taxonID	type	description	language	source
03AF8788F721FFAE91C0BA3F903477D9.taxon	distribution	Distribution — Borneo, Philippine Is., Java, Celebes, Molucca Is., New Guinea, Solomon Is.; Australia (?). Habitat & Ecology — In forest, 0 – 500 (– 1650?) m altitude. Notes — Recognized especially by its chartaceous, symmetrically ovate leaves, and short, stout, fully concrescent fruits. The leaves are generally glabrous but there may be a few slender, pale, patent hairs on the stipule, petiole and nerves below. A denser bristly to subvillous indument is seen in a number of specimens from West New Guinea, e. g. BW 7367, Vogelkop Peninsula. The species seems to be uncommon in the Bismarck Archipelago. It is absent from the Highlands Region, and I am sure of only one New Guinea collection made from above c. 500 m: LAE 58792, New Britain, Mengen Massif, 900 m. (Another collection, NGF 8485 (Morobe District, Skindewai, c. 1650 m), may be P. abbreviatum – its inflorescences are too young to be informative – but its 9 4.5 cm leaves seem too large for it to be credible that the stated altitude is correct). The New Guinea plants are not nearly as conspicuously red-glandular as those from the Philippines. Borneo is included in the species’ range solely because of the citations of Quisumbing (1930: 62) and Beaman & Anderson (2004: 253). A recently described species P. fungiforme (Spokes 2007: 234, 457), endemic to northern Queensland, resembles P. abbreviatum in some respects, notably in its short, fully concrescent fruits (as illustrated by Cooper 1994: 223, under the tag-name Piper sp. ‘ Leo Creek’). It is puzzling though that males are said to have their anther locules separated by a swollen connective, just as in P. celtidiforme. An re-examination of P. fungiforme needs to be undertaken to resolve whether it might represent a mixture.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F721FFA991C0BF4C92D876D1.taxon	description	Piper arfakianum C. DC. (1917) 127; Chew (2003) 15. — Type: Gibbs 5525 (iso K, L n. v.), Arfak Mts, Angi Lake, [c. 1800 m]. Piper pilosulinodum C. DC. (1917) 128; Chew (2003) 15. — Type: Gibbs 5624 (holo BM; iso K n. v.), Arfak Mts, Koebre Ridge. Fertile-shoot internodes c. 2 mm diam, nearly smooth. Vegetative parts (at least, stipule, newest internodes and petiole) with patent pale to mid-brown bristly-flexuose hairs to c. 1 mm long. Stipule to c. 1 cm long. Leaf blade chartaceous to subcoriaceous, ovate (or elliptic-oblong), 4 – 7 (– 10) by 1.5 – 4.5 cm; base symmetrical, usually shortly cordate to rounded, subequal at petiole and usually shortly incurved there, apex long-acuminate; main lateral nerves 1 – 2 pairs, basal, narrowly prominent above; surfaces of blade sometimes red-glandular. Petiole 0.5 – 1.5 cm long, usually c. 1 / 6 as long as blade. Male inflorescence not seen. Infructescence a spike c. 4 – 8 cm long, c. 0.5 cm diam, on a peduncle c. 1 – 2 cm long; rachis sparsely hirsute, bracts subsessile, glabrous to villous, bract-heads 0.5 – 1.3 mm diam, held at or just below apex of fruitlets. Fruitlets free, 1.2 – 2 mm diam, broadly rounded to flattened above but usually beaked by a stout style c. 0.25 mm long; stigmas 3, broadly oblong, together c. 0.35 – 0.5 mm diam. Distribution — New Guinea: Arfak Mts to Milne Bay Province. Habitat & Ecology — Small climber or scrambler in montane moss forest, ridge thickets, and Nothofagus - dominated forest; c. 1900 – 2900 m altitude. Notes — This species seems to be an uncommon one. I accept twelve of the thirteen extra-typical collections listed by Chew (2003: 16). They are mainly from Papua New Guinea, and I cannot add any new localities there or for the island as a whole. With respect to the thirteenth listing, Schodde 1556, I place this in P. macropiper (Fig. 6 j). Recognized by its small to medium-sized, ovate-triangular, long-acuminate leaves, which are usually coarsely bristly on the petiole and venation below. The major nerves all depart the midrib from 0 – 5 mm above the blade base, and the blade’s basal margins are usually shortly incurved before the midrib channel is reached. These features help distinguish the species from P. subcanirameum (shortly acuminate apex, glabrous, nerves departing from up to 1.5 cm from petiole base, blade margins decurrent down into sides of petiole). Both species have rather coriaceous and glossy leaves but there is a textural difference: in P. arfakianum the lesser venation is seldom as prominent above as it is in P. subcanirameum. Possibly the leaves of the former are, in life, relatively fleshy, but label-notes are inadequate on this point. Material from the western part of the island has consistently smaller leaf blades (to c. 6 2.5 cm) than those from Mt Kaindi and further eastwards. Kostermans 2382, from the type locality, differs in its leaves being appressed-villous above and below. Chew (2003) saw isotypes of P. pilosulinodum C. DC. and found them not to differ significantly from the present species. Having seen Van Royen & Sleumer 7456 (Tamrau Mts, Mt Nettoti, CANB), said by Chew (2003: 15) to be a good match for the P. pilosulinodum types, I agree.