/ V

BULLETIN OF

THE BRITISH MUSEUM

(NATURAL HISTORY)

ZOOLOGY

VOL. 17

1968 1969

TRUSTEES OF

THE BRITISH MUSEUM (NATURAL HISTORY)

LONDON: 1973

DATES OF PUBLICATION OF THE PARTS

No. i ...... 19 November 1968

No. 2 ...... 19 November 1968

No. 3 . . . . . 19 November 1968

No. 4 ...... 4 February 1969

No. 5 ...... 4 February 1969

No. 6 ...... 7 February 1969

No. 7 ...... 14 March 1969

No. 8 29 April 1969

No. 9 30 May 1969

Printed in England by Staples Printers Limited at their Kettering, Northunts, establishment

2 9

CONTENTS

ZOOLOGY VOLUME 17

PAGE

No. i. Notes on woodpeckers (Picidae). By D. GOODWIN I

No. 2. A review of the iguanid lizard genera Uracentron and Strobilurtis.

By R. ETHERIDGE 45

No. 3. Nigerian lizards of the genus Agama (Sauria: Agamidae). By ALICE

G. C. GRANDISON (Pis. 1-6) 65

No. 4. A revision of the amphipod genus Microdeutopus Costa (Gammaridea:

Aoridae). By A. A. MYERS (PI. i) 91

No. 5. Convergence in the structure of the head and cuticle of Euchromadora

species and apparently similar nematodes. By W. GRANT INGLIS 149

No. 6. A nomenclatural index to "A history of the British marine Polyzoa"

by T. Hincks (1880). By J. S. RYLAND 205

No. 7. The clupeoid fishes described by Bloch and Schneider. By P. J. P.

WHITEHEAD (Pis. 1-3) 261

No. 8. Cyclophyllidean cestodes from birds in Borneo. By MICHAEL D. B.

BURT 281

No. 9. Type material of the families Lysianassidae, Stegocephalidae,
Ampeliscidae and Haustoriidae (Crustacea: Amphipoda) in the
collections of the British Museum (Natural History). By M. H.
THURSTON & ELIZABETH ALLEN 347

Index to Volume 17 389

NOTES ON
WOODPECKERS (PICIDAE)

D. GOODWIN

_ BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 17 No. i

LONDON: 1968

NOTES ON
WOODPECKERS (PICIDAE)

BY

D. GOODWIN

Pp. 1-44: 10 text-figures.

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 17 No. i

LONDON: 1968

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NOTES ON WOODPECKERS (PICIDAE)

By D. GOODWIN

THESE notes are based on examination of the woodpeckers, Picidae, in the British
Museum (Natural History) during revision and re-arrangement of the collection.
The only species I know in life are the Green Woodpecker Picus viridis, the Greater
Spotted Woodpecker Dendrocopos major and the Lesser Spotted Woodpecker
D. minor; I have also briefly observed the Grey-headed Woodpecker Picus canus, the
Black Woodpecker Dryocopus martius, the Syrian Woodpecker Dendrocopos syriacus,
the Middle Spotted Woodpecker D. medius, the Hairy Woodpecker D. villosus, the
Downy Woodpecker D. pubescens, the Sapsucker Sphyrapicus varius, the Red-bellied
Woodpecker Melanerpes carolinus and the Golden-shafted Flicker Colaptes auratus
during visits to Austria and North America.

Suggestions or conclusions presented here are, therefore, based mainly on external
morphological characters. For this reason I have not suggested alterations from the
most recent comprehensive check list (Peters, 1948) except when the evidence in
favour of such a course appears overwhelming.

Section i : General notes on the family

REMARKS ON AFFINITIES, CONVERGENCE AND COLORATION

Peters (1948) divides the true woodpeckers (Picinae) into two major groups on the
basis of correlated foot and bill characters : those in which the outer hind toe is not
longer than the outer front toe, the nasal shelf narrower and the nostril near the
culmen ; and those which have the outer hind toe longer than the outer front toe, a
wider nasal shelf, and the nostril mid-way between the culmen and the edge of the
upper mandible or nearer the latter. When the woodpeckers are thus divided into
two groups two significant correlations emerge. First, most, if not all, genera,
many or all of whose members are known to be partly or entirely ground feeders, or
to feed largely by picking insects from the surface of trunks, branches or leaves or by
fly-catching, come into the first group. This is the case with Nesoceleus, Picus,
Dryocopus, Melanerpes, Asyndesmus, and Micropternus. Second, some " pairs " of
genera whose geographical distributions, colour patterns and general similarity
suggest close relationships between them Geocolaptes and Mesopicos, Micropternus
and Blythipicus, and Dinopium and Chrysocolaptes are separated. Thus it seems
likely that these bill and foot differences may be rather labile adaptive characters
and that the toe formation may have become differentiated in closely related forms
in the course of adaptive radiation.

In Europe woodpeckers divide easily into four taxonomically distinct genera
Picus, Dryocopus, Dendrocopus and Pico'ides with no troublesome intermediate
forms. The same does not hold true for most other geographical regions. Even
such visually striking differences in coloration as those between " pied " woodpeckers
and " green " woodpeckers are bridged by " pied " woodpeckers that show consider-
able yellow or green suffusion, such as Dendrocopos temminckii and Melanerpes

ZOOL. iy, I. J

4 D. GOODWIN

striatus; and by "green woodpeckers, such as Veniliornis spilogaster, which, show
an underlying pattern virtually identical to that of some of the black and white
forms. As in other groups of birds there is evidence that relatively concolorous
forms with a simplified colour pattern can derive from forms with a more complex
pattern, although in the woodpeckers most of these are still at a stage where their
affinities are obvious; for example Melanerpes herminieri and Sapheopipo noguchii.

The following general trends in coloration and colour pattern are evident. Forest
forms tend to have boldly patterned black and white colour patterns, usually with
red or yellow signal or display markings. Forms inhabiting more open woodland are
usually more cryptic ; either through a predominantly olive-green colour, at least on
the upper parts; or, when predominantly black and white in coloration, through a
barred (" ladder-backed ") or profusely mottled pattern. Forms that have become
largely terrestial tend to be more fully cryptically coloured, and show reduction or
absence of the red or yellow markings on crown or nape that are otherwise so nearly
universal in the woodpeckers. In view of the number of species whose behaviour and
ecology are little known the above remarks are, of course, tentative. On the other
hand, apparent exceptions to those general rules may prove not to be so when we have
further information. For example, the Syrian Woodpecker D. syriacus which is
black and white with a red nuchal patch in the male inhabits, at least in its
European range, rather open cultivated and settled regions, but this choice of open
habitat may be, in an evolutionary sense, a recent development. Again, the Green
Woodpecker P. viridis feeds largely on the ground through it appears colourful enough
at close quarters. When, however, it is on the ground in typical feeding sites, such
as grassy slopes, parkland, and woodland openings, its plumage is highly procryptic
and even the red on its head, which is darker and less " fiery " in tone than that
of many more arboreal species, does not show up very conspicuously. It must be
mentioned that conspicuous red, yellow or white rumps can be concealed by the
folded wings, and they usually are when their owners are alarmed, so that their
possession by such species as the Ground Woodpecker Geocolaptes olivaceus and the
Andean Flicker Colaptes rupicola, does not significantly affect the cryptic character
of the rest of their plumage.

There are widely separated allopatric forms that show considerable general
resemblance to each other. In some cases the details of their respective colour
patterns make it fairly certain that the resemblance is due to convergence. This
is so, for example, with the two ground-living woodpeckers, Colaptes rupicola of
South America and Geocolaptes olivaceus of South Africa, whose affinities are clearly
with other American and African forms respectively, not with each other. The
Chestnut-coloured Woodpecker Celeus castaneus of Central America is related most
closely to other Celeus species, and thence to other American genera, and is, obviously,
not at all closely allied to the Asiatic Rufous Woodpecker Micropternus brachyurus,
to which its rufous, black-barred plumage gives it considerable superficial resem-
blance. The small green woodpeckers of Central and South America, of the genus
Veniliornis, show such close resemblance to the African genera Dendropicos and
Campethera that I feel uncertain whether this is due to genuine close affinity or to
convergence, although I think the former less likely.

NOTES ON WOODPECKERS 5

The Ethiopian region has no arboreal woodpecker that is very large in size.
Although the smallest woodpeckers in the Nearctic, Neotropical and Oriental regions
are all about the same size, the largest arboreal African woodpeckers (Thripias)
are much smaller than the largest arboreal species in all the other woodpecker-
inhabited regions. The largest African species, the terrestial Geocolaptes olivaceus, is,
however, about the same size as, or only a little smaller than species that are equally
(Colaptes rupicola) or to some considerable extent (other Colaptes species, some Picus
species) terrestial in the Nearctic, Palearctic and Neotropical regions. Also no
African woodpecker is boldly black and white in colour although the African barbets
have produced some predominantly black and white species.

Another interesting fact is that although all other continents inhabited by wood-
peckers have several very similarly coloured " green " woodpeckers, there are no
green species in North America, where their ecological equivalents would seem to be
the brownish and barred flickers, Colaptes, and the " ladder-backed " species of
Melanerpes (Centurus).

FIG. i. (a to e) Diagrammatic outline sketches of the largest species of arboreal wood-
pecker in, respectively, the Nearctic, Palearctic, Oriental, Neotropical and Ethiopian
regions ; f shows comparative size of the smallest woodpecker species (piculets excluded)
in all of the above regions. It should be noted that the largest Nearctic species, the
Imperial Woodpecker Campephilus imperialis, is rare and restricted in range; the largest
widespread species in the Nearctic, the Pileated Woodpecker, Phloeoceastes pileatus, is
approximately the same size as the largest species in each other region, the Ethiopian
excepted.

SEXUAL DIMORPHISM

The great majority of woodpeckers are sexually dichromatic. In nearly all the
difference consists of the male having red or, less often, yellow areas on the head that
are absent in the female. This may involve only the male having red on the head or
both sexes may have heads that are predominantly red in colour but still with the

6 D. GOODWIN

female having a little less than the male. Even in species such as Colaptes rupicola and
Micropternus brachyurus, in which the male has only a little rather dull red on the
malar region, the sexual distinction is clear cut and not just a matter of the male
being slightly brighter or having his bright colours slightly more extensive than the
female's. This latter form of sexual dichromatism, common in so many birds, is
found in only a very few woodpeckers.

Only in five species are the sexes alike or nearly so. These are the Middle-spotted
Woodpecker Dendrocopos medius, the Sapsucker Sphympicus varius (the predomi-
nantly red-headed races daggetti and nuchalis) Lewis's Woodpecker Asyndesmus
lewis, the Red-headed Woodpecker, Melanerpes erythrocephalus and the Porto
Rican Woodpecker M. portoricensis . In D. medius, both sexes have the crown red,
a very common feature of males of allied Dendrocopos species. There is, however,
some sexual difference in that the red on the female is not quite so bright as on the
male and does not usually reach quite so far back on the nape where it is tinged with
yellow. This is the kind of sexual dimorphism common to most bird species in which
the sexes are usually said to be alike. It would, presumably, not give so immediate
or positive a clue to sex as the type of difference usual in woodpeckers. In the red-
headed form of 5. varius there is also a marginal tendency for the male to be more
intensely coloured, but in M. erythrocephalus there is no constant difference in brilliance
between the sexes. There does not seem any obvious or likely reason why any of
these three forms should be able to dispense with the usual type of sexual dimorphism
which, since it is nearly universal, must have, in most species, strong selective value.
D. medius is, however, sympatric with congeneric and rather similar species, and
in its case selection for specific distinctness may have operated at the expense of
sexual differentiation. One species that is in many places sympatric with it,
D. leucotos, is extremely similar in coloration and colour pattern except that the
female has a black instead of a red crown ; it is also larger in size.

Asyndesmus lewis is a rather aberrant species which differs much in feeding habits
from the more typical woodpeckers (see Bent, 1939). Its colour pattern, although
showing some connections with those of some Melanerpes species, to which it is
clearly allied, is unusual and it has no red on crown or nape. Typically the female
is marginally duller than the male. M. portoricensis shows in extreme degree the loss
of distinctive markings common in isolated island forms, being all black except for a
dark reddish tinge on the underparts. The sexes are alike.

Colaptes auratus is the only species which shows both conspicuous red markings
(a red nuchal band) and clear cut sexual dimorphism that does not involve a lack of
red in the female, the male alone having black malar stripes. Other flickers, leaving
aside for the moment the aberrant Fernandina's Flicker Nesoceleus, differ sexually
in the male having red or partially red malar stripes and the female not. In C.
pitius the red on the malar stripes of the male is reduced to a pale, dull pink on the
tips of the feathers, whose bases are blackish, so that the sexual difference in this
species is somewhat intermediate in character between that of auratus and other
flickers although less conspicuous than in either.

The only woodpeckers in which neither sex shows a trace of red or yellow in the
plumage are Nesoceleus fernandinus and the Heart-spotted Woodpecker Hemicircus

NOTES ON WOODPECKERS 7

canente. In Nesoceleus the sexes differ in the male having a black and the female a
speckled malar stripe; in H. canente the male has the forehead and crown white, the
female has a spotted forehead and crown.

Besides the colour differences described above female woodpeckers, like the females
of most other birds, often average slightly smaller in bill size than the males. In
some island forms this difference and the correlated difference in body size is consider-
able. In a recent important paper on this subject (Selander, 1966) it has been shown
that such differences are correlated with sexual differences in feeding ecology and
serve to lessen or prevent feeding competition between male and female.

The piculets, sub-family Picumninae, show " typical " woodpecker sexual
dimorphism except that in two species the " male's " colour is chestnut, not pure
red or yellow.

JUVENILE PLUMAGES

Juvenile woodpeckers, like juvenile barbets, Capitonidae, but unlike most other
birds, usually show similar bright red or yellow pigments to those of the adults. In
some species, of which the Greater Spotted Woodpecker Dendrocopos major is an
example, the juvenile male may have differently placed and more extensive red
markings than his father, and the juvenile female show red areas that are lacking in
the adult female.

Many juvenile woodpeckers have a greater number of attenuated tail feathers
similar in character to the stiff, central feathers used to prop the bird when resting or
working on a tree trunk than have the adults. The juvenile's wing may show a
reduction in size of the two innermost primaries and consequently rather more
pointed wing. These facts prompted Kipp (1956) to suggest that the juvenile
plumages of woodpeckers represented a probable future stage in their evolution as
they could, he thought, neither represent a recapitulation of a prior evolutionary
stage nor be of value to the juvenile as distinct from the adult. Verheyen (1957)
disagreed with this conclusion; pointing out that, in many other birds, the juvenile
rectrices are to some degree narrower and more pointed than the adult's and that the
reduction in size of the inner primaries was probably connected with the need for
quick growth to enable the early onset of the first moult. This was also discussed by
Sibley (1957), who pointed out that this reduction of the innermost primaries occurs in
some species only, and is almost certainly adaptive. The Stresemanns (1966) in their
comprehensive study on moult in birds agree with this conclusion.

Verheyen also emphasized, correctly, that the juvenile plumage of woodpeckers is,
like that of most other birds, usually of a weaker and more downy character than the
adult plumage. He suggests that this might be of particular disadvantage to a
woodpecker and hence one reason for the quick onset of the first moult. This, in
some species, begins before fledging, at which time the two reduced inner primaries
above mentioned, may have been replaced by adult feathers. Verheyen suggested
that the bright red or yellow pigments of juvenile woodpeckers, and the fact that in
many woodpeckers the juveniles show the same sexual dimorphism as the adults,
might be due to an original (presumably duller and less dimorphic) juvenile plumage

ZOOL. 17, I. I

8 D. GOODWIN

having been completely suppressed in the course of evolution. He considered that
the present juvenile plumage was originally the first post- Juvenal dress. I think this
is an unlikely hypothesis. Except for the bright red or yellow pigments often present,
this plumage has the typical characteristics commonly found in juvenile plumages in
other groups: a looser and more downy or woolly texture of the contour feathers;
weaker shafts of the quill feathers; a tendency for transverse barring, dark or light
shaft streaks or pale feather tips (not present or not so pronounced in adult) to be
present or more prominent and to give a more spotted or barred general appearance
and a tendency for black areas to be browner and less glossy. All these characters
are not, of course, usually found in the juvenile plumage of any one species.

The red or yellow pigments themselves may be less fully developed in the juvenile
plumage. The red is often lighter or more orange-red than that of the adult; and
in many species the red tips of the feathers are less extensive, so that the juvenile
appears to have a scattering of red speckles where the adult, if in unworn plumage,
appears uniform red. In a few cases the lighter red of the juvenile may make it
appear more brilliant. Similarly, in those species which have golden yellow instead
of red on the head, this is often of a darker and more orange tint in the juvenile. It
is possible, however, that in both these instances the juveniles may not appear more
impressive to avian as they do to human eyes.

Verheyen divided the African and Eurasian species he studied into four groups:
those in which the juveniles resembled the respective adults in their degree of sexual
dimorphism; those in which the juveniles resembled neither parent; those in which
both sexes resembled the adult male, and those in which both sexes resembled the
adult female. Most other woodpeckers fall into one or other of Verheyen's categories,
which do not, however, cover the whole range of juvenile plumages. It seems
worthwhile to describe briefly the likenesses and differences (vis a vis the adult)
found in the coloration and sexual dimorphism of the juvenile plumages, and their
generic distribution. In so-doing I shall not take account of such minor differences as
are generally characteristic of juvenile plumage. Thus a juvenile may be said to be
" like the adult " if it has the same colour pattern and general coloration, even
though it may, for example, have an orange-red instead of a scarlet cap.

It seems pertinent to digress here to mention the hazards when ascertaining the
juvenile plumage in species of which few reliably sexed juvenile specimens are
available. It has become very evident in the course of this study, that in the past
many collectors have labelled young woodpeckers as males or females according to
whether their head coloration most resembled that of the male or female adult.
With well-known European and American species this guess-sexing is immediately
obvious, but is is much less easy to detect with species represented only by one or two
specimens. Another hazard is that in some species, of which the Three-toed Wood-
pecker, Pico'ides tridactylus is an example, the feathers of the forehead and f orecrown
(sexually differentiated in the adult) are moulted very soon indeed after fledging,
so that specimens in almost complete juvenile plumage, but with adult plumage on
front and top of the head, may be much commoner in collections than birds in complete
juvenile plumage.

In general colour pattern and the (usually sexually dimorphic) head markings,

NOTES ON WOODPECKERS 9

juvenile plumages, come into one or more of the following categories. Categories 2
to 6 involve coloration of male juveniles, 7 to 9 of females, and 10 to 12 of both sexes.

(1) The juvenile male and female are like respective adults in the extent of red or
yellow on the head.

In this category are some species of Colaptes, some (possibly all) species of Piculus
and Chrysoptilus , some species of Melanerpes (cruentatus, formicivorus , candidus and
possibly some others), Trichopicus (although in this species the yellow on the throat,
which is not a sexual character, is absent in the juvenile), Celeus flavescens (most
probably other species of Celeus also), some species of Phloeoceastes (lineatus,
pileatus and, probably, schulzi and galeatus), Dryocopus, Picus (with two possible
exceptions and a tendency for restriction of lipochrome pigment in the juveniles of
the yellow-naped species), Gecinulus (females of both species and males of one),
Dinopium benghalense and D. javanense (but in the latter the front of juvenile's
forehead is brownish so that it is intermediate in this character between benghalense
and the other Dinopium species) , Chrysocolaptes (festivus only ; although the male of
validus is very similar to the adult but has yellowish brown throat and malar region),
Dendrocopos (males of all the red-crowned species and females of a very few species) ,
Meiglyptes (but juvenile males often show a reddish tinge on forehead, which is
rarer in adult males), Blythipicus (Pyrrhotis], Micropternus (but a few juveniles show
a trace of red on the nape).

(2) The juvenile male has less extensive red or yellow areas on head than has the
adult male.

This broad category is best subdivided into the following groups.

(a) Juvenile male has red (or yellow] only on the hind crown and lor nuchal region,
adult male has entire top of head, and sometimes forehead also, red or yellow. In
this group belong the Campethera species. In nubica, punctuligera and bennettii
juveniles resemble the adult female in having red only on the hind crown and nape,
the forehead and forecrown is, however, more or less dull grey or blackish, not
spotted or streaked with white as in adult female. In maculosa and permista the
differences between adult and juvenile males are similar. Our few specimens of
juvenile females of these species show less extensive red on the nuchal region than
males. In caroli and nivosa, in which the adult females has no red and the adult
male has red only on the nuchal region, the juvenile males appear to have no red
and thus resemble the female. This statement is, however, based on only one
juvenile male specimen of each species in our collection (which I have reason to
believe were reliably sexed) and on Verheyen's description of caroli. In Dinopium
rafflesii the juvenile male has the forehead brownish, shading to blackish on the
crown with a few red flecks, narrow greenish gold fringes to some of the feathers
and only the hind crown and long nuchal crest red; in D. shorii the juvenile male
has the forehead streaked dark brown and dull cream; in both these species the
adult males have the top of the head entirely red.

In Chrysocolaptes lucidus (guttacristatus and allied mainland races) the young
male also lacks red on the forehead. This species shows, however, an interesting
situation. The juvenile male at first has a brownish black forehead, unspotted or
with only obscure spots. Before the juvenile plumage as a whole is shed some of the

D. GOODWIN

forehead feathers are replaced by boldly spotted ones similar to those on the head
of the adult female. These spotted feathers may extend well onto the crown
where they have, presumably, replaced juvenile red feathers. Later these spotted
feathers are replaced by red ones but some birds show feathers intermediate
between the adult red feathers and the black and white spotted feathers, that is
red feathers somewhat intermediate in form and with a white spot. These spotted
feathers on the forehead do not seem to represent an intermediate plumage in
any true sense but they do give the owner's head some " female " characteristics.
From the available specimens it is difficult to judge whether differences are due to
age or individual variation and observations on living birds are needed.

FIG. 2. Diagrammatic sketches of heads of Hemicircus concretus to show differences in
colour pattern between adults (left) and juveniles (right) of both sexes; males above,
females below. Shaded areas are dark grey, stippled areas bright red or orange-red,
unshaded areas buff; the scale-like markings on the crest of the juvenile female represent
grey feather edges.

Hemicircus concretus shows a striking difference between the juvenile and adult
male. The latter has the entire top of the head, from forehead to nuchal crest,
bright red, the rest of its head is dark grey. The juvenile male has the forehead
and crown buff and only the nuchal crest (ends of the long head feathers) red. In
the race sordidus the adult has the nuchal area largely grey and the juvenile male
has much less red than in the juvenile of the nominate form. In both the juvenile
shows a suggestion of a buff malar stripe which is lacking in the adult. It may
be mentioned here that although the adult female's head and crest are grey the
juvenile female is like the juvenile male except that her nuchal region is buff like
her crown, not red.

NOTES ON WOODPECKERS II

(b) Red on juvenile male does not extend to, or so far down, nape as in adult male.
In this group belong only the species of Mesopicos.

(c) Adult male has head largely or entirely red; juvenile male has less red on head.
Phloeoceastes : in guatemalensis and melanoleucos , juveniles are like the adult

female, thus having black forehead and forecrown and much less extensive red
areas that adult males. In robustus (but this is surmised from only one specimen)
the juveniles also, probably, resemble the adult female. In haematogaster ,
however, in which, as in robustus, the adult female has the forecrown red, the
juveniles of both sexes have the forehead and forecrown blackish. They also
have buff instead of red rumps although both on rump and head the juvenile feathers
are soon replaced. In leucopogon the juveniles of both sexes are said (Laubmann,
1930) to have entirely black heads but this statement was based on one moulting
juvenile. In pollens, the only form in which the female has no red on the head, an
unsexed fledgling is like the adult female.

FIG. 3. Diagrammatic sketches to show difference in colour patterns of heads of adults
(right) and juvenile male (left centre) of Ipocrantor magellan icus ; the distribution of red
and black on juvenile female is similar to adult female's.

In Ipocrantor (magellanicus] the male has the head entirely red; the juvenile
male has a black head with forehead, crown, nape, malar stripes and chin red.
Both juvenile and adult females have only the chin and loral region red.

In the red-headed forms of Sphyrapicus varius (ruber and daggetti] the generally
smoky-brown juvenile has the head extensively tinged with red but this red is not
so extensive and not nearly so bright as that of the adult. So far as adult- juvenile

12 D. GOODWIN

differences are concerned the situation is the same with those races of varius in
which the adults have less red on the head.

(d) Adult male has some red on head, juvenile male has none.

In Campethera nivosa and C. caroli the adult males (but not the females) have red
on the nape, although in carola this is reduced to a scattering of red-tipped feathers.
If a male and a female juvenile of carola and a male juvenile of nivosa are r rrrectly
sexed, as I believe to be the case, then in neither of these species has the juvenile
male any red on the head. The juvenile of caroli (both sexes) also differs in
having less yellow pigment, in the top of its head being blackish brown instead of
dark green and the yellow, dark-speckled throat and eye-stripe of the adult being
replaced by pale brownish.

In the nominate form of Sphyrapicus varius the adult male has red crown and
throat, the mainly brownish (instead of black and white) juvenile male has no red
but it usually moults some or all of the feathers on its forehead and crown before it
attains other parts of the adult dress. The juvenile male of S. thyroideus has the
same black and white colour pattern as the adult male but has a white instead of a
red throat.

There appears to be only one published description of a juvenile male of
Campephilus principalis (Tanner, in Bent, 1939) . This bird resembled in colour the
adult female and lacked the red nuchal crest of the male (which extends forwards
on sides of head). It did not begin to show red feathers until it was about 3
months old. It is highly likely this is the normal situation and possible that some
of the few other juveniles seen or collected, and sexed as females were, in fact,
males also. It is also probable that the closely similar and possibly conspecific
C. imperialis agrees with principalis in this.

(3) Juvenile male has red (or yellow) on forehead and /or crown, adult male has this
colour only on nuchal region.

In this category are many species of Dendrocopos, one species (rubiginosus) of
Blythipicus and, but to a marginal degree only, Dendropicosfuscescens. In this latter
the red on the adult male's nape extends forward nearly to the crown.

(4) Juvenile male has red on crown extending a little further towards forehead than
red of adult male.

In this category come all species of Mesopicos (3), both species of Gecinulus and,
marginally, Colaptes auratus in which the juvenile male has a strong tinge of red on
the crown.

(5) Juvenile male and adult male both have red (or yellow) crown, this colour sometimes
extending further over the nuchal region in the adult.

This includes those species (many) of Dendrocopos in which the adult male has a
red crown; Mesopicos (these come into category 4 also), Thripias, Pico'ides, Dryocopus,
Chrysocolaptes; most species of Dinopium and Picus; some species of Dendropicos
and some (possibly all?) species of Veniliornis.

(6) Adult male has red only on malar region; juvenile male has red on forehead and
crown also.

Mulleripicus pulverulentus. I think it highly likely that the case is the same in
M. fuliginosus, of which I have been unable to find either juveniles or descriptions

NOTES ON WOODPECKERS 13

of them. Meiglyptes may show a suggestion of this condition, some male juveniles
being tinged with red on the forehead.

FIG. 4. Diagrammatic sketches to show colour patterns of heads of adult males (left)
and juvenile males (right) of Mulleripicus pulverulentus and M. julvus.

(7) Juvenile female resembles adult female in having no red on head, in species in which
males have red on head.

Those species of Colaptes in which the males have red malar stripes ; a few species
of Dendrocopos, including the aberrant D. albolarvatus ; some species of Picus
(squamatus and its allies), Meiglyptes, Chrysocolaptes and Dinopium and in two
related and rather aberrant species of Campethera (nivosa and caroli) .

(8) Juvenile female and adult female differ appreciably in colour of head but neither
has any red although the male has.

One species only, Hemicircus concretus, in which the adult female has the head
(and most of the body) a concolorous olivaceous grey but the juvenile has the top of
head and crest buff.

(9) Juvenile female has some red (or yellow) on head; none present in adult female.
Most species of Dendrocopos, Pico'ides, Mesopicos, Thripias, most species of

Dendropicos, Veniliornis callonotus and, in all probability, other species of Veniliornis
also.

(TO) Juveniles of both sexes have strikingly different coloration and colour pattern from
adults.

14 D. GOODWIN

Melanerpes erythrocephalus. The juvenile appears predominantly greyish brown
except on wings and tail owing to extensive pale brownish to brownish white fringes,
central streaks and/or cross bars on the blackish cover feathers. There is only a
little red around the eye, in contrast to the entirely red head of both sexes of the
adult. The juveniles of the nominate form of Spyrapicus varius (and to a lesser
extent those of other forms of the species) are very similar in general coloration (and
in the plumage patterns by which this is achieved) to the juvenile of M. erythrocephalus
described above.

Hemicircus concretus comes less definitely into this category. The colour pattern
of the juveniles (but not their coloration) is very close to the adult male's (see Text-fig.
2) but they also show some indication of a buff malar stripe which is lacking in the
adults.

(n) Adults differ in coloration and colour pattern, juveniles very like respective
adults but juvenile male lacks the red throat of the adult male.

Sphyrapicus thyroideus. This species is of interest in that the differences between
the male and female (at all ages) closely parallel the differences between adults and
juveniles in the related S. varius.

(12) Neither adults nor juveniles have red or yellow on head; juveniles resemble
respective adults (allowing for very minor differences) .

Hemicircus canente. It seems probable that Nesoceleus comes into this category
but I can find neither juveniles of this species nor descriptions of them.

The aberrantly coloured Asyndesmus lewis does not fit readily into any of the above
categories. Its juveniles are similar to but generally duller than the adults. They
lack the silver collar and upper breast, have only traces of red on the face and have
white sub-terminal spots (not present in the adult) on the feathers of the hind neck.

DISCUSSION. Thus, taking the woodpeckers as a whole, juveniles tend to be less,
although often only a little less, conspicuously coloured than the adults. Sexual
dimorphism is usually present in the juvenile plumage but is often much less marked
than in the adult plumage. In species in which the adult males or adults of both
sexes have the head almost entirely red the juveniles (where known) have appreciably
less red on the head. Juveniles, taking all species into consideration, approximate
more closely to the familial mean in the amount of red on their heads than do adults.
This is well exemplified, within a small genus, by Mulleripicus pulverulentus and
M. fulvus (see Text-fig. 4) and certainly suggests that the juvenile colour pattern
resembles a previous stage in the species' evolutionary history, as Voous (1947)
claimed for the red crowns of those species of Dendrocopos which show this feature
only in their juvenile plumage. Certainly speciation in woodpeckers has involved
both reductions and increases in the red areas of the adult plumages.

It is likely, however, that many features of the juvenile plumages may be functional
and hence of selective value. This seems obvious in Melanerpes erythrocephalus and
Sphyrapicus varius, whose much less conspicuous juvenile plumage must give some
protection against aerial predators ; both these species are partly migratory and often
inhabit relatively open, although wooded, country. Under such conditions it may
be of great advantage to the inexperienced juveniles to be less conspicuous. The

NOTES ON WOODPECKERS 15

same is probably true for some other species in which the differences between young
and adults are much less marked. In Picus viridis, for example, the more or
less speckled and barred (but otherwise similarly coloured) juveniles are even less
conspicuous when feeding on the ground than are the adults.

In the many species in which the juveniles are like the adults it is evident that
here any advantages of a less conspicuous juvenile plumage are outweighed by
others. Probably adult coloration is advantageous in the acquisition of territory or
use of feeding areas in competition with adults. On the other hand the tendency in
so many species, some of them congeneric with those showing strong sexual dimor-
phism, for the male and female juveniles to be almost alike, differing only in the female
showing less extensive red or yellow areas than those of the male, may perhaps give
other advantages which are more important for these species. It might, perhaps,
lessen any tendency that might otherwise exist for either parent to " favour "
young of one sex or it might better enable the female juvenile to hold her own in
competitive strife with her brothers. Young Dendrocopos major, for example, may
fight even in the nest hole (Sielmann, 1959). Where they are numerous, one sees
much aggression and frequent fights between fledged young before (as well as after)
they have moulted their juvenile plumage.

Where the juvenile lacks red on the head or has it more restricted than the adult it
is tempting to suggest that this might function to reduce or inhibit aggressiveness on
the adult's part, particularly in those species in which the juvenile lacks the red
precisely on those parts, forehead and forecrown, which are presented to the adult
when soliciting or taking food from it. This may be so but against this hypothesis
is the fact that in some other species, including a large number of Dendrocopos, it is
precisely these areas (forehead and forecrown) which are red in juveniles and not in
the adult. It is often implied that the juveniles of those species of Dendrocopos that
have a red nuchal patch have the entire top of the head red but in fact the young
male, although having a red forehead and forecrown, is black on the nape where the
adult male is red. The same is of course, true of the young female who has less red
on her head (as a rule) than the young male. In these species of Dendrocopos the
" replacement " of the red on the napes of adults by red on foreheads and crowns of
juveniles reaches an almost bizarre state in the Red-cockaded Woodpecker D. borealis
in which the adult male has a tiny red mark (the " cockade ") on the sides on the
hindcrown and the juvenile male an equally small red mark in the middle of his
forecrown. It is, of course, feasible that in some species some inhibition of adult
agression could best be achieved by the young not presenting red markings to the
feeding parent and in other species, in which the relative balance of fear and aggression
differed, in their doing precisely that!

As Dendrocopos major is often instanced as a supposed example of a species in which
the young are more brightly coloured that the adult it seems pertinent to digress
here to say it is, in my opinion, very doubtful if the red foreheads of the juveniles
make them more conspicuous to predators than are their parents. The same is
probably true for the other species showing this character. All the boldly-marked
black and white woodpeckers are, at least to human eyes, relatively conspicuous
as compared with olivaceous or " ladder backed " species.

ZOOL. 17, I. !

16 D. GOODWIN

Section 2 : Taxonomic notes on genera and species

MELANERPES, CENTURUS, TRIPSURUS AND LEUCONERPES

Peters (1948) included all the above genera in Melanerpes with the exception of
Leuconerpes, which he retained as a monotypic genus for the White Woodpecker
candidus. Hargitt (1890) had used a similarly broad conception of Melanerpes but
had also included candidus therein. De Schauensee (1966) retains the genus
Leuconerpes but notes that it "is perhaps not separable from Melanerpes. "
Selander & Giller (1963) give cogent reasons for treating Tripsurus as a synonym of
Centurus but say that they do not feel justified in including both these genera in
Melanerpes. However, I think that that is the most reasonable decision, unless all
three genera are to be maintained. Both in its morphological characters and what is
recorded of its ecology (see esp. Selander & Giller) Tripsurus seems perfectly to
bridge the gap (such as it is) between Centurus and Melanerpes (sensu strictu). I
provisionally maintain the genus Melanerpes as used by Peters, except as hereunder
discussed.

The species candidus does not seem to me to be sufficiently distinct to warrent
upholding the monotypic genus Leuconerpes. Its very distinct coloration, with
entirely white head, parallels that of the few white-headed African barbets of the
genus Lybius, the differences between most Lybius species and the white-headed
forms being comparable to those between candidus and related species. Wetmore
(1926) gave information on field habits of candidus, including its gregariousness (a
feature also found in some Melanerpes species) and added that the genus Leuconerpes
" in external characters . . . seems to be only slightly differentiated from Tripsurus. "
Sharncke (1931) concluded from studies of its tongue musculature that candidus
was a primitive form and closely linked with Sphyrapicus. He found, however,
several similarities between its tongue and that of Melanerpes aurifrons. Pending
further and more detailed studies it is, I think, best to include candidus in the genus
Melanerpes.

Melanerpes striatus

The Hispaniolan Woodpecker is a very distinct species which stands out among
other species of Melanerpes by reason of its red rump, greenish yellow and black
barred upperparts, yellow-green belly and, on closer investigation, its rather long
tail and the marked sexual dimorphism in length of bill. For these and other reasons
Selander & Giller (1963) argued that the monotypic genus Chryserpes should be
used for this species. Later, however, after a detailed study of the species in the
field in Hispaniola, the senior author (Selander, 1966) changed his opinion and was in
favour of including it in Centurus (Melanerpes} .

I do not know this species in life but on its museum characters I am entirely in
favour of Selander's second decision. The barred pattern of its upperparts only
differs essentially from other barred Melanerpes (Centurus) species in the greenish
yellow instead of white on the paler parts of the feathers ; this would seem to involve
only the spread of lipochrome pigments to these areas. In this connection it may

NOTES ON WOODPECKERS 17

be significant that M. superciliaris from Cuba has the white parts of the mantle
suffused with yellow and M. radiolatus from Jamaica has much dark yellow on the
lower breast and belly. The head colour pattern of striatus is rather like that of
superciliaris. Its largely yellowish green coloration gives striatus a certain resem-
blance to some species of Chrysoptilus but it shows no trace of the conspicuously
barred and spotted underparts of these. The red patch on the rump is shared by
no other New World woodpecker except Veniliornis kirkii but I do not think any of
the Chrysoptilus species or V. kirkii are very close relatives of striatus.

M. flavifrons, M.formicivorus & M. chrysauchen

The Yellow-fronted Woodpecker M. flavifrons and the Golden-naped Woodpecker
M. chrysauchen are allopatric and seem best given specific rank although they
certainly comprise a superspecies. The differences of the colour patterns of the
heads of these two are very similar to those between the North and Central American
forms of the Acorn Woodpecker M. formicivorus and the South American form
(flavigula) of that species. The differences between nominate chrysauchen from
Costa Rica and the Pacific slope of western Panama and the race pulcher from
northern Colombia may be in part or whole due to character displacement in reference
to the different races of formicivorus with which they are sympatric (see Text-fig. 5).

Melanerpes rubrifrons

Peters (1948) gives the Red-crowned Woodpecker rubrifrons specific rank but others
(Stresemann, 1924, Greenway & Griscom, 1941, Todd, 1946 & Haverschmidt, 1956)
have concluded that is a colour phase (morph) of M. cruentatus. I fully agree with
this latter opinion which is substantiated by the specimens in the B.M. collection.
Besides phenotypically " pure " specimens of each form there are several that are
intermediate, to varying degrees.

The African Woodpeckers

All the sub-Saharan African woodpeckers seem to be more closely allied to one
another than to any non- African species or genus. They are, I think, representatives
of the stock ancestral also to Picus and Dendrocopos. Adaptive radiation has,
however, resulted in similar but less marked divergences to those shown by wood-
peckers in Eurasia and America. The second possibility: that the African forms are
polyphyletic, with Thripias and Dendropicos closer to Dendrocopos, and Mesopicos
and Campethera to Picus, is, I think, rather less likely. In spite of some remarkable
general resemblances in coloration I think that the South and Central American
green woodpeckers are more closely related to other New World forms, in the genera
Colaptes and Melanerpes, than they are to African forms. When comparing, for
example, species of Dendropicos and Veniliornis that look much alike, one finds that
some details of their colour patterns differ from each other and are, respectively,
closer to those of other African and South American species.

There have been divergencies of opinion as to how many genera of African wood-

i8

D. GOODWIN

FIG. 5. Sketches to show colour patterns of heads of males (left) and females (right) of
Melanerpes formicivorus (top) which overlaps in range with M. chrysauchen (second row)
in Central America and of the form of M. formicivorus (third row) which overlaps with
M. chrysauchen in northern South America. (See text).

peckers to recognize. Hargitt (1890) recognized the genera Campethera, Dendropicos,
Mesopicos, Thripias and Geocolaptes, mainly on structural characters with emphasis on
shape and size of bill. Other authors, notably Chapin (1939) recognized further
genera but Mackworth-Praed & Grant (1957) followed Hargitt except for resurrecting
the genus Ipophilus for the two species obsoletus and stierlingi. Peters (1948) put
elliotii and johnstoni (now reckoned as a race of elliotii) in the genus Polipicos
(following Chapin) but otherwise recognized the same genera as Hargitt and Praed
and Grant although not with precisely the same allocation of species within them.
I have followed Peters' nomenclature except where otherwise stated.

NOTES ON WOODPECKERS 19

DENDROPICOS

Peters includes in this genus fuscescens, stierlingi, elachus, abyssinicus, poecilolaemus ,
gabonensis and lugubris. I concur with him in thinking that these species are probably
more closely allied to each other than any is to species in another genus. This
assumes, as I think is likely, that some differences of general coloration such as a
tendency towards light, barred or spotted plumage in dry savannahs; darker,
greener coloration in forests and the acquisition of a red rump have arisen indepen-
dently in different (but related) stocks.

Their head markings suggest the Dendropicos species are nearer to Thripias and
Mesopicos than to Campethera. The extreme similarity between Dendropicos
lugubris and Campethera nivosa is, I think, due to parallel development but it may
indicate true affinity in spite of differences in bill shape.

" Dendrocopos " obsoletus

Most writers on African woodpeckers (Bates, 1937, Chapin, 1939, Hargitt, 1890,
Peters, 1948, Voous, 1947) put the Brown-backed Woodpecker, obseletus, in the genus
Dendrocopos (or in Yungipicus which is now usually regarded as a synonym of
Dendrocopos) but Grote (1928) and Bannerman (1933) placed it in Dendropicos.
Voous in his monograph on Dendrocopos says that Grote and Bannerman had
" abusively referred (it) to Dendropicos " and adds that this latter genus is quite
distinct in having reddish or yellowish shafts to rectrices or remiges and usually in
addition a distinct greenish tinge of the plumage. He considered, however, that
stierlingi was correctly referred to Dendropicos. Mackworth-Praed & Grant, 1957,
place obsoletus and stierlingi together in the genus Ipophilus.

I think that Bannerman and Grote were right to include obsoletus in Dendropicos.
Among them the Dendropicos species show varying amounts of green coloration and/
or yellow or reddish quill shafts. In two of them, stierlingi and elachus, this is
reduced to a not very pronounced yellow tinge in the shafts of the wing and tail
quills (stierlingi) or tail quills only (elachus). If either of these two species are com-
pared with any other Dendropicos species it will be seen that the " gulf " between
them, so far as yellow pigmentation is concerned, is greater than that between
either one of them and obsoletus.

Voous links obsoletus with the pygmy woodpeckers of south Asia, Dendrocopos
nanus and its allies, but in plumage pattern obsoletus is closer to Dendropicos species
than it is to any of the Asian forms. In its coloration it is nearest to elachus although
it lacks the latter's scarlet rump which is, in all probability, a species recognition
mark serving to isolate obsoletus and elachus in the areas where their ranges overlap.
Such species as obsoletus and elachus undoubtedly indicate the close relationship
between " green " and " pied " woodpeckers or the potentiality for " green " and
" pied " forms to evolve, in different geographical regions, from a common stock. I
think, however, there is little doubt but that the true affinities of obsoletus are, as
Bannerman and Grote decided, with Dendropicos.

20 D.GOODWIN

Campethera bennettii, C. scriptoricauda & C. nubica

Benson (1952) gives evidence for the conspecifity of Bennett's Woodpecker
C. bennettii and the Tanganyika Woodpecker C. scriptoricauda. In C. bennettii
vincenti (Grant & Mackworth-Praed 1953) the throat colour of females is intermediate
between the chocolate brown of bennettii and the creamy white of scriptoricauda,
although it does not have the black spots of the latter. Some males of vincenti also have
the ear coverts a less pure white than those of other forms of bennettii. Also a female
scriptoricauda, from Blantyre, shows some approach to bennettii in colour, having
fewer and smaller spots than usual, on a beige-tinted throat. It is unlikely that these
specimens merely indicate occasional hybridization between two " good " species as
there seem to have been no " pure " specimens of either bennettii or scriptoricauda
collected within the range of vincenti and, more importantly, Mr Benson informs me
(pers. comm.) that the calls and ecology of bennettii and scriptoricauda do not differ.
I therefore agree with him that, on present evidence, scriptoricauda is best considered
as a race of bennettii.

The very distinctive chocolate and white markings on the face and throat of the
females in most forms of bennettii and, possibly, also the very white cheeks and
intense red malar stripes of the male, probably function as isolating mechanisms
in reference to C. abingoni, with which the chocolate-throated forms of bennettii
overlap widely in range whereas scriptoricauda (and C. nubica} overlap only marginally
with abingoni.

The Nubian Woodpecker C. nubica does not seem to differ in any significant
plumage character from C. bennettii scriptoricauda; they differ only in the unspotted
throat of nubica. Localities of specimens in the British Museum collections do not
suggest any actual overlap of distribution. I provisionally maintain them as
separate species, however, because Mr. C. W. Benson informs me that their calls differ,
or at any rate those calls most usually uttered, and they differ in ecology.

Campethera nubica & P. punctuligera

I provisionally treat the Nubian and the Fine-spotted Woodpeckers, nubica and
puctuligera, as members of a superspecies but I think it likely that they may prove to
be conspecific. Although their respective ranges come close together in the southern
Sudan and the eastern Congo I can find no evidence of both having been taken or
identified in the same area (see also Chapin, 1939).

The form of punctuligera, C. p. balia, from the south-eastern Sudan and eastern
Congo is largely intermediate in colour and colour pattern. The differences between
the two forms are, chiefly, the size and formation of the spots on the breast and the
presence (in punctuligera} or absence (in nubica} of spotting on the throat. One
female of balia from Baginzi, Bahr el Gazal (B.M. No. 1919. 10.7.23.) comes closer to
nubica than most, having no spots on the throat, very dull underparts (less yellow tinge
than usual in punctuligera} with the breast spots closely approaching those of nubica
in pattern and general appearance although still much smaller than those of nubica.
A juvenile from Angba, on the river Welle, also much resembles nubica in its breast
spots.

NOTES ON WOODPECKERS 21

Chapin (1939) says that the calls and habits of punctuligera and nubica are extremely
similar and that " adherents of the Formenkreise theory " would surely consider
them as forms of one species. A main difficulty and the one that chiefly prompts me
still to treat them as species is that in appearance punctuligera could equally well
be a form of bennettii, with which it is also allopatric. In fact there is an even closer
resemblance in colour pattern between punctuligera and bennettii scriptoricauda than
between the former and nubica. Present evidence suggests differences between the
calls of bennettii and those of scriptoricauda but this negative evidence is of no great
significance until the vocalizations of all three forms have been recorded in more
detail. I suspect they may prove all to be members of a superspecies or even
conspecific.

Campethera notata

I think Clancey (1958) is, right in suggesting that the Knysna Woodpecker notata
and the Golden-tailed Woodpecker abingoni are possibly conspecific and certainly
members of a superspecies. In spite of their great similarities, however, the pattern
of the breast spots in notata is closer to those of nubica, bennettii and punctuligera,
with none of which is notata sympatric, than to abingoni although such breast spots
as those of notata could easily drive from the streaks of abingoni or vice versa.

Campethera cailliautii, C. taeniolaema & C. maculosa

These three species are all very closely allied, the Little Spotted Woodpecker,
cailliautii and the Fine-banded Woodpecker taeniolaema being mainly allopatric.
Their ranges overlap in the eastern Congo and adjacent areas where they appear,
however, to be ecologically isolated, taeniolaema inhabiting the highlands and
cailliautii the lowlands. They are very similar in coloration but cailliautii has the
throat and face buffish with blackish, speckly-looking, barring whereas taeniolaema
has the face and throat greenish white very finely barred with greyish olive.

From the Golden-backed Woodpecker maculosa, with which it is sympatric,
cailliautii differs in the male having a slightly brighter red crown, greener and les
gold-tinged upperparts and less boldly spotted on the throat. The females differ
rather more markedly in maculosa lacking the red nape of cailliautii; she also has her
entirely blackish crown spotted with deep buff, the spots on the head of cailliautii
being pale buffish to white.

Campethera tullbergi

The rather distinct Tullberg's Woodpecker appears to be closest to cailliautii
(permista) with which it is sympatric. The carmine patch on the carpel area of the
wing probably serves as a species-specific recognition mark in reference to cailliautii.
I fully concur with Peters (1948) that C. wellsi should be considered a race of tullbergi.

Campethera nivosa

The Efulen race of the Buff-spotted Woodpecker, efulensis Chubb, appears to be
inseparable from nominate nivosa. One of three specimens from the type locality

22 D. GOODWIN

(Efulen, Cameroon) can be matched by a specimen of nivosa from Portuguese Guinea.
The large series from Kumba, Cameroons, vary from typical " nivosa " to typical
" efulensis " in colour. The races poensis and herberti are slightly differentiated but
recognizable.

MESOPICOS

The Grey Woodpecker M. goertae and the Olive Woodpecker M. griseocephalus are
very closely related and could certainly be considered as members of a superspecies or
even as conspecific but for the fact that their ranges overlap in parts of Kenya and
north-eastern Tanzania. We have, however, no records of both species from precisely
the same place although there are specimens of both from north-eastern Tanzania,
goertae from Nabara and griseocephalus from Same. In and near the area of apparent
overlap they appear to be ecologically separated, griseocephalus being confined to
highland forest.

The only absolute colour differences between the adults are the darker grey head
and golden-olive breast of griseocephalus. In goertae juveniles of both sexes have
red on the head, the male usually rather more than the female. Some works on
African birds imply that in griseocephalus the young are, except for being duller, like
the respective adults. There are only six juvenile griseocephalus in the British
Museum collection, some unsexed, others sexed as males. AU of them have red on
the head ; in some of them the red is less extensive than in others and I suspect these
may be females. Juveniles of griseocephalus are, incidently, paler and greyer in
colour than adults and thus come even closer to goertae in appearance than do the
adults.

A few remarks on geographical variation in both species: From West Africa
(Senegal, Gambia, etc.) across to Uganda and the southern Sudan M . goertae shows no
striking differentiation, if allowance is made for individual differences (from the same
area some birds may have an orange-red belly patch and others not) and those due
to wear (which are often considerable). Within this region there is a tendency for
birds from the more northern arid areas to be palest and to have more distinct
barring on the wings and upperparts, those from the southern areas to be darker and
duller. Specimens from Gambia to Uganda tend to have little or no orange-red,
only yellowish, on the belly. Those from the extreme north-east of this area (parts of
Southern Sudan) are more like those of Senegal in coloration than are the latter to
Gambian birds!

I think it best to recognize three races within the above areas : nominate goertae from
Senegal, the paler konigi (Neumann, 1903) from the Lake Chad regions, parts of
French Niger and French Sudan and centralis (Reichenow 1900) and agree with
Mackworth-Praed & Grant (1957) in thinking agmen (Bates, 1932) and oreites (Grote,
1923) best treated as synonyms of centralis. This is an admittedly wide but I think
workable interpretation of centralis but it must be borne in mind that no hard and
fast line can be drawn between any of the above-mentioned forms of goertae. To
the east of its range centralis intergrades with abessinicus (Reichenow, 1900).

M. griseocephalus persimilis (Neumann, 1933), described from the Highlands of

NOTES ON WOODPECKERS 23

Angola (Benguella) was recognized by Peters but has since been commonly treated as
a synonym of ruwenzori (Sharpe, 1902). In the original description persimilis was
separated on its smaller size. Besides the rather slight size difference our specimens
from Angola, southern Congo and Tanzania mostly show quite conspicuous paler
barring on the flanks which is absent from specimens of ruwenzori from Ruwenzori
and Lake Kivu. I have, therefore, recognized persimilis.

Mesopicos elliotii

Chapin (1939) & Peters (1948) both use the genus Polipicus Cassin for Elliot's
Woodpecker which was previously (Hargitt, 1890) and subsequently (Mackworth-Praed
& Grant, 1957) included in Mesopicos. In structure of bill elliotii (with which johnstoni,
formerly given specific rank, is now reckoned conspecific) is nearer to Mesopicos than
to other African genera but in its other external characters it seems to be as close to
Campethem. The two genera are, however, closely allied. It seems preferable to
leave elliotii, at least provisionally, in Mesopicos rather than to place it in a monotypic
genus.

Mesopicos elliotii sordidatus (Bates, 1928) was described from a juvenile from Oku,
west of Kumbo, Cameroons. Its supposed racial characters are merely those of the
juvenile plumage. There are now two adults from Oku in the British Museum
collection, these do not differe from examples of M. e. johnstoni of which I therefore
consider M. e. sordidatus a synonym.

THRIPIAS

Peters (1948) includes the Bearded Woodpecker namaquus, the Fire-bellied
Woodpecker pyrrhogaster and the Golden-crowned Woodpecker xanthocephalus in
the genus Thripias. Many other authors, e.g. Hargitt (1890), Chapin (1939) and
Grant & Mackworth-Praed (1957) have put Pyrrhogaster and xanthocephalus in
Mesopicos, presumably mainly on account of their proportionately shorter wings
(than those of namaquus) and maintained Thripias as a monotypic genus.

I think Peter's arrangement is best. These three species all agree in having a
relatively long and straight bill, black nuchal region in both sexes and similar
characteristic facial markings. Against these I do not think that the proportionately
longer bill and tail of namaquus warrants its generic separation. T. namaquus seems
to link the other two Thripias species (as reckoned here) with Campethera rather than
with Mesopicos. In this connection it may be mentioned that the female of
Campethera bennettii has facial markings rather similar to, although not identical
with, those of the Thripias species.

Geocolaptes olivaceus

I think the Ground Woodpecker is most closely related to other African forms and
that its resemblance to some of the American flickers, Colaptes, (near which it is
usually placed in systematic lists,) is due to convergent adaptations for terrestial or
semi-terrestial life. It differs from the Colaptes species but very closely resembles

24 D. GOODWIN

Mesopicos goertae and M. griseocephalus in colour pattern except for the barred tail
and the vestigial red malar stripes of the male. Both these are, however, very
widespread characters in " green " woodpeckers and occur, inter alia, in some
Campethera species.

It seems likely that olivaceus evolved from some form ancestral both to itself and to
goertae and griseocephalus. It may well have retained (or developed) red malar
stripes as a species-specific recognization mark in reference to griseocephalus (or its
ancestor). The present apparently obsolescent state of the malar markings might
indicate that the development of greater divergence in size and ecology has now
obviated the need for such distinguishing markings between these two forms.

Notes on Picus

Peters (1948) includes in Picus the former genera Chrysophlegma species mentalis
andflavinucha, and the monotypic Callolophus species mineaceus. Hargitt (1890)
included the latter in Chrysophlegma also although he placed the smaller yellow-
crested forms, puniceus, chlorolophus and chlorigaster with the " typical " green
woodpeckers in the genus Gecinus. Within Peter's broad and inclusive conception
of Picus the species fall into three rather distinct groups which are not precisely
the same as those suggested by former allocations of species between Gecinus
(or Picus} and Chrysophlegma. The latter genus was defined as having " wing
rounded . . . nasal ridge almost obsolete, culmen ridge very blunt, tail two thirds
length of wing " and later by Stuart Baker as having "... bill more curved and nasal
ridge almost obsolete, well-developed nuchal crest. " These definitions have,
apparently, restrained those who recognized Chrysophlegma from including in it the
species chlorolophus chlorigaster and puniceus, which have rather straighter culmens
that the other yellow-crested species.

The group of most typical Picus species comprises viridis, canus, vaillantii,
awokera, viridanus, vittatus, squamatus, xanthopygaeus , erythropygasus and rabieri. All
these are relatively long-billed forms with fairly straight culman, nuchal crest absent
or slight, primaries marked whitish and black or grey, and except in one race of canus,
no red or chestnut colour on wings. P. rabieri has a more rounded wing and rather
shorter bill than the others and approaches the next group in these features.

The species in the second group are characterized by having yellow nuchal crests,
shorter bills and, one species excepted, red coloration or at least a reddish tinge, on
the wings. They comprise flavinucha and mentalis, which in addition to the characters
listed above also have black and chestnut primaries and a spotted patch on the throat,
and puniceus, chlorolophus and chlorigaster which have rather straighter culmens, no
chestnut on their primaries which are dark and whitish, varying amounts of red
on the head, and are smaller in size. To some degree these three forms link the
original " Chrysophlegma " species to the first group but they are, I think, much more
closely allied to the former.

The third group consists of the single species mineaceus. This has a long and
broad occipital crest, relatively short, broad bill, barred plumage pattern both above
and below and rather harsh, coarse-feeling plumage. Although it may be justifiable

NOTES ON WOODPECKERS 25

to include mineaceus in the present enlarged conception of Picus, (and certainly this
is more reasonable than allying it with flavinucha and mentalis and yet not putting
puniceus, chlorolophus and chlorigaster in the same genus), it is a very distinct species.
It appears to be a link between Picus, via Picus puniceus, and the Rufous Woodpecker
Micropternus to which it shows some approach in proportions, coloration, plumage
pattern and plumage texture.

