Bull. nat. Hist. Mus. Lond. (Zool.) 67(1): 1-24

Issued 28 June 2001

Freshwater nematodes from Loch Ness,

Scotland Part I. The orders Tylenchida

JUN

Thorne, 1949 and Rhabditida Chitwood, 1933.: RAL USR

(Nematoda, Secernentea). ee

F. R. WANLESS AND R. HUNTER’

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

CONTENTS

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SYNOPSIS.

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During a survey of the nematodes of Loch Ness, Scotland (Project Urquhart 1993-94) 94 nematode species were

identified from benthic core samples collected from the profundal (90-211 m) and from river mouth areas (< 0.5 m). This paper

presents a taxonomic review of the nematodes of the two orders Tylenchida Thorne, 1949 and Rhabditida Chitwood, 1933. Thirty

five species are described and figured; some morphological features are illustrated by Normarski interference-contrast

microscopy.

INTRODUCTION

This study is based on nematodes collected in Loch Ness, Scotland

during Project Urquhart (1993-94). The first part presented here

deals with species that belong in the orders Tylenchida Thorne, 1949

and Rhabditida Chitwood, 1933. The second part, now in prepara-

tion, will cover species of the remaining orders. The results of

ecological studies (R. David,1998 — Ph.D. thesis) will be published

elsewhere. The following brief profile of Loch Ness is essentially

based on the literature Maitland (1981) and Shine & Martin (1987).

Loch Ness is the largest body of freshwater in the United King-

dom. It runs in a north-east/ south-west direction and is 39 km in

length, but only 2.4 km at its widest point with an average depth of

132 m. A new maximum depth of 239.5 m was found during a

Project Urquhart sonar survey. The loch basin has in most places

very steep, sometimes precipitous shelving sides, thus it has a very

small shore zone for its surface area Maitland (1981). The flat bed of

the loch is divided into two basins of 220 m depth as a result of

fluvial deposits of material opposite the entrance to the River

Foyers. The floor is extensively covered by black lake sediments

which occupies over 50% of the area of Loch Ness greater than 150

m depth. Shine & Martin (1987) found that a 4 m core only just

penetrated to clay, suggesting that the organic sediment is at least of

that depth. Sediments analysed from a single core obtained from the

profundal area of Loch Ness were found to consist of fine size

fractions with clay (< 39 um, 45.8%), fine silt (39-63 um, 18.1%)

*R. Hunter nee R. David.

and sand (63-125 um, 36.4%) (Smith B. D. et al., 1981).

Water levels remain relatively constant throughout the year al-

though stream levels vary because of snow melt in spring, but more

importantly because of fluctuations in rainfall. The loch has a

catchment area of 1775 km? providing 84 m* s"' mean inflow with

the majority of the catchment being rough ground or forest, and only

7.5 km? urbanized. Surface water is slightly acidic with an annual

mean pH 6.7; it has low conductivity 30 uS cm"! at 20°C and is

stained with humics. Low conductivity levels in lake water are often

associated with dark colouration and a high concentration of organic

compounds because they have been drained through swampy/peaty

regions. The total organic carbon concentrations of the surface water

is 3 mg Cl Maitland (1981). In many systems of lake classifica-

tion, Loch Ness falls within the nutrient-poor Spence (1967) and

oligotrophic category Vollenweider (1968). However, Bailey et al.,

1981 debate whether Loch Ness along with Loch Awe should be

described as dystrophic (acid), a category considered by Jarnefelt

(1958) as supplementary to, rather than comparable with oligotropic

and eutrophic categories.

The dissolved oxygen content of the Loch Ness water column

was, along with other lochs studied, consistently found to be over

80% saturated (often approaching 100%) even during periods of

stratification Smith, I. R. et al., (1981)

Under the classification system of Yoshimura (1936), Loch Ness

is a warm monomictic lake which means that the temperature never

falls below 4°C at any depth, there is one circulation of the water

column every year in winter and the loch is stratified during the

summer Murray & Pullar (1910). From late autumn to the spring the

loch is the same temperature throughout, with progressive develop-

ment of stratified conditions from May to the following autumn. The

temperature range in midsummer is typically 14.5°C at the surface to

5.5°C at 220 m (Mackal & Love, 1969; Murray & Pullar, 1910).

Maitland (1981) gives a detailed account of human activities

which influence the environmental parameters inherent in scientific

studies effecting Loch Ness. The loch is relatively unpolluted despite

arable, tourist and urban influences. The Scottish Environmental

Protection Agency has supplied details of sewerage inputs to Loch

Ness which indicate that there are seven direct septic tank inputs

from small communities or hotels and tourist attractions, only two of

which have monitored flows. Another septic tank inflow enters

Loch Ness indirectly via the River Moriston. There are two activated

sludge plants for full biological treatment, one at Drumnadrochit

and another at Fort Augustus. When maximum flow is reached,

sewerage by-passes the treatment plant and discharges untreated

into the water course. Seasonal information on the loading variation

was not available. However, Drumnadrochit and Fort Augustus are

known to frequently breach their consent limits. The water regime of

Loch Ness is affected directly by an hydro-electric pump storage

scheme based at Foyers and indirectly by four more in the catchment

area. These influences may have an effect on the flora and fauna of

Loch Ness.

NEMATODE FAUNA

So far as is known the only study of nematodes from freshwater

lakes of the United Kingdom is that of Martin et al., (1993) who

published a full listing of macrobenthic and meiofaunal species from

profundal sediments of both Loch Ness and Morar. Their list of

nematodes from Loch Ness profundal cores (2 200 m) although not

exhaustive includes /ronus tenuicaudatus de Man, Tripyla glomerans

grp, Tobrilus gracilis grp, Tobrilus cf. pellucidus and Dorylaimid

‘A’. Additional species listed (Eumonhystera filiformis grp,

Eumonhystera cf. longicaudatula, Ethmolaimus pratensis grp,

Aphanolaimus sp. and Dorylaimus cf. stagnalis) were collected at

lesser depths such as 50-170 m.

In addition Coles (1996) records Chromadorita tenuis Schneider,

1906 and Aphanolaimus ? aquaticus Daday, 1894 from ‘trickle of

freshwater spring from beside footpath from Goyes Hotel, Foyers

Harbour, Loch Ness, Scotland’.

F.R. WANLESS AND R. HUNTER

MATERIAL AND METHODS

SAMPLING AND FIXATION

Profundal sampling: cores were taken with a Bowers and Connolly

Multicorer deployed from the research vessel Seol Mara. This

device takes four core tubes (each of 5.9 cm diameter giving a core

surface area of 27.34 cm’), allowing sediments to be recovered

without disturbing the sediment water interface.

Sediments were sampled at four stations: Fort Augustus Bay,

Deep South Basin, Foyers Plateau and Deep North Basin covering a

range of depths between 91 and 211 m (Fig. 1, Table 1). At each

station sampling was concentrated within approximately 10-20 m?

between 10—13th July 1993, allowing for boat drift. Eight cores

were collected from each station except for the Deep North Basin

where only four cores were taken because of poor weather conditi-

ons. Of the eight cores collected six were used for faunal analysis; of

these five were sectioned into two, one centimeter layers. One was

sectioned into five one centimeter layers i.e. 0-1 cm, 1—2 cm, 2-3

cm, 3—4 and 4-5 cm. The seventh core was used to take redox

potential readings immediately on recovery. The top two centimeters

of the remaining core was placed in a cold box for carbon content

analysis.

River sampling: sediments were collected by hand with a coring

tube of the same dimensions as used for collecting profundal sam-

ples. The cores were sectioned as above. Three stations i.e. Rivers

Oich, Foyers and Moriston were selected by surveying the area for

soft sediment similar in grain size to the profundal samples where

core tubes could used with ease. Samples were then taken randomly

within an area of 10 m?. All samples were sublittoral ranging from

0.08-0.39 m depth. Eighteen cores, six from each station were

collected from 7—8th August 1994.

After slicing, all samples were immediately fixed using 8%

formaldehyde, at ambient temperature, giving a final dilution of 4%

after mixing. A trace of rose bengal was added to the fixative to

temporarily stain the nematodes, thus making them easier to see

when counting and sorting.

In addition to the stations listed in Table 1, four sub-littoral

qualitative cores were taken from a sandy beach at Borlum Bay, Fort

Augustus (NH 085 385) on 22 July 1992 from depths of 0.10-0.38

m. Some of the nematode species found did not reoccur during the

1992-93 surveys, so we have taken this opportunity to include them

in this paper. They are identified in the descriptions as Borlum Bay

Table 1 Summary of sampling stations, core depths and sediment characteristics.

Core Depth

Station No’s Position Date (m) Sediment

Fort Augustus Bay (FA) 2-7 57°08..80° N 4°39.45° W_ 10/7/93 91-105 Organic matter (OM) floc, occasionally ferrugineous mixed with

grey siliceous sand

Deep South Basin(DSB) 8-13 57°12.55° N 4°34.55° W_11412/7/93 194-210 As above but with slight sulphurous smell

Foyers Plateau (FP) 14-19 57°15.80° N 4°29.60° W_ 12+13/7/93 150-161 As above

Deep North Basin(DNB) 20,22 57°16.40° N 4°29.10°W 13/7/93 211211 As above

River Oich (RO) 31,32 NH094 382 7/8/94 0.31 Sublittoral. River mouth.Algae and OM on surface, coarse sand below

33,34 NH094 382 7/8/94 0.20 Small stones and gravel

35 NH 094 382 7/8/94 0.33 Coarse sand and small pebbles

36 NH 094 382 7/8/94 0.17 Coarse sand

River Foyers (RF) 40-45 NH 211 495 7/8/94 0.14-0.39 Stagnant sublittoral. 50 m from river mouth. OM and grey siliceous

sand containing black streaks. Strong sulphurous smell

River Moriston (RM) 50 NH 211 495 8/8/94 0.15 100 m from river mouth Sand between boulders

(account of full data set 51,52 NH 211495 8/8/94 0.100.08 Coarse sand small pebbles

in prep.) 53 NH 211 495 8/8/94 0.20 Sand and gravel, some pebbles

54 NH 211 495 8/8/94 0.15 Coarse sand

55) NH 211 495 8/8/94 0.16 Sand and gravel

FRESHWATER NEMATODES FROM LOCH NESS

als

0 5km

R. Moriston

Caledonian

Canal

R. Tarff

Fig. 1 Map of Loch Ness showing sampling stations.

(1992 collection). Also, they have not been included in the ecologi-

cal part of this study which is based entirely on the 1992-93

samples.

EXTRACTION AND SLIDE PREPARATION

Nematodes were extracted by the Ludox centrifugation-floatation

technique (Persmark et al., 1992). However, after extracting the first

core it was found that the nematodes were still obscured by large

200m contour

R. Ness

R. Enrick 5S

R. Coiltie 200m contour

1. River Oich (RO)

2. Fort Augustus Bay (FA)

3. River Moriston (RM)

4. Deep South Basin (DSB)

. River Foyers (RF)

. Foyers Plateau (FP)

. Deep North Basin (DNB)

** Activated sludge treatment

* Significant septic tank input

amounts of organic matter of very similar density to that of the

nematodes. In order to reduce sample sorting time, it was decided to

subsample the remaining | cm core slices using the “Asko sample

splitter’ Elmgren (1973). Two hundred nematodes were picked out

from each core; if less than 200 nematodes were present in the first

subsample, further subsamples were extracted until the required

number was found. Extracted nematodes were then dehydrated to

glycerine by the Seinhorst method (Seinhorst 1962).

FIGURES, PHOTOGRAPHS AND MEASUREMENTS

Specimens were examined, figured and photographed with an Ol-

ympus BH-2 interference contrast microscope fitted with camera

lucida. The drawings are original and based entirely on Loch Ness

specimens. All absolute measurements of nematodes are given um.

SEDIMENT HORIZON

This can be taken as 0—1 cm unless a statement is made to the contary

in the descriptions.

ABBREVIATIONS USED

a = body length + greatest body width

b = body length + distance from anterior end to junction of

esophagus and intestine

b’ = body length + distance from anterior end to posterior

end of oesophageal glands

c = body length = tail length

(ei = tail length + body width at anus

IL, = total body length

MB = distance of median bulb from anterior end + oesopha-

geal length x 100

R = total number of body annules

Roes = number of annules in oesophageal region.

Rex = number of annules between anterior end of body and

excretory pore

Rv = number of annules between anterior end and vulva

Rvan = number of annules between vulva and anus

Ran = number of annules on tail

tail/V—a = tail length +distance between vulva and anus

Vv = distance of vulva from anterior end + body length x 100

We = distance of vulva from anterior end + distance of anus

from anterior end x 100

VL/VB = distance between vulva and posterior end of body +

body width at vulva

SPECIES LIST

The systematic classification of the order Tylenchida is based on that

of Fortuner, Geraert, Luc, Maggenti and Raski (1987-1988) ‘A

reappraisal of Tylenchina (Nemata)’. That of the order Rhabditida is

after Lorenzen (1981).

Order TYLENCHIDA Thorne, 1949 S* P

Family TYLENCHIDAE Orley, 1880

Aglenchus agricola (de Man, 1884)

Coslenchus polonicus Brzeski, 1982 +

Coslenchus sp. | =

Coslenchus sp. 2 + —

Filenchus cf. discrepans (Andrassy, 1954) +

Filenchus facultativus (Szczgiel, 1969) -

Filenchus thornei (Andrassy, 1954) +

Filenchus sp. 1 -

Filenchus sp. 2 =

Filenchus sp. 3 -

Lelenchus leptosoma (de Man, 1880) -_

Lelenchus sp. -

Malenchus acarayensis Andrassy, 1968

Malenchus bryophilus (Steiner, 1914)

Malenchus pachycephalus Andrassy, 1981 -

Malenchus pressulus (Kazachenko, 1975) —

+++

t++tteteteeettzret

F.R. WANLESS AND R. HUNTER

Miculenchus salvus Andrassy, 1959 —

Tylenchus rex Andrassy, 1979 -

Tylenchinae sp. +

Family ANGUINIDAE Nicoll, 1935

Ditylenchus sp. +

Family BELONOLAIMIDAE Whitehead, 1960

Geocenamus cf. nanus (Allen, 1955) +

Tylenchorhynchus sp. -

Family HOPLOLAIMIDAE Filipjev, 1934

Helicotylenchus pseudorobustus (Steiner, 1914) +

Family HETERODERIDAE Filipjev, 1934

Meloidogyne ardenensis Santos, 1968 +

Meloidogyne kralli Jepson, 1983 -

Family CRICONEMATIDAE Taylor, 1936

Criconema annuliferum (de Man, 1921) -

Criconema demani (Micoletzky, 1915) +

Criconema sp. =

Discocriconemella sp. -

Hemicycliophora sp. —

Ogma sp. +

Family TYLENCHULIDAE Skarbilovich, 1947

Paratylenchus sp. =

Family APHELENCHOIDIDAE Skarbilovich, 1947

Aphelenchoides sp. | +

Aphelenchoides sp. 2 +

Aphelenchoides sp. 3 +

Order RHABDITIDA Chitwood, 1933

Family RHABDITIDAE Chitwood, 1933

cf. Caenorhabditis sp. +

Family DIPLOGASTERIDAE Micoletzky, 1922

Diplogasteritus nudicapitatus (Steiner, 1914) +

Family CEPHALOBIDAE Filipjev, 1934

Eucephalobus oxyuroides (de Man, 1876) iP

Eucephalobus sp. +

* § =. Sublittoral, P = Profundal.

+++

FRESHWATER NEMATODES FROM LOCH NESS

DESCRIPTIONS OF SPECIES

Aglenchus agricola (de Man, 1884) Andrassy, 1954

(Fig. 2)

MATERIAL EXAMINED. Fort Augustus Bay: 92, 105 m; cores 2, 3;

299. Foyers Plateau: 150-161 m; cores 14-17; 222 44d. River

Foyers: 0.14—0.39 m; cores 40, 43, 45; 229, 2d.

HORIZON. Core 2 (2—3 cm), core 14 (3-4 cm), core 40 (2-3 cm).

FEMALES. (n=5).L=650 um (625-667); a= 31 (26.4-34.4); b=

6.6 (6.3-6.9); c = 3.3 (3.2-3.4); c’ = 14.5 (12.7-15.8); V =55.1 (53-

56); V’ = 77.6 (76.5-78.4); tail/V—a = 1.9 (1.7-2.0); tail = 194 um

(188-200).

Head continuous, not annulated. Body annules 1.6—1.8 um wide

at mid-body. Lateral fields with three lines; outer lines sometimes

appear double, whereas the inner line which is usually weakly

expressed, could be interpreted as two lines very closely set as they

appear as either gully or ridge-like (dependent on optical settings).

Spear moderately robust, 11.0—12.8 um long, basal knobs distinct

about 3.2 um wide. Oesophagus 92.7—101.7 um long; median bulb

ovoid with usually distinct valves at 44-50% of oesophageal length.

Vulva with flaps about 5.0 um long covering about three annules;

vagina curved anteriorly with swollen pyriform walls; anterior

Fig. 2 Aglenchus agricola (de Man, 1884). A-E, female. A, oesophageal

region; B, posterior end of reproductive system; C, tail; D, lateral field:

E, habitus; F, male spicular region. Scale bars a = 20 um, b = 40 um, c =

100 um.

genital branch 123—146 um long; spermatheca partly offset, usually

poorly defined, ovoid to elongate-ovoid, sometimes appearing

double, length variable 20—25.6 um; postuterine sac lacking.

MALES. (n=6). L=591.5 um (469-670); a = 38.9 (33-43); b (n

=) — 10.2, (0—01) 1c — 5. (229=3'3)2 ce = ior (la.3—=19):

gubernaculum (n= 5) = 6.4 um (6.3-6.4); spicules (n=5) = 14.7 um

(14-15.3); tail = 193.8 um (153-219).

Similar to female. Annules 1.6—1.96 um wide at mid-body. Spear

12—12.8 um long, with rounded knobs about 2.6 um wide. Oesopha-

gus 81.6-101 um long; median bulb ovoid with usually distinct

valves at 46-48% of oesophageal length. Testis 144—153.6 um long

occupying 24-25% of body length; cloacal lips tube-like, somewhat

asymmetrical; bursa adanal 32—38.4 um long with smooth or very

finely crenate margins.

DISTRIBUTION AND HABITAT. Cosmopolitan, in various soils, lives

on or near plant roots, in mosses and freshwater.

REMARKS. Morphometric measurements of this population fall

midway between those of two very similar species, A. agricola and

A. muktii Phukan & Sanwal, 1980. A. muktii, according to the

original authors, is distinguished from A. agricola ‘in having a non-

annulated lip region, 3 incisures in the lateral field, and a very long

filiform tail. (A. agricola has 3 annules in the lip region, | lateral

incisures and a tail which is not as filiform as in A. muktii).’ Of these

characters the first two are mistaken because in A. agricola the head

is not annulated and the lateral field is comprised of three lines. In

practice A. mucktii is distinguished from A. agricola by its very long

filiform tail. Tail length in A. agricola ranges from 134 to 179 um,

whereas in A. muktii the range is 178-276 um (Geraert & Raski,

1988). Female tail length of this population varies between 153-197

um. Brzeski (pers. comm.) notes tail lengths of A. agricola varying

from 134-208 um. In males of both species there would appear to be

a distinction between tail lengths and ratio c’. Tail length 140-169

um in A. agricola vs 178-330 um in A. muktii; ratio c° 13-15 vs 22—

25 in A. muktii. However the Loch Ness males bridge the gap in

respect of tail length 153-219 um and fall between the range of

values in the case of ratio c’, c° = 15.3-19.

The overlap in tail lengths and other morphometric measurements

suggest that the specific status of A. muktii is doubtful. This popu-

lation is therefore identified as A. agricola.

Coslenchus polonicus Brzeski, 1982

(Fig. 3)

MATERIAL EXAMINED. Deep South Basin: 204 m; core 9; 1.

River Foyers: 0.22 m; core 40; 1 &.

Core 40, (1-2 cm)

FEMALES. (n=2). L=635, 744 um; a= 41.5, 46.5; b=5.4, 6.2; c

=5.8,5.4;c°=11, 12; V=65.8, 64; V'=79.6, 80; Rex =47, 52; Roes

= 62, 63; Rv = 203; Rvan = 44, 49; Ran =55, 65; tail/V—a = 1.0; tail

= 110, 137.6 um.

Head more or less continuous with body contour, 6.4—6.7 um

wide with 4 very fine annules. Annules 2.3, 2.8 um at mid-body.

Lateral fields with 4 lines, the two inner lines closely spaced, thus

sometimes appearing as a single line. Cuticle with 18—20 longitudi-

nal ridges (difficult to count) excluding lateral fields. Spear robust

15, 13.4 um long; knobs rounded, 3.8, 3.2 um wide. Oesophagus

118, 120 um long; median bulb ovoid, valve at about 45% of

oesophageal length. Vulva, with lateral vulval membranes, 6.4 um

long or about 3 annules wide; vagina inclined anteriorly with thick,

somewhat pyriform, walls; postuterine sac small; spermatheca

HORIZON.

Fig. 3 Coslenchus polonicus Brzeski, 1982. A-F, female. A, oesophageal

region; B, lateral field; C, posterior end of reproductive system; D,

habitus; E, tail tip; F, tail. Scale bars a = 20 um, b = 100 pm.

elongate, 38, 41 um long, packed with sperms about 2.0 um in

diameter. Distance from vulva to anus 107—128 pm. Tail tip finely

rounded with annules in one specimen; rounded without discernible

annules in other.

DISTRIBUTION AND HABITAT. ‘Collected in several places in Po-

land and in Andreevskoe, Karelia, USSR, always in low peat soil, in

small number of specimens’ (Brzeski, 1987).

REMARKS. These nematodes were not originally identified as C.

polonicus because they have fewer cuticular ridges (18-20 vs 26 in

C. polonicus). However, Brzeski (pers. comm.) recognized these

females, and suggested they appeared to be C. polonicus. As there

are no other significant differences between these females and

descriptions of C. polonicus (Brzeski, 1982, 1987) it is assumed that

the lower number of cuticular ridges lies within the range of varia-

tion that might be expected in this species.

Coslenchus sp. |

(Fig. 4)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 4; 12

Foyers Plateau: 157 m; core 16; 1 8.

FEMALES. (n=2).L =520, 527 um; a= 23.2, 24.7; b=5.5, 5.4; c

= 6) OFC t—F/-2,1619- Vi 1641695 Vie a aIRexe— So /e-oaROES

= 42, 46; Rv = 136, 138; Rvan = 40, 37; Ran = 59, 53; tail/V—a =

0.92, 0.93; tail = 93.4, 86.4 um.

Head offset in one specimen; 7.0, 7.5 um wide with 4 annules.

Annules about 2.6, 2.7 um at mid-body. Lateral fields with four

lines. Cuticle with 14 longitudinal ridges (excluding lateral fields).

Spear moderately robust 11.5, 12.8 um long, knobs rounded, slightly

F.R. WANLESS AND R. HUNTER

Fig.4 Coslenchus sp. 1. A-E, female. A, oesophageal region; B, lateral

field; C, vulval region; D, habitus; E, tail. Scale bars a = 20 um, b = 100

um.

sloping posteriorly, 2.6, 3.0 um wide. Oesophagus 96.3, 94.7 um

long; median bulb ovoid; valve well developed at about 46% of

oesophageal length. Vulva with lateral vulval membranes 7.0, 7.6

um or 3 adjacent annules long; vagina perpendicular or slightly

inclined anteriorly, with thick walls; postuterine sac not evident;

spermatheca without sperm. Distance from vulva to anus 100, 120

um. Tail tip finely rounded.

REMARKS. Using Brzeski’s (1987) and Geraert & Raski’s (1988)

keys, these nematodes come close to C. andrassyi Brzeski, 1987, but

the mid-body annules are marginally too wide 2.6, 2.7 vs 2.1—2.5 um

in C. andrassyi, also the number of annules between the vulva and

the anterior end of the body are too low, 136, 138 vs 157-186 in C.

andrassyi.

Coslenchus sp. 2

(Fig. 5)

MATERIAL EXAMINED. River Foyers: 0.22 m; core 40; 12

HORIZON. Core 40, (1-2 cm).

FEMALE. (n=1).L=546 um; a=24; b=4;c=5.4;c° =7; V=65;

V°=79.5; Rex = 40; Roes = 63; Rv = 152; Rvan =41; Ran = 66; tail/

V—a = 1.1; tail = 102 um.

Head narrower than adjacent body; 7.7 um wide with 5

annules.Annules 2.7 um wide at mid-body. Lateral fields with 4

lines. Cuticle with longitudinal ridges and additional short ridges

distributed irregularly, giving an estimated total of 15-16 ridges

excluding lateral fields. Spear slender, 13.5 um long; knobs small,

FRESHWATER NEMATODES FROM LOCH NESS

Fig.5 Coslenchus sp. 2. A-E, female. A, oesophageal region; B, lateral

field; C, vulval region; D, tail; E, habitus. Scale bars a = 20 um, b = 100

um.

rounded about 2.6 um wide. Oesophagus 136 um long; median bulb

ovoid, valve well developed at 45.5% of oesophageal length. Vulva

with lateral vulval membranes, 7.6 um long or about 3 annules wide;

vagina slightly inclined anteriorly with thickened walls; postuterine

sac short; spermatheca empty. Distance from vulva to anus 91 um.

Tail damaged, but annules up to finely rounded terminus.

REMARKS. Although the tail of this specimen is damaged, it is

clearly different from the other species of Coslenchus found in Loch

Ness. Ratios effected by this damage are considered to be close to

measurements that would have been obtained had the specimen been

undamaged. It keys out close to C. pastor Andrassy, 1982, but the

morphometrics are a poor fit.

Filenchus cf. discrepans (Andrassy, 1954) Raski &

Geraert, 1987

(Fig. 6)

MATERIAL EXAMINED. Fort Augustus Bay: 98, 102 m; cores 5, 6;

222. Deep South Basin: 196, 204 m; cores 8, 10; 229. Foyers

Plateau: 158,160 m; cores 14, 15, 18; 3 9. Deep North Basin: cores

20, 22; 211 m, 2 2°. River Foyers: 0.25—-0.39 m; cores 41, 42, 44; 16,

329

HORIZON. Core 14, (1-2 cm).

FEMALES. (n= 10). L=429 um (337-526); a = 35 (27.0-45.8); b

= 5.3 (5.0-6.1); c=3.8 (3.44.5); c’ = 14.1 (12.8-17.6); V = 60 (57-

64); V~ = 82.3 (80.5—84); tail/V—a = 2.1 (1.7-2.5); tail = 109 um

(91.4-140).

Head 3.8—5.0 um wide at base. Annules 1.2 um (0.8-1.6) wide at

Fig.6 Filenchus cf. discrepans (Andrassy, 1954). A-E, female. A,

oesophageal region; B, lateral field; C, reproductive system; D, tail; E,

habitus. F, male spicular region. Scale bars a = 20 um, b = 10 um, c =

50 um.

mid-body. Lateral fields with two lines. Spear slender, 6.0—7.6 um

long with small basal knobs, about 1.2 um wide. Oesophagus 69-92

um long; median bulb somewhat fusiform, valve weak at 35-42% of

oesophageal length. Anterior genital branch 68-156 um long.

Spermatheca axial, oblong with rounded sperms about 1.3 um in

diameter. Postuterine sac 7.0—-13.4 um long. Vagina more or less

perpendicular to body axis.

MALE) (n= 1) 453m; a — 475 b— 510764556 = 45:

gubernaculum = 3.0 um; spicules = 13.8 um; tail = 101 um.

Similar to female, except body more slender with narrower lateral

fields. Testis 152 um long, occupying 33% of body length. Bursa

adanal, 19 um long.

DISTRIBUTION AND HABITAT. Widely distributed in Europe with

additional records from Mexico, Russia, USA (terrestrial soils) and

New Guinea (terrestrial and fresh water).

REMARKS. This population keys out to F: discrepans. The mor-

phology of the female reproductive system closely resembles that

figured for the type specimen, especially in respect of the oblong

spermatheca in females. However, the cephalic region is not makedly

narrower than the adjacent body region as figured by Andrassy

(1954) and described by Troccoli & Geraert (1995). Brzeski (pers.

comm.,) suggests that these nematodes show affinities to F. misellus

(Andrassy, 1958), but in F: misellus tail length (28—71 pm) and ratio

tail/V—a (0.7-1.6) are different (Brzeski, 1979); tail shape also

differs.

Filenchus facultativus (Szczygiel, 1969) Raski & Geraert,

1987

(Fig. 7)

MATERIAL EXAMINED. Fort Augustus Bay: 91, 98 m; cores 2, 5;

322,

HORIZON. Core 2, (2-3 cm).

FEMALES. (n=3).L=515 um (434-557); a= 36 (35-37.8); b=6.1

(6.0-6.3); c=4.0 (3.9-4.1); c’ = 13 (11.6-14.5); V = 63.2 (62-64.6);

V ‘= 84.8.(83.7-85.6); tail/V—a = 2.1 (2.0-2.2); tail = 127 um (105—

142).

Head about 4.6 um wide at base with 2-3 annules. Amphids

sinuous. Annules 1.7 um (1.5-1.9) at mid body. Lateral fields with

two lines. Spear slender 9.0 um (n = 2) long with small basal knobs,

about 1.2um wide. Oesophagus 83.8 um (73.6—91) long; median

bulb ovoid, valve weak at 39-42% of oesophageal length. Excretory

pore 62.4, 68.7 um (n= 2) from anterior. Anterior genital branch 145

um (127-159) long. Spermatheca axial or slightly offset, elongate

23, 26 um long with rounded sperms (n= 2). Postuterine sac 7.7—12

um long. Vagina 7.0—8.0 um long, more or less perpendicular to

body axis. Tail tip pointed to finely rounded.

DISTRIBUTION AND HABITAT.

restrial soils.

A widespread species found in ter-

REMARKS. F. facultativus is a variable species and these nema-

todes are within the range of measurements of other populations

described in the literature. However, the spear at 9.0 um long is at the

extreem range recorded for this species (6.0—9.0 um.). A range of

6.0-8.0 um is given by Raski & Geraert (1987) in their key to

Fig. 7 Filenchus facultativus (Szczygiel, 1969). A—G, female. A,

oesophageal region; B, head showing amphid; C, posterior end of

reproductive system; D, lateral field; E, reproductive system; F, tail

terminus; G, habitus. Scale bars a = 10 um, b = 30 um, c = 100 um.

F.R. WANLESS AND R. HUNTER

species of Filenchus, the same range is also given by Karegar &

Geraert (1988) in their summary of characters distinguishing F.

facultativus from other very similar species, whereas Torres &

Geraert (1996) record spear lengths of 7.5—9.0 um in a population

described from Argentina.

Filenchus thornei (Andrassy, 1954) Andrassy, 1963

(Fig. 8)

MATERIAL EXAMINED. Deep South Basin: 161 m; core 19, 1.

River Foyers: 0.22 m; core 40, 1 9. Foyers Plateau: 159 m; core 14,

Nice

HORIZON. Core 14, (2—3 cm); core 19, (1-2 cm); core 40, (2-3

cm).

FEMALES. (n= 2). L=997, 1003 um; a = 38, 44.7; b= 6.9, 6.5; c

=)5.0, 4c = 14255 182 Vi = 595562 735 Sata 0195)

1.16; tail = 131.5, 191.5 um.

Head about 7.6, 8.0 um wide at base with 3-4 annules; height

about 3.5 um. Amphids not observed. Annules 1.3—1.6 um wide at

mid-body. Lateral fields with four lines, inner lines very faint and

not always evident. Spear distinct, moderately slender, 13.4, 12 um

long with small basal knobs about 2.0 um wide. Oesophagus 144—

155 um long; median bulb more or less ovoid, valves moderately

distinct at 39.7, 42% of oesophageal length. Excretory pore

sclerotized at 112, 122 um from anterior. Anterior genital branch

432 um (n= 1); spermatheca 42 um long, offset. Vagina perpendicu-

lar to body axis about 13.5 um long. Tail tip finely pointed.

Fig. 8 Filenchus thornei (Andrassy, 1954). A-E, female. A, habitus; B,

posterior end of reproductive system; C, tail; D, head region; E,

oesophageal region. F, male spicular region. Scale bars a = 200 um, b =

30 pm, c = 20 pm.

FRESHWATER NEMATODES FROM LOCH NESS

MATE Gn) — O60) a— 442 bi—7 le = Sale = 128:

gubernaculum = 6.0 um; spicules = 22.6 um; tail = 188 um.

Similar to female; body slightly sinuous and curved distally, also

narrowed in region of cloaca — possibly an artefact as the body is

slightly twisted in this region. Bursa 38 um long, with crenate

borders; cloacal lips protruding. Tail tip finely rounded.

DISTRIBUTION AND HABITAT.

and freshwater habitats.

Cosmopolitan, found in damp soils

REMARKS. This species was recently redescribed by Brzeski (1997)

and these specimens fall well within the range of measurements

presented for F. thornei. The height of the head is perhaps slightly

lower, but this is not thought to be of any significance.

Filenchus sp. |

Fig. 9

MATERIAL EXAMINED. Foyers Plateau: 159 m; core 14, 2 9°. Deep

North Basin: 211 m; core 22, 1 2.

Core 14, (1-2 cm; 3-4 cm).

FEMALES. (n=3).L=531 um (439-615); a = 40.6 (39.7-41.8); b

= 6.0 (5.5-6.5); c = 3.4 (3.2-3.6); c” = 18.5 (15.8—20.2); V = 58.6

(57-60); V° = 83.3 (82.6-84.5); tail/V—a = 2.5 (2.2-2.7); tail = 157

uum (132-191.5).

Head 5.0-5.7 um wide at base with two annules. Amphids

sinuous. Annules about 1.6 um wide at mid-body. Lateral field (only

evident in two specimens) with four lateral lines, the inner two being

very faint and close to outers. Spear indistinct, slender, 9.6 um long

with small rounded knobs about 1.3 um wide. Excretory pore 64 um

HORIZON.

Fig.9 Filenchus sp. 1. A-F, female. A, oesophageal region; B, head

showing amphid; C, lateral field; D, tail tip; E, habitus; F, posterior end

of reproductive system. Scale bars a = 10 um, b = 100 um.

(56-73) from anterior. Oesophagus 85 um (74—94.2) long; median

bulb with poorly developed valves at 40.4% (35-45.6) of oesopha-

geal length. Anterior genital branch 163 um (115-198) long.

Spermatheca offset, 27.7 um (22.4—32) long. Postuterine sac 8.9 um

(8.3—-9.6) long. Vagina 7.0—7.7 um long, gently inclined anteriorly

with thickened walls. Tail terminus pointed to finely rounded.

REMARKS. The specific identity of these females is uncertain.

They show affinities with F facultativus described above and may

be conspecific. However, in these nematodes the spear is slightly

longer and the walls of the vagina are slightly thickened.

Filenchus sp. 2

(Fig. 10)

MATERIAL EXAMINED.

RENAE een (hel) le O20 mea At elle D929 — syn 1910):

V = 65.8; V’ = 80; tail/V—a = 1.0; tail = 108 um;

Head 5.7 tum wide at base. Amphids not observed. Annules fine,

1.3 um wide at mid-body. Lateral field with four lines, inner two

faint. Spear indistinct, 9.3 um long with small basal knobs about 1.3

uum wide. Excretory pore 82.5 um from anterior. Oesophagus 119

um long; median bulb ovoid, valves weak at 40.7% of oesophageal

length. Anterior genital branch 175 um long; spermatheca axial,

about 22 um long. with rounded sperms about 1.9 um in diameter.

Postuterine sac 6.4 um long, occupying 41% of corresponding body

diameter. Vagina about 5.8 um long, perpendicular to body axis. Tail

terminus finely rounded.

Foyers Plateau: 157 m; core 14, 1 9.

Fig. 10 Filenchus sp. 2. A-E, female. A, oesophageal region; B, habitus;

C, lateral field; D, tail; E, posterior end of reproductive system. Scale

bars a = 20 um, b = 100 um.

REMARKS. This nematode is close to F. butteus (Thorne & Malek,

1968) Brzeski, 1997, but the tail is longer and differs in shape.

Filenchus sp. 3

(Fig. 11)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 3, 1d.

MALE. (n=1).L=434 um; a=35.7;b=6.4; c=4.4; c’ = 10.3;

gubernaculum = 5.8 um; spicules = 15 um; tail = 99 um.

Head 5.0 um wide at base with 3-4 annules. Amphids not

observed. Annules fine about 0.9 um at mid-body. Lateral field with

four lateral lines, hardly evident. Spear moderately slender, 11.2 um

long with small basal knobs 1.9 um wide. Excretory pore 61.4 um

from anterior. Oesophagus poorly preserved, about 68 pm long;

median bulb ovoid, valves not observed, centre at about 59% of

oesophageal length. Testis 138 um long, occupying 32% of body

length, bursa adanal, about 19 um long, with fine crenated margins.

Tail tip pointed.

REMARKS. No conclusion can be reached on the specific identity

of this specimen. It is close to Fthornei, but differs by its shorter

body length, spicules and tail.

Lelenchus leptosoma (de Man, 1880) Raski & Geraert,

1985

(Figs 12, 36B)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 3; | 9. Deep

South Basin: 194 m; core 13; 1 2. Foyers Plateau: 159, 160 m; cores

1 ee a

FEMALES. (n= 5). L = 546 (487-583) um; a = 46.6 (40-49.8);

D='951(4.9=6)3) se = Sal (273539) ie = 242 822 —27/).9) 2 Ve

Fig. 11 Filenchus sp. 3. A-E, male. A, oesophageal region; B, lateral

field; C, spicular region; D, habitus; E, tail and spicular region. Scale

bars a = 20 um, b = 100 um.

F.R. WANLESS AND R. HUNTER

Fig. 12 Lelenchus leptosoma (de Man, 1880). A-E, female. A,

oesophageal region; B, head showing amphid; C, posterior region; D,

habitus; E, reproductive system. Scale bars a = 20 um, b = 10 um, c =

60 um, d = 100 um.

(4854.8); V° = 77.6 (75.5—78); tail/V—a = 2.2 (1.9-2.5); tail = 180

um (156-215).

Cephalic region narrowed dorso-ventrally. Cuticle smooth by

light microscopy. Amphids sinuous. Lateral field absent. Spear

slender, 7.7—9.3 um long with small rounded knobs about 1.4 um

wide. Oesophagus 88-108 um long; median bulb spindle-shaped

with valves at 41-43% of oesophageal length. Anterior genital

branch 96.7—121.5 um long. Spermatheca more or less bispherical

with rounded sperm (absent in one specimen) about 1.3 um in

diameter. Postuterine sac not evident in any specimen, possibly

collapsed. Vagina about 5.5 um long with thin walls, curved

anteriorly.

DISTRIBUTION AND HABITAT. Cosmopolitan; soils and freshwater.

REMARKS. These specimens, for the most part, agree with the

description and morphometrics presented in the literature (Raski &

Geraert, 1985). Minor differences slightly extending the range of

several ratios are not considered to be of any significance.

Lelenchus sp.

(Figs 13, 36A)

MATERIAL EXAMINED. Deep South Basin: 195, 204 m; cores 9, 11;

2 29. Foyers Plateau: 157, 159 m; cores 14, 16; 146,12.

FEMALES. (n=3).L=620 (512-763) um; a=47.4 (42-54.5); b=

6.0 (5.5-6.5); c = 3.5 (3.3-3.8); c” = 22.8 (21.4-24); V =57.6 (57—

58); V’ = 81 (80-82); tail/V—a = 2.1 (1.8—2.4); tail = 173 (154-200)

um.

FRESHWATER NEMATODES FROM LOCH NESS

Fig. 13. Lelenchus sp. A-C, F, female. A, oesophageal region; B, tail; C,

habitus; F, vulval region. D, E, male. D, habitus; E, spicular region.

Scale bars a = 20 um, b = 30 um, c = 100 um.

Cephalic region narrowed dorso-ventrally. Cuticle smooth by

light microscopy. Lateral field absent. Spear slender, poorly defined,

9.0-9.3 um long (n = 2) with small rounded knobs about 1.3 um

wide. Oesophagus 92-118 um long; median bulb spindle-shaped,

valves very indistinct, at about 48% of oesophageal length. Anterior

genital branch 175-250 um long. Spermatheca elongate ovoid

without sperm. Egg present in two specimens about 62 um long by

10 um wide. Postuterine sac about 11.5 um long or 80% of vulval

body width (n= 2). Vagina more or less perpendicular, about 7.7 um

long with slightly thickened walls.

MAGE (i |), = 920 um; a= 58; b= 6:4; c= 4.5; ci = 19.5;

spicules = 23 um; gubernaculum = 6.5 pm; tail = 203 um.

Similar to female except body longer and more slender. Spear

indistinct, about 10 um long, with small, possibly deformed, rounded

knobs. Oesophagus 145 tum long; median bulb spindle-shaped with

indistinct valves at 40% of oesophageal length. Testis 300um long,

occupying 32.6% of body length; bursa adanal about 38 um long

with smooth margins.

REMARKS. These nematodes are possibly conspecific with L.

leptosoma, but for the present they are treated as a separate species.

In females the vagina walls are thicker and generally more pro-

nounced. The male differs from males of L. leptosoma described in

the literature by its greater body length and longer, strongly curved,

almost bent spicules.

Malenchus acarayensis Andrassy, 1968 Geraert & Raski,

1986.

(Fig. 14)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 3, | 2. Foy-

ers Plateau: 159, 160 m; cores14, 15, 32°. River Foyers: 0.39 m;

core 42, 12.

HORIZON. Core 14 (2-3 cm).

FEMALES. (n=5). L = 352 um (323-373); a = 24.7 (23-26.6); b

(n=3) = 4.4 (4.3-4.6); c = 4.4 (4.0-4.9); c” = 9.1 (8-10.1); V = 63.3

(61-66.4); V° = 80.8 (79-82); tail/V—a = 1.5 (1.25—1.73). tail = 78.9

um (71.7—92).

Head narrow, about 4.0—-5.0 um at base, with very fine annules.

Amphids not clear. Annules 1.0—1.3 um at mid-body. Lateral fields

with two lines appearing smooth or crenate, depending on fine

focus; originating between 6.4—7.0 um posterior to spear base,

ending at 27-33% of tail length. Spear fine, especially conus, 8.3—

9.0 um long, knobs small, sloping posteriorly, about 1.3 um wide.

Excretory pore 56—66 um from anterior. Oesophagus 73.6—86 um (n

= 3) long; median bulb ovoid, valve hardly evident, but positioned at

44-47.8% of oesophageal length. Anterior genital branch 101—112

um long. Spermatheca usually rounded, ovoid in one specimen, 9.6—

12.8 um wide. Postuterine sac 6.4 um long or 43% of vulval body

width (n=1), collapsed in other specimens. Vagina with thickened

walls, perpendicular to body axis or slightly tilted anteriorly. Dis-

tance from vulva to anus 45—56.9 um. Tail tip slightly to strongly

curved, tapering to a fine point, but not thread-like.

DISTRIBUTION AND HABITAT. Described from N. and S. America,

EGE

Fig. 14 Malenchus acarayensis Andrassy, 1968. A—-F, female. A,

oesophageal region; B, habitus; C, posterior end of reproductive system;

D, tail; E, F, lateral field at different focal levels. Scale bars a= 10 um,

b=50 um.

India, Malawi, Holland, Poland and Belgium. Terrestrial, found in

various soils, forest litter and sand dunes.

REMARKS. These specimens are all identified as M. acarayensis

using Geraert & Raski’s key (1986); furthermore they compare well

with the range of measurements presented for M. acarayensis.

However, they possibly differ by the more anterior origin of the

lateral field, 6.4-7.0 um behind spear base in these females, com-

pared with 11-15 um ina Spanish population (Gomez Barcina et al.,

1992). Other populations are more difficult to compare in the same

manner as the start of the lateral field is given as originating at a level

of 1/3 to about middle of the oesophageal procorpus (Geraert &

Raski, 1986).

