Redescription of Mesokalliapseudes crassus (Menzies, 1953) (Crustacea: Tanaidacea: Kalliapseudidae):...

Redescription of Mesokalliapseudes crassus (Menzies, 1953) (Crustacea:Tanaidacea: Kalliapseudidae): the first record of

a hermaphroditic kalliapseudid

D. T. Drumm* and R. W. Heard

(DTD) Department of Coastal Sciences, University of Southern Mississippi, Gulf Coast

Research Laboratory Campus, Ocean Springs, Mississippi 39566-7000,

e-mail: [email protected], [email protected]

Abstract.—The type species of the genus Mesokalliapseudes Lang 1956,

M. crassus (Menzies 1953), which was originally incompletely described and

illustrated, is redescribed and a key to the species of the genus is provided. A

study of type and non-type material indicates that M. crassus is a protandric

hermaphrodite. All of the adult specimens of M. crassus examined,including presumptive males and preincubatory and incubatory females,

had a male genital cone. In all other aspects, except for the development of

oostegites in the female phase, this species expresses no marked sexual

dimorphism.

Kalliapseudes crassus Menzies 1953 was

described based on 34 specimens, includ-

ing ovigerous females and ‘‘juveniles,’’

from San Quintin Bay, Lower (5Baja)

California, Mexico. Lang (1956) divided

Kalliapseudes into four subgenera based

on characteristics of the cheliped and

pereopods. At that time, he designated K.

crassus as the type species for the sub-

genus Mesokalliapseudes Lang 1956,

which is characterized by the dactylus of

the second and third pereopods having

a well-developed sensory organ and the

absence of an exopod on the cheliped and

first pereopod. In a recent treatment of

the genus Kalliapseudes, Gutu (2006)

elevated all four of Lang’s subgenera to

full generic rank and erected the genus

Alokalliapseudes to receive K. macsweenyi

Drumm, 2003, which had been placed in

the subgenus Mesokalliapseudes.

Examination of K. crassus type materi-

al from the Los Angeles County Museum

(LACM), and non-type material loaned

from the National Museum of Natural

History, Smithsonian Institution, Wash-

ington, D.C. (USNM), revealed that all

adult specimens appear to be hermaph-

roditic, i.e., ovigerous and preparatory

individuals possess a genital cone. Be-

cause Menzies (1953) provided only a brief

diagnostic description and limited illus-

trations for M. crassus, its redescription is

warranted, especially as it is the type

species for the genus. Gutu (2006) placed

five additional species, [M. viridis (Men-

zies, 1953), M. bahamensis (Sieg, 1982),

M. brasiliensis (Bacescu, 1986), M. sonia-

dawnae (Bamber, 1993), M. thalasispeleus

Gutu, 2006], within the genus Mesokal-

liapseudes.

Unless otherwise indicated the termi-

nology used here follows that of Larsen

(2003).

Family Kalliapseudidae Lang, 1956

Subfamily Kalliapseudinae Lang, 1956

Genus Mesokalliapseudes Lang, 1956

Diagnosis (modified after Gutu 2006).—

Antenna with peduncle having few simple*1 Corresponding author.

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON120(4):459–468. 2007.

setae on last article. Cheliped without

exopodite; propodus slender and very

long, much longer than carpus; fixed

finger shorter than dactylus. Pereopod 1

without exopodite. Dactylus of pereopods

2 and 3 long and thin with small outer

proximal digitiform prolongation having

several setae or ‘‘aesthetascs.’’ Dactylus of

pereopods 4 and 5 short and thick with

numerous aesthetascs; unguis absent. Pe-

reopod 6 with normal dactylus (claw-

shaped). Pleopod with exopodite biarticu-

late. Pleotelson with 2 terminal long

plumose setae. Male similar to female.