I think it possible that more comprehensive studies might indicate that the species
in this second group are less closely allied to those in the first group than are the
latter to such Ethiopian genera as Campethera and Mesopicos.

Picus canus

In the eastern part of its range, where it and viridis do not overlap, the Grey-headed
Woodpecker canus much more nearly approaches the Green Woodpecker viridis
in size and proportionate length of bill. The only similarly-sized green woodpeckers
which it overlaps in eastern Asia are P. erythropygius and P. rabieri, which are very
distinct from canus, with striking colour differences. Picus dedemi of Sumatra
seems rightly put as a form of canus in spite of its different colouring, with bronzy
carmine, with a hint of green, replacing the green and olive of canus. The racial
variation within the Paleartic populations of canus has been very fully discussed by
Vaurie (1959).

Picus vaillantii

Vaurie (1965) put North African Green Woodpecker vaillantii, as a race of viridis
and earlier (1959) anticipated this decision on the grounds that in everything except
the black malar stripe of the male, which he considered an " alternate character "
to the red and black malar stripe of the male viridis, P. viridis sharpei of the Iberian
peninsula " bridges the gap " between viridis and vaillanti. He pointed out
vaillantii had already been listed as conspecific with viridis by Goutenoire (1955) and
that Dr. David Snow, who has observed this species in the wild, was also in favour of
this decision.

I cannot, however, see that sharpei is intermediate between [other races of]
viridis and vaillantii except in bill shape and I do not think that minor details of
size, size of bill or tone of general colour, in all of which vaillantii comes nearer to
viridis than it does to European forms of canus, are of much significance when dealing
with a geographically isolated " green woodpecker ". Especially in view of the
amount of geographic variation shown by both viridis and canus and the obvious
close relationship of the above three forms and the Japanese Green Woodpecker
P. awokera. The female of P. viridis sharpei has the feathers of the forehead and
crown red (or, to be more precise, these areas all tipped with red) as in other races of
viridis, whereas these areas are without any red in females of vaillantii. Incidently,
the red tips on the nape feathers of vaillantii, appear longer than and somewhat
different in texture from those of viridis or canus.

While vaillantii may be more closely allied to viridis than it is to canus or awokera,
the differences in colour patterns of their heads, lack of red on malar stripe of male
vaillantii and black and grey instead of red forehead and crown of female, might be

26

D. GOODWIN

FIG. 6. Diagrammatic sketches to show colour patterns of heads of males (left) and
females (right) of Picus viridis (top, Picus vaillantii (middle) and P. canus (bottom).

pre-adapted to act as isolating mechanisms should the two ever overlap in range.
Of its more striking plumage characters vaillantii shares one (crown and nape of male
red) with viridis, two (no red on malar stripe of male, forehead and crown of female
grey or grey and black) with canus and two (red on nape only of female, forehead and
crown of female grey or black and grey) with the more geographically distinct
awokera. I think it best, therefore, to give vaillantii specific status although it can,
at least provisionally, be put in the same superspecies as viridis.

Picus rabieri

This is a very distinct species but, in spite of its rounded wings and somewhat
different colour pattern, I think it is an offshoot of the viridis group although in wing
shape and in its rather short bill it shows some approach (convergence) towards the
flavinucha group.

NOTES ON WOODPECKERS 27

Picus squamatus, P. viridanus, P. vittatus & P. xanthopygaeus

These species form a closely related group and are essentially alike in coloration
and colour pattern. The Red-rumped Woodpecker P. erythropygius is related to
this group, as its colour pattern especially in the eye-striped morph, clearly indicates.

The Scaly-bellied Green Woodpecker P. squamatus overlaps in part of its range
with the Little Scaly-bellied Woodpecker P. xanthopygaeus from which it differs in
being much larger, having very conspicuous black and white malar stripes, the upper
breast plain greyish green without squamate markings and conspicuously barred
central (and other) tail feathers.

P. xanthopygaeus overlaps with the Burmese Scaly-bellied Woodpecker P. viridanus
and marginally with the Laced Green Woodpecker P. vittatus from both of which it
differs in having inconspicuous (almost obsolescent) malar stripes, more streaked
(longitudinal squamate markings) throat and yellow or greenish yellow instead of
yellowish green rump and upper tail coverts.

P. vittatus and P. viridanus are extremely similar, differing in appearance only in
viridanus having the streaky-looking squamate markings of the underparts extending
forward onto the upper breast and throat where they are, however, less clearly
denned than lower down. As given in many standard works the ranges of the two would
appear to overlap in parts of Eastern Burma and south-western Siam. In the fairly
extensive series of both in the British Museum (Natural History) there are no
specimens of both species from any one area with the exception of one specimen of
viridanus, allegedly from Tenasserim, where only vittatus normally occurs. I think
this probably represents some error in labelling.

Robinson & Kloss (1923) say that " where the ranges of the two species touch or
approach there is not the slightest sign of intergradation ". In view of the extremely
slight differences between them I do not find this statement very convincing. There
are two specimens in the British Museum, No. 1887.8.10.1556 from Mergui and
No. 1887.11.1.95 from " Burmah ", which are absolutely intermediate between
" typical " vittatus and viridanus at first appearance. On closer examination they
are seen to have squamate markings on the upper breast although these are fainter
than usual in viridanus. It is significant, perhaps, that in all specimens of viridanus
these squamate markings are least developed on the upper breast and throat, contrary
to what one might expect if they functioned as a species recognition mark in reference
to vittatus.

Deignan (1963) puts viridanus as a race of vittatus and, although he does not there
give his reasons for so-doing, all the evidence I have been able to find suggests that
he is quite right in this decision.

Picus chlorolophus & P. chlorigaster

The number of races of the Small Yellow-naped Woodpecker chorolophus that
has been described tends to obscure the main difference which is between the popu-
lations in peninsular India and Ceylon and those from elsewhere. The former, at
one time given specific status as P. chlorigaster, are not only geographically isolated but
in some characters come closer to the related Crimson-winged Woodpecker P. puniceus

28 D. GOODWIN

resembling it and differing from typical chlorolophus in having blackish lores, instead
of whitish as in chlorolophus, and the top of the head entirely red. In the amount
of red on their wings they are intermediate.

The three forms, chlorigaster , chlorolophus and puniceus are allopatric except that
an isolated race of chlorolophus (rodgeri) occurs in the mountains of Perak. There is
thus a very limited geographical overlap of this form with puniceus although there
is apparently no ecological or altitudinal overlap (Robinson, 1928). This montane
race is rather dark in colour and thus superficially somewhat intermediate between
chlorolophus and chlorigaster but in its main features it agrees with the former, with
which its affinities clearly lie. I think it is preferable to treat chlorigaster as a member
of a superspecies, together with chlorolophus and puniceus, rather than as a race of
chlorolophus.

DENDROCOPOS

Voous (1947) shows that the American and Asiatic " ladder-backed " woodpeckers
in this genus all inhabit areas that were not covered with Pleicestocene land-ice and
which, in North America, represent the glacial forest refuges. He considers all
these ladder-backed species to be (relatively) closely related in spite of their present
discontinuous ranges.

FIG. 7. Diagrammatic sketches to show colour patterns of heads of a typical Old World
Dendrocopos (major], a typical New World ditto (villosus) and Pico'ides trydactylus (left
to right) (In all species the heads shown are of females).

All American ladder-backs agree with other American Dendrocopos species, and
differ from most Old World Dendrocopos species, ladder-backed and otherwise, in
lacking red on the ventral regions or under tail coverts. They also have a similar
head pattern, with a dark post-ocular band, to other American Dendrocopos species.
I think these facts indicate that all the New World Dendrocopos species are more
closely related to each other phylogenetically than any one of them is to any Old
World form.

It seems more likely that the " ladder-back " pattern (so common in woodpeckers
generally) has developed, or been retained if it was a primitive feature, independently
in both Old and New Worlds, as an adaption to relatively open or well-lighted
habitats in which it probably has protective value, than that some factor in the
American environment has caused two or more phylogenetically distinct Dendrocopos
stocks to lose all trace of red on the underparts and to acquire similar facial markings
subsequent to their arrival in America. In this connection it is pertinent that in
some species of Melanerpes the adults are plain backed and the juveniles ladder-backed

NOTES ON WOODPECKERS

29

and in the Three-toed Woodpecker Picoides tridactylus some races are ladder-backed
and others not (see Text-fig. 8).

Of Old World species the " dwarf " woodpeckers, D. minor, D. kizuki, D. nanus
et al., come nearest to the American forms in their colour patterns and are, I think,
closest to them phylogenetically.

FIG. 8. Diagrammatic sketches to show variation in back pattern within a species:
Females of Pico'ides trydactylus from Sweden (left), South-eastern Tibet (middle) and
eastern North America (right).

The Hairy Woodpecker D. villosus and the Downy Woodpecker D. pubescens are
now very largely sympatric. They presumably derived from a common ancestor
at a relatively recent date as their colour patterns are virtually identical. They also
show similar geographical variation although this, by itself, is of course, no argument
for close relationship.

Nuttall's Woodpecker D. nuttallii and the Cactus Woodpecker D. scalaris are
also close relatives. They appear not to be sympatric and descriptions of their calls
do not suggest any great difference of voice. They may prove to be conspecific but
provisionally are, perhaps, best treated as members of a superspecies. The Red-
cockaded Woodpecker D. borealis is, I think, closely allied to them. Its strikingly
different facial pattern seems to have been achieved by the extreme reduction of
the usual (in American Dendrocopos) black post-ocular stripe so that only a vestige
of this marking remains. I think, however, that the great similarity in position
and extent of the red on the head in this species and the Old World D. kizuki is due to
convergence, within related stocks, and does not indicate that borealis is closer to

30 D. GOODWIN

kizuki than are other American species. The juveniles of borealis and kizuki do not,
apparently, resemble one another in position of red on the head as kizuki is said to
resemble the adult in this respect. The Arizona Woodpecker D. arizonae and
Strickland's Woodpecker D. stricklandi are allopatric, they are close to one another
in appearance, voice, behaviour and ecology (Davis, 1965). On its upperparts
stricklandi has a colour pattern intermediate between that of completely ladder-
backed species and those, such as arizonae, with uniform dark back and rump.
Individuals of arizonae which show traces of a barred dorsal pattern do, however,
occur (Davis, 1965). The two are otherwise similar in colour but differ in stricklandi
having dark streaks on the underparts with some indication of barring on the flanks,
whereas arizonae is spotted, but the differences are less impressive when the markings
on the individual feathers are compared. I think arizonae and stricklandi are best
considered members of a superspecies.

The two South American forms the Striped Woodpecker D. lignarius and the
Chequered Woodpecker D. mixtus differ from all North American Dendrocopos
species in having their central tail feathers strongly barred with white and they are
generally lighter and more " spotty " in appearance. They show much resemblance
in plumage pattern to Veniliornis spilogaster and I think this may indicate close
relationship between Veniliornis and Dendrocopos. They agree, however, with the
northern forms in general overall colour pattern. From their visual characters one
would certainly suspect them of being conspecific but they apparently overlap in
range, both being found in Nequen and the eastern Rio Negro in Argentina (Peters,
1948, Olrog, 1963) . Two specimens in the British Museum from Cordova, Argentina,
(B.M. Nos. 1889.2.26.112 and 113) appear to be intermediate between D. lignarius
and D. mixtus berlepschi. The male of the two agrees with mixtus in the extent of
the red on its head but both are closer to lignarius in plumage pattern, which is,
however, extremely similar in the two species (?) in any case. These two have longer
bills than our specimens of berlepschi but we have only two of the latter and, at least
in lignarius, this feature seems prone to individual variation.

The White-headed Woodpecker Dendrocopos albolarvatus is rather aberrant in
colour pattern and its tongue is said (Voous, 1947) to be less extensible than in other
(all other?) Dendrocopos species. Its external features, particularly the typically
dendrocopine correlation of a red nape in the adult adult male with a red crown and
black nape in the juvenile indicate that Peters was right to include it in Dendrocopos.

Dendrocopos darjellensis

Measurements of our specimens seem to indicate a less clear-cut difference between
nominate specimens of the Darjeeling Pied Woodpecker, darjellensis, and
D. darjellensis desmursi (]. Verreaux) than did those measured by Vaurie (1959)
We have, however, no topotypical specimens of desmursi from Sikang and only a few
from Yunnan. Our only specimen of fumidus, from the Naga Hills, does not show
the characters claimed for this race (Ripley, 1951) and thus tends to confirm Vaurie 's
opinion that fumidus should be considered a synonym of darjellensis.

This species overlaps widely with the Crimson-breasted Pied Woodpecker

NOTES ON WOODPECKERS 31

D. cathpharius. The two are nearly identical in coloration but differ much in size.
Their yellow-tinged cheeks and underparts, red breast bands and heavy streaking
on the underparts are considered by Voous to be primitive characters. Whether
this is so or not these characters might function now as species-specific signal markings.

Dendrocopos leucopterus

I concur with Voous (1947) and Vaurie (1959) in thinking that it is better to give
leucopterus, provisionally, specific rank. This, however, is mainly on grounds of
convenience. I think Vaurie has rather over-emphasised the differences between
leucopterus and major. I have seen no juvenile specimens of leucopterus so do not
know if I should think them so distinct from juvenile major as they are said to be. It
seems, however, pertinent to remark that although, as Vaurie says, juvenile major
of both sexes have red on the crown, that on the female is normally less extensive
than the male's. So the fact of the juvenile female of leucopterus having (always?)
an entirely black crown may not be of specific significance.

Dendrocopos dorae

Although at first glance the Arabian Woodpecker is rather like some of the species
in the African genus Dendropicos it seems most closely allied to the Eurasian forms
of Dendrocopos and is now rightly included in that genus.

I think that dorae may be closest to medius ; to which it bears much resemblance
when allowance is made for " fading " of the black areas to darkish brown (already
often shown to some extent in some individuals of medius) and for reduction of the
white areas. In size, texture and colour of the red crown feathers, and the streaking
on its flanks it is very like medius. Indeed, the differences between these two
woodpeckers are very similar in kind, although greater in degree, to those between
the Arabian form of the Magpie Pica pica and its more northerly representatives. In
dorae, like most Dendrocopos species but unlike medius, the female lacks red on the
head but I do not think this feature of medius, which is unique within the genus,
argues against their close relationship. D. medius and D. leucotos are unquestionably
close relatives, but differ in this feature as they do in size and bill shape.

Dendrocopos analis, D. atratus & D. macei

I provisionally follow Peters & Deignan (1945) in treating analis as a form of the
Fulvous-breasted Pied Woodpecker macei. They differ slightly in size, in analis
having a spotted rather than a streaked breast, white ground colour on underparts
and central tail feathers and upper tail coverts barred with white instead of uniformly
black. They do not, however, differ in colour pattern of the head. Intermediates
occur (fide Deignan, 1945) in the Chiang Mai area of Siam.

Voous (1947) suggests that macei and the Stripe-breasted Pied Woodpecker
atratus may be conspecific, quoting Standford & Ticehurst (1939) that where they
appear geographically to overlap in range in northern Burma macei is a lowland and

32 D. GOODWIN

atratus a highland form. He mentions specimens of macei obtained by Hume in
Manipur, Assam and implies that these showed some intergradation with atratus.

At least some of these Manipur specimens are in the British Museum collection
and do not seem to differ from other specimens of macei, collected elsewhere. When
they are compared with specimens of atratus, also collected in Manipur, the differences
between them are at least as great as between specimens of both from elsewhere.
That is, macei differs in being smaller, having an unstreaked lower throat, the
underparts creamy rather than dusky yellowish and with not very prominent dark
streaks, as against very prominent dark streaks in atratus. The red on the ventral
region does not usually extend so far forwards, the red of the head is a little less
bright and the red feathers are a little dissimilar in texture. These differences also
hold for analis, which has, in addition, tail and rump feathers barred with white.
D. analis and D. atratus overlap widely, geographically, but may be ecologically
isolated. Thus some of the differences between analis and macei might have
originated as species-specific recognition marks in reference to atratus.

Dendrocopos major

Vaurie (19590) puts the race parroti (Hartert) from Corsica as a synonym of the
Sardinian form harterti (Arrigoni) . I prefer to recognize parroti as there appears to
be very little overlap in bill length and the bills of all our specimens of parroti are
noticeably more massive than those of specimens of harterti of the same sex. In
view of the only slightly differentiated and intergrading continental races of this
woodpecker that are currently recognized it seems better not to lump these two
island forms.

Vaurie considers it most probable that D. m. tianshanicus represents inter-specific
hybridization between major and leucopterus. With one exception our specimens
all seem nearer to major and I provisionally treat them as a form of major and not
as hybrids. The exception to this, mentioned above, is a female from the Tekkes
Plain (B.M. No. 1931.7.8.259) which was originally identified as leucopterus and
later re-identified by Vaurie as tianshanicus, because it had a wing measurement
of 137 mm. as against 120-129 mm. for 23 females of leucopterus measured by Vaurie
and because he considered the white spots on its wings too small for leucopterus. I
think this bird was correctly identified originally. The very large measurement of
its right wing appears to be due to the make-up of the skin. The left wing measures
only 130 mm. and n other females of leucopterus, picked at random, have wing
measurements ranging from 125 to 131 mm. The white spots on its wings do not
appear to be significantly smaller than those of some other specimens.

PICOIDES

I agree with Delacour that this genus is very close to Dendrocopos. Its lack of
red pigments, or rather their replacement by yellow; its flatter bill and its lack of a
fourth toe are, in combination, perhaps just sufficient grounds for keeping it in a
separate genus. It is certainly convenient to do so and the two Pico'ides species are,
unquestionably, much more closely related than either is to any other.

NOTES ON WOODPECKERS 33

However, although Pico'ides seems to have diverged a little further from the
Dendrocopos species in the course of evolution than these have from one another it
is, I think, likely that it may be phylogenetically closer to the American Dendrocopos
species than these are to Old World forms.

The colour pattern of Pico'ides tridactylus is very close to that of Dendrocopos
villosus. This is especially so with the northern Old World forms of tridactylus. The
relatively greater amount of difference between the New World forms of tridactylus
and villosus may be due to character displacement although in some other Holarctic
species or superspecies there is a tendency for the New World representatives to show a
greater amount of melanin pigment than the Old World forms (e.g. Numenius
phaeopus, Branta bernicla). In this connection it is of interest that nominate
P. t. trydactylus has a broad white stripe down its back like D. villosus whereas the
American forms, e.g. P. t. fasciatus, show white cross-barring very like that of the
" ladder-backed " Dendrocopos species and similar in position and extent to that of
D. stricklandi.

The Arctic Three-toed Woodpecker P. arcticus, which is entirely American in
distribution, has a wholely black back with, sometimes, a hint of barring indicated
by white tips to one or two feathers. In the large amount of black in its plumage
generally arcticus stands in about the same relation to American forms of tridactylus
as these latter do to most of the Old World forms of their species. Although now
largely sympatric with the smaller tridactylus, arcticus must, presumably, have
evolved in isolation from tridactylus or proto-tridactylus . The very dark, and now
apparently geographically isolated, Tibetan form of tridactylus, P. t. funebris, is
suggestive in this connection as it shows similar, and nearly as great, contrast to the
northern Old World forms of its species as does arcticus to American forms of
tridactylus.

Sapheopipo noguchii

The Okinawa Woodpecker, originally included in the genus Picus (Seebohm,
1887), was placed by Hargitt (1890) in the monotypic genus Sapheopipo, which he
having: " wing very long and pointed; tail only equal to the second primary and
falling short of the longest by about an inch ". It is found only in Okinawa, in the
central Ryukyu Islands, south of Japan.

The general coloration of noguchii is dark olive, more or less suffused with red,
especially on the tips of the feathers, on back, rump and underparts; with dark
brownish-black wings and tail. It is thus, superficially, most like the dark red
Sumatran form of Picus canus, P. c. dedemi. It has, however, very clear indications
(see Text-fig. 9) of head and breast markings nearly identical to those of many
Dendrocopos species and quite unlike those of any species of Picus or Dinopium. Also
those parts of the primaries that are not visible, or at any rate would not be conspicu-
ous, when the wings are closed have white markings similar to and I think homologous
with those of D. leucotos owstoni, the very dark race of D. leucotos from the northern
Ryukyu islands. The legs and irides of noguchii and owstoni agree in colour although
their bills do not (Kuroda, 1925).

34

D. GOODWIN

The difference of coloration between leucotos and noguchii is very similar to what
one finds between many other species and their island derivatives, for example
the pigeons Columba arquatrix and C. pollenii. They seem to have involved, assuming
noguchii to be derived from Dendrocopos stock, simply a spread of dark pigment
together with some lessening of its intensity and the spread into some of these dark
areas of red or yellow pigments; or, perhaps, their revelation through their no longer
being completely masked by melanin pigments. The pale colour of the bill might be
similarly caused. In proportion and in the shape and structure of its bill noguchii is
quite close to leucotos although its bill is proportionately longer, a common feature
with island forms. Its type of sexual dimophism is identical with that of leucotos
and several other Dendrocopos species. D. leucotos owstoni is, presumably, a more
recent invader of the Ryukyu Islands that is already beginning to evolve in the same
direction as has noguchii.

FIG. 9. Diagrammatic sketches to show resemblances and differences of colour patterns of
Dendrocopos leucotos from northern Europe (left), same species from northern Ryu-kyu
Islands (middle) and Sapheopipo noguchii (right).

I think that noguchii is an offshoot of Dendrocopos stock and is rather more likely
to have derived from Dendrocopos leucotos subsequent to the speciation of the latter
than from a more primitive common ancestor to them both. It is said (Kuroda, 1925)
to be found on or near the ground in damp woods or bamboo jungle and never in
tall trees during the day (he does not say whether it roosts in tall trees but this is,
perhaps, implied). This suggests that noguchii' s ecology may have already diverged
considerably from that of typical Dendrocopos species and this, together with its
unusual coloration, may, perhaps, justify retention of the monotypic genus Sapheopipo.

NOTES ON WOODPECKERS 35

If, however, further studies should show that it does not differ so markedly in habits
as Kuroda's notes suggest and that there are no striking anatomical differences I
would be in favour of treating Sapheopipo as a synonym of Dendrocopos.

CHRYSOCOLAPTES & DINOPIUM

Delacour (1951) suggested that the absence of the fourth toe is not of great phylo-
genetic significance in woodpeckers and that in this character Brachypternus with its
greatly reduced fourth toe, placed by Peters (1948) in the synonymy of Dinopium,
forms a link between the three-toed Dinopium species and Chrysocolaptes with four
well-developed toes. I agree with his opinion on both points; there can be no
doubt that the species comprising the above genera are all closely allied. I
provisionally maintain the genus Chrysocolaptes mainly on grounds of taxonomic
convenience. This can, perhaps, be further justified by the fact that the three species
involved lucidus , festivus and validus seem very close to each other. All, apart from
having four toes, agree in having some white on hind neck and mantle, admittedly
vestigial in lucidus, almost identical facial markings which are, however, approaching
obsolescence in validus, and bills that are longer and more massive than those of
Dinopium. Except in the case of Dinopium javanense and Chrysocolaptes lucidus,
wherever two species of either (or both) of these two genera overlap widely in range
they show fairly striking differences of colour pattern.

The very similarly-coloured Golden-backed Three-toed Woodpecker, D. javanense
and the Golden-backed Woodpecker C. lucidus are extensively sympatric and, in
general, show similar geographic variation. They are most alike in western India,
Burma and much of Malaysia where, except for the not very striking difference in the
pattern on face and throat, colour differences between them are slight or non-existant.
C. lucidus is, however, a larger bird with proportionately much larger bill. In other
areas where they overlap the two show greater differences in coloration. On Java
lucidus (race strictus) has a female with the top of the head yellow and rather obscurely
spotted, and both sexes have only a tinge of red on the rump, whereas the javanense
female has a black and white streaked crown and both sexes have a conspicuous
red rump.

On Palawan lucidus has a very distinct race (erythrocephalus) in which both sexes
have a red face and pink throat, with the usual dark striations rather reduced, the
female has the top parts of her head a mixture of red and yellow-olive with light dull
yellow spots, the individual feathers being red near the base, then olive with a yellow
terminal spot. The Palawan form of D. javanense (everetti) is very distinct, having a
boldy striped buffish and black face and the female a black forehead and crown and
red nape. It is noteworthy, however, that, compared with other forms of javanense,
this race shows parallel development to lucidus on Palawan, in having more red on
the head than is usual in the species : females of other forms of javanense have no red
on the nape and the males no red on the malar region as has everetti. They also
agree in having the dark and light facial markings less pronounced than in most other
forms of their respective species.

36 D. GOODWIN

Chrysoclaptes lucidus

I provisionally follow Peters (1948) in his wide conception of this species because,
on external characters, it is not possible to make any logical division among the
many races or presumed races of lucidus. It is, however, quite possible that some
of the island forms, especially in the Philippines, may have already reached specific
level.

Like previous workers, I cannot separate specimens from South India from those
from southern Malaya although the latter are, on average, a slightly lighter golden-
yellow on the upperparts. There seems no alternative, therefore, but to continue
to employ the name chersonesus for both these populations although it is likely that,
in a phylogenetic sense, chersonesus from southern India are more closely allied to
guttacristatus from Bengal than they are to chersonesus from the Malay Peninsula.

On the different Philippine islands the races (?) of lucidus show great variation of
head colour (besides other differences). These are comparable to the differences of
head colour between undoubted species of the Chrysocolaptes-Dinopium group
elsewhere. This is no doubt due at least partly to the lack of any closely related
sympatric species on most of these islands. Nominate lucidus from Mindanao is,
perhaps rather unfortunately, a rather nondescript or intermediate form and as it has
indistinct facial markings, mixed red and yellow upperparts and the top of the female's
head yellowish and spotted it is difficult to ally it with any one, or even any one of the
main groups of Philippine forms rather than another. The interesting sequence
of the immature plumages of guttacristatus (and other races of lucidus!} is described
in the section on juvenile plumages.

Dinopium shorii & D.javanense

The Himalayan Golden-backed Three-toed Woodpecker and D. javanense are very
closely allied and must have derived relatively recently from a common ancestor but
they appear now to overlap without interbreeding in northern Burma. There are
specimens of both from Thayetmayo and the Arrakan Hills, in the British Museum.
They appear to keep or even slightly to accentuate their minimal plumage differences
pale bases to the red head feathers in shorii, slightly different pattern of streaks on
the throat where they overlap. Admittedly hybrids between the two would be
difficult to recognize. There is a slight difference of size and bill length between them
and, no doubt, this correlates with differences of ecology.

Dinopium rafflesii

I concur with Peters in thinking it preferable to put the monotypic genus
Gauropicoides (species rafflesii) into the synonomy of Dinopium. The Olive-backed
Three-toed Woodpecker, rafflesii, seems to link Dinopium with Gecinulus, agreeing
with the former in most characters but with Gecinulus to some extent in its softer
feather texture and generally dark and rather concolorous body plumage. The
wing markings are similar in both genera.

NOTES ON WOODPECKERS

The genus GECINULUS

37

I maintain this genus for the Pale-headed Woodpecker grantia and the Green
Bamboo Woodpecker viridis. These two closely allied forms differ from the green
woodpeckers of the genus Picus in having proportionately small bills and only three
toes ; also in some plumage characters in which they are closer to Dinopium. Although
rather discrete they appear, as has been mentioned above, to be linked with
Dinopium by rafflesii and probably represent a link between Picus and the Dinopium-
Chrysocolaptes group.

DRYOCOPUS & PHLOEOCEASTES

Peters (1948) placed the American species pileams, lineatus, erythrops and galeatus
in the genus Dryocopus together with the old world forms martius and javensis.
Presumably he did this because their outer hind toes are not longer than their outer
front toes (Peters, 1948, introduction, p. 6). The characters of a narrower nasal
shelf and nostrils nearer the culmen, which he says are correlated with the shorter
outer hind toe, do not appear to be valid as there is much variation between species;
on these characters lineatus, at least, seems to fit better with those species which have
a longer outer hind toe. I do not think this relative length of the outer hind toe is,

FIG. 10. Diagrammatic sketches to show colour patterns of heads of Phloeoceastes lineatus
(top left), Dryocopus javensis (bottom left), Picus viridis (bottom middle), Phloeoceastes
haematogaster (top right) and P. melanoleucos (bottom right).

38 D. GOODWIN

by itself, a very important character. Their distribution and their plumage
characters, especially the colour patterns of their heads all suggest that the American
species placed by Peters in Dryocopus are closer to the species included by him in the
genus Phloeoceastes than they are to Old World forms. I therefore think it best to
place them in Phloeoceastes and restrict Dryocopus to the two Old World species
martins and javensis.

I think it possible that Dryocopus, as above restricted, might prove to be more
closely related to other Old World genera, such as Mulleripicus and Picus. Kilham
(1959) was impressed by the great similarity between some of the behaviour patterns
he had observed in pileatus and some described for martins (Blume, 1956) but
comparisons of these two with both (other) Phloeoceastes species and Mulleripicus
are needed. Most of sketches and descriptions in Blume's comparative study of
European woodpeckers (Blume, 1961) suggest to me that the displays, and possibly
even the calls, of martins bear a more close resemblance to those of Picus viridis than
to those described for pileatus by Kilham.

Phloeoceastes divides readily into the lineatus species-group comprising lineatus
(syn. erythrops), pileatus, schulzi and galeatus, all of which have rather softer, denser
plumage with consequently longer nuchal crests and tend to have proportionately
smaller bills, and the melanoleucos species-group, comprising melanoleucos ,
guatemalensis , pollens, haematogaster , rubricollis, robustus and leucopogon.

Within the lineatus group pileatus, lineatus and schulzi are closer to each other in
colour patterns than any of them is to galeatus. This may give a false impression,
as the very differently coloured (largely buffish brown) head and underwing of
galeatus may be the result of selection for species-specific markings in reference to
lineatus which it overlaps widely in range. P. pileatus does not overlap any congener
and schulzi would appear to have only a marginal overlap with galeatus and lineatus
at the northern edge of its range. Both pileatus and schulzi have concolorous black
underparts unlike any other close relatives. This might, however, be due to some
environmental factor as these two are in predominently temperate climates north
and south, respectively, of the ranges of other Phloeoceastes species.

The species in the melanoleucos group all appear to be very closely related to each
other. Within this group it is possible that relatively greater differences in colour
pattern between any two species may not indicate comparable phylogenetic disparity
as such differences may have evolved as isolating mechanisms in reference to each
other. Although all species show differences in colour pattern these are, on the
whole, most marked, and often correlated with differences of body proportions, in
species that overlap widely in range. It is notable that both this and the lineatus
group have evolved some forms predominantly white and others predominantly
buff or chestnut on the under wing.

I follow Peters in thinking it best to give specific rank to both guatemalensis and
melanoleucos, although they can certainly be considered as members of a superspecies.
Further studies may, however, show that they are in fact conspecific and if so the
rather striking differences in colour patterns of their heads may be a case of character
displacement as melanoleucos overlaps in range with robustus whose male has an
almost entirely red head like that of guatemalensis.

NOTES ON WOODPECKERS 39

Pholeoceastes lineatus & P. erythrops

Peters (1948) says that it has been suggested that erythrops is a subspecies of the
Lineated Woodpecker lineatus but that, if specimens have been correctly identified,
they overlap over a wide area. He treats erythrops as a good species but remarks
that " The inter-relationships of the lineatus-erythrops-schulzi group are quite involved
and require additional field study ".

Van Rossem (1934) and De Schauensee (1966) have both presented evidence that
erythrops and lineatus are conspecific. I have compared specimens of both from
Sapucay, Paraguay and they differ in no character except the presence (in lineatus)
or absence (in erythrops) of white on the scapular region. Van Rossem states that
some specimens of erythrops show white markings on the shoulder region and one of
our specimens has one white-edged feather in this region. I think the evidence
suggests that lineatus is dimorphic over the southern-most part of its range and that
erythrops is merely a black-shouldered morph of this species.

CAMPEPHILUS & IPOCRANTOR

Hargitt (1890) followed Cabanis & Heine (1863) in placing the Magellan Woodpecker
in the monotypic genus Ipocrantor; Peters (1948) placed it with the ivory-billed
woodpeckers in the genus Campephilus. Peters was, presumably, led to this because
its rather large size (in comparison with Phloeoceastes species), soft, glossy black
plumage, large white wing patches and the long crest of the female combine to give
it a very similar general appearance to an ivory-bill. Although it is possible that
magellanicus does represent a divergence from the same stock as produced the ivory-
bills, subsequent to the latter's splitting off from ancestral Phloeoceastes, I think this
rather unlikely.

The coloration and colour-patterns of the heads of both magellanicus and the ivory-
bills are closer to those of Phloeoceastes than they are to each other. In both
magellanicus and the ivory-bills the colour pattern of the head seems to represent a
modification and some degree (considerable in magellanicus) of simplification of a
pattern typical of Phloeoceastes. This simplification has involved a reduction in
the amount of red in the male ivory-bills and its increase in the male of magellanicus.
The white wing patch of magellanicus might represent a restricted version of that found
in the ivory-bills but could equally (in both) be a development from the white or
light bars that some Phloeoceastes species show on (some of) the inner webs of the
primaries and secondaries. The under wing pattern of magellanicus is closer to that
of Phloeoceastes melanoleucos and P. guatemalensis than it is to that of Campephilus.
The bill of magellanicus is dark, not white or cream-coloured. Some species of
Phloeoceastes have pale bills (others have dark bills).

It is, I think, likely that the features in which magellanicus agrees more closely with
the ivory-bills than it does with Phloeoceastes species: length of crest of female,
relatively lax, glossy plumage and rather long tail (all, probably, correlated charac-
ters) are due to convergence in related stocks but that each is, phylogenetically, at
least as closely related to Phloeoceastes as to the other. I think, therefore, that it is
preferable to retain the monotypic genus Ipocrantor for magellanicus. The only

40 D. GOODWIN

reasonable alternative, and one that seems to have much to recommend it, would be
to place all these large, related, predominantly black, white and red woodpeckers in
the single genus Campephilus. I feel, however, that it would be premature for
me to do so now in view of the work currently being done in the U.S.A. on the habits
and ecologies of the American woodpeckers which will, doubtless, soon provide other
and perhaps firmer grounds for taking or refraining from this course.

Variant individuals

The commonest plumage variation in woodpeckers, occurring in individuals of
many green-backed and golden-backed species, is for the green or golden parts of the
plumage, especially on the mantle, to be more or less suffused or mottled, with red.
This correlates with the existence of several red-backed forms closely related to or
even conspecific with green-backed and golden-backed forms. Other variants in
the British Museum (Nat. Hist.) collection are listed and described below.

Picus vittatus

A male from Pulau Langkawi, Malay Peninsula (B.M. No. 1936.4.12.586) shows
an apparent reduction of red and yellow pigments. Its breast, underparts and sides
of face are paler and more buffy than in normal specimens. The green of its upper-
parts is a colder hue, less tinged with yellow, and the red parts of its head a light
flame-orange instead of the deep scarlet-crimson of normal birds.

Dendrocopos assimilis

A male from Khipri, Sind (B.M. No. 1898. 12. 12. 331) has the red on the head
replaced by straw yellow except for the nuchal region and a line extending thence
above each eye, where it is orange-red, not the normal dark scarlet.

Dendrocopos himalayensis

A supposed male from near Simla (B.M. No . 1887 .8.10.7) has the belly, flanks and
much of the lower breast red, instead of this colour being confined to the under tail
coverts and post-cloacal area as in normal specimens. Surprisingly, if it is correctly
sexed, it shows much less red on the head than normal, this being confined to the tips
of only about a third of the crown feathers. It would seem strange if the bird
exhibited an excess of red pigment in one area yet a decrease of it elsewhere and
I am inclined to think that it may be a female, in which case (the female having
normally no red on the head) its head coloration would, like that of its underparts,
represent an increase of red.

Dendrocopos syriacus

An adult female (B.M. No. 1888. 11.1.564) shows partial albinism. It has many
white or partially white feathers on normally black areas of head, back and wings.
The distribution of red is normal.

NOTES ON WOODPECKERS 41

Dendrocopos atratus

A presumed female from the Southern Shan States of Burma (B.M. No. 1903. 12.
24.53) shows an interesting coloration as a result of partial loss of melanin pigment.
It gives a general impression of having most of the normally black areas replaced with
light grey. More closely examined it shows, on its upperparts, a narrow edging of
more intensely pigmented (darkish grey) plumage posterior to every white marking
(except where this comes to the end of the feather) and a similar but broader dark
edge immediately anterior to it. This is most clearly seen on the secondaries but
holds good for all other feathers. On the outer webs of the outer primaries the dark
grey areas are more extensive. The tail shows ill-defined light and darker grey
transverse barring but the modified, supporting, central feathers have their tips and
the edges of both webs more heavily pigmented, a dark brownish grey. The top of
the head is noticeably darker than most of the back, being, at least so far as the tips
of most feathers are concerned, a very brownish grey, almost blackish. Due,
presumably, to the reduction of melanin the underparts look much paler and more
yellowish than normal, the dark streaks are greyish brown, not brownish black, and
the red under tail coverts lighter and brighter.

It would be interesting to know whether the areas (tail tips, primary edges] etc.)
which show stronger pigmentation in this individual also have in fact more melann
in normal birds where it is not visually evident. As they all seem to be areas that
are likely to need " extra " strengthening, this would seem probable.

Dinopium benghalense

A male from southern Konkan (B.M. No. 1887. 8. 10.1957) and a male from
Khandeish (B.M. No. 1880. 1.1.225) both have many red-tipped feathers in the
malar region, giving much the impression of the type of black-enclosed red malar
stripe found in males of Picus viridis. Normally D. benghalense has no red in the
malar region in any plumage phase.

A female from Sind (B.M. No . 1941 . 5 . 30 . 1391) shows reduction of red and yellow
pigments. The normally golden parts of the mantle are a pale straw yellow, the
normally scarlet nuchal crest more or less cream, intermixed with some partially
orange-pink feathers.

A female from Lucknow (B.M. No . 1887 . 8 . 10 . 1932) although normal in coloration,
seems worth mentioning here. It has an abnormally long bill with decurved culmen
which looks more like a Hoopoe's bill than a woodpecker's. The culmen measures
64 mm., as against 36 to 38 in normal females, and overlaps the under mandible by
7 mm. The bill tips are narrow and flexible and it is impossible to believe that the
bill could have been used at all for wood-pecking. The bird had evidently been able
to obtain food as is adult and in good plumage.

Phloeoceastes lineatus

Male from eastern Ecuador in post-juvenal moult (B.M. No . 1940 . 12 . 5 .93). Has
the central part of the crown and nuchal region bright straw yellow, tinged orange,

42 D. GOODWIN

instead of red. The red on the forehead and sides of head is a little lighter in hue
than usual.

Dinopium javanense

Three females of the race intermedium show some trace of red in the head plumage.
No. 1921.12.31.92, from western Siam has several male-type red feathers, two of
them long nuchal feathers, on the left side of its head. No. 1927.6.5.419, from
Tonkin, has one red-tipped but otherwise female-type feather above its left eye and
No. 1927 .6.5. 1587, from Cochin-China, has several red-tipped but otherwise sexually
intermediate-looking feathers.

Many of the above variants involve partial loss of pigmentation and are comparable
in appearance, and probably in origin, to relatively common varieties in other groups.
It is, however, of interest that where this involves only the partial substitution of
red by yellow, as in the Dendrocopos assimilis and the Phloeoceastes lineatus, it is
comparable to one of the more conspicuous differences in coloration often found
between related species of woodpeckers. This, of course, is also true of the second
type of variant included above, where individuals show red on areas not normally
so-coloured in their sex or species.

SUMMARY

Some aspects of taxonomy and relationships of the woodpeckers are described and
discussed. Certain close resemblances in coloration and ecology are due to conver-
gence. In all geographic regions inhabited by woodpeckers the smallest species are
about the same size ; the same is true for the largest species except that there are no
large arboreal species in the Ethiopian region. The typical forms of sexual dimor-
phism and the exceptions to them are described.

Juvenile woodpeckers usually have bright pigments similar to, but not always
identical with, those of the adults. When all species are considered, the juveniles
approximate more closely to the family average in the amount of red or yellow on the
head than do the adults. In no instance, however, is the juvenile plumage considered
to be more conspicuous than the adult male's.

Leuconerpes is treated as a synonym of Melanerpes. The species obsoletus is
considered to be referable to Dendropicos not Dendrocopos. Geocolaptes is related
to other African genera, not to Colaptes. It is thought that all the New World
species of Dendrocopos are more closely related to each other than any one of them is
to any Old World species. Sapheopipo is a recent derivative of Dendrocopos. Some
of the Philippine Island races of Chrysocolaptes lucidus show colour differences
comparable to those shown by different species of the Chrysocolaptes- Dinopium
group elsewhere. The New World forms currently placed in Dryocopus seem better
placed in Phloeoceastes. Unless all the large black, white and red American wood-
peckers are to be treated as congeneric, it is best to retain the monotypic genus
Ipocrantor for the species magellanicus.

NOTES ON WOODPECKERS 43

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WETMORE, A. 1926. Observations on the birds of Argentina, Paraguay, Uruguay, and

Chile. U.S. Nat. Mus. Bull. No. 133.

,rx.

* SK <*

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IB
A REVIEW OF THE IGUANID

LIZARD GENERA URACENTRON
AND STROBILURUS

R. ETHERIDGE

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 17 No. 2

LONDON: 1968

A REVIEW OF THE IGUANID LIZARD GENER
URACENTRON AND STROBILURUS

BY

R. ETHERIDGE

Department of Zoology, San Diego State College, San Diego, California, U.S.A.

Pp. 45-64 : 2 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 17 No. 2

LONDON: 1968

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In 1965 a separate supplementary series of longer
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This paper is Vol. 17, No. 2 of the Zoological
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A REVIEW OF THE IGUANID LIZARD
GENERA URACENTRON AND STROBILURUS

By RICHARD ETHERIDGE

SYNOPSIS

The external morphology, distribution, and systematics of the iguanid lizard genera Uracentron
and Strobilums are reviewed. Four species of Uracentron and one species of Strobilurus are
recognized, and a key is given to the species of Uracentron. Two species erroneously referred to
Uracentron are discussed .

INTRODUCTION

Uracentron and Strobilurus are South American iguanid lizards belonging to a group
of genera that I have called the " tropidurines ", (Etheridge, 1964 : 629). Within
this group Uracentron is easily identified by its very large interparietal scale, and
short, spiny tail, and by the absence of a row of scales aligned mid-dorsally. Strobil-
urus is closely allied to it, and like Uracentron has a very large interparietal scale
and a spiny tail ; it differs in having a row of scales aligned middorsally. Also,
the tail of Uracentron is flat, nonautotomic, and scarcely more than half as long as
the head and body, whereas the tail of Strobilurus is nearly cylindrical, autotomic,
and about as long as the head and body.

Of the species we now know under the genus Uracentron the first was described as
Lacerta azurea by Linnaeus (1758 : 202). Prior to this the species had been figured
by Seba (1734 : 2 : 62 : 6), and Linnaeus (1754 : 42) had described in some detail
specimens then in the Museum of King Adolf Fredrik of Sweden. Lacerta azurea
was subsequently combined with diverse other lizards that also have a spiny tail
under the genera Stellio (Latreille, 1802 : 29 and 34 ; Daudin, 1802 : 36 and 40 ;
Fitzinger, 1826 : 49), and Uromastyx (Merrem, 1820 : 56-7). The generic name Ura-
centron was introduced by Kaup (1826 : 88), and in the following year he included
Uracentron in his list of the New World lizards (Kaup, 1827 : 612). The latter
publication was cited as the original description of the genus by Burt & Burt
(1933 : 48). Wagler (1830 : 145) changed the spelling to Urocentron, and with few
exceptions (Mertens, 1925 : 75 ; Dunn, 1944 : 89 ; Valdivieso & Tamsitt, 1963 : 31 ;
Peters, 1967 : 36) all later authors either adopted the change, or altered it still
again to Uranocentron (Gray, 1831 : 42, 1845 : 225; O'Shaughnessy, 1881 : 245),
or to Urocentrum (Boulenger, 1894 : 729; Werner, 1900 : 4). Cuvier (1829 : 34)
proposed Doryphorus as a substitute name, and for a time it was also in use (Schinz,
1835 : 9 2 i Guerin-Meneville, 1829-1838 : 8 ; Dumeril & Bibron, 1837 : 369 ;
Guichenot, 1855 : 26 ; Dumeril, 1856 : 559 ; Cope, 1870 : 556).

Boulenger (1885 : 182-184) recognized three species of Uracentron : azureum
(Linnaeus, 1758), flaviceps (Guichenot, 1855), and castor (Cope, 1870), and later he
described a fourth species, U. guentheri (Boulenger, 1894 : 729). A fifth species
was described by Mertens (1925 : 75), U. werneri. Burt & Burt (1933 : 48-9)
recognized five species of Uracentron in their Checklist of South American Lizards.
They included U. meyeri (Werner, 1900 : 4), and U. palluma (Tschudi, 1845 : 35),

ZOOL. iy, 2. 2

48 R. ETHERIDGE

neither of which is correctly referred to the genus Uracentron, and they failed to
include U. flaviceps and U. werneri.

In the present study U. meyeri and U. palluma are removed from the genus Uracen-
tron, U. castor is placed in the synonomy of U. flaviceps, and this species together
with U. azureum, U. guentheri, and U. werneri are considered valid species.

Strobilurus is, and has always been, considered monotypic, with the single species
5. torquatus (Wiegmann, 1834 : I 8).

ACKNOWLEDGEMENTS

During the course of this study I have had the good fortune to work in a number
of museums in Europe and in the United States. I should like to express my appre-
ciation to those curators who have allowed me to examine the collections in their
care : Mr. C. M. Bogert, the American Museum of Natural History, New York ;
Dr. E. E. Williams, the Museum of Comparative Zoology, Harvard ; Dr. R. F. Inger,
the Field Museum of Natural History, Chicago ; Dr. A. Leviton, the California
Academy of Sciences ; Dr. J. A. Dixon, the Los Angeles County Museum ; Miss
A. G. C. Grandison, British Museum (Natural History), London ; Monsieur J.
Guibe, Museum National d'Histoire Naturelle, Paris; Dr. K. Klemmer, Natur-
Museum und Forschungs-Institut Senckenberg, Frankfurt ; Dr. J. Eiselt, Natur-
historischen Museum, Vienna ; Dr. C. Edelstam, Naturhistoriska Riksmuseet,
Stockholm ; Dr. H. Wermuth, Staatliches Museum fur Naturkunde, Ludwigsburg ;
Dr. J. A. Peters, United States National Museum, Washington ; Dr. G. Peters,
Institut fur Spezielle Zoologie und Zoologisches Museum der Humbolt Universitat,
Berlin ; Dr. W. Ladiges, Zoologisches Staatsinstitut und Zoologisches Museum,
Hamburg ; Dr. F. W. Braestrup, Zoologiske Museum, K0benhavn ; Dr. W. Hellmich,
Zoologisches Sammlung des Bayerischen Staates, Munich. I should also like to
thank Mr. E. V. Malnate, Academy of Natural Sciences of Philadelphia, for providing
me with information on the holotype of U. castor.

I am especially grateful to Miss A. G. C. Grandison and her staff at the British
Museum (Natural History) where most of this work, including the preparation of
this manuscript, was undertaken.

The following abbreviations are used :

A.M.N.H. American Museum of Natural History, New York.

A.N.S.P. Academy of Natural Sciences of Philadelphia.

B.M.N.H. British Museum (Natural History), London.

C.A.S. California Academy of Sciences, San Francisco.

F.M.N.H. Field Museum of Natural History, Chicago.

L.A.C.M. Los Angeles County Museum, Los Angeles.

M.C.Z. Museum of Comparative Zoology, Harvard University,

Cambridge.

M.H.N.P. Museum National d'Histoire Naturelle, Paris.

N.M.W. Naturhistorischen Museum, Wien (Vienna).

N.R.M.S. Naturhistoriska Riksmuseet, Stockholm.

S.M.F. Natur-Museum und Forschungs-Institut Senckenberg, Frank-

furt.

IGUANID LIZARD URACENTRON AND STROBILURUS 49

S.M.N.L. Staatliches Museum fur Naturkunde, Ludwigsburg.

U.S.N.M. United States National Museum, Washington.

Z . M . B . Zoologisches Museum der Humboldt Universitat , Berlin .

Z.M.H. Zoologisches Staatsinstitut und Zoologisches Museum, Ham-
burg.

Z.M.K. Zoologiske Museum, K0benhavn (Copenhagen).

Z.S.B.S. Zoologisches Sammlung des Bayerischen Staates, Miinchen.

URAGENTRON Kaup, 1826, Isis (von Oken), 19 : 88.

Type species Lacerta azurea Linnaeus.

DIAGNOSIS. Uracentron belongs to that group of South American- West Indian
iguanids that is distinguished by the presence of a large sternal fontanelle and the
absence of femoral pores, the " tropidurines " (Etheridge, 1964 : 629). Within the
tropidurines Uracentron is probably most closely allied to Strobilurus, Tropidurus,
Plica, and Platynotus, all of which have a very large interparietal scale. Uracentron
differs from these genera in having a short, spiny, nonautotomic tail that is about
one half as long as the head and body ; in the others the tail is at least as long as the
head and body, and is autotomic. Among South American iguanids only the
unrelated Hoplocercus has a tail similar to Uracentron's, but Hoplocercus may be
easily distinguished by its small interparietal and heterogeneous dorsal squamation.

CHARACTERISTICS. Upper head scales polygonal, juxtaposed, slightly to moder-
ately convex, with a distinctly granular surface. Supraorbital semicircles poorly
differentiated ; scales in frontal and prefrontal regions similar to other snout scales.
Three or four pair of frontals, of which one to three may be in contact medially
between the orbits, or separated by a median row of small scales. One to three
rows of enlarged supraoculars, at least some of which are transversely expanded.
Enlarged supraoculars usually separated from frontals and frontoparietals by two
rows of small scales. Interparietal scale very large, one third to one fourth as wide
as the head, narrower in front, usually longer than wide, with a central " eye ".
Nasal large, dorsal, separated from rostral and upper labials by a row of scales ;
opening large, directed dorsally in the posterior part of the scale. One or two large,
convex, overlapping canthals. Canthals followed by six elongate superciliaries,
each overlapping the one behind, followed by three shorter scales that overlap in the
opposite direction. Two or three large scales in loreal region. A wide, elongate
subocular scale with a keel along its upper margin, preceded by two or three shorter
but similar scales that border the orbit anteriorly. A single row of rather large
loreolabials, ending below the subocular, which posteriorly forms a short suture
with the last upper labial. Upper labials large, more or less rectangular. Temporals
polygonal, juxtaposed, smooth or keeled. Lower labials similar to upper labials
but larger still. No enlarged postmental scales. Gulars small, mostly hexagonal,
convex, juxtaposed or subimbricate, reduced in front of the anterior transverse
gular fold, enlarged and imbricate in front of the posterior transverse gular fold.
Tympanum about as large as the eye, without projecting scales along its anterior
margin.

5 R. ETHERIDGE

No scale row aligned mid-dorsally. Dorsal and lateral scales of neck and anterior
body small, convex and juxtaposed, or keeled and imbricate. Remaining dorsal
and lateral scales of body larger, imbricate, smooth or keeled. Ventral body scales
imbricate, smooth or faintly keeled, larger than dorsals.

Tail short, not autotomic, about one half as long as the head and body, moderately
or strongly depressed, with equal whorls of large, spinose scales.

Limbs covered with rhomboidal, imbricate scales, smooth or keeled, those on the
posterior surface of the thighs reduced. Scales of palms and soles imbricate, denti-
culate, smooth or faintly keeled. Subdigital lamellae mostly tricarinate and
tridenticulate, the keels becoming faint on the distal lamellae.

Two transverse gular folds present, the anterior one just posterior to a line even
with the lower border of the tympanum, and with normal gular scales ; the posteror
fold formed by ventromedial extensions of the antibrachial folds, and enclosing
reduced but not granular scales. Antibrachial folds extend up and back over the
forelimbs, then slant back and downward along the side of the body, fading out
about midway to the hind limbs. Sides of neck with well developed, irregular folds.
A ventrolateral longitudinal fold present between the forelimb and hind limb
insertions.

Scales on top and sides of head and on chin with numerous, closely set scale
organs. Body scales with a single scale organ or none at all. Caudal scales with
several prominent scale organs along posterior edge on each side of keel. No
femoral or preanal pores.

Uracentron azureum (Linnaeus)

Lacerta azurea Linnaeus, 1758, p. 202 (type locality, Africa; here restricted to the vicinity of

Paramaribo, Surinam).

Stellio brevicaudata Latreille, 1802, p. 29 (type locality, interior of Guiana and Surinam).
Stellio azureus Latreille, 1802, p. 34.
Stellio brevicaudatus Daudin, 1802, p. 40.
Stellio azureus Daudin, 1802, p. 36.

Uromastyx caeruleus Merrem, 1820, p. 56 (substitute name for Stellio azureus}.
Uromastyx azureus Merrem, 1820, p. 57.
Uracentron azureum Kaup, 1826, c. 88.
Uracentron coeruleus Kaup, 1826, c. 88.
Doryphorus brevicaudatus Cuvier, 1829, p. 34.
Doryphorus azureus Cuvier, 1829, p. 34.
Urocentron azurea Wagler, 1830, p. 45.
Ophessa (Uranocentron) brevicaudatus Gray, 1831, p. 42.
Ophessa (Uranocentron) azureus Gray, 1831, p. 42.
Urocentron brevicaudatum^Wiegma.n-n, 1834, p. 48.
Urocentron azureum Fitzinger, 1843, p. 77.
Urocentron azureum Boulenger, 1885, p. 182.

SYNTYPES. No. KaF 1900 : 113 (2 exs) in the Naturhistoriska Riksmuseet,
Stockholm, Sweden.

TYPE LOCALITY. In listing Africa as the type locality of Lacerta azurea, Linnaeus
(1758 : 202) probably was misled by Seba (1734 : 2 : 62 : 6), who had figured this

IGUANID LIZARD URACENTRON AND STROBILURUS 51

species and given Africa as its origin. Nevertheless, Linnaeus did have before
him two specimens upon which his description was based, and there is reasonably
good evidence that they had come from the neighbourhood of Paramaribo,
Surinam.

Many of the species of South American plants and animals described by Linnaeus
were collected by Carl Gustaf Dahlberg, a man of Swedish nationality, who arrived
in Surinam in 1746, where he acquired plantations on Peruca Creek, and Cottica
River, within 100 km of Paramaribo. In 1754, during one of his return trips to
Sweden, Dahlberg presented his collections to King Adolf Fredrik and in the same
year Linnaeus (1754 : 42) described specimens of Lacerta azurea in the museum of
the King. When Dahlberg returned to Surinam in 1755 he took with him Daniel
Rolander at the request of Linnaeus, who at least partly financed Rolander's trip.
Rolander was a former student of Linnaeus, and private tutor to his son. Rolander
is known to have collected in the neighbourhood of Paramaribo, and also went up
the Commewijne River. The unrest caused by the revolt of escaped negro slaves
prevented him from penetrating deeper into the interior of Surinam, and in 1756
he returned to Sweden. Part of Rolander's collection was purchased by the Swedish
Baron de Greer, who in turn presented the specimens to Linnaeus (Holthuis, 1959 :
17-21).

Most of the South American species of lizards described by Linnaeus, including
Uracentron azureum, are widely distributed in the northern part of the continent,
and are known to occur at Paramaribo. It seems probable that many, if not all,
of Linnaeus' specimens were from the collections of Dahlberg and Rolander, made in
the vicinity of Paramaribo. I therefore propose the restriction of the type locality
of Uracentron azureum to the vicinity of Paramaribo, Surinam.

CHARACTERISTICS. The supraoculars are variable in shape ; one, two, or two and
a half rows of enlarged scales may be present. The scales of the inner row are always
to some extent transversely widened, but never strap-like. The enlarged supra-
oculars are always separated from the superciliaries by two rows of small scales.

