Pteroptyx maipo Ballantyne, Fu, Shih, Cheng & Yiu, 2011, sp. nov.
Creators: name: Ballantyne, Lesley
name: Fu, Xin Hua
name: Shih, Chun-Hat
name: Cheng, Chui-Yu
name: Yiu, Vor
Publisher: name: Zenodo
Year: 2011
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Coleoptera
Lampyridae
Pteroptyx
Pteroptyx maipo

description: Pteroptyx maipo sp. nov.
(Figs 1–6, 8, 14–17, 33–93)
Type. Holotype male, Hong Kong, China, 22.3° N, 114° E, Hong Kong Wetland Park, 26.v.2010, sweeping, C.H. Shih (AFCD).
Paratypes. Hong Kong, China, 22.3° N, 114° E, Hong Kong Wetland Park, 26.v.2010, sweeping, C.H. Shih 4 males, 2 females, 2 larvae; iii.2010, J. Cheng, 3 males, 1 female, 2 larvae. Mai Po Nature Reserve, 2.vi.2010, C.H. Shih, 4 males, 2 females; 13.viii.2010, V. Yiu, male. Tin Shui Wai, 7.viii.2010, V. Yiu, 4 males, 2 females. Lau Fau Shan, 13.x.2010, C.H. Shih, 2 males, 2 females (AFCD).
Etymology. The specific name maipo is a Cantonese word considered here a noun in apposition (Mai meaning rice, Po meaning a plain). The Mai Po Inner Deep Bay (1540 hectares of wetlands) was designated as a wetland of international importance under the Ramsar Convention in 1995. 2010 is the 15th anniversary of the designation.
Diagnosis. One of 13 species of Pteroptyx from the Oriental region having a metafemoral comb, aedeagal sheath with paraprocts, aedeagal lateral lobes shorter than median lobe and separated for less than half their length dorsally (Table 1); sharing a pale dorsal colouration (Figs 1–4) with black elytral apices with 10 species; most similar to Pteroptyx valida, and P. sulawesiensis, distinguished from both by shortened deflexed elytral apices, and contiguous LO in V7; distinguished from P. asymmetria, P. decolor and P. similis in lacking elongate lobes arising from median posterior margin beside median emargination of tergite 8; differing from P. t r un c a ta, which has shortened elytral apices, LO halves in V7 separate and apices of PLP of V7 produced and obliquely truncate, by LO halves contiguous in mid line and PLP rounded, scarcely produced. Females macropterous, bursa with elongate plates on upper and lower surfaces on left side. Species produces spermatophores. Larvae (Figs 5, 6) of form of Pteroptyx valida (Ballantyne & Menayah 2002), elongate slender, lacking laterally explanate tergal margins; differing from P. v a l i d a by pale abdominal terga 8, 9.
Description. Male. 6.8–7.5 mm long; 2.6–2.9 mm wide; W/L 0.37–0.4.
Colour (Figs 1, 2): Pronotum pale yellow, semitransparent with underlying white fat body contributing to light colour; retraction of fat body in middle of disc and across posterior margin gives misleading impression of darker median markings; fat body at sides outlining edge of flattened hypomeron. Mesoscutellum and mesonotal plates coloured like elytra. Elytra semitransparent, very light brownish-yellow; fat body scattered evenly beneath cuticle; in ethanol preserved specimens underlying hind wing often causing wings to appear darker; dark apical portion narrow, deflexed apex dark brown. Head, antennae and palpi dark brown. Thorax ventral surface yellow; legs faint brownish yellow, with brown posterior face of tibiae of legs 1, brown apices of tibiae of legs 2, and all tarsi brown. Abdomen with basal ventrites yellow, ventral abdomen more brightly yellow than thorax; posterior margin of V5 narrowly white (apparently fat body only as this area does not show any illumination); white LO occupying all of V6; LO in V7 white, apparently bipartite with halves contiguous in median line and barely separated in anterior midline, separation appearing as small triangular slightly darker area; luminous LO material extending into PLP but not to apices; LO retracted from posterior margin in area of emarginations separating PLP and MPP; LO barely extending into MPP with a further area of non luminous fat body extending about half its length; V7 very narrowly yellow across posterior margin with tips of PLP sometimes brownish; all tergites yellowish and semitransparent; dorsally reflexed lateral margins of ventrites brownish yellow except for white V6 and 7; dorsal surface of margins of posterior 1/3 of PLP light brown.
