MARJORIE ESTELLF CART

and JANET M. BRADFORJ

Postembryonic Development

of Three Species of A

Freshwater Harpacticoid

Copepoda

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 119
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 119

Marjorie Esteik Carter Postembryonic Development

and Janet M. Bradford Q f Xhre e Species of

Freshwater Harpacticoid

Copepoda

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972
ABSTRACT

Carter, Marjorie Estelle, and Janet M. Bradford. Postembryonic Development of

Three Species of Freshwater Harpacticoid Copepoda. Smithsonian Contributions

to Zoology, number 119, 26 pages, 14 figures. 1972.—The postembryonic development of three species of freshwater harpacticoid copepods from southern Georgia

and southwestern Virginia has been studied. All of the larval stages (six nauplii

and six copepodid stages) of Elaphoidella bidens coronata and Bryocamptus

zschokkei alleganiensis, and all of the nauplii and two copepodid stages of Moraria

virginiana were obtained, fully described, and figured. Where possible, the anatomical details were determined from the exuviae of a single individual.

A key to the identification of the six naupliar stages of harpacticoids is given as

well as a chart summarizing the major features of each copepodid stage.

It is suggested that all life cycles of harpacticoid copepods in which fewer than

six naupliar stages have been reported should be reexamined.

FOREWORD

This report represents the major portion of a dissertation submitted by Dr. Marjorie

Estelle Carter to the faculty of the University of Virginia in partial fulfillment of the

requirements for the Ph.D. degree that was awarded to her in June 1952. Her teaching

duties at Valdosta State College and involvement in student affairs for some 16 years

delayed the revision needed before submitting it for publication,, and not until 1968,

with the encouragement of Dr. Thomas E. Bowman and Miss Gayle A. Heron, did she

begin work on a revised draft. Following her untimely death later in the year, the

manuscript, together with her illustrations, was sent to me, and through the efforts of

Dr. Janet M. Bradford, it has been brought up-to-date, and pertinent references to studies

that have been made subsequent to Dr. Carter's investigations included.

As Dr. Carter's major professor, I should like to thank Dr. Bradford for her interest

and the work that she has devoted to revising the copy supplied to her. I am sure that

Dr. Carter would have concurred in my insistence that Dr. Bradford's name appear as a

coauthor. I am also grateful to Dr. Bowman for his many helpful suggestions and his

critical reading of the final draft. Thanks are also due Miss Margaret A. Daniel for

arranging and refurbishing the figures.

The acknowledgments within the body of the text are quoted, almost verbatim, from

the dissertation (original manuscript).

HORTON H. HOBBS, JR.

Library of Congress Cataloging in Publication Data

Carter, Marjorie Estelle, 1905-1968.

Postembryonic development of three species of freshwater Harpacticoid Copepoda.

(Smithsonian contributions to zoology, no. 119)

A revision of M. E. Carter's thesis—University of Virginia, 1952.

Bibliography: p.

1. Elaphoidella bidens. 2. Bryocamptus zschokkei. 3. Moraria virginiana. I. Bradford, Janet M.,

1940- joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions

to zoology, no. 119.

QL1.S54 no. 119 [QL444.C7] 591'.08s [595'.34] 72-277

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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

Washington, D.C. 20402 - Price 40 cents (paper cover)
Esteiu Carter Postembryonic Development

and Janet M. Bradford Qf Thre e SpeCieS of

Freshwater Harpacticoid

Copepoda

Introduction

Few complete studies on larval and postlarval development of harpacticoid copepods have been made,

and only three observations exist on the genera (Elaphoidella, Bryocamptus, and Moraria) considered

here (Lang, 1948).

There is disagreement as to the number of naupliar

stages in the development of harpacticoids. Although

six naupliar stages have been found in some (Chappuis, 1916; Borutskii, 1925; Ewers, 1930; Johnson

and Olson, 1948; Krishnaswamy, 1951; Grig, 1960;

Fehrenbach, 1962; and Haq, 1965), only five naupliar

stages have been described for the genera Elaphoidella, Bryocamptus, and Moraria (Chappuis, 1916;

Borutskii, 1925; and Gurney, 1932).

The present report traces the postembryonic development in three species, two from the first naupliar

stage through the adult, the other to the second copepodid stage.

All figures except those with individual scales are

drawn to the scale indicated in the first component.

Systematic Status of Species Investigated

FAaphoidella bidens coronata (SARS, 1904).—I have

followed Coker (1934) and Wilson (1956) in assigning the specimens used in this study to the subspecies

coronata, described originally from the Hawaiian Islands. For a discussion of the somewhat involved

Marjorie Estelle Carter, Valdosta State College, Valdosta,

Georgia (deceased 1968). Janet M. Bradford, Collaborator,

Division of Crustacea, National Museum of Natural History,

Smithsonian Institution, Washington, D.C. 20560.

taxonomic status of North American specimens of E.

bidens, see Wilson (loc. cit.).

In order that no confusion arise as to the identity

of the animals used in the present investigation, detailed illustrations are given of the adult male (Figures 2i, j ; 3/; An; 5/, m; 6e, k, n, p) and the adult

female (Figures 1g, h; 3g, m; 4d, j , o, s; 5g, n; 6f, j ,

o, q).

Bryocamptus zschokkei alleganiensis COKER, 1934.—

The harpacticoids used here agree, for the most part,

with Coker's (1934:93) description of the subspecies

Bryocamptus zschokkei alleganiensis. However, the

number of segments of the endopods of the second

and third legs of the female varies in individuals from

the same locality. Some specimens have two segments

as described by Coker, others have the endopod threesegmented on the second leg but only two-segmented

on the third leg, and in some the endopods of both

legs have three segments. Similar variations have been

observed by Gurney (1932:148) and by Kiefer

(1929:321).

The adult male (Figures 8g; 10/, m; 11/, m) and

female (Figures 8/, 9a-c, j ; \0g, n; lie, n) are

figured.

Moraria virginiana CARTER, 1944.—This species

was described (Carter, 1944) from only two specimens, both female. Drawings of the hitherto undescribed male and most of its appendages are included

here (Figures I3e, We, g, h, j-p).

Materials and Methods

The animals used in this study were obtained from

several localities. Elaphoidella bidens coronata was

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

collected from two localities in the vicinity of Valdosta, Lowndes County, Georgia. Late in 1948, I collected several females and a single male from a small

artificial pool on the campus of the Valdosta State

College. The next year, I could find no harpacticoids

in this habitat but found them abundant in a small

stream at the north end of town. No males were

taken from the stream, but several females with eggs

were found, and from one of these I succeeded in

rearing three generations of females parthenogenetically.1

Bryocamptus zschokkei alleganiensis is found in

abundance in nearly every small stream in the vicinity

of Mountain Lake, Giles County, southwestern Virginia, and is usually associated with growths of algae

and moss. Animals from several locations around the

Mountain Lake Biological Station were collected and

reared. Those used in the drawings and descriptions

were reared from individuals taken from a small

stream that flows into Little Stony Creek, about two

miles from the Station, while those taken from other

localities were used for comparison.

Moraria virginiana was originally collected from a

small group of springs, locally known as Twin

Springs, near the Mountain Lake Biological Station

(Carter, 1944). Since then, I have taken many animals, both male and female, from swampy places

where there is a heavy growth of sphagnum moss

along the trail leading from the Station to Twin

Springs.

