Crustorhabdsaditis Chitwoodi Sp. Nov. (Nematoda Rhabditidae)

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Crustorhabditis chitwoodi sp. nov.(Nematoda: Rhabditidae): An intertidalspecies from the coast of New South

Wales, Australia, with observations onits ecology and life historyWarwick L. Nicholas

a

a Division of Botany and Zoology , Australian National University ,

Canberra, ACT, 0200, Australia E-mail:

Published online: 30 Mar 2010.

To cite this article: Warwick L. Nicholas (2004) Crustorhabditis chitwoodi sp. nov. (Nematoda:Rhabditidae): An intertidal species from the coast of New South Wales, Australia, with observations

on its ecology and life history, New Zealand Journal of Marine and Freshwater Research, 38:5,

803-808, DOI: 10.1080/00288330.2004.9517279

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New Zealand Journal

of

Marine

and

Freshwater Research, 2004,

Vol

38Nicholas—Crustorhabditischitwoodisp.n

0 0 2 8 - 8 3 3 0 / 0 4 / 3 8 0 5 - 0 8 0 3

©

T h e

R o y a l S o c i e t y

of

N e w

Z e a l a n d 2 0 0 4

803

rustorhabditis

chitwoodi sp nov.

Nematoda:

Rhabditidae):

an

intertidal

spec ies from the coast of New

South

Wales,

Australia, with observations on its ecology and life history

WAR WICK L. NICHOLAS

Division of Botany and Z oology

Australian National University

Canberra

ACT 0200, Austral ia

email: [email protected]

Abstract A new species, Crustorhabditis chit-

woodi ,

w as discovered in nutrient agar plates inocu-

lated with beach sand co llected from an o cean beach

in New South Wales, Austral ia.

The

three other

known species

of

the genus have

all

been found

as

commensal inhabitants

of

ocypodid shore crabs.

It

is suggested that the sand inoculum was seeded w ith

dauer larvae of the nematod e that had been dispersed

by scavenging crabs. The new species is distin-

guished from the three previously described species

by possessing a smaller number of differently ar-

ranged bursal papillae, eight rather than

10. The

species

is

amphim ictic, oviparous,

and

produces

abundant dauer larvae

in

culture, which can be sus-

tained indefinitely

on sea

water nutrient agar

in

which

it

feeds

on

associated bacteria.

K e y w o r d s

Crustorhabditis;

Brach yura; in tertidal

beach; marine; nematode

M0 4115; Online publication date

24

November 2004

Received 1 June 2004; accepted

9

September 2004

INTRODUCTION

The name Crustorhabditis was given by Sudhaus

(1974) to an ew subgenus of Rhabditidae to accom-

modate three species that are intimately associated

with beach-inhabi t ing ocypodid crabs . The new

species, described in this paper, was c ollected from

a sandy beach

in

south-eastern A ustralia awa y from

crabs

and can

reproduce

in the

laboratory

in

their

absence.

The

ghost crab,

Ocypode cardimana is

found o n the same beach, but with burrows near the

high t ide mark. Sudhaus (1974) described

C

riemanni, the type species, from nem atodes cultured

in the laboratory from specimens isolated from the

mouthparts and branchia l chamber of Ocypode

kuhlii, where it apparently lives as a saprobiont. T he

crab cam e from a beach at Kikambala, K enya, E ast

Afr ica . Most Rhabdi t idae

are

bacteria-feeding

terrestrial nematodes and, al though

a

number

of

bacteria-feeding Rhad itidae are comm only found in

the intert idal zone, only

the

three previously

described species

of

Crustorhabd itis

are

known

to

be closely associated with crabs.

Chitwood (1935) originally described

C. ocypo-

dis, which

he

found

on the

eggs

of O albicans

inhabit ing

a

sandy beach

in

North Carol ina ,

as

Parasitorhabditis ocypodis, but

later changed

the

n a me to Rhab ditis ocypodis (Chitwood, 1951).

Sudhaus (1974) redescribed the species from fixed

material obtained by R iemann from O. albicans from

a sandy Caribbean beach in Colum bia, South

Am erica. Sudhaus also redescribed

C. scanica,

origi-

nally named

Rhabdit is scanica by

Allgén (1949),

from specimens found between the mo uthparts of the

same beach crab that hosted

C. riemanni

when

the

crab was collected

at

Kikambala

in

Kenya.

Crustorhabditis is morphologically very close to

the subgenus M esorhabditis Osche, 1952. Osche's

mon ograph sets out to review the morphology of all

the then described species of Rhabd itis, but over-

looked Allgén's description of R. scanica. Osche

assigned

R. ocypodis

Chitwood, 1935,

to R habdit is

(Mesorhabditis) ocypodis.

