Observations on Amphora Species (Bacillariophyceae) in the British Museum (Natural History)

Observations on Amphora species (Bacillariophyceae) in the British Museum (Natural History). V. Some species from the subgenus Amphora

F.R. Schoeman and R.E.M. Archibald National Institute for Water Research, Council for Scientific and Industrial Research, Pretoria

Six Amphora species [A. affinis KOtzing, A. copulata Introduction (KOtzing) nov. comb., A. marina W. Smith, A. ova/is (KOtzing) This is the penultimate paper in a series on Amphora species KOtzing, A. proteus Gregory, A. robusta Gregory) belonging (Schoeman & Archibald 1986a, 1986b, 1986c, 1986d, 1986e) to the subgenus Amphora Cleve were observed on strewn slides in the British Museum (Natural History). Light of which the type specimens are kept in the British Museum microscope photographs of examples corresponding to (Natural History). We have dealt with these species in order their descriptions have been included. Comments on the to verify the identification of taxa recorded from southern authenticity of the material examined and the suitability of Africa_The purpose of these papers is also to provide helpful the observed specimens as types for the species are made. information to those who may not have the opportunity of S. Afr. J. Bot. 1986, 52: 425-437 examining the British Museum (Natural History) material. Six taxa from the subgenus Amphora Cleve are considered Ses Amphora-spesies [A. affinis KOtzing , A. copulata (KOtzing) nov. comb., A. marina W. Smith, A. ova/is here. Three are treated in the format adopted in earlier papers (KOtzing), KOtzing, A. proteus Gregory, A. robusta Gregory) (e.g. Schoeman & Archibald 1986a). The other three taxa wat tot die subgenus Amphora Cleve behoort is op form a group of closely related species, whose taxonomy and diatoomstrooimikroskoopplaatjies in die versameling van die nomenclature are more complex. This group comprises Am­ British Museum (Natural History) waargeneem. Ligmikro­ phora affinis, A. copulata and A. ova/is, which were all skoopfoto's van voorbeelde wat ooreenstem met hul originally described by Kiitzing (1833, 1844), but his concept beskrywings word ingesluit. Kommentaar word gelewer oor die egtheid van die ondersoekte materiaal en oor die of these taxa is confused. Our research shows that Kiitzing geskiktheid van die eksemplare as tipes van die (1833) originally distinguished two species, Frustulia ova/is and respektiewelike spesies. Frustulia copulata, which he subsequently united as Amphora S.-Afr. Tydskr. Plantk. 1986, 52: 425-437 ova/is (Kiitzing 1844). At the same time he (Kiitzing 1844) distinguished two more species, Amphora libyca Ehrenberg Keywords: Amphora, Bacillariophyceae, light microscopy, (1840) and Amphora ajjinis Kiitzing_ Later authorities (Van type material Heurck 1880- 85; Cleve 1895) recognized that these two species are conspecific, but treated them as a variety of A. ova/is_ However, it appears from the literature that there has been no re-investigation of the identity of F. copulata. Through the courtesy of the British Museum (Natural History) we obtained a recently prepared slide of an unnumbered sample in the Kiitzing collection, labelled as containing both F. ova/is and F. copulata. Having examined this slide we conclude that the differences between the two are sufficient to resurrect F. copulata as a valid species. On comparing examples of F. copulata with specimens on the type slide of A. ajjinis it was evident that the two species are identical. As a result some nomenclatural and taxonomic changes are proposed. These are discussed in detail in our treatment of the 'Amphora ova/is complex' (consult paragraph 4 below). Unfortunately we have not been able to examine Ehrenberg's type material of Amphora libyca (kept in Berlin) to confirm whether it is identical to A. ajjinis. Nevertheless, for the time being we accept the commonly held opinion that they are conspecific. F.R. Schoeman* and R.E.M. Archibald Materials National Institute for Water Research, Council for Scientific and Industrial Research, P.O. Box 395, Pretoria, 0001 Republic of Following the style of our previous papers in this series South Africa (Schoeman & Archibald 1986a), the slides examined in this *To whom correspondence should be addressed study are listed separately under the species or species complex involved_ In addition to the British Museum slides (indicated Accepted 14 April 1986 by the abbreviation BM prefixed to their serial number) we 426 S.-Afr. Tydskr. Plantk., 1986, 52(5)