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F726FFA9930ABFBA97A77225.taxon	description	Piper betle L. (1753) 28; Quisumb. (1930) 85; R. O. Gardner (2006) 579; (2010) 8. — Type: Herb. Hermann 3: 32, 4: 9 (BM n. v.). Distribution — Widely cultivated in Asia, Malesia, Micronesia, and in Melanesia east to Fiji. Habitat & Ecology — In gardens and persisting after cultivation, also naturalized; to c. 750 m altitude. Notes — If the distinctive fruits are not present this species might be confused with P. caninum or P. interruptum, but its leaves tend to dry brownish rather than grey- or olive-brown and do not have large silvery glands. It almost always lacks long hairs, but near the base below (sometimes across both surfaces, best seen on new leaves) there is a scattering of minute (0.05 mm long) appressed hairs. Labels for New Guinea specimens of P. betle are generally unclear as to naturalization status. However, for the Bismarck Archipelago Peekel (1984: 124) says: “ Everywhere wild in scrub ... “. Quisumbing (1930: pl. 42 (8 )) described the stamen of P. betle as having a short stout filament c. 0.6 – 1 mm diam, with the two rather large anther locules colinear at the filament apex. Dehiscence takes place at or just below the bract-heads, and is by a continuous apical slit. As in other regions, female spikes can also bear stamens, e. g. in Darbyshire 1018.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F726FFA991C0BB1E90BC7733.taxon	distribution	Distribution — New Guinea, Solomon Is. Habitat & Ecology — In forest, to 400 (– 750) m altitude. Notes — Recognized by its symmetrical, subcoriaceous, greyish, glabrous leaves that may be conspicuously silverglandular (as in P. caninum and P. interruptum), and especially, by the very short fruiting spike and large ovoid fruitlets. Of male material from New Guinea I have seen only Docters van Leeuwen 11202 and Kanehira & Hatusima 12103. They contrast with three male Solomon Is. collections in having relatively large stamens, and in these being paired rather than solitary (Gardner 2010: 8). Also, the leaf and the female spike tend to be larger in the Solomons, and possibly the fruitlets are larger there too, being, according to Chew (2003: 17), “ c. 10 6 mm ”. The basal leaf nervation, solitary anther (sometimes) and short female spike with large free fruitlets, give this plant a resemblance to P. quinqueangulatum Miq. (P. korthalsii Miq.) of Borneo, Java and the Philippine Is. The latter, however, tends to dry more intensely orange-brown and its stems are more strongly ridged.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F726FFA491C0BE20912B7284.taxon	distribution	Distribution — Western Malesia to the Solomon Is. and northeastern Australia. Habitat & Ecology — In forest, 0 – 500 (– 1600) m altitude. Notes — Recognized by its chartaceous, greyish to olive-brown leaves which usually have a light cover of short pale bristly hairs. Rarely, the indument is subvillous (Streimann 8368, Morobe Province, near Lae, 250 m). Because of the similarity of leaf shape, texture, glandularity and nervation in P. caninum and P. interruptum, and because of the variability of the latter’s indument, sterile specimens can be hard to place as one rather than the other of these two species. The two highest-altitude specimens I have seen are both from 1600 m: Brass 24814, Milne Bay Province, Goodenough I.; NGF 37258, Western Highlands, Kopiango. Van Royen (1982: 1269, f. 403) incorrectly described P. caninum as reaching “ the upper subalpine shrubberies [at] 3290 m ”. This altitude, and his f. 403, are based just on Vink 17365, which belongs to P. rodatzii of the P. macropiper species-complex. Johns et al. (2006: 401) cite a Kloss specimen collected from “ Camp IX – XIII ” [c. 1700 – 3150 m] on Mt Jaya (Carstenz). I have not been able to check this specimen at BM, and regard even 1700 m as an anomalously high altitude. The leaf blades of Hoogland & Craven 10805 (East Sepik Province, Hunstein River, c. 150 m) measure only 7 by 2 cm. With this exception, and even at altitudes above 500 m, narrow-leaved plants seem not to occur in New Guinea. This contrasts greatly with the situation in the Philippines (Gardner 2006). Three collections have leaves that are unusually coriaceous and glossy: Brass 24814 (cited above); Clemens 4520, Morobe Province, Ogeramnang; NGF 20253, Central Province, Woitape. They are not otherwise unusual. Takeuchi 14166 (Morobe District, Guam River, c. 80 m, A) has a 30 cm long infructescence (in typical P. caninum, 8 (– 12) cm) and elongate bracts, as in P. interrruptum. But the densely hairy leaves suggest P. caninum, as do the hairy rachis and stipitate fruitlets. I therefore leave this collection undetermined.