Head: GHW 1.5–1.7mm; SIW 0.2–0.3mm; ASD (0.11–0.17mm)> ASW (0.09–0.14); vertex quite deeply depressed between eyes; no demarcation between median frontal area above antennal sockets and vertex; apical labial palpomere ovate, all margins entire, and 1/4 as long as ovoid apical maxillary palpomere. Antennal length (Figs 1, 2) approximately 2 x GHW, all segments simple, elongate, no FS dilated; FS decreasing very slightly in length and width from FS1 (longest and widest) to apically rounded FS 9.
Pronotum (Fig. 1): 1.9–2.2 mm wide; 1.3–1.4 mm long; W/L 1.5–1.6; pronotal width less than humeral width; median anterior margin broadly rounded, projecting slightly beyond rounded obtuse anterolateral corners; lateral margins convex sided (pronotum widest across middle); posterolateral corners narrowly rounded, slightly exceeding 90°; posterior margin largely straight, median posterior area projecting slightly beyond corners, not separated from corners by emarginations; dorsal surface fairly smooth, not greatly elevated, lateral areas flat (outline partly defined by retraction of fat body), median sulcus depressed slightly along its length, more widely in median area of disc; narrow depressed area running across posterior margin and covering 2/3 width of pronotum in this area; narrow hypomeron closely adpressed at sides, more widely so in middle.
Elytra (Fig. 1): 5.5–6.2 mm long; lateral margins subparallel-sided for approximately 1/2 their length, beginning just behind humerus, and converging posteriorly in posterior 3/8; interstitial lines not well defined, line 3 at base (near humerus) sometimes visible; sutural margins in closed elytra contiguous for most of length; deflexed apex rounded (Fig. 2), A longer than B, both longer than C.
Legs (Fig. 2): all tibiae straight; no femora swollen or curved; no basitarsi of any legs with excavated inner margins; MFC present (Figs 49, 50), with 5 strong apically curved, and 4 weaker setae (the functional extent of the MFC not reliably established, is unlikely to include any more of the weaker hairs along this part of the femoral apex).
Abdomen (Figs 33–42) with posterior margin of narrow V3 recurved, anterior margin of V4 extending into it; V3 about 1/3 median length of V4; LO occupying all of V6, bipartite in V7, closely approximate in median line with small median anterior triangular area devoid of LO; light production (Fig. 35) indicates LO material extending into PLP and slightly into MPP; from below MPP slightly longer than wide, from side slightly deeper than wide; truncate along its ventral edge; much deeper at apex than at base with lateral margins developed as pointed projections at their posterior end and sloping ventrally towards the anterior end; posterior face largely flat, from behind along longitudinal axis ovoid in outline with dorsal surface broadly emarginated; ventral area of emargination with very short paired rounded projections, visibility depending on orientation of abdomen; dorsal surface of MPP with median longitudinal concavity; separated by shallow emarginations from narrower PLP which do not project posteriorly as far as the MPP.
Tergite 8 (Figs 36, 37) lacking any projections at sides of median posterior emargination; posterior margin with corners rounded, slightly produced, and margins between corners and median emargination slightly sinuate; ventral surface with median longitudinal trough housing sheath and aedeagus margined by well defined raised areas terminating anteriorly in short, narrow, apically pointed flanges; anterior paired prolongations of tergite 8 rounded, subequal in length to posterior half tergite 8, wide in horizontal plane, lacking lateral projections and projecting anteriorly beneath tergite 7.
Aedeagal sheath (Figs 46–48) like the Oriental Pteroptyx (Ballantyne & Lambkin 2009 Figs 86–89) with bulbous lateral paraprocts (extensions of sheath tergite) enveloping sides of sheath sternite just behind its widest point; sheath sternite posterior to tergite articulations with lateral margins sloping obliquely and partially enclosing aedeagus at sides; muscle attachment onto these pieces may cause deformation if aedeagus is removed. Aedeagus (Figs 43–45) with narrow apices of lateral lobes) not visible at sides of median lobe; lateral lobes shorter than median lobe and separated apically for 2/5 their median dorsal length (measured from anterior dorsal margin). Paired spiracles and tracheal trunks lying at sides of sheath and beneath tergite 8 assumed to belong to segment 8.
