PEABODY MUSEUM OF NATURAL HISTORY YALE UNIVERSITY ON FILINIA TERMINALIS (PLATE) AND F. PEJLERI SP. N. (ROTATORIA: FAMILY TESTUDINELLIDAE)

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PEABODY MUSEUM OF NATURAL HISTORY YALE UNIVERSITY Number 81 June 1, 1964 New Haven, Conn. ON FILINIA TERMINALIS (PLATE) AND F. PEJLERI SP. N. (ROTATORIA: FAMILY TESTUDINELLIDAE) G. E. HUTCHINSON DEPARTMENT OF BIOLOGY, YALE UNIVERSITY In 1886, Plate described, as Triarthra terminalis, a rotifer found in the spring, in the vicinity of Bonn, which differed from T. longiseta Ehrenberg, now referred to Filinia, in having the posterior appendage inserted apically, rather than ventrally some little distance from the posterior end of the body. Apart from usually lacking spines on the appendages (such spines were present on one specimen), the new species appears to have resembled Filinia longiseta in shape and general characters. Plate gave no figure, but Caiman (1892) who may be regarded as the first reviser relative to T. terminalis, illustrated a single specimen from Dundee tap water (fig. lb), which clearly belongs, in spite of subsequent erroneous statements for which I am partly responsible, with the European species later discussed by Carlin and Pejler, and referred by the latter to F. terminalis (Plate). Pejler (1957a, b) gives the most complete account of the species. It appears in Europe to be a cold stenotherm organism, known from Swedish Lapland southward to Switzerland,

2 Postilla Yale Peabody Museum No. 81 % Figure la. Filinia pejleri sp. n. Ootacamund, Nilghiri Hills, S. India (holotype), X230. Figure lb. F. terminalis (Plate), first figured specimen, Dundee, Scotland; after Caiman (1892). and as Voigt (1957) correctly indicates, recorded as F. longiseta by Edmondson and Hutchinson (1934) from Ladakh and Kashmir. In F. terminalis the insertion of the posterior appendage, if not terminal, is less and usually much less than

June 1, 1964 Filinia terminalis and F. pejleri 3 IOJJL from the posterior end of the body. Such animals were regarded as a cyclomorphotic winter form of F. longiseta by Slominski (1926), who seems to have found F. limnetic a during the summer and F. terminalis during the rest of the year in the Polish locality that he studied. It is evident, however, that terminalis can occur as the only planktonic member of the genus in a lake, as in the Mansfelder See (Colditz, 1914). Both Carlin (1943) and Pejler (1957a, b) make an excellent case for regarding terminalis as distinct from longiseta, though Carlin, following Edmondson and Hutchinson's (1934) misidentification of terminalis, believed the first valid name of the species to be major Colditz. There is also in Europe an array of forms in which the posterior appendage is inserted well in front of the posterior apex of the body, the distance between insertion and posterior end varying from rather over IOJJL in small to over 30[JL in large specimens. In Scandinavia these animals can be separated into two discontinuous groups; in one the anterior appendages are less than 350fji long, in the other more than 400JJL long. The ratio of posterior to anterior appendage length is greater in the first than in the second group. Carlin (1943) regarded the two groups as species; namely, F. longiseta (Ehrenberg) living in ponds and F. limnetica (Zacharias) living in lakes. Voigt (1957) accepted Carlin's separation, though it is very probable that Voigt's conception of longiseta would include specimens of limnetica. Plotting the length of the posterior appendage against the mean length of the two anterior appendages for all specimens of longiseta, terminalis and limnetica from Sweden, Pejler found evidence of two regression lines converging in an area occupied by points defining longiseta s. str. When, however, a double logarithmic plot is made it appears that the Scandinavian data give envelopes around two parallel straight lines with a slope of about 1.33. One line runs through the envelopes of terminalis and longiseta, the other through that of limnetica. Pejler was doubtful as to the specific separation of longiseta and limnetica, since a few specimens, marked by saltires (X) in figure 2, taken in ponds and rivers in central Europe, appeared to be intermediate. In view of the great number of points

