Scyphiphora hydrophylacea C.F.Gaertn., Suppl. Carp. 1(2): 91-92, tab. 196, fig. 2. 1806.

Berger, Andreas, 2023, Solved at last: The Philippine endemic Psychotria philippensis is a synonym of Scyphiphora hydrophylacea (Rubiaceae, Scyphiphoreae), PhytoKeys 221, pp. 127-148 : 127

Scyphiphora hydrophylacea C.F.Gaertn., Suppl. Carp. 1(2): 91-92, tab. 196, fig. 2. 1806.

Epithinia malayana = Epithinia malayana Jack, Malayan Misc. 1(5): 12-13. 1820. ≡ Scyphiphora malayana (Jack) Bedd., Fl. Sylv. S. India Forester’s Man. Bot.: cxxxiv-3, tab. 29., fig. 5. 1874, nom. inval. Type: Singapore. Singapore Island [protologue: "Found in Mangrove swamps on the Island of Singapore."], s.d., W. Jack s.n. (lectotype, designated by Wong et al. 2019: 283: L [L 0001344]).

Psychotria philippensis = Psychotria philippensis Cham. & Schltdl., Linnaea 4(1): 21-22. 1829a. ≡ Uragoga philippensis (Cham. & Schltdl.) Kuntze, Revis. Gen. Pl. 2: 962. 1891. Type: Philippines. Luzon, Calabarzon Region, Cavite Province: Noveleta [protologue: "Legimus in maritimis circa Tierra-alta Luçoniae.”], 1817-1818, L. K. A. von Chamisso s.n. (type, B†). Neotype: Philippines, Palawan Province: Culion Island, August 1913, L. Escritor s.n. in Merrill: Species Blancoanae 635 (neotype, here designated: US [00624079]; isoneotypes: L [L.2962064], P [P03972577], W [W 0131765]).

Ixora manila = Ixora manila Blanco, Fl. Filip.: 60-61. 1837. Type: Philippines. Palawan Province: Culion Island, August 1913, L. Escritor s.n. in Merrill: Species Blancoanae 635 (first-step neotype, designated by Merrill 1918: 364; second-step neotype, here designated: US [00624079]; isoneotypes: L [L.2962064], P [P03972577], W [W 0131765]).

Hydnophytum costatum = Hydnophytum costatum Drake, J. Bot. (Morot) 9: 240-241. 1895. Type: Vietnam. Quảng Ninh Province: Surroundings of Quảng Yên [protologue: "Environs de Quang-Yen, au milieu des palétuviers (685)."], August 1885, B. Balansa 685 (lectotype, designated by Wong et al. 2019: 283: P [P00836559]).

Type.

Indonesia. Java, Jawa Barat: Anyer [protologue: " Hydrophylax . Collect. Banks."], 2. Oct. 1770, unknown collector in J. Banks s.n. (lectotype, here designated: BM [BM000945301]) .

The type of Psychotria philippensis

Psychotria philippensis was published in a series on the botanical results of the Romanzoffian expedition on the Rurik prepared by Chamisso and Schlechtendal (1829a). The protologue gives "in maritimis circa Tierra-alta Luçoniae” as a rather minimalistic collection locality. Fortunately, more detailed information can be derived from the diaries and travelogues of Chamisso (1836a, b) and Kotzebue (1821). According to them, the Rurik stayed in the port of Cavite from 18 December 1817 to 28 January 1818 for refurbishments, the most important Philippine fortress and an arsenal of the Spaniards at that time. Knowing that Cavite had little to offer a botanist, Chamisso quickly settled in Tierra Alta, then a small village on the high shore of Manila Bay, where the sandy headland of Cavite joins the adjacent mainland.

Chamisso noted that the lush forests around Tierra Alta extend from the mountains to the coast, where " Rhizophora and other trees reach into the sea" ( Chamisso 1836b: 118). He spent most of his time in the Philippines here and roamed the area around Tierra Alta, where he also collected the type specimen of Psychotria philippensis ( Chamisso and Schlechtendal 1829a). Chamisso first passed through Tierra Alta on 27 December 1817, when he travelled overland from Manila back to Cavite. The French nobleman Don San Jago de Echaparre offered him hospitality there, and he returned to work in Tierra Alta a few days later. He stayed there until 12 January 1818, when he first left for an eight-day expedition to Taal Lake and Volcano in the interior, returning to Cavite shortly thereafter.