examined two slides from Van Heurck's 'Types du Synopsis J.!m; dorsal striae near the centre 15-20 in 10 J.!ID, and at des Diatomees de Belgique' (prefixed by the abbreviation the poles 16-20 in 10 J.!ID; ventral striae 12-16 in 10 J.!ID. VHS) in the National Institute for Water Research (NIWR) slide collection. A corresponding slide may be found in the 4. The 'Amphora ova/is complex' Van Heurck collection in the British Museum (Natural Hist­ Rather than discussing each species individually, we will deal ory). In the text below (paragraph 6) we have used the symbol instead with a group of taxa which have close, but sometimes MF to denote Maltwood Finder coordinates used to locate confusing affinities. Different opinions on their taxonomic some specimens. status and nomenclature have produced a complex of species or varieties, whose taxonomy needs clarification. The group ObseiVations and Discussion comprises Amphora ova/is (Kiitzing) Kiitzing (1844, p.107, 1. Amphora affinis KUtzing = A. copulata (KUtzing) nov. pl.5, figs 35, 39), A. affinis Kiitzing (1844, p.107, pl.30, comb. fig.66), A . libyca Ehrenberg (1840, p.205), A. ova/is var. See comments under 'Amphora ova/is complex' (consult ajfinis (Kiitzing) Van Heurck (1880- 85, p.59, pl.l, fig .2), paragraph 4). and A. ova/is var. libyca (Ehrenberg) Cleve (1895, p.104). It is therefore appropriate that we should determine first how 2. Amphora copulata (Kutzing) nov. comb. many taxa are involved and then their correct nomenclature. See comments under 'Amphora ova/is complex' (consult As far as we are aware, the British Museum (Natural paragraph 4). History) does not have a slide designated as the type for A. ova/is, but there are several slides prepared from Kiitzing's 3. Amphora marina W. Smith herbarium material, marked as containing A. ova/is. However, W. Smith 1857, p.9, pl.l, fig.2. Cleve 1895, p.103. Van after examining these slides, we became confused as to what Heurck 1880-85, p.58, pl.l, fig.16. Kiitzing really called A. ova/is, since some slides contained forms fitting our present concept of A. ova/is (cf. Patrick and Slides examined: BM 1033 Biarritz, France, 1856. Coli. Reimer 1975, p.68, pl.l3, figs 1, 2), while others only had Greville. (Figures 1 - 7). examples fitting the description of A. affinis, and yet others VHS 101 (NIWR 6/101) 'Carrieghills' (? Corriegills), Arran, contained both species. Scotland. (Figures 8- 10). A. ova/is was first described by Kiitzing (1833, p.539, pl.l3, Notes: A. marina was described by W. Smith (1857, p.9, pl.l, fig.5) under the name Frustulia ova/is and was recorded as fig.2) but his illustration only depicts the species in girdle view. occurring 'Einzeln unter Frustulia subulata und acuminata etc. We have therefore no illustration of the valve to aid its identifi­ bei Weisenfels'. Referring to the descriptions of the latter two cation. Our identification of this species is based on slide BM species in the same publication, Kiitzing (1833, p.538 and 1033 from the Greville collection in the British Museum (Natu­ p.555) cited Decas IX of his 'Algarum aquae dulcis German­ ral History), which was prepared from material collected at icarum' (Kiitzing 1833- 36) as the depository for his type Biarritz in France, one of the two type localities cited by W. material (Habitat: Leucopetram = Weisenfels) of these two Smith in his diagnosis. The authenticity of this slide is further species, and hence also for A. ova/is. We therefore examined attested to by W. Smith's initials engraved on the glass. We two slides (BM 78085, BM 78086) made from Decas IX, can therefore regard it as the type slide for A. marina. In No.81 (F. subulata), and one (BM 78084) prepared from Figures 1-7 we have illustrated several examples of A. marina Decas IX, No.84 (F. acuminata) expecting to find A. ova/is. from this slide, which contained only frustules. These specimens This expectation was encouraged by an illustration of A. ova/is show the valve characteristics of the species reasonably well. from the Decas IX, No.84 material produced by Reimer (in In addition toW. Smith's slide, we also observed a number Patrick & Reimer 1975, pl.l3, fig .1). However, despite a very of examples of A. marina on Van Heurck's (1884- 87) slide careful search, we were unsuccessful and observed only a few no. 10 from 'Carrieghills', Arran (e.g. Figures 8-10). They specimens of A. ajfinis (mainly on Decas IX, No.84) and are identical to the Biarritz examples. some odd examples of Amphora pediculus (Kiitzing) Kiitzing Cleve (1895, p.103) doubted the validity of A. marina and (vide Schoeman & Archibald 1978). We were therefore still suggested that it may be a form of Amphora proteus. Accord­ uncertain as to what Kiitzing described as A. ova/is and which ing to our observations (see paragraph 5), the latter is a larger material could be used to typify this species . species with much coarser striation. In this respect a comment It was not until the British Museum (Natural History) sent by Gregory (1857, p.518) on the variability in striae density us a recently made strewn slide (BM 81032), prepared from of A. proteus is noteworthy. Gregory cited ca. 9 striae in 10!!m an early Kiitzing gathering kept in an unnumbered packet, for A. proteus, but remarked that 'there are very great varia­ that we felt we had a solution to the problem. Written on tions in this species, . . . . In some of the smaller specimens this packet in Kiitzing's own handwriting is a list of species the striae are at least twice as numerous as in some of the found in the material. This list includes Frustulia copulata larger, . . .. ' This raises the possibility that Gregory may have and Frustulia ova/is, both described and illustrated by Kiitzing observed specimens of A. marina, which according to W. (1833, pp. 541 and 539 respectively). Lower on this list an Smith (1857, p.9) is not infrequent on the British coast, but annotation, probably written later by Kiitzing, indicated that that Gregory considered them to fall within the A. proteus he had combined F. copulata and F. ova/is under the name range. W. Smith (op. cit.) pointed out the resemblance of A. Amphora ova/is (vide Kiitzing 1844, p.107). On examining marina to A. affinis ( = A . copulata, see our comments this slide (BM 81032) we did, indeed, find two Amphora taxa, below), but the two species can be readily distinguished on one corresponding with our present concept of A. ova/is (vide the basis of raphe structure and construction of the dorsal Patrick & Reimer 1975, p.68, pl.l3, figs 1, 2) and the other, central area (compare Figures 1-10 of A. marina with Figures which has a very distinct dorsal central area, agreeing with 25-53 representing A. copulata). A. affinis as depicted in Van Heurck (1880- 85, pl.l, fig.2). Dimensions of specimens examined: Length 24,0 - 46,5 J!m; Kiitzing (1833) illustrated only two Amphora species (as F. breadth of frustules 14,5-23,0 J!m; breadth of valves 7,0 - 8,5 ova/is and F. copu/ata) in his 'Synopsis Diatomearum'. The S. Afr. J. Bot., 1986, 52(5) 427

