R.W Jordan , A. Winter: An unusual coccolithophorid ...... , p. 8 1 - 83.

Journal of Nannoplankton Research, 20, 2, 1998 .

AN UNUSUAL COCCOLITHOPHORID (HAPTOPHYTA) SPECIMENFROM PUERTO RICO
Richard W Jordan* & Am os Winterf*Dept. of Earth & Environmental Sciences, Yamagata University, Yamagata , 990 8560 Japan; fDept. ofAlarine
Scie,;ces, University of Puerto Rico, PO Box 5000, Jvfayaguez, Puerto Rico 00681*Author to contact for reprints- e-mail: sh08l@kdw.kj.yamagata-u.ac.jp

Abslraci:During routine phytoplankton counting of offshore Puerto Rican samples, an unusual spec imenof Calcioso /enia murrayi Gran was encountered. C. murrayi typicall y has monomorphi c body-cocco liths,
but the new spec imen bore several types of body-coccolith : at lea st two types of rh ombolith, and anarrow, elliptical cocc olith new to sc ience. This latter type is not rhombi c like the others, but shares
several rhombolith features. However, it dilfcrs by being longer, relativel y flatt ent:d, sli ght ly twi sted, andwith distally-pointed ends. Although it is poss ib le th at the sin gle specimen rcprest:nts a coccospherc
composed mainl y of ma lformed coccol ith s, it is sugges ted here th at the new coccolith type ma y beevidence of an alternate phase. As thousands of onl y rhombolith-bearing coccospheres have been previouslyobserved by coccolithophorid workers, this new coccolith typ e ma y be produced for only a shortduration .

Introduction
Since the advent of electron microscopy and the improvement of single-cell isolation and culture techniques, ourknowledge of algal life-cycles has increaseO tremendously.
This is even true for the coccolithophorids, calcareous
haptophytes. which are predominantly restricted to openoceans. However, despite tl1ese advances, coccolithophorid
life-cycles are still poorly understood and those that arepresently known are mainly neritic species (e .g.
Pleurochrysis carterae (Braarud & Fagerland)
Christensen) or species that are easily grown (e.g.
Emiliania huxleyi (Lolunann) Hay & Mohler) (Billard,1994 ). On the other hand, when observing wild material,
coccolithophorid workers sometimes encounter convincing
examples of coccospheres seemingly bearing different
phases of a life-cycle. Considering the paucity of direct
observations on axenic cultured material. these records may
give useful information on possible species combinations.
In this way a number of associations have been witnessed
and reported in the literature (Thomsen et al., 1991 ; Kleijne,1993; Alcober &Jordan, 1997; Young et al., 1998, t11is issue;
Cros et al. , in press). Most of the known coccolithophorid
life-cycles involve either a motile holococcolithophorid andnon-motile heterococcolithophorid phase (e.g. Coccolithus
pelagicus (Wallich) Schiller: Parke & Adams, 1960), or amotile scale-bearing and non-motile coccolith-bearing
phase (e.g. E. huxleyi : Braarud, 1963). These motile andnon-motile phases are now known to represent t11e haploidand diploid stages respectively (Rayns, 1962; Green et al. ,1996). The interesting possibility raised by Alcober & Jordan(1997) was of an association between two non-motile
phases, Ceratolithus cristatus Kamptner and Neosphaera
coccolithomorpha Lecai -Schlauder. Recently, whilststudying ilie phytoplankton assemblages on the filters fromthe La Parguera time-series station in Puerto Rico (Jordan& Winter, in press), further examples of CeratolithusNeosphaera associations were encountered (Young et al. ,1998, this issue) , as well as an unusual specimen of
Calciosolenia murrayi, bearing several coccolith types,
one of which has not been reported previously in theliterature.

