Bijdragen tot de Dierkunde, 60 (3/4) 137-143 (1990)

SPB Academie Publishingbv, The Hague

The origin of the anchialine cave fauna — the “deep sea” versus the

“shallow water” hypothesis tested against the empirical evidence of the

Thaumatocyprididae (Ostracoda)

Dan L. Danielopol

Limnological Institute, Austrian Academy of Sciences, A-5310 Mondsee, Austria

Keywords: Anchialine cave fauna, Thaumatocyprididae, stygobionts

Abstract For the historical biogeography a very intriguing

problem arose concerning the fauna of these an-

Two alternative hypotheses explaining the origin of some of the chialine caves of oceanic islands. As Iliffe et al. oceanic anchialine cave faunas, viz. those of Iliffe et al. (1984) (1984) noted, many ancient and/or primitive taxo- and Stock (1986a) are critically reviewed. nomie groups (e.g. the Remipedia or Data pertaining to the phytogeny, ecology and biogeography absent from epigean marine of the myodocopid belonging to the Thaumatocyprid- Thermosbaenacea)

habitats both old land and idae () show close affinities between the caver- occur on masses young nicolous and the marine shallow water dwelling representatives. oceanic islands. It is difficult to explain how such

"living fossils" dispersed to oceanic islands, like

which Résumé Bermuda or the Galápagos Islands, were

never connected to continental land masses and

Deux hypothèses alternatives qui expliquent l’originede certains which are separated from the continental coasts éléments faunistiques des grottes anchialines océaniques, c’est- through deep oceans. Different hypotheses have à-dire celles d’Iliffe et al. (1984) et celle de Stock (1986a), sont been for such revues d’une manière critique. proposed biogeographic patterns.

Iliffe al. when the fauna of Des données concernant la phylogenie, l’écologie et la bio- et (1984) analyzing géographie des Ostracodes Myodocopides appartenant aux the Jámeos del Agua cave, on Lanzarote, Canary Thaumatocyprididae (Halocyprida) indiquent des affinités Islands, proposed that some of the troglobionts oc- étroites entre les Thaumatocyprididés cavernicoles et les repré- curring on oceanic islands of volcanic origin could sentants marins vivants dans des eauxpeu profondes en dehors originate by immigration of deep sea elements liv- des grottes.

ing below 200 m depth through crevicular systems

(sensu Hart et al., 1985: 291). Stock (1986a, 1986b)

Introduction challenged the hypothesis of the "deep sea" and

suggested thatthe fauna of anchialine marine caves

In the last twenty years there was a tremendous in- originates from benthic, shallow water, crease in knowledge of the diversity of the subterra- whose colonization of the oceanic islands occurred

fauna in from the Atlantic nean (see Botosaneanu, 1986, for a review). not so remote times (e.g. is-

It was asurprise to discover therich marine fauna lands in the last 30-40 million years). Both which lives in anchialine caves (sensu Stock et al., hypotheses are historical-narrative explanations

1986: 9) of oceanic islands. But not only new taxa (sensu Bock, 1981) and their validity should be test- have been also ed described, new biogeographical pat- against additional empirical observations. terns and hypotheses explaining the origin and the Despite the inductive character of statements with distribution of the subterranean fauna emerged. low predictive power, these historical-narrative 138 D.L. Danielopol - The origin of the anchialine cave fauna

scientific for the bio- models offer us explanations waters from a deep sea stock, because both environ-

and allow have the low conditions. A geographical patterns the development of ments same temperature better research programmes. similar scenario is conceived for the water mites of

of the Thau- The halocyprid ostracods family the genus Bathyhalacarus (Bartsch, 1988). matocyprididae are recorded from the to A cladistic phylogenetical analysis of the Mis-

Recent. As living representatives they are distribut- ophrioida, a group of copepods with species occur- ed in deep sea and anchialinecave habitats.The two ring either in the deep sea or in marine caves of alternative hypotheses mentioned will be tested oceanic islands suggested to Boxshall (1989) that against the empirical evidence of the Thauma- the colonization route of anchialine subterranean

habitats islands tocyprididae. on took place through crevicular

corridors which connect the deep sea with the caves.

However, noted there is Newman (1985) that up to

The alternative hypotheses under test today no clear evidence for such direct crevicular

connections.