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72BFFA4930ABBF193E77443.taxon	distribution	Distribution — Philippine Is., New Guinea including Bismarck Archipelago, Solomon Is.; probably also Celebes. Habitat & Ecology — In forest, 0 – 2000 (– c. 2700) m altitude. Notes — Recognized by its chartaceous, ovate-oblong, nearly glabrous (never coarsely hairy), pinnate-veined leaves. The usual red subepidermal mottling is often sufficient to give the whole leaf a brownish red cast. Male P. celtidiforme has uniquely-formed stamens (see ‘ Spot Characters’; Gardner 2006: f. 5). The female flowers are very distinctive too, their narrow, smooth-surfaced stigmas being seen elsewhere only in P. versteegii. These fragile structures are mostly lost as the fruit ripens and the fruitlet apex then just shows a ring of eroded tissue. Also, female specimens sometimes have a granulose papillosity around the base of the stigmas, making a whitish band conspicuous enough to be visible to the naked eye (Fig. 4 a). Less frequently, the central third or so of a female’s bract-heads also bears a patch of granulosity. The specimen NGF 34383 (West New Britain, Fulleborn Harbour, 50 m, A) is unusual in having 4 – 6 very short, broadly ovate, strongly papillose stigmas. Also, the stigmas of Takeuchi 15243 (Morobe Province, Tabare (Tabili) River, sea level, A) have the usual narrow character but are also notably papillose. Neither though seems otherwise unusual for P. celtidiforme. The occurrence on Celebes seems likely, since de Candolle (1923: 278) synonymized one of his names from there, P. sinkojan C. DC. (as ‘ P. sinkgian ’) with P. corylistachyon C. DC. of the Philippines, and the latter has, correctly I believe, been placed by Quisumbing (1930: 177) under P. celtidiforme. De Candolle never saw the PR holotype of P. celtidiforme – as the annotation-free photograph of Quisumbing (1930, pl. 22) proves – and in his 1923 account he treated the species just as a ‘ Eupipera non satis nota ’. For the Bismarck Archipelago Peekel (1984: 129) says, under P. singkojang [sic]: “ Easily the most common species of pepper in the region; widespread on the trunks of forest trees. The red fruit-spikes, erect from horizontal twigs on the tree-trunks, are particularly conspicuous. Where the plants find little support, the stems and branches spread widely over the ground and the densely-set leaves form there a regular turf. The leaves of such plants are usually smaller and their bases deeply cordate or reniform ”.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72BFFA491C0BA3F9612706E.taxon	distribution	Distribution — Celebes, Molucca Is., Philippine Is., New Guinea (western part of island, also Sepik and Madang regions); possibly also Micronesia. Habitat & Ecology — In forest, to c. 500 m altitude. Notes — This species is apparently uncommon in New Guinea, just as it is in the Philippines, where Quisumbing (1930: 42) saw only four collections, all from ‘ medium altitude’. The New Guinea specimens I know of – those I have seen myself, and others cited by Chew (1972: 7) – are from the northern side of the island, from the Vogelkop east to Madang Province (NGF 24752, near Aiome). I have seen only a few specimens of the Micronesian (Palau Is.) taxon P. hosokawae Fosberg (P. decumanum var. palauense Hosok.), none of which have mature infructescences. Their leaves, e. g. as in Takamatsu 1784 (B) are relatively small (c. 18 by 10 cm) compared to those typical of Malesian P. decumanum but are not otherwise unusual. Fosberg & Sachet (1975) do not mention anything distinctive about the female parts of P. hosokawae. The stomata of P. decumanum leaf are borne singly on low pustules scattered across the blade underside (Fig. 4 b).	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72BFFA591C0B9DD93B473E8.taxon	distribution	Distribution — Molucca Is., Philippine Is., New Guinea (northern coast, also Milne Bay and Bismarck Archipelago), Solomon Is., Vanuatu, Micronesia. Habitat & Ecology — Coastal forest and scrub, mainly on limestone and coral sands, perhaps only to c. 100 m altitude. Notes — De Candolle (1869: 336) based his P. barclayanum on a BM collection from the H. M. S. ‘ Sulphur’ voyage: Barclay 4021, island of Japen (‘ In ins. Tobia’), West New Guinea. He placed it at that time in his sect. Pothomorphe, which included taxa like P. peltatum L. and Macropiper spp. Subsequently, in his Candollea treatment (1923: 171), he placed it in sect. Macropiper. Smith (1975: 35) was unable to locate a type for P. barclayanum and commented that there was nothing in its description to suggest it was a Macropiper. He might have added that a New Guinea occurrence would be a notable regional range extension westwards from the Santa Cruz Is. A sheet of Barclay 4021 (BM 000624274, image!) has been found in the present study, but bears no De Candollean annotation. It represents P. fragile. Piper fragile might be confused with P. abbreviatum, but its leaves are glabrous and thicker and tend to dry olive yellowish rather than dark grey. Also, its leaves (mainly just the climbingshoot ones) are subpeltate to peltate. In New Guinea this species seems to occur just along the northern coasts and on the islands there. Fosberg & Sachet (1975) say it is common in Micronesian lowland forests. Peekel! Fig. 4 Piper species, various features. — a. P. celtidiforme Opiz. Infructescence, showing ring of white papillae around base of stigmas. – b. P. decumanum L. Leaf underside, showing pustulate stomata. – c. P. macropiper Pennant