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

SPB Academie Publishing bv, The Hague

The origin of the anchialine cave fauna —

the “deep sea” versus the

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

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 of the

some
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 critically reviewed.

(1986a) are

nomie groups (e.g. the crustacean Remipedia or

Data pertaining to the phytogeny, ecology and biogeography
absent from epigean marine
of the myodocopid ostracods belonging to the Thaumatocyprid- Thermosbaenacea)

idae (Halocyprida) show close affinities between the caver- habitats occur on both old land masses and 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

Résumé Bermuda or the Galápagos Islands, which were

never connected to continental land masses and

Deux hypothèses alternatives qui expliquent l’origine de 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. et celle de Stock (1986a), sont
(1984)
been proposed for such biogeographic patterns.
revues d’une manière critique.

Des données concernant la phylogenie, l’écologie et la bio- Iliffe et al. (1984) when analyzing the fauna of

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 eaux peu 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 that the fauna of anchialine marine caves

In the last twenty years there was a tremendous in- originates from benthic, shallow water, animals

crease in knowledge of the diversity of the subterra- whose colonization of the oceanic islands occurred

fauna for review). in times from the Atlantic is-

nean (see Botosaneanu, 1986, a not so remote (e.g.

It was a surprise to discover the rich 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 described, also new biogeographical pat- ed against additional empirical observations.

terns and hypotheses explaining the and the Despite the inductive character of statements with
origin
distribution of the subterranean fauna emerged. low predictive power, these historical-narrative
 138 D.L. Danielopol -
The origin of the anchialine cave fauna

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

geographical patterns and allow the development of ments have the same low conditions. A
temperature

better research programmes. similar scenario is conceived for the water mites of

The ostracods of the family Thau- the genus

halocyprid Bathyhalacarus (Bartsch, 1988).

matocyprididae are recorded from the Permian to A cladistic of the Mis-

phylogenetical analysis

Recent. As living representatives they are distribut- ophrioida, a group of copepods with species occur-

ed in deep sea and anchialine cave habitats. The two ring either in the deep sea or in marine caves of

alternative hypotheses mentioned will be tested oceanic islands Boxshall that

suggested to (1989)

against the empirical evidence of the Thauma- the colonization route of anchialine subterranean

tocyprididae. habitats islands took crevicular

on place through
corridors which connect the deep sea with the caves.

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

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

connections.

1. The “deep hypothesis” Stock criticized the sea"

sea (1986a, 1986b) "deep
Iliffe et al. (1984) consider that bathyal and/or scenario. He considered that of the
some 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

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

those of volcanic origin) and migrated upwards because extended anoxia

habitats, an period oc-

through the reaching the anchialine curred in the especially during the
macropores oceans, Upper

caves. Documentation of phylogenetical affinities Cretaceous 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 low food There faunistical indications closer

darkness, re- are pointing to

and thermal stability (Margalef, phylogenetical between anchialine

sources, 1976). relationships
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 al., 1983; Hart al., 1985; Manning of Thermosbaenacea which colo-
et et et al., representatives

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

the deep fauna is accepted by these the sea, but have closer affinities littoral in-
cause sea
deep to

authors to be very old, the migrations to anchialine terstitial species (Stock, 1986c).

caves could occur since more than 100 million Stock considers that the anchia-
years (1986a, 1986b)

(Iliffe et al., 1983). Wilkens et al. (1986: 223) con- line cave fauna of the oceanic islands derived from

sider that the of of the endemic animals which lived

origin some oxiphilous or hypoxic during

hypogean fauna of the Islands the the continental shelf.

Canary "probably Neogene on

derived from widely spread deep sea ancestors".

The plausibility that the fauna

deep sea can ex-
Empirical evidence for the Thaumatocyprididae

pand upwards in habitats with similar ecological

conditions can be documented with 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 isopod Ilyarachnidae recorded

sea matocypris orghidani was by Danielopol

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

Fig. 1. Distribution of living and fossil Thaumatocyprididae: Squares =

Thaumatocypris echinata; dots =
Thaumatoconcha, various

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

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

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

the Kornicker & of the Antarctic shelf close

genus Danielopolina 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. Thaumatoconcha polythrix 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 Island and Th. elonga-

Mexico, D. bahamensis and D. mexicana Kornicker 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-Chilean coasts, thus not so far remote from the

also in the shallow crevicular system (2 to 8 m

below Galápagos Islands.

the surface) of Lanzarote, located closely to the Triebel (1941) and Bartenstein found
(1949) two

coast, and up to 300 m inland (Wilkens et al., 1986). thaumatocypridid species (now attributed to the ge-
There is species of viz. D. in the lower Jurassic sedi-
one Danielopolina known, nus Pokornyopsis Kozur)
carolinae Kornicker & Sohn, 1976a, occurring in a ments of & The
Germany (Kornicker Sohn, 1976b).
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 genus Thaumatocypris has only species, tinental and the with well-calcified
one sea ostracods,

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

1100 and 2000 The and

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

sea thaumatocypridid species included in the has rounded With the of the
genus a shape. exception
Thaumatoconcha Kornicker & Sohn, 1976a were pelagic species Thaumatocypris echinata which has

collected in 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

(1976a) and Kornicker (1985). Hartmann Considering these morphological fea-

(1985) processes.

found one female of Thaumatoconcha sp. in 150 m tures one could hypothesize that the species can bur-
140 D.L. Danielopol -
The origin of the anchialine cave fauna

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

while the former has the

spaces
of coarse grain sediments or alternatively families, only one family:

swim free in water above the bottom. Thaumatocyprididae.

in the anchialine caves Kornicker & Sohn consider that the

The Danielopolina species (1976b)

have been caught swimming in free waters (Danie- Thaumatocyprididae derive from the Devonian os-

lopol, 1976; Kornicker & Iliffe, 1989a, 1989c). It is tracod group Checotonomus Kesling (Entomocon-

mention the morphological and eco- chidae). The oldest known thaumatocypridid,
interesting to

analogies of this with the ostracod Thaumatomma piscifrons Kornicker & Sohn,
logical genus

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-

live endo- and epiben- 1941 and T.