The dorsal and lateral scales of the neck and anterior part of the body are very
small, smooth, convex, and juxtaposed. Posteriorly the dorsal and lateral scales
of the body become larger, imbricate, and obtusely keeled, largest in the sacrolumbar
region. The ventral scales are smooth. Scale counts are given in Table I.

The tail is moderately depressed, with about 21 whorls of large, spinose scales.
A whorl halfway between the vent and the tip of the tail contains about 13 scales.
The tail is 0.48 to 0.65 (mean, 0.58) times as long as the head and body.

The snout-vent length of the largest male examined is 87 mm, of the largest
female 86 mm.

Colour in preservative : the upper surfaces are bluish-grey, with bold, black mark-
ings. The medial half of the supraocular region is black, and there is usually a
median, V-shaped mark in front of the frontal region. A W-shaped mark on the
back of the head has its median apex on the interparietal scale, and its horns extend
down to the suboculars. Two crescentric bands cross the neck, the extremities of
the anterior one curving down through the tympani to the angle of the mouth.

R. ETHERIDGE

IGUANID LIZARD URACENTRON AND STROBILURUS 53

A series of similar crescentric bands cross the back, the extremities of the first curving
down into the antihumeral folds. Five or six bands may be present between the
shoulders and the sacrum, or the posterior one, two, or three bands may be broken
and off-set at the dorsal midline, or may be broken up into a bold reticulum. A
black reticulum covers the upper surfaces of the limbs, and is most intense on the
hind limbs. The tail has irregular black marks above, and about one half of the
spines along the sides of the tail are tipped with yellow. Below, the throat is dark
bluish-grey. The belly is a lighter grey, and the ventral surface of the tail is dark
bluish-grey along the sides, with a yellowish area down the middle.

Mr. Marinus S. Hoogmoed has furnished me with colour notes on a living individual
captured along Feticreek, Litani River, in Surinam. The colour is described as
" head and back yellow-green mottled, tail with yellow points, throat yellow-green ;
belly, underside of legs grey-blue ". Cott (1926) has published an excellent coloured
illustration of this species.

RANGE. (Text-fig, i). Records of Uracentron azureum are from British Guiana,
French Guiana, and Surinam, and from along, or near the Amazon River in north-
eastern Brazil, as far west as Manaus. U. azureum and U. flaviceps occur together
at Manaus.

MATERIAL EXAMINED. British Guiana : no specific locality B.M.N.H.
1905.10.21.1. Surinam : Bergenaal N.M.W. 13926 (2 exs) ; no specific locality.
M.H.N.P. 2515, Z.S.B.S. 534/0, M.C.Z. 4500. French Guiana : Cayenne M.H.N.P.
196, 2514, 2396 ; Saint Laurent M.H.N.P. 24.121 ; Mornes du bas Mahurg M.H.N.P.
03.20 ; no specific locality M.H.N.P. 8211, 2516, B.M.N.H. 1920.1.20.1334,
Z.S.B.S. 534/0. Brazil : Amazonas, Manuas B.M.N.H. 1922.6.15.1, C.A.S. 84247 ;
Amazonas, Manjuru River A.M.N.H. 101945 ; Para, Ilha Marajo, Soute S.M.F.

24930 ; Para, Ilha Marajo, Dist. Calderao Z.S.B.S. 37/1924 (5 exs), Z.M.H. 4421
(5 exs), 4496 ; Para, Ilha Marajo B.M.N.H. 1923.11.9.68-72, 1926.5.5.5, S.M.F.

24931 ; Para, within 50 miles of Belem M.C.Z. 53216 ; Para, Amnuathena, Rio
Tocantins Z.S.B.S. 674/1920 ; Amapa, Serra do Navio S.M.F. 59580 ; Maranhao
S.M.F. 11202 ; no specific locality M.H.N.P. 92.290, 99.215, N.M.W. 13928.1,
B.M.N.H. 51.7.17.45, A.M.N.H. 58277-9, 60330-2. South America : no specific
locality B.M.N.H. 47.2.19.6, 59.12.28.2, 66.8.14.302, N.M.W. 13927 (2 exs),
N.R.M.S. Kaf 1900 : 113 (syntypes), S.M.N.L. unnumbered.

Additional records from Surinam have been provided to me (in litt. )by Mr. Marinus
S. Hoogmoed : Litani Fetikreek, Sipaliwini, Paloemeu, Berg en Dal, and Moengo.

Uracentron guentheri Boulenger

Urocentrum gttentheri Boulenger, 1894, P- 7 2 9 (type locality : Iquitos, Peru).
Urocentr on guentheri Burt and Burt, 1933, p. 49.

HOLOTYPE. No. 93 . 7 . 10 . 13 (RR 1946 . 8 . 29 . 85) in the British Museum (Natural
History), London. Collector, A. E. Pratt.

CHARACTERISTICS. Four or five transversely widened, band-like supraocular
scales are present in a single row, all in contact laterally with the superciliaries, or

54 R. ETHERIDGE

the first two or three may be separated from the superciliaries by a row of very small
scales.

The dorsal and lateral scales of the neck and anterior part of the body are very
small, smooth, convex, and juxtaposed. Posteriorly the dorsal and lateral scales
of the body become larger, imbricate, and weakly, obtusely keeled, largest in the
sacrolumbar region. The ventral scales are smooth. Scale counts are given in
Table I.

The tail is moderately depressed, with about 21 whorls of large, spinose scales.
A whorl halfway between the vent and the tip of the tail contains about 13 scales.
The tail is 0.52 to 0.57 (mean 0.54) times as long as the head and body.

The snout-vent length of the largest male examined is 75 mm., of the largest
female 75 mm.

Colour in preservative : the upper surfaces are bluish-grey with black markings.
Many of the head scale sutures are edged in black, and the supraocular region on
each side is outlined in black, with a black bar across the middle. On the frontals
is a median, longitudinal black bar. Five or six narrow, somewhat irregular,
crescentric bands cross the posterior part of the head and neck, with the horns of
each crescent pointing downward and forward. Across the shoulders is a similar,
narrow crescent that descends on each side into the antihumeral folds. The upper
surface of the limbs and most of the back are covered by a bold reticulum of black
lines. In the middle of the anterior part of the back the reticulum tends to form short,
transverse bars. On the hind leg the reticulum extends onto the posterior surface of
the thigh. The caudal scales and the ends of their spines are outlined in yellow.
The ventral surface of the head and body is light bluish-grey, becoming yellowish
in the preanal area and under the tail. There are no records of the colour in life
of this species, but is seems probable that the bluish-grey background of preserved
animals is green in life. A black-and-white illustration of the pattern may be found
in Boulenger, 1894 : pi. 47, fig. 3.

RANGE. (Text-fig, i). Records of Uracentron guentheri are from the Rio Ucayali
system in the Department of Loreto in eastern Peru, and from the western Amazonian
basin in northwestern Brazil. The range of U. guentheri apparently overlaps that
of U. flaviceps in eastern Peru and Brazil, and the two are definitely known to occur
together at Iquitos, Peru.

REMARKS. Boulenger (1894 : pi. 47, fig. 3) shows the tip of the tail of the holo-
type of Uracentron guentheri blunt. Mertens (1925 : 76) used this as a character to
distinguish it from U. werneri. However, the blunt tail of the holotype of
U. guentheri is due to injury; the tip is missing and the end of the tail is healed over
with scar tissue. The tail has i6| rows of spinose scales ; about 5^ rows are missing.

MATERIAL EXAMINED. Peru : Dept. Loreto, lower Rio Cushdbatay A.M.N.H.
56410 ; Dept. Loreto, Roaboya, Rio Ucayali A.M.N.H. 57204 ; Dept. Loreto, Rian
Rian, Contamana region, Rio Sahnaya Valley A.M.N.H. 57205 ; Dept. Loreto,
Iquitos B.M.N.H. 93.7.10.3 (RR 1946.8.29.85) (Holotype). Brazil : Amazonas,
Rio Jutai Z.M.B. 30974 ; Amazonas, Rio Solimoes, Lago Calado S.M.F. 30304/5.2.

IGUANID LIZARD URACENTRON AND STROBILURUS 55

Uracentron werneri Mertens
Uracentron werneri Mertens, 1925, p. 75 (type locality : upper Orinoco, Venezuela).

HOLOTYPE. No. 11203 in the Natur-Museums und Forschungs-Institutes, Frank-
furt am Main. Collector, G. Hubner.

CHARACTERISTICS. The shape and size of the supraoculars are variable ; two or
two and a half rows of enlarged scales with four or five scales in each, those of the
inner row somewhat transversely widened, and those of the outer row separated
from the superciliaries by two rows of small scales.

The dorsal and lateral scales of the neck and anterior part of the body are very
small, smooth, convex, and juxtaposed. Posteriorly the dorsal and lateral scales
of the body become larger, imbricate, and remain smooth. The ventral scales are
smooth. Scale counts are given in Table I.

The tail is moderately depressed, with about 21 whorls of large, spinose scales.
A whorl halfway between the vent and the tip of the tail contains about 13 scales.
The tail is 0-56 to 0-60 (mean, 0-57) times as long as the head and body.

The largest male examined is 60 mm. snout-vent length, the largest female is
58 mm.

Colour in preservative : all upper surfaces are dark greyish-brown or bluish-black,
somewhat lighter on the snout and sides of the head. A pattern of light spots and
a dark nuchal collar is very faintly indicated in some specimens, including the holo-
type. The holotype has a bold, yellowish, asymetrical spot on the top of the head
that covers the interparietal and adjacent scales on both sides and to the rear, and
extends forward on the right side only, narrowing towards the superciliary ridge.
This mark appears to be anomalous, for it is absent in all specimens other than the
holotype. According to Valdivieso and Tamsitt (1963:31) living animals of this
species are bright green.

RANGE. (Text-fig, i). Records of Uracentron werneri are from the upper Rio
Orinoco valley in Venezuela, and from near the western tributaries of the Orinoco
in Colombia.

MATERIAL EXAMINED. Venezuela : Amazonas, Alto Orinoco S.M.F. 11203
(holotype). Colombia : Meta, near Macarena, Rio Guayabero A.M.N.H. 91755 ;
Vaupes, Cerro Yapopoda M.C.Z. 67982. Published records from Colombia are
Caqueta (Dunn, 1944 : 89), and near the Macarena Mountains, Meta (Valdivieso
and Tamsitt, 1963 : 31).

Uracentron flaviceps (Guichenot)

Doryphorus Azureus, variety Dumeril, 1851, p. 85.

Doryphorus flaviceps Guichenot, 1855, p. 26 (type locality : Sarayacu, Peru).

Doryphorus castor Cope, 1870, p. 556 (type locality : Pebas, " Ecuador ", now Peru).

Uranocentron flaviceps O'Shaughnessy, 1881, p. 245.

Urocentr on flaviceps Boulenger, 1885, p. 183.

Urocentron castor Boulenger, 1885, p. 184.

HOLOTYPE. No. 6682 in the Museum National d'Histoire Naturelle, Paris.
Collectors, Castelnau and DeVille.

CHARACTERISTICS. The supraocular scales are variable ; usually there is an inner

56 R. ETHERIDGE

row of four or five transversely widened scales, and an outer row of three or four
scales that are about half as large as those of the inner row, with a single row of
scales between the outer row and the superciliaries.

The dorsal nuchal scales are somewhat conical and keeled, with tiny granules in
between. These grade laterally into smaller, but otherwise similar scales on the
sides of the neck. The dorsal body scales are larger, imbricate, and distinctly
keeled, largest in the sacrolumbar region. The lateral body scales are a little smaller,
but otherwise similar to the dorsals. The ventrals are smooth, or faintly keeled,
especially in the pectoral region. Scale counts are given in Table I.

The tail is very flat and wide, almost leaf-like, with 30 to 38 whorls of moderately
large, spinose scales. A whorl halfway between the vent and the tip of the tail
contains 9 to n scales. The tail is 0-48 to 0-63 (mean 0-50) times as long as the head
and body.

The largest male examined is 128 mm. snout-vent length, the largest female is
87 mm.

Colour in preservative : juveniles and females are a rich, dark brown above, with
numerous small, light spots; the spots are light bluish on the head, forelimbs, and
anterior part of the body, becoming yellowish on the posterior part of the body,
hind limbs, and tail. In juvenile males the light spots tend to be absent from a
crescentric band across the nape, leaving a solid, dark band. The throat in females
and juvenile males is light bluish with brown spots that may be arranged in irregular,
oblique rows ; other ventral surfaces are light grey. In adult males the light bluish
spots of the head and neck increase in extent, and join one another until the head and
neck have small brown spots on a bluish background. The dark brown nuchal
band becomes conspicuous, and is bordered behind by a light, bluish band. On the
remainder of the body, limbs, and tail the light spots become obscure or disappear
entirely, leaving a uniform dark brown surface. The ventral surfaces become
brown as well, except for the chin and throat, and the ventral surface of the tail
where some yellow spotting usually remains.

According to my field notes of yth August, 1961, an adult male of 88 mm. snout-
vent length, obtained 39 km. NNE of Oxapampa, Dept. Pasco, Peru (now L.A.C.M.
No. 39161) had the following colour in life : dorsum of body, limbs, and tail very
dark brown, finely punctate with yellow on limbs and sides of body ; head reddish-
brown mottled ; collar a wide, dark brown band bordered behind by a yellow band ;
chest and venter of limbs iridescent blue-green ; tail reddish below ; throat dirty
yellow. Another male, 114 mm. snout- vent length, from the same locality, had
the same pattern, but with colours more intense, and the head was bright reddish.
After six years in preservative the reddish and yellow colours have become light
bluish-green. Guichenot (1855, pi. 3, fig. 2) gives a coloured illustration of the
holotype.

RANGE. (Text-fig. 2). Records of Uracentron flaviceps are from the western
part of the Amazonian Basin in northwestern Brazil, southeastern Colombia,
eastern Ecuador, and eastern Peru. The species occurs together with U. guentheri
in Peru, and with U. azureum in Brazil.

57

WEST LONGITUOC

FIG. 2. Map of northern South America showing localities for Uracentron flaviceps.

REMARKS. In his description of Doryphorus castor, Cope (1870 : 55) used as the
principle diagnostic character the presence of the nostril between two scales rather
than within a single scale. Boulenger (1885 : 184) recognized U. castor as a valid
species, but called attention to its close similarity with U. flaviceps. Mr. E. V. Mal-
nate has examined the holotype of U. castor (A.N.S.P. 11303), and compared it with
specimens of U. flaviceps from eastern Peru. He has informed me (in litt.} that
contrary to Cope's description each nasal opening is within a single scale, and in this
and in all other details of scalation and colour pattern the specimen is indistinguish-
able from U .flaviceps.

In his list of the type specimens of lizards in the Paris Museum, Guibe (1954 : 42)
incorrectly considered Uracentron flaviceps to be a synonym of U. azureum.

58 R. ETHERIDGE

MATERIAL EXAMINED. Colombia : Amazonas, Leticia F.M.N.H. 83038, 78384-5
(2 exs) ; Putumayo, Santa Rosa de los Kofanes, upper Rio Guamues F.M.N.H.
165828. Ecuador : Sarayacu M.H.N.P. 6882 (holotype), B.M.N.H. 80.12.8.56 ;
Canelos B.M.N.H. 80.12.8.57-9 ; Pastaza River M.C.Z. 37270 ; Napo, Santa
Cecilia M.C.Z. 92519 ; no specific locality Z.M.B. 9988. Brazil : Amazonas, Rio
Madiera A.M.N.H. 57207 ; Amazonas, Manaus N.H.M.P. 92.291 ; Amazonas, Rio
Jutai Z.M.B. 30973 ; Amazonas, Rio Solimoes N.R.M.S. 3342 ; Amazonas, Lago
Calado S.M.F. 30304/5.1. Peru : Loreto, Iquitos A.M.N.H. 56408, N.M.W. 13929
(2 exs), 13930, F.M.N.H. 45487 ; Loreto, Prov. Ucayali, Yarinacocha F.M.N.H.
45484, 56064 ; Loreto, Prov. Ucayali, Pucalpa F.M.N.H. 56061-3 ; Loreto, Prov.
Bajo Amazonas, Quidtococha F.M.N.H. 45485-6 ; Loreto, upper Ucayali A.M.N.H.
71101-2 ; Loreto, Pebas A.N.S.P. 11303 (holotype Doryphorus castor) ; Pasco,
39 km NNE of Oxapampa L.A.C.M. 39161 ; Loreto, Pongo Manseriche, Marafion
valley A.M.N.H. 56409 ; between Apaga and Nieva A.M.N.H. 57206. No specific
locality B.M.N.H. 69 . 5 . 21 . 57.

Species Referred Erroneously to Uracentron

Urocentrum meyeri was described by Werner (1900 : 4) from Lima, Peru, and
included by Burt & Burt (1933 : 49) in their list of South American lizards. The
holotype and only specimen was destroyed in the Dresden Museum (No. 1764)
during the last days of World War II. However, the description contains details
that are sufficient to identify the specimen as belonging to the genus Stenocercus
rather than Uracentron.

The upper head scales are smooth, the interparietal scale small, and there is a
projecting auricular scale. The tail is flattened, but scarcely wider than the sacral
region, with spiny scales in rings, and is a little longer than the head and body
length : total length 131 mm., tail length 77 mm. These are all characteristics of
the spiny-tailed species of Stenocercus: atrigularis, roseiventris, marmoratus , crassi-
caudatus, carrioni, and simonsi. Additional details in the description, and the
type locality indicate that the specimen may have been one of Stenocercus crassi-
caudatus. The sides of the neck have two converging folds, a postympanic and an
antihumeral fold, with a horizontal fold between them ; the dorsal scales are small
and granular. A median dorsal crest is not mentioned, and presumably was lacking.
The ventral scales are said to be similar to the dorsals, but faintly keeled, almost
smooth. Only the faintly keeled ventrals are not characteristic of 5. crassicaudatus.
Although there are no records of S. crassicaudatus from Lima, the species does occur
inland from Lima at higher elevations. The exact identity of Werner's species may
never be known, but in any event it is clearly not referrable to Uracentron.

Burt & Burt (1933 : 49) also included in their checklist of South American lizards
the name Urocentronpalluma, which they credited to Tschudi (1846 : 35). However,
it is clear from Tschudi's own treatment in his Fauna Peruana (incorrectly cited by
Burt & Burt as 1845), and in an earlier work (Tschudi, 1845 : 157-8) that he is
referring to Phymaturus palluma, which Tschudi correctly attributes to Molina
(1782 : 217). Tschudi listed Phymaturus Gravenhorst as a subgenus of Urocentron,

IGUANID LIZARD URACENTRON AND STROBILURUS 59

and gave as its only species in Peru " U. palluma Tschudi ". Although he followed
the species with his own name, he listed Lacerta palluma Molina in its synonomy.
Burt & Burt's error is clearly that of accepting Tschudi's placement of Phymaturus
palluma in the genus Urocentron, and, apparently because Tschudi followed the
species name with his own, considering Tschudi to be its author. Phymaturus is
a valid, monotypic genus, containing only the species palluma, and quite unrelated
to Uracentron. It differs from the latter in many ways, including the presence of
tricuspid premaxillary teeth, preanal pores in males, a small interparietal scale,
and a rather slender, somewhat spiny, autotomic tail.

Relationships Within Uracentron

Uracentron flaviceps stands apart from the other three forms of Uracentron as the
most distinctive species. Compared with the others its tail is much flatter and
wider, with smaller, less distinctly spinose scales. Its body scales are larger and
consequently fewer in number around the middle of the body, and more distinctly
keeled. U. flaviceps also attains a greater maximum size.

Uracentron azureum, U. guentheri, and U. werneri are very similar in the form of
their body and tail, and in scalation. U. guentheri differs from azureum and werneri
in having greatly expanded, band-like supraoculars, and U. werneri differs from
azureum and guentheri in having smooth dorsal scales in the sacrolumbar region.
The most obvious differences among the three are their colour patterns : bold black
cross-bands usually followed by a bold reticulum in azureum, a finer reticulum with
narrow cross-bands on the neck in guentheri, and solid or faintly spotted in werneri.
The most extreme example of breaking up of the posterior cross-bands into a reti-
culum in azureum does not approach the pattern of guentheri, but transitional
stages between the two are not difficult to imagine. The pattern of werneri might
result from an overall darkening of the pattern of guentheri.

The ranges of U. azureum, U. guentheri, and U. werneri are not known to contact
one another, or to overlap. With the acquisition of more specimens from inter-
mediate areas it may become possible to determine whether or not any two, or all
three of these species intergrade. Until that time the best course would seem to be
recognition of three separate species.

1 Tail very strongly depressed. More than 25 whorls of caudal scales from base to tip

of tail. Dorsal body scales, including nuchals, distinctly keeled . . flaviceps

- Tail moderately depressed. Not more than 25 whorls of caudal scales from base to

tip of tail. Dorsal scales of neck and anterior part of body smooth ... 2

2 Four or five band-like supraoculars separated from superciliaries by one row of small

scales, or in contact with superciliaries ....... guentheri

- Supraoculars variable, but those enlarged separated from superciliaries by at least

two rows of small scales .......... 3

3 Dorsal body scales in sacrolumbar region obtusely keeled. A bold dorsal pattern of

black cross-banks or anterior cross-bands and posterior reticulum . . azureum

- Dorsal body scales of sacrolumbar region smooth. Dorsum solid gray-brown, or

faintly spotted werneri

60 R. ETHERIDGE

STROBILURUS Wiegmann, 1834, Herp. Hex., p. 18

Type species Strobilurus torquatus Wiegmann.

DIAGNOSIS. Strobilurus is a member of the tropidurine group of South American-
West Indian iguanid lizards. It was not initially included in that group because
I had no data on the structure of the skeleton (Etheridge, 1964 : 629). I have
since determined, however, that a large sternal fontanelle is present, and on the
basis of this and other osteological and integumentary characteristics, Strobilurus
is clearly a member of the tropidurine group. Within the tropidurines Strobilurus
is probably most closely allied to Uracentron, Tropidurus, Plica, and Platynotus,
all of which have a very large interparietal sacle. The tail of Strobilurus is auto-
tomic, about as long as the head and body, and provided with unequal whorls of
spinose scales. In Uracentron the tail is about half as long as the head and body,
and is not autotomic. In the other genera the tail is considerably longer than the
head and body, and autotomic. Some species of Tropidurus have a moderately
spiny tail due to sharp, projecting mucrons, but the spines are not nearly as well
developed as they are in Strobilurus. Among other South American iguanids some
species of Stenocercus also possess a tail about equal to the head and body length,
and provided with unequal whorls of stout, spinose scales. However, the inter-
parietal scale of Stenocercus is very small, or absent entirely.

CHARACTERISTICS. Upper head scales polygonal, juxtaposed, with a distinctly
granular surface. Supraorbital semicircles not well differentiated as such. Three
pairs of frontals in contact between the orbits, preceded by two or three larger
prefrontals and frontonasals on each side. Two rows of enlarged supraoculars,
four or five scales in each row. Those on the inner row a little wider than long and
separated from the supraorbital semicircles by two rows of smaller scales ; those on
the outer row separated from the superciliaries by two rows anteriorly and one row
posteriorly. Interparietal scale very large, about one third as wide as the head,
narrowing anteriorly, and as wide as to a little wider than long. A median " eye "
present anteriorly in the interparietal scale, and often a median pit followed by a
median groove posteriorly. Nasal scale large, dorsolateral in position, with a small,
dorsally directed nostril in the posterior part of the scale. Nasal separated from
rostral and upper labials by a single row of scales. Two rather small but strongly
convex canthals. Canthals followed by five or six elongate, curved superciliaries,
each of which overlaps the one behind, these in turn followed by three shorter super-
ciliaries that overlap in the opposite direction. Several polygonal scales in the
postnasal-loreal-preocular region. An elongate subocular narrowing anteriorly,
with a keel along its upper margin. Loreolabials in a single row below the nasals,
two rows below the loreal region, and one row between the subocular and upper
labials. Temporals imbricate, obtusely keeled. Tympanum large, about equal to
eye opening, with two flat, projecting scales in front. Upper labials more or less
rectangular. Lower labials about one and a half times wider than upper labials.
No enlarged postmentals. Gulars smooth, imbricate, reduced in size medially and
posteriorly.

A row of scales aligned mid-dorsally from occiput to base of tail, its scales slightly

IGUANID LIZARD URACENTRON AND STROBILURUS 61

larger than, but otherwise similar to adjacent scales, forming a low denticulation in
individuals over 100 mm. snout-vent length. Dorsal nuchals rhomboidal, imbricate,
sharply keeled, the keels rising to sharp mucrons. Lateral nuchals smaller, spinose
along the lateral nuchal folds. Dorsal scales of body rhomboidal, keeled, imbricate,
shortly mucronate, the keels forming oblique lines that converge posteriorly toward
the dorsal midline. Lateral body scales slightly smaller, but otherwise similar to
the dorsals. Ventral scales smooth, imbricate, about one and a half times as large
as the dorsals.

Tail about as long as the head and body, and autotomic. Base of tail slightly
depressed and a little wider than the sacral region, with eight equal whorls of
large, sharply keeled, and strongly spinose scales. Remainder of tail cylindrical and
much narrower than the base, with scales of unequal whorls that correspond to the
autotomy segments of the vertebral column. Anteriorly each segment with a
ventrally incomplete anterior ring of small, smooth scales, and posteriorly an
additional ventrally incomplete ring of small scales added to the anterior border of
each segment.

Dorsal scales of forelimb rhomboidal, keeled, imbricate, about as large as dorsal
body scales. Ventral scales of forelimb smaller, smooth, imbricate. Dorsal scales
of hind limb sharply keeled and spinose, similar to but much smaller than spinose
scales of tail. Postfemoral scales reduced, keeled, imbricate. Scales of palms and
soles sharply keeled and mucronate, some of them tricarinate and tridenticulate.
Subdigital lamellae tricarinate and tridenticulate.

A transverse fold extends across the throat just posterior to a line even with the
tympani, but scales within fold not differentiated from adjacent gulars. A pair
of short, deep, antihumeral folds enclosing very small scales, widely separated vent-
rally by about 12 anterior pectoral scales. A dorsolateral fold extends posteriorly
from upper border of tympanum about halfway back to shoulder ; a ventrolateral
fold begins as a pair of converging folds that meet just posterior to tympanum, and
extend back to intercept antihumeral fold.

Scales on top and sides of head with numerous scale organs. Dorsal scales of
body and limbs with one to three scale organs on the free edge, or none at all. Caudal
scales with three to eight scale organs on each side of the keel, along the free edge.
No femoral pores or preanal pores.

Strobilurus torquatus Wiegmann

Strobilurus torquatus Wiegmann, 1834, p. 18 (type locality, Brazil).
Steironotus (Strobilurus) torquatus Fitzinger, 1843, p. 71.
Doryphorus spinosus Guichenot, 1855, p. 27 (type locality, Bahia, Brazil).
Strobilurus torquatus Boulenger, 1885, p. 181.

SYNTYPES. Nos. 672-3 and 9215 in the Zoologisches Museum der Humboldt
Universitat, Berlin.

CHARACTERISTICS. Since Strobilurus is a monotypic genus the characteristics of
torquatus are the same as those of the genus. The vertebral scales counted from the

62

R. ETHERIDGE

occiput to a line even with the anterior surface of the thighs when the limbs are
extended at right angles to the body number 39 to 57 (mean, 55-6) ; the paravertebral
scales counted in the same way are 49-72 (mean, 67-6). There are 19 to 29 (mean,
24-1) lamellae under the fourth finger, and 23 to 34 (mean 28-3) under the fourth
toe. The tail is 0-93 to 0-95 (mean, 0-94) times as long as the snout-vent length.
The largest male examined is 97 mm. snout-vent length, the largest female is
106 mm.

Colour in preservative : the head and neck above are bluish-grey, with bold, black
markings. There is a narrow, black crescent on each side where the supraoculars
contact the frontals and frontoparietals. A wide crescent extends across the anterior
part of the neck from ear to ear, and from this crescent there extends anteriorly a
medial bar. On each side a black crescent extends from the interparietal scale
down to the angle of the mouth. A wide, black, crescentric collar about six or seven
scales wide middorsally, extends down on each side into the antihumeral fold.
Behind the black collar the dorsal surface of the body, limbs, and tail is somewhat
darker greyish-green, with obscure lighter cross-bands. The chin and throat are
marbled light and dark grey, and the belly is light grey. A coloured illustration may
be found in Guichenot (1855, pi. 7, fig. la), labelled Doryphoms spinosus.

RANGE. Strobilurus torquatus is known from the states of Pernambuco, Alagoas,
and Bahia, Brazil.

REMARKS. Wiegmann described Strobilurus torquatus in a footnote (1834 : ~&]
and the subsequent taxonomic history of the genus and species has been relatively
uncomplicated. The species has remained rare in collections.

MATERIAL EXAMINED. Brazil : Alagoas, Sao Miguel M.C.Z. 59275 ; Bahia, no specific
locality Z.M.B. 9215, 672-3 (syntypes), 8272, M.H.N.P. 5085 (2 exs, syntypes of
Doryphorus spinosus}, N.M.W. 13908 (2 exs),B.M.N.H. 62.11.23.50, 1903.10.16.23;
Pernambuco, no specific locality B.M.N.H. 88.4.18.6. South America : no
specific locality M.H.N.P. 6880, N.M.W. 13909 (2 exs), B.M.N.H. xxiii.io4a.

Scales along middle

of black
Scales around middle

of body
Subdigital lamellae

of fourth finger
Subdigital lamellae

of fourth toe

azureum

92-(lO7'O)-I2O

TABLE I

guentheri
N = 6

werneri

N = 3

!27-(i23-3)-i 4 i

98-(iio-4)-i28

2 4 -(27- 4 )-30

3 i-(3i-6)-34

flaviceps
N = 29

74-(8o-o)-88
66-(78-2)-88

25-(29'7)-33
26-(30-3)-34

TABLE i . Some scale counts of Uracentron. Scales along middle of back counted from occi-
put to line even with anterior margin of hind limb at right angles to body ; scales around
middle of body counted halfway between limb insertions. Mean figures in parentheses.
N = number of specimens examined.

IGUANID LIZARD URACENTRON AND STROBILURUS 63

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NIGERIAN LIZARDS OF THE GENUS
AGAMA (SAURIA : AGAMIDAE)

HIS?

ALICE G. C. GRANDISON

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 17 No. 3

LONDON: 1968

18

NIGERIAN LIZARDS OF THE GENUS AGAMA
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BY

ALICE G. C. GRANDISON

*" " " "*"/ t%',A

British Museum (Natural History)

Pp. 65-90; 6 Plates: 2 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 17 No. 3

LONDON : 1968

THE BULLETIN OF THE BRITISH MUSEUM

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

Parts will appear at irregular intervals as they become
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within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 17, No. 3 of the Zoological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

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

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NIGERIAN LIZARDS OF THE GENUS AGAMA
(SAURIA : AGAMIDAE)

By ALICE G. C. GRANDISON

SYNOPSIS

The external morphology, distribution and systematics of the species of Agama occurring in
Nigeria are reviewed. The number and position of the dermal sense organs on the trunk scales
are used to help clarify the status and relationships of the five species recognized. Agama
agama benueensis Monard is shown to be a complex comprising two distinct species, one of
which is described as new, the other sympatric with but unrelated to Agama agama (Linnaeus).
Agama boensis Monard is not a valid form; the types are a mixed series of Agama sankaranica
Chabanaud and Agama weidholzi Wettstein.

INTRODUCTION

UNTIL a few years ago it was believed that only three species of Agama occurred in
Nigeria: the ubiquitous Agama agama (Linnaeus) and the two smaller and less
common species A . sankaranica Chabanaud and A . gracilimembris Chabanaud. But
in 1963 Dr. Hilary Fry of the Ahmadu Bello University, Zaria drew my attention
to the existence of a form which he described as being of similar adult size to Agama
agama and as occurring on the smaller inselbergs in the Zaria region of northern Nigeria.
The adult male was said to be distinguished by four indigo or black, clearly defined
blotches on the throat. It was immediately recognized from an examination of an
adult male which Fry submitted that this Agama was not conspecific with any of the
three known species. With the co-operation of members of the staff of his depart-
ment intensive efforts were made to obtain further examples as well as ecological
data. The material resulting from these field studies contained more adult male
" blue throats " but not adult females belonging to the same form, instead examples
of Agama sankaranica, which in the field had been confused with the apparently
undescribed form, and gravid females of yet another apparently undescribed agama.
The absence in collections of females and immature individuals of this " blue throat "
continued until the following year when a Danish zoologist, Dr. Arne Schi0tz, who
was working in the British Museum recognized the male blue-throated agama as
being similar to ones he had collected in the Western Region. His large series,
obtained on inselbergs near Idanre and Igbetti, included both sexes and juveniles
and were in the collections of the Zoologiske Museum Copenhagen. A detailed study
there of his material enabled me to appreciate the difficulty that other collectors
had encountered in the field in distinguishing the females and immature individuals
from the superficially similar A. sankaranica and to elucidate the intraspecific
variation of all the Nigerian forms of Agama. The study was extended by referring
to type and other material in several institutions and to the British Museum's collec-
tions including those acquired during the 1962 Northern Nigeria expedition. Finally
the specimens on which Monard based the name Agama agama benueensis were
borrowed.

ZOOL. 17, 3. 3

68 A. G. C. GRAND I SON

The identity and relationships of Monard's Agama agama benueensis had not been
questioned until Wermuth (1967) in a footnote pointed out that benueensis is probably
no more than a local variety of Agama agama agama differing from it only in
minute details of toe length and pattern. The original description of benueensis,
extremely brief and inadequate, turned out to be based on a mixed series, some of
the individuals being conspecific with the Nigerian " blue-throated " form, while
others, strikingly different in their pattern and in the position of the nostril and with
a lower average midbody count are closely related to Agama agama and apparently
conspecific with a Nigerian population. The name benueensis is here applied to the
" blue-throat " form and a lectotype is designated. The information now available
on the ecology, morphology, distribution and variation of Agama agama, A . benue-
ensis, A . sankaranica and A . gracilimembris is given and a new species is described.
Descriptions are based on the observed variation in the examined material.
Throughout this paper the following abbreviations are used to refer to specimens
or collections.

B.M. British Museum (Natural History)

M.H.N.C.F. Museum d'Histoire Naturelle, La Chaux de Fonds
M.N.H.N. Museum National d'Histoire Naturelle, Paris
Z . M . C . Uni versitets Zoologiske Museum Copenhagen

METHODS AND TERMINOLOGY

The characters used in this study are defined as follows:

Body scales. The shape, size and direction of imbrication of the trunk and flank
scales, the extent of the keel relative to the axial length of each scale, from its anterior
border to its apex.

Microscopic structure of body scales. The number and position of the " hair-
bearing " scale organs on the scales of the mid trunk region were studied by removing
a series from the vertebral to the mid ventral row and mounting them according
to the method described by Underwood (1957).

Midbody scale count. The number of longitudinal scale rows around the middle
of the trunk.

Vertebral count. The number of transverse scale rows along the vertebral line
from forelimbs (on a level with the axillae) to hind limbs (groin).

Direction of imbrication of temporal and supratemporal scales. Direction of overlap
of the scales in these regions, the regions being defined as : temporal, the area between
the anterior rim of the ear and the postocular scales, overlying the temporal fossa
and the squamosal and the posterior half of the jugal bones; supratemporals, the
scales overlying the supratemporal arcade and the postorbital bone.

Body length. The distance from the tip of the snout to the anterior margin of the
vent, measured with dial calipers.

Head length. The distance measured with dial calipers from the tip of the snout
to the angle of the lower jaw at the posterior extremity of the articular.

Upper labials. The number of scales bordering the upper lip, counted from the
scale adjacent to the rostral to the corner of the mouth.

NIGERIAN AGAMA LIZARDS

69

Lower labials. The number of scales bordering the lower lip, counted from the
scale adjacent to the mental to the corner of the mouth.

Lamellae under fourth toe. The number of transversely enlarged scales under the
fourth toe from the base of the claw to the junction of the fourth toe with the third
toe.

Relative lengths of the toes. Comparisons of the first and fifth and of the third and
fourth are expressed as the number of lamellae contained in the difference in lengths
when the toes are laid flat and parallel to each other.

C

Q

FIG. i. Supratemporal and temporal areas of Agama; underlying bones indicated by

broken lines.

E Eye

J Jugal

M Maxilla

P Parietal

Po Postorbital bone

Q Quadrate

S Squamosal

St Supratemporal arcade

T Temporal fossa

Ty Tympanum

DERMAL SENSE ORGANS

Cohn (1914) was the first person to report on the occurrence of dermal sense organs
in Agama and he described in detail their histology as well as their external appearance
and distribution on various parts of the body. His study was restricted to Agama
colonorum (= A. agama). Subsequently Schmidt (1920) and Preiss (1922) examined
the scale organs of other species and genera of lizard, Schmidt extending his studies
on reptile receptors to the Gekkonidae, Iguanidae and Agamidae but limiting his
investigation of Agama to an Asian species A. sanguinolenta, which however exhibited
several differences from A . agama, as described by Cohn, in the number and position
of the organs on each trunk scale. It was not until 1937 and 1941 when Scortecci
made an extensive survey of the lenticular and " hair-bearing " sense organs in the
Agamidae that the density of these receptors and their positions on each scale were
shown to have some value as a species character. (The agamid material which

yo A. G. C. GRANDISON

Scortecci used belonged to nineteen genera and eighteen species, his nine species of
Agama all being African forms but none occurring in West Africa.) But despite the
breakthrough afforded by Scortecci's meticulous comparative studies the potential
of the receptor character for helping to distinguish agamid species was passed by,
yet Underwood (1957) in surveying the scale organs in pygopod lizards had stressed
that they would repay further study and Wayne King (1962) had effectively demon-
strated their usefulness in sorting out the taxonomy of Caribbean sphaerodactyl
geckos. The only attention recently given to these scale organs is by Miller and
Kasahara (1967) who have studied their innervation and who remark on the tactile
function of the " hair-bearing " receptor.

In species whose mode of life is known correlations might be established between
habits and the frequency with which the " hair " bearing organs occur, their position
on the scales, their distribution on the body surface and the extent to which the
" bristles " project beyond the edge of the scale. An investigation along these
lines seems merited but unfortunately through lack of adequate field observations
this could not be carried out on West African species of Agama.

It has been found in the present study that the species number and the position
of the " hair-bearing " organs on the trunk scale do not vary ontogeneticaly, nor
sexually, but that there is some individual variation which is discussed under the
species. Nor does this character vary geographically, at least not in four of the
Nigerian species where material from throughout their known ranges has been
examined. Whether there is geographic variation in Agama agama receptors has
still to be established but none is evident in the Nigerian specimens.

Another apparently reliable taxonomic character not previously employed in
keying out agamas is introduced into this study, namely the direction of imbrication
of the temporal and supratemporal scales. These two characters, receptors and
direction of imbrication, both of which are probably related to the animal's ecology,
seem to have greater significance in classification than some of the characters so
rigidly adhered to in the past in keys and descriptions of species of agama ; certainly
the results of studying them in the Nigerian species are encouraging and an extension
of study to other groups of agamid lizards may help elucidate the status and relation-
ships of the many forms that have been described.

Agama agama (Linnaeus)
(Map, PI. i, figs, c, d; Pis. 4, 5)
Lacerta agama Linnaeus, 1758 : 207.

MATERIAL EXAMINED. NIGERIA, Sokoto Prov., i mile north of Kware, 15 miles
north of Sokoto B.M. 1962.1609; Katsina Prov., Rimi village, 13 miles South-east
of Katsina B.M. 1962.1608; Kano Prov., Birnin Kudu B.M. 1962.1610; BornuProv.,
Maiduguri B.M. 1962.1611, Galtimare village, South of Maiduguri B.M. 1962.1612;
Zaria Prov., Kawo, 4 miles north of Kaduna B.M. 1962.1596-98, Kujama, 25 miles
South-east of Kaduna B.M. 1962.1599-1603, Zonkwa B.M. 1962.1604 and 1606,
Birnin Gwarri B.M. 1967.2207, Galma Fadama B.M. 1967.2208, Old City Zaria

NIGERIAN AGAMA LIZARDS 71

B.M. 1967.2209-2211, River Amethyst, Kaduna Road, Zaria B.M. 1967.2212;
Plateau Prov., Jos B.M. 1962.1613-54, 10 miles south of Jos on Miango Road B.M.
1962.1593-95, Shendam B.M. 1962.1574-92; Oyo Prov., Igbetti Z.M.C. R3656i,
36567, 36569, 36573, 36565, 36644, 36655, 36745, 36286-9; Ondo Prov., Idanre
Z.M.C. R36577-8, 36584, 36586.

DESCRIPTION. Body subcylindrical, little depressed above, between 3-3 and 3-8
times the head length. The body of adult males rather triangular in section, the
apex mid dorsal. Nostril on a line connecting the canthal supraciliaries with the
rostral/first labial suture, that is on the canthal ridge ; situated in the posterior half
or third of a strongly swollen, pear-shaped nasal shield and directed upwards, or
upwards and outwards. A keel usually present between the nostril and the anterior
edge of the nasal. On its posterior margin the nasal forms a suture, generally a
broad suture, with the first canthal supraciliary. The supraorbital scales and the
scales overlying the frontal and anterior portion of the parietal quite smooth, the
other scales on the head keeled, particularly so in juvenile specimens. The direction
of imbrication of the scales overlying the supratemporal vacuity is forwards but the
direction of the scales covering the postorbital bone alters to downwards and slightly
backwards. The scales overlying the temporal fossa, imbricate in the reverse
direction and between the two areas and along an oblique line, which marks the
position of the squamosal and the posterior limb of the jugal, is a single row of quite
strongly keeled juxtaposed scales (PI. I, fig. c and Text-fig i). Occipital enlarged, of
variable size, its greatest width generally half to three quarters the diameter of the
tympanum. Seven to twelve upper labials; a similar number of lower labials. A long,
low nuchal crest of approximately fourteen scales but no dorsal crest. A poorly
developed caudal crest present in adult males. The shape of the ear roughly a right-
angled triangle, the tympanic membrane rather superficial. No conspicuous fringe of
long, pointed scales along the anterior border of the tympanum but a few erect
scales which in juveniles may be only slightly raised. A row of erect or somewhat
spinous scales from the commissure of the lips to immediately behind and below the
tympanum where it joins with one or two (in adults invariably two) round groups of
spinose scales on the swollen area at the angle of the jaws. At the upper posterior
corner of the tympanum a less elevated and more elongated group of erect scales is
present. Midway between a point half way along the nuchal crest and the lower
posterior corner of the tympanum lies a small oval group of spinose scales; in front
of and also behind this group a very small group of erect scales usually insignificant
but if prominent the longest scales less than half the width of the tympanum. All
these groups of erect scales are considerably more pronounced and the scales longer
and more spinous in adult specimens, particularly in adult males, and in the juveniles
the groups may be quite inconspicuous. A strong gular fold present in both sexes.

Body scales homogeneous, those in the mid dorsal area flattened dorsoventrally,
their distal margins rounded, often only feebly keeled and with short mucrones.
Scales on the flanks more rhomboidal, particularly in adult males, the keels more
pronounced and terminating in upward turning short mucrones ; distal margins some-
what denticulate. Scales on either side of the nuchal crest small, barely mucronate,
approximately equal in size to those between the neck pit and ear.

72 A. G. C. GRANDISON

The sense organs of each trunk scale are situated at the distal margin or just below
the dorsodistal edge of the scale. They are not confined to the area around the base
of the mucrone but may extend along the entire distal aspect of the scale. No scale
organ on a trunk scale has more nor less than one " bristle ". The number of
receptors per scale varies from one to eight and while there is a tendency for the
number to increase on the flanks and then diminish towards the venter no definite
correlation could be established between the number of organs per scale and the
longitudinal scale row although the body scale rows bearing the highest density are
invariably between rows ten and sixteen. It is in this region of the trunk that the
scales often have a rhomboidal shape with a somewhat denticulated distal margin
and each organ is usually situated in an identation. Harris' (1963) statement that
sense organs are absent from the ventral body scales is incorrect. Each ventral
scale has one but no more than one " hair-bearing " sensory organ at the apex. The
smoother scales of the middorsum generally have one to four sensory organs per
scale.

Cohn (1914) found only three organs on each of the scales of the back and describes
the central one as placed somewhat to one side of the projecting tip of the keel.
He reports a similar number of receptors on flank scales and says that the scales on
the vertebral line may lack one or both lateral organs. He gives no indication of
the origin of his material which he identifies as Agama colonorum, and there seems
to be a strong possibility that a misidentification accounts for the much lower density
of trunk scale organs in his specimens.

Midbody scale counts of sixty-four Nigerian examples 59-77, the mean for thirty-
five females being 68-2 and for thirty males 66-2; vertebral count 38-55. Mature
males with eight to twelve, usually ten preanal pores. Eighteen lamellae under the
fourth toe. First toe shorter than the fifth by two to seven (usually three to six)
lamellae. Third toe shorter than fourth toe by one to two lamellae, very exceptionally
subequal. Tail approximately twice the body length, depressed at the base and
particularly so in adult males, its scales strongly keeled and mucronate and arranged
in whorls.

INTRASPECIFIC VARIATION. Loveridge (1936) in discussing intraspecific variation
in Agama agama remarked on the extraordinary wide range of midbody count,
60-80, quoted by Boulenger (1885). Presumably Boulenger's counts were based
solely on the material that was available in the British Museum at that time, namely
twenty-one West African examples many of which bore no more precise locality
than " West Africa ", and eight Angola and South West Africa examples, in other
words a small sample from only a very limited portion of the range of the species.
In the 1936 publication Loveridge demonstrated without regard to any sex difference
a gradual reduction in the same count from east to west (Uganda to Senegal) and
in the belief that populations occurring west of Ghana have a considerably lower
average count he suggested recognizing these individuals as belonging to a distinct
race, savatieri Rochebrune. The recognition of a western race based solely on this
character was ill conceived and hasty, particularly in view of Loveridge having at
his disposal only one individual from Ghana and none from the 1,000 mile stretch
between Ghana and the Cameroun. In 1956 Grandison in reporting on over one

NIGERIAN AGAMA LIZARDS

73

hundred Agama agama obtained in West Africa west of Ghana remarked on a south-
west to northwest reduction from 68 to 60 in their average midbody count (range
of the entire series 56-76). She assigned the individuals to A. agama savatieri,
having overlooked the statement in Loveridge's later paper (1941) that Hallowell's
name africana has priority over savatieri.

Daniel's (1961) counts (M 61-6 <$ : 65-1 $) on twenty adult Liberian individuals
which he identified as A. agama africana agree favourably with those given by
Loveridge (1936, 1941) for Liberian material but suggest that Loveridge's samples
in which the average number of scales round the body was sixty-two probably
consisted mainly of males.

The range of variation in midbody count in the sixty-five Nigerian examples
studied here runs almost the whole gamut of the variation said to occur in africana
but the average number of scales in males and females are intermediate between
those quoted by Loveridge for Cameroun and Uganda individuals of the nominate
form and those given by Loveridge, Daniel and Grandison for examples identified
as africana and obtained west of Ghana. The Nigerian, Ghanaian and eighteen
of the Senegal counts shown in the map are for adult specimens in the British Museum
collection.

U<^ 68-74(

m, 21, m 45 ) (

57-68(n 10 o; m 6/6) -72(n 35, m 66)
S8-72(n 10?,

\ 62-80(n 130*, m 70 V -4) /

<~ 68-74(n 7? 'm 71-41 [

n21, m65) 7 \ I !***

FIG. ^ Geographic varation in midbody count in Agama agama.

Thys van den Audenaerde (1963) in his study of Congolese species of Agama
examined large samples of A . agama from the Congo and compared his findings with
those of other authors for specimens from all regions in the range of the species.
His thorough review of the pertinent literature available on Agama agama emphasizes
the clinal sequence in midbody count from east to west although his argument is
to some extent weakened by blind acceptance of identifications by others, to unequal
sampling and to his not distinguishing between the counts for males and for females.
He includes in his range of variation in A . agama a specimen from the Bauchi Plateau
with ninety-two midbody scales that Grandison (1956) identified as A. sankaranica.
While the individual is not available for re-examination it seems more than likely
that it was incorrectly identified and is actually an example of A. benueensis, a
species closely related to sankaranica and in its juvenile livery resembling sank-
aranica but with a considerably higher midbody count. An A'ir specimen with a

74 A. G. C. GRANDISON

low count of 58 was also mentioned by Grandison (1956) but excluded from the table
of means; it too was probably incorrectly identified and may be an example of the
large scaled species Agama boueti Chabanaud. This species was described from Gao
on the Niger. Examination of type material shows that its trunk receptors are
similar in number and position to those of A. bibronii Dumeril of North West
Africa and are quite unlike any other West African form.

Thys van den Audenaerde maintains that development of the keel and mucrone is
correlated with body count, the lower the count the stronger the keeling, but I can
establish no such link in the Nigerian material where variation in this character
seems to be related to the proximity of sloughing. He further comments on the
geographical variation that occurs in A . agama colouration, in particular of the head,
throat and tail, which is perhaps correlated with the clinal gradient in scaling and
he draws attention to Angolan intermediates between the nominate race and the
Angolan race mucosoensis. A similar cline in the development of the nuchal crest
which both Stejneger (1893) and Thys van den Audenaerde (1963) have suggested
exists, has not yet been effectively demonstrated. Much more quantitative data
on large samples from throughout the range of the species would have to be compared,
sex with sex, if a true picture is to emerge. Meanwhile the only East African figures
available are those of Stejneger based on only six Kenya examples, and of Barbour
and Loveridge (1928) for eighteen adult Usambara specimens and in neither case
is the sex or the proportion of the sexes stated. Since in Nigeria at least, there is a
sex difference in the length and height of the crest, the spines being fewer and shorter
in the females, the East African figures are not necessarily comparable with those
for twenty adult male Nigerian individuals (13-17 : M 14-9).

Thus the abundance of races of A. agama that have been described reflects not
the occurrence of legitimate geographically localized subdivisions but accidents of
sampling or more likely the work of splitters. There would appear to be no evidence
of geographic isolates but only of marked individual variation within a population
and Thys van den Audenaerde's conclusion that Agama agama is a large polymorphic
species rather than a polytypic species would seem to be the logical one.

COLOUR. Immature individuals and adult females are similar in colouration.
In their reproductive colour phase the head is a medium brown or brown with green
tinges, the back fawn or brown with darker brown marks of variable shape and
position but often in the form of rather indistinct diamond-shaped outlines along
the vertebral region. The upper part of the head, the temporal region and the neck
have pale green spots and short longitudinal stripes of the same colour. The
undersurfaces are a grey-white. Mature males that are dominant over territories
are characterized in their reproductive colour phase by an orange head and neck,
indigo blue body and by a median orange segment on a black-tipped tail. Other
colour phases occur and these as well as the reproductive colour phase are admirably
and fully described by Harris (1964).

ECOLOGY. In Nigeria wherever the savanna and forest have been disturbed by
human agency, bridges built, houses and shacks erected, trees felled, rubbish dumped,
Agama agama seems to have established territories. It is very much a commensal

NIGERIAN AGAMA LIZARDS 75

of man and is common around native houses, farm buildings, litter, wood piles and
along paths. The species is also common in the boulder strewn country flanking
the road from Jos to Miango. Schi0tz (personal communication) says that its
habitat preferences do not include flat uniform savanna, nor large rocks, nor dense
forest but as soon as such areas acquire human habitations or are in other ways
disturbed by man Agama agama takes up residence. While this is largely true, the
rocky mass of Kujama Hill south of Kaduna supports a number of Agama agama yet
on the bare rock surfaces of the vast gneiss domes 1,000-3,000 ft. high at Igbetti
and Idanre Agama agama is absent, its place being taken by Agama benueensis.

RANGE. Senegal to Ethiopia, southwards to Tanzania and Angola.

Agama paragama sp. n.

(PI. i, figs, a, b; Pis. 4, 5)

Agama agama benueensis Monard 1951 : 131 (part).

MATERIAL EXAMINED. HOLOTYPE: NIGERIA, Zaria Prov., mile Southwest of
Ahmadu Bello University, Zaria City, collected by C. H. Fry from Parkia tree on
17.11.1964, B.M. 1967.2215 $.

PARATYPES: NIGERIA, Zaria Prov., same locality as holotype B.M. 1967.2216,
Mile 7, Zaria Road B.M. 1967.2214, Mile 5, Zaria Road B.M. 1967.2213, Ahmadu
Bello University campus B.M. 1967.2217, no information, probably in vicinity of
Zaria City B.M. 1967.2218, Birnin Gwarri, 62 miles Southwest of Funtua B.M. 1968.
497, Zaria City B.M. 1968.482-483, Veterinary Unit, Zonkwa B.M. 1962.1605,
B.M. 1962.1607; BornuProv., Potiskum B.M. 1968.498, Yo B.M. 1968.496; Plateau
Prov., Naraguta, Jos B.M. 1968.484-485. CAMEROUN, Bangouve M.H.N.C.F.
1162, Rei Bouba M.H.N.C.F. 1432, 1440, 1459, Mayo Sala M.H.N.C.F. 1374,
Ngaouyanga M.H.N.C.F. 965 (six of the paralectotypes of Agama agama benueensis.}

These individuals differ from A . agama not only in body size, proportions, greater
spinosity and in lower average midbody and vertebral counts but in an absence of
sexual dimorphism in size. They resemble A . agama in the position of the nostril being
on the canthal ridge and in having multiple sense organs on the distal edges of the
scales of the flanks and the two forms are undoubtedly closely related but being
sympatric in Nigeria specific status is given to paragama.

Relatively few examples of paragama have been collected and the form is so far
known only from northern Cameroun and from the Northern Region of Nigeria,
although one specimen obtained by C. H. Fry in a crevice 10 ft. up a Terminalia
tree in the Botanical Gardens, University College, Ibadan was thought by him to
belong to this form; the specimen has not been seen by me. Although situated in
the high forest belt Ibadan has many clearings that have the character of cultivated
savanna woodland and support typical savanna species. Schi0tz (1963) describes
the vegetation of the University College campus as " dry forest and culture savanna ".
It is possible that where suitable habitats are available this species, like Agama
benueensis, penetrates the Western Region's forest belt.

76 A. G. C. GRANDISON

DESCRIPTION OF HOLOTYPE. Body subcylindrical, depressed above, 3-6 times the
head length. Nostril on the canthal ridge, situated in the posterior half of a swollen,
pear-shaped nasal shield and directed laterosuperiorly. Nasal keeled from its
anterior tip to the upper rim of the nostril, forming a suture on its posterior margin
with the first canthal supraciliary. The supraorbital scales and the scales overlying
the frontal and the anterior portion of the parietal quite smooth, remaining head
scales keeled. The direction of imbrication of the supratemporal scales is forwards
and slightly upwards; anteroventrally the scales overlying the postorbital bone
face downwards and backwards. The scales immediately in front of the ear, that
is those overlying the temporal fossa, imbricate in the reverse direction to those
covering the lower postorbital and between the two areas and along an oblique line
marking the position of the squamosal and posterior limb of the jugal is a single row
of large, keeled, juxtaposed scales. Occipital enlarged, its greatest width equal
to three-quarters the horizontal diameter of the tympanum. Upper labials 8 : 9
(one divided) ; lower labials 9 : 10. A short but conspicuous nuchal crest formed
from nine, long, conical scales, the longest equal to three-quarters the width of the
tympanum; no dorsal nor caudal crest. Ear triangular. On anterior margin of
ear a fringe of two to three long, pointed scales, on upper posterior border one long,
pointed scale surrounded by several erect scales (this group barely developed on
left side of head). At the posterior extremity of the lower jaw, that is behind and
below the ear, a small, round group of spinose scales of which one scale is three to
four times longer than the others. A similar group immediately above. Between
a point midway along the nuchal crest and the lower, posterior corner of the tymp-
anum lies a prominent round tuft of long spinose scales the longest scale three-
quarters the tympanic diameter; in front (right side only), and also behind, a small,
round but insignificant group of erect scales. (The lengths of the scales comprising
the nuchal crest and the groups of spinose neck and ear scales are considerably longer
and more pronounced in male specimens : see note on variation.) A well developed
gular fold.