s. str. Specimen from Morobe Province, leaf underside, showing dendritic hairs. – d. P. macropiper var. macrophylla R. O. Gardner. Leaf underside, showing indument of nerves near blade base. – e. P. mestonii F. M. Bailey. Infructescence, longitudinal section, showing coriaceous surface of the fused ovaries, long styles and ‘ 2 - lipped’ stigmas. – f. P. versteegii C. DC. Infructescence, showing the long styles and very narrow stigmas. – g. P. subcanirameum C. DC. Infructescence (rehydrated), showing the mostly-free fruitlets. – h. P. cf. amboinense C. DC. Leaf base below, showing ant-sac (a: NGF 32677; b: Darbyshire & Hoogland 8124; c: Takeuchi 4572; d: Takeuchi 8675; e: NGF 40944; f: Brass 7000; g: Takeuchi 5756, B; h: Kanehira & Hatusima 11500; all from A except as noted). — Scale bars = 1 mm. (1984: 129) describes it in the Bismarck Archipelago: “ Frequent; climbing on tree-trunks and on coral rocks behind beaches. The natives gaily crown themselves with the decorative, often mottled, leaves ”.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72AFFA5930ABB65909073AC.taxon	description	Piper philippinum sensu Quisumb. (1930), non Miq. (1843 – 1844) 322. Distribution — Taiwan, Philippine Is., New Guinea, Solomon Is., New Caledonia, Fiji and Samoa. Habitat & Ecology — In forest, near the sea or some way inland but at low altitude. Notes — For P. insectifugum in New Guinea De Candolle (1925: 219) cited Riggenbach 16 [Mamberamo region] and Moskowski 16 [no locality given]. Chew (2003), under P. austrocaledonicum, cited two collections: Peekel 322 as in the synonymy above, and Schlechter 14381, ‘ Torricelli’, BO. I have only seen Peekel 322, and can confirm its identity. I have also seen some of the material maintained by Chew (2003) as P. melula and do not think this taxon differs from P. insectifugum in any important way (compare Fig. 3 c with Gardner 2010: f. 6 a, b). Chew (2003: 20) in the notes following his full description suggested that P. melula appeared “ to be related to P. austrocaledonicum ”.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72AFFA591C0BA9C961A71D9.taxon	distribution	Distribution — Taiwan, Philippine Is., New Guinea, Solomon Is., and possibly Vanuatu (Gardner 2006: 582); also Australia. Habitat & Ecology — In forest, to c. 700 (– 1300) m altitude. Notes — Two specimens have the anomalously high altitude of c. 1300 m: Carr 13458, Central Province, Boridi; NGF 29214, Eastern Highlands Province, Kassam Pass. The leaves of New Guinean and Solomons specimens have a palmate-pinnate nervation, in contrast to the usual strictly palmate (basally nerved) condition in plants from the Philippines (Gardner 2006). The taxon described under this name by Van Royen (1982) is P. bolanicum Schltr. ex R. O. Gardner, a montane shrub.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72AFFA091C0B94292E370BC.taxon	distribution	Distribution — Philippine Is., Celebes (f. Chew 2003: 19), New Guinea, Solomon Is. Habitat & Ecology — In forest, perhaps from near sea level to c. 820 m altitude at least. Notes — Chew (2003: 18) pointed out that the authority for P. lessertianum should be solely that of C. de Candolle, who published the name without referring in any way to Chavica lessertiana Miq. In my treatment of this species in the Philippines (Gardner 2006) I neglected to comment on the altitudinal variation there: lower-altitude specimens, and the type collection Cuming 1343 (no altitude stated in its protologue) have relatively large but narrow, deeply cordate-auriculate, shortly petiolate leaves, and the peduncles of their inflorescences (male or female) are much longer than the spikes themselves. Nor did I mention that Quisumbing (1930) placed at least some of the smaller-leaved, higher-altitude plants under other names, e. g., P. delicatum C. DC. and P. halconense C. DC. Nevertheless, I believe that the uniformity of the fruitlets of all this material (free, sometimes with a short tapering stylar beak, c. 2 × 1.2 – 1.8 mm; stigmas 3 (– 4), together c. 0.5 mm diam) makes it reasonable to suppose that only one taxon is represented. Also, Chew (2003: 19) noted the extreme variability in leaf form, from elongate to broadly cordate, and pointed out that one collection, BW 8871 (West New Guinea) shows such variation ‘ on the same plant’. I have not myself seen any P. lessertianum specimen from New Guinea that closely resembles the Philippines higher-altitude form. With respect to lower-altitude (elongate-leaved) plants, I have seen two such specimens, both cited by Chew (2003): NGF 28829 (Milne Bay District, nr Mayu I., 350 m) and Pullen 5924 (Northern [Oro] Province, Sibium Range, c. 820 m). I therefore think it reasonable to accept the three other of his citations as belonging to this species: BW 8871, Wissel Lakes; Ijiri & Niimura 674, Mapon District, Wati and NGF 24403, Morobe Province, Asubazo. I also include on my own cognizance the relevant part of the mixed sheet Kloss s. n., K, Mt Carstenz [Jaya], ‘ Camp I 700 feet, Nov. – Dec. 1913 ’. Chew (2003: 18) also synonymized three New Guinea-typified taxa with P. lessertianum. They are: Biro 32, B, “ near Malanaku ”, type of P. biroi K. Schum. & Lauterb.; Roemer 962, L n. v., Mt Hellwig, type of P. lineatipilum C. DC; Brass 1370, A, BRI n. v., Gulf District, Mowabula, type of P. viridibaccum Trel. I cannot confirm this synonymy, because of the complication outlined below. Chew (2003) accepted as a species “ closely similar ” to P. lessertianum a larger-leaved plant, P. pseudamboinense C. DC. In addition to the type, which is from the Ramu River (see Incertae Sedis for locality, etc), he cited two collections, also from lowland forest in Morobe Province: NGF 25678, Buso River and Hartley 10564, Oomsis Creek. I have seen several similar collections from this part of New Guinea (LAE 70724, Takeuchi 5629, 5669, 14024) and one from c. 100 km away (Platts-Mills 51, Madang Province, Adelbert Range, 650 m). As well as cohering geographically these plants present a uniform appearance in their large (c. 25 by 13 cm), chartaceous, palmate-pinnately nerved leaves. The three collections with mature fruit (Takeuchi 5629, 5669; Platts-Mills 51) have long-pedunculate female spikes composed of smallish, free, ovoid, shortly beaked fruitlets (Fig. 8 a). A major point of difference with P. lessertianum is, as Chew (2003) has noted, the fruitlets’ two-lipped stigmas. However, the differences as outlined above are not applicable to male collections, nor to sterile ones with leaves of intermediate size, e. g., Takeuchi 14455, Bulili Ridge, Morobe Province, 215 m and Takeuchi 14828, Arawiri River, Morobe Province, nr sea level. Also, I have seen one Morobe Province specimen of intermediate leaf size (Takeuchi 14432, Tabili Creek, 100 m) whose fruitlets have three small oblong stigmas as in P. lessertianum. For the above reasons then P. lessertianum is maintained here to include P. pseudamboinense. However, for the convenience of those who might want to take the matter further, the ‘ List of Collections’ indicates which plants fit a rigorous definition of P. pseudamboinense (“ leaves greater than c. 20 × 10 cm; stigmas 2 - lipped ”). As mentioned above, Chew (2003) synonymized three New Guinea-typified names with P. lessertianum. I do not know whether any of those names might better fit P. pseudamboinense. For example, the protologue for P. viridibaccum (Trelease 1928) has reference to characteristics of both species: “ leaf blades 15 – 17 cm long … stigmas 2 ”.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72FFFA0930AB9AA908176E5.taxon	distribution	Distribution — Taiwan, throughout Malesia, also Vanuatu, Micronesia, Australia, and the Pacific Ocean region (Wallis & Futuna Is., Samoa). Habitat — In forest, 0 – 2000 (– c. 3300) m altitude. Notes — As noted in the Introduction, the New Guinea plants under this name are treated here as a species-complex. The four major named variants are as follows. (For types and publication data see ‘ Incertae Sedis’. The ‘ List of Collections’ indicates which specimens best fit the circumscriptions here). See Fig. 6 for the variation in leaf size and shape, etc. Additionally, a new large-leaved variety is described further below. 1. Piper macropiper s. str. – Leaves with a unilateral basal lobule, nerves basal or nearly so, glabrous or hirsute to villous; throughout New Guinea incl. Bismarck Archipelogo, 0 – 1500 (– 2500) m altitude. 2. Piper breviantherum – Leaves relatively short and narrow, without a basal lobule, nerves basal or nearly so, indument usually sericeous; infructescence relatively slender (6 – 9 (– 13) by 0.35 cm; central part of New Guinea, from the Star Mts (West Sepik Province) to the Eastern Highlands and Morobe Provinces, (490 –) 2000 – 2500 (– c. 3000) m altitude. 3. Piper novoguineense – Leaves without a basal lobule, nervation usually palmate-pinnate (i. e., usually one or more nerves suprabasal), usually conspicuously red-glandular, glabrous; mainly in Morobe Province (absent from the Bismarck Archipelago and islands of the Milne Bay District), c. 75 – 2800 m altitude. 4. Piper rodatzii – Leaves relatively broad, without a basal lobule, nervation palmate-pinnate (uppermost nerve from up to c. 1 / 3 way along midrib), chartaceous, glabrous; infructescence usually relatively short and stout (to c. 10 (– 14) by 0.7 cm), bracts with red-brown hairs spreading from near top of bract-stalk for up to c. 0.4 mm beyond margin of head, surface of bract-heads usually glaucous; throughout New Guinea, but apparently rare in the western half of island and in the Bismarck Archipelago and islands of the Milne Bay District, (100 –) 500 – 2500 (– 3290) m altitude. These four variants, if interpreted strictly according to the given characters, take in much of the variation in this complex, but nevertheless a considerable number of specimens cannot be placed into one or the other. This is not to say that all kinds of intermediates occur: a basal lobule, for example, is never found in a rodatzii - or breviantherum - kind of leaf blade. Another difficulty is that I have seen far too few specimens from West New Guinea to be sure that the last three taxa, all typified on Papuan New Guinea collections, properly represent the range of