Malenchus bryophilus (Steiner, 1914) Andrassy, 1980

(Fig. 15)

MATERIAL EXAMINED. Fort Augustus Bay: 91, 105 m; cores 2, 7,

19,236. Foyers Plateau: 158, 160 m; cores 15, 18, 12, 1d. River

Oich: 0.31 cm; core 31, 222.

Core 2 (1—2 cm) Core 31 (3-4 cm).

FEMALES. (n=4). L = 349.5 um (334-364); a= 21.3 (18-24.7); b

(n=2)=4.5 (4.44.5); c=4.9 (4.6-5.1); c°=7.9 (7.1-8.6); V = 64.4

(63.8-65.3); V° = 81.2 (63.8-65.3); tail/V—a = 1.4 (1.2-1.5); tail 72

um (69.1—73.6).

HORIZON.

Fig. 15 = Malenchus bryophilus (Steiner, 1914). A, C-G, female. A,

oesophageal region; C, tail; D, habitus; E, posterior end of reproductive

system: F, G, lateral field at different focal levels. B, H, male. B, head

region showing amphid and origin of lateral field in respect of spear

knobs; H, spicular region. Scale bars a = 10 um, b = 60 um.

F.R. WANLESS AND R. HUNTER

Head slightly narrowed, about 5 um wide at base with 4-5

annules. Amphids not clear. Annules 1.4-1.5 um at mid-body.

Lateral field with two lines appearing smooth or crenated depending

on fine focus; originating at level of spear knobs to two annuli

posterior, or 0Q—2.5 um from spear knobs, ending between 25-35% of

tail length. Spear fine, especially conus, 9-9.6 um long, knobs

small, about 2.56 um wide. Excretory pore 61—71 um from anterior.

Oesophagus 78.8 tum long (n = 2); median bulb ovoid, valve poorly

developed, at 45-50% of oesophageal length (n = 2). Vulva sunken.

Vagina perpendicular or slightly tilted anteriorly with thickened

walls; anterior genital branch 96—113 um long; spermatheca rounded

to ovoid, packed with sperms. Postuterine sac about 5 um long (n=

2), possibly collapsed in other females. Distance from vulva to anus

48.6—57 um. Tail tip finely rounded to pointed.

MALES. (n=3).L =362 um (337-379); a= 21.1 (20.6—21.5); b=

4.7 (4.4-5.0); c =4.5 (4.14.7); c’ = 7.3 (6.6-8.3); gubernaculum =

5.5 um (5.3-5.76); spicules = 16.5 um (16-17); tail = 80.6 um

(71.6-91)um.

Similar to female. Head narrowed, about 5 um wide at base with

5 fine annules. Amphids sinuous. Body annules 1.4—1.5 um at mid-

body. Spear 9.0—-9.6 um long. Oesophagus 71-85 um long; median

bulb ovoid, valve weak, at 48.6-53.8% of oesophageal length. Testis

189-213 um long, occupying 41-56% of body length. Cloacal lips

pronounced; bursa adanal, 27—32 um long, with smooth margins.

DISTRIBUTION AND HABITAT. Cosmopolitan, occurs in soil, near

or in roots; in humus, forest litter and moss.

REMARKS. The morphometrics and morphology agree for the

Fig. 16 Malenchus pachycephalus Andrassy, 1981. A—-F, female. A,

oesophageal region; B, anterior region showing amphid and origin of

lateral field; C, posterior end of reproductive system; D, tail; E, lateral

field; F, habitus. Scale bars a = 20 um, b = 10 um, c = 100 pm.

FRESHWATER NEMATODES FROM LOCH NESS

most part with the original description. Minor differences include

slightly lower values for tail length, tail/V—a and ratio c’. These

measurements however, fall within the range of measurements of

females of M. bryophilus examined by Brzeski (pers. comm.).

Malenchus pachycephalus Andrassy, 1981

(Fig. 16)

MATERIAL EXAMINED. Foyers Plateau: 157 m; core16, 1 &.

FEMALE. (n=1).L=497 um; a= 20; b=4.9;c=6.1; c° =7.0; V

= 69; V’ = 81.6; tail/V—a = 1.0; tail = 81.2 um.

Head slightly offset, 6.4 um wide at base with 4—5 fine annules.

Amphids not evident. Annules 2.1 um wide at mid-body. Lateral

field with two lines appearing smooth or strongly crenate depending

on fine focus; originating at 3 annules or 4.0 um anterior to spear

knobs; terminating at about 46% of tail length. Spear 12 um long

with knobs about 2.4 um wide. Excretory pore 93 um from anterior,

almost at level of posterior margin of basal bulb. Oesophagus 100

um long; median bulb slender ovoid; valve hardly evident located at

44% of oesophageal length. Phasmid like structure not seen. Ante-

rior genital branch 170 um long. Spermatheca bilobed 24 um long

with rounded sperms about 1.3 tm in diameter. Postuterine sac 9.6

um long. Vagina slightly tilted anteriorly with thickened walls.

Vulva — anus distance 76.8 um. Tail tip tapering to a fine point.

DISTRIBUTION AND HABITAT. Recorded from USA, Spain, Hun-

gary and Bulgaria; occurs in soil around plant roots.

REMARKS. Using the keys of Andrassy (1981) and Geraert &

Raski (1986), this specimen can be identified as M. pachycephalus,

but the anterior part of the oesophagus is not distinctly longer than

the posterior as described in the original description, however, this

feature is not evident in the accompanying illustration of the anterior

end (Andrassy, 1981: Fig. 13c). It also differs in having a longer tail

81 um vs (56-72); this range includes measurements from the

original description and a Spanish population (Gomez Barcina ef al.,

1992).

Malenchus pressulus (Kazachenko, 1975) Andrassy, 1981

(Fig. 17)

MATERIAL EXAMINED. Deep South Basin: 204 m; core 9, 1 9.

BEMALE. (0=1). 1 = 460 um; a= 25; b=5.1;c =5.9; ¢ =8.1; V

= 66.9; V = 80.5; tail/V—a = 0.95; tail = 77.7.

Head not offset, 7.0 um wide at base with 3-4 minute annules at

focal level of dorso-ventral margin, otherwise appearing smooth by

light microscopy. Amphids sinuous. Annules 1.5 um at mid body.

Lateral fields with two lines appearing smooth or crenate depending

on fine focus; originating at level of spear knobs, terminating at

about 32% of tail length. Spear delicate, 11.8 um long with spear

knobs 2.5 um wide. Excretory pore 86 um from anterior at level of

posterior margin of basal bulb. Oesophagus 89 um long; median

bulb slender ovoid, valve not evident; centre of median bulb located

at about 51% of oesophageal length. Phasmid like structure located

at 14 annules anterior to vulva. Anterior genital branch 170 um long.

Spermatheca elongate ovoid, 19 um long with indistinct rounded

sperms about 1.3 wm in diameter. Postuterine sac 12.8 um long.

Vagina slightly tilted anteriorly, with thickened walls. Vulva slightly

sunken. Vulva — anus distance 74.5 um. Tail tip pointed.

DISTRIBUTION AND HABITAT. Originally described from Russia in

soil of a coniferous forest with an additional record from soil around

grass roots (Andrassy, 1981).

Fig. 17 Malenchus pressulus (Kazachenko, 1975). A—D, female. A,

oesophageal region; B, posterior end of reproductive system; C, tail; D,

habitus. Scale bars a = 20um, b = 100 um.

REMARKS. This single female largely agrees with the redescription

of M. pressulus by Andrassy (1981) minor differences in ratios c, c”

and tail/V—a are not considered to be of specific importance.

Miculenchus salvus Andrassy, 1959

MATERIAL EXAMINED. Fort August Bay: 91, 92, 98 m; cores 2, 4,

5,322, 1d. Deep South Basin: 197 m; core 12, 2 29. Foyers Plateau:

157-160 m; cores 14-16, 18, 629, 1d.

REMARKS. These specimens were described and figured by Wanless

& David (1997).

Tylenchus rex Andrassy, 1979

(Figs 18, 36D)

MATERIAL EXAMINED. Deep South Basin: 194 m; core 13, 1d.

MATE (ni 1)) = 97 AP ims a — 362351D: 107 G— Onl ioa—1910?

gubernaculum = 5.9 uum; spicules = 24 um; tail = 160 um.

Head 8.5 um at base with 4-5 annules. Amphid opening a gently

curved slit. Annules distinct about 2.1 um at mid-body. Lateral field

with four lines, outers weakly crenate. Spear robust, 20.5 um long,

with large basal knobs 3.5 um wide. Excretory pore 130 um from

anterior. Oesophagus 154 um long; median bulb ovoid, valves

distinct at 47.5% of oesophageal length. Testis 262 tm long occupy-

ing 27% of body length. Bursa adanal about 38 um long with

crenated margins. Anterior margin of cloacal lips with two spicate

projections (arrowed, fig.36D). Tail tip rounded.

DISTRIBUTION AND HABITAT. The type population was found in

moss, Ben Hedi, Scotland. Also known from Poland and Australia.

REMARKS. This specimen largely agrees with the original descrip-

tion except that the basal bulb is more or less elongate rather than

pyriform; the outer lateral lines of the lateral field are weakly crenate

G a

iS mah re

SU Ye

Fig. 18 Tylenchus rex Andrassy, 1979. A-F, male. A, anterior region; B,

oesophageal region; C, head region showing amphid; D, lateral field; E,

tail and spicular region; G, spicular region. Scale bars a = 20 um, b = 50

um, c = 10 um, d= 100 um.

rather than smooth; the tail is longer (160 um vs 130-140 um); ratio

c is lower (6.1 vs 6.3-6.6) whereas c’ is slightly higher (9.0 vs 7-8).

However, these ratios and tail length fall within the range of meas-

urements given for T. rex by Brzeski (1996).

Tylenchinae sp.

(Figs 19, 36C)

MATERIAL EXAMINED. Foyers Plateau: 159 m; core 14; 1 9. River

Foyers: 0.39 m; core 42, 1 8.

Core 14, (1-2 cm).

FEMALES. (n= 2). L= 468, 520 um; a= 43.3, 40.6; b= 4.5, 4.7;

= 4.7, 5.0; c° = 12, 12.5; V =65, 62; V’ = 80; tail/V—a = 74.8, 94; tail

= 99.2, 104 um.

Head not set off, about 6.6 um wide at base. Amphid apertures not

seen. Annules fine about 0.8 um at mid-body. Lateral fields narrow

with two lines. Spear moderately robust, 11.5 um long with rounded

basal knobs 2.5 um wide; shaft slightly longer than conus. Oesopha-

gus 103, 110 um long; median bulb ovate, valve at about 45% of

oesophageal length. Excretory pore 71.4, 78.7 um from anterior.

Anterior genital branch 115, 109 um long. Spermatheca empty.

Postuterine sac not seen, possibly collapsed. Vagina about 6.0 um

long, slightly inclined or curved anteriorly. Tail elongate, not fili-

form; tip rounded.

HORIZON.

REMARKS. ‘The generic position of these nematodes is uncertain.

They are placed in Tylenchinae because they are close to Tylenchus

F.R. WANLESS AND R. HUNTER

Fig. 19 Tylenchinae sp. A-E, female. A, oesophageal region; B, tail; C,

lateral field; D, vulval region; E, habitus. Scale bars a = 10 um, b = 100

um.

Bastian, 1865 in which the anterior conus and posterior shaft of the

spear are about equal in length. However, the presence of two lines

in the lateral fields is not compatible with the generic definition of

Tylenchus, which is characterized by the presence of four lines

Additional material and SEM studies are required in order to resolve

the status of these interesting nematodes.

Ditylenchus sp.

(Fig. 20)

MATERIAL EXAMINED. _ River Foyers; 0.39 m; core 44, 1d.

MALE. (= 1). L=754 um; a=58.9:b=5.7; ¢ =74ce = 9:5;

spicule = 16 um; gubernaculum = 5.2 um; tail = 102 um.

Head 4.2 um wide, annuli not seen. Cephalic framework weak.

Lateral fields with five lines at mid-body, apparently four lines on

anterior and posterior regions, but for the most part difficult to

observe. Body annuli fine, less than 1.0 um wide. Spear delicate, 7.6

uum long; cone shorter than shaft; knobs rounded about 2.0 um wide.

Excretory pore 91 um from anterior. Oesophagus 131 pm long;

median bulb oval with distinct valve at 39.6% of oesophageal

length; basal bulb and isthmus indistinct. Caudal alae about 30.7 um

long, 30% of tail length. Tail terminus finely rounded.

REMARKS. Ditylenchus species are rather uniform and do not

show many characters helpful for identification (Brzeski, 1991b).

This single male cannot be named with any confidence and addi-

tional specimens, particularly females, are required for proper

identification.

FRESHWATER NEMATODES FROM LOCH NESS

Fig. 20 Ditylenchus sp. A-D, male. A, oesophageal region; B, spicular

region; C, habitus; D, lateral field. Scale bars a = 20 um, b = 10 pm, c =

100 pm.

Fig. 21 Geocenamus cf. nanus (Allen, 1955). A-C, female. A,

oesophageal region; B, tail; C, habitus. Scale bars a = 20 um, b = 100

um.

Geocenamus cf. nanus (Allen, 1955) Brzeski, 1991

(Fig. 21)

MATERIAL EXAMINED. River Oich: 0.31 m; core 31, 1 9.

RENAL Ents) — OM a —20.5.1b) Ose — llc = Sr:

V = 54; tail/ V—a = 0.24; tail annules 54; tail = 44.8 um.

Head slightly narrower than adjacent body with 5 or 6 fine

annules; labial frame work weak. Spear 12 um long with backward-

sloping knobs, 2.5 um wide. Cuticle annules fine and moderately

distinct, about 0.9 um wide at mid-body. Lateral fields with 6 lines

at mid-body, inner lines faint and not always evident throughout the

entire length of the lateral fields. Deirid at level of excretory pore.

Excretory pore 77 um from anterior. Oesophagus (possibly con-

tracted) 80 um long; median bulb ovoid, valve weak at 49.5% of

oesophageal length. Vulva and vaginal structures poorly defined.

Genital branches stretched out; anterior and posterior branches do

not extend into the oesophageal or anal regions respectively. Phasmids

at 40% of tail length. Tail annules continue almost to terminus.

DISTRIBUTION AND HABITAT. Originally described from the

Netherlands with further records from Iran, Turkey, Belgium, and

Poland. Terrestrial, in soils around roots of various plants (Saltukoglu

et al., 1976); also in meadow and peat soils (Brzeski, 199 1a).

REMARKS. Provisionally identified as G. nanus on account of the

number of tail annuli and the value of ratio c*. Other very similar

species have fewer tail annules and ratio c’ is generally lower, but

observed values overlap. For a discussion on relationships between

G. nanus (Allen, 1955), G. alboranensis (Tobar Jimenéz, 1970) and

G. microdorus (Geraert, 1966) see Brzeski (199 1a).

Tylenchorhynchus sp.

MATERIAL EXAMINED. Fort Augustus Bay: 105 m; Core 7, |

juvenile. Foyers Plateau: 157 um; core 16, | juvenile.

Helicotylenchus pseudorobustus (Steiner, 1914) Golden,

1956

(Fig. 22)

MATERIAL EXAMINED. Fort Augustus Bay: 91 m; core 2, | °. River

Oich: 0.33 m; core 35, 1 9.

FEMALES. (n= 2). L= 800, 830 um; a= 26.6, 34; b = 6.8, 6.9; b”

=) Sali 49: 257s 6e— Load = OONO2 tally Sen 22.4. tm:

Head hemispherical 6.4, 7.0 um wide with 4-5 annules. Spear

27.5, 26.8 um long, with distinct knobs about 6.0 um wide. Cuticle

annules about 1.6 um wide at mid-body. Lateral field with 4 lines,

outers weakly crenate. Dorsal oesophageal gland opening 9.6, 10.0

uum behind spear knobs. Excretory pore 110 um from anterior.

Oesophagus 160, 162 um long. Spermathecae offset without sperm.

Phasmid 6—8 annules before anus. Tail with 12 ventral annules,

dorsally convex and terminating in a long projection that has a fine

irregular outline.

DISTRIBUTION AND HABITAT. Cosmopolitain; found in soils and

also fresh water.

REMARKS. These females are in close agreement with descriptions

of H. pseudorobustus by Sher (1966) and Siddiqi (1972).

Meloidogyne ardenensis Santos, 1968

(Fig. 23)

MATERIAL EXAMINED. Borlum Bay (1992 collection): 0.18 m;

core 4, second-stage juvenile.

Fig. 22 Helicotylenchus pseudorobustus (Steiner, 1914). A-C, female.

A, anterior region; B, tail; C, habitus. Scale bars a= 10 um, b = 100 pm.

Fig. 23. Meloidogyne ardenensis Santos, 1968. A—D, second stage

juvenile. A, anterior region; B, tail; C, lateral field; D, habitus. Scale

bars a= 10 um, b= 50 pm.

F.R. WANLESS AND R. HUNTER

SECOND-STAGE JUVENILE. (n= 1). L=403 um; a= 26.8; b=6.1;

c = 9.8; c° = 4.0; tail 40.9 um..

Head not offset from body with two faint post labial annules.

Cuticular annulation distinct but fine, about 0.9 um at mid-body.

Lateral field with four lines, the outer ones finely crenate and giving

the impression of two vague additional lines in the post oesophageal

region. Spear slender, 11.5 um long with small backward-sloping

knobs. Dorsal gland opening 2.5 um behind spear knobs. Length of

oesophagus (from anterior to base of median bulb) 65.3 um; median

bulb ovoid. Excretory pore 75.8 um from anterior. Hemizonid not

seen. Tail with rounded tip; phasmid not seen; Length of hyaline tail

terminus 12.1 um.

DISTRIBUTION AND HABITAT. Reported from Belgium, France,

Germany, Poland, Russia and the United Kingdom (Karssen & Van

Hoenselaar, 1998). Infective, migratory second-stage juveniles have

been found in a wide variety of soils including dunes; adults are

commonly associated with roots of vegetable and herbaceous crops

(Jepson, 1987).

REMARKS. Although the hemizonid and phasmids could not be

seen, this second-stage juvenile is otherwise in close agreement with

descriptions of M. ardenensis.

Meloidogyne kralli Jepson, 1983

(Fig. 24)

MATERIAL EXAMINED.

stage juvenile.

Foyers Plateau: 159 m; core 14, second-

SECOND-STAGE JUVENILE.

= 6.7; c° = 6.8; tail 64.6 um.

@=1)sb =435 ima — Seb 7c

Fig. 24 Meloidogyne kralli Jepson, 1983. A—D, second stage juvenile. A,

anterior region; B, lateral field; C, tail; D, habitus. Scale bars a = 10 um,

b = 100 um.

FRESHWATER NEMATODES FROM LOCH NESS

Head not offset from body, with labial cap and one postlabial

annule. Cuticular annulation distinct but fine, about 0.9 um wide at

mid-body. Lateral fields with four lines. Spear slender, 11.5 um long

with small backward-sloping knobs. Dorsal gland opens 5.0 um

behind spear knobs. Length of oesophagus (from anterior to median

bulb) 61.4 um; median bulb more or less ovoid; excretory pore 81.2

um from anterior. Hemizonid immediately anterior to excretory

pore. Phasmid not seen. Tail terminus tapers sharply with a short

terminal portion ending in a finely rounded tip. Length of hyaline

tail terminus 16.5 um.

DISTRIBUTION AND HABITAT. Recorded from Estonia, Russia, Po-

land and the United Kingdom, all in wet sandy, peat and silt soils

(Karssen & Van Hoenselaar, 1998).

REMARKS. The present second-stage juvenile agrees well with

descriptions of the species.

Criconema annuliferum (de Man, 1921) Raski & Luc,

1984

(Fig. 25)

MATERIAL EXAMINED.

BEMAUE (n= 1). =598 um: a= 11.2;b=3.9;c=25; V=88;R

= 60; Rex = 20; Rvan = 5; Ran = 4; VL/VB = 1.4; tail = 22.4 um.

Labial disc low. Head region offset, collar-like. Annule two

Foyers Plateau: 160 m; core 15, 1 9.

Fig. 25 Criconema annuliferum (de Man, 1921) A-C, female. A,

habitus; B, anterior region; C, posterior region. Criconema sp. D, E,

female. D, anterior region; E, posterior region. Scale bars a = 100 um,

b = 30 um.

slightly smaller and narrower than annule three, both distinctly

smaller than annules one and four. Annules 10.8 um wide at mid-

body, outer edges smooth, rounded or slightly retrorse; interspaces

sometimes filled with fine detritus. Spear straight, 101 um long;

knobs 11.5 um wide, anterior surfaces indented. Vulva closed,

anterior lip well developed, overlapping entire posterior lip. Ovary

not reflexed, oocytes arranged in two rows. Spermatheca not

observed. Anus inconspicuous. Tail terminus peg-like.

DISTRIBUTION AND HABITAT. Throughout Europe, associated with

plant roots in both dry and wet soils.

REMARKS. This nematode agrees with descriptions and

morphometrics presented for C. annuliferum.

Criconema sp.

(Fig. 25)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 3, | 2.

FEMALE) ‘(= 1) 91 = 670iumsa= 12.3: b= 4.15c= 21-8; V —90;

R = 65; Rex = 21; Rvan = 3; Ran=5; VL/VB = 1.5; tail = 30.7.um.

Labial disc low. Head region offset, collar-like; annules two

and three of equal size, distinctly smaller than annules one and

four. Body annules 10.2 tm wide at mid-body, outer edges

smooth, rounded or slightly retrorse; interspaces with fine scatter-

ing of detritus. Spear 120 um long; shaft gently curved, conus

straight; knobs 12.2 um wide, anterior surfaces indented. Vulva

closed, anterior lip well developed, overlapping entire posterior

lip. Ovary poorly preserved; oocytes arranged in two rows.

Spermatheca not evident. Anus conspicuous. Tail terminus bifur-

cate.

REMARKS. Similar to C. annuliferum and keys out to that species

(de Grisse & Loof, 1965) but the spear is longer with a gently curved

shaft, possibly an artefact; the tail terminus is bifurcate, also the 2nd

and 3rd head annules are equal in size.

Criconema demani (Micoletzky, 1915) Raski & Luc, 1984

(Fig. 26)

MATERIAL EXAMINED. River Oich: 0.31, 0.18 m; cores 31, 33,22

FEMALES. (n=2).L=401, 417 um; a= 8.3, 8.7; b = 3.8, 4.0; c=

10.6, 10; V = 84.5, 82.2; R=73, 71; Rex (n= 1) =20; Rvan =5; Ran

=) DAVIN NES! —Alefis CALs her Ree 2

Labial disc low. Head region offset with second annule broader

than first. Annules 5.8, 6.4 um wide at mid-body; outer edges

smooth, rounded or moderately retrorse; with fine, scattered, foreign

detritus but nevertheless margins apparently minutely crenate. Spear

70.4, 71.6 um long; knobs 8.3, 8.6 um wide, anterior surfaces

indented. Vulva closed; anterior lip well developed, overlapping

entire posterior lip. Ovary reflexed distally. Spermatheca poorly

preserved, without sperms. Anus inconspicuous. Tail with terminus

straight or curved dorsally.

DISTRIBUTION AND HABITAT. Belgium, Denmark, Netherlands and

North America. Terrestrial, in moist soils and alongside riverbanks,

ditches and lake margins.

REMARKS. Keys out to C. demani and generally fits in well with

descriptions of the species (Taylor, 1936), (Bongers, 1988), (de

Grisse & Loof, 1965); except that ratio a is slightly lower, as is V in

one specimen. Differences between the tail termini are not consid-

ered in this instance to be of specific importance.

i e

_——

SSE

<a) Samana

————

ee a

D E

Fig. 26 Criconema demani (Micoletzky, 1915). A, B, D, female 1. A,

anterior region; B, posterior region; D, tail. C, E, female 2. C, posterior

region; E, tail. Scale bars a = 30 um, b = 10 pm.

Discocriconemella sp.

(Fig. 27)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 4, 12.

FEMONES. (= 1) Jb 257/ line A= O22 = SF © = MIs W = ee

R= 98 (ventral side); Rex = 38; RV = 13; Ran=8; VL/VB = 1.0; tail

= 14.1 um.

Labial annule discoid, offset from body. Body annules retrorse

except fifth to twelfth which are anteriorly directed on ventral side;

annules with anastomoses and finely crenate posterior edges; out-

line somewhat angular. Spear 54 um long, knobs 7.4 um wide with

anterior surfaces indented. Oesophagus about 82 um long. Vulva

apparently closed. Anterior genital branch poorly preserved.

Spermatheca filled with sperms. Tail terminus bluntly rounded.

REMARKS. This female closely agrees with the description of D.

limitanea Luc, 1959 by Rashid et al., (1987) but differs by having

fewer anteriorly directed annules (Sth to 12th in this female vs 4th to

23rd annules in their population). Possibly this is the first record of

this tropical genus in Europe.

Hemicycliophora sp.

MATERIAL EXAMINED. Foyers Plateau: 159 m; core 14, 1 8.

REMARKS. Specimen in very poor condition and not worth des-

cribing.

Ogma sp.

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 3, 1 9. River

Oich: 0.31 m; core 31, 19.

F.R. WANLESS AND R. HUNTER

Fig. 27. Discocriconemella sp. A-C, female. A, habitus; B, posterior

region; C, anterior region. Scale bars a = 30 um, b = 20 um.

REMARKS. Neither specimen is in good condition, the spear knobs

are missing; the oesophagus and genital structures are poorly pre-

served.

Paratylenchus sp.

(Fig. 28)

MATERIAL EXAMINED. Fort Augustus Bay: 92 m; core 4, 1 9.

FEMALE. (@= 1) 5 =318iim:eia = 23:72 b= 4679 seen

V =77; tail = 34.2 um.

Fig. 28 Paratylenchus sp. A-D, female. A, oesophageal region; B,

habitus; C, lateral field; D, tail. Scale bars a= 10 um, b = 50 um.

FRESHWATER NEMATODES FROM LOCH NESS

Head rounded with truncate anterior end, and minute protrusions

at margins of lip region Annules 1.0 um at mid-body. Lateral field

with four lines. Spear 14 um long with laterally directed knobs

3.2um wide. Dorsal oesophageal opening about 5.0 um behind

spear knobs. Excretory pore 69 um from anterior. Hemizonid just

anterior to excretory pore. Oesophagus 84 um long, poorly pre-

served. Vulva flaps prominent about 5.0 um long. Vagina curved

anteriorly. Anterior genital branch outstretched. Spermatheca not

seen. Vulva-anus distance 36.5 um. Tail tip finely rounded.

REMARKS. Using Raski’s (1975) key to, Paratylenchus spp with a

spear < 38 um long, this female comes to either P. italiensis Raski,

1975 or P. tateae Wu and Townshend, 1973. Unfortunately these

species are difficult to separate from one another because of overlap

or continuous sequences in respect of their measurements. Head

structures are also remarkably similar. The present female differs

slightly in respect of ratios c, V and spear length, also the head is not

offset. More specimens are required to assess variation and identify

this female with any confidence.

Aphelenchoides sp. |

(Fig. 29)

MATERIAL EXAMINED. River Oich: 0.31 m; core 31, 19. River

Foyers: 0.22 m; core 40, 1 9.

HORIZON. Core 40, (1—2 cm); Core 31, (2—3 cm).

FEMALES. (n= 2). L=387, 492 um; a= 35, 33.4; b= 6.7, 8.2;c=

eles 9ce— 4.5, 4.0; V ='67.7, 69; tail= 31:5, 35.4 um.

Head region smooth, slightly offset. Cuticle with fine striations

about 0.9 um wide. Lateral fields with four lateral lines, inner pair

faint. Spear 10.2, 10.8 um long with small basal thickenings.

Procorpus of oesophagus narrow; median bulb ovoid, valve well

a a

Fig. 29 Aphelenchoides sp. 1. A-E, female. A, anterior region; B, tail; C,

habitus; D, vulval region; E, lateral field. Scale bars a = 10 um, b = 50

um.

developed, located posterior to middle of bulb. Excretory pore less

than one body width posterior to median bulb. Vagina slightly

inclined anteriorly. Ovary outstretched anteriorly 116, 152 um long.

Spermatheca not seen. Postvulval uterine sac apparently absent.

Anus distinct, anterior lip slightly raised. Tail conical, terminating in

a pointed ventral mucro.

REMARKS. Using Sanwal (1960) these females key out as A.

goeldi Steiner, 1941 primarily because the postvulval uterine sac is

absent. However, these females clearly differ from A. goe/di in both

head and tail shape.

Aphelenchoides sp. 2

(Fig. 30)

River Oich: 0.31 m; core 31, 12.

Core 31, (1-2 cm).

BEVALLEN On — I) —A70limea—3351b 161): e—62160—19-5;

Vi= 72 xtail = 29) (im.

Head region smooth, offset by constriction. Cuticle with fine

annules about 0.9 um wide. Lateral fields with four lines, inners very

faint. Spear 10.2 um long with small basal thickenings. Procorpus of

oesophagus moderately wide; median bulb ovoid, valve well devel-

oped, located slightly posterior to middle of bulb. Excretory pore

opening slightly anterior to base of median bulb. Vagina slightly

tilted anteriorly. Anterior genital branch outstretched 204 um long;

spermatheca 177 um long with disc-like sperms. Postvulval uterine

sac 43.5 um long without sperm. Anus moderately distinct.

MATERIAL EXAMINED.

HORIZON.

REMARKS. Additional specimens are needed in order to identify

this nematode with confidence. Morphometric measurements and

the anterior position of the excretory pore suggests that it may be

Fig. 30 Aphelenchoides sp. 2. A-E, female. A, anterior region; B, tail; C,

vulval region; D, habitus; E, lateral field. Scale bars a = 10 um, b = 50

um.

close to A. cyrtus Paesler, 1957 originally described from mush-

room-beds of the Champignonzuchtinstitut Dieskau near Halle,

Germany.

Aphelenchoides sp. 3

(Fig. 31)

MATERIAL EXAMINED. River Foyers: 0.22 m; core 40, 1 8.

FISIWANOS, (Giles HN) Ib SWS ing AS S72 lo = NS ARCS N22 e = sa22

V = 68; tail = 50 um

Head region smooth and slightly offset. Cuticle with fine striations

about 0.9 um wide. Lateral fields with four lines, the inner pair

indistinct and fading away in some regions. Spear 10.8 um long with

pronounced basal thickenings. Procorpus of oesophagus hardly

visible; median bulb more or less ovoid with well developed valve

located slightly posterior to middle of bulb. Excretory pore about 1.5

body widths posterior to median bulb. Vagina directed anteriorly.

Anterior genital branch outstretched, about 290 um long.

Spermatheca 58 um long with globular sperms. Postvulval uterine

sac 123 um also with globular sperms. Anus distinct. Tail concoid

with terminal mucro.

REMARKS. This female is very similar to A. fragariae (Ritzema

Bos, 1890), but differs by having more robust basal spear thickenings;

described as minute but distinct in A. fragariae, Siddiqi (1975).

Fig. 31 Aphelenchoides sp. 3. A-E, female. A, anterior region; B, vulval

region; C, lateral field; D, tail; E, habitus. Scale bars a = 10 um, b = 100

um.

F.R. WANLESS AND R. HUNTER

cf. Caenorhabditis sp.

(Fig. 32)

MATERIAL EXAMINED. River Oich: 0.31 m; core 31, 12.

HORIZON. Core 31 (1-2 cm).

FEMALES (i — 1) S62 5i ima — 30s b= 519 %c— oO nce— clo

= 50; tail = 105 um.

Head not offset, 7.0 um wide with minute labial papillae. Amphids

not seen. Cuticle finely annulated; annules about 0.9um wide at

mid-body. Stoma 14.1 um long; cheilostome distinct, lightly

sclerotised, barely setoff from long prorhabdions; metastome with

two small denticles. Oesophageal collar not observed. Oesophagus

159 um long with swollen midbulb. Excretory pore not seen.

Ovaries paired, anterior branch reflexed. Spermatheca filled with

more or less rounded sperms about 2.2 um.in diameter. Tail straight,

tapering to fine tip. Phasmids inconspicuous.

REMARKS. Additional specimens, especially males, are required

in order to properly identify this nematode.

Diplogasteritus nudicapitatus (Steiner, 1914)

(Figs 33, 36H)

MATERIAL EXAMINED.

core 4,10.

Borlum Bay (1992 collection): 0.38 m;

Fig. 32 cf. Caenorhabditis sp. A-D, female. A, oesophageal region; B,

vulval region; C, tail; D, habitus. Scale bars a = 20 um, b = 50 um.

FRESHWATER NEMATODES FROM LOCH NESS

Fig. 33 Diplogasteritus nudicapitatus (Steiner, 1914). A-C, male. A,

habitus; B, spicular region; C, oesophageal region. Scale bars a = 20

um, b = 50 um.

MADEN I). lo = 637 im; a= 25; b = 6.3; c = 4.0; c = 8.5;

gubernaculum = 25.6 um; spicule = 33 um; tail = 155 um.

Head not offset; labial papillae minute. Amphids not seen. Cuticle

with ca. 20 prominent longitudinal lines. Annules minute, less than

}

Fig. 34 Eucephalobus oxyuroides (de Man, 1876). A, B, E, F, female 1.

A, tail; B, anterior region; E, vulval region; F, habitus. C, D, female 2.

C, habitus; D, tail. Scale bars a = 20 um, b = 100 um.

0.8um wide. Oesophagus with large median bulb and smaller basal

bulb; 105 um long. Excretory pore 98um from anterior, at mid-level

of basal bulb. Testis anteriorly directed with reflexed tip. Spicula 1.8

times anal body width. Gubernaculum, distally with two pairs of

mound-like swellings (arrowed, fig. 36H). Bursa absent. Nine pairs

of caudal papillae; three pairs pre-anal and six post-anal.

DISTRIBUTION AND HABITAT. A common species found in Europe

and South Africa in terrestrial and aquatic habitats.

REMARKS. Morphology and measurements closely agree with the

descriptions of D. nudicapitatus provided by Bongers (1988) and

Zullini (1982).

Eucephalobus oxyuroides (de Man, 1876) Steiner, 1936.

(Figs 34, 36E-G)

MATERIAL EXAMINED. Foyers Plateau: 161 m; core 19, 1 9. River

Oich: 0.17 m; core 36, | .

FEMALES. (n=2).L=558, 615 um; a= 23.5, 19.6; b=3.6, 3.9;c

—G.55 ieee 0.0, 4.05 V.—= 55.5, Ol: tailis55 79-3 um.

Cuticle 1.9 um thick with annules 1.4, 2.1 um wide at mid-body.

Lips six, pointed. Lateral fields with three lines ending at level of

phasmids. Stoma (n= 1) 14.7um long, rhabdions distinct. Oesopha-

gus cylindrical, 153, 157 um long, basal bulb ovate. Excretory pore

(n=1) 91 um from anterior end at level of hemizonid. Reproductive

system monodelphic, prodelphic. Reflexed at oviduct. Spermatheca

and postuterine sac not seen. Vagina length about one half of vulval

body width. Vulva not protruding. Rectum 23, 26 um long or 1.8, 1.6

times anal body width. Tail 80-86um long. Phasmid at 28, 32% of

tail length.

Fig. 35 Eucephalobus sp. A-D, female. A,oesophageal region; B,

reproductive system; C, tail; D, habitus. Scale bars a = 20 um, b = 50

uum.

DISTRIBUTION AND HABITAT. Cosmopolitan, prefers dry ground,

but also mentioned from freshwater.

REMARKS. These specimens agree with descriptions of this species

by Thorne (1937) and Rashid et al., (1984). Note variation in length

of the slender tail terminus and appearance of the lateral field at

different focal levels.

Eucephalobus sp.

(Fig. 35)

MATERIAL EXAMINED. River Oich: 0.31 cm; core 31, 1 2.

FEMALE. (n=1).L=430 pm; a= 19.8; b=3.5; c= 10.2; c¢ =3.4;

V = 63. tail 42 um.

Cuticle 1.2 um thick. Annules 1.6 um wide at mid-body. Lips six,

pointed. Lateral field with three lines in region of oesophageal basal

bulb, not evident elsewhere. Stoma 12.8 um long, rhabdions indis-

tinct. Oesophagus cylindrical, 122 um long. Excretory pore 73.6 um

from anterior end, hemizonid not seen. Reproduction system

monodelphic, prodelphic. Ovary with double flexure. Egg 38 um

long by 16 um wide. Spermatheca not seen. Postuterine sac 9.6 um

long. Vulva protruding. Length of vagina about one quarter of vulval

body width. Rectum 10.2 um long, or 0.8 times anal body width. Tail

terminus without distinct annules, broadly spear-shaped. Phasmid

not evident.

REMARKS. The distinctive spear-like tail terminus of this female is

not known to occur in other species of Eucephalobus and this

specimen may represent a new taxon. It cannot however, be described

until additional material becomes available for study.

ACKNOWLEDGEMENTS. This work was made possible by the financial

support of The Natural History Museum, London and by sponsorship of the

all-documentory television station The Discovery Channel under the aus-

pices of Project Urquhart. Additional support was provided by Simrad, the

electronics company which was the main sponsor of the hydrological survey

of the loch. Special thanks are due to Mr Nicholas Witchell, the founder and

Chairman of Project Urquhart. We also thank the crew of the research vessel

Soel Mara who’s skills contributed greatly to the success of the profundal

coring programme. The late Professor M. W. Brzeski, Museum and Institute

of Zoology, Polish Academy of Sciences, Warszawa, Poland kindly read

through early descriptions of these nematodes and his comments are greatly

appreciated. Several of our colleagues also deserve many thanks; Nicola

Mitchell gave invaluable help assisting with field and laboratory work; Dr T,

J Ferrero produced the map of Loch Ness and ‘sorted’ many of the word

processing problems encounted by the first author. Translation of various

German and Dutch texts by Dr R. Huys is also gratefully acknowledged.

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Dy a" Ex6 S| ®.

Taxonomy of Oncaeidae (Copepoda,

Poecilostomatoida) from the Red Sea.

II. Seven species of Oncaea s.str.

RUTH BOTTGER-SCHNACK

Institut fiir Meereskunde an der Universitat Kiel, Diisternbrooker Weg 20, D-24105 Kiel, Germany. E-mail

address: dschnack @ ifm.uni-kiel.de

SYNOPSIS. Seven species of Oncaeidae (one of which is new to science) are described from the Red Sea, including a description

of the hitherto unknown male of Oncaea clevei Friichtl. They all belong to Oncaea sensu stricto as defined in a phylogenetic study

of oncaeid species by Bottger-Schnack and Huys in 1998. The genus is defined by a combination of characters including the

absence of a conical process on the distal margin of the endopod of swimming leg 4, ornamentation of the labrum and mandibular

blade and sexual dimorphism in antennary setae and in endopods of swimming legs. Oncaea s. str. includes two subgroups, the

venusta-subgroup and the c/evei-subgroup, females of which can be separated by the absence or presence of a dorso-posterior

projection on the P2-bearing somite. Species of the venusta-subgroup in the Red Sea include O. venusta Philippi, O. mediterranea

(Claus), O. media Giesbrecht, O. scottodicarloi Heron & Bradford-Grieve and O. waldemari Bersano & Boxshall; the clevei-

subgroup consists of O. clevei Friichtl and O. paraclevei sp. nov. Oncaea praeclara Humes is a junior subjective synonym of O.

venusta. O. philippinensis (Kazatchenko & Avdeev) as well as Oncaea sp. | and sp. 2 sensu Ferrari are regarded as species

inquirendae in Oncaea s.str. The type species of the genus, O. venusta, has two form variants, forma fypica and forma venella

Farran, which can be distinguished by differences in size and in a few minor morphological characters. Their spatio-temporal

distribution differs considerably in the Red Sea, which may indicate reproductive isolation. The world-wide records of O. venusta

forms are reviewed and discussed.

Issued 28 June 2001

CONTENTS

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Description of species .

Oncaea venusta Philippi forma typica Farran

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INTRODUCTION

Small copepod species of the family Oncaeidae are widely distrib-

uted in marine pelagic areas, extending from the epipelagic zone

(e.g. Boxshall, 1977a; Dessier & Donguy, 1985; Cowles ef ai.,

1987) to the deep-sea (Wishner, 1979; Humes, 1988) and from

tropical (Tsalkina, 1970; Deevey & Brooks, 1977) to polar regions

(Hopkins, 1985; Richter, 1994; Metz, 1995). Due to their small size,

oncaeid copepods are often neglected or under represented in marine

plankton investigations, which are usually conducted with conven-

tional nets of 0.3 or 0.5 mm mesh sizes. In recent years, increasing

attention has been paid to the study of smaller mesozooplankton

organisms sampled with fine mesh nets of 0.1 mm mesh size or less.

In particular, small oncaeid copepods were sampled in high abun-

dance by these nets (e.g. B6ttger, 1982; Paffenh6fer, 1983; Schnack

et al., 1985; Cowles et al., 1987; Bottger-Schnack, 1987) and

subsequent research topics have focussed on their community struc-

ture and biology [see Bottger-Schnack (1999) for a review of the

literature], thereby providing new insights into the largely unknown

ecological importance of this copepod group.

Apart from ecological issues, the taxonomy of oncaeid species

has been the subject of increasingly detailed studies over the past

two decades (e.g. Boxshall, 1977b; Heron, 1977; Malt, 1982a, b,

1983a, b, c, d; Heron ef al., 1984; Krsini¢é & Malt, 1985: Boxshall &

Bottger, 1987; KrSini¢, 1988; Bottger-Schnack & Boxshall, 1990;

Heron & Bradford-Grieve, 1995; Bottger-Schnack & Huys, 1997a,

b, in press; Itoh [in: Chihara & Murano (1997)]). Despite these

efforts, however, the taxonomy of many oncaeid species is still

inadequate and particularly those species that are smaller than 0.5

mm have not been described in sufficient detail. Moreover, many

oncaeid species are morphologically very similar, and thus can be

distinguished only by a few characters, including microstructure of

the mouthparts and on the exoskeleton, which require a level of

detail not generally adopted in most taxonomic descriptions. Such

integumental microstructures have been used to differentiate bet-

ween closely related species of cyclopoid (Ueda et al., 1996; Rocha,

1998) and oncaeid (Bottger-Schnack, 1999; Bottger-Schnack &

Huys, in press) copepods.

The phylogenetic relationships of the Oncaeidae are not well

understood. The diphyletic status of the family has recently been

recognized by Huys & Bottger-Schnack (1996/97), who proposed

the new family Lubbockiidae to accommodate Lubbockia Claus,

1862 and related genera, retaining only Oncaea Philippi, 1843,

Conaea Giesbrecht, 1891 and Epicalymma Heron, 1977 in the

Oncaeidae. The large type genus Oncaea sensu lato, which cur-

rently contains over 70 validly described species (Heron &

Bradford-Grieve, 1995), has been recognized as a paraphyletic

assemblage (Huys & Bottger-Schnack, 1996/97). A detailed phyloge-

netic analysis of the three oncaeid genera at the species level is

currently in progress and will result in the recognition of numerous

new genera. Some preliminary results of the phylogenetic study

were briefly presented by B6ttger-Schnack & Huys (1998).

The Red Sea is an extreme environment, with constant and

unusually high deep-sea temperatures (21.7°C) and salinities (40.5

PSU) (Edwards, 1987). Due to these unique environmental conditi-

ons an impoverished zooplankton fauna is found in the Red Sea,

which is characterized by the absence of true deep sea species

(Weikert, 1982, 1987). This phenomenon was also reported for

oncaeid copepods (Bottger-Schnack, 1994, 1999) and their esti-

mated species number in the Red Sea (about 26) appears to be low in

comparison to the adjacent Arabian Sea, where the vast majority of

the about 70 recorded oncaeid species and forms is made up by deep-

living representatives (Bottger-Schnack, 1994). The surface waters

in the Red Sea show a considerable increase in salinity from the

southern entrance to the central-northern areas (Morcos, 1970). Due

to the less favourable environmental conditions in the north, species

numbers of planktonic taxa generally decrease from south to north

(e.g. Halim, 1969; Kimor, 1973; Almeida Prado-Por, 1983; Bottger-

Schnack, 1995). For oncaeid copepods, however, a corresponding

regional decrease in species number from south to north is less

pronounced (Bottger-Schnack, 1995). This unexpected result was

assumed to be at least partly attributable to the taxonomic difficul-

ties encountered in the identification of oncaeids, since the proportion

of unidentified form variants recorded in the south was much higher

than in the north. In a subsequent taxonomic study five new species

of oncaeids were described, four of which occurred mainly or

exclusively in the southern area (Bottger-Schnack, 1999), thereby

indicating some latitudinal difference in species numbers for this

copepod group.