Body scales homogeneous, the longitudinal rows converging towards the midline,
scales on the dorsum and flanks equally strongly keeled and mucronate ; the mucrones
of the flank scales rather broad at their base, tapering rapidly and rising at an angle
to the scale. Distal margins of the body scales rounded on each side of the mucrones.
Ventral scales smooth. Scales on either side of the nuchal crest with strong,
compressed keels and long, almost erect, mucrones, the scales at least twice the size of
those between the neck pit and ear.

The maximum number of sense organs on the middorsal and flank scales is nine,
the minimum three. They are situated just under the dorsodistal margin and may
extend as far as the " shoulders " of the scale. No scale organ on a trunk scale has more
nor less than one " bristle ". The highest density of receptors on midtrunk scales
occurs on longitudinal rows 3-5. Each ventral scale bears a receptor at its apex.

Midbody scale count 60, vertebral count 30, eighteen lamellae under the fourth toe.
First toe four or five lamellae shorter than the fifth toe, third toe half to one lamella
shorter than the fourth.

Tail one and a quarter times body length (tip of tail missing), depressed at base,

NIGERIAN AGAMA LIZARDS 77

only very slightly compressed distally, its scales keeled and strongly mucronate
and arranged in whorls.
Length of body 94-0 mm.

VARIATION. The following variation in meristic characters in the paratype series
was noted. Nigerian individuals: midbody scale count 50-66 (M 58-5), vertebral
count 26-34 (M 30-1), number of lamellae under the fourth toe seventeen to twenty-
one, first toe shorter than fifth toe by between three and five lamellae, third toe half to
two, usually one to two lamellae shorter than fourth toe. Differences in the develop-
ment of the rosettes of neck and ear spines and in the lengths of the scales forming
the nuchal crest are attributable to sex and maturity, in adult males the longest of
the pointed nuchal crest scales being as much as equal to the width of the tympanum,
likewise the most prominent scales in the neck tufts, whereas in females they are
less than three-quarters the diameter of the tympanum. The number of scales form-
ing the nuchal crest varies from eight to ten, the usual number being eight. The
density and distribution of " hair-bearing " receptors on the midbody scales of the
paratypes is similar to the holotype. Sex dichromatism is discussed below. The
tail of the adult male is considerably more compressed than that of the female and
has a slight crest ; the base of the tail is depressed but not to the same extent as in
Agama agama.

Juvenile A. par agama in which the neck and ear rosettes and nuchal crest are
insignificant can be readily distinguished from young individuals of A. agama by
the relative size of the scales above and below the central neck rosette and by the
roundness of the rosette. Further distinguishing features are the pattern, lower
vertebral count, gradual tapering of the tail and less depressed base to the tail.

All the Cameroun examples, except an adult male (Mayo Sala M.N.H.C.F. 1374),
fall within the range of variation in midbody count of the Nigerian individuals but
they have a slightly higher number of transverse rows on the trunk, their vertebral
count being 36-38. The Mayo Sala example with seventy-two midbody scale rows
is well outside the normal range of variation in the species yet in its other morpho-
logical characters it agrees with par agama.

COLOUR. Male. In life the upper surfaces of the body and limbs of the adult <$
are a metallic blue with white flecks. On the back the flecks tend to be arranged
in narrow transverse lines, no more than one scale in width, with approximately
eight lines between fore and hind limbs. The vertebral zone is whitish, the head
chalk white, the proximal two-thirds of the tail whitish, the tail tip black. Except
for the whitish areas the general appearance of the $ is rather similar to the reproduc-
tive colour phase of a male Agama agama. Harris (1964) describes in detail the
various colour phases of both <$ and $ Agama agama and while he refers to a geograph-
ical variation in the colour of the head of adult males, northern individuals (Kano)
having a deep yellow while more southern populations have either orange or vermilion,
he makes no mention of any adult male agama in which the orange or yellow tones
are replaced by white.

FEMALE. None of the colour phases described by Harris for female A. agama
occurring in Nigeria corresponds to, or in any way approaches the colour pattern

78 A. G. C. GRANDISON

of female A. paragama which is characterized by four or five pairs of more or less
clearly defined, large, round, brown spots on a lighter brown (orange in life) background
on the dorsum between fore and hind limbs. In those specimens in which the spots
are ill-defined the lateral halves of the spots are indicated by narrow, dark brown
outlines. The flanks and the upper surfaces of the limbs and tail are of a similar
shade of brown to the spots; the head is mottled brown. In both sexes the throat,
and in some specimens also the sides of the neck and chest, has a dark network on a
cream ground which takes the form of isolated, round, cream spots and is dissimilar
to the gular pattern of A. agama which tends rather to consist of a longitudinal
arrangement of dark lines or blotches (Harris, 1964). The distinctness of the gular
spotting does not appear to bear any relation to the maturity or sex of the individual,
for example one gravid female has a pronounced pattern while in another a pattern
is barely discernible. No pattern whatsoever is present on the throat of one adult
male but a distinct network is present in another mature male

Adult female paragama are larger and considerably more robust than female
agama. Three gravid female paragama vary in body size from 96-107 mm. while
the gravid female agama examined in this study vary from 86-91 mm. Harris
(1964) records 97 mm. as the average body length of sixty-eight Nigerian adult
female agama and Chapman & Chapman (1964) regard all Ghanaian female A . agama
of about 90 mm. as mature. Adult male paragama are considerably smaller
(99-108 mm.) than mature male agama obtained in the same general area (107-137
mm.).

ECOLOGY. Little information on the ecology of paragama is available. Fry
(in litt.) states that the species is to be found on the trunks and lower limbs of shade
trees and collectors have suggested that it occupies a different ecological niche
from A. agama but only the specimens collected at Mile 7 Zaria, Potiskum and Birnin
Gwarri bear information that suggest an arboreal existence (Mango and Parkia
trees) ; the Yo specimen, a juvenile, was taken on the walls of a house and the rest
of the examined material have no biotope data.

RANGE. West Africa: Northern Region of Nigeria to northern Cameroun.

Agama benueensis Monard
(PL 2, figs, a-d; Pis. 4, 5)

Agama agama benueensis Monard, 1951 : 131 (part).

Agama agama boensis Monard, 1951 : 130 (part: males).

? Agama sankaranica: Grandison, 1956 : 231 (part: Bauchi specimen).

MATERIAL EXAMINED. CAMEROUN, Upper Benue, Ngaouyanga M.H.N.C.F.
956-7, Sakdje M.H.N.C.F. 1269, Mayo Sala M.H.N.C.F. 1373, Rei Bouba M.H.N.C.F.
1442, 1460, 1517 (seven of the paralectotypes of A. a. benueensis), Ngaouyanga
M.H.N.C.F. 961 (designated lectotype of A. a. benueensis). NIGERIA, Oyo Prov.,

Igbetti Z.M.C. R 36342, 36345, 36347-51. 36353, 36389, 3639 2 > 36394, 36397, 36399*
36401, 36599, 36601-12, 36614-20, 36661-2, 36702-3; Ondo Prov., Orosuta, Idanre
Z.M.C. no number, Idanre Z.M.C. R 36576, 36579, 36580-2, 36585, 36588-9; Zaria

NIGERIAN AGAMA LIZARDS 79

Prov., R. Amethyst, Kaduna Road B.M. 1967.2234-8, B.M. 1967.2263, Ahmadu
Bello University campus, Zaria B.M. 1967.2239-44, Siberia, A.B. University campus
Zaria B.M. 1967.2254-7, Old City, Zaria B.M. 1967.2245-7, B.M. 1967.2258-61,
Mile 5, Zaria Road B.M. 1967.2248-52, B.M. 1967.2253, Zaria B.M. 1967.2262,
Near Oomawa, at foot of Kwatarakwashi rock B.M. 1968.490-495; Plateau Prov.,
Jos B.M. 1968.486-487, Hillcrest School, Jos B.M. 1968.488.

REMARKS. The whereabouts of four of the type specimens of Agama a. benueensis
(959, 1372, 1441 and 955) listed by Monard (1951) are not known at present. Thys
van den Audenaerde tells me he believes that part of the Monard material originally
lodged in the herpetological collection at La Chaux de Fonds was transferred to other
European institutions and that the missing Cameroun agamas may have been among
it. The thirteen individuals from the benueensis type series that are available and
have been re-examined belong to two morphologically very different forms, and as
referred to in the introduction to this paper, the name benueensis is applied to the
blue-throated form. The other form is described as A. paragama sp. n. The
Monard specimens here assigned to A. benueensis are those males from the Upper
Benue that are described by him (1951) as having " une tache noiratre a la gorge ",
" une tache foncee a la gorge " and " une raie vertebrale claire ", as well as four
females and a juvenile male that have a feebly developed pattern characteristic of
immature and female Nigerian benueensis, and two males from Tibati, Adamaoua
Plateau which he assigned to A, agama boensis. The Tibati males, M.H.N.C.F. 673,
784, have been examined by Dr. Braestrup who reports (in litt.) that both individuals
have chin as well as gular spots. The presence of the gular spots, even if faint,
identifies them as benueensis for this species is the only West African agama that
has a bilobed blotch in this region in the adult male. In the Tibati males the midbody
count is about 74, the vertebral counts 45 and 47 which accord with Cameroun
examples of benueensis. Neither I nor Dr. Braestrup has seen the female, M.H.N.C.F.
726, which Monard also referred to A. a. boensis and its identification remains in
doubt.

DIAGNOSIS. A large sized Agama (body length of $ up to 113-0 mm., of $ to
74-0 mm.) related to A. sankaranica and characterized by a high average number of
midbody scale rows, 74-98 (M 88 : N 71) in Nigerian populations, ventrally directed
temporal scales and marked sexual dimorphism. Dorsal trunk scales each with
one to three single " hair-bearing " scale organs close to base of mucrone, nostril below
canthal ridge; in these respects similar to A. sankaranica.

DESCRIPTION OF LECTOTYPE M.H.N.C.F. 961, <$. Body subcylindrical, little
depressed above, 3-7 times the head length. Nostril below a line connecting the
canthal-supraciliaries with the rostral-labial suture, its aperture round, situated in
the posterior third of an elongated, pear-shaped nasal, directed superior-laterally.
From the upper rim of the nostril to the front of the nasal shield is a distinct ridge.
Canthal-supraciliaries 8:9, the anterior forming broad sutures with the nasals at
their posterior borders. Supraorbitals quite smooth, supratemporals distinctly
keeled, temporals only very slightly keeled. The scales overlying the supratemporal
arcade and the postorbital bone directed forwards and slightly upwards; ventrally

8o A. G. C. GRANDISON

the imbrication of the scales covering the posterior part of the jugal, the broad
portion behind and below the eye, as well as those over the temporal fossa directed
downwards and slightly backwards; no row of keeled juxtaposed scales. Occipital
plate enlarged, broadly oval, its length equivalent to the horizontal diameter of the
tympanum. Seven to ten upper labials, eight to twelve lower labials. Tufts of
enlarged, conical, rather flat scales around the ear, one, consisting of three scales,
on the anterior border, one on the upper posterior border, on the lower posterior
border a group of about four to five more prominent conical scales and below this
group a curved row of about five flattish conical scales. At the side of the neck
midway between the tympanum and a shallow neck pit, but a little above, a rosette
of spines, the longest scale in the rosette two-thirds the length of the nasal. Behind
and a little below this rosette and situated above the neck pit an oblique or vertical
line of conical scales. Above and slightly behind the tympanum a prominent
erect scale surrounded by a few lower but erect scales. Ear subtriangular, its
horizontal diameter equal to the distance from the tip of the snout to the middle
of the nasal. A gular pouch present. Nuchal crest low, formed from about ten
conical scales; no dorsal nor caudal crest. Dorsal scales imbricate, homogeneous,
strongly converging on the vertebral line, strongly keeled but the keel not markedly
compressed, mucrone finely tapering, ventrals and subcaudals smooth.

A maximum of two scale organs on each body scale but usually only one and
situated under the dorsodistal edge of the scale close to the base of the mucrone.
Each scale organ bears a single " hair ".

Midbody scale count 78; vertebral count 46; preanal pores in two transverse
rows, 13 anteriorly, 15 posteriorly. The adpressed hind limb reaches to posterior
border of the tympanum. Nineteen lamellae under the fourth toe. First toe three
or four lamellae shorter than fifth toe, third toe one lamella shorter than fourth toe.

Tail slightly depressed at the base; broken. Caudal scales keeled and mucronate,
arranged in distinct whorls.

Body size 94-0 mm.

COLOUR IN ALCOHOL. Apart from a conspicuous whitish vertebral line approxi-
mately three scales wide and extending from the nape to the base of the tail there is
little colour pattern discernible. Any pattern that may have been present on the
flanks of the living animal has become obscured by preservation and sloughing and
only a slight mottling of cream and dark brown is now evident. The throat bears
a large, blackish, bilobed patch at the level of the gular fold ; in front of this patch
and extending on to the chin are several small, dark spots on a cream background.
The rest of the undersurfaces are a grey-cream.

COLOUR IN LIFE OF NIGERIAN EXAMPLES. Dorsum grey or brown with whitish
dark-edged ocelli on the flanks which are arranged in two or three longitudinal rows,
the spots in the outer, or middle in the case of individuals with three rows on each
side, generally larger than the others. Across the back from the shoulders to the
tail a series of dark chocolate brown bands which, particularly in juveniles, are
considerably broader and often diamond-shaped on the vertebral line and enclose a
narrow, longitudinal, whitish streak. The streaks become confluent in some adult

NIGERIAN AGAMA LIZARDS 81

males and females (see sex dimorphism) and form a diffuse yellowish vertebral stripe.
Schi0tz (personal communication) describes the entire dorsum and belly of adult
males as having a distinct bluish tinge, but not of such an intense blue as the Agama
agama dominant male, while the chin and throat are yellow with two to four dark
blue or black patches. The upper surface of the male's head is irregularly mottled
green; the tail is greenish with indistinct yellow bands.

The ground colour of the female varies according to the colour of the rocks or
soil in its habitat. At maturity the transverse bands on the trunk are reddish brown
and the sides of the head are greenish. The ventral surface of both the female and
the juvenile is white but the chin and gular region of most individuals has a network
of dark lines forming irregular spots.

INTRASPECIFIC VARIATION. An analysis of meristic variation in the material
examined shows some geographical differences. The samples from the western
region (Igbetti and Idanre) have the highest midbody and vertebral counts, midbody
84-98 (M 91-3 : N 34), those from the Jos Plateau and the Cameroun the lowest,
68-82 (M 76 : N n) while the gradation in the Zaria Province population almost
spans these two extremes, 74-96 (M 85-8 : N 34). The variation in these characters
for the entire series is : midbody 68-98 (M 87-0 : N 79), vertebral 41-56 (486 :
N 76). The correctness of the identification of the individuals at the extremes of
the variation is not in doubt, a combination of other characters such as throat mark-
ings, position of nostril, trunk scale organs and direction of imbrication of the post-
orbital/temporal scales assigning them unquestionably to benueensis. Variation in
the number of fourth toe lamellae (nineteen to twenty-three) and in the relative
lengths of the first and fifth, and third and fourth toes is independent of the variation
observed in midbody and vertebral counts and appears to have no geographical nor
ontogenetic significance. The third toe may equal the fourth but generally it is
shorter by half to one lamella. Variation in the relative lengths of the first and fifth
toes is greater; the first is always shorter than the fifth but the difference may be as
many as four lamellae.

In the majority of the examples studied the lateral head scales between the eye
and the ear all imbricate downwards or downwards and slightly backwards but in a
few individuals, juveniles as well as adults, the scales in the lower posterior quarter
of this area, that is those overlying the temporal fossa are directed forwards and
upwards, in other words in a similar direction to those of Agama agama; however
they are not as strongly keeled as in agama nor are they separated from the post-
orbital /supratemporal scales by an oblique row of juxtaposed scales.

Scales on either side of the nuchal crest are not or only slightly mucronate and are
only slightly keeled. They are considerably larger than the scales below the central
group of spines on the side of the neck.

SEXUAL DIMORPHISM AND DICHROMATISM. Some sexual dichromatism is evident,
notably in the conspicuous gular and chin spots which are a characteristic feature
of the adult male. The twin gular spots develop before the chin spots and first
appear as a darkening of the median gular region which later becomes an intense
black or navy bilobed blotch; occasionally the lobes are separated on the midline.

ZOOL. 17, 3. 4

82 A. G. C. GRANDISON

The gular spots do not develop until the male reaches a body size of over 73 mm.,
although there is one exception among the fifty-six males examined, a 69-7 mm.
individual collected at Igbetti. By 80 mm. most of the males have developed two
additional dark spots; these are on the chin and somewhat oval and parallel to each
other on either side of the median line. The development of these gular and chin
spots in adult male benueensis does not seem to be seasonal; they are present in
Nigerian individuals collected in January, February, March, June, July and August.
The three largest males in the Cameroun series (72-5, 78-3 and 74-0 mm.) which were
obtained in July and in September have only gular spots.

Adult males retain some evidence of the juvenile pattern; generally the two longi-
tudinal rows of ocelli along each flank are discernible, although feebly, as well as
remnants of the five dark transverse lines in the mid dorsal region on each side of the
midline from shoulders to groin but the light coloured vertebral streaks, a character-
istic feature of the juvenile livery, invariably coalesce and form an ill defined light
vertebral zone extending from the nape to the tail. The female sample is insufficient
to be certain whether a similar light vertebral zone always develops at maturity.
Only six of the fifteen females are gravid and of these six only the two Cameroun
examples (M.N.H.C.F. 1460, 1269) have a clearly defined continuous light line.
In the other examples the vertebral pattern is obscure in all but one of the Igbetti
examples (Z.M.C. 36351). In both sexes the dorsolateral series of ocelli becomes
progressively less distinct as the individuals mature. Egg bearing females were
obtained in July, August and September. Preanal pores are usually ten to twelve
in a single series but in three Idanre males, the only male benueensis collected there,
and in the lectotype the pores are in double series 13 + 14, 10 + 13, 8 + 13, 13 + 15
respectively.

ECOLOGY. In the western region of Nigeria Schi0tz (personal communication)
says that A. benueensis is strictly confined to large rocky outcrops such as the vast
gneiss inselbergs where it is common both in the savanna (Igbetti) and in the forest
(Idanre). Other ecological niches that he searched in southern Nigeria, such as
mounds of boulders, small rocky areas, produced no examples. He noted that
benueensis hides in rock crevices and among vegetation at the perimeter of an insel-
berg but conducts its feeding and sexual activities on the actual surface of the rock.
Farther north on the Jos Plateau the species is much less abundant but it is again
quite common in Zaria Province on the inselbergs around Zaria City as well as in a
variety of other habitats such as on mounds of laterite on open farmland in the
Galma Fadama, a locality some miles from the nearest inselbergs, on sandy river
banks (Amethyst River) and in sandy stream beds (R. B. Walker, personal communi-
cation) . Near Sabon Gari, in an area of scrap cars and garbage tips, both A . benueensis
and A. agama were taken and no specific habitat preference was noted but Schi0tz
maintains that at Igbetti A. benueensis, A. sankaranica and A. agama can be seen
virtually only a few feet from each other yet each still adhering to its preferred
habitat.

A careful search by Schi0tz of the Shai Hills in Ghana and suitable localities in the
Eastern Region of Nigeria failed to produce any benueensis and he believes the species

NIGERIAN AGAMA LIZARDS 83

may also be absent from the Bamenda region of the Cameroun. Agama benueensis
seems to be endemic to the Western and Northern regions of Nigeria and the upper
Benue valley in the Cameroun.

Monard (1951) does not record the biotopes for the type series of benueensis and
he describes only the general areas around the villages. The I : 1,000,000 maps of the
Upper Benue show the villages where benueensis were taken as being in an area
deeply dissected by the Benue and its tributaries and at altitudes varying from
740 ft. to 1,400 ft.; only Ngaouyanga is on a high steep-sided wooded platform,
the other localities are in or at the sides of valleys in woodland savannah.

RANGE. West Africa: Western and Northern Regions of Nigeria and northern
Cameroun.

Agama sankaranica Chabanaud
(PI. 3, figs, a, b; Pis. 4, 5)

Agama sankaranica Chabanaud 1918 : 105.

Agama boensis Monard 1940 : 155 (part: adults only).

MATERIAL EXAMINED. PORTUGUESE GUINEA, Madina Boe M.H.N.C.F. 865, 867-8
(three of the syntypes of Agama boensis). REPUBLIC OF GUINEA, Beyla M.N.H.N.
21.309-316, Kankan M.N.H.N. 21.297-298, Kerouane M.N.H.N. 21.299-308,
B.M. 1921.11.12.1-5. MALI, Moussaia, Sankaran M.N.H.N. 1901.395 (holotype of
Agama sankaranica) . GHANA, B.M. 1930 .6.9.8 Korley Bu, Accra B.M. 1931 .5.6.3,
B.M. 1932.6.1.5-7, Tamale B.M. 1927.9.27.199, Tafo B.M. 1962.910. NIGERIA,
Benue Prov., Makurdi B.M. 1937.12.4.14; Plateau Prov ., Shendam B.M. 1962.1573;
Kano Prov., Kano B.M. 1961.2069, B.M. 1962.567 (no. 567 later skeletonized);
Zaria Prov., B.M. 1967.2206, Rimi, South-east of Kaduna B.M. 1962.1572, Zonkwa
B.M. 1962.1571, Ahmadu Bello University campus, Zaria B.M. 1967.2205, 14 miles
South of Zaria B.M. 1967.2204; Lagos Prov., Lagos B.M. 1960.1.7.28; Oyo Prov.,
Igbetti Z.M.C. R 36386, 36388, 3639. 36393, 36395-6, 36398, 36403, 36405, 36407,
36597, 36607, 36697-8, 36691, 36758.

DIAGNOSIS. A medium sized Agama (body length of adult males averaging
66 mm., of mature females 76 mm.) belonging to the group of agamas that have the
nostril below the canthal ridge, homogeneous scaling and dermal sense organs not
exceeding three in number on each trunk scale. Related to Agama benueensis
Monard but distinguished by a lower average midbody count (69-6) and by dorsally
directed temporal scales.

DESCRIPTION. Body subcylindrical, scarcely depressed above, 3-3-7 times the
head length. Nostril below the canthal edge, situated in the posterior half of a
slightly swollen, oval nasal, directed laterally. No post nasal separating the first
canthal supraciliary from the nasal. Scales of the head keeled; the interorbital
scales considerably smaller than the supraorbitals ; imbrication of temporals directed
upwards and of supratemporals directed upwards and forwards (PI. 3 fig. b)
Occipital large, its greatest width equivalent to or as much as one and a half times
greater than the diameter of the tympanum. Eight to twelve upper labials; a

ZOOL. 17, 3. 4

84 A. G. C. GRANDISON

similar number of lower labials. Usually three tufts of long, conical spines on pos-
terior border of ear ; behind and above these a single group of spines and yet farther
back and just above a shallow neck pit another group of somewhat shorter spines.
A weakly developed gular pouch. A low nuchal but no dorsal or caudal crest.
Dorsal scales homogeneous, broadly rounded, strongly keeled and mucronate, the
sharp high keel extending the length of each scale, the mucrone very slender. Scales
on the flanks slightly smaller than dorsals, similarly keeled and mucronate. Ventrals
keeled only in juveniles. " Hair-bearing " sensory pits on the dorsal midtrunk
scales confined to around the base of the mucrones, at or just under the dorsodistal edges
of the scales. The maximum number of scale organs on each scale is three but
invariably no more than two are present. In the vertebral region at midbody there
are generally two " hair-bearing " pits, each with a single hair-like projection, the
one pit close to the base of the mucrone or under the mucrone and concealed in dorsal
view, the other on the opposite side of the mucrone but near the apex of the scale.
On more lateral scales there is a tendency for only the more lateral, that is lateral
with respect to the mucrone, to develop.

The number and position of the scale organs on the dorsal scales bear a close
similarity to the condition obtaining in East African rueppelli and described and
figured by Scortecci (1937). Midbody scale count 64-78 (M 69-6 : N 61); vertebral
count 32-46 (M 38-8 : N 61). Mature males with eight to twelve usually ten preanal
pores in a single transverse row. Fifteen to twenty-one lamellae under the fourth
toe. First and fifth toes subequal or first toe one lamella shorter. Third toe usually
subequal to fourth toe, rarely half to one lamella shorter or longer than fourth.
Tail one and three quarter times the body length, depressed at the base, slender and
cylindrical distally, its scales strongly keeled and very distinctly mucronate, not
arranged in distinct whorls.

COLOUR IN ALCOHOL. Upper parts reddish brown. Invariably a conspicuous
yellowish or russet vertebral stripe from behind the head to the proximal third of the
tail, much broader and always prominent in the cervical region but narrowing rapidly
to a width of approximately two scales on the rest of the body. On each side of this
stripe and from nape to tail broad, dark brown, transverse bands, six between the
nape and hind limbs. At the lateral extremities of these bands light-coloured ocelli
or elongated blotches which in some specimens coalesce to form a light dorsolateral
stripe. In some examples the vertebral stripe widens at the level of each transverse
band to form light diamond-shaped areas. Halfgrown individuals occasionally
have indistinct marbling on the flanks below the ocelli or dorsolateral stripe. In
one mature male, collected at Zaria, the transverse bands are broken up into irregu-
larly shaped blotches. Between the eyes a thin dark transverse line is present;
four similar lines radiating from the eyes to the lips are usually evident. Limbs
indistinctly cross-barred with narrow dark lines.

SEXUAL DIMORPHISM. There are no obvious external differences between the
females and immature males. Adult males have eight to twelve preanal pores and
also broken, longitudinal, blue stripes on the chin and throat which become more
prominent with maturity and often enclose a large, dark blue, median gular spot

NIGERIAN AGAMA LIZARDS 85

and a lozenge-shaped chin blotch. The rosettes of neck spines and the lips may also
be blue in mature males. Adult females are on average rather larger than adult
males. The body length of mature females varies from 68-85 mm - (average 76-2 mm.)
while males vary from 62-76 mm. (average 66-4 mm.).

REMARKS. The examined material consists of examples from throughout the
known range of the species. No evidence of geographical variation can be demon-
strated. The examples from the extreme west of the geographical range of sank-
aranica are the three adults, two males and one female, in the series of syntypes of
Agama boensis Monard. These individuals unquestionably belong to sankaranica.
The throats of the males have the blue blotches and stripes that are typical of mature
male sankaranica, furthermore the scale counts and other morphological characters
including the density and distribution of trunk receptors agree well with sankaranica
although one male with a midbody count of 64 is exceptional in having 20 : 21
lamellae under the fourth toe, which is outside the range for all other specimens
examined (fifteen to nineteen). The fourth syntype of boensis, a juvenile, lacks a
nuchal crest and clusters of neck spines and has a single apical scale organ on each
body scale; it is conspecific with Agama weidholzi Wettstein. (Grandison, in
press.) The Cameroun individuals assigned by Monard (1951) to A. a. boensis are
discussed in the section on A. benueensis.

ECOLOGY. In the field this species has often been confused with female and imma-
ture Agama benueensis and as a result some ecological information claimed to be on
A. sankaranica has had to be disregarded or treated with suspicion in this study.
Dr. Arne Schi0tz' reliable field notes and comments make it clear that he readily
distinguishes sankaranica from benueensis and his notes correspond with my own
field observations in stating that unlike all other Nigerian species of Agama, with the
exception of gracilimembris, sankaranica does not form family groups but is a solitary
creature. The species has been observed and captured in ploughed farmland and
in maize and cassava plantations when running along the rutted ground, also on
paths in grass covered savanna. Schi0tz' notes state that he saw the species on the
ground only in grass covered savanna or in rather dense tree savanna and never in
forest nor on rocks. The localities at which examples of A. sankaranica have been
collected are, with one exception, in the Doka and True Guinea woodland belts from
Nigeria westwards to Portuguese Guinea; the exception is an individual taken at
Kano which is towards the southern limit of Sudan woodland.

RANGE. West Africa: Portuguese Guinea to Nigeria.

Agama gracilimembris Chabanaud

(PI. 3, ngs. c, d; PI. 6)
Agama gracilimembris Chabanaud, 1918 : 106.

MATERIAL EXAMINED. DAHOMEY, M.N.H.N. 04.114-5 (syntypes). NIGERIA,
Benue Prov., Lafia B.M. 1938.3.1.47, Wukari B.M. 1938.3.1.48-9; Plateau Prov.,
Shendam B.M. 1962.1570; Zaria Prov., Zonkwa B.M. 1961.949, B.M. 1962.1569,
Samara Bush, Zaria (on exchange to Vienna Mus.) ; OyoProv., Igbetti Z.M.C. R 36654,
R 36701, R 36263; Kano Prov., Kano B.M. 1961.2067-8, B.M. 1962.566, CENTRAL

86 A. G. C. GRANDISON

AFRICAN REP., " Pays des Senoussi " M.N.H.N. 17.191 (according to information
received from Dr. J. Guibe this specimen was collected in the neighbourhood of
Ndelle, 8 25' N : 2036'E).

DIAGNOSIS. A small sized Agama (body length of males up to 47 mm., of females
to 57 mm.) lacking tufts of erect spiny scales behind the ear and bearing only one
receptor on each body scale, related to Agama weidholzi Wettstein but distinguished
by heterogeneous body scales, strongly keeled head scales and a low fourth toe
lamellar count (13-14).

DESCRIPTION. Body slightly depressed, 3-3-6 times the head length. Nostril
below the canthal edge, situated in the posterior half of a convex, often keeled, oval
or pear-shaped nasal; directed laterally. Interorbital scales as large or larger than
the supraorbitals. Above the nasal a continuous series of three clearly defined,
somewhat elongated scales runs from the rostral to the canthal-supraciliaries. The
first canthal-supraciliary is separated from the nasal by a small postnasal. Scales of
the head strongly keeled, somewhat rugose ; imbrication of temporal scales directed
downwards and slightly backwards. Occipital large, its greatest width equivalent
to the diameter of the tympanum. Nine to twelve, usually ten upper labials;
eight to eleven, usually ten lower labials. No tufts of spines around the ear, instead
single, short, conical scales close to the border of the ear and three to four tubercles
on the side of the neck in a slightly curved row extending from the upper edge of
the ear to above a point midway between the neck pit and the arm insertion ; addi-
tional tubercles irregularly scattered below this row and between the ear and the
neck pit, the lowest at the jaw angle. No gular fold. No trace of a nuchal or dorsal
crest. Dorsal scales heterogeneous, those in the vertebral region a little larger
than the lateral series; on the flanks, and irregularly disposed, some large scales
which are as big and occasionally bigger than the vertebrals.

The scales of the midtrunk region broadly oval or pear-shaped with pronounced
" shoulders ", dorso ventrally flattened on each side of a high keel, the edges of which
rise abruptly from the scale. The keel extends half to two-thirds the length of the
scale but does not project beyond the scale apex to form a mucrone. A single sensory
organ is present on the dorsodistal surface of each of the midbody scales just under the
keel tip or very slightly to one side and sometimes evident only when the scale is
viewed from behind. Generally each of these organs bears one but never more
than one " hair ", which projects slightly beyond the apex of the scale and by macro-
scopic inspection may be mistaken for a mucrone. Gular and ventral scales strongly
keeled. Midbody scale count 70-85 (M 74-8 : N 16) vertebral count 30-46 (M 37 :
N 15). Mature males with eight to twelve preanal pores in a single transverse row.
The adpressed hind limb reaches to the tympanum. Thirteen to fourteen lamellae
under the fourth toe. The first toe is usually either one lamella longer than the
fifth toe or equal to it; rarely (two examples) is it shorter (half lamella). The third
toe exceeds the fourth toe in length by half to two lamella. Tail one and a half
times the body length, depressed at the base, slender and cylindrical distally its
scales strongly keeled and slightly mucronate, and not arranged in whorls.

Body length of gravid females 49-0-56-7 mm., of mature males 44-0-47-0 mm.

NIGERIAN AGAMA LIZARDS
COMPARISON OF CHARACTERS OF NIGERIAN SPECIES OF Agama

agama*

paragama* benueensis*

sankaranica

gracilimembris

Dorsal body Homogeneous
scales

Homogeneous Homogeneous

Homogeneous

Heterogeneous

Position of On canthus

On canthus Below canthus

Below canthus

Below canthus

nostril

Dorsal body +
scales keeled

+ (strongly) +

+ (strongly)

Keeled but not
mucronate

and mucronate

Keeled
ventrals

+ (juveniles
only)

+

Midbody 59~77
count (M68 2 ? :
M66 2 <J)

50-66 74-98
(M 5 8- 5 ) (M 9 i- 3 W.R.:

M85 8 Zaria)

64-78
(M6g 6)

70-85
(M 74 -8)

Vertebral count 38-55 (M^6)

26-34 (^30) 42-56 (M5o)

32-46 (M 39 )

30-46 (M37)

Nuchal crest +

+ +

+

Imbrication of \
scales in \
temporal region
on left side of

\ /rarely \
\ \ \\ /

\

\

\

head

Number of 1-8
receptors per

3-9 1-2 (rarely

3)

1-2 (rarely
3)

I

dorsal scale

Keeling of Faint
dorsal head

Faint Faint

Keeled

Strongly
keeled

scales

Fourth toe

lamellae 18-25

17-21 18-23

15-21

I3-I4

First to fifth toe < (3-6)
(difference
expressed in
number of

<(3-5) <(34)

Subequal

<ft), >(i)

lamellae)

Third to fourth

toe = or<(i-3)

<(i-2) = or <(i-i)

or<(i) or

>(I~2)

Body length 84-91 mm.
of gravid .

96-107 mm. 65-74 mm.

68-85 rnm.

49-57 mm.

Body length 107-137 mm.
of adult c?

99-108 mm. 80-113 mm.

62-76 mm.

44-47 mm.

Tufts of spines +
behind ear and

+ +

+

on side of

neck

* Based on Nigerian individuals only.

88 A. G. C. GRANDISON

COLOUR IN ALCOHOL. <$ Upper surfaces of body and limbs greyish brown ; usually
a clearly defined light vertebral stripe which extends from behind the occiput to the
base of the tail. Straddling the vertebral stripe from nape to base of tail are nine
< > shaped, dark chocolate brown marks which in some specimens are rather
obscure. lu the same region and alternating with these marks, diamond-shaped
brick orange zones occasionally present. The lips, snout, supraorbital and
tympanic and parietal regions invariably dark brown or blue black but a lighter
area over the occipital is always present. Dark, irregular, longitudinal lines are
usually present on the throat, chest and belly but they may extend no farther back-
wards than the gular region. In a sexually mature male these lines are dark brown
and closely set to give a brownish appearance to the entire belly; the throat has a
dark median area.

9- The entire dorsal surfaces of the body, tail and limbs brown with obscure, irreg-
ularly placed lighter areas. An ill defined light vertebral stripe occasionally present.
Snout, supra-orbital area, cheeks and olbique stripe from below eye to commissure
of lips dark brown; occipital, temporal and nuchal regions light fawn. Above and
behind each ear a large ill defined blackish patch in which the small pointed scales
stand out clearly as light bluish or greyish spots. Lower surfaces greyish white with
faint longitudinal dark lines from throat to belly. In some specimens these lines
do not extend farther than the chest.

ECOLOGY. A. gracilimembris is the rarest of the West African savannah species
and has been recorded from only a few localities all of which are in the Doka and
True Guinea woodland vegetation belts and on the extreme southern limit of Sudan
woodland (at Kano). Its biotope has not been described.

RANGE. West Africa : Dahomey to Ubangi Chari.

ACKNOWLEDGEMENTS

The stimulating enthusiasm, interest and collecting ability of R. B. Walker and
Dr. C. H. Fry of the Ahmadu Bello University is gratefully acknowledged and thanks
are extended to Prof. A. P. Mead for his cooperation. Thanks are also due to Dr.
G. T. Dunger who co-operated in obtaining samples from the Jos area. I am
especially indebted to Dr. A. Schi0tz, Danmarks Akvarium for graciously making
available his field notes on his extensive collections and for valuable discussions and
hospitality during my Copenhagen visit. Dr. F. W. Braestrup, Copenhagen Museum
has not only taken a keen interest in this study but has been a source of encourage-
ment and assistance. To Professor J. Guibe, Paris Museum, Dr. V. Aellen, Geneva
and Dr. R. Matthey, La Chaux de Fonds I am also indebted for the loan of speci-
mens, for supplying information on specimens of Agama in their care or for museum
facilities. I wish to thank too A. F. Stimson, Paula D. Jenkins and Belinda A. Brindley
for their help in scale counting and in preparing slides, skeletons and line drawings.

Summary oftaxonomic changes :

Agama boensis Monard part = A . sankaranica Chabanaud

Agama boensis Monard part (juvenile) = A. weidholzi Wettstein
Agama a. benueensis Monard part = A . benueensis Monard

Agama a. benueensis Monard part = A . paragama sp. n.

NIGERIAN AGAMA LIZARDS 89

REFERENCES

BARBOUR, T. & LOVERIDGE, A. 1928. A comparative study of the herpetological faunae of

the Uluguru and Usambara Mountains, Tanganyika Territory, with descriptions of new

species. Mem. Mus. Comp. Zool. Harv. 50 (2) : 87-265, 4 pis.
CHABANAUD, P. 1917. Enumeration des reptiles non encore etudies de 1'Afrique occidentale,

appartenant aux Collections du Museum, avec la description des especes nouvelles. Bull.

Mus. Hist. nat. Paris, 23 : 83-105, 13 text figs.
1918. Etude complementaire de deux Agama de 1'Afrique occidentale et description de

quatre especes nouvelles de reptiles de la meme region. Bull. Mus. Hist. nat. Paris, 24 :

104-112.
CHAPMAN, B. M. & CHAPMAN, R. F. 1964. Observations on the biology of the lizard Agama

agama in Ghana. /. Zool. Lond. 143 : 121-132, 6 figs.
COHN, L. 1914. Die Hautsinnesorgane von Agama colonorum. Zool. Anz. 44 : 145-155,

7 text-figs.
DANIEL, P. M. 1961. Notes on the life history of Agama agama africana (Hallowell) in Liberia.

Spec. Publs. Ohio herpet. Soc. no. 3 : 1-5, 2 tables.
GRANDISON, A. G. C. 1956. On a collection of lizards from West Africa. Bull. Inst. fr. Afr.

noire (A) 18 : 224-245, 4 text-figs, 2 maps.
1969. Agama weidholzi (Sauria: Agamidae) of West Africa and its relationship to Agama

gracilimembris. Ibid, (in press).
HARRIS, V. A. 1963. The anatomy of the Rainbow Lizard. Hutchinson Tropical Monographs,

London: 1-104, 39 text-figs.
1964. The life of the Rainbow Lizard. Hutchinson Tropical Monographs, London:

1-174, 39 text-figs.
KEAY, R. W. J. 1959. Vegetation map of Africa south of the Tropic of Cancer. Oxford

University Press, London.
KING, W. 1962. Systematics of lesser Antillean lizards of the genus Sphaerodactylus. Bull.

Fla. St. Mus. biol. Sci. 7 : 1-52, 17 text-figs.

LINNAEUS, C. 1758. Systema Naturae. Tenth Edition: 824 pp.
LOVERIDGE, A. 1936. African reptiles and amphibians in Field Museum of Natural History.

Publs. Field Mus. nat. Hist. Zool. ser. 22 : i-m.
1941. Report on the Smithsonian-Firestone Expedition's collection of reptiles and

amphibians from Liberia. Proc. U.S. natn. Mus. 91 : 113-140, i text-fig.
1952. Mission A. Villiers au Togo et au Dahomey (1950) XII. Tortoises and lizards.

Bull. Inst. fr. Afr. noire (A) 14 : 229-242.
MILLER, M. R. & KASAHARA, M. 1967. Studies on the cutaneous innervation of lizards.

Proc. Cal. A cad. Sci. 34, 16 : 549-568, 20 text-figs.
MONARD, A. 1940. R6sultats de la Mission scientifique du Dr. Monard en Guinee Portugaise

( I 937- I 93^)- VIII Reptiles. Archos. Mus. Bocage, 11 : 147-180, 2 pis., 5 text-figs.
1951. Resultats de la mission zoologique suisse au Cameroun. Mem. Inst.fr. Afr. noire,

1 : 1-244, ! 3 figs.
PREISS, F. 1922. Uber Sinnesorgane in der Haut einiger Agamiden. Jena Z. Naturw.

58 : 25-76, 9 text-figs.
ROCHEBRUNE, A-T, DE, 1884. Faune de la Senegambie. Reptiles. Octave Doin, Paris,

89 : 1-22 1, 20 pis.
SCHIOTZ, A. 1963. The Amphibians of Nigeria. Vidensk. Medd. dansk. naturh. Foren.

125 : 1-92, 28 text-figs., 4 pis.
SCHMIDT, K. P. 1919. Contributions to the herpetology of the Belgian Congo based on the

collection of the American Congo Expedition, 1909-1915. Bull. Am. Mus. nat. Hist.

39 : 385-624, 27 figs., 32 pis.
SCHMIDT, W. J. 1920. Einiges uber die Hautsinnesorgane der Agamiden insbesondere von

Calotes, nebst Bemerkungen uber diese Organe bie Geckoniden und Iguaniden. Anat. Anz.

53 : 113-139, 1 6 text-figs.

go A. G. C. GRANDISON

SCORTECCI, G. 1937. G"li organ! di senso della pelle degli Agamidi. Memorie Soc. ital. Sci.
nat. 10 (u) : 159-206, 39 text-figs., 2 pis.

1941. I recettori degli agamidi. Memorie Soc. ital. Sci. nat. 10 (m) : 209-326, 80 text-
figs.

STEJNEGER, L. 1893. On some collections of reptiles and batrachians from East Africa and
the adjacent islands, recently received from Dr. W. L. Abbott and Mr. William Astor
Chanler, with descriptions of new species. Proc. U.S. natn. Mus. 16 : 711-741.

THYS VAN DEN AUDENAERDE, D. F. E. 1963. Les Agamidae du Congo: les especes et leur
distribution geographique. Rev. Zool. Bot. afr. 68 : 203-250, 4 text-figs.

UNDERWOOD, G. 1957. O n lizards of the family Pygopodidae. A contribution to the morph-
ology and phylogeny of the Squamata. /. Morph. 100 (2) : 207-268, n text-figs.

WERMUTH, H. 1967. Liste der rezenten Amphibien und Reptilien: Agamidae. Tierreich,
86 : 1-127.

WETTSTEIN, O. 1932. Eine neue Eidechse aus Senegambien. Zool. Anz. 99 : 303-305, i
text-fig.

PLATE I

a. Agama paragama sp. n. Holotype B.M. 1967.2215.

b. Agama paragama sp. n. Lateral view of the head of the holotype.

c. Agama agama L. Lateral view of the head of a juvenile female B.M. 1967 .2212.

d. Agama agama L. Dorsal view of same individual.

Bull. BY. Mus. nat. Hist. (Zool.) 17, 3

PLATE i

ZOOL. 17, 3.

PLATE 2

a. Agama benueensis Monard. Dorsal view of an immature male B.M. 1967.2239.

b. Agama benueensis Monard. Dorsal view of an adult male B.M. 1967.2234.

c. Agama benueensis Monard. Throat of a sexually mature male B.M. 1967.2253.

d. Agama benueensis Monard. Laterial view of the head of an immature male B.M. 1967 . 2239.
Note developing dark gular patch.

Bull. Br. Mus. nat. Hist. (Zool.) 17, 3

PLATE 2

ZOOL. 17, 3.

5

PLATE 3

a. Agama sankaranica Chabanaud. Dorsal view of an adult female B.M. 1967.2204.

b. Agama sankaranica Chabanaud. Lateral view of the head of the same individual.

c. Agama gracilimembris Chabanaud. Lateral view of the head of an adult male B.M. 1961 . 949.

d. Agama gracilimembris Chabanaud. Dorsal view of same individual.

Bull. Br. Mus. nat. Hist. (Zool.) 17, 3

PLATE 3

PLATE 4

Stereoscan electron microscope photographs of dorsal midbody scales. Arrows point to the
sense organs.

Upper row left to right. Agama agama B.M. 1962.1595 x 48

Agama par agama sp. n. Holotype X 28

Lower row left to right. Agama benueensis B.M. 1967.49 x 52

Agama sankaranica B.M. 1967.2204 x 47

Bull. Br. Mus. nat. Hist. (Zool.) 17, 3

PLATE 4

PLATE 5

Stereoscan electron microscope photographs of the apices of the same trunk scales as illustrated
in Plate 4 showing the positions of the sense organs relative to the keel and dorsodistal margin
and the single " hair " protruding from each receptor.

Upper row left to right. Agama agama X 70

Agama paragama X 70

Lower row left to right. Agama benueensis X 131

Agama sankaranica X 260

Bull. BY. Mus. nat. Hist. (Zool.) 17, 3

PLATE 5

PLATE 6

Stereoscan electron microscope photographs of a dorsal midbody scale of Agama gracili-
membris B.M. 1961 .949 showing the position of the single " hair-bearing " sense organ directly
below the termination of the keel.

Left X no. Right X 580.

Bull. BY. Mus. nat. Hist. (Zool.) 17, 3

PLATE 6

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

A REVISION OF THE AMPHIPO:
GENUS MICRODEUTOPUS COSTA
(GAMMARIDEA : AORIDAE)

A. A. MYERS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 17 No. 4

LONDON: 1969

A REVISION OF THE AMPHIPOD GENUS
MICRODEUTOPUS COSTA
(GAMMARIDEA : AORIDAE)

BY

A. A. MYERS

Department of Zoology
University College of Swansea

Pp. 91-148; i Plate, 22 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 17 No. 4

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

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

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 17, No. 4 of the Zoological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 4 February, 1969 Price i 55.

A REVISION OF THE AMPHIPOD GENUS
MICRODEUTOPUS COSTA
(GAMMARIDEA : AORIDAE)

A. A. MYERS

SYNOPSIS

The systematic position of the Aoridae and the phylogenetic relationships of its members are
discussed. All the known species of Microdeutopus and one new species are described and figured,
with keys for their identification. The synonymy of each species of Microdeutopus is reviewed,
and details of their ecology and distribution are given. A catalogue of material studied in the
present work, and the location of type material, is given in the appendices.

INTRODUCTION

THE need for a revision of a number of genera in the Aoridae has long been indicated.
Enequist (1949) pointed out that there is uncertainty about the delimitation of the
species of the genera Microdeutopus and Lembos. Andersson (1954) elucidated the
relationships of Microdeutopus gryllotalpa Costa, M. anomalus (Rathke) and
M. propinquus Sars, using material collected in the Gullmarfjord and collections in
the Swedish State Museum. His findings have been substantiated in the present
work, which included studies on laboratory-reared material. Generally, however,
field workers have experienced difficulties in the identification of aorid material, as is
clear from the numbers of wrongly identified specimens that exist in many institu-
tions. The literature is so widely scattered, and the synonymy, particularly of the
European species, so confused, that a thorough revision of the synonymy and review
of the literature are needed.

During the present investigations, fresh material was examined wherever possible
together with preserved material from the British Museum and a number of other
institutions and specimens from private donors (see Appendix i). Laboratory reared
material of M. gryllotalpa and M. anomalus was also studied.

Identification of Aoridae is frequently hindered by their fragility, specimens collec-
ted in dredge samples often being devoid of all appendages except gnathopoda,
pleopoda and uropoda. Fortunately male Microdeutopus species can readily be
identified by the structure of the gnathopoda, uropoda and telson, and by the form
of the processes which arise from the peraeon segments in the mid- ventral line.
These ventral processes have been shown to occur in all species of Microdeutopus
which have been examined for this character. The only species not examined is
M. haswelli Stebbing, which at this stage seems better not dissected since it is repre-
sented only by the single holotype specimen obtained on loan from the Australian
Museum, Sydney.

In general, the structure of the mouthparts is not a valuable taxonomic criterion,
except at a generic level and above, though in some species, certain minor differences

ZOOL. 17, 4 6

94 A. A. MYERS

can be recognized as in the structure of the mandibular processes of the paragnath
in M. schmitti Shoemaker and M . hancocki Myers.

Kinne (ig63a, 1963^ 1964) has shown that temperature, salinity, food, and com-
binations of these factors affect maximum size in some organisms. Personal
observations in the laboratory (Myers, J_968d) indicate that some, if not all these
factors, also affect growth rates and maximum size in M. gryllotalpa and presumably,
therefore, in other Microdeutopus species. Field observations also suggest that the
various species reach differing maximum sizes, in different localities, and it is therefore
not possible to give an overall indication of the size at which the male of any
Microdeutopus species reaches maturity and is hence identifiable. In particular it
should be noted that high latitude specimens are generally much larger than low
latitude specimens of the same species. In the systematic section the maximum
recorded size, measured from the anterior margin of the cephalosome to the tip of
the telson, is given for each species and each sex.

Male Microdeutopus are readily identifiable, and descriptions of all known species
are given below, together with keys for their identification. The identification of
females presents considerable difficulty, however, since aorid genera are based almost
entirely upon adult male characters, of forms which show extensive sexual dimor-
phism. There are no characters exhibited by females which are diagnostic of the
genus Microdeutopus. Many resemble the male in the structure of the antennules,
antennae, uropoda and telson, thus exhibiting features which differ in one way or
another from those of other Microdeutopus females, and probably from those of other
female Aoridae. However, until the females of all the known Aoridae are studied,
identification of solitary females within this family must remain questionable. Full
descriptions and figures of ovigerous females, are given here, to aid provisional
identification and also to prevent the confusion of females in samples containing
more than one Microdeutopus species.

SYSTEMATIC POSITION OF THE AORIDAE

The Aoridae are closely related to the Photidae, from which they differ in having
the first pair of gnathopoda more markedly developed than the second. Aorcho
Barnard is intermediate in having the first and second gnathopoda of equal size
and can be assigned to either family (Barnard, 1964). Also intimately related to
the Aoridae are the Corophiidae, which differ from the Aoridae only in the degree of
depression of the pleon. The loss of the accessory flagellum of the antennule has
been described as an advanced character in Amphipoda (Barnard, 1962), and it is
suggested here that the corophiid tendency towards the loss of a uropod 3 ramus is
also advanced, so does not necessarily indicate close relationship between the posses-
sors of this character. Barnard (1962) has pointed out that it has become increas-
ingly difficult to assign, between the two families, various members of the Photidae
and Corophiidae, because here too the differences involve quantitative aspects of
depression or compression of the pleon. It seems probable that the family Coro-
phiidae is polyphyletic and includes, within its framework, advanced members of
both the Aoridae and Photidae. It is therefore not possible to indicate any circum-

REVISION OF MICRODEUTOPUS 95

scribed diagnosis for the family Aoridae, which can only be described as those mem-
bers of the aorid/photid/ischyrocerid/corophiid complex which exhibit greater
development of gnathopod I than gnathopod 2, coupled with a tendency towards
retention of such primitive characters as an accessory flagellum on the antennule,
uropod 3 with two rami, contiguous coxae, and no marked depression of the pleon.

The subdivision of the family Aoridae is also problematical since the complex
permutations of characters among the species makes a systematic classification,
based on possible phylogenetic relationships, exceedingly difficult. The accessory
flagellum of the antennule exhibits a complete gradation of stages, from (a) primitively
multi-articulate as in Aora typica Kroyer, Microdeutopus anomalus (Rathke),
Lembopsis spinicarpus Pearse, Lembos viguieri Chevreux and others, to (b) reduced
to two or three articles of which the terminal one is usually vestigial, e.g. in Micro-
deutopus armatus Chevreux, Lemboides afer Stebbing, and Neomegamphopus roose-
velti Shoemaker, or microscopic as in Aoroides columbiae Walker. The rami of uropod
3 may be (a) more or less equally developed as in Amphideutopus oculatus Barnard,
Microdeutopus versiculatus (Bate), Lembos longipes Lilljeborg, and Aorcho delgadus
Barnard; (b) the endopodite may be greatly reduced as in Acuminodeutopus heter-
uropus Barnard ; (c) both rami may be minute, as in Paradryope orguion Stebbing and
Dryopoides westwoodi Stebbing; or (d) there may be a single ramus only as in
Neomicrodeutopus cabindae Schellenberg. The antennae exhibit considerable
variation in form. They may be (a) shorter than the antennules stout, sub-pediform,
with few flagellar articles, resembling those of the Corophiidae, as in Microdeutopus
haswelli Stebbing; (b) less stout, as in M. gryllotalpa Costa, Aora typica Kroyer,
Lembos websteri Bate and Lemboides afer Stebbing; (c) slender though still shorter
than the antennules, as in Microdeutopus stationis Delia Valle, Lembopsis spinicarpus
Pearse and Lembos macromanus (Shoemaker) ; or (d) very long and slender, exceeoling
the length of the antennules, with multiarticulate flagella, as in Paradryope orguion
Stebbing and Amphideutopus dolichocephalus Myers. The development of median
spines on the ventral surface of the peraeon segments is of frequent occurrence in
the Aoridae, but such spines are not restricted to the Aoridae, being also exhibited
by various members of the related family Corophiidae, including certain species of
Unciola Say and Grandidierella Coutiere.

Most classifications of the Aoridae have been based upon the structure of the male
gnathopod i, the gross development of which is so characteristic a feature of the
Aoridae, though it is not diagnostic. Two main tendencies are discernible in this
appendage: first, the great development of one or more of the articles, frequently
accompanied by reduction of some of the others, and secondly, the production of
toothlike processes on one or more of the articles. Classifications based on either
of these characters are arbitrary, since it would be unreasonable to assume that either
of these tendencies has occurred only once in the evolution of the family. Never-
theless, a good workable classification is that of Barnard (1958, 1962, 1964) which is
adopted here with two important modifications suggested by present work. Firstly
Microdeutopus tridens Schellenberg and M. kraemmeri Reid are transferred to the
genus Lembopsis Pearse (see Myers I968c); and secondly, the genus Coremapus
Norman is incorporated in the genus Microdeutopus (see p. 102).

96 A. A. MYERS

Genus MICRODEUTOPUS Costa 1853

Microdeutopus Costa, 1853 : 178.
Autonoe Bruzelius, 1859 (pro-parte) : 23.
Stimpsonia Bate, 1862 : 162.
Microdeuteropus Norman, 1868 : 281.
Stimpsonella Delia Valle, 1893 : 421.
Coremapus Norman, 1905 : 78.

TYPE SPECIES : Microdeutopus gryllotalpa Costa.

DIAGNOSIS. Head lobes moderately produced, obtuse; antennules longer than
the antennae (with the exception of male M. chelifer (Bate)) ; article i of mandibular
palp the shortest, article 3 the longest, article 3 broad centrally, narrowing terminally;
mandibular process of paragnath acute ; gnathopod i of male larger than gnathopod
2, complexly chelate, article 5 considerably larger than article 6 ; females with the
first and second gnathopods dissimilar; uropod 3 with two more or less subequal
rami; telson simple.

HISTORICAL. The genus was established by Costa (1853) to include the single
species M. gryllotalpa Costa, independently described by Bruzelius under the name
Autonoe grandimana. Bate (1856) erected the genus Lembos to include the species
L. cambriensis Bate, L. damnoniensis Bate, L. versiculatus Bate and L. websterii
Bate, but later (1862) relegated all these species to the genus Microdeutopus Costa
(corrected from Microdentopus in the text, to Microdeutopus in the appendix).
In the same year he erected the genus Stimpsonia, to include as monotype chelifera
Bate, while admitting that " the genus probably bears too close a resemblance to
Microdeutopus to be retained as generically distinct ". Delia Valle (1893) noted the
preoccupation of Stimpsonia among the Nemertina (Stimpsonia Girard, 1853)
and erected Stimpsonella, including in the genus, armatus Chevreux with chelifera
Bate. Norman (1868) altered Microdeutopus to Microdeuteropus for grammatical
reasons but later (Norman, igo5a) reverted to the original spelling. Norman
(i905b) erected the monotypic genus Coremapus for versiculatus Bate.

