Integrated taxonomy of three new species of Glyphiulus Gervais, 1847 (Diplopoda, Spirostreptida, Cambalopsidae) from Laos Author Likhitrakarn, Natdanai Author Jeratthitikul, Ekgachai Author Jirapatrasilp, Parin Author Wesener, Thomas text Raffles Bulletin of Zoology 2024 2024-01-26 72 42 61 journal article 10.26107/RBZ-2024-0003 2345-7600 10788647 DC2582A7-C5DD-4726-9B59-3F2C00E9485E Glyphiulus pseudocostulifer , new species ( Figs. 1 , 2 ) Holotype . Male ( ZFMK MYR-10009 A ( TW411 )), Laos , Oudomxay Province , Ban Chom Ong , Tham Ketlin (F47- 120-011), 20°43′19″N , 101°45′59″E , 28 December 2014 , leg. M. Vandermeulen. Paratypes . 1 female ( ZFMK MYR-10009 ), same locality, together with holotype . 1 male ( CUMZ MYR-10008 A ( TW414 )), same locality, 27 December 2014 , leg. M. Vandermeulen. 1 male ( ZFMK MYR-9994 A ( TW400 )) 1 female ( ZFMK MYR-9994 B ( TW401 )), Laos , Oudomxay Province , Ban Houay Soy , Tham Houay Soy (F47-120- 015), 20°42′37″N , 101°45′50″E , 29 December 2014 , leg. H. Steiner and I. Ermakova. 1 male ( ZFMK MYR-10004 A ( TW412 )), Laos , Oudomxay Province , Ban Chom Ong , Tham Hou Nguak (F47-120-010), 20°42′54″N , 101°45′38″E , 27 December 2014 , leg. D. Fröhlich and M. Laumanns. Etymology. To emphasise the obvious similarities to G. costulifer Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 . Noun in apposition. Diagnosis. This new species is especially similar to both G. steineri , new species , and G. costulifer , from Luang Prabang Province , Laos ( Golovatch et al., 2007b ). With G. costulifer , it shares the following diagnostic characters: the unique legs 1, and anterior and posterior gonopod structural details. It differs from G. costulifer by the carinotaxic formula of the collum: 1–5+6a+pc+ma ( Fig. 1A, B ) (vs. 1–6+7a+pc+ma) and the carinotaxic formula of the midbody rings/segments: 2/2+I/i+3/3+I/i+2/2 ( Fig. 1A, B, D–H ) (vs. 2+I/i+3/3+I/i+2), coupled with the paramedian processes on male coxae 1 being much higher than the prefemur ( Fig. 2B, C ) (vs. lower than the prefemur), the apicomesal processes (d) on the anterior gonopods being long and slender ( Fig. 2H, I ) (vs. shorter and digitiform) and the presence of a very long and slender flagellum process (f) on the posterior gonopods ( Fig. 2J, K ) (vs. a shorter flagellum process). The differences between G. pseudocostulifer , new species , and G. steineri , new species , are found in the carinotaxic formula of the midbody rings as well as the gonopods (see further details below). Table 2. Matrix of the average uncorrected p-distance (%) based on 671-bp COI barcoding region between 17 Glyphiulus species and some related taxa. Interspecific divergences (and intraspecific distances in the case of G. pseudocostulifer ) are below the diagonal; intraspecific divergences are marked in bold.