The ecology of oncaeid copepods in the Red Sea has been

investigated intensively during the past years within the framework

of multidisciplinary environmental research programmes (Thiel et

al., 1986; Weikert, 1988). The species diversity, vertical distribution

and diurnal vertical migration, as well as some biological parame-

ters, such as variation in body length and breeding activity or

feeding of oncaeid copepods were studied on the basis of routine

sampling with fine nets of 0.1 mm or 0.05 mm mesh size in the upper

500 m or 1000 m of the water column during different seasons and

in various regions [see Bottger-Schnack (1999) for a review of the

literature]. The taxonomy of Red Sea Oncaeidae is less well known,

however. Boxshall & B6ttger (1987) and Bottger-Schnack &

Boxshall (1990) described four new oncaeid species from the central

Red Sea, and provided a redescription of Oncaea atlantica Shmeleva.

More recently, O. mediterranea (Claus) was redescribed from this

R. BOTTGER-SCHNACK

area, including a thorough review of the taxonomic history of this

allegedly cosmopolitan species (B6ttger-Schnack & Huys, 1997). In

arecent taxonomic study, 11 species of Oncaeidae were described or

redescribed from the Red Sea, which belong to the conifera/similis-

group as defined in the phylogenetic study of B6ttger-Schnack &

Huys (1998). A new genus, Triconia, was proposed to accommodate

all oncaeid species belonging to this group (Bottger-Schnack, 1999).

As part of an ongoing taxonomic study of Red Sea Oncaeidae, the

present paper describes seven species, which belong to the venusta-

group as defined by Bottger-Schnack & Huys (1998). In their

cladogram of oncaeid species it is equivalent to species group 2.

This core-group includes Oncaea venusta Philippi, the type species

of the genus, and is considered here as Oncaea sensu stricto. Oncaea

s.str. 1s defined by a combination of characters including the absence

of a conical projection on the distal margin of P4 endopod, ornamen-

tation of the labrum and of the dorsal blade of the mandible, sexual

dimorphism in the antenna and in the endopods of the swimming

legs. Oncaea s. str. is considered as the sister-group of Triconia

Bottger-Schnack, 1999, and the establishment of the revised genus

is briefly substantiated in the present paper. A detailed discussion of

phylogenetic relationships of Oncaea s.str. within the Oncaeidae

will be published separately (Huys & Bottger-Schnack, in prep.).

Oncaea s.str. currently includes 8 species, 6 of which had previ-

ously been reported from the Red Sea (Bottger-Schnack, 1994) and

are redescribed in the present account. For O. mediterranea,

redescribed recently by Bottger-Schnack & Huys (1997b), some

short corrective notes will be given. A new species similar to O.

clevei Friichtl will be described. The taxonomic status of O. curta

Sars, for which no material was available, will be discussed on the

basis of literature data.

The type species of Oncaea s.str., O. venusta, was recently

redescribed in detail by Heron & Bradford-Grieve (1995), based on

specimens collected in the Gulf of Naples, near the type locality in

the Mediterranean Sea. Their account does not contain information

about the two size variants of the species, forma typica and forma

venella (Farran, 1929), which are generally known to occur in

tropical and temperate areas (Malt, 1983b), but are poorly docu-

mented morphologically. Apart from a distinct size difference, only

few minor differences in body morphology have been reported for

the two forms so far (Farran, 1929; Sewell, 1947: Ferrari, 1975:

Boxshall, 1977a). In the Red Sea, both size variants of O. venusta

occur. They differ considerably in spatio-temporal distribution

(Bottger-Schnack, 1990b, 1995), which may indicate reproductive

isolation. Both forms of O. venusta will be redescribed in detail in

the present account, including observations with scanning electron

microscope (SEM). Particular attention is paid to microstructures in

the mouthparts and on the exoskeleton, which might provide new

and helpful information for the separation of the two forms. The

relationship of the two Red Sea forms with the species occurring in

the Mediterranean Sea is defined upon re-examination of specimens

from Heron & Bradford-Grieve’s material. The present knowledge

on the world-wide distribution of the two venusta forms is reviewed.

The two size variants of O. media as defined by Sewell (1947)

have recently been recognized as distinct species by Heron &

Bradford-Grieve (1995). They assigned forma major to O. media

Giesbrecht (except for his Plate 47, Fig. 11), and described forma

minor as anew species, O. scottodicarloi. Both species occur in the

Red Sea and are briefly redescribed in the present paper, including

some important morphological details not noted by these authors. O.

waldemari, which is very similar to O. media and O. scottodicarloi,

had recently been described from Brazilian waters by Bersano &

Boxshall (1994). Due to some descriptive errors, however, the

authors did not notice the close relationship of O. waldemari with

SEVEN SPECIES OF RED SEA ONCAEA

species of Oncaea s.str. and erroneously placed it in a group

containing O. petila Heron, O. ovalis Shmeleva and others. In the

present account, O. waldemari is completely redescribed based on

Red Sea specimens (figures) and the type material from Brazilian

waters, with additional comparisons on specimens from different

localities in the Atlantic and Pacific Ocean.

MATERIAL AND METHODS

Oncaeids were collected using a multiple opening-closing net with

a mesh size of 0.05 mm during cruise 5/5 of R/V Meteor (Weikert,

1988) in various locations of the Red Sea (Fig. 1). Samples collected

during cruise 29 of R/V Valdivia with the same sampling gear, but

equipped with nets of 0.1 mm mesh, were also examined. A station

list and sampling data are given in Table I. The plankton was initially

fixed in a 4% formaldehyde-seawater solution buffered with

hexamethylene tetramine and transferred after ca 2 years into a

preservation fluid of 5% propylene glycol, 0.5% propylene pheno-

xetol, and 94.5% filtered seawater (Steedman, 1976). Specimens

were dissected in lactic acid, mounted on slides in lactophenol and

sealed with transparent nail-varnish. All figures have been prepared

using a camera lucida on a Leitz Dialux differential interference

contrast microscope.

Total body length and the ratio of prosome to urosome (excluding

caudal rami) were calculated as the sum of the middorsal lengths of

individual somites measured in lateral view. In the case of telescop-

ing somites these lengths are measured from the anterior to the

Table 1 Sampling data of multiple opening-closing net in the Red Sea and adjacent areas (D = Day, N = Night).

Stn. No. Date Time

Red Sea + Gulf of Aden

R/V Valdivia Cruise 29

130 28.10.1980 D

664 21. 2.1981 D

R/V Meteor Cruise 5/5

63la 11.7.1987 N

663 20.7.1987 D

703 3.8.1987 D

708 5.8.1987 D

Northern Arabian Sea

R/V Meteor Cruise 32/3

247 14. 5.1995 D

Eastern Mediterranean Sea

R/V Meteor Cruise 5/1

35 20. 1.1987 N

M@Port Said

cae, }

4 Jiddan

Port Sudan ‘gl,

30° OS

Total water depth (m)

Mesh size (mm) Geographic position (°N, °E)

1960 0.1 PLe2S Si A308 201.9!

2000 0.1 INP Plas StF 0S

1400 0.05 Iter ye wars lo hase /)

1200 0.05 22°58.4', 37°19.4'

970 0.05 15°34.8', 41°54.9'

190 0.05 13°40.0', 42°37.4'

3000 0.05 ca 19°, ca 65°

3400 0.05 34°25.3', 26°14.8'

AS: Karachi

27 | i We

oe (eet

60° 70°

Fig. 1 Location of stations in the Red Sea and adjacent areas. A = Valdivia-Cruise 29, October 1980, February 1981; Hl = Meteor-Cruise 5/1, January

1987; @ = Meteor-Cruise 5/3a, April 1987; W = Meteor-Cruise 5/5, July/August 1987; ® = Meteor-Cruise 32/3, May 1995.

posterior margin. This approach differs from that traditionally used

in oncaeid taxonomy, where the telescoping of somites is not

considered in length measurements. Traditional length data of Red

Sea oncaeids as given by Bottger-Schnack et al. (1989) are only up

to 70% of the sizes presented in this paper, due to the excessive

telescoping of somites in the sorting medium. In order to make sizes

of the species in this paper comparable to those of previous taxo-

nomic descriptions (e.g. Heron & Bradford-Grieve, 1995; Bersano

& Boxshall, 1994), length data of each species were obtained by the

traditional method as well (i.e. measured dorsally from the tip of

prosome to the distal end of caudal ramus) and are given in square

brackets.

Descriptive terminology for body and appendages follows that of

Huys and Boxshall (1991). Abbreviations used in the text are: ae =

aesthetasc; CR = caudal rami; enp = endopod; exp = exopod;

exp(enp)-1(-2, -3) = to denote the proximal (middle, distal) segment

of aramus; P1—P4 = swimming legs 1-4. Species of Oncaeidae have

a number of pores and other integumental structures (e.g. pits,

scales) on body surfaces, but only those discernible with a light

microscope were figured or mentioned.

O. venusta was examined with a Philips XL30 scanning electron

miscroscope. Specimens were prepared by dehydration through

graded ethanol, critical point dried, mounted on stubs and sputter-

coated with palladium.

Type and other material is deposited in the collections of The

Natural History Museum, London (BMNH), Smithsonian Institu-

tion (USNM) and the Zoologisches Institut und Museum der

Universitat Hamburg (ZMH). Paratypes or other material retained in

personal collection, R. Bottger-Schnack designated (RBS).

GENERIC DIAGNOSIS

Order POECILOSTOMATOIDA Thorell, 1859

Family ONCAEIDAE Giesbrecht, 1892

Oncaea sensu stricto

DIAGNOSIS. Body cyclopiform, prosome stout or broad-oval. P2-

bearing somite with (clevei-subgroup) or without (venusta-subgroup)

dorsoposterior projection in female. Cephalosome without lateral

lobate extensions. Genital double-somite female slightly flask-

shaped, not particularly swollen dorsally. First and second postgenital

somites shorter than anal one. Anal somite with wide anal opening;

operculum with small spinules.

Exoskeleton well or heavily chitinized, usually ornamented with

numerous pores, ridges and scales.

Sexual dimorphism in antennule segmentation and armature,

antennary setae, maxilliped, genital segmentation and ornamenta-

tion, endopods of P1—P3 (sometimes also in P4), P5 and P6, and in

caudal ramus.

Antennule 6-segmented in female with armature formula 1-[3], 2-

[8], 3- [5], 4-[3 + ae], 5-[2 + ae], 6-[6 (1 +ae)]; 4-segmented in male

with formula 1-[3], 2-[8], 3-[4], 4-[11 + (1 + ae) + 2ae]. Distalmost

seta of female segment 3 absent in male. Aesthetascs slender.

Antenna. Enp-1 with triangular projection on outer margin form-

ing concavity distally; inner margin with 1—2 denticular rows. Enp-2

shorter than enp-1; posterior surface with double row of spinules;

lateral armature consisting of 3 well developed, bare or minutely

pinnate setae (I, I] and IV) and 1 pinnate spine (III) in female; distal

armature consisting of 4 pinnate setae, | long bare or minutely

pinnate seta, and 2 bare setae; posterior seta well developed. Sexual

R. BOTTGER-SCHNACK

dimorphism in lateral armature of enp-2, with seta III much stouter

and seta IV spiniform and curved in male, both elements shorter than

in female; often also expressed in coxobasal seta, being short and

naked in male.

Labrum distinctly bilobate, forming paired semi-circular

posteroventral lobes. Lobes without marginal teeth but with row of

minute denticles around outer ventral margin, with row of long fine

setules latero-distally (except for clevei-subgroup) and dentiform

processes converging and decreasing in size medially. Lobes sepa-

rated by semicircular vertex covered anteriorly by overlapping rows

of small hyaline petaloid flaps, flanked by slit-like pores located on

proximal part of each lobe; posterior face with median sclerotized

teeth and usually with paired patches of long fine setules (absent in

media). Anterior face with paired spinular patches or denticulate

rows (except for venusta); paired integumental pockets usually well

developed, free margin of pockets serrated or ornamented with

denticle row.

Mandible with 2 blades, 2 setae and | spine. Dorsal blade with 3

dentiform processes along distal margin and up to 2 additional

processes along dorsal margin.

Maxillule weakly bilobate; praecoxal arthrite with innermost

element proximally displaced and outermost element spiniform and

bearing transverse row of spinules.

Maxilla. Allobasis shorter than syncoxa.

Maxilliped (female). Large, moderately ovoid. Basis without

ornamentation on posterior surface; anterior surface with spinular

row and spinular fringe along palmar margin half the distance

between distal seta and endopod; both palmar elements long,

spiniform and spinulose, similar in length. Enp-1 completely sepa-

rated. Enp-2 with long, minutely pectinate claw, rudimentary outer

setule and fused unipinnate inner spine.

Maxilliped (male). Palmar margin forming shallow longitudinal

cleft bordered by anterior denticulate fringe and posterior multiple

rows of coarse, blunt spinules; not developed into distal flap.

Anterior surface of basis with patch of transverse spinular rows.

Palmar setae short, smooth. Endopodal claw curved, naked, with

hyaline apex.

PI exopod. Outer and terminal spines with subapical tubular

extensions.

P1 endopod. Enp-3 distal spine stout, with broad serrate hyaline

flange; base of distal inner seta concealed beneath long anterior

spinous outgrowth of segment; length of outgrowth (often) sexually

dimorphic, relatively longer in the male.

P2—P4 exopods. Outer spines stout, with broad, serrate hyaline

flanges; lateral spines not markedly increasing in size distally.

P2—P4 endopods 3-segmented. Enp-3 twice as long as enp-1 and

-2 combined or longer; with large conical processes on P2—P3, not

on P4. Enp-3 distal spine short in P2—P3, longer in P4; distal outer

spine small in P2—P3, always longer than conical process in female.

Sexual dimorphism usually expressed in length of conical proc-

esses, being longer in the male, and in spine length of enp-3, being

reduced in the male.

Swimming leg armature formula:

Leg Coxa Basis Exopod Endopod

Pl 0- 1-I I-0;I-1;01,14 0-1;0-1;1,5

P2 0-0 1-0 J-0;I-1;0,15 0-1;0-2;LIL3

P3 0-0 1-0 I-0;I-1;11,1,5 0-1;0—2;1, 11,2

P4 0-0 1-0 I-0;1-1;IL,1,5 0-1;0-2:1,11,1

P5 small; represented by outer basal seta and small exopod with

2 spiniform setae; exopod delimited at base in females, fused to

SEVEN SPECIES OF RED SEA ONCAEA

somite in males; sometimes additional sexual dimorphism in shape

and length of exopodal setae. Genital apertures of female large;

located near midregion of dorsal surface of genital double-somite

but usually in proximal half; each operculum with small spine and 2

minute spinules or spinous processes.

Male P6 membranous flaps produced posterolaterally into spinous

process; without armature.

Caudal ramus about 3-4 times as long as wide or shorter, without

conspicuous dorsal expansion surrounding base of seta VII. Seta I

absent; setae II and III unipinnate; setae IV and V not resilient,

relatively rigid and bipinnate; seta VII plumose and distinctly shorter

than V; seta VI spiniform and sparsely pinnate. None of setae

displaced. Sexual dimorphism expressed in length to width ratio of

CR, being smaller in male, and in proportional lengths of caudal setae.

TYPE SPECIES. Oncaea venusta Philippi, 1843 (by monotypy)

OTHER SPECIES

O. mediterranea (Claus, 1863)

O. media Giesbrecht, 1891

O. curta Sars, 1916 [not O. curta sensu Boxshall, 1977b]

*O. clevei Frichtl, 1923

[ O. praeclara Humes, 1988] syn. of O. venusta

O. waldemari Bersano & Boxshall, 1994

O. scottodicarloi Heron & Bradford-Grieve, 1995

*O. paraclevei sp. nov.

SPECIES INQUIRENDAE

Oncaea sp. | Ferrari, 1975

Oncaea sp. 2 Ferrari, 1975

O. philippinensis (Kazatchenko & Avdeev, 1977)

[Species marked with an asterisk (*) belong to the clevei-subgroup,

remaining species belong to venusta-subgroup. |

REMARKS

Within the Oncaeidae Oncaea s.str. belongs to a lineage comprising

the conifera/similis-group (Triconia Bottger-Schnack, 1999), the

notopus-group and the brocha-group (including O. brocha Heron

and O. olsoni Heron). These four groups correspond with species

groups 2, 3+4, 6 and 7 recognized by Bottger-Schnack & Huys

(1998) and together are considered to form a monophyletic lineage

on account of the structure of the labrum (median concavity with 4

posterior dentiform processes) and the male maxilliped (palmar

margin with multiple overlapping rows of blunt spinules). Oncaea

s.str. is considered here as the sistergroup of Triconia on the basis of

the presence of integumental pockets on the anterior surface of the

labrum and the formation of the dorsoposterior projection on the

second pedigerous somite. The fact that the latter character is not

expressed in all members of both genera is interpreted as the result

of secondary loss which happended convergently in each genus.

Oncaea s.str. differs from Triconia in the sexual dimorphism of the

antenna (seta II and IV modified in the male), the presence of paired

slit-like pores on the anterior surface of the labrum and the absence

of a conical process on the distal endopod segment of P4. A more

detailed discussion of the phylogenetic relationships of Oncaea

s.str. within the Oncaeidae is beyond the scope of this paper and will

be published separately (Huys & B6ttger-Schnack, in prep.).

The revised genus includes two subgroups of species, the venusta-

subgroup and the c/evei-subgroup. Females of the clevei-subgroup

are characterized by a dorso-posterior projection on the P2-bearing

somite, which is lacking in the venusta-subgroup. The dorsal projec-

tion on the prosome (‘hump’) is a sexually dimorphic character,

which is absent in males. Thus males of both subgroups are very

hard to distinguish. A further morphological character separating

the 2 subgroups is found in the ornamentation of the labrum, the row

of long setules on the latero-distal margin of the lobes being absent

in the clevei-subgroup.

Oncaea curta Sars, 1916 (p. 228, Plate IV, as Once@a curta) was

originally described from the western Mediterranean, near the

Moroccoan coast. The species has also been widely recorded at low

latitudes in the Atlantic and Pacific [as compiled by Malt (1983a)

and Razouls (1996)]. Recently, Heron & Bradford-Grieve (1995,

p.41) recorded the species from the Gulf of Naples, but at the same

time noted in their samples several as yet undescribed species close

to O. media, O. scottodicarloi and O. curta indicating that a complex

of species close to O. curta exists within Oncaea s.str. Thus, a

number of different species might have been recorded in the litera-

ture under the wrong name curta (e.g. Boxshall 1977b, see remarks

below) and the geographical records of O. curta appear to be doubt-

ful. Due to the identification problems, the species had erroneously

been placed into the bowmani-group in the preliminary cladogram

of Bottger-Schnack & Huys (1998). The species could not be re-

examined during the present study, since no material was available.

Therefore, Sars’ description was taken as a basis to compare the

morphology of O. curta with the closely related O. media, O.

scottodicarloi and O. waldemari (see under O. scottodicarloi, Re-

marks). A thorough revision of O. curta is needed to clarify the

taxonomic confusion surrounding this species.

Oncaea curta sensu Boxshall (1977b: p.141—143, Table 1-2, Fig.

21a—k) does not belong to Oncaea s. str. Malt (1983a) has already

pointed out the similarity of the species with O. i/lgi Heron, which

belongs to the bowmani-group as defined in the phylogenetic study

of Bottger-Schnack & Huys (1998; their species group 5). Common

characteristics of this group are very elongate conical processes on

the distal endopod margins in P2—P3 and a very robust maxillipedal

basis, armed with 2 relatively short setae. Boxshall (1977b)

synonymized O. curta with O. ovalis Shmeleva, O. longiseta

Shmeleva and O. latimana Gordeyeva, as well as with Oncaea sp. |

and 2 of Ferrari (1975), but his opinion was not followed by Malt

(1983a) nor in the present account.

Oncaea sp. | and sp. 2 described by Ferrari (1975; Oncaea sp. 1:

p. 228, Figs. 6E, F, 7A—D; Oncaea sp.2: p.228, Figs. 6G, H, 7E—H)

are placed in Oncaea s.str. on the basis of the swimming leg

armature, the lack of a conical process on the distal margin of P4

endopod, and the size and position of the maxillipedal setae. Malt

(1983a) tentatively assigned Oncaea sp. 1 to O. media Giesbrecht

forma minor and Oncaea sp. 2 to both O. curta Sars and O. venusta

Giesbrecht f. venella, reflecting the author’s undecisiveness on this

matter. Since the original description of both species lacks sufficient

detail necessary for unequivocal identification, they are regarded

here as species inquirendae in Oncaea s.stt.

Myspictosum philippinensis Kazatchenko & Avdeey, 1977 (p.

44-47, Fig. 9a-1, 10a—g) has been synonymized with Oncaea s.1. by

Malt (1982a; erroneously spelled Myctospictosum by Malt) and her

opinion was followed by Huys & Bottger-Schnack (1996/97). The

single male of O. philippinensis was found on the gills of the deep-

sea fish Myctophum spinosum collected in the Philippine Trench at

7255 m, which is the deepest record of an oncaeid species so far. The

unusually deep record has been regarded as accidental rather than

real (Huys & B6ttger-Schnack, 1996/97). O. philippinensis might

well belong to Oncaea s.str., based on the modified seta IV on the

antenna, which is hook-like as in most other species of the genus.

However, the leg armature in Kazatchenko & Avdeev’s description

is incomplete in P2 (endopod missing) and very unusual in P1, the

endopod showing 4 inner setae and 3 outer and distal spines, with a

conical process at the distal margin. This combination of armature

elements is not found in any oncaeid species known thus far and the

species is regarded here as species inquirenda in Oncaea s.str.

Oncaea praeclara Humes is regarded as a synonym of O. venusta,

on grounds that will be discussed below under O. venusta, Remarks.

DESCRIPTION OF SPECIES

Oncaea venusta Philippi, 1843

Oncaea venusta Philippi (1843): 62-63, Tafel II, Fig. 2a—d.

Oncda venusta Giesbrecht, 1892

Oncea venusta Farran, 1929

Oncaea praeclara Humes, 1988

RELIABLE DESCRIPTIONS. Giesbrecht (1892): 590-604, 755, 756,

774, 789, Plate 2, Fig: 5; Plate 3, Fig: 7, Plate} 47, Fig.

2,5,13,19,39,44,48,50,54,58 [as Oncda venusta]; Farran (1929):

284-285, Fig. 33 [as Oncea venusta]; Wilson (1932): 353-354, Fig.

213a—d; Mori (1937, reprinted 1964): 119-120, Plate 66 Figs. 1-9;

Dakin & Colefax (1940): 116, Fig. 205A a-f; Sewell (1947): 263-

264; Olson [MS] (1949): 101-104, Plate XXIV Figs.1—8 (9), Plate

XXV Figs. 11-13 (d); Tanaka (1960): 71-72; Corral Estrada (1970):

216-217; Chen ert al. (1974): 40-41, Plate 6 Figs. 1-5; Ferrari

(1975): 225-228, Figs. 5I-K, 6A—D; Boxshall (1977b): 124-128

Figs. 1la—k (9), 12a—-d (d); Ho (1984): 41-44, Figs. 12-14; Humes

(1988): 475-485, Figs. la,b,d [not Fig. lc, e], 2a, 3a—g (9), 4a-g

(3) [as Oncaea praeclara]; Huys & Boxshall (1991): 286,289,445

Figs. 2.10.8.B, 2.10.10.A, 2.10.21.B, 2.10.21.D, 2.10.24.A; Heron

& Bradford-Grieve (1995): 33,36, Figs. 14e—-l, 15a—j, 27b, 28; Itoh

[in: Chihara & Murano 1997]: 981—982, Plate 223 Figs. 371 (left) a—

b (f. venella), 371 (right) a—e (f. typica).

DOUBTFUL DESCRIPTIONS. Razouls (1974): 236-237, Figs. 1A—H

(9), 2A-C (d); Humes (1988): only Fig. lc, e (as Oncaea praeclara).

TYPE LOCALITY. Western Mediterranean Sea, near Palermo.

MATERIAL EXAMINED.

Red Sea: see below under O. venusta f. typica and f. venella

Other areas:

(1) Gulf of Naples, Italy; collected 1 February 1967; depth 0-100

m; leg. B. Scotto di Carlo; identified by G.A. Heron: 1 9, 1d in

alcohol (RBS).

(2) Pacific Ocean, off Tanega Islands; R/V Toyoshio-maru; col-

lected 5 November 1994 with ORI-plankton net, mesh size 0.33

mm; oblique haul, depth 1600 m; leg. S. Ohtsuka: several 29

and 3d (RBS).

(3) Sea of Japan, Tassha Bay, Sado Island; collected 1978 from

colonies of Solandria secunda (Inaba), a hydroid; depth 10m;

leg. and identified by J.-s. Ho; 4 22 in alcohol (RBS).

(4) (a) North Pacific Ocean, Galapagos Rift, 00° 48.0'N, 086°

13.0'W; collected 7 December 1979; DSRV Alvin dive no.

990; depth 2451 m; 1 specimen in alcohol, labelled Oncaea

praeclara, Humes, 1988, holotype 2 (National Museum of

Natural History, Smithsonian Institution, Washington, DC,

reg. no. USNM 234 109): this vial contains Oncaea venusta

(1 9); parts of the specimen broke off during re-examination

and are mounted on slides in lactophenol [right antennule

(segment 4-6), left antenna (endopod segments 1 and 2),

right maxilliped (claw)].

(b) North Pacific Ocean, Galapagos Rift; further sampling data

not specified; 1 specimen in alcohol labelled: Acc.no. 37 66

47; O. praeclara, 13 August 1987 (USNM 234 111): this

vial contains Oncaea venusta (1 ®).

R. BOTTGER-SCHNACK

13.0'W; collected 5 December 1979; DSRV Alvin dive no.

998; depth 2450 m; 1 specimen in alcohol labelled Oncaea

praeclara, Humes, 1988, 1 d (USNM 234 112): this vial

contains Oncaea venusta (1 6, possibly f. typica).

(d) North Pacific Ocean, Galapagos Rift, 00° 47'N, 086° 08'W;

collected 24 January 1979; DSRV Alvin dive no. 883; depth

2493 m; 4 specimens in alcohol labelled Oncaea praeclara,

3 dd, 1 2(USMN 234114): this vial contains a mixture of

O. venusta (1 ovigerous 9, 1 3), Oncaea sp. (1 3, possibly

venusta f. venella), Triconia sp. (1 3).

(e) North Pacific Ocean, Galapagos Rift, 00° 48.25'N, 086°

13.48'W; collected 30 November 1979; DSRV Alvin dive

no. 983; (material donated to R. Bottger-Schnack by A.

Humes); 11 specimens in alcohol labelled Oncaea praeclara,

Humes 1988, 4 22,7 dd: this vial contains a mixture of O.

venusta (2 22, 3 3d), O. media (2 29), O. clevei (1 3),

Oncaeidae indet. (2 dd), Corycaeidae indet. (1 juvenile);

(RBS).

(f) East Pacific Rise, 12° 48.52'N, 56°48'W; collected 22

November 1987, Hydronaut Cruise, Nautile dive 221;

depth 2630 m (material donated to R. Huys, NHM, by A.

Humes); 9 specimens in alcohol labelled O. praeclara,

Humes, 5 29,5 dd: this vial contains 5 22(1 ovigerous)

and 4 d6 of O. venusta (all specimens empty exoskeletons

with no internal tissue).

(5) Sargasso Sea, near Bermuda, 31° 37.94'N, 64° 09.45'W; col-

lected 9 July 1998 with 1 m net, mesh size 0.15 mm; depth 210

m; leg. D. Steinberg: several 22 (ovigerous and non-ovigerous),

3 dé in alcohol, for molecular analysis (A. Bucklin, Durham,

New Hampshire); 2 22, 2 CV @@ in alcohol (RBS).

Philippi’s (1843) description of O. venusta was the first account of

an oncaeid species. It was based on a single male specimen col-

lected in the western Mediterranean, near Palermo. Unfortunately,

the specimen was lost by accident before the author could com-

plete the description. Giesbrecht (1892) redescribed O. venusta in

more detail from the Gulf of Naples and recently, Heron & Brad-

ford-Grieve (1995) provided an excellent redescription of the

species based on material from the Gulf of Naples as well as from

other localities in the Atlantic and Pacific Ocean. Both authors do

not mention the existence of different size variants among O.

venusta, although the range of size variation was quite large in

Giesbrecht’s specimens from the Pacific. In 1929, Farran

described two distinct size morphs of O. venusta, forma typica and

forma venella, from various Atlantic and Pacific locations, which

were separated mainly by their size and some minor morphologi-

cal characters. The two forms were distinguished in some of the

subsequent taxonomic descriptions (e.g. Tanaka, 1960; Ferrari,

1975; Boxshall, 1977b) and a third, ‘robust’ form was added by

Boxshall (1977b). In the Red Sea, both form variants of O. venusta

sensu Farran occur. They differ considerably in temporal and spa-

tial distribution (Béttger-Schnack, 1990b, 1995), thus indicating

the existence of reproductively isolated populations. A detailed

morphological examination of both size variants was undertaken

during the present study, including microstructures on the append-

ages and on the exoskeleton (pores, scales), partly by using SEM

analyses. The results pointed out several morphological diffe-

rences between the two forms not noted in the literature before,

but these were not regarded as sufficient to warrant recognition of

the two forms as separate species. In order to provide the

morphological basis required for future taxonomic analyses on O.

venusta, both forms are redescribed. A further approach to

differentiate the two forms using alternative methods, e.g. mole-

cular analysis, is currently in progress.

SEVEN SPECIES OF RED SEA ONCAEA

Oncaea venusta Philippi, 1843 forma typica Farran, 1929

Oncaea venusta Farran (1929): 284 [as Oncea venusta forma typica]

Oncea venusta forma typica Farran, 1929

TYPELOCALITY. Not specified, various locations in the temperate

and tropical Atlantic, as well as south of New Zealand.

MATERIAL EXAMINED.

Central Red Sea, 21° 22.09'N, 38° 05.09'E: Stn. 664; R/V Valdivia

leg 29: collected 21 February 1981 with MSN 0.1 mm net (Haul 218/

5); depth 0-50 m; total water depth ca 2000 m.

(a) 5 22,5 dé in alcohol (BMNH 1998.2777-2786).

(b) 5 22,5 3c in alcohol (ZMH K-39586).

1 3 in alcohol; 1 2, 1 d in mating position in alcohol (RBS).

DESCRIPTION. Note illustrations are based on (c).

ADULT FEMALE (Figs. 2-5, 8A—E, 9A-F).

Body length (measured in lateral aspect; from anterior margin of

rostral area to posterior margin of caudal rami, calculated as sum of

individual somites): 1615 um [traditional method 1130 um, range:

1000-1230 um, based on 14 specimens (Bottger-Schnack ef al.,

1989)].

Exoskeleton heavily chitinized, covered with numerous granules,

forming long raised structures (lines), especially along lateral part of

cephalic shield (Fig. 8A). Prosome 2.1 times length of urosome,

excluding caudal rami, 1.7 times urosome length including caudal

rami. P2-bearing somite without dorso-posterior projection in lat-

eral aspect (Fig. 2B). P3-bearing somite with conspicuous raised

pore protruding laterally (Fig. 2A). Other integumental pores on

prosome as indicated in Fig. 2A, B. Pleural areas of P4-bearing

somite with rounded posterolateral corners.

Proportional lengths (%) of urosomites 12.2 : 57.7: 8.5: 9.1:

12.5. Proportional lengths (%) of urosomites and caudal rami 10.0 :

47.0: 6.9: 7.3: 10.2: 18.6.

Genital double-somite 1.5 times as long as maximum width

(measured in dorsal aspect) and 1.9 times as long as postgenital

somites combined (Fig. 2C); largest width measured at anterior two-

thirds, lateral margins of genital double-somite rounded, posterior

part tapering gradually. Paired genital apertures located at about 2/5

the distance from anterior margin of genital double-somite; arma-

ture represented by | spine and 2 minute spinous processes (Fig.

8B), only 1 of which discernible under light microscope (Fig. 2G).

Double-curved sclerotization between, but slightly anterior to, geni-

tal apertures, pore pattern on dorsal surface as indicated in Fig. 2C.

Anal somite 1.6 times wider than long; about half the length of

caudal rami (Fig. 2C). Secretory pore discernible on either side of

anal opening and additional one near posterior margin. Anterior

margin of anal opening (vestigial anal opening) with transverse row

of minute denticles. Posterior margin of somite finely serrate ven-

trally and laterally (Fig. 2D). Ventral face with paired secretory pore

near posterior margin (Fig. 2C, cf. also Fig. 6E).

Caudal ramus (Fig. 2F) about 3.5 times as long as wide. Armature

consisting of 6 elements: antero- and posterolateral setae (II+III)

long, spiniform and unipinnate along medial margin; outer terminal

seta (IV) long and plumose; inner terminal seta (V) longest and

plumose; terminal accessory seta (VI) more than 2/3 the length of

outer terminal seta and 1.4 times longer than caudal ramus; dorsal

seta (VII) about same length as posterolateral seta (III) and less than

half the length of terminal accessory seta, plumose and bi-articulate

at base. Inner margin of somite with fringe of long setules. Dorsal

anterior surface (Fig. 2F) with secretory pore near insertion of seta

II. Dorsal surface covered with numerous small scales (Fig. 2F).

Antennule 6-segmented (Fig. 2E), relative lengths (%) of seg-

ments measured along posterior non-setiferous margin 8.4 : 27.6:

39.3 : 10.7 : 4.7 : 9.3. Armature formula: 1-[3], 2-[8], 3-[5], 4-

[3+ae], 5-[2+ae], 6-[6+(1+ae)]. Small element on segment 4

ornamented with row of scales along entire length, tip with tubular

extension (SEM observation, not figured). Small element on seg-

ment 6 (arrowed in Fig. 2E) with tubular extensions (Fig. 8C),

indicating sensory function.

Antenna 3-segmented, distinctly reflexed (Fig. 3A). Coxobasis

with row of long, fine spinules or setules near outer margin and with

few additional denticles on proximal and distal part of outer

(exopodal) margin, curved row of denticles on posterior face; with

bipinnate seta at inner distal corner. Endopod segments unequal in

length; proximal endopod segment subtriangular forming outer

lobate outgrowth bearing patch of branched tubular extensions (Fig.

8D, E), with row of denticles along posterior inner margin. Distal

endopod segment distinctly shorter than proximal endopod seg-

ment, with narrow cylindrical base articulating; with two patches of

branched tubular extensions along outer margin (Fig. 8D); lateral

armature with 4 elements, numbered using Roman numerals in Fig.

3A: | unipinnate spiniform seta (III) and 3 curved setae (I, II, IV),

setae I and II sparsely pinnate, seta I shortest; distal armature

consisting of 7 elements: | long curved unipinnate seta (E), 4

slightly curved unipinnate setae of graduated length (A—D), seta D

being shortest, and 2 slender naked setae (F and G), similar in length

and shorter than seta D; none of armature elements spiniform or

geniculate.

Labrum (Fig. 3B,C) distinctly bilobed. Each lobe with row of

minute denticles around outer ventral margin, row of long fine

setules latero-distally and dentiform processes converging and de-

creasing in size medially. Lobes separated by median concavity

covered anteriorly by overlapping rows of broad hyaline petaloid

flaps, flanked by paired slit-like pores on proximal part of each lobe

(Fig. 9A, B; position of pore arrowed in Fig. 3B). Anterior surface

with well developed integumental pockets (Fig. 9C) either side of

median swelling, free margin of pockets serrate (Fig. 9D); median

swelling with large secretory pore posteriorly. Posterior part of

medial incision ornamented with four rounded integumental

thickenings (Fig. 3C). Posterior surface with paired patch of very

long fine setules and 2 large secretory pores located on proximal part

of each lobe.

Paragnaths (Fig. 5, 9C) with small lateral extensions, anterior

margin densely covered with several rows of long setules, median

bulge unornamented.

Mandible (Fig. 3D) with few minute setules on surface of coxa;

gnathobase with 5 elements, indicated by capital letters in Fig. 3D:

1 at subdistal ventral -corner, 2 along distal margin and 2 along

subdistal dorsal margin; ventral element (A) shorter than ventral

blade, with long, fine setules along dorsal side; ventral blade (B)

strong and spiniform, with row of setules on posterior side; dorsal

blade (C) strong and broad, with 3 dentiform processes along distal

margin; dorsal elements setiform, the shorter (D) hyaline, flat and

densely setose, the longer (E) multipinnate.

Maxillule (Fig. 3E) indistinctly bilobed, with numerous spinules

on anterior and posterior surfaces. Inner lobe subcylindrical, with 3

elements: outermost one spiniform, swollen at base, fringed with

coarse spinules, others setiform and bipinnate; innermost one located

along concave inner margin at some distance from other elements.

Outer lobe with 4 elements; outermost element spiniform, curved

and bipinnate along inner proximal margin, unipinnate along distal

margin, longer than the following; other elements bipinnate or

naked, element next to the innermost shortest.

Maxilla (Fig. 3F) 2-segmented, comprising syncoxa and allobasis.

R. BOTTGER-SCHNACK

Ke Ky

——=

‘

RAN

SSS

————

Fig. 2 Oncaea venusta f. typica, female (Red Sea) (A) Habitus, dorsal (lateral raised pore enlarged); (B) same, lateral (appendages omitted); (C) urosome,

dorsal; (D) urosome, lateral; (E) antennule, small sensory element arrowed; (F) caudal ramus, dorsal, setae are numbered using Roman numerals; (G) P6.

SEVEN SPECIES OF RED SEA ONCAEA 33

Fos

Fig. 3 Oncaea venusta f. typica, female (Red Sea) (A) Antenna, posterior, lateral elements are numbered using Roman numerals, distal elements indicated

by capital letters; (B) labrum, anterior, slit-like pores arrowed; (C) same, posterior; (D) mandible, showing individual elements, identified using capital

letters; (E) maxillule; (F) maxilla; (G) maxilliped.

R. BOTTGER-SCHNACK

LL LIL LLL PLLA TP

SRE Ll

OS LE SS LEAS /

SE LaaassS Sisson Nye

Sa ERE R PLL wi AP LPL ne, :

Sei eer EP A Mee TT

LEMON Mea wl ELT PAMELA

SSS SRR NCES ~ ET Lekbthy SOLIS

IIS LIG SS 4

a ppivce L ty

SOK OS Z,

SRP Xe SN BEEK

LIE RSS SES,

LER OY,

SAS Ss ges Ca

Sn on ae

ERR

DROS

se"

50"

Weert

fens

iil

pte

G ee

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SSS ee

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SSS << =

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SSS

Fig. 4 Oncaea venusta f. typica, female (Red Sea) (A) P1, anterior [a, third endopod segment, showing aberrant spine number]; (B) P2, anterior; (C) P3,

anterior; (D) P4, anterior.

SEVEN SPECIES OF RED SEA ONCAEA

HHS

+50 um

Fig.5 Oncaea venusta f. typica, female (Red Sea) Paragnaths, ventral

view.

Syncoxa unarmed, surface ornamented with 2 spinular rows and 2

large secretory pores. Allobasis produced distally into slightly

curved claw bearing 2 rows of very strong spinules along medial

margin; outer margin with strong seta extending almost to tip of

allobasal claw, ornamented with few strong spinules distally and a

thin hyaline lamella bilaterally, tip of seta with tubular extension;

inner margin with slender pinnate seta and strong basally swollen

spine with double row of very strong spinules along the medial

margin and single row of shorter spinules along outer margin.

Maxilliped (Fig. 3G) 4-segmented, comprising syncoxa, basis

and 2-segmented endopod. Syncoxa ornamented with few spinules

on anterior surface, unarmed. Basis robust, inner margin with 2

spiniform spinulose elements nearly equal in length; fringe of long

setules half the distance between distal seta and articulation with

endopod; anterior surface with 2 rows of long spinules along palmar

margin, additional longitudinal row of spinules near outer margin as

in Fig. 3G. Proximal endopod segment unarmed. Distal endopod

segment drawn out into long curved claw, with pinnules along

proximal two-thirds of concave margin; accessory armature consist-

ing of minute, naked seta on outer proximal margin and unipectinate

spine fused basally to inner proximal corner of claw.

Swimming legs 1-4 biramous (Fig. 4A—D), with 3-segmented

rami. Spine and setae formula as for genus. Intercoxal sclerites well

developed, ornamented with 3 posterior denticles at distal corners in

P1—P3. Coxae and bases of legs 1-4 with surface ornamentation as

in Fig. 4A—D. Coxae of P1—P4 with posterior raised secretory pore

near outer distal corner (not figured, cf. Fig. 13A—D). Bases with

short naked (P1—P3) or plumose (P4) outer seta; with anterior

secretory pore near outer proximal corner; inner portion slightly

produced medially in P2—P4 (Fig. 4B—D). Inner basal seta on P1

spiniform and minutely pinnate. Respective legs without distinct

length differences between exopod and endopod (P1) or with endopod

slightly longer than exopod (P2—P4). Bases of spines on exopod and

endopod segments anteriorly surrounded by small spinules. Ante-

rior face of all segments ornamented with small scales (not figured),

similar to those on caudal ramus (Fig. 2F).

Exopods. Outer margin of exopod segments with well developed

serrated hyaline lamella, interrupted by small gap at about half the

distance, longitudinal concavity below serrated lamellar margin

(Fig. 9E); inner margin of proximal exopod segments with long

setules. Secretory pore present on posterior surface of distal seg-

ments. Hyaline lamellae on outer spines well developed; outer and

terminal spines of Pl with subapical tubular extension (Fig. 9F).

Terminal spine shorter than distal exopod segment in all legs.

Endopods. Outer margin of endopod segments with fringe of long

setules. Inner seta of proximal endopod segment slightly swollen at

base. Distal endopod segments with several secretory pores on

posterior surface; distal margin of P2 and P3 produced into conical

projection ornamented with minute spinules anteriorly (Fig. 4B,C).

Outer subdistal spine nearly equal in length to (P2—P3) or shorter

than (P4) outer distal spine, always shorter than distal spine. Outer

distal spine about 3/4 the length of terminal spine in P4. Outer

margin of distal segment of Pl terminating in a long process

obscuring insertion of distalmost inner seta (Fig. 4A). Inner setae of

distal endopod segments with spinule comb along proximal inner

margin; this comb also present on distal inner seta of middle

endopod segment in P3—P4.

PS as figured for O. venusta f. venella (cf. Fig. 11G), comprising

small plumose seta arising from lateral surface of somite, and small

free unornamented segment representing exopod. Exopod slightly

longer than wide, bearing 2 naked setae nearly equal in length,

subapical one spiniform.

P6 (Fig. 2G) represented by operculum closing off each genital

aperture; armed with a spine and 2 small spinous processes (Fig.

8B), only 1 of which discernible under light microscope (Fig. 2G).

Egg-sacs paired, oval-shaped; each sac containing appr. 30-40

eggs (diameter 50-60 tum).

ADULT MALE (Figs. 6, 7A—C, 8F, 10).

Body length: 1158 um [traditional method: 910 um, range: 880-950

um, based on 9 specimens (B6ttger-Schnack ef al., 1989)]. Sexual

dimorphism in antennule, antenna, maxilliped, genital segmentation

and ornamentation, P1—P3 (endopod), P5—P6, and in caudal ramus.

Prosome 2.0 times the length of urosome, excluding caudal rami,

1.7 times urosome length, including caudal rami.