Mostof the recent polycopids matocypris” species, T. feifeli Triebel,

thically or in interstitial habitats (Neale, 1983). The bettenstaedtiBartenstein, 1949, now assigned to the

of the genus Metapolycope in anchia- Pokornyopsis, have affinities with Thau-

species occur genus

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

the marine shallow environment outside the caves genera Thaumatocypris and Thaumatoconcha

continuity of sandy form separate phylogenetical lineage within the

presume that
one can a some a

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. Such example is documented by is between 1.4—2.3 living in the sea

gean an range mm) deep
Maddocks & lliffe for Anchistrocheles hart- (Kornicker & Sohn, 1976a; Kornicker, 1985).
(1986)
manni Maddocks, 1976 (Bairdiacea) from an an- Within the genus Danielopolina, D. mexicana

chialine cave 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)

the of Bermuda (Maddocks, and Bellonci organ of the halocyprids have

along coast 1976). a (most

Danielopolina orghidani could also be intersti- this species which show

an organ). Pokornyopsis
tial species which penetrated from the sandy coast phylogenetical affinities with Danielopolina have

into the Grietas Matansas, Cuba, where such sandy of medium size, viz. P. bettenstaedti,
carapaces

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-

Danielopolina the Bellonci be clustered in

mexicana (see further) occurs on void of a organ
can two

continent in the Yucatan Peninsula while the other one living in anchialine caves (D. orghida-
groups,

less primitive cave dwelling Danielopolina species ni, D. styx, D. wilkensi, and D. bahamensis) with

occur on islands of different ages like Cuba, an old small carapace size (0.4 to 0.6 mm length) and a

land Galápagos and which with size

mass, or Lanzarote, are group 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

order with cladistic Jurassic be the origin of the

Halocyprida a analysis two to at thaumatocy-
main the Cladocopina pridids which rise the Danielopolina
phylogenetical groups: con- gave to group.

of the and the Some of the Danielopolina species colonized the

taining the group Polycopacea,

Halocypridina containing the Thaumatocypridacea subsurface environment of the marine caves, while

and the The latter has others migrated into the Strangely
Halocypridacea. superfamily deep sea.
 de Dierkunde, 60 1990 141
Bijdragen tot (3/4) -

because of their cross the

enough, no Danielopolina species display any adap- habitats, inaptitude to

environments. All the Thau- thermocline of the oceanic layered waters

tationto aphotic living zone

blind. But this is due al., 1984, 1985). The Thau-

matocyprididae are not to a (Benson et genus

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

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

less than 200 but none in the

cyprida. waters at m depth,

shallow warm water habitats (Hartmann, 1985).

In conclusion the brief review on the evolution-

and biogeography of the

Discussion ary history, ecology,

does corroborate the

Thaumatocyprididae not

the Iliffe al. hypothesis". The present data suggest

Following et (1984) hypothesis one "deep sea

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

size their from shallow

matoconcha or Danielopolina species of large Danielopolina species water

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

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

ic islands fromthe One would expect these This scenario is in accordance with the "shallow
deep sea.

in Bermuda and the Galápagos Islands water hypothesis" of Stock (1986a).

especially

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

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

from anchialine derive to Jan H. Stock

Danielopolina species caves Homage

from deep sea ancestors which spread through

crevicular systems be in the of The stimulated by Stock's ideas

can questioned light present note was

the and of these and his research on the subterranean

morphology ecology species. Why (1986a) by

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

miniaturized (small) roborate Stock's "shallow water" hypothesis. But

carapace size evolve carapaces

as we see
in the anchialine cave dwelling Danielopo- I consider that the best homage that I can bring to

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

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

Such the explanation of the diverse and original subsur-

cea. an example is, e.g., the remipedian La-

sionectes entrichoma & 1986; it has face fauna of oceanic will be fur-
Yager Schram, aquatic islands)

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

A is that the late people intensify the search of Thau-

large! more parsimonius explanation to

anchialine caves have been colonized by shallow matocyprididae. If my intuition is correct, then one

water dwelling Danielopolina with medium or 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

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

tion low ed that the interstitial Turbellariafrom the Ca-

to 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-

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

in the its interstitial Xestobleberidae occurring on volcanic

pecially Atlantic, changed drastically com-

position. During the last 40 million years

the deep islands like Galápagos in the Pacific, and Stromboli

sea benthic ostracod fauna, mainly in the Atlantic, in the Mediterranean.

is characterized These do The possibility of a combination of dispersal

by psychrophilic species.

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

conceived for marine interstitial

persal by rafting is Benson, R.H., 1984. Estimating greater paleodepths with ostra-

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

press). We need certainly more field observations

1334-1336.

and studies considering subsurface

experimental 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).
oped in many of us both excitement and wonder.
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: 76) when he looked retrospectively to his
Boxshall, G. A., 1989. Colonisation of inland marine caves by

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

confronted with the Danielopol, D.L., 1972. Sur la présence de Thaumatocypris or-
gists as well, 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. Comparative morphology 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 origin and distri-

bution of the interstitial Ostracoda of the species

Some of the ideas here took after discussions group
presented shape
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

C. Juberthie L. Kornicker and geogra-
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Fuchs, T., 1894. Über Tiefseetiere in Höhlen. Annin. k.-k.

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