alpheus soror, a new snapping shrimp cryptospecies from sri lanka

THE RAFFLES BULLETIN OF ZOOLOGY 1999 47(2): 453-463 

© National University of Singapore 

ALPHEUS SOROR, A NEW SNAPPING 

SHRIMP CRYPTOSPECIES FROM SRI LANKA 

(CRUSTACEA: DECAPODA: ALPHEIDAE) 

a;: A. J. Bruce 

Crustacea SeC{lon, Queensland Museum, P.O.Box 3300, South Brisbane 4101, Australia 

. Email: abruce@broad.net.au 

ABSTRACT. - A new species of snapping shrimp from Sri Lanka is described and 

illustrated. Alpheus soror is a free living cryptospecies, closely related to A. gracilipes 

Coutiere, an apparently widely distributed member of the A. diadema species group. 

The status of some alpheid cryptospecies is discussed. 

KEY WORDS. - Natantia, Alpheidae, Alpheus soror, new species, Sri Lanka, 

cryptospecies. 

The recent recognition of the importance of sibling species amongst decapod crustaceans, 

particularly on coral reefs, raises some difficult taxonomic problems. Information on the 

distributions of some widely distributed tropical species is not infrequently based on historical 

records of considerable antiquity: the original specimens are often in a poorly preserved 

state or even lost, and frequently without data on live colour patterns or habitat. Close 

examination of some of these taxa has indicated that complexes of closely similar species 

are involved. Where molecular data is available the existence of closely related species may 

be indicated (sibling species). In the absence of molecular information, the relationship 

may be convergent morphological resemblance only (cryptospecies). The resemblance may 

also be due simply to very close parallel morphological evolution. The importance of sibling 

species for systematics and zoogeography are discussed by Knowlton (1993) and Knowlton 

& Weigt (1997). The number of endemic species for restricted regions can be expected to 

rise significantly. Alpheus gracilipes Stimpson, 1860, is such a species, reported to range 

from the Red Sea to the Hawai'ian Islands. Using existing identification keys (e,g., Banner 

& Banner, 1981a), the present specimen from Sri Lanka would be identified asA. gracilipes. 

Attention was drawn to the potential distinctiveness of the Sri Lankan specimen through its 

conspicuous colour pattern, fortunately recorded on film. Further examination then revealed

minor morphological differences that can be used to separate specimens for which no colour 

information is available. The species is now described as new, but it is quite probable that 

some of the specimens reported in the literature of A. gracilipes may belong to this species 

and should be re-examined. The collection of further specimens from Tahiti, the type locality 

of A. gracilipes, with photographic records of their colour patterns, would facilitate the 

clarification of the details of the species' distribution and its morphology. 

The holotype specimen is deposited in the Zoological Reference Collection (ZRC), National 

University of Singapore. The carapace length (CL) refers to the post-frontal carapace length. 

CLASS CRUSTACEA 

ORDER DECAPODA LATREILLE, 1803 

FAMILY ALPHEIDAE RAFINESQUE,1815 

Alpheus soror, new species 

(Figs. 1-3) 

Material examined. - I adult female, holotype, off Trincomalee, Sri Lanka, coIl. native collectors for 

R. Pethiyagoda, scuba, 1993, ZRC 1999.0089. 

Carapace smooth, glabrous; rostrum slender in lateral view, acute, horizontal, reaching to 

middle of intermediate segment of antennular peduncle, broadly triangular in dorsal view, 

2.0 x longer than width at level of frontal margin, centrally slightly depressed, concave, 

without median carina, distally T-shaped, ventral border concave, without setae, adrostral 

carinae sharply defined, uniformly slightly concave, reaching posteriorly well behind orbital 

hoods, orbital hoods well developed, unarmed, separated from rostrum by deep adrostral 

groove, with small feeble vertical anterior carina; frontal margin feebly produced, convex, 

unarmed, supraorbital teeth absent; pterygostomial angle rounded, cardiac notch distinct. 

Abdomen without special features, smooth, glabrous. Telson c. 0.4 of CL, sparsely setose, 

1.7 x longer than anterior width, broad, feebly concave lateral margins, posterior margin c. 

