and reproduction of Cherbonniera utriculus

and Molpadia blakei from the N.E. Atlantic. Journal of the Marine Biological Association of the United

Kingdom 67: 385-398.
& Gage, J. D. 1 9840. The reproductive biology of echinothuriid and cidarid sea urchins from the deep sea

(Rockall Trough, NE Atlantic). Marine Biology 80: 63-74.
& 19846. Seasonal reproduction of Echinus affinis (Echinodermata: Echinoidea) in the Rockall

Trough, NE Atlantic Ocean. Deep-Sea Research 31: 387-402.

— , Gage, J. D. & Billett, D. S. M. 1985a. Life-history biology of Peniagone azorica and P. diaphana

(Echinodermata: Holothurioidea) from the northeast Atlantic Ocean. Marine Biology 89: 71-81.
, Muirhead, A., Billett, D. S. M. & Gage, J. D. 19856. Reproductive biology of the deep-sea holothurians

Laetmogone violacea and Benthogone rosea (Elasipoda: Holothurioidea). Marine Ecology Progress Series

23: 269-277.
-, Pain, S. L., Gage, J. D. & Billett, D. S. M. 1984. The reproductive biology of deep-sea forcipulate sea

stars (Asteroidea: Echinodermata) from the N. E. Atlantic Ocean. Journal of the Marine Biological
Association of the United Kingdom 64: 587-601.

Verrill, A. E. 1 895. Distribution of the echinoderms of Northeastern America. American Journal of Science (3)
49: 127-141 & 199-212.

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 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. 0 565 00982 6 £17.50.

Titles to be published in Volume 54

The crania] 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. Billett, A. M. Clark & G. L. J. Paterson

A morphological atlas of the avian Uropygial gland. By David W. Johnston

Primed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset

INAIUKAL

PRESENTED
GENERAL I

Bulletin of the

British Museum (Natural History)

A morphological atlas of the avian uropygial
gland

David W. Johnston

Zoology series Vol 54 No 5

24 November 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
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
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,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on'either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
England.

World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.)

© Trustees of the British Museum (Natural History), 1 988

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 0565 05041 9

ISSN 0007- 1 498 Zoology series

Vol54 No. 5 pp 199-259
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 24 November 1988

A morphological atlas of the avian uropygial gland

David W. Johnston*

Department of Zoology, University of Florida, Gainesville, Florida 3261 1 USA

Dedicated to PIERCE BRODKORB, whose encouragement , friendship',
and scholarship influenced the preparation of this monograph

Contents

Introduction

Materials and methods

Systematic accounts of gland morphology
Weights and sizes of glands . . . .

Feathers on glands

Naked and tufted glands ....

Gland absence

Glands in flightless birds ....
General taxonomic considerations of glands .
Functions of glandular secretions .

Future studies

Acknowledgments

Summary

Literature Cited

bKITISH M
(NATURAL HI

25NOV

PRESEN
GENERAL L

Introduction

The uropygial gland (also known as the uropygium, preen-, oil-, rump-, tail-, and coccygeal gland;
glandula uropygii; the 'eloeodochon' of Coues 1890) is the only compact gland in the avian -
integument. Lying medially and dorsal to the levator muscles of the tail, this gland is usually
bilobed and partly covered by skin and body feathers. It secretes a chemically complex oil through
ducts in a papilla which often bears a feather tuft (circulus uropygialis of Baumel et al. 1979). Most
reports and investigations on the gland for at least 1 50 years have concentrated on specific topics
such as histology, morphology, function, chemical nature of secretions, and development.

The earliest published references to the uropygial gland focused almost entirely on function.
Emperor Frederick II of Hohenstaufen about 1260 (Wood & Fyfe 1943) in his treatise on falconry
gave the view that birds of prey transfer oil from the gland to their talons via the mandibles, the oil
presumably being noxious and capable of killing the prey more quickly. Willughby (1678) noted
that the 'oyly pap . . . recomposes and places them [feathers] in due order.' The French anatomist
Cuvier (1799-1805) provided the first description of the gland's internal anatomy. Burton (1822)
considered size as well as function of the gland in Fregata aquila. Audubon (1829) discussed the
use of the glandular secretion in lubricating the plumage of 'The Bird of Washington' (the Bald
Eagle, Haliaeetus leucocephalus). From 1832-1836, F.O. Morris and Charles Waterton exchanged
acrimonious and unscientific letters in which they debated the functions ('office') of the gland

"Present address: 5219 Concordia St., Fairfax, Virginia 22032 USA.

Bull. Br. Mm. nat. Hist. (Zool.) 54(5): 199-259

Issued 24 November 1988

200 DAVID W. JOHNSTON

mostly in domestic birds, and they even questioned the ability of birds to preen with oil.
Macgillivray's (1837) interest in the gland was also largely functional, although he reported an
increase in gland size at the time of moulting. Crisp (I860) questioned this report, and found no
experimental evidence for such seasonal size change. Hussey (1860), Waterton (1860), and
Matthews (1861) continued to be concerned with preening and the use of oil in 'barn-door' fowls.

Although these earlier authors were pre-occupied with the gland's functional attributes, some of
their reports contained passing references to gland morphology in various birds (e.g., duck, fowl,
dipper, robin). Nitzsch ( 1 840) published the first account to include morphological information for
glands in many taxa. He discussed for many groups of birds the gland's presence or absence, tufted
vs. naked condition, relative sizes, and shapes. This hallmark publication by Nitzsch was followed
by subsequent authors who either copied Nitzsch or provided information on gland morphology in
additional species: Crisp (1860, 1862), Owen (1866), Kossmann (1871), Garrod (1874a, 6), Coues
(1890), and Newton (1893-1896). Beddard (1898) then published his renowned classification of
birds in which he included some morphological notes on glands in many major taxa. Papers and
books by Pycraft (1900, 1910), Lunghetti (e.g., 1906), Granvik (1913), and Paris (e.g., 1913)
reported much new information about gland morphology in selected species. Thus, information
about gland morphology in different groups and species was scattered among these and other
publications through the first part of the 20th century. More recently, Elder (1954), Lucas &
Stettenheim (1972), and Jacob & Ziswiler (1982) have provided information on gland morphology
in additional taxa. These three publications and Stettenheim (1972) contain a wealth of references
to various biological attributes of uropygial glands (histology, functions, and chemistry).

The degree to which any gland attribute can be used in avian taxonomy remains controversial.
Early classifications of birds (Nitzsch 1840, Beddard 1898, Paris 1913, Verheyen 195 5- 1960) often
employed gland morphology as diagnostic properties of given taxa, but many more modern
schemes (e.g., Hancock &Kushlan 1984) have not. Jacob (e.g., 1978), Jacob & Ziswiler (e.g., 1982),
and von Jacob & Hoerschelmann (1985) provide a chemotaxonomic approach in which they relate
the qualitative chemical composition of gland secretions to the systematic positions of avian taxa.
Their results seem to show qualitative differences at some ordinal, family, and subfamily levels.
Perhaps it is too soon to evaluate this approach, i.e. whether it is any more fundamental in avian
taxonomy than a scheme employing only gland morphology, as a taxonomic characteristic.

Despite a plethora of publications, to date no one has compiled a single, complete, comprehen-
sive monograph on the gland's morphological variations in all major avian taxa. The present
monograph covers, both generally and specifically, the morphology of the gland in all families and
subfamilies, a review of the pertinent morphological literature, and corrections of erroneous and
incomplete information about glands, much of this perpetuated from author to author over the
years without questioning and adequate documentation. This comprehensive survey is deemed
necessary prior to an accurate assessment of the function(s) of the gland's secretion and to the
possible use of gland morphology as a character in avian systematics.

Materials and methods

Most birds examined in this study were preserved in alcohol or other fluids in museum collections.
For a few species, where such specimens were unavailable, I examined museum study skins but
only for rare species that were believed to have tufted glands. Supplementing the museum studies
were ( 1 ) birds caught in mist-nets, or collected with a shotgun in Florida, (2) road kills in Virginia,
England, Ireland, Belize, and Malawi and (3) some freshly-dead birds from zoos (e.g., parrots,
toucans, and hornbills). I dissected a gland from one or more species in each family intact from the
alcoholic specimen, and freed it from connective and other non-glandular tissues. That gland was
then used for the artist's illustrations and studies of feathers. These gland examples currently
remain in my collection.

Anatomical nomenclature for the gland's major parts follows that of Baumel et al. (1979) as
illustrated in Figure 1 .

AVIAN UROPYGIAL GLAND 201

TUFT
PAPILLA

LOBES

Fig. 1 External morphological nomenclature of the uropygial gland.

Body and gland weights were taken, for the most part, from freshly killed birds, but some frozen
birds (e.g., penguins, ducks, hawks) were thawed, weighed, and their glands dissected free for
weighing. Linear dimensions of glands were usually not made. In only a few cases were zoo birds
used for weights because I did not know whether the bird might have been emaciated at the time of
its death. Glands from fresh birds were never compressed before weighing in order to preserve the
gland's oil content at the time of death. The gland tuft, if present, was included in the gland weight,
as was any oil in the tuft.

For the microscopic study of gland feathers, two or three feathers were dissected from a gland,
cleared in xylene, and mounted in Canada balsam on a microscope slide. Each feather preparation
could then be studied under the microscope at magnifications up to SOX. I did not consider gland
histology or development (see reviews in Lucas & Stettenheim 1972 and Jacob & Ziswiler 1982) or
the number of orifices in the papilla's tip (see Jacob & Ziswiler 1982).

This study resulted in the examination of representative glands from every family and all
subfamilies (except one) as identified in Peters (1931-1986). In all, I examined 3011 individuals
from 1433 species and 883 genera. The objectives of this examination were to determine for each
individual the presence or absence of the gland, its shape, relative size, weight, presence or absence
of the papilla and feather tuft. Any individual variations in these characteristics are noted in the
systematic accounts to follow.

Systematic accounts of gland morphology

Classification here follows that of Peters (1931-1986). The terminology for gland morphology (see
Fig. 1) has been adapted from Baumel et al. (1979). At the end of the Morphology section for each
family is the gland feather type (see section on Feathers on glands for definitions). Unless otherwise
indicated, all specimens were considered to be adults. Format for the Material examined: for each
species, the number of individuals examined; in parentheses, when available, sex, body weight in
grams followed by gland weight as a percent of body weight. Z = zoo bird; SK = study skin.
Accompanying gland illustrations contain a linear scale that equals 1 cm.

Order Struthioniformes

General characteristics. Absent in adult Ostrich, Rhea, Cassowary, and Emu, but present (naked)
in Rhea and Emu chicks. Present (naked or with minute feathers) in Apterygidae.

202 DAVID W. JOHNSTON

Family Struthionidae (Ostriches)

MORPHOLOGY. Absent in all age groups. Pycraft (1900) also found no glands in any age group of

this species.

MATERIAL EXAMINED. Struthio camelus 1 1 (9 chicks of various ages, 2 ad.).

Family Rheidae (Rheas)

MORPHOLOGY. Present (naked) and very small in all chicks examined; absent in adult. Pycraft
(1900) found the gland in the 'embryo and nestling' but absent in the adult of Rhea americana.
MATERIAL EXAMINED. Rhea americana 14(11 chicks up to 4 months, 3 ad.). Pterocnemia pennata,
lad.

Family Casuariidae (Cassowaries)

MORPHOLOGY. Absent in all age groups (absent in adult fide Pycraft 1900).
MATERIAL EXAMINED. Casuarius casuarius 4(1 chick, 1 immature, 2 ad.).

Family Dromaiidae (Emus)

MORPHOLOGY. Present (naked) in all chicks examined; probably absent in adults (first reported by

Pycraft 1900).

MATERIAL EXAMINED. Dromaius novaehollandiae 1 3 (chicks up to 2 weeks).

Family Apterygidae (Kiwis)

MORPHOLOGY. More terminally located than in any other family, apparently single-lobed, papilla

conical, naked (or 1-2 minute 'bristle-like feathers' in A. australis mantellifide Beddard 1898,

1899).

NOTE. Beddard ( 1 898, 1 899) was apparently the first to report a gland not only in Apteryx but also

in any ratite bird. The kiwi gland is clearly different from every nonratite gland examined because

of its single-lobed appearance and terminal location. Its presence only in adults distinguishes kiwis

from other ratites. The presence of the gland in kiwis supports a suggested affinity between

Apterygidae and Tinamidae (see Cracraft 1981).

MATERIAL EXAMINED. Apteryx australis 5 (1 chick, 4 ad.); A. owenii 1.

I 1

Rhea americana Dromaius novaehollandiae Apteryx australis

(chick) (chick)

Order Tinamiformes
General characteristics. Tufted with long feathers.

Family Tinamidae (Tinamous)

MORPHOLOGY. Indistinctly bilobed, papilla small or lacking, tufted. Verheyen ( 1 960a) regarded the

gland as always present in the family although 'sometimes vestigial.' In Crypturellus spp. the four

AVIAN UROPYGIAL GLAND

203

feathers, 2 long and 2 short, are 4-5, 3-4 mm in length; in Eudromia spp. the four feathers are 13,
12 mm; shorter (1 -5 mm) in Rhynchotus rufescens (Jacob & Ziswiler 1982). Type II.
MATERIAL EXAMINED. Crypturellus soui 1; C. undulatus 1; C. cinnamomeus 2; C. tataupa 1;
Rhynchotus rufescens 1 ; Nothoproctaperdicaria 1 ; N. pentlandii 1 ; Eudromia elegans 3; E.formosa 1 ;
Tinamotis pentlandii 1.

Order Procellariiformes
General characteristics. Densely tufted in all families.

Family Diomedeidae (Albatrosses)

MORPHOLOGY. Distinctly bilobed, papilla rounded and slightly raised, tufted (32 feathers in

Diomedea exulans, Jacob & Ziswiler 1982). Type I.

MATERIAL EXAMINED. Diomedea exulans 1; D. nigripes 2; D. immutabilis 1; D. melanophrys 1;

Phoebe tria palpebrata 1 .

Family Procellariidae (Fulmars, Petrels, Shearwaters)

MORPHOLOGY. Distinctly bilobed, papilla small or lacking, tufted (24-36 feathers, Paris 1913;

36-42 feathers in 2 species, Jacob & Ziswiler 1982). Type I.

MATERIAL EXAMINED. Macronectes giganteus 1 ; Fulmarus glacialoides 1 ; F. glacialis 2 (F im.: 642-4,

0-67); Thalassoicaantarctical', Daption capense 1; Pagodromanivea 1 (M: 245-0, 0-25); Pterodroma

hasitata 5 (M: 481-3, 0-36; 364.9, 0-34. F: 459-0, 0-35); P. hypoleuca 1; Halobaena caerulea 1;

Pachyptila desolata 1; Bulweria bulwerii 1; Calonectris diomedea 1 (F:410-0, 0-40); Puffinus gravis

7(M: 636-6, 0-44. F: 654-5, 0-39); P. griseus 3; P. puffinus l(unsexed: 450-0, 0-53); P. Iherminieri 2.

Eudromia elegans

Diomedea immutabilis

Puffinus Iherminieri

Family Hydrobatidae (Storm Petrels)

MORPHOLOGY. Indistinctly (Oceanites) or distinctly (Oceanodroma) bilobed, papilla small, tufted

(20 feathers in O. melania). Type I.

204

DAVID W. JOHNSTON

MATERIAL EXAMINED. Oceanites oceanicus 5 (M: 35-6, 0-40; 31-1, 0-46. F: 35-0, 0-40; 33-6, 0-38);
Oceanodroma leucorha 1 (M: 29-5, 0-23); O. melania 1 (unsexed: 53-8, 0-69); O. homochroa 1
(M: 34-0, 0-31); O.furcata \ (unsexed: 54-4; 0-39).

Family Pelecanoididae (Diving Petrels)

MORPHOLOGY. Distinctly bilobed, papilla small, tufted (20 feathers). Type I.

MATERIAL EXAMINED. Pelecanoides magellani \ ; P. georgicus 1 ; P. urinator \ .

Order Sphenisciformes
General characteristics. Densely tufted.

Family Spheniscidae (Penguins)

MORPHOLOGY. Distinctly bilobed, flattened and raised papilla, tufted, ('about 50' feathers, Paris
1913; 44-48 feathers in 2 species, Jacob & Ziswiler 1982). Illustrations in Grasse (1950: 286) and
Jacob & Ziswiler (1982, Fig. 4a, p. 216) of Spheniscus demersus, lacking a feather tuft, are inac-
curate, copied from Paris (1913) who illustrated one gland from which the tuft had undoubtedly
been removed. Type Ha.

MATERIALS EXAMINED. Aptenodytes patagonica 1; A.forsteri 1; Pygoscelis papua 1; P. adeliae 2
(M: 4990, 0-06; 5348, 0-12); P. antarctica 1; Eudyptes crestatus 1; E. chrysolophus 1; Eudyptula
minor 1 ; Spheniscus humboldti 1 .

Oceanodroma melania

I 1

Pelecanoides magellani

Aptenodytes forsteri

Order Gaviiformes
General characteristics. Deeply bilobed, densely tufted.

Family Gaviidae (Loons)

MORPHOLOGY. Distinctly bilobed and elongated, small papilla, tufted (30^40 feathers, Paris 1913;

26-28 feathers in 2 species, Jacob & Ziswiler 1982). Type I.

AVIAN UROPYGIAL GLAND

205

MATERIAL EXAMINED. Gavia stellata 5 (M: 1597, 0-19; 1638, 0-17. F: 1351, 0-21); G. arctica 2
(M: 1598, 0-30. 1 unsexed: 2082, 0-23); G. immer 15 (M: 2780, 0.09; 3180, 0-11; 3490, 0-11);
G. adamsii 1 (unsexed: 41 15, 0-19).

Order Podicipediformes
General characteristics. Deeply bilobed, densely tufted.

Family Podicipedidae (Grebes)

MORPHOLOGY. Distinctly bilobed and somewhat flattened papilla raised nearly perpendicular to

the two lobes, tufted (14-18 subterminal feathers in 2 species, Jacob & Ziswiler 1982). Verheyen

(\959d) reported the gland as 'voluminous and crowned with long plumes.' Type I.

MATERIAL EXAMINED. Rollandia rollandl; R. microptera 1 ; Tachybaptus ruficollis 1 ; T. rufolavatus 1 ;

T. dominions 1; Podilymbus podiceps 8 (M: 358-0, 0-42; 380-2, 0-19. F: 312-0, 0-22; 301-2, 0-26);

P. gigas 1 ; Podiceps major 1 ; P. auritus 3 (M : 410-0, 0-23; 342-7, 0-22); P. grisegena 1 ; P. nigricollis 2;

P. occipitalis 2; P. taczanowskii \\Aechmophorus occidentalis 1 .

Spheniscus humboldti

Gavia immer

Podiceps auritus

Order Pelecaniformes
General characteristics. Densely tufted.

Family Phaethontidae (Tropicbirds)

MORPHOLOGY. Distinctly bilobed, papilla tufted (40 feathers in P. lepturus}. Verheyen (19606)

stated (p. 12) that the gland 'lacks a nipple except in Phaethon.' Type I.

MATERIAL EXAMINED. Phaethon aethereus 1 (F: 496-0, 0-43); P. rubricauda 1; P. lepturus 3.

206

DAVID W. JOHNSTON

Family Fregatidae (Frigatebirds)

MORPHOLOGY. Indistinctly bilobed, papilla broad and flattened, tufted (ca 30 feathers in F.

magnificens). Type I.

MATERIAL EXAMINED. Fregata magnificens 4 (M: 1365-5, 0-07; 1336, 0-07. F: 1512, 0-06); F. aquila

\ ; F. minor 1 ; F. ariel 1 .

Family Phalacrocoracidae

Subfamily Phalacrocoracinae (Cormorants)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, tufted (36-52 feathers in 2

species, Jacob & Ziswiler 1982). Papilla and feather tuft subterminal. Type I.

MATERIAL EXAMINED. Phalacrocoraxharrisi\;P. auritus24(M: 1660,0-25; 1710, 0-25. xofSF: 1048,

0-25); P. aristotelis 1; P. magellanicus 1; P. bougainvillii 1; P. albiventer 1.

Phaethon lepturus

Fregata ariel

Phalacrocorax auritus

Subfamily Anhinginae (Darters)

MORPHOLOGY. Bilobed and somewhat flattened, papilla absent, short tuft of 14 feathers. Type I.
MATERIAL EXAMINED. Anhinga anhinga 11 (M: 1230, 0-16; 1352, 0-12; 1230, 0-16. F: 1307, 0-15;
1 1 78, 0- 1 7); A . melanogaster 2.

Family Sulidae (Boobies, Gannets)

MORPHOLOGY. Distinctly flattened and bilobed, papilla absent, tufted with 70 short feathers in

S. bassana (Jacob & Ziswiler 1982). Type I.

MATERIAL EXAMINED. Sula bassana 3 (M subad.: 2200, 0-38; S. dactylatra 2; 5. sula 1 .

Family Pelecanidae (Pelicans)

MORPHOLOGY. Large and bilobed, papilla very short and broad, tufted (70 feathers, Paris 1913;

66 feathers in P. onocrotalus, Jacob & Ziswiler 1982). Tuft and openings obviously subterminal,

Type I.

MATERIAL EXAMINED. Pelecanus onocrotalus 1; P. philippensis 1; P. erythrorhynchos 2 (M: 4700,

0-30; P. occidentalis 9 (M im.: 2730, 0-43; 3060, 0-32; unsexed ad.: 3320, 0-37).

AVIAN UROPYGIAL GLAND

207

Anhinga anhinga

Sula bassana

Pelecanus onocrotalus

Order Ciconiiformes

General characteristics. Much inter- and some intrafamilial variation: naked, sparsely or densely

tufted.

NOTE. The large morphological differences among glands examined here lend support to Olson's

(1979) view that the Ciconiiformes is not a natural order.

Family Ardeidae (Herons, Bitterns)

Subfamily Ardeinae (Day Herons)

MORPHOLOGY. Although considerable variation occurs in the family, the gland is generally small
(see also Paris 1913: 192), bilobed, lacks a papilla (or 'very short,' Paris 1913: 192), and is tufted
(4-18 feathers) or naked. Although 'small' in many species, the gland cannot be regarded as
'rudimentary' as described by Jacob (1978: 168). Verheyen (19596) noted that in the Ardeae, the
gland is naked or has a few vestigial feathers. Miller (1924) described variations in the tuft among
13 species, then reported the tuft as absent in Ardea goliath, A. herodias, A. cocoi, A. occidentalis,
Notophoyx novaehollandiae , N. pacifica, Egretta candidissima, and Hydranassa tricolor. I found
the tuft absent only in Pilherodias pileatus, Ardea pacifica, Egretta rufescens, E. tricolor,
E. novaehollandiae, E. garzetta and E. sacra. In A. herodias (contra Miller 1924), all 10 specimens
examined here had extremely small feathers, often as few as 4. Beddard (1898) and Gadow (1893)
reported that all Ardeidae have feathered glands. Thus in the family, the tuft might be absent,
represented by only 4-8 feathers (Ardea herodias, Agamia agami), or tufted with 16-18 feathers
(Ixobrychus minutusfide Jacob & Ziswiler 1982). Type I.

MATERIAL EXAMINED. Pilherodias pileatus 1*; Ardea cinerea 1; A. herodias 10 (M: 2695-0, 0-21;
1 28 1 -0, 0-05; 1 790, 0- 1 4. F: 1 250-0, 0. 1 7; 1 507, 0- 1 0); A . pacifica \*;A. melanocephala \\A. purpurea
1; A. alba 8 (M: 478-5, 0-15; 579-0, 0-12); Egretta rufescens 1*; E. tricolor *; E. ibis 1 1 (M: 342-6,
0-03. x of 5F: 286-8, 0-04); E. novaehollandiae 1*; E. caerulea 3 (F: 372-0, 0-07); E. garzetta 1*; E.
sacra 1 *; Ardeola speciosa \;A. striata 1 (M: 220-3, 0-07; 220- 1 , 0-07; 1 95-4, 0-06); Agamia agami 2.
NOTE. Ligon (1967: 1) believed 'that the storks and herons are dissimilar' in many features
(osteology, myology, pterylography and others); the dissimilar uropygial glands of the two groups
add another difference.

*naked gland, present study.

208

DAVID W. JOHNSTON

Subfamily Nycticoracinae (Night Herons)

MORPHOLOGY. Indistinctly bilobed, no papilla, tufted (Nycticorax, Nyctanassa) or naked

(Cochlearis).

MATERIAL EXAMINED. Nyctanassa violacea 5 (M: 546-0, 0-04); Nycticorax nycticorax 2; Cochlearius

cochlearius 2.

NOTE. Peters regarded Cochlearis as comprising a separate family, the Cochlearidae (Peters 1931,

Vol. I, 1st ed.). Mayr & Cottrell, (1974 in their 2nd ed. of Peters' Vol. I) and Hancock & Kushlan

(1984), however, included Cochlearis in a subfamily (Nycticoracinae) of the family Ardeidae. The

gland of Cochlearis differs markedly from that of all other ardeids because of its relatively large

size, distinctive appearance (see figure), and absence of papilla and feather tuft. Beddard (1898)

and Miller (1924) both noted this distinctiveness of the gland in Cochlearis.

Subfamily Tigrisomatinae (Tiger Herons)
MORPHOLOGY. Bilobed, short papilla, tufted. Type I.
MATERIAL EXAMINED. Tigrisoma mexicanum 1 .

Subfamily Botaurinae (Bitterns)

MORPHOLOGY. Indistinctly bilobed, no papilla, small tuft. Type I.

MATERIAL EXAMINED. Ixobrychus exilis 2 (M: 68-0, 0-05. F: 38-6, 0-03); Botaurus lentiginosus 5

(M: 564-0, 0-31; 789-0, 0-42; 720-0, 0-38. F: 599-3, 0-38).

Family Scopidae (Hammerhead)

MORPHOLOGY. Indistinctly bilobed, papilla short and somewhat flattened, and tufted (18 long

feathers). The gland of this species is distinct from those of ardeids, more closely resembling that of

Balaeniceps and the Ciconiidae (see figures). Type I.

MATERIAL EXAMINED. Scopus umbretta 2.

Family Ciconiidae (Storks)

MORPHOLOGY. Distinctly bilobed and large, papilla small, tufted (36 feathers in C. ciconia, Jacob

& Ziswiler 1982). Unlike the illustration of Mycteria here, in Anastomas, Ciconia nigra, and

Ardea herodias

Cochlearius cochlearius

Scopus umbretta

AVIAN UROPYGIAL GLAND

209

Leptoptilos the gland appears to be separated into right and left portions, each with a separate lobe,
feather tufts and orifices, features implied by Nitzsch (1867: 131). Type I.

MATERIAL EXAMINED. Mycteria americana 3 (F: 2490-0, 0-05); Anastomus oscitans 1; Ciconia nigra
1 ; C. abdimii 1 ; C. episcopus 1 ; C. ciconia 1 ; Ephippiorhynchus asiaticus 1 ; Leptoptilos crumeniferus 1 .

Family Balaenicipitidae (Shoebill)

MORPHOLOGY. Indistinctly bilobed and relatively small (see figure and Bartlett 1861), papilla small,

tufted. With the exception of the large tuft in Scopus, I agree with Miller's (1924: 322) comment

that 'in Balaeniceps the tuft is very much larger than in any heron,' although I did not count the

feathers. Type II.

MATERIAL EXAMINED. Balaeniceps rex 1Z.

NOTE. Gland morphology sheds no light on the controversy over the affinities of Balaeniceps

(Cottam 1957, Olson 1979, Cracraft 1981).

Family Threskiornithidae (Ibises, Spoonbills)

Subfamily Threskiornithinae (Ibises)

MORPHOLOGY (family). Distinctly bilobed, papilla absent, tufted (26-28 feathers, Paris 1913; 30
feathers in E. ruber, Jacob & Ziswiler 1 982). Nitzsch ( 1 867) implied that ibises have glands 'divided
in half.' In the present study one specimen each of Eudocimus albus and Platalea leucorodia had
glands with nearly separate lobes, separate orifices, and separate feather tufts. Type I.
MATERIAL EXAMINED. Eudocimus albus 2 (M, Z: 999, 0-12); E. ruber 2; Plegadis falcinellus 3
(M: 530-0, 0-18; F: 420.0, 0-19); Threskiornis aethiopicus 2.

Subfamily Plataleinae (Spoonbills)

MATERIAL EXAMINED. Platalea leucorodia 1; P. ajaja \ (M: 985-0, 0-11).

Mycteria americana

Balaeniceps rex

Threskiornis aethiopicus

2 1 0 DAVID W. JOHNSTON

Order Phoenicopteriformes
General characteristics. Densely tufted.

Family Phoenicopteridae (Flamingos)

MORPHOLOGY. Bilobed, papilla absent, tufted (30 feathers, Paris 1913). Type Ha.

NOTE. Paris (1913: 190) believed that the gland of Phoenicopterus roseus closely resembles that

of the Anseriformes. In contrast, I found marked differences in the gland of Phoenicoparrus

from both the Anseriformes and Ciconiiformes (see figures), relationships suggested by Sibley

(1967), and Sibley et al. (1969). The gland resembles that of the Recurvirostridae, suggesting a

charadriiform relationship as proposed by Olson & Feduccia (19806).

MATERIAL EXAMINED. Phoenicopterus ruber 1; Phoeniconaias minor 2; Phoenicoparrus andinus 1;

P.jamesi 1.

Order Falconiformes

General characteristics. Inter- and some intrafamilial variation: naked or sparsely to densely
tufted.

Family Cathartidae (American Vultures)

MORPHOLOGY. Indistinctly bilobed, round papilla, naked; two separate, distinct orifices. Nitzsch's

( 1 867) report that 'vultures of the New World' have a short circlet of feathers at the gland apex

appears to be in error. I did not confirm Fisher's (1943) statement that down is often present on the

oil gland in Coragyps atratus.

NOTE. My findings support Ligon's (1967: 1) view 'that the Cathartidae are not at all closely related

to the remainder of the Falconiformes.' The naked glands of the Cathartidae, however, differ

markedly from the heavily tufted glands of the Ciconiidae, to which cathartids might otherwise be

related (Ligon 1967).

MATERIAL EXAMINED. Coragyps atratus 5 (M: 2221-0, 0-04; 2245-0, 0-04. F: 2135-0, 0-05; 2238-0,

0-04); Cathartes aura 7 (M: 1939-0, 0-02. x of 5F: 2019-0, 0-02); Gymnogyps calif ornianus 2 (1 SK);

Vultur gryphus 2; Sarcoramphus papa 2.

Phoenicoparrus jamesi

Cathartes aura

Pandion haliaetus

AVIAN UROPYGIAL GLAND

211

Family Accipitridae

Subfamily Pandioninae (Osprey)

MORPHOLOGY. Large and distinctly bilobed, indistinct papilla, tufted (18 feathers, Jacob &

Ziswiler 1982). Type I.

NOTE. Sometimes included in a separate family, Pandionidae (Cracraft 1981), the Osprey's gland

differs from those of the Accipitrinae by being much heavier and having a longer, usually denser

feather tuft.

MATERIAL EXAMINED. Pandion haliaetus 5 (M: 1433-0, 0-28; 1363-0, 0-31. F: 1500-0, 0-32).

Subfamily Accipitrinae (Hawks, Eagles)

MORPHOLOGY. Bilobed, papilla moderately developed, sparsely (Ictinid) to densely (Haliaeetus)
tufted (12-20 feathers in 7 species, Jacob & Ziswiler 1982). Type 1 .

MATERIAL EXAMINED. Aviceda leuphotes 1; Elanoides forficatus 2 (M: 492-0, 0-13); Harpagus
bidentatus 1; Ictinia plumbea 1; /. mississippiensis 1; Haliaeetus leucocephalus 3 (ad. sex?: 3625,
0-07); Neophron percnopterus 2; Gyps fulvus 1; Circus hudsonius 2 (F: 324-6, 0-03); Melierax
canorus 1; Accipiter nisus 1; A. striatus 10 (subad. M: 84-2, 0-05; 98-0, 0-04; 88-9, 0-06. x of 4 ad. F:
157-3, 0-06); A. cooperii 4 (ad. F: 41 1 -7, 0-07; 390-0, 0-07)M. gentilis 4 (subad. M: 816-0, 0-06; 775-0,
0-05; 918-0, 0-07. Ad. F: 930-0, 0-03); Geranospiza caerulescens 1; Buteo lineatus 9 (ad. M: 612-3,
0-05; 595-7, 0-06. Ad. F: 601-0, 0-05; 566-7, 0-07); B. platypterus 2 (subad. M: 489-4, 0-04; 309-0,
0-05); B. swainsoni 1 (ad. M: 874-0, 0-04); B.jamaicensis 14 (ad. M: 1307-3, 0-04. Subad. M: 856-3,
0-06. Subad. F: 1210-0, 0-04; 1272-0, 0-08); B. lagopus 3 (ad. M: 860-0, 0-04); Pithecophagajefferyil;
Aquila chrysaetos 1.

Family Sagittariidae (Secretarybird)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, tufted (20 feathers). In most

specimens the gland appears to be nearly separated into two distinct lobes, with separate papillae

and feather tufts. Although writing about 'Les Cariamiformes,' Verheyen (1957c) stated

erroneously that the gland is absent in Sagittarius. Type I.

MATERIAL EXAMINED. Sagittarius serpentarius 6.

Buteo jamaicemis

Sagittarius serpentarius

Falco columbarius

2 1 2 DAVID W. JOHNSTON

Family Falconidae

Subfamily Polyborinae (Caracaras)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, densely tufted.

MATERIAL EXAMINED. Daptrius ater 1; Polyborus plancus 2 (unsexed im.: 900-5, 0-07).

Subfamily Falconinae (Falcons)

MORPHOLOGY. Distinctly bilobed, papilla moderately to well developed, tufted (17 feathers in
Falco tinnunculus, Jacob & Ziswiler 1982), or rarely, naked. Both Miller (1924) and Verheyen
(1959c) reported naked glands in Microhierax fringillarius and Nitzsch (1867) noted naked glands
in two specimens of M. caerulescens. In my study all glands of 1 1 individuals of four species of
Microhierax were naked, but all other genera and species in this subfamily had tufted glands. Type
I.

MATERIAL EXAMINED. Micros tur ruficollis 1 : Spiziapteryx circumcinctus 1 ; Polihierax semitorquatus
1 ; P. insignis 4; Microhierax (mostly SK) caerulescens 7; M. fringillarius 2; M. erythrogenys 1 ;
M. melanoleucus 1; Falco sparverius 16 (x of 6 M: 100-3, 0-05. x of 6 F: 1 19-2, 0-08); F. tinnunculus
1; F. columbarius 2 (F: 191.0, 0-09); F. mexicanus 1; F. rufigularis 1; F. rusticolus 1; F. peregrinus 3
(M: 61 7-0, 0-08).

Order Anseriformes

Gland characteristics. Densely tufted.

MORPHOLOGY (order). Large and distinctly bilobed, papilla moderately developed, tufted (22-90
feathers in 15 species, Jacob & Ziswiler 1982). The Anhimidae differ from the Anatidae only by
having a gland that is less distinctly bilobed. Type I.

Family Anatidae

Subfamily Anseranatinae (Pied Geese)
MATERIAL EXAMINED. Anseranas semipalmata, 1 .

Subfamily Dendrocygninae (Whistling Ducks)
MATERIAL EXAMINED. Dendrocygna bicolor 1 .

