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Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

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Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 104 Band

May 1995 Mei

Part 8 Deel

REVISION OF THE FAMILY PARAMELITIDAE

(CRUSTACEA, AMPHIPODA)

FROM SOUTH AFRICAN FRESH WATERS

BARBARA A. STEWART

CHARLES L. GRIFFITHS

Cape Town Kaapstad

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D3903

REVISION OF THE FAMILY PARAMELITIDAE

(CRUSTACEA, AMPHIPODA)

FROM SOUTH AFRICAN FRESH WATERS

BARBARA A. STEWART

South African Museum, P.O. Box 61, Cape Town, 8000, South Africa

CHARLES L. GRIFFITHS

Zoology Dept, University of Cape Town, Rondebosch, 7700, South Africa

(With 29 figures and 2 tables)

[MS accepted 6 February 1995]

ABSTRACT

Twenty-five species of freshwater amphipod, including one new species, are recognized

within the family Paramelitidae. These are allocated to three genera, Paramelita, Aqua-

dulcaris gen. nov. and Mathamelita gen. nov. Each genus and species is diagnosed and all

species are illustrated. Distribution records are updated, and new keys for the identification

of the species are provided. Morphological similarities between the species are discussed.

CONTENTS

PAGE

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MP ALCTAAIS ANG SIMIC OUS erie reese ee ao eee ete ee ers SEN Nc Aetna RSA Raye 183

MB IAGISEICLANALYSIS We Soren. trines cise seasons cSen ee swe snl oe Se oelace mc aR Mees ERR el rcieminwereataee 184

PARCHELE SAM ALY SIS access ta eens oe se ce cero es ate sere nae teats Saree eels nig each athe emai ane arses 190

CS RITSTI RITES She beac. Cac ic tobi cea n Ue Nc aoe aac BEC BC Horns HOSES Tapers amt ci clin aetna ao 192

AC MHOWICOP CMENES ccnaae cece re creel eae ioe ioc ite eeee eee ce bee ne Taman aes Hee aise tote See 246

RR LEE NCES et raga cree a ata GS AY tae aks ed His DING SP aviedie ah Sy mad aL rete a 246

INTRODUCTION

South African freshwater amphipods first received attention from Barnard

(1916) who described four species from Table Mountain, Cape Peninsula, plac-

ing them in the genus Gammarus. Schellenberg (1926) erected the genus Para-

melita to accommodate the new species P. ctenodactyla, collected from the

Cape Peninsula, but a year later, Barnard (1927) listed this species as a syn-

onym for ‘G. capensis’. Thus, although it is clear that Barnard (1927) had seen

Schellenberg’s (1926) paper, he did not recognize, nor discuss, the validity of

Schellenberg’s proposed new genus Paramelita. Instead, he extended the known

ranges of two of his existing ‘Gammarus’ species and added six further new

species and one variety. Schellenberg (1937) again argued that South African

freshwater species assigned to Gammarus were sufficiently different from those

181

Ann. S. Afr. Mus. 104 (8), 1995: 181-247, 29 figs, 2 tables.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the Palaearctic and Nearctic regions to warrant separate generic status under

the name Paramelita. Thus, when Thurston (1973) described a new cave-

dwelling amphipod from the Cape Peninsula, he placed it in this genus, recog-

nizing the transfer of South African freshwater ‘Gammarus’ species to Para-

melita. A further new species of Paramelita was added to the fauna by Griffiths

(1981), bringing the total to 12 species and one variety.

Present collections by the authors in 1989 and 1990 have doubled this num-

ber of species. Four of the most distinctive species were described by Stewart &

Griffiths (1992a) and, after further morphological and isozyme analysis, eight

other new taxa were added by Stewart (1992), Stewart & Griffiths (1992b,

1992c) and Stewart et al. (1994). Thus, since Griffiths (1981) last revised the

genus, 12 new species have been added, bringing the total to 24.

Despite the fact that a number of species of Paramelita did not fit his

original generic diagnosis, Schellenberg (1937) did not extend this; neither did

Griffiths (1981) comment on or rediagnose the genus in his revision. Barnard &

Barnard (1983) provided a rediagnosis of the genus in their study of freshwater

amphipods of the world, but this contains some inaccuracies and also does not

adequately accommodate all of the species described to date. Thus, a relatively

large assemblage of morphologically variable paramelitid species from the

south-western regions of South Africa is currently assigned to the single, poorly

diagnosed genus. Phylogenetic relationships between these species are

unknown.

The closest relatives to the South African genus Paramelita are 21 species

of Australian crangonyctoids, which are placed in seven genera in the family

Paramelitidae, all rediagnosed or described in a recent detailed account of the

Australian crangonyctoids by Williams & Barnard (1988). Although a cladistic

analysis of the family Paramelitidae has yet to be undertaken, Williams &

Barnard (1988) did comment on relationships between the genera in their

revision of the Australian crangonyctoid amphipods. Thus, they regarded

Austrogammarus as the most primitive of Australian paramelitids, Antipodeus as

being ‘much closer to Austrocrangonyx than to Austrogammarus’ (p. 54),

Hurleya as having ‘more affinities with paramelitid genera than with neoni-

phargid and perthiid genera’ (p. 86), and Uroctena as having ‘strong affinities

with Paramelita’ (p. 91).

The only other freshwater gammarid amphipods from the south-western

Cape are five species assigned to the genus Sternophysinx (Holsinger 1992).

There is some dispute as to how closely related Sternophysinx is to Paramelita.

Bousfield (1983) placed this genus in the Paramelitidae, but Williams & Barnard

(1988) did not recognize its inclusion in the family in their revision of Austral-

ian crangonyctoids. Holsinger (1992) also concluded that sternophysingids are

sufficiently different from the other paramelitids to warrant recognition as a

separate family, the Sternophysingidae.

The aim of this paper is to provide a revision of the taxonomic status of

southern African species falling within the family Paramelitidae. Relationships

between the South African paramelitid species are investigated by means of a

cladistic analysis of morphological data. The polarity of the characters used was

determined by outgroup comparison (Watrous & Wheeler 1981), and the com-

puter program HENNIG86 (Farris 1988) used to generate most parsimonious

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 183

trees. The genus Paramelita is redefined and new genera are created to

accommodate species falling outside this definition. Keys are provided to adult

males, and all species, some of which have not been adequately depicted in

earlier papers, are illustrated. Full distribution records are provided for all

forms, many of which extend the ranges previously recorded. Finally, the

opportunity is taken to describe a newly discovered and remarkable new

species, which is allocated separate generic status under the name Mathamelita

gen. nov.

MATERIALS AND METHODS

This analysis is based both on an extensive series of samples gathered by the

authors between 1989 and 1993 and deposited in the South African Museum

(SAM), and on existing collections of the South African Museum. Specimens of

both sexes were examined from all populations collected to assess the level of

morphological variability. When necessary, specimens were partially dissected

to facilitate measurement and illustration of the limbs. Drawings were made

with the aid of a camera lucida attached either to Wild dissecting or compound

microscopes.

A total of 24 characters, referring to the external morphology of the amphi-

pods, could be successfully polarized. Quantitative characters were gap-coded

by plotting histograms of all quantitative characters, and coding the character

states according to ‘identifiable’ gaps (Conlan 1988; Notenboom 1988). Hom-

ologous characters were recognized by similarity in positions and connection

with other body parts. Character states were polarized using outgroup compari-

son, where character state distributions in other paramelitid genera were deter-

mined largely by a survey of the literature (e.g. Williams & Barnard 1988). In

the case of quantitative characters not supplied in the descriptions, these were

scored from illustrations (see also Conlan 1988). Unknown polarities were

coded in the data matrix with a question mark. Unique character states found

only in one species were considered to be autapomorphies, and were excluded

from the numerical analysis.

The data matrix was analysed by means of the HENNIG86 package (Farris

1988). Most parsimonious trees, which minimize the number of changes in

character states needed to explain the pattern of character state distribution

among the taxa, were derived from the character state matrix by means of the

‘mh*’ and ‘bb’ commands. Although the ‘ie*’ command is certain to find all

trees of minimal length, this command proved to be prohibitively time-

consuming for the analysis of Paramelita. The ‘mh*’ command was therefore

selected as the next-best choice, as recommended by Farris (1988) in the docu-

mentation accompanying the program. A consensus tree was constructed by

means of the ‘nelsen’ command.

Morphological similarity between species was also investigated pheiuetically

by calculating a matrix of Simple Matching Coefficients based on 29 variables.

A cluster analysis based on this matrix was performed by means of the UPGMA

algorithm. This analysis was performed with the aid of the NTSYS-pc computer

program (Rholf 1989).

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

CLADISTIC ANALYSIS

Selection of outgroups

It is commonly held (e.g. Ridley 1986) that the most suitable outgroup to

choose is that of a closely related species or genus. Obvious candidates would

therefore be other genera within the Paramelitidae. Unfortunately, the choice of

suitable outgroups is complicated by the fact that the composition of Paramelit-

idae is still under question (e.g. Bousfield 1983; Williams & Barnard 1988).

Notenboom (1988: 160) encountered a similar situation in his study of the

phylogeny of Pseudoniphargus, and commented that ‘An important obstacle in

phylogenetic studies of amphipods at lower taxonomic levels is the highly

debated classification into families and superfamilies’. For the phylogenetic

analysis of Paramelita, Austrogammarus and Austrocrangonyx were chosen as

outgroups. Like Paramelita, these two genera possess sternal gills, a coxal gill

on pereopod 7, a short second segment on the outer ramus of uropod 3 and a

cleft telson. Williams & Barnard (1988) regard Austrogammarus as the most

plesiomorphic genus of Australian paramelitids. Although these authors have

suggested that Uroctena has strong affinities with Paramelita, a Uroctena type

ancestor for Paramelita would have involved losing and regaining the coxal gill

on pereopod 7, which is a plesiomorphic crangonyctoid ‘marker’. Similarly,

Hurleya, Protocrangonyx or Giniphargus type ancestors would also involve the

loss and regaining of a coxal gill on pereopod 7, and an Antipodeus type ances-

tor, the loss and regaining of sternal gills. Thus, it seems more probable that

Austrogammarus and Austrocrangonyx are closest to the hypothetical ancestor

of Paramelita.

Selection and polarity of characters

The characters that differentiated species of Paramelita and that were used

in the numerical analysis are listed in Table 1, and the distributions of character

states over the species are given in Table 2. Characters were either ‘qualitative’,

such as presence or absence or differences in shape, ‘quantitative’ and ‘discon-

tinuous’, such as counts of spinules, or ‘quantitative’ and ‘continuous’, such as

the relative lengths of limbs. As in Notenboom’s (1988) and Conlan’s (1988)

cladistic studies on amphipods, ratios and counts were included in the present

analysis, despite reservations by some authors (e.g. Pimental & Riggins 1987)

about quantitative data. These characters avoided the use of subjective character

state definitions, and were needed because of the shortage of usable characters.

Care was taken to identify clear gaps when coding these data.

The structure of antenna 2 in adult males showed interesting differences

between the paramelitid species. In eight species, the ‘pediformity’ alluded to by

Williams & Barnard (1988) in the Australian genus Uroctena, was clearly evi-

dent, with articles 3, and particularly 4, strongly swollen in adult males (e.g.

Figs 3C, 8C). Based on outgroup analysis, this condition was considered apo-

morphic in southern African paramelitid species. The presence of teeth

(Figs 6C, 20C, 26C), lobes (Figs 3C, 4C, 8C, 14C) and ridges (Fig. 23C) on

antenna 2 in eight Paramelita species was also apomorphic, as was the elonga-

tion of the peduncle of this antenna in six Paramelita species (Figs 17C, 18C,

DUC. 23. 24C.28C).

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 185

TABLE |

Descriptions of the 24 characters used for cladistic analysis of paramelitid species from South

Africa. Characters apply to adult males, and have been polarized using Austrogammarus and

Austrocrangonyx as outgroups.

No. Character States

Antenna 1

Q Ratio of lengths of antenna 1 : antenna2 (0) > 1.2; (1) 0.7-1.0

1 Ratio of length : width of article 1 (0) 1.8-2.3; (1) 2.6-4.3

Antenna 2

2 Number of articles in flagellum (O) 12-30+; (1) < 12

3 Ratio of length : width of article 3 (0) 0.7-0.9; (1) 1.1-1.9

4 Lobe on article 3 (0) absent; (1) present

5 Shape of article 4 (0) normal; (1) elongate and stout;

(2) strongly laterally swollen

Gnathopod 1

6 Medial spines on article 2 (0) absent; (1) present

Gnathopod 2

7 Angle of palm (0) transverse to slightly oblique;

(1) moderately to strongly oblique

8 Medial spines on article 2 (0) absent; (1) present

Pereopod 3

9 Medial spines on article 2 (0) absent; (1) present

10 Width of article 4 (0) normal; (1) widening distally

11 Projection on article 4 (0) absent; (1) present

12 Tooth-like spines on article 5 (0) absent; (1) present

13. Shape of article 6 (0) normal; (1) arched

14 Spinules on dactyl of article 7 (0) 1; (1) 2-6; (2) 7-8

Pereopod 4

15. Posterior margin of coxa 4 (0) distinctly excavate; (1) slightly emarginate

16 Spinules on dactyl of article 7 (0) 1; (1) 2-6

Pereopods 5-7

17 Spinules on dactyl of article 7 (O) 1; (1) 2-10; (2) > 10

Uropod 1

18 Setation of outer ramus (0) absent; (1) present

Uropod 2

19 Setation of inner ramus (0) absent; (1) present

20 Setation of outer ramus (0) absent; (1) present

Uropod 3

21 Ratio of length of inner : outer ramus (0) 0.6-0.7; (1) 0.1-0.4; (2) 1.0

22 Article 2 on outer ramus (0) 7-20% of article 1; (1) < 7%;

(2) rudimentary or absent

Telson

23 Spination (0) 0-1 spines per lobe; (1) > 1 per lobe

ANNALS OF THE SOUTH AFRICAN MUSEUM

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REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 187

Gnathopod 2 differed amongst Paramelita species in terms of spination of

article 2 and the nature of the palm in article 6. Like Austrogammarus and

Austrocrangonyx, 15 species of Paramelita had transverse to slightly oblique

palms (e.g. Fig. 4D), whereas nine Paramelita species had moderately to

strongly oblique palms (e.g. Fig. 12D). The presence of spines on article 2 of

gnathopod 2 was common amongst South African paramelitids, with 11 Para-

melita species sharing this condition (c.g. Fig. 8D). A survey of the literature

(e.g. Williams & Barnard 1988) revealed that none of the six Austrogammarus

or two Austrocrangonyx species have spines of this type. Their presence was

thus considered as apomorphic in Paramelita.