10pm

Figures 1-29 (1-7) A. marina W. Smith. BM 1033, Biarritz (type). (8 -10) A. marina W. Smith. Van Heurck Slide No. 101 (NIWR 6/101), Carrieghills, Arran. (11-13) A. copulata (Klitz.) nov. comb. BM 81032, Kiitzing material (as F. copulata). (14, 15) A. copulata. BM 18933, Dieskau (as A. ova/is on slide and F. copulata on packet). (16-21) A. copulata. BM 78084, Kiitzing Alg. Aq. dulc. Germ. Decas IX. (22, 23) A . copulata. BM 18936, Wiirzburg (as A. ova/is). (24) A. copulata. BM 78086, Weisenfels (as F. ova/is.) (25- 28) A. copulata. BM 18938, Falaise (as A . ajjinis.) (29) A. copulata. BM 18934, Tennstiidt (as A. ova/is 13 minor). Figures I, 22, 26, 27a, 28: oblique bright field illuminaton (O.B.F. Ilium.). Figures 2- lOa, II- 15, 18 -21, 23, 24, 29 : bright field illumination (B.F. Ilium.). Figures lOb, 25, 27b: phase contrast illumination (P.C. Ilium.). All figures x 1500. 428 S.-Afr. Tydskr. Plantk., 1986, 52(5)

two taxa observed on slide BM 81032 could be correlated with correlate with the illustrations of F. ova/is (Kutzing 1833, the original illustrations of F. ova/is and F. copulata, using pl.l3, fig .5). the presence or absence of the central nodule as the distinctive We may thus conclude that at the time Kutzing (1833) wrote feature. In doing so, the A. affinis forms conformed to the his 'Synopsis Diatomearum' he identified two Amphora drawings of F. copulata (Kutzing 1833, pl.l3, fig.6) having a species, the one being A. ova/is (F. ova/is) as we know it clearly defined central area, and the A. ova/is forms therefore today, and the second being the taxon he called F. copulata.

36 37 38a 38b ' 39

10pm 35b

40 41a 41b 42 43 51 44

l!) 53

Figures 30-56 (30- 34) A . copulata (Kiitz.) nov. comb. Van Heurck Slide No.I (BM 26312), Falaise (mixed with A. ova/is). (35- 39) A . copulata. Van Heurck Slide No.2 (BM 26313), Dieghem (as A. affinis). (40 - 45) A. copulata. Van Heurck Slide No.424 (BM 26735), La Hulpe (as A. ova/is var. afjinis). (46- 49) A. copulata. Van Heurck Slide No.424 (NIWR 22/424), La Hulpe (as A. ova/is var. affinis). (50-53) A. copulata. BM 23137, Lewes (as A. minutissima W. Smith, type) . (54) A. ova/is Kiitzing. BM 81032, Kiitzing material (as F ova/is). (55, 56) A. ova/is. BM 81029 and BM 81030, Kiitzing material (as F. ova/is) . Figures 30- 35a, 41a, 42 - 49, 54-56: B.F. Ilium. Figures 35b, 36, 38b-40, 4lb, 50 - 53 : P .C. Ilium. Figures 37, 38a: O.B.F. Ilium. All figures : x 1500. S. Afr. J. Bot., 1986, 52(5) 429

Further research into members of this complex (see comments wntmg. This list included Frustulia ova/is and Frustulia on slides below) has shown that Kiitzing's taxon (F. copulata) copulata, which are now called Amphora ova/is and Amphora is identical to A. af.finis Kiitzing, and that it should be ac­ copulata respectively. corded species status on the grounds of morphological and Slides examined: BM 81029 Early Frustulia Sps. Kiitzing structural differences. It is difficult to understand how in a material. Unnumbered packet filed under Pleurosigma atte­ single work Kiitzing (1844, p.107) united this taxon (as F. nuatum W. Smith (Diat. Herb. 1976). Diatomaceae. Selected copulata) with A. ova/is on the one hand, and on the other slide. (Figure 55). recognized it as a new species under the name A. af.finis, while BM 81030 As above. Diatomaceae. Selected slide. (Figure 56). recording it as Amphora libyca Ehrenberg, which has sub­ BM 81031 As above. Diatomaceae. Selected slide. sequently been recognized as synonymous with A. af.finis (cf. BM 81032 As above. Diatomaceae. Strewn slide. (Figures Cleve 1895). This paradoxical situation does explain, however, 11 - 13, 54). why many of the Kiitzing slides marked as A. ova/is contained only forms identifiable as F. copulata or A. af.finis. Notes: Four slides, prepared from this early Kiitzing material, As a result of our observations on the material examined were sent to us from the British Museum (Natural History). (see detailed comments below) as can identify two species. Of these, three (BM 81029- 81031) were of selected speci­ The first is Amphora ova/is (Kiitzing) Kiitzing, which has no mens, and the fourth (BM 81 032) was a strewn slide. The nomenclatural complications, since it is based on Frustulia latter is of the greatest interest, since it forms the basis of ova/is Kiitzing (1833). The second species requires more careful our discussion above. It contained the two Amphora species, consideration with regard to its correct nomenclature, since A. ova/is (Figure 54) and A. copulata (Figures 11 -13), which in the past different taxonomic opinions have resulted in a Kiitzing united, causing the confusion when some of the slides large number of names being given to the same taxon, depend­ dealt with below were examined. The specimens of A. copula­ ing on the authors' concept. As shown above Kiitzing des­ fa were all frustules, showing the typical dorsal central area, cribed it as Frustulia copulata (Kiitzing 1833, p.541) and which is here closed on the ventral side by a row of puncta. Amphora af.finis (Kiitzing, 1844, p.107). Ehrenberg (1840, One frustule (Figure 13), however, showed one valve without p.205) recognized it as Amphora libyca. Van Heurck (1880- a central area. This is probably an atypical form, as we have 85, p.59) considered it a variety of A. ova/is, calling the taxon found only one other specimen with a similar structure (see A. ova/is var. af.finis, while Cleve (1895, p.105) agreed with paragraph 4.2.9). The examples of A. ova/is, also observed Van Heurck's taxonomic ranking, but preferred the combina­ on this slide, were typical of our present concept of this tion A. ova/is var. libyca. Since we recognize the taxon as species. a discrete species, only the first three nomenclatural combina­ The selected specimens on the remaining three slides, BM tions above come under consideration. Of these, F. copulata 81029 (Figure 55), BM 81030 (Figure 56) and BM 81031 were Kiitzing (1833), despite being a forgotten name since its all examples of A. ova/is, agreeing completely with the modem unification with A. ova/is (vide Kiitzing 1844, p.107) is the concept of the species (cf. Patrick & Reimer 1975). oldest valid name for the taxon. Therefore, following the International Code of Botanical Nomenclature (Voss et a!. 4.2. Slides containing Amphora copulata 1983) the correct name for this species should be Amphora 4.2.1. BM 18933 Amphora ova/is Kiitzing. 'Daskau' (Dieskau) copulata (Kiitzing) nov. comb. Accordingly, in our discussion 468 b. Coil. Kiitzing (Figures 14, 15) of specimens observed in the material described below, this Notes: The Dieskau material is important because it provides combination is used. clear confl.l1Tlation of Kiitzing's concept of F. copulata ( = A. A complete list of synonyms with their citations follows :­ copulata). Although this slide (BM 18933) is marked as Am­ phora ova/is, the packet containing the material from which Amphora copulata (Kutzing) Schoeman & Archibald, nov. it was prepared is labelled 'Frustulia copulata Kiitz. (1833) comb. = Amphora ova/is' . The only Amphora species we observed Frustu/ia copu/ata Kiitzing 1833, p.541, pl.13, fig.6. on this slide was A . copulata. Two examples (valves) are Amphora libyca Ehrenberg 1840, p.205. illustrated in Figures 14 and 15. These are larger than the Amphora affinis Kiitzing 1844, p.107, pl.30, fig. 66. forms from the unnumbered Kiitzing material (Figures 11 - Amphora ova/is var. affinis (Kiitzing) Van Heurck 1880 - 13), but clearly display the characteristics of the species. Note 85, p.59, pl. I' fig. 2. the striae structure and the construction of the central area, which in both examples is closed on the ventral side by a row Amphora ova/is var. /ibyca (Ehrenberg) Cleve 1895, p.I04. of puncta. Amphora minutissima W. Smith 1853, p.20, pl.2, fig . 30. 4.2.2. BM 78084 Kiitzing, Alg. Aq. dulc. Germ. Decas IX, Having established that we are dealing with two distinct No. 84. Frustulia acuminata. (Figures 16-21). species, A. ova/is and A. copulata, we can now comment on the slides we examined in this study. Slides made from the Notes: As mentioned above (paragraph 4) we expected to fmd unnumbered packet of early Kiitzing material (consult para­ A. ova/is on this slide. However, our search for this species graph 4.1) which we used to define these two species will be was fruitless, but we did observe a number of examples of considered first. We then comment on all slides illustrating A. copulata, some of which are illustrated in Figures 16- A. copulata (paragraph 4.2), and end with those depicting 21. This was somewhat puzzling, since Patrick & Reimer A. ova/is (paragraph 4.3). (1975, p.68, pl.l3, fig.1) depict a frustule of the true A. ova/is from this material. Since slide BM 78084 (prepared 10.5.1978) 4.1. Kiitzing material: unnumbered packet with no locality was not available to them when their book was published, given we assume that they have a separate slide made from the The only information given on the packet is a list of species exsiccata material. It is possible, therefore, that the true A. occurring in this material written in Kiitzing's own hand- ova/is is very rare in this material, while A . copulata is slightly 430 S.-Afr. Tydskr. Plantk ., 1986, 52(5)