Subfamily Anserinae (Swans, Geese)

MATERIAL EXAMINED. Cygnusolor 1 (im. M: 10435,0-16); C. columbianus 1; Coscoroba coscoroba 1;

Anser albifrons 1 (F: 2556, 0-11);^. caerulescens 2 (M: 2330, 0-13; 2154, 0-12); A. rossii 1 (M: 1616,

0-15); A. canagicus \ (M: 1855, 0-10); Branta canadensis 2 (M: 2435, 0-1 1. F: 3929, 0-15); Cereopsis

novaehollandiae 1 .

Subfamily Tadorninae (Shelducks)

M ATERIAL EXAMINED. Alopochen aegyptica 1; Tachyeres pteneres 1; T.patachonicus 1.

Subfamily Anatinae (Typical Ducks)

MATERIAL EXAMINED. Rhodonessa caryophyllacea 1; Aix sponsa 3 (M: 642-0, 0-26; 573-1, 0-32. F:
595-2, 0-32); A. galericulata \\Anasstrepera 1 (M: 849-5, 0-20); A. crecca 3 (M: 251-0, 0-31; 295-5,
0-27. F: 262-0, 0-29); A. aucklandica 1; A. platyrhynchos 2 (M: 880-0, 0-23); A. acuta 1 (F: 708-5,
0-26); A. discors 8 (M: 341-5, 0-40; 368-2, 0-33. F: 365-5, 0-36; 384-0, 0-35; 430-0, 0-32); A. clypeata
2 (M: 672-0, 0-27); Aythya valisineria 3 (M: 893-3, 0-27. F: 792-0, 0-30; 884-2, 0-24); A. americana
2 (F: 1 172-0, 0-28); A. collaris 1 1 (x of 8 M: 749-6, 0-32. F: 697-5, 0-31; 745-0, 0-30); A. marila 2
(F: 843-0, 0-22; 991-6, 0-20); A. affinis 2 (M: 882-8, 0.22. F: 783-4, 0-22).

Subfamily Merginae (Sea Ducks)

MATERIAL EXAMINED. Somateria mollissima 1 (M: 2255, 0-18); S. spectabilis 1 (M: 1540, 0-26);
Melanitta perspicillata 1 (F: 703-1, 0-20); Bucephala clangula 1; B. albeola 1; Mergus cucullatus
5 (M: 571-0, 0-34; 671-0, 0-37. F: 671-3, 0-30; 526-0, 0-28); M. senator 1 (F: 599-5, 0-36);
M. merganser 2 (M: 1577, 0-21. F: 1027, 0-29); M. australis 1.

AVIAN UROPYGIAL GLAND 2 1 3

Subfamily Oxyurinae (Stifftailed Ducks)

MATERIAL EXAMINED. Oxyurajamaicensis 1 (M: 596-7, 0-27).

Family Anhimidae (Screamers)

MATERIAL EXAMINED. Anhima cornuta 1 (M, Z: 2600, 0-13); Chauna chavaria 1.

Order Galliformes

Gland characteristics. Inter- and intrafamilial variation: sparsely to densely tufted, or, rarely,
naked.

NOTE. Glands of most galliform families do not resemble those of the Anseriformes. These differ-
ences do not support an anseriform-galliform relationship (see also Olson & Feduccia 1 980«). I did
not confirm Pettingill's (1985) statement that 'certain species' of Galliformes lack a gland.

Family Megapodiidae (Megapodes)

MORPHOLOGY. Bilobed, papilla large, naked or tufted (6 feathers). The present study and the

reports of Miller (1924) and Clark (1964) demonstrate naked glands in Leipoa, Alectura, Tallegalla

jobiensis, and Aepypodius arfakianus, whereas tufted glands are known from five species of

Megapodius and Macrocephalon maleo. Beddard (1898: 302) reported that megapodes have 'oil

gland nude,' and both Sharpe and Ogilvie-Grant regarded the glands of these birds as nude (fide

Miller 1924). Type I.

MATERIAL EXAMINED. Megapodius nicobariensis 2; M.freycinet 3; M. pritchardii 6; Alectura lathami

1 ; Aepypodius arfakianus 1 .

Anas discors

Anhima cornuta

Megapodius pritchardii

Family Cracidae (Curassows, Guans, Chachalacas)

MORPHOLOGY. Distinctly bilobed, papilla large, tuft usually short and sparsely feathered (6-12

feathers, 2-4 mm). Miller (1924: 322) reported an 'apparently bare' gland in one specimen of

Ortalis vetula, and noted a 'virtually vestigial' tuft in all the Cracidae, this last conclusive statement

confirmed in the present study. Type I.

MATERIAL EXAMINED. Crax nigra 1 ; C. alberti 1 ; C. globulosa 1 ; Penelope jacu-caca 1 ; Ortalis guttata

2; O. vetula 1; Pipile pipile 1.

2 1 4 DAVID W. JOHNSTON

Family Tetraonidae (Grouse, Ptarmigans)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted (10-12 feathers, Jacob & Ziswiler 1982).

Type I.

MATERIAL EXAMINED. Tetrao urogallus 2; Lyrurus tetrix 1; Lagopus lagopus 1 (M: 576-0, 0-18);

Canachites canadensis 1 (M: 552-4, 0-02); Bonasa umbellus 3 (M: 552-2, 0-03. F: 591-5, 0-03);

Pedioecetes phasianellus 1 (F: 664-0, 0-03); Tympanuchus cupido 1 (M: 863-5, 0-02); Centrocercus

urophasianus 1 (M: 2221-7, 0-04).

Family Phasianidae (Quails, Pheasants, Peacocks)

MORPHOLOGY (family). Usually distinctly bilobed, papilla large, tuft variable (6-12 feathers, Paris
1913; 5-10 feathers in 6 species, Jacob & Ziswiler 1982; only 2 in Rollulus) or, rarely, absent (one
specimen of Crossoptilon mantchuricum was naked as were five specimens of Argusianus). Earlier,
Nitzsch (1840), Newton (1893-1896), Beddard (1898), Grasse (1950), and Verheyen (\956d) had
reported the absence of a gland in Argusianus ( = Argus). Beddard (1898) found a tuft in one
specimen of Callipepla squamata but a naked gland in another specimen. Type I.

Subfamily Odontophorinae (American quail)

MATERIAL EXAMINED. Callipepla squamata 2 (F: 209-4, 0-06); Lophortyx californica 3 (M: 133-2,
0-09. F: 169-4, 0-08); Colinus virginianus 9 (x of 4 M: 151-8, 0-15. x of 4 F: 156-5, 0-16); Cyrtonyx
montezumae 1 .

Subfamily Phasianinae (Partridges, Quails, Pheasants)

MATERIAL EXAMINED. Francolinus adspersus 1; F. ahantensis 1; Perdix per dix 1 (F: 360-0, 0-09);
Coturnix coturnix 1; Excalfactoria chinensis 3; Arbor ophila torqueola 1; A. brunneopectus 1;
Rollulus roulroul 1; Tragopan temmincki 1; Lophophorus impejanus 1; Crossoptilon auritum 1;
C. mantchuricum 1 ; Lobiophasis bulweri 1; Callus gallus 5 (M: 2270, 0-02); Callus gallus x Meleagris
gallopavo 1; Catreus wallichii 1 (M, Z: 1340, 0-05); Phasianus colchicus 2 (M: 1374.7, 0-03; 1292,
0-02); Syrmaticus reevesii 1 ; Chrysolophus pictus 1 ; Argusianus argus 5; Pavo cristatus 3 (M, Z: 3350,
0-02); Afropavus congensis 3.

Crax alberti Lyrurus tetrix Phasianus colchicus

Family Numididae (Guinea fowl)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted (8 feathers in N. meleagris, Jacob & Ziswiler

1 982; only 2 in Phasidus). Type I.

MATERIAL EXAMINED. Phasidus niger 1 ; Numida sp. 1 ; N. meleagris 1 ; Guttera pucherani 1 ; Acryllium

vulturinum 1.

AVIAN UROPYGIAL GLAND

215

Family Meleagrididae (Turkeys)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted (7, 9 feathers, Jacob & Ziswiler 1982). Type

I.

MATERIAL EXAMINED. Meleagris gallopavo 5 (ad. M: 7400, 0.02. Subad. M: 3740, 0-02. Subad. F:

1870,0-04).

Family Opisthocomidae (Hoatzin)

MORPHOLOGY. Bilobed, papilla small, tuft (up to 12 feathers) variable in size, or gland naked.

Beddard (1898), Gadow (1893), and Verheyen (1956c) described the gland as feathered. I found

that 9 nestlings, 'young,' or 'juveniles' had minute tufts, 2 'subadults' were naked, and 3 'adults'

had tufted glands. Type I.

MATERIAL EXAMINED. Opisthocomus hoazin 14.

Numida sp.

Meleagris gallopavo

Opisthocomus hoazin

Order Gruiformes

Gland characteristics. Inter- and intrafamilial variation: gland absent, naked, or sparsely to densely
tufted.

Family Mesoenatidae (Mesites, Monias)

MORPHOLOGY. Absent (also reported as such by Gadow 1893, Miller 1924, Verheyen 1958a, Van
Tyne & Berger 1976). Beddard (1898: 381) erroneously reported the gland as present and nude.
MATERIAL EXAMINED. Mesoenas variegata 1; M. unicolor 1; Monias benschil.

Family Turnicidae (Bustard-Quails)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, tufted (also reported by

Verheyen 1958«; ca 10 feathers). Type I.

MATERIAL EXAMINED. Turnix sylvatica 2; T. tanki 1; T.suscitator 1; Ortyxelos meiffrenii 1.

Family Pedionomidae (Collared Hemipodes)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, tufted (12 feathers). Type I.

NOTE. Gadow ( 1 89 1 ) and Beddard ( 1 898) reported a tufted gland in this family. Olson & Steadman

(1981) believe that Pedionomus is a charadriiform, but the tufted gland supports no specific

relationship for this family.

MATERIAL EXAMINED. Pedionomus torquatus 2.

Family Gruidae (Cranes)

MORPHOLOGY (family). Large and bilobed, papilla small with the end large and tufted (14-16

216

DAVID W. JOHNSTON

feathers, Paris 1913; 32 feathers in G. grus, Jacob & Ziswiler 1982; 20 feathers in Grus, Nitzsch
1867). Gadow (1893) described the gland of Grus grus as naked, but that report was probably
erroneous because of the tufted glands now known from all other species examined. Type I.

Subfamily Gruinae

MATERIAL EXAMINED. Grus canadensis 9 (ad. M: 3520, 0-05. Subad. M : 2360, 0-03. Ad. F: 3880, 0-05;
3560, 0-05); G. antigone 1; Anthropoides paradisea 2.

Turnix suscitator

Pedionomus torquatus

Grus canadensis

Subfamily Balearicinae

MATERIAL EXAMINED. Balearica pavonina 2.

Family Aramidae (Limpkin)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, tufted (14 feathers). Type I.

MATERIAL EXAMINED. Aramus scolopaceus 9 (x of 6 M: 1 128-6, 0-28. F: 1300, 0-29).

Family Psophiidae (Trumpeters)

MORPHOLOGY. Indistinctly bilobed, papilla absent, sparse and short tuft (20 feathers). Nitzsch

(1867) made the contradictory statement (p. 123), '. . . of the naked oil-gland, which is furnished

with a circlet of feathers at the tip.' Type I.

MATERIAL EXAMINED. Psophia crepitans 1; P. leucoptera 3.

Family Rallidae (Rails, Coots, Gallinules)

MORPHOLOGY (family). Distinctly bilobed, papilla usually large, tuft variable (Verheyen 19576)
(1 1-17 feathers in 5 species, Jacob & Ziswiler 1982; 6 in Porphyrio) or gland naked. Miller (1924),
Verheyen (1957Z>), and Ripley (1976) stated that Himantornis has a naked gland, a condition that I
confirmed. Beddard (1898: 321) stated that Ralli 'have as a rule a tufted oil gland but Porzana
Carolina is an exception.' Miller ( 1 924) and I each found that 5 different specimens of this species all
had tufted glands. In Atlantisia rogersi 2 specimens at the British Museum had naked glands, but
2 specimens at the American Museum of Natural History and Museum of Comparative Zoology
each had tufted glands. Type I.

Subfamily Rallinae

MATERIAL EXAMINED. Rallus longirostris 3 (F: 1 86-3, 0-23); R. elegans 3 (F: 298-0, 0-16; 372-7, 0-29);
R. limicola 6 (M: 66-3, 0-34. F: 65-3, 0-09), R. owstoni 1; R. wakensis 1; Atlantisia rogersi
5; Tricholimnas sylvestris 1; Dryolimnas cuvieri 1; Rallina eurizonoides 1; Cyanolimnas cerverai
1; Gallirallus australis 2; Himantornis haematopus 1; Crecopsis egregia 1; Crex crex 1; Anurolimnas

AVIAN UROPYGIAL GLAND

217

castaneiceps 2; Limmocorax flavirostra 2; Porzana Carolina 5 (M: 66-9, 0-15. F: 65-9, 0-22); P.
albicollis 1; Porzanula palmeri 2; Later allus albigular is 1; Micropygia schomburgkii 1; Coturnicops
noveboracensis 1; Sarothrura rufa 1; Poliolimnas cinerus 1; Tribonyx mortierii 1; Amaurornis
phoenicurus 1 ; Gallicrex cinerea 1 ; Gallinula chloropus 8 (M: 364-2, 0- 1 1 ; 21 1 -5, 0-23. x of 4 F: 274-3,
0-18; Porphyriornis nesiotis 1; P. comeri 1; Porphyrula alleni 1; P. martinica 4 (F: 203-6, 0-16);
Porphyrio porphyrio 1, P . poliocephalus 1; Notornis mantelli 1.

Aramus scolopaceus

Psophia leucoptera

Porphyrio porphyrio

Subfamily Fulicinae

MATERIAL EXAMINED. Fw//ca americana 8 (M: 625-0, 0-1 1. F: 395-4, 0-16; 386-5, 0-17).

Family Heliornithidae (Sun-Grebes)

MORPHOLOGY. Broad and bilobed, papilla moderately developed, tufted (16 feathers). Type I.

MATERIAL EXAMINED. Podica senegalensis 1; Heliopais per sonata 1; Heliornisfulica 1.

Family Rhynochetidae (Kagu)

MORPHOLOGY. Apparently single-lobed, indistinct papilla, naked. The gland is not 'rudimentary'

as stated by Jacob (1978: 168).

MATERIAL EXAMINED. Rhynochetos jubatus 1 .

Family Eurypygidae (Sun-Bittern)

MORPHOLOGY. Indistinctly bilobed, papilla moderately developed, tufted (10 1-2 mm feathers).

Miller (1924: 322) concluded: 'Gadow gives the oil-gland ofEurypyga as bare; Beddard states that

it is generally nude but occasionally tufted. In each of my two fresh examples, . . . there was a small

tuft present.' Type I.

MATERIAL EXAMINED. Eurypyga helias 3.

Family Cariamidae (Cariamas)

MORPHOLOGY. Apparently single-lobed, papilla large, naked. The unusual shape is described by

Nitzsch (1867) as distinctly 'of a conical pyriform shape.'

MATERIAL EXAMINED. Cariama cristata 1 .

Family Otidae (Bustards)

MORPHOLOGY. Gland absent in all species examined, a condition previously noted by Nitzsch
(1840), Gadow (1893), Beddard (1898), Paris (1913), Grasse (1950), Verheyen (19576) and Van
Tyne & Berger (1976). Paris (1913) reported 'well marked outlines of the gland' in embryos.

218

DAVID W. JOHNSTON

Heliopais per sonata

Rhinochetosjubatus

Eurypyga helias

MATERIAL EXAMINED. Choriotis kori 2; C. australis 1 ; Lophotis ruficrista 1 ; Eupodotis senegalensis 2;
Lissotis melanogaster 1 .

Order Charadriiformes

Gland characteristics. Heavily tufted.

NOTE. 1 0 families were described by Verheyen ( 1 9586) as being tufted. All individuals of 1 6 families

in the present study had tufted glands. Type I.

Family Jacanidae (Jacanas)

MORPHOLOGY. Bilobed, papilla moderately developed (contra 'without a well-developed nipple'

Verheyen \951d), tufted (12 feathers).

MATERIAL EXAMINED. Jacana spinosa 2.

Family Rostratulidae (Painted Snipe)

MORPHOLOGY. Bilobed, papilla indistinct, tufted (12 feathers).

MATERIAL EXAMINED. Rostratula benghalensis 1 .

Cariama cristata

Jacana spinosa

Rostratula benghalensis

AVIAN UROPYGIAL GLAND

219

Family Haematopodidae (Oyster-catchers)

MORPHOLOGY. Bilobed, papilla small, tufted (28 feathers, Jacob & Ziswiler 1982).

MATERIAL EXAMINED. Haematopus ostralegus 2.

Family Charadriidae (Lapwings, Plovers)

MORPHOLOGY (family). Distinctly bilobed, papilla moderately developed, tufted (12-24 feathers,

Paris 1913; 1 2-1 4 feathers, Jacob & Ziswiler 1982).

Subfamily Vanellinae

MATERIAL EXAMINED. Vanellus vanellus 2; Hoplopterus spinosus 2; Hoploxypterus cay anus 1;
Zonifer tricolor 1 .

Subfamily Charadriinae

MATERIAL EXAMINED. Squatarolasquatarola2(M: 205-3,0-22. F: 216-2,0-14); Charadrius hiaticula
\ ; C. vociferus 4 (F: 92-0, 0-08; 94- 1 , 0-07); Eupoda montana 1 .

Family Scolopacidae (Woodcock, Sandpipers)

MORPHOLOGY (family). Distinctly bilobed, papilla small, tufted (12-24 feathers in 4 species, Jacob

& Ziswiler 1982).

Subfamily Tringinae

MATERIAL EXAMINED. Bartramia longicauda 1; Numenius minutus 1; Tringa totanus 1; Actitis
macularia 2 (F: 28-5, 0-12); Catoptrophorus semipalmatus 5 (M:297-8, 0-19).

Haematopus ostralegus

Charadrius vociferus

Catoptrophorus semipalmatus

Subfamily Arenariinae

MATERIAL EXAMINED. Arenaria interpres 5.

Subfamily Scolopacinae

MATERIAL EXAMINED. Limnodromus scolopaceus 8 (x of 4M: 104-2, 0- 1 8. F: 121 • 1, 0-20); L. griseus 6
(F: m-9,Q-\2;S5-2,Q-\2y,Capellagallinago3(F: 101-5, 0-09); Philohela minor 5 (M: 106-0, 0-09.X
of4F: 179-2,0-09).

Subfamily Eroliinae

MATERIAL EXAMINED. Calidris canutus 2 (M: 94-5, 0-08); Crocethia alba 2 (F; 49-9, 0-07); Ereunetes
pusillus 2; Erolia minutilla 1; E.fuscicollis 1; E. alpina 2.

220

DAVID W. JOHNSTON

Family Recurvirostridae (Avocets, Stilts)

MORPHOLOGY. Bilobed, papilla small, tufted (ca 20 feathers).

MATERIAL EXAMINED. Himantopus himantopus 3; Recurvirostra americana 1 (F: 289-4, 0-20).

Family Phalaropodidae (Phalaropes)

MORPHOLOGY. Distinctly bilobed, papilla small, tufted.

MATERIAL EXAMINED. Phalaropusfulicarius 4 (unsexed: 40-5, 0-62; 40-4, 0-53. F: 42-0, 0-79); Lobipes

Family Dromadidae (Crab-plovers)

MORPHOLOGY. Distinctly bilobed, papilla apparently absent, tufted (16 feathers).

MATERIAL EXAMINED. Dromas ardeola 1 .

Himantopus himantopus

Phalaropusfulicarius

Dromas ardeola

Family Burhinidae (Thick-knees)

MORPHOLOGY. Indistinctly bilobed, broad papilla, tufted (14 feathers in B. oedicnemus, Jacob &

Ziswiler 1982).

MATERIAL EXAMINED. Burhinus oedicnemus 1; B. senegalensis 1; Esacus recurvirostris 1.

Family Glareolidae (Pratincoles, Coursers)

MORPHOLOGY (family). Indistinctly bilobed, large and round papilla, tufted (14 feathers).

Subfamily Cursoriinae

MATERIAL EXAMINED. Cursorius cursor 1 .

Subfamily Glareolinae

MATERIAL EXAMINED. Stiltia Isabella 1; Glareola pratincola 1.

Family Thinocoridae (Seed-snipe)

MORPHOLOGY. Indistinctly bilobed, papilla moderately developed, tufted (16 feathers).

NOTE. Based upon comparisons of gland morphologies, the present study confirms the belief of

Sibley et al. (1968: 243) that seed-snipe 'are more like ... the Charadriiforms than any other

group.'

MATERIAL EXAMINED. Thinocorus orbignyianus 1; T. rumicivorus 1.

AVIAN UROPYGIAL GLAND

221

Burhinus oedicnemus

H h

Cursorius cursor

Thinocorus rumicivorus

Family Chionididae (Sheath-bills)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, tufted (16-18 feathers, Paris

1913).

MATERIAL EXAMINED. Chionis alba \ .

Family Stercorariidae (Skuas, Jaegers)

MORPHOLOGY. Distinctly bilobed, papilla apparently absent, tufted.

MATERIAL EXAMINED. Stercorarius pomarinus 2 (F: 616-0, 0-37); S. Iongicaudus4(ad M: 333-9, 0-23.

Subad.F: 271-5, 0-28).

Chionis alba

Stercorarius pomarinus

Larus atricilla

222 DAVID W. JOHNSTON

Family Laridae

Subfamily Larinae (gulls)

MORPHOLOGY. Distinctly bilobed and broad (see also Verheyen 1954a), papilla moderately devel-
oped, tufted (18-26 feathers in 4 species, Paris 1913; 22-29 feathers, Jacob & Ziswiler 1982).
MATERIAL EXAMINED. Larus delawarensis 5 (ad. M: 330-0, 0- 1 5. Ad. F: 440-0, 0- 1 3); L. atricilla 8 (ad.
M: 368-5, 0-29. Ad. F: 320-0, 0-20); L. Philadelphia 2 (ad. F: 179-8, 0-21); Rissa tridactyla 2.

Subfamily Sterninae (Terns)

MORPHOLOGY. Similar to Larinae except gland is more compact (see figures); 6-8 feathers in 4

species, Jacob & Ziswiler 1982.

MATERIAL EXAMINED. Chlidonias nigra 2; Hydroprogne caspia (tschegrava of Peters) 4 (F: 686-0,

0'\%;69S-Q,Q-1S); Sterna hirundo 9 (M: 117-4,0-26. F: 137-7,0-33); S. paradisea 2 (M: 110-9,0-16);

S.forsteri4(F: 147-0, 0-33); S. anaethetus 6 (M: 130-0, 0-43; 135-0, 0-31); S.fuscata 8 (M: 218-0,

0-29; 218-6, 0-25. F: 141-0,0-38; 156-0,0-35; 174-0, 0-38); S. albifrons 5 (M: 46-1, 0-46; 51-3, 0-57);

Thalasseus maximus 1 (M: 385-9, 0-35. F: 353-2, 0-26); T. sandvicensis 3; Larosterna inca 2 (M, Z:

153-3, 0-21); Anous stolidus 4\ Gygis alba 2 (M: 117-9, 0-43; 113-3, 0-47).

Family Rynchopidae (Skimmers)

MORPHOLOGY. Bilobed, papilla broad and apparently double, densely tufted (24 feathers).

MATERIAL EXAMINED. Rynchops niger 6 (M: 211-4, 0-20).

Family Alcidae (Auks, Murres, Puffins)

MORPHOLOGY. Elongated and bilobed, papilla moderately developed, tufted (Verheyen \958d;
30-50 feathers, Paris 1913; 2-8 (sic?) feathers, Jacob & Ziswiler 1982; 20-40, present study).
MATERIAL EXAMINED. Plautus alle 2; Pinguinus impennis (mounted bird) 1 ; Uria lomvia 1 ; U. aalge 1 ;
Cepphus grylle 1 (F: 363-0, 0-22); C. columba 3 (M: 380-0, 0-18. F: 371-5, 0-19); Synthliboramphus
antiquus 2 (M: 180-5, 0-25); Ptychoramphus aleuticus 2 (F: 207-9, 0-47); Aethia cristatella 1;
Cerorhinca monocerata 2 (M: 631-5, 0-31); Lunda cirrhata 2 (F: 673-5, 0-30; 792-4, 0-20).

Sterna fuscata

Rynchops niger

Lunda cirrhata

AVIAN UROPYGIAL GLAND 223

Order Columbiformes
Gland characteristics. Naked or absent.

Family Pteroclididae (Sand-grouse)

MORPHOLOGY. Indistinctly bilobed, papilla broad and well developed, naked.

MATERIAL EXAMINED. Syrrhaptes paradoxus 1; Pterocles namaqua 1; P. decoratus 1; P.

lichtensteinii 1 .

Family Columbidae (Pigeons and Doves)

Subfamily Treroninae (Fruit pigeons)

MORPHOLOGY. Absent or, when present, indistinctly bilobed, papilla large, naked.

MATERIAL EXAMINED. Sphenurus apicauda 1 *; S1. oxyura 1 *; Treron curvirostra 2*; T. pompadora 2*;

T. olax 1*; T. vernans 2*; T. bicincta 1*; T. s. thomae 1*; T. australis 1*; T. calva 1*; T. waalia 1*;

Phapitreron leucotis 2; P. amethystina 1; Leucotreron occipitalis 1; Ptilinopus** dupetithouarsii 1;

P. regina 1; P. insular is 1*; P. raratongensis 1; P. huttoni 1; P. porphyraceus 1; P. greyii 1;

P. richardsii 1; P. perousii 1; P. superbus 3; P. pulchellus 2*; P. coronulatus 2*; P. monacha 1;

P. iozonus 1 *; P. r/vo// 3*; P. eugeniae 1 *; P. hypogastra 1 *; P.jambu 1 ; P. aurantiifrons 1 ; P. ornatus

2; P. tannensis 1*; Chrysoena victor 1*; Alectroenas pulcherrima 1; ,4. madagascariensis 2;

Megaloprepia magnified 1 *; Ducula oceanica 1 ; Z). pacifica 1 ; Z). #e«eo 1 ; Z). bicolor 1 ; Z). spilorrhoa

1; Z). fow//'0 1*; Z). rufigaster 1; Z). zoeae 1*.

Subfamily Columbinae (Pigeons, Doves)

MORPHOLOGY. Except for some individuals or varieties ofColumba livia, the gland is present in all
genera and species of Columbinae thus far examined. Indistinctly bilobed, papilla large, naked.
Reported by Beddard (1898) as absent in Ptilopaspuella( = Columbapuella of Peters), Starnoenas
(also absent fide Garrod 1874#), and Turacoena, all genera and species unavailable for the present
study.

MATERIAL EXAMINED. Columba livia^ 2 (M: 268-0, 0-05. F: 312-8, 0-08); C. palumbus 1; C. leuco-
cephala 3 (F: 205-2, 0-10; 264-0, 0-05); C. guinea 1; C. fasciata 1; Macropygia unchall 1; M.
amboiensis 2; M. ruficeps 1 ; M. phasianella \ ; M. nigrirostris 2; Ectopistes migratoria 2; Zenaidura
macroura 8 (M: 123-0, 0-01; 134-4, 0-03; 105-7, 0-02. F: 123-4, 0-01; 117-1, 0-03); Z. auriculata 1;
Zenaida asiatica 1 (M : 1 73 -2, 0-02); Nesopeliagalapagoensis 1 ; Streptopelia orientalis 1 ; S. capicola 1 ;
S. senegalensis 3; Geopelia humeralis 1; G. striata 2; G. cuneata 1; Metriopelia melanoptera 1; M.
aymara 1; Scardafella inca 1; Columbigallina passer ina 1 (F: 41-5, 0-02); C. talpacoti 1; C. minuta 1;
Claravispretiosa 3; Oena capensis 6; Turtur afer 1 ; T. chalcospilos 2; Chalcophaps indica!>; C. stephani
1; Henicophaps albifrons 1; Phaps chalcoptera 1; Ocyphaps lophotes 3; Lophophaps ferruginea 1;
Geophaps smithii 2; Aplopelia larvata \; A. simplex 1; Leptotila verreauxi 1; L. ruf axilla 1; L.
plumbeiceps 3; L. cassini 5;Oreopelia caniceps 1 ; Geotrygon versicolor 1 ; Gallicolumba luzonica 1 ; G.
beccarii 1; G. rubescens 2; Otidiphaps nobilis 1; Caloenas nicobarica 2.

Subfamily Gourinae (Crowned Pigeons)

MORPHOLOGY. Absent

MATERIAL EXAMINED. Gowra cristata 1; G. scheepmakeri 1; G. victoria 1.

Subfamily Didunculinae (Tooth-billed Pigeons)

MORPHOLOGY. Absent. Jacob & Ziswiler (1982) reported a gland in 2 specimens of Didunculus, an

inexplicable difference from the present and all previous reports (Newton 1893-1896, Beddard

1898,Verheyenl957tf).

MATERIAL EXAMINED. Didunculus strigirostris 3.

*gland absent, present study; absent in Treron (Garrod 1 8740). Jacob & Ziswiler ( 1 982 and V. Ziswiler in litt.) found glands

in adult Treron pompadora, T. vernans, T. waalia.

**gland 'very small in Ptilinopus' fide Garrod 1874a.

tabsent in some varieties such as Fantail, Oriental, Roller, Maltese, White Carneau (Darwin 1900, Johansson 1927, Levi

1941,Verheyenl957a).

224 DAVID W. JOHNSTON

Order Psittaciformes*
Gland characteristics. Tufted or absent.

Family Psittacidae (Lories, Parrots, Macaws)

Subfamily Strigopinae (Owl Parrots)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted.

MATERIAL EXAMINED. Strigops habroptilus 1.

Subfamily Nestorinae (Keas)

MORPHOLOGY. Distinctly bilobed, papilla well developed, tufted (13 feathers, Jacob & Ziswiler

1982).

MATERIAL EXAMINED. Nestor notabilis 3.

Subfamily Loriinae (Lories)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted (5-8 feathers in 3 species, Jacob & Ziswiler

1982). Type I.

MATERIAL EXAMINED. Chalcopsitta atra 1; Eos cyanogenia 1; E. squamata 1; E. bornea 3;

Trichoglossus ornatus 1; T. haematodl; T. chlorolepidotus 1; Psitteuteles Johns toniae 1; Domicella

garrula 3; Vini stepheni 1; Glossopsitta porphyrocephala 1; Charmosyna josefinae 1; C. papou 1;

Oreopsittacus arfaki 1 ; Neopsittacus musschenbroekii 1 .

Subfamily Micropsittinae (Pigmy Parrots)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted (3-4 feathers, Jacob & Ziswiler 1982).

MATERIAL EXAMINED. Micropsitta pusio 1.

Subfamily Kakatoeinae (Cockatoos)

MORPHOLOGY. Absent or when present, distinctly bilobed, papilla large, tufted (4-8 feathers in 2
species, Jacob & Ziswiler 1982).

NOTE. Newton (1893-1896: 653) stated that the gland 'exists, though hardly in a functional con-
dition, in ... Cacatua cristata (Cockatoo) . . .' =Kakatoe sulphured citrino-cristata of Peters.
Nitzsch (1867), Garrod 18746, Gadow (1893), and Grasse (1950), noted no gland in Cacatua
sulfurea and its absence in C. roseicapella was reported by Paris (1913). I found that specimens of
both of these species had tufted glands.

P (erodes lichtensteinii

Zenaidura macroura

Ara chloroptera

"many zoo and captive birds.

AVIAN UROPYGIAL GLAND 225

MATERIAL EXAMINED. Probosciger aterrimus 4f; Calyptorhynchus baudinii 1; Callocephalon
fimbriatum 2; Kakatoe galerita 7; K. sulphured 4; K. alba 3; K. moluccensis 1; K. haematuropygia 1;
K. leadbeateri 1; K. sanguinea 1; K. tenuirostris 1; /T. roseicapella 3; Nymphicus hollandicus 4.

Subfamily Psittacinae (Macaws, Parrots)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted (12 feathers, Paris 1913; 3-1 1 feathers in 33
species, Jacob & Ziswiler 1982) or gland absent. Type I.

NOTE. Miller ( 1 924: 324) reported no gland in Orthopsittaca and Diopsittaca ( = Ara of Peters), but
listed no species. Jacob (1978: 168) stated that the gland is absent in Ara but indicated on species.
MATERIAL EXAMINED. Anodorhynchus hyacinthus 5*; A. leari 1 *; Ara** ararauna 2; A. militaris 2; A.
macao 4 (F, Z: 996-5, 0-05); A. chloroptera 3; A. auricollis 1; A. severa 1; A. manilata 3*; Aratinga
acuticaudata 1; A. guarouba 1; A. leucophthalmus 1; A. holochlora 1; A.jandaya 1; A. solstitialis 2;
A. canicularis 1; A. aurea 2; Nandayus nanday 1; Conuropsis carolinensis 1; Rhynchopsitta
pachyrhyncha 1; Cyanoliseus patagonus 1; Pyrrhura rhodogaster 1; P. molinae 1; P. hoffmanni 1;
Myiopsitta monachus 1; Psilopsaigon aurifrons 1; Forpus conspicillatus 1; Brotogeris tirica I*; B.
versicolurus 2*; 5. pyrrhyopterus 1*; B. jugularis 2*; 5. cyanoptera 2*; fi. chrysopterus 4*; 5. s/.
thoma 1*; Pionites melanocephala 2; Graydidascalus brachyurus 2*; Pionus*** menstruus 2*; P.
maximiliani 1*; P. senilis 1*; P. chalcopterus 1*; Amazona leucocephala \*\ A. ventralis \*\ A.
xantholora 1*; ^4. albifrons 1*; .4. ag//w 1*; ^4. vittata 1*; /I. viridigenalis 1*; ,4. autumnalis 1*; ,4.
barbadensis 1*; ,4. aestiva 1*; ^4. ochrocephala 3*; ^4. amazonica 1*; A.farinosa 1*; /I. vinacea 1*;
y4. guildingii 1*; A imperialis 2*; Triclaria malachitacea 1; Poicephalus senegalus 2; P. meyeri 1;
P. ruppellii 2; Psittacus erithacus 1; Coracopsis nigra 1; Psittrichas fulgidus 1; Lorius roratus 1;
Prioniturus discurus 1; Psittacula krameri 5; P. alexandri 1; P. cyanocephala 1; Polytelis swainsonii
2; P. alexandrae 2; Aprosmictus erythropterus 3; Psittinus cyanurus 2; Agapornis roseicollis 2; ^4.
fischeri \; A. lilianae 3; Loriculus vernalis 1; Platycercus elegans 1; P. eximius 3; P. icterotis 2;
P. zonarius 2; Psephotus haematonotus 3; P. varms 3; Neophema elegans 3; TV. chrysostomus 1; TV.
petrophila 1; TV. pulchella 7; TV. splendida 3; TV. bourkii 6; Cyanoramphus auriceps 2; Melopsittacus
undulatus 1 .

Order Cuculiformes
Gland characteristics. Tufted or naked.

Family Musophagidae (Plantain-eaters)

MORPHOLOGY. Flattened and distinctly bilobed, papilla moderately developed, tufted (8 feathers).

Verheyen (19566) was evidently in error when he noted (p. 2) that touracos have a naked gland.

Type I.

MATERIAL EXAMINED. Tauraco corythaix 1; J1. leucolophus 1; Gallirex porphyreolophus 1;

Musophaga violacea 3; Crinifer leucogaster 1; C. africanus 2.

Family Cuculidae (Cuckoos, Roadrunner, Anis)

MORPHOLOGY (family). Flattened and more or less distinctly bilobed, papilla large and often

appearing double, naked.