Postilla Yale Peabody Museum No. 81 defining the envelopes, seventy for terminalis, forty-one for limnetica and seventeen for longiseta, it seems likely that these points, probably not related to the ordinary growth patterns of the individual species involved, represent introgressive hybridisation, or perhaps very large specimens of longiseta with broken posterior appendages. The specimens recorded as longiseta from the lake at Ootacamund by Edmondson and Hutchinson (1934) are certainly referable to F. limnetica, as indicated by the open circles in figure 2. Parise (1961) has considered several Italian populations, which must be discussed in the present context (figure 3). 700f 600h UJ 500 CD < Q 2 400 LU Q_ Q_ < 300 DC O O 200 Q_ L. _.._.! _ i. h E TERMINALIS a o X R PEJj-ERI R LIMNETICA R CF. LONGISETA KASHMIR AND LADAKH ALL RECORDS NILGHIRI HILLS C.EUROPE y. y / u% / ) / y / TERMINALIS-^/ / / / LONGISETA; 7 '*//''' ^LIMNETICA /A A/ /// SWEDEN / / /' y x (// i''' / 1 /// / //? y / 100 -L 1 1 i i i i 1 I00L 200 300 400 500 600 800 1000 MEAN ANTERIOR APPENDAGES J. Figure 2. Relationship of length of posterior appendages to mean length of the anterior in F. longiseta (Ehrenberg), F. terrninalis (Plate), F. limnetica (Zacharias) and F. pejleri sp. n., based on measurements of Pejler and the present author.

June 1, 1964 Filinia terminalis and F. pejleri 5 700 600-500 =3 " 400 l±j O < O 300 UJ a. Q. < O 200 01 UJ h- c/> o a..pejleri / Y / } NEMIH NEMIItt / V \ j S / ///''"/'yj/^7^^emi LONGISETA/7/ / /^LIMNETICA SWEDEN/ 1 \ / I / /^TERMINALIS V 100 I00LI 200 300 400 500 600 700 800 1000 MEAN ANTERIOR APPENDAGES J. Figure 3. Envelopes of figure 2 with those of the three populations (dotted lines) of Lake Nemi, from Parise. In Lake Nemi three populations have occurred during the history of partial drainage and refilling of the lake. One of these (Nemi I), when the appendage lengths are plotted logarithmically gives a set of points falling within an envelope on the upper side of that of F. limnetica. Parise says nothing definite about the insertion of the posterior appendage in this population. Apart from a graphical presentation of appendage measurements and a statement that the appendages carry barely visible spinules, he remarks only that "la forme du lac n'accorde pas avec Filinia limnetica Zacharias" though the basis of this statement is far from clear. The other two populations are both tentatively considered in relation to F. terminalis, having a clear apical insertion of the posterior appendage. One (Nemi II) was present only in April, 1934, and consisted entirely of mictic females. Most specimens fall within the range of F. terminalis as established by Pejler, though a few have relatively slightly longer anterior

6 Postilla Yale Peabody Museum No. 81 appendages. The third population (Nemi III), amictic and with a relatively longer posterior appendage, occurred sporadi cally between 1922 and 1926. It is compared by Parise with a cold water population from Lago di Garda, which is presumably terminalis. From its position on the diagram of figure 3, however, one might suspect that the Nemi III population really belonged with the warm stenotherm species to be named below and that at different times all three of the limnoplanktonic species here discussed have occurred in the lake. In default of information on the body shape and on the seasonal occurrence of this population, no further conclusions are possible. In a population from a pond in the vicinity of Padua, the distance between the insertion of the posterior appendage and the apex of the body is said to be variable and of no value as a taxonomic character. The population would fall entirely within the envelope of Nemi III in figure 3, but is doubtless referable to a large long-spined form of the true F. longiseta. Hutchinson, Pickford and Schuurman (1932) recorded from South Africa, on the strength of an identification by the late David Bryce, a species that they called F. terminalis but which is obviously very different from the cold stenotherm species discussed in the preceding paragraphs. With the possible exception of the Nemi III population, no European specimens comparable to those from South Africa appear to be recorded (Hauer in lift.; Edmondson, 1935; Voigt, 1957). As Pejler points out, Hutchinson, Pickford and Schuurman (1932) were clearly in error as to their identification, as were Edmondson and Hutchinson (1934) when they recorded the same species from the lake at Ootacamund, and Edmondson (1935) when he noted the species from Mormon Lake, Arizona. In view of the necessity of having a valid name for this species in the discussion of the rotifers of the zooplankton in the forthcoming second volume of my Treatise on Limnology, I feel justified in putting forward as new, Filinia pejleri sp. n. Filinia terminalis Hutchinson, Pickford and Schuurman (1932), Edmondson and Hutchinson (1934), Edmondson (1935), Voigt (1957).