commoner. Furthermore, as we have seen above, Klitzing may specimens with A. copulata we are convinced that the two have identified A . copulata as A . ova/is at this stage, and species are conspecific. A . ajfinis must therefore be considered hence the lack of reference to the presence of A. copulata. a synonym of A . copulata. Specimens of A . copulata observed in this material showed 4.2.6. BM 18934 Amphora ova/is Klitz. - B minor Klitz. some variability in the construction of the dorsal central area. Tennstadt, Mat. No. 171. Coli. Klitzing. (Figure 29). Some specimens (Figures 17, 18, 21) were quite typical in having a row of pores closing the ventral side at the central Notes: Only five specimens of a form identical to A. copulata nodule, while in others (Figures 16, 19, 20) the central area were found on this slide. One of these is illustrated in Figure 29. was open at the central nodule. This variety (?) described by Klitzing (1844, p.107) is not mentioned by VanLandingham (1967) in his synonymy of A . 4.2.3. BM 78086 Klitzing, Alg. Aq. dulc. Germ. Decas IX, ova/is, and does not appear to have been recorded elsewhere No . 81. Leucopetram ( = Weisenfels). Frustulia subulata (not since its determination by Klitzing. If these specimens truly a diatom according to VanLandingham 1971 , p.1850, as F. represent Klitzing's concept of the var.(?) minor, it further subulator). (Figure 24). complicates understanding Klitzing's separation of the various Notes: Since Klitzing (1833, p.539) commented that Frustulia taxa in the A. ova/is complex (see paragraph 4). ovalis ( = A. ova/is) occurred with F. subulata in material 4.2.7. BM 26312 Amphora ova/is Klitzing. Falaise. Van from Weisenfels, we concluded that representatives of A. Heurck Type slide no. 1. (Figures 30- 34). ova/is should be present on this slide. However, having scanned the whole slide, the only specimens of Amphora present were Notes: The slide is marked Amphora ova/is, but among true a frustule of Amphora pediculus (Klitzing) Klitzing (cf. representatives of this species (Figures 63 - 69) are large num­ Schoeman & Archibald 1978), and a broken valve of A. bers of A. copulata (Figures 30- 34). The latter have a greater copulata (Figure 24). Using an England Finder this specimen size range (length 21,5-51,0 !liD and valve breadth 6,5- may be found on the slide at coordinate 0 3312. It differs 10,0 !liD) than recorded on previous slides. Nevertheless, these from specimens described above by having an open central specimens are structurally quite typical of A . copulata. In these area, and ventral striae each composed of two puncta. The the dorsal central area is closed by puncta or short striae. latter has also been observed in other specimens (see below), 4.2.8. BM 26313 Amphora affinis Klitzing. Dieghem (Bel­ but is very distinct in this example. gium). Van Heurck Type slide No. 2. (Figures 35- 39). 4.2.4. BM 18936 Amphora ova/is Klitz. Wurzburg 470. Coli. Notes: Specimens on this slide are identical to A. copulata. Klitzing. (Figures 22, 23). The dorsal central area was again either open at the central Notes: This slide is also marked as containing A . ova/is, but nodule (Figures 36, 37) or closed by puncta (Figures 35, 38, we only found a few valves of A . copulata. In the examples 39). illustrated (Figures 22, 23) the dorsal central area is closed 4.2.9. BM 26735 Amphora ova/is var. affinis (Klitzing) van by a row of puncta. Otherwise they agree entirely with the Heurck. La Hulpe (Belgium). Van Heurck Type Slide No. forms on slide BM 81032 (consult paragraph 4.1), the type 424. (Figures 40 - 45). slide of A . copulata, as well as with the types of A . affinis on slide BM 18938 (see paragraph 4.2.5). Notes: Together with examples (Figures 40-45) from this slide in the British Museum (Natural History) collection, we have 4.2.5. BM 18938 A . affinis Klitz. Falaise 436. Type. Coli. included specimens (Figures 46- 49) from the corresponding Klitzing. (Figures 25- 28). slide in the NIWR collection (NIWR No. 22/424 = Van Notes: Klitzing (1844, p.107) described A. affinis from materi­ Heurck Type slide No. 424). All the specimens illustrated here al gathered by de Brebisson at Falaise, and this slide has been belong to A. copulata, again showing a range in the dorsal designated its type slide. Despite the fact that Reimer examined central area construction. In addition to both open and closed this particular slide (see annotation on the slide label), it does central areas, one specimen (Figure 48) had no central area not seem that he (vide Patrick & Reimer 1975, p.69). included at all (cf. one valve of Figure 13 from the type slide). This these specimens in his circumscription of A. ova/is var. affinis. appears to be an abnormal condition, somewhat reminiscent The dimensions and striae counts that we obtained for speci­ of the central striae in A. ovalis. mens on this slide (BM 18938) closely fit Reimer's (op. cit.) description for A . ova/is var. pediculus (Klitzing) Van Heurck. 4.2.10. BM 23137 Amphora minutissima W. Smith. Lewes, It is therefore interesting that Reimer himself suggested that December 1850. Type. Coli. W. Smith (Figures 50- 53). the forms he designated as var. affinis and var. pediculus may be Notes: Figures 50- 53 illustrate specimens from the type slide identical. In our opinion this should be the case, and we of Amphora minutissima W. Smith. They are identical to A. (Schoeman & Archibald 1978) have previously commented copulata, although in all the observed examples the dorsal in greater detail on the taxonomy of A. pediculus, A. per­ central areas are open. A . minutissima W. Smith is therefore pusilla and A. ova/is. considered a synonym of A . copulata. We examined a number of specimens of A. affinis from slide BM 18938, and have illustrated four examples in Figures 4.3. Slides containing Amphora ova/is. 25-28. The dimensions were:- length 21,0-40,5 !liD, breadth 4.3.1. BM 18937 Amphora ova/is Klitzing. Battaglia (ex Mene­ 5,5-8,0 !liD, dorsal striae near the centre (13)14-16(17) in ghini). Coli. Klitzing No. 444. (Figures 57, 58). 10 !liD, reaching 16- 19 in 10 11m at the poles; ventral striae (13)14 -16(17) in 10 !liD. The dorsal central area was generally Notes: We have illustrated two specimens (Figures 57, 58) of open at the central nodule (Figures 25, 27, 28), but in some A. ova/is from material collected by Meneghini at Battaglia specimens (e.g. Figure 26) it was closed by two puncta. A in Italy. These agreed completely with the present concept of similar situation was noticed in examples of A. copulata on this species (cf. Patrick & Reimer 1975), and conformed slide BM 78084 (see paragraph 4.2.2). Having compared these precisely to the examples of F. ovalis (Figures 54- 56) from S. Afr. J. Bot., 1986, 52(5) 431

