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Zoology Publications from Victoria University of Wellington—Nos. 78, 79 and 80

A Guide to the the Larvae of New Zealand Shallow Water Caridea (Crustacea, Decapoda, Natantia)

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A Guide to the Larvae of New Zealand Shallow Water Caridea
(Crustacea, Decapoda, Natantia)

Abstract

A guide to the identification of New Zealand's shallow water caridean larvae of the families Alpheidae, Crangonidae, Hippolytidae, Ogyrididae, and Palaemonidae is presented. It gives combinations of characters for identifying the larvae to the level of family, with brief descriptions or keys to genera, and to species where possible.

Introduction

New Zealand's shallow water1 caridean fauna comprises species belonging to five families: Alpheidae Bate, 1888, Crangonidae Bate, 1888, Hippolytidae Bate, 1888, Ogyrididae Holthuis, 1955 and Palaemonidae Samouelle, 1819. Larvae of many of these shallow water species have now been described, although much of this information is yet to be published (see Wear 1985). The guide presented here summarises this knowledge. The larvae of New Zealand's offshore Caridea are, however, too poorly known to enable construction of a single, comprehensive key to their larvae. This guide is intended as a preliminary step toward such a key.

The first part of the guide comprises a list of characters which, taken in combination, distinguish caridean larvae from those of other decapods. Larvae of each of the five families are then similarly characterised. Genera or individual species within each family are considered in the form of a key or, in the case of the Hippolytidae, are briefly described.

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Records of the New Zealand caridean fauna, and data on egg size and number, were obtained from Yaldwyn (1954, 1959, 1960, 1971), Richardson & Yaldwyn (1958) and Hayashi & Miyake (1970). The lists of characters distinguishing larvae of the Caridea and of each family were based principally on Gurney's (1942) "Larvae of Decapod Crustacea" and the keys of Williamson (1957, 1982).

1 Defined as 0-20m after Yaldwyn (1959)

Terminology

Terminology used in this guide essentially follows that of Gurney (1942). However, terminology used here to define developmental type and stage is given below.

Unabbreviated development

Species with four or more planktonic zoeal instars in their life history.

Key to zoeal stage
A. Eyes sessile zoea 1
Eyes stalked B
early stage
B. Uropods not freed from telson (Fig. 2F) zoea 2
Uropods freed from telson C
C. Uropod endopod rudimentary (Fig. 2E) zoea 3
Uropod endopod and exopod developed D
D. Pleopods absent or rudimentary (Fig. 2D) zoea 4+
late stage
Pleopods setose and functional postlarvae

Early and late stages are arbitrarily defined. The terms are useful when referring to larvae in which the number of instars is variable or unknown, or when more than one instar is referred to. Early larvae are those in which there are no pleopod buds, and usually refers to zoeae 1-3. Late larvae are those in which all the appendages are present and well formed, and the pleopod buds are large but not yet setose. Intermediate classification into mid-stage larvae is only necessary when page 3there are a large number of instars and the degree of morphological change between them is small. Mid-stage larvae are zoea 4+ larvae in which pleopod buds are absent or small.

Abbreviated development

Larvae with development abbreviated to three or fewer instars follow the above pattern in development of the eyestalks and uropods. However, all other appendages including pleopod buds are present from zoea 1 onwards. For example, a zoea 3 larva with abbreviated development has rudimentary uropod endopods but also has large pleopod buds and is similar in other respects to late larvae with unabbreviated development.

Caridea

Larvae of the Caridea can be distinguished from those of other decapods by the following combination of characters.

Caridean larval characters

1.Carapace and abdomen usually without large numbers of spines.
2.Posterolateral margins of carapace smooth, not produced posteriorly as spines and rarely overlapping more than abdominal somite 1.
3.Abdomen in mid- and late larvae with pleura of somite 2 overlapping those of somites 1 and 3.
4.Antennal exopod unsegmented (e.g. Fig. 2A) or in early larvae semented near distal end only (Fig. 3E).
5.Maxillipeds 1-3 present and with functional natatory exopod from hatching (except in some species with abbreviated development); endopod of maxilliped 1 shorter than that of maxilliped 2 which is shorter than that of maxilliped 3; basis of maxilliped 1 broad and flattened, with endopod at distal margin.
6.Telson dorsoventrally flattened; usually triangular in early larvae and parallel sided or tapering in mid- and late larvae; seta 2 not reduced to a small hair (Fig. 2E).