Dougherty (1955),

in a

nomenclatorial review

of

the Rhabd it idae Örley,


4/8

804

New Zealand Journal of Marine and Freshwater Research, 2004, Vol. 38

1880, assigned both R. scanica and R. ocypodis to

the subgenus M esorhabditis . In a mono graph on the

taxonomy of the Rhabdi t idae, Andràssy (1983)

raised O sche's (1952) subgenera to generic rank.

METHODS

A culture of C. chitwoodi sp. nov. was established

fortuitously when a small amo unt of beach sand was

scattered on a 5 0 cm agar plate. The medium was 2

agar in sea water enriched w ith 5 m g Na

2

S iO

3

,5 mg

K

2

HPO

4

and 5 mg Fe(NH

4

)

2

(S O

4

)

2

per litre.

The intention had been to culture algae and

diatoms as part of ongoing studies of the dietary

habits of nematodes, but instead brown clumps of

glutinous bacteria-forming rosettes of cocci formed,

later to be dispersed and fed upon by the nematodes.

Some nematod es were transferred to 0.5 bacterio-

logical nutrient agar in sea water. Dense cultures and

many gen erations of the nem atodes were ma intained

by serial subculture on the same nutrient agar

medium. At this agar concentration the nematodes

move freely through the medium and are not

restricted to the surface.

Individual larval nematodes, or small groups of

eggs,

or newly hatched larvae were transferred to

small 5 cm closed agar plates and incubated at 20°C

to study the life cycle. All the nematodes fed on

bacteria transferred with the nema tode inoculum.

Nematode type specimens were fixed in 5

formalin plus 10 acetic acid for 24 h then washed

in distilled water and transferred to 5 aqueo us

glycerol. Water was evaporated in an oven at 40°C

and the nem atod es were then transferred with a fine

needle to anhydrous glycerol on glass slides and a

cover sl ip was supported by fine glass beads

(ballatini) and ringed with Glyceel (G urr). M easure-

ments were made of nematodes after fixation and

mounting in this way.

SYSTEMATICS

A diagnosis of genus Crustorhabditis is taken from

Andràssy's (1983) taxonomic monograph on the

Rha bditoidea, w hich places the genus hierarchically

in the Rhabd itidae, and M esorhabditinae. The body

is relatively large, 0.8 to 4 mm long, cuticle with

transverse annules and fine longitudinal striae. Head

slightly offset, lips well separated with small rod -like

papillae, amphids very small, on lateral lips. Stoma

(buccal cavity) 1.5 to 2 times as long as head

diameter. Cheilostom not cuticularised, promesos-

tom tubular with parallel walls; metastom consisting

of three hemispherical swellings, each bearing two

minute den ticles; pharyngeal sleave do es not invest

promesostom . Pharygeal corpus swollen, muscular,

isthmus long and non -muscular, nerve ring encloses

isthmus posteriorly, metacorpus with cuticular valve,

muscular, cardia small tripartite enclosed by anterior

intestine. Female gonad unpaired prodelphic, vulva

far posterior, close to anu s, tail conical. Ma le gona d

unpaired, spicules strong, the posterior two thirds

fused, bursa pelloderean, open anteriorly, anterior

half with crenulated margin, 10 pairs of genital

papillae present, two of them lying prean al, tail very

short, phasmids open close to anus.

Sudhaus & Fitch (2001) give a very detai led

review of the genus

Crustorhabditis

based on

C .

ocypodis

and

C. r iemanni,

but ignore

C. scanica.

Crustorhabditis

is morphologically very close to

Mesorhabdit is

and two of the three previously

described species were transferred by Sudhaus

(1974) to the newly created genus Crustorhabditis.

In distinguishing the two genera, Sudhaus draws

attention to their association with beach-inhabiting

ocypodid crabs, their large size, the crenulated

anterior bursal velum, the strong spicules fused for

the posterior two thirds of their length (in

Mesor-

habditis

usually fused for less than two thirds of their

length).

Crustorhabditis scanica

is smaller than the

other two species and the dependen ce of this species

on ocypod id crabs needs qualification.

Type species

Crustorhabditis chitwoodi sp. nov.

