Two New Caridean Shrimps,

One Representing a New Family, from

Marine Pools on Ascension Island

(Crustacea: Decapoda: Natantia)

FENNER A. CHACE, JR.

and

RAYMOND B. MANNING

I

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 131
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 13 1

Two New Caridean Shrimps,

One Representing a New Family,

from Marine Pools on Ascension Island

(Crustacea: Decapoda: Natantia)

Fenner A. Chace, Jr.

and Raymond B. Manning

SMITHSONIAN INSTITUTION PRESS

City of Washington

1972
ABSTRACT

Chace, Fenner A., Jr., and Raymond B. Manning. Two New Caridean Shrimps,

One Representing a New Family, from Marine Pools on Ascension Island

(Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology,

number 131, 18 pages, 11 figures, 1972.—Procaris ascensionis, new genus and

new species, and Typhlatya rogersi, new species, are described from isolated

marine pools on Ascension Island. The habitats are described and the possible

origins of the shrimps discussed. Procaris, which also represents a new superfamily, may be one of the most primitive living members of the Caridea yet

discovered. A key to the caridean superfamilies is presented. The Typhlatya

(family Atyidae) is the first species of that genus to be collected outside of the

Gulf of Mexico and Caribbean Sea and the first known from salt water. A key

to the species of Typhlatya is included.

Official publication date for each title is handstamped in a limited number of initial copies and

is recorded in the Institution's annual report, Smithsonian Year.

Library of Congress Cataloging in Publication Data

Chace, Fenner Albert. 1908-

Two new caridean shrimps.

(Smithsonian contributions to zoology, no. 131)

Bibliography: p.

1. Procaris ascensionis. 2. Typhlatya rogersi. 3. Crustacea—Ascension Island (Atlantic Ocean).

I. Manning, Raymond B., 1934- joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 131

QL1.S54 no. 131 [QL444.D3] 591'.08s [595'.3843] 72-3784

For sale by the Superintendent of Documents, U. S. Government Printing Office

Washington, D.C. 20402 - Price 30 cents (paper cover)
Two New Caridean Shrimps,

One Representing a New Family,

from Marine Pools on Ascension Island

(Crustacea: Decapoda: Natantia)

Fenner A. Chace, Jr.

and Raymond B. Manning

Introduction

In June 1970, Storrs S. Olson, a graduate student

at Johns Hopkins University, visited Ascension

Island, South Atlantic Ocean, to seek evidence of

an extinct rail-like bird that had been reported

from the island. During that visit, he made several

collections of marine animals for the Department

of Invertebrate Zoology, National Museum of

Natural History. Among the specimens brought to

the Museum by Mr. Olson was a series of a small

shrimp collected from an inland salt-water pool

by Douglas S. Rogers, an employee of Pan American World Airways and Curator of the Fort Hayes

Museum of the Ascension Historical Society. The

existence of these shrimps was known to several

people on the island, and their occurrence in the

pools was reported by John E. Packer (1968) in

his handbook on Ascension Island. The shrimps

proved to belong to the caridean genus Typhlatya,

the four other known members of which live in

fresh water, usually subterranean, habitats in

Yucatan and on some Caribbean islands. Subsequently, additional collections from the salt-water

pools on Ascension were sent to us through Mr.

Fenner A. Chace, Jr., and Raymond B. Manning, Department

of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Blake Lorenz of the National Aeronautics and

Space Administration and, inadvertently, through

the District of Columbia Police Department which

recovered the samples after they were stolen from

Mr. Lorenz's automobile.

After examining these collections, we decided

that the decapod and stomatopod crustaceans of

Ascension needed more intensive study. With the

help of Helena Weiss, then Registrar of the Smithsonian Institution, a trip to Ascension was arranged

for one of us (RBM) in May 1971, and 10 days were

spent making collections there.

In correspondence exchanged during preparation

for the trip, Mr. Rogers forwarded a sketch of the

area in which were located two series of small

pools containing Typhlatya. The first sample examined was taken from what we shall refer to as

the "marl pool." Subsequent lots were taken from

that pool as well as from the seaward member of

a nearby, larger series of pools, one containing

living coral that we shall refer to as the "coral

pool." The coral pool was the habitat of both

Typhlatya and a second, larger shrimp, Procaris,

belonging to a previously unknown family. This

report is based on the shrimps found in these

pools. A more extensive report on the marine

decapods and stomatopods of Ascension Island is

in preparation.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Acknowledgments

Many persons have contributed to this study.

We thank Storrs S. Olson for making the collections

that originally kindled our interest and led to the

discovery of the two unusual shrimps in the Shelly

Beach habitat. Mr. Olson and Blake Lorenz both

contributed by delivering collections made by

Douglas Rogers. The visit by one of us to Ascension was facilitated by Miss Helena Weiss, and

permission to visit the island was granted by its

Administrator, Brigadier H. W. D. McDonald.

Valuable background information on Ascension

Island was provided prior to the trip by John E.

Packer, who was stationed there for many years.

During the visit to Ascension, dormitory and mess

facilities were made available by Major Jack Couch,

Base Commander of the U. S. Auxiliary Air Force

Base. The success of the brief visit resulted in

large measure from the interest and valuable assistance of Douglas S. Rogers, who provided transportation on the island and introduced the second

author to a variety of habitats. Ken Double of

Pan American World Airways also assisted on

several excursions. We thank J. Tuzo Wilson of

Erindale College, University of Toronto, and J. D.

Bell of the University of Oxford for information

on the geology of Ascension. Among our colleagues

at the Smithsonian, Isabel Perez Farfante, Horton

H. Hobbs, Jr., and Austin B. Williams discussed

several aspects of the study with us. We thank

Joseph Rosewater for the identification of the

mollusks, F. M. Bayer for identifying the coral,

and Thomas E. Bowman for identifying the amphipod and reviewing the manuscript.