Adult animals were collected by taking small

amounts of water and debris from the bottom of

small streams and pools. This material was examined

with the aid of a wide-field stereoscopic microscope,

and the animals placed in culture dishes where they

could be observed more easily. Females with eggs

were isolated in culture dishes until the eggs hatched,

whereupon some of them were isolated, and others

were left in the dish with the mother. When only one

species of harpacticoid was found in a collection, the

water and detritus contained in it were strained

through bolting silk and placed in culture dishes, and

the larvae and adults introduced into the filtrate

seemed to thrive. In some instances, an artificial cul1

 A discussion of the rearing of this fpecies parthenogenetically is given by Roy (1931), and the scarcity of males

reported by Lang (1948).

ture medium, similar to that used by Banta (1921)

in raising Daphnia, was used.

Square watch glasses were most convenient for culturing the copepods, but small chemical watch glasses,

placed in Syracuse dishes to support them, were satisfactory. All culture dishes were covered with small

squares of glass and kept in a moist chamber.

The entire larva is difficult to study when stained,

and most of my observations are based on the exuviae. This procedure enabled the development of a

single individual to be followed through its various

stages. Although the tests are very difficult to find in

the culture dishes because of their transparency, I

found that directing light on a mirror on which the

culture dish had been placed reflected just enough

light on the tests so they could be seen more readily.

The adults and the larvae which died were treated

with 5 to 10 percent potassium hydroxide in order to

remove the tissues.

The adults and copepodid larvae, stained with

alum cochineal, provide satisfactory material for

mounting; however, Reynold's Stain gives better results. The test of the naupliar stages are difficult to

stain because of the very thin cuticle; best results

were obtained when the larvae were left in acid

fuchsin for 24 to 48 hours and then treated with

glacial acetic acid and clove oil before mounting in

balsam. The adults and copepodids were mounted in

glycerine sealed with paraffin so that they could be

removed for dissection.

Drawings were made with the aid of a camera

lucida, and the figures of all stages were made, insofar

as possible, from the exuviae of a single individual.

Postembryonic Development

Elaphoidella bidens coronata (Sars)

First Naupliar Stage

FIGURE \a, g

Body inflated, with subcircular and characteristically

ornamented dorsal shield. Two median lobes present

posterior to midlength; anterior lobe semicircular

with posteromedian emargination; posterior lobe

bearing posteromedian knob (Figure lg).

Labrum subspatulate, very narrow anteriorly; anterior margin distinctly emarginate. Narrow portion
NUMBER 1 I 9

of labrum lying between first antennae; posterior to

first antennae, lateral margins abruptly diverging

before turning posteriorly. Posterior margin fringed

with fine hairs and with tuft of longer median hairs.

ANTENNA I.—Segment 1 short and naked. Segment

2 about twice length of proximal segment, bearing

three terminal spinules and three setae: proximal and

median setae short and rather delicate; subterminal

seta heavy and extending beyond end of terminal

segment. Segment 3 about two-thirds length of segment 2, bearing one subterminal group of spinules

and two subterminal and one terminal seta; latter

about three times length of subterminal setae.

ANTENNA II.—Coxopodite broader than long with

semicircular row of spinules on distoventral surface,

its proximopostaxial angle produced into curved acute

process, bordered by dentations from apex to beginning of curve, projecting above posterior portion of

labrum. Basipodite broader than long but narrowing

into short postaxial protuberance bearing one seta.

Exopodite three-segmented; segment 1 almost as long

as combined length of distal segments, bearing short

postaxial seta at midlength and terminal seta nearly

four times length of segment; segment 2 one-third

length of segment 1, naked; segment 3 with subterminal seta slightly exceeding length of segment and

terminal seta more than twice length of entire exopodite. Endopodite consisting of one long segment bearing very delicate seta at midlength, group of subterminal spinules, and two terminal setae; one of latter

narrow and very short, other stout, almost as long

as segment, and apparently movably articulated at its

base.

MANDIBLE.—Coxopodite dorsal to basipodite, its

median border produced into rounded lobe. Basipodite subrectangular with two oblique rows of spinules

on ventral surface and plumose spine on proximoventral surface. Exopodite two-segmented: segment 1

slightly longer than basipodite, bearing small spine at

midlength and subterminal seta; segment 2 about

one-third length of segment 1, bearing short subterminal seta and terminal seta, latter about threefourths length of entire animal. Endopodite one-segmented, bearing two curved serrate spines and four

slender setae.

Ventral plate (Figure la) about one-third broader

than long with two rows of spinules across ventral

surface, one near midlength and other on caudal margin. Caudal margin rounded but produced posterolaterally in paired protuberances, each terminating

in single fine seta.

Second Naupliar Stage

FIGURE \b

Second nauplius differing from first in following

respects: ornamentation of dorsal shield indistinct except for retention of anterior median lobe.

ANTENNA I.—Segment 3 with additional group of

spinules just distal to midlength, and subterminal seta

about one-third length of terminal seta present in first

nauplius.

ANTENNA II.—Segment 1 of exopodite with additional spine on preaxial surface near midlength, its

size similar to that of corresponding postaxial spine.

Large simple terminal seta of endopodite with two

small spinules at midlength.

MANDIBLE.—Basipodite with second seta, slenderer

than first, on postaxial surface. Endopodite with one

additional spine lateral to those present in first

nauplius.

MAXILLA I.—First maxillae, absent in first nauplius, represented by pair of feathered setae on anterior third of ventral plate. Ventral plate emarginate

caudally; posterior rows of spinules lacking but replaced by short lateral spinules.

Third Naupliar Stage

FIGURE lc

Third nauplius differing from second in following

respects:

ANTENNA I.—Segment 3 with two additional subterminal setae proximally about half length of subterminal seta of second nauplius; distalmost subterminal seta subequal in length to terminal seta.

ANTENNA II.—Protuberance on postaxial margin

of postaxial extension of coxopodite bearing short

feathered seta instead of tuft of hairs characteristic

of second nauplius. Segment 3 of exopodite bearing

subterminal seta approximating length of terminal

seta. Endopodite with additional group of spinules

at about midlength.

Ventral plate lengthened, with posterior part of

body extending well beyond dorsal shield; caudal

margin of plate with second pair of protuberances

mesial to first pair.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 1.—Elaphoidella bidens coronata: a, first nauplius, ventral view; b, second nauplius,

ventral view; c, third nauplius, ventral view; d, fourth nauplius, ventral view; e, fifth nauplius,

ventral view; /, sixth nauplius, ventral view; g, first nauplius, dorsal view; h, fourth nauplius,

dorsal view; i, fifth nauplius, dorsal view; ;', sixth nauplius, dorsal view.
NUMBER 119

Fourth Naupliar Stage

FIGURE Id, h

Fourth nauplius differing from third in following

respects: Antenna I with segment 3 bearing additional small seta. Ventral plate with single pair of

lobes bearing seta of first maxilla. Posterior region of

body longer, its caudal margin bearing two setae lateral to each of original pair; pair of prominent setae

present on protuberances first appearing in third

nauplius. Anal plate with fine spinules along posterior

border.

Fifth Naupliar Stage

FIOURE \e, i

Fifth nauplius differing from fourth in following

respects: Antenna I with segment 3 bearing one additional seta; maxilla I with lobe more clearly delineated, and bearing small protuberances. Pygidial

area with additional ventral seta on each side.

Sixth Naupliar Stage

FIGURE If, }

Sixth nauplius differing from fifth in following respects: Ventral plate bearing small seta on lobe of

second maxilla; rudiment of maxilliped represented

by lobe. Posterior region consisting of two segments

indicated by two pairs of swimming legs and by dentate margins dorsally, and pygidial area.