Kiitzing's unnumbered packeted material (see slides BM 4.3.3. BM 19509 Amphora ova/is Kiitzing. Falaise, France. 81029-81032 in paragraph 4.1). A few specimens of A. Eulenstein Diat. Spec. typ. No. 95. Coli. Roper. (Figures pediculus were also observed (see Schoeman & Archibald 60-62). 1978). Notes: This slide was prepared by Eulenstein (1867) from 4.3.2. BM 18935 Amphora ova/is Kiitzing. Falaise. Coli. Kiitzing herbarium material. It is another gathering from Kiitzing No.l680. (Figure 59). Falaise, France, and contains many fine examples of the Notes: This slide is made from de Brebisson material sent to typical A. ova/is. Three are illustrated in Figures 60- 62. Kiitzing, and appears to be similar to the Falaise material from 4.3.4. BM 26312 Amphora ova/is Kiitzing. Falaise, France. which Van Heurck ( 1880- 85; 1884- 87) produced his Type Van Heurck Type slide No.I. (Figures 63- 69). slide No. I (see comments on slide BM 26312, paragraphs 4.2.7 and 4.3.4). Both A. ova/is and A. copulata are present Notes: This is yet another slide made from Falaise material. It on the slide. Unfortunately the examples of A. ova/is are not is, however, not clear whether they have a common origin, or very good, and only one specimen has been illustrated (Figure whether they are different gatherings. Again there were many 59) to show that it agrees with examples from Kiitzing's examples illustrating the typical characteristics of A. ova/is. unnumbered material (consult paragraph 4.1). A few are represented in Figures 63-69. This slide also