Mesokalliapseudes crassus

(Menzies, 1953)

Figs. 1–3

Kalliapseudes crassus Menzies, 1953:474–

479, figs. 17–20.—Lang, 1956:205, 207,

209, 223.—Bacescu, 1961:164.—Sieg

1982: 3.—Gutu, 2006:141.

Kalliapseudes (Mesokalliapseudes) cras-

sus Lang, 1956:211, 216, 217, fig. A4.—

Drumm, 2003:1, 2, 11.

Mesokalliapseudes crassus Gutu, 2006:

142.

Material examined.—Holotype: oviger-

ous hermaphrodite, 8 mm, San Quintin

Bay, lower (5Baja) California, Mexico, 7

Apr 1950, leg. Chas Horvath, Don Reish

and R. J. Menzies (LACM CR 1950-

076.1). Paratypes: 26 ovigerous and 2

preparatory hermaphrodites, 6–7 Apr

1950, depth 1.5–4.5 m, same locality and

collectors as holotype (LACM CR1950-

076.2). Non-types (USNM 262506): SQ

13: 1 ovigerous hermaphrodite, San Quin-

tin Bay, Baja California, Mexico, North

Pacific Ocean, 22 Apr 1960, leg. J.

Barnard, depth 0–13 m; SQ 28: 5 oviger-

ous, 2 preparatory (1 partially dissected)

and 1 juvenile, San Quintin Bay, Baja

California, Mexico, North Pacific Ocean,

21 Apr 1960, leg. J. Barnard, depth 5–7 m.

Diagnosis (adult).—Rostrum acute.

Medial extension of antenna with numer-

ous (12 or 13) plumose setae; second

(third?) article with distinctive narrow,

spinulate process. Cutting edge of fixed

finger of cheliped with numerous (,10)

bipectinate setae and cutting edge of

dactylus with numerous (,11) serrate setae

increasing in length distally; fixed finger

more than two-thirds length of dactylus.

Pereopod 6 propodus with numerous

(,12) outer serrate spiniform setae. Hypo-

sphaenia absent. Hermaphroditic.

Description (adult with oostegites and

genital cone).—Body length (Fig. 1A)

,8 mm. Carapace as broad as long, 1

pair of mid-lateral setae and few short

posterolateral setae; rostrum acute. All

pereonites lacking hyosphaenia. Pereo-

nites 4 and 5 (Fig. 1A) longest and

subequal; pereonites 1, 2, and 6 shorter

than pereonite 3 and subequal; 1 to several

simple setae on anterior corners and

several simple setae on posterior corner

of first 3 pereonites; genital cone (Fig. 2H)

on last pereonite. Pleonites (Fig. 1A)

subequal; epimera rounded, with several

plumose setae and 1 pair of simple

dorsolateral setae; hyposphaenia absent.

Pleotelson (Fig. 1A) posteriorly rounded,

with several lateral and posterior simple

setae and a few pair of dorsal simple setae;

2 long terminal plumose setae.

Antennule (Fig. 1B). First peduncle

article approximately twice as long as

second and third articles combined and

approximately 2.7 times as long as

maximum width, with spinules on inner

proximal margin and numerous simple

setae on outer margin. Inner flagellum

with 3 articles, distal article with 3 simple

setae and 1 broom seta; outer flagellum

with 9 articles, 1 aesthetasc each on

articles 4, 5, and 7, and 4 setae on distal

article.

Antenna (ovigerous adult; Fig. 1C).

First peduncle article with medial exten-

sion bearing 13 plumose setae. Squama

with 7 long simple setae. Second article

with structure bearing a narrow, spinulate

process (Fig. 1D). Third article with 1

460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

simple seta on inner margin. Last pedun-

cle article shorter than flagellum, with

numerous long, simple setae on inner

margin. Flagellum with 7 articles bearing

long, simple setae; distal article with 5

simple setae.

Labrum (Fig. 1E) subrectangular bear-

ing numerous hairs.