The 1. “deep sea hypothesis” Stock (1986a, 1986b) criticized the "deep sea"

Iliffe al. consider that and/or scenario. considered et (1984) bathyal He that some of the examples abyssal organisms penetrate into the subsurface given by Iliffe and his associates (e.g. the amphi- habitats through crevices. Deep sea biota colonized pods Paradaliscidae) are based on taxa that could

in crevicular systems formed solid rocks (especially not have a long history in the bathyal/abyssal

of and those volcanic origin) migrated upwards habitats, because an extended anoxia period oc-

the the anchialine curred in the through macropores reaching oceans, especially during the Upper caves. Documentationof phylogenetical affinities and/or the Paleocene (Benson, 1979, between deep sea and marine cave taxa exists since 1984; Arthur & Schlanger, 1979). Therefore, their more than 100 years (Fuchs, 1894; Racovitza, 1912; origin should be better searched in a shallow water

Vandel, 1965; Riedl, 1966; Margalef, 1976). Migra- fauna. tion of organisms from the deep sea to shallow water anchialine caves is considered possible be- 2. The “shallow water hypothesis”, Stock’s (1986a) cause of the similarities between these two types of scenario environment like total darkness, low food re- There are faunistical indications pointing to closer sources, and thermal stability (Margalef, 1976). phylogenetical relationships between anchialine

Evidences for this hypothesis have been provided cave taxa and marine taxa living outside the caves by Iliffe and/or his associates in various papers (Il- in the shallow water zones. For instance, the iffe et al., 1983; Hart et al., 1985; Manning et al., representatives of Thermosbaenacea which colo-

1986; Wilkens et al., 1986; Boxshall, 1989). Be- nize oceanic anchialine caves have no relatives in

the fauna is cause deep sea accepted by these the deep sea, but have closer affinities to littoral in- authors to be very old, the migrations toanchialine terstitial species (Stock, 1986c). caves could occur since more than 100 million Stock years (1986a, 1986b) considers that the anchia-

(Iliffe et al., 1983). Wilkens et al. (1986: 223) con- line cave fauna of the oceanic islands derived from sider that the origin of some of the endemic oxiphilous or hypoxic animals which lived during

fauna of hypogean the Canary Islands "probably the Neogene on the continental shelf. derived from widely spread deep sea ancestors".

The plausibility that the deep sea fauna can ex- Empirical evidence for the Thaumatocyprididae pand upwards in habitats with similar ecological conditions can be documentedwith analogical ex- 1. Ecology and biogeography (Fig. 1)

& amples. For instance, Hessler Thistle (1975) con- The first anchialine cave thaumatocypridid Thau- sider that some deep sea isopod Ilyarachnidae matocypris orghidani was recorded by Danielopol

able the shallow from (Janiroidea) were to invade polar (1972) Cuba. This species is now attributed to Bijdragen tot de Dierkunde, 60 (3/4) - 1990 139

1. Fig. Distribution of living and fossil Thaumatocyprididae: Squares = Thaumatocyprisechinata; dots = Thaumatoconcha, various

species (A = T. elongata, B = T. polythrix); black stars = anchialine cave species, white star = deep sea dwellingspecies (1 = Danie-

lopolinaorghidani, 2 = D. carolinae. 3 = D. wilkensi, 4 = D. mexicana 5 = D. bahamensis, 6 = D. hemicircle = Thaumatom- , styx);

mapiscifrons; triangle = Pokornyopsis, two fossil species. (Map adapted from Kornicker & Sohn, 1976a; Kornicker & Iliffe, 1989a, c.)

the Kornicker of the Antarctic shelf close genusDanielopolina & Sohn, 1976a. depth to King George Is-

Four other species of Danielopolina occur in an- land. All these species are epibenthic dwelling os- chialine caves, viz. in Lanzarote (Jámeos del tracods. Thaumatoconchapolythrix Kornicker &

Agua), Canary Islands, D. wilkensi Hartmann, Sohn, 1976a, occurs in the deep sea (below 2000 m

1985; in the Bahamas and the Yucatan Peninsula, depth) not far from Bermuda Islandand Th. elonga-

Mexico, D. bahamensisand D. mexicanaKornicker ta Kornicker & Sohn, 1976a, was found at

& Iliffe, 1989a;and finally in the Galápagos Islands, 3750-4125 m depth in the South Pacific off the

D. s/j'-X'Kornicker&Iliffe, 1989c. D. wilkensi occurs Peru-Chileancoasts, thus not so far remote from the also in the shallow crevicular 8 below Islands. system (2 to m Galápagos the surface) of Lanzarote, located closely to the Triebel (1941) and Bartenstein (1949) found two coast, and up to 300 m inland (Wilkens et al., 1986). thaumatocypridid species (now attributedto the ge-