LITERATURE. The best general account is that of Delia Valle (1893) who gives
ten species. Of these, however, M. gryllotalpa Costa and M. minax Smith are
synonymous, M. tennis Dana is not a species of Microdeutopus and M. titii Heller
is a dubious species, while the distinct species M . versiculatus is treated as a synonym
of M. anomalus Rathke. The keys, which apply to males only, are adequate.
Stebbing (1906) gives a good account of eight species, and not eleven as it would
seem, for M. anomalus (Rathke) and M. propinquus Sars are synonymous,
M. megnae Giles is attributable to Grandidierella Coutiere, and the doubtful M. titii
Heller is listed with reservation. Descriptions of the females are inadequate, as
in most literature, but the keys which apply to males only are adequate. Chevreux
and Fage (1925) give seven species and their keys are modified after Stebbing (1906).
Barnard (1958) gives a catalogue of species, and other useful taxonomic works are
those of Haswell (1882) describing i sp. from Australia, Sars (1894) 2 spp. from
Norway, Stephenson (i92ga) 2 spp. North Sea and Baltic, Shoemaker (1938) i sp.
Mexico, Schellenberg (1942) 2 spp. Germany, Gurjanova (1951) 3 spp. U.S.S.R. and
Myers (i968b) 3 spp. Central America.

REVISION OF MICRODEUTOPUS 97

KEY TO THE MALE MICRODEUTOPUS OF THE WORLD

N.B. in most figures of limbs, article i is omitted so article 2 is the most proximal shown.

1 a. Gnathopod 2 chelate ........... 2

b. Gnathopod 2 not chelate .......... 3

2 a. Gnathopod 2 with articles 5 and 6 subequal in length and breadth, article 2

expanded, crenulated on anterior margin (Text-fig. i8d) . M. armatus (p. 129)

b. Gnathopod 2 with article 6 much shorter and broader than article 5, article 2

unexpanded, anterior margin smooth, (Text-fig. IQC) . . M. chelifer (p. 132)

3 a. Gnathopod i with the anterior margin of article 5 produced into a single tooth.

Accessory tooth if present, arising within the posterior margin (Text-fig. 8b) . 4

b. Gnathopod i with article 5*8 anterior margin produced into 2 or more teeth (Text-
fig. 2a) . 7

4 a. Gnathopod 2 densely setose, the setae finely pectinate; article 4 extending over

greater length of article 5 (Text-fig. 31) . . . M. versiculatus (p. 101)

b. Gnathopod 2 moderately setose, the setae not pectinate; article 4 relatively short

(Text-fig, gc) (" anomalus " group) 5

5 a. Gnathopod 2 with articles 5 and 6 short and broad, palmar angle of article 6

almost transverse (Text-fig, ga) ; uropod 3 with rami relatively short and broad

M. damnoniensis (p. 114)

b. Gnathopod 2 with articles 5 and 6 slender, elongate, palmar angle of article 6
oblique (Text-fig, gc) ; uropod 3 with rami relatively elongate and slender
(Text-fig, ge) 6

6 a. Gnathopod i with the anterior distal margin of article 5 produced into a slender,

straight or inwardly curved tooth, generally with an accessory tooth arising
near its base within the posterior margin (Text-fig. 8b) . M. anomalus (p. no)

b. Gnathopod i with article 5*3 anterior margin produced into a broad based stout

tooth, never with accessory tooth (Text-fig. 8h) . . . M. algicola (p. 117)

7 a. Gnathopod i with article 5 having the proximal (outer) of two teeth longer than

the other (Text-figs. 2e, 22a) ......... 8

b. Gnathopod i 's article 5 not as above ........ 9

8 a. Antenna stout, subpediform, with the first flagellar article grossly developed

(Text-fig. 4b); gnathopod 2 with article 2 elongate and slender (Text-fig. 3h)

M. haswelli (p. 107)
b. Antenna relatively slender, the first flagellar article not as above (Text-fig. 21),

gnathopod 2 with article 2 broad and short (Text-fig. 22g) . M. sporadhi (p. 136).

9 a. Gnathopod i with the distal tooth on article 5 the longest (Text-fig. 2b) . . 10
b. Gnathopod i with the central tooth on article 5 the longest (Text-fig. 2d)

M. stationis (p. 104)
10 a. Gnathopod 2 with article 2 convex anteriorly, crenated on the anterior margin

(Text-fig. 3g) M. gryllotalpa (p. 98)

b. Gnathopod 2 with article 2 concave anteriorly and its anterior margin smooth

(Text-fig. 14!) (" schmitti " group) n

n a. Gnathopod i having article 5 with a tooth on the anterior margin (Text-fig. I3a) 12
b. Gnathopod i having article 5 without a tooth on the anterior margin (Text-fig. I3b)

M. schmitti (p. 120)

12 a. Gnathopod i having article 6 with the palmar margin produced into a forward
directed tooth opposable to article 7; posterior margins of articles 4, 5 and 6 with
very long setae (Text-fig. 136) ..... M. trichopus (p. 124)

b. Gnathopod i having article 6 with the palmar margin at most produced into a
small lobe, which is never directed forward; posterior margin of article 4 only
having very long setae, those of articles 5 and 6 being relatively short (Text-fig.
i3a) .......... M. hancocki (p. 124)

98 A. A. MYERS

Microdeutopus gryllotalpa Costa
(Text-figs. la, 2a-c, 3a-c, g, m, 5b, 6c-d, 2oa)

Microdeutopus gryllotalpa Costa, 1853 : 178; Costa, 1857 : 231, pi. 4, fig. 10; Boeck, 1870 : 156;
Boeck, 1876 : 565, pi. 29, fig. 6; Blanc, 1884 : 75, pi. 4, figs. 82-90; Hoek, 1889 : 226; Delia
Valle 1893 : 411, pi. i, fig. 12, pi. n, figs. 25-43; Sars 1894 : 543. P^ I 9 2 > n g- 2 > Sowinsky
!895 : 237, pi. 4, fig. 6; Sowinsky, 1898 : 480; Stebbing, 1906 : 590; Norman, 1907 : 368,
pi. 16, fig. 3, pi. 17, figs. 6-7; Chevreux & Fage, 1925 : 299, fig. 310; Stephenson, 1927: 124;
Stephensen. igzqa : 151, fig. 273; Oldevig, 1933 : 213, fig. 95; Cecchini & Parenzan, 1934 :
213, fig. 43; Loven, 1934: 68, fig. 2; Miloslavskaia, I939a : 121; Schellenberg, 1942 : 186,
fig. 154; Soika, 1949 : 199; Gurjanova, 1951 : 831, fig. 580; Barnard, 1958 : 29.

[Nori] Microdeutopus gryllotalpa Bate, 1862 : 163, pi. 30, fig. i.

[Nori] Microdeutopus gryllotalpa Nebeski, 1880 : 45, fig. 41.

[Nori] Microdeutopus grillotalpa [sic] Sowinsky, 1880 : 125, pi. 5, fig. 17 a-d.

Autonoe grandimana Bruzelius, 1859 : 26, pi. i, fig. 5.

Microdeutopus grandimanus (Bruzelius) Bate, 1862 : 378.

Microdeutopus minax Smith, 1874 : 562.

Microdeutopus bidens Sowinsky, 1880 : 129, pi. 5, fig. 18.

[?]Amphithoe salenskiiCarus, 1885 : 396.

Microdeuteropus gryllotalpa (Costa) Norman, 1886 : 16; Chevreux & Bouvier, 1893 : 133.

Microdeuteropus minax (Smith) Norman, 1886 : 17.

TYPE LOCALITY. Lago di Fusaro, Italy.

DIAGNOSIS OF MALE. Antennule about half the body length, the first peduncular
article slightly shorter than the second, the third about one third the length of the
second ; flagellum longer than the peduncle normally with up to twenty- two articles
(maximum recorded twenty-six) ; accessory flagellum with two articles, the second
article rudimentary. Antenna slightly under two thirds the length of the antennule,
the fourth and fifth peduncular articles subequal; flagellum shorter than the fifth
peduncular article, normally with up to nine articles (maximum recorded eleven).
Peraeon segments 2-7 each with a process in the mid-ventral line, those of segments
2 and 3 short, straight, those of segments 4-7 short, robust, posteriorly directed;
processes 6 and 7 with short setae arising from their posterior margins. Gnathopod

1 with article i moderately produced anteriorly, acute ; article 2 short and expanded
distally; article 5 immensely expanded, in some specimens as broad as long, the
posterior distal margin produced into two to four teeth, of which the most distal
is always the longest ; article 6 short, the posterior margin irregularly lobed ; article
7 sometimes short and robust, sometimes relatively long and slender. Gnathopod

2 with article 2 greatly expanded, the anterior margin convex and crenated; articles
5 and 6 subequal in length, 5 slightly the broader, articles 4, 5 and 6 with long pectin-
ate setae on the anterior margins. Uropod 3 with rami equal in length to the peduncle
exopodite slightly the longer of the two rami; each ramus with a transverse row of
three closely associated spines on the outer dorsal margin, together with a number of
paired and solitary spines of variable distribution, terminal setae relatively long.
Telson with each terminal crest bearing on its distal dorsal margin, a group of up to
six setae.

Maximum size of male : 8 mm.

REVISION OF MICRODEUTOPUS

99

FEMALE. As the male except for the absence of ventral peraeon processes and
the structure of the sexually dimorphic gnathopoda.

Gnathopod i with article 2 moderately expanded; articles 5 and 6 subequal in
length, 5 slightly the broader, 6 with the anterior and posterior margins roughly
parallel; article 7 equal in length to the angular palmar region. Gnathopod 2 with

FIG. i. Microdeutopus gryllotalpa Costa, (a) Male head, Swansea, Wales. Microdeutopus
versiculatus (Bate), (b) Male head, Falmouth Harbour, England.

ioo A. A. MYERS

article 2 slightly expanded, crenulated on the anterior margin; articles 5 and 6
elongate, subequal; articles 4, 5 and 6 with pectinate setae on their anterior margins.
Maximum size of female: 10 mm.

DISCUSSION. M. gryllotalpa is closely related to M. versiculatus but differs in
both sexes by the second gnathopod (Text-figs. 3g, 5b) not having the fourth article
greatly produced along the posterior border of the fifth, nor the fifth and sixth

FIG. 2. Microdeutopus gryllotalpa Costa, (a) Male gnathopod i, Swansea, Wales, (b) Male
gnathopod i, Swansea, Wales, (c) Male gnathopod i, Isefjord, Denmark. Microdeuto-
pus stationis Delia Valle. (d) Male gnathopod i, St. Peter's Port, Guernsey, (f) Male
gnathopod i, Naples, Italy. Microdeutopus haswelli Stebbing. (e) Male gnathopod i,
Port Jackson, Australia (Holotype).

REVISION OF MICRODEUTOPUS 101

articles so markedly elongate and slender. Females of M. gryllotalpa are readily
identifiable by the structure of the fifth article of the second gnathopod (Text-fig. 5b) ,
which has long pectinate setae arising from the entire length of the anterior margin,
whereas in all other Microdeutopus species (except M. versiculatus) the long setae
arise from the anterior corner only of the fifth article.

ECOLOGY. In the intertidal zone in rock pools amongst Chaetomorpha and other
algae; amongst Zoster a spp.; and in salt marshes (Chevreux & Page, 1925), often
associated with Ruppia maritima as at Roscoff (Truchot, 1963). Also occurring
sub-littorally to a depth of about 150 metres on oyster beds, and amongst algae,
tunicates, sponges, polyzoans, Mytilus etc., always in areas of high detritus accumu-
lation. It particularly favours docks and other man-made installations, and is
frequently associated with Corophium acherusicum Costa in both the Palearctic
and Nearctic regions.

DISTRIBUTION. From the Lofoten Islands southwards along the coasts of Norway,
Sweden, Denmark, including the Baltic, and Holland, around the coasts of the British
Isles, chiefly in the south-west, along the Atlantic coasts of France, Spain and
Portugal, and in the Mediterranean Ligurian, Tyrrhenean, Adriatic, Aegean and
Black Seas. Also, on the north-eastern seaboard of the United States of America
(Massachusetts, Rhode Island and Connecticut) .

Microdeutopus versiculatus (Bate)
(Text-figs, ib, 3i, k, 5a, 6a-b, 2oc)

Lembos versiculatus Bate, 1856 : 58; Bate, 1857 : 142.

Microdeutopus versiculatus Bate, 1862 : 165, pi. 30, fig. 5; Bate & Westwood 1863 : 295; Walker,

1895 : 469; Chevreux 1900 : 89; Stebbing, 1906 : 593.
Microdeuteropus versiculatus (Bate) Norman, i86ga : 282; Stebbing, 1874 : 12, pi. i, figs. 2,

2a-f. Chevreux & Bouvier, 1893 : 134.
Autonoe longipes (Lilljeborg) Boeck, 1876 (pro-parte) : 574.
Autonoe versiculata (Bate) Norman, 1886 : 17.

Microdeutopus anomalus (Rathke) Delia Valle, 1893 (pro-parte) : 418.
Coremapus versiculatus (Bate) Norman, igo5b : 78; Chevreux & Page, 1925 : 301, fig. 312;

Stephensen, i92ga : 152, fig. 276; Cecchini & Parenzan, 1934 : 217, fig. 45; Miloslavskaia,

I939a : 125, fig. 25; Soika, 1949 : 200; Gurjanova, 1951 : 835, fig. 585; Barnard, 1958 : 29.

TYPE LOCALITY. Plymouth, Devon, England.

DIAGNOSIS OF MALE. Antennule about equal to the body length, the first pedun-
cular article slightly shorter than the second, the third about one third the length
of the second; flagellum almost twice the length of the peduncle, normally with up
to twenty articles (maximum recorded twenty-two); accessory flagellum with two
articles, the second article rudimentary. Antenna little more than half the length
of the antennule, the fourth peduncular article slightly longer than the fifth ; flagel-
lum shorter than the fifth peduncular article, normally with up to eight articles
(maximum recorded nine). Per aeon segments 2-7 each with a process in the mid-
ventral line; that of segment 2 slender, spiniform, directed anteriorly; those of
segments 3 and 4 slender, spiniform, straight; those of 5-7 robust, decreasing in
size antero-posteriorly, with processes 6 and 7 having setae arising from their posterior

loa A. A. MYERS

margins. Gnathopod i with article I moderately produced anteriorly, acute;
article 2 considerably expanded; article 5 much enlarged, longer than broad, the
posterior distal angle produced into a single, robust tooth; article 6 short, the post-
erior margin irregularly lobed; article 7 short and robust. Gnathopod 2 with article
2 somewhat expanded ; article 4 well developed, extending over the greater length of
the posterior margin of article 5 ; article 5 longer than article 6, elongate and slender,
about three times as long as broad; article 6 also very long and slender, about three
and a half times as long as broad; article 7 very short, long pectinate setae arising
from the anterior margins of articles 4, 5 and 6. Uropod 3 with rami sub-equal, equal
in length to the peduncle ; exopodite with a transverse row of three closely associated
spines on the outer-dorsal margin, basal to which is a solitary spine ; endopodite with a
similar transverse row of closely associated spines on the outer-dorsal margin, and
in addition on the inner margin, a series of well separated spines; terminal setae of
both rami relatively long. Telson with each terminal crest bearing on its distal
dorsal margin a group of two to three setae.
Maximum size of male: 7 mm.

FEMALE. As the male except for the structure of the sexually dimorphic gnatho-
poda and absence of ventral peraeon processes. Gnathopod i with article 2 short
and considerably expanded, about two thirds as broad as long; article 5 scarcely
longer than article 6, with a dense clothing of comb setae along the entire length of
the posterior border; article 6 almost as broad as 5, with the anterior and posterior
margins roughly parallel; article 7 moderately long, equal in length to the palmar
region. Gnathopod 2 with article 2 elongated and slender, otherwise identical with
the corresponding appendage of the male.

Maximum size of female : 8 mm.

DISCUSSION. First designated as a nomen nudum by Bate (1856) the species was
later described from the female (Bate, 1857, 1862). The male was first described
by Norman (1868) and later by Stebbing (1874) who also gave figures. The sugges-
tion that a monotypic genus should be erected for this species (Norman, i905b) is not
accepted. The present species bears a close relationship with M. gryllotalpa Costa
in the similarity of the male first gnathopod of the two species. That of M. versicu-
latus (Text-fig. 3k) differs in having a single tooth on the posterior distal angle of
article 5, whereas in M. gryllotalpa (Text-figs. 2a-c) there are two to four. The
two species are also similar in the structure of the male second gnathopod. Article
2 of that appendage is expanded, though less markedly in M. versiculatus (Text-fig.
3i) than in M. gryllotalpa (Text-fig. 3g) and article 5 has pectinate setae arising from
the entire anterior margin (also a character of the female). Finally, males of both
species have ventral processes on all peraeon segments except the first. The
remarkable slender, setose second gnathopod of M. versiculatus (Text-fig. 5a),
which led Normon (iQ05b) to erect a monotypic genus for the species, is an extreme
modification of the type found in M. gryllotalpa and serves to distinguish M.
versiculatus in both sexes from all other Microdeutopus species.

ECOLOGY. In the intertidal zone (Oldany, Sutherland) , but much more frequently
in the sub-littoral to a depth of about 150 metres (off the Shetland Islands) amongst

REVISION OF MICRODEUTOPUS

103

Fig. 3. Microdeutopus gryllotalpa Costa, (a) Male gnathopod i, 1-8 mm. <$, Swansea,
Wales, (b) Male gnathopod i, 2-9 mm. <$, Swansea, Wales, (c) Male gnathopod i,
3-9 mm. cJ, Swansea, Wales, (g) Male gnathopod 2, Swansea, Wales, (m) Male telson,
Swansea, Wales. Microdeutopus stationis Delia Valle. (d) Male gnathopod i, 1-9 mm.
<J, Naples, Italy, (e) Male gnathopod i, 2-0 mm. <J, Naples, Italy, (f) Male gnathopod
i, 3-5 mm. $, Naples Italy, (j) Male gnathopod 2, Naples, Italy. (1) Male telson,
Naples, Italy. Microdeutopus haswelli Stebbing. (h) Male gnathopod 2, Port Jackson,
Australia (Holotype). Microdeutopus versiculatus (Bate), (i) Male gnathopod 2, Ply-
mouth, Devon, (k) Male gnathopod i, Plymouth, Devon.

104 A. A. MYERS

algae, especially cystoseires and corallines, polyzoans, Mytilus etc. in areas of high
detritus accumulation.

DISTRIBUTION. Along the south and west coasts of the British Isles to the Shet-
lands, in the Channel Islands and the adjoining French coast, Bay of Biscay (Houat,
Arcachon), the Atlantic coasts of Spain and Portugal, Senegal and the Azores.
Also around the coasts of the Mediterranean, Tyrrhenian and Adriatic seas, and in the
Black sea (north-east of Cape Caliacra, Roumania).

Microdeutopus stationis Delia Valle
(Text-figs. 2d, f, 3d-f, j, 1, 4a, 5c, 6e-f, 2oe)

Microdeutopus gryllotalpa Nebeski, 1880 : 45, fig. 41.

Microdeutopus stationis Delia Valle, 1893 : 415, pi. 5, fig. 2, pi. 10, figs. 31-41; Stebbing, 1906 :

590; Norman, 1907 : 368, pi. 16, fig. 4, pi. 17, figs. 8-n ; Chevreux & Fage, 1925 : 300, fig. 311 ;

Cecchini & Parenzan, 1934 : 2I ^> fig- 44>' Miloslavskaia, i939a : 122; Gurjanova, 1951 : 832,

fig. 581 ; Barnard, 1958 : 29.
[Wow] Microdeutopus stationis Sowinsky, 1895 : 237, pi. 4, figs. 1-5; Sowinsky, 1898 : 480.

TYPE OF LOCALITY. Naples, Italy.

DIAGNOSIS OF MALE. Antennule about equal to the body length, the first pedun-
cular article slightly shorter than the second, the third less than one third the length
of the second; flagellum twice the length of the peduncle, normally with up to
twenty-six articles (maximum recorded twenty-nine) ; accessory flagellum with
two to five articles, the terminal article rudimentary. Antenna two thirds the
length of the antennule, the fourth and fifth peduncular articles subequal; flagellum
equal in length to the fifth peduncular article, normally with up to twelve articles
(maximum recorded fourteen). Per aeon segments 2-4 each with a process in
the mid-ventral line, that of segment 2 slender spiniform, straight, those of
segments 3 and 4 becoming progressively shorter and more robust. Gnathopod ~L
with article I strongly produced anteriorly, very attenuated; article 2 scarcely
expanded; article 5 very enlarged, longer than broad, the posterior distal angle
produced into three teeth of which the central is the longest, the inner two slightly
recurved; article 6 short, the posterior margin convex distally, concave basally;
article 7 long and slender. Gnathopod 2 with article 2 not expanded; article 5
longer and slightly broader distally than article 6. Uropod 3 with the rami slightly
shorter than the peduncle, the expedite slightly the longer of the two rami, bearing
on its outer distal margin a group of three closely associated elongate spines, with a
pair of spines (of which one is elongate) basal to these and not infrequently a further
solitary spine basal to the pair; inner margin of exopodite with one or two solitary
spines; endopodite outer margin with a pair of spines, basal to which is a solitary
spine; inner margin with up to two pairs of spines and a solitary spine; terminal
spines and setae of both rami very short. Telson with each terminal crest bearing
on its distal dorsal margin, a group of three short stout spines and a single fine seta.

Maximum size of male : 10 mm.

FEMALE. As the male, except for the structure of the sexually dimorphic gnatho-
poda, and absence of ventral peraeon processes. Gnathopod i with article 2 scarcely

REVISION OF MICRODEUTOPUS

105

FIG. 4. Microdeutopus statlonis Delia Valle. (a) Male head, St. Peter's Port, Guernsey.
Microdeutopus haswelli Stebbing. (b) Male head, Port Jackson, Australia (Holotype).

io6

A. A. MYERS

expanded, articles 5 and 6 subequal in length, the sixth distinctly broader distally,
narrowing at the junction with the fifth article, the palmar angle smoothly rounded;
article 7 elongate and slender. Gnathopod 2 with article 2 relatively slender, article
5 slightly shorter than article 6, setae not pectinate.
Maximum size of female : 12 mm.

DISCUSSION. The species was first described by Nebeski (1880) from the Adriatic,
and erroneously ascribed to M. gryllotalpa Costa. That author's illustrations
(Nebeski, 1880, fig. 41) of the growth stages of the male gnathopod i clearly indicate

FIG. 5. Microdeutopus versiculatus (Bate), (a) Female gnathopod 2, Plymouth, Devon.
Microdeutopus gryllotalpa Costa, (b) Female gnathopod 2, Swansea, Wales. Micro-
deutopos stationis Delia Valle. (c) Female gnathopod 2, Naples, Italy.

that he was dealing with the species later described by Delia Valle under the name
M. stationis.

M. stationis is easily distinguishable in the male from all other described species
in the genus by having the posterior distal angle of article 5 of the first gnathopod
(Text-figs. 2d, f) produced into three teeth, of which the central is the longest.
Females have few diagnostic characters, but important characters taken in combina-
tion are the telson (Text-fig. 31) bearing both spines and setae, uropod 3 (Text-fig. 6f)
with groups of spines on the rami, accessory flagellum with two to four articles
and long slender antenna with multi-articulate flagellum.

ECOLOGY. Recorded in Sacchoriza holdfasts, among Posidonia, on oyster beds
and over fine sands, in regions of relatively low detritus contamination, in the sub-
littoral to a depth of 50 metres.

REVISION OF MICRODEUTOPUS 107

DISTRIBUTION. Guernsey (St. Peter's Port), France (Arcachon, Roscoff, Perros
Guirec), Corsica (Ajaccio, Bonifacio), Algeria (Bone, La Calle), Tunisia (Golfe de
Gabes), Italy (Naples, Trieste), Greece (Khios).

Microdeutopus haswelli Stebbing
(Text-figs. 2e, 3h, 4b)

Microdeuteropus chelifer Hasell, 1879 : 340, pi. 22, fig. 3; Delia Valle, 1893 : 421.
Microdeutopus chelifer Haswell, 1882 : 265.

Microdeutopus haswelli Stebbing, 1899 : 350; Stebbing, 1906 : 591; Stebbing, 1910 : 647;
Sheard, 1937 : 2 ^; Barnard, 1958 : 29.

TYPE LOCALITY. Clark Island, Port Jackson, Australia.

The present species is represented by a single male specimen in Haswell's collec-
tions, at the Australian Museum, Sydney. Haswell did not designate types for
any of his material, but this specimen is almost certainly the holotype, agreeing in
the main with Haswell's description, and is here considered as such. An expanded
description of the more relevant features of the holotype is given. The range of
variation of this species is not known, as further material has not been forthcoming.

DESCRIPTION OF MALE HOLOTYPE. Head lateral lobes slightly produced, obtuse;
eyes round. Antennule slightly less than half the body length, the first and second
peduncular articles subequal, the third slightly over a third the length of the second ;
flagellum longer than the peduncle, with seventeen articles; accessory flagellum
with five articles, the fifth article rudimentary. Antenna shorter than the antennule,
sub-pedif orm, the fifth peduncular article slightly longer than the fourth ; flagellum
with four articles, the first article long and robust, over half the length of the fifth
peduncular article, the second short, the third and fourth articles progressively
shorter, obscurely articulate. Gnathopod I with article I moderately produced
anteriorly, obtuse; article 2 short and expanded, the anterior margin excavate;
article 5 much enlarged, longer than broad, the posterior distal angle produced into
two teeth, of which the outer is the longer and recurved; article 6 short, the posterior
margin concave centrally; article 7 very stout. Gnathopod 2 with article 2 very
elongated and slender, concave anteriorly, articles 5 and 6 slender with article 5
slightly the longer. Epimeral plate 3 with the posterior free corner produced into a
tooth, above which is inserted a short seta. Uropod 3 with the peduncle exceeding
the length of the very short and stout rami, of which the exopodite is very slightly
the longer; each ramus with a terminal group of setae which exceed the length of
the ramus. Telson with the terminal crests moderately well developed, each bearing
on its distal, dorsal margin, a group of three setae, of which the central is the longest.

Length : 4-5 mm.

DISCUSSION. The very robust, sub-pediform antenna, (Text-fig. 4b) with the
flagellum very reduced and the first flagellar article robust, approaches the condition
found in many Corophiidae. However, the lateral compression of the urosome,
biramous third uropoda, and moderately well developed coxae, are characteristic
of aorid/photid stock. M. haswelli differs from all other Microdeutopus species,

ZOOL. 17, 4 7

io8

A. A. MYERS

except M. sporadhi sp. nov. (p. 136), in having the posterior distal angle of article 5
of the male first gnathopod (Text-fig. 2e) produced into two teeth, of which the outer
is the longer. The elongated, slender article 2 of the male second gnathopod (Text-
fig. 3h) is also a distinctive feature.

FIG. 6. Microdeutopus versiculatus (Bate), (a) Female gnathopod i, Plymouth, Devon,
(b) Male uropod 3, Falmouth, Cornwall. Microdeutopus gryllotalpa Costa, (c) Female
gnathopod i, Swansea, Wales, (d) Male uropod 3, Arcachon, France. Microdeutopus
stationis Delia Valle. (e) Female gnathopod i, Naples, Italy, (f) Male uropod 3, Naples,
Italy.

REVISION OF MICRODEUTOPUS 109

ECOLOGY. Unknown.

DISTRIBUTION. Not yet recorded from other than the type locality.

THE " ANOMALUS " GROUP OF SPECIES

The systematics of the anomalus group of species has long been under dispute.
In 1856 Bate erected two species, Lembos damnoniensis and L. cambriensis, but gave
no descriptions and only in the following year attributed to the former species
" first hand with a thumb on carpus " to the latter " first hand without thumb ".
White (1857), no doubt in error, transposed the " mn " in L. damnoniensis, to
" nm " in L. danmoniensis , and later Bate (1862) and Bate and Westwood (1863)
relegated L. cambriensis to a synonym of M. anomalus (Rathke) and L. damnoniensis
to a synonym of M. gryllotalpa Costa. Norman (1868) considered that M. gryllo-
talpa Bate represented a young male M. anomalus, but later (Norman iQOSb) decided
that M. damnoniensis Bate, was a distinct species, and that M. gryllotalpa Bate (non
M. gryllotalpa Costa) represented that species. Sars (1894) figures a species under
the name M. propinquus Sars, but in the text relegates it to a synonym of
M . danmoniensis (Bate) (altered presumably in error, or following White, 1857, from
M. damnoniensis}. St ebbing (1906) considered that M. propinquus Sars was not
synonymous with M. damnoniensis (Bate) and reinstated it as a distinct species with
synonym M. danmoniensis Sars (non M. damnoniensis (Bate)). Andersson (1954)
considered M. propinquus Sars to be a juvenile form of M. anomalus (Rathke) and
in the synonymy also listed L. damnoniensis Bate. Delia Valle (1893) described a
new species M. algicola, and placed M. damnoniensis in the synonymy of M. anomalus
(Rathke).

Present investigations suggest that there are three distinct species in the
"anomalus" group; M. anomalus (Rathke), M. damnoniensis (Bate) and
M. algicola Delia Valle.

Males of the " anomalus " group of species are distinguished from those of all
other Microdeutopus species, except M. versiculatus (Bate) M. chelifer (Bate) and
M. armatus Chevreux by having the posterior margin of gnathopod I produced into a
single marginal tooth. In M. anomalus an accessory tooth is usually present, but
this always arises within the posterior margin. They differ from the above three
species in the structure of gnathopod 2, which is neither chelate as in M. chelifer
and M. armatus not densely setose as in M . versiculatus. Females of the " anomalus "
group of species are notable for their lack of diagnostic features, and are therefore
identifiable only by negative characters. They are distinguishable from
M. versiculatus, M. gryllotalpa and M. stationis by the absence of paired spines from
the rami of the third uropod, from the first two species by the absence of pectinate
setae on gnathopod 2. They differ also from M. stationis and from the " schmitti "
group of species by the lack of any spines as distinct from setae on the telson terminal
crests, and from M. armatus by the lack of any pronounced projection at the antero-
distal corner of article 2 or gnathopod 2. Within the " anomalus " group of species
the identification of females is even more difficult, particularly in the absence of
males. In practice, when two or more of the " anomalus " group occur together

no A. A. MYERS

in a sample, the females have been separated with some degree of accuracy and
grouped together with the corresponding males, according to characters summarized
in Table i. Owing to confusion between the three species, records are unreliable
and so the ecology and distribution of the " anomalus " group of species is poorly
known.

Character

1. Maximum size

2. Antennule

a. Ratio of peduncular

articles

b. Average number of

flagellar articles

c. Number of articles to

accessory flagellum
excluding vestigial
terminal article

3. Uropod 3

a. Peduncle

b. rami

M. anomalus
10 mm.

23

3-4

Flanges poorly
developed, dorsal
and inner
longest
slender,
elongate

M. damnoniensis
4 5 mm.

14

Flanges well
developed inner
much the
longest
broad, robust

M. algicola
4-5 mm.

i*

3

as damnoniensis

as anomalus

TABLE i. The main identification characters for females of the "anomalus" group of species.
* = From Delia Valle (1893).

Microdeutopus anomalus (Rathke)
(Text-figs. 7a, 8a-e, QC, e, loc, f, 2ob, d)

Gammarus anomalus Rathke, 1843 : 63, pi. 4, fig. 7; Lilljeborg, 1855 : 457.

Autonoe anomala (Rathke) Bruzelius, 1859 : 25, fig. 4.

Autonoe sp. Smith, 1874 : 562.

Microdeutopus anomalus (Rathke) Boeck, 1870 : 157; Catta, 1875 : 167; Boeck, 1876 : 567,
pi. 25, fig. 5; Delia Valle, 1893 : 417, pi. 56, fig. 41 ; Sars, 1894 : 540, pi. 191 ; Sowinsky, 1898 :
480, pi. 10, figs. 20-24; Stebbing, 1906 : 591; Kunkel, 1910 : 76, fig. 29; Chevreux & Fage,
1925 : 298, fig. 309; Stephensen, 1927 : 124; Stephensen, i92ga : 152, fig. 275; Oldevig,
1933 : 212, fig. 94; Miloslavskaia, i939a : 123; Carausu & Carausu, 1942 : 74, fig. 5; Schellen-
berg, 1942 : 188, fig. 155; Soika, 1949 : 198; Gurjanova, 1951 : 833, fig. 582; Andersson,
1954 : 252, figs. 1-4; Barnard, 1958 : 29.

[Nori] Microdeutopus anomalus Bate, 1862 : 164, pi. 30, fig. 3.

Microdeuteropus anomalus (Rathke) Norman, i86ga : 281; Norman, 1886 : 16.

Microdeutopus propinquus Sars, 1894 : 542, pi. 192; Stebbing, 1906 : 592; Stephensen, 1927 :
125; Stephensen, i92ga : 152, fig. 274; Oedevig, 1933 : 212, fig. 95; Schellenberg, 1942 : 189,
fig. 156; Dahl, 1946 : 5; Gurjanova, 1951 : 834, fig. 583.

Microdeutopus danmoniensis [sic] Sars, 1894 : 542, pi. 192.

Microdeutopus stationis Sowinsky, 1895 : 237, pi. 4, figs. 1-6.

TYPE LOCALITY. Christiansund, Norway.

REVISION OF MICRODEUTOPUS in

DIAGNOSIS OF MALE. Antennule about two thirds the body length, the first
peduncular article shorter than the second, the third about one third the length of
the second; flagellum slightly less than twice the length of the peduncle, normally
with up to twenty-three articles (maximum recorded twenty-seven) ; accessory
flagellum with four to five articles, the terminal article of which is rudimentary.
Antenna two thirds the length of the antennule, the fourth and fifth peduncular
articles subequal; flagellum about equal to the fifth peduncular article, normally
with up to six articles (maximum recorded eight) . Per aeon segments 2-4 each with
a spine in the mid-ventral line, those of segments 2 and 3 the longest and most
slender, that of segment 4 the shortest and most robust ; segment 5 in some specimens
with a vestigial spine. Gnathopod i with article i strongly produced anteriorly,
very attenuated; article 2 elongate and relatively slender; article 5 very enlarged,
longer than broad, the posterior distal angle produced into a straight or slightly
recurved tooth, at the base of which in mature specimens is characteristically an
accessory tooth, which arises within the posterior margin and is not an extension
of it as in other Microdeutopus species ; article 6 with the posterior margin concave,
excepting the palmar angle; article 7 variable, usually relatively long and slender.
The accessory tooth on article 5 varies considerably in size, sometimes being so redu-
ced that it does not project over the posterior margin and appears obsolete; occasion-
ally in adults and frequently in juveniles it is absent. The aberrant form of this
species described and figured by Chevreux and Fage (1925) with two accessory teeth,
has not been observed in the present investigations. Gnathopod 2 with article 2
unexpanded; articles 5 and 6 elongate, 5 slightly the broader; article 6 almost three
times as long as broad, with the palmar angle oblique ; article 7 relatively long. Uropod
3 with the peduncle slender, its outer, inner and dorsal margins each produced
into a flange of which the dorsal and inner are the most pronounced ; rami equal to
or slightly exceeding the length of the peduncle ; endopodite slightly shorter than the
exopodite, with four spines on the inner margin and two on the outer, none of which
are closely associated; exopodite with two spines on the outer margin only. Telson,
with each terminal crest bearing on its distal dorsal margin, a pair of unequal
setae.

Maximum size of male : 8 mm.

FEMALE, As the male, except for the structure of the sexually dimorphic gnatho-
poda, and absence of ventral peraeon spines. Gnathopod i with article i somewhat
produced anteriorly, rounded; article 2 unexpanded; article 6 longer than article
5, distinctly broader distally, and constricted at the junction with article 5 ; article
7 longer than the palmar region. Gnathopod 2 with article 2 elongate and slender;
article 6 considerably longer and more slender than article 5.

Maximum size of female : 10 mm.

DISCUSSION. M. anomalus is distinguishable in the male, in its typical form, from
all other Microdeutopus spp. by the presence of an accessory tooth, arising within
the posterior margin of article 5 of gnathopod i (Text-figs. 8a, b, d, e). Past workers
have described M . anomalus as having this accessory tooth (or teeth) always present,
although this is by no means the case, particularly in immature material, and have
frequently utilized this character as a basis for a key (Stebbing, 1906, Chevreux &

A. A. MYERS

FIG. 7. Microdeutopus anomalus (Rathke). (a) Male head, Plymouth, Devon. Micro-
deutopus damnoniensis (Bate), (b) Male head, Plymouth, Devon.

REVISION OF MICRODEUTOPUS

FIG. 8. Microdeutopus anomalus (Rathke). (a) Male gnathopod i, Falmouth, Cornwall,
(b) Male gnathopod i, Bonifacio, Corsica, (c) Male gnathopod i, Falmouth, Cornwall,
(d) Male gnathopod i, Isle of Mull, Scotland. (e) Male gnathopod i, Isle of Mull, Scot-
land. Microdeutopus damnoniensis (Bate), (f) Male gnathopod i, Plymouth, Devon.
Microdeutopus algicola Delia Valle. (g) Male gnathopod i, Naples, Italy, (h) Male
gnathopod i, Bone, Algeria.

H4 A. A. MYERS

Fage, 1925) . This has often led to the attributing of immature material of the present
species to M. damnoniensis (Bate), the separation of which must be made with great
care. In its immature form, or in mature specimens lacking an accessory tooth,
M. anomalus is distinguishable from M. algicola Delia Valle by the form of the
primary tooth which arises at the posterior distal angle of article 5 and is relatively
slender, though never as slender as in M. damnoniensis. In M. algicola (Text-figs,
8g-h) a considerable area of the posterior margin of article 5 contributes to the
development of the tooth, which is hence very robust. M . anomalus differs from
M . damnoniensis also in the form of the tooth on article 5 of gnathopod I, but is more
easily distinguished from that species by the structure of the second gnathopod,
which is slender and elongate with an oblique palm in M. anomalus (Text-fig. QC),
relatively short and broad, with an almost transverse palm in M. damnoniensis
(Text-fig, ga). In addition, the rami of uropod 3 are long and slender in M. anomalus
(Text-fig. 96) ,whereas they are short and broad in M. damnoniensis (Text-fig. 9g).

ECOLOGY. A sub-littoral species in depths ranging down to 200 metres, but
occasionally extending into the intertidal zone. In areas influenced by detritus,
often among arborescent weeds, but also in Zostera beds, and among shells, poly-
zoans, sponges, tunicates and Mytilus. In the Black Sea, associated with Modiola
phaseolina and Amphiuraflorifera.

DISTRIBUTION. Norway south of the Lofoten Islands, Sweden, Denmark,
including the Baltic, the British Isles, particularly the South and West coasts, the
channel coasts of France, including the Channel Islands, along the Atlantic coasts
of France, Spain, Portugal and North West Africa to the Canary Isles. Throughout
the Mediterranean, Adriatic, Aegean, Ionian and Black seas, in the Bermudas and
on the north-eastern seaboard of the United States of America (Rhode Island and
Massachusetts).

Microdeutopus damnoniensis (Bate)
(Text-figs, yb, 8f, ga, g, zoa-b, 2oh, j)

Lembos damnoniensis Bate, 1856 (nomen nudum): 58; Bate, 1857 : 142.

Lembos danmoniensis [sic] (Bate) White, 1857 : 180.

Microdeutopus gryllotalpa Bate, 1862 : 163, pi. 30, fig. i; Bate & Westwood, 1863 : 289.

Microdeutopus damnoniensis (Bate) Norman, i9O5a : 24; Stebbing, 1906 (pro-parte) : 593;

Chevreux & Fage, 1925 (pro-parte): 297, fig. 308; Barnard, 1958 : 29.
Microdeutopus danmoniensis [sic] (Bate) Norman & Scott, 1906: 83.
[Non] Microdeutopus danmoniensis [sic] Sars, 1894:542, pi. 192.

TYPE LOCALITY. Plymouth, Devon, England.

DIAGNOSIS OF MALE. Antennule slightly over one half the body length, the first
and second peduncular articles subequal, the third almost one half the length of the
second; flagellum a little longer than the peduncle, normally with up to fourteen
articles (maximum recorded fifteen) ; accessory flagellum with two articles the
terminal article rudimentary. Antenna shorter than the antennule, the fourth
and fifth peduncular articles subequal ; flagellum about equal to the fifth peduncular
article normally with up to seven articles (maximum recorded eight). Per aeon

REVISION OF MICRODEUTOPUS 115

segments 2-4 each with an anteriorly directed spine in the mid-ventral line, that of
segment 2 the longest. Gnathopod i with article i moderately produced anteriorly,
less acute than in M. anomalus or M. algicola ; article 2 expanded somewhat distally ;
article 5 very enlarged, longer than broad, the posterior distal angle produced into a
single, long, slender, inward curved tooth which in fully mature specimens consider-
ably exceeds half the length of article 6 ; article 6 with the posterior margin smoothly
rounded; article 7 relatively long with accessory teeth on the posterior margin.
Gnathopod 2 with article 2 expanded; articles 5 and 6 short, 5 slightly the longer,
but scarcely broader than article 6, which is scarcely twice as long as broad, and has
the palmar angle almost transverse; article 7 short and robust. Uropod 3 having
the peduncle with its lateral and dorsal flanges very distinct, as in M. algicola;
rami short and very stout, not exceeding the length of the peduncle, the exopodite
the longer of the two rami, with two separated spines on the outer margin, the more
distal of which is much the longer, endopodite with a single median spine on the
outer margin, and a single more distal spine on the inner, each ramus terminating in a
group of spines of which one is relatively long. Telson as M. anomalus.
Maximum size of male : 4-5 mm.

FEMALE. As the male, except for the structure of the sexually dimorphic
gnathopoda, and the absence of ventral peraeon spines. Gnathopod i with article
i not markedly produced anteriorly, very obtuse; article 2 somewhat expanded;
article 6 slightly longer than article 5 and narrowing somewhat at the junction with
it; article 7 longer than the palmar region. Gnathopod 2 with article 2 unexpanded
and article 6 somewhat longer and more slender than article 5, with the palmar
margin angular.

Maximum size of female : 4-5 mm.

DISCUSSION. Andersson (1954) using material of Swedish origin concluded that
M. propinquus Sars and M. damnoniensis (Bate) were both synonyms of M. anomalus
(Rathke). Present investigations, whilst supporting the contention that
M. propinquus is synonymous with M. anomalus, indicate that M. damnoniensis
certainly is not. Since there are no substantiated records of M. damnoniensis
north of latitude 51 N. it is probable that Andersson was working entirely with a
population of M. anomalus. The distinction between M. anomalus and M. damnoni-
ensis is not simply one of presence or absence of an accessory tooth, and most so-
called M. damnoniensis in the literature are in fact immature M. anomalus. Never-
theless, true M. damnoniensis does exist and males are distinguishable from those of
M. anomalus and M. algicola by a number of important characters, including the
very slender, inward curved tooth on article 5 of gnathopod i (Text-fig. 8f), the short,
broad articles 5 and 6 of gnathopod 2 (Text-fig, ga), and the short, robust uropod
3 rami (Text-fig, gg). In addition, the number of articles to the accessory flagellum
of the antennule remains constant from eclosion to maturity, in contrast with the
condition in M. anomalus where the number of articles increases with age.

ECOLOGY. Unlike M. anomalus, the present species occurs most commonly in
the intertidal zone, among sponges, polyzoans and corallines, especially in rock
pools, or in the shallow sub-littoral.

u6

A. A. MYERS

FIG. 9. Microdeutopus damnoniensis (Bate), (a) Male gnathopod 2, Plymouth, Devon,
(g) Male uropod 3, with transverse section across peduncle at z, Plymouth, Devon.
Microdeutopus algicola Delia Valle. (b) Male gnathopod 2, Naples, Italy, (d) Male
telson, Naples, Italy, (f) Male uropod 3, with transverse sections across peduncle at
x and y, Naples, Italy. Microdeutopus anomalus (Rathke). (c) Male gnathopod 2,
Plymouth, Devon, (e) Male uropod 3, with transverse section across peduncle at w,
Plymouth, Devon.

REVISION OF MICRODEUTOPUS 117

DISTRIBUTION. Very difficult to ascertain, due to confusion in the literature with
immature M. anomalus. Present investigations have confirmed material from
Drake's Island and Wembury Bay, Plymouth, Jersey and Guernsey, and Bandol,
France.

TYPE MATERIAL. A lectotype has been erected from Bate's syntypic series (see
Appendix 2) .

Microdeutopus algicola Delia Valle
(Text-figs. 8g-h, gb, d, f, lod-e, 2of)

[?] Microdeutopus grillotalpa Sowinsky, 1880 : 125, pi. 5, fig. lya-d.

Microdeutopus algicola Delia Valle, 1893 : 4 1 **, pi. i, fig- 3. pi- n, figs. 1-12; Norman, igc^a :

25-

Microdeutopus damnoniensis (Bate) Stebbing, 1906 (pro-parte) : 593; Chevreux & Page, 1925
(pro-parte): 297, fig. 308; [?] Miloslavskaia, I939a : 124, fig. 24; [?] Soika, 1949 : 199, [?] Gur-
janova, 1951 : 834, fig. 584.

TYPE LOCALITY. Mergellina, Naples, Italy.

DIAGNOSIS OF MALE. Antennules and antennae have been missing in material
examined in present investigations, and the description of these appendages here is
extracted from the description and figures of Delia Valle (1893). Antennules with
the first and second peduncular articles sub-equal, the third about one third the
length of the second; flagellum longer than the peduncle with twenty articles;
accessory flagellum possibly with two articles, but almost certainly three, of which
the terminal is rudimentary as figured (for M. grillotalpa) by Sowinsky (1880).
Antenna shorter than the antennule, the fourth peduncular article slightly longer
and more robust than the fifth, flagellum about equal to the fifth peduncular article,
with eight articles. Per aeon segments 2-4 each with an anteriorly directed spine
in the mid- ventral line, that of segment 2 the longest; segment 5 with a vestigial
spine. Gnathopod I with article I moderately produced anteriorly, acute, though
less attenuated than in M. anomalus; article 2 relatively robust, expanded somewhat
distally; article 5 very enlarged, the posterior margin produced into a single, robust,
broad based tooth not exceeding half the length of article 6 ; article 6 with the posterior
margin convex distally, straight or slightly concave basally; article 7 robust, with
accessory teeth on the posterior margin. Gnathopod 2 with article 2 having the
anterior margin produced into a flange which terminates distally in a blunt process ;
articles 5 and 6 slender, subequal in length, article 5 slightly the broader; article 6
over twice as long as broad, with the palmar region oblique ; article 7 short and robust.
Uropod 3 with the peduncle broad, due to the marked development of the lateral
flanges, particularly the inner, which exceeds in length both the outer flange and
the dorsal crest; rami long and slender, about equal in length to the peduncle, the
exopodite slightly the longer, not differing in spination from M. anomalus of a similar
size. Telson as in M. anomalus.

Maximum size of male : 4-5 mm.

FEMALE. As the male, except for the structure of the sexually dimorphic gnatho-
poda, and absence of ventral peraeon spines. Gnathopod I with article I not
markedly produced anteriorly, obtuse ; article 2 robust, otherwise scarcely differing

n8

A. A. MYERS

from that of M. damnoniensis. Gnathopod 2 with article 6 having the palmar angle
rounded, otherwise as that of M. damnoniensis.
Maximum size of female 4-5 mm.

DISCUSSION. M. algicola is the most robust member of the " anomalus " group
of species. In facies it more closely resembles M. stationis, than it does M. anomalus

FIG. 10. Microdeutopus damnoniensis (Bate), (a) Female gnathopod i, Plymouth, Devon.

(b) Female gnathopod 2, Plymouth, Devon. Microdeutopus anomalus (Rathke).

(c) Female gnathopod i, Plymouth, Devon, (f) Female gnathopod 2, Plymouth, Devon.
Microdeutopus algicola Delia Valle. (d) Female gnathopod 2, Naples, Italy, (e) Female
gnathopod i, Naples, Italy.

or M. damnoniensis, and it is noteworthy that a male M. algicola was discovered
during present re-examinations of M. stationis material in the Chevreux collections.
Male M. algicola are distinguishable from those of M. damnoniensis, in having the
tooth on article 5 of gnathopod i (Text-figs. 8g-h) broad-based and triangular,
whereas in M. damnoniensis (Text-fig. 8f) it is slender throughout its length, arising

REVISION OF MICRODEUTOPUS 119

at the posterior distal angle only. The present species also differs from M. damnon-
iensis in the structure of gnathopod 2 (Text-fig, gb), which has articles 5 and 6
slender, as in M. anomalus and in the form of uropod 3 (Text-fig, gf), which has slender
elongate rami. In lacking an accessory tooth on article 5 of the male gnathopod i,
M. algicola differs from typical M. anomalus which usually possesses such a tooth.
In specimens of the latter species lacking an accessory tooth (including immature
specimens) the primary tooth is structurally very different, being longer and more
slender, with the posterior margin more smoothly rounded, and with a narrower
base, than it is in M. algicola. In addition, the flange on the anterior margin of
article 2 of gnathopod 2 terminates distally in a more markedly developed process
in M. algicola than in M . anomalus.

ECOLOGY. At Mergellina (Delia Valle, 1893) among attached algae on the shore,
and at Bone in 15-20 metre depth.

DISTRIBUTION. Difficult to ascertain, due to the scarcity of reliable records.
The present author has seen material from Naples, Italy and Bone, Algeria, and
considers that the M. grillotalpa Sowinsky from Sevastopol in the Black Sea, refers
to this species. The species may prove to be widely distributed in the Mediterranean
region, and Black Sea.

THE " SCHMITTI " GROUP OF SPECIES

The " Schmitti " group of species occurs over a wide geographical area, around
both the Pacific and Atlantic coasts of the Americas. So far material has been
examined from a number of isolated geographical localities, and regional morpholo-
gical differences are difficult to assign to varietal, subspecific and specific status.
Nevertheless there seems at present to be justification for grouping these forms into
at least three species, of which M. schmitti Shoemaker and M. hancocki Myers are
sympatric, overlapping in their distribution and maintaining their separate identities.
It is clear that ecological studies on the " schmitti " group of species would be of
considerable value.

Males of the " schmitti " group, with the exception of M. schmitti itself, differ
from those of all other species of Microdeutopus by the development of a tooth on
the anterior margin of article 5 of gnathopod i. M. schmitti differs from M. gryl-
lotalpa Costa (the only species with which it could be confused) by having article 2
of gnathopod 2 concave and smooth on the anterior margin, whereas in M. gryllo-
talpa the anterior margin is convex and crenated. Females of M. schmitti differ
from those of all other known species in the genus (but see also Hansenella longi-
cornis Chevreux, p. 138) by the development of a tooth or teeth on the posterior
margin of article 5 of gnathopod i. Females of other species in the " schmitti "
group are without recognizable diagnostic features and their identification in the
absence of males cannot be attempted with certainty. Most apparent differences
in structure between M. schmitti and the smaller species M. hancocki and
M. trichopus are due to neoteny. The rami of the third uropod of mature
female M. hancocki and M. trichopus, for example, correspond to those of immature
specimens of M. schmitti of a similar size.

I2O

A. A. MYERS

Microdeutopus schmitti Shoemaker
(Text-figs, n, i3b, d, i4a-c, f, g, 1, 15, i6a-b, d, 2om, pi. ib)

Microdeutopus schmitti Shoemaker, 1942 : 18, fig. 6; Barnard, 1958 : 29; Barnard, 1959 : 32,
pi. 9; Myers, 19685 : in press.

TYPE LOCALITY. Bahia de Magdalena, Baja California, Mexico.

DIAGNOSIS OF MALE. Pamgnaths with the mandibular processes relatively short
and stout. Antennule slightly over one third the body length, the first and second
peduncular articles subequal, the third a little less than one half the length of the

FIG. ii. Microdeutopus schmitti Shoemaker. Male head, Bahia de Salinas, Costa Rica.

second; flagellum about equal in length to the peduncle, normally with up to ten
articles (maximum recorded twelve); accessory flagellum with three articles, the
third article rudimentary. Antenna about two thirds the length of the antennule,
the fourth and fifth peduncular articles subequal; flagellum shorter than the fifth
peduncular article, with four articles, of which the first is equal to the combined
length of the terminal three. Peraeon segments 3-6 each with a short, robust,

REVISION OF MICRODEUTOPUS

FIG. 12. Microdeutopus hancocki Myers, (a) Male head, Bahia de Salinas, Costa Rica
(Paratype). Microdeutopus trichopus Myers, (b) Male head, Isabela Island, Galapagos
(Paratype).

122 A. A. MYERS

anteriorly directed spine in the mid- ventral line. Gnathopod i with article i moderately
produced anteriorly, rounded ; article 2 expanded, oval, article 4 roughly triangular,
with relatively short setae on the posterior margin ; article 5 longer than broad, with-
out teeth on the anterior margin, the posterior distal angle produced into a large,
stout tooth, at the base of which are from one to four small forward projecting
teeth; article 6 short, the posterior margin with a distal and proximal lobe; article
7 relatively long, with accessory teeth on its posterior edge. Gnathopod 2 with article

2 somewhat concave on its anterior margin, the anterior distal corner produced
into a rounded lobe; article 5 slightly longer than article 6 and expanded distally,
where it considerably exceeds the width of article 6; articles 4, and 5 with numerous
long, finely pectinate setae arising from the anterior and posterior margins. Uropod

3 with the rami subequal, slightly longer than the peduncle; endopodite with two
to four equally spaced spines on the inner margin and one or two on the outer;
exopodite with two well separated spines on the inner margin, and on the outer a
solitary spine, basal to which in fully mature adults is a pair of closely associated
spines, the outer of which is the longer. Telson with each terminal crest bearing
on its distal dorsal margin a group of three or four setae, two or three of which are
short and spiniform. Additional setae may appear spiniform during growth, which
is in contrast with the condition in M. stationis where the spines are always quite
distinct from the setae.

Maximum size of male : 5 mm.

FEMALE. As the male except for the absence of ventral peraeon spines, and the
sexually dimorphic gnathopoda. Gnathopod i with article 2 unexpanded; articles
5 and 6 subequal in length, article 5 distinctly the broader, with the posterior distal
angle produced into one or two small teeth, article 6 with the anterior margins
roughly parallel; article 7 slender, considerably longer than the palmar region with
accessory teeth on the posterior edge. Gnathopod 2 with article 2 unexpanded;
articles 5 and 6 subequal in length, article 5 roughly pentagonal with the anterior
margin almost straight, broadening distally, where it considerably exceeds the breadth
of article 6 ; anterior and posterior margins of articles 4 and 5 bearing long pectinate
setae.

Maximum size of female : 6 mm.

DISCUSSION. M. schmitti is closely related to M. hancocki and M. trichopus but
differs from these in the male, by having the anterior margin of article 5 of gnathopod
i (Text-figs. I3b, d) devoid of any toothlike processes and in the female by the pres-
ence of teeth on the posterior distal angle of article 5 of gnathopod i (Text-fig.
i6b,d).

ECOLOGY. In the sub-littoral to a recorded depth of 42 metres (off Point Loma
Light, California). Tolerant of a wide range of conditions, on mud, fine or course
sand, and rock substrata, among various vegetation, shells, and coral.

DISTRIBUTION (Text-fig. 15). Along the coasts of California (Cayucos, Point
Conception, Point Loma, San Clemente, and Newport), Baja California (El Coyote,
Bahia de San Quintin, Bahia de Magdalena and Cape San Lucas) and Costa Rica
(South Viradores Islands, Playa Blanca and Bahia de Salinas).

REVISION OF MICRODEUTOPUS

123

FIG. 13. Microdeutopus hancocki Myers, (a) Male gnathopod i, Bahia de Salinas, Costa
Rica (Holotype). (c) Male gnathopod i, Salango Island, Equador. Microdeutopus
schmitti Shoemaker, (b) Male gnathopod i, Bahia de San Quintin, Baja California.

(d) Male gnathopod i, Bahia de Salinas, Costa Rica. Microdeutopus trichopus Myers.

(e) Male gnathopod i, Isabela Island, Galapagos (Holotype). Microdeutopus sp. nov.?

(f) Male gnathopod i, Tortugas, Florida, U.S.A. (g) Male gnathopod i, Tortugas, Florida,
U.S.A.