None of the eight species examined in the two Australian genera possessed

a claw-like pereopod 3 of the type found in adult males of five Paramelita

species (Figs 3E, 4E, 7E, 8E, 23E). Closer examination of this condition

revealed that the ‘claw’ is achieved in several different ways through various

modifications of either articles 4 or 5, and 6. This condition has obviously

arisen in some of these species as a result of convergent evolution, so that the

possession of a claw-like pereopod 3 is not necessarily evidence of close affinity

between them.

Coxa 4 varies from being quadrate (e.g. Fig. 24F) to having its posterior

margin strongly excavate (e.g. Fig. 13F) in Paramelita species. In Austro-

gammarus and Austrocrangonyx, this coxal plate is excavate. When Barnard

(1916, 1927) first described Paramelita species, he was unable to decide which

condition was plesiomorphic. However, outgroup analysis suggests that a

strongly excavate coxa 4 is plesiomorphic in Paramelita.

Eighteen Paramelita species were characterized by a multispinose dactyl in

pereopods 3-7 (e.g. Fig. 16F) and, in at least two species, as many as 14 spin-

ules were counted on the dactyl of pereopod 6! This multispinose condition is

apomorphic, as all Austrogammarus and Austrocrangonyx species have at most

one spinule on the margin of the dactyl of each pereopod.

Uropods 1 and 2 differed mainly in the degree of setation of the rami. In

Austrogammarus and Austrocrangonyx, the outer rami of these uropods are

never setose, but in six Paramelita species, setae were present on one or both of

these rami (e.g. Fig. 17H-I). It is likely, however, that this setose condition has

evolved more than once, so that the sharing of this condition could represent an

example of convergent evolution.

The relative length of the inner ramus of uropod 3 differed markedly

between the outgroups. In Austrogammarus, this appendage is usually about

60-70 per cent of the length of the outer ramus, but in Austrocrangonyx, the

inner ramus is only about 10 per cent the length of the outer ramus. This figure

varied from 10-40 per cent in Paramelita. Ten of the Paramelita species were

characterized by the virtual absence of a second segment on the outer ramus of

uropod 3 (e.g. Figs 31, 4J), whereas the remaining species had a small, but

distinct, article 2 on the outer ramus of uropod 3 (e.g. Fig. 13J).

On the whole, relatively few characters could be polarized, and thus only

these were usable. Many characters, such as eye colour, setation of the antennae

and limbs, and the relative lengths of the peduncle and flagellum in the

antennae, had variable character states within the outgroups. Notenboom (1988)

also complained of a shortage of usable characters in his study of the amphipod

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

genus Pseudoniphargus, and pointed out that this was a general problem when

working at a low taxonomic level.

Phylogeny of Paramelita

In the initial analysis, use of the ‘mh*’ and ‘bb’ commands resulted in

116 most parsimonious trees with consistency indices of 0.40. A consensus tree

was constructed using the ‘nelsen’ command (Fig. 1). Six of the species formed

a monophyletic group (group A) defined by the possession of a strongly laterally

swollen article 4 in antenna 2. Relationships within this group of six species

were fully resolved. The monophyletic subgroup comprising P. pheronyx,

P. andronyx and P. auricularia was defined by two synapomorphies—the pos-

session of a posterior lobe on article 3 of antenna 2, and a characteristic distal

widening of article 4 of pereopod 3. Two of these species are known from

adjacent streams, but P. andronyx is known only from an isolated mountain

massif over 75 km north of the P. auricularia and P. pheronyx localities. This

clade, thus, has a rather disjunct distribution.

The remaining species in group A—P. crassicornis, P. dentata and P. mar-

unguis—have been found in streams on the Cape Peninsula that are at most

15 km apart. The geographical distribution of these species, therefore, supports

the proposed existence of this clade.

It is proposed that species falling into group A be recognized as belonging to

a new, monophyletic genus, Aquadulcaris gen. nov., the description of which

follows in the systematic section. Species in this proposed genus do not fit the

original diagnosis of the genus Paramelita Schellenberg, 1926. Schellenberg

(1926: 367) described dactyls with a row of spinules, and also alluded to the

second segment on the outer ramus of uropod 3 as being well formed in species

of Paramelita. All of the six species falling in group A usually have only a

single spinule on the dactyl of pereopods 3-7, and the second segment on the

outer ramus of uropod 3 is rudimentary or absent.

The only other species that have article 4 of antenna 2 markedly laterally

swollen are P. tulbaghensis and P. spinicornis. The former species is in fact

very similar to the group A species, and shares features such as a moderately

emarginate (rather than distinctly excavate) coxa 4 and the absence of an outer

segment on the outer ramus of uropod 3. It differs, however, in the possession

of more than one spinule (2-4) on the dactyl of each pereopod. Preliminary

analysis of electrophoretic data (Stewart in prep.) also suggests the inclusion of

P. tulbaghensis in Aquadulcaris gen. nov. However, P. tulbaghensis has been

left in the genus Paramelita in this review, pending further investigations

regarding the phylogenetic relationships of this species.

The ‘pediformity’ of antenna 2 in P. spinicornis is more likely to represent

an example of convergent evolution. This species has an excavate rather than

poorly emarginate coxa 4, 3-4 spinules on the dactyl of pereopods 3-4 and 5-8

spinules on pereopods 5-7, and a distinct but small second article on the outer

ramus of uropod 3. It is, therefore, more likely that lateral swelling of article 4

of the peduncle of antenna 2 has occurred more than once in different lineages,

and does not suggest that P. spinicornis and the species of Aquadulcaris are

related.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 189

Austrogammarus |

Austrocrangonyx

sdnoi63no

. dentata

. Crassicornis

marunguis

. auricularia

vy dnoiy

andronyx

pheronyx

flexa

. kogelensis

nigroculus

. seticornis

. Spinicornis

tulbaghensis

. aurantia

granulicornis

. pillicornis

q dnoi5

. capensis

parva

barnardi

validicornis

magna

magnicornis

. odontophora

wv VD DHDDBD DDH DDVD® DD HDDH DN DDN

. pinnicornis

a°)

platypus

Fig. 1. Cladistic analysis of southern African paramelitids.

Aquadulcaris gen. nov. is almost certainly most closely allied to the Aus-

tralian genus Uroctena, which also includes members with ‘pediform’ second

antennae and poorly emarginate fourth coxal plates (Williams & Barnard 1988).

The unresolved nature of group B can be attributed to a combination of the

relative shortage of ‘good’ apomorphies and the possibility that the genus, as

currently composed, is not monophyletic. For example, it is possible that “weak’

synapomorphies are shared due to parallel evolution rather than due to common

descent. A good example of this is the possession of medial spines on article 2

of pereopod 3, a condition shared by Paramelita granulicornis and P. capensis.

Myers (1988) also expressed concern about the effect of parallel evolution on

the development of a synapomorphic scheme and concluded, from his study of

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

amphipods in the family Aoridae, that parallel evolution was a far more com-

mon phenomenon than previously believed. He suggested that in amphipods, in

general, complex character states are rare and variation morphoclinal, thus

making it difficult to detect cases of parallel evolution in these animals (Myers

1988). In addition, Notenboom (1988) has pointed out that at low taxonomic

levels, species share similar gene pools, thus increasing the chances of the

occurrence of parallelisms.

Within group B, the cladogram shows that the species P. aurantia and

P. granulicornis are closely related. The fact that P. aurantia and P. granuli-

cornis are geographically close provides additional evidence for the validity of

this clade. Paramelita aurantia and P. granulicornis share many apomorphic

conditions, such as an almost quadrate coxa 4, medial spines on article 2 of

gnathopod 2, and the absence of a second article on the outer ramus of uro-

pod 3. Griffiths (1981: 82) has suggested that these ‘Hottentots Holland Moun-

tain’ species, along with P. kogelensis and P. seticornis, form a ‘closely related

group’. When Barnard (1927) first described P. kogelensis, he considered it to

be ‘closely allied’ (p. 172) to P. seticornis. It is highly probable that the present

analysis does not include all the extant species of this group, so that a fully

resolved cladogram is not possible at this stage.

The group consisting of P. magna, P. magnicornis, P. odontophora, P. pin-

nicornis and P. platypus is characterized mainly by the possession of elongate

and stout second antennae. Paramelita magna and P. magnicornis occur, often

in sympatry, in streams in the southern part of the Cape Peninsula. Paramelita

odontophora and P. pinnicornis are also relatively close geographically to each

other, with the most easterly known population of P. pinnicornis only about

15 km away from the nearest population of P. odontophora. Paramelita pinni-

cornis is also known from two localities on the Cape Peninsula.

The most unlikely monophyletic group is that of P. parva and P. barnardi,

as these species occur over 400 km apart, with the latter species known only

from a single cave on the Cape Peninsula.

Although the assumption has been made that all existing species have been

included in the cladistic analysis, it is possible that a more thorough search of

streams in the mountainous areas of the south-western Cape could reveal more

as yet undescribed forms.

PHENETIC ANALYSIS

A phenetic analysis of the 25 known paramelitid species, including the new

species described in this paper, confirmed the existence of morphologically

distinct groups within the species (Fig. 2). Six of the species fell into a rela-

tively ‘tight’ and distinct cluster (cluster A), and were easily distinguished from

the remaining species by a combination of characteristic features. These

included the possession of a sparsely setose urosome and a poorly emarginate

coxa 4; the presence of only a single spinule on the dactyl of pereopods 3 and 4,

and (with the exception of one population of Paramelita auricularia) on the

dactyl of pereopods 5-7; a strongly swollen article 4 in antenna 2, and the

absence of a second segment on the outer ramus of uropod 3. This cluster

represents the six species belonging to Aquadulcaris gen. nov.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE

191

Seventeen species grouped together to form ‘cluster B’. This cluster was

clearly divisible into smaller distinct ‘subclusters’. Like the Aquadulcaris

species (cluster A), Paramelita aurantia and P. granulicornis are characterized

by the possession of a poorly emarginate coxa 4, and the absence of a second

article on the outer ramus of uropod 3. However, these species have a multi-

spinose dactyl on each pereopod, and the peduncle of antenna 2 is never mark-

edly swollen laterally. With the exception of P. seticornis and P. tulbaghensis,

all of the remaining ‘cluster B’ species have an excavate coxa 4 and a distinct

second segment on the outer ramus of uropod 3. Although it is possible that

these 16 species should be considered as belonging to more than one genus, it is

proposed that, until further detailed cladistic analysis is undertaken, they remain

in the genus Paramelita.

ba 8) a a) eee ae Ae yea) al a) hea) ae) a)

a aL ae Lan cL

0.4 0.5 0.6 0.7 0.8 0.9 1.0

Simple matching coefficient

andronyx

. pheronyx

. auricularia

. Crassicornis

. dentata

marunguis

aurantia

granulicornis

flexa

. kogelensis

. Spinicornis

tulbaghensis

barnardi

parva

pillicornis

seticornis

magna

odontophora

pinnicornis

validicornis

capensis

magnicornis

nigroculus

M. aequicaudata }

P. platypus Jj

Fig. 2. Phenetic analysis of southern African paramelitids.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

In January 1992, a collection of unusual paramelitid specimens was made

from the Outeniqua Mountains near Knysna by CLG. This population is rep-

resented by cluster C in the phenogram. In these specimens, there is no coxal

gill on pereopod 7, and the inner and outer rami of uropod 3 are of equal length

in adult males. All other South African paramelitids have a coxal gill on pereo-

pod 7, as do species of the Australian paramelitid genera Austrogammarus,

Austrocrangonyx and Antipodeus. The inner ramus of uropod 3 is either absent

or shorter than the outer ramus in all known species of paramelitids both in

South Africa and Australia. The occurrence of rami of equal length in uropod 3

in these specimens is thus a highly diagnostic feature, and this, together with the

absence of a coxal gill on pereopod 7, supports the recognition of a new, mono-

specific genus, Mathamelita gen. nov., which is described below.

A unique combination of character states in Paramelita platypus is reflected

in the position of this species in a separate cluster D in the phenogram. This

species is characterized by the possession of an almost quadrate coxa 4, a

strongly convex palm with a palmar tooth in gnathopod 2, lateral expansion of

article 4 in both pereopods 3 and 4, and a small but distinct second segment on

the outer ramus of uropod 3. The combination of a distinctly quadrate coxa 4

and a palmar tooth in gnathopod 2 is shared with P. granulicornis and, although

article 4 of pereopod 4 in this species is not as markedly posterodistally pro-

truded as in P. platypus, Barnard (1927: 176), in his description of P. granuli-

cornis, remarked that this article was ‘rather strongly expanded distally’ in this

species. These two species also occur in close proximity geographically.

SYSTEMATICS

Superfamily CRANGONYCTOIDEA Bousfield, 1973

Family Paramelitidae Bousfield, 1973

KEY TO THE SOUTH AFRICAN PARAMELITID GENERA

ACS Pereopodi7/iicoxalepillkabsenti.-ssss.-teoe eee see Mathamelita gen. nov.

IB* Pereopod’ 7, coxal’ gill present: %;....22.0.sse0sccs+oceseener sone cee ee eee Zz

2A. Antenna 2, article 4 always laterally swollen (e.g. Figs 3C, 4C, SC);

pereopods 3 and 4, dactyl with a single spinule; pereopods 5-7, dactyl

usually with one spinule, rarely two .................. Aquadulcaris gen. nov.