geographical variation on the island as a whole. For this reason especially, I am not willing to change their taxonomic status (say, to subspecies rank). Piper macropiper in the strict sense is common in New Guinea up to c. 1500 m, but above that one or other of the three variants seem to predominate. The four can certainly grow in the same general area – I have seen them thus in the Kaironk Valley, Schrader Range, between c. 1500 and 2500 m altitude. Specimens of climbing pipers are sometimes labelled as ‘ epiphytes’ or ‘ shrubs’. This is most often said about P. rodatzii – I have seen more than a dozen such examples, including two of my own from the Kaironk Valley at 1700 – 2200 m: Gardner 7078, “ low epiphyte on relict streamside-forest tree ”; 9681, “ bushy plant c. 1 m tall on open gully floor in primary forest, perhaps originally climbing? ”. Similarly, the label of Grubb & Edwards 187 (Fatima River, 2600 m) has: “ Scrambler / climber up lower boles, or, apparently, independent shrub ”, and that of Takeuchi 10541 (Bismarck Range, c. 2400 m): “ Epiphytic shrublet ”. Only a very few specimens elsewhere in the P. macropiper complex (e. g. P. breviantherum, LAE 65805 and P. macropiper s. str., Brass 13976) note such a habit. Specimens with small dendritic hairs (Fig. 4 c), these sometimes accompanied by a typical coarser indument, are from low to moderate altitude (to c. 1000 m) in Morobe District, e. g. Clemens 1257, 1721, 7959, 10824, 40849, Hartley 9765, NGF 26030, Takeuchi 4572) with one from nearby East Sepik District (LAE 73630). All belong to P. macropiper s. str. Other members of the complex from this region, including the new large-leaved variety described below, lack such hairs. In addition to the above-described variation in P. macropiper across the island there is one well-marked local variant, newly described as follows.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72FFFA191C0BC4D90E8750E.taxon	description	Piper macropiper var. macrophylla R. O. Gardner. — Type: Brass 32543 (holo A; iso CANB), Morobe Province, Markham Valley, Umi River, 480 m, locally common in flood-plain forest. Diagnosis: Distinctive in P. macropiper by its large leaves and by the indument on the nerves near the base of the blade below, which is of scattered long hairs over numerous short patent ones. Fertile-shoot internodes to c. 4 mm diam. Vegetative parts (newest stem, stipule, petiole and nervation of blade underside near base), and peduncle of the inflorescence, hirsute to subvillous with dense short pale multicellular hairs (to c. 0.5 mm) among which are few patent to straggling long hairs (to c. 2 mm). Leaf blades subcoriaceous, drying brown to yellowish, ovate, 20 – 35 by 8 – 17 cm; base rounded-truncate to shortly cordate, nearly equal at petiole, basal lobule small; apex shortly acuminate; main lateral nerves 2 – 3 pairs, all basal or nearly so; lower surface not conspicuously glandular; petiole c. 1 cm long. Male inflorescence a spike c. 20 cm long on a 5.5 cm long peduncle; anthers 0.4 mm diam, obscurely 2 - locular (locules nearly continuous at apex of short filament), dehiscing apically, at level of bract-heads. Infructescence c. 20 – 40 cm long, 6 mm diam, on a 4 – 10 cm peduncle; bracts stalked, sparsely pale-villous, bract-heads suborbicular, 0.75 mm diam; fruitlets free, oblong, c. 2 by 0.8 mm, flattened or low-rounded at apex; stigmas 3 (– 4), sessile, more or less oblong, together 0.25 – 0.4 mm diam. Distribution — Papua New Guinea, Morobe Province. Habitat & Ecology — In forest, c. 400 – 900 m altitude. Other specimens seen. Clemens 8216 (A, B), Boana, c. 750 m [‘ 2 – 3000 feet’]; Conn 100, South of Boana, disturbed forest, c. 900 m; Takeuchi 8675 (A, BRIT), North of Busu River, mature forest, 400 m. Notes — The nature of the infructescence allows no doubt that these plants represent a local development of P. macropiper. A basal lobule to the leaf blade seems to be present (though small and obscured by hairs) in at least the A duplicates of Clemens 8216 and Takeuchi 8675. For the holotype and Conn 100 it is unclear, because of the way in which these large-leaved plants have been mounted, whether or not a lobule is present. The unique indument of the proximal part of the nervation is shown in Fig. 4 d.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72EFFA191C3BC7F96DE74F7.taxon	distribution	Distribution — Java (and presumably elsewhere in central Malesia), Philippine Is., New Guinea (perhaps mainly in the western part of island), Solomon Is. Habitat & Ecology — In forest, sea level to c. 100 (– 840 m) altitude. Note — In the older literature this species was confused with P. decumanum (as noted by Quisumbing 1930: 47, Chew 1972: 7). The two are generally distinguishable by their colour on drying: the former becomes dark greyish and is often flushed dull maroon below, while the latter becomes greenish yellow.