First Copepodid Stage

FIGURES 2a,b; 3a,h; 4a,e,k,p; 5a,h

Body consisting of cephalothorax and four free segments.

ANTENNA I (Figure 3a).—As in sixth naupliar

stage except segment 3 bearing 11 setae.

ANTENNA II (Figure 3h).—Similar to that of

adult: segment 1 very short, naked, about one-third

length of following segment; segment 2 bearing palp

with one subterminal and three terminal setae; segment 3 with three groups of spinules—one proximally, another at midlength, and third subterminally—also one subterminal and three terminal setae.

MANDIBLE (Figure 4a).—As in adult.

MAXILLAE AND MAXILLIPEDS (Figure 4e,k,p).—

Similar to those of adult.

FIRST LEG2 (Figure 5a).—Basipodite with lateral

protuberance bearing-short seta; preaxial surface with

two groups of spinules. Exopodite one-segmented with

three lateral spines and three terminal setae; groups

of spinules present at bases of lateral spines; distal

row of spinules extending obliquely nearly to mesiodistal border of preaxial surface. Endopodite onesegmented, bearing four spinules on lateral border,

group of spinules proximally on mesial border, two

terminal setae, and short subterminal spine on mesial

border.

SECOND LEG (Figure 5h).—Basipodite bearing short

lateral seta with group of spinules at base and row of

spinules near mesiodistal border. Exopodite one-segmented, bearing two lateral spines, both as long as

exopodite, one subterminal lateral spine slightly longer

than exopodite, one laterodistal seta more than twice

length of exopodite, and mesiodistal seta more than

three times length of exopodite. Endopodite onesegmented, bearing four lateral spinules, two terminal

setae, one subterminal seta on mesial border, and two

groups of spinules on proximomesial border.

THIRD LEG (Figure 2a).—Represented by lobe with

three setae on second free segment.

Second Copepodid Stage

FIGURES 2C; 3b,i; 4b,f,l; 5b,i; 6a

Second copepodid differing from first in following

respects:

Body consisting of cephalothorax and five free

segments.

ANTENNA I (Figure 3b).—Five-segmented.

FIRST LEG (Figure 56).—Exopodite two-segmented:

segment 1 bearing one laterodistal spine; segment 2

with lateral margin bearing one spine at midlength,

one subterminally, one terminally, and two terminal

setae. Endopodite two-segmented: segment 1 bearing

oblique row of spinules on lateropreaxial surface and

3

 Each pair of legs joined together by copula extending

between coxopodites. Exopodites of all legs have group of

spinules situated at bases of lateral spines, and all segments

of both exopodites and endopodites have oblique subterminal row of spinules extending onto preaxial surface.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Elaphoidella bidens coronata: a, first copepodid; b, posterior of first copepodid,

lateral view; c, second copepodid; d, third copepodid; e, fourth copepodid; /, fifth copepodid;

g, posterior of adult female, dorsal view; h, posterior view of adult female, lateral view; i, posterior of adult male, dorsal view; j , posterior of adult male, lateral view.
NUMBER 1 1 9

FIGURE 3.—Elaphoidella bidens coronata, a—g, antenna 1: a, first copepodid; b, second copepodid; c, third copepodid; d, fourth copepodid; e, fifth copepodid; /, adult male; g, adult

female, h-m, antenna 2: h, first copepodid; i, second copepodid; /, third copepodid; k, fourth

copepodid; I, fifth copepodid; m, adult female.

group of very small spinules on mesial border at midlength; segment 2 with oblique row of spinules on

lateropreaxial surface and one spine, one very long

seta, and one short spine terminally.

SECOND LEG (Figure 5i).—Exopodite two-segmented: segment 1 bearing lateral spine at midlength; segment 2 with one lateral spine, one mesial

spine, one subterminal spine, and two terminal setae.

Endopodite two-segmented: segment 1 naked; segment 2 with four spinules on lateral border, one terminal spine and one terminal seta, and mesial border

with one subterminal spine and short spine near midlength.

THIRD LEG (Figure 6a).—Basipodite bearing one

lateral seta. Exopodite one-segmented, with two lateral spines, subterminal spine, terminal spine, and

terminal seta. Endopodite one-segmented, bearing

three terminal spines.

FOURTH LEG (Figure 2c).—Represented by lobe

bearing four spines.

Third Copepodid Stage

FIGURES Id; 3c,;; 4g; be,}; 6b,g

Third copepodid differing from second in following

respects:

Body consisting of cephalothorax and six free segments.

FIRST LEG (Figure 5c).—Basipodite with additional

spine on mesiodistal border.

SECOND LEG (Figure 5;).—Segment 2 of exopodite

bearing additional lateral spine slightly smaller than,

and proximal to, lateral spine present in second copepodid stage, and additional very small spine on mesial

border. Segment 1 of endopodite with small spine on

mesial border.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 4.—Elaphoidella bidens coronata, a-d, mandible: a, first copepodid; b, second copepodid; c, fourth copepodid; d, adult female, e—j, maxilla 1: e, first copepodid; /, second copepodid; g, third copepodid; h, fourth copepodid; i, fifth copepodid; j , adult female, k—o,

maxilla 2: h, first copepodid; /, second copepodid; m, fifth copepodid; n, adult male; o, adult

female, p-s, maxilliped: p, first copepodid; q, fourth copepodid; r, fifth copepodid; s, adult

female.

THIRD LEG (Figure 6b).—Exopodite two-segmented; segment 1 bearing lateral spine and oblique

row of spinules distally; segment 2 with one lateral

spine, one subterminal spine, two terminal setae, and

two mesial spines. Exopodite two-segmented: segment 1 naked; segment 2 with three terminal setae

and subterminal spine on mesial border.

FOURTH LEG (Figure 6g).—Exopodite one-segmented, bearing two lateral spines and three terminal

spines. Endopodite one-segmented, with two terminal

setae.

FIFTH LEG (Figure 2d).—Represented by small

lobe on caudolateral margin of fourth free segment

bearing two setae.

Fourth Copepodid Stage

FIGURES 2e; 3d,k; 4c,h,q; 5d,k; 6c,h,l

Fourth copepodid differing from third in following

respects:

Body consisting of cephalothorax and seven free

segments.
NUMBER 1 19

FIGURE 5.—Elaphoidella bidens coronata, a-g, first leg: a, first copepodid; b, second copepodid; c, third copepodid; d, fourth copepodid; e, fifth copepodid; /, adult male; g, adult female, h—n, second leg: h, first copepodid; i, second copepodid; ;', third copepodid; k, fourth

copepodid; /, fifth copepodid; m, adult male; n, adult female.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ANTENNA I (Figure 3d).—Six-segmented.

FIRST LEG (Figure 5d).—Segment 2 of exopodite

bearing spine on mesial border at midlength. Both

segments of endopodite with mesial spine.

THIRD LEG (Figure 6c).—Segment 2 of exopodite

bearing additional spine on lateral border and one

spine on mesial border.

FOURTH LEG (Figure 6h).—Exopodite two-segmented: segment 1 with one spine laterally and row

of spinules distally; segment 2 bearing two lateral

spines, one distal spine, two terminal setae, one subterminal mesial spine, and two small mesial spines.

Endopodite two-segmented: segment 1 naked; segment 2 with one terminal spine, one terminal seta,

and one small mesial spine.