Family Alpheidae

The following species have been recorded from the New Zealand region: Alpheopsis garricki Yaldwyn, 1971, Alpheus nouvaezealandiae Miers, 1876, Alpheus richardsoni Yaldwyn, 1971, Alpheus socialis Heller, 1865 and Betaeopsis aequimanus (Dana, 1852). However, Yaldwyn (1959) noted that Alpheus novaezealandiae had not been found in New Zealand for 50 years. Early larvae of the other four species have been hatched and reared by the present author, and although not yet studied page 4in detail, it is possible to distinguish between them on the basis of the key provided below.

Larvae of the family Alpheidae can be distinguished by the following combination of characters.

Alpheid larval characters

1.Supraorbital spines when present without spinules (Fig. 1A).
2.Pereiopod 5 develops before pereiopods 3 and 4, terminal spine of dactylus extremely long (extends at least beyond eyes) and provided with spinules distally (Figs. IB, C).
3.Protopod of maxilla with 3 endites only, exopod shorter than endopod in early larvae (Fig. 1H).
4.Endopod of maxillule small and unsegmented.
5.Endopod of maxilliped 1 with two or three incomplete segments (as indicated by setation), never with indication of four segments (Figs. ID, E).
6.Maxillipeds 2-3 and pereiopods raptorial in form, with spiniform serrate and simple setae, dactylus produced into a strong spine (e.g. Figs. IF, G).

Key to larvae of the New Zealand Alpheidae1
A. Rostrum as long as or longer than antennular peduncle; with supraorbital spines from zoea 2; strong posterolateral spines on abdominal somite 5 from zoea 3; pereiopod 5 not developed until zoea 3 (Fig. 1A) Betaeopsis aequimanus
Rostrum short, does not extend beyond eyes; without supraorbital spines or posterolateral abdominal spines in zoeae 1-3; pereiopod 5 developed in zoea 2 (Fig. IB) B
B. Endopod of maxilliped 1 very small, with two incomplete segments (as indicated by setation) (Fig. ID) Alpheus richardsoni
Endopod of maxilliped 1 with three incomplete segments (as indicated by setation) Cpage 5
C. Ischium of maxilliped 3 without spiniform seta (Fig. IF) Alpheus socialis
Ischium of maxilliped 3 with spiniform seta (Fig. 1G) Alpheopsis garricki

1 Excludes Alpheus novaezealandiae

Family Crangonidae

The following species have been recorded from the New Zealand region: Pontocaris lacazei (Gourret, 1888), Pontophilus australis (Thomson, 1879), P. acutirostratus Yaldwyn, 1960, P. challengeri Ortmann, 1893, P. chiltoni Kemp, 1911, P. hamiltoni Yaldwyn, 1971, P. indicus de Man, 1918, P. pilosoides Stephenson, 1927, P. quadrispinosus Yaldwyn, 1971, P. yaldwyni Zarenkov, 1968, Prionocrangon curvicaulis Yaldwyn, 1960, Sclerocrangon knoxi Yaldwyn, 1960, S. richardsoni Yaldwyn, 1960. In addition, the Family Glyphocrangonidae may be represented in deep water off New Zealand (Richardson & Yaldwyn 1958), and larvae of species belonging to this family could possibly be confused with those of the Crangonidae on the basis of characters given below.

Early larvae of Pontocaris lacazei have been described by De Simón (1979), and larvae of Pontophilus australis, P. chiltoni, P. hamiltoni and P. pilosoides by Packer (1983). No other larvae of the New Zealand Crangonidae are known. However, on the basis of their relatively large egg size and on published information for larvae from outside the New Zealand region, the species of Prionocrangon and Sclerocrangon almost certainly have abbreviated larval development. This is probably also true for the Glyphocrangonidae.

Larvae of the Glyphocrangonidae and Sclerocrangon with abbreviated development, described from beyond the New Zealand region, have more than the usual 8+8 telsonic setae (e.g. Dobkin 1965, Makarov 1968). In larvae of Sclerocrangon the posterior margin of the telson is either smoothly rounded posteriorly (Fig. 2F) or weakly indented (Sars 1890, Wollebaek 1906, Makarov 1968) whereas in Glyphocrangon spinicauda the posterior margin of the telson bears a median spine or "prominence" (Dobkin 1965) (Fig. 2G). To the author's knowledge no larvae of Prionocrangon have been described.

