Contributions to Zoology, 65 (2) 79-99 (1995)
SPB Academic Publishing bv, Amsterdam
New perspectives on the evolution of the genus Typhlatya (Crustacea,
Decapoda): first record of a cavernicolous atyid in the Iberian Peninsula,
Typhlatya miravetensis n. sp.
Sebastián Sanz & Dirk Platvoet
1 Unitat d'Ecologia, Facultat de Ciències Biologiques, Universitat de Valencia, E-46100 Burjassot,
2 Valencia, Spain; Institutefor Systematics and Population Biology (Zoological Museum, Amsterdam),
University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands
Keywords: Typhlatya, Decapoda, Spain, subterranean waters, systematics, zoogeography, vicariance,
evolution, key to genus
Abstract historia geológica de la zona y la distribución mundial del
género, del grupo de géneros, y la familia.
On several occasions, shrimps belonging to a new species ofthe
genus Typhlatya were collected in a cave in the province of
Castellón, Spain. This is the first record of the in the genus Introduction Iberian Peninsula. The species is described and the validity, dis-
tribution, and zoogeography of the genus, as well as the status In 1993 and were on several of the discussed. 1994, shrimps caught genus Spelaeocaris, are Former models for the occasions in in the evolution of the genus Typhlatya and its genus group are re- a cave near Cabanes, province
viewed, as well asthe system ofinner classification of the Atyidae of Castellón, eastern Spain. The specimens belong
and its For the and evolution of biogeographical meaning. age the to genus Typhlatya Creaser, 1936, a genus the genus we developed a new model based on vicariance prin- with members known from the Galápagos Islands, ciples that involves further evolution of each species after the Ascension and the Caribbean of the ancestral This allows estimations Island, Bermuda, disruption range. new
for the of area Dominican Mona age the genus. Accordingly, we suppose that other (Mexico, Cuba, Republic,
proposals, such as recent dispersal through the sea, should be Island, Barbuda, Puerto Rico, Caicos Islands,
disregardedfor this genus. Theevolutionary developmentofthis Bonaire, and Curaçao) (cf. Holthuis, 1986; species is discussed in the context of the geologicalhistory ofthe Banarescu, 1990; Stock, 1993). Up to the present, and the world distribution ofthe the area genus, genus group, members of the have not been recorded in and the family. genus
Eurasia.
Several works have dealt with the origin and bio- Resumen geography of the genus group, the genus, or some
of its species (Monod & Cals, 1970; Croizat et al., En diversas ocasiones se capturaron camarones pertenecientes a 1974; Monod, 1975; Rosen, 1976; Iliffe et al., 1983; una nueva especie del género Typhlatya en una cueva de la Banarescu, 1990; Stock, provincia de Castellón (España). Esta constituye la primera cita 1993).
del la Península Ibérica. In of the coherent géneroen La especie es descrita y se dis- spite vicariance models sug-
cute sobre la validez, la distribución la del y zoogeografía gested by several authors cited above, a general género, así como sobre el estatus del género Spelaeocaris. Se agreement on the evolutionary history of the genus revisan anteriores modelos sobre la evolución del género y la is far from being reached. In other works, a recent serie, así como el sistema de clasificación interna de los atiídos
the sea has been considered su la edad evolución dispersal through y significado biogeográfico. Respecto a y & Hart del género, desarrollamos un nuevo modelo basado en princi- (Chace Hobbs, 1969; Peck, 1974; et al., pios de vicariancia considera la evolución de cada que posterior 1985; Stock, 1986). The occurrence of a member of especie después de la del ancestral. Esto disrupción rango per- the genus in the Iberian Peninsula made us recon- mite aportar nuevas estimaciones respecto a la edad del género. sider the differenttheories and formulateour ideas De acuerdo con este modelo, otras propuestas, como dispersión about the reciente evolution of both this new species and the por el mar, deberían desestimarse para este género. Se comenta whole el desarrollo evolutivo de esta especie en base a la genus.
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ischium shorter than merus. Exopodite of Systematic part always
fifth pereiopod reaching beyond ischium (Fig. 4a).
4. Epipodites present on pereiopods 1 to Dactylus Typhlatya miravetensis n. sp. of fifth pereiopod distinctly longer than that of (Figs. 1-4) pereiopods 3 and 4. Pereiopod 5 longest.
First pleopod (Fig. 4b) with short inner ramus; Material examined. - Male holotype, 20.7 mm, female allotype,
no interna Second pleopod (Fig. 18.5mm,and two female paratypes,11.9 and 13.3 mm (coll. no. appendix present.
Rambla de and short ZMA De. 201475, a, b, c). Cave "Ullal de la 4e) with appendix interna appendix mas-
Miravet" between the towns of Cabanes and Orpesa, province culina with nine setules implanted near apex. Pleo- ofCastellón, eastern Spain (UTM coordinates: 30T YK 504447), pods 3 (Fig. 4c) to 5 with subequal rami and appen- chloride 25 30 June 1993. Conductivity 615.2 nmhos; mg/1, pH dix interna. fauna: 7.01; water temp. 24°C. Accompanying Typhlocirolana Uropods (Fig. 4d) long and slender. Diaeresis sp. (Isopoda: Flabellifera).
March 1994. Other material: same locality, 10 specimens, with short spine on posterodistal corner and three
setules. Telson (Fig. 2d) with one pair of lateral
inter- Description of holotype. - Habitus illustrated(Fig. spines and 13 spines on posterior margin,
mixed with small Two la). Length 20.7 mm. Rostrum short, not reaching two setae. longitudinal rows beyond eyestalks, almost absent. Eyes without pig- of setules on dorsal surface. ment. First peduncular segment of antennule as
and third sub- Female: for the second where the long as second combined, flagella Except pleopod,
of females lack the sexual equal in length, betweenhalfand two-thirds body appendix masculina, no length (Figs, la, b), outer flagellum with proximal dimorphism was found. fourteen segments swollen. Stylocerite slender, not
Remarks. - This fits well within the defini- reaching beyond first peduncular segment. Anten- species nal scaphocerite (Fig. le) reaching well beyond tionof the genus(see Creaser 1936; Monod & Cals,
Hobbs absence of on peduncle; distal tooth on straight outer margin at 1970; et al., 1977): spines
antennular 80% of total length. Inner flagellum 35% longer carapace, rostrum never overreaching
first fifth than body. Outer flagellum one-segmented and peduncle and lacking spines, through
with first to fourth pereio- extremely reduced. pereiopods exopodites,
Mandibles with row of hairs between molar and pods with epipodites. incisor (Figs, lc, d). Palp absent. Small projection The species diagnosis is based on a combination
of first maxilla of characters that makes it differentfrom all other present at base of upper lobe (Fig.
of setules base of of the short If), inner margin with row near species genus(see below): very rostrum, of teeth. Inner of lower lobe with absence of ischium and merus strong edge long eye pigment,
of fifth pereiopod setae of varying length. Palp one-segmented with pereiopods not fused, exopodite
well A to the of three small setae near apex. Membranous structure developed. key species Typhlatya
second maxilla is in the at mid-length on palp of (Fig. 2b), given Appendix. near connection between scaphognathite and palp.