2B. Antenna 2, article 4 sometimes elongate and stout (e.g. Figs 17C, 18C),

but rarely laterally swollen; pereopods 3 and 4, dactyl usually with

2-8 spinules, rarely one; pereopods 5-7, dactyl usually with 2-14 spinules,

RaAnelyfOne: <sccs6ak eke ti See sca Saded are dela nin tse eee eee Paramelita

Aquadulcaris gen. nov.

Diagnosis

Eyes white. Antenna | longer than 2, peduncle sparsely setose, article |

1.2-1.5 length of article 2, 3.0-3.8 longer than wide, flagellum sparsely to

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 193

densely setose, 16- to 30-articulate, accessory flagellum 3- to 5-articulate.

Antenna 2 sparsely setose, peduncle either shorter or longer than flagellum,

article 4 laterally swollen, 1.9-2.7 longer than wide, semicircular lobe on

article 3 present or not, flagellum 8- to 20-articulate. Gnathopod 2, article 2

medially spinose or not, palms slightly to markedly convex, transverse to

slightly oblique. Pereopod 3 modified or not, article 4 either posterodistally

protruded or not, article 5 lobed or not, often bearing 1-4 tooth-like spines.

Coxa 4, posterior margin transverse to slightly emarginate. Pereopods 3 and 4,

dactyl usually with a single spinule, pereopods 5-7, usually with one, but

occasionally with two spinules. Segments 2-7 with 1-4 sausage-shaped sternal

gills, coxal gill present on pereopod 7. Uropod 3, second segment of outer

ramus rudimentary to absent.

Etymology

From the Latin aqua dulcis, meaning fresh water, and caris, meaning

shrimp. The gender is feminine.

Type species. Gammarus crassicornis Barnard, 1916: 207-209, pl. 27

(figs 24-25).

KEY TO SPECIES OF AQUADULCARIS

Pane bercopods unmodified (Fig. 2GE)ia.saesctees cote ee coke mca nae oe A. dentata

1B. Pereopod 3 modified, either article 4 posterodistally protruded to form a

lobe or spur (Figs 3E, 8E), or article 5 posteriorly lobed (Fig. 4E), or with

i—Atooth-like’spines i (Fics SE a E)ice. tse eaee ec eee eeeec mtneen carats 2

2A. Antenna 2, article 3 bearing a semicircular lobe (Figs 3C, 4C, 8C) ....... 3

DEP Amtcina? articles lackinpsallobe’ ye-eaecntes anes estee coos: < aennaeae ce sen emt >)

3A. Pereopod 3, article 4 not posterodistally protruded, article 5 usually

posteriorly lobed, always bearing a tooth-like spine (Fig. 4E) ................

4A. Pereopod 3, article 4 short, posterodistally protruded into a long, narrow

‘spur’; antenna 2, article 3 strongly swollen and enlarged (Fig. 8A, C, E)

8 ICE ORCS SEC OO ERED COE OA SINE Ee SA. or A AIA a Uae A. pheronyx

4B. Pereopod 3, article 4 long, posterodistally protruded into a triangular-

shaped lobe; antenna 2, article 3 moderately swollen (Fig. 3A, C, E) ......

ee Sa eR ees LTA S18. SatiOee SRO AUNT ae heuiN tag meen wane fits A. andronyx

5A. Pereopod 3, article 5 attached normally to article 6 (Fig. SE) ................

By ee are eRe een oes oebroh cin ere ce Sitios ae ae eee ne eee Wao A. crassicornis

5B. Pereopod 3, article 5 attached at right angles to article 6 to form a ‘claw’

CERO RIE I eset eee cw nccises Saas nae eon Chere ence miine eked A. marunguis

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Aquadulcaris andronyx, SAM-A40017, holotype, male, 16.1 mm. A. Lateral

aspect. B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Pereopod 6.

G. Uropod 1. H. Uropod 2. I. Uropod 3. J. Telson. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 195

Aquadulcaris andronyx (Stewart & Griffiths, 1992)

Fig. 3

Paramelita andronyx Stewart & Griffiths, 1992a: 148-153, figs 5-6.

Material examined

Types. Holotype, SAM-A40017; paratypes, SAM-A40018, from a tributary

of the Riebeek’s River, above the farm Waterval, Kasteelberg.

Other material. SAM-A40019, from a stream above the farm Wynkelders-

berg, draining the slopes of Kasteelberg.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum about 20- to 30-articulate,

accessory flagellum 3- to 5-articulate. Antenna 2 shorter than 1, moderately

setose, article 3 of peduncle bearing a semicircular lobe posteriorly and article 4

laterally swollen in males, flagellum 13- to 20-articulate. Coxa 4 slightly emar-

ginate posteriorly. Gnathopod 2, article 2 strongly spinose medially, palm

slightly oblique, with 3-5 defining spines. Pereopod 3 moderately setose, modi-

fied in males, article 2 strongly spinose medially, article 4 posterodistally pro-

jected into a large lobe, article 5 short and stout, article 6 bent at right angles to

article 5, dactyl with a single spinule. Pereopods 4-7 sparsely to moderately

setose, unmodified, dactyl with a single spinule. Uropods 1 and 2, peduncle

spinose and setose, rami usually with marginal spines and setae and apical

spines. Uropod 3, inner ramus 0.3 length of outer, apically spinose, sometimes

with a seta, outer ramus with marginal and apical spines, sparsely to moderately

setose, second segment rudimentary or absent. Telson deeply cleft, each lobe

with one spine and 4-5 setae.

Remarks

One of three species with article 3 of antenna 2 lobed and pereopod 3 modi-

fied to form a ‘claw’, Aquadulcaris andronyx is easily distinguished by the

manner in which the claw-like structure is achieved.

Distribution

From streams draining the slopes of Kasteelberg, north of Malmesbury

(Fig. 29).

Aquadulcaris auricularia (Barnard, 1916)

Fig. 4

Gammarus auricularius Barnard, 1916: 209-210, pl. 27 (figs 26-28); 1927: 169-170.

Paramelita auricularis (Barnard) Thurston, 1973: 166. Griffiths, 1981: 82-85, fig. 3A-C.

Paramelita auricularius (Barnard) Stewart & Griffiths, 1992b: 166-179, figs 1-3.

Material examined

Types. Syntypes, SAM-A2599, top of Table Mountain, Cape Peninsula.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Aquadulcaris auricularia, SAM-A2599, syntype, male, 6.7 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2, medial view. E. Pereopod 3. F. Coxa 4.

G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 197

Other material. SAM-A2634, A2962, A3882, A4559, A5907, A40251 and

A40800, all from various localities on the top of Table Mountain. SAM-

A40252, from a stream draining Constantiaberg, Cape Peninsula.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 20- to 25-articulate,

accessory flagellum 3- to 4-articulate. Antenna 2 moderately setose, shorter than

1, article 3 with a posterodistal lobe extending forward to middle of swollen

article 4 in males, flagellum 8- to 13-articulate. Coxa 4, posterior margin trans-

verse to very slightly emarginate. Gnathopod 2, palm transverse, with

1-3 defining spines. Pereopod 3 sparsely to moderately setose, modified in

males, article 4 widened distally, article 5 with a rectangular lobe and a single

tooth-like spine posteriorly, article 6 elongate and arched and folded back

against the lobed posterior margin of 4, dactyl with a single spinule. Pereopod 4

sparsely to moderately setose, unmodified, dactyl with one spinule. Pereo-

pods 5-7 sparsely to moderately setose, dactyl with 1-2 spinules. Uropod 1,

peduncle spinose, sometimes with 1-2 setae, rami subequal, with marginal and

apical spines, inner ramus sometimes with a single seta. Uropod 2, peduncle

spinose and setose, inner ramus slightly longer than outer, both rami with mar-

ginal and apical spines, lacking setae. Uropod 3, inner ramus about 0.3 length

of outer, apically spines, outer ramus with marginal and apical spines, sparsely

setose, second segment rudimentary to absent. Telson deeply cleft, each lobe

with about 6-8 apical setae, lacking spines.

Remarks

This species most closely resembles Aquadulcaris andronyx and A. pher-

onyx, but can be distinguished from these species by the manner in which the

claw-like structure of pereopod 3 is achieved. Article 5 of this pereopod can

vary from having a small projection, to having a large, rounded to rectangular-

shaped lobe posteriorly.

Distribution

Cape Peninsula, in streams draining the upper slopes of Table Mountain in

the north to Constantiaberg in the south (Fig. 29).

Aquadulcaris crassicornis (Barnard, 1916)

Fig. 5

Gammarus crassicornis Barnard, 1916: 207-209, pl. 27 (figs 24-25).

Paramelita crassicornis (Barnard) Thurston, 1973: 166. Griffiths, 1981: 85-86, fig. 3E-G.

Stewart & Griffiths, 1992b: 166-179, figs 1, 4-5.

Material examined

Types. Syntypes, SAM-A3031, from Table Mountain.

Other material. SAM-A3865 and A40220, Grotto Ravine, Table Mountain.

SAM-A3864, Platteklip Gorge, Table Mountain. SAM-A3881, Slangolie,

Table Mountain. SAM-A4367 and A4368, unknown localities on Table

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

y s

ee >

Fig. 5. Aquadulcaris crassicornis, SAM-A40223, male, 6.9 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereo-

pod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 199

Mountain. SAM-A4868, Stinkwater, Table Mountain. SAM-A40222, Blink-

water, Table Mountain. SAM-A40223, Echo Valley, Table Mountain. SAM-

A40225, Rhodes Memorial, Table Mountain. SAM-A40991, Table Mountain,

above Camps Bay. SAM-A41220, Woodhead Reservoir, Table Mountain.

Diagnosis

Eyes white. Antenna | sparsely to moderately setose, flagellum 19- to 25-

articulate, accessory flagellum with 3-5 articles. Antenna 2 moderately setose,

peduncle articles 3 and, to a greater extent, 4, strongly swollen in males, nor-

mal in females, flagellum 10- to 13-articulate. Coxa 4, posterior margin slightly

emarginate. Gnathopod 2, palm transverse to slightly oblique, with 2-4 defining

spines. Pereopod 3, article 5 in males usually with 1-4 stout, tooth-like spines,

dactyl with one spinule. Pereopods 4-7 sparsely to moderately setose, unmod-

ified, dactyl with a single spinule. Uropod 1, peduncle spinose and setose, rami

subequal, with marginal and apical spines, inner ramus usually with a few setae.

Uropod 2, peduncle spinose and setose, inner ramus longer than outer, both

with marginal and apical spines, lacking setae. Uropod 3, inner ramus 0.3

length of outer, apically spinose, outer ramus with marginal and apical spines,

sparsely setose, second segment rudimentary to absent. Telson deeply cleft,

each lobe with a few setae but no spines.

Remarks

The swollen peduncle in antenna 2, poorly emarginate coxa 4, absence of a

second segment on the outer ramus of uropod 3, and the presence of only a

single spinule on the dactyl of each pereopod suggests a strong link between this

species and Aguadulcaris marunguis, A. dentata, A. auricularia, A. andronyx

and possibly Paramelita tulbaghensis.

Distribution

Endemic to streams draining the upper slopes of Table Mountain (Fig. 29).

Aquadulcaris dentata (Stewart & Griffiths, 1992)

Fig. 6

Paramelita dentata Stewart & Griffiths, 1992b: 179-183, figs 7-8.

Material examined

Types. Holotype, SAM-A40244; paratypes, SAM-A40245, from a tributary

of the Sandvlei River on Ou Kaapse Weg, Cape Peninsula.

Other material. SAM-A40249, from a tributary of the Silvermine River,

Cape Peninsula. SAM-A40794, Noordhoek lookout, Silvermine Nature

Reserve.

Diagnosis

Eyes white. Antenna 1, peduncle sparsely setose, flagellum moderately to

densely setose posteriorly, 18- to 21-articulate, accessory flagellum 3- to 5-

articulate. Antenna 2 shorter than 1, sparsely setose, articles 3 and 4 of

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ocal ae ie

Fig. 6. Aquadulcaris dentata, SAM-A40244, holotype, male, 6.9 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereo-

pod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 201

peduncle strongly laterally swollen and enlarged and article 5 with a postero-

distal tooth in adult males, flagellum 9- to 11-articulate. Coxa 4 slightly emar-

ginate posteriorly. Gnathopod 2, palm transverse to slightly oblique, with 3-4

defining spines. Pereopods 3-7 sparsely setose, unmodified, dactyl with a single

spinule. Uropods 1 and 2, peduncles spinose and setose, rami with marginal and

apical spines, lacking setae. Uropod 3, inner ramus 0.3-0.4 length of outer,

apically spinose, lacking setae, outer ramus with marginal and apical spines,

poorly setose, second segment rudimentary to absent. Telson deeply cleft, each

lobe with 1-2 spines and 1-3 setae.

Remarks

Although obviously related to Aquadulcaris crassicornis, A. dentata is dis-

tinguished from it by the presence of a posterodistal tooth on article 5 of the

peduncle of antenna 2. Paramelita spinicornis, with its swollen article 4 of

antenna 2 and occasional tooth on article 5 is also superficially similar, but this

species can be distinguished by the possession of multispinose dactyls, an exca-

vate coxa 4, and the presence of a small, but distinct second article on the outer

ramus of uropod 3.

Distribution

Known from small streams draining the Kalk Bay Mountains, as well as

Chapman’s Peak Mountains above Noordhoek, Cape Peninsula (Fig. 29).

Aquadulcaris marunguis (Stewart & Griffiths, 1992)

Bigs

Paramelita marunguis Stewart & Griffiths, 1992b: 183-187, figs 9-10.

Material examined

Types. Holotype, SAM-A40224; paratypes, SAM-A40246, from a tributary

of the Burgersbos River, Cape Peninsula.

Other material. SAM-A40221, from a tributary of the Disa River, Cape

Peninsula.