Subfamily Cuculinae

MATERIAL EXAMINED. Clamator glandarius 1; Cuculus canorus 1; Cacomantis merulinus 1;
Chrysococcyx cupreus 1 ; C. fc/aas 1 ; C. caprius 9; Chalcites basalts 1 .

tgland absent, this study and Beddard (1898).

*gland absent, present study; Jacob & Ziswiler (1982, V. Ziswiler in litt.) found a gland in adult Pionus fuscus.
**gland present in /I. ambigua and maracana (fide Garrod 18746).
''gland also absent in P. sordidus (fide Garrod 1 8746).

***

226

DAVID W. JOHNSTON

Subfamily Phaenicophaeinae

MATERIAL EXAMINED. Coccyzus americanus 4 (unsexed: 46-7, 0-09); Piaya cayana 2; Saurothera
vetula 2; Ceuthmochares aereus 1; Rhopodytes diardi 1; R. tristis 2; Rhamphococcyx curvirostris 2;
Dasylophus super ciliosus 1 .

Subfamily Crotophaginae (Anis, Guiras)

MATERIAL EXAMINED. Crotophaga ani 3 (M: 113-9, 0-03); C. sulcirostris 7; Guiraguira 1.

Subfamily Neomorphinae (Roadrunners, Ground Cuckoos).

MATERIAL EXAMINED. Taper a naevia 1 ; Morococcyx erythropygus 2; Geococcyx californiana 3.

Subfamily Couinae (Couas)
MATERIAL EXAMINED. Coua cristata \ .

Subfamily Centropodinae (Coucals)

MATERIAL EXAMINED. Centropus viridis 2; C. toulou 1; C. benegalensis 2.

Order Strigiformes

Gland characteristics. Naked or minutely tufted.

NOTE. Glands of various strigiform species have usually been described as 'naked' or 'nude'
(Gadow 1893, Beddard 1898, Jacob & Ziswiler 1982). Nitszch (1840), Miller (1924), and I used
magnification and identified 1 to 12 'rudimentary,' 'vestigial,' or 'minute' feathers on the papilla's
tip in some individual specimens.

Family Tytonidae (Barn Owls)

Subfamily Tytoninae

MORPHOLOGY. Indistinctly bilobed, papilla moderately developed, tufted with minute feathers
(1 or 2 5-mm feathers) or naked. Nitzsch (1867: 71) noted minute feathers on the papilla's apex in
Strix flammea, S. perlata, and S.furcata (all = Tyto alba). Type I.
MATERIAL EXAMINED. Tyto alba 7* (M: 502-0, 0-07; 490-0, 0-04. F: 530-0, 0-12; 488-4, 0-11).

Subfamily Phodilinae

MORPHOLOGY. Like Tytoninae. Nitzsch (1867: 71) reported minute feathers at the gland apex in
Strix badia ( = Phodilus badius), but none were seen on 3 specimens in the present study.
MATERIAL EXAMINED. Phodilus badius 3.

Tauraco corythaix

Saurothera vetula

Tyto alba

•minute feathers in 3 specimens.

AVIAN UROPYGIAL GLAND 227

Family Strigidae (Typical Owls)

MORPHOLOGY (family). More superficial than in any other avian family, appearing to lie on top of
the skin ('almost standing up,' Paris 1913: 180), bilobed, papilla large, tufted (up to 10 1-mm
minute feathers) or naked. Type I.

Subfamily Buboninae

MATERIAL EXAMINED (naked unless otherwise indicated). Otus spilocephalus It; O. scops 1;
O. bakkamoena 1; O. asio 26ft, ttt (M: 103-2, 0-07; 107-7, 0-08. F: 149-3, 0-07; 108-4, 0-11);
O. trichopsis It; O. guatemalae If; O. choliba If, ttt; O. watsonii 1; O. leucotis 1; Lophostrix
cristata 2; Bubo virginianus 8ftt (M: 1207-0, 0-04; 1407-0, 0-04. F: 1887-0, 0-04; 1670-0, 0-03); B.
bubo 1; B. africanus Ittt; B. lacetus Ittt; Ketupa^ketupu 1; Pulsatrix perspicillata Ittt; Nyctea
scandiaca 2ftt; Surnia ulula 3 (M: 310-0, 0-07. F: 355-7, 0-04); Glaucidium brasilianum 2; G. brodiei
1; Micrathene whitneyi \\Ninoxnovaeseelandiae Ittt 5 N.philippensis 1; Athene noctual; A. brama
1; Speotyto cunicularia 4; Ciccaba virgata It; C. nigrolineata 1; C. woodfordii \.

Subfamily Striginae

MATERIAL EXAMINED (naked unless otherwise indicated): Strix aluco 1*; S. varia 20**, ****
(M: 762-9, 0-09. x of 9 F: 775-4, 0-07); S. nebulosa 1*; Rhinoptynx clamator 1; Asio otus 5***, ****
(F: 306-0, 0-09); A. madagascariensis 1*; A.flammeus 3*****; Pseudoscops grammicus 1; Aegolius
acadicus5(M:91-l,Q-\Q).

Order Caprimulgiformes
Gland characteristics. Naked or rarely absent.

Family Steatornithidae (Oil-bird)

MORPHOLOGY. Indistinctly bilobed, papilla large, naked (first reported by Garrod 1873).

Described by Paris (1913: 177) and Newton (1893-1896: 653) as 'large.' (See section on Weights

and sizes of glands.)

MATERIAL EXAMINED. Steatornis caripensis 1 .

Family Podargidae (Frogmouths)

MORPHOLOGY. Podargus — absent (see also Gadow 1893, Verheyen 19560, Grasse 1950, Miller

1924). Batmchostomus — indistinctly bilobed, papilla large, naked. The implication by Van Tyne &

Berger (1976) that the gland is absent in (all) Podargidae is incorrect.

MATERIAL EXAMINED. Podargus strigoides 2; P. papuensis 1; P. ocellatus 2; Batrachostomus auritus

1; B. septimus 2; B. stellatus 1; B.javensis 1.

Family Nyctibiidae (Potoos)

MORPHOLOGY. Very small, indistinctly bilobed, papilla large, naked.
NOTE. Miller (1924: 324) reported 'the loss of the oil-gland' in Nyctibius.
MATERIAL EXAMINED. Nyctibius griseus 3.

tminute tuft, present study.

ttminute tuft, up to 8 1-mm feathers in 8 specimens.

ttfsome specimens with minute tuft fide Miller (1924: he also reported tufts in Ketupa zeylonensis, Bubo bubo, Gymnoglaux

lawrencii).

minute tuft, present study.

*minute feathers in 6 specimens.

**minute feathers in 3 specimens.

***some specimens with minute tuft (Nitszch 1840, Beddard 1898, Miller 1924, or Verheyen 1956a).

****'one or 2 very small white feathers,' Paris 1913: 182.

228

DAVID W. JOHNSTON

Asioflammeus

Steatornis caripensis

Batrachostomus septimus

Family Aegothelidae (Owlet-nightjars)

MORPHOLOGY. Broad, flattened and bilobed, papilla large, naked.

MATERIAL EXAMINED. Aegotheles insignis 1.

Family Caprimulgidae (Nighthawks, Goatsuckers)

MORPHOLOGY (family). Very small not apparently bilobed (see also Paris 1913: 173), papilla large,

naked. I did not confirm the report by Arnall & Keymer (1975) that the gland is absent 'in

nightjars.'

Subfamily Chordeilinae (Nighthawks)

MATERIAL EXAMINED. Lurocalis semitorquatus 1; Chordeiles minor 6 (M: 64-8, 0-01; 67-5, 0-01.

F: 75-4, 0-01; 87-5, 0-01; 79-6, 0-01); Podager nacunda 2.

Subfamily Caprimulginae (Goatsuckers)

MATERIAL EXAMINED. Eurostopodus macrotis 1 ; Nyctidromus albicollis 4; Phalaenoptilus nuttallii 2;
Otophones yucatanicus 1; Caprimulgus carolinensis 10 (M: 113-0, 0-01; 80-7, 0-01; 124-8, 0.01. x of
4 F: 1 14-2, 0-01); C. vociferus 2 (M: 55-5, 0-02. F: 52-5, 0-02); Scotornis climacurus 1; Semeiophorus
vexillarius 1 ; Hydropsalis brasiliana 1 .

Nyctibius griseus

Aegotheles insignis

Podager nacunda

AVIAN UROPYGIAL GLAND 229

Order Apodiformes

Gland characteristics. Naked.

NOTE. The reference by Elder (1954) and Pettingill (1985) to gland absence in 'certain species' of

Apodiformes was unsubstantiated by me.

Family Apodidae (Swifts)

MORPHOLOGY (family). Indistinctly bilobed, papilla moderately developed, naked.

Subfamily Chaeturinae (Spine-tailed Swifts)

MATERIAL EXAMINED. Collocalia inexpectata 1; C. vanikorensis 1; Hirund-apus giganteus 2;

Streptoprocne zonaris 1; Chaetura pelagica 3; C. rutilus 1 .

Subfamily Apodinae (Typical Swifts)

MATERIAL EXAMINED. Apus apus 1 ; Aeronautes saxatalis 1 ; Reinarda squamata 1 ; Cypsiurus parva 1 .

Family Hemiprocnidae (Crested Swifts)

MORPHOLOGY. Indistinctly bilobed, papilla absent, naked.

MATERIAL EXAMINED. Hemiprocne mystacea 1 ; H. comata 1 .

Family Trochilidae (Hummingbirds)

MORPHOLOGY. Distinctly bilobed with lobes greatly separated, papilla large, naked.
MATERIAL EXAMINED. Glaucis hirsuta 1 ; Phaethornis superciliosus 2; P. eurynome 1 ; P. longuemareus
2; Eutoxeres condamini 2; Phaeochroa cuvierii 2; Campylopterus curvipennis 8; C. hemileurcurus 1 ;
C. ensipennis 1 ; Eupetomena macroura 1 ; Florisuga mellivora 1 ; Colibriserrirostris 1 ; Anthracothorax
nigricollis 1 ; Chrysolampis mosquitus 1 ; Stephanoxis lalandi 1 ; Chlorestes notatus 1 ; Thalurania
furcata 1; Hylocharis chrysura 1; Chrysuronia oenone 1; Leucochloris albicollis 1; Amazilia Candida
\\A. versicolor \;A. cyanocephala \;A. rutila \;A. tzacatl 1 3; Patagona gigas 1 ; Ensifera ensifera 1 ;
Archilochus colubris 3; Selasphorus rufus 1 .

Hirund-apus (Chaetura) Hemiprocne comata Glaucis hirsuta

giganteus

Order Coliiformes
Gland characteristics. Tufted.

Family Coliidae (Colics)

MORPHOLOGY. Distinctly bilobed, papilla large, tufted. Verheyen (19566') makes the unsubstan-
tiated comment that the gland of Urocolius ( = Colitis indicus and C. macrourus of Peters) is naked.
Both Nitzsch (1867) and Garrod (1876) reported that the gland of Colius is tufted. Type I.
MATERIAL EXAMINED. Colius striatus 4; C. colius 1 ; C. indicus 2; C. macrourus 1 .

230 DAVID W. JOHNSTON

Order Trogoniformes
Gland characteristics. Naked.

Family Trogonidae (Trogons)

MORPHOLOGY. Indistinctly bilobed, papilla large, naked.

MATERIAL EXAMINED. Pharomachrus mocino 2; Priotelis temnurus 1; Temnotrogon roseigaster 1;

Trogon strigilatus 1 ; T. citreola 1 ; Apaloderma marina 1 ; Harpactes erythrocephalus 1 .

Patagona gigas Colius macrourus Pharomachrus mocino

Order Coraciiformes*
Gland characteristics. Much inter- and intrafamilial variation: naked, or sparsely to densely tufted.

Family Alcedinidae (Kingfishers)

Subfamily Cerylinae

MORPHOLOGY. Indistinctly bilobed, papilla absent or small, tufted (16 feathers in C. alcyori).
MATERIAL EXAMINED. Ceryle torquata 1; C. alcyon 6 (M: 102-5, 0-25. F: 104-5, 0-22); C. rudis 7;
Chloroceryle americana 6; C. aena 4.

Subfamily Alcedininae

MORPHOLOGY. Like Cerylinae (12 feathers, Paris 1913, Jacob & Ziswiler 1982).

MATERIAL EXAMINED. Alcedo atthis \;A. meninting \;A. euryzona 1; A. leucogaster 1; Ispidina picta

1; /. madagascariensis 1; Ceyx argentatus 1; C. azureus 1; C. erithacus 1 .

Subfamily Daceloninae

MORPHOLOGY. Indistinctly or distinctly bilobed, papilla absent (Pelargopsis) to large (Tanysiptera},
tufted (small in Lacedo, large in Halcyon) or gland naked (Tanysipterd) (12 feathers in Dacelo}.
Tanysiptera species apparently have no distinctive ecological or behavioral traits that might be
correlated with the unusual naked gland condition (Fry 1980). Type I.

MATERIAL EXAMINED. Pelargopsis capensis 1; Lacedo pulchella 2; Dacelo novaeguineae 3; D. leachii
1; Clytoceyx rex 1; Melidora macrorrhina 1; Halcyon coromanda 1; H. smyrnensis 1; H. pileata 1;
H. senegalensis 1; H. malimbica 1; H. albiventris 1; H. macleayii 1; H. cinnamomina 1; H. chloris 1;
Tanysiptera galatea 2; T. sylvia 1 .

Family Todidae (Todies)

MORPHOLOGY. Indistinctly bilobed, papilla large, tufted (6 feathers in T. subulatus). Nitzsch (1867:
88) erroneously stated that Todus has a naked oil-gland, a point corrected by Forbes (1 882). Type I.
MATERIAL EXAMINED. Todus angustirostris 1; T. subulatus 2.

"morphology reported here is, with exceptions noted below, consistent with the descriptions in Verheyen (1955, a, b, c).

AVIAN UROPYGIAL GLAND

231

Family Momotidae (Motmots)

MORPHOLOGY. Flattened and distinctly bilobed with lobes widely divergent, papilla moderately to

well developed, minutely tufted or naked. Much difference of opinion is found in the literature

concerning the feathered condition of glands in this family probably because some investigators

failed to use magnification in their examinations of glands. By combining here the comments of

Garrod ( 1 878), Forbes ( 1 882), Newton ( 1 893-1 896), Beddard ( 1 898), Miller ( 1 9 1 5), and Verheyen

( 1 955fl) plus microscopic examinations in the present study, it is apparent that any specimen of any

species might have a gland that is naked or one that is tufted with 1-8 'vestigial,' 'rudimentary,' or

very small feathers (ca. 1 mm). Type I.

MATERIAL EXAMINED. Hylomanes momotula 1 ; Electron platyrhynchum 1 ; Eumomota super ciliosa 5;

Baryphthengus ruficapillus 3; Momotus momota 5.

Dacelo novaeguineae

Todus subulatus

Momotus momota

Family Meropidae (Bee-eaters)

MORPHOLOGY. Indistinctly bilobed, papilla large, naked (also reported by Paris 1913: 175).

MATERIAL EXAMINED. Melittophagus pusillus 1 ; Merops apiaster 2; M. viridis 1 ; Nyctyornis amicta 1 .

Family Leptosomatidae (Cuckoo-rollers)

MORPHOLOGY. Indistinctly bilobed, papilla large, naked. The gland, about 10 mm in length, does
not conform to Nitzsch's description of 'atrophy and almost total disappearance . . .' (1867:161).
MATERIAL EXAMINED. Leptosomus discolor 2.

Family Coraciidae (Rollers)

MORPHOLOGY (family). Flattened and indistinctly bilobed, papilla large, naked.

Subfamily Brachypteraciinae

MATERIAL EXAMINED. Brachypteracias leptosomus 1; Uratelornis chimaera 1.

Subfamily Coraciinae

MATERIAL EXAMINED. Coracias garrulus \\Eurystomusorientalis 1.

Merops apiaster

Leptosomus discolor

Coracias garrulus

232

DAVID W. JOHNSTON

Family Upupidae (Hoopoes)

MORPHOLOGY. Distinctly bilobed with widely diverging lobes, papilla large, tufted (10 feathers,

Paris 1913, Grasse 1950; 14 feathers, Jacob & Ziswiler 1982). Type I.

MATERIAL EXAMINED. Upupa epops 2.

Family Phoeniculidae (Wood-hoopoes)

MORPHOLOGY. Small and not apparently bilobed, papilla large, tufted (10 feathers). Type Ha.

MATERIAL EXAMINED. Phoeniculus purpureus 1 ; P. bollei \ ; Rhinopomastus minor 1 ; R. cyanomelas 1 .

Family Bucerotidae (Hornbills)

MORPHOLOGY. Distinctly bilobed (lobes completely separated), papilla small, tufted (50 feathers,

Paris 1913; 32-48 feathers in T. erythrorhynchus, Jacob & Ziswiler 1982). In Tockus hartlaubi the

gland and its feather tuft are 'vestigial' (Verheyen \955a). Type II.

MATERIAL EXAMINED. Tockus alboterminatus 1; T. erythrorhynchus 1; T.flavirostris 1; T. deckeni 1;

Aceros undulatus 1 ; A.plicatus 1 ; Anthracoceros malabaricus \;A. coronatus 1 ; Ceratogymna atrata

1; Buceros bicornis 1; B. hydrocorax 1; Bucorvus abyssinicus 2.

Upupa epops

Phoeniculus purpureus

Tockus erythrorhynchus

Order Piciformes

Gland characteristics. Much inter- and intrafamilial variation: absent (rarely), naked, or sparsely
to densely tufted.

NOTE. Differences (see figures) in gland morphology among the six families lend evidence to a
polyphyletic origin of the Piciformes as suggested by Olson (1983).

Family Galbulidae (Jacamars)

MORPHOLOGY. Indistinctly bilobed, papilla moderately developed, naked.

MATERIAL EXAMINED. Galbalcyrhynchus leucotis 1; Brachygalba lugubris 1; Galbula albirostris 5;

G. galbula 1; G. ruficauda 3; Jacamerops aurea 1 .

Family Bucconidae (Puff-birds)

MORPHOLOGY. Distinctly bilobed, papilla large, naked. Gadow (1893), Nitzsch (1867: 94), and
Beddard (1898) each refer to some bucconids (e.g., Malacoptilafusca, Bucco, Monasa) as having
glands with 'a few fine hairs at the apex' or 'feathered.' However, Miller (1915) and I found that all
species and individuals in the Bucconidae that we examined had naked glands.

AVIAN UROPYGIAL GLAND 233

MATERIAL EXAMINED. Notharchus macrorhynchos 3; Nystalus maculatus 1; Hypnelus bicinctus 1;
Malacoptila striata 1 ; M.fusca 1 ; M. panamensis 6; Monasa nigrifrons 1 ; M. atra 1 ; M. morphoeus 1 ;
Chelidoptera tenebrosa 2.

Family Capitonidae (Barbels)

MORPHOLOGY. Distinctly bilobed, papilla moderately developed, naked or sparsely tufted (8-12
feathers in 2 species, Jacob & Ziswiler 1982). In addition to the species marked * below, the
following species were reported by Miller (1924: 323) as having naked glands: Stactolaema,
Pogoniulus duchaillui, Trachyphonus cafer, and T. margaritatus. Individual differences (naked vs.
tufted) have been found in Trachyphonus vaillantii, T. darnaudii, and Lybius torquatus. A feathered
gland was reported for Pogonias (Lybius) by Nitzsch (1867: 93). Beddard (1898: 168) noted that
(all) capitonids have feathered glands. I did not confirm the statement by Verheyen (\955b) that
different species in the Capitonidae lack a gland. Type I.

MATERIAL EXAMINED (tufted unless otherwise indicated). Semnornis frantzii 1; S. ramphastinus 1;
Psilopogonpyrolophus 2; Megalaima rafflesii 2; M. mystacophanos 1 ; M.flavifrons 1 ; M. asiastica 1 ;
M. henricii 1; M. haemacephala 1; Gymnobucco bonapartei 2*; Smilorhis leucotis 1*; Pogoniulus
simplex I; P. bilineatus \;P. subsulphureus 1; Tricholaema leucomelan \*;T. diadematum 1*; Lybius
guifsobalito 1*; L. leucocephalus 1*; L. dubius 2*; Trachyphonus purpuratus 1; T. vaillantii 2*;
T. darnaudii 1*.

Galbula ruficauda Notharchus macrorhnychos Megalaima rafflesii

Family Indicatoridae (Honey-guides)

MORPHOLOGY. Indistinctly bilobed, papilla moderately developed, tufted (2 feathers). Miller

(1924: 323) correctly noted that the Indicatoridae are invariably tufted 'but the tuft is vestigial in

Prodotiscus.' I did not confirm the statement by Verheyen (1955Z?) that different species in the

Indicatoridae lack a gland. Type I.

MATERIAL EXAMINED. Prodotiscus insignis 2; Indicator exilis 1; /. minor 1; /. maculatus 2;

Melichneutes robustus 1 .

Family Ramphastidae (Toucans)

MORPHOLOGY. Distinctly bilobed, papilla poorly developed, tufted (8 feathers). Type I.
MATERIAL EXAMINED. Aulacorhynchus prasinus 1 ; Pteroglossus torquatus 2 (M, Z: 1 83-3, 0- 14; 1 86-3,
0-12); Andigena hypoglauca 1; Ramphastos vitellinus 1; R. discolorus 1; R. sulfur atos \;R. swainsoni
1; R. tucanus 1; R. cuvieri 1; R. inca 1; R. toco 2.

Family Picidae (Wryneck, Piculets, Woodpeckers)

MORPHOLOGY (family). Absent or distinctly bilobed with widely separated lobes, papilla usually

moderately developed, naked or tufted (8-12 feathers in 3 species, Jacob & Ziswiler 1982). In some

*naked, present study.

234

DAVID W. JOHNSTON

North American species (e.g., Dryocopus, Colaptes) each lobe narrows down to an extremely small
'band' before joining at the papilla, making dissection and removal of an intact gland difficult.
Miller ( 1 924) noted the gland's absence in Campethera maculosa, permista, caroli, and nivosa, these
in addition to C. cailliartii in the present study. The gland is naked in Dinopium and Gecinulus and
naked or tufted in specimens of Chrysocolaptes validusfide Miller (1924).

Subfamily Jynginae (Wrynecks)

MORPHOLOGY. 'Well developed and clearly bilobed' (Paris 1913: 168), tufted (8 feathers, Paris

191 3). Type II.

MATERIAL EXAMINED. Jynx torquilla 1 .

Subfamily Picumninae (Piculets)

MORPHOLOGY. Tufted.

MATERIAL EXAMINED. Picumnus cirratus 1 ; Nesoctitesmicr omegas 1 ; Sasia ochracea 2; 5". abnormis 1 .

Subfamily Picinae (Woodpeckers)

MORPHOLOGY. Tufted, naked, or absent. Type I.

MATERIAL EXAMINED (tufted unless otherwise indicated). Colaptes auratus 1 (M: 100-9, 0-12. F:

131-9, 0-12; 137-7, 0.11; 98-5, 0-12); Piculus simplex 1; Campethera punctuligera 2; C. nubica 4;

C. bennettii 1; C. cailliautii 2*; C. abingoni 2; C. permista 1*; C. caroli 2**; C. nivosa 7*; Celeus
flavescens 2; Micropternus brachyurus 1; Picus viridis 1; Dinopium beneghalense 2***; D.javanense

1 ***; Dryocopus pileatus 5 (M: 240-8, 0- 1 5. F: 220-8, 0- 1 2); Asyndesmus lewis 1 ; Melanerpes erythro-
cephalus 2; M. carolinus 1 (M: 76-0, 0-09; 72-2, 0-14. F: 54-9, 0-12); M. aurifrons 3; M.flavifrons 1;
Leuconerpes candidus 1; Sphyrapicus varius 4 (M: 43-4, 0-12; 50-3, 0-08. F: 45-5, 0-14); Trichopicus
cactorum 2; Veniliornis fumigatus 2; V. passerinus 1 ; V. affinis 1 ; Dendrocopos hyperythrus 1 ;

D. villosus 1 (F: 54-3, 0-11); D.pubescens 2; Picoides arcticus 1; Xiphidiopicus percussus 1; Thripias
pyrrhogaster 1; Hemicircus canete 1; Blythipicus pyrrhotis 1; B. rubiginosus 1; Chrysocolaptes
validus 2***; C. lucidus 8; Phloeoceastes guatemalensis 1; P. melanoleucus 1; P. leucopogon 1;
P. haematogaster 1 ; Campephilus principalis 1 ; C. magellanicus 1 .

NOTE. In his comprehensive study of woodpeckers of the world, Short ( 1 982) presents no ecological,
structural, or behavioral information that might correlate with gland presence/absence, tufted/
naked condition in different species of Campethera, Dinopium, or Chrysocolaptes.

Indicator maculatus

Ramphastos toco

Colaptes auratus

*gland absent, this study; also absent in C. maculosa (Miller 1924).

**gland present (tufted) or absent in some specimens, this study.

***gland naked, this study. Tuft is individually variable in specimens of Chrysocolaptes (Miller 1924: 324).

AVIAN UROPYGIAL GLAND 235

Order Passeriformes

Gland characteristics. Naked.

MORPHOLOGY (order). Indistinctly or distinctly bilobed, papilla moderately or well developed,
naked. Although varying slightly in shape ('kidney-vs heart-shaped'), weight, and relative length
of papilla (Jacob & Ziswiler 1982), glands of all passerines have been uniformly described by all
authors as being present and naked. In the present comprehensive study, representatives of all
passerine families (68, Peters 1931-1986) and subfamilies were examined: 1 187 individuals of 349
genera and 482 species. Except for relative size (see Weights and sizes of glands section), I found
no consistent, major morphological differences between or among any taxa. Paris (1913: 67) in his
extensive study reported only slight variations in shape among at least 1 1 passerine families.

Suborder Eurylaimi
Family Eurylaimidae (Broadbills)

Subfamily Eurylaiminae

MATERIAL EXAMINED. Smithornis capensis 1 ; Eurylaimus javanicus 1 ; Psarisomus dalhousiae 1 .

Subfamily Calyptomeninae

MATERIAL EXAMINED. Calyptomena whiteheadi 1 .

Suborder Tyranni
Superfamily Furnarioidea

Family Dendrocolaptidae (Wood-hewers)

MATERIAL EXAMINED. Dendrocincla anabatina 8; D. homochroa 4; Sittasomus griseicapillus 4;
Glyphorhynchus spirurus 10; Drymornis bridgesii 1; Dendrocolaptes certhia 7; Xiphorhynchus
ocellatus 1; X. guttatus 2; X. flavigaster 3; Lepidocolaptes souleyetii 3; Campy lor hamphus
trochilirostris 1 ;

Family Furnariidae (Ovenbirds)

MATERIAL EXAMINED. Geositta cunicularia 1; Upucerthiavalidirostris 1; Cinclodesfuscus \\Furnarius
leucopus 1 ; Aphrastura spinacauda 2; Phleocryptes melanops 2; Schizoeacafuliginosa 1 ; Synallaxis
albescens 1 ; S. erythrothorax 2; Poecilurus candei 1 ; P. scutatus 1 ; Asthenes hudsoni 1 ; Phacellodomus
striaticollis 1; Coryphistera alaudina 2; Anumbius annumbi 2; Margarornis squamiger 1; Pseudosei-
sura lophotes 2; Ancistrops strigilatus 1; Syndactyla rufosuperciliata 2; Philydor erythrocercus 1;
Automolus infuscatus 1; A. albigularis I; A. ochrolaemus 7; Heliobletus contaminatus 1; Xenops
minuta 1 1 ; Sclerurus guatemalensis 4.

Family Formicariidae (Ant-thrushes)

MATERIAL EXAMINED. Taraba major 1; Thamnophilus doliatus 5; T. aethiops 1; Myrmotherula
surinamensis 1 ; Microrhopias quixensis 2; Formicivora grisea 1 ; Drymophila caudata 1 ; Cercomacra
tyrannina 5; C. nigricans 1; Hypocnemis cantator 1; Myrmeciza longipes 1; Formicarius colma 1;
F. analis 5; Chamaeza ruficauda 1; Pithy s albifrons 1; Gymnopithys leucaspis 1; Hylophylax
naevioides 1 ; Grallaria guatimalensis 1 .

Family Conopophagidae (Ant-pipits)

MATERIAL EXAMINED. Conopophaga lineata 1; C. castaneiceps 1; Corythopis torquata 1.

Family Rhinocryptidae (Tapaculos)

MATERIAL EXAMINED. Pteroptochos tarnii 1; Scelorchilus rubecula 1; Rhinocrypta lanceolata

1 ; Teledromasfuscus 1 ; Melanopareia maximiliani 1 ; Scytalopus latebricola 1 .

236 DAVID W. JOHNSTON

Superfamily Tyrannoidea

Family Tyrannidae (Tyrant Flycatchers)

Subfamily Elaeniinae

MATERIAL EXAMINED. Sublegatus modestus 1; Myiopagis viridicata 2; Elaenia flavogaster 3; E.
pallatangae 1; Mionectes olivaceus 1; M. oleagineus 28; Leptopogon amaurocephalus 7; Oncostoma
cinereigulare 8; Todirostrum sylvia 1; T. cinereum 2; Rhynchocyclus brevirostris 1; Tolmomyias
sulphurescens 4; Platyrinchus cancrominus 4; P. mystaceus 4.

Subfamily Fluvicolinae

MATERIAL EXAMINED. Onychorhynchus coronatus 3; Terenotriccus erythrurus 1; Myiobius barbatus
2; Contopus virens 1; C. cinereus 1; Empidonax flaviventris 3; £". virescens 1; £. minimus 5; Sayornis
phoebe 2; Ochthoecafumicola 1; Myiotheretes striaticollis 1; Xolmis irupero 1; Muscisaxicola sp. 1;
Knipolegus aterrimus 1; Fluvicola pica 1.

Subfamily Tyranninae

MATERIAL EXAMINED. /J////0 spadiceus 7; Rhytipterna simplex 1; Myiarchus tuber culifer 2; Af.
nuttingi 2; M. crinitus 2 (M: 42-4, 0-08); Pitangus sulphuratus 3; Megarhynchus pitangua 2;
Myiodynastes bairdii 1 ; Tyrannus tyrannus 2 (F: 40-0, 0-10); T7. melancholicus \ .

Subfamily Tityrinae

MATERIAL EXAMINED. Pachyramphus cinnamomeus 1; Tityra semifasciata 2;
r. inquistor 3.

Family Pipridae (Manakins)

MATERIAL EXAMINED. Schiffornis turdinus 3; Chloropipo uniformis 1; Xenopipo atronitens 1;

Manacus manacus 10; Chiroxiphia lanceolata 1; Piprafilicauda 1; P. mentalis4Q; P. chloromeros 1.

Family Cotingidae (Cotingas)

MATERIAL EXAMINED. Ampelion rubrocristatus 1; Pipreola arcuata 1; P. chlorolepidota 1; Lipaugus
vociferans 1 ; Gymnoderus foetidus 1 ; Querula purpurata 1 ; Pyroderus scutatus 1 ; Cephalopterus
ornatus 1 ; Perissocephalus tricolor 1 ; Procnias nudicollis 1 ; Rupicola peruviana 1 .

Family Oxyrunicidae (Sharpbills)
MATERIAL EXAMINED. Oxyruncus cristatus 1 .

Family Phytotomidae (Plantcutters)
MATERIAL EXAMINED. Phytotoma rutila 1 .

Family Pittidae (Pittas)

MATERIAL EXAMINED. P/fte erythrogaster 1; P. granatina 1.

Family Philepittidae (Asitys)

Subfamily Philepittinae

MATERIAL EXAMINED. Philepitta castanea 1 .

Subfamily Neodrepanidinae

MATERIAL EXAMINED. Neodrepanis coruscans 1.

Family Acanthisittidae (New Zealand Wrens)

MATERIAL EXAMINED. Acanthisitta chloris 2; Xenicus longipes 5; A', gilviventris 1 .

NOTE. Most specimens available for examination were poorly preserved. I identified a gland in

Acanthisitta, in only 1 of the 5 Xenicus longipes, and not in A', gilviventris.

AVIAN UROPYGIAL GLAND 237

Suborder Menurae
Family Menuridae (Lyrebirds)
MATERIAL EXAMINED. Menura novaehollandiae 2.

Family Atrichornithidae (Scrub-birds)

MATERIAL EXAMINED. Atrichornis clamosus \ .

NOTE. B. Gillies (in litt., 4 April 1985) reported that this specimen (Rl 1353) has a naked gland;

another specimen (A 15926) is illustrated in Zusi (1985) as having a naked gland.

Suborder Oscines

Family Alaudidae (Larks)

MATERIAL EXAMINED. Mirafrajavanica 1 ; M. assamica 1 ; Eremopterix signata 1 ; Alaemon alaudipes
1; Melanocorypha yeltoniensis 1; Calandrella cinerea 1; Galerida cristata 1; Lullula arborea 1;
Eremophila alpestris 1 .

Family Hirundinidae (Swallows)

MATERIAL EXAMINED. Tachycineta bicolor 2; Progne subis 1 (F: 63-1, 0-03); Hirundo rustica 3;

H. smithii 1 .

Family Motacillidae (Wagtails, Pipits)

MATERIAL EXAMINED. Dendronanthus indicus 1; Motacilla alba 2; M. aguimp 1; Macronyx croceus
\\Anthusspinoletta 1 (M: 19-2, 0-13).

Family Campephagidae (Cuckoo-shrikes)

MATERIAL EXAMINED. Coracina novaehollandiae 1; C. striata 1; C. morio 1; C. panayensis 1;
C. melaschistos 1; Lalage nigra 1; Campephaga phoenicea 1; Pericrocotus cinnamomeus 1;
P.flammeus 1; Hemipus picatus 1.

Family Pycnonotidae (Bulbuls)

MATERIAL EXAMINED. Pycnonotus barbatus 8; P. goiavier 1 ; Chlorocichlaflaviventris 2; Bleda eximia

1; Criniger phaeocephalus 1; Setornis criniger 1; Hypsipetes everetti 1.

Family Irenidae (Leaf Birds)
MATERIAL EXAMINED. Irenapuella 1.

Family Laniidae (Shrikes and Allies)

Subfamily Prionopinae

MATERIAL EXAMINED. Eurocephalus ruppelli 1; Prionops plumata 1.

Subfamily Malaconotinae

MATERIAL EXAMINED. Dryoscopus cubla 3; D. sabini 1; Tchagra senegala 1; T. australis 2; Laniarius
ferrugineus 3; L. barbarus 1; Telophorus sulfureopectus 1; T. multicolor 1.

Subfamily Laniinae

MATERIAL EXAMINED. Corvinella corvina 1; Lanius collurio 1; L. ludovicianus 2.

Subfamily Pityriasinae

MATERIAL EXAMINED. Pityriasis gymnocephala 1.

Family Vangidae (Vangas)

MATERIAL EXAMINED. Calicalicus madagascariensis 1 ; Vanga curvirostris 1 .

Family Bombycillidae (Waxwings)

Subfamily Bombycillinae

MATERIAL EXAMINED. Bombycilla garrulus 1; B. cedrorum 1 (F: 29-2, 0-10).

238 DAVID W. JOHNSTON

Subfamily Ptilogonatinae

MATERIAL EXAMINED. Ptilogonys cinereus 1 .

Subfamily Hypocoliinae

MATERIAL EXAMINED. Hypocolius ampelinus 1 .

Family Dulidae (Palm Chat)
MATERIAL EXAMINED. Dulus dominions 1 .