All known larvae of the genus Pontophilus Leach can be separated into three distinct groups (Packer 1983). The first group includes larvae referred to the genus Pontophilus. The second group includes larvae referred to Philocheras Stebbing, which was synonymized with Pontophilus Leach by Kemp (1911) - a revision followed by the majority of authors with regard to the adults (Lebour 1954, Williamson 1982) but not accepted by authors working with larval material (see Williamson page 6(1960) for synopses of larval data and keys to the larvae forming these two groups). These groups have been referred to as 'Pontophilus-group A' and 'Pontophilus-group B' respectively (Packer 1983). Larvae of Pontophilus australis, P. chiltoni, P. hamiltoni and P. pilosoides, from the New Zealand region, form the third group. However, larvae of the five species from New Zealand waters which have yet to be described could belong to any of these three groups, and so provision for this possibility is made in the key given below.

Larvae of the family Crangonidae can be distinguished by the following combination of characters.

Crangonid larval characters

1.Eyes oviform, close together (e.g. Fig. 2A).
2.Carapace with ventral margins convex, without supraorbital spines (Fig. 2D).
3.Inner flagellum of antennule in the form of a stout rod (e.g. Fig. 2A) or of many segments (Fig. 2I).
4.Antennal exopod not distally segmented (except in early larvae of Pontocaris lacazei (Fig. 2I).
5.Exopod of maxilla without proximal extension (Fig. 2C, cf. Figs. 3K, 4I).
6.Pereiopod 1 with rudimentary subchela in late larvae, propodus broad (Fig. 2D).

Key to larvae of New Zealand Crangonidae
A. Larvae with all appendages present except the uropods, or uropods present but endopod rudimentary; in latter case pleopod buds large, i.e. abbreviated development Sclerocrangon Prionocrangon Fam. Glyphocrangonidae
Larvae with unabbreviated development B
B. Abdomen less than twice length of carapace, without dorsomedial spine on somite 3 (although this somite may have paired postero-lateral spines); posterior margin of telson weakly concave or straight Cpage 7
Abdomen more than twice length of carapace, with dorsomedial spine on somite 3; posterior margin of telson strongly concave D
C. Base of rostrum with concave lateral margins (Fig. 2A); exopods of maxillipeds 2-3 and pereiopods 1-2 with no more than six setae (Fig. 2D); posterior margin of telson straight from zoea 4 onward Pontophilus australis P. chiltoni P. hamiltoni P. pilosoides (other Pontophilus spp.?)
Base of rostrum with convex lateral margins (Fig. 2B); exopods of maxillipeds 2-3 and pereiopods 1-2 with more than six setae; posterior margin of telson weakly concave in ll larvae 'Pontophilus-group B'
D. Abdominal somite 5 with strong posterolateral spines; inner flagellum of antennule horter than carapace (Fig. 2H). 'Pontophilus-group A' (? Including P. indicus & P. challengeri from group 1 of Kemp 1916)
Abdominal somite 5 without posterolateral spines; inner flagellum of antennule longer than carapace, of many segments from zoea 2 onward (Fig. 2I) Pontocaris lacazei

Family Hippolytidae

Seven species have been recorded from the New Zealand region: Alope spinifrons (H. Milne-Edwards, 1837), Bathyhippolyte yaldwyni Hayashi & Miyake, 1970, Hippolysmata morelandi Yaldwyn, 1971, Hippolyte bifidirostris (Miers, 1876), H. multicolorata Yaldwyn, 1971, Nauticaris marionis Bate, 1888, Tozeuma novaezealandiae Borradaile, 1916. Larvae of five of these species have been described by Packer (1983). The remaining two species are Hippolysmata morelandi, a shallow water species from northern New Zealand, and Bathyhippolyte yaldwyni, which has been taken in deep water off the east cost of the South Island.

The eggs of B. yaldwyni are large and few in number, and so this species almost certainly has abbreviated development, and the larvae page 8should easily be distinguished from those of the other New Zealand Hippolytidae.

