Lacandodrilus paludosus

Septa all thin and membranous. Paired septal glands in V, VI, VII, VIII and IX (in X incipient), attached to corresponding posterior septum, mixed with lateral vascular commissures. Gizzard, caeca and typhlosole absent. Oesophagus moniliform, with no distinct transition to intestine. Dorsal vessel single; sometimes increasing from XIV posteriad. Vascular commissures long and contorted in segments V?, VI – XI; embedded in septal glands. Ventral vessel present. Ventral holonephridia from XI backwards; covered by a granular peritoneal layer giving them an ovoid, flat appearance; this ovoid portion floating in the coelom, and not attached to septa. Nephridia gradually increasing in size and from XV attaining maximal size; exonephric (opening in ab) and probably open (stomate). Holandric; iridescent male funnels in X and XI; those of XI the larger; testes not seen. Both funnels enclosed in seminal vesicles. Male gonoduct coming from male funnels of X entering the body wall in XI; a second gonoduct coming from funnels of XI entering the body wall in XII. Both male ducts intraparietal over XI, XII and 3 / 4 of XIII (deduced from the externally observed, undulating cream band). Accordingly, male pore deduced to be in XIII, postsetal and in BC. One pair of sausage – shaped vesicles in XI; projecting in posterior direction from XII until segments XVIII or XX and running dorsolateral of the digestive tract, completely covering it. In X another pair of vesicles that also contain male funnels that run in anterior direction before turning over 180 ° and which, apparently, are limited to XII. Two pairs of lobulate, rosette – shaped glands internally on the ventral body wall XIII and XIV, the anterior the larger; composed of several short lobules, deduced to be prostatic glands. No tubular, atrium – like glands present. Large ovaries in XIII projecting from the ventral portion of 12 / 13 at AB and extending over the entire segment. Female funnels on the ventral wall of XIII very close to 13 / 14, in AB, and with a duct opening in XIV before setae ab. Septum 13 / 14 apparently modified as dorsolateral paired ovisacs that contain the corresponding seminal vesicles of XII and extending backwards until XVIII or XX. One pair of adiverticulate spermathecae in X opening ventrally in 9 / 10, at AB, the duct turned 180 ° to connect with a sausage – shaped ampulla. Total length of spermathecae 580 µm, 370 µm the ampulla, and 210 µm the duct.

Remarks. This species was considered as gen. nov. sp. nov. 4, but not named, by Fragoso & Lavelle (1987). Fragoso and Rojas (2033) suggest that future morphological and molecular evidence might render Lacandodrilus the type of a new family within the order Alluroidida. The absence of normal atria and presence of two pairs of prostates not associated with the male pores lend some support to this proposition.

Distribution. Mexico, Chiapas, Lacandon tropical rain forest in semi – inundated gley soils at 30 – 40 cm depth.

Ecology: This species, considered as gen. nov. sp. nov. 4 by Fragoso & Lavelle (1987), inhabited only the swampy gley soils of tropical rain forest at the Lacandon region. It was found at 40 cm depth in flooded, sandy soils, characterized by its acidity (pH = 4.5), and low values of organic matter (0.9 %), Ca (2.32 meq / 100 g) and Mg (0.72 meq / 100 g), and with an increase of clay in deep strata. The site was dominated by the palms Bactris balanoidea and Bactris trichofila. All the individuals were found in a single sample in the month of December. Abundance and biomass at the entire forest were very low, less than one individual by square meter (Fragoso & Lavelle 1987); however, if only samples taken in gley soils are considered, the average abundance is higher (1.6 ind / m 2).