Family Cinclidae (Dippers)

MATERIAL EXAMINED. Cinclus cinclus 1 ; C. pallasii 1 ; C. mexicanus 4 (unsexed: 63-6, 0-48; 58-3, 0-65;

59-7,0-71).

NOTE. Nitzsch (1867: 73) reported that the gland of Cinclus 'bears small down-feathers upon its

surface,' but it is not clear that his 'surface' refers to the papilla's tip. All specimens examined in the

present study had naked glands.

Family Troglodytidae (Wrens)

MATERIAL EXAMINED. Campylorhynchus rufinucha 1; Cistothorus platensis 2; C. palustris 2;
Thryothorus pleurostictus 2; T. maculiectus 2; T. ludovicianus 1; T. rufalbus 1; Troglodytes aedon 2;
Uropsila leucogastra 1; Henicorhina leucosticta 3.

Family Mimidae (Mockingbirds and Allies)

MATERIAL EXAMINED. Dumetella carolinensis 1 7 (F: 35-4, 0- 1 8); Mimuspolyglottos 5 (M: 47-6, 0- 1 6);

Toxostoma rufum 5 (M: 55-3, 0-07. F: 69-1, 0-14).

Family Prunellidae (Accentors)
MATERIAL EXAMINED. Prunella collaris 1.

Family Muscicapidae*

Subfamily Turdinae (Thrushes)

MATERIAL EXAMINED. Zeledonia coronata 4; Sialia currucoides 1; Catharus fusee scens 2 (F: 28-1,
0-09); C. minimus 2; C. ustulatus 3; C. guttatus 2; Hylocichla mustelina 28 (F: 47-4, 0-09; 59-7, 0-07);
Turdus merula 1; T. iliacus 1; T.philomelos 1; T. viscivorus 1; T.grayi4; T. migratorius 4 (M: 85-3,
0-09. F: 72-4, 0-09; 85-7, 0-12).

Subfamily Orthonychinae (Logrunners)
MATERIAL EXAMINED. Cinclosoma cinnamomeum 1 .

Subfamily Timaliinae (Babblers)

MATERIAL EXAMINED. Trichastoma bicolor 1; Malacopteron magnum 1; Pomatorhinus schisticeps
1; Napothera brevicaudata 1; Chamaeafasciata 2; Turdoides squamiceps 1; Garrulax leucolophus 3;
Actinodura ramsayi 1; Alcippe castaneceps 1.

Subfamily Panurinae (Parrotbills)
MATERIAL EXAMINED. Paradoxornis heudei 1 .

Subfamily Picathartinae (Picathartes)
MATERIAL EXAMINED. Picathartes oreas 1.

Subfamily Polioptilinae (Gnatcatchers and allies)

MATERIAL EXAMINED. Ramphocaenus melanurus 2; Polioptila caerulea 1 .

"nomenclature and inclusive taxa according to Peters (Vol. X, 1964).

AVIAN UROPYGIAL GLAND 239

Family Sylviidae (Old World Warblers)

MATERIAL EXAMINED. Locus tella lanceolata 1; Acrocephalus scirpaceus 1; Cisticola erythrops 1;
Sylvietta rufescens 1; Hylia prasina 1; Abroscopus schisticeps 1; Sylvia communis 1; S. hortensis 1;
Regulus calendula 6; R. satrapa 1 .

Family Muscicapidae (Old World Flycatchers)**
MATERIAL EXAMINED. Muscicapa dauurica 1 .

Family Platysteiridae (Puffback Flycatchers)
MATERIAL EXAMINED. Batis molitor \ .

Family Maluridae (Australo-Papuan Wrens)
MATERIAL EXAMINED. Malurus lamberti 1 .

Family Acanthizidae (Australasian Warblers)

Subfamily Acanthizinae

MATERIAL EXAMINED. Sericornis magnirostris 1 .

Subfamily Mohouinae

MATERIAL EXAMINED. Unavailable.

Family Monarchidae (Monarch Flycatchers)

Subfamily Monarchinae

MATERIAL EXAMINED. Terpsiphone viridis 1; T. atrocaudata 1; Chasiempis sandwichensis 1.

Subfamily Rhipidurinae

MATERIAL EXAMINED. Ripidura albicollis \ .

Family Eopsaltriidae (Australasian Robins)

MATERIAL EXAMINED. Petroica phoenicea 1; P. vittata 1; Tregellasia leucops 1.

Family Muscicapidae***

Subfamily Pachycephalinae (Whistlers)
MATERIAL EXAMINED. Pachycephala lanioides 1 .

Family Aegithalidae (Long-tailed Tits, Bush Tits)
MATERIAL EXAMINED. Aegithalos caudatus 2.

Family Remizidae (Penduline Tits)
MATERIAL EXAMINED. Auriparusflaviceps 2.

Family Paridae (Titmice)

MATERIAL EXAMINED. Parus atricapillus 1; P. carolinensis 1; P. bicolor 1; Hypositta corallirostris 1.

Family Sittidae

Subfamily Sittinae (Nuthatches)

MATERIAL EXAMINED. Sittapusilla 1; S. canadensis 1; S. carolinensis 1.

Subfamily Daphoenosittinae (Treerunners)

MATERIAL EXAMINED. Neositta chrysoptera 3; Daphoenositta miranda 2.

* ""nomenclature and inclusive taxa according to Peters (Vol. XI, 1986).
***nomenclature and inclusive taxa according to Peters (Vol. XII, 1967).

240 DAVID W. JOHNSTON

Subfamily Tichodromadinae ( Wallcreepers)
MATERIAL EXAMINED. Tichodroma muraria \ .

Family Certhiidae (Creepers)

Subfamily Certhiinae (Treecreepers)
MATERIAL EXAMINED. Certhiafamiliaris 2.

Subfamily Salpornithinae (Spotted Creeper)
MATERIAL EXAMINED. Salpornis spilonotus 1 .

Family Rhabdornithidae (Philippine Creepers)
MATERIAL EXAMINED. Rhabdornis mysticalis 1 .

Family Climacteridae (Australian Treecreepers)
MATERIAL EXAMINED. Climacteris melanura 1 .

Family Dicaeidae (Flowerpeckers)

MATERIAL EXAMINED. Rhamphocharis crassirostris 1 ; Prionochilus olivaceus 1 ; Dicaeum concolor 1 ;

D. cruentatwn 1; Oreocharis arfaki 1; Pardalotus rubricates 1.

Family Nectariniidae (Sunbirds)

MATERIAL EXAMINED. Anthreptes malacensis 1; Hypogramma hypogrammicum 1; Nectarinia
olivacea 1 ; N. senegalensis 7; N. sericea 1 ; N. jugularis 2; N. asiatica 1 ; N. venusta \ ; N. talatala 2;
N. habessinica 1; Aethopyga boltoni 1; Arachnothera longirostra 1 .

Family Zosteropidae (White-eyes)
MATERIAL EXAMINED. Zosterops griseotincta 1 .

Family Meliphagidae (Honeyeaters)

MATERIAL EXAMINED. Oedistoma iliolophum 1; Myzomela sanguinolenta 1; M eliphaga fusca 1;
M. pencillata 1; Melithreptus brevirostris 1; Philemon citreogularis 1; Melidectes fuscus 1;
Acanthorhynchus tenuirostris 1; Anthochaera carunculata \.

Family Emberizidae

Subfamily Emberizinae (Buntings and American Sparrows)

MATERIAL EXAMINED. Emberiza flaviventris 1; Calcarius lapponicus 1; Zonotrichia melodia 4
(M: 17-6, 0-12); Z. georgiana 3; Z. albicollis 7; Junco hyemalis 1; Ammodramus sandwichensis 3;
A. savannarum 3; Spizella passerina 1; S. pusilla 1; Pooecetes gramineus 1; Aimophila aestivalis 2;
Sicalis olivascens 1; Volatinia jacarina 2; Sporophila torqueola 6; S. telasco 1; Camarhynchus
crassirostris 1; Pipilo erythrophthalmus 3 (M: 43-8, 0-26. F: 36-7, 0-22); Arremon aurantiirostris 3;
Arremonops rufivirgatus 3; A. chloronotus 1.

Subfamily Catamblyrhynchinae (Plush-capped Finch)
MATERIAL EXAMINED. Catamblyrhynchusdiadema 1.

Subfamily Cardinalinae (Cardinal-grosbeaks)

MATERIAL EXAMINED. Pheucticus ludovicianus 1; P. melanocephalus 1 (F: 40-5, 0-09); Cardinalis
cardinalis 5 (M: 40-3, 0-09; 33-0, 0-07. F: 40-0, 0-09); Saltator atriceps 1; S. maximus 2;
S. aurantiirostris 1; Passerina cyanoides 8; P. caerulea 2; P. cyanea 6.

Subfamily Thraupinae (Tanagers)

MATERIAL EXAMINED. Eucometis pencillata 2; Lanio aurantius 3; Tachyphonus luctuosus 1; Habia
rubica 3; H.fuscicauda \ 6; Piranga rubra 2; .P. olivacea 3 (F: 26-5, 0-06); Ramphocelus sanguinolentus
4; R. passer inii 4\ Thraupis episcopus 1 ; 7\ bonariensis 1 ; Euphonia affinis 2; Dacnis cyana 1 ; Cyanerpes
cyaneus 4; Diglossa carbonaria 1 .

AVIAN UROPYGIAL GLAND 241

Subfamily Tersininae (Swallow-tanager)
MATERIAL EXAMINED. Tersina viridis 1 .

Family Parulidae (Wood Warblers)

MATERIAL EXAMINED. Mniotilta varia 3; Vermivora peregrina 2; Parula americana 1; Dendroica
petechia 4; D. magnolia 1; D. coronata 3; D. cerulea 1; D.fusca 3; D. pensylvanica 2; D. castanea 2;
D. striata 1 ; D.pinus 1 ; D.palmarum 1 ; Setophaga ruticilla4; Seiurus aurocapillus 6; S. noveboracensis
9 (M: 1 6-2, 0- 1 5); S. motacilla 2; Helmitheros vermivorus 6; Protonotaria citrea 1 ; Geothylpis trichas
4 (M: 11-5, 0-09); G. poliocephala 1; G.formosa 20; Wilsonia pusilla 4; Icteria virens 4; Coereba
flaveola 2.

Family Drepanididae (Hawaiian Honeycreepers)

MATERIAL EXAMINED. Himatione sanguinea 1; Palmeria dolei 1; Vestiaria coccinea 1; Loxops virens

1.

Family Vireonidae (Peppershrikes, Shrike- Vireos)

Subfamily Cyclarhinae

MATERIAL EXAMINED. Cyclarhis gujanensis 1 .

Subfamily Vireolaniinae

MATERIAL EXAMINED. Vireolanius pulchellus 1.

Subfamily Vireoninae

MATERIAL EXAMINED. Vireo griseus 3; V.flavifrons2\ V. solitarius 2; K. olivaceus 3; V.flavoviridis3;
V. gilvus 1 ; Hylophilus ochraceiceps 4; //. decurtatus 1 .

Family Icteridae (American Orioles and Blackbirds)

Subfamily Icterinae

MATERIAL EXAMINED. Psarocolius montezuma 2; Amblycerus holosericeus 1; Icterus galbula 1; /.
spurius 2; /. dominicensis 1; Agelaius phoeniceus 26 (x of 16 M: 55-4, 0-17. x of 9 F: 42-8, 0-18);
Sturnella magna 4 (M: 105-5, 0- 1 1 . F: 78-6, 0- 1 3): S1. neglecta 1 ; Quiscalus mexicanus 1 ; (2- ma/or 23
(x of 14M:192-9,0-17.xof9F:91-0, 0-21); Q.quiscula6(M: 113-4, 0-16; 117-4, 0-14; 119-1, 0-14.F:
84-2, 0-17); Euphagus carolinus 1 (M: 66-0, 0-13); Molothrus ater 15 (x of 6 M: 47-5, 0-13. x of 8 F:
37-3,0-14).

Subfamily Dolichonychinae

MATERIAL EXAMINED. Dolichonyx oryzivorus 12 (x of 10 M: 39-0, 0-09).

Family Fringillidae

Subfamily Fringillinae (Chaffinches and Brambling)
MATERIAL EXAMINED. Fringilla coelebs 1; F. montifringilla 1 .

Subfamily Carduelinae (Serins, Goldfinches, et al.)

MATERIAL EXAMINED. Serinus mozambicus 14; Carduelis pinus 2 (M: 10-5, 0-10); C. tristis 1;
Carpodacus purpureus 2 (M: 26-2, 0-03); C. mexicanus 2 (F: 23-2, 0-11); Pinicola enucleator 4
(F: 55-2, 0-06; 56-8, 0-05; 60-2, 0-03); Coccothraustes vespertinus 1 (M: 56-6, 0-05).

Family Estrildidae (Waxbills, Grass Finches, and Mannikins)

MATERIAL EXAMINED. Pytilia melba 2; Uraeginthus angolensis 8; Estrilda caerulescens 1; Poephila

acuticauda 1; P. cincta 1; Chloebia gouldiae 1; Lonchura cucullata \\Amadinafasciata 1 .

Family Ploiceidae

Subfamily Viduinae (Indigo-birds and Whydahs)
MATERIAL EXAMINED. Vidua paradisaea 1 .

242 DAVID W. JOHNSTON

Subfamily Passerinae

MATERIAL EXAMINED. Passer domesticus 14 (M: 24-5, 0-09; 24-8, 0-20. F: 22-3, 0-18; 25-1, 0-18);
P. griseus 3.

Subfamily Bubalornithinae

MATERIAL EXAMINED. Dinemellia dinemelli 1 .

Subfamily Ploceinae

MATERIAL EXAMINED. Amblyospiza albifrons 1; Ploceus subaureus 3; P. xanthops 1; P. velatus 4;
P. cucullatus 25; Euplectes hordeaceus 1 ; E. orix 1 1 .

Family Sturnidae (Starlings)

Subfamily Sturninae

MATERIAL EXAMINED. Sturnus vulgaris 5 (M: 78-3, 0-11; 80-5, 0-13; 89-7, 0-11); Sarcops calvus 1;
Gracula religiosa 1 .

Subfamily Buphaginae

MATERIAL EXAMINED. Buphagus erythrorhynchus 2; Buphagus sp. 1 .

Family Oriolidae (Orioles)

MATERIAL EXAMINED. Oriolus oriolus 1; O. chinensis 1; O. xanthornus 1.

Family Dicruridae (Drongos)

MATERIAL EXAMINED. Dicrurus remifer 1; D. hottentottus 1 ; D. paradiseus 1.

Family Callaeidae (New Zealand Wattlebirds)

MATERIAL EXAMINED. Callaeas cinerea 1 ; Creadion carunculatus 1 ; Heteralocha acutirostris 1 .

Family Grallinidae (Australian Mud Nest Builders)

Subfamily Grallininae

MATERIAL EXAMINED. Grallina cyanoleuca 1 .

Subfamily Corcoracinae

MATERIAL EXAMINED. Corcorax melanorhamphos 1; Struthidea cinerea 2.

Family Artamidae (Wood-swallows)

MATERIAL EXAMINED. Artamus fuscus 1; A. leucorhynchus 1; A. super ciliosus 1; A. cinereus 1;

A. minor 1.

Family Cracticidae (Australian Butcherbirds)

MATERIAL EXAMINED. Cracticus nigrogularis 1 ; Gymnorhina tibicen 1 ; Strepera graculina 1 .

Family Ptilonorhynchidae (Bowerbirds)

MATERIAL EXAMINED. Ailuroedus crassirostris 1; Amblyornis macgregoriae 2; Sericulus

chrysocephalus 1; Ptilonorhynchus violaceus 1; Chlamydera nuchalis 2.

Family Paradisaeidae (Birds of Paradise)

MATERIAL EXAMINED. Manucodia comrii 1 ; Semioptera wallacei 1 ; Astrapia stephaniae 1 ; Lophorina

superba 1 ; Cicinnurus regius 1 ; Diphyllodes respublica 1 ; Paradisaea apoda 1 .

Family Corvidae (Crows, Magpies, Jays)

MATERIAL EXAMINED. Cyanocitta cristata 14 (M: 77-7, 0-20; 78-4, 0-10. F: 59-4, 0-08); Aphelocoma
coerulescens 2 (M: 70-3, 0-1 1; 72-0, 0-07); Garmlus glandarius 1; Pica pica 2; Corvus monedula 1;
C.frugilegus 2; C. brachyrhynchos 39 (x of 14 M: 563- 1 , 0-05. x of 1 1 F: 495-7, 0-05); C. ossifragus 4
(M: 330-0, 0- 15. F: 232-0, 0- 1 3); C. corone 2.

AVIAN UROPYGIAL GLAND 243

Dendrocolaptes certhia Cindus mexicanus Corvus brachyrhynchos

Weights and sizes of glands

Early accounts of uropygial glands included brief comments about relative size ('small,' 'large,'
'smaller than'; Willughby 1678; Burton 1822, Macgillivray 1837, Bartlett 1861). Such relative
adjectives and phrases persist in the more contemporary literature (Austin 1961, Thomson 1964,
Shortt 1977). Edwards Crisp (1860, 1862) was probably the first person to publish gland weights,
and, by weighing birds and their glands separately, he presented the relative proportion of gland
weight to the bird's body weight. He (1860: 258) presented data on 34 species of aquatic and
terrestrial birds, showing relatively lightest glands in pigeons and heaviest ones in Cindus aquaticus
(C. cinculus aquaticus of Peters) and six species of waterfowl. The frequent assertion that the preen
gland of water-birds is relatively larger than that of land-birds (e.g., Kennedy 1971) is probably
derived from Crisp's results (see also Coues 1890), even though Frederick II in 1260 reported large
glands in aquatic species (Wood and Fyfe 1943).

Subsequent authors have presented absolute or relative gland weights for many more species
(Kossman 1871, Paris 1913, Kar 1947, Grasse 1950, Elder 1954, Kennedy 1971, Johnston 1979,
Jacob & Ziswiler 1982). In these reports, differences in relative gland weights have variously been
attributed to season (Kossman 1871, Kennedy 1971), habitat (Crisp 1860, Jacob & Ziswiler 1982),
intergeneric body weight (Johnston 1 979), nutrition (Kossman 1871), individual variation (present
study and others), and sex (Groebbels 1 932). Although Elder ( 1 954) suggested that glands of diving
ducks (Aythyd) are relatively heavier than those of dabbling ducks (Anas), some of Jacob &
Ziswiler's data (1982: 214) 'clearly refute this hypothesis.' Subsequently I compared relative gland
weights from 7 species of dabbling ducks (N = 21, x = 0-30%, SD = 0-047) with weights from 5
species of diving ducks (N = 20, x = 0-29, SD = 0-065); the differences were not statistically
significant (d.f. = 40, t = 0-7560, p > 0-05).

Jacob & Ziswiler (1982) presented gland weights from 574 individuals in 183 species, and I
obtained gland weights from 544 individuals in 200 species. All these weights are presented in Table
1 . A comparison of these two data sets shows reasonable agreement for the same taxon especially
as regards mean values. Also apparent are variations in relative gland weights within and between
species, variations that I attribute largely to individual body weight differences. The latter are
probably due to sexual differences (see, for example, data for three species of Icterinae in the
Systematic accounts) and variations in the amounts of subcutaneous fat.

From their analysis of relative gland weights, Jacob & Ziswiler concluded that 'the only thing
that can be said with certainty regarding the size of the uropygial gland is that birds that swim and
dive have, without exception, a large uropygial gland' (1982: 214). This statement should be
expanded to include the earlier demonstrated correlations with season, nutrition, and sex by the
other authors mentioned above.

I was able to examine the habitat-habit issue more thoroughly because of a much larger sample
size, including birds living in most major habitat types. By grouping relative weights of birds at the
family level and to broad habitat-habit categories (Fig. 2), I found that the largest (relative) glands

244

DAVID W. JOHNSTON

Table 1
Summary of uropygial gland weights

PRESENT STUDY

JACOB & ZISWILER (1982)

No. of
species

No. of

glands

Relative gland
weight*

No. of
species

No. of
glands

Relative gland
weight*

Tinamidae

1

12

0-18

Diomedeidae

1

1

0-30

Procellariidae

5

9

0-46 (0-30-0-67)

2

4

0-24-0-27

Hydrobatidae

4

6

0-16(0-07-0-37)

Spheniscidae

1

2

0-08(0-06-0-12)

2

3

0-21-0-26

Gaviidae

4

9

0-18(0-09-0-30)

2

3

0-14-0-20

Podicipedidae

2

6

0-26(0-19-0-42)

2

7

0-48-0-61

Phaethontidae

1

1

0-43

Fregatidae

1

3

0-07 (0-06-0-07)

Phalacrocoracinae

1

10

0-25(0-20-0-35)

3

12

0-21-0-31

Anhinginae

1

8

0-15(0-12-0-17)

Sulidae

1

1

0-38

1

2

0-35

Pelecanidae

2

4

0-36(0-30-0-43)

1

1

0-19

Ardeidae

9

20

0-14(0-01-0-42)

3

7

0-05-0-07

Ciconiidae

1

1

0-05

1

2

0-08

Threskiornithidae

3

4

0-15(0-11-0-19)

3

4

0-03-0-11

Phoenicopteridae

2

4

0-18-0.21

Cathartidae

2

10

0-04(0-01-0-05)

Pandioninae

1

2

0-25(0-18-0-31)

1

1

0-36

Accipitrinae

11

25

0-06(0-01-0-13)

7

17

0-04-0-22

Falconidae

4

15

0-07 (0-03-0-09)

1

4

0-08

Anatidae

25

55

0-26(0-10-0-42)

15

55

0-08-0-54

Anhimidae

1

1

0-13

Megapodiidae

1

1

0-18

Tetraonidae

6

7

0-05(0-02-0-18)

3

11

0-09-0-15

Phasianidae

8

17

0-07(0-02-0-16)

5

12

0-08-0-11

Meleagrididae

1

3

0-03 (0-02-0-04)

1

2

0-05

Numididae

2

3

0-05-0-18

Gruidae

1

4

0-05 (0-03-0-05)

1

1

0-05

Aramidae

1

7

0-28(0-17-0-42)

Rallidae

6

27

0-19(0-11-0-34)

5

14

0-09-0-24

Haematopodidae

1

1

0-20

Charadriidae

2

4

0-18(0-07-0-22)

1

2

0-16

Scolopacidae

9

25

0-12(0-05-0-20)

4

5

0-11-0-18

Recurvirostridae

1

1

0-24

Phalaropodidae

2

4

0-59 (0-42-0-79)

Burhinidae

1

1

0-09

Chionididae

1

1

0-17

Stercorariidae

2

3

0-29 (0-28-0-37)

Larinae

3

5

0-20(0-13-0-29)

4

19

0-12-0-20

Sterninae

9

20

0-33(0-18-0-57)

4

5

0-19-0-44

Rynchopidae

1

1

0-20

Alcidae

5

7

0-27(0-18-0-47)

3

7

0-17-0-29

Pteroclididae

1

1

0-02

Columbidae

5

11

0-04(0-01-0-10)

4

5

0-02-0-04

Psittacidae

2

1

0-11(0-05-0-16)

41

125

0-04-0-19

Cuculidae

2

2

0-06 (0-03-0-09)

2

4

0-12-0-16

Tytonidae

1

4

0-09 (0-04-0- 12)

1

7

0-08

Strigidae

6

22

0-07(0-03-0-11)

4

9

0-04-0-09

Steatornithidae

1

0-22

Caprimulgidae

3

11

0-01 (0-01-0-02)

1

0-01

Apodidae

2

11

0.04-0-05

Alcedinidae

1

2

0-24(0-22-0-25)

2

0-18

Upupidae

2

0-14

Meropidae

1

0-11

Bucerotidae

2

4

0-08-0-11

AVIAN UROPYGIAL GLAND

245

Ramphastidae

1

2

0-1 3 (0-1 2-0- 14)

2

5

0-08-0-10

Picidae

5

13

0-1 2 (0-08^)- 14)

4

9

0-09-0-12

Tyrannidae

2

2

0.09(0-08-0-10)

Alaudidae

1

1

0-28

Hirundinidae

1

1

0-03

1

6

0-21

Motacillidae

1

1

0-13

1

2

0-27

Bombycillidae

1

1

0-10

Cinclidae

1

3

0-61 (0-48-0-71)

Troglodytidae

1

3

0-58

Mimidae

2

4

0-14(0-07-0-18)

Prunellidae

1

1

0-28

Turdinae

3

6

0-09(0-07-0-12)

2

7

0-08-0-14

Aegithalidae

1

2

0-21

Paridae

2

23

0-14-0-15

Sittidae

1

1

0-12

Emberizinae

2

3

0-20(0-12-0-26)

2

6

0-26-0-31

Cardinalinae

2

4

0-09 (0-07-0-09)

1

2

0-18

Thraupinae

1

1

0-06

Parulidae

2

2

0-1 2 (0-09^)- 15)

Icterinae

6

55

0-15(0-ll^)-24)

Dolichonychinae

1

10

0-09(0-06-0-13)

Fringillidae

5

7

0-06(0-03-0-11)

3

26

0-22-0-25

Estrildidae

6

24

0-17-0-23

Passerinae

1

4

0-16(0-09-0-20)

Ploceidae

3

17

0-19-0-28

Sturnidae

1

3

0-12(0-11-0-13)

1

8

0-10

Corvidae

4

32

0-10(0-04-0-20)

7

40

0-08-0-12

*gland weight as percent of body weight; mean (extremes)

are found in nonpasserines that swim, dive or rest on water (N= 18 families, x = 0-28%). The
smallest glands occur in terrestrial (non-aquatic) birds: nonpasserines (N= 15, x = 0-07%) and
passerines (N = 1 6, x = 0-04%).

I found the largest relative gland weights in the Procellariidae (Oceanodroma melania, 0-69%
Fulmarus glacialis, 0-67%) Phalaropodidae (Phalaropus fulicarius, 0-79%), Sterninae (Sterna
albifrons, 0-57%), and Cinclidae (Cinclus mexicanus, 0-71 %). The largest glands reported by Jacob
& Ziswiler (1982) were for Tachybaptus ruficollis (0-61%) and Troglodytes troglodytes (0-58%).
Because of its habit of plunging into water for fish, the Osprey expectedly has a larger (0-25%)
gland than any other of the Falconiformes (Accipitrinae, 0-06% and Falconidae 0-07%).
The smallest glands, which could be accurately weighed in the present study, were found in
Caprimulgidae (11 individuals averaging 0-01%), Meleagrididae (3 individuals, x = 0-03%) and
Columbidae (1 1 individuals, x = 0-04%).

Burton (1822: 4; and quoted by Murphy 1936) reported that the uropygial gland of Fregata
aquila is a 'trifling size.' In my study, fresh weights of birds and glands of Fregatidae were available
only for F. magnificens , so I could not compare relative gland weights among frigatebirds. How-
ever, the length of glands (sans feathers) were as follows: F. magnificens, 1 5 mm; F. aquila, 13 mm;
F. ariel, 1 3 mm. The gland of F. aquila is thus no smaller than that of F. ariel which is the smallest
species of Fregata (Nelson 1975). Frigatebirds have smaller glands relative to body weight (0-07%)
than other nonpasserine birds that live on or in water (e.g., Procellariidae, 0-44%; Phaethontidae,
0-30%; Pelecanidae, 0-28%; Anatidae, 0-27%) with the exception of Spheniscidae (0-08%). The
putative relationship between a small, 'insufficient' gland and feathers becoming so wet that
frigatebirds drown (first proposed by Burton in 1822 and paraphrased by Welty 1962) lacks
scientific verification (see related discussion on spread-wing posture in Clark 1969).

My findings enable me to correct several unverified statements in the literature on gland sizes.
Gurney's (1913) assertion, paraphrasing Ticehurst, that the gland of Sula bassana 'is the largest
proportionally' of all birds, now turns out to be incorrect. One relative gland weight for this species
was only 0-38% (present study), compared with relatively much heavier glands in procellarids,

246

DAVID W. JOHNSTON

CO

E

LJJ
CO
CO

TERRESTRIAL -

SWIM IN WATER -

TERRESTRIAL -

<0 FLY OVER WATER -

LU

1
LU

$ WALK IN WATER -

Ma

1

SWIM, DIVE, OR REST
ON WATER

oooooo o

O 000000*10 o oooo
M

00 O OO. OO O o

o = AVERAGE FOR A FAMILY
M = MEAN

oo o o ooo ooo o ooo o o

3000

M

.00 .10 .20 .30 .40 .50 .60 -70

UROPYGIAL GLAND WEIGHT AS PERCENT OF BODY WEIGHT
Fig. 2 Relationships between uropygial gland weights and avian habitat-habits.

anatids, phalaropes, and others (above). Austin (1961) (see also Shortt 1977) wrote that dippers
(Cinclidae) have a 'tremendous preen gland, ten times the size of that of any other passerine bird.'
To be sure, the gland of Cinclus mexicanus (x = 0-61; 0-48-0-7 1%) is the largest yet reported for
any passerine, but other passerine families (e.g., Emberizidae, Icteridae) have glands as large as
0-24-0-26% and Troglodytes troglodytes has a large gland (0-56-0-58%, Kennedy 1971; Jacob and
Ziswiler 1982).

My data show that large (i.e., heavy) birds have absolutely large uropygial glands. For 670
individuals, representing 61 families of passerine and nonpasserine birds, I found a significant
correlation between body weight and gland weight (r = 0-694, P ^ 0-0 1 ). This correlation is import-
ant especially because of the inverse relation of plumage weight (as a percent of body weight) with
body weightier se (Kossman 1871, Turcek 1966). Kennedy (1971: 370) correctly cautioned that
'this parallel could result from a functional connection between the preen gland [size and] secretion
and the area of feathers which require anointing with it. Additionally, it is possible that relative to
body weight, water-birds have a larger area of feather surface requiring anointing with secretion
than land-birds of similar size, which may partly explain their larger glands.'

Conclusions drawn from size and weight relationships of glands must still be tentative. Despite
the large numbers of weights and broad taxonomic coverage presented in this study and others
(Kennedy 1971, Jacob & Ziswiler 1982), gland weights have never been reported from many
birds — e.g., Apterygidae, wild Psittacidae, Trochilidae, Coliidae, most of the coraciiform and
piciform families, and most of the passerines.

Feathers on uropygial glands

Feathers attached to the papilla at the end of the uropygial gland are collectively termed circulus
uropygialis by Lucas & Stettenheim (1972) and Baumel et al. (1979). Through the years these
feathers have been variously described in different birds as 'contour' (Nitzsch 1867), 'down' or
'downy' or 'modified down' (Nitzsch 1867, Newton 1893-1896, Beddard 1898, Verheyen 1956/,
1958c, d, Grasse 1950, Lucas & Stettenheim 1972, Baumel et al. 1979), 'semiplumes' (Nitzsch

AVIAN UROPYGIAL GLAND

247

1840), 'plumules' or 'plumulets' (Paris 1913), 'plumes' (Beddard 1898), with or without a rachis
and/or hyporachis (Paris 1913, Verheyen 1959c). Some of the earlier publications (e.g., Nitzsch
1867) even described 'fine hairs' at the tip of certain glands. Miller (1924) was apparently the first
investigator to use magnification in determining the number and type of feathers on a gland.

The number of feathers per gland ranges from 1 (minute) to 90 (Jacob & Ziswiler 1982, the
present study). Jacob & Ziswiler provided a thorough discussion of the number, arrangement,
density, and length of the feathers. Because of some individual variation in number of feathers and
other considerations, Jacob & Ziswiler (correctly in my opinion) cautioned against the use of
feather number for taxonomic or diagnostic criteria. Rather, from a functional standpoint, they
noted a general tendency for waterbirds to have more and longer feather tufts than landbirds. They
also believed that the proportional length of the papilla to that of the tuft is taxonomically specific
(see also Schumacher 1919).

Lucas and Stettenheim (1972) classified uropygial gland feathers as 'modified down,' defining
down as feathers with a rachis shorter than the longest barbs and semiplumes having a rachis that
exceeds the longest barbs. My microscopic study of gland feathers from 70 families containing
tufted glands revealed the presence of three feather types (Fig. 3). Most of the family representa-
tives (62) had feathers of type I which, by the definition of Lucas & Stettenheim (1972), are down.

TYPE I

-4-* T jTT-v- (TRUNCATE)

BARB

CALAMUS

TYPE II

TYPE lla

Fig. 3 Diagrams of typical uropygial gland feathers.

CALAMUS

248 DAVID W. JOHNSTON

Three families each had feathers of Types II and Ha, both defined here as semiplumes. Nodal
structures on barbules on these feathers differed from those on true down and contour feathers
(Douglas Deedrick and Roxie Laybourne, pers. comm.). Therefore, the most appropriate terms
for describing the circulus uropygialis are modified down or modified semiplumes.

Contrary to the reports of some other investigators (e.g., Lucas & Stettenheim 1972), I found no
afterfeathers on any gland feathers. Even in groups (e.g., Galliformes) renowned for having
afterfeathers on body contour feathers, afterfeathers were not found. This difference might be
attributable to the criterion for identifying an afterfeather. Lucas & Stettenheim (p. 252) regarded
any group of outgrowths on the rim of the superior umbilicus as an afterfeather; outgrowths were
not identified in the present study.

Naked and tufted glands

Most early investigators such as Nitzsch (1867), Beddard (1898), and Paris (1913) generally cate-
gorized glands as either tufted or naked (nude, bare), that is, with or without feathers on the
papilla. For the most part, the glands that they examined were 'obviously' (unmagnified) tufted or
naked, although occasional references were made to a 'fine hair' at the tip of some glands (Nitzsch
1 867). This dichotomous difference apparently served well until some putative 'naked' glands were
examined with magnification by Miller (1924) and were found to possess 1-2 mm feathers.

It is now desirable to establish three categories of glands with respect to the degree of feathering
on the papilla: naked (no feathers observable, even with magnification), minutely tufted (feathers
detected only with magnification), and tufted (feathers observable without magnification). Glands
previously considered to be naked by Paris (1913) and others but now known to be 'minutely
tufted' include species in the families Apterygidae, Opisthocomidae, Tytonidae, Strigidae, and
Momotidae. In each of these families, considerable individual variation has been found between
the naked and minutely tufted conditions (see Systematic accounts).

Naked glands are also morphologically variable, particularly as regards the length and width of
the papilla. In all passerine birds, for example, the papilla is distinct and well defined. Most
nonpasserine naked glands, on the other hand, either have no papilla (Rhinochetidae, Columbidae,
Hemiprocnidae, Galbulidae) or the papilla is so ill-defined and broad that it appears to be
continuous with the glandular lobes (e.g., Apterygidae, Cathartidae, Cuculidae). Although I
believe the passerine gland shape is distinctive, some nonpasserine glands superficially resemble the
passerine type — Cariamidae, Steatornithidae, Batrachostomus. Close examinations of figures in
the Systematic accounts will show the distinctiveness of the naked passerine gland as opposed to
the several naked nonpasserine ones. Jacob & Ziswiler (1982) recognized different shapes among
many passerine glands (heart-shaped, kidney-shaped, etc.), but these designations were so variable
between and within families that I could not use them.

The majority of nonpasserines have obviously tufted glands, whereas passerine glands are
uniformly naked. From an examination of all nonpasserine gland types, I conclude that the
tufted gland is primitive. Derived types include those that are (1) minutely tufted, (2) naked
(nonpasserine), and (3) naked (passerine).

