Corresponding author: Jans Morffe (
Academic editor: H.-P. Fagerholm
The genus
At present, the male is known only in two species:
In this work, we discuss the taxonomical status of
All the passalids for this study were collected by hand from rotting logs in Cuba. Two specimens of
Two additional specimens of
Measurements of the nematodes were made with a calibrated eyepiece micrometer and are given in millimeters. De Man’s ratios a, b, c and V% were calculated. Micrographs were taken with an AxioCam digital camera attached to a Carl Zeiss AxioScop 2 Plus compound microscope. Line drawings were made on the basis of micrographs using the softwares CorelDRAW X3 and Adobe Photoshop CS2. Scale bars of all Figures are given in millimeters.
The material examined is deposited in the Colección Helmintológica de las Colecciones Zoológicas (CZACC), Instituto de Ecología y Sistemática, Havana, Cuba.
First cephalic annule notably long, surpassing the stoma length, inflated, forming a truncated cone-like structure. According with
Posterior end finished in a sharply pointed tail, instead of the ventral expansion of the tail cuticle in
These discrepancies support the possibility that the males attributed to
♀ holotype; Cuba, La Habana Province, San José de Las Lajas, La Jaula; in
4 ♂♂ vouchers; Cuba, Holguín Province, Nipe-Sagua-Baracoa, La Melba; in
3 ♀♀ paratypes of
Holotype (female) a = 23.78, b = 4.63, c = 7.04, V% = 51.70, total length = 3.520, maximum body width = 0.148, first cephalic annule (length×width) = 0.028×0.035, stoma length = 0.078, procorpus length = 0.640, isthmus length = 0.045, diameter of basal bulb = 0.070, total length of oesophagus = 0.760, nerve ring to anterior end = 0.233, excretory pore to anterior end = 0.980, vulva to posterior end = 1.700, anus to posterior end = 0.500, eggs = 0.123–0.128×0.045–0.053 (0.125 ± 0.003×0.049 ± 0.004 n = 3).
Female adult paratypes (n = 11) a = 23.61–28.28 (25.00 ± 1.57 n = 11), b = 4.49–5.45 (4.81 ± 0.29 n = 10), c = 6.24–8.54 (7.37 ± 0.66 n = 11), V% = 49.73–58.01 (52.98 ± 2.34 n = 11), total length = 3.500–4.525 (3.863 ± 0.277 n = 11), maximum body width = 0.138–0.173 (0.155 ± 0.010 n = 11), first cephalic annule (length×width) = 0.023–0.028×0.035–0.038 (0.024 ± 0.002×0.036 ± 0.001 n = 9), stoma length = 0.073–0.085 (0.079 ± 0.005 n = 11), procorpus length = 0.630–0.720 (0.678 ± 0.027 n = 10), isthmus length = 0.040–0.058 (0.053 ± 0.005 n = 10), diameter of basal bulb = 0.065–0.078 (0.071 ± 0.004 n = 10), total length of oesophagus = 0.760–0.860 (0.806 ± 0.030 n = 10), nerve ring to anterior end = 0.230–0.250 (0.242 ± 0.007 n = 9), excretory pore to anterior end = 0.970–1.125 (1.050 ± 0.047 n = 10), vulva to posterior end = 1.600–1.925 (1.813 ± 0.097 n = 11), anus to posterior end = 0.470–0.590 (0.526 ± 0.038 n = 11), eggs = 0.110–0.143×0.050–0.060 (0.133 ± 0.006×0.053 ± 0.003 n = 28).
Female immature paratypes (n = 5) a = 24.31–27.02 (24.96 ± 1.16 n = 5), b = 4.13–4.42 (4.28 ± 0.12 n = 5), c = 7.07–8.86 (7.74 ± 0.68 n = 5), V% = 53.54–56.56 (54.99 ± 1.07 n = 5), total length = 3.050–3.325 (3.165 ± 0.104 n = 5), maximum body width = 0.118–0.135 (0.127 ± 0.006 n = 5), first cephalic annule (length×width) = 0.023–0.025×0.035 (0.024 ± 0.001×0.035 ± 0.000 n = 5), stoma length = 0.070–0.073 (0.072 ± 0.001 n = 5), procorpus length = 0.610–0.680 (0.638 ± 0.028 n = 5), isthmus length = 0.055–0.058 (0.056 ± 0.001 n = 5), diameter of basal bulb = 0.063–0.068 (0.065 ± 0.002 n = 5), total length of oesophagus = 0.690–0.760 (0.740 ± 0.029 n = 5), nerve ring to anterior end = 0.223–0.233 (0.227 ± 0.004 n = 5), excretory pore to anterior end = 0.910–0.980 (0.942 ± 0.031 n = 5), vulva to posterior end = 1.325–1.500 (1.425 ± 0.068 n = 5), anus to posterior end = 0.350–0.470 (0.412 ± 0.045 n = 5).