Mandibles (Fig. 1F, G). Right mandi-

ble (Fig. 1F) with tridentate incisor pro-

cess and setal row with 5 spiniform setae.

Left mandible (Fig. 1G) with approxi-

Fig. 1. Mesokalliapseudes crassus, adult with oostegites and genital cone (USNM 262506, SQ-28). A,

dorsal view of body; B, antennule; C, antenna (ovigerous hermaphrodite); D, spinulate process of antenna;

E, labrum; F, right mandible; G, incisor process, lacinia mobilis and setal row of left mandible; H, labium.

Scale bars: A 5 1.0 mm; B, C 5 0.2 mm; E, F, H 5 0.1 mm.

VOLUME 120, NUMBER 4 461


maxillule; B, maxilla; C, maxilliped; D, cheliped (inner surface); E, chela (inner surface); F, pereopod 1

(outer surface); G, pereopod 2 (outer surface); H, genital cone. Scale bars: A, B 5 0.1 mm; C, D, G 5

0.2 mm; F 5 0.4 mm.



pereopod 3 (outer surface); B, pereopod 4 (inner surface); C, merus and carpus of pereopod 5 (outer surface);

D, pereopod 6 (inner surface); E, pleopod; F, uropod. Scale bars: B, D, E 5 0.2 mm; F 5 0.1 mm.


mately 9 denticulations on incisor pro-

cess; lacinia mobilis with approximately 6

denticulations; setal row with 4 spiniform

setae. Mandibular palp uniarticulate with

row of long plumose setae and terminat-

ing in a long, naked spiniform seta.

Labium (Fig. 1H). Distal margin pro-

vided with short hairs and spinulate on

outer margin. Palp with long hairs and

terminating in a short, triangular spini-

form seta.

Maxillule (Fig. 2A). Inner endite bear-

ing 4 terminally setulate setae and dense

row of hairs on outer and inner margins.

Outer endite with 7 long serrate, 4 long

naked and 1 short spiniform setae, 2

subterminal setae and dense row of hairs

on outer and inner margins.

Maxilla (Fig. 2B). Inner lobe of fixed

endite with posterior row of 6 serrate

setae and with long anterior row of setae

(,45). Outer lobe of fixed endite with 3

serrate setae and several simple setae.

Inner lobe of moveable endite with

several simple setae; outer lobe of move-

able endite with 2 serrate and 3 plumo-

denticulate (bearing setules and denticles)

setae. Inner margin spinulate.

Maxilliped (Fig. 2C). Basal article

fringed with plumose setae on outer

margin, 3 simple and 1 plumose setae on

inner margin. First article of palp with

several simple setae on inner margin. Last

3 articles of palp with double row of long

plumose setae on inner margin; second

article with 1 distal simple setae on outer

margin; last article terminating in 2 long

simple setae. Endite with 20 pappose setae

along margin and 3 long setae distally; 2

coupling hooks.

Cheliped (Fig. 2D, E). Basis with 1

short simple ventrodistal seta. Merus with

1 long and 1 short simple setae. Carpus

approximately 2.5 times as long as broad,

with double row of long, plumose setae

and 4 short simple setae on dorsodistal

corner. Propodus slender and very long,

approximately 1.5 times as long as carpus,

with diagonal row of long plumose setae

on inner face; palm region with several

long simple and serrate setae; fixed finger

more than 0.67 length of dactylus, with

several simple setae; cutting edge with 10

bipectinate spiniform setae. Dactylus with

3 long setae on inner face; cutting edge

with 11 serrate spinules increasing in

length distally. Exopodite absent.

Pereopod 1 (Fig. 2F). Basis with acute,

proximal process, 2 short simple setae on

dorsal margin and 2 long simple setae on

ventrodistal corner. Ischium short and

naked. Merus longer than broad, shorter

than basis and approximately 3 times

length of carpus, with several simple setae

on ventral margin and spiniform seta

ventrodistally. Carpus short with 3 ven-

trodistal and 1 dorsodistal spiniform

setae, outer surface with row of small

tubercles. Propodus shorter than carpus,

5 ventral and 1 dorsodistal spiniform

setae; outer surface with small tubercles.