Proportional lengths (%) of urosomites (excluding caudal rami)

10.1 : 64.5: 6.0: 4.4: 5.0: 10.1; proportional lengths (%) of uroso-

mites (caudal rami included) 8.6 : 55.0: 5.1 : 3.7: 4.3: 8.6: 14.7.

Length to width ration of genital somite 1.5 : 1. Caudal rami about

2.5 times longer than wide, shorter than in female. Caudal setae with

proportional lengths as in female, except for seta VI, which is about

2/3 the length of seta IV and 2.2 times the length of caudal ramus.

Dorsal and ventral surface of caudal ramus covered with minute

scales as in female. Surface of genital flaps and ventral surface of

anal segment ornamented with several rows of small spinules.

Ventral face of anal somite with paired secretory pore near posterior

margin (Fig. 6E).

Antennule (Fig. 6B) 4-segmented; distal segment corresponding

to fused segments 4-6 of female; relative lengths (%) of segments

measured along posterior non-setiferous margin 8.0 : 27.1 : 43.3:

23.6. Armature formula: 1-[3], 2-[8], 3-[4], 4-[11+2ae+(1+ae)].

Antenna (Fig. 6H) with seta on coxobasis naked and shorter than

in female. Distal endopod segment with seta III much stouter than in

female, seta IV spiniform and curved, both elements shorter than in

female (Fig. 8F).

Maxilliped (Fig. 6C) 3-segmented, comprising syncoxa, basis

R. BOTTGER-SCHNACK

Cw SL

LE KK

SSSS:

SKE

SIRS yr \

AVN

1 oy

LLLé fy y

y =o RK

Fig. 6 Oncaea venusta f. typica, male (Red Sea) (A) Habitus, dorsal, arrows indicating position of lateral raised pores; (B) antennule; (C) maxilliped,

anterior; (D) urosome, dorsal; (E) urosome, ventral; (F) same, lateral (spermatophores fully developed); (G) P5, dorsal; (H) antenna, posterior.

val ed,

SEVEN SPECIES OF RED SEA ONCAEA

20 um

ABCDEF

Fig. 7 Oncaea venusta f. typica, male (Red Sea) (A) P1, distal part of endopod; (B) P2, distal part of endopod; (C) P3, distal part of endopod. — Oncaea

venusta f. venella, male (Red Sea) (D) P1, distal part of endopod; (E) P2, distal part of endopod; (F) P3, distal part of endopod.

and |-segmented endopod. Syncoxa without surface ornamentation,

except for several secretory pores, unarmed. Basis robust, particu-

larly inflated in proximal half forming bulbous swelling; anterior

surface with 2—3 transverse spinular rows in addition to row of short

flat spinules along inner margin (Fig. 6C); posterior surface with 3

rows of short spatulated spinules of graduated length along palmar

margin (Fig. 6C); with 2 small naked setae within the longitudinal

cleft, nearly equal in length. Endopod drawn out into long curved

claw, concave margin unornamented; accessory armature consisting

of short, unipectinate spine basally fused to inner proximal corner of

claw; tip of claw with minute hyaline apex.

Swimming legs 1-4 with armature and ornamentation as in

female; sexual dimorphism expressed in terminal process on P| enp-

3, being longer than in female, reaching half the length of distal

spine (Fig. 7A) and in conical projections on distal endopod segment

of P2—P3, being longer than in female, reaching half the length of

outer distal spine (Fig. 7B, C).

P5 (Fig. 6G) exopod not delimited from somite, general shape and

armature as in female, except for subapical seta spiniform and

shorter than in female, ornamented with row of minute spinules

along outer margin.

P6 (Fig. 6E) represented by posterolateral flap closing off genital

aperture on either side; covered by pattern of denticles as shown in

Fig. 6E and 10A; with receptor (pores) at inner edge of posterola-

teral corners (Fig. 10B); these corners not protruding laterally so that

they are hardly discernible in dorsal aspect (Fig. 6D).

Spermatophore oval (Fig. 6F), of variable size according to state

of maturity; swelling of spermatophore during development not

affecting shape and relative size of genital somite.

Oncaea venusta Philippi, 1843 forma venella Farran, 1929

Oncaea venusta Farran (1929): 284-285, Fig. 33 (female only) [as

Onceea venusta forma venella].

Onceea venusta forma venella Farran, 1929

TYPE LOCALITY. Not specified, various locations in the temperate

and tropical Atlantic, as well as south of New Zealand (Farran,

1929).

MATERIAL EXAMINED

(1) Central Red Sea, 21° 22.09'N, 38° 05.09'E: Stn. 664; R/V

Valdivia leg 29: collected 21 February 1981 with MSN 0.1 mm

net (Haul 218/5); depth 0-50 m; total water depth ca 2000 m.

(a) 5 29,5 dd in alcohol (BMNH 1998.2787-2796)

(b) 5 29,5 dd in alcohol (ZMH K-39587)

(2) Southern Red Sea, 15° 34.8'N, 41° 54.9'E: Stn. 703; R/V Meteor

leg 5/5: collected 03 August 1987 with MSN 0.05 mm net (Haul

39/5); depth 0-50 m; total water depth 970 m: 1 6 in alcohol

(RBS).

(3) Southern Red Sea, 15° 34.8'N, 41° 54.9'E: Stn. 703; R/V Meteor

leg 5/5: collected 03 August 1987 with MSN 0.05 mm net (Haul

39/4); depth 50-100 m; total water depth 970 m: 2 99,2 dd

dissected on slides, 1 2, 1 3 in alcohol (RBS).

DESCRIPTION. Note illustrations are based on (3).

ADULT FEMALE (Figs. 11-13)

Body length (measured in lateral aspect; from anterior margin of

38 R. BOTTGER-SCHNACK

» Magn:

/ B000x

Fig. 8B = Oncaea venusta f. typica, female (Red Sea) Genital aperture, right;

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 8C Oncaea venusta f. typica, female (Red Sea) Antennule, 6th segment, short sensory element.

Fig. 8D Oncaea venusta f. typica, female (Red Sea) Antenna, endopod segment | + 2, concavity and patches of tubular extensions arrowed.

40 R. BOTTGER-SCHNACK

Fig. 8E Oncaea venusta f. typica, female (Red Sea) Antenna, endopod segment 2, branched tubular extensions (arrowed).

Fig. 8F Oncaea venusta f. typica, male (Red Sea) Antenna, endopod segment 2, lateral armature.

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 9A Oncaea venusta f. typica, female (Red Sea) Labrum, anterior, showing integumental pockets either side of median swelling and right slit-like

pore (arrowed).

Fig.9B Oncaea venusta f. typica, female (Red Sea) Labrum, anterior, slit-like pore, right.

42 R. BOTTGER-SCHNACK

Fig.9C Oncaea venusta f. typica, female (Red Sea) Oral area, ventral.

Fig.9D Oncaea venusta f. typica, female (Red Sea) Labrum, anterior, integumental pocket, right.

SEVEN SPECIES OF RED SEA ONCAEA 43

Fig.9E Oncaea venusta f. typica, female (Red Sea) P2—P4, exopod-1, showing lateral concavity (arrowed).

Fig. 9F Oncaea venusta f. typica, female (Red Sea) P1, exopod-1, tip of spine, showing tubular extension.

R. BOTTGER-SCHNACK

Fig. 10A Oncaea venusta f. typica, male (Red Sea) Caudal ramus and P6, ventral.

Fig. 10B Oncaea venusta f. typica, male (Red Sea) P6, posterolateral corners, showing receptor pores at inner edge.

SEVEN SPECIES OF RED SEA ONCAEA

Loo7

Fig. 11 Oncaea venusta f. venella, female (Red Sea) (A) Habitus, dorsal: (B) same, lateral (appendages omitted; dorsoposterior swelling on P2-bearing

somite arrowed); (C) urosome, dorsal; (D) urosome, lateral; (E) antennule; (F) caudal ramus, dorsal: (G) P5, dorsal.

rostral area to posterior margin of caudal rami, calculated as sum of

individual somites): 1096 um [traditional method: 800 um, range:

750-880 um, based on 36 specimens (B6ttger-Schnack et al. 1989)].

Body proportions different from f. typica, with prosome 2.5 times

length of urosome, excluding caudal rami, 2.0 times urosome length

including caudal rami. P2-bearing somite with insignificant

dorsoposterior swelling in lateral aspect (arrowed in Fig. 11B).

Integumental pores on prosome as indicated in Fig. 11A, B. Gran-

ules and raised structures on exoskeleton less pronounced than in f.

typica.

Proportional lengths of urosomites similar to f. typica. Genital

double-somite 1.8 times as long as maximum width (measured in

dorsal aspect); largest width measured at anterior third, lateral

margins of genital double-somite weakly rounded at anterior third,

posterior part nearly straight. Double-scalloped sclerotization

between genital apertures, pore pattern on dorsal surface as indi-

cated in Fig. 11C.

Anal somite (Fig. 11C) as in f. typica.

Caudal ramus (Fig. 11F) about 2.8 times as long as wide, shorter

than in f. typica. Proportional length of caudal setae as in f. typica,

except for seta VI 1.6 times longer than caudal ramus.

Antennule (Fig. 11E) and antenna (Fig. 12A) as in f. typica,

except for slight differences in the ornamentation of setae on second

endopod segment in the antenna (seta I; setae C and D).

Labrum (Fig. 12B, C) similar to f. typica, except for integumental

pockets with free margin serrate along half the distance only (Fig.

12B).

Mandible (Fig. 12D), maxillule (Fig. 12E), maxilla (Fig. 12F) and

maxilliped (Fig. 12G) as in f. typica.

Swimming legs 1-4 (Fig. 13A—D) as in f. typica, except for P4

enp-3 outer distal spine only 2/3 the length of distal spine. Distal

spines on endopods of P1—P4 somewhat longer than in f. typica.

PS (Fig. 11G) and P6 as in f. typica.

Egg-sacs paired, containing appr. 30-35 eggs each (diameter 40—

50 um).

ADULT MALE (Fig. 7D-F, 14).

Body length: 985 um [traditional method: 590 um, range: 550-650

um (single extreme value 730 um), based on 17 specimens (Bottger-

Schnack ef al. 1989)]. Sexual dimorphism in antennule, antenna,

maxilliped, PI—P3 (endopod) and P5—P6, caudal ramus and in

genital segmentation.

Prosome 2.3 times length of urosome, excluding caudal rami, 2.0

times urosome length including caudal rami. Proportional lengths

(%) of urosomites similar to f. typica. Length to width ratio of

genital somite 1.7 : 1, longer than in f. typica. Ornamentation of

genital flaps as in Fig. 14E. Caudal rami about 1.9 times longer than

wide, shorter than in f. typica. Caudal setae with proportional

lengths as in f. typica, except for seta VI less than 2/3 the length of

seta IV and 2.4 times longer than caudal ramus.

Sexual dimorphism in antennule (Fig. 14B), antenna (Fig. 14G),

maxilliped (Fig. 14C) and in PS—P6 (Fig. 14D, E), as well as in enp-

3 of P1—P3 (Fig. 7C, D) similar to f. typica. Seta on body near P5

naked.

TAXONOMY

The original description of O. venusta by Philippi (1843) was rather

poor and the single male specimen was lost by accident before the

mouthparts could be documented. Dana’s subsequent (1849, 1852)

records of Antaria obtusa and A. crassimana were included under the

synonymies of both O. venusta and O. mediterranea by Giesbrecht

(1892), reflecting the author’s undecisiveness on this matter. Lubbock

(1860) described the species (as Oncea pyriformis) from the equato-

rial Atlantic and the southern Indian Ocean, and synonymized it with

R. BOTTGER-SCHNACK

Dana’s A. obtusa. He was the first to record male-female pairs

(‘couples’) of the species and erroneously believed that he had also

been the first author to report the males; however, as the original

account of O. venusta by Philippi was based on a male specimen,

Lubbock’s opinion was erroneous. Claus’ (1866) description of

Antaria coerulescens from Nice has been synonymized with O.

venusta by Giesbrecht (1892). Claus described several characters,

such as the heavily sclerotized exoskeleton and its surface ornamen-

tation, which are typical for the species, but at the same time recorded

the P4-bearing somite as being heart-shaped and pointed, which is not

the case in O. venusta. Brady’s (1883) illustrations [Challenger

Expedition] of Antaria obtusa (Dana), were also synonymized with

O. venusta by Giesbrecht (1892), but he suspected that Brady’s

specimens might include O. mediterranea as well. According to

Brady’s illustration (Fig. 11), the P4 endopod exhibits a length ratio

of distal spine to outer distal spine (1.2: 1) more similar to O. venusta

(1.3: 1) than to O. mediterranea (1.55: 1). His illustration of the male

urosome (Fig. 3), however, shows laterally produced genital flaps,

which is not typical for O. venusta, but can be observed in O.

mediterranea (Bottger-Schnack & Huys, 1997, their Fig. 4A, D).

Giesbrecht (1892) redescribed O. venusta on the basis of material

from Naples, reviewed the earlier literature on Antaria and Oncaea

and summarized the synonymies of the respective species known at

that time (see above). He stated that, judging from its general

habitus, Dana’s form variety of A. gracilis would also resemble O.

venusta. However, as the P4-bearing somite of Dana’s specimen is

pointed in lateral view (Plate 86, Fig. 12) as in O. mediterranea, and

not rounded as in O. venusta, Giesbrecht’s opinion is not followed

here. [Dana’s typical A. gracilis (Plate 86, Fig. 11b) is figured with

egg-sacs attached, which conceal great parts of the urosome. Thus

its unusual, narrow form cannot be used as a specific character for

identification as was proposed by Giesbrecht.]

Recently, an excellent redescription of O. venusta was provided

by Heron & Bradford-Grieve (1995), based on material from the

Gulf of Naples and from various locations in the Atlantic and

Pacific. The authors did not figure the minute element on the 6th

segment of the antennule, and did not report on the sexual dimor-

phism in the coxobasal seta on the antenna and in the endopodal

spines of P1—P3. Re-examination of their material (1 and 1 d from

Naples, kindly put at my disposal by G.Heron) showed, however,

that these characters are also present in O. venusta from Naples and

that the specimens from the Red Sea are conspecific. A further

discussion of their material will be given below under Form variants.

FORM VARIANTS

Farran (1929) distinguished two form variants of O. venusta, f.

typica and f. venella, which differed mainly in size: the typical form

measured 1|.08—1.16 mm (9) and 0.78-0.81 mm (<) in total body

length, while females of the venella form measured 0.91—1.07 mm.

No males of the venella form were recorded by Farran (1929).

Females of the two forms differed furthermore in general habitus, f.

typica being more ‘pear-shaped’ than f. venella, and in the length

ratio of prosome : urosome, which was smaller in f. typica (1.3:1)

than in f. venella (1.5:1).

Sewell (1947) recorded two size variants among female O. venusta

in the northern Arabian Sea and found no overlap in size between the

two groups (1.18—1.25 mm and 0.85—0.91 mm, respectively). He

described some morphological differences other than size, which

included (1) length to width ratio of prosome, which was larger in f.

venella than in f. typica, (2) length to width ratio of caudal ramus,

which was larger in f. typica than in f. venella and (3) slight

differences in the proportions of urosomites and caudal ramus

between the two forms.

SEVEN SPECIES OF RED SEA ONCAEA

“3

N35

Fig. 12 Oncaea venusta f. venella, female (Red Sea) (A) Antenna, anterior; (B) labrum, anterior; (C) same, posterior; (D) mandible, showing individual

elements; (E) maxillule; (F) maxilla; (G) maxilliped.

R. BOTTGER-SCHNACK

WERK

pee

SOY

St,

Q EES

SPER PEELE

PP ERSERS SS

usin

Z Z 5 SI ALLE}, /,

WES ye SLE Lie hig PUES ora

mes 2S = ee

LOS

<= ‘

female (Red Sea) (A) P1, posterior; (B) P2, posterior, intercoxal sclerite omitted; (C) P3, anterior; (D) P4, posterior.

Fig. 13. Oncaea venusta f. venella

SEVEN SPECIES OF RED SEA ONCAEA 49

Fig. 14 Oncaea venusta f. venella, male (Red Sea) (A) Habitus, dorsal; (B) antennule; (C) maxilliped, anterior; (D) urosome, dorsal (spermatophores not

fully developed); (E) urosome, ventral; (F) same, lateral; (G) antenna, posterior.

Specimens of O. venusta from the Red Sea exhibited a corre-

sponding difference in size (Table 2) with no overlap between the

two groups. No distinct regional or seasonal differences in the

lengths of the two forms were observed, although specimens of f.

venella in the central Red Sea tended to be smaller in summer as

compared to autumn and winter (Table 2). Generally, Red Sea

specimens of f. venella are smaller than elsewhere, which might be

related to the extreme environmental conditions in this basin (Bottger-

Schnack etal., 1989). Sizes of f. typica, on the other hand, correspond

well to those reported from other regions (B6ttger-Schnack ef al.,

1989). A possible explanation for this may be that the large morph

penetrates the Red Sea during a limited period only (see below under

Ecological notes) and does not survive as an indigenous population

in this area.

Alternative explanation could be that smaller individuals of f.

venella may not have been collected during earlier investigations,

because nets with fairly large mesh sizes of about 0.3 mm were used

(e.g Boxshall, 1977b). However, this would not explain the absence

of intermediate sizes, measuring between 0.9—1.0 mm ( 2) and 0.75—

0.85 mm (d) in length, in O. venusta from the Red Sea.

Morphological differences other than size between the two forms

of O. venusta females from the Red Sea were similar to those

recorded by Farran (1929) and Sewell (1947). Additional differen-

ces in body morphology found in the present study include (1) the

length to width ratio of female genital double-somite, which is

smaller in f. typica (1.5:1) than in f. venella (1.8:1), (2) the small

dorso-posterior protrusion of the P2-bearing somite in female f.

venella, which is not found in f. typica, (3) small differences in the

relative lengths of distal spines on P4 endopod, which are likewise

found in both sexes, and (4) the length to width ratio of male genital

somite, which is smaller in f. typica (1.5 : 1) than in f. venella (1.7:

1). The latter two characters are the only differences other than size,

by which males of the two forms can be separated. The dorsoposterior

swelling on the P2-bearing somite in f. venella was consistent for all

specimens from the Red Sea and was also found in f. venella from

the northern Arabian Sea. Altogether, the results pointed to several

additional morphological differences between the two forms, which

had not been noted in the literature before, but these were not

Table 2 Body length (mm) of O. venusta in the Red Sea.

Form variant Sex n xX R

AUTUMN Northern Red Sea

f. venella F 15 0.80 0.75-0.85

M 6 0.58 0.57-0.59

Central Red Sea

f. venella F 24 0.80 0.75-0.85

M 4 0.61 0.57-0.73

WINTER Central Red Sea

f. venella F 12 0.80 0.75-0.88

M 16 0.59 0.55—0.65

f. typica F 14 ii 1.00—1.23

M 7 0.92 0.88—0.95

SUMMER ~ Central Red Sea

f. venella F 10 0.76 0.70-0.80

M 1 0.56

Gulf of Aden + Strait of Bab al Mandab

f. venella F 14 0.83 0.74-0.92

M 3 0.60 0.58—0.63

f. typica F 5 Ie JS) 1.10-1.22

n=no. of individuals measured; X = mean; R = range

R. BOTTGER-SCHNACK

regarded as sufficient to warrant distinction of the two forms as

separate species.

Identification of O. venusta f. typica males during routine counts

in plankton samples is facilitated by their great size and overall

robust appearance. Males of f. venella, on the other hand, are very

similar to males of O. clevei, which are described in the present

account for the first time (see below).

COMPARISON OF O. VENUSTA FORM VARIANTS WITH SPECIMENS

FROM THE MEDITERRANEAN

Total body length of specimens from Naples recorded by Giesbrecht

(1892) and Heron & Bradford-Grieve (1995) ranged between 1.09—

1.27 mm for females, and 0.8-0.95 mm for males. This size range

corresponds to that of O. venusta f. typica and most previous authors

have regarded the mediterranean specimens as being conspecific

with the typical form (e.g. Sewell, 1947; Tanaka, 1960). A compa-

rison of morphological characters other than size between the two

form variants from the Red Sea and specimens from the Gulf of

Naples are summarized in Table 2. The Mediterranean specimens in

fact seem to be more similar to the typical form than to the venella

form. However, they also share some characters with f. venella, such

as the form of the sclerotized structure between female genital

apertures and the pore pattern of the male urosome. Two morpho-

logical characters of the Naples specimens were intermediate between

the two Red Sea forms, the length to width ratio of the caudal ramus

in both sexes and the length to width ratio of the genital somite in the

male (Table 2). Based on these observations, it cannot be confirmed,

which of the two form variants from the Red Sea is conspecific with

O. venusta Giesbrecht sensu Heron & Bradford-Grieve. The length

to width ratio of caudal ramus has been found to be very variable

among specimens of O. venusta (Boxshall, 1977b) and this might

also apply to the pore pattern. Malt (1983c) investigated the

integumental pore patterns of females of the two venusta forms,

based on material collected in the Atlantic, however, did not find any

significant differences between them and/or the third ‘robust’ form

variety. Males were not investigated during her study.

OTHER RECORDS OF O. VENUSTA FORM VARIANTS

The geographical distribution of the size morphs of O. venusta is

poorly documented (Malt, 1983a). Farran (1936) reported a great

size variation in specimens from the Great Barrier Reef, but could

not distinguish the two form variants, which he previously had

described from the temperate and tropical Atlantic and off New

Zealand, because many specimens intermediate in size occurred.

Sewell (1947) recorded two variants of O. venusta from the

northern Arabian Sea (discussed above), and concluded that they

might have slightly different breeding seasons, since both exhibited

different proportions of ovigerous females and of females bearing

spermatophores in the samples. He summarized the geographical

distribution of O. venusta known at that time and concluded that the

smaller f. venella form was absent in the Mediterranean. However,

both forms of O. venusta were recorded from Lebanese waters by

Malt et al. (1989) without further descriptive details. In the quanti-

tative study of Bottger-Schnack (1996) conducted in the Eastern

Mediterranean, O. venusta was totally absent.

Tanaka (1960) recorded two size groups (1.13—1.39 mm and

0.90-1.0 mm, respectively) among female O. venusta from the

Indian Ocean and off Cape of Good Hope, as well as from Antarctic

waters; specimens from Japanese waters (South China Sea) be-

longed to the typical form only. He did not find any structural

differences between the two forms, except for a somewhat more

slender prosome in f. venella as compared to f. typica. The caudal

rami were four times longer than wide in both forms, irrespective of

sex; this is unusual for species of Oncaea s. str., which typically have

~~ os

SEVEN SPECIES OF RED SEA ONCAEA

Table 3. Comparison of morphological characters of Oncaea venusta Giesbrecht from the Gulf of Naples with two forms, f. typica and f. venella, from the

Red Sea.

Gulf of Naples Red Sea

Species/form f. typica f. venella

Female

Ornamentation on surface of prosome (ridges, etc) very strong strong present, less strong

P2-bearing somite with dorso-posterior swelling no no yes

Genital double-somite

1: w ratio 14:1 [oral 1.8:1

anterior part produced dorsally yes yes no

sclerotization between gen.ap.

—form double-scalloped paired s-shaped double-scalloped

— location between gen.ap. anterior to gen.ap. between gen.ap.

Caudal ramus, | : w ratio SO Sieleill Poieall

P4 enp-3, ratio of DS:ODS Nes wei teehee ifesyel

Male

Genital segment

1: w ratio eel is) | Ia

no. of pores on dorsal surface 5 I]

Caudal ramus, | : w ratio BED ish PFs ra ROR

DS = distal spine; ODS = outer distal spine; gen.ap. = genital apertures; P2, P4 = swimming legs 2, 4; enp-3 = third endopod segment; | = length; w = width; no. = number

asmaller length to width ratio of caudal ramus in the male (see above

under ‘Generic diagnosis’). The length to width ratio of the male

genital segment was greater inf. venella (1.6: 1) than inf. typica (1.3

: 1), which is in accordance with results from the Red Sea. The body

lengths of males in Tanaka’s study ranged between 0.74—1.07 mm,

with no separation given for the two groups.

Corral Estrada (1970) recorded two forms of female O. venusta

from the NE Atlantic, near Tenerife, which differed mainly in size

(1.05—1.25 mm and 0.87—0.95 mm). Small differences were also

found in overall body proportions, the venella form being more

slender.

Ferrari (1975) recorded two size groups for both sexes of O.

venusta from the Gulf of Mexico, measuring 1.1—1.2 mm or 0.92—

0.99 mm (females) and 0.76—0.86 mm or 0.57—0.63 mm (males).

Both groups occurred over the entire period (4 yrs) of his study,

without exhibiting any overlap in size. No morphological differ-

ences other than size were noted by the author. His illustration of the

male antenna of the typical form (his Fig. 6D) shows a long, plumose

seta on the coxobasis, not the small naked one usually found in O.

venusta (cf. Fig. 6H, 14G).

Boxshall (1977b) reported both size morphs of female O. venusta

from the NE Atlantic, near the Cap Verde Islands, and in another

report gave detailed information on their vertical distribution and

diurnal vertical migration (Boxshall 1977a). The two forms differed

only in size, with a mode length of 1.13 mm for f. typica and 0.98

mm for f. venella. No other detectable differences were recorded.

The length frequency distribution of both forms (his Fig. 13) shows

very little overlap in size between the two groups. Males were not

distinguished into size groups during his study, their length ranged

between 0.96-1.08 mm with a mean of 1.01 mm. Both sexual

dimorphic characters of the male antenna (naked seta on coxobasis

and modified seta on lateral armature) were illustrated by Boxshall

(his Fig. 12b). The terminal accessory seta on the male caudal

ramus, however, was figured as being less than twice the length of

CR, whereas it is more than twice the length in O. venusta from

Naples and the Red Sea. In addition to the two forms known at that

time, Boxshall also recorded a few female specimens of a third

‘robust form’, which was more squat in general appearence and

differed in the length to width ratio of the caudal ramus. The length

of the specimens ranged between 0.88—1.4 mm, thus covering the

length range of both other varieties of O. venusta. The ‘robust form’ ,

however, seemed to have a limited distribution range, because it was

not discovered again in subsequent collections from other North

Atlantic localities (Malt 1983c).

Recently, Itoh [in: Chihara & Murano (1997)] recorded both size

morphs of O. venusta from Japanese waters. His descriptions of

female and male habitus exhibit the same differences in overall body

morphology between the two morphs as recorded here for Red Sea

specimens. Also, Itoh’s illustration of the male antenna [Plate 223,

Fig. 371 (right) c] shows the two sexual dimorphic characters typical

for the species. No overlap in size between the two groups was noted

by Itoh, females measured 1.09-1.23 mm (f typica) or 0.86—0.94

mm (f. venella) and males 0.89-0.98 mm (f. fypica) or 0.62—0.65

mm (f. venella). Specimens of female O. venusta from the Sea of

Japan examined during the present study, however, covered a wide

size range from 0.80—1.3 mm, and many specimens intermediate in

size (about 0.96 mm) occurred. Specimens at the upper and lower

end of the size range exhibited nearly all morphological characters

of f. typica and f. venella, respectively, while specimens intermedi-

ate in size could be assigned to neither morph. [The same phenomenon

was observed in O. venusta specimens from Australian waters

(McKinnon material). |

In summary, the two distinct size morphs of O. venusta, f. typica

and f. venella, are widespread in tropical and temperate regions of

the Atlantic (Corral Estrada, 1970; Ferrari, 1975; Boxshall, 1977b)

and the western Indian Ocean and adjacent seas, including the Red

Sea (Sewell, 1947; Tanaka, 1960; this report). Records of two

corresponding size groups from the Eastern Indian Ocean and the

Pacific are rare, because intermediate size forms occur, which make

a clear separation of the groups more difficult. This had already been

noted by Farran (1936), who found specimens of O. venusta from

the Great Barrier Reef covering a wide size range, without being

able to separate them into the two groups. Further studies on O.

venusta from Pacific localities are required in order to define their

morphological relationship to Atlantic and western Indian Ocean

populations. The present results, based on both light and SEM

microscopy, pointed to several morphological differences between

the two venusta forms, which had not been noted before. However,

these were not regarded as sufficient to warrant recognition of the

two forms as separate species. Alternative taxonomic techniques,

such as molecular analysis, may permit examination of any genetic

differentiation between the forms of this ubiquitous species. A study

on the molecular genetics of O. venusta size variants from the

Atlantic is in progress and future morphological studies in other

areas are recommended.

OTHER RECORDS OF O. VENUSTA

O. venusta is widely distributed at mid- and low latitudes, between

approximately 50°S and 65°N (Malt, 1983a, b). Due to its relatively

large size and the characteristic habitus of the female, it is one of the

best documented oncaeid species in the world.

Halim (1969) listed O. venusta as one of the ‘perennial-indig-

enous’ species of the Red Sea, being distributed throughout the main

basin and the Gulf of Suez all year round. His compilation of earlier

records did not differentiate between the two forms of the species,

which were found to exhibit considerable differences in regional and

seasonal distribution in the Red Sea (see below under Ecological

notes).

Wilson (1932) recorded O. venusta as the most abundant species

of the genus in the Woods Hole region. The length range of his

specimens (9%. 1.1-1.27 mm, dd: 0.8-0.95 mm) indicates that he

had collected the typical form, however, his illustration of dorsal

aspect of the female (Fig. 213A; from W.M. Wheeler) shows the

form of genital double-somite to be more similar to f. venella.

In the Western Pacific area, Mori (1937, reprinted 1964) figured

O. venusta from Japanese waters and Dakin & Colefax (1940)

recorded it as the commonest species in the coastal waters of

Australia (New South Wales). Chen et al. (1974) described the

species from the Yellow Sea and the East China Sea. Their illustra-

tion of P4 endopod (Plate 6, Fig. 3) shows 2 inner setae on distal

segment which is not found in any species of Oncaea s. str. This

requires confirmation. The length range reported by Mori (22 1.0-

1.28 mm, dd: 0.8-1.0 mm) was smaller than the sizes reported by

Chen et al. (22 1.2-1.35 mm, dc: 0.9-1.1 mm), but the dorsal

habitus of females illustrated by these authors indicate that both

were dealing with f. typica.

Olson (1949) recorded O. venusta from the East Pacific, off

Oregon, which according to its size (29. 1.25 mm, dd: 0.9 mm) and

female habitus might belong to the typical form. The author did not

note any sexual dimorphism on the antenna or on the swimming

legs.

Razouls (1974) figured O. venusta from the region of Banyuls-

sur-mer (Golfe du Lion) and summarized previous records from the

Mediterranean Sea. His illustrations lack many details, such as the

genital apertures on the female genital double-somite, several ele-

ments on the antennule, antenna, P5 and the basal seta on P2—P4;

moreover, the distal endopod spines in P2—P4 of his specimens are

figured much longer than usually found in O. venusta (especially in

P4). Further differences between his specimens and typical characters

of O. venusta are found in the female caudal ramus, which is less

than 3 times longer than wide and the length of caudal seta VI, which

is unusually short in both sexes. The coxobasal seta on the male

antenna was figured as long and plumose by Razouls, which is not

the case in O. venusta. In summary, positive identification of his

specimens can only be given after re-examination of his material.

Ho (1984) redescribed O. venusta based on several females

collected from colonies of Solandria secunda (Inaba), a hydroid

found at 10 m depth in Tassha Bay, Sado Island (Japan). His

description differs from the present account in the setal formula of

P1 endopod (4 instead of 5 setae on distal segment), in the armature

of the antennule (some elements missing on segment 4 and 5), and in

the armature of P6 (2 minute spinous processes not mentioned).

Four females from his collection were kindly made available by J.-

s. Ho. Re-examination showed that all specimens from Sado Island

exhibit the typical setal formula on P! enp-3 (5 setae). The insertion

R. BOTTGER-SCHNACK

of the distalmost seta is hidden beneath the long terminal process (cf.

Fig. 4A) and thus can easily be overlooked. Also, the armature of the

antennule and on P6 is the same as described here. Due to their large

size, Ho ascribed his specimens to f. typica and his opinion is

followed here, although some characters, such as proportional spine

lengths on P4 enp-3 were inconclusive in the two smaller specimens

(0.92—0.96 mm) examined.

Humes (1988) described both sexes of a new species of Oncaea

s.str., O. praecalara, collected with slurp guns or box corers during

deep dives by manned deep-sea submersibles from the vicinity of

deep-sea hydrothermal vents in the eastern Pacific. The main

characters for the new species as summarized by Humes are: (1) its

relatively large size [1.01—1.3 mm in the female], (2) extremely long

caudal rami, about twice longer than anal somite and exhibiting a

length to width ratio between 4.96-6.7 : 1 (measured halfway down

the CR length), (3) sexual dimorphism on third segment of male

antenna (not on fourth segment, as erroneously stated in his ab-

stract), (4) labrum with nearly straight posteroventral margins. The

female holotype as well as female and male paratypes of O. praeclara

from the collection of the National Museum of Natural History,

Smithsonian Institution, were re-examined, in addition to several

samples of the species kindly made available by A.G. Humes. The

following measurements taken of the female holotype gave strong

evidence that O. praeclara is conspecific with Oncaea venusta: (1)

length to width ratio of CR 4.2 : 1 (measured halfway down the CR

length), 3.5 : 1 (measured proximally) [as reported in the present

account], (2) maxillipedal basis with spinular row along palmar

margin (not smooth as described by Humes, his Fig. 2h), (3) labrum

bilobate [ornamentation on lobes not discernible without dissection

of specimen], integumental pockets and slit-like pores on anterior

face present. The male paratype of O. praeclara can be assigned to

O. venusta on the basis of (1) the sexual dimorphism of antenna,

with coxobasal seta short and naked (not noted by Humes), seta I on

second endopod segment curved, and (2) the pore pattern on dorsal

surface of genital segment (11 pores).

Humes’ account of O. praeclara may have been based on two

different species: the form of the genital double-somite and the

position of the genital apertures differ considerably between his Fig.

la (habitus, dorsal) and Fig. 1c (urosome, dorsal). The extremely

elongate CR noted by Humes (his Fig. lc, e) was found neither in the

female holotype nor in any of the specimens re-examined, but may

have been present in other specimens. Fig. 1c and le (caudal ramus)

as well as Fig. 2h (maxilliped) of Humes (1988) are regarded here as

belonging to a different species, which, however, could not be traced

in his material, although some of the samples contained a mixture of

oncaeid species (see ‘Material examined’ above). The remaining

figures of Humes seem to be identical with O. venusta. Like Ho

(1984), Humes erroneously figured the distal endopod segment of

P1 with 4 outer setae, instead of 5. He also missed one seta on the

distal armature in the male antenna.

Huys & Boxshall (1991) illustrated several appendages and a

dorsal view of the female urosome of O. venusta, based on speci-

mens from the North Atlantic. Their Fig. 2.10.10.A of the mandible

shows 4 dentiform processes along the distal margin, whereas only

3 processes are found in specimens from the Red Sea (Fig. 3D, 7D)

and in those from the Gulf of Naples (Heron & Bradford-Grieve

1995, their Fig. 14J). The authors erroneously figured the female P5

with | exopodal seta only (their Fig. 2.10.21.B), not with 2 setae as

usually found in O. venusta.

Many more records of O. venusta from different localities of the

world ocean are known [see Malt (1983a) for a review], but are not

considered here, because they did not include figures or a descrip-

tion that positively identified the species.

- - oe

SEVEN SPECIES OF RED SEA ONCAEA

ECOLOGICAL NOTES

O. venusta f. venella is much more abundant than f. typica in the Red

Sea, exceeding the population densities of the latter by a factor of

between 3 to 100, when both forms co-occur (Bottger-Schnack,

1990b, 1995).

GEOGRAPHICAL DISTRIBUTION

In the Red Sea, O. venusta f. venella is more widespread than O.

venusta f. typica, it occurs throughout the main basin (Bottger-

Schnack, 1990a, b, 1995) and was also found in samples from the

northernmost part of the Gulf of Aqaba (unpubl. data). O. venusta

f. typica is mainly restricted to the southernmost Red Sea and the

Gulf of Aden, reaching the central Red Sea only during the winter

months, when a strong seasonal inflow of southern Red Sea waters

influences the plankton fauna in the central area (BOttger-Schnack,

1990b; see also Weikert, 1987; Beckmann, 1996).

VERTICAL DISTRIBUTION AND VERTICAL MIGRATION

O. venusta is mainly distributed in the epipelagic zone in the Red

Sea, with maximum abundances in the 0-100 (150) m depth layer

(B6éttger-Schnack, 1990 a, b). Occasional finds of the species in the

bathypelagic zone (unpubl. data, see also Beckmann, 1996) were

usually regarded as moribund specimens or may be due to contami-

nation of the nets.

When co-occurring, the two forms of O. venusta tend to be

vertically separated: Both sexes of O. venusta f. typica were concen-

trated in the upper epipelagic zone at 0-20 m (Strait of Bab al

Mandab) or at 0-40 m (Gulf of Aden) during summer, whereas

female f. venella occurred deeper in the water column, with maxi-

mum concentrations at 20-60 m (Bab al Mandab) and (20)80—100 m

(Gulf of Aden), respectively. Male f. venella occurred at the same

depth horizon as females in the Gulf, but were concentrated some-

what shallower than females (O40 m) in the Strait, thereby extending

into the depth horizon where f. typica dominates. No corresponding

vertical separation of the two forms became obvious in the central

Red Sea during winter, when both forms stayed in the upper 50 m

during day and night (Bottger-Schnack, 1990b). However, the sam-

pled depth strata in the epipelagic zone were broader during winter

(SO m-intervals) than during summer (20 m-intervals) and thus a

vertical segregation may not have been detected due to the limita-

tions in the sampling strategy.

In the central and northern Red Sea, O. venusta f. venella was

mainly concentrated in the lower epipelagic zone, within and

below the strong seasonal thermocline, during autumn (Bottger-

Schnack, 1990a). Diurnal vertical migration of moderate intensity

was observed during this season, with specimens showing a

stronger tendency to concentrate within the depth range of maxi-

mum temperature gradients during the night than during the day.

From autumn to winter, a conspicuous shift of the population

centre from the lower epipelagic zone (40— 100 m) to shallower

depths (0-50 m) was observed, which coincided with the weaken-

ing of the thermocline during the winter season (Bottger-Schnack,

1990b). Males of O. venusta f. venella generally exhibited the

same depth distribution as females, but were evaluated semi-quan-

titatively during the autumn survey only.

SEASONAL VARIATION IN ABUNDANCE (CENTRAL RED SEA)

Seasonal variation in abundance in the central Red Sea was most

conspicuous for O. venusta f. typica, which occurred in the central

area only during winter, but was absent during summer and autumn

(B6ttger-Schnack, 1990b, 1995). By this, a strong inflow of south-

ern Red Sea populations into the central area was indicated, and the

species has been regarded as an indicator species of southern Red

Sea waters, similar to other copepod species, such as species of

Eucalanus (Beckmann, 1984, 1996). Abundances of O. venusta f.

venella in the central Red Sea were highest during winter, but

moderately high densities were also found during autumn, thereby

indicating a lesser influence of the southern inflow than was observed

for f. typica.

Oncaea mediterranea (Claus, 1863)

Antaria mediterranea Claus (1863): 159-160, Tafel XXX, Fig. 1-6

(D273):

Oncea mediterranea (Claus, 1863); Oncda mediterranea (Claus,

1863).

OTHER DESCRIPTIONS. Giesbrecht (1892) [as Oncda mediter-

ranea); Heron (1977); Heron & Bradford-Grieve (1995): for further

details see Bottger-Schnack & Huys (1997b).

TYPE LOCALITY. Tyrrhenian Sea; Messina.

PRELIMINARY NOTE. A detailed redescription of O. mediterranea,

based on material from the Red Sea and the Eastern Mediterranean,

has recently been published by Bottger-Schnack & Huys (1997b),

including a review of the taxonomic history of the species. The

following corrective note describes additional morphological details,

which were not mentioned in the previous account, but might be of

importance for constructing phylogenetic relationships within the

genus Oncaea s. str.

CORRECTIVE NOTE

(1) The labrum of O. mediterranea exhibits an additional patch of

long spinules on posterior face at posterior part of each lobe,

similar to O. venusta (cf. Fig. 3C), which was not figured by

Bottger-Schnack & Huys.

(2) The endopodal spines of Pl exhibit a subapical tubular exten-

sion, similar to O. venusta (cf. Fig. 4A). These extensions seem

to be widespread among oncaeids as they have been found in

distantly related species, such as Archioncaea arabica (Bottger-

Schnack & Huys, 1997a), species of Triconia (BOottger-Schnack,

1999), O. subtilis (Bottger-Schnack & Huys, in press) and

species of the atlantica-group (unpubl. data).

(3) The coxa of P4 is ornamented with a patch of long setules on

posterior face, similar to O. venusta (cf. Fig. 4D).

(4) The female P6 is ornamented with 1 spine and 2 spinous

processes, as in all other species of Oncaea s.str., not only 1

spinous process as erroneously figured by Bottger-Schnack &

Huys.

(5) In addition to the sexually dimorphic characters described by

Bottger-Schnack & Huys, sexual dimorphism is expressed in

the endopods of P1—P3, in the ornamentation of P5, and in the

length of caudal setae: (1) in P1, the spinous outgrowth at the

distal margin of the endopod is relatively longer in the male,

reaching half the length of the distal spine, (2) in P2—P3, the

conical projections on enp-3 are relatively longer in the male

as compared to the female, similar to O. venusta (cf. Fig. 7B,

C, E, F), (3) the outer long seta on P5 exopod is ornamented

with triplicate row of minute spinules along entire length, not

naked as in female, (4) caudal setae VI and IV are relatively

shorter in the male as compared to the female, which was

correctly figured by Bottger-Schnack & Huys (their Fig. 4A),

but erroneously described as being equal in length to the

female in the text.

Oncaea media Giesbrecht, 1891

Oncaea media Giesbrecht (1891): 477.

Oncda media Giesbrecht (1892)

RELIABLE DESCRIPTIONS. Giesbrecht (1892): 591-600, 602, 603,

756, 757, 774, Plate 47, Fig. 1 (not Fig. 11), 29-33, 40 [as Oncda

media]; Tanaka (1960): 69,70, Plate XXXI, Figs. 4-9; Heron &

Bradford-Grieve (1995): 36, 39, Figs. 15k, 16a—k, 17a—-1, 26b; Itoh

{in: Chihara & Murano 1997]: 980, Fig. 365a-f.

DOUBTFUL DESCRIPTIONS. Dakin & Colefax (1940): 117, Fig.

205C a [2 only]; Chen er al. (1974): 41-42, Plate 6, Figs. 12-15;

Mori (1937; reprinted 1964): 120-121, Plate 66, Figs. 14-18.

TYPE LOCALITY. not specified; original description based on

material from various locations near the equator in the tropical

Pacific.

PRELIMINARY NOTE. Géiesbrecht’s original material of O. media

was not available for study, because it is not allowed to be sent out

on loan (A. Ianora, Zoological Station Naples, pers. comm.). Heron

& Bradford-Grieve (1995) gave an excellent redescription of O.

media based on specimens from the Gulf of Naples and provided a

summary of its distribution in the Pacific Ocean and other areas.

They pointed to the great similarity between O. media and O.

scottodicarloi, which they described as a new species, and cleared

up the confusion in Giesbrecht’s (1892) redescription with regard to

these two species. Specimens from the Red Sea agreed in almost

every detail with the redescription of Heron & Bradford-Grieve

(1995). However, some morphological characters are described

below, which were not noted by Heron & Bradford-Grieve and/or

appeared to differ between the two areas. Also, the body dimensions

of the species from the Red Sea calculated by the different methods

used throughout this study are provided.

MATERIAL EXAMINED.

(1) Northern Red Sea, 22° 58.4'N, 37° 19.4'E: Stn. 663; R/V Meteor

leg 5/5: collected 20 July 1987 with MSN 0.05 mm net (Haul 17/

4); depth 50-100 m; total water depth ca 1200 m.

