Protozapotecia acaxetlensis and Protozapotecia oyametlensis, two new earthworm species (Crassiclitellata: Acanthodrilidae) from Mexican central mountains Author Cervantes, Gabriela Author Fragoso, Carlos text Zootaxa 2018 2018-10-05 4496 1 431 439 journal article 29252 10.11646/zootaxa.4496.1.32 a2d943e5-355c-42f9-b53d-276769d20dca 1175-5326 1446951 87981062-CA21-40E3-80FA-1622CD7F9864 Protozapotecia oyametlensis sp. nov. ( Figures 2 , 3E–H ) Localities and material. Mexico : (1) Veracruz , Municipality of Perote. Cofre de Perote National Park, Road El Conejo–La Cima. Small patch of fir forest, within a dark soil (andolsol) at 30–40 cm depth. N19°31'16.5" , W97°9'29.90" , 3475m asl. Two clitellate adults, June 24th, 2008 (Collectors C. Fragoso and A. Angeles). Holotype . Clitellate adult, IEOL 4198 ( Fig. 2 ). Complete specimen, last 10 segments ( 3.3 mm ) amputated to obtain DNA . Paratype. One clitellate adult, IEOL4199. Complete specimen, last 10 segments ( 3.3 mm ) amputated to obtain DNA. Description. External. Length 66–85 mm (mean= 75.5 mm , n= 2; holotype = 66.5 mm ); post-clitellar diameter 2.7mm (n=2). Segments 99–101 (mean= 100, n= 2; holotype = 99). Colour absent in living specimens, beige after fixation in formalin. Prostomium closed epilobous, reaching 1/2–2/3 of the first segment. Secondary annulation: one postsetal in 4–6 , one presetal and one or two postsetal in 7–12 , multiple behind clitellum. Setae closely paired, eight per segment, visible from segment 2 in ventral position. Setal formula ( aa:ab:bc:cd:dd ) (n=2) in 10 : 4.5:1.0:5.4:1.3:21.1; in 30 : 5.5:1.0:4.5:1.3:18.6; in the last segments: 5.3:1.0:5.6:1.5:20.6. Genital setae absent. Penial setae present in 18 and 20 but difficult to observe ( Fig. 3 E, F ); small ( 0.69 mm length) thin compared to length of worms, bracket shaped, without ornamentations; distal part almost straight and with an acute delicate apex ( Fig. 3G, H ). Saddle-shaped clitellum in ½ 13 – ½ 18 ventrally interrupted in AB , clitellum color dark orange. Dorsal pores present all along the body, the first dorsal pore in 11/12, 12/13. Spermathecal pores paired in 8 and 9 , presetal and slightly outside B . Female pores in 14 , presetal in AB , within a papilla. Two pairs of prostatic pores in 18 and 20 in AB , joined by almost straight and superficial seminal belts, which run along AB . Male pores at the equator of 19 in AB . Setae ab absent in 18 ; in 19 and 20 internally present as penial setae. Genital markings: five unpaired and midventral ovoid papillae: in 10/11, 11/12, 12/13, 15/16 and 22/23, reaching B or AB ( holotype ) or two in 11/12 and 22/23 reaching AB ( paratype ). Internal. Funnel-shaped septa in 5/6 , 6/7 , 7/8 , 8/9 and 9/10 ; 10/11 and 11/12 not funnel-shaped. Those of 5/6–12/13 slightly muscular, in holotype septa 6/7–9/10 more thick and muscular; all muscular septa joined by dorsal and lateral connective tissue fibers. From 13/14 and backwards septa thin. Two gizzards, in 5 and 6 , with same width but different length (largest in 6) ; both gizzards separated by a muscular septum. Esophageal lamellae present in 8–12 , small in 10–12 , larger in 8 , 9 , where almost occluding lumen. Esophagus with abundant vascularization in 9 , less in 8 and scarce or absent in 10–13 . Irregular presence of septal glands in 14 . Calciferous glands absent, although esophageal lamellae, vascularization and dilatation of walls in segments 8 and 9, suggest absorption function. Intestine beginning in 15/16 (holotype and paratype). Laminar typhlosole starting as a thin line from 17 to 20 ; in 21–39 reaching its maximal size, ending smaller in segment 40 . Lateral typhlosoles absent. Intestinal caeca absent. Sacculated intestine in 16–19 . Lateral hearts in 7–9 . Latero-esophageal hearts in 10–12 . Supra-esophageal vessel visible in 10–12 . Single dorsal vessel visible from segment 5 . Holoic; nephridia without vesicles, parietal from 12 and backwards, opening to the exterior in C and with a nephrostome in AB ; in 8–11 also parietal but closer to midventral line and occluded by funnel septa. Holandric. Testes and iridescent, slightly plicated male funnels in 10 and 11 ; those of 11 slightly larger. Two pairs of acionous seminal vesicles in 11 and 12 covering all esophagus and joining in dorso-lateral position 10/11 