10pm

Figures 57-69 (57, 58) A. ova/is Kiitzing. B.M. 18937, Battaglia. (59) A. ova/is. BM 18935, Falaise. (60- 62) A. ova/is. Eulenstein Slide No. 95 (BM 19509), Falaise. (63- 69) A. ova/is. Van Heurck Slide No.I (BM 26312), Falaise. Figures 57-59, 61, 64, 66, 67, 69: B.F. Ilium. Figures 60, 62: P.C. Ilium. Figures 63, 65, 68: O.B.F. Ilium. All figures: x 1500. 432 S.-Afr. Tydskr. Plantk., 1986, 52(5)

Table 1 Summary of dimensions (in J.lm) and striae counts (in 10 J.lm) recorded in this study for A. copu/ata and A. ova/is

Dorsal striae Breadth Breadth Ventral Taxon Length (valve) (frustule) (near centre) (poles) striae A . copulata 12 - 51 4 - 10 12-18 (13)14 - 16(18) (14)16-18(20) (13)14-17(20) A. ova/is 38 - 86 9,5-14 25 - 45 10- 12(13) (11)12 - 14 10 - 12

Table 2 Summary of the reference slides for A. Notes: Gregory (1857, p.518) described this species from copulata and A. ova/is in the British Museum (Natural material collected both in Lamlash Bay (Arran) and in neigh­ History) collections bouring Loch Fine. According to Hendey (1964, p.262) slide BM 1196, prepared from Arran material in the Greville Slide No. Our Figure No. Notes collection of the British Museum (Natural History), is the type Amphora copulata slide for A. proteus. Four ringed frustules of this species were BM 18933 14, 15 examined on this slide and are presented in Figures 70- 73 BM 18934 29 to show their typical characteristics. They display some of the BM 18935 mixed with A. ova/is variability in frustular shape described by Gregory, and agree BM 18936 22, 23 with his description of the striae with respect to the undulate BM 18938 25 - 28 longitudinal disposition of the puncta (see Figures 70, 71). BM 23137 50 - 53 BM 26312 30 - 34 mixed with A. ova/is There is, however, some discrepancy with regard to the ventral BM 26313 35 - 39 striae ('striae of the inner compartments'). Gregory described BM 26735 40 - 45 Van Heurck slide No. 424 and depicted two rows of striae separated by a blank longi­ NIWR 22/ 424 46 - 49 Van Heurck slide No. 424 tudinal band. In the examples illustrated here, only the ventral BM 78084 16 - 21 striae in Figure 70 approach this. In the remaining three BM 78086 24 with frustule of A. pediculus specimens (Figures 71 - 73) the ventral striae are either unin­ BM 81032 II - 13 mixed with A. ova/is terrupted or a row of puncta is cut off, either on the inner Amphora ova/is side along the raphe (Figure 73) or on the ventral side (Figure BM 18935 59 mixed with A. copula fa 71). Gregory does not mention the dorsal central area, which, BM 18937 57, 58 mixed with A. pediculus in the four ringed specimens, is a relatively small lens-shaped BM 19509 60-62 area above the central nodule, closed on the ventral side by a BM 26312 63 - 69 mixed with A. copulata row of puncta. Nevertheless we have accepted these examples BM 81029 55 as representative of Gregory's species. BM 81030 56 A number of ringed specimens on a further four slides BM 81031 BM 81032 54 mixed with A. copulata prepared from Lamlash (Arran) material were examined. These specimens (Figures 74- 79) are all clearly conspecific with the type examples, but show a wide variation in the contains examples of A. copulata (see paragraph 4.2.7). arrangment of the ventral striae. The individuals approaching Table 1 summarizes the dimensions and striae counts we most closely Gregory's description of the ventral striae are recorded for A. ova/is and A. copulata from the above­ those on slide BM 1216 (Figures 75, 76). They have two more mentioned slides. Table 2 presents a summary of reference or less complete rows of pores. Figure 77 illustrates a specimen slides in the British Museum (Natural History) bearing exam­ from slide BM 1340 which very closely resembles Gregory's ples of these two species. (1857) figure 8le on plate 13. We also examined a ringed specimen from slide BM 1455 5. Amphora proteus Gregory (Figure 80) made from Loch Fine material, one of the type Gregory 1857, p.518, pl.l3, figs 81 a-e. Cleve 1895, p.l03. localities mentioned by Gregory (1857). This individual is Peragallo & Peragallo 1897- 1908, p.200, pl.44, figs 24- 27. basically the same as the other Lamlash examples, including Hendey 1964, p.262. the types, but is considerably larger and differs slightly in the Slides examined: BM 1196 Arran. Gregory 1857. Type. Coli. construction of the central area. The latter is open on the · Greville. Ring Nos 2- 5 (Figures 70- 73). ventral side, i.e. it is not bounded by a row of pores just above BM 1213 Lamlash, Arran. Gregory 1857. Coli. Greville. Ring the central nodule. No.I (Figure 74). It is interesting to note that, contrary to Gregory's (1857, BM 1216 Lamlash, Arran. Gregory 1857. Coli. Greville. Ring p.518) claim of great variation in striae numbers in A. proteus, Nos 1,2 (Figures 75, 76). we found that the above specimens have rather constant striae BM 1340 Lamlash, Arran. Gregory 1857. Coli. Greville. Ring counts, with relatively narrow limits. As suggested in our No.2 (Figure 77). discussion of A. marina, it is possible that Gregory included BM 1359 Arran, 1856. Coli. Greville. Two ringed frustules such specimens in A. proteus, which would account for the (Figures 78, 79). doubling of the striae counts in the smaller specimens he BM 1455 Loch Fine. Gregory. Coil. Greville. Ring No.4 mentioned. (Figure 80). Finally, two slides from the F.W. Payne collection in the BM 38691 Caloundra, Queensland. Coli. F.W. Payne. British Museum, reported to contain A. proteus, were exam­ (Figures 81- 83). ined. One (BM 38691: Figures 81 - 83) is prepared from a BM 38693 Colon (Panama), Pinna shells. Coli. F.W. Payne. sample collected at Caloundra in Queensland, Australia, while (Figure 84- 86). the other (BM 38693; Figures 84- 86) is made from the S. Afr. J . Bot., 1986, 52(5) 433

contens of Pinna shells collected at Colon (Panama). While The Caloundra examples (Figures 81- 83) differ mainly in there is some similarity in frustular shape, there are certain that the ventral striae completely fill the ventral part of each distinctions casting doubt on their identification as A. proteus. valve in contrast to the broad structureless areas on the ventral