Gland absence

The absence of uropygial glands in certain species of birds has been known at least since Nitzsch
(1840). As more species were examined over the years, more were found to lack glands. For
example, in the Columbidae, Garrod ( 1 8740) noted gland absence in only 4 genera, Beddard ( 1 898)
and Grasse (1950) in 6 genera, and Verheyen (19570) added 'new' species in Treron and Goura. I
made a special effort to examine as many species and individuals as available in the Columbidae,
Psittacidae, and Picidae because of earlier discrepancies in reports for genera and species in these
families. Nearly every published account containing any information on the absence of uropygial

AVIAN UROPYGIAL GLAND 249

glands (Beddard, Paris, Grasse, Elder, VanTyne & Berger, and others) include at least one factual
error on the subject, sometimes simply by omission and frequently by uncritically copying a
statement from an earlier author.

The glandless condition varies markedly at every taxonomic level: absent from entire orders,
families, genera, species, and individuals. A gland might be present in some species of a genus,
yet absent in others. At the individual level, for example in each of three species of Ptilinopus
(coronulatus, pulchellus, rivoli), some individuals possess glands, whereas others do not. Darwin
(1900) and Levi (1941) reported the gland's absence in certain varieties ofColumba livia.

I found the gland to be absent in the following taxa (see Systematic accounts for details and
pertinent comments): Struthionidae (all age groups), Rheidae (adults), Casuariidae (all age
groups), Dromaiidae (adults), Mesoenatidae (all 3 species), Otidae (all 5 species examined),
Columbidae (9 genera, 28 species), Psittacidae (6 genera, 31 species), Podargus (3 species
examined), and Picidae (1 genus, 4 species).

Some minor discrepancies exist between my findings and earlier reports. Although Garrod
(1 874/?) reported no gland in Cacatua sulfur (Kakatoe sulphured), a live bird that I examined had a
conspicuous, tufted gland. Nitzsch ( 1 840), Beddard ( 1 898), and others noted no gland in Argusianus
(Argus); 5 specimens in the present study contained a gland. I suspect these discrepancies, as well
as those in Ara and Cacatua roseicapella, can be attributed to individual variation among the
specimens examined, most or all of them being captive birds.

Because the glandless condition is found in such a wide diversity of species and other taxa, a
quest for causal relationships is appropriate. Why, for example, do some parrots have glands
whereas others do not? No single attribute (distribution, climate, ecology, flight, etc.) has been
found to be consistent as an explanation, a conclusion also reached by Kossmann (1871) who
stated that he could find no relationship between gland absence and 'way of life of the bird.' Some
flightless birds lack glands; others do not. Some insular pigeons have glands; others do not. Some
neotropical parrots lack glands; others have well-developed glands. In my opinion, uropygial
glands have apparently been secondarily and independently lost in a variety of birds, but these
losses remain unexplained.

Although Beddard (1 898: 232) stated that 'presence or absence cannot be made use of as a fact of
great systematic importance,' he (pp. 3 1 3-3 1 4 and elsewhere) nonetheless used the 'fact' as a family
characteristic. Similarly, Garrod (1874&), Nitzsch (1840), Paris (1913) and many others have used
presence or absence of glands (in addition to naked vs. tufted conditions) as distinctive family and
generic characteristics. This is a valid use of uropygial gland data except in cases of known
individual variation. Gland absence in all species of the Mesoenatidae is just as good a family
characteristic as is their singular limited distribution. On the other hand, in the Picidae, and
especially in only a few species of Campethera, gland absence is probably of little taxonomic
importance at the family level.

Glands in flightless birds

Because some authors (e.g., Elder 1954) have suggested relationships among uropygial gland
secretions, normal feather functions, and flight capabilities, an analysis of gland presence/
absence in flightless birds is desirable. Elder's experiments on ducks essentially showed that gland
extirpation resulted in reduced feather waterproofing, thus rendering the birds flightless.

The first consideration has been to determine if a relationship exists between a flightless con-
dition and gland presence in nature. In Table 2, flightless species identified from several literature
sources are listed along with the presence or absence of the gland. Except for most of the ratites
(Struthionidae, Rheidae, Casuariidae, Dromaiidae), only one other taxon (all three species of
Mesoenatidae) is known wherein flightless species lack a gland. Overall, this analysis reveals
virtually no correlation between a flightless condition and gland absence: for 42 flightless species
examined, only 8 lacked a gland in the adult.

250 DAVID W. JOHNSTON

Table 2

The Relationship between the flightless condition*
and presence of uropygial glands

Struthionidae**

Struthio camelus — absent in adult

Rheidae**

Rhe a americana — absent in adult
Pterocnemia pennata — absent in adult

Casuariidae**

Casuarius bennetti — unavailable
Casuarius casuarius — absent in adult
Casuarius unappendiculatus — unavailable

Dromaiidae**
Dromaius novaehollandiae — absent in adult

Apterygidae

Apteryx australis — gland present, essentially naked (but see Beddard 1898, 1899)
Apteryx owenii — gland present, naked
Apteryx haastii — gland present, naked

Spheniscidae

16 species — gland present and tufted in all 10 species examined

Podicipedidae

Rollandia microptera — gland present, tufted
Podilymbus gigas — gland present, tufted

Phalacrocoracidae

Phalacrocorax harrisi — gland present, tufted

Anatidae

Tachyeres pteneres — gland present, tufted

Tachyeres brachypterus — gland present, tufted

Anas aucklandica — gland present, tufted

Mergus australis — gland present, tufted (but perhaps capable of flight, see Weller 1980)

Mesoenatidae

Mesoenas variegata — gland absent
Mesoenas unicolor — gland absent
Manias benschi — gland absent

Rallidae***

Rallus owstoni — gland present, tufted
Rallus wakensis — gland present, tufted
Cabalus modestus — unavailable
Atlantisia rogersi — gland present, tufted or naked
Tricholimnas lafresnayanus — unavailable
Tricholimnas sylvestris — gland present, tufted
Dryolimnas cuvieri aldabranus — gland present, tufted
Cyanolimnas cerverai — gland present, tufted
Nesoclopeus poeciloptera — unavailable
Gallirallus australis — gland present, tufted
Habropteryx insignis — unavailable
Habroptila wallacii — unavailable
Megacrex inepta — unavailable
Porzanula palmeri — gland present, tufted
Pennula san dwichensis — unavailable
Aphanolimnas monasa — unavailable
Tribonyx mortierii — gland present, tufted
Porphyriornis nesiotis — gland present, tufted
Porphyriornis comeri — gland present, tufted
Notornis mantelli — gland present, tufted

AVIAN UROPYGIAL GLAND 25 1

Rhynochetidae

Rhynochetos jubatus — gland present, naked

Alcidae

Pinguinus impennis — gland present, tufted

Psittacidae

Strigops habroptilus — gland present, tufted

Acanthisittidae
Xenicus lyalli — unavailable (see Systematic accounts for other species).

*Flightless species names taken from Thomson (1964), Greenway (1958), Austin (1961), Van Tyne & Berger (1976), Olson

(1973fl, b\ Weller (1980), and Mlikovsky (1982).

**Several authors (e.g., Beddard 1898, Jacob 1978) have reported the absence of a gland in all ratites except Apteryx,

although the species examined were usually not identified by the author.

""•""Opinions differ on the flight capability of some of these species. Ripley and Beehler (1985: 7), for example, reported that

Rallus owstoni 'can fly as high as one or two meters above the ground, but they seldom do so.'

A second consideration concerns a possible relationship between gland size and a flightless
condition. Although fresh gland weights from flightless species were unavailable to me, sizes (linear
measurements) of glands of flightless species in the Podicipedidae, Phalacrocoracidae, Anatidae,
and Rallidae were compared with glands from closely related (often congeners) species that fly.
These comparisons revealed no major size differences in glands between flightless and flying birds.

General taxonomic considerations of glands

The use of the uropygial gland as a character in avian systematics has been both commonplace and
controversial for many years. As early as 1840, Nitzsch identified general gland features (e.g.,
tufted vs. naked conditions) as characteristics of different avian taxa, and the gland was subse-
quently much used in classification by ornithologists such as Coues (1890) and Beddard (1898).
This use in taxonomy has continued to date by some investigators (e.g., Olson & Steadman's 1981
characterization of Pedionomus), but others have excluded gland morphology in taxonomic con-
siderations (e.g., Cracraft, 1985). Thomson (1964) stated that the gland is 'unsatisfactory as a
taxonomic character,' and Jacob & Ziswiler (1982) noted that the gland 'has little systematic
importance '

From the systematic accounts of this monograph, it can be seen that the gland's presence or
absence, tufted vs. naked condition (and variations thereof) might vary at any taxonomic level:
intraspecific to interordinal. At the ordinal level, the gland is present and naked (with distinctive
papilla) in all the Passeriformes, thus adding, as it were, another passerine characteristic. On the
other hand, the several morphological variations in glands of the piciform families lend credence, I
believe, to Olson's (1983) suggestion for a polyphyletic origin of the Piciforms (see also Burton
1984).

The morphological gland characteristics that could be used in taxonomic analyses are:

1 . ontogeny — e.g., gland present in embryos and young of some ratites, but absent in
all age groups of other ratites.

2. presence or absence of the gland — e.g., absent from families (Mesoenatidae in the
Gruiformes) and genera (Amazona in the Psittacidae).

3. lobe shape — e.g., cf. Apodidae and Trochilidae in the Apodiformes.

4. tufted vs. naked condition —

a. degree of feathering (cf. Momotidae and Meropidae)

b. shape, size, and length of papilla (cf. Leptosomatidae and Coraciidae).

5. histology — little is known about histological variations, but features such as the
number of gland openings are mentioned by Jacob & Ziswiler (1982).

252 DAVID W. JOHNSTON

The value for a cladistic taxonomic scheme would depend on the number of variable features of the
gland that could be analyzed and the incidence of multiple evolutions of those features. The recent
chemo-taxonomic approach of Jacob & Ziswiler ( 1 982), based on chemical differences in uropygial
secretions among different taxa, has as yet received little attention in avian taxonomic schemes.

I believe that gland morphology is just as important a diagnostic taxonomic character as are
muscle variations, osteological minutiae, incubation patterns, syringeal structures, and the like.
The question is, of course, the degree of importance that one assigns to gland morphology in a large
suite of taxonomic characters. Because of significant variations in gland morphology in different
taxa, as identified in the present study, the least that could be said here is that gland morphology
should be considered especially in cladistic taxonomic approaches.

Functions of glandular secretions

Uropygial gland functions (actually, the functions of glandular secretions) have been controversial
ever since the gland was first described in the 13th century. Form and function of glands are
biologically interrelated features, but, because the present report concentrates on gland mor-
phology, only a brief summary of secretion functions is included here. (More detailed accounts can
be found in Law (1929), Elder (1954), Thomson (1964), and Jacob & Ziswiler (1982)). At least 8
functions have been ascribed to the gland, and the interested reader is referred to the appropriate
publications:

1. water-repellent action (Stubbs 1910, Elder 1954, Rijke 1970),

2. preserve physical structure of feathers (Rutschke 1960),

3. maintain horny sheath of bill (Thomson 1964),

4. as a scent organ (Giebel 1857, Jackson 1938, Mackworth-Praed and Grant 1970),

5. pheromone-producing (Balthazar! and Schoffeniels 1974),

6. antirhachitic action (Hou 1928),

7. prevent growth of skin microorganisms (King and McLelland 1984),

8. dislodge feather lice (Morris 1836#).

Apparently any one or some combination of these functions could be ascribed to the secretions of
an individual species but also the functions might not be identical for all birds. Most of the research
on water-repellency ('waterproofing') has been appropriately conducted on aquatic birds, but
virtually nothing is known about 'waterproofing' in landbirds. Indeed, Rutschke (1960) believed
that the gland is only indirectly involved in 'waterproofing' of plumage in aquatic birds (see also
Clark 1969), and Spearman (1971) made the unsupported comment that the glandular products
are 'not essential for terrestrial birds.' Hou's studies (1928) on rickets and vitamin D were
conducted only on chickens, pigeons, and, later, ducks. A scent-organ function for the gland has
been reported for a variety of birds (e.g., Anas moschata, Phoeniculus bollei), but it is not clear how
the 'foul-smelling' (to humans) secretions actually function. Contrary to the research reported by
Elder (1954) on ducks, a number of reports have indicated that some birds from which glands had
been surgically removed nonetheless had 'normal, bright plumages' (Arnall and Keymer 1975). As
early as 1910, Pycraft expressed 'grave doubts' as to the function of the gland primarily because
(1) he believed that some birds (e.g., Anastomus) presumably could not remove oil from the gland
because of their peculiar bill structure and (2) birds lacking glands presumably keep their feathers
in as good condition as those species possessing those glands.

Throughout much of the literature on uropygial glands, one finds the recurring suggestion that
powder down somehow fulfills the function of oil from glands in those species where the gland is
small or absent (Bartlett 1861, Nitzsch 1867, Newton 1893-1896, Verheyen 1956/, Voitkevich
1966, Jacob 1978, Goodwin 1983). This presumed correlation arose, I believe, because observers
were seeking functional replacements in those birds either lacking glands or possessing small
glands. Goodwin (1983: 27), for example, reports for pigeons, The powder down . . . appears to

AVIAN UROPYGIAL GLAND 253

function in lieu of preen oil to aid in waterproofing of the feathers.' Verheyen (1956/, 19570)
variously describes the glands of Psittaciformes and Columbidae as being in a 'phase of regression'
or 'deficient,' somehow compensated by 'a lot of powder.' This correlation argument contains a
number of basic flaws, i.e., unproven assumptions (1) that a small gland produces insufficient oil
and (2) that the oil and powder down are used more or less interchangeably for waterproofing
feathers especially in land birds. It should be emphasized that virtually nothing is known about
the quantity, rate of production, or rate of secretion of uropygial oils. Without experimental
documentation, it cannot be assumed that small glands have any reduction in rate or quantity of
secretion, whether the bird has powder down or not.

It is true that birds known to produce a significant amount of powder down (Gadow 1891,
Chandler 1916, Thomson 1964, Jacob 1978, Baumel et al. 1979) tend to have relatively small
glands: (gland weights as a percent of body weight) Ardeidae (N = 20, x = 0-29%), Psittacidae
(N = 41, x = 0- \Q%,fide Jacob &Ziswiler 1982 forzoo birds), Ramphastidae(N = 3,x = 0-12%), and
Tinamidae (N= 1, 0-\8%fide Jacob & Ziswiler 1982). Relative gland weights are unavailable for
other species that produce powder down: Podargidae, Cotingidae, Leptosomatidae, Artamidae,
Ptilonorhynchidae, and others. The Mesoenatidae, which have five pairs of powder down patches
(Olson 1 978), lack a gland. According to Schuz (1 927), 'powder downs are lacking, or nearly so, in
ratite birds;' among the ratites, only Apteryx possesses a gland as an adult. Powder downs,
produced in various amounts are known from a wide variety of other birds including Columbidae,
Rhynochetidae, Eurypigidae, Podargidae, Otididae, and Accipitridae. This body of circumstantial
evidence lends some support to the view that birds with well-developed powder down production
have reduced (or no) uropygial glands.

A cause-and-effect functional relationship remains unproven, however. Furthermore, the func-
tion of powder remains conjectural probably because several types of powder (down) are known:
as a waterproof dressing (Bartlett 1861), preserving feathers (Welty 1962), and cleaning feathers
(Thomson 1964). Although the powder has a nonwettable property, it is composed largely of
keratin, so its functional equivalence to uropygial oils must await experimental proof (see also
Lucas & Stettenheim 1972).

All this information strongly argues that more research on functions of glandular secretions is
badly needed before physiological generalizations can be asserted. Particularly open to question is
the function of secretions in terrestrial birds and in bird taxa containing some glandless members
(e.g., doves, parrots).

Future studies

Several biological aspects of uropygial glands merit further investigations because results
therefrom could help to explain some of the morphological variations identified in the present
monograph. Johansson's studies (1927) indicated a strong genetic component in the inheritance
of uropygial glands in Columba livia, as is also suggested from the fact that certain varieties
of this pigeon lack glands (Darwin 1900, Levi, 1941, Goodwin 1983). Hutt (1949) reported that
mutation of a dominant gene in chickens causes bifurcation of the gland's papilla and that
most heterozygotes have no uropygial gland at maturity. The rumpless chickens of Waterton
(1836a) presumably had no glands and might have been genetic mutants. Inheritance of double
gland papillae were discussed by Kessel (1945) for domestic fowl. Apparently these are the only
investigations pertaining to the inheritance of uropygial glands, and further genetic studies might
reveal biological relationships to the absence of glands in taxa of wild birds.

Another aspect in need of experimental studies is the physiology of gland production, secretion,
and its relationship to preening. Nothing is known about either the quantity or rate of secretion
of uropygial oils. Some information is available on histology, vascular supply and innervation
(Kossman 1871, Paris 1913, Kanwar 1961). Many additional questions are unanswered, however:
(1) do birds with large glands (e.g., waterbirds) produce more oil than birds with small glands (e.g.,
landbirds); (2) is gland oil production stimulated, and at what rate, by physical manipulation with
the bird's bill; (3) is there either seasonal or daily variation in the quantity or rate of secretion?

254 DAVID W. JOHNSTON

Despite some papers that address preening activities in birds (e.g., in penguins, Bekoftet al. 1979,
and references therein), little is known about the relationship between gland secretion and types or
rates of preening activities that involve this gland. A case in point was the radical statement by
Gurney (1913) that Sula bassana does not use its gland in preening, a statement since disputed by
Nelson (1978).

As indicated in the previous account, attention should be given to functional attributes of
glandular secretions, especially in terrestrial birds.

Little is known about any relationship between gland shape and the underlying muscles and
rectrices. Future research could focus on explaining the several different gland shapes identified in
this study especially as those shapes might be related to muscle differences or to placement of the
rectrices.

A final research need is in embryology, especially post-hatching development as it might relate to
function in certain species (a review of the gland's embryology is found in Jacob and Ziswiler 1982).
Prior to the report by Pycraft in 1900, it was believed that ratites, with the exception of Apteryx,
lack uropygial glands. He found, however, that in both Dromaius novaehollandiae and Rhea
americana, a gland exists in both the embryo and nestling, but is absent in the adult. (Those
conditions have been verified in the present study). Apparently no one has examined those
embryonic glands histologically or functionally. Although no glands have been found in any age
group ofStruthio or Casuarius, might some trace or anlage be found by an embryonic-histological
study? In other adult birds lacking glands (e.g., Mesoenatidae, Otidae) is there any early
embryological development of a gland?

Acknowledgments

Several organizations provided travel funds from 1970-1986 for this study — National Science
Foundation (DEB-79-03687, BSR-82- 14603, suppl. to K. R. McKaye), American Museum of
Natural History (Frank M. Chapman Memorial Fund), University of Florida Department of
Zoology, and George Mason University Graduate School. Museums visited for examination of
spirit collections were: National Museum of Natural History (R. L. Zusi), British Museum
(Natural History) in Tring (P. J. K. Burton, G. Cowles, I. C. J. Galbraith), American Museum of
Natural History (L. L. Short, A. V. Andors), Field Museum of Natural History (J. W. Fitzpatrick),
Museum of Comparative Zoology (R. A. Paynter, Jr.), University of Michigan Museum of
Zoology (R. W. Storer), Peabody Museum of Natural History (C. G. Sibley), Manomet Bird
Observatory (K. S. Anderson, T. L. Lloyd-Evans), Charles R. Conner Zoology Museum (R. E.
Johnson), North Carolina State Museum of Natural History (D. S. Lee), University of Georgia
Museum of Zoology (J. E. Cadle), Stovall Museum of Science and Technology (G. D. Schnell,

D. W. Mock), and California Academy of Sciences (L. F. Baptista). Some specimens were
provided by D. G. Ainley, O. L. Austin, Jr., C. Dau, J. B. Faro, D. Forrester, M. S. Foster, M. J.
Gilroy, R. Heath, H. W. Kale, II, B. Kessel, A. M. Lindahl, C. D. Marti, B. W. Miller, D. G.
Matthiesen, S. A. Nesbitt, D. S. Peters, W. B. Robertson, Jr., R. W. Schreiber, R. Spenser, and G.

E. Woolfenden. Additional specimens and data on glands came from the Tall Timbers Research
Station (R. L. Crawford), Busch Gardens (Fla.), Sea World (Fla.), Louisiana State University
Museum of Zoology (J. P. O'Neill), and Santa Fe Community College Teaching Zoo (Fla.). The
microscopic study of feathers was enhanced by the labours of Ray Bienert and by discussions
with Roxie Laybourne. Belinda Gillies supplied information on Atrichornis clamosus, and J. A.
Bartle assisted in gathering information on the Acanthisittidae. Julian J. Baumel offered advice
on several anatomical details. Caroline Carboni-Vetter kindly translated articles in French, and
Elisabeth Egghart, articles in German. Through the years countless valuable specimens and much
encouragement came from Pierce Brodkorb and Tony Gilyard. Anne Vale was generous with her
time and expertise in locating references in the Rothschild Library at the British Museum of
Natural History in Tring. Earlier drafts were read by Storrs Olson, Peter Stettenheim, Vincinz
Ziswiler, and Richard Zusi; their suggestions helped to improve the manuscript. Anne O'Malley
spent many hours typing and retyping several drafts of the entire manuscript.

AVIAN UROPYGIAL GLAND 255

This investigation has also benefitted from numerous discussions with Storrs Olson especially on
matters dealing with systematics and pertinent references. I am profoundly indebted to Richard L.
Zusi who, over many years, freely offered his time, advice, and suggestions on virtually every aspect
of the study. Finally, the study would have been incomplete without the superb illustrations
meticulously executed by Esta L. Johnston.

Summary

The primary goal of this study has been to assemble a complete analysis of the uropygial gland's
morphology in representatives of all bird families and subfamilies. Particular attention has been
given to correcting erroneous information about glands in the existing literature. Morphological
data are included from many avian taxa not previously reported.

The largest glands, relative to body weight, are found in birds that swim, dive or rest on water.
Progressively smaller glands occur in birds that walk in water, those that habitually only fly over
water, and lastly, terrestrial species. Glands are now known to be absent in the Struthionidae (all
age groups), Rheidae (adults only), Casuariidae (all age groups), Dromaiidae (adults only),
Mesoenatidae, Otidae, Columbidae (9 genera, 28 species), Psittacidae (6 genera, 31 species),
Podargus spp., and Picidae (1 genus, 4 species). The absence of glands in these taxa is believed to be
a secondary and independent loss. Because only 17 percent of flightless species lack a gland, this
study revealed no significant correlation between gland absence and a flightless condition.

Many nonpasserine taxa possess tufted glands, whereas others have manifestly naked (non-
tufted) glands. Apparently naked glands of others, e.g. Strigidae, actually might bear minute
feathers. All species of the Passeriformes have naked glands. Feathers attached to the uropygial
gland are more numerous in waterbirds than landbirds and are of two principal types, modified
down or modified semiplumes. Only circumstantial evidence was found to support the
oft-expressed hypothesis that power down is a substitute for gland secretions in glandless species.

The present complete study provides sufficient morphological characteristics for their
consideration in avian taxonomic schemes.

Future investigations should focus on gland function, especially in terrestrial birds, the quantity
and rate of secretion of uropygial oils, relationships between gland production and preening
activities, details of feather structure, genetics, and embryological development.

Literature cited

Arnall, L. & I. F. Keymer. 1975. Bird diseases. Neptune City, New Jersey: T.F.H. Publications.

Audubon, J. J. 1 829. Notes on the bird of Washington (Falco Washingtoniand) or Great American Sea Eagle.

Magazine of Natural History 1: 1 15-120.

Austin, O. L. Jr. 1961 . Birds of the world. New York: Golden Press.
Balthazart, J. & E. Schoffeniels. 1979. Pheromones are involved in the control of sexual behaviour in birds.

Naturwissenschaften 66: 55-56.
Bartlett, A. D. 1861. On the affinities of Balaeniceps. Proceedings of the Zoological Society of London,

pp. 131-134.
Baumel, J. J., A. S. King, A. M. Lucas, J. E. Breazile, H. E. Evans (eds.). 1979. Nomina anatomica avium.

New York: Academic Press.
Beddard, F. E. 1898. The structure and classification of birds. London: Longmans, Green & Co.

— 1899. Notes on the anatomy of the genus Apteryx. Novitates Zoologicae 6: 386—402.
Bekoff, M., D. G. Ainley & A. Bekoff. 1979. The ontogeny and organization of comfort behavior in Adelie

Penguins. Wilson Bulletin 91: 255-270.
Burton, E. 1822. Observations on the natural history and anatomy of the Pelecanus Aquilus of Linnaeus.

Transactions of the Linnaean Society of London, 13: 1-1 1.
Burton, P. J. K. 1984. Anatomy and evolution of the feeding apparatus in the avian order Coraciiformes and

Piciformes. Bulletin of the British Museum (Natural History) (Zoology) 47: 331^43.

256 DAVID W. JOHNSTON

Chandler, A. C. 1916. A study of the structure of feathers, with reference to their taxonomic significance.

University of California Publications in Zoology 13: 243-446.
Clark, G. A., Jr. 1964. Ontogeny and evolution in the Megapodes (Aves: Galliformes). Postilla No. 78: 1-37.

— 1969. Spread-wing postures in Pelecaniformes, Ciconiiformes, and Falconiformes. Auk 86: 136-139.
Cottam, P. A. 1957. The pelecaniform characters of the skeleton of the Shoe-bill Stork, Balaeniceps rex.

Bulletin of the British Museum (Natural History) (Zoology) 5(3): 49-72.
Coues, E. 1890. Handbook of field and general ornithology. London: Macmillan & Co.
Cracraft J. 1981. Toward a phylogenetic classification of the recent birds of the world (Aves). Auk 98:

681-714.

- 1985. Monophyly and phylogenetic relationships of the Pelecaniforms: a numerical cladistic analysis.
Auk 102: 834-853.

Crisp, E. 1860. On the structure, relative size, and use of the tail-gland in birds. Proceedings of the Zoological
Society of London, Part 28: 254-260.

— 1862. Exhibition of the enlarged tail-glands of two domestic hens. Proceedings of the Zoological Society
of London, p. 219.

Cuvier, G. 1799-1805. Paris: Lecons d'Anatomie Comparee.

Darwin, C. 1900. The variation of animals and plants under domestication. Vol. I. New York: D. Appleton

and Co.

Elder, W. H. 1954. The oil gland of birds. Wilson Bulletin 66: 6-31.
Fisher, H. I. 1943. The pterylosis of the King Vulture. Condor 45: 69-73.
Forbes, W. A. 1 882. On some points in the anatomy of the todies (Todidae), and on the affinities of that group.

Proceedings of the Zoological Society of London, pp. 442-450.

Fry, C. H. 1980. The evolutionary biology of kingfishers (Alcedinidae). Living Bird, 18: 1 13-160.
Gadow, H. 1891. Notes on the structure of Pedionomus torquatus, with regard to its systematic position.

Records of the Australian Museum, 1: 205-21 1 .

— 1893. Vogel. I. Anatomischer Theil in Bronn's Klassen undOrdnungen des Thiereichs, 6: 1-1008, Leipzig.
Garrod, A. H. 1873. On some points in the anatomy of Steatornis. Proceedings of the Zoological Society of

London, pp. 526-535.

— 1874a. On some points in the anatomy of the Columbae. Proceedings of the Zoological Society of London,
pp. 249-259.

- 18746. On some points in the anatomy of the parrots which bear on the classification of the suborder.
Proceedings of the Zoological Society of London, pp. 586-598.

- 1876. Notes on the anatomy of the colics (Colies). Proceedings of the Zoological Society of London,
pp. 41 6-420.

1878. On the systematic position of the Momotidae. Proceedings of the Zoological Society of London,

pp. 100-102.
Giebel, C. 1857. Zur Anatomic des Wiedehopfs, Upupa epops, nach Chr. L. Nitzsch's Untersuchungen

mitgetheilt. (Zeitschrift Gesammt, Naturwissenschaften: 236-244).

Goodwin, D. 1983. Pigeons and doves of the world. 3rd ed. New York; Cornell Univ. Press, Ithaca.
Granvik, H. 1913. Untersuchungen uber glandula uropygii. Arkivfoer Zoologi, 8: 19pp.
Grasse, P. P. 1950. Oiseaux. Traite de zoologie 15: 285-289.
Greenway, J. C., Jr. 1958. Extinct and vanishing birds of the world. New York. American Committee for

International Wild Life Protection, Special Publication 13.

Groebbels, F. 1932. Der Vogel. Bau, Funktion, Lebenserscheinung. Part 1. Berlin: Borntrager.
Gurney, J. H. 1913. Thegannet. London: Witherby & Co.

Hancock, J. & J. Kushlan. 1984. The herons handbook. New York: Harper and Row.
Hou, H. C. 1928. Studies on glandula uropygialis of birds. American Journal of Physiology 85: 380.
Hussey, A. 1860. What is the use of the oil-gland at the base of the tail of birds. Zoologist 18: 7049-7050.
Hutt, F. B. 1949. Genetics of the fowl. New York: McGraw-Hill.
Jackson, F. J. 1938. The birds of Kenya Colony and the Uganda Protectorate. Vol. II. London: Gurney and

Jackson.
Jacob, J. 1978. Uropygial gland secretions and feather waxes. Chap. 6 in Chemical Zoology (A. H. Brush, ed.),

Vol. X. New York: Academic Press.
Jacob, J. & V. Ziswiler. 1982. The uropygial gland. Chap. 4 in Avian biology, D. S. Earner, J. R. King, K. C.

Parkes (eds), Vol. VI. New York: Academic Press.
Johansson, I. 1927. Studies on inheritance in pigeons. VI. Number of tail-feathers and uropygial glands.

Genetics 12: 93-107.
Johnston, D. W. 1979. The uropygial gland of the Sooty Tern. Condor 81: 430-432.

AVIAN UROPYGIAL GLAND 257

Kanwar, K. C. 1 96 1 . Morphological and histochemical studies on the uropygial glands of pigeon and domestic

fowl. Cytologia26: 124-136.
Kar, A. R. 1947. The hormonal influence in the normal functioning of the uropygial gland in the fowl.

Anatomical Record 99: 75-89.

Kennedy, R. J. 1971. Preen gland weights. Ibis 113: 369-372.
Kessel, E. L. 1945. Inheritance of cleft and double uropygial gland papillae in domestic fowl. Wasmann

Collector 6: 84-87.
King, A. S. & J. McLelland. 1984. Birds, their structure and function, 2nd edition. Philadelphia, Pennsylvania:

Bailliere Tindall.

Kossmann, R. 1871. Ueber Talgdrusen der Vogel. Zeitschrift fur Wissenschaftliche Zoologie: 568-599.
Landouer, W. & L. C. Dunn. 1925. Two types of rumplessness in the domestic fowls. Journal of Heredity 16,

pp. 153-160.

Law, J. E. 1929. The function of the oil-gland. Condor 31: 148-156.
Levi, W. M. 1941. The pigeon. Columbia, South Carolina: R. L. Bryan Co.
Ligon, J. D. 1967. Relationships of the cathartid vultures. Occasional Papers of the Museum of Zoology

(University of Michigan) No. 651, pp 1-26.
Lucas, A. M. & P. R. Stettenheim. 1972. Uropygial gland, in Avian anatomy, Part II. United States

Department of Agriculture , Agricultural Handbook 362: 613-626.
Lunghetti, B. 1906. Konformation, Struktur und Entwicklung der Burzeldruse bei verschiedenen Vogelarten.

Archiffur Mikroskopische Anatomic 69: 264-321.

Macgillivray, W. 1837. A history of British birds. Vol. 1. London: Scott, Webster, and Geary.
Mackworth-Praed, C. W. & C. H. B. Grant. 1970. Birds of West Central and Western Africa. African

Handbook Birds, Series HI, Vol. 1. London: Longman Group Ltd.
Matthews, H. S. R. 1861. Oil gland in birds. Zoologist 19: 7439.
Miller, W. D. 1915. Notes on ptilosis, with special reference to the feathering of the wing. Bulletin of the

American Museum of Natural History 34: 129-140.

— 1 924. Further notes on ptilosis. Bulletin of the American Museum of Natural History 50, Art. V: 305-33 1 .
Mlikovsky, J. 1982. Evolution of flightlessness in birds: an ecological approach. Evolution and Environment:

693-730.

Morris, F. O. 1 836a. Touching the question whether birds oil their plumage. Magazine of Natural History 9:
159-164.

— 18366. The question of the office of the gland upon the rump of birds. Magazine of Natural History 9:
269-271.

1836c. The question of the office of the gland upon the rump of birds. Magazine of Natural History 9:

434-437.
Murphy, R. C. 1936. Oceanic birds of South America. Vol. 2. New York: American Museum of Natural

History.

Nelson, B. 1978. The gannet. Berkhamsted: T. and A. D. Poyser.

Nelson, J. B. 1975. Breeding biology of frigatebirds — a comparative review. Living Bird 14: 1 13-155.
Newton, A. 1893-1896. A dictionary of birds. London: Adam and Charles Black.
Nitzsch, C. L. 1840. System der Pterylographie. Burmeister Halle.

- 1867. Nitzsch's Pterylography. Translated into English by W. S. Dallas and edited by P. L. Sclater.
London: Ray Society.

Olson, S. L. 1973a. Evolution of the rails of the South Atlantic Islands (Aves: Rallidae). Smithsonian
Contributions to Zoology, 152: 1-53.

- 19736. A classification of the Rallidae. Wilson Bulletin 85: 381^416.

- 1979. Multiple origins of the Ciconiiformes. Proceedings 1978 Conference of the Colonial Waterbird
Group: 165-170.

— 1983. Evidence for a polyphyletic origin of the Piciformes. Auk 100: 126-133.

— & A. Feduccia. 19800. Presbyornis and the origin of the Anseriformes (Aves: Charadriomorphae).
Smithsonian Contributions to Zoology, 323: 1-24.

- & - - 19806. Relationships and evolution of Flamingos (Aves: Phoenicopteridae). Smithsonian
Contributions to Zoology, 316: 1-73.

& D. W. Steadman. 1981 . The relationships of the Pedionomidae (Aves: Charadriiformes). Smithsonian

Contributions to Zoology, 337: 1-25.
Owen, R. 1866. On the anatomy of vertebrates. Vol. 2. London: Longmans, Green, and Co.
Paris, P. 1913. Recherches sur la Glande Uropygienne des Oiseaux. Archives de Zoologie Experimentale et

Generate 53: 139-276.

258 DAVID W. JOHNSTON

Peters, J. L., et al. 1931-1986. Check-list of the birds of the world. Vols. 1-15. Cambridge, Massachussetts:

Harvard University Press.

Pettingill, O. S., Jr. 1985. Ornithology in laboratory and field. 5th ed. Orlando, Florida: Academic Press.
Pycraft, W. P. 1900. On the morphology and phylogeny of the Paleognathae (Ratitae and Crypturi) and

Neognathae (Carinatae). Transactions of the Zoological Society of London 15: 149-290.

— 1910. Animal life. Vol. 2. A history of birds. London: Methuen and Co.
Rijke, A. M. 1970. The phylogenetic development of water repellency in water bird families. Ostrich

Supplement 8: 67-76.
Ripley, S. D. 1976. Rails of the world. American Scientist 64: 628-635.

- & B. M. Beehler. 1985. Rails of the world, a compilation of new information, 1975-1983 (Aves:

Rallidae). Smithsonian Contributions to Zoology, 417: 1-28.
Rutschke, E. 1960. Untersuchungen iiber Wasserfestigkeit und Struktur des Gefieders von Schwimmvogeln.

Zoologische Jahresbuecher, Abteilungfur Systematiks 87: 441-506.
Schumacher, S. 1919. Der Burzeldocht. Anatomischer Anzeiger 52: 291-301.