There is viz. one species of Danielopolina known, D. nus Pokornyopsis Kozur) in the lower sedi- carolinae Kornicker & Sohn, 1976a, occurring in a ments of Germany (Kornicker & Sohn, 1976b). The benthic deep sea zone off the Brazilian coast in the paleo-environment is a silty sediment deposited in

Atlantic Ocean at 3459 m depth. shallow waters not deeper than 200 m of an epicon-

The has genus Thaumatocypris only one species, tinental sea and the ostracods, with well-calcified

echinata which benthic forms Thaumatocypris Müller, 1906, was carapaces, are (see also Neale, 1983). found in the pelagic waters of Indonesia between The Thaumatocyprididae have antennae with

1100 and 2000 m depths (Poulsen, 1969). The deep long natatory setae and a strong furca. The carapace

included sea thaumatocypridid species in the genus has a rounded shape. With the exception of the

Thaumatoconcha Kornicker & Sohn, 1976a were pelagic species Thaumatocypris echinatawhich has collectedin the Indian, Atlantic, and Pacific Oceans on the carapace very long tubular processes, the and have been described by Kornicker & Sohn other Thaumatocyprididae species have short blunt

and Kornicker these (1976a) (1985). Hartmann (1985) processes. Considering morphological fea- found female of in could that the one Thaumatoconchasp. 150 m tures one hypothesize species can bur- 140 D.L. Danielopol - The origin of the anchialine cave fauna

fine sediment live in interstitial clustered in several row into the and/or many pelagic representatives,

of coarse sediments or alternatively families, while the former has only one family: the spaces grain swim free in water above the bottom. Thaumatocyprididae.

The Danielopolina species in the anchialinecaves Kornicker & Sohn (1976b) consider that the have been caught swimming in free waters (Danie- Thaumatocyprididae derive from the os- lopol, 1976; Kornicker & Iliffe, 1989a, 1989c). It is tracod group Checotonomus Kesling (Entomocon- interesting to mention the morphological and eco- chidae). The oldest known thaumatocypridid, logical analogies of this genus with the Thaumatomma piscifrons Kornicker & Sohn,

Polycopidae. These species have also round cara- 1976a, was found in Permian sediments in Greece paces, strong natatory antennae and a strong furca. (Kornicker & Sohn, 1976a). Two Jurassic “Thau-

Mostof therecent polycopids live endo-and epiben- matocypris” species, T. feifeli Triebel, 1941 and T. thically or in interstitial habitats (Neale, 1983). The bettenstaedtiBartenstein, 1949, nowassigned to the

in anchia- have affinities with Thau- species of the genus Metapolycope occur genus Pokornyopsis, line caves (in Bermuda), in the deep sea and in polar matomma Kornicker & Sohn and with the Recent shallow waters (Kornicker & Iliffe, 1989b). cave dwelling thaumatocypridids of the genus

Because the anchialine caves are in contact with Danielopolina (cf. Kornicker & Sohn, 1976b). The

outside the Thaumatoconcha the marine shallow environment caves genera Thaumatocypris and

of form within the one can presume that a continuity some sandy a separate phylogenetical lineage gravelly or silty sediment habitats existed between Thaumatocyprididae (Kornicker & Sohn, 1976b). the epigean and hypogean systems. The benthic os- These are benthic species of large size - the cara- tracods could therefore easily colonize the hypo- pace has generally more than 1.4 mm length - (the

habitats. is documented is in sea gean Such an example by range between 1.4—2.3mm) living the deep

Maddocks & lliffe (1986) for Anchistrocheles hart- (Kornicker & Sohn, 1976a; Kornicker, 1985). manni Maddocks, 1976 (Bairdiacea) from an an- Within the genus Danielopolina, D. mexicana chialinecave in Bermuda. This is an interstitial spe- appears more primitive than the other species, i.e. cies initially found living in the sublittoral zone it has a carapace of medium size (0.8 mm length)

of of have along the coast Bermuda (Maddocks, 1976). and a Bellonci organ (most the halocyprids

could also be intersti- this which show Danielopolina orghidani an organ). Pokornyopsis species tial species which penetrated from the sandy coast phylogenetical affinities with Danielopolina have into the Grietas Matansas, where such of medium viz. P. Cuba, sandy carapaces size, bettenstaedti, gravelly sediments exist (Juberthie et al., 1977, and 0.8-1 mm length and P. feifeli, 1-1.2 mm length

Orghidan & Juberthie, pers. comm.). (Triebel, 1941; Bartenstein, 1949; Kornicker &

It is interesting that the more primitive species Sohn, 1976a). The other Danielopolina species de-

the Bellonci be clustered in two Danielopolina mexicana (see further) occurs on void of a organ can continent in the Yucatan Peninsula while the other in anchialine caves groups, one living (D. orghida- less primitive cave dwelling Danielopolina species ni, D. styx, D. wilkensi, and D. bahamensis) with occur on islands of differentages like Cuba, an old small carapace size (0.4 to 0.6 mm length) and a land size mass, or Galápagos and Lanzarote, which are group with very large carapace (1.85 mm young volcanic islands. length) composed of the deep sea dwelling D.

carolinae.