0.6 of anterior width, anterior dorsal spines small, 0.08 of tel son length, at 0.5 of tel son 

length, posterior spines similar to anterior, asymmetrical, 2 on right, 0 on left, posterior 

margin with small lateral spine with larger spine medially, subequal to dorsal spines, with 

c. 15 marginal spinules, c. 24 long plumose setae. 

Antennular peduncle short, robust; proximal segment c. as long as wide, with acute 

ventromedial tooth, stylocerite acute, exceeding anterior dorsal margin of segment; 

intermediate segment subcylindrical, 2.0 x longer than central width, subequal to length of 

proximal segment, distal segment c. 1.5 x longer than wide, 0.6 of proximal segment length, 

upper flagellum feebly biramous, with 22 proximal segments fused, shorter free ramus 

indistinctly segmented, c. 0.15 of fused rami length, lower ramus long, filiform, with c. 30 

groups of aesthetascs.

Fig. 1. A. Alpheus soror, new species, holotype female, Trincomalee. Live specimen, photo Peter 

Ng. B. Alpheus gracilipes Stimpson, Dudley Point Reef, Darwin, Australia, NTM Cr.002507, photo 

A.J.Bruce

f-I 

5.0mm 

A-H 

1.0 mm 

I 

Fig. 2. Alpheus soror. new species, holotype female, Trincomalee. A. anterior carapace and 

appendages, lateral. B. same, dorsal. C. antennular peduncle, dorsal. D. antenna. E. third maxilliped. 

F. second pleopod. G. telson. H. same, right posterior spines. 1. uropod.

Antenna with stout basicerite with strong lateral tooth, carpocerite robust, 2.6 x longer than 

central width, reaching to end of intermediate segment of antennular peduncle, not exceeding 

scaphocerite lamella, flagellum well developed, slender; scaphocerite reaching to end of 

antennular peduncle, lamella well developed, 2.0 x longer than wide, broadest proximally, 

lateral margin concave, with short stout distal tooth, slightly exceeding lamella. 

Eyes with well pigmented cornea, completely covered by orbital hoods, without anteromedial 

process. With conspicuous blunt bee oeellaire. 

Mouthparts: second and third maxillipeds only removed. Generally similar to those of A. 

bannerorum (Bruce,1987) ••..Second maxilliped with much larger, oval epipod, and third 

maxilliped with mu~ larger better developed arthrobranch than in that species; endopod 

exceeding carpocerite by half terminal segment length, terminal segment 5.0 x longer than 

proximal width, tapering distally, without terminal spine, lateral margin sparsely setose, 

medial margin densely covered by groups of short spiniform setae; carpus c. 0.5 of terminal 

segment length, sparsely setose; ischiomerus and basis fully fused, combined segment c. 1.5 

x terminal segment length, sparsely setose, exopod well developed, lateral margin slightly 

expanded with c. 18 long minutely setulose setae, numerous shorter plumose setae distally; 

coxa with large distally acute lateral plate, with hamate epipod laterally; with large 

arthrobranch. 

First pereiopods with chelae strongly asymmetrical. Major (left) chela c. 1.15 x CL, palm 

slightly compressed, smooth, glabrous, tapering slightly distally, 2.3 x longer than maximal 

width, with very feeble transverse groove on superior margin proximal to dactylus, inferior 

margin without transverse groove, dactylus c. 0.5 of palm length, slightly exceeding fixed 

finger, semicircular, compressed, distally subacute, cutting edge shallow concave, with low, 

feebly developed molar process posteriorly, ratios of dactylar length, 1 ; height, 0.45 ; molar 

process height, 0.1 ; angle, 157 0 (see Banner & Banner, 1981, p. 23); posterior molar 

process with few short setae, fixed finger distally subacute, exceeded by dactylus, deeply 

channelled with large fossa proximally; carpus very short, deeply excavate, unarmed; merus 

stout, c. 0.5 of palm length, ventrally excavate, c. 2.8 x longer than central width, distodorsal 

margin with stout acute tooth, with smaller acute medial distoventral, tooth, ventral medial 

border with 2 minutely spinulate tubercles; ischium, basis and coxa without special features. 