Schuz, E. 1927. Beitrag zur Kenntnis der Puderbildung bei den Vogeln. Journal fur Ornithologie 75: 86-224.
Short, L. L. 1982. Woodpeckers of the world. Delaware Museum of Natural History, Greenville, Delaware

Monograph Series, No. 4.

Shortt, T. M. 1977. Wild birds of the Americas. Boston, Massachusetts: Houghton Mifflin Co.
Sibley, C. G. 1967. Proteins: history books of evolution. Discovery 3: 5-20.

— , K. W. Corbin & J. E. Ahlquist. 1968. The relationships of the seed-snipe (Thinocoridae) as indicated by

their egg white proteins and hemoglobins. Banner Zoologische Beitrage, 3/4: 235-248.

& J. H. Haavie. 1969. The relationships of the flamingos as indicted by the egg-white proteins and

hemoglobins. Condor 71: 155-179.
Spearman, R. I. C. 1 97 1 . Integumentary system. In Physiology and biochemistry of the domestic fowl, D. J. Bell

and B. M. Freeman (eds.) Vol. 2: 609. New York: Academic Press.
Stettenheim, P. 1972. The integument of birds. Chap. 1 in Avian biology. Vol. II (D. S. Farner and J. R. King,

eds.). New York: Academic Press.

Stubbs, F. J. 1910. The mechanism of plumage in water birds. The Zoologist, 4th series, 14: 201-206.
Thomson, A. L. 1964. A new dictionary of birds. New York: McGraw-Hill.
Turcek, F. J. 1966. On plumage quantity in birds. Ekologia Polska Ser A 14: 1-7.
Tyson, E. 1 863. Anatomy of the Mexico musk-hog. Philosophical Transactions of the Royal Society of London,

Ser. A 13: 359-385.

Van Tyne, J. & A. J. Berger. 1976. Fundamentals of ornithology. 2nd edition. New York: Wiley & Sons.
Verheyen, R. 1955a. Analyse du Potential Morphologique et Considerations sur la Systematique des

Coraciiformes (Wetmore 1934). Bulletin. Institut Royal des Sciences Naturellesde Belgique, Vol. 31, No. 31:

1-19.

- 19556. Analyse du Potential Morphologique et Considerations sur la Systematique des Coraciiformes
(Wetmore 1934). Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 31, No. 92: 1-16.

- 1955c. Analyse du Potential Morphologique et Considerations sur la Systematique des Coraciiformes
(Wetmore 1934). Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 31, No. 94: 1-16.

— \955d. Contribution a la Systematique des Piciformes Basee sur 1' Anatomic Comparee. Bulletin. Institut
Royal des Sciences Naturelles de Belgique, Vol. 31, No. 51: 1-19.

- 1956o. Les Striges, les Trogones, et les Caprimulgi dans la Systematique Moderne. Bulletin. Institut
Royal des Sciences Naturelles de Belgique, Vol. 32, No. 3: 1-31 .

- 19566. Contribution a 1' Anatomic et a la Systematique des Touracos (Musophagi) et des Coucous
(Cuculiformes). Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 32, No. 23: 1-28.

- 1956f. Note Systematique sur Opisthocomus hoazin (St.-Muller). Bulletin. Institut Royal des Sciences
Naturelles de Belgique, Vol. 32, No. 32: 1-8.

- \956d. Contribution a T Anatomic et a la Systematique des Galliformes. Bulletin. Institut Royal des
Sciences Naturelles de Belgique, Vol. 32, No. 42: 1-24.

- \956e. Note sur F Anatomic et la Classification des Coliiformes. Bulletin. Institut Royal des Sciences
Naturelles de Belgique, Vol. 32: No. 47: 1-7.

- 1956/. Analyse du Potentiel Morphologique et Projet d'une Nouvelle Classification des Psittaciformes.
Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 32, No. 55: 1-54.

- 1957a. Analyse du Potentiel Morphologique et Projet de Classification des Columbiformes (Wetmore
1934). Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 33, No. 3: 1-42.

- 19576. Contribution au Demembrement de 1'Ordo Artificiel des Gruiformes (Peters 1934). Part I. Les

Ralliformes. Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 33, No. 21: 1-44.

AVIAN UROPYGIAL GLAND 259

— 1957c. Contribution au Demembrement de 1'Ordo Artificial des Gruiformes (Peters 1934). Part II. Les
Cariamiformes. Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 33, No. 39; 1-7.

— \957d. Contribution au Demembrement de 1'Ordo Artificiel des Gruiformes (Peters 1934). Part III. Les
Jacaniformes. Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 33, No. 48: 1-19.

— 1958a. Contribution au Demembrement de 1'Ordo Artificiel des Gruiformes (Peters 1934). Part IV. Les
Turniciformes. Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 34, No. 2: 1-18.

- 19586. Analyse du Potentiel Morphologique et Projet d'une Nouvelle Classification des Charadrii-
formes. Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 34, No. 18: 1-35.

— 1958c . Note sur la Classification des Procellariiformes (Tubinares). Bulletin. Institut Royal des Sciences
Naturelles de Belgique, Vol. 34, No. 30: 1-22.

— \95Sd. Contribution a la Systematique des Alciformes. Bulletin. Institut Royal des Sciences Naturelles de
Belgique, Vol. 34, No. 45: 1-15.

— 1959a. Note sur la Systematique de Base des Lariformes. Bulletin. Institut Royal des Sciences Naturelles
de Belgique, Vol. 35, No. 9: 1-16.

- 19596. Contribution a 1' Anatomic et a la Systematique de Base des Ciconiiformes (Parker 1868).
Bulletin. Institut Royal des Sciences Naturelles de Belgique, Vol. 35, No. 24: 1-34.

— 1959c. Revision de la Systematique des Falconiformes. Bulletin. Institut Royal des Science Naturelles de
Belgique, Vol. 35, No. 37: 1-51.

— \959d. Les Plongeons (Gaviae) et les Grebes (Podicipitides) dans les Systems de Classification. Bulletin.
Institut Royal des Sciences Naturelles de Belgique, Vol. 35, No. 44: 1-12.

— 1960a. Les Tinamous dans les Systems Ornithologiques. Bulletin. Institut Royal des Sciences Naturelles
de Belgique, Vol. 36, No. 1:1-11.

19606. Les Pelecaniformes et le Paille-en-queue (Phaethon). Bulletin. Institut Royal des Sciences

Naturelles de Belgique, Vol. 36, No. 25: 1-18.
Voitkevich, A. A. 1966. The feathers and plumage of birds. London: Sidgwick and Jackson.
von Jacob, J. & H. Hoerschelmann. 1985. [Classification of flamingos (Phoenicopteriformes) by comparative

analysis of their uropygial gland secretions.]. Zeitschrift fur Systematik Evolution 23: 49-58.
Waterton, C. 1 832. On birds using oil from glands 'for the purpose of lubricating the surface of their plumage.'
Magazine of Natural History 5: 412—415.

- 1836tf. On the Robin and Dipper, and rumped, and rumpless birds. Magazine of Natural History 9:
158-159.

18366. The question of the office of the gland upon the rump of birds. Magazine of Natural History 9:

266-269.

1860. The oil-gland in birds. Zoologist 18: 7103-7104.

Weller, M. W. 1980. The island waterfowl. Ames, Iowa: Iowa State University Press.
Welty, J. C. 1962. The life of birds. Philadelphia, Pennsylvania: W. B. Saunders Co.
Willughby, F. 1678. The ornithology of Francis Willughby. Translated by J. Ray. London: A.C. for John

Martyn.
Wood, C. A. & F. M. Fyfe. 1943. The art of falconry, being the De Arte Venandi Cum Avibus of Frederick II of

Hohenstaufen. California: Stanford Unversity Press.
Zusi, R. L. 1985. Muscles of the neck, trunk and tail in the Noisy Scrub-bird, Atrichornis clamosus, and

Superb Lyrebird, Menura novaehollandiae (Passeriformes: Atrichornithidae and Menuridae). Records of

the Australian Museum 37: 229-242.

Manuscript accepted for publication 29 February 1988

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 Afro tropical 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. 0565 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 Haplocaulm 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. Billett, A. M. Clark & G. L. J. Paterson

A morphological atlas of the avian Uropygial gland. By David W. Johnston
Miscellanea

Printed in Great Britain by Henry Ling Ltd . at the Dorset Press. Dorchester. Dorset

25NOV1938

PRESENTED
GENERAL LIBRAR

Bulletin of the

British Museum (Natural History)

Miscellanea

Zoology series Vol 54 No 6

24 November 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. V ,

Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
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,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:

Publications Sales,

British Museum (Natural History),
Cromwell Road,

London SW7 5BD,
England.

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 : MrJ.F. Peake
Editor of Bulletin : DrC.R. Curds
Assistant Editor : Mr C. G. Ogden

ISSN 0 565 05042 7

ISBN 0007 1498 Zoology series

Vol54 No 6 pp 26 1-307
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 24 November 1 988

Miscellanea

Contents

A review of the copepod endoparasites of brittle stars (Ophiuroida). By G. A. Boxshall .
A new genus of tantulocaridan (Crustacea: Tantulocarida) parasitic on a harpacticoid

copepod from Tasmania. By G. A. Boxshall

Unusual ascothoracid nauplii from the Red Sea. By G. A. Boxshall & R. Bottger-Schnack
New nicothoid copepods (Copepoda: Siphonostomatoida) from an amphipod and from

deep-sea isopods. By G. A. Boxshall & K. Harrison

A new genus of Lichomolgidae (Copepoda: Poecilostomatoida) associated with a

phoronid in Hong Kong. By G. A. Boxshall & A. G. Humes

Page
261

271
275

285
301

BRITISH MUSEU
(NATURAL HISTORY)

25NOVJ988

PRESENTED
GENERAL LIBRAI

Geoffrey A. Boxshall

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

Summary

A new family, the Chordeumiidae, is proposed for six genera of copepods that live as endoparasites of brittle
stars. The status of these genera, Chordeumium, Arthrochordeumium, Lernaeosaccus , Ophioicodes, Ophioika
and Parachordeumium is reviewed. The genera Ophioithys and Amphiurophilus are recognised as subjective
synonyms of Parachordeumium. The sole species of Lernaeosaccus is reinterpreted and redescribed.
Examination of the holotype of L. ophiacanthae revealed that it was originally described upside down. Codoba
discoveryi is also redescribed from the types but cannot be placed in any of the existing families of the
Siphonostomatoida at present.

Introduction

In his review of copepods associated with invertebrate hosts Gotto (1979) listed eight genera of
endoparasites that utilise brittle stars as hosts. None of these has formally been placed in a family
and even their ordinal placement is uncertain. There are obvious taxonomic problems concerning
these genera which need to be resolved before their phylogenetic relationships with other copepods
can be understood. Most of the species contained in these 8 genera have highly transformed
females with bizarre body morphology in the adult and few, if any, recognisable limbs. How-
ever, in some genera either males or developmental stages are known and these provide more
taxonomically useful information. The monotypic genus Codoba Heegaard is much less modified,
still retaining a more-or-less cyclopiform facies in the adult female (Heegaard, 1951). Codoba is
redescribed, based on the type material, and the taxonomic status and phylogenetic relationships
of the other genera endoparasitic in ophiuroids are reassessed. The morphology of the monotypic
genus Lernaeosaccus Heegaard is reinterpreted and a new description and diagnosis of the genus is
provided.

Descriptions
Genus CODOBA Heegaard, 1951

DIAGNOSIS. Adult female prosome 4-segmented, slightly swollen; urosome 5-segmented. Caudal
rami distinct, bearing 6 setae. Antennule 20-segmented, lacking aesthetascs. Antenna reduced to
unarmed process. Mouth tube well developed. Mandible stylet-like, without teeth; palp absent.
Maxillule bilobed, inner lobe with 1, outer with 2 setae. Maxilla with distal recurved claw.
Maxilliped 3-segmented. Legs 1-4 biramous, rami medially directed, intercoxal sclerites absent.
Leg 5 lacking. Male unknown.

TYPE SPECIES. Co doba discoveryi Heegaard, 1951 bymonotypy.

Codoba discoveryi Heegaard, 1951

ADULT FEMALE. Body (Fig. 1 A) slightly transformed cyclopiform, but with distinct segmentation.
Prosome swollen, comprising cephalothorax incorporating first pedigerous somite, and 3 free
pedigerous somites. Total body length of figured syntype 1-97 mm. Each free pedigerous somite
with a pair of posterolaterally directed epimeral processes. Urosome short (Fig. IB), comprising

Bull. Br. Mus. not. Hist. (Zool.) 54(6): 261-270 Issued 24 November 1988

262

G. A. BOXSHALL

COPEPOD ENDOPARASITES OF BRITTLE STARS 263

somite of leg 5 (95 x 398 urn), the genital complex (130xl30um), 2 postgenital somites
(40x235(im, 25xl90um) and the anal somite (115xl50um). Genital apertures unarmed;
located ventrolaterally on genital complex. Anal somite with row of spinules along posterior
margin. Caudal rami (Fig. 1 B) about 1 -5 times longer than wide (80 x 53 um); armed with a lateral
seta about at midlength, a naked dorsal seta and 4 distal margin setae, 3 of which are pinnate.

Antennule (Fig. 1C) 20-segmented; lengths of segments measured along posterior margin 13, 9,
5, 10, 11, 10, 13, 12, 12,9,9,9, 18, 15, 14, 11, 15, 14, 11 and 61 um; armature elements as follows:
1—3, II— 1, III— 1, IV— 2, V— 2, VI— 1, VII— 1, VIII— 2, IX— 2, X— 2, XI— 1, XII— 1, XIII— 2,
XIV— 1, XV— 1, XVI— 1, XVII— 1, XVIII— 1, XIX— 1, XX— 12. No aesthetascs present. The
size and number of armature elements on the apical segment indicate that it is derived by fusion of 2
or more segments. Antenna vestigial; reduced to small unarmed process located between base of
antennule and mouth tube (Fig. 2F).

Mouth tube (Fig. ID) well developed, formed from labrum and labium. Mandible (Fig. IE)
stylet-like, unarmed; palp lacking. Maxillule bilobed (Fig. 1 F); inner lobe represented by single
hirsute seta; outer lobe short, armed with 2 hirsute setae. Maxilla (Fig. 1G) comprising unarmed
syncoxa and distal basis. Basis drawn out into recurved claw, armed distally with patches of
denticles as figured. Maxilliped (Fig. 2A) 3-segmented; basal segment armed with a row of
spinules; second segment elongate, armed with a single seta and several rows of spinules; third
segment bearing a tiny seta proximally on outer margin, a small inner seta and 2 unequal apical
spines. Spines dentate, 51 um and 41 jim in length. Fine spinules present around apex.

Swimming legs 1 to 4 biramous, positioned on ventral surface of somite with rami directed
medially (Fig. IB); intercoxal bars (sclerites) absent. Leg 1 (Fig. 2B) with unarmed protopod;
exopod unsegmented, bearing 4 pinnate setae along outer margin, 2 at apex and 2 distally on inner
margin; endopod indistinctly 2-segmented, bearing 4 pinnate setae along inner margin, 2 at apex
and 1 distally on outer margin. Leg 2 (Fig. 2C) with unarmed protopod. Exopod 2-segmented; first
segment with outer pinnate seta, second with 4 naked setae on outer margin, 1 at apex and 2 on
inner margin. Endopod unsegmented; with 5 pinnate setae along inner margin and 2 at apex. Leg 3
(Fig. 2D) with unarmed protopod; both rami indistinctly 2-segmented. Exopod with outer seta on
first segment, second with 2 outer and 2 apical setae, and a row of spinules medially; endopod with
2 inner margin setae on first segment; second with 1 inner, 1 outer and 2 apical setae. Leg 4 (Fig. 2E)
with unarmed protopod; exopod with single apical seta; endopod with 2 setae on inner margin and
3-branched spinous projection apically. Leg 5 lacking.

Egg sacs lacking, eggs loose inside capsule with adult female. Capsule of specimen 1987.244
containing 44 eggs; eggs large, lecithotrophic, diameter approximately 190 um.

MATERIAL EXAMINED. Syntypes: Dissected syntype (BM(NH) No. 1987.243) collected from
Ophiura meridionalis (Lyman) at Discovery Stn 123, in 230-250 m off South Georgia, 15.12.1926.
Intact syntype (BM(NH) No. 1987.244) from O. meridionalis at Discovery Stn 156, in 236m off
South Georgia, 20.1.1927. Third syntype an empty capsule (BM(NH) No. 1987.245) collected
from Amphiura belgicae Koehler at Discovery Stn 1 60, in 1 77 m off South Georgia, 7.2. 1 927. These
are the data on the labels with the specimens and do not correspond in some details with those data
published by Heegaard (1951).

REMARKS. When Heegaard (1951) erected the genus Codoba he indicated that new families would
probably have to be established to accommodate the copepods parasitic on echinoderms but felt
that it would be premature to do so in view of the lack of knowledge of the group. The high number
of antennulary segments in Codoba is a plesiomorphic character and is typical of the family
Asterocheridae but the genus cannot be referred to the Asterocheridae as it lacks a mandibular
palp and an aesthetasc on either the penultimate or antepenultimate segment of the antennule.

Fig. 1 Codoba discoveryi Heegaard, 1951. Adult female syntype. A, dorsal; B, Fourth pedigerous
somite and urosome, ventral; C, Antennule, ventral; D, mouth cone and maxillule, anteroventral; E,
Mandible, anterior; F, Maxillule, lateral; G, Maxilla, posterior. Scale bars lOOum unless otherwise
stated: A = 1 mm, B = 200 um, c = 50 um.

264

G. A. BOXSHALL

B

mnd

mxl

ant

COPEPOD ENDOPARASITES OF BRITTLE STARS 265

These are regarded as diagnostic characters for the Asterocheridae by Stock (1987). The very
reduced antenna is rare in siphonostomatoids, occurring mainly in species of the Nicothoidae.
However, Codoba is not related to the Nicothoidae which primitively have reduced antennules
and a well developed antenna. The typical subchelate antenna of siphonostomatoids is primarily
an organ for grasping the host and its reduction is here interpreted as an adaptation to
endoparasitism. Subject to a full review of the families of siphonostomes parasitic on invertebrates,
which is in progress, the genus Codoba is left unassigned to any of the families recognised at
present.

Review of other endoparasitic genera and species
AMPHIUROPHILUS Delamare Deboutteville, 1962

In his revision of the members of the family Philichthyidae parasitic on European fishes Delamare
Deboutteville (1962) correctly removed Philichthys amphiurae Herouard, 1906 from the genus
Philichthys Steenstrup, 1862 on the basis of differences in male morphology. He established a new
genus, Amphiurophilus, to accommodate Philichthys amphiurae. Amphiurophilus is a junior
objective synonym of Ophioithys which was established by Heegaard in 1951 with the same type
species, Philichthys amphiurae of Herouard (1906). It is a subjective synonym of Parachordeumium
Le Calvez, 1938 (see below).

ARTHROCHORDEUMIUM Stephensen, 1918

DIAGNOSIS. Postmetamorphosis adult female highly transformed; body somewhat dorsoventrally
compressed, indistinctly segmented and with short, paired lateral processes. Two pairs of limbs
present; antennules and maxillae. Antennules unsegmented, bifid at tip. Maxillae 3-segmented,
including terminal claw. Egg masses irregularly wrapped around body.
Male and copepodid unknown.

TYPE SPECIES. Arthrochordeumium appendiculosum Stephensen, 1918 (by subsequent designation,
Stephensen, 1933).

This is a valid genus established by Stephensen (1918) for A. appendiculosum. A second species,
Arthrochordeumium asteromorphae, was described by Stephensen (1933).

CHORDEUMIUM Stephensen, 1918

DIAGNOSIS. Postmetamorphosis adult female moderately transformed; body cylindrical with
paired lateral swellings on genital complex only; segmentation distinct. Median process present
posterodorsally on genital complex. Three pairs of cephalic appendages present. Antennules
unsegmented, bifid at tip. Antennae reduced to papilliform processes. Maxillae 3-segmented,
including terminal claw. Legs 1 to 4 represented by slender, laterally-directed processes. Egg mass
extruded posteriorly into cyst surrounding parasite.

Adult male similar to female in general fades but with better defined segments. Posterodorsal
median process absent. Testes paired.

Copepodid stage without discrete caudal rami, caudal setae located on margin of anal somite.

TYPE SPECIES. Chordeumium obesum (Jungersen, 1912).

This genus was proposed by Stephensen (1918) as a replacement name for Chordeuma of Jungersen
(1912). Chordeuma was preoccupied for a genus of Myriapoda (Kock, 1847). The type species,
Chordeuma obesum, described by Jungersen (1912,1914) becomes the type species of Chordeumium
by monotypy.

Fig. 2 Codoba discoveryi Heegaard, 1951. Adult female syntype. A, Maxilliped, posterior; B, Leg 1,
ventral; C, leg 2, ventral; D, leg 3, ventral; E, leg 4, ventral; F. Area between mouth cone and antennule
showing antenna (ant), mandible (mnd) and maxillule (mxl). All scale bars 100 urn.

G. A. BOXSHALL

B

mx

Fig. 3 Lernaeosaccus ophiacanthae Heegaard, 1951. Holotype female. A, lateral; B, ventral view
showing antennules (atl), presumed maxillae (mx) and legs 1-4 (1-4); C, Posterior end of body,
posterodorsal view showing median abdominal process and position of detached egg. Scale bars 1 mm
unless otherwise stated: c = 0-25 mm.

LERNAEOSACCUS Heegaard, 1951

DIAGNOSIS. Postmetamorphosis adult female sac-like, lacking external segmentation. Single pair
of lateral processes present on trunk. Posterior end concave containing a median posterior lobe.
Lobate antennules and vestigial maxillae present anteriorly. Swimming legs 1-4 reduced to tiny,
posteriorly directed lobes. Eggs large, released into masses surrounding female.
Male and copepodid stages unknown.

TYPE SPECIES. Lernaeosaccus ophiacanthae Heegaard, 1951 (by monotypy).

Lernaeosaccus ophiacanthae Heegaard, 1951
POSTMETAMORPHOSIS FEMALE. Body (Fig. 3A) highly transformed, lacking distinct segmentation

COPEPOD ENDOPARASITES OF BRITTLE STARS 267

and tagmosis. Body swollen, rounded anteriorly and dorsally, flattened ventrally and concave
posteriorly. Total body length of holotype 2-87 mm. Posterior part of body of holotype reflexed
dorsally, probably due to fixation. Single pair of lateral lobes present posteriorly behind fourth legs
(Figs 3 A-B). Concave posterior end containing genital openings, a median abdominal process and
a median lobe. The median abdominal process originates close to the genital apertures (Fig. 3C).
Eggs located at genital openings removed during examination. Eggs large; maximum diameter
173 um. Egg masses not contained within sacs; wrapped around body within capsule produced by
host.

Antennules represented by unarmed lobes (Fig. 3B, atl). Oral area obscured on holotype but
apparently comprising a median structure and a pair of more posteriorly located limbs (probably
maxillae, Fig. 3B, mx). Legs 1-4 present, reduced to unsegmented, unarmed and posteriorly
directed lobes on ventral body surface (Fig. 3B, 1-4).

MATERIAL EXAMINED. Holotype: Adult female (BM(NH) No. 1982.242) collected from
Ophiacantha disjuncta (Koehler) at Discovery Stn 190, in 316m off the Palmer Archipelago on
24.03 . 1 926. The depth and date on the label differ from those given by Heegaard (1951), which were
278 m and 14.03.1926 respectively.

REMARKS. This species was described by Heegaard (1951) upside down. The paired structures
referred to by Heegaard as maxilliped 2 (Heegaard, 1951: Fig. 4b) are eggs. Dissected from the
holotype, they were found to contain a nauplius at an early stage of development but with 3
recognisable pairs of appendages. The structure identified as a mouth cone containing mandibles is
here reinterpreted as a median abdominal lobe. The thickening of the cuticle was misinterpreted by
Heegaard as the paired mandibles. Close inspection of the trunk of the holotype revealed 4 pairs of
posteriorly directed legs that were overlooked by Heegaard (1951). This parasite is closely related
to Chordeumium obesum. The body morphology and appendage positioning are very similar. They
are maintained as separate genera because of the differences in structure of the posterior part of the
body.

OPHIOICODES Heegaard, 1951

DIAGNOSIS. Postmetamorphosis adult female highly transformed; body asymmetrical, irregularly
shaped and provided with numerous lateral and dorsal processes. No traces of segmentation
visible. Female with midventral groove.

Male highly transformed; body elongate, unsegmented and with a pair of long, slender lateral
processes. Testes paired. Male lies in midventral groove of female.

Copepodid stage unknown.

TYPE SPECIES. Ophioicodes asymmetrica (Pyefinch, 1940).

This is a valid genus. It was established by Heegaard (1951) on the basis of differences in male
morphology between Ophioica asymmetrica Pyefinch, 1940 and the other 3 species of Ophioika
Stephensen, 1933. He designated Ophioica asymmetrica Pyefinch, 1940 as type.

OPHIOIKA Stephensen, 1933

DIAGNOSIS. Postmetamorphosis adult female highly transformed. Body unsegmented, apparently
globular due to positioning of about 5 pairs of long lateral processes which curve ventrally. Four to
six egg masses held within space enclosed by processes. Pair of small anterior processes possibly
representing antennules. No other limbs recognisable. Median conical process may represent oral
cone.

Males degenerate body form with single well developed testis, male penetrating body of female.

Copepodid unknown.

TYPE SPECIES. Ophioika ophiacanthae Stephensen, 1933.

This genus was established by Stephensen (1933) to accommodate Ophioika ophiacanthae

268 G. A. BOXSHALL

Stephensen 1933. This becomes the type species by monotypy. Stephensen (1935) added a second
species, Ophioica appendiculata, misspelling the generic name Ophioika. This is corrected here.

OTHER SPECIES. Ophioikia appendiculata Stephensen, 1935; Ophioika tenuibranchia Heegaard, 1 95 1 .

0P///0/77/FS Heegaard, 1951

Heegaard (1951) recognised the close relationship between Philichthys amphiurae and the genus
Ophioika. He removed this species from Philichthys and proposed a new genus, Ophioithys, with P.
amphiurae Herouard, 1906 as type, "i his name is a senior objective synonym of Amphiurophilus as
both genera have the same designated type species. This genus is, however, recognised herein as a
synonym of Parachordeumium Le Calvez, 1938 (see below).

PARACHORDEUMIUM LeCtivez, 1938

DIAGNOSIS. Postmetamorphosis female highly transformed; body symmetrical, unsegmented,
lacking limbs except for maxillae. Body with 4 pairs of major lateral processes, sometimes
branched, forming enclosure containing egg masses; several paired and/or median papillae present
dorsally; abdominal process well developed. Genital apertures paired. Maxillae 3-segmented,
including terminal claw.

Adult male small, highly transformed, living in permanent association with female; body
unsegmented, drawn out into long abdominal process posteriorly, anteriorly with pair of long
lateral processes. Maxillae 3-segmented, including terminal claw.

Copepodid hatching with 3-pairs of developed biramous legs; lacking discrete caudal rami,
caudal setae located on margin of anal somite. Antennules 5-segmented. Antenna 3-segmented,
lacking exopod. Maxillae 3-segmented with terminal claw. Other cephalic appendages absent.

This is a valid genus established to accommodate a single species, P. tetraceros Le Calvez 1938,
which was the type species by monotypy. The dorsal and ventral figures of the adult postmetamor-
phosis female P. tetraceros given by Le Calvez (1938) do not differ significantly from the figures in
the detailed redescription of Amphiurophilus amphiurae by Goudey-Perriere (1979). Both these
species inhabit the genital bursae of the Amphipholis squamata Delia Chiaje in European waters
and it is here proposed that they be synonymised. The oldest available name for this species is
Parachordeumium amphiurae (Herouard, 1906) and this is the type species of Parachordeumium.

TYPE SPECIES. Parachordeumium amphiurae (Herouard, 1906) new combination, (syn. P. tetraceros
Le Calvez, 1938).

The three new species described by Goudey-Perriere (1979) and placed within the genus
Amphiurophilus are here transferred to the genus Parachordeumium. These new combinations are:
Parachordeumium bocqueti (Goudey-Perriere. 1 979); Parachordeumium humesi (Goudey-Perriere,
1979); Parachordeumium hendleri (Goudey-Perriere, 1979).

REMARKS. The third appendage of the copepodid stage of P. amphiurae (as Amphiurophilus
amphiurae was identified as the mandible by Goudey-Perriere (1979). This limb is retained in the
adult of both sexes. Its morphology (the segmentation and possession of a terminal claw) is most
atypical for a mandible and this limb is here reinterpreted as the maxilla.

CHORDEUMIIDAE New Family

DIAGNOSIS. Copepods endoparasitic in ophiuroids; adult females more-or-less highly transformed
typically losing external segmentation and often with paired lateral and median dorsal or posterior
processes. Lateral processes where fully developed forming 'cage' enclosing egg masses. Cephalic
appendages reduced or absent; antennules and maxillae typically retained in adult. Antenna
sometimes present as vestige in adult. Maxillipeds absent. Legs 1-4 reduced to uniramous pro-
cesses or absent. Eggs or eggs masses retained within cyst of host origin enclosing female. Adult
male typically transformed, with at most one pair of lateral processes. Maxillae typically retained

COPEPOD ENDOPARASITES OF BRITTLE STARS 269

by adult male, sometimes antennules and antennae also. Copepodid larva lacking discrete caudal
rami; caudal setae present on margin of anal somite.

TYPE GENUS. Chordeumium Stephensen, 1918.

OTHER INCLUDED GENERA. Arthrochordeumium, Lernaeosaccus , Ophioicodes, Ophioika and
Parachordeumium.

Discussion

It is difficult to produce a meaningful diagnosis for the new family because so many of the species
are extremely modified for their endoparasitic mode of life. Comparison between the better known
species, Chordeumium obesum and Parachordeumium amphiurae, provides the best apomorphy for
the family, based on the developmental stages. Both C. obesum and P. amphiurae lack discrete
caudal rami in the copepodid stage (Jungersen, 1914; Goudey-Perriere, 1979). This character is
rare in copepods and serves to link Chordeumium and Parachordeumium. This relationship is
central to the definition of the family because the former genus shares several characters (the
structure of the antennules and maxillae in the adult female, for example) with Arthrochordeumium
and the latter, several characters of gross female morphology (the possession of paired lateral
processes enclosing the egg masses, for example) with the genera Ophioicodes and Ophioika. The
genus Lernaeosaccus, now reinterpreted, is apparently closely related to Chordeumium. Both
genera have retained legs 1-4 as tiny lobes in the adult female, both have a pair of lateral processes
posteriorly and both have a median abdominal process located just posterior to the gonopores.

The new family is placed in the order Siphonostomatoida on the basis of the reports of a mouth
cone in Ophioika by Stephensen (1935). The mouth cone described by Heegaard (1951) in
Lernaeosaccus is an abdominal process. It is probable that the highly modified female body from
has been derived independently within the family, from a siphonostomatoid precursor that moved
freely over the surface of the brittle stars. The species of the family Cancerillidae are typically
ectoparasites of brittle stars (Emson et al., 1985) and the family Asterocheridae contains species
that live in the stomach of basket stars (Humes, 1986). A common ancestor shared with either of
these families could have made the step towards a specialised endoparasitic existence. A relation-
ship with the highly transformed families of fish parasites, such as the Chondracanthidae,
Lernaeopodidae and Philichthyidae, is regarded as extremely improbable.

The endoparasitic Cucumaricola notabilis Paterson, 1958 may be closely related to the
Chordeumiidae. The highly transformed morphology of the adult female, with its paired lateral
processes, resembles that of the chordeumiids. Also, this species retains only the antennules,
antennae and maxillae in the adult female (Paterson, 1958), as in Chordeumium itself. It differs
from the new family in having discrete caudal rami in the copepodid stage, although the
presence of 5-segmented antennules at this stage is another similarity between Cucumaricola and
Parachordeumium.

Acknowledgement

The author is grateful to Prof. Jan Stock (University of Amsterdam) for his valuable comments on the
manuscript.

References

Delamare Deboutteville, C. 1962. Prodrome d'une faune d'Europe des Copepodes parasites de poissons. Les

Copepodes Philichthyidae (Confrontation des donnees actuelles). Bulletin du Musee Oceanographique de

Monaco 59: 3-44.
Emson, R. H., Mladenov, P. V. & Wilkie, I. C. 1985. Studies of the biology of the West Indian copepod

Ophiopsyllus reductus (Siphonostomatoida: Cancerillidae) parasitic upon the brittle star Ophiocomella

ophiactoides. Journal of Natural History 19: 151-171.

270 G. A. BOXSHALL

Gotto, R. V. 1979. The association of copepods with marine invertebrates. Advances in Marine Biology 16:

1-109.
Goudey-Perriere, F. 1979. Amphiurophilus amphiurae (Herouard). Crustace Copepode parasite des bourses

genitales de 1'Ophiure Amphipholis squamata Delia Chiaje, Echinoderme: morphologic des adultes et etude

des stades juveniles. Cahiers de Biologic Marine 20: 201-230.
Heegaard, P. 1951. Antarctic Parasitic Copepods and an Ascothoracid Cirriped from Brittle-stars.

Videnskabelige Meddelelserfra Dansk Naturhistorisk Forening i Kjobenhavn 113: 171-190, pi. I-II.
Herouard, E. 1906. Sur un nouveau Copepode parasite d'Amphiura squamata. Compte Rendu Hebdomadaire

des Seances de V Academic des Sciences, Paris 142: 1287-1289.
Humes, A. G. 1986. Two new species of Copepoda associated with the basket star Astroboa nuda

(Ophiuroidea) in the Moluccas. Zoologica Scripta 15, 323-332.
Jungersen, H. F. 1 9 1 2. Chordeuma obesum, a new parasitic copepod endoparasitic in Aster onyx loveni. Report.

British Association for the Advancement of Science, 82nd Meeting (1912): 505-506.

— 1914. Chordeuma obesum, a new parasitic copepod endoparasitic in Asteronyx loveniM. Tr. Mindeskrift

for Japetus Steenstrup. 1 (16): 1-19, pi. I-II.
Le Calvez, J. 1938. Parachordeumium tetraceros n. gen. n. sp., Copepode gallicole parasite d'une ophiure de

Villefranche-sur-Mer. Compte Rendu du Congres des Societes savantes de Paris, Section des Sciences 71:

259-263.
Paterson, N. F. 1958. External features and life cycle of Cucumaricola notabilis nov. gen. et sp., a copepod

parasite of the holothurian Cucumaria. Parasitology 48: 269-290.
Pyefinch, K. A. 1940. The anatomy oWphioica asymmetrica, sp. n., a Copepod endoparasitic in an Ophiuroid.

Journal of the Linnean Society of London 41: 1-19.
Stephensen, K. 1918. On a gall-producing parasitic Copepod infesting an Ophiurid. Videnskabelige

Meddelelserfra Dansk Naturhistorisk Forening i Kjobenhavn 69: 263-275.

- 1933. Some new Copepods, parasites of Ophiurids and Echinids. Videnskabelige Meddelelserfra Dansk

Naturhistorisk Forening i Kjobenhavn 93: 197-213.

1935. Two Crustaceans (a Cirriped and a Copepod) endoparasitic in Ophiurids. Danish Ingolf-

Expedition3(\2):\-\8.

Stock, J. H. 1 987. Copepoda Siphonostomatoida associated with West Indian hermatypic corals 1 : associates
of Scleractinia: Faviinae. Bulletin of Marine Science 40: 464-^83.