(a) 2 29in alcohol (BMNH 1998.2797-2798).

(b) 1 3 in alcohol (ZMH K-39584).

slides, 2 dd in alcohol (RBS).

(2) Gulf of Aden, 11° 55.5'N, 43° 37.9'E: Stn. 631; R/V Meteor leg

5/5: collected 11 July 1987 with MSN 0.05 mm net (Haul 3/5);

depth 0-50 m; total water depth ca 1400 m.

(a) 2 29,2 dé in alcohol (BMNH 1998.2799-2802).

(b) 2 22,2 3¢ in alcohol (ZMH K-395835).

DESCRIPTION. Note illustrations are based on | (c).

ADULT FEMALE (Fig. 15).

Body length: 884 um [traditional method: 710 um, range: 650-770

uum, based on 22 specimens (Bottger-Schnack ef al., 1989)].

Exoskeleton moderately chitinized. Prosome 2.6 times length of

urosome, excluding caudal rami, 2.2 times urosome length includ-

ing caudal rami. P2-bearing somite without conspicuous

dorso-posterior projection visible in lateral aspect (Fig. 15B).

Integumental pores on prosome as indicated in Fig. 15A, B. Pleural

areas of P4-bearing somite with rounded posterolateral corners.

Proportional lengths (%) of urosomites 13.0: 62.7: 8.5: 6.1 : 9.7.

Proportional lengths (%) of urosomites and caudal rami 11.4 : 55.2

BoD) BD)-8) 85) BAIT

Genital double-somite 1.9 times as long as maximum width

R. BOTTGER-SCHNACK

(measured in dorsal aspect) and 2.6 times as long as postgenital

somites combined (Fig. 15C). Genital apertures with armature

represented by 1 spine and 2 minute spinous processes (Fig. 151)

[the latter 2 not mentioned by Heron & Bradford-Grieve]. Double-

curved sclerotization between genital apertures, pore pattern on

dorsal surface as indicated in Fig. 15C.

Anal somite 2.0 times wider than long; about 2/3 length of caudal

rami (Fig. 15C). Ornamentation as in O. venusta.

Antennule with armature as for genus, small element on 6th

segment (arrowed in Fig. 15 D) not mentioned by Heron & Brad-

ford-Grieve.

Antenna as figured by Heron & Bradford-Grieve (their Fig. 16c),

except for seta I of lateral armature on first endopod segment slightly

longer than figured by these authors; additional surface ornamenta-

tion on coxobasis as in O. waldemari (cf. Fig. 25A).

Labrum (Fig. ISE, F) as for O. venusta, but lacking patch of

setules on posterior face at posterior part of each lobe. [O. media is

the only species of Oncaea s.str. that misses these setules.]

Mandible mainly as figured by Heron & Bradford-Grieve (their

Fig. 16e), except for dorsal blade (C) ornamented with 4 dentiform

processes at distal margin, 1 of them inserted subdistally, and 1

minute process halfway at dorsal margin (Fig. 16E). Maxillule,

maxilla and maxilliped mainly as figured by Heron & Bradford-

Grieve (their Figs. 16f—h), but with additional surface ornamentations

and microstructures, such as 2 large secretory pores (not 1) on

surface of maxilla and third single row of shorter spinules along

outer margin of strong maxillary spine (similar to O. waldemari, cf.

Fig. 25F). [Complete pattern of microstructures on surface of ex-

oskeleton not additionally figured in present account, but generally

similar to those observed for O. venusta and O. waldemari].

Swimming legs with armature as for genus and surface ornamenta-

tion similar to O. waldemari (cf. Figs. 26A—D). P1—P4 with spines on

exp-1 longer than figured by Heron & Bradford-Grieve (their Figs.

16i-k, 17a), reaching beyond half length of spine on exp-2. P1 with

inner basal element minutely pinnate (Fig. 15G), not naked as figured

by Heron & Bradford-Grieve; distal margin of exp- 1 and-2 ornamented

with long spinules anteriorly as in O. scottodicarloi (cf. Fig. 23C).

P5 (Fig. 15H) with exopod longer than wide, length to width ratio

eVaealle

P6 (Fig. 151) armed with a spine and 2 small spinous processes

[the latter 2 not mentioned by Heron & Bradford-Grieve].

Egg-sacs not observed.

ADULT MALE (Fig. 16).

Body length: 672 um [traditional method: 560 um, | specimen].

Sexual dimorphism in antennule, antennary setae, maxilliped, P1—

P3 (endopod) and P5—P6, caudal ramus and in genital segmentation.

Prosome 2.2 times the length of urosome, excluding caudal rami,

1.9 times urosome length, including caudal rami.

Proportional lengths (%) of urosomites (excluding caudal rami)

12.3 : 66.4 : 4.3 : 3.7 : 4.3 : 8.6; proportional lengths (%) of

urosomites (caudal rami included) 10.9 : 58.7 : 3.8: 3.3:3.8:7.6:

11.9. Caudal rami about as long as wide, much shorter than in

female. Caudal setae with proportional lengths as in female, except

for seta VI, which is about 2/3 the length of seta IV and 3 times the

length of caudal ramus (Fig. 16A).

Antennule with armature as for genus.

Antenna (Fig. 16D) as in female, except for seta on coxobasis

naked and shorter than in female [not mentioned by Heron &

Bradford-Grieve]; lateral armature on distal endopod segment dif-

fering from female, with spiniform seta III much stouter and seta IV

spiniform and curved, with row of dentiform processes along outer

distal margin; both elements shorter than in female.

=. =

SEVEN SPECIES OF RED SEA ONCAEA

LAID

LEE: sea

LEE

——

Fig. 15 Oncaea media, female (Red Sea) (A) Habitus, dorsal; (B) same, lateral (appendages omitted); (C) urosome, dorsal; (D) antennule, 6th segment,

small sensory element arrowed; (E) labrum, anterior: (F) same, posterior; (G) inner basal seta of P1; (H) P5, dorsal; (I) P6.

SSS

A7 ay,

: L

SSS ¢

> a :

=. v

Fig. 16 Oncaea media, male (Red Sea) (A) Habitus, dorsal; (B) P5,

dorsal; (C) P6, aberrant posterolateral corner arrowed. (D) antenna,

anterior; (E) mandible, blade C.

R. BOTTGER-SCHNACK

Maxilliped as figured by Heron & Bradford-Grieve (their Fig.

17f).

Swimming legs 1-4 with armature and ornamentation as in

female; sexual dimorphism expressed in (1) terminal process on P1!

endopod (obscuring insertion of distalmost seta), being relatively

longer than in female (similar to O. scottodicarloi, Fig. 231), (2)

terminal conical projections on distal endopod segment of P3 (not on

P2) longer than in female, reaching 2/3—3/4 the length of outer distal

spine and (3) reduced spine lengths in endopod spines of P2—P4

(Table 4).

Table 4 Sexual dimorphism in spine length (um) on distal endopod seg-

ment of P2—P4 in O. media from the Red Sea. (Data represent single

measurements).

Spine length

Female Male Male : female (%)

Outer subdistal spine 20.0 10.0 50

Outer distal spine 18.1 Tes 41

Distal spine 26.3 11.3 43

P3 left/right

Outer subdistal spine 18.8 10.6/11.3 57/60

Outer distal spine 19.4 10.6/13.8 Soy

Distal spine 32.5 25.6/25.6 79

Outer subdistal spine 24.3 16.3 67

Outer distal spine 26.3 20.6 78

Distal spine 43.8 SiS 72.

P5 (Fig. 16B) exopod fused to somite, length of segment shorter

than in female; proportional lengths of exopodal setae as in female.

P6 represented by posterolateral flap closing off genital aperture

on either side; covered by pattern of denticles as in Fig. 16C;

posterolateral corners with single pointed tip, occasionally with

bifid tip (arrowed in Fig. 16C).

TAXONOMY

Giesbrecht (1891: 477) presented a short latin diagnosis of O. media

and subsequently described the species in more detail from the

Mediterranean, Naples area (Giesbrecht, 1892). Recently, Heron &

Bradford-Grieve (1995) redescribed O. media from the Gulf of

Naples and in the same account described a closely related species,

O. scottodicarloi, which co-occurred with O. media in the Gulf. The

authors pointed out an important discrepancy in Giesbrecht’s (1892)

description of O. media: all of his figures of the species matched O.

media as re-examined by Heron & Bradford-Grieve, except for his

Plate 47, Fig. 11, dorsal view of female urosome, which resembled

that of the newly described O. scottodicarloi. [In this context it

should be noted that Heron & Bradford-Grieve (1995) recorded

several undescribed species similar to O. media and O. scottodicarloi

in their Gulf of Naples samples, indicating that a complex of species

close to O. media exists, which is as yet undescribed. One of the

species belonging to this complex is undoubtedly O. waldemari,

whose taxonomic position will be re-evaluated below. ]

Specimens of O. media from the Red Sea met all important

characters described by Heron & Bradford-Grieve, including pat-

terns of integumental pores on the prosome of both sexes. Slight

differences in Red Sea specimens included (1) the armature on 6th

segment of antennule, the minute sensory element not noted by

Heron & Bradford-Grieve, (2) additional surface ornamentation on

coxobasis of female antenna and (3) spine lengths on P2—P4 enp-3 in

the male. The first two characters are difficult to discern and may

have been overlooked in the previous description. The reduction of

spine lengths on P2—P4 enp-3 in male O. media was less pronounced

SEVEN SPECIES OF RED SEA ONCAEA

in specimens from the Red Sea than had been reported by Giesbrecht

(1892, Pl. 47, Figs. 32, 33) and Heron & Bradford-Grieve (their Fig.

17g—). In particular the outer distal spine on P2 enp-3 was longer in

Red Sea specimens, measuring about 3/4 the length of the outer

subdistal spine (cf. Table 4), whereas it was figured being only 1/3—

1/2 that length by the authors mentioned above. Proportional spine

lengths on P2 enp-3 of O. media males from Japanese waters as

recorded by Itoh (in: Chihara & Murano, 1997: his Fig. 365f) were

similar to those found in Red Sea specimens.

O. media is closely related to O. scottodicarloi, with which it has

often been confused. According to Heron & Bradford-Grieve (1995),

important distinctions between females of the two species are found

in (1) proportional lengths of urosome segments, (2) form and

location of sclerotization between genital apertures, and (3) relative

lengths of endopod spines of P4. Comments on the first two characters

will be given below. A further character separating females of the

two species as observed during the present study is the proportional

length of exopodal setae on PS, the outer one being relatively shorter

in O. media than in O. scottodicarloi.

(1) The typical elongate form of the genital double-somite of O.

media as figured by Heron & Bradford-Grieve appears to be of

great significance, because it enables unequivocal separation of

O. media from other species of the media complex. In the original

account of Giesbrecht (1892), however, the urosome is concealed

underneath the paired egg-sacs of the ovigerous specimen in the

illustration of female habitus (Plate 47, Fig. 1) and his illustration

of the female urosome in dorsal aspect (Fig. 11) was assigned to O.

scottodicarloi by Heron & Bradford-Grieve. Therefore it cannot

be decided definitely, whether this character had also been present

in Giesbrecht’s specimen and the conspecificity of Giesbrecht’s

O. media with O. media sensu Heron & Bradford-Grieve can only

be confirmed after re-examination of his type material. The lateral

view of O. mediaas figured by Giesbrecht (Plate 2, Fig. 12) shows

the genital segment being twice as long as the remaining part of the

urosome, which is similar to O. media sensu Heron & Bradford-

Grieve; also, the proportional lengths of spines on P4 enp-3 P4

figured by Giesbrecht are more similar to O. media than to O.

scottodicarloi. Since both characters support Heron & Bradford-

Grieve’s conclusion, their opinion is followed in the present

account.

(2) The integumental sclerotization located between the genital

apertures is a reinforced attachment site for the insertion of the

dorsal longitudinal trunk muscles (R.Huys, pers. commn.). These

usually insert on the anterior rim of each somite, or in the case of

the female genital double-somite near the genital apertures,

marking the original plane of fusion. The sclerotization is

always located at about 1/3 to 40% the distance from the anterior

margin and does not differ between species, but as the position

of genital apertures on the genital double-somite is different, the

sclerotization changes its position relative to the genital aper-

tures. In O. media, the genital apertures are located more laterally

than in O. scottodicarloi, and thus the sclerotization changes its

position accordingly.

Males of O. media can be separated from those of O. scottodicarloi

most easily by the sexual dimorphism in the coxobasal seta of the

antenna, which is short and naked (Fig. 16D) in O. media, but long

and plumose as in the female in O. scottodicarloi. Giesbrecht and

Heron & Bradford-Grieve did not notice the sexual dimorphism of

this seta in O. media. In the present study, however, this character

was found in all species of Oncaea s.str., except for O. scottocarloi

and O. waldemari. The sexual dimorphism in spine lengths of P2—P4

enp-3, on the other hand, which had already been reported for males

of O. media in both previous account mentioned above, cannot be

used to distinguish between males of both species, since it was also

found in O. scottodicarloi during the present study (Fig. 23J, M) and

seems to be a typical character for males of Oncaea s.str. (cf. O.

venusta, O. waldemari).

OTHER RECORDS OF O. MEDIA

A summary of the world-wide records of O. media has been com-

piled by Heron & Bradford-Grieve (1995). They point to the great

confusion caused by Sewell (1947), who described two size vari-

ants, f. major and f. minor, from the Arabian Sea. The major form (

2: 0.73-0.82 mm, dd: 0.53—0-62 mm) exhibited a smaller length to

width ratio of caudal ramus (2:1) than the minor form (3:1), which

would correspond to the observed differences between O. media and

O. scottodicarloi. However, Sewell described the distal spine on P4

enp-3 of f. major measuring about 2/3 the length of distal endopod

segment (p.261), which is longer than reported by Giesbrecht (1892)

and Heron & Bradford-Grieve for O. media, the spine being only

Table 5 List of important characters separating O. media and related species. Data of O. curta after Sars (1918), remaining data from present study.

Species media scottodicarloi curta waldemari

Female

GDS

— |:w ratio 1E9=1 oe 15:1 We 7/e

— Form very el. oval-el. squarish elongate

—ratio GDS : rest of urosomites! (excl.CR) 3.9:1 Se Sil 1.9:1

— position gen.ap., distance from lateral margin 1/4 1/3 ca 1/5 1/4

L. ratio caudal setae

—seta VI:IV 2/3 1/2 ca 1/2 3/4

—seta VI:CR 3.8:1 2.221 ca 1:1 1.6:1

—seta VIL:II slightly longer slightly longer ? longer

Labrum, anterior face, paired patch of setules no yes 2 yes

PS exopod

— |:w ratio el ley cal.5:1 38

— outer:inner seta equal slightly longer 1/2length equal

Male

Antenna

— coxobasal seta short naked long, plumose 2 long, plumose

—endopod 2, element IV strong-curved curved 2 slightly curved

]. ratio caudal seta VI: VII longer longer u equal

GDS = genital double-somite; CR = caudal ramus; gen.ap.= genital apertures; el. = elongate; | = length; w = width

‘Calculated by traditional method (i.e. telescoping of segments not considered)

1/2 length of the segment. Confusingly, Sewell refers to his figures

of swimming legs, but these are not given in his account. A relatively

long distal spine on P4 enp-3 is found in O. scottodicarloi, and it

might be that Sewell had mixed up the characters of both species in

his report. Since all three species of the O. media-complex (O.

media, O. scottodicarloi and O. waldemari are found in the Arabian

Sea (Bottger-Schnack, 1996, as O. media f. major, O. media f. minor

and Oncaea sp. B, respectively), it is conceivable that he included

the latter species in the lower size range of females as well.

Mori (1937, reprinted 1964) described the species from Japanese

waters and Chen et al. (1974) recorded two size groups of O. media

from the East China Sea and the Yellow Sea. In both accounts, the

female genital double-somite is much shorter than in O. media sensu

Heron & Bradford-Grieve and also seta VI on caudal ramus is too

short. The same argument applies for the record of Dakin & Colefax

(1940) from Australian waters. Although the illustration of the male

antenna by Mori (his Plate 66, Fig. 17) shows a naked coxobasal

seta, positive identification cannot be made without a closer exam-

ination of the specimens, especially in view of the extremely wide

size range that was recorded for females (0.5—0.92 mm) in his

account. However, the occurrence of O. media in the marine plank-

ton of Japan has been confirmed in a recent excellent account by Itoh

[in Chihara & Murano, 1997].

Tanaka (1960) recorded O. media from various locations in the East

China Sea, Indian Ocean and south of Cape of Good Hope. He

mentions the occurrence of two size groups, f. major and f. minor,

which covered an overall size range of 0.55—0.79 mm. Sizes of the two

groups were not recorded separately. His Plate XX XI, Fig. 4 clearly

shows the dorsal view of female O. media sensu Heron & Bradford-

Grieve, although he identified them as f. ‘minor’. The different length

ratio between subdistal and distal spines on P4 enp-3, which accord-

ing to Heron & Bradford-Grieve (1995) separate O. media from O.

scottodicarloi, is not apparent between Tanaka’s f. major and f. minor

(his Figs. 5 and 6). So it cannot be judged whether his small sized

specimens are identical to O. scottodicarloi or probably belong to

another closely related species, such as O. waldemari.

O. media f. minor sensu Malt (1982b) was assigned to O.

waldemari during the present study and will be discussed below. In

the identification key of oncaeids (Malt, 1983b) the separation

between males of O. media and O. venusta needs to be revised, since

the sexual dimorphism on antenna is found in both species, not only

in O. venusta, as was erroneously stated by Malt.

GEOGRAPHICAL DISTRIBUTION

Oncaea media had been reported as O. media f. major in the

previous quantitative accounts of Béttger-Schnack (1990b, 1994,

1995, 1996, 1997). The species is distributed throughout the Red

Sea, but exhibits very variable abundances both seasonally and

regionally. During summer, the species occurred in minimal num-

bers in the southern Red Sea, but exhibited higher values to the

north, in the central-northern Red Sea, as well as to the south, in the

Gulf of Aden and Bab al Mandab area (BO6ttger-Schnack, 1995).

In the northern Arabian Sea, O. media was found in appreciable

numbers, with abundances being one to two orders of magnitude

higher than in the Gulf of Aden and/or in the central Red Sea

(Bottger-Schnack, 1996). The species also occurred in the Eastern

Mediterranean Sea (Bottger-Schnack, 1997) in comparably low

numbers.

SEASONAL VARIATION IN ABUNDANCE (CENTRAL RED SEA)

O. media was recorded from the central-northern Red Sea during

winter (Bottger-Schnack, 1990b) and summer (B6ttger-Schnack,

1995), but not during autumn, when it was completely absent or

occurred as solitary finds only (Bottger-Schnack, 1990a, b). Due to

R. BOTTGER-SCHNACK

its seasonality, the species had previously been assumed to be of

southern origin (Bottger-Schnack, 1990b), but subsequent data from

the summer season did not point to a consistent seasonal variation in

the central Red Sea (B6ttger-Schnack, 1995).

VERTICAL DISTRIBUTION AND VERTICAL MIGRATION

O. media was generally confined to the epipelagic zone (0-150 m)

in the Red Sea, with few, isolated finds down to 900 m. The species

had a unimodal distribution pattern with maximum densities in the

upper epipelagic (0-50 m) and was classified as non-migratory

during winter in the central Red Sea (Bottger-Schnack, 1990b). In

the Gulf of Aden and Strait of Bab al Mandab, maximum densities

were found at a depth of 0-20 m and 0-60 m, respectively, during

summer (unpubl. data). By this, the species tended to be vertically

separated from the two related species, O. scottodicarloi and O.

waldemari, which generally occurred deeper and over a much wider

depth range in that area.

Oncaea clevei Friichtl, 1923

Oncea clevei Friichtl(1923): 455, Tafel 26, Figs. 19-22 (2 only).

Oncaea conifera Cleve, 1901

RELIABLE DESCRIPTIONS. Friichtl (1924): 22-23, 89-91, Figs.

14,15, 60-70 (2 only); Itoh [in: Chihara & Murano (1997)]: 979,

Fig. 361a, d, f (Qonly).

DOUBTFUL DESCRIPTIONS. Sewell (1947): 258 [as Oncea clevei);

Tanaka (1960): 66, Plate XX VIII, Figs. 7-13; Chen et al. (1974): 42,

Plate 7, Figs. 4-7.

TYPELOCALITY. Aru Archipelago, Indo-Pacific area.

PRELIMINARY NOTE. The original description of Friichtl (1923,

1924) lacks many details, particularly in the mouthparts, of which

he described only the maxilliped. A detailed redescription of O.

clevei from the Red Sea is given below, including a description of

the hitherto unknown males. During the course of the study a

closely related species was found, which is described as a new

species, O. paraclevei sp. nov. A comparison of morphological

characters separating the two species is included under O.

paraclevei see Remarks.

MATERIAL EXAMINED.

(1) Central Red Sea, 21° 25.53'N, 38° 01.91'E: Stn. 130; R/V

Valdivia leg 29: collected 28 October 1980 with MSN 0.1 mm

net (Haul 117/5); depth 0-20 m; total water depth 1960 m.

(a) 2 22in alcohol (BMNH 1998.2803—2804).

(b) 2 22 in alcohol (ZMH K-39574).

in alcohol (RBS).

(2) Southern Red Sea, 15° 34.8'N, 41° 54.9'E: Stn. 703; R/V Meteor

leg 5/5: collected 03 August 1987 with MSN 0.05 mm net (Haul

39/5); depth 0-50 m; total water depth 970 m.

(a) 1 6 partly dissected (urosome on slide), remaining speci-

men in alcohol (BMNH 1998.2805), 3 dd in alcohol (BMNH

1998.2806—2808).

(b) 1 6 dissected on 10 slides (ZMH K-39575a-i), 1 2.3 dd in

alcohol (ZMH K-39576).

(3) Gulf of Aden, 11° 55.5'N, 43° 37.9'E: Stn. 631a; R/V Meteor leg

5/5: collected 11 July 1987 with MSN 0.05 mm net (Haul 3/5);

depth 0-—50m; total water depth 1400 m.

(a) 1 2 1 do (mating position, d lacking urosome) in alcohol

(BMNH 1998.2809-2810).

(b) 1 9, 1 d (mating position) in alcohol (ZMH K-39577).

== qi em,. «

————

SEVEN SPECIES OF RED SEA ONCAEA

from mating pair) in alcohol, 1 9, 1 d (mating position) in

alcohol (RBS).

(4) Northern Arabian Sea, 19° N, 65° E: Stn. 247; R/V Meteor leg

32/3: collected 14 May 1995 with MSN 0.05 mm net (coll. L.

Postel); depth 0-50m; total water depth ca 3000 m: 1 2dissected

on 11 slides, 1 Qin alcohol (RBS).

(5) Northern Arabian Sea, near Oman, 20° 44.3'N, 59° 40.5'E: Stn.

347; R/V Meteor leg 5/3a: collected 05 April 1987 with MSN

0.05 mm net (Haul 8/1); depth O—S0 m; total water depth ca 2500

m: 6 29 in alcohol (RBS).

(6) Pacific Ocean, Great Barrier Reef; further sampling data not

specified; leg G.P. Farran, 1928-1929, cf. Farran (1936); 10

specimens in alcohol, labelled Oncaea clevei (BMNH

1949.12.31.516): this vial contains 5 29 of O. clevei, 1 prosome

with dorsal hump and 4 damaged prosomes; 2 urosomes (1 of O.

clevei, 1 possibly of O. paraclevei sp. nov.), and 2 bits of

calanoid copepods).

(7) Pacific Ocean, Great Barrier Reef; further sampling data not

specified; leg G.P. Farran, 1928-1929, cf. Farran (1936); more

than 10 specimens in alcohol, labelled Oncaea clevei (BMNH

1948.4.28.140): this vial contains 22 22 of O. clevei, 1 3

Corycaeidae indet., | calanoid copepod.

DESCRIPTION. Note illustrations are based on (2a—2c) and (4).

ADULT FEMALE (Figs. 17-19, 21A).

Body length: 810 um [traditional method: 640 um, range: 620-680

tum, based on 4 specimens].

Exoskeleton well chitinized. Prosome 2.4 times length of urosome,

excluding caudal rami, 2.1 times urosome length including caudal

rami. P2-bearing somite with conspicuous dorso-posterior projec-

tion in lateral aspect (Fig. 17B), not varying in size between specimens

examined. Integumental pores on prosome as indicated in Fig. 17A,

B. Pleural areas of P4-bearing somite with rounded posterolateral

corners.

Proportional lengths (%) of urosomites 11.4: 55.3: 10.1: 10.1:

13.1. Proportional lengths (%) of urosomites and caudal rami 9.6 :

BOs s.). 8.) 2 11:0): 16:0.

Genital double-somite almost rectangular in dorsal aspect, 1.4

times as long as maximum width (measured in dorsal aspect) and 1.6

times as long as postgenital somites combined (Fig. 17C); largest

width measured near anterior margin, lateral margins of genital

double-somite rounded at anterior quarter, posterior part tapering

slightly. Paired genital apertures located very close to dorsolateral

margin at about 2/5 the distance from anterior margin of genital

double-somite; armature represented by | spine and 2 minute spinous

processes (Fig. 17H). Sickle-shaped sclerotization between, but

slightly posterior to genital apertures, pore pattern on dorsal surface

as indicated in Fig. 17C.

Anal somite 1.4 times wider than long; slightly shorter than

caudal rami (Fig. 17C). Surface ornamentation mainly as in O.

venusta.

Caudal ramus (Fig. 17F) about 2.3 times as long as wide. Seta VI

more than 4/5 length of seta IV and 2.5 times longer than caudal

ramus; seta VII about half length of seta VI and longer than seta III

(Fig. 17C, F). Inner margin of somite with few long setules.

Antennule (Fig. 17E) with relative lengths (%) of segments

measured along posterior non-setiferous margin 6.7 : 26.1 : 38.3:

11.1: 4.4: 13.3. Armature as for genus.

Antenna (Fig. 18A) similar to O. venusta, except for coxobasis

with fewer surface ornamentation and seta III on second endopod

segment bipinnate.

Labrum (Fig. 18B,C) as in O. venusta, except for long fine setules

missing latero-distally on outer margins of lobes. Anterior surface

(Fig. 18B) with row of stout denticles on proximal part of each lobe,

which are not found in O. venusta.

Mandible (Fig. 18D) as in O. venusta, except for blade C

ornamented with | additional dentiform process halfway on dorsal

margin. Maxillule (Fig. 18E), maxilla (Fig. 18F), and maxilliped

(Fig. 18G) mainly as in O. venusta.

Swimming legs (Fig. 19A—D) with armature as for genus and

surface ornamentation similar to O. venusta, except for less surface

ornamention on coxae and bases, most obvious in P4 coxa, lacking

patch of long setules on posterior surface. Endopodal and exopodal

spine lengths mainly as in O. venusta, except for terminal spines on

endopods of Pl, P2 and P4 relatively longer, particularly in P4,

reaching more than half the length of distal endopod segment. P2

with outer subdistal spine reaching insertion of outer distal spine; P4

with outer distal spine shorter than in O. venusta, about 1/2 length of

distal spine. Inner basal seta on P1 naked; outer basal seta on P3

plumose.

P5 (Fig. 17G) comprising small naked seta arising from lateral

surface of somite, and free unornamented segment representing

exopod. Exopod twice as long as wide, bearing 2 spiniform, naked

setae equal in length, which are straight.

P6 (Fig. 17H) represented by operculum closing off each genital

aperture; armed with a spine and 2 small spinous processes.

Egg-sacs paired, oval-shaped; each sac containing appr. 15-20

eggs (diameter 40-48 um) (Bottger-Schnack er al., 1989).

ADULT MALE (Fig. 20, 21A-C).

Body length: 630 um [traditional method: 540 um, range: 490-570

uum, based on 4 specimens]. Sexual dimorphism in antennule,

antenna, maxilliped, P2—P3 (endopod) and P5—P6, caudal ramus and

in genital segmentation.

Prosome 2.5 times the length of urosome, excluding caudal rami,

2.2 times urosome length, including caudal rami.

Proportional lengths (%) of urosomites (excluding caudal rami)

12.5 : 66.9 : 4.3 : 4.3 : 4.3 : 7.6; proportional lengths (%) of uro-

somites (caudal rami included) 11.0: 58.9: 3.8: 3.8:3.8: 6.7: 12.0.

Caudal rami about 1.8 times longer than wide, shorter than in

female. Caudal setae with proportional lengths as in female, except

for seta VI, which is about 3/5 the length of seta IV and 2.2 times

length of caudal ramus; seta VII longer than in female, reaching

about 2/3 length of seta VI (Fig. 20A).

Antennule (Fig. 20B) relative lengths (%) of segments measured

along posterior non-setiferous margin 10.0 : 24.1 : 38.2 : 27.6.

Armature formula as for genus.

Antenna (Fig. 20H) as in female, except for seta on coxobasis

naked and shorter than in female; lateral armature on distal endopod

segment differing from female, with element III being much stouter

and element IV spiniform and distinctly curved; both elements

shorter than in female.

Maxilliped (Fig. 20C) similar to O. venusta.

Swimming legs 1-4 with armature and ornamentation as in

female; terminal process on Pl endopod as in female; P4 with outer

distal spine half length of distal spine as in female; sexual dimor-

phism expressed in size of terminal conical projections on distal

endopod segment of P2—P3, being relatively longer than in female,

reaching 2/3 length (P2) or almost same length (P3) of outer distal

spine (Fig. 21B1, B2, C). Variation in spine length as shown in Fig.

ZIBIE B2:

P5 (Fig. 20G) exopod not delimited from somite, general shape

and armature as in female, except for exopodal setae shorter than in

female.

60 R. BOTTGER-SCHNACK

v us

l°

r 200 um i

SALE Z

COLA KKK Wh f/

Fig. 17 Oncaea clevei, female (Red Sea) (A) Habitus, dorsal; (B) same, lateral (appendages omitted); (C) urosome, dorsal; (D) urosome, lateral; (E)

antennule; (F) caudal ramus, dorsal; (G) P5, dorso-lateral; (H) P6.

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 18 Oncaea clevei, female (Red Sea) (A) Antenna; (B) labrum, anterior; (C) same, posterior; (D) mandible, showing individual elements; (E)

maxillule; (F) maxilla; (G) maxilliped.

R. BOTTGER-SCHNACK

50 um

A-D

Fig. 19 Oncaea clevei, female (Red Sea) (A) P1, anterior; (B) P2, anterior; (C) P3, anterior; (D) P4, posterior.

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 20 Oncaea clevei, male (Red Sea) (A) Habitus, dorsal; (B) antennule; (C) maxilliped, medial; (D) urosome, dorsal (spermatophores immature); (E)

urosome, ventral; (F) same, lateral; (G) P5, dorsal; (H) antenna, anterior.

ENS \

Fig. 21 Oncaea clevei (Red Sea) (A) Female and male, mating position,

lateral, appendages of female and swimming legs of male omitted.

Oncaea clevei, male (Red Sea) (B1, B2) P2, distal part of endopod,

showing variation in spine length. (C) P3, distal part of endopod.

R. BOTTGER-SCHNACK

P6 (Fig. 20E) represented by posterolateral flap closing off

genital aperture on either side; covered by pattern of denticles as

shown in Fig. 20E.

Spermatophore oval (Fig. 20E, F), of variable size according to

state of maturity.

TAXONOMY

Friichtl(1923) gave a first short diagnosis of O. clevei based on a

single female collected near the Aru Archipelago, off Wokam, and

subsequently described it in more detail (Friichtl,1924). The second

account included additional specimens of O. conifera sensu Cleve

(1901) from the Indo-Malayian Archipelago, which had been sent to

Friichtl by G.O. Sars (Friichtl, 1924, footnote to p.89 (111)), and

which he had synonymized with O. clevei after re-examination.

Friichtl& figures of the female urosome (1923: Tafel 26, Fig. 19;

1924: Fig. 24) show an extremely lateral position of the genital

apertures and a sickle-shaped sclerotization between, but slightly

posterior to, the genital apertures. This combination of characters

enables unequivocal identification of the Red Sea specimens with

Friichtl$ O. clevei. The closely related O. paraclevei sp. nov., which

co-occurs with O. clevei in the Red Sea, differs in the location of

genital apertures and in the form of sclerotization (see below under

‘Remarks’ of O. paraclevei). Red Sea specimens of O. clevei differ

slightly from the original account in the length of the outer distal

spine on P4 enp-3, which is about half the length of the distal spine,

whereas it is shorter in O. clevei sensu Friichtl. The length of the

outer subdistal spine on enp-3 in P2 (shorter in Red Sea specimens)

and P3 (longer in Red Sea specimens) is also slightly different

between the two descriptions.

Friichtl (1924) erroneously recorded the armature of Pl enp-3

with 4 inner setae (his Fig. 66 and p. 91), but in the same account

described the ‘aberrant’ condition of the type specimen from Wokam,

which exhibited 5 inner setae (his Fig. 15). The 5th inner seta on P1

enp-3 has repeatedly been overlooked in previous descriptions of

this and other species of Oncaea s.str. (see above e.g. O. venusta),

because the base of the seta is concealed beneath the long anterior

spinous outgrowth of the segment (cf. Fig. 19A).

Males of O. clevei were not recorded by Friichtlor in any reliable

subsequent account. Their description here is based on specimens

taken from male-female pairs in mating position (Fig. 21A), which

represents the highest probability of encountering a conspecific

male of a given species (but see Heron & Bradford-Grieve, 1995 for

exceptions). O. clevei males are very similar in size and habitus to

those of O. venusta f. venella. However, differences in the ornamen-

tation of the labrum (see below) as well as in proportional lengths of

caudal setae separate the two species.

O. clevei is closely related to O. paraclevei sp. nov., from which

it can be distinguished mainly by the location of genital apertures,

the form of the genital double-somite and by further morphological

characters summarized under Remarks, O. paraclevei. Together, the

two species form a well-defined group within Oncaea s.str., which is

characterized by the presence of a dorso-posterior projection (“hump )

on the P2-bearing somite in the female and by the absence of long

setules on the latero-distal margin of the lobes in the labrum. As the

hump is a sexually dimorphic character, which is not found in males,

males of the two subgroups can only be distinguished by differences

in the ornamentation of the labrum, otherwise they are very similar.

OTHER RECORDS OF O. CLEVEI

Malt (1983a) summarized the published records of O. clevei and

found it restricted to surface waters in low latitudes of the western

Pacific and Indian Ocean. In view of the existence of a hitherto

undescribed species, which is extremely close to O. clevei, it may be

assumed that both O. clevei and O. paraclevei may have been

SEVEN SPECIES OF RED SEA ONCAEA

recorded under the name O. clevei in previous records.

Sewell (1947, p. 258) recorded the species from the northern

Arabian Sea, but it is not clear whether his material included O.

paraclevei as well, because he stated that ‘the dorsal projection on

the 2nd thoracic segment varies considerably in its development’,

which is typical for the latter species.

Tanaka (1960) recorded both sexes of O. clevei from the South

China Sea and off Cape Good Hope. His drawing of the female

urosome (Plate XXVIII, Fig. 7) resembles that of O. clevei rather

than O. paraclevei, but his description of the male is meagre and

could also be assigned to any other oncaeid.

Chen et al. (1974) described the species from the Yellow Sea and

the East China Sea. Their figure of female urosome lacks genital

apertures and/or the sclerotization between them, which is necessary

to separate O. clevei from the closely related O. paraclevei. Thus, a

positive identification cannot be given without examination of their

specimens.

More recently, Itoh [in: Chihara & Murano (1997)] recorded the

species from Japanese waters; his dorsal view of the female shows

the two characters typical for O. clevei.

Several other records of O. clevei from different localities in the

Indo-Pacific are known [see Malt (1983a) for a review], but are not

considered here because they did not include figures or a description

that positively identified the species. In the Eastern Mediterranean

Sea, neither O. clevei nor related species were found (B6ttger-

Schnack, 1997), thus corroborating the limitation of this subgroup to

the Indo-Pacific area. The fact that O. clevei was not recorded in the

detailed account of the oncaeid fauna from the New Zealand area by

Heron & Bradford-Grieve (1995), confirms its restriction to low

latitudes (Malt, 1983a).

GEOGRAPHICAL DISTRIBUTION

Oncaea clevei is distributed throughout the Red Sea, with lowest

abundances in the northern area (B6ttger-Schnack, 1990a, b,

1995). It was not found in small mesh net samples from the

northernmost part of the Red Sea, in the Gulf of Aqaba (unpubl.

data). In the southernmost Red Sea and at Bab al Mandab, abun-

dances of the species were up to two orders of magnitude higher

than in the central part (Bottger-Schnack, 1995). The data from

both areas include an unknown number of O. paraclevei sp.nov.,

however, which was not separated from O. clevei during the quan-

titative counts. Thus, the actual regional difference in abundance

remains uncertain.

In the northern Arabian Sea, O. clevei was recorded from the

epipelagic zone by Bottger-Schnack (1996), however, the potential

co-occurrence of O. paraclevei was not investigated.

VERTICAL DISTRIBUTION AND VERTICAL MIGRATION

The depth distribution of O. clevei remains uncertain, as the species

was counted together with O. paraclevei during the earlier quantita-

tive investigations in the Red Sea (Bottger-Schnack, 1988, 1990a, b,

1995). Generally, the two species were confined to the epipelagic

zone (0-100 m) with occasional occurrences below that depth.

Individual specimens found down to 950 m depth during summer

(unpubl. data) might be regarded as contaminants from shallower

depths or as moribund specimens. Within the epipelagic zone, O.

clevei and O. paraclevei exhibited a unimodal distribution in the

upper epipelagic zone (0—20 or 0-40 m), staying above the strong

seasonal thermocline, which usually develops during autumn

(Boéttger-Schnack, 1990a). No indication of a significant diurnal

vertical movement became apparent and also no seasonal variation

in the depth distribution of the two species was noted in the central

Red Sea (Bottger-Schnack, 1990b).

In the deep southern Red Sea and in the Strait of Bab al

Mandab/Gulf of Aden area, the depth distribution of the two

species was similar to that observed in the central area, with

maximum abundances in the upper 20 to 40 m layer (unpubl.

data). In the shallow southern Red Sea, however, two population

centres were found, situated at depths of 0-20 m and 100-125 m.

Re-investigations of the southern plankton samples are required to

find out whether O. clevei might be vertically separated from O.

paraclevei in that area. For other oncaeid species or forms, which

are closely related to each other, such as O. media and O. scotto-

dicarloi and the two forms of O. venusta, a corresponding vertical

separation had been observed in the shallow parts of the southern

Red Sea.

SEASONAL VARIATION IN ABUNDANCE (CENTRAL RED SEA)

O. clevei / O. paraclevei exhibited a strong seasonal variation in

abundance in the central Red Sea, with highest abundances during

winter, but low numbers during summer and autumn (Bottger-

Schnack, 1995). This indicates a substantial recruitment for

populations of the clevei-subgroup in the central area due to the

inflow of southern Red Sea waters during the NE monsoon, which

is similar to that observed for O. venusta f. typica. It remains

uncertain, however, whether this effect applies likewise for both

species of the clevei-subgroup, as the two species were not sepa-

rated earlier.

Oncaea scottodicarloi Heron & Bradford-Grieve, 1995

Oncaea scottodicarloi Heron & Bradford-Grieve (1995): 39-41,

Figs. 17j-r, 18a—k, 27a.

Oncda media Giesbrecht, 1892 (partim); Giesbrecht 1892, Pl. 47,

Fig. 11 only; H.Itoh [in: Chihara & Murano (1997)]: 981, Fig.

369a-f.

TYPE LOCALITY. Gulf of Naples, western Mediterranean Sea.

MATERIAL EXAMINED

(1) Southern Red Sea, 13° 40.0'N, 42° 37.4'E: Stn. 708; R/V Meteor

leg 5/5: collected 05 August 1987 with MSN 0.05 mm net (Haul

47/3); depth 20-40 m; total water depth ca 190 m.

(a) 2 29,1 d in alcohol (BMNH 1998.2811—2813).

(b) 2 22,2 32 in alcohol (ZMH K-39581).

slides, 1 ¢ in alcohol (RBS).

(2) Southern Red Sea, 13° 40.0'N, 42° 37.4'E: Stn. 708; R/V Meteor

leg 5/5: collected 05 August 1987 with MSN 0.05 mm net (Haul

47/2); depth 40-60 m; total water depth ca 190 m.

(a) 2 dd in alcohol (BMNH 1998.2814—2815).

(b) 1 9, 1 3 (mating position) in alcohol (ZMH K-39582).

(3) Central-northern Red Sea, 22° 58.4'N, 37° 19.4'E: Stn. 663; R/V

Meteor leg 5/5: collected 20 July 1987 with MSN 0.05 mm net

(Haul 17/2); depth 150-200 m; total water depth 1200 m: 1 on

slide in lactophenol, numerous 22 and d for length measure-

ments (RBS).

North-East Atlantic, upwelling area off Northwest-Africa, 17°

36'N, 16° 26'W: Stn. 262 (DIV); R/V Meteor leg 64: collected

March 1983 with Messhai [= multiple opening-closing modified

Gulf III type sampler, Pommeranz et al. (1979), Pommeranz &

Moser (1987)] mesh size 0.05 mm; depth 80 m; total water depth

100 m: 1 dissected on | slide in polyvinyl-lactophenol (RBS).

(5) North-East Atlantic, upwelling area off Northwest-Africa, 16°

09,N, 16° 48'W: Stn. 284 (DV); R/V Meteor leg 64: collected 27

March 1983 with Messhai [cf. (4)], mesh size 0.05 mm; depth

10-20-40 m; total water depth 100 m: 1 2dissected on 1 slide in

polyvinyl-lactophenol (RBS).

(6) Eastern Indian Ocean, NW Cape Australia, 21° 49.86'S, 114°

30.3,E: Stn. B; NWC 005/2; RV ‘Lady Basten’ leg 1630;

collected 26 October 1997 with 0.5 m WP-2 net with 0.073 mm

mesh, vertical haul; depth 0-20 m (leg. D. McKinnon): 4 29

(RBS).

DESCRIPTION. Note illustrations are based on 1(c).

ADULT FEMALE (Figs. 22, 23A—E).

Body length (measured in lateral aspect; from anterior margin of

rostral area to posterior margin of caudal rami, calculated as sum of

individual somites): 803 um [traditional method: (a) 600 um, range:

540-610 um, based on numerous specimens from southern Red Sea

and Gulf of Aden; (b) 510 um, range: 480-520 um, based on 12

specimens from central Red Sea (Bottger-Schnack ef al. (1989)].

Exoskeleton moderately chitinized. Prosome 2.8 times length of

urosome, excluding caudal rami, 2.4 times urosome length includ-

ing caudal rami. Integumental pores on prosome as indicated in Fig.

22A, B.

Proportional lengths (%) of urosomites 10.6 : 63.8 : 7.7: 7.2:

10.6. Proportional lengths (%) of urosomites and caudal rami 9.2 :

55.3) 8 O51 3 O.3 3973 IS4'.

Genital double-somite 1.5 times as long as maximum width

(measured in dorsal aspect) and 2.5 times as long as postgenital

somites combined (Fig. 22C); straight sclerotization between geni-

tal apertures, pore pattern on dorsal surface as indicated in Fig. 22C.

Armature of genital apertures represented by | spine and 2 minute

spinous processes (Fig. 22G).

Anal somite 1.8 times wider than long; about 3/4 length of caudal

rami (Fig. 22C). Ornamentation as for O. venusta.

Caudal ramus (Fig. 22F) about 2.3 times as long as wide, shorter

than reported by Heron & Bradford-Grieve. Dorsal seta (VII) half the

length of terminal accessory seta (VI), not 2/3 the length as reported

by Heron & Bradford-Grieve, plumose and bi-articulate at base.

Antennule with minute element on 6th segment (arrowed in Fig.

22E), which was not mentioned in the original account.

Antenna as figured by Heron & Bradford-Grieve (their Fig. 17m),

except for additional surface ornamentation on coxobasis similar to

O. waldemari (cf. Fig. 25A) and 2 patches of spinules (not only 1) on

anterior surface of distal endopod segment.