Larvae of Hippolysmata morelandi should also easily be identified. Known larvae of the genus Hippolysmata are characterised by the enormous size and oar-like propodus of pereiopod 5 (Fig. 3C). Gurney (1937) summarised larval characters of this genus.

Only the zoea 1 of Nauticaris marionis is known, and no other larvae of the genus have been described. Hence, without information on later larvae of this genus, it is not possible to construct a key to larvae of the New Zealand Hippolytidae. Therefore, in place of such a key, characteristic features of the known New Zealand hippolytid larvae are provided below.

There is such a wide variation of form among larvae of this family that it is difficult to provide a list of characters by which they can be recognised as a group. The following combination of characters is shared by the larvae of all New Zealand species except those of Hippolysmata morelandi for which characters 1, 3 and 7 do not apply:

Hippolytid larval characters

1.Eyestalks short, cylindrical rather than tapering proximally.
2.Rostrum extends beyond eyes (except in late larvae of Alope), without spines.
3.Maxillule with outer plumose seta and with distal lobe of endopod bearing three setae (Fig. 3J).
4.Exopod of maxilla in mid- and late larvae with proximal extension, and with distal margin almost straight rather than smoothly rounded, always with setae on outer proximal margin (Fig. 3K).
5.Maxillipeds 2 and 3 with flexible rather than spiniform setae (e.g. Fig. 3I, cf. Fig 4E).
6.Both pereiopods 1 and 2 with a well formed chela in late larvae.
7.Antennules not separated at base by more than width of one of them (Fig. 3D).
8.Antennal exopod distally segmented in early larvae (Fig. 3E).
9.Never more than 8+8 telsonic setae.

Diagnostic characters of New Zealand Hippolytidae larvae

Alope spinifrons (Fig. 3G)

Carapace with one pair of anterolateral spines in zoea 1, and two pairs from zoea 2 onward (of which the more dorsal is as long as the antennal protopod in late larvae), with supraorbital spines from zoea 2 page 9onwards; abdominal somites 1-5 without spines; pereiopods 1-4 with exopod bearing six setae when first developed but with up to 12 setae in late larvae; pereiopods 3-5 raptorial in form and pereiopod 5 shorter than pereiopods 2-4 when all developed.

Hippolyte bifidirostris and H. multicolorata (Figs. 3A, B)

Carapace with 3-5 pairs of anteroventral spines and one pair of anterolateral spines; region above latter minutely serrated; also with supraorbital spines from zoea 3; abdominal somite 5 with posterolateral spines, also with small posterodorsal spines in H. multicolorata only; pereiopods 1-2 with exopod, 3-5 poorly segmented and probably not functional; exopods of pereiopods and maxillipeds with no more than six setae.

Nauticaris marionis (Figs. 3D-F)

Only zoea 1 known; carapace with one pair of anterolateral spines; abdomen with small posterodorsal spines on somite 5 and no other spines; antennal endopod almost twice length of exopod, with small accessory seta, and with long tapering distal portion bearing two rows of widely spaced spines.

Tozeuma novaezealandiae (Fig. 3H)

Long and slender larvae with very prominent, forward-pointing dorsomedial spine on somite 3; also with strong posterolateral spines on abdominal somite 5.

Family Ogyrididae

This family is represented in New Zealand waters by a single species, Ogyrides delli Yaldwyn, 1971. Early larvae of O. delli have been reared by the author, although they have not yet been described formally.

The only larvae of an ogyridid which have been described in the literature are those of O. limicola Williams, 1955, described by Sandifer (1974). The larvae of both O. delli and O. limicola are similar in many respects to those of the Hippolytidae. O. delli larvae should easily be identified by substituting the following for characters 1-4 given above for the Hippolytidae:

Ogyridid larval characters

1.Eyestalks short, oviform, close together (Fig. 4A).page 10
2.Carapace with one pair of anterolateral spines (at pterygostomial angles), and one pair of anteroventral spines; rostrum extends beyond eyes in O. delli (but not in O. limicola), with subrostral spines from zoea 2 onward (Fig. 4A).
3.Maxillule without outer plumose seta, and with distal lobe of endopod bearing two setae.
4.Exopod of maxilla from zoea 3 onward with proximal extension, with distal margin smoothly rounded, and with reduction in number of setae on outer proximal margin (Fig. 4B).