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72DFFA2930ABA3F91FB7551.taxon	distribution	Distribution — New Guinea, east to Milne Bay Province (Goodenough I.); also northern Australia. Apparently rare or absent from the Bismarck Archipelago; absent from the Solomon Is. Habitat & Ecology — In forest, 0 – 1000 (– 3200) m altitude. Notes — The highest-altitude collections I have seen are from the Owen Stanley Range: Hopkins 946, Mt Kenevi, 3200 m and Brass 4680, Murray Pass, 2840 m. In the Highlands Region (Simbu and Southern Highlands Provinces only) the species is known from just a few collections and attains only c. 2550 m (NGF 6389). Van Royen (1982) accepted P. mestonii as a member of the montane / subalpine flora, but I have seen no specimen that would confirm his stated uppermost altitude of 3500 m. Peekel (1984: 124, f. 201) apparently records this species for two localities on the eastern side of New Ireland, under the synonym P. rueckeri C. DC. His description, especially of the fruit-spikes being spiny-tuberculate, does suggest P. mestonii, but his figure is inconclusive. I have seen no specimen from the Bismarck Archipelago myself, nor did Chew (1972, 1992) cite any. Chew (1972, 1992) gave numerous synonyms based on types from New Guinea and Australia. The most salient of these is P. stenocarpum C. DC., a higher-altitude form distinguished only by its short narrow leaves and smaller fruit. In all but the most coriaceous leaves the nerves are narrowly and sharply prominent above. This, and the stipule usually being less than 1 cm long, help distinguish P. mestonii from glabrous members of the P. macropiper complex, where the nerves are usually rounded-prominent above and the stipules 1 – 2 cm long. In ripe fruits the surface of the fused ovaries has a coriaceous, nearly glossy character (Fig. 4 e). The styles too are stiffened and are 0.3 – 1 (– 2.5) mm long. The 2 (– 3) stigmas are usually short and broad (‘ 2 - lipped’) but occasionally are recurved and elongated, each branch being almost 1 mm long (e. g., Carr 16262, Northern Province, Kokoda and Clemens 10525, Morobe Province, Markham Valley). I have not been able to determine stamen number from any of the ten or so male specimens I have seen. However, in his description of P. stenocarpum Chew (1972: 17) states that its male flowers are “ 2 - staminate, filaments much longer than anthers at maturity ”. Dowe & Broughton (2007) have elucidated the circumstances of the name’s publication. See under P. versteegii for a comparison with that species.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72DFFBC91C0BA3F91FD708D.taxon	description	Piper subcanirameum (‘ subcaniramum ’) C. DC. (1923) 196, (1925) 221; Chew (1992) 163. — Type: Boorsma 7 (holo BO n. v.), New Guinea.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F72DFFBC91C0BA3F91FD708D.taxon	distribution	Distribution — New Guinea, mainly the Highlands Region, also Finisterre Range (Madang Province) and Morobe Province; apparently absent from the Bismarck Archipelago. Habitat & Ecology — In montane forest, (2000 –) 2500 – 3620 m altitude. Notes — The species resembles P. arfakianum in its rather small, coriaceous leaves; see Notes under that species for some distinguishing features. The two are found together in the vicinity of Mt Kaindi and Edie Creek (Morobe Province), from c. 2050 m (LAE 60357, NGF 30885) to c. 2450 m (P. arfakianum; NGF 32863) and 2650 m (P. subcanirameum; Durand & Nelson 146). The female inflorescences differ especially in that the fruitlets of P. arfakianum are usually free. However, Takeuchi 5756 (Mt Wilhelm, 2800 m, A, B, BISH) and Takeuchi 20136 (Porgera, 2900 m, A) are clearly P. subcanirameum by their leaves, but have mature spikes with substantially free fruitlets (Fig. 4 g). I accept both as representing this species, though possibly they are aberrant. De Candolle (1925) simply noted the type of P. subcanirameum as being from “ New Guinea ”, and gave no date of collection. Chew (1992: 163) assumed it came from West New Guinea (Irian Jaya), making it the only such collection he cited for that part of the island. I have not seen any from there myself, nor is the species recorded for Mt Jaya (Carstenz) by Johns et al. (2006). Van Royen (1982) did accept for his P. trombek two collections from Lake Habbema in West New Guinea, Brass 9134 and 9308. I am not sure of their identity but at least the former is not P. subcanirameum. a c With respect to a possible type locality for P. subcanirameum we should consider the statement by Van Steenis-Kruseman (1950: 71) that “ New Guinea Pipers were collected on his [W. G. Boorsma, fl. 1891 – 1922] behalf ”. Unfortunately, consideration of the identities of the nine consecutively numbered Piper specimens under Boorsma’s name (De Candolle 1925, Chew 1972: 22) does not narrow the field. Nevertheless I speculate that the type is from the eastern rather than the western part of the island, with the most likely such locality being the Huon Peninsula mountains, explored botanically by C. Keysser in 1909 – 1916 (Van Steenis-Krusemann 1950: 279). Chew (1992) compared the types of P. subcanirameum and P. trombek and found no significant differences. Van Royen (1982) included in his P. trombek two collections from the Mt Suckling complex (Northern Province), LAE 54062 and 55677. I have seen a duplicate of the latter at A (Tantam Plateau, 1980 m). It is small-leaved and completely glabrous and does not have the usual texture of P. subcanirameum. I cannot identify it (the inflorescences are in poor condition), and so omit both these collections from the mapped distribution. Van Royen (1982) described the fruits of P. trombek as orange or yellow, in agreement with other label-notes for P. subcanirameum, which generally just state ‘ fruit green’ (at least, I have seen none that say ‘ fruit red’). Three collections with what seem to be ripe fruits describe their colour as ‘ dull yellow’ (Clemens 7605), ‘ dull buff’ (Clemens 7556) or ‘ orange’ (NGF 24994).	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