10pm

Figures 70 - 76 (70 - 73) A. proteus Gregory. BM 1196, Arran. (74) A. proteus. BM 1213, Lamlash, Arran. (75, 76) A . proteus. BM 1216, Lamlash, Arran. Figures 70 - 76: B.F. Ilium. All figures: x 1500. 434 S.-Afr. Tydskr. Plantk., 1986 , 52(5)

side of true A . proteus. The specimens from Colon (Figures 6. Amphora robusta Gregory 84- 86) are quite obviously a different taxon, distinguished Gregory 1857, p.516, pl.l3, fig .79. Cleve 1895, p.103. Peragal­ from A . proteus by the ventral striae structure and the very lo & Peragallo 1897- 1908, p.202, pl.44, figs 33, 34. Hendey prominent conopeum on the dorsal side of the raphe. 1964, p.262, pl.38, fig. 7. Dimensions of specimens examined: Arran specimens: Length Slides examined: BM 1124 Loch Fine, coarse sand. Gregory 56,0- 83,0 J.!m; breadth of frustule 27,0-43,0 J.!m; dorsal 1857. Coli. Greville. Ring Nos 3, 5 (Figure 87- Ring No. 5). striae at and near the centre (10)11- 12 in 10 J.!m, at the poles BM 1193 Lamlash, Arran. Gregory 1857. Coli. Greville. Ring (11)12- 13(14) in 10 J.!m; ventral striae 9- 12 in 10 J.!m; striae No.4. on valve mantle 10-14 in 10 J.!m. BM 1216 Lamlash, Arran. Gregory 1857. Coli. Greville. Ring Loch Fine example: Length 113,5 J.!m; breadth of frustule Nos 3, 6. 53 ,0 J.!m; dorsal striae at and near the centre 9 in 10 J.!m, at BM 3290 Arran. Coli. Greville. MF 26/ 35 (Figure 88). the poles 10 in 10 J.!m; ventral striae 8- 9 in 10 J.!m. BM 3295 Arran. Coli. Greville. MF 30/ 34 (Figure 89).

78

10pm

Figures 77 - 80 (77) A . proteus Gregory. BM 1340, Lamlash, Arran. (78, 79) A . proteus. BM 1359, Arran. (80) A. proteus. BM 1455, Loch Fine. Figures 77 - 80: B. F. Ilium . All figures : x 1500. S. Afr. J. Bot., 1986, 52(5) 435

BM 3297 Arran. Coli. Greville. MF 32/ 12. example, unsuitable for photography as it lies obliquely, and simultaneous focus of the raphe structures is impossible. On Notes: Gregory (1857, p.516) cited two localities, Loch Fine the other hand, the frustule in Ring No.5 (Figure 87) lies in and Lamlash Bay, for this species. According to Hendey such a plane that most of the characteristic features are (1964, p.262) slide BM 1124, made from the Loch Fine reasonably clear. It agrees very closely with Gregory's descrip­ material, has been designated the type slide for this species. tion, although modern terminology would help to clarify There are two ringed specimens (Ring Nos 3 and 5) on this certain statements. Gregory (op. cit.) referred to a peculiar slide. Unfortunately, the frustule in Ring No.3 is a poor aspect of the valve in which a triangular blank space is formed

10pm

Figures 81 - 86 (81 - 83) 'A. proteus Gregory' (sensu Payne). BM 38691, Ca1oundra, Queensland. (84- 86) 'A. proteus Gregory' (sensu Payne). BM 38693, Colon, Panama. Pinna shell s. Figures 81 - 83a, 84-86: B.F. Ilium. Figure 83b: P .C. Ilium. All figures: x 1500. 436 S.-Afr. Tydskr. Plantk., 1986, 52(5)

10pm

Figures 87-89 (87) A. robust a Gregory. BM 1124, Loch Fine (type). (88) A. robusta. BM 3290, Arran. (89) A. robust a. BM 3295, Arran. Figures 87, 88: B.F. Ilium. Figure 89: O.B.F. Ilium. All figures: x 1500.

by the proximal ends of the raphe (curve lines) and a straight Dimensions of specimens examined: Length 74,0-120,0 J.!m; line. On examination of the specimen in Figure 87, it would breadth of frustules 39,0-42,5 J.!m; breadth of valves 17,0- appear that Gregory's 'straight line' refers to the dorsal edge 24,0 J.!m; dorsal striae at the centre 7- 8 in 10 J.!m, near the of a very broad conopeum, developed on both sides of the centre 6- 7 in 10 J.!m and at the poles 7 - 8 in 10 J.!m; ventral raphe, which masks the ends of the striae below it. The striae striae 6- 7(8) in 10 J.!m. It should be noted here that the themselves appear to be deep grooves in the basal siliceous frustules recorded by Gregory (1857, p.516) are broader layer or lie between transverse costae with a single row of (45,5- 61,0 J.!m) that those observed by us . circular to oval puncta within each groove. Because of the relative clarity of the valve characteristics in this particular Acknowledgements specimen (Ring No.5 - Figure 87) we have designated it the Once again, the authors wish to express their sincere thanks holotype of the species. to Mr J.F.M. Cannon, Keeper of Botany, British Museum We examined five other slides from material gathered in (Natural History) for granting permission to the senior author Arran, two of which are labelled Lamlash. Unfortunately, to conduct his studies in the British Museum, and to Ms P .A. most of the marked specimens on these slides were of poor Sims for invaluable assistance with the diatom collections and quality, and unsuitable for photographic reproduction. There her helpful criticism of the manuscript. Thanks are also were, nevertheless, two specimens that demonstrate the diag­ extended to Mr R. Ross (c/o British Museum, Natural His­ nostic features of this species. Figure 88 shows a frustule from tory) for checking the nomenclature of Amphora copulata. slide BM 3290 located at T39/3 using an England Finder. This Finally, we wish to thank Ms V.H. Meaton for producing the specimen is identical to the holotype, and shows the edge of fmal prints used to illustrate the specimens observed. Financial the conopeum even more distinctly. The other example, which assistance in the form of a British Council grant partly can be found at S40/4 using an England Finder, is a valve sponsored these investigations, and is gratefully acknowledged. (Figure 89) on slide BM 3295. This example is valuable as This paper is published with the approval of the National it shows the shape of the valve, and the abrupt dorsal deflec­ Institute for Water Research, Pretoria, Republic of South tion of the proximal ends of the raphe branches. Africa. S. Afr. J. Bot., 1986, 52(5) 437