ZOOL. 17, 4

8

124 A. A. MYERS

Microdeutopus hancocki Myers
(Text-figs. I2a, I3a, c, I4d, h, m, 15, i6c, e, 2ok, pi. la)

Microdeutopus hancocki Myers, ig68b : In press.
TYPE LOCALITY. Bahia de Salinas, Costa Rica.

DIAGNOSIS OF MALE. Antennules and antennae not noticably different from those
of M. schmitti. Paragnaths with the mandibular processes longer and more slender
than in M. schmitti. Per aeon segments 3-5 each with a short robust, anteriorly
directed spine in the mid- ventral line, segment 6 with a small or vestigial spine.
Gnathopod I with article I moderately produced anteriorly, rounded; article 2
markedly expanded anteroproximally ; article 4 roughly triangular, with very
long setae on the posterior margin ; article 5 longer than broad, oval, with a median
dentiform process on the anterior margin, and at the posterior distal angle, three
stout teeth, the most distal of which is the longest; article 6 short, with a single lobe
on the posterior margin ; article 7 relatively long, with accessory teeth on the posterior
edge. Gnathopod 2 with article 2 concave on the anterior margin ; article 5 consider-
ably longer than article 6 but scarcely broader; articles 4 and 5 with numerous
long, finely pectinate setae arising from the anterior and posterior margins. Uropod
3 and telson scarcely differing from those of M. schmitti of a comparable size.

Maximum size of male 3-8 mm.

FEMALE. As the male except for the absence of ventral peraeon spines, and the
sexually dimorphic gnathopoda. Gnathopod i similar to that of M . schmitti but
with the posterior margin of article 5 without teeth at the posterior distal angle.
Gnathopod 2 scarcely differing from that of M. schmitti, but with the pentagonal
shape of article 5 less well marked.

Maximum size of female 4-1 mm.

DISCUSSION. M. hancocki is very closely related to M. schmitti, from which it
differs in the male having a tooth-like process on the anterior margin of article 5
of gnathopod i (Text-fig. I3a, c), and article 4 of this appendage bearing very long
setae. In addition the structure of the ventral peraeon spines (Text-fig. 2ok)
and paragnath (PI. la) and the shape of the second gnathopod (Text-fig. I4m)
distinguishes the present species from M. schmitti. It differs from M. trichopus
by having article 6 of gnathopod i devoid of any forward projecting teeth on its
posterior margin.

ECOLOGY. Apparently more habitat-specific than M. schmitti, having been
recorded at present only from sandy bottoms to a depth of about 18 metres.

DISTRIBUTION (Text-fig. 15). Costa Rica (Bahia de Salinas), Panama (Bahia
Honda), Equador (Salango Island), and Galapagos (Isabela Island).

crest of male telson, Bahia de Salinas, Costa Rica (Paratype). (m) Male gnathopod 2,
Bahia de Salinas, Costa Rica (Paratype). Microdeutopus trichopus Myers. (e) Male
uropod 3, Isabela Island, Galapagos (Holotype). (i) Dorsal view of left terminal crest
of male telson, Isabela Island, Galapagos (Holotype). (j) Male gnathopod 2, Isabela
Island, Galapagos (Paratype). Microdeutopus sp. nov.? (k) Male gnathopod 2, Tortugas,
Florida.

REVISION OF MICRODEUTOPUS

125

FIG. 14. Microdeutopus schmitti Shoemaker, (a)-(c) Male uropod 3, all figured the same
size (and therefore each at a different scale) for easy comparison, (a) 3-0 mm. $ (b)
4-5 mm $ (c) 5-0 mm. $, Bahia de San Quintin, Baja California, (f) (g) Dorsal view of
left terminal crest of male telson, Bahia de San Quintin, Baja California. (1) Male gnatho-
pod 2, Bahia de San Quintin, Baja California. Microdeutopus hancocki Myers, (d) Male
uropod 3, Bahia de Salinas, Costa Rica (Holotype). (h) Dorsal view of left terminal

126 A. A. MYERS

Microdeutopus trichopus Myers
(Text-figs. I2b, 136, 146, i, j, 15, i6f-g, 20g)
Microdeutopus trichopus Myers, ig68b : In press.

TYPE LOCALITY. East of south end of Isabela Island, Galapagos.

DIAGNOSIS OF MALE. Antennule slightly over one half the body length, the first
peduncular article slightly shorter than the very slender second article, the third
about a third the length of the second article; flagellum somewhat longer than the
peduncle, normally with up to twelve articles (maximum recorded thirteen) ;
accessory flagellum with three articles, the third rudimentary. Antenna about two
thirds the length of the antennule, the fourth and fifth peduncular articles elongate,
slender subequal; flagellum shorter than the fifth peduncular article with four, rarely
five articles, the first article long, about equal to the combined length of the terminal
three or four. Peraeon segments 3-6 each with a long and slender spine in the mid-
ventral line, those of segments 3 and 4 directed anteriorly those of segments 5 and 6
straight or slightly recurved. Gnathopod I with article I moderately produced
anteriorly, rounded ; article 2 expanded anteroproximally ; article 4 short and bulky ;
article 5 oval, longer than broad, with a median dentiform process on the anterior
margin and at the posterior distal angle, a short stout tooth, basal to which on the
posterior margin is a further small tooth ; article 6 over one half the length of article
5, the palmar angle produced into a short, blunt ending, forwardly projecting tooth,
opposable to article 7; article 7 of moderate length, with accessory teeth on the
posterior margin ; the posterior margins of articles 4, 5 and 6 densely setose, the setae
exceptionally long, in the largest males measuring 0-5 mm., 0-4 mm. and 0-3 mm.
on each article respectively. Gnathopod 2 with article 2 somewhat expanded distally,
the anterior margin slightly concave; articles 5 and 6 subequal in length, article
5 slightly the broader. Uropod 3 with the rami long and slender, longer than the
peduncle, the exopodite slightly the longer of the two rami; each ramus with a single
spine on each of the inner and outer margins. Telson with each terminal crest
bearing on its distal dorsal margin a long seta and a short stout spine.

Maximum size of male : 3-8 mm.

FEMALE. As the male except for the absence of ventral peraeon spines, and
sexually dimorphic gnathopoda. Gnathopod I with article 2 unexpanded, articles 5
and 6 slender, subequal in length, article 6 somewhat broader distally than at the
junction with article 5; article 5 without teeth at the posterior distal angle; article
7 longer than the palmar region, with accessory teeth on the posterior edge. Gnatho-
pod 2 with article 2 unexpanded; articles 5 and 6 subequal in breadth, article 6
slightly the longer, broadening distally; article 7 short and stout.

Maximum size of female : 4-0 mm.

DISCUSSION. M. trichopus differs in the male from all other known Microdeutopus
species (but see following form and Lembopsis Pearse, (p. 139), in having the posterior
margin of article 6 of gnathopod I (Text-fig. 130) produced into a forward projecting
tooth opposable to article 7.

ECOLOGY. M. trichopus appears to be a relatively deep water species having

REVISION OF MICRODEUTOPUS

127

been absent from samples of M. hancocki taken in shallow waters in geographical
areas where the present species is known to occur. Recorded from 58-110 metres
over mud, and also among nullipores on sandy bottoms.

DISTRIBUTION. Galapagos (two localities off Isabella Island).

Microdeutopus sp. nov.?
(Text-figs. i3f-g. i4k, 15, 20)

Two males and a single female of a species of Microdeutopus of the " schmitti "
group, were examined, from Tortugas, Florida. This material most closely resembles
M. trichopus from the Galapagos archipelago but it differs from that species particu-
larly in the structure of the male gnathopod i. This has article 4 elongate and

M. schmitti Shoe.
M. hancocki Myers
M.trichopus Myers
] Microdeutopus sp. nov. ?

FIG. 15. The recorded distribution of the " schmitti " group of species in the Americas.

rectangular in the present material, whereas it is short and triangular in M. trichopus,
in addition article 6 is short and article 7 opposable to the teeth of both articles 5
and 6 in present material, whereas in M. trichopus article 6 is over half the length
of the article 5, with article 7 opposable to the teeth of article 6 only. Lastly, the
setation of gnathopod i is considerably more sparse in present material than in
M . trichopus. On gnathopod 2 article 5 is slightly longer than article 6 in present
material, whilst the same two articles are subequal in M. trichopus.

128

A. A. MYERS

FIG. 16. Microdeutopus schmitti Shoemaker, (a) Female gnathopod 2, Bahia de San Quin-
tin, Baja California, (b) Female gnathopod i, Bahia de San Quintin, Baja California,
(d) Female gnathopod i, Bahia de Salinas, Costa Rica. Microdeutopus hancocki Myers,
(c) Female gnathopod 2, Bahia Honda, Panama (Paratype). (e) Female gnathopod i,
Bahia Honda, Panama (Paratype). Microdeutopus trichopus Myers, (f) Female gnatho-
pod 2, Isabela Island, Galapagos (Paratype). (g) Female gnathopod i, Isabela Island,
Galapagos (Paratype).

REVISION OF MICRODEUTOPUS 129

The present form is separated geographically from M. trichopus by the Central
American isthmus, which has not been severed since the late Miocene, so one might
expect the two forms to be specifically distinct. This possibility is strengthened by
what is known so far of the ecology of the two forms, M. trichopus occurring in, and
apparently restricted to, muddy and sandy bottoms in over 50 metres depth, while
present material is found in relatively shallow waters amongst weeds.

More abundant material from a wider range of localities is required before the
true taxonomic status of the present form can be elucidated.

DESCRIPTION OF MALE. Antennules and antennae missing. Per aeon segments
3-6 each with a spine on the mid- ventral line, those of segments 3 and 4 directed
anteriorly, those of segments 5 and 6 straight or slightly recurved. Gnathopod I
with article i moderately produced anteriorly, rounded ; article 2 markedly expanded
anteroproximally; article 4 very elongate, roughly rectangular; article 5 oval,
longer than broad, with a median dentiform process on the anterior margin, and at
the posterior distal angle, a short stout tooth, basal to which, on the posterior margin,
are two further small teeth; article 6 about one third the length of article 5, the
palmar angle produced into a short, blunt ending, irregular, forward projecting
tooth, basal to which is a small lateral projection; article 7 relatively long, opposable
to the teeth of both articles 5 and 6; the posterior margins of articles 4, 5, and 6
bearing long setae. Gnathopod 2 with article 2 very slender basally, markedly
expanded proximally ; article 5 somewhat longer and broader than article 6. Uropod 3
with the rami long and slender, longer than the peduncle, the exopodite the longer
of the two rami, each ramus with a single spine on its inner margin. Telson with
each terminal crest bearing on its distal dorsal margin, a long seta, and a short
spine.

Length of males : 3-4 mm. and 3-5 mm.

FEMALE. Antennules missing. Antennae about one third the body length, the
fourth and fifth peduncular articles elongate, slender, subequal; flagellum shorter
than the fifth peduncular article, with four articles, the first article long, slightly
shorter than the combined length of the terminal three. Gnathopods i and 2 not
differing from those of M. trichopus.

Length of female : 3-3 mm.

MATERIAL, i $ and i $; station 48-30, Qth August, 1930, east and south of
Loggerhead Light, Dry Tortugas, Monroe Co., Florida, in 20-20 metres, i <$;
station 12-31, 26th June, 1931, Tortugas, Florida (24 36' N. 82 56' W. approx.
from chart) in 18-20 metres (low water) net tow-weedy haul.

Microdeutopus armatus Chevreux
(Text-figs, ryd, 18, 20!)

Microdeutopus armatus Chevreux, 1886 : XLI; Chevreux, i88jb : 312, pi. 5, figs. 11-12;

Chevreux, iSSya : 92; Stebbing, 1906 : 589; Chevreux & Page, 1925 : 296, figs. 303, 307;

Barnard, 1958 : 29.
Stimpsonella armata (Chevreux) Delia Valle, 1893 : 422, pi. 4, fig. 8, pi. n figs. 13-24; Chevreux,

1900 : 89; Norman i905a : 25.

130

A. A. MYERS

TYPE LOCALITY. North East of Basse-Kikerie, France.

DIAGNOSIS OF MALE. Antennule between one half and two thirds the body length,
the second peduncular article longer than the first, the third a little over one third
the length of the second; flagellum longer than the peduncle, normally with up to
fourteen articles (maximum recorded sixteen) ; accessory flagellum with two articles,

FIG. 17. Microdeutopus chelifer (Bate), (a) Male head, with (b) a transverse section and
(c) a lateral view of the third peduncular article of the antenna, Jersey, Channel Islands.
Microdeutopus armatus Chevreaux. (d) Male head, Villefranche, France.

REVISION OF MICRODEUTOPUS 131

the second article rudimentary. Antenna about two thirds the length of the antennule,
the fourth and fifth peduncular articles subequal ; flagellum slightly shorter than the
fifth peduncular article with five articles. Peraeon segments 2-5 each with a spine in
the mid- ventral line, that of segment 2 slender hook-shaped, directed anteriorly, those
of segments 3-5 robust anteriorly directed, that of segment 5 the shortest. Gnathopod

1 with article I moderately produced anteriorly, acute; article 2 short and greatly
expanded ; article 5 almost as broad as long, the posterior distal angle produced into a
large, broad, blunt ended tooth; article 6 short, the posterior margin irregularly lobed;
article 7 relatively long. Gnathopod 2 with article 2 expanded, the anterior margin
produced into a crenated flange; articles 5 and 6 roughly rectangular, subequal in
length and breadth, article 6 with the palmar angle produced into a short tooth.
Uropod 3 with the rami subequal, slightly shorter than the peduncle; endopodite
with a pair of spines on the inner dorsal margin near the apex, and one long and one
short terminal seta, exopodite with a similar pair of terminal setae. Telson with
each terminal crest bearing, on its distal dorsal margin, a long seta.

Maximum size of male : 3 mm.

FEMALE. As the male except for the structure of the sexually dimorphic gnatho-
poda, and absence of ventral peraeon spines. Gnathopod i with article 2 somewhat
dilated; article 6 longer than article 5, with the anterior and posterior margins roughly
parallel, article 7 very elongate, exceeding in length the palmar region. Gnathopod

2 with article 2 expanded, the antero-distal corner produced into a lobe; article 6
considerably longer than article 5, with the palmar angle produced into a broad-
based triangular process.

Maximum size of female : 4 mm.

DISCUSSION. M. armatus differs in the male from all other species in the genus,
except M. chelifer, in having the second gnathopod chelate. It differs from the
latter species by having the fifth and sixth articles of gnathopod 2 (Text-fig. i8d),
subequal in length and breadth, whereas in M. chelifer (Text-fig. IQC), article 6 is
much shorter and much broader than article 5. There is no justification for placing
these two species together in a distinct genus (Stimpsonella Delia Valle) as suggested
by Delia Valle (1893) on the basis of the chelate male second gnathopoda, since in
other respects they exhibit no greater affinity with each other than they do with other
Microdeutopus species. The most generally useful character enabling the identi-
fication of females of M. armatus is the structure of the second gnathopod (Text-fig.
i8e), which has a pronounced projection at the antero-distal corner of article 2,
and a triangular process at the palmar angle of article 6. In addition article 2 of the
first gnathopod (Text-fig. i8a) is somewhat dilated.

ECOLOGY. On soft grey mud (type locality), fine or coarse sands, in the sub-
littoral, in 10-90 metres. At Naples (Delia Valle, 1893), associated with Urothoe
and Ampelisca.

DISTRIBUTION. France, from the type locality, southwards along the coasts of
the Bay of Biscay (Le Croisic, Gulf of Gascony), Portugal, and in the Mediterranean
off the coasts of France (Antibes, Villefranche) Corsica (Bonifacio) and Italy (Naples).

A. A. MYERS

Microdeutopus chelifer (Bate)
(Text-figs. I7a-c, 19, 2on)

Stimpsonia chelifera Bate, 1862 : 162, pi. 29, fig. 9; Bate & Westwood, 1863 : 285; Stebbing,

1878 : 34, pi. 5, figs. 2-3; Norman, 1886 : 17.
Microdeutopus chelifer (Bate) Stebbing, 1888 : 334; Stebbing, 1906 : 589; Chevreux & Page,

1925 : 295, fig. 306; Barnard, 1958 : 29.

[Nori\ Microdeutopus chelifer Haswell, 1879 : 340, pi. 22, fig. 3.
Stimpsonella chelifera (Bate) Delia Valle, 1893 : 424, pi. 56, figs. 42-45; Norman, igosa : 25;

Norman, I905b : 82; Norman & Scott, 1906 : 84; Norman, 1907 : 368.

TYPE LOCALITY. Salcombe, Devon, England.

DIAGNOSIS OF MALE. Antennule slightly over one half the body length, the second
peduncular article a little longer than the first, the third over one third the length
of the second; flagellum longer than the peduncle, normally with up to fifteen articles
(maximum recorded seventeen); accessory flagellum with three articles, the third

FIG. 1 8. Microdeutopus armatus Chevreux. (a) Female gnathopod i, Le Croisic, France,
(b) Male gnathopod i, Le Croisic, France, (c) Male uropod 3, Le Croisic, France, (d)
Male gnathopod 2, Le Croisic, France, (e) Female gnathopod 2, Le Croisic, France,
(f) Male telson, Le Croisic, France.

REVISION OF MICRODEUTOPUS

133

article rudimentary. Antenna longer than the antennule, the third peduncular
article with its lower outer margin produced into a flattened lobe, the fourth pedun-
cular article very slightly shorter than the fifth; flagellum longer than the fifth
peduncular article, normally with up to ten articles (maximum recorded eleven).

FIG. 19. Microdeutopus chelifer (Bate), (a) Male gnathopod i, Paignton, Devon, England,
(b) Male gnathopod i, articles 4 and 5, Khios, Greece, (c) Male gnathopod 2, Paignton,
Devon, England, (d) Male (Immature) gnathopod 2, articles 6 and 7, Khios, Greece,
(e) Male uropod 3, Brittany, France, (f) Female gnathopod i, Guernsey, Channel
Islands, (g) Female gnathopod 2, Guernsey, Channel Islands.

134

A. A. MYERS

Peraeon segments 2-4 each with an anteriorly directed process in the mid-ventral
line, those of segments 2 and 3 long, slender, spiniform, that of segment 4 short,
robust. Gnathopod i with article I strongly produced anteriorly, very acute;
article 2 scarcely expanded; article 5 very enlarged, but longer than broad, the
posterior margin produced sub-distally into a long, narrow, inward curving tooth,
distal to which in some, but not all specimens, is a further small tooth; article 6
relatively long, the posterior margin undulate; article 7 robust. Gnathopod 2 with
article 2 narrow and elongate, with a small projection at the antero-distal corner;
article 6 shorter and broader than article 5, with the palmar region produced into a
process, which in young males is short and broad, bearing a stout spine at its
tip, but in mature males is drawn out into a long inward curving tooth, when the
spine is usually obsolete; article 7 very long, opposable to the palmar process.
Uropod 3 with the rami elongate and slender, subequal in length with the peduncle ;

FIG. 20. Ventral peraeon processes in Microdeutopus species, (a) M. gryllotalpa Costa
Isefjord, Denmark, (b) M. anomalus (Rathke) Falmouth, Cornwall, England, (c) M.
versiculatus (Bate) Falmouth, Cornwall, England, (d) M. anomalus (Rathke) Plymouth,
Devon, England, (e) M. stationis Delia Valle Guernsey, Channel Islands, (f) M. algi-
cola Delia Valle Bone, Algeria, (g) M. trichopus Myers Isabela Island, Galapagos,
(h) M. damnoniensis (Bate) Bandol, France, (i) M. sp. nov. ? Tortugas, Florida, U.S.A.
(j) M. damnoniensis (Bate) Plymouth, Devon, England, (k) M. hancocki Myers Bahia
de Salinas, Costa Rica. (1) M. armatus Chevreux Villefranche, France, (m) M. schmitti
Shoemaker Bahia de Salinas, Costa Rica, (n) M. chelifer (Bate) Jersey, Channel Islands.

REVISION OF MICRODEUTOPUS 135

exopodite slightly the longer of the two rami, with two isolated spines on the outer
margin, of which the more distal is particularly long; endopodite with two well
separated spines on the inner margin and one on the outer; each ramus with a term-
inal cluster of spines, of which one is greatly elongated. Telson with each terminal
crest bearing on its distal dorsal margin, a group of up to 3 setae.
Maximum size of male : 8 mm.

FEMALE. As the male except for the absence of ventral peraeon spines and the
sexually dimorphic antennae and gnathopoda. Antenna with the third peduncular
article unmodified, and the entire appendage less well developed than that of the
male, such that it does not exceed the length of the antennule. Gnathopod i with
article 2 moderately expanded; article 6 longer than article 5, about twice as long
as broad, with the anterior and posterior margins roughly parallel; article 7 robust,
equal in length to the palmar region. Gnathopod 2 similar to gnathopod I out more
slender, the palmar region more transverse.

Maximum size of female : 8 mm.

DISCUSSION. As in the preceding species, the second gnathopoda of the male
are chelate. In the present species, however, article 6 of this appendage (Text-fig,
igc) is much shorter and broader than article 5, whereas in M. armatus (Text-fig.
i8d) the fifth and sixth articles are more or less subequal in length and breadth.
M. chelifer is unique among the known Microdeutopus species in having sexually
dimorphic antennae, but this feature is not considered to be a sufficient reason for
separating the present species from the genus Microdeutopus (see also Stebbing,
1888). Females have few diagnostic characters, but important identification features
when taken in combination are : the accessory flagellum of the antennule having
three articles (Text-fig. lya), the third uropod rami having their sparse spination
of solitary spines, the more distal of two spines on the outer margin of the exopodite
being exceptionally long (Text-fig. IQC) and the telson bearing setae, but no
spines.

ECOLOGY. In the scarcely tidal Mediterranean the present species occurs in
shallow waters, among Cystoseira, Chondrus, Saccorhiza etc. On the Atlantic and
English Channel coasts, the species is also found in the shallow sub-littoral among
Laminaria holdfasts and other weeds, but it most frequently inhabits the littoral
zone, among algae in rock pools often high up on the shore, particularly in sheltered
places with a high accumulation of silt, often where there is some brackish water
influence.

DISTRIBUTION. Along the south-western coasts of the British Isles (Studland,
Torbay, Wembury Bay, Salcombe, Port wrinkle) , the Channel Islands and the adjoin-
ing French coast, and thence at a number of localities along the coasts of the Bay of
Biscay to Guethery in the extreme south. Recorded from Cadiz, and probably
occurs in a number of favourable localities around the coasts of Spain and Portugal.
In the Mediterranean it has been recorded from the coast of France (Sete, Monaco,
Antibes, Villefranche) and in the Aegean from the Greek Island of Khios, off the
Turkish coast. The IStimpsonia sp. of Sowinsky (1898) from the Black sea may
refer to this species.

136

A. A. MYERS

Microdeutopus sporadhi sp. nov.
(Text-figs. 21, 22)

DESCRIPTION OF MALE HOLOTYPE. Mouthparts not differing significantly from
those of other members of the genus. Head lateral-lobes moderately well developed,
obtuse; eye reniform. Antennule about two thirds the body length, the first pedun-
cular article shorter than the second, the third less than one half the length of the

FIG. 21. Microdeutopus sporadhi sp. nov. Anterior end of male holotype, Emborios Bay,

Khios, Greece.

second; flagellum about one and one half the length of the peduncle, with thirteen
articles; accessory flagellum with two articles, the second article minute, shorter
than the first article of the primary flagellum. Antenna shorter than the antennule,
the fourth and fifth peduncular articles subequal ; flagellum scarcely longer than the
fifth peduncular article, with seven articles. Peraeon segments 2 and 3 each with a
robust, anteriorly directed spine in the mid-ventral line, that of segment 2 the longer.
Gnathopod i with article I strongly produced anteriorly, somewhat attenuated,
article 2 elongate, relatively slender, but broadening distally, its anterior margin
excavate for the reception of the carpus when folded, article 4 slender, scarcely over
lapping the posterior margin of article 5, article 5 grossly expanded, longer than broad,

REVISION OF MICRODEUTOPUS

137

slender basally, broad distally, with groups of long fine setae arising from within the
distal and postero-distal margins, and with the posterior distal angle produced
into two distinct teeth, the proximal (outer) tooth curved inward, relatively long
and slender, terminating acutely, the distal (inner) short, robust, obtuse, article 6
elongate and slender, broadening somewhat distally, but with the palmar region
almost obsolete, the posterior margin almost straight, article 7 very long, though
slightly shorter than article 6, with accessory teeth on the posterior margin. Gnatho-
pod 2 with article 2 relatively short and broad, the anterior margin almost straight,
articles 5 and 6 subequal in length and breadth, article 7 relatively short. Peraeo-
pods 3-7 of the typical Microdeutopus structure. Uropod 3 with the rami short,
not exceeding the length of the peduncle, exopodite slightly the longer of the two
rami, devoid of spines on the lateral margins, terminating distally in a group of
five long setae, of which the two most distal arise from a vestigial second article;
endopodite with a single median spine on the inner margin and a group of four

FIG. 22. Microdeutopus sporadhi sp. nov. Male paratypes. (a) Gnathopod i. (b) Telson.
(c) Uropod 3. (d) Paragnath. (e) Terminal article of maxillule palp, (f) Maxillule.
(g) Gnathopod 2. (h) Ventral peraeon processes, (i) Mandible.

138 A. A. MYERS

terminal setae. Telson of the usual Microdeutopus form, each terminal crest bearing
on its distal dorsal margin a pair of long setae of unequal length.
Length : 3-0 mm.

MALE PARATYPES. Agree closely with the holotype in all diagnostic characters.
Antennal flagellum of one paratype with 8 articles, otherwise this appendage also
invariable.

Length of paratypes 2-8-3-2 mm.

FEMALE. Unknown.

TYPES, <J Holotype B.M. (N.H.) Registration Number 1968.1.8.1

1 c? Paratype B.M. (N.H.) Registration Number 1968 .1.8.2

2 <$ Paratypes U.S.N.M. Catalogue number 122544.

TYPE LOCALITY. Emborios Bay, Khios, Greece.

MATERIAL. 6 $ from the type locality, August 8-nth, 1967.

DISCUSSION. M. sporadhi does not show close affinity with any of the previously
described species of Microdeutopus. Males differ from those of all other Micro-
deutopus species except M. haswelli Stebbing in the structure of gnathopod I (Text-
fig. 22a), which has the posterior distal angle of article 5 produced into two teeth,
of which the proximal (outer) is the longer. They differ from M . haswelli in the
structure of the antenna (Text-fig. 21), which is slender, with a relatively elongate
and slender flagellum, whereas in M. haswelli (Text-fig. 4b) it is sub-pediform, with
the first flagellar article stout and the terminal articles compressed and obscurely
articulate. In addition article 2 of gnathopod 2 is broad and relatively short in
M. sporadhi (Text-fig. 22g), but elongate and very slender in M. haswelli (Text-fig. 3h).

ECOLOGY. Type material from among Cystoseira in 0-2 metres where relatively
little water movement has allowed accumulation of detritus. Absent from Cysto-
seira growing nearby in deeper waters where there is relatively little detritus accumu-
lation.

INCERTAE SEDIS
Genus Hansenella Chevreux 1909

Hansenella longicornis Chevreux, 1909 : 5, fig. 3.

The genus Hansenella was established by Chevreux (1909) for the reception of
three ovigerous females obtained off the Azores at a depth of 1360 metres. The
genus is characterized by the form of the female gnathopod I, which is similar to
that of male Microdeutopus species. In the light of the present review, this feature
alone is not sufficient to erect a new genus, since development of teeth on article
5 of the female gnathopod i also occurs in M. schmitti Shoe. When males of H.
longicornis are known, it is likely that the species will have to be transferred to the
genus Microdeutopus, but it should be emphasized that the female gnathopod i is
far more massively developed in H. longicornis than in any known Microdeutopus
female.

REVISION OF MICRODEUTOPUS 139

SPECIES DUBIA
Microdeutopus tittii Heller

Microdeutopus tittii Heller, 1866 : 48, pi. 4, fig. 8.

It is doubtful if this form described from a single specimen collected at Pirano
in the Adriatic is a distinct species. Delia Valle (1893) considered it to be a much
mutilated specimen of M. gryllotalpa Costa, but even allowing for mutilation, the
undilated article 2 of gnathopod 2 would appear to precluds that species The
relative lengths of the antennules and antennae could apply to M. chelifer (Bate),
but the non-cheliferous gnathopod 2 precludes that species. Allowing for the mutila-
tion of the antennules and antennae, which is of frequent occurrence in the delicate
Aoridae, this species is most likely to be M. anomalus (Rathke) as suggested by
Stebbing (1906) or M. stationis Delia Valle. The long slender antenna with a
flagellum of eleven articles would most favour the latter species.

Species not attributable to Microdeutopus Costa

1. Microdeutopus anomalus Bate, 1862 : 164. = $ Aora typica Kroyer, 1845 : 328.

2. Microdeutopus arcticus Hansen, 1887 : 231. = Lembos arcticus (Hansen) Steb-

bing, 1895 : 207.

3. Microdeuteropus australis Haswell, 1879 : 27 1 = Lemboides australis (Haswell)

Stebbing, 1899 : 350.

4. Microdeuteropus bidentatus Stebbing, 1876 : 73. = Lembos websterii Bate,

1856 : 58.

5. Microdeutopus kraemmeri Reid, 1951 : 251. = Lembopsis kraemmeri (Reid)

Myers, ig6Sc : In press.

6. Microdeutopus longipes (Lilljeborg) Bate, 1862 : 166. = Lembos longipes

(Lilljeborg) Stebbing, 1895 : 207.

7. Microdeutopus maculatus Thomson, 1879 : 331. = $ Aora typica Kroyer, 1845

328.

8. Microdeutopus megnae Giles, 1888 : 231. = Grandidierella megnae (Giles) Chilton,

1921 : 548.

9. Microdeuteropus mortoni Haswell, 1879 : 339 = <$Aora typica Kroyer, 1845 : 328.

10. Microdeuteropus tenuipes Haswell, 1879 : 339 = %A oratypica Kroyer, 1845 : 328.

11. Microdeutopus tenuis (Dana) Bate, 1862 : 165. Lembos tenuis (Dana) Stebbing,

1895 : 207.

12. Microdeutopus tridens Schellenberg, 1938 : 74 = Lembopsis tridens (Schellen-

berg) Myers, 19680 : In press.

13. Microdeutopus websterii (Bate, 1862) : 164. = Lembos websterii Bate, 1856 : 58.

ACKNOWLEDGEMENTS

My thanks are due to Dr. E. Naylor for much helpful discussion and criticism, to
N.E.R.C. for a grant with which to carry out the work, to the Khios Committee for
defraying travel costs and to Professor E. W. Knight- Jones for laboratory

ZOOL. 17, 4. 9

i 4 o A. A. MYERS

facilities. I am indebted to the Trustees of the British Museum (Natural History)
for allowing me to study the collection, to Drs. J. L. Barnard, T. E. Bowman and
R. B. Manning (United States National Museum), Dr. J. S. Garth (Hancock Founda-
tion, University of Southern California), Dr. J. C. Yaldwyn (Australian Museum,
Sydney), Dr. M. Amanieu (Station Biolgique D'Arcachon), Dr. J. Forest (Museum
National D'Histoire Naturelle, Paris) and to Dr. E. Rasmussen and Dr. T. Wolff
(Universitetets Zoologiske Museum, Copenhagen) for invaluable loans of material,
and to the Director of the Plymouth Laboratory for facilities and access to collections.

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A. A. Ostroumovim v 1892 e 1893 gg. Zap. Kiev. Obshch. Estest. 15 (2) : 447-518, pis. 8-13.

144 A. A. MYERS

STEERING, T. R. R. 1874. Ampbipodous Crustacea. A new species and some items of Des-
cription and Nomemclature. Ann. Mag. Nat. Hist. s. 4, 14 : 10-15, P^ 8 - I ~ 2 -
1876. Description of a new species of sesile-eyed crustacean and other notices. Ann.

Mag. Nat. Hist. s. 4, 17 : 73-80, pis. 4-5.
1878. Notes on sessile-eyed crustaceans, with description of a new species. Ann. Mag.

Nat. Hist. s. 5, 1 : 31-37, pi. 5.
1888. Report on the Amphipoda collected by H.M.S. Challenger during the years 1873-76.

Rep. Challenger, 29 : 1-1737, pis. 1-210.
1895. Notes on Amphipoda old and new. Ann. Mag. Nat. Hist. s. 6, 16 : 205-213, pis.

8-10.

1899. Revision of Amphipoda. Ann. Mag. Nat. Hist. s. 7, 3 : 350.

1906. Amphipoda, i. Gammaridea. Tierreich, 21 : 1-806, figs. 1-127.

1910. Scientific results of the trawling expedition of H.M.C.S. " Thetis " off the coast of

New South Wales in February and March 1898. Crustacea. 5: Amphipoda. Mem. Aust.

Mus. 4 (12) : 565-658, pis. 47-60.
STEPHENSEN, K. 1927. Revideret Fortegnelse over Danmarks Alter af Amphipoda. 3.

Gammaridea: Gammaridae Podoceridae; Caprellidea. Vidensk. Meddr. dansk naturh.

Foren. 84 : 107-150.

I929a. Amphipoda. Tierwelt N.-u. Ostsee, 14 (lof) : 188 pp., 43 figs.

ig29b. Marine Crustacea Amphipoda. Zoology Faroes 23 : 1-40.

LE SUEUR, R. F. 1954. The Cumacea, Mysidacea, Amphipoda, Isopoda and Tanaidacea

of the Channel Islands. Bull. a. Soc. jersiaise, 16 (2) : 207-216.
TESCH, J. J. 1915. De amphipoden der zuidelijke Noordzee, verzameld met de " Wodan "

Rapp. Verhand. Rijksinst. Vissch. -Onderz. 1 (3) : 319-374, pis. 10-12.

1922. Amphipoden. Flora Fauna Zuiderzee, 326-336, figs. 1-7.

THOMSON, G. M. 1879. Additions to the amphipodous Crustacea of New Zealand. Ann.

Mag. Hist. s. 5, 4 : 329-333, pi. 16.
TRUCHOT, J. 1963. Etude faunistique et e"cologique des amphipodes des facias roucheux

intertidaux de Roscoff. Cah. Biol. mar. 4 : 121-176, figs. 1-16.
VALCANOV, A. 1936. Belejki verhu nasite brachicini vodi, 2. God. sof. Univ. 32 (3) : 209-

341, figs. 1-31.
WALKER, A. O. 1895. The Amphipoda of Bate & Westwood's " British sessile-eyed

Crustacea ". Ann. Mag. Nat. Hist. s. 6, 15 : 464-476.
1897. Malacostraca from the west coast of Ireland. Proc. Trans. Lpool biol. Soc. 12 : 159-

172.
WHITE, A. 1852. A popular history of British Crustacea. 358pp. 20 pis. London.

APPENDIX I

Catalogue of material examined in the present work.
KEY TO ABBREVIATION OF SOURCES:

A.A.M. Personal collections

A.M.S. Australian Museum Sydney

B.M.N.H. British Museum (Natural History)

E.R. Dr. E. Rasmussen

M.B.A.P. Marine Biological Association, Plymouth

M.N.H.N. Museum National D'Histoire Naturelle, Paris.

S.B.A. Station Biologique D'Arcachon

U.S.C. University of Southern California

U.S.N.M. United States National Museum (Smithsonian Institution)

U.Z.M.D. Universitets Zoologiske Museum, Denmark.

* Indicates material emanating from the collections of the original author of the species.

REVISION OF MICRODEUTOPUS

145

Location No. of samples

Microdeutopus gryllotalpa Costa

England

Chaldon Herring, Dorset . U.Z.M.D.

Weymouth, Dorset . . B.M.N.H.

Plymouth, Devon . . B.M.N.H.
Wales

Swansea, Glamorgan . . A.A.M.
Denmark

" Denmark "... B.M.N.H.

Bohuslen .... U.Z.M.D.

Isefjord .... E.R.

Roskilderjord . . . E.R.
France

LeCroisic . . . B.M.N.H.

Arcachon .... S.B.A.
Spain

Valencia .... B.M.N.H.
Italy

Naples .... B.M.N.H.
Greece

Khios .... A.A.M.
U.S.A.

" N.E. America "

Wood's Hole, Mass.

Marion Harbour, Mass

Vinyard Sound, Mass. .

Newport, Rhode Island

Noank, Connecticut

Microdeutopus versiculatus (Bate)

England

Plymouth, Devon

Salcombe, Devon

Mouth of river Yealm, Devon

Falmouth, Cornwall

Scotland

Isle of Mull

Oldany Harbour, Sutherland
Eke

Birterbuy Bay, Co. Galway .

Roundstone, Co. Galway

Westport, Co. Mayo
Channel Islands

Guernsey ....

A.A.M.

B.M.N.H.

U.Z.M.D.

M.B.A.P.

B.M.N.H.

B.M.N.H.

B.M.N.H.

U.Z.M.D.

B.M.N.H.
U.Z.M.D.

B.M.N.H.
B.M.N.H.
B.M.N.H.

B.M.N.H.

i
i

18
3

i
i

B.M.N.H.

I

U.S.N.M.

7

U.S.N.M.

i

U.S.N.M.

7

U.S.N.M.

4

U.S.N.M.

4

Microdeutopus stationis Delia Valle

Channel Islands

St. Peter's Port, Guernsey . B.M.N.H.

Material

76*18?
i<J 2?

1 <J i ? 3 immature

500 <J 500 ? 1000 immature

2 6* 2 ?
26*

43 6* 56 $ 13 immature

36* 4?

36* 3$

31 (J 21 ?

3?

56*9?
17 6* 26?

20 6* 37 ?

2?

72 $ 102 ? 4 immature

4 (J 3 ? i immature

8 3* 13 ? 8 immature

36*7?

io*i?

16*1?

i?

5 <J 10 ? 2 immature

21 <J 19 ? 4 immature

76*6?

36*6?
3?

4 cJ 8 ? 2 immature
9 # 12 ? 2 immature
36*8?

96*15?

146*6?

146

Corsica

Bonifacio .
Italy

*Naples
Algeria

B6ne
Greece

Khios

A. A. MYERS
Location No. of samples

I

M.N.H.N.
B.M.N.H.
M.N.H.N.
A.A.M.

Microdeutopus haswelli Stebbing

Australia

Port Jackson . . . A. M.S.

Microdeutopus anotnalus (Rathke)

England

Plymouth, Devon
Falmouth, Cornwall

Scotland

Argyll

Isle of Mull
Eire

Birterbuy Bay, Co. Galway

Westport, Co. Mayo

Valentia, Co. Kerry

Killiebegs, Co. Donegal
Norway

" Norway "
Denmark

Bohuslen .

France

Cette
Corsica

Bonifacio .
U.S.A.

Wood's Hole, Mass.

Marion Harbour, Mass.

Vinyard Sound, Mass. .

Rivers' Island, Mass. .

Provincetown, Cape Cod. Mass. U.S.N.M

Newport, Rhode Island

No Data .

Microdeutopus damnoniensis (Bate)

England

Plymouth, Devon

Drake's Island, Plymouth
Devon

Material

i?

4 5* ii ? i immature

i 6*2?
36*1$

i < HOLOTYPE

A.A.M.
B.M.N.H.
U.Z.M.D.

i
i

127 < 264 $ 149 immature
64 o* 67 ? 21 immature
16*

B.M.N.H.
B.M.N.H.

2
2

56*5?
I36*6i?

B.M.N.H.
B.M.N.H.
B.M.N.H.
B.M.N.H.

I
I

I
I

40 5* 50 ? 45 immature

i 6*1?
4 <J 5 ? i immature
1 6* 9 ? 3 immature

B.M.N.H.

I

i?

B.M.N.H.
U.Z.M.D.

I

I

16*1?
i6*i?

M.N.H.N.

I

26*

M.N.H.N.

I

7 (J 26 ? 12 immature

U.S.N.M.
U.S.N.M.
U.S.N.M.
U.S.N.M.

. U.S.N.M.

4

i
8

i
i

H6*i7?

2?
156*18?
2?
2 <? 3 2

U.S.N.M.
U.S.N.M.

[i., 4 . ,\

4
i

** U J T

5 6* 23 ? 2 immature

i?

Datej

B.M.N.H.
B.M.N.H.

i
i

i <$ LECTOTYPE
4 ? PARALECTOTYPES
2 ? 2 immature

U.Z.M.D.

i

8 6* 7 ? 3 immature

Channel Islands

Jersey

Guernsey .
France

Bandol

REVISION OF MICRODEUTOPUS
Location No. of Samples

B.M.N.H.
B.M.N.H.

M.N.H.N.

Microdeutopus algicola Delia Valle

Italy

*Naples .... B.M.N.H.
Algeria

B6ne .... M.N.H.N.

Microdeutopus schmitti Shoemaker

U.S.A.

Point Conception, California

Point Loma, California

San Clemente, California

Newport, California
Mexico

El Coyote, Baja California .

Bahia de San Quintin,
Baja, California

Bahia de Magdalena .

Baja California
Costa Rica

Bahia de Salinas

PlayaBlanca

South Viradores Islands

Microdeutopus hancocki Myers
Costa Rica

Bahia de Salinas
Panama

Bahia Honda
Equador

Salango Island .
Galapagos

Isabela Island

Microdeutopus trichopus Myers

Galapagos

Isabela Island U.S.N.M.

Microdeutopus sp. nov?
U.S.A.

Tortugas, Florida

U.S.N.M.

Microdeutopus armatus Chevreux
France

*LeCroisic . . . B.M.N.H.

*Antibes .... M.N.H.N.

* Villefranche M.N.H.N.

Material

10 $ 2 immature

16*

U.S.C.

2

24 C

* 64 $ 24 immature

u.s.c.

2

56*

5 $ 2 immature

u.s.c.

I

36*

2$

u.s.c.

6

3 o

6 $ i immature

U.S.N.M.

i

i 6*

U.S.N.M.

i

76*

19$

U.S.C.

i

2$

U.S.N.M.

9

5M

<J 261 $ 50 immature

U.S.N.M.

2

36*

U.S.N.M.

I

i 6*

U.S.N.M.

2

21 <J TYPE MATERIAL

U.S.N.M.

2

12 c

J 23 2 4 immature

U.S.N.M.

I

16*

U.S.N.M.

I

21 C

J 23 2 4 immature

38 <J 3 $ TYPE
MATERIAL

36*3$
26*8?

136*17$

148

A. A. MYERS

Microdeutopus chelifer (Bate)

England

Salcombe, Devon

Wembury Bay, Plymouth,
Devon ....

Paignton and Torbay, Devon

Studland, Dorset
Channel Islands

Guernsey ....

Jersey ....
France

Brittany ....

LeCroisic
Greece

Khios ....

Microdeutopus sporadhi sp. nov.
Greece

Khios

Location No. of samples

B.M.N.H.

U.Z.M.D.
B.M.N.H.
U.Z.M.D.

B.M.N.H.
U.Z.M.D.

B.M.N.H.
B.M.N.H.

A.A.M.

A.A.M.

Material

HOLOTYPE

3?
3?

39 6* 43 ?

66*

APPENDIX II

Location of type material of Microdeutopus species

M. haswelli Stebbing
M. damnoniensis (Bate)

M. schmitti Shoemaker
M. hancocki Myers

M. trichopus Myers

M. chelifer (Bate)
M. sporadhi sp. nov.

Holotype

Lectotype

Paralectotypes

Holotype

Holotype

Paratypes

Holotype
Paratypes

Holotype
Holotype
Paratypes

A.M.S. 3497

B.M.N.H. 1967.11.1.1

B.M.N.H. 1967.11.1.2-5

U.S.N.M. 79375
112797
112798-9
1967.7.2.1-4
112800
112801-2

U.S.N.M.

U.S.N.M.

B.M.N.H.

U.S.N.M.

U.S.N.M.

B.M.N.H.

B.M.N.H.

B.M.N.H.

B.M.N.H.

U.S.N.M.

1967

1952

1968

1968,

122544

7-2

5-7
i. 8. i
1.8.2

5-6

129

Type material of M. versiculatus (Bate) is not present in Bate's collections at the British
Museum (Natural History) and is therefore probably non-existant. Type material of species
described by Costa, Delia Valle, Rathke and Chevreux has been untraceable.

PLATE i

Microdeutopus hancocki Myers

(a) Male paragnaths, Bahia de Salinas, Costa Rics.

Microdeutopus schmitti Shoemaker

(b) Male paragnaths, Bahia de Salinas, Costa Rica.

Bull. BY. Mus. nat. Hist. Zoology 17, 4

PLATE i

B

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

<;

CONVERGENCE IN THE STRUCTURING
OF THE HEAD AND CUTICLE OF
EUCHROMADORA SPECIES AND
APPARENTLY SIMILAR NEMATODES

W. GRANT INGLIS

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 17 No. 5

LONDON: 1969

$wm

CONVERGENCE IN THE STRUCTURE OF THE L_* F *i*
HEAD AND CUTICLE OF EUCHROMADORA
SPECIES AND APPARENTLY
SIMILAR NEMATODES

BY

W. GRANT INGLIS

Department of Zoology, British Museum (Natural History), London, S.W.y.

and

Western Australian Museum, Perth, W.A.
(Now : Director, South Australian Museum, Adelaide, S.A.)

Pp. 149-204; 109 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY VoL 17 No. 5

LONDON: 1969

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in Jive series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 17, No. 5 of the Zoological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

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

Trustees of the British Museum (Natural History) 1969

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 4 February, 1969 Price i 2s.

CONVERGENCE IN THE STRUCTURE OF THE

HEAD AND CUTICLE OF EUCHROMADORA

SPECIES AND APPARENTLY

SIMILAR NEMATODES

By W. GRANT INGLIS

CONTENTS

Page

SYNOPSIS ........... 152

INTRODUCTION ........... 153

MATERIAL AND METHODS . . . . . . . . . 154

SPECIES STUDIED ......... 155

THE STRUCTURE OF THE HEAD ........ 156

THE STRUCTURE OF THE CUTICLE ....... 158

Euchromadora DE MAN, 1886 ........ 159

HEAD STRUCTURE ......... 159

Euchromadora vulgaris . . . . . . . . 160

Euchromadora gaulica . . . . . . . . 162

Euchromadora striata . . . . . . . . 162

Euchromadora eileenae . . . . . . . . 162

CUTICLE STRUCTURE . . . . . . . . . 163

DIAGNOSIS OF Euchromadora . . . . . . , 167

SPECIES REFERRED TO Euchromadora . . . . . 168

Steineridora GEN. NOV. ......... 168

DIAGNOSIS OF Steineridora . . . . . . . .170

Parapinnanema GEN. NOV. . . . . . . . . 171

DIAGNOSIS OF Parapinnanema . . . . . . . 175

Protochromadora GEN. NOV. . . . . . . . . 177

DIAGNOSIS OF Protochromadora . . . . . . . 179

Graphonema COBB, 1898 ......... 180

DIAGNOSIS OF Graphonema ........ 183

DIAGNOSIS OF Innocuonema GEN. NOV. ...... 183

Nygmatonchus COBB IN COBB, 1933 . . . . . . . 184

DIAGNOSIS OF Nygmatonchus . . . . . . . 184

Austranema GEN. NOV. ......... 185

DIAGNOSIS OF Austranema . . . . . . . . 187

UNCERTAIN Euchromadora AND Euchromadora-LiKv SPECIES . . . 188

REMARKS ON THE CLASSIFICATION OF THE CHROMADORIDAE . . . 189

DESCRIPTIVE SECTION . . . . . . . . . 191

Euchromadora striata (EBERTH, 1863) ...... 191

Euchromadora eileenae SP. NOV. ....... 193

Parapinnanema wilsoni GEN. ET SP. NOV. ..... 194

Graphonema georgei SP. NOV. . . . . . . . 195

Austranema alii (MURPHY, 1965) COMB. NOV. .... 195

Hypodontolaimus slacksmithi SP. NOV. ...... 197

SUMMARY OF GENERA AND SPECIES RECOGNIZED ..... 199

LIST OF NOMENCLATURAL CHANGES PROPOSED ..... 2OI

ACKNOWLEDGEMENTS ......... 202

REFERENCES ........... 202

ZOOL. 17, 5. ii

I 5 2 W. G. INGLIS

SYNOPSIS

The comparative anatomy and structure of the head and cuticle is described in several species
of Euchromadora and a number of other morphologically very similar species in which the cuticle
is complex and heterogeneous and the lateral pieces of the gubernaculum are massive and typi-
cally L-shaped. Euchromadora species are characterized by a solid dorsal onchium, a series of
denticles in transverse rows on both the lateral and ventral walls of the oesophastome, a cuticle
in which the large punctations are due to the presence of processes arising from one side of each
cuticular annule and there are no markings on the cuticle along the dorsal and ventral surfaces
over much of the length of the body. Other Euchromadora-like species which in the past have
been described as having an anteriorly directed dorsal onchium are shown to possess hollow-type
dorsal onchia; flanges on the lateral walls of the oesophastome; anteriorly directed, cone-like
onchia on the ventral sectors of the oesophastome; a cuticle in which the large punctations are
frequently due to processes developed from both sides of each cuticular annule and there are
markings on the cuticle of the dorsal and ventral surfaces along the whole length of the body.
As a result the following major taxonomic changes are proposed. Euchromadora de Man, 1866
retains seven species: E. vulgaris (Bastian, 1865), type species of the genus; E. gaulica Inglis,
1962; E. meadi Wieser & Hopper, 1967; E. permutabilis Wieser, 1954; - striata (Eberth, 1863);
E. tokiohai Wieser, 1955; E. eileenae sp. nov. STEINERIDORA Gen. Nov. is very similar to
Euchromadora but has large, sickle-like onchia arising from both the lateral and ventral walls
of the oesophastome and a distinct posterior oesophageal bulb. It contains the following four
species: S. loricata (Steiner, 1916), type species of the genus; S. adriatica (von Daday, 1901);
S. archaica (Steiner & Hoeppli, 1926); S. (?) dubia (Steiner, 1918) species inquirenda. In the
remaining genera the dorsal onchium is hollow. Graphonema Cobb, 1898 is rediagnosed on the
basis of Australian specimens which are undoubtedly congeneric with those described by Cobb
so that this genus is characterized by a swollen, set-off head, and two circles of cephalic setae.
It contains the following species: G. vulgaris Cobb, 1898, type species of the genus; G. georgei
sp. nov.; G. amokurae (Ditlevsen, 1921) Wieser, 1954. The rediagnosis of Graphonema displaces
species in which the dorsal onchium, although hollow, is controlled by a massive development
of the dorsal oesophageal musculature, there is a posterior oesophageal bulb and the guber-
naculum is small and not L-shaped. The four species involved are referred to a new genus
Innocuonema: I. flaccida (Wieser, 1959) type species of the genus; /. clovosa (Wieser, 1959);
/. tentabunda (de Man, 1890); /. chilensis nom. nov. pro G. amokuroides of Wieser, 1954. The
discovery of Nygmatonchus alii Murphy, 1965 led to a reassessment of Nygmatonchus Cobb in
Cobb, 1933 which is considered to contain four species: N. scriptus Cobb in Cobb, 1933, type
species of the genus; N. fossiferus Wieser, 1954; N. minutus Gerlach, 1967; N. bicornata (Wieser,
1959) comb, nov., in all of which the cuticle is laterally differentiated, the amphid has a prom-
inent double contour and there is a (?) pre-cloacal modification on the male body. N. alii
is referred to a new genus Austranema with three other species: A. colesi (Inglis, 1968), type
species of the genus; A. alii (Murphy, 1965); A. pectinata (Weiser & Hopper, 1967); A. shirleyae
(Coles, 1965), characterized by the possession of a circle of ten cephalic setae, a head which is
not set-off and the presence of a median pre-cloacal thickening on the ventral surface of the
males. A new genus and species, Parapinnanema wilsoni, is described in which the cuticle is
very thick and obviously battlement-like, the dorsal onchium is hollow and the lateral flanges
of the oesophastome are not dentate anteriorly, the cephalic sense organs are in three circles
of which the intermediate and outer are setae and there is a region of pre-cloacal modification on
the males. Finally a new genus, Protochromadora, is proposed for three species: P. scampae
(Coles, 1965), type species of the genus; P. mediterranea (Allgen, 1942); P. parafricana (Gerlach,
1958), in which there are no structures on the lateral walls of the oesophastome, the cuticle
lacks punctations on the dorsal and ventral surface and there is no pre-cloacal modification on
the ventral surface of the male. Five species are described from the coasts of Western Australia,
thus: Euchromadora striata (Eberth, 1863); E. eileenae sp. nov.; Graphonema georgei sp. nov.;
Parapinnanema wilsoni gen. et sp. nov. ; and Hypodontolaimus slacksmithi sp. nov. The classi-
fication of the Chromadoridae is briefly discussed in view of the recognition that complex cuticle,

EUCHROMADORA AND SIMILAR NEMATODES 153

structure of the dorsal onchium, form and distribution of cephalic sense organs and shape of lateral
pieces of the gubernaculum are not covariant. Nomenclatural changes are listed on page 201.

INTRODUCTION

THE classification of the nematode family Chromadoridae Filipjev, 1917 is largely
based upon the appearance of the cuticle and its associated lateral differentiation,
the shape of the amphids and the gross form of the dorsal onchium which occurs in
the buccal cavity. Thus de Coninck (1965) recognizes three subfamilies within the
family : Chromadorinae with a solid dorsal onchium and non-spiral amphids ; Hypo-
dontolaiminae de Coninck, 1965 with a hollow dorsal onchium and non-spiral
amphids; and Ethmolaiminae Filipjev & Schuurmans Stekhoven, 1941 with a hollow
dorsal onchium and spiral amphids. Within each of these subfamilies genera are
distinguished very largely by the distribution and form of the cephalic sense organs,
the appearance of the cuticle, the presence or absence of lateral differentiation and
the dentition of the buccal cavity.

Using such criteria the genus Euchromadora de Man, 1886 is apparently easily
recognized by a blunt head, a prominent solid dorsal onchium, a characteristically
dark and complicated cuticle and a gubernaculum of a unique L-shape. Several
authors have drawn attention to the possible value of the detailed structure of the
cuticle as a source of taxonomic data within the genus (Steiner & Hoeppli, 1926;
Inglis, 1962 ; Coles, 1965 ; Wieser & Hopper, 1967) but it has rarely been studied in
detail in one species (Steiner & Hoeppli, 1926; Inglis, 1964; Watson, 1965) and never
comparatively in a number of species.

The peculiar and characteristic shape of the gubernaculum has generally been
well described and appears to supply " One of the most decisive features of this genus
. . ." (Wieser, 1954, page 101). However it has never been shown that the character-
istic gubernaculum is consistently associated with the elaborate cuticle and the
large solid onchium. Some doubt about this assumed covariance is raised by
Murphy's (1965) description of Nygmatonchus alii Murphy, 1965 in which an
L-shaped Euchromadora-type gubernaculum is associated with a complex cuticle and
a hollow dorsal onchium. It will be shown below that this doubt is well justified.

The difference between a hollow and a solid onchium is spectacular and obvious
when the extremes of both conditions are compared but intermediate forms occur
which I have found difficult, if not impossible, to classify in this way with accuracy.
It was, therefore, thought necessary to establish whether the hollow and solid onchia
are associated with other more easily distinguished features before accepting this
difference as a basis for classification.

The difficulty of distinguishing between the solid and hollow condition has arisen
with others in the past since both Wieser (1954) and de Coninck (1965) refer Spilo-
phorella Filipjev, 1918 to the solid-onchia group while to me all the illustrations of
species referred to that genus, and all the specimens I have studied, are best described
as possessing hollow onchia.

Similar confusion has arisen in another way since both Wieser (loc. cit.) and de
Coninck (loc. cit.) group Odontocricus Steiner, 1918 and Dicriconema Steiner & Hoeppli,
1926, in both of which the cuticle is complex, with those genera in which a solid
dorsal onchium is definitely present. This can only have been based on an assump-

154 W. G. INGLIS

tion of covariance between the solid onchium and the complex cuticle since in neither
case is any information available about the structure of the dorsal onchium. In
Dicriconema the illustration of the anterior end of the body given by Steiner &
Hoeppli (1926, PI. i, fig. i) suggests that the dorsal onchium is hollow while the
uninformative description of 0. hupferi Steiner, 1918 simply mentions the presence
of a dorsal onchium. However this description is preceded by a description of
Euchromadora luederitzi Steiner, 1918 in which the dorsal onchium is illustrated as
hollow.

