Steinernema ceratophorum
Jian, Reid & Hunt, 1997
DESCRIPTION
Females:
Body spiral or C-shaped when heatkilled. Cuticle with fine transverse striae
under SEM. Lateral field and phasmids inconspicuous. Head truncated to
slightly round, continuous with body; mouth opening circular. Six labial
and 4 cephalic papillae arranged as in other species. Stoma partially collapsed:
pharyngeal collar absent; esophagus extending near to mouth opening. Cheilorhabdions
located beneath lips and composed of thick, cuticularized ring, appearing
as two dark dots under a compound microscope . Below this is another cuticularised
ring, presumably prorhabdions; meso-, meta- and telo-rhabidions are vestigial
and presumably occur in collapsed area of stoma. Esophagus typical of family;
procorpus cylindrical and muscular; metacorpus slightly swollen, non-valvate;
isthmus distinct; basal bulb muscular, with small, distinct valve. Nerve-ring
surrounding isthmus. Esophago-intestinal valve almost bilaterally symetrical
and projecting into intestine. Excretory pore opening usually anterior
to nerve-ring; somewhat variable in location. First generation females
larger (average length: 5.37 mm) than second generation females (average
length: 1.7 mm). Vulva a transverse slit; vulval lips usually protruding
slightly from body surface. Vagina short, leading into amphidelphic genital
tracts. Females lay eggs initially, but later eggs hatch inside female
body with juveniles boring their way out. First generation female tail
shorter than anal body width, with small mucronate projection on
tip. In contrast, posterior region of second generation female tapering
evenly posteriorly, and ending with a fine mucron about 5.5 micrometers
long.
Male:
Similar to females but smaller and with male reproductive system. Body
curved posteriorly, when heat-killed. First generation male ( 1.38
mm long) larger than second generation male ( 0.9 mm long). Gonad monorchic,
reflexed. Spicules paired, symmetrical, curved, with distinct dark yellow
coloration; spicule head elongate; head length about 1.5 times or more
than head width, somewhat angular in shape; small rostrum present;
velum extending nearly to tip of lamina. Each spicule has 2 internal ribs.
Gubernaculum boat-shaped in lateral view and distally forked in ventral
or dorsal view; anterior part thin with or without a proximal hook, ventrally
curved, cuneus simple in dorsal view. Bursa absent.There are 23 genital
papillae (11 pairs plus one single) arranged as: 6 precloacal. subventral
pairs, most posterior being near cloaca; one single ventral papilla located
just anterior to cloaca; one adcloacal subdorsal pair located near the
proximal section of gubernaculum; 2 postcloacal subventral pairs and 2
subdorsal pairs near tail tip. Tail short, rounded conoid and lacking mucron
in both first and second generations.
Measurements (Male, first generation,
n=35): L=1358 SE=134 (1136-1694) um, W=146 SE=21 (104=185)um, stoma length=12
SE=1 (9-15), stoma width=7 SE=1 (5-8) um, EP=85 SE=11 (50-104)um, width
at excretory pore=56 SE=6 (45-72)um, NR=123 SE=14 (90-147) um, ES=165 SE=10
(149-190) um, testis reflexion=393 SE=94 (163-574) um, anal body width=52
SE=5 (45-70) um, tail length=30 SE=4 (23-38) um, spicule length=71 SE-7
(54-90) um, spicule width at head=11 (9-16) um, at neck=9 (7-11) um, at
lamina=11 (7-14) um, gubernaculum length=40 SE=4 (25-45) um, gubernaculum
width=7 SE=1 (5-9) um, D%=51.4 SE=7.2 (32.8-64.8) EW=1.5 SE=0.3 (07-2.0),
SW=1.4 SE=0.2 (1.0-2.0), GS=0.6 SE=0.1 (0.4-0.8), c'=0.6 SE=0.1 (0.4-0.8).
