EM lose -/ —» Zoology Series SZ THE NATURAL HISTORY MUSEUM VOLUME 67 NUMBER1 28 JUNE 2001 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology Prof P.S. Rainbow Editor of Bulletin: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Email: intercept @ andover.co.uk Internet: http://www.intercept.co.uk Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) © The Natural History Museum, 2001 Zoology Series ISSN 0968-0470 Vol. 67, No. 1, pp. 1-107 The Natural History Museum Cromwell Road London SW7 5BD Issued 28th June 2001 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset 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 TNR OUCRU CULO Wiieeteas cca ecaeen ee» faierc Seve vacesvaedeccotapa- ve sesh raa-coesvanasav'es = = = Z ZA B 2 Fos 2 » 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 34 i JL 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 os Sy Yj 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 Ae a if ms s se" so 50" S wy Weert mh \ FA fens [7 iil pte G ee SS S SSE REO aN Ss SEER IT SON 5 \\ SSS ORK LTE —< ~ a IO, SSS ee LLG SSS 52a 3 SRS SSSES SOLE. SSS << = >> SSAA 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 oN HHS y +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 35 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 36 R. BOTTGER-SCHNACK Cw SL LE KK SSSS: EE 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) (c) numerous 29 and dd (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: 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. 39 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. 4] 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. 44 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 bs | he 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. 45 46 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 \ Q “3 a: se ~ N35 N: we rs 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 48 IK WERK SS pee SOY Zi 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 <= ‘ O 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. 50 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 51 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 2 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 58 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. 54 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). (c) 1 Qdissected on slides, 2 22 in alcohol; 1 ¢ dissected on 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). (c) 2 22(1 ovigerous), 2 dd in alcohol, (RBS). 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. Ee =. = SEVEN SPECIES OF RED SEA ONCAEA 2" LAID ee LEE: sea a LEE a —— 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. 55 56 SSS A7 ay, : L SSS ¢ > a : =. v 2 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 (%) p2 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 P4 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. 3) (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) 58 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). (c) 1 @partly dissected (maxilliped and maxilla on slides), 1 2 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). (c) 2 29,3 36 in alcohol (RBS). (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). le == qi em,. « ———— SEVEN SPECIES OF RED SEA ONCAEA (c) 1 do (from mating pair) dissected on 9 slides, | 2(ovigerous, 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. 59 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/ ZL 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. 61 R. BOTTGER-SCHNACK 62 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. 63 64 ~ 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 65 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). (c) 2 22 dissected on slides, 1 Qin alcohol; 1 3 dissected on 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). (c) 2 dé in alcohol (RBS). (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 (4 wa 66 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. 67 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 GZ CDEIJM age Ue. = Se meres renwal <7 SS SSS SS=s SSS =y SESS & < SESS are a 68 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 69 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. 70 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). (c) 2 22,3 dé dissected on slides; 2 29, 1 d 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 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). 71 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. 72 R. BOTTGER-SCHNACK eyo; G ° 7 ES ) ‘ ° Chun , +100 “ER fe” Bp lye oS 4 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. 73 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 74 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. 76 R. BOTTGER-SCHNACK = ZZ S FES SS SS - 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 77 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 78 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). (c) additional paratypes: 1 d dissected on 10 slides, 1 @ dis- 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). (c) 1 paratype @ dissected on 10 slides (RBS). (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. W 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. 