First maxilliped (Fig. 2a) with long flagellar lobe, Derivatio nominis. - The specific name, mirave-
de distal lobe of exopodite reduced. Epipodite small tensis, refers to the mediaevalname of "Tinença
located. but distinct, thumb-shaped. Second maxilliped as Miravet", where the cave is illustrated (Fig. 2c). Flagellum of exopodite of third maxilliped (Fig. 3a) with podobranch and small Ischium-merus fusion. - In some species of
the epipodite. Segmentation between basis and ischium Typhlatya, fusion of the ischium and merus in
such indistinct. pereiopods occurs (Table I). In many groups,
like this is Exopodites of pereiopods decreasing in relative as amphipods, a character considered to
length from anterior to posterior (Figs. 3b-e, 4a). be of generic value. This could imply that the genus
be dividedinto six sub- Ischium and merus separate in all pereiopods, Typhlatya might as many as
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Fig. 1. Typhlatya miravetensis n. sp.: a, habitus; b, first antenna with enlarged seta from segment 1 (scales III and V); e, left
mandible (I); d, right mandible (I); e, second antenna with enlarged seta from antennal scale and detail of distal part of peduncle (III);
f, first maxilla (IV).
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Fig. 2. Typhlatya miravetensis n. sp.: a, first maxilliped (scale IV); b, second maxilla (IV); c, second maxilliped (IV); d, telson (II).
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Fig. 3. Typhlatya miravetensis n. sp.: a, third maxilliped (scale II); b, first pereiopod(II); c, second pereiopod (II); d, third pereiopod
(II); e, fourth pereiopod (II).
since characters rank groups. However, other (rostrum, Typhlatya pretneri (Matjasic, 1956) new
eye pigmentation, exopodite P5 reduction) do not
support such a division we consider the genus Typh- The genus Spelaeocaris (S. pretneri Matjasic,
for workable which is found in latya, the time being, as a taxon. 1956), Hercegovina (former
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second Fig. 4. Typhlatya miravetensis n. sp.: a, fifth pereiopod(scale II); b, first pleopod (II); c, third pleopod (II); d, uropod (II); e, pleopod with details (II, IV, V).
Yugoslavia), is distinguished from Typhlatya by the T. pearsei Creaser, 1936, the exopodites of the pe-
of the from P5. reduction exopodites of the pereiopods reiopods are also gradually reduced PI to
(Monod & Cals, 1970). Nevertheless, in theirreview Another character used by Monod & Cals for re-
of the "série Typhlatyienne", these authors did not taining the genus Spelaeocaris is the number of dis-
point out that in Typhlatya monae Chace, 1954and tal spines on the telson. Since their study, two more
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characters in the = 2 = Table I. Fusion of ischium and merus of pereiopods and other genus Typhlatya (exop. exopodite). Keys: (a)
= = = = = s not fused, 1 fused; (b) - eyes unpigmented, + eyes pigmented; (c) l = long, s short; (d) exopodite P5, l = long, short;
distal setae of (e) presence ( + ) of on exopodites pereiopods.
rostrum setae species fusion of ischium and raerus eye exop. P5 exop.
PI, 2 P3,4 P5
campecheae 2 2 2 - s I +
garciai 2 2 2 + s 1
miravetensis 2 2 2 - s 1 +
monae 2 2 2 - s s + pearsei 2 2 1-1 s +
mitchelli 2 12-1 s +
1 1 - 1 galapagensis 2 s
consobrina 12 2+1 1 + rogersi 12 2+1 I +
iliffei 1 1 1 + I 1 + pretneri 111-1 s
Key: (a) (a) (a) (b) (c) (d) (e)
Fig. 5. Map ofthe area in which the cave is located. The solid dot represents the mouth ofthe cave.The crossed dots indicate two karstic
holes that possibly represent the water inlets ofthe system involved. The surface hydrologicalnetwork of the area isrepresentedby dotted
lines. This network is solely formed by temporary rivers. The arrows indicate the direction of the water flow.
species have been assigned to the genus Typhlatya: overlap with the genus Spelaeocaris.
T. iliffei Hart & Manning, 1981 and T. mirave- With this in mind the status of the genus Spelaeo-
reductions tensis. These two bear ten or more distal spines on caris becomes uncertain. Convergent of
the these show occurred in After telson; so, Typhlatya species some exopodites frequently decapods.
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6. Fig. Profile of the cave. The dotted lines with the dates represent the water level at every exploration. The number of shrimps caught
is also given.
the Table II. Environmental variables recorded at the first success- Description of study area
ful sampling in the cave and general geographical information (Figs. 5, 6 and Tables II, IV)
about the cave mouth.
In February 1993, members of the Grupo Espeleo-
UTM coordinates 30T YK 504447 lógico Oropesa del Mar (G.E.O.M.), belonging to
altitude 140-160 m above sea level the Valencian Speleological Federation, collected
distance to the sea two shrimps at the mouth of a cave, called by them
- in direct line 7.75 km "Ullal de la Rambla de Miravet", near Cabanes, in
— by the temporary stream 10.35 km the province of Castellón, eastern Spain. In June water temperature (°C) 24.0 first author descended into the pH 7.01 1993, the cave, ac-
redox potential -5.4 companied by Julio Perpiñá (G.E.O.M. member),
dissolved (mg/1) 6.3 oxygen and collected two further specimens. In February salinity (%o) 0.25 and March 1994, the cave was visited again and conductivity (nmhos) 615.2 several shrimps were caught. total alkalinity (mmol/1) 5.15 is of phenophtaleinalkalinity (mmol/1) 0.0 The cave mouth located S.S.E. of the town
chloride (mg/1) 25.0 Cabanes in eastern Spain (see Fig. 5), next to the
+ + + + total hardness (Ca plus Mg ) 6.45 (meq/1) Cabanes-Orpesa road. The mouth (140-160 m 18.0 (°dH) above sea level) is on thebed of a temporary stream date 30/06/93 known as Rambla de Miravet, or Riu de Xinxilla. hour 9:00 a.m.
flooded Since part of the cave is permanently
the it has been throughout year, only explored par-
tially. In Fig. 6 we given an approximate profile of
of careful examination two specimens of S. pretneri the cave. Before 1992, the entrance was only a nar-
(from Kifinoselo, Hercegovina) in the National row cleft from which water flowed after heavy rain.
Museum of Natural History at Leiden, we decide to Then, G.E.O.M. members removed rocks, which
of them to the The water and consider Spelaeocaris as a junior synonym gave access cave. level
Typhlatya. number of shrimps caught on each occasion are
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Table III. Habitat and geographicalrange, with number of known localities, of the species of Typhlatya. References (both for habitat and morphological features): 1, Hobbs et al., 1977; 2, Holthuis, 1986; 3, Botosaneanu & Holthuis, 1970; 4, Monod & Cals, 1970;
5, Chace & Manning, 1972; 6, Hart & Manning, 1981; 7, Matjašič, 1956.
species habitat distribution No. local. réf.