The type collection of Psychotria philippensis (apparently mentioned by him as one of the "other trees" in the remark quoted above) can thus be dated and localized to the mangroves around Tierra Alta and a period of about two weeks. According to information on the neotype, US 00624079, Scyphiphora hydrophylacea was already extinct in the entire Manila Bay region by around 1913, where it was once widespread ( Blanco 1837: 60-61, as Ixora manila ). This is even more tragic as Blanco reported that the native Tagalog names he attributed to the species, nilad and manilad or may-nilad, places where nilad is abundant, ultimately led to the name Manila. This notion is, however, rejected by modern authors (e.g., Baumgartner 1975).

After the destruction of B, where Schumann and Merrill had seen the only known type specimen, there should have been additional original material of Psychotria philippensis found in HAL and LE, which hold the largest extant parts of Chamisso’s herbarium (see above). However, no specimens are currently known in either of these herbaria (HAL: Braun and Wittig 2003; LE: Larisa Orlova, personal comment) or other collections (JSTOR Global Plants database, http://plants.jstor.org; accessed February 2023). Searches in the herbarium of the Natural History Museum, Vienna (W), where three Chamisso specimens of Psychotria recently resurfaced in the extensive private herbarium of Stephan Ladislaus Endlicher (1804-1849), professor of botany and director of the Botanical Garden and Botanical Museum of Vienna, were also unsuccessful ( Berger 2018; see also Bräuchler et al. 2021).

Therefore, the name Psychotria philippensis is neotypified here, fixing the application of the name after nearly two centuries of uncertainty (ICN, Art. 9.8, 9.13; Turland et al. 2018). The specimen designated as neotype was collected by Leonicia Escritor on Culion Island, Palawan, and is no. 635 of the exsiccatae series "Species Blancoanae" issued by Merrill. In this series, he distributed selected specimens that he considered particularly characteristic of the species published by Francisco Manuel Blanco (1778- 1845) in the first two editions of his "Flora de Filipinas" ( Blanco 1837, 1845) and the third edition edited by Andrés Náves ( Blanco 1877). Blanco did not preserve specimens, and the interpretation of some of his names remained problematic until Merrill (1918) published his critical revision "Species Blancoanae" after a thorough study of the publication. In 1916, he issued a corresponding set of "illustrative specimens" for each of Blanco’s names as he understood them.

For example, Merrill (1918: 364) identified Ixora manila ( Blanco 1837) as the previously published Scyphiphora hydrophylacea and distributed the specimens mentioned above to clarify the application of the name Ixora manila (see also Naves and Fernández-Villar 1880: 109, tab. 277). Thus, he anticipated the concept of the neotype, and his "illustrative specimens" can be understood today as first-step neotypifications of the respective names. In a subsequent second-step neotypification the designation may be narrowed to a single specimen (ICN, Art. 9.17, Turland et al. 2018; see also Nicolson and Arculus 2001). Indeed, many of Blanco’s names were later neotypified with specimens from Merrill’s "Species Blancoanae" (e.g., Nicolson and Arculus 2001).

According to a letter preserved at the United States National Herbarium (US), Merrill prepared 15 sets of his "Species Blancoanae", each with 1046 specimens, and sent the first to US. Furthermore, he stated that the set at US was the only set that included original data such as field labels or notes and typewritten drafts of the treatments in the "Species Blancoanae". According to Nicolson and Arculus (2001), specimens from these sets are now represented in many herbaria including A, B, BM, BO, CAL, F, GH, K, L, MO, NSW, NY, P, U, UC, US and W. These usually have minimal labels stenciled "Merrill: Species Blancoanae No." in black ink followed by a number stamped in blue ink ( Nicolson and Arculus 2001).