The larvae of O. delli are further characterised by the absence of spines on abdominal somites 1-5 (as are O. limicola larvae), and by the loss of telsonic setae 2 and 3 in zoea 4.

Family Palaemonidae

The following species have been recorded from New Zealand waters: Leander tenuicornis (Say, 1818), Palaemon affinis H. Milne-Edwards, 1837, Periclimenaeus novaezealandiae (Borradaile, 1916), Periclimenes yaldwyni Holthuis, 1959 and three as yet unnamed species of Periclimenes (see Richardson & Yaldwyn, 1958).

Larvae of L. tenuicornis have been described by Gurney (1938, 1939) and Gurney & Lebour (1941). Larvae of Palaemon affinis, Periclimenes yaldwyni and one of the unnamed species of Periclimenes have been described by Packer (1983). Larvae of the other species are not known, although Gurney (1924) described larvae from northern New Zealand which could belong to one of the unnamed species of Periclimenes (see Packer 1983, p. 187). Larvae of the two species of Periclimenes described by Packer (1983) were too similar to separate readily in planton samples.

Very little is known of larvae of the genus Periclimenaeus Borradaile. Those described from beyond New Zealand waters are similar in overall morphology to larvae of Periclimenes spp. They may or may not have a toothed rostrum and more conspicuous spinules on the dactylus spine of maxilliped 2 by which they could be distinguished from larvae of the genus Periclimenes (see Gurney & Lebour 1941 (as Periclimenes, subgenus Periclimenaeus) and Fig. 4F).

As a group the larvae of this family are superficially similar to those of the family Alpheidae. They can be identified by substituting the following for characters 1-3 given above for the Alpheidae.

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Palaemonid larval characters

1.Carapace with supraorbital spines bearing retrorse spinules from zoea 2 onward (Figs. 4C, D, K).
2.Pereiopod 5 not much longer than pereiopod 4 when latter developed (Figs. 4C, K), except in Leander (Fig. 4D).
3.Protopod of maxilla with three endites only (Figs. 41, J), exopod longer than endopod except in early larvae of Periclimenes.

Key to larvae of New Zealand Palaemonidae
A. Body with double bend, especially in late larvae; carapace without dorsomedial spines; rostrum shorter than half the length of the antennular peduncle; abdomen without spines on somite 5 (Fig. 4C) Periclimenes spp.
Periclimenaeus novae zealandiae?
Body straight or abdomen curved ventrally; carapace with dorsomedial spines bearing spinules from zoea 2 onward; rostrum subequal in length to antennular peduncle; abdomen with posterolateral spines on somite 5 in all zoeae or from zoea 2 onward (Figs. 4D, K) B
B. Carapace never with more than two dorsomedial spines; pereiopods 1-3 with exopod; pereiopod 5 develops before pereiopod 3, much longer than pereiopods 3-4 when latter developed; Maxilliped 1 with inner margin of basis strongly protuberant in all zoeae; endopod of maxilla without basal lobe (Figs. 4D, H, J) Leander tenuicornis
Carapace with three dorsomedial spines from zoea 4 onwards; pereiopods 1-4 with exopod; pereiopod 5 not developed before pereiopod 3, not much longer than pereiopods 3-4 when latter developed; maxilliped 1 without inner margin of basis strongly protruberant in early larvae; endopod of maxilla with basal lobe (Figs. 4G, I, K) Palaemon affinis
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Acknowledgements

I wish to express my gratitude to Drs R.G. Wear and M.R. Clark (Victoria University of Wellington) for their constructive criticism of the manuscript. I also wish to thank Dr Wear for providing larvae of Tozeuma novaezealandiae and for his help with finding and capturing ovigerous Ogyrides delli. Mr P.E. Roberts (Marine Sciences Laboratory, Queenscliff, Australia) kindly provided larvae of Nauticaris marionis. The Victoria University of Wellington assisted with travel.

References

De Simón, M. , 1979. Primeros estadios larvarios de Pontocaris lacazei (Gourret) (Decapoda, Macrura, Crangonidae) obtenidos en laboratorio. Investigación pesquera 43 : 565-580.

Dobkin, S. , 1965. The early larval stages of Glyphocrangon spinicauda A. Milne-Edwards. Bulletin of Marine Science 15 : 872-884.