03AF8788F733FFBC930AB9F896517559.taxon	description	Piper versteegii C. DC. (1910) 415; Chew (1972) 19. — Type: Versteeg 1136 (isolecto BO, L image!), West New Guinea, Noord-rivier. Ultimate leafy internodes c. 5 – 8 mm diam, usually with distinct narrow ridges c. 1 mm apart. Vegetative parts glabrous except for short patent hairs on stipule. Stipule to c. 1.5 cm long. Leaf blade (chartaceous-) subcoriaceous, drying mid-brown or greyish, broadly ovate (- elliptic), c. 13 – 25 by 10 – 20 cm; base rounded to cordate and equal at petiole; apex shortly acuminate; main lateral nerves usually 3 – 4 pairs, the lower 2 – 3 pairs basal, the upper pair from c. 1 / 3 way along midrib (or rarely, replaced by several relatively weak equal pairs from middle part of blade), all nerves rounded- to sharply prominent above; surfaces of blade not gland-dotted. Petiole stout, to c. 20 cm long, usually c. 1 / 3 – 1 / 2 as long as blade. Male inflorescence a fascicle of up to 7 short-peduncled spikes, at its base a congested group of stipule-like structures c. 0.5 – 1.5 cm long, sometimes 1 – several internodes c. 2 – 5 cm long interpolated between this grouping and the spikes; rachis glabrous, bracts of inflorescence sessile, orbicular, 0.4 – 0.8 mm diam; stamens 2 per flower, filament slender, c. 0.25 mm long, anther 0.25 – 0.4 mm diam, dehiscing laterally. Infructescence a stout spike to c. 8 cm long, 0.8 – 1.5 cm diam, on a peduncle c. 1 – 2 cm long, spikes solitary or 2 – 3 fascicled together as in the male, dull yellowish to orange when ripe; rachis and bracts as in male. Fruitlets usually fully concrescent, tapering above for c. 0.5 mm into a terete style c. 1 (– 2) mm long; stigmas 2 (– 4), narrow-ovate, glabrous or only minutely papillose (40), together (when fully spread) 0.8 – 1.5 mm diam. Distribution — New Guinea, apparently throughout; perhaps rare in the Bismarck Archipelago. Habitat & Ecology — In forest, to c. 1350 m altitude. Notes — This species appears to be widely but sparsely distributed in the New Guinea lowlands; I have seen fewer than twenty collections of it. I know of only one collection from the Bismarck Archipelago, (NGF 21973, West New Britain; BRI, image!). Presumably the species is uncommon there, since Peekel (1984) does not mention it. De Candolle’s protologue is based on three specimens, Versteeg 1136, 1350 and 1768, all obtained on the Noord-rivier (Lorentz River, West New Guinea) during the First Lorentz Expedition to Dutch Southern New Guinea, 1907. It contains information about both sexes. Chew (1972) chose the first collection as lectotype, without comment. The L duplicate of this appears to be female. Chew (1972: 20) described the fruitlets as only ‘ partially concrescent’ but I have seen only one such specimen (Takeuchi 12682, Eastern Highlands, Crater Mountain, 1400 m, A); all others have fully concrescent fruit. In the ripe fruit of P. versteegii (Fig. 4 f) the surface of the ovaries appears to be softer and somewhat granular as compared to the coriaceous, smooth and glossy texture seen in P. mestonii. Also, the fruits of these two species seem to be differently coloured, those of P. mestonii generally being described as ‘ red’ or ‘ dark red’ or ‘ bright red’. The stipule-like structures grouped into a kind of rosette between the inflorescence (s) and the leaf-tipped shoot below can be interpreted to suggest that the architecture of this species is not so very different from that of the other New Guinea climbing pipers: the distinctive feature of P. versteegii would be that its fertile shoots have become greatly shortened and have lost their leaves, all that remain being the stipules. The observation by Chew (1972) that P. versteegii has leaves whose petiole is conspicuously sheathing is then correct (the implicit comparison being with the fertile-shoot leaves of P. mestonii); note though that the latter’s sterile-shoot leaves are, as usual in climbing pipers, strongly sheathing. Chew (1972) saw a “ strong superficial similarity ” between P. versteegii and P. mestonii, especially in leaf venation, and Van Royen (1982) placed the former under the latter without comment. However, there is no equivalent in P. versteegii of the narrow-leaved, higher-altitude form of P. mestonii.	en	Gardner, R. O. (2013): Piper (Piperaceae) in New Guinea: the climbing species. Blumea 57 (3): 275-294, DOI: 10.3767/000651913X665053, URL: https://doi.org/10.3767/000651913x665053