Although the above-mentioned gathering "Species Blancoanae" 635 is not from the Manila Bay, it serves as an excellent neotype for Psychotria philippensis because the gathering has duplicates in many herbaria and agrees with both Merrill’s concept of Scyphiphora hydrophylacea and his interpretation of the type of P. philippensis as its synonym. Therefore, a specimen of the gathering is here designated as the neotype of P. philippensis and the second-step neotype of Ixora manila .

The type of Scyphiphora hydrophylacea

With the application of Psychotria philippensis fixed, some further notes on Scyphiphora and Scyphiphora hydrophylacea seem useful. As to the type of genus and species, the names were published by the German botanist Carl Friedrich von Gaertner (1772-1850, original German orthography Karl Friedrich von Gärtner) in a “Supplementum” ( Gaertner 1806) to his father Joseph Gaertner’s (1732-1791, original German orthography Joseph Gärtner) pioneering work on fruit and seed morphology "De fructibus et seminibus plantarum" ( Gaertner 1788, 1790-1792). In writing his “Supplement”, C. F. Gaertner made extensive use of his father’s fruit and seed collection, which was largely based on specimens received from contemporary botanists. One of the most important sources J. Gaertner consulted was Banks’s herbarium in London in 1778, and large parts of his ‘Carpologia’ were based on material that he received on loan or as duplicates through Banks’ generosity ( Deleuze 1805: 23-24; Stafleu 1969). Following in his father’s footsteps, C. F. Gaertner also travelled to London in 1802 with the same aims ( Stafleu 1969).

J. Gaertner’s collection of fruits and seeds is kept in the herbarium of the University of Tübingen (TUB), as is the herbarium of his son, who added to his father’s carpological collection and kept his own herbarium ( Stafleu 1969). Yet, there is no extant original material of Scyphiphora hydrophylacea in the Gaertner herbarium at TUB (Uta Grünert, personal comment). However, the protologue (" Hydrophylax . Collect. Banksian.") indeed gives reference to traveler, naturalist and patron of science, Sir Joseph Banks (1743-1820), whose extensive private herbarium later became the foundation of the herbarium at the Natural History Museum, London (BM). There are a number of specimens and drawings of S. hydrophylacea at BM associated with Banks, his herbarium and endeavors.

Two specimens, both mounted together on one sheet, are curated there as types of the name. The sheet is annotated in pencil as type of Scyphiphora hydrophylacea with the place of publication of the name. The two specimens are labelled ‘1’ and ‘2’ in black ink on the mounting tape at the base of the shoots. The numbers correspond to meagre annotations of locality and collector on the verso of the sheet in the upper left corner, written in black ink, as was common practice in many herbaria at that time. These annotations are here-interpreted as being in the hand of Samuel Törner, who was employed as amanuensis in the Banks herbarium from 1792 to 1797 ( Marshall 1978).

Specimen ‘1’, BM 000945301, mounted on the upper left side of the sheet and annotated on the verso '1. Java prope Angerpoint. J. B.', is a small fragment of a fruiting branch with a single leaf. 'J. B.' refers to Joseph Banks and links the specimen to James Cook’s first voyage aboard HMS Endeavour from 1768 to 1771, which is confirmed by a printed label of a later date reading 'Java 1770-71 Banks & Solander’. The specimen was collected at ‘Angerpoint’, here identified as the present-day coastal town of Anjer or Anyer on Java, a natural harbor and important victualling station during the passage of the Sunda Strait. The diaries and travelogues of Cook and Sydney Parkinson (1745-1771), Banks’ illustrator ( Hawkesworth et al. 1773: 705; Parkinson 1773: 171; Cook 1893: 349), reported that a boat was sent ashore at Anger Point on 2 October 1770, to gather supplies, and the specimen was most likely collected then. However, neither Banks nor botanist Daniel Carlsson Solander (1733-1782) went ashore that day, as can be seen from Banks’s travelogue published by Hooker (1896: 363). For that reason, the gathering must have been made by one of the sailors gathering supplies, which probably explains the scrappy condition of the specimen.