Dactylus represented by a sensory organ

with numerous distal sensory setae and

inner surface with 1 seta. Exopodite

absent.

Pereopod 2 (Fig. 2G). Basis more than

2.5 times as long as broad, several short

simple setae on proximal dorsal margin, 1

long and 2 short simple ventrodistal setae.

Ischium with 3 simple ventrodistal setae.

Merus approximately as long as carpus,

with several simple and 1 spiniform setae

on ventral margin. Carpus approximately

1.2 times as long as broad, with 6

spiniform setae and several long simple

distal setae. Propodus approximately 0.33

length of basis, with 5 spiniform setae on

ventral margin, 3 spiniform setae on outer

surface, 1 long simple seta and 1 long

serrate spiniform setae dorsodistally, 1

broom seta midway on dorsal margin.

Dactylus long and slender, shorter than

basis, with a thin outer digitiform lobe

terminating in 3 aesthetascs.

Pereopod 3 (Fig. 3A). Similar to pe-

reopod 2. Carpus with 10 spiniform setae.

Propodus with 5 spiniform setae on outer

surface.


Pereopod 4 (Fig. 3B). Basis approxi-

mately 3 times as long as broad with 2

simple setae on dorsal margin and 1 short

simple seta on ventrodistal corner. Ischi-

um with 1 long and 2 short simple setae.

Merus shorter than carpus with 1 distal

spinform seta, 4 simple setae on ventro-

distal corner and 1 spiniform seta on outer

surface. Carpus with 7 spiniform setae on

inner surface and 8 spiniform setae on

outer surface, 1 long simple dorsodistal

seta. Propodus with 10 spiniform setae on

each side increasing in length distally, 1

short pectinate seta on inner surface and 1

broom seta on dorsal margin. Dactylus

represented by a sensory organ with

numerous distal sensory setae.

Pereopod 5 (Fig. 3C). Similar to pereo-

pod 4. Merus with 2 distal and 2 inner

spiniform setae. Carpus with 8 spiniform

setae on outer surface and 9 spiniform

setae on inner surface.

Pereopod 6 (Fig. 3D). Basis with 3 long

plumose setae on dorsal margin, 4 long

plumose setae on ventral margin and 3

simple ventrodistal setae. Ischium with 2

long and 2 short ventrodistal simple setae.

Merus with several simple setae on

ventral margin, 1 ventrodistal and 1

dorsodistal plumose setae. Carpus ap-

proximately 2 times as long as merus with

long simple setae distally and on ventral

margin. Propodus with 12 outer spini-

form setae and approximately 33 inner

bipectinate setae. Dactylus longer than

propodus with 1 subterminal seta.

Pleopods (Fig. 3E). 5 biramous pairs.

Basal article with 5 long plumose setae.

Exopodite biarticulate with 1 plumose

seta on first article and 22 plumose setae

on distal article. Endopodite uniarticulate

with 30 plumose setae.

Uropod (Fig. 3F). Exopodite with

a short basal ‘‘pseudoarticle’’ (see ‘‘Re-

marks’’ section) followed by two longer

articles, the last being the longest; second

article with 1 long simple setae; last article

with 4 long simple setae. Endopodite with

approximately 18 articles.

Distribution.—Known only from the

type locality, San Quintin Bay, Baja

California, Mexico, 0–13 m.