Location and collection
The RV Pezmar (belonging to the University of PuertoRico) was regularly used over a four year period (19921995) to collect biweekly samples off the shelf edge ofPuerto Rico. The sampling point (- 18°N, 67°W) lies westof La Parguera, the site of the university's marine biologicalstation. Water samples were collected systematically froma number of depths using a single five-litre Niskin bottle,
which could be closed manually using a messenger system.
Sampling depths were calculated by wire out. After eachcast, the water from the bottle was transferred to a plastic
carboy, and stored until the water could be filtered at theresearch station. Only samples collected during January
to May, 1995 have been analysed for coccolithophorids(see Jordan & Winter, in press, for further details) .
Preparation techniques
A known volume of each water-sample was filtered tluougha Millipore 47nun-diamcter, 0.45J..Un porosity polycarbonate
filter, washed with buffered distilled water, air dried, andstored in a plastic Petri dish. A piece of each filter, - 8x8mm,
was attached to an AI EM stub using a small amount ofglue, and t11en coated witl1 eitl1er Au or Pt!Pd in an Eiko IB3 ion coater. The filters were then examined in a Hitachi S2250N SEM. Photographs were taken either directly by acamera attached to the SEM or images were stored on
optical disc prior to being sent to the camera.

Calciosolenia murrayi
This species is regularly reported from the middle photiczone (- 70-100 m water-depth) of subtropical-tropical waters(Winter et al., 1994; Jordan & Winter, in press), where ilie
long, cylindrical shape of its coccosphere possibly
enhances its ability to photosynthesise in low lightconditions (Man ton, 1986). Man ton & Oates ( 1985)
compared t11e tluee living members of the Calciosoleniaceae(C. murrayi, Anoplosoienia brasiliensis (Lohmann)
Deflandre and Naviso/enia apri/ei Lecal) and concluded
that the body-coccoliths of these species were always
rhombic, even when they became very narrow at the poles.
Thus, t11ese coccolitl1s are tenned rhombolitl1S in preference

81Journal of Nannoplankton Research, 20, 2, 1998.

R.W Jordan. A. Winter: An unusual coccolithophorid ..... , p. 81 - 83.

to the other commonly-used tenn, scapholith, which means
boat-shaped (Jordan et al., 1995). Furthermore, it appears
that the rhomboliths on an individual coccosphere aremonomorphic despite variation in shape and size.
A collapsed specimen of C. murrayi was encountered (on the l5m filter of 13th February, 1995) which bore
two types of rhomboliths and a second body-coccolithtype (Plate 1, Figure I). C. murrayi is nonnally characterisedby monomorphic body-coccoliths, so our specimen may
be malformed. However, the nature of the new coccolithtype suggests that the specimen is truly exhibiting
polymorphic body-coccol iths. The rhomboliths of the newspecimen are either constructed of fine , slightly
overlapping laths with spaces in between (labelled ,a' inPlate I, Figure 2), or are seemingly more heavily calcified,with much broader laths with no gaps in between (labelled,b' in Plate I, Figure 2). This latter type was figured by
Manton & Oates ( 1985) and tentatively identified as C. aff.
murrayi. One of their specimens (their pl.4, fig. 14) clearly
showed that the body-coccoliths were monomorphic. The
third type of body-coccolith is typically longer andnarrower than most of the rhomboliths, bears more laths,
is slightly twisted at the ends, and is not rhombic (Plate 1,Figure 3). Another characteristic of this new coccolith type
is the raised, almost pointed, ends which can be clearlyseen in side-view. Whilst rhomboliths of C. murrayi andA. brasiliensis do not possess this feature, theunmineralised rhombic scales of Nm>isolenia Lecal appear
to do so (see Leadbeater &Mort on, 1973, pl.l , fig.4).
Nm>isolenia was recorded by Leadbeater & Morton(1973) from the coast ofYugoslavia between 0-10m waterdepth. Whether the unmineralised Navisolenia or ourmineralised new cocco1ith type represent phases in theCalciosolenia life-cycle is not known at present. However,
it is true to say that these haptophytes are unlike anyother living coccolithophorids, and should be certainly
separated at the family level. Interestingly, the rhombolithsof Calciosolenia and Anoplosolenia (or their ancestors)
can be found as far back as the Hauterivian of the Early
Cretaceous, and their morphology has remained seemingly
unchanged for over a 100m.y. (Perch-Nielsen, 1985). Priorto their first appearance, similar-looking coccolithsoccurred. Ofthese, Truncatoscaphus Rood et al. (JurassicEarly Cretaceous) of the Stephanolithiaceae may be theancestor of Calciosolenia , in particular T delftensis(Stradner & Adamiker) Rood et al. (Perch-Nielsen, 1985;Bown & Young, 1997). Given that the lineage of theCalciosoleniaceae, geologically speaking, is very long andits morphology has remained unchanged throughout thistime, it may prove to be a very useful genetic marker for theseparation of the haptophytes from other algae. Thus, inthe future it would be very interesting to compare thegenetics of these calciosolenioid species with other extant
coccolithophorids.
Acknowledgements
The authors would like to thank Dr. Jeremy Young, for
discussions relating to the evolution of rhombolith-bearing
coccolithophorids, Mark Reigle, Angie McGeehe, Jackie
Bonilla, Brett Stuart and Salvador Rivera Casiano for
helping in the collection and filtration of water-samples,82