Labrum (Fig. 23A, B) similar to O. waldemari, but fewer denti-

form processes medially on each lobe and free margin of integumental

pockets distinctly serrate (Fig. 23A).

Mandible generally as figured by Heron & Bradford-Grieve,

dorsal blade ornamented with | additional minute dentiform process

at dorsal margin (Fig. 22H). Maxillule, maxilla and maxilliped

similar to O. waldemari, some ornamentation elements on syncoxa

of maxilla and on basis of maxilliped missing in the account of

Heron & Bradford-Grieve.

Swimming legs 1-4 with armature as for genus and surface

ornamentation similar to O. waldemari (Fig. 26A—D), except for

anterior face of Pl enp-1 and -2 with stronger spinules on distal

margin (Fig. 23C). Pl exp-3 with outer distal spine slightly longer

than figured by Heron & Bradford-Grieve. P4 exp-3 and enp-3 (Fig.

23E) with outer distal spines somewhat longer than in original

account, outer endopodal spine reaching almost 1/2 length of distal

spine.

P5 (Fig. 22F) with exopodal segment longer than figured by

Heron & Bradford-Grieve, ornamented with 3 spinules ventrally;

inner one of exopodal setae stouter and slightly shorter than outer

one.

P6 (Fig. 22G) represented by operculum closing off each genital

R. BOTTGER-SCHNACK

aperture; armed with a spine and 2 small spinous processes.

Egg-sacs paired, oblong-oval; each sac containing 6 eggs (diameter

40-45 um) (Boéttger-Schnack, 1989, as O. media f. minor).

ADULT MALE (Fig. 23F—M).

Body length: 607 um [(a) 440-480 um, based on several specimens

from southern Red Sea and Gulf of Aden; (b) 410 um, range: 410-

440 um, based on 7 specimens from central Red Sea (B6ttger-

Schnack et al. (1989)]. Sexual dimorphism in antennule, antenna,

maxilliped, endopods of P2—P4, P5—P6, caudal ramus and in genital

segmentation.

Prosome 2.5 times length of urosome, excluding caudal rami, 2.2

times urosome length, including caudal rami.

Proportional lengths (%) of urosomites (excluding caudal rami)

11.2 : 69.2 : 3.8 : 3.8 : 3.8 : 8.3; proportional lengths (%) of

urosomites (caudal rami included) 10.0: 61.1 : 3.3: 3.3:3.3:7.3:

11.6. Caudal rami about 1.6 times longer than wide, shorter than in

female. Caudal setae with proportional lengths as in female, except

for seta VI 2.6 times the length of caudal ramus, and dorsal seta VII

2/3 the length of seta VI.

Antennule with armature as for genus.

Antenna with seta on coxobasis long and plumose as in female;

lateral armature on distal endopod segment differing from female,

element III much stouter and element IV spiniform and curved,

ornamented with dentiform processes along distal margin (arrowed

in Fig. 23H); both elements shorter than in female.

Maxilliped as figured by Heron & Bradford-Grieve, except for

endopodal segment (claw) with concave margin naked, not

ornamented with short spinules pinnate as figured in their Fig. 18}.

Swimming legs 1-4 with armature and ornamentation as in

female; terminal conical projections on P2—P3 enp-3 as in female.

Sexual dimorphism expressed in reduced spine lengths on enp-3,

most obvious in P3 (Fig. 23D, J) and in outer subdistal spine of P4

(Fig. 23E, M). Spine lengths on P2 enp-3 less reduced.

P5 (Fig. 23K) exopod not delimited from somite, shape and

armature as in female, except for exopodal setae more spiniform and

shorter than in female; small plumose seta arising from lateral

surface of somite shorter than in female.

P6 represented by posterolateral flap closing off genital aperture

on either side; covered by pattern of denticles as shown in Fig. 23L.

Spermatophore not observed.

TAXONOMY

O. scottodicarloi is the species previously referred to as O. media f.

minor in the quantitative studies of Bottger-Schnack (1990a, b,

1994, 1995). Specimens from the Red Sea agreed in almost every

detail with the original account of Heron & Bradford-Grieve (1995)

from the Mediterranean, except for their smaller size, which is

discussed below under ‘Size variation’. The only other remarkable

difference in female morphology between the two regions was the

length to width ratio of caudal ramus, which was smaller in Red Sea

specimens (2.3:1) than in the original account (about 3:1). [In the

closely related O. waldemari, the length to width ratio of the caudal

ramus turned out to be a variable character at closer examination

(see below), which might also apply to O. scottodicarloi]. Other

slight differences between the two descriptions, such as the surface

ornamentation on mouthparts and antenna, as well as the minute

element on 6th segment of antennule, which was not mentioned by

Heron & Bradford-Grieve, are not regarded as substantial and/or

might have been overlooked in the original account.

Males of O. scottodicarloi from the Red Sea differed from the

original account in the ornamentation of maxillipedal claw, which

was naked, while it had been described as having short spinules

along the concave margin by Heron & Bradford-Grieve (1995, p. 40,

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 22 Oncaea scottodicarloi, female (Red Sea) (A) Habitus, dorsal; (B) same, lateral (appendages omitted); (C) urosome, dorsal; (D) caudal ramus,

dorsal; (E) antennule, 6th segment (short sensory element arrowed); (F) P5, dorsal: (G) P6; (H) mandible, blade C.

R. BOTTGER-SCHNACK

ZO"

SSS SS

ZZ, as ae os SE

Zo SSS a FORRES

a Z = SSR REELS ==

t =e ~ FE SO = =

‘ a a ae Se ee

SSS

CDEIJM

age

Ue. =

meres

renwal

SS SSS

SS=s SSS

=y SESS

& < SESS

are a

anterior; (D) P3, distal endopod segment,

ramus, dorsal (seta V not drawn); (H) antenna, lateral

armature of second endopod segment (ornamentation of seta IV arrowed); (I) P1, distal part of endopod; (J) P3, distal endopod segment; (K) P5, dorsal;

(L) P6; (M) P4, distal part of endopod.

; (C) Pl, endopod,

Fig. 23. Oncaea scottodicarloi, female (Red Sea) (A) Labrum, anterior; (B) same, posterior

anterior; (E) P4, anterior. — Oncaea scottodicarloi, male (Red Sea) (F) Habitus, dorsal; (G) caudal,

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 18j). Males of oncaeid species usually exhibit a naked

maxillipedal claw, with few exceptions only [Epicalymma schmitti

and Oncaea englishi (Heron, 1977); O. mollicula (unpubl. data)]. In

the case of O. englishi the ornamentation seems to be variable, as it

was lacking in some individuals (Heron, 1977). A further difference

between the original description and the present account is the

sexual dimorphism in spine lengths on enp-3 in P2—P4, which was

not reported for O. scottodicarloi before. The sexual dimorphism of

this character seems to be typical for species of the genus Oncaea

s.str., however, as it was found in almost all species described in the

present account. Nonwithstanding these differences between the

two descriptions, both sexes of the species found in the Red Sea are

regarded as conspecific with O. scottodicarloi.

O. scottodicarloi is closely related to O. media, O. waldemari,

and O. curta. Major distinctions between the four species have been

summarized in Table 5. Female O. media can most easily be

separated from the three other species by the length ratio of urosome

somites, particularly of the genital double-somite, which is much

longer. This character enables rapid identification of the species

during quantitative counts. The three remaining species are morpho-

logically very similar, except for differences in the length ratio of

urosome somites, the length of exopodal seta on PS and the relative

lengths of caudal setae. The morphological relationships of O. curta

within this species complex remain uncertain, however. Sars’ origi-

nal account of O. curta is the only reliable description of the species.

He described several important characters, such as the unequal

exopodal setae on P5, which need to be affirmed, as they are difficult

to discern. The species could not be re-examined during the present

study due to the lack of material. Heron & Bradford-Grieve (1995)

recorded O. curta from the Gulf of Naples, but their summary of

morphological distinctions between the species and O. scottodicarloi

was based on Sars’ original account, rather than on own observ-

ations (G. Heron, pers. commn to R. Huys). The authors noted

several as yet undescribed species close to O. scottodicarloi in their

samples from Naples, one of which probably had been O. waldemari,

which is also present in the Mediterranean. However, Heron &

Bradford-Grieve were not aware of this species, because the original

description of O. waldemari (dated 1994) did not appear until 1996.

Males of O. scottodicarloi can easily be separated from those

of O. media by the lack of sexual dimorphism in the coxobasal

seta on the antenna, which is long and plumose as in the female.

Males of O. waldemari, however, share this character and can be

distinguished from O. scottodicarloi only by minor details in the

form and ornamentation of seta IV on the antenna as well as in

proportional lengths of caudal setae (Table 5). For males of O.

curta corresponding sexual differences still need to be investi-

gated.

SIZE VARIATION

Female O. scottodicarloi from the central and northern Red Sea

were considerably smaller (0.48—0.52 mm, Bottger-Schnack et al.,

1989) than those from the southern Red Sea and Gulf of Aden, which

measured between 0.54—0.61 mm (numerous specimens measured

by the traditional method used by Bottger-Schnack et al., 1989). No

morphological differences other than size became apparent for

specimens from the two regions. Re-examination of specimens from

the central populations confirmed that the small morph had not been

confused with O. waldemari, which is similar in size. The observed

regional size difference of O. scottodicarloi (which occurred also in

the males) does not seem to be a temporary phenomenon, as the

lengths of specimens in the central Red Sea did not vary during

different seasons. Generally, O. scottodicarloi seems to exhibit a

wide range in body length: Heron & Bradford-Grieve reported

females between 0.59-0.72 mm, with distinct differences between

different oceanic areas. Female O. scottodicarloi from the northern

Arabian Sea and Eastern Mediterranean Sea as well as those col-

lected of Australia’s NW Cape (eastern Indian Ocean) measured

between 0.58—0.62 mm (unpubl. data) and were thus in the upper

size range of specimens from the Gulf of Aden. Itoh [in: Chihara &

Murano (1997)] recorded females between 0.50—0.64 mm and males

between 0.38—0.54 mm in size, however, the author was not aware

of O. waldemari, which might have been included in the lower size

range of his specimens. [A recent examination of two forms of

female O. scottodicarloi from the Western Subtropical Pacific kindly

made available by H. Itoh and S. Nishida showed that females of the

smaller form (0.49-0.62 mm) were conspecific with O. waldemari,

while the larger form (0.58—0.71 mm) belonged to O. scottodicarloi.

Compared to these data, specimens of O. scottodicarloi from the

central Red Sea appear to be even smaller than the lowest values

reported for this species so far. The occurrence of the small morph

seems to be a local phenomenon for this area. A possible explanation

for the reduced size of central populations may be the more extreme

environmental conditions in the central Red Sea as discussed by

Bottger-Schnack et al. (1989). Similar to O. venusta f. venella, the

small morph of O. scottodicarloi in the central Red Sea seems to be

a self-sustaining population, with no substantial recruitment from

the South.

OTHER RECORDS OF O. SCOTTODICARLOI

Heron & Bradford-Grieve (1995) recorded O. scottodicarloi from

various regions in the Atlantic and Pacific and recently Itoh [in:

Chihara & Murano (1997)] described it from Japanese waters. The

species was also found in great abundance in the Eastern Mediter-

ranean as well as in the northern Arabian Sea (Bottger-Schnack,

1994, 1996, as O. media f. minor). Sewell (1947) had reported O.

media f. minor from the Indian Ocean, however, the identity of his

size morph with O. scottodicarloi cannot be confirmed as the

specimens of Sewell are not available. The specimens might have

been confused with O. waldemari, which was found to co-occur

with O. scottodicarloi in both areas (see below). The identity of O.

media f. minor (sensu Malt, 1982b) will be discussed below under

O. waldemari.

GEOGRAPHICAL DISTRIBUTION

Oncaea scottodicarloi is distributed throughout the Red Sea proper

(Bottger-Schnack, 1988, 1990a, b, 1995, as O. media f. minor) and

can also be found in the northernmost part of the Gulf of Aqaba (own

unpubl. data). The species is one of the most abundant oncaeids in

the central Red Sea, usually accounting for 2-10% of the total

number of Oncaeidae in the water column down to 450 or 1050 m

depth, depending on region and season. In narrower depth layers,

even higher percentages, up to 15%, of all oncaeids are found

(epipelagic zone, winter, central Red Sea). Similar to O. media, O.

scottodicarloi exhibited a 2-tailed horizontal distribution during

summer, with high abundances in the central-northern Red Sea as

well as in the Strait of Bab al Mandab and Gulf of Aden, but low

abundances in the intermediate southern parts of the Red Sea

(Bottger-Schnack, 1995: Table 6). Consequently, its relative abun-

dance in the southern Red Sea was rather low (0.4/1.6%), but

strongly increased again further south. The proportion of O.

scottodicarloi in the Gulf was possibly underestimated, since sam-

ples in that area covered only the upper 250 m of the water column,

thereby excluding a potential mesopelagic population. In Table 8 of

Bottger-Schnack (1995), the abundance values of O. scottodicarloi

in the uppermost depth zone (0O—100 m) were inadvertently changed

between stations. The correct data for O. scottodicarloi are given in

Table 6 of the present account.

R. BOTTGER-SCHNACK

Table 6 Abundance (individuals beneath 0.25m?2) of O. scottodicarloi adults in the Gulf of Aden and different regions of the Red Sea during summer

1987.

Red Sea Main Basin

Gulf of Aden Bab al Mandab South, shallow South, deep Central North

Stn. 631 633 WA 708 703 682 663

Sampling Time N D D D D D D

Total water depth (m) 1400 1300 250 190 970 1890 1200

Depth layer (m)

0-100 2800 2000 5000 380 300 770 850

100-250 160 - 330 620* 81 220 380

250-450 - - 91 350 620

450-1050 ~ - The 28 130

Corrected data of Table 8 in B6ttger-Schnack (1995)

D = Day; N = Night; — = no data; Stn. = Station number

* = Depth range 100-175m

** = Depth range 450-950m

VERTICAL DISTRIBUTION AND VERTICAL MIGRATION

In the Red Sea, O. scottodicarloi had an epi- and mesopelagic

distribution mainly between the surface and 600 m depth in the Red

Sea; individual finds occurred down to 1250 m. Females exhibited a

bimodal vertical distribution pattern, with maximum abundances in

the epipelagic zone (0O-100m) and in the mesopelagic zone, at 200—

400(450) m depth (Bottger-Schnack, 1990a, b and unpubl. data from

summer 1987). The depth horizon of the two population centres

varied seasonally, with the epipelagic mode shifting to shallower

depth during winter, due to absence of a strong thermocline, as well

as regionally, with the mesopelagic mode shifting to shallower depth

in the South as compared to the central-northern area. In general,

however, the bimodal pattern was consistent. Diurnal vertical mi-

gration of moderate intensity was observed only for the epipelagic

part of the female population during autumn, which migrated up-

wards at night. During winter, the vertical migration behaviour of

the species was not investigated.

In contrast to females, males of O. scottodicarloi exhibited a

unimodal distribution pattern in the epipelagic zone, with highest

densities in the upper 5O m (winter, summer) or in the 50-100 m

zone (autumn). Within this depth zone, their diurnal vertical migra-

tion was stronger than in the females.

The mesopelagic population centre of O. scottodicarloi did not

show a regional downward shift between the central and northern

Red Sea in relation to the regional differences in oxygen profiles, as

had been observed for other oncaeids exhibiting bimodal distri-

bution patterns in the area, such as O. mediterranea and Lubbockia

squillimana (B6ttger-Schnack, 1990b). During summer, however,

the lower part of the population occurred somewhat shallower in the

southern Red Sea (200-300 m) than in the central-northern area

(250-400/450 m) (own unpubl. data). This might be related to a

corresponding regional shift in the depth of the oxygen minimum

(Bottger-Schnack, 1995). However, as the sampling variability

within each area was not investigated during the summer, the data

are inconclusive.

In the shallow part of the southern Red Sea (SRS) and in the Strait

of Bab al Mandab (BaM), both sexes of O. scottodicarloi were found

more or less evenly distributed between 40 m and the near-bottom

layer at 175 m (SRS) or occurred in maximum concentrations

between 20-60 m depth (BaM) during summer (unpubl. data). In

this area, the population occurred at the same depth as O. waldemari,

but the two species were vertically separated from O. media, which

was concentrated much shallower, at 0-20 m (BaM).

SEASONAL VARIATION IN ABUNDANCE (CENTRAL RED SEA)

O. scottodicarloi exhibited no consistent seasonal variation in

abundance in the central Red Sea, indicating that the species is not

markedly influenced by seasonal inflow of southern Red Sea waters

(Bottger-Schnack, 1995, see also above under ‘Size variation’).

Oncaea waldemari Bersano & Boxshall, 1994

Oncaea waldemari Bersano & Boxshall (1994): 29-41, Figs.1-6.

Oncaea media f. minor Malt (1982b): 130-133, Figs. lai, 2a-j,

3a—z’, 4a—o, Sa—h.

TYPELOCALITY.

52°20'W) .

PRELIMINARY NOTE. The type description by Bersano & Boxshall

(1994) includes some inconsistencies between text and figures, in

particular with regard to P5 exopod, that severely hampers a positive

identification of the species. Re-examination of the type material of

O. waldemari during the present study gave unequivocal evidence

that specimens from the Red Sea are conspecific. The species is

redescribed based on Red Sea specimens (figures) and the type

material. It includes several morphological details which were not

mentioned in the original account (e.g. labrum). Also, body dimen-

sions of specimens from the Red Sea, calculated by the different

methods used throughout this study, are provided. The urosome

segmentation of a typical female Copepodid Stage V, which has

been incorrectly described for oncaeid copepods in the literature

before, is described and briefly discussed.

Southern Brazilian shelf (3 1°40'-33°45'S, 51°00'—

MATERIAL EXAMINED.

(1) Southern Red Sea, 13° 40.0'N, 42° 37.4'E: Stn. 708; R/V Meteor

leg 5/5: collected 05 August 1987 with MSN 0.05 mm net (Haul

47/3); depth 20-40 m; total water depth 190 m.

(a) 2 29 in alcohol (BMNH 1998.2816—2817).

(b) 2 22, 1 6 in alcohol (ZMH K-39583).

(2) Southern Red Sea, 13° 40.0'N, 42° 37.4'E: Stn. 708; R/V Meteor

leg 5/5: collected 05 August 1987 with MSN 0.05 mm net (Haul

47/2); depth 40-60 m; total water depth 190 m.

(a) 2 dd in alcohol (BMNH 1998.2818—2819).

(b) 3 223 dd (RBS).

(3) Southern Brazilian shelf, 31° 40'-33° 45'S, 51° 00'-52° 20'W,

collected between 1988-1991 with cylindrical-conical closing

net (mesh size 0.15 mm); depth 0-25m: holotype 2 (Stn. 319,

BMNH 1994.6213), 2 2¢ paratypes (Stn. 309), 2 dd paratypes

(Stn. 319) (BMNH 1994.6214-6223).

(4) Eastern Mediterranean Sea, 34° 25.3'N, 26° 14.8'E, Stn. 35; R/V

Meteor leg 5/1: collected 20 January 1987 with MSN 0.055 mm

SEVEN SPECIES OF RED SEA ONCAEA

net (Haul 24/5); depth 0-50 m; total water depth 3400 m: 1 ?in

alcohol (RBS).

(5) Northern Arabian Sea, near Oman, 20° 44.3'N, 59° 40.5'E: Stn.

347; R/V Meteor leg 5/3a: collected 05 April 1987 with MSN

0.05 mm net (Haul 8/1); depth 0-50 m; total water depth ca 2500

m:5 22,2 dd for length measurements (RBS).

(6) Plymouth Sound, BMNH 1981.229, labelled Oncaea media

(sensu Malt, 1982): 3 29, 2 dé in alcohol.

(7) Adriatic Sea, 42° 38.5'N, 18° 02.0'E: Stn. ‘Lokrum 100’; col-

lected 26 September 1996 with 0.053mm Nansen type net; depth

0-25 m, vertical haul (leg. F. KrSini¢): 2 29(1 ovigerous), 1CV

2.1 3 in alcohol (RBS).

(8) Eastern Indian Ocean, NW Cape Australia, 21° 49.86'S, 114°

30.3,E: Stn. B; NWC 005/2; RV ‘Lady Basten’ leg 1630; col-

lected 26 October 1997 with 0.5 m WP-2 net with 0.073 mm

mesh, vertical haul; depth 0-20 m (leg. D. McKinnon): 2 22 in

alcohol (RBS).

(9) Western Subtropical Pacific, off Sagami Bay (Japan), 34° 15.0’N,

139° 45.0°E: Stn. 1; R/V Tansei-Maru leg KT-00-5; collected 09

May 2000 with Norpac net with 0.1 mm mesh; depth 0-200 m,

vertical haul (leg. S. Nishida; H. Itoh): > 30 29 in alcohol for

molecular analysis (RBS)

REDESCRIPTION.

(CV 9).

ADULT FEMALE (Figs. 24-26).

Body length: 637 um [Traditional method: 420-500 um, based on

21 specimens].

Exoskeleton well chitinized. Prosome 2.7 times length of urosome,

excluding caudal rami, 2.3 times urosome length including caudal

rami. P2-bearing somite without conspicuous dorso-posterior pro-

jection in lateral aspect (Fig. 24B). Integumental pores on prosome

as in Fig. 24A, B.

Proportional lengths (%) of urosomites 10.3 : 56.6: 10.7 : 8.7:

12.6. Proportional lengths (%) of urosomites and caudal rami 8.8 :

omenome ee = L1G 215.1.

Genital double-somite (GDS) of Red Sea specimens 1.7 times as

long as maximum width (measured in dorsal aspect) and 1.7 times as

Note illustrations are based on | (c) and (7)

| long as postgenital somites combined (Fig. 24C); GDS length to

/ width ratio of three female paratypes from Brazilian waters ranging

between 1.6-1.7 : 1, larger than reported by Bersano & Boxshall

| (1.4:1). Armature of genital apertures represented by | spine and 2

| small spinous processes, as figured by SEM-micrograph by Bersano

| & Boxshall (their Fig. 5D), the second minute one hardly discernible

| with light microscope in Red Sea specimens (Fig. 24H).

Anal somite shorter than caudal rami (Fig. 24C), 3/4 the length of

_ caudal rami in Red Sea specimens; variable in paratype specimens

| due to differences in length of caudal rami (see below).

Caudal ramus (Fig. 24F) about 2.3 times as long as wide in Red

| Sea specimens, longer in paratypes, varying between 2.4~2.8 : 1.

| Seta VI longer than dorsal seta VII and less than twice the length of

caudal ramus; seta VII distinctly longer than seta II.

Antennule (Fig. 24E) with armature formula as for genus, some

/ elements missing in original account of Bersano & Boxshall (their

| Fig. 2A); pore pattern as figured (Fig. 24E).

Antenna 3-segmented (Fig. 25A). Coxobasis with row of long,

_ fine spinules or setules near outer margin and with few additional

denticles on proximal part of outer (exopodal) margin, not figured in

| the original account of Bersano & Boxshall.

Labrum (Fig. 25B, C) as for O. venusta, except for additional

surface ornamentation (paired row of denticles) on anterior face, and

| integumental pockets either side of median swelling more pro-

| nounced (Fig. 25B).

Paragnaths (not figured) as in O. venusta.

Mandible (Fig. 25D) with surface of coxa unornamented; arma-

ture elements as for O. venusta, but dorsal blade with 2 additional

dentiform processes along dorsal margin (not only one as figured by

Bersano & Boxshall).

Maxillule (Fig. 25E) as for O. venusta, with the 3 innermost

elements on outer lobe similar in length, not innermost one shortest,

as erroneously figured by Bersano & Boxshall (their Fig. 3C).

Maxilla (Fig. 25F) and maxilliped (Fig. 25G) similar to O.

venusta. Proximal basal seta of maxilliped slightly longer than distal

one (Fig. 25G), not shorter, as erroneously figured by Bersano &

Boxshall (their Fig. 3E).

Swimming legs 1-4 with armature as for genus, surface ornamen-

tation as shown in Fig. 26A—D. Exopods and endopods with

ornamentation and spine lengths similar to O. venusta, except for

terminal spine on P4 endopod relatively longer and hyaline lamella

more narrow (Fig. 26D).

P5 with small exopodal segment clearly delimited from somite

(Fig. 24G), not fused as stated by Bersano & Boxshall (p. 35).

Exopod slightly longer than wide, ornamented with minute subapi-

cal pore and bearing 2 naked setae nearly equal in length, inner one

spiniform. Small naked seta arising from lateral surface of somite.

P6 (Fig. 24H) represented by operculum closing off each genital

aperture; armed with a spine and 2 small spinous processes, smaller

one hardly discernible in Red Sea specimens.

Egg-sacs paired, oval-shaped; each sac containing appr. 10-12

eggs (diameter ca 50 um).

ADULT MALE (Fig. 27).

Body length: 454 um [Traditional method: 340-360 um, based on 6

specimens]. Sexual dimorphism in antennule, antenna, maxilliped,

Pl and P3 (endopod), P5—P6, caudal ramus and in genital segmen-

tation.

Proportional lengths (%) of urosomites (excluding caudal rami)

9.3 : 66.8 : 3.3: 4.7: 4.2: 11.5; proportional lengths (%) of uro-

somites (caudal rami included) 8.3 : 59.6: 3.3: 4.2: 3.8: 10.0: 10.8.

Caudal rami about 1.4 times longer than wide, shorter than in

female. Caudal setae with proportional lengths as in female, except

for seta VI shorter than in female, about same length as seta VII and

2.0 times length of caudal ramus. The extreme shortness of seta VI

recorded by Bersano and Boxshall (about half the length of seta VII,

according to their Fig. 1D) could not be confirmed upon re-exam-

ination of male paratypes. Surface of genital flaps ornamented with

several rows of small spinules and area of minute pits as in Fig. 27E.

Antennule (Fig. 27B) with armature as for genus, pore pattern as

figured [distalmost element on segment 2 erroneously figured as

belonging to segment 3 in original account, Fig. 2B of Bersano &

Boxshall].

Antenna as in female, except for third spiniform element on

lateral armature being somewhat stouter than in female and fourth

element curved and ornamented with vestigial dentiform processes

along distal margin (arrowed in Fig. 27H). Seta on coxobasis long

and plumose as in female.

Maxilliped (Fig. 27C) similar to O. venusta.

Swimming legs 1-4 with armature and ornamentation as in

female; sexual dimorphism expressed in terminal process on P| enp-

3, being relatively longer than in female (Fig. 27J), and in longer

terminal conical projection on P3 enp—3, reaching 2/3 the length of

outer distal spine (Fig. 27K).

P5 (Fig. 27G) exopod not delimited from somite, general shape

and armature as in female, except for setae slightly shorter than in

female; small naked seta arising from lateral surface of somite as in

female.

R. BOTTGER-SCHNACK

eyo;

7 ES

)

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Chun

+100 “ER

fe” Bp

lye

Le es

Fig. 24 Oncaea waldemari, female (Red Sea) (A) Habitus, dorsal; (B) same, lateral (appendages omitted); (C) urosome, dorsal; (D) urosome, lateral; (E)

antennule; (F) caudal ramus, dorsal; (G) P5, dorsal; (H) P6.

SEVEN SPECIES OF RED SEA ONCAEA

posterior; (D) mandible, showing individual

Fig. 25 Oncaea waldemari, female (Red Sea) (A) Antenna, posterior; (B) labrum, anterior; (C) same,

elements; (E) maxillule; (F) maxilla; (G) maxilliped.

R. BOTTGER-SCHNACK

Fig. 26 Oncaea waldemari, female (Red Sea) (A) P1, anterior; (B) P2, posterior; (C) P3, anterior; (D) P4, anterior.

SEVEN SPECIES OF RED SEA ONCAEA 5

Fig. 27 Oncaea waldemari, male (Red Sea) (A) Habitus, dorsal; (B) antennule; (C) maxilliped, anterior; (D) urosome, dorsal; (E) urosome, ventral

(spermatophores immature), modified tip of posterolateral corner on P6 arrowed; (F) same, lateral; (G) P5, dorsal; (H) antenna, lateral armature of distal

endopod segment (ornamentation of seta IV arrowed); (I) caudal ramus, dorsal; (J) P1, distal endopod segment, anterior; (K) P3, distal endopod

segment, anterior.

R. BOTTGER-SCHNACK

FES

- a

10 um

Fig. 28 Oncaea waldemari, female copepodid stage Vv (Adriatic Sea) (A) Habitus, dorsal; (B) same, lateral (appendages omitted); (C) urosome, dorsal,

slit-like structures arrowed; (D) urosome, lateral; (E) slit-like structure; (F) P5, dorsal.

SEVEN SPECIES OF RED SEA ONCAEA

P6 (Fig. 27E) represented by posterolateral flap closing off

genital aperture on either side; covered by pattern of denticles and

area of minute pits as in Fig. 27E; posterolateral corners rounded,

generally with bifid tips, as in Fig. 27F (aberrant stunted corner on

right side arrowed in Fig. 27E).

Spermatophore oval (Fig. 27E, F), of variable size according to

state of maturity; swelling of spermatophore during development

not affecting shape and relative size of genital somite.

FEMALE COPEPODID STAGE V (Fig. 28).

Body length: 640 um [Traditional method: 550 pm].

Prosome 2.4 times length of urosome, excluding caudal rami,

about 2.1 times urosome length including caudal rami. Integumental

pores on prosome as in Fig. 28A, B.

Urosome 5-segmented; proportional lengths (%) of urosomites

11.7:51.7: 11.5: 8.9: 16.2. Proportional lengths (%) of urosomites

and caudal rami 10.0 : 44.0: 9.8: 7.6: 13.8: 14.8.

Second urosomite 1.8 times as long as maximum width (meas-

ured in dorsal aspect) and 1.4 times as long as postgenital somites

combined (Fig. 28C). Paired slit-like structure located close to

dorso-lateral margin at about midlength along second urosomite

(Fig. 28C), representing precursors of adult genital apertures (Fig.

28E). Pore pattern on dorsal surface similar to adult female; entire

surface covered with numerous small pits.

Anal somite shorter than caudal rami.

Caudal ramus with length to width ratio 2.0: 1. Caudal setae with

proportional lengths as in adult female.

P5 (Fig. 28F) similar to adult female.

Antennule, antenna, mouthparts and swimming legs not examined.

TAXONOMY

Oncaea waldemari has been referred to as Oncaea sp. AD (Bottger-

Schnack, 1994, 1995) or Oncaea sp. B (Bottger-Schnack, 1996) in

previous ecological studies. The original description of O. waldemari

by Bersano & Boxshall (1994) shows an inconsistency between text

and figures with regard to the exopod segment of female P5, which

is described as being fused to the somite (p. 35), but is figured as

being delimited (their Fig. 1B). Re-examination of the holotype and

female paratypes of the species during the present study gave clear

evidence, that the exopod of female PS is free (as in Fig. 24G), and

that specimens from the Red Sea are conspecific. Slight differences

in body dimensions between the two descriptions are mainly brought

about by the different methods used throughout this study (telescop-

ing of somites) or appear to be variable between specimens (e.g.

length to width ratio of caudal ramus as discussed below).

Based on the erroneous assumption of a fused PS exopod in the

female, Bersano & Boxshall (1994) placed O. waldemari in a small

group of oncaeids together with O. petila, O. ovalis, O. brocha and

O. compacta. The present redescription undoubtedly shows that the

species belongs to Oncaea s.str. and within this genus is most closely

related to O. curta, O. scottodicarloi and O. media. Major distinc-

tions between the three species are summarized in Table 5. Further

differences between O. waldemari and O. scottodicarloi include the

relative lengths of endopodal spines on P4, which are shorter in O.

waldemari. Males of O. waldemari can be separated from those of

O. media by the sexually dimorphic coxobasal seta of the antenna

(Table 5). Males of O. scottodicarloi lack this character; they can be

distinguished from O. waldemari by slight differences in element IV

on the antenna, the length of seta VII on the caudal ramus and the

ornamentation of the P6.

SIZE VARIATION

The body length of O. waldemari varies considerably in different

geographic regions (Table 7). Lengths of specimens from the Red

Table 7 Body length (mm) and length to width ratio of female caudal

ramus (CR) of O. waldemari from various locations.

Region g 3 CR

South Atlantic, 0.49-0.58 0.36-0.41 2.42.8: 1

Brazilian waters*

Mediterranean

(a) Adriatic Sea 0.51—0.60 0.40-0.46 2.4:1

(b) Eastern Medit. 0.50

Plymouth Sound 0.62-0.69 0.50-0.52 21

Red Sea 0.42-0.50 0.34—0.36 23E M1

Arabian Sea 0.47-0.52 0.34;0.38 no data

Eastern Indian Ocean, 0.48-0.54 ~ 2.2-2.4: 1

NW Cape Australia

Western Subtropical Pacific, 0.49-0.62 - no data

off Sagami Bay (Japan)

*after Bersano & Boxshall (1994)

Sea were in the lower end of the size range observed for other

regions or were smaller.

Beside the difference in size, the length to width ratio of caudal

ramus (Table 7) and the form of the genital double-somite seemed to

be variable within an area as well as outside. Female paratypes of O.

waldemari from Brazilian waters exhibited a variable length to

width ratio of caudal ramus, which was not noticed the Red Sea

material. Specimens from Plymouth (see below) had a somewhat

shorter genital double-somite than the type collection, and within

the populations from the Adriatic and from Australian waters,

differences in length to width ratio of the genital double-somite, as

well as of the caudal ramus, were noted. This indicates considerable

intraspecific variation (morphs) in O. waldemari, which makes it

difficult to identify the species. More detailed taxonomic studies on

this small and widespread species, including alternative methods

such as molecular analyses, are necessary to ascertain the taxonomic

status of these morphs.

The form of the bifid corners of genital lappets on male P6 was

slightly variable in Red Sea specimens. A corresponding variation in

male morphology was observed in O. media (cf. Fig. 16C) and had

previously been observed for another oncaeid, Triconia hawii

(Bottger-Schnack, 1999). It may be more widespread among oncaeids

than previously known.

OTHER RECORDS OF O. WALDEMARI

Malt (1982b) described the developmental stages of O. media from

Plymouth Sound and assigned the species to f. minor on the basis of

size. Re-examination of adult specimens from her collection indi-

cated that they are conspecific with O. waldemari, based on the

following characters (cf. Table 5): (1) ratio of genital double-somite

to rest of urosomites (2.0: 1), (2) position of genital apertures (1/4

the distance from lateral margin), (3) length ratio of seta VI on CR

(3/4 length of seta IV), and (4) length to width ratio of P5 exopod

(1.3: 1). The length to width ratio of genital double-somite of Malt’s

specimens was somewhat smaller (1.5 : 1) than in typical O.

waldemari and its form was not as elongate as in specimens from the

Red Sea, but resembled more that of O. scottodicarloi. However,

such variation was also observed in specimens from the Adriatic

during various seasons, and thus may indicate some intraspecific

variation. Males of O. media sensu Malt were assigned to O.

waldemari on the basis of antennary seta IV, which was slightly

curved and setiform (cf. Fig. 27H), rather than being spiniform as in

O. scottodicarloi.

The records of O. waldemari from very distant localities in

tropical and temperate regions of the three great oceans indicate a

world-wide distribution of the species. However, some morphological

differences were observed between these morphs, which need to be

clarified.

DEVELOPMENTAL STAGE CV 2 OF 0. WALDEMARI

The developmental stages (NI-NVI, CI-CVI @ and 3) of O.

waldemari were described by Malt (1982b, as O. media f. minor),

based on individuals reared in the laboratory and collected at sea.

According to her report, the female CV exhibits a 4-segmented

urosome (1 segment between genital double-somite and anal somite),

which is very unusual among copepods exhibiting the full comple-

ment of urosomites in the adult. Typically, the number of somites

between female CV and CVI does not change in female copepods

possessing a genital double-somite because the addition of one

abdominal somite from CV to CVI is compensated for by the fusion

of the genital somite and the first abdominal somite in the female

(e.g. Itoh & Nishida, 1995). Remarkably, the only other taxonomic

report of copepodid stages of oncaeids by Bjornberg (1972) also

describes the female CV of an unidentified Oncaea sp. with a 4-

segmented urosome. So both Malt and Bjornberg must in fact have

been dealing with a CIV. In the present study, late female copepodid

stages of O. waldemari and related species, such as O. venusta, O.

mediterranea, and O. scottodicarloi, were found with a 5-seg-

mented urosome, which exhibited no genital apertures (cf. Fig. 28C,

E). According to the general rule mentioned above, these stages

should be regarded as the typical female CV. A more detailed re-

examination of developmental stages of O. waldemari and other

oncaeid species is in progress.

GEOGRAPHICAL DISTRIBUTION

O. waldemari was confined to the Gulf of Aden / Strait of Bab al

Mandab area and the southernmost, shallow parts of the Red Sea. It

was not found in the deep oceanic area of the southern Red Sea or

further north in the central/northern regions (Bottger-Schnack, 1995,

as Oncaea sp. AD). Thus, it cannot be regarded as a typical Red Sea

species. In the Arabian Sea, it was present in somewhat greater

numbers near Oman than in the central Arabian Sea (Bottger-

Schnack, 1996, as Oncaea sp. B). The species is also present in the

Eastern Mediterranean, but was not separated from the closely

related O. scottodicarloi during earlier quantitative counts in that

area (Bottger-Schnack, 1997, as O. media f. minor).

VERTICAL DISTRIBUTION

Both sexes of O. waldemari occurred in the lower epipelagic zone,

with maximum concentrations between 20-60 m in the Gulf of Aden

and the Strait of Bab al Mandab. In the southern, shallow region of

the Red Sea (total water depth 190 m), the populations showed a

strong tendency to submerge: they were generally found between

20-175 m, with maximum concentrations between 125-175 m

(females) or 100—150 m (males).

Oncaea paraclevei sp. nov.

TYPE LOCALITY. Southern Red Sea, oceanic area, 15° 34.8'N, 41°

54.9'E, at O-50 m; total water depth 970 m.

MATERIAL EXAMINED.

(1) Southern Red Sea, 15° 34.8'N, 41° 54.9'E: Stn. 703; R/V Meteor

leg 5/5: collected 03 August 1987 with MSN 0.05 mm net (Haul

39/5); depth 0-50 m; total water depth 970 m.

(a) holotype 2 dissected on 11 slides (urosome partly damaged

after documentation (ZMH K-39578a-}); paratypes: 6 29in

alcohol, some ovigerous (ZMH K-39579).

(b) additional paratypes: 1 2 dissected on 10 slides (BMNH

1998.2820), 5 29 in alcohol (BMNH 1998.2821—2825).

R. BOTTGER-SCHNACK

sected on 11 slides, 6 29(1 ovigerous, 2 with spermatophores,

1 with small dorsal projection, 1 without dorsal projection)

in alcohol (RBS).

(2) Central Red Sea, 21° 25.53'N, 38° 01.91'E: Stn. 130; R/V

Valdivia leg 29: collected 28 October 1980 with MSN 0.1 mm

net (Haul 117/5); depth 0-20 m; total water depth ca 1960 m.

(a) 1 paratype @ in alcohol (BMNH 1998.2826).

(b) 2 paratype 22 in alcohol (ZMH K-39580).

(3) Northern Arabian Sea, near Oman, 20° 44.3'N, 59° 40.5'E: Stn.

347; R/V Meteor leg 5/3a: collected 05 April 1987 with MSN

0.05 mm net (Haul 8/1); depth 0-50 m; total water depth ca 2500

m: 2 22 in alcohol (RBS).

DESCRIPTION. Note illustrations are based on the holotype, except

for habitus, maxilliped and labrum.

ADULT FEMALE (Figs. 29-31).

Body length (measured in lateral aspect; from anterior margin of

rostral area to posterior margin of caudal rami, calculated as sum of

individual somites): 770 um [traditional method: 650-660 pum,

based on 2 specimens].

Exoskeleton well chitinized. Prosome 2.1 times length of urosome,

excluding caudal rami, 1.8 times urosome length including caudal

rami. P2-bearing somite with dorso-posterior projection in lateral

aspect (Fig. 29B), variable in conspicuousness according to degree

of telescoping of somites (Fig. 29a—c). Integumental pores on

prosome as indicated in Fig. 29A, B. Pleural areas of P4-bearing

somite elongate and with rounded posterolateral corners.

Proportional lengths (%) of urosomites 9.4 : 58.5: 9.9: 10.4:

11.8. Proportional lengths (%) of urosomites and caudal rami 7.9 :

ANOVA) 213-3) 3 tell 2 SS) E ISL

Genital double-somite oval-elongate, 1.6 times as long as maxi-

mum width (measured in dorsal aspect) and 1.9 times as long as

postgenital somites combined (Fig. 29C); largest width measured at

anterior third, lateral margins of genital double-somite rounded

anteriorly, tapering posteriorly. Paired genital apertures located near

dorsolateral margin at about 1/3 distance from anterior margin of

genital double-somite; armature represented by | spine and 2 minute

spinous processes (Fig. 29H). Double-curved sclerotization between

genital apertures, varying in form as in Fig. 28d, e; pore pattern on

dorsal surface as in Fig. 29C.

Anal somite 1.3 times wider than long; about 3/4 length of caudal

rami (Fig. 29C). Anterior margin of anal opening (vestigial anal

opening) with transverse row of minute denticles, which are variable

in number, ranging between 4 (holotype) and 10. Other surface

ornamentation as in O. venusta.

Caudal ramus (Fig. 29F) about 2.3 times as long as wide, variation

within (Fig. 29F) and between specimens ranging between 2.1—2.4

: 1 (4 specimens). Surface ornamentation and length ratios of caudal

setae as in O. clevei, except for longer outer terminal seta (IV),

reaching 4/5 the length of inner terminal seta (V).

Antennule (Fig. 29E) with relative lengths (%) of segments

measured along posterior non-setiferous margin 6.0 : 24.5 : 42.9:

10.9 : 4.3 : 11.4. Armature formula as for genus.

Antenna (Fig. 30A) as in O. clevei, except for seta III on second

endopod segment unipinnate.

Labrum (Fig. 30B,C) as in O. clevei, except for anterior surface

lacking row of denticles on proximal part of each lobe.

Mandible (Fig. 30D) similar to O. clevei, except for number of

dentiform processes along distal margin of blade C varying between

3 or 4 (Fig. 30D, d).

Maxillule (Fig. 30E), maxilla (Fig. 30F), and maxilliped (Fig.

30G) as in O. clevei.

SEVEN SPECIES OF RED SEA ONCAEA

Fig. 29 Oncaea paraclevei sp. nov., female (Red Sea) (A) Habitus, dorsal; (B) same, lateral (appendages omitted) [a—c: different degrees of telescoping of

somites, causing variation in conspicuousness of dorso-posterior projection on P2-bearing somite]; (C) urosome, dorsal, setae V and VI (right) not figured

[d, e: variation in form of sclerotization between genital apertures]; (D) urosome, lateral; (E) antennule; (F) caudal ramus, dorsal; (G) PS; (H) P6.

80 R. BOTTGER-SCHNACK

Fig. 30 Oncaea paraclevei sp. nov., female (Red Sea) (A) Antenna, anterior; (B) labrum, anterior; (C) same, posterior; (D) mandible, showing individual

elements [d: blade C, showing variation in number of dentiform processes]; (E) maxillule; (F) maxilla; (G) maxilliped.

os a

ee = Za

(B) P2, anterior; (C) P3, anterior; (D) P4, anterior.

<= Ae;

3S <<

Li ES

ZZILEZ

Fig. 31 Oncaea paraclevei sp. nov., female (Red Sea) (A) P1, anterior;

SEVEN SPECIES OF RED SEA ONCAEA

Swimming legs 1—4 (Fig. 30A—D) with armature as for genus and

surface ornamentation as in O. clevei. Spine length of exopodal and

endopodal segments not substantially different between the two

species, except for outer distal spine on P4 endopod, which is

slightly longer, measuring > 1/2 to 3/5 length of distal spine (Fig.