Taxa	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28	29	30	31	32	33	34	35	36
Glyphilulus polytrichus (JXK222)
Hypocambala zizhongi (JXK165)	0,192
Pericambala sinica (JXK155)	0,230	0,214
Pericambala aramulus (JXK392)	0,228	0,214	0,002
Cambala annulata (JXK517)	0,236	0,223	0,215	0,212
Plusioglyphiulus saksit (CUMZ CAM021)	0,204	0,206	0,228	0,224	0,214
Plusioglyphiulus erawan (CUMZ CAM031)	0,197	0,188	0,208	0,208	0,205	0,136
Trachyjulus singularis (CUMZ CAM107)	0,225	0,228	0,247	0,247	0,231	0,201	0,216
Trachyjulus magnus (CUMZ CAM070)	0,188	0,208	0,223	0,231	0,246	0,206	0,200	0,215
Trachyjulus unciger (CUMZ CAM079)	0,232	0,245	0,230	0,239	0,223	0,243	0,246	0,238	0,206
Trachyjulus phylloides (CUMZ CAM027)	0,214	0,203	0,217	0,220	0,215	0,216	0,206	0,207	0,200	0,195
Trachyjulus bifidus (CUMZ CAM061)	0,219	0,241	0,243	0,246	0,218	0,210	0,219	0,215	0,201	0,203	0,151
Glyphiulus duangdee (CUMZ CAM022)	0,195	0,195	0,223	0,227	0,222	0,212	0,195	0,235	0,237	0,227	0,213	0,212
Glyphiulus impletus (SCAUWL39)	0,221	0,251	0,245	0,239	0,234	0,203	0,220	0,197	0,227	0,247	0,233	0,224	0,233
Glyphiulus calceus (SCAUWL37)	0,224	0,230	0,254	0,248	0,221	0,208	0,213	0,216	0,219	0,262	0,210	0,215	0,231	0,142
Glyphiulus zorzini (SCAUWL23)	0,166	0,190	0,243	0,242	0,206	0,178	0,174	0,239	0,221	0,248	0,207	0,199	0,181	0,210	0,211
Glyphiulus speobius (SCAUG62)	0,175	0,188	0,197	0,202	0,179	0,202	0,183	0,210	0,215	0,233	0,183	0,199	0,149	0,212	0,204	0,167
Glyphiulus proximus (SCAUG61)	0,171	0,195	0,212	0,215	0,202	0,183	0,175	0,227	0,189	0,228	0,181	0,198	0,181	0,212	0,196	0,170	0,122
Glyphiulus deharvengi (SCAUG15)	0,206	0,214	0,230	0,237	0,217	0,207	0,192	0,227	0,207	0,233	0,207	0,224	0,193	0,221	0,199	0,195	0,164	0,151
Glyphiulus granulatus (JXK196)	0,208	0,193	0,227	0,227	0,214	0,206	0,201	0,225	0,210	0,225	0,208	0,201	0,206	0,207	0,215	0,179	0,160	0,186	0,180
Glyphiulus quadrohamatus (JXK072)	0,201	0,204	0,234	0,234	0,206	0,192	0,184	0,227	0,208	0,245	0,179	0,201	0,169	0,229	0,215	0,177	0,171	0,182	0,158	0,199
Glyphiulus calceus (JXK061)	0,219	0,228	0,256	0,256	0,241	0,212	0,223	0,221	0,217	0,256	0,214	0,219	0,236	0,146	0,021	0,225	0,219	0,214	0,212	0,217	0,221

Taxa	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	21	22	23	24	25	26	27	28	29	30	31	32	33	34	35	36
Glyphiulus impletus (JXK002)	0,212	0,239	0,228	0,228	0,230	0,195	0,214	0,192	0,221	0,238	0,215	0,217	0,225	0,022	0,129	0,210	0,203	0,201	0,214	0,201	0,212	0,131
Glyphiulus guangnanensis (JXK051)	0,184	0,197	0,219	0,219	0,201	0,197	0,177	0,236	0,195	0,215	0,190	0,192	0,199	0,216	0,206	0,157	0,177	0,169	0,186	0,199	0,193	0,208	0,206
Glyphiulus foetidus (JXK059)	0,221	0,243	0,247	0,247	0,234	0,228	0,247	0,228	0,221	0,228	0,227	0,230	0,228	0,245	0,221	0,227	0,199	0,236	0,238	0,221	0,225	0,228	0,230	0,221
Glyphiulus formosus (JXK278)	0,215	0,219	0,247	0,239	0,231	0,211	0,233	0,224	0,208	0,271	0,217	0,219	0,220	0,166	0,168	0,205	0,210	0,205	0,227	0,215	0,210	0,168	0,157	0,221	0,225
Plusioglyphiulus steineri (SMF)	0,199	0,199	0,227	0,229	0,232	0,161	0,159	0,227	0,203	0,230	0,207	0,231	0,203	0,223	0,205	0,189	0,189	0,172	0,210	0,197	0,210	0,204	0,217	0,204	0,236	0,228
Plusioglyphiulus deharvengi (SMF)	0,199	0,203	0,208	0,207	0,214	0,122	0,128	0,204	0,210	0,228	0,206	0,194	0,194	0,205	0,209	0,177	0,183	0,175	0,195	0,188	0,186	0,212	0,197	0,196	0,216	0,208	0,167
Glyphiulus pseudocostulifer , new species (ZFMK MYR9994A)	0,184	0,223	0,230	0,236	0,222	0,210	0,206	0,235	0,227	0,247	0,204	0,222	0,225	0,233	0,231	0,198	0,175	0,193	0,225	0,212	0,217	0,238	0,212	0,175	0,221	0,261	0,216	0,212
Glyphiulus pseudocostulifer , new species (ZFMK MYR9994B)	0,186	0,223	0,238	0,243	0,223	0,218	0,209	0,240	0,230	0,251	0,209	0,224	0,228	0,240	0,233	0,203	0,184	0,200	0,230	0,218	0,218	0,244	0,218	0,179	0,231	0,260	0,222	0,217	0,012