Material examined

Holotype male, AN IC slide 32 specimen

4 1:

Allo-

type female, slide 32 specimen 4 2; 9 paratype m ales:

slide 33 specimen 45, slide 35 specimens

5 0 - 5 3 ,

slide 36 specimens 5 7-5 9; slide 37 specimen 6 1; 9

paratype females: slide 32 specimens 43–44, slide

33 specimen 4 6, slide 34 specimens 4 7–49, slide 36

specimen 60, slide 37 specimens

6 2 - 6 3 .

All type

specimens taken from same agar plate culture

inocula ted with beach sand from the Brou lee beach,

New South Wales on 4 April 2004. Type specimens

are deposited in the Austral ian National Insect

Collection, Nematode Collection (ANIC), CSIRO

Entomology, G.P.O Box 1700, Canberra , ACT,

Australia.


5/8

Nicholas— Crustorhabdit is chitwoodi sp. nov

805

Description

Measurements in µm are given in Table 1, see also

Fig. 1.

Holotype male Body predominantely cylindrical,

tapered towards the head and truncated abruptly at

the posterior end; cuticle annulated; six slightly

offset lips bearing papilliform outer labial and

cephalic papillae, inner labial papillae not visible.

Cheilostom not cuticularised, Promesostom cylin-

drical, parallel sided, strongly cuticularised; a nterior

pharynx muscular, expanded to form muscular

corpus, followed by non-m uscular narrower isthmus,

then expanded to form muscular metacorpus;

metacorpus with semilunar cuticular valves,

terminating in tripart i te short muscular cardia.

Straight intest ine, anteriorly encloses cardia,

terminates in short narrower rectum , about level with

spicule capitulum, that terminates in cloaca. Single

testis, vas deferens ventral to gut, opens ven trally to

cloaca. Paired, long strongly built spicules, with

distinct capitulum, grooved shaft, fused for posterior

two thirds of their length, with sharp straight tip.

Bursa open a nteriorly, velum crenulated for anterior

half eight paired bursal rays, two short pair

pre-cloacal, one pair at level of cloacal opening,

paired group of three post-cloacal then two more

pairs slightly separated from their group of three, at

extreme posterior of bursal velum, beside very short

tail, triangular from dorsal view, no terminal notch

in velum (Fig 1).

Five paratype m ales do not differ significantly

from holotype.

Allotype female Essentially similar to male apart

from sexual organs and po sterior end of body. Single

ovary, uterus and oviduct ventral to gut, oviduct

terminating in a posterior vulva, close to but anterior

to an us. Oviduct with two uncleaved eggs. Tail initially

conical but drawn out into a terminal filament.

The excretory system described by Sudhaus &

Fitch (2001) was not identified in any of the types

or paratypes. A small perturbation of the cuticle

posterior to the nerve ring observed in several

specimens may be the excretory pore, but no

terminal cuticular duct was evident so the identity

of the cuticular structure is doubtful. Perhaps the

excretory system is less prominent in specimens

cultured in a saline medium than in typical terrestrial

rhabditids.

Table

Measurements of type specimens of Crustorhabdit is chitwoodi sp. nov. All measurements in µm.

Type

Length

Buccal cavity

Prostom

Pharynx

Head to corpus

Corpus length

Head to nerve ring

Head to bulb

Bulb length

Head to intestine

Head to gonad

Spicule length

Gubernaculum

Bursa length

Head to vulva

Vulva to anus

Head to anus

Tail length

Max. width

Width at vulva

Width at anus

de M an's a

de M an's b

de M an's c

de M an's V

Holo.

Male

970

22

17

194

98

41

149

182

34

223

317

67

48

53

95 3

17

43

23

23

5.0

40

Males

Mean

1020

23

17

194

93

35

144

174

35

21 8

32 8

62

37

58

989

24

43

23

24

5.3

4 2

n = 10

SD

57

1.66

1.48

9.6

8.61

3.6

10.4

10

4.0

10.4

31

4.6

6.4

5.6

56

2.3

4.9

2.07

2.91

0.23

2.1

Range

970-1084

19-24

15-19

196-202

77-101

31–41

134-158

161-188

30–43

199-226

283-379

53-67

26–48

53-70

944-1065

2 4 -3 6

36–48

2 0 -2 6

2 1 -3 0

5.0-5.6

37-57

Allo.

Female

1090

22

17

196

91

41

1581

185

41

22 6

468

960

41

1001

89

43

43

26

25

5.6

12.3

88

Females

Mean

1086

22 2

18

193

93

3.5

158

180

38

218

34 9

961

49

946

89

40

36

27

28

5.6

12.3

89

n= 10

SD

64

3.0

2.4

10.9

4.0

8.84

10.4

2.9

12.4

41

58

4.5

8.62

67

3.1

6.2

3.1

5.63

4.61

0.24

1.15

1.8

Range

–10841090

17-24

17-22

178-213

86-96

29–41

132-136

161-195

35–41

197-228

332-467

883-990

4 3 - 5 3

869-1039

82-101

36–48

30-46

19-41

2 2 -3 4

5.2-5.9

9.7-13.1

86-89


6/8

80 6


50ftm

X

Fig. 1 Crustorhabdit is sp. nov.