and 11/12 , respectively; those of 12 larger. Two pairs of tubular prostates in 18 and 20 , small, coiled with two or three folds, usually limited to their corresponding segments and occasionally extending a posterior segment. Muscular duct narrower and shorter than the glandular part. Male gonoduct over parietes, running almost straight outside B until 18 , where it enters body wall. Ovaries one pair in 13 , fan folded shaped, and composed of long strings of eggs; female funnels attached to the floor close 13/14. Two pairs of spermathecae with sessile diverticulum, in 8 and 9 ( Fig. 2 I –IV). Diverticulum with a bilobed iridescent chamber, perpendicular to the axis of duct and ampulla; ampulla longer than duct and diverticulum. FIGURE 2. Protozapotecia oyametlensis sp. nov. A. Holotype IEOL 4198. Anterior region, external ventral view. Spermathecae in dorsal view: left spermatheca from segment 8 (I), right spermatheca from segment 8 (II), left spermatheca from segment 9 (III), right spermatheca from segment 9 (IV). di=diverticulum, am= ampulla, gm=genital marking, pp=prostatic pore, mp=male pore. Scale bar 1 mm. FIGURE 3. Protozapotecia acaxetlensis sp. nov. (A–D) and Protozapotecia oyametlensis sp. nov. (E–H), penial setae of segment 18 . A–D, scanning electron micrographs, E–H, photograph of optical microscope. A. Penial seta a . B. Penial seta b . A, B in lateral view. C. Apex of seta a . D. Apex of seta b . E. Penial seta b . F. Penial seta a . G. Apex of seta b . H. Apex of seta a . Scale bars = 100µm (A, B, E, F), 1µm (C), 2µm (D), 20µm (G, H). Etymology. The word oyametl means "fir" in nahuatl language, and it alludes to the type of vegetation where this species was found and to which it is presumably restricted. Remarks. By the presence of prostatic pores in 18 and 20 , P. oyametlensis sp. nov. is closely related to P. acaxetlensis sp. nov. , and clearly separated from the other species of the genus. From the last species it is separated by the position of seminal vesicles (in segments 11 , 12 in P. oyametlensis sp. nov . vs. in 9 , 11 , 12 in P. acaxetlensis sp. nov. ) and the position of large esophagus lamellae (segments 8, 9 vs. 11–14 ). Differences between P. oyametlensis sp. nov. and the other species of the genus are presented in Table 1 . This species has previously been referred to IEOL4199 " Protozapotecia sp. nov. 1" ( Cervantes et al. 2014 ), and " Zapotecia ( Protozapotecia ) sp. nov.1 " ( Cervantes 2012 ). TABLE 1. Morphological differences among species of the genus Protozapotecia . Characters common to all species are not listed (e.g., body colour absent, spermathecae at 8, 9, testes holandric).
Species Length (mm) Prostomium 1 st dorsal pore Last hearts Lamella Acinous septal e glands Intestine beginning
Protozapotecia acaxetlensis 55–78 closed 10/11 12 9–14 present 15/16
sp. nov. epilobous
Protozapotecia angelesae Rodríguez & Fragoso, 2017 40–67 open epilobous 8/9 12 11– 12 absent 14
Protozapotecia aquilonalis James, 1993 53–95 open epilobous 9/10 12 11–13 * present 15 or 14
Protozapotecia australis James, 1993 54–73 open epilobous 6/7 12 11–13 * present 15
Protozapotecia centralis 28–70 closed 4/5 12 13 absent 14
Rodríguez & Fragoso, 2017 epilobous
Protozapotecia cubensis 95–130 closed 4/5 13 13–15 absent 17
Rodríguez & Fragoso, 2017 epilobous
Protozapotecia oyametlensis 66–85 closed 11/12, 12/13 12 8, 9 present 15/16
sp. nov. epilobous
continued.
Species Typhlosole Seminal Genital vesicles setae Penial Tubular setae prostates Genital markings
Protozapotecia acaxetlensis sp. nov. from 17 9 , 11 , 12 absent present 18 and 20 10/11–15/16, 22/23,23/24
Protozapotecia angelesae Rodríguez absent & Fragoso, 2017 11 , 12 absent absent 17 and 19 22 , 23
Protozapotecia aquilonalis James, 1993 from 19 11 , 12 absent present 17 and 19 10/11
Protozapotecia australis James, 1993 from 19 11 , 12 present present 17 and 19 10/11
Protozapotecia centralis Rodríguez & Fragoso, 2017 from 17 11 , 12 absent present 17 and 19 6 , 14 , 16 , 17 , 20
Protozapotecia cubensis Rodríguez & Fragoso, 2017 from 20/21 11 , 12 absent present 17 and 19 absent or not developed
Protozapotecia oyametlensis sp. nov. from 17 11 , 12 absent present 18 and 20 10/11, 11/12, 12/13, 15/15 and 22/23
* Character uncertain ( James 1993: "Pebbly internal texture in 11–13" ).