FIFTH LEG (Figure 6/).—Represented by three

lobes; mesial lobe with one small spine, large lobe

lateral to it with two spines and three setae, and lateral lobe bearing long seta.

seta on mesial border, and one mesial spine near

midlength. Endopodite two-segmented: segment 1

bearing one small mesiodistal spine; segment 2 with

two additional mesial spines.

FOURTH LEG (Figure 6i).—Exopodite three-segmented: segment 2 bearing two subterminal spines,

one laterally and one mesially, and distal row of

spinules; segment 3 with one lateral spine, one distolateral spine, one terminal spine, one terminal seta,

one distomesial spine, and one mesial spine at midlength. Endopodite two-segmented: segment 2 bearing one terminal spine, one terminal seta, and two

very thick mesial spines.

FIFTH LEG (Figure 6m).—Three lobes large, appearing as distinct segments, mesial lobe bearing three

additional spines.

SIXTH LEG (Figure 6m).—Represented by distinct

lobe bearing two setae on fifth free segment. Leg not

discernible in adult female.

Fifth Copepodid Stage

FIGURES 2/; 3e,l; M,m,r; 5e,l; 6d,i,m

Fifth copepodid differing from fourth in following

respects:

Body consisting of cephalothorax and eight free

segments.

FIRST LEG (Figure 5^).—Exopodite three-segmented: segment 1 bearing lateral spine and semicircular row of subterminal spinules; segment 2 with

lateral spine, row of spinules, and one seta on mesiodistal border; segment 3 bearing lateral subterminal

spine, terminal spine, and two terminal setae. Endopodite two-segmented, one-fourth longer than exopodite: segment 2 with additional small spine on

mesial border.

SECOND LEG (Figure 5/).—Exopodite three-segmented; segment 2 bearing two subterminal spines,

one laterally and one mesially, and distal row of

spinules; segment 3 with one lateral, one mesial, and

one subterminal spine and two terminal setae. Endopodite two-segmented: segment 2 with additional

spine on mesial border.

THIRD LEG (Figure 6d).—Exopodite three-segmented: segment 2 bearing two subterminal spines,

one laterally and one mesially, and row of subterminal

spinules; segment 3 with one lateral spine, one subterminal spine, two terminal setae, one subterminal

Bryocamptus zschokkei alleganiensis Coker

First Naupliar Stage

FIGURE 7<Z

Dorsal shield subcircular, smooth, with anterior portion often broken in exuviae.

Labrum subspatulate, very narrow anteriorly. Narrow portion lying between bases of first antennae,

margins diverging posterolaterally posterior to antennae; fine hairs bordering posterior margin and with

tufts of longer hairs on median line.

ANTENNA I.—Segment 1 short and naked; segment

2 about twice length of segment 1, bearing three

subterminal spinules and three setae, proximal and

median setae short and delicate, terminal seta rather

heavy and about twice length of segment 2; segment

3 about two-thirds length of segment 2 bearing two

long terminal setae, one subterminal seta about half

length of terminal setae, and two groups of spinules,

one group situated at midlength, other subterminally.

ANTENNA II.—Coxopodite broader than long with

semicircular row of spinules on lower distal surface;

proximopostaxial angle produced into curved acute

process bearing hairs along postaxial margin from

apex to beginning of curve, process projecting dorsally above posterior portion of labrum. Basipodite

wider than long, narrowing postaxially forming short
NUMBER 1 1 9 11

FIGURE 6.—Elaphoidella bidens coronata, a-f, third leg: a, second copepodid; b, third copepodid; c, fourth copepodid; d, fifth copepodid; e, adult male; /, adult female, g-k, fourth leg:

g, third copepodid; h, fourth copepodid; i, fifth copepodid; j , adult female; k, adult male.

l-m, fifth leg and sixth lobe: /, fourth copepodid; m, fifth copepodid. n-o, fifth leg: n, adult

male; o, adult female, p, spermatophore; q, seminal receptacle.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

protuberance bearing three terminal setae. Exopodite

three-segmented: segment 1 subequal in length to

basipodite, bearing one short postaxial seta at midlength and terminal seta nearly four times length of

segment; segment 2 one-half length of segment 1 and

bearing one distal seta subequal in length to exopodite; segment 3 two-thirds length of segment 1, bearing one short subterminal seta and another terminally

about one and one-half times length of exopodite.

Endopodite one-segmented with group of spinules

near distal end, very delicate seta at midlength, and

two terminal setae; one of latter very short and narrow, other almost as long as endopodite, very stout,

and articulated basally.

MANDIBLE.—Median border of coxopodite produced into rounded lobe. Subrectangular basipodite

bearing semicircular row of spinules preaxially and

plumose seta on proximomesial surface. Exopodite

two-segmented: segment 1 slightly longer than segment 2, bearing small spine at midlength and seta

terminally; segment 2 about one-third length of segment 1, bearing short subterminal seta and another

terminally about three-fourths length of entire animal. Endopodite one-segmented with two curved

spines and four setae.

Ventral plate about one-third wider than long,

bearing two rows of long, slender spinules across ventral surface, one row near midlength and other near

caudal margin. Caudal margin with paired protuberances, each terminating in single fine seta.

Second Naupliar Stage

FIGURE 7b

Second nauplius differing from first in following respects :

ANTENNA I.—Segment 3 with additional group of

spinufes and single short, stout, tufted seta subterminally.

ANTENNA II.—Coxopodite bearing protuberance

with small tuft of hairs on postaxial margin of postawal m*mmam. Exopodite with additional small seta

«n preaxial surface of segment 1 near midlength.

EruAopodite one-segmented with additional group of

spinuies proximally.

MANDIBLE.—Endopodite with one additional spine

lateral to two present in first nauplius.

MAXILLA I.—Represented by pair of plumose setae

on anterior third of ventral plate.

Caudal portion of ventral plate emarginate and

posteromedian row of spinules lacking but replaced

by short lateral spinules.

Third Naupliar Stage

FIGURE 7c,g

Third nauplius differing from second in following

respects:

ANTENNA I.—Segment 3 with two additional

small setae at midlength.

ANTENNA II.—Protuberance on postaxial margin

of postaxial extension of coxopodite with one plumose

seta instead of tuft of hairs found in second nauplius.

Segment 3 of exopodite with additional slender seta

immediately proximal to, and about twice as long as,

subterminal seta.

Caudal margin of ventral plate bearing second pair

of setae anteromesial to first pair about one-third

their length, and pair of protuberances mesial to

setae. Plate considerably lengthened with posterior

region of body extending well beyond dorsal shield.

Fourth Naupliar Stage

FIGURE Id

Fourth nauplius differing from third in following

respects:

ANTENNA I.—One additional seta on segment 3.

MAXILLA I.—Represented by bifurcate lobe with

two wtae on mesial lobe and one very small seta on

smaller lateral lobe.

Posterior region of body lengthened, its caudal

margin bearing two delicate setae lateral to setae present in third naupliar stage; median protuberances of

third naupliar stage lacking. Anal plate with very fine

spinuies along posterior border.

Fifth Nmupiimr Stmge

FIGU«E 7e

Fifth aauplius differing from fourth in following

respects:

ANTENNA I.—An additional very small seta present

on segment 3.

MAXILLA I.—Setae larger and lobe more prominent

and two-segmented.