50 nm A, head and pharyngeal region; B,

head and buccal cavity; C, hind

end of female in lateral view; D,

hind end of male in lateral view;

E, hind end of male in dorsal view.

chitwoodi

riemanni

scanica

ocypodis

Fig . 2 Com parison of the bursal

region and bursal papillae of all

four known species of the genus

Crustorhabdit is. C. riemanni, C.

ocypodis,

and

C. scanica

from

Sudhaus (1974); C. chitwoodi, n.

sp .

original. Drawin gs not to same

scale.


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Nicholas— Crustorhabdit is chitwoodi sp. nov

807

Dauer larvae, 536-78 7 µm long 21- 24 µm wide,

accumulate in cultures. These are a non-growing

dispersive stage in the life cycle. In common with

other Rhabdit idae, these dauer larvae retain the

cuticle of the second instar, the lips and lumen of the

alimentary canal are closed, but they retain m otility.

Differential diagnosis

Significant differences between C. chitwoodi sp .

nov.,

C. r iemanni, C . ocypodis ,

an d

C. scanica

are

listed in Table 2.

C . r iemanni

an d

C. ocypodis

are

much larger than the other two species, which are

of comparable size. De M an 's ratios of pharyngeal

length to body length,

b ,

and tail length to body

length, c, as well as the ratio of the distance b etween

the vulva and an us to tail length and the absolute tail

length are greater in the former two, probably related

to their greater size. De Man's rat ios are not

independent of body length. C. scanica has shorter

spicules than any of the other three species.

C .

scanica

an d

C. chitwoodi,

sp. nov. are more alike

than the other two. However, the most significant

difference between the four species is the difference

in the number and disposition of the bursal papillae

(Fig. 2). Unlike the other species with 10 pairs, C .

chitwoodi sp. nov. has only 8 pairs. Baird (2001)

shows that in the rather closely related Caeno-

rhabditis briggsae the bursal papillae are a good

indicator of species, although some variation can

occur in strains.

Type locality

Mid-tide level on a sandy beach at Broulee on the

South East Coast of New South Wales, Australia,

La titu de 35º55¢ , Lon gitu de 150º9¢ .

Etymology

The specific epithet nam ed in hono ur of the greatest

nematologist and the describer of

Crustorhabditis

ocypodis .

Reproductive biology

Crustorhabditis chitwoodi sp. nov. is amphimictic,

with approximately equal numbers of male and

female progeny. Females are normally oviparous,

but sometimes eggs hatch in utero and the larvae

devour the mother, i.e., endotokia matricida. Chen

& Caswell-Chen (2003) show that in the related

rhabditid

Caenorhab ditis elegans

this behaviour is

an adaptive response to failing food resources. In the

absence of ma les, females die with several unhatched

eggs

in utero.

Protandrous hermaphrodi t i sm,

comm on in some rhabditids, has not been observed.

The generation time at 20°C is c. 7 days.

Ecology

Crustorhabditis ocypodis was recovered from the

eggs of the shore crab

Ocypode albicans

in North

Carolina in North Am erica by Chitwood (1935) and

again from the branchial chamber of the same

species from Columbia in South America

(Rieman n, 1970).

C. riemanni

has been found in the

branchial chamber and between the mouth parts of

the shore crab Ocypode kuhlii in Ken ya, East Africa,

together with C. scanica. How ever, C. chitwoodi sp.

nov. was not found in direct association with crabs

when it was recovered from sand at mid-tide level,

although

Ocypode cardimana

occurs on the same

beach clo se to the high tide mark. Con sequently this

species is not totally restricted in its distribution to

the immed iate presence of crab s. It can multiply for

Table 2

Com parison of significant taxono mic measurements of three previously described species of

Crustorhah bdit is

an d

C . chitwoodi

sp. nov. All measurements in µm.