Diagnosis

Eyes white. Antenna 1 sparsely to moderately setose, flagellum 24- to 26-

articulate, accessory flagellum 4-articulate. Antenna 2 shorter than 1, sparsely

to moderately setose, articles 3 and 4 laterally swollen and enlarged in adult

males, flagellum with 12-15 articles. Coxa 4, posterior margin weakly emargin-

ate posteriorly. Gnathopod 2, palm transverse and markedly convex, with

3-5 spines. Pereopod 3 moderately to densely setose posteriorly, articles 5 and

6 modified in males, article 5 with two large teeth on posterior margin, article 6

bent backwards against toothed, posterior margin of 5, with two spines, dactyl

with one spinule. Pereopods 4-7 unmodified, moderately setose, dactyl with a

single spinule. Uropods 1 and 2, peduncle spinose and setose, inner rami with

marginal spines and setae and apical spines, outer rami with marginal and apical

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Aquadulcaris marunguis, SAM-A40224, holotype, male, 10.8 mm. A. Lateral

aspect. B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4.

G. Pereopod 6. H. Uropod 1. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 203

spines, lacking setae. Uropod 3, inner ramus 0.3 length of outer, apically

spinose, outer ramus with marginal and apical spines, sparsely setose, second

segment rudimentary. Telson deeply cleft, each lobe with 8-10 setae.

Remarks

This species is very similar to Aquadulcaris crassicornis, from which it is

distinguished by the ‘claw-like’ nature of pereopod 3. In A. marunguis, article 6

of this limb is attached ‘at right angles’ to article 5, whereas in A. crassicornis,

these articles are attached normally.

Distribution

This species has so far been collected from two streams draining the

southernmost parts of Table Mountain, Cape Peninsula (Fig. 29).

Aquadulcaris pheronyx (Stewart & Griffiths, 1992)

Fig. 8

Paramelita pheronyx Stewart & Griffiths, 19925: 187-191, figs 11-12.

Material examined

Types. Holotype, SAM-A40247; paratypes, SAM-A40248, from a stream

draining the slopes of Constantiaberg, Cape Peninsula.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 16- to 21-articulate,

accessory flagellum 3- to 4-articulate. Antenna 2 sparsely setose, shorter than 1,

articles 3 and 4 of peduncle strongly swollen and enlarged in males, article 3

with a large lobe posteriorly, flagellum with 10-14 articles. Coxa 4, posterior

margin slightly emarginate. Gnathopod 2, article 2 strongly spinose medially,

palm transverse, with 3-4 defining spines. Pereopod 3 sparsely setose, article 2

medially setose, articles 4 and 5 modified in males, article 4 short, widening

distally, with a long, narrow posterodistal projection, article 5 elongate and

enlarged, curved, bearing a stout spine at point of attachment with 4, dactyl

with one spinule. Pereopods 4-7 sparsely setose, unmodified, dactyl with a

single spinule. Uropods 1 and 2, peduncle spinose and setose, inner rami with

marginal spines and setae, outer rami with marginal spines, lacking setae, all

rami with apical spines. Uropod 3, inner ramus 0.3-0.4 length of outer, apically

spinose, outer rami with marginal and apical spines, sparsely setose, second

segment rudimentary to absent. Telson deeply cleft, each lobe with 4-5 setae,

lacking spines.

Remarks

The ‘spur-like’ projection on article 4 of pereopod 3 makes A. pheronyx

unmistakable. Aquadulcaris andronyx also has a projection on this article but, in

this species, the projection is wide and triangular-shaped. Both species have a

lobe on article 3 of the peduncle of antenna 2.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

_—_________

Fig. 8. Aquadulcaris pheronyx, SAM-A40247, holotype, male, 7.2 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2, lateral and medial views. D. Gnathopod 2, medial view.

E. Pereopod 3. F. Coxa 4. G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3.

Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 205

Distribution

This species is known only from the type locality, a stream draining the

southern slopes of the Constantiaberg, above the Hout Bay Hotel (Fig. 29).

Mathamelita gen. nov.

Diagnosis

Eyes white. Antenna | shorter than 2, sparsely setose, accessory flagellum

with about four articles. Antenna 2 peduncle not toothed or lobed. Gnathopod 2

palm oblique, excavate. Coxa 4 not excavate posteriorly. Pereopod 3 unmodi-

fied, not subchelate. Pereopods 3-7, dactyls with a single spinule. Pereon 2 with

a single sternal gill, pereon segments 3-7 each with a pair of sternal gills.

Pereopod 7, coxal gill absent. Uropod 3, second segment of outer ramus pres-

ent, inner ramus as long as outer in adult males, reduced in juveniles.

Etymology

This genus is named after the author’s (CLG) young son Matthew. His

request for a comfort stop during a scenic drive in the Outeniqua Mountains

provided his father with the opportunity to explore a roadside stream, leading to

the discovery of the type species.

Type species. Mathamelita aequicaudata sp. nov.

Mathamelita aequicaudata sp. nov.

Figs 9, 10

Etymology

The specific name refers to the equal rami of the third uropod, which is the

most unusual feature of this species.

Material examined

Types. Holotype, SAM-A40990; paratypes, SAM-A41189, from stream

situated between Kruisvallei and Knysna (Fig. 29).

Description of holotype (male, 8 mm)

Body off-white in colour. Head 1.5 length of pereon segment 1, eyes white,

invisible in preserved specimens.

Antenna | about one-third body length, peduncle short, segment 1 50 per

cent longer and broader than 2, 2 twice length of 3, flagellum twice length of

peduncle, 18-articulate, accessory flagellum 4-articulate. Antenna 2 consider-

ably longer and more robust than 1, 60 per cent body length, articles 4 and 5 of

peduncle elongate, not bent or toothed, flagellum 20 per cent longer than ped-

uncle, consisting of 16 broadened segments.

Left mandible with strongly triturative molar and 3-articulate palp, second

article of palp more than three times length of first and slightly longer than

third, which ends in eight long setae. Cutting edge of five strong teeth, lacinia

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Mathamelita aequicaudata sp. nov., SAM-A40990, holotype, male,

6.0 mm. A. Antenna 1, peduncle. B. Lateral aspect. C. Coxa 4. D. Pereopod 3.

E. Pereopod 5. F. Pereopod 6. G. Pereopod 7.

mobilis large and 4-toothed, spine row of one simple and four spine-setae. Right

mandible, cutting edge of five teeth, lacinia mobilis slender and bifurcate, spine

row of two thick and two slender spine-setae.

Maxilla 1, inner plate with four terminal setae, outer plate with 10 strong

serrate spines, palp bi-articulate, reaching beyond outer plate, ending in seven

short spines.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 207

S ~

Fig. 10. Mathamelita aequicaudata sp. nov., SAM-A40990, holotype, male,

6.0 mm. A. Gnathopod 1. B. Gnathopod 2. C. Left mandible. D. Lower lip.

E. Maxilla 1. F. Maxilla 2. G. Maxillipeds. H. Uropod 1. I. Uropod 2.

J. Uropod 3. K. Telson. L. Uropod 3 of juvenile, 2.7 mm.

Maxilla 2 with about 23 hooked setae on inner plate, 21 on outer plate.

Maxilliped, inner plate with about 14 pectinate setae along margin, outer

plate medially lined by 19-20 close-set spine-setae that increase in length

distally, palp 4-articulate.

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pereon segments dorsally smooth, coxae 1-3 quadrate, setose ventrally,

coxa 4 quadrate, not excised posteriorly, coxae 5 and 6 bilobed, 7 semicircular.

A single sausage-shaped sternal gill occurs centrally on pereon segment 2 and a

lateral pair on each of pereon segments 3-7. Coxal gills on pereopods 2-6 only,

absent from pereopod 7.

Gnathopod 1 subchelate, article 2 with long setae posteriorly, article 5 two-

thirds length of 6, 6 rectangular, palm oblique, fairly straight, lined by minute

close-set setae, defined by a slender spine. Gnathopod 2 much larger and more

robust than 1, article 5 much smaller than the broad quadrate 6, palm slightly

concave, defined by three strong spines.

Pereopods 3 and 4 not modified, 5 and 6 about equal, dactyl with a single

spinule. Pereopod 5 short, only about 25 per cent body length, article 2 moder-

ately expanded posteriorly, spinose anteriorly, dactyl with a single spinule.

Pereopods 6 and 7 each progressively longer, article 2 only slightly lobed

posteriorly, spinose anteriorly, not strongly tapering, distal articles strongly

spinose, but with few setae, dactyl with a single spinule.

Pleon segments sparsely setose dorsally, pleonal epimera rounded, with

groups of spines along ventral and posterior margins. Uropod 1 reaching to tip

of 2, peduncle with a row of seven spines on dorsal margin, rami equal, two-

thirds length of peduncle, strongly spinose terminally. Uropod 2 two-thirds

length of 1, rami equal. Uropod 3 reaching beyond 2 by full length of its rami,

length about 11 per cent of body, peduncle quadrate, outer ramus twice length

of peduncle, rectangular, with three clusters of spines on lower edge, five on

upper, minute second article ends in one spine and two setae. Inner ramus equal

in length to outer but more slender, tapering and without lateral spines, five

spines at apex.

Telson 60 per cent cleft, apex rounded, each lobe with a single, short apical

spine.

Variation

The inner ramus of uropod 3 appears to start life small and tapering, and

increases in size relative to the outer ramus later in life. In a juvenile of 2.7 mm

(Fig. 10L), the inner ramus is triangular and only one-third the length of the

outer—not dissimilar to that of Paramelita species but, by 4 mm, males had the

inner ramus 85 per cent as long as the outer.

In females, antenna 2 is much shorter than that of males, reaching only

about 90 per cent of the length of antenna |. Gnathopod 2 is also much smaller

than that of males, being similar in shape and structure to gnathopod 1. Uro-

pod 3 of females has unequal rami, the inner ramus being tapering and about

50 per cent of the length of the outer.

Paramelita Schellenberg, 1926

Diagnosis

Eyes white or black. Antenna | 0.7-1.8 length of 2, peduncle sparsely

setose, article 1 1.1-1.6 length of 2, 2.6-3.7 longer than wide, flagellum

sparsely setose, 18- to 80-articulate, accessory flagellum 3- to 8-articulate.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 209

Antenna 2, sparsely to densely setose, rarely toothed or lobed, peduncle either

shorter or longer than flagellum, often stout in males, article 4 2.1-4.5 longer

than wide, flagellum 11- to 35-articulate. Gnathopod 2, article 2 medially

spinose or not, palm slightly to strongly oblique. Pereopod 3 usually unmodi-

fied, rarely subchelate. Coxa 4, posterior margin slightly emarginate to strongly

excavate. Pereopods 3 and 4, dactyl usually with 2-8 spinules. Pereopods 5-7,

dactyl usually with 3-14 spinules. Pereon segments 2-7 with 1-4 sausage-

shaped sternal gills, coxal gill present on pereopod 7. Uropod 3, inner ramus

0.1-0.4 length of outer, second segment on outer ramus present or absent.

Type species. Paramelita ctenodactyla Schellenberg, 1926: 367-370,

fig. 57 (= Paramelita capensis (Barnard, 1916)).

KEY TO SPECIES OF PARAMELITA

IPP CSU AC Keres oP otters ciate erasers acteristics aaiare Asim asic siosislSn ome alesis Bae eee ULE 2

ESE! VCSHWUN CO ec chan oe enc need eee cate cyte tune eos Gene como cue ate atiasaer | eet ORT areet 3

2A. Antenna 2, posterior margins and pereopods 3-7 densely setose

DORSEY Mancugascccossosnaseeneoncaners meen nants P. nigroculus var. persetosa

2B. Antenna 2, posterior margins and pereopods 3-7 sparsely to moderately

setose posteriorly. lacking setalsbrushes: <...2---..2:-04-05e50 2 P. nigroculus

3A. Antenna 2, peduncle either toothed, lobed or ridged (Figs 14C, 20C,

DCR OE) rs res reeset sce a et hck Mee ed Senter eee om REET 4

3B. Antenna 2, peduncle lacking teeth, lobes or ridges (e.g. Fig. 11C) ...... vi

4A. Antenna 2, articles 4 or 5 of peduncle toothed (Figs 20C, 26C) .......... 5

4B. Antenna 2, article 3 posteriorly lobed (Fig. 14C) or article 5 with lateral

YORE (CEO IIEY 8) can ais 4 ah ae a HBR een ai aan, PR. Re ener NEN So BO 6

5A. Antenna 2 shorter than 1, article 4 of peduncle strongly laterally swollen,

with a posterodistal, terminal tooth and a proximal, medial lobe

GEO OAD) eae ieee tica te eee Aas ONG ron euceved seen Cae ms P. spinicornis

5B. Antenna 2 extremely elongate, exceeding | in length, article 4 of ped-

uncle with a posterodistal, subterminal tooth (Fig. 20A) . P. odontophora

6A. Antenna 2, article 3 of peduncle posteriorly lobed, pereopod 3 normal

GReTL Awe ements ase cies oc eran se eee tins a niaiore ere ecclesinaetee ane P. flexa

6B. Antenna 2, article 5 of peduncle with lateral ridges, pereopod 3 sub-

HCL ALC ALS ae) 5 AN) eam rie ch sre eee eee oe ce les coe nese P. pinnicornis

ae eAsteniias densely setose (Figs 226. 25 ©) yack om, wclens yen eeis ee seenee ae 8

7B Antenna 2 spatsely to moderately setose (e.g. Figs 11€) -.. 2... 9

8A. Antenna 2, peduncle stout, flagellum shorter than peduncle, with 8-12

articles; pereopods 5-7, article 2 moderately expanded; uropod 1, outer

ramus lacking setae; uropod 2, inner ramus usually with some setae

GE aD IAL) eo ace I eaten P. seticornis

210

8B.

9A.

OB.

10A.

10B.

LIA.

11B.

12A.

12B.

13A.

13B.

14A.

14B.

ISA.

SB:

16A.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Antenna 2, peduncle elongate and slender, flagellum as long as peduncle,

with 13-16 articles; pereopods 5-7, article 2 markedly poorly expanded;

uropod 1, outer ramus with some setae; uropod 2, inner ramus lacking

Setae(h1gi22 Amie s x oie eeateay oa eisetee meen evade eee eee P. pillicornis

Coxa 4, posterior margin poorly emarginate (Figs 11F, 15A, 24F, 27F) .