References Observations on Amphora species (Bacillariophyceae) in CLEVE, P.T. 1895. Synopsis of the Naviculoid diatoms, Part II. the British Museum (Natural History). II. Some species from K. svenska VetenskAkad. Hand!. 27: I - 219. the subgenus Psammamphora Cleve. Nova Hedwigia (in press). EHRENBERG, C.G. 1840. Charakteristik von 274 neuen Arten SCHOEMAN, F.R. & ARCHIBALD, R.E.M. 1986c. Observations von Infusorien. Ber. Bekanntm. Verh. K. Preuss. Akad. Wiss. on Amphora species (Bacillariophyceae) in the British Museum Berlin 1840: 197-219. (Natural History). III. Two species from the subgenus EULENSTEIN, T. 1867. Diatomacearum Species Typicae Amblyamphora Cleve. Nova Hedwigia (in press). (Exsiccata), Part I, 2 p. & 100 slides. Stuttgartiae. SCHOEMAN, F.R. & ARCHIBALD, R.E.M. 1986d. GREGORY, W. 1857. On new forms of marine Diatomaceae Observations on Amphora species (Bacillariophyceae) in the found in the Firth of Clyde and in Loch Fine. Trans. R. Soc. British Museum (Natural History). IV. Some species from the Edinb. 21: 473-542. subgenus Diplamphora Cleve. Cryp!OJlamie: Algologie 7(1): 9-21. HENDEY, N.J. 1964. Bacillariophyceae (Diatoms). In: An SCHOEMAN, F.R. & ARCHIBALD, R.E.M. 1986e. Observations introductory account of the smaller algae of British coastal on Amphora species (Bacillariophyceae) in the British Museum waters, Fishery Investigations, series 4 part 5, 317 p. & 45 pis. (Natural History). VI. Some species from the subgenus Her Majesty's Stationery Office, London. Halamphora Cleve. Nova Hedwigia (in press). KUTZING, F.T. 1833. Synopsis Diatomearum oder Versuch einer SMITH, W. 1853. A synopsis of the British Diatomaceae; with systematischen Zusammenstellung der Diatomeen. Linneae 8: remarks on their structure, functions, and distribution, Vol. I. 529 - 620, pis 13- 19. 89 p. & 31 pis. John Van Voorst, London. KUTZING, F.T. 1833 -36. Algarum aquae dulcis Germanicarum, SMITH, W. 1857. Notes of an excursion to the Pyrenees in search Decas I - XVI. Exsiccatae, text without page numbers (39 p.). of Diatomaceae. Ann. Mag. nat. Hist., Ser. 2, 19: 3 - 15. C.A. Schwetschkii et Fil., Halis Saxonum. VAN HEURCK, H. 1880 - 85. Synopsis des Diatomees de KUTZING, F.T. 1844. Die kieselschaligen Bacillarien oder Belgique. Atlas: 132 pis. Text: 235 p. & 3 pis. J. Ducaju & Diatomeen. 152 p. & 30 pis. Nordhausen. Co., Anvers. PATRICK, R. & REIMER, C.W. 1975. The diatoms of the VAN HEURCK, H. 1884 - 87. Types du Synopsis des Diatomees United States exclusive of Alaska and Hawaii, Vol. 2, Part I. de Belgique. Serie I - XXII. Anvers, 123 p. & 550 sl ides. Monogr. Acad. nat. Sci. Phi/ad. 13: 1-213. VANLANDINGHAM, S.L. 1967. Catalogue of the fossil and PERAGALLO, H ." & PERAGALLO, M. 1897 - 1908. Diatomees recent genera and species of diatoms and their synonyms, part marines de France et des districts maritimes voisins. I: Acanthoceras through Bacillaria, pp. I - 493 . J. Cramer, Micrographe-Editeur, Grez-sur-Loing (S.-et M.), 491 & 48 p., Lehre. 137 pis. VANLANDINGHAM, S.L. 1971. Catalogue of the fossil and SCHOEMAN, F.R. & ARCHIBALD, R.E.M. 1978. The diatom recent genera and species of diatoms and their synonyms, part flora of southern Africa. No.4, April 1978. CSIR Special 4: Fragilaria through Naunema, pp. 1757 - 2385 . J. Cramer, Report W AT 50. No pagination; series of plates with text, 68 Lehre p. National Institute for Water Research, CSIR, Pretoria. VOSS, E.G., BURDET, H.M., CHALONER, W.G., SCHOEMAN, F.R. & ARCHIBALD, R.E.M. 1986a. DEMOULIN, V., HIEPKO, P., McNEIL, J., MEIKLE, R.D., Observations on Amphora species (Bacillariophyceae) in the NICOLSON, D.H., ROLLINS, R.C., SILVA, P.C. & British Museum (Natural History). I. Some species from the GREUTER, W. 1983 . International code of botanical subgenus Oxyamphora Cleve. Nova Hedwigia (in press). nomenclature, 472 p. Bohn, Scheltema & Holkema, SCHOEMAN, F.R. & ARCHIBALD, R.E.M. 1986b. Utrecht/ Amwerpen.