Remarks.—Mesokalliapseudes crassus

can easily be distinguished from its

congeners by its acute rostrum, distinct

spinulate process on the antenna and

fixed finger of the cheliped, which is

approximately 0.67 the length of the

dactylus. All other congeners have

a rounded rostrum and the fixed finger

of the cheliped 0.5 or less the length of the

dactylus. It also appears that M. crassus is

more setose and spinose than other

member of the genus, especially the

antenna, antennule, chela and pereopod

6. It must be noted that in many

kalliapseudids (and perhaps apseudo-

morphs in general) the exopodite of the

uropod has a small round basal article. It

is unclear if this is a true article or if the

presumed articulation is an artifact,

representing a fracture line due to bend-

ing of the uropod. We believe there is

much confusion with this character; for

example, Sieg (1982) figures the exopod

of M. bahamensis (Sieg, 1982) as biarti-

culate, but examination of type (para-

types; USNM 181901) and non-type

specimens of this species reveals the

presence of a small round basal article.

Sieg (1982) also figures the exopod of M.

viridis (Menzies, 1953) as biarticulate, but

an indentation is shown instead of a line

between the round basal article and the

‘‘second’’ article. Most researchers have

treated this character as a true article, but

it can easily be missed if the uropod is

viewed dorsally. Thus, this character has

been treated differently by different re-

searchers and it is therefore recommended

that it should not be used to characterize

a species or in diagnostic keys until

a consensus is reached on how to properly

designate this ‘‘article.’’

Hermaphroditism has been recorded in

several tanaidacean groups, including all

three suborders. Lang (1953), in a study

of Apseudes hermaphroditicus Lang, 1953,


recorded specimens which possessed both

mature ova and a male genital cone.

Wolff (1956) indicated that Apseudes

gracillimus Hansen, 1913 is a protandric

hermaphrodite and suggested that this

condition may be more widespread. Fors-

man (1956) described protogynous her-

maphroditism in Heterotanais oerstedi

(Krøyer, 1842), the first record of proto-

gyny among Crustacea. Lang (1958)

found the same sexual development in

Tanais stanfordi Richardson, 1901 and

described two male forms, one of which

develops from a juvenile and one which

develops from a mature female. Apseudes

bermudeus Bacescu, 1980 was thought to

be a hermaphroditic species by Bacescu

(1980), and the observations of Heard et

al. (2004) indicate this species is a pro-

tandric hermaphrodite. Knight & Heard

(2006) also suggest the possibility of

hermaphroditism in A. larseni Knight &

Heard, 2006. Simultaneous hermaphro-

ditism has been suggested for the apseu-

did genera Androgynella Gutu, 2006 and

Muramurina Gutu, 2006 (Gutu 2006,

2007). Leite & Leite (1997) suggest the

possibility of protogynous hermaphrodit-

ism in the kalliapseudid Monokalliap-

seudes schubarti (Mane-Garzon, 1949)

based on the presence of two distinct

male forms, but additional material is

needed to determine if one or both of

these ‘‘forms’’ represent non-hermaphro-

ditic primary or secondary (protogynous)

males, or simply subterminal and termi-

nal male stages. Large secondary copula-

tory males can be derived from large

primary copulatory males so the presence

of two forms does not necessarily prove

hermaphroditism. The presence of proto-

gynous hermaphroditism has also been

suggested for some neotanaidomorphan

species (Kudinova-Pasternak 1965a, b;

Gardiner 1975). These life history recon-

structions have all been based upon

preserved material. Buckle-Ramirez’s

(1965) detailed developmental study on

live specimens of H. oerstedi revealed that

this species is a potential hermaphrodite

because the presence/absence of male

pheromones determines the sex of an

individual. Females of this tanaidomor-

phan species can either be truly gono-

choristic or they can be protogynous. This

type of development has also been shown

experimentally for the tanaids Leptochelia

dubia (Krøyer, 1842) (Highsmith, 1983),

Hargeria rapax (Harger, 1879) (Modlin &

Harris, 1989) and Leptochelia forresti

(Stebbing, 1896) (Stoner 1986) and is

presumably a reproductive strategy when

males are either absent or in low abun-

dance in populations.