Description. External. Dimensions: Length 50, 50, 58 mm (average = 53, n = 3), holotype 28 mm (fragmented, only anterior half; Fig 1 A). Width, middle body: 0.46, 0.50, 0.64, 0.64 mm (average = 0.56, n = 4); holotype = 0.48 mm. Number of segments: 208, 225, 248 (average = 227, n = 3). Furrows: One pre- and one post-setal along all the body; in some individuals up to two pre- and two post-setal in some segments. Pigment: Absent. Prostomium; Prolobic. Setae: Eight per segment, present from segment 2 (Fig. 1 A); closely paired throughout. Setae of preclitellar segments, conspicuous and with replacements. Setae a and b of preclitellar segments measuring 130 – 150 µm length, almost straight, slightly bent distally and with a slight nodulus close to its basal end; apex smooth and without ornamentations (Fig 2 A, 2 B). Setae a, b of segments 11 and 12 larger, but with the same shape of anterior ones; replacement setae measuring 64, 93 µm (Fig. 2 A, 2 B); seta c (112 µm) and d smaller than seta a and b. After clitellum all setae similar (slightly curved and without ornamentations) measuring 105 – 117 microns. Setal formula (averages, n = 6, relative to ab; d’d’ absolute) (aa: ab: bc: cd: dd) at 10: 3.7: 1: 4.7: 0.9: 9.2 and 0.54 d’d’ = 1 / 2 C (= cd dorsal); at 30: 5.7: 1: 5.3: 1: 14.2 and 0.7 d’d’ = 1 / 2 C (= cd slightly dorsal); ten segments before anus (only entire individuals): 2.9: 1: 2.4: 1: 14.4 and 1.2 d’d’ = 1 / 2 C (= cd ventral). Clitellum: Annular, from ½ 12 – ½ 16, very thin (Fig. 1 A). Dorsal pores: absent. Spermathecal pores: not visible. Female pores: In 14, presetal in AB. Male pores: In two aclitellate individuals (Paratypes IEOL 6565, 6567; Fig 1 C, 1 D) it was observed from the outside, a thin band running throughout segment 11 in BC, which is assumed that corresponds to the gonoduct from male funnels of 10. In these two individuals and in the clitellate holotype (3190) (Figs 1 B, 1 C, 1 D) a more thick, clearly visible and slightly undulating band of cream color was also observed extending throughout segments 12 – ¾ 13, in BC, which corresponds to male gonoducts from the male funnels of 10 and 11, that run together. If male pores are located at the end of this duct, then they are in segment 13, postsetal and in BC. Genital marks: None, only setae ab of segment 15 enclosed within small and barely visible protuberances (seen only in the clitellate holotype). Nephridial pores: not visible. Lateral bands: In all non clitellated individuals (paratypes IEOL 2312, 6565, 6566, 6567, 6568) lateral bands were observed in BC close to C, from segments 2 or 3 to segments 14, 16 or 20; in one individual they enclosed CD and were observed in more posterior segments. They are seen as depressions of external cuticle, and probably correspond to longitudinal muscle layers. Internal (Fig. 3 A, B). Septa: All thin and membranous. Septal glands: In segments 5, 6, 7, 8 and 9 (in 10 incipient) a kind of paired glands were observed in dorsolateral walls, attached to corresponding posterior septum (Fig. 3 B, 4 A, 4 B). They were profusely mixed with lateral vascular commissures. Gizzard, Caeca and Typhlosole: Absent. Esophagus: Moniliform, with the same structure along all the body; even that no distinction in the transition to intestine was observed, in one individual and from segment 11 backwards there was an increase in length of the digestive tract in each segment (Fig. 3 B). Dorsal vessel: Single, visible throughout (Fig. 3 B, 4, 5); in one individual size increases from segment 14 backwards. Vascular commissures: Long and contorted vessels are observed in segments 5?, 6 – 11; they are embedded with septal glands (Fig. 5 A). Ventral vessel: Present. Nephridia: Ventral holonephridia observed from segment 11 backwards. They are covered by a granular peritoneal layer that gives them an ovoid, flat appearance (Fig. 5 B, C, D); this ovoid portion is free, floating in the coelom, and not attached to septa. Nephridia gradually increase in size and from segment 15 they attain its maximal size. They