Fen aOR RET RRO RAT OME re HUNG 5o 500000000000 10

MOE oe eiitece suaaisd ae doing alicia s audbreiel oe-aelnioeley cts calcio Se eee eee 13

Gnathopod 2, palm moderately convex, lacking tooth at defining angle;

pereopod 3, article 4 only moderately longer and wider than 5 (Figs 11A,

Po EN) Naded aa aE ROC es PEI oe CME oe P NGA Nim Sido bev cov0GeD 5000 ih

Gnathopod 2, palm strongly convex, defining angle forming a small

projecting rounded tooth; pereopod 3, article 4 often considerably longer

and wider than 3 (Fig. 24B). 30.3 ccc ccc scatoscoenssce ses cect ete eee eRe EeEe 12

Antenna 2, peduncle laterally swollen, flagellum 9- to 12-articulate

CRISS DIC) eo tetcn de tate eee ceatane amore sh ae renee ts eee a ieme P. tulbaghensis

Antenna 2, peduncle not markedly laterally swollen, flagellum 12- to 13-

articulate (Fig. TG) Wess ven coccn deca hona. tee denies ote een eee eee P. aurantia

Pereopod 3, article 4 greatly expanded laterally; uropod 3, inner ramus

about/02 Jencthiof outer ramus (Rig. 242A) se... teen eae P. platypus

Pereopod 3, article 4 not markedly expanded laterally; uropod 3, inner

ramus about 0.4 length of outer ramus (Fig. 15A) ........ P. granulicornis

Antenna 2 as long as, or exceeding | in length, peduncle markedly stout

(Bigs 7; 18C 28 Cr css s oc Geese ct aes cae sos cieine eee eee 14

Antenna 2, distinctly shorter than 1, peduncle slender to moderately stout

(es Figs I2€, 13 CG) se ss seve ierace ten eachore cos tsenena te aa nee eee 16

Uropod 3, outer ramus 3.0 length of peduncle; uropods 1 and 2, inner

rami always with a few setae, outer rami lacking setae (Fig. 28A) .........

Bite class fe Gata eaie Ute arn nee cua Serie ear tcea nen een an eer P. validicornis

Uropod 3, outer ramus 2.0-2.6 length of peduncle; uropods 1 and 2,

inner rami with or without setae, outer rami sometimes with setae

(Bigs. 7H=J,A8SGED) « essusce cas cas ste asteeoasene stele aeaeea des eaae ee oe eee 15

Pereopod 3, article 4 unmodified; urosome densely setose dorsally; uro-

pods 1 and 2, inner and outer rami with setae; body colour brown

CIS TTA ccs cet aiare Sku lasagne so aSeicidion sake w delecanuseeee ae Oe ea see tee P. magna

Pereopod 3, article 4 posterodistally protruded to form a ‘tooth’; urosome

moderately setose dorsally; uropod 1, inner ramus with a few setae,

outer ramus without; uropod 2, rami lacking setae; body colour white

(Rip BIB AY ee ccs ences eed nec sees aecee snence eee eee P. magnicornis

Antenna 1, flagellum with 22-27 articles; antenna 2, flagellum 11- to 18-

articulate (Figs 16B-—C,21B-€)) 2.03.2. sccwet onder oes tee ee eee 17

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 211

16B. Antenna 1, flagellum with 33-80 articles; antenna 2, flagellum 15- to 35-

articulate (Rigs) (2B —C. Al SIB=C)) si ecc sate cts ons cate abiosinteons bein tone pawns « 18

17A. Pereopods 3 and 4, dactyl with 2-3 spinules; uropod 2, inner ramus lack-

ing marginal setae; uropod 3, outer ramus moderately to densely setose

CEI A) eas noe tcnctee ah cece eticace ante edd s Pie aeeecide wane seems nace P. parva

17B. Pereopods 3 and 4, dactyl with four spinules; uropod 2, inner ramus with

a few marginal setae; uropod 3, outer ramus, poorly setose (Fig. 16A) ...

SBC Ie Aco ARCO a SG SUES SEES CS RSE ie P. kogelensis

18A. Antenna 2, flagellum with 15-17 articles; pereopods 3 and 4, dactyl with

2-3 spinules; pereopods 5-7, dactyl with 5-7 spinules; coxa 4 distinctly,

but moderately excavate posteriorly; uropods 1 and 2, rami lacking setae;

uropod 3, outer ramus poorly setose (Fig. 12A) ................ P. barnardi

18B. Antenna 2, flagellum usually with more than 17 articles; pereopods 3 and

4, dactyl with 3-6 spinules; pereopods 5-7, dactyl with 8-13 spinules;

uropods 1 and 2, inner rami with marginal setae; uropod 3, outer ramus,

Sion elyesctOSex (ell ewla A! eens cane ae ae eee Oe aee P. capensis

Paramelita aurantia (Barnard, 1927)

Fig. 11

Gammarus aurantius Barnard, 1927: 173-174, pl. 10 (figs 6, 16).

Paramelita aurantius (Barnard) Thurston, 1973: 167. Griffiths, 1981: 82, fig. 2J.

Material examined

Types. Syntypes, SAM-A3997, from Landdrost Kloof, Hottentots Holland

Mountains.

Other material. SAM-A4005, from valley at foot of Vallei Berg, Hottentots

Holland Mountains. SAM-—A4014, from Moordenaars Kop, Hottentots Holland

Mountains. SAM-A4869, from Caledon side of Landdrost Kloof, Hottentots

Holland Mountains. SAM-A40234, from a tributary of the Du Toit’s River on

Franchhoek Pass, Villiersdorp side. SAM-A40235, from a stream flowing

through the Nuweberg State Forest on Viljoen’s Pass.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum about 20- to 32-articulate,

accessory flagellum 3- to 4-articulate. Antenna 2 sparsely to moderately setose,

shorter than 1, only slightly stouter than 1 in males, flagellum with about 12-13

articles. Coxa 4, posterior margin with a slight emargination. Gnathopod 2,

article 2 sometimes weakly spinose medially, palm transverse to slightly

oblique, with 2-5 defining spines. Pereopods 3-7 unmodified, dactyl with 2-10

spinules. Uropod 1, peduncle spinose, sometimes with a few setae, rami

subequal, with marginal and apical! spines, lacking setae. Uropod 2, peduncle

spinose, sometimes bearing at least one seta, inner ramus slightly longer than

outer ramus, both with marginal and apical spines, lacking setae. Uropod 3,

inner ramus about 0.3 length of outer, apically spinose, outer ramus with

212 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. Paramelita aurantia, SAM-A3997, syntype, male, 6.8 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereo-

pod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. K. Telson. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 213

marginal and apical spines, sparsely setose, second segment rudimentary.

Telson deeply cleft, each lobe bearing one spine and a few setae.

Remarks

This species is identified by the possession of an unmodified, sparsely to

moderately setose, and relatively short antenna 2 in males, an almost quadrate

coxa 4, the relative lack of setae on uropods | and 2, and its orange colour

when alive. It most closely resembles Paramelita granulicornis, but can be

distinguished from this species by the lack of a tooth on the palm of gnatho-

pod 2, and the absence of spines on the medial surface of article 2 of

pereopods 3 and 4.

Distribution

From streams draining the slopes of Hottentots Holland and Franchhoek

mountains (Fig. 29).

Paramelita barnardi Thurston, 1973

Fig. 12

Paramelita barnardi Thurston, 1973: 159-168, figs 1-3. Griffiths, 1981: 85, fig. 2A-C.

Material examined

Types. Allotype, SAM-A16808, from Boomslang Cave, Kalk Bay

Mountains.

Other material. SAM-A40239, from Boomslang Cave, Kalk Bay Moun-

tains. SAM-A40796, Avernus Crack, Cave Peak, Kalk Bay.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 33- to 36-articulate,

accessory flagellum with 4-5 articles. Antenna 2 shorter than 1, moderately

setose, flagellum with 15-17 articles. Coxa 4 distinctly but shallowly excavate

posteriorly. Gnathopod 2, palm oblique, with 4-5 defining spines. Pereopods 3

and 4 moderately setose, unmodified, dactyl with 2-3 spinules. Pereopods 5-7

moderately setose, dactyl with 5-7 spinules. Uropod 1, peduncle spinose, some-

times with a single seta, rami subequal, with marginal and apical spines, lacking

setae. Uropod 2, peduncle spinose, inner ramus slightly longer than outer, both

rami with marginal and apical spines, lacking setae. Uropod 3, inner ramus

about 0.4 length of outer, apically spinose, outer ramus very poorly setose, with

several groups of marginal and apical spines, second segment short, about 4 per

cent length of first. Telson deeply cleft, left lobe bearing a single spine and

some setae, right lobe with 1-2 spines and some setae.

Remarks

This species is morphologically similar to Paramelita capensis, P. kogel-

ensis and P. parva, but can be separated from these species based on its rela-

tively weakly excavate coxa 4, its poorly setose uropod 3, and the number of

articles in the flagellum of antenna 2.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

— —

Fig. 12. Paramelita barnardi, SAM-A40239, male, 9.5 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereo-

pod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. K. Telson. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 215

Distribution

Known only from caves in the Kalk Bay Mountains, Cape Peninsula

(Fig. 29).

Paramelita capensis (Barnard, 1916)

Fig. 13

Gammarus capensis Barnard, 1916: 203-205, pl. 27 (figs 20-22) (part., non SAM-A3083);

1927: 169.

Paramelita ctenodactyla Schellenberg, 1926: 367, fig. 57.

Paramelita capensis (Barnard) Griffiths, 1981: 85, fig. 4. Stewart, 1992: 288.

Material examined

Types. Syntypes, SAM-A2259, from Table Mountain.

Other material. SAM-A195, A2258, A2459, A2552, A2598, A2963,

A2967, A2968, A3033, A3866, A4008 and A41009, all from various localities

on Table Mountain. SAM-A2960, from Muizenberg Mountains. SAM-A4565,

from Hout Bay. SAM-A6604, from the Cedarberg. SAM-A7328, from Noord-

hoek forest. SAM-A40242, Echo Valley, Table Mountain. SAM-A40524,

Bokkeman’s Kloof, Hout Bay. SAM-A40532 and A41006, Constantiaberg.

SAM-A40813, The Baths, Citrusdal. SAM-A40814, Grotto Ravine, Table

Mountain. SAM-A40815, Blackburn Ravine, Hout Bay. SAM-A40817, Platte-

klip Gorge, Table Mountain. SAM-A40818, tributary of the Spansemat River,

Cape Peninsula. SAM-A41221, source of Disa River, above Hout Bay.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum approximately 40- to 80-

articulate, accessory flagellum 5- to 8-articulate. Antenna 2 usually shorter than

1, moderately setose, peduncle long and slender, stouter than 1, flagellum with

about 15-35 articles. Coxa 4 strongly excavate posteriorly. Gnathopod 2,

article 2 either with long spine-like setae, or with stout spines on medial,

posterior margin, palm strongly oblique, with 3-5 defining spines. Pereopods 3

and 4 moderately setose, unmodified, article 2 usually either with spine-like

setae or strong spines medially, dactyl with 3-6 spinules. Pereopods 5-7

moderately setose, dactyl with 8-13 spinules. Uropod 1, peduncle spinose, with

very few setae, rami subequal, inner ramus with marginal spines and setae,

outer ramus with marginal spines, rarely with setae, both ending in apical

spines. Uropod 2, peduncle spinose, inner ramus slightly longer than outer,

both with marginal and apical spines and marginal setae. Uropod 3, inner ramus

0.2 length of outer, apically spinose, outer ramus with marginal and apical

spines, strongly setose, distinct second segment. Telson deeply cleft, each lobe

bearing one spine and many apical and subapical setae.

Remarks

This species closely resembles Paramelita barnardi, P. kogelensis and

P. parva, but is distinguished from these species by its large size at maturity,

strongly oblique palm of gnathopod 2, deeply excavate coxa 4, and densely

setose uropod 3.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Paramelita capensis, SAM-A2259, syntype, male, 16.3 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereo-

pod 5. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE

ZAG

—

ay ee oe

Fig. 14. Paramelita flexa, SAM-A40782, male, 7.9 mm. A. Lateral aspect. B. Antenna 1.

C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereopod 6.

H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Apparently widespread, collected from streams in the Cedarberg area in the

north, to the Cape Peninsula in the south (Fig. 29).

Paramelita flexa Griffiths, 1981

Fig. 14

Paramelita flexa Griffiths, 1981: 86-89, fig. 5.

Material examined

Types. Holotype, Albany Museum MISC 52B, from a tributary of the Pal-

miet River between Elgin and Grabouw. Paratypes, SAM-A16776, from the

same locality as the holotype.

Other material. SAM-A40782 and A40788, Highlands State Forest,

Kleinmond.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum about 26-articulate, access-

ory flagellum 5-articulate. Antenna 2 shorter than 1, in males, article 3 strongly

lobed posterodistally, article 4 curved ventrally and article 5 bent at right angles

to 4, flagellum 16-articulate. Coxa 4 distinctly excavate posteriorly. Gnatho-

pod 2, palm slightly oblique, with three defining spines. Pereopods 3 and 4

unmodified, dactyl with 3-4 spinules. Pereopods 5-7, dactyl with 4-9 spinules.

Uropod 1, peduncle spinose, lacking setae, rami subequal, with marginal and

apical spines, no setae. Uropod 2, peduncle spinose and setose, inner ramus

slightly longer than outer, both with marginal and apical spines, lacking setae.

Uropod 3, inner ramus 0.3 length of outer, with some spines on apex, outer

ramus with marginal and apical spines and setae, second segment distinct.

Telson deeply cleft, each lobe with one spine and some setae.

Remarks

The combination of a protruded posterior margin in article 3 of antenna 2,

an excavate coxa 4, multispinose dactyls and the presence of a second segment

on the outer ramus of uropod 3 makes this species distinctive. It is highly

unlikely that the ‘lobe’ on article 3 of antenna 2 is homologous to the semi-

circular lobe found in Aquadulcaris auricularia, A. andronyx and A. pheronyx.