MAXILLA II.—Represented by small lobe caudal to

maxilla I.
NUMBE R 11 9 13

FIGURE 7.—Bryocamptus zschokkei alleganiensis: a, first nauplius, ventral view; b, second

nauplius, ventral view; c, third nauplius, ventral view; d, fourth nauplius, ventral view; e,

fifth nauplius, ventral view; /, sixth nauplius, ventral view; g, third nauplius, lateral view.

Setae on caudal margin of ventral plate larger;

one additional small seta present anteromesial to

most lateral seta of distal group.

Sixth Naupliar Stage

FIGURE If

Sixth nauplius differing from fifth in following respects :

MANDIBLE.—Usually no apparent difference but in

one shed (with distinct intersegmental sutures across

entire body), coxopodite with teeth on median lobe

as in copepodids and adult.

MAXILLA I.—Lobes distinct: intersegmental suture

present laterally between lobes of first maxilla and

those of second maxilla.

Body posterior to maxillae divided laterally into

two distinct segments with rudiments of first two pairs

of swimming legs and pygidial area.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

First Copepodid Stage

FIGURES 8a; 9e; \0a,h

Body consisting of cephalothorax and four free segments.

ANTENNA I (Figure 9e).—As in sixth nauplius but

segment 2 with distal lobe bearing one seta and one

asthete.

FIRST LEG (Figure 10a).-—Basipodite bearing lateral protuberance with one spine and two rows of

spinules on lateral surface. Exopodite one-segmented,

bearing two lateral spines, one subterminal spine, one

terminal spine, and one terminal seta. Groups of

spinules present at bases of lateral spines, and distal

row of spinules extending obliquely on preaxial surface. Endopodite one-segmented, with four pairs of

spinules along lateral border, row of subterminal

spinules, one terminal spine, and one terminal seta.

SECOND LEG (Figure 10/z).—Basipodite bearing one

lateral spine with group of spinules at base and row

of spinules near mesiodistal border. Exopodite onesegmented, bearing two lateral and two terminal

spines. Endopodite one-segmented, with four pairs of

spinules along lateral border, one terminal spine, and

one terminal seta.

THIRD LEG (Figure 8a).—Represented by caudolateral lobe with two spines on second free segment.

FIGURE 8.—Bryocamptus zschokkei alleganiensis: a, first copepodid; b, second copepodid;

c, third copepodid; d, fourth copepodid; e, fifth copepodid; /, posterior of adult female, dorsal

view; g, posterior of adult male, dorsal view. Figures 8a-e are on scale indicated on Figure 8a.
NUMBER 1 19 15

Second Copepodid Stage

FIGURE S 86 ; 9/ ; \Ob,i; ll a

Second copepodid differing from first in following

respects:

Body consisting of cephalothorax and five free segments.

ANTENNA I (Figure 9/).—Five-segmented.

FIRST LEG (Figure 10b).—Basipodite bearing additional spine on mesiodistal surface. Exopodite twosegmented: segment 1 with lateral spine near distal

end; segment 2 with lateral spine at midlength, one

subterminal spine, one terminal spine, and two terminal setae. Endopodite two-segmented: segment 1

bearing row of spinules on lateral border; segment 2

with lateral row of spinules, one terminal spine and

terminal seta.

SECOND LEG (Figure 10z').—Exopodite two-segmented: segment 1 with laterodistal spine; segment 2

bearing lateral spine, two terminal spines, and one

small terminal seta. Endopodite two-segmented: segment 1 naked; segment 2 with three pairs of spinules

on lateral border, one subterminal spine, and two

terminal setae.

THIRD LEG (Figure lla).—Basipodite bearing one

lateral seta and laterodistal group of spinules. Exopodite one-segmented, bearing lateral spine near midlength, one subterminal spine, and three terminal

spines. Endopodite one-segmented, with four lateral

spinules and three terminal spines.

FOURTH LEG (Figure 8b).—Represented by lobe

bearing two spines and one seta.

Third Copepodid Stage

FIGURE S 8C ; 9g; 10c,; ; llb,g

Third copepodid differing from second in following

respects:

Body consisting of cephalothorax and six free segments.

SECOND LEG (Figure 10;').—Segment 2 of exopodite with one additional lateral spine proximal to

lateral spine present in second copepodid, one additional very small spine on mesial border, and mesioFIGURE 9.—Bryocamptus zschokkei alleganiensis: a, adult female, antenna 2; b, adult female,

mandible: c, adult female, maxilliped; d, spermatophore. e-j, antenna 1: e, first copepodid;

/, second copepodid; g, third copepodid: h, fourth copepodid; i, fifth copepodid; ;'. adult

female.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

distal seta more than four times length of that of

second copepodid.

THIRD LEG (Figure lib).—Exopodite two-segmented : segment 1 bearing laterodistal spine; segment 2 with one lateral spine, one subterminal spine,

one terminal spine, one terminal seta, and one small

mesial spine. Endopodite two-segmented: segment 1

naked; segment 2 bearing one subterminal spine, two

terminal setae, and two lateral spinules.

FOURTH LEG (Figure llg).—Exopodite one-segmented, with one lateral spine and four terminal

spines. Endopodite one-segmented, bearing two terminal spines and one terminal seta.

FIFTH LEG (Figure 8c).—Represented by small

lobe bearing two small spines on caudolateral margin

of fourth free segment.

Fourth Copepodid Stage

FIGURES 8d; 9h; \Od,k; llc,h,j

Fourth copepodid differing from third in following

respects:

Body consisting of cephalothorax and seven free

segments.

ANTENNA I (Figure 9h).—Six-segmented.

FIRST LEG (Figure 10c?) .—Segment 2 of exopodite

bearing one spine on mesial border at midlength.

Segment 2 of endopodite with two stout terminal

spines.

THIRD LEG (Figure lie).—Segment 2 of endopodite bearing two additional terminal setae, subterminal

spine enlarged into very stout, terminally rounded

spine equivalent in length to that of segment.

FIGURE 10.—Bryocamptus zschokkei alleganiensis, a-g, first leg: a, first copepodid; b, secotid

copepodid; c, third copopodid; d, fourth copepodid; e, fifth copepodid; /, adult male; g, adult

female, h-n, second leg: h, first copepodid; i, second copepodid; ;, third copepodid; k, fourth

copepodid; I, fifth copepodid; m, adult male; n, adult female.
NUMBER 1 1 9 17

FOURTH LEG (Figure \lh).—Exopodite two-segmented: segment 1 bearing one laterodistal spine;

segment 2 with two lateral spines, two distal spines.

one terminal seta, and one subterminal mesial spine.

Endopodite two-segmented: segment 1 naked; segment 2 bearing one subterminal spine, one terminal

spine, and one terminal seta.

FIFTH LEG (Figure 11;).—Represented by three

lobes: mesial lobe bearing two small spines, large

lobe lateral to it with three spines and two setae, and

lateral lobe bearing single long seta.

SIXTH LEG (Figure 11/).—Represented by pair of

small lobes on caudolateral margins of fifth free segment, each bearing one spine and one seta.

Fifth Copepodid Stage

FIGURES 8«; 9«; lOe,l; \\d,i,k,l

Fifth copepodid differing from fourth in following

respects:

FIGURE 11.—Bryocamptus zschokkei alleganiensis, a-f, third leg: a, second copepodid; b, third

copepodid; c, fourth copepodid; d, fifth copepodid; e, adult female; /, adult male, g-i, fourth

leg: g, third copepodid; h, fourth copepodid; i, fifth copepodid. ;, fourth copepodid, fifth and

sixth legs; k, fifth copepodid, fifth legs; /, fifth copepodid, sixth leg; m, adult male, fifth and

sixth legs; n, adult female, fifth leg.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Body consisting of cephalothorax and eight free

segments.