Specimen ‘2’, BM 000945302, consists of four flowering branches and is annotated '2. Paolo Candor. Dav. Nelson’. It was collected by David Nelson (1740-1789), who participated in Cook’s third voyage aboard HMS Resolution (1776-1780) on behalf of Banks. The modern transcription of 'Paolo Candor’ is Pulo Condor, now Côn Sơn Island, Vietnam. The gathering can be dated to 21-28 January 1780, based on King’s (1784: 450-464) account of the Resolution’s visit to the island.

Banks and his collaborators had already collected Scyphiphora hydrophylacea in flower and fruit in Australia, which the Endeavour expedition visited before returning home via the Dutch East Indies and the Cape of Good Hope ( Banks et al. 1901: 46). Based on these gatherings, illustrations were produced under the unpublished name 'Ixoroides littoralis’, and the corresponding artwork is held in the "Cook First Voyage Artwork Collection" in the library of the Natural History Museum in London. These materials include a first pencil sketch with color notes by Parkinson, made in 1770 somewhere at the mouth of the Endeavour River (plate number A4/169A). After Parkinson died on the return from the voyage in 1771 (see Parkinson 1773), the watercolors were completed by James Miller (plate number A4/169B, A4/169 2) and engraved by Charles White (plate number A4/169C, also water colored A4/169 5), both under the patronage of Banks. The illustration remained unpublished for over a century until a posthumous publication on Banks’s Australian plants from the expedition, edited by James Britten ( Banks et al. 1901: 46, tab. 143). Corresponding specimens that could be located are BRI-AQ0450760, P00836560 and W0131767, all received from BM, and the former two are curated as types of S. hydrophylacea .

Another notable holding of Scyphiphora hydrophylacea material at BM is a long-unpublished watercolor by Ferdinand Bauer, made during the 1801-1803 circumnavigation of Australia on HMS Investigator under Captain Matthew Flinders. The watercolor shows a complete specimen with analysis of flowers and fruits, and was based on gatherings from the Northern Territory, also in the library of the Natural History Museum in London (Botany Library no. 70, Admiralty Library no. 122; see Mabberley and Moore 1999: 122, plate 70).

The materials of the Investigator expedition (1801-1803) were definitely not available at the Banksian herbarium during C. F. Gaertner’s visit in 1802. It is also unlikely that the Australian materials of the Endeavour expedition (1768-1771) were seen by him, although they were surely accessioned in a timely manner after Banks’ return. The specimens were gathered under the herbarium name 'Ixoroides littoralis’, and there is no reference to that name in the protologue of Scyphiphora hydrophylacea , where the material seen by Gaertner in the Banks herbarium is instead referred to the genus Hydrophylax L. f. ( Rubiaceae : Spermacoceae ). The only species currently assigned to that genus, Hydrophylax maritima , inhabits sea shores in India, Sri Lanka, Bangladesh and the western Thai part of the Malay peninsula. It has indehiscent fruits that superficially resemble Scyphiphora (e.g. Groeninckx et al. 2009; Puff 1986; Puff et al. 2021), which explains the use of the generic name for the undescribed material in the Banksian Herbarium, and also lends its name to the species epithet. Hence, the two specimens curated as types of Scyphiphora at BM remain to be considered as original material.

C. F. Gaertner described and illustrated the fruits of Scyphiphora hydrophylacea , but made no mention of the flowers ( Gaertner 1806: "Cor. … Stam. …”) and the vegetative parts as for many other species that he described. Therefore it seems more likely that C. F. Gaertner either received isolated fruits on loan and did not study one or both of the complete specimens, or that the material had already been studied or received on loan by J. Gaertner during his visit in 1778, two years before the collection of Nelson’s specimen, and that C. F. Gaertner built upon sketches, manuscripts or other materials inherited from his father. For these reasons, the flowering specimen gathered by Nelson (BM000945302) is not considered original material here, and the single fruiting specimen, BM000945301, is here designated as lectotype of the name S. hydrophylacea .