Infective stage juveniles:
Infective juveniles often enclosed in the 2nd stage cuticle. Body slender,
gradually tapering from base of espophagus to anterior end and from anus
to terminus. Cuticle with fine transverse striae; lateral field with
6 or 8 longitudinal ridges in mid-body region as seen under light microscope
and with SEM. Marginal and submarginal ridges are less distinct than other
ridges, and space (groove) between ridges are not even. Lip region
continuous, rounded and hemispherical with pair of lateral horn-like structures
near the oral opening ; mouth and anus closed; esophagus and intestine
collapsed. Excretory pore located in anterior region of esophagus (D% =
45.6). Anterior portion of intestine with pouch containing cells of symbiotic
bacteria. Tail straight, conoid, tapering evenly to a pointed terminus;
terminal spine absent. Hyaline region about half tail length.
Measurements(n=45): L=706 SE=62 (591-800)
micrometers (um), W=27 SE=3 (23-34) um, head to excretory pore=55 SE=5
(47-70) um, width at excretory pore=19 SE-1 (16-23), head to nerve ring=92
SE=6 (79-103) um, esophagus length=123 SE=7 (108-144), anal body width=15
SE=2 (9-18), tail length=66 SE=5 (56-74), a=25.9 SE=1.1 (23.7-27.9), c=10.6
SE=0.9 (8.8-12.9), D%= 45 SE=3.1 (40.0=55.8), E%=84.2 SE=6 (73.8-96.4),
c'=4.2 SE=0.4 (3.3-5.1).
DIAGNOSIS
AND RELATIONSHIPS
Steinernema ceratophorum is characterised by the following
combination of characters: infective stage juvenile with a body length
of about 706 (591-800) um; juvenile lip region with a double horn-like
structure and lateral field with six or eight longitudinal ridges; tail
tip of both first and second generation males lacking a cuticular mucron;
spicules curved with a distinct dark yellow coloration, about 71 and 54
um long in first and second generation respectively; first generation female
tail conoid with a small mucronate projection; and second generation female
tail tapering evenly posterior to salient posterior anal lip and ending
in a fine mucronate process about 5.5 um long. The RFLP banding pattern
is also unique when compared to a library of 33 other Steinernema
spp. The new species is morphologically similar to S. affine, S. intermedium,
S. riobrave and S. bicornutum, but can be separated from these four
species by a combination of morphological, DNA and cross-breeding techniques.
Only one other species is recorded as having two horn-like structures on
the infective juvenile head, viz. S. bicornutum. In addition,
S. ceratophorum can be further distinguished by the following characters:
the spicule length of both first and second generation males is longer
than S. bicornutum (71 vs 65 um; 54 vs 51 um) whereas the gubernaculum
is shorter (40 vs 48 um; 31 vs 33 um) and second generation male tail lacks
a small mucron, recorded as present in S. bicornutum; from S.
affine in lacking a mucron on the male tail tip; from S. intermedium
by the first generation male having a shorter spicule (71 vs 91 um) which
is dark-yellow rather than grey or colourless, a smaller average length
of gubernaculum. (40 vs 62 um), and in lacking a ventral concavity in the
tail (Poinar, 1988); from S. riobrave by the first generation male
spicules being longer (71 vs 67 um), the shorter gubernaculum (40 vs 51
um), the GS ratio (length of gubernaculum divided by spicule length) being
lower (0.6 vs 0.7) (Cabanillas et al., 1994) and the negative cross-breeding
test.
Etymology: Derived from the Greek keratos = a horn
and phoreus = a bearer (horn-bearer).
REFERENCE
JIAN, H., A. P. REID & D. J. HUNT, 1997.
Steinernema ceratophorum n. sp. (Nematoda: Steinernematidae) a new
entomopathogenic from north east China. Systematic Parasitology 37:115-125.
This document was constructed and is maintained by KHUONG
B. NGUYEN
Entomology & Nematology Department
University of Florida