The Habitats

The pools in which Procaris and Typhlatya

were collected are situated on a rough lava flow

extending from South Gannet Hill to Shelly Beach

near the southwestern corner of the island; the

general location, but not the exact shapes, of the

pools is shown in Figure 1. The area is known

locally as Wideawake Fairs, the breeding ground

of the Wideawake or Sooty Tern. John Packer

(in litt.) gave the following account: "The pools

are in fractured and jagged basalt, they connect

by percolation with the adjacent ocean and there

may be subterranean cavities of some size, but the

species [Typhlatya] is not truly spelean as they are

always present in numbers in the open surface

pools." These habitats have been found only in the

Shelly Beach area. Unfortunately, there is no information as to the nature of subterranean marine

passages on Ascension. Both series of pools are

situated from 55 to 90 meters inland from the

beach proper (Figure 1), from which they are

separated by a shelly ridge rising 4-6 meters above

sea level. There was no evidence that extreme

high or storm tides had recently reached as far

inland as the pools; flotsam was common on the

seaward side of the beach ridge, but was not observed on the landward side. However, the coarse

shelly material forming the beach and beach ridge

extended inland from the ridge to the seaward

pool in both series.

The marl pool is the largest and seawardmost

member of a series of three or four pools (Figure

2). It is perhaps 6 meters in major diameter, with

a maximum water depth of one-half meter and a

bottom composed of a deep, soft, marl-like material,

the flocculent surface layer of which readily goes

into suspension. It is the only one of the marl

pool series in which shrimps were observed, and

only Typhlatya was collected there. Other organisms observed and collected included numerous

sphaeromatid isopods, a gastropod, ~Nodilittorina

melearis (Quoy and Gaimard), a large burrowing

anemone, burrowing worms, copepods, and a

tanaid. On 18 May, the salinity (measured by

salinometer) in this pool was 39 °/oo

 a t

 29°C;

on 23 May, it was 35 °/oo at 25°C. On 23 May,

salinity in tide pools on the open beach was

40 °/00 at 27 °C. The landward members of the

marl pool series are completely dry at low tide,

and no organisms were observed in them. Three

subterranean openings were located in the marl

pools by Mr. Rogers, and, during periods of tidal

change, water flow can be observed at these openings.

The coral pool, in which both Procaris and

Typhlatya were found, is the seaward pool in the

larger series of about 10 pools situated approximately 125 meters southeast of and parallel with

the marl pool series (Figure 3). The narrower and

shallower members of this partially disconnected

series are lined with marl and sand, but the primary substrate of the very irregularly shaped coral

pool is rock. The deeper parts of this pool, with a
NUMBER 131

FIGURE 1.-Portion of southwestern coastal region of Ascension Island, showing locations of

(a) marl pool and (6) coral pool series inland from Shelly Beach (prepared from U.S. Navy

Topographic Map of Ascension Island, Sheet 14, 1962): Inset: outline map of Ascension Island,

showing most recent lava flows (after Atkins, Baker, Bell, and Smith, 1964) and site of enlarged

area.

water depth of about 1 meter, are protected from

direct sunlight by irregular rocky overhangs and

large boulders. It is the only one of the series in

which shrimps were collected. The most conspicuous other organisms in the pool were extensive colonies of algae (Valonia) and irregular beds

of coral (Favia); also present were the limpets

Fissurella nubecula (Linnaeus) and Diadora gibberula (Lamarck), a black gammaridean amphipod

of the genus Melita, and polychaete worms. Formalin washes of the corals and algae yielded primarily

amphipods and worms. On 18 May, the salinity

in the coral pool was 40 %o at 29°C, and, on 23

May, it was also 40 °/Oo at 27°C.

No other macroinvertebrates or fishes were observed or collected in any of these pools. On one

visit to the area, a single Grapsus grapsus (Linnaeus), the commonest shore crab on the island,

was seen on the rocks adjoining the pool, but none

was seen in or near the water.

The shrimps were active in both pools by day

and by night; at night, however, greater activity was

apparent in the more open marl pool than in the

coral pool. Both shrimps tried to evade collecting

nets, but Procaris was more agile than Typhlatya.

A sweep of the net in the coral pool might yield

dozens of Typhlatya, but the less abundant

Procaris had to be sought and collected individually. Tt took refuge under boulders and in crevices

in the irregular bottom and was difficult to catch.

The extent to which the Wideawake or Sooty

Tern preys upon these shrimps, if at all, is not
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Left, seaward portion of marl pool series, showing largest pool. Right, narrower landward extension of marl pool series. Photos by D. S. Rogers.

FICURF. 3.-Left. seaward pool of coral pool series. Right, larger pool in coral pool series showing

coral colonies near center. The tripod is in the same position in each figure. Photos by D. S.

Rogers.
NUMBER 131

known; in both pools, the shrimps would seem to

be vulnerable to these and other sea birds. The

activities of man may have some adverse overall

effects on the shrimp populations. During World

War II, the area in which the pools are located was

a practice bombing range, and there are several

bomb craters in the immediate vicinity of the pools.

At the present time, aquarium enthusiasts among

the residents of Ascension visit the area to collect

the shrimps to feed aquarium fishes. The inaccessibility of the site and its relative isolation

from the populated areas of the island, however,

help to protect the animals living there.

Origins of the Shrimps

The fragmentary fossil record of caridean

shrimps offers little evidence for the origin and

affinities of either Typhlatya or Procaris. According to Glaessner (1969), fossil Caridea, not identifiable to family, are known from the Lower

Jurassic and Cretaceous, and members of the fossil

family Udorellidae, as well as species possibly of

the Oplophoridae, also existed during those periods. Udorellids are known only from the Upper

Jurassic, but fossils identified as oplophorids are

known from both the Upper Jurassic and Upper

Cretaceous. Other caridean families are not known

until the middle Tertiary, when members of the

families Atyidae and Palaemonidae appeared.