Distribution

Known from a stream flowing through the Highlands State Forest and also

from a tributary of the Palmiet River on the Grabouw-Elgin road, between

Hottentots Holland and Groenland mountains (Fig. 29).

Paramelita granulicornis (Barnard, 1927)

Fiss5

Gammarus granulicornis Barnard, 1927: 175-177, pl. 10 (figs 10-11, 20).

Paramelita granulicornis (Barnard) Thurston, 1973: 167. Griffiths, 1981: 89, fig. 2H-I.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE

Fig. 15. Paramelita granulicornis, SAM-A40236, male, 10.3 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3.

F. Pereopod 6.

G. Uropod 1. H. Uropod 2. I. Uropod 3. Scale lines represent 1 mm.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material examined

Types. Syntypes, SAM-A4874 and A5178, from Steenbras River.

Other material. SAM-A5182, A5183 and A5185, all from the Hottentots

Holland Mountains. SAM-A40236, from a stream on Houhoek Pass. SAM-

A40237, from a tributary of the Palmiet River flowing near the Orchard’s Farm

Stall. SAM-A40238 and A40787, from a tributary of the Palmiet River below

Elephant Rock. SAM-A40781 and A40786, Grabouw Forest. SAM-A40779,

between Kleinmond and Betty’s Bay. SAM-A40945, Gordon’s Bay.

Diagnosis

Eyes white. Antenna 1, sparsely setose, flagellum about 27- to 40-articulate,

accessory flagellum 4- to 5-articulate. Antenna 2 sparsely to moderately setose,

shorter than 1, moderately stout, peduncle sometimes elongate in males, flagel-

lum with about 12-16 articles. Coxa 4 quadrate. Gnathopod 2, article 2

posteriorly spinose, palm transverse, defining angle forming a small protruding

rounded tooth, with three spines. Pereopods 3 and 4, article 2 strongly spinose

posteriorly, article 4 often considerably longer and wider than 5, dactyl with

2-3 spinules. Pereopods 5-7, dactyl with 4-8 spinules. Uropod 1, peduncle

spinose, lacking setae, rami subequal, bearing marginal and apical spines,

lacking setae. Uropod 2, peduncle with spines and usually at least one seta,

inner ramus longer than outer, both with marginal and apical spines, lacking

setae. Uropod 3, inner ramus about 0.4 length of outer ramus, with some apical

setae and at least one seta, outer ramus with marginal and apical spines and

setae, second segment rudimentary. Telson deeply cleft, each lobe bearing one

spine and several setae.

Remarks

This species is most like Paramelita aurantia, from which it is distinguished

by the possession of a strongly convex palm with palmar tooth in gnathopod 2.

Paramelita granulicornis shares this condition with P. platypus, and it is poss-

ible that these two species are closely related. Both species have almost quad-

rate fourth coxal plates, and Barnard (1927: 176) commented on how article 4

in pereopods 3 and 4 is ‘strongly expanded distally’, and article 5 is ‘noticeably

shorter’ than article 4 in P. granulicornis. This condition is extremely well

developed in P. platypus. Any further decisions regarding the position of

P. granulicornis will be taken once genetic analysis of Paramelita is complete.

Distribution

Known from the Hottentots Holland Mountains and adjacent areas (Fig. 29).

Paramelita kogelensis (Barnard, 1927)

Fig. 16

Gammarus kogelensis Barnard, 1927: 172-173, pl. 10 (figs 9, 21).

Paramelita kogelensis (Barnard) Thurston, 1973: 167. Griffiths, 1981: 89, fig. 2G.

Material examined

Types. Syntypes, SAM-A4873, west of Kogelberg.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 221

Fig. 16. Paramelita kogelensis, SAM-A40243, male, 8.0 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Pereopod 6.

G. Uropod 1. H. Uropod 2. I. Uropod 3. Scale lines represent 1 mm.

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

Other material. SAM-A5174, Kogelberg. SAM-A5190, on way to Kogel-

berg from Steenbras. SAM-A40243, Viljoen’s Pass, Nuweberg State Forest.

SAM-A40529, Steenbras River.

Diagnosis

Eyes white. Antenna 1 sparsely setose, peduncle sometimes spinose, flagel-

lum 27- to 34-articulate, accessory flagellum 3- to 5-articulate. Antenna 2

shorter than 1, sparsely to densely setose, flagellum 12- to 16-articulate.

Coxa 4, posterior margin, distinctly excavate. Gnathopod 2, article 2 not

medially spinose, palm slightly to moderately oblique, with 2-3 defining spines.

Pereopods 3 and 4 unmodified, dactyl with 2-4 spinules. Pereopods 5-7, dactyl

with 5-8 spinules. Uropod 1, peduncle spinose, lacking setae, rami subequal,

with marginal and apical spines, inner ramus rarely with setae. Uropod 2,

peduncle spinose, rarely with one seta, inner ramus slightly longer than outer,

both with marginal and apical spines, inner ramus rarely with 1-2 setae.

Uropod 3, inner ramus 0.2-0.3 length of outer, apically spinose and sometimes

with 1-2 setae, outer ramus with marginal and apical spines, sparsely setose,

distinct second segment present. Telson deeply cleft, each lobe with one spine

and 3-4 setae.

Remarks

One of four morphologically similar species, Paramelita kogelensis is

distinguished from P. parva, P. barnardi and P. capensis by the number of

articles in the flagella of antennae | and 2, spination of the dactyl of pereo-

pods 3 and 4, and the setation of the inner rami of uropod | and the outer ramus

of uropod 3.

Distribution

Collected from localities on the Hottentots Holland and adjacent mountains

(Fig. 29).

Paramelita magna Stewart & Griffiths, 1992

Fig. 17

Paramelita magna Stewart & Griffiths, 1992c: 491-494, figs 2-3.

Material examined

Types. Holotype, SAM-A40208; paratypes, SAM-A40209, from a tributary

of the Krom River in the Cape of Good Hope Nature Reserve.

Other material. SAM-A3083, Kalk Bay (= Barnard’s 1916: 205, SAM=

A3084— typographical error). SAM-A4563, from Noordhoek. SAM-A40210,

from the Booiskraal River, and SAM-A40211 and A41011, from the Buffels

River, both in the Cape of Good Hope Nature Reserve. SAM-A40212, from

Nellies Pool, and SAM-A40213 and A40798, from the Silvermine River, both

in the Silvermine Nature Reserve. SAM-A40515, Klassjagers River, Cape

Point. SAM-—A40930 and A40932, De Goede Hoop Estate, Noordhoek (latter

incorrectly assigned to A40214 in Stewart & Griffiths (1992c)).

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 223

Fig. 17. Paramelita magna, SAM-A40208, holotype, male, 22.3 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2, medial view. E. Pereopod 3. F. Coxa 4.

G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

Eyes white. Antenna 1 sparsely setose, flagellum 37- to 42-articulate,

accessory flagellum 6- to 8-articulate. Antenna 2 sparsely to moderately setose,

in males, peduncle stout and elongate so that antenna 2 equal to, or exceeding

antenna | in length, flagellum with 16-19 articles. Coxa 4, posterior margin

excavate. Gnathopod 2, medial posterior margin of article 2 strongly spinose in

males, palm slightly oblique, defined by five spines. Pereopods 3 and 4 unmodi-

fied, moderately to densely setose posteriorly, dactyl with 4-6 spinules.

Pereopods 5-7 strongly setose anteriorly, dactyl with 8-10 spinules. Uropod 1,

peduncle spinose, usually lacking setae, rami subequal, with marginal spines

and setae and apical spines. Uropod 2, peduncle spinose, sometimes with a few

setae, inner ramus longer than outer, both with marginal spines and setae and

apical spines. Uropod 3, inner ramus 0.3 length of outer, apex spinose and

setose, outer ramus with marginal and apical spines and setae, second segment

distinct, about 6 per cent length of first. Telson deeply cleft, each lobe usually

with one spine and many setae, right lobe sometimes with two spines.

Remarks

Some of the largest specimens of Paramelita collected are members of this

species. Paramelita magna is easily recognized by its dark brown colour,

markedly setose urosome and pereopods, and the possession of stout elongate

second antennae in males. It is distinguished from P. validicornis by the relative

length of the peduncle and outer ramus in uropod 3, and from P. magnicornis

by the setation of the uropods, body colour, and the lack of modification of

pereopod 3.

Distribution

In streams draining mountainous areas in the southern part of the Cape

Peninsula (Fig. 29).

Paramelita magnicornis Stewart & Griffiths, 1992

Fig. 18

Paramelita magnicornis Stewart & Griffiths, 1992a: 144-148, figs 3-4.

Material examined

Types. Holotype, SAM-A40009; paratypes, SAM-A40010, from a stream

draining the Swartkop Mountains near Miller’s Point.

Other material. SAM-A40011 and SAM-A40015, from a stream draining

Chapman’s Peak. SAM-A40012, from a stream in the Kalk Bay Mountains near

Clovelly. SAM-A40013, from a stream near Miller’s Point. SAM-A40014,

A40016, and A40940, from Peck’s Valley on Boyes Drive. SAM-A40773,

Swellendam Mountains.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 27- to 40-articulate,

accessory flagellum 4- to 6-articulate. Antenna 2 sparsely to moderately setose,

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 225

Fig. 18. Paramelita magnicornis, SAM-A40009, holotype, male, 15.0 mm. A. Lateral

aspect. B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Pereopod 7.

G. Uropod 1. H. Uropod 2. I. Uropod 3. J. Telson. Scale lines represent 1 mm.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

in males, articles 3, 4 and 5 of peduncle elongate and stout, with articles 4 and

5 distally swollen, antenna 2 exceeding 1 in length, flagellum 13- to 21-

articulate. Coxa 4, posterior margin excavate. Gnathopod 2, article 2 spinose on

medial posterior margin in males, palm oblique, defined by 3-4 stout spines.

Pereopods 3 and 4 moderately to densely setose posteriorly, in males, article 4

posterodistally protruded into a triangular tooth, dactyl with 4-6 spinules.

Pereopods 5-7, moderately to densely setose, dactyl with 6-9 spinules. Uro-

pod 1, peduncle spinose and setose, rami subequal, both with marginal and

apical spines, inner ramus with a few setae. Uropod 2, peduncle spinose and

setose, inner ramus slightly longer than outer, both with marginal and apical

spines, inner ramus sometimes with a few setae, outer ramus lacking setae.

Uropod 3, inner ramus 0.3 length of outer, apex spinose, outer ramus with

marginal and apical spines and setae, second segment distinct but small, about

5 per cent length of first. Telson deeply cleft, each lobe with 1-2 spines and

many setae.

Remarks

This species is usually distinguished by the elongation of antenna 2, and the

posterodistal projection of article 4 in pereopods 3 and 4. Populations in which

this ‘tooth’ on article 4 is absent or poorly developed are distinguished from

Paramelita magna by setation of the uropods and body colour and size.

Distribution

Originally thought to be confined to the Cape Peninsula, this species has

recently been discovered in the Swellendam Mountains (Fig. 29).

Paramelita nigroculus (Barnard, 1916)

Fig. 19

Gammarus nigroculus Barnard, 1916: 206-207, pl. 27 (fig. 23); 1927: 168-169.

Paramelita nigroculus (Barnard) Thurston, 1973: 166. Griffiths, 1981: 89-90, fig. 6.

Paramelita nigroculus var. persetosus Barnard, 1927: 168-169.

Material examined

Types. Syntypes, SAM-A3059, from a stream above Oranjezicht, Table

Mountain. Syntypes, Paramelita nigroculus var. persetosa, SAM-A4877, from

Sneeuwgat near Tulbagh.

Other material. SAM-A1270, Devil’s Peak, Table Mountain. SAM-A2461

and A4009, Platteklip Gorge, Table Mountain. SAM-A2966 and A3060-

A3062, Table Mountain. SAM-A3038, Kirstenbosch. SAM-A4002, north of

Landdroskloof, Caledon side. SAM-A4016 and A4871, Steenbras River.

SAM-A4560, Jonkershoek, opposite Diep Gat. SAM-A4876, Tulbagh.

SAM-A4878-A4884, all from localities near Sneeuwgat, north of Tulbagh.

SAM-A4885, Franchhoek Mountains. SAM-A4887, Vlakte, Ceres. SAM-—

A5188, Hottentots Holland Moutains. SAM-A6054, A6055 and A8273, Zon-

derend Mountains. SAM-A6296, Montagu. SAM-A6602, Middelberg plateau,

Cedarberg. SAM-A6603 and A6965, Tafelberg, Cedarberg. SAM-A6936-

A6938, Swellendam Mountains. SAM-A6944, Simonsberg. SAM-A6945,

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 227

Witte River. SAM-A7335, Schuiffenberge, east of Citrusdal. SAM-A8196,

Krom River, Cedarberg. SAM-A12308, Porterville. SAM-A40264, A40266

and A40271, Du Toit’s Kloof. SAM-A40267, Bain’s Kloof Pass. SAM-

A40269, Steenboks Nature Reserve. SAM-A40270, Paarl Rocks. SAM-

A40273, Dwarsrivierhoek, near Stellenbosch. SAM-A40274 and A40941,

Wemmershoekdam. SAM-A40275, Franchhoek Pass. SAM-A40519, A40522

and A40937, Cedarberg Mountains. SAM-A40520, A40790, A40791 and

A40792, Piketberg. SAM-A40523, Tradouw Pass, Barrydale. SAM-A40527,

Grey’s Pass. SAM-A40533 and A41008, Kasteelberg. SAM-A40774 and

A40793, Grootvadersbos, east of Swellendam. SAM-A40776, between Klein-

mond and Betty’s Bay. SAM-A40926 and A40943, Mitchell’s Pass, Ceres.

SAM-A40931, Keyers River, Cape Peninsula. SAM-A40938, Winterhoek

Mountains, Tulbagh. SAM-—A40944, Swellendam Mountains.