2. Phytogeny and evolution Kornicker & Sohn (1976a) consider the shallow

Kornicker & Sohn (1976a) distinguished within the benthic taxa like Pokornyopsis species from the

Jurassic of order Halocyprida with a cladistic analysis two to be at the origin the thaumatocy- main the which rise phylogenetical groups: Cladocopina con- pridids gave to the Danielopolina group.

of the and the Some of the colonized the taining the group Polycopacea, Danielopolina species

Halocypridina containing the Thaumatocypridacea subsurface environmentof the marine caves, while and the Halocypridacea. The latter superfamily has others migrated into the deep sea. Strangely - 141 Bijdragen tot de Dierkunde, 60 (3/4) 1990

because of their cross the enough, no Danielopolina species display any adap- habitats, inaptitude to

of the oceanic waters tationto aphotic environments. All the living Thau- thermocline zone layered

due The Thau- matocyprididae are blind. But this is not to a (Benson et al., 1984, 1985). genus

in regressive evolution which occurred in the caves or matoconcha, for instance, has ten species living

characteristic of the Halo- the and in antarctic cold in the deep sea, but a deep sea one species occurs

in the cyprida. waters at less than 200 m depth, but none

shallow warm water habitats (Hartmann, 1985).

In conclusion the brief review on the evolution-

and biogeography of the Discussion ary history, ecology,

Thaumatocyprididae does not corroborate the

The data Following the Iliffe et al. (1984) hypothesis one "deep sea hypothesis". present suggest

anchialine oceanic islands derived should expect to find in anchialine caves the Thau- that the caves on

size their from shallow matoconcha or Danielopolina species of large Danielopolina species water

that have either surface or interstitial located (more than 1 mm carapace length) could habitats, ones,

oceanic islands. penetrated into the crevicular systems of the ocean- along the continental margins or

This scenario is in accordance with the "shallow ic islands fromthe deep sea. One would expect these especially in Bermuda and the Galápagos Islands water hypothesis" of Stock (1986a).

(Fig. 1). But this is not the case. The contentionof

Wilkens et al. (1986) and Boxshall (1989) that

derive Jan H. Stock Danielopolina species from anchialine caves Homage to from deep sea ancestors which spread through crevicular be in the of The stimulated Stock's ideas systems can questioned light present note was by the morphology and ecology of these species. Why (1986a) and by his research on the subterranean

of oceanic islands. data should a deep sea thaumatocypridid with large fauna Incidentally my cor-

miniaturized roborate Stock's "shallow water" But carapace size evolve (small) carapaces hypothesis. as we see in the anchialine cave dwelling Danielopo- I consider that the best homage that I can bring to

research direc- lina? The crevicular corridors are macroporous sys- this scientist is to show him that the

tion within which he worked so tems which enables the life of many large Crusta- successfully (viz.

the of the diverse and subsur- cea. Such an example is, e.g., the remipedian La- explanation original sionectes entrichoma Yager & Schram, 1986; it has face aquatic fauna of oceanic islands) will be fur-

intend stimu- 22 mm length and the cephalon is more than 2 mm ther followed using theostracods. I to

is the search of Thau- large! A more parsimonius explanation that the late people to intensify

been colonized shallow If intuitionis then anchialine caves have by matocyprididae. my correct, one water dwelling Danielopolina with mediumor small could find representatives of this group along the

inter- continental in interstitial habitats. New in- carapace sizes which live either in surface or coasts stitial habitats. Ecological difficulties with the vestigations of the dispersal mechanisms of os-