June 1, 1964 Filinia terminalis and F. pejleri 7 nee Triarthra terminalis Plate (1886), Caiman (1892). nee Filinia terminalis Pejler (1957a, b) Body fusiform, from two and a quarter to over three times as long as deep, hardly rounded dorsally, appendages minutely spinulose, posterior seta with a broad oblique base inserted terminally at the hind end of the body (fig. 1). Length of body Dorsoventral Depth Length Depth Right Left Anterior Anterior Posterior Appendage Appendage Appendage Ootacamund, S. India 138/** 138 142 54/* 50 58 2.56 2.60 2.45 342/* 300/* 242/* 330 330 262 308 333 242 Ruitkuil Pan, Transvaal 138 145 124 56 56 56 2.50 2.67 2.25 4.56(±, stuck together)456 318 (±) 408 401 415 422 325 Mormon Lake, 200 Arizona 60 3.33 432 480 360 * Dimensions in the first line refer to the holotype. Holotype: (YPM: Aschelminthes 25) Artificial Lake, Ootacamund, Nilghiri Hills, S. India; townet collection, 8 Nov. 1932, ph 6.6, temp. 17.5 C. (figure la; previously also figured by Edmondson and Hutchinson, 1934, fig. 2C). As Edmondson and Hutchinson point out, the largest specimens of the true F. terminalis, referred by them to longiseta, have a ratio of body length to depth overlapping that of pejleri. The latter, however, may always be separated by its more spindle-shaped body, with the dorsal surface hardly more rounded than the ventral. In contrast even the longest terminalis have a gibbous dorsal profile. It is also probable that the larger pejleri are proportionately narrower than the smaller, so that for any absolute size the ratio of length to depth would prove diagnostic. If the posterior appendage length is plotted against the mean length of the anterior appendages, the points for pejleri

8 Postilla Yale Peabody Museum No. 81 fall along the upper edge of the envelope defining this relationship in terminalis. F. pejleri is probably eurytopic chemically, occurring in the neutral waters of the type locality and in somewhat alkaline waters in the Transvaal. Its distribution suggests that it requires a warm temperate climate. It can occur, as at Ootacamund? sympatrically with F. limnetica. REFERENCES Caiman, W. T., 1892. On certain new or rare rotifers from Forfarshire. Ann. Scot. Nat. Hist. 240-245. Carlin, B., 1943. Die PlanktonrotatoTien des Motalastrom. Zur Taxonomie und Okologie der Planktonrotatorien. Medd. Lunds Univ. limnol. Instn. 5, 256 p. Colditz, F. V., 1914. Beitrage zur Biologie des Mansfelder Sees mit besonderen Studien iiber das Zentrifugenplankton und seine Beziehungen zum Netzplankton der pelagischen Zone. Z. wiss. Zool. 108:520-630. Edmondson, W. T., 1935. Some Rotatoria from Arizona. Trans. Amer. microscop. Soc. 54:301-306. Edmondson, W. T. and Hutchinson, G. E., 1934. Yale North India Expedition. Report on Rotatoria. Mem. Conn. Acad. Arts Sci. 10:153-186. Hutchinson, G. E., Pickford, G. E. and Schuurman, J. F. M., 1932. A contribution to hydrobiology of pans and other inland waters of South Africa. Arch. Hydrobiol. 24:1-154. Parise, A., 1961. Sur les genres Keratella, Synchaeta, Polyarthra et Filinia d'un lac italien. Hydrobiologia 18:121-135. Pejler, B., 1957a. Taxonomical and ecological studies on planktonic Rotatoria from northern Swedish Lapland. K. svenska Vetensk Akad. Handl. ser. 4 Bd. 6, no. 5; 68 p. Pejler, B., 1957b. On variation and evolution in planktonic Rotatoria. Zool. Bidrag fran Uppsala 32:1-66. Plate, L., 1886. Beitrage zur Naturgeschichte der Rotatorien. Jena Z. Naturw. 19 (N. F. 12): 1-20. Slominski, P., 1926. Sur la variation saissoniere chez Triarthra (Filinia) longiseta E. C. R. Soc. Biol., Paris. 94:543-545. Voigt, M., 1957. Die Radertiere Mitteleuropas. Berlin, Geb. Borntraeger I. 508 p.