30D).

P5 (Fig. 29G) as for O. clevei, except exopodal setae slightly

curved, and small seta arising from lateral surface of somite plumose.

P6 (Fig. 29H) represented by operculum closing off each genital

aperture; armed with a spine and 2 small spinous processes.

Egg-sacs as in O. clevei.

ADULT MALE. Unknown.

ETYMOLOGY. The Greek prefix para-, meaning beside, refers to

the great similarity of the species with O. clevei.

TAXONOMY

O. paraclevei is closely related to O. clevei, from which it can be

distinguished by differences in the form of the genital double-

somite, which is more elongate, location of genital apertures, which

are more medial and somewhat more posterior than in O. clevei, and

by the resulting differences in form and location of sclerotization

between genital apertures. The combination of these characters

enables rapid separation of the two species during routine counts.

Further minor differences are found in ornamentation of the labrum,

in proportional spine lengths on P4 enp-3, and in P35 setae.

Males of O. paraclevei were not encountered during the present

study, although ovigerous females and specimens carrying

spermatophores were observed in the samples. Those males co-

occurring with females of O. clevei and O. paraclevei were all

assigned to the first species at closer examination, based on the

proportional length of endopodal spines on P4. However, as this is a

variable character in O. paraclevei, it might not be adequate to

separate males of these two species. Further investigation using

alternative methods, such as molecular analysis, may help to iden-

tify males of O. paraclevei, which cannot be distinguished

morphologically.

SIZE VARIATION

O. paraclevei exhibited variable body morphology, most obviously

in the form and size of the dorso-posterior projection on P2-bearing

somite, which varied greatly in size and conspicuousness according

to the degree of telescoping of somites (cf. Figs. 29a—c). Variation

was also expressed in the form of sclerotization on the dorsal surface

of genital double-somite, in the ornamentation of the anal opening,

in length to width ratio of caudal ramus and in proportional lengths

of terminal spines on P4 endopod.

GEOGRAPHICAL DISTRIBUTION

O. paraclevei was recorded from the central and southern Red Sea as

well as from the Gulf of Aden and the northern Arabian Sea. Its

potential distribution in the northern Red Sea still has to be con-

firmed, as the species was not separated from the closely related O.

clevei during the earlier quantitative counts.

VERTICAL DISTRIBUTION

No quantitative data on the vertical distribution of O. paraclevei are

available, because the species was not distinguished from O. clevei

by Bottger-Schnack (1990a, b, 1995). It generally co-occurred with

this species in the upper epipelagic zone, at 0-20 or 0-50 m depth,

and was absent in the meso- and bathypelagic zones.

ACKNOWLEDGEMENTS. Iam very grateful to Dr R. Huys (The Natural

History Museum, London) for his encouragement, fundamental help and

indefatigable advice in the study of oncaeid taxonomy and for arranging the

R. BOTTGER-SCHNACK

loan of O. waldemari and O. media sensu Malt. His critical comments helped

to improve the manuscript. Sincere thanks are due to Prof. J. Lenz, Prof. D.

Schnack and Drs H. Weikert and K. Schulz for continuous support during the

study. I wish to thank H. Itoh (Kawasaki, Japan) for translating the Japanese

text of his recent taxonomic study on oncaeids into English. I appreciate

assistance from the following persons or institutions, who provided speci-

mens, samples or information pertinent to the study: Dr G.A. Heron (University

of Washington, Seattle); Prof. J.-s. Ho (California State University, Long

Beach); the late Prof. A.G. Humes (Boston University Marine Program,

Woods Hole); Dr F. Krsinié (Laboratory of Plankton Ecology, Dubrovnik);

Dr D. McKinnon (Australian Institute of Marine Science, Townsville); Dr S.

Nishida (Ocean Research Institute, Tokyo); Dr S. Ohtsuka (Hiroshima Uni-

versity); Dr H. Postel (Institut fiirOstseeforschung, Warnemiinde);Dr D.K.

Steinberg (Bermuda Biological Station); Dr C.T. Walter (National Museum

of Natural History, Smithsonian Institution). The financial support for a visit

to The Natural History Museum, London, under the EC-funded TMR Pro-

gramme Bioresource LSF is gratefully acknowledged. This study was

supported by Deutsche Forschungsgemeinschaft grant Le 232/18 to J. Lenz.

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Review of the False Smooth snake genus

Macroprotodon (Serpentes, Colubridae) in

Algeria with a description of a new species

E. WADE

Middlesex University, Cat Hill, Barnet, Hertfordshire, EN4 8HT

CONTENTS

EG CAE) Gt CN MMe ee ace rc ede clr vs ccna ns sweat sade ucawauaseite eavenes setae icGnntnsanasseset ucidesceituss sbuadnsxinenunaisiseuoWeetadessvaoeddewssectvantetascerdsvoseases 85

DOP Sire LPT CIMOGRE NOC Gi tehame. sews suncde ceeres aad awdceutn aes nate aN Ca on caas coacet sueeaasssacetettancavousicdereass sas saneudsvavavavaudiveravuncncorestucueisecwansnecsensos 87

MOO rea aN Ste SeHET ND a eee a ere ec cree eee ane ce Reon ance sata dananrcbenestder asaacocanuanmrenanuevercanctacossuddnuaattetansttcivevcvcertstiaees 87

eC ET ANCA OLLI Caer cia ane tae gM ont aad ar soy Seams ane ee Mae ee ac aactnsod sea craneuceseaycesesna see cactar se ducsoe us suatath'scuaeetdsvendeeandsecadsoniecnivevs 89

BMeaOMECUIEN NCL ce tos. cease nina t a sane aaas samosas aMsiangeact aes ancagcosyasiosceace=eaeoter wughetrcesensaessaceeasernantevsnseritdzentendestfsusessdeveasestavsvectot des 100

MBS CAEISIS1 OM ibe ae street's sacs rcai ps seaeahin actos’ a dete utbadiar'cxe gah tar ac onianea «anaes causetsvoaaterceatees diane sageetanavdswaccarsdcudsusdvusaevantervugdeaciuvssaavaubsdubepeuseads 100

eee SEU U LCE GIMCMIUS) cccaetenecasosewcsceoneccearescauccden cer seeratoss concur dese suanasscaescee teat aaudes tae dkte voces Sunni tdacvetbse se essesvastersasipeadesstidiitdeverterssts 105

NOSE GUI GC Surceacecsstavauetecrarss rece ecesteess tttaas See mM ervas ovastraavles\oaisacevivenun vabtb ae ceieet teceeeaes seuttieessdecusuatesaih va Svacuavad saves vesedsseccesusviveeseveniviteuns 105

SyNopsis. The characters used to define Macroprotodon cucullatus mauritanicus Guichenot are re-evaluated. The taxa, M. c.

cucullatus, M. c. brevis and M. c, mauritanicus are considered to be full species. The populations occurring in northern Algeria-

from Algiers eastwards to Northern Tunisia are retained as M. mauritanicus. Those populations from Algiers westwards into

Morocco as far as Melilla are recognised as a new species, M. abubakeri. The populations inhabiting the regions further south are

morphologically closer to M. cucullatus Geoffroy Saint-Hilaire and are assigned to that species. The status of M. c. ibericus

Busack & McCoy is discussed.

INTRODUCTION

The subspecies of Macroprotodon cucullatus were first investigated

across its range by Pasteur & Bons (1960) and Bons (1967) in which,

on the basis of midbody scale differences, M. c. brevis was separated

from M. c. cucullatus. The former race was restricted to the Iberian

peninsula, Morocco exclusive of the east and northeast of that

country and the Western Sahara; the populations elsewhere were

retained in the nominate form (Fig. la).

Wade (1988), recognised the northern Algerian, northern Tuni-

sian and Balearic populations as distinct from the nominate race

mainly on the head and body pattern. He applied the available name,

M. c. mauritanicus Guichenot 1850, to which the eastern and

northeastern Moroccan populations were also assigned (without

seeing any of these specimens). In all other respects there was

agreement with Bons (Fig. 1b).

The latest review by Busack and McCoy (1990) rearranged the

distribution of the subspecies and added a fourth (Fig. lc). The

distributions of the four subspecies are as follows:

M. c. cucullatus (Geoffroy Saint-Hilaire, 1827) is restricted to

Libya, Egypt and southern Israel.

M. c. mauritanicus Guichenot, 1850, to which were assigned

populations of M. c. cucullatus sensu Wade, 1988, occupies the

whole of northern Algeria (south to 23°16'N), Tunisia (south to

33°42'N), and the Balearic Islands (Spain). The isolated populations

of Macroprotodon (M. c. cucullatus, sensu Wade, 1988) from the

Hoggar (Algeria) and Lampedusa Island (Italy) were assigned to M.

c. mauritanicus.

M. c. brevis (Giinther 1862) was excluded from Iberia but stated

to inhabit the whole of Morocco. The Western Saharan form of M. c.

cucullatus (sensu Wade, 1988) together with the eastern Moroccan

population of M. c. mauritanicus (sensu Wade, 1988) in the extreme

northeast of that country were assigned to M. c. brevis.

M. c. ibericus Busack & M.Coy, 1990 (M. c. brevis auct.) inhabits

the Iberian peninsula except the extreme north.

The sets of characters used by Busack & McCoy have been

reappraised. M. c. mauritanicus has been redefined. The analysis

revealed that more than one taxon is involved; the populations from

Algiers eastwards comprising the nominate form, those to the west

anew form extending into the extreme northeastern Morocco. (Fig.

7). The Balearic populations are considered to be sufficiently dis-

tinct as to merit separation at subspecific level and will be described

by Dr J. M. Pleguezuelos.

The main aim of this work is the resolution of the conflict of

opinions on the forms currently said to comprise Macroprotodon

cucullatus mauritanicus, the bulk of which occur in Algeria. The

value of the analysis would be diminished had the forms under

investigation not been compared with extralimital forms in regions

where the putative taxa meet. The Moroccan M. c. brevis whilst not

being part of the present analysis is here nevertheless taken into

account insofar as the populations which meet the western form of

M. c. mauritanicus sensu lato (=the new species) and the adjacent

populations of the nominate form are concerned.

With regard to M. c. cucullatus, the present analysis addresses

only those populations (sensu Wade, 1988) which occur in Algeria,

Tunisia and Lampedusa i.e. those which have been synonymised

with M. c. mauritanicus by Busack & McCoy, 1990 and those

extending to the Moulouya valley, northeast Morocco. Bons (1967:

Carte no.16) depicts a good number of localities but unfortunately

T =n

18° 24°

HM. c. cucullatus

2) M. c. brevis

aa)

20, 325 p

ee Be 1 36°

b 4

; 30°4

Bons, 1967

3) M. c. brevis

S88 M. c. mauritanicus

E22] M.c. brevis

M88 ™. c. manritanicus

Fig. 1 Concepts of the distribution of Macroprotodon of various authors.

\ 3

E. WADE

REVIEW OF ALGERIAN MACROPROTODON

the deposition of these specimens is not indicated. M. c. brevis, with

which the western population of M. c. mauritanicus (sensu Wade,

1988) is synonymised by Busack & McCoy (1990), is similarly

poorly represented in collections from areas where it would meet

neighbouring taxa. A full revision of M. c. cucullatus and M. c.

brevis is postponed pending study of material from eastern and

southern Morocco.

The subspecies are accorded full species rank. The balance of

evidence suggests that the distinctness between them is maintained

with little suggestion of any intergradation between the taxa.

MATERIALS AND METHODS

The material examined in the course of this work is listed at the end

of each species treatment. Abbreviations used are as follows: Acad-

emy of Natural Sciences, Philadelphia (ANSP); Natural History

Museum, London (BMNH); California Academy of Sciences, San

Francisco (CAS); Carnegie Museum of Natural History, Pittsburgh

(CM); Field Museum of Natural History, Chicago (FMNH); Mu-

seum of Comparative Zoology, Harvard (MCZ); Museo Nacional de

Ciencias Naturales, Madrid (MNCN); Muséum @ Histoire naturelle,

Genéve (MHNG); Muséum National d’Histoire Naturelle, Paris

(MNHN); Naturhistorisches Museum Basel (NMB); Rijksmuseum

van Natuurlijke Historie, Leiden (RMNH); Senckenberg Museum,

Frankfurt am Main (SMF); University of Colorado Museum, Boul-

der (UCM); National Museum of Natural History, Washington, D.C.

(USNM); Museum fiirNaturkunde, Universitat Humboldt, Berlin

(ZMB); Zoologische Staatssammlung, Miinchen (ZSM);

Zoologisches Museum der Universitat Hamburg (ZMH);

Naturhistorisches Museum, Wien (NMW); personal collection of

the author (EW).

The primary source for Fig. 7 has been the bioclimatic map of

Emberger et al. (1962) from which relevant data has been extrapo-

lated and forms the substance of Fig. 12. Other maps have also been

consulted. The bioclimatic map utilises meteorological data and

quantifies dry season drought taking into account not only tempera-

ture and precipitation but also atmospheric humidity as the

xerothermic index (x) or number of physiologically dry days in the

dry season (Emberger ef al., 1962, pp. 12-19). The area under

consideration broadly translates into north-south divisions of seven

subregions with their respective xerothermic indices ranging from,

in the more northern latitudes, the sub-mediterranean (x =<40) to

those in the south, the desertic (x =300+). The xerothermo-

mediterranean subregion (x =150—200) is seen as a transitional zone

between the Mediterranean and the Arid-Saharan regions. Translit-

eration of indigenous place names mainly follows the Michelin

(tourist) map 958 “Algérie-Tunisie’. The convenient symbols of

Busack and McCoy (1990) have been used.

Each synonymy includes most of the generic and specific combi-

nations. Pages refer to citations of the taxa. Most of the references in

which either sources are not given or the species is just mentioned,

have been excluded.

CHARACTERS EXAMINED

All specimens were sexed. Morphological characters used by Wade

(1988) and those used and considered by Busack & McCoy (1990)

were re-evaluated. Of those of the latter the sum of the ventral +

subcaudals, anterior temporals and postocular counts were not

analysed; all their other characters were found to be useful in

varying degrees. Additional characters such as maxillary tooth

counts and scale row reductions were incorporated into the analysis.

Dorsal scales rows. These are normally understood to mean those

counted arounud midbody. The reductions were recorded in the

manner proposed by Dowling (1951b). They occur behind the head

from 25 in four stages to 19 rows until anterior to the vent where the

count falls to 17 or more rarely to 15. Occasionally after 19 there

may be no further reduction. These are mostly due to fusion between

the 4th and 5th rows but those of the 4th and even the 2nd and 3rd are

not infrequently involved. Exceptionally, especially in individuals

with aberrant dorsal scalation, fusion may occur in other rows, in

particular the vertebral and paravertebral. For practical purposes

only the reduction from 21 to 19 on the neck and 19 to 17 anterior to

the vent, not taking into account any irregular fluctuations, have

been considered. The length of the head is defined in this work as the

distance from the snout to the retroarticular process of the mandible.

The Ist ventral is situated at or slightly posterior to this point in

vertical alignment.

In addition to dorsal counts at midbody Busack & McCoy (1990)

considered counts at a point one head length (not specified) behind

the head and one of same value anterior to the vent. These characters,

especially the latter, were found to be significant. The histograms

(Figs. 8 & 9) show (ordinate) each point of reduction at the relevant

ventral scute expressed as a percentage of the total ventral number.

A mean to the nearest integer was taken when the reductions were

unequal on either side. Subsequent fusions and divisions after the

main points of reduction were not quantified.

Ventral and subcaudal counts. These were counted in the manner

of Dowling (1951a) and are summarised in Table 1. Addition of the

ventrals to the subcaudals (Busack & McCoy, 1990) whilst not

completely eliminating sexual dimorphism did, nevertheless,

diminish it, whereas as discrete states they revealed clear sexual

dimorphism in all the taxa (Fig. 11). With the exception of M. c.

cucullatus (sensu Wade, 1988) interpopulational variation in these

characters was usually slight or too discordant to permit determination

of any geographic trends in either of the northern forms.

Supralabials. Nearly all the material examined here possessed 8

supralabials on each side of which the 4th & 5th meet the eye: one

individual of M. mauritanicus had 9 on both sides.

Supralabial-parietal contact. (Table 2). In the North African

material this character was useful in differentiating M. mauritanicus

from M. cucullatus .

Infralabials. (Table 2). The anterior series, i.e. those in contact

with the genials, normally numbered 6, there being few departures.

Significant differences were found in the posterior or ‘free’ series of

infralabials.

Maxillary teeth. (Table 2). The maxilla bears a series of six teeth

increasing in size followed by a diastema and a second numbering

from three to five of smaller size that precede the fangs. Dumeril and

Bibron (1854) gave the number of maxillary teeth as “6+3 ou 5+2’

which is understood here to mean 6, followed by 3 or 5+II. Significant

differences occurred in the second series of teeth, i.e. that which

precedes the fangs.

Head pattern. The variability in the elements of head pattern first

noted by Boulenger has been analysed by Wade (1988) and Busack

and McCoy (1990). From the confusing array of configurations in

E. WADE

Table 1 Variation (sample size, range, mean, and one standard deviation [SD]) in ventral and subcaudals in the species of Macroprotodon.

M. cucullatus

M. mauritanicus M. abubakeri

Males Females Males Females Males Females

Character:

Ventrals

Sample size 23 28 44 39 16 15

Range 153-163 163-182 162-178 168-193 159-175 172-185

Mean 158.17 Aya 170.55 183.51 167.5 180.33

SD 6225) 4.04 3.99 4.56 5.54 3.72

Subcaudals

Sample size 22 28 44 34 15 15

Range 42-54 39-53 46-59 43-54 45-50 41-51

Mean 48.41 44.57 57.91 48.38 49.7 45.27

SD 2.86 3.35 2.79 2.81 1.76 3.08

Ventrals + subcaudals

Sample size 22 28 44 34 15 1S

Range 196-217 207-226 212-235 222-241 209-224 213-237

Mean 207.05 216.36 222.45 231.65 216.74 2257

SD Sy) 5.49 55) Spa) 5.74 6.23

Table 2 Variation in character states the species of Macroprotodon.

M. cucullatus

M. mauritanicus M. abubakeri

Sample size: 51

Character:

Labial-parietal contact (L/R)

+/+ 1 (1.96%)

+/— or —/+ 0

== 50 (0.98%)

Posterior infralabials (L/R)

3/3 5 (9.8%)

3/4 or 4/3 0

4/4 or more 46 (90.2%)

Posterior maxillary teeth

3 0

4 24 (47.1%)

4/5 or 5/4 2 (3.9%)

5 25 (49%)

Nuchal collar

Entire 1 (1.96%)

Divided 30 (58.8%)*

Partially melanocephalic

Melanocephalic

‘Pale’ collar

5 (9.8%)

15 (29.4%)

Present (uninterrupted) 5 (9.8%)

Present (interrupted) 1 (2%)

Absent 24 (47.1%)

Intermediate or indeterminate 4 (7.8%)

Obscured 17 (33.3%)

Postorbital streak

Short 0

Interrupted or fragmented 13)@529 9a) ="

Entire 30 (58.8%)***

Embraces last supralabial 2 (3.9%)

Obscured or part. absorbed 8 (15.7%)

83 31

74 (89.2%) 13 (41.9%)

7 (8.4%) 3 (9.7%)

2 (2.4%) 14 (42.2%)

46 (54.4%) 2 (6.5%)

14 (16.9%) 1 (3.2%)

23 (27.7%) 28 (90.3%)

83 (100%) 0

0 30 (100%)

0 0

3 (3.6%) 22 (71%)

78 (93.4%) 6 (19.4%)

0 1 (3.2%)

2 (2.4%) 2 (6.5%)

81 (97.6%) 20 (64.5%)

0 10 (32.3%)

2 (2.4%) 0

0 0

22 (2.4%) 1 (3.2%)

63 (75.9%) 0

3 (3.6%) 5 (16.1%)

17 (20%) 26 (83.9%)

82 (98.8%) 27 (87.1%)

1 (1.2%) 0

* Laberal portions in most cases more forwardly situated and often united to the “V’-mark; ** occasionally rather short but never less than two supralabials in length;

*** highly variable in shape (see p. 89).

the pattern essentially three characters, the nuchal collar, the

interspace between it and the head pattern elements (the ‘pale

collar’) and the postorbital streak presented states which were useful

in the differentiation of the taxa: these are summarised in Table 2. In

some individuals, from north west Algeria in particular, there occurs

a bar, more or less at the level of the mouth, connecting the posterior

head pattern elements to the nuchal collar. The state of the forward

position of the lateral portions of the nuchal collar (1.e. occupying

the space of the ‘pale collar’) still serves to distinguish the taxa but

there is some reduction in reliability where it has been used to

differentiate some individuals of M. cucullatus from the eastern

Hauts Plateaux + southern Tunisia from the populations further west

and the mauritanicus complex to the north.

The V-mark is an inverted V or Y-shape which commences from

around mid-frontal its posterior ‘arms’ extending, entire or frag-

mented to the last supralabials, or further. The state where it either

REVIEW OF ALGERIAN MACROPROTODON

embraces the last supralabial — where it may unite with the postor-

bital streak — or extends on to the ‘pale collar’ has significance in

differentiating the taxa, in particular those of eastern Algeria and

Tunisia.

The postorbital streak is highly variable in the forms from the

southern and more arid regions (M. c. cucullatus sensu Wade, 1988)

and less so in those from the north. In Table 2 the categories are as

follows: ‘short’ denotes the condition in which the streak extends

not much further than the 6th supralabial and is always associated

with absence of labial smudging; there may occasionally be vestiges

of the streak on the 7th or 8th supralabials but should this condition

be unaccompanied by labial smudging it is counted as ‘short’:

‘interrupted’ or ‘fragmented’ covers the variety of conditions other

than ‘short’ or ‘entire’ (Fig. 3): ‘Embraces last supralabial’ denotes

the condition in which the the postorbital streak joins the posterior

extensions of the V-mark; in the ‘short’ condition the partial em-

bracing of the last supralabial is made up of the V-mark: ‘Entire’

denotes the uninterrupted condition. The short state (Fig. 4c & d) is

of some taxonomic significance in evaluating the northern forms.

Body pattern. This essentially consists of a conspicuous series of

vertebral spots or flecks, on one or both sides of each scale upon

which they are situated (Fig. 6c & f), with less prominent ones

alternating on the sides (Fig. 6a & e). These in varying degrees are

connected diagonally: they are sometimes manifest only when the

body is bent the imbrication being reduced thus exposing the parts of

the scales which are normally hidden. This condition, the

‘undifferentiated’ pattern, may merge into one in which the spots

expand to form a reticulated-tessellated pattern consistent with some

darkening of the ground colour leaving the spaces so formed as

discrete pale areas, the ‘textilis’ pattern. A third condition sometimes

occurs in which the ground colour differentiates into pale.and dark

longditudinal stripes, the ‘taeniatus’ pattern. This character is of

little value when applied to intraspecific differences between the

northern forms but very useful in differentiating those from the

southern populations.

Belly pattern. The chequered pattern and its variations and the

immaculate condition are present in all the taxa. In the northern

populations the progression from one to the other is by reduction in

number and also the size and density of the spots whereas in those

from the south the change is more often by progressive fading.

SYSTEMATIC ACCOUNT

Macroprotodon cucullatus (Geoffroy Saint-Hilaire)

Coluber cucullatus 1. Geoffroy Saint-Hilaire, 1827:151, Pl. 6, Fig.

3. Type locality, “Lower Egypt’ Type lost.

?Coronella laevis (part), Schlegel, 1837:65 .

Coronella cucullata: Ginther (part), 1858:35. Strauch, 1862:55.

Lallemant, 1867:28. ?Olivier (part), 1894: 25; ?1896:124.

Lycognathus cucullatus: Duméril & Bibron (part), 1854:926.

?Gervais, 1857:511, Pl. 5, fig. 6; 21869: 200, pl. XLIV, 2.

Lycognathus textilis: Duméril & Bibron, 1854:931.

Coronella textilis: Strauch, 1862b:67. Lallemant, 1867:29.

Psammophylax cucullatus: Jan, 1862:312.

Psammophylax cucullatus: Jan & Sordelli, 1866:312; Pl. i, Fig. 3A.

Lataste, 1881: 399.

Psammophylax cucullatus var textilis: Jan & Sordelli, 1866:312; PI.

i, Fig. 4

= §

/ \ | S

po str \ i retrartproc 1stv

post inf labs

Fig. 2 Head of Macroprotodon abubakeri (MNHN 1899.273 Laghouat,

Algeria) showing the characters used in the study): par = parietal, 6th

sup lab = 6th supralabial, post inf labs = posterior infralabials, Ist v =

Ist ventral, retr art proc = retroarticular process of the mandible, V-mk =

V-mark, temp str = temporal streak, po str = postorbital streak, nuc col =

nuchal collar, pal col = pale collar, lab smud = labial smudging, hl =

head length.

Macroprotodon cucullatus: Rochebrune, 1884:160. Boulenger (part),

1891:149-150. (part), 1896:175—176. Mayet, 1903:24. Boulenger,

1920a (1919):290; 1920 (1919b):305. Pellegrin, 1926:161.

Mosauer (part), 1934:58. Laurent, 1935:347. DeWitte, 1930:618.

Angel & L’ Hote, 1938: Tab. Angel, 1944b:419. Villiers, 1950:98.

Chpakowski & Schnéour, 1953:135 (forme méridionale), PI.

XVI, 2. Domergue (part), 1959: 166-167, photos. 16&17. Pasteur

(part), 1959 (1960):136-7?. Lanza & Bruzzone, 1960:55; Fig. 2,

Pl. xli. Davidson, 1964:15 (southern form). Slavtchev & Chadli,

1984:417.

Macroprotodon textilis: Rochebrune, 1884a:160.

Macroprotodon cucullatus melanocephala Mosauer & Wallis,

1927:305-310, Fig. 1. Type locality: Gafsa.

Macroprotodon cucullatus cucullatus: Pasteur & Bons (part),

1960:87. Bons & Girot (part), 1962:50. Kramer & Schnurrenberger

(part), 1963:506. Villiers, 1963:133+134. Bons (part), 1967:29,

84, 287, 90, 93, 162, 191,192, 231, fig. 16, map 16. Papenfuss,

1969:308. Bons (part), 1972:116; 1973:94. Villiers, 1975:134.

Wade, 1988:242—243, Figs. 3b—c & 4b. Gruber (part), 1989: 149—

151. (part) Sindaco, 1990:151.

Macroprotodon cucullatus mauritanicus: Busack & McCoy (part),

1990:268-269, Fig. 2 (Codes 2&4).

Macroprotodon cucullatus brevis: Busack & McCoy (part),

1990:269, 271.

DIAGNOsIS. A species of Macroprotodon characterised by four or

five maxillary teeth in the series preceeding the fangs. Head pattern

E. WADE

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REVIEW OF ALGERIAN MACROPROTODON

variable, body pattern usually tessellated-reticulated (the textilis)

pattern. Midbody scales in 19 rows. Sixth supralabial usually not in

contact with parietal. Usually four posterior infralabials. Separable

from M. mauritanicus mainly in the maxillary tooth count; 4 or 5 as

opposed to 3 and from that race and the new species (see below) in

its possession of the ‘fextilis’ pattern in contrast to the undifferenti-

ated or the ‘taeniatus’ pattern.

REDESCRIPTION OF THE HOLOTYPE OF Lycognathus textilis Duméril

& Bibron 1854. Adult female MNHN 849, (Figs. 3a & 6a): snout-

vent length 287 mm; tail 57 mm. Type locality, ‘Deserts of Western

Algeria’, collected by F. Schousboé.

Scalation. Scales at midbody 19, ventrals 168, subcaudals 45+1.

Dorsal scale row reduction formula:

1 3+4 (8) 4+5 (119)

3+4 (7) 4+5 (128)

Supralabials 8/8 the 6th well separated from the parietals on both

sides; infralabials 6+4 /4+4.

17 168

Maxillary teeth. 6+4+II; a diastema occurs between the Ist and

2nd tooth in the second series (both sides) i.e. that which precedes

the fangs.

Coloration. Spirit-faded, cream coloured the darker ground colour

only just discernible; dark markings undiminished.

Head pattern. | V-mark indistinct; postorbital streak fairly thin

extending obliquely from the eye to the 6th supralabial whereupon

it abruptly assumes a horizontal course, narrowing and petering out

on the 7th; nuchal collar fragmented, consisting of a median and two

lateral portions.

Body pattern. Conspicous black dorsal and dorsolateral markings

in varying degrees on the margins of the scales they occupy are

connected diagonally by black slivers usually confined to the inner

margins of the scales: the ground colour faded and only just

discernable thus poorly differentiating these from the pale patches.

Belly pattern. Immaculate, no trace of any markings visible.

Remarks. Doumergue (1901) believed the type locality to be in the

region of El Aricha (34°13'N, 1°01'W). This is supported to a large

extent by the character states of the holotype agreeing with almost

all those of the nearest populations, allowing for variation in small

sample sizes (Berguennt n=3; Tadmit-Djelfa n=3). Although the

condition of 4 maxillary teeth in the second series is normal for

material from eastern Morocco and western Algeria the diastema

between the Ist and 2nd is but a small step from a socket into which

an additional tooth would easily fit suggesting a recent departure

from the 5-toothed condition and that its provenance may lie closer

to the 5-toothed populations the nearest of which in Algeria is Oum

Ali.

DESCRIPTION. Based on 51 specimens including the holotype of

Lycognathus textilis. Restricted to material from north Africa between

3°W and 10°33'E. Lampedusa Island (Italy) is included.

Dorsal scale rows. 19 rows at midbody commencing from the 3rd

to the 11th ventral. In one individual from the Hoggar for a distance

of 20 ventrals and another from Gafsa for a distance of | the count

dropped to 17 before the 40th ventral: in two from southern Tunisia

the count was 18 in stretches of 4 and 5 ventrals. The count anterior

to the vent varied from 17 to 15. Seventeen in 40% (n=19), decreasing

to 15 in 30% (n=14) and reverting from 15 to 17 in 30% (n=14).

Ventrals. 153-182 (males 153-163, females 167-182).

Subcaudals. 43-54 (males 43-55, females 40-54).

Supralabials. Eight supralabials the 6th, with one exception (CAS

132803), always out of contact with the parietals.

Infralabials. Usually ten (6+4): one (USNM 195463b from

Berguennt) exhibited 7+3. Four (MNHN1916.31 from Kebili, SMF

57648, CAS 132803 from near Gabés and MNHN 1930.181 from

the Hoggar possessed nine (6+3).

Maxillary teeth. 6+4+II or 6+5+II.

Pattern. Nuchal collar, in specimens in which the pattern elements

are sufficiently discrete, is usually divided and sometimes frag-

mented; the lateral portions are fragments may be either vertically

aligned in relation to the median spot a few scales distant from the

last supralabial displaying a ‘pale collar’, or are anteriorly situated

so as to exclude it. Anterior head pattern very variable. V-mark

conspicuous, indistinct or even absent; connecting the nuchal

elements, discrete or fragmented. Postorbital streak highly variable:

it may be curved, oblique-straight, or straight; thick, thin or tapering;

entire or fragmented; a ‘spur’ may be present on its lower margin.

Melanocephalism present in varying degrees: jet black or diluted,

and/or partial to virtually complete.

Body pattern of the ‘rextilis’ type. Typically this consists of a

vertebral and two lateral series of spots connected diagonally by

black markings situated on the unexposed parts of the scales in-

volved and expands to form a reticulated-tessellated pattern with

some darkening of the ground colour leaving the spaces so formed as

discrete pale areas. The black markings may be conspicuous or they

may be occasionally reduced even to a condition in which the spots

are barely discernible except when the scales have been parted.

Belly pattern presents conditions in which the spots may oppose,

alternate or coalesce; they may progressively fade — often from the

centre — be uniform or be completely absent.

REMARKS ON THE SYNONYMY. ‘Two of the synonyms listed above,

Lycognathus textilis Duméril & Bibronand Macroprotodoncucullatus

melanocephala Mosauer & Wallis pertain to the form of M. c.

cucullatus from the regions under consideration in this work. Only the

holotype of Lycognathus textilis has been examined. With regard to

Macroprotodoncucullatus melanocephalaMosauer and Wallis (1927)

neither indicated the institution in which the single specimen was

deposited nor did they give a full description. The general appearance

of the specimen, its possession of the ‘fextilis’ pattern and its having

originated in Gafsa confirm its belonging with M. cucullatus .

GEOGRAPHIC VARIATION. Some of the character states show dis-

tinct geographical variation. In specimens from the west, in the

Hauts Plateaux as far as the Moulouya valley the nuchal collar

fragments when sufficiently discrete, i.e. not obscured by encroach-

ing melanocephalism, are often disposed to show a ‘pale collar’.

This state is also accompanied by that of 6+4+II maxillary teeth.

There is an eastward trend towards exclusion of the ‘pale collar’ and

an increase in the number of maxillary teeth, 6+5+II being the

condition most frequently met with in populations from southern

Tunisia. There is a noticable east-west reduction in the degree of

chequering in the belly patterns. Supralabial-parietal contact was

lacking in all except one individual CAS 132803 from near Gabés

and 4 posterior infralabials, or more, were manifest in all except for

three in USNM 196463b (near Berguennt, Morocco), MNHN

1916.31 (Kebili, Tunisia), SMF 47648 and CAS 132803 (near

Gabés, Tunisia). From Libya eastwards the incidence of 6+5+II

maxillary teeth increases, 6+4+II being of rare ocurrence in Egypt;

labial-parietal contact in 33-50% of specimens (Busack & McCoy,

1990) and in none did the posterior infralabial count exceed three. It

is likely that, given the magnitude of the range and the degree of

variation in M. cucullatus, further work will result in the description

of an additional subspecies. Should the character states which

distinguish any of the western forms from that occuring further east

be considered sufficiently trenchant as to warrant separation the

name available for the Algerian and Tunisian population is M. c.

textilis (Duméril & Bibron): it is to this form that those of the

northeastern Moroccan steppe from Berguennt to the Moulouya

valley belong. M. c. melanocephala Mosauer & Wallis would be a

synonym of M. c. textilis (see also p. 91).

AFFINITIES. The infralabial count is an exceptionally stable

character with respect to this species not only in the populations of

the nominate form in the area of this study but in those from Libya

eastward (Busack & McCoy, 1990). Any departures from the norm

are regarded as significant. Thus specimens from Kebili and Gabés

are seen as belonging to the eastern form of M. c. cucullatus.

The demarcation line drawn by Busack & McCoy (1990) between

these populations and those further east is the Tunisian-Libyan

border, approximates that which Cherlin (1990b) separates popul-

ations of Echis arenicola from those of E. pyramidum.

LOCALITIES

MOROCCO - Taourirt (34°25'N, 2°53'W) MNHN 1927.130-1: El Agreb

(34°20'N, 3°,08'W); Bons & Geniez, 1996: Ain Fritissa (34°09'N, 3°14'W);

Bons & Geniez, 1996: Rchida (33°55'N, 3°13'W); Bons & Geniez, 1996: El

Ateuf (33°51'N, 3°02'W) 1.5km S, aprox I1km S, aprox 201m S; Bons &

Geniez, 1996: Berguennt (34°01'N, 2°01'W); Pellegrin, 1926; Werner,

1929; Bons, 1960 MNHN 1925.219; 25km S; Bons, 1960; Bons & Geniez,

1996 and 43km W, Bons & Geniez, 1996: between Missour (33°03'N,

4°01'W) and Talsinnt (32°32'N, 3°26'W); Bons, 1967: Ain Benimathar

(34°0S'N, 2°00'W) 10 km S; Busack & McCoy, 1990 USNM 196463a-b:

Meridja (34°02'N, 2°24'W) approx 8kmW, approx 5km E; Bons & Geniez,

1996.

ALGERIA — ‘Deserts of Western Algeria’ =?E] Aricha (34° 13'N, 1°16'W)

MNHN 849 (holotype of Lycognathus textilis), Duméril & Bibron, 1854;

Gervais, 1857, 1869; Olivier, 1894: Wade, 1988; Busack & McCoy, 1990

Ain Sefra (32°45'N, 0°35'W); Werner, 1929; Busack & McCoy, 1990 MCZ

27501: Beni Ounif (32°3'N, 0°15'W), Foley, 1922: Méchéria (33°33'N,

0°17'W), Doumergue, 1901: Djelfa (34°40'N, 3°15'E) NMB 13774:

Douis=Bordj Douis (34°22'N, 2°43'E) MNHN 1927.149: Talmit=Tadmit

(34°17'N, 2°59'E) MHNG 899.55: Biskra (34°51'N, 5°44'E), Jan & Sordelli,

1866; Busack & McCoy, 1990 MNCN 1802: Bir Oum Ali=Oum Ali (35°01'N,

8°19'E), Wade, 1988; Busack & McCoy, 1990; BMNH 1920.1.20.3859:

Djebel Ilaman, Hoggar (23°16'N, 5°31'E), Angel & Lhote, 1938; Witte,

1930 MNHN 1930.181: ‘Algeria’ Busack & McCoy, 1990 USNM 10940.

TUNISIA — no locality MNHN 1894; ZFMK 4844: Fériana (35°57'N,

8°34'E2), Slavtchev & Chadli, 1984; Wade, 1988; Busack & McCoy, 1990;

BMNH 1920.1.20. 3109; MNHN 1885.447-8: Redeyef (34°24'N, 8°09'E)

MNHN 1961.664—S: Gafsa (34°25'N, 8°48'E) (Oasis), Busack & McCoy,

1990 UCM 37371: Djebel Hattig nr Gafsa, Mayet, 1903: between Gafsa

and Redeyef (“Bled Douara’ ), Busack & McCoy, 1990 UCM 37345-37358,

37360, 37363-37370, 48083; CM 54559: Gabés (33°53'N, 10°07'E) 15 km

N, Busack & McCoy, 1990 SMF 57648 and16 km N, ibid CAS 132803:

Chott Fédjédj (33°55'N, 9°20'E), Lataste, 1881: Kebili (33°42'N, 8°58'E),

Chabanaud, 1916b MNHN 1916.31: Tozeur (33°55'N, 8°08'E); Lavauden,

1926: ‘Mateur’ MHNG 1379.84.

SPAIN -— Southern Spain, Busack & McCoy, 1990 SMF 20170.

ITALY -— Lampedusa Island (35°31'N, 12°35'E), Camerano, 1891;

Lanza & Bruzzone, 1959; Busack & McCoy, 1990; Sindaco, 1990 UCM

31078.

E. WADE

Macroprotodon mauritanicus Guichenot

Macroprotodon mauritanicus Gervais 1848 (nomen nudum).

Guichenot 1850:22, Pl. i1. Fig. 2. Type locality: Algeria. Syntypes,

MNHN 2172, 1994.2339-41. Miiller 1882b:169.

Macroprotodon mauretanicum: Miller 1878:467; Macroprotodon

mauret. {abbreviation for mauretanicum], ibid: 666.

?Coronella austriaca: Gervais (part), 1836:312 .

?Coronella laevis: Schlegel (part), 1837: 65 .

Lycognathus taeniatus: Duméril & Bibron, 1854:930. Wade,

1988:242. Busack & McCoy, 1990:268.

Coronella taeniata: Strauch, 1862b: 57.

Lycognathus textilis: Busack & McCoy, 1990:268.

Lycognathus cucullatus: Duméril & Bibron (part), 1854: 926. Bosca,

1877:55. Miiller 1890:692.

Coronella cucullata: Giinther 1958:35. Strauch, 1862b:055.

Lallemant (part), 1867:29. Boettger (part), 1883; 1885:457-458:

1896:297. ?Olivier, 1894: 25; ?1896:124. Le Cerf (part), 1907:24.

Psammophylax cucullatus: Jan, 1862:312; Jan & Sordelli, 1866, Pl.

i, fig. 3B.

Macroprotodon cucullatus: Tristram, 1859; Boulenger (part),

1891:149-150; (part), 1896:175—176. Escherich, 1896:279. Gough

(part), 1903:468. ?Mayet, 1903:24. Chaignon, 1904:20-22.

Gadeau de Kerville, 1908:95. Werner, 1892:352; 1894:85; (part),

1909:620-621. Lavauden (part), 1926:158, 159. Hediger (part),

1929:24—26. Mosauer (part), 1934:58 . Chpakowsky & Schnéour:

1953:134 (forme septentrionale), Pl. XVI, 3, XVII, 7. Mertens &

Miiller (part), 1940:53. Bellairs & Shute, 1954:226. Lanza &

Bruzzone, 1954:55 Fig.2, Pl. xli. Mertens & Wermuth (part),

1960:184. Kramer & Schnurrenberger (part), 1963:508. Davidson,

1964:14,15. Schneider, 1969:250, Pl. ii. Johann, 1977:320-321.

Sura,1983: 29. ?Blanc, 1988:22. Bischoff & In den Bosch,

1991:170.

Macroprotodon cucullatus cucullatus: Bons (part), 1967:192, Fig.

16. Sochurek (part), 1985:71. Sindaco (part), 1990:151.

Macroprotodon cucullatus melanocephala: Busack & McCoy,

1990:269.

Macroprotodon cucullatus mauritanicus: Wade (part), 1988:242,

Figs. 3a, 5 & 4c. Busack & McCoy (part), 1990:268—269, Fig. 2,

codes 2 & 4. Gruber (part), 1989. Joger & Bischoff, 1989:100,

RIES:

DIAGNOsIS. A species of Macroprotodon characterised by a series

of three maxilliary teeth preceding the grooved fangs; divided

nuchal collar; a short, or less frequently, an entire postorbital streak.

Dorsal pattern of the ‘undifferentiated’ or the ‘taeniatus’ state.

Supralabial-parietal contact on both sides in the majority of speci-

mens. Dorsal scales in 19 rows, sometimes 20 or 21 at points

irregularly along the body but never in 21 rows continuously.

Ventral scutes 162-193, subcaudals 43-61. Readily distinguishable

from M. abubakeri and M. cucullatus by the number of maxillary

teeth in the series preceeding the fangs (3 as opposed to 4 and 4 or 5

respectively). From M. cucullatus it is separable by the high

incidence of supralabial — parietal contact (as opposed to virtually no

contact) and possession of the ‘undifferentiated’—‘taeniatus’ as

opposed to the “fextilis’ body patterns.

DESCRIPTION OF THE SYNTYPES. Males: MNHN 2172 (Fig. 4a),

1994.2341; snout-vent length 386-462 mm, tail 83-95 mm. Fe-

males: MNHN 1994.2339 & 2340 (Figs. 4b-c & 6c) snout-vent

length 384-392 mm, tail 68—95 mm.

Scalation. Dorsal rows 19 at midbody, reduction formula simple.

MNHN 2172 (male)

————

REVIEW OF ALGERIAN MACROPROTODON

1 1+2 (5) 44+5 (137) v 174

2+3 (5) 4+5 (135)

MNHN 1994.2341 (male)

1 34+4 (9) 4+5 (144) 164

3+4 (8) 4+5 (146)

MNHN 1994.2339 (female)

4+5 (12) 3+4 (171) 3=34+4(173) 3+4(174) 184

44+5 (12) 3+4 (169)

MNHN 1994.2340 (female)

4 4 l

TEI g OOS) 4, 186

4+5 (6) 4+5 (165)

Ventrals. males 164-174, females 184-186 . Subcaudals: males

51-54, females 47-51. Three possessed eight supralabials, one

MNHN 1994.2339 exhibited 9/8; the 6th in contact with parietal.

Ten infralabials (6+4) in three specimens one of which, MNHN

1994.2339, possessed an incomplete suture between the Ist and 2nd

shield in the second series on the left side (Fig. 4b). MNHN 1994.

2340 exhibited nine (6+3) infralabials.

Pattern. The nuchal collar is divided in all four specimens. A

subsidiary study of the mouth, in Pl. 2 (Guichenot, 1850), clearly

refers to MNHN 1994.2340 in which the infralabials number 9 (6+3)

without any partial fusion. Pileal part of V-mark indistinct in

MNHN 1994.2339 and 2341, discernible in MNHN 1994.2340 and

most conspicuous in MNHN 2172 in which it unites with the

temporal streak. Postorbital streak entire in MNHN 2172 and

1994.2339; short, terminating at the suture of the Sth and 6th

supralabial in MNHN 1994, 2340 and 2341, (Fig. 4c).