FIRST LEG (Figure lOe).—Basipodite bearing one

mesiodistal spine. Exopodite three-segmented: segment 2 with one laterodistal spine and one mesiodistal

spine; segment 3 with one subterminal spine, one terminal spine, and two terminal setae. Endopodite twosegmented: segment 1 bearing one mesial spine,

group of spinules on mesial border proximal to spine,

and three pairs of spinules on lateral border; segment

2 with one terminal spine, two terminal setae, one

spine on mesial border at midlength, and two spinules

on lateral border at midlength; segment appearing

to be divided at midlength.

SECOND LEG (Figure 10/).—Exopodite three-segmented: segment 1 bearing one laterodistal spine and

one mesiodistal spine; segment 2 with one laterodistal

spine and one mesiodistal spine; segment 3 bearing

one lateral and one subterminal spine, two terminal

spines, one terminal seta, and one mesial spine. Endopodite

 3 three-segmented: Segment 1 bearing one

mesiodistal spine; segment 2 with one laterodistal and

one mesiodistal spine; segment 3 bearing two subterminal spinules, one terminal spine, two terminal

setae, and one mesial spine at midlength.

THIRD LEG (Figure lid).—Exopodite three-segmented: segment 2 bearing one laterodistal and one

mesiodistal spine; segment 3 with two lateral spines,

one subterminal spine, one terminal spine, one terminal seta, one subterminal mesial spine, and one

small spine on mesial border at midlength. Endopodite three-segmented: segment 1 bearing one mesiodistal spine; segment 2 with one laterodistal spine;

segment 3 bearing one lateral and two mesial spinules,

stout subterminal spine longer, and two terminal setae

thicker at their bases.

FOURTH LEG (Figure lli).—Exopodite three-segmented: segment 2 bearing one laterodistal and one

mesiodistal spine; segment 3 with one lateral spine,

one subterminal spine, two terminal spines, one terminal seta, and two mesial spines. Endopodite twosegmented: segment 1 bearing two lateral spinules,

3

 Adult females from the same locality also have threesegmented endopods on the second and third legs; however,

from other localities, some adults have two-segmented endopods on both second and third legs while others have the

second endopod two-segmented and the third three-segmented. Similar variations have been observed in the adults

by Kiefer (1929:321) and Gurney (1932:148).

two terminal spines, one terminal seta, and two mesial

spines.

FIFTH LEG (Figure Ilk).—Two mesial lobes larger

and more distinct.

SIXTH LEG (Figure 11/).—Represented by pair of

small lobes on caudolateral margins of fifth free segment, each lobe bearing one spine and two setae. Leg

not present in adult female.

Moraria virginiana Carter

First Naupliar Stage

FIGURE \2a,b

First nauplius inflated, bearing subcircular dorsal

shield characteristically ornamented: one dorsomedian lobe with four points projecting caudodorsally,

one pair of pointed lateral lobes, and one pair of lobes

posterior to median lobe.

Labrum subspatulate, very narrow anteriorly with

paired lobes on anterolateral angles, posterior margin

bordered by fine hairs with tuft of longer hairs projecting from lobe on median line. Narrow portion

situated between bases of antennae, from which margins diverging posterolaterally.

ANTENNA I.—Segment 1 short and naked. Segment 2 about twice length of segment 1, bearing distal group of long spinules, proximal group of shorter

spinules, and three setae: distal seta long and slender,

remaining two short, stout, and tufted with fine hairs

distally. Segment 3 about two-thirds length of segment 2, bearing two terminal setae, one subterminal

seta, and two subterminal groups of very long

spinules.

ANTENNA II.—Coxopodite broader than long,

bearing semicircular row of spinules on ventral surface; proximopostaxial angle produced into slightly

curved extension ending in tuft of hairs, extension also

bearing protuberance with one plumose seta. Basipodite wider than long, narrowing postaxially forming short protuberance bearing two setae. Exopodite

three-segmented: segment 1 with one preaxial seta;

segment 2 bearing one subterminal seta on short protuberance; segment 3 with two terminal setae. Endopodite one-segmented, bearing one delicate seta at

midlength, group of subterminal spinules, and two

terminal setae: one short and slender, other almost as

long as segment, stout, and movably articulated with

segment.
NUMBER 1 1 9 19

MANDIBLE.—Preaxial median border of coxopodite

produced into small rounded lobe. Basipodite bearing row of spinules on lower surface and one plumose

seta on proximal lower surface. Exopodite two-segmented: segment 1 slightly longer than basipodite,

bearing one small spine at midlength and one terminal seta; segment 2 with one short subterminal

seta and one terminal seta about twice length of

exopodite. Endopodite one-segmented, bearing two

curved spines and several delicate setae.

Ventral plate bearing two rows of very long slender

spinules across ventral surface, one row near midlength and other near caudal margin. Caudal margin

rounded but bearing paired lateral protuberances

each with single fine seta.

Second Naupliar Stage

FIGURE \2c,d

Second nauplius differing from first in following

respects:

ANTENNA I.—Segment 3 of first antenna bearing

one additional subterminal, very stout, and tufted

seta.

ANTENNA II.—Endopodite with additional group

of spinules proximally.

MANDIBLE.—Basipodite with one additional seta on

mesial border.

MAXILLA I.—Represented by one pair of lateral

lobes on anterior surface of ventral plate.

Caudal portion of ventral plate emarginate, median

posterior row of spinules replaced by two groups of

very long, slender, hairlike spinules situated at base of

caudal protuberances.

Third Naupliar Stage

FIGURE \2e,f

Third nauplius differing from second in following

respects:

ANTENNA I.—Segment 3 bearing one additional

seta.

ANTENNA II.—Segment 1 of exopodite bearing one

additional seta. (Endopodite absent in single test

examined.)

MAXILLA I.—Each lobe present in second nauplius

bearing one plumose seta.

Caudal margin of ventral plate with one pair of

protuberances mesial to lobes present in second nauplius, median anterior row of spinules replaced by

lateral spinules.

Fourth Naupliar Stage

FIGURE \2g,h

Fourth nauplius differing from third in following respects :

ANTENNA II.—Postaxial extension of coxopodite

bearing one additional lobe, and lobe present in third

nauplius with one plumose seta.

MAXILLA I.—Bearing one additional seta. Intersegmental suture present posterior to maxillary lobes.

MAXILLA II.—Represented by one pair of indistinct lobes anterior to intersegmental suture.

MAXILLIPED.—Represented by one pair of indistinct lobes posterior to maxillary lobes.

Caudal portion of body produced into two large

prominences corresponding to caudal rami of adult,

each prominence bearing four setae: lateral and

mesial setae on caudal margin, remaining two setae

on lobe just anteroventral to posterior margin.

Fifth Naupliar Stage

FIGURE \2ij

Fifth nauplius differing from fourth in following

respects:

ANTENNA I.—Segment 3 bearing nine setae, two of

which short, blunt, and plumose.

LABRUM.—Broader posteriorly than anteriorly,

broader portion bearing short hairs along lateral and

caudal margins.

MAXILLA I.—Lobes more prominent, setae longer,

and one additional very small spinule present mesial

to other two setae.

Caudal margins of caudal prominences directed

caudolaterally instead of caudally, resulting in four

setae on each side appearing on lateral instead of

posterior border.