Female (Figs 3, 4). Macropterous with flight observed. 7.5–8.2 mm long, 3.0–3.1mm wide; W/L 0.37–0.4. In common with other Pteroptyx females lacking MFC and deflexed elytral apex, and with head and eyes smaller than those of male. Coloured as for male except for white LO restricted to V6 only, V7, 8 orange yellow; all tergites orange yellow; lateral margins of V7 and V8 tapering posteriorly, with V8 about half as wide as V7, slightly shorter and medially narrowly emarginated along its posterior margin (Figs 51, 52); anterior margin of V8 very narrowly prolonged beneath V7, with concealed anterior 1\4 of V8 pale semitransparent (Fig. 54); posterior margin of V7 shallowly, broadly emarginated, slightly trisinuate (Figs 51, 52).
Reproductive system (Figs 53–60): Calder (1996) outlined the numerous difficulties in homologising the structures of the female genital tract. Interpretations here are based on Wing et al. (1983), Yahiro (1996:539) and Zacharuk (1958). In particular, we adopted Tandon’s (1970:81) definition of the extent of the bursa as—“the vagina…extends anteriorly to form a broad blind cylindrical tube the bursa copulatrix …..the junction of the bursa copulatrix with the vagina is marked by the opening of the median oviduct.” In Pteroptyx maipo sp. nov., an elongate cylindrical vagina leads anteriorly from the vulva, just anterior to the styli, and enlarges anterior to the entry of the common oviduct to form an elongate, blind ending muscular bursa copulatrix which contains paired plates. A spermathecal digesting gland (which may not be expanded and is often difficult to see unless immersed in water) opens from the anterior end of the bursa. Remnants of one elongate spermatophore were found in the most gravid female, partly in the digesting gland and partly within the bursa, with the anterior apices of the lateral bursa plates closely pressed to its sides (Figs 58, 60). We assume that the digesting gland was closely pressed around the spermatophore because its outline was not visible. A small stalked spermatheca arises from the side of the bursa (Figs 57, 58). The median oviduct branches into two lateral oviducts which branch into numerous well filled ovarioles occupying most of abdomen (Fig. 55). A small plate defines the entry point of the median oviduct (Fig. 56). The bursa is muscular, deep grey in freshly killed specimens and dorso-ventrally flattened with position of the anterior margin varying in the four females dissected depending on the gravidity of the abdomen, the most gravid abdomen having the bursa close to the posterior end of the abdomen; bursa with elongate plates on upper and lower surfaces on left side; plates elongate, curved, with pointed posterior ends (resembling fish hooks), bearing very short spines on their inner surfaces; bursa otherwise somewhat C shaped with strong ridged cuticle covering remainder of surface; posterior margin narrowing and bearing a small plate which apparently surrounds the area of the vagina into which the median oviduct discharges; dissected bursa lacked spermatophore in 3 of 4 specimens. Genitalia (Figs 53, 54) with external short styli, scarcely differentiated coxites and elongate valvifers/baculi which attach both to sides of the vagina and inner surface of anterior prolongation of V8, increasing the vaginal width during egg laying; these muscle attachments when dried may lead to irregularities in external cuticle of V 8 in pinned females (Ballantyne observations).
Larva (Figs 61–76). Similar to Pteroptyx valida (Ballantyne & Menayah 2002; Ohba & Sim 1994) and this description highlights differences; body form also like Colophotia praeusta Eschsch. (Ballantyne & Lambkin 2009), and some Australian Luciola (Ballantyne & Lambkin 2000); terrestrial; soft bodied; elongate slender tapering a little in front and behind (widest across abdominal segments 2 – 4); with 3 thoracic and 9 obvious abdominal segments (a 10th segment may be represented by a narrow ring of dark cuticle behind tergum 12); lacking laterally explanate margins on terga; lacking eversible glands and gills along the sides of the body.