1 campecheae pools in a cave Campeche, Mexico 1,2 garciai subterranean lake (fresh water) Cuba 2 1,2,3 miravetensis freshwater cave Castelló, Spain 1 present
study monae pools in caves or phreatic wells Mona Is. (Puerto 6 1
(mixo- or mesohaline) Rico), Barbuda &
Dominican Republic
1 pearsei freshwater caves & cenotes Yucatán Peninsula, 13
Mexico mitchelli cenotes (fresh waters) Yucatán, Mexico 9 1 galapagensis from fresh (altitude 50 m, 2 km Galápagos Islands
from the coast) to brackish waters, (Sta. Cruz & Isabela) 5 4
lava pools and caves
consobrina subterranean lake Cuba and Caicos Isls. 3 1,2,3 rogersi anchialine habitat of lava Ascension Island 2 2.5
pools, salinity 35-40%o
1 iliffei anchialine habitat in caves Bermuda 2.6 pretneri caves (fresh water) Hercegovina 2 2,1
2.5 km the northwest Table IV. Results from a water chemistry analysis performed on quick; (3) some to (inland)
March 1994. No “in situ” measured. Con- 7, parameters were there is an endorheic area called Pla de Cabanes ductivity values are given corrected to 20°C. (altitude about 260 m), that does not have surface
drainage, with two big karstic holes in the centre, North siphon South siphon
where rainwater disappears (see Fig. 5). A more chloride (mg/1) 20.0 20.0 detailed description of the area is given in a bulletin alkalinity (meq COj /l) 6.4 5.45 of the "Espeleo Club Castelló" (1985). The cave total hardness described in this is probably a secondary + + paper (Ca + plus Mg+ ) in °dH 20.0 17.0
exit of the same hydrological in meq/1 7.2 6.1 water system. conductivity (timhos) 610.0 550.0 salinity (%o) 0.3 0.2
The non-anchialine character of the cave
indicated in Anchialine Stock also Fig. 6. The water level was quite (see et al., 1986; Iliffe, 1992) variable. Some environmentalparameters recorded stands for bodies of haline water, usually with re-
during the first sampling are shown in Table II as stricted exposure to open air, showing both marine well and terrestrial influences. of as some geographical data. Many species Typhla-
Although the cave is only partially explored, tya occur in such habitats (see Table III).
is sufficient there information to suggest that it is In March 1994, after a period of severe drought,
flooded of the not solely by water infiltrating through the the lowest level since exploration cave began
but direct soil, probably mainly by water penetra- was recorded. Then, the cave was sampled again to tion: (1) even though the water was quite transpar- search for a possible marine influence. For that
the ent, sediment contained organic matter, not, purpose only main water ions were analyzed. The
fine but also seeds and shown only particulate matter, results of these water analyses are in Table fragments of leaves; (2) the response to rain is very IV. No evidence for a marine connection was
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found: neither a tunnel nor water chemistry clues. earliest Tertiary, these sedimentswere uplifted with
The altitude would have caused water to flow to relatively littlefolding. During therest of the Palae-
if the this Iberian the sea a connection existed; however, water ogene, corner of the Peninsula repre-
is is also that sented elevated contained stagnant. It possible complex topogra- an area that many inner
allowed water to in In the basins with to the at phy stay siphons. unlikely poor drainage sea. As present,
event of a marine connection, it is probably of shallow-water conditions (freshwater marshes,
minor and with influ- shallow importance, temporary, no seas) probably have also prevailed in the
ence on the main body of the cave, neither on coastal areas.
faunalmovements, nor on water chemistry changes. Neogene tectonics affected the entireIberian and
the cave is the location within Catalan but this Tectonic Also, entrance only ranges, especially area.
that exhibits a wide area the characteristic water compressions and distensions in E.S.E.-N.N.W.
surging pattern of strong output in rainy periods direction during this period led to the renewed
and none in dry periods. We believe that the mouth uplift of Mesozoic rocks. These rocks now form a
where we penetrated is the main (and probably the series of mountains that run parallel to the coast
exit sole) to the surface of a subterranean drainage (S.S.W.-N.N.E.) from Cabanes, in our study
fed from endorheic The system an upper area. area, throughout the Gandesa/Flix area, some washing of this area by rainfall appears to supply 150 km to the north (Simón Gómez, 1983; Anadón, the the Between these mountains further organic matter supporting ecosystem. Any- 1992). and areas how, the cave does not fit with the definitionof an inland, a series of marshes were trapped. Sediments anchialine habitat. of the Neogene period were limnic. In the Quater-
nary, during a period of sea-level regression in com-
bination with dryer conditions, most of these areas
Geology, geological history, and paleogeography emptied, some of them into the sea, others by being
transversed by rivers (such as the Ebro), or desic-
from sediments carried the Apart some by stream, cated by evaporation. thebed of Rambla de Miravet is made Creta- The Pla de up by Cabanes is a plain area surroundedby ceous materials. The Pla de Cabanes is covered by an area of complex topography (see Fig. 5). The
slender a layer of Quaternary continental deposits area is not transversed by rivers. Surface drainage
older and thicker lying concordantly on Tertiary is very difficult. Since the bed underlying the Ter- limnic endorheic deposits. This depression is sur- tiary and Quaternary deposits is formed by car- rounded mountains formed folded and ele- bonate it is this by by rocks, very likely that drainage of vated Mesozoic marine started deposits (IGME, 1973). area to occur through a subterraneankarstic
This whole area represents a transition between complex that channels water to the mouth we ex-
Iberian and Catalan these ranges. Although ranges plored. have differentorientations, they were caused by the same orogeny, one that affected the entire N.E.