Carideans are represented by four genera in the

Jurassic and three genera in the Tertiary. The fact

that more than 170 Recent genera are recognized

might seem to indicate that the Caridea evolved

rapidly after the Tertiary, but it is more likely

that this group, generally of small size and soft

integument, did not fossilize well and that the

fossil record does not reflect accurately the abundance and diversity of these shrimps before the

Recent.

According to several sources, Ascension is a

relatively young oceanic island of volcanic origin.

In one of the more complete accounts of the

geology of the island, Daly (1925:10) notes that

"geologically the island is extremely young." J.

Tuzo Wilson, in a semi popular article (1963),

points out that the age of Atlantic islands tends

to increase with increasing distance from the MidAtlantic Ridge. Thus, Ascension Island, on the

ridge, is estimated to be a million years old, whereas Fernando de Noronha, several hundred miles to

the west, is thought to be as much as 120 million

years old. J. D. Bell, a participant in a geological

expedition to Ascension in 1964, kindly gave us

(in litt.) an approximate age of the island of 1.0

to 1.5 million years, based on potassium-argon

dating methods, but he pointed out that the accuracy of this estimate might be questioned. In

any case, the island is probably no more than 2

million years old; in other words, it must have

appeared during the late Pliocene or early

Pleistocene.

This estimated age seems to suggest that the

stocks from which Procaris and Typhlatya descended were free-living marine forms that could

have colonized the island in relatively recent times.

It seems unlikely, however, that the present habitat

of these shrimps, in salt-water pools in the lava flow

from South Gannet Hill, is the original site of

colonization, for the sea level during the Pleistocene glacial periods almost certainly was lower than

it is at present, and the South Gannet Hill lava

flow is comparatively recent (Atkins, et al., 1964;

Bell, in litt.).

The availability of Ascension for colonization

by Typhlatya in the late Pliocene or early Pleistocene seems to suggest, as did Creaser (1936), that

Typhlatya populated its present habitats during

or shortly before the Pleistocene. If this is true,

and assuming that the directions of oceanic currents during that time were similar to those of

existing currents, then the Ascension Typhlatya

precursor possibly was an amphi-Atlantic marine

shrimp, perhaps pelagic, which existed at least

into the Pleistocene but did not survive in the

oceanic habitat into the present; living atyids are

confined almost entirely to fresh water. The close

similarities of the known species of Typhlatya

suggest a common origin, but their relationships

may be obscured by convergence.

This explanation of the origin of the Typhlatya

on Ascension may not be applicable to Procaris,

for the presumably primitive features of that

shrimp suggest that it was derived from a more

ancient stock. It is certainly possible that Procaris

or a Procaris ancestor existed as a wide-ranging

form, possibly pelagic as suggested by the long

pereopodal exopods, at least until the Pleistocene.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

This possibility is supported to some extent by the

orange coloration of P. ascensionis, which might

have persisted from the reddish pigmentation of an

ancestral pelagic shrimp; many mesopelagic

natantians are characteristically red in color. On

the other hand, reddish coloration is found in many

land-locked or subterranean shrimps that do not

have pelagic relatives, such as species of Melabetaeus, Ligur, and Barbouria (Holthuis, 1963).

An alternative explanation of the origin of the

Ascension shrimps was proposed by J. Tuzo Wilson

(in litt.): "another possibility which I think much

more likely and intriguing from your point of

view is that Ascension is only the latest in a series

of islands whose remains form scattered seamounts

and ridges from Ascension Island to the Cameroons

in one direction (The Guinea Rise) and in the

other direction to the north-east corner of Brazil.

The idea that I proposed in the Scientific American

[Wilson, 1963] was that there had been a continuously active centre from the time that Recife separated from the Cameroons and that these two

chevron-shaped ridges formed as a result of continuous volcanic action at the centre now represented by Ascension Island. If that is so, it is just

conceivable that forms of life might have survived

on Ascension from the time when the Atlantic was

very narrow and the forerunners of Ascension were

in contact with Brazil and the Cameroons." Thus,

the present populations of these shrimps on Ascension may be relicts of earlier populations that inhabited islands that have since subsided. This

explanation could well apply to either or both of

the shrimps found in the Shelly Beach habitats.

Superfamily PROCARIDOIDEA, new superfamily

(For diagnosis, see Key to Recent Superfamilies of

Caridea, p. 13.)

Family PROCARIDIDAE, new family

Third maxillipeds composed of seven segments.

None of pereopods chelate or subchelate. Epipods

on four anterior pereopods very large, forming

right angle and extending dorsad well into branchial chamber.

This family contains only the following genus.

Genus Procaris, new genus

Rostrum small, unarmed. Mandible with fused

molar and incisor processes, palp large, 3-jointed.

Dactyl of 2nd maxilliped articulating obliquely

on propodus. Maxillipeds and pereopods each

with large exopod. Anterior four pereopods with

pleurobranch, setobranch, and massive simple

epipod. Pleopods with short endopods, lacking

appendices internae or masculinae.

TYPE-SPECIES.—Procaris ascensionis, new species.

ETYMOLOGY.—From the Latin, "pro," before, and

"caris," shrimp. The gender is feminine.

Procaris ascensionis, new species

FIGURES 4-9

MATERIAL.—Coral pool back of Shelly Beach,

Ascension Island; 25 October 1970, 12:00-2:00 p.m.;

D. S. Rogers: 4 specimens.—Same; 18 May 1971,

3:00 p.m.; R. B. Manning: 2 specimens (1 is holotype, USNM 139461).-Same; 23 May 1971, 12:00-

2:00 p.m.; R. B. Manning: 14 specimens.

DESCRIPTION.—Integument thin, rather fragile.

Rostrum short, acutely triangular, unarmed, not

reaching distal margins of eyes. Carapace (Figure

4) completely unarmed; anterior margin slightly

convex, nearly vertical, anterior portion of ventral

margin forming shallow sinus below hepatic region,

trending posteroventrally to nearly horizontal ventral margin of branchiostegite; surface evenly inflated except (1) deep furrow extending from near

anterior margin below orbit posteriorly and slightly

ventrally to anterior branchial region in line with

base of 1st pereopod and (2) less prominent cervical groove extending from just above furrow on

hepatic region nearly to dorsal midline but not

quite continuous across dorsum.