A few years after Gaertner’s (1806) publication, the plants were again described as a monotypic genus, Epithinia , by Jack, based on one of his gatherings from Singapore ( Jack 1820: 12-13). Epithinia malayana was first synonymized with Scyphiphora hydrophylacea by Korthals (1851: 203-204), and from Miquel (1857: 238-239) and especially Gray (1859: 307), onwards the names were consistently considered synonyms. The combination S. malayana (Jack) Bedd. was ascribed to Beddome and considered a valid name in various publications and databases. Although the name indeed first appears in the caption accompanying the respective figure analysis in his rare Foresters Manual ( Beddome 1874: tab. 29, fig. 5), it is not accepted in the corresponding text ( Beddome 1874: cxxxiv-3, using a combination of Roman and Arabic pagination). There, Beddome explains that the name used in the figure should be changed to S. hydrophylacea ("Under the name of Scyph. Epithinia malayana , which should be altered as above."). Apparently, the plates were engraved with the name before the text was finished, and Beddome changed his mind on the identity of the material during the completion of the text, as with few other species in the text. S. malayana is therefore not accepted by Beddome and is not valid according to Art. 36.1 (ICN, Turland et al. 2018). Wong et al. (2019: 283) designated the only currently known duplicate of the respective gathering W. Jack s.n. at L as lectotype.

Finally, the plants were again described as Hydnophytum costatum Drake ( Drake del Castillo 1895) and based on B. Balansa 685 gathered in Vietnam. Again, the single known specimen was designated as lectotype by Wong et al. (2019: 283).

Bibliography of Scyphiphora

Selected and more or less useful historic and contemporary information and illustrations on Scyphiphora hydrophylacea can be found in chronological order in the following publications, some of them under synonyms added in parentheses:

Gaertner (1806: 91-92, tab. 196, fig. 2), Jack (1820: 12-13, as Epithinia malayana ), Blume (1826-1827: 955), Chamisso and Schlechtendal (1829a: 21-22, as Psychotria philippensis ), de Candolle (1830: 477-478, as E. malayana , 505-506, as P. philippensis , 577), Richard (1830: 79, tab. 4, fig. 1), Wight and Arnott (1834: 423-424, as E. malayana ), Blanco (1837: 60-61, as Ixora manila ), Endlicher (1838: 525-526, 545, on the latter page as E. malayana ), Korthals (1851: 203-204), Griffith (1854a, b: 269-271, tab. 478, as E. malayana ; 1854b: tab. 644A, is the same figure in different composition but named Lumnitzera pentandra , the corresponding text accompanying the figure - 1854a: 684 - apparently describes the actual Lumnitzera ), Hasskarl (1855: 16-17), Miquel (1857: 238-239), Gray (1859: 307), Bentham and Mueller (1867: 417-418), Beddome (1874: 134-3, tab. 29, fig. 5), Kurz (1877: 4), Hooker (1880: 125), Naves and Fernández-Villar (1880: 109, tab. 277), Vidal (1883a, b: 29, tab. 57, fig. G, 1886: 154), Trimen (1894: 337), Schumann (1897: 80, fig. 29 B, C), Banks et al. (1901: 46, tab. 143), Koorders and Valeton (1902: 124-127), King and Gamble (1904: 227-228), Elmer (1906: 32), Koorders (1912: 258), Koorders and Valeton (1915: fig. 563), Merrill (1918: 364, 1923b: 533), Pitard (1922-1924: 280-282, tab. 23, figs 5-8), Ridley (1923: 88-89), Watson (1928: 84-87, tab. 43), White (1929), Valeton (1930: 303), Corner (1940: 559), Henderson (1950: 217-218, fig. 196), Bakhuizen van den Brink (1955: 101-102, 1975: 34-35), Backer and Bakhuizen van den Brink (1965: 316), Wong (1988: 197), Wong (1989: 408), Keng (1990: 161), Hộ (1993: 206, fig. 7607), Puff and Rohrhofer (1993, richly illustrated), Turner (1995: 446), Ridsdale (1998: 235), Mabberley and Moore (1999: 122, pl. 70), Banerjee et al. (2002: 262), Puff et al. (2005: 82, pl. 3.1.19), Tao et al. (2011: 323) and Wong et al. (2019: 282-285, fig. 71).