The unreliability of the gross appearance of the cuticle as an indicator of other
similarities is understandable when we recall the functional importance in the
Nematoda of a cuticle which is longitudinally flexible while tangentially strong
(Inglis, 1964). Attention has already been drawn to the narrow morphological
range within which the cuticle could be, or has been, modified (Inglis, 1965) so that
it is easy to appreciate that massive convergence in the appearance of the cuticle
is more likely to be the rule than the exception. Such convergences are unlikely to
be recognized so long as the structure of the cuticle is considered under such simple
headings as homogeneous/heterogeneous, simple /complex. However, the complexity
of the cuticle in some forms leads to very great difficulties in analysing its structure
in any greater detail than is implied by such simple groupings and here again it is
necessary, if possible, to establish the co-occurrence of other more easily studied
features.

It is clear that, as de Coninck (1965) stresses, the taxonomy of the Family Chroma-
doridae is in need of examination but it is difficult to see that any such revision can
be based upon a reassessment of the kind of information generally available. Too
much of the present classification is fortuitous, too many of the groupings recognized
are based on bibliographical information and too many of the morphological studies
have been carried out with the sole aim of delimiting species. The major difficulties
in overcoming this are the time needed to carry out comparative studies and the
small size of the specimens involved.

I have been able to carry out some such comparative studies on specimens collected
from the coasts of Western Australia. The specimens are in some cases confusingly
Euchromadora-like. in appearance but detailed examination disclosed discrepancies
which led me to study such species of the genus Euchromadora as were easily available
in large numbers. The unexpected results are presented here.

MATERIALS AND METHODS

Almost all the specimens were killed and fixed by gathering seaweed by hand and
putting it into plastic bags with seawater. Sufficient formalin was then added to
the bags to make a 7% formalin solution. The bags were shaken and nematodes
picked out later by hand under a stereoscopic dissecting microscope. Specimens
were then studied under a compound microscope ; uncleared and mounted in alcohol
or cleared and mounted in glycerine (prepared by a slow method involving the
evaporation of alcohol from an alcohol : glycerine mixture) ; or in lactophenol. The
structure of the cuticle was examined in squashes, in whole mounts, by transmitted
light, by phase contrast and after treatment with sodium hypochlorite. This latter

EUCHROMADORA AND SIMILAR NEMATODES 155

method produces separate rings of the sclerotized annules and rods of the cuticle.
Finally the rings of sclerotized cuticle were studied through crossed polaroid filters
when their birefringence makes them stand out spectacularly.

The heads were studied by mounting in glycerine jelly in a few cases but, more
commonly, they were mounted in liquid glycerine or lactophenol under cover-slips
supported by lens cleaning tissue. This latter method allows the head to be tilted
to any angle so that it can be studied in detail. In a few cases it was possible to
evert the anterior end of the oesophagus through the mouth opening so that the
dentition could be studied in great detail.

All whole mounts were fluid and temporary with the coverslips supported by
paper so that it was possible to get a view of the animals from any angle by rolling.

The use of Cobb slides or other form of permanent mount is to be deprecated for
morphological studies. Such slides are undoubtedly useful and simple for routine
identification but are a grave handicap in establishing the taxonomy of any group
of nematodes. The use of any permanent mount leads to the treatment of the
animals being examined as two dimensional and to the description of plates, seen in
optical section, as rods; to the description of flanges as teeth and to the analysis of
solid curved structures as hollow or pierced by holes or channels.

Studies of the kind reported here are generally time consuming and sometimes
difficult but experience in other fields of nematology has shown that en face views
of the head, although not always essential in the delimitation of species, are essential
in establishing any meaningful classification above the species level.

Most of the specimens studied were collected from various localities on the coast
of Western Australia during October, 1966, but some were collected on the coasts
of Europe, particularly the British Isles, and one sample originated from South
Africa. The holotype males of all new species are deposited in the collections of the
Western Australian Museum, Perth, Western Australia. Paratypes are deposited
in the same institution and in the collections of the British Museum (Natural
History), London, S.W.y.

Species Studied

Euchromadora adriatica (Von Daday, 1901)
Euchromadora colesi Inglis, 1968
Euchromadora eileenae sp. nov.
Euchromadora gaulica Inglis, 1962.
Euchromadora loricata (Steiner, 1916)
Euchromadora mediterranea Allgen, 1942.
Euchromadora scampae Coles, 1965.
Euchromadora shirleyae Coles, 1965.
Euchromadora striata (Eberth, 1863)
Euchromadora vulgaris (Bastian, 1865).
Graphonema georgei sp. nov.
Hypodontolaimus slacksmithi sp. nov.
Nygmatonchus alii Murphy, 1965
Parapinnanema wilsoni gen. et sp. nov.

156 W. G. INGLIS

THE STRUCTURE OF THE HEAD

The head of any nematode is derived developmentally from two sources, from the
anterior end of the region of primary embryonic invagination, the stomodaeum,
which becomes the oesophagus (there is some reasonable doubt about this inter-
pretation of the oesophagus as a stomadaeum, which is generally overlooked, and
the term is used here for convenience) and from a region of secondary overgrowth
derived from the hypodermis. These two regions are generally easily distinguished
in histological sections because the primary oesophageal region is lined by simple,
structurally uniform cuticle while the secondary region is lined by cuticle which is
continuous with and similar in appearance to the cuticle covering the outer surface
of the body.

The entire mouth cavity of nematodes I have proposed should continue to be
called the stoma; the region of secondary overgrowth continues to be known as the
cheilostome while any region of specialization at the anterior end of the oesophagus
(i.e. region of primary invagination) is the oesophastome (Inglis, 1965; 1967). The
wall of the cheilostome is the cheilorhabdion and that of the oesophastome is the
oesopharhabdion. Following Cobb's (1919) terminology as modified by Chitwood
(1950) I use the term odontium for any tooth-like structure developed from the
cheilostome and the term onchium for any similar structure developed from the
oesophastome. For convenience the general term denticle will also be used in this
paper to imply a small solid cuticular tooth-like structure developed from the
oesopharhabdion .

The cheilostome in the Chromadoridae is typically shallow, without odontia but
with a circular mouth opening surrounded by twelve rugae. In some cases the
terminal processes of the rugae are lacking and there are twelve flange-like supporting
structures which project into the cheilostome as curved wedges which may warrant
description as odontia (see page 198).

The cuticle covering the anterior end of the body is either without punctations
or, more posteriorly, the punctations are simple and rod-like. It is worth noting
that when the cuticle in Euchromadora is stained with Mallory's Triple Stain this
lightly punctate region stains various shades of light blue while the more elaborate
punctate component posteriorly stains a dense orange-red. This difference in
staining reaction appears to reflect a difference in the degree of sclerotization and
flexibility of the cuticle, the blue regions being flexible while the orange-red are
relatively rigid (Inglis, 1961).

In Euchromadora and similar forms, most of the variation in the structure and
elaboration of the anterior end of the body is restricted to the oesophastome, into
which a large dorsal onchium almost invariably projects. The oesophastome is in
the form of a cup formed by an expansion of the lumen of the oesophagus at
its anterior end. The cuticular walls of this oesophastomal cup are thick and
prominent and form curved sheets which blanket the cup tangentially. The
thickening of the cup extends posteriorly into the true tri-radiate lumen of the
oesophagus where it becomes increasingly thin posteriorly until it ceases to be obvious
(Text-figs. 3, 19, 43, 46).

EUCHROMADORA AND SIMILAR NEMATODES 157

The oesophastomal cup is roughly rectangular in transverse section (i.e. when
viewed en face). The dorso-lateral radii of the oesophagus arise from the dorso-
lateral corners of this rectangular cavity while the ventral arm of the lumen arises
from the mid-point of the ventral side of the rectangle. The boundaries between
the dorsal and lateral walls are, therefore, easily established by the dorso-lateral
arms of the oesophageal lumen. The boundaries between the lateral and ventral
walls (more correctly sub-walls for uniformity but called walls for convenience)
are frequently set-off by somewhat similar ventro-lateral extensions of the lumen
of the oesophagus. These extensions can be spectacularly obvious in some species.
In all cases, however, the limits can be established not only by the shape of the oeso-
phastome but also by the distribution of the modifications in the form of onchia
and plates present on the lateral and ventral walls (Text-figs. I, 6, 12, 13, 27, 30,
39, 42, 45, 47, 52, 58).

In typical Euchromadora species as exemplified by E. vulgaris, the type species
of the genus, the dorsal wall of the oesophastome is fairly straight, in an en face
view, and bears a series of wholly cuticular onchia (or denticles) along its anterior
edge (Text-figs. 12, 13). Posterior to this level is the large, wholly cuticular onchium
which projects into the oesophastome and articulates posteriorly on the thickened
dorsal wall of the oesophastome.

The anterior, most dorsal, edge of the lateral walls projects inwards over the
cavity of the oesophastome as a pointed, somewhat onchium-like flange which arises
from the ventral edge of the dorso-lateral radii of the lumen of the oesophagus.
Posterior to this extreme anterior edge the lateral walls bear a series of wholly
cuticular, generally tiny, denticles which run roughly dorso-ventrally (i.e. trans-
versely when viewed from the lateral aspect of the body) in distinct rows, of which
four can usually be recognized. In some cases one or more of these onchia is larger
than the others and the rows may be straight or bent (Text-figs. 2, 17, 24, 34).

The ventral walls of the oesophastome also bear a series of small, wholly cuticular
denticles with, usually, a series of much more prominent solid onchia arising from a
basal plate low down in the oesophastome or from the border between the ventral
walls and the median- ventral extension of the lumen of the oesophagus (Text-figs,
i, 6, 12, 13, 25, 27 etc.).

In all the species in which the dorsal onchium is unequivocably hollow, and in many
in which no decision can be easily reached, the conditions are consistently different.
The oesophastome is still in the form of a rectangular cup with thickened cuticular
walls but the dorsal onchium is not, or not obviously, solid. This means that the
musculature controlling it passes into the lumen of the onchium and does not attach
to its base as in solid onchia. The anterior edge of the oesophastomal cup, which is
generally strongly curved dorsally, is not dentate along its anterior edge, while
ventrally and laterally there are no rows of small denticles. Instead two large
cone-like cuticular onchia arise from the ventral floor of the oesophastome with, in
some species, another smaller similar pair which arises from the edge of the ventral
lumen (Text-figs. 30, 39, 42, 45, 52).

The lateral walls of the oesophastome do not bear series of denticles, instead,
there is a cuticular flange which stands independent of, but parallel to each lateral

158 W. G. INGLIS

wall (Text-figs. 44, 45, 52, 54). The anterior edge of this flange is generally higher
towards the dorsal surface of the body than the ventral and in some species is divided
along its anterior edge to produce a series of tooth-like points which project wholly
anteriorly. When viewed from the ventral aspect this flange looks like a tooth
arising from the lateral wall of the oesophastomal cup (Text-fig. 54; see also Fig.
21 in Coles, 1965). Its shape in en face views is very characteristic because it stands
free more posteriorly on its ventral edge than it does on its dorsal. This means
that when viewed in optical section en face it gives the appearance of being a ventrally
curved hook-like derivative from the wall of the oesophastome. The shape of this
lateral flange and the presence and distribution of the ventral cone-like onchia is
shown by Murphy (1965) in his figure of the head of Nygmatonchus alii (Murphy's

In all species the head bears sixteen cephalic sense organs of which six are arranged
in a circle near the edge of the mouth opening while the remainder may be arranged
in one circle of ten or as an intermediate circle of six and an outer circle of four.
In all the definitely Euchromadora species usually the outer sense organs only are
in the form of setae while in the others the intermediate circle is also composed of
setae.

THE STRUCTURE OF THE CUTICLE

The structure of the cuticle will be considered in detail later but it is only fair to
stress at this point that its analysis is very difficult. However the following features
are common to all those species with complex cuticles. The cuticle, except the
extreme anterior end, is dark in colour, thick and marked by large, elongate puncta-
tions anteriorly, at least. In addition, in many forms, the components of the cuticle
appear to move when the focus is adjusted under very high magnifications. This
elaborate structure is best understood against the functional background imposed
by the design of the nematode body but I would point out that I have never been
able to find any indication of fine " lines " connecting the punctations in a network
or any other arrangement.

The long, thin cylindrical nematode body with its system of longitudinal muscles
operating against an internal hydrostatic pressure produces two inevitable and
conflicting mechanical requirements: longitudinal flexibility combined with radial
or tangential strength (Inglis, 1964). This has been overcome during the evolution
of various groups of nematodes in one of, or a combination of both of, two ways:
by the evolution of either a system of cross spiral, helically arranged fibres or a series
of transverse rings of dense material within the cuticle.

The spiral fibre method resolves both problems at the same time but the transverse
ring method introduces further problems, particularly a loss of flexibility and the
presence of regions of relative weakness in the cuticle between the strong, dense
rings of sclerotized material. The latter problem has been resolved in a number of
ways such as the development of overlapping flaps of dense material which lie over
the weaker, less dense inter-annular material or by the development of tiling (de
Coninck, 1942). The most complicated method, however, appears to be that evolved
in some of the Chromadoridae, particularly those referred to the genus Euchromadora,

EUCHROMADORA AND SIMILAR NEMATODES 159

in which the inter-annular regions are strengthened by processes bridging the gaps
between the annules like the teeth of a comb.

In the typical Euchromadora species, in which the dorsal onchium is massive and
solid and the oesopharhabdion bears series of small denticles, the cuticle always
has non-punctate regions running down the dorsal and ventral surfaces of the body
(see Coles, 1965, Fig. i and discussion) while in the hollow onchium forms, in which
there are flanges laterally in the oesophastome, the cuticle does not have clear strips
dorsally and ventrally. Instead elongate punctations occur over the whole surface
of the body along its whole length, except on four regions, dorso- and ventro-lateral
in position, where the punctations are replaced by prominent articulating plates
(see Text-fig. 68 and also Steiner & Hoeppli, 1926, PI. i, 2, figs, i, 3, 4, 5).

The non-punctate region of the cuticle at the anterior end of the body is shallow
and, as a corollary, the amphids tend to lie near the anterior end in Euchromadora
species, while in the other group of species this non-punctate region is relatively
extensive and the amphids lie relatively far posteriorly. The amphids themselves
differ in being rather indistinct in Euchromadora, where they are bordered by slightly
raised fringes of cuticle, while in the other forms they are distinct with very prominent
wide cuticular fringes round their edges.

In all the descriptions of the cuticle that follow it must be remembered that the
conditions are reversed roughly half way along the length of the body. Thus when
anteriorly directed processes are described as occurring within the cuticle on the
anterior end of the body, those on the posterior half of the body will be directed
posteriorly.

Finally in all the descriptions of the cuticle the modifications referred to are repre-
sented by a dense, dark component lying within the cuticle and embedded in a less
distinct matrix. This dense component is generally in the form of transverse annules
which, in longitudinal section, are roughly rectangular with the short side of the
rectangle lying parallel to the antero-posterior axis of the body. (Text-figs. 60, 67,
69.)

EUCHROMADORA de Man, 1886

It is usual to defer using new names for taxa until the evidence on which the taxa
are based has been presented. Here there are so many changes in nomenclature
and so much discussion of evidence that to do so would make the entire presentation
even more complicated than it already is, or will become. I shall, therefore, present
the argument and evidence under various headings, usually the name of the genus
I finish by delimiting.

The genus which is central to the morphological argument is Euchromadora
which I consider first.

Head Structure

The immediate result of the detailed study of the structure of the head and
oesophastome is to suggest that it is of little value in delimiting species within the
genus. One difficulty is the technical problem imposed by the small size of some of
the specimens and another is that the appearance of the dentition varies dependent

160 W.G.INGLIS

upon the degree to which the month is open or closed. The following features are,
however, common to all the specimens, of all the species studied.

There is always a row of denticles along the dorsal edge of the oesopharhabdion
anterior to the large, solid dorsal onchium which is usually obviously bent about its
mid-length. The anterior edge of the oesophastome is always developed as an
anteriorly directed process on the most dorsal edge of the lateral wall of the oeso-
phastome, and sometimes a similar development appears to be present on the lateral
edges of the dorsal wall also.

The lateral and ventral walls of the oesophastome are always distinct and the
lateral bear at least four rows of transversely arranged small solid cuticular denticles
of which the most posterior (i.e. those lying deepest or most posterior within the
oesophastome), Row-IV, tend to be largest and generally arise from a common
thickened cuticular base or ridge. In some species the distribution of the denticles
on the lateral walls appears to be characteristic but it is impossible to be sure as yet.
Nevertheless there is almost invariably what looks like a swollen pad bearing two
fairly prominent denticles with, frequently, a few smaller denticles between them,
on the extreme anterior limit of the lateral wall (Row-I) with, posterior to this level,
a further two rows, Rows II and III, anterior to the most posterior Row-IV.

The distribution of the denticles on the ventral walls of the oesophastome appears
to be a little more valuable in delimiting species but here also some uncertainty
remains. The anterior limit of each of the ventral walls is frequently obvious in
en face views as a distinct curved rod-like ridge. This is followed more deeply, or
posteriorly, by a number of small solid denticles with, at the extreme posterior
depth of the oesophastome, a number (usually three) of large solid tooth-like struc-
tures which arise, typically, from a massive plate (Text-figs. 9, 10).

In detail the dentition in the various species I have studied is as follows.

Euchromadora vulgaris (Text-figs. 1-5).

Lateral walls. Row-I : two large denticles, no smaller ones seen; Row-II: denticles
small, row bent anteriorly (Text-fig. 2) ; Row-Ill : straight and prominent with most
dorsal denticles larger than others; Row-IV: denticles prominent, row slightly
L-shaped, largest most ventral (Text-figs. 2, 4).

Ventral walls. There are two rows of small denticles near the anterior edge,
three very prominent and massive denticles posteriorly (i.e. deep within oesophas-
tome) which arise from a distinct plate and one or two prominent denticles on the
edge of the wall where it meets the ventral extension of the lumen. These denticles
vary in appearance, possibly because of different angles of examination.

This description, based on light microscope studies, has been confirmed by electron
scan (Stereoscan) examination.

Figs i-n contd.

gaulica. 6. En face view of head. 7. Lateral oesophastomal dentition, en face
(sketch), dorsal towards top. 8. Left ventral dentition of oesophastome en face (sketch).
9-10. Inner views of ventral walls of oesophastome on one specimen. Ventral extension
of oesophastomal lumen lies between figures, n. Lateral dentition of oesophastome
viewed from inner surface, dorsal to left.

EUCHROMADORA AND SIMILAR NEMATODES

161

FIGS. i-n. 1-5. Euchromadora vulgaris. i. w face view of head. 2. Oesophas-
tomal dentition (sketch, detail). 3. Lateral view of anterior end of oesophagus, dorsal
to right. 4. Lateral dentition of oesophastome en face (sketch), dorsal towards top.
5. Left ventral dentition of oesophastome en face (sketch). 6-n. Euchromadora

162 W. G. INGLIS

Euchromadora gaulica (Text-figs. 6-n).

Lateral walls. Row-I : tiny denticles between large dorsal and ventral denticles ;
Row-II: short, straight with tiny denticles; Row-III: L-shaped with short leg
directed posteriorly and long leg transversely; Row-I V: very prominent with denticles
arising from a distinct, curved, basal flange. In addition there are a number (three
seen) of thorn-like denticles arising from the most posterior level of the wall. I am
not absolutely certain but these latter structures appear to be additional to the
more usual four rows of denticles. (Note that in Text-fig, n the ventral surface of the
body is towards the right and the drawing shows the inner surface of the wall.)

Ventral walls. The small anterior denticles are arranged in a crude spiral (Text-
figs. 8, 9. 10) while there are three massive, basal onchia arising from a very promi-
nent plate. Three illustrations of the conditions are given of which Text-fig. 8 is
taken fully en face, while Text-figs. 9 and 10 show the two ventral walls of one speci-
men. In this latter case the walls are viewed from the inner surface and are taken
from a wholly dissected specimen. Text-fig. 9 shows the left wall and Text-fig. 10
the right. It should be noted that the spiral of denticles occurs towards the lateral
surface of the body. In other words the right-hand side of Text-fig. 8 corresponds
to the median ventral extension of the lumen while the left-hand side is contiguous
with the lateral wall of the oesophastome.

Euchromadora striata (Text-figs. 12-24).

Lateral walls. Row-I: usual two prominent denticles with series of smaller
denticles between them; Row-II: curves anteriorly and all denticles are about same
size; Row-III : very slightly bowed anteriorly, denticles all about same size; Row-IV:
denticles relatively massive, few in number and arising from very prominent basal
plate.

Ventral walls. The occurrence of some variation could be established possibly
because of the relatively large size of specimens of this species. The small, anterior
denticles lie in three rows of which those of the most anterior row are consistently
smaller than those of the others. In addition a varying number of tiny denticles
can occur (or be seen) in some specimens but these are always restricted to the more
lateral region of the wall (Text-figs. 14, 20). There are two, sometimes three, large
denticles deep on the ventral wall of which two are consistently present near the
edge of the ventral lumen (Text-figs. 15, 20). This may vary because of the contrac-
tion of the mouth opening so that in some specimens the large denticles are not quite
on the edge (Text-figs. 14, 16). The large posterior onchia are very obvious in
lateral view (Text-figs. 19, 24).

Euchromadora eileenae (Text-figs. 31-37).

The dentition of the oesophastome in this species is very similar to that in E.
gaulica.

Lateral walls. All rows of denticles slope posteriorly from dorsal side (Text-fig. 34).
Row-I: two prominent denticles only; Row-II: relatively few, small denticles in an
almost straight line; Row-III: rather short and slightly bowed anteriorly; Row-IV:

EUCHROMADORA AND SIMILAR NEMATODES 163

straight, denticles prominent of which most dorsal is larger than the remainder, all
arise from a definite plate, at least in one specimen studied en face (Text-fig. 37).
Posterior to Row-IV are two, sometimes three, prominent dorsal edge of the walls.
Ventral walls. The denticles form a definite curved or spiral arrangement with
the spiral towards the lateral surface of the body. Towards the mid-ventral exten-
sion of the lumen there are two large denticles but there does not appear to be any
definite plate associated with them (Text-fig. 36) nor are they always present, or
at least have not always been seen (Text-fig. 35).

Cuticle Structure

The detailed structure of the massively punctate cuticle occurring over the anterior
region of the body in Euchromadora has been described elsewhere (Inglis, 1964) but,
because at that time I was only considering the modifications of the punctation sys-
tem, the structure of the unpunctate cuticle was not discussed nor were the lateral
plates considered. Briefly, the cuticle consists of a series of annules of dense material
embedded in a general covering of less dense material and all the structure usually
described as cuticular modification or simply as cuticular structures are represented
by these dense components.

Each annule is represented on the surface of the cuticle by a transverse striation
and the annules themselves can be extremely complicated. At the anterior end of
the body each annule can be thought of as composed of three subsiduary annules
which together form a somewhat rectangular block in longitudinal section (Text-
fig. 60). The outermost sub-annule is developed anteriorly as a thin, wedge-shaped
flange which lies over the posterior edge of the next most anterior annule and
posteriorly is curved to form a region covered by the flange from the anterior edge
of the next most posterior annule.

The innermost sub-annule is more regularly rectangular in longitudinal section
but from about the middle of the oesophagus length posteriorly it bears a lateral
peg or plate which projects anteriorly to lie under the next annule anteriorly. There
is a slight concavity on the posterior edge of this innermost sub-annule, immediately
posterior to the anteriorly directed peg, into which the anteriorly directed peg from
the next more posterior annule fits. These pegs I have called " lateral plates " and
it should be noted that they lie within the lateral chords of the hypodermis and are
below the lateral rod-like punctations when the cuticle is viewed from the surface
(Text-figs. 60-61).

In some specimens the lateral plates appear to be represented by a series of granules
of dense material but it is uncertain whether or not this is due to studying pegs in
optical section as I have not had sufficient specimens to warrant treating them with
sodium hypochlorite. It may be noted in passing that this is not the system to
which Steiner (1918) refers, as he is dealing with the true punctation rods which lie
above the lateral plates and that Wieser & Hopper (1967) appear to have confused
the two sets of structures in their discussion of Euchromadora gaulica.

The massive punctation blocks arise from the region between the outermost and
innermost sub-annules and can be pictured as a third sub-annule. These blocks
which are circular in transverse section project anteriorly on the anterior half of the

W. G. INGLIS

FIGS. 12-24. Euchromadora striata. 12. En face view of head. 13. En face view of
head of another specimen. 14. Dorsal-oblique view of inner surface of ventral oeso-
phastomal walls (sketch). 15. As fig. 14, but extremely regular condition with only two
large onchia (sketch). 16. Right ventral wall of oesophastome en face (sketch).
17. Inner surface of lateral wall of oesophastome, dorsal to left (sketch). 18. Right
ventral, wall of oesophastome en face (sketch), dorsal towards top. 19. Lateral view of

EUCHROMADORA AND SIMILAR NEMATODES 165

body to lie between the similar blocks developed on the next most anterior annule.
The blocks are further connected to both the outermost sub-annule and the inner-
most over their posterior halves as I have illustrated elsewhere (Inglis, 1964, Figs.
13, 15). The conditions described above apply to the anterior half of the body
Over the posterior half the conditions are reversed so that the outer sub-annules
are extended posteriorly instead of anteriorly and the punctation blocks and lateral
plates are also directed posteriorly instead of anteriorly.

To revert to the anterior cuticle, the large punctations disappear over two strips
both dorsally and ventrally so that a V-shaped strip of punctations projects posteriorly
along the dorsal and ventral surfaces of the body (see Coles (1965) particularly Fig. i),
until by about the posterior end of the oesophagus the dorsal and ventral cuticle is
marked only by transverse striations. Meanwhile the large elongate punctations
become slimmer and longer and are restricted to a fairly narrow zone running down
the lateral surfaces of the body (Text-figs. 60-63).

The structure of the cuticle in Euchromadora vulgaris has been studied by light
and electron microscopy by Watson (1965) but most of what she reports is based on
an examination of the dorsal or ventral region of non-punctate cuticle. This explains
the apparent discrepancy referred to by Lee (1966). In particular Watson describes
the middle layer in this non-punctate region as consisting of overlapping plates,
with which I agree, but it is the origins of these plate-like structures (a description
which only applies to the view in longitudinal section) which is particularly interesting.

The lateral surface cuticle is relatively thick and is marked by the obvious punc-
tations which correspond to the layer of rods sandwiched between the two other layers
like teeth of a comb (Text-figs. 60, 65). Dorsally and ventrally the cuticle, which
lacks such punctations, is much thinner. Each annule, in longitudinal section, is a
thin wedge-shape with the thinner edge outermost and overlying the next more
anterior or posterior annule (Text-fig. 63). The region of transition between these
two zones of modification is most interesting since it explains, among other things,
some of the features of the cuticle described and illustrated by de Man (1886).
Before describing this transition zone I should stress that I have only been able to
confirm this detail in E. vulgaris, E. gaulica and E. loricata.

The change from the lateral region with long punctations to the dorsal and ventral
region without such punctations occurs very sharply and involves (on the anterior
end of the body) the displacement of the innermost sub-annule of the dense compon-
ent of the cuticle posteriorly relative to the remainder of the annule. At about the
same level the punctation sub-annule becomes solid and it also is displaced posteri-
orly, but not to the same extent as the innermost. The combination of these displace-
ments is to produce a wide, thin wedge-shaped (in section) annule, so that although
the annules only overlap each other slightly on the lateral surface of the cuticle

Figs. 12-24 contd.

anterior end of oesophagus, dorsal to right. 20. Right ventral wall of oesophastome
en face (sketch). 21-23. En face optical sections through lateral wall of oesophastome,
dorsal towards top (sketches). 21. Denticles of Row-I. 22. Denticles of Rows-II and
III 23. Denticles of Row-IV. 24. Lateral wall of oesophastome viewed from outer
surface (sketch), dorsal to right.
ZOOL. 17, 5. I2

1 66

W. G. INGLIS

the overlap is considerable on the dorsal and ventral surfaces (Text-figs. 60-64).
As a consequence each annule of dense material when considered alone is not a
simple ring but is a narrow thick sector with punctation processes laterally and a
thin, wide, wedged sector dorsally and ventrally. This is illustrated in a very simpli-
fied way in Text-fig. 65. One consequence of this relative displacement of the

26

29

30

FIGS. 25-30. 25-27. Steineridora loricata. 25. En face view of head. 26. Lateral
wall of oesophastome viewed from outer outer surface (sketch), dorsal to left. 27. En
face detail of dentition of oesophastome (sketch). 28-30. Graphonema georgei. 28.
Lateral view of anterior end of body. 29. En face view of head. 30. Oesophastomal
dentition en face (sketch) dorsal towards top.

EUCHROMADORA AND SIMILAR NEMATODES 167

constituent sub-annules is that the analysis of the modification in whole mounts is
very difficult because one edge of one sub-annule moves posteriorly relative to the
others while the others, which can still be recognized in the specimen, remain at the
same level. Over the whole diameter of the body the anterior edge of each annule
remains at the same transverse level and appears as a straight striation running round
the body. The general appearance of this cuticle is shown in Text-fig. 64, but it should
be remembered that this illustration is taken from a much compressed specimen.

There are still some minor discrepances between this description and that given
by Watson (1965), particularly the condition shown in her Fig. 3 A in which are illus-
trated what she interprets as the " rod-like bodies " of de Man. However I suspect
that what is shown is the anterior region just posterior to the amphids in which the
cuticle is supported by rod-like punctations. This interpretation is supported by the
fact that the cuticle shown in Fig. 3A is obviously considerably thinner than that
shown in Watson : Fig. lA which in turn is, I already know, considerably thinner
than that in which the typical, massive punctation blocks or processes occur.

To complicate matters even further I suspect that the illustrations of the over-
lapping plate-like cuticle were taken from the transition zones at the anterior end
of the body so that there are structures present which are definitely not present
more posteriorly. I have, in fact, seen structures by the light microscope which
resemble the transverse rings of dense material in Layer i in Watson's Fig. 2 but
they cannot be analysed in my material. This feature deserves further study.
Finally it is perhaps worth pointing out that the direct attachment of the muscles
to the cuticle described by Watson is the feature responsible for the description of
the basal layer by several other workers as longitudinally striated or with longitudinal
fibres.

In view of these observations the genus may be rediagnosed thus :

EUCHROMADORA de Man, 1886

Chromadoridae : Cuticle: complex with large, prominent, elongate hexagonal or
ovid punctations anteriorly and posteriorly which correspond to processes developed
within cuticle from one side of each annule only; similar, but slimmer markings restric-
ted to lateral surface over middle length of body; no punctations on dorsal or ventral
surfaces over middle length of body; "lateral plates" generally well developed;
no lateral differentiation.

HEAD : large, solid dorsal onchium preceeded by denticles on anterior dorsal edge
of oesophastome ; transverse rows of small, solid denticles on lateral walls of oeso-
phastome; similar denticles anteriorly on ventral walls of oesophastome with,
frequently, larger denticles posteriorly arising from a distinct plate ; cephalic sense
organs in three circles of which the outer is composed of four setae; mouth surrounded
by twelve rugae ; amphids elongate transverse slits without marked fringes of cuticle
surrounding them; oesophagus without definite posterior bulb.

MALE: gubernaculum with prominent, generally hammer-shaped or L-shaped
lateral pieces; no pre-cloacal supplements or other modifications of cuticle; tail
relatively short and stout.

168 W. G. INGLIS

TYPE SPECIES: Chromadora vulgaris Bastian, 1865. Described above in part and
reference confirmed by de Man's (1886) and Coles' (1965) descriptions in addition to
the study of specimens.

OTHER SPECIES. The following species may be definitely referred to the genus
Euchromadora as restricted by this diagnosis because they possess or appear to
possess this combination of characters.

E. gaulica Inglis, 1962. Redescribed above in part. The gubernaculum has
been re-examined in European specimens and the presence of the two minute denticles
near the posterior end which are figured and described by Wieser and Hopper (1967)
has been confirmed.

E. striata (Eberth, 1863). Redescribed below, page 191).

E. meadi Wieser & Hopper, 1967. Retained on the basis of the original description.

E. permutabilis Wieser, 1954. Retained on the basis of the original description.

E. tokiokai Wieser, 1955. Retained on the basis of the original description.

E. eileenae sp. nov. Described below, page 193.

The disposition of other species usually referred to Euchromadora is listed on page

201.

STEINERIDORA gen. nov.

This new genus is proposed for some of those species which are generally referred
to the genus Euchromadora but which differ most obviously from the species discussed
above in possessing a distinct posterior oesophageal bulb. In this group are seven
nominal species: E. archaica Steiner & Hoeppli, 1926; E. amokurae (Ditlevsen, 1921)
(although Ditlevsen points out that his illustration tends to exaggerate the oeso-
phageal bulb) ; E. loricata (Steiner, 1916) ; E. adriatica (von Daday, 1901) ; E. dubia
Steiner, 1918; E. denticulata Cobb, 1914 and E. stateni Allgen, 1930.

E. loricata, of which I have studied specimens in detail, differs from the typical
non-bulbed Euchromadora species in several other morphological features, many of
which were reported by Steiner and Hoeppli in 1926 in describing E. archaica. The
punctations in the cuticle, which are processes exactly as in Euchromadora, remain
much the same size over the whole length of the body in contradistinction to the
condition in Euchromadora (s.s.) where they become markedly thinner and longer
over the middle length of the body. This is, I think, what Steiner and Hoeppli
mean when they describe the cuticle of E. archaica as "... rather undifferentiated
structure compared with other species; . . ." (Steiner & Hoeppli, 1926, page 571).
There are clear non-punctate regions running down the dorsal and ventral surfaces
of the body.

The structure of the oesophastome is recognizably Euchromadora-]ike but, although
the typical dorsal onchium is present preceded by a row of denticles along the
anterior dorsal edge of the oesophastome, it is a characteristic square shape (Text-figs.
25-27). The dentition of the lateral and ventral walls of the oesophastome is also
similar to that of Euchromadora species with at least three transverse rows of small
denticles on the lateral walls (Text-fig. 26). However the lateral walls also bear
very prominent, sickle-like dorsally directed cuticular onchia which arise from distinct

EUCHROMADORA AND SIMILAR NEMATODES

FIGS. 31-40. 31-37. Euchromadora eileenae. 31. En face view of head. 32. En face
view of head from another specimen. 33. Lateral view of head. 34. Lateral dentition
of oesophastome viewed from inner surface (sketch), dorsal to left. 35. Oesophastomal
dentition en face (sketch) from specimen with mouth wide open, shown in fig. 32.
36. Inner surface of left ventral wall of specimen shown in fig. 31 (sketch). Note
presence of large paired denticles and spiral arrangement of small denticles. 37. Lateral
dentition of specimen illustrated in fig. 31 en face (sketch). 38-40. Austranema alii.
38. Anterior end of body. 39. Dentition of oesophastome en face (sketch). 40. En
face view of head.

1 70 W. G. INGLIS

ridges, the edges of which bear very tiny onchia (Text-figs. 26,27). This ridge,
and associated minute denticles, represents the middle of the three lateral rows of
denticles.

Ventrally a similar pair of sickle-like onchia occur near the ventral edges of the
ventral walls and these also arise from distinct transverse blade-like ridges which
bear minute denticles along their anterior edges (Text-fig. 27). In addition on the
ventral walls there is, possibly, a row of very tiny denticles anterior to the basal
flanges of the large onchia while there is a set of three large onchia fairly deep in the
oesophastome which arise from a plate-like base, exactly as in some of the
Euchromadora species.

The same dentition is illustrated and described by Steiner and Hoeppli (1926)
for E. archaica and I have established that the same conditions occur in E. adriatica,
so far as is possible in whole mounts from the lateral aspect.

I therefore propose to refer these species to a distinct genus :

STEINERIDORA gen. nov.

Chromadoridae : Cuticle: complex with relatively stout, elongate punctations
anteriorly and posteriorly which correspond to processes developed within the cuticle
from one side of each annule only; similar and equally large, punctations continue
on lateral surfaces of cuticle only over the middle region of body; no punctations
on dorsal and ventral surfaces over much of middle body length where each annule is
wider than it is laterally; " lateral plates " poorly developed; no lateral differentiation.

HEAD : large, squarish, solid dorsal onchium ; denticles on anterior, dorsal edge of
oesophastome ; rows of small denticles on both ventral and lateral walls of oesophas-
tome and large, curved, horn-like onchia projecting into oesophastome; cephalic
sense organs in three circles of which outer is composed for four setae; mouth sur-
rounded by twelve rugae; amphids elongate, transverse slits, not bounded by
prominent fringe of cuticle; oesophagus with definite posterior bulb.

MALE: gubernaculum with prominent L-shaped lateral pieces; no pre-cloacal
supplements or other modifications of cuticle; tail relatively short and stout.
TYPE SPECIES : Spilophora loricata Steiner, 1916.

OTHER SPECIES : 5. adriatica (von Daday, 1901) ; S. archaica (Steiner & Hoeppli,
1926) ; S. (?) dubia (Steiner, 1918) sp. inq.

The position of the remainder of the seven nominal species with a posterior bulb
to the oesophagus is difficult to establish, largely because of the poverty of the origi-
nal descriptions. However, Spilophora amokurae Ditlevsen, 1921 fairly clearly has a
hollow dorsal onchium (PI. 2, fig. 4 in Ditlevsen, 1921) and has a very extensive
unmarked region of cuticle at the anterior end of the body (see further discussion
below, page 182) ; Euchromadora dubia Steiner, 1918 is insufficiently described for
any certain opinion to be reached but appears to be referable to Steineridora although
only as a species inquirendum; E. denticulata Cobb, 1914 is poorly described but
because it appears to have a hollow or hollow-type dorsal onchium and the inter-
mediate circle of cephalic sense organs is setose it is not referable to either

EUCHROMADORA AND SIMILAR NEMATODES 171

Euchromadora or Steineridora (see below, page 188) ; E. stateni Allgen, 1930 is treated
as a synonym of E. denticulata by Wieser (1954) but is better treated as a species
dubia as suggested by Coles (1965). Nevertheless it should be noted that the
illustration of the head shows that E. stateni is not referable to Steineridora nor to
Euchromadora since it has a hollow-type anteriorly directed dorsal onchium, an
intermediate circle of six setae and an extensive non-punctate region at the anterior
end of the body.

PARAPINNANEMA gen. nov.

In a new species from Western Australia, which is later named Parapinnanema
wilsoni, the cuticle is thin and marked by a series of fine annules, in which there are
fine elongate punctations, for about one head diameter posterior to the anterior end
of the body. Posterior to this level the cuticle thickens very rapidly and is very
prominently marked by obvious " battlement-like " punctations (Text-figs. 66, 97).
The cuticle becomes thinner, rather slowly, over the posterior quarter of the length
of the oesophagus and the markings become smaller and less pronounced but are
still obviously battlement-like. Such a region of thick cuticle is a characteristic
feature of most of the species in which the cuticle is battlement-like, although never
as strongly as in P. wilsoni where it is one of the most startlingly obvious features
of the species even under a low-power stereoscopic microscope (Text-fig. 97).

It should be noted that it is possible to get the impression that the cuticle is
modified by a series of rods and dots over some areas but this depends upon the
focusing and the extent to which the specimen being studied is under pressure. An
additional complication in interpreting the structure of the cuticle from whole
specimens is that, as in Euchromadora, each annule of the dense component of the
cuticle consists of three or two sub-annules which are generally skewed so that one
never lies immediately above another. As a result it is easy to interpret the condi-
tions as representing two inner annules to each outer annule. This never appears
to be true when the annules are separated by using sodium hypochlorite.

The punctation markings occur over the whole surface of the body, both dorsally
and ventrally as well as laterally and the battlement-like markings become more
pronounced again over the region just anterior to the cloacal opening or the anus
and continue like this over the tail. The annulation is wide and the transverse
striations on the surface of the cuticle are deep and pronounced (Text-figs. 73-75).

In detail the cuticle resembles Euchromadora in that each annule consists of two,
or three (depending on interpretation) sub-annules of which the outermost is curved
anteriorly (on the anterior half of the body) to form a strip overlying the posterior
edge of the next immediately anterior annule. The innermost is similar to that in
Euchromadora, consisting of a solid sub-annule which is slightly rounded along both
the anterior and posterior edges. The major difference between the Euchromadora
and Parapinnanema cuticles is that in the latter there are processes developed on
both sides of the annules, on the middle of the body-length, about the middle of the
thickness of the cuticle (Text-fig. 72), instead of on only one side as in Euchromadora,
and there do not appear to be clear channels passing completely through the middle
sub-annule.

172

W. G. INGLIS

41

FIGS. 41-50. 41-43. Austranema colesi. 41. En face view of head. 42. Oesophastomal
dentition en face (sketch). 43. Lateral view of anterior end of oesophagus, dorsal to
right. 44-46. Austranema shirleyae. 44. Ventral view of oesophastome showing
ventral onchia and lateral flanges in optical section (sketch). 45. Oesophastomal
dentition en face (sketch). 46. Lateral view of anterior end of oesophagus, dorsal to
right. 47-50. Protochromadora scampae. 47. Oesophastomal dentition en face (sketch)
48. Detail of dorsal onchium (sketch) 49. En face view of head. 50. Lateral
view of anterior end of body.

EUCHROMADORA AND SIMILAR NEMATODES 173

This modification is apparent in a different way in the thick cuticle at the anterior
end of the body where the battlement-like markings are very obvious. Here when
the cuticle begins to become very thick the anterior edge of the outer annule becomes
notched so that it looks like battlements. Along the posterior edge of the annule
posteriorly directed peg-like processes are present which arise from the middle level
of the annule (Text-fig. 67). In other words the annules are articulated together
by a system derived from both the anterior and the posterior edges (see also Fig.
5ob in Filipjev, 1918 and Fig. 52c).

As the cuticle becomes thinner posteriorly the dorso- and ventro-lateral articula-
ting plates (see below) appear about halfway along the length of the oesophagus
and the pegs on the posterior edges of the annules move relatively nearer the surface
of the cuticle until the modfication is restricted to the outer surface of the cuticle.
Meanwhile, as the middle pegs move outwards a series of grooves develop in the
innermost sub-annule of the cuticle so that there are, in surface view, a series of
large battlement-like modifications on the outer level and a series of very fine, short,
elongate punctations on the innermost level of the cuticle (Text-figs. 68-71).

Posterior to the posterior end of the oesophagus the battlement-like modifications
of the outer surface become smaller, and less distinct, until they almost disappear
and are indistinct over roughly the middle third of the body length. At the same
distance from the anterior end the fine striations of the inner sub-annule move
relatively nearer the middle level of the cuticle as a series of anteriorly and posteriorly
directed pegs (Text-fig. 72) .

On passing further posteriorly the conditions are reversed; the middle layer of
punctation pegs moves inwards to form fine elongate grooves on the inner sub-annules
and the battlement-like indentations reappear on the outer surface and those along
the anterior edge of each annule move inwards, relative to the surface of the cuticle, to
form a series of pegs arising from the middle depth of each annule. In other words
the conditions are reversed relative to those on the anterior part of the body.

At the level on the length of the body where the punctations are wholly due to
the innermost modifications the annules are very skewed (Text-fig. 72) so that on
focusing from the surface there appear to be two inner annules to each outer annule,
and this condition reverses rather sharply about the middle of the body length.
Over this same region of the body there is a distinct straight striation of the surface
of the cuticle which runs round the middle of each annule and the annules are
delimited anteriorly and posteriorly by wavy lines corresponding to the suppressed
battlement-like processes (Text-fig. 75) .

Therefore in contrast to the condition in Euchromadora there are two systems of
punctation rods involved and the annules never widen dorsally and ventrally as in
Euchromadora.

In addition to all this modification there are interlocking plates along the dorso-
and ventro-lateral aspects of the body which are due to the development of inter-
locking extensions from the innermost level of the cuticle and are covered externally
by the elongate punctations found on the rest of the cuticle (Text-fig. 68). The
cuticle has no clear zones, without punctations, dorsally and ventrally.

The dorsal onchium is hollow (Text-figs. 53, 55) although in some specimens it is

W. G. INGLIS

FIGS. 51-59. 51-55. Parapinnanema wilsoni. 51. En face view of head. 52. Oeso-
phastomal dentition en face (sketch). 53. Lateral view of head. 54. Dorsal view of
oesophastome showing lateral flanges in optical section (sketch). 55. Detail of dorsal
onchium (sketch). 5659. Hypodontolaimus slacksmithi. 56. En face view of head.
57. Dorsal view of head. 58. Oesophastomal dentition en face (sketch). 59. Lateral
view of head.

EUCHROMADORA AND SIMILAR NEMATODES 175

difficult to reach a decision. However, in addition to the structure of the cuticle
and the form of the dorsal onchium, the specimens are characterized by no denticles
along the anterior dorsal edge of the oesophastome, the intermediate circle of cephalic
sense organs is setose, there are three circles of sense organs, the lateral walls of the
oesophastome bear cuticular flanges which are not divided to form anteriorly directed
denticles along their anterior edges so that each flange stands out in en face view as a
simple, blade-like structure (Text-figs. 51-54). Ventrally there are two anteriorly
directed cone-like denticles while in some specimens there appears to be a single
denticle on the lateral walls of the oesophastome anterior to the lateral flanges
(Text-figs. 51-53).

The tail is long and thin, the lateral pieces of the gubernaculum are L-shaped
and there is a mid-ventral, thickened, pre-cloacal modification of the cuticle on the
males as in E. colesi and N. alii (see Inglis, 1968 and Murphy, 1965).

The generic name Odontocricus Steiner, 1918 is already available for specimens
with this massively battlemented type of cuticle. This name was proposed for a
new species which was considered sufficiently distinct to warrant reference to a
distinct subgenus of Euchromadora, E. (0.) hupferi Steiner, 1918. The remarkably
obvious battlement-like structure of the cuticle is illustrated by Steiner (1918 : Fig.
4) but the species description is based on a juvenile female specimen and it is uncertain
that it could ever be recognized again. Equally it is not possible to forecast the
continuing uniqueness of the strongly battlemented cuticle so that rather than use a
generic name of which the type species is a species dubia I propose to erect a new
genus for the Australian specimens:

PARAPINNANEMA gen. nov.

Chromadoridae : CUTICLE: without markings on anterior end until posterior to
prominent amphids where a series of rods and dots appear; posterior to this level
very prominent battlement-like appearance due to presence of processes within
cuticle on both anterior and posterior edges of annulations ; posterior to oesophagus,
over most of body length, punctations slim and elongate but due to second inner
layer, most outer layer now reduced to rather wavy lines on outer surface of cuticle ;
battlement-like cuticle reappears over posterior end of body from slightly anterior
to anus or cloacal opening; annulation of cuticle very pronounced and wide; cuticle
very thick over roughly length of oesophagus but region just posterior to head thin;
no clear zones dorsally and ventrally and annules same width all round body; four
files of articulating plates dorso- and ventrolaterally.

HEAD : slightly set-off by appearance of thin covering cuticle relative to conditions
more posteriorly; dorsal onchium hollow; no denticles on anterior dorsal edge of
oesophastome; flanges on lateral walls of oesophastome, without dentate anterior
edges; one, solid, cone-like anteriorly directed onchium on each ventral sector at
base of oesophastome; cephalic sense organs in three circles of which intermediate
consists of six setae and outer of four setae; mouth surrounded by twelve rugae;
amphids prominent transverse elongate slits bordered by prominent fringes of
cuticle ; oesophagus without prominent posterior bulb.

1 7 6

W. G. INGLIS

62

63

66

FIGS. 60-72. Structure of cuticle. 60-65. Euchromadora vulgaris. 60-63. Successive
longitudinal sections through one annule from wholly laterally to wholly ventrally or
dorsally showing relative displacement of subannules. 64. Surface view of two annules
showing relative displacement and widening of inner subannule. 65. Reconstruction
of one annule about middle of body. 66-72. Parapinnanema wilsoni. 66. Anterior

EUCHROMADORA AND SIMILAR NEMATODES 177

MALE: gubernaculum with L-shaped lateral pieces; elongate, narrow, thickened,
mid- ventral pre-cloacal cuticular modification ; tail long and slim.

TYPE SPECIES : Parapinnanema wilsoni sp. nov.

PROTOCHROMADORA gen. nov.

Euchromadora scampae Coles, 1965 is characterized by an intermediate circle of
six non-setase cephalic sense organs and four setae in the outer circle. The dorsal
onchium is apparently hollow in some specimens and apparently solid in others
(Text-figs. 48, 50). However the oesophastome has no denticles along the anterior
dorsal edge, there are two ventral cone-like onchia and there is no trace of any lateral
oesophastomal modification of any kind, neither in the forms of transverse rows of
denticles nor as lateral flanges (Text-fig. 47).

The cuticle just posterior to the outer circle of cephalic sense organs is marked by
tiny, slightly elongate, punctations which become more prominent posteriorly.
They are large oval punctations over the anterior half of the length of the oesophagus
which lie directly within the annulations. In other words, on focusing down through
the annules the punctations lie directly over the inner sub-annules and wholly below
the outer sub-annules so that there is no appearance of movement.

The punctation blocks are not, however, simple in structure since on focusing
through them, from the surface, they become somewhat dumb-bell-shaped in section
and finally at the deepest level are battlement-like in appearance (Text-figs. 76-79).
The punctation blocks become slimmer on studying them more posteriorly and
become much less prominent, very slim and close together. At the same time they
begin to project from the anterior edge of each annule. As this projection becomes
more pronounced notches appear in the posterior edges of each annulation to accept
the processes projecting from the next more posterior annule (Text-figs. 77, 78).
These notches on the posterior edge become deeper and more pronounced until the
annules appear to be battlemented on both their edges (Text-fig. 79) with a distinct
transverse striation running between them. I am not certain at which level these
punctations lie within the cuticle but those on the anterior end appear due to processes
in the middle depth while those on the middle of the body appear to be due to
indentations on the extreme outer subannule of the cuticle (Text-fig. 79).

The conditions are reversed over the posterior half of the body.

Tiny lateral plates are present starting on the anterior end of the body about the

FIGS. 60-72 contd.

end of body, surface view. Black region is outer surface, white processes lie inwards.
67. Reconstruction of anterior annule. 68. Dorso- and ventro-lateral articulating
processes. 69-72. Annules in longitudinal section, anterior to left. 69. Thick
cuticle over anterior region of oesophagus. 70. Middle of oesophagus length, posterior
process displaced outwards. 71. Posterior end of oesophagus, posterior process now on
outer surface, anterior processes replaced by grooves on anterior edge. 72. One-third
of body length from anterior end. Note distortion of annules with dominance of inner
system of processes; outer processes reduced to irregularities on outer edges of annules.

i 7 8

W. G. INGLIS

Win) QQ

Q Q Q

77

1 1 ' i "i 11 1 1 ii

76

78

&JM)&*?*teJ^^

75

79

80

84

81

82

muuiiiii

83

89 90

FIGS. 73-90. 73-75. Parapinnanema wilsoni: surface appearance of cuticle. Anterior
end towards top. 73. Just posterior to amphids. 74. Over oesophagus length.
75. On most of body posterior to oesophagus. 76-79. Protochromadora scampae:
cuticle. 76. Punctation just posterior to amphids. From left to right: surface, about
middle level, deep view (see description, page 177). 77. Just posterior to amphids.
78. Over first quarter of oesophagus. 79. Over most of body, plan on left, longitudinal
section on right. 80-84. Gvaphonema georgei: surface appearance of cuticle. 80. Just
posterior to amphids. 81. Three annules posterior to 80. 82. First third of oesophagus

EUCHROMADORA AND SIMILAR NEMATODES 179

level of the middle of the length of the oesophagus but over the middle region of the
body they are represented by a series of granulations lying between the innermost
parts of the dense annules of the cuticle.

The cuticle is marked by punctations laterally, but is clear dorsally and ventrally
over the middle length of the body. In the male there is no precloacal mid-ventral
modification of the cuticle and the tail is relatively short and stout.

This species is obviously distinct from those grouped in the genera Euchromadora
and Steineridora and I propose to refer it to a new genus the validity of which will
become clearer below.

PROTOCHROMADORA gen. nov.

Chromadoridae : : CUTICLE: complex with large, prominent, elongate punctations
anteriorly and posteriorly which appear to correspond to processes developed on
one side of each annule only; similar, but slimmer, punctations restricted to lateral
surfaces of body over much of middle length; cuticle without punctations dorsally
and ventrally over similar middle length of body; " lateral plates " represented by
granules.

HEAD : dorsal onchium possibly solid but impossible to classify rigidly; no denticles
on anterior dorsal edge of oesophastome ; no denticles or flanges or any other struc-
tures on lateral walls of oesophastome ; one cone-like, anteriorly directed onchium on
each ventral sector at base of oesophastome; cephalic sense organs in three circles
of which outer consists of four setae; mouth surrounded by twelve rugae; amphids
elongate, transverse slits, not bordered by prominent fringes of cuticle; oesophagus
without prominent posterior bulb.

MALE : gubernaculum with L-shaped lateral pieces ; no pre-cloacal supplements or
other modification of cuticle; tail relatively short and stout.

TYPE SPECIES: Euchromadora scampae Coles, 1965.

OTHER SPECIES: P. mediterranea (Allgen, 1942); P. parafricana (Gerlach, 1958).

The reference of E. mediterranea Allgen 1942 to Protochromadora is based on a
study of some specimens in a rather poor condition. Nevertheless it is possible to
confirm their specific identity and to establish with a fair degree of certainty their
similarities to P. scampae.

The inclusion of Euchromadora parafricana Gerlach, 1958 in Protochromadora is
slightly less certain as it is based on the published description. Gerlach (1958) gives

FIGS. 73-90 contd.

to first third of body length. 83. Remainder of body. 84. Longitudinal section,
about level of figure 82. Anterior to left. 85-88. Austranema shirleyae: cuticle.
85. Surface appearance, anterior end of body. 86. Detail at same level as 85. 87. Body
posterior to oesophagus. 88. Transverse section at same level as 87 showing origins
of punctation processes. 89. Longitudinal section at anterior end. Note marked
anterior displacement of outer subannule. 90. Longitudinal section about middle
of body (anterior to right in 89 and 90).

180 W. G. INGLIS

two figures of the head, one of which shows a hollow dorsal onchium (Gerlach, Fig.
4a) while the other (4b) shows a possibly hollow or solid dorsal onchium, reflecting
the difficulty of distinguishing between the two forms. In addition an intermediate
and an outer circle of setae is shown in Fig. 4b while only an outer circle of four setae
is shown in Fig. 4a. The tail is long and thin, there are no pre-cloacal modifications
or supplements and the oesophagus does not end in a bulb. The illustrations of
the cuticle suggest the condition in P. scampae and I refer E. parafricana to
Protochromadora with little hesitation.

GRAPHONEMA Cobb, 1898

Graphonema Cobb, 1898 is, in some ways, the most distinctive genus which remains
to be considered and was erected by Cobb (1898) for G. vulgaris Cobb, 1898 with a
reference to a further forthcoming species, G. pachyderma, which was never described.
G. vulgaris was based on specimens collected from algae and sea sand on the coasts
of New South Wales and Victoria, Australia. In describing it Cobb specifically
draws attention to the following points: " The conoid neck terminates in a truncate
head, conspicuous because of a distinct constriction behind it, and because of the
sudden diminution in size on it of the cuticular markings . . ."; " The six cephalic
setae opposite the apex of the dorsal tooth, and the four sub-cephalic setae . . .".
In addition he refers to a minute dorsal tooth ; linear, arcuate spicules of equal width
throughout their lengths; a gubernaculum with lateral pieces; no oesophageal bulb
and no pre-cloacal supplements.

I have collected specimens from the west coast of Australia, described below,
which conform to this description with very great accuracy in all respects, although
they appear to represent a different species from that described by Cobb (1898).
As a consequence it is possible to reassess the status and diagnosis of the genus
Graphonema.