Abdomen (Figure 4) with 3rd somite produced

posterodorsally as narrowly rounded cap over anterior half of 4th somite; pleura not deep, barely

concealing ventral surface of abdomen from lateral

view, those of four anterior somites rounded, of

5th subrectangular posteroventrally; 6th somite

with posterior lobes embracing base of telson

rounded, unarmed. Ventral surface of abdomen

armed with sharp spine or blunt tubercle in midline between bases of each pair of pleopods (Figure

5b); 6th somite with bulbous tubercle armed with
NUMBER 131

FIGURE A.—Procaris ascensionis. x7.

sharp spine directed posteriorly on ventral surface

between bases of uropods (Figure 5c). Telson

(Figure 5d), not including posterior spines about

1% times as long as 6th somite, armed with two

subequally spaced pairs of dorsal spines; posterior

margin (Figure be) subtriangular, armed typically

with four pairs of spines, sublateral pair very long,

two mesial pairs about one-half as long, subequal,

lateral pair shortest.

Eyestalk (Figure 5a) produced into two distal

triangular lobes, mesial lobe longer, directed

distodorsally, lateral lobe directed distally, containing irregular mass of black pigment but without discrete cornea.

Antennular peduncle (Figure 5a) short and

broad, not overreaching proximal limit of distal

one-third of antennal scale; stylocerite situated in

nearly vertical plane, with subparallel margins in

proximal two-thirds, tapering distally to acute tip

overreaching distal margin of 2nd segment of

peduncle, ventrolateral margin strengthened by

marginal rib; segments subequal in length, basal

segment with anterior margin deeply incised

dorsomesially and with proximomesial portion of

segment raised, suggesting nonexistent extra segment. Lateral flagellum (Figure 4) fully twice as

long as body of animal, irregularly setose throughout, extreme basal portion swollen and bearing

row of long setae. Mesial flagellum much shorter,

about as long as carapace and two anterior abdominal somites combined.

Antennal scale (Figure 5/) slightly less than 214

times as long as wide, distal margin rounded, produced distomesially, without distolateral tooth.

Antennal peduncle reaching about as far as distal

margin of scale, armed with acute but not sharp

tooth beneath base of scale. Flagellum slightly

longer than body of animal, irregularly setose on

mesial surface.

Mandible (Figures 6a-c) massive, with prominent

3-jointed palp, incisor and molar processes fused;

incisor process subtriangular, scooplike, with only

two obscure marginal teeth in addition to acute

apex; molar process low, indistinct, without apparent grinding surface. Paragnaths (Figure 6rf)

sinuous, distally pointed, partially enveloping

bases of mandibles. First-maxilla (Figure 6e) with

endites well formed, palp simple with long slender

terminal seta and shorter stouter subterminal one.

Second maxilla (Figure 6/) with two unequally

cleft endites, prominent slender palp, and relatively

small scaphognathite. First maxilliped (Figure 6g)

with subsemicircular endite, strong palp, base of

exopod little expanded, and epipod simple, sub-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE b.—Procaris ascensionis, holotypc: a, anterior region in dorsal view; b, ventral portions

of posterior thoracic and two anterior abdominal somites, viewed from right side, showing projections in midline between bases of pleopods and posterolateral spine on coxa of 5th pereopod;

c, ventral surfaces of posterior end of (ith abdominal somite and anterior portions of telson

and uropods; d, telson and uropods; e, end of telson; /, right antennal peduncle, ventral view;

g, posterolateral angle of right lateral mopod. Magnifications: a-d, f, xl8; e, x36; g, x91.
NUMBER 131

FIGURE 6.—Procaris ascensionis, holotype: a, right mandible, extensor surface; b, same, flexor

surface; c, same, lateral view; d, paragnaths; e, right 1st maxilla; /, right 2nd maxilla; g, right

1st maxilliped; h, right 2nd maxilliped. Magnifications: g, It, xl8; a-f, x36.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

triangular. Second maxilliped (Figure 6h) 7-

segmented, with large exopod and simple epipod,

segments of endopod not expanded, dactyl attached

diagonally to propodus. Third maxilliped (Figure

la) overreaching antennal scale by dactyl and extreme distal end of propodus, 7-segmented, with

exopod nearly as long as endopod and with rather

large simple epipod; dactyl (Figure 76) short, subtruncate, bearing one long spine and five long

strong setae.

All five pairs of pereopods similar, each with

large exopod and with long simple setae on flexor

margins, especially of carpus and propodus; dactyls

short and stout, armed with eight or nine long

curved spines in addition to numerous long setae.

Four anterior pairs each provided with pleurobranch, setobranch, and massive simple epipod

(Figure 9), in addition to exopod. First pereopod

(Figure 7c) overreaching antennal scale by dactyl,

propodus, and one-third of carpus; propodus

slightly shorter than carpus, merus twice as long

as carpus; dactyl (Figure Id) armed with eight long

spines, longest about \y4 times as long as dactyl

proper. Second pereopod (Figure 7e) distinctly

longer than 1st but overreaching antennal scale

only by dactyl and propodus; propodus considerably shorter than carpus, merus about 1.8 times as

long as carpus; dactyl (Figure If) armed with nine

long spines, longest about li/j times as long as

dactyl proper. Third pereopod (Figure 7g) subequal in length to 1st, overreaching antennal scale

by dactyl and three-fourths of propodus; propodus

sensibly shorter than carpus, merus 1.9 times as long

as carpus; dactyl (Figure 7h) armed with eight

moderately long spines, longest no longer than

dactyl proper. Fourth pereopod (Figure 8a)

noticeably shorter than 1st and 3rd, reaching little

beyond midlength of antennal scale; propodus

somewhat shorter than carpus, merus 1.8 times as

long as carpus; dactyl (Figure 86) armed with eight

spines, longest two-thirds as long as dactyl proper.

