Travisia pupa Moore, 1906

Tovar-Hernández, María Ana, Burgess, Dany E., León-González, Jesús Angel De & Hendrickx, Michel E., 2025, Travisiid worms from deep water off Western Mexico with the establishment of a new species (Annelida: Travisiidae), Zootaxa 5729 (2), pp. 349-369 : 355-359

Travisia pupa Moore, 1906

( Figs 4‒5 )

Travisia pupa Moore, 1906: 228‒231 , Pl. XI, fig. 23; 1923: 221.— Uschakov, 1950: 209; 1955: 324, fig. 120F‒G.— Berkeley & Berkeley, 1952: 89, figs 181‒182.— Levenstein, 1970: 214.— Dauvin & Bellan, 1994: 173.— Hobson, 1976: 139.— Hobson & Banse, 1981: 62, fig. 13j. — Blake, 2000: 163, fig. 7.8.— Imajima, 2009: 139‒140, fig. 45A‒E.— Alalykina, 2020: 193.

Material examined. Washington State Department of Ecology. Puget Sound , Washington, USA: Elliott Bay , Sta. 187, 103 m, 47°36’26”N, 122°21’32”W, coll. PSEMP Urban Bays Monitoring, 16 June 2021 ( 1 specimen) . Hood Canal , north of Seabeck, Sta. 222, 126 m, 47°40’42”N, 122°48’53”W, coll. PSEMP Long-term Monitoring, 01 June 2022 ( 3 specimens) . Saratoga Passage , Sta. 19, 126 m, 48°5’53”N, 122°28’17”W, coll. PSEMP Long-term Monitoring, 01 May 2023 ( 1 specimen) .

Diagnosis. Travisia with annulate branchiae from chaetiger 2, 24‒25 pairs of branchiae and 26‒27 chaetigers. Mid-ventral groove present. Nuchal organs small. Parapodial lappets present along body except for the last body quarter.

Description. Specimens 23‒75 mm long, 9‒18 mm wide with 26‒27 chaetigers (n: 5). Body pale to cream-colored in alcohol; robust, grub-like with rugose epidermis ( Fig. 4A‒B ). Prostomium small, with rounded margin, smooth ( Figs 4C , 5B ). Nuchal organs dorsolateral, small narrow splits ( Fig. 5B ). Eyes absent. Mouth ventral between chaetigers 1 and 2: upper lip with elongate annulations of chaetiger 1, lower lip with short annulation of chaetiger 2 ( Figs 4B , 5A, D ). Mid-ventral groove present along body, more visible in anterior and posterior segments ( Figs 4B , 5A, D, F ).

Peristomium uniannulate. Chaetiger 1 biannulate. Chaetigers 2 to 13‒14 triannulate. Chaetigers 14‒15 to 17‒ 18 biannulate. Following chaetigers to the end uniannulate. Peristomium triangular shaped dorsally, lateral sides oblique, distal margin rounded, basal margin four times wider than anterior margin, covered with small, spherical papillae ( Figs 4C , 5B ).

First chaetiger with dorsal and lateral sides covered by rounded packages of equal size spherical papillae ( Figs 4C , 5B ). From chaetiger 2 towards posterior end, each segment with spherical papillae around body as follows: chaetigers 2‒16 with spherical papillae dorsally ( Fig. 5B ), large papillae only on posterior edges of annulations ( Fig. 5C ), anterior papillae to posterior edges considerably smaller; chaetigers 17‒18 to body end with epithelium fully covered with spherical papillae, increasing gradually in size from anterior margin of each segment to posterior margin ( Fig. 5E ). Last 11 segments with inferior margins flap-like, with ventral margin longer than dorsal margin ( Fig. 5E ).

Branchiae slightly annulate, 24‒25 pairs starting at chaetiger 2 and extending to chaetigers 24‒26. First pair short, increasing gradually in size towards chaetiger 12, decreasing in size gradually to posterior end ( Figs 4A‒B , 5A ).

Parapodia biramous. Parapodial lappets present on neuropodia of chaetigers 2‒15 ( Figs 4A‒B, D , 5A, D, F ). Each neuropodial lappet triangular, composed of aglutinated papillae with largest papillae on top ( Figs 4D , 5D ). Notopodial lappets absent ( Fig. 5C ). Chaetigers 15 or 16 with a gradual transition of parapodial lappets rather than abrupt transition to end without parapodial lappets.All noto- and neurochaetae iridescent, spinulose capillaries. Last 4‒5 segments achaetous.

Interamal sensory pores small, between notochaetae and neurochaetae from chaetiger 1 until near last segments. Lateral neuropodial nephridiopores present along chaetigers 3 to 14.

Pygidium short, terminal or shifted to side, thick, with 10 lobes ( Figs 4A‒B , 5G ).

Methyl green staining. Branchiae and mid-ventral groove unstained. Rest of the body is uniformly stained ( Fig. 5 ).