Glyphiulus pseudocostulifer , new species (ZFMK MYR10009A)	0,169	0,217	0,227	0,231	0,215	0,212	0,199	0,240	0,221	0,234	0,201	0,212	0,228	0,241	0,233	0,193	0,186	0,192	0,219	0,201	0,221	0,234	0,214	0,166	0,208	0,250	0,222	0,215	0,057	0,064
Glyphiulus pseudocostulifer , new species (ZFMK MYR0004A)	0,188	0,227	0,243	0,246	0,228	0,215	0,210	0,246	0,235	0,252	0,212	0,231	0,232	0,242	0,239	0,202	0,183	0,199	0,237	0,221	0,225	0,247	0,221	0,184	0,230	0,271	0,222	0,221	0,024	0,036	0,069
Glyphiulus pseudocostulifer , new species (ZFMK MYR10008)	0,169	0,214	0,228	0,234	0,215	0,210	0,198	0,238	0,221	0,234	0,207	0,209	0,224	0,238	0,241	0,195	0,186	0,193	0,224	0,199	0,221	0,239	0,212	0,171	0,214	0,255	0,221	0,215	0,057	0,064	0,009	0,066
Glyphiulus pseudocostulifer , new species (ZFMK MYR10008A)	0,169	0,217	0,227	0,231	0,215	0,209	0,197	0,240	0,221	0,238	0,206	0,215	0,228	0,241	0,238	0,192	0,184	0,192	0,225	0,204	0,223	0,239	0,215	0,171	0,212	0,254	0,219	0,213	0,055	0,066	0,010	0,058	0,010
Glyphiulus steineri , new species (ZFMK MYR6203)	0,184	0,204	0,227	0,229	0,234	0,201	0,212	0,219	0,237	0,261	0,215	0,231	0,197	0,232	0,221	0,201	0,198	0,196	0,218	0,210	0,206	0,221	0,227	0,166	0,212	0,222	0,209	0,207	0,206	0,209	0,204	0,216	0,204	0,203
Glyphiulus semicostulifer (CUMZ CAM057)	0,186	0,188	0,208	0,215	0,217	0,191	0,173	0,242	0,204	0,237	0,210	0,219	0,209	0,235	0,229	0,168	0,204	0,172	0,209	0,184	0,192	0,225	0,225	0,149	0,234	0,224	0,200	0,185	0,181	0,185	0,188	0,187	0,188	0,188	0,176
Glyphiulus subbedosae (CUMZ CAM058)	0,197	0,203	0,247	0,244	0,225	0,210	0,200	0,242	0,222	0,257	0,225	0,228	0,166	0,242	0,217	0,189	0,190	0,195	0,209	0,204	0,201	0,221	0,232	0,188	0,197	0,230	0,219	0,208	0,216	0,216	0,214	0,223	0,219	0,219	0,193	0,188

Fig. 1. Glyphiulus pseudocostulifer , new species , male holotype (ZFMK MYR10009A). A–C, anterior part of body, lateral, dorsal, and ventral views, respectively; D, E, midbody rings, dorsal and lateral views, respectively; F, cross-section of a midbody ring; G–I, posterior part of body, lateral, dorsal, and ventral views, respectively. Fig. 2. Glyphiulus pseudocostulifer , new species , male holotype (ZFMK MYR10009A). A, gnathochilarium, ventral view; B, C, leg pair 1, oral and caudal views, respectively; D, leg pair 2, caudal view; E, leg pair 3, caudal view; F, midbody leg; G, antenna, lateral view; H, I, anterior gonopods, oral and caudal views, respectively; J, K, posterior gonopods, oral and caudal views, respectively. Abbreviations: ap, apicolateral process; dp, apicomesal processes; f, flagellum process; l, lamelliform lobe. Scale bar: A–G 0.1 mm, H–K 0.02 mm. Description. Length of holotype 26.5 mm ; adult paratypes 17.5–29.5 (males) or 20.8–27. 5 mm long (females); midbody rings/segments round in cross-section ( Fig. 1F ), their width (horizontal diameter) and height (vertical diameter) being similar; width in holotype 0.9 mm ; in paratypes 1.0–1.3 (males) or 1.1–1.5 (females). Colouration in alcohol ( Fig. 1 ), after nine years of preservation, uniformly greenish-golden to grey-brown, dorsal crests and porosteles usually dark brownish ( Fig. 1A, B, D, E, G, H ). Head light yellow, vertex dark brown to red brownish ( Fig. 1A, C ). Antennae, venter and legs yellowish to yellow brown ( Fig. 1A–C, E–G, I ). Eyes blackish ( Fig. 1A, C ). Body with 60p+2a+T ( holotype ) rings; paratypes with 34–52p+5a+T (males) or 44–57p+2–6a+T (females). Eye patches transversely ovoid, each composed of 7–11 blackish, rather flat ommatidia in 3–5 irregular longitudinal rows ( Fig. 1A ). Antennae rather short and clavate ( Figs. 1A, C , 2G ), extending to ring 4 laterally, antennomeres 5 and 6 each with a small distoventral group or corolla of bacilliform sensilla ( Fig. 2H ). Gnathochilarium with a clearly separated promentum ( Fig. 2A ). Head width = ring 2 <collum = midbody ring (close to 13th to 15th) >> ring 4 <5 <6 <7 <3 <8 <9 <10; body abruptly tapering towards telson on a few posterior-most rings ( Fig. 1H ). Postcollar constriction evident ( Fig. 1B ). Collum with low evident crests, carinotaxy formula 1–5+6a+pc+ma ( Fig. 1A–C ), with 6+6 longitudinal crests starting from anterior edge, but both median crests interrupted at about caudal 1/4–1/3, being replaced there by 1+1+1 crests starting from posterior edge, lateral crests being longer than median one. Following metaterga similarly strongly crested ( Fig. 1A–H ), especially from ring 5 on, whence porosteles commence ( Fig. 1A, B ), smaller tubercles in their stead on legless rings in front of telson due to lack of ozopores ( Fig. 1G, H ). Porosteles large, conical, round, directed caudolaterad, wider than high, ozoporiferous crests distinctly divided into two about midway, their frontal halves being higher ( Fig. 1A, B, D–H ). Carinotaxy formulae 2+I/i+3/3+I/i+2 on rings 2 and 3, as well as on the last 1 or 2 leg-bearing and legless rings ( Fig. 1A, B, G, H ); midbody rings showing all dorsal crests subdivided transversely (carinotaxy formulae 2/2+I/ i+3/3+I/i+2/2) and sharper, especially so lateral crests ( Fig. 1D, E, G, H ). Tegument finely alveolate- areolate ( Fig. 1A, B, D, E, G, H ), dull throughout. Fine longitudinal striations in front of stricture between pro- and metazonae, remaining surface of prozonae very delicately shagreened ( Fig. 1D, E ). Metatergal setae absent. Rings 2 and 3 with long pleural flaps. Epiproct ( Fig. 1G–I ) simple, regularly rounded caudally, faintly convex medially, with a rounded ridge in caudal part and an evident axial rib dorsally. Paraprocts regularly convex, each with premarginal sulci medially and a row of sparse setae at medial margin ( Fig. 1I ). Hypoproct slightly concave caudally, with 1+1 strongly separated setae near caudal margin ( Fig. 1I ). Ventral flaps behind gonopod aperture on male ring 7 evident, distinguishable as rather low swellings with rounded flaps bent abruptly caudad ( Fig. 1C ). Legs rather short and stout, on midbody rings about 3/4–3/5 as long as body diameter ( Figs. 1A, C, G, I , 2F ). Claw at base with an evident accessory claw about 1/3–1/4 the length of main claw. Male leg pair 1 highly characteristic ( Fig. 2B, C ) in showing nearly fully developed, 5-segmented telopodites and a pair of rather small, subdigitiform, medially contiguous, but apically diverging coxal processes, higher than prefemur ( Fig. 2B, C ), with a group of a few strong setae at base. Male leg pair 2 nearly normal, claw long and slender, coxa somewhat reduced, femur abbreviated on frontal face; penes broad, oblong-subtrapeziform, each with 4 or 5 setae distolaterally ( Fig. 2D ). Male leg pair 3 modified, coxa especially slender and elongate ( Fig. 2E ). Anterior gonopods ( Fig. 2H, I ) with a typical shield-like coxosternum, this being rather densely microsetose on caudal face ( Fig. 2I ), with a long and high, digitiform, apicomesal process (dp). Telopodite typical, rather small, slender, movable, 1-segmented, lateral in position, with 3 or 4 strong apical setae and a field of microsetae at base ( Fig. 2I ). Posterior gonopods ( Fig. 2J, K ) compact, subtrapezoid, micropapillate medially on oral face ( Fig. 2K ); with a very long and slim flagellum process (f), a pointed tip ( Fig. 2J, K ); a lamelliform lobe (l) higher than caudal piece of coxite ( Fig. 2J ), membranous, wrinkled frontolaterally, with an apical field of coniform microsetae caudally ( Fig. 2K ). Coxite rather smooth, each mediolateral part of coxite with 13–17 strong, short and curved setae ( Fig. 2J, K ).