The belly pattern varies from moderately strongly chequered as

exemplified by MNHN 2172 in which the spots tend to be square and

arranged alternately to immaculate in MNHN 1994.2341. In MNHN

1994.2440 the markings are somewhat reduced, squared rather than

barred with more gaps; in MNHN 1994.2339 the spots are longer than

deep, often with spaces 1—3v or more and positioned centre and off

centre.

Guichenot’s figure (1850, Pl. 2) appears to be a composite draw-

ing. The head incorporating character states of MNHN 1994.2339,

namely the slender median portion of the divided nuchal collar, the

complete postorbital streak and the lack of the discernable V-mark.

The belly pattern is portrayed somewhat fancifully as being entirely

black except the distal portions of the ventrals uniting at intervals

with transverse lateral bands enclosing ocelli. This is most closely

approximated by belly pattern of MNHN 2172. which is the most

densely chequered.

DESCRIPTION. Based on 83 specimens including the syntypes: the

dubious specimen, (MHNG 1379.68 from ‘Bordj Bou Arréridj’) is

excluded and is discussed below.

Scalation. Dorsal scales normally in 19 rows along the body but

occasionally reaching 20 and 21 in short stretches (1-8 ventrals).

Scale row reduction usually simple: 2nd + 3rd, 4th + Sth and

sometimes | st + 2nd, 5th + 6th and even vertebral and paravertebrals

are involved.

In ten specimens fusions, divisions and reformations of scale

rows at irregular intervals along the body occurred, where the counts

should have been 19, reaching 20 and 21 from the position of the Ist

ventral to virtually the entire distance . In two individuals, ZSM

1985 (‘Annabes’, Algeria) and ZMB 143381 (Tunis), the reduction

on the neck from 21 to 19 was due to fusion of the 3rd + 4th and 4th

+ Sth rows; thereafter divisions and fusions occured erratically

along the body involving the vertebral and paravertebral series alone

until the count dropped to 17 where again fusion of the 3rd + 4th and

4th + 5th rows occurred.

Ventrals 162—193 (males 162-178, females 175—193); subcaudals

43-61 (males 46-61, females 43-59).

Supralabials. Normally eight but occasionally an additional scute

may arise. Contact with parietal in seventy four (89%) of which fifty

seven (74%) exhibited sutural contact, sixteen ‘point’ contact on one

side and one ‘point’ contact on both sides. Contact was lacking on

one side in seven and on both sides in two specimens, one of which

possessed nine supralabials on both sides.

Infralabials. 6+3/6+3 in 44 individuals (53%); twenty four

intermediates with mostly 6+3 or 6+4 on one side; 6+4/6+4, in

thirteen (16%): two possessed 7+4/7+4.

Maxillary teeth. 6+3+II, 6+2+II (one specimen).

Pattern. nuchal collar divided in all specimens except three in

which the collar was entire (BMNH 59.3.29.17, Algiers; NMB

2016, Médéa and BMNH 47.10.30.203A, Tunis) and two which

were black headed (Médjana, Algeria). V-mark inverted *Y’, *V’, or

lyre-shaped sometimes originating as a T-shaped mark on frontal

and supraocular scutes, bifurcating, the ‘arms’ uniting posteriorly

with the postorbital streak around the last supralabial. Anteriorly the

V-mark may be in varying degrees obscured by the occasional dense

concentrations of the pileal vermiculations and occasionally the

‘arms’ are separated from the pileal part.

Postorbital streak short in 76%, reaching the suture of the Sth &

6th supralabial but, save for a few instances, not extending much

beyond it, recommencing at and embracing the last supralabial

posteriorly, sometimes uniting with the temporal streak. It is always

accompanied by unmarked anterior lips save for slight overspill

beyond its suture with the loreal. Entire postorbital streak uniting

with the V-mark together with the anterior part of the mouth along

its line smudged with black in only 17 individuals (20.5%).

In one of the black headed individuals (Fig. 4, d.) most of the head

above is ink-black save for a pale streak in the canthal region,

involving the preocular, supraocular and prefrontal as far as the

suture of that shield with the internasal. The lower part of the head,

supralabials and adjacent lower parts of the temporals are creamy

white: the postorbital streak terminates at the suture of the 5th + 6th

supralabial and there is no smudging on the mouth. In the other black

headed specimen the whole of the top of the head is black; the

supralabials below the position of the postorbital streak and the

infralabials are white, the underside of the head nearly so.

The body pattern usually consists of a vertebral series and three

longditudinal rows of spots on each side of the body disposed at

intervals of 1-3 scales. The vertebral spots vary in size from flecks

barely covering a single scale to clusters of as many as 6 scales;

where a whole scale is involved it is more or less bisected

longditudinally. The dorsolateral and flank series of spots are less

conspicuous and tend to occupy the margins of scales, in some

specimens small spots less than a scale wide, paler than the ground

colour distributed in conjunction with the dark spots dorsally and

dorsolaterally sometimes giving the impression of imperfect ocelli.

The ground colour may be uniform but is often differentiated into

darker striae on the 2nd and 3rd scale rows from the ventral, and

sometimes the adjacent portion of the fourth, and a darkening of the

median 7-9 rows leaving a pale stripe some 2-2.5 scales wide.

These spots are obliquely connected by diagonal series of black

marks at the scale pockets, being often seen only when the scales are

E. WADE

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REVIEW OF ALGERIAN MACROPROTODON

separated. The outermost row of spots takethe form of oblique bars

which border the ventrals.

Belly chequered in varying degrees (Fig. 6c, d & e). These

markings vary from broad, angled bars and/or rectangles, centred or

otherwise, with few separations to conditions in which the marks

may be reduced and widely spaced and ultimately intergrading to the

immaculate state. Belly immaculate or nearly so in about 40 indi-

viduals (48%): this condition is never associated with that in which

labial smudging or entire postorbital streaks occur.

REMARKS ON THE SYNONYMY. The author was unable to locate the

holotype of Lycognathus taeniatus Duméril & Bibron 1854. The

specimen has been lost (Brygoo, in litt., Wade, 1988) for well over a

century. At least neither Strauch, (1862) nor Doumergue, (1901)

claim to have seen the specimen, although they both presented

extracts from the original description. The type locality was given as

‘Deserts of Western Algeria’ (?=El Aricha) as for L. textilis. El Aricha

is aregion of subsaharan steppe in which neither of the two forms of

the M. mauritanicus complex are likely to occur. The original

description is principally concerned with the body pattern, the

‘taeniatus’ pattern sensu Wade, (1988) shades of which may occasion-

ally be found in any of the taxa. Clear evidence of any diagnostic

features is lacking. F. Schousboé who collected the holotype was

principal translator for the French Army in Algeria and was closely

associated with the ornithologist Commandant V. Loche whotravelled

extensively in that country. In consequence Schousboé was unlikely

to haverestricted his activities solely to western Algeria: the specimen

therefore could have originated from anywhere in Algeria. As defined

here Macroprotodon mauritanicus is restricted to northeastern Alge-

ria (Algiers eastwards) and northern Tunisia.

GEOGRAPHICAL VARIATION. Scarcely any geographic variation

discernible: the only two melanocephalic individuals were found in

the Médjana region, Algeria (38°04'N, 4°40'E).

AFFINITIES. Resembles most closely the new species described

below (p. 105).

DISTRIBUTION. Northern Algeria from Algiers eastwards and south

as far as Biskra (34°51'N, 5°44'E): northern Tunisia south to Sidi

Bou Zit (32°02'N, 9°30'E).

LOCALITIES

ALGERIA - no locality Boulenger, 1896; Duméril & Bibron, 1854; Gervais,

1857; 1869; Olivier, 1894; Busack & McCoy, 1990 BMNH 93.11.30.2;

MNHN 2172, 1994.2339-41 (syntypes of Macroprotodon mauritanicus)

Algiers (36°47'N, 03°03'E); Giinther 1858; Boulenger, 1896; Hediger,

1935; Busack & McCoy, 1990 BMNH 59.3.29.17; MNHN 3735; ANSP

3484; 3488; NMB 2422; USNM 56437 as M. c. mauritanicus fide Busack &

McCoy, 1990=Coronella girondica; Maison-Carrée=El Harrach, 36°43'N,

30°08'E, Le Cerf, 1907: Médéa (36°16'N, 02°45'E); Hediger, 1935;

Miiller1882 NMB 2016: Ben Arous=Bordj ben Arous (36°01'N, 3°25'E)

ZFMK 14316: Lalla Khadidja (36°27'N, 4° 14'E) MHNG 847.69: Médjana

(34°08'N, 4°40'E) 18 km NW EW 92.5, 92.9.1-6, 93.9.1—3: Guellal (36°02'N,

705°21'E); Sura,1983: Sétif (36°12'N, 05°24'E); Werner, 1909: Bougie=

Bejaia (36°45'N, 5°05'E); Jan & Sordelli, 1866; Bellairs & Shute, 1954; Sura,

1983 BMNH uncatalogued: Massif de I'Aurés; Sindaco, 1990: Col de

Tibharine, (35°21'N, 6°25'E) Gruber, 1989; Bischoff & In den Bosch, 1991

ZFMK 49602: Batna (35°34'N, 6°11'E); Wade, 1988; Busack & McCoy,

1990 BMNH 1920.120.1525 (skeleton), 1920.1.20.3108; MCZ 6421; ZFMK

4841-2: Biskra (34°51'N, 5°44'E) Jan & Sordelli, 1866; Boettger, 1885;

Busack & McCoy, 1990 MHNG 2031.98; SMF 20169 Constantine (36°22'N,

6°37'E); Werner, 1929; Busack & McCoy, 1990 MCZ 144.371: Guelma

(36°28'N, 7°26'E); Gough, 1903: Hammam Meskoutine (36°27'N, 7°16'E);

Anderson, 1882; Busack & McCoy, 1990 BMNH 91.5.4.14:

Bugeaud=Seraidi (36°55'N, 7°40'E); Busack & McCoy, 1990 BMNH

1920.1.20.1186; CM 58420-1: Bone, Bona, ?7Annabes=Annaba (36°54'N,

7°46'E); Hediger, 1935; Wade, 1988; Busack & McCoy, 1990 BMNH

1920.1.20.1620; NMB 2018-2024; MNHN 1981.31; 30km E ZSMH 77/

1985.14: Mt Edough (36°53'N, 7°37'E), Werner, 1892: Teniet (locality

unknown) MNHN 8507. The specimens NMW 25840 Algiers and MHNG

1379.68 Bordj Bou Arréridj (36°4'N, 04°46'E) have been identified as

Macroprotodon brevis:

TUNISIA —- no locality Busack & McCoy, 1990 MNHN

1891.316,1908.88; ZFMK 4843, 4845; FMNH 83650-1: fles Galita

(37°32'N, 8°56'E); D’Albertis, 1878; Davidson, 1964; Lanza & Bruzzone,

1960; Schneider, 1969): Ain Drahm (36°47'N, 8°42'E); Gadeau de Kerrville,

1908; Busack & McCoy, 1990 BMNH 1906.8.29.20-21, MNHN 1891.316—

322: road to Sedjenane (37°03'N, 9°14'E) just past junction to Cap Serrat

(37°14', 9°13'E); Busack & McCoy, 1990 USNM 165874: Bizerte (37°17'N,

9°52'E); Busack & McCoy, 1990: Bulla Regia (36°33'N, 8°44'E); Davidson,

1964: Mateur (37°03'N, 9°40'E) MHNG 1379.81-83: Mégrine (36°46'N,

10°14'E); Chpakowski & Chnéour, 1958: Zaghouan (36°24'N, 10°09'E);

Chaignon, 1904: Tunis (36°48'N, 10°11'E); Giinther 1858; Boulenger, 1896;

Escherich, 1896; Werner, 1909; Busack & McCoy, 1990 BMNH 47.10.203.1—

3; ZMB 14338, 14381, 14854, 51650—-1; between Sidi bou Said (36°47'N,

9°49'E) and Hammam Lif (36°44'N, 10°20'E); Busack & McCoy, 1990

SMF 34340: Cap Bon (36°45'N, 10°45'E); Blanc, 1988: Soukra (36°53'N,

10°15'E; Mosauer, 1934: Bir el Bey=Bordj el Bey (36°02'N, 10°18'E);

Davidson, 1964: ?Sfax (34°44'N, 10°45'E) MNHN 1982.115: Sidi bou Zit

(35°02'N, 9°30'E); Chaignon, 1904; Busack & McCoy, 1990 FMNH 75967.

NMW 19193.3 Tunis has been identified as Macroprotodon brevis.

NORTH AFRICA - no locality ZMH R04321.

‘LEVANT’ — MNHN 3736b.

Macroprotodon abubakeri sp. nov.

Lycognathus cucullatus: ?Gervais, 1857: 511, Pl. 5, Fig. 2.

Coronella cucullata: Strauch (part), 1862: 55. Lallemant (part),

1867:28.

Macroprotodon cucullatus: Doumergue (part), 1901: 282-288; PI.

XXI, fig.12a. Gough (?part), 1903: 468 . Werner (part), 1909 161;

1929: 11-12, ?20; (part), 1931: 300-301. Zulueta (part), 1909:

354. ?Maluquer, 1917b: 564. Hediger (part), 1935: 24-26.

Sochurek, 1956: 87+ pl. (part), 1979:221.

Macroprotodon cucullatus cucullatus: Pasteur (part), 1959

(1960):136—7 ?. Bons (part), 1960: 64, 74; (part), 1967: 29, ?87,

192, Fig. 16, Map no.16; (part), 1972: 116. Pasteur & Bons (part),

1960: 116. Bons & Girot, 1962: 50.

Macroprotodon cucullatus brevis: Busack & McCoy (part), 1990:

269-271.

Macroprotodon cucullatus mauritanicus: Wade (part), 1988: 242,

Fig.1. Busack & McCoy (part), 1990: 268-269. Bons & Geniez

(part), 1996: 226-228, (314, appendix in English).

HOLOTYPE. BMNH 1913.7.3.14 (Figs. 5a & 6f) from Oran, Alge-

ria; collected by the Hon. Walter Rothschild and Dr. Ernest Hartert.

Named in honour of M. Aboubakeur Sid-Ahmed, a naturalist from

Tlemcen, Algeria.

PARATYPES. MNHN 3734 (Fig. 5b); MCZ 27502, 29919; FMNH

42840: all from the region of Oran (see end of species description for

locality data).

DIAGNOSIS. A species of Macroprotodon characterised by a series

of four maxillary teeth preceding the fangs, nuchal collar entire or

sometimes divided; postorbital streak usually entire uniting with the

‘arms’ of the V-mark. Dorsal body pattern of the ‘undifferentiated’

or “taeniatus’ state. Belly usually chequered. Supralabials separated

from the parietals or in contact with them. Four posterior infralabials

in the majority of specimens.

Close to M. mauritanicus from which it is readily separated by

possession of an additional maxillary tooth. Postorbital streak usually

unbroken. Separable from M. c. brevis by 19 dorsals or in patches of

21 along the body (cf 21 uninterrupted rows); sutural contact

E. WADE

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REVIEW OF ALGERIAN MACROPROTODON

between supralabials and parietal in less than half; 4 as opposed to 3

in the posterior series of infralabials; postorbital streak usually

separate from the ‘arms’ of the V-mark. The entire nuchal collar and

lack of the ‘textilis’ pattern distinguishes it from the parapatric M.

cucullatus .

DESCRIPTION OF THE HOLOTYPE. Adult male snout-vent length

386 mm; tail 78 mm from Oran, Algeria.

Scalation. Scales at midbody 19, ventrals 170, subcaudals 51+1.

Dorsal scale row reduction formula:

1 3+4 (6) 44+5 (115)

3+4 (6) 4+5 (121)

Supralabials 8/8, the 6th falling short of contact with the parietal

on both sides; infralabials 6+4/6+4.

17 — vertebral (170) 16 170

Maxillary teeth. Arranged ina series of six teeth increasing in size

followed by a diastema and a series of four smaller in size preceding

the fangs on both sides of the maxilla.

Coloration. Colour (in alcohol) grey with black and light brownish

markings.

Pattern. V-mark poorly defined at its apex becoming more distinct

posteriorly where it unites with the postorbital streak around the 8th

supralabial and the 10th infralabial extending also beyond the suture

on the 9th. Postorbital streak dark, black on its upper margin

commencing on the Sth, occupying most of the 6th and 7th

supralabials and extending on to the 9th and the whole of the 10th

infralabial where it unites with the V-mark. From the anterior

margin of the eye it continues as a widening, forwardly directed

emarginate band its lower margin crossing the apices of the 3rd—Sth

supralabials and enclosing the loreal and most of the nasal including

the nostril, extending on to the anterior parts of the prefrontals and

most of the internasals but failing to contact the rostral. Anteriorly,

the rostral, supralabials and less conspicuously the infralabials are

smudged with black along the mouth. Temporal streak ill defined.

Nuchal collar not divided dorsolaterally being about three scales

wide at the narrowest point, extending forwards sagittally and

terminating to a point some two fifths along the suture dividing the

parietals.

Body pattern consists of a uniform ground colour; a vertebral

series of somewhat darker, emarginate, more or less ‘x’ shaped

spots, at points of articulation of 4—6 scales, is disposed at intervals

of about two scales distance along the body. Two other series of

alternating dorsolateral and lateral spots reduced in size, not visible

much beyond the imbrications of the scales, are imperfectly con-

nected by markings situated at the scale pockets: these being best

seen on exposure of the scales by bending the specimen. The belly

is marked by rectangular bars sometimes alternating, occasionally

disposed sequentially in single or fused units; these are occasionally

separated by unmarked ventrals (Fig. 6f).

DESCRIPTION. Based on 31 specimens including the holotype: the

dubious specimen, (MHNG 1214.40 *Chellala’ ) is excluded and is

discussed below (p. 105).

Scalation. Dorsal scales in 19 rows along the body but reaching 20

and 21 in stretches of 1 to 51 and in one male (BMNH 1931.2.9.2

from Algeria, without precise locality) the count, save for two short

intervals (1v) where the count dropped to 19, the dorsals numbered

21 from the 3rd to the 118th ventral. As in M. mauritanicus the

counts can fluctuate as much as fifteen times. Ventrals in males

(n=16), 154-175; females (n=15), 172-185. Subcaudals in males

(n=15), 46—53; females (n=15), 42—52.

Supralabials. Eight supralabials on each side. in which thirteen

individuals exhibited contact between the 6th supralabial and the

parietal, in eleven of these contact was sutural, two exhibited ‘point’

— contact; three showed contact on one side and in fourteen contact

was lacking.

Infralabials. Contact on both sides (n=25), single specimens with

6+5/64+4, 6+5/6+4s, 5+4 on both sides, 6+4/64+3 and two individuals

with 6+3 on both sides.

Maxillary teeth. 6+4+II in all the material examined.

Pattern. Nuchal collar entire n=22 (71%), divided n=6 (19%). V-

mark variable; indistinct, solid or obscured by vermiculations,

fragmented and/or discrete in varying degrees but often

(approximately 40%) entire. In all except four individuals the V-

mark unites with the postorbital streak. Postorbital streak embracing

the 8th supralabial (n=28), either entire (n=21) or interrupted

(n=3).The V-mark-postorbital streak connection is separated from

the nuchal collar by 2-4 scales (n=18) but may be connected to the

nuchal collar by a bar at the level of the mouth of 0.25-2 scales

thickness.

Extensive melanocephalism occurs in two individuals: the top of

head and snout is black, save for a white streak on the first 5 or so

supralabials and another, oblique, occupying the upper part of the

supralabials+the lower parts of the temporals extending from the

eye to the corner of the mouth, or beyond. In one (BMNH 1931.2.4.23

from Algeria without locality) the white streak commencing at the

eye continuing uninterrupted beyond the 8th supralabial for 3 scale

rows abruptly changing course downwards and forwards termin-

ating near the last infralabial from which it is separated by a single

scale. The throat is heavily smudged with black: the penultimate

infralabial save for its posterior margin is white. The other speci-

men, (MCZ 27502 from Djebel Mourdjadjo) is similar but differing

in that the white streak is shorter, more oblique and terminates at the

posterior margin of the 8th supralabial; the black pigment on the last

two infralabials and the postorbital streak formes its lower margin.

the black pigmentation is concentrated on the anterior part of the

lower jaw. The configuration of the pattern of the melanocephalic

specimen purportedly from ‘Bona’ (NMB 2017) approximates the

last specimen very closely.

Body pattern very similar to that in M. mauritanicus but slightly

more variable, consisting of a vertebral series of dark spots 2—2.5

scales apart, eachl—3 or more scales wide; each of the contributing

scales is longditudinally bisected and emarginate. These are

accompanied by a dorsolateral series of spots or flecks of reduced

size. As in M. mauritanicus these spots are diagonally connected by

smaller series which are often obscured by imbrication: pale flecks

may accompany the vertebral or paravertebral spots. Ground colour

uniform or darkening at the median 7 or so rows and the 2nd and/or

3rd from the ventrals leaving a paler stripe the equivalent of two

scales wide. A melanocephalic and partially melanocephalic speci-

men, both from Algeria but lacking further locality data exhibited

diffuse pale interspaces between the vertebral spots.

Belly pattern likewise similar to that of M. mauritanicus, consist-

ing of squares, bars, occasionally chevrons of varying size but with

asomewhat greater tendency towards central disposition, n=3 (9.7%)

ef n=1 (1.2%).

GEOGRAPHIC VARIATION. None discernible.

98 E. WADE

ORR

Fig. 6 Dorsal patterns in Macroprotodon. Macroprotodon cucullatus. a. MNHN 849, Deserts of Western Algeria (holotype of Lycognathus textilis). b.

MNHN 1930.181, Djebel Ilaman, Hoggar, Algeria. Macroprotodon mauritanicus. e. MNHN 1994.3439 (syntype). d. BMNH 1920.1.20.3108, Batna,

Algeria. e. ZMB 51650, Tunis. Macroprotodon abubakeri. f. BMNH 1913.7.3.17, Oran, Algeria (holotype).

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100

AFFINITIES. Close to the preceding species with which it shares the

frequent irregularities in dorsal scalation and identical body pattern.

In scale characters (supralabial-parietal contact, ventral, subcaudal

and infralabial counts, points of reduction) it appears intermediate

between M. mauritanicus and M. cucullatus . However, the con-

dition of the head and body patterns suggest intermediacy between

M. mauritanicus and M. brevis.

DISTRIBUTION. Northwestern Algeria from the coast towards the

Hauts Plateaux at altitudes below 1,000 metres, Extends eastwards

as far as Algiers, westwards to northeastern Morocco as far as

Melilla (35°19'N, 2°57'W). Specimens having been taken from

localities as far south as El Kreider (34°09'N, 0°04'E) and Laghouat

(33°48'N, 2°53'E). With respect to the natural and bioclimatic range

of the species the last named locality is unexpected and may be

erroneous: it is possible that the specimen (MNHN 1899.273 pre-

sented by Secques) could have originated from further north.

LOCALITIES

SPAIN (North African possessions) — Melilla (35°19'N, 2°57'W); Zulueta,

1909 MNCN 1795.

MOROCCO - Ras el Ma (35°08'N, 4°26'W); Bons & Geniez, 1996:

Berkane (34°56'N, 2°40'W); Bons, 1960; 1967; Bons & Geniez, 1996

MNHN 1912.221: Guenfouda (34°29'N, 2°03'W) 19 km S (photographs +

exuviate); D. Donaire in litt.. Taforalt (Beni Snassene) (34°49'N, 2°14'W):

Werner, 1931; Bons & Geniez, 1996 MCZ 29920.

ALGERIA - no locality BMNH 1931.2.4.23; MNHN 3731; MHNG

524.59: Sebdou (34°38'N, 1°20'W); Doumergue, 1901: Beni Snous=Khemis

(34°38'N, 1°35'W); Doumergue, 1901: Arlal=Aghlal (35°12'N, 1°04'W);

Doumergue, 1901: Nemours=Ghazaouet (35°06'N, 1°51'W); Busack &

McCoy, 1990 MCZ 144369-70: Oued Sefioun (34°59'N, 0°07'W);

Doumergue, 1901: Saint Leu=Bettioua (35°48'N, 0°16'W); Doumergue,

1901: Beni Saf (35°18'N, 1°23'W); Doumergue, 1901: Mostaganem

(35°56'N, O°OS'E); Strauch, 1862: Mascara (35°24'N, 0°08'E); Doum-

ergue,1901: Djebel Mourdjadjo (35°40'N, 0°45'W); Werner, 1929; 193;

Busack & McCoy, 1990 MCZ 27502 (paratype): Iles Habibas (35°44'N,

1°08'W); Doumergue, 1901: Oran (35°43'N, 0°38'W); Doumergue, 1901;

Werner, 1909; Busack & McCoy, 1990 BMNH 1913. 7.3.14 (holotype);

MNHN 3734 (paratype); Oran, Battérie Espagnole; Werner, 1931; Busack

& McCoy, 1990 MCZ 29919 (paratype): Arcole=Bir el Djir (35°43'N,

0°34'W); Sochurek, 1956: Es-Sénia (35°39'N, 0°38'W) Busack & McCoy,

1990 FMNH 42840 (paratype): El Kreider (34°9'N, 0°04'E): Werner, 1929;

Busack & McCoy, 1990 MCZ 27500: Chellala=Ksar Chellala (35°13'N,

2°41'E) MHNG 1379.85-86, 1379. 88-89: Miliana (36°19'N, 2°14'E);

Strauch, 1862: Hammam Righa (36°23N, 2°24'E) ZMH RO4327: Algiers

(36°47'N, 3°03'E); Wade, 1988; Busack & McCoy, 1990 BMNH 53.2.4.23,

MNHN 3732, ANSP 3486, RMNH 212a—b, ZMH RO4322, ZSMH 2095/0,

NMB 9401: 100 km W EW 91.1: Laghouat (33°48'N, 2°53'E) MNHN

1899.273: ‘Bona’= ‘Annaba’(36°54'N, 7°46'E); Hediger, 1935 NMB 2017.

MHNG 1214.40 Chellala=Ksar Chellala (35°13'N, 2°41'E) has been identi-

fied as Macroprotodon brevis. For NMB 2015 (Oran) see under

Macroprotodon mauritanicus.

“LEVANT?” — MNHN 3736a.

KEY TO THE SPECIES

1 Usually lO }scaleimow sratenni cline chy eens cee eee nee eee 2

(2Wor23\scalemowsjatmmidbodiy eeceescceeecsseeceesteeeeeees M. brevis )

ine)

Body with ‘fextilis’ pattern; maxillary teeth 4—5 in series preceeding the

fangs; ‘pale’ collar usually indistinct or absent ............. M. cucullatus

Body with undifferentiated or ‘taeniatus’ pattern; 3 or 4 maxillary teeth

in series preceeding the fangs; ‘pale’collar nearly always present .... 3

3. Three maxillary teeth in series preceeding the fangs; nuchal collar

divided; postorbital streak often short... M. mauritanicus

E. WADE

Four maxillary teeth in series preceeding the fangs; nuchal collar entire

in majority; postorbital streak never short ..............0000-+- M. abubakeri

DISCUSSION

The distribution pattern of the forms of Macroprotodon cucullatus

as understood by Wade (1988) (Fig. 1b) followed naturally that of

Bons (1967) (Fig. la). However, that conceived by Busack &

McCoy (1990) (Fig. 1c), purportedly in accord with bioclimatic

parameters was probably the result of dismsissal of certain characters

and application of heavier weighting in others, notably that of

infralabial counts. With the exception of midbody counts characters

states used were shared at least in part by most of the taxa. The

diagnoses of Busack & McCoy (op. cit. pp 268-271) do not ad-

equately differentiate the subspecies. Their conclusions, insofar as

Morocco is concerned, have been accepted by Fahd & Plegezuelos

(1992) but rejected by Bons & Geniez (1998): Schleich et al. (1996)

present both conceptions. The results of this analysis supplement

that of Wade (1988) and resolve the uncertain status of many of the

populations from areas from which hitherto no material had been

seen by the author. The subspecies cucullatus, brevis, mauritanicus,

are raised to full species, the population from northwestern Algeria

and extreme northeastern Morocco (mauritanicus auct.), is recog-

nized as a new distinct species, abubakeri and the forms textilis and

ibericus are for the present retained as populations (or varieties) of

cucullatus and brevis respectively (Fig. 10).

Of the species of Macroprotodon, M. mauritanicus presented the

greatest areas of conflict of opinions in respect of its distribution and

composition. This is due in part to different interpretations of some

of the character states.

The position on the neck at which the count reduces from 21 to 19

is variable in all of the taxa (Fig. 8). The variability is most

10 Macroprotodon cucullatus

Macroprotodon abubakeri

art re ie Ea

li ee

i Macroprotodon mauritanicus

fo)

ee SN Eee ee

T T rT I i j T T T T T T T Lt 1

<) 10

percentage of ventral number

Fig. 8 Histogram showing anterior reductions (nuchal) from 21 to 19

scale rows. Percentage of ventral number (ordinate); number of

specimens (abscissa). Open bars = males, solid bars = females.

101

REVIEW OF ALGERIAN MACROPROTODON

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a am

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fo)

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102

12" VIR 18° 24° [30° “an

Ba . cucullatus Pease

M. cucullatus “textilis”

M. brevis

M. brevis “ibericus”

MG ™. mauritanicus

M. abubakeri

Fig. 10 The author’s concept of the distribution of the species of Macroprotodon.

pronounced in M. mauritanicus the reduction occuring at the 3rd to

the 16th ventral averaging at about the 10th (5.4%). M. abubakeri

exhibits a similar condition. The range in the position of the reduc-

tion for M. cucullatus in the region under investigation is narrower,

being from the 3rd to the 11th ventral and averaging at around the 7th

ventral (3.7%). For the Libyan populations and those further east the

average positions of reduction occur progressively closer to the head

(Wade, pers. obs.) suggesting that at least between these populations

the differences are clinal.

The differences in the position of reduction from 19 to 17 anterior

to the vent between the taxa, however are much sharper (Fig. 9). In

M. mauritanicus the position ranges from zero (i.e. no reduction) to

38 ventrals distance from the vent (x =10.5%), the majority exhibit-

ing positions of reduction much less distant. The range is widest in

M. abubakeri from the 3 to the 54 ventrals and the positions more

evenly distributed (x=15.7%). The positions of reduction are the

most distant from the vent in M. cucullatus ranging from 25 to 71

ventrals, the most frequent being from 40 to 47 (kx=28.6%).

Characters such as labial-parietal contact dismissed by Wade

(1988), Busack & McCoy (1990) and infralabial counts overlooked

by Wade were found to be exceptionally valuable in the resolution of

the mauritanicus problem. The sample of Busack & McCoy, 1990

(Fig. 1c) of ‘M. c. mauritanicus’ (n=73) is a heterogeneous assem-

blage which includes material from the Balearics (n=8), northwestern

Algeria =M. abubakeri (n=9) and that which is assigned in this work

to M. cucullatus (n=33): only twenty two of the specimens from

northern Algeria and northern Tunisia are referrable to M.

mauritanicus. Populations here recognised as M. cucullatus (n=3)

and M. abubakeri (n=2) where they extended into Morocco were

assigned to M. c. brevis. These authors found supralabial-parietal

contact in 40/47% (L/R) of ‘M. c. mauritanicus’. This character in

the present study was found to sharply differentiate M. mauritanicus

from M. cucullatus . The species showed no intermediacy in any of

the diagnostic character states where they occur at Biskra, the only

known point of contact. The scatter diagram (Fig. 11) also shows the

species to be well differentiated. Two specimens of M. mauritanicus,

SMF 20169 (closely resembling syntype MNHN 1994.2340, Fig.

4c.) and MHNG 2031.98 showed, in addition to undifferentiated

body patterns and the state of 6+3+II maxillary teeth, 6+3 infralabials

in conjunction with good supralabial-parietal contact. Two black

headed individuals of M. cucullatus MNCN 1802 (Fig. 3d) and one

originally from the Museo Civico di Storia Naturale di Milano (Jan

& Sordelli, 1966 Livr. 19e, Pl. I, Fig. 3A) shared the following

states: a pronounced ‘fextilis’ pattern, 6+4 infralabials and

supralabials separated widely from the parietals: the former pos-

sessed 6+4+II maxillary teeth as did presumably also the latter (lost

due to destruction of Jan’s collection during the 2nd World War, Dr

M. Podesta, in litt., 1998). Busack & McCoy (1990) found 23%

(n=16) of ‘M. c. mauritanicus’ to possess ‘entirely’ black heads.

Melanocephalism is rare in M. mauritanicus but common in M.

cucullatus (Table 2).

Three specimens of M. mauritanicus from a sample of 83 were

found to possess an entire nuchal collar, the common state in M.

abubakeri, of which two (BMNH 59.3.29.17 Algiers and NMB

2016 Médéa) originated in boundary zones between the two species.

However both possessed the states of short postorbital streak and

less than 4 maxillary teeth in the series preceding the fang; indeed

the Médéa specimen exhibited 6+2+II on both sides. In four other

specimens from Algiers, NMB 2422, MNHN 3735, ANSP 34847

and 34848 the nuchal collar was divided, supralabial-parietal con-

tact was pronounced but the infralabial state was equivocal (6+3

n=3, 6+4 n=2). By contrast in the nine specimens of M. abubakeri

from the same locality the nuchal collar was entire in all except

MNHN 3732 in wich it was divided: all possessed 6+4 infralabials

(6+5 on one side in one individual)whereas the supralabial-parietal

condition was equivocal (contact n=3, non-contact n=4 contact one

side n=2).

The evidence presented above shows that M. mauritanicus and M.

abubakeri are distinct taxonomic entities and behave as good species

without any evidence of intergradation: both being distinguishable

REVIEW OF ALGERIAN MACROPROTODON

subcaudals

103

ventrals

Fig. 11 Scatter diagram showing intraspecific variation for ventrals (ordinate) and subcaudals (abscissa) in Macroprotodon. Open symbols=males, closed

symbols=females, symbols enclosing stars refer to type specimens. Symbols as in Fig. 7.

from the M. cucullatus populations in the south, and M. abubakeri

from M. brevis in the west. Almost inevitably there are a few

exceptional cases where the odd individual may be divergent in one

of the character states.

Aside from the nuances of the nebulous concept of ‘general

appearance’ the characters by which M. abubakeri can be distin-

guished from M. brevis (excluding the Iberian populations) are the

lower number of dorsals and the usual state of the postorbital streak

meeting the ‘arms’ of the V-mark around the last supralabial. M.

mauritanicus exhibits the greatest degree of homogeneity in its

character states and the distinction from M. abubakeri is similarly

clear cut. Compared with M. mauritanicus, M. abubakeri is some-

what less homogeneous in its character states. In five specimens the

dorsal count rose to 21 in stretches from 2 to 24 ventral scales. Two

of these from the extreme northeast of Morocco, MNHN 1912.221

Berkane and MCZ 29920 Taforalt (Beni Snasséne Mountains) whilst

exhibiting counts of 21 in maximum stretches of 24 and 21 ventrals

respectively conformed in all other respects to the diagnostic states

of M. abubakeri. The single specimen of this species out of a total of

four from Melilla the boundary zone, MNCN 1795, makes no

approach to M. brevis in any of its characters: it has an uninterrupted

run of 19 rows, 344 infralabials and no contact between the

supralabials and parietals. The other three, MNCN 1783, 1796, 1797

all have 21 rows, good supralabial-parietal contact and, except for

the last in which the infralabials number 7+4, the count is 6+3. Bons

& Geniez (1996) *...examined numerous specimens from eastern

Morocco all of which possessed 19 rows’. The count anterior to the

vent is normally 17 but may decrease further: in one specimen which

originated from Dj. Bou Keltoum (south of Guenfouda) the count

actually fell to 15 for a distance of 2 ventrals before settling at 16. In

three individuals (including the holotype and one of the paratypes)

it decreased to 16. Other characters the states of which may resemble

those of neighbouring taxa such as infralabial counts and nuchal

collar are scattered within the body of the populations.

M. cucullatus and M. brevis (including populations of ibericus)

have wide geographic ranges and possess intraspecific variations

which often lack consistencey from one locality to another. Al-

though M. cucullatus shows little of the variability in dorsal counts

found in M. brevis (19 as opposed to 19-25) it exhibits a polymor-

phism in head pattern at least as great, being unique in that a

significant number of its populations lack the ‘pale collar’. By

contrast M. abubakeri and more particularly M. mauritanicus are

restricted in their ranges showing considerably less intraspecific and

virtually no geographic variation when their ranges are compared to

those of similar dimensions of populations of M. cucullatus .

Most of the character states which could be said to define M. c.

ibericus, e.g. Supralabial-parietal contact, 3 posterior infralabials,

complete nuchal collar, occasional melanocephalism apply also to

some populations of M. brevis occuring east of Tangiers including

those parapatric with M. abubakeri (Wade, unpublished).

Pleguezuelos (1998) noted melanocephalism in examples bordering

the strait of Gibraltar. The variations are greatest in M. brevis of

which the Tangiers-Melilla populations form only a part. The dis-

tinction between these ibericus-like forms from M. abubakeri is

readily made but from M. brevis elsewhere in Morocco it is not easy

to make as there is considerable overlapping of character states over

wide areas. Nineteen midbody scale rows has been found to occur in

some of the Iberian populations and very exceptionally in the

Moroccan but this state has not been found in any of the populations

from Tangiers to Melilla. Busack & McCoy (1990) presented a case

for M. c. brevis giving rise to M. c. ibericus in Iberia largely on

evidence of electrophoresis. It is regrettable that they restricted their

samples for that analysis to populations from either side of the Strait

Fig. 12 Scatter diagram showing the position of the species of

E. WADE

104

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REVIEW OF ALGERIAN MACROPROTODON

of Gibraltar and not from the other regions in which they altered the

ranges. Definition of the states that constitute M. c. ibericus and

determination of its range requires further investigation.

There is to a large extent correlation between the distribution of

the species and bioclimatic parameters. The xerothermomediterra-

nean demarcation line (x=150—200) separating the Mediterranean

from the arid-Saharan regions is a convoluted band of variable width

which commences at the Atlantic and extends eastwards to the

Tunisian coast (Fig. 7). The line becomes particularly narrow as it

reaches the Moulouya valley to the west of which lie the Anti Atlas,

an area of moderate humidity (mesomediterranean) where M. brevis

is found: to the east of it stretches the arid region along the Moulouya

valley to less tan 50 kilometres from the Mediterranean coast. From

thence the line extends along the the Hauts Plateau for some 400

kilometres whereupon it reverses to almost as far as Ain Sefra only

to again reverse and continue along the Atlas Saharien from whence

it extends in an arc across the Monts du Hodna to the southern part

of the Massif de I’ Aurés before proceeding eastwards to the coast.

M. cucullatus inhabits largely the arid or Saharan regions to the

south of the line and the abubakeri-mauritanicus assemblages the

Mediterranean to the north of it (Fig. 7).

M. abubakeri could be viewed as an eastern mesomediterranean-

thermomediterranean extension of the northern Moroccan

populations of M. brevis which it resembles at least as closely as it

does to M. mauritanicus. Further east M. abubakeri is replaced by

M. mauritanicus which is most densly concentrated in the

mesomediterranean but extends into the more humid submediterra-

nean subregions. The two northern forms extend southward, although

with considerably less frequency, into the xerothermomediterranean

and enter the desertic subregions (Fig. 12). M. abubakeri, unlike M.

cucullatus or M. mauritanicus which do not eschew the high

ground, is restricted to land below 1,000 metres, at least in Algeria.

The localities Taforalt, Djebel Bou Keltoum (Morocco), Khemis

and Sebdou (Algeria) although appearing on the map (Fig. 7) in the

shaded area showing land above 1,000 metres do in fact lie below

that altitude. The scale of the map is not large enough for the smaller

valleys and similar low lying areas to be shown.

Individual specimens that do not conform to the taxon or popu-

lation from which they are supposed to have originated occasionally

appear: this may be due to error of attribution or more likely as a

result of human introduction. Four specimens of M. brevis three of

which possess 23 midbody scale rows from ‘Algeria’ (NMW

25840.5), Bordj Bou Arréridj (MHNG 1379.68) and Tunis (NMW

19193.3) —1.e. well within the range of M. mauritanicus — agree with

material from the Essaouira-Marrakesh region; the other with 21

uninterrupted rows from Ksar Challala (MHNG 1214.40) most

closely resembles the northern Moroccan population. There is no

evidence of intergradation but instead there is some suggestion of

character displacement thus arguing against expanding the descrip-

tion of M. mauritanicus to accomodate the additional states: rather,

it provides further support for the recognition of the taxa at specific

level.

The melanocephalic specimen mentioned by Hediger (1935) p.

24 from ‘Bona’ NMB 2017 with four posterior infralabials and four

teeth preceding the fangs, presents a head configuration resembling

so closely that from Djebel Mourdjadjo that its discovery in the

eastern sector of Algeria is most likely to be due to accidental

introduction. This individual is listed in the current NMB catalogue

as one of eight specimens collected by Hagenmiiller Miiller(1890 p.

692) lists only seven specimens presented by that collector from

around Annaba. Although noteworthy particulars are recorded no

individual is singled out as being black headed. A further specimen,

NMB 2015 also listed by Miiller 1878: 567 & 666 under ‘Algerien

105

und Aegypten’, the provenance of which is stated in the actual

catalogue to be Oran. However, possession of sutural labial-parietal

contact, 6+3 infralabials and 3 posterior maxillary teeth suggests

that it originated from further east: it is identical to M. mauritanicus.

It is of some significance that both individuals were collected from

long established seaports rather than from inland.

A specimen (SMF 20170) collected in ‘southern Spain’ which is

identical to material from Egypt is surely the result of an accidental

introduction. Macroprotodon not infrequently occurs near human

habitations. Some of the specimens from Médjana were rescued,

both as adults and particularly as hatchlings, from domestic chick-

ens. Secretive, adventitious little snakes abroad at dusk or early

morning hiding in human artefacts, could easily be transported by

accident.

ACKNOWLEDGEMENTS. [am indebted to Drs E. N. Arnold and C. McCarthy

for permitting me to re-examine the Macroprotodon collection in the NHM

and for numerous other courtesies. For the loan and making available

material for examination I thank Drs V. Mahnert and B. Schatti (MHNG), E.

Stéckli (NMB), J. Rosado (MCZ), G. Zug (USNM), J. Cadle (ANSP), E. J.

Censky (CM), R. L. Humphreys (UCM), R. Giinther(ZMB), G.E. Gonzales

(MNCN), M. Hoogmoed (RMNH), I. Ineich (MNHN), W. Bohme (ZFMK),

M. Laudahn (SMF), U. Gruber (ZSMH), H. Koepecke (ZMH), A. Resetar

(FMNH), J. Vindum (CAS), Prof. F. Tiedemann (NMW). I thank Dr G. Vogel

for the donation of a specimen and Mr D. Donaire for photographs and an

exuviate. Dr G. Underwood and Mr B. Hughes provided constructive criti-

cism and Mr J. Pether provided advice and assistance during the course of this

work.

I thank my senior colleagues at Middlesex University, the late Professor J.

Lansdown who authorised the study and particularly Professor lan McLaren

who approved leave for travel and provided invaluable help. The visits to

Algeria were financed by grants RO31/1109 and RO31/6180. Colin Rattray

and Phil Wilson took over my responsibilities during my absence.

I owe a debt of gratitude to Dr Nebbache Mounir and M. Médani Kerim for

providing facilities and assistance whilst I was in Algeria and Dr Mohamed

Bey and M. Nebbache Djemai for hospitality. Messrs Aboubakeur Sid-

Ahmed, Nebbache Riyadh, Ouali Nour ed-Dine and O. Abbas provided

hospitality and companionship in the field. To all | extend warmest thanks.

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