Colour (Figs 5, 6, 61, 62, 70): dorsal surface dark brown (mottled) with areas surrounding hair bases paler than rest and irregular pale areas on surface of terga 1–10; tergum 11 pale and 12 with pale areas at the sides; differing most obviously from P. v a l i d a by the pale dorsal surface of abdominal tergum 8, the pale sides of tergum 9 and lacking any pale markings along sides or posterior margins of most terga; dark colour of tergal plates extending to sides of plates (sides of body sometimes appearing pale if laterotergites visible; Fig. 6 larva dead preserved, Figs 61, 62 living larva); pale median line arising from the anterior margin of tergum 1 extending posteriorly to posterior margin of abdominal tergum 8; median line expanding at its posterior end in each thoracic segment and abdominal segments 1–5; widest across abdominal segments 2–4 and narrowing posteriorly in abdominal segments 5–7. Except for largely pale yellow venter of segments 11, 12 ventral surface dingy cream with extensive dark markings (Figs 6, 62).
Head (Figs 71–76): flattened, parallel-sided, prognathous, smooth, largely hairless except for hairs surrounding mouthparts, enclosed at rest within flexible extensible “neck membrane” inside prothorax; with median dorsal frontoclypeus bounded at sides and behind by an epicranial suture, with posterior arm very short; parietals reflexed ventrally; mouthparts occupying most of ventral head area as a compound plate between edges of reflexed parietals, consisting of maxillae at sides and median labium (cardines and stipes on each side fused with posmentum by membranous areas). Homologies of mouthparts follow Ballantyne and Menayah (2002). Apex of antennal segment 2 oblique, surmounted by much shortened segment 3 and adjacent sense cone; flagellar segment 3 elongate, terminated by hairs; sense cone rounded, subequal in length to segment 3 (single apparently anomalous larva with 2 sense cones to left antenna Fig. 75; instar not determined). Sensory organs at apices of apical maxillary and labial palpomeres (indicating a terrestrial mode of life; Fu & Ballantyne in prep.). Mandibles lack retinaculum.
Dorsal surface: all terga except for tergum 1 and 12 wider than long; tergum 12 parallel-sided, longer than wide and terminated by a series of eversible pygopodia; terga slightly biemarginate along posterior margin with rounded nodules at posterolateral corners and to either side of the median line (nodules becoming more pronounced towards end of abdomen; nodules and emarginations arrowed in Fig. 61); irregular elongate depressed areas on most terga; tergum of prothorax longer than wide, widest across posterior margin; very slightly emarginated along anterior margin at origin of midline, lateral margins diverging posteriorly with posterior half slightly wider than anterior half; posterolateral corners rounded, terga of meso and metathorax shorter than tergum of prothorax, wider than long with lateral margins converging slightly in anterior 1/3; abdomen with anterior terga subequal in length, slightly shorter than metathoracic tergum.
Ventral surface: elongate pleural suture running from anterior margin of segment 2 to posterior margin of segment 11 delimits laterotergites bearing spiracles above, and in the abdomen laterosternites below (Fig. 62, 70; see also Ballantyne & Menayah 2002 Fig. 1); laterotergites may be visible at sides of body when viewed from above (Fig. 61); episterna and epimera of segments 1–3 represented by narrow cuticular strips above and to the sides of the coxae of legs 1–3 (as in Ballantyne & Menayah 2002 Fig. 1); ventral surface of meso and metathorax composed of two sternal areas in each segment, an anterior presternum and posterior median sternum bearing legs (Fig. 62); pair of laterotergites at sides corresponding to each sternal element, anterior pair bearing spiracles in mesothorax. Legs: 5 segmented, short, coxae widely separated at their bases, terminated by a single apical claw (tarsungulus) as described for P. v a l i d a (Ballantyne & Menayah 2002). Abdominal segments 1–8 (body segments 3–11; Figs 62, 70) with single spiracle bearing laterotergites at sides, wide median coloured sternal plate separated from laterotergites by elongate slender plates—laterosternites; LO present beneath segment 11; ventral area of terminal segment not differentiated into areas.
Pupa (Figs 65–70, 77–84). Bred in captivity from field collected larvae. Before pupation, larvae provided with soil burrowed into the soil, and made a cell (10 mm in diameter), the inside surface of which was smoothed by the larva, but no coating was found (soil was not necessary for pupation). The prepupa assumed a curled shape and produced light continuously (without flashing) from the 8th abdominal segment (source bipartite) (Figs 69, 70). Duration of prepupa: 6–8 days. Duration of pupal period:13 days. Length of male pupae: 6.5 mm–8.5 mm (measured from abdominal apex to frons, curled posture, cerci not included).