Minor (right) chela c. 0.9 of CL, 0.8 of major chela length; palm subcylindrical, slightly 

compressed, 3.0 x longer than deep, smooth, glabrous, dactylus c. 0.4 of palm length, elongate, 

c. 4.2 x longer than maximal depth, with bilateral balaeniceps setal ridges, tip acute, slightly 

exceeding fixed finger, fixed finger cannulate, with acute hooked tip, cutting edges sparsely 

setose, without distinct setal fringes; Proximal segments similar to major chela. 

Second pereiopods slender, exceeding carpocerite by 4 distal carpal segments; with palm 

subcylindrical, slightly compressed, c. 1.8 x longer than wide, fingers similar, c. 1.2 x palm 

length, slender, cutting edges entire, with small acute hooked tips, sparsely setose; carpus 

5 segmented, slender, c. 3.0 x chela length, segments in ratio 2: 1.5: 1: 2.8: 4.2, proximal 

segment longest; merus c. 22.0 x chela length, subcylindrical, c. 6.6 x longer than central 

width; ischium subequal to merus length, 6.4 x longer than central width; basis and coxa 

without special features. 

Third pereiopod moderately slender, exceeding carpocerite by distal half of propod and dactyl; 

dactyl slender, simple, compressed, c. 4.0 x longer than proximal depth, unguis clearly

2.0mm 

C 

1.Omm 

H 

2.0mm 

A 

5.0mm 

B-G 

Fig. 3. Alpheus soror, new species, holotype female, Trincomalee. A. first pereiopod, major chela, 

(left). B. same, dactylus. C. same, merus. D. first pereiopod, minor chela, (right). E. second pereiopod. 

F. third pereiopod. G. same, propod and dactyl. H. same distal propod and dactyl. I. third maxilliped, 

lateral coxal plate and epipod.

demarcated, c. half of corpus length; propod 4.3 x dactyl length, 0.45 of CL, slender, tapering 

slightly distally, 9.0 x longer than proximal depth, dorsal margin with numerous simple 

setae, pair of spiniform setae distally, ventral margin with 7 spines, of decreasing size 

proximally, ventrolaterally with 4 smaller spines, sparsely setose; carpus c. 0.6 of propod 

length, tapering proximally, c. 4.5 x longer than distal width, unarmed; merus 1.1 x propod 

length, uniform, 6.0 x longer than central depth, unarmed; ischium c. 0.4 of merus length, 

with very small ventral spine; basis without special features; coxa robust, with setobranch. 

Fourth and fifth pereiopods similar, with reduced spinulation. 

Pleopods normal. Second pleopod with slender appendix interna on endopod, basipodite 

with ovigerous setae proximomedially. 

lW 

Uropod with protophdite with large acute process dorsolaterally, with similar slightly smaller 

process medially, tuft of simple setae arising between teeth; exopod broad, c. 1.6 x longer 

than greatest width, lateral margin straight, with acute distal tooth, stout spine medially, 

with smaller medial tooth, dieresis well marked, posterior border without spinules, with row 

of long simple setae dorsolaterally; endopod suboval, c. 1.6 x longer than wide, posterior 

margin with c. 20 minute spinules, with short row of long simple setae medially, dorsal 

surface with central row of similar setae. 

Measurements .• Total body length (approx.), 45 mm, postrostral carapace length, 15 mm, 

major chela, 17 mms; minor chela, 13.5 mm. 

Colouration. - General body colour pale orange, antennae and flagella, pleopods and caudal 

fan similar. Cornea black, readily visible through carapace. Carapace with. small scattered 

white patches dorsally, with similar larger patches on upper branchiostegite. Abdomen similar 

dorsally, with larger white spot on lower first, third, fourth, fifth and sixth pleura, two on 

second. Second segment with large conspicuous lateral ocellus: large central spot deep red, 

almost black centrally, surrounded by broad white ring, enclosed by a narrow orange-red 

border. First pereiopods with fingers and distal palm orangish, rest of palm purple, carpus 

purple and white, merus purple banded with white. Second pereiopods purple, distal carpal 

segments orangish. Third to fifth pereiopods paler purple, dactyls orange. 

Systematic Position. - A member of the diadema species group, closely related to Alpheus 

gracilipes Stimpson, 1861. Banner & Banner (1981) have provided a recent illustrated account 

of Australian specimens of A. gracilipes. 