are exonephric (Fig. 5 C) (opening in ab) and probably open (stomate), even though nephrostomes were not seen. Testicles and male funnels: Holandric. Iridescent male funnels in 10 and 11; those of 11 bigger than those of 10 (Fig 3 B, 6 A); testes not seen. Both funnels are enclosed by seminal vesicles. Male gonoduct: A gonoduct that comes from male funnels of s. 10 enters the body wall in 11, very close to 10 / 11; a second gonoduct coming from s. 11 funnels, enters the body wall in 12, very close to 11 / 12. Both male conducts run intraparietal over segments 11, 12 and 3 / 4 of segment 13 (deduced from the external observed, undulating cream band) (Fig. 1 B, C, D). Accordingly, the male pore would be in 13, postsetal and in BC. Seminal vesicles: In the clitellate holotype (IEOL 3190) and one semiadult paratype (IEOL 6565), there is one pair of sausage-shaped vesicles in segment 11; these vesicles contain male funnels, project in posterior direction from segment 12 until segments 18 or 20 and run dorso-lateral of the digestive tract, completely covering it (Fig 3 A). In segment 10 there is another pair of vesicles that also contain male funnels, that run in anterior direction before they turn over 180 ° and which, apparently, are limited to the following segment 11. In a semiadult, we observed dense seminal coagulum in segments 11 and 12 that could be incipient seminal vesicles; they completely covered the esophagus. Genital glands: Two pairs of lobulate, rosette-shaped glands lie internally on the ventral body wall of segments 13 and 14 (Fig 3, 6), the anterior being the larger one. They are composed of several short lobules almost fixed to the ventral wall; they could be the equivalent of prostatic glands found in other crassiclitellate worms. We were unable to see any intraparietal connection with male gonoduct. They are superficially like the septal glands present in the anterior segments. No tubular, atrium-like glands were present. Ovaries and female funnels: Large ovaries in segment 13 projecting from the ventral portion of 12 / 13 at AB row level, and extending over the entire segment (Fig 6 B, 7 A). No strings of oocytes or mature ovules were observed. In one individual the ovaries presented a large neck with a kind of developing eggs in the distal part. Female funnels on the ventral wall of segment 13 very close to 13 / 14, in AB, and with a duct opening in segment 14 before seta ab. Ovisacs: Apparently the septum 13 / 14 is modified as dorsolateral paired sacs that contain the corresponding seminal vesicles of 11, and that extends backwards until segments 18 or 20 (Fig. 3 A). No oocytes were observed within the posterior end of these membranous sacs. Spermathecae: One pair of spermathecae without diverticulum in segment 10 (Fig. 3 A, 7 B) opening ventrally in 9 / 10, at AB. In the holotype the left spermatheca was empty. Conversely, the right spermatheca presented iridescence near the union with duct (Fig 7 B) and was completely covered by the seminal vesicles and fixed to male funnels. The duct of this spermatheca turned out 180 ° to connect with a sausage-shaped ampulla. Width and length of ampulla was twofold and shorter than, respectively, width and length of duct. Total length of spermathecae was 580 µm (probably more, as the duct was not complete), 370 µm of this corresponding to ampulla, and 210 µm to duct.

Etymology: The name refers to the swampy habitat of the species.

Localities and material. Mexico, Chiapas, Chajul village, municipality of Marques de Comillas, 1 km south of town, close to the water tank. A patch of small palms (Bactris balanoidea and B. trichofila) within a tropical rain forest over semi-inundated gley soils at 30 – 40 cm depth, 16 ° 06 ' 24 " N, 90 ° 55 ' 24 " W, 190 m asl (Figure 8), one clitellate adult, three non-clitellate adults, two juveniles 12 / 30 / 1982, C. Fragoso. Holotype. Fragmented clitellate adult: IEOL 3190. Paratypes. Three entire semi-adults with some internal genitalia: IEOL 2312, 6565, 6567; two juveniles: IEOL 6566, 6568.