"deep sea" hypothesis applied to the Thaumato- tracods to oceanic islands from remote coasts have

looked for. Ehlers cyprididae occur when one considers the specializa- also to be Ehlers & (1980) point-

ed that the interstitial Turbellariafrom the Ca- tion to low temperature in the oceanic deep sea out waters of the benthic ostracods. Benson (1975) nary Islands could originate independently from showed that during the Late Eocene / Early Oligo- continental interstitial taxa. This scenario is dis-

also Bonaduce for cène the thermospheric deep sea ostracod fauna, es- cussed by Danielopol & (in press)

its Xestobleberidae volcanic pecially inthe Atlantic, changed drastically com- interstitial occurring on

last million the islands like in the and position. During the 40 years deep Galápagos Pacific, Stromboli sea benthic ostracod fauna, mainly in the Atlantic, in the Mediterranean. is characterized by psychrophilic species. These do The possibility of a combination of dispersal not colonize the shallow warm waters of the shelf along the continental margins and short passive dis- - 142 D.L. Danielopol The origin of the anchialine cave fauna

persal by rafting is conceived for marine interstitial Benson, R.H., 1984. Estimating greater paleodepthswith ostra-

polychaetes (Westheide, 1977; Westheide& Rieger, codes, especially in past thermospheric oceans. Palaeogeogr.

Palaeoclimat. 48: 107-141. could Palaeoecol., 1987). Such a possibility of dispersal apply to Benson, R.H., R.E. Chapman& L.T. Deck, 1984. Paleooceano- ostracods as well (Danielopol & Bonaduce, in graphic events and deep-sea ostracodes. Science, 224: We need field observations press). certainly more 1334-1336.

and experimental studies considering subsurface Benson, R.H., R.E. Chapman & L.T. Deck, 1985. Evidence

dwelling Ostracoda! from the Ostracoda of major events in the South Atlantic and

world-wide over the past 80 million years. In: K.J. Hsii&H.J. Finally, 1 want to stress that the investigation of Weissert (eds.), South Atlantic Paleooceanography: 325-350 the subterranean fauna of oceanic islands devel- (Cambridge Univ. Press, Cambridge). in of both excitement and wonder. oped many us Bock, W.J., 1981. Functional-adaptiveanalysis in evolutionary

The excitement is produced by acquisition of new classification. Amer. Zool., 21: 5-20.

wonder "increases Botosaneanu, L. (ed.), 1986. Stygofauna mundi. A faunistic, knowledge. A sense of ever distributional and ecological synthesis of the world fauna in- more with the development of our knowledge". habiting subterranean waters (including the marine intersti- This has been noted by A. Einstein (quoted by Hol- tials): 1-740 (E.J. Brill, Leiden). ton, 1986: when he looked to his 76) retrospectively Boxshall, G.A., 1989. Colonisation of inland marine caves by

fieldof research, but the remark applies to us biolo- misophrioid copepods. J. Zool., Lond., 219: 521-526.

with the Danielopol, D.L., 1972. Sur la présence de Thaumatocypris or- gists as well, confronted unexpected

ghidani n. sp. (Ostracoda, Myodocopida) dans une grotte de diverse and interesting subsurface aquatic fauna

Cuba. C. r. hebd. Séanc. Acad. Sei., Paris, 247: 1390-1393. that we discovered in the last years. Danielopol, D.L., 1976. Comparativemorphology of the deep

sea Thaumatocypris echinata G.W. Müller 1906, and the cave

species Thaumatocypris orghidani Danielopol, 1972 (Os-

Acknowledgements tracoda, Myodocopida). Vie Milieu, (C) 26: 9-20.

Danielopol, D.L. & G. Bonaduce, in press. The originand distri-

bution of the interstitial Ostracoda of the species Some of the ideas presented here took shape after discussions group Xestoleberis arcturi Triebel (Crustacea). Cour. Forsch.-Inst. and written correspondence I had with R. Rouch (Moulis), J. Senckenberg, 114. Notenboom (Bilthoven), T. Iliffe (Bermuda), the late T. Or- Ehlers, B. & U. Ehlers, 1980. Zur Systematik und ghidan (Bucharest), C. Juberthie (Moulis), L. Kornicker and geogra- phischen Verbreitung interstitieller Turbellarien der T.B. Bowman (Washington, D.C.). My visit to the Smithsonian

Kanarischen Inseln. Mikrofauna Meeresboden, 80: 1-23. Institution (Washington, D.C.) was possible due to D. Culvers' Fuchs, T., 1894. Über Tiefseetiere in Höhlen. Annin. k.-k. (Washington, D.C.) arrangements. Thanks are also due to F. naturh. Hofmus. Wien, 9: 54-55. Pieters (Amsterdam) and three anonymous reviewers. A. M.

Hart, Jr. C.W., R.B. Manning& T.M. Iliffe, 1985. The fauna Henning and I. Gradl (Mondsee) typed the manuscript.

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