81 < os a es RS ee = Za So 4 (B) P2, anterior; (C) P3, anterior; (D) P4, anterior. <= Ae; 3S << Li ES ZZILEZ x , . Fig. 31 Oncaea paraclevei sp. nov., female (Red Sea) (A) P1, anterior; SEVEN SPECIES OF RED SEA ONCAEA 82 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. REFERENCES Almeida Prado-Por, M.S. 1983. The diversity and dynamics of Calanoida (Copepoda) in the northern Gulf of Elat (Aqaba), Red Sea. Oceanologica Acta, 6: 139-145. Beckmann, W. 1996. Der Einfluf der grofrdumigen Wasseraustauschvorgdnge auf den Zooplanktonbestand des Roten Meeres und sein trophisches Gefiige Disserta- tion Fachbereich Biologie, Universitat Hamburg, 167pp. Bersano, J.G.F and Boxshall, G.A. 1994. 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Copepods of the Woods Hole Region Massachusetts. Bulletin of the United States National Museum, 158: 1-623. Wishner, K.F. 1979. The biomass and ecology of the deep-sea benthopelagic (near- bottom) plankton. Ph.D. Dissertation, University of California, San Diego, 144 pp. Bull. nat. Hist. Mus. Lond. (Zool.) 67(1): 85—107 = Issued 28 June 2001 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 © The Natural History Museum, 2001 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 86 T =n 18° 24° ie HM. c. cucullatus 2) M. c. brevis Y aa) q oS 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 (4 Fig. 1 Concepts of the distribution of Macroprotodon of various authors. \ \ Se \ 3 NY \ \ 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 87 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 88 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 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 89 = § | / \ | 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 90 “(WILIG=9[89G) “"RLIOSTY “VXYSIG ‘7081 NONI ‘P eHes|y “eayag uly -1OSL7T ZOIN “9 “RLIOS|V “‘sInod [piog ‘6P 1 L761 NHNW 4 “(S1pxay SNYIDUSOIKT jo adAjojoy) RLIOs|V UIOISIMA JO S}IOSOC] ‘68 NHNW “® *SNID]JNINI uopojoido.ovy ul usjoyjed oadeu pure peoH € “Sly i Qe OOO Ce en S OSVernoen Oe) 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 9] 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 wy 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 o8 + 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 94 ‘(utUIG=9]/k9§) “‘eLOSTY ‘vuelpslA ‘SC T661'MA “P “(eLAsTYy ‘sadAuds = 3-8) OP€E7'P661 NHNW "9 6£€7'P661 NHNW “4 ‘ZLIZ NHN ‘8 “s7121UDAnDW Uopojosdosovyy Ut UIE adeu pue peoH =p ‘Sly ae ) ) i ORR Kee te ON ) SNe. K) os ieeeee oe 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 95 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 96 MOY eLIOSTY “TELE NHN *P “(MUI g=a]k9S) “Ay[eD0] astoaId ‘SISIB|V “1076 GINN “9 “(eHe8]Yy ‘ueIC “q-e) (edAiered) pe7E NHN “4 ‘(2dM 004) HIE LE 161 HNN “8 /2¥DqGnqQv uopojordosoyy ut usayed adeu pur peop s “Bq 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 97 (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). ‘(uosvas Arp ayy ut sXep Asp AqyeorsojorsAyd Jo soquinu) XopUl STWIAYJOIOX=K *(9]LOS IWS dy} 0}) UOTSOI IeSS0H oy} :JOSuT ‘sotoads ay] JO sasuvs oy) Ul JUAUINGe Jo saovid puke santfeso] adA] 0} JaJo1 soweU ddeId :s}10da1 sinyesO}T] ‘sjoquuAs uado ‘pourwexe susuttdads Uaseida1 sjoquiAs pr]Os ‘vopojoido.sopP Jo uoNNgINsIq LL “BLY 99 r T Ww OO€ 002 sauepunog |eonijod 00c=* OSL=* Supp] 12galq | Ni S81]8W 000‘ JAA0 pur] Sy RS Ne —~, : a oy R 4 y See gS | OO SEN ( aS Sanu a SIOIB]V Vas NVHYNVHYHXALIGCAWN tuayoqnqn uopojosdosnvpy @ snoiupjlinou uopojosdo.IvY snjojynond uopojoidoszsvy @ av. ies s1daig uopojosdosnvy W a. : i : 0 M\. SHEMEEH TT die Ze S S i S & a © = = Z ps Z na Oo Bs z ta Oo Ss = > a 4 100 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 = oO art re ie Ea li ee i Macroprotodon mauritanicus a fo) ee SN Eee ee oa = T T rT I i j T T T T T T T Lt 1 <) 10 15 percentage of ventral number 0 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 ie bd bb ALEC ee ed GEEEELESEERETR ERASE a am es aan | | oy _ 7. Oe Se Se ee ee ea ee a ee "Q “SL JO] se sanjeA pur sjoquikg *sMOI g]ROS /] 0} 6] WOIZ SUOMONpal JOLA\sod SurMoys wIeISO}STE] «6 “SE JOquINU [eNUIA JO asv}UdoIEd Se oe Se Ov | [ rm + wo SNI1UDJ1ANDU ~UOpoJOAdOsIDIY (fo) = = LJ wo 14aypqnqn uopojosdosavpy [o) = a i gee Re aes ae il y ceeg generat is i a Lk ies Ss LP — Tin 'aan elie aon lihaenleaian lea a lS ee lags Senate pein cnc wo fo) = snjpjjnInd uopojosdosavpy J 102 12" VIR 18° 24° [30° “an Wi Ba . cucullatus Pease M. cucullatus “textilis” M. brevis M. brevis “ibericus” MG ™. mauritanicus M. abubakeri 2] 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 *suoutoods Jo saquinu Juasosdar sjoquIAs UIYIIM SANS °/ “BLY UI se S[oquIAg “(essiosqe) apnyne] Seaisap ‘(9}eUIPIO) SooIpUT STULIOYIOIOX :sUOISoIqNs puL sUOIS9I NRUNT|DOIG a4) 0} UONEIAI UT uopojoidoiovy T Af T T ib al a F al T il T T T 7] 00 0sz 002 OS| 00! 