The genus is easily recognized because the head is, as reported by Cobb, distinctly
set-off as a swollen, almost globular, region on which there are very fine dot-like
punctations (Text-fig. 28). More posteriorly the cuticle becomes thicker over the
region of the oesophagus but always remains relatively thin and delicate in appear-
ance when compared to the conditions in Euchromadora and the other genera consid-
ered here. Nevertheless the annulation of the cuticle is fairly marked and obvious.
The punctations at the anterior end of the body, just posterior to the swollen region,
are of a definite battlement-like form although the battlement processes are small
relative to the width of the annules from which they arise (Text-fig. 80). On
passing posteriorly the processes rapidly increase in relative size and notches develop
in the posterior edges of the annules of dense material (Text-fig. 81). By the level
of the posterior end of the first third of the length of the oesophagus the annules bear
a distinct system of double battlement-like processes. The processes on the anterior
edge are longer and nearer the surface of the cuticle than those on the posterior
edge (Text-figs. 82, 84). Over the whole of this region the annules of dense
cuticular material are roughly rectangular in longitudinal section with slight flanges
developed along their anterior edges (Text-fig. 84).

EUCHROMADORA AND SIMILAR NEMATODES 181

The double-battlement markings on the cuticle over the posterior two-thirds of the
oesophagus continue over the region posterior to the oesophagus virtually unchanged
until about one-third of the total body length from the anterior end of the body where
the battlement-like processes on the posterior edge of the annules move relatively
nearer the surface of the cuticle, the cuticle becomes thinner and the punctations
now look like long thin blocks (Text-fig. 83).

There are four clear zones running down the cuticle on the dorso- and ventro-
lateral aspects so that the cuticle is marked by punctations dorsally and ventrally
along the whole length of the body. These punctations are thinner and less obvious
than those occurring on the lateral surface of the body. In the males there is no
mid-ventral pre-cloacal modification of the cuticle and the gubernaculum is small
and slightly L-shaped.

The mouth opening is surrounded by the usual twelve rugae and the cephalic
sense organs are in three distinct circles of which the members of the outermost two
circles are setose (Text-fig. 29). The amphids are very prominent, transverse slits
bordered by obvious thin fringes of cuticle which stand out from the general surface
of the cuticle. The circles of cephalic setae are far apart and the members of the
outer circle lie on the same level as the amphids which are themselves positioned
relatively far posteriorly from the anterior end of the body.

The dorsal onchium is small, hollow, directed anteriorly and there are no denticles
developed on the anterior dorsal edge of the oesophastome. Each of the lateral
walls of the oesophastome bears a flange which projects anteriorly most markedly
towards its dorsal edge so that it resembles an anteriorly directed onchium when the
level of focus is high. However on focusing down through the head, in an en face
preparation, the flange appears progressively. There are no distinct separate
denticles or processes developed from the effectively continuous edge of the flange
(Text-fig. 30).

Each ventral sector bears one fairly prominent cone-like onchium. There is
another less prominent pair, each member of which arises from, or near to, the ventro-
lateral corners of the oesophastomal funnel. These structures are definitely present
(Text-fig. 30) and there is a possibility that each ends anteriorly as a double point.
This cannot be established with certainty but when this head is studied en face
the dentition of the oesophastome is immediately obvious as seven bright points of
light (Text-fig. 29) of which one is large and dorsal (the dorsal, anteriorly pointing
onchium), two fairly large are lateral in position (representing the anterior edges of
the lateral flanges), and four occur in an almost straight line across the ventral wall
of the oesophastome (the onchia on the ventral sectors and those arising from the
ventral corners of the oesophastome). With more careful focusing and studying
the less certain second points to the ventro-lateral onchia can be made out.

The oesophagus expands evenly posteriorly but there is no distinct posterior bulb.
The tail is long and slim (Text-fig. 99).

These features conform so exactly to Cobb's verbal picture of G. vulgaris that there
can be little doubt that we are dealing with a species which is at least congeneric.
As a consequence it is possible to reassess the genus which is certainly distinct.

The discovery of unequivocable specimens of Graphonema has further taxonomic

ZOOL. 17, 5. I3

182 W. G. INGLIS

consequence since Wieser (1954) "... thinks I am able to revive Cobb's doubtful
genus ..." and refers four species to that genus while in 1959 he adds another
four species of which two are new.

As a result the genus currently contains the following nine species in addition to
the type species: Chromadora sabangensis Steiner, 1915; Spilophora amokuroides
Allgen, 1927 (redescribed by Wieser, 1954) ; Spilophora norvegica Allgen, 1932 ;
Chromadora paraheterophya Allgen, 1932; with Spilophora pusilla Allgen, 1947;
Chromadora suilla Allgen, 1947 and Chromadora spectabilis Allgen, 1932 referred to
the synonymy of these species (Wieser, 1954); in 1959 Wieser adds the following
species: G. flaccida Wieser, 1959; G. divosa Wieser, 1959; Chromadora tentabunda
de Man, 1890; Chromadorita crassa Timm, 1952 and Chromadorita chitwoodi Wieser,

1954-

However of these species Chromadora tentabunda, which is redescribed by Gerlach

(1951), has a simple gubernaculum without L-shaped lateral pieces, a post-oesophageal
bulb, a prominent hollow dorsal onchium with a well developed dorsal expansion
of the oesophagus associated with it, the head is not set off and the intermediate
circle of cephalic sense organs is not setose. Timm's (1952) Chromadorita crassa, a
new name for C. tentabunda of Chitwood (1951), is I agree with Wieser (1959) almost
certainly de Man's species.

Both G. flaccida and G. clivosa have a similar combination of characters ; the dorsal
onchium is large and hollow, the intermediate cephalic sense organs are not setose,
there is a posterior oesophageal bulb, the gubernaculum lacks lateral pieces and the
heads are not set off.

Spilophora amokuroides Allgen, 1927 is redescribed by Wieser (1954) and referred
to the genus Graphonema with S. pusilla Allgen, 1947 and Chromadora suilla Allgen,
1947 as synonyms. As Wieser (1954) points out, Allgen's descriptions of 5. amoku-
roides and 5. pusilla are " insufficient " and he later (Wieser, 1959) adds that the
use of the name 5. amokuroides for his redescription is "... hypothetical, since
Allgen's inexact figures and descriptions could very well fit any species of this group."
I concur and propose to rename Graphonema amokuroides of Wieser, 1954 and will
treat all four of Allgen's species as species dubia.

Chromadora sabangensis Steiner, 1915 is referred to Graphonema by Wieser (1954)
but is originally described from females which makes discussion difficult. The
intermediate circle of sense organs is not setose, the dorsal onchium is small and
hollow, the punctations on the anterior end of the body are very light or lacking,
the punctations on the general surface are thin and elongate and the posterior end
of the oesophagus does not form a distinct bulb. It is, at best, a species inquirendum.

Finally Spilophora amokurae Ditlevsen, 1921 as redescribed by Wieser (1954) under
the name Euchromadora amokurae and according to the original description, has a
clear zone of cuticle anteriorly, the dorsal onchium is hollow, the intermediate circle
of cephalic sense organs appears to be setose, there is no report of any pre-cloacal
modification of the cuticle on the ventral surface of the male, the lateral pieces of
the gubernaculum are L-shaped, and the tail is long and slim. In many of these
respects the species differs from those referred to Euchromadora and, although some
doubt remains, it appears most probably referable to Graphonema as re-diagnosed.

EUCHROMADORA AND SIMILAR NEMATODES 183

The redescription of a Graphonema species, sensu Cobb, means that the species
referred to that genus by Wieser are displaced and I propose to refer them to a
new genus, Innocuonema. This new genus and Graphonema may be differentially
diagnosed thus :

GRAPHONEMA Cobb, 1898

Chromadoridae : CUTICLE : anterior end of body swollen, bulb-like without puncta-
tions anteriorly and with tiny dot-like punctations posteriorly ; anterior punctations,
just posterior to bulb region, prominent and battlement-like ; more posteriorly puncta-
tions long and slim; punctations on both dorsal and ventral surfaces of body;
dorso- and ventro-lateral strips of articulating processes.

HEAD: markedly set off as swollen, bulb-like region; dorsal onchium hollow;
no denticles on anterior dorsal edge of oesophastome ; flanges on lateral walls of
oesophastome, without dentate anterior edges; two prominent, cone-like anteriorly
directed onchia on each ventral sector at base of oesophastome ; cephalic sense organs
in three circles of which intermediate and outer are far apart and composed of setae ;
mouth surrounded by twelve rugae ; amphids prominent transverse slits bounded by
prominent fringes of cuticle ; oesophagus without prominent posterior bulb ;

MALE: gubernaculum with poorly developed L-shaped lateral pieces; no pre-
cloacal supplements or other modification of cuticle; tail long and slim.

TYPE SPECIES: Graphonema vulgaris Cobb, 1898.

OTHER SPECIES: G. georgei sp. nov. ; G. amokurae (Ditlevsen, 1921).

INNOCUONEMA gen. nov.

Chromadoridae: CUTICLE: punctations complex and obvious; no lateral differen-
tiation; no other information available.

HEAD : head not set off as swollen region ; dorsal onchium fairly obvious and hollow,
with associated dorsal swelling of oesophagus; structure of oesophastome unknown;
only outer circle of four sense organs setose ; amphid not prominent ; definite posterior
oesophageal bulb.

MALE: gubernaculum without prominent lateral pieces; no pre-cloacal supple-
ments or other pre-cloacal modifications ; tail long and slim.

TYPE SPECIES: Graphonema flaccida Wieser, 1959.

OTHER SPECIES : /. clovosa (Wieser, 1959) ; /. tentabunda (de Man, 1890) ; I. chilensis
nom nov pro Graphonema amokuroides of Wieser, 1954 non Spilophora amokuroides
Allgen, 1927.

The following species are considered dubia: Chromadora suilla Allgen, 1947;
C. paraheterophya Allgen, 1932; Spilophora norvegica Allgen, 1932; S. amokuroides
Allgen, 1927; 5. pusilla Allgen, 1947 and Chromadora spectabilis Allgen, 1932.

184 W. G. INGLIS

NYGMATONCHUS Cobb in Cobb, 1933

Before considering the remaining Euchromadora-type species I have seen it is
necessary to consider the delimitation and diagnosis of the genus Nygmatonchus
Cobb in Cobb, 1933. This genus was erected with one contained species, N. scriptus
Cobb in Cobb, 1933, which was described, without illustrations, by reference to the
genus Rhips Cobb, 1920. Using a combination of the verbal description of N.
scriptus and the illustrations of Rhips ornata Cobb, 1920 it is possible to establish
the following characteristic features for Nygmatonchus. The dorsal onchium is
small, hollow and anteriorly directed, the amphid is prominent with a double contour,
the cuticle is complex with lateral differentiation and basket-work markings anteri-
orly but only dots about the middle of the body, the cephalic sense organs of the
intermediate and outer circles are setose and lie at the same level, the oesophagus
lacks a definite posterior bulb, the tail is long and slim, and there is possibly a region
of raised pre-cloacal modification on the males (not mentioned by later authors).

Wieser (1954) describes a new species N. fossiferus but delimits Nygmatonchus as
" Distinguished from Neochromadora by having ten instead of the usual four cephalic
setae". As the descriptions in this paper show, such a condition obtains in a wide
range of forms. Nevertheless Wieser's description agrees with that given by Cobb
in Cobb (1933) fairly well although the amphid is not obviously like that of Rhips,
and the gubernaculum is not obviously L-shaped as in Rhips, a condition suggested
by Cobb in Cobb's description.

Andrassy (1959) describes what he considers to be a female of Nygmatonchus
scriptus and although I consider it unestablished that it is referable to that species
it is, nevertheless, recognizably the same genus. Here, also, the dorsal onchium is
small and hollow, the intermediate and outer circles of setae occur at the same level,
the amphid is prominent with a double contour, there is definite lateral differentiation
of the cuticle, and the tail is long and slim. Most recently Gerlach (1967) describes
a new species N. minutus, in which the cephalic setae are in two circles but which
otherwise agrees with the diagnosis of the genus.

These characters may be summarized in a diagnosis as:

NYGMATONCHUS Cobb in Cobb, 1933

Chromadoridae : cuticle complex anteriorly, elongate punctations posteriorly on
middle of body; distinct lateral differentiation; intermediate and outer circles of
cephalic sense organs on one or two levels, setose; amphid prominent with double
contour; oesophagus without definite posterior bulb; dorsal onchium small and
hollow ; tail long and slim.

MALE : pre-cloacal modification on male (?) ; gubernaculum not L-shaped.

TYPE SPECIES: Nygmatonchus scriptus Cobb in Cobb, 1933.

OTHER SPECIES: AT", fossiferus Wieser, 1954; N. bicornata (Wieser, 1959) comb,
nov. ; N. minutus Gerlach, 1967.

EUCHROMADORA AND SIMILAR NEMATODES 185

The reference of the three latter species to the genus is slightly doubtful. Although
the first appears to be a true member of the genus the illustration of the amphid
given by Wieser (1954) does not match that described by Cobb since it is short and
rather indistinct (compare Wieser's Fig. I4ia of N. fossiferus with Fig. I42a of
Actinonema pachydermatum Cobb, 1920 a much more Rhyps-like amphid), and Wieser,
as Murphy (1965) points out, does not mention any pre-cloacal modification of the
cuticle.

Neochromadora bicornata was described by Wieser (1959) from females only so that
it is difficult to refer it to any genus with certainty, but here again the amphid is
small and indistinct, and the cephalic sense organs of the intermediate and outer
circles are setose. Nevertheless the structure of the cuticle appears to match that
of Nygmatonchus, particularly when Andrassy's (1959) and Gerlach's (1967) illus-
trations are considered, and I will refer Wieser's species to Nygmatonchus as a
provisional measure, at least until males have been described.

Murphy (1965) describes a new species Nygmatonchus alii of which he says " My
species conforms so closely to Cobb's generic description that there is little room for
doubt as to generic placement". (Murphy, 1965; page 208.) However, the dis-
covery of several species with roughly the same combination of characters as N.
alii raises difficulties since if Wieser's interpretation of Nygmatonchus is correct and
his illustration of the gubernaculum accurate the genus is characterized, among other
features, by a small Neochromadora-type gubernaculum while if Murphy is correct
the gubernaculum is Euchromadora-like.

However Cobb in Cobb's (1933) description of N. scriptus does not say that the
lateral pieces of the gubernaculum (telamons) resembles those of Rhips (which are
L-shaped) which one might have expected in view of the other direct reference to
that genus. I, therefore, continue to accept Wieser's (1954) interpretation of the
genus and cannot accept that Murphy's species is referable to it, particularly as the
lateral differentiation he illustrates is not lateral and does not suggest " wings " as
mentioned by Cobb in Cobb (see below, page 200).

AUSTRANEMA gen. nov.

The remaining Euchromadora-like species I have studied are all characterized by
the presence of flanges, developed from the lateral walls of the oesophastome, with
dentate anterior edges ; the cephalic setae are in one circle of ten and lie anterior to
the level of the amphids ; the amphids are wide and prominent with cuticular fringes ;
the dorsal onchium is generally hollow; the cuticle is thick over the oesophageal
region of the body; the lateral pieces of the gubernaculum are L-shaped; the tail is
long and slim ; there is a region of pre-cloacal modification on the mid- ventral surface
of the body in the form of a raised strip of cuticle ; the cuticle is marked by puncta-
tions dorsally and ventrally; the posterior end of the oesophagus swells evenly but
there is no distinct posterior bulb; there are no denticles along the dorsal anterior
edge of the oesophastome and there are anteriorly directed, cone-like onchia ventrally
within the oesophastome.

ZOOL. 17, 5. i3

i86 W. G. INGLIS

Nygmatonchus alii Murphy, 1965 falls neatly into this group and the only apparent
discrepancies are that Murphy does not report any dentition on the anterior edge of
the lateral flanges of the oesophastome, although he illustrates the presence of such
flanges in optical section (Murphy, 1965, Fig. I, D) and describes a definite lateral
differentiation of the cuticle. However, I have a few specimens from Western
Australia which appear to be indistinguishable from those described by Murphy
and can establish that the lateral oesophastomal flanges are dentate and that there
is no lateral differentiation. I suspect that Murphy has, instead, illustrated the
dorso- and ventro-lateral files of articulating plates common on this kind of nematode.
Similarly a re-examination of the original specimens of Euchromadora shirleyae
Coles, 1965 and E. colesi Inglis, 1968 shows them to have this combination of charac-
ters and, to that extent, cannot remain as members of the genus Euchromadora
(The description of the cephalic sense organs of E. colesi in Inglis (1968) is wrong.)
It may be stressed again that when the mouth opening is closed and the oesophastome
somewhat collapsed, the dorsal onchium folds posteriorly upon itself and gives every
appearance in lateral view of being solid. However, when the mouth is open and the
dorsal onchium fully extended it is clearly hollow.

I have been able to study the cuticle in some detail in E. colesi, E. shirleyae and
N. alii, and it is structurally the same in all three. The sole difference of note is that
the punctation blocks are much larger and the cuticle is thicker in E. shirleyae
than in N. alii and E. colesi. As a result most of the following description is based
on E. shirleyae. The structure of the cuticle in these three species differs from that
in all the other species described in this paper in that each annule of dense material,
when studied in transverse section, is markedly skewed, even at the extreme anterior
end of the body so that the outer sub-annule lies fairly far anterior to the innermost
(Text-fig. 89) . This condition continues over the whole length of the body with the
conditions reversed on the posterior half, i.e. the outer sub-annule is then posterior
relative to the innermost.

The cuticle for about one head diameter posterior to the anterior end is marked by
simple, dot-like punctations and then elongate punctations appear which are, as
usual, due to the presence of anteriorly directed processes lying in the middle level
of the cuticle. Initially these punctation-blocks are restricted to the anterior edge
of each annule to give the appearance of a battlement-like cuticle (Text-figs. 85, 86)
but after only two or three annules the punctations become elongate rectangles when
viewed from the surface (Text-fig. 87). When single annules are studied, however,
the punctations do not lie as strips in one plane but appear to be higher anteriorly
than they are posteriorly. This appears to be caused by the anterior half (approxi-
mately) of each punctation being produced by the anteriorly directed process while
the posterior half is due to grooves in the innermost, relatively posterior, sub-annule
(Text-fig. 89). The processes are, as in Euchromadora, circular in transverse section
and, although completely free nearer their outer ends, attached to the surface sub-
annule by a thin strip of cuticle while more posteriorly a second similar strip attaches
them to the innermost sub-annule (Text-fig. 88). As a result there are channels
running through each annule between the punctation processes which continue as
grooves in the innermost sub-annule.

EUCHROMADORA AND SIMILAR NEMATODES 187

Steiner & Hoeppli (1926) create a new genus Dicriconema, for a new species
D. tennis, based on the study of ". . .an apparently young female ..." from Japan.
The genus was diagnosed largely on the structure of the cuticle and, although both
Wieser (1954) and De Coninck (1965) treat it as a " solid toothed " form, the original
authors give no explicit information about the structure of the dorsal onchium and
their figure (Tafel i, Fig. i) suggests that it was in fact hollow. The description
shows that the cuticle is marked by elongate punctations over the whole surface,
except for dorso- and ventro-lateral strips, and it appears possible that they studied
a specimen similar to those discussed here.

I prefer not to use the name Dicriconema because it is extremely doubtful that
D. tennis could ever be unequivocably recognized. The difficulty is that mentioned in
discussing Odontocricus. It is not yet possible to be sure that the structure of the
cuticle is always covariant with the same features of other parts of the animal and,
even if this could be established, it is doubtful that D. tennis could ever be anything
other than a species inquirenda, as I propose to treat it. It therefore appears advis-
able, because fairly drastic changes are involved anyway, that the knot be cut by
introducing a new name for a generic group containing E. shirleyae, E. colesi, N. alii
and related species.

AUSTRANEMA gen. nov.

Chromadoridae : CUTICLE: small, elongate punctations anteriorly which are not
obviously battlement-like on all specimens but which are due to processes within
cuticle developed from both sides of annules; cuticular annules markedly skewed
anteriorly and posteriorly; punctations present on dorsal and ventral surfaces,
although sometimes slimmer than those laterally at same level; dorso- and ventro-
lateral files of articulating processes ;

HEAD: not obviously set off; dorsal onchium difficult to classify but usually
hollow; no denticles along anterior dorsal edge of oesophastome ; flanges on lateral
walls of oesophastome with dentate anterior edges; prominent, cone-like, anteriorly
directed onchia on ventral sectors at base of oesophastome; outer and intermediate
circles of cephalic sense organs at same level, all setose; mouth surrounded by twelve
rugae; amphids prominent, transverse slits bounded by prominent cuticular fringes;
oesophagus without posterior bulb, but slightly expanded posteriorly.

MALES: gubernaculum with prominent L-shaped lateral pieces; mid- ventral,
elongate, pre-cloacal modification present; tail long and slim.

TYPE SPECIES: Euchromadora colesi Inglis, 1968.

OTHER SPECIES: A. shirleyae (Coles, 1965); A. alii (Murphy, 1965); A. pectinata
(Wieser & Hopper, 1967).

Euchromadora pectinata Wieser & Hopper, 1967 is described as having a pre-cloacal
mid-ventral modification of the cuticle and an oesophagus with a dilated posterior
end but no distinct bulb; but the intermediate circle of cephalic sense organs is
papillose. I shall, therefore, refer it to Austranema because of the first two features
and full confirmation must depend upon a re-examination of the specimens. It

i88 W. G. INGLIS

should be noted that Wieser & Hopper (1967) refer to the presence of a post-cloacal
thickening of the cuticle. This kind of modification occurs in some specimens of all
the species I have seen but does not appear to be consistent in its occurrence.
Perhaps I have simply been unable to see it.

Euchromadora meadi Wieser & Hopper, 1967 is left in the genus Euchromadora
although it looks suspiciously like a member of Austranema. Here again a
re-examination of the specimens is called for.

Uncertain Euchromadora and Euchromadora-like Species

Some nominal species which have been described as members of Euchromadora
and some species which appear similar to the various genera recognized above remain
to be considered. Some of these species may be easily dismissed because of poor
descriptions but others have to be considered because the convergence demonstrated
above makes their generic placement difficult or uncertain. In such cases there is
little doubt that the species concerned could be recognized again but there is
insufficient information at present to allow them to be referred to one or other of the
Euchromadora-like genera recognized here.

Cobb (1914) describes four new species from the Antarctic, which he refers to
Euchromadora. More recently two of them are referred to other genera by Wieser
(1954) : E. antarctica to Prochromadorella and E. septentrionalis to (?) Hypodontolai-
mus ; while the other two E. denticulata and E. meridiana, are kept in Euchromadora
as doubtful species which are treated as species dubiae by Coles (1965). While
agreeing with this, it is worth pointing out that it is now clear that neither of these
latter species is a Euchromadora-type. Both have " hollow " dorsal onchia, both
have two circles of setae, both have very prominent amphids and both have
extensive, non-punctate or lightly punctate regions anteriorly. Unfortunately the
descriptions make it impossible to establish their relationships more fully and it
remains doubtful that either could be recognized again.

A large number of nominal species of Euchromadora have been named, described,
redescribed or reported by Allgen in a large number of publications. Some of these
have been " recognized " and redescribed by other authors but the general standard
of Allgen's work is so poor that many such redescriptions are based on guess work.
The majority of Allgen's species have been treated as unrecognizable by several
authors (summarized by Coles, 1965), and will not be reconsidered here.

Steiner (1918) describes five new species of Euchromadora ah 1 of which are best
treated as dubia, as proposed by Coles (1965) who, however, treats E. hupferi as a
species inquirenda. As argued above, although it is possible to recognize the genus
to which this species should be referred it is most unlikely that the species could ever
be recognized. As a consequence E. hupferi is also considered a species dubia.

It is worth noting that E. longicaudata Steiner, 1918, in which the amphid is
prominent with, apparently, a double contour and a dorsal onchium which appears
to be of the hollow-type, is referred to Actinonema by Wieser (1954). Wieser, never-
theless, points out that " the male genital apparatus of this species is insufficiently
described but it clearly differs in structure from that of the following species."

EUCHROMADORA AND SIMILAR NEMATODES 189

(i.e. A. pachydermatum Cobb, 1920). This is certainly not true since Steiner only
studied a female and naturally says nothing about the male and it is extremely
unlikely that Steiner's species could be unequivocably recognized again or differen-
tiated from other species of any genus to which it might be thought to belong.

Wieser's (1954) comment on A. longicaudata must refer to the descriptions of
Pareuchromadora fragilis of Allgen, 1942 and P. setifer Schuurmans Stekhoven, 1943
both of which he refers to the synonymy of E. longicaudata. There is no obvious
justification for any of this since P. fragilis is, like E. longicaudata, known only from
a female while P. setifer is recognizably an Actinonema species. As a consequence
I treat E. longicaudatum Steiner, 1918 as a species dubia; P. fragilis of Allgen, 1942
as a species dubia and recognize P. setifer as a species inquirenda and refer it to
Actinonema.

Also in 1943 Schuurmans Stekhoven describes Euchromadora inflatispiculum
which is clearly not referable to that genus and must remain a species incertae sedis
until further specimens have been studied. From the descriptions it appears to
have a hollow dorsal onchium and the redescription of Schuurmans Stekhoven (1950)
does not appear to be the same species as that described in 1943, as is pointed out
by both Wieser (1954) and Coles (1965).

Euchromadora arctica Filipjev, 1946 is a species with a hollow dorsal onchium, an
oesophagus without a posterior bulb, with L-shaped lateral pieces to the guberna-
culum, four cephalic setae, an extensive clear zone of cuticle at the anterior end of the
body and, according to Filipjev (1946), there is a mid- ventral pre-cloacal papilla
on the male. If this be correct the species is very different from any of the others
described here or by others. In addition the form of the dorsal onchium is very
cyatholaimid in appearance. As a result I propose to treat this species as incertae
sedis although it shows some similarity to Austranema.

REMARKS ON THE CLASSIFICATION OF THE CHROM ADORID AE

Almost all discussion of the classification of free-living marine nematodes, in
particular, and all non-parasitic forms in general, starts or finishes by stressing the
doubtful value of much of the present grouping. Such comments are undoubtedly
justified. The classification of the Family Chromadoridae is no exception and is
probably as unreliable as that of any other family. Its limits and divisions float in a
cloud of taxonomic uncertainty lightened only by flashes of morphological ignorance.
As De Coninck says, "Toute lafamille des Chromadoridae devra etre soigneusement
revisee" (De Coninck, 1965, page 637.)

Although it is easy to offer destructive criticism of the classification, constructive
suggestions are much more difficult to formulate because of our extensive lack of
knowledge about the animals involved. The weakness of this position is well
exemplified by the present work. It is only too clear that there is no sound basis
on which to erect a classification ; and there is no background of morphology against
which to assess new information. The immediate consequence is that it is difficult to
suggest easily observed characters or, more usefully, characters which have pre-
viously been described, which can be relied upon to be co variant with those structures
of the head and cuticle described above.

igo W. G. INGLIS

The recognition of one group with a solid dorsal onchium and one with a hollow
onchium is easy when the extreme and clear cut conditions are compared. The
division is difficult, if not impossible, to apply when a solid " hollow " onchium is
examined. Equally the two groups, so far as they have been studied, appear to
differ in several covariant characters of the cuticle and dentition but it is difficult
to apply this to those groups with simple punctate cuticles and we do not know, as
yet, whether all the " hollow " toothed forms are similar in their other morpholo-
gical characters. In other words it has been established that some " solid " toothed
forms are, in other characters, most similar to " hollow " toothed but we do not yet
know if forms which look obviously " hollow " toothed may not later be found to have
great similarities with the obviously solid toothed.

In an attempt to overcome some of this uncertainty I have looked at a very small
number of other species. In Spilophorella paradoxa (de Man, 1888), for example,
there is obvious difficulty in describing the structure of the dorsal onchium. I
would call it hollow but most groupings presuppose it is solid (e.g. Wieser, 1954;
De Coninck, 1965). Be that as it may, there is a pair of small, ventral, anteriorly
directed cone-like onchia in the oesophastome and there appear to be lateral flanges
arising from the lateral oesopharhabdions. In these respects, therefore, Spilophorella
falls into the hollow-toothed group of genera.

The conditions in Hypodontolaimus are much clearer, because the specimens of
H . slacksmithi (described below) are relatively large. Here the mouth is surrounded
by twelve, inwardly pointing, flanges developed from the wall of the cheilostome and
there are no rugae as in typical Chromadoridae, the dorsal onchium is very prominent
and hollow, there are no denticles along the dorsal anterior edge of the oesophastome,
the oesophastome is rectangular in transverse section with two ventral, cone-like,
anteriorly directed onchia on the ventral sectors and there are typical lateral flanges
developed from the lateral walls of the oesophastome. These flanges are most
anterior towards their ventral edges so that they look like pointed onchia on focusing
up on en face preparations (Text-figs. 57-59).

In addition, in Hypodontolaimus the cuticle is of a battlement-type and a similar
condition is suggested for various other " hollow " toothed species. For example
Filipjev's (1918) illustrations of Chromadora sabulicola (Plate 8, fig. 5ob) and C.
poecilosomoides (Plate 8, figs. 52a, b, c) [both now referred to Neochromadora] show
a battlement-like cuticle co-occurring with a hollow dorsal onchium. Equally many
of Gerlach's (1951) illustrations show a similar association between a hollow onchium
and a battlement-like cuticle.

To extrapolate, it appears that the presence of a hollow dorsal onchium is fre-
quently associated with the presence of lateral flanges and cone-like ventral onchia
in the oesophastome, and with the possession of a battlement-like modification of
the dense component within the cuticle. There is, as yet, insufficient information
on the structure of the cuticle and oesophastome in those species with an unequi-
vocably solid dorsal onchium to allow any conclusions to be reached and the relation-
ships of those species with simple punctate cuticles may always be difficult to estab-
lish. In spite of this uncertainty it appears that Wieser's (1954) treatment of the
Chromadoridae in two groups distinguished crudely by the form of the dorsal

EUCHROMADORA AND SIMILAR NEMATODES 191

onchium is of significance but it is doubtful that enough information yet exists to
make De Coninck's (1965) division of the family into subfamilies on the same character
acceptable.

For example there appear to be at least two distinct groupings possible for those
genera in which the dorsal onchium is hollow. One of the groups consists of the
genera considered above, which appears to culminate in a form such as Parapinnan-
ema wilsoni, in which the cuticle is elaborate, thick and strong but the dorsal onchium
is small. The other group contains those nematodes in which the dorsal onchium
is very prominent and obviously hollow, frequently with a massive development of
the oesophageal musculature associated with it, the cuticle is not thick, very elaborate
and dark, and the gubernaculum is relatively simple. This group contains genera
such as Hypodontolaimus and Nygmatonchus.

I would, therefore, expect that further study will lead to the recognition of at
least three distinct groups within the subfamilies Chromadorinae and Hypodonto-
laiminae as recognized by De Coninck (1965). This does not, however, take into
consideration those forms grouped by De Coninck in the Ethmolaiminae, Cyatho-
laimidae and even the Desmodoridae, the analysis of whose relationships is closely
tied to the Chromadoridae. In all these groups there appear to be various indepen-
dent lines of modification all of which lead to a stronger and more elaborate cuticle.
As a result there is almost certainly a large amount of concealed convergence which
is, as yet, unrecognized so that much of the current classification continues to be
suspect.

DESCRIPTIVE SECTION

In agreement with the arguments of Wieser & Hopper (1967) I only give absolute
measurements of specimens and have not given ratios.

Euchromadora striata (Eberth, 1863), Filipjer, 1918
(Text-figs. 12-24; 91-93)

Weed and associated hold-fasts and sand; silty and sheltered. In 60 cm. water.
Cheyne Beach, Nr. Albany, W.A.

Weed and associated hold-fasts and sand; on exposed rocks. Goode Beach,
Albany, W.A.

Weed and associated hold-fasts and sand; no silt, on exposed beach in 60 cm.
water. Sarge Bay, Cape Leeuwin, W.A.

Among weed in rock pools. Robert Point, Mandurah, W.A.

Among weed on coral reef in 60 cm. water. Radar Reef, Rottnest Island, W.A.
measurements (mm.).

MALES. Body length: 1-61; 1-64 ; 1-67; 1-86; 2-37. Body breadth: 0-078
0-078, 0-076; 0-068; 0-082. Diameter of head: 0-031; 0-030; 0-033; 0-033; '33-
Length of cephalic setae: 0-009; o-oio; o-on; o-on; o-on. Oesophagus length:
0'3i; 0'33i 0-33; 0-35; 0-36. Length of spicules: 0-075; 0-083; 0-085; 0-085; 0-079.
Gubernaculum length; 0-052; 0-047; '5 2 i 0-048; 0-046. Tail length: 0-185;
0-238; 0-232; 0-212; 0-215. Cloacal diameter: 0-066; 0-057; 0-061; 0-052; 0-059.

1 92

W. G. INGLIS

94

96

98

EUCHROMADORA AND SIMILAR NEMATODES IQ3

FEMALES. Body length: 2-13; 2-16; 2-63. Body breadth: o-no; 0-098; 0.131.
Diameter of head: 0-036; 0-036; 0-037. Length cephalic setae: 0-013; o-on; 0-013.
Oesophagus length: 0-40; 0-39; 0-43. Tail length: 0-208; 0-221; 0-234. Distance
of vulva from anterior end: 1-18; 1-09; 1-25. Size of eggs: 0-069 x 0-046; 0-073 X
0-063.

This species is the same as that described by Coles (1965) and Filipjev (1918)
and it should be noted that Coles' figure of the spicules (Fig. 26) shows the spicules
as rather too narrow posteriorly and the " serrated " posterior end to the lateral
piece of the gubernaculum is three denticles (Text-figs. 91, 92) as in the Australian
specimens.

The only major differences between the specimens from Australia and England are
that the setae are slightly shorter in the Australian forms. The detailed structure
of the head is described on page 162.

This species is wide-spread and common on the coasts of Western Australia and
I have collected it from as far north as Perth, south down the coast to Cape Leeuwin
and along the south coast as far as Cheyne Beach about 30 miles east of Albany.

Euchromadora eileenae sp. nov.
(Text-figs. 31-37; 94-95)

Weed on coral in 60-100 cm. water; Radar Reef, Strickland Bay, Rottnest, W.A.
(Type locality).

Weed and sand brought up in trawl from 12 m. ; Shark Bay (midway between
Denham and Dirk Hartog Island), W.A.

Weed and sand associated with hold-fasts, silty, sheltered, in 60 cm. water.
Cheyne Beach, Nr. Albany, W.A.

Measurements (mm.)

MALES. Body length: 1-63; 1-79; 1-96; 2-05. Body breadth: 0-048; 0-064;
0-062; 0-059. Diameter of head: 0-023; 0-026; 0-026; 0-024. Length of cephalic
setae: 0-008; 0-009; 0-009; 0-008. Length of oesophagus: 0-269; 0-281; 0-316;
0-296. Spicule length: 0-049; '53i '58; 0-061. Gubernaculum length: 0-034;
0-035; 0-040; 0-038. Tail length : 0-225; 0-189; 0-221; 0-236. Cloacal diameter:
0-039 ; '43 i 0-041 ; 0-044.

FEMALES: Body length: 1-74; 1-87; 1-93. Body breadth: 0-093; 0-089; 0*091.
Diameter of head: 0-026; 0-029; ' 02 9- Length of cephalic setae: 0-009; 0-009;
0*10. Length of oesophagus: 0-310; 0-314; 0-326. Tail length: 0-188; 0-208;
0-223. Anal diameter; 0-039; 0-040; 0-043. Distance of vulva from anterior end of
body: 0-88; 0-91; 0-97. Diameter of eggs (spherical): 0-047-0-049.

FIGS. 9198. 9193. Euchromadora striata. 91-92. Gubernaculum and spicules.
93. Lateral view of male tail. 9495. Euchromadora eileenae. Gubernaculum and spicules.
Difference in appearance due to different angles of view. 96. Graphonema georgei.
Gubernaculum and spicules. 97-98. Parapinnanema wilsoni. 97. Anterior end of body.
Black region is thick cuticle in section. 98. Gubernaculum and spicules.

194 W. G. INGLIS

The structure of the head is described on page 162 and the cuticle is typical of the
genus. The lateral plates are tiny and a series of four files of longish setae occur over
the oesophageal region of the body.

The lateral pieces of the gubernaculum are stout with a very slight ventral curve
at the distal end. On this curved region are two tiny denticles. The spicules are
slightly curved and bear alae. This detail of both the spicules and the gubernaculum
has been confirmed by dissection and is difficult to establish from whole mounts.

E. eileenae differs from all other members of the genus in the shape of the lateral
pieces of the gubernaculum in which the distal end is hardly L-shaped.

The species is fairly common on the coasts of Western Australia with a wide range
from Shark Bay in the north to Cheyne Beach in the east, on the south coast.

Parapinnanema wilsoni gen. et sp. nov.
(Text-figs. 51-55; 66-72; 73-75; 97-98)

From weed and associated sand without silt, on exposed beach in 60 cm. water.
Sarge Bay, Cape Leeuwin, W.A. (Type locality).

From among finger-like green algae in rock-pools, just east of Cape Naturaliste
(Bunker Bay), Geographe Bay, W.A.

Measurements (mm.).

MALES. Body length: 1-34; 1-47; 1-50; 1-78. Body breadth: 0-039; 0-034;
0-044; 0-046. Head diameter: 0-016; 0-016; 0-016; 0-018. Length of cephalic
setae: 0-007; 0*007; 0-008; 0-006. Oesophagus length: 0-23; 0-24; 0-29; 0-27.
Length of spicules: 0-041; 0-040; 0-039; 0-046. Length of gubernaculum: 0-023;
0-024; 0-022; 0-026. Length of tail: 0-143; 0-153; 0-148; 0-151. Cloacal diameter:
0-038; 0-047; 0-042; 0-041.

FEMALES: Body length: 2-34; 2-76. Body breadth: 0-073; 0-069. Head dia-
meter: 0-023; 0-026. Length of cephalic setae: 0-006; 0-007. Oesophagus length:
0-42; 0-46. Length of tail: 0-257; 0-261. Anal diameter: 0-047; 0-049. Size of
eggs: 0-053 X 0-042.

The structure of the head and of the cuticle is described on pages 171, 175 but it
may be stressed that this species is easily recognized in collections by the extreme
thickness of the cuticle over the oesophageal region of the body (Text-fig. 97).

The spicules are evenly curved, without proximal swellings or alae and terminate
distally in sharp points. The lateral pieces of the gubernaculum are fairly broad
and L-shaped. The short, distal arm of the gubernaculum carries a slight bulge
on its inner surface which can be difficult to see on whole specimens but is immediately
obvious on dissection. There is a median raised region of the cuticle anterior to
the cloacal opening.

This species cannot be confused with any other because of the features diag-
nostic of the genus Parapinnanema (page 175). The discovery of further species
referable to that genus must be awaited before a more restricted diagnosis can be
established.

EUCHROMADORA AND SIMILAR NEMATODES 195

Graphonema georgei sp. nov.
(Text-figs. 80-84; 96; 99)

Weed and associated sand in hold-fasts on exposed rocks, Goode Beach, Albany
W.A.

Measurements (mm.).

MALES. Body length: 1-30; 1-42. Body breadth: 0-052; 0-044. Diameter of
head: 0-014; 0-014. Oesophagus length: 0-187; 0-198. Length of cephalic setae;
intermediate /outer : 0-0026/0-0040; 0-0026/0-0040. Spicule length: 0-039; 0-043.
Gubernaculum length: 0-017; 0-017. Tail length: 0-150; 0-164. Cloacal diameter:
0-036; 0-039.

The structure of the head is described above (page 180) and that of the cuticle
on page 178. It may be stressed that the species is easily recognized by the swollen
anterior end, the relatively posterior position of the amphids, the intermediate and
the outer circles of setose cephalic sense organs, and the rather thin and delicate
appearance of the cuticle.

The spicules are equal in length and identical in structure, fairly massive in appear-
ance with narrow alae and slightly barbed posterior ends. The lateral pieces of the
gubernaculum are relatively small and are a slightly curved L-shape (Text-fig. 96).
The tail is long and slim and there is no pre-cloacal thickening on the mid-ventral
surface of the body (Text-fig. 99).

The only other species referred to this genus is G. vulgaris Cobb, 1898 which is
described without illustrations. All the cephalic setae are described as lying rela-
tively near the anterior end of the body, while in the specimens from Western
Australia they are far posterior to the same level. This difference could be because
Cobb studied specimens in which the head was somewhat contracted but this appears
unlikely as he refers to a number of papillae (eighteen) near the border of the mouth
opening. These I interpret as the rugae which surround the mouth opening and the
fact that they could be seen implies that the mouth was not fully closed.

Cobb describes the spicules as "... of uniform size throughout, and are twice
as long as the anal body diameter." while in the Western Australian specimens
the spicules narrow markedly towards their extreme posterior ends and are about
one cloacal width in length.

As a result of these differences I propose to treat the specimens from Western
Australia as a distinct, new species, G. georgei sp. nov.

Austranema alii (Murphy, 1965) comb. nov.
(Text -figs. 38-40; 100-101)

Weed and sand from trawl from 12 metres in Shark Bay, W.A. ; midway between
Dirk Hartog Island and Denham.

W. G. INGLIS

FIGS. 99-109. 99. Graphonema georgei. Lateral view of male tail. 100-101. Austra-
nema alii. 100. Gubernaculum and spicules. Fig. 101. Lateral view of male tail.
102-109. Hypodontolaimus slacksmithi. 102-106. Surface appearance of cuticle.
102. Between cephalic setae and anterior end of lateral differentiation. 103. Anterior
end of lateral differentiation. 104. At anterior end of oesophageal bulb. 105. At
posterior end of oesophagus. 106. At level of cloacal opening. 107. Gubernaculum
and spicules. 108. Male tail. 109. Oesophagus.

EUCHROMADORA AND SIMILAR NEMATODES 197

Measurements (mm.).

MALES. Body length: 170; 178; 2-16. Body breadth: 0-052; 0-046; 0-057-
Head diameter: 0-016; 0-017; 0-019. Setae length: 0-005; '5; 0-005. Oeso-
phagus length: 0-22; 0-24; 0-27. Spicule length: 0-044; '38; 0-052. Gubernaculun
length: 0-026; 0-021; 0-035. Pre-cloacal supplement, length /distance anterior to
cloacal opening: 0-048/0-098; 0-034/0-091; 0-047/0-130. Tail length: 0-19; 0-19;
0-22. Cloacal diameter: 0-038; 0-033; 0-049.

FEMALES. Body length: 1-90; 2-31. Body breadth: 0-051; 0-057. Head dia-
meter: 0-019; 0-021. Setae length: 0-004; 0-005. Oesophagus length: 0-27; 0-32.
Tail length: 0-19; 0-21. Anal diameter: 0-033; o - 37- Vulva from anterior end:
0-90; 1-06. Size of eggs: 0-089 x '33i 0-074 x 0-036.

The spicules are equal and identical (Text-fig. TOO). The lateral pieces of the
gubernaculum are L-shaped with each arm about the same length. The distal end
of the gubernaculum is slightly pointed and bears a slight notch on the under edge
near the tip. There is a median pre-cloacal ventral modification in the form of a
raised region of the cuticle. The three or four cuticular projections which "...
appear to form a kind of bursa ..." (Murphy, 1965, page 207) are due to the dense
annules within the cuticle at the level of the cloacal opening projecting above the
general body surface. This is because the cloacal opening itself is surrounded by
an unannulate region and the projections represent the outer border of this region.

There are distinct granules lying on the lateral surface of the body between the
annules, in a position corresponding to the lateral plates in Euchromadora species.

A . alii differs from the other species referred to the genus most particularly in the
notched distal end to the lateral pieces of the gubernaculum and in the characteristic
shape of the spicules.

Hypodontolaimus slacksmithi sp. nov.
(Text-figs. 56-59; 102-109)

Sand and weed brought up in trawl from 12 m. ; Shark Bay, midway between
Dirk Hartog Island and Denham, W.A. (Type locality).
Weed and hold-fasts on rocky flat in 12 cm. water; Cowaramup Bay, W.A.

Measurements (mm.).

MALES. Body length: 1-34; 1-46. Body breadth: 0-051; 0-052. Length of
cephalic setae: 0-020; 0-022. Diameter of head: 0-036; 0-038. Length of oeso-
phagus: 0-19; 0-23. Length of spicules: 0-039; o '44- Length of gubernaculum:
0-030; 0-031. Length of tail: 0-13; 0-15; Cloacal diameter: 0-033: 0-042.

FEMALES. Body length: 1-38; 1-43. Body breadth: 0-075; 0-080. Length of
cephalic setae: 0-021; 0-022. Diameter of head: 0-033; '37- Length of oeso-
phagus: 0-22; 0-24. Distance of vulva from anterior end of body: 0-57; 0-63.
Length of tail: 0-14; 0-17. Anal diameter: 0-039; 0-041.

The cuticular markings consist of prominent ovoid punctations just posterior to
the cephalic setae (Text-fig. 102) and remain like this until the lateral differentiation

ig8 W. G. INGLIS

begins about half-way along the oesophagus length. Here the lateral differentiation,
which is raised above the general surface of the body, consists of a bar flanked by
large dots in turn flanked by battlement-like punctations which occur on both sides
of the annules (Text-fig. 103). By the level of the anterior end of the oesophageal
bulb, the lateral markings have become wholly battlement-like along the anterior
edge of each annule. At the posterior end of the oesophagus, the lateral bars of the
differentiation are narrow, the large dots flanking them are relatively small and the
markings outside these again are now in the form of small, squarish punctations of
which one row is level with the large dots of the lateral differentiation while one row
lies between the large dots (Text-figs. 104-105) . The punctations continue to become
simpler posteriorly and less prominent until by the level of the cloaca or anus they
consist of rows of small dots while the lateral differentiation consists of a clear strip
flanked by a series of punctations which are only very slightly larger than those found
elsewhere on the surface of the body (Text-fig. 106). There appear to be four files
of ventro- and dorso-lateral setae on the body but they are small and indistinct.

The mouth opening is bounded by twelve lobes, without rugae, which are sup-
ported by twelve flanges of the cheilorhabdion. These flanges project inwards and
over the cavity of the cheilostome as rather hook-like structures in optical section.
The cephalic sense organs consist of two circles of six papillae and an outer circle
of four long setae. The amphid was not seen but the head is set-off from the body.
The tail narrows fairly rapidly and ends in a long, pointed spinerette.

The oesophastome contains a very large, hollow dorsal onchium with a prominent
dorsal apophysis in the male. The dorsal side of the oesophastome is very thick
and slightly irregular in optical section longitudinally (Text-fig. 59). There is a
lateral flange on each lateral wall of the retangular oesophastome, in en face view,
which is highest towards the ventral side, and there is a single cone-like, anteriorly
directed onchium on each ventro-lateral sector (Text-figs. 57-59).

The oesophagus ends posteriorly in a very prominent double bulb with a prominent
cuticular lining (Text-fig. 109), of which the anterior bulb is slightly smaller than the
more posterior.

The spicules are evenly curved and taper posteriorly to finely rounded ends.
The gubernaculum is about two-thirds of the length of the spicules and is prominent
with anterior, lateral processes which lie over the spicules laterally. There is a
pair of slight bumps near the distal end of the gubernaculum on the side away from
the spicules (Text-fig. 107).

This species is referable to the subgenus Ptycholaimellus Cobb, 1920 of the genus
Hypodontolaimus de Man, 1888 as recognized by Gerlach (1955) and as followed by
Wieser & Hopper (1967). It would probably be better to treat the group as a full
genus, if it is to be recognized at all. There is little value in recognizing subgenera
within free-living marine nematodes since they simply add another name to be
considered.

Be that as it may, H. slacksmithi is, therefore, similar to H. carinatus Cobb, 1920 ;
H. poncticus Filipjev, 1922; H. pandispiculatus Hopper, 1961 and H. macrodentatus
Timm, 1961, from all of which it differs in the relatively stout, smoothly curved
spicules and the massive gubernaculum.

EUCHROMADORA AND SIMILAR NEMATODES 199

SUMMARY OF GENERA AND SPECIES RECOGNIZED

Given here are brief summaries of the major diagnostic characters of the genera
recognized and lists of the species referred to them. All genera are referred to the
family Chromadoridae sensu lato.

EUCHROMADORA de Man, 1886

Head blunt with outer four sense organs setae; amphids without surrounding
cuticular fringe; dorsal onchium solid; rows of denticles on lateral and ventral
walls of oesophastome ; oesophagus without posterior bulb ; cuticle complex, unmarked
dorsally and ventrally, not markedly thick over oesophageal region; lateral plates
always present; no pre-cloacal modifications on male; tail short and stout.

Species :

E. vulgaris (Bastian, 1865); E. gaulica Inglis, 1962; E. meadi Wieser & Hopper,
1967; E. permutabilis Wieser, 1954; E. tokiokai Wieser, 1955; E. striata (Eberth,
1863) ; E. eileenae sp. nov.

STEINERIDORA gen. nov.

Generally similar to Euchromadora ; differs in massive squarish dorsal onchium
and sickle-like prominent onchia laterally and ventrally; oesophagus with posterior
bulb; cuticular markings always about same size over full length of body laterally.

Species :

S. loricata (Steiner, 1916) ; 5. adriatica (von Daday, 1901) ; 5. archaica (Steiner
& Hoeppli, 1926); 5. (?) dubia (Steiner, 1918) sp. inq.

PARAPINNANEMA gen. nov.

Head set off as narrow region because of very thick cuticle over oesophagus
length of body; intermediate circle of sense organs six setae and outer four setae;
amphids with surrounding fringe of cuticle; dorsal onchium hollow; flanges on lateral
walls and onchia ventrally in oesophastome; oesophagus without posterior bulb;
cuticle very thick over oesophagus region and very battlement-like; cuticle with
punctations dorsally and ventrally; tail long and slim; raised region of pre-cloacal
modification on male.

Species:
P. wilsoni sp. nov.

PROTOCHROMADORA gen. nov.

Slim with rounded head; outer sense organs setae; dorsal onchium difficult to
classify but frequently hollow looking; no elaborations on lateral walls of oeso-
phastome but single onchium on each ventral sector; oesophagus without posterior
bulb ; amphid without cuticular fringe ; cuticle not markedly thick over oesophageal

200 W. G. INGLIS

region; cuticle complex (?) without punctations dorsally and ventrally; tail short
and stout ; no pre-cloacal modification.

Species :

P. scampae (Coles, 1965) ; P. mediterranea (Allgen, 1942) ; P. parafricana (Gerlach,
1958).

GRAPHONEMA Cobb, 1898

Head set off as swollen, bulb-like region; cephalic sense organs in three well
separated circles, outer of four setae and intermediate of six setae ; amphid relatively
far posterior on head, with cuticular fringe; dorsal onchium small and hollow;
lateral flanges and ventral onchia in oesophastome ; oseophagus without posterior
bulb; cuticle fairly thin with distinct annulations and punctations; tail long and
slim; no pre-cloacal modification on male.

Species :

G. vulgaris Cobb, 1898; G. georgei sp. nov. ; G. amokurae (Ditlevsen, 1921).

INNOCUONEMA gen. nov.

Dorsal onchium hollow; with associated dorsal swelling of oesophageal muscu-
lature ; outer circle of cephalic sense organs setae ; amphid not prominent ; posterior
oesophageal bulb present ; cuticle complex ; tail long and slim ; no pre-cloacal modifi-
cation on male.

Species :

/. flaccida (Wieser, 1959) ; /. clovosa (Wieser, 1959) ; /. tentabunda (de Man, 1890) ;
I. chilensis nom. nov.

NYGMATONCHUS Cobb in Cobb, 1933

Cuticle complex anteriorly, elongate punctations posteriorly; lateral differen-
tiation present ; intermediate and outer circles of cephalic sense organs setae ; dorsal
onchium hollow; amphid with prominent double contour (?) ; no posterior oesophageal
bulb; tail long and slim with pre-cloacal modification(?).

Species :

N. scriptus Cobb in Cobb, 1933 ; N. fossiferus Wieser, 1954 ; N. bicornata (Wieser,
1959) comb. nov. ; N. minutus Gerlach, 1967.

AUSTRANEMA gen. nov.

Head not set off or blunt ; dorsal onchium difficult to classify but generally hollow
and anteriorly directed; outer and intermediate cephalic sense organs in one circle
of ten setae; flanges laterally and onchia ventrally in oesophastome; oesophagus
without posterior bulb, although expanded evenly posteriorly; cuticle complex,

EUCHROMADORA AND SIMILAR NEMATODES 201

marked dorsally and ventrally, and thickened over oesophagus region ; tail long and
thin; pre-cloacal mid- ventral modification present on males.

Species :

A. colesi (Inglis, 1968); A. shirleyae (Coles, 1965); A. alii (Murphy, 1965); A.
pectinata (Wieser & Hopper, 1967) .

LIST OF NOMENCLATURAL CHANGES PROPOSED

Chromadora

paraheterophya Allgen, 1932
sabangensis Steiner, 1915
spectabilis Allgen, 1932
suilla Allgen, 1947
tentabunda de Man, 1890

Dicriconema

tenuis Steiner & Hoeppli, 1926

Euchromadora

archaica Steiner & Hoeppli, 1926

arctica Filipjev, 1946

colesi Inglis, 1968

denticulate Cobb, 1914

dubia Steiner, 1915

hupferi Steiner, 1918

inflatispiculum Schuurmans

Stekhoven, 1943
mediterranea Allgen, 1947
meridiana Cobb, 1914
parafricana Gerlach, 1958
pectinata Wieser & Hopper, 1967
scampae Coles, 1965
shirleyae Coles, 1965
stateni Allgen, 1930

Graphonema

amokuroides Wieser, 1954
clovosa Wieser, 1959
flaccida Wieser, 1959

Neochromadora

bicornata Wieser, 1959

Nygmatonchus

alii Murphy, 1965

Odontocricus

hupferi Steiner, 1918

= species dubia

= species inquirenda

= species dubia

= species dubia

Innocuonema t. comb. nov.

= species inquirenda

= Steineridora a. comb. nov.

= incertae sedis

= Austranema c. comb. nov.

= species inquirenda

= Steineridora (?) d, comb. nov.

= species dubia

= incertae sedis

= Protochromadora m. comb. nov.

= species inquirenda

= Protochromadora p. comb. nov.

= Austranema p. comb. nov.

= Protochromadora s. comb. nov.

= Austranema s. comb. nov.

= species dubia

= Innocuonema chilensis nom. nov.
= Innocuonema c. comb. nov.
= Innocuonema f. comb. nov.

= Nygmatonchus b. comb. nov.
= Austranema a. comb. nov.
= species dubia

202 W. G. INGLIS

Pareuchromadora

fragilis Allgen, 1942 = species dubia

setifer Schuurmans Stekhoven, 1943 = species inquirenda
Spilophora

adriatica v. Daday, 1901 = Steineridora a. comb. nov.

amokurae Ditlevsen, 1921 = Steineridora a. comb. nov.

amokuroides Allgen, 1927 = species dubia

loricata Steiner, 1916 = Steineridora I. comb. nov.

norvegica Allgen, 1932 = species dubia

pusilla Allgen, 1947 = species dubia

ACKNOWLEDGEMENTS

This work was started in the Western Australian Museum while I was Exchange
Curator in 1966-67 and I would express my thanks to all those who helped me
during my time there. In particular Mr. R. Slack-Smith, Western Australian
Department of Fisheries for arranging a visit to Shark Bay where some of the speci-
mens described here were collected. Also Miss C. M. Chambers (now Mrs. Martin)
for assistance in Perth. In London thanks are due to Miss Eileen A. Mitchell who
sorted so many samples for me.

Hypodontolaimus slacksmithi is named in appreciation of Mr. Slack-Smith's efforts
on my behalf; Euchromadora eileenae is a small token of appreciation for Miss
Mitchell's endurance in sorting formalin samples while Austranema wilsoni and
Graphonema georgei are named in honour of Dr. B. R. Wilson and Dr. R. W. George
respectively, W.A. Museum for the many kindnesses shown to me by them and their
families during my stay in Perth, W.A.

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Ledokolvn Parokhode " G. Se