Male pupa (Figs 65–68, 77–79). Exarate; glabrous, body slightly curled with flexible abdomen; legs not movable.
Colour: whole body light yellow in 1-day old pupa, except for light brown crescentic area at the side of each compound eye, formed by the pigmented, developing ommatidia; dark colouration extending to entire compound eyes after 2 days, continuing to darken subsequently, becoming almost completely black by day 8. Frons, antennae, mandibles, maxillary palpi, labial palpi, hind wing pads, all tarsi started to darken in day 10; remainder of body whitish yellow remaining unchanged during development.
Head: large, well defined, not concealed within prothorax; eyes large; antennae subequal in length to GHW and extending posteriorly to sides of pronotum; mouthparts well defined with apical palpomeres visible in front of head.
Thorax: with large pronotum not covering head from above; pronotum with median line (Cicero 2008 terms this the DML—dorsomedian longitudinal line); mesonotum reduced, wider than long, with short wide mesoscutellum; metanotum large, about as long as wide; wing pads well defined, apex of elytral pads not deflexed; curvature of pupa allows wing pads to reach as far as 4th last ventrite; both sets of wing pads clearly visible from beneath; elytra bearing a depressed area on the outer preapical margin (Figs 77, 78); femora of legs 1, 2 strongly projecting sideways; femora of legs 3 largely concealed behind metathoracic wing pads; tarsi extending well into ventrite 6.
Abdomen: abdominal segments wider than long; with 8 visible tergites, posterolateral corners of tergite 1 weakly prolonged, of tergites 2–7 strongly prolonged; corners of tergite 8 not prolonged and posterior margin essentially trisinuate; ventrites 1–7 all visible (V1 largely covered by hind wing pads, visible from the side; V7 is the terminal light organ segment; no recognisable sign of ventrite 8, and based on visual examination only, the structures behind the posterior margin of V7 (Fig. 79) are attributed to the aedeagal sheath and aedeagus; membranous pleural line running between dorsal tergites and ventral plates; spiracles at sides of ventrites just below pleural line (Fig. 68); elongate segmented cerci arising from the sides of segment 9 (Fig. 79).
Light formation (Fig. 66): As soon as pupation occurred any slight stimulus caused light to be continuously (without flashing) emitted from the area above V7 and near the V7 spiracles (light passing through V7 giving the misleading impression the source is immediately beneath the V7 cuticle; light organ source at this stage bipartite but areas appearing to overlap; from day 4 light emitted from second last abdominal ventrite (V6) in two lateral patches; no light observed from terminal segment from that stage on).
Female pupa (Figs 80–84) like male except: head and compound eyes smaller, abdomen with 8 ventrites, light organ beneath V6 visible; no depressed area on outer preapical margin of elytra; posterolateral corners of tergite 1– 7 less produced, especially those of tergite 7; all abdominal ventrites visible; terminal ventrites not trisinuate across posterior margins.
Eclosion. Deflexion of elytral apices began after the adult male (pharate adult) emerged from the pupal case. Full colouration was achieved in 12 hours. Glowing light was produced only from V 6 in the first few hours. Twelve hours after emergence two restricted light spots in V7 were observed; these gradually enlarged reaching full size with the two halves touching in the median line at 36 hours after emergence, when both V6 and V7 light organs commenced flashing. Before that, only a continuous glow was observed.
Remarks. The description of this new species of firefly contains an unusual wealth of features including morphological and ecological characters of all life cycle stages. Additionally association of males, females and larvae has been established by rearing.
Biology and Ecology. Habitat (Figs 94–96). The new species is probably mangrove dependent as it was recorded in mangrove habitats, including the Hong Kong Wetland Park and Mai Po Nature Reserve, in Hong Kong. The sites are located at intertidal rivers which are affected by daily tidal changes, and have brackish muddy soil and dense mangrove vegetation on the banks. The habitats are dominated by mangrove plant species Acanthus ilicifolius, Aegiceras corniculatum and Kandelia obovata.