of the Iberian corner Peninsula. They were formed Distribution in the early Tertiary. During this period, tectonic movements related to Alpine orogeny approached Hitherto the family Atyidae, to which Typhlatya the Ebro and rise Iberian, European plates, giving belongs, was represented in the fresh waters of the to the Iberian and Catalan mountain ranges Iberian Peninsula by two members only: Atyae-
(Capote, 1983; Capote & Carbó, 1983; Santanach, phyra desmaresti (Millet, 1831) and Dugastella
Martínez 1983; López, 1989). These ranges had valentina(Ferrer Galdiano, 1924). Both are epigean been ofmarine mainly areas and lacustrine environ- freshwater species. The first is distributed in most
sedimentation ments with strong processes during Iberian basins, draining into both the Atlantic and the Mesozoic, of a character more shallow than the Mediterranean. D. valentina is an endemism of bathyal (Rincón et al., 1983; Canérot, 1991). In the the eastern Iberian Peninsula where it occurs in a
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of the Gulf of Valencia he that fit far better narrow zone in the vicinity points out this genus seems to
(Sanz & Gómez, 1984). in Rosen's model, except for the fact that one spe-
The is known from the cies is known from Ascension island. genus Typhlatya Galápa-
All freshwater but gos Islands, Mexico (Yucatán), Caribbean region, recent Atyidae are species,
Bermuda, and Ascension Island. The genus group, some of them show high salinity tolerances. Hunte
documentedthe larval or "série Typhlatyienne" as defined by Monod & (1976a, b) complete develop-
Cals (1970), consists of the genera Antecaridina, ment of the atyids Mycratya poeyi (Guérin
Typhlatya, the former Spelaeocaris, Typhlopatsa, Méneville, 1856) and yAtya innocouss (Herbst, 1792), and Stygiocaris, and has a disjunct Tethyan distri- reared in the laboratory. He found the optimum
bution: and in the western larval at salinities of 32 and 30%o Typhlatya Spelaeocaris development ,
results he part (Mediterranean region, Caribbean area, and respectively. After these laboratory sug-
Galápagos Islands), while Antecaridina, Typhlo- gested marine stages in the evolution of these spe-
Larval patsa, and Stygiocaris occur in the Indo-Pacific cies and perhaps other Atyidae. hatching
in estuaries has been for striolata area (see Monod, 1975; Holthuis, 1986; Banarescu, suggested Atyoida
1990). (McCulloch & McNeill, 1923) (cf. Smith &
distribu- At present, the new species has only been found Williams, 1982). On the other hand, the
that reached in this particular cave. tion of /4/ya-related genera, remote
islands in the Pacific, certainly does not suggest
vicariance events from old marine ranges (see
Discussion Chace, 1983, for distribution in the Indo-Pacific
region).
of Members of the genus Typhlatya are found in Even ifall recent members the family Atyidae waters ranging from marine (T. rogersi Chace & are freshwater species, the evolutionary euryhalini-
of each Manning, 1972) to completely fresh ((T. miraveten- ty, the biogeographical layout, and the age sis, T. garciai Chace, 1942, T. pearsei), but general- genus do not necessarily have to be the same.
the first who stated that ly they occur close to the sea. Although they show Banarescu (1973) was (part the osmotic ability to adapt to fresh water, Typh- of) the family Atyidae evolved much earlier into
fur- fresh than such the latya species do not seem to be able to penetrate waters groups as palaemonids. ther inland. At first sight the distribution pattern At least some of the atyid species are rather salt- seems to be linked to modern shore lines and seas. tolerant and "the dispersal of some of them (e.g.,
Much confusionand controversy exists about the the circumtropical and those reaching remote archi-
in in recent biogeography of the genus Typhlatya. The genus pelagos the Pacific) occurred probably has been widely used for the development of bio- times" (Banarescu, op.cit.: 19). On the contrary geographical models and theories: Rosen's vicari- "a Tethys origin and age must be assumed" for
and ance model for Caribbean biogeography (Rosen, Spelaeocaris, Dugastella, Troglocaris Palae-
1976) and Stock's reaction to it (Stock, 1986), a monias and perhaps also for Parisia (: 19).
think that short review of classifica- model to explain colonizationof anchialinecaves in We a atyid
Atlantic islands (Hart et al., 1985), and an alterna- tion could throw some light on this apparent con- tive approach by Stock (1993). troversy.
We think that part of the confusion is the result of the assumption that the evolution of Typhlatya
the “series” followed the same path as in other genera. For ex- Classification of Atyidae: concept and ample, in Rosen's model, Typhlatya follows a sim- its biogeographical meaning ilar track (however not exactly the same) as other atyids (Atya) and even palaemonids (Macrobrachi- The main review concerning the family Atyidae is
Stock from Bouvier This author classified the um). (1986) refutes the use of these decapods (1925). because of physiological features like marine dis- genera known at that time into two kinds of persal abilities (see below). However, for Typhlatya "formes": "formeacanthéphyroïde", with its sole
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and "formes authors fullreview of the genus Xiphocaris, atyiennes ty- provided a morphological
features of these clear their piques". He divided the "formes atyiennes ty- genera, making coher- piques" inthree "séries": "paratyienne", "caridel- ence and closest proximity to the Paratya series lienne", and "caridienne".To these forms and ser- rather than the Caridina or Caridiella series. Even ies he gave an evolutionary meaning. The "forme though they suggested that the morphology and dis-
would tributionof the series calls for ancient for acanthéphyroïde" (viz. Xiphocaris) repre- an origin sent the more primitive form, the closest to the Typhlatya galapagensis auct. Monod & Cals (1970) ancestral atyid. Among the typically atyan forms, proposed a recent (Miocene or Pliocene) dispersal the Caridellaand Caridinaseries wouldbe the more by the sea from Central America. evolved and whilst the series Monod modern, Paratya Later, (1975) gave a new biogeographical would represent an intermediate stage. Among the significance to this newly built series. He concluded recent families of the suborder Caridea, he placed that the morphology and distributional patterns
relatives of the closest the family Atyidae withinthe arose from a common and old origin for the mem-
"des known family Acanthéphyrés", nowadays as bers of this series related to Triassic paleogeo-
Oplophoridae. In fact, Oplophoridae and Atyidae, graphy, refuting his older work by including Typh- together with Nematocarcinidae, often have been latya galapagensis in this scheme. grouped within the same superfamily, named Banarescu (1990) revised the series concept, es-
Atyoidea (Bowman & Abele, 1982) or Oplopho- tablishing a different biogeographical layout for roidea (Schram, 1986). Nevertheless, Chace (1992) each of the four series. He assumed a tropical origin placed these three familiesinto differentsuperfami- for the entire family, relating the distribution of lies. He also removed Xiphocaris (the only represen- each series to the position of the tropical zone and tativeof the "forme acanthéphyroïde" in Bouvier's the distributionof seas and land masses during the
from scheme) the Atyidae and elevated it to a new period at which the ancestors of each series left the
series family, Xiphocarididae. He placed this new family sea. For example, the Atya/Caridina (Bou- within the superfamily Nematocarcinoidea. vier's "série caridienne") is pantropical and more
From the publication of Bouvier's work until or less corresponds to the present-day position of
number of 1970, a new atyid genera have been the tropical zone (op.cit.: 511). He assumed that described: Candines Calman, 1926, Antecaridina this series didnot leave the sea untilat least the Plio-
1954 Mesocaris the the Edmonson, (= Edmonson, 1935), cene or Pleistocene (: 250). In case of Typh-
Typhlatya Creaser, 1936, Parisia Holthuis, 1956, latya series (Monod & Cals' "série Typhlatyienne")
Typhlopatsa Holthuis, 1956, Spelaeocaris Matjasic, matters would be different; the whole series would
Potimirin 1956, Holthuis, 1957 and Stygiocaris have a "Tethyan disjunct range" (: 238).