Fifth pereopod (Figure 8c) shorter than any of

preceding pairs, reaching at most as far as level of

anterior margin of carapace, without pleurobranch,

setobranch, and epipod; propodus distinctly shorter

than carpus, merus somewhat less than \y2 times

as long as carpus; dactyl (Figure Sd) especially

short and stout, armed with eight comparatively

short spines, longest barely two-thirds as long as

dactyl proper.

All pleopods (Figures Se-i) with short, poorly

developed endopods, none with appendices internae

or masculinae, but each of four anterior pairs

armed with three to five small, slender, inconspicuous spines near midlength of mesial margin;

that of 5th pair without spines, only long setae on

mesial margin. Uropod branches (Figure 5d) subequal in length, not reaching posterior margin of

telson, each branch with sharp longitudinal submedian carina, lateral branch with two unequal

spines in deep notch at juncture of lateral and

posterior margins (Figure 5g).

SIZE.—Holotype with total length about 22 mm,

carapace length 7.8 mm. Paratypes ranging in

carapace length from 4.7 to 8.6 mm.

COLOR.—Uniformly pale orange.

REMARKS.—The absence of obvious secondary

sexual characters and of ovigerous specimens in

the limited series available to us precluded the

recognition of males and/or females from external

characters. There is a remote possibility, of course,

that our specimens are juveniles, but they display

no evident postlarval characters. Certainly the absence of appendices internae on the pleopods is

not associated with immaturity necessarily, for they

are present in the postlarvae of most carideans,

and even appendices masculinae are sometimes

developed at the post larval stage.

The unusual combination of characters manifested by Procaris may well affect the accepted classification of the "macruran" decapods in ways that

we cannot fully comprehend at the present time.

This little shrimp may possibly revive the concept

of the Natantia as a natural group—a concept that

has apparently been abandoned in recent years

(Burkenroad, 1963:4; Glaessner, 1969:R444). The

7-segmented 3rd maxilliped, the subterminal rather

than subcentral attachments of the pleurobranchs,

and the lack of appendices internae on the pleopods are characters that are commonly associated

with penaeideans and stenopodideans, rather than

carideans, and the massive epipods extending well

into the branchial chamber are similar to those

found in penaeids and quite unlike any occurring

in other caridean families. There seems little doubt,

on the other hand, that Procaris is more closely

related to the carideans than it is to either of the

other natantian sections, as indicated by the anteriorly expanded and overlapping pleuron of the 2nd

abdominal somite, the form of the telson and uro-
NUMBER 131

11

FIGURE T.—Procaris ascension is, holotype: a, right 3rd maxilliped; b, same, dactyl; c, right 1st

pereopod; d, same, dactyl (denuded); e, right 2nd pereopod; /, same, dactyl (denuded); g, right

3rd pereopod; h, same, dactyl (denuded). Magnifications: a, c, e, g, xl3.5; b, d, f, h, x68.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Procaris ascensionis, holotype: a, right 4th pereopod; b, same, dactyl (denuded); e,

right 5th pereopod; d, same, dactyl (denuded); e, right 1st pleopod; /, right 2nd pleopod; g,

right 3rd pleopod; h, right 4th pleopod; /, right 5th pleopod. Magnifications: a, c, e-i, xlS.5;

b, d, x68.

pods, as well as of the mandibles and other anterior mouth parts, and, especially, by the phyllobranchiate structure of the pleurobranchs.

As a means of indicating some of the differences

and similarities between the Procarididae and the

caridean families known heretofore, we include

below a key to the Recent superfamilies recognized

by Holthuis (1955:10-12) and the Procaridoidea.

There is still much to be said for the stand taken

by Kemp (1910:36) that superfamilies should not

be established in the Caridea until the family limits

and relationships are better understood, but recognition of superfamilies does offer an opportunity

for generalized comparisons that would not be

possible otherwise. We agree completely with the

FIGURE 9.-Procaris ascensionis, paratype with carapace length

of 7.0 mm. Right side of thorax with branchiostegite removed to show epipods and points of attachment of pleurobranchs. xl4.
NUMBER 1S1 13

Key to Recent Superf amilies of Caridea

(Modified from Holthuis. 1955)

I. Third maxillipeds composed of seven segments; none of pereopods chelate or subchelate;

epipods on four anterior pereopods very large, forming right angle and extending dorsad

far into branchial chamber. (All maxillipeds and pereopods with prominent exopods;

1st pereopod not stouter than 2nd; 2nd pereopod with undivided carpus; all pleopods

with endopods poorly developed without appendices internae.) PROCARIDOIDEA

Third maxillipeds composed of four or five segments; 1st and/or 2nd pereopods chelate or

subchelate; epipods when present, small, not extending vertically into branchial

chamber 2

2.(1) First pair of pereopods subchelate; pleopods with endopods poorly or moderately developed, with or without appendices internae. (Pereopods without epipods; 1st pereopod

much stouter than 2nd; 2nd peTeopod chelate, simple, or absent, with carpus subdivided

or not.) CRANCONOIDEA

First pair of pereopods chelate or simple, not subchelate; 2nd to 5th pleopods usually with

well-developed endopods and appendices internae 3

3. (2) First and 2nd pereopods with fingers pectinate on opposable margins. (Pereopods with

exopods, without epipods; 1st and 2nd pereopods chelate, similar, 1st usually shorter and

not much stouter than 2nd.) PASIPHAEOIDEA

Fingers of chelae not pectinate on opposable margins 4

4.(3) Second pereopod with undivided carpus; 1st pereopod with well-developed chela 5

Second pereopod with carpus usually subdivided into two or more joints; if not, 1st

pereopod simple, not chelate 9

5. (4) Second maxilliped with two terminal segments articulated side by side on preceding

segment; 1st and 2nd pereopods with fingers of chelae extremely long and slender.

(Pereopods without exopods or epipods; 1st and 2nd pereopods similar, subequal.)