Examination of type and non-type

material of M. crassus revealed that every

specimen with oostegites or a fully de-

veloped brood pouch also possessed

a genital cone. Unfortunately, the type

material from the LACM is in poor

condition; it appears that the specimens

had been dried and subsequently returned

to alcohol because every specimen was

shriveled to some degree. Some recovery

was made possible by placing them in

glycerol to soften. After this, the major

characters, including the genital cone,

could be studied. Because specimens

which did not have fully developed

oostegites had a genital cone and exam-

ination of a juvenile revealed the presence

of a forming genital cone, it is believed

that this species is a protandric hermaph-

rodite; however, the study of more

material and histological studies to de-

termine the presence of functional testes

in the ‘‘subadults’’ lacking developing of

fully developed oostegites are needed to

confirm this. There also remain two less

likely possibilities, at least for what is

previously known for the Tanaidacea, or

simultaneous hermaphroditism or parthe-

nogenesis. Notwithstanding, any of these

scenarios could account for the lack of

distinctive males in M. crassus.

It appears that the other congeners are

gonochoristic. Examination of the type

material of M. viridis [holotype adult


female with emptied marsupium (LACM1951-020.2); allotype juvenile (LACM

1951-020.5)] revealed the absence of

a genital cone. Menzies (1953) stated that

the juvenile was an immature male, but

we did not see a genital cone on this

specimen and the antennule did not have

any more aesthetascs than the adult

female. Since no males are known forthis species, hermaphroditism cannot be

ruled out until additional material is

examined. It must be noted that the

juvenile possessed hyposphaenia on the

pereonites and pleonites while the adult

female did not (probably resorbed). Sieg

(1982) did not recognize any male M.

bahamensis specimens in his collection,but Bamber (1993) examined the para-

types and noticed that approximately half

of them were males; examination of this

material (USNM 181901) confirms Bam-

ber’s observation. Mesokalliapseudes so-

niadawnae (Bamber 1993) and M. thali-

sispeleus Gutu, 2006 have been shown to

be gonochoristic (Bamber 1993; Gutu2006, respectively).

Key to species of Mesokalliapseudes

1. Pleonites with lateral plumose setae . . 2

Pleonites without lateral plumose

setae . . . M. brasiliensis (Bacescu, 1986)

2. Rostrum acute; fixed finger of cheli-

ped 0.67 or more length of dacty-

lus . . . . . . . M. crassus (Menzies, 1953)

Rostrum rounded; fixed finger of

cheliped 1/2 or less length of dacty-

lus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Anterolateral corners of pereonites

2–4 with apophysis . . . . . . . . . . . . . . 4

Anterolateral corners of pereonites

2–4 without apophysis . . . . . . . . . . . 5

4. One long seta on ventrodistal corner

of pereopod 1 basis; spinulate pro-

cess on second antenna article . .

. . . . . . . . . . . M. viridis (Menzies, 1953)

One short pectinate spiniform seta on

ventrodistal corner of pereopod 1

basis; no spinulate process on second

antenna article . . . . . . . . . . . . . . .

. . . . . . M. soniadawnae (Bamber, 1993)

5. Dactylus cutting edge of adult female

chela with more than 10 long spini-

form setae increasing in length dis-

tally . . . . . M. bahamensis (Sieg, 1982)

Dactylus cutting edge of adult female

chela with less than 10 short spini-

form setae . . . . . . . . . . . . . . . . . . .

. . . . . . . . . M. thalasispeleus Gutu, 2006

Acknowledgements

We would like to thank George Davis

and Joel Martin (LACM) for the loan of

type material and Charles Coleman (Ber-

lin Museum) for lending additional ma-

terial, which was borrowed from USNM.

We also thank two anonymous reviewersfor providing useful suggestions and

directing us to some relevant literature.

This research was supported by NSF

grant DEB-0529749.

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Associate Editor: Christopher B. Boyko


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