1960. Some of these new dif- Holthuis, generawere Following Banarescu (1990), both the Paratya ficult to place within Bouvier's scheme. Typhlatya and Typhlatya series would have been derived from was put into the "Caridinienne" series by Creaser ancient marine stocks. Since these stocks were
Holthuis (1936). (1956: 98) stated that if one strictly formed by shallow sea populations, the separation
Bouvier's follows key, Typhlatya, Antecaridina, of land masses which characterizes the Mesozoic and would fall within the "série cari- Typhlopatsa period made the prawns "land-locked when the
but these three land while those which remained marinelater dellienne", generaare perhaps more rose,
related the of Bouvier's "série became extinct" For of closely to genera (: 250). the case Typhlatya, paratyienne". Later, Holthuis (1965) paid atten- he stated: "In a similar manner the disjunct range tion to the homogeneity of these genera and their of Typhlatya (Antilles, Yucatán, Galapagos and peculiar in Bouvier's classification and place pro- Ascension Islands) can be explained by coloniza-
the idea of Monod & Cals tion of brackish posed a separate series. the and freshwater caves by
(1970) based their "série Typhlatyienne", com- representatives of the same marine stock that in- posed of Typhlatya, Spelaeocaris, Antecaridina, habited the shallow waters, and not by direct con-
and and Typhlopatsa, Stygiocaris on this idea. These tinental contact or drifting of Galapagos
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Ascension Islandfrom the Caribbeanarea" (: 250). freshwater lineage would depend on: (1) the posi-
It should be kept in mind that these supposed dis- tion of shore lines at the time that evolution
the from this "ma- towards fresh started take the persal abilities concern species waters to place; (2)
rine stock", viz., the ancestors of therecent species, marine distribution of these ancestors; (3) the
and not the recent (cavernicolous) species them- length of the period of transition.
while the selves. Here we want to point out that, facing
problem of explaining the distribution of animals
like Atyidae, or at least its older series, we should
Family framework better think in ancient coastal marine patterns
rather than pelagic marine dispersal, an unlikely
Relatively little is known about the origin and bio- supposition, at least for long distances (transocean- geography of the Atyidae, except for the fact that ic dispersal) and/or for lineages adapted to fresh
of each series different if faces the development seems to be waters for a long time. Furthermore, one
(Banarescu, 1973, 1990). the explanation of the distribution of a genus or
almost in fresh it is the series the "série Recent Atyidae range exclusively species, (e.g., Typhla- waters of both tropical and, considerably less, tem- tyienne") that is the highest possible framework in
Their marine believed which perate regions. ancestors are we can move.
(Bouvier, 1925) to be related to the family Oplo- phoridae, a groupof bathyal shrimps. These primi- tive marine Atyidae probably occurred in benthic Evaluation of former proposals regarding the
the coastal habitats, along warm seas. From here, they evolution of genus wouldhave given rise to the present four freshwater lineages of the Atyidae. Croizat, Nelson & Rosen ( 1974) dealt with the distri-
bution of of their "vicari- In contrast, recent Palaemonidaehave at least as Typhlatya as a paradigm
model" of distribution many marinerepresentatives (Palaemon, Pontonia) ance viz., as apattern species
freshwater and from the of old distribu- as brackish-water genera (Macro- arising fragmentation an brachium, Palaemonetes). Following Banarescu tional range. Rosen (1976), studying the so-called
(1990: 249-251), the distribution of most fresh- "tracks" for the Caribbean area, established four water lineages within this family corresponds with kinds of "transoceanic generalized tracks", among the recent distribution of sea and land. Further- which the Typhlatya track represents the oldest.
This track would have been derived from more, freshwater palaemonids are more generally an old,
late marine 1976: associated with coastal areas, while atyid shrimps Mesozoic, range (see Rosen, figs. tend to be found more in rivers and inland. 5E, 6F). He came to this since the Typhlatya track
The question when this evolution of Atyidae into was both amphi-American (Galapagos - Caribbean) fresh waters took place and how simultaneously and amphi-Atlantic (Caribbean-western Africa). these for the different series Our of the in the Mediterranean developments were discovery genus remains open. As discussed above, the situation region agrees with these conclusions, but, as willbe
discussed seems far from being coherent and a model cannot below, we place the origin earlierthan late
Cretaceous. the think that be provided for the entire family. If one accepts the At same time, we only present situation of palaemonids as a kind of model now, after the discovery of the genus in Europe, is
it kind of track for past developments in atyids, it becomes clear possible to assign this for Typh-
the from the basis that evolutionary processes are far being latya. Rosen developed his ideas on of the
of T. in "West synchronous: amongpalaemonids some groups are occurrence rogersi Africa", mean-
freshwater Ascension true species whilst other groups consist ing Island, on which this species occurs.
is of marine, coastal, or intermediatespecies (brack- This island located west of the Mid-Atlantic ish, estuarine). Ridge and probably more related to America than
from of As a result, the distributional patterns of each to Africa a tectonic point view. Other
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from Hart al. authors also supported vicariance models, but A more dispersalist model came et
distributional while the fauna of Atlantic based on morphology and the pat- (1985) who, reviewing terns of the entire series (Monod & Cals, 1970; caves, proposed for most of the taxa a dispersive
Monod, 1975; Banarescu, 1990). evolutionfrom (very old) ancestors, mainly bathyal
these marine in their All models would correspond to a or pelagic. Including Typhlatya species ancestor inhabiting shallow waters in the Mesozoic. review, they came to conclude: "however, it is even
The later disruption of its range would have led to more difficult to create a scenario for their disper- isolation of the populations due to a low capacity sal" (op.cit.: 289). As will be discussed below, we for dispersal. Those races would have given rise to do not think that this is the case in Typhlatya. the present cavernicolous species as well as to sur- The model provided by Stock (1993) seems to fit face forms that later became extinct. As discussed better the Typhlatya situation. This author pays below, these proposals agree with our findings. attention to the fact that "many stygobionts have
of in Atlantic relatives in shallow marine and The occurrence Typhlatya islands, congeneric waters",
in islands like Ascension he the occurrence of several includ- especially young Island, explains taxa, led some authors to different conclusions. Some ing Typhlatya, on Atlantic islands: "the young is- based their ideas on dispersion, others on the exis- lands must have existed as shallow banks or sea-
in when the Atlantic started tence of ancestral populations as "marine stocks". mounts the period to
Chace Hobbs and Peck and before the of the Sea" & (1969) (1974), dealing open, disruption Tethys with the Antilles, stated that the species dispersed (Stock, 1993: 807). It is important to pay attention from Central America to these islands in recent to the two paleogeographical conditions argued by times. To explain the occurrence of Typhlatya in this author, since they are important to establish an
estimated for the the AtlanticOcean Ascension Island, Stock (1986) gave two options: age genus: was
(1) even though the island is young, it is theremnant opening and the Tethys was not completely closed. of an older archipelago; (2) the genus has marine Our findings agree with this, especially for Typh-
dispersal abilities at a larval stage. Since most latya. Furthermore, this takes the Typhlatya track
Typhlatya species described at the time (but certain- from Stock's third track (amphi-atlantic/eastern ly not all) were from anchialine habitats (connected Pacific) to an older track (amphi-atlantic/eastern with the sea), he supported the second option, Pacific/Mediterranean). Further development of without disregarding the first. Chace & Manning these ideas can be found in Stock (1994).