Gurney, R. , 1924. Crustacea. Part 9. Decapod larvae. British Antarctic Terra Nova Expedition, 1910. Natural History Reports, Zoology 8 : 37-202

Gurney, R. , 1937. Larvae of decapod Crustacea. Part 4. Hippolytidae. 'Discovery' Reports 14 : 351-404.

Gurney, R. , 1938. The larvae of the decapod Crustacea. Palaemonidae and Alpheidae. Scientific Reports of the Great Barrier Reef Expedition 6 : 1-60

Gurney, R. , 1939. A late larval stage of the Sargas sum prawn, Leander tenuicornis (Say), and a note on the statocyst of the adult. Annals and Magazine of Natural History , series 2, 3 : 120-126.

Gurney, R. , 1942. Larvae of decapod Crustacea . London, Ray Society, pp. 306.

Gurney, R. & Lebour, M.V. , 1941. On the larvae of certain Crustacea Macrura, mainly from Bermuda. Journal of the Linnean Society, Zoology 41 : 89-181.

Hayashi, K-I. & Miyake, S. , 1970. Bathyhippolyte yaldwyni n.gen., n.sp., a deepsea hippolytid (Decapoda, Natantia) from New Zealand. Transactions of the Royal Society of New Zealand, Biological Sciences , 12 : 41-47.

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Kemp, S.W. , 1911, Notes on Decapoda in the Indian Museum. 2. Descriptions of two new Crangonidae with observations on the mutual affinities of the genera Pontophilus and Philocheras . Record of the Indian Museum 6 : 5-12.

Kemp, S.W. , 1916. Notes on Crustacea Decapoda in the Indian Museum. 6. Indian Crangonidae. Record of the Indian Museum 12 : 355-384.

Lebour, M.V. , 1954. The position of Pontophilus echinulatus (M. Sars) in the Crangonidae. Journal of the Marine Biological Association of the United Kingdom 33 : 587-588.

Makarov, R.R. , 1968. On the larval development of the genus Sclerocrangon G.O. Sars (Caridea, Crangonidae). Crustaceana , supplement 2 : 27-37.

Packer, H.A. , 1983. Larval morphology of some New Zealand shallow water shrimps (Crustacea, Decapoda, Caridea) of the families Crangonidae, Hippolytidae, and Palaemonidae. M.Sc. thesis, Victoria University of Wellington, pp. 232 (unpublished).

Pillai, N.N. , 1974. Laboratory reared larval forms of Hippolysmata (Exhippolysmata) ensirostris Kemp (Decapoda: Hippolytidae). Journal of the Marine Biological Association of India 16 : 594-608.

Richardson, L.R. & Yaldwyn, J.C. , 1958. A guide to the natant decapod Crustacea (shrimps and prawns) of New Zeland. Tuatara 7 : 17-41.

Sandifer, P.A. , 1974. Larval stages of the shrimp, Ogyrides limicola Williams, 1955 (Decapoda, Caridea) obtained in the laboratory. Crustaceana 26 : 37-60.

Sars, G.O. , 1890. Bidrag til Kundskaben om Decapodernes Forvandlinger. 3. Fam. Crangonidae. Archiv for Mathematik og Naturvidenskab 14 : 132-195.

Wear, R.G. , 1985. Checklist and annotated bibliography of New Zealand decapod crustacean larvae (Natantia, Macrura Reptantia, and Anomura). Zoology Publications from Victoria University of Wellington 79 : 1-15.

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Williamson, D.I. , 1957. Crustacea, Decapoda: Larvae, 1. General. Conseil International pour l'Exploration de la Mer. Fiches d'identification du zooplancton 67 : 1-7.

Williamson, D.I. , 1960. Crustacea, Decapoda: Larvae, 7. Caridea, Family Crangonidae, Stenopodidea. Conseil International pour l'Exploration de la Mer. Fiches d'identification du zooplancton 90 : 1-5.

Williamson, D.I. , 1982. Larval morphology and diversity. In: Bliss, D.E. & Abele, L.G., eds., The Biology of Crustacea. Vol. 2. Embryology, Morphology, and Genetics , pp. 43-110. New York, Academic Press.

Wollebaek, A. , 1906. Le developpement du genre Sclerocrangon (G.O. Sars). Bergens museums aarbog 11 : 1-9.