The new species can be distinguished from typical A. gracilipes by the following features: 

1. Longer rostrum, reaching to middle of intermediate segment of antennular peduncle. 

2. Adrostral carinae uniformly concave, more broadly divergent, (more narrowly convergent, 

biconcave in A. gracilipes). 

3. Frontal margin convex, (not subacute as in A. gracilipes). 

4 Orbital hoods less elevated, with poody developed anterior carina, (strongly elevated 

with well marked anterior carina in A. gracilipes). 

5 Distolateral tooth of scaphocerite only slightly exceeding lamella (far exceeding in A. 

gracilipes); distal lamella broad (narrow in A. gracilipes). 

6. Major chela with dactyl only very slightly exceeding fixed finger, semicircular, palm 

with only very feebly developed dorsal transverse groove (strongly exceeding fixed finger,

lower than semicircular, palm with well marked dorsal transverse sulcus, in A. gracilipes). 

Dactylus with height 0.45, depth 0.1, and angle 1570 (as opposed to 0.3, 0.95 and 1630 7. 

in A gracilipes). 

Minor chela palm and fixed finger without long setae, (palm with numerous 

in A. gracilipes). 

long setae 

8. Carpus of second pereipods with only minutely spinulate ventral tubercles, (with well 

marked spinuliferous tubercles in A. gracifipes). 

9. Third pereiopod with very small ventral ischial spine, propod without long spiniform 

distodorsal seta, dactyl less than quarter of propod length, (with well developed ischial 

spine, with long spiniform distodorsal seta and greater than one fourth of propod length 

in A. gracilipes). •• 

11. Anterior dorsal ~lson spines at 0.5 of tel son length, (distinctly less than 0.5 in A. 

gracl 'f') lpes . i. 

The easiest character with which to distinguish A. soror from A. gracifipes is the large ocellar 

spot on the side of the second abdominal segment. 

Etymology. - From Latin soror, a sister, with regard to the close relationship of this species 

with Afpheus gracilipes. 

Remarks. - Afpheus gracilipes Stimpson has been noted as one of the common alpheids 

of coral reefs (Miya, 1974) and also one of the most widely distributed, with a range extending 

from the Red Sea, East Africa and western Indian Ocean islands to Japan, Australia, east 

1.0mm 

D 

O.5mm 

E 

2.0mm 

AD 

2.0mm 

C 

Fig. 4. Alpheus gracilipes Stimpson, ovigerous female, Hibernia Reef, Coral Sea, stn. HI-31, 0.5 m, 

QM W17956. A. anterior carapace and appendages, lateral. B. same, dorsal. C. scaphocerite. D. 

third pereiopod, propod and dactyl.

to the Society Islands (type locality: Tahiti), Marquesas Islands and the Hawai'ian Islands, 

occurring largely among dead corals. The type material is no longer extant: the original 

description was brief, without note of colouration and unillustrated. A longer unillustrated 

redescription was also provided by Banner and Banner (1967). The colouration of the 

topotypic specimen, a small male, was not noted. Poupin (1998) recently noted that both 

type and neotype material have been destroyed by fire, the former in the 1871 Chicago 

conflagration and the latter in the Hawai'i Marine Laboratory fire in 1961. However, the 

latter specimen was only redescribed, as an aid "until a neotype can be described". There 

is still no neotype for this species at the present moment. The longer detailed and illustrated 

description provided by Banner & Banner (1981a), based on a specimen from Australia, is 

accepted as valid for fo~A. gracilipes Stimpson, but without the original type material 

complete certainty ~s not possible. 

I 

Heller (1865) and Ortmann (1891) also reported A. gracilipes from Tahiti, and, more recently, 

by Banner & Banner (1967). Specimens have also been reported from Moorea by Naim 

(1980) and Kropp & Birkeland (1981). Most recently the French Polynesian records have 

been reviewed by Poupin (1998). Several authors, notably Miya (1974) and Banner & Banner 

(1967, 1981b) have commented on the variability in both morphology and colour pattern 

of specimens referred to this taxon. The specimen described by Coutiere (1898) from Jibuti 

has conspicuous abdominal ocelli and "beau bleu" pereiopods and may belong to A. soror, 

as may the the ocellate material reported on by Banner (1953) from Hawai'i, suggesting a 

very extensive geographical distribution. No specimens A. gracilipes from Jibouti are 

identifiable in the collections of the Museum National d'Histoire Naturelle, Paris (A. Anker, 

pers. comm., 14 April 1999). The Banners (1967), referring to their material from the Society 

Islands, including Tahiti, report that two morphological forms exist, with and without a 

balaeniceps chela in the male. This variability suggests there may well be further concealed 

species residing under the name of Alpheus gracilipes. The specimens reported by Miya 

(1974), with dark blue upper and lower spots on the first abdominal segment, small central 

lateral spot on the second, upper spot on the third and lateral spots on fourth and fifth segments, 

suggest that at least one further species is involved. The specimen illustrated in his pI 30C 

may well belong to A. soror, suggesting again a wider distribution of this species. 

Recently it has been demonstrated that some "well known species" of alpheid shrimps actually 

consist of complexes of closely related taxonomically cryptic or sibling species. The widely 

distributed coral-associate Alpheus lottini Guerin ranges from the Red Sea to the western 

American seaboard. Knowlton & Weigt (1997) have recently demonstrated, for example, 

that the Pacific Ocean representatives of this taxon actually consist of several species. Two 

readily recognisable colour patterns were apparent but on the basis of molecular biological 

data three species of specific rank were distinguishable. Similar results have also been 

obtained from appropriate studies on other alpheids: Anemone associates Alpheus armatus 

Rathbun and three related sibling species (Knowlton & Keller, 1983, 1985); sponge associated 

Synalpheus rathbunae Coutiere and two related sibling species together with another closely 

related species, S.jilidigitus Armstrong (Duffy, 1996a); S. brooksi Coutiere and three more 

sibling species (Duffy, 1996b). Free-living Alpheus parvirostris Dana and A. bannerorum 

Bruce also appear to be a further pair of sympatric cryptospecies distinguished most readily 

by their colour patterns (Bruce, 1987). The existence of free-living Alpheus sibling species 

has been demonstrated by Knowlton et aI., (1993). These authors indicate that the species 

studied showed habitat preferences, which may have helped maintain genetic isolation. These 

preferences can exist, in free living species, on a very small scale. In Hong Hong, at low 

water spring tide level on one beach, A brevicristatus De Haan was found in a small area

of about 25 square meters of soft deeper mud, surrounded by a region of shallower sandier 

mud occupied by other Alpheus species (personal observation). Distinguishing sympatric 

and allopatric species can present subltle problems. Knowlton and Jackson (1994) have 

recently suggested that the actual species diversity on coral reefs is three to five times greater 

than currently recognised. It would therefore not be surprising to find that similar species 

complexes occurred in some other well established "species". The principle difference in 

the present case is that, unlike the first three examples mentioned above, which all have 

commensal associations, there is no evidence of any such behaviour in any species of the 

A. diadema group of the Alpheidae, which all appear to be free-living inhabitants of the 

coral reef substrate. As no biochemical information is available, the present species is not 

called a sibling species, foj.lowing the proposal of Knowlton (1986). 

Several authors havlcommented on the importance of live colour pattern as an important 

criterion in the recognition of closely related species of carideans (Knowlton, 1986; Bruce, 

1998). It may be mentioned here that alpheid shrimps recently identified as A. gracilipes 

Stimpson were of common occurrence on the shallow water reefs around Darwin, Northern 

Territory, Australia. These were of nondescript pale orangish colouration, flecked with white 

patches, without any conspicuous lateral abdominal or other eyespots. Numerous specimens 

were collected, which were very consistent in their colouration and morphology. To facilitate 

morphological comparison with A. soror some new illustrations of A. gracilipes, from 

Hibernia Reef, Western Australia, (fig. 4) are provided. 

I am most grateful to Dr Peter Ng, for providing me with the opportunity to study this specimen 

and for the use of his colour slide. The specimen was collected by Mr R.Pethiyagoda, of 

the Wildlife Heritage Trust of Sri Lanka. John Short kindly provided the specimens of 

Alpheus gracilipes from the collections of the Queensland Museum. This study was also 

facilitated by a grant from the Australian Biological Resources Survey. 

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1-50. 

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alpheus soror, a new snapping shrimp cryptospecies from sri lanka

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