Allotype (male) a = 15.68, b = 3.63, c = 49.67, total length = 1.490, maximum body width = 0.095, stoma length = 0.038, procorpus length = 0.330, isthmus length = 0.033, diameter of basal bulb = 0.045, total length of oesophagus = 0.410, nerve ring to anterior end = 0.160, excretory pore to anterior end = 0.560, cloaca to posterior end = 0.030, distance from the mammiform papillae to the posterior end = 0.130, distance from the dorso-lateral papillae to the posterior end = 0.065.
Male paratypes (n = 5) a = 11.56–14.10 (12.40 ± 1.04 n = 5), b = 3.13–3.60 (3.38 ± 0.17 n = 5), c = 43.33–51.27 (47.13 ± 3.54 n = 5), total length = 1.220–1.510 (1.366 ± 0.111 n = 5), maximum body width = 0.100–0.120 (0.111 ± 0.010 n = 5), stoma length = 0.040–0.048 (0.043 ± 0.003 n = 5), procorpus length = 0.300–0.340 (0.322 ± 0.018 n = 5), isthmus length = 0.030–0.033 (0.031 ± 0.001 n = 5), diameter of basal bulb = 0.045–0.048 (0.047 ± 0.001 n = 5), total length of oesophagus = 0.390–0.420 (0.404 ± 0.015 n = 5), nerve ring to anterior end = 0.150–0.168 (0.159 ± 0.007 n = 5), excretory pore to anterior end = 0.480–0.550 (0.512 ± 0.028 n = 5), cloaca to posterior end = 0.028–0.030 (0.029 ± 0.001 n = 5), distance from the mammiform papillae to the posterior end = 0.128–0.148 (0.137 ± 0.010 n = 3), distance from the dorso-lateral papillae to the posterior end = 0.058–0.065 (0.062 ± 0.003 n = 5).
Males (n = 4) a = 11.55–14.00 (13.00 ± 1.12 n = 4), b = 3.10–4.20 (3.50 ± 0.48 n = 4), c = 33.87–61.09 (44.96 ± 11.49 n = 4), total length = 1.170–1.680 (1.348 ± 0.227 n = 4), maximum body width = 0.085–0.120 (0.104 ± 0.015 n = 4), stoma length = 0.038–0.040 (0.039 ± 0.001 n = 4), procorpus length = 0.283–0.320 (0.303 ± 0.016 n = 4), isthmus length = 0.028–0.038 (0.032 ± 0.004 n = 4), diameter of basal bulb = 0.050–0.063 (0.054 ± 0.006 n = 4), total length of oesophagus = 0.378–0.400 (0.384 ± 0.010 n = 4), nerve ring to anterior end = 0.133–0.158 (0.145 ± 0.013 n = 3), excretory pore to anterior end = 0.430–0.490 (0.463 ± 0.025 n = 4), cloaca to posterior end = 0.028–0.038 (0.031 ± 0.005 n = 4), distance from the mammiform papillae to the posterior end = 0.115–0.133 (0.125 ± 0.009 n = 3), distance from the dorso-lateral papillae to the posterior end = 0.055–0.070 (0.063 ± 0.006 n = 4).
Females (n = 3) a = 17.20–23.94 (21.67 ± 3.87), b = 4.79–4.86 (4.83 ± 0.05), c = 6.68–7.63 (7.25 ± 0.51), V% = 45.99–52.19 (50.04 ± 3.51), total length = 3.580–3.890 (3.737 ± 0.155), maximum body width = 0.150–0.218 (0.177 ± 0.036), first cephalic annule = 0.018–0.025×0.035–0.038 (0.021 ± 0.005×0.036 ± 0.002), stoma length = 0.060–0.075 (0.066 ± 0.008), procorpus length = 0.650–0.660 (0.655 ± 0.007), isthmus length = 0.050–0.058 (0.053 ± 0.004), diameter of basal bulb = 0.070–0.073 (0.071 ± 0.002), total length of oesophagus = 0.780–0.800 (0.790 ± 0.014), nerve ring to anterior end = 0.225–0.235 (0.230 ± 0.005), excretory pore to anterior end = 0.990–1.05 (1.020 ± 0.042), vulva to posterior end = 1.720–2.020 (1.867 ± 0.150), anus to posterior end = 0.480–0.560 (0.517 ± 0.040), eggs = 0.128–0.140×0.045–0.060 (0.133 ± 0.005×0.053 ± 0.006 n = 5).