Manuscript accepted for publication 3 May 1988

A new genus of tantulocaridan

(Crustacea: Tantulocarida) parasitic on a harpacticoid

copepod from Tasmania

Geoffrey A. Boxshall

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

Introduction

There are now eleven described species of Tantulocarida placed in five genera (Boxshall & Lincoln,
1987). One of these, Basipodella Becker, contains two species both of which are parasitic on
copepods. The other four genera parasitise tanaid, isopod, cumacean and ostracod hosts. B.
harpacticola Becker was described from unidentified harpacticoid hosts caught at depths of 2000
to 5000 m in the Peru Trench in the eastern Pacific (Becker, 1975). B. atlantica Boxshall & Lincoln
was found at a depth of about 3000 m in the North Atlantic to the southwest of the Azores, on a
copepod belonging to the harpacticoid family Tisbidae (Boxshall & Lincoln, 1983). Whilst
examining a collection of harpacticoids from the Bass Strait, off Tasmania, a single specimen of a
Stenhelia species was found bearing a tantulocaridan on the side of its urosome (Fig. 1). This
specimen, a tantulus larva containing a developing male, is described below as a new genus.

Description
AUSTROTANTULUS gen. n.

DIAGNOSIS. Class Tantulocarida. Tantulus larva with first thoracic tergite partly concealed beneath
posterior margin of dorsal cephalic shield; cephalic shield ornamentation comprising longitudinal
lamellae and pores; thoracopods 1-5 of tantulus larva biramous, with well developed endites,
uniramous leg 6 with coupling spines on protopod; abdomen of tantulus 2-segmented; adult male
formed within trunk sac originating posterior to sixth thoracic tergite of preceding stage.

TYPE SPECIES. Austrotantulus lincolni gen. et sp. n.

ETYMOLOGY. The generic name is derived from the Latin australis meaning South, and tantulus
which forms part of the name of the class Tantulocarida.

Austrotantulus lincolni gen. et sp. n.

TANTULUS LARVA. The body (Fig. 2A) comprises the cephalic shield, 6 free thoracic somites and a
2-segmented abdomen. The body length is 125 um, measured from the tip of the cephalic shield to
the posterior margin of the abdomen, excluding the caudal setae. This may be an overestimate
because expansion of the trunk sac may have caused separation of the thoracic tergites. The
cephalic shield (Fig. 2B) is longer than wide (47 x 32 um) and tapers anteriorly. The rostrum is
absent. The oral disc has a diameter of about 1 5 um and is positioned anteriorly so that it is visible
in dorsal view. The surface ornamentation of the head, as seen by light microscopy, consists of
longitudinal lamellae dorsally and oblique lamella on the downturned ventrolateral margins.
Associated with these lamellae are at least 5 pairs of pores, as marked on Figure 2B. Internally a
pair of chitinous bars leads towards the central pore of the oral disc. The cephalic stylet is very
slightly curved and is about 22 um long.

Bull. Br. Mus. not. Hist. (Zool.) 54(6): 271-274 Issued 24 November 1988

272

G. A. BOXSHALL

Fig. 1 Austrotantulus lincolni gen. et. sp. n. Holotype tantulus male undergoing metamorphosis
attached to its harpacticoid host. Scale bar in um.

The 6 free thoracic tergites are not conspicuously ornamented. The first tergite is partly
concealed beneath the rear margin of the cephalic shield. Each thoracic somite bears a pair of
well developed legs.

The 2-segmented abdomen is 27 um in length (Fig. 2C) and is deflected ventrally by the
expansion of the trunk sac posterior to the sixth tergite. The first abdominal somite is wider than
long (5x13 um), the second longer than wide (21x13 um). The caudal rami are each represented
by 1 short and 2 long setae arising from a common base.

There are no cephalic appendages. Thoracopods 1 to 5 have a large unsegmented protopod
bearing a single endite which originates at the proximal rim of the limb. The armature of the endites
cannot be discerned by light microscopy, but probably resembles that of Deoterthron harrisoni (see
Boxshall & Lincoln, 1987). Thoracopods 1 to 5 are biramous. The exopod is 2-segmented and
carries 2 apical setae in leg 1 (Fig. 2D). It bears 4 setae apically in legs 2 (Fig. 2E) to 5. The endopod
is more than twice as long as the exopod and is indistinctly 2-segmented. The endopod of leg 1 is
armed with 2 apical spines only. The endopods of legs 2 to 5 each have the 2 apical spines and, in
addition, the proximal segment bears 2 long setae distally on its lateral margin. Leg 6 (Fig. 2F) is
uniramous and has a large protopod armed with 2 complex coupling spines on the medial margin.
There is no endite. The ramus is 1 -segmented and armed with 2 long, curved setae.

MALE. The trunk sac of the holotype contained a male at an intermediate stage of development.
This developing male was still attached via the umbilical cord. The tagmosis of the adult male could
be discerned only in part, through the wall of the trunk sac. The first 2 pedigerous thoracic somites
were fused to the head to produce a cephalothorax of 7 somites and there were 4 free thoracopod-
bearing somites. However, the segmentation of the folded abdomen was not visible. A cluster
of aesthetascs was present anteriorly on the cephalothorax, as described for the adult male of
D. harrisoni. The male was not in a sufficiently advanced state of development for it to be dissected
out of the enclosing trunk sac.

ETYMOLOGY. The species is named after Roger Lincoln in recognition of his work on the
Tantulocarida.

MATERIAL EXAMINED. Holotype: <$ tantulus, parasitic on the harpacticoid Stenhelia sp. from fine
calcareous mud collected at about 22m depth in the Bass Strait (at approximately 41°00'S
146°00'E), off Round Hill Point, near Burnie on the north coast of Tasmania. BM(NH)
Registration No. 1987.418.

TANTULOCARIDA

273

Fig. 2 Austrotantulus lincolni gen. et sp. n. A, Holotype, lateral view; B, cephalic shield, dorsal; C,
abdomen, dorsal; D, leg 1, posterior; E, leg 2, posterior; F, leg 6, posterior. All scale bars in \am.

274 G. A. BOXSHALL

REMARKS. The new genus can be readily distinguished from Basipodella species, which are known
to parasitise harpacticoids, by the segmentation of the abdomen. The abdomen is 2-segmented
in the tantulus larva of the former and 6-segmented in the latter. Other differences include the
general pattern of the surface ornamentation of the cephalic shield. In the new genus this basically
consists of a system of longitudinal lamellae whereas in Basipodella species there are conspicuous
transverse and longitudinal lamellae. The combination of 2-segmented abdomen in the tantulus,
well developed endites and .ami on the tantulus thoracopods, and of longitudinal ornamentation
on the cephalic shield is found only in the family Deoterthridae, in species of Deoterthron Bradford
& Hewitt. However, the familial position of Austrotantulus is problematical because, whilst the
larval characters agree with the familial diagnosis of the Deoterthridae (Boxshall & Lincoln, 1987)
the position of the male trunk sac (posterior to the sixth tergite) is typical for the families Basipodel-
lidae and Microdajidae. The families Deoterthridae and Microdajidae may have been established
prematurely by Boxshall & Lincoln (1987), formalising morphological gaps between taxa which
represented a lack of data more than a phylogenetic reality. The new genus is provisionally placed
in the family Deoterthridae, although as more taxa are discovered it may become necessary to
revise the farrilial arrangement of the tantulocaridans.

The area around Tasmania and New Zealand is rich in tantulocaridans. Deoterthron aselloticola
Boxshall & Lincoln was described from an isopod host, Hydroniscus lobocephalus Lincoln, caught
at 3250-3340 m in the Tasman Sea (Boxshall & Lincoln, 1983) and D. megacephala Lincoln &
Boxshall was also found on an isopod, Haploniscus tangaroae Lincoln, taken at 1386m in the
Tasman Sea (Lincoln & Boxshall, 1983). The genotype, D. dentatum Bradford & Hewitt, was
reported from the ostracod Metavargula mazeri Kornicker collected in 384 m to the east of New
Zealand (Bradford & Hewitt, 1980). It is probable that this apparent species richness compared to
other geographical regions can be attributed to sampling effort in this recently discovered taxon,
rather than to any distinct zoogeographical pattern.

Acknowledgement

I would like to thank Dr Roger Lincoln for his comments on the manuscript.

References

Becker, K.-H. 1975. Basipodella harpacticola n. gen., n. sp. (Crustacea, Copepoda). Helgolander Wissens-

chaftliche Meeresuntersuchungen 27: 96-100.
Boxshall, G. A. & Lincoln, R. J. 1983. Tantulocarida, a new class of Crustacea ectoparasitic on other

crustaceans. Journal of Crustacean Biology 3: 1-16.

— & — - 1987. The life cycle of the Tantulocarida (Crustacea). Philosophical Transactions of the Royal

Society, London, series B 315: 267-303.
Bradford, J. M. & Hewitt, G. C. 1980. A new maxillopodan crustacean, parasitic on a myodocopid ostracod.

Crustaceana 38: 67-72.
Lincoln, R. J. & Boxshall, G. A. 1983. A new species of Deoterthron (Crustacea: Tantulocarida) ectoparasitic

on a deep-sea asellote from New Zealand. Journal of Natural History 17: 881-889.

Manuscript accepted for publication 11 May 1988

Unusual ascothoracid nauplii from the Red Sea

Geoffrey A. Boxshall

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

Ruth Bottger-Schnack

Institut fur Meereskunde, Abtlg. Fischereibiologie, Diisternbrooker Weg 20, D-2300 Kiel, Federal
Republic of Germany

Introduction

The study of larvae is important for elucidating the phylogenetic history of crustacean groups such
as the maxillopodans where macroevolutionary events involving progenesis and neoteny may have
occurred (Newman, 1983; Boxshall, 1983). It can also reveal homologies between apparently
disparate structures such as the attachment disc on the antennule of a cirriped cyprid and the
attachment claw on the same limb of an ascothoracid larva (Grygier, 1987a). One maxillopodan
taxon, the Facetotecta, is still only known from larvae (see Grygier, 1985, 1987/>). It is therefore,
of interest when larvae with clear maxillopodan affinities are discovered which cannot easily be
placed in a known group. Grygier (19870) described a 'metanauplius incertae sedis' from the South
China Sea which he tentatively identified as a postbrooding ascothoracid, possibly of a laurid
(Ascothoracida: Lauridae). During recent plankton studies in the central Red Sea (Bottger, 1987)
similar metanauplii were discovered in near surface waters, together with an earlier naupliar stage
of what appears to be the same species. These and a similar metanauplius that clearly represents a
related but distinct species are described in this paper.

Description

Nauplius type I

Body discoid (Fig. 1 A), convex dorsally and concave ventrally; entirely covered with dorsal shield.
Surface of shield (Fig. 1 B) punctuated with pores arranged in bilaterally symmetrical pattern
around lateral margins and in irregular rows either side of dorsal midline. Nauplius eye present.
Conspicuous dorsal pore present on swelling about in middle of shield. Margins of shield down-
turned, provided with extensive ventral band of pores around entire circumference. Posteriorly
shield drawn out into paired caudal processes each bearing a small ventral papilla. Papilla with
apical pore. Caudal spine originating on ventral surface near posterior margin, 120 urn in length,
armed with setules bilaterally. Body dimensions given in Table 1.

Paired frontal filaments simple, setiform but with a blunt tip (Fig. 1 A, ff); positioned either side
of midline just anterior to bases of antennules; median pore present between filaments.

Antennule (Fig. 1 A, all; Fig. 1C) uniramous, 4-segmented. First segment unarmed; second with
1 naked seta; third with 1 plumose seta; fourth segment showing some signs of subdivision near its
apex, armed with 5 plumose and 2 naked setae on and around apex.

Labrum (Fig. 1A, la) an elongate muscular lobe extending posteriorly from between bases of
antennae; bearing several rows of fine spinules around apex.

Antenna (Fig. 1A, ant; Fig. 2A) biramous, comprising protopod, 3-segmented endopod and
9-segmented exopod. Protopod with 3 endites; proximal endite small, armed with a few spinules;
middle endite strongly developed, produced into 2 spiniform processes with 3 slender setae; distal
endite strongly developed, produced into 2 divergent spiniform processes, each armed with
spinules, 1 naked seta present on anterior surface and a slender plumose seta distally on medial
surface. First endopod segment with 3 naked seta, second with 3, third with 4 at tip. First exopod
segment unarmed, second to eighth each armed with a long naked seta, ninth segment with 2 setae,
that on apex with a swollen base possible representing a tenth segment.

Bull. Br. Mus. not. Hist. (Zool.) 54(6): 275-283 Issued 24 November 1988

276

G. A. BOXSHALL & R. BOTTGER-SCHNACK

ff

Fig. 1 Red Sea nauplius type I. A, ventral view, showing frontal filaments (ff), antennule (atl),
antenna (ant), mandible (mnd), labrum (la), maxillule (mxl) and caudal furca (cf); B, dorsal view; C,
antennule, ventral. All scale bars 100 um.

Mandible (Fig. 1 A, mnd; Fig. 2B) biramous, comprising protopod, 3-segmented endopod and
6-segmented exopod. Protopod with 2 endites; proximal endite armed with a large hirsute seta, a
short spine and some spinules; distal endite armed with 4 stout setae, 3 of them plumose, and a row
of long spinules. First endopod segment with 2 rows of long spinules, a naked seta and 2 stout setae

ASCOTHORACID NAUPLII 277

bearing long setules, second with 3 naked setae, third with 4 naked setae. First to fifth exopod
segments each with a single long seta, that on segment 4 being unusually robust and unilaterally
spinulate; sixth segment with 3 naked apical setae.

Paired lobes located on ventral body surface just posterior to mandibles; armed with rows of
medially directed spinules. Maxillule (Fig. 1A, mxl) represented by a small lobe bearing a long,
posteriorly directed, apical seta, a short incurved spine and rows of medially directed spinules.

Caudal furca (Fig. 1 A, cf) comprising paired lobes each bearing an inner naked seta and a short
outer spine.

Table 1 Body dimensions and depth distributions of ascothoracid larvae type I in the central Red Sea.

STAGE

DEPTH (m)

DATE (time)

LENGTH (urn)

WIDTH (^im)

Nauplius

0-50

28.10.80(00:20)

443

350

0-50

05.11.80(15:20)

463

370

20-^0

04.11.80(15:20)

468

344

mean

458

355

Metanauplius

0-50

27.10.80(11:00)

732

591

0-50

05.11.80(00:13)

704

561

0-50

05.11.80(00:13)

728

564

50-100

27.10.80(11:00)

710

609

50-100

19.10.80(13:00)

774

589

50-100

05.11.80(00:13)

743

553

mean

732

578

Metanauplius type I

Body discoid (Fig. 2C) convex dorsally, concave ventrally; entirely covered with dorsal shield.
Surface of shield ornamented with fine lamellae anteriorly and posteriorly, and punctuated with
numerous pores arranged more or less symmetrically in an irregular row either side of the midline
and over entire dorsolateral area. Tripartite nauplius eye well developed. Middorsal pore present
but less conspicuous than in early nauplius. Posteriorly shield bearing 3 pairs of sensory setules.
Equatorial ring of marginal pores on ventral surface of downturned shield margins present.
Caudal processes bearing marked papillae both dorsally and ventrally; ventral papilla with an
apical pore. Caudal spine 278 um long. Body dimensions given in Table 1 . Frontal filaments as in
early nauplius.

Antennule (Fig. 3B) uniramous, 6-segmented. First segment unarmed; second and third each
with 2 plumose setae, fourth with 3 plumose setae and a prominent angled spine passing obliquely
across next segment; fifth segment with 4 long plumose setae, sixth with 4 long naked setae.
Labrum as in early nauplius.

Antenna (Fig. 4A) biramous, comprising 2-segmented protopod, 3-segmented endopod and 12-
segmented exopod. First protopodal segment (coxa) with single well developed endite bearing 2
slender naked setae basally and 2 curved spines apically, one of which is armed with small denticles
distally. Second segment (basis) produced into strong endite bearing 2 setulate spines apically, 2
plumose setae present on anterior surface at base of endite. First endopod segment with 3 naked
setae and a row of spinules; second with 7 marginal setae and a patch of spinules; third with 4 apical
setae. First exopod segment unarmed, second to eleventh each with a long seta, twelfth segment
with 1 medial and 1 apical seta, the latter with a swollen base probably representing a thirteenth
segment.

278

G. A. BOXSHALL & R. BOTTGER-SCHNACK

Fig. 2 Red Sea nauplius type I. A, antenna, medial; B, mandible, medial. Red Sea metanauplius type
I. C, dorsal view. Scale bars 100 jam unless otherwise stated.

Mandible (Fig. 4B) biramous, comprising 2-segmented protopod, 3-segmented endopod and
7-segmented exopod. First protopod segment (coxa) armed with a large plumose seta, a short spine
and some spinules; second armed with 5 stout setae, 4 of which are plumose, and some spinules.
First endopod segment bearing 3 stout spinulate setae, second with 6 naked setae, third with 3

ASCOTHORACID NAUPLII

279

unilaterally plumose and 2 naked setae. First exopod segment unarmed; second to fourth each with
a long seta; fifth with a robust, unilaterally spinulate seta; sixth with a plumose seta and seventh
with 3 plumose setae.

Maxillule (Fig. 3C) 2-segmented; first segment fused with body surface, armed with rows of

Fig. 3 Red Sea metanauplius type I. A, ventral view, with setae omitted from anterior limbs on left
side; B, antennule, medioventral; C, maxillule, ventral. Scale bars 100 ^m unless otherwise stated.

280

G. A. BOXSHALL & R. BOTTGER-SCHNACK

Fig. 4 Red Sea metanauplius type I. A, antenna, medial; B, mandible, medial. All scale bars 100 \im.

medially directed spinules that extend onto body; second segment bearing 2 short medial setae, 3
subapical setae on ventral surface and 3 distal margin setae.

Seven pairs of limb buds visible through body wall, marked externally only by tiny sclerotised
lobes at the tip of each bud. Limb buds presumably representing maxillae and first to sixth
thoracopods of subsequent stage. Caudal furca comprising paired lobes, each bearing a long inner
seta and a shorter outer seta.

ASCOTHORACID NAUPLII 281

MATERIAL EXAMINED. 3 nauplii and 6 metanauplii collected at and around Valdivia Stn 1 77 (cruise
29) in the central Red Sea (21°25-96'N 38°04-22'E) between 0 and 100 m (see Table 1), in a multiple
opening-closing net of mesh 0-1 mm. 1 nauplius and 2 metanauplii stored in the collections of the
BM(NH), Reg. Nos 1988.100-102.

REMARKS. The metanauplius type I from the Red Sea is very similar to that described from the
South China Sea by Grygier (19870). There are small differences in the armature and segmentation
of the antenna, mandible and maxillule, in the size of the caudal processes, and in the relative
lengths of the caudal spine and setae on the lobes of the caudal furca. However, the basic organis-
ation of these larvae is the same. As established by Grygier (19870), their possession of rudiments
of 6 pairs of thoracopods implies a maxillopodan affinity. The flattened, bowl-shaped dorsal shield
is typical of the Ascothoracida and the possession of an equatorial ring of pores around the shield
margin which is found only in some families of Ascothoracida (Grygier, 19870) is here regarded as
a derived character. These Red Sea larvae are referred to the Ascothoracida and, on the basis of
similarities in the caudal region, possibly to the family Lauridae. Ascothoracid larvae of this type
were distributed irregularly in small numbers in the upper 100 m (see Table 1) in the central Red
Sea. None was found further north, above the Kebrit Deep.

The free swimming nauplius does not contain an obvious store of yolk. This, combined with its
highly setose antenna and mandible each bearing well developed gnathobases, suggests that it is a
planktotrophic feeder. The metanauplius is about 1-6 times longer than the nauplius and shows a
marked increase in the numbers of limb segments. It is inferred from this that there is at least one
stage, intermediate in size and limb segmentation, between the two stages described above. The
metanauplius is clearly equivalent to the sixth nauplius (NVI) of cirripedes. The primordia of the 6
pairs of thoracopods visible through the integument indicate that the next moult will be to the
equivalent of the cyprid stage of development. The presence of a well developed, setose maxillulary
rudiment tends to confirm that the metanauplius is equivalent to the cirripede NVI which may have
a similar maxillulary rudiment (Moyse, 1987).

The earlier nauplius is more difficult to equate precisely with a given nauplius stage of cirripedes. It
appears to be a mid-development stage, most likely a NTV because it possesses 2 preaxial setae on the
antennule. Whilst antennule segmentation varies within the cirripedes the sequence of appearance
of preaxial setae on the antennule is remarkably constant. Typically the number of preaxial setae is
0 in NI-NII, 1 in NIII, 2 in NIV and 3 in NV-NVI. This pattern occurs repeatedly in both
lepadomorph cirripedes, such as Capitulum mitella Linnaeus (Yasugi, 1937), and balanomorphs,
such as Balanus eburneus Gould (Costlow & Bookhout, 1957) and Tetraclita serrata Darwin
(Griffiths, 1979). Also the presence of 4 setae on the apical segment of the endopod of both the
antenna and mandible is regarded as typical of the cirripede NIV, although this number does vary
within the cirripedes.

Thus, the free-living, postbrooding phase of ontogeny appears to comprise 3 or more stages. The
length of any brooded phase, prior to release of the larvae, is unknown but may be relatively short.
This presumed ontogenetic pattern is much less modified than that of any other ascothoracid (data
from Grygier, 1984). The discovery of the nauplius stage, probably equivalent to the cirripede
NIV, confirms Grygier's (19870) prediction that when ascothoracids release feeding naupliar
stages these will resemble late instar cirripede larvae more closely than when nauplii are brooded.

Metanauplius type II

A single metanauplius (Fig. 5) was found which differed significantly from that described above. It
is 808 um in length and has the same basic organisation. The primordia of the maxillae and 6 pairs
of thoracopods are visible through the integument but are smaller and more anteriorly located
than in the above metanauplius. Other differences include: the limbs are relatively smaller, the
antennary exopod is only 8-segmented, the mandibular exopod is only 5-segmented, the nauplius
eye is lacking, the labrum is broader and truncate, the maxillules have 5 setae, the caudal processes
are very small, the caudal furca bears 2 subequal pairs of setae, and there is no trace of any median
caudal spine.

282

G. A. BOXSHALL & R. BOTTGER-SCHNACK

Fig. 5 Red Sea metanauplius type II, ventral view. Scale bar 200 \im.

MATERIAL. A metanauplius collected on 23 February 1981 in the central Red Sea (21°23TN
38°04-7'E) during Valdivia cruise 29 between 0 and 50 m, in a multiple opening-closing net of mesh
0-1 mm. Stored in the collections of the BM(NH), Reg. No. 1988.198.

REMARKS. This metanauplius is also identified as that of an ascothoracid probably of the family
Lauridae, but there are sufficient differences to indicate that it represents a second species from the
Red Sea. It differs from the Red Sea metanauplius type I and resembles the metanauplius described
by Grygier (1987a) from the South China Sea in the following respects: the segmentation of the
antennary and mandibular exopods, the maxillules have 5 setae, the caudal processes are small and
the 2 pairs of setae on the caudal furca are subequal in length. It differs from the South China Sea
specimen in the lack of a median caudal spine.

Acknowledgements

Cruise 29 of the RV Valdivia was funded by the Red Sea Joint Commission, Jeddah, and by the Bundesminis-
terium fur Forschung und Technologic, Federal Republic of Germany. We are grateful to Dr Mark Grygier
for his valuable comments on an earlier draft of the manuscript.

References

Bottger, R. 1987. The vertical distribution of micro- and small mesozooplankton in the central Red Sea.
Biological Oceanography 4: 383-402.

ASCOTHORACID NAUPLII 283

Boxshall, G. A. 1983. A comparative functional analysis of the major maxillopodan groups. In: F. R. Schram

(Ed.), Crustacean Phylogeny, A. A. Balkema, Rotterdam. 121-143pp.
Costlow, J. D. & Bookhout, C. G. 1957. Larval development ofBalanus eburneus in the laboratory. Biological

Bulletin 112: 3\3-324.
Griffiths, R. J. I. 1979. The reproductive season and larval development of the barnacle Tetraclita serrata

Darwin. Transactions of the Royal Society of South Africa 44: 97-1 1 1 .
Grygier, M. J. 1 984. Comparative morphology and ontogeny of the Ascothoracida, a step towards a phylogeny of

the Maxillopoda. Ph.D. Thesis, University of California at San Diego, 417pp.

— 1985. Comparative morphology and ontogeny of the Ascothoracida, a step towards a phylogeny of the

Maxillopoda. Dissertation Abstracts International 45(8): 2466B-2467B.

- 1987a. Nauplii, antennular ontogeny, and the position of the Ascothoracida within the Maxillopoda.

Journal of Crustacean Biology 7: 87-104.

19876. New records, external and internal anatomy, and systematic position of Hansen's Y-larvae

(Crustacea: Maxillopoda: Facetotecta). Sarsial2: 261-278.
Moyse, J. 1987. Larvae of Lepadomorph barnacles. In: A. J. Southward (Ed.), Barnacle Biology, A. A.

Balkema, Rotterdam. 329-362pp.
Newman, W. A. 1983. Origin of the Maxillopoda: urmalacostracan ontogeny and progenesis. In: F. R.

Schram (Ed.), Crustacean Phylogeny, A. A. Balkema, Rotterdam. 105-1 19pp.
Yasugi, R. 1937. On the swimming larvae of Mitella mitella L. Botany and Zoology. Theoretical & Applied.

Tokyo 5: 792-796.

Manuscript accepted for publication 10 May 1988

New Nicothoid copepods (Copepoda:
Siphonostomatoida) from an amphipod and from
deep-sea isopods

Geoffrey A. Boxshall & Keith Harrison

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

Summary

One new genus, Cephalorhiza, and eight new species of parasitic copepods of the family Nicothoidae are
described. One new species, Sphaeronella australis, is parasitic in the brood pouch of a lyssianasoid amphipod
from southern Australia. All the other new taxa are parasites of deep-sea asellote isopods, from the North
Atlantic and Indian Oceans. Two new species belong to the genus Rhizorhina, 4 belong to Diexanthema and
one, Cephalorhiza flaccida, to the new genus.

Introduction

The family Nicothoidae contains eighteen genera of small, highly transformed copepods, all
of which are parasitic on other crustaceans. Several of these genera are known either from amphi-
pods (Stenothocheres Hansen) or from isopods (Diexanthema Ritchie, Choniorhiza Boxshall &
Lincoln, Nicorhiza Lincoln & Boxshall), or from both amphipods and isopods (Rhizorhina
Hansen, Sphaeronella Salensky). The present account describes a new species of Sphaeronella
found in the marsupium of an amphipod from Australia and several new taxa from deep-sea
asellote isopods collected in the North Atlantic and Indian Oceans. During an ecological study of
the asellotes of the Rockall Trough, off the west coast of Scotland, about 15,500 asellote isopods
were examined. Only four specimens harboured nicothoid parasites, which were found to represent
four new species. Examination of additional material from the Porcupine Seabight and from the
collections of the Centre National de Tri d'Oceanographie Biologique (CENTOB, Brest) made off
the island of Reunion in the Indian Ocean, revealed another three new species, one belonging in a
new genus.

Descriptions
Family NICOTHOIDAE

Rhizorhina hystrix sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, simplified to a globular, almost spherical,
trunk portion (Fig. IB) and an intricate branching holdfast (Fig. 1 A). Maximum width of globular
trunk 859 urn, maximum length 788 um; trunk featureless except for raised gonopores located about
353 um apart on posterior surface. Small anterior swelling tapers towards branching holdfast.
Holdfast 4-branched at origin, processes branching irregularly along length. Maximum extent of
holdfast within host about 680 um from origin. Arrangement of holdfast branches more or less
2-dimensional within host.

MATERIAL EXAMINED. Holotype ?, parasitic on a preparatory female of Eurycope complanata
Bonnier (sensu Wilson, 1982). Locality: Discovery Stn 50602 # 2 in the Porcupine Seabight
(51°1-0'N 13°7-2'W), depth 1955-1980 m, 07.vii.1979. Parasite located on arthrodial membrane
between tergites of pereon segments 3 and 4. Holotype stored in BM(NH), Reg. No. 1987.435.

Bull. Br. Mus. nat. Hist. (Zool.) 54(6): 285 299 Issued 24 November 1988

286

G. A. BOXSHALL & K. HARRISON

B

Fig. 1 Rhizorhina hystrix sp. n., Holotype female. A, Posterodorsal view of oral rootlet system; B,
Trunk sac, showing point of attachment to rootlet system. Rhizorhina aesthetes sp. n., Holotype
copepodid. C, Dorsal; D, Maxillule, lateral; E, Maxilla, posterior. Scale bars 200 urn, unless otherwise
stated: C = 50 um, D = 10 urn, E = 20 urn.

NICOTHOID COPEPODS 287

ETYMOLOGY. The species is named after the Porcupine seabight where it was collected.

REMARKS. It is difficult to characterise parasites that have undergone such a metamorphic reduc-
tion as Rhizorhina species. Four species of Rhizorhina have been described: R. ampeliscae Hansen
from the amphipods Ampelisca diadema (Costa) and A. brevicornis (Costa), R. serolis Green from
the isopod Serolis bromleyana Suhm, R. leptostracae Gotto from the leptostracan Nebaliella caboti
Clark and R. tanaidaceae Gotto from the tanaid Leviapseudes hanseni (Lang). Small differences
in size and shape of the trunk are given in Table 1 . The new species is much smaller than R. serolis
and also differs from this species in shape (Green, 1959). It is considerably larger than both R.
leptostracae and R. tanaidaceae, which also differ in shape, the former being somewhat flattened
dorsoventrally and the latter subquadrate in dorsal aspect (Gotto, 1984) in comparison with the

Table 1. Morphometrics of the adult females of Rhizorhina species.

GONOPORES

MAY

WIDTH

LENGTH

SHAPE

POSITION

SEPARATION

R. ampeliscae

1-05 mm

0-96 mm

subspherical

posterior

187 urn

R. serolis

2-5 mm

2-0 mm

subrectangular

posterior

1-2 mm

R. leptostracae

552 urn

521 urn

discoid

posterior

205 urn

R. tanaidaceae

594 um

500 n.m

subquadrate

posterior

258 urn

R. hystrix

859 urn

788 urn

subspherical

posterior

353 urn

R. aesthetes

282 urn

304 urn

subspherical

ventral

132 urn

almost spherical shape of R. hystrix. In size and shape the new species most closely resembles R.
ampeliscae (Hansen, 1892). They can be distinguished on the basis of the much wider separation of
the gonopores in R. hystrix. The branching holdfast system is probably not a good taxonomic
character. It has rarely been described and may well vary according to position in the host, as has
been described for the holdfasts of other mesoparasitic copepods (Fryer, 1961; Boxshall, 1989). It
is noteworthy that the holdfast is basically 4-branched in both the new species and in R. serolis
according to Green (1959). These branches are probably derived from the divided labrum and
labium as found by Lincoln & Boxshall (1983) in the closely related Nicorhiza.

Rhizorhina aesthetes sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, comprising globular trunk and branching
holdfast. Trunk (Fig. 2A) subspherical, longer than wide; maximum width 282 urn, maximum
length 304 um. Trunk lacking appendages, featureless except for gonopores located 132 urn apart
on ventral surface, just posterior to midlevel. Holdfast broken off in host.

COPEPODID STAGE. Body comprising 2-segmented prosome and 3-segmented urosome (Fig. 1C).
Total body length 136 um, maximum width 75 um. First urosome somite bearing pair of setules
dorsally and pair of setae posterolaterally representing the third swimming legs. Caudal rami
about as long as wide; armed with a dorsal seta, lateral seta and 3 distal margin setae, all naked.

Antennules 3-segmented (Fig. 2B); armature as follows: 1-1,11-1 + 1 aesthete, III-6+ 1 aesthete.
Aesthete on second segment inflated proximally and distally, with mid-level constriction. All setae
naked. Antenna absent. Mandible stylet-like, without apical teeth, palp absent. Maxillule (Fig.
ID) reduced to simple lobe bearing 2 apical naked setae. Maxilla (Fig. IE) 2-segmented; basal
segment unarmed, second segment bearing a terminal claw 19um long and seta 9 urn long.
Maxilliped (Fig. 2C) 5-segmented, including terminal claw. Fourth segment bearing seta 7 um
long and the terminal claw 24 um in length.

288

G. A. BOXSHALL & K. HARRISON

Fig. 2 Rhizorhina aesthetes sp. n. A, Paratype female trunk, ventral; B, Holotype copepodid antennule,
ventral; C, Maxilliped, posterior; D, leg 1, anterior; E, leg 2, anterior. Scale bars 20 um, unless
otherwise stated: A = 100 um

Swimming legs 1 and 2 biramous, with 1 -segmented rami. Leg 1 (Fig. 2D) exopod bearing 2
spines and 5 setae; endopod with 5 setae. Leg 2 (Fig. 2E) exopod bearing 2 spines and 4 setae,
endopod with 4 setae. Leg 3 represented by posterolateral setae on surface of first urosomal somite.

MATERIAL EXAMINED. Holotype copepodid, 1? paratype. Parasitic on an unidentified ischnomesid
(a fragment only). Locality: the Scottish Marine Biological Association's (SMBA) 'Permanent

NICOTHOID COPEPODS 289

Stn', sample ES172, in the southern Rockall Trough (54°39'N 12°17'W), depth about 2910m,
27. v. 1980. Adult female attached dorsolaterally to tergite of second pereon somite. Types stored in
BM(NH), Reg. Nos 1987.436 (Holotype) and 1987.437 (paratype).

ETYMOLOGY. The species name refers to the large aesthete present on the second antennulary
segment.

REMARKS. The new species is the only Rhizorhina described in which the adult female is longer than
broad (see Table 1). This difference in shape, together with the small size of the trunk and the
ventral position of the gonopores are sufficient to distinguish the female from other known species.
The copepodid larva has 3-segmented antennules as in other species of Rhizorhina but, in addition
to the usual apical aesthete, there is a conspicuous aesthete on the second segment.

Diexanthema nudum sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, comprising a swollen, slightly dorsoven-
trally flattened prosome and an unsegmented urosome (Figs 3A, B). Total body length in ventral
view 302 urn, maximum width 267 um. Urosome length 75 urn, maximum width 86 um. Prosome
bearing well developed mouth cone anteroventrally containing stylet-like mandibles. Paired
irregular lobes just anterior to base of mouth cone may represent modified antennae. Other mouth-
parts and legs 1-3 absent. Leg 4 represented by pair of minute papillae on ventral body surface.

Urosome bearing leg 5 ventrolaterally. Leg 5 comprising a simple unsegmented lobe armed with
3 distal spines (Fig. 3B). Gonopores large and unarmed, opening on posterior surface of urosome.
Postgenital abdominal segments reduced to small, median lobe on posterior surface of urosome.
Caudal setae lacking.

MATERIAL EXAMINED. Holotype 9, parasitic on a species of the desmosomatid Mirabilicoxa,
probably M. acuminata Hessler. Locality: SMBA Stn ES10 in the central Rockall Trough (56°37'N
11°04'W), depth about 2540m, 04.vii.1973. Only one host specimen was infected out of a total
of 61 examined from the Rockall Trough. Holotype stored in BM(NH), Reg. No. 1987.438.

ETYMOLOGY. The species name refers to the absence of cuticular hairs from the surface of the
prosome.