and tl1e crew of the RV Pezmar for tl1eir cooperation during
sampling.
Refen~nccs

Alcober, J. & Jordan, R.W. 1997 . An intc:rcsting association
between Neosplwera coccolithomorpha and Ceratolitlms
crista/us (Haptophyta). Eur. J. Phycol., 32: 91-93.
Billard, C. 1994. Life cycles. In: J.C. Green & 13.S.C. Lcadbcatcr(Eds). The Haptophyte Algae. Th e Systematics Association
Special Volume, 51: 167-186. Clarcndon Press, Oxford.
Bown, P.R. & Young , J.R. 1997 . Meso zo ic calcareous
nannoplankton classification. J. Nmmoplankton Res. , 19: 2136 .
Braarud, T. 1963. Reproduction in the marine coccolithophorid
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press . New examples of holococcolith-heterococcolith
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ploidy levels analysed by flow cytometry. J. A.far. Syst. , 9:33-44.
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glossary of the extant Haptophyta of the world . J. mar. bioi.
Ass. U.K., 75: 769-814.
Jordan, R.W. & Winter, A. In press. Living microp1ankton
assemblages off the coast of Puerto Rico during JanuaryMay 1995. Mar. Aficropaleontol.
Klcijne , A. 1993. Morphology, taxonomy and distribution ofextant coccolithophorids (calcareou s nannoplankton).
Drukkcrij FEBO B. V, Enschcde: 321 pp.
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Manton , I. 1986 . Functional parallels between calcified anduncalcificd periplasts . In: B.S.C. Leadbeater & R. Riding(Eds). Biomineralization in lower planls and animals. The
Systematics Association Special Volume, No.30: 157-172.
Clarendon Press, Oxford.
Manton, I. & Oates, K. 1985. Calciosoleniaceac (coccolithophorids) from the Galapagos Islands. Unmincraliscd
components and coccolith morphology in Anoplosolenia andCalciosolenia , with a comparative analysis of equivalents inthe unmineraliscd genus Navisolenia (Haplophyceac ~Prymnesiophyccac). Phi/. Trans. R. Soc. London, 8309: 461477.
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R.W: Jordan. A. Wm ter: An unusual eo coli thophorid ........ p. 81 - 83 .

Journal ofannoplankton Research. 20, 2. 1998.

Plate I
ScaJe bars = I ~unFigures 1-3: An unusua l specimen of Calciosolenia murrayi. I. The collapsed coccosphere. 2. Hig h magni fication oftwo types of rhomboliths : a = a narrow. light I calcified rhombolith (note the open laths), and b = a broad, heavily
calcified rhombolith . 3. High magnification of the twisted elongate body-coccoliths. Note the pointed ends.

83