(1972), referring to the species from Ascension,
that due of pointed out to the age the island T.
must rogersi be a newcomer in these caves, which Troglobiosis did not disperse from the continentbut originated from a "marine stock". They didnot point out how Withregard to the occupation of freshwater subter- ancient this stock might be, if it was bathyal or ranean habitats by crustaceans, several mechanisms
with pelagic, or without dispersal capabilities, and and evolutionary scenarios have been suggested. how its geographical distributionwould have been. For interstitial and crevicular subterranean fauna
Iliffe et al. (1983), dealing with this problem and the proposed models (Stock, 1980; Coineau & with the occurrence of T. iliffei in Bermuda, stated Boutin, 1992; Notenboom, 1991) assume troglo- that both species could originate from a marine bites evolved from marine (coastal) interstitial or-
of shoreline stock associated to the Mid-Atlantic Ridge, where ganisms thatoccurred in areas frequent they had persisted since "the separation of the Afri- changes. However, free-swimming freshwater ani- can and American continental masses". They did mals from caves may not have evolved directly from not think that this stock was as ancient as Tethyan, marine ancestors, but rather from surface fresh- even though they stressed that in these caves true water forms (Barr & Holsinger, 1985; Holsinger,
Tethyan relicts occur, as the hippolytid shrimp 1988).
Somersiella sterreri Hart & Manning, 1981. A differentexplanation can be given for animals
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occurring in former anchialine habitats. It is pos- (Stampfli et al., 1991). The progressive breakup of
sible that anchialine environments could have lost Pangea reached the Iberian Peninsula during the
with for mid Triassic. their connection the sea, example by sea-
level changes. If freshwater influenceoccurred, this In the early Jurassic, a rift valley system was
would have become the organizing factor for the developing in the central North Atlantic (Jansa,
system. In this case, it is not a surface population 1991). As a result, in the Pliesbachian (194-200 mY
evolved towards subterranean marine America that a species, but ago) an incipient passage through
rather it is the cave itself that changed from marine was opened, connecting the western Tethys and the to freshwater conditions. Subsequently, the popu- eastern Pacific (Riccardi, 1991). From this point
either osmotic barriers the Central lations break the or they dis- and during the rest of the Jurassic,
appear. Since true anchialine habitats are usually Atlantic continued to widen, resulting in a change
variable in fresh- of circulation from estuarine quite salinity (depending on tides, pattern an to an water input, etc.), the hypothesized populations "open channel" type (Jansa, 1991). The Iberian
in of this may have been pre-adapted to change (Hutchinson, Peninsula was the middle passage, sur-
Iberia and 1960; Peck, 1974). rounded by a channel between Europe
(Biscay Seaway) and another between Iberia and
Africa (Rift Seaway) (see San Román & Aurell,
A modelfor the evolution of the genus 1992; Schwentke & Kuhnt, 1992; Reicherter et al.,
1994).
Now, let us try to provide a model for theevolution In the early Cretaceous, the Central Atlantic- of the genus largely based on the ideas exposed by Caribbean connection between the Mediterranean the authors citedin previous sections. We based our Tethys and the eastern Pacific was fully opened, model on Tethyan paleogeography and paleocea- and a connection with the Arctic by the just opened
the of the North Atlantic made nography, ecology recent species, and on was (Scotese, 1991). As a con-
of of in finding a member the genus Europe, all sequence, following Winterer (1991: 261): "the within the "series" framework. circum-global west-flowing equatorial current sys-
The occurrence of the genus in Europe is a strong tem established in Late Jurassic time was now less clue to link the origin of the genus Typhlatya to restricted through the Mediterranean, Central At- the Tethys Sea, following Monod, Banarescu, and lantic and Caribbean, and the most typical of all
Stock. The ancestor probably was a marine, coastal Tethyan faunas, the rudists, spread widely around
in low latitudinal such the the from about Also in shrimp, ranging seas, as globe, 30°N to 30°S". the
Mediterranean, the mid Atlantic, the Gulfof Mexi- early Cretaceous, the South Atlantic started its rift-
and the eastern Pacific western of the from south to north. This co (the part ing phase, seaway was
Tethys Sea). These seas covered a much smaller not opened until the Albian-Cenomanian (90 mY area during the Mesozoic than at present. The only ago) (Riccardi, 1991; Scotese, 1991). Later, in the living descendants of this species are found in cave Turonian, faunal exchanges between the opened habitats. Monod (1975) related thepresent range of South Atlantic and the Mediterranean-Central
Triassic Typhlatya to paleogeography. The paleo- Atlantic-Caribbean-East Pacific system, were geographical reconstructions we have consulted do possible both through the establishment of a cur- not support this. In order to determinethe period rent system in the whole Atlantic and the Sahara at which this ancestor may have occupied its con- Seaway. The late Cretaceous paleogeography (from
tinuous before it look into Santonian - range was disrupted, a to Maastrichtian, 87 65 m Yago), marks
Tethys paleogeography is needed. the acme of the global Tethyan Ocean. From here
In the early Triassic, when the Pangea supercon- on the trend is toward constriction in low latitudes tinent started to break up, the western part of the (Winterer, 1991).
Tethys Sea more or less corresponded with the We think that the maximal development of the present eastern Mediterranean-Middle East area ancestral range of Typhlatya should be situated
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somewhere in the Turonian (where the South At- nellacea), and cyprinodontiform fishes (for a re-
latest lantic was opened) and the Maastrichtian/ view, see Banarescu, 1991).