Yaldwyn, J.C. , 1954. A preliminary survey of the New Zealand Crustacea Decapoda Natantia with an appendix on the reptant genus Nephrops . M.Sc. thesis, Victoria University of Wellington, pp. 544 (2 vols.) (unpublished).

Yaldwyn, J.C. , 1959. The New Zealand natant decapod Crustacea: Systematics, distribution and relationships. Ph.D. thesis, Victoria University of Wellington, pp. 435 (unpublished).

Yaldwyn, J.C. , 1960. Crustacea Decapoda Natantia from the Chatham Rise: a deep water bottom fauna from New Zealand. New Zealand Department of Scientific and Industrial Research bulletin 139 (New Zealand Oceanographic Institute Memoir 4) : 13-53.

Yaldwyn, J.C. , 1971. Preliminary descriptions of a new genus and twelve new species of natant decapod Crustacea from New Zealand. Record of the Dominion Museum, Wellington 7 : 85-94.

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Abbreviations in Figures 1-4

A1antennule
A1Pantennular peduncle
A2antenna
ALSanterolateral spine
AS1-5abdominal somites 1-5
AVSanteroventral spine
BLbasal lobe
DMSdorsomedial spine
DSdactylus spine
E1-4endites 1-4
endendopod
exexopod
feexternal flagellum
fiinternal flagellum
M1-3maxillipeds 1-3
OSoutside seta
P1-5pereiopods 1-5
PDSposterodorsal spine(s)
PEproximal extension
PLBpleopod bud
PLSposterolateral spine
Rrostrum
S1-8telsonic setae 1-8
SOSsupraorbital spine
Ttelson
Uuropod
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Legends for Figures 1-4
(scale in mm)

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Figure 1: A - Betaeopsis aequimanus zoea 3, carapace and abdomen, lateral view. B,C,G,H - Alpheopsis garricki zoea 3, lateral view (B), enlarged tip of pereiopod 5 dactylus spine (C), maxilliped 3 (exopod omitted) (G), maxilla (H). D - Alpheus richardsoni zoea 3, maxilliped 1 (exopod omitted). E, F - Alpheus socialis zoea 3, maxilliped 1 (E) and 3 (F).

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Figure 2: A - Pontophilus australis zoea 5, anterior dorsal view. B -Cheraphilus (=Pontophilus) echinulatus zoea 5, anterior dorsal view, showing rostrum shape in 'Pontophilus-group B' species. C-E - Pontophilus chiltoni, zoea 5 maxilla (C), zoea 5 lateral view (D), zoea 3 telson and uropods (E). F - Sclerocrangon boreas zoea 1, telson. G - Glyphocrangon spinicauda zoea 1, telson. H - Pontophilus spinosus zoea 5, anterior dorsal view showing the length of antennullar flagellum in 'Pontophilus-group A' species. (B, H after Sars 1890, F after Makarov 1968, G after Dobkin 1965, I after De Simón 1979)

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Figure 3: A, B - Hippolyte multicolorata zoea 8, lateral view (A), enlarged dorsal view of spines on abdominal somite 5 (B). C -Hippolysmata ensirostris zoea 7, lateral view of cephalothorax (note enormous length of pereiopod 5; other thoracic appendages omitted). D-F - Nauticaris marionis zoea 1, dorsal view (D), antenna (E), enlarged dorsal view of spines on abdominal somite 5 (F). G -Alope spinifrons zoea 5, lateral view. H, K - Tozeuma novaezealandiae zoea 1, lateral view (H), maxilla (K). I, J - Hippolyte bifidirostris zoea 3, maxilliped 3 (I), maxillule (J). (C after Pillai 1974)

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Figure 4: A, B - Ogyrides delli zoea 4, dorsal view of cephalothorax (A), maxilla (B). C - Periclimenes yaldwyni zoea 8, lateral view. D, H, J - Leander tenuicornis zoea 5(?), lateral view (D), maxilliped 1 (H), maxilla (J). E, G, I, K - Palaemon affinis zoea 6, maxilliped 2 (E), maxilliped 1 (G), maxilla (I), lateral view (K). F - Periclimenaeus (?) wilsoni last larva, maxilliped 2 (exopod omitted). (D, H, J after Gurney 1938, F after Gurney & Lebour 1941, F, H, J scale unknown.)