Female. Body long and relatively slender. Cervical cuticle without spines. Subcuticular longitudinal striae present. Lateral alae well developed, arising from about two body widths posterior to the basal bulb up to the level of the anus. Head rounded, set-off from body by a single, deep groove, bearing eight prominent, paired papillae. First cephalic annule slightly inflated, about 2 to 2.5 head-lengths long. Stoma long, slender, about 3.5 first cephalic annule lengths long, surrounded by an esophageal collar. Oesophagus consisting of a very elongated, subcylindrical, muscular procorpus less set-off from the isthmus. Basal bulb sub-spherical, valve plate well developed. Intestine simple, sub-rectilinear, its fore region slightly inflated. Rectum short, anus not prominent. Nerve ring encircling procorpus at about 20% of its length. Excretory pore located at about a body width posterior to basal bulb. Vulva a median transverse slit, near midbody, lips not prominent. Vagina muscular, strongly developed, forwardly directed. Genital tract monodelphic-prodelphic. Ovary reflected at about two body widths posterior to bulb, distal flexure about two body widths long. Eggs ovoid, with eight, rough, very prominent, longitudinal ridges not reaching the poles. Oocytes in a single row. Tail conical, subulate, ending in a fine tip.
Male. Body comparatively stout, shorter than female. Posterior region of body ventrally curved. Cervical cuticle unarmed, visibly annulated up to a little distance before the nerve ring. Sub-cuticular longitudinal striae present. Head set off from body by a single groove and bearing eight small, paired papillae. First cephalic annule not conspicuous. Stoma long, surrounded by an esophageal collar. Oesophagus slightly sub-fusiform, diminishing its diameter toward its second half until equaling the diameter of the isthmus. Basal bulb rounded, valve plate developed. Nerve ring encircling procorpus at about 35% of its length. Excretory pore situated at little more than a body-width posterior to basal bulb. Testis single, full of spermatids, commencing at about 1.5 body-widths posterior to bulb. Spicule absent. A large, median, mammiform precloacal papillae situated at a little less than a body width before the posterior end. A pair of dorso-lateral precloacal papillae located at about 0.5 body-widths before the tail tip. Dorsal cuticle of posterior end thickened from level of the latter pair of papillae to the tail tip. There is a cuticular bursa-like structure in the tail end, almost reaching the level of the tail tip.
Named after the Latin
Gut caeca
La Jaula, San José de Las Lajas, La Habana Province, Cuba.
All the records of females of
Comparative measurements of the females and males of
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La Jaula, La Habana Province (type locality) (n = 12) | El Salón, Pinar del Río Province (n = 3) | La Melba, Holguín Province (n = 6) | La Platica, Granma Province (n = 6) |
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Total length | 3.500–4.525 | 3.580–3.890 | 3,390–4,090 | 3,620–4,150 |
Body width | 0.138–0.173 | 0.150–0.218 | 0,150–0,170 | 0,100–0,140 |
Stoma length | 0.073–0.085 | 0.060–0.075 | 0,055–0,060 | 0,058–0,070 |
Procorpus length | 0.630–0.720 | 0.650–0.660 | 0,640–0,740 | 0,600–0,690 |
Isthmus length | 0.040–0.058 | 0.050–0.058 | 0,050–0,063 | 0,048–0,060 |
Diameter basal bulb | 0.065–0.078 | 0.070–0.073 | 0,073–0,080 | 0,060–0,073 |
Esophagus length | 0.760–0.860 | 0.780–0.800 | 0,770–0,950 | 0,730–0,820 |
Nerve ring-head end | 0.230–0.250 | 0.225–0.235 | 0,225–0,250 | 0,235–0,250 |
Excretory pore-head end | 0.970–1.125 | 0.990–1.05 | 0,950–1,040 | - |
Vulva-tail end | 1.600–1.925 | 1.720–2.020 | 1,720–2,040 | 2,000 |
Anus-tail end | 0.470–0.590 | 0.480–0.560 | 0,480–0,610 | 0,600–0,640 |
Eggs | 0.110–0.143×0.050–0.060 | 0.128–0.140×0.045–0.060 | 0,135–0,153×0,050–0,070 | 0,125–0,140×0,050–0,058 |
a | 23.61–28.28 | 17.20–23.94 | 21,19–27,27 | 29,36–36,20 |
b | 4.49–5.45 | 4.79–4.86 | 4,27–5,13 | 4,83–5,41 |
c | 6.24–8.54 | 6.68–7.63 | 6,70–7,90 | 6,03–6,92 |