is that the the early Cenozoic (end of the global Tethys). It most We suppose split up of distributional likely that these ancestors existed before this period area of the ancestor into several isolated areas (both
of (ranging in the Mediterranean-Caribbean-East- by plate tectonics and by the change global cur-
more or less simultaneous- ern Pacific) and it is then that they achieved their rent systems) happened
does that further de- maximal expansion, once they had colonized shal- ly. This not mean evolutionary
of each of these either low areas in the South Atlantic. It is not very likely velopment populations was
the much earlier than this simultaneous of the kind. Once iso- that genus developed or even same
lated in coastal period: there are two genera of the series in the populations, separated by deep western Tethys ((Typhlatya and the former Spelaeo- seas, each local population could have evolved in
and three in the different conditioned and caris) more eastern Tethys (Ante- ways by: (1) physical geo- caridina, Typhlopatsa, andStygiocaris). logical factors: presence of caves (limestone or lava
of fresh brackish We do not think that the genus Typhlatya could types) near the coast, presence or
the have evolved later than this period. In that case we water bodies, depth and morphology of coasts
shallow of lands face new obstacles: the mentioned seas were sepa- or banks, presence emerged (in
with restricted circula- of shallow biotic factors: rated by deep ocean basins case banks), etc.; (2) pres- tion between them.The finding of a memberof the ence of competitors in the sea, presence of competi-
avail- genus at the eastern side of the Iberian Peninsula, tors in the fresh waters near the shore, food
in etc. isolated in this period from the Tethyan global cur- ability the caves,
In 7 schematic of rent, makes this option very unlikely. To overcome Fig. we give a representation
would have for the of the A this, we to postulate a pelagic or bathy- our proposal evolution genus. ma- al atyid shrimp with good dispersal abilities giving rine species would have ranged along the eastern origin to the present-day Typhlatya. As discussed, Tethys coasts about theend of the Cretaceous. This
shared ofthe the supposition of bathyal or pelagic ancestors for species represents the nearest ancestor
The full of the Atlan- any member of the family (and even more of the present-day species. opening series) is far from likely. tic Ocean and the end of the global Tethyan cur-
Additionalproblems connected with the supposi- rents divided its range in at least three populations.
American tion of a relatively recent evolution from a marine European and central populations were
would while Mid-Atlantic atyid be: (1) a more pronounced morpholog- still linked to the coasts, Ridge ical similarity between geographically close species populations remained in shallow sea banks. Like would be expected; (2) even with exceptions, as several authors (Iliffe et al., 1983; Banarescu, 1990;
Munidopsis polymorpha Koelbel, 1892, there are Stock, 1993), we believe that those shallow sea
few marine this and sank relatively anchialineanimalsrelatedwith banks may have existed from period
revision of the bathyal or pelagic taxa (Iliffe, 1992; Stock, 1993, later on. Winterer (1991) made a
seafloor is evolution of reefs and seamounts the 1994); (3) dispersal through the only pos- guyots, on sible for cold-adapted species and Typhlatya is oceanic crust of the Pacific. He concluded that
old the tropical or temperate as are most of the Atyidae. many of these seamounts were as as Barre-
Since anchialinecaves are rich in "ancient" fauna mian (Early Cretaceous). During the late Creta-
of (Iliffe, 1992; Iliffe et al., 1984; Stock, 1994), this ceous "there was an important episode develop-
of reefs and carbonate banks around and early origin should not come as a surprise. On the ment on other hand, the fauna of the Iberian Peninsula is top of the subsidiary seamounts". Furthermore, rich both in taxa that are undoubtedly Tethyan or during the Eocene, those Cretaceous seamounts have renewed vulcan- amphi-Atlantic/Mediterranean tracks, e.g., "were rejuvenated and uplifted by
Atyidae of the Paratya series, Stenasellidae, Ciro- ism" (Winterer, 1991: 264). lanidae (Isopoda), Bogidiellae, Hadziidae, Ingolf- The period of coastal existence must have been iellidae (Amphipoda), Parabathynellidae (Bathy- short for the European species, otherwise the find-
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T. miravetensis
T. pretneri
T. iliffei
T. rogersi
T. mitchelli
T. campecheae
T. pearsi
T. monae
T. consobrina
T. garciai
T. galapagensis
Summarized scenarios for the evolution of the marine Fig. 7. genus Typhlatya. Rectangles represent (coastal) populations or species.
Ellipses are fresh- or brackish-water forms. Hexagons are troglobitic species. Major geologicalevents: (1) opening of the Atlantic Ocean;
(2) division, by plate tectonics, of ancestral Central America; (3) uplifting of Bermuda; (4) uplifting of Ascension Island. The time scale
For is not proportional. The time of events is only approximative. a detailed explanation, see text.
ing of more recent Typhlatya in this area would be A representation of the ecological evolution of expected. For unknown reasons (cold weather, the species of the genus is provided in Fig. 8. Fol- competition, etc.) these populations soon disap- lowing this model, there wouldhave been two ways
survived environment: into anchia- peared. Only those species that evolved to evolve to the cave (1) earlier into a cave environment. In Central Ameri- line (or paleoanchialine) caves directly from the ca, the marine stock was again divided by plate tec- sea; (2) evolving first to brackish waters, then to tonics into three main populations: continental fresh waters and finally to caves, like T. miraveten-
Central America, proto-Antilles, and Galápagos Is- sis in our scenario (see below). It must be stressed lands (for a revision of the geological history of the that as soon as a species evolves to a cave habitat, area see Rosen, 1976). These populations quickly genetic isolation from the rest is produced. Many
after rise the recent members of the do not inhabit anchia- disappeared, giving to present Typh- genus latya of the area. In some cases, especially in the line habitats (for example the Yucatan and Europe- species from the continent, it would have been pos- an species, as can be seen in Table III). We have no
for true sible that these species first passed through a fresh- evidence to prove it, but even anchialine water phase. The populations thatremainedmarine species it is hard to think of mechanisms of disper- for a longer period of time were those linked to the sal from one cave to another because: (a) in most
Mid-Atlantic Ridge. They reached the caves in a cases the caves are separated from each other by
when islands This relatively recent period, emerged. long distances and deep sea arms (and we are dealing may explain why T. rogersi (Ascension) and T. il- with shallow and warm-temperate water adapted iffei (Bermuda) still retaineye pigmentation and are animals); (b) large quantities of offspring wouldbe found among the more haline species (see Tables I needed; (c) there are "couples" of differentspecies
& III). inhabiting geographically nearby localities (e.g.,
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Perhaps the proposed model could provide an
explanation for the absence of patterns. One reason
to explain this could be that isolation seems to have
taken place early, not only between the three main
lineages, but also early within each lineage. The
other reason could be that, after division, the inter-
mediate ancestors of each recent species could have
been affected by quite different selection pressures,
much time in the depending on how was spent seas,
which events led to troglobiosis, etc. As a result,
8. Summarized scenarios for the ecological evolution of the Fig. some species have retained plesiomorphous char-
further genus Typhlatya. For explanation, see text. acter states, whilst others became apomorphous
(fusion of ischiumand merus,reduction ofrostrum,
T. garciai and T. consobrina Botosaneanu& Holt- reduction of fifth pereiopod, loss of eye pigment, huis, 1970). Nevertheless, for very nearby caves, etc.).