Remarks. Travisia pupa was originally described by Moore (1906) from the Gulf of Georgia, the Behm Canal, and Kasaan Bay ( Alaska, USA), in sediments from 32‒438 m depth. Unfortunately, Moore (1906) did not include illustrations of body structures or papillation, except for a short, anterior area of a chaeta ( Moore 1906: pl. XI, fig. 23). Posteriorly, T. pupa has been reported in Alaska, British Columbia, California, Sea of Okhotsk, Bering Sea, Japan Trench, and Kuril-Kamchatka Trench, in shelf and slope depths ( Moore 1923; Uschakov 1950, 1955; Berkeley & Berkeley 1952; Levenstein 1970; Hobson 1976, Hobson & Banse 1981; Blake 2000; Imajima 2009), though in all cases poorly illustrated. Among all these records, that of Imajima (2009: fig. 45A‒E, Honshu/ Japan) includes detailed drawings regarding body papillation.

Moore (1906) noted that the size of T. pupa was 24‒82 mm long, 8‒32 mm width, 31‒32 segments, stating “... the number is not correlated with the size of the worm, the largest two having thirty-one and the smallest thirty-two”, 22‒25 pairs of branchiae, and interamal sensory pores starting at chaetiger 1. While examining the type material, Hobson (1976) reported the following features: worms 40‒85 mm long; 30‒32 segments: 1 achaetous anterior segment, 25‒27 chaetigerous segments, and 3‒6 achaetous pre-anal segments; and 21‒25 pairs of branchiae. Specimens from Puget Sound examined herein measure 23‒75 mm long, 9‒18 mm width, 31‒32 segments, 26‒27 chaetigerous, 4‒5 achaetous pre-anal segments, and 24‒25 pairs of branchia.

In her contribution, Hobson (1976) declared T. foetida Hartman, 1969 , a species described from Redondo Canyon, California to 256 m depth, a junior synonym of T. pupa , a synonym followed by Blake (2000) in his report of T. pupa to Santa Maria Basin and Western Santa Barbara Channel ( California, USA) in 91‒197 m depth; however, T. foetida was still included as a valid name in Blake & Maciolek (2019), Rizzo & Salazar-Vallejo (2021) and Read & Fauchald (2025).

Fauchald (1972) reported the presence of T. foetida in many localities from western Mexico (off Baja California to Acapulco, and in the southern part of the Gulf of California, from 1985‒3383 m depth), and it was therefore suggested that the distribution of T. pupa reaches western Mexico ( Blake 2000; Imajima 2009). Fauchald (1972) reported a uniform number of 27 chaetigers in all specimens he reviewed and identified as T. foetida ( 47 specimens), and the presence of a dorsally and anteriorly directed peristomial lappet. In his drawing ( Fauchald 1972: pl. 49, fig. d) this structure is triangular, as in T. pupa here reported from Puget Sound, but the position and size of nuchal organs are peculiar in Fauchald´s specimens: circular, large (as long as prostomium) and placed on the base of the peristomium versus nuchal organs small, as narrow splits located on the anterior margin of the peristomium in T. pupa . This difference is significant and provides support to reject the synonymy proposed by Hobson (1976). It is desirable, however, to carefully revise the types of T. foetida and T. pupa and Fauchald´s material to prove the validity of T. foetida and its presence in Western Mexico.

Specimens here identified from Puget Sound as T. pupa have a mid-ventral groove running along the body (also see Burgess 2018b). Travisia pupa was included in the subgroup B2 (species without mid-ventral groove) by Plathong et al. (2023), but the presence of a mid-ventral groove clearly indicates that T. pupa belongs to subgroup B1 (species with mid-ventral groove). However, we believe that the use of this feature as a diagnostic character demands further study. In the case of Travisia amoyanus Yang, Wu, Wang, Zhao, Hwang & Cai, 2022 , described from the intertidal-subtidal, in Xiamen, China ( Yang et al. 2022), the original description suggests that some specimens lack a midventral groove: “Midventral groove absent, if have, present from last four segments”. In Travisia pupa from Puget Sound the mid-ventral groove is clearly visible in anterior and posterior segments only. This raises the question about the presence or absence of a midventral groove. It might represent a variable feature according to ontogeny or fixation, making this character questionable for its current use in classification or recognition of taxa. A further argument in favor of using this character with caution is provided by Santos (1977), who stated that “The relative development of the midventral groove in T. hobsonae [ Santos, 1977] appears to be related to the preparation of the specimens. Those that are incompletely narcotized before fixation show a pronounced groove, whereas the specimens that are properly narcotized exhibit a very slight midventral groove”.

Figures

FIGURE 4. Travisia pupa Moore, 1906.A) Body, lateral view, B) same, ventral view, C) prostomium and anterior segments, dorsal view, D) parapodial lappets of chaetigers 5, 6 and 7. Abbreviations: mo—mouth, no—nuchal organ, pr—prostomium, vg—ventral groove, 1, 2, 5, 6, 7— refers to the number of chaetiger. Scale bars: A–D) 1 mm. A–D: specimen from Puget Sound, Washington.

FIGURE 5. Travisia pupa Moore, 1906 stained with methyl green.A) body, ventral view, B) prostomium and anterior segments, dorsal view, C) anterior segments, dorso-lateral view, D) anterior segments, ventral view, E) posterior segments, lateral view, F) segments from mid-body, ventral view, G) posterior segments and pygidium, lateral view. Abbreviations: dm—dorsal margin, mo—mouth, no—nuchal organ, pr—prostomium, vg—ventral groove, vm—ventral margin, 1, 2—refers to the number of chaetiger. Scale bars: A–D, F) 1 mm, E, G) 0.5 mm. A–G: specimen from Puget Sound, Washington.