The Mai Po Nature Reserve is part of the Mai Po Inner Deep Bay Ramsar Site, a wetland of international importance, while close to it is the Hong Kong Wetland Park. These wetlands are situated in an estuarine bay under the tidal influence from Inner Deep Bay in northwestern Hong Kong. The average water depth of the area is about 2.9 m and the mean tidal range is 1.4 m. The Inner Deep Bay receives water and sediment from surrounding rivers that carry clay and silt to form the core part of the intertidal mudflat. Hong Kong is a sub-tropical region under strong influence of monsoons. Rainfall occurs mainly from April to September. Salinity of the intertidal water varies (3.5–6.6 ppt) depending on season with lower salinity common during summer.
Adults were found in rare cases up to 50 m from the mangrove. They did not seem to exhibit any preference for different plant species. Larvae were found in wet soil that would be flooded by sea water at high tide. It is highly unlikely that the larvae could survive the periodic immersion and probable that they climb up the mangroves or move to areas above high tide mark. Further work is necessary to determine preferred habitat.
Seasonality. The seasonality of the new species is yet to be ascertained, but it appears that the adult stage of this firefly persists throughout summer. Adults of the new species have been recorded on the following dates, and the number of individuals observed peaked in May and August.
Of the eight larvae collected in August 2010 and reared in captivity, five larvae pupated in late October to early November and emerged one after another from mid-November.
Behaviour. Shortly after dusk flashing males were observed flying up from the vegetation (Table 2). On 16th and 17th August 2010, flashing began at 19:23hrs and 19:25hrs respectively with numbers of flashing males increasing later in the evening. Flashing males usually flew not higher than 1.5m above vegetation with single short flashes of 0.90±0.21 sec (mean±SD) duration (Fig. 93), and longer interflash intervals of 4.95±0.28 sec (mean±SD). The females usually perched on leaves and stems of the vegetation and have quick flashes with 0.17±0.06 sec (mean±SD) duration and interflash intervals of 0.1±0.06 sec (mean±SD). Therefore this species differs from the well known fireflies of Malaysia (P.t e ne r and P. malaccae) which congregate (separately) along mangrove lined tidal rivers and males flash in synchrony. P valida in Malaysia does not synchronise but males are often observed in the same trees as populations of P. malaccae. Observations on about 50–60 flying or stationary individuals of P. m a i p o sp. nov. randomly checked at different times and at different locations revealed all were male. The only females observed were mating. Around 3 hours after sunset most males had settled on the vegetation and many stopped flashing.
Date Location Lat. Long. Elevation Temp R.H Sunset 1st Sight of 1st Sight of time Flashing Flight
16/8/2010 Hong Kong Wet- 22° 28N 114° 00E 12m 28 88% 18:55 19:23 19:27
land Park
17/8/2010 Mai Po Nature 22° 29N 114° 02E 13m 25 95% 18:56 19:25 19:30
Reserve Area
Mating, Copulation Clamp and Egg Laying (Figs 85–88). Initially males mounted the female facing in the same direction holding the edges of the female elytra primarily with the tarsal claws, and the tip of the abdomen was inserted from one side of the female. No function for the MFC in the mating process was observed. Once the mating pair turned to the tail to tail position, the male elytra of one pair were positioned under those of the female with the deflexed apex pressing on the female abdomen in the area of tergite 5–6 (and thus probably anterior to the muscular bursa). In the three mating pairs Yiu observed in the field, one had the male's terminal abdominal segments extended posteriorly such that the elytral tips did not reach the posterior end of the female; the second had the elytral tips placed above the female's elytra (Figs 85–92). Both these females produced fertilised eggs. The third pair, demonstrating a copulation clamp stood on a v-shaped platform where they could easily push against each other (Fig. 85). This pair was sectioned (Figs 91, 92).
Pairs may remain coupled for up to three hours. During a 44 minute observation of a mating pair by Yiu in the field, the female emitted light throughout the period; the male either emitted no light or luminesced only from the light organ in V6.
Females can lay at least two batches of eggs (in one instance seven days after mating) with up to 106 eggs in each batch. On average, the larvae emerge after 17 to 34 days.
description: Published as part of Ballantyne, Lesley, Fu, Xin Hua, Shih, Chun-Hat, Cheng, Chui-Yu & Yiu, Vor, 2011, Pteroptyx maipo Ballantyne, a new species of bent-winged firefly (Coleoptera: Lampyridae) from Hong Kong, and its relevance to firefly biology and conservation, pp. 8-34 in Zootaxa 2931 on pages 13-31, DOI: 10.5281/zenodo.278036