after be assumed as an and dispersal speciation may For clarifying these patterns schedule, every
detailed cladis- hypothesis. particular evolutionary event and a
much characters In our model, we allow dispersalist explanations tic analysis, using as as possible, only in the evolutionary stages previous to troglobi- would be needed. It would be even more important
able to of osis. Cretaceous ancestors were disperse, to study the adaptative significance every single and they probably occupied the South Atlantic character in detail. when it opened from its older Caribbean-Central
Atlantic-Mediterranean range. But as the areas were separating further, the more the currents be- Evolutionary scenario for T. miravetensis came local, and the species evolved into brackish,
transoceanic freshwater or cavernicolous forms, The previously espoused ideas are very important
dispersal became impossible. Within each of the as background to place the evolution of this new three mainareas (see Fig. 7), dispersal at local scale species in space and time. Our model is based on the could have occurred, for example between islands paleogeographical evolution of the area, and the in the Antilles. But here we mainly mean marine, fact that the ancestors of T. miravetensis were iso- brackish, or even freshwater ancestral populations lated, from the late Cretaceous onwards. These an- in evolutionary stages where the haline behaviour cestors would have occupied the shallow waters of could have allowed low-range dispersal through the the present-day Iberian/Catalonian ranges during
to in recent times the sedi- sea, as it seems have happened the Cretaceous. Alpine orogenesis uplifted within waters led to the birth species of the genus Atya. ments of these shallow and
The shoreline of the Iberianand Catalonianranges.
its location. moved towards a position near present
Morphological patterns From here we have two options: (1) some popula-
in low- tions remained trapped swamp areas or
Some characters of the morphological recent spe- land rivers (the situation at Pla de Cabanes during cies shown of the genusare in TableI. Even if there the Paleogene) inside those ranges or between the are features of not too many listed, theabsence of any mountains and the sea; (2) the ancestors T.
be pattern can easily recognized. Geographically miravetensis followed the sea in its retreat and oc- distant often show species more similarity than spe- cupied the Mediterranean shore.
In cies from the same area. this case there are no Neogene tectonics led to the elevation of sur-
linked to habitats. The patterns rounding areas, making drainage difficult.
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karstic in the lime- AMB-0430 from the C.I.C.Y.T. The first swamp started to open a system project Spanish
author received a pre-doctoral grant from the Conselleria stone rocks underneaththe bed of limnic sediments.
d'Educació i Ciència of the Generalitat Valenciana. This induced the slow drainage of the swamp. Per- manent water bodies developed in the resulting cave system. If the swamp kept being occupied by References shrimps (first option), the population evolved to- wards a cavernicolous species into this more stable Anadón, P., 1992. Les fosses i depressions neögenes, sector habitat. Even if relatively quick (Ginés & Ginés, meridional. In: J. Guimerà (coord, ed.), Geología II. Historia of karstification would have 1992), the duration Natural dels Paisos Catalans, 2: 340—345 (Enciclopèdia been slow enough for the shrimps to evolve towards Catalana, S.A. Barcelona).
1973. On Tethys marine remnants in fresh the occupation of cavernicolous habitats. Banarescu, P.,
waters. Revue roum. Biol. Zool., 18(1): 15—21. It is also possible that after alpine orogenesis the Banarescu, P., 1990. Zoogeographyof fresh waters, 1. General did not retain shrimps and colonizationof swamps distribution and dispersal of freshwater animals: 1-511
the cave system (once it was opened in the Neogene) (Aula-Verlag GmbH, Wiesbaden).
1991. Zoogeographyoffresh 2. Distribu- took place from the Mediterranean shore, as in Banarescu, P., waters, tion and dispersal of freshwater animals in North America other Typhlatya populations in limestone caves or and Eurasia: 524-1087 (Aula-Verlag GmbH, Wiesbaden). lava tubes connectedto the sea in otherparts of the Barr, T.C., Jr. & J.R. Holsinger, 1985. Speciation in cave world. The evolution wouldbe the geological same faunas. Ann Rev. Ecol. Syst., 16: 313-337.
L. & L.B. 1970. Subterranean as in the other option. The only difference may be Botosaneanu, Holthuis, shrimps
from Cuba (Crustacea Decapoda Natantia). Trav. Inst. that subterranean drainage did not lead water to a Spéol. Emile Racovitza, 9: 21-133. low lying area but directly to the sea, giving rise to Bouvier, E.L., 1925. Recherches sur la morphologie, les varia-
an anchialine habitat of the "limestone cave" type. tions et la distribution systématique des Crevettes d'eau douce
A further marine regression (perhaps a conse- de la famille des Atyidés. Encyclopédie Entomologique, (A)
iso- 4: 1-370 (Lechevalier, Paris). quence of Neogene tectonics too) would have Bowman, T.E. & L.G. Abele, 1982. Classification ofthe recent lated the mouth from the sea. The surface water Crustacea. In: L.G. Abele (éd.), The biology of Crustacea, 1. source ensures the permanence of water and, subse- Systematics, the fossil record, and biogeography: 1-27 (Aca- of the quently, shrimps. demic Press, New York).
eastern Both options are in agreement with geological Canérot, J., 1991. Comparative study of the Iberides
(Spain) and the western Pyrenees (France) Mesozoic basins. data. There is only a minor difference: theexistence Palaeogeogr., Palaeoclimatol.,Palaeoecol., 87: 1-28. of a former connection of the subterranean system Capote, R., 1983. La tectónica de la Cordillera Ibérica. In: J.A. with the But here crucial informa- sea. geological Comba (coord, ed.), Libro Jubilar J.M. Ríos. Geología de
tion is missing. España, 2: 108-120 (Instituto Geológico y Minero de España,
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b Rostrum reaching well beyond second antennular seg- calendar for geologicals events. In: R.H. Gore & K.L. Heck ment 9 (eds.), Crustacean biogeography: 195-203 (A.A. Balkema, of all 7 6 a Ischium and merus pereiopods separate Rotterdam). b Ischium and merus in P3-4 fused 8 in Stock, J.H., 1993. Some remarkable distribution patterns telson with 7 a Rostrum not reaching beyond eyestalk, two stygobiont Amphipoda. J. nat. Hist., 27: 807-819. dorsal T. miravetensis lateral and no spines n. sp. Stock, J.H., 1994. Biogeographic synthesis of the insular b Rostrum reaching beyond eyestalk, telson with four dorsal groundwater faunas of the (sub)tropical Atlantic. Hydro- spines T. campecheae Hobbs & Hobbs, 1976 biologia, 287: 105—117. 8 a Ischiomeral articulation in P5 Stock, J.H., T.M. Iliffe & D. Williams, 1986. The concept "an- T. mitchelli Hobbs & Hobbs, 1976 chialine" reconsidered. Stygologia, 2(1/2): 90-92. P5 b Ischium and merus fused Jurassic Winterer,E.L., 1991. The Tethyan Pacific duringLate T. galapagensis Monod & Cals, 1970 and Cretaceous times. Palaeogeogr., Palaeoclimatol., Palae- 9 well ischiomeral articula- a Exopodite P5 reaching beyond oecol., 88: 253-265. tion T. consobrina Botosaneanu & Holthuis, 1970
b Exopodite P5 reduced, shorter than total length of basi-
podite 10
PI—4 T. Received: 14 September 1994 10 a Exopodites setose pearsei Creaser, 1936
Accepted: 18 May 1995 b Exopodites PI-4 not setose ... T. pretneri (Matjašič, 1956)
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