0S OF [O Ar ir oc + 6 ~ a2 - - - of 4 1Ox oO : ld be - = 1% B Ps one eats a 98 29 oL€ — LE 00 osz 002 OSL 00! 0S OF [es [oa =< | L ll | eile “‘eseq “‘aesspqns ‘PSWOLWJSUJO1EX “PSWOUWISY | “pswoss|/| “*pesWgns NVYVHVS-CIYV NVANVYHALIGCAN 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. REFERENCES Anderson, J. 1892. On a small collection of mammals, reptiles and batrachians from Barbary. Proceedings of the Zoological Society, London (1): 3-24. Angel, F. 1944b, Contribution al’ Etude de la Faune herpétologique du Sahara Central. Bulletin du Muséum Nationald Histoire naturelle, Paris (2) 16: 418-419. & H. L’Hote. 1938. 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Preference is given to original contributions in English whose contents are based on the Collections, or the description of speci- mens which are being donated to enhance them. Acceptance of manuscripts is at the discretion of the Editor, on the understanding that they have not been submitted or published elsewhere and become the copyright of the Trustees of the Natural History Mu- seum. All submissions will be reviewed by at least two referees. Submission of manuscripts. Initially three clear, complete cop- ies should be submitted in the style and format of the Bulletin. The text must be typed double-spaced throughout, including references, tables and legends to figures, on one side of A4 paper with 2.5 cm margins. All pages should be numbered consecutively, beginning with the title page as p. 1. SI units should be used where appropriate. Whenever possible a copy of the text, once the paper has been accepted, should also be provided on floppy disc (see below). 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In: Simpson, T.L. & Volcani, B.E. (eds) Silicon and Siliceous Structures in Biological Systems. Springer- Verlag, New York. Tables. Each table should be typed on a separate sheet designed to extend across a single or double column width of a Journal page. It should have a brief specific title, be self-explanatory and be supple- mentary to the text. Limited space in the Journal means that only modest listing of primary data may be accepted. Lengthy material, such as non-essential locality lists, tables of measurements or details of mathematical derivations should be deposited in the Biological Data Collection of the Department of Library Services, The Natural History Museum, and reference should be made to them in the text. Illustrations DRAWINGS — Figures should be designed to go across single (84 mm wide) or double (174 mm wide) column width of the Journal page, type area 235 x 174 mm. Drawings should be in black on white stiff card with a line weight and lettering suitable for the same reduction throughout, ideally not more than 40%. After reduction the smallest lettering should be not less than 10 pt (3 mm). Tracing paper should ideally be avoided because of the possibility of shadows when scanned. All artwork must have bulletin, author and figure number included, outside of the image area, and must be free of pencil, glue or tape marks. PHOTOGRAPHS — All photographs should be prepared to the final size of reproduction, mounted upon stiff card and labelled with press-on lettering (eg Letraset). They can be mounted on white or black background; a black background must be evenly black all over; any background must be free of all pencil and glue marks within the image area. All figures should be numbered consecutively as a single series. Legends, brief and precise, must indicate scale and explain symbols and letters. Photos, when components of figure- plates should be abutted, trimmed as regular rectangles or close trimmed up to edge of specimen. Joins etc. can be removed at the scanning stage but at extra cost. Cropping instructions, if any, should be indicated on an overlay or marked on a photocopy of the figure. SIZE — Maximum size of artwork for use of flatbed scanners is A3. Larger artwork has to be reduced photographically prior to scan- ning, therefore adding to expense. Symbols in text. Male and female symbols within the text should be flagged within curly brackets to enable setter to do a swift global search. Reprints. 25 reprints will be provided free of charge per paper. Orders for additional reprints can be submitted to the publisher on the form provided with the proofs. Later orders cannot be accepted. CONTENTS 1 Freshwater nematodes from Loch Ness, Scotland Part I. The orders Tylenchida Thorne, 1949 and Rhabditida Chitwood, 1933 (Nematoda, Secernentea) FR. Wanless and R. Hunter 25 Taxonomy of Oncaeidae (Copepoda, Poecilostomatoida) from the Red Sea. Il. Seven species of Oncaea s.str. R. Bottger-Schnack 85 Review of the False Smooth snake genus Macroprotodon (Serpentes, Colubridae) in Algeria with a description of a new species E. Wade Bulletin of The Natural History Museum ZOOLOGY SERIES Vol. 67, No. 1, June 2001