STYLODACTYLOIDEA

Second maxilliped with distal segments articulating serially; fingers of chelae not extremely long 6

6. (5) First pereopod with two movable crossing fingers; 2nd pereopod not chelate, terminating

in tuft of setae. (Pereopods without exopods or epipods.) PSALIDOPODOIDEA

First pereopod with single movable finger (dactyl); 2nd pereopod chelate 7

7. (6) First pereopod stouter, although often shorter, than 2nd; if not clearly so, tips of fingers

dark colored. (Pereopods with or without exopods, with or without epipods.)

BRESILIOIDEA

First pereopod not much stouter, usually more slender, than 2nd; fingers not dark

colored 8

8. (7) Pereopods usually bearing exopods; if not, chelae of first two pairs terminating in brushes

of long hairs. (Pereopods with or without epipods.) OPLOPHOROIDEA

Pereopods without exopods; chelae not terminating in dense brushes of long hairs.

(Pereopods without epipods.) PALAEMONOIDEA

9. (4) First pereopod distinctly chelate, at least on right side, usually stouter than 2nd, often

remarkably so ALPHEOIDBA

First pereopod with chela microscopically small or absent, usually no stouter than 2nd

pereopod PANDALOIDEA

qualifications expressed by Holthuis (1955:10) in

introducing his key to the caridean families and

superfamilies. Certainly full consideration must be

given to the mouth parts, gill formulae, secondary

sexual characters, and, especially, larval development in any potentially stable classification of the

Caridea, but the one proposed by Holthuis, based

largely on the form of the two anterior pairs of

pereopods, seems to be as tenable as any that have

been proposed.

Glaessner (1969:R452) has noted the futility of

attempting to integrate the few-known fossil carideans into a classification based on characters usually

discernible only in Recent animals. For that reason,

we have excluded fossil taxa from the key. Also, we

have not included the superfamily Amphionelloida
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tentatively proposed by Balss (1957:1525) and

adopted by Heegaard (1969:67) as the subtribe

Amphionidea. Even though Heegaard considered

Amphion reynaudi "as possibly the most primitive

of the known Caridea . . . ," we are not yet fully

convinced that it is a caridean. Neither do we

accept the superfamily Heterocarpodoidea and family Heterocarpodidae (more correctly "Heterocarpoidea" and "Heterocarpidae") proposed by

Thompson (1967:323); the terminal pegs on the

fingers of the chelae that Thompson believed distinguished Heterocarpus from other genera assigned to the family Pandalidae are just as

prominent, sometimes more so, in the typical

genera Pandalus and Pandalopsis.

Typhlatya rogersi, new species

FIGURES 10, 11

MATERIAL.—Marl pool, back of Shelly Beach,

Ascension Island; 17 May 1970; D. S. Rogers: 104

males, 115 females, 14 juveniles (including male

holotype, USNM 139465).-Same; 25 October 1970,

12:00-2:00 p.m.; D. S. Rogers: 135 males, 700 females, 84 juveniles.-Same; 18 May 1971, 2:20 p.m.;

R. B. Manning and D. S. Rogers: 4 males, 11 females.-Same; 23 May 1971, 9:00-10:00 a.m.; R. B.

Manning, D. S. Rogers, and K. Double: 3 males,

7 females, 2 ?.—Coral pool, near marl pool; 25

October 1970, 12:00-2:00 p.m.; D. S. Rogers: 7

males, 12 females.—Same; 18 May 1971, 3:00 p.m.;

R. B. Manning: 4 males, 14 females.—Same; 23 May

1971, 12:00 p.m.; R. B. Manning: 2 females.

DESCRIPTION.—Rostrum (Figures 10a, b) triangular, unarmed, extending beyond eyes but at most

only slightly beyond first segment of antennular

peduncle. Carapace completely unarmed, suborbital lobe forming obtuse angle projecting anteriorly distinctly beyond remainder of anterior

margin.

Abdomen (Figure 10<r) with pleura of first two

somites marginally rounded, of 3rd somite somewhat angulate, of 4th and 5th somites subrectangular posteroventrally. Sternites of first two somites

bearing triangular, flaplike projection between pleopods in both sexes. Sixth somite about 1 \/% times

as long as 5th, with lobe on either side of telson

rounded but bearing minute denticle at apex and

with sharp, posteriorly recurved pre-anal tooth at

posterior end of ventral midline between bases of

uropods (Figure lOrf). Telson (Figure 10^), not

including marginal spines, slightly longer than 6th

somite, armed in posterior half with two pairs of

small dorsolateral spines; posterior margin (Figure

10/) rounded but with shallow median sinus and

armed with four unequal pairs of spines and mesial

pair of slender setae.

Eyes (Figure 106) distally subconical, without

cornea but with irregular dark pigment spot.

Antennular peduncle (Figure lOg) robust and

short, not nearly reaching level of distal margin

of antennal scale. Stylocerite triangular distally,

extending about as far as distal margin of basal

segment of peduncle. Antennular flagella subequal,

slightly more than twice as long as carapace.

Antennal scale (Figure IO/I) slightly less than

twice as long as wide; lateral margin little convex

in proximal two-thirds, distal tooth short and

broad but distinct. Antennal peduncle nearly reaching distal one-third of blade; basal segment with

prominent tooth near base of scale. Flagellum 5i/2

times as long as carapace, extending posteriorly

beyond telson by about two-fifths length of flagellum.

Mouth parts as figured (Figures lOi-m, lla).

Mandible with five small, crowded teeth at extremity of incisor process. Scaphognathite of 2nd

maxilla broadly truncate distally; palp composed

of two segments. First maxilliped with caridean

lobe not strongly produced distomesially. Second

maxilliped with penultimate segment distally truncate, not produced along lateral margin of terminal

segment. Third maxilliped overreaching antennal

scale by about one-half length of terminal segment.