Sixth Naupliar Stage

FIGURE 12*,/

Sixth nauplius differing from fifth in following respects:

ANTENNA I.—Additional small seta on segment 3.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 12.—Moraria virginiana: a, first nauplius, ventral view; b, first nauplius, dorsal view;

c, second nauplius, dorsal view; d, second nauplius, ventral view; e, third nauplius, ventral

view; /, third nauplius, posterior part of dorsal shield; g, fourth nauplius, ventral view;

h, fourth nauplius, dorsal view; i, fifth nauplius, ventral view; j , fifth nauplius, posterior part

of dorsal shield; k, sixth nauplius, posterior part of dorsal shield; /, sixth nauplius, ventral view.

MAXILLA I.—Additional seta lateral to two setae

present in fifth nauplius.

MAXILLA II.—Lobes more prominent and intersegmental suture posterior to lobes no longer evident.

MAXILLIPED.—Represented by two pairs of lobes:

one pair near median line, and other situated lateral

to first pair and just mesial to second maxillae.

Posterior part of body divided, by incomplete in-
NUMBER 1 19 21

tersegmental sutures, into two segments: anterior with

rudiments of first two pairs of swimming legs, and

posterior constituting pygidial area. Setae of large

caudal prominences longer.

First Copepodid Stage

FIGURES 13a,b; \Aa,d

In the two available tests of the first copepodid, the

cephalic appendages are so poorly preserved that they

cannot be described or figured.

Body consisting of cephalothorax and four free

segments.

FIRST LEG (Figure 14a).—Basipodite bearing one

lateral spine and three rows of spinules on distal border. Exopodite one-segmented, bearing two lateral

spines, one subterminal spine, one terminal spine, and

two terminal setae. Endopodite one-segmented, bearing subterminal spine, one terminal spine, one terminal seta, and two lateral spinules.

SECOND LEG (Figure 14<i).—Basipodite bearing one

lateral spine with group of spinules at its base and

one row of spinules on its distal border. Exopodite

one-segmented, with one lateral spine, one subterminal spine, and three terminal spines. Endopodite onesegmented, bearing three terminal spines.

THIRD LEG (Figure 13a).—Represented by one pair

of caudolateral lobes on second free segment, each

lobe bearing two setae. Caudal segment with ventral

surface bearing two rows of spinules extending from

lateral border almost to median line: dorsal surface

with slightly rounded and toothed anal plate.

FIGURE 13.—Moraria virginiana: a, first copepodid; b, posterior part of first copepodid, lateral

view; c, second copepodid, lateral view; d, posterior part of second copepodid, dorsal view;

e, adult male, lateral view.
22 SHITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Caudal ramus with dorsal protuberance bearing

one slender seta about three times length of ramus,

two lateral setae, one row of subterminal spinules.

and three terminal setae: mesial seta very short,

middle one about half length of entire body, and

lateral seta half length of middle one.

Second Copepodid Stage

FIGURES 13c,d; l-ib,e,f,i

Body consisting of cephalothorax and five free segments.

ANTENNA I.—Five-segmented: segment 1 bearing

one seta; segment 2 with five setae and one esthete;

remaining three segments bearing total of twelve

setae.

FIRST LEG (Figure 146).—Basipodite bearing one

laterodistal and one mesiodistal spine, with spinules

at bases of each, and two rows of subterminal spinules. Exopodite two-segmented: segment 1 bearing

one lateral spine near distal end; segment 2 with one

lateral spine at midlength, one subterminal spine, one

terminal spine, and two terminal setae. Endopodite

two-segmented: segment 1 bearing one subterminal

row of spinules: segment 2 with two terminal spines

and one terminal seta.

SECOND LEG (Figure 14^).—Basipodite bearing one

lateral spine near distal border with group of spinules

FIGURE 14.—Moraria virginiana, a-c, first leg: a, first copepodid; b, second copepodid; c, adult

male, d—e, second leg: d, first copepodid; e, second copepodid. f-g, third leg: /, second

copepodid; g, adult male, h, adult male, fourth leg. i-j, antenna 1: i, second copepodid;

;, adult male, k, posterior part of adult male, dorsal view; I, adult male, mandible; m, adult

male, maxilliped; n, adult male, maxilla 2; o, adult male, antenna 2; p, adult male, fifth leg.
NUMBER 1 1 9 23

at its base and one row of subterminal spinules. Exopodite two-segmented: segment 1 bearing one laterodistal spine; segment 2 with one subterminal spine

and three terminal spines. Endopodite two-segmented: segment 1 naked; segment 2 bearing one

lateral spinule and three terminal spines.

THIRD LEG (Figure 14/).—Basipodite bearing one

lateral spine with group of spinules at its base and

one subterminal row of spinules. Exopodite one-segmented, bearing one lateral spine and four terminal

spines. Endopodite one-segmented, bearing three terminal spines and three subterminal spinules.

FOURTH LEG (Figure 13c).—Represented by lobe

bearing two setae.

Anal plate more round than that of first copepodid.

teeth on margin less prominent.

Caudal rami similar to those of first copepodid but

innermost terminal seta much longer.

Discussion

Most early reports on the stages in postembryonic development of harpacticoids are based on the association of one or two larval stages with adults, and few

investigators attempted to rear an individual from

the first nauplius to the first copepodid stage as did

Dietrich (1915) for Canthocamptus staphylinus.

More recently (Fraser, 1936; Johnson and Olson,

1948; and Haq, 1965), attempts were made at rearing the larvae of harpacticoid copepods to study their

development, and some success was achieved during

the present study in recording the development of a

single individual.

The Naupliar Stages

Several workers have reported five or fewer naupliar

stages for a number of harpacticoid copepods (Dietrich, 1915; Tesch, 1915; Chappuis, 1916; Brian,

1922; Borutskii, 1925; Gurney, 1932; and Fraser,

1936) but other work, much of it more recent, indicates six naupliar stages (Chappuis, 1916; Borutskii,

1925; Ewers, 1930; Johnson and Olson, 1948; Krishnaswamy, 1951: Grig, 1960; Fahrenbach, 1962; and

Haq, 1965).

Six naupliar stages are characteristic of the development of all three species considered in the present

study: Elaphoidella bidens coronata, Bryocamptus

zschokkei alleganiensis, and Moraria virginiana.

In the reports in which only five nauplii have been

observed in the life history of harpacticoids, there is

no uniformity as to which stage is lacking. This lack

of consistency seems particularly significant in view

of the following: (1) Few of the investigators have

based their reports on the exuviae of a single individual, and only a few workers have isolated and

reared the nauplii of a single species. (2) The fourth

and fifth stages are so similar that had not the exuviae

of an isolated individual been obtained in the present

study, these two nauplii probably would not have been

distinguished.

If it may be assumed that all freshwater harpacticoids have six naupliar stages in their life cycles and

that the features exhibited by each stage of the species

observed in the present study (Figures 1, 7, 12) are

characteristic, then in most instances it seems apparent that the fourth or fifth nauplius was overlooked

(e.g., Dietrich, 1915; Borutskii, 1925, and others);

however, in the description of the naupliar development of Canthocamptus pygmaeus, Gurney (1932:

89-92) probably failed to obtain the third nauplius.

In light of the careful work of Ewers and others,

my own observations, the fact that six nauplii are

characteristically present in the life cycles of all calanoids and many cyclopoids, and that few detailed

studies have been made of the naupliar development

of harpacticoids, it seems desirable that the postembryonic development of all species in which fewer

than six stages have been repoued should be reexamined.