V% | 49.73–58.01 | 45.99–52.19 | 45,65–55,67 | 49,24 |
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Total length | 1.220–1.510 | - | 1.170–1.680 | - |
Body width | 0.095–0.120 | - | 0.085–0.120 | - |
Stoma length | 0.038–0.048 | - | 0.038–0.040 | - |
Procorpus length | 0.300–0.340 | - | 0.283–0.320 | - |
Isthmus length | 0.030–0.033 | - | 0.028–0.038 | - |
Diameter basal bulb | 0.045–0.048 | - | 0.050–0.063 | - |
Esophagus length | 0.390–0.420 | - | 0.378–0.400 | - |
Nerve ring-head end | 0.150–0.168 | - | 0.133–0.158 | - |
Excretory pore-head end | 0.480–0.560 | 0.430–0.490 | ||
Cloaca-tail end | 0.028–0.030 | - | 0.028–0.038 | - |
Median papilla-tail end | 0.115–0.133 | - | 0.128–0.148 | - |
Dorso-lateral papilla-tail end | 0.055–0.070 | - | 0.058–0.065 | - |
a | 11.56–15.68 | - | 11.55–14.00 | - |
b | 3.13–3.63 | - | 3.10–4.20 | - |
c | 43.33–51.27 | - | 33.87–61.09 | - |
♂ voucher; Cuba, La Habana Province, Jaruco, Escaleras de Jaruco; in
Male (n = 1) a = 15.66, b = 4.15, c = 54.80, total length = 1.370, maximum body width = 0.088, stoma length = 0.045, procorpus length = 0.260, isthmus length = 0.028, diameter of basal bulb = 0.040, total length of oesophagus = 0.330, nerve ring to anterior end = 0.145, excretory pore to anterior end = 0.500, cloaca to posterior end = 0.025, distance from the mammiform papillae to the posterior end = 0.153, distance from the dorso-lateral papillae to the posterior end = 0.068.
Male body comparatively slender, shorter than female. Cuticle thin, finely annulated. Sub-cuticular longitudinal striae present. Head short, rounded, set-off from body by a single groove, papillae not observed. First cephalic annule not differentiated. Stoma as in the female. Oesophagus with its second half slender than first, hardly set-off from the isthmus. Basal bulb rounded, valve plate developed. Intestine simple, slender, its fore region dilated. Anus inconspicuous. Nerve ring encircling procorpus at about 40% of its length. Excretory pore located at about 1.5 body widths posterior to basal bulb. Testis single, full of spermatids, commencing just behind excretory pore, its second third slightly inflated. Spicule absent. A large, mid-ventral, precloacal, mammiform papilla situated at about 1.5 body widths before the tail tip. A pair of dorso-lateral, precloacal papillae at little less than a body width before the tail tip. Dorsal cuticle of the posterior end thickened from the level of the pair of papillae to almost the tail tip. Ventral cuticle expanded, forming a bursa-like structure with the distal end terminating before the tail tip. Tail short, conical, ending in a rounded tip. Posterior end slightly reflexed toward dorsal region.
The male of
From
Gut caeca.
Escaleras de Jaruco, Jaruco, La Habana Province, Cuba.
Note: males are described only in two of the five Cuban
1 | Isthmus long and bent | |
– | Ishmus short and not bent | 2 |
2 | First cephalic annule not inflated | |
– | First cephalic annule more or less inflated | 3 |
3 | Cervical cuticle without wide annule, ovary reflected at about two body widths beyond the excretory pore | |
– | Cervical cuticle with wide annule, ovary reflected at little less than a body width beyond the excretory pore |
Authors are indebted to Lic. René Barba (Instituto de Ecología y Sistemática) and Lic. David Ortiz for the collection of the hosts from La Melba. To Lic. Elier Fonseca and Lic. Gunnary León (Universidad de La Habana) and the undergraduate student Facundo Alvarez for their help during field work. To Dr. Luis F. de Armas (Instituto de Ecología y Sistemática) for the revision of the early version of the manuscript and Dr. Pedro Herrera (Instituto de Ecología y Sistemática) for the corrections of the English manuscript. We also thank MSc. Yamir Arias (Instituto de Ecología y Sistemática) for his technical assistance with the micrographs. This work was supported by IDEAWILD and the project DB-06 “Colecciones Zoológicas, su Conservación y Manejo” of the Ministerio de Ciencia, Tecnología y Medio Ambiente (CITMA), Cuba.