All pereopods with well-developed exopods. First

pereopod (Figure 116) reaching to about distal

one-third of antennal scale; carpus, even including

flexor extension, short and stout, distinctly shorter

than chela; ischiomeral articulation not discernible.

Second pereopod (Figure lie) reaching about as

far as 1st; carpus short, in overall length slightly

shorter than chela; ischiomeral articulation obscure. Third pereopod (Figure lid) overreaching

antennal scale by length of dactyl and one-third

of propodus; dactyl (Figure We) with four accessory spines, increasing in length distally, on flexor

margin; propodus slightly more than three times

as long as dactyl, nearly t;wice as long as carpus;

merus nearly one-third again as long as propodus,
NUMBER 131 15

FIGURE 10.—Typhlatya rogersi, holotype: a, anterior region in lateral view; b, anterior part of

carapace and eyes in dorsal view; c, abdomen; d, juncture of 6th abdominal somite and telson,

with right uropod removed to show pre-anal spine; e, telson and uropods; /, end of telson; g,

right antennule; h, right antenna; i, right mandible; ;', right 1st maxilla; k, right 2nd maxilla;

/, right 1st maxilliped; m, right 2nd maxilliped. Magnifications: c, x9 a, b, d, e, g, h, xl8; /, i-m,

x36.

armed with two movable spines laterally; ischium

shorter than dactyl, armed with one lateral spine.

Fourth pereopod (Figures 11/, g) similar to 3rd,

but shorter, reaching about to end of antennal

scale. Fifth pereopod (Figure ll/i) longer than but

reaching anteriorly about as far as 4th; dactyl

(Figures Hi, ;) elongate, armed on flexor margin

with nearly 50 denticulate spines; propodus fully

twice as long as dactyl and about twice as long as

carpus; merus about as long as propodus, armed

with one slender spine on lateral surface; ischium

less than one-third as long as merus, armed with

single slender spine near distal margin.

Endopod of 1st pleopod of male (Figures 11 k, I)

subtriangular with microscopic distal denticle,

without conspicuous marginal setae. Appendix masculina (Figure lln) on endopod of 2nd pleopod

shorter than appendix interna, armed with about
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

seven apical and subapical spines. Lateral branch

of uropod (Figure \0e) with rather prominent

movable spine inserted between distolateral angle

and margin of blade.

Gill series consisting of five pleurobranchs on

pereopodal somites, arthrobranch on 3rd maxilliped, and strap-shaped epipods on 3rd maxilliped

and all but 5th pereopod.

SIZE.—Carapace lengths of males, 2.5—4.2mm

(holotype, 3.45 mm); of females, 2.5-4.5 mm; of

juveniles, 1.6—2.5 mm. Smallest recognizable males,

with appendix masculina no more than bud, with

carapace lengths 2.5- or 2.6 mm. None of females

ovigerous.

COLOR.—White except for orange stomach contents.

REMARKS.—As indicated in the key presented below, this species differs most noticeably from the

four species previously known, from the West

Indies and Yucatan, by its shorter and stouter pereopods; this is especially apparent in the proportions of the carpus of the 1st and 2nd pairs. It

FIGURE H.— Typhlatya rogersi, holotype: a, right 3rd maxilliped; b, right 1st pereopod, c, right

2nd pereopod; d, right 3rd pereopod; e, same, dactyl; /, right 4th pereopod; g, same, dactyl;

h, right 5th pereopod; i, same, dactyl; ;, same, spines on flexor margin; k, right 1st pleopod;

/, same, endoped; m, right 2nd pleopod; n, appendices masculina and interna. Magnifications:

ad, f, h, k, m, xl8; e, g, i, I, n, x91; /, x375.
NUMBER 131 17

resembles the type-species from Yucatan, T. pearsei,

and its Cuban analogue, T. consobrina, in having

the rostrum overreaching the eyestalks, but it differs from those species most obviously in the presence of four, rather than three, pairs of spines on

the posterior margin of the telson and a pigment

spot in the eyestalks, both of which are characters

that T. rogersi shares with T. garciai from Cuba

and the latter's analogue, T. monae, from Isla

Mona and Barbuda. It differs from T. garciai, however, in having only four, rather than five or more,

accessory denticles on the flexor margin of the

dactyls of the 3rd and 4th pereopods. It may be

distinguished from T. monae by the longer rostrum and by the anterior pair of dorsolateral spines

situated in the posterior, rather than the anterior,

half of the telson, as well as by the fully developed

exopod of the 5th pereopod. In addition, all five

species of Typhlatya seem to display rather subtle

differences in the mouth parts.

In view of the possibility expressed by Botosaneanu and Holthuis (1970) that T. consobrina

may prove to be only a subspecies of T. pearsei,

we have re-examined a series of about ten paratypes of the latter species from Balam Canche Cave

and can verify that five of the six differences mentioned as means of distinguishing the Cuban from

Yucatan populations are based on errors in the

original description of T. pearsei or are probably

too variable to be useful. The telson is misrepresented in Creaser's figure; the antennal scale has a

somewhat obscure, but definite, anterolateral tooth;

the 3rd maxilliped bears only one arthrobranch;

and the 3rd pereopod is armed with three meral

spines. All of the specimens examined, however,

have the exopod on the 5th pereopod reduced,

and it seems best to treat T. pearsei and T. consobrina as specifically distinct on the basis of this

character for the time being, as in T. garciai and

T. monae.

We take pleasure in naming this species for

Douglas S. Rogers, who was responsible for bringing these two remarkable shrimps to our attention

and who followed up this initial motivating effort

Key to Species of Typhlatya

1. Rostrum overreaching eyestalks 2

Rostrum not reaching distal margin of eyestalks. (Telson armed with five pain of spines

and/or setae on posterior margin; eyestalks with inclusion of black pigment; 1st pereopod

with extensor surface of carpus much longer than palm of chela; 2nd pereopod with

extensor surface of carpus considerably longer than entire chela not including terminal

setae; 3rd pereopod with merus seven or eight times as long as wide.) 4

2. (1) Telson with posterior margin bearing four pairs of spines and mesial pair of setae; eyestalks with irregular inclusion of black pigment; 1st pereopod with extensor surface of

carpus shorter than palm of chela; 2nd pereopod with extensor surface of carpus much

shorter than entire chela not including terminal setae; 3rd pereopod with merus less

than seven times as long as wide. (Fifth pereopod with well-developed exopod.)