Key to the Naupliar Larvae of Harpacticoids Having Five Molts Preceeding the

First Copepodid Larva

1. Caudal margin rounded and bearing row of hairs First Nauplius

Caudal margin not rounded and without row of hairs 2

2. Caudal margin emarginate and bearing groups of hairs on posterolateral margin.

Second Nauplius

Caudal margin cleft and bearing no hairs 3
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Naupliar Larvae of Harpacticoids Having Five Molts Preceeding the

First Copepodid Larva—Continued

3. Caudal margin with slight median cleft and with fingerlike lobe on each side of cleft.

Third Nauplius

Caudal margin with deep median cleft and fingerlike lobes lacking 4

4. Each caudal lobe bearing four setae Fourth Nauplius

Each caudal lobe bearing five setae 5

5. No trace of swimming legs Fifth Nauplius

Two pairs of swimming legs represented by setae-bearing lobes Sixth Nauplius

The naupliar stages of each of the three species of

copepods treated in this study are similar in most respects. Differences were noted in the second antennae

and in the development of the maxillae and of the

maxillipeds.

On antenna II, the postaxial extension of the

coxopodite of E. bidens coronata is dentate, but bears

hairs in B. zschokkei alleganiensis and M. virginiana.

This extension bears two lobes in M. virginiana but

only one in the other two species. The setae on the

protuberance of the basipodite vary in number: one

in E. bidens coronata, three in B. zschokkei alleganiensis, and two in M. virginiana. The first maxilla of

the second nauplius is evident in E. bidens coronata

and B. zschokkei alle ganiensis as a single seta; however, in M. virginiana, there is a distinct naked lobe,

and setae are not present until the third stage. The

second maxilla does not appear until the fifth stage

in E. bidens coronata and B. zschokkei alle ganiensis,

whereas there is a distinct lobe in the third stage of

M. virginiana. This lobe is more distinct in the fifth

stages of B. zschokkei alle ganiensis and M. virginiana

than in the sixth nauplius of E. bidens coronata.

The rudiments of the maxillipeds are scarcely evident until the sixth stage of E. biden; coronata, but

a lobe is present in the fifth stage of B. zschokkei

alleganiensis and in the fourth stage of M. virginiana.

The caudal margin of the third nauplius bears two

pairs of setae in B. zschokkei alleganiensis, but only

the original pair are present in that of the other two

species.

Except for slight variations and discrepancies between some accounts, there appears to be a remarkable degree of uniformity, even in details in the

naupliar development of freshwater harpacticoid copepods. In many instances, the similarities are so

marked that distinction between the early nauplii of

different species and even between some genera,

would appear to be difficult, if not impossible. Because the naupliar stages of so few species in a single

genus have been studied, it is impossible at present

to distinguish between the nauplii of different genera

with any degree of assurance.

Harpacticoid nauplii may be distinguished from

nauplii of other groups by the following characteristics which have been compiled from my work and the

studies of others: the first antennae directed laterally

when at rest; the large spatulate labrum; the stout,

prehensile seta on the endopodite of the second antenna; the pincerlike spines on the endopodite of the

mandible; and the long terminal spine on the exopodite of the mandible.

The Copepodid Stages

All free-living copepods, insofar as is known, undergo

six molts following the last naupliar stage, the last of

which gives rise to the adult. These latter immature

stages, the copepodid larvae, resemble the adult in

their general facies, and the adult cephalic appendages are only slightly modified from those of the first

copepodid. Some attempts have been made (Brian,

1922; Gurney, 1932; Krishnaswamy, 1951; and

others) to trace the later larval development of harpacticoids. Gurney's account of the copepodid development, as expressed in his chart, applies equally as

well to the two copepods, Elaphoidella bidens coronata and Bryocamptus zschokkei alleganiensis, of

which I have a complete series of stages, and to the

first two stages I obtained of M. virginiana.

The following chart presents a complete resume of

the pattern of development as it occurs in Elaphoidella bidens coronata, Bryocamptus zschokkei alleganiensis, and the first two stages of M. virginiana.

Although Gurney's report for C. staphylinus is not so

complete, there is agreement between the features he

mentioned and those cited here.
NUMBER 1 19 25

TABLE 1.—Segmentation in body and appendages of copepodids I—VI

Stage

Copepodid

segs. I

Body Ant.

segs.

Segments of exopodite Segments of endopodite

P. P.

I

II

III

IV

V

adult female

adult male .

12

13

14

15

16

16

17

rud.

1

2

2

3

3

3

rud.

1

2

3

3

3

1

2

2

2

2**

2**

rud.

1

2

2

2»*

2**

rud.

1

2

2

2

rud. rud.*

rud. rud.

present

present

* Sixth leg of fourth copepodid in E. bidens coronata represented by very small lobe.

** Endopodites of second and third legs of B. zschokkei alleganiensis sometimes consisting of three segments.

The copepodid stages can be distinguished by

counting the number of free segments; i.e., those not

included in the cephalothorax. The first copepodid

stage has four, the second five, the third six, the

fourth seven, the fifth eight, and the adult male nine.

The adult female has only eight visible segments;

although another is added posteriorly, the fourth and

fifth from the caudal end are fused, either partly or

completely. The fifth copepodid stage can be distinguished from the adult by the presence of a long segment at the posterior end of the abdomen, whereas

the long segment in the adult female is the fourth

anterior to the caudal rami.

The addition of segments in the first antenna progresses more irregularly so that while there are three

in the first copepodid larva, there are five in the second and third, six in the fourth and fifth, and eight

in the adult. It is also noteworthy that the addition

of segments in this appendage occurs both distally and

proximally. The fourth segment from the proximal

end in the adult, which may be recognized in all of

the stadia by the presence of a protuberance on which

is borne a seta and an esthete, corresponds to the second of the three-segmented appendage of the first

copepodid. Thus there are two additional segments

added basally: the first segment appears in the fourth

copepodid larva, and the second segment appears in

the adult.

Because of the difficulty of locating the exuviae in

the culture dishes, and because of the variation in the

rate of development, the time between each stage has

not been accurately determined. However, the time

required for the newly hatched nauplius to reach the

first copepodid stage is approximately 10 days, and

the time required for the first copepodid stage to

reach the adult is about 20 days.

Acknowledgments

It is with great pleasure that I express my appreciation to Dr. Horton H. Hobbs, Jr., for his help and

guidance, for his corrections of the manuscript, and

for his encouragement which has made this work

possible. I also wish to acknowledge my indebtedness

to the late Dr. B. D. Reynolds, Director of the Mountain Lake Biological Station, for sending me materials; to Dr. J. Rappaport of the Miller School of

Biology for translation of difficult passages in German; to Dr. Beatrice Nevins and Miss Betty Gunter

of Valdosta State College for their assistance in arranging the drawings and reading of the manuscript;

and to the faculty and members of the Miller School

of Biology for their help and encouragement.

Literature Cited

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Borutskii, E. V.

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Brian, A.

1922. L'AUheutha depressa Baird (copepodo harpacticoide) e i suoi larvali. Monitore Zoologico Italiano, 33:8-14.
26 SHITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Carter, Marjorie E.

1944. Harpacticoid Copepods of the Region of Mountain Lake, Virginia. Journal of the Elisha Mitchell Scientific Society, 60(2): 158-166, plates

65-67.

Chappuis, P. A.

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Coker, R. E.

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Kiefer, F.

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Boktryckeri, Lund. 1682 pages, 610 figures.

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