Diagnosis

Eyes black. Antenna 1 sparsely setose, flagellum 20- to 70-articulate,

accessory flagellum 4- to 5-articulate. Antenna 2 shorter than 1, sparsely to

densely setose, slender to stout, flagellum 15- to 22-articulate. Coxa 4, posterior

margin strongly excavate. Gnathopod 2, article 2 not medially spinose,

articles 5 and 6 markedly elongate or not, palm slightly oblique, with 2-4

defining spines. Pereopods 3 and 4 unmodified, moderately to densely setose,

dactyl with 3-7 spinules. Pereopods 5-7 moderately to densely setose, dactyl

with 5-10 spinules. Uropod 1, peduncle spinose and setose, rami subequal, both

with marginal and apical spines, inner ramus setose or not. Uropod 2, peduncle

spinose and setose, both rami with marginal and apical spines, inner ramus

setose or not. Uropod 3, inner ramus 0.2-0.3 length of outer, apically spinose,

margins and apex setose, outer ramus with marginal and apical spines, sparsely

to densely setose, second segment present or not. Telson deeply cleft, each lobe

with one spine and many apical and dorsal setae.

Remarks

All populations of Paramelita that have black eyes have been considered

members of a single widespread species, P. nigroculus, despite morphological

variation between them. For example, some populations have individuals with

elongate, stout second antennae, whereas in others these antennae are relatively

slender and short. Barnard (1927) recognized a variety, P. nigroculus var.

persetosa, based mainly on specimens from the Sneeuwgat valley north of

Tulbagh. These animals have densely setose second antennae, pereopods and

uropods. Setation in P. nigroculus can vary considerably between populations as

well as within one population, and is usually related to maturity, with the larger

older individuals more setose than younger specimens. Therefore, Barnard

(1927) was reluctant to consider these populations with highly setose second

antennae aS a Separate species. There is a need for a thorough, morphological

and genetic investigation of all black-eyed Paramelita populations.

Distribution

Widely distributed from the Cedarberg in the north to Swellendam in the

east (Fig. 29).

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

St pM

Fig. 19. Paramelita nigroculus, SAM-A8273, male, 18.0 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4. G. Pereo-

pod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 229

Fig. 20. Paramelita odontophora, SAM-A40241, paratype, male, 11.1 mm. A. Lateral

aspect. B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4.

G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paramelita odontophora Stewart, Snaddon & Griffiths, 1994

Fig. 20

Paramelita odontophora Stewart et al., 1994: 191-194, figs 5-6.

Material examined

Types. SAM-A40240, holotype, SAM-A40241; paratypes, from a tributary

of the Palmiet River, near Elgin.

Other material. SAM-A40250, tributary of the Palmiet River, near Klein-

mond. SAM-A40783, A40939, tributary of the Palmiet River, near Elgin.

SAM-A40936, Grabouw Forest.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 35- to 38-articulate,

accessory flagellum 4- to 5-articulate. Antenna 2 sparsely setose, in males, both

peduncle and flagellum extremely elongate so that antenna 2 is considerably

longer than 1, article 4 with a subterminal posterodistal tooth, flagellum with

19-22 articles. Coxa 4, posterior margin excavate. Gnathopod 2, article 2

spinose on posterior, medial margin in males, palm slightly oblique, with four

defining spines. Pereopods 3 and 4 unmodified, moderately setose, dactyl with

4-7 spinules. Pereopods 5-7 moderately to densely setose, dactyl with 11-13

spinules. Uropod 1, peduncle spinose and setose, rami subequal, both with mar-

ginal and apical spines, inner ramus sometimes with a single seta. Uropod 2,

peduncle spinose and setose, rami approximately subequal, both with marginal

and apical spines, inner ramus with a few marginal setae. Uropod 3, inner

ramus Q.3 length of outer, with two apical spines, outer ramus with marginal

and apical spines and setae, second segment distinct but small, about 4-5 per

cent of first. Telson deeply cleft, each lobe with a single spine and about

6-8 setae.

Remarks

The extremely elongate antenna 2 with a subterminal ‘tooth’ on article 4 of

the peduncle in males makes this species unmistakable.

Distribution

Known from tributaries of the Palmiet River (Fig. 29).

Paramelita parva Stewart & Griffiths, 1992

Fice2t

Paramelita parva Stewart & Griffiths, 1992c: 501-504, figs 8-9.

Material examined

Types. Holotype, SAM-A40226; paratypes, SAM-A40227, from a tributary

of the Storms River, eastern Cape.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 231

Fig. 21. Paramelita parva, SAM-A40226, holotype, male, 8.7 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2, medial view. E. Pereopod 3. F. Coxa 4.

G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Other material. SAM-A40228, A40229, A40230 and A40775, all from

tributaries of the Storms River, eastern Cape.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 22- to 26-articulate,

accessory flagellum 4- to 5-articulate. Antenna 2 sparsely to moderately setose,

shorter than 1, peduncle not enlarged in males, flagellum with 11-18 articles.

Coxa 4, posterior margin excavate. Gnathopod 2, palm moderately oblique,

with 3-4 defining spines. Pereopods 3 and 4 moderately setose, unmodified,

dactyl with 2-3 spinules. Pereopods 5-7 moderately setose, dactyl with 4-7

spinules. Uropods 1 and 2, peduncle spinose, lacking setae, rami with marginal

and apical spines, lacking setae. Uropod 3, inner ramus 0.3 length of outer,

apically spinose, outer ramus with marginal and apical spines, moderately to

densely setose, second segment small but distinct. Telson deeply cleft, each lobe

with 1-2 spines and 1-4 setae.

Remarks

This species is morphologically similar to Paramelita kogelensis, from

which it is distinguished by the number of spinules on the dactyl of pereopods 3

and 4, and the setation of the uropods.

Distribution

The most isolated of all the paramelitid species, P. parva has been collected

from the Storms River catchment, eastern Cape (Fig. 29).

Paramelita pillicornis Stewart & Griffiths, 1992

Fig. 22

Paramelita pillicornis Stewart & Griffiths, 1992c: 494-497, figs 4-5.

Material examined

Types. Holotype, SAM-A40214; paratypes, SAM-A40215, from a tributary

of Waboomsrivier on Gydo Pass, north of Ceres.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 18- to 33-articulate,

accessory flagellum 3- to 4-articulate. Antenna 2 shorter than 1, densely setose

posteriorly in males, flagellum with 13-16 articles. Coxa 4, posterior margin

excavate. Gnathopod 2, palm distinctly oblique, with three defining spines.

Pereopods 3-7 moderately to densely setose. Pereopods 5-7, article 2 markedly

poorly expanded, dactyl with 3-5 spinules. Uropod 1, peduncle spinose and

setose, both rami with marginal and apical spines and sometimes marginal setae.

Uropod 2, peduncle spinose, lacking setae, inner ramus with marginal spines,

outer ramus with marginal spines and sometimes setae, both with apical spines.

Uropod 3, inner ramus 0.3-0.4 length of outer, apically spinose and setose,

outer ramus with marginal and apical spines, moderately setose, second segment

small but distinct. Telson deeply cleft, each lobe with one spine and 3-8 setae.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 233

7m]

SF;

Fig. 22. Paramelita pillicornis, SAM-A40214, holotype, male, 10.8 mm. A. Lateral

aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2, medial view. E. Pereopod 3.

F. Coxa 4. G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. K. Telson.

Scale lines represent 1 mm.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

One of two white-eyed Paramelita species with a highly setose antenna 2,

P. pillicornis is distinguished from P. seticornis by the relative length of the

peduncle and the number of articles in the flagellum of this antenna, the width of

article 2 in pereopods 5-7, and the setation of the rami in uropods 1 and 2.

Distribution

Known only from the type locality, Gydo Pass, north of Ceres (Fig. 29).

Paramelita pinnicornis Stewart & Griffiths, 1992

Fig. 23

Paramelita pinnicornis Stewart & Griffiths, 1992a: 140-144, figs 1-2.

Material examined

Types. Holotype, SAM-A40004; paratypes, SAM-A40005, from a tributary

of the Burgersbos River, Cape Peninsula.

Other material. SAM-A10017, from Newlands, Cape Peninsula. SAM-

A40008, from Kenilworth Race Course, Cape Peninsula. SAM-A40006 and

A40007, from adjacent streams in the Cape Hangklip area, east coast of False

Bay.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 31- to 46-articulate,

accessory flagellum 5- to 6-articulate. Antenna 2 sparsely to moderately setose,

of equal length to 1, articles 4 and 5 of peduncle in males extremely elongate

and outer margin and tip of article 5 extended into an elongate triangular flange,

flagellum with 16-23 articles. Coxa 4, posterior margin excavate. Gnathopod 2,

article 2 strongly spinose medially, palm oblique, with 3-5 defining spines.

Pereopod 3 moderately setose, modified in males, article 4 elongate, article 5

with a posterior lump and a few long, blade-like spines, article 6 curved,

attached at right angle to 5, dactyl with 6-7 spinules. Pereopod 4 unmodified,

dactyl with 6-7 spinules. Pereopods 5-7, dactyl with 10-14 spinules. Uropods 1

and 2, peduncle spinose and setose, rami with marginal spines and setae and

apical spines. Uropod 3, inner ramus 0.1-0.2 length of outer, apically spinose,

Outer ramus with marginal and apical spines, densely setose, second segment

small but distinct, apically spinose. Telson deeply cleft, each lobe with one

spine and several setae.

Remarks

The unusual form of antenna 2 and pereopod 3 is unique to this species.

Distribution

This species has a rather disjunct distribution, and is known from the north-

ern and eastern parts of the Cape Peninsula and also the Cape Hangklip area,

along the east coast of False Bay (Fig. 29).

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 235

Fig. 23. Paramelita pinnicornis, SAM-A40004, holotype, male, 13.5 mm. A. Lateral

aspect. B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Coxa 4.

G. Pereopod 7. H. Uropod 1. I. Uropod 2. J. Uropod 3. Scale lines represent 1 mm.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paramelita platypus Stewart & Griffiths, 1992

Fig. 24

Paramelita platypus Stewart & Griffiths, 1992a: 153-157, figs 7-8.

Material examined

Types. Holotype, SAM-A40020; paratypes, SAM-A40021, from a tributary

of the Fernkloof River in the Fernkloof Nature Reserve.

Other material. SAM-—A40022 and A40516, from a stream near Stanford.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 28- to 41-articulate,

accessory flagellum 4- to 5-articulate. Antenna 2 sparsely setose, of equal length

to 1 in adult males, both peduncle and flagellum extremely elongate and stout,

flagellum with 16-22 articles. Coxa 4, posterior margin very slightly emargin-

ate. Gnathopod 2, article 2 sparsely medially spinose, palm transverse, mark-

edly convex, with a tooth and 2-5 spines. Pereopods 3 and 4 densely setose

posteriorly, modified in males, article 4 greatly expanded laterally, dactyl with

3-5 spinules. Pereopods 5-7 moderately to densely setose, dactyl with 6-10

spinules. Uropods 1 and 2, peduncle spinose and setose, inner rami with mar-

ginal spines and setae, outer rami with marginal spines, lacking setae, each rami

with apical spines. Uropod 3, inner ramus 0.2 length of outer, apically spinose,

outer ramus with marginal and apical spines, moderately setose, second segment

small but distinct. Telson deeply cleft, each lobe with one spine and 6-8 setae.

Remarks

The combination of an extremely elongate antenna 2, a markedly convex

palm with a distinct palmar tooth in gnathopod 2, laterally expanded article in

pereopods 3 and 4, almost quadrate coxa 4, and a distinct, albeit small, second

segment on the outer ramus of uropod 3 make this species unmistakable.

Distribution

This species has been collected from streams draining the slopes of the

Kleinriviersberge between Hermanus and Stanford (Fig. 29).

Paramelita seticornis (Barnard, 1927)

Fig. 25

Gammarus Seticornis Barnard, 1927: 171-172, pl. 10 (figs 7, 17).

Paramelita seticornis (Barnard) Thurston, 1973: 166-167. Griffiths, 1981: 90, fig. 2D-F.

Material examined

Types. Syntypes, SAM-A3994, from Landdrost Kloof, Caledon side,

Hottentots Holland Mountains.

Other material. SAM-—A40933, from Sir Lowry’s Pass, Somerset West side,

Hottentots Holland Mountains. SAM-A40946, from Malkopvlei, Betty’s Bay.

SAM-A40525, Outeniqua Mountains. SAM-A40777 and A40947, Betty’s Bay.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 237

‘ (RU ape /}

Lee

ikke SE

Ly 1) \ my

EW, rae N

KRG

ev? &Q AA .

—IN mY

Fig. 24. Paramelita platypus, SAM-A40020, holotype, male, 12.8 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2, right side, medial view. E. Pereopod 3.

F. Coxa 4. G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3.

Scale lines represent 1 mm.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘oc

nae

¥,

Zig

ry)

ANN -

)

INN

Fig. 25. Paramelita seticornis, SAM-A40933, male, 9.0 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2, lateral and medial views. E. Pereopod 3.

F. Coxa 4. G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3. K. Telson.

Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 239

Diagnosis

Eyes white. Antenna 1 sparsely setose, flagellum 20- to 30-articulate,

accessory flagellum with 4-5 articles. Antenna 2 stout and densely setose in

males, moderately stout and setose in females, flagellum 8- to 12-articulate.

Coxa 4, posterior margin with a shallow but distinct emargination. Gnatho-

pod 2, palm oblique, with 2-4 palmar spines. Pereopods 3 and 4 unmodified,

dactyl with 2-4 spinules. Pereopods 5-7, dactyl with 4-10 spinules. Uropod 1,

peduncle with spines and setae, rami subequal, with marginal and apical spines,

inner ramus with some setae. Uropod 2, peduncle with spines and setae, inner

ramus longer than outer, both with marginal and apical spines, inner ramus with

some setae. Uropod 3, inner ramus about 0.3 length of outer ramus, with 2-3

apical spines, outer ramus with marginal and apical spines and setae, second

segment rudimentary. Telson deeply cleft, each lobe with a single spine and

several setae.

Remarks

This species is distinguished from an allied Hottentots Holland form, Para-

melita kogelensis, by its densely setose antenna 2, a condition it shares with

P. pillicornis.

Distribution

Known from Hottentots Holland and adjacent mountain ranges (Fig. 29).

Paramelita spinicornis (Barnard, 1927)

Fig. 26

Gammarus spinicornis Barnard, 1927: 174-175, pl. 10 (figs 8, 18-19).

Paramelita spinicornis (Barnard) Thurston, 1973: 166-167. Griffiths, 1981: 91, fig. 3D.

Stewart et al., 1994: 179-190, figs 1-4.

Material examined

Types. Syntypes, SAM-A5177, from Hottentots Holland Mountains.

Other material. SAM-A5180, Hottentots Holland Mountains. SAM-A5186,

Steenbras Valley. SAM-A6053, Zonderend Mountains. SAM-A6939, Swellen-

dam Mountains. SAM-A6940, Swellendam Mountains. SAM-A6941, Zuur-

braak Peak. SAM-A6942, Tradouw Peak. SAM-A6943, south of Barrydale.

SAM-A40253, Betty’s Bay. SAM-A40254, Disa Kloof, Betty’s Bay. SAM-

A40255, Fernkloof Ravine, Hermanus. SAM-A40256, Harold Porter Gardens,

Betty’s Bay. SAM-A40257, Lamloch Stream, Kleinmond. SAM-A40258,

between Betty’s Bay and Kleinmond. SAM-A40517, Nuweberg State Forest.

SAM-A40526, Grabouw Forest. SAM-A40528, Swartberg, near Caledon.

SAM-A40784 and A40785, Betty’s Bay. SAM-A40948, between Hermanus

and Stanford. SAM-A41010, Grotto Beach, Hermanus.

Diagnosis

Eyes white. Antenna 1 sparsely setose, flagellum 20- to 25-articulate,

accessory flagellum 4-articulate. Antenna 2 shorter than 1, sparsely setose,

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

—

g .N

f lA \

a | q \

<Fi \ \ NESS i

Fig. 26. Paramelita spinicornis, SAM-A40253, male, 7.0 mm. A. Lateral aspect.

B. Antenna 1. C. Antenna 2. D. Gnathopod 2. E. Pereopod 3. F. Pereopod 6.

G. Uropod 1. H. Uropod 2. I. Uropod 3. Scale lines represent 1 mm.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 241

article 4 of peduncle strongly laterally swollen and with a proximal medial lobe

and a posterodistal tooth in males, flagellum 9- to 11-articulate. Coxa 4,

posterior margin excavate. Gnathopod 2, article 2 not medially spinose, palm

slightly oblique, with 3-4 defining spines. Pereopods 3 and 4 sparsely setose,

unmodified, articles 4, 5 and 6 strongly spinose, dactyl with 3-4 spinules.

Pereopods 5-7 sparsely to moderately setose, dactyl with 5-8 spinules.

Uropod 1, peduncle spinose, usually with a single seta, rami subequal, both

with marginal and apical spines, lacking setae. Uropod 2, peduncle spinose,

usually with 1-2 setae, outer ramus longer than inner, both rami with marginal

and apical spines, inner ramus sometimes with a single seta. Uropod 3, inner

ramus 0.3 length of outer, apically spinose, outer ramus with marginal and

apical spines, sparsely setose, small but distinct second segment present. Telson

deeply cleft, each lobe with one spine and 3-5 setae.

Remarks

The possession of a terminal, posterodistal tooth on the laterally swollen

article 4 of antenna 2 in males, excavate coxa 4, multispinose dactyls on the

pereopods, and the presence of a distinct second segment on the outer ramus of

uropod 3 make this species unmistakable, despite its superficial resemblance to

Aquadulcaris dentata and Paramelita odontophora.

Distribution

Collected from Hottentots Holland to Swellendam mountains (Fig. 29).

Paramelita tulbaghensis (Barnard, 1927)

Fig. 27

Gammarus tulbaghensis Barnard, 1927: 170-171, pl. 10 (figs 5, 15).

Paramelita tulbaghensis (Barnard) Thurston, 1973: 166-167. Griffiths, 1981: 91, fig. 3H-I.

Material examined

Types. Syntypes, SAM-A4875, from the Sneeuwgat Valley near Tulbagh.

Other material. SAM-A40934, from a stream on the path to Sneeuwgat

Peak above the farm Bergplaas, foot of the Winterhoek Mountains, near

Tulbagh. SAM-A40935, from a stream in the Ceres municipal campsite. SAM-

A40232, from a tributary of the Molenaar’s River, Worcester end of Du Toit’s

Kloof Pass.

Diagnosis

Eyes white. Antenna | sparsely setose, flagellum 16- to 25-articulate,

accessory flagellum 3- to 4-articulate. Antenna 2 shorter than 1, sparsely setose,

slender in females, articles 3, 4 and 5 of peduncle swollen in males, article 4

the longest, flagellum 9- to 12-articulate. Coxa 4, posterior margin with a

distinct but shallow emargination. Gnathopod 2, palm transverse to slightly

oblique, with 2-4 spines. Pereopods 3 and 4 unmodified, dactyl usually with

two, but sometimes with one spinule. Pereopods 5-7, dactyl usually with 2-4,

B. Antenna 1.

pod 6. H. Uropod 1

C. Antenna 2

ANNALS OF THE SOUTH AFRICAN MUSEUM

“ES

“ih

<< — — —

D. Gnathopod a),

. I. Uropod 2.

Fig. 27. Paramelita tulbaghensis, SAM-A4875, syntype, male, 6.3 mm. A. Lateral view

5» Pereopod seat: Coxa 4. G. Pereo-

J. Uropod 3. Scale lines represent 1 mm

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 243

but sometimes with one spinule. Uropod 1, peduncle with spines and setae, rami

subequal, usually with marginal, and always with apical spines, sometimes with

a few marginal setae. Uropod 2, peduncle with spines and setae, inner ramus

longer than outer, both with marginal and apical spines, inner ramus with at

least one seta. Uropod 3, inner ramus about 0.3 length of outer, with some

apical setae, outer ramus with a few marginal and apical spines and setae,

second segment rudimentary. Telson deeply cleft, each lobe usually with a

single spine and a few setae.

Remarks

Paramelita tulbaghensis is most similar to Aquadulcaris crassicornis, from

which it is most easily distinguished by the possession of usually 2-4 spinules

on the, dactyl of pereopods 3-7, and by the lack of tooth-like spines on article 5

of pereopod 3.

Distribution

This species has been collected from the Winterhoek Mountains near Tul-

bagh in the north to Du Toit’s Kloof on the Dutoitsberg in the south (Fig. 29).

Paramelita validicornis Stewart & Griffiths, 1992

Fig. 28

Paramelita validicornis Stewart & Griffiths, 1992c: 497-501, figs 6-7.

Material examined

Types. Holotype, SAM-A40216; paratypes, SAM-A40217, from a stream

flowing into Kleinriviervlei, near Hermanus.

Other material. SAM-A7394, from near Bredasdorp. SAM-A40218, from

a tributary of the Afdaksrivier. SAM-A40219, A40514, and A40789, from

Fernkloof Nature Reserve, Hermanus. SAM-A40518, Salmonsdam Nature

Reserve. SAM-A40924, Hermanus Yacht Club. SAM-A41007, Grotto Beach,

Hermanus.

Diagnosis

Eyes white. Antenna 1 sparsely setose, flagellum with 44-48 articles,

accessory flagellum 5- to 6-articulate. Antenna 2 sparsely setose, peduncle

elongate and stout in males, flagellum with 19-22 broad, flattened articles.

Coxa 4, posterior margin excavate. Gnathopod 2, article 2 medially spinose,

palm distinctly oblique, with 4-5 spines. Pereopods 3-7 moderately setose,

unmodified, dactyl with 4-9 spinules. Uropods 1 and 2, peduncle spinose,

1 sometimes with 1-2 setae, inner rami with marginal spines and setae, outer

rami with marginal spines, lacking setae, all rami with apical spines. Uropod 3,

inner ramus 0.2 length of outer, apically spinose and setose, outer ramus

with marginal and apical spines, moderately to densely setose, second segment

small but distinct. Telson deeply cleft, each lobe with 1-2 spines and

6-10 setae.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 28. Paramelita validicornis, SAM-A40216, holotype, male, 13.8 mm. A. Lateral

aspect. B. Antenna 1. C. Antenna 2. D. Gnathopod 2, medial view. E. Pereopod 3.

F. Coxa 4. G. Pereopod 6. H. Uropod 1. I. Uropod 2. J. Uropod 3.

Scale lines represent 1 mm.

245

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE

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246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The most distinguishing features of this species are the stout and elongate

antenna 2, particularly in males, and the relatively long outer ramus in

uropod 3. The latter condition, along with differences in setation of the rami of

uropods | and 2, is used to separate Paramelita validicornis from P. magna and

P. magnicornis.

Distribution

Although this species is known from Bredasdorp in the west to the Klein-

riviersberge in the east, it is possibly more widespread (Fig. 29).

ACKNOWLEDGEMENTS

Special thanks go to all those willing hands who assisted in the field.

Michelle van der Merwe kindly assisted with the drafting of figures. Financial

support was provided by the Foundation for Research Development through a

grant from the Inland Waters Ecosystems Programme, and through the Special

Programme on Rivers directed by Professor Bryan Davies.

REFERENCES

BARNARD, K. H. 1916. Contributions to the crustacean fauna of South Africa. 5. The

Amphipoda. Annals of the South African Museum 15 (3): 105-302.

BARNARD, K. H. 1927. A study of the freshwater isopodan and amphipodan Crustacea of

South Africa. Transactions of the Royal Society of South Africa 14 (2): 139-215.

BARNARD, J. L. & BARNARD, C. M. 1983. Freshwater Amphipoda of the world. I. Evol-

utionary patterns. II. Handbook and bibliography. Mt. Vernon: Hayfield Associates.

BOUSFIELD, E. L. 1983. An updated phyletic classification and palaeohistory of the

Amphipoda. Jn: SCHRAM, F. R. ed. Crustacean phylogeny: 257-277. Rotterdam: A. A.

Balkema.

CONLAN, K. E. 1988. Phenetic and cladistic methods applied to a small genus (Corophi-

oidea: Ischyroceridae: Microjassa) and a larger outgroup. Crustaceana. Supplement 13:

143-166.

Farris, J. S. 1988. Hennig86. Version 1.5.

GRIFFITHS, C. L. 1981. The freshwater Amphipoda (Crustacea) of South and South West

Africa. Annals of the South African Museum 83: 79-97.

HOLSINGER, J. R. 1992. Sternophysingidae, a new family of subterranean amphipods

(Gammaridea: Crangonyctoidea) from South Africa, with description of Sternophysinx

calceola, new species, and comments on phylogenetic and biogeographic relationships.

Journal of Crustacean Biology 12: 111-124.

Myers, A. A. 1988. A cladistic and biogeographic analysis of the Aorinae subfamily nov.

Crustaceana. Supplement 13: 167-192.

NoTENBOOM, J. 1988. Phylogenetic relationships and biogeography of the groundwater-

dwelling amphipod genus Pseudoniphargus (Crustacea), with emphasis on the Iberian

species. Bijdragen tot de dierkunde 58: 159-204.

PIMENTAL, R. A. & RIGGINS, R. 1987. The nature of cladistic data. Cladistics 3: 201-209.

RIDLEY, M. 1986. Evolution and classification: The reformation of cladism. London:

Longman.

ROHLF, F. J. 1989. NTSYS-pce. Numerical taxonomy and multivariate analysis system.

Version 1.5. Stony Brook: Exeter Publishing Ltd.

SCHELLENBERG, A. 1926. Die Gammariden der Deutschen Siidpolar-Expedition 1901-

1903. Deutsche Stidpolar-Expedition, 1901-1903 18: 235-414.

REVISION OF THE AMPHIPOD FAMILY PARAMELITIDAE 247

SCHELLENBERG, A. 1937. Kritische Bemerkungen zur Systematik der Siisswasser-

gammariden. Zoologische Jahrbiicher. Abteilung fiir Systematik, Geographie und

Biologie der Tiere 69: 469-516.

STEWART, B. A. 1992. Morphological and genetic differentiation between allopatric popu-

lations of a freshwater amphipod. Journal of Zoology, London 228: 287-305.

STEWART, B. A. & GRIFFITHS, C. L. 1992a. Four new species of the genus Paramelita

(Amphipoda, Crangonyctoidea) from South Africa. Annals of the South African Museum

101: 139-158.

STEWART, B. A. & GRIFFITHS, C. L. 1992b. A taxonomic re-examination of freshwater

amphipods in the Paramelita auricularius-P. crassicornis complex, with descriptions of

three additional species. Crustaceana 62: 166-192.

STEWART, B. A. & GRIFFITHS, C. L. 1992c. Further new species within the freshwater

amphipod genus Paramelita (Crangonyctoidea: Paramelitidae) from South Africa.

Journal of Natural History 26: 489-506.

STEWART, B. A., SNADDON, C. D. & GRIFFITHS, C. L. 1994. Morphological and genetic

differentiation among populations of the freshwater amphipod Paramelita spinicornis

(Amphipoda: Crangonyctoidea), with description of a new species. Zoological Journal

of the Linnean Society 111: 179-195.

THuRSTON, M. H. 1973. A new species of Paramelita (Crustacea: Amphipoda) from South

Africa. Annals of the South African Museum 62: 159-168.

Watrous, L. E. & WHEELER, Q. D. 1981. The outgroup comparison method of character

analysis. Systematic Zoology 30: 1-11.

WILLIAMS, W. D. & BARNARD, J. L. 1988. The taxonomy of crangonyctoid Amphipoda

(Crustacea) from Australian fresh waters: Foundation studies. Records of the Australian

Museum. Supplement 10: 1-180.

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particu-

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. °.. . the Figure depicting C. namacolus .. .’: ‘. . . in C. namacolus (Fig. 10). . .’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by

initials or full names

e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

not be abbreviated at the beginning of a sentence or paragraph.

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.