Species:

Sex:

Body length

Vulva-anus/tail

Tail length

Spicules length

spicule fused

Bursal formula

d eM a n s ' a

de Ma n's b

de Ma n's c

C. riemanii

Male

1895-2820

0.96

27-54

53-62

66

2 .2 .6= 10

16.5-21.8

6.5-9.1

39.2-64.7

Female

2422-3890

91-158

14.5-23

7.9-11.8

19.6-35.8

C. ocypodis

Male

2015-2380

0.68

24-30

65-71

69

2.1.7 = 10

26.7–40.4

6.1-7.4

71.1-98.5

Female

2 4 2 2 3 8 9 2 1 5 – 2 3 8

101-162

25.4-38.9

6.8-8.7

14.2-17.8

C. scanica

Male

7 8 5 - 1 1 7 0

17-26

38–48

54

2.8 = 10

15.0-22.8

4 . 0 - 6 . 5

31 .4-49 .3

Female

857-1516

0.56

8 3 - 1 2 5

15.0-22.816

6.1-7.46.8

8.7-16.4

C .

chit

Male

970-1084

24-36

53-67

57-60

2.1.3.2 = 8

21-30

5.0-5.6

37.3-57

woodi

Female

950-1090

0.40-0.63

82-101

22-34

5.2-5.9

8.7-1


8/8

80 8


many generations on nutrient agar, feeding on

associated bacteria.

The dependence of C. scanica for reproduction

on the presence of crabs is uncertain. Schuurmans

Stekhoven (1943) reported Rhabdit is monhystera,

which seems from its description to be a synonym

for

C. scanica,

from sand together with the macro-

algae

Codium

and

Caulerpa

in the Mediterranean.

According to Sudhaus (1974), Allgén (1934)

reported a single male as

Rhab ditis marina

with the

bursal papilla formula of 2/8 so that this specimen

was in all probability C. scanica. Allgén (1949),

again according to Sudhaus (1974), described C .

scanica as a new species from a single male derived

from marine algae. Sudhaus (1974) equates the

single female figured by Gerlach (1953) from c oastal

ground water in Madagascar as

C scanica,

but its

identi ty is by no means certain. In summary,

although

C. scanica

has been found in the mouth

parts of ocypode crabs it has also been found in the

absence of crabs. Sudhaus suggests its dauer larvae

may have been dispersed by c rabs.

Although

C. chitwoodi

sp. nov. was not found in

the proximity of crabs, ocypode crabs do inhabit the

beach from which it was found. It is not a comm on

inhabitant of the beach m eiofauna bec ause it has not

previously been found in studies of this and neigh-

bouring beaches during several years of sampling

(Nicholas 200 1). The m ost likely explanation is that

Ocypode cardimana

crabs scavenge at night at low

tide on the beach and disperse dauer larvae, as

Sudhaus (1974) suggests, and though these are

capable of feeding and reproducing on bacteria, the

habitat does not facilitate sustained reproduction

without direct association with ocypode crabs. It

would be interesting to examine a number of the

local ocypodid crabs for nem atodes, but the author

sees no prospect of doing so in the near future.

REFERENCES

Allgén, C. 1949: Uber einiger sudschwedische Brack-

wasser und Eardnematoden.

Kunglia Fysiogra-

fiska Säliskkapets i Lund Förhandlingar19: 1—19.

Andràssy, I. 1983: A taxonomic review of the suborder

Rhabditina (Nematoda: Secernentia). Edit ions de

l Office de la Re sea rch e Scientifiqu e et Tec h-

nique outra-mer. Paris. Pp 1-240.

Baird, S. E. 2001: Strain specific variation in the pattern

of caudal papillae in

Caenorhabdit is briggsae

(Nematoda: Rhabditidae); implications for spe-

cies identification. Nematology 3: 373-376 .

Chen, J.; Caswell-Chen, E.

2003:

W hy

Caenorhabdit is

elegans adults sacrifice their bodies to progeny.

Nematology 5: 641-645.

Chitwood, B . G. 1935: Nema todes parasitic in, and asso-

ciated with, Crustacea, and description of some

new species and a new variety.

Proceedings of

the Helminthological Society of Washington 2:

93-96 .

Chitwood, B. G. 1961: North American marine nema-

todes.

Texas Journal of Science 3:

617-672.

Dougherty, E. C. 1955: The genera and species of the

subfamily Rhabditinae Micoletzky, 1922 (Nema-

toda):

a nom enclatorial analysis-including an ad-

dendum on the composit ion of the family

Rhabditidae Örley, 1880. Journal o f Helminthol-

ogy 29:

105-152.

Gerlach, S. A. 1953: Recherches sur la faune des eaux

interstitielles de Mad agascar.

M émoires de l Institut

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Crustorhabdsaditis Chitwoodi Sp. Nov. (Nematoda Rhabditidae)

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