REMARKS. The new species is closely related to D. bathydiaita Ritchie, a parasite of a species of
Nannoniscus found in deep water off the western coast of Africa (Ritchie, 1975). The postmetamor-
phosis female of both species have the same gross morphology and both possess a mouth cone
containing mandibles, an irregular branching structure derived from the antennae and a lobate
fifth leg armed with 3 spines. The species differ in the presence of caudal setae and of a covering of
minute hairs over the cuticle of the prosome in D. bathydiaita (Ritchie, 1975). The tiny papillae
representing the fourth legs of the new species are not figured for D. bathydiaita but they may have
been overlooked.

The host isopod is probably referable to M. acuminata described from the Gay Head-Bermuda
transect off the eastern coast of the U.S.A. at depths of 3834 to 4800 m (Hessler, 1970). It differs
only in having a lower pereopodal setal count in the preparatory female.

Diexanthema corrugatum sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, comprising a swollen, globular prosome
and a small unsegmented urosome (Fig. 3C). Total body length in ventral view 536 um, maximum
width 507 um. Urosome length 107 um, maximum width 134 um. Oral area of holotype damaged,
mouth cone not observed. Irregular branching structure present in a semicircle around oral area
probably representing modified antennae. Other mouthparts absent. Leg 1 represented by pair of
minute papillae on ventral surface of prosome posterior to oral region. Leg 3 represented by pair of
naked setae located ventrolaterally. Leg 4 a free segment bearing 2 apical setae. Leg 5 absent.
Gonopores unarmed, opening on posterolateral surface of urosome. Caudal setae lacking.

290

G. A. BOXSHALL & K. HARRISON

MATERIAL EXAMINED. Holotype ?, parasitic on an as yet undescribed new species of the eurycopid
Acanthocope. Locality; SMBA 'Permanent Stn' in the southern Rockall Trough
(54°39'N12°17'W), depth about 2900m, 07.iv.1977. Only one host specimen was infected out of a
total of 1 1 1 examined from the Rockall Trough (84 from the Permanent Stn). Holotype stored in
BM(NH), Reg. No. 1987.439.

B

Fig. 3 Diexanthema nudum sp. n., Holotype female. A, Ventral view; B, Lateral view. Diexanthema
corrugatum sp. n., Holotype female. C, Ventral view. Scale bars A, B= 100 urn, C = 200 jam.

NICOTHOID COPEPODS

291

ft

Fig. 4 Diexanthema ritchiei sp. n., Holotype female. A, Ventral view. Diexanthema apoda sp. n.,
Holotype female. B, Dorsal view of female attached to pereopod of host, showing 4 oral rootlets inside
limb; C, Lateral view; D, Urosome, ventral. Scale bars A, D= 100 jim, B, C = 200 \im.

ETYMOLOGY. The species name refers to the highly furrowed cuticle of the prosome.

REMARKS. The new species is placed in Diexanthema because it possesses a globular prosome and
an unsegmented urosome as in the preceding species. It differs from the three described species of
Diexanthema in the structure of the third and fourth legs of the adult female. These legs are absent

292 G. A. BOXSHALL & K. HARRISON

from D. desisloma Ritchie and D. bathydiaita (Ritchie, 1975) and absent (leg 3) or reduced to
minute papillae (leg 4) in D. nudum. The third and fourth legs strongly resemble those ofNicorhiza
species but the new species cannot be placed in this genus because of its truncated urosome.
Nicorhiza species have a long, 4-segmented urosome (Lincoln & Boxshall, 1983).

The host occurs widely in the southern Rockall Trough between 2076 and 2925 m. It is clearly
distinguishable from other described species of Acanthocope.

Diexanthema rltchiei sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, comprising a swollen, globular prosome
and an unsegmented, slightly dorsoventrally flattened urosome (Fig. 4A). Total body length in
ventral view 426 urn, maximum width 307 um. Urosome length 61 urn, maximum width 108 um.
Prosome without recognisable mouthparts or legs. Irregular branching structure present anterior
to oral area may represent antennae as in other species of Diexanthema. Conical structure present
in oral region interpreted as broken base of oral rootlets. Urosome bearing leg 5 anteroventrally.
Leg 5 a simple lobe bearing 3 spines distally. Gonopores on posterolateral surface, each armed with
2 tiny setules. Posterior margin of urosome slightly concave, without trace of postgenital segments
or caudal setae.

MATERIAL EXAMINED. Holotype 9, parasitic on juvenile female of the ischnomesid Haplomesus
tenuispinus Vanhoffen. Locality: Discovery Stn 50604 # 1 in the Porcupine Seabight (50°6.2'N
13°52'W), depth 3490-3550 m, O4.vii. 1979. Holotype stored in BM(NH). Reg. No. 1987.440.

ETYMOLOGY. The species is named after the late Larry Ritchie, who established the genus
Diexanthema.

REMARKS. The new species is placed in Diexanthema because of the gross body form, a swollen
prosome and an unsegmented urosome. It possesses no obvious appendages on the prosome. The
irregular branching structure anterior to the oral area is found in all other Diexanthema species
except D. desistoma. D. ritchiei differs from other species in the structure of the feeding apparatus.
D. desistoma, D. bathydiaita, and D. nudum all possess a typical nicothoid mouth cone containing
stylet-like mandibles. The oral region of D. corrugatum was obscured (see above). The conical
structure in the oral region of this species was not a typical oral cone. It appeared to be the broken
stump of a rootlet system, as found in genera such as Rhizorhina, Choniorhiza and Nicorhiza. No
rootlets were found in the host although this was in poor condition. The generic concept of
Diexanthema is considerably broadened by the inclusion of this species as it now contains species
with an oral cone and species with oral rootlets. The evolution of a rootlet system appears to have
occurred independently several times within the Nicothoidae and, in our opinion, the presence of
rootlets alone is insufficient to justify generic separation when the gross body morphology is the
same.

The host, H. tenuispinus, was first described from the Davis Strait and from off the south coast of
Greenland (Hansen, 1916). In the present study 228 specimens were examined from depths of 1993
to 2925 m in the southern Rockall Trough. None was infected. Only the specimen taken in the
Porcupine Seabight was infected.

Diexanthema apoda sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, comprising a swollen prosome and a small
unsegmented urosome (Fig. 4B). Total body length in lateral view (Fig. 4C) 529 urn, maximum
width 495 um. Prosome extended anteriorly into a subrectangular 'hood'. Four oral rootlets
originating on midventral surface of 'hood'. Rootlets unbranched, between 486 and 659 um in
length and about 1 2 um in diameter. No other appendages or attachment structures present
externally on prosome. Urosome (Fig. 4D) bearing unarmed gonopores ventrolaterally and small
abdominal process posteromedially. Leg 5 absent. Caudal setae absent.

MATERIAL EXAMINED. Holotype ?, parasitic on a preparatory male of the ilyarachnid Bathybadistes
spinosissima (Hansen). Locality: SMBA 'Permanent Stn' in the southern Rockall Trough (54°39'N

NICOTHOID COPEPODS

293

Fig. 5 Cephalorhiza flaccida gen. et sp. n., Holotype female. A, Lateral view; B, Tip of head process,
dorsal; C, Tip of head process, ventrolateral; D, Urosome, ventral. Scale bars 100 urn, unless otherwise
stated: A = 500 urn.

12°17'W), depth about 2910m, 27. v. 1980. A total of 839 specimens of B. spinosissima was
examined from the entire Rockall Trough (783 from the 'Permanent Stn'), only 1 was infected.
Holotype stored in BM(NH), Reg. No. 1987.441.

ETYMOLOGY. The species name refers to the complete absence of any recognisable limbs.

REMARKS. This species, like D. ritchiei, has rootlets instead of a typical siphonostomatoid mouth
cone. It is even more reduced than D. ritchiei as it has lost the fifth legs but the general configuration
of the urosome is the same as in other Diexanthema species. The fifth legs have also been lost in D.

294 G. A. BOXSHALL & K. HARRISON

corrugatum. D. apoda differs from all of its congeners in the presence of an anterior 'hood' distinct
from the prosome.

The host, B. spinosissima, was first described from deep water (2702-3521 m) in and around the
Davis Strait (Hansen, 1916). Chardy (1979) also recorded this species (as Ilyarachna spinosissimd)
from the Bay of Biscay. It occurred widely in the southern Rockall Trough between 1993 and
2925 m and at the 'Permanent Stn' it was the commonest of the 79 species of asellotes recorded,
accounting for 14-7% of the asellote population.

Cephalorhiza gen. n.

DIAGNOSIS. Nicothoidae. Postmetamorphosis female highly transformed, comprising swollen
prosome and small, unsegmented urosome. Prosome bearing stout, somewhat twisted, head
process. Head process ornamented with 2 transverse chitinous lamellae dorsally. Oral apparatus
carried distally on head process, apparently consisting of rootlets. No recognisable limbs present
on prosome or urosome. Urosome unsegmented, bearing gonopores posterolaterally. Median
abdominal process present but lacking caudal setae.

TYPE SPECIES: Cephalorhiza flaccida gen. et sp. n.

REMARKS. The new genus is the nineteenth in the family. It differs from all other genera in the
possession of the stout head process which is embedded in the host up to the level of its base. It can
be placed in the Rhizorhina group of genera identified by Boxshall & Lincoln (1983). This group
now comprises five genera. Rhizorhina, Choniorhiza, Nicorhiza, Diexanthema and Cephalorhiza,
and 14 species including those taxa described in the present account. Members of this group have
highly transformed adult females and they exhibit the tendency to lose all cephalic appendages and
develop oral rootlets. The structure of the urosome of the new genus is similar to that of some
Diexanthema species and this is probably its closest relative within the group.

Cephalorhiza flaccida gen. et sp. n.

POSTMETAMORPHOSIS FEMALE. Body highly transformed, comprising swollen prosome and small,
unsegmented urosome (Fig. 5A). Body length 2-34 mm, measured in lateral view round curve
from tip of head process to end of urosome. Prosome globular with maximum diameter of
1-01 mm, bearing at its anterior extremity a stout, somewhat twisted, head process. Head process
ornamented with 2 transverse chitinous lamellae dorsally (Fig. 5B). Oral apparatus carried distally
on head process, apparently consisting of rootlets (Fig. 5C), but broken off in holotype. No
recognisable limbs present on prosome or urosome. Irregular branching structure present near
oral rootlets similar to the modified antennae of Diexanthema species but located posteriorly to
oral region and unlikely to be homologous. Urosome unsegmented; maximum length 129um,
maximum width 260 (am. Gonopores unarmed, opening on posterolateral surface (Fig. 5D).
Median abdominal process present but lacking caudal setae.

MATERIAL EXAMINED. Holotype $, parasitic on preparatory female of Ilyarachna antarctica
Vanhoffen; collected during the MD 32 cruise organised by the Terres Australes et Antarctiques
Francaises (TAAF, Paris); chef de Mission Alain Guille and sorted by CENTOB, Brest. Locality:
Marion-Dufresne Stn DS 106, off Reunion Island (20°47'5S 55°04'5E), depth 1710-1730 m.
Holotype stored in Museum National d'Histoire Naturelle, Paris, No. MNHN Cp39.

ETYMOLOGY. The genus name is derived from the Greek kephale, meaning a head, and rhiza,
meaning root. The species name, derived from the Latin flaccidus meaning flaccid, refers to the
posture of the head process.

REMARKS. The host has the form of Ilyarachna bicornis as described by Hansen (1916). /. bicornis
was synonymised with /. antarctica by Thistle (1980).

Sphaeronella australis sp. n.

ADULT FEMALE HOLOTYPE. Body subspherical (Fig. 6A) consisting of a small, somewhat dorsoven-
trally flattened head and a swollen, almost spherical trunk. Total body length 831 jam, maximum

NICOTHOID COPEPODS

295

Fig. 6 Sphaeronella australis sp. n., A, Holotype female, ventral view; B, Paratype male, lateral view.

Scale bars A = 200 ^m, B = 100 ^m.

width 623 (im. Size variable according to reproductive condition, trunk often much larger when
fully gravid. Head dorsoventrally flattened, slightly concave ventrally. Anterior rim of head
provided with fine marginal strip of hyaline membrane. Trunk covered with dense coat of tiny
denticles.

Antennules, mandibles, maxillules and maxillae as for male (see below). Isolated seta present
anterior to mouth cone may represent the antenna, absent in male. Ornamentation on surface of

296

G. A. BOXSHALL & K. HARRISON

maxilliped arranged into rows on female, rather than irregular as in male. Legs 1-2 comprising a
single free segment bearing 2 unequal setae apically. Caudal rami slightly longer than wide, bearing
3 distal setae.

ADULT MALE. Body highly transformed (Figs 6B, 7), dorsoventrally flattened, divided into
head and small trunk. Body length 477 um, maximum width 311 um. Head naked dorsally and
dorsolaterally, trunk covered with a dense coat of fine setules. Setules longer at junction between
head and trunk, and at posterior extremity of trunk (Fig. 6B). Anterior margin of head (Fig. 7)
complex, with deep indentations separating the median pseudorostrum from the lateral cephalic
processes on either side. Margin of lateral cephalic processes armed with fine setules. Lateral
margins of dorsal cephalic shield deflected ventrally, armed with row of submarginal setules.

Antennule (Fig. 8 A) 3-segmented; first segment with 2 setae, second with 1, third segment with 9
simple setae and a double seta at the apex. Antenna absent. Mandible (Fig. 8B) a simple stylet
armed with a marginal membrane distally; palp absent. Maxillule (Fig. 8C) comprising a simple

Fig. 7 Sphaeronella australis sp. n., Paratype male, ventral view. Scale bar 100 u

NICOTHOID COPEPODS

297

Fig. 8 Sphaeronella australis sp. n., Paratype male. A, Antennule, ventral; B, Mandible, posterior; C,
Maxillule, ventral; D, Maxilla, posterior; E, Maxilliped, anterior; F, Maxilliped, posterior; G, Tip of
maxillipedal claw, ventral; H, Leg 1, ventral; J, Leg 2, lateral; K, Leg 2, ventral; L, Caudal ramus,
ventral. Scale bars 50 urn, unless otherwise stated: A-C = 25 urn.

298 G. A. BOXSHALL & K. HARRISON

tapering process, bearing a naked seta proximally, and an isolated seta situated on ventral surface
of head near base of maxillulary process. Maxilla (Fig. 8D) 2-segmented; syncoxa robust, armed
with single row of stout spinules, basis claw-like, bearing 3 teeth distally and a row of fine setules.
Maxilliped (Figs 8E, F) 4-segmented, including the terminal claw; first segment robust armed with
many stout spinules on anterior and medial surfaces, and a row of slender setules distally on
posterior surface. Second segment unarmed, third bearing the terminal claw and a subapical
tricuspid spine. Terminal claw 33 um long, with complex quindentate tip.

Two pairs of uniramous swimming legs present. Leg 1 (Fig. 8H) slender, comprising 2 incomple-
tely fused segments with the indentation at midlevel marking the line effusion. Proximal segment
bearing a naked seta, distal segment armed with 2 transverse rows of fine setules, a short (20 um)
subapical spine and a long (70 um), sparsely pinnate, apical spine. Leg 2 (Fig. 8J, K) indistinctly
2-segmented, with deep indentation most apparent in lateral view (Fig. 8J). Proximal part bearing
a single seta on a small ventral swelling; distal part with a patch of fine setules, a subapical seta
(22 um) and a long (68 um), sparsely pinnate, apical spine. Caudal rami (Fig. 8L) bearing 3 naked
setae and a few setules distally.

A large female paratype (Museum of Victoria J 1 1 809) had produced 1 9 egg sacs with a mean size
of 795 x 681 um (range 699 to 883 um x 589 to 810 um based on 19 measurements). Typical sac
measuring 810 x 681 um contained 216 eggs.

MATERIAL EXAMINED. Holotype 9, 399 and 1 $ paratypes all parasitic in brood chamber of a species
of the lysianassoid genus Amaryllis. Locality: Holotype 9, Port Arthur, Tasmania (43°09'S
147°51E), 29 vii. 1909, Australian Museum (AM) No. P35852. Paratypes: I? (AM No. P35853) off
end of South Mole, Arthur Head, Freemantle, Western Australia, depth 6m, 25.xii.1983, coll.
J. K. Lowry; 1 9, 1 $ (Victoria Museum (VM) No. J 1 1 809), and 1 9 (J 1 1 8 1 0), FV 'Sarda', Bass Strait
Stn 112 (40°22-2'S 145°17'E), depth 40 m, 3.xi.l980, coll. G. C. B. Poore.

REMARKS. Sphaeronella contains 76 species, the majority (42 species) parasitic on amphipod hosts.
The new species belongs to the S. leuckartii group which Hansen (1897) recognised for the
following 9 species, S. antillensis Hansen, S. atyli Hansen, 5". chinensis Hansen, S. danica Hansen,
S. leptocheiri Hansen, S. leuckartii Salensky, S. elegantula Hansen, S. messinensis Hansen, and S.
vestita Hansen. This number was increased by the description of S. aorae by Scott (1905) and of S.
devosae and S. ecaudata by Stock (in Stock & De Vos, 1960). Green (1958) regarded S. elegantula
and S. aorae as synonymous with S. leuckartii. This group is characterised by the presence in the
male of a conspicuous rectangular projection in the middle of the frontal margin of the head (here
referred to as the pseudorostrum), by the rudimentary or absent antenna, and by the presence of a
tuft of hairs at the base of the maxillule. Within the group the new species is most closely related to
S. chinensis Hansen, which is known from the marsupium of Corophium bonelli Milne-Edwards
from Hong Kong. The morphology of the males is very similar, especially in the configuration of
the frontal margin of the head. However, the ventral sternal processes in the maxillipedal region of
the male are better developed and more widely divergent in S. australis than in S. chinensis. Also the
terminal seta on the apex of leg 2 is longer than the limb itself in S. chinensis but shorter than the
limb in S. australis.

The oral area of the male of the new species had collapsed inwards, thereby retracting the mouth
cone so that it is less visible in ventral view. In life it would protrude more, as in the lateral view of
S. chinensis given by Hansen (1897).

Acknowledgements

We would like to thank Dr J. D. Gage (SMBA) for making available material from the Rockall Trough
investigations, the Insitute of Oceanographic Sciences for donating the material from the Porcupine Seabight,
and Dr M. Segonzac for allowing us to examine CENTOB material. Joan Ellis sorted the parasitised isopods
from the Porcupine Seabight material. Thanks are also due to J. K. Lowry (Australian Museum) and G. C.
Poore (Victoria Museum) for the loan of S. australis.

NICOTHOID COPEPODS 299

References

Boxshall, G. A. 1989. Parasitic copepods of fishes: a new genus of Hatschekiidae from New Caledonia, and

new records of Pennellidae, Sphyriidae, and Lernanthropidae from the South Pacific and South Atlantic.

Systematic Parasitology 13: in press.
Boxshall, G. A. & R. J. Lincoln, 1983. Some new parasitic copepods (Siphonostomatoida: Nicothoidae) from

deep-sea asellote isopods. Journal of Natural History 17: 891-900.
Chardy, P. 1979. Structure of deep sea asellota assemblages in the Bay of Biscay; relationships with the abyssal

environment. Ambio Special Report 6: 79-82.

Fryer, G. 1961 . Variation and systematic problems in a group of Lernaeid copepods. Crustaceana 2: 275-285.
Gotto, R. V. 1984. Two new species of Rhizorhina (Copepoda: Siphonostomatoida) from leptostracan and

tanaidacean hosts. Journal of Natural History 18: 81 1-817.
Green, J. 1958. Copepoda parasitic on British Amphipoda (Crustacea), with a description of a new species of

Sphaeronella. Proceedings of the Zoological Society of London 131: 310-313.
- 1959. Sphaeronella serolis Monod, and a new species of Rhizorhina, copepods parasitic on the isopod

Serolis bromleyana Suhm (Crustacea). Proceedings of the Zoological Society of London 132: 647-654.
Hansen, H. J. 1892. Rhizorhina ampeliscae n. gen., n. sp. En ny til Herpyllobiidae, n. fam., horende Copepod,

snyltende paa Amp. laevigata Lilljb. Entomologiske Meddelelser 3: 207-234.

— 1897. The Choniostomatidae . A family of Copepoda, parasites on Crustacea Malacostraca. Copenhagen,

205pp, 13pl.

1916. Crustacea Malacostraca III.-V. The order Isopoda. The Danish Ingolf-Expedition, 3(5): 1-262.

Hessler, R. R. 1970. The Desmosomatidae (Isopoda, Asellota) of the Gay Head-Bermuda Transect. Bulletin

of the Scripps Institution of Oceanography 15: 1-185.
Lincoln, R. J. & G. A. Boxshall, 1983. Deep-sea asellote isopods of the north-east Atlantic: the family

Dendrotionidae and some new ectoparasitic copepods. Zoological Journal of the Linnean Society 79:

297-318.
Ritchie, L. 1975. A new genus and two new species of Choniostomatidae (Copepoda) parasitic on two deep sea

jsppods. Zoological Journal of the Linnean Society 57: 1 55-1 78.
Stock, J- H. & De Vos, A. P. C. 1960. Einige wirbellose tiergruppen des Dollart-Ems-estuarium. Verhandel-

ingen van het Koninklijk Nederlandsch Geologisch Mijnbouwkundig Genootschap (Geologische serie) 19:

203-220.
Thistle, D. 1980. A revision ofllyarachna (Crustacea, Isopoda) in the Atlantic with four new species. Journal

of Natural History 14: 1 1 1-143.
Wilson, G. D. F. 1982. Systematics of a species complex in the deep-sea genus Eurycope, with a revision of six

previously described species (Crustacea, Isopoda, Eurycopidae). Bulletin of the Scripps Institution of

Oceanography 25: 1-64.

Manuscript accepted for publication 3 May 1988

A new genus of Lichomolgidae (Copepoda:
Poecilostomatoida) associated with a phoronid in Hong
Kong

Geoffrey A. Boxshall

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

Arthur G. Humes

Boston University Marine Program, c/o Marine Biological Laboratory, Woods Hole,
Massachusetts 02543, U.S.A.

Introduction

Copepods are rarely reported as parasites or associates of the lophophorate phyla Bryozoa,
Brachiopoda and Phoronida. Two new harpacticoids, Tegastes knoepffleri and Peltobradya
bryozoophila, were described by Medioni & Soyer (1967) as associates of the bryozoans
Schizobrachiella sanguined (Norman) and Schizomavella linearis (Hassall) respectively, at
Banyuls-sur-Mer in southern France. Recently Humes & Boxshall (1988) described a new genus of
the family Myicolidae which occurs on the intertidal brachiopod Lingula anatina Lamarck col-
lected at Starfish Bay, New Territories, Hong Kong. This is the first known copepod associate of a
brachiopod. Examination of other invertebrates collected at the same time in Starfish Bay has
revealed a copepod associated with a phoronid, Phoronis australis Haswell, which is itself an
associate of the burrowing anemone Cerianthusfiliformis Carlgren. The phoronid lives embedded
in the mucilaginous tube of the Cerianthus (Morton & Morton, 1983). Three copepods were found
in the lophophore of the phoronid and are described below as a new genus of the family
Lichomolgidae.

Material and Methods

The phoronid was collected by a diver at a depth of 3 to 5 m in Hoi Sing Wan (Starfish Bay). The
phoronid was placed in a plastic bag, transported to the laboratory and examined live. The
copepods were picked off and fixed in 70% ethanol. All figures were drawn with the aid of a camera
lucida.

Descriptions
Genus PHORONICOLA gen. nov.

DIAGNOSIS. Body cyclopiform. Urosome in 9 4-segmented, in $ 5-segmented. Caudal ramus with 6
setae. Rostrum broad. First antenna 7-segmented. Second antenna 4-segmented, with the formula
1 , 1,4+1 articulated claw, and 4 + 3 articulated claws.

Labrum incised medially. Mandible tapering smoothly into lash. Paragnath a small hairy lobe.
First maxilla with 3 setae. Second maxilla with long slender lash, 2 setae near base of lash, and a
short proximally directed seta. Maxilliped in ? 3-segmented with pointed tip, in £ 4-segmented.

Legs 1-4 with 3-segmented rami, except for leg 4 endopod which is 1 -segmented. Leg 4 exopod
armed 1-0; 1-1; I, I, 5: endopod armed with an apical serrate spine. Leg 5 with a free segment
bearing 1 apical spine and 1 subapical seta. Leg 6 in $ represented by 2 setae situated on a small lobe
at each genital aperture, in £ represented by single seta on genital flap.

Associated with Phoronids.

Bull. Br. Mus. nat. Hist. (Zool.) 54(6): 301-307 Issued 24 November 1988

302 G. A. BOXSHALL & A. G. HUMES

TYPE SPECIES. Phoronicola spinulatus gen. et sp. n.

ETYMOLOGY. The generic name is derived from Phoronis, the host genus, and the Latin -cola,
meaning inhabiting.

Phoronicola spinulatus gen. et sp. n.

TYPE MATERIAL. 1 9 and 2^ from 1 Phoronis australis collected in 3-5 m in Hoi Sing Wan (Starfish
Bay), on the southern shore of Tolo Harbour, New Territories, Hong Kong by Dr P. G. Oliver on 3
April 1986. Holotype $ (BM(NH) Reg. No. 1987.415), and 2^ paratypes (BM(NH) Reg. Nos
1987.416-417) deposited in the British Museum (Natural History), London.

FEMALE. Body (Fig. 1 A) with stout prosome and slender urosome. Length (excluding caudal setae)
1-17 mm and greatest width 0-49 mm. Length to width ratio of prosome 1-41:1. First pedigerous
segment fused with cephalosome. Ratio of length of prosome to that of urosome 1-47:1.

Segment bearing leg 5 (Fig. IB) 62 x 139 um. Genital segment elongate, 189 um long, anterior
half with convex lateral margins and broader (1 33 um) than posterior half (straight sides and width
of 80 um). Genital areas located dorsolaterally about in middle of segment. Each area bearing a
small process armed with 2 smooth setae (Fig. 1 B), an apical seta 23 um long and a subapical seta
29 um long. Postgenital segment 52 x 72 um, anal segment 82 x 65 urn.

Caudal ramus (Fig. 1 A) 3-0 times longer than wide (84 x 28 um), bearing 6 setae. Outer lateral
seta 63 um long, naked and positioned on mid-dorsal surface. Dorsal seta naked, 55 um long.
Outermost terminal seta 84 um long, innermost terminal seta 106 um and 2 median terminal seta
377 um (outer) and 501 um (inner); all 4 of these setae plumose.

Rostrum (Fig. 1C) broad based, moderately well defined. First antenna 7-segmented as in male
(Fig. ID); formula for armature: 4, 12, 6, 3, 4+ 1 aesthete, 2+ 1 aesthete and 7+ 1 aesthete. Two
setae on first segment, 3 on fourth segment, 1 on sixth segment and 5 on seventh segment plumose.
Second antenna (Fig. IE) 4-segmented; armature 1, 1,4+1 articulated claw, and 4 + 3 articulated
claws.

Labrum (Fig. 1C) with 2 broad, tapering lobes. Mandible (Fig. IF) with a slender base carrying
distally a bipectinate blade. Paragnath a small hairy lobe. First maxilla (Fig. 1G) conical, with 3
elements. Second maxilla (Fig. 2A) 2-segmented. First segment unarmed. Second segment drawn
out into a slender apical lash with a comb of strong spinules along outer margin, and fine spinules
along inner margin. Second segment armed with a stout barbed spine on medial margin, a naked
spine near its base and a fine, proximally directed setule near the base of the outer margin.
Maxilliped (Fig. 2B) 3-segmented. First segment unarmed; second segment bearing a naked seta
and a stout spinose seta. Third segment armed with a naked spine, terminating in slightly curved
spiniform process bearing 3 spinules. Ventral area between bases of maxillipeds (Fig. 2C) and first
legs only slightly protuberant.

Legs \~4 (Figs 2D-F, 3A) biramous with 3-segmented rami except for endopod of leg 4 being
1 -segmented. Spine and seta formula of legs as follows:

PI coxa 0-1 basis 1-0 expl-0; 1-1; III , 1,4

enpO-l;0-l;I, 5
P2 coxa 0-1 basis 1-0 exp 1-0; 1-1 ; III, I, 5

enpO-l;0-2;I, II, 3
P3 coxa 0-1 basis 1-0 exp 1-0; 1-1 ; II, I, 5

enp 0-1; 0-2; I, II, 2
P4 coxa 0-0 basis 1-0 exp 1-0; 1-0; I, I, 5

enp I

Outer distal angle of legs 1-2 coxae with patch of fine spinules. Inner margin of basis of legs 3-4
with rows of long hairs. Rows of hairs present on inner margins of all exopod segments and outer
margins of all endopod segments. Leg 4 (Fig. 3 A) with 1 -segmented endopod 42 um in length,
bearing a few spinules proximally on inner margin and a longer row of spinules on outer margin.
Apical element spiniform, 62 urn long, armed with serrate membrane on both sides.

LICHOMOLGIDAE

B

303

Fig. 1 Phoronicola spinulatus gen. et sp. nov. A, Holotype female, dorsal; B, Fifth pedigerous and
genital segments, dorsal; C, Rostrum and labrum, ventral; D, Male first antenna, anteroventral (with
aesthetes absent in female marked with asterisks); E, Female second antenna, posterior; F, Mandible,
posterior; G, First maxilla, anterior. All scales in um.

304

G. A. BOXSHALL & A. G. HUMES
A

B

Fig. 2 Phoronicola spinulatus gen. et sp. nov. Female, A, Second maxilla, posterior; B, Maxilliped,
posteromedial; C, Ventral body wall between maxillipeds and swimming legs; D, Leg 1 , posterior; E,
Leg 2, posterior; F, Leg 3, posterior. All scales in urn.

LICHOMOLGIDAE

305

Fig. 3 Phoronicola spinulatus gen. et sp. nov. A, Female leg 4, posterior; B, Paratype male, dorsal; C,
Urosome, ventral; D, Maxilliped, posteromedial. All scales in jam.

306 G. A. BOXSHALL & A. G. HUMES

Leg 5 (Fig. IB) free segment about 1-2 times longer than wide (21 x 18um), with a small
spiniform projection at outer distal angle. Strong spiniform apical element 49 urn long, armed with
serrate membrane both sides; subapical seta 56 um long, with tiny spinules along anterior margin.
Dorsal seta 35 um. Leg 6 represented by process bearing 2 setae located in genital area (Fig. 1 B).

Egg sacs not seen.

Colour of live specimen white.

MALE. Body (Fig. 3B) similar to that of female in general form. Length 1 -04 mm excluding caudal
setae, greatest width 0-33 mm; length to width ratio of prosome 1 -73 : 1 . Ratio of length of prosome
to length of urosome 1-25:1.

Segment of leg 5 (Fig. 3C) 43 x 101 um. Genital segment 214 x 166 um. Three postgenital
segments from anterior to posterior 49 x 51, 41 x 52 and 54 x 51 um.

Caudal ramus similar to that of female but shorter, 56 x 20 um, ratio 2-8:1.

First antenna (Fig. ID) 7-segmented; lengths of segments measured along posterior border 26,
61, 19, 34, 44, 29 and 18 um respectively. Armature formula as for female except for additional
aesthetes on segments 2 and 4 (marked with asterisks on Figure ID).

Rostrum, second antenna, labrum, mandible, paragnath, first and second maxilla like those of
female. Maxilliped (Fig. 3D) slender, 4-segmented. First segment stout, unarmed. Second segment
with 2 naked inner setae, a row of spinules along inner surface and a shorter row of fine spinules
proximally. Third segment unarmed. Claw comprising short basal section representing the fourth
segment, armed with 2 unequal setae. Claw curved, 146 um in length, bearing a row of fine spinules
along concave margin.

Legs 1-4 similar to those of female. Leg 5 (Fig. 3C) free segment more elongate than in female,
1 -7 times longer than wide (17 x 10 um). Apical serrate element 38 um long, subapical seta 40 um.

Leg 6 (Fig. 3C) forming the posteroventral genital flap closing the genital aperture, armed with a
single apical seta 34 um long.

Spermatophores not seen.

Colour of live specimens white.

ETYMOLOGY. The specific name spinulatus refers to the spiniform nature of the single serrate
element on the endopod of leg 4.

REMARKS. The new genus belongs to the family Lichomolgidae. The lichomolgid genera
Aspidomolgus Humes, Haplomolgus Humes & Ho, Kelleria Gurney, Lichomolgella Sars,
Octopicola Humes, Paramacrochiron Sewell, Pseudomacrochiron Reddiah, Sewellochiron Humes
and Telestacicola Humes & Stock share with the new genus the 1 -segmented endopod of leg 4
(Humes & Stock, 1973). Most species of Macrochiron Brady also exhibit this 1 -segmented con-
dition. However, none of these genera has only a single apical element on the leg 4 endopod.
Aspidomolgus (II, I), Kelleria (II, 1), Octopicola (2, 1) and Telestacicola (II, 1) all have 3 elements,
Lichomolgella (II), Macrochiron (II), Paramacrochiron, Pseudomacrochiron (II) and Sewellochiron
(II) have 2 apical elements and Haplomolgus has none.

The presence of one articulated claw on segment 3 and 3 claws on segment 4 of the second
antenna is recorded in only a single species of Acaenomolgus Humes & Stock, within the
Lichomolgidae. Members of this genus, however, have a 2-segmented leg 4 endopod. Kelleria has 1
and 2 articulated claws on the third and fourth segments of the second antenna respectively and has
a 1 -segmented endopod on leg 4 but differs from Phoronicola in possessing an additional urosome
segment in both sexes and in the formula for the third exopod segment of leg 4 which is II, I, 5
compared with I, I, 5 in the latter.

No phoronid has previously been recorded as host to a copepod. Cerianthus, with which the
Phoronis is associated, is host to the copepod Boholia cerianthiphila Kossmann in the Philippine
Islands (Kossmann, 1877). This lichomolgid has not been recorded since its original description
and was regarded as insufficiently described or of uncertain position by Humes & Stock (1973) in
their revision of the family. The original description of B. cerianthiphila shows that it possesses 2
strong claws at the apex of the second antenna whereas Phoronicola has 3 slender articulated
setiform claws. The segmentation of the body is also different in these two genera. These differences
are sufficient to distinguish between them.

LICHOMOLGIDAE 307

Acknowledgements

We would like to thank Professor Brian Morton (University of Hong Kong) for the opportunity to collect this
material during the Second International Workshop on the Flora and Fauna of Hong Kong and Southern
China held during April 1986. We are also grateful to Dr P. G. Oliver (National Museum of Wales) who
collected the phoronid host.

References

Humes, A. G. & Boxshall, G. A. 1988. Poecilostome copepods associated with bivalve molluscs and a brachio-

pod at Hong Kong. Journal of Natural History 22: 537-544.
Humes, A. G. & Stock, J. H. 1973. A Revision of the Family Lichomolgidae Kossmann, 1877, Cyclopoid

Copepods mainly associated with Marine Invertebrates. Smithsonian Contributions to Zoology, 127: 1-368.
Kossmann, R. 1 877. Entomostraca (1 . Theil: Lichomolgidae). In: Zoologische Ergebnisse einer im Auftrage der

Koniglichen Academic der Wissenschaften zu Berlin ausgefuhrten Reise in die Kustengebiete des Rot hen

Meeres. IV: 1-24, pi. 1-6.
Medioni, A. & Soyer, J. 1967. Copepodes Harpacticoides de Banyuls-sur-Mer: Quelques formes recoltees sur

des Bryozoaires. Vie et Milieu, 18: 317-343.
Morton, B. S. & Morton, J. E. 1 983. The sea shore ecology of Hong Kong. Hong Kong University Press: 350pp.

Manuscript accepted for publication 6 May 1988

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. 0 565 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. Billett, A. M. Clark & G. L. J. Paterson

A morphological atlas of the avian Uropygial gland. By David W. Johnston
Miscellanea

Primed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset