12) (7) Body highly compressed; carapace with knife-like mid-dorsal keel for its entire length

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Silver plated necklace with horse shoe Horses-store.com 12) (7) Body highly compressed; carapace with knife-like mid-dorsal keel for its entire length

7. 11 8. …………………………………………………………. ………………..

Betaeus (I :347) Rostrum present, of various development; dactylus of large chela carried in superior or lateral position………………………………………………….. 9. 9 (8) Fingers of chelae of second legs about one-tenth length of palm and bearing dense tuft of scaley or setiferous bristles; sixth abdominal segment lacking articulated pleura ………………………………………………

Batella (p. 15) Fingers of chelae of second legs approximately equal to length of palm, without dense setae; sixth abdominal segment with articulated pleura.. 10 (9) Mouthparts as normal for family, mandible with molar process and palp ………………………………………………………… ………..

Alpheopsis (I :336) Mandible without molar process and palp, incisor process greatly expanded and bearing few, but strong, teeth; other mouth parts except third maxilliped modified ………………………………………

Prionalpheus (p. 12) (7) Body highly compressed; carapace with knife-like mid-dorsal keel for its entire length ……………………………………………………………Racilius (I :350) Body not markedly compressed; if carapace bearing keel, keel not knife-like posteriorly…………………………………………………………….. 12 10. 11. 12. (11) With pterygostomial margin usually produced; without anal tubercles; without mastigobranchs and setobranchs ………………… …

Synalpheus (11 :271) With pterygostomial margin rounded, never angular; usually with anal tubercles; bearing mastigobranchs and setobranchs at least on anterior thoracic legs ………………………………………………………………………. 13 13. (12) With normal, not protuding mouth parts; third maxilliped with trigonal, not flattened, basal article; appendix masculina of second pleopods of male not reaching beyond end of endopod . ……………………. .Alpheus (p. 18) THE AlPHEID SHRIMP OF AUSTRALIA 9 With mouth parts enlarged, with labrum protruding and enclosed by incisor process of mandible; third maxilliped with basal article expanded and flattened; appendix masculina of second pleopod of malft reaching beyond end of endopod . ………………………………………….

Metalpheus (p. 280) Genus Athanopsis Coutiere — Fig. 3 Batella parvimanus (‘Bate) a.

Shrimp, lateral view; h.

Antennule; c, third maxilliped; do second leg; e.

Distalend of second leg. (All figures after Bate) Batella bifurcata Miya and Miyake f.

Anterior region, dorsal view. (After Miya and Miyake). ‘I –” 18 DORA M.


BANNER Confer: Miya and Miyake, 1968b:113.

DIAGNOSIS: Rostrum acute, reaching to end offirst antennular article; acute orbital teeth nearly as long as rostrum. “Opthalmus …

Is visible in front when viewed anteriorly.” Viewed laterally, first antennular article nearly as long as second and third combined.

Stylocerite acute, reaching well beyond end of first article.

Outer f1agellum simple, not bifurcate.

Scaphocerite reaching end of antennular peduncles; squamous portion broad, lateral tooth small but acute.

Carpocerite reaching to middle of third antennular article and bearing a slender flagellum about as long as entire animal.

Third maxilliped with proximal article of endopod shown with strong subterminal spine and several spinules.

Only chela known slender, sub-cylindrical, five times as long as broad with fingers occupying the distal third (from plate).

Fingers turned slightly inward, dactylus markedly curved at tip, overhanging propodal finger when closed.

Other chela missing.

Ischium of second leg somewhat swollen in middle and tapering distally:Carpus of five articles, a little longer than merus; articles with approximate ratio of 10 :6: 6:6:9 (from figure).

Chela is almost as long as carpus and tapers markedly distally.

Dactylus and propodal finger minute, about 0.1 length of palm (from figure) and almost obscured by a distal tuft of long hairs.

Bate states: “These hairs appear to be the same diameter from base to apex, but near the base and for about half their length the surface appears to consist of scales which gradually pass into minute hairs forming a closely packed fur towards the extremity.” Third leg longer and heavier than second, with no armature described or figured, merus seven times as long as broad (from plate).

Dactylus simple.

Telson long, slender and tapering, similar to that of the genusAlpheus.

Uropods are longer than telson and outer branch with articulation. (Adapted from description and figures of Bate.) DISCUSSION: Miya and Miyake separate their species, B.

Bifurcata, on the basis of its possession of: (1) a protuberance on the carapace in the midline ofthe gastric region; (2) a bifurcation of the outer antennular flagellum; (3) biunguiculate dactyli on the third to fifth legs; and (4) a relatively longer rostrum and stylocerite.

Inasmuch as we had no specimen of B.

Parvimanus we are presenting a copy of Bate’s original drawings (Challenger Report pl.96, Fig. 2).

With the permission of Drs Miya and Miyake we are also including a dorsal view of the carapace of B.


BIOLOGICAL NOTES: Bate’s specimen was 13 mm long and it was collected at eight fathoms.


Bifurcata was longer, 19.6 mm, and was dredged at a depth of 156 m.

AUSTRALIAN DISTRIBUTION: Bate’s specimen, caught off Cape York, Qld., is the only specimen of this genus reported from Australia. Genus Alpheus Fabricius Crangon Weber, 1795:94. (Use of name suppressed; nec Crangon Fabricius 1798:387, 409).

Alpheus Fabricius, 1798:380,404. [For other synonyms see Holthuis 1955a:89] THE ALPHEID SHRIMP OF AUSTRALIA 19 TYPE SPECIES; Alpheus avarus Fabricius, 1798 [selected by Latrielle, 1810, according to Holthuis loc.


J. (See under A.

Avarus, Appendix 111.) DIAGNOSIS; Carapace continued anteriorlyto form orbital hoods which completely enclose the eyes except on ventral side.

Orbital hoods frequently projecting as rounded to acute teeth or ridges and usually de marked interorbitally from rostra I base by shallow or marked depressions.

Rostrum usually present and continued posteriorly as carina.

Pterygostomial margin of carapace rounded; with cardiac notch.

Orbitorostral process almost always present.

Antennules usually short, frequently with basal peduncular article and stylocerite reduced.

Squame of scaphocerite at times reduced; basicerite usually bearing inferolateral tooth; carpocerite usually reaching to or beyond end of scaphocerite.

Mouthparts without enlarged labrum.

Pars incisiva of mandible never expanded nor bearing long teeth; pars molaris and palp always present; subsequent mouth parts as usual for family but with distal articles of third maxilliped bearing variously bristles or spines.

Chelae of first pair always asymmetrical in form and usually in size, never carried folded against merus.

Large chela of variable form in the species, from smooth and sub-cylindrical to compressed and twisted with sculpturing of palm at times deep and strong.

Dactylus with “plunger”, a piston-like process on oppositive surface fitting into cavity on propodal pollex, usually well-developed, but at times reduced to slight confluent ridge; always with palmar and digital adhesive plaques.

Large cheliped always with short, hemispherical carpus, with merus triangular in section.

Small chela usually of simple form with conical fingers, but at times showing marked sexual dimorphism, at times with dactylus expanded and carrying dense setae on crests.

Carpus of second legs always with five articles of variable proportions.

Third and following legs robust, at times with merus triangular in section; armature and proportions of various articles variable; dactylus usually simple and conical, in some species biunguiculate, in others sub-spatulate.

Fifth legs with “brush” distally on propodus.

Abdomen usually without lateral compression, with pleura in females larger than those of ma.les, at times with acute projections on margins.

Pleura of sixth abdominal segment not articulated.

Endopod of second pleopods of males carrying an appendix masculina in addition to the usual appendix interna.

Telson with posterolateral angles not projecting or acute, posterior margin arcuate and slightly projecting, neither indented nor projecting as a tooth; two pairs of dorsal spines.

Anal tubercles almost always well-developed.

Outer uropod almost always with distal articulation; inner uropod at times bearing short heavy spines on distal margin.

Branchial formula usually with five pleurobranchs, one arthrobranch, eight epipodites and at times a supplementary arthrobranch on third maxilliped.

DISCUSSION: CHANGE FROM CRANGON TO ALPHEUS For over 100 years almost all carcinologists used ‘Fabricius’ 1798 name to designate this genus (except for those who split off certain species to which they applied what are now considered to be junior synonyms, or simply mispelled the name – see Holthuis, 1955a).

However, in 1904 M.


Rathbun, the leader in carcinology in the United States, 20 DORA M.


BANNER published upon the names used in Weber’s “Nomenclator entomologicus … ” of 1795.

She pointed out that both Weber and Fabricius had available to them unpublished manuscript copies of Daldorf’s report on Asiatic crustaceans, and each had interpreted this manuscript in a different way.

As Weber’s publicatio!l had priority over that of Fabricius, she made changes in generic names in 7 cases.

Concerning us here are only 2 changes in nomenclatu re: Weber had placed the species that Fabricius had earlier named as Astacus malabaricus in his new genus that he called Crangon.

Fabricius, in his later publication, ignored Weber’s work and created the genus Alpheus for malabaricus and other species, and Crangon for the Linnean species, Cancer crangon.

All subsequent workers, until the Rathbun report, were either unfamiliar with Weber’s work or chose to ignore it.

By the principle of priority, Rathbun showed that the nameAlpheus should be suppressed, the genus previously known as Alpheus should be called Crangon, and the commercially important genus known by the Fabrician name of Crangon must be given a new name, for which she chose Crago Lamarck.

Most Americans and some Australians followed Rathbun’s lead while most Europeans ignored her changes for the next 50 years.

It was the situation in which if the name Crangon was used in the scientific literature, one would have to know the nationality and the preferences of the author to determine if it was Crangon =Alpheus or Crangon =Crago.

Finally, in 1955 the International Commission for Zoological Nomenclature, in Opinion 334, acting upon an application by L.


Holthuis, used its plenary powers to suppress Weber’s usages and to return as officially accepted the names Alpheus Fabricius and the family name Alpheidae. (We have not listed Crangon in synonymy except when a significant contribution to the knowledge of the species was made under that generic name; all citations from Australia made under Crangon have been listed in Appendix 111.) RELATED GENERA AND SPECIES A number of genera were created for species once included in the genus Alpheus.

The first major and single most important split was the creation of the genus Synalpheus by Bate in 1888; this genus, as redefined by Coutiere (1899:334), is the second most important genus in terms of number of species within the family.

It was dealt with in Part 11 of this study, and the characteristics for the separation of the two genera were there discussed.

Other new genera which now include members previously placed in Alpheus, or contain related species, are: Pomagnathus Chace, 1937, created for a new American species, P.

Corallinus Chace; Thunor Armstrong, 1949, created for Alpheus rathbunae Schmitt, an Atlantic species and once extended to include various Indo-Pacific species which were subsequently transferred back to Alpheus – see B&B, 1962: 162 et seq.

Metalpheus Coutiere, 1908a, redefined by Chace, 1972 – see p. 280 for discussion. SUBGENERIC GROUPS Coutiere (1899) divided the genus into “Groups” and later (1905a) divided one ofthe groups into three “Sub-Groups” which he was able to define and to which he applied the name of a characteristic species of the group, as the “Edwardsii Group”.

These groups were not given the name or status of subgenera, and lie beyond the International Rules of Zoological Nomenclature.

As new species that were added to the groups varied in this or that of the group THE ALPHEID SHRIMP OF AUSTRALIA 21 characteristic, all definitions of the groups became ambiguous.

This was discussed in 1953 (B., pA7) and 1966b (B&B., p.75) – in this publication we elevated “sub-groups” to be coequal to the four original groups – and decided that while the groups could not be considered as subgenera they were of definite utility in the separation of the large number of species found within the genus.

Apparently all other contemporary workers utilize these groups, some (as Crosnier and Forest, 1966) by formally dividing the species by “Group” captions, others by referring to the “Groups” in their discussions (Holthuis, 1951, Chace, 1972).

We will also continue to use this aid to classification and will give a short characterization and notes before each group given below.

We have followed the order of placement of the groups given by de Man.

We have not been able to arrange the key so that the species within each group lie in sequence.

VARIATION IN THE GENUS ALPHEUS Within this genus, as in Synalpheus, we find some species that are remarkably constant across the span of the Indo-Pacific realm, and others that vary markedly on the same coral reef.

As with Synalpheus, we do not know how to handle this variability.


Collumianus Stimpson for example, some 25 years ago we divided the specimens from the Marianas into three separate subspecies; in this paper we are eliminating the subspecific designations; similarly, we have found sufficient variation inA.

Gracilis Helier to believe that its three subspecies (or varieties) are not warranted.

Variation is especially marked in the species in the Obesomanus Group that dwell as pairs in galleries under coralline algae and coral crust; fortunately with these the extent of variation can be determined by comparing the members of cohabiting pairs.

On the other hand, very small characteristics in the sculpturing of the palm of the large chela such as the overhanging shoulder above the superior saddle in A.

Pacificus Dana, or its sharp prolongation in A.

Chiragricus Milne Edwards seem to be consistent throughout the entire range of the species.

Some few characteristics vary with maturity, especially the secondary sexual characteristics found in the small chela of males.

Even this is not constant and recent1ly we found we could separate A.

Lobidens de Haan into two apparently geographically separated subspecies upon whether the adult males showed the development of these sexual characteristics or not (B&B, 1975:429; see also below, p. 252).

Other characteristics, especially highly modified characteristics like the hoof-shaped unguis of the third leg of A.

Lottini Guerin, may be slow in development (B, 1958:164).

We are suggesting that a young rather aberrant specimen in the present collection may be the young of A.

Novaezealandiae Miers (p. 148).

With the variation that can be observed in many of the species, we have doubts as to whether some of the forms we have described below as new species or subspecies will continue to be considered as valid when greater collections are made of these and related species.

We have taken the attitude that where in the related species a continuum of variation in a characteristic is seen and the differing form presents a slight to moderate extension of the continuum, it should not be described as new.

On the ·other hand, where the extent of variation is not known in the described or related species, or where the new form shows a condition far beyond any variation previously reported, it should be described as new simply to facilitate dealing with it, as a named form, in the future literature.

NOTES ON SOME SPECIAL CHARACTERISTICS “Balaeriiceps” chela: As far as we have been able to determine, it was Coutiere in 22 DORA M.


BANNER 1899 who first used the word “balaeniceps” to describe the development of the small chela found in certain of the species of the genus, especially those in the Brevirostris and Edwardsii Groups.

The word is obviously derived from the Latin bafaena, the name of the baleen whales, and the Latin caput, head, and refers to a fringe of setae that apparently reminded Coutiere of the rows of whalebone in a cetacean’s mouth*.

In the most extreme form, the dactylus of the small chela is broadened and somewhat excavate on the oppositive face and bears a row of regularly placed stiff setae on a low ridge starting on the lateral face near the articulation and proceeding distally; towards the tip, but short of it, the setiferous crest withdraws from the margin of the fingers and crosses over the superior surface to proceed proximally to near the dactylar articulation on the medial face (see Fig. 75, e, f, g).

The fringe demarks an area on the superior face that is approaching triangular.

Beyond the fringe of setae the tip of the dactylus usually extends as a curved, heavy and acute tooth.

This is usually a sexually dimorphic characteristic and is found in mature males, with the immature males and usually the mature females usually bearing only simple conical fingers.

Not all species, even when mature, bear this extreme condition.

Instead, the completely befringed dactylus may show lesser degrees of lateral expansion down to a narrow and tapering article (cf.

Figs. 77f and 79f).

To this condition we still apply the term “balaeniceps”.

If, however, only the medial and lateral faces of the chela bear the characteristic rows of setae, but they are not continuous over the superodistal portion, we apply the term “sub-balaeniceps” (fig. 70e).

If a setiferous crest is found on only one side of the dactylus, we have not applied either term (fig. 60f).

The term does not refer to a generalized setiferous condition, nor to the existence of tufts or bunches of setae (fig. 57f). Plunger and socket of large che/a: Characteristically in the genera Racilius, Synalpheus, Alpheus and Metalpheus, the dactylus of the large chela bears a somewhat cylindrical “plunger” that rests in a concavity in the fixed finger, or pollex, when closed.

On the superoproximal surface of the dactylus, and on the distal end of the upper palma surface, there are two “adhesive plaques”, circular areas of extremely smooth chitin.

The two plaques adhere when the chela is opened and, with the straining of the large muscles found in the palm, the adhesive bond is finally broken and the plunger is shot into the socket with such force as to make the characteristic clicking sound that gives the shrimp their common name, “snapping” or “pistol” shrimp.

This production of sound and the use of the mechanism in the shrimp’s behaviour has often been discussed in the literature (for example, Knowlton and Moulton, 1963:311, 20 figs.; Pope, 1949:326, 3 figs.; Ritzmann, 1973:459, 2 figs.

According to Dr F.


Chace Jr., judging from sketch of the chela (fig. 2a) the species discussed in Ritzman’s paper “is notAlpheus californiensis” [personal communication].) The characteristics of this plunger have seldom been used for specific separation within the genus.

Early in our study of this genus we were struck by how the apparatus varied between species.

For example, it is lacking entirely inA.

Hailstonei Coutiere, and is developed as a triangular tooth without an opposing socket in the related A.

Crockeri (Armstrong).

Of those that have the piu nger-socket apparatus (fig. 4) there are some, like A.

Pacificus Dana that have a massive plunger, with height about half of that of maximum height of the dactylus (when seen in side view) and with its margins set off abruptly from axis of the dactylus at an angle of about 115°.

In A.

Lottini the plunger is about one-third ‘Or F.


Chace Jr.

Of the Smithsonian Institution has suggested in a letter to us that Coutiere might not have been comparing the fringe of setae to the baleen, but rather the “triangular boss between the rows of setae on the extensor surface of a balaeniceps dactyl” to the “beak” of a whalebone whale when seen from above. I — (p. 72) With heavy, acute teeth flanking dactylar articulation of large chela …………………………………………………………………A.

Collumianus (p. 45) Without acute teeth flanking dactylar articulation of large chela ……… 18 ‘Underthis couplet comesA.


Novo U (ms. 1977:17; see our discussion p.24); it apparently may be distinguished from both A.

Socialis and A.

Parasocialis by the smooth rather than pustulate faces of both chelae. THE ALPHEID SHRIMP OF AUSTRALIA 18. (17) large chela rounded or at most slightly compressed in section, entire or with slight sculpturing……………. ……………………………………………

Large chela, if rounded in section, with definite grooves; otherwise, laterally compressed, with or without sculpturing……………………….. 19. (18) Dactylus of large chela either tapering to tip, or heavy and rounded, but not in shape of a double-headed hammer …………………………………

Dactylus of large chela in form of a rounded double-headed hammer, with depression on superior margin of palm to accommodate superior head of dactylus when flexed………………………….. ……………………. 20. (19) Fi rst carpal article of second legs equal in length to or longer than second First carpal article of second legs about half as long as second article, or less………………………………………………………………………………… 21. (20) Merus of large cheliped with heavy, acute subterminal tooth on inferior internal margin …………. ………………………………………………………

Merus of large cheliped with or without terminal tooth on inferior margin, never with tooth located subterminally………………………….. 22. (21) 27 19 35 20 33 21 27 22 23 Fingers of small chela about twice length of palm ……………A.

Labis (p. 127) Fingers of small chela equal to length of palm ……….A paracrinitus* (p. 129) 23. (21) Rostrum strong, acute………………………………………………………….

Rostrum absent, or very small obtuse triangle…. ………………………… — 38 DORA M.


BANNER Alpheus hailstonei Coutiere Fig. 6 Alpheus hailstonei Coutiere, 1905a:879, pI. 74, fig. 18.

Alpheus hailstonei laetabilis De Man, 1908:98; 1911 :333, fig. 64d. Alpheus hailstonei assimulans De Man, 1908:99; 1911 :331, fig. 64-64c.

Miya, 1974:116, pI. 18. Crangon hailstonei paucispinata Banner, 1953 :51, fig. 16. SPECIMENS EXAMINED: 1 specimen from AM G. 2190; 1, AMP. 7224; 3, AM P. 7377; 1, AM P. 7557; 1 AM P. 9448; 1, WM 45-65; 1 WM 63-65; 2, WM 64-65; 1, WM 69-65; 4, WM 71-65; 3, WM 90-65; 11, WM 94-65; 2, WM 132-65a; 1, WM 145-65; 14, WM 184-65; 2, WM 213-65; 2, WM 246-65; 2, WM 255-65; 5, WM 270-65; 2, WM 4985.

DIAGNOSIS: Rostrum acute, 1.5 times as long as broad at its base, reaching to middle of visible part of first antennular article.

Visible part of antennular article 0.8 as long as second article, second article 4 times as long as broad, third article 0.4 as long as second.

Stylocerite acute, reaching near end of first antennu lar article.

Scaphocerite with lateral margin concave, squamous portion very narrow, reaching short of middle of third antennular article; lateral tooth reaching to end of third article.

Carpocerite slender, 8 times as long as broad, reaching well past end of antennular peduncle.

Basicerite with strong tooth on inferolateral margin.

Large chela 2.4 times as long as broad with fingers occupying distal 0.25; dactylar articulation displaced laterally to proximal portion of chela.

Promixal superior margin of palm continued as rounded ridge leading to palmar adhesive plaque, separated from distal superior ridge by flattened area broadly rounded with slight depression in middle.

Superomedial ridge terminating distally in strong tooth at dactylar articulation, de marked proximally by transverse groove that extends a slight distance on to medial face; shoulder proximal to groove overhanging groove.

Ridge to plaque de marked on inferior side by shallow rounded groove reaching to mid-palm.

Ridge of lower margin of groove flattened and terminating in rather small but acute tooth flanking dactylar articulation.

Inferior margin of palm bearing strong, rounded shoulder proximal to broad depression at level of superior transverse groove.

Dactylus extremely compressed, curved with tip expanded especially on medial face; tip bearing patches of setae.

Cutting edge of dactylus knife-like, somewhat curved and without trace of plunger.

Cutting edge of pollex similar, but more concave and bearing promixally an angular tooth.

Merus 1.5 times as long as broad superior margin projected in a rounded tooth bearing on its margins several short setae.

Inferointernal margin bearing several small spines and an acute tooth distally.

Ischium bearing one spine distally on inferior margin.

Small chela not sexually dimorphic, 4.5 times as long as broad, fingers and palm almost equal.

Palm bearing distally an acute tooth on either side of articulation of dactylus.

Dactylus displaced somewhat laterally.

Superior and lateral surfaces rounded, glabrous.

Medial face concave its entire length and with many setae.

Tips cross when closed leaving no gape between fingers.

Merus 3 times as as long as broad, inferointernal margin bearing several small spines and an acute tooth distally.

Articles of third maxilliped with ratio 10:4.4:6.5.

Carpal articles of second leg with ratio: 10:5.6:3.1 :4.7:4.7.

Ischium of third leg with spine.

Merus inermous, 8 timeas as long as broad.

Carpus 0.6 as long as merus, superodistal margin projected ,as an obtuse tooth.

Propodus 0.7 as long as merus, bearing on its inferior margin 9 pairs of spines and a pair distally; medial THE ALPHEID SHRIMP OF AUSTRALIA 39 surface, near superior margin of all three walking legs, bearing distally 2 or 3 setiferous bristles.

Dactylus slender, 0.2 times as long as propodus, usually biunguiculate with inferior unguis small and acute; superior margin bearing tufts of setae.

Telson 2.6 times as long as posterior margin is broad.

Inner spine of posterolateral pair over twice as long as outer, anterior pair of dorsal spines placed well anterior to middle.

DISCUSSION: Since Coutiere’s original description of A.

Hailstonei from the Maldives, De Man named two new varieties and Banner, raising De Man’s names to subspecific raQk, described a third (citations above).

Banner also presented a table of 31 characteristics, some of which would serve to separate the subspecies and some to separate the species from related species.

Miya (1974:118) chose from the earlier table those characteristics he believed could be used to separate the nominal subspecies, but he cautioned that “the validity of each subspecies might be questionable … ” In his descriptions De Man emphasized the proportions of the chelae and third legs, ratio of the carpal articles of the second legs, the armature of the merus and ischium of both chelipeds and the biunguiculation of the dactylus of the third legs.

As the differences in the proportions were close and the number of intact specimens available to De Man was small, Miya chose to ignore these characteristics as reliable for use in separations; to this we agree.

This left the number of spines on the merus and ischium of the chelipeds and the biunguiculation of the dactylus of the third leg.

None of the specimens in the present or in the Hawaiian collection had spines on the superior margin of the ischium or merus of either cheliped, but on the merus of the large cheliped there would be frequently one or more setae in positions comparable to the locations where the spines were found.

All the Australian specimens carried 3-8 spines of variable size on the inferior margin of the meri of the large and small chelipeds.

The inferior margin of the ischium of the large cheliped carried one to two spines, the small cheliped carried none.

In as much as in other species such armature varies considerably, often with setae substituting for spines, and the number of spines or patches of setae also varies, we believe that this characteristic is unreliable.

Remaining then is the biungiuculation of the dactylus.



The inferior unguis, never large, varies in size.

At our request Dr J.

Forest of the Museum National d’Histoire Naturelle of Paris kindly re-examined the 4 specimens that Coutiere had marked “Type” and found all had “un minuscule ongle” (Coutiere had not shown or mentioned the secondary unguis).

Dr Forest stated that in other specimens it was not possible to distinguish the secondary unguis from a strong swelling.

In the Australian specimens the size of the secondary unguis was variable although always small, and in one 17 mm specimen the extra unguis was lacking.

In view of the variability here, and in view of the variability in other species (A.

Gracilis Helier, p.60 below and A.

Diadema Dana, Banner, 1959:142, fig. 7), we believe that this criterion cannot be relied upon for the separation of the subspecies.

Therefore we return all to the nominate species without distinctions.

It should be noted in passing that as these specimens are always dredged, each person working upon the species has reported that many, if not most, of the specimens available were in only fragmentary condition.

Possibly the lack of sufficient intact specimens has prevented earlier workers and ourselves from discovering the extent of variation within the species.

BIOLOGICAL NOTES: This species is a deep water form and has been collected from 25 metres to 536 metres.

The colour is unknown.

Our specimens ranged in size up to 30 mm. .j::.. o ~~’O — THE ALPHEIO SHRIMP OF AUSTRALIA 41 AUSTRALIAN DISTRIBUTION: In Western Australia we have specimens from Cape Naturaliste to Bluff Pt.; in eastern Australia from off Cooktown Qld.

To Cape Green N.S.W.

We also have one specimen from Victoria.

GENERAL DISTRIBUTION: This species has been reported, either as the nominate form or its subspecies, from the Maldives, Amirantei, Seychelles, Indonesia, Japan and Hawaii. (?)Alpheus staphylinus Coutiere Fig. 7 Alpheus staphylinus Coutiere, 1908a:204; 1921 :418, pI. 62, fig. 13.

Alpheus crockeri Miya, 1974, pI. 21. (Nec Armstrong, 1941.) SPECIMEN EXAMINED: 1 specimen from AM 305 (AM P. 28114).

DIAGNOSIS: Rostrum triangular, a little longer than broad at base, reaching to last quarter of visible part of first antennular article.

Anterior carapace dorsally rounded, without trace of orbitorostral grooves, orbital teeth half as long as rostrum.

Second antennular article 1.5 times as long as visible part of first and nearly twice as long as broad, third article 0.5 as long as second.

Stylocerite reaching as far forwards as rostrum.

Lateral margin of scaphocerite almost straight, lateral tooth strong, reaching to end of antennular peduncle, squamous portion reaching only to last quarter of third antennular article.

Carpocerite a little longer than antennular peduncle.

Lateral tooth of basicerite strong.

Large chela three times as long as broad with dactylus less than 0.3 as long as entire chela.

Crest leading to superolaterally placed adhesive plaque heavy and rounded, arising from proximal portion of superior margin of palm and curving laterally to plaque.

Superomedial crest separated from plaque crest by shallow flattened area and terminating in strong tooth at medial margin of dactylar articulation; crest interrupted proximally and re-arising as low, short crest, rounded in profile.

Heavy crest on lateral face proximally flattened and demarked from plaque crest by deep groove; terminating in acute tooth with rounded tip lying on lateral side of dactylar articulation; inferior side of crest de marked by deep flattened area.

Rounded concavity on inferior margin of palm somewhat proximal to fingers.

Dactylus heavy, somewhat compressed, with rounded tip closing across end of pollex; plunger developed as a conical outstanding tooth with rounded apex, socket round and complete.

Merus 2.5 times as long as broad, superior margin terminates distally in a rounded tooth, inferointernal margin bearing several small spines and terminating distally in an acute tooth.

Small chela 5.4 times as long as broad, palm and fingers equal.

Palm without sculpturing and terminated both medially and laterally proximal to dactylus in an acute tooth.

Opposing margins of fingers bearing a row of short stiff setae.

Merus similar to that of large chela.

DISCUSSION: This sole specimen is fragmentary: the body and the appendages appear as if they had been soaked in a chemical (such as a household bleach or sodium hydroxide) that ate away most of the flesh and softened the exoskeleton.

Most of the legs were already gone and the soft body was badly distorted when we first examined the specimen.

In the process of examination the small chela was lost and the dactyl was broken off the large chela.

From the parts that are intact, we can see no differences between th’is specimen and 42 DORA M.


BANNER the species Coutiere described asA.

Staphylinus from the Chagos Archipelago, the type of which we have examined at the Museum National d’ Histoire Naturelle of Paris.

The orbital hoods and the relationship of the antennular and antennal peduncles are the same; both the large and small chelae are similar, but the interruption of the ridge leading to the medial articular tooth of the large chela which we show in fig. 7c is not shown in Coutiere’s figure 12a – that is merely because of rotation for it appears in the holotype.

This specimen is also related to A.

Crockeri (Armstrong) and A.

Astrinx (described above), but can be separated from them by lack of modification of the plunger of the dactylus of the large chela; to A.

Albatrossae (Banner), but from it this species differs by having a relatively heavier and second antennular article; and to A.

Oahuensis (Banner), but that species differs by having the fingers of the small chela straight and heavy, not curved and simple.

Dr Miya reported some specimens from the Ryukyus asA.

Crockeri (Armstrong) (Ioc.


From his description and figure we suspected that the specimens might be A.


We sent specimens of A.

Staphylinus from our Philippine collections to him and he compared them to his specimens from the Ryukyus and confirmed our tentative diagnosis; he has requested that we indicate the correction of his identification in this paper (personal communication).

AUSTRALIAN DISTRIBUTION: The sole specimen was collected from Murray Island, Torres Straits, in 1907.

GENERAL DISTRIBUTION: This species is known only from the Chagos Archipelago in the central Indian Ocean; it may also occur in Japan.

We have specimens in our collections from the Philippines.

Alpheus deuteropus Hilgendorf Fig. 8 Alpheus deuteropus Hilgendorf, 1878:834, pI. 4, figs. 8-10; Coutiere, 1899:215, figs. 254, 255; Banner and Banner, 1966b:80, fig. 26; Tiwari, 1963:281, fig. 6. Crangon deuteropus Banner, 1953:70, fig. 22. Previous Australian records: O’Loughlin, 1969:36.

Houtman Abrolhos, W.A.

SPECIMENS EXAMINED: 1 specimen from AC C-54; 1, AM 467 (AM P. 27410); 3, BAU 15; 3, BAU 17; 5, BAU 30; 2, BAU 31; 7, BAU 47; 1, BAU 55.

DIAGNOSIS: Rostrum acute, curved abruptly upward toward tip, tip reaching to middle of visible portion of first antennular article; rostral carina pronounced, rounded dorsally, and continuing to base of eyes.

Orbital hoods inflated, with acute teeth about as long as rostrum.

Medial margins of orbital hoods expanded and shelflike.

Orbitorostral grooves moderately deep.

Second article of antennular peduncle slender, three times as long as broad, 1.5 times as long as visible portion of first and about twice as long as third article.

Antennular peduncles hirsute.

Lateral spine of stylocerite reaching to distal third of first antennular article.

Outer margin of scaphocerite markedly concave, lateral tooth strong, reaching to end of antennular peduncle, squamous portion narrow, reaching only to end of second antennular article.

Carpocerite as long as antennular peduncle.

Basicerite bearing strong lateral tooth and adjacent shorter acute tooth.

Large and small chelae so densely hirsute on inner faces and superior margins of outer faces that their form is obscured; hirsute surfaces heavily papillose.

Large chela THE ALPHEID SHRIMP OF AUSTRALIA 43 Fig. 8 Alpheus deuteropus Hilgendorf 25 mm male from BAU 15.


Anterior region, dorsal view; b.

Basicerite; c.

Third maxilliped; d.

Large cheliped, medial face (dense layer of hairs not fully indicated); e, f.

Large chela and detail of dactylus, lateral face; g, h.

Small chela and merus, lateral face; i.

Second leg; j.

Third leg; k.

Telson and uropods.

C, d, e, g, h, i, j scale a; a, b, f, k scale b. 44 DORA M.


BANNER strongly compressed, ovate in shape and about twice as long as high.

Fingers approximately 0.2 as long as entire chela.

Superior longitudinal crest rounded and bounded on either side by grooves; groove on lateral face extending to distal third of palm.

Superior groove reaching to linea impressa with papillose superior margin of serrulate appearance.

Superior margin interrupted by a deep transverse groove proximal to dactylar articulation.

Dactylar articulation flanked by two heavy and acute teeth; superior being a continuation of superior margin, lateral arising below lateral groove; both reaching almostto end of shortened pollex.

Inferior margin ofchela rounded, much less hirsute than superior margin and bearing small shoulder opposite transverse groove.

Tip of pollex truncate; cavity for plunger with “slot” in distal margin.

Dactylus rotated somewhat laterally, with bulbous tip overhanging pollex; plunger short.

Merus 1.5 times as long as broad, inferointernal margin projecting as slight acute tooth, superodistal margin forming a triangular projection; inferoexternal margins not projected.

Small chela 0.8 as long as large chela, ovate, fingers overO.5 as long as palm.

Dactylar articulation flanked by acute teeth, with medial prominent and lateral tooth small.

Superior margin of palm bearing a transverse groove near dactylar articulation with proximal margin overhanging groove.

Merus similar to that of large chela.

Third maxilliped densely hirsute.

Ratio of articles: 10:4:8.

Last article 6.6 times as long as broad, tapering distally.

Ratio of carpal articles of second legs: 10:6:2:2:4.

Ischium of third leg with spine.

Merus 3.5 times as long as broad, with strong acute tooth distally and numerous hairs on superior and inferior margins.

Carpus 0.5 as long as merus, both margins terminating in strong acute teeth.

Propodus as long as carpus, bearing on inferior margin four pairs of small spines and a pair distally.

Dactylus simple, curved.

Telson stout, anterior margin 2.2 times as wide as posterior margin.

Lateral margins anteriorly convex, posteriorly concave.

Anterior pair of dorsal spines at 0.5 of length.

Distolateral margins of inner uropods bearing five strong spines.

DISCUSSION: Our specimens showed the same variation in characters as those discussed by Banner (1953:72) for 50 specimens from Hawaii. 1.

The rostrum varied from slightly shorter to slightly longer than the orbital teeth. 2.

The frontal margin of the carapace between the orbital teeth varied from almost straight to definitely arcuate. 3.

The length of the stylocerite and relative lengths of scaphocerite, carpocerite and antennular peduncle varied.

Usually the scaphocerite reached to somewhat past the middle of the third antennular article. 4.

The second carpal article of the second legs varied from 0.55 to 0.85 times the length of the first article, usually being about 0.6-0.7 its length.

The distal articles appeared to bear a more fixed relationship to the length of the second article than to the length of the first article. 5.

The number of spines on the propodus of the third legs varied from four to seven pairs.

BIOLOGICAL NOTES: This species characteristically is found in fissures in massive heads of living coral.

The following forms have been reported as hosts: Astreopora myriophthalma (Lamarck), Porites lobata Dana (B&B, 1964:88); Porites spp., Acropora sp. (B&B, 1966b:82); Porites evermanni Verrill, Montipora verrucosa (Lamarck), and Pavona varians Verrill (Vaughan, 1973:37 et seq.).

It may continue to live in fissures persisting in heads after the coral has died and it has been found in dead bases of living coral heads.

The species has been found from the low intertidal to depths of 35 m (Vaughan, op.

Cit.) but this was the limit of depth of his observations. THE AlPHEID SHRIMP OF AUSTRALIA 45 Vaughan (op.

Cit.) made an extensive study of A.

Deuteropus, its fissures, distribution and behaviour in Hawaii.

He found that the fissures inhabited were from 0.3-0.9 cm wide and up to 25 cm long and that the longerfissures had the greater depths (circa 3 cm).

Off the deeper parts of the U-shaped crevice were the burrows, rounded cul-de-sacs reaching deeper into the head and exactly fitting the shrimp that would retire there when disturbed, lying with its hairy chela effectively blocking the entrance.

Vaughan found that 85 percent of the fissures he studied contained a cohabiting pair, each with its separate burrow; a few crevices less than 5 cm long had only one inhabitant and one burrow; in two cases he found in longer fissures two berried females with a male and in one other longer fissure he found two pairs living.

While Vaughan could not prove that the shrimp actually excavated the crevice-burrow unit, he could show that the fissure was not pre-existing in the growth form of the coral, that the groove was not made by any other known biological agent, and that it might be possible for the shrimp to inhibit the growth of coral and possibly excavate the coral with its large chela.

He made sections through the coral head at the site of the groove and was able to count up to five annual growth rings to the sides of the groove; whether that meant that the shrimp had been in position for five years or whether it had actively burrowed through five years growth his studies did not show.

He found the rim of the groove to bear a band of small hydroids of various families embedded in a mass of filamentous alga dominated by the red alga, Spermothamnion that extended 2 cm from the lip.

He speculated that the hydroids might with their nematocysts protect the inhabitants of the groove and might also eat the shrimp larvae when released.

He was able to show that when the shrimp were removed from their dwellings and given fragments of the algal mat under laboratory conditions they would graze on the mat.

He found the same algae in the gut contents of the shrimp freshly removed from their burrows, so the shrimp use the mat as a standing food crop. (All information from Vaughan summarised with his permission.) It is noteworthy that when we were collecting in Indonesian waters in 1975 we found many massive heads of coral with grooves that appeared to be superficially similar to those made by A.

Deuteropus, but in almost all cases they were inhabited by A.

Acutofemoratus Dana (full collections not yet studied and will be reported in a future paper).

We have no colour observations on the Australian specimens, but in Hawaii the species has pink eyes, surrounded by a circlet of broken red lines and the body and appendages are largely transparent with a sprinkling of small red chromatophores; the gut shows th rough the carapace as brown, and the inner face and the tips of the fingers of the chelae are brownish.

The embryos in maturing eggs are brown with black eyes.

The specimens ranged from 8 to 30 mm in length.

Australian distribution: One specimen was found in the Houtman Abrolhos in Western Australia, the remainder were found from off Port Douglas, Qld.

To as far south as Heron Island in the Capricorn Group.

General distribution: South and East Africa, Maldive Archipelago; Thailand, Vietnam; Japan, Marianas; Marshall Islands; Samoa; Line Islands and Hawaii. Alpheus collumianus Stimpson Fig. 9 Alpheus collumianus Stimpson, 1861 :30; Banner, 1953:67, fig. 21.

Alpheus collumianus probabilis Banner, 1956:338, fig. 10. 46 DORA M.




BANNER Fig. 10 Alpheus villosus (Olivier) 40 mm male from WM 296-65.

A, b.

Anterior region, dorsal and lateral view; c.

Third maxilliped; d, e.

Large cheliped, medial and lateral face; f, g.

Small cheliped, medial and lateral face; h.

Second leg; i.

Third leg; j.

Telson and uropods. 38 mm male from WM 247-65.

K, I.

Third leg and enlarged dactylus.

A, b, c, h, i, j, k scale a; d, e, f, g scale b; 1, scale c. THE ALPHEID SHRIMP OF AUSTRALIA Paralpheus diversimanus Bate, 1888:568, pI. 102.

Previous Australian records: Haswell, 1882b:187.

Southern Australia.

Miers, 1884:290.



Ortmann, 1894:14.

Thursday Is., Torres Straits.

Coutiere, 1898f:206. (discussion of type) Nobili, 1899:233.

Beagle Bay, Dampier Land.

Sayce, 1902:155.

Port Phillip, Victoria.

Balss, 1921 :9.

Cape Jaubert, W.A.

Hale, 1927a:46, fig. 37.


Australia (as Crangon vil/osus).

Hale, 1927b:307.

Baeres Pt., Kangaroo Is., S.

Australia [as C.


Hale, 1941 :265.


Australia [as C.

Vil/osus]. 51 SPECIMENS EXAMINED: 1 specimen from AC 40; 5, AM 15 (AM P. 27798); 1, AM 18 (AM P. 27785); 3,AM21 (AM P. 27793); 2, AM 34 (AM P. 27822); 1, AM 122 (AM P. 28145); 1, AM 128 (AM P. 27823); 3,AM 139 (AM P. 27820); 2,AM 224 (AM P. 27819); 1,AM248 (AM P. 27829); 2, AM 303 (AM P. 28146); 2, AM 308 (AM P. 27784); 4, AM 382 (AM P. 27834); 1, AM E. 4496; 4, AM E. 4497; 1, AM G. 614; 1, AM P. 2055; 1, AM P. 2344; 2, AM P. 2768; 6, AM P. 3014; 8, AM P. 3956; 1, AM P. 8702; 1, AM P. 9422; 1, AM P. 13583; 4, AM P. 14959; 5, AM P. 27884; 12, AM P. 27885; 1, BAU 28; 2, SM C-501 ; 2, 5M C-502; 1, SM C-504; 4, SM C-505; 1, 5M C-514; 1, SM C-1074; 1, TM G1461; 1, TM G1511; 1, TM G1528; 1, VM 1; 4, VM 10; 13, VM 11; 2, VM 17; 1, VM 18; 2, VM 22; 1, VM 30; 1, VM 33; 1, VM 25S; 2, VM 335; 2, VM 41N; 1, WM 38-65; 3, WM 85-65; 1, WM 105-65; 1, WM 187-65; 2, WM 212-65; 2, WM 226-65; 2, WM 247-65; 1, WM 274-65; 1, WM 293-65; 6, WM 296-65; 1, WM 301-65; 1, WM 298-65; 1, WM 6052; 1, WM 8752; 1, WM 9982; 1, WM 10468; 1, WM 10570; 1, WM 108-60; 1, WM 11100/1; 1, WM 115-33.

DIAGNOSIS: Entire body hispid with short stiff setae interspersed with scattered long setae, all arising from small bosses.

Rostrum acute, 1.8 times as long as broad at base, slanted upward and reaching end of first antennular article; carina sharp, carrying acute, forward-directed tooth at level of posterior margin of eyes, and continued posteriorly as sharp ridge to mid-carapace.

Orbital hoods inflated, each with acute tooth overhanging anterior margin of hood.

One small triangular tooth on orbitorostral margin between orbital hoods and base of rostrum.

Eyes never brown to black, at times appearing chalky or ranging from a light yellow to a dark red in alcohol.

Second antennular article about 2 times as long as broad and 1.3 times length of first and almost twice length of third article; all articles with scattered long setae.

Stylocerite with prominent lateral tooth reaching beyond end of first antennular article.

Scaphocerite with lateral margin concave, lateral tooth reaching beyond end of antennular peduncle, almost to end of carpocerite; squamous portion narrow, reaching middle of third antennular article.

Carpocerite reaching more than half length of third antennular article past that article.

Basicerite with lateral tooth heavy, cylindrical and reaching to end of second antennular article; inferior to major tooth lies a minute but acute tooth (not seen from angle of fig. 10b).

Third maxilliped stout, third article 2.2 times as long as broad.

Medial surface of all articles bearing long hairs with those of second and third articles long, thick and brushlike.

Large chela compressed, 2.2 times as long as broad with fingers occupying the distal 0.2.

All surfaces of chela bearing small tubercles, lateral face with short bristles, but medial face with thickly set stiff setae that sweep forward.

Medial face without grooves. 52 DORA M.


BANNER Superior face with obliquely directed longitudinal groove extending from carpal to dactylar articulation.

Lateral margin of groove smooth, medial serrate.

Lateral face with smooth broad groove arising mid palm and extending to dactylar articulation.

Near inferolateral margin is a moderately deep longitudinal groove that extends from inferior shoulder nearly to distal portion of pollex.

Dactylar articulation flanked by acute teeth on either margin.

Tip of dactylus truncate, overhanging end of pollex.

Inferior margin of ischium with three spines.

Merus 2 times as long as broad, surface covered with bosses carrying hairs.

Inferointernal margin bearing two or three spines, distally bearing a small acute tooth.

Superior margin projected into a broad rounded tooth and bearing a tuft of setae.

Small chela not sexually dimorphic, 2.7 times as long as broad, fingers and palm nearly equal.

Dactylar articulation flanked by teeth, with medial tooth longer and curved upward.

Superior margin bearing groove di rected obliquely toward lateral face similar to that of large chela.

Merus similar to that of large chela.

Carpal articles of second leg with ratio: 10:5:2:2:5.

Chela as long as last three articles.

Ischium of third leg armed with spine.

Merus 2.7 times as long as broad, inferior margin bearing about seven patches of short stiff setae, or alternatively with definite spines, and terminating distally in a strong acute tooth.

Superior margin terminates in a patch of moderately long stiff setae.

Carpus 0.6 as long as merus, inferior margin bears two to three patches of setae or spines simi lar to those on merus and terminating in acute tooth; superior margin projected but rounded.

Propodus a little longer than carpus, with inferior margin beari ng six pai rs of spines and a pai r distally.

Superior margi n beset with a series of patches of long stiff setae with a spine distally.

Dactylus biunguiculate, inferior unguis two-thirds as long as superior.

Telson 1.7 ti mes as long as broad, anterior margin 1.3 ti mes as wide as posterior.

Anterior pair of dorsal spines placed anterior to middle.

Lateral margin of inner uropod bearing a row of spines.

DISCUSSION: Olivier in 1811 described among other species collected by Perron (or Peron) two species that are now believed to be identical.

One was named Pa/aemon villosus from the “mer des Indies” but bearing the label in the vial of “Port du Roi Georges”, presumably King George Sound at Albany, W.A., and the other was named P.

Diversimanus from “Nouvelle Hollande”.

The specimens of “P.

Diversimanus” evidently have been lost, and the exact locality of collection cannot be ascertained as Peron accompanied Nicolas Baudon, who in 1800-1804 explored and mapped the south and west coasts of Australia from near Sydney around to Melville Island.

In his terse descriptions, Olivier did not contrast the characteristics or otherwise distinguish between the two named forms.

In 1837 Milne-Edwards re-described A.

Villosus and stated (p. 354) that A.

Diversimanus “ne me parait pas devoir etre distingue specifiquement de l’Alphee velu … ” But he did state that A.

Diversimanus had a little more abundant hai rs which were a little stiffer.

Others accepted this synonymy until Bate (1888) revived the specific name diversimanus and created it for a new genus, Paralpheus; his specimen came from off Cape York.

In 1898f Coutiere re-examined the specimens of Bate and decided that his Paralpheus diversimanus was actually Alpheus villosus.

In his 1899 Thesis Coutiere provided some excellent figures of the species (see above).

Since that date the name has stood except for that interim when the name Crangon was substituted for Alpheus.

This species shows variation on a number of characteristics.

In preservation and THE AlPHEID SHRIMP OF AUSTRALIA 53 possibly in life there is a variation noted above in the eye colour.

The rostrum may reach considerably short of the end of the first antennular article to somewhat beyond; the carinal tooth at the level of the posterior margin of the eyes may vary from a slight protuberance to a large acute tooth.

The lengths relative to the antennular articles of the carpocerite itself and the large tooth of the basicerite both vary.

The ratio of lengths of the fi rst two carpal articles of the second legs ranges from 10:5 to 10:8.

In Bate’s figure (pI. 102, fig.

V) he showed two spines on the di stal portion of the outer u ropod; two of ou r speci men s were si mi larly armed, but all other specimens had but a single tooth.

Other minor differences in proportions were seen.

However, the armature of the merus and carpus of the third legs can be used to separate the species into two distinct forms.

One form, mostly from south temperate waters, carries tufts of setae on the inferior borders of these articles; the other, always from tropical waters, carries distinct spines in place of the setae (contrast figs. 10i and 10k).

Between the two forms we have found no intermediates, such as thicker setae or finer spines approaching setae.

This difference has not been noted by earlier workers.

Coutiere, in his figure of the third leg (1899; fig. 319) showed spines on the type specimens, but our examination of the same specimen (Olivier’s, from Port du Roi Georges) showed these were not spines but setae.

Bate’s specimen of “diversimanus” (pI. 102, fig. 1) was shown to have spines as would be expected from its tropical location, but Coutiere (1898f) in his re-examination of the specimen did not remark on the difference.

We were initially inclined to consider the form bearing the spinose articles as a distinct and geographically separated sub-species.

The northern form did not extend beyond the Torres Straits (about 11 °S) in the east nor beyond Exmouth Gulf (about 22°5) in the west, while the southern form did not reach further north than 37°5 in the east nor 32°5 in the west save for one specimen from Houtman Abrolhos (at about 28°43’5).

However, after we had made the decision two additional specimens were received that upset this geographical separation: the specimen from SM C505, from “Northern Territory” noted above, and one specimen from the Sulu Archipelago, Philippines (WM 216-65), neither of which have the spines on the third legs characteristic of the other tropical specimens.

We therefore are deferring the separation of the species into geographical or ecological sub-species until further studies are made.

Two other minor mistakes have appeared in the literature.

Bate described the dactylus of the third leg as being “single pointed”; this was corrected by Coutiere in his re-description.

Coutiere (1899, fig. 383) did not show the row of small spines on the margins of the inner uropod, but our re-examination of the holotype showed these were present and they were found on all other specimens we have examined.

BIOLOGICAL NOTES: This species has been reported to reach 67 mm in length.

It has been dredged as deep as 24 fathoms and taken from corals collected intertidally.

The specimens for which the habitat was noted in the collection data, were recorded from cryptic locations; the sole specimen we ourselves collected was from deep within a head of coral.

It is probable that the lack of normal pigmentation of the eyes is associated with this dimly lit or lightless habitat.

The colour notes on the northern and southern forms show similar colouration.

The specimen we collected from the Torres Straits region was lemon yellow with stellate red chromatophores, with red and white striped antennules and antennae and red eyes.

The colour notes on a specimen supplied by the Fisheries and Wildlife Department of Victoria (VM 335) from Western Port stated “All over pale light orange, spotted with red stellate 54 — ‘—_ _—l’–_ _–I. C Fig. 15 Alpheus lottini Guerin 29 mm male from AMP 8026.

A, b.

Anterior region, dorsal and lateral view; c.

Third maxilliped; d, e.

Large cheliped and enlarged distal region; f.

Small cheliped, lateral face; g.

Second leg; h, i.

Third leg and enlarged dactylus, inferior face; j.


A, b, c, e, g, h, j scale a; d, f scale b; i scale c. THE ALPHEID SHRIMP OF AUSTRALIA 67 8567; 3, AM P. 10322; 2, AM P. 11401; 1, AM P. 13547; 1, AM P. 13573; 1, BAU 3; 1, BAU 4; 1, BAU 15; 1, BAU 16; 1, BAU 24; 1, BAU 27; 1, BAU 32; 2, BAU 38; 4, BAU 44; 2, BAU 47; 3, BAU 48; 1, BAU 52; 1, BAU 57; 1, )C23; 2, MM 111; 1, MM 263; 1, MM421; 2, QM W. 999; 2, VM 2; 1, WM 23-65; 1, WM 118-65; 2, 75 LlZ-S (AM P. 27918).

DIAGNOSIS: Rostrum slender, acute, reaching to end of first article of antennular peduncle.

Rostral base broadened, flattened, not carinate dorsaliy, separated from carapace by deep and narrow sulci on each side.

Orbital hoods rounded laterally, bearing acute teeth on more medial portion of hood, teeth directed slightly inward.

Antennular peduncle slender, with second article variable but usually 1.7 times as long as broad, only slightly longer than visible portion of first or third.

Stylocerite reaching to middle of second antennular article.

Scaphocerite long with narrow squamous portion reaching just beyond end of antennular peduncle and with lateral tooth reaching well past end.

Carpocerite subequal to length of scaphocerite.

Lateral spine of basicerite acute, equal to or exceeding length of stylocerite.

Third maxilliped stout, articles with a ratio: 10:3:8.

Inferior margin of both first and second article bearing spines, superior margins bearing coarse setae distally.

Large chela compressed, margins rounded, without grooves or crests, 2.5 times as long as broad.

Dactylus not strongly arcuate, in males 0.3 length of entire chela with tip acute and hooked; in females dactylus rounded at tip, only 0.2 length of entire chela.

Merus with both superior and inferointernal angles rounded, although projecting; inferointernal margin with about 5 spines.

Small chela almost as long as large chela but more slender, varying from 2.5 to 3.0 times as long as broad; fingers about equal in length to palm.

Dactylus strongly curved at tip, crossing fixed finger when closed.

Inner margin of cutting face of fixed finger with distinct lamellar ridge along entire length.

Merus similar to that of large cheliped.

Ratio of carpal articles of second leg: 10:6:6:6:5; last four articles almost as broad as long.

Ischium of third legs inermous.

Merus inermous, over 3 times as long as wide.

Carpus with both margins continued as heavy projections.

Propodus with 5-7 spines, with broadened, flattened tips.

Dactylus heavy, blunt, laterally compressed, with thick longitudinal ridge on inferior face continuing around blunt tip as a curving ridge, the blunt tip thus carrying a ridge of hard chitin similar in form to a horses hoof; portion of dactylus surrounded by ridge of soft flexible chitin.

Telson 3.3 times as long as broad at posterior end.

First pair of dorsal spines placed just anterior to middle.

Inner pair of posterior spines nearly same size as dorsal spines.

DISCUSSION: This well-known species has been figured and described many times and has been known by4 names.

Coutiere in 1899 (p. 12) placedA/aevis into synonymy toA ventrosus; Holthuis in 1958 (p. 12) reviewed the synonymy and accepted Kingsley’s (1882:113) decision thatA ventrosus was a junior synonym of A.

Lottini Guerin.

Holthuis (1957,1961) also discussed the dates of publication of Guerin’s plate bearing the name Lottini (in his 1838 description he spelled it Lottinii).

The junior author also created a synonym, Crangon latipes, based on a juvenile specimen markedly different from the adult (1953:82); this was corrected in a subsequent paper (1958:165).

Finally we have reported that the largely unused name A thetis White is a synonym of A.

Lottini (B&B, 1977:282).

This species is most easily recognized and the most surely identified in its adult form in the whole genus by the unique development of the dactyls of the walking legs; smaller 68 DORA M.


BANNER specimens found in symbiosis with living pocilloporid corals which do not have this development should be carefully compared to the description of the juvenile form (loc.

Cit.) and with the other symbiont of the same corals, Synalpheus charon (Helier) (B&B, 1975:369).

BIOLOGICAL NOTES: This species is an obligate symbiont, living exclusively in living heads of the members of the Pocilloporidae, especially Pocillopora meandrina var.

Nobilis Verrill, P.

Ligulata Dana, large colonies of P.

Damicornis (Linnaeus) (=P.

Cespitosa Dana) and some of the species of the genus Seriatopora (the last reported by Patton, 1966).

This species, together with crabs of the genus Trapezia, is found in various parts of the branching colonies, but retire deeply between the branches when disturbed.

Also in the same heads, but apparently confined to the unbranched base is the smaller Synalpheus charon (Helier).

All three symbionts are of similar colour, a mottled bright orange-red ground color with a dark red mottling.


Lottini also may bear a mid-longitudinal stripe of deep red that may appear almost black.

In most of the Australian collections the exact depth at which the specimens were collected was not often clearly indicated, but as the host corals cannot survive much intertidal exposure, most must have been subtidal, and extending down to the depth limits of the host.

The specimens range in size up to 38 mm.

AUSTRALIAN DISTRIBUTION: We have a few specimens from western Australia from near Perth to the Dampier Archipelago.

In North Australia we have specimens from Darwin and the Torres Straits, but the majority of specimens range from Princess Charlotte Bay in north Queensland to Sydney.

We also have 4 specimens from Lord Howe Island.

There were no specimens from South Australia or Tasmania in our collections.

GENERAL DISTRIBUTION: This species is one of the most widespread species of the Indo-Pacific appearing, itwould seem, wherever the host corals appear, from the head of the Red Sea to east and South Africa, through the Indian and Pacific Oceans, and even extending beyond the Eastern Pacific Barrier to the mouth of the Gulf of California.

It would be expected to occur in the Ryukyus and Southern Japan as the genus Pocillopora appears there, but it has not yet been so reported. Alpheus socialis Helier Fig. 16 Alpheus socialis Helier, 1865:106, pI. 10, fig. 1.

Thomson, 1903:436, pI. 27, figs. 6-12.

Alpheus doto White, 1847:75. (nomen nudum). Previous Australian records Haswell, 1882b:190.


Sayce, 1902:155.

Port Phillip, Victoria.

SPECIMENS EXAMINED: 1 specimen from AM 32 (AM P. 27872); 4, AM 49 (AM P. 27867); 1, AM 76 (AM P. 27845); 7, AM 93 (AM P. 27890); 6, AM 122 (AM P. 27873); 1, AM 150 (AM P. 27843); 2, AM 192 (AM P. 27846); 2, AM214 (AM P. 27933); 1 AM 220 (AM P. 27866); 1, AM 233 (AM P. 27844); 1, AM 289 (AM P. 27847); 2, AM 383 (AM P. 27876); 1, AM 388 (AM P. 27868); 1, AM 395 (AM P. 27848); 1, AM 398 (AM P. 27849); 2, AM P. 3072; 1, AM P. 5029; 1, AM P. 5711; 1, AM P. 6309; 2, AM P. 6526; 3, AM P. 6912; 1, AM P. 8438; 3, AM P. 10092; 1, AM P. 10114; 15, AM P. 11734; 1, AM P. 13546; 1, AM P. 13560; 4, AM P. 13580; 1, AM P. 27874; 2, AM P. 27877.

DIAGNOSIS: As this species is so like A.

Parasocialis sp.

Nov., described and figured I — THE ALPHEIO SHRIMP OF AUSTRALIA 73 thin ridge; socket distally with narrow but deep ‘V’-shaped incision to accommodate ridge.

Carpus cup-shaped, 004 as long as merus, bearing just proximal to inferior margin of chela a small acute tooth, at times of minimal development.

Merus twice as long as broad with distosuperior margin projecting as subacute tooth; inferior margin bearing 7-8 small spines artd a small acute tooth distally.

Ischium bearing 3 spines on inferior margin similar to those of merus.

Small chela with slight sexual dimorphism, varying from 3.0 to 3.6 times as long as broad, with that of female usually more slender than that of male.

Fingers and palm nearly equal.

Palm bearing an acute tooth over dactylar articulation.

Inferior and superior margin bearing bosses similar to those of large chela.

Finger hirsute, at times with hair so dense as to obscure cutting surface on medial face.

Merus similar to that of large chela, but distosuperior margin is not projected.

Ratio of carpal articles of second leg: 10:5:2:2:4.

Ischium of third leg bearing small spine.

Merus 4.6 times as long as broad, usually with inferior margin projecting as small acute tooth.

Carpus 0.5 as long as merus, both distal margins projecting.

Propodus 0.7 as long as merus, bearing on inferior margin 7 spines and a pair distally.

Dactylus 0.3 as long as propodus, biunguiculate.

Telson 3.2 times as long as posterior margin is broad.

Anterior pair of dorsal spines placed anterior to middle.

Posterolateral spines the same size as dorsal spines.

Articulation on outer uropod scalloped.

DISCUSSION: This species is plainly closely related to A.

Sulcatus Kingsley and to other species so related; the separation of these is given in Table 1.

Its closeness to A.

Socialis Helier is most great, and it can be separated by only four characteristics, three of which may be variable.

The most reliable and important is the form of the distal articles of the third maxilliped.


Parasocialis the second article is only slightly expanded distally and the third has a slight taper from the base to the tip.

In A.

Socia/is the development of these articles is unique, with the second bearing a massive, diagonally-truncate lobe, and the third article expanding broadly in the initial third and distally tapering rapidly to a narrow tip; both the lobe and the inferior side and tip of the thi rd article bears many setae about as long as third article.

In addition the carpus of the large chela bears a tooth on the inferodistal margin that is lacking in A.

Socialis, the small chela of the male and female varies from 204-3.0 times as long as broad in this species and bears a meral tooth like the large cheliped while inA.

Socia/is it is stouter, 3.0-4.0 times as long as broad, and lacks the distal tooth.

We believe this species, as it is more generalized and more broadly spread (with the exception of the New Zealand range of A.

Socia/is) to be the form from which A.

Socia/is was derived.

If our hypothesis about A.

Parasocialis being derived from A.

Sulcatus is correct (see discussion under that species) it is notable that this species, having adapted to the cold Australian waters, has re-entered warmer waters both in eastern and western Australia where its range overlaps that of A.


Its range also overlaps with A.

Socialis in New South Wales and extends on either side of A.

Austra/osu/catus sp.

Nov., known only from Tasmania.

Of the 130 specimens in our collection, 60 complete specimens were carefully studied for variation.

It was found that the third maxilliped was at times much more hirsute on the inferior margin than at other times.

The distal tip ofthe dactylus ofthe large chela varied from heavily rounded (fig. 17d) to a definite but rounded tooth.

The small chela varied from 3.0-3.B times as long as broad and usually that of the female was more slender.

The merus of the third leg varied from 4.2 to 4.6 times as long as broad and the 74 DORA M.


BANNER distal tooth varied from a small acute tooth to a sharp angle and in 14 of the specimens the tooth was entirely lacking.

One specimen from Lancelin Is., W.A. (WM 143-65) and one from Tasmania (TM G1538), both males, differ from the usual A.

Parasocialis in the form of their chelae.

On the large chela the palm is normal for the species, but the dactylus is acute at its tip and the plunger is massively developed, and instead of being cylindrical with a thin distal ridge, tapers only slightly distally from maximum diameter to broadly rounded edge.

When the article is viewed from the superior aspect, the distal portion of plunger is seen to make a heavy shoulder on the medial side of the article.

Correspondingly the narrow distal incision of the socket is here a broad and deep “U”-shaped depression to accommodate the heavy distal extension.

In the small chela the fingers were much more hirsute than is usual and appear almost sub-balaeniceps (fig. 17p) but did not have the orderly rows of setae as in A.

Australosulcatus, and the constriction on the inferior margin opposite the articulation of the dactylus was more pronounced than is usual.

However, the rest of the characteristics were li ke those of the holotype and the Tasmanian male was in the same collection as a normal female.

We therefore suggest that these differences may be an extreme variation.

In one specimen from Houtman Abrolhos (WM 185-65) the plunger of the dactylus of the large chela was only of minimal development (fig. 17q) and the fingers were nearly 0.8 as long as the palm instead of 0.4 as in the typical specimen.

The rest of the specimen was usual.

This may be due to a regenerating large chela.

BIOLOGICAL NOTES: Most of these specimens have been collected intertidally, but a few came from as deep as 16 fathoms.

One specimen was reported from a sponge.

The following colour notes were supplied by james and Davemport for 2 specimens from Houtman Abrolhos (AC C-1) “Transparent body.

Orange chela with red tips.

Red to orange on each abdominal segment.

Orange to red carapace”.

Our largest specimen was 35 mm.

AUSTRALIAN DISTRIBUTION: The species ranges from Moreton Bay south and west along the entire southern coast of Australia, including Tasmania, and northward on the western coast to Shark Bay.

No specimens came from the coast of Victoria, but it certainly should be expected to occur there.

DISTRIBUTION OF TYPE MATERIAL: The holotype and allotype will be returned to the Western Australian Museum and the paratypes to their respective museums. Alpheus amir·antei sizou Banner and Banner Fig. 18 Alpheus amirantei sizou Banner and Banner, 1967:265.

Crangon amirantei Banner, 1953:87, fig. 29. Confer: Alpheus amirantei Coutiere, 1908a: 15, 1921 :421, pI. 63, fig. 16.

SPECIMEN EXAMINED: 1 specimen from AM 109 (AM P. 27507).

DIAGNOSIS: Rostral carina high, sharp and narrow between eyes, abruptly curving ventrally to short free portion of rostrum; tip of rostrum reaching to end of first third of visible part of first antennular article, posteriorly carina reaching only to posterior end of orbital hoods.

Orbital hoods inflated, projecting anteriorly into round vertical keel.

Orbitorostral margin almost straight, orbitorostral grooves flattened.

Carapace inflated over antennular bases, leaving groove between bases THE ALPHEID SHRIMP OF AUSTRALIA 2 mm L–_-‘—_—l’ 75 b — m b Fig. 25 Alpheus frontalis Milne-Edwards 37 mm male from AM P. 13572.

A, b.

Anterior region, dorsal and lateral view; c, d.

Large chela, lateral face, and merus, medial face; e.

Small cheliped, lateral view, merus held flat; f.

Small chela, lateral face, held flat; g, h.

Small chela medial and superior face; i.

Small chela, inferior face; j.

Small chela, inferolateral face; k.

Second leg; I.

Third leg; m.

Telson. 45 mm female from AM 432.


Small cheliped, lateral face.

A, b, i, j, k, I, m scale a; c, d, e, f, g, h, n scale b. THE ALPHEID SHRIMP OF AUSTRALIA 101 antennular article.

Lateral tooth of scaphocerite strong, tip curved slightly medially and reaching to end of antennular peduncle; squamous portion broad, reaching past middle of third antennular article.

Carpocerite reaching length of third article past that article.

Inferior margin of basicerite with a small acute tooth.

Third maxillipeds stout, ratio of articles: 10:4:7.

Second article broader than long, with large setiferous lobe on inferior margin.

Third article 1.7 times as long as broad, compressed and acutely triangular in section with inferior margin knife-like and superior margin somewhat thicker; long setae on both margins, with most equal to article in length; setae more dense on inferior margin.

Large chela massive, subcylindrical, without grooves, 2.3 times as long as broad, fingers occupying the distal 0.3.

Plunger of dactylus long, tip of dactylus strongly curved and extending slightly beyond pollex when closed.

Entire surface of chela covered with minute papillae.

Merus nearly as long as broad, superodistal margin not projected.

Inferointernal margin with fine irregular teeth and bearing long fine setae; distal angles not projecting.

Small chelipeds sexually dimorphic.

Small chela of males about 0.6 as long as large chela, stout, almost 2.4 times as broad, with fingers approximately equal in length to palm.

Palm compressed, margins rounded with slight rounded longitudinal depressions on either side of low, rounded crest leading to palmar adhesive plaque, and similar shallow depression on lateral face of base of pollex; palm with only scattered setae.

Articular hinge of dactylus set at 45° to vertical axis of palm, closing on pollex from sLiperolateral angle; dactylus broadly expanded, almost petal-like with slight superior ridge distally that terminates in strong, sharply curved terminal tooth; tooth continued on oppositive face of dactylus as a heavy “V”-shaped ridge; tip of tooth with sharp shearing edge that slides past comparable edge on terminal tooth of pollex.

Rounded margin of dactylus on either side of terminal tooth with dense row of setiferous bristles; other bristles along margins and scattered softer setae on flattened oppositive face superior to ridge.

Pollex slightly expanded proximally, tapering to curved terminal tooth, with groove on oppositive face to accommodate ridge of dactylus; inferior margin near palm with numerous long setae; inner margin with row of setiferous bristles of moderate length and with row of shorter setae on proximal portion of lateral margin.

Palm and dactylus with adhesive plaques.

Merus similar to that of large cheliped.

Small chela of female unmodified, less than half length of chela of male of corresponding size, with palm cylindrical, 3.7 times as long as broad, fingers slender and tapering; fingers about half length of palm.

Merus 3.3 times as long as broad, slightly longer than chela and almost 3 times as long as carpus.

Carpal articles of second leg with ratio: 10:4:2:2:3.

Ischium of third leg with spine.

Merus unarmed, 3.5 times as long as broad.

Carpus 0.4 as long as merus, terminating inferiorly in acute, superiorly in obtuse tooth.

Propodus 0.5 as long as merus and bearing on its inferior margin 7 pairs of spines and a pair distally.

Dactylus simple, curved.

Posterior margin offirst abdominal pleura of male rounded, but more narrow than in female.

Telson 2.3 times as long as posterior margin is wide.

Spines on dorsal surface small, anterior pair placed at middle.

Outer spine of posterior pair very small.

Articulation of outer uropod straight, not scalloped.

DISCUSSION: The shape of the anterior margin of the carapace is variable in this 102 DORA M.


BANNER species.

It is sometimes shorter than in the specimen figured, sometimes slightly emarginate and the sharpness of the carina is also variable.

De Man (1911 :fig. 79, 79a, b) adequately illustrates this variation.

In immature males in this collection the small chelae have only a minimal development of sexual dimorphic characters, a condition already remarked upon (B&B, 1968:286).

The usual development of the anterior region of the carapace in spite of its variation makes this species easy to separate from all other species.

The 10 mm female from Ambon mentioned by Zehntner (1894:200) was notA.

Frontalis as the first article of the second leg is a third the length of the second article, a condition never approached by this species.

BIOLOGICAL NOTES: For the tube dwelling habits of A.

Frontalis see discussion under the Crinitus Group.

As this species appears to be an obligatory commensal in tubes of algae, we suggest that the label of one of Miya’s specimens (1974:138) which stated that it was found as a commensal with Tridacna crocea was in error.

There appears to be some variation in the colour patterns.

Bruce (1975:24, fig. 4) published a beautiful colour photo of a specimen from Mombasa, Kenya which was dark purple and violet and sprinkled with pale lavender and white spots.

In a personal communication he stated that this species is ” …

Quite common here on Heron Island (Australia) and the colour pattern is still essentially the same, possibly slightly paler only”.

This is similar to the colour pattern reported by Miya (1974: 137) for specimens from the Ryukyus.

Fishelson reports his specimens from the Red Sea have ” …

Pale transparent bodies, and the only blackish spots are the eyes and stomach.

On both sides along the dorsum, there are two rows of white spots on each tergite.

The chelae are marbly … ” However, we cited (1968:286) a report from Dr Cadet Hand that specimens of A.

Frontalis from Kapingamarangi in the Caroline Islands were “purple with a red tail and had blue spots on their bodies”.

Our specimens from the Philippines were pale brown.

This species ranges in depth from the immediate subtidal to appreciable depths, 130 m according to Miya (1974:138), and from a dredge haul from 20-80 fathoms according to Coutiere (1921 :425).

We have specimens up to 45 mm in length.

AUSTRALIAN DISTRIBUTION: In western Australia we have a specimen from Northwest Cape, in eastern Australia we have specimens from the Torres Straits and along the Great Barrier Reef from the Coral Sea to Heron Island.

GENERAL DISTRIBUTION: This species ranges across the Indo-Pacific from East Africa to the Society Islands.

It does not occur in Hawaii. Alpheus pachychirus Stimpson Figs 23 j, k; 26 Alpheus pachychirus Stimpson, 1861 :30.

De Man, 1890:116, pI. 6, fig. 14.

Ortmann, 1890:487, pI. 36, figs. 17 a-k.

Cowles, 1913: 121, figs. 1,2. Previous Australian records: Coutiere, 1900:414.

Murray Is., Torres Straits.

SPECIMENS EXAMINED: 1 specimen from AM 74 (AM P. 27501); 1, AM 305 (AM P. 27780); 2, AM P. 7521; 2 AM P. 13568; 2, BAU 31.

DIAGNOSIS: Frontal margin projecting and broadly convex across front from orbital hood to orbital hood, with on Iy a trace of orbitorostral concavity and only a slight obtuse triangle for rostrum.

Rostrum bearing low, but distinct, carina reaching posterior to eyes.

Orbital hoods inflated, orbitorostral area almost flat. THE AlPHEID SHRIMP OF AUSTRALIA — THE ALPHEID SHRIMP OF AUSTRALIA 129 and with its tip projecting, and to our knowledge none have the almost complete lack of orbitorostral grooves.

This species is plainly different from all those in the Diadema Group that have deep orbitorostral grooves, with or without associated teeth such as A.

Diadema Dana or A.

Bidens (Olivier), and from those in which the large chela is more or less round in section.

However, the species does resembleA.

Paracrinitus Miers in many ways: the form of the anterior carapace is quite similar, the lack of sculpturing on the large chela is the same (but it is only 1.4 times as broad as thick in A.

Paracrinitus); the armature of the merus of the large chela is almost identical, even the third maxillipeds, second legs and telson show similarities, but those are of rather generalized development.

Other species in the Diadema Group showing some, but fewer, similarities areA.

Mitis Dana and A.

Tenuipes De Man.

From all members of the Diadema Group this species may be differentiated by the development of the squame of the scaphocerite, the compression of the large chela and the proportions between the fingers and palm in that appendage and especially the long fingers of the small chela.

We have decided to assign it to the Diadema Group because it shows none of the specialized adaptations for life on a muddy or silty bottom found usually in the Brevirostris Group; we suspect that the walking legs, when other specimens are found, will also show similar lack of adaptation.

Nothing is known of the habitat beyond the facts given on the label, reproduced above.

The name is derived from labis, Greek for forceps and refers to the form of the small chela.

The holotype will be placed in the Australian Museum.

Alpheus paracrinitus Miers Fig. 36 Alpheus paracrinitus Miers, 1881 :365, pI. 16, fig. 6.

Chace, 1962 :609.

Crosnier and Forest, 1966:253, fig. 15.

Banner and Banner, 1967:278, 2 tbls. Alpheus paracrinitus bengalensis Coutiere, 1905a:901, pI. 82, fig. 37.

Alpheus bengalensis Holthuis, 1958:25.

Crangon paracrinita bengalensis Banner, 1953:110, fig. 40.

Crangon togatus Armstrong, 1940:2, fig. 1. SPECIMENS EXAMINED: 1 specimen from AM 109 (AM P. 28119); 2, BAU 46.

DIAGNOSIS: Rostrum triangular, acute, somewhat longer than wide at base, tip almost reaching middle of visible part of first antennular article, rounded dorsally; base separated from anterior orbital hoods by short shallow and rounded depressions.

Orbital hoods not inflated, rounded anteriorly.

Orbitorostral margin only slightly concave.

Second antennular article 2 times as long as wide, 1.5 times longer than third article.

Tip of stylocerite reaching to end of first antennular article.

Lateral spine of scaphocerite not reaching end of thi rd antennular article, squame slightly shorter.

Carpocerite reaching by length of third article past that article.

Lateral spine of basicerite small but acute.

Large chela slightly compressed, without sculptu ring except for a slight concavity on inferior margin in region of dactylar articulation.

Chela 3 times as long as broad, fingers about 0.3 length of entire chela.

Plunger of dactylus fully developed.

Merus slender, 3.4 times as long as wide, armed with strong tooth on inferointernal margin slightly distal to middle, superodistal margin not projected. -” w o CJ o ~ 5 mm Ib — mm ,a Fig. 36 Alpheus paracrinitus Miers 17 mm male from BAU 46.

A, b.

Anterior region, dorsal and lateral view; Co large cheliped, medial face; d.

Small chela, lateral face; e.

Merus, small cheliped, medial face; f.

Second leg; g.

Third leg; h.


A, b, h scale a; c, d, e, f, g scale b. THE ALPHEIO SHRIMP OF AUSTRALIA 131 Small chela sexually dimorphic.

Male chela 4.2 times as long as broad with fingers occupying half entire length, dactylus variable, at times balaeniceps.

Female chela more slender, tapering.

Meri in both sexes with spine on inferointernal margin similar to that of large chela.

Carpal articles of second legs with fi rst and second article varyi ng in relative lengths: 10:8-18:3:3:4. (From B&B, 1966b:116).

Ischium of third leg armed with small spine, merus 7 times as long as broad, inermous.

Carpus half as long as merus, both distal margins projected but rounded.

Propodus 0.7 as long as merus and armed with 6-8 movable spines on inferior margin.

Dactylus simple, curved and elongate.

Telson 3 times as long as posterior margin is broad.

Anterior pair of dorsal spines placed anterior to middle.

DISCUSSION: Miers type locality was Senegambia, West Africa; Coutiere described the variety bengalensis from the Maldives in the Indian Ocean.

Holthuis believed the Atlantic form was separate from that found in the I ndo-Pacific and raised A.

Bengalensis to specific level; finally, Chace, comparing West African specimens to some from Clipperton Island, came to the conclusion that there was but a single circumtropical species without distinct characteristics that would warrant the separation of subspecies.

Crosnier and Forest placed into synonymy A.

Togatus that Armstrong had described from Bermuda.

In 1967 we reviewed the extent of variation in our central Pacific collections which included over 200 specimens from the Cook and Society Islands alone, and found extensive variation in the armature of the large and small cheliped, and the degree of balaeniceps development on the small chela of the male in the ratio of the first two carpal articles of the second leg, and in the carpal-propodal length ratio of the third legs.

As these characteristics were those that were supposed to separate the two subspecies or species, we confirmed Chace’s conclusion.

We have examined the type of A.

Togatus at the American Museum of Natural History in New York and agree with the action of Crosnier and Forest.

BIOLOGICAL NOTES: The species apparently is confined to the intertidal and the upper subtidal with the greatest depth reported to be 18 m; it occurs under boulders in dead coral heads and in coralline algae.

Miya (1974:159) reported that it has also been found from heads of Tubipora.

No one has published colour notes on the species, but our notes made in Hawaii showed that the entire specimen was transparent white with transverse bright red bands.

The bands were broad and diffuse on the large and small chelae, slight on the anterior carapace, more defined but confined to the middle on the posterior carapace, and narrow and well-defined on each abdominal segment, running from pleuron to pleuron; both the antennular bases and the tip of the telson and uropods were splashed with red stellate chromatophores.

We have observed similar colouration in other parts of the Pacific.

The largest specimen we have seen was 20 mm in length.

AUSTRALIAN DISTRIBUTION: Only 3 specimens have been collected, one from the Herald Cay in the Coral Sea and 2 from Heron Island on the Great Barrier Reef; considering how common it has been found to be in the central Pacific, it is surprising that more specimens have not been collected in Australian waters. 132 DORA M.


BANNER GENERAL DISTRIBUTION: As indicated, the species appears to be circumtropical, in the Indo-Pacific running from the Red Sea through the Hawaiian and Society Islands to Clipperton Island lying west of Central America; from the Ryukyus in the north to the Capricorn Group in the south; in the Atlantic from the Gulf of Mexico and the Caribbean to West Africa.

Alpheus ehlersii De Man Fig. 37 Alpheus ehlersii* De Man, 1909c:663, pI. 70.

Banner and Banner, 1966b:114, fig. 40. Alpheus macrochirus De Man, 1888a:519. (Nec Richters, 1880). SPECIMENS EXAMINED: 1 specimen from BAU 16; 1, BAU 20; 7, BAU 29; 1, BAU 43; 2, BAU 44.

DIAGNOSIS: Rostrum acute, reaching well past middle of visible part of first antennular article.

Rostral carina rounded, reaching to posterior margin of slightly inflated orbital hoods, orbitorostral grooves not pronounced.

Orbitorostral margin bearing slight arcuate prominences.

Visible part of first antennular article and second article equal, second article almost 2 times as long as broad, and 1.5 times as long as third article.

Stylocerite acute, reaching past end of first antennular article.

Lateral tooth of scaphocerite reaching to end of antennular peduncle, somewhat longer than narrow squamous portion and turned inward at tip.

Carpocerite reaching beyond end of antennular peduncles.

Basicerite with strong tooth.

Large chela slightly compressed, 2.7 times as long as broad, fingers not quite 0.3 of total length; superior margin proxi mal to dactylus beari ng shallow transverse groove that extends obliquely toward medial face; groove so faint that chela often must be rotated to discern it; inferior margin bearing slight constriction opposite articulation of dactylus.

Plunger of dactylus well developed.

Merus 2.2 times as long as broad with inferointernal margin armed with 6-9 spines and bearing acute tooth distally.

Superior margin not projecting.

Merus of female more slender, but with similar armature.

Small chelipeds not sexually dimorphic, chela 4 times as long as broad with fingers a little longer than palm; rounded tooth on medial side of dactylar articulation.

Merus 2.6 tim::.’ as long as broad, bearing several small spines on inferointernal margin and distally a rounded tooth.

Carpal articles of second leg with ratio: 10:5:2:2:4.

Ischium of third and fourth legs with spine.

Merus ofthird leg inermous, 4.3 times as long as broad.

Carpus 0.6 as long as merus, superior margin projected into a rounded distal tooth, inferior margin truncate distally.

Propodus 0.7 as long as merus, bearing on its inferior margin 5 spines and a pair distally.

Dactylus simple, 0.3 as long as propodus.

Telson 2.5 times as long as posterior margin is broad.

Anterior pair of dorsal spines placed just anterior to middle.

Inner spines of posterolateral pair the same size as dorsal spines.

DISCUSSION: The holotype of this species is apparently missing so we were unable *Spelled ehlersi according to the International Code of Zoological Nomenclature of 1961; ehlersii by the code of 1964, but not again changed in the revisions of 1974.

These changes in rules have also been applied to the names Synalpheus stimpsonii (B&B, 1975:292) and Alpheus edwardsii (p. 404) Rule 32 (a)(ii) also requires that Athanas haswelfi Coutiere as used by Hale, 1927:47 and B&B, 1973:316 be returned to its origi.nal spelling of A.

Hasswelfi. THE ALPHEID SHRIMP OF AUSTRALIA 133 — Fig. 44 Afpheus acutocarinatus De Man 35 mm female from UQ 18.

A, b.

Anterior region, lateral and dorsal view; Co third maxilliped; d, e.

Large chela and merus, medial face; f.

Small chela, medial face; g.

Second leg; h.

Third leg; i.


After De Man, 1911, fig. 94D.


Cheliped of male.

All drawings same scale except j. THE AlPHEIO SHRIMP OF AUSTRALIA 153 Ischium of third legs bearing small spine.

Merus inermous, 8 times as long as broad.

Carpus 0.5 as long as merus, superior margin projected as an obtuse tooth.

Propodus 0.6 as long as merus, inferior margins without spines, but both superior and inferior margins bearing long fine setae; two other rows of short setae on flattened faces, parallel to margins.

Dactylus 0.4 as long as propodus, spatulate, slightly excavate on its inferior surface.

Telson elongate, 33 times as long as posterior margin is wide, 1.9 times as wide anteriorly as posteriorly; lateral margins concave; strongly arcuate tip extending by 0.15 of total telsal length beyond small posterolateral spines.

DISCUSSION: We have compared our specimens to De Man’s holotype in the Zoologisch Museum in Amsterdam and find only one slight difference.

The third leg on our 3 specimens had a merus that was 9 rather than 8 times as long as broad.

De Man noted, butfailed to show, the ischial spine on thethird leg; it is present in the holotype as well as the Australian specimens.

The extent of variation in this species cannot be projected from the 8 specimens ever reported.

BIOLOGICAL NOTES: The specimens from Moreton Bay were from 10 fathoms and De Man’s 5 specimens were dredged from 26-72 m.

He reported them from muddy bottoms and presumably the habitat in Moreton Bay was similar.

The Thai specimens were also dredged and probably from sandy-to-muddy bottoms.

AUSTRALIAN DISTRIBUTION: The only record is from Moreton Bay, Queensland.

GENERAL DISTRIBUTION: Indonesia and Gulf of Thailand.

Alpheus stephensoni Banner and Smalley Figs. 45, 46 Alpheus stephensoni Banner and Smalley, 1969:43, fig. 2. Previous Australian records: Banner and Smalley, (lac cit.J.

Moreton Bay, Qld.

SPECIMENS EXAMINED: 1 specimen from AM P. 12927; 1, AM P. 12936; 1, QM W 2238; 1, UQ 27; 21, UQ 30; 32, UQ 31; 39, UQ 32; 3, UQ 34; 1, VM 23.

DIAGNOSIS: “Rostrum acute, awl-shaped, reaching past end of first antennular article.

Rostral carina pronounced, knife-edged, disappearing abruptly at 0.4 length of carapace.

Orbital hoods inflated, forming moderately deep grooves between hoods and high rostral carina.

Orbitorostral area set off distinctly from lateral regions of carapace, more narrow in relation to carapace than usual for Alpheus.

Carapace covered with small bosses.

Second article of antennular peduncle 3 times as long as broad.

Visible part of fi rst antennular article and third article sub-equal, first article about one-third as long as second.

Lateral margins of antennular peduncles beset with dense, fine hair.

Antennular flagella somewhat shorter than body length; antennal flagella somewhat longer than body.

Lateral margin of scaphocerite straight, lateral spine only slightly longer than squamous portion.

Squamous portion narrow distally.

Carpocerite reaching to end of second antennular article, antennae long, often as long as entire shrimp.

Stylocerite acute, reaching to end of first antennular article, lateral margin fringed with fine hairs.

Basicerite with strong lateral spine.

Tip of third maxilliped reaching beyond end of antennular peduncles when extended. “large chela compressed, 3.7 times as long as broad.

Palm covered with small bosses, less abundant near distal end; superior margin rounded, bearing towards the 154 DORA M.


BANNER < ., ~g ,'". ""~~ t<~t, ... """~" L--_ _ _ _ -- ..... Fig. 46 Alpheus stephensoni, regenerating chelae 75 mm male from AM 347.


Large cheliped, medial face; b.

View of oppositive face of pollex showing developing socket; c.

Small cheliped (fingers broken).

A, c scale a; b scale b. V1 V1 156 DORA M.


BANNER inner face a row of forward-sweepi ng hairs; inferior margin flattened and bearing rows of hairs similar to superior margin; inner face flat; outer face a little rounded toward the superior margi n but longitudinally slightly grooved near the inferior margin; groove not extending into finger.

Dactylus 0.38 as long as entire chela, superior margin slightly flattened bearing a row of fine setae on inner margin of flattened portion.

Fingers slightly crossing; surfaces with a few long setae, but no bosses.

Merus 3.7 times as long as broad, surface covered with small bosses.

Distal end of superior margin not armed, inferior internal margin bearing row of fine hairs, distal end with strong acute tooth.

Inner face bearing a shallow longitudinal groove near superior margin.

Large chela offemale slightly smaller than male chela. “Small chela of male highly compressed, 4.8 times as long as broad, entire surface covered with small bosses.

Fingers only slightly longer than palm.

Inner face of chela beset with forward-sweeping hairs, superior and inferior margins rounded.

Superior margin of dactylus with sharp crest.

Fingers bearing dense fringe of setae along lateral margins of opposing faces.

Fingers gaping, tips crossing.

Merus 3.6 times as long as broad, surface covered with small bosses.

Inferointernal margin bearing fine hairs, distal end armed with pronounced tooth.

Inner face bearing a narrow longitudinal groove near superior margin, similar to groove on merus of large chela. “Small chela of female highly compressed, about one-third as large as that of male, 6.5 times as long as broad, fingers a little longer than palm.

Dactylus slightly shorter than fixed fingers.

Fingers with patches of short stiff setae near opposing surfaces, but no fri nge of setae as fou nd in male.

Pal mar su rface with bosses si mi lar to large chelae.

Inner face partially covered with forward-sweeping setae set into superior and inferior margins.

Merus similar to that for large chela with tooth on inferodistal margin and a slight narrow groove longitudinally near the superior margin. “Carpal articles of second leg with ratio: 10:8:3:3:3. “Ischium of third leg with small spine.

Merus 6 times as long as broad, unarmed.

Carpus almost half as long as merus.

Propodus 0.6 as long as merus, without movable spinules but with 3 longitudinal rows of stiff setae.

Dactylus spatulate, 0.3 as long as merus. “Telson 3.2 times as long as posterior margin is broad.

Posterior margin strongly arcuate with rounded tip extending well beyond lateral spines.” (Original description).

DISCUSSION: The variation in this species is discussed in the 1969 paper.

The chelae on the 75 mm male specimen from AM 347 are anamolous (fig. 46).

The large chela is 2.8 times as long as broad, with the fingers slightly longer than the pal m; the socket of pollex is well demarked and relatively deep, but the plunger of the dactylus is only slightly developed and cannot reach the bottom of the socket.

The distal portions of the fingers carry knife-like edges, meeting only in the middle third and gaping in the distal third; tips curved and crossing.

The large chela is covered with rounded bosses and hairs.

The small chela is simple with distal portion of fingers broken off; the fingers, where broken, are oval in section with oppositive faces slightly angular and bearing rows of short stiff setae that cross.

The palm is relatively smooth and bears only slightly developed rows of setae.

Otherwise the specimen falls well within the range of variation of A.


We were originally going to give the specimen a separate description as a questionable A.

Stephensoni, but Dr.

John C.

Yaldwyn has suggested to us that it was likely that the specimen had lost its large chela and was in a series of moults in which the small chela was changing to the form of the large chela and the regenerating chela was being transformed into a normal small chela.

This phenomenon is well known, but THE AlPHEID SHRIMP OF AUSTRALIA seldom have we seen chelae of such intermediate development. 157 BIOLOGICAL NOTES: This species has been largely caught in shrimp trawls in water up to 10 fms.

It has been caught especially at night and seems to be more abundant in the summer months.

Specimens caught in early spring often are heavily encrusted with hydroids and tube worm cases.

All alpheids caught on muddy grounds east of Redcliffe, Qld.

Are likely to be this species. “The carapace is blue-green dorsally; rostral carina is brown.

Abdomen olive dorsally, and often red laterally; along lower edges of pleura is a white band continuous with the white branchiostegites.

Tergum markings usually include brown transverse band along posterior edge of each tergum, various small colourless patches assuming bilateral symmetry, and a medial colourless line on anterior of first tergum and on sixth tergum continuing posteriorly on telson.

Telson also with two white patches proximally and a white spot around base of each of the four spines. “Antennules and antennae mostly brown, but blue-green at bases.

Inner faces of large and small chelipeds with proximal ends of fingers, palm, and carpus, and distal end of merus all olive mottled on pale pink background, but for blue inferior internal edge of palm; most of fingers blue over pale pink with extreme tips white.

Outer faces of chelae purplish or pink.

Uropod reddish with blue and brown borders.

Other appendages red distally, white proximally.” (Colour notes by Smalley, lac.


AUSTRALIAN DISTRIBUTION: This species has been reported in eastern Australia from Bundaberg, Qld.

To Port Jackson, N.S.W.

GENERAL DISTRIBUTION: This species has never been reported out of Australia. Alpheus distinguendus De Man Fig. 47 Alpheus distinguendus De Man, 1909b: 155, pI. 7, figs. 9-14.

Banner and Smalley, 1969:47, fig. 3. Alpheus rapax De Haan, 1850:177, pI. 45, fig. 2.? Bate, 1888:552, pI. 99, fig. 1.

De Man, 1888b:264; 1892:404.

Ortmann, 1890:481.

Nobili, 1903:7. (Nec Fabricius, 1798.) ? Alpheus brevirostris De Man, 1907:427, pI. 33, figs. 51, 52. ? Alpheus digitalis De Haan, 1850:178, pI. 45, fig. 4.

Coutiere, 1898h:249, fig. 2 (=A rapax De Haan).

Previous Australian record: Banner and Smalley, lac.


Moreton Bay, Qld.

SPECIMENS EXAMINED: 1 specimen from AM 99 (AM P. 27771); 1, AM 125; (AM P. 27530); 1, AM 148; (AM P 27869); 1, AM 354; (AM P. 27536); 1, AM P. 14630; 1 each from CS 50,51;1, MM 355; 1, QM W 2381; 1, UQ 17; 1, UQ19, 1, UQ 20; 32, UQ 28; 20, UQ 29; 1, UQ 33; 1, WM 134-65; 1, WM 162-65; 1, WM 199-65; 2, WM 237-65; 1, WM 241-65; 1, WM 242-65; 5, WM 284-65; 1, WM 131-76.

DIAGNOSIS: Rostrum awl-shaped, reaching to near end of first antennular article.

Dorsal carina high and narrow, extending into carapace beyond orbits.

Eye hoods inflated, forming broad deep grooves between eye hoods and dorsal carina.

Visible part of first antennu lar article equal to thi rd.

Second article 3 times as long as third article and 3 times as long as wide.

Stylocerite reaching to end of first antennular article, acute tip not — 10 mm la Fig. 47 Alpheus distinguendus De Man a.

Anterior region, dorsal view; b, c.

Large cheliped, lateral and medial face; d, e.

Small chela and merus, medial face; f.

Small cheliped, lateral face; g, h.

Small chela and merus, female, medial face; i.

Second leg; j.

Third leg.

A, b, c, d, e, f.

H, i, j scale a; g, h scale b. (Figures of 74 mm male and 70 mm female.

After Banner and Smalley, 1969, fig. 3.) THE AlPHEID SHRIMP OF AUSTRALIA 159 directed medially, lateral margin of stylocerite and antennules beset with row of dense, fine setae.

Scaphocerite with outer margin slightly concave, reaching well beyond antennular peduncle with tip of lateral tooth slightly longer than squamous portion.

Carpocerite as long as antennular peduncle.

Basicerite bearing small acute tooth.

Large chela compressed, 3 times as long as wide, with fingers occupying distal 0.3.

Superior margin without transverse groove, but bearing along its entire length a flattened angled slant toward the outer face at the distal end of palm.

Inner edge of flattened area bearing fine hairs di rected forward.

Lateral face bearing a diffuse longitudinal ridge about the middle of the face.

Medial face flat.

Inferior margin of chela knife-like, and bearing fine hai rs directed forward along its entire length.

Entire chela bearing fine bosses.

Merus 2.7 times as long as broad.

Superior margin terminating in an acute tooth.

Inferointernal margin lightly serrate and bearing 4 movable spinules, terminating distally in an acute tooth.

Inferoexternal margin without spinules, but bearing fine serrations along margin.

Small chela of male slender, 4.5 times as long as wide, palm covered with minute bosses.

Fingers 2.8 times as long as palm, fingers crossing at tips.

Dactylus longer than fixed finger.

Opposing margins of fingers bearing brush of forward directed hairs.

Outer faces of fingers show gape only near distal end when closed, but inner face shows wide arched gape along its entire length giving an excavate appearance.

Opposing faces of fingers heavily hirsute.

Merus similar to that of large chela.

Small chela of female about half the size of male small chela.

Fingers 1.7times as long as palm, fingers slightly gaping, crossing at tips.

Opposing margins bearing slight row of cri ss-cross hairs.

Patches of setae on lateral margins of fingers.

Superior and inferior margin carrying long fine, forward-sweeping hairs.

Merus similar to that for male but slightly more slender.

Ratio of carpal articles ot second leg: 10:10:5:4:4.

Ischium of third legs with spine.

Merus 7 times as long as wide, without spines.

Carpus half as long as merus; propodus a little longer than carpus, bearing 2 rows of short hairs on its lateral face and inferior margin.

Dactylus 0.5 as long as propodus, spatulate.

Telson 2.5 times as long as posterior margin is broad, proximal margin 1.5 times wider than distal.

Distal margin broadly rou nded. (Description modified from Banner and Smalley, loc.


DISCUSSION: When we redescribed A.

Distinguendus from specimens from Australia in 1969 we suggested that the overhang of the pollex of the dactylus of the small ch~la of the male might separate the Australian form from the Japanese form, the type locality. (The differences in finger length appearing in fig. 47d,f are the result of differing angles of view.) Since that time Dr.

Yasuhiko Miya of the Institute of Biology, Nagasaki University, Japan has informed us that the specimens in Japan also have the overhanging dactylus (personal communication).

The variation in the length-breadth ratio of the chelae that Miya found was also similar to that found in the Australian specimens.

After over 20 years of study during which time he was trying to separate various authors’ use of the names A.

Brevirostris (Olivier) and A.

Rapax Fabricius, De Man came to the conclusion that almost all of the specimens he had placed under A.

Brevirostris were in actuality a different species to which he gave the name A.


As we indicate under A.

Brevirostris, we have sincere doubts about the validity of this separation, but are continuing to use part of De Man’s separation between the two nominal species.

Coutiere after studying the specimens of De Haan from Japan, reported (1898h) that 160 DORA M.



Digitalis De Haan, whose distinguishing characteristic was a strange large chela that Coutiere decided was an anomalous form due to regeneration, was the form that De Haan had named A.

Rapax Fabricius (1850:177), but that De Haan’sA.

Rapax was actually A.

Brevirostris (Olivier).

Later, De Man (1909b:155) in his reconsideration of the Brevirostris Group, stated that De Haan’sA.

Rapax and part of the specimens Coutiere considered to beA.

Brevirostris belonged to his, De Man’s, new species,A.


Presumably, therefore, A.

Digitalis is a synonym of A.

Distinguendus, but this synonymy was never listed by De Man.

BIOLOGICAL NOTES: The specimens discussed in 1969 (lac.

Cit.) from Moreton Bay as well as most of the abovementioned specimens were caught in prawn trawls in less than 20 fathoms.

Miya reports that specimens from Japan are caught on summer nights in prawn trawls, further he stated that in” …

Hakata Bay, Fu kuoka City, they are caught with the mantis shrimps Oratosquilfa oratoria (De Haan) … ” (Personal communication).

The collecting notes indicated in the Australian specimens that they are commonly found in brackish water at the mouth of rivers.

This species appears to live on the bottom or slightly above it.

One 54 mm specimen from the Gulf of Carpentaria (AM 345) had an abnormal growth on the carapace giving it a “furry” appearance.

The growth was tentatively identified by Dr.

Michael Hadfield of the University of Hawaii as the bryozoan, Triticella sp.

This genus is known to be attached to the exoskeletonof crustaceans (see: Hyman, 1959:431), but we previously have not seen any alpheids with this epizoic.

In life the shrimp is green to brown dorsally, colourless to pale blue laterally, with longitudinal streaks on abdomen; tips of chelae orange to pink; legs lighter in colour (for full colour notes see Banner and Smalley, lac.


Our specimens range up to 80 mm in length.

AUSTRALIAN DISTRIBUTION: This species has been collected in Western Australia from near Perth to Shark Bay and Kuri Bay in northwestern Australia; in northern Australia in the Gulf of Carpentaria and in eastern Australia from c:ooktown, Qld.

To Botany Bay, N.S.W.

GENERAL DISTRIBUTION: Mergui Archipelago; Singapore; China; Japan. (De Man was of the opinion that many of the earlier records of A.

Rapax Fabricius were actually A.


As there can be no way to retroactively sort them, we have ignored them here.

The range, therefore, may be much broader). Alpheus rapacida De Man Fig. 48 Alpheus rapacida De Man 1908:105; 1911:394, fig. 91.

Barnard, 1950:750, figs. 142 a-f. Tiwari, 1963:302, fig. 21,22.

Lewinsohn and Holthuis, 1964:47, fig. 1.

Banner and Banner, 1966b:118, fig. 42. Alpheus rapax Bate, 1888:552, pI. 99, fig. 1. (Nec Fabricius , 1798.) — , b Fig. 48 Alpheus rapacida De Man 57 mm female from AM 288.

A, b.

Anterior region, lateral and dorsal view; c.

Third maxilliped, medial face; d, e.

Large chela and merus, medial face; f, g.

Small chela and merus, medial face; h.

Second leg; i.

Third leg; ;.


A, b, c, d, e, f, g, j scale a; h, i scale b. 162 DORA M.


BANNER reaching to middle of third antennular article.

Basicerite with small acute tooth on i nferio r margi n.

Ratio of articles of third maxilliped: 10:3:7.

Inferodistal margin of second article bearing patch of long setae most of which surpass tip of distal article.

Tip of distal article bearing long setae.

Third maxilliped exceeds length of antennules by half length of distal article.

Large chela compressed, 4 times as long as broad with fingers occupying distal 0.4.

Upper margin of palm with flattened area, not cut by a transverse groove; medial margin of area carrying a row of long, forward-sweeping bristles; outer margirl with row of sparsely set shorter hairs.

Inferior margin thin but rounded bearing a row of setae directedanterolaterally.

Entire surface of palm and proximal section of propodal finger granular.

Dactylus smooth, superior margin carinate, tip truncate and slightly shorter than pollex.

Plunger of dactylus of minimal development.

Carpus short, slightly granular and sparsely hi rsute.

Merus 3.4 times as long as broad, su rface slightly gran ular.

Superior margin bearing obtuse tooth distally; inferointernal margin dentate, bearing a number of small spines and an acute tooth distally.

Inferior margin of ischium also bearing a few small spines.

Medial surface of both merus and ischium with narrow, deep, longitudinal groove located slightly superior to middle.

Small chelae of males and females similar, about as long as large chela, without ridges or grooves, 5.3 times as long as broad, fingers 1.8 times longer than palm.

Inferior and superior margin bearing long forward sweeping hairs similar to large chela; surface of fingers bearing sparsely set, short, stiff setae.

Opposing edges of fingers beset with short forward-sweeping hairs that cross in middle.

Tips of fingers cross when closed, leaving a gape proximally.

Merus similar to that of large chela.

Ratio of articles of second leg: 10:10:3:3:3.

Ischium of third leg with spine.

Merus inermous, 5 times as long as broad.

Carpus 0.4 as long as merus, superior margin projected slightly as a rounded tooth.

Propodus 0.6 as long as merus, without spines on inferior margin, but with row of long setae; a second row of setae on lateral face with short and stiff bristles set in groups of 2 or 3; superior margin bearing similar bristles, but set singly.

Dactylus spatulate and more than 0.4 length of propodus.

Telson 2.5 times as long as posterior margin is broad.

Posterior margin arcuate, projecting far behind posterolateral spines.

Anterior pair of dorsal spines set well anterior to middle. DISCUSSION: The Australian specimens differ from De Man’s in the following characters: 1.

The rostrum reaches to near the end of the first antennular article instead of the middle. 2.

The second’antennular article is much longer in relation to the first and third with a ratio of 10:30:10 instead of De Man’s ratio of 10:12:6. 3.

The third maxillipeds reach beyond the antennules in our specimens instead of equal as in De Man’s.

In addition, specimens in our collections from Hawaii and Madagascar have a slight transverse groove behind the dactylus of the large chela, a condition not noted in the Australian specimens.

This seemed also to be indicated in Tiwari’s figure of a specimen from Vietnam. THE ALPHEID SHRIMP OF AUSTRALIA 163 The first three differences were remarked upon by Lewinsohn and Holthuis (lac.

Cd.) with specimens from Israel and by Tiwari with his specimens from Vietnam; we find the same from Hawaii.

We do not attach importance to any of these differences.

We should remark that the condition of the large chela shown in figure 48d in which the dactylus is shorter than the pollex is not common.

We have examined the specimen from Hong Kong collected by the Challenger Expedition at the British Museum (Natural History) and identified by Bate as A.

Rapax, who probably based his identification upon De Haan’s figure of what he, De Haan, thought was that species.

Later, when De Man separated A.

Rapax Fabricius from the form De Haan had described, giving De Haan’s form the nameA.

Distinguendus, he did not know what to do with Bate’s reference so merely listed it “?Alpheus rapax Spence-Bate” in the synonymy of A.


The only difference between Bate’s specimen and A.

Rapacida was the very long lateral spine of the scaphocerite as shown in his figures; our examination of the specimen shows the drawing to be inaccurate and we find this specimen to be A.

Rapacida. BIOLOGICAL NOTES: This species appears to be a form that burrows in sandy to muddy bottoms.

It has been collected as deep as 56 m.

The specimens in our collections from Hawaii were from the intertidal region and were found in association with the gobiid fish Psilogobius mainlandi Baldwin.


Randall of the Bishop Museum, Honolulu, Hawaii, caught this species in 2 metres of water near One Tree Island, Great Barrier Reef, in association with an, as yet to be identified, goby.

We have no colour notes from Australian specimens butthe specimens we have from Hawaii are almost transparent with widely dispersed red stellate chromatophores.

The lateral margins of the carapace are delicately coloured with a shifting band of rainbow colours and the anterior portion of the carapace carry a sprinkling of lemon-green pigment spots.

The eyes are pale violet and the eggs green.

When a living specimen is twisted in the light the whole animal displays a delicate iridescence.

Our largest Australian specimen is 57 mm in length. AUSTRALIAN DISTRIBUTION.

We have specimens from Shark Bay in Western Australia; in northern Australia from the Gulf of Carpentaria and from eastern Australia from One Tree Island in the Capricorn Group to Yamba, N.S.W.

GENERAL DISTRIBUTION: South Africa; Red Sea; Mediterranean coast of Israel (a migrant species through the Suez Canal – Steinitz, 1967:167); Singapore; Thailand; Vietnam; Indonesia and Hawaii. Alpheus barbatus Fig. 49 — -1,b Fig. 72 Alpheus strenuus cremnus subspec.

Novo Holotype (male).


Anterior region, dorsal view; b.

Basicerite, lateral view; e, d, e.

Large cheliped, medial and lateral face and dactylus; f, g.

Small cheliped, lateral and medial face; h.

Second leg; i, j.

Third leg and enlarged propodus and dactylus; k.


B, c, d, e, f, g, h, i scale a; a, j, k scale b. THE AlPHEID SHRIMP OF AUSTRALIA spine of scaphocerite.

Basicerite bearing acute slender tooth.

Ratio of articles of third maxilliped: 10:3:7. 231 Large chela compressed, massive, 2.5 times as long as broad with fingers occupying approximately the distal third.

Superior saddle deep and quite narrow; proximal shoulder slightly overhanging saddle, distal shoulder prominent and initially abrupt.

Lateral palmar depression well defined, quadrangular, reaching proximally to linea impressa.

Medial palmar depression a narrow triangle with apex reaching to proximal third of chela.

Inferior shoulder heavy and rounded; inferior notch prominent.

Inferolateral depression continued up face of palm 0.2 total width, disappearing into pollex distally.

Inferomedial depression small, but continuing proximally as a narrow longitudinal groove almost reaching to articulation at proximal E!nd of chela.

Medial face of chela sparsely hirsute, lateral face glabrous.

Plunger of dactylus only moderately developed.

Merus 1.6 times as long as broad; and inferoexternal and superior margins rounded distally; inferointernal margin with acute tooth distally.

Small chela of both male and female of balaeniceps form.

Cheliped 3.4 times as long as broad; fingers a little shorter than palm.

Superior and inferior margin of palm bearing shoulders, grooves and depressions similarto palm of large chela but not as pronounced.

Articulation of dactylus flanked medially by a small obtuse projection.

Adhesive plaque of dactylus borne on flattened triangular areas, laterally rounded, but demarked medially by rounded ridge that continues distally, disappearing slightly proximal to point of union of balaeniceps setiferous crests; ridge proximally bearing scattered setae.

Medial face of entire chela bearing long forward-sweeping hairs, more distally than proximally; lateral face glabrous.

Merus similar to that of large chela, 2.7 times as long as wide.

Second leg with ratio of carpal articles: 10:8:2:2:5.

Ischium of third leg bearing tooth.

Merus 3.8 times as long as broad, inermous; carpus 0.6 as long as merus, distosuperior and distoinferior margins slightly projected.

Propodus 0.7 as long as merus, inner face bearing 14 spines, roughly paired.

Dactylus triangluar in section, simple, 0.4 as long as propodus.

Telson 1.8 times as long as posterior margin is broad; posterolateral spines small, spines on dorsal surface of telson prominent.

DISCUSSION: This subspecies is identical to the nominate subspecies except in two morphological characteristics and apparently in its ecology.

The most pronounced and consistent difference in the structure in is in the orbital grooves.



Strenuus they are shallow and rounded, with the margins confluent with curvature of rostrum and the orbital hoods; they extend only slightly posterior to the pigmented portions of the eyes.

In A.


Cremnus, they are deeper, broader and extend further posteriorly; medially the sides of the rostrum are more abrupt, and laterally, posterior to the pigmented portion of the eyes, a ridge of the orbital hood overhangs the concave margin of the groove; even at the posterior end the grooves are firmly demarked from the carapace.

The second morphological characteristic is in the sculpturing of the palm and the ridge of the dactylus of the small chela of both males and females.

As remarked under A.


Strenuus, the degree of sculpturing of the palm of the small chela was found to be variable but usually slight in the Central Pacific specimens; those from Australia are also variable but usually have a greater degree of sculpturing.

However, this difference between the Australian and the Central Pacific specimens is not great enough or constant enough to warrant in our opinion the designation of the Australian form as a separate subspecies.

On the other hand the difference in sculpturing of the small chelae between A.


Strenuus and A.


Cremnus is marked and almost always present, with that of A.

S. 232 DORA M.


BANNER cremnus much more like that of the large chela- contrast figs. 71e and nf.

In only a few specimens of A.


Strenuus, as determined by the nature of the orbital grooves, did the sculpturing approach the extreme conditions characteristic of this subspecies.

As these occurred in the waters of New South Wales where evidently the two subspecies coexist, they may be hybrids.

We believe there may be an ecological separation between the two subspecies, as well, but this cannot be proven by the collection data available with the two subspecies.

The two forms overlap geographically.

However A.


Strenuus occurs definitely on coral reefs and coral cays, and along more continental shores, but apparently in clean sand and cleaner waters, whileA.


Cremnus may occur more commonly in muddy and estuarine environments, such as Moreton Bay, the mouth of the Clarence River, near the mouth of Tuggerah Lake, etc.

Yet both subspecies have been collected from Long Reef, Collaroy, on the ocean coast of north Sydney.

If the two forms we are here calling subspecies are indeed found to overlap ecologically, then A.


Cremnus should be considered to be a separate species.

The name is derived from the Greek word cremnos, which means, in part, an overhanging wall or bank and refers to the lateral margins of the orbitorostral grooves.

The holotype and allotype as well as some paratypes will be deposited at the Australian Museum in Sydney, N.S.W.

Paratypes will be placed in the Western Australian Museum, Perth, W.A.

BIOLOGICAL NOTES: The evidently cohabiting pair we selected for the holotype and allotype had the female larger, but in other pairs this size relationship did not hold.

Healy and Yaldwyn, (1970, pI. 18) published a colour picture of a specimen from Long Reef, N.S.W. !AM 399); it shows an overall red cast with the cephalic and thoracic appendages definitely light red.

The carapace is a brownish-red and the abdomen with transverse bands separated by pinkish-white, brown and white irregular bands on caudal fan.

Yaldwyn described the holotype and allotype as “purplish-red with transverse abdominal bands of the same colour”.

He described a similar coloration for four other specimens.

However, one specimen from the Clarence River was noted by Cameron, the collector, as “striped with olive green and white, brick reEl hands.” (AM 132).

AUSTRALIAN DISTRIBUTION: Most of the specimens came from near Sydney and on the other parts of the coast of New South Wales; however, collections of itwere made near Hopetoun (near Albany) from Cape Leveque and Yampi Sound, W.A.

And from Darwin, N.T., the Gulf of Carpentaria and Moreton Bay, Qld.

Alpheus euphrosyne euphrosyne De Man Fig. 73 Alpheus euphrosyne De Man, 1897:745, fig. 64 a-d; 1898b:317, pI. 4, fig. 2.

Banner and Banner, 1966b:130, fig. 49. Alpheus eurydactylus De Man, 1920:109; 1924:48, fig. 17. SPECIMENS EXAMINED: 4 specimens from AM 81 (AM P. 28127); 2, AM 96 (AM P. 28128); 1, AM 162 (AM P. 28129); 1, AM 202 (AM P. 28130); 1, AM 279 (AM P. 28131); 1, AM 310 (AM P. 28132); 1, QM W 2248.

DIAGNOSIS: Rostrum triangular, short and variable in length, not reaching further than middle of visible part of first antennular article.

Rostral carina low, rounded, extending past posterior end of orbital hoods; orbitorostral grooves shallow, at times non-existent.

Second antennular article almost twice as long as broad and 1.5 times as THE ALPHEIO SHRIMP OF AUSTRALIA — 247 scaphocerite almost straight; lateral tooth slightly exceeding third article, varying from slightly longer than, to well past squame.

Carpocerite thick, exceeding length of antennular peduncle by length of third article.

Large chela 2.6 times as long as broad, fingers occupying the distal third.

Superior saddle “U”-shaped with distal shoulder gently rounded and proximal shoulder varying from gently rounded to forming a right angle to margin of palm.

Medial palmar depression small and “Uff-shaped, lateral palmar depression triangular with apex reaching linea impressa.

Inferior shoulder low and rounded, not projecting; inferior notch small, extending as faint depression into both lateral and medial faces.

Plunger of dactylus well developed.

Merus 2 times as long as broad, inferointernal margin rounded, bearing 2-3 short spines.

Small chela of male 3.6 times as long as broad, fingers slightly shorter than palm with medial tooth flanking dactylar articulation.

In most males the dactylus is balaeniceps-shaped with characteristic rows of setae on lateral and medial faces that meet on dorsal surface.

In other males these crests of hair are imperfectly developed.

Pollex also bearing a slight fringe of hairs proximally.

The female chela is more slender sometimes bearing a slight fringe of setae on dactylus (fig. 77k) and in others (fig. 77 n,o) the fringe of setae is lacking and the dactylus bears only random patches of hairs.

Merus similar to that of large chela.

Carpus of second leg with ratio of articles: 10:7:3:3:5.

Third leg usually without spine on ischium.

Merus unarmed and distally rounded, varying from 3.5-4.5 times as long as broad.

Carpus 0.45 as long as merus.

Propodus 0.6 as long as merus and bearing on its inferior ma·rgin about 7 spines and a pair distally.

Superodistal margin bearing either a few long setae or one or two spines.

Dactylus simple, occasionally bearing on ventral surface a slight thickening suggesting a secondary unguis.

Telson 2.3 times as long as broad posterioriy, posterior border slightly arcuate.


Edwardsii leviusculus was applied by Dana to a specimen collected at Wake Island in the Pacific.


Bouvieri was applied by Milne-Edwards to specimens collected from the islands of Cape Verde Archipelago in the central Atlantic.

Bate in 1888 applied the name A.

Leviusculus to a specimen from the Challenger Expedition that all future workers decided was not similar to that described by Dana.

Coutiere dealt with the complex twice.

First (1898b:133) he reported the extension of the range of A.

Bouvieri to the west coast of Africa, to Fernando de Noronha in the Western Atlantic and to the Pacific coast of Panama.

In the same paper he also described a new variety, A.

Bouvieri bastardi, from Madagascar, Djibouti and Panama (presumably also in the Pacific coast).

In his second paper (1898h :249) he stated thatA.

Edwardsii of Dana was not that of Audouin, butwasA.

Bouvieri Milne-Edwards, and that Dana’sA.


Leviusculus was merely an A.

Bouvieri with an anomalous chela.

In spite of his rejection of A.

Leviusculus in this paper, Coutiere recognized A.

Leviusculus as well as A.

Edwardsii and A.


Bastardi as occurring at Djibouti (1899:486).

In this listing it should be noted that Lockington in 1878 (p. 474) created a homonym (or near homonym) by naming A.

Laeviusculus from California; according to Coutiere (1909:21) this species was actually in the genus Synalpheus and to it he applied the name s.


Subsequent references to the species complex were rather rare: Coutiere (1905a: 907) said that he could find no differences between A.

Bouvieri from the Maldives and the Milne-Edward’s type from the “Canaries” (sic), and he describedA.

Bastardi more fully from the Maldives, and raised it to species level.

In the same paper he pescribed A. 248 DORA M.



Hululensis as a new variety and stated (p. 915) “CA.

Leviusculus Danaest probablement, comme le dit cet auteur, une simple varilHe de fA.

Bouvieri . .. ” De Man recognized one specimen from the Siboga Expedition asA.

Leviusculus (raising it also to the species level) and suggested that while it was definitely separate from A.

Bouvieri, A.

Bastardi might be a synonym.

Coutiere (1921 :427) listed both A.

Bouvieri andA.

Bastardi as being collected by the Percy Sladen Trust Expedition in the Indian Ocean.

Edmondson (1925:15) lists A.

Leviusculus as being collected from Wake, Hult (1938:3) from the Galapagos, and Barnard (1950:740) from Mauritius.


Bouvieri was listed by Holthuis in the Atlantide Reports (1951 :81), but the name of this form was subsequently changed toA.

Holthuisi by Ribiero (1964:1).

Forest and Guinot (1958:9) gave a partial redescription of Milne Edward’s type series.

Finally, to conclude this period, Holthuis (1958:28) reported A.

Leviusculus from the Red Sea with the remarks: “It is doubtful whether the Indo-West Pacific specimens assigned by Coutiere to A.

Bouvieri actually can be distinguished from A.

Leviusculus” and suggested that the separation between the three species being here discussed should be revised.

I n the late 1950’s as we lauQched ou r studies of the Pacific alpheids we discovered we had a collection of 45 reasonably intact specimens that we considered to be A.

Leviusculus, including 5 specimens from Wake Island, the type locality, and 5 specimens in better condition from Johnston Island already reported upon by Edmondson (1925 :15).

To clarify the definition of the species we selected a 9.8 mm male to be designated a neotype and redescribed the species, and then on the basis of the other specimens at hand we made a study of the extent of variation in the series of specimens.

This entire series of specimens and some of our original notes upon them were destroyed in a laboratory fire late in 1961 (1962:238) so we could not designate a neotype.

Fortunately the complete manuscript which gave the description and the discussion had been prepared and was lying unpublished at the Bishop Museum in Honolului it was subsequently published (B&B, 1964).

We should note in passing that we failed to report the collection localities from the specimens other than the 13 reported in the 1964 paper.

In going over the fire-ravaged and water-stained original notes we discover that some, if not most of the 32 specimens came from the Marshalllslands, principally Bikini and Enewetak (old spelling, Eniwetok) and at least one collection from Western Samoai some may have been collected in other archipelagoes.

In the 1964 paper we compared the variation in A.

Leviusculus to the reported differences between the AltanticA.

Bouvieri and the Indian Ocean A.

Bastardi, and found that we could not distinguish between the nominal forms.

We therefore placed the two younger names in synonymYi we reserved judgment on A.

Bouvieri hululensis.

In 1966 we recorded and described the species from Thailand but did not discuss it further.

Also in 1964 Crosnier and Forest published their preliminary notes on the collections of the Calypso made in tropical eastern Atlantic, and stated that the specimens reported under the name of A.

Bouvieri from the Indo-Pacific were probably a different species.

In their final report on the collections they gave excellent descriptions and figures of A.

Bouvieri and A.

Bouvieri hululensis (1966:273, 282 et seq.)i they also gave a table separati ng seven species and forms they considered to be closely related, but they did not mention A.


They gave 3 criteria that would separate A.

Bastardi from A.

Bouvieri: A.

Bastardi has a shorter rostrum with a more rounded and shorter carina, a broader and more rounded squame on the scaphocerite and finally A.

Bastardi has spines on the ischia of the third and fourth legs which are lacking in A.


They suggested that Coutiere’s report of A.

Bastardi from Panama should be confirmed. THE ALPHEID SHRIMP OF AUSTRALIA 249 Chace (1972:63) recorded 26 specimens of A.

Bouvieri from various parts of the Caribbean and gave certain differences he found between ou r description and the range of variation we had given for A.

Leviusculus and the Caribbean specimens.

These were: large cheliped, merus palm, length/breadth Small cheliped, male Tooth of dactylar articulation Second legs, carpus, ratio of 2nd to 5th article Third legs, spine on ischium A.

Leviusculus small tooth 1.90 rarely balaeniceps sharp second shorter A.

Bouvieri rounded 1.48-1.80; average 1.66 always balaeniceps subrectagular or bluntly acute.

Second 0.98 to 1.6 times fifth present absent Chace also discussed some specimens from the Galapagos and Clipperton Island which he felt lay between A.

Leviusculus and A.

Bouvieri, and concluded it seems best for the time being to treat all forms (including the Eastern Pacific forms) as separate species.” H ••• We wish again to discuss these separations.

We have some notes preserved of our original study on the 45 specimens, the 16 specimens listed above, approximately 10 specimens from Madagascar (some fragmentary) and one each from the Philippines, the Maldives and Hong Kong.

We also examined specimens from the West Indies at the Smithsonian Institution.

ROSTRUM: Again we have found in the Australian specimens variation that encompasses the supposed differences set forth, with some specimens from the Coral Sea (AM 64 and AM 226) corresponding to the condition reported for A.

Bastardi, while some of the specimens from Heron Island (BAU 54) are similar to Coutiere’s figure for A.


Most of the specimens again were intermediate.

SCAPHOCERITE: The variation in the breadth of the squamous portion and the curvature and length of the lateral spine in the Australian specimens again was like that reported in 1961, and encompasses the differences cited by other authors.

LARGE CHELIPED: In our original tabulation of the 45 specimens we measured only the total length-breadth ratio of the chela, not the length-breadth ratio of the pal m, but ou r figures give an indication of the variation: of 36 specimens with the chelae intact, the average length-breadth ratio was 2.63, but the range was 2.1-3.0.

In the figure we drew of the “neotype-to-be” the length-breadth ratio of the entire hand was 2.76 and that of the palm 1.88.

If we are to presume that the other 35 specimens had the same relative finger lengths (which, of course, they would not have), then the average palmar ratio would have been 1.78 and the range would have been 1.43-2.05.

This average is within 0.12 of Chace’s average and the range extends on either side beyond Chace’s range.

Similar variation was found in the Australian specimens, but they were not measured.

The small tooth on the merus usually was present in the central Pacific and Australian specimens, but it varied in size and at times was lacking; in one pair, probably cohabiting (AM 64), the end of the merus in the female was rounded while it had an acute tooth in the male.

SMALL CHELA: The male Australian specimens showed a greater range of variation in the degree of the balaeniceps condition than did those originally studied.

For example, 2 250 DORA M.


BANNER males from the Coral Sea had a definite balaeniceps-type fringe of setae, but did not have the great lateral expansion found in other specimens, while none of the 5 males with small chelipeds present collected from Heron Island bore more than short setiferous crests as described for the Wake Island specimens.

In our notes about the original 45 specimens we wrote “spine at (dactylar) articulations somewhat blunt, sometimes sharp, always present”.

We are not sure whether this means any were “sub rectangular”, but it does overlap Dr Chace’s “bluntly acute”.

CARPUS, SECOND LEGS: We did not measure the relative lengths of the second and fifth carpal articles in our initial study, but we did measure the relative lengths of the first two articles in 36 specimens with these legs intact.

Here the ratio was 10:4.4.

To 10:6.3 with the average of 10:5.7.

It is worthy to note that Crosnier and Forest stated that they fou nd the fi rst article to be 1.65-2.5 as long as the second in A.

Bouvieri, which if converted to the ratio we are usillg, it would be 10:4.0-10:6.1. ,We measured the second to fifth articles of 5 Australian specimens taken at random and found the second articles to be equal to the fifth in 2 specimens and 1.06, 1.15 and 1.19 times as long in the other three – thus the Australian specimens have a longer second article than the one drawn from Wake.

In six specimens from the Caribbean studied atthe Smithsonian, 3 had the second and fifth articles equal, two had the ratio of 1.1 and one had an anomalous 1.46.

A re-examination of the original data showed another interesting thing: 6 specimens collected in Western Samoa (BBS6) all had markedly shorter second articles when compared to the first: their ratios ran from 10:4.4-10:5.8, with an average of 10:4.8, and if these were removed from the 36 previously averaged, the average would be 10:5.9.

This may indicate a geographically distinct race but we would be loathe to consider it of taxonomic importance.

ISCH I UM OF TH IRD LEGS: Both Crosnier and Forest and Chace regard the presence of a spine on the ischium inA.

Leviusculus or A.

Bastardi, and its absence inA.

Bouvieri as important; indeed, Chace stated that the characteristic is “perhaps of most importance”.

Unfortunately, we did not consider this in our original study and we now do not know how consistently the specimens in the Central Pacific were in the characteristic.

The specimens we drew from Thailand lacked the spine.

In 13 randomly selected specimens from Australia, 5 carried ischial spines, 8 did not.

Thus again we find that there are no firm characteristics to differentiate the specimens from the Atlantic and from the Indo-Pacific.

Yet it is inconceivable that in recent geological time this species, like other circumtropical species of non-pelagic marine life could have been able to maintain a common gene pool.

One of us (DMB) after examining the specimens from the Caribbean at the Smithsonian, was struck with subtle differences between these and the familar Indo-Pacific specimens and wrote in her working notes “Think FC (Fenner Chace) is right about the species from the Caribbean.” Thus, we conclude thatA.

Leviusculus should be divided into at least two geographically separate subspecies, A.

Leviusculus leviusculus and A.

Leviusculus bouvieri which probably can be distinguished by differing norms in distribution curves of variable characteristics, but not by any single firm difference.

In 1964 we renamed as A.

Batesi the specimen from the Philippines that Bate had listed in the Challenger Report as A.

Leviusculus; it evidently has not been collected since that time.

In the same paper we suggested thatA.

Bouvieri hululensis might be a species separate from A.


Since that time, through the courtesy of the Museum National d’Historie Naturelle of Paris, we were able to examine Coutiere’s holotype from THE ALPHEID SHRIMP OF AUSTRALIA 251 lmm ‘———“c 2mm ‘–_’—–‘, b — DORA M.


BANNER 27893); 5, AM P. 3127; 2, AM P. 5215; 4,AM P. 8009; 3AM p, 13561; 2,AM P. 13569; 1,AM P. 27452; 13, BAU 2; 12, BAU 8; 19, BAU 9; 1, BAU 23; 1, BAU 25; 7, BAU,26; 34, BAU 34; 2, BAU 36; 8, BAU 41; 13, BAU 45; 4, BAU 46; 8, BAU 59; 3, BAU 72; 5, BAU 73; 4, BAU 74; 13, BAU75; 6,JB1; 2,JC1; 6,JC2; 2,JC9; 2,JC13; 1,JC19; 2,JC21; 1,JC26; 3,JC27; 9,JG 6-73; 1, jG 7-73; 3, jG 10-73; 1, JG 17-73; 4, MM 72; 2, QM W1224; 3, QM W2234; 4, QM W 2240; 15, QM W 2391; 1, UQ 21; 3, UQ 23; 1, UQ 27; 1, US 106163; 1, US 106166; 2, US 123567; 6, US 123568; 1, US 123602; 20, US 123603; 1, WM 36-65; 1, WM 142-65; 7, WM 275-65.

DIAGNOSIS: Rostrum acute, triangular, varying from 1.1 to 1.7 times as long as broad reaching almost to end of first antennular article.

Orbitorostral grooves shallow and rounded.

Second antennular article usually about 2 times as long as broad and varying from 1.3-2.0 times length of first; third varying from 0.6 to equal length of first.

Stylocerite acute, reaching to end of first antennular article.

Scaphocerite with lateral tooth reaching just beyond antennular peduncle; squamous portion reaching end of antennular peduncle.

Tip of carpocerite reaching to end of lateral tooth of scaphocerite.

Large chela similar to A.

Australiensis sp.

Novo (see below p. 256).

Small chela sexually dimorphic.

Male chela balaeniceps, varying from 3.1 to 4.7 times as long as broad.

In fully mature specimens the palm usually bears sculpturing similar to that of large chela but reduced; in smaller males sculpturing is greatly reduced and may be almost entirely absent.

Female chela not balaeniceps, varying from 3.5-4.7 times as long as broad.

Sculpturing on palm varying with maturity of female, with larger specimens bearing superior indentation and inferior shoulder strong but less developed than in large males while in smaller females all sculpturing may be lacking.

Ratio of carpal articles of second legs varying as indicated: 10: (6-8): (3-4): (3-4): (4-5).

Ischium of third leg usually with movable spine.

Merus inermous, varying from 3.5-5.0 times as long as broad.

Distal margins of carpus not produced into acute processes.

Propodus usually with about 10 spines.

Dactylus simple, slightly curved.

Telson 2.3 times as long as posterior margin is wide, spines on upper surface small.

DISCUSSION: After examining topotypes of A.

Lobidens we foundA.

Crassimanus to be a junior synonym.

Then, examining collections of A.

Lobidens from the Indo-Pacific area, including Australia, we separated the species into two geographic subspecies: A.


Lobidens and A.


Polynesica (1974 :429).

The pri ncipal difference between the two rested in the sculpturing of the small chela of the mature males.

In A.

Lobidens the superior margin of the palm proximal to the dactylar articulation is notched similar to that of the large chela and opposite this on the inferior margin is a strong shoulder with the groove extending into the medial face.

In A.


Polynesica these grooves are only slight constrictions in the outlines: However, the smaller males of A.


Lobidens are similar in sculpturing to A.





Lobi dens occurs in the western Pacific and Indian Ocean, A.


Polynesica has only been collected in the central Pacific.

This same type of variability was described by us for specimens of A.


Euphrosyne from Thailand (B&B, 1966b:130).

In this paper we are describing a new species, A.

Australiensis (p. 256), as closely related to A.


Lobidens and we are questioning the separation of A.

Inopinatus Holthuis and Gottlieb from A.


Lobidens (p. 241).

In a group of 11 specimens from near Brisbane, Qld. (AM 350) the inferointernal margin of the merus of the large cheliped was inermous distally.

However, as they resembled in every other character A.


Lobidens, we interpret this as a variation. \ THE ALPHEID SHRIMP OF AUSTRALIA 255 Tiwari (1963:307) described 2 males as A.

Crassimanus which we believe are two different species.

The “short male” he described exactly agrees with A.

Sudara, a species we described from Thailand (1966b:153, fig. 59; see also p.372 above).

In his plates, figures 25a, d and 26b, c, d, e are figures of A.


The “long male” he describes is probably the trueA.


Lobidens and is represented in his plates by figu res 25b, 25c and 26f.

Fourmanoir (1958:118, fig. 5) records on some specimens as A.

Crassimanus from Nosy Be Madagascar.

He figures the large chela with the distal margin of the grooves on both the superior and inferior margins as being projected and subacute, a condition that never occurs in A.


We feel these specimens were, in all probability, Alpheus edwardsii (Audouin).

BIOLOGICAL NOTES: Barnard (1r.J50:758) remarks on the colour of his Irve specimens, “Greeny-brown, olive green, or smoky-grey, anterior parts of abdominal segments often white (producing a banded appearance), with or without longitudinal stripes (a median and 2 lateral) on each segment, the lower lateral stripe runs along the lower margins of the pleura and is often edged with black, a black spot in middle of the side on segments 2 and 4; telson and uropods apically blackish; chelae greeny-orange or greeny-brown, finger and thumb of large chela orange, tips dull violet, palm with a more or less brilliant cobalt-blue patch on inner (upper) surface; other legs dull pinkish” Kemp (1915:301) also remarked on the black spots on each side of the second and fourth abdominal segments.

Kemp (loc.


P.300) found in his specimens from Chilka Lake thatthe species has an adaptability to extreme salinity changes.

He found that although the species lives under rocks it does not produce an elaborate burrow, but simply makes a horizontal tunnel not more than a few inches in length.

Farrow (1971 :482), on the other hand, reports that at Aldabra the burrows in the carbonate sand are elaborate with the main tunnel lying horizontal 8-13 cm below the sand and with several sets of dichotomously branched entrance burrows reaching to the surface.

He does not give the length of the horizontal portion of the burrow, but if one estimated from an approximatley 10 cm depth in his figure 17a, the burrow is near a half metre long.

McNae (1957:361) reports in South Africa they inhabit radiating burrows to the depth of 25 cm.

Farrow, McNae and others have reported that this species lives in associaton with gobiid fish (see also discussion on p. 182).

Almost all ou r ou r speci mens were collected i ntertidally, but we do have 4 speci mens that were captured in a prawn trawl at 14 fathoms in the Gulf of Carpentaria (AM 13), presumably they had been living in the mud on the bottom.

This species is also occasionally found in the bases of dead coral heads.

Our largest specimen attained the length of 44 mm but Barnard (1950:758) reports on specimens up to 55 mm in length.

AUSTRALIAN DISTRIBUTION: Our specimens have come from all warmer parts of Australia: Houtman Abrolhos, Darwin, Gulf of Carpentaria, Torres Straits and on down the east coast to Sydney, N.S.W.

We also have 17 specimens from Lord Howe Island.

GENERAL DISTRIBUTION: This species ranges throughout the entire Indo-Pacific area from the Red Sea to Hawaii, but the subspecies does not occur in the central Pacific area.

Forest and Guinot (1956:102) reported the species under A.

Crassimanus from Tunisia; it may have reached the Mediterranean via the Suez Canal. 256 DORA M.


BANNER Alpheus australiensis sp.

Novo Fig. 79 HOLOTYPE: 33 mm male from Caloundra, Qld., collected by A.


Livingstone 14/8/22. (AM P. 6352).

ALLOTYPE: 26 mm female from the same locality as the type. (AM P. 27264).

PARATYPES: 1 specimen from AM 46 (AM P. 27205); 11, AM 126 (AM P. 27221); 2, AM 144 (AM P. 27220); 1, AM 150 (AM P. 27202); 4, AM 167 (AM P. 27222); 3, AM +16 (AM P. 27203); 1, AM 349 (AM P. 27201); 10, AM 350 (AM P. 27248); 2, AM 406 (AM P. 27204); 1, AM P. 10980; 1, AM P. 27254; 13, AM P. 27255; 16, BAU 63.

DESCRIPTION: Rostrum conical, about as long as wide at base, reaching somewhat past middle of visible part of first antennular article.

Rostral carina rounded, extending posteriorly to base of eyes.

Orbital hoods not markedly inflated with frontal margi n somewhat convex; orbitorostral grooves moderate.

Ratio of antennular articles beginning with visible part of first antennular article 10:13:10; second antennular article 1.4 times as long as broad.

Stylocerite acute, reaching to end of first antennular article.

Squamous portion of scaphocerite moderately wide, reaching end of antennular peduncle; lateral tooth a little longer, outer margin straight.

Carpocerite reaching length of third antennular article past that article.

Basicerite with acute lateral tooth.

Ratio of articles of third maxilliped: 10:4:6.

Second article bearing only long hairs on inner face.

Large chela 2.4 times as long as broad, fingers occupying the distal 0.4.

Superior saddle broad and relatively shallow, proximal shoulder usually gently rounded but at times almost abrupt, distal shoulder always gently rounded.

Medial palmar depression a well-marked triangle whose apex reaches half the distance from saddle to proximal end of palm.

Lateral palmar depression quadrangular, reaching to linea impressa.

Inferior shoulder heavy and rounded; inferior notch broadly “U”-shaped, continuing on lateral face of palm as a well-defined but small triangular depression with rounded apex, and on medial face as a longer, broader, but less well-defined depression.

Plunger pronounced.

Merus a little longer than broad, bearing no teeth distally on inferointernal margin.

Small chela of male 3.0 times as long as broad, dactylus balaeniceps, fingers only slightly shorter than palm, both superior and inferior margins of palm with shallow, rounded indentations proximal to fingers.

Medial side of dactylar articulation bearing acute tooth.

Superior margin of dactylus with a slight subacute carina that disappears where the crest of hairs meet on the superior surface.

Merus 1.7 times as long as broad with distal margins inermous.

Small chela of female similar to male, 3.4 times as long as broad with fringe of setae well developed on medial and lateral margins of dactylus, but not reaching beyond two-thi rds length of dactylus and not meeti ng at crest.

Merus similar to male but 2.3 times as long as broad.

Ratio of carpal articles of second leg: 10:9:3:4:4.

Ischium of third leg armed with spine.

Merus inermous 4.3 times as long as broad.

Carpus 0.5 as long as merus; superior and inferior margins slightly projected distally.

Propodus 0.7 as long as merus bearing 7 spines on inferior margin and a pair distally.

Dactylus simple, slightly curved, 0.3 as long as propodus.

Telson 2.5 times as long as posterior margin is broad.

Dorsal spines of moderate size.

DISCUSSION: It is with considerable reluctance that we are naming this as a species separate from A.

Lobidens lobidens De Haan and A.

Lobidens polynesica Banner and THE AlPHEID SHRIMP OF AUSTRALIA 257 Banner (1974:429).

We separated the nominate species from the subspecies on the basis of the small chelae of mature males which carry heavy sculpturing in the nominate species that is entirely lacking in the subspecies.

The two subspecies are geographically separated, with the nominate form being found in the far western Pacific (including Australia) to the Red Sea and A.


Polynesica being confined to the archipelagoes of the central Pacific.

The central Pacific subspecies never reaches the large size at maturity attained by the western subspecies. A.

Australiensis can be firmly separated from the two subspecies of A.

Lobidens only by the characteristics of the small chelae of the males and females.

Like A.


Polynesica, the males of mature size lack the sculpturing on the small chela, and unlike both subspecie·s the chelae of the females show balaeniceps development.

These and other characteristics of less reliability are set fourth in Table 6. — ‘—__-‘-__-‘, a 2mm Fig. 91 Synalpheus tijou sp.

Novo Holotype (female).

A, b.

Anterior region, dorsal and lateral view; c, d.

Large chela and merus, lateral face; e.

Small cheliped, medial face; f.

Second leg; g, h.

Third leg and dactylus, enlarged; t telson and uropods.

A, b, c, d, e, f, g, i scale a; h, scale b. 298 DORA M.


BANNER second as broad as long and equal in length to third.

Stylocerite minimal in size, reaching only 0.5 length of first antennular article.

Statoliths plainly visible beneath the stylocerite.

Squamous portion of scaphocerite broad, reaching near end of antennular peduncle, lateral tooth acute but markedly shorter than squame.

Carpocerite 5 times as long as broad when viewed laterally, not reaching end of antennular peduncles.

Basicerite without teeth.

Ratio of articles of third maxilliped: 10:2.4:5.1.

Third article with a few heavy setae distally. ~arge chela 3.2 times as long as broad, surface wrinkled from dessication, but normal chela is undoubtedly without grooves and may be thicker.

No tooth on palm flanking dactylar articulation.

Fingers 0.3 as long as entire chela.

Plunger well developed.

Merus 2 times as long as broad and without inferior and distal teeth.

Small chela 5 times as long as broad, palm 2 times as long as fingers.

Dactylus not broadened, tips of fingers acute, somewhat hooked and crossing when closed.

Merus unarmed, slender, 6times as long as broad in middle. Second legs with only 4 carpal articles.

Ratio of carpal articles: 10:3:3:5.

Ischium of third legs inermous, almost half as long as merus.

Merus also inermous, 5 times as long as broad.

Carpus 0.48 as long as merus, distal angles rounded and not projecting.

Propodus as long as merus, bearing on its inferior margin 6 strong spines and a pair distally.

Dactylus biunguiculate, 0.14 as long as propodus, tip curved at right angles to axis of propodus; inferior unguis arising in basal third of dactylus.

Superior unguis 4.5 times as wide at the base as the inferior unguis and 4 times as long.

Telson 4.6 times as long as broad posteriorly and almost 4 times as broad anteriorly as posteriorly; posterior margin convex.

No spines visible on superior surface; posterolateral spines small.

Outer uropod with a distinct articulation.

DISCUSSION: The lack of oribitorostral process, the anterior region of the carapace advanced so far distal to the eyes, the markedly curved superior unguis of the dactylus of the thoracic legs and its association with a crinoid places this species firmly within the Comatularum Group of the genus Synalpheus. “From all species within the group it differs by the extreme length of its rostrum and the great reduction of the orbital teeth; by the slender and elongate palm of the small chela; by the presence of 4 articles in the carpus of the second legs (if this is an adult condition); by the form of the dactylus of the third legs; and by the shape of the telson with its extreme taper and narrow tip.

It may be separated from other species by other specific characteristics, as by the lack of a tooth on the base of the propodal finger of the large chela as found in S.

Odontophorus de Man, or the lack of a hooked daqylus of the small chela as in S.comatularum (Haswell), but the major differences listed above are more than sufficient.

It appears to us that the species may be related most closely in the group to S.

Stimpsonii (de Man) but that species always has strong and distinct orbital teeth.

This specimen was evidently ready to moult or had recently moulted for the integument was soft and transparent, permitting one to see through it in places.

Visible were slightly more pronounced orbital teeth and possibly one pair of dorsal spi nes on the telson.

The soft exoskeleton had also caused the palms of both chelae to wrinkle, so that they may be more plump in a fully hardened specimen.

The presence of four articles in the carpus of the second legs may be an indication of immaturity in the specimen (see discussion in B&B, 1975:298), but in no other members of the genus does one find immature traits in a 14 mm long specimen. THE ALPHEID SHRIMP OF AUSTRALIA 299 This species may be inserted into the key to Synalpheus by deleting the name s.

Stimpsonii from dichotomy 5 on p.279 (1975) and inserting the following: 5A (5) Rostrum not reaching near end of second antennular article; orbital teeth prominent and acute; carpus of second legs of 5 articles …


Stimpsonii If:’ 292 Rostrum reaching near end of third antennular article; orbital teeth reduced to low obtuse angles; carpus of second legs of 4 articles .. 5.

Tijou This species may also be inserted in the key to the alpheids known to inhabit crinoids (1975:389) by converting the dichotomy under 6 to a trichotomy with the addtiion of: Rostral carina absent; tip reaching to near end of third article …… .5.

Tijou The name is derived from the type locality.

The holotype will be placed in the Australian Museum, Sydney N.S.W.

Synalpheus paralaticeps sp.

Novo Fig. 92 HOLOTYPE: 10 mm female from Rudder Reef, off Port Douglas, Qld.

From outer reef flat, 200 yards from reef edge.

Reef subject to heavy surf with S.


Winds. (BAU 30) (AM P.30810).

ALLOTYPE: 9 mm male from same collection. (AM P.30811).

DIAGNOSIS: Rostrum slender, 1.8 times as long as broad, reaching to last quarter of visible part of first antennular article, tip rounded.

Orbital teeth as long as broad at base and almost as long as rostrum.

Tip of rostrum and orbital teeth bearing a few short stiff setae.

Rostral base with orbitorostral process.

Second antennular article slightly longer than wide, 0.8 as long as visible part of first, and 1.5 times longer than third.

Styl”ocerite reaching near middle of second antennular article.

Scaphocerite with very narrow squame reaching to end of second antennular article, lateral tooth reaching nearly length of third article past that article.

Carpocerite 5.6 times as long as broad, viewed laterally, and reaching well past end of lateral tooth of scaphocerite.

Inferior tooth of basicerite reaching near end of second antennular article and superior tooth as long as orbital teeth.

Ratio of the articles of third maxilliped 10:1 :6.

Distal tip beset with a circlet of short spines.

Large chela cylindrical, 2.6 times as long as broad, with fingers occupying distal 0.3.

Tip of dactylus abruptly rounded.

Palm bearing one obtuse tubercle above dactylar articulation.

Merus twice as long as broad, bearing a small acute tooth on superodistal margin.

Small chela 2.8 times as long as broad with fingers 0.4 as long as palm.

Dactylus bears 2 rounded curved teeth distally; pollex terminating in a single acute tooth flanked by two abrupt shoulders lying at right angles to axis of tooth.

Merus 2.8 times as long as wide, unarmed.

Carpus cup-shaped, 0.3 as long as chela.

Carpal articles of second legs with a ratio: 10:2:2:2:5; middle articles broader than long.

Ischium of third leg over half length of merus, unarmed.

Merus 3 times as long as broad and bearing two small spines on inferior margin in distal third with a few hairs proXimally.

Carpus 0.4 as long as merus with superodistal margin bearing obtuse projection and inferodistal margin a small spine.

Propodus 0.7 as long as merus, inferior margin with 6 pairs of spines interspersed with a few hairs.

Dactylus biunguiculate, inferior unguis a little less than half as long as superior; apex of notch subacute.

Telson 3 times as long as tip is broad, 2.2 times as wide anteriorly as at tip.

Anterior 300 DORA M.


BANNER O.5mm 2mm lmm — 2mm lmm ‘——–” b Fig. 93 Athanas polynesia B&B 8 mm male from AM P. 25161.

A, b.

Anterior region, lateral and dorsal view; e, d.

Large cheliped, lateral and medial face; e.

Small cheliped, lateral face; f.

Second leg; g, h.

Third leg and enlarged dactylus; i.

Telson and uropod. 8 mm female from 75·LlZ-V.


Anterior region, dorsal view; k.

Cheliped, lateral face.

A, b, c, d, e, f, g, i scale a; h, j, k scale b. 304 DORA M.


BANNER Telson 3.7 times as long as posterior margin is broad.

Posterior margin arcuate, anterior pair of dorsal spines placed anterior to middle.

Uropods elongate with outer uropod distally bearing articulation.

Sympodite with two teeth, outer longer and broader than inner.

DISCUSSION: Unfortunately, most of these specimens are fragmentary.

The seven specimens with the “LlZ” collection records are especially bad, with only 4 walking legs and one cheliped remaining attached; the male specimen from AM P. 25161, also from Lizard Island, is however complete.

Extra appendages in the “LIT’ bottles agreed with the appendages that were attached.

There was only one female cheliped in the entire collection and females of this species have not been previously described.

We suggest that the females probably have symmetrical chelae as has been reported in related species.

The rostrum in one specimen is short and somewhat distorted (fig. 92j); we suggest that it may be a malformation from heredity or accident, as we reported in a specimen of A.

Borradailei (Coutiere) from Samoa (1966a:152).

Otherwise in the intact parts only minor variation was noted, such as the range in the relative lengths of the carpal articles reported above.

The specimens agree well with the specimens we described from Samoa; the similarities in the male chelae are especially striking.

There is a group offour nominal species that lie intermediate between the old genera of Athanas and Arete; they are A.

Borradailei (Coutiere), A.

Ghardaqensis (Ramadan), A.

Verrucosus B&B, and this species.

All carry the chela in an inverted position, with the dactylus in inferior orientation; in all males the palm is greatly widened and compressed, with associated modification of the merus and carpus to permit flexion; in all the bases of the antennules and antennae are heavy and in all of these species the mouthparts are greatly protrudant (the mouth parts for A.

Ghardaqensis have not been described, but we have specimens of it as yet unreported in our collections from Madagascar and the Red Sea).

The only species that can be firmly separted from the others isA.

Verrucosus, for it has 5 carpal articles in the second leg rather than four.

The other species, described on the basis of one or few specimens, often partially fragmentary, are separated by more subtle and variable characteristics and should be reviewed when a larger series of intact specimens are available (as we pointed out in 1960:149 and in 1966a:152).

This compact group of species also should be considered in their generic relationship.

In 1960 we considered them as intermediate between the previously separated genera of Athana$ and Arete, so we combined the two genera.

On the other hand, it may be more useful to consider them to be a separate genus, yet to be deSignated, which would permit the two previous genera to stand.

With the present collections, however, nothing further can be done.

This species may be inserted into our key to the genus Athanas (B&B, 1973:303) by substituting the following for dichotomy 10: “10 (1) With supracorneal spines; dactylus of third to fifth legs simple …………………………………………………………………………. .A.

Polynesia Without supracorneal spines; dactylus of third to fifth legs biunguiculate ……………………………………………………………………..11″ and renumbering the present dichotomy from 10 to 11.

BIOLOGICAL NOTES: All of these specimens were reported as coming from areas largely covered with encrusting coralline algae in water from 6-20 ft deep.

AUSTRALIAN DISTRIBUTION: These Lizard Island specimens are the only ones THE ALPHEID SHRIMP OF AUSTRALIA known from Australia.

GENERAL DISTRIBUTION: Samoa. Salmoneus sibogae (De Man)* Fig. 94 305 Jousseaumea sibogae De Man, 1910b:303; 1911 :158, fig. 9. SPECIMENS EXAMINED: 1,14 mm male and 1,10 mm ovigerous female from North Reef, Heron Island, Capricorn Group.




Bruce, 411/77.

DIAGNOSIS: Surface of carapace finely granulate, bearing a slight pubescence of short stiff setae.

Rostrum triangular, with somewhat concave margins, 1.3 times as long as broad at base, tip reaching to middle of third antennular article; carina slight extending prosteriorly from tip to level of middle of eyes.

Extracorneal teeth triangular, about one-fifth as long as rostrum, with inner margins parallel to medial plane of body.

Margin between extracorneal teeth and rostrum narrow but rounded.

Antennular peduncle stout, with second antennular article slightly broader than long; antennular articles nearly equal in length.

Stylocerite with acute tip reaching slightly beyond end of second antennular article.

Scaphocerite with squamous portion broad, reaching just past antennular peduncle; outer margin straight; lateral tooth as long as squamous portion.

Carpocerite stout, reaching just past end of second antennular article.

Lateral tooth of basicerite broad and acute, reaching to level of tips of extracorneal teeth.

Pterygostomial angle produced but rounded.

Large chela 3.0 times as long as broad, with fingers occupying distal 0.4, rotated about 90° with fingers opening laterally.

Palm somewhat quadrangular but rounded in section without excavations, with heavy proximal shoulder and notch at carpal articulation, distally constricting opposite dactylar articulation.

Fingers compressed, bearing 11 teeth that mesh exactly when closed; tips hooked and crossing.

Carpus bearing 2 strong teeth on lower (medial) side, otherwise projecting in a cyathiform manner.

Merus 8.3 times as long as broad in inferior view, unarmed; inferior face broadened, flattened to excavate, and curved to accommodate palm when carpal articles are flexed.

Ischium about 0.4 length of merus (in female specimen; that of male broken).

Small cheliped of minimal size, when all articles are extended it is about 0.9 length of large chela proper, not showing sexual dimorphism.

Ischium unarmed, 0.6 as long as merus.

Merus 6 times as long as broad; carpus 0.8 as long as merus, almost 3 times as broad distally as proximally.

Chela 0.7 as long as merus, with fingers a little shorter than palm.

Second leg with ratio of carpal articles: 10:5:3:2:5.

Ischium of third leg with 2 spines, merus 5.8 times as long as broad.

Carpus 0.8 as long as merus, not projected distally; propodus 0.7 as long as merus bearing on its inferior margin 5 spines and a pair distally.

Superior margin bearing a slender spine distally.

Dactylus simple, 0.3 as long as propodus, bearing on its dorsal surface near the tip a small notch from which protrudes 2 fine hairs.

Telson 4 times as long as posterior margin is broad; posterior notch narrow, V-shaped, with depth equal to 0.5 breadth of tip.

With usual posterolateral pairs of spines and long plumose setae arising in middle.

Inner part of spines as long as outer (in female, spines on male telson missing).

Anterior pair of dorsal spines set posterior to middle.

DISCUSSION: It is the 14 mm male that is the basis of this description unless *This name will be reduced to a synonym of s.

Serratidigitus(Coutiere, 1896b) in a forthcoming paper on the alpheids of the Red Sea and Gulf of Aden. 306 — Fig. 95 Australia (sinusoidal projection), showing faunal provinces and percentages ot endemism of alpheid and ogyridid shrimp.

The location of the boundaries is discussed in the text, as well as the distinction between the Solanderian and Banksian provinces; in the figures, the numerator is the total number of species reported for the province, the denominator is the total number of endemic species found in the province, and the quotient is the percentage of endemism. THE ALPHEID SHRIMP OF AUSTRALIA 323 Finally, in fig. 95 and Table 8, we have treated the subspecies found in Australia as equal to the species, have considered the unnamed form Alpheus sp., as a separate species, and were forced to leave out A.

Brevirostris (Olivier) because there is no indication as to where it was collected in half the perimeter of Australia (see p.226).

The two species of Ogyrides are also included in the computations.

As to be expected, the tropical fauna is best found in, but not confined to, the Dampierian, Solanderian and Banksian provinces.

Many species penetrate further south than the Tropic of Capricorn.

Along the east coast we find species especially of the Banksian province, which are species typically not associated with coral reefs, reaching as far south as Sydney, N.S.W.

The limits of the tropical fauna on the west coast are more difficult to delimit both by the lack of large numbers of collections, and by the lack of extensive coral reefs.

However, as we found that 7 tropical species had reached the Houtman Abrolhos slightly north of 29°S, these islands should be near the zone of transition.

This is further north than Griffin and Yaldwyn found for the majid crabs (op.

Cit., map 1).

The large number of species shared by the Solanderian, Banksian and Dampierian provinces (see Table 8) would indicate that if the same habitats were found in the three provinces, they could be expected to have much the same species composition.

The temperate fauna of the Peronian and Flinderian provinces have many fewer species (fig. 94) and like the tropical fauna, many of these species reach into the adjacent parts of other provinces.

For example, a number of species that appear to have arisen in the temperate fauna and are endemics (see below) reach up to Moreton Bay in the east and to Shark Bay in the west; two species, at least, reach from the south to the Gulf of Carpentaria. Fauna! Affinities and Endemism As Griffin and Yaldwyn found for all decapods, almost all species found in the Dampierian, Banksian and Solanderian provinces are of Indo-Pacific relationship, with 101 of the 142 species and subspecies tabulated appearing elsewhere in the Indo-Pacific tropics.

The group listed as “Indo-Pacific” included only those which were known to occur both in the Pacific north or east of Indonesia and in the Indian Ocean; the number of species in this group is likely to increase when more extensive collections are made in the western Pacific that are comparable to the collections made from the Maldives and Laccadives in the Indian Ocean.

The number of species shared only between Indonesia and Australia is what one would expect considering, first, that most zoogeographers have found the great shallow-water triangle between Indonesia and the southern Philippines to be the richest area in species in the entire Indo-Pacific with many species that apparently have not spread far from the triangle, and, second, considering that the Indonesian shelf area is bridged by a tight island chain to the shelf area between New Guinea and northern Australia.

Some of the non-Australian distributional patterns shown in Table 8 and listed in the text under the distribution of each species appear to be sharply discontinuous.

However, the discontinuities may be those of collection, not distribution.

For example, the species shown as shared alone between Japan and the Dampierian province isAthanas japonicus Kubo, a species that was collected intertidally under rocks on mud-flats in southern Japan and in a mangrove swamp near Darwin, N.T.

We are quite certain that if similar habitats were searched in the Philippines and Indonesia, the species would also be found there.

We also believe that the large number of apparent endemics in the tropical waters of Australia is also a reflection of the intensity of collection.

Most appear in the extensive Australian collections as a single or several specimens; presumably if collections as exhaustive were made elsewhere in the Indo-Pacific, these species might be found there Table 8.

Faunal Relationships of Australian Alpheids and Ogyridids.

Australian Distribution 0..

E ro 0..

E ro “0 w ~ ‘u il) Vl ~ il)

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    Horses-Store.com - 12) (7) Body highly compressed; carapace with knife-like mid-dorsal keel for its entire length