T. rogersi, new species

Telson with posterior margin bearing three pairs of spines; eyestalks without pigment spot;

1st pereopod with extensor surface of carpus considerably longer than palm of chela;

2nd pereopod with extensor surface of carpus about as long as, or longer than, entire

chela not including terminal setae; 3rd pereopod with merus more than nine times as

long as wide S

3. (2) Fifth pereopod with exopod extending beyond ischiomeral articulation

T. consobrina Botqsaneanu and Holthuis, 1970

[Provincias de Pinar del Rio and Camagiiey, Cuba; subterranean fresh-water "lakes."]

Fifth pereopod with exopod reduced, barely overreaching basis T. pearsei Creaser, 1936

[Estado de Yucatan, Mexico; subterranean fresh water.]

4.(1) Fifth pereopod with exopod extending far beyond ischiomeral articulation

T. garciai Chace, 1942

[Provincia de Oriente, Cuba; subterranean fresh-water "lakes".]

Fifth pereopod with exopod vestigial, barely visible T. monae Chace, 1954

[Isla Mona (Puerto Rico) and Barbuda (Leeward Islands); fresh water chiefly subterranean.]
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

by furnishing invaluable operational assistance to

our general survey of the decapod and stomatopod

faunas of Ascension Island. Perhaps we should take

this opportunity to note that it is still possible for

the amateur naturalist to make significant contributions to knowledge, especially if he or she has

the chance to observe and collect in a part of the

world that has not yet succumbed to the careless

misdeeds of civilized man.

Literature Cited

Atkins F. E., P. £. Baker, J. D. Bell, and G. W. Smith

1964. Oxford Expedition to Ascension Island, 1964. Nature, 204 (4960):722-724, figure 1.

Balss, H.

1957. Decapoda, Part VIII: Systematik. In H. G. Bronn,

Klassen und Ordnungen des Tierreichs, Funfter

Band, I Abteilung, 7 Buch, 12 Lieferung: 1505-1672.

figures 1131-1199. Leipzig: Akademische Verlagsgesellschaft.

Botos£neanu, L., and L. B. Holthuis

1970. Subterranean Shrimps from Cuba (Crustacea:

Decapoda: Natantia). Travaux de I'Institut de

Spiologie £mile Racovitxa," 9:121—133, figures 1, 2.

Burkenroad, Martin D.

1963. The Evolution of the Eucarida (Crustacea, Eumalacostraca), in Relation to the Fossil Record. Tulane

Studies in Geology, 2 (1):3—17, figure 1.

Chace, Fenner A., Jr.

1942. A New Cave Shrimp from Cuba. Proceedings of the

New England Zoological Club, 19:99-102, plate 29.

1954. Two New Subterranean Shrimps (Decapoda: Caridea) from Florida and the West Indies, with a Revised Key to the American Species. Journal of the

Washington Academy of Sciences, 44(10):318-S24,

figures 1, 2.

Creaser, E. P.

1936. Crustaceans from Yucatan. In Pearse, Creaser, and

Hall, The Cenotes of Yucatan: A Zoological and

Hydrographic Survey. Carnegie Institution of Washington Publication, 457:117-132, figures 1-43.

Daly, R. A.

1925. The Geology of Ascension Island. Proceedings of

the American Academy of Arts and Sciences, 60(1):

1-80, figures 1-8, plates 1-21.

Glaessner, M. F.

1969. Decapoda. In Moore, editor, Treatise on Invertebrate Paleontology, Part R, Arthropoda 4, volume

2:R399-R533, figures 217-340.

Heegaard, Poul

1969. Larvae of Decapod Crustacea: The Amphionidae.

Dana-Report, number 77:1-82, figures 1-165, mapi

I, II.

Holthuis, L. B.

1955. The Recent Genera of the Caridean and Stenopodidean Shrimps (Class Crustacea, Order Decapoda, Supersection Natantia) with Keys for Their

Determination. Zoologische Verhandelingen Uitgegeven door het Rijksmuseum van Natuurlijke Historie te Leiden 26:1-157, figures 1-105.

1963. On Red Coloured Shrimps (Decapoda, Caridea)

from Tropical Land-Locked Saltwater Pools. Zoologische Mededelingen Uitgegeven door het Rijksmuseum van Natuurlijke Historic te Leiden, 38 (16):

261-279, figures 1, 2.

Kemp, S.

1910. The Decapoda Natantia of the Coasts of Ireland.

Fisheries, Ireland, Scientific Investigations, 1908,

number 1:1-190, plates 1-23.

Packer, John E.

1968. The Ascension Handbook: A Concise Guide to Ascension Island, South Atlantic: 1-68. Georgetown,

Ascension Island: Privately printed by the author.

Thompson, John R.

1967. Comments on Phylogeny of Section Caridea (Decapoda: Natantia) and the Phylogenetic Importance

of the Oplophoroidea. Proceedings of Symposium

on Crustacea, Marine Biological Association of

India, part 1:314-326, figure 1.

Wilson, J. Tuzo

1963. Continental Drift. Scientific American, 208(4):86~

100.

Addendum

The description of Typhlatya galapagensis and the detailed discussion of related genera and

species by Theodore Monod and Philippe Cals (1970) in Mission zoologique beige aux ties

Galapagos et en Ecuador (N. et J. Leleup, 1964-196!), 2: 57-103, figures 1-67, came to our

attention after this paper was in press and too late to permit revision of the key to the species

of Typhlatya and of our remarks about the possible origin of the genus.

•